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MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

VOLUME XXxIV
1987

BOARD OF EDITORS

Class of:

1987—J. RZEDOwsKI, Instituto de Ecologia, A.C., Mexico
DorOTHY DOUGLAS, Boise State University, Boise, ID

1988—SusAN G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA

1989—FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley

1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson

1991—Davip J. KEL, California Polytechnic State University, San Luis Obispo
JAMES HENRICKSON, California State University, Los Angeles

Editor— WAYNE R. FERREN JR.

Associate Editor—BARRY D. TANOWITZ
Department of Biological Sciences
University of California, Santa Barbara, CA 93106

Published quarterly by the
California Botanical Society, Inc.
Life Sciences Building, University of California, Berkeley 94720

Printed by Allen Press, Inc., Lawrence, KS 66044

Few botanists have had a more profound impact on the fields of
plant reproductive biology and evolutionary ecology than Herbert
and Irene Baker. For almost half a century, they have pioneered the
study of plant reproductive systems, with major contributions rang-
ing from Herbert’s classical early work on the evolutionary impor-
tance of variation in plant breeding systems to Herbert and Irene’s
definitive recent work on the ecological significance of pollen, nectar,
and fruit chemistry. In addition, Herbert has contributed many sem-
inal ideas toward understanding the evolutionary ecology of colo-
nizing species and weedy species. Yet, in many ways, the most

11

enduring impact that Herbert and Irene have had is on the many
students, colleagues, and friends who have taken their legendary
ecology courses at the University of California, accompanied them
on field trips in California, Costa Rica, and elsewhere, or attended
the stimulating and pun-filled evening meetings at their home in the
Berkeley hills. Their contagious enthusiasm, quick wit, and unlim-
ited interest in sharing insights about plant ecology and evolutionary
biology have served as a continual source of inspiration. In return,
it is with fondest affection that we dedicate this volume of Madrono
to them.

rab

TABLE OF CONTENTS

AIKEN, SUSAN G. (see ARGUS, GEORGE W. and SUSAN G. AIKEN)

ALLEN, LINDA (see CURTO, MICHAEL et al.)

ALMEDA, FRANK, Message from the past CBS president 22.2... ccccccsesccccccscssseseseeeeeeeens

ARGUS, GEORGE W., Noteworthy collection Of S@liX Z@VCVIANA o....iccccccceeecccccseeeeen

ARGUS, GEORGE W. and SUSAN G. AIKEN, Noteworthy collection of Festuca
TRUETULEL L/L OV 0 op mht ss te To foe eee ee

ARGUS, GEORGE W. and T. C. BRAYSHAw, Noteworthy collection of Salix
ICCA Het Pie entre eer ec Sate eta aes tate Alte. einer ne

AYALA, RICHARD (see BULLOCK, STEPHEN H. et al.)

BAKER, HERBERT G. (see BULLOCK, STEPHEN H. et al.)

BAKER, IRENE (see BULLOCK, STEPHEN H. et al.)

BISWELL, HAROLD H. (see EVANS, RAYMOND A. et al.)

Bowers, JANICE E. (see MCLAUGHLIN, STEVEN P. et al.)

BOWMAN, RoBERT N., Clarkia concinna subsp. automixa (Onagraceae), a new
subspecies from the South Bay region, central California 00.

BRAYSHAW, T. C. (See ARGUS, GEORGE W. and T. C. BRAYSHAW)

BREEDLOVE, DENNIS F., Review of Flora Fanerogamica del Valle de Mexico.
Volumen ITI. Dicotyledonae (Euphorbiaceae—Compositae (by Jerzy Rze-
dowski and Graciela C. de RZCGOWSKA) o2....ii.e.e..ccicesccssseececsssssessesssessssssveessessssueesessssseseee

BROYLES, PAULEEN, A flora of Vina Plains Preserve, Tehama County,
G71 U0) cl: Raeeetae ee een ane aR ee i ne OeaE aT Re Ae Se RAINE ER ER ele

BULLOCK, STEPHEN H., RICHARD AYALA, IRENE BAKER, and HERBERT G. BAKER,
Reproductive biology of the tree Jpomoea wolcottiana (Convolvula-
CAE) ree eet arr a2 see OME ee eee OU een ee er me eee ee

Busu, J. K. and O. W. VAN AUKEN, Some demographic and allometric char-
acteristics of Acacia smallii (Mimosaceae) in successional communities ..

CARTER, ANNETTA, Review of Xdntus, The Letters of John Xantus to Spencer
Fullerton Baird from San Francisco and Cabo San Lucas, 1854-1861 (by
TaN 0) ct We A','3 001 wo) omen eee anim nee tae RD Rennie Deora pre-e) =) BY eres BRAD ure a, AWE

CONSTANCE, LINCOLN, Review of Marcus E. Jones: Pioneer western geologist,
mining engineer & botanist (by Lee W. Lenz) oo ccieeeeeeceseseeeeeeeeeeeeneeeeeeeeeenenee

CORNETT, JAMES W., Cold tolerance in the desert fan palm, Washingtonia filifera
CATO CACC AS) ae cca ee aes ree cr

CUMMINGS, ROBERT, Review of Poisonous plants of California (by Thomas C.
Fuller and Elizabeth McC imtock) oi. ccssseeccssseecssseeeessueessssueesssseesseuesessuesssnseesees

CurRTO, MICHAEL, LINDA ALLEN, and GEOFFREY LEVIN, Noteworthy collection
OE FCSTUCE OCCLQCTECITS eee eae ea i ane eel

Day, MAGGIE, Review of Botanical illustration: Preparation for publication (by
Noel H. Holmeren and Bobbie Angell) .......0 eee

EvANS, RAYMOND A., HAROLD H. BISWELL, and DEBRA E. PALMQUIST, Seed
dispersal in Ceanothus cuneatus and C. leucodermis in a Sierran oak-
WV OOCIATICUSA Vala nts cn oh Bee) Ah i oe, Ss eal 2 ane caea e

EWAN, JOSEPH, Roots of the California Botanical Society 0...

FERNANDEZ, CARMEN F., LOWELL E. URBATSCH, and GENE SULLIVAN, Alloi-
spermum insuetum (Asteraceae: Heliantheae), a new species from Colom-
| ace emcee ann OO PEN TORO ONTO POSE SEED ASO Ng ce oe eo

FERREN, W. R., JR., Editor’s report for VOlUMEe 34 ooo eecceeecseeecseeesceeeesseeeeeeeeeneees

FIEDLER, PEGGY LEE and RoBErRT A. LEIDy, Plant communities of Ring Moun-
tain. Preserve, Marin County, Califomig 2... ee

GE-LIN, CHU (see STUTZ, HOWARD C. et al.)

HALL, KENNETH R. F. (see MCLAUGHLIN, STEVEN P. et al.)

HALVORSON, WILLIAM L. and RICHARD E. KosKE, Mycorrhizae associated with
an invasion of Erechtites glomerata (Asteraceae) on San Miguel Island,
CNT OTA AS Bc oe a Bcc eran a ence he,

269

268

41

270

209

304

250

269
72

a7

HELMS, JOHN A., Invasion of Pinus contorta var. murrayana (Pinaceae) into
mountain meadows at Yosemite National Park, California 200000000...

HELMS, JOHN A. and RAYMOND D. RATLIFF, Germination and establishment
of Pinus contorta var. murrayana (Pinaceae) in mountain meadows of
Yosemite National Park, CalifOrmia 2... ees eecceeeeeneeeeesneeessneesseneesssneesesueeseeneess

HOLLAND, DAN C., Prosopis (Mimosaceae) in the San Joaquin Valley, Cali-
fornia: vanishing relict Or reCemt VAT ccc eee cece eeececeenneeeeceecnnnnnnnennnne

HUFFORD, LARRY D., Inflorescence architecture of Eucnide (Loasaceae) ............

KEELEY, JON E., Fruit production patterns in the chaparral shrub Ceanothus
CTAS SUV OVELLS eer ea aoe ener een hte oles cus nis trceince, ae ee eee

KEELEY, JON E. and STERLING C. KEELEY, Role of fire in the germination of
chaparral herbs and suffrutescemts 2. eeessneeeesnneeesennessennneesesnneeseenneesecs

KEELEY, STERLING C. (see KEELEY, JON E. and STERLING C. KEELEY)

Ket, DAvip J., MELISSA A. LucKow, and DONALD J. PINKAVA, Cymophora
(Asteraceae: Heliantheae) returmed to Tiida x ee... eeeeecccccccseeeeeeeeeeeeeeeetnneeeeeeeeenenn

KELLEY, WALTER A. (see SWANSON, JOHN R.)

KNIGHT, WALTER, Review of Vascular plants of Upper Bidwell Park, Chico,
GAADY Viermonabie OSwald ) e120 sree i ee a se nine

KNIGHT, WALTER, Review of 4 flora of Dry Lakes Ridge, Ventura County,
Gali ormid (Dye DAVid Won NUACICY)) essa ee eee a ce

KoskKE, RICHARD E. (see HALVORSON, WILLIAM L. and RICHARD E. KOSKE)

KOWALSKI, DONALD T., New records of Myxomycetes from California. VI .....

LAJTHA, KATE, JOHN WEISHAMPEL, and WILLIAM H. SCHLESINGER, Phosphorus
and pH tolerances in the germination of the desert shrub Larrea tridentata
RZ OD VAC AS) reas ear e cco etter Ao a taco eA eee ar

LAWESSON, JONAS E. (see NORMAN, ELAINE M. and JONAS E. LAWESSON)

LeIDY, ROBERT A. (see FIEDLER, PEGGY LEE and RoBeErT A. LEIDy)

LENIHAN, JAMES M. (see SUGIHARA, NEIL G. et al.)

LEVIN, GEOFFREY A., Noteworthy collections of Astragalus hypoxylus and Mim-
CTI ISS CLON A) 200) 1 sca Aree EEO a OE RO

LEVIN, GEOFFREY A. (see CURTO, MICHAEL et al.)

LEVIN, GEOFFREY A. (see ZEDLER, PAUL H. et al.)

LisTON, AARON, Noteworthy collection of Peterid thOMpPSONGE 0....ccccccccoveeeeeeeeeeeee-

Luckow, MELISSA A. (see KEIL, DAvIpD J. et al.)

MAGNEY, DAvID L., The range and two new locations of Boschniakia strobilacea
(COTES) oF20 a6) 01 eter? ay ) ee ae ae neat as nes ce ND mv bse cts Westnet rar aon: MUUna RON IR so

Mayor, JAcK, A review of Uinta Basin flora (by Sherel Goodrich and Elizabeth
ANSE See rn sah cee eta Aces Pena Arr Ate one nat as vis oe ore nee er tek ae a ee

MARRS-SMITH, GAYLE and JAN NACHLINGER, Noteworthy collections of As-
tragalus gilmanii, Erigeron ovinus, and Polygala subspinosa var. hetero-
FIL V CI Co ere ee ak eres grecoeMb seen sdeebensoaeent spas kechc nascar aris os aN

McARTHUR, E. DURANT (see STUTZ, HOWARD C. et al.)

MCLAUGHLIN, STEVEN P., JANICE E. BOwErRS, and KENNETH R. F. HALL, Vas-
cular plants of eastern Imperial County, California 2000

MCNEAL, DALE W., Allium shevockii (Alliaceae), a new species from the crest
of the southem Sierra Nevada, Califomia... 2

MINNICH, RICHARD A., The distribution of forest trees in northern Baja Cali-
i Eo ob Fe Vcd ys [25-016 0 x eee ete oe RU ON RRTIVE re ARE RA ge NOE a OS aa eee

MoorING, JOHN, Range extension, chromosome count, and mephitism in Les-
SUPPOTO ST GH UTS CE OMI DS OSIUAS) tee Dae Ble Gl, ooh ee Le,

MorRAN, VIRGINIA (See ZEDLER, PAUL H. et al.)

NACHLINGER, JAN (see MARRS-SMITH, GAYLE and JAN NACHLINGER)

NAKAI, KEI M., Some new and reconsidered California Dudleya (Crassulaceae)

NORMAN, ELAINE M. and JONAS E. LAwEsson, Noteworthy collection of Bud-
CALC RCUTI VET NCCT | ee i cates re ae ne 2 en, Pa cass, Dey ee soe

|

qa

324
18

2D

240

354

169
Pat |

48

63

170

381

579

172

382

S50

150

98

168

334

PALMQUIST, DEBRA E. (see EVANS, RAYMOND A. et al.)

PARFITT, BRUCE D., Noteworthy collection of Lygodesmia grandiflora ................

PARKER, KATHLEEN C., Seedcrop characteristics and minimum reproductive
size of organ pipe cactus (Stenocereus thurberi) in southern Arizona ..........

PINKAVA, DONALD J. (see KEIL, DAvID J. et al.)

RATLIFF, RAYMOND D. (see HELMS, JOHN A. and RAYMOND D. RATLIFF)

RAY, MARTIN F., Soliva (Asteraceae: Anthemideae) in California 0.00...

REED, Lois J. (see SUGIHARA, NEIL G. et al.)

RILLING, TRUDY (see ZEDLER, PAUL H. et al.)

SANDERSON, STEWART C. (see STUTZ, HOWARD C. et al.)

SCHLESINGER, WILLIAM H. (see LAJTHA, KATE et al.)

SHOWERS, MAry ANN T., A systematic study of Silene suksdorfi, S. grayi, and
S.. Sargent (Caryophylaceae) ss.cc5 etn ee eee

SMITH, CLIFTON, Review of Flora of the Santa Monica Mountains, California
(by Peter H. Raven, Henry J. Thompson, and Barry A. Prigge) ...............

SPIRA, TimoTHy P., Alpine annual plant species in the White Mountains of
Caster, Calformla: 2255-5 ore eee het oh eel ented ca

STUART, JOHN D., Fire history of an old-growth forest of Sequoia sempervirens
(Taxodiaceae) forest in Humboldt Redwoods State Park, California ..........

STUTz, HOWARD C., STEWART C. SANDERSON, E. DURANT MCARTHUR, and
CuHu GE-LIn, Chomosome races of Grayia brandegei (Chenopodiaceae) ..

SUGIHARA, NEIL G., Lois J. REED, and JAMES M. LENIHAN, Vegetation of the
bald hills oak woodlands, Redwood National Park, California 0000000...

SULLIVAN, GENE (see FERNANDEZ, CARMEN F. et al.)

SWANSON, JOHN R. and WALTER A. KELLEY, Claytonia palustris (Portulacaceae),
anew Species trom, Cantornia: 22 ate, o ee ee eee

Topson, THOMAS K., Noteworthy collection of Penste@Mmon raMoOSUs -.00..2....-0000----

TURNER, B. L., A new species of Axiniphyllum (Asteraceae: Heliantheae) from
| DV Vash ot10 my ws (op 6 (a,c Rameau nR es TORRE Iae Ob Senter totes ate Ne, Pee ORD etn eee. ee

URBATSCH, LOWELL E. (see FERNANDEZ, CARMEN F. et al.)

VAN AUKEN, O. W. (see BusH, J. K. and O. W. VAN AUKEN)

WEBER, WILLIAM A., Noteworthy collection of Byrur DUINLL oo.........c..cccseccces cee

WEISHAMPLE, JOHN (see LAJTHA, KATE et al.)

WHITTEMORE, ALAN T., Noteworthy collections of Athalamia hyalina, Cam-
pylium halleri, Entodon schleicheri, Manni fragrans, Mylia anomala,
Preissia quadrata, Riccia albida, and Timmia megapolitana subsp. ba-
(0) 16? Me ae aa te ER ete eeareeP lien esse te eeTR ied Em era ees MMM ed Me a kas an

ZEDLER, PAUL H., VIRGINIA MORAN, TRUDY RILLING, and GEOFFREY A. LEVIN,
Noteworthy collections of Agrostis avenaceae and Scribneria bolanderi ...

v1

294

228

29

70
315
128
142
3
Es)
171

165

171

69

381

NEW PAGE CHARGES FOR MADRONO AUTHORS

The Executive Council of the California Botanical Society has initi-
ated a new policy for page charges. Starting with papers received by the
editor after 31 Dec 1987, each author will receive five free pages per
volume of Madrono. The decrease from 20 free pages every two volumes

is necessary because of the escalating costs of publication. We wish to
maintain the size and quality of issues, and this will be possible only
with greater financial commitment from authors. We greatly appreciate
the many papers contributed to Madrono and look forward to serving
the membership and botanical community with quality volumes in the
future.

DATES OF PUBLICATION OF MADRONO, VOLUME 34

Number 1, pages 1-76, published 31 March 1987
Number 2, pages 77-172, published 30 June 1987
Number 3, pages 173-272, published 30 September 1987
Number 4, pages 272-392, published 20 January 1988

Vil

ANNOUNCEMENT

MADRONO BACK ISSUES AVAILABLE

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is the time to inform your periodical manager to acquire back issues to complete
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To order, provide desired volume and issue numbers to:

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% Department of Botany

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Inst. Ind.

Cost per issue

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a;
| VOLUME 34, NUMBER 1 C K JANUARY-MARCH 1987

MA

A WEST AMERICAN JOURNAL OF BOTANY

{
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RONO

Si,
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Contents ss ‘~

' ROOTS OF THE CALIFORNIA BOTANICAL SOCIETY sj WUE F 4 1989

ape
—

Joseph Ewan
INFLORESCENCE ARCHITECTURE OF Eucnide (LOASACEAE)\ : ite ee

Larry D. Hufford a ae ISR, PRS crt 18
A SYSTEMATIC STuDY OF Silene suksdorfii, S. grayi, AND S. ee eel cian

(CARYOPHYLLACEAE)

Mary Ann T. Showers 29
Clarkia concinna SUBSP. autoxima (ONAGRACEAE), A NEW SUBSPECIES FROM THE

SOUTH BAY REGION, CENTRAL CALIFORNIA

Robert N. Bowman 4]
NEw RECORDS OF MyYXOMYCETES FROM CALIFORNIA. VI.

Donald T. Kowalski 48

COLD TOLERANCE IN THE DESERT FAN PALM, Washingtonia filifera (ARECACEAE)

James W. Cornett ov
PHOSPHORUS AND PH TOLERANCES IN THE GERMINATION OF THE DESERT SHRUB Larrea

tridentata (ZYGOPHYLLACEAE)

Kate Lajtha, John Weishampel, and William H. Schlesinger 63
NOTEWORTHY COLLECTIONS

CALIFORNIA 69

MEXICO 69
REVIEWS 70
ANNOUNCEMENTS 17, 28, 40, 47, 62, 68, 74, 75, 76

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
ciety, Inc., and is issued from the office of the Society, Herbarium, Life Sciences
Building, University of California, Berkeley, CA 94720. Subscription rate: $25 per
calendar year. Subscription information on inside back cover. Established 1916.
Second-class postage paid at Berkeley, CA, and additional mailing offices. Return
requested. POSTMASTER: Send address changes to James R. Shevock, Botany Dept.,
California Academy of Sciences, San Francisco, CA 94118.

Editor—WaAyne R. FERREN, JR.
Associate Editor—BARRY D. TANOWITZ

Department of Biological Sciences
University of California
Santa Barbara, CA 93106

Board of Editors
Class of:

1987—J. RZEDowsKI, Instituto de Ecologia, A.C., Mexico
DoroTHy DouGLas, Boise State University, Boise, ID
1988—SusAn G. CoNARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989—FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991—Davip J. KEL, California Polytechnic State University, San Luis Obispo
JAMES HENRICKSON, California State University, Los Angeles

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1986-87

President: FRANK ALMEDA, Department of Botany, California Academy of Sciences,
San Francisco, CA 94118

First Vice President: PATRICK E. ELVANDER, 273 Applied Sciences, University of
California, Santa Cruz, CA 95064

Second Vice President: KINGSLEY R. STERN, Department of Biology, California State
University, Chico, CA 95929

Recording Secretary: V.THOMAS PARKER, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
Francisco, 50 Phelan Ave., San Francisco, CA 94112

The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, CHARLES F. QUIBELL, Department of Biology,
Sonoma State University, Rohnert Park, CA 94928; the Editor of MADRONO; three
elected Council Members: THOMAS FULLER, 171 Westcott Way, Sacramento, CA
95864; ANNETTA CARTER, Department of Botany, University of California, Berkeley,
CA 94720; JOHN Moorina, Department of Biology, University of Santa Clara, Santa
Clara, CA 95053; and a Graduate Student Representative, NIALL F. MCCARTEN,
Department of Biological Sciences, San Francisco State University, San Francisco,
CA 94132.

ROOTS OF THE CALIFORNIA BOTANICAL SOCIETY*

JOSEPH EWANT
Department of Biology, Tulane University,
New Orleans, LA 70118

ABSTRACT

The historic roots of the California Botanical Society are traced from 1853, when
the California Academy of Natural Sciences was founded, to the organizational meet-
ing in 1913 called by W. L. Jepson. Twenty-five organizations and events more or
less influenced the Society’s origins. Natural history events, especially botanical, in
the East, the role of the California Botanical Club, the Sierra Club, and rivalries are
noted particularly.

‘“*Some time there will be here in Berkeley a wild-flower protection
society, just as in the older states... . Some time, gentle reader, the
call will come down from the mountain top... and everywhere the
expression of the overmastering desire—the love of life.”’ This from
a short essay published by Jepson in 1898 entitled ““The Love of
ite 2

The historian of the sciences must divest prejudices and special
interests of his own age, so far as that is possible, and migrate into
a strange land to bring back as unbiased an account as he can of
what he has learned. He must not treat the past as one in spirit with
the present. To paraphrase William Ferguson,’ we must be alive to
the existence of many different pasts, leading without predetermined
succession, much less a progression. The historian pursues the facts
and fastens upon them. “‘The dead were and are not. Their place
knows them no more and is ours today. Yet they were once as real
as we and we shall tomorrow be shadows like them.’’? Perhaps we
should remember Montesquieu: in the infancy of societies, the chief
shapes the institution; later the society shapes the chief.

Were there differences between the Eastern and Californian de-
votees of botany? Botanists beach-combed Boston harbor for Eu-
ropean ballast weeds and excitedly published their trophies in the
Bulletin of the Torrey Club or Rhodora. Californians found stow-
aways from Chile or Mazatlan. Sometimes they asked “‘is it native?”’
or “‘are birds carrying the seeds?” Compared with the flora of the
Eastern United States, California proved rich in endemics. Indian
uses of plants had been studied, too. In the Mother Lode it was
Digger Indians amid the Digger pines. Jepson called anthropologist
Pliny Earle Goddard “‘my traveling companion on an expedition to

* Presented in shorter form at the Annual Banquet of the Society at Berkeley on
22 February 1986. + Present address: Missouri Botanical Garden, St. Louis, MO
63166-0299.

MADRONO, Vol. 34, No. 1, pp. 1-17, 1987

2 MADRONO [Vol. 34

the South Fork of the Salmon River in 1902’’, where Goddard was
noting the food and ceremonial plants of the Hupa tribe.

ALBERT KELLOGG AND THE ACADEMY

Roots of the California Botanical Society (CBS) may be traced
back to the pioneering spirit of Albert Kellogg, who with three other
doctors, a real estate agent, and a school superintendent met to
organize the California Academy of Natural Sciences (Fig. 1). From
Kellogg’s taste, botanical discoveries dominated the Academy’s de-
liberations. During the first decade 123 papers were published of
which 43 concerned trees and flowering plants.

When Dr. Kellogg arrived in Sacramento on 8 August 1849, he
surely brought the argonaut spirit, borne in the sailing ship around
the Horn, the year “the world rushed in’’. As a youth, he had enjoyed
natural history in Connecticut and brought that sharing of enthu-
slasms with amateurs so important in scientific societies. Jepson
relates the welcome reception Kellogg gave him as a young man‘ on
his first visit to the old Academy. The first of the many endemic
species Kellogg described was the Channel Island mallow, Lavatera
assurgentiflora. With a certain patriotism, he described the noble
Washington Lily and with almost missionary zeal, the oracle oak,
Quercus morehus. Occasionally, novelties slipped away from the Bay
Region botanists to be first published by the Eastern Establishment
and foreigners. Kellogg became almost militant and endeavored to
publish “‘new species”’ more promptly through the Academy’s Bul-
letin. He had seen John Lindley herald the Sierra sequoia in London
before American scientific circles awoke.° Kellogg, who lived 38
years in “the bosom of the urgent West’’, was the pioneer spirit.

TRANSCONTINENTAL AND STATE SURVEYS

Three enterprises brought California to the attention of botanists
in the East and beyond. Locally, the California Geological Survey,
commonly referred to as the “‘State Survey”’, staked out the limits
of knowledge for the three kingdoms. The Botany of California in
two volumes was especially important for the field work of William
Henry Brewer, with his attention to detail and careful numbering of
collections, and for his journal that documents his travels. From the
surviving copies with their marginal notes and queries, it is evident
how that important reference-work served California botanists.® If
you have not read Brewer’s account, edited by Francis Farquhar
under the title Up and Down California, you have a rewarding ex-
perience ahead.’ On one occasion John Muir wrote, ““blessed Brewer
of a thousand speeches and stories and merry ha-has.”’

Secondly, the Pacific Railroad Surveys carried out by the U.S.
Army Corps of Engineers seeking a practicable route from the Mis-

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY 5

ROOTS OF THE CALIFORNIA BOTANICAL SOCIETY

International California
Phytogeographic Botanical Pacific Division
Excursion Society of A.A.A.S.
Sept 1913 12 Apr 1913 12 Apr 1913
1910
Harriman
New England Alaska
ey ay PAE Botanical Club ee
28 May 1892 ie
New York University
Botanical Stanford California of California
Garden University Botanical Club Botanic Garden
1891 1891 7 Mar 1891 1891
1890
Bureau of
Biological
New Orleans Survey
San Diego Naturalists’ Cotton Centennial 1885-1940
Society of Directory 1884
Natural History 1877
1874
California San Francisco
Arnold College of Microscopical
Arboretum Pharmacy Society
1872 1872 1872, 1893
1870
Torrey
University American Overland Botanical
of California Naturalist Monthly Club
1868 1867 1868-1876 1867
Harvard University
Gray Herbarium
1864
California California
Pacific Academy of Geological
RR Surveys Natural Sciences Survey
1853-1859 1853 1853-1864
1850

Fic. 1. Roots of the California Botanical Society. Events, institutions, and or-
ganizations with botanical connections to the Society are identified by the leaf symbol.
Publications are printed in italic. Dates represent the day or year in which an event
occurred or was initiated or when an institution, organization, or journal was founded.

4 MADRONO [Vol. 34

sissippi River to the Pacific? was another enterprise in the growth
of California botanical knowledge, but the naturalists who accom-
panied the surveyors in their buckboard wagons and on horseback
delivered collections to John Torrey in New York and Asa Gray at
Harvard.

The essential reference herbarium delivered by Asa Gray to Har-
vard in 1864° was the third event. The “‘“Gray Herbarium” and the
comprehensive library was to stand beside the Hooker Herbarium
at Kew in calling for necessary visits by California students through
the years.

TORREY BOTANICAL CLUB

Three years later (1867) and twelve years after the founding of
the California Academy of Sciences, John Torrey, beloved by the
botanical community, became the center piece for the Club that,
against his wish, was to bear his name.!° There were 31 founding
members. William Henry Leggett, a mainstay of its early years, began
distributing a four-page monthly sheet in 1870 as a privately funded
venture to which he gave the name “Bulletin of the Torrey Botanical
Club”. He admitted the Club was “rather informal and somewhat
fluctuating’’. Prominent in the Club was George Thurber, editor and
publisher of horticultural titles, who was instrumental in incorpo-
rating the Club. Today the Club of Leggett and Thurber flourishes
as the oldest exclusively botanical society in America. Thurber’s
herbarium came to the Academy in 1893, only to be lost in the
holocaust of 1906. John Strong Newberry, who had collected in
California with the Pacific Railroad Survey, was president of the
Club for ten years. Newberry’s interest was fossil plants and he taught
paleontology at Columbia College from 1866 to 1890.

American Naturalist AND Overland Monthly

How does the American Naturalist published in Salem, Massa-
chusetts, feed the roots of a California society? Following the trans-
continental railroads and the opening of the West, the signing of the
Morrill Act in 1862, and the building of land-grant colleges, there
followed a growing interest about the living things of this wilderness
and wide-open spaces. Varied, informative, entertaining essays on
the prairie dogs, pronghorns, burrowing owls, and locoweeds—life
forms never seen in the East—appeared in the nineteen volumes of
the American Naturalist of the late 19th century. Classes in acade-
mies and female seminaries were reading George Perkins Marsh’s
Man and Nature. To learn more about the West there was the Amer-
ican Naturalist with first-hand descriptions by Elliott Coues, Edward
Palmer, William Henry Brewer, James Graham Cooper, and others.
Cooper, for example, who had been an army surgeon with the Pacific

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY 5

Railroad Surveys, wrote the ornithology volume for the California
State Geological Survey— Brewer had collaborated with Asa Gray
and Sereno Watson in the botany volume— after which Cooper moved
to California.

The American Naturalist was founded in 1867 by four pupils of
Louis Agassiz led by Frederic Ward Putnam (1839-1915), who in
turn would be the teacher of David Starr Jordan. Putnam published
on the fishes of Salem Harbor at 16 and became a most important
ichthyologist of the 19th century. Described as “a wiry, nervous,
black-haired, black-eyed, intense little fellow,’ Putnam guided mu-
seums in Salem, Boston, and Cambridge, and for 25 years led the
American Association for the Advancement of Sciences.!! E. G.
Conklin related in his “Early history of the American Naturalist”
that, from its first printing of 250 copies, ““Putnam was indefatigable
in his work for the Naturalist.”’ Later in our story we shall see how
Putnam also attended the birth of the Naturalist’s Directory.

About this time, San Francisco’s Overland Monthly, the “true
pulse of a pioneer society’’, as Franklin Walker!* characterized it,
was on Sale in cities and towns. The cover of the first issue carried
a grizzly bear, his feet planted on the iron rails, a snarling muzzle
turned towards the oncoming westbound locomotive. He repre-
sented the independent spirit of the West. Anton Roman, its man-
ager, had arrived in the gold mines in 1850, peddled books to the
miners, then opened a bookstore in San Francisco in 1859. He
published the Overland Monthly for eight years. Besides short sto-
ries—some gained lasting fame: for example, Bret Harte’s Luck of
Roaring Camp appearing in installments—scientific articles were
featured. Andrew Jackson Grayson told about his Mexican jornadas;
Josiah Whitney and Clarence King wrote their rockbound opinions;
C. C. Parry,!* and separately, J. G. Lemmon, on their botanical
excursions. In 1870 Anton Roman published Nicholas Bolander’s
Catalogue of plants growing in the vicinity of San Francisco.

BEHR, DAVIDSON, AND HARKNESS

A newcomer escaped from the German Revolution of 1848, and
arrived in San Francisco in 1850 when wild columbines (Aquilegia
truncata) grew on Telegraph Hill.'* Dr. Hans Herman Behr joined
the Academy in 1854 and served as its vice president from 1864 to
1904. When the California College of Pharmacy opened in 1872 he
began teaching botany and in 1884 published a Synopsis of the
Genera of Vascular Plants in the Vicinity of San Francisco, with an
Attempt to arrange them according to evolutionary principles,'> for
his botany students. In 1896, he published his “Botanical rem-
iniscences of San Francisco” in Jepson’s Erythea. Hearty, generous,
witty, Dr. Behr was popular in the Bohemian club, an association
that Prof. Setchell later enjoyed.

6 MADRONO [Vol. 34

Dr. Behr found a friend in George Davidson. Born in 1825 in
Nottingham, England, Davidson spent his boyhood in Philadelphia.
He entered the U.S. Coast Survey in 1845, preparing charts that
guided the Gold Rush vessels that were soon to converge on the
coast. He prepared successive editions of the Coast Pilot, known to
the mariners as ““Davidson’s Bible’’. For sixty years he was the best
known scientist on the Pacific Coast—a member of over forty sci-
entific and learned societies. Fame is fickle. Davidson is not men-
tioned in the Dictionary of Scientific Biography. His 6.4 inch tele-
scope in Lafayette Park, San Francisco, was the first observatory in
the state, and he was pivotal in the history of the observatory funded
by James Lick. His library was so rich that Robert Louis Stevenson
was advised to check, on his visit to San Francisco, whether his wanted
books on the South Pacific were in Davidson’s library. They were.
We do not think of this astronomer and geographer as a botanist,
but both Asa Gray and Professor Greene commemorated Davidson
for his plant collections.

Davidson was president of the California Academy of Sciences
from 1871 to 1887. If you peruse the records of the Academy in
that decade you will note two alliances: The Harkness-Brandegee
vs. the Davidson-Behr alliance. Harvey Willson Harkness was born
in 1821 in Pelham, Massachusetts, took his medical degree in 1847,
and then fled to California where he practiced first at Bidwell’s Bar.!°
After his retirement in 1869, he devoted his years to fungi. Both M.
C. Cooke and P. A. Saccardo named genera for Harkness, who
described 108 new and old species of hypogeous fungi. He visited
Sonoma County for truffles during the years Mrs. Curran (later Mrs.
Katherine Brandegee) was botanizing there. David Starr Jordan
characterized Harkness as “‘a physician of prominence’’, that he and
Davidson “were vigorous and rather intolerant, a combination of
qualities which was not rare in pioneer days.’’!’ Davidson and Hark-
ness were born of different temperaments and their discord was
fueled by an Academy problem. Davidson’s comrade, James Lick,
had endowed the Academy handsomely and the funds were invested
in a large office building on Market Street, where the Academy’s
museum occupied cramped quarters in the rear. Harkness, evidently
joined by the Brandegees, wished to devote the entire building to
the museum. Davidson proposed that an income property be de-
veloped and the Academy move to another site. He had been pres-
ident of the Academy since 1871, but was defeated in 1887 by Dr.
Harkness in a vote of 80 to 102. Tensions flared and by 1891 the
supportive Brandegees commented that Dr. Harkness has been “‘sac-
rificing to the Academy’s interest and advancement all his time,
attention and energy.”’ Soon after Harkness’ victory, an argument
arose with Dr. Behr at one of the Academy directors’ meetings.
Unable to stand the force of Behr’s points, Harkness shouted, “‘Oh,

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY y

go to hell!’? Behr answered politely, ““After you, my dear sir.”” David
Starr Jordan succeeded Harkness as president, but the rancor per-
sisted. Within a decade, the three combatants had left the stage: first,
Harkness in 1901, Behr in 1904, and finally, Davidson died in 1911
one year after he received an honorary LL.D. from the University
of California.

CASSINO, ORCUTT, LEMMON, AND MERRIAM

Whereas the American Naturalist stimulated field biologists by its
articles of discovery, the Naturalist’s Directory matched collectors
who might live on the opposite edges of the continent. For example,
fern enthusiast John Gill Lemmon living in Sierra Valley might
locate a fellow collector at the foot of the Adirondacks through the
columns of the Directory. Although we associate the Naturalist’s Di-
rectory with Samuel Edson Cassino, who first published it in book-
form in 1877, it was Frederic W. Putnam, one of the founders of
the American Naturalist, who initiated the idea of a directory. Put-
nam first listed 402 persons by “department of study”? (Geology,
etc.) and invited those and others to cooperate in a “Naturalist’s
Directory’’, as a feature of the Proceedings of the Essex Institute in
1865. Putnam managed and supported the “Directory” as part of
his Salem Press, which published American Naturalist, until he dis-
posed of the Directory to Cassino (1856-1937), then 22 years old.
As a lad, Cassino had collected moths in company with entomologist
A. S. Packard, author of a book on insects (also published by the
Salem Press). Cassino’s Naturalist’s Directory as it came to be known
appeared as his enterprise at intervals through the 30th edition (1936);
under various managements it has continued to the present 44th
edition (1985).'® The San Franciscans Kellogg, Behr, and Bolander
appeared in the first Cassino Naturalist’s Directory of 1877. Other
Californians included Mrs. Ellwood Cooper of Santa Barbara and
J. G. Cooper of Haywood, Alameda Co. By 1880, the “‘California”’
section of the Directory listed 140 names.

Daniel Cleveland came to San Diego in May 1869 at the age of
31 to carry on his law practice. In 1874, he was one of the founders
of the San Diego Society of Natural History and soon was corre-
sponding with Asa Gray, who named a delightfully fragrant sage,
Audibertia clevelandi, which was collected in the mountains near
Potrero east of San Diego. Parry visited Cleveland in 1882. Probably
Parry and C. G. Pringle encouraged another San Diego naturalist,
Charles Russell Orcutt, who was only 21, to launch a modest “‘pop-
ular monthly”, the West American Scientist, in 1884. By 1890, his
Scientist was identified on the cover as the “‘official organ”’ of the
Society, selling for 10 cents per number or one dollar for the year.
Its influence was wide indeed; contributors included Parry, Edward

8 MADRONO [Vol. 34

Palmer, and T. D. A. Cockerell.'!? San Diego, after San Francisco,
continued as a Pacific Coast center of natural history into the 20th
century, although Orcutt’s West American Scientist remains a bib-
liographic relic.

The “‘good Doctor Parry”’ made influential friends. Among them
were Leland Stanford and Charles Crocker, railroad builders who
provided a pass for Parry and other botanists in the West. Parry
likely spoke for John Gill Lemmon, who with his wife was thereby
enabled to exhibit at the World’s Industrial and Cotton Centennial
Exposition that opened in New Orleans on 16 December 1884. The
Southern Pacific Railroad had completed its route eastward from
California to New Orleans in 1883. The booth at that Exposition
may well have looked like a photograph labelled in Lemmon’s hand
“‘Lemmon Herbarium and its occupants’’, dated 23 June 18957° (Fig.
2). In this photograph, decorated with the Darlingtonia standing at
the lower right, the portfolio on the floor, and behind the seated
““occupants’’, is their precious curtained herbarium, the sheets rest-
ing as bolts of fabric in a department store. At the lower left we
speculate that the Botany of California (in two volumes) or the Pacific
Railroad Reports occupy the lowest shelf. Certainly Sara Allen Plum-
mer Lemmon “energized his life’, and was his helpmate in all his
enterprises.*! Jepson knew and understood the Lemmons and in his
characteristic esoteric way, fittingly labelled a Lomatium described
from their collection the ‘“‘Love Parsnip’’. The Naturalist’s Directory
of 1905 reads ‘““Lemmon, Mrs. J. G., Artist and Explorer, Lemmon
Herbarium, 5985 Telegraph Ave., Oakland, Calif. Bot., Cryptogams,
Eth[nology]’, concluding with an asterisk indicating that Cassino
had heard from the person since the last Directory edition. The next
entry, “Lemmon, Prof. John Gill. . .’’ concludes “‘Bot., Mic[roscopy],
Forestry, Zool. C[ollection]’’ and the asterisk as above. Sara survived
John and attended the natal meeting of the Society called by Jepson
in 1913, but her name does not appear in the list of members; she
died in 1923 at the age of 87.

Although Clinton Hart Merriam did not join the CBS, his “Life
Zones”’ and the concepts it engendered continue to provoke discus-
sions in the laboratory and around the campfire. Merriam knew the
Adirondacks and then collected in the Yellowstone with the Hayden
Survey when he was 16.7? His Bureau of Biological Survey, funded
under various names, figured prominently in the West from 1885
to 1940. He organized the Death Valley Expedition of 1891 and its
botanist, F. V. Coville, then 24, named the endemic poppy Arcto-
mecon merriami. Merriam’s Life Zones with “‘Lower Sonoran’’,
‘“‘Upper Sonoran’’, etc., up to “‘Arctic-Alpine’’, were parts of the
language that Merriam’s boys adopted in the North American Fau-
nas. Forty years after Merriam first proposed them, Jepson intro-
duced his Manual with a review of “‘Life Zones’’. Merriam began

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY 9

Fic. 2. The Lemmon Herbarium and its occupants (J. G. and S. P. Lemmon),
Oakland, CA (23 Jun 1895).

mapping the Indian tribes of California and their uses of plants in
1904. He organized the Harriman Alaska Expedition, advised on
its scientists, and finally edited its massive reports. He worked with
his friend John Muir in conservation missions.

KATHERINE BRANDEGEE’S BOTANICAL CLUB

Competition is a growth-promoting substance for societies, as for
American business. It is no misconception that the California Bo-
tanical Society had been delivered by Katherine Brandegee, M.D..,
assisted by Dr. Harkness in the “herbarium room”’ of the California
Academy of Sciences on 7 March 1891. For the physicians’ report
read the journal Zoe,’> itself brought into the world by the Bran-
degees in the 1890’s. The Club had enrolled 99 charter members in
the first month, including such names as Parish, Palmer, Cleveland,**
Hasse, Sonne, Shockley, Carl Purdy, and John McLaren, Mrs. Ell-
wood Cooper of Santa Barbara,?> and Mary Elizabeth Parsons—
difficult to name a California enthusiast not in the Club. Prof. Dudley
was president of the Club in 1893 and Parish of San Bernardino,
vice president. Strong sentiments against Harkness and, by associ-
ation, against Katherine Brandegee, are seen in Jepson’s writings.
In his admiring sketch of Edward Lee Greene published in 1918—
Kate was still alive—Jepson related his first visit to the Academy
when “‘an unkempt woman” with an “‘unpleasant voice”’ introduced
Dr. Kellogg and Mr. Greene, whom she then labelled “‘a very won-
derful man’’. Someday a biography of Willis Linn Jepson (Fig. 3)

10 MADRONO [Vol. 34

' i

WILLIS LiInNUe a Sor
i
‘

Fic. 3. Willis Linn Jepson. Portrait by Peter Van Valkenburgh (Feb. 1927). “‘Tel-
escope Peak; Panamint Range (11,000 ft.), looking toward Sierra Nevada.”

will be written. A chapter may well be titled ““Bold Kate, Jepson’s
“Viper Parsnip’ ’’.?°
SENATOR STANFORD’S DR. STILLMAN AND PROFESSOR DUDLEY

Just as during the 1850’s, the 1890’s were bustling meristematic
years for botany in the Bay Region. Growing points included Stan-

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY 11

ford University, energized by President David Starr Jordan who
brought William Russel Dudley to teach botany. Do not forget that
Senator Stanford’s private physician, Jacob Davis Babcock Stillman,
had collected plants for John Torrey and that Stanford, on behalf
of the Southern Pacific, befriended Parry with railroad passes for
his botanizing junkets. Stanford had supported botany in small but
important ways before Dudley came from Cornell. The Timothy
Hopkins Seaside Laboratory, a “perfect paradise for the marine
biologist’’ opened at Pacific Grove. Professor O. P. Jenkins?’ of
Stanford was the director. Professor Setchell later found it a good
collecting site for algae.

BRITTON-ABRAMS ALLIANCE

Far from California, but influencing its botanical future, was the
founding of the New York Botanical Garden in 1891. The impact
of its director and guiding spirit, Nathaniel Lord Britton, was to
grant wide acceptance to the so-called American Code of Nomen-
clature that sought to enforce among botanists strict priority for the
adoption of plant names. Professor LeRoy Abrams of Stanford, who
succeeded Dudley, adopted the Brittonian code for his writings on
the flora of the Pacific Coast. As David Keck has mentioned,’
Jepson followed Harvard in adopting the policy of Kew in accepting
plant names established by wide use among authors. Jepson viewed
Abrams with his important //lustrated Flora of the Pacific States as
a competitor and, in a way, this impelled Jepson to establish a
California Botanical Society to advance his position of leadership
in the botany of the state. Again, competition played its part here,
as with Asa Gray versus Alphonso Wood in his bid for the market
in introductory botany books for the East.

JOHN MUIR’S MEETING WITH JEPSON

John Muir’s interest in learning plant affinities beyond the folk
names was not acquired first in California—he had been botanizing
in the savannahs of the South—but was fostered when he took the
Sierras to heart. It was the Yosemite and the exploration of the great
Tuolumne Canyon that he wrote about as early as 1871 in, to be
sure, the Overland Monthly! Muir’s journals demonstrate that he
was acquainted with the botany of the Pacific RR Reports. In 1871,
he wrote, “I made my camp in a grove of Williamson spruce” — that
was an early name for the Mountain hemlock. There are dozens of
such botanical identifications. Remember that Albert Kellogg, Galen
Clark, and artist Billy Sims were with Muir as they camped on the
way to Mount Whitney. He collected the alpine cinquefoil, named
by Asa Gray Ivesia muiri, on Mount Hoffmann. In 1877, Muir was
with Gray and Joseph Dalton Hooker on Mount Shasta. He was a

12 MADRONO [Vol. 34

guest at the Bidwell home, Rancho Chico, when the majestic Hooker
Oak that C. C. Parry wrote about in the Overland Monthly”? was
spreading its canopy. In 1888, Muir camped with Parry for more
than a week on the shores of Lake Tahoe. Afterwards, Muir would
recall, “I had him all to myself— precious memories.”

One of the roots of this Society was Jepson’s meeting with John
Muir in the founding of the Sierra Club. When and where Jepson
first met Muir, I cannot say, but on 28 May 1892, they joined
Joachim Henry Senger, a professor of German, William Dallam
Armes,*° who taught American literature at Berkeley, and William
E. Colby,*! who was later prominent in the Sierra Club, at the office
of attorney Warren Olney in San Francisco, to draft the articles of
incorporation of the Sierra Club. Professor Armes, a bachelor, had
been teaching at Berkeley since 1882 and was living in the Faculty
Club. Later, Armes edited Joseph LeConte’s Autobiography (1903)
and published a critique of More’s Utopia (1912). Clearly, the two
Berkeley professors were congenial friends of Jepson.

HARRIMAN ALASKA EXPEDITION, FARLOW AND SETCHELL

Alaska took the front stage in 1899 with businessman Edward H.
Harriman in the lead role and a supporting cast of 25 scientists. The
Harriman Alaska Expedition was a success that produced multi-
volumed reports edited by Merriam. The only University of Cali-
fornia faculty man to accompany the Expedition was William Ritter,
although Jepson and Setchell met the Harriman party that summer.

William Albert Setchell had arrived in Berkeley to succeed Pro-
fessor E. L. Greene in 1895. That year, the New England Botanical
Club had been founded by Setchell’s mentor, W. G. Farlow, and six
associates and in four years the Club numbered 46 gentlemen “of
leisure but not of idleness”’.** Farlow had been an assistant to Asa
Gray in 1871 and later left his stroma of mycologia and an endow-
ment to Harvard for cryptogamic botany, with the stipulation that
“no part thereof shall be used to pay for lectures or instruction of
any kind.’’?> Setchell has related how a small separate expedition to
Alaska had been planned in 1898, evidently while the Harriman
Expedition was being planned.** Four of the members of the Uni-
versity of California had made reasonably definite arrangements to
attempt some limited botanical exploration in the same general field,
and especially had set their eyes on the region of the Island of
Unalaska, at the southeastern corner of the Bering Sea. Besides
Setchell and Jepson, then Assistant Professor of Botany, Anstruther
Abercrombie Lawson, who had graduated in botany in 1897, and
Loren Edward Hunt, Instructor in Civil Engineering, participated in
the expedition. Jepson recorded in his Field Book that Hunt was
““soing along as supercargo and handy member of the party.” Jepson

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY 13

noted that the Harriman Expedition arrived at Unalaska 12-13 July
1899 and on that occasion he met several members of the Expedition
including Merriam, Coville, Muir, and John Burroughs. Anstruther
Lawson later became Professor of Botany at University of Sydney,
and studied plant embryogeny, Psi/otum, and gymnosperms. He was
at Stanford from 1900 to 1906 where he wrote the “‘Life history of
Sequoia’’. Anstruther Lawson was brother of the witty combative
geologist of Berkeley, Andrew Cowper Lawson.

JEPSON’S CALIFORNIA BOTANICAL SOCIETY

Jepson confided in his 1938 anniversary address to the CBS that
the ‘“‘idea of a Society was definitely conceived in the year 1902”’.°°
It is easy to see how the stimulating meetings with outside scientists
and the comraderie that then had existed with Setchell would have
given rise to the merit of such a Society, but then Jepson added,
““some one may note the lapse between that date and the year 1913
when the Society was founded.’’ He then suggested that a certain
disassociation of the Stanford and Berkeley botanists may have been
responsible for the delay. I suggest a certain competitive spirit that
existed between the botanists of Stanford and Berkeley, and differ-
ences, for example, in the adoption of opposing codes of nomen-
clature as practiced by Abrams and Jepson, may have dissociated
them. Then, too, the conspicuous vigor of the California Botanical
Club and its associates in San Francisco, all contributed to Jepson’s
compelling interest in founding the CBS directed from Berkeley.

Two Berkeley faculty members were prominently attending the
birth of the Society. William Frederic Badé was temporary chairman
of that founding meeting of the Society on 12 April 1913, and then
was elected second vice president. Badé, an archeologist, linguist,
and literary executor of John Muir had edited Muir’s letters.*° He
was a distinguished figure. Curiously, Jepson did not mention Badé
in his Annual Dinner address of 1938, although he had died only
two years before. By contrast and in lengthy prose, Jepson praised
Cornelius Beach Bradley, professor of rhetoric at Berkeley who had
missed being a full-time botanist, in Jepson’s words, “‘by only a
narrow margin.”

Then on the same April day in 1913 on which the Society was
being born at the Oakland Public Museum, another meeting was
under way on the Berkeley campus. In preparation for the coming
Pan Pacific International Exposition of 1915, the original AAAS
founded and based in the East determined that it should be repre-
sented by a Pacific Division. Botanist Daniel Trembly MacDougal
prompted that action. A committee of twenty scientists met to in-
augurate a Pacific Division with W. W. Campbell in the chair.*’
Some of the other Berkeley faculty present were E. W. Hilgard, C.

14 MADRONO [Vol. 34

A. Kofoid, A. L. Kroeber, A. C. Lawson, Ritter and Setchell. Pro-
fessor Setchell was enrolled a charter member of the CBS, although
he evidently was not present at the Oakland meeting.

INTERNATIONAL PHYTOGEOGRAPHICAL EXCURSION

The International Phytogeographical Excursion (IPE) of 1913, when
European and Eastern botanists visited California after field trips in
the Rocky Mountains and Crater Lake, was another root nourishing
the growth of botany in the Bay Region.** Although the IPE took
place five months after the founding of the Society, preparations had
been going forward before the April meeting in Oakland. Botanists
from Amsterdam, Copenhagen, Berlin, Munich, Zurich, and Cam-
bridge, England, met their colleagues from Berkeley, San Francisco,
and Stanford, later to be joined by MacDougal from Tucson and
Samuel Bonsall Parish from San Bernardino. A new language of
flowers was heard when the excursionists discovered that “‘Sail-ix’’
grows in America, “‘Sall-ix”’ in Europe. Whatever animus may have
separated faculties was lost for the days the excursionists tramped
California’s chaparral and shared their experiences. The IPE was a
high point for Jepson. He related some details in the first issue of
Madrono, which appeared three years after the founding of the So-
ciety.

On Friday 12 September 1913, Professor Jepson presided at the
dinner for the IPE. His closing words were:

““Now there arises a school of botanists, the plant ecologists, who
are leading us back to the fields and woods, taking with them the
experience of all other schools, and in addition making important
use of the observations of the old-time naturalists. California 1s a
glorious field for such work, and we welcome them here to help us
appreciate our own flora, and to help Californians to an appreciation
of it.”

Montesquieu was right: Jepson shaped the Society but the Society
in turn shaped Jepson’s dream. On one of my 5 x 8 half-sheets for
15 March 1937, I wrote, “I learn tonight of a boyhood dream or
aircastle, which Dr. Jepson himself now says was ‘preposterous.’*?
In Vaca Valley, his boyhood home, stood a two-story brick building
occupied by a small college dating from the Gold Rush days with
something of a classic demeanor, a courtly flavor. This stood on a
low hill with creek bed beside it deeply filled with rich alluvium. As
a lad he envisioned devoting the structure to an herbarium and of
surrounding it with a botanic garden. As a boy he did not have

9°99

money to think of such a reality but he had the ‘desire’.

ACKNOWLEDGMENTS

My thanks for assistance from Dr. Lawrence Heckard, Jepson Herbarium; Barbara
Lekisch, Librarian, Sierra Club; from Wayne R. Ferren, Jr., Editor, Madrono; two

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY 15

Tulanians: Milton G. Scheuermann, Professor of Architecture, and Dolores Gunning,
Department Secretary. First and last, my wife Nesta.

NOTES

A complementary essay to this subject is by J. Ewan (San Francisco as a mecca
for 19th century naturalists with a roster of biographical references to visitors and
residents, Jn Century of progress in the natural sciences, 1853-1953. pp. 1-63. Cal-
ifornia Academy of Sciences, San Francisco, 1955), hereinafter cited as Century.

1. Carlin, Eva V., ed. 1898. A Berkeley year. A sheaf of nature essays. pp. 61-64.
Women’s Auxiliary of the First Unitarian Church, Berkeley.

2. William S. Ferguson, Greek Imperialism, 1913, quoted in: Charles P. Curtis, Jr.
and Ferris Greenslet. 1962. Practical Cogitator. ed. 3. New York. p. 134.

3. G. M. Trevelyan, ibid. p. 131.

4. Carew, Harold D. 1928. High priest of Flora. A glance at the life and works of
Willis Linn Jepson, California’s foremost botanist. Touring Topics 20(12):32-
34, 50. p. 31.

5. Ewan, J. 1973. William Lobb, plant hunter for Veitch and messenger of the Big
Tree. Univ. Calif. Publ. Bot. 67:1—36. p. 7. For the ““smouldering resentment” of
California botanists see: A. H. Dupree. 1959. Asa Gray. Harvard Univ. Press.
pp. 395-396.

6. See W. L. Jepson (Flora Calif. 2:6. 1936). Volume 1 of Botany California (1876)
was reprinted with additions in 1880, the year vol. 2 was published in a smaller
printing because of reduced funding.

7. Yale Univ. Press. 1930. Jepson assisted Farquhar with plant identifications. For
a charming vignette of Brewer, ““omnivorous devourer of facts in every field’,
with a “‘tentacular mind’’, see: Rudolph Schevill (Recollections of a golden age.
Pittsburgh, pp. 174-175, 1985). Ian Jackson provided this trail sign.

8. For a consummate analysis see: Max Meisel. 1929. Bibliography of American
Natural History. New York. 3:189—220; and for critical commentary: I. M. John-
ston. 1943. J. Arnold Arbor. 24:237-—242.

9. Dupree, A. Hunter. op. cit. pp. 327-328. John Torrey was present at the Harvard
occasion. The herbarium contained ‘“‘at least 200,000 specimens’’, valued es-
pecially for the types.

10. Barnhart, John H. 1918. Historical sketch of the Torrey Botanical Club. Mem.
Torrey Bot. Club 17:12-21.

11. Conklin, E. G. 1944. Early history of the American Naturalist. Amer. Naturalist
78:29-37; Ralph W. Dexter. 1982. F. W. Putnam as secretary of the A.A.A.S.
(1873-1898). Essex Inst. Historical Collections 118:106—118, especially 109.

12. Walker, Franklin. 1939. San Francisco’s literary frontier. Knopf, New York. pp.
256-283, especially 279-280.

13. See: J. Ewan. 1950. Rocky Mountain Naturalists. Univ. Denver Press. pp. 34—
44. Portrait and suppl. refs. Jn J. and N. Ewan. 1981. Biographical dictionary
of Rocky Mountain naturalists. Utrecht. pp. 168-169.

14. Zoé 2:3. 1891.

15. Century, pp. 12 and 43. A long overdue biography of Behr, a “‘Forty-eighter”’
and friend of Alice Eastwood, would profile a half century of California history.

16. T.S. Brandegee’s biography of Harkness (Zoé 2:1—2 and portrait, 1891) was clearly
an apology that appeared ten years before the death of Harkness.

17. Century, p. 37.

18. 44th edition of the Naturalist’s Directory (Flora and Fauna Publs., Gainesville,
FL, 1985) includes a portrait of S. E. Cassino (p. vi) and the first history of this
important chapter in American natural history by Ralph W. Dexter (pp. 1-7)
and additional notes by Ross and Mary Arnett. References kindly supplied by
Dale Johnson (MO).

19. For references on Theodore Dru Alison Cockerell (1866-1948) see: J. Ewan and

Z0:

yal

22:

23%

24.

Za.

Z6:

PRIS

28.

Pao)

30.

MADRONO [Vol. 34

N. Ewan. 1981. op. cit. pp. 44-45. Also see: W. A. Weber. 1976. Theodore D.
A. Cockerell. Colorado Assoc. Univ. Press. Most recent account of C. R. Orcutt
(1864-1929) by Helen DuShane (Baja Calif. Travel Series, 23, Dawson’s, Los
Angeles, 1971) extends Jepson’s account (Madrofio 1:273-274, 1929). Lee W.
Lenz (Marcus E. Jones. Rancho Santa Ana Botanic Garden, Claremont, CA. pp.
52-57, 1986) reviews the Orcutt-Parry-Pringle-Jones episode.

J. G. Lemmon was certainly the author of the anonymous “Catalogue of the
plants and paintings of the Lemmon Herbarium” that appeared in Charles B.
Turrill (Catalogue of the products of California exhibited by the Southern Pacific
Company .. . Nov. 10, 1885—April 1, 1886. New Orleans, pp. 55-62, 1886).
Jepson annotated this unpublished record (Fig. 2) as an “interesting photograph
because it tells so much of what Lemmons were personally and botanically”’ and
that it was communicated by Dr. Rimo Bacigalupi, ca. Sep 1932. For Lemmon
references see Century, pp. 23-24, 54, Ewan, op. cit. (1981, p. 132) and F. S.
Crosswhite (Desert Plants 1:12—21 and portraits, 1979).

Jepson, W. L. 1946. Dict. Amer. Biog. 11:162; J. Ewan. 1944. Amer. Mid.
Naturalist 32:513-518.

Clinton Hart Merriam (1855-1942) was the “‘central figure in a dynamic era
connecting the pioneer period of exploration with the present time of experi-
mentation and interpretation” (Wilfred H. Osgood. 1945. Biogr. Mem. Natl.
Acad. Sci. USA. 24:1-57, with bibliog. of his publs.). ““Merriam, Vernon Bailey,
and the cyclone trap fleshed out mammalogy as a science in America” (p. 9).
Zoé 2:93-96. 1891; Century, pp. 36-37. Alice Eastwood presided over the Cal-
ifornia Botanical Club for more than sixty years from her first meeting of 26 Apr
1892 (Leafl. W. Bot. 7:59, 1953). A useful account of K. Brandegee is in: Notable
Amer. Women 1:228-229, 1971.

Daniel Cleveland (1838-1929) and associates are described in: Elizabeth C.
MacPhail. 1976. Kate Sessions, pioneer horticulturist. San Diego Hort. Soc., San
Diego. pp. 44 and 69; also see: Century, pp. 37 and 46.

Sarah Paxen Moore Cooper (died 1908) married Ellwood Cooper (1829-1918)
in August, 1853 (teste J. H. Barnhart). Cooper was a pioneer in eucalyptus
cultivation in California. His friend Lorenzo Gordin Yates (1837-1909) who
came to Santa Barbara in 1881, became president of the Santa Barbara Natural
History society that was founded in 1876 (Charles L. Camp. 1963. J. Soc. Bibliogr.
Nat. Hist. 4:178-193). Also see: Clifton F. Smith. 1976. Flora of the Santa Barbara
Region, California. Santa Barbara Mus. Nat. Hist., Santa Barbara, pp. 42 and
59. He tells me (litt. 1972) that J. G. Lemmon and Henry Bolander visited the
Cooper ranch near Goleta.

Jepson portrayed K. Brandegee (Newman Hall Review 1:24, 1918). Type of
“Viper parsnip” (Leptotaenia anomala Coult. & Rose) was taken by Katherine
Curran near Carbondale, Amador Co., CA (W. L. Jepson, Flora Calif. 2:634,
1936). Other accounts: Century, p. 32-33; F. S. and C. D. Crosswhite (Desert
Plants 7:128-139, 158-162, and portraits, 1985).

Zoé 4:58—63 and pl. 26. 1893. Oliver Peebles Jenkins (1850-1935), physiologist
and ichthyologist was noticed in: David Starr Jordan. 1922. Days of a man.
New York. 1:399-—400.

Keck, D. D. 1948. Place of Willis Linn Jepson in California botany. Madrono
9:223-228.

Parry, C. C. 1888. Rancho Chico. Overland Monthly ser. 2. 11:561-576. Pho-
tographs of “‘Sir Joseph Hooker Oak” on p. 565 and in: Rockwell D. Hunt. 1942.
John Bidwell. Caldwell, Idaho. opp. p. 273. Parry is noticed on pp. 209, 211-
212, 278-279.

“Organization and early conservation activities of the [Sierra] Club” included
W. Olney, Joseph LeConte, J. H. Senger, W. D. Armes, and C. B. Bradley,
according to W. F. Badé (Life and Letters of John Muir, Boston, 2:256, 1924),
but Jepson was not mentioned. Also, Holway R. Jones (John Muir and the Sierra
Club, Sierra Club, San Francisco, pp. 7-9, 1965) did not mention Jepson. Jepson,

1987] EWAN: CALIFORNIA BOTANICAL SOCIETY Ia

Si,

32.
33:

34.

35.

36:

37.
. Madrono (1:12-18, 1916) includes a photograph of 14 identified participants

39:

however, was present at the meeting of 28 May 1892, according to Linnie Marsh
Wolfe (John of the mountains: the unpublished journals of John Muir, New
York, p. 299, 1938; and Son of the Wilderness: the Life of John Muir, New
York, pp. 254 and 360, 1947). Professor Senger taught German at Berkeley 1886-
1913 and died in 1926. “‘Billy’’ Armes was evidently a popular professor, judging
by comment of Loye Miller (Lifelong Boyhood. Berkeley, p. 58, 1950). Armes
died in 1926.

William E. Colby (1875-1964) a lawyer and native of Benicia, California, joined
the Sierra Club in 1898 and was active (president 1917-1919) to his death. Warren
Olney (1870-1939), a lawyer and native of San Francisco, is quoted by Badé (op.
cit. 2:376-377). Mrs. Edward T. Parsons, Muir’s friend, of that letter, was a
charter member of the California Botanical Society.

Williams, E. F. 1899. New England Botanical Club. Rhodora 1:37-39.
Setchell, W. A. 1927. William Gilson Farlow. Biogr. Mem. Natl. Acad. Sci. USA.
21(4):1-22. portrait.

Setchell, W. A. 1907. Univ. Calif. Publ. Bot. 2:309-311. Note on A. A. Lawson
(1870-1926) occurs in: Francis E. Vaughan. 1970. Andrew C. Lawson. Glendale,
CA. pp. 38-39. Loren Edward Hunt, who contributed photographs to Jepson’s
Silva of California, accompanied him on a field trip to Blue Lakes, Lake Co., 16
Jul 1897, when the type of Godetia amoena f. huntiana Jeps. was collected.
Jepson, W. L. 1938. Viae felicitatis: the beginning years of the California Botanical
Society. Madrono 4:276—286. see p. 282.

William Frederic Bade (1871-1936), archeologist, professor, and acting president
of the Pacific Theological Seminary, later Pacific School of Religion, Bancroft
Way, Berkeley, and outdoorsman, “one of the most able and devoted preser-
vationists’”” (James Mitchell Clarke. 1980. Life and adventures of John Muir.
Sierra Club, San Francisco. pp. 304 and 320). Also see C. C. McCown in: Dict.
Amer. Biog. suppl. 2, 1958. Cornelius Beach Bradley (1843-1936) was a mis-
sionary and professor of English, UC Berkeley (1882-1911).

Robert C. Miller (Science 108:220—221, 1948) lists members of the committee.

taken at Mariposa Grove of Big Trees. Also see G. E. Nichols (International
Phytogeographic Excursion in America. Torreya 14:55-64, 1914); A. G. Tansley
(New Phytol. 12:322-336, 1914; and 13:30-—41. 83-92, 1914); the F. E. Clements
records in Univ. Wyoming archives, Laramie; and a note by Paul B. Sears (Plant
Ecology. Jn J. Ewan, Short history of botany in the United States, Hafner, New
York, pp. 130-131, 1969).

During my four years with Jepson (see his Flora Calif. 2:10, 1936) I recorded
conversations and events on half-sheets.

(Received 27 Jun 1986; revision accepted 1 Oct 1986.)

ANNOUNCEMENT

JOINT ANNUAL MEETING OF
AIBS, ASPT, BSA, AND ESA

9-13 August 1987, Ohio State University

Two symposia in botany will be presented this year: one on the Generic
Concept, and one on the Reproductive Ecology of Aquatic Angiosperms.

INFLORESCENCE ARCHITECTURE OF
EUCNIDE (LOASACEAE)

LARRY D. HUFFORD
Department of Botany,
University of California, Berkeley,
Berkeley 94720

ABSTRACT

Terminal inflorescences and axillary flowers have been reported in recent revisions
of Eucnide (Loasaceae). Developmental studies show, however, that axillary flowers
are not present. All flowers are terminal: the initial shoot axis terminates in a flower,
and lateral branches that terminate in flowers after producing only one or two leaves
arise from the distal nodes of each initial and renewal axis. The branching pattern
in the inflorescence region of each axis is complicated by the apparent displacement
of subtending leaves outward onto lateral floral branches during their extension.
Renewal axes that first arise in the nodes subjacent to lateral floral axes reiterate the
pattern of the initial axis.

Recent systematic treatments of Eucnide (Loasaceae) have largely
underestimated the architecture of the inflorescence. Waterfall (1959)
used floral position to separate “‘two natural, but somewhat inter-
grading series in the genus.” His first series, comprised of E. bar-
tonioides, E. xylinea, and E. urens, was characterized by solitary
flowers in leaf axils. The second series had “terminal inflorescences
more or less developed.” The most recent revision by Thompson
and Ernst (1967) distinguished three sections in Eucnide (including
Sympetaleia at the sectional level; Waterfall did not consider Sym-
petaleia to be congeneric with Eucnide). Thompson and Ernst did
not retain the informal division based on floral position proposed
by Waterfall, and inflorescence data does not appear to have had a
major role in their sectional circumscriptions. They observed ter-
minal inflorescences of a few flowers in most species and noted
axillary flowers in more than half of the species. The only inflores-
cence data given for the new species E. durangensis (Thompson and
Powell 1981) was in the Latin description: “‘Jnflorescence pauci-vel
multiflorae’’.

My investigations show that our current knowledge of inflores-
cence morphology of Eucnide is oversimplified. The purpose of this
paper is to clarify the inflorescence architecture of Eucnide by placing
it in the context of whole plant morphology and development.

MATERIALS AND METHODS

Eucnide bartonioides (seven plants), E. cordata (four plants), E.
hirta (two plants), and E. lobata (three plants) of Thompson and
Ernst’s sect. Eucnide and E. aurea (three plants) of sect. Sympetaleia

MADRONO, Vol. 34, No. 1, pp. 18-28, 1987

1987]

TABLE 1.

HUFFORD: EUCNIDE INFLORESCENCES 19

PERTINENT COLLECTIONS OF Eucnide USED FOR THIS INVESTIGATION.

* = collections that were sources of seeds for glasshouse populations.

Species

E. bartonioides
Zucc.

E. cordata
(Kell.) Kell.
ex Curran

E. durangensis
Thompson &
Powell

E. floribunda S.
Wats.

E. grandiflora
(Groenl.)
Rose

E. hirta (G.
Don) Thomp-
son & Ernst

E. hypomalaca
Standl.

E. lobata
(Hook.) A.
Gray

E. aurea (A.
Gray)
Thompson &
Ernst

E. rupestris
(Baill.)
Thompson &
Ernst

E. tenella (1. M.
Johnst.)
Thompson &
Ernst

E.. urens Parry

Collection

Sect. Eucnide

USA, Texas: Big Bend National Park, 26 Jun 1962, Thompson
and Ernst 3283 (LA)*. MEX, Tamaulipas: Jaumave, 25 Nov
1962, Moran 10031 (LA, UC); Nuevo Leon: Cuesta de Ma-
miluque, 14 Aug 1942, Gentry 6729 (UC).

MEX, Baja California Sur: La Paz, 29 Dec 1958, Porter 106 (LA);
Sierra de la Gigantea, 9 Nov 1961, Carter 4278 (UC); Isla
Monserrate, 27 Mar 1977, Cody (Thompson 3829 seed collec-
tion) (LA)*.

MEX, Durango: Gomez Palacio, 25 Mar 1973, Johnston, Wendt,
and Chiang 10417 (LA); Torreon, 14 Aug 1973, Henrickson
12405 (LA).

MEX, Coahuila: Cuatro Cienegas, 10 Jun 1968, Lehto, Keil, and
Pinkava 5360 (LA); 4 Apr 1969, LaBounty, Lehto, and Pinkava
5927 (LA); Las Delicias, 12 Aug 1973, Henrickson 12240 (LA).

MEX, Oaxaca: Tomellin Canyon, 17 May 1894, Pringle 4645
(GH); Jayacatlan, 4 Nov 1973, Breedlove 35885 (RSA); Teo-
titlan del Carmen, 22 Aug 1975, Webster, Armbruster, and
Holstein 20036 (GH).

MEX, Jalisco: San Cristobal de la Barranca, 11 Nov 1962,
McVaugh 22140 (NY, LL); Tizapan, 30 Jun 1957, McVaugh
15108 (Thompson 3319 seed collection)*; Veracruz: Cerca de
Puenta Nacional, 13 Jan 1973, Hernandez, Dorantes, and Do-
rantes 1819 (NY).

MEX, Chihuahua: Batopilas, 15 Apr 1948, Hewitt 272 (GH);
Chiapas: Chiapa de Corzo, 24 Feb 1973, Breedlove 33828 (RSA).

MEX, Hidalgo: Barranca de Toliman, 27 Nov 1962, Moran 10048
(LA); Nuevo Leon: Monterrey, 10 Aug 1959, Waterfall 15324
(F) (Thompson 3298-6 seed collection) (LA)*; Coahuila: Car-
men Pass, 6 Aug 1978, Fryxell 3023 (ASU)*.

Sect. Sympetaleia

MEX, Baja California: Sierra de la Gigantea, 25 Nov 1953, Carter
and Kellogg 3266 (UC); Isla Carmen, 10 Mar 1960, Carter and
Ferris 3710 (UC); Idlefonas Island, 2 Apr 1962, Moran 9056
(ASU)*.

MEX, Baja California: Mexicali, 22 Feb 1960, Raven 14802 (LA);
San Estaban Island, 22 Mar 1962, Moran 8858 (LA); Bahia de
Los Angeles, 26 Feb 1963, Thorne and Henrickson 32694 (LA).

MEX, Baja California Sur: Mission Los Delores, 5 Dec 1951,
Wiggins, Carter, and Ernst 260 (LA); Sierra de la Gigantea, 31
Oct 1971, Moran 18845 (UC); Sierra de la Gigantea, 5 Nov
1971, Moran 19017 (UC).

Sect. Mentzeliopsis

USA, California: Whipple Mountains, 20 Mar 1936, Clary 2580
(JEPS); near Death Valley National Monument, 17 Mar 1984,
Hufford 1114 (UC); Trona, 18 Mar 1984, Hufford 1116 (UC);
MEX, Baja California: Okie Landing, 4 May 1966, Moran 13124
(UC).

20 MADRONO [Vol. 34

Fics. 1-2. Recaulescence in Eucnide. Fic. 1. Axillary bud (arrow) displaced onto
petiole of leaf in E. aurea. Fic. 2. After extension of the lateral branch, the subtending
leaf (P indicates the petiole of the leaf) appears to be inserted on the lateral branch
(B) derived from its axil because of extension of the common basal portion of both
leaf and branch. Arrow indicates position of insertion of the leaf (P) on the axis (S)
where it arose in E. cordata. Scale bars equal 5 mm.

were grown under glasshouse conditions in Berkeley, California.
Eucnide urens, comprising sect. Mentzeliopsis, was examined under
natural conditions (Table 1). I examined herbarium specimens of
all species, except E. xylinea C. H. Muller. Collection data for per-
tinent herbarium specimens and sources of seeds for glasshouse
populations are given in Table 1.

The term inflorescence is used in this paper in the sense of Steenis
(1963), who defined it as “‘‘the specialized fertile part(s) of an in-
dividual plant which post anthesin does (do) not participate in the
vegetative extension of the individual’, and is hence either shed or
withering away.”

RESULTS

All species of Eucnide are perennial. The shoot system is sym-
podial; each shoot axis eventually terminates in a flower. A slight
concaulescence (adnation of the pedicel of the terminal flower with
the uppermost lateral branch, Troll 1964) is common among all of
the species, although it is not a consistent feature of all individuals
of any species. Plants grown under glasshouse conditions develop
approximately five to ten leaves following the cotyledons and before
terminal flower formation. The distal-most leaves of each axis are
recaulescent (sensu Troll 1964, see also Kuyt 1981) with their ax-
illary buds. Recaulescence implies that an axillary bud is somewhat
displaced onto the petiole of the subtending leaf (Fig. 1; this con-

1987] HUFFORD: EUCNIDE INFLORESCENCES 72)

dition is also called epipetioly, Dickinson 1978). When the axillary
bud begins extension the basal region common to the subtending
leaf and axillary bud also begins outgrowth. At full extension of a
lateral floral branch, the ‘subtending leaf’ appears to be inserted on
the lateral axis rather than on the axis on which it was produced
(Fig. 2).

Sect. EUCNIDE. In glasshouse populations of FE. bartonioides, E.
hirta, and E. lobata, the three leaves and axillary buds subjacent to
the terminal flower are recaulescent (Figs. 3, 4). Of these three up-
permost nodes, lateral floral axes usually arise in the axils of the two
distal leaves (forming a dichasial inflorescence, Figs. 3, 4), although
only the uppermost node may form a lateral floral axis (a mono-
chasial inflorescence). The terminal flower and either the monocha-
sial or dichasial lateral floral axes form the inflorescence of each
shoot axis. Lateral floral axes in these species do not undergo ex-
tensive internodal elongation. Internodes of these lateral floral axes
elongate only as flowers begin to mature and then most of the elon-
gation is in the pedicel. Each lateral floral axis usually produces two
leaves (Fig. 5) or may form only one leaf (Fig. 6) and a terminal
flower. Each leaf of this primary lateral axis becomes recaulescent
with the bud in its axil. Each of these axillary buds (secondary lateral
axes) repeats this pattern of producing one or two leaves (each be-
coming recaulescent with its axillary bud) and a terminal flower (Figs.
5, 6). The lateral floral axes may be either dichasial (Fig. 5) or
monochasial (Fig. 6). Some herbarium specimens show that inflo-
rescence development changed from one condition to the other dur-
ing ontogeny. Lateral floral axes were not observed (in neither glass-
house populations nor on herbarium specimens) to convert back to
vegetative growth (1.e., back to production of more than two leaves
before terminal flower formation). This pattern of inflorescence ar-
chitecture appears to be common to all species of section Eucnide
(except EL. xylinea, which was unavailable, and EF. cordata, which is
discussed below) as ascertainable from examination of herbarium
specimens (Table 1).

Leaf form undergoes a gradual transformation in the transition to
the flowering region. Leaf laminas with a lobate margin and a cordate
base are produced in the vegetative portion of the plant. At the few
nodes proximal to the inflorescence region, where the internodes do
not extensively elongate, the leaves are smaller. Leaves lose regularly
lobed margins and cordate bases with transition into the region of
lateral floral axes. Leaves on some primary, secondary, and tertiary
axes of the inflorescence often occur as tiny, lanceolate bracts.

The recaulescent ‘node’ (involving leaf and axillary bud) on the
main shoot, which was not immediately floral, is the location of the
first renewal shoot (Figs. 3, 4) or innovation shoot (sensu Weberling

a2 MADRONO [Vol. 34

‘ MALE

Fics. 3-10. Renewal growth and inflorescence architecture of Eucnide. Fic. 3.
Monochasial renewal, one renewal branch arises from the node proximal to the lateral
flowering axes. Fic. 4. Dichasial renewal, vegetative branches arise from the two
nodes proximal to the lateral flowering axes. Fic. 5. Flower and leaf positions on
dichasial floral axes that would be located at positions indicated by solid arrows in
Figs. 3 and 4. Terminal flowers of successive lateral axes are indicated. Fic. 6. Flower
and leaf positions on monochasial floral axes that would be located at positions
indicated by solid arrows in Figs. 3 and 4. Terminal flowers of successive lateral axes
are indicated. Fic. 7. Shoot terminus of E. cordata prior to secondary branching of
the inflorescence. Fic. 8. Architecture of E. cordata inflorescence. Terminal flowers
of primary, secondary, and tertiary axes are indicated for one of the three floral
branches. Fic. 9. Architecture of shoot system of E. tenella, showing dichasial branch-
ing on left side and monochasial branching on right. Fic. 10. Architecture of inflo-
rescence of E. urens, showing displacement (concaulescence) of terminal flowers to-
ward subjacent lateral branches. A = terminal flower, B = floral branch, C = vegetative
branch, D = leaf produced by axis on which it is inserted, E = leaf produced by axis

1987] HUFFORD: EUCNIDE INFLORESCENCES ZS

1983). The subjacent node at which there is usually no recaulescence
has an axillary bud that subsequently begins renewal growth. The
sympodial growth pattern appears to be primarily, although not
strictly, dichasial (Fig. 4). Monochasial renewal (Fig. 3; especially
in E. lobata) and also pleiochasial renewal growth (usually three
branches in E. bartonioides) are common. Following transition to
flowering on a renewal axis, buds in leaf axils (although these leaves
have withered) of the previous axis also will begin growth. Outgrowth
of buds along this axis is basitonic (1.e., beginning near the base of
an axis with acropetal progression). The first few internodes of re-
newal axes elongate. As an axis nears flowering, internodes remain
largely unextended. Each renewal shoot produces only five to ten
leaves before forming a terminal flower. The inflorescence of any
shoot axis in Eucnide may be considered to be the region above the
uppermost renewal branch because this portion of the plant usually
dries and withers after flowering (it does not contribute to further
vegetative extension). Renewal growth patterns were difficult to de-
termine for species that were not grown under glasshouse conditions
because herbarium specimens seldom have enough of the plant for
evaluation.

Eucnide cordata differs from all species described above. It has
longer renewal shoots that produce more leaves before conversion
to flowering. The internode, between the lowest node with a leaf that
becomes recaulescent and a nonrecaulescent leaf, becomes elongated
when flowering begins. The next distal internode also becomes quite
elongated and effectively segregates an inflorescence region that is
more distinct than in the other species of sect. Eucnide. These pen-
ultimate internodes in the other species remain compact.

The basic architectural pattern of the flowering region in E. cordata
is similar to that described above for other species in sect. Eucnide;
however, there are some distinctions. Three lateral floral axes (Fig.
7) are produced on the initial axis and each renewal branch, rather
than the two most commonly produced in Eucnide (Figs. 3, 4). Each
of these primary lateral axes (Fig. 8) produces two or occasionally
more leaves and a terminal flower. The leaf subtending a primary
lateral axis becomes recaulescent as is common in Eucnide. The first
leaf produced by a primary floral axis often remains at nearly the
same level as the subtending leaf because the internode between
them does not elongate. An axillary bud may or may not form in
association with this first leaf. If an axillary bud does form it is floral

—

subjacent to the one on which it is inserted (recaulescence). T = terminal flower, 1° =
flower of primary, 2° = flower of secondary, 3° = flower of tertiary, and 4° = flower
of quaternary floral branches.

24 MADRONO [Vol. 34

(a secondary floral axis). This secondary floral axis produces two
leaves, but only the uppermost produces an axillary bud that is floral
(a tertiary lateral axis). The uppermost leaf of the primary lateral
axis also has an axillary bud. This again becomes a secondary floral
axis that produces two leaves, only one of which subtends a floral
bud (another tertiary lateral axis). Each tertiary axis present (there
may be two associated with each primary lateral axis) produces two
leaves (one of which has an axillary bud) and a flower. Leaves on
these floral axes, which subtend axillary buds, usually become re-
caulescent with that lateral axis when the axis begins growth. The
lateral axes in the inflorescence region of E. cordata have greater
internodal growth than is common among the other species of sect.
Eucnide.

Renewal growth in E. cordata is dichasial. Renewal branches arise
in the axils of two leaves subjacent to the floral nodes. It is usually
below the region where extensive internodal elongation occurred
concurrent with flowering. Each renewal axis may or may not be
recaulescent with its subtending leaf.

Sect. SYMPETALEIA. Eucnide aurea and E. rupestris have an inflo-
rescence pattern similar to that identified as the most common among
the species of sect. Eucnide. Each lateral floral axis of E. aurea and
E. rupestris, examined on herbarium specimens, may be monocha-
sial (Fig. 6) or dichasial (Fig. 5) (as also was true of most species of
sect. Eucnide). Eucnide tenella (Fig. 9), unlike these other species
of sect. Sympetaleia, lacks distinct lateral floral branches and renewal
branches. Axes (Fig. 9; initial axes were unavailable on herbarium
specimens) produce either one or two leaves and a terminal flower.
Each leaf of an axis usually is recaulescent with its axillary bud. Each
of these axillary buds reiterates the pattern of producing one or two
leaves (each of which usually will be recaulescent with its axillary
bud) and a terminal flower.

Unlike the other species grown under glasshouse conditions, E.
aurea is likely to have up to six flowers that open concurrently on
a single renewal axis. In the other species, usually only one or two
flowers associated with any one renewal axis were observed to be
open concurrently. The internodes of the lateral floral axes of E.
aurea also elongate to a greater extent over the period of flowering
than do the corresponding internodes in most species of sect. Euc-
nide.

Renewal growth in E. aurea appears to be primarily monochasial
(Fig. 3). As in the other species, the primary renewal axis is one of
the recaulescent axillary buds subjacent to the terminal flower. Other
reiterative lateral shoots with a vegetative phase (i.e., producing
more than two leaves before forming a terminal flower) begin growth
after the renewal shoots from an axis have begun to flower. Renewal

1987] HUFFORD: EUCNIDE INFLORESCENCES 25

growth of E. rupestris was impossible to determine from herbarium
specimens.

Sect. MENTZELIOPSIS. Eucnide urens (Fig. 10) is distinct in having
terminal flowers displaced from the notch between the two subjacent
lateral flowering axes. In some of the other species, the terminal
flowers of vegetative axes are sometimes somewhat confluent with
the uppermost lateral axis that is flowering (concaulescence, sensu
Troll 1964). The actual developmental process that causes this dis-
placement in E. urens is unclear, although it also appears to be
concaulescence. The lateral floral shoots in EF. urens (Fig. 6) largely
are the same as in most species of sect. Eucnide, except for the
distinctive terminal flower displacement. Eucnide urens also differs
from the species in sects. Eucnide and Sympetaleia because the leaf
directly beneath a terminal flower is clasping. Renewal growth data
for E. urens is unavailable because I could not obtain adequate
growth of this species in glasshouse populations.

DISCUSSION

Eucnide has monotelic axes (sensu Troll 1964) because each shoot
terminates in a flower. Troll (1964) suggested that the Loasaceae is
among a group of families characterized by monotelic synflores-
cences (i.e., monotelic shoots associated with the initial and each
renewal axis).

Inflorescences in Eucnide were first described as cymes (Urban
1886). Urban (1892) later described inflorescence patterns in various
loasaceous species, including EF. bartonioides. For E. bartonioides,
he described two or three floral branches beneath the terminal flower.
Each floral branch was observed to be adnate with its subtending
leaf and to produce two prophylls before terminating in a flower.
Each prophyll subtended a similar branch and was likewise adnate
with it. His set of observations concur with the patterns I have
described for E. bartonioides, and generally characterize the patterns
found among most of the species (EF. cordata, E. tenella, and E.
urens are divergent the most notably).

No major distinctions in inflorescence architecture seem to dif-
ferentiate sect. Sympetaleia from sect. Eucnide. Within both sections
there are variations (E. cordata in sect. Eucnide and E. tenella in
sect. Sympetaleia) from the commonly expressed patterns. Gilg (1925)
described Eucnide as having flowers arranged in cymes (presumably
implying dichasia in this instance) and monochasia. He described
the genus Sympetaleia (synonymized with Eucnide by Thompson
and Ernst, although they segregated these species into sect. Sym-
petaleia) as having flowers arranged in few-flowered cymes. Gilg’s
use of the term cyme appears to imply a dichasial branching pattern

26 MADRONO [Vol. 34

in the inflorescence, and this concurs with my observations of di-
chasial inflorescences in E. aurea and E. rupestris. Although the
initial and renewal axes tend to form two lateral floral axes (dichasia)
in E. aurea and E. rupestris, | have observed that the successive
iterations of floral branches from each of these lateral floral axes
may be either dichasial or monochasial.

Eucnide tenella is the only species of Eucnide that probably should
not be considered to have either a monochasial or dichasial cyme.
This species was described (Johnston 1924) only a year before Gilg
(1925) published descriptions of Eucnide and Sympetaleia, and E.
tenella was not included among them. The growth pattern of E.
tenella appears to be a simplification of that found in the other
species of Eucnide because it has neither distinct lateral floral branch-
es nor distinct renewal branches that produce more than two leaves
before conversion to flowering. The branches in E. tenella are similar
to the lateral floral branches of the other species because they produce
only one or two leaves and then a terminal flower. They differ from
these branches because they do not die back after a flush of flowering;
instead, they appear to continue producing one or two leaves (each
with iterative axillary buds) and a terminal flower. Thus, the inflo-
rescence in E. tenella is limited to the terminal flower produced by
each axis. This alteration implies that whole plant architecture in
E. tenella would differ significantly from the other species of the
genus.

Among the species of sect. Eucnide, E. cordata has the most di-
vergent inflorescence architecture because its penultimate internodes
are distinctly elongated, as are those of the lateral floral branches.
Waterfall (1959) noted this distinct architectural pattern in E. cor-
data and characterized the inflorescence as “‘lifted above the leaves
on a short peduncle.”’ He considered this to be the greatest tendency
toward a terminal inflorescence in the genus.

Eucnide aurea and E. cordata may invest more heavily in flowers
than other species. Both have a number of flowers that approach
maturity simultaneously on inflorescence systems with extensive
internodal elongation. In other species, floral buds remain small and
inflorescence internodes are unextended until a particular flower
begins to mature. Eucnide cordata and E. aurea, along with E. ru-
pestris (the development of which I have not examined), are the
only species that Thompson and Ernst (1967) reported to have many,
crowded flowers in their inflorescences. These three species and E.
tenella are primarily centered in Baja California, whereas the other
Eucnide species are distributed throughout mainland Mexico and
southwestern United States. Whether these similarities among species
distributed in Baja California represent common ecological adap-
tations, phylogenetic constraints, or merely coincidental conver-
gences should be investigated further. I have shown previously

1987] HUFFORD: EUCNIDE INFLORESCENCES Zi

(Hufford 1986) that individual flowers of E. aurea tend to be longer-
lived following anthesis than flowers of other species of Eucnide (the
same species as grown in glasshouse populations) that were inves-
tigated for this study. The persistence of individual flowers also may
make this species appear to have many flowers that mature simul-
taneously.

The displaced terminal flowers of FE. wrens appear to be an extreme
modification of the slight concaulescence that was observed com-
monly among the other species. Eucnide urens also differs from the
other species because the leaf subtending a terminal flower is clasp-
ing. Waterfall (1959) noted that the ““uppermost leaves [were] some-
times sessile and slightly amplexicaul’’. In the other species, leaves
in the inflorescence region were often quite reduced, but they re-
mained petiolate and were never clasping. These divergent features
in the inflorescence region of E. urens support Thompson and Ernst’s
(1967) segregation of this species into its own section. Cladistic
analysis of Eucnide (Hufford 1986) has shown that the EF. urens
complex is probably a sister group to the rest of the genus. The
inflorescence features are among a suite of unique characteristics
possessed by E. urens within the genus.

When Waterfall (1959) delineated two series in Eucnide based on
inflorescence positions (the first series had solitary flowers in leaf
axils and the second series had terminal inflorescences), he noted,
“collections from young plants beginning to flower might be con-
fused with the first group”’ (1.e., those thought to have solitary flowers
in leaf axils). This observation is likely to be true because Waterfall
and Thompson and Ernst (1967) each characterized at least some
of the species as possessing axillary flowers. Axillary flowers have
not been present in any of the material I examined. It is likely that
the analyses presented in these systematic revisions were confused
by the condensed internodes in the flowering region, the extended
developmental period of the lateral floral branches, and the recau-
lescence in the floral branches. An accurate analysis of the inflores-
cence pattern in Eucnide would have been difficult without obser-
vation of the growth patterns of living plants. It is exceptionally
difficult to determine inflorescence and branching patterns from her-
barium specimens. Further comparative studies of 1) the develop-
mental origin of the recaulescence common to most or all of the
species, 2) the elongated internodes associated with the inflorescence
region of E. cordata, and 3) the displacement of terminal flowers of
E. urens are warranted.

ACKNOWLEDGMENTS

I thank Henry J. Thompson and Bruce Parfitt for providing the seeds of Eucnide
species that I have used in my developmental studies. I thank the following herbaria
for the loan and use of specimens: GH, JEPS, LA, LL, NY, RSA, TEX, UC. I
graciously acknowledge Rudolf Schmid, Donald R. Kaplan, Kevin Padian, Pamela

28 MADRONO [Vol. 34

Diggle, Maynard F. Moseley, and Christopher Davidson for helpful criticisms of this
manuscript during its preparation. This investigation was funded partially by a Grant-
in-Aid of Research provided by Sigma Xi, The Scientific Research Society.

LITERATURE CITED

Dickinson, T. A. 1978. Epiphylly in angiosperms. Bot. Rev. 44:181-232.

GILG, E. 1925. Loasaceae. Jn A. Engler and K. Prantl, eds., Pflanzenfamilien III,
21:522-543.

HUuUFFORD, L. D. 1986. Floral ontogeny and the divergence of reproductive mor-
phologies in Eucnide (Loasaceae). Ph.D. dissertation, Univ. of California, Berke-
ley.

JOHNSTON, I. M. 1924. Expedition of the California Academy of Sciences to the
Gulf of California in 1921. The botany (vascular plants). Proc. Calif. Acad. Sci.
IV, 12:951-1218.

KuutT, J. 1981. Inflorescence morphology of Loranthaceae—an evolutionary syn-
thesis. Blumea 27:1-73.

STEENIS, C. G. G. J. VAN. 1963. Definition of the concept “inflorescence” with
special reference to ligneous plants. Flora Malesiana Bull. 4:1005—-1007.

THOMPSON, H. J.and W. R. ERNST. 1967. Floral biology and systematics of Eucnide
(Loasaceae). Jour. Arnold Arb. 48:56-88.

and A. M. Powe. 1981. Loasaceae of the Chihuahuan desert region.
Phytologia 49:16-32.

TROLL, W. 1964. Die Infloreszenzen. Vol. 1. Gustav Fischer Verlag, Stuttgart.

URBAN, I. 1886. Die Bestaubungseinrichtungen bei den Loasaceen. Jahrb. Konigl.
Bot. Gart. 4:364—-388 + 1 pl.

. 1892. Die Bliithenstande der Loasaceen. Ber. Deutsch. Bot. Ges. 10:220-
225 + 1 pl.

WATERFALL, U. T. 1959. A revision of Eucnide. Rhodora 61:231-—243.

WEBERLING, F. 1983. Fundamental features of modern inflorescence morphology.
Bothalia 14:917—922.

(Received 23 Jan 1986; revision accepted 11 Jul 1986.)

ANNOUNCEMENT

THIRD ANNUAL SOUTHWESTERN BOTANICAL
SYSTEMATICS SYMPOSIUM

‘**Advances in Plant Systematics and Ecology”

For information write to: Rancho Santa Ana Botanic Garden, Botanical
Systematics Symposium, 1500 N. College Ave., Claremont, CA 91711;
phone: (714) 625-8767. Date: 23 May 1987.

A SYSTEMATIC STUDY OF SILENE SUKSDORFII,
S. GRAYI, AND S. SARGENTII
(CARYOPHYLLACEAE)

MARY ANN T. SHOWERS
California Department of Parks and Recreation,
P.O. Box 942896, Sacramento, CA 94296-0001

ABSTRACT

The taxonomic and geographic limits of Silene suksdorfi, S. grayi, and S. sargentii
are not well defined in California. Specimens of Si/ene from subalpine and alpine
regions have been regarded traditionally as either S. grayi or S. sargentii. Several
collections from northern California possess features characteristic of S. suksdorfii.
Evidence from morphological, ecogeographical, and phytochemical examinations is
presented that clarifies the taxonomic relationships, delimits the differences among
these species, and corroborates the existence of 8. suksdorfii in California.

Silene has been the subject of several regional and worldwide
taxonomic revisions (Williams 1896, Robinson 1897, Hitchcock and
Maguire 1947, Chowdhuri 1957). Although many species are cir-
cumscribed clearly, others are polymorphic or possess characters
that do not permit clear distinctions to be made, and information
about the distribution and habitat(s) of these species often is lacking
or defined poorly. Historically, subalpine and alpine specimens of
Silene from the southern Cascade Range of California have been
considered to be either S. grayi Wats. or S. sargentii Wats. The
occurrence of S. suksdorfii Robins. on Mount Shasta has been over-
looked generally, and examination of herbarium records and per-
tinent literature indicates that the distributional ranges of these three
species are not well understood.

Robinson (1891), Hitchcock and Maguire (1947), and Chowdhuri
(1957) commented on the similar morphology of S. suksdorfi, S.
grayi, and S. sargentii. Traits that typically distinguish S. suksdorfii
from S. grayi and S. sargentii are its shorter stature, shorter basal
leaves, bilobed petal blades, and trichomes of the calyx that possess
purple septal walls. Silene suksdorfi is differentiated further from
S. grayi by the presence of anastomosing calyx veins, and from S.
sargentiil by epapillose seeds.

Hybridization studies by Kruckeberg (1955, 1961) showed that
S. suskdorfii, S. grayi, and S. sargentii are distinct species. Krucke-
berg (1955) ascribed the high degree of sterility in hybrids to meiotic
abnormalities during microspore formation and observed little chro-
mosome pairing in hybrids. Kruckeberg (1961) concluded that veg-
etative and reproductive features showed that extensive genetic dif-

MADRONO, Vol. 34, No. 1, pp. 29-40, 1987

30 MADRONO [Vol. 34

TABLE 1. POPULATION DESIGNATIONS USED IN FIGs. 1—4. * = putative populations
of Silene grayi and/or S. sargentii determined to be S. suksdorfii.

SILENE SUKSDORFII

BT Broken Top, Three Sisters Wilderness Area, OR
SS South Sister, Three Sisters Wilderness Area, OR
BU* Bumpass Mt., Lassen Volcanic National Park, CA
CRE* Crescent Crater, Lassen Volcanic National Park, CA
LA* Lassen Peak, Lassen Volcanic National Park, CA
SP* South Plug, Lassen Volcanic National Park, CA
RB* Red Butte, Mount Shasta, CA

SILENE GRAYI
MM Marble Mountain, Marble Mountains Wilderness Area, CA
LMH Little Mount Hoffman, Medicine Lake Highlands, CA
PSM Pumice Stone Mountain, Medicine Lake Highlands, CA
PAN Panther Creek Meadows, Mount Shasta, CA
SHA Glacial basin north of Red Butte, Mount Shasta, CA
TAL Talus at base of Red Butte, Mount Shasta, CA

ferentiation in these and most other western North American Silene
species was complete. In spite of their genetic isolation, these three
species have similar morphology and ecological preferences, which
have contributed to the confusion in their delimitation in California.
The purpose of the present study is to examine these species by the
use of morphological, ecogeographical, and phytochemical analyses
to circumscribe them clearly.

MATERIALS AND METHODS

Specimens of S. suksdorfii and S. grayi were collected in Oregon
and California (Table 1). Extensive collections of S. sargentii were
not made during this study because the taxon is represented amply
in California herbaria.

Field collections of S. suksdorfii and S. grayi were used in chemical
analyses. Flavonoid compounds were extracted from dried flowers,
purified, and identified using two-dimensional paper chromatogra-
phy following standard techniques (Harborne 1967, Mabry et al.
1970). The first phase was developed in 4:1:5 butanol: glacial acetic
acid: water (BAW); the second, in 15% glacial acetic acid. Rutin
provided a reference marker. For spectrophotometric analyses, ex-
tracts from dried flowers were streaked onto Whatman 3 mm paper
and developed in four solvent systems: BAW, 15% glacial acetic
acid, BEW (4:1:2.2 butanol: ethanol: water), and water. Each band
in each system was examined using UV light, with and without
exposure to ammonia fumes. Identification of compounds was made
using UV spectroscopy. Sugar moieties were not identified during

1987]

SHOWERS: SYSTEMATICS OF SILENE

31

TABLE 2. MORPHOLOGICAL CHARACTERS AND DIAGNOSTIC FEATURES OF Silene
suksdorfi, S. grayi, AND S. sargentii. Adapted in part from Hitchcock and Maguire

5

(1947). | = definitive for species; 7 = variable, but definitive in combination with
other characters. All measurements are in millimeters except where noted.

Stature’, cm
Stem glandu-
larity

Leaves?

Leaf glands
Leaf length
Leaf width
Calyx length
Calyx
trichomes!

Calyx nerves!

Petal claw
Petal blade
Petal lobing?

Carpophore
Style number
Seed length!
Testa character!

S. SUKSDORFII

3-10(—15)

glandular above;
increases from
base

matted; linear to
linear-oblanceo-
late

present
(5—)15(-50)
1.5—2(-4)
10-14(-18)
purple-septate

anastomosing;
purple

(7-)8—1 1(-13)

(3-)3.5(—5)

bilobed with occa-
sional small lat-
eral teeth

(2-)2.5-3.5

3(4)

1.1-2.0 (k = 1.6)

tessellate

S. SARGENTII

10-15(—20)
glandular above

tufted, marcescent;
linear-oblanceo-
late

present
15-25(-—40)
(1-)1.5(-3)
(8—)10-14(-17)
hyaline-septate,
rarely purple-
septate
anastomosing;
green-purple
(8—)10—14(-17)
2.5-3.5(-5)
bilobed with small
lateral teeth

1.5-3

3(4.5)

1.2—2.0 (kX = 1.5)

tessellate with
marginal pa-
pillae

S. GRAYI

10-—20(-30)

glandular above;
can be eglandular
below

tufted, thickened, +
fleshy; linear-ob-
lanceolate to ob-
lanceolate-spatu-
late

absent

(15-)20-40(-60)

(1-)2-5S(-7)

(7-)10-12(-13)

hyaline-septate

nonanastomosing;
green-purple

8-11

(3-)3.5(-5)

more or less four-
lobed, rarely bi-
lobed

(1.5-)2-3

3

1.8-—3.0 (KX = 2.3)

tessellate

this study. Flavonoids isolated from putative populations of S. suks-
dorfli were compared to those extracted from different populations
of S. grayi and S. suksdorfii. Flavonoids of S. sargentii were not
examined because this species possessed distinctive morphological
traits.

Morphological measurements were made using field and herbar-
ium specimens to corroborate those given by Hitchcock and Maguire
(1947). Statistical tests included F-test, Student’s t-test, and arcsin
transformation methods (Sokal and Rohlf 1973). Seeds were mea-
sured using a micrometer calibrated for use with a dissecting mi-
croscope.

Specimens were examined from CAS, DS, F, GH, JEPS, MO, NY,
ORE, OSC, PH, UC, US, WS, and WTU. Voucher specimens are
deposited in CAS, SFSU, and WTU. Nomenclature conforms to
Munz (1959) and Hitchcock et al. (1969).

32 MADRONO [Vol. 34

100

80

60

40

PERCENT

20

BT SS BU CRE LA_ RB MM PSM PAN SHA TAL

SUK SDORFIl GRAYI
POPULATIONS

Fic. |. Percent anastomosing calyx veins in populations of Silene suksdorfii and
S. grayi (n = 40).

RESULTS

Morphology. Morphological characters were studied with specific
reference to those given by Hitchcock and Maguire (1947) and are
shown in Table 2. Traits that were valuable in the differentiation of
S. suksdorfli from S. grayi and S. sargentii were the presence of
purple-septate calyx trichomes, the absence of corolla lobes lateral
to primary lobes, seed size, and lack of marginal papillae on seeds.
Silene grayi and S. sargentii possess hyaline-septate calyx trichomes
and corolla lobes lateral to primary lobes. Seeds of S. grayi are larger
than those of S. sargentii or S. suksdorfii, and seeds of S. sargentii
possess marginal papillae. Calyx venation is an additional feature
by which S. suksdorfili and S. sargentii can be distinguished from S.
grayi.

Robinson (1891) distinguished S. grayi and S. suksdorfii using
calyx venation. He reported veins of S. grayi as simple and those
of S. suksdorfii as anastomosing. An examination of specimens of
S. sargentii also indicates that calyx veins are primarily anasto-
mosing in this taxon. Although this trait is variable in these three
species, presence or absence of anastomosing calyx veins can be a

1987] SHOWERS: SYSTEMATICS OF SILENE 33

30

25

20

CM
°

oO

ir

BT SS BU CRE LA RB MM PAN SHA
SUK SDORFII GRAYI

POPULATIONS

Fic. 2. Comparison of stem length in populations of Silene suksdorfii and S. grayi
[n = 15; 1.96 + 1.3 (K + s.e.)].

significant feature. I found that the presence of anastomosing calyx
veins in S. suksdorfii was significant in the differentiation of S.
suksdorfii from S. grayi (p < 0.05). The differences between these
species was greater than the natural variation within each species
(Fig. 1). Anastomosing calyx veins in S. sargentii were not analyzed
statistically due to inadequate sample sizes. Differences in calyx
length/width between S. suskdorfii and S. grayi were not significant
Statistically (p > 0.05). In general, calyx size is not a reliable taxo-
nomic trait in the differentiation of species of Si/ene because the
developing capsule deforms the calyx, which renders field measure-
ments inaccurate (Bocquet and Baehni 1961). Robinson (1891),
Hitchcock and Maguire (1947) and Chowdhuri (1957) attributed
shorter stature to S. suksdorfii than to S. grayi or S. sargentii. Leaves
of S. suksdorfii were described similarly as being smaller than those
of the other two species. In the examined populations, S. grayi is
taller than S. suksdorfii (p < 0.05; Fig. 2). Stature in S. sargentii is

34 MADRONO [Vol. 34

3.0

2.0

BT SS BU LA SP RB MM PSM PAN SHA TAL
SUK SDORFIl GRAYI

POPULATIONS

Fic. 3. Comparison of seed length in populations of Silene suksdorfii and S. grayi
[n = 20; 1.96 + 0.09 (kK + s.e.)].

similar to that of S. grayi. Differences in the size of basal leaves of
S. suksdorfili and S. grayi were not significant (p > 0.05). The leaves
of S. suksdorfii and S. sargentii appear narrower than those of S.
grayi. Seed length in S. suksdorfii and S. sargentii was similar in
range and average size. Silene suksdorfii had significantly different
seed lengths in comparison to S. grayi (p < 0.05; Fig. 3).

Flavonoid analysis. The two-dimensional spot configuration was
similar for S. suksdorfii and S. grayi, but iso-orientin, a c-glycosyl-
flavonoid, was present on chromatograms of S. grayi.

Distribution. Silene suksdorfii occurs at elevations of 1800-3000
m, and is restricted to alpine environments on volcanic peaks in the
Cascade Range. Prior to the present study, S. suksdorfli was reported
solely from the major Cascade Range peaks of Oregon and Wash-
ington, and although Merriam (1899) reported correctly that S. suks-
dorfli occurred on Mount Shasta, its existence in California has been

1987] SHOWERS: SYSTEMATICS OF SILENE 35

Fic. 4. Distribution of Silene suksdorfii (%), S. grayi (*), and S. sargentii (@).
Locations are based on herbarium specimens and from literature, and represent the
corrected distribution of each species.

largely overlooked. In California, S. suksdorfii is represented by
populations on Mount Shasta (Red Butte and Lake Helen) and on
several peaks within Lassen Volcanic National Park (Table 1). Silene
grayi, found at 1200-2900 m elevations, is an element of several
disparate ecosystems: montane chaparral, montane forest, subalpine
forest, and alpine. Substrates on which it occurs include gabbro,
granite, marble, serpentine, andesite, and rhyolite pumice. It occurs

36 MADRONO [Vol. 34

in the Klamath Mountains Province of Oregon and California; in
California, it also occurs in the Medicine Lake Highlands and on
Mount Shasta, the type locality. Silene grayi is sympatric with S.
suksdorfii on Mount Shasta. Silene sargentii, also an alpine species,
occurs at elevations of 2400-3800 m, and is found primarily on
granitic, metamorphic, and volcanic substrates. In California, it oc-
curs on Sierra Nevada peaks from Inyo to Plumas cos.; in Nevada,
it occurs on Mount Rose, and in the Monitor, Toiyabe, and Toquima
ranges.

The distributions of S. suksdorfi, S. grayi, and S. sargentii re-
ported previously in the literature are inaccurate due to misidenti-
fication of herbarium specimens. Collections reported as S. sargentii
from the Cascade Range are either S. suksdorfii or S. parryi (Wats.)
Hitch. & Maguire. Specimens identified as S. sargentii from the
Klamath Mountains Province are collections of S. grayi. Similarly,
individuals identified as S. grayi from the Cascade Range of Oregon
and Washington are either S. suksdorfii or S. parryi. The report of
S. grayi from the Webber Lake Mountains, Sierra Co., California,
by Hitchcock and Maguire (1947) is based on a collection of S.
bernardina Wats. ssp. maguirei Bocquet (=S. montana Wats.). Col-
lectors (Gillett et al. 1961) also have reported S. grayi from Lassen
Volcanic National Park, California, but examination of specimens
from this location revealed they are collections of S. suksdorfii.
Several collections of S. douglasii Hook. from the Cascade Range
and from Mount Olympus have been misidentified as S. suksdorfii.
Some collections of S. sargentii also have misidentified as S. suks-
dorfii. The corrected distributional ranges for S. suksdorfi, S. grayi,
and S. sargentii are illustrated in Fig. 4.

DISCUSSION

Morphology. There is considerable overlap of many morpholog-
ical characters among S. suksdorfi, S. grayi, and S. sargentii, but
seed size and morphology, calyx trichomes and venation, and petal
lobing constitute a suite of taxonomic features useful in their dif-
ferentiation.

The data presented here, in addition to the phylogenetic study of
Williams (1896), offer several conclusions about the value of these
morphological characters: 1) The use of external seed microsculp-
turing is valuable in delimiting S. suksdorfii, S. grayi, and S. sar-
gentii (see Crow 1979, Prentice 1979, Wofford 1981). The prominent
papillae on seeds of S. sargentii are diagnostic. 2) Seed length in S.
grayi is 1.8-3.0 mm, not 1.5 mm as reported by Hitchcock and
Maguire (1947). 3) The presence of purple-septate trichomes is valu-
able in delimiting S. suksdorfii. Purple-septate trichomes are not
known in S. grayi and are rare in S. sargentii. The occasional oc-

1987] SHOWERS: SYSTEMATICS OF SILENE 3M

currence of these trichomes in S. sargentii does not diminish the
importance of this trait in circumscribing S. suskdorfii. 4) Calyx
veins are primarily anastomosing in S. suksdorfii and S. sargentii
and are nonanastomosing in S. grayi. 5) Petal blades in flowers of
S. suksdorfii typically are bilobed and lack lateral teeth; those in S.
sargentii usually possess small lateral teeth, and lateral lobes com-
monly are prominent in S. grayl.

Chemotaxonomy and pollination. The presence of iso-orientin in
S. grayi floral extracts and its absence in those of S. suksdorfii pro-
vides additional evidence to support the present taxonomic and
distributional circumscriptions of these species. Because flavonoids
may function in UV absorption as nectar guides, differences in floral
chemistry may result from adaptations to different pollinators
(Thompson etal. 1972, Harborne 1975). In higher plants, differences
in floral morphology that attract different types of pollinators to
different species or reduce the possibility of cross pollination between
two species are effective prezygotic isolating mechanisms (Stebbins
1977). If floral structure is similar, then diurnal or vespertine flow-
ering also may serve as an isolating mechanism. Silene suksdorfii is
a diurnal species, whereas S. grayi is vespertine. Sphyngid moths
(after sunset) and syrphid flies (at dawn) were observed visiting S.
grayi. No pollinators have been observed on S. suksdorfii.

Ecology. Detailed ecological information is not widely available
for S. suksdorfi, S. grayi, or S. sargentii. Excerpts from phytoso-
ciological studies (Whittaker 1960, Pemble 1970, Hamann 1972,
Taylor 1976, Burke 1982), herbarium collections, regional floras
(Ireland 1968, Hunter and Johnson 1983), and field observations,
however, provide insight into the types of habitats in which they
occur. Silene suksdorfii and S. sargentii are found typically on well-
drained substrates in similar habitats in the alpine: in soil pockets
on talus slopes, in soil around boulders, and on open, windswept
ridges and plateaus. Density and duration of snowpack is variable
in areas in which S. suksdorfii is found. Populations on vertical cliff
faces, such as those on Red Butte and Broken Top, are subject to
winter desiccation and exposure to extreme cold. The ability of S.
suksdorfii to grow on cliff faces may depend on the snowmelt water
from snowbeds in crevices of rocks and on the ability of poikilo-
hydric mosses to rapidly absorb snowmelt and rainfall, as well as
to trap soil (Billings and Mooney 1968, Grime 1979, Walter 1979;
D. W. Showers, pers. comm.). Areas in which S. sargentii occurs
often are not free of snow until midsummer and dry out rapidly
following snowmelt. Populations of S. grayi occur largely on well-
drained substrates, but several alpine populations occur in moist,
poorly-drained soils located commonly in glacial basins (e.g., Mount
Shasta), or on slopes below snowfields. For example, Ferlatte (1974)

38 MADRONO [Vol. 34

reported a population of S. grayi growing on a granitic slope below
a permanent snowfield on Thompson Peak in the Trinity Alps. The
presence of other moisture-tolerant taxa at this site is indicative of
the poorly-drained nature of the soil.

Certain populations of S. suksdorfi, S. grayi, and S. sargentii
occur in alpine habitats below climatic timberline. These popula-
tions occur in areas where localized climatic and edaphic conditions
result in the formation of alpine-like microhabitats at elevations
lower than typical of alpine habitats (Daubenmire 1954, Tranquillini
1979, Walter 1979). The existence of azonal alpine is significant in
understanding the distribution of S. suksdorfii in California because
climatic timberline in northern California occurs at 2800 m. The
presence of suitable azonal alpine environments at Lassen Volcanic
National Park and on Mount Shasta explains the occurrence of SS.
suksdorfii and other species typical of the alpine below that elevation.

KEY TO SPECIES

Blades of petals 2-lobed or with 4 unequal lobes.

Calyx 8-10 mm long; calyx trichomes hyaline-septate; blades of
petals 3—5 mm long; basal lvs. 2-5 mm broad; seeds tessellate,
ca. 2-3 mm long. n. CA ands. OR ............. S. grayi

Calyx 10-14 mm long; blades of petals 2.5—3.5 mm long; basal
lvs. 1-2 mm broad; seeds ca. 1.5 mm long.

Calyx trichomes hyaline-septate (rarely purple-septate); seeds
with tessellate faces and marginal papillae (visible with
hand lens). Plumas to Inyo cos., CA; NV . S. sargentii

Calyx trichomes purple-septate; seeds tessellate, without mar-
ginal papillae. Volcanic peaks, Shasta and Siskiyou cos.,
CAPM. fae de eee Oe ees eee S. suksdorfii

ACKNOWLEDGMENTS

This paper is the result of a study submitted to the Department of Biological
Sciences, San Francisco State University, in partial fulfillment of the requirements
for the Master of Arts degree. I thank Drs. R. W. Patterson, V. T. Parker, and J. R.
Sweeney for their interest, support, and careful analyses of this study. I also thank
the National Park Service, Lassen Volcanic National Park, for assistance during the
course of this study.

LITERATURE CITED

BILLINGs, W. D. and H. A. Mooney. 1968. The ecology of arctic and alpine plants.
Biol. Rev. 43:481-529.

BocquET, G. and C. BAEHNI. 1961. Les Caryophyllacées—Silenoidees de la flore
Suisse. Candollea 17:191-202.

BurKE, M. T. 1982. The vegetation of the Rae Lakes basin, southern Sierra Nevada.
Madrono 29:164-176.

CHOWDHURI, P. K. 1957. Studies in the genus Si/ene. Notes Royal Bot. Gard.,
Edinburgh 22:221-278.

1987] SHOWERS: SYSTEMATICS OF SILENE 39

CookE, W. B. 1940. Flora of Mt. Shasta. Amer. Midl. Nat. 23:497-572.

Crow, G. E. 1979. The systematic significance of seed morphology in Sagina
(Caryophyllaceae) under scanning electron microscopy. Brittonia 31:52-63.
DAUBENMIRE, R. 1954. Alpine timberlines in the Americas and their interpretation.

Butler Univ. Stud. 11:119-136.

FERLATTE, W. J. 1974. A flora of the Trinity Alps of northern California. Univ.
California Press, Berkeley.

GILLETT, G. W., J. T. HOWELL, and H. LESCHKE. 1961. A flora of Lassen Volcanic
National Park, California. Wasmann J. Biol. 19:1-185.

GRIME, J. P. 1979. Plant strategies and vegetation processes. John Wiley & Sons,
Chichester.

HAMANN, M. J. 1972. Vegetation of alpine and subalpine meadows of Mt. Rainier
National Park, Washington. M.A. thesis, Washington State Univ., Pullman.
HARBORNE, J. B. 1967. Comparative biochemistry of the flavonoids. Academic

Press, New York.

1973. Phytochemical methods. Chapman & Hall, London.

. 1975. The biochemical statistics of flavonoids. Jn J. B. Harborne et al.,
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HITCHCOCK, C. L. and B. MAGuIRE. 1947. A revision of the North American species
of Silene. Univ. Wash. Publ. Biol. 13:1-73.

, A. CRONQUIST, M. OwnsBey, and J. W. THompson. 1969. Silene L. In
Vascular plants of the Pacific Northwest, vol. 2, p. 281-296. Univ. Washington
Press, Seattle.

Hunter, K. B. and R. E. JOHNSON. 1983. Alpine flora of the Sweetwater Mountains,
Mono County, California. Madrono 30:89-105.

IRELAND, O. L. 1968. Plants of the Three Sisters region, Oregon Cascade Range.
Bull. Mus. Nat. Hist. 12:1-117. Univ. Oregon Press, Eugene.

KRUCKEBERG, A. R. 1955. Interspecific hybridizations of Si/Jene. Amer. J. Bot. 42:
373-378.

1960. Chromosome numbers in Si/ene (Caryophyllaceae): II. Madrono 15:

205-215.

. 1961. Artificial crosses of western North American silenes. Brittonia 13:
305-333.

Mapsry, T. J., K. R. MARKHAM, and M. B. THOMAS. 1970. The systematic identi-
fication of flavonoids. Springer-Verlag, New York.

MERRIAM, C. H. 1899. Results of a biological survey of Mt. Shasta, northern Cal-
ifornia. North Amer. Fauna 16:1-179.

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

PEMBLE, R. H. 1970. Alpine vegetation in the Sierra Nevada of California as lith-
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PRENTICE, H. C. 1979. Numerical analysis of infraspecific variation in European
Silene alba and S. dioica (Caryophyllaceae). J. Linn. Soc., Bot. 78:181-212.
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43-45.

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Robins. (Caryophyllaceae). M.A. thesis, San Francisco State Univ.

SOKAL, R. R. and F. J. ROHLF. 1973. Introduction to biostatistics. W. H. Freeman
& Co., San Francisco.

STEBBINS, G. L. 1977. Processes of organic evolution. Prentice-Hall, Inc., Englewood
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TAYLOR, D. W. 1976. Ecology of timberline vegetation at Carson Pass, Alpine
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THOMPSON, W. R., J. MEINWALD, D. ANESHANSLEY, and T. EISNER. 1972. Flavonols:

40 MADRONO [Vol. 34

pigments responsible for ultraviolet absorption in nectar guides of flowers. Sci-
ence 177:528—530.

TRANQUILLINI, W. 1979. Physiological ecology of the alpine timberline. Springer-
Verlag, Berlin.

WALTER, H. 1979. Vegetation of the Earth and the ecological systems of the geo-
biosphere. Springer-Verlag, Berlin.

WHITTAKER, R. H. 1960. Vegetation of the Siskiyou Mountains, Oregon and Cal-
ifornia. Ecol. Monogr. 30:279-338.

WILLIAMS, F.N. 1896. A revision of the genus Silene. J. Linn. Soc., Bot. 32:1-196.

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the southeastern United States. Syst. Bot. 6:126-135.

(Received 11 Mar 1985; revision accepted 8 Oct 1986.)

ANNOUNCEMENT
MARY DEDECKER SYMPOSIUM

Co-sponsored by the
University of California White Mountain Research Station
The California Native Plant Society
The Bristlecone Chapter of the CNPS
30 April, 1-3 May 1987
Bishop, California

The University of California White Mountain Research Station with
the California Native Plant Society and its Bristlecone Chapter are co-
sponsoring a symposium and field trip honoring Mary DeDecker. The
topic of the symposium is the flora and plant biology of the eastern
Sierra, Owens Valley, White-Inyo Mountains and western Basin and
Range Provinces. Palynology and legislation related to preservation of
plants in eastern California and western Nevada also are included as
symposium topics.

A field trip will be lead by Mary DeDecker, a member of the BLM
staff, and others to the Eureka Dunes on Saturday, 2 May. A cookout-
BBQ will be held in the Eureka Dunes following the field trip.

For additional information contact Dr. Clarence A. Hall, Jr., White
Mountain Research Station, 6713 Geology Building, University of Cal-
ifornia, Los Angeles, CA 90024; phone: (213) 825-2093.

CLARKIA CONCINNA SUBSP. AUTOMIXA (ONAGRACEAB),
A NEW SUBSPECIES FROM THE SOUTH BAY
REGION, CENTRAL CALIFORNIA

ROBERT N. BOWMAN
Botany Department, Colorado State University,
Fort Collins 80523

ABSTRACT

Clarkia concinna subsp. automixa Bowman is described from the South Bay Region
of central California. Unlike typical C. concinna, this subspecies is not protandrous
and is highly modified for selfing.

Clarkia concinna (Fischer & Meyer) Greene was described in 1835
from material collected near Fort Ross, Sonoma Co., California.
Despite generic revision (Greene 1891), the species has remained
intact, based on its distinctive petal configuration and possession of
four anthers, a condition unique within sect. Eucharidium. The
section contains two species, C. concinna and C. breweri (Gray)
Greene (Lewis and Lewis 1955); both are endemic to northern and
central California. Investigations of breeding systems within sect.
Eucharidium have revealed two dissimilar forms within C. concin-
na. The southern form is strongly, if not exclusively, selfing. Further
field investigations and examination of herbarium material support
the taxonomic distinctness of this group, which I propose herein as
a new subspecies.

Clarkia concinna (Fischer & Meyer) Greene subsp. automixa Bowman
subsp. nov.

Subspecies fabricata, antheris distinctis. Flores nonproterandri,
automixi, claudentes noctu; petala 12 mm longa. Chromosomata
numero 2n = 14 (Figs. 1-5).

Annual, erect to diffusely branched, to 4 dm tall; stems glabrate
or with minute, upwardly curled hairs. Leaves elliptic to ovate,
entire, 2—-3(—4.5) cm long, 6-20 mm broad, narrowed to petioles 5—
25 mm long. Rachis of the inflorescence erect. Flowers erect in bud,
becoming deflexed; sepals linear or narrowly oblanceolate, 1-2 cm
long, 1-2 mm wide, commonly remaining united at the tips at an-
thesis, sharply deflexed at the middle, deep red and petaloid at the
base; petals deep bright pink, the claw not streaked with white or
purple, 8—12(—17) mm long, 4-8 mm broad, the blade 3-lobed, the
lobes shallow, (2—)3—4 mm deep, petals closing the flower at night;

Maprono, Vol. 34, No. 1, pp. 41-47, 1987

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1987] BOWMAN: A NEW SUBSPECIES OF CLARKIA 43

stamens 4, roughly equal, surrounding the style, filaments pinkish-
lavender; anthers without hairs or only slightly ciliate, curling after
dehiscence; pollen bluish with copious viscin threads; stigma white,
obscurely 4-lobed, appearing capitate or bifid at maturity, receptive
prior to or at anthesis, positioned with the anthers, clearly not prot-
androus. Chromosome number 2n = 14. Flowering from mid-May
to late June.

Type: USA, California, Santa Clara Co.: Mt. Hamilton Range,
along Kincaid Rd., 11.1 km n. of Smith Creek Ranger Station,
w.-facing slope 3 m e. of road in grassy oak woodland and ca. 300
m s. of locked gate across road, R3E T6S S29 nw.'4 nw.'4 (Mt. Day
quad.), 37°23'13”"N, 121°39'26’W, elev. 735 m, 29 May 1986, Bow-
man 7001 (Holotype: UC; isotypes: CAS, CS, CSUC, GH, LA, MO,
RSA, SJSU).

PARATYPES: USA, California, Alameda Co.: 2.4 mi w. of Sunol,
14 May 1938, Constance 2233 (CAS, GH, RSA, UC). Santa Clara
Co.: Montebello Rd., 3.1 miw. of Stevens Canyon Rd., 16 Jun 1983,
Bowman 3202 (CS, CSUC, UC); 11.3 mis. of Alum Rock Ave. on
Mt. Hamilton Rd., 16 May 1985, Bowman 6035 (CS, CSUC, UC);
jct. Stevens Canyon Rd. and Redwood Gulch Rd., 24 May 1985,
Bowman 6076 (CS, CSUC, UC); Soda Spring Canyon, 28 May 1895,
Dudley 4029 (RSA); Congress Springs, 13 May 1904, Heller 7412
(DS, GH, UC); Booker School near Saratoga, 19 May 1906, Pen-
dleton 346 (POM, UC); Poverty Flat, Henry Coe St. Park, 10 May
1972, Powers 581 (SJSU); Smith Creek, 7 May 1934, Sharsmith
1016 (DS, RSA, UC); headwaters of Stevens Creek, 3 Jun 1961,
Thomas 9517 (DS, RSA).

Distribution. Clarkia concinna subsp. automixa is limited to Santa
Clara and southern Alameda cos., California (Fig. 6). Although it
occurs extensively in the foothills surrounding the Santa Clara Val-
ley, this subspecies is known only from Sunol Canyon in Alameda
Co. It is common in mesic, shaded oak woodlands. In contrast,
subsp. concinna ranges from extreme northwestern California south-
ward to the Oakland Hills (Alameda Co.) and Mt. Diablo (Contra
Costa Co.). The two subspecies are allopatric and are separated by
a minimum distance of ca. 33 km.

—

Fics. 1-5. Floral variation in subspecies of Clarkia concinna. Specimens grown
from field collections as indicated. 1. Top row, subsp. concinna, |. to r., Bowman
6070, 3336, 6074; bottom row, subsp. automixa, Bowman 3685, 3687, 3302; all at
1x. 2. Stigma exertion in subsp. concinna, Bowman 6073; 2x. 3. Stages in the
protandrous development of subsp. concinna flowers. The interval between anthesis
(top) and stigmatic receptivity (bottom) is 1-2 days; ca. 2x. 4, 5. Self-pollination in
subsp. automixa, Bowman 6036; 2 x . Stigmas in subsp. automixa are not protandrous
and are positioned with the anthers at dehiscence.

44 MADRONO [Vol. 34

Santa Clara

Fic. 6. Geographical distribution of Clarkia concinna subsp. concinna and subsp.
automixa in northern and central California.

Morphology and pollination. Clarkia concinna subsp. automixa
is distinguished by a combination of characters that promote selfing.
Its flowers are smaller and markedly less variable (Fig. 1) than those
of subsp. concinna, which are predominately outcrossed. Through-
out its entire range, subsp. concinna is strongly protandrous (Figs.
2, 3); its anthers and stigmas are isolated temporally and spacially.
The flowers of subsp. automixa are not at all protandrous (Figs. 4,
5) because the stigma becomes pollen-receptive before the bud opens
and anther dehiscence occurs as the flower opens. The stigma is
positioned in close proximity to the anthers, thus, facilitating self
pollination. The corolla of subsp. automixa also lacks white streaks,
particularly along the base of the claw, which are characteristic of
subsp. concinna. MacSwain et al. (1973) noted that flowers in C.
concinna remain open at night, an observation correct for all known
populations of subsp. concinna. In contrast, all populations of subsp.
automixa possess corollas that close tightly at night and appress
anthers against the stigma, thus, promoting selfing. Undisturbed
greenhouse plants of subsp. concinna generally fail to set seed, where-
as those of subsp. automixa routinely set full complements of seeds.
This confirms that the combination of morphological characters
unique to subsp. automixa serves to facilitate selfing.

Figure 7 depicts the distribution of stamen/style ratios measured
from all specimens of Clarkia concinna available at CAS, CS, CSUC,
DS, GH, HSC, JEPS, POM, RSA, SACT, SJSU and UC. The dis-

1987] BOWMAN: A NEW SUBSPECIES OF CLARKIA 45

re CLARKIA CONCINNA

subsp. CONCINNA

| | subsp. AUTOMIXA

20

15

NUMBER OF COLLECTIONS

10

45 '.5so ' 55 ' 60 ' 65 ' 70 ' 75 ' 80 '.85 ' 90 ' 95 |! 1.0 ! 1.05
STAMEN/STYLE RATIO

Fic. 7. Distribution of stamen/style ratios in Clarkia concinna. Stamens were
measured from the base of the filament to the tip of the anther. Styles were measured
from the distal tip of the ovary to the stigma surface. Only collections with three or
more mature flowers were measured; values reported are averages of ratios from all
available flowers on a specimen. Although the number of collections is clearly de-
pendent on the total number of specimens examined, the two subspecies are defined
by discontinuous stamen/style ratios.

continuous displacement of stamen/style ratios between the. two
subspecies agrees with evidence discussed previously. Curiously,
those specimens of subsp. concinna (stamen/style ratio of 0.80—0.85)
that most closely approach subsp. automixa are not from sympatric
areas between the subspecies, but rather, from the extreme northern
limits of the range of subsp. concinna. Thus, selfing may occur at
both the extreme northern and southern limits of the species with
allogamy predominating elsewhere. A similar pattern with selfing
predominating at the margins of a species’ range has been reported
in Lycopersicon pimpinellifolium (Solanaceae) by Rick et al. (1977).
Unlike the well developed autogamy in subsp. automixa, the north-
ern populations of subsp. concinna show no signs of morphological

46 MADRONO [Vol. 34

modification promoting selfing, other than stamen/style ratios ap-
proaching unity.

Taxonomic recognition of subsp. automixa is justified by com-
parison with other treatments in the genus. At least five Clarkia
species have been described principally on their status as derived,
primarily selfing, neospecies (Vasek and Harding 1976, Lewis 1973).
Several subspecies in the genus, such as C. tembloriensis Vasek
subsp. calientensis (Vasek) Holsinger (Holsinger 1985), C. gracilis
(Piper) Nelson and Macbride subsp. gracilis, and C. purpurea (Cur-
tis) Nelson and Macbride subsp. guadrivulnera (Douglas) Lewis and
Lewis (in Lewis and Lewis 1955), also are based predominately on
their selfing habit, even though the subspecies are not isolated from
their conspecifics by geographic or other strong reproductive bar-
riers. Subspecific taxa in C. concinna are differentiated by absolute
geographical barriers and an assemblage of morphological charac-
ters. Although possible, gene flow between the two subspecies, as
demonstrated by morphological continuity, is not in evidence. Gene
flow is not likely, furthermore, because pollination of subsp. con-
cinna is dependent on Lepidoptera and long-tongued Diptera
(MacSwain et al. 1973). These insects are not known for long distance
dispersal.

Sectional affinity. The geographical range of subsp. automixa is
perhaps not based simply on ecological preference. Its range nearly
matches the northern range of C. breweri, the only other species in
sect. Eucharidium. Both species occur sympatrically at Congress
Springs, Loma Prieta, Mt. Hamilton, and Cedar Mountain. Even
though C. concinna and C. breweri overlap geographically, strong
autogamy in subsp. automixa within this region prevents gene flow.
Otherwise, all taxa within sect. Eucharidium (all 2n = 14) are easily
hybridized by artificial means and the progeny are morphologically
intermediate. The identical base numbers and the comparative ease
with which artificial hybrids can be produced suggest that chro-
mosomal repatterning may not have been important in evolution
of the section, although it is prevalent elsewhere in the genus (Lewis
1962). Regardless of the mechanisms that enforce reproductive iso-
lation in the section, lack of hybrids among herbarium specimens
or in extensive field reconnaissance indicates that the integrity of
each taxon is conserved. The factors governing evolution and re-
productive isolation in the section remain to be elucidated.

ACKNOWLEDGMENTS

I thank curators of the herbaria herein cited for access to their specimens. Appre-
ciation also is extended to R. Gardner, L. Main, and B. O’Brien for field locality
information.

1987] BOWMAN: A NEW SUBSPECIES OF CLARKIA 47

LITERATURE CITED

GREENE, E. L. 1891. Epilobiaceae. Flora Franciscana. Payot, Upham and Co., San
Francisco.

Ho LsInNGceER, K. E. 1985. A phenetic study of Clarkia unguiculata Lindley (Onagra-
ceae) and its relatives. Syst. Bot. 10:155-165.

Lewis, H. 1962. Catastrophic selection as a factor in speciation. Evolution 16:257-
271.

. 1973. The origin of diploid neospecies in Clarkia. Am. Nat. 107:161-170.

and M. E. Lewis. 1955. The genus Clarkia. Univ. Calif. Publ. Bot. 20:241-
392.

MacSwaln, J. W., P. H. RAVEN, and R. W. THorp. 1973. Comparative behavior
of bees and Onagraceae. IV. Clarkia bees of the western United States. Univ.
Calif. Publ. Ent. 70:1-80.

Rick, C. M., J. F. ForsBes, and M. HoLie. 1977. Genetic variation in Lycopersicon
pimpinellifolium: evidence of evolutionary change in mating systems. PI. Syst.
Evol. 127:139-170.

VASEK, F. S. and J. HARDING. 1976. Outcrossing in natural populations. V. Analysis
of outcrossing, inbreeding, and selection in Clarkia exilis and C. tembloriensis.
Evolution 30:403-411.

(Received 6 Mar 1986; revision accepted 3 Sep 1986.)

ANNOUNCEMENT
NEW PUBLICATION
Flora of the Great Plains

BARKLEY, T. M. (ed.), R. E. BRooks, E. K. SCHOFIELD (assoc. eds.), R.
L. McGREGor (coordinator), and 11 other members of the Great
Plains Flora Association (W. T. Barker, M. Bolick, S. P. Churchill,
R. L. Hartman, R. B. Kaul, O. A. Kolstad, G. E. Larson, D. M.
Sutherland, T. Van Bruggen, R. R. Weedon, D. H. Wilken), Flora of
the Great Plains, Univ. Kansas Press, 329 Carruth St., Lawrence‘
66045, 1986, vil, 1392 pp., illus., ISBN 0-7006-0295-xX, $55.00
(hardbound). [Covers Kansas, Nebraska, North and South Dakota,
e. Montana, e. Wyoming, e. Colorado, ne. New Mexico, the Texas
panhandle, nw. Oklahoma, w. Missouri, w. Iowa, w. Minnesota.]

NEW RECORDS OF MYXOMYCETES FROM
CALIFORNIA. VI.

DONALD T. KOWALSKI
Department of Biological Sciences,
California State University, Chico 95929

ABSTRACT

Twenty-two additional species of Myxomycetes are reported from California. Nine
of these, Comatricha ellae, C. longipila, C. penicillata, Didymium bahiense, D. ver-
rucosporum, Licea lucens, Macbrideola argentea, Paradiacheopsis cribrata and P.
microcarpa, appear to be new records for the United States.

In the last paper of this series (Kowalski 1973), I reported that 231
species of slime molds had been reported from California. Since
then, an additional 32 species have been listed from the state. These
additional records resulted mainly from the work of Whitney (1980,
1982) on corticolous species and Cox (1981) on coprophilous taxa.
In the present paper, 22 new records are discussed, bringing the
number of species of Myxomycetes reported from California to 285.

All collections listed have been deposited in the Herbarium of the
University of California (UC). The nomenclature generally follows
that of Martin and Alexopoulos (1969) and, unless otherwise stated,
the collection numbers are my own.

LICEACEAE

LICEA LUCENS Nannenga-Bremekamp.— Butte Co.: Chico, Upper
Bidwell Park, on Live Oak bark, 24 Jan 1977, Whitney 368; Lower
Bidwell Park, on bark of Quercus lobata Neé, 23 Feb 1977, Whitney
401. Both collections were obtained from bark placed in moist cham-
bers to allow the fructifications to develop. The minute (ca. 50 um
in diameter) stalked, bright orange sporangia, containing strongly
warted spores, are the hallmarks of this distinctive species. In her
original description, Nannenga-Bremekamp (1981) states the spores
are 8-10 um in diameter; in the California collections, however,
they are usually 10-11 wm in diameter.

Because of its small size, L. /ucens appears most similar to L.
perexigua Brooks & Keller. Both taxa have brightly shining, often
stalked sporangia that are less than 100 wm in diameter. The major
differences between the species are as follows: in L. perexigua the
sporangia are yellow to dark bluish gray and the spores smooth,
whereas in L. /ucens the sporangia are orange and the spores are
distinctly warted. Licea lucens was described originally from France,

MADRONO, Vol. 34, No. 1, pp. 48-56, 1987

1987] KOWALSKI: MY XOMYCETE RECORDS 49

and the present report marks only its second known occurrence. As
more workers cultivate bark in damp chambers for slime molds, I
predict this species will be found in many localities.

LICEA OPERCULATA (Wingate) Martin.—Alameda Co.: Oakland,
Redwood Park, on bark, 4 Feb 1970, Duran; Berkeley, Strawberry
Canyon, on bark, 29 Jan 1972, Duran. As with Licea lucens, both
collections were obtained by the moist chamber technique. The
stalked, urniform, operculate sporangia separate this taxon from all
other members of the genus. Martin and Alexopoulos (1969) give
the height of the fructifications as 0.4—1.0 mm and state that the
spores are colorless by transmitted light. In the California material,
the total height of the fruiting bodies reaches 1.2 mm and the spores
are pale yellow by transmitted light. Licea operculata has been re-
ported from numerous locations around the world and appears to
be common on the bark of living trees.

RETICULARIACEAE

LYCOGALA EXIGUUM Morgan.— Butte Co.: Covered Bridge, Hon-
eyrun Road, on decayed wood, 15 Apr 1967, 5974. This collection
of the pseudocapillitium are approximately 5 wm in diameter and
the spores are small, averaging 4.5—5.0 wm in diameter, and smooth
to weakly reticulate. This taxon has been reported from many lo-
calities beyond California, but in all cases it appears to be uncom-
mon.

ENTERIDIUM MINUTUM Sturgis.— Plumas Co.: Humbug Summit,
6700 ft., on decayed wood, 4 Jun 1966, 3320; Siskiyou Co.: Mt.
Shasta, Panther Meadows Campground, 7600 ft., on decayed wood,
6 Jul 1965, 7821, 1878. Confusion as to the exact status of this
species has existed in the past. Both Lister (1925) and Hagelstein
(1944) reluctantly accepted the taxon as valid, whereas Martin and
Alexopoulos (1969) thought it was a small form of Enteridium oli-
vaceum Ehrenberg [Reticularia olivacea (Ehrenb.) Fries]. The prob-
lem originated because all workers stressed the size of the fructifi-
cations. Lister thought E. minutum had aethalia 1-2 mm in diameter,
whereas those of E. olivaceum were usually over 1 cm. Martin and
Alexopoulos, however, thought the size overlapped and, thus, did
not believe that E. minutum was worthy of recognition. I agree with
Martin and Alexopoulos that the size of the fructifications overlap.
If one looks at the spores, however, the two taxa can be distinguished
easily. In E. olivaceum, the spores are olivaceous, occur in large
clusters, usually 6—20 in number, and the ornamentation consists of
large warts covering about one-half of the surface of the spores. In
E. minutum, the spores are yellow, the clusters are small, usually
consisting of 2—4 spores, and the ornamentation is minutely spi-
nulose or warted, covering most of the surface of the spores. Enteri-

50 MADRONO [Vol. 34

dium minutum is an extremely rare species. Previously it had been
known only from the type locality at Eldora Lake, Colorado, and
from Yorkshire, England.

CRIBRARIACEAE

CRIBRARIA FERRUGINEA Meylan.— Mendocino Co.: Simpson Lane,
2 mie. of State Hwy 1, on decayed wood, 15 Apr 1976, 13467. The
large, brick-red sporangia, 1.0—1.5 mm in diameter, with their perid-
ial nets lacking distinct nodes and calyculi, delimits this distinct,
apparently rare species. Martin and Alexopoulos (1969) give its
distribution as Switzerland, Tennessee, Oregon, and New Mexico.

TRICHIACEAE

ARCYRIA MAGNA Rex.— Butte Co.: Chico, Lower Bidwell Park, on
decayed wood, 4 Feb 1967, 5225; Humboldt Co.: Humboldt Red-
woods State Park, on decayed wood, 26 Jan 1966, 24/2. The nu-
merous, densely clustered, grayish sporangia, which often attain a
length of more than a centimeter after becoming fully expanded,
separate this species from other members of the genus. Although
occurring worldwide, A. magna is found infrequently.

TRICHIA MACBRIDEI M. E. Peck.— Butte Co.: Philbrook Reservoir,
5500 ft., on dead bark, 13 Jul 1966, 3809. This taxon can be dif-
ferentiated by the dark, sessile sporangia that contain brownish spores
11-13 wm in diameter. Trichia brunnea Cox 1s the only other species
in the genus that has brown spores, but it has stipitate sporangia
and the spores are 10—11 um in diameter. An unusual feature of the
California collection is the capillitial threads that often terminate in
depressed expansions, which resemble minute suction cups, a char-
acteristic that has not been reported previously. Trichia macbridei
is rare and it apparently has been reported previously only from
Oregon.

TRICHIA SUBFUSCA Rex. — Humboldt Co.: Patrick’s Point State Park,
on decayed wood, 29 Mar 1969, 9910, and on decayed bark, 1 Apr
1969, 9967; Mendocino Co.: MacKerricher Beach State Park, on
decayed wood, 12 Apr 1968, 8258. Trichia subfusca often has been
included in Trichia botrytis (G. F. Gmel.) Pers., which may explain
why this relatively common species has not been reported from
California. The best features to use in distinguishing the two taxa
are the capillitium and spores. In 7. botrytis, the individual elaters
taper gradually to long, slender, pointed tips, and the spores are 9-
11 wm in diameter. In 7. subfusca, the elaters end abruptly, often
in curved, pointed tips, and the spores are (11—)12—14(—15) um in
diameter. The hypothallus in 7. subfusca is huge, often uniting the
individual sporangia into a unit, whereas in 7. botrytis it is usually
much smaller and much less extensive, rarely uniting the sporangia.

1987] KOWALSKI: MYXOMYCETE RECORDS |

STEMONITACEAE

AMAUROCHAETE COMATA G. Lister & Brandza.— Butte Co.: Chico,
Lower Bidwell Park, on dead bark, 23 Nov 1966, 3956, and 12 Feb
1969, 9899: 14 mi n. of Chico, Pine Creek Ranch, 1 Feb 1973,
12587. Amaurochaete comata is the only species in the genus with
a capillitium consisting of flaccid, circinate threads, approximately
1-2 um in diameter; therefore, it is distinct. The collections cited
above differ somewhat from the description given in Martin and
Alexopoulos (1969). They give the diameter of the aethalia as 5—10
mm and describe the spores as prominently warted. In the California
material, the aethalia are up to 5 cm in diameter and the spores are
distinctly spinose. These differences may indicate that the California
collections represent a different taxon. At this time, however, I do
not believe that these differences are large enough to warrant the
description of a new taxon. Amaurochaete comata was described
originally from Romania. Eliasson (1977) reported it from France
and Sweden and Farr (1982) reported it from Alaska. Thus, although
A. comata seems to be rare, it has a wide distributional pattern.

COLLODERMA OCULATUM (Lipert) G. Lister.—Mendocino Co.:
MacKerricher Beach State Park, on rotting wood covered with leafy
liverworts, 31 Mar 1972, 12318, 12322, and 23 Mar 1972, 12316.
In each of these collections, the substrate was originally collected
because it had another, larger myxomycete on it. The small sporangia
of C. oculatum were discovered later in the laboratory while the
substrate was being scanned with a stereoscopic microscope. The
outer gelatinous layer, unique to the genus, is scantily developed in
these collections. Additionally, the spores are warted rather than
echinulate as given in Martin and Alexopoulos (1969). Colloderma
oculatum is a poorly known species; until more material is available
for study, I believe it is best to take a conservative taxonomic ap-
proach and to enlarge the species concept to include the California
collections. Colloderma oculatum is known only from a few collec-
tions in the United States. This is due undoubtedly to its small size
and to it growing among bryophytes on dead wood, hidden from
view. When investigators study lignicolous bryophytes thoroughly
for slime molds, I predict that C. oculatum will be found more
commonly than heretofore thought.

MACBRIDEOLA ARGENTEA Nannenga-Bremekamp & Yamamo-
to.— Butte Co.: 2 mi ne. of Magalia, on bark of Cupressus macnab-
iana A. Murray in moist chambers, 28 Oct 1978, Whitney 997,
1006, 1012 and 7 Mar 1979, Whitney 1062; Deer Creek Canyon,
7.5 mi w. of Ponderosa Way, on bark of Vitis californica Bentham
in moist chamber, 4 Apr 1977, Whitney 418; Los Angeles Co.: Santa
Catalina Island, Avalon, on bark of Cypressus sp. in moist chamber,
2 Sep 1978, Whitney 963. Within the genus, M. argentea can be

52 MADRONO [Vol. 34

distinguished by its persistent peridium, long stalks that are three-
fourths to four-fifths the total height, and spores having clusters of
larger and darker warts. As Nannenga-Bremekamp and Yamamoto
(1983) have indicated, this species probably is most closely related
to Lamproderma biasperosporum Kowalski. Both species have hol-
low stipes, persistent peridia, and spores with warts of two sizes.
They can be differentiated most easily on the bases of the capillitium
and sporangial size. In M. argentea, the sporangia are approximately
0.1 mm in diameter and the capillitium is reduced, consisting of
two or three branches of the columella that also branch 3-5 times
and infrequently anastomose to form a weak but dark brown net.
In L. biasperosporum, the sporangia are 0.25—0.5 mm in diameter
and the capillitium is more highly developed, being formed from
numerous branches of the columella, which radiate and branch many
times in all directions. These branches also anastomose infrequently
to form a weak, but decidedly whitish, net. Previously, M. argentea
had been known only from moist chamber developments made from
several locations in Japan. It may have been collected in the past,
however, and incorrectly identified as L. biasperosporum. Hence, it
could be much more common than previously thought.

MACBRIDEOLA MARTINII (Alexopoulos & Beneke) Alexop. — Lassen
Co.: Eagle Lake Field Station, developed on bark of Juniperus oc-
cidentalis Hooker in a damp chamber, 23 May 1977, Whitney 426.
This collection was assigned to M. martinii because it is the only
species in the genus with a completely evanescent peridium and
spores with clusters of larger and darker warts. Whitney’s collection,
however, may represent an undescribed taxon because it differs in
several respects from the published descriptions of M. martinii. In
typical M. martinii, the stipes are long, usually 5-10 times the di-
ameter of the sporangia, the columella is tapering, the capillitium is
smooth, and the spores are 6.5—8.0 wm in diameter. In Whitney’s
material, however, the stipes are only 2—4 times the diameter of the
sporangia, the columella is broad and scarcely tapering, the capil-
litium bears conspicuous bead-like outgrowths, and the spores are
much larger, 10.5—12.0 wm in diameter. Because typical M. martinii
has been found apparently only in Jamaica, Dominica, Gambia, and
Kentucky, and it is rare, I believe it prudent to consider Whitney’s
material an extreme variant of M. martinii and to wait for additional
material to become available before reaching a final conclusion as
to the disposition of the California form.

COMATRICHA ELLAE Harkonen.— Butte Co.: 5 mie. of Stirling City,
4000 ft., on decayed wood, 19 Aug 1965, 2006; Chico, Lower Bid-
well Park, on decayed wood, 18 Nov 1966, 3907; Lassen Co.: Eagle
Lake Field Station, on bark of Juniperus occidentalis in a moist
chamber, 20 Jun 1977, Whitney 511; Los Angeles Co.: Santa Cat-
alina Island, Avalon, on bark of Cupressus sp. in a moist chamber,

1987] KOWALSKI: MYXOMYCETE RECORDS 53

2 Sep 1978, Whitney 962. The major features of this taxon are the
small size (less than 1.0 mm in total height), globose sporangia,
capillitium that forms a distinct surface net with few free ends, and
the relatively long stipes that are 3-4 times the diameter of the
sporangia. The California material is typical in all respects. When
originally described by Hark6nen (1977, as C. nannengae), C. ellae
was known only from Norway and Finland. It has been reported
since from Spain (Nannenga-Bremekamp and Lado 1985) and prob-
ably will be discovered wherever bark of living trees is cultured.

COMATRICHA LONGIPILA Nannenga-Bremekamp.—Sutter Co.:
Sutter Buttes, on bark of living Quercus sp., 29 Jan 1969, 9841. This
collection was made in the field, not in a moist chamber. Comatricha
longipila appears to be most closely related to C. /axa Rost. Both
taxa are relatively common on the bark of living trees and have
sporangia that are globose to elongate in shape. Additionally, the
main branches of the capillitium arise at right angles to the columella,
and a distinct surface net is lacking. There are two major differences
between these species: in C. /axa, the spores are 7-11 wm in diameter
and the capillitium terminates in numerous short, free ends; in C.
longipila, the spores are 6—7 wm in diameter and the capillitium has
long free ends at the periphery. Although reported from several
localities in Europe, this report appears to be the first record for
North America.

COMATRICHA PENICILLATA Nannenga-Bremekamp & Yamamo-
to.— Nevada Co.: Donner Summit, 7200 ft., on dead wood, 24 Jun
1971, 11537. The distinctive characteristics of this species include
jet-black, globose sporangia that are less than 0.2 mm in diameter
with unbranched, or sparsely branched, capillitial threads ending in
slight expansions. As Nannenga-Bremekamp and Yamamoto (1983)
indicate, it is similar to Paradiacheopsis fimbriata (G. Lister) Hertel,
and the two species can be differentiated as follows: in P. fimbriata,
the columella is stout and not tapering, the capillitium radiates in
all directions, and the spores are 10-14 um in diameter; in C. pen-
icillata, the columella tapers, the capillitium is brush-like, and the
spores are 7.0—9.0 um in diameter. In the original description, the
height of the sporangia is listed as 0.8—1.0 mm and the spores are
given as 7.0—8.5 wm in diameter. The California collection differs
from the type collection, which was made in Japan (apparently the
only other known collection), by having sporangia up to 2.0 mm in
height and spores 8—9 um in diameter.

PARADIACHEOPSIS CRIBRATA Nannenga-Bremekamp.—Sonoma
Co.: 3 mi ne. of Asti, Thompson Property, on bark of Quercus sp.
in moist chamber, 22 Jul 1978, Whitney 930; 25 Nov 1978, Whitney
1022. Both P. cribrata and P. acanthodes (Alexopoulos) Nannenga-
Bremekamp have small sporangia, 0.6 mm or less in total height,
and strongly spinose spores, 12—14 wm in diameter. In P. cribrata,

54 MADRONO [Vol. 34

the capillitial threads are numerous, stout, and anastomosed at the
surface so that a rigid network is formed. In P. acanthodes, the
capillitial threads are fewer in number, finely pointed, and do not
fuse to form a peripheral net. This appears to be the first report of
P. cribrata from the Western Hemisphere.

PARADIACHEOPSIS MICROCARPA (Meylan) Mitchell.— Butte Co.: 2
mi ne. of Magalia, developed on bark of Cupressus macnabiana in
moist chamber, 28 Oct 1978, Whitney 1019; Lassen Co.: Eagle Lake,
2 mis. of Little Troxel Point, developed on bark of Juniperus oc-
cidentalis in moist chamber, 18 Jul 1977, Whitney 699. The major
features of this taxon are the small sporangia, less than 1.0 mm in
total height, the primary branches of the capillitium that arise at
right angles to the columella and end in fine threads bearing short,
spine-like processes, and the spinulose or minutely warted spores,
11-—13(—14) um in diameter. Meylan (1921) originally described this
taxon (as Comatricha laxa var. microcarpa) from three collections
made from the same tree in the Jura Mountains of Switzerland.
Thus, these two collections appear to represent only the fourth and
fifth known specimens and the first from the Western Hemisphere.

PARADIACHEOPSIS RIGIDA (Brandza) Nannenga-Bremekamp.—
Butte Co.: Chico, corner of Arcadian and Sowilleno avenues, on
palm stem in moist chamber, 17 Nov 1966, 3887; Chico, Lower
Bidwell Park, on bark of Juglans sp. in moist chamber, 10 Jan 1977,
Whitney 295; 2 mine. of Magalia, on bark of Cupressus macnabiana
in moist chamber, 28 Oct 1978, Whitney 1013; Marin Co.: San
Rafael, Lucas Valley Road and US Highway 101, on dead wood, 13
Jan 1977, Whitney 248. Paradiacheopsis rigida appears to be related
most closely to P. microcarpa. Sporangia of the two taxa look very
similar under a stereoscopic microscope. They can, however, be
distinguished by numerous characters. In P. rigida, the sporangia
attain 1.25 mm in total height; the base of the stipe is distinctly
yellowish; most of the capillitium originates from the apex of the
columella; the branches are dichotomous, flexuose, and occasionally
anastomosed; and the spores are 9-10 um in diameter and minutely
spinulose. In P. microcarpa, however, the sporangia are smaller,
rarely reaching | mm; the stipe is black along its entire length; the
capillitium arises evenly along the length of the columella; the
branches are extremely rigid, completely free, and not dichotomous;
and the spores are | 1—13(—14) um in diameter and distinctly warted.
Paradiacheopsis rigida has been reported in the United States only
from Minnesota (Hagelstein 1944).

DIDYMIACEAE

DIDERMA EFFUSUM (Schweinitz) Morgan.— Alameda Co.: Berke-
ley, University of California Campus, on dead leaves and herbaceous

1987] KOWALSKI: MYXOMYCETE RECORDS =P)

stems, E. E. Morse, 22 Jan 1930. This distinctive species is char-
acterized by flat sporangiate to plasmodiocarpous fructifications that
contain minutely warted spores with clusters of larger, darker warts.
Diderma effusum is generally common throughout its worldwide
distribution; therefore, it is interesting that I have never found it in
over 20 years of collecting in California. This collection appears to
be the only record for the state.

DIDYMIUM BAHIENSE Gottsberger. — Humboldt Co.: Trinidad, Col-
lege Cove, on fallen leaves, 20 Apr 1973, 12753; Mendocino Co.:
MacKerricher Beach State Park, on dead leaves, 29 Dec 1967, 7629,
7641; Albion, on fallen leaves, 11 Apr 1968, 8205, 8225. This
species was originally described from Brazil (Gottsberger 1968) and
has been reported from the Netherlands (Nannenga-Bremekamp
1972) and England (Mitchell 1977). These collections appear to be
the first reports from North America. The features that delineate
this taxon are the yellowish stipes, the distinctive whitish to yellow-
ish pseudocolumellae, and the minutely warted spores with clusters
of larger, darker warts. It appears to be similar to D. megalosporum
Berkeley & Curtis, but in that species the pseudocolumella is usually
spiny or roughened and the spores lack the distinct clusters of larger
and darker warts.

DIDYMIUM VERRUCOSPORUM Welden.— Butte Co.: Chico, Upper
Bidwell Park, on dead grass leaves, 17 Jan 1969, 9828; Glenn Co.:
13 mis. of Hamilton City, on dead dicot leaves, 24 Feb 1968, 786/,
7864. The nodding, globose sporangia containing white, globose
columellae and warted spores with clusters of larger warts easily
separates this species from others in the genus. Didymium verruco-
sporum was described originally from Panama (Welden 1954) and
has been found in various European and Asian localities. These
collections appear to be new records for the United States. The
California material differs from the type in that, in addition to the
warts, the spores bear a wide-meshed reticulum with four or five
meshes per spore. The reticulum is very similar to the one illustrated
for Didymium nigrisporum by Nannenga-Bremekamp et al. (1984).

LEPIDODERMA AGGREGATUM Kowalski.— Tehama Co.: Wells Cab-
in Campground, 6300 ft., on decaying bark near the melting snow,
18 Jun 1966, 3570. This distinctive alpine slime mold is character-
ized by sessile, clustered, buff sporangia 1.5—3.0 mm in diameter,
which contain spores with widely scattered spines. Until now, it was
known only from the state of Washington, where it is extremely
common in the spring near melting snowbanks.

ACKNOWLEDGMENTS

I thank T. Duncan and I. I. Tavares (UC) and K. D. Whitney (Univ. Texas,
Arlington) for loan of material during the course of this investigation.

56 MADRONO [Vol. 34

LITERATURE CITED

Cox, J. J. 1981. Notes on coprophilous Myxomycetes from the western United
States. Mycologia 73:741-747.

ELIASSON, U. 1977. Ecological notes on Amaurochaete Rost. (Myxomycetes). Bot.
Notiser 129:419-425.

FARR, M. L. 1982. Notes on Myxomycetes. III. Mycologia 74:339-343.

GOTTSBERGER, G. 1968. Myxomyceten aus Bahia und Goias. Nov. Hedw. XV:361-
370.

HAGELSTEIN, R. 1944. The Mycetozoa of North America. Publ. by the author,
Mineola.

HARKONEN, M. 1977. Comatricha nannengae, a new species of Myxomycetes. Kar-
stenia 17:87-89.

KowaALsKI, D. T. 1973. New records of Myxomycetes from California. V. Madrono
22:97-100.

LisTER, A. 1925. A monograph of the Mycetozoa, ed. 3. Revised by G. Lister. Brit.
Mus. Nat. Hist., London.

MarTINn, G. W. and C. J. ALEXOPOULOS. 1969. The Myxomycetes. Univ. Iowa
Press, Iowa City.

MEYLAN, C. 1921. Contribution a la connaissance des Myxomycétes de la Suisse.
Bull. Soc. Vaud. Sci. Nat. 53:451-463.

MITCHELL, D. 1977. Kentish Myxomycetes. Trans. Kent Field Club 6(2):91-100.

NANNENGA-BREMEKAMP, N. E. 1972. Notes on Myxomycetes. XVIII. A new Di-
dymium and some comments on the Didymium species with long-stalked spo-
rangia. Proc. K. Ned. Akad. C, 75(4):352-363.

1981. Notes on Myxomycetes. XX. A new Licea and its associates from

France. Proc. K. Ned. Akad. C, 84(3):285-288.

and C. Labo. 1985. Notes on some Myxomycetes from Central Spain. Proc.

K. Ned. Akad. C, 88(2):219-231.

, K. G. MUKERJI, and R. PAsSRICHA. 1984. Notes on Indian Myxomycetes.

Three new species, and comments on others. Proc. K. Ned. Akad. C, 84(4):471-

482.

and Y. YAMAMOTO. 1983. Additions to the Myxomycetes of Japan. I. Proc.
K. Ned. Akad. C, 86(2):207-241.

WELDEN, A. L. 1954. Some Myxomycetes from Panama and Costa Rica. Mycologia
46:93-99.

WHITNEY, K. D. 1980. The Myxomycete genus Echinostelium. Mycologia 72:950-
987.

. 1982. A survey of the corticolous Myxomycetes of California. Madrono
29:259-268.

(Received 22 Jan 1986; revision accepted 6 Aug 1986.)

COLD TOLERANCE IN THE DESERT FAN PALM,
WASHINGTONIA FILIFERA (ARECACEAE)

JAMES W. CORNETT
Natural Science Department, Palm Springs Desert Museum,
101 Museum Drive, Palm Springs, CA 92263

ABSTRACT

Natural populations of Washingtonia filifera tolerate temperatures down to — 11°C
and subfreezing temperatures for at least 22 hours. Nonacclimatized seedlings survive
temperatures as low as —21°C for one hour. Seeds are hardy, germinating readily
after 36 hours of exposure to a temperature of —21°C. Available climatic data suggests
that the absence of W. filifera from springs and seeps in the eastern Sonoran Desert
and portions of the Mojave Desert is for reasons other than simple intolerance to
subfreezing temperatures.

The desert fan palm, Washingtonia filifera (Lindl.) Wendl., of the
Sonoran Desert is known to tolerate subfreezing temperatures. Muir-
head (1961) stated that adults survived temperatures to at least —9°C
with small plants showing leaf burn at temperatures below —4°C.
Blombery and Rodd (1982) believed that desert fan palms survived
winter temperatures down to —5°C. Although these references pro-
vide some information on the minimum temperature tolerance of
W. filifera, no data have been presented on the duration of the
subfreezing temperatures. The purpose of this paper is to present 1)
new information on the minimum temperatures and the duration
of subfreezing temperatures tolerated by natural populations of W.
filifera, and 2) the results of tests on the cold tolerance of both seeds
and seedlings. I use these data to evaluate the effect of temperature
on the present distribution of W. filifera (Fig. 1).

METHODS

Minimum temperatures that occurred in the vicinity of six palm
oases were obtained from seven meteorological shelters within the
Sonoran Desert of southeastern California and western Arizona
[Agave Hill, Pinyon Crest, and Taylor Site data obtained from the
Boyd Deep Canyon Desert Research Center, Palm Desert, CA; Kofa
and Castle Creek data from Sellers and Hill (1974); Oasis of Mara
data obtained from Joshua Tree National Monument Headquarters;
Indio data from U.S. Date and Citrus Station (1981)]. Locations of
oases and shelters are given in Table 1. With two exceptions, each
palm oasis was situated within 2 km and at approximately the same
elevation as one of the weather shelters. Owl Hole was approximately

MADRONO, Vol. 34, No. 1, pp. 57-62, 1987

58 MADRONO [Vol. 34

* DESERT FAN PALM OASIS

erro tte

THE COLORADO DESERT

Fic. 1. The Colorado Desert subdivision of the Sonoran Desert (shaded area).
* = location of desert fan palm oases.

40 m higher and 7 km north of the weather shelter at Indio, Riverside
Co., CA. Kofa Palm Canyon was 144 m higher and 12 km north of
the weather shelter located in the settlement of Kofa, Yuma Co.,
AZ. Three of the shelters (Agave Hill, Taylor Site, and Pinyon Crest)
enclosed continuous recording thermographs and, thus, recorded the
duration of subfreezing temperatures.

Minimum cold tolerance of W. filifera seeds was determined by

1987] CORNETT: COLD TOLERANCE IN WASHINGTONIA 59

TABLE 1. MINIMUM TEMPERATURES AND DURATION OF SUBFREEZING TEMPER-
ATURES EXPERIENCED BY POPULATIONS OF Washingtonia filifera IN S1xX PALM OASES
LOCATED IN THE SONORAN DESERT. D = duration; BDCDRC = Boyd Deep Canyon
Desert Research Center, Univ. California, Riverside.

Min. Ele-
temp. D Palm oasis vation Date of Shelter
"©. (hi) location (m) minimum location

—03 15 CA, Riverside Co.: 938 28 Jan 1979 CA, Riverside Co.:
Santa Rosa Moun- BDCDRC, Agave
tains, Hidden Hill
Palms Canyon

—04 14 Hidden Palms Can- 938 2Jan 1974 Agave Hill
yon

—04 14 CA, Riverside Co.: 1066 20 Jan 1983 CA, Riverside Co.::
Santa Rosa Moun- BDCDRC, Pin-
tains, Dos Palmas yon Crest
Spring

—05 ? AZ, Yuma Co.: Kofa 685 2 AZ, Yuma Co.: Kofa

Palm Canyon
—06 18 Dos Palmas Spring 1066 29 Dec 1982 Pinyon Crest

—08 ? AZ, Yavapai Co.: 621 ? AZ, Yavapai Co.:
Castle Creek Castle Hot Springs

—09 22 Dos Palmas Spring 1066 29 Jan 1979 BDCDRC, Taylor

Site

—11 ? CA, San Bernardino 600 3 Jan 1974 CA, San Bernardino
Co.: Joshua Tree Co.: Twentynine
Nat. Mon., Oasis Palms, Mon. Head-
of Mara quarters

—11 ? CA, Riverside Co.: 37 22 Jan 1937 CA, Riverside Co::
Indio Hills, Owl Indio
Hole

placing them in a freezer for the durations and temperatures shown
in Table 2. Following cold exposure, the seeds were planted in plastic
containers filled with a mixture of compost and vermiculite in equal
proportions. The containers were placed on a heating pad that main-
tained a constant temperature of 32°C. The bedding medium was
saturated with water on alternate days. Daily inspections were made

TABLE 2. DURATION OF MINIMUM TEMPERATURE EXPOSURES OF Washingtonia
filifera SEEDS.

Germination
success
Min. temp. °C Duration (h) #/total %
07 24 38/40 95
(07 96 18/20 90
=) 06 38/40 95

=| 36 17/20 85

60 MADRONO [Vol. 34

TABLE 3. DURATION OF MINIMUM TEMPERATURE EXPOSURES OF Washingtonia
filifera SEEDLINGS.

Survival rate

Min. temp. °C Duration (h) #/total %
le 1.0 6/20 30
12 3.0 0/19 0
= 6.0 0/19 0
=15 0.5 20/20 100
=?) 1.0 4/19 21

to determine whether sprouting had occurred. Cold tolerance of
seedling palms was determined by placing 20-30 day-old sprouts in
a freezer for the durations and temperatures shown in Table 3.

RESULTS

Weather records indicate the minimum temperature tolerance of
natural populations of W. filifera to be at least — 11°C, two degrees
lower than the known minimum of —9°C (Table 1). In addition, it
appears that adult trees can withstand up to 22 h of subfreezing
temperatures.

Palm seeds germinated readily after exposure to subfreezing tem-
peratures. In these studies, seeds tolerated exposures as low as — 21°C
for 36 h (Table 2). Seeds exposed to such low temperatures also
tended to germinate earlier (kx = 21.5 days; s.d. = 2.6) than seeds
not exposed to subfreezing temperatures (x = 27.5 days; s.d. = 5.1).
These means are significantly different (t-test, p < 0.001). Seeds also
germinated readily following exposure to freezing temperatures for
seven days with temperatures dropping to — 30°C (A. Stumpf, pers.
comm.). Palm seedlings were less tolerant of cold, although four
seedlings survived one hour exposure at —21°C (Table 3). Percent
survival may have increased had the seedlings been acclimatized
prior to exposure to freezing temperatures.

DISCUSSION

At least three populations of W. filifera probably experience even
colder temperatures and for longer durations than do the palm oases
listed in Table 1. Fortynine Palms oasis in Joshua Tree National
Monument is located 6 km west of the Oasis of Mara and 206 m
higher at 878 m. Because of its higher elevation and decreased ex-
posure to direct sunlight due to its canyon location, temperatures
are probably colder for longer periods at Fortynine Palms than at
the Oasis of Mara. Mopah Spring, located in the Turtle Mountains
of San Bernardino Co., CA, lies 24 km north and 100 m higher than
the Oasis of Mara. Some individuals in Munsen Canyon (Joshua

1987] CORNETT: COLD TOLERANCE IN WASHINGTONIA 61

Tree National Monument) occur at approximately 1015 m, which
is the second highest elevation recorded for adult W. filifera. Al-
though no climatic data exist for this site, the upper end of the
Munsen Canyon palm grove may experience colder temperatures
than do the high-altitude palms at Dos Palmas Spring in the Santa
Rosa Mountains.

Low winter temperature is a probable factor in the exclusion of
W. filifera from certain regions adjacent to its present range. Desert
fan palms line many of the eastward-trending canyons that drain
the Peninsular Ranges of southern California and adjacent Baja Cal-
ifornia Norte. Perennial streams exist in over a dozen of these can-
yons. Palms occur at the lowest point where water appears on the
surface and they grow along the streams up to an elevation of ap-
proximately 900 m; they are absent above 1000 m even though the
streams continue well above this elevation. Palms also are absent
from most of the Mojave Desert where winter temperatures occa-
sionally drop to — 13°C or lower (U.S. Weather Bureau 1951, 1980,
Sellers and Hill 1974). These observations, combined with the data
from Table 3, suggest that the distribution of W. filifera is affected
by low winter temperatures, but not to the degree that the tropical
association of its family might indicate. How cold temperatures
affect the palms is not known. Possibly, the seedlings may fail to
establish because of a reduction in competitiveness that results from
retarded growth or because of frost damage to the apical meristem.

The information presented on low temperature tolerances of W.
filifera indicates, insofar as temperature is concerned, that the dis-
tribution of this species could be broader than it is at present. For
example, there are no low temperature records that preclude the
widespread occurrence of desert fan palms at springs and streams
within the Sonoran Desert of Arizona and northern Mexico (Sellers
and Hill 1974, Steinhauser 1979). The presence of two palm oases
in Arizona, at the Kofa Mountains and Castle Creek, indicates that
climatic conditions are suitable for W. filifera in the Sonoran Desert.
Potential habitat also exists in the Colorado River drainage region
of southern Nevada and the Death Valley area in the Mojave Desert.
Geographical and ecological barriers, the lack of efficient dispersal
agents, or insufficient time may account for the absence of the desert
fan palm in these regions.

ACKNOWLEDGMENTS

This study was made possible through a grant from the Richard King Mellon
Foundation of Pittsburgh, Pennsylvania. The author wishes to thank Allan and Vic
Muth of the Boyd Deep Canyon Desert Research Center for providing important
climatic data. Theo Glenn and Jule Anne Huffnagle assisted in the germination
studies.

62 MADRONO [Vol. 34

LITERATURE CITED

BLomBeErRY, A. and T. Ropp. 1982. Palms. Angus and Robertson Publishers, Lon-
don.

MUIRHEAD, D. 1961. Palms. Dale Stuart King Publisher, Globe, AZ.

NATIONAL CLIMATIC CENTER. 1980. Local climatological data: Las Vegas, NV. U.S.
Dept. of Commerce, Asheville, NC.

SELLERS, W. D. and R. H. HILL. 1974. Arizona climate 1931-1972. Univ. Arizona
Press, Tucson.

STEINHAUSER, F. 1979. Climatic atlas of North and Central America. World Me-
teorological Organization, Geneva, Switzerland.

U.S. DATE AND CITRUS STATION. 1981. Climatological summary for Indio. U.S.
Date and Citrus Station, Indio, CA.

U.S. WEATHER BUREAU. 1951. Climatological summary for Barstow. U.S. Dept.
Commerce, Asheville, NC.

(Received 6 Jan 1986; revision accepted 14 Aug 1986.)

ANNOUNCEMENT
NEw PUBLICATIONS

Cope, E. A., Native and cultivated conifers of northeastern North Amer-
ica: A guide, Cornell Univ. Press, 124 Roberts Place, Ithaca, NY
14850, 1986, 231 pp., illus., ISBN 0-8014-1721-X, $39.95 (hard-
bound), ISBN 0-8014-9360-9, $17.95 (paperbound).

Creso, I., Vascular plants of western Washington, Irene Creso Publisher,
[Creso Herbarium, Pacific Lutheran Univ., Tacoma, WA], 1984, xviii,
532 pp., illus., ISBN 0-9613916-0-X, $14.95 (paperbound).

GiBSON, A. C. AND P. S. NoseL, The cactus primer, Harvard Univ.
Press, 79 Garden St., Cambridge, MA 02138, 1986, 1x, 286 pp.., illus.,
ISBN 0-674-08990-1, $39.95 (hardbound).

MartTIN, W. C. AND C. R. HUTCHINS, Summer wildflowers of New Mex-
ico, Univ. New Mexico Press, Albuquerque, NM 87131, 1986, v, 318
pp., 16 pls. (color), illus. (B&W), ISBN 0-8263-0859-7, $24.95 (hard-
bound), ISBN 0-8263-0860-0, $12.95 (paperbound). [=New Mexico
natural history series. |

PHOSPHORUS AND PH TOLERANCES IN THE
GERMINATION OF THE DESERT SHRUB
LARREA TRIDENTATA (ZYGOPHYLLACEAE)

KATE LAJTHA,! JOHN WEISHAMPEL, AND WILLIAM H. SCHLESINGER
Department of Botany, Duke University,
Durham, NC 27706

ABSTRACT

Seeds of Larrea tridentata, a dominant shrub of deserts in the southwestern U.:S.,
were germinated on both a pH and phosphorus (P) gradient to determine if require-
ments for germination can help explain the field distribution of Larrea. Germination
decreased significantly above pH 8, which is consistent with the conspicuous absence
of Larrea from high pH sodic or saline desert soils. Although Larrea tends to be
absent from noncalcareous soil, seed germination was not inhibited in acidic solutions.
Germination showed no response to P or to interactions of pH and P. In contrast,
recent literature has suggested that Larrea may be restricted to calcareous soils of
low phosphorus availability due to toxicity of high concentrations of P to seedlings.

Larrea tridentata (Sessé & Moc. ex DC.) Cov. (creosote bush) is
one of the most abundant and widely distributed shrubs of south-
western deserts (Runyon 1934); its limits have been used to define
the warm desert region of North America (Benson and Darrow 1954).
Within its range, however, Larrea-dominated communities often
exhibit sharp boundaries, and a complete transition to other com-
munities may be seen within 5-10 meters (Barbour 1969). Age dis-
tributions of Larrea in mature communities and observations of
germination in the field indicate that germination and survival of
seedlings are rare events under natural conditions (Barbour 1969,
Ackerman 1979, Boyd and Brum 1983, Goldberg and Turner 1986)
and suggest that germination could affect the distribution of Larrea.

Several authors have shown that soils in areas dominated by Lar-
rea are porous and have greater drainage and aeration than do soils
of adjacent areas (Yang and Lowe 1956, Fosberg 1940, Lunt et al.
1973). Others have observed that Larrea is found on soils that are
generally calcareous throughout the profile (Hallmark and Allen 1975,
Gardner 1958). Larrea apparently has no unusual physiological de-
mands for calcium (Ca) (El-Ghonemy et al. 1978), but soil CaCO,
may modify physical and chemical soil properties that are essential
to the survival of Larrea (Hallmark and Allen 1975, Johnson 1961).

One potential effect of free CaCO, in soil is the fixation of available
phosphorus (P) onto carbonates. Chemical interactions between Ca
and P have been well documented, both in experimental solutions
and in natural systems (Griffin and Jurinak 1973, Avnimelech 1983,

' Current address: Dept. Botany, Ohio State Univ., Columbus 43210.

MADRONO, Vol. 34, No. 1, pp. 63-68, 1987

64 MADRONO [Vol. 34

Cole and Olsen 1959). Carbonates affect P levels in solutions through
ion pairing with Ca, physical sorption, and the precipitation of cal-
cium phosphate minerals (Marion and Babcock 1977). Fixation of
P by soil carbonates may lower crop response to P fertilization in
the southwest (McCaslin and Gledhill 1980, Chang 1953).

Musick (1978) found that P was toxic to Larrea seedlings at fairly
low external concentrations (10 uM) in slightly acid solution cultures
(pH 6). Because there was no toxicity response at pH 8, Musick
suggested that Larrea is adapted to alkaline, calcareous soils of low
P availability. In a study of the germination requirements and tol-
erances of Larrea seeds, Barbour (1968) found no differences in
germination success in the pH range of 7-10. Barbour, however,
used phosphate buffers of an unreported P concentration to establish
these solution pH values. In the light of Musick’s (1978) finding of
a significant effect of P in seedling growth, we have reevaluated the
germination tolerance of Larrea with respect to both pH and various
levels of P.

METHODS

Seeds of Larrea tridentata were collected in July 1984 from shrubs
along a bajada of the Jornada Experimental Range of New Mexico
State University near Las Cruces, New Mexico. Fruits from ca. 30
shrubs were mixed and stored in paper bags at room temperature
until germination experiments began.

Experimental methods followed those of Barbour (1968), with
several modifications. Whereas Barbour used whole mericarps in
his germination trials, the high incidence of empty mericarps in our
collections prohibited this technique. Mericarps were cracked open,
and only mature seeds whose lengths were approximately 3 to 5 mm
were used in the experiment.

Seeds were germinated along both a pH and a P gradient in a
2-way factorial design. The Modified Universal Buffer (MUB) of
Skujins et al. (1963), a phosphate-free buffer used widely in phos-
phatase enzyme studies (Tabatabai and Bremner 1969), was used
to establish the pH of the experimental solutions. Solutions of pH
7,8, 9, and 10 were used as in Barbour (1968), as well as an additional
treatment of pH 4.5. Buffers were mixed 1:1 with half-strength mod-
ified Hoagland’s solution (Downs and Hellmers 1975) with a sub-
sequent readjustment of pH. Treatment levels of P were 1) 1 uM,
derived from the quarter-strength modified Hoagland’s solution with
no additional P added; 2) 10 uM; and 3) 100 uM PO,-P, added as
KH,PO,. These concentrations span the range of P levels used by
Musick (1978).

In each treatment, we used 100 seeds grouped into 5 lots of 20
seeds each. As in Barbour (1968), seeds were soaked for 3 hours in

1987] LAJTHA ET AL.: GERMINATION REQUIREMENTS IN LARREA _ 65

TABLE 1. NUMBER OF SEEDS THAT GERMINATED AFTER 5 AND 10 DAYS IN EACH
PH x PHOSPHORUS TREATMENT. (abcde) values with the same letter within the pH x
phosphorus factorial cross on each day are not significantly different by Duncan’s
multiple range test (5% level). (ABC) values with the same letter for pH treatments
on each day are not significantly different by Duncan’s multiple range test (5% level).
* = one-way ANOVA indicates no significant differences (5% level) between values
for phosphorus treatments on each day.

Day 5
Mean
Phosphorus concentration over all
a a ee ee ee DP ROSphonus
pH 1 uM 10 uM 100 uM treatments
4.5 12.0% 15282 14.42 1421“
q ee ee PieG? 1220" 12.44
8 13-622" Oss 16.2? 13.44
9 13:07 8.0% 829° 9.78
10 4.8° 6.84 6.8% ropa lee
Mean over all
pH treatments | et [0237 Se
Day 10
Mean
Phosphorus concentration over all
ee es i Se ee ee ae = SO DNOSPNOLUS
pH 1 uM 10 uM 100 uM treatments
4.5 152872 7A? 16.62 16.64
fi joy Olas 1627 14. 4abed 15:65
8 16.62 o.2°s4 17.02 522°
9 14 feted jslie steers 11.4¢¢ [Pees be
10 8.6° 8.6° 8.8° 8.75
Mean over all
pH treatments 14.1% 13.4* 1326"

the appropriate treatment solution, and each set of 20 seeds was
transferred to a petri dish filled with sand moistened with the same
solution. All dishes were incubated in darkness at 25°C. We tallied
germinations after both 5 and 10 days, and observed no germination
after this period.

Statistical analyses were performed using the ANOVA procedure
of SAS (SAS Institute Inc. 1982). The number of seeds that ger-
minated in each lot of 20 seeds was treated as one observation.
When the ANOVA indicated statistical significance, Duncan’s mul-
tiple range test was used to distinguish differences among treatments.

RESULTS

Although pH had a significant effect on germination (p < 0.0001),
there was no effect of P concentration (Table 1). There were no

66 MADRONO [Vol. 34

significant differences in germination among pH 4.5, 7, and 8, but
the number of seeds that germinated declined sharply at pH 9 and
10. The ANOVA for day 5 counts showed a significant interaction
between pH and P (p < 0.0004), possibly due to a high mean ger-
mination value for pH 9 (1 uM PO,-P solutions) and a high value
for pH 8 (100 uM solutions). The interaction was not significant on
day 10, and we believe that it occurred as a result of random vari-
ation.

DISCUSSION

Differences in methodology may account for the disparity between
our results and those of Barbour (1968), who found no effect of pH
on germination over the range of pH 7-10. Although his germination
trials lasted 5 days, we found significant germination occurring be-
tween 5 and 10 days in seeds extracted from mericarps. Barbour
(1968) found that root growth decreased greatly with increasing pH,
especially above pH 8. The response curve of root growth to pH
(Fig. 5 in Barbour 1968) is remarkably similar to the response of
germination to pH found in this study. The lack of response of
germination to P concentration probably reflects the high internal
stores of P in seeds of Larrea (Barbour 1968, Musick 1978). Thus,
nutrient absorption does not become significant until seedling emer-
gence.

Our data show that relatively acid solution (pH 4.5) did not inhibit
the germination of Larrea seeds. Barbour (1968) found that root
growth was greater in slightly acidic solutions (pH 6) than in those
of higher pH. Thus, germination and early root development in
response to pH cannot be used to explain the distribution patterns
of Larrea found by Hallmark and Allen (1975), who showed that
shrubs were restricted to soils that were calcareous in the upper
10 cm.

Solutions of pH 9 to 10 frequently reduced germination to less
than 50% of maximum, certainly an important reduction for a species
that has no significant seed bank (Boyd and Brum 1983) and few
years that are favorable for germination and establishment in the
field (Ackerman 1978). This result is consistent with known Larrea
distribution patterns. Calcic soils have pH’s in the range of 8-8.4,
and soil pH will be higher only when significantly sodic or saline.
Larrea is conspicuously absent from saline soils near topographic
lows and playas (Barbour 1969, Fosberg 1940, Went and Wester-
gaard 1949).

Many of the environmental variables that appear to affect Larrea
distributions are correlated, and thus it is difficult to discern cause-
and-effect relationships in nature. Calcareous soils tend to have a
well-buffered pH range and are often coarse-grained with good in-

1987] LAJTHA ET AL.: GERMINATION REQUIREMENTS IN LARREA_ 67

ternal drainage, whereas soils of a higher pH tend to be saline and
fine-grained, with a lower permeability. Lunt et al. (1973) showed
that Larrea has a relatively high oxygen requirement for root growth,
and the correlation between CaCO, and Larrea occurrence may be
due to the improved aeration and root penetration in CaCO,-rich
soils rather than to the direct presence of CaCO, or to the buffering
of soil pH by CaCO, (Lunt et al. 1973, Johnson 1961). Our data
suggest that soil pH does not limit the germination of Larrea in acid,
non-calcareous soils of southwestern deserts, although soil pH may
interact with other factors to determine successful seedling estab-
lishment and growth.

ACKNOWLEDGMENTS

This work was supported by a Sigma Xi Grant-in-Aid of Research to KL and NSF
Grant BSR 8212466 to WHS. We thank Peter Vitousek, Brad Musick and anonymous
reviewers for many helpful comments on the manuscript.

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ACKERMAN, T. L. 1979. Germination and survival of perennial plant species in the
Mojave Desert. Southwestern Natur. 24:399-408.

AVNIMELECH, Y. 1983. Phosphorus and calcium carbonate solubilities in Lake Kin-
neret. Limnol. Oceanogr. 28:640-645.

BARBOUR, M. G. 1968. Germination requirements of the desert shrub Larrea di-
varicata. Ecology 49:915-926.

. 1969. Age and space distribution of the desert shrub Larrea divaricata.
Ecology 50:679-685.

BENSON, L. and R. A. DARROW. 1954. The trees and shrubs of the southwestern
deserts, 2nd ed. Univ. Arizona Press, Tucson.

Boypb, R. S.and G. D. BRUM. 1983. Postdispersal reproductive biology of a Mojave
desert population of Larrea tridentata (Zygophyllaceae). Amer. Midl. Natur. 110:
25-36.

CHANG, C. W. 1953. Chemical properties of alkali soils in Mesilla Valley, New
Mexico. Soil Science 75:233-242.

Coe, C. V. and S. R. OLSEN. 1959. Phosphorus solubility in calcareous soils: I.
Dicalcium phosphate activities in equilibrium solutions. Soil Sci. Soc. Amer.
Proc. 23:116-118.

Downs, R. J. and H. HELLMeErRS. 1975. Environment and the experimental control
of plant growth. Academic Press, New York.

EL-GHONEMY, A. A., A. WALLACE, and E. M. ROMNEY. 1978. Nutrient concentra-
tions in the natural vegetation of the Mojave Desert. Soil Sci. 126:219-229.
FOSBERG, F.R. 1940. The aestival flora of the Mesilla Valley Region, New Mexico.

Amer. Midl. Natur. 23:573-593.

GARDNER, J. L. 1958. Ecobiology of the arid and semiarid lands of the southwest.
New Mexico Highlands Univ. Bull. 212.

GOLDBERG, D. E. and R. M. TURNER. 1986. Vegetation change and plant demog-
raphy in permanent plots in the Sonoran Desert. Ecology 67:695—7 12.

GRIFFIN, R. A. and J. J. JURINAK. 1973. The interaction of phosphate with calcite.
Soil Sci. Soc. Amer. Proc. 37:847-850.

HALLMARK, C. T. and B. L. ALLEN. 1975. The distribution of creosotebush in west
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JOHNSON, D. C. 1961. Edaphic factors affecting the distribution of creosote bush
(Larrea divaricata) (DC.) Cov. in desert grassland sites of southern Arizona. MS
thesis, Univ. Arizona, Tucson.

Lunt, O. R., J. LETEY, and S. B. CLARK. 1973. Oxygen requirements for root growth
in three species of desert shrubs. Ecology 54:1356-1362.

MARION, G. M. and K. L. BABcock. 1977. The solubilities of carbonates and
phosphates in calcareous soil suspensions. Soil Sci. Soc. Amer. J. 41:724-728.

McCASLIN, B. D. and R. J. GLEDHILL. 1980. Alfalfa fertilization in New Mexico.
Bulletin 675, Agricultural Experiment Station, Las Cruces, NM.

Musick, H. B. 1978. Phosphorus toxicity in seedlings of Larrea divaricata grown
in solution culture. Bot. Gaz. 139:108-111.

RuNYON, E. H. 1934. The organization of the creosote bush with respect to drought.
Ecology 15:128-138.

SAS INsTITUTE INc. 1982. SAS user’s guide: statistics, 1982 edition. SAS Inst. Inc.,
Cary, NC.

SKUJINS, J. J., L. BRAAL, and A.D. MCLAREN. 1963. Characterization of phosphatase
in a terrestrial soil sterilized with an electron beam. Enzymologia 25:125-133.

TABATABAI, M. A. and J. M. BREMNER. 1969. Use of p-nitrophenyl phosphate for
assay of soil phosphatase activity. Soil Biol. Biochem. 1:301-307.

WENT, F. W. and M. WESTERGAARD. 1949. Ecology of desert plants. III. Devel-
opment of plants in the Death Valley National Monument, California. Ecology
30:26-38.

YANG, T. W. and C. H. Lowe, Jr. 1956. Correlation of major vegetation climaxes
with soil characteristics in the Sonoran Desert. Science 123:524.

(Received 10 Feb 1986; revision accepted 5 Aug 1986.)

ANNOUNCEMENT
NEW PUBLICATION

RZEDOWSKI, J. and G. C. DE RZEDOwSKI (eds.), Flora Fanerogamica del
Valle de México, Vol. 2, Dicotyledoneae (Euphorbiaceae-Compositae),
Instituto de Ecologia, AP 18-845, Deleg. Miguel Hidalgo, CP 11800,
Mexico, D. F., 1985, 674 pp., illus., ISBN 968-7213-02-7, US $35.00
(hardbound). [The second volume of a proposed three-volume flora.
Volume | (publ. Mar. 1979, reprinted 1984, 432 pp., US $28.00,
source above) provided introductory information on topography, ge-
ology, climate, plant communities, etc., and a floristic treatment of
gymnosperms and of dicotyledons from Saururaceae to Polygalaceae.
Volume 3 will include the monocotyledons. ]

NOTEWORTHY COLLECTIONS

CALIFORNIA

MANNIA FRAGRANS (Balb.) Frye & Clark (HEPATICOPSIDA: AYTONIACEAE).— Tuo-
lomne Co., mountain on n. side of Tioga Pass, Yosemite National Park, 10,000 ft,
4 Jul 1932, M. S. Baker 5809 (UC). Mono Co., H. M. Hall Natural Area near Stanford
Research Area, TIN R24E S1, moist diffusely lit soil under boulder, in lodgepole
pine forest and adjacent meadows, ca. 10,000 ft, 3 Jun 1977, D. H. Norris 48446
(HSC); Tioga Jct. Campground about 2.5 mi n. of Tioga Pass on hwy. 120, TIN
R25E S19, moist, sunny soil between boulders, in open meadows and adjacent rock
outcrops, ca. 9500 ft, 2 Jun 1977, D. H. Norris 48355 (HSC). Inyo Co., slopes
above Treasure Lake, w. of Big Pine, T9S R31E S27, on temporarily moist, sunny
soil at base of boulder, in alpine fell field with scattered Pinus albicaulis, ca. 11,500
ft, 6 Sep 1975, D. H. Norris 47006 (HSC, JE) (verified by R. Grolle, JE).

Previous knowledge. Eurasia; e. North America w. to Colorado, and perhaps Idaho
and the Yukon (Schuster, American Midl. Nat. 59:274, 1958).

Significance. First records for CA. This species should be looked for farther n.; I
have seen sterile specimens apparently belonging here from Modoc Co., CA and
Josephine Co., OR (Norris 22473 and 52495, respectively, both HSC).

MYLIA ANOMALA (Hook.) S. F. Gray (HEPATICOPSIDA: JUNGERMANNIACEAE). — Hum-
boldt Co., Sphagnum bog at Big Lagoon Co. Park, T9N R1W S13, near sea level, 2
Jun 1977, D. H. Norris 48316 (HSC).

Previous knowledge. Circumboreal, s. on the Pacific coast to WA (Schuster, The
Hepaticae and Anthocerotae of North America East of the Hundredth Meridian
2:1040, 1969).

Significance. First record for CA.

PREISSIA QUADRATA (Scop.) Nees (HEPATICOPSIDA: MARCHANTIACEAE). — Siskiyou
Co., Salmon River near Big Flat, T37N R9W S18, on moist sunny seepage on schis-
tose wall of stream, ca. 5000 ft, 4 Aug 1968, D. H. Norris 9185 (HSC); in moist
canyon along Jaynes Creek to headwaters (nw. of Klamath River), T47N R3W S13,
very moist diffusely lit boulder along stream in white fir forest, ca. 5000-6000 ft, 10
Jul 1977, D. H. Norris 48690 (HSC). Mono Co., meadow above Lake Mildred, Convict
Creek drainage, 1 18°52.3'W, 37°32.6'N, bedrock limestone, 9800 ft, 20 Sep 1981, A.
T. Whittemore 1496 (DAV), 1498A (CAS), 1516 (to be distributed), 1521] (to be
distributed), 1540 (CAS).

Previous knowledge. Circumboreal, s. in the Pacific states to OR (Hong, Bryologist
81(3):441, 1978).

Significance. First records for CA.

MExICco

ATHALAMIA HYALINA (Sommerf.) Hatt. HEPATICOPSIDA: CLEVEACEAE). — Nuevo Leon,
Puente de Dios, ca. 8 km n. of Galeana on road to Rayones, 100°05.5'W, 24°53’N,
sheltered grotto in canyon wall, arid scrub with Cupressus, 1500 m, 27 Aug 1984, A.
T. Whittemore 2469, M. Lavin, and T. Atkins (MEXU).

Previous knowledge. Circumboreal, s. in the Rocky Mountain system to CO and
AZ (Schuster, Amer. Midl. Nat. 59:298, 1958, as Clevea hyalina).

Significance. First record for Mexico.

RICCIA ALBIDA Sull. (HEPATICOPSIDA: RICCIACEAE). — Nuevo Leon, junction of Mon-
terrey bypass and hwy. 40, ca. 16 mi w. of Monterrey, 100°33.5'W, 25°42'N, rather

MADRONO, Vol. 34, No. 1, pp. 69-70, 1987

70 MADRONO [Vol. 34

open soil near shrubs, low land above banks of river, in thorn scrub, 880 m, 12 Mar
1982, A. T. Whittemore 1599, T. Ayers, F. Barrie, and D. Lemke (MEXU).
Previous knowledge. Edwards Plateau of Texas (McGregor and Menhusen, Bryol-
ogist 64:71-74, 1962) and e. LA (Guerke, Bryologist 74:203, 1971).
Significance. First record for Mexico.

CAMPYLIUM HALLERI (Hedw.) Lindb. (BRYOPSIDA: AMBLYSTEGIACEAE). — Nuevo Leon,
cliffs just ne. of the microwave station, summit of Cerro Potosi, ca. 17 km nw. of
Galeana, 100°14'W, 24°52'4'N, on rock of cliffin Pinus hartwegii forest, 3700 m, 23
Aug 1984, A. T. Whittemore 2373, M. Lavin, and T. Atkins (MEXVU) (verified by C.
Delgadillo M., MEXU).

Previous knowledge. Circumboreal, s. in the Rocky Mountains to CO (Crum &
Anderson, Mosses of Eastern North America, p. 947, 1981).

Significance. First record for Mexico.

ENTODON SCHLEICHERI (Schimp.) Demeter (BRYOPSIDA: ENTODONTACEAE). — Nuevo
Leon, on road up Cerro Potosi 1.5 mi below the lower microwave tower, ca. 17 km
nw. of Galeana, 100°13.5’W, 24°53’N, on rock in grove of Populus tremuloides, Pinus-
Abies forest with grazed herbaceous understory, 3100 m, 24 Aug 1984, A. T. Whitte-
more 2396, M. Lavin, and T. Atkins (MEXU, CAS).

Previous knowledge. Eurasia and Lincoln Co., NM (Buck & Crum, Bryologist 81:
429-432, 1978).

Significance. First record for Mexico.

TIMMIA MEGAPOLITANA Hedw. subsp. BAVARICA (Hessl.) Brassard (BRYOPSIDA: TIM-
MIACEAE).— Coahuila, atop high cliffs on n. side of summit of Sierra Coahuilon, ca.
25 km (by air) e. of San Antonio de las Alazanas, 100°20’W, 25°15’N, in rock
crevice beneath overhang in open Pinus-Picea forest with Lupinus and Senecio on
soil and Heuchera on rock faces, 3500 m, 23 Jul 1985, A. 7. Whittemore 2686, A.
McDonald, S. Boykin, and S. Ginzbarg (MEXU, CAS, TEX) (verified by C. Delgadillo
M., MEXU).

Previous knowledge. Circumboreal, s. in the Rocky Mountains to CO and AZ; also
a single collection from the state of Mexico, Mexico (Brassard, Lindbergia 10:33-—40,
1984).

Significance. Fills in a gap of almost 2000 km between the single Mexican report
and the nearest sites in the s. Rocky Mountains.— ALAN T. WHITTEMORE, Dept. of
Botany, Univ. of Texas, Austin 78712.

REVIEWS

Flora of the Santa Monica Mountains, California. By PETER H. RAVEN, HENRY J.
THOMPSON, and BARRY A. PRIGGE. Southern California Botanists, Special Publication
No. 2. Second Edition, June 1986. $8. (paperbound).

This flora was published first in 1966 as an introductory manual for beginning
students at UCLA, etc. and was revised in 1977. Since 1966, 72 species have been
added, of which 45 are native, now bringing the total to 880 species. A major revision
was undertaken by the junior author, Barry A. Prigge, and resulted in many nomen-
clatoral changes in the species, genera, and even at the family level.

MaApDrRONO, Vol. 34, No. 1, pp. 70-74, 1987

1987] REVIEWS ()

There is a map of the Santa Monica Mountains, a special area close to some five
million people. Concerned citizens have set aside many local government parks,
including 33,000 acres in State Parks and 70,000 acres as a National Recreation Area.

The introduction follows with an in-depth explanation of the Mediterranean cli-
mate, and a summation of the geology. The vegetation is ably discussed with black
and white photographs and a table of Raunkiaer’s life forms. There is a statistical
summary of the flora. The catalogue is alphabetical, with families at the top of each
page for easy reference, an asterisk before introduced species with countries of origin,
and numbers for months indicating the flowering periods. A glossary is added with
an extensive list of literature cited, and an index is included.

The main part of the flora has ample keys to the divisions, classes, families, genera,
and species of vascular plants. Obvious characters are used to simplify the keys when
applicable and they are not unduly technical unless needed. The local distribution is
usually given in an easy flowing simple declarative sentence. A few are vague: Lep-
idospartum squamatum (A. Gray) A. Gray is quoted as “Occasional in coastal sage
scrub and southern oak woodland”, . . . ending with “‘Often in washes and along
streams in sandy soil.’ In my opinion, this plant is rarely found outside washes in a
given plant community. I would like to see every plant with an exact habitat(s) when
possible. The authors shine here and there with such descriptions as “‘shallow soil
over rocks in open grassland”; I then know that they know about this specific plant.
Synonyms are given for many taxa, although I would like to see more in a scientific
publication such as this. A few typographical errors occur here and there besides the
included errata sheet.

When I read about special taxa that literally shoot up out of the pages as disjunct
species, etc., without an explanation, I am left with a void. Nothing is given about
Asplenium vespertinum Maxon, Lewisia rediviva Pursh var. rediviva, Silene verecunda
S. Watson subsp. platyota (S. Watson) Hitch. and Maguire, Myrica californica Cham.,
Perityle emoryi Torr., Adenostoma sparsifolium Torr., Batis maritima L., etc. But I
can see why. Most of these field notes are in McAuley (1985).

Another minor criticism but important to me is the use of “‘extinct” and ‘“‘extir-
pated”’ without historical data. Under Astragalus pycnostachys A. Gray var. la-
nosissimus (Rydb.) Munz and McBurne ex Munz (a plant that I have heard and
wondered about for years along the seaward side of the Santa Monica Mountains), I
read “Very rare if not already extirpated from our area; not actually known in our
area but may occur at Mugu Lagoon.” This gives me mixed reactions. Why not
shorten this to the facts by merely citing the last collection(s), e.g., a hypothetical
case: ““Last collected by Parish at Pt. Mugu in 1901,” and not seen since. Then one
can speculate where one might look for it. An interesting entry is Myrica californica
Cham. and Schlecht. “Presumably in tangled streamside vegetation; Santa Monica
Canyon.”’ This same explanation is in the 1966 flora. It seems like someone would
have checked this out in 20 years. McAuley (1985) states it is ““Found in canyons
and moist slopes at low elevations. Uncommon.”

Now, to bring this flora into perspective. The Santa Monica Mountains have been
of much interest to me because I became involved in a flora to the immediate north
and was interested in the relation of the plants of the Santa Monica Mountains to
the northern Channel Islands, because they are visually an extension of this range. I
was first introduced to its flora by correspondence with Henry L. Bauer of Santa
Monica Junior College, who was also interested in these mountains and published
(mimeographed) in 1952 a “Check List and Distribution of the Plant Species Growing
Naturally in the Santa Monica Mountains, California’’, by O. H. Kappler, consisting
of 13 pages. In 1983, several popular checklists on the flora of the Santa Monica
Mountains were published by Bob Muns, and in 1985 the Wildflowers of the Santa
Monica Mountains by Milt McAuley appeared with 544 pages and 496 colored plates,
followed in 1986 by Flowering Plants: the Santa Monica Mountains by Nancy Dale,
with 239 pages and 214 colored plates. These two beautiful publications are irrevo-
cably woven into this article.

1p MADRONO [Vol. 34

I have the impression that cooperation is uncoordinated among the leading floristic
authors, plant collectors, and the professional botanists. Barry Prigge states that for
new species to be incorporated in future revisions, ‘““Voucher specimens have to be
made with collection data and deposited at a local herbarium.”’ Several McAuley
species are not listed as he (Prigge) has not seen them. Bob Muns in an October 1983
checklist states “I have found 20 more species growing in the Santa Monica Moun-
tains.”’ We in Santa Barbara also are guilty and can add two more species, Lavatera
arborea L. and Nicotiana clevelandii Gray. Cordylanthus maritimus Nutt. ex Benth.
subsp. maritimus has been surveyed at the Mugu Lagoon by Julie M. Vanderwier
and Judith C. Newman (Madrono 31:185—186. 1984), and others.

This flora is the only publication for these mountains with dichotomous keys and
should remain the “‘scientific master list’’. It can only grow in scope with the help of
many who have diverse interests. —CLIFTON F. SMITH, Santa Barbara Botanic Garden,
1212 Mission Canyon Rd., Santa Barbara, CA 93105.

Marcus E. Jones: Pioneer Western Geologist, Mining Engineer & Botanist. By LEE
W. LENz. xv + 486 pp., 9 pp. black and white photographs. Rancho Santa Ana
Botanic Garden, Claremont, CA. 1986. ISBN 0-9605808-2-4, $28.00 plus tax and
$1.50 shipping.

If Lee Lenz had done no more than compile and make generally available the
excellent gazetteer of his subject’s collecting localities and chronology for nearly a
half century of field work, the botanical community would be greatly in his debt. For
anyone who, like this reviewer, tends to identify Marcus E. Jones, A.M., solely with
the Great Basin, the map on page 292 is a revelation, showing that he was active in
every western state but Alaska and Hawaii, although his coverage was considerably
more modest than his claims. His unconsummated “Flora of the Great Plateau’”’ was
to have embraced “‘the region west of the Plains” and extend well into both Canada
and Mexico, in short, the American West.

But the author also has assumed the task of presenting a biography of Jones. This
is a difficult assignment because the man and the scientist have become almost
obscured by his reputation as a choleric, fiercely independent, feisty, rough-and-
tumble combatant in a notably polemic era. Lenz allows Jones to speak for himself,
but this does little to dispel the reputation. Jones was a born-again Protestant and
frequently a lay preacher, but he was notably lacking in the Christian virtues of
humility and charity. His strong religious commitment, however, did not pose an
obstacle to his full acceptance of evolution, as it did for many of his contemporaries.
The panoply of Nature was God’s handiwork.

Born in Ohio, educated to a Master’s degree at Iowa College (predecessor of Grin-
nell), he was prepared to teach Classics or, as was the custom in the West of his day,
any other subject for which there was an opportunity. His precarious health drew
him to outdoor life and his inclination to a career as a field naturalist. He did not
succeed in establishing a firm institutional connection until late in life. He and his
family, centered after 1880 in Salt Lake City, endured a grinding hand-to-mouth
existence based on his irregular teaching assignments, preaching, free-lance writing
on a variety of issues, and, increasingly, geological consulting. Mining investments
seldom proved profitable and his performance as an expert witness for the prosecution
in smelter-pollution law suits, while admirable, was not financially rewarding. His
long-suffering wife augmented or supplied the family income with kindergarten teach-
ing and operation of a rooming and boarding house.

Jones’ most productive period botanically was undoubtedly from about 1880 to
1912. Every summer was devoted to collecting somewhere in the West, the winters
to identifying, labeling, and distributing the resulting specimens for sale, and working
on his “‘Flora.”’ By 1885, access to the few existing botanical journals had been cut
off, as it was to anyone who refused to have his proposals approved by members of
the Eastern establishment. Launching of the journal Zoé by the Brandegees’ provided

1987] REVIEWS 73

the opportunity to initiate his extensive series of ‘““Contributions to Western Botany”
in 1891, but a few years later he was doing his own publishing. The “‘Flora” was
planned to be the first fully illustrated botany of the area. The manuscript for volume
one was essentially completed by 1907, but the second remained incomplete, and
there is no indication that he worked on it after he left Utah for California in 1923.
Neither volume was ever published, and a “Trees and Shrubs of Utah” and a sub-
stantial ‘“‘Flora of Flathead Lake” also remained in typescript.

Because Jones published relatively little of substance besides new species and trained
no disciples, the book inevitably becomes in large part the history of his herbarium.
By sale of collections, diversification of teaching stints, and consulting trips, he was
able to cover an astonishing amount of territory, and amass a steadily growing private
herbarium, which was clearly his first priority. From the early seventies until his
death in 1934, he was continuously and vigorously adding specimens. Much of his
concern in later years was finding a suitable home for it and, for the few active years
remaining, for himself. In this he fortunately was successful. After brief flirtations
with the California Academy of Sciences and the University of Utah, the herbarium
was obtained for Pomona College by Phil Munz. Although terms of the sale provided
for Jones to publish his “‘Flora’’, and he was now freed of financial worries for the
first time in his life, much of the original area of his interest had long been preempted
by other authors. He turned instead to field work in northern Mexico— further adding
to his herbarium.

The author attempts a “‘summing up” of Jones’ contributions, which particularly
pairs him with Greene. Certainly the two were fellow mavericks and adepts at in-
venting colorful invective and generating hostility, but I think a closer comparison
might have been made between Jones and Katharine Brandegee, whom he so greatly
admired. Both had considerable gifts of critical analysis, but their special insights
largely died with them.

The volume includes a list of published writings, diary and field notes, 1894 (Ap-
pendix I), the 112-page annotated gazetteer (Appendix II), and a list, with explanatory
notes, of the nearly 800 new taxa that Jones proposed during his lifetime (Appendix
Ill).

Incorporation of the Jones Herbarium into the Pomona-Claremont-Rancho Santa
Ana research complex at last provides its creator the central role in West American
botany that he never quite succeeded in attaining in life. — LINCOLN CONSTANCE, Dept.
Botany, Univ. California, Berkeley 94720.

Botanical Illustration: Preparation for Publication. By No—EL H. HOLMGREN and BOBBIE
ANGELL. 74 pp. The New York Botanical Garden, Bronx, NY 10458. 1986. ISBN
0-89327-272-8, $12.

A quote the authors use in the beginning of this book could just as well apply to
the authors themselves: ‘“‘Well-ordered presentation is the sign of a well-ordered
mind” (Alfred A. Blaker, 1977). This book is small, but it covers concisely and
thoroughly all aspects necessary to produce a botanical illustration that is correct and
aesthetically pleasing. I have done quite a bit of botanical illustrating, yet I was able
to learn from this book. Holmgren and Angell not only cover basic illustrating guide-
lines, they also offer shortcuts and technical tips that can save both the illustrator
and the author valuable time, money and energy.

Much has been written about scientific illustration (some of the better publications
are included in a helpful Annotated Bibliography at the end of this book), but very
little good information exists on the very special needs encountered in botanical
drawings. From the first chapter, ““Working Relationships’’, through others that in-
clude some useful information on tools, sizes and proportions of plates (including
instruction on reductions), labels, captions, illustrations, maps and graphs, photo-
graphs and halftones, to preparing art for shipping, this volume contains a wealth of
information in a very succinct and informative way. There is a good section on the

74 MADRONO [Vol. 34

U.S. Copyright law that was changed in 1978. An understanding of this law is im-
portant, especially if either the artist or the author have any interest in maintaining
rights to the artwork.

The illustrations used throughout the book are clear and nicely done. I thought the
samples used in the chapter on ‘Plant Illustrations” were especially well-rendered.
It’s nice to see that the authors are quality artists as well as good writers. Their writing
style 1s straight-forward and very clear; they stay away from verbal embellishments
that might only tend to confuse. The Table of Contents is well done, making it a
simple matter to look up particular information (chapters are indicated in bold type,
whereas sub-headings with page numbers show specific areas contained in those
chapters).

In short, this book would be a welcome addition to any botanist’s or artist’s library.
Botanists who do their own illustrating will find this book especially helpful. — MAGGIE
Day, Dept. Biological Sciences, Univ. California, Santa Barbara 93106.

ANNOUNCEMENT
NEW PUBLICATIONS

VAN BRUGGEN, T., The vascular plants of South Dakota, 2nd ed., Iowa
State Univ. Press, 2121 S. State Ave., Ames 50010, 1985, xxv, 476
pp., illus., ISBN 0-8138-0650-X, $28.95 (paperbound). [First edition
1976; treatment of 1608 native and adventive species found in South
Dakota and adjacent areas. ]

SCAGEL, R. F., D. J. GARBARY, L. GOLDEN, and M. W. HAwkKEs, 4
synopsis of the benthic marine algae of British Columbia, northern
Washington and southeast Alaska, Dept. of Botany, Univ. British
Columbia, Vancouver V6T 2B1, 1986, vi, 444 pp., ISSN 0831-4861,
ISBN 0-88865-460-X, Can $15.00 (paperbound).

STUBBENDIECK, J., S. L. HATCH, and K. J. Hirscu, North American range
plants, 3rd ed., Univ. Nebraska Press, 901 North 17th St., Lincoln
68588, 1986, xv, 465 pp., illus., ISBN 0-8032-9 162-0, $18.95 (paper-
bound).

1987] ANNOUNCEMENTS Ti)

ANNOUNCEMENT
NEW PUBLICATIONS

Cooke, W. B., The fungi of our mouldy earth, Beihefte zur Nova Hed-
wigia, heft 85. J. Cramer, Berlin, West Germany, 1986, vi, 468 pp.,
114 figs., 2 pl. [This volume includes chapters on the collection,
preparation, isolation, and identification of fungi, in addition to those
on habitats, classification, and a systematic list of the species.]

WELSH, S. L., N. D. ATwoop, L. C. Hicains, and S. Goopricu, A Utah
flora, Great Basin Naturalist Memoir 9, Brigham Young Univ., 290
M. L. Bean Life Science Museum, Provo, UT 84602, 1986. [4 Utah
flora is a comprehensive treatment of the vascular flora of the state,
including 2572 native species, 355 intraspecific entities, and 580 in-
troduced species. Taxa are described, ecological data are given, and
geographic information is provided. ]

WIARD, L. A., An introduction to the orchids of Mexico, Comstock Publ.
Assoc., Cornell Univ. Press, Ithaca, NY 14850, 1986, 216 pp., 80
pls. (color), ISBN 0-8014-1833-X, $75.00 (hardbound). [A sampling,
in 21.6 x 28 cm format, of 154 of the over 800 native orchid species
of Mexico; all 154 species are illustrated by Wiard, mostly from plants
in the wild or from collections of Wiard and friends. The main part
of the book describes the 60 genera treated, with species descriptions
giving habitat and other details.]

ANNOUNCEMENT
FIFTH WILDLAND SHRUB SYMPOSIUM

The Shrub Research Consortium is sponsoring the Fifth Wildland
Shrub Symposium 30 June—2 July 1987 at Utah State University, Logan,
Utah. The symposium, “‘Shrub Ecophysiology and Biotechnology’’, will
feature invited and contributed papers. Contributed presentations will
be 20 minutes. The proceedings will be published by the USDA Forest
Service Intermountain Research Station. If you would like to present a
paper, send a title and abstract by 31, March 1987, to: Dr. Arthur
Wallace, Laboratory of Biomedical and Environmental Sciences, UCLA,
900 Veteran Avenue, Los Angeles, CA 90024.

To receive preregistration materials and information please contact:
Michael B. Price, Eccles Conference Center, Room 103 F, Logan, UT
84322-5005; phone: (801) 750-1696.

ANNOUNCEMENT
THIRD CALIFORNIA ISLANDS SYMPOSIUM

Hosted by
Santa Barbara Botanic Garden (SBBG), Santa Barbara Museum of Nat-
ural History (SBM), and the Southern California Academy of Sciences.

This symposium took place in Santa Barbara on 2-6 March 1987.
Contributed papers covered the following topics: history and resource
management; birds; fishes and marine botany; anthropology; ocean-
ography; terrestrial and marine invertebrates; terrestrial botany; marine
and terrestrial vertebrates; and geology and geography. A collected set
of full length manuscripts and extended abstracts of papers presented
at the symposium will be published in book form. For information
concerning the event contact Dr. F. G. Hochberg, Santa Barbara Mu-
seum of Natural History, 2559 Puesta del Sol Rd., Santa Barbara CA
93105.

ANNOUNCEMENT
ELEVENTH GRADUATE STUDENT MEETINGS

The California Botanical Society will sponsor the Eleventh Graduate
Student Meetings on 25, 26 April 1987 at the University of California,
Davis.

The presentation categories (proposed research, research in progress,
and finished research) allow for the sharing of ideas and knowledge
among the graduate student community. Awards for each of these cat-
egories will be presented at the banquet on 26 April.

For information contact Niall F. McCarten, Graduate Student Rep-
resentative, Dept. of Biology, San Francisco State Univ., 1600 Hollo-
way, San Francisco, CA 94132.

Volume 34, Number 1, pages 1-76, published 31 March 1987

SUBSCRIPTIONS — MEMBERSHIP

Membership in the California Botanical Society is open to individuals ($18 per year;
students $10 per year for a maximum of seven years). Members of the Society receive
MaproNo free. Family memberships ($20) include one ten-page publishing allot-
ment and one journal. Emeritus rates are available from the Corresponding Secretary.
Institutional subscriptions to MADRONO are available ($25). Membership 1s based on
a calendar year only. Applications for membership (including dues), orders for sub-
scriptions, and renewal payments should be sent to the Treasurer. Requests and rates
for back issues, changes of address, and undelivered copies of MADRONO should be
sent to the Corresponding Secretary.

INFORMATION FOR CONTRIBUTORS

Manuscripts submitted for publication in MADRONO should be sent to the editor.
All authors must be members, and membership is prerequisite for review.

Manuscripts and review copies of illustrations must be submitted in triplicate for
all articles and short items intended for NOTES AND NEWS. Follow the format
used in recent issues for the type of item submitted and allow ample margins all
around. All manuscripts MUST BE DOUBLE SPACED THROUGHOUT. For ar-
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CALIFORNIA BOTANICAL SOCIETY

STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION
(Required by Title 39, U.S.C. 3685)

MaproNo, A West American Journal of Botany, is published quarterly at Berkeley,
California. Annual subscription price is $25.00.

The Publisher is the California Botanical Society, Inc., Life Sciences Building,
University of California, Berkeley, CA 94720.

The editor is Wayne R. Ferren, Jr., Department of Biological Sciences, University
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The owner is the California Botanical Society, Inc., Life Sciences Building, Uni-
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I certify that the statements made by me above are correct and complete.

January 26, 1987 WAYNE R. FERREN, JR., Editor

“VOLUME 34, NUMBER 2 ; i APRIL-JUNE 1987

i vA

MADRONO

i WEST AMERICAN JOURNAL OF BOTANY

Contents

| GERMINATION AND ESTABLISHMENT OF Pinus contorta VAR. murrayana NaN aad in
| MOUNTAIN MEADOWS OF YOSEMITE NATIONAL PARK, Moe ;
John A. Helms and Raymond D. Ratliff ? ‘ae Ti.

INVASION OF Pinus contorta VAR. murrayana (PINACEAE) INTO: Monaro iemowt
AT YOSEMITE NATIONAL PARK, CALIFORNIA

Se

John A. Helms 91
_ THE DISTRIBUTION OF FOREST TREES IN NORTHERN BAJA CALIFORNIA, MEXICO
Richard A. Minnich 98

|
Fire History OF AN OLD-GROWTH ForEST OF Sequoia sempervirens (TAXODIACEAE)
|

FoREST IN HUMBOLDT REDWOODS STATE PARK, CALIFORNIA

John D. Stuart 128
| CHROMOSOME RACES OF Grayia brandegei (CHENOPODIACEAE)
_ Howard C. Stutz, Stewart C. Sanderson, E. Durant McArthur, and Chu Ge-Lin 142
| Allium shevockii (ALLIACEAE), A NEw SPECIES FROM THE CREST OF THE SOUTHERN

SIERRA NEVADA, CALIFORNIA
_ Dale W. McNeal 150
Claytonia palustris (PORTULACACEAE), A NEw SPECIES FROM CALIFORNIA
| John R. Swanson and Walter A. Kelley Ss
| Alloispermum insuetum (ASTERACEAE: HELIANTHEAE), A NEW SPECIES FROM COLOMBIA

Carmen F. Fernandez, Lowell E. Urbatsch, and Gene Sullivan 162
| A New SPECIES OF Axiniphyllum (ASTERACEAE: HELIANTHEAE) FROM DURANGO,

MEXICO

B. L. Turner 165
/ NOTES
| RANGE EXTENSION, CHROMOSOME COUNT, AND MEPHITISM IN Lessingia tenuis
_ (COMPOSITAE)
John Mooring 168
‘NOTEWORTHY COLLECTIONS

ARIZONA 170

CALIFORNIA 170

COLORADO 171
| New Mexico Leal
) REVIEWS 169, 172

ANNOUNCEMENTS 90, 141, 149, 154, 164, 167

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MaAprRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
ciety, Inc., and is issued from the office of the Society, Herbarium, Life Sciences
Building, University of California, Berkeley, CA 94720. Subscription rate: $25 per
calendar year. Subscription information on inside back cover. Established 1916.
Second-class postage paid at Berkeley, CA, and additional mailing offices. Return
requested. POSTMASTER: Send address changes to James R. Shevock, Botany Dept.,
California Academy of Sciences, San Francisco, CA 94118.

Editor—WAyNE R. FERREN, JR.
Associate Editor—BARRY D. TANOWITZ
Department of Biological Sciences

University of California
Santa Barbara, CA 93106

Board of Editors
Class of:

1987—J. RZEDoOwsKI, Instituto de Ecologia, A.C., Mexico
DoroTHy DouGLas, Boise State University, Boise, ID
1988—SusANn G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989—FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991—Davip J. Ket, California Polytechnic State University, San Luis Obispo
JAMES HENRICKSON, California State University, Los Angeles

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1986-87

President: FRANK ALMEDA, Department of Botany, California Academy of Sciences,
San Francisco, CA 94118

First Vice President: PATRICK E. ELVANDER, 273 Applied Sciences, University of
California, Santa Cruz, CA 95064

Second Vice President: KINGSLEY R. STERN, Department of Biology, California State
University, Chico, CA 95929

Recording Secretary: V.THOMAS PARKER, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
Francisco, 50 Phelan Ave., San Francisco, CA 94112

The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, CHARLES F. QuUIBELL, Department of Biology,
Sonoma State University, Rohnert Park, CA 94928; the Editor of MADRONO; three
elected Council Members: THOMAS FULLER, 171 Westcott Way, Sacramento, CA
95864; ANNETTA CARTER, Department of Botany, University of California, Berkeley,
CA 94720; JOHN Moorina, Department of Biology, University of Santa Clara, Santa
Clara, CA 95053; and a Graduate Student Representative, NIALL F. MCCARTEN,
Department of Biological Sciences, San Francisco State University, San Francisco,
CA 94132.

GERMINATION AND ESTABLISHMENT OF
PINUS CONTORTA VAR. MURRAYANA (PINACEAE)
IN MOUNTAIN MEADOWS OF YOSEMITE
NATIONAL PARK, CALIFORNIA

JOHN A. HELMS
Department of Forestry and Resource Management,
University of California, Berkeley 94720

RAYMOND D. RATLIFF
USDA Forest Service, Pacific Southwest Forest and
Range Experiment Station,
2081 E. Sierra Ave., Fresno, California 93710

ABSTRACT

Pinus contorta var. murrayana (lodgepole pine) commonly invades mountain
meadows. Studies were made on a wet and a dry meadow in Yosemite National Park,
California, to determine differences in meadow habitat as defined by differences in
composition and cover of herbaceous vegetation, soil characteristics, and physiog-
raphy. Site ordination identified species on sites of decreasing moisture content. In
both fall and spring, P. contorta seeds were sown on both covered and uncovered
plots in each of 12 meadow sites. Germination and survival of seeds were measured
for three consecutive years. Invasion of pine was uncommon on both the wetter
meadow, dominated by Aster alpigenus, Carex nebraskensis, and Deschampsia caes-
pitosa, and on the driest sites characterized by Lupinus confertus and Horkelia fusca
subsp. capitata. Most invasion occurred on moderately dry sites dominated by Aster
occidentalis, Trifolium longipes, and Danthonia californica. Differences in germina-
tion from fall and spring seeding were not significant. Depredation by rodents and
birds reduced germination and survival on the uncovered seed plots by approximately
50%.

Pinus contorta Dougl. ex Loud. var. murrayana (Grev. & Balf.)
Engelm. (lodgepole pine) commonly invades mountain meadows in
the Sierra Nevada of California. Invasion is sporadic and the factors
that limit germination and establishment of pine seedlings in mead-
Ows are not well understood. The invasion is important in meadow
succession and is of concern to resource managers who are respon-
sible for meadow conservation and management.

Mountain meadows differ substantially in topography, water
availability, and microclimate. These differences occur as within-
and between-meadow variation in timing of snow melt, drainage
patterns, and vegetative cover. These factors probably also influence
the timing and extent of lodgepole pine seedling establishment in
meadows.

Germination of lodgepole pine is abundant in full sunlight, on
bare mineral soil or disturbed duff, in the absence of competing

MADRONO, Vol. 34, No. 2, pp. 77-90, 1987

78 MADRONO [Vol. 34

vegetation, and with adequate soil moisture (Lotan 1964, Shepperd
and Noble 1976, Lotan and Critchfield in press). Seedling mortality
is associated commonly with high soil surface temperature, drought,
soils with low water-holding capacity, unincorporated organic mat-
ter, and grazing (Cochran 1969, Lotan and Perry 1977, Lotan and
Critchfield in press). Specific sites within meadows that are more
favorable for pine establishment are indicated by the presence of
‘outlier’ trees. These trees are associated commonly with exposed
rocks, logs, and groups of shrubs (Leonard et al. 1968, 1969) that
are thought to provide higher soil temperature, more favorable soil
texture and drainage, earlier snow melt, and protection from brows-
ing.

This study was designed to contribute to the knowledge of lodge-
pole pine germination and to examine more closely within- and be-
tween-meadow variability in establishment of pine seedlings. Spe-
cific objectives were to determine the extent to which successful
establishment is associated with: 1) availability of seeds and possible
losses over the winter, 2) vegetative cover and soil water content,
and 3) depredation by rodents and birds.

STUDY AREA

Two meadows that had not been grazed recently by range cattle,
one large and wet and the other small and dry, were studied in
California’s Yosemite National Park. These meadows are located at
2100 m near Glacier Point Road on the trail to Lost Bear Meadow
(Fig. 1). Both meadows are surrounded by lodgepole pine stands,
have vegetated rather than sandy margins, and are montane rather
than subalpine. The larger site (3.37 ha) has topography of type A,
formed in a basin; the smaller site (1.83 ha) is type C, formed along
a permanent stream (Ratliff 1985). The larger meadow ranges from
good to excellent condition, i.e., having no abnormal erosion and
with herbage production near the climatic potential. The smaller
meadow ranges from good to very poor condition. The poorest
conditions occur near the stream channel, which is a continuous
gully 1-2 m deep where erosion has lowered the water table.

Several study sites (areas that differ in species composition; Ratliff
1982) occur in each meadow. They represent different meadow series
as defined by their hydrologic and vegetative classifications (Ratliff
1985).

METHODS

Study sites. In late summer of 1981, five sites (sites 1-5) on the
large meadow and four (sites 6—9) on the small meadow were selected
to represent different microenvironments that could influence the

1987] HELMS & RATLIFF: PINUS CONTORTA VAR. MURRAYANA. I. 79

Yosemite
Hwy Village
140 YOSEMITE |
4, — Glacier
Point
VALLEY
Hwy
120
1km
37°40
Badger vif iq Study
Pass Bridalveil Site
Campground °.
240’ ‘ Lost Bear
Uiehe Meadow

Fic. 1. Location of study.

success of lodgepole pine seedling establishment. Size and shape of
sites varied with vegetation boundaries, but contained no less than
100 m?. Because of small floristic differences, sites 2, 6, and 9 were
subdivided for vegetation analysis. Rather than measure topography
and microclimate directly, we evaluated site differences using flo-
ristic characteristics. This approach presumes that frequency and
cover of particular herbaceous species are related to differences among
microenvironments within the meadows, and that different species
and their relative cover influence the capacity of lodgepole pine to
invade meadows.

On each site, species frequencies were estimated using 100 ran-
domly located 10 <x 10 cm quadrats. Foliar cover (proportion of
area under live aerial parts) and species composition were estimated
at each seed plot. Site and species ordinations were derived from
these data by reciprocal averaging (Hill 1973).

Soil characteristics near the center of each site were compared by
taking a soil sample from the 10—20 cm depth. Samples were ana-
lyzed for: 1) percent sand, silt, and clay (estimated by the hydrometer
method; Bouyoucos 1936); 2) percent organic matter [estimated by
the loss on ignition method (ignition was for 4 hr at 600°C), Nelson
and Sommers 1982]; 3) pH (estimated by the soil-water paste meth-
od with ionanalyzer; McLean 1982); and 4) bulk density at 0-10 cm
and 10-20 cm depths [estimated by the cylinder (known volume)
method, Cook and Stubbendieck 1986].

Soil depth to consolidated material was estimated using a probe,
and depth to the water table was determined prior to fall precipi-
tation in October 1983.

Seed plots. Ten replications of both uncovered and covered 0.1

80 MADRONO [Vol. 34

m? plots containing approximately 50 seeds were established on each
of the 12 sites in the fall of 1981 and again in the spring of 1982.
Seeds were obtained from cones collected in 1981 from trees in an
adjacent area. Seeds sown in the fall were unstratified. Spring sowing
after snow melt utilized stratified seeds (soaked one day, held at
1.5°C in plastic bags for 28 days, sown wet; USDA 1974) to simulate
the treatment that seeds receive as they overwinter on the meadows.
Covers were made from 1.3-cm-mesh wire screen.

The number of seedlings in each plot was counted four times
during the growing season in 1982, and also at the end of summer
in both 1983 and 1984. Tests for significance of differences between
mean survival at the end of three growing seasons were made using
the Kruskal-Wallis non-parametric test for differences in means of
ranked data grouped by single classification (Conover 1980). All
plant nomenclature follows Munz (1959).

RESULTS
Vegetation

Twenty-three herbaceous species were found on both meadows;
an additional 27 species occurred only on the large meadow and an
additional 20 occurred only on the small meadow. Species with
frequencies of 20% or more on at least two sites were used for
vegetation analyses. The single axis reciprocal averaging ordinations
of species frequency and composition data described similar arrays
of herbaceous species on sites of decreasing moisture content. Con-
sequently, species composition of the seed plots adequately defined
the microenvironments of each site (Table 1).

Species most representative of wetter conditions (sites 4, 3, 5)
were Aster alpigenus, Carex nebraskensis, C. rostrata, Deschampsia
caespitosa, Juncus orthophyllus, and Phalacroseris bolanderi (Table
1). The moderately dry sites (sites 2, 6, 7, 1) were characterized by
abundant Aster occidentalis, Danthonia californica, and Trifolium
longipes. Species most representative of dry conditions (sites 8, 9)
were Horkelia fusca subsp. capitata, Lupinus confertus, and Penste-
mon rydbergii.

Soils

Ordination of the sites from wettest to driest generally reflected
increasing depths to the water table (Table 2). This shows a close
relationship between species frequency, composition, and soil water
availability. Also, with the gradient from wet to dry sites, organic
matter decreased, and both bulk density (OQ—10 cm depth) and acidity
increased. Soil textures and depths showed no apparent relationship
to the moisture gradient.

HELMS & RATLIFF: PINUS CONTORTA VAR. MURRAYANA. I. 81

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1987] HELMS & RATLIFF: PINUS CONTORTA VAR. MURRAYANA. I. 83

TABLE 3. THE NUMBER OF SURVIVING LODGEPOLE PINE SEEDLINGS PER SEED PLOT
AT THE END OF THE THIRD GROWING SEASON AFTER SOWING 50 SOUND SEED PER
PLot. Site ordination corresponds to a decreasing soil moisture gradient (left to right)
shown in Table 1.

Site
4 3 5 2. 6 1 1 8 9 Mean
Fall-sown: covered 0 O 04 5.6 3.1 29.5 0.0 10.7 O.1 5.48
uncovered 0 O O 0.1 0.8 7.3 0.0 3.5 O 1.30
Spring-sown: covered 0 0 O 3.8 1.4 19.6 7.8 8.7 0.3 4.62
uncovered 0 0 O OF) Mey Vid C2 3.5 O 213
Seed Plots

Fall seeding: covered seeds. On the large meadow, survival during
the first year after sowing on sites 2, 4, and 5 was 20-30% and was
substantially greater than the 2—4% survival on sites 1 and 3 (Fig.
2a). In the second year, survival on all sites except site 2 decreased
rapidly. By year 3, survival on sites 1, 3, and 4 was zero (Table 3).
On site 5, survival of 0.8% was lower (p = 0.10) than the 11.2% on
site 2.

On the small meadow, seedling survival at the end of the first year
was generally much higher (72.4, 48.8, 37.4, and 0%, respectively
on sites 7, 8, 6, and 9). In subsequent years, survival decreased until
by year three it was 59.0, 21.4, 6.2, and 0.2%, respectively. Mean
survival on site 7 was significantly greater (p = 0.05) than on sites
8 and 6, and survival on site 8 was greater (p = 0.10) than on site 6.

Fall seeding: uncovered seeds. On the large meadow at the end of
the first year, survival was 28.0, 15.6, 4.0, 3.6, and 0%, respectively
on sites 4, 5, 3, 2, and 1 (Fig. 3a). By the end of year two, survival
on all sites had become close to zero.

On the small meadow, survival at the end of the first year was
28.0, 18.6, 7.8, and 0%, respectively on sites 6, 7, 8, and 9 (Fig. 3b).
Mean survival on site 7 was greater (p = 0.10) than that on site 8;
however, the means of survival on sites 6 and 7 were not different
at that same level of significance. By the end of year three, survival
was 14.6, 7.0, 1.6, and 0%, respectively on sites 7, 8, 6, and 9. This
ranking of sites was the same as that on the covered seed plots.

Spring seeding: covered seeds. On the large meadow, only sites 1
and 2 were sown because sites 3, 4, and 5 were under water until
late summer. First-year survival on these two sites was 6.8 and
11.8%, respectively (Fig. 4). Second-year survival on site | increased
to 28.1%, indicating additional germination of seeds on this driest
site in the second year after sowing. At the end of year three, the

84

Survival (%)

MADRONO [Vol. 34

FALL SEEDING -- covERED SEED PLOTS

a. Large Meadow

40

| WA

10

b. Small Meadow

_—
8

Ss
9

JJA O A

A
1982 1983 1984
Calendar Date

80

70

60

$0

40

30

20

10

1987] HELMS & RATLIFF: PINUS CONTORTA VAR. MURRAYANA. I. 85

mean survival on plots 1 and 2 was 15.6 and 7.6%, respectively,
and these means were not different (p = 0.10).

On the small meadow, survival at the end of the first year was
47.4, 45.2, 25.4, and 0% on sites 8, 6, 7, and 9, respectively (Fig.
4b). Additional germination of seeds in the second year occurred on
the driest sites 7 and 9. By the end of year three, survival was 38.2,
17.8, 2.8, and 0.7%, respectively in sites 7, 8, 6, and 9. Using p =
0.10 as a test of significance, the means of survival on sites 6 and
8, and on sites 7 and 8, were different, but means of survival on
sites 6 and 9 were not.

Spring seeding: uncovered seeds. On the large meadow, the extent
and pattern of survival were similar to those in the covered spots
(Fig. 5a), but survival at the end of the third year was only 4.2% on
site 1 and 1.4% on site 2, respectively; the difference between these
means was not significant (p = 0.10).

On the small meadow, survival at the end of the first year on sites
6, 8, 7, and 9 was 18.0, 16.4, 14.0, and 0%, respectively (Fig. 5b).
Sites 7 and 9 showed additional germination during the second year
after sowing. By the end of year three, survival was 22.4, 7.0, 3.4,
and 0% for plots 7, 8, 6, and 9, respectively. The difference between
mean survival on sites 7 and 8 was significant (p = 0.05), whereas
that between sites 6 and 8 was not.

DISCUSSION

Seed availability. Direct seeding of lodgepole pine showed that
pine establishment in meadows 1s associated less with seed avail-
ability than with meadow type. On the wetter meadow, applying
seeds in the fall to simulate natural seed fall showed that, despite
abundant seeds, no seedlings survived through the second year (Fig.
3). On the drier meadow, up to 14% survival by the end of the third
year indicates that invasion of this meadow type is much more likely
(Fig. 3). Generally, for each meadow site, differences in survival
between seeding in the fall and spring (Figs. 2 and 4; Figs. 3 and 5)
were non-significant. This suggests that overwinter decreases in ger-
minability and physical movement of seed off the site by melting
snow are not important factors in the establishment of lodgepole
pine.

Depredation. The similarity of pine seedling survival in covered

_

Fic. 2. Survival of lodgepole pine seedlings on fall-seeded, covered plots on sites
1-9 for the large and small meadows. Bars indicate standard errors of means; letters
indicate months.

86 MADRONO [Vol. 34

FALL SEEDING -- UNCOVERED SEED PLOTS

a. Large Meadow

40
30
; Q 4
20
vas 4 5
10
At > ae
: "4 2 — a

b. Small Meadow

Survival (%)

JJsa 0 A A
1982 1983 1984

Calendar Date

Fic. 3. Survival of lodgepole pine seedlings on fall-seeded, uncovered plots on
sites 1-9 on the large and small meadows. Bars indicate standard errors of means;
letters indicate months.

and uncovered fall-seeded plots on the large meadow (except for the
drier site 2) (Figs. 2 and 3) suggests that depredation by rodents and
birds on wet sites is unimportant. Covered, spring-seeded plots on
the large meadow, however, had approximately twice the survival
of the corresponding uncovered plots at each measurement date
(Figs. 4 and 5).

Depredation was most apparent on the drier small meadow where
germination and survival on uncovered fall-seeded plots were re-
duced by approximately 50% (Figs. 2 and 3). The effect of covering
seeds and seedlings was less apparent on spring-seeded plots, with

1987] HELMS & RATLIFF: PINUS CONTORTA VAR. MURRAYANA. 1. 87

SPRING SEEDING --CoveRED SEED PLOTS

a. Large Meadow
40

30

20

10

b. Small Meadow

Survival (%)

60

40

30

20 a

10 =

JJa O A A
1982 1983 1984

Calendar Date
Fic. 4. Survival of lodgepole pine seedlings on spring-seeded, covered plots on

sites 1-9 on the large and small meadows. Bars indicate standard errors of means;
letters indicate months.

88 MADRONO [Vol. 34

SPRING SEEDING -- UNCOVERED SEED PLOTS

a. Large Meadow

b. Small Meadow

40

Survival (%)

30

20

10

JJIA 0 A A
1982 1983 1984

Calendar Date
Fic. 5. Survival of lodgepole pine seedlings on spring-seeded, uncovered plots on

sites 1-9 on the large and small meadows. Bars indicate standard errors of means;
letters indicate months.

the notable exceptions of site 7 where seeds were sown on soil bare
from gopher activity, and of site 8 that had only 30% foliar cover
(Figs. 4 and 5). The greater survival of seedlings on the small meadow
corresponds with its having a much larger number of previously-
established outlier lodgepole pine trees.

Increased number of seedlings in the second year on the drier
spring-seeded plots, 1, 7, and 9, may be associated with possible
delayed germination of some seeds. This increase of approximately
8-25 seedlings per 0.1 m? plot probably is too large to be explained
entirely by additional naturally-dispersed seeds. No new seedlings
were observed adjacent to the plots.

1987] HELMS & RATLIFF: PINUS CONTORTA VAR. MURRAYANA.I. 89

Meadow type. Differences in frequency and cover of herbaceous
species across meadows reflect differences in moisture regimes and
potential for pine invasion. High potential occurs on moderately dry
sites. These sites are usually too wet early in the year for much seed
and seedling depredation. Later, these sites are not saturated but
remain moist, which improves chances for establishment. Low po-
tential occurs in the wettest and driest sites where either continuous
saturation or moisture stress tends to prevent establishment.

ACKNOWLEDGMENTS

This study was supported by the USDA Forest Service, Pacific Southwest Forest
and Range Experiment Station, Berkeley, California, Project Number PSW-80-0022.
It also was part of the University of California’s Agricultural Experiment Station
Project 2942-MS. We thank Drs. W. Critchfield and F. Vasek for their constructive
reviews and suggestions for improving the manuscript.

LITERATURE CITED

Bouyoucos, C. J. 1936. Directions for making mechanical analysis of soils by the
hydrometer method. Soil Science 42:225-228.

COCHRAN, P. H. 1969. Thermal properties and surface temperatures of seedbed: a
guide for foresters. U.S.D.A. Forest Serv. Pacific Northw. Forest and Range Exp.
Sta., Portland, OR.

CONOVER, W. J. 1980. Practical nonparametric statistics. John Wiley and Sons,
New York.

Cook, C. W. and J. STUBBENDIECK, eds. 1986. Range research: basic problems and
techniques. Soc. for Range Management, Denver, CO.

HILL, M. O. 1973. Reciprocal averaging: an eigenvector method of ordination. J.
Ecology 61:237-249.

LEONARD, R., D. HARKIN, and P. J. ZINKE. 1968. Ecological study of meadows in
lower Rock Creek, Sequoia National Park: progress report for 1967. Copy on
file at U.S.D.A. Forest Serv. Pacific Northw. Forest and Range Exp. Sta., Fres-
no, CA.

, C. M. JOHNSON, and P. J. ZINKE. 1969. Ecological study of meadows in
lower Rock Creek, Sequoia National Park: progress report for 1968. Copy on
file at U.S.D.A. Forest Serv. Pacific Northw. Forest and Range Exp. Sta., Fres-
no, CA.

LOTAN, J. E. 1964. Initial germination and survival of lodgepole pine on prepared
seedbeds. U.S.D.A. Forest Serv. Res. Note INT-29.

and W. B. CRITCHFIELD. In press. Pinus contorta. In Silvics of forest trees

of North America. U.S.D.A. Forest Serv. Pacific Southw. Forest and Range Exp.

Sta., Berkeley, CA.

and D. A. Perry. 1977. Fifth-year seed: seedling ratios of lodgepole pine
by habitat type and seedbed preparation technique. U.S.D.A. Forest Serv. Res.
Note INT-148.

McLegan, E. O. 1982. Soil pH and lime requirement. Jn A. L. Page, R. H. Miller,
and D. R. Keeney, eds., Methods of soil analysis, Part 2. Chemical and micro-
biological properties— agronomy monograph 9 (2nd ed.), p. 199-224. ASA-SSSA,
Madison, WI.

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

NELson, D. W. and L. E. Sommers. 1982. Total carbon, organic carbon and organic
matter. Jn A. L. Page, R. H. Miller, and D. R. Keeney, eds., Methods of soil

90 MADRONO [Vol. 34

analysis, Part 2. Chemical and microbiological properties—agronomy mono-
graph 9 (2nd ed.), p. 539-579. ASA-SSSA, Madison, WI.

RATLIFF, R. D. 1982. A meadow site classification for the Sierra Nevada, California.
U.S.D.A. Forest Serv. Pacific Southw. Forest and Range Exp. Sta. Gen. Tech.
Rep. PSW-60.

. 1985. Meadows in the Sierra Nevada of California: state of knowledge.
U.S.D.A. Forest Serv. Pacific Southw. Forest and Range Exp. Sta. Gen. Tech.
Rep. PSW-84.

SHEPPERD, W. D. and D. L. Nose. 1976. Germination, survival, and growth of
lodgepole pine under simulated precipitation regimes: a greenhouse study. U.S.D.A.
Forest Serv. Res. Note RM-328.

USDA. 1974. Seeds of woody plants in the United States. Agric. Hdb. No. 450.
U.S.D.A. Forest Serv., Washington, DC.

(Received 6 Sep 1985; revision accepted 15 Dec 1986.)

ANNOUNCEMENT

WOMEN BOTANISTS RECOUNT CAREER HARDSHIPS AND HIGHLIGHTS

Three women botanists describe the highlights of their careers as herbarium
curators, collectors of native and ornamental plants, and conservationists of
flora and habitat in this initial volume of a new series of oral histories on
California Women in Botany. Produced by the University of California’s Re-
gional Oral History Office, the volume challenges the traditional view of botany
as the pastime for the ‘“‘weaker sex.”

The vivid tales of UC Herbarium botanist Annetta Carter, describing col-
lecting trips to Baja California with the eighty-year-old Annie Alexander, attest
to the hardships and joys of life in the field. Owens Valley botanist Mary
DeDecker’s recounting of her battles to protect the fragile habitat of her beloved
desert plants reflects great strength of purpose and fearless, informed persis-
tence. Elizabeth McClintock’s dedicated work for the California Academy of
Sciences herbarium reveals a third aspect of the many contributions of Cali-
fornia women botanists.

California Women in Botany is available for study at The Bancroft Library,
UC Berkeley, and at the UCLA Library Department of Special Collections.
The volume may be purchased from Friends of the Bancroft Library, Regional
Oral History Office, Univ. of California, Berkeley 94720. Price: $45.00.

INVASION OF PINUS CONTORTA VAR. MURRAYANA
(PINACEAE) INTO MOUNTAIN MEADOWS AT
YOSEMITE NATIONAL PARK, CALIFORNIA

JOHN A. HELMS
Department of Forestry and Resource Management,
University of California, Berkeley 94720

ABSTRACT

Stands of Pinus contorta var. murrayana (lodgepole pine) at the edge of many
meadows have a tiered appearance due to bands of trees of increasing size class.
Factors that might contribute to these waves of encroachment are seed availability
and seedling establishment. Seed fall and distribution were monitored on two mead-
ows in Yosemite National Park, California. Approximately 10,000—135,000 seeds/
ha were distributed annually across the meadows. This probably represents 2—13%
of seeds that fall commonly within lodgepole pine stands. First-year seedlings were
estimated to be 550—10,000/ha annually. None of the naturally-established seedlings
in the sample plots survived through the third growing season. Thus, waves of en-
croachment are more likely the result of success in establishment than of inadequate
numbers of seeds or short dispersal distance. Nine 26—40 m transects from the
meadow edge into the adjacent forest showed distinct periods of encroachment. On
the wetter meadow these were 1950-1962 and 1918-1936. A comparison with records
of precipitation available since 1907 shows that the two most recent periods of
encroachment are associated broadly with periods of lesser precipitation. On the drier
meadow, periods of encroachment were less distinct, but occurred during 1948-1973
and 1905-1931. Drier meadows are more conducive to pine establishment and en-
croachment is influenced less by patterns of precipitation.

Many mountain meadows in the Sierra Nevada, particularly the
smaller, drier ones, are being invaded gradually by Pinus contorta
var. murrayana (Grev. & Balf.) Engelm. (lodgepole pine) (De-
Benedetti and Parsons 1979a). Invasion of meadows is a natural,
dynamic process, but the factors that influence the rate of meadow
invasion are not well understood. The phenomenon is of managerial
and ecological interest. Conversion of meadows to stands of lodge-
pole pine reduces ecological diversity with adverse impacts on sce-
nic, recreational, habitat, and grazing values.

Lodgepole pine in the Sierra Nevada is a prolific seed producer
with crops of non-serotinous cones produced annually (Critchfield
1980). Seed fall in south-central Oregon has varied from 12,000 to
over 2 million seeds/ha (Dahms and Barrett 1975). Although lodge-
pole pine has small seeds that are among the most dispersible of
any North American conifers (Critchfield 1980), density of seedfall
at a distance of 20 m from the timber edge may be only 10-30% of
that within the stand, and most seeds fall to the ground within a
distance of about 60 m (Lotan 1975, Lotan and Critchfield in press).

MADRONO, Vol. 34, No. 2, pp. 91-97, 1987

92 MADRONO [Vol. 34

Invasion of lodgepole pine into meadows appears to be associated
with warm-dry weather, grazing, and fire. The relative dryness of
meadow soils is associated with the amount of snow and timing of
snowmelt, which in turn are affected by the size and orientation of
meadows (Anderson 1967). Similarly, Wood (1975) suggests that
lodgepole pine seedlings tend to become established in years of low
snowpack and that invasion patterns are affected by water table
fluctuations. In the southern Sierran region, for example, changes
in meadow vegetation are commonly associated with geological in-
stability that in turn causes changes in water status (Benedict 1982).
Vale (1981) concluded that although warm-dry weather is cited often
as a major factor influencing lodgepole pine invasion into meadows,
climatic fluctuations are typically less important than grazing or fire.
Meadow disturbances from livestock grazing and trampling favors
the establishment of pine seedlings, but also retards their develop-
ment. Pines rapidly encroached into meadows when intensive sheep
grazing ceased in 1900. Similarly, pine invasion into meadows of
Kings Canyon National Park followed after the expiration of cattle
grazing permits in the mid-1950’s (Sharsmith 1959). Fire due to
lightning strikes in grazed areas or due to the activities of Indians
also influenced pine regeneration in meadows (DeBenedetti and Par-
sons 1979b).

I conducted this study to better understand the factors that influ-
ence the rate and timing of lodgepole pine invasion into meadows.
My objectives were to determine 1) the dispersion of lodgepole pine
seed into meadows from neighboring stands, 2) the extent of seed
germination and establishment, and 3) the patterns ofrate and timing
of past lodgepole pine invasion into meadows.

METHODS

The large and small meadows selected for this study are located
at 2100 min Yosemite National Park, California, near Glacier Point
Road on the trail to Lost Bear Meadow. They are the same meadows
used for a concurrent study of lodgepole pine germination and es-
tablishment (Helms and Ratliff 1987). Detailed description of mead-
ow physiography, soils, and vegetation are presented in that paper.

Natural seed dispersal. A sampling system of 195 and 95 seed
traps was installed in the large and small meadows, respectively, in
the fall of 1981, 1982, 1983, and 1984. They were in position from
mid-August until snowfall was imminent in October. Traps were
0.1 m? in size, made of 2.5-mm-mesh wire screen, and positioned 1
m above the ground ona 10 X 10 m grid. The grids extended from
the southern to the northern edge, and were located in the eastern
half of each meadow.

1987] HELMS: PINUS CONTORTA VAR. MURRAYANA. II. oS

TABLE 1. NUMBERS OF SEEDS/HA (= s.e.) DISTRIBUTED IN EACH MEADOW IN FOUR
SUCCESSIVE YEARS.

Year Large meadow Small meadow

1981 28,830 + 6080 134,560 + 14,940
1982 23,240 + 4090 125,945 + 14,830
1983 9620 + 2850 24,760 + 6060
1984 20,861 + 8800 26,911 + 9300

Natural seedling establishment. A 0.1 m? plot was established 1
m north of each seed trap location. This provided a sample of 195
plots on the large meadow and 95 plots on the small meadow. Each
fall, from 1982 through 1984, plots were examined for the presence
of lodgepole pine seedlings.

Encroachment of lodgepole pine into meadows. Five transects were
established in the large meadow and four in the small meadow. Each
transect was 26-40 m long and extended from the meadow edge
into the adjacent forest. Along each transect, 15—20 trees were mea-
sured in terms of height and distance from meadow edge. The age
of each tree was determined by extracting a core from the stem at
ground level and counting annual rings. Approximately 20 repre-
sentative outlier trees within each meadow also were measured.

RESULTS

Natural seed dispersal. In each study year, a substantial cone crop
was observed on the dominant trees with exposed crowns. Cone
opening occurred on warm fall days. In the large meadow, the pro-
portions of seed traps containing seeds in 1981, 1982, 1983, and
1984, were 17, 16, 6, and 23%, respectively. In the small meadow,
the corresponding proportions in each year were 65, 88, 18, and
12%, respectively. The average annual seed fall on the large and
small meadow over the four year period was 10,000—29,000 and
25,000—135,000 seeds/ha, respectively (Table 1). In the 80-m-wide
small meadow, substantially more seeds were distributed within 30
m of the meadow edges than in the center of the meadow. This trend
was not found in the large 220-m-wide meadow where the seeds
were distributed more evenly.

Natural seedling establishment. In the large meadow, the total
numbers of seedlings found in the 195 plots in 1982, 1983, and 1984
represent 550, zero, and 2200/ha, respectively. This is equivalent
to two, zero, and 22% of seeds dispersed in the previous year (Table
1). On the small meadow, the total numbers of seedlings found in
the 95 plots in the same years represent 4500, 4500, and 10,200/

94 MADRONO [Vol. 34

Height (m)

0 5 10 15 20 25 30 35
Distance from Meadow Edge into Adjacent Forest (m)

Fic. 1. Typical distribution of height- and age-classes in meadow-edge lodgepole
pine of the large meadow.

ha, respectively, or 3%, 3%, and 41% of seeds dispersed in the pre-
vious year (Table 1).

In both meadows, the highest first-year survival occurred in 1984,
which followed the smallest seed crop measured during the 4 yr
study. All seedlings observed over the 3 yr of the study were ephem-

4 SMALL MEADOW

Number of Trees

LARGE MEADOW

(@) 20 40 60 80 100 120 140 160 180
Age (yr)

LET aa ag ae FETA mr SEURETT, (ES) (SSR Fh (ee Fa FN TT A
1980 1950 1900 1850 1800

Calendar Date

Fic. 2. Frequency distribution of age classes of meadow-edge trees (solid bars)
from a total of nine 26-32 m transects, and all outlier lodgepole pine trees (hatched
bars) in both the large and small meadows. Arrows indicate periods during which
most encroachment of meadows occurred.

1987] HELMS: PINUS CONTORTA VAR. MURRAYANA. I. 95

ANNUAL PRECIPITATION

Yosemite National Park

eee

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et

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ae

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ogi Saas
= a! Gee
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= 7.27
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50

1920 40 60 1980
Calendar Date
Fic. 3. Complete record of annual precipitation at the valley floor, Yosemite

National Park, California. Hatched areas show periods during which most lodgepole
pine encroached on the larger, wetter meadow.

eral. In only one case did a seedling survive into year two, but it
did not survive through the third season.

Encroachment of lodgepole pine into meadows. The typical pattern
of meadow-edge trees contained distinctly different age and height
classes (Fig. 1). The large meadow was bordered by trees of two age
classes: 21-33 and 47-65 yr and some older individuals that ranged
from 80-170 yr (Fig. 2). These waves of encroachment occurred in
the periods: 1950-1962 and 1918-1936, with older trees having
become established prior to 1900. The small meadow was bordered
by two, less distinct age classes: 10-35 and 50-80 yr and some older
individuals that ranged from 115-150 yr. These age classes represent
waves of encroachment occurring in 1948-1973 and 1905-1931,
and some older trees established prior to 1860.

In the large meadow, two classes of outlier trees were identified:
1) 23.8 + 1.3 yr and 6.02 + 0.53 m in height, and 2) 49.0 + 4.1 yr
and 11.97 + 0.79 m in height. These two classes correspond to the
two youngest age classes of meadow-edge trees (Figs. 1 and 2). In
the small meadow, the corresponding two classes were: 1) 13.6 +
0.5 yr and 1.5 + 0.13 min height, and 2) 34.5 + 1.1 yrand 12.68 +

96 MADRONO [Vol. 34

0.46 m in height. These two classes occur within the youngest age
class of meadow-edge trees (Fig. 2).

No relationship was found between age of outlier tree and distance
from the meadow edge. Outliers were located on slightly higher,
drier locations within the meadows.

Correlation between encroachment and precipitation patterns. The
periods during which most encroachment occurred on the larger,
wetter meadow (Fig. 2) were superimposed on a complete record of
annual precipitation (1907-1984) in Yosemite Valley (Fig. 3). Av-
erage precipitation during periods of no encroachment was 97.1 +
5.4 cm. In comparison, average precipitation during periods of suc-
cessful regeneration, was 80.5 + 3.2 cm. The difference between
these means was significant (p = 0.02). The smaller, drier meadow,
in which channel erosion had lowered the water table, showed no
apparent correlation between encroachment and precipitation.

DISCUSSION

Encroachment of pine seedlings into meadows was limited by lack
of seedling survival rather than by inadequate seed supply or insuf-
ficient numbers of seedlings. In the large meadow, seeds distributed
in the fall that escape predation may be washed away by free-flowing
water that generally covers parts of the meadow until mid-summer.
Invasion of lodgepole pine into meadows, therefore, is more likely
at the meadow edge where there is higher seed fall and less flooding.
The similar ages of both edge trees and outlier trees within the
meadows suggests that conditions favorable for pine establishment
in both locations occur concurrently.

The association between periods of most encroachment of lodge-
pole pine and periods of less precipitation (Fig. 3) is not strong, but
provides support for the concept that invasion occurs under drier
conditions.

In a study of lodgepole pine invasion near the Tioga Pass entrance
to the park, the oldest tree found was established in 1866 (Vale
1981). Most trees were established from 1910-1975. No distinct
pattern of invasion was found; however, most trees were established
in 1925. Other periods reported to be associated with invasion are
1853-1875 and 1898-1909 (Boche 1974), 1910 (Vankat and Major
1978), and 1903, 1906, and 1924 (Wood 1975). Results of the pres-
ent study suggest that variability in reported periods during which
invasion occurred may be associated with wetness and size of mead-
Ow.

Rates of encroachment from 1868-—present, determined by the
relationship between age of trees and distance from the meadow
edge, were 0.19 and 0.22 m yr! for the large and small meadow,

1987] HELMS: PINUS CONTORTA VAR. MURRAYANA. II. oF

respectively. At a similar rate of encroachment and with current
meadow conditions the larger and smaller meadows will be invaded
completely within ca. 580 and 180 yr, respectively.

ACKNOWLEDGMENTS

This study was supported by the USDA Forest Service, Pacific Southwest Forest
and Range Experiment Station, Berkeley, California, Project Number PSW-80-0022.
It also was part of the University of California’s Agricultural Experiment Station
Project 2942-MS. I thank Drs. W. Critchfield and F. Vasek for their helpful suggestions
to improve the manuscript.

LITERATURE CITED

ANDERSON, H. W. 1967. Snow accumulation as related to meteorological, topo-
graphic, and forest variables in the Central Sierra Nevada, California. Int. Assoc.
Sci. Hydrology Publ. No. 78:215-224.

BENEDICT, N. B. 1982. Mountain meadows: stability and change. Madrono 29:148-
153:

BocuHE, K. E. 1974. Factors affecting meadow-forest borders in Yosemite National
Park, California. M.S. thesis, Univ. California, Los Angeles.

CRITCHFIELD, W. B. 1980. Genetics of lodgepole pine. U.S.D.A. Forest Serv. Res.
Pap. WO-37, Washington, DC.

DauHMs, W. G. and J. W. BARRETT. 1975. Seed production of central Oregon pon-
derosa and lodgepole pines. U.S.D.A. Forest Serv. Res. Pap. PNW-191. Pacific
Northw. Forest and Range Exp. Sta., Portland, OR.

DEBENEDETTI, S. H. and D. J. PARSONS. 1979a. Mountain meadow management
and research in Sequoia and Kings Canyon National Parks: a review and update.
In R. Linn, ed., Proc. First Conf. on Sci. Res. in the National Parks, Nov. 9-
12, 1986. U.S.D.I. Natl. Park Serv. Trans. and Proc. Series No. 5, Washington,
DC 11:1305-1311.

and . 1979b. Natural fire in subalpine meadows: a case description
from the Sierra Nevada. J. Forest. 77:477-479.

HeEtms, J. A. and R. D. RATLIFF. 1987. Germination and establishment of Pinus
contorta var. murrayana (Pinaceae) in mountain meadows of Yosemite National
Park, California. Madrono 34:77-90.

LoTAN, J. E. 1975. Regeneration of lodgepole pine forests in the northern Rocky
Mountains. Jn Management of lodgepole pine ecosystems: symposium proceed-
ings, vol. 2:516-535. Coop. Ext. Serv., Washington State Univ., Pullman.

and W. B. CRITCHFIELD. In press. Pinus contorta. In Silvics of forest trees
of North America. U.S.D.A. Forest Serv. Pacific Southw. Forest and Range Exp.
Sta., Berkeley, CA.

SHARSMITH, C. 1959. A report on the status, changes, and ecology of back country
meadows in Sequoia and Kings Canyon National Parks. Natl. Park Service.
(unpubl.)

VALE, T. R. 1981. Ages of invasive trees in Dana meadows, Yosemite National
Park, California. Madrono 28:45-47.

VANKAT, J. L. and J. MAyjor. 1978. Vegetation changes in Sequoia National Park,
California. J. Biogeogr. 5:377—402.

Woop, S. H. 1975. Holocene stratigraphy and chronology of mountain meadows,
Sierra Nevada, California. Ph.D. dissertation, California Inst. Tech., Pasadena.

(Received 6 Sep 1985; revision accepted 15 Dec 1986.)

THE DISTRIBUTION OF FOREST TREES IN NORTHERN
BAJA CALIFORNIA, MEXICO

RICHARD A. MINNICH
Geography Program, Department of Earth Sciences,
University of California, Riverside 92521

ABSTRACT

This survey includes maps and provides descriptions of the distribution of 22
Pacific Coast temperate trees, including two endemics, in northern Baja California,
from the international border south to latitude 30°. Ranges were discerned from aerial
photographs and verified by field reconnaissance, botanical collections, and a low-
altitude aerial flight. With the exception of mixed conifer and pinyon forests, most
forests comprise single tree species that represent fragments of more diverse ecosys-
tems from more mesic areas of California. The rapid decline in the diversity of Pacific
Coast temperate trees below the international border reflects strong precipitation
gradients associated with orography. Several California tree species have been re-
ported erroneously in northern Baja California due to the misidentification of spec-
imens, or the misinterpretation of common plant names or place names.

The southern geographic limits of many Pacific Coast temperate
trees are in the mountains and coastal valleys of northern Baja
California, Mexico. From various sources, Griffin and Critchfield
(1976) mapped the ranges of these trees for California, but detailed
maps stop at the international border. This survey includes detailed
maps and provides descriptions of the natural distribution of 22
trees, including two endemics, in northern Baja California, from the
international border south to latitude 30° (Fig. 1).

METHODS

Many localities are derived from botanical collections and field
observations. Extensive collections, particularly by Wiggins (1980,
DS), and Moran (SD) have probably recorded most of the flora of
northern Baja California. I also consulted other botanical collections
from SD, RSA, UCLA, and UCR. On each map I have plotted each
collection that had definite locality data. Point records, however, are
difficult to extrapolate into broader geographic distributions, because
collections are invariably non-random and reflect particular interests
and access. Indeed, many remote areas have never been visited.

Important locality data come from the extensive library of aerial
photography at the Department of Earth Sciences, University of
California, Riverside (Table 1). These photographs show the forests
of northern Baja California, which are conspicuous in this semiarid
shrubland landscape. I studied photographs on a Bausch and Lomb

MADRONO, Vol. 34, No. 2, pp. 98-127, 1987

a9

MINNICH: TREES OF BAJA CALIFORNIA

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102 MADRONO [Vol. 34

roll film stereoscope with 3 and 8 X magnification. Tree species were
recognized from gross characters that included crown perimeter and
shape, vertical structure, shadows, and color (hue) (Table 2).

Interpretation was verified by field reconnaissance and botanical
collections. The process of developing diagnostic identification of
trees required progressive cross-referencing between aerial photog-
raphy and ground observations. Field data that were plotted on work
maps or recorded on ground photographs also were correlated with
aerial photographic signatures in the laboratory, and photographic
information was verified in the field. I traveled through many for-
ested areas of the Sierra Juarez, Sierra San Pedro Martir, and several
coastal sites (Minnich 1986). I flew one low-altitude reconnaissance
(within 600 m of ground) in May 1986 from Tijuana to Cerro Bola,
Valle Guadalupe, Cerro Los Pinos, Sierra San Pedro Martir, Sierra
Juarez, Valle las Palmas, and back to Tijuana.

Tree ranges were transferred from photographs onto 1:250,000
topographic quadrangles using a Bausch and Lomb Zoom Transfer
Scope. The ranges shown on Figs. 2—16 will be modified by subse-
quent field research because the scale of aerial photography results
in omissions, particularly local outposts and among species forming
scattered small stands. Subtropical trees such as Washington filifera
Lindl., Erythea armata Wats., and Prosopis juliflora (SW.) DC. are
excluded from this treatment.

PHYSICAL SETTING

Physiography. The coastal valleys and mountains of northern Baja
California extend into the southern part of the Mediterranean cli-
matic zone along the Pacific Coast. Although the region is at the
southern margin of reliable winter cyclonic storms, seasonally moist
habitats are widespread because the large relief of mountainous ter-
rain provides cooler temperatures with altitude and encourages oro-
graphic precipitation.

The physiography of the region can be subdivided into three ranges
of the peninsular range province (Fig. 1) (Gastil et al. 1975). The
Sierra Juarez, an extension of the Laguna Mountains of southern
California, is an undissected plateau (elev. 1200-1800 m) of mostly
granite substrate from the international border southeastward to near
Santa Catarina. The southern third of the Sierra Juarez, south of
Santa Catarina, is an extensive tableland of mesas capped by Mio-
cene volcanics, with summits to nearly 2000 m.

Toward the Pacific coast, there is a discontinuous series of dis-
sected, lower ranges (1000-1400 m), termed here the coastal Sierra
Juarez, that extends southeastward from an unnamed range south-
west of Valle las Palmas (Cerro Bola, 1280 m) to Ensenada Bay.
Substrates include a granitic batholith, and extensive prebatholithic

1987] MINNICH: TREES OF BAJA CALIFORNIA 103

(Cretaceous) undifferentiated volcanics, metavolcanics, and marine
sediments (Alistos formation). The coastal and interior Sierra Juarez
are separated by alluvial basins and low plateaus, including Valle
las Palmas, Valle Guadalupe, Valle Ojos Negros, and an extensive
high basin between Santa Catarina and El Alamo. South of Ensenada
Bay, the coastal Sierra Juarez increases in altitude (1200-1500 m)
and turns eastward along the Agua Blanca fault.

South of Valle la Trinidad, these transverse coastal ranges join
the Sierra San Pedro Martir batholith, characterized by a steep west-
ern scarp (vertical relief 700-1000 m) and extensive plateaus along
the crest, the elevations of which decrease in steps from 2500 m in
the north to 1800 m at Cerro Matomi. The area from the west scarp
to the ocean contains extensive low foothills and mesas. These in-
clude a series of low northwest-southeast trending coastal ranges
from Valle Santo Tomas to Colonet. The substrate is derived mostly
from the Alistos formation, with Cretaceous marine sedimentary
rocks (Rosario formation) outcropping along the coast. The faulted
eastern scarps of the Sierra Juarez and Sierra San Pedro Martir are
rugged with numerous small canyons.

Climate. Annual precipitation results mostly from frontal storms
that occur between December and March. It ranges from 200-300
mm in the coastal valleys to 500 mm in the mountains, and only
100 mm on lee slopes at the margin of the Sonoran Desert (Table
3). Heaviest precipitation occurs on the highest peaks of the coastal
ranges, and on the western slopes of both the interior central Sierra
Juarez and Sierra San Pedro Martir. Despite their altitude, the south-
ern Sierra Juarez tablelands are arid due to rainshadows created by
the relatively high coastal peaks south of Ensenada. Because high
relief becomes discontinuous along the peninsula south of the Sierra
San Pedro Martir, the rainfall, which is dependent primarily upon
orography, becomes unreliable south of lat. 30°, where the Sonoran
Desert extends west to the Pacific Coast.

Winters are mild from the coast inland to the interior valleys
(Table 3). Ground inversions, however, produce cold nights and
hard frosts in high mountain plateaus. Snowfall may account for no
more than ca. 15% of the annual precipitation in the Sierra Juarez,
but perhaps as much as 50% in the higher Sierra San Pedro Martir
(Minnich 1986a).

During summer, near-coastal valleys and western slopes of the
coastal Sierra Juarez are cool, humid, and foggy because of the
onshore flow of marine air with sea breezes and valley wind systems,
as in California. Interior mountains and valleys are warm and dry
except for occasional afternoon thundershowers that mostly occur
along the eastern walls of the Sierra Juarez and Sierra San Pedro
Martir and are caused by surges of tropical moisture that move north
along the Gulf of California from the subtropical Pacific Ocean.

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104

1987] MINNICH: TREES OF BAJA CALIFORNIA 105

SPECIES DISTRIBUTIONS

The geographic distributions of northern Baja California trees
reflect a number of factors, including altitudinal zonation, topog-
raphy, substrate, and climatic gradients associated with orography,
distance from the Pacific Coast, and latitude. With the exception of
mixed conifer forests (Pinus jeffreyi, P. lambertiana, Abies concolor,
Calocedrus decurrens) in the Sierra San Pedro Martir and xeric pin-
yons (Pinus quadrifolia, P. monophylla), most forests include single
tree species with highly localized distributions that represent frag-
ments of more diverse ecosystems in more mesic areas of California.

ABIES CONCOLOR (Gord. & Glend.) Lindl. (Fig. 2). White fir is
common above 2200 m in the mixed conifer forest that covers the
Sierra San Pedro Martir. Individuals have been found as low as 1900
m along Arroyo los Pinos near Corral de Sam (Table 4). Although
Pinus jeffreyi is the dominant tree of most forests in the range, A.
concolor is locally dominant with P. lambertiana on steep northern
exposures in the vicinity of Cerro Venado Blanco, Cerro la Botella
Azul, and upper headwalls of the eastern escarpment, including Pi-
cacho del Diablo; a photo by Clyde (1975:85) records a sapling at
the summit (3095 m). I have not seen A. concolor in extensive forests
of P. jeffreyi south of La Grulla and La Encantada meadows. This
tree has short, thick, wide leaves of the southern California-Rocky
Mountain variety (Vasek 1985). The nearest stand is 180 km north
in the Cuyamaca Mountains of San Diego Co., California (Griffin
and Critchfield 1976), and, thus, is not known from the Sierra Juarez.

CALOCEDRUS DECURRENS (Torr.) Florin (Fig. 3). Incense cedar is
rare even in the highest mountains of northern Baja California. In
the Sierra San Pedro Martir it grows mostly near streams from 1350-
2400 m on the northern and eastern scarps of the plateau. It also
has been collected along several arroyos to the south, as at La Corona,
Valladares Creek, La Vibora, and La Encantada. The southernmost
locality I have seen is along a gully 5 km south of La Grulla. Beyond
stream habitats, incense cedar is occasional in mixed conifer forests
at Vallecitos, including the largest tree of any species I have seen in
the range (3 m dbh, 45 m height).

Moran found few C. decurrens groves in moist habitats within
Pinus jeffreyi forest in the central Sierra Juarez, including La Matanza
meadow, and Arroyo El Tule. Aerial photographs confirm his ob-
servation that the tree is relatively abundant along the canyon at El
Tule. Federal foresters in Baja California have seen the tree along
the arroyo that drains Laguna Juarez.

CUPRESSUS ARIZONICA Greene var. STEPHENSONII (C. B. Wolf)
Beauchamp (Fig. 4). This variety of Arizona cypress was believed
to be endemic to the Cuyamaca Mountains in San Diego Co. (Griffin
and Critchfield 1976) until another larger cluster of populations was

[Vol. 34

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discovered by Broder (1963) in the southern Sierra Juarez along
Mesa Huicual, Mesa Valle Seco, and adjacent Canada El Rincon
southwest to near Santa Catarina (1200-1545 m). Trees along Cana-
da El Rincon are apparently old with large bole diameters (to 2 m),
whereas stands on mesas are young due to chaparral fires (Moran
1972). A vaquero told Moran (pers. comm.) that the cypress also
occurs in the next canada south of El Rincon. In the 1986 aerial
reconnaissance, I saw small colonies of this species to the southeast
on an unnamed mesa south of Canada El Rincon, on Mesa la Vinata
Romero, in Canon Alamito, and in Canada la Esperanza. All stands
grow on Miocene volcanics or Miocene postbatholithic continental
sedimentary rocks (Gastil et al. 1975). Also, they occur in an area
having a bimodal precipitation regime with limited winter rains,
due to a rainshadow effect of the coastal Sierra Juarez, and with
summer thundershowers. These conditions are similar to those of
Arizona where C. arizonica occurs.

CUPRESSUS GUADALUPENSIS Wats. subsp. FORBESII (Jeps.) Beau-
champ (Fig. 4). Cupressus guadalupensis subsp. forbesii occurs in
San Diego County at Tecate Mountain, Otay Mountain, and near
Guatay southwest of Cuyamaca Mountain (Griffin and Critchfield
1976). Botanical collections and aerial photographs show that these
border populations are the northern limit of an extensive disjunct
pattern of small groves. They are far more numerous than shown
by Wolf (1948) and Little (1971) and occur for 150 km along coastal
foothills of northern Baja California.

The northernmost stands in Mexico are extensions of U.S. pop-
ulations at the border. Numerous colonies occur in near-coast foot-
hills and mesas of the Cerro Bola range, especially at the south,
between Cerro San Felipe and Canada El Golpe. Interior outliers
occur at Cerro Grande and 10 km south of Tecate. Another cluster
of groves extends along the coast ranges north and east of Ensenada,
from Rancho de la Cruz to Cerro Los Pinos, including Cerro Mira-
cielo, Canon los Cipreses, | km north of Cerro El Toro, and along
the wash and adjacent northern exposures of Canon San Carlos. To
the south, isolated populations occur in coastal canyons with Pinus
muricata west of San Vicente, above Rancho los Zaguaritos, an
inland site at the summit of Cerro El Cipres (N), the upper head-
waters of Canon Nueva York, and at another Cerro El Cipres (S).
Many groves are found at locations with “‘cipres”’ in place names
on 1:50,000 scale topographic sheets.

Most stands grow in chaparral and are even-aged because of brush-
fires (see review by Vogl et al. 1977); scattered trees often occur in
adjacent arroyos. The species grows between 200 and 1200 m, mostly
on the Alistos or Rosario formations, although some inland and
southern populations are on granodiorites. Nearly all stands grow
on northern exposures, as in southern California (Vogl et al. 1977).

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Cupressus guadalupensis also forms a significant forest on Guadalupe
Island.

CUPRESSUS MONTANA Wigg. (Fig. 4). Cupressus montana is en-
demic to the Sierra San Pedro Martir. Most stands are associated
with a Pinus lambertiana-Abies concolor forest on steep granite walls
along the eastern rim above 2200 m in the upper headwaters of
Canon del Diablo, Canon la Providencia, and on the massive faces
of Picacho del Diablo to near the summit (3095 m). Unlike many
species of cypress, this tree develops great girth at the bole (1-2 m
diam) and, thus, may be long-lived because forest productivity at
these altitudes is diminished by cold climate. Juveniles may be seen
down to 1400 m along Canon del Diablo with Quercus chrysolepis,
Calocedrus decurrens, and Salix spp. Scattered trees also are found
along arroyos on the plateau, near Los Llanitos, above La Encantada,
and on the main drainage between this meadow with La Grulla.
Moran found one tree at La Vibora.

FRAXINUS VELUTINA Torr. var. CORIACEA (Wats.) Rehd. (Fig. 13).
Arizona ash has been recorded in botanical collections in the south-
ern Sierra San Pedro Martir and adjacent coastal foothills, the Uribes,
and along an unnamed arroyo near Canada El Alamoso (800-1000
m). Wiggins collected the tree at Arroyo de Agua Amarga and at
Agua Caliente de Guadalupe on the desert slope. Fraxinus velutina
also occurs in San Julio Canyon in the Sierra San Francisco of central
Baja California. The absence of records for Arizona ash in more
mesic parts of the Sierra San Pedro Martir and Sierra Juarez is
peculiar. Several colonies in San Diego Co. at the international bor-
der (Griffin and Critchfield 1976) suggest that more intensive ex-
ploration will uncover other stands in northern Baja California.

PINUS ATTENUATA Lemmon (Fig. 5). Until recently, knobcone pine
was collected only in the vicinity of Rancho de la Cruz, Canon Arce,
and Canada Dona Petra (250-500 m) on the west flank of Cerro
Miracielo (1100 m), north of Ensenada (cf. Map 58 in Critchfield
and Little 1966). Aerial photographs from 1938 and 1956, however,
show numerous stands along the north slope of this peak to near
Cerro Blanco and Canon Borreguero. Three groves were discovered
by aerial reconnaissance 20 km southeast on a ridge at Cerro los
Pinos. Pinus attenuata was recently collected in Canon El Carmen,
west of Valle Guadalupe. Occasional stands also may be present
among numerous groves of Cupressus guadalupensis subsp. forbesii
in the southern Cerro Bola range.

Similar to C. guadalupensis subsp. forbesii, P. attenuata grows in
dense chaparral on the Alistos formation. Stands have an even-aged
structure associated with canopy fires that are characteristic of the
closed-cone pines (Vogl et al. 1977). Aerial photographs from 1972
show that most groves of P. attenuata on Cerro Miracielo were
burned in a large fire in the late 1960’s, but saplings were observed

"SUOTIOI]

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throughout the burn in the 1986 aerial reconnaissance. The nearest
stand in California is 200 km to the north, on Pleasants Peak in the
Santa Ana Mountains (Griffin and Critchfield 1976).

PINUS CONTORTA Dougl. ex Loud. (Fig. 6). Lodgepole pine is con-
fined mostly to the Vallecitos and La Tasajera basins in the Sierra
San Pedro Martir (2300-2400 m), where it typically becomes the
dominant tree at edges of meadows. Smaller colonies descend ad-
jacent arroyos. This subalpine forest tree is absent from mixed co-
nifer forests covering higher ridges east of Vallecitos (2700-2900 m)
except for scattered trees on northern exposures near the summits
of Cerro la Botella Azul, Cerro “2828” (observatory), and Cerro
Venado Blanco. A few individuals were reported recently from Pi-
cacho del Diablo at 3095 m (M. Hamilton, pers. comm.).

Characteristically, P. contorta prefers to grow on poorly drained
sites where potential competitors cannot grow (Fowells 1965). Strong
nocturnal ground inversions with temperatures as low as — 15°C in
winter and O°C in summer (Alvarez 1981) also may permit this
subalpine species to grow in basin floors. Its absence from the highest
peaks may be due to rainshadow effects that extend from the wetter
western rim of the plateau, where scattered populations are found
along watercourses down to 2200 m. The nearest forest of P. contorta
in California is in the San Jacinto Mountains (270 km north).

PINUS COULTERI D. Don (Fig. 7). Coulter pine is rare in northern
Baja California (Minnich 1986b). Most colonies grow in mixed chap-
arral, often with Quercus chrysolepis, on highly resistant bedrock in
mesic parts of the interior sierra. The only stand (100 ha) in the
coastal Sierra Juarez is on Sierra Blanca (1250 m), southeast of Valle
Guadalupe (cf. Griffin and Critchfield 1976). In the interior Sierra
Juarez, isolated colonies grow on granites that occur southwest of
Rancho San Faustino (1500 m) and hillsides immediately west and
northwest of Laguna Juarez (1800 m). Aerial photographs indicate
a number of small colonies in similar habitats between these local-
ities. To the south, Moran (1977) found small populations at 1600
m on the Miocene volcanic tablelands on the north and south rims
of Canada El Rincon. Large colonies on the western margin of Mesa
Huicual are clearly evident on aerial photographs. Moran (1972)
stated that the cones in these populations were unusually small for
the species, but did not mention whether there was evidence of
hybridization with P. jeffreyi, which grows within 2 km in Arroyo
El Rincon.

Several groves of P. coulteri were found by aerial reconnaissance
in the Sierra San Pedro Martir. Stands as large as 100 ha occur on
northern exposures of the main divide immediately north of the
plateau; another is on the headwaters of Arroyo la Palizada. In the
southern part of the range, several stands are on the northeast and
northwest flanks of Cerro “2040”. The San Pedro Martir stands are

~

MADRONO

[Vol. 34

MMasjjal Snug

114

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poddew svoie popeys ‘Moaiffal snuig JO UOTINGLINSIP 94 |,

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1987] MINNICH: TREES OF BAJA CALIFORNIA 115

all on northern exposures from 1900-2150 m. They also form even-
aged stands that established after fires recorded on 1956 aerial pho-
tographs. According to Griffin and Critchfield (1976), three other P.
coulteri localities occur in the range: northeast of Corral de Sam,
upper La Corona Meadow (a single tree), and 8 km northwest of
Santa Rosa Meadow. Although I have not seen any P. coulteri at
these sites, Rojas-Gomez recently collected it at the northeast Corral
de Sam site.

PINUS JEFFREYI Grev. & Balf. (Fig. 8). Jeffrey pine is the most
important tree of mixed conifer forests along the crests of the Sierra
Juarez and Sierra San Pedro Martir (Minnich 1986b). Except for a
few colonies of C. decurrens near watercourses, P. jeffreyi is the only
tall tree in the Sierra Juarez, and forms extensive forests along mead-
ows, basin floors, and watercourses above 1400 m from Valle Los
Pinos to Arroyo El Rincon.

The northernmost stands in the Sierra San Pedro Martir are five
groves of ca. 50 trees on Cerro San Matias, an isolated peak (2100
m) 10 km north of the plateau. It recurs in several nearby basins
(1600 m) in association with woodlands of Quercus peninsularis in
the La Palizada, El] Huico, and San Rafael drainages. On the Sierra
San Pedro Martir plateau above 2100 m, P. jeffreyi forms a zonal
forest on slopes and basins above the chaparral belt in association
with Pinus lambertiana, Abies concolor, Quercus chrysolepis and
scattered understory of montane chaparral (Arctostaphylos patula
Greene var. platyphylla Wells, A. pungens HBK., A. pringlei Parry,
A. peninsularis Wells, Ceanothus cordulatus Kell., Rhamnus cali-
fornica Esch., and Quercus peninsularis). It occurs up to 2900 m on
the south face of Cerro Botella Azul. Scattered stands descend ar-
royos on the eastern scarp where it grows with Calocedrus decurrens.
South of La Grulla and La Encantada meadows, P. jeffreyi again
retreats to edges of meadows, basin floors, and arroyos down to ca.
1400 m. Moran found a few trees at 650 m near Rio San Antonio
on the west slope. The southernmost stands occur on Arroyos Fresnal
and San Simon.

PINUS LAMBERTIANA Dousgl. (Fig. 9). Sugar pine grows on the Valle-
citos surface of the Sierra San Pedro Martir, mostly on steep rocky
slopes and cliffs in association with mixed conifer forest. It descends
below elevations of Abies concolor (2100 m), with outposts extending
further south beyond La Grulla and La Encantada meadows to Cerro
Picacho la Vibora and scattered northern exposures near the Arroyo
El Cajon jumpoff. A single tree at 1700 m was collected by Moran
at Arroyo los Pinos near Rancho San Pedro Martir. Sugar pine is
locally dominant on the upper headwaters of the precipitous eastern
rim and on Picacho Del Diablo up to 3000 m. The nearest stands
in California are 200 km north, in the Cuyamaca Mountains of San
Diego Co. (Griffin and Critchfield 1976).

116 MADRONO [Vol. 34

PINUS MONOPHYLLA Torr. & Frém. (Fig. 10). Singleleaf pinyon
grows almost exclusively along the arid eastern scarps of the Sierra
Juarez and Sierra San Pedro Martir. The geographic extent of P.
monophylla was underestimated by Critchfield and Little (1966),
who based their report on limited botanical collections.

In the Sierra Juarez, P. monophylla forms extensive forests above
1000 m in association with desert chaparral characteristic of the
peninsular ranges (Rhus ovata Wats., Quercus dunnii Kell., Q. cor-
nelius-mulleri Nixon and Steele, Rhamnus crocea Nutt., Yucca schi-
digera Roezl ex Ortges., Y. whipplei Torr., Juniperus californica
Carr.). A few pinyons cross the border at Jacumé into southeastern
San Diego Co. A major forest with P. guadrifolia occurs on a large
plateau surface from La Rumorosa to El Topo, with P. quadrifolia
dominant on the wetter western margin and P. monophylla dominant
on the eastern rim. In the wetter central Sierra Juarez, P. monophylla
forests decrease to a narrow belt along the eastern scarp. Scattered
colonies occur on both Pacific and desert flanks of the arid southern
Sierra Juarez tablelands and northern foothills of the Sierra San
Pedro Martir adjacent to Valle la Trinidad.

Extensive forests with desert chaparral understory dominated by
Arctostaphylos peninsularis and Quercus peninsularis are found on
the eastern flank of the Sierra San Pedro Martir from 1200-2000
m, above which it is gradually replaced by P. quadrifolia. As the
elevation of the range decreases south of 30°50’N, P. monophylla
decreases to scattered outposts on high ridges within the east scarp.
It was collected on the north slope of Cerro Matomi. The southern
limit of the species is near Cerro San Luis in north central Baja
California.

PINUS MURICATA D. Don (Fig. 5). In northern Baja California, this
closed-cone pine is known only from the foggy coastal foothills
southwest of San Vicente, including Arroyo San Isidro (see Map 59,
Critchfield and Little 1966), Arroyo Hediondo, and near Punta San
Isidro. All stands grow in chamise chaparral or succulent coastal
sage scrub (Mooney 1977) in the upper Cretaceous Rosario for-
mation. A few colonies grow with Cupressus guadalupensis subsp.
forbesii. Pinus muricata stands reported on Cedros Island (Critch-
field and Little 1966) were recently named P. radiata D. Don var.
cedrocensis (Howell) Axelrod (Axelrod 1980).

PINUS QUADRIFOLIA Parl. ex Sudw. (Fig. 11). Four-needled pinyon
is the most widespread coniferous tree in northern Baja California,
but rarely occurs below 1200 m. Over most ofits range, P. quadrifolia
forms scattered groves within relatively dense chaparral of Adenos-
toma fasciculatum H. & A. and A. sparsifolium Torr. on the mesic
western flank of the interior Sierra Juarez or continuous forests with
desert chaparral along the crest and east rim. Pinus quadrifolia is
often allopatric with P. monophylla, which is confined to the arid

PLT.

MINNICH: TREES OF BAJA CALIFORNIA

1987]

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118 MADRONO [Vol. 34

eastern scarps, similar to their distributions in the Peninsular Ranges
of southern California (Map 16, Critchfield and Little 1966).

A few populations occur west of Jacumé near the international
border. The most extensive forest overlaps with P. monophylla on
the arid La Rumerosa-El Topo plateau, with smaller stands extend-
ing into shallow basins to the west. Numerous colonies pepper the
chaparral belt on the Pacific slope of the wetter central Sierra Juarez,
southward to Arroyo Barbon where the tree nearly drops out. The
P. quadrifolia belt on desert drainages also narrows between the east
margin of P. jeffreyi forests and P. monophylla forests on the eastern
rim. Pinus quadrifolia then expands to form scattered cover on the
Pacific and desert faces of the volcanic tablelands in the southern
part of the range. Populations also extend westward along arid lee
slopes on the coastal Sierra Juarez, including the Santa Catarina
basin, E] Alamo, and northwest to the southern edge of Valle Ojos
Negros.

In the Sierra San Pedro Martir, numerous groves of P. quadrifolia
grow in mostly Adenostoma chaparral in higher basins north of the
plateau. It then decreases to infrequent small colonies in dense chap-
arral on the mesic west flank of the range above the Meling Ranch.
Stands become more frequent in the drier southern part of the range.
Small outposts extend locally westward on higher spurs such as Mesa
Barreal and major arroyos. On the eastern scarp, P. quadrifolia forms
an extensive forest with Arctostaphylos peninsularis, Quercus pen-
insularis, and Q. chrysolepis understory from Cerro Venado Blanco
to Arroyo El Cajon (1500-2500 m), where it meets Pacific slope
groves. Although P. quadrifolia was collected up to 2700 m near the
observatory, it is almost absent from the mixed conifer forest belt
above 2100 m on the plateau. The southern limit is near Cerro
Matomi.

Lanner (1974) provides evidence of extensive hybridization be-
tween Pinus quadrifolia and P. monophylla on the La Rumerosa-El
Topo plateau. He suggests that P. quadrifolia should be replaced by
Pinus juarezensis Lanner in this area.

PLATANUS RACEMOSA Nutt., POPULUS FREMONTII Wats. (Fig. 12).
On aerial photographs, these riparian trees cannot be separated, but
can be distinguished from other trees by their deciduous habit, can-
opy structure, and row-like stand arrangement along streams. Field
reconnaissance indicates that most riparian forests consist primarily
of P. fremontii.

The size of riparian forests is broadly proportional to streamflow.
Stands are intermittent in the coastal Sierra Juarez where surface
water is rarely permanent, except at Canon Agua Escondida, and
streams that cut through the ranges from the interior valleys. More
continuous gallery forests follow the major arroyos descending the
Pacific slope of the interior Sierra Juarez plateau below 1500 m,

119

MINNICH: TREES OF BAJA CALIFORNIA

1987]

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= V ‘Mmjuowudad ‘gq JO SUOTIOT[OO [voTUL1OG = x ‘sydeisojoYd [else
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120 MADRONO [Vol. 34

except in the arid southern tablelands. The largest forests follow
Arroyo Santo Domingo and Rio San Rafael that drain the high Sierra
San Pedro Martir.

Botanical collections show that P. fremontii, the only tree to span
the Baja California peninsula (Wiggins 1980), is occasional in desert
canyons of both the Sierra Juarez and Sierra San Pedro Martir,
including Arroyo El Cajon, Canada El Diablito, and Arroyo El Tajo.
Platanus racemosa occurs mostly below 1000 m on the Pacific slope
of the Sierra Juarez and western foothills of the Sierra San Pedro
Martir, and south to Arroyo El Socorro.

POPULUS TREMULOIDES Michx. (Fig. 13). Numerous groves of
quaking aspen occur in the Sierra San Pedro Martir above 2300 m
along watercourses, on edges of meadows, and near springs. Most
trees are small (<10 m), but some reach 25 m at permanent wet
sites. Only the largest groves could be mapped and are recognized
by their deciduous habit and compact grove structure on aerial pho-
tographs. These include stands near Cerro Venado Blanco, along the
east ridge from the observatory to east of La Encantada Meadow,
and margins of Vallecitos Meadow. On the eastern scarp, groves
occur on the headwalls just northeast of the observatory and the
upper north face of Picacho del Diablo. The nearest stands north of
Baja California are two groves at Fish Creek and Gocke Valley, 350
km north in the eastern San Bernardino Mountains (Griffin and
Critchfield 1976).

POPULUS TRICHOCARPA T. & G. (Fig. 13). Black cottonwood occurs
at only two localities in the Sierra San Pedro Martir: along Arroyo
la Grulla (1400 m), 4 km southwest of La Grulla Meadow, and along
Rio San Rafael (1325 m).

QUERCUS AGRIFOLIA Neé (Fig. 14). Coast live oak, the most wide-
spread hardwood tree in northern Baja California, grows mostly near
stream banks, on meadow perimeters, and on basin floors within
the chaparral belt. It occasionally grows on north exposures, espe-
cially near the international border.

Widespread stands in adjacent San Diego Co. (Griffin and Critch-
field 1976) continue south along the mesic coastal flank of the interior
Sierra Juarez below 1300 m. Quercus agrifolia is particularly abun-
dant between Tecate and Neji. Stand frequency decreases southward
along the range except for large gallery forests along arroyos Barbon
and El Ranchito. It drops out at Canada El Piquillo, avoiding the
arid southern Sierra Juarez tablelands. Quercus agrifolia is occa-
sional in arid interior valleys north of Valle Ojos Negros.

In the coastal Sierra Juarez, QO. agrifolia is abundant in all the
subranges from Canon La Presa to Ensenada Bay and Valle Santo
Tomas and continues southward along the near-coast foothills to
Canon Santa Cruz. At Valle Santo Tomas, scattered populations
swing inland along the arroyos of the transverse coastal ranges par-

MINNICH: TREES OF BAJA CALIFORNIA 21

1987]

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122 MADRONO [Vol. 34

alleling the Agua Blanca fault to the east end of Canon Dolores (cf.
Orcutt 1887).

A single grove of Q. agrifolia at Los Encinos is the only population
within a 25 km span between coastal Sierra Juarez forests and those
at Canon El Carrizo at the northwest end of the Sierra San Pedro
Martir. It is common along the major arroyos on the mesic western
face of the high Sierra San Pedro Martir from 1200-1700 m. The
southernmost locality is a single grove on a stream bank near the
coast along Arroyo Santo Domingo.

QUERCUS CHRYSOLEPIS Leibm. (Fig. 15). Most stands of Quercus
chrysolepis consist of small-leaved shrubs to small trees (<8 m) (Myatt
1975) that grow on steep, well-drained slopes and canyons in the
higher sierra above 1500 m. In the interior Sierra Juarez, Q. chry-
solepis occurs mostly in chaparral on the highest peaks of the plateau.
The largest stands concentrate around Rancho San Faustino, north-
west of Laguna Juarez, near Caballo Muerto, and northern exposures
of mesas and peaks in the southern tablelands. It is found at lower
altitudes (1000-1400 m) on northern exposures of several peaks in
the coastal Sierra Juarez (e.g., Cerro Bola, Sierra Blanca, and Cerro
los Pinos).

In the Sierra San Pedro Martir, outposts occur on Cerro San Matias
(2100 m) and adjacent peaks at the north end of the range. A small
grove of QO. chrysolepis is found on the north slope of Cerro Blanco
(1900 m) near Mike’s Sky Ranch. It is widespread in the highest
part of the range above 1900 m, mostly as understory to P. quad-
rifolia forests on the east scarp and mixed conifer forests on the
plateau, with scattered stands locally entering canyons and northern
exposures at the upper margin of the chaparral belt on the west
slope. It grows as a large tree (ca. 15 m) near streams along the
headwaters of Arroyo la Palizada, and most deep canyons within
the eastern scarp north of Picacho del Diablo that includes upper
Canon la Providencia. Quercus chrysolepis is absent from cold Valle-
citos basin, and highest peaks above 2300-2700 m. South of La
Grulla and La Encantada meadows, Q. chrysolepis is restricted to
northern exposures of highest peaks. The southern limit is Cerro
Chato.

QUERCUS ENGELMANNII Greene (Fig. 14). Although numerous pop-
ulations of O. engelmannii have been recorded near the international
border in San Diego Co. (Griffin and Critchfield 1976), only a few
trees have been found in northern Baja California (4 km south of
Tecate). Although the partly winter-deciduous habit helps differ-
entiate this tree from Q. agrifolia, aerial photographs in winter do
not show recognizable populations elsewhere in Baja California.
Scattered trees undoubtedly will be discovered at new localities,
perhaps in the vicinity of Tecate and the Cerro Bola range.

125

TREES OF BAJA CALIFORNIA

MINNICH

1987]

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124 MADRONO [Vol. 34

QUERCUS PENINSULARIS Trel. (Fig. 16). An endemic to inland ranges
of Baja California, Q. peninsularis is closely related to QO. emoryi
(Muller 1965), widespread in northern Mexico, Arizona, and New
Mexico. In the interior Sierra Juarez, Q. peninsularis is mostly a
shrub to small tree in association with P. jeffreyi forests. The north-
ernmost stand is near Cerro El Topo. It is occasional with pinyon
forests along the east rim and the arid volcanic tablelands to the
south. Botanical collections show that locally it descends arroyos on
the Pacific slope down to 1200 m, avoiding the chaparral.

Quercus peninsularis is infrequent throughout the mixed conifer
forest belt of the Sierra San Pedro Martir; it is particularly abundant
with P. jeffreyi forest at the south end of the range, where it occa-
sionally grows into a robust tree 15 m in height and spread at wet
sites. It is absent from the chaparral belt on the Pacific slope except
along arroyos or forest margins. On the desert scarp, it is a zonal
component of desert chaparral understory of P. monophylla and P.
quadrifolia forests from 1000-2500 m from Canada la Esperanza to
Canon Huatamote. Its abundance perhaps reflects summer rain on
the eastern scarp, as in the area of Q. emoryi in mainland Mexico.
The southernmost collection of Q. peninsularis in the Sierra San
Pedro Martir is from Cerro Chato. The tree occurs on the summits
of Sierra San Luis, and Sierra San Borja, its southern limit.

QUERCUS WISLIZENII A. DC. var. FRUTESCENS Engelm. (Fig. 16).
Field reconnaissances and botanical collections show that interior
live oak is rare in northern Baja California and was not listed by
Wiggins (1980). All known stands, usually a few individuals, occur
at the conifer forest-chaparral ecotone (1200-1700 m) in the wettest
parts of the Sierra. In the Sierra Juarez, it was seen or collected on
Sierra Blanca and above Laguna Juarez. In the Sierra San Pedro
Martir, it was found on a steep northern exposure near Arroyo Los
Pinos, at Arroyo La Corona, southeast of Oak Pasture, and in Q.
agrifolia woodland below the Parque Nacional entrance. Quercus
wislizenii is unusually large (to 10 m) at the latter site, confirming
Brandegee’s (1893) observation that it formed large bushes in the
Sierra San Pedro Martir. Further botanical collecting should expand
the known range of this tree in northern Baja California.

OTHER SPECIES

Three tree species known primarily from California occur south
of the border only in central Baja California. Quercus tomentella
Engelm. occurs in an arroyo 3 km east of Mt. Augusta along the
coast and on Guadalupe Island (Wiggins 1980); Prunus lyonii (Eastw.)
Sarg. is found in the Sierra San Francisco northwest of San Ignacio.
It appears that both trees survive at these localities in part through
isolation from Quercus chrysolepis and Prunus ilicifolia (Nutt.) Walp.,

1987] MINNICH: TREES OF BAJA CALIFORNIA 125

with which they freely hybridize (Muller 1965; Everett 1957). Pinus
radiata D. Don grows in the summer stratus fog-drip zone of Cedros
and Guadalupe Islands (700-900 m) (Critchfield and Little 1966).

NEAR MISSES

Several California trees have southern limits in the Cuyamaca
Mountains within 50 km of the international border, including Acer
macrophyllum Pursh, A. negundo L. subsp. californicum (T. & G.)
Wesmael, Alnus rhombifolia Nutt., Cornus nuttallii Aud., Pinus pon-
derosa Dougl. ex P. & C. Lawson, Quercus kelloggii Newb., and
Umbellularia californica (H. & A.) Nutt. (Griffin and Critchfield
1976). Although the biogeography of organisms seems to be influ-
enced by the intensity of collecting and field surveys, the rapid de-
crease in forest diversity at the border may be no coincidence because
of the strong precipitation gradients associated with relief. The Cu-
yamaca Mountains have a steep western face and no upwind rain-
shadows toward the Pacific Ocean. Precipitation from winter cy-
clonic storms concentrates along a narrow zone at the crest of the
range, where annual amounts approach 1000 mm (California 1980).
In the Sierra Juarez, orographic lift of rain-bearing air masses is
spread over a wider area along gentle west-facing slopes. Rainshad-
ows extend over the range from the coastal Sierra Juarez. Favorable
orography on the steep west face of the Sierra San Pedro Martir is
compensated by decreased winter storm activity southward. Thus,
few areas in northern Baja California have more than 500 mm annual
precipitation, or half the amount in the Cuyamaca Mountains. In
southern California, nearly all trees with southern limits in San Diego
Co. grow in mesic habitats compared to trees with ranges extending
into Baja California.

Several California trees have been erroneously reported in Baja
California as a result of the misidentification of specimens and mis-
interpretation of common plant names and place names. Reports
of Pinus edulis Engelm. in the Sierra Juarez and P. cembroides Zucc.
in the southern Sierra San Pedro Martir (Wiggins 1980) appear to
be based on collections of P. quadrifolia. According to Moran, claims
by rangers that P. ponderosa also occurs in the central Sierra Juarez
are based on invalid taxonomic criteria (see also Dufheld and Cum-
ming, 1949; Wiggins, 1980). The only evidence for Umbellularia
californica (cf. Wiggins 1980) appears to be a ranch named Tres
Laureles, 5 km south of Tecate. References to Pseudotsuga macro-
carpa (Vasey) Mayr in northern Baja California may have resulted
from descriptions of “‘spruce”’ in the Sierra San Pedro Martir or from
confusion of a valley named San Felipe in Baja California with one
in San Diego County (Minnich 1982). The reports of Arbutus men-
ziesli Pursh (Wiggins 1980) are doubtful. Perhaps they are based on

126 MADRONO [Vol. 34

18th century diaries by Arrillaga (Tiscareno 1969) and Longinos-
Martinez (Simpson 1938) who use the name madrono in areas where
Arctostaphylos spp. now occur, including hillsides above La Encan-
tada (Longinos-Martinez), near San Pablo, west of Santa Catarina,
ca. 5 km north of Valle La Trinidad, La Encantada, southern Valle
San Rafael and several locations on the Sierra Juarez plateau (Arri-
llaga). Spanish diarists also may have confused the more rare Het-
eromeles arbutifolia with the Mediterranean Arbutus unedo, which
is still called madrono in Spain. Likewise, references to ‘“‘alder’’
(Alnus rhombifolia) by these diarists, as well as Serra (Tibesar 1955)
and Crespi (Bolton 1927), appears to be a mistranslation for Platanus
racemosa, which also was called “‘aliso”. Thus, 18th century diarists
probably saw sycamores when they traveled through northern Baja
California.

ACKNOWLEDGMENTS

I am most grateful to Reid Moran, William B. Critchfield, and Frank C. Vasek for
their careful reviews of the manuscript. Appreciation also is given to Paulino Rojas-
Gomez for enlightening trips to closed-cone forests near Ensenada.

LITERATURE CITED

ALVAREZ, M. 1981. Aspectos Climaticos del Observatario Astronomico Nacional:
Reporte del Ano de 1969. Instituto de Astronomica. Universidad Nacional Au-
tonoma de México. Reporte No. 5.

AXELROD, D. I. 1980. History of the maritime closed-cone pines, Alta and Baja
California. Univ. California Publ. Geology, Vol. 20. Berkeley.

BoLTon, H. E. 1927. Fray Juan Crespi, missionary explorer on the Pacific Coast
1769-1774. Univ. California Press, Berkeley.

BRANDEGEE, T.S. 1893. Southern extension of the California flora. Zoe 4:199-—210.

BRODER, R. E. 1963. A flora of the Paipai Indian Reservation, Sierra Juarez, Baja
California, México. M.S. thesis, Univ. California, Santa Barbara.

CALIFORNIA, Department of Water Resources. 1980. Monthly total precipitation,
1849-1980. The Resources Division, Sacramento, CA.

CLYDE, N. 1975. El Picacho del Diablo: the conquest of Lower California’s highest
peak, 1932 and 1937. Dawson’s Book Shop, Los Angeles.

CRITCHFIELD, W. B. and E. L. LITTLE, JR. 1966. Geographic distribution of the pines
of the World. U.S.D.A. Forest Serv. Misc. Publ. 991.

DUFFIELD, J. W. and W. C. CUMMING. 1949. Does Pinus ponderosa occur in Baja
California? Madrono 10:22-24.

EvereTT, P.C. 1957. A summary of the culture of California plants at the Rancho
Santa Ana Botanic Garden, 1927-1950. Rancho Santa Ana Botanic Garden,
Claremont, CA.

Fowe.is, H. A. 1965. Silvics of forest trees of the United States. U.S.D.A. Agri.
Hdb. 272.

GASTIL, R. G., R. P. PHILLips, and E. C. ALLISON. 1975. Reconnaissance geology
of the State of Baja California. The Geological Society of America, Inc. Memoir
140.

GRIFFIN, J. R. and W. B. CRITCHFIELD. 1976. The distribution of forest trees in
California. U.S.D.A. Forest Serv. Pacific Southw. Forest and Range Exp. Sta.
Res. Pap. PSW-82.

HELLER, R. C., G. E. DoVERSPIKE, and R. C. ALDRICH. 1964. Identification of tree

1987] MINNICH: TREES OF BAJA CALIFORNIA 127

species on large scale panchromatic and color aerial photographs. U.S.D.A. Forest
Serv. Agri. Hdb. No. 261.

LANNER, R. M. 1974. A new pine from Baja California and the hybrid origin of
Pinus quadrifolia. Southw. Naturalist 19:75-95.

LitT_e, E. L., JR. 1971. Atlas of United States trees. Vol. I, Conifers and important
hardwoods. U.S.D.A. Misc. Publ. 1146.

Mexico, Secretaria de Agricultura y Recursos Hidraulicos. n.d. Manuscript cli-
matological data. Division hydrometrica. Ensenada, Baja California.

MINNICcH, R. A. 1982. Pseudotsuga macrocarpa in Baja California? Madrono 29:
22-31.

. 1986a. Snow levels and amounts in the mountains of southern California.

Journal of Hydrology 89:37-58.

1986b. Range extensions and corrections for Pinus jeffreyi and P. coulteri
(Pinaceae) in northern Baja California. Madrono 33:144—146.

Mooney, H. A. 1977. Southern coastal scrub. Jn M. G. Barbour and J. Major, eds.,
Terrestrial vegetation of California, p. 471-490. Wiley-Interscience, New York.

Moran, R. 1972. Plant notes from the Sierra Juarez of Baja California, Mexico.
Phytologia 35:205-214.

MuLter, C. H. 1967. Relictual origins of insular endemics in Quercus. In R. N.
Philbrick, ed., Proceedings of the Symposium on the Biology of the California
Islands. Santa Barbara Botanic Garden, Santa Barbara, CA.

Myatt, R. G. 1975. Geographic and ecological variation in Quercus chrysolepis
Liebm. Ph.D. dissertation, Univ. California, Davis.

Orcutt, C. R. 1887. The oaks of southern and Baja California. W. Amer. Sci. 3:
135-139.

Simpson, L. B. 1938. Californiain 1792. The expedition of Jose Longinos- Martinez
Transl. The Huntington Library, San Marino, California.

TIBESAR, A. 1955. Writings of Junipero Serra. Academy of American Franciscan
History, Washington, DC.

TISCARENO, F. 1969. José Joaquin Arrillaga. Diary of his Surveys of the Frontier,
1796. Transl. Dawson’s Book Shop, Los Angeles.

VASEK, F.C. 1985. Southern California white fir. Madrono 32:65-77.

VoGL, R. J., W. P. ARMSTRONG, K. L. WHITE, and K. L. Cote. 1977. The closed-
cone pines and cypresses. Jn M. G. Barbour and J. Major, eds., Terrestrial
vegetation of California, p. 295-358. Wiley-Interscience, New York.

WiaaIns, I. 1980. Flora of Baja California. Stanford Univ. Press.

WoLF, C. B. 1948. Taxonomic and distributional studies of New World cypresses.
Aliso 1:1—250.

(Received 6 Jun 1986; revision accepted 24 Dec 1986.)

FIRE HISTORY OF AN OLD-GROWTH FOREST OF
SEQUOIA SEMPERVIRENS (TAXODIACEAE)
FOREST IN HUMBOLDT REDWOODS
STATE PARK, CALIFORNIA

JOHN D. STUART
Department of Forestry, Humboldt State University,
Arcata, CA 95521

ABSTRACT

Establishment dates of basal sprouts of Sequoia sempervirens were found to be
reliable estimates of known fires in Redwood National Park and Prairie Creek State
Park in Humboldt Co., California. Fire history of an old-growth S. sempervirens
forest in the Bull Creek watershed in Humboldt Redwoods State Park, northwestern
California, was determined by analyzing S. sempervirens basal sprouts, fire scars, and
dates of establishment of Pseudotsuga menziesii and Abies grandis. Fire frequency
was estimated for watershed zones, for a clearcut area having fire scars, and for the
entire study area. Conservative and liberal estimates of the fire cycle were made. Pre-
settlement fire intervals were: 24.6 yr for watershed zones, 31 yr for the area with
fire scars, and 13.3 yr for the entire study area. The conservative estimate of the fire
cycle was 51.6 yr and the liberal estimate was 26.2 yr. Settlement fire intervals were:
15.6 yr for watershed zones and 7.5 yr for the entire study area. The conservative
and liberal estimates of the fire cycle were 16 and 10 yr. Post-settlement fire intervals
were: 7.8 yr for the watershed zones, 14 yr for the area with fire scars, and 4.5 yr for
the entire study area. The conservative and liberal estimates of the fire cycle were 16
yr and 9.5 yr. Statistically significant differences (p < 0.05) were found between the
fire interval means for all three settlement periods. No statistically significant differ-
ences (p > 0.05) were found between settlement period and fire size. Fire size was
not correlated with fire frequency.

Throughout the 800 km range of Sequoia sempervirens there is
abundant evidence of fire. In nearly all groves, trees exhibit fire scars,
hollowed-out bases (goose pens), and/or bark char that extend many
meters up the bole. The role of fire in S. sempervirens has been
discussed (Fisher 1903, Fritz 1931, Stone 1966, Veirs 1982). One
approach to deduce fire’s natural role has been to determine fire
history by aging fire scars on stumps. Near the southern end of the
range of S. sempervirens, mean fire intervals of approximately 50
years (Greenlee 1983), and 22-27 yr (Jacobs et al. 1985) have been
reported. Fire intervals near the northern end of the range have been
reported to vary from 50-500 yr, that increase along a continuously
mesic east to west gradient (Veirs 1982). Fritz (1931) concluded that
on a 12 ha area, to the east of Weott, California, there was an average
of 4 major fires per century over the past 1100 yr.

These studies provided good estimates of historic fire frequency,
but did not establish exact calendar dates for fires and did not in-

MADRONO, Vol. 34, No. 2, pp. 128-141, 1987

1987] STUART: FIRE AND SEQUOIA SEMPERVIRENS 129

dicate what size the fires may have attained. The determination of
exact calendar dates is difficult because of 1) weathering of the outer
rind of sapwood on old S. sempervirens stumps that obliterates the
most recent annual rings, 2) the difficulty in determining when trees
were cut, and 3) the production of discontinuous rings or absence
of rings at stump height for large trees during drought or stress
periods (Fritz 1931, LaMarche and Wallace 1972). An alternate
method of determining fire frequency is to date S. sempervirens basal
sprouts that presumably developed following fire. By using basal
sprouts, the first two problems are eliminated and the latter problem
would be minimized because the sprouts would be younger, more
vigorous trees. Therefore, they would be less likely to have discon-
tinuous or missing rings (LaMarche and Wallace 1972). Missing
rings, however, are possible on young, vigorous sprouts if adjacent
sprouts have grown together. Fritz (in Douglass 1928) observed that
the interior radii of joined stump sprouts had fewer annual rings
than did the exterior radii or those radii above the junction of the
sprouts. This phenomenon can be avoided by sampling solitary
sprouts. Lack of certainty that a sprout developed following fire is
another possible problem in using basal sprouts as an indicator of
historic fires. Although it has been well established that S. semper-
virens sprouts following fire (Fritz 1931, Wiant and Powers 1967,
Daubenmire 1975, and many others), it also sprouts following me-
chanical injury to its base (Fisher 1903, Wiant and Powers 1967,
Lindquist 1979, and many others). When there is a ring of basal
sprouts around a fire-damaged old-growth parent tree, it can be
assumed that the sprouts developed because of fire.

The present study was undertaken to 1) verify that basal sprouts
can be used to determine fire frequency; 2) determine the historic
fire frequency in an old-growth S. sempervirens forest by dating basal
sprouts and fire scars on stumps of S. sempervirens, and by dating
other conifer species (Pseudotsuga menziesii and Abies grandis), and
3) estimate the area burned by each fire so that a fire cycle can be
calculated.

FIRE RECORDS

Pre-1940 agency fire records are poor (Wallis et al. 1963). Archival
fire records of the California Department of Forestry are incomplete
for the 1920’s and 1930’s and for some years no records exist. There
are no pre-1920 fire records on file. Gripp (1976) reviewed exten-
sively the northwestern California newspapers and various other
documents and found that large fires in Humboldt and Del Norte
cos. were common. He concluded that between 1880-1939 the mean
interval between severe fire seasons was 3.3 + 0.79 (s.e.) yr.

In a study of large fires occurring in Humboldt and Del Norte cos.

130 MADRONO [Vol. 34

To Eureka
60km

/ o S
[ © Oo iy
\ = m4 ) i =
> S = ss 7
\ 2 &€ /
, / KS cS) /
7
7 oo” WA
{ 7 N
| a Weott
/ 7
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Kilometers

Fic. 1. Location of the study area in the Bull Creek watershed, Humboldt Red-
woods State Park, northwestern California.

between 1955-1974, Gripp (1976) found that 89% were associated
with three major synoptic weather systems: the Pacific High (post-
frontal), the Great Basin High, and the Subtropical High Aloft pat-
tern. These weather patterns can be expected to occur 50-55 days
per summer fire season (Hull et al. in Gripp 1976). The greatest
number of days of critical fire weather can be expected to occur in
July, August, and September. Sixty-nine percent of the large fires
(1955-1974) in Humboldt and Del Norte cos. occurred during Au-
gust and September (Gripp 1976).

STUDY SITE

This study was conducted in a ca. 3500 ha old-growth S. sem-
pervirens and Pseudotsuga menziesii forest growing in the Bull Creek
watershed of Humboldt Redwoods State Park, California (Fig. 1).
The forest includes the Rockefeller Forest that occurs 35 km east of
the coast and experiences none of the moderating influence of the
Pacific Ocean on summer temperature and humidity to the same
extent as do the coastal S. sempervirens groves to the north. Inland
sites typically have summer temperatures between 20-—30°C and rel-
ative humidities between 40-50%, whereas coastal sites experience

1987] STUART: FIRE AND SEQUOIA SEMPERVIRENS 131

summer temperatures of 15—18°C and relative humidities between
80-90% (Horn 1966, Azevedo and Morgan 1974, Elford and
McDonough 1974). Summer fog flowing up the Eel River Valley
often drifts into the Bull Creek watershed. Fog, however, occurs less
frequently in the Rockefeller Forest than it does in S. sempervirens
groves along the Eel River or closer to the ocean (Waring and Major
1964, Azevedo and Morgan 1974). As a result, the Bull Creek drain-
age experiences greater diurnal and annual temperature extremes.
Summer droughts are common with nearly all of the 760—2500 mm
of precipitation falling as rain from October to April.

The watershed’s topography is varied with slopes from 0% to more
than 50%. Elevations range from 50 m at the mouth of Bull Creek
to roughly 1030 m. Potential fire hazard is higher in the Bull Creek
watershed than in coastal S. sempervirens because of the steep slopes
and relatively severe summer fire weather.

Nearly pure stands of S. sempervirens are found along the alluvial
flats of Bull Creek. Occasional overstory associates include Abies
grandis and P. menziesii. Understory associates include Lithocarpus
densiflorus, Umbellularia californica, Vaccinium ovatum, Gaultheria
shallon, Polystichum munitum, and Oxalis oregana.

On the slopes above the alluvial flats, the density and basal area
of S. sempervirens declines with increasing elevation, although this
species dominates (greatest basal area) nearly everywhere it is found.
At lower elevations, S. sempervirens 1s associated with P. menziesii,
A. grandis, L. densiflorus, A. menziesii, and U. californica. On rocky
sites or near prairies, Quercus garryana and Q. kelloggii are found.
At higher elevations, A. grandis, U. californica, Q. garryana, and Q.
kelloggii are absent or rare, whereas L. densiflorus and A. menziesii
increase in density and basal area. Lithocarpus densiflorus saplings
and V. ovatum thickets are found in the understory at all slope
elevations.

LAND USE HISTORY

Pre-settlement period (pre-1875). Pre-settlement fires in the Bull
Creek watershed probably were caused mostly by Indians. Lightning
activity in this part of the range of S. sempervirens is relatively low,
but it is often accompanied by rainfall when it does occur (Fritz
1931). The Sinkyone Indians were the primary inhabitants of this
region. They migrated through the watershed on their way from the
South Fork of the Eel River to the Pacific Ocean and had established
villages in and near the watershed. Regular burning by the Sinkyones
has been reported as a means to drive out insects and rodents for
food and to keep the forest understory open for travel (Fritz 1931,
Gilligan 1966).

132 MADRONO [Vol. 34

Settlement period (1875-1897). European man arrived in the vi-
cinity of Bull Creek in 1848 when four members of Dr. Josiah Gregg’s
expedition from Weaverville to San Francisco proceeded up the
South Fork of the Eel River. It was not until the early 1870’s, how-
ever, that settlement occurred. One of the earliest settlers was To-
saldo Johnson who grazed sheep and cattle on prairies in what is
now known as the Rockefeller Forest. By 1895, most of the Bull
Creek watershed had been claimed under provisions of the Home-
stead Act. Fire was used by early settlers for the maintenance and
enlargement of pastures and for land clearing (Gilligan 1966). Many
fires escaped into the forests because of the lack of organized fire
suppression.

Post-settlement period (1898-1940). The major land use activities
from 1895-1945 were livestock grazing, farming, debarking of L.
densiflorus for tannin production, and logging of P. menziesii and
S. sempervirens. Broadcast burning was used regularly during this
period to maintain pastureland (Gilligan 1966), and to facilitate
logging activities. Logged areas were burned commonly prior to log
skidding to reduce the impediments of logging debris and understory
vegetation (Fritz 1931). Many of these fires burned into old-growth
S. sempervirens stands, but apparently were extinguished naturally
because of a combination of lower temperatures, higher relative
humidities, and high fuel moisture contents. In years when the fuels
were especially dry and the weather hot or windy, however, fire
readily spread through the S. sempervirens forest. For example, in
1936 a fire spread southward from a broadcast burn on property of
the Pacific Lumber Company into an old-growth S. sempervirens
forest in the Bull Creek watershed (unpubl. fire report, Calif. Dept.
of Forestry 1936). In 1936, there were many other fires burning in
old-growth S. sempervirens forests on the north coast of California.
Although fire archives of the California Department of Forestry
probably do not contain all of the fire reports for 1936, the existing
reports indicate that there were fires that covered from 86—4856 ha
in areas of old-growth S. sempervirens in Humboldt and Del Norte
cos. All of these fires originated from escaped broadcast burns or
other incendiary activities.

After 1945, land use patterns changed with the implementation
of the State Forest Practices Act and with more vigorous fire suppres-
sion. As a result, the number of escaped fires from agricultural or
logging activities has been reduced greatly.

METHODS

The test to verify that basal sprouts can be used to determine fire
frequency was conducted in northern Humboldt Co., 130 km north

1987] STUART: FIRE AND SEQUOIA SEMPERVIRENS 133

of the main study area. All other data were collected in the Bull
Creek watershed, Humboldt Redwoods State Park.

Basal sprout ages and fire frequency. Establishment dates of basal
sprouts were determined in three areas of old-growth S. sempervirens
that are known to have experienced fires. One of the areas burned
in October 1974 along a tributary of Redwood Creek in Redwood
National Park (RNP), and the other two areas burned during a fire
of late September 1936 near the northern boundary of Prairie Creek
State Park. The exact location of the 1974 fire was determined from
records on file at RNP and by eyewitness accounts (Veirs, pers.
comm. 1983). Locations of the two areas in the 1936 fire were
determined using a fire report (unpubl. fire report, Calif. Dept. of
Forestry 1936), and re-establishing photo points used in the taking
of photographs following the fire (photos on file at RNP). A total of
20 basal sprouts from different parent trees were dated from the
three areas: 10 from the 1974 fire, and five from each of the two
1936 fire areas. Basal sprouts were sampled next to trees that had
bark char or fire scarring, and those sprouts that had no external
evidence of fire. Increment corings were extracted at a height of 15
cm, cross sections were removed approximately 10 cm above ground
level, and both were sanded and examined under a microscope. A
correction factor of one year was added to the dates determined by
counting annual rings.

Fire history study. Fifty-nine sample points were established in a
0.80 km? grid pattern in the 3500 ha old-growth S. sempervirens
and P. menziesii forest that occurs in the eastern portion of the Bull
Creek watershed. All trees of S. sempervirens within a 150 m radius
of the sample point were examined for basal sprouts and evidence
of past fires. Any sprout whose parent had a fire scar or bark char
was assumed to have resulted from a fire. There were often two and
three generations of basal sprouts evident from one parent tree (Fig.
2). The number of sprouts age class"! parent tree”! was variable, but
usually was only one or two. Increment cores were extracted from
basal sprouts of all apparent age classes as close to the base of their
stems as possible, although usually within 25 cm of ground level.
The cores were mounted in permanent holders, sanded, and ex-
amined under a microscope.

Dates of sprout establishment were determined by summing the
number of years counted on the extracted increment cores to the
mean annual height growth of basal sprouts. Mean basal sprout
height growth for the first year following three prescribed burns near
Look Prairie was 37 + 11.7 (s.e.) cm (Stuart 1986). Crossdating was
attempted, but was found to be ineffective primarily because of
strong competitive interactions and variable radial growth patterns.

134 MADRONO [Vol. 34

Fic. 2. Two generations of Sequoia sempervirens basal sprouts. Parent tree has a
fire scar at its base, and charred bark extending approximately 10 m up its stem.

1987] STUART: FIRE AND SEQUOIA SEMPERVIRENS 135

Studies in S. sempervirens by Douglass (in Schulman 1940), Schul-
man (1940), and LaMarche and Wallace (1972) have found similar
difficulties in crossdating S. sempervirens cross sections. Fire dates
were recognized if there was synchrony among basal sprout dates.
To help substantiate the basal sprout fire dates, increment corings
were extracted from adjacent P. menziesii and A. grandis. It was
assumed that these species became established following the liber-
ation of growing space by a fire. A similar pattern was observed by
Veirs (1982) in S. sempervirens stands that occur 150 km to the
north of the study site, where P. menziesii and Tsuga heterophylla
became established after ground fires.

Additional fire dates were obtained by dating fire scars on stumps
found in an old clearcut. All but one of the several clearcuts that
were scattered throughout the forest were cut too long ago and yield-
ed rotten, unusable stumps. A 35-year-old clearcut unit on a north-
west aspect, however, was found whose stumps were sound enough
to be used. Following a reconnaissance, five stumps were found
showing two or more fire scars. Crossdating was performed for those
stumps with uninterpretable sapwood rings using fire intervals es-
tablished with S. sempervirens basal sprouts.

The original objective of determining exact historic fire boundaries
could not be achieved because equal-aged basal sprouts from indi-
vidual trees were dispersed too widely. Some reasons for this were:
1) the density of S. sempervirens on slopes was relatively low (10-
40/ha); 2) not every S. sempervirens sprouted following fire, leading
to highly dispersed equal-aged basal sprouts; and 3) some basal
sprouts were killed by subsequent fires, which led to even greater
dispersion of equal-aged basal sprouts. In spite of these factors, two
estimates of fire size were attempted based on aspect and position
within the Bull Creek watershed. Eleven zones were delimited (Table
1) to represent areas of similar aspect and watershed position. The
first estimate of fire size was a liberal one based on the assumption
that an entire watershed zone burned if there was any evidence of
fire in that zone. Fire size was then estimated by summing the areas
of those fire affected zones. Although this technique overestimates
fire size for small local fires, it is probably realistic for many pre-
settlement and unsuppressed fires. Those fires likely burned for weeks
at a time, especially during drought periods, and were extensive.
Many of the large 1936 fires, for example, in S. sempervirens forests
on the north coast of California, burned for 2—3 weeks even with
fire suppression (unpubl. fire reports, Calif. Dept. of Forestry 1936).
The second estimate of fire size was more conservative. I assumed
that, for a watershed zone to burn completely, it must have had
more than one plot with evidence of fire. For those watershed zones
having only one plot with evidence of fire, I assumed that the fire

136 MADRONO [Vol. 34

TABLE 1. ZONES REPRESENTING AREAS OF SIMILAR ASPECT AND WATERSHED PoOsI-
TION WITHIN THE OLD-GROWTH PORTION OF THE BULL CREEK DRAINAGE. Mean pre-
settlement fire intervals for each zone are presented (data are x + s.e.).

Fire zones Area (ha) Aspect Mean fire interval
Harper Creek 368 S 21.8 (6.7)
Calf Creek 337 S 31.0 (12.1)
Western Cow Creek 397 SE 23.4 (11.6)
Eastern Cow Creek 433 SW, W 34.8 (8.4)
Tepee Creek 286 NE 19.5 (4.9)
Connick Creek 190 N 43.7 (14.2)
Miller Creek 223 N 36.3 (25.3)
Lower Eastern Squaw Creek 261 NW 11.3 (6.7)
Upper Eastern Squaw Creek 410 NW, W 21.0 (5.1)
Lower Western Squaw Creek 257 NE, N 18.2 (3.3)
Upper Western Squaw Creek 348 NE, E 21.8 (9.2)
3510 24.6 (2.8)

burned %, of the entire study area, 1.e., 3500 ha/59 plots = 59.3
ha. Once fire size was estimated, I calculated conservative and liberal
estimates of the fire cycle for the post-settlement (1898-1940), set-
tlement (1875-1897), and pre-settlement periods (pre-1875). The
fire cycle is defined as the length of time necessary for an area equal
to the entire area of interest to burn (Romme 1980).

RESULTS

Basal sprout ages and fire frequency. Dates of basal sprout estab-
lishment in the three sampling areas verified the known fire dates.
In the area of the late season 1974 fire, nine out of 10 basal sprouts
were determined to have been established in 1975, with the other
basal sprout dated at 1976. Similar results were found in the two
sampling areas in the area of the late season 1936 fire. Eight out of
10 basal sprouts were dated to 1937; one was dated to 1938; and
one was found to have been established in 1940.

Fire history study. Fire frequencies for the pre-settlement, settle-
ment, and post-setthement periods were calculated for individual
watershed zones, the entire study area, and for the 35-year-old clear-
cut (fire scars only). The pre-setthement mean fire interval for the
11 watershed zones ranged from 11.3 + 6.7 (s.e.) yr to 43.7 + 14.2
(s.e.) yr (Table 1). There were no statistically significant differences
(p > 0.05) between the pre-settlement mean fire intervals of the
watershed zones. The average of the mean pre-settlement fire inter-
vals of the 11 watershed zones was 24.6 + 2.8 (s.e.) yr. Shorter mean
fire intervals were found for the settlement [15.6 + 1.5 (s.e.) yr] and
the post-settlement [7.8 + 0.6 (s.e.) yr] periods. No statistically sig-
nificant differences (p > 0.05) were found between the mean fire

1987] STUART: FIRE AND SEQUOIA SEMPERVIRENS 137

TABLE 2. FIRE DATES AND FIRE INTERVALS DETERMINED FROM Sequoia semper-
virens BASAL SPROUTS, CONIFER (Pseudotsuga menziesii AND Abies grandis) EstA-
BLISHMENT DATES, AND FIRE SCARS IN THE OLD-GROWTH PORTION OF THE BULL CREEK
DRAINAGE.

No. of | No.of No. of Fire

Fire fire basal conifers interval
year scars sprouts est. (yr)
Post-settlement period 1940 2 3 1 4
1936 — 11 2 6
1930 — 8 — 3
1927 _ 4 1 4
1923 l 5 — 4
1919 — 13 6 6
1913 1 3 l 4
1909 — 6 4 5
1904 — 8 5 6
1898 2 6 1 3
Settlement period 1895 — 4 2 7
1888 — 15 4 5
1883 — 6 3 8
1875 —_ 4 5 10
Pre-settlement period 1865 l 6 4 8
1857 — i 5 11
1846 — 5 6 11
1835 4 5 4 9
1826 — 3 3 9
1817 _ 4 l 15
1802 1 7, 5 18
1784 _~ 5 — 20
1764 2 — _ 16
1748 — 5 1 9
1741 2 2 — 15
1726 — 6 — 19
Incomplete data 1707 2 — — 12
1695 — 2 — 9
1686 — 2 — 38
1648 — 2 — 37
1611 _ 2 — 13
1598 1 l — 12
1586 _ 1 — 17
1569 l — — 63
1506 _— 1 — 11
1495 _ 1 — 26
1469 — 1 — 139
1330 l _ — —

intervals of the eleven watershed zones for either the settlement or
post-settlement periods.

Considerable variability was found in fire intervals within wa-
tershed zones and between the fire intervals of all watershed zones.

138 MADRONO [Vol. 34

Pre-settlement fire intervals varied from 8-87 yr. The coefficients
of variation for pre-settlement fire intervals for individual watershed
zones ranged from 36—120% and for the means of fire intervals of
all watershed zones it was 37.8%.

Fire intervals calculated for the entire study area in each fire year
and averaged for the three time periods are shown in Table 2. The
post-settlement period had the lowest mean fire return interval [4.5 +
0.6 (s.e.) yr], with the settlement period, and pre-settlement periods
having longer intervals [7.5 + 0.8 (s.e.) yr, and 13.3 + 1.2 (s.e.) yr,
respectively]. Statistically significant differences (p < 0.05) were found
between the fire interval means for all three time periods.

The fire intervals calculated from the fire scar data exhibited a
similar trend as the other two estimates, with a longer fire interval
[31 + 3.1 (s.e.) yr] in the pre-setthement period and a shorter fire
interval [14 + 2.0 (s.e.) yr] for the post-settlement period. There
were no fire scars found for the settlement period.

No statistically significant differences (p > 0.05) between time
periods and either the conservative or liberal estimates of fire size
were found. Mean fire size based on the conservative estimate for
the post-settlement period was 918 + 162 (s.e.) ha, for the settlement
period 1097 + 362 (s.e.) ha, and for the pre-settlement period 786 +
139 (s.e.) ha. The mean fire size based on the liberal estimate for
the post-settlement period was 1748 + 121 (s.e.) ha, for the settle-
ment period 2018 + 151 (s.e.) ha, and for the pre-settlement period
1629 + 167 (s.e.) ha. Fire frequency was not correlated with either
the conservative or the liberal estimates of fire size (r = —0.215,
p = 0.100; r= —0.254, p= 0.100, respectively). Fire cycles calculated
using the conservative estimate of fire size were 51.6, 16.0, 16.0 yr
for the pre-settlement, settlement, and post-settlement periods. The
fire cycles using the liberal fire size estimates were 26.2, 10.0, and
9.5 yr for the same periods. A comparison of the five estimates of
fire frequency are presented in Table 3.

DISCUSSION

Fire frequency based on watershed zones, the entire study area,
fire scars, and two estimates based on the fire cycle were all similar.
Each of these methods revealed a pattern of relatively long fire
intervals during the pre-settlement period, shorter fire intervals dur-
ing the settlement period, and still shorter fire intervals during the
post-settlement period. The mean pre-settlement fire intervals for
all the watershed zones (24.6 yr), and for the area with fire scars (31
yr) were similar to those reported by Fritz (1931; ca. 25 yr) and
Jacobs et al. (1985; 22—27 yr). The shorter pre-settlement fire interval
based on the entire study area (13.3 yr) was an artifact of the size
of the reference area. A large sampling area is likely to include more

1987] STUART: FIRE AND SEQUOIA SEMPERVIRENS 139

TABLE 3. ESTIMATES OF FIRE FREQUENCY FOR THE OLD-GROWTH PORTION OF THE
BULL CREEK WATERSHED. Frequency estimates are presented for the post-settlement,
settlement, and pre-settlement periods. Fire interval data are X + S.e.

Post-settlement Settlement Pre-settlement

Fire interval (yr)

Based on entire study area 4.5 (0.6) 7.5 (0.8) 13.3 (1.2)

Based on scars from | site 14.0 (2.0) — 37,0: G.1)

Based on watershed zones 7.8 (0.6) 15.6 (1.5) 24.6 (2.8)
Fire cycle (yr)

Conservative estimate 16.0 16.0 51.6

Liberal estimate 9.5 10.0 26-2

evidence of past fires than a smaller one (Arno and Petersen 1983).
My entire study area, for example, was 3500 ha, whereas the largest
watershed zone was only 433 ha. The studies by Fritz, Veirs (1982),
and Jacobs et al. were conducted in areas less than 100 ha.

Fire frequency estimates based on the fire cycle should be con-
sidered only as estimates because I was unable to reconstruct exact
fire perimeters. The liberal estimate of the fire cycle (26.2 yr) should
be considered the probable minimum time; and the conservative
estimate (51.6 yr) should be considered as the probable maximum
time. It is unlikely that the true fire cycle, especially for the pre-
settlement period, is greater than my conservative estimate because
pre-settlement fires were not suppressed and, therefore, were prob-
ably more extensive.

Pre-settlement fire frequencies were highly variable and no sig-
nificant differences were found between pre-settlement mean fire
intervals among the eleven watershed zones. These phenomena are
probably due to large variances within and between watershed zones
and the small number of intervals in each watershed zone. Ecological
and land management inferences based solely on pre-settlement mean
fire intervals would be simplistic. Any prescribed burning program
designed to recreate pre-European man fire regimes should incor-
porate variable intervals between fires (McBride et al. 1986) within
and between watershed zones.

The longer fire intervals (SO—500 yr) reported by Veirs (1982) can
be attributed to the relatively mild fire weather conditions found in
coastal S. sempervirens forests (Elford and McDonough 1974, Gripp
1976). Differences in fire frequency throughout the range of S. sem-
pervirens are apparently a function of the steep climatic gradient
extending from the cool, moist coastal sites to the relatively warm,
dry inland sites.

ACKNOWLEDGMENTS

This research, College of Natural Resources contribution #85-6, was supported by
the California Department of Parks and Recreation and MclIntire-Stennis grant num-

140 MADRONO [Vol. 34

ber 81. David Boyd, Marla Hastings, Stephen Matthews, Tom Gilmour, and Ray
McCay provided invaluable assistance. I thank Stephen Veirs, Ron Mastrogiuseppe,
and Dale Thornburgh for critical reviews of the manuscript. I am grateful to Stephen
Veirs and Lyman Abbott for providing cross sections of basal sprouts from the 1974
Redwood Creek fire.

LITERATURE CITED

ARNO, S. F. and T. D. PETERSEN. 1983. Variation in estimates of fire intervals: a
closer look at fire history of the Bitterroot National Forest. U.S.D.A. Forest Serv.
Res. Pap. INT-301.

AZEVEDO, J.and D.L. MORGAN. 1974. Fog precipitation in coastal California forests.
Ecology 55:1135-1141.

DAUBENMIRE, R. and J. DAUBENMIRE. 1975. The community status of the coastal
redwood, S. sempervirens. Report to Redwood National Park, U.S. Natl. Park
Serv.

Douatass, A. E. 1928. Climatic cycles and tree growth. Vol. II. A study of the
annual rings of trees in relation to climate and solar activity. Publ. Carnegie Inst.
Wash. 289. Washington, DC.

ELFORD, R. C. and M. R. McDoNouGH. 1974. The climate of Humboldt and Del
Norte Counties. Univ. California, Humboldt and Del Norte Counties, Agri.
Extension Serv. Eureka, CA.

FISHER, R. T. 1903. A study of redwood. Jn The redwood. U.S. Forest Serv. Bull.
38:3-28. Washington, DC.

Fritz, E. 1931. The role of fire in the redwood region. J. Forest. 29:939-950.

GILLIGAN, J. P. 1966. Land use history of the Bull Creek Basin. Jn Proc. Symp.
Mgt. for park preservation: a case study at Bull Creek, Humboldt Redwoods
State Park, California, p. 42-57. School of Forestry, Univ. California, Berkeley.

GREENLEE, J. M. 1983. Vegetation, fire history, and fire potential of Big Basin
Redwoods State Park, California. Ph.D. dissertation, Univ. California, Santa
Cruz.

Gripp, R. A. 1976. An appraisal of critical fire weather in northwestern California.
M.S. thesis, Humboldt State Univ.

Horn, W. L. 1966. Climate and hydrology. /n Proc. Symp. Mgt. for park preser-
vation: a case study at Bull Creek, Humboldt Redwoods State Park, p. 13-31.
School of Forestry, Univ. Calif., Berkeley.

HuLt, M. K., C. A. O’DELL, and M. J. SCHROEDER. 1966. Critical fire weather
patterns—their frequency and levels of fire danger. U.S.D.A. Pacific Southw.
Forest and Range Exp. Sta. Berkeley, CA.

Jacoss, D. F., D. W. CoLe, and J. R. McBribe. 1985. Fire history and perpetuation
of natural coast redwood ecosystems. J. Forest. 83:494-497.

LAMARCHE, V. C. and R. E. WALLACE. 1972. Evaluation of effects on trees of past
movements on the San Andreas Fault, northern California. Geol. Soc. Amer.
Bull. 83:2665-2676.

LINDQuIST, J. L. 1979. Sprout regeneration of young-growth redwood: sampling
methods compared. U.S.D.A. Forest Serv. Res. Note PSW-337.

McBrIDE, J. R., D. F. JAcoss, and D. W. Cote. 1986. Use of fire history data in
planning fire intervals for controlled burning. Jn J. P. Long, ed., Proc. Symp.,
Fire Mgt.: the challenge of protection and use, p. 279-286. Utah State Univ.,
Logan.

RomME, W. 1980. Fire history terminology: report of the ad hoc committee. Jn
Proceedings of the fire history workshop. October 20-24, 1980. Tucson, Arizona.
U.S.D.A. For. Serv. Gen. Tech. Rep. RM-81.

SCHULMAN, E. 1940. Climatic chronology in some coast redwoods. Tree-ring Bull.
6(3):22-23.

STONE, E. C. 1966. Ecology of the watershed. /n Proc. Symp., Mgt. for park pres-

1987] STUART: FIRE AND SEQUOIA SEMPERVIRENS 141

ervation: a case study at Bull Creek, Humboldt Redwoods State Park, p. 65-78.
School of Forestry, Univ. California, Berkeley.

STUART, J. D. 1986. Redwood fire ecology. Final report submitted to the California
Department of Parks and Recreation.

VeEIRS, S. D., JR. 1982. Coast redwood forest: stand dynamics, successional status,
and the role of fire. Jn Proc. Symp., Forest Succession and Stand Development
Research in the Northwest, p. 119-141. Oregon State Univ., Corvallis.

WALLIs, J. R., K. L. Bowben, and J. D. LENT. 1963. Area burned by wildfire in
California watersheds, 1940-1959. U.S.D.A. For. Serv. Res. Note PSW-30.

WARING, R. H. and J. Mayor. 1964. Some vegetation of the California coastal
region in relation to gradients of moisture, nutrients, light, and temperature. Ecol.
Monogr. 34:167-215.

WIANT, H. V., JR. and R. F. Powers. 1967. Sprouting of old-growth redwood. Proc.
Soc. Amer. For. 1966, p. 82-90.

(Received 9 Nov 1985; revision accepted 6 Jan 1987.)

ANNOUNCEMENT

VASCULAR PLANTS OF ARIZONA

Taxonomists interested in the flora of Arizona have convened several times
since summer 1986 to organize a working group to produce a new plant iden-
tification manual for the state. Because Arizona Flora by Kearney and Peebles
is out-of-print, a seminar intended for potential users was hosted by the Her-
barium of the University of Arizona at Tucson, addressing the subject, “‘Re-
vising Arizona Flora: What do you want?” in discussion format.

As a result of the seminar an editorial board, consisting of Frank S. Cross-
white, editor-in-chief, Richard S. Felger, Charles T. Mason, Jr., Donald J.
Pinkava, John R. Reeder, and Rebecca K. Van Devender, has been established.
It will govern all stages of production from planning to actual printing and
publication. Several decisions have been made. The new book will be entitled
Vascular Plants of Arizona and will be consistent in size, shape and complexity
with Gray’s Manual of Botany, although not necessarily consistent in style and
format. The editors wish the book to include the works of the most highly
qualified experts for each taxonomic group and solicit communication with
all interested in contributing. Treatments accepted and published will be ac-
knowledged with an authorship by-line. Guidelines for authors are being pre-
pared and will be available from Rebecca K. Van Devender, Herbarium,
College of Agriculture, University of Arizona, Tucson 85721.

CHROMOSOME RACES OF GRAYIA BRANDEGEI
(CHENOPODIACEAE)

HOWARD C. STUTZ
Department of Botany and Range Science,
Brigham Young University, Provo, UT 84602

STEWART C. SANDERSON and E. DURANT MCARTHUR
Shrub Sciences Laboratory, Intermountain Research Station,
735 North 500 East, Provo, UT 84601

CHU GE-LIN
Institute of Botany, Northwest Teachers College,
Lanchou, Gansu, The People’s Republic of China

ABSTRACT

Diploid (2n = 18) and tetraploid (2n = 36) races of Grayia brandegei differ in
phenotype and geographic distribution. Diploids have narrow, linear leaves and most-
ly are restricted to south-central Utah and northeastern Arizona. Tetraploids are
larger statured, bear larger, more oval leaves and mostly occur in isolated populations
in northeastern Utah, south-central Wyoming, eastern Colorado, and northwestern
New Mexico. Because of their morphological and distributional differences, the tet-
raploid form is designated as a new variety, Grayia brandegei var plummeri.

Grayia brandegei was described by Asa Gray (1876) from speci-
mens obtained by Brandegee on the banks of the San Juan River in
Utah, near the border of Colorado. It differs from the only other
species in the genus, G. spinosa Mogq., in several morphological
features and geographic distribution. Whereas G. spinosa has a wide-
spread distribution throughout western United States, G. brandegei
is limited mostly to the drainage of the Colorado River in eastern
Utah and adjacent states. Their ranges overlap in several places and
appear to be separated primarily by edaphic differences.

Grayia brandegei usually grows on heavy clay soils or sandy loams,
but is not restricted to the Chipeta formation, as suggested by Col-
lotzi (1966). Like most other shrubs of the Chenopodiaceae, except
Atriplex, it lacks Kranz-type leaf anatomy and presumably has the
C, photosynthetic pathway. Contrary to previous reports, it is mon-
oecious rather than dioecious. Leaves appear in early April, before
flowering, and abscise in late summer.

Some populations of G. brandegei contain large robust plants with
broad, oval leaves, whereas others have smaller-statured plants with
narrow, linear leaves. To determine the basis for interpopulational
differences, comparative morphological and cytological studies were

MADRONO, Vol. 34, No. 2, pp. 142-149, 1987

1987]

TABLE 1.

STUTZ ET AL.: GRAYIA BRANDEGEI

143

LOCATIONS OF POPULATIONS OF Grayia brandegei EXAMINED FOR

CHROMOSOME NUMBER AND MORPHOLOGICAL FEATURES. Soil properties were ex-
amined for those populations marked with *.

Diploids

*AZ, Apache Co., 16 km n. of Many
Farms
*CO, Mesa Co., 16 km w. of Mack
*UT, Grand Co., 8 km nw. of Moab
*UT, San Juan Co., 16 km s. of Bland-
ing
UT, San Juan Co., 20 km se. of Hites
Crossing
UT, San Juan Co., 1 km w. of Ismay
Trading Post
*UT, San Juan Co., 6 km n. of Monte-
zuma Creek

Tetraploids

*CO, Garfield Co., 12 km nw. of Rifle
NM, San Juan Co., 10 km n. of Aztec
NM, San Juan Co., 20 km se. of

Bloomfield

*NM, San Juan Co., 8 km e. of La Plata

UT, Emery Co., 16 km e. of Fremont
Jct.

UT, Garfield Co., 1 km e. of Henrie-
ville

*UT, Kane Co., Cottonwood Wash, 1

km s. of Cannonville

*UT, Sanpete Co., 10 km nw. of Sterling

*UT, Uintah Co., 18 km e. of Roosevelt
UT, Wayne Co., 3 km e. of Notom
Exit 24
UT, Wayne Co., 10 km n. of Baker
Ranch, Thousand Lake Mt.
*WY, Carbon Co., 3 km nw. of Baggs

made on plants that grow in each of 19 populations distributed
throughout the range of G. brandegei.

METHODS

Chromosome counts were made from microsporocytes in male
buds taken from plants in natural populations or from mitotic cells
in root tips taken from actively growing seedlings that were ger-
minated on blotter paper in Petri dishes. In each case, the tissues
were fixed in aceto-alcohol (1:3, v/v) and stained in aceto-carmine
for microscopic examination. Numbers were determined for at least
two plants in each of the 19 populations listed in Table 1.

Measurements of plant height and width, and leaf length and width
were made on 20 plants in each population. Because the plants are
mostly iso-diametric, plant volume was determined by plant height
x a(plant diameter/2)”. Plants measured were taken at random at
intervals of six feet along a linear transect through each population.
Each leaf measured was the largest leaf on a randomly selected twig
taken from each plant. Morphological characteristics were compared
between ploidy levels using one-way ANOVA (Table 2). Voucher
specimens from each population are deposited at Brigham Young
University (BRY).

Soil samples were obtained from the site within each population
by mixing together five shovelfuls of soil taken at six-foot intervals
in a randomly selected linear transect. Cations of the soils were

144 MADRONO [Vol. 34

TABLE 2. MEAN AND STANDARD DEVIATIONS OF PLANT STATURE AND LEAF DI-
MENSIONS OF DIPLOID (n = 140) AND TETRAPLOID (n = 240) Forms OF Grayia bran-
degei. Measurements are from 20 plants in each of the populations listed in Table 1.
All values significant at p < 0.0001 except when indicated.

2n 4n F (df = 378)
Plant ht. (cm) 31.23 + 10.14 37.03 + 13.35 19.78
Plant w. (cm) Tie2 28:28 84.46 + 33.59 15.41
Plant vol. (dm?) 127.22 297-21 307.66 + 263.04 13.69
Leaf 1. (mm)"s 34.86 + 10.17 33.87 + 13.04 0.60
Leaf w. (mm) 3.83 + 1.47 913 = 521 138.62
Leaf 1/w ratio 9,59 2 2.27 4.18 + 1.67 708.21

analyzed in a Perkin Elmer 5000 Atomic Absorption Spectropho-
tometer. A saturated water extract was used for the analysis of the
macronutrients Ca, Mg, K, and Na. The micronutrients Cu, Fe, Mn,
and Zn were extracted in DTPA. P content was determined color-
imetrically from a Na,CO, extract. Na absorption ratio (SAR) was
calculated from the values obtained for Na, Ca, and Mg. Cation
exchange capacity (CEC) was determined by the ammonia distilla-
tion method. Texture was determined by the hydrometer method.

RESULTS

Cytological studies showed that plants in the population first col-
lected by Brandegee in southeastern Utah near Ismay Trading Post
were diploid with 2” = 18 chromosomes. Diploid chromosome
counts also were obtained from plants in six other populations (Table
1). In each population, the plants had narrow, linear leaves (Fig. 1,
Table 2). In 12 other populations of G. brandegei, plants were more
robust and had considerably wider leaves (Figs. 1, 2, 3, and Table
2). Those individuals examined were found to be tetraploid with
2n = 36.

Tetraploid populations are more widely distributed than diploid
populations (Fig. 3). Putative diploid populations, in which the plants
are small-statured and bear narrow, linear leaves, are found along
the San Juan River in southeastern Utah. Putative tetraploid pop-
ulations, in which the plants are larger statured and bear wider leaves,
are common in central and northeastern Utah, northwestern New
Mexico and northwestern Colorado.

The only known population of G. brandegei outside the Colorado
River drainage is located west of Sterling, Sanpete County, Utah.
This sizeable tetraploid population is growing on Tertiary, Green
River clays that are locally abundant, but are otherwise rare in the
Great Basin drainage of Utah. This geological formation is common

1987] STUTZ ET AL.: GRAYIA BRANDEGEI 145

tt
) N
YW) Vy
Va
BAN

Fic. 1. Twigs of diploid (a, b) and tetraploid (c, d) Grayia brandegei. a. AZ, Apache
Co., 16 km n. of Many Farms. b. UT, San Juan Co., 6 km n. of Montezuma Creek.
c. WY, Carbon Co., 3 km nw. of Baggs. d. UT, Uintah Co., 18 km e. of Roosevelt.

in the northern portion of the Colorado Plateau, and some of the
tetraploid populations in eastern Utah and western Colorado occur
on it.

All macro- and micronutrients among populations noted in Table
1 showed no significant differences. Similarly pH, CEC, SAR, and
soil texture showed no significant differences.

ANOVA for between-population differences for plant height and
width and leaf length and width of 12 tetraploid populations were
all significant (F-test, p < 0.0001).

DISCUSSION

The distribution pattern of diploid and tetraploid populations of
G. brandegei in Utah, Wyoming, Colorado, Arizona and New Mex-
ico suggests that the tetraploids were derived from the diploids.
Because there are no other species that appear to have been involved
in their origin, tetraploids are probably autotetraploids. Because of
the conspicuous inter-populational differences among them, they
probably arose polyphyletically.

Repeated establishment of autotetraploid populations from dip-
loid progenitors could be best explained as originating via triploids.
Occasional triploids, arising in a diploid population from fertiliza-

146 MADRONO [Vol. 34

Fic. 2. Leaves of diploid (a—f) and tetraploid (g—l) Grayia brandegei. a. UT, Grand
Co., 8 km nw. of Moab. b. UT, San Juan Co., 6 km n. of Montezuma Creek. c. UT,
San Juan Co., 16 km s. of Blanding. d. CO, Mesa Co., 16 km w. of Mack. e. AZ,
Apache Co., 16 km n. of Many Farms. f. UT, San Juan Co., 1 km w. of Ismay Trading
Post. g. UT, Uintah Co., 18 km e. of Roosevelt. h. WY, Carbon Co., 3 km nw. of
Baggs. 1. UT, Sanpete Co., 10 km nw. of Sterling. j. UT, Emery Co., 16 km e. of
Fremont Jct. k. CO, Garfield Co., 12 km nw. of Rifle. 1. UT, Kane Co., 30 km s. of
Cannonville.

tion of unreduced gametes, could generate bursts of additional trip-
loids and tetraploids, because most aneuploid gametes from triploids
are expected to fail. Progeny from triploids, therefore, would be
diploids, triploids and tetraploids. The diploids would breed nor-
mally; the tetraploids would leave only triploid progeny; the triploids
would again leave diploids, triploids and tetraploids. Consequently,
there could arise, from an occasional triploid, self-accelerating bursts
of triploids and tetraploids until sufficient tetraploids occurred to
permit autonomous perpetuation.

Tetraploid plants of G. brandegei differ from diploids phenotyp-

1987] STUTZ ET AL.: GRAYIA BRANDEGEI 147

IDAHO

WYOMING

|?
Sake Rive, °POCATELLO

114°

42°

ROCK SPRINGS

River

Fic. 3. Distribution of verified diploid and tetraploid forms of Grayia brandegei.
Symbols: O = 2X, @ = 4X.

148 MADRONO [Vol. 34

ically, particularly in stature and leaf-width, and have different geo-
graphic distributions, and so we propose them as a distinct variety.

Grayia brandegei A. Gray var. plummeri Stutz and Sanderson, var.
nov.

Similis var. brandegei sed foliis latioribus, plantis altioribus, la-
tioribus et tetraploideis differt, chromosomatum numerus 2n = 36.

Erect shrub 6-14 dm high, 2.4-6 dm broad; branches erect or
ascending, densely and finely pubescent. Leaf blades elliptic, ovate,
obovate or oblanceolate, 2.5—6 cm long, 5—22 mm wide, apex obtuse
or rounded. Leaf-anatomy non-Kranz. Heterodichogamously mon-
otecious. Male inflorescence glomerulate, the glomerules borne in
axils of leaves or small bracts; perianth 4-(5) parted, segments mem-
branaceous, obovate; stamens 4(5), filaments subulate, anthers di-
dymous, included. Female inflorescence paniculate, the flowers borne
in axils of leaves or small bracts; bibracteolate, bractlets 5-6 mm
wide, orbicular, completely united, margins extended into two wings
4-8 mm broad; perianth lacking; stigmas 2, filiform; utricle orbic-
ular, compressed, included in the two bracts; pericarp membrana-
ceous, free. Seed orbicular, 2-4 mm broad, erect; testa thin, mem-
branaceous; embryo annular; endosperm copious; radicle inferior.
Chromosome number: 2n = 36.

Type: USA, UTAH, Duchesne Co.: ca. 18 km e. of Roosevelt, 31
Aug 1984, Stutz 9325 (Holotype: BRY).

PARATYPES (all deposited in BRY): USA, CO, Garfield Co.: ca. 12
km nw. of Rifle, 18 Aug 1979, Stutz 8478. Moffat Co.: Sand Wash
near Dugout Draw, TION R97W S28, 31 Aug 1982, Parks 908. Rio
Blanco Co.: 25 km e. of Rangely on US 64, 7 Jun 1965, Collotzi
551. NM, San Juan Co.: Angel’s Peak badlands, ca. 20 km se. of
Bloomfield, 5 Jun 1985, Stutz 9438. UT, Daggett Co.: East Grind-
stone Spr., e. end of Antelope Flat, T3N R22E S24, 6600 ft, 29 Aug
1978, Neese and England 6648. Emery Co.: I-70, ca. 16 km e. of
Fremont Jct., 10 Sep 1983, Stutz 9153. Garfield Co.: 1 km e. of
Henrieville, 20 Jun 1986, Stutz 94337. Kane Co.: Cottonwood Wash,
ca. 12 km n. of US hwy 89, 12 Sep 1971, Atwood and Kaneko 3319.
Sanpete Co.: clay hills w. side antelope valley, 10 km nw. of Sterling,
21 May 1984, Stutz 9245. Uintah Co.: ca. 18 km e. of Roosevelt,
25 May 1985, Stutz 9430. Wayne Co.: 3 km e. of Notom Exit, Hwy
24, 20 Jun 1984, Stutz 9374. WY, Carbon Co.: 3 km nw. of Baggs,
1 Sep 1984, Stutz 9330.

The varietal name is chosen to honor A. Perry Plummer who has
pioneered numerous important studies of shrubs in western North
America. He also discovered the only known population of Grayia
brandegei growing in the Great Basin (Utah, Sanpete Co., 10 km w.
of Sterling, 21 May 1984, Stutz 9245).

1987] STUTZ ET AL.: GRAYIA BRANDEGEI 149

ACKNOWLEDGMENTS

Appreciation is expressed to Utah International, Inc., Brigham Young University,
and USDA Forest Service (Cooperative Agreement 22-C-4-INT-45 Brigham Young
University and USDA Forest Service) for financial support of the study, and to Bruce
L. Webb, Mildred R. Stutz, and R. Craig Stutz for technical assistance.

LITERATURE CITED

CoLiotzi1, A. W. 1966. Investigations in the genus Grayia, based on chromato-
graphic, morphological, and embryological criteria. M.S. thesis, Utah State Univ.,
Logan.

Gray, A. 1876. Miscellaneous botanical contributions. Proc. Amer. Acad. Arts 11:
101-103.

(Received 21 Aug 1985; revision accepted 15 Dec 1986.)

ANNOUNCEMENT

GRADUATE STUDENT AWARDS

The Eleventh Graduate Student Meetings of the California Botanical Society
were held Saturday 25 April 1987 at the University of California, Davis.
Sixteen papers were presented and awards were presented to the following
individuals:

Completed Research
First Place—Jon Hart, UC Davis
Second Place— George Robinson, UC Davis

Research in Progress
First Place— Bruce Baldwin, UC Davis
Second Place— Herb Saylor, San Francisco State University
Third Place—Sam Hammer

Proposed Research
First Place—Stacy Giles, San Francisco State University
Second Place— Barbara Gartner, Stanford University
Third Place— Mike Wood, San Francisco State University

Naill McCarten
Graduate Student Representative

ALLIUM SHEVOCKII (ALLIACEAE), A NEW SPECIES
FROM THE CREST OF THE SOUTHERN
SIERRA NEVADA, CALIFORNIA

DALE W. MCNEAL
Department of Biological Sciences,
University of the Pacific,
Stockton, CA 95211

ABSTRACT

Allium shevockii, a new species from the crest of the southern Sierra Nevada on
the Kern Plateau in Kern County, California is described and illustrated. The new
species shows morphological similarities to A. atrorubens, A. fimbriatum, and A.
monticola in the A. sanbornii alliance. It differs from these species in its obovate to
oblanceolate perianth segments that are distinctly reflexed distally, and its long thread-
like rhizomes that terminate in bulblets in addition to those produced at the base of
the parent bulb.

A new species of A//ium was discovered by James R. Shevock
during floristic work in botanically unexplored and remote areas of
the southern Sierra Nevada. Populations of this species are scattered
over a relatively limited geographical area on Spanish Needle Peak
along the crest of the southern Sierra Nevada. Due to the rugged
nature of the habitat and limited access to this area, the full range
of this new taxon can only be surmised. Review of A//ium in major
U.S. herbaria (CAS, DAV, DS, GH, JEPS, MO, NY, POM, RSA,
UC, US, WS) failed to locate additional collections of it.

Allium shevockii McNeal, sp. nov.

Tunica exterior bulbi brunnea, reticulatione cellulari carens, tu-
nice interiores luteolae, in sicco rubescentes; bulbi facientes rhizo-
mata filiformia 3-10 cm longa facientia bulbos terminales vel fa-
cientes 1—2 bulbillos basales parientes rhizomata filiformia. Scapus
(7—)10—20(—29) cm longus. Folium singulum, teres, 15-33 cm longa.
Umbella 12-30 vel pluribus floribus. Segmenta perianthii alba ad
pallide viridia infra, marronina in triente superiore; segmenta ex-
teriora 12-14 mm longa et 4.5—6 mm lata, erecta, obovata ad ob-
lanceolata, acuta ad mucronata, marginibus irregulariter et non pro-
funde dentatis, reflexa et crispa ad apicem; segmenta interiora 11-
13 mm longa et 4—4.5 mm lata, ovata, acuta, marginibus integris,
latescentia ad apicem; stylus trilobus; ovarium manifeste cristatum
6 anguste triangularibus processibus, margines exteriores proces-
suum undulatae ad non profunde et irregulariter dentatas (Fig. 1).

MaproNo, Vol. 34, No. 2, pp. 150-154, 1987

f shor, |
BN\l
hay

\ (
SN

Fic. 1. Allium shevockii McNeal. a. Entire plant. b. Habit. c. Inner perianth
segment. d. Outer perianth segment. e. Flower with anther. f. Ovary with prominent
crests. g. Bulblet with rhizome and new bulb at tip (the lower bulb is the new one).
h. Bulb with stipitate bulblet that has developed a rhizome. Drawn from Shevock
11219 and 35 mm transparencies.

152 MADRONO [Vol. 34

Bulbs subglobose, 10-15 mm long, outer bulb coat brown, lacking
cellular reticulation, inner coats light yellow, turning reddish on
drying, with obscure + quadrate cellular markings; bulbs forming
thread-like rhizomes 3-10 cm long that develop terminal bulblets
or forming | or 2 basal bulblets that produce thread-like rhizomes.
Scape terete, succulent, fragile when fresh, (7—)10—20(—29) cm long;
leaf one, terete, 15-33 mm long. Umbels 12-30 or more flowered;
bracts usually 3 (rarely 2), 16-20 mm long, 5-8 mm wide, lanceolate,
apiculate; pedicels 10-16 mm long. Outer perianth segments 12-14
mm long and 4.5-6 mm wide, erect, oblanceolate, acute to mu-
cronate with irregularly shallow toothed margins, white to light green
below, maroon on the reflexed, curled distal one-half; inner segments
11-13 mm long and 4—4.5 mm wide, ovate, acute, margins entire,
white to light pink or maroon on the upper one-third, outwardly
flared at the tip; stamens 3—'4 as long as the perianth, anthers ca. 1
mm long, yellow, elliptic, mucronate; styles ca. equalling the sta-
mens, three-lobed, ovary prominently crested with 6 narrowly tri-
angular, radially oriented processes, processes emarginate, outer
margins undulate to shallowly and irregularly toothed. Seed coat
black with hexagonal, minutely pustuliferous cells. Chromosome
number n = 7 (from the type collection).

Type: USA, CA, Kern Co.: W. slope of Spanish Needle Peak near
summit, ca. 1.2 air km s. of the Tulare-Kern-Inyo Co. line, T25S
R37E S4, 2315 m, 15 Jun 1985, Shevock 11219 (Holotype: CAS;
isotypes: CPH, NY, RSA).

PARATYPES: USA, CA, Kern Co.: From the type locality, 10 Jun
1986, McNeal and Boyd 3155 (CAS, CPH, NY, RSA, UC); e. slope
of Spanish Needle Peak, 2300 m, 10 Jun 1986, Shevock, Norris, and
Bagley 11636 (CAS, CPH, RSA, MO, NY, US).

Distribution, habitat, and phenology. Allium shevockii occurs in
soil pockets on dark colored metamorphic (chlorite-chloritoid schist)
outcrops, an adjacent igneous (aplite) intrusion, and on steep col-
luvial talus slopes between 2200-2350 m on Spanish Needle Peak
in the southern Sierra Nevada, Kern Co., California. Bulbs mainly
occur along the margins of outcrops where the slope is more stable.
Reproduction appears to be primarily vegetative. Few mature flow-
ers with developing capsules have been observed.

Vegetation in the general area is an open, mixed evergreen forest.
Associates in the immediate vicinity are sparse due to the steep,
unstable slopes. Associated species include: Arabis davidsonii Greene,
Arabis sp., Caulanthus pilosus S. Wats., Cercocarpus intricatus S.
Wats., Dudleya calcicola Bartel & Shevock, Epilobium canum
(Greene) Raven subsp. Jatifolium (Hook.) Raven, Eriogonum breed-
lovi (J. T. Howell) Reveal var. shevockii J. T. Howell, Eriogonum
nudum Dougl. ex Benth., s.l., Eriogonum umbellatum Torr., s.1.,

1987] McNEAL: NEW ALLIUM Wes)

Eriogonum wrightii Torr. ex Benth. subsp. subscaposum S. Wats.,
Eriophyllum ambiguum (Gray) Gray var. paleaceum (Bdg.) Ferris,
Eriophyllum confertiflorum (DC.) Gray, s.l., Juniperus occidentalis
Hook., Keckiella breviflora (Lindl.) Straw, Leptodactylon pungens
(Torr.) Rydb. subsp. pulchriflorum (Brand) Mason, Mimulus sp.,
Pellaea mucronata (D. C. Eat.) D. C. Eat., Pinus monophylla Torr.
& Frem., Quercus chrysolepis Liebm., Scrophularia desertorum
(Munz) R. Shaw, and Symphoricarpos parishii Rydb.

Allium shevockii is yet another highly restricted endemic from the
southern Sierra Nevada. Habitat with similar slope, aspect, geology,
and elevation occurs along the crest south of Spanish Needle to Mt.
Jenkins, a distance of 6.5 air km. Access to this area will be enhanced
greatly with the completion of the Pacific Crest Trail section that is
currently under construction between Spanish Needle and the Owens
Peak-Mt. Jenkins saddle. Suitable habitat north of Spanish Needle
toward Sawtooth Peak and south of Mt. Jenkins toward Walker Pass
along the crest appears to be lacking due primarily to a change in
geology. In addition to the type locality, A. shevockii was located in
six small, adjacent canyons on the west face and two canyons on
the east face of Spanish Needle. These populations are composed of
several thousand individuals. Approximately 10% of the suitable
habitat has been surveyed at this time. All populations are free of
human disturbances and are likely to remain so due to the rugged
nature of the habitat.

Relationships. Allium shevockii belongs to the A. sanbornii alliance
(Saghir et al. 1966) that is characterized by taxa having a single terete
leaf per scape and a prominent ovarian crest with six processes (two
per lobe). Allium shevockii possesses several features unique in the
alliance including: 1) obovate to oblanceolate perianth segments, the
outer series of which are strongly reflexed to coiled in the distal half;
2) the light-lemon yellow fresh bulb coats; 3) the long filamentous
secondary rhizomes that develop from the main bulb or more com-
monly from basal bulblets that form on short, stout primary rhi-
zomes at the base of the main bulb. These characters make A. shev-
ockii distinctive and easily recognized.

The position of A. shevockii within the A. sanbornii alliance is
problematical. In its formation of basal bulblets and more or less
entire crest processes, it is similar to A. atrorubens S. Wats. and A.
monticola A. Davids., but otherwise appears not to be related closely
to either taxon that lack the secondary, filamentous rhizomes and
distinctively reflexed outer perianth segments. In stature and the
three-lobed style, it resembles A. fimbriatum S. Wats., s.1., but it
seems not to be closely related to this taxon, which lacks both basal
bulblets and filamentous rhizomes and has dentate to lacinate crest

154 MADRONO [Vol. 34

processes. Allium shevockii apparently has evolved a series of unique
features independent of other members of the 4. sanbornii alliance.

ACKNOWLEDGMENTS

I thank James R. Shevock for specimens, information, photographs and field as-
sistance, Dr. Robert Smutny of the Department of Classics, University of the Pacific,
for assistance in preparing the Latin diagnosis and Sandra McNett-McGowan for
preparing the drawings. I appreciate helpful reviews of the manuscript by Drs. J.
Henrickson and L. V. Mingrone. Financial support for this research from the Faculty
Research Committee and F. R. Hunter Memorial Fund of the University of the
Pacific is acknowledged gratefully.

LITERATURE CITED

SAGHIR, A. R. B., L. K. MANN, M. OwnBeEy, and R. Y. BERG. 1966. Composition
of volatiles in relation to taxonomy of American Alliums. Amer. J. Bot. 53:477-—
484.

(Received 27 May 1986; revision accepted 6 Jan 1987.)

ANNOUNCEMENT

NEw PUBLICATION

WEBER, W. A., Colorado Flora: Western Slope, Colorado Associated Univ.
Press, 1334 Grandview Ave., Box 480, Univ. Colorado, Boulder 80309, 1987,
550 pp., 107 pl., 64 color pl., $20.50. [Illustrated manual of vascular plants
of the entire hydrological western slope of Colorado. Same format as Rocky

Mountain Flora, with type faces reduced to accommodate more text. Intro-
duction contains essays on floristic zones, pronunciation (European recom-
mended!), common names (discouraged!), the Colorado-Altai plant geography
connection, generic concepts (non-traditional!), eponymy. Keys to families,
genera and species with derivations, glossary. Statements of habitat and en-
demic status. Families and genera alphabetical. Non-numerical index uses
three-letter family acronyms. Controversial? Yes, indeed! Dedicated to Greene,
Rafinesque, Rydberg, Camp, Shinners, and Love.

CLAYTONIA PALUSTRIS (PORTULACACEAB),
A NEW SPECIES FROM CALIFORNIA

JOHN R. SWANSON
Department of Biology,
California State University, Northridge,
Northridge, CA 91330

WALTER A. KELLEY!
Biology Department, Mesa College,
Grand Junction, CO 81501

ABSTRACT

A new species of Claytonia (Portulacaceae) is described from the Sierra Nevada of
California. Morphology, chromosome number, and distribution indicate a close re-
lationship with C. sibirica.

Montia heterophylla (Torr. & Gray) Jepson was transferred to
Claytonia by Swanson (1966) in his assessment of systematic rela-
tionships in the Montioideae (Portulacaceae). Jepson (1914) had
indicated that the combination was based on C. unalaschkensis B
heterophylla Nuttall. This was a manuscript name published by
Torrey and Gray (1838) as asynonym of C. alsinoides y heterophylla
Torr. & Gray. Examination of the type of heterophylla (Nuttall s.n.,
“Oregon”, NY!), shows it belongs to C. sibirica L., and differs from
California collections upon which Jepson, Swanson, and others have
based their usage of this epithet. The two collections cited by Jepson
(Jepson 4884, JEPS; Hall and Chandler 304, UC) are stoloniferous,
whereas the Nuttall specimen is not. Our further studies of the
California plants, which in the past have been referred to C. (Montia)
heterophylla, show that they represent a distinct species. This new
species is differentiated easily from C. sibirica and from members
of Claytonia sect. Rhizomatosae (C. cordifolia S. Watson, C. neva-
densis S. Watson, C. sarmentosa C. A. Meyer, and C. scammaniana
E. Hulten).

Claytonia palustris Swanson and Kelley, sp. nov.

Herbae perennes, glabrae. Caules simplices, ca. 10—20 cm alti.
Radix verticalis, stolones ex axillis foliorum producens. Folia cau-
lesque virides; radix stolonesque albidi. Inflorescentia terminalis,
racemosa, floribus in axillis bractearum, numerosis. Ovarium pos-
tremo uniloculare, ovulis 3 (Fig. 1).

MADRONO, Vol. 34, No. 2, pp. 155-161, 1987
' Request reprints from Walter A. Kelley.

156

MADRONO

[Vol. 34

Fic. 1.

Claytonia palustris. Drawn from Swanson 490.

1987] SWANSON AND KELLEY: NEW CLAYTONIA 157

Perennial glabrous herbs with white fleshy rootstalks 2-3 mm
wide, 4-15 mm long. Stolons white, terete, 1-2 mm in diameter, 5-
15 cm long, arising from the axils of the basal leaves. Basal leaves
1-10, lamina oblanceolate 3—4 cm long, 1—1.5 cm wide, petioles 5—
10 cm long. Flowering branches several, 10-20 cm long; inflores-
cence terminal, racemose, 5-10 cm long, subtended by a pair of
opposite, subsessile, slightly unequal, oblanceolate to narrowly el-
liptic leaves 0.5-1.5 cm wide, 2—5 cm long, sessile or tapering into
a winged petiole. Flowers 5-12, each subtended by a sessile, ovate
to elliptic bract, 1-3 mm wide, 3-8 mm long; sepals 2, 1-3 mm
wide, 3—4 mm long; petals 5, oblong, white, emarginate, 5-9 mm
long; stamens 5, opposite the petals, filaments 4-7 mm long, pollen
3-colpate; ovary unilocular at maturity, 2-valved, ovoid, 2-3 mm
long; 1.5—2.5 mm wide; ovules 3; styles 3-5 mm long, style branches
3, each branch only partially stigmatic. Seeds 3, glossy, black. Chro-
mosome number n = 6.

Type: USA, CA, Butte Co.: Jonesville, marshy, sloping, spring-fed
meadow above s. bank of Jones Creek, 0.4 km e. of the Jonesville
Hotel, 1550 m, 4 Jul 1959, Swanson 490, (Holotype: OSC; isotypes:
CAS, CS, Mesa College, RSA, NY, SFV, UC).

PARATYPES: CA, Plumas Co.: Lake Almanor, 18 Jun 1920, Cle-
mens s.n. (CAS); Butterfly Valley, 23 Jun 1967, Howell 42704 (CAS,
OSU). Butte Co.: Chico Meadows, Sierra Nevada, 25 Jun 1915,
Heller 12022 (CAS); Jonesville, s. bank of Jones Creek, 0.4 km e.
of Jonesville Hotel, 13 Jun 1959, Swanson 477 (SFV). Fresno Co.:
Pine Ridge, 15-25 Jun 1900, Hall and Chandler 304 (UC); Frying
Pan Meadow, s. fork of Kings’ River, 12 Jul 1940, Munz 15960
(RSA, UC). Tulare Co.: Sequoia National Park, vicinity of Alta Peak,
4 Aug 1896, Dudley 1568 (DS); Jordan Hot Sprs., 18 Jul 1906, Hall
and Hall 8392 (UC); Cliff Creek, near junction of Timber Gap and
Black Rock Pass trails, 31 Jul 1943, Ferris and Lorraine 10944 (UC);
Tule River, Sierra Nevada, Pierson 1860 (RSA); Crescent Meadow,
Sequoia National Park, 17 Jun 1956, Tillett 464 (RSA); s. fork of
middle fork of Tule River, 6 Jun 1974, Gordon et al. 255 (SFV).

Claytonia palustris is characterized by a fleshy perennial stem 2-—
3 mm in diameter from which basal rosette leaves, stolons, scapose
inflorescence branches, and adventitious roots arise. A distinctive
feature of the species is the unequal length of the two opposite leaves
that subtend the inflorescence, a phenomenon not found in any other
Claytonia species.

Distribution and ecology. Claytonia palustris is endemic to Cali-
fornia. It is known from two disjunct regions at opposite ends of the
Sierra Nevada (Fig. 2) and from a third disjunct region in Siskiyou
Co. (see Miller et al. 1984, Fig. 4). The species is largely restricted

158 MADRONO [Vol. 34

to sunny areas, in wet meadows, marshy slopes, and streamside
vegetation at mid elevations (1025-1650 m) in the north, and to
mid and high elevations (1550-2450 m) in the south. Throughout
most of its range, C. palustris does not overlap with the known
distribution of C. sibirica s.1., the taxon with which it was confused
by Jepson (1914) when he described the distribution of M. hetero-
phylla as ‘““Southern Sierra Nevada, 5700-7000 feet. Oregon to Alas-
ka.”” The Nuttall type locality, ““Columbia Woods’’, is within the
range of C. sibirica, which is widespread in the northwestern United
States, coastal British Columbia, Alaska, and the Aleutian and Com-
mander Islands. Members of the C. sibirica complex extend south-
ward into the California coast ranges to Santa Cruz Co., and eastward
to Siskiyou Co. in the northern part of the state, where the complex
overlaps the range of C. palustris.

The habitats in which C. sibirica s.l. is found in California and
Oregon are moist but not marshy. Claytonia sibirica is adapted to
growing in shaded habitats, whereas C. palustris is not. Many of the
habitats occupied by C. sibirica are disturbed by natural causes or
by man and it often grows as a ruderal plant. In Great Britain where
C. sibirica was introduced in the last century, it has become wide-
spread and is considered a weed (Salisbury 1961). Ruderal tendencies
are not apparent in any of the populations of C. palustris.

The distribution of C. nevadensis approximates that of C. palustris
(Fig. 2), although the individual populations are isolated geograph-
ically and ecologically. At similar latitudes, Claytonia nevadensis is
found at higher elevations than C. palustris. The former is restricted
to relatively pure stands, whereas C. palustris is found embedded in
a dense tangle of perennial vegetation.

Claytonia cordifolia is widespread in the pacific northwest and
extends southward into Siskiyou Co., California. Ecologically this
species is similar to C. sibirica in its capacity to grow in shaded
habitats, and to C. palustris in its ability to withstand competition
from other vegetation. Claytonia cordifolia also is generally found
in marshy or very moist situations.

Variation. Field and herbarium studies indicate that species of
sect. Rhizomatosae are relatively uniform throughout their range.
Chambers (1963) also noted the uniformity of C. nevadensis pop-
ulations. Claytonia sibirica, however, is quite variable within and
between populations. Miller et al. (1984) demonstrate that a euploid
series exists in the taxon. The five species of sect. Rhizomatosae rely
largely upon asexual reproduction by the use of branching rhizomes
or stolons. Asexual reproduction is an adaptation common in plants
living in climatically severe environments or where seedling estab-
lishment is difficult due to competition in closed communities (Steb-
bins 1950).

1987] SWANSON AND KELLEY: NEW CLAYTONIA 159

OREGON

CALIFORNIA NEVADA

© C. nevadensis

@c palustris

Fic. 2. Known localities for Claytonia nevadensis and C. palustris.

Chromosome counts. Previously unreported chromosome counts
are given here; vouchers are deposited at SFV. Four collections of
C. palustris (Butte Co., Swanson 477, 490; Plumas Co., Swanson
1047; Tulare Co., Swanson 1162) are all n = 6. Three populations
of C. nevadensis are n = 7 (Lassen Co., Swanson 509; Mono Co.,
Swanson 1055, 1056). Counts were performed on three plants from
each collection. All counts were made from anthers fixed in 3:1
ethanol-acetic acid and stored in 70% ethanol at 4°C until used.
Meiosis was regular in all microsporocytes examined.

160 MADRONO [Vol. 34

TABLE 1. CHARACTERS USED TO DIFFERENTIATE C. palustris AND CLOSELY RELATED
Species. See Lewis (1967) and Miller et al. (1984) for references to chromosome
numbers not reported in this paper.

Maxi-
mum
Ovule seeds Inflo-
num- ma- rescence Chromosome Runners or
Species ber __ tured bracts numbers stolons
C. cordifolia 6 3 none n= 5, no
2n = 10,. 20%
C. nevadensis 6 6 one n=7, yes
2n = 14,
C. palustris 3 3 several n= 6, yes
2n= 125
C. sarmentosa 6 6 none n= 7, 8, yes
2n = 14, 164,
28, 324
C. scammaniana 6 6 none unknown unknown
C. sibirica 3 5 several n= 6, infrequent
2n = 12), 24n,
36n

Relationships. In this paper, binomials are used to represent the
nominal taxonomic species although different ploidy levels are known
to exist within some of them and reproductive isolation is presumed
to exist. The principal features we used to separate species of Clay-
tonia are: 1) ovule number; 2) seed number; 3) ovule abortion; 4)
number of bracts in the inflorescence (e.g., none, one subtending the
lowermost flower, or several and each one subtending a flower); 5)
base chromosome number; and 6) the presence or absence of stolons
(Table 1).

Claytonia palustris shows similarities in these characters with
members of sect. Rhizomatosae (C. nevadensis, C. sarmentosa, C.
scammaniana and C. cordifolia) and members of C. sibirica s.1. (sect.
Caudicosae). Jepson (1914, p. 474) referred to C. palustris accurately
when he wrote (of M. heterophylla), “Stems. . . rising from tuberous
rootstocks or cormlets, these sending out slender stolons that pro-
duce terminal cormlets, the secondary cormlets promptly producing
leaves and flowers... .” In these respects, C. palustris most closely
resembles C. nevadensis and occasional plants of C. sibirica. Mor-
phologically, there are greater differences between these species and
C. cordifolia (Table 1). Our data suggest the close relationship of C.
palustris and C. sibirica. In addition, similarities in flower size and
morphology, leaf shape, leaf and stem texture, and plant and flower
color support this suggested relationship. The occasional formation
of stolons by C. sibirica (Swanson 1966) is quite similar to that in

1987] SWANSON AND KELLEY: NEW CLAYTONIA 161

C. palustris. The same base number (” = 7), six-ovuled flowers,
pigmentation of vegetative parts and flowers, and texture of leaves
suggest the affinity of C. nevadensis and C. sarmentosa and possibly
C. scammaniana (chromosome number unknown). Claytonia cor-
difolia is isolated in sect. Rhizomatosae on the basis of chromosome
number (n = 5). The regular abortion of three of the six ovules
produced by each flower of C. cordifolia is an example of an evo-
lutionary phenomenon that may have occurred independently in at
least three groups in the genus (i.e., C. sibirica, C. perfoliata Donn
ex Willd., and C. spathulata Dougl. ex Hook. species complexes).

Chambers (1963) has noted a number of anatomical differences
between C. cordifolia and C. nevadensis. Further anatomical and
morphological studies of C. sibirica s.1., C. palustris, and members
of sect. Rhizomatosae are necessary before the relationship and tax-
onomy of these species can be clarified.

ACKNOWLEDGMENTS

Thanks are due R. C. Bacigalupi for correcting the Latin diagnosis and Francis
Runyan for the illustration. Special thanks goes to Kenton Chambers for providing
insight on relationships in Claytonia and the helpful comments on the manuscript.
John Miller and Charles Fellows were the source of a number of stimulating discus-
sions.

LITERATURE CITED

CHAMBERS, K. L. 1963. Claytonia nevadensis in Oregon. Leafl. W. Bot. 10:1-8.

HUuLTEN, E. 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press,
Stanford.

JEPSON, W. L. 1914. Portulacaceae. Jn W. L. Jepson, A flora of California. Part 5,
p. 465-480. Univ. California, Berkeley.

Lewis, W. J. 1967. Cytocatalytic evolution in plants. Bot. Rev. 33:105-115.

MILLER, J. M., K. L. CHAMBERS, and C. E. FELLows. 1984. Cytogeographic patterns
and relationships in the Claytonia sibirica complex (Portulacaceae). Syst. Bot. 9:
266-271.

SALISBURY, E. J. 1961. Weeds and aliens. Collins, London.

STEBBINS, G. L. 1950. Variation and evolution in plants. Columbia Univ. Press,
New York.

SWANSON, J. R. 1966. A synopsis of relationships in Montioideae (Portulacaceae).
Brittonia 18:229-241.

TorREY, J. and A. Gray. 1838. A flora of North America. 1:198-202. New York.

(Received 14 Nov 1985; revision accepted 29 Oct 1986.)

ALLOISPERMUM INSUETUM
(ASTERACEAE: HELIANTHEAB),
A NEW SPECIES FROM COLOMBIA

CARMEN F. FERNANDEZ, LOWELL E. URBATSCH, and GENE SULLIVAN
Department of Botany, Louisiana State University,
Baton Rouge 70803

ABSTRACT

A description and illustration are provided for Alloispermum insuetum, a new
species from Colombia.

Alloispermum Willd. (Heliantheae: Galinsoginae) includes ap-
proximately 10 species of suffrutescent shrubs and a few truly her-
baceous species that occur in dry to moist mountainous regions from
Mexico to northern South America. The generic name was proposed
by Willdenow (1807), but not used until revived by Robinson
(1978a,b, 1979). The previously undescribed species was discovered
among specimens examined as part of our ongoing investigations in
the subtribe.

Alloispermum insuetum Fernandez, Urbatsch, and Sullivan, sp. nov.

Suffrutex, 1-2 m altus. Folia lanceolata, 5—10 cm longa. Capitu-
lescentia ad 15 cm lata, capitulae ca. 10, pedunculis 2.5—7 cm longis.
Involucra hemisphaerica, ca. 9 mm alta, 15 mm lata. Flores radii
ca. 15, ligulae albae tincta apice roseae, flores disci 40-65. Achenia
radii 2.5 mm longa, glabrescentia; achenia disci 2.7 mm longa, pu-
bescentia sparsim, pappi radiorum et discorum similes, squamae ca.
15, plerumque 5—5.5 mm longae (Fig. 1).

Weak, suffrutescent shrubs, 1—2 m tall. Leaves subsessile; blades
lanceolate, 5-10 cm long, 2.5 cm wide, basally obtuse, apically long
acuminate, abaxially pilose, with uniseriate, 5—6-celled trichomes,
mainly along the veins, adaxially pubescent, with uniseriate, 2-3-
celled trichomes; margins remotely serrate. Capitulescence corym-
bose, ca. 15 cm broad, ca. 10-headed; peduncles 2.5—7 cm long,
densely pilose. Capitula 55—80-flowered; involucres hemispheric, ca.
9 mm long, ca. 15 mm wide, with phyllaries 2—3-seriate, obovate,
6.5-8.5 mm long, 4—6 mm wide, pubescent, apically acute; receptacle
conical, 4.5 mm long, 4 mm wide. Ray flowers ca. 15; corollas 14—
18 mm long, ligules 12—15 mm long, up to 5.5 mm wide, white with
the apex tinged pinkish-red; corolla tube 2.5—4.2 mm long, densely
pilose; disc flowers 40-65; corollas 6.4 mm long; ray achenes 2.5

MADRONO, Vol. 34, No. 2, pp. 162-164, 1987

1987] FERNANDEZ ET AL.: NEW ALLOISPERMUM 163

f
7] A
Jy) Y

x aa
oS ye
=i. = iG

Fic. 1. Alloispermum insuetum. A. Habit. B. Capitulum. C. Disk flower. D. Ray
flower. Drawn from Schlim 359.

mm long, mostly glabrous. Disc achenes 2.7 mm long, sparsely:
pubescent; pappus of both ray and disc achenes ca. 15 linear-lan-
ceolate, fimbriate scales, 5—5.5 mm long.

TYPE: Colombia, Norte de Santander, Provincia de Ocana: 8000—
10,000 ft, Jan 1852, Schlim 359 (Holotype: K!; isotype: BM). The
collection date for the type material can not be ascertained directly
from the specimen label data. According to Linden (1867), Schlim
explored the Ocana region in 1851 until the beginning of 1852.

164 MADRONO [Vol. 34

PARATYPE: Colombia, Norte de Santander, around Ocana, Schlim
440 (K!).

Alloispermum insuetum is known only from the type material and
one other specimen; it grows from 2400-3200 m and flowering oc-
curs in January. This species is similar in habit, disc pappus, leaf,
phyllary, and achene features to the other species in the genus. It is
distinguished easily from the other South American species by its
larger heads, longer ligules, and the presence of a pappus in the rays.
The specific epithet “‘insuetum’”’ (=unusual) was chosen to call at-
tention to these features. A//oispermum insuetum is similar to larger-
headed forms of A. caracasanum (Kunth) H. Robinson, but the
former has more than 40 disc flowers and about 15 ray flowers per
head, whereas the latter has fewer than 35 and 8, respectively.

ACKNOWLEDGMENTS

We thank Elizabeth Harris for preparing the illustration, J. Pruski (NY) for infor-
mation on the isotype at BM, and the various herbaria for their loans of specimens.
Financial assistance was provided by the Botany Department, Louisiana State Uni-
versity and NSF Grant No. DEB78-04265 to L.E.U.

LITERATURE CITED

LINDEN, J. J. 1867. Les explorations botaniques de la Colombie. Belgique Hort. 17:
235-256.

ROBINSON, H. 1978a. Studies in the Heliantheae (Asteraceae). IX. Restoration of
the genus Alloispermum. Phytologia 38:41 1-412.

1978b. Studies in the Heliantheae (Asteraceae). XV. Various new species

and new combinations. Phytologia 41:33-38.

1979. Studies in the Heliantheae (Asteraceae). X XI. Additions to Allois-
permum, Galinsoga, and Tridax. Phytologia 44:425-435.

WILLDENOw, C. L. 1807. Einige Bemerkungen tiber die Pflanzen der Klasse Syn-
genesia. Ges. Naturf. Freunde Berlin Mag. Neuesten Entdeck. Gesammten Na-
turk. 1:132-141.

(Received 19 Nov 1985; revision accepted 10 Nov 1986.)

ANNOUNCEMENT

REVISED EDITION

TERRELL, E. S., S. R. Hitt, J. H. WieRSEMA, and W. E. Rice, A checklist of
names for 3,000 vascular plants of economic importance, Revised ed., U.S.D.A.,

Agricultural Handbook, no. 505, pp. [i-1i], 1-241, Oct. 1986, no ISBN, paper-
bound, price unknown (from Superintendent of Documents, Government
Printing Office, Washington, DC 20402). [First edition = 1977; a useful com-
pilation of common names and accepted scientific names for 1241 genera and
3296 species, subspecies, and varieties, with inclusion of 983 rather common
synonyms. ]

A NEW SPECIES OF AXINIPHYLLUM
(ASTERACEAE: HELIANTHEAE) FROM
DURANGO, MEXICO

B. L. TURNER
Department of Botany, University of Texas,
Austin 78713

ABSTRACT

Axiniphyllum durangense from southern Durango, Mexico, is related to A. cor-
ymbosum of Guerrero and adjacent Oaxaca, but is distinguished readily by its rhom-
bic-ovate leaves and much longer outer involucral bracts.

In spite of a recent revisional treatment of the relatively small,
uncommon genus Axiniphyllum (Turner 1978), I have observed
recent collections that represent an additional novelty from south-
western Durango. This brings to five the number of species now
recognized. It is noteworthy that McVaugh (1984) omitted the genus
from his Flora Novo-Galiciana, but the proximity of collections of
this genus to the north of Jalisco suggests that it also will be found
ultimately in that floristic region.

Axiniphyllum durangense B. Turner, sp. nov.

A. corymbosum accedens sed foliis tenuioribus rhombeo-ovatis,
phyllariis exterioribus multo longioribus (Fig. 1).

Erect perennial herb 50-75 cm high, arising from short, corm-like
rhizomes; the root-system fibrous. Stems terete, 2—5 mm thick be-
low; moderately to densely hirsute with crisp, spreading hairs; the
upper stems less pubescent and soon beset with a dense array of
short, glandular trichomes. Leaves opposite, rhombic-ovate, 5-8 cm
long, 1-3 cm wide, gradually tapering into a slender petiole that
abruptly flares below into a stipule-like, perfoliate appendage; blades
pubescent above and below principally along the venation, 3-nerved
from above the base, the margins irregularly serrulate. Heads 6-8,
in an open corymbose panicle, the ultimate peduncles 1—5 cm long.
Involucres hemispheric, phyllaries 2—3-seriate, the outer series of 4—
6 loose, leaf-like, mostly lanceolate bracts 10-15 mm long, 1.5—2.5
mm wide; the inner series of 8—11 + scarious, broadly ovate bracts,
6-7 mm long, 3—4 mm wide, puberulent. Receptacle convex, ca. 2
mm across, the bracts obovate, 5—6 mm long, ca. 2 mm wide, 3—4-
nerved, the apices acute. Ray florets absent. Disk florets 40-50,
yellow; corollas ca. 5 mm long, the tube ca. 1 mm long, pubescent

MADRONO, Vol. 34, No. 2, pp. 165-167, 1987

166 MADRONO [Vol. 34

Fic. 1. Axiniphyllum durangense (from holotype). a. Head. b. Base of stem with
roots. c. Mid-stem leaves. d. Capitulescence. e. Floret.

with both coarse hairs and short-stipulate glands, the limb ca. 4 mm
long, the lobes ca. 1 mm long, acute. Anthers yellow, ca. 2.5 mm
long, eglandular. Style branches hispid with abrupt conical append-
ages ca. 0.5 mm long. Achenes epappose, quadrate, black, smooth,
2.5—3 mm long, ca. 1 mm wide.

1987] TURNER: NEW AXINIPHYLLUM 167

TYPE: Mexico, Durango, Mpio. de Mesquital: “‘Alrededores de
Platano Tatemado (a 12 km de La Guajolota).... Margenes de
arroyo en Bosque de pinoencino”’;:ca. 23°30'N 104°30’W, 12 Sep
1985, I. Solis 294 (Holotype: TEX; isotypes: to be distributed).

PARATYPE: Durango, Mpio. Mezquital: ca. 11 km from La Gua-
jolota, 28 Sep 1985, Solis 379 (TEX).

In my treatment (Turner 1978), A. durangense will key near A.
corymbosum, an eradiate species of Guerrero and adjacent Oaxaca.
It differs markedly from that taxon in leaf shape, vestiture and tex-
ture. In addition, the involucral bracts of A. corymbosum are smaller
and the much narrower, outer series is only 3-5 mm long.

ACKNOWLEDGMENTS

I am grateful to Dr. M. C. Johnston for the Latin diagnosis, Dr. Linda Vorobik
for providing the illustration, and especially to Dr. J. Strother for calling to my
attention several glaring errors in an earlier draft of this paper.

LITERATURE CITED

McVauau, R. 1984. Flora Novo-Galiciana 12:1-1129.
TURNER, B. L. 1978. Taxonomy of Axiniphyllum (Asteraceae— Heliantheae). Ma-
drono 25:46—-52.

(Received 25 Sep 1986; revision accepted 6 Jan 1987.)

ANNOUNCEMENT

NEw PUBLICATIONS

JENKINS, D. I., Amanita of North America, Mad River Press, Route 2, Box
151B, Eureka, CA 95501, 1986, vi, 198 pp., illus. (color), ISBN 0-916422-
55-0 (paperbound), price unknown. [With keys to and descriptions of 128

species and varieties. ]

LAMPE, K. F., and M. A. McCann, AMA Handbook of Poisonous and Injurious
Plants, American Medical Association, Chicago, 1985, xi, 432 pp., illus. (most-
ly color), ISBN 0-89970-183-3 (flexibound), $24.95 (from Chicago Review
Press, 814 North Franklin, Chicago, IL 60610). [On plants of the United States,
Canada, and the Caribbean, with 437 color photographs.]

NOTES

RANGE EXTENSION, CHROMOSOME COUNT, AND MEPHITISM IN Lessingia tenuis
(COMPOSITAE).— Recent field work in Central California revealed a range extension
for Lessingia tenuis (A. Gray) Cov.: CA, Santa Clara Co., s. side of ridge ca. 4 airline
km s. of Los Gatos, near Priest Rock and power lines along a dirt road from Lexington
Reservoir to Sierra Azule Ridge, ca. 517 m, Mooring 3517, 3530, 3538 (UC). (fide
J. Strother, UC).

First observed here in 1984, these annuals were mostly 2-3 cm tall, relatively
narrow-leaved, and sparsely branched. They did not conform to descriptions (Munz,
A California Fl., 1959; Abrams and Ferris, Illustr. Fl. Pacific States, 1960) nor did
they closely resemble the herbarium specimens I examined. Thus, an experimental
study seemed appropriate.

Achenes collected at the site and sown in vermiculite in an unheated greenhouse
in November readily germinated ca. 7 days later. Twenty seedlings that were planted
in pots flowered an average 175 days after germination. Mature plants were 15-23
cm tall, relatively broad-leaved, and diffusely branched, in sharp contrast to their
wild parents. They matched the description of Lessingia tenuis, and resembled some
of the herbarium specimens examined. Analysis of microsporocytes stained in ace-
tocarmine showed in three plants that 2n = 5, and that meiosis was regular. This
count agrees with those for populations of L. tenuis in San Luis Obispo and Ventura
cos. [Spence, A biosystematic study of the genus Lessingia Cham. (Compositae),
Ph.D. diss., Univ. California, Berkeley, 1963]. Stainability of fresh pollen in cotton
blue-lactophenol (minimum of 300 pollen grains from each of 10 plants) ranged from
76-100% (k = 95%). Greenhouse plants differed conspicuously from wild progenitors
in that each had a pungent, skunk-like odor that is unreported in this genus. Spence
(pers. comm.) grew large numbers of L. tenuis and other species of Lessingia, and
did not notice this odor in the genus.

The presence of L. tenuis in the Santa Cruz Mountains represents a 60 km westward
range extension. I found the population while looking for local populations of the
Western Whiptail Lizard (Cnemidophorus tigris), which is another example of a
coastward outlier of an Inner Coast Range species (Mooring, Herp. Rev. 14:123,
1983). Lessingia tenuis may be a recent introduction here, from nearby power lines
or road, or may have increased its numbers rapidly. I have hiked through the Los
Gatos site intermittently for 20 years without seeing this species. I first noticed the
Los Gatos lizard populations during the high-precipitation years associated with El
Nino weather, and the discovery of L. tenuis came about 2 years later. The numbers,
density, and distribution of the population have varied. In 1984, an estimated 500
plants occurred along ca. 150 m and within 2 m of a north-to-south trail through
chaparral that is dominated by Adenostoma fasciculatum. In 1985 most of the ca.
150-200 individuals were within the northernmost 100 m, and also were close to the
trail. In 1986, however, several thousand were scattered in clumps over 358 m,
including hundreds in bare or thinly vegetated soil up to 20 m from the trail. The
population was less dense in the northernmost section than it had been before, possibly
due to the increased use of the trail by mountain bicycles. It has expanded southward,
however, and now abuts a bulldozed area adjacent to a 1985 burn that may allow
further expansion.

The skunk-like odor of the Los Gatos population should be looked for in other
populations of this and other species of Lessingia. Presence-absence patterns might
be a useful taxonomic character in a group where keys (Jepson, A Man. FI. Plants
California, 1925; Munz, op. cit.; Abrams and Ferris, op. cit.) emphasize vegetative
features. Pungent mephitism in L. tenuis seems to be associated with relatively lux-

MADRONO, Vol. 34, No. 2, pp. 168-169, 1987

1987] NOTES 169

uriant growth, and may not be obvious in wild plants. I did not notice it when
collecting and observing them in 1984. In 1986 I had to sniff the wild plants to detect
it, whereas half-grown wild plants that were transplanted to the greenhouse that year
were, when full-grown, almost as odorous as I remembered the 1984 greenhouse-
grown seedlings to be.

Mephitism may be an anti-herbivore adaptation in the Los Gatos population of
L. tenuis. They often occur in clumps; some occur under Adenostoma fasciculatum,
Arctostaphylos (glandulosa?), or Baccharis pilularis subsp. consanguinea. The shrubs
have no surrounding bare zones unlike the situations described previously (Bartho-
lomew, Science 170:1210—1212, 1970; Halligan, Bioscience 23:429-—432, 1973). Evi-
dence of grazing by mammals is present, but the lessingias and co-occurring annuals
[including Navarettia squarrosa(?), ““skunkweed”’] show no evidence of being grazed.
The lessingias, however, do not have an unpleasant taste (at least to me), and green-
house plants are attacked by whiteflies. Mephitism has been reported in Navarettia
squarrosa (see Abrams & Ferris, op. cit.) and I had noticed it in a Santa Cruz Co.
population growing in a relatively mesic site. I have not observed it among local
populations, however, but neither have I sniffed the plants. Perhaps mephitism, and
other strong scents, might be found to be more widespread if it is looked for in
greenhouse plants derived from dense populations of apparently ungrazed herbaceous
species of dry habitats.

I appreciate the comments of reviewers Pinkava and Tanowitz, and have extended
the discussion of mephitism.—JOHN Moorina, Biology Department, Santa Clara
Univ., Santa Clara, CA 95053. (Received 25 Jun 86; revision accepted 9 Oct 1986.)

REVIEW

Vascular Plants of Upper Bidwell Park, Chico, CA. By VERNON H. OSWALD. vi + 98
pp. The Herbarium, Department of Biological Sciences, California State University,
Chico, Publication No. 3. 1986. $5.95 plus tax and mailing.

This book has an attractive sketch of Polygonum bidwelliae on its soft yellow cover.
The content is formatted professionally and has appeal to anyone who wishes to deal
with basic botany with the assistance of keys, glossary, map, and bibliography. The
preface explains the three plant communities involved and has a synoptic geological
presentation. The nomenclature is up-to-date and keys involve major plant groups,
divisions, families, genera, species, and subspecific taxa. Although undoubtedly in-
complete (as is any other new checklist), 748 species and subspecific taxa have been
tabulated. About 30% of these are introduced and this is about 50% higher than on
the county list.

Two thousand acres are included in the study area. Elevations extend from 260
feet to 1520 feet. Twelve plants are listed in various categories in the C.N.P.S. rare
plant inventory. There is no mention of climatology and there are no illustrations or
photographs. A short addenda and errata are enclosed.

Even for amateurs who have only a superficial knowledge of botany, this profes-
sionally presented text will be found more enjoyable, and certainly more educational,
than a plant list keyed to the color of the flowers. — WALTER KNIGHT, Field Associate,
California Academy of Sciences, San Francisco, CA 94118.

NOTEWORTHY COLLECTIONS

ARIZONA

ASTRAGALUS HYPOXYLUS S. Wats. (FABACEAE).— Santa Cruz Co., Patagonia Mts.,
Coronado Natl. For., ca. 4.5 km s. of Harshaw on rd. to Washington Camp, 31°26.5'N,
110°43.5'W, stony openings in pine-oak-juniper woodland, restricted to limestone-
derived soil, sw. exposure, ca. 1600 m, 20 Apr 1986, Levin and Levin 1633 (NY,
RSA, SD), T. R. and R. K. Van Devender 86-140 (ARIZ) (det. R. C. Barneby). On
a subsequent visit, T. R. Van Devender (pers. comm.) found 107 plants on this
hillside and a few plants ca. 1 km s.

Significance. First collections since the type collected by J. G. Lemmon (Cochise
Co., Huachuca Mts., Maloney’s Ranch, Jul 1882, 2656) about 30 km e. The plants
are caulescent, with stems of the season up to 6 cm long, as described by Barneby
(Mem. N.Y. Bot. Gard. 13:1028-1029, 1964); Kearney and Peebles (Arizona F1., p.
469, 1951) incorrectly described the plant as acaulescent and caespitose. To Barneby’s
description, I add that the petals are whitish with bluish-purple tips, the banner is
recurved through about 45°, and the ovules may be up to eight in number; the seeds
and pod dehiscence remain unknown.— GEOFFREY A. LEVIN, see California notes
below.

CALIFORNIA

FESTUCA OCCIDENTALIS Hook. (POACEAE).—San Diego Co., Palomar Mtn. State
Park, Doane Valley Nature Trail ca. 250 m w. of Doane Pond, T10S RIE S5 ne.%4
of nw.%4, 1400 m, 18 Jun 1986, Curto and Allen 330 (SD); Los Coyotes Indian Res.,
Hot Springs Mtn., T10S R4E S15 nw. of se.4, 1790 m, 21 Jun 1986, D. Clemons,
Levin, and Curto 1509 (SD); Santa Ysabel Indian Reserv., Volcan Mts., n. and nw.
slopes of Oak Ridge, T12S R3E S12, 1585 m, 3 Jul 1986, Curto 336 (SD); Cuyamaca
Rancho State Park (CRSP), Engineers Rd. sw. of North Peak, T13S R4E, 1465 m, 3
Jun 1986, Curto and Allen 308 (SD); CRSP, w. slope of Middle Peak, along Middle
Peak Loop Fire Rd. | km n. of Milk Ranch, T14S R4E, 1585 m, 29 May 1986, Curto
and Allien 300 (SD, TAES, US); CRSP, Japacha Fire Rd. at Japacha Creek, T14S
R4E, 1330 m, 4 Jun 1986, Curto and Allen 310 (SD). At all sites, common in mixed
conifer or pine-oak woodlands growing on loamy soils derived from gabbro or gran-
odiorite.

Significance. A range extension of ca. 300 km se. from Santa Barbara Co. The
abundance of this grass in the Peninsular Ranges of San Diego Co. indicates that it
should be looked for in the intervening mountain ranges. Known previously from
B.C., Canada, s. to Santa Barbara Co., CA, e. across Canada to Ont., and in ID, MT,
WY, WI, and MI.—MIcHAEL CurRTO and LINDA ALLEN, California Dept. of Parks
and Recreation, 1333 Camino del Rio S., Suite 200, San Diego, CA 92108; and
GEOFFREY A. LEVIN, see notes below.

MIMULUS CONGDONII Rob. (SCROPHULARIACEAE). — San Diego Co., Cuyamaca Ran-
cho State Park, oak woodland border n. of meadow near Merrigan Fire Rd. ca. 0.8
km n. of Viejas Blvd., 32°52'N, 116°36.75'W, 1050 m, 24 Mar 1986, Curto 255 (SD).
Several populations of about 100 plants along seasonal creeks.

Significance. A range extension of ca. 300 km se. from the Coast Ranges in Ventura
Co. and ca. 350 km s. from the Greenhorn Mts., Kern Co. Known previously in the
Coast Ranges from Mendocino Co. to Ventura Co. and in the Sierra Nevada from
Mariposa Co. to Tulare Co.

MADRONO, Vol. 34, No. 2, pp. 170-171, 1987

1987] NOTEWORTHY COLLECTIONS 171

RHUS TRILOBATA Nutt. ex Torr. & A. Gray var. SIMPLICIFOLIA (Greene) Barkley
(ANACARDIACEAE).—San Diego Co., Pinon Peak [probably highest point in Pinyon
Mts., T13S R6E S11], 24 Sep 1938, Stover and Harbison s.n. (SD); Vallecito Mts.,
T13S R6E S14 ne.'4 of se.%4, 1355 m, 15 May 1983, D. Clemons and E. Jonsson 590
(SD); T13S R6E 823, 1220 m, 10 Apr 1985, Clemons and Jonsson 1012 (SD), 1013
(SD). Apparently uncommon in pinyon-juniper woodland.

Significance. First records of this variety for CA, a n. range extention of 70 km
from near La Rumorosa, Baja California Norte, Mexico. Known previously from
Baja California, s. UT, n. AZ, sw. CO, and OK. Beauchamp (A fl. San Diego Co.,
California, p. 82, 1986) incorrectly cited the Pinyon Mts. specimen as var. anisophylla
Jepson and he (op. cit., p. 170) used the Clemons and Jonsson specimens as the basis
for listing Ribes cereum Dougl. in San Diego Co. The latter species remains unknown
in the county.— GEOFFREY A. LEVIN, Botany Dept., San Diego Natural History Mu-
seum, P.O. Box 1390, San Diego, CA 92112.

COLORADO

BRYUM BLINDII B.S.G. (BRYOPSIDA: EUBRYA: BRYACEAE).— Grand Co., Arapahoe
Natl. For., Fraser River valley at base of Berthoud Pass, T2S R75W S34, 3100 m,
in saturated sand and gravel on steep slope just above roadside on e. side of lowest
hairpin turn, 14 Sep 1986, W. A. Weber & H. Dahnke 91852 (COLO; to be distributed
in Krypt. Exsicc. Vindob.).

Previous knowledge. Northern and central Europe, northern Asia and Japan; rare
in North America: Ontario (Thunder Bay), New Brunswick, northern Manitoba,
British Columbia, and southeastern Alaska (Crum & Anderson, Mosses of Eastern
North America, Vol. 1, 1981).

Significance. First record for contiguous United States. — WILLIAM A. WEBER, Univ.
of Colorado Museum, Boulder 80309.

New MEeExIco

LYGODESMIA GRANDIFLORA (Nutt.) Torr. & A. Gray (ASTERACEAE). —San Juan Co.,
Cutter Canyon, T29N R8W S21, silt slope, 1950 m, 9 Jun 1973, J. T. Wynoff 492
(ASU) (det. A. S. Tomb).

Significance. First record for NM. When L. grandiflora was redefined (Tomb, Syst.
Bot. Monogr. 1:1—51, 1980) to include only material from CO, UT, and WY, material
from AZ and NM was assigned to L. arizonica Tomb. Recently, L. grandiflora s.s.
was found in ne. AZ (Parfitt, J. Ariz. Nev. Acad. Sci., in press.) — BRUCE D. PARFITT,
Dept. Botany, Arizona State Univ., Tempe 85287.

PENSTEMON RAMOSUS Crosswhite (SCROPHULARIACEAE). — Dona Ana Co., Sierra de
las Uvas, base of Ponciello Peak, T20S R3W, 1500 m, 11 Jun 1977, Weber s.n.
(NMC); Dona Ana Mts., sheltered ne. slopes of Summerford Mt., T21S RIE S3, 1350
m, 27 May 1984, Todsen 8406-1 (NMC). Luna Co., ca. 1.5 km n. of Cooke’s Peak,
open slopes with juniper, 1800 m, T20S R8W, < 1) Oct 1979, Spellenberg, Isaacs, and
Soreng 5436 (NMC). Sierra Co., arroyo e. of Rio Grande and 0.5 km s. of Caballo
Dam, 1275 m, T16S R4W, 22 Jun 1986, Todsen A119 (NMC).

Significance. Known only from the US, P. ramosus was previously reported from
Cochise, Graham, Greenlee, and Pima cos. in se. AZ and Grant and Hidalgo cos. in
NM. Above are all new county records with the Dona Ana Mts. location 180 km e.
of the nearest previously reported site. — THOMAS K. TopsENn, Dept. Biol., New Mexico
State Univ., Las Cruces 88003.

REVIEW

Uinta Basin Flora. By SHEREL GOODRICH and ELIZABETH NEESE. 320 p. U.S.D.A.
Forest Service— Intermountain Region, Ashley National Forest and U.S.D.I. Bureau
of Land Management— Vernal District. 1986.

A good local flora makes field study of plants easier. The more local, the easier,
and probably better.

This is an excellent local flora. It treats an area of almost 40,000 km? that contains
about 1650 species. The locale is in northeastern Utah, including the east-west trend-
ing Uinta Mts., the Uinta Basin and the adjoining Tavaputs Plateau, and a strip of
adjacent Colorado about 80 km wide. Elevations range from 1288 m (4255 ft) on
the Green River to 4130 m (13,528 ft) on Kings Pk. The plant communities of the
area are correspondingly diverse, from cold desert and lowland riparian to alpine.
Geological diversity further complicates the mosaic of habitats and plant commu-
nities. These habitats are described succinctly, accurately, and undogmatically by
means of some of the plant species they contain. For individual species, habitats and
abundance are described, again simply but directly. Altitudinal limits for species are
given. According to the authors, the book is intended for field identification and extra
keys using vegetative or fruit characters are provided often. One feature is interpolated
keys that separate easily confused species such as Carex elynoides and Kobresia
myosuroides. The book has a glossary, references cited, and index.

A nomogram of feet versus meters would be useful. No one outside the U.S. is
going to put up with the English system of measures. We in the U.S. are stuck with
altitudes in feet so long as the U.S. Geological Survey topographic maps continue
their backward progress.

The U.S. Forest Service has been taking some hard knocks lately for overcutting
timber, roading areas to sell timber instead of developing logging methods not re-
quiring so many roads, foot-dragging on Wilderness classification, neglecting trails,
pushing resort and helicopter skiing instead of back-country, etc. The BLM was
originally set up to respond to local grazing interests, and only recently has it been
able to even start an inventory of its forage resources. The Forest Service usually acts
as local people want it to act. Add a Washington administration that is compulsively
anti-conservationist and the public lands in the National Forests and those admin-
istered by the BLM are in deep trouble. So is the Forest Service itself. The “good
guys” really need overt support when they do something that furthers good land
management. Goodrich and Neese’s book is just such a good thing. It is invaluable
for managing vegetation, either timber or range forage, because it provides necessary
help for the first question a land manager asks, What is out there? Ecological rela-
tionships can be known only when vegetation is known, and acquaintance with the
flora is necessary to understand the ecology of vegetation.

The Uinta Mts. and its environs is a beautiful, rich, interesting ecosystem. This
book will be a treasured aid to botanists sampling that ecosystem.—JACK MAJor,
Botany Dept., Univ. California, Davis 95616.

Volume 34, Number 2, pages 77-172, published 30 June 1987

MaproNno, Vol. 34, No. 2, p. 172, 1987

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CALIFORNIA BOTANICAL SOCIETY

VOLUME 34, NUMBER 3 GR JULY-SEPTEMBER 1987

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Cok :

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

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| Pee: ~ Sy,
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Contents 9

PLANT COMMUNITIES OF RING MOUNTAIN PRESERVE, Nea COUNTY, “CALIFORNIA /
Peggy Lee Fiedler and Robert A. Leidy ren 173

_ VEGETATION OF THE BALD HILLS OAK WOODLANDS, REDWOOD National! PARK, ©
CALIFORNIA Serereggpemnepreens
Neil G. Sugihara, Lois J. Reed, and James M. Lenihan 193
A FLORA OF VINA PLAINS PRESERVE, TEHAMA COUNTY, CALIFORNIA
— Pauleen Broyles 209

Soliva (ASTERACEAE: ANTHEMIDEAE) IN CALIFORNIA
Martin F. Ray 228

_ ROLE OF FIRE IN THE GERMINATION OF CHAPARRAL HERBS AND SUFFRUTESCENTS
Jon E. Keeley and Sterling C. Keeley 240

~ SOME DEMOGRAPHIC AND ALLOMETRIC CHARACTERISTICS OF Acacia smallii
(MIMOSACEAE) IN SUCCESSIONAL COMMUNITIES
J. K. Bush and O. W. Van Auken 250
MYCORRHIZAE ASSOCIATED WITH AN INVASION OF Erechtites glomerata (ASTERACEAE)

ON SAN MIGUEL ISLAND, CALIFORNIA
William L. Halvorson and Richard E. Koske 260

NOTEWORTHY COLLECTIONS

BRITISH COLUMBIA 268

_ New Mexico 269
REVIEWS 269
_NEW MADRONO POLICY 208
- ANNOUNCEMENTS 192, 249, 259

| CBS MEETING PROGRAM FOR 1987-88 272

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
ciety, Inc., and is issued from the office of the Society, Herbarium, Life Sciences
Building, University of California, Berkeley, CA 94720. Subscription rate: $25 per
calendar year. Subscription information on inside back cover. Established 1916.
Second-class postage paid at Berkeley, CA, and additional mailing offices. Return
requested. POSTMASTER: Send address changes to James R. Shevcck, Botany Dept.,
California Academy of Sciences, San Francisco, CA 94118.

Editor—WAyNnE R. FERREN, JR.
Associate Editor—BARRY D. TANOWITZ

Department of Biological Sciences
University of California
Santa Barbara, CA 93106

Board of Editors
Class of:

1987—J. RZEDOwSKI, Instituto de Ecologia, A.C., Mexico
DoroTHY DOUGLAS, Boise State University, Boise, ID
1988—SusAn G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989— FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991—Davip J. KEIL, California Polytechnic State University, San Luis Obispo
JAMES HENRICKSON, California State University, Los Angeles

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1987-88

President: DALE MCNEAL, Department of Biological Sciences, University of the
Pacific, Stockton, CA 95211

First Vice President: PEGGY FIEDLER, Department of Biology, San Francisco State
University, 1600 Holloway Ave., San Francisco, CA 94132

Second Vice President: STEVEN TIMBROOK, Ganna Walska Lotusland Foundation,
695 Ashley Rd., Montecito, CA 93108

Recording Secretary: V.THOMAS PARKER, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
Francisco, 50 Phelan Ave., San Francisco, CA 94112

The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, FRANK ALMEDA, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118; the Editor of MADRONO; three elected
Council Members: ANNETTA CARTER, Department of Botany, University of Califor-
nia, Berkeley, CA 94720; JOHN MooriInGc, Department of Biology, University of
Santa Clara, Santa Clara, CA 95053; BARBARA ERTTER, University Herbarium, De-
partment of Botany, University of California, Berkeley, CA 94720; and a Graduate
Student Representative, NIALL F. MCCARTEN, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132.

PLANT COMMUNITIES OF RING MOUNTAIN PRESERVE,
MARIN COUNTY, CALIFORNIA

PEGGY LEE FIEDLER
Department of Biology, San Francisco State University,
San Francisco, CA 94132

ROBERT A. LEIDY
U.S. Environmental Protection Agency, 215 Fremont Street,
San Francisco, CA 94105

ABSTRACT

Vegetation on Ring Mountain Preserve, Marin Co., California, was sampled and
plant communities were determined according to natural but dynamic boundaries.
Six plant communities were identified primarily by their dominant life form: ser-
pentine bunchgrass, mixed broadleaf evergreen forest, non-native grassland, northern
coyote brush, freshwater marsh, and freshwater seep. Species composition, percent
cover, and frequency of occurrence were recorded, and the percentage of the total
flora and areal cover of exotic species was determined for each community. The
serpentine bunchgrass community occupied the greatest areal extent and contained
the smallest percentage of exotic species, both in frequency of occurrence and cover.
In contrast, the non-native annual grassland community, although second in areal
coverage, was dominated by exotic grasses and broad-leaved forbs. The remaining
four plant communities are lesser components of the total vegetation. These vary
greatly in percent cover and frequency of exotic species, which in turn are dependent
on the phase of the community and the degree of past disturbance.

Ring Mountain Preserve (RMP) is an isolated remnant of the
perennial bunchgrass communities that formerly characterized much
of the San Francisco Bay landscape. Established by The Nature
Conservancy (TNC) in 1983, the Preserve is located in southern
Marin Co., California, and covers approximately 148 hectares on
the summit of the Tiburon peninsula (Fig. 1). Elevation ranges from
1-200 m above sea level.

Ring Mountain is distinguished by its extensive serpentine soils
that were derived from ultramafic intrusions into the widespread
Franciscan sandstone formation (Taliafero 1943). These soils tend
to be thin and rocky and are accompanied by glaucophane schist
outcrops. They support a highly endemic flora and are accompanied
by bunchgrasses typical of deeper ultramafic soils. Seven rare plant
species are found on this preserve (Smith and York 1984, Cal. Dept.
Fish and Game 1986). Calochortus tiburonensis Hill (Tiburon mar-
iposa) is endemic to Ring Mountain.

Although much has been written about the ultramafic soils and
vegetation of California (e.g., Kruckeberg 1984), little has been pub-
lished specifically on the plant communities of the Tiburon penin-

MADRONO, Vol. 34, No. 3, pp. 173-192, 1987

174 MADRONO [Vol. 34

Pacific

Ocean
Vicinity Map
O 10 25
kilometer

Fic. 1. Location of Ring Mountain Preserve in the San Francisco Bay area.

sula. Penalosa (1963) and Howell (1970) contain general descriptions
of the major plant communities of the Tiburon peninsula and Marin
Co., respectively. Certain unique features of Ring Mountain have
been published (e.g., Ellman 1975), as well as general descriptions
of Coastal Range vegetation types (Barbour and Major 1977).

Climate at RMP is mediterranean. Summer fog with strong winds
is frequent; annual precipitation averages 71 cm, occurring mostly
between November through April (Rantz 1971).

The objectives of this study were: (1) to characterize the vegetation
of RMP; (2) to assess the degree of human disturbance evidenced
by the percentage of exotic species that comprise each plant com-
munity; and (3) to provide data for the future management of the
plant communities at the Preserve.

METHODS

Plant communities were determined using the California natural
community classification system of the Natural Diversity Data Base

1987] FIEDLER & LEIDY: RING MOUNTAIN PRESERVE 175

(Jensen and Holstein 1983). This vegetation classification, a recent
improvement on the more widely used system of Cheatham and
Haller (1975), was used because of its wide acceptance and usage by
TNC and public organizations. The terms plant community and
vegetation type are used interchangeably throughout the text, and
one term is not used preferably over the other.

Plant communities were mapped for RMP from March through
July 1983 from aerial photographs taken the same spring. These
communities were delineated first on acetate overlays on aerial con-
tact and composite photographs, and then verified in the field. Areal
extent of each community was determined with a Lasico compen-
sating polar planimeter (Series 40). Samples of each unit were located
randomly on a working map for study in the field.

Each community was sampled from March through May 1984
and April 1986. As a result of an early spring sampling period, some
conspicuous species that flower later in the spring and summer (e.g.,
Danthonia californica) were not recorded. Species composition,
ground slope, aspect, and topographic position were obtained for
each community. For the serpentine bunchgrass, non-native grass-
land, freshwater marsh, and freshwater seep communities, a stan-
dard 1% areal sample was achieved, using the point intercept method
(Mueller-Dombois and Ellenberg 1974). Sampling transects were run
along slope contours from randomly chosen points along a line-
transect that was oriented perpendicular to the contours. The north-
ern coyote brush community was sampled by the line-intercept
method (Canfield 1941) to achieve a 5% sample of the total area.

Random points were located within a random sample of the mixed
broadleaf evergreen forest, and data were obtained by the point-
centered quarter method (Mueller-Dombois and Ellenberg 1974). A
5% areal sample was attained for this community. At each point,
data on tree and sapling composition, shrub species composition,
and percent cover were estimated on four 25 m? subplots. For herb
species, percent cover was estimated on four 5 m? plots at each tree
subplot. Botanical nomenclature follows that of Munz (1968) with
only a few exceptions.

After field sampling, the serpentine bunchgrassland, non-native
grassland, northern coyote brush, and mixed broadleaf evergreen
forest communities were divided into ‘phases’ according to aspect,
topographic position, ground slope, and canopy layer and represent
variations within community types.

Percent exotic species within the flora of each community and
percent total plant cover by exotic species for each community were
computed and tested with a chi-square statistic with a continuity
correction (Grieg-Smith 1983). This test determined whether any
significant differences in the extent of exotic species within the var-
ious phases of each vegetation type occurred. The Jaccard Index was

176 MADRONO [Vol. 34

= rRESHWATER MARSH
NORTHERN COYOTE BRUSH
WM -ResHwaTeR SEEP

vc] MIXED BROADLEAF EVERGREEN
FOREST

[__] SERPENTINE BUNCHGRASSLAND
E NON-NATIVE GRASSLAND

(0) 15 30
t———_—_—__+4
kilometers

Fic. 2. Distribution of the six plants communities on Ring Mountain Preserve.

computed to examine the floristic similarities among communities
(Grieg-Smith 1983).

RESULTS

Six communities were identified on RMP: serpentine bunchgrass
(SBG), non-native (annual) grassland (NNG), mixed broadleaf ev-
ergreen forest (MBEF), northern coyote brush (NCB), freshwater
marsh (FWM), and freshwater seep (FWS) (Fig. 2). The SBG com-
munity is the dominant vegetation type, representing 70 ha (47% of
the total area). NNG is slightly less extensive on RMP, comprising
49 ha (33%) that are primarily on the lower slopes.

MBEF covers approximately 16 ha (11%) that occur in irregularly
scattered stands throughout the Preserve. The FWM, FWS, and NCB
communities are of much smaller distribution. These represent 1.4
ha (<1%), 4.7 ha (3%), and 7.3 ha (5%) of RMP, respectively.

Serpentine bunchgrass. This community occurred most frequently
on the upper slopes and ridgetops (Fig. 3). Ground slope varied from
zero at the upper elevations to greater than 25% at midslope. Fifty-
six species that represent a predominately native flora were recorded
in this plant community (Table 1). The exotic Lolium multiflorum

1987] FIEDLER & LEIDY: RING MOUNTAIN PRESERVE 177

Fic. 3. Serpentine bunchgrass community type, which is typified by many boul-
ders strewn among various perennial grasses and bulbous plant species.

Lamk. and the ubiquitous native Chlorogalum pomeridianum (DC.)
Kunth were the species most frequently encountered. Perennial
bunchgrasses, particularly Stipa pulchra Hitche. and Melica torrey-
ana Scribn., also were abundant. A high frequency (14%) of exposed
bedrock and boulders typified this vegetation type on RMP.

The richness of the flora is shown by the low frequency of occur-
rence of most herbs. A little greater than 71% (40 species) of the
flora occurred less than 1% of the time, whereas no species occurred
more frequently than 13%. Infrequent species were represented by
both annual and perennial native forbs.

All seven rare species on RMP were found in this community,
but only those most abundant in the early and mid-spring were
recorded during this sampling period. These species are: Calochortus
tiburonensis Hill, C. umbellatus Wood, Castilleja neglecta Zeile,
Calamagrostis ophitidis (J. T. Howell) Nygren, Eriogonum caninum
(Greene) Munz, Hesperolinon congestum (Gray) Small, and Hemi-
zonia congesta DC. subsp. vernalis Keck. Of these, Calamagrostis
ophitidis is most frequently encountered. Eriogonum caninum was
localized throughout the Preserve, and not represented in the sample.
Hesperolinon congestum also was not recorded. Although wide-
spread, it is rarely abundant locally.

Mixed broadleaf evergreen forest. This community was found in
the most mesic habitats on RMP. These include the drainages near
groundwater seeps (Fig. 4), and the large rock outcrops (Fig. 5).
Species composition in these two habitats were similar, and, thus,
MBEF was treated as a relatively homogeneous plant community
that is characterized by three canopy layers.

[Vol. 34
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183

FIEDLER & LEIDY: RING MOUNTAIN PRESERVE

1987]

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184 MADRONO [Vol. 34

Fic. 4. Mixed broadleaf evergreen forest community, drainage phase. Note the
linear aspect of this type. Darker regions in the grasslands are the freshwater seeps.

A small grove of eucalyptus (Eucalyptus globulus Labill.) was found
on the west-southwest arm of RMP, and is mapped as part of the
MBEF community. Eucalyptus is not considered a natural plant
community (Jensen and Holstein 1983), represents only a small
portion of the preserve, and was omitted from the sample.

The tree layer of MBEF contains four species: Quercus agrifolia
Nee, Lithocarpus densiflora (H. & A.) Rehd., Umbellularia califor-
nica (H. & A.) Nutt., and Arbutus menziesii Pursh (Table 1). The
latter two species occurred slightly less frequently (26.3% and 10.5%,
respectively) than the former two (31.6% and 31.6%, respectively).
Other tree species typical of the mixed broadleaf evergreen forest

Fic. 5. Mixed broadleaf evergreen forest, rock outcrop phase, surrounding large,
often exposed boulders, and typically found on the upper slopes of Ring Mountain.

1987] FIEDLER & LEIDY: RING MOUNTAIN PRESERVE 185

Fic. 6. Northern coyote brush community, found above the broadleaf evergreen
forests, on the lower slopes of the Preserve and adjacent urban areas.

(cf. Sawyer et al. 1977), such as Aesculus californica (Spach) Nutt.,
are infrequent on RMP, and were not recorded in this sample.

Sapling frequency differed from that of tree frequency. Umbellu-
laria californica was the most frequent tree species as a sapling
(52.6%), whereas the remaining tree species were at a lower frequency
as saplings (Q. agrifolia 15.8%; L. densiflora 21.1%). Arbutus men-
ziesii was absent from the understory as a sapling.

The shrub layer of MBEF was more species rich than its overstory.
Those species found in this layer include: Corylus cornuta Marsh.
var. californica (A. DC.) Sharp, Physocarpus capitatus (Pursh) Ktze.,
Pteridium aquilinum (L.) Kuhn var. pubescens Underw., Rhamnus
californica Esch., Rubus ursinus C. & S., Symphoricarpos rivularis
Suksd., and Toxicodendron diversiloba Greene (Table 1). Toxico-
dendron diversiloba was a common understory species (100% fre-
quency) and represented 19% of the total understory shrub cover.
Corylus cornuta var. californica was less frequently encountered
(16%), and its mean cover was slightly less than 14%. Conversely,
S. rivularis was common in the understory (52%), but its average
cover was typically low (8%). The remaining four species were both
sparse in coverage and infrequent in occurrence (Table 1).

The herb layer of this community was more species rich than
either the tree or shrub layers. Weedy species characteristic of a
variety of habitats have invaded this herbaceous layer. Eighteen herb
species were recorded in the MBEF flora. Avena spp. showed the
highest mean cover (37%). Eighty-three percent of the species showed
less than 1% areal cover and reflected the openness of this stratum.
The range in frequency of occurrence for most herbs was 5—10%
(Table 1).

186 MADRONO [Vol. 34

a ee EE ub
i rhc na a

Fic. 7. Non-native annual grassland lacks obvious rock outcrops and is charac-
terized by the smooth texture of the grassland slopes. Note coyote brush and mixed
broadleaf evergreen forest in the drainage at center.

Northern coyote brush. The NCB community was found primarily
along the lower to middle slopes of RMP, although it also occurred
at the higher elevations within the eastern boundaries of the Preserve
(Fig. 6). This community was found typically on slopes that ranged
from 20-30%. Baccharis pilularis DC. subsp. consanguinea (DC.)
C. B. Wolf comprises between 41-59% of the total cover, depending
on the slope location (Table 1). Within this shrub community, the
NNG community occupied slightly less of the total cover (31-45%).
The remaining species recorded in this vegetation type are largely
broad-leaved forbs (e.g., Chlorogalum pomeridianum, Carduus pyc-
nocephalus L.) that are distinctive, locally abundant members of the
grassland communities. Additional shrub species (e.g., Thermopsis
macrophylla H. & A., Toxicodendron diversiloba, and Rosa califor-
nica C. & S.), were recorded occasionally, and do not represent a
significant component of the community.

With the exception of C. pomeridianum, there were few elements
typical of the serpentine bunchgrass community in this vegetation
type. Those that were found (e.g., Melica californica Scribn.) occur
in sparse amounts in the Baccharis scrub and only on the upper
slopes of RMP.

Freshwater seep. The FWS community lines many of the ephem-
eral drainages, mostly on the southern portion of RMP (Fig. 4). This
vegetation type is characterized by the presence of surface water,
although some drainages typically dry completely by June.

Carex densa Bailey and Juncus phaeocephalus Engelm. var. pa-
niculatus Engelm. were the most frequently encountered species in
the freshwater seeps. No species occurred more frequently than 20%.
Some exotic grasses such as Polypogon monspeliensis (L.) Desf.,

1987] FIEDLER & LEIDY: RING MOUNTAIN PRESERVE 187

elie

a bes 7 ‘ ‘ ee a= eau ae Peyeee 4
ie Se nS PS ; A ORIOL Re ERTL Demet Mal oot, rf a Joby
Byhcse (aay \ Re yer OV OAR Set: mR popcugiey SP gts he a 4
wi fi . “ > . o a Salah tec, V oar claly Rig Dh %
ee" ny,

Fic. 8. Freshwater marsh is characterized by the light-colored, exotic grass Festuca
arundinacea in the center and by the ring of dark-colored, native species.

Cynodon dactylon (L.) Pers., and Cortaderia selloana (Schult.) Asch.
& Graebn., have established extensively throughout these seep areas.

Non-native grassland. NNG is the second largest plant community
in areal extent (Fig. 7). This grassland was found on all aspects, with
ground slope varying from less than 5% to nearly 25%. South- and
north-facing slopes appeared different with respect to species com-
position (Table 1). For example, southern slopes of the non-native
grassland had a relatively high concentration of exotic weedy species
[e.g., Carduus pycnocephalus (9.27%); Bromus diandrus Roth.
(4.30%); Briza maxima L. (29.16%); Silybum marianum (L.) Gaertn.
(0.16%)], whereas the northern slopes still harbored remnants of the
SBG community [e.g., Stipa pulchra (3.08%); Melica californica
(0.44%); Calochortus umbellatus (2.20%)].

Of the 43 plant species recorded for this grassland, more species
exhibited higher frequencies of occurrence than in SBG. For ex-
ample, in NNG we recorded a 20% frequency for Briza maxima
and Lolium multiflorum (Table 1). By contrast, the most abundant
species in the native SBG community was Lolium multiflorum at
13%. Both grasslands had an approximately equal portion of the
flora exhibiting less than 1% frequency of occurrence (67% for NNG;
70% for SBG). The NNG community, however, is extremely vari-
able throughout the preserve, and is difficult to characterize gener-
ally.

Freshwater marsh. This community occurs at the northern en-
trance to the Preserve (Fig. 8). It is a permanent wetland of less than
5% slope, and historically may have been of brackish or saltwater
origin (Josselyn, pers. comm.). It is now separated from an adjacent
tidal marsh by a road.

188 MADRONO [Vol. 34

TABLE 2. MATRIX OF THE JACCARD COEFFICIENT OF SIMILARITY FOR THE SIX PLANT
COMMUNITIES OF RING MOUNTAIN PRESERVE. SBG = serpentine bunchgrass; NNG =
non-native grassland; MBEF = mixed broadleaf evergreen forest; NCB = northern
coyote brush; FWM = freshwater marsh; FWS = freshwater seep.

SBG NNG MBEF NCB FWM FWS
SBG — 24.69 6.17 4.23 137 9:59
NNG — _ P23 15.09 202 1552
MBEF — — — 15.00 0.00 bey
NCB _ _ _ — i 13.33
FWM _ — — — — 5.26

The freshwater marsh was dominated by the exotic Festuca arun-
dinacea (26%; Table 1), whereas the marsh periphery was surround-
ed by a variety of freshwater plants (e.g., Carex spp., Typha angus-
tifolia L.; Table 1) that occurred less commonly. Several ubiquitous
exotic species (e.g., Rumex acetosella L., Picris echioides L.) occurred
frequently in this community (Table 1).

Community similarities. A simple measure of floristic similarity
among the five plant communities on RMP was computed using
Jaccard’s coefficient of similarity (Grieg-Smith 1983; Table 2). In
general, overall similarities are moderate.

The two grassland communities, SBG and NNG, exhibited the
greatest similarity among all the plant communities present on RMP.
The SBG community, however, did not share many species with
any of the other plant communities. NCB was most similar to MBEF
and NNG, two communities with which it interdigitates. NNG also
showed similar floristic affinities with FWS, the plant community
typically found in the wetter portions of this exotic grassland. MBEF
did not appear to share many species with the other vegetation types
other than NNG. Finally, FWM showed little floristic affinity to any
of the other plant communities on RMP.

Exotic species. Percent total number of species and total cover of
each plant community was tested to determine whether the different
phases of each type differed significantly. The percentage of exotic
flora differed greatly among the six plant communities (Table 3).

Less than 14% of the total flora of the SBG community was exotic,
whereas NNG possessed 44% exotic species. The NCB, MBEF, and
FWM communities possessed 32%, 33%, and 41% exotic species,
respectively (Table 3).

The NCB community differed significantly among its phases in
percentage of exotic flora. The upper slope possessed 9% exotic
species, midslope 50%, and the lower slope 38%. All slope sections
were significantly different from each other (Table 3).

MBEF also showed a significant difference in percentage of exotic

1987] FIEDLER & LEIDY: RING MOUNTAIN PRESERVE 189

TABLE 3. PERCENT OF TOTAL FLORA AND COVER OF ExOTIC SPECIES FOR EACH
PLANT COMMUNITY AT RING MOUNTAIN PRESERVE. * = p < 0.05; ** = p < 0.01;
x? = chi-square statistic with a continuity correction.

No.
exotic % flora % cover
Plant community species exotic exotic x? (flora) x* (cover)
Serpentine bunchgrass
North slope (N) 6 12.50 2.76 N vs. S N vs. S
0.4033 2.4061
South slope (S) 3) 16.67 8.90 Svs. R S vs. R
0.0076 2.1141
Ridgetop (R) 7 15322 3.03 N vs. R N vs. R
0.1239 0.0948
Total 8 13.79 1.26 — —
Non-native grassland
North slope (N) 9 32.14 55.50 N vs. S N vs. S
2.7020 18.8316**
South slope (S) 16 44.44 84.60 _ —
Total 19 44.19 80.30 — —
Mixed broadleaf evergreen forest
Tree layer (T) 0 0.00 0.00 T vs. S T vs. S
Shrub layer (S) 0) 0.00 0.00 S vs. H S vs. H
37.6313** 85.3261**
Herb layer (H) 6 33.33 61.20 T vs. H T vs. H
37631327 85.3261**
Total 6 33.33 61.20 _ _
Northern coyote brush
Upper slope (U) 1 9.11 31.03 U vs. M U vs. M
38:2135** 6.3445*
Midslope (M) 5 50.11 49.50 M vs. L M vs. L
2.2346** 0.8777
Lower slope (L) 5 38.50 41.90 L vs. U L vs. U
22:21814t 2.1024
Total 31.60 38.90 — —
Freshwater marsh 7 41.18 45.73 — —
Freshwater seep 6 26.09 14.05 — —

flora between the herb and shrub layers, and between the herb and
tree layers. These differences were the result of an entirely native
tree and shrub layer, and an herb layer with a rather high component
of exotic species. Finally, neither of the other two communities tested
(SBG, NNG) differed significantly between community phases with
respect to percent of exotic flora.

When the plant communities were analyzed according to the per-
centage of the total cover occupied by exotic species, a different
pattern emerged. NCB showed no differences between cover occu-

190 MADRONO [Vol. 34

pied by exotics among the three slope positions, whereas the north
and south exposures of NNG were significantly different. The south-
ern slopes of this grassland had a much greater cover by exotics than
the northern slope (84% and 55%, respectively; Table 3). As ex-
pected, in MBEF, the herb layer in this community had significantly
greater cover by exotic species than either the shrub or tree layer.

DISCUSSION

The flora of ultramafic soils has been a perennial source of interest
to California botanists. The literature concerning ultramafic com-
munities in California recently has been summarized by Kruckeberg
(1984). He suggested that the San Francisco Bay region does not
harbor unusual range extensions of ultramafic species. We have found
that the flora of RMP provides no exceptions.

The SBG community of RMP is a diverse, native, perennial grass-
land that is important because of its rather undisturbed condition
in which seven rare plant species occur. In an areal context, the
native SBG type remains the dominant plant community on the
Preserve. The small percentage of exotic species in this community,
both with respect to percent frequency of occurrence and cover,
contrasts sharply with the adjacent NNG community.

The NNG community harbors certain bunchgrass species of SBG,
such as Stipa pulchra and Melica californica, but it remains domi-
nated by opportunistic exotic grasses and broad-leaved forbs. Cer-
tain areas, such as the colluvial soils and shaded portions of this
community, are dominated by Lolium multiflorum, a grass that
presumably was seeded when Ring Mountain was managed privately
for grazing by dairy cattle during the 1950’s (Lozier, pers. comm.).
Although the variability of this grassland is extensive, the dominant
species (L. multiflorum, Briza maxima, and Plantago lanceolata L.)
are predictably the same for the two slope phases. On RMP, the
NNG community most clearly reflects the mountain’s recent land
use history of cattle grazing, fire suppression, and off-road vehicular
use.

MBEF represents a smaller portion of the RMP flora, and will
likely remain so because of its restriction to the drainages and larger
rock outcrops. Species composition of the tree and shrub portions
of this community is typical of the mixed evergreen forests through-
out the North Coast Ranges of California (Sawyer et al. 1977). The
herbaceous stratum, however, consists largely of exotic species.

The FWS community is significant in that it reflects the complex
geology of Ring Mountain, where ultramafic intrusion, shearing, and
orogenic activities have created a complex soil and bedrock profile.
Typical freshwater plants are found in the seeps and areas where
ground water is not visible, but is present immediately below the

1987] FIEDLER & LEIDY: RING MOUNTAIN PRESERVE ee

soil surface. Several of the seeps have been disturbed greatly by
cattle grazing and more recently by off-road vehicular trafic. We
suggest that the high proportion of exotic species in this community
is the result of such long-term disturbances.

FWM contains a flora that is distinctive compared with the re-
maining preserve. This marsh, historically a brackish wetland bor-
dering San Francisco Bay, is no longer influenced by tidal cycles.
Therefore, under present hydrologic conditions, it 1s expected that
this community will maintain a flora of exotic and native species
typical of disturbed freshwater marshes.

NCB may be a recently established plant community in the study
area. Its probable origin lies in the cessation of grazing by cattle and
of fires set by Native Americans (Lewis 1973). We have observed
this as a mid-successional sere that is invading NNG. Extensive
areas of NNG interdigitating with NCB support the observation that
B. pilularis subsp. consanguinea is invading and changing the com-
position of the NNG community. Baccharis cover is rarely greater
than 50% cover anywhere along the hillsides. Under existing con-
ditions, i.e., the absence of fire and grazing, percent cover of coyote
brush is likely to increase on the lower slopes.

The importance of RMP lies not only in its distinctive floristic
composition, but also in the relatively pristine condition of the SBG
community. The Preserve’s flora, however, reflects both its unusual
geology and its history of cattle grazing, fire suppression, and other
recent forms of site disturbance. Management of this preserve will
require baseline information as provided in this community study.
Finally, periodic monitoring to record changes in the structure and
composition of the vegetation will provide information to support
future management decisions.

ACKNOWLEDGMENTS

An earlier version of this manuscript was submitted by PLF as a consultant’s report
to the California Field Office of The Nature Conservancy, San Francisco. We thank
Lynn Lozier and Greg Wolley for unlimited access to the Preserve, Winifred Fiedler
for photographic assistance, and two reviewers for their comments. Because The
Nature Conservancy’s policy did not permit the collection of voucher specimens, we
do not include them here. However, Table | represents the majority of the taxa
observed on Ring Mountain. A complete list is available from the authors on request.

LITERATURE CITED

BARBOUR, M. G. and J. MAjor. 1977. Terrestrial vegetation of California. John
Wiley and Sons, New York.

CALIFORNIA DEPARTMENT OF FISH AND GAME. 1986. Designated endangered or rare
plants. The Resources Agency.

CANFIELD, R. 1941. Application of the line intercept method of sampling in range
vegetation. J. Forestry 39:388-394.

CHEATHAM, N. D. and R. HALLER. 1975. An annotated list of California habitat
types. Univ. California Natural Lands and Water Reserve System [Natural Re-
serves System].

192 MADRONO [Vol. 34

ELLMAN, P. 1975. Let’s save Ring Mountain. Fremontia 3:10-14.

GRIEG-SMITH, P. 1983. Quantitative plant ecology. Studies in Ecology, Vol. 9, 3rd
ed. Univ. California Press, Berkeley.

Howe LL, J.T. 1970. Marine flora. 2nd ed. with supplement. Univ. California Press,
Berkeley.

JENSEN, D. and G. HOLSTEIN. 1983. Natural diversity data base natural communities.
California Dept. Fish and Game.

KRUCKEBERG, A. R. 1984. California Serpentines: flora, vegetation, geology, soils,
and management problems. Univ. California Publications in Botany Vol. 78.
Univ. California Press, Berkeley.

Lewis, H. T. 1973. Patterns of Indian burning in California. Ballena Press Anthro-
pological Papers 1:1-101.

MUELLER-DomBolIs, D. and H. ELLENBERG. 1974. Aims and methods of vegetation
ecology. John Wiley and Sons, New York.

Munz, P. 1968. A California flora and supplement. Univ. California Press, Berkeley.

PENALOSA, J. 1963. A flora of the Tiburon peninsula, Marin County, California.
Wasmann J. Biol. 21:1-74.

RANTZ, S. E. 1971. Precipitation depth-duration-frequency relations for the San
Francisco Bay Region, California, with isohyetal map showing mean annual
precipitation. U.S.D.I., Geological Survey, Professional paper 750-C.

SAWYER, J. O., D. A. THORNBURGH, and J. R. GRIFFIN. 1977. Mixed evergreen
forest. In M. G. Barbour and J. Major, eds., Terrestrial vegetation of California,
p. 359-381. John Wiley and Sons, New York.

SMITH, J. P., JR. and R. YORK. 1984. Inventory of rare and endangered vascular
plants of California. California Native Plant Society Special Publication No. 1
(3rd ed.), Berkeley.

TALIAFERO, N. L. 1943. Franciscan-Knoxville problem. Bull. Amer. Assoc. Petro-
leum Geol. 27:109-219.

(Received 14 May 1985; revision accepted 13 Jan 1987.)

ANNOUNCEMENT

NEw PUBLICATIONS

Bowers, J. E., A career of her own: Edith Shreve at the Desert Laboratory,
Desert Plants 8:23—29, 1986. [Interesting account of Shreve (1878-1956), whose
career in plant physiology was sidetracked by motherhood and, typical for
those times, womanhood. Shreve’s husband was Forrest Shreve, author with
I. L. Wiggins of Vegetation and Flora of the Sonoran Desert, 1964.]

CARTER, A., Aspectos generales de la flora de Baja California, Cactaceas y
suculentas Mexicanas 31:79-96, 1986. [With 7-page English version. Since the
submission of this article some years ago, various nomenclatural changes were
made that have not been brought up to date in this paper; nor did the author
have the opportunity of including in the bibliography I. L. Wiggin’s Flora of
Baja California (1980), which was published after the paper was written. In
addition, because of the tremendous time lag for mail between Mexico and
the United States, there was no opportunity to correct page proof—hence such
inadvertencies as Hystis for Hyptis (p. 88) and Dalia for Dalea (p. 89).]

VEGETATION OF THE BALD HILLS OAK WOODLANDS,
REDWOOD NATIONAL PARK, CALIFORNIA

NEIL G. SUGIHARA and LoIs J. REED
Redwood National Park, P.O. Box 7, Orick, CA 95555

JAMES M. LENIHAN
Department of Geography, Oregon State University,
Corvallis 97331

ABSTRACT

Composition and structure are determined for stands of bald hills Quercus garryana
(Oregon white oak) woodland in Redwood National Park, California. Seven distinct
plant community types are found. Distribution of the three most widespread types
is related to moisture, slope position, and fire history: 1) Quercus/Cynosurus (xeric
woodlands); 2) Quercus/Dactylis (mesic woodlands); and 3) Quercus/Symphoricarpos
(dense, young mesic stands). Four types occupy specific habitats within the park: 4)
Ribes/Phacelia (rock outcrops); 5) Arrhenatherum/Sherardia (glades); 6) Quercus/
Delphinium (seasonally moist areas within xeric woodlands); and 7) Philadelphus/
Cystopteris (stream channels).

Quercus garryana Dougl. (Oregon white oak) ranges from British
Columbia to the Santa Cruz Mountains of California (Griffin and
Critchfield 1972). Optimum development is reached in the Willam-
ette Valley of Oregon, where Q. garryana dominates oak woodlands
that occupy over 400,000 ha (Franklin and Dyrness 1973). In the
North Coast Ranges of California, Q. garryana dominates the north-
ern oak woodland and is a minor component of several forest types.
The northern oak woodland consists of two distinct elements, a
continuation of the interior foothill woodland and a coastal com-
munity type that is structurally distinct and known as “‘bald hills”
oak woodlands (Griffin 1977).

Bald hills oak woodlands occur in the Coast Ranges of California
from Humboldt and Trinity cos., southward to Napa Co. Approx-
imately 19% of this area supports oak or oak-grassland vegetation
(Wieslander and Jensen 1948, Storie and Wieslander 1952). Al-
though the woodlands occur between 75—1600 m elevation through-
out the region, they are best developed along ridgetops and upper
south-facing slopes in Humboldt and Mendocino cos. The bald hills
oak woodlands are structurally distinguished by a patchy mosaic
pattern of dominance by either oak or grasses and not the balanced
mixture of oaks and grassland found elsewhere in California (Clark
1937).

Thilenius (1968) describes the vegetation of Willamette Valley
oak woodlands as seral and derived from open oak savannahs by

MADRONO, Vol. 34, No. 3, pp. 193-208, 1987

194 MADRONO [Vol. 34

edwood National Park

0 10 20

ote ee cee eel
Kilometers

— Study Area

Fic. 1. Location of the study area within Redwood National Park and California.

the exclusion of fire. He defines four plant communities primarily
by their shrub layers. Sugihara et al. (1983) found three stand struc-
tural types in the oak woodlands of Redwood National Park (RNP):
1) open savannah stands composed of all size classes and dominated
by a few, large, widely scattered individuals; 2) closed-canopy stands
of numerous, uniformly medium-sized, clustered trees; and 3) dense
closed-canopy stands with uniformly small, single-stemmed indi-
viduals. Hektner et al. (1983) describe the vegetation composition
and dynamics after disturbance of the open bald hills grassland
outside the oak-grass mosaic. No other studies describe the flora or
vegetation succession of the coastal bald hills. The purpose of this

1987] SUGIHARA ET AL.: BALD HILLS WOODLANDS 195

Tact

pats SR UR i ae eS a ee
hs i eames
i
‘

my
; i. ot Ute Ae Ser
Fe cL Wa aie) Lats

Fic. 2. A view, looking northwest from Schoolhouse Peak, of the oak/grass mosaic
and study area. Best development of the woodland is near the ridges, with redwood
forests downslope.

study is to describe and classify the present vegetation in the northern
range of the bald hills oak woodlands. Results of this study of present
conditions will contribute to the potential for restoration and main-
tenance of the woodlands as a natural ecosystem. These baseline
data will help assess the effects of the future management of the
ecosystem.

STUDY AREA

Location. The 250 ha study area (Figs. 1, 2) is located within the
Redwood Creek basin of RNP in Humboldt Co. and is representative
of the northern extent of the bald hills oak woodlands. The area lies
8-22 km from the coast and 85-95 km south of the Oregon border.
Redwood Creek flows from the southeast to the northwest and emp-
ties into the ocean at Orick, California. The grassland/woodland
mosaic ranges in elevation from the banks of Redwood Creek (75
m) to near the top of Schoolhouse Peak (945 m) on the northeast
slope above Redwood Creek.

Climate. Regional climate is Mediterranean, with strong oceanic
influence at lower elevations in the northwest portion of the study
area where summer fog frequently occurs. Approximately 90% of
the total annual precipitation falls between October and May. Av-
erage annual rainfall ranges between 178 cm and 203 cm with snow

196 MADRONO [Vol. 34

rarely falling except at higher elevations (Coghlan 1984). The mean
daily maximum temperature in July is 25°C with absolute maxima
rarely exceeding 38°C. The mean daily minimum in January is 2°C.
Absolute minima rarely go below —7°C (Humboldt State Univ.
1974).

Geology and soils. Geologic substrate of the study area is residuum
and colluvium from Franciscan siltstone, sandy siltstone, and gray-
wacke sandstone. The landscape is characterized by numerous earth-
flows. A complex pattern of Inceptisol, Alfisol, and Ultisol soils
underlies the woodlands, adjacent forests, and grasslands. Subsoil
properties largely reflect geologic substratum and relief. Forest soils
lack the umbric epipedon found in woodland and prairie soils, but
all of these subsoils have a similar range of properties. Consistent
soil patterns that correspond with forest/woodland/grassland bound-
aries have not been established (J. Popenoe pers. comm.).

Historical use. Native Americans regularly set fire to the bald hills
for at least 6000 years prior to 1864, and these fires profoundly
affected vegetation patterns (Thompson 1916, Lewis 1973, Bickel
1979, King and Bickel 1980, Benson 1983, Hayes 1985). Livestock
grazing that was initiated by European settlers resulted in the estab-
lishment of many aggressive, non-native range plants, and was dis-
continued by the National Park Service in 1982. Extensive logging
of adjacent redwood and Douglas-fir forests has been the primary
disturbance factor affecting vegetation in the Redwood Creek basin
from the 1940’s until acquisition by the Park Service in 1978.

METHODS

Field reconnaissance of the study area revealed several plant as-
semblages within the oak woodland with distinct structure and com-
position. These assemblages were sampled by placing a total of 56
relevé plots averaging 750 m in homogeneous vegetation within
uniform habitats (Mueller-Dombois and Ellenberg 1974). A list of
all vascular plant species in each plot was compiled by height strata.
Visual estimates of cover for the canopy, shrub, and herbaceous
layers, as well as of each species, were made using the Braun-Blan-
quet (1932) cover scale: 1 = <1-5%, 2 = 5—25%, 3 = 25-50%, 4 =
50-75%, 5 = 75-100%. Aspect, slope, slope position, topographic
configuration, oak stand type, exposure, and elevation of each plot
also were recorded. Sampling was completed from May-July 1983,
which coincides with the flowering and fruiting periods of most
species. Nomenclature generally follows Munz (1973). Voucher
specimens are on file at RNP.

Floristic characterization 1s based on species cover, percent oc-
currence, and fidelity. The stand classification is interpreted at a
division level where the stand groups best represented the vegetation

SUGIHARA ET AL.: BALD HILLS WOODLANDS 197

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200 MADRONO [Vol. 34

types observed in the field. One-hundred thirty-five species with
frequencies greater than 5% were entered into TWINSPAN, a com-
puter analysis procedure in the Cornell Ecology Program (CEP) series
(Hill 1979a). TWINSPAN is a program for two-way indicator species
analysis, a polythetic divisive method for community classification.
The program was run with all default options except for the definition
of pseudospecies cut-levels. Four cut-levels were defined as follows:
level 1 = 1-—5% cover, level 2 = 6—25% cover, level 3 = 26-50%
cover, and level 4 = 51—100% cover.

The classified stands and 135 species are ordinated by detrended
correspondence analysis using the DECORANA program in the CEP
series (Hill 1979b) to reveal any additional dimension of the stand-
group relationship. The DECORANA procedure was run with all
default values and options. Comparisons of physiographic data with
distribution of community types on the ordination graph and field
observations produced environmental interpretations of plant com-
munity relationships. Plant community descriptions were then de-
veloped and used to classify and map the vegetation within the study
area.

RESULTS

Three-hundred five species were found during sampling. Analysis
by TWINSPAN identified seven plant communities with three dis-
tinct structural forms: tree-dominated, shrub-dominated, and grass-
dominated (Table 1). Four communities contain Q. garryana as the
main structural unit. Two communities have shrubs composing the
main structural unit with some Q. garryana present. The other com-
munity is dominated by grasses with mature Q. garryana absent.
Within each life form category, species are ordered by relationships
to one another. In general, species with mesic habitats are followed
by those occurring in more xeric habitats. The distribution of the
classified stands in floristic ordination space is shown in Fig. 3.
Environmental features of the areas supporting the seven vegetation
types are presented in Table 2.

Descriptions of the plant communities are based on Tables 1 and
2. Vegetation types are named for a combination of two species. The
first is a dominant member of the main structural element. The
second is a characteristic species with high cover and presence. The
names reflect the mosaic pattern where oaks are either dominant or
nearly absent. Important associates are species with high modal
cover and presence.

Quercus/Symphoricarpos (Qu/Sy): This woodland type is found
mid-slope in uniform, extremely dense stands of 25—40 yr-old, small-
diametered oaks. Symphoricarpos rivularis forms most of the well
developed low shrub layer. This type has the densest understory
with the greatest number of shrub species and highest shrub cover

1987] SUGIHARA ET AL.: BALD HILLS WOODLANDS 201

YN 0=x> DOU

DCA Axis 1

Fic. 3. Ordination graph showing the floristic relationships between plots.
1 = Quercus/Symphoricarpos, 2 = Quercus/Dactylis, 3 = Quercus/Delphinium, 4 =
Philadelphus/Cystopteris, 5 = Ribes/Phacelia, 6 = Quercus/Cynosurus, 7 = Arrhen-
atherum/Sherardia.

(30%) of any oak-dominated type. Forbs dominate the herbaceous
layer with only scattered grasses present. Important associates in the
perennial forb-dominated herb layer include Polystichum munitum,
Satureja douglasii, Fragaria californica, and Bromus carinatus. Li-
gusticum aplifolium, Rubus vitifolius, Cynoglossum grande, Ceras-
tium arvense, and Festuca occidentalis are the more commonly en-
countered species that characterize this type.

Quercus/Dactylis (Qu/Da): This mesic woodland type is extensive
on lower concave slopes associated with uniform, medium-sized oak
stands. A mixture of tall, perennial grasses and perennial forbs dom-
inates the understory. The shrub layer is very sparse. Important
associates include Dactylis glomerata, S. douglasii, Osmorhiza chi-
lense, F. californica, Sanicula crassicaulis, and Vicia americana.
Galium nuttallii, Lonicera hispidula, and Stachys rigida are common
characteristic species.

[Vol. 34

MADRONO

202

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1987] SUGIHARA ET AL.: BALD HILLS WOODLANDS 203

Quercus/Cynosurus (Qu/Cy): This xeric woodland type is domi-
nated by shorter perennial and annual grasses with forb cover rel-
atively low. The shrub layer is not well developed. This xeric type
was the most heavily disturbed by grazing and occupies the upper,
convex, south-facing slopes with oak stands containing a wide range
of sizes and ages. Important associates include Cynosurus echinatus,
Holcus lanatus, Elymus glaucus, Poa pratensis, Arrhenatherum ela-
tius, S. crassicaulis. Taraxacum officinale is the most common char-
acteristic species.

Quercus/Delphinium (Qu/De): Found on upper, concave slopes
in uniform, medium-diametered oak stands, this type is restricted
to concave topography on otherwise xeric slopes. In the spring and
early summer, perennial forbs heavily dominate the understory of
this distinctive woodland type. Forbs die back and grasses become
dominant as the soil dries by mid-July, but the shrub layer remains
sparse. Delphinium trolliifolium is the strong, early season dominant
and Dentaria californica, Lithophragma affine, Claytonia perfoliata,
and Jsopyrum stipitatum are characteristic early season species. Im-
portant late season associates include Galium aparine, Melica su-
bulata, and E. glaucus.

Arrhenatherum/Sherardia (Ar/Sh): Open glades dominated by pe-
rennial and annual grasses are found as narrow openings running up
the slope within the oak stands. Shrubs are present only in scattered
patches. These glades and the surrounding oaks form the distinctive
oak/grass mosaic characteristic of the bald hills oak woodlands.
Important associates include A. elatius, H. lanatus, Festuca viridula,
C. echinatus, and Rumex acetosella. Sherardia arvense, Lotus mi-
cranthus, Viola praemorsa, Aira caryophyllea, and Bromus mollis
are the common characteristic species. All of these species also are
components of the open bald hills prairies. Many of the more abun-
dant woodland species, however, such as G. aparine, O. chilensis,
and S. crassicaulis, are absent from the Ar/Sh type.

Philadelphus/Cystopteris (Ph/Cy): This rocky stream channel type
is composed of a dense shrub layer and a scattered herb layer of
perennial forbs. The oak canopy is composed of a variety of stand
types that range from very dense, small-diametered to large, broadly
branched individuals. Canopy trees are found on the banks above
the incised stream channels with their crowns extending over the
channel but not rooted in the channel. Philadelphus lewisii and Hol-
odiscus discolor dominate the tallest shrub layer found in these bald
hills oak woodlands, and often reach a height of 7 m. Sanicula
crassicaulis and D. trolliifolium are important associates in the herb
layer. Characteristic shrubs include Rosa pisocarpa, Amelanchier
pallida, and Osmaronia cerasiformis. Cystopteris fragilis, Polypo-
dium glycyrrhiza, and Tellima grande are characteristic members of
the herb layer.

204 MADRONO [Vol. 34

Ribes/Phacelia (Ri/Ph): This rock outcrop type is composed of a
moderately dense shrub layer and a scattered herb layer of perennial
and annual forbs and grasses. The oak canopy is generally present
although often not well developed. When trees are present they
usually grow adjacent to the outcrops and frequently shade them.
Ribes roezlii and Rhus diversiloba dominate the shrub layer. Im-
portant associates in the herb layer include G. aparine, Caucalis
microcarpa, and Bromus rigidus, Phacelia heterophylla, Silene cal-
ifornica, Cheilanthes gracillima, and Avena barbata are the common
characteristic species. Notably uncommon on the rock outcrops are
the grasses such as H. lanatus, Agrostis hallii, M. subulata, and
Trisetum cernuuwm, which are abundant in the understory of the
open woodlands.

DISCUSSION

The indirect ordination (Fig. 3) resulted in clustering of plots in
two dimensional space corresponding to the seven plant community
types. The left end of DCA axis-one represents dense stands in mesic
locations with less historic grazing disturbance and a lower repre-
sentation by introduced species. The Ar/Sh type occurs at the far
right. These open glades without summer shade sustained the great-
est grazing impact. The lower half of DCA axis-2 represents well
developed continuous soils. Rock outcrops and rocky stream chan-
nels occur at the top of the ordination. The Qu/De occurs on rocky
soils and appears intermediate between the types characteristic of
outcrops and well developed soils.

Oak/grassland mosaic. The distinctive oak/grassland mosaic char-
acteristic of the bald hills oak woodlands is best developed in the
southeast corner of RNP. This pattern occurs primarily on higher
ridges, but is continuous downslope on earthflows with southern
exposures. Mid-slope oak woodlands are extensive and associated
with occasional glades and a large central open prairie. At low ele-
vations and closer to the coast, the prairie becomes the primary
feature, with Q. garryana stands restricted to forest margins and
narrow projections into the grasslands. Oak woodlands extend
downslope to Redwood Creek at elevations of less than 100 m in
several locations. In mesic low elevation and coastal areas within
the fog zone, P. menziesii forest has colonized former oak woodlands
during the past 130 years. Low elevation stream channels and rock
outcrops are converted completely to this conifer forest, and only
the open woodlands are left intact. The remaining outcrops and
streamside vegetation occur only at mid- to upper elevations in the
study area. The remaining open woodlands are the Qu/Da type that
is correlated with the mesic nature of the concave lower slopes within
the fog belt.

1987] SUGIHARA ET AL.: BALD HILLS WOODLANDS 205

Open woodlands. The Qu/Da, Qu/Sy, and Qu/Cy types comprise
most of the area within the well developed open woodlands. Dis-
tribution of these three types is related to topography, slope position,
grazing, and fire history. Qu/Da and Qu/Sy are found exclusively
under closed-canopy oak stands on lower slopes and mid- to upper
north-facing slopes. Aspect, sheltered topography, and frequent sum-
mer fog make these relatively mesic sites. The Qu/Da type is pre-
dominant on the lower slopes under stands of 70-100 yr-old oaks.
Qu/Sy occurs under very dense stands of small-diameter oaks on
upper north-facing slopes. These stands originated following a fire
in 1948 (Sugihara et al. 1983). The two mesic woodland types are
closely related floristically. Qu/Sy is probably a fire sere of Qu/Da.
Qu/Sy is clearly distinguished from Qu/Da by the dense S. rivularis-
dominated shrub layer characteristic of the Qu/Sy type. The third
major woodland type, Qu/Cy, is found in oak stands composed of
all size classes but dominated by widely-spaced, large-diameter trees.
This association is found on upper, convex, south-facing slopes and
along the ridgeline where moisture conditions are more xeric and
grazing was the most intense.

Specialized habitats. The remaining four plant community types
are confined to specialized habitats within the study area. Ar/Sh
occurs in glades among the oak stands and includes many species
characteristic of the continuous open grassland adjacent to the study
area. These include weedy introduced grasses and forbs such as A.
elatius, C. echinatus, A. caryophyllea, Trifolium dubium, Trifolium
subterraneum, Linum bienne, H. lanatus, R. acetosella, Plantago
lanceolata, Hypochoeris radicata, and Pteridium aquilinum (Hektner
et al. 1983). Glades also are related floristically to the xeric Qu/Cy
woodland, but not limited to xeric topographic positions within the
study area. Qu/De is found on relatively moist concave slopes within
xeric Qu/Cy woodlands. Dominance of native forbs, especially D.
trolliifolium, is especially evident in the early summer when the
adjacent Qu/Cy type is dominated by immature grasses. Lack of
canopy cover results in dominance by characteristic xeric woodland
species in mesic physiognomic positions. Two minor community
types are found in areas that have very little soil. The Ph/Cy type
occupies incised, boulder-strewn stream channels on upper slopes.
The numerous dry, exposed, rock outcrops are occupied by the R1/
Ph type. Both Ph/Cy and Ri/Ph are dominated by tall shrubs that
thrive on the bare soil and rock surfaces not covered by a thick
herbaceous mat.

Introduced species. Success of introduced species in the seven
community types appears to be influenced by past grazing and fire.
The most heavily grazed communities are grass-dominated and
composed of the highest percentage of introduced species. Only 50%

206 MADRONO [Vol. 34

of the high presence species found in Ar/Sh and 53% in Qu/Cy are
native. Qu/De is dominated by native forb species with 68% of the
high presence species native. Grazing impact on Qu/De was reduced
by the lack of early summer grazing, which was restricted due to the
toxicity to cattle of the early season dominant Delphinium trolli-
ifolium. Mesic woodlands supporting the Qu/Da type had 74% na-
tive species. Stream channels and rock outcrops were more protected
from grazing because of inaccessibility. This is reflected in the 75%
native species for Ri/Ph and 82% for Ph/Cy. The highest represen-
tation of native species in any vegetation type was the 85% found
in the Qu/Sy community that was recently influenced by fire.

Relationship to other woodlands. Ecologically and floristically, these
QO. garryana woodlands are more similar to those of the Willamette
Valley of interior Oregon than to any other California woodland
type. The bald hills oak woodlands, however, are distinct from the
interior Oregon woodlands in structure, composition, and their coastal
habitat (Thilenius 1968). Shrubs dominate all Willamette Valley
plant communities, but in bald hills woodlands only the Qu/Sy type,
stream channels, and rock outcrops support well developed shrub
layers. In the Willamette Valley and the bald hills, stand structure
was determined largely in the past by burning. Savannahs with grassy
openings between the individual trees that are characteristic of the
Willamette Valley were not typical of woodlands in this area. His-
torical accounts indicate that Q. garryana was well-spaced in the
pre-settlement stands of the bald hills, but the canopy was closed
and alternated with the grasslands.

Fire and succession. Reduction of fire frequency during post-set-
tlement times has altered succession in both the Willamette Valley
and the bald hills. Succession is from oak savannah to oak forest,
and then to Pseudotsuga menziesii forest in Oregon. In the bald hills,
succession is from oak forest to mixed evergreen forest in the xeric
interior areas, and to Sequoia sempervirens/P. menziesii at low el-
evation and coastal mesic areas (Sawyer et al. 1977). With the ces-
sation of burning by Native Americans and introduction of wildfire
suppression, succession to P. menziesii has progressed without nat-
ural control. Subsequent succession to S. sempervirens 1s seen in
older mesic stands of P. menziesii. The absence of redwood forest
on potential redwood sites supports archaeological evidence of con-
stantly high fire frequency over several thousand years prior to Eu-
ropean settlement.

Although allied more closely to the Willamette Valley woodlands
than any California oak type, the bald hills oak woodlands are a
unique feature of California’s redwood region. The National Park
Service has allowed the vegetation of bald hills to develop ungrazed
for the first time in over a century. This study provides a description

1987] SUGIHARA ET AL.: BALD HILLS WOODLANDS PAU

of existing vegetation patterns in the northern bald hills. This base-
line information is essential for the monitoring of vegetational changes
occurring in response to management of oak woodlands in Redwood
National Park.

ACKNOWLEDGMENTS

We express appreciation to all of the people who reviewed the manuscript, especially
James Agee and John Sawyer. We thank the Biology Department of Humboldt State
University for the use of their herbarium. Special thanks to Mary Hektner, Don
Reeser, James Popenoe, Roy Martin, and the staff at RNP.

LITERATURE CITED

BENSON, J. R. 1983. Archaeological test excavations at four sites in Redwood Na-
tional Park, Humboldt County, California. Redwood National Park, Arcata, CA.

BICKEL, P. M. 1979. A study of cultural resources in Redwood National Park.
U.S.D.I. National Park Service, Denver Service Center, Denver, CO.

BRAUN-BLANQUET, J. 1932. Plant sociology, the study of plant communities. (Transl.
by G. D. Fuller and H. S. Conard.) McGraw-Hill, New York.

CLARK, H. W. 1937. Association types in the North Coast Ranges of California.
Ecology 18:214—230.

COGHLAN, M. 1984. Aclimatologically-based analysis of the storm and flood history
of Redwood Creek. Tech. Rep. No. 10, Redwood National Park, Arcata, CA.

FRANKLIN, J. F. and C. T. DyrneEss. 1973. Natural vegetation of Oregon and Wash-
ington. U.S.D.A. For. Serv. Gen. Tech. Rep. PNW-8.

GRIFFIN, J.R. 1977. Oak woodland. Jn M. G. Barbour and J. Major, eds., Terrestrial
vegetation of California, p. 384-415. Wiley Interscience, New York.

and W. B. CRITCHFIELD. 1972. The distribution of forest trees in California.
U.S.D.A. For. Serv. Res. Paper PSW-82.

Hayes, J. F. 1985. An analysis of Redwood National Park artifacts. Redwood
National Park, Crescent City, CA.

HEKTNER, M. M., R. W. Martin, and D. R. DAVENPORT. 1983. The Bald Hills
prairies of Redwood National Park. Jn C. Van Riper III, L. D. Whittig, and M.
L. Murphy, eds., Proceedings of the first biennial conference of research in Cal-
ifornia’s National Parks, p. 70-78. Cooperative Park Studies Unit, Univ. Cal-
ifornia, Davis.

Hitt, M.O. 1979a. TWINSPAN: a FORTRAN program for arranging multivariate
data in an ordered two-way table by classification of the individuals and attri-
butes. Ecology and Systematics, Cornell Univ., Ithaca, NY.

. 1979b. DECORANA: a FORTRAN program for detrended correspondence
analysis and reciprocal averaging. Ecology and Systematics, Cornell Univ., Ith-
aca, NY.

HUMBOLDT STATE UNIVERSITY. 1974. Simpson’s Bald Hills ranch: range economics
and planning. Range Management 170. Arcata, CA.

Kina, A. G. and P. McW. BIcKEL. 1980. Resource evaluation at nine archeological
sites, Redwood Creek basin, Redwood National Park, California. Redwood Na-
tional Park, Arcata, CA.

Lewis, H. T. 1973. Patterns of Indian burning in California: ecology and ethno-
history. Ballena Press, Ramona, CA.

MUELLER-DomBsols, D. and H. ELLENBERG. 1974. Aims and methods of vegetation
ecology. John Wiley and Sons, Inc., New York.

Munz, P. A. 1973. A California flora with supplement. Univ. California Press,
Berkeley.

SAWYER, J. O., D. A. THORNBURGH, and J. R. GRIFFIN. 1977. Mixed evergreen

208 MADRONO [Vol. 34

forest. Jn M. G. Barbour and J. Major, eds., Terrestrial vegetation of California,
p. 359-381. Wiley Interscience, New York.

STORIE, R. E.and A. E. WIESLANDER. 1952. Dominant soils of the redwood—Douglas-
fir region of California. Soil Sci. Soc. Amer. Proc. 16:163-167.

SUGIHARA, N. G., M. M. HEKTNER, L. J. REED, and J. M. LENIHAN. 1983. Oregon
white oak woodlands of Redwood National Park: description and management
considerations. /n C. Van Riper III, L. D. Whittig, and M. L. Murphy, eds.,
Proceedings of the first biennial conference of research in California’s National
Parks, p. 177-182. Cooperative Park Studies Unit, Univ. California, Davis.

THILENIUS, J. F. 1968. The Quercus garryana forests of the Willamette Valley,
Oregon. Ecology 49:1124—1133.

THOMPSON, L. 1916. To the American Indian. Cummins Print Shop, Eureka, CA.

WHITTAKER, R. H. 1960. Vegetation of the Siskiyou mountains, Oregon and Cal-
ifornia. Ecol. Monog. 30:279-338.

WIESLANDER, A. F. and H. A. JENSEN. 1948. Forest areas, timber volumes, and
vegetation types in California. Forest Survey Release No. 1, California Forest
and Range Experiment Station.

(Received 2 Jul 1985; revision accepted 23 Jan 1987.)

NEW MADRONO POLICY

Madrono now accepts manuscripts written in Spanish. The first paper to be
published under this new policy will appear in 35(1), and a limited number
will be published in Spanish in each volume. All contributors of these manu-
scripts should follow general Madrono conventions, and also should include
an English language abstract.

Members of the California Botanical Society have a long-term and apparently
increasing interest in the botany of Mexico and Central and South America.
Because of this interest, the editors believe our new policy will provide broader
communication among scientists, will open the journal to a wider readership,
and perhaps will increase membership in the CBS. We also hope it will extend
a gesture of goodwill to our Hispanic colleagues and neighbors. We look forward
to a successful bilingual journal and trust the membership will support this
new policy.

UNA NUEVA POLITICA DE MADRONO

A partir de esta fecha Madrono aceptara manuscritos redactados en Espanol.
El primer articulo que se publicara dentro de esta nueva politica aparecera en
el volamen 35(1); en cada volimen se publicaran un numero limitado de
articulos escritos en Espanol. Los autores deberan utilizar las convenciones
editoriales de Madrono e incluir un resumen del trabajo en Inglés.

Los miembros de la Sociedad Botanica de California tienen un interés a
largo plazo en la botanica de México, Centro y Sudameérica; dicho interés
aparentemente se esta incrementando. Por esta razon, los editores creemos
que esta nueva politica ampliara la comunicacion entre los cientificos, la revista
estara al alcance de un mayor numero de lectores y probablemente se extender
un gesto de buena voluntad para con nuestros colegas y vecinos hispano-
parlantes. Confiamos en el éxito de esta revista bilingiie y en el apoyo hacia
esta nueva politica por parte de los socios.

A FLORA OF VINA PLAINS PRESERVE,
TEHAMA COUNTY, CALIFORNIA

PAULEEN BROYLES
Department of Biological Sciences,
California State University, Chico 95929

ABSTRACT

The 626 hectare Vina Plains Preserve is located in southernmost Tehama Co.,
California, and is a remnant of various native grassland habitats. Geologically, this
region is a weathered fanglomerate formed from alluvial deposits of the Tuscan
formation and later deposits of silt. A botanical survey of the Preserve between
January 1982 and May 1987 resulted in the identification of 53 vascular plant families
that included 287 taxa. Native species comprised 67% of the taxa, and annuals
comprised 77%. Eight rare plant species were found, their distributions noted, and
numbers estimated. All the plants occur in one or more of the six habitats, which
include vernal pools, hogwallows, seeps, vernal marshes, uplands, and outcrops.
Families with the most species include Poaceae (50), Asteraceae (36), Fabaceae (17),
Boraginaceae (13), and Amaryllidaceae (10).

The original species composition of the pristine Californian grass-
lands is largely unknown (Bartolome and Gemmill 1981). The first
direct evidence for the replacement of native perennial bunchgrass
by introduced annual species on Californian grassland has been pro-
duced by a recent study of opal phytoliths at Jepson Prairie Reserve
in Solano Co. (Bartolome et al. 1986). There is little detailed infor-
mation on the composition of other similar Californian grasslands
(Jokerst 1983). Published studies are available for the following areas
of the northern Sacramento Valley: Richvale Vernal Pool Site, Butte
Co. (Schlising and Sanders 1983), Jepson Prairie Nature Conser-
vancy Preserve, Solano Co. (Holland 1981), Maidu Park, Placer Co.
(Holland 1982), and Table Mountain, Butte Co. (Jokerst 1983).

The vernal pools of Californian grasslands contain a high pro-
portion of endemic plants (Holland 1976). With increasing urban-
ization and the expansion of agribusiness, these habitats are dis-
appearing except where they are protected by specific organizations.
One such area of grassland and vernal pools is Vina Plains Preserve
in Tehama Co., which is owned and managed by The Nature Con-
servancy. The purposes of this study were to conduct an inventory
of all vascular plant species and their habitats on the Preserve, and
to provide detailed information on the occurrence of rare plants.

STUDY AREA

Location. The Vina Plains Preserve is located in southernmost
Tehama Co., in the northern portion of the great Central Valley of

MADRONO, Vol. 34, No. 3, pp. 209-227, 1987

[Vol. 34

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MADRONO

210

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1987] BROYLES: VINA PLAINS PRESERVE pA |

Fic. 2. Aerial view of the Vina Plains Preserve from the south, showing several
large vernal pools and a barn located on the access road.

California (Fig. 1). The area is bordered by Tuscan volcanic mud-
flows of the Cascade Range to the east and the Sacramento River
to the west. The elevation is about 66 m, and topographic relief
varies by about 4.5 m from north to south. The Wurlitzer addition
in 1984 (176 ha) is not included in the present study.

The study area is composed of flat or mounded, rolling grassland,
interspersed by several kinds of wet areas (Fig. 2). These wet areas
consist of: 1) four drainages that were deepened for irrigation in the
past, and that traverse the parcel in a roughly north/south direction;
2)a number of vernal pools of varying sizes; and 3) many hogwallows
(Fig. 3), smaller depressions, and natural drainages that hold water
for short periods of time.

Geology and soils. The following theory of the formation of parent
materials for the soils is based on information from J. W. Guyton
(pers. comm.), a geologist at California State University, Chico. The
oldest rock type in the area is an alluvial fan deposit. It was derived
from the Tuscan formation composed of old volcanic mudflows that
make up a considerable portion of the foothills from Red Bluff

212 MADRONO [Vol. 34

Fic. 3. A large hogwallow that displays “‘rings”’ of flowering species. Mt. Lassen
is visible in background.

southward to the Vina area (Harwood et al. 1981). This deposit
consists of coarse sands and gravels that were transported from the
foothills on the northeast and east, deposited in alluvial fans, and
cemented. This “fanglomerate”’ is 1 to 2 million yr old, mainly
andesitic and basaltic in composition, and well cemented into rock
(conglomerate) by materials in the ground water. At Vina Plains
Preserve, the strata are exposed in such places as the waterways and
edges of the pools. About 100,000 yr ago, weathering weakened the
cement, converting the conglomerate back to sand, silt, and gravel
to a variable depth of 2-5 m. Subsequently (10,000—20,000 yr ago)
the deposition of silt occurred either by wind or by flooding of the
Sacramento River during the period of glacial climate. The fine-
grained sediment is evident in isolated mounds. Finally, strong winds
associated with a dry climate during the period that ended about
4000 yr ago (known as the altithermal) could have removed much
of the silt layer and excavated pools in the weathered fanglomerate.
The pools hold water because of the basic, slightly permeable fan-
glomerate floor, and because of the clay that washed in from the
higher surroundings. Milder erosion by gentle runoff could have
produced the many gullies that occur naturally.

The existing soils are largely of andesitic and basaltic composition,
and are described as the “‘Tuscan series” (U.S.D.A. 1967). The sur-
face is dark brown and cobbly; the subsoil is more reddish, clayey,

1987] BROYLES: VINA PLAINS PRESERVE PAE

and gravelly, and is often exposed at the margins of pools and other
eroded areas. Other series include the Keefers, similar to Tuscan,
and Anita and Berrendos, that are more clayey. The deeper (Anita
and Berrendos) soils are found commonly where pools have formed
or in the drainage channels. All soils and pools have an underlying
hardpan.

HABITATS AND VEGETATION

General trends during one field season indicated that the plants
on the Preserve tend to group themselves into six general habitats,
with some overlap or transition between habitats. For purposes of
this study, these were termed upland, pool, hogwallow, seep, vernal
marsh, and outcrop.

Upland. The majority of the terrain, consisting of rather well
drained areas, is considered upland. The soil is mainly Tuscan loam:
it is slightly acidic, with depth from a few centimeters to about one
meter (U.S.D.A. 1967), and is slightly cobbly. Water runs off quickly
and collects in the internal drainages. Species found in uplands are
mainly annual grasses and forbs, such as Lasthenia californica, Layia
fremontii, Orthocarpus erianthus, Lepidium nitidum, Hemizonia
fitchii, several species of Erodium, Navarretia, Vulpia, and Bromus,
and such perennials as Dodecatheon clevelandii subsp. patulum and
Brodiaea californica.

Seeps. Areas that have at least some moisture supplied by slow-
moving water during most of the year are considered seeps. These
include margins of irrigation ditches and adjoining low areas. The
soil is often deep and contains much silt. Such habitats are found
along (east to west) Singer Creek Ditch, Sheep Camp Ditch, and two
other ditches traversing the west pasture (Fig. 1). Typical species
are: Callitriche heterophylla subsp. bolanderi, Centaurium floribun-
dum, Elatine heterandra, Limosella acaulis, Lythrum hyssopifolia,
Mimulus guttatus, Crassula saginoides, and Ranunculus muricatus.
In some places, the plants occur in zones. For example, at one site
Eleocharis macrostachya occurs in standing water of a ditch, Am-
mannia coccinea and Diplachne fascicularis on the muddy shore,
Echinochloa crusgalli on adjacent damp grounds, and Eragrostis
cilianensis with the last two.

Hogwallows. The hundreds of potholes, depressions, and internal
drainages that have ephemeral standing water are considered hog-
wallows. The soil is usually thinner than in seeps. Hogwallows may
have rocky bottoms and varying amounts of sand and silt. When
silt is present, species occur that also are found in pools. Typical
species of hogwallows are: Lasthenia fremontii, Limnanthes doug-
lasii var. rosea, Plagiobothrys stipitatus var. micranthus, and Down-

214 MADRONO [Vol. 34

ingia ornatissima or D. bicolor. Under optimum conditions, hog-
wallows display concentric colored “‘rings’’ of flowering species.

Pools. A “pool” is deeper and larger than a hogwallow and is
presumed to have been formed by the blow-out process. The bottom
is composed of clay or silt underlain by impermeable fanglomerate.
Water accumulated during winter rains remains into late spring or
early summer. The maximum depth of the different pools varies
from 0.3—1.0 m; however, depth also fluctuates with seasonal rainfall
and with drydown. Ten large pools were found to contain Orcuttia
pilosa and O. tenuis.

The largest of these (located in the northern portion of the west
pasture) measures about 345 x 142 m. The smallest (located in the
northeast portion of the south pasture) measures 80 x 65 m. Most
are in areas of Anita clay loam, but three are on Tuscan loam. All
have some degree of silt or clay accumulated by erosion from sur-
rounding higher ground. The deepest pool retained water (in 1982)
into June, but most had no standing water by early to mid-May. In
some, the margins are sandy, whereas in others there are many
cobbles with heavy varnish that indicates great age. Typical species
are: Asclepias fascicularis, Eryngium vaseyi var. vallicola, Marsilea
vestita, Navarretia leucocephala, Psilocarphus brevissimus, Down-
ingia bella, D. bicornuta, Orcuttia pilosa, O. tenuis, Tuctoria gree-
nei, and Chamaesyce hooveri. Although there is a greater species
richness on the upland, the species listed for pools (except C. hooveri
and Orcuttia tenuis) usually occur in all the large pools.

Vernal marshes. The vernal marsh differs from the hogwallow
habitat in its greater area and from the pool habitat in that it lacks
deep, standing water. Water remains in the marsh habitat longer
than in the hogwallows, but not as long as in the pools. The soil is
thin Tuscan loam and often contains additional clay deposited from
erosion. Species found here include primarily native annual forbs
found in the pools and hogwallows, and some native annual grasses,
which include those that are generally found in upland habitats and
hogwallows. Boisduvalia cleistogama and B. glabella, native forbs
found in the pools, are present in this habitat. Other typical species
include: A/opecurus saccatus, Limnanthes douglasii var. rosea, and
Plagiobothrys stipitatus var. micranthus.

Outcrops. This is a habitat of thin, poor, rocky or sometimes
gravelly soil. These outcrops usually occur on the upiand, but also
occasionally are found within the dried pools. Usually, the outcrop
is the driest habitat, and the plants are small, native or introduced
annuals. Typical outcrop species are: Crassula connata, Parvisedum
pumilum, Koeleria phleoides, and occasionally Plantago erecta and
P. bigelovii.

1987] BROYLES: VINA PLAINS PRESERVE ZS

FLORA

Summary. Fifty-three vascular plant families were found that con-
sist of 287 taxa. Two other species (Populus fremontii and Salix sp.)
have been extirpated by the razing of a man-made reservoir. Of the
total taxa, 192 (67%) are native and 95 (33%) are introduced; 221
(77%) are annuals and 66 (23%) are perennials.

Rare plants. Eight species at Vina Plains Preserve are mentioned
in the California Native Plant Society (CNPS) Jnventory of Rare and
Endangered Plants of California (Smith and York 1984). A ninth,
Paronychia ahartii, is too recently described for listing, but is being
considered for inclusion.

Astragalus pauperculus has been found scattered at a few sites on
thin Tuscan soil where gaps in the vegetation occur. It is incon-
spicuous, and may occur more often than indicated. Smith and York
(1984) list it as rare but not endangered, and endemic to California.
Its Federal status is ““‘widespread, not threatened.”

Cuscuta howelliana was found in three of the pools and a hog-
wallow. Due to the twining, meandering nature of the plant, counts
must be based on the number of host plants; these vary from a few
(3-15) to many, covering about one fourth of one pool. Although
not endangered (CNPS, Federal), this species is rare and endemic
to California, and requires monitoring.

Chamaesyce hooveri was found in six of the pools and a hogwallow.
Population size varied from a single plant in the hogwallow to about
3000 in the largest pool. Mature plants form mats up to one meter
in diameter by late summer. It is rare and endemic to California
and is endangered throughout its range (CNPS) because of loss of
vernal-pool habitat. Insufficient data, however, are available for Fed-
eral listing.

Fritillaria pluriflora has been found at seven sites; all but one of
these occurred in deep clay soil. Population size was from 15-30,
and usually approached the latter. Although rare and endemic to
California, it is considered endangered (CNPS) in only a portion of
its range and is not on the Federal list.

Lepidium latipes has been found in a few hogwallows. A census
in 1985 at one large hogwallow revealed a population of 584 plants.
This species is rare, but not endangered in California, and is not on
the Federal list.

Orcuttia pilosa was found in six pools, but occurred in large num-
bers in only the four deepest ones. Numbers were estimated in the
thousands. It was found in association with Tuctoria greenei in only
one pool. It is rare and endangered in California throughout its range
(CNPS) due to reduction of habitat. Data are on file to support a
Federal listing.

216 MADRONO [Vol. 34

Orcuttia tenuis occurs in one small pool at the northeast corner
of the Preserve. Numbers were estimated at 5000—10,000 plants. A
second germination took place following mid-summer rains in 1982,
and this population was estimated at about 1000 plants. The status
of this species is similar to O. pilosa.

Paronychia ahartii Ertter was found at one location in the eastern
portion of the Preserve in April 1984. It was known previously only
from a few, widely separated areas in northern California, from
collections made by J. T. Howell, and from the Lowell Ahart Ranch
in Butte Co. (J. Jokerst pers. comm.). It is being considered for CNPS
listing; more data are needed to support a Federal listing.

Tuctoria greenei is more widespread than Orcuttia pilosa or O.
tenuis. Most populations of it are smaller and usually consist of a
few hundred plants; however, about 30,000 plants have occurred in
one exceptional pool. Tufting makes accurate counts difficult to
obtain. This is a rare endemic of California and is endangered
throughout its range due to reduction of habitat (CNPS). Data are
on file to support a Federal listing.

Annotated catalogue. The following list includes all species col-
lected between late January—October 1982 and March—May 1983.
Seven additions have been made from 1984-1987. Fifty-two visits
were made in the original study, most frequently during the peak
flowering season. Voucher specimens are housed at CHSC. Nomen-
clature follows Munz (1959, 1968). Current regional authorities are
followed whenever possible. Synonyms are included if names differ
from Munz. Each entry includes information on habitat and flow-
ering phenology at the Preserve, and whether the species is annual
(A), biennial (B), perennial (P). If the plant is not native to the area,
it is marked with an asterisk. Occurrence of most species is noted
as abundant, common, occasional, and uncommon. Rare plants list-
ed in Smith and York (1984) are noted.

VASCULAR PLANTS OF VINA PLAINS PRESERVE

LEPIDOPHYTA
Isoetaceae

Isoetes howellii Engelm. P; standing water of seeps, hogwallows, and pool margins.
Isoetes nuttallii A. A. Eat. P; habitat as for J. howellii, but appears earlier in spring.

PTEROPHYTA
Marsileaceae

Marsilea vestita Hook. & Grev. P; abundant in vernal pools, occasional in hogwallows.
Pilularia americana A. Br. P; standing water of irrigation ditches, occasional at
margins of pools.

1987] BROYLES: VINA PLAINS PRESERVE PANE

ANTHOPHYTA — DICOTYLEDONEAE
Amaranthaceae

*4maranthus albus L. A, common; dried pools and low places; Jul-Sep.
Amaranthus blitoides Wats. A; disturbed areas; Jun—Aug.

Apiaceae

*Anthriscus scandicina (Weber) Mansf. A, occasional; upland; Feb.

Eryngium vaseyi Coult. & Rose var. vallicola (Jeps.) Munz. B or P, locally abundant;
vernally wet areas; Jun—Jul.

Lomatium humile (Coult. & Rose) Hoov. ex Math. & Const. P, common; upland,
on gentle slope or depression; Tuscan loam; Mar-Apr.

Sanicula bipinnatifida Dougl. ex Hook. B or P, common; upland slope or shallow
depression; Tuscan loam or deeper soils; Mar—Apr.

Asclepiadaceae

Asclepias eriocarpa Benth. P, occasional; upland; Jun.
Asclepias fascicularis Dene. in A. DC. P; abundant in dried pools, occasional long
internal drainages and ditches; Anita clay or Tuscan or Keefers loam; Jun—Sep.

Asteraceae

Achyrachaena mollis Schauer. A; hogwallows and upland where heavier soils hold
moisture; Apr.

Blennosperma nanum (Hook.) Blake. A, common; hogwallows and vernal marshes,
shallow to deep; thin or cobbled soils or clay; Feb—Mar.

*Centaurea solstitialis L. A, widespread, especially on disturbed soil; May—Aug.

*Conyza canadensis (L.) Cronq. A; scattered locations of upland; Tuscan and Keefers
loam; Aug-—Sep.

Evax acaulis (Kell.) Greene. A, occasional; upland or hogwallow; May—Apr.

Evax caulescens (Benth.) Gray. A; vernally moist areas, especially pool edges and
hogwallows; clay soil; Apr-Jun.

*Filago gallica L. A, occasional; disturbed Tuscan loam; Apr—May.

*Gnaphalium luteo-album L. A, common, especially on disturbed sites; Jul.

Hemizonia fitchii Gray. A, common; entire Preserve except wet areas; May—Sep.

Hemizonia luzulaefolia DC. subsp. rudis (Benth.) Keck. A; hogwallow areas and, to
a lesser extent, borders of seeps; more gregarious than H. fitchii but less widespread;
Jul-Sep.

*Hypochoeris glabra L. A, common; upland, vernal marsh, and hogwallow; May—
Jun.

*Lactuca saligna L. A; scattered locations of upland, especially in disturbed areas;
Jul-Aug.

*Lactuca serriola L. A; more common than the previous species but similar habitat;
Aug.

Lagophylla glandulosa Gray subsp. glandulosa. A, abundant; upland on thin or clayey
Tuscan loam; Jun—Sep.

Lagophylla glandulosa Gray subsp. serrata (Greene) Keck. A, common; upland open
slopes; thin Tuscan loam; May-Jun.

Lasthenia californica DC. ex Lindl. [L. chrysostoma (Fisch. & Mey.) Greene] A,
common; widespread in many habitats except wettest or most drained; Mar—May.

Lasthenia fremontii (Torr. ex Gray) Greene. A, common; widespread in hogwallows
and margins of pools, less so on drained upland; Tuscan loam or sometimes deeper
soils; Mar—May.

Lasthenia glaberrima DC. A; dried pools; May-Jun.

Lasthenia glabrata Lindl. subsp. coulteri (Gray). A; upland; Apr.

218 MADRONO [Vol. 34

Lasthenia platycarpha (Gray) Greene. A, locally common; hogwallows or wet upland;
Feb—Apr.

Layia fremontii (T. & G.) Gray. A; ubiquitous on upland and bordering hogwallows;
Feb—May.

*Leontodon leysseri (Wallr.) G. Beck. A, common; Tuscan or deeper soil of upland,
or near seeps; Jun.

*Leontodon taraxacoides (Vill.) Meart. A; Tuscan loam or heavier soils of upland,
or near seeps; May.

Lessingia nana Gray in Benth. A, abundant; upland; Jul-Aug.

Matricaria matricarioides (Less.) Porter. A, common on disturbed sites; Mar-—Jul.

Micropus californica F. & M. A, common; dried hogwallows or open upland; Tuscan
loam; Apr—May.

Microseris acuminata Greene. A; upland; Apr.

Microseris douglasii (DC.) Sch.-Bip. subsp. douglasii. A; upland or hogwallows; Tus-
can loam; Apr.

Microseris douglasii (DC.) Sch.-Bip. subsp. tenella (Gray) Chamb. A, occasional;
upland; Tuscan loam or deeper soils; May.

Psilocarphus brevissimus Nutt. A, common; dried vernal pools, hogwallows, and
vernal marshes; Apr—Jun.

Psilocarphus oregonus Nutt. A, less common than the preceding species; habitat and
phenology similar.

*Senecio vulgaris L. A; disturbed areas; Jan—Sep.

*Silybum marianum (L.) Gaertn. A, occasional; near irrigation ditches; May.

*Sonchus asper L. A, occasional; Jul.

*Sonchus oleraceus L. Similar to S. asper.

*Xanthium strumarium L. var. canadense (Mill.) T. & G. A; dried vernal pools,
abundant in some; Jul-Aug.

Boraginaceae

Amsinckia intermedia F. & M. A; found on a grassy slope near a fence; Apr.

Amsinckia menziesii (Lehm.) Nels. & Macbr. A; found with A. intermedia; Apr.

*Heleotropium europaeum L. A; scattered locations on upland; Tuscan or clay loam;
Aug-Sep.

Plagiobothrys austinae (Greene) Jtn. A; vernally wet, slight depressions; Tuscan or
deeper soils; Mar—Apr.

Plagiobothrys canescens Benth. A; occasional populations on upland; May-Jun.

Plagiobothrys fulvus (H. & A.) Jtn. var. campestris (Greene) Jtn. A; occasional pop-
ulations on upland; Mar-Apr.

Plagiobothrys glyptocarpus (Piper) Jtn. A; seeps; Tuscan or Keefers loam; Apr—May.

Plagiobothrys greenei (Gray) Jtn. A; hogwallows and other slight, vernally wet depres-
sions; Tuscan or deeper soils; Mar—Apr.

Plagiobothrys humistratus (Greene) Jtn. A; phenology and habitat as for P. scriptus.
According to Schlising (1984), may be conspecific.

Plagiobothrys leptocladus (Greene) Jtn. A; seeps and shallow hogwallows; Tuscan or
heavier soils; Apr-May.

Plagiobothrys scriptus (Greene) Jtn. A, uncommon; upland on thin Tuscan or deeper
soils, sometimes hogwallows; prostrate and inconspicuous; Feb—Mar.

Plagiobothrys stipitatus (Greene) Jtn. var. micranthus (Piper) Jtn. A; pools (mostly)
and hogwallows; Mar—Jun.

Plagiobothrys stipitatus (Greene) Jtn. var. stipitatus. A; hogwallows; Mar-Apr.

Brassicaceae

Athysanus pusillus (Hook.) Nutt. A; occasional populations in many habitats except
wettest; Feb—Mar.
*Brassica campestris L. A; occasional on disturbed soil; Apr-May.

1987] BROYLES: VINA PLAINS PRESERVE 219

*Capsella bursa-pastoris (L.) Medic. A, common; disturbed areas; Feb—Apr.

Cardamine oligosperma Nutt. A, occasional; Tuscan loam or deeper soils; Mar—Apr.

Draba verna L. A, scattered locations on vernally moist upland; thin soil; Mar—Apr.

Lepidium lasiocarpum Nutt. A, occasional; clayey soil; Apr.

Lepidium latipes Hook. A; found at two sites: several small, scattered populations in
hogwallows near barn in south pasture; single, larger population in deep soil near
irrigation ditch in south pasture; Mar.

Lepidium nitidum Nutt. A, common and widespread; Feb—Mar.

*Raphanus raphanistrum L. B, common; disturbed areas; Jan—Apr.

*Raphanus sativus L. Similar to R. raphanistrum.

Rorippa palustris (L.) Bess. subsp. glabra (O. E. Schultz) Stuckey. A or B, occasional;
disturbed areas; Apr.

* Sisymbrium officinale (L.) Scop. A, occasional; disturbed soil near an irrigation ditch;
Apr.

Callitrichaceae

Callitriche hermaphroditica L. var. hermaphroditica. A, occasional; irrigation ditches;
Apr.

Callitriche heterophylla (Pursh.) subsp. bolanderi (Hegelm.) Calder & Taylor. P, com-
mon and widespread in pools, hogwallows, and ditches, less common in vernal
marshes; Mar.

Callitriche longipedunculata Morong. A, occasional; hogwallows; Apr.

Callitriche marginata Torr. A, occasional; pools and hogwallows; Apr.

Campanulaceae

Downingia bella Hoov. A, abundant; pools, hogwallows, and seeps; Apr-May.

Downingia bicornuta Gray. A; hogwallows, seeps, and pools; Apr-May.

Downingia cuspidata (Greene) Greene. A; pools; May.

Downingia ornatissima Greene. A; seeps, hogwallows, and shallow pools or vernal
marshes; Apr-May. This species occurs with D. bella, but with D. bicornuta at
only one location, a wet depression.

Githopsis specularioides Nutt. A, uncommon; upland; Apr.

Caryophyllaceae

*Cerastium glomeratum Thuill. A, widespread on upland, and common in disturbed
areas; Mar—Apr.

Minuartia californica (Gray) Mattf. [Arenaria california (Gray) Brew.] A, common;
thin soils and cobbled areas of Tuscan loam, and especially in dried hogwallows;
Feb—May.

Paronychia ahartii Ertter. A; small population found on thin soil of well-drained
upland at northeast corner of Preserve; Mar—Apr.

*Petrorhagia velutina (Gussone) Ball & Heywood. [Tunica prolifera L., Kohlrauschia
velutina (Guss.) Reichb.] A, common; upland or less moist seeps; Apr.

*Sagina apetala Ard. A; dried thin or rocky soil of disturbed areas; Apr.

Sagina decumbens (Ell.) T. & G. subsp. occidentalis (Wats.) Crow. [S. occidentalis
Wats.] A, common; scattered in low or disturbed areas of Tuscan loam or deeper
soils; Apr.

*Silene gallica L. A, occasional; disturbed areas; Apr.

*Spergularia bocconii (Scheele) Foucaud. A, occasional; disturbed areas; Apr.

*Spergularia rubra (L.) J. & C. Presl. A; disturbed, low and dried areas; Apr-May.

*Stellaria media (L.) Vill. A; disturbed areas; Apr.

Chenopodiaceae

*Chenopodium vulvaria L. A; disturbed Tuscan loam; Jul.

220 MADRONO [Vol. 34

Convolvulaceae

*Convolvulus arvensis L. P, abundant; disturbed areas, becoming invasive in pools;
May-Aug.

Crassulaceae

Crassula connata (Ruiz. & Pav.) Berger var. eremica (Jepson) Bywater & Wickens
[C. erecta H. & A.] A; dried hogwallows, outcrops, and thin soil of upland; Feb-
Mar.

Crassula saginoides (Maxim.) Bywater & Wickens [Tillaea drummondii Torr. & Gray
var. bolanderi (Wats.) Jepson. A; hogwallows and seeps; Mar-Apr.

*Crassula tillaea Lester-Garland [C. muscosa (L.) Roth] A; shallow depressions on
upland; Tuscan loam; Mar-Apr.

Parvisedum pumilum (Benth.) Clausen. A; outcrops on Tuscan soils; Apr-May.

Cuscutaceae

Cuscuta howelliana Rubtzoff. A, uncommon; hogwallows and shallow, dried pools;
parasitic on Navarretia leucocephala, Boisduvalia cleistogama, Eryngium vaseyi
var. vallicola, and Downingia species; May-Jun.

Elatinaceae

*Elatine heterandra Mason. A, common; wet mud and sand of seeps; May—Jun.

Euphorbiaceae

Chamaesyce glyptosperma (Engelm.) Small. [Euphorbia glyptosperma Engelm.] A;
found at one site: disturbed soil near an irrigation ditch at crossing under Highway
99; Jun.

Chamaesyce hooveri (Wheeler) Burch. [Euphorbia hooveri Wheeler] A, rare; dried
vernal pools; Jun—Sep.

*Chamaesyce maculata (L.) Small [Euphorbia maculata L.] A; found in a hogwallow
on Tuscan loam; Aug.

Chamaesyce ocellata (Dur. & Hilg.) Millsp. [Euphorbia ocellata Dur. & Hilg.] A,
common; pools, hogwallows, or flat upland; Jul—Oct.

Eremocarpus setigerus (Hook.) Benth. A, ubiquitous; dried vernal pools and flat
upland; thin Tuscan or clay loam; Jun—Aug.

Fabaceae

Astragalus gambelianus Sheld. A, occasional; upland; Tuscan or deeper soil; Mar-—
Apr.

Astragalus pauperculus Greene. A, rare; thin soil of well-drained Tuscan upland;
Mar-Apr.

Lupinus bicolor Lindl. subsp. pipersmithii (Heller) D. Dunn. A; upland, drained sites
or shallow hogwallows; thin Tuscan or deeper soils; Mar-Apr.

Lupinus nanus Dougl. in Benth. subsp. apricus (Greene) Ell., Hard., & Mank. [L.
vallicola Heller subsp. apricus (Greene) D. Dunn.] A; upland; Tuscan loam; Mar-—
Apr.

Lupinus polycarpus Greene. A; upland; Tuscan loam; Mar-Apr.

Lupinus subvexus C. P. Sm. A, occasional; low areas of Anita clay; Apr-May.

* Medicago polymorpha L. var. polymorpha. A, common, upland, edges of hogwallows,
pools, and outcrops; Mar-Apr.

* Medicago polymorpha L. var. brevispina (Benth.) Heyn. A; with var. polymorpha;
Mar-Apr.

Trifolium albopurpureum T. & G. A, occasional; upland; Tuscan loam; Apr—May.

Trifolium amplectens T. & G. A, common; Tuscan loam or deeper soils; Apr-May.

1987] BROYLES: VINA PLAINS PRESERVE 224

Trifolium depauperatum Desv. A, abundant; flat or drained areas of upland; Tuscan
or deeper soils; Mar—Apr.

*Trifolium fragiferum L. P; damp edges of irrigation ditches and disturbed areas;
Mar-Apr.

*Trifolium hirtum All. A; upland; thin, baked Tuscan loam or clay loam; Apr-May.

Trifolium microcephalum Pursh. A, uncommon; Tuscan loam; May.

*Trifolium repens L. P; damp areas along irrigation ditches; Jun—Jul.

Trifolium tridentatum Lindl. var. tridentatum. A; upland or less moist areas of seeps;
Tuscan loam or deeper soils; Apr—May.

Trifolium variegatum Nutt. A; edges of irrigation ditches; Apr-May.

Gentianaceae

Centaurium floribundum (Benth.) Rob. A; less moist portions of seeps; often forms
a zone between more typical seep species and those of dry upland; Jun—Aug.

Centaurium venustum (Gray) Rob. subsp. abramsii Munz. A; open, flat upland;
Tuscan or Keefers loam; Jun—Jul.

Cicendia quadrangularis (Lam.) Griseb. A; open flats of upland, and depressions;
Mar-Apr.

Geraniaceae

*Erodium botrys (Cav.) Bertol. A, common outside of wettest areas; Feb—Apr.

*Frodium brachycarpum (Godr.) Thell. [E. obtusiplicatum (Maire, Weiller & Wilcz.)
T. J. Howell] A, abundantly weedy outside of wettest areas; Mar—May.

*Erodium cicutarium (L.) L’Her. A, less common than E. brachycarpum; upland in
disturbed areas but outside of wettest parts; Feb—Apr.

*Erodium moschatum (L.) L’Her. A, least common Erodium species; disturbed areas;
Apr-May.

*Geranium dissectum L. A, occasional on upland, more common near seeps or vernal
pools on deeper soil; Apr—Jun.

Hydrophyllaceae

Nemophila pedunculata Dougl. ex. Benth. A; upland; thin Tuscan loam, on fresh soil
of pocket gopher mounds; Apr.

Hypericaceae

Hypericum anagalloides Cham. & Schlecht. A, occasional; along edge of irrigation
ditch in rich clay loam; Jul.

Lamiaceae

Pogogyne zizyphoroides Benth. A, common in hogwallows, occasional on upland,
and later in pools; Apr—May.
Trichostema lanceolatum Benth. A; upland; thin Tuscan loam; Aug-—Sep.

Limnanthaceae

Limnanthes douglasii R. Br. var. rosea (Hartw. in Benth.) C. T. Mason. A, common;
vernal marshes and hogwallows or edges of pools; Feb—Apr.

Lythraceae

Ammannia coccinea Rottb. A; wet mud along irrigation ditches; May—Aug.
Lythrum hyssopifolia L. A, common; wet mud of irrigation ditches; May—Aug.

229 MADRONO [Vol. 34

Malvaceae

*Malva nicaeensis All. A; disturbed area around barn; Tuscan loam; Jun.

Sidalcea diploscypha (T. & G.) Gray. A, common but widely scattered on upland
slopes or near vernally wet flats; May—Jun.

Sidalcea hartwegii Gray ex Benth. A; scattered on upland; Tuscan loam; Apr.

Sidalcea hirsuta Gray. A, occasional along irrigation ditches, sometimes abundant
in pools; May-Jun.

Martynaceae

*Proboscidea louisianica (Mill.) Thell. A, common in larger vernal pools, occasional
near irrigation ditches and near barn; Jun—Jul, and in Sep after late rain.

Molluginaceae

*Mollugo verticillata L. A; edges of drying pools, seeps, and hogwallows, especially
where soil is rocky, sandy, or gravelly; late May—Aug.

Onagraceae

Boisduvalia cleistogama Curran. A, common in pools and vernal marshes, occasional
in hogwallows; May-Jun.

Boisduvalia densiflora (Lind1.) Wats. A, occasional; seeps; Jun—Aug.

Boisduvalia glabella (Nutt.) Walp. A, occasional; small pools and hogwallows; Jun—
Jul.

Boisduvalia stricta (Gray) Greene. A, common; seeps, hogwallows, and edges of pools;
Apr-May.

Clarkia purpurea (Curt.) Nels. & Macbr. subsp. quadrivulnera (Doug.) Lewis & Lewis.
A, occasional; upland; Tuscan loam; May-Jun.

Ludwigia palustris (L.) Ell. P, occasional; standing water of irrigation ditches; Aug.

Ludwigia peploides (HBK.) Raven. P, common; standing or slowly moving water of
irrigation ditches; May—Aug.

Papaveraceae

Eschscholzia lobbii Greene. A, uncommon; upland; Tuscan loam; Mar.

Plantaginaceae

Plantago bigelovii Gray. A; upland, hogwallow or margins of pools, especially on
thinner soils; Mar.

*Plantago coronopus L. A, occasional; Tuscan loam; May.

Plantago erecta Morris. [P. hookeriana F. & M. var. californica (Greene) Poe.] A,
common except in wettest habitats; Mar—Apr.

*Plantago lanceolata L. P, occasional between dry upland and seep; Jun.

Polemoniaceae

Gilia tricolor Benth. A, common; upland; Tuscan loam or deeper soils; Apr-May.

Linanthus bicolor (Nutt.) Greene. A, common; upland; Tuscan loam; Mar-Apr.

Navarretia heterandra Mason. A; upland; low but drained Tuscan loam or deeper
soils; May.

Navarretia intertexta (Benth.) Hook. A; low areas; Tuscan loam or deeper soils; May.

Navarretia leucocephala Benth. A, common; hogwallows, pools, and vernal marshes;
Mar-—Jun.

Navarretia nigellaeformis Greene. A; found at only one site on Tuscan loam; May.

Navarretia pubescens (Benth). H. & A. A; scattered on upland; Tuscan loam; May-—
Jun.

Navarretia tagetina Greene. A; scattered on upland; Tuscan loam; May-Jun.

1987] BROYLES: VINA PLAINS PRESERVE 225

Polygonaceae

Chorizanthe polygonoides T. & G. A; upland; thin Tuscan loam; Apr-May.

*Polygonum aviculare L. A; upland; disturbed Tuscan loam; Jun—Sep.

Polygonum californicum Meissn. A; upland; disturbed Tuscan loam; Jun.

Polygonum hydropiperoides Michx. var. asperifolium Stanf. P; bordering seeps; May.

*Rumex crispus L. P, becoming common on disturbed sites of upland and seeps, and
invading some pools; May-Jun.

Portulacaceae

Calandrinia ciliata (R. & P.) DC. var. menziesii (Hook.) Macbr. A, occasional; drained
or seep habitats; Mar.

Claytonia perfoliata Willd. [Montia perfoliata (Donn.) Howell var. perfoliata] A; a
damp, disturbed site on Keefers loam; Apr.

Montia fontana L. subsp. amphoritana Sennen. [H. hallii (Gray) Greene] A; vertical
north surface of exposed fanglomerate near an irrigation ditch; Mar—Apr.

Montia linearis (Doug.) Greene. A; a large, shallow hogwallow; Anita clay; Mar.

*Portulaca oleracea L. A, occasional; margins of pools or in hogwallows; Jul.

Primulaceae

*Anagallis arvensis L. A; hogwallows, vernal marshes, and margins of ditches on
moist soil; Mar—Apr.

Anagallis minima (L.) Krause. A; seeps and pool margins; May.

Dodecatheon clevelandii Greene subsp. patulum H. J. Thomps. P, abundant; upland;
Tuscan loam; Jan—Mar.

Ranunculaceae

Delphinium variegatum T. & G. f. emiliae (Greene) Ewan. P; upland; Tuscan loam;
Mar-Apr.

Myosuros minimus L. var. filiformis Greene. A; hogwallows; Mar-Apr.

Myosuros minimus L. subsp. apus (Greene) Campb. var. sessiliflorus (Huth.) Campb.
A; scattered in low places; Apr.

Ranunculus aquatilis L. var. hispidulus E. Drew. P, common; irrigation ditches; Mar-—
Apr.

*Ranunculus muricatus L. A, common on moist banks of ditches, scattered in low
vernally wet areas; Apr.

Ranunculus occidentalis Nutt. var. eisenii (Kell.) Gray. P, occasional between seep
and upland; Tuscan loam; Jun.

Ranunculus pusillus Poir. A; seeps; May.

*Ranunculus sceleratus L. A, uncommon; along irrigation ditches; Apr.

Rosaceae

Alchemilla occidentalis Nutt. A, occasional; upland; thin Tuscan or deeper soils; Mar.

Rubiaceae

*Galium aparine L. A; found at one site, on disturbed Tuscan loam; Jul.

Saxifragaceae

Saxifraga nidifica Greene. A, occasional on upland slopes; Mar.

Scrophulariaceae

*Dopatrium junceum (Roxb.) Buch.-Ham. in Benth. A; standing or slow-moving water
of irrigation ditches; Jul.

224 MADRONO [Vol. 34

Limosella acaulis Ses. & Moc. A, abundant; mud or deeper water of irrigation ditches;
May.

Lindernia dubia L. var. anagallidea (Michx.) Cooperider. A; mud of irrigation ditches;
Aug.

Mimulus guttatus Fisch. ex DC. A or P, common in seeps, less common at margins
of pools; Apr—Aug.

Mimulus tricolor Hartw. ex Lindl. A; hogwallows and margins of pools; Tuscan loam
or deeper soils; Apr.

Orthocarpus attenuatus Gray. A; upland; Tuscan loam; Mar-Apr.

Orthocarpus erianthus Benth. A, abundant; upland; Feb—Apr.

*Verbascum blattaria L. B or P; found at one site, on disturbed soil; Jul.

Veronica peregrina L. subsp. xalapensis (HBK.) Penn. A, abundant; seeps, hogwal-
lows, and vernal marshes; Mar—May.

*Veronica persica Poir. A; found at one site, on disturbed soil; Mar.

Solanaceae

Physalis angulata L. var. lanceifolia (Nees) Waterfall. A, occasional; upland; Tuscan
loam; Jul-Sep.

Violaceae

Viola douglasii Steud. P, occasional; upland; Mar.

Zygophyllaceae

*Tribulus terrestris L. A, uncommon; disturbed sites; Jul-Sep.

ANTHOPHYTA— MONOCOTYLEDONEAE
Alismataceae

Alisma triviale Pursh. P; in a few places in seeps, standing water of irrigation ditches;
Jul-Aug.

Echinodorus rostratus (Nutt.) Engelm. A, occasional; sandy mud of seeps along ir-
rigation ditches; Aug.

Sagittaria calycina Engelm. P, occasional; seeps of irrigation ditches; Aug.

Amaryllidaceae

Allium amplectens Torr. P, abundant; upland; Apr.

Brodiaea californica Lindl. P, common; upland, including minor drainages, near
seeps, and in thin, gravelly or disturbed soil; May—Jun.

Brodiaea coronaria (Salisb.) Engler. P, occasional; upland; May.

Brodiaea elegans Hoover. P, common; well-drained upland sites; May.

Brodiaea minor (Benth.) S. Wats. P, abundant; thin soils; Apr—May.

Dichelostemma multiflorum (Benth.) Heller. [Brodiaea multiflora Benth.] P, common
and widespread; Apr—May.

Dichelostemma pulchellum (Salisb.) Heller. [Brodiaea pulchella (Salisb.) Greene] P,
common; Mar.

Triteleia hyacinthina Greene. [Brodiaea hyacinthina (Lind1.) Baker] P, common; Tus-
can or deeper soils; Apr. Some specimens appear to be intermediate between this
species and 7. /ilacina; there are no clearcut differences in habitat.

Triteleia laxa Benth. [Brodiaea laxa (Benth.) Wats.] P, common; upland; Apr-May.

Triteleia lilacina Greene. [Brodiaea hyacinthina (Lindl.) Baker var. greenei (Hoov.)
Munz.] P; Anita clay loam. Phenology as for 7. hyacinthina (see for comments).

Cyperaceae

*Cyperus difformis L. A, occasional along seeps; Jul.
Cyperus eragrostis Lam. P, common along seeps; May-Jun.

1987] BROYLES: VINA PLAINS PRESERVE 225

Cyperus niger R. & P. var. capitatus (Britton) O’Neill. P, occasional along seeps; Jun—
Jul.

Cyperus strigosus L. P; seeps; Jun—Jul.

Eleocharis acicularis (L.) R. & S. P; seeps; Jun—Jul.

Eleocharis bella (Piper) Svenson. A; seeps; May-Jun.

Eleocharis macrostachya Britton. P, abundant along seeps and margins of some pools
(does not flower in the latter); Apr.

Scirpus acutus Muhl. P, occasional along ditches; not seen to flower.

*Scirpus mucronatus L. P; seeps; Jul.

Juncaceae

Juncus acuminatus Michx. f. sphaerocephalus Herm. A or P; margins of irrigation
ditches in rich, clay loam; May-Jun.

Juncus balticus Willd. P; habitat as for J. acuminatus; May.

Juncus bufonius L. A; seeps and margins of pools and hogwallows; May.

Juncus dubius Engelm. P, occasional; margins of irrigation ditches in rich, clay loam;
Jul.

Juncus uncialis Greene. A, uncommon; hogwallows, seeps, and pools; Apr.

Lilaeaceae

Lilaea scilloidea (Poir.) Haum. A, common; seeps, emergent in ditches or drainage
of pools; May—Jun.

Liliaceae

Calochortus luteus Doug. ex Lindl. P, uncommon; upland; Tuscan loam; Apr.

Chlorogalum angustifolium Kell. P; upland; Tuscan loam, clay, or Keefers loam; Apr-—
May.

Chlorogalum pomeridianum (DC.) Kunth. P, occasional; upland; May-Jun.

Fritillaria pluriflora Torr. in Benth. P; occasional populations on upland, heavier clay
soils and nearly always in association with Zigadenus fremontii;,; Mar.

Odontostomum hartwegii Torr. P, common; upland; Tuscan loam or deeper soils;
May.

Zigadenus fremontii Torr. P, abundant; low areas of upland in heavier soils; Feb—
Mar.

Poaceae

*Agrostis avenacea Gmel. A, occasional; upland; May-Jun.

*Aira caryophyllea L. A; upland; Mar-Apr.

Alopecurus carolinianus Walt. A, occasional; upland; May.

Alopecurus saccatus Vasey. A; vernal marshes, pools, and hogwallows; Mar-Apr.

Aristida oligantha Michx. A; upland; Jul.

*Avena barbata Brot. A, common; upland or flat places; Mar—Apr.

*Avena fatua L. A, uncommon; upland; Apr—May.

*Briza minor L. A; upland; Apr—May.

*Bromus diandrus Roth. A; upland; Mar-Apr.

*Bromus madritensis L. A, occasional; upland; May.

*Bromus mollis L. A, common; widespread but infested with smut where invading
small pools; Mar—Apr.

*Bromus rubens L. A, common; upland; Mar-Apr.

*Crypsis schoenoides (L.) Lam. [Heleochloa schoenoides L.] A; margins of pools, vernal
marshes; Jun.

*Crypsis vaginiflora (Forsk.) Opiz. [C. niliaca Fig. & DeNot.] A; habitat similar to
C. schoenoides; Jul.

*Cynodon dactylon (L.) Pers. P, common in disturbed areas; Apr.

226 MADRONO [Vol. 34

Deschampsia danthonioides (Trin.) Munro ex Benth. A, common; hogwallows, small
pools, and vernal marshes; Apr.

Diplachne fascicularis (Lam.) Beauv. [Leptochloa fascicularis (Lam.) Gray] A, abun-
dant; seeps; Jun—Jul.

*Echinochloa colonum (L.) Link. A, abundant; seeps; Jul.

*Echinochloa crusgalli (L.) Beauv. var. crusgalli. Similar to E. colonum.

*Echinochloa crusgalli (L.) Beauv. var. oryzicola (Vasing) Ohwi [E. oryzicola (Vasing)
Vasing] Occasional; seeps; phenology as for E. colonum.

*FEragrostis cilianensis (All.) E. Mosher. A, abundant; seeps; Jul.

*Gastridium ventricosum (Gouan) Schinz. & Thell. A; upland or hogwallows; May.

*Hordeum geniculatum Allioni. [H. hystrix Roth.] A; upland or seep; Tuscan loam;
Apr-May.

*Hordeum leporinum Link. A; upland, especially disturbed areas; Apr—May.

*Koeleria phleoides (Vill.) Pers. A; upland, especially outcrops; Apr-May.

*Lolium multiflorum Lam. P, common; upland or seep; Apr—May.

*Lolium perenne L. P; upland or seep; Jun.

Melica imperfecta Trin. A, uncommon; upland; Apr.

Orcuttia pilosa Hoover. A, rare; dried vernal pools; abundant in large pools; Jun—
Jul.

Orcuttia tenuis Hitchc. A, rare; dried vernal pools; found in one pool, where it was
abundant; May, or Jul after late rain.

Panicum dichotomiflorum Michx. A, common; muddy margins of irrigation ditches;
Jun—Jul.

*Paspalum dilatatum Poir. P; mud or standing water of irrigation ditches; Jun—Jul.

Paspalum paspaloides (Michx.) Scribn. [P. distichum L.] P; muddy margins of irri-
gation ditches; Jun.

*Phalaris paradoxa L. A; upland; Tuscan loam or deeper soils; May.

*Poa annua L. A; upland, outcrop, and shallow pools and hogwallows; Mar—May.

Poa scabrella (Thurb.) Benth. ex Vasey. P, occasional; Anita clay; May.

Poa tenerrima Scribn. P; upland, especially near or in hogwallows; Tuscan loam;
Mar-Apr.

*Polypogon interruptus HBK. P; found at one site, near a seep on Tuscan loam; May.

*Polypogon maritimus Willd. A, common; damp areas near irrigation ditches; Tuscan
or Keefers loam; May.

*Polypogon monspeliensis (L.) Desf. A, common near seeps and in hogwallows; Tus-
can loam or deeper soils; May.

Scribneria bolanderi (Thurb.) Hack. A, occasional; Tuscan loam; Apr.

*Sorghum halapense (L.) Pers. P, occasional; gravelly areas; Jun.

Stipa pulchra Hitchce. P, occasional on upland Tuscan loam, more common on deeper,
clay soils; Apr-May.

Tuctoria greenei (Vasey) J. Reeder [Orcuttia greenei Vasey] A, rare; dried vernal
pools; common but not abundant in pools on Preserve; Jul.

*Taeniatherum caput-medusae (L.) Nevski. [T. asperum (Simonkai) Nevski, Elymus
caput-medusae L.] A; scattered on upland, especially disturbed areas; Tuscan loam
or Anita clay; May.

*Vulpia bromoides (L.) S. F. Gray [Festuca dertonensis (All.) Asch & Graebn.] A;
upland or near seep or hogwallow; Tuscan loam; Apr.

Vulpia microstachys (Nutt.) Benth. var. ciliata (Beal) Lonard & Gould. [Festuca
eastwoodae Piper] A, common; well-drained upland; Apr.

Vulpia microstachys (Nutt.) Benth. var confusa (Piper) Lonard & Gould. [Festuca
confusa Piper] A, common; thin soils of upland; Apr.

*Vulpia myuros (L.) K. C. Gmelin var. hirsuta Hack. [Festuca megalura Nutt.] A,
common; upland; Tuscan or deeper soils; Mar—Apr.

*Vulpia myuros (L.) D. C. Gmelin var. myuros [Festuca myuros L.] A, common;
similar to V. myuros var. hirsuta; Mar-Apr.

1987] BROYLES: VINA PLAINS PRESERVE psoag |

Potomogetonaceae

Potomogeton diversifolius Raf. P; slow-moving water of irrigation ditches; May.

Typhaceae

Typha angustifolia L. P, occasional; irrigation ditches; Jun.
Typha latifolia L. P, occasional; irrigation ditches; Jun.

ACKNOWLEDGMENTS

I am grateful to Robert Schlising for his continuing encouragement and help. I also
thank James Jokerst for help in identifying the Juncaceae and Cyperaceae. The field
work was supported in part by The Nature Conservancy.

LITERATURE CITED

BARTOLOME, J. W. and B. GEMMILL. 1981. The ecological status of Stipa pulchra
(Poaceae) in California. Madrono 28:172-184.

, S. E. KLUKKERT, and W. J. BARRY. 1986. Opal phytoliths as evidence for
displacement of native Californian grassland. Madrono 33:217—222.

HARwoop, D., E. KELLEY, and M. DouKAs. 1981. Chico Monocline and north-
eastern part of the Sacramento Valley, California [Geological]. U.S. Geol. Surv.
No. 1-1238.

HOLLAND, R. F. 1976. The vegetation of vernal pools: a survey. /n S. K. Jain, ed.
Vernal pools: their ecology and conservation: proceedings of a symposium. In-
stitute of Ecology Publ. No. 9:11-14.

1981. Botanical study of the Dozier Tract of the Jepson Prairie Preserve,

Solano County, California. Report prepared for The Nature Conservancy, 156

Second St., San Francisco, CA.

. 1982. Plant and wildlife study of Maidu Park. Prepared for City of Roseville,
316 Vernon St., Roseville, CA.

JOKERST, J. D. 1983. The vascular plant flora of Table Mountain, Butte County,
California. Madrono 30(Suppl.):1—18.

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

1968. Supplement to A California flora. Univ. California Press, Berkeley.

SCHLISING, R. A. 1984. Boraginaceae of Butte County, California. Studies From the
Herbarium, California State Univ. Chico, No. 1.

, and E. L. SANDERS. 1983. Vascular plants of the Richvale Vernal Pools,
Butte County, California. Madrono 30(Suppl.):19—30.

SMITH, J. P. and R. York. 1984. Inventory of rare and endangered vascular plants
of California. Calif. Native Plant Soc. Spec. Publ. No. 1 (3rd ed.).

UNITED STATES DEPARTMENT OF AGRICULTURE. 1967. Soil Survey: Tehama County,
California. U.S. Govt. Printing Off., Washington, DC.

(Received 28 Apr 1986; revision accepted 15 Dec 1986.)

SOLIVA (ASTERACEAE: ANTHEMIDEAE)
IN CALIFORNIA

MARTIN F. RAy
Department of Biological Sciences, Stanford University,
Stanford, CA 94305-2493

ABSTRACT

Soliva sessilis, S. pterosperma, and S. daucifolia have been distinguished from each
other on achenal characteristics and habitat, and have been listed as members of the
naturalized flora of California. Examination of over 310 collections from California
documented a continuum of achene morphology and an indiscriminate distribution
of morphs that form a single taxonomic species. Synonymy of S. sessilis also is
extended to include S. neglecta and S. valdiviana.

Soliva Ruiz Lopez & Pavon is a genus of low growing annuals
first described in 1794 from Chile. Native to South America, it was
naturalized in California by 1836 when Nuttall visited Santa Barbara
and collected what he later (1841) described as S. daucifolia. Soliva
may have come to California from Chile in shipments of hides
(Cabrera 1949, Healy 1953), but probably not on livestock (Raven
1963).

Achenes of Soliva are well adapted for long-distance dispersal, as
suggested by their unique morphology. They are small and light-
weight with stiff, appressed pubescence, and disperse by adhering to
animals or other objects that move. More importantly, however,
they have a sharp, persistent stylar spine that easily becomes imbed-
ded in dispersal agents. So/iva is found most often in hard-packed
soil or near well-beaten paths or roadsides. In addition, it is found
in planted lawns. Human activities are probably a major means of
dispersal for Soliva, which occurs mainly in areas with large human
populations or along major travel routes. Soliva spp. are adventive
and have become established world-wide in many such locations.

Soliva sessilis, S. pterosperma, and S. daucifolia have been rec-
ognized for California (Crampton 1954), although two other species,
S. neglectaand S. valdiviana, could be recognized based on published
descriptions by Cabrera (1949) and Philippi (1864-65). This group
of species was called subgenus Eusoliva (=Soliva; see Voss et al.
1983) by Cabrera (1949) and includes about half of the genus. The
remaining five species do not occur in North America and differ
markedly from the species considered here in both achene and veg-
etative characters (Cabrera 1949). Cabrera (1949) treated S. dau-
cifolia as a synonym of S. sessilis, and distinguished between it and
the remaining species only by means of achene characters. Crampton

MADRONO, Vol. 34, No. 3, pp. 228-239, 1987

1987] RAY: SOLIVA 229

(1954) distinguished between S. sessilis, S. pterosperma, and S. dau-
cifolia by achene characters and habitat. All published descriptions
for the five taxa are basically identical with regard to vegetative and
floral morphology; both kinds of characters are deemphasized be-
cause they are not used to distinguish the species. Floral morphology
is of limited use as a descriptive or distinguishing character because
of the short duration and minute size of the flowers.

The generally accepted (Cabrera 1949, Crampton 1954, Munz
1959) achene morphologies that characterize the five taxa under
discussion are illustrated in Fig. | (A-E). Achenes of S. sessilis are
pubescent and have wide, entire wings that usually have large wing-
tips. Achenes of S. pterosperma are pubescent and have wide wings,
long, tapering, curved tips, and a large sinus between upper and
lower wing-lobes that is positioned about one-third of the distance
from the base to the top of the wing (excluding the wing-tip). Achenes
of S. daucifolia are pubescent, have no wings, but have small wing-
tips. Those of S. neglecta are similar to those of S. sessilis, except
they are glabrous rather than pubescent. Soliva valdiviana usually
includes plants with wingless, wing-tipless, glabrous achenes (Ca-
brera 1949), but Philippi’s (1864-65) original description does not
specifically mention a lack of pubescence and refers to the presence
of wing-tips. Notes on many specimens of Soliva suggest that the
achene characters are unreliable. This paper presents an examination
of Soliva collections from California to determine the species pres-
ent, and a study of achene and general morphology to review the
distinguishing characteristics for the species considered.

MATERIALS AND METHODS

To examine achene and vegetative morphology and determine
which species of So/iva occur in California, I studied over 250 spec-
imens from CAS, CHSC, DS, JEPS, POM, RSA, and UC. I also
made 60 collections of So/iva in various parts of the state. A list of
specimens examined in this study is included in Ray (1984), which
is available at Stanford University and CAS.

Some terms used in this paper relative to achene morphology
are illustrated in Fig. 1 (F—R). Wing-tips are usually pointed pro-
jections that occur in pairs on either side of the central stylar spine
(see Fig. 1 and below), and often the outside edges are continuous
with those of the wing. The wing is a region flattened in the plane
of the achene that occurs on either side of the achene. The stylar
spine, derived from the persistent style, arises from the center of the
achene and is continuous with the central, thicker body of the achene.
A sinus is a region where the edge contour of the wing is broken
sharply, like a “bite’’ out of the wing, or sometimes appears as a
crack. An incurved region of the wing is a kind of sinus with a very

230 MADRONO [Vol. 34

why
Youn

N\yvliy

Fic. 1. Achene morphologies for five ‘“‘taxa”’ of Soliva (after Cabrera 1949, Cramp-
ton 1954). A. S. daucifolia. B. S. sessilis. C. S. neglecta. D. S. valdiviana. E. S.
pterosperma. Morphological terms used in the text: F. Long wing-tip. G. Short wing-
tip. H. Typical short, stiff, appressed pubescence on achene “body” and wings. I.
Sinus. J. Narrow sinus. K. “Below” (lower lobe). L. ““Above’”’. M. Stylar spine. N.
Incurved region of wing. O. Wing entire. P. Achene “body” (in this case glabrous).
Q. Narrow wing (=margin). R. Hyaline area of wing (thin and transparent).

shallow, smooth interruption of the wing edge contour.

Although vegetative morphology has not been used in the liter-
ature to distinguish the species of Soliva under consideration here,
I examined vegetative characters in view of possible relationships
to achene characters. These characters included typical habit and
size of plants; leaf position, shape, divisions, surface texture, and
pubescence; internode characters; and aspects of the inflorescence.

I examined achenes from each collection by stereoscope. One or
more (depending on variation occurring on sheet) representative
achenes were drawn in detail and described. Achene body and total
length were measured, because this character has been used to rec-
ognize S. daucifolia (Crampton 1954). Vegetative characteristics from

1987] RAY: SOLIVA 251

each collection were described. Based on the accumulated data, 15
achene morphology categories (Fig. 2) were designated, each rep-
resented by a specimen. I then assigned all specimens to one or more
(depending on variation) of these morphological categories.

The range of So/iva achene variation is described here by reference
to the above mentioned series of 15 numbered, artificial achene
morphs. The features of each morph are described in detail in the
caption for Fig. 2. Although the artificial morphs are discrete, many
further intermediates occur, and in fact every achene is somewhat
unique. The morphs are not necessarily spaced equally over the range
of variation. Some morphs are of more general form than others,
and thus contain more internal variation (intermediates) and rep-
resent more collections. Achene morphology was not always uniform
in a particular collection and I noted many cases in which variation
occurred on individual plants or even within capitulae. The generally
recognized species (Cabrera 1949, Crampton 1954) correspond ap-
proximately with the artificial achene morphs (Fig. 2) as follows: S.
valdiviana, morphs 1-2; S. daucifolia, morphs 4—6; S. sessilis, morphs
9-10; and S. pterosperma, morphs 13-15. Soliva neglecta is found
among S\. sessilis morphs. In the following text, the phrase “‘achene
morph” or “‘achene morph number’”’ refers to an actual observed
morphology corresponding to that particular numbered artificial
morph from Fig. 2.

RESULTS AND DISCUSSION

Achene morphology. Achenes in all specimens have a central por-
tion or “‘body”’ of more or less similar shape, including a sharp,
persistent stylar spine (Fig. 1P), and most also have wings of various
shapes and sizes, and/or wing-tips. A few achenes are wingless and
wing-tipless. A carina on the achene body toward the base on the
convex (abaxial) side is usually more pronounced in the drier col-
lections from late in the season. Achenes and their appendages vary
in color from light green or tan to dark brown. They vary in the
distribution of typically short, stiff, appressed pubescence, and often
have minute purple spots variously distributed on the body, stylar
spine, and wings. The achene body is usually bilaterally symmetrical,
but the two wings and wing-tips sometimes differ from one another
in shape and size. Wings vary in shape, width, thickness, and edge
characters, such as splits, cracks, and sinuses. Some wings are trans-
lucent or hyaline in limited regions. Wing-tips vary in width, length,
curvature, and degree of furcation (Figs. 1, 2). Wing-tips on a given
achene are sometimes dissimilar.

Total achene length varies from 3.5—5.2 mm. Crampton (1954)
reported that S. daucifolia had achenes with consistently shorter

232 MADRONO [Vol. 34

1987] RAY: SOLIVA 233

bodies than those of either S. sessilis or S. pterosperma. I observed
no consistent relationship between achene body length and a par-
ticular achene morphology. Achene body length appeared to vary
with overall length.

I observed no consistent relationship of any vegetative character
or group of such characters to any achene character. In fact, vege-
tative morphology was relatively uniform in all specimens. Some
previously unreported minor details of habit and leaf morphology
were observed, and I have included these in the description below
(compare with Cabrera 1949, Munz 1959).

A number of specimens that I examined corresponded with the
description (with illustration) of S. neglecta Cabrera [e.g., Bacigalupi
1527 (DS), Breedlove 4405 (DS), Cerrate 2515 (UC), Eastwood and
Howell 2561 (CAS), Knight 626 (CAS), Mason 4315 (DS), and Raven
19734 (DS, RSA)]. Achenes from these specimens also usually
matched the generalized morphology of S. sessilis (Fig. 1), but are
glabrous or nearly so. This condition also was found in other col-

—

Fic. 2. Achene variation in Soliva sessilis. Achenes shown are from representative
collections that are cited below and that form the basis for artificial achene morphs,
which describe the range of wing and wing-tip variation. Morphs are numbered 1-
15, and are followed by a description and citation. Compare with Fig. 1, A-E. Note
that each artificial morph contains internal variation; as discussed in the text, some
artificial morphs are circumscribed more broadly in terms of actual achene variation,
than others. Morphs are in a proposed order of complexity of wing and wing-tip
features. Scale in mm is shown with morph 15.

1. Achenes with no wings, no wing-tips; Wiggins 12352 (DS). 2. No wings, vestigial
tips; Tracy 6684 (UC). 3. Wings vestigial above, no tips; Raven 10668 (CAS). 4.
Wings narrow above, absent below, tips evident; Eastwood 143 (UC). 5. Wings
extremely narrow, tips vestigial to evident; Linsdale s.n. (CAS). 6. Wings to 0.25 mm
wide, tips longer than in #5; Raven 6933 (CAS). 7. Wings wider above, smoothly
curving to narrow below, or slightly lobed below; Howell 41480 (CAS). 8. Wings of
medium width, about halfway (ca. 0.75 mm wide) between narrow (#5) and wide
(#9, 10, and beyond) with many edge and tip variations. May be incurved below;
Eastwood and Howell 2561 (CAS). 9. Wings | mm or more wide, rounded, tips long
with a number of variations; Lee and Mason 9105 (UC). 10. Wings wide, more or
less rounded with many edge and tip variations; Howell 42163 (CAS). 11. Wings
wide above, narrower below, with the upper lobes curving into the lower, but the
lower not protruding beyond the edge line once it is vertical such that a sinus is not
formed; Howell 2981 1c (CAS). 12. Wings wide above, curving to small sinuses below,
blending into small lobes that protrude below these, the lobes not as wide as the
wings above; Jepson 18856 (JEPS). 13. Wings wide above and about the same below,
divided by a sinus that is wide and fairly deep. Contour of upper and lower lobes
more or less continuous; many edge and tip variations; Raven 6624 (CAS). 14. Wings
wide above, about the same or less below, divided by a relatively narrow sinus that
is quite deep. Contour of upper and lower lobes more or less continuous; Jepson
18018 (JEPS). 15. Wings wide, upper lobes wider than the lower, divided by a deep
sinus, entire achene rather arrowhead-shaped, the contour of upper and lower lobes
not continuous; Hoover 1996 (JEPS).

234 MADRONO [Vol. 34

lections that exhibited different morphologies [e.g., Ashwin 535 (CAS),
Howell 45554 (CAS), Jepson 11563 (JEPS), Ray 33 (DS), and Thom-
as 7101 (DS)]. Glabrous achenes are found occasionally throughout
the range of wing-character morphology. Degree and distribution of
pubescence also varies over that range. Howell 45554 has both pu-
bescent and glabrous achenes, but not on the same plant. Thus,
pubescence and wing morphology appear to vary independently. In
view of this variability, the distinction between S. sessilis and S.
neglecta is unclear.

The description of S. valdiviana (Philippi 1864-65) includes no
specific reference to presence or lack of achenal pubescence, so it
corresponds with a number of the specimens examined in this study
[e.g., Ashwin 535 (CAS), Howell 21653 (CAS), Koch 812 (UC), Rose
39154 (UC), Tracy 6684 (UC), and Wiggins 12352 (DS, UCO)]. Nut-
tall (1841) described S. daucifolia as “‘slenderly margined” and ‘“‘mi-
nutely bidentate at the summit’. Philippi (1864-65) described SS.
valdiviana as “‘haud alatis” (having no wings) and “‘spinoso coro-
natis” (spined crown). Cabrera (1949) includes an illustration of S.
valdiviana that shows no wing-tips. Because the wings have been an
important descriptive character in the literature, the assumption that
the term “‘margin’’ was used by Nuttall (1841) to refer to the wing
is a logical one. The descriptions “‘slender margin” (1.e., very narrow
wings) and “‘haud alatis”’ (i.e., lacking the wide wings some other
achenes have) are not greatly distinct. The descriptions “minutely
bidentate at the summit” and “‘spinoso coronatis’’ also are similar;
both refer to the small wing-tips that occur in achene morphs 2, 4,
and 5 (Fig. 2). Therefore, the original published descriptions for S.
daucifolia (Nuttall 1841) and S. valdiviana (Philippi 1864-65) are
similar. Significant variation occurs in wings and wing-tips for ar-
tificial morphs 1—5 (Fig. 2), which is the range described for achenes
of both S. daucifolia and S. valdiviana. The distinction between the
two taxa is unclear.

The specimens cited by Crampton (1954) to support recognition
of S. daucifolia differ morphologically from each other and, in some
cases, from Nuttall’s (1841) description. Tracy 1089 (UC) has achenes
with narrow wings and sinuate edges, often wider above, with long
wing-tips; Rose 39154 (UC) and Wiggins 12352 (DS, UC) have
achenes with no wings and no wing-tips (morph 1, Fig. 2); and
Eastwood 143 (UC) has achenes with the widest portion of the wing
above, the wings absent below, and the wing-tips curved sharply
inward (obscured behind the stylar spine in Fig. 2). Crampton’s
(1954) illustration of Eastwood 143 is similar to achenes I examined,
but differs in details of wing morphology. Crampton cited a specimen
he called ““Les Koch 812’, which probably corresponds to Leo F.
Koch 812 (UC). Examples of achene morphs 1-10 were found on
this sheet. Crampton (1954) provided illustrations of Crampton 1121

1987] RAY: SOLIVA 235

and Crampton 1223, with no discussion of these in the text. On
examination, I found achenes from these specimens to correspond
loosely with the illustrations, but they differ substantially in detail.
Apparently the achenes examined by Crampton (1954) were different
from those seen in the present study, which again suggests taxonomic
unreliability in achene characters.

Soliva collections that show variation of achene morphology be-
tween plants on the same sheet or within individual plants or ca-
pitulae are listed below. Each collection is listed with the artificial
achene morph numbers (see Fig. 2) corresponding to achenes found
on that sheet. An asterisk indicates variation within the same plant:
Alameda Co.: Lee 70] (JEPS), 5, 7, 8. Amador Co.: Hansen 1054
(UO), 8, 11, 12, 14*. Butte Co.: Ahart s.n. (CAS), 10, 14*. Humboldt
Co.: Davy 5684 (UC), 9, 11*. Mendocino Co.: Koch 812 (UC), 1-
10. Monterey Co.: Howell 41480 (CAS), 6, 7*. San Mateo Co.: Abrams
2423 (DS), 6, 10, 14; Dudley s.n. (RSA), 11, 12*; Ferris 4157 (DS),
11, 12, 13*; Ray 37 (DS), 7, 12*; Ray 38 (DS), 4, 7, 8*; Ray 40 (DS),
6, 12*, Thomas 4283 (DS), 4, 7, 8*. Santa Clara Co.: Dudley s.n.
(DS), 6, 12; Thomas 4822 (RSA), 8, 11*. Santa Cruz Co.: Ray 36
(DS), 6, 7*. Sonoma Co.: Brandegee s.n. (POM), 5, 10, 14*. Tuol-
umne Co.: Howell 40693 (CAS), 14, 15*; Johannsen 883 (UC), 9,
hal lis:

Dispersal. Humans appear to be a major factor in the spread of
Soliva in its role as an adventive species. It consistently occurs in
either hard-packed paths, waste ground, dirt roads, or cultivated
lawns. In collecting, I observed that achenes easily became imbedded
in my hands. Achenes are probably dispersed on shoes or clothes,
or on the tires of cars or other machines. It seems likely that in lawns
on the Stanford campus, where Soliva occurs frequently and is
spreading, achenes are moved on the tires of large lawnmowers. To
test the hypothesis that achenes can move in tires, I rode a balloon-
tired bicycle through some mature patches of So/iva in a lawn, and
then checked both tires after about 150 m riding distance. Five
achenes were found, two of which were firmly imbedded. This in-
dicates that not only can short distance dispersal occur in tires, but
also longer distance movement because the firmly imbedded achenes
might remain so for some time. Soliva, therefore, is a genus well
adapted to dispersal within an area in which the plants are already
established, and to locations that may be quite remote. Such new
locations are generally in well-travelled areas, where the achenes are
most likely to become detached from the dispersal agent and to be
pressed into the ground by subsequent traffic. The possibility exists
that the distribution pattern in well-travelled and populated areas
results from sampling. That is not sufficient reason to discredit over-
all collection evidence because of the additional strong evidence of

MADRONO

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(fe)
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—_ wo oat io

1987] RAY: SOLIVA 237

achene dispersal adaptations and the many collections from similar
microhabitats.

Distribution. Soliva is well established in watered lawns and plant-
ings in many places in California. There are few collections from
urban southern California (Los Angeles basin and vicinity), although
I have collected Soliva in lawns in Santa Monica, Arcadia, Newport
Beach, and rural Orange Co., and I have observed it growing in
lawns near USC. It is unlikely that the genus is established in many
natural cr unwatered areas in southern California, because of low
seed carryover (Johnson and Lovell 1980) and the relatively long
wet period required for growth. In contrast, Soliva is probably es-
tablished in more locations in the San Francisco Bay region and
northern California than records indicate.

Distribution maps (Fig. 3) show collection locations for many
Soliva specimens examined in this study, with a symbol for each
indicating an artificial achene morph that corresponded most closely
with achenes from that specimen (variation in the same collection
not indicated). The three maps are designed to show the distribution
of each of the 15 artificial achene morphs. This is necessary because
species of Soliva have been recognized by alleged localization of
particular achene morphologies. Examination of the maps (Fig. 3)
may give the impression that achene morphs 11-15 (Fig. 2) occur
more often in the Sierra Nevada foothill areas and that morphs 1-
6 occur more often on the southern coast. Crampton (1954) made
similar observations and used them to support recognition of Soliva
pterosperma (Sierra foothills) and Soliva daucifolia (southern coast).
I observed, however, that all achene morphologies were distributed
randomly and in high density in the San Francisco Bay region, from
which there are more collections than any other region in the state.
Also, morphs 5-8 have been collected in the Sierra foothills (Fig.
3). Because of the low frequency of collections in the Sierra foothills
and the southern areas (relative to the Bay region), and the random
distribution of morphs in the Bay region, localization of particular
achene morphologies in the Sierra or southern areas does not seem
plausible. Thus, there appears to be no separation of achene mor-
phologies throughout the range of So/iva in California.

Conclusion. My observations of achene morphology show that a
continuum of variation exists for the achene wing and wing-tip
characters in the California collections of Soliva. Based on these
observations, and the lack of any other consistent variations or major

—_—

Fic. 3. Distribution of Soliva sessilis in California. Three maps are provided for
clarity in areas of high specimen density. Keys match symbols on maps to artificial
achene morph numbers (1-15) in Fig. 2.

238 MADRONO [Vol. 34

separations of habit or habitat, I conclude that only one species of
Soliva occurs in California. Because this species also occurs in other
parts of the world (such as Australia, New Zealand, and South Amer-
ica), and indeed has been introduced into California, this conclusion
also applies on a wider basis. By priority of publication, Soliva sessilis
is the name for this species, and thus the names S. pterosperma and
S. daucifolia are synonyms. The names S. neglecta and S. valdiviana
also appear to be referable to S. sessilis.

TAXONOMIC TREATMENT

SOLIVA SESSILIS Ruiz Lopez & Pavon, Syst. Veg. Fl. Per. Chil. 215.
1798.—Type: Chile, “‘Habitat in plateis et pratis Conceptionis
(sic) Chile, praesertim ad Mochita, Hualpen, Andalien et Ga-
vilan tractus’’; no specimens seen; description and earlier illus-
tration (Ruiz Lopez & Pavon, Syst. Veg. Fl. Per. Chil. 113, tab.
24, 1794) fix application of the name. See Cabrera (1949) for
additional synonymy.

Gymnostyles pterosperma A. L. Juss., Ann. Mus. Natl. Hist. Nat. 4:
262, tab. 61, f. 3. 1804.—Soliva pterosperma (A. L. Juss.) Less.,
Synop. Gen. Compos. 268. 1832.—Type: Argentina, Buenos
Aires, ““Ex Bonaria. Car. ex sicca in herb. Commers.’’, no spec-
imens seen; illustration fixes application of the name. See Ca-
brera (1949) for additional synonymy.

Soliva daucifolia Nutt., Trans. Amer. Philos. Soc., ser. 2. 7:403.
1841.—Type: California, “*. .. within the limits, and in the im-
mediate vicinity of St. Barbara’’; no specimens seen; description
allows certain application of the name.

Soliva valdiviana Philippi, Linnaea 33:168. 1864—-65.— Type: Chile,
‘‘Frequens in prov. Valdivia’’; no specimens seen; description
allows certain application of the name.

Soliva neglecta Cabrera, Notas Mus. La Plata 14:128. 1949.—Type:
Argentina, Jujuy, Santa Ana, 3100 m, 29 Feb 1940, A. Burkart
and N. S. Troncoso 11665 (LP; isotype: SI) (not available).

Herbaceous annuals with fibrous roots. Ascending, spindly-
stemmed plants to nearly acaulescent plants, or clumpy and compact
plants with glomerate leaves and capitulae, or spreading plants with
decumbent to prostrate stems and elongate internodes. Compact
plants 2—7 cm tall, spreading plants 25 cm diam. Stems 1-10 from
base, light- to dark-colored, often purple-spotted, sparsely pubescent
to villous. Leaves to 5 cm long, petioled, the bases broad, + clasping;
once-pinnate, the pinnae with 2-8 + palmate narrowly lanceolate
lobes, often one lobe smaller, the terminal pinna sometimes single;
puberulent to sericeous or villous. Capitulae sessile in axils; disci-
form; receptacle convex or low-conic; involucre of 5—12 subequal
phyllaries, broadly ovate to lanceolate, abruptly acute, in 1-2 series,

1987] RAY: SOLIVA 239

2-3 mm long, green to hyaline, pubescent to villous; disc flowers 4—
6, perfect, minute, greenish-translucent (yellow stamens within),
4-merous, probably functionally staminate, surrounded by 10-12
naked pistillate flowers. Achenes 3.5—5.2 mm long including stylar
spine, the ovate to lanceolate central body + carinate, style persis-
tent, becoming hard and sharp, stigmas persistent or deciduous.
Achenes wingless (with or without toothlike wing-tips) to wide-
winged, the wings thin, opaque to regionally hyaline, the edges
notched, incurved, split, cracked, or sinuate; the wing-tips from
short, blunt, toothlike projections to long curving tips with thin
edges; achenes light green to dark brown, often with minute purple
spots, glabrous to variously pubescent. Probably self-fertile. Feb—
Jul. Disturbed, hard-packed and weedy areas that receive sufficient
water for seed set, especially paths, dirt roads, roadsides, and other
well-travelled areas; also in watered lawns.

ACKNOWLEDGMENTS

I thank the following individuals for their assistance: Drs. L. R. Heckard, P. H.
Lovell, Margery Marsden, Peter M. Ray, Kingsley R. Stern, John L. Strother, and
John H. Thomas.

LITERATURE CITED

CABRERA, A. L. 1949. Sinopsis del genero Soliva (Compositae). Notas Mus. La
Plata, Bot. 14:123-139.

CRAMPTON, B. 1954. Observations on the genus So/iva in California. Leafl. W. Bot.
7:196-198.

HEALY, A. J. 1953. Contributions to a knowledge of the naturalized flora of New
Zealand No. 3. Trans. Roy. Soc. New Zealand 81:23-26.

JOHNSON, C. D. and P. H. LOvELL. 1980. Germination, establishment and spread
of Soliva valdiviana. New Zealand J. Bot. 18:487-—493.

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

NuTTALL, T. 1841. Descriptions of new species and genera of plants in the natural
order of the Compositae. Trans. Amer. Philos. Soc. (New Series) 7:403.

PHILIPPI, R. A. 1864-65. Plantarum novarum Chilensium. Linnaea 33:168-169.

RAVEN, P. H. 1963. Amphitropical relationships in the floras of North and South
America. Quart. Rev. Biol. 38:151-177.

Ray, M. F. 1984. The genus Soliva (Asteraceae) in California. M.S. thesis, Stanford
Univ., Stanford, CA.

Voss, E. G. etal. 1983. International Code of Botanical Nomenclature, adopted by
13th International Botanical Congress, Sydney. Regnum Vegetabile 111.

(Received 5 Nov 1984; revision accepted 9 Mar 1987.)

ROLE OF FIRE IN THE GERMINATION OF CHAPARRAL
HERBS AND SUFFRUTESCENTS

JON E. KEELEY
Department of Biology, Occidental College,
Los Angeles, CA 90041

STERLING C. KEELEY
Department of Biology, Whittier College,
Los Angeles, CA 90608

ABSTRACT

Fifty-seven herbaceous and suffrutescent species common after fire in chaparral
were tested for their response to charred wood and heat shock of 120°C for five
minutes. Over half of the species germinated readily without either treatment. These
included all of the herbaceous perennial monocots, most herbaceous perennial dicots,
and a number of annuals. In most species, the heat treatment reduced germination
and only one species was stimulated significantly by heat. Forty-two percent of the
species showed significant enhancement of germination with charred wood. For some
perennials, such as Penstemon spectabilis and Romneya coulteri, and an annual,
Papaver californicum, there was a near obligatory requirement for charred wood.
Significant enhancement of germination in the presence of charred wood is now known
for species in 10 plant families: Asteraceae, Boraginaceae, Brassicaceae, Caryophyl-
laceae, Hydrophyllaceae, Onagraceae, Papaveraceae, Polemoniaceae, Rubiaceae, and
Scrophulariaceae. Several fire-following species, Eucrypta chrysanthemifolia and Di-
centra spp., failed to germinate under any treatment.

Since the early observations of Brandegee (1891), botanists have
been impressed by the often spectacular wildflower displays that
occur the first growing season after wildfire in chaparral. Over 200
species of annuals, herbaceous perennials, and short-lived suffru-
tescents have been recorded from chaparral burns. This abundance
and diversity of herbs is in marked contrast to the generally depau-
perate herbaceous vegetation in mature chaparral (Sweeney 1956,
Stocking 1966, Keeley et al. 1981).

Nearly all of this temporary vegetation arises from seed or veg-
etative parts present in the soil prior to burning. This fact has resulted
in two theories accounting for the breaking of seed dormancy after
fire: 1) seeds are inhibited from germinating by the mature chaparral
vegetation (allelopathy) and fire releases seeds from this inhibition,
or 2) seed germination is stimulated by fire.

The question of allelopathic inhibition of seeds by chaparral has
been examined in numerous s*udies (Sweeney 1956, McPherson and
Muller 1969, Christensen and Muller 1975a,b, Kaminsky 1981, Kee-
ley et al. 1985), but its role in inhibiting seed germination is unclear.

MADRONO, Vol. 34, No. 3, pp. 240-249, 1987

1987] KEELEY AND KEELEY: FIRE AND GERMINATION 241

There is strong evidence, however, that many chaparral species
have seeds that, under natural conditions, require a stimulus from
fire for germination. Germination of some species is stimulated by
heat shock from fire that ruptures the seed coat (Sweeney 1956,
Christensen and Muller 1975a,b, Keeley et al. 1985). Germination
of other fire-following herbs is stimulated by a chemical leached
from charred (but not ashed) wood (Wicklow 1977, Jones and Schle-
singer 1980, Keeley et al. 1985). Many species have been tested for
their germination response to heat shock; however, the vast majority
of chaparral herbs and suffrutescents have not been tested for their
response to charred wood.

Although germination of many chaparral species is apparently
dependent upon one or the other of these fire-related cues, a number
of species are known to germinate readily without such cues (e.g.,
Sweeney 1956, Keeley et al. 1985).

The purpose of this study was to test the germination response of
57 species, representing all of the life-histories and growth forms
present in the temporary postfire vegetation. Specific questions ad-
dressed were: 1) How widespread is charred wood stimulated ger-
mination? 2) For species with charred wood stimulated germination,
will heat shock produce a similar stimulation in germination? 3) To
what extent can generalizations be drawn concerning the relationship
of growth form and germination response?

METHODS

Selection of species was based on availability of plants with mature
seed crops that were present in recently burned chaparral. Collections
were made between elevations of 500-1500 m in Los Angeles, Riv-
erside, San Diego, San Bernardino and Ventura cos., California.
Vouchers have been deposited at LOC (Occidental College). No-
menclature is according to Munz (1974). Seeds of each species were
collected from a single population of 25 or more plants during the
spring and summer of 1982 and stored in paper bags under room
conditions for 14-18 months. Although there is little data on seed
longevity of these species, the fact that many species may be absent
on a site for decades prior to fire suggests the seed pool in the soil
is quite long-lived (Sweeney 1956).

Seeds were sown into 60 <x 15 mm petri dishes filled with 15 g
(fresh weight) of commercial potting soil (Gro-Lite, see Keeley 1984
for chemical analysis of this soil). Charred wood was made by char-
ring (but not ashing) 1-2 cm diameter stems of the chaparral shrub
Adenostoma fasciculatum and grinding in a Wiley Mill to pass a 1
mm screen. Charred wood treatments received 0.5 g of this powdered
charred wood. For comparison, a heating treatment of seeds was
included. The treatment of 120°C for five minutes was selected be-

242 MADRONO [Vol. 34

TABLE 1. GERMINATION OF SELECTED CHAPARRAL HERBS AND SUFFRUTESCENTS IN
RESPONSE TO 120°C FOR FIVE MINUTES OR APPLICATION OF POWDERED CHARRED
WOOD TO THE GERMINATION MEDIUM (N = 5 DISHES OF 50 SEEDS EAcu). A = annual,
B = biennial, Hp = herbaceous perennial, S = suffrutescent, * = non-native. # =
Gilia capitata seeds from two populations were tested, a first year burn and an adjacent
mature chaparral stand. ns = no significant difference between treatments (p > 0.05);
for species with significant difference, treatments with the same superscript are not
significantly different at p > 0.05.

Percentage germination
Growth Con-- 120°C Charred

form trol 5 min wood p
Dicots
Apiaceae
Daucus pusillus (A) 30 102 18? <0.01
Lomatium dasycarpum (Hp) 12 l 4 <0.01
Asteraceae
Agoseris heterophylla (A)
Beaked achenes 79 86 88 ns
Non-beaked achenes 62 78 92 <0.001
Gnaphalium californica (A/B) 46? 672» T9e <0.05
Heterotheca grandiflora (A/B) be 44 86? <0.001
Lactuca serriola (A*) 53 44 54 ns
Madia gracilis (A) 86? 45 86? <0.05
Malacothrix clevelandii (A) 9a 102 35 <0.001
Microseris linearifolia (A) 98 96 95 ns
Perezia microcephala (Hp) 35 l 9 <0.001
Porophyllum gracile (S) hee 26 13 <0.01
Rafinesquia californica (A) 4a 38 55 <0.01
Stephanomeria virgata (A) 454 40 56 <0.05
Boraginaceae
Cryptantha intermedia (A) 58 5aF 74 <0.05
Brassicaceae
Lepidium nitidum (A) 23 Bh 22, <0.001
Sisymbrium orientale (A*) 94 798 748 <0.01
Streptanthus
heterophyllus (A) ite gi: 25 <0.001
Caryophyllaceae
Silene gallica (A*) 642 662 34 <0.01
S. multinervia (A) G* 9a 44 <0.01
Fabaceae
Lotus salsuginosus (A) 62 24? 2 <0.01
L. strigosus (A) 35° 38? 24 <0.05
Hydrophyllaceae
Eucrypta
chrysanthemifolia (A) 0 0 0 ns
Phacelia minor (A) 0? 0? 13 <0.001
Onagraceae

Camissonia californica (A) 32 6 49 <0.001

1987] KEELEY AND KEELEY: FIRE AND GERMINATION 243

TABLE 1. CONTINUED.

Percentage germination

Growth Con- 120°C Charred

form trol 5 min wood p
Clarkia epilobioides (A) 4) 54 75 <0.001
C. purpurea (A) 402 408 [o =().05
C. unguiculata (A) 61 65 68 ns
Papaveraceae
Dicentra chrysantha (Hp) 0 0 0 ns
D. ochroleuca (Hp) 0) 0 0) ns
Papaver californicum (A) 0? 0? 89 <0.001
Romneya coulteri (S) 0? 0? 40 <0.001
Polemoniaceae
Gilia australis (A) 312 323 80 <0.001
G. capitata (A)
Mature chaparral# 8? 22° 83 <0.001
Burned chaparral# 207 25% 69 <0.001
Polygonaceae
Chorizanthe fimbriata (A) 37 29 45 ns
Pterostegia
drymarioides (A) 68 30 47 <0.01
Ranunculaceae
Delphinium cardinale (Hp) 0° Z 402 <0.001
D. parryi (Hp) 68" = 30 61 <0.0
Rubiaceae
Galium angustifolium (Hp/S) De 17 43 <0.001
G. parisiense (A*) 85 89 100 ns
Scrophulariaceae
Antirrhinum
coulterianum (A) 23 3 42 <0.001
A. kelloggii (A) 392 454 63 <0.01
A. nuttallianum (A) 69 56 58 ns
Collinsia parryi (A) 24 12 (a) <0.001
Cordylanthus filifolius (A) We 2h 627 <0.05
Penstemon
centranthifolius (Hp) 0? De 16 <0.001
P. heterophyllus (Hp) 54 4 74 <0.001
P. spectabilis (Hp) ie 33 61 <0.001
Scrophularia
californica (Hp) 823 67 25 <0.001
Solanaceae
Solanum douglasii (S) 823 852 63 <0.01
Amaryllidaceae
Allium praecox (Hp) 18 14 18 ns
Bloomeria crocea (Hp) 5D 46 60 ns

Dichelostemma pulchella (Hp) 100 l 64 <0.001

244 MADRONO [Vol. 34

TABLE |. CONTINUED.

Percentage germination
Growth Con- 120°C Charred

form trol 5 min wood p
Liliaceae
Calochortus concolor (Hp) 84 66 82 <0.05
C. splendens (Hp) 89 9 66 <0.001
Chlorogalum
parviflorum (Hp) 46 15 46 <0.01
Poaceae
Melica imperfecta (Hp) 42 51 34 ns
Stipa lepida (Hp) 7 64 64 ns

cause it stimulates germination of many chaparral herbs (Keeley et
al. 1985). Seeds were heated in a forced convection oven prior to
sowing. For both treatments and a control, in which seeds were not
heated and charred wood was not applied, five replicate petri dishes
of 50 seeds each were tested. The experiment was initiated by ad-
dition of 8 ml of deionized water to all dishes except charred wood
treatments, which received 10 ml because of water absorption.

Seeds of some species require a period of low temperature treat-
ment in order to overcome embryo dormancy. Periods of two weeks
to two months are commonly employed (Atwater 1980), with the
longer periods being required for species from higher elevations and
latitudes. Many chaparral species from southern California do not
require stratification (J. Keeley, unpubl. data). In this investigation,
stratification requirement was not studied; however, all dishes were
maintained at 5°C for three weeks prior to incubation at 23°C for
two weeks, under a 12 hour photoperiod at approximately 350 umol
m~*s_!. Germination was scored after the pre-chilling treatment and
each week at 23°C. To determine if some species might require a
longer cold treatment, this cycle of three weeks cold and two weeks
at 23°C was repeated once before ending the experiment.

Treatments, including controls, were compared with 1-way AN-
OVA on arcsin transformed data and the Student-Newman-Keuls
multiple range test.

RESULTS

Fifty-seven herbs were tested for their response to charred wood
and heat shock (Table 1). Two-thirds of the 22 herbaceous perennial
and suffrutescent species germinated readily under ‘control’ condi-
tions and showed no enhancement with either treatment. These
included all of the monocot species tested. The heat treatment of
120°C for five minutes did not stimulate germination of any of the

1987] KEELEY AND KEELEY: FIRE AND GERMINATION 245

herbaceous perennials, but the possibility of seeds being stimulated
by other heating treatments cannot be ruled out. Heating, however,
tended to reduce germination of many herbaceous perennials. In
these species, heating was apparently lethal because many of the
seeds had rotted by the end of the experiment. Germination of five
herbaceous perennial and suffrutescent species was stimulated sig-
nificantly by charred wood; this response was particularly striking
in Penstemon spectabilis and Romneya coulteri, but also was ob-
served in Galium angustifolium, Penstemon centranthifolius, and P.
heterophyllus.

Germination of 20 of the annual species was enhanced signifi-
cantly by charred wood (Table 1). Some species, e.g., Papaver cal-
ifornicum and Phacelia minor, showed a nearly complete depen-
dence on charred wood. For other species, e.g., Antirrhinum
coulterianum, Camissonia californica, Gilia capitata, Lepidium ni-
tidum, Rafinesquia californica, Silene multinervia, and Streptanthus
heterophyllus, the presence of charred wood resulted in nearly an
order of magnitude greater germination. In others, such as Agoseris
heterophylla, Antirrhinum kelloggii, Collinsia parryi, Clarkia spp.,
Cryptantha intermedia, Gilia australis, Gnaphalium californica,
Malacothrix clevelandii, and Stephanomeria virgata, there was often
substantial ‘control’ germination, but an additional 20-50% ger-
mination with charred wood.

Heat treatment stimulated the germination of Lotus salsuginosus,
but reduced the germination of seven other annuals, including species
with charred wood stimulated germination. Agoseris heterophylla
had polymorphic germination behavior related to achene morphol-
ogy; non-beaked achenes had significantly greater germination with
heat and charred wood treatments in contrast to the beaked achenes.

Several common fire-following species, Dicentra chrysantha, D.
ochroleuca, and Eucrypta chrysanthemifolia, failed to germinate,
despite having seeds that appeared filled and viable (tetrazolium
testing was inconclusive due to the very small or rudimentary em-
bryos characteristic of these species).

Timing of germination was variable and not related clearly to
growth form or germination response. For example, 90% of the total
germination of Calochortus splendens, an herbaceous perennial, had
occurred by the end of the three week pre-chilling treatment; this
pattern also was observed for Dichelostemma pulchella and annuals
such as Gilia capitata, Heterotheca grandiflora, Pterostegia dryma-
rioides, and Rafinesquia californica. Other herbaceous perennials,
e.g., Allium praecox, Bloomeria crocea, Chlorogalum parviflorum,
Penstemon spectabilis and Scrophularia californica, and annuals such
as Silene multinervia and Stephanomeria virgata failed to germinate
in the cold, but the vast majority germinated within the first week
at 23°C. Some species (Lotus strigosus and Silene gallica) had more

246 MADRONO [Vol. 34

or less equal germination percentages at each scoring period through-
out the 10 weeks. Delphinium cardinale was particularly slow to
germinate, none germinated until the second cold treatment eight
weeks after the beginning of the experiment.

DISCUSSION

Germination behavior of fire-following herbs and suffrutescents
can be categorized into species with no apparent dormancy (except
perhaps a cold ‘stratification’ requirement) or ones with varying
degrees of dormancy that can be overcome, under natural conditions,
only by fire-related stimuli such as heat shock or charred wood.

Species with non-dormant seeds. Chaparral species with non-dor-
mant seeds include all herbaceous perennial monocot species, both
bulb-forming geophytes and bunch grasses, and many herbaceous
perennial dicots, such as Delphinium spp., Lomatium spp., Marah
macrocarpus, Paeonia californica, Perezia microcephala, and
Scrophularia californica (Table 1; also Sweeney 1956, Everett 1957,
Emery 1964, Keeley et al. 1985). The presence of these species on
recently burned sites is the result of resprouting from underground
vegetative parts; seedlings are uncommon at this time. Unlike most
perennials that colonize burned sites via seedlings, these resprouting
herbs flower vigorously during the first growing season after fire. We
predict that the timing of seedling establishment is most likely in
subsequent years after fire and up until the time the area is dominated
by shrubs. These species survive in gaps in the shrub cover or under
the canopy as dormant bulbs that occasionally produce depauperate
growth, but seldom flower (Stone 1951, Stocking 1966, Christensen
and Muller 1975a).

Non-dormant seeds also are characteristic of some annual species
found commonly on burned sites. Some of these, such as Agoseris
heterophylla, Galium parisiense, Heterotheca grandiflora, Lactuca
serriola, and Microseris linearifolia (Table 1), are relatively weedy
and produce diaspores capable of distant dispersal. Their presence
on first-year burns can be accounted for by colonization from nearby
disturbed areas such as road-cuts or natural disturbances. Many of
these annuals have heat sensitive seeds and, thus, it is of interest
that several disperse seeds in the fall and winter, after the time of
most chaparral wildfires. Some of these species produce polymorphic
achenes with different germination responses (e.g., Agoseris hetero-
phylla, see Table 1, and Heterotheca grandiflora, see Flint and Palm-
blad 1978) that may promote colonization of burned sites.

Other less weedy, annual species also have non-dormant seeds.
Antirrhinum nuttallianum, Clarkia unguiculata, Cordylanthus fili-
folius, Madia gracilis, Pterostegia drymarioides (Table 1), and Fes-
tuca megalura (Keeley et al. 1985) are often abundant in gaps in the
mature canopy. The seeds of these species are dispersed during the

1987] KEELEY AND KEELEY: FIRE AND GERMINATION 247

summer dry season and do not germinate until the following winter
wet season. Their presence on burned sites may be from seeds in
the soil that were produced by ‘gap’ plants the previous season or
from seeds under the canopy that, due to allelopathic compounds
from the shrub overstory, were dormant prior to the fire.

Species with heat-stimulated germination. Lotus salsuginosus was
the only species in this study with a significant increase in germi-
nation following heat treatment (Table 1). Other studies have re-
ported heat-stimulated germination for annuals such as Apiastrum
angustifolium, Brassica nigra, and Camissonia hirtella, as well as
for suffrutescents such as Helianthemum scoparium and Lotus sco-
parius (McPherson and Muller 1969, Christensen and Muller 1975a,
Keeley et al. 1985), and some shrubs such as Ceanothus spp. (Quick
1935). These species commonly are described as being ‘hard-seeded’
due to the heavily sclerified seed coats and thick cuticle that hinders
imbibition (Atwater 1980). Heat melts or cracks the cuticle, com-
monly around the hilum or strophiole, and this is sufficient to allow
germination because artificial scarification of the seed coat will pro-
duce the same stimulatory effect as heating. High soil temperatures
may produce the same stimulus as a heat shock during fire and,
thus, germination may be stimulated in gaps in the mature canopy
as well as on disturbed sites.

Species with charred wood stimulated germination. Germination
stimulated by charred wood is a far more specific means of timing
seedling establishment to burned sites than is heat. Not surpris-
ingly, such species are strongly associated with burns; sometimes
they appear in abundance the first year after fire and then disappear
until the next fire (true ‘pyrophyte endemics’). Charred wood stim-
ulated germination is widespread in the Hydrophyllaceae. It was first
discovered in Emmenanthe penduliflora (Wicklow 1977) and later
in many species of Phacelia (Keeley et al. 1985) and the shrub
Eriodictyon crassifolium (Keeley 1987). In terms of environmental
cues that they are likely to encounter in the field, these species exhibit
a nearly complete dependence upon charred wood. Complete de-
pendence upon charred wood also is found in both annual and
perennial species of Papaveraceae (Table 1). Other families with
species having a significant level of germination stimulated by charred
wood include the Asteraceae, Boraginaceae, Brassicaceae, Cary-
ophyllaceae, Onagraceae, Polemoniaceae, Rubiaceae, and Scroph-
ulariaceae (Table 1). Some of these species, e.g., the annual Gilia
capitata (Table 1), may persist in gaps during fire-free periods and
produce seeds that are polymorphic in their germination response.
A fraction of the seeds germinate each year and a larger portion
remain dormant until after fire when germination is stimulated by
charred wood (see also Grant 1949).

Based on the taxonomic distribution of germination responses

248 MADRONO [Vol. 34

observed here, we suggest that the mechanism behind germination
stimulated by charred wood is different than that for germination
stimulated by heat. Species in these two groups differ in several
respects. Unlike seeds that are stimulated by heat shock, which have
a smooth thick cuticle impermeable to water, seeds stimulated by
charred wood have highly sculptured, reticulate seed coats that are
not cutinized heavily. Emmenanthe penduliflora, for example, pro-
duces dormant seeds that will imbibe water (Sweeney 1956). Thus,
it seems probable that the chemical leached from charred wood
(apparently an oligosaccharide, Keeley and Pizzorno 1986) acts on
some internal component, and affects gas permeability of mem-
branes or provides a chemical stimulus to embryo development.
The former hypothesis is supported by the fact that scarification will
produce the same effect as charred wood (Wicklow 1977). The latter
hypothesis is supported by the fact that artificial application of gib-
berellic acid can duplicate the charred wood stimulus. For example,
germination of Penstemon spectabilis and Romneya coulteri in-
creased from 1-61% and from 0-—40%, respectively, in the presence
of charred wood (Table 1) and Atwater (1980) reported increases
from 2—70% for P. spectabilis and from 0—42% for R. coulteri with
the addition of gibberellic acid.

Future research will focus on those fire-following species, e.g.,
Eucrypta chrysanthemifolia, Dicentra crysantha, D. ochroleuca (TYa-
ble 1), and Phacelia brachyloba (Keeley et al. 1985), that we have
been unable to germinate. All of these species are restricted largely
to postfire conditions. Atwater (1980) has found that germination
of D. crysantha can be accomplished with the addition of gibberellic
acid. In light of the fact that in other species gibberellic acid can
simulate the effect of charred wood, it is likely that under natural
conditions, germination of these species is also cued by charred
wood, but apparently in conjunction with some other unknown fac-
tor.

ACKNOWLEDGMENTS

This work was supported in part by NSF grant RII-8304946. We thank Susan
Conard and Richard Minnich for helpful comments on the manuscript.

LITERATURE CITED

ATWATER, B. R. 1980. Germination, dormancy and morphology of the seeds of
herbaceous ornamental plants. Seed Sci. Technol. 8:523-573.

BRANDEGEE, T. S. 1891. The vegetation of “‘burns”’. Zoe 2:118-122.

CHRISTENSEN, N. L. and C. H. MULLER. 1975a. Effects of fire on factors controlling
growth in Adenostoma chaparral. Ecol. Monogr. 45:29-55.

and . 1975b. Relative importance of factors controlling germination
and seedling survival in Adenostoma chaparral. Amer. Midl. Naturalist 93:
71-78.

EmMerRY, D. 1964. Seed propagation of native California plants. Lflts. Santa Barbara
Bot. Gard. 1:81-96.

1987] KEELEY AND KEELEY: FIRE AND GERMINATION 249

EVERETT, P. C. 1957. A summary of the culture of California plants at the Rancho
Santa Ana Botanic Garden 1927-1950. Rancho Santa Ana Bot. Gard., Clare-
mont, CA.

FLINT, S. D. and I. G. PALMBLAD. 1978. Germination dimorphism and develop-
mental flexibility in the ruderal weed Heterotheca grandiflora. Oecologia 36:
33-43.

GRANT, V. 1949. Seed germination in Gilia capitata and its relatives. Madrono 10:
87-93.

JONES, C. S. and W. H. SCHLESINGER. 1980. Emmenanthe penduliflora (Hydro-
phyllaceae): further consideration of germination response. Madrono 27:122-
125.

KAMINSKY, R. 1981. The microbial origin of the allelopathic potential of Adenos-
toma fasciculatum H. & A. Ecol. Monogr. 51:365-382.

KEELEY, J. E. 1984. Factors affecting germination of chaparral seeds. Bull. So. Calif.
Acad. Sci. 83:113-120.

. 1987. Role of fire in seed germination of woody taxa in California chaparral.

Ecology 68:434-443.

, B. A. Morton, A. PEDROSA, and P. TROTTER. 1985. Role of allelopathy,
heat and charred wood in the germination of chaparral herbs and suffrutescents.
J. Ecology 73:445-458.

KEELEY, S. C., J. E. KEELEY, S. M. HUTCHINSON, and A. W. JOHNSON. 1981. Postfire
succession of the herbaceous flora in southern California chaparral. Ecology 62:
1608-1621.

and M. Pizzorno. 1986. Charred wood stimulated germination of two fire-
following herbs of the California chaparral and the role of hemicellulose. Amer.
J. Bot. 73:1289-1297.

McPHERSON, J. K. and C. H. MULLER. 1969. Allelopathic effects of Adenostoma
fasciculatum, ‘“‘chamise,” in the California chaparral. Ecol. Monogr. 39:177-198.

Munz, P.A. 1974. A flora of southern California. Univ. California Press, Berkeley.

Quick, C. R. 1935. Notes on the germination of Ceanothus seeds. Madrono 3:135-
140.

STOCKING, S. K. 1966. Influences of fire and sodium-calcium borate on chaparral
vegetation. Madrono 18:193-203.

STONE, E. C. 1951. The stimulative effect of fire on the flowering of the golden
brodiaea (Brodiaea ixiodes Wats. var. lugens Jeps.). Ecology 32:534—537.

SWEENEY, J. R. 1956. Responses of vegetation to fire. A study of the herbaceous
vegetation following chaparral fires. Univ. California Publ. Bot. 28:143-216.

WickLow, D. T. 1977. Germination response in Emmenanthe penduliflora (Hy-
drophyllaceae). Ecology 58:201—205.

(Received 8 Feb 1986; revision accepted 22 Dec 1986.)

ANNOUNCEMENT

NEw PUBLICATION

PETERSON, P. M., A flora of the Cottonwood Mountains, Death Valley Na-
tional Monument, California, Wasmann J. Biol. 44:73-126, 1986. [On 60
fam., 252 gen., 543 taxa of vascular plants.]

SOME DEMOGRAPHIC AND ALLOMETRIC
CHARACTERISTICS OF ACACIA SMALLIT
(MIMOSACEAE) IN SUCCESSIONAL COMMUNITIES

J. K. BUSH and O. W. VAN AUKEN
Division of Life Sciences,
The University of Texas at San Antonio,
San Antonio 78285

ABSTRACT

Diameter distributions of Acacia smallii Isley were examined in the South Texas
Plains Region in a series of successional communities ranging in age from 15 yr to
mature stands >150 yr. In the 15 yr stand, all individuals were in the sapling stage
and the size class distribution was non-normal and positively skewed. Mid-succes-
sional and older woodland communities had distributions that approached normal
curves. Mean live basal area of A. smailii increased in stands up to 25-29 yr, and
then declined. There was no recruitment of A. smallii in stands 19 yr or older. The
mean number of stems per plant increased and then decreased in the progression
from younger to older stands. Dead stems increased from zero in the 15 yr stand to
66% of the A. smallii density in the 33 yr stands. Of the total A. smallii basal area,
ca. 70% was dead tissue in the 33 yr stands. No individuals of A. smallii were present
in the mature forest community. Possible cause of this lack of A. smallii recruitment
was the low light environment caused by canopy closure. Demographic analyses of
A, smallii diameter distributions suggest it is an early successional species.

Acacia smallii Isley (huisache) is found throughout the southern
United States from northern Florida to California (Correll and John-
ston 1970). In south Texas, it is reported on approximately 1.1
million ha with more than 20% cover occurring in some places
(Smith and Rechenthin 1964). Acacia smallii is a heliophyte (Bush
and Van Auken 1986a), and is tolerant of low levels of soil nitrogen
(Van Auken et al. 1985). Acacia smallii has been reported as an
early successional species in south Texas (Van Auken and Bush
1985); however, there are no reports of demographic or allometric
characteristics of this species.

Studies of demographic characteristics of long-lived woody plants
in natural plant populations are scarce (Harper 1977). Whittaker
(1975) suggested that early successional species, those that would
not remain in a mature forest community, developed normal dis-
tributions, whereas the mature forest species had negative exponen-
tial distributions. Mohler et al. (1978) proposed idealized frequency
distributions of trunk diameters for selected stages in succession.
During stand establishment, the frequency distribution is a nega-
tively skewed function. After establishment, but before the start of
thinning, the distribution approaches a normal curve; at the start of

MADRONO, Vol. 34, No. 3, pp. 250-259, 1987

1987] BUSH AND VAN AUKEN: ACACIA SMALLIT 251

Fic. 1. Habitats of Acacia smallii during several stages of secondary succession.
A. Multi-stemmed growth form of A. smallii. B. Early open savanna or woodland
stage with a high grass density in the understory. C. Canopy closure during the late
savanna stage. D. Early mature forest stage with most A. smallii dead, and the presence
of other deciduous species in the overstory.

thinning the distribution is positively skewed. Finally, during late
thinning the curve again approaches a normal distribution. Other
studies, although less comprehensive, showed that disturbance se-
quence, site or niche difference, and initial density or spacing were
all factors that had a role in determining the demographics of a
species (Koyama and Kira 1956, Davidson and Donald 1958, Yoda
etal. 1963, Leak 1975, Crisp and Lange 1976, Crisp 1978, Harcombe
and Marks 1978, Ross et al. 1982, Knowles and Grant 1983, West
and Borough 1983).

In addition to changes in the demographics of a plant population,
increased competition causes alterations in the morphology of in-
dividual plants (Harper 1977). Morphological changes occur in whole
plants, for example, as shown in the annual diameter growth of Picea
sitchensis (Bong.) Carr. at various spacings (Jack 1971). Additional
changes in whole plant morphology were demonstrated in Pseudo-
tsuga menziesii (Mirb.) Franco, which showed a larger number of
suppressed individuals at higher densities (Curtis and Reukema 1970).
Jack (1971) demonstrated a reduction in the number of branches in
Picea sitchensis grown at high densities.

The purposes of this study were to examine some demographics
of population development and decline of Acacia smallii, and to

[Vol. 34

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MADRONO

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NIN WN =

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AGE (years)

1987] BUSH AND VAN AUKEN: ACACIA SMALLII 253

examine some allometric changes of this species associated with
temporal change.

METHODS

This study was conducted in the northern portion of the South
Texas Plains Region of Texas (Gould 1969). Fifteen communities
in various stages of secondary succession were selected for detailed
analyses. The sites were located on flood plain terraces of the San
Antonio River, which potentially support riparian woodland or for-
est communities (for locations see Van Auken 1982, Bush and Van
Auken 1984). Climate of the upper San Antonio River area is dry
subhumid (Thornthwaite 1948). This area receives approximately
71 cm/yr rainfall, with a mean annual temperature of 15.5°C (Carr
1967, Arbingast et al. 1976). Soils are deep loams (240-310 cm),
that usually are well drained, calcareous, and friable (Taylor et al.
1966, Taylor 1977).

Candidate stands were examined during a series of field surveys.
Stands also were located on aerial photographs and soil survey maps,
and local residents were interviewed concerning past history of the
sites. Selection was based on stand area, lack of additional distur-
bance other than light grazing in the mature communities, uniform-
ity of physical features, and vegetation. Selected stands depicted
various Stages in a chronological sequence from open fields to mature
forests.

The stands selected for study were 1-5 ha in size, and the area
sampled in each stand ranged from 0.1—0.5 ha. Most of the stands
were cleared in the 1950’s. The mature stands were estimated to be
in excess of 150 yr, as based on tree ring analyses.

Stands were sampled using belt transects, with 5m x 5 m quadrats
(Greig-Smith 1983). Density and basal area for all species except A.
smallii were pooled. All stems greater than 1.0 cm in diameter at
breast height were used to construct diameter distribution plots.

Demographic characteristics for the various stands were analyzed
by size class (diameter). The width of the size classes in each stand
was determined by dividing the range of diameters into 12 equal
size classes (Mohler et al. 1978). Statistical tests included Chi-square
analysis to test for normal distributions and a “‘t’’ statistic to test
for skewness (g,, asymmetry) and kurtosis (g,, peakedness) (Sokal

—

Fic. 2. Live (@ and dead (C) standing stem density (A) and basal area (B) for
Acacia smallii and all other woody species combined (Z) for a series of successional
stands. Asymmetry (skewness = O) and peakedness (kurtosis = @) of the 4. smallii
populations also are presented (C) with the number (N) of stands sampled (D). M =
mature stands greater than 150 yr old.

254 MADRONO [Vol. 34

FREQUENCY (%)

O 5 10 15 20 25 30 35 40
DIAMETER (cm)

Fic. 3. Frequency distribution of live stems of Acacia smallii in secondary succes-
sion including a 15 yr stand (A), a 27 yr stand (B), and a 33 yr stand (C). n = the
total number of plants measured in each stand, the distributions are not normal (Chi-
squared, p < 0.05).

and Rohlf 1981). An example of a 15, 27, and 33 yr stand is pre-
sented. Several morphological characteristics of A. smallii were ex-
amined including the number of stems per plant and the number
and percent of dead stems per plant for each stand.

For determination of stand age, 52 plants of A. smallii (of various
sizes and from various stands) were cored using the increment coring

1987] BUSH AND VAN AUKEN: ACACIA SMALLIT Z55

technique (Fritts 1976). Regression analysis (Steel and Torrie 1980)
was performed between tree diameter and tree age using all 52 sam-
ples (r = 0.88, p = 0.001). Next, the diameters of the five largest
trees in each stand were used to estimate the stand age. Stand age
was determined by adding 5 yr to the mean tree age (from regression)
to account for seed recruitment and an additional 3 yr to account
for growth from ground level to breast height (measured differences).

RESULTS

Some typical flood plain terrace habitats that include 4. smallii
are shown in Fig. 1. Secondary succession begins with a community
disturbance. Within 5 yr, A. smallii and other woody species colonize
the area (Fig. 2A). Acacia smallii increased in density and basal area
for the next 25 yr and dominated the stands (Fig. 2A,B). Dead A.
smallii first appeared in 25 yr stands. Of the total standing basal
area in 25 yr stands, ca. 16% was dead A. smallii. This value increased
to ca. 39% of the total standing basal area in the 33 yr stand. In the
33 yr stand, 74% of the A. smallii basal area was dead tissue. Acacia
smallii density followed a similar trend. Species composition shifted
and stands greater than 30 yr had a high density and basal area of
Celtis laevigata Willd. Acacia smallii was not present in the mature
stands, which were dominated by C. /aevigata, Carya illinoinensis
(Wang.) K. Koch. and Ul/mus crassifolia Nutt. (see Bush and Van
Auken 1984, Van Auken and Bush 1985).

None of the stands examined had statistically normal distributions
for A. smallii (Chi-square analyses p <= 0.05). Skewness decreased
from the highest value (1.08) in the 15 yr stand and approached zero
in older stands (Fig. 2C). Kurtosis also was highest (1.90) in the 15
yr stand, but decreased to ca. —1 in the 25-29 yr stands and ap-
proached zero in the oldest stands.

The diameter distribution for the 15 yr stand was a significant,
positively skewed function (g, = 1.08; t = 4.41, p < 0.001) (Fig.
3A). The distribution was also leptokurtotic (peaked) (g, = 1.90;
t = 3.88, p < 0.001). The largest individual in this stand was ca. 10
cm in diameter. The 27 yr stand was not skewed, but was platy-
kurtotic (flattened); however, neither value was statistically signifi-
cant (Fig. 3B). The largest individual was ca. 37 cm in diameter or
3.7 times larger than the largest tree in the 15 yr stand. In the 33 yr
stand, the distribution was slightly, positively skewed and was platy-
kurtotic (Fig. 3C). No individuals were found in the smallest size
classes. The largest individual was 26 cm in diameter. No 4. smallii
plants were found in the mature stands (>150 yr).

The number of stems per plant (both live and dead) for A. smallii
was time dependent (Fig. 4). In the 15 yr stand, number of stems
per plant ranged from 1-17 with no dead stems (Fig. 4A). In the 27

256 MADRONO [Vol. 34

FREQUENCY(%)

0 4 8 12 16 20 24 28
NUMBER OF STEMS/PLANT

Fic. 4. Frequency distribution of number of stems/plant in a 15 yr stand (A), a
27 yr stand (B), and a 33 yr stand (C). HM = live stems; LD = dead stems.

yr stand, number of stems per plant ranged from 2-18. There were
no single stemmed plants and those standing with 2 stems were
dead. In general, almost all other plants had 50% dead stems (Fig.
4B). In the 33 yr stand, number of stems was reduced to 1-8 per
plant, more than 50% of the stems per plant were dead, and many
of the larger trees were dead (Fig. 4C).

DISCUSSION

In many areas of southern Texas, Acacia smallii is a pioneer woody
species that colonizes abandoned farmland or rangeland. Although
it increases in density and basal area, community dominance of A.
smallii only lasts approximately 30 yr, at which time it is replaced
by mature community species (Van Auken and Bush 1985). De-
mographic characteristics examined showed the fate of A. smallii in

1987] BUSH AND VAN AUKEN: ACACIA SMALLII 25)

older stands. Density decreased and basal area increased, as has been
shown for other early successional forest species (e.g., Spurr and
Barnes 1973). Furthermore, a self-pruning of stems occurred; thus,
the tendency of growth form was toward single stemmed trees in
older communities. Finally, with overtopping by Celtis laevigata
and subsequent additional shading, the remaining trees died.

Acacia smallii seems to follow the pattern of frequency distribu-
tion that was proposed by Mohler et al. (1978) for Prunus pennsyl-
vanica L. In the present study, the 15 yr stand was well beyond their
‘establishment’ and “transition”’ stages, but was similar to their
“start of thinning’’ stage, which is a positively skewed function.
Phytosociological data showed an increase in the basal area of A.
smallii from the 15-27 yr stands, but a 46% decrease in density,
which supports the start of thinning hypothesis. Although none of
the frequency distributions in the present study were normal, the
distribution in the late successional community did approach a nor-
mal curve similar to the “late thinning”’ stage.

The differences between the present study and the frequency dis-
tributions of single species proposed by Mohler et al. (1978) are
probably due to several factors. In addition to intraspecific com-
petition that causes thinning in plant populations, interspecific com-
petition was probably also playing a critical role in shaping the
frequency distribution of A. small/ii. In the early stand, thinning was
most likely through the death of the smallest individuals; however,
in the mid- and late stands, death occurred to parts of whole plants
before the individuals died. Whereas in the early stands none of the
individuals had dead stems, the mid- and late stands showed in-
creased death to stems of individuals. In addition, in the mid- and
late stands, there was no recruitment of A. smallii.

Whittaker (1975) suggested that colonizing species would develop
bell-shaped frequency distributions, whereas mature community
species would have negative exponential distributions. This is prob-
ably an oversimplification, and dependent on episodic seedling es-
tablishment. Furthermore, his data suggested that, as the population
of a colonizer ages, the distribution remained normal, but frequency
of individuals in each size or age class would be reduced by deaths.

Changes in the available resources in a temporal sequence greatly
affect the competitive ability of the species involved and, therefore,
affect the frequency distributions. Acacia smallii is a heliophyte and
tolerates low levels of soil nitrogen (Van Auken et al. 1985, Bush
and Van Auken 1986a). Consequently, early in succession, A.
smallii is a better competitor than late community species, which
require higher levels of soil nitrogen (Van Auken et al. 1985, Bush
and Van Auken 1986a). In early stands, therefore, intraspecific com-
petition is probably more intense for A. smaillii than interspecific
competition; hence, smaller individuals often die. In older stands,

258 MADRONO [Vol. 34

light becomes limiting and soil nitrogen levels favor the late com-
munity species (Bush and Van Auken 1986a,b). These conditions
apparently do not allow recruitment of A. smallii (Van Auken and
Bush 1985), whereas interspecific competition affects the larger in-
dividuals through the death of stems and, finally, the death of the
trees. In the mature forest, A. sma/llii disappears and species such
as Celtis laevigata dominate because they are better competitors in
the canopy shade.

ACKNOWLEDGMENTS

This study was supported in part by the U.S. Department of the Interior, National
Park Service. We especially thank Ms. M. Bush Thurber and Mr. J. Cisneros of the
San Antonio Missions National Historical Park for their encouragement. We are
grateful to Mr. and Mrs. W. Southern for access to their land along the San Antonio
River and to Mr. D. Riskind of the Texas Parks and Wildlife Department for access
to undeveloped parkland in the same area. We also thank Dr. D. C. Randall and an
anonymous reviewer for many helpful comments and corrections.

LITERATURE CITED

ARBINGAST, S. A., L. G. KENNAMER, R. H. RYAN, J. R. BUCHANAN, W. L. HEZLEP,
L. T. Etuis, T. G. JORDAN, C. T. GRANGER, and C. P. ZLATKOVICH. 1976. Atlas
of Texas. Univ. Texas Bur. Bus. Res., Austin, TX.

Busu, J. K. and O. W. VAN AUKEN. 1984. Woody-species composition of the upper
San Antonio River gallery forest. Tex. J. Sci. 36:139-148.

and . 1986a. Light requirements of Acacia smallii and Celtis laevigata

in relation to secondary succession on floodplains of south Texas. Amer. Mid.

Nat. 115:118-122.

and 1986b. Changes in nitrogen, carbon and other soil properties
during secondary succession. Soil Sci. Soc. Amer. J. 50:1597-1601.

CARR, J. T. 1967. The climate and physiography of Texas. Texas Water Develop.
Board Reps., Austin, TX.

CorRRELL, D. S. and M. C. JOHNSTON. 1970. Manual of the vascular plants of Texas.
Tex. Res. Found., Renner, TX.

Crisp, M. D. 1978. Demography and survival under grazing of three Australian
semi-desert shrubs. Oikos 30:520-528.

and R. T. LANGE. 1976. Age structure, distribution and survival under
grazing of the arid-zone shrub Acacia burkittii. Oikos 27:86—92.

Curtis, R. O. and D. L. REUKEMA. 1970. Crown development and site estimates
in a Douglas fir plantation spacing test. For. Sci. 16:287-301.

Davipson, J. L. and C. M. DONALD. 1958. The growth of swards of subterranean
clover with particular reference to leaf area. Aust. J. Agri. Res. 9:53-72.

Fritts, H. C. 1976. Tree rings and climate. Academic Press, New York.

GOULD, F. W. 1969. Texas plants: a checklist and ecological summary. Texas Agri.
Exp. Sta. Bull. MP 505a.

GREIG-SMITH, P. 1983. Quantitative plant ecology. Butterworth, London.

HARCOMBE, P. A. and P. L. MARKS. 1978. Tree diameter distributions and replace-
ment processes in southeast Texas forests. For. Sci. 24:153-166.

HARPER, J. L. 1977. Population biology of plants. Academic Press, London.

JAcK, W. H. 1971. The influence of tree spacing on Sitka spruce growth. Irish
Forestry 28:13-33.

KNOWLES, P. and M. C. GRANT. 1983. Age and size structure analyses of Engelmann
spruce, ponderosa pine, lodgepole pine, and limber pine in Colorado. Ecology
64: 1-9.

KOYAMA, H. and T. Kira. 1956. Intraspecific competition among higher plants.

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VIII. Frequency distribution of individual plant weight as affected by the inter-
action between plants. J. Inst. Polytech. Osaka Cy. Univ. 7:73-94.

LEAK, W. B. 1975. Age distribution in virgin red spruce and northern hardwoods.
Ecology 56:1451-1454.

MOHLER, C. L., P. L. MARKS, and D. G. SPRUGEL. 1978. Stand structure and
allometry of trees during self-thinning of pure stands. J. Ecol. 66:599-614.
Ross, M. S., T. L. SHARIK, and D. W. SMitH. 1982. Age-structure relationships of
tree species in an Appalachian oak forest in southwest Virginia. Bull. Torrey Bot.

Club 109:287-298.

SMITH, H. N. and C. A. RECHENTHIN. 1964. Grassland restoration: the Texas brush
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SOKAL, R. R. and F. J. ROHLF. 1981. Biometry. W. H. Freeman, San Francisco, CA.

SpurR, S. H. and B. V. BARNES. 1973. Forest ecology. Ronald Press, New York.

STEEL, G. D. and J. H. Torrie. 1980. Principles and procedures of statistics: a
biometric approach. McGraw-Hill, New York.

TAYLOR, F. B. 1977. Soil survey of Wilson County, Texas. U.S. Dept. Agri., Soil
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, R. B. Harvey, and D. L. RICHMOND. 1966. Soil survey of Bexar County,
Texas. U.S. Dept. Agri., Soil Cons. Serv., Washington, D.C.

THORNTHWAITE, C. W. 1948. Anapproach toward a rational classification of climate.
Geogr. Rev. 38:55-94.

VAN AUKEN, O. W. 1982. Landscape study of the San Antonio Missions: botanical
aspects. Final report submitted to the U.S. Dept. Int., Nat. Park Ser., San Antonio
Missions Nat. Hist. Park.

and J. K. BusH. 1985. Secondary succession on terraces of the San Antonio

River. Bull. Torrey Bot. Club 112:158-166.

, E. M. Geese, and K. Connors. 1985. Fertilization response of early and late
successional species: Acacia smallii and Celtis laevigata. Bot. Gaz. 146:564—-569.

WEsT, P. W. and C. J. BoRouGH. 1983. Tree suppression and the self-thinning rule
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WHITTAKER, R. H. 1975. Communities and ecosystems. MacMillan, New York.

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(Received 7 Sep 1986; revision accepted 22 Apr 1987.)

ANNOUNCEMENT
NEw PUBLICATION

GRAYSON, A. J., Birds of the Pacific Slope, The Arion Press, 460 Bryant St.,
San Francisco, CA 94107, 1986, $45.00. [Includes a portfolio of 165 bird
portraits (full-scale facsimiles), reproducing all surviving paintings (California
and Mexico from 1853-1869) from the Bancroft Library (UC); a book with
preface by S. D. Ripley; and a biography of the artist and naturalist by Lois
C. Stone (long-time member of the California Botanical Society), with bird
biographies and field notes by Grayson and current ornithological descriptions
for each plate. Excellent plant and landscape paintings are associated with the
portraits. This work is considered by some authorities to be the most important
contribution to American Ornithology (i.e., illustrations and biographies) next
to the work of Audubon.]

MYCORRHIZAE ASSOCIATED WITH AN INVASION OF
ERECHTITES GLOMERATA (ASTERACEAE) ON
SAN MIGUEL ISLAND, CALIFORNIA

WILLIAM L. HALVORSON
U.S. National Park Service, Channel Islands National Park,
Ventura, CA 93001

RICHARD E. KOSKE
Department of Botany, University of Rhode Island,
Kingston 02881

ABSTRACT

Erechtites glomerata (Australian fireweed) is a perennial alien species that recently
has invaded San Miguel Island, an island off the coast of southern California. It is
presently advancing into a grassland dominated by Distichlis spicata and with scat-
tered shrubs. As is typical of many weedy species, E. glomerata is facultatively
mycotrophic on the island. Levels of colonization by vesicular-arbuscular mycorrhizal
(VAM) fungi ranged from 0-30% of the root system. Nine species of VAM fungi were
recovered from its root zone.

Australian fireweed, Erechtites glomerata (Poir.) DC. (Asteraceae),
is a native of Australia and New Zealand that has been advancing
slowly into the southern California region. It is a perennial that grows
to 2 m in height. The common name “‘fireweed”’ comes from Aus-
tralia and refers to the invasive nature of the plant in burn areas;
however, it also can easily invade cleared or otherwise disturbed
sites (Taylor 1964). On San Miguel Island, Santa Barbara Co., Cal-
ifornia, it has invaded and spread through a stable native grassland
community (Fig. 1).

Vesicular-arbuscular mycorrhizae (VAM) are intimate, mutualis-
tic associations formed between certain Zygomycetous fungi and
plant roots. The fungi apparently are obligate symbionts, obtaining
the bulk of their nutritional requirements from the “host” plant
(Harley and Smith 1983). The fungi occupy the cortical cells of roots
and produce hyphae that grow a few cm into the surrounding soil
where they absorb phosphate that is beyond the root’s depletion
zone. Numerous studies (e.g., Nelsen and Safir 1982, Harley and
Smith 1983, Fitter 1985) have shown that VAM ameliorate the
effects of water stress and reduced availability of phosphorus in the
soil. Growth improvements of up to 1100% have been achieved
when plants growing in phosphorus-deficient soils were inoculated
with VAM fungi (Mosse 1972).

VAM associations are extremely common, occurring in 95% of

MADRONO, Vol. 34, No. 3, pp. 260-268, 1987

1987] HALVORSON AND KOSKE: ERECHTITES GLOMERATA 261

Fic. 1. Erechtites glomerata in a grassland community on San Miguel Island,
California.

all plant families (Trappe in press). Although present in most soils,
VAM fungi sometimes are absent from certain sites. The presence
or absence of VAM fungi in soil can influence the ability of different
plant species to establish in an area (Janos 1981, Miller 1979, Reeves
et al. 1979). Plant species that require association with VAM fungi
to complete their life cycle (=obligate mycotrophs) are unable to
successfully invade and persist in sites that lack propagules of VAM
fungi. Such VAM-free sites, therefore, are preferentially colonized
by plant species that do not have an absolute requirement for VAM.
These plant species are classified as non-mycotrophs or facultative
mycotrophs, depending upon their ability to form VAM when the
appropriate fungi are present. Because they grow well whether or
not VAM fungi are present in the soil, facultative mycotrophs in-
clude many of the most troublesome weedy species (Trappe in press).

We examined plants of E. glomerata to determine their mycor-
rhizal status and to help explain their ability to be so invasive on
San Miguel Island, a natural area managed by the National Park
Service that functions under policy that calls for the removal of all
such invasive, noxious weeds.

STUDY AREA

San Miguel is a 4000 ha island off the coast of southern California.
It is the westernmost of the northern Channel Islands, occurring

262 MADRONO [Vol. 34

re)
i) 120 © Fireweed locations
Lompoc
@
°° ©
0)
© 9
iC) Santa Barbara
6
@ Ventura
Oxnard

Northern Channel Islands
a
34°

O 20
a |
kilometers

.~)

san Miguel Island

kilometers

Fic. 2. Location of San Miguel Island in the southern California Bight area,
showing the distribution (shaded area) of E rechtites glomerata (Australasian fireweed).

about 45 km south of Point Conception and 100 km west-southwest
of Ventura (Fig. 2). Bedrock on the island is composed primarily of
Cretaceous and early to mid-Tertiary conglomerates, sandstones,
siltstones, shales, and volcanics. Structurally, the island represents
the north flank of a folded and faulted anticline, whose axis trends
northwest-southeast (Johnson 1979, Weaver et al. 1969). Much of

1987] HALVORSON AND KOSKE: ERECHTITES GLOMERATA 263

the island is covered with sand, both stabilized and unstabilized. In
the area that is the subject of this report, the soils are of the vertisol
type with a high level of expandable clay and shrink-swell charac-
teristics (Johnson 1979).

The specific weather/climate characteristics of the island are rel-
atively undefined due to a lack of adequate data. The island lies in
the dry-summer, subtropical climate, commonly called Mediterra-
nean (Trewartha 1954). Rainfall is in the range of 330-355 mm per
year and the mean annual temperature is 13.7°C with an annual
range of 3°C. The two most characteristic features of the weather
are wind and fog. The wind is almost constant and comes principally
out of the northwest. The winds commonly blow 30—40 km/hr with
gusts up to 60 km/hr; strong northwest flows during the period of
winter storms, bring winds of 70-80 km/hr. Morning fog is common
throughout the year, but it is most constant during the summer
months (Dunkle 1950, Weissman and Rentz 1977, NPS files).

The most important plant community on the island is grassland.
There are two types: those dominated by introduced Avena (A. fatua
L. and A. barbata Brot.) and Bromus (B. mollis L., B. rubens L. and
B. diandrus Roth) and those dominated by the native Distichlis
spicata (L.) Greene. Other community types include scrub domi-
nated by Haplopappus venetus (HBK.) Blake, and coastal sage scrub,
coastal bluff, coastal dunes, and a small coastal salt marsh (Hochberg
et al. 1979).

METHODS

The island was surveyed carefully in January 1985, and the total
areal extent of the Erechtites population was mapped. One hundred
quadrats (1 x 1 m) were placed randomly throughout the invasion
area to assess the density of the population and to determine the
species composition of the grassland community.

Root and soil samples were collected in July and November 1985,
to determine the status of mycorrhizae. Roots of six specimens of
E. glomerata were fixed in the field in formalin: acetic acid : ethanol:
water (2:1:5:7, v/v/v/v). In the laboratory, roots were cleared and
Stained using a modification of the method of Phillips and Hayman
(1970). The fixed roots were cleared by autoclaving for 3 minutes
in 10% KOH. Cleared roots were rinsed in a dilute HCl solution,
and mycorrhizae were stained by autoclaving the roots for 3 minutes
in 0.05% trypan blue in lactic acid: glycerol: water (1:2:1, v/v/v).
Roots were destained by autoclaving for 3 minutes in the above
solution without trypan blue.

The extent of colonization of roots by VAM fungi was determined
by estimating the percent (to nearest 10%) of the length of the ab-
sorbing root system that contained arbuscules, vesicles, hyphal coils,
or internal hyphae of VAM fungi.

To determine the species of VAM fungi associated with the plants,

264 MADRONO [Vol. 34

TABLE 1. COMPOSITION OF THE GRASSLAND COMMUNITY INTO WHICH Frechtites
glomerata Is INVADING ON SAN MIGUEL ISLAND. Presence (%) was derived by dividing
the number of plots in which a species was found by the total number of plots sampled.
* = species that are considered alien to the San Miguel Island flora.

Species Presence (%)

Distichlis spicata (L.) Greene 94
*Medicago polymorpha L. 58
Amsinckia intermedia F. & M. 32
*Stellaria media (L.) Vill. 18
Malacothrix incana (Nutt.) T. & G. 16
*Galium aparine L. 14
*Sonchus oleraceus L. 14
Calystegia macrostegia (Greene) Brummitt 10
Lupinus succulentus Dougl. ex Koch. 8
Dichelostemma pulchellum Heller
Eschscholzia californica Cham.
Atriplex californica Mog. in DC.
* Atriplex semibaccata R. Br.
*Daucus pusillus Michx.
Astragalus curtipes Gray
Chenopodium californicum (Wats.) Wats.
*Erodium moschatum (L.) L’Her.

NNNAHRHHAD

soil samples (ca. 500 cc) were collected from the root zones of two
plants. A 75 cc subsample composed of 20-30 smaller subsamples
withdrawn from the 500 cc sample was processed to recover spores.
Spores were extracted from the soil by a water-sucrose centrifugation
technique (Walker et al. 1982). Following centrifugation, spores were
collected on a 5.5 cm filter paper (Whatman no. 1) in a Buchner
funnel. The filter paper was examined at 30 with a dissecting
microscope, and spores were removed, mounted in a polyvinyl al-
cohol mountant (Koske and Tessier 1983) and identified with the
aid of a compound microscope at 400-1000 x. Identifications were
confirmed by comparison with type or authenticated specimens and
by consultation with VAM taxonomists. Voucher specimens have
been deposited in the mycological herbarium at the University of
Rhode Island. Nomenclature of higher plants follows Munz (1968)
except for Erechtites glomerata, which follows Barkley (1981). No-
menclature for fungi follows original authors that are given in Ta-
ble 2.

RESULTS

The grassland that Erechtites glomerata is invading on San Miguel
Island (Table 1) is dominated by Distichlis spicata with scattered
patches of forbs, particularly Amsinckia intermedia, Eschscholzia
californica, Calystegia macrostegia, Chenopodium californicum,
Sanicula arguta, and Dichelostemma pulchellum. Scattered shrubs,

1987] HALVORSON AND KOSKE: ERECHTITES GLOMERATA 265

Fics. 3, 4. VAM fungi in roots of Erechtites glomerata. Stele is indicated (‘‘S’’),
scale bar is 50 wm. 3. Hyphae and hyphal coils. 4. Arbuscules and hyphae in cortical
cells.

including Baccharis pilularis subsp. consanguinea and Solanum
douglasii, also are present.

The flora of San Miguel Island was surveyed in 1978-79 (Hochberg
et al. 1979), and no plants of Australasian fireweed were found.
Plants of E. glomerata were first observed in May 1984, and the
species was well established at that time (Junak pers. comm.). In
January 1985, we determined that it covered an area of approxi-
mately 70 ha to the west of Green Mountain. Densities of stems
within the population showed a pattern of spread from north to
south in response to the prevailing winds. Density at the point of
origin was 8800/ha. This decreased to 2100/ha and finally 500/ha
with increasing distance southward.

TABLE 2. SPECIES OF VESICULAR-ARBUSCULAR MYCORRHIZAL (VAM) FunNGaI
ISOLATED FROM THE Root ZONE OF Erechtites glomerata.

Acaulospora laevis Gerd. & Trappe (Gerdemann and Trappe 1974)

Entrophospora infrequens Ames & Schneider (Ames and Schneider 1979)
Gigaspora calospora (Nicol. & Gerd.) Gerd. & Trappe (Gerdemann and Trappe 1974)
Glomus aggregatum Schenck & Smith (Schenck and Smith 1982)

Gl. intraradices Schenck & Smith (Schenck and Smith 1982)

Gl. pansihalos Berch & Koske (Berch and Koske 1986)

Gl. scintillans Rose & Trappe (Rose and Trappe 1980)

Gl. 598 (spores yellow-brown to red-brown, 70-140 wm diam., thick-walled)

Gl. 2163 (spores pale yellow, 60-120 um diam., thin-walled)

266 MADRONO [Vol. 34

Five of the six plants of Australasian fireweed sampled possessed
vesicular-arbuscular mycorrhizae (Figs. 3, 4), with levels of VAM
colonization ranging up to 30% (x = 14%). Nine species of VAM
fungi were isolated from the root zone of E. glomerata (Table 2).
Species that produced the most numerous spores in association with
this host were Glomus pansihalos and Gigaspora calospora. Two of
the species, Glomus 598 and G/. 2163, could not be assigned to
existing taxa and apparently are undescribed new species.

DISCUSSION

Erechtites glomerata is an aggressive alien that newly inhabits San
Miguel Island. Introduction apparently resulted from seeds being
carried across the Santa Barbara Channel from the mainland (Fig.
2). At this time, it is found nowhere else on the Channel Islands.
Although in its native Australia it is most common in burned or
disturbed areas, on San Miguel Island this fireweed is invading an
established native grassland. Its spread has been rapid and effective
with fireweed becoming a major component of the grassland com-
munity.

We have found eight of the nine species of VAM fungi recovered
in the present study occurring in association with native plant species
on San Miguel Island, a typical situation because VAM fungi usually
have wide host ranges (Harley and Smith 1983). Glomus scintillans,
the one species that has not been found thus far in the root zones
of other plants on the island, was described originally from a shrub
desert site in eastern Oregon (Rose and Trappe 1980), where it was
associated with shrubs that harbor nitrogen-fixing actinomycetes in
their roots.

Of the six other species that have been described previously, four
have been found in the southwestern U.S.: Entrophospora infrequens
in mainland Ventura Co., California (Ames and Schneider 1979,
Nemec et al. 1981); Acaulospora laevis, Entrophospora infrequens,
Gigaspora calospora, and Glomus intraradices from the Sonoran
Desert in Arizona (Bloss 1986); and Glomus intraradices from Anza
Borrego State Park in southern California (Bethlenfalvay et al. 1984).
The other two species (Glomus aggregatum and Glomus pansihalos)
have been found in the Great Lakes Region and on the east coast
of the United States (Koske 1987, Koske and Tews in press, P.
Olexia, pers. comm.).

Erechtites glomerata appears to be a facultative mycotroph that
fits within the pattern of tropical weedy species in which those plants
that produced light seeds were shown to be the least dependent upon
mycorrhizae (Janos 1980). This characteristic explains in part why
E. glomerata is such a successful invader, and should be considered
in any management strategy that the National Park Service might
develop for its control or removal.

1987] HALVORSON AND KOSKE: ERECHTITES GLOMERATA 267

ACKNOWLEDGMENTS

We thank Steve Junak for his assistance in determining the location of E. glomerata,
for help in the field, and in reviewing this manuscript, and Chris Walker and Joe
Morton for their assistance in identifying the fungi. Field assistance was provided by
Ronie Clark and Frank Ugolini. The study was supported by the National Park Service
Science Program.

LITERATURE CITED

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BARKLEY, T. M. 1981. Senecio and Erechtites (Compositae) in the North American
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Bercy, S. M. and R. E. Koske. 1986. Glomus pansihalos, a new species in the
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BETHLENFALVAY, G. J., S. DAKESSIAN, and R. S. PACovsky. 1984. Mycorrhizae in
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Boss, H. E. 1986. Symbiotic microflora and their role in the ecology of desert
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DUNKLE, M. B. 1950. Plant ecology of the Channel Islands of California. Allen
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Fitter, A. H. 1985. Functioning of vesicular-arbuscular mycorrhizae under field
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GERDEMANN, J. W. and J. M. TRAPPE. 1974. The Endogonaceae in the Pacific
Northwest. Mycol. Mem. 5:1-76.

HARLEY, J. L. and S. E. SmitH. 1983. Mycorrhizal symbiosis. Academic Press,
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HOcHBERG, M., S. JUNAK, R. PHILBRICK, and S. TIMBROOK. 1979. Botany. Jn D.
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JANOS, D. P. 1980. Vesicular-arbuscular mycorrhizae affect lowland topical rain
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JOHNSON, D. L. 1979. Geology, soils, and erosion. Jn D. M. Power, ed., Natural
resources study of the Channel Islands National Monument, California, p. 3.1-
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KosKE, R. E. 1987. Distribution of VA mycorrhizal fungi along a latitudinal tem-
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and B. Tessier. 1983. A convenient, permanent slide mounting medium.

Mycol. Soc. Amer. Newsl. 34(2):59.

and L. L. Tews. In press. Vesicular-arbuscular mycorrhizae of Wisconsin’s
sand soils. Trans. Brit. Mycol. Soc.

MILLER, R. M. 1979. Some occurrences of vesicular-arbuscular mycorrhizae in
natural and disturbed ecosystems of the Red Desert. Canad. J. Bot. 57:619-623.

Mosse, B. 1972. Effects of different Endogone strains on growth of Paspalum no-
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Munz, P. A. 1968. A California flora and supplement. Univ. California Press,
Berkeley.

NELSEN, C. E. and G. R. SAFiR. 1982. Increased drought tolerance of mycorrhizal
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NEMEC, S., J. A. MENGE, R. G. PLATT, and E. L. V. JOHNSON. 1981. Vesicular-
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268 MADRONO [Vol. 34

REEVES, F. B., D. WAGNER, T. MOORMAN, and J. KeIL. 1979. The role of endo-
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WALKER, C., C. M. Mize, and H. S. McNAsB. 1982. Populations of edogonaceous
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(Received 5 May 1986; revision accepted 9 Mar 1987.)

NOTEWORTHY COLLECTIONS

BRITISH COLUMBIA

SALIX TWEEDYI (Bebb) C. R. Ball (SALICACEAE).— Bolean Lake, northeast of Falk-
land. 50°32'N, 119°30’W, 1440 m, in a Salix, Carex swamp at s. end of lake in front
of resort, associated with S. barclayi, 25 Jul 1986, 7. C. Brayshaw 86-23, -24, -26,
-27, -28, -29, -30 (CAN, V).

Previous knowledge. This species was first collected in Canada at this locality in
1941 by C. L. Hitchcock and J. S. Martin. Their collection number 7524 was dis-
tributed as S. barclayi Anderss. In 1962, A. Cronquist recognized that a duplicate at
NY was actually S. tweedyi.This specimen evidently was the basis for the inclusion
of BC in the distribution of the species in Hitchcock et al. (Vascular Plants of the
Pacific Northwest 2:69, 1964). Specimens of 7524 in RM and WTU alsoare S. tweedyi,
but the specimen in A was correctly named S. barclayi.

Significance. These collections confirm the occurrence of S. tweedyi in Canada at
a locality about 200 km n. of its nearest locality in Washington (Okanogan Co.,
Tiffany Mt.). This species is rare in BC.—GEORGE W. ArGus, National Herbarium,
Museum of Natural Sciences, Ottawa, ON K1A 0M8 and T. C. BRAyYsHAW, British
Columbia Provincial Museum, Victoria, BC V8V 1X4, Canada.

NEw MExIco

SALIX GEYERIANA Anderss. (SALICACEAE). — Catron Co., Mogollon Mountains, Gilita
Cr. at confluence of Indian Cr., ca. 31 km e. of Mogollon, 33°24'N, 108°34’W, 8000

MaproNo, Vol. 34, No. 3, pp. 268-269, 1987

1987] NOTEWORTHY COLLECTIONS 269

ft, dominant in Salix thicket along creek, 27 Jun 1986, G. W. and J. N. Argus 12258,
12263 (CAN). Luna, on US 180, 3.4 km w. of town at crossing of San Francisco
River, 33°50'N, 109°01’W, 7500 ft, Populus angustifolia thicket on creek margin, 2
Jul 1986, G. W. and J. N. Argus 12394 (CAN). Luna, on US 180, 2 km w. of town,
33°50'N, 108°59'W, 7100 ft, Salix irrorata dominated thicket in wet meadow, 2 Jul
1986, G. W. and J. N. Argus 12398 (CAN).

Previous knowledge. Occurs in the Rocky Mountains from southern British Co-
lumbia to Colorado and in California with disjunct localities in western Nebraska
and the White Mountains of Arizona.

Significance. New to the flora of New Mexico. This occurrence in the Mogollon
Mts. parallels the disjunction in the White Mts. of Arizona.—GEORGE W. ARGUS,
National Herbarium, Museum of Natural Sciences, Ottawa, ON K1A OM8, Canada.

FESTUCA MINUTIFLORA Rydb. (POACEAE).— Rio Arriba Co., Pecos Wilderness Area,
North Truchas Peak, w. slope of mountain, 35°59'N, 105°37’W, 12,000 ft, alpine
vegetation on talus slope, 4 Jul 1986, G. W. and J. N. Argus 12404 (CAN) (identified
by Susan Aiken).

Previous knowledge. Scattered throughout the w. states (AZ, CA, CO, OR, UT,
WY) at elevations between 3000-4000 m. It is relatively common in Colorado, but
poorly known elsewhere (Frederiksen, Bot. Notiser 132:315-318, 1979).

Significance. New to the flora of New Mexico.— GEORGE W. ARGUS and SUSAN G.
AIKEN, National Herbarium, Museum of Natural Sciences, Ottawa, ON K1A 0OM8,
Canada.

REVIEWS

Xantus, The Letters of John Xdntus to Spencer Fullerton Baird from San Francisco
and Cabo San Lucas, 1854-1861. Introduction, Notes and Illustrations by ANN H.
ZWINGER. 442 pp. Dawson’s Book Shop, Los Angeles. 1986. $69.00.

Any biologist concerned with natural history in Baja California, Mexico, is familiar
with the specific epithets xanti or xantusii. John Xantus de Vesey sailed from San
Francisco in March 1859 during our spring and arrived to Cabo San Lucas in early
April, at the height of the dry season there. It is small wonder that in his first letter
he said, “‘There is not a drop of water for a distance of 28 miles (San Jose) only Mr.
Ritchie has a well, of very indifferent brackish water, and there is not a tree for many
miles, if we except the Cactuses, of which there is infinite variety... .”” Xantus installed
a tidal gauge, which was the reason for the U.S. Coastal Survey having sent him to
the tip of Baja California, and began to collect natural history specimens for the
Smithsonian Institution. This was a field of endeavor in which he excelled and one
that he much preferred to that of recording tidal data.

These letters to Mr. Baird, the newly appointed Assistant Secretary to Smithsonian
Institution in Washington, DC, show the difficulties under which Xantus carried on
his work. He had to take all scientific equipment with him; mail sometimes took six
months or more to reach him. Shipment of his scientific specimens depended upon
unscheduled arrival of whalers or ships that were bound for San Francisco or eastern
seaboard ports. His letters contain meticulous reports on the contents of each shipment

MADRONO, Vol. 34, No. 3, pp. 269-271, 1987

270 MADRONO [Vol. 34

and of collecting conditions, but the most that they say of the people in the little
pueblo of Cabo San Lucas is that boys sometimes brought him specimens. That there
were people there is brought out by Professor Emeritus Herbert Mason’s story about
his brief time ashore at Cabo San Lucas in 1925, when the California Academy of
Sciences’ expedition stopped there during its return trip from the Revillagigedos
Islands. An elderly paisano who was watching while specimens were being put into
a plant press remarked, “Mi papa tenia uno de estos.”’ ““‘Who was your papa?”
‘*Xantus.” The man couldn’t have been much more than a baby at the time Xantus
went from Cabo San Lucas on very short notice; so Xantus must have left more than
collecting equipment behind! In 1940 when Steinbeck and Ricketts touched at Cabo
San Lucas (cf. Zwinger footnote, p. 324) the manager of the cannery, pointing to three
little Indian children said, ‘““Those are Xanthuses great-grandchildren,” and “in the
town there is a large family of Xanthuses.”’

The paucity of detail about life of the people is undoubtedly due to the fact that
Xantus’ letters to Baird were business letters detailing progress of his work and
difficulties encountered. In one letter, however, Xantus includes a list of 14 donors
of scientific material. Typical of these is item No. 8, ““Donnas Juana & Pachita Dodero,
10 nests, with 34 eggs, & several bottles of insects.’ In contrast to his usual letters
is that of 28 December 1860 in which he says, ‘““The Christmas day I spent in San
Jose, amongst bullfights, cockfights, & dancing. There was a great concurse [sic] of
people. .. . The whole fiasta [sic] went off however very decently & with great order,
more so than a 4th of July in a small American village.” His letters to his mother in
Hungary were replete with exciting accounts of his expeditions and adventures—
many of them undoubtedly more fancy than fact.

Ann Zwinger’s introductory chapter (36 pp.) provides a biographical background
for Xantus and brings out the important role that Spencer Fullerton Baird played in
building up natural history collections at Smithsonian. Zwinger’s copious footnotes
to the letters contribute important historical data as to the identity of people men-
tioned in the letters as well as clarification of some of the scientific names that Xantus
cited in his lists. These footnotes not only add to the interest of the book, but also
make it historically valuable. An extensive bibliography of the works cited and an
unusually full index add to the usefulness of this volume. Because of her long interest
in and association with the Cape Region of Baja California, as evidenced by her book
A Desert Country Near the Sea, Ann Zwinger is especially well-fitted to present this
treatment of Xantus.

The Castle Press is to be complimented on a good job of printing the difficult
material. This is a worthy addition to Glen Dawson’s series on Baja California. —
ANNETTA CARTER, Dept. of Botany, Univ. of California, Berkeley 94720.

Flora Fanerogamica del Valle de Mexico. Volumen II. Dicotyledonae (Euphor-
biaceae—Compositae). Edited by JERZY RZEDOWSKI and GRACIELA C. DE RZEDOWSKI.
Instituto de Ecologia, Apartado Postal 18-845, Delegacion Miguel Hidalgo, 11800
Mexico, D.F., Mexico. ISBN 968-7213-02-7. 1985. 674 pp. $35? (cloth).

The valley of Mexico is thought of as an area full of people (one of the world’s
largest metropolitan regions) and, therefore, quite denuded of vegetation. In reality,
there is a lot of plant life in the region. The Rzedowskis are in the process of producing
an excellent three volume flora of this valley: Vol. I (1979) Gymnosperms and dicots
up to Polygalaceae; Vol. II (1985), the remainder of the dicots, Euphorbiaceae through
Compositae (here reviewed with about 1040 species treated); Vol. III, to be published,
monocots.

The format is clear and very easy to use with the families arranged in an order that
seems to be of the editors’ design with similarities to some modern systematic treat-

1987] REVIEWS oi

ments. This volume has a hard cover and a small but clear type face, which are
distinct improvements over Vol. I that was printed by a different publisher.

The families, genera, and species are described fully with economic and distribu-
tional notes at the end of the family and generic descriptions. The species are presented
alphabetically within the families except for the Compositae, which are alphabetical
within the tribes. The species descriptions are concise with many unique observations,
and are followed by the plant’s range within the area of the flora with localities and
then in general terms for its total range. A statement of the habitat and associations
where the plant occurs in the valley also is given.

The keys are indented and easy to use, although some couplets in the keys to large
genera are very involved, use many characters, and are a little confusing. There are
good drawings with habit and details of some species in all families. One would like
more, but cost and space are a valid concern.

The boundaries for this flora include the slopes of all of the various sierras that
form the Valley of Mexico and range up to 5452 m. A very nice biproduct of this
fact is that this flora has a utility for a much wider range, as many of the plants occur
at high altitudes from Durango to Oaxaca.

Throughout the work, references are cited where they are drawn upon in the prep-
aration of the treatments. This is reflected in the conservative and usually current
species concepts presented. Although there are 46 authors of various groups in this
volume alone, the presentation is uniform and reflects considerable effort by the
editors to accomplish this.

The Rzedowskis are to be congratulated on the completion of this volume of the
flora of the Valley of Mexico, which is a valuable addition to our knowledge of the
plants of Mexico and will be a useful tool for many years to come.— DENNIS E.
BREEDLOVE, Dept. of Botany, California Academy of Sciences, Golden Gate Park,
San Francisco, CA 94118.

A Flora of Dry Lakes Ridge, Ventura County, California. By David L. Magney, vii
+ 110 pp. The Herbarium, Dept. of Biological Sciences, Univ. of California, Santa
Barbara, Publication No. 5, 1986. $8.00

This botanical research results from a four year study during every month of the
year. Data include details regarding soil analyses, geology, climatology, land use,
floristic history, botanical resources of special concern, environmental sensitivity,
and recommendations for management procedures.

The area is north of Ojai at the headwaters of the North Fork of Matilija Creek.
The ridge’s north flank is drained by Sespe Creek. Access is available by foot, with
some difficulty, from two directions via bulldozed firebreak/trails.

The effects on the vegetation by fires during 1932, 1948, and 1985 are explained.
There are two habitat groups: 1) wetlands—consisting of stream, seeps and slopes;
2) upland—basins along the ridge at the summit with finer-grained soils.

Each principal plant species is discussed as to percentage of cover and how it
dominates or persists in localized areas. Post-fire vegetation is listed. Erosion control
plantings are evaluated. There are five plants endemic to the general region. Paleo-
botanical aspects are dealt with. Fossil evidence and woodrat middens are discussed.
The book concludes with an annotated catalogue of vascular plants.

There are several black-and-white illustrations of good quality. Graphs, charts, line
drawings, and maps are uncomplicated and quite understandable. The text is double-
spaced and easily readable. The work is well-prepared and deals with a little-known
botanical area.-WALTER KNIGHT, Research Associate, The Carnegie Museum, Section
of Botany, Pittsburgh, PA.

CALIFORNIA BOTANICAL SOCIETY

SCHEDULE OF SPEAKERS
1987-1988
8:00 pm University of California, Berkeley LSB 2503

SPEAKER & TOPIC

V. T. Parker, San Francisco State Univ.
“Seed banks of California chaparral’’

Niall McCarten, San Francisco State Univ.

“Taxonomy of the genus Hesperolinon”

Eloy Rodriquez, Univ. California, Irvine
‘‘Pharmacopeia of wild champanzees”’

P. L. Fiedler, San Francisco State Univ.
“Hierarchical thinking in rare plant biology”

TO BE ANNOUNCED

Connie Millar, Univ. California, Berkeley
‘The role of genetics in conservation biology”

Daryl Koutnik, Huntington Library and Botanical Garden
‘The genus Euphorbia in Southern Africa”

Richard D. Laven, Colorado State University
‘‘Subalpine fir succession in the Colorado Rockies”

* Annual Banquet—location and speaker to be announced.

Volume 34, Number 3, pages 173-272, published 30 September 1987

MADRONO, Vol. 34, No. 3, p. 272, 1987

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CALIFORNIA BOTANICAL SOCIETY

OK
VOLUME 34, NUMBER 4 } OCTOBER-DECEMBER 1987
072
| MiSF
| Bot

MADRON

A WEST AMERICAN JOURNAL OF BOTANY

Contents

FRUIT PRODUCTION PATTERNS IN THE CHAPARRAL SHRUB ;Ceanothus crassifolius
Jon E. Keeley 278
SEED DISPERSAL IN Ceanothus cuneatus AND C. leucodermis IN A SIERRAN; : 1 re)
OAK-WOODLAND SAVANNA A NE enol?
Raymond A. Evans, Harold H. Biswell, and Debra E. Palmaut nanee 283
SEEDCROP CHARACTERISTICS AND MINIMUM REPRODUCTIVE SIZE OF ORGAN PIPE
Cactus (Stenocereus thurberi) IN SOUTHERN ARIZONA

Kathleen C. Parker 294
REPRODUCTIVE BIOLOGY OF THE TREE /pomoea wolcottiana (CONVOLVULACEAE)
Stephen H. Bullock, Ricardo Ayala, Irene Baker, and Herbert G. Baker 304
_ ALPINE ANNUAL PLANT SPECIES IN THE WHITE MOUNTAINS OF EASTERN CALIFORNIA
Timothy P. Spira 35

Prosopis (MIMOSACEAE) IN THE SAN JOAQUIN VALLEY, CALIFORNIA: VANISHING RELICT
OR RECENT INVADER?

Dan C. Holland 324
_ Some NEw AND RECONSIDERED CALIFORNIA Dudleya (CRASSULACEAE)
Kei M. Nakai 334
| Cymophora (ASTERACEAE: HELIANTHEAE) RETURNED TO Tridax
David J. Keil, Melissa A. Luckow, and Donald J. Pinkava 354
| VASCULAR PLANTS OF EASTERN IMPERIAL COUNTY, CALIFORNIA
_ Steven P. McLaughlin, Janice E. Bowers, and Kenneth R. F. Hall 359
_ NOTES
THE RANGE AND Two NEw LocatTIons OF Boschniakia strobilacea (OROBANCHACEAE)
| David L. Magney 379
_ NOTEWORTHY COLLECTIONS
CALIFORNIA 381
ECUADOR 381
NEVADA 382
“REVIEW 382
~-ANNOUNCEMENTS 293, 333, 353, 383, 392
~NEW MADRONO EDITOR 358
LETTERS AND COMMENTARY
LETTER 384
MESSAGE FROM THE PAst CBS PRESIDENT 384
EDITOR’S REPORT FOR VOLUME 34 385

(continued on back cover)

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
ciety, Inc., and is issued from the office of the Society, Herbarium, Life Sciences
Building, University of California, Berkeley, CA 94720. Subscription rate: $25 per
calendar year. Subscription information on inside back cover. Established 1916.
Second-class postage paid at Berkeley, CA, and additional mailing offices. Return
requested. PosTMASTER: Send address changes to James R. Shevock, Botany Dept.,
California Academy of Sciences, San Francisco, CA 94118.

Editor— WAYNE R. FERREN JR.
Associate Editor—BARRY D. TANOWITZ

Department of Biological Sciences
University of California
Santa Barbara, CA 93106

Board of Editors
Class of:

1987—J. RZEDOwSKI, Instituto de Ecologia, A.C., Mexico
DoroTHY DOUGLAS, Boise State University, Boise, ID
1988—SusANn G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989—FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991—Davip J. Kem, California Polytechnic State University, San Luis Obispo
JAMES HENRICKSON, California State University, Los Angeles

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1987-88

President: DALE MCNEAL, Department of Biological Sciences, University of the
Pacific, Stockton, CA 95211

First Vice President: PEGGY FIEDLER, Department of Biology, San Francisco State
University, 1600 Holloway Ave., San Francisco, CA 94132

Second Vice President: STEVEN TIMBROOK, Ganna Walska Lotusland Foundation,
695 Ashley Rd., Montecito, CA 93108

Recording Secretary: V. THOMAS PARKER, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
Francisco, 50 Phelan Ave., San Francisco, CA 94112

The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, FRANK ALMEDA, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118; the Editor of MADRONO; three elected
Council Members: ANNETTA CARTER, Department of Botany, University of Califor-
nia, Berkeley, CA 94720; JOHN MoorinGc, Department of Biology, University of
Santa Clara, Santa Clara, CA 95053; BARBARA ERTTER, University Herbarium, De-
partment of Botany, University of California, Berkeley, CA 94720; and a Graduate
Student Representative, NIALL F. MCCARTEN, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132.

FRUIT PRODUCTION PATTERNS IN THE
CHAPARRAL SHRUB CEANOTHUS CRASSIFOLIUS

JON E. KEELEY
Department of Biology, Occidental College,
Los Angeles, CA 90041

ABSTRACT

Annual fruit production patterns in Ceanothus crassifolius varied significantly over
a period of six years and significant differences occurred between sites. Plants on
southern exposures out-produced similar aged shrubs on northern exposures. Whether
or not the causative factors are associated with aspect or differences in density of
vegetative cover are unknown. Significant differences occurred among shrubs on the
same slope. Some plants produced, on average, nearly an order of magnitude more
fruits (per unit of areal coverage) than other plants. Fruit production patterns in
Ceanothus species, particularly members of subg. Cerastes, are, in part, a function of
conditions in the year prior to flowering and fruiting because flower buds are produced
nearly 12 months prior to flowering. In years of heavy fruit production, internal
competition for resources may limit flower bud production for the following year.
This pattern is particularly likely if heavy fruit production occurs in a year of sub-
normal precipitation. During the course of this study, rainfall levels were above
normal in five of six years. In plants on south-facing slopes there were consecutive
years of large fruit crops. Stepwise multiple regression analysis revealed that total
fruit production per slope face was correlated significantly with high precipitation in
February, just prior to flowering. Fruit production by individual shrubs, however,
was correlated with different parameters on north-facing vs. south-facing slopes. On
the former aspect, fruit production was higher in years with warmer March temper-
atures and on the latter aspect larger fruit crops were correlated positively with October
temperature and December precipitation.

Chaparral vegetation dominates the drier slopes and foothills
throughout cismontane California. Its fire-prone nature and ability
to regenerate rapidly after such disturbance is well documented (Hanes
1977). Many of the dominant shrub species are capable of regen-
eration by vegetative sprouts from basal burls and rootcrowns,
whereas others depend upon seedling establishment. Noteworthy
among the latter type of shrub are the majority of species in Cea-
nothus and Arctostaphylos. These species have no adaptations for
vegetative regeneration and are referred to commonly as obligate
seeding shrubs (Wells 1969). The shrubs begin fruit production be-
tween five and 10 years of age and continue flowering and fruiting
throughout their lifespan. Although seeds are dispersed soon after
maturation, they remain dormant in the soil and duff until germi-
nation is stimulated by fire, or occasionally by other disturbances.

Previous studies of Ceanothus and Arctostaphylos have shown that
the bulk of the seed crop is lost to predation soon after dispersal

MaproNo, Vol. 34, No. 4, pp. 273-282, 1987

274 MADRONO [Vol. 34

(Keeley 1977, Davey 1982). Thus, the soil seed banks are relatively
dynamic and have large fluxes in and out following dispersal of seed
crops. Over long periods of time there may be relatively little increase
in size of the seed banks (Keeley 1987). Consequently, postfire re-
production for obligate seeding shrubs may be tied intimately to
seed production in the preceding years.

Flower and fruit production has been studied in some chaparral
shrubs. In general, for both obligate and facultative seeding species
of Ceanothus and Arctostaphylos, substantial fruit crops are not pro-
duced every year (Keeley 1977, Baker et al. 1982, Davey 1982,
Schlesinger et al. 1982). The size and periodicity of fruit crops re-
ported for various chaparral shrubs 1s highly variable and the causes
for such variability are unknown. The purpose of the present study
was to document fruit production patterns for one such species over
a period of many years. Specific questions addressed were: 1) What
is the extent of year to year variability? 2) Is this variability related
to patterns of precipitation? 3) What effect does slope aspect have
on fruit production patterns? 4) To what extent do individual plants
vary in fruit production? and 5) Is there any relationship between
the size of fruit crops in one year and the size the following year?

Ceanothus crassifolius (nomenclature for all species follows Munz
1974) was selected because it is an obligate seeding shrub and is
restricted to chaparral vegetation. It is common throughout much
of southern California and dominates slopes away from the im-
mediate coast from northern San Diego Co. to San Bernardino and
Santa Barbara cos. Ceanothus crassifolius is similar in life history
to C. megacarpus, which was described in detail by Schlesinger et
al. (1982). Seed germination and seedling establishment of C. cras-
sifolius is restricted to the first spring after fire; thus, it normally
forms even-aged stands. Flowering may begin as early as five years
of age, although substantial flower and fruit production may not
occur until 15 years of age or older. Flowering continues sporadically
until the next wildfire, which typically occurs every 20-40 years in
southern California.

STUDY SITES AND METHODS

Study sites. This study was conducted in the San Dimas Experi-
mental Forest maintained by the U.S. Forest Service on the coastal
front of the San Gabriel Mountains in Los Angeles Co. Two pop-
ulations on north-facing and two populations on south-facing slopes
with inclinations of 15—20° were selected. The slopes were on op-
posite sides of two east—west running ridges adjacent to Dalton Can-
yon Road between elevations 725 and 750 m. These stands had
burned in 1960 and, thus, the plants were 17 years old at the be-
ginning of this study in 1978.

1987] KEELEY: CEANOTHUS CRASSTIFOLIUS 243

Methods. Vegetative cover on each slope was characterized by
measuring shrub coverage on each slope with two 25 m line transects.
Fifteen shrubs were selected randomly on each slope and tagged. A
few tagged shrubs died during the study and were replaced with
adjacent individuals, so that the sample size remained the same
throughout the study. Height and diameter of the nearly circular
canopy were recorded in the third year of the study and did not
change appreciably through the remainder of the study.

Fruit production was measured on each shrub in late spring prior
to seed dispersal. A 0.25 m?’ hoop was placed randomly on each
shrub and all fruits within the hoop were counted. This was done
for six consecutive years (all sites were destroyed by a controlled
burn in 1983).

Precipitation and temperature data were derived from the NOAA
(1977-1983). The nearest station at this elevation with complete
precipitation data was Big Tujunga Dam (710 m), which is 40 km
to the west of the study site. This site, however, lacked temperature
data. The Mt. Wilson Station (1740 m) had complete temperature
and precipitation data and was used in the stepwise regression anal-
ysis. Although this site is at a substantially higher elevation than Big
Tujunga, the yearly variations in weather patterns are similar. For
example, over the period 1977-1983 monthly precipitation was cor-
related significantly between Big Tujunga and Mt. Wilson (r = 0.96,
p < 0.001, n = 84).

A one-way ANOVA was used to test for differences in level of
fruit production between years and among individuals on the same
slope. Within a given year, a two-tailed t-test was used to test for
differences between the different aspects on the same ridge and, for
the same aspect, for differences between ridges. The Kendall coef-
ficient of rank correlation analysis was used to test for a relationship
between the size of the fruit crop on an individual shrub in a given
year with the size of the fruit crop the following year. A stepwise
multiple regression analysis was made between fruit crop size and
monthly precipitation and mean temperature.

RESULTS

Cover. Total shrub coverage of all species (percentage ground sur-
face covered) was markedly greater on the two north-facing slopes;
120% and 130% for Ridges 1 and 2, respectively, vs. 70% and 90%
on the southern exposures. Ceanothus crassifolius comprised nearly
all of the cover on the south-facing slopes, but only approximately
three-fourths of the cover on the north-facing slopes. The mean
height of the C. crassifolius shrubs was slightly more than 2.5 m
on all slope faces, but generally the shrubs on the south-facing ex-
posures had larger canopies. The largest individual shrubs were on

276 MADRONO [Vol. 34

TABLE 1. COMPARISON OF Ceanothus crassifolius INDIVIDUALS ON NORTH-FACING
AND SOUTH-FACING SLOPES IN THE SAN GABRIEL MOUNTAINS. Fruit production was
measured on 15 individuals on each of two north-facing and two south-facing slopes
for six years (slope aspects were compared with a 2-tailed t-test, ns = p > 0.05, ** =
p < 0.01). Each fruit potentially can mature three seeds.

Annual fruit

production
Average areal cover (fruits/m? areal
Height (m) (m?/shrub) _coverage)
Slope aspect X + s.d. (n) X + s.d. (n) X + s.d. (n)
Ridge |
North 2.5 + 0.3 (15) 3.0 se 2515) 284 + 497 (90)
ns ** KK
South 2.6 + 0.3 (15) 7.9 + 3.4(15) 576 + 515 (90)
Ridge 2
North 2.6 + 0.6 (15) 22 2 1.015) 223 + 515 (90)
ns ns led
South 2.6 £°0.3'(15) 3.22 21 (15) 386 + 405 (90)

the south face of Ridge 1, which was the most open of the four slope
faces.

Fruit production. Due to the different sizes of shrubs on these
slopes, fruit production is expressed on an areal coverage basis.
Averaged over the six years of this study, the number of fruits
produced per m? of areal coverage was significantly greater on south-
facing exposures than on north-facing exposures (Table 1). Level of
fruit production was similar between the two north-facing slopes,
but the south face of Ridge 1 had significantly greater annual fruit
production than that exposure on Ridge 2 (p < 0.01). Because there
was much plant to plant variation and yearly variation in size of
fruit crops, the variance in fruit production was generally high on
all slopes (Table 1).

Annual patterns of fruit production are illustrated in Fig. 1. For
all exposures there was a significant difference in fruit production
between years. For all slopes, 1978 wasa year of high fruit production
and, for the south-facing exposures, this was followed by another
year of high fruit production. The shrubs on the southern exposures
significantly (p < 0.01) out-produced shrubs on the northern ex-
posures in all years except 1978 on Ridge | and 1979, 1982, and
1983 on Ridge 2. In all years, fruit production between the two
north-facing slopes was not significantly different (p < 0.05). In 1980
and 1981, production on the south-facing slope on Ridge 1 was
significantly higher than the same exposure on Ridge 2 (p < 0.01).

Within a given year the variation in fruit production between
plants on a single slope was high; the coefficient of variation usually

1987] KEELEY: CEANOTHUS CRASSIFOLIUS a)

Ppt. (mm) 1493 803 1086 395 723 1478

Ridge #1
Wi North
YA South
800 \
600 \ N \
Y § ,
a Y NS 8
3s 400 \ \ \ N
5 oof BN EN SS aX
: , IY oY I
3 S EX iN EX
= 1978 1979 1980 1981 1982 1983
£ nicge re mE North
= GA South
2 800
600 : \
X ¥ ;
400 N \ \
y | \
200 N \ . \
\ a \ \
1978 1979 1980 1981 1982 1983
Year

Fic. 1. Annual fruit production for years 1978-1983 for Ceanothus crassifolius
on north- and south-facing exposures in Dalton Canyon in the San Gabriel Mountains
of southern California (n = 15 shrubs per slope). Statistical analyses are given in the
text. Precipitation data is for the period July—June and is from Big Tujunga Dam;
average = 660 mm per year.

exceeded 80% and often was above 100%. During the period of this
study there was a significant difference between individuals on the
south face of Ridge 1 (p < 0.01). On this slope, over the six-year
period, one shrub averaged 1140 + 747 fruits m~? areal coverage
per year whereas another produced nearly an order of magnitude
fewer (173 + 80 fruits m~? areal coverage). On the other slopes,
there was much variation in size of fruit crops, but across all six
years there was no significant difference among shrubs.

An analysis of the size of the fruit crop on an individual shrub in
a given year with the size in the following year showed that there
was a Significant positive relationship in fruit production between
1978 and 1979, and between 1980 and 1981 (p < 0.01). Thus, for

278 MADRONO [Vol. 34

those years, shrubs on south-facing slopes that produced the largest
fruit crops in 1978 and 1980 produced the largest crops in the suc-
ceeding year. Shrubs on the northern exposure showed no significant
correlation between any years.

Precipitation and temperature patterns. During the course of this
study, five of the years had above average precipitation levels (Fig.
1). The year of highest fruit production, 1978, was also the year of
highest precipitation (the 1977-1978 rainfall season had more than
double the average). There also was much greater than average rain-
fall during the 1982-1983 season, yet fruit production, particularly
on north-facing exposures, was not high in 1983. In three years of
this study the summer drought was interrupted by measurable pre-
cipitation resulting from unusual subtropical storms; >30 mm pre-
cipitation was recorded in either August or September of 1977, 1978,
and 1982. On south-facing slopes these were followed by years of
high fruit production.

A stepwise multiple regression analysis was made between annual
fruit crop size and precipitation and mean temperature for all months
from March of the previous year through June of the year of the
fruit crop, and including annual precipitation total as well as sum-
mer, fall, winter, and spring precipitation totals. There was a highly
significant positive correlation between the mean fruit crop size per
slope and February precipitation (r = 0.61, p < 0.005, n = 24).
However, this relationship did not hold up if slopes were compared
separately. Using fruit crop size for each shrub on the two north-
facing slopes revealed that only March temperature was positively
correlated with fruit crop size (r = 0.52, p < 0.001, n = 180). On
the south-facing slopes, the stepwise regression included both Oc-
tober temperature and December precipitation (r = 0.35, p < 0.001,
n = 180), both of which were positively correlated with fruit crop
SIZe.

DISCUSSION

Annual variation in fruit production by chaparral shrubs is likely
dependent upon environmental conditions during the season of flow-
ering and fruiting. Flower production by species of Ceanothus (subg.
Cerastes) also may be dependent upon conditions during the pre-
vious growing season. This is because flower buds are produced at
the end of the previous year’s growing season, and as a consequence,
flowering is on old growth branchlets (Keeley 1977, Kummerow et
al. 1981; found in all species of subg. Cerastes, although not easily
recognized, cf. Baker et al. 1982).

I propose a model in which fruit production is a function of the
number of nascent flower bud primordia produced at the end of the
previous growing season, flowering success, and the level of pho-

1987] KEELEY: CEANOTHUS CRASSIFOLIUS 219

Spring of Year (t-1) Spring of Year (t)

Early Mid Late Early Mid Late
Se SSS

Temperature/ Precipitation Temperature/Precipitation

|

Carbohydrate stores | Carbohydrate stores |

; Nascent I Nascent =>
j flower buds | flower buds

Flower Flower
production production |
Fruit crop Fruit crop

Fic. 2. Schematic model of factors influencing the size of fruit crops in Ceanothus
species. Fruit production in year (t) will be a function of the number of nascent flower
buds produced the previous year (t — 1), flowering success, and the level of carbo-
hydrates available during fruit maturation.

tosynthates available during fruit maturation (Fig. 2). This model
may be useful for interpreting annual fruit production patterns in
other chaparral shrubs that produce floral primordia in the year prior
to fruiting [viz., all species of Arctostaphylos, Garrya, and Rhus,
excluding R. (Malosma) laurina, Keeley unpubl. data].

In the model, nascent flower bud production is a function of the
level of carbohydrate stores available at the end of the growing season
and carbohydrates are a function of photosynthate production and
demands by carbohydrate sinks. Photosynthate production by chap-
arral shrubs is insensitive to annual variations in temperature rel-
ative to the role of soil moisture (Oechel et al. 1981). Thus, in years
of high precipitation, carbohydrate production should be higher than
in years of low precipitation. Carbohydrate sinks include growth and
maintenance as well as fruit production. The latter represents a
substantial, and annually variable, carbohydrate drain. Because Ce-
anothus flower buds arise from axillary meristems in the nodes of
leaves, nascent flower bud production also would be an indirect
function of the extent of terminal branch and leaf production during
that growing season.

Flower production should be a function of the number of nascent
flower buds produced the previous year and the available carbo-
hydrate stores present at the time of flowering. Because flowering
takes place in early spring, temperature, through its effect on polli-
nators, presumably plays a role in determining success.

Although I recognize that other factors are involved, I suggest that
this model is conceptually useful in understanding annual variation

280 MADRONO [Vol. 34

in size of fruit crops observed for species of Ceanothus. For example,
in a study of C. greggii in San Diego Co. massive fruit crops were
reported for 1974 (Keeley 1977), a year with a 60% precipitation
deficit (all references to annual precipitation are for the rainfall sea-
son, in this instance | Jul 1973 to 30 Jun 1974, most of which falls
in winter and spring). The considerations discussed above would
predict that 1) the 1973 season should have had high precipitation
(requisite for flower bud production), and 2) poor fruit production
in 1975 (due to limited flower bud production in 1974, a conse-
quence of high fruit production combined with limited precipitation
in 1974); both of these predictions were true. A four-year study of
C. megacarpus in Santa Barbara Co. reported that the highest fruit
production occurred in 1978, which was a year of very high rainfall
(Schlesinger et al. 1982). Although it followed a year of subnormal
precipitation, massive flower bud production in 1977 apparently
was possible as a result of a lack of fruit production that year. This
pattern was seen also for C. crassifolius in the present study, where
the year of highest fruit production (1978) was preceded by a year
of subnormal precipitation, but also one in which fruit production
was low (reported by Davey in 1982 for a nearby site).

It is also apparent that on different slope faces fruit production
may be sensitive to different environmental factors. Higher March
temperatures are correlated with higher fruit production on north-
facing slopes. March is the peak month for flowering and, thus,
warmer temperatures may be critical to pollinator success on these
cooler north-facing slopes. On south-facing sites, higher fruit pro-
duction is correlated with higher December precipitation and Oc-
tober temperatures. These factors may affect seasonal carbon gain
and, thus, overall carbohydrate stores on these south-facing slopes.

One implicit factor in the model (Fig. 2) is the effect of summer
and fall drought on nascent flower bud survival. Baker et al. (1982)
suggested this to be an important factor controlling fruit production
in Arctostaphylos. Bud survival might be affected by total seasonal
precipitation and also by atypical summer thunderstorms. As noted
here, significant summer precipitation occurred in 1977, 1978, and
1982, and these years were followed by high fruit production on
south-facing slopes.

An understanding of the factors responsible for temporal variation
in fruit production is complicated in that there is significant spacial
variation (Fig. 1). Throughout this investigation, the shrubs on
southern exposures out-produced those on northern exposures. The
regression analysis suggests that cooler temperatures during flow-
ering may play a role in controlling fruit production on north-facing
slopes. Soil moisture levels, however, may play a role because results
from other studies would predict higher soil moisture levels for the
sparsely vegetated south-facing slopes (Poole et al. 1981).

1987] KEELEY: CEANOTHUS CRASSIFOLIUS 281

Elevational differences also may affect level of fruit production.
Davey (1982) documented C. crassifolius seed production on a south-
facing slope (last burned in 1960) within | km of my sites, but at
approximately 200 m higher elevation. For the years 1978-1980,
she reported seed fall of 6000-8000 seeds m ~? ground surface, which
for her sites translates into 4000-5500 seeds m~? areal coverage.
Assuming each fruit dispersed the maximum number of seeds pos-
sible (i.e., three), the highest seed fall observed at my sites during
those same years would have been 1500-2500 seeds m ~? areal cov-
erage.

Even within a site there is much interplant variation. On the south-
facing exposures studied here, some shrubs out-produced (by an
order of magnitude) others that were only meters away. That some
shrubs consistently produced larger fruit crops than others nearby
suggests either inherent genetic differences in shrubs, or important
microhabitat differences. Small scale differences in soil depth could
produce different soil moisture regimes (Miller and Hajek 1981);
however, nothing is known about the subsoil conditions at these
sites.

More and longer term studies of fruit production will be needed
before we can elucidate all factors responsible for the annual vari-
ation in magnitude of fruit crops in these chaparral shrubs. Future
studies will need to consider microhabitat characteristics and other
environmental parameters in order to fully account for fruit pro-
duction patterns in these species.

ACKNOWLEDGMENTS

I thank Sterling Keeley for help in sampling, Mary (Hochberg) Carroll for many
helpful comments on Ceanothus biology and reviews of several versions of the manu-
script, and Frank Vasek for helpful manuscript review of an earlier version.

LITERATURE CITED

BAKER, G. A., P. W. RUNDEL, and D. J. PARSONS. 1982. Comparative phenology
and growth in three chaparral shrubs. Bot. Gaz. 143:94-100.

Davey, J. R. 1982. Stand replacement in Ceanothus crassifolius. M.S. thesis, Cal-
ifornia State Polytechnic Univ., Pomona.

HAnes, T. L. 1977. California chaparral. Jn M. G. Barbour and J. Major, eds.,
Terrestrial vegetation of California, p. 417-470. John Wiley, New York.

KEELEY, J. E. 1977. Seed production, seed populations in soil, and seedling pro-
duction after fire for two congeneric pairs of sprouting and nonsprouting chaparral
shrubs. Ecology 58:820-829.

1987. Ten years of change in seed banks of the chaparral shrubs, Arc-
tostaphylos glauca and A. glandulosa. Amer. Midl. Nat. 114: in press.

KUMMEROwW, J., G. MONTENEGRO, and D. KRAuSE. 1981. Biomass, phenology, and
growth. Jn P. C. Miller, ed., Resource use by chaparral and matorral, p. 69-96.
Springer-Verlag, New York.

MILLER, P. E and E. HAJeEK. 1981. Resource availability and environmental char-
acteristics of mediterranean type ecosystems. /n P. C. Miller, ed., Resource use
by chaparral and matorral, p. 17-41. Springer-Verlag, New York.

282 MADRONO [Vol. 34

Munz, P. A. 1974. A flora of southern California. Univ. California Press, Berkeley.

NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION. 1977-1983. Climatolog-
ical data annual summary—California. Vols. 81-87 (No. 13). Environmental
Data and Information Service, National Climatic Center, Asheville, NC.

OECHEL, W. C., W. LAWRENCE, J. MUSTAFA, and J. MARTINEZ. 1981. Energy and
carbon acquisition. /n P. C. Miller, ed., Resource use by chaparral and matorral,
p. 151-183. Springer-Verlag, New York.

PooLg, D. K., S. W. RoBErRTs, and P. C. MILLER. 1981. Water utilization. Jn P. C.
Miller, ed., Resource use by chaparral and matorral, p. 123-149. Springer-Verlag,
New York.

SCHLESINGER, W. H., J. T. GrAy, D. S. GILL, and B. E. MAHALL. 1982. Ceanothus
megacarpus chaparral: a synthesis of ecosystem processes during development
and annual growth. Bot. Rev. 48:71-117.

WELLS, P. V. 1969. The relation between mode of reproduction and extent of
speciation in woody genera of the California chaparral. Evolution 23:264—267.

(Received 23 Jan 1986; revision accepted 9 Mar 1987.)

ANNOUNCEMENT

SANTA CRUZ ISLAND RESEARCH PROJECTS

The Nature Conservancy (TNC) proposes a number of research ques-
tions for which they would like to receive proposals for funding. Projects
will be funded by the Santa Cruz Island Research Fund jointly admin-
istered by the Santa Barbara Museum of Natural History and The Nature
Conservancy. The projects should address the specific question(s) and
provide management recommendations. All projects with management
recommendations should include a section on the possible impacts and
effects that mitigation measures might have on current operations of
the Santa Cruz Island Company. The inclusion of recommendations
does not imply that TNC will act upon them in the immediate future.

The list of questions is divided into two categories: projects dealing
with general baseline studies and inventories of widespread general
utility, and projects dealing with immediate biological and/or manage-
ment concerns. We will review all proposals and grant funding on the
merit of the individual project and the appropriateness of the topic to
the Conservancy’s needs. Several projects have potential for collabo-
ration with the Channel Islands National Park Service or for contrib-
uting new data to the established geographical information system.

For a copy of the list of research questions or for more information,
contact the Preserve Director, Peter Schuyler, Santa Cruz Island Project,
213 Sterns Wharf, Santa Barbara, CA 93101.

SEED DISPERSAL IN CEANOTHUS CUNEATUS
AND C. LEUCODERMIS IN A
SIERRAN OAK-WOODLAND SAVANNA

RAYMOND A. EVANS
USDA/ARS, 920 Valley Road, Reno, NV 89512

HAROLD H. BISWELL
Department of Forestry, University of California,
Berkeley 94720

DEBRA E. PALMQUIST
University of Nevada, Reno 89512

ABSTRACT

Seed dispersal of Ceanothus cuneatus and C. leucodermis was studied in an oak-
woodland community in the central Sierra Nevada of California. As the capsule of
Ceanothus (usually containing three seeds) matures and dries, it opens with force and
ejects its seeds at varying distances. Seed-casting, in relation to date, showed a skewed
polynomial distribution that peaked early in July and gradually tapered off. The active
seed-casting period lasted two weeks in the one year of study. Phenology of fruit
ripening, temperature, and humidity were related directly to time and rate of seed-
casting. About one-third of the seeds fell beneath the canopy, whereas the remainder
were cast away from the shrub in an exponential density distribution. Forty-two
percent of the seeds (average density of 2850 m ”) fell at the edge of the shrub and
1.9% (average density of 10 m~’) at 9 m. The probabilities that seeds would be cast
within specific distances from the shrub were 29% (at the edge of the shrub to 2 m),
33% (4-6 m), and 21% (8-9 m).

Ceanothus comprises 55 species restricted to North America, and
most are found along the Pacific coast of the United States (Reed
1974). Californian species are ecologically diverse and, among other
communities, are found in the chaparral, oak-woodland savannas,
and lower coniferous forests of the Sierra Nevada. Ceanothus is
important as wildlife feed and habitat, and because of its nitrogen-
fixing properties also is important in soil development and conser-
vation (Zavitkovski and Newton 1968, Youngberg and Wollum
1976). Among the oak-woodland savannas of the Sierra Nevada
foothills, C. cuneatus and C. leucodermis are considered weeds to
livestock producers because they compete with the herbaceous vege-
tation and reduce the yield of forage for cattle (Biswell 1974).

As the capsule of Ceanothus matures and dries, it opens with force
and ejects the seeds to varying distances from the parent shrub.
Explosively dispersed seeds have been reported for other genera,
including Dendromecon, Oxalis, Viola, Phlox, Geranium, Alstroe-
meria, Lupinus, Impatiens, Millettia, and Hura (Swaine and Beer

MADRONO, Vol. 34, No. 4, pp. 283-293, 1987

284 MADRONO [Vol. 34

1977). Individuals of these genera cast their seeds 1-45 m, depending
on species and size of plant. Little is known about the advantages
of explosive seed dispersal in terms of seed germination or plant
establishment. Many questions remain unanswered about the adap-
tive advantages conferred on a species by investment in dispersal
structures (Howe and Smallwood 1982).

The primary objectives of this study were to investigate seed dis-
persal patterns of C. cuneatus and C. leucodermis in order to more
fully understand seed bank characteristics that appear advantageous
to the establishment of new plants, and to examine seed dispersal
patterns in light of theoretical models of explosively dispersed seeds
(Beer and Swaine 1977) and the theoretical distribution and prob-
ability distributions of seeds (Peart 1985).

METHODS

Studies were conducted in a California oak-woodland savanna
community, situated in the foothills of the central Sierra Nevada,
70 km northeast of Fresno, at 910 m and with an average rainfall
of 760 mm. Dominant tree species are Pinus sabiniana, Quercus
douglasii, QO. wislizenii, and Aesculus californica, and dominant shrub
species are Ceanothus cuneatus and C. leucodermis. An understory
of herbaceous plants includes the annual grasses, Bromus mollis, B.
diandrus, B. rubens, Avena barbata, A. fatua, and Festuca spp.; and
the broadleaf plants, Erodium botrys, E. cicutarium, Medicago poly-
morpha, and Trifolium spp.

Eight representative plants of Ceanothus cuneatus and C. leuco-
dermis were selected in the study area (C. cuneatus, 1-3.5 m tall and
C. leucodermis, 2—4 m tall; Munz 1973). Transects 10 m long and
0.6 m wide, radiating out in eight compass directions from two large
shrubs, one of each species, were cleared of herbaceous vegetation
and plant litter (Fig. 1). One meter intervals along the transects were
marked by large nails beginning at the edge of the shrub canopy.
There was one sampling point beneath the canopy on each transect
positioned 0.3 m inside the canopy edge. On smaller shrubs, one to
four transects were made along compass directions radiating out
from each shrub, based on the proximity of adjacent shrubs and
terrain. These transects were cleared and marked in a similar fashion
to the large shrubs. No transects were made adjacent to neighboring
shrubs to avoid measuring areas of overlap of seed dispersal from
two or more shrubs.

Sampling for seed dispersal consisted of counting seeds within a
0.1 m? frame every other day during the active casting period. After
seeds were counted, the transects were swept clean with a whisk
broom. The counts were made in early evening or early morning
when seeds were not dehiscing. To minimize seed predation, ants

1987] EVANS ET AL.: SEED DISPERSAL IN CEANOTHUS 285

Fic. 1. Ceanothus cuneatus with cleared transects radiating out from the shrub
in eight cardinal directions.

were poisoned in the transect areas. No indication of seed predation
by rodents or birds was noted during the course of the study.

When the capsules of Ceanothus burst open and dehisce their seeds
an audible pop is heard. At the height of casting on 20 July, seed
dehiscing on the large shrubs of both species was monitored every
30 minutes from 0700-2130 hr by counting the number of pops
heard by an observer standing close to the individual shrub for three
60 second intervals. At the same time, measurements of relative
humidity and temperature were made using a sling psychrometer.
Seasonal weather data were taken from the nearest station, North
Fork Ranger Station, located 16 km south of the study area at 800 m.

Results were analyzed statistically by ANOVA with Duncan’s
multiple range test to determine differences in seed density in relation
to compass direction. Correlation and curvilinear regression tech-
niques were used to portray and analyze seed dispersal relative to
distance from the shrub canopies and to calendar date.

RESULTS

Seed-casting in relation to date. Seed-casting for both C. cuneatus
and C. /eucodermis occurred from 13 July to 11 August 1952. The

286 MADRONO [Vol. 34

w= 099 «x

A -
Y =

30 11.78 +14.83X-4.78X 2+ 0.48X 3 -0.02x 4

25

NO
oO

SEEDS CAST (%)
a

10

16 18 20 22 24 26 28 30
JULY
Fic. 2. Fourth-degree polynomial distribution curve of percentage of Ceanothus

cuneatus seeds cast in relation to calendar date. Multiple correlation coefficient (r) is
significant at p < 0.01.

period of active seed-casting, when more than 95% of the observed
total seeds were dispersed, lasted 14 days (16-30 July). Average
maximum and minimum daily temperatures during active seed cast-
ing were 35.7 + 2.1°C and 15.1 + 2.5°C, respectively. Rain showers
occurred 25 July (2.3 mm) and 30 July (2.5 mm). Partly cloudy
weather accompanied the showers, which increased the humidity
and decreased air temperatures during these periods.

Maximum seed-casting occurred on 18 and 20 July (two and four
days after the active seed-casting period began) for C. cuneatus and
C. leucodermis, respectively (Figs. 2, 3). Seed casting in relation to
date formed a skewed fourth-degree polynomial distribution in which
the numbers of seeds cast increased rapidly from the initiation of
casting and decreased gradually over the following 10-12 days. In
C. cuneatus, a marked decrease in seed dispersal was noted during

1987] EVANS ET AL.: SEED DISPERSAL IN CEANOTHUS 287

N = 0.99 x*x

30 = 0.67+21.3X -5.28xX2 +0.45xX3 -0.01x 4

A
ry

SEEDS CAST (%)

16 18 20 22 24 26 28 30
JULY

Fic. 3. Fourth-degree polynomial distribution curve of percentages of Ceanothus
leucodermis seeds cast in relation to calendar date. Multiple correlation coefficient
(r) is significant at p < 0.01.

the period of higher humidity and rainfall toward the end of active
seed casting and then a subsequent increase with higher temperatures
and lower humidity (Fig. 2). In C. leucodermis, this response was
similar, but of less magnitude (Fig. 3).

Seed-casting in relation to time of day. On 20 July, seed casting
of both species occurred from 0930-2130 hr (Fig. 4). Seed-casting
in C. cuneatus reached a maximum at 1200 hr and continued at a
high level (20/min being counted) until 1700 hr. Seed-casting in C.
leucodermis increased to a high level at 1230 hr and reached similar
maxima at 1730 and 1830 hr. Total seeds cast on 20 July was almost
identical for monitored shrubs of both species (Figs. 2, 3). At the
onset of seed-casting, temperature was 27°C and relative humidity
(RH) 30%. In the most active period of seed-casting, temperatures

288 MADRONO [Vol. 34

40 — TEMPERATURE
--- RELATIVE HUMIDITY

35

30

25

TEMPERATURE (C)
(%) ALIGINNH SAILV 1344

20

—— C.CUNEATUS

100 2-=- C. LEUCODERMIS

60

40

NUMBER OF CAPSULE POPS

20

TIME OF DAY

Fic. 4. Number of capsule pops of Ceanothus cuneatus and C. leucodermis per
three minute period and air temperature and relative humidity for each one-half hour
on 20 July.

ranged from 35—38°C with RH from 15-20%. At cessation of casting,
temperature was 25°C and RH was 37%.

Seed-casting in relation to distance from shrub. Of the total seeds
cast, 32% and 36% fell beneath the shrub canopy of C. cuneatus and
C. leucodermis, respectively. This presumably resulted from seeds
striking branches or leaves or because the seed capsules were oriented
downward or inward. The remainder of seeds cast by both species
formed an exponential density distribution from a high of 42% at
the edge of the shrub to 1.9% at 9 m from the shrub (Fig. 5). Seed

1987] EVANS ET AL.: SEED DISPERSAL IN CEANOTHUS 289

50 25
—— DENSITY FUNCTION |
—-—-— PROBABILITY FUNCTION

f= 0.99 * *

: -0.6645X
> = 1.814+ 39.8426 °°

DENSITY (%)
(%) ALINIGVEOdd

DISTANCE (m)

Fic.5. Density and probability functions of seed-casting distribution of Ceanothus
cuneatus and C. leucodermis in relation to distance from edge of the parent shrub.
Data for both species are combined. Multiple correlation coefficient (r) of the density
function is significant at p < 0.01. Probability function should be interpreted as a
measure of probability of a seed landing within a specific distance range from edge
of the parent shrub.

densities averaged 2850 m ? at the edge of the shrub to 10 m~? at
9 m. Overall dispersal patterns of different shrubs were consistent
in both species and were not significantly different (p <= 0.05) between
species (Table 1). Highest seed densities, those near the shrubs,
agreed with densities reported by Keeley (1977) for C. leucodermis.

The probability of a seed escaping from the parent plant and
landing within a specific distance range was derived by P(r) =

290 MADRONO [Vol. 34

TABLE 1. CALCULATED SEED DENSITIES OF Ceanothus cuneatus AND C. leucodermis
AT VARIOUS DISTANCES FROM THE PARENT SHRUBS WITH CONFIDENCE INTERVALS
DERIVED FROM MULTIPLE REGRESSION. Correlation coefficient for multiple regres-
sion = 0.99; n = per 0.1 m7?.

Distance from edge C. cuneatus C. leucodermis — Confidence interval
of shrub (m) (n) (n) p = 0.05
0 273 oT 16
| 163 167 10
2 92 96 8
3 53 a7 8
4 31 35 8
5 19 23 8
6 16 8 9
7 8 12 9
8 6 10 9
9 5 9 9

2nmrD(r)dr (Peart 1985), where P(r) = probability of a seed escaping
to an angular area at distance (r) from the point of release; D(r) =
a density-dispersal function based on actual sampling data of the
two species; and dr = sample size of 0.1 m. Results indicated that
the probability of a seed landing within specific distance ranges from
the parent shrub varied from 29% (edge of the shrub to 2 m), to
33% (4-6 m), and to 21% (8-9 m) (Fig. 5).

Seed-casting in relation to direction from shrub. Over the active
period of casting, most C. cuneatus seeds were cast in a southwesterly
direction and fewest in a westerly direction. Early in the casting
period and at maximum seed dispersal more seeds were cast to the
north and east, whereas later in the casting period most seeds were
cast in the southwesterly direction. In C. leucodermis, most seeds
were cast in a southeasterly direction and fewest were cast in a
westerly direction. Most seeds were cast to the southeast during the
10 days of most active seed dispersal. In both species, fewer seeds
were cast to the west throughout the dispersal period (Table 2).

DISCUSSION

Seed-casting in Ceanothus cuneatus and C. leucodermis occurs
during a short but active period (14 days in this study), when the
capsules ripen and expel their seeds (usually three per capsule) to a
distance of 9 m or more. Seeds are dispersed mainly beneath or near
parent shrubs, but explosive dispersal permits a wider distribution
of some seeds. Both diurnal and seasonal trends of seed-casting seem
to indicate that specific thresholds of temperature and moisture were
critical for seed-casting in Ceanothus. Our results suggest that com-
mencement and rate of seed-casting are functions of the phenological

291

EVANS ET AL.: SEED DISPERSAL IN CEANOTHUS

1987]

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292 MADRONO [Vol. 34

stage of fruit ripening and of the temperature and relative humidity
as they affect plant moisture status.

Ceanothus seeds require relatively high temperatures (70—100°C)
for germination (Reed 1974). Most germination and seedling estab-
lishment is associated with fire (Quick and Quick 1961, Schlesinger
and Gill 1978). Between fires, large seed banks of C. cuneatus and
C. leucodermis beneath and around shrubs can accumulate in the
Sierran oak-woodland savanna.

Effects of the interaction of seed dispersal at varying distances
from the parent shrubs and seed mortality in relation to fire tem-
perature as influenced by type (shrub vs. herbaceous) and amount
of fuel create a mosaic of safe sites (Harper et al. 1965) for seed
germination and plant establishment. Seedling establishment of Ce-
anothus in relation to competition from herbaceous species (Schultz
et al. 1955) that grow among shrubs increases heterogeneity of the
seed bank in terms of safe sites and restricts opportunities for suc-
cessful establishment of seedlings. Furthermore, wide dispersal of
long-lived seeds of Ceanothus increases the probability for estab-
lishment of new plants by reducing the effects of seed predation by
insects, predation and herbivory by rodents, and intraspecific seed-
ling competition (Peart 1985).

ACKNOWLEDGMENTS

The senior author acknowledges the tireless efforts and exceptional ability of his
wife, Dorthea B. Evans, in assisting in the field sampling of the study. I also thank
her for encouragement and inspiration during all phases of the study.

LITERATURE CITED

BEER, T. and M. D. SwaIne. 1977. On the theory of explosively dispersed seeds.
New Phytol. 78:68 1-694.

BISWELL, H. H. 1974. Effects of fire on chaparral. Jn T. T. Kozlowski and C. E.
Ahlgren, eds., Fire and ecosystems, p. 321-364. Academic Press, New York.

HARPER, J. L., L. T. WILLIAMS, and G. R. SAGAR. 1965. The behavior of seeds in
soil. I. The heterogeneity of soil surfaces and its role in determining the estab-
lishment of plants. J. Ecology 539:273-286.

Howe, H. F. and J. SMALLWoop. 1982. Ecology of seed dispersal. Ann. Rev. Ecol.
Syst. 13:201-228.

KEELEY, J. E. 1977. Seed production, seed populations in soil, and seedling pro-
duction after fire for two congeneric pairs of sprouting and nonsprouting chaparral
shrubs. Ecology 58:820-829.

Munz, P. A. 1973. A California flora with supplement. Univ. California Press,
Berkeley.

PEART, D. R. 1985. The quantitative representation of seed and pollen dispersal.
Ecology 66:1081-1083.

Quick, C. R. and A. S. Quick. 1961. Germination of Ceanothus seeds. Madrono
16:23-30.

Reep, M. J. 1974. Ceanothus. In C. S. Schopmeyer, tech. coord., Seeds of woody
plants in the United States, p. 284-290. Agriculture Handbook No. 450, Forest
Service, U.S.D.A., Washington, DC.

1987] EVANS ET AL.: SEED DISPERSAL IN CEANOTHUS 293

SCHLESINGER, W. A. and D. S. GILL. 1978. Demographic studies of the chaparral
shrub, Ceanothus megacarpus in the Santa Ynez mountains, California. Ecology
59:1256-1263.

ScHuLtTz, A. M., J. L. LAUNCHBAUGH, and H. H. BISweLL. 1955. Relationship
between grass density and brush survival. Ecology 36:226-238.

Swaln, M. D. and T. BEER. 1977. Explosive seed dispersal in Hura crepitans L.
(Euphorbiaceae). New Phytol. 78:695-708.

YOUNGBERG, C. T. and A. G. WoLLuM II. 1976. Nitrogen accretion in developing
Ceanothus velutinus stands. Soil Sci. Soc. Amer. Proc. 40:108-112.

ZAVITKOVSKI, J. and M. NEwTon. 1968. Ecological importance of snowbrush Ce-
anothus velutinus in the Oregon Cascades. Ecology 49:1134-1145.

(Received 14 Apr 1986; revision accepted 9 Mar 1987.)

ANNOUNCEMENT

THE 1987 JESSE M. GREENMAN AWARD

The 1987 Jesse M. Greenman Award has been won by Geoffrey A.
Levin for his publications “‘Systematic Foliar Morphology of Phyllan-
thoideae (Euphorbiaceae). I. Conspectus”, “Systematic Foliar Mor-
phology of Phyllanthoideae (Euphorbiaceae). II. Phenetic Analysis’,
which appeared in the Annals of the Missouri Botanical Garden, volume
73, number 1, and “Systematic Foliar Morphology of Phyllanthoideae
(Euphorbiaceae). III. Cladistic Analysis’, which was published in Sys-
tematic Botany, volume 11, number 4. This series of papers is derived
from a Ph.D. dissertation from the University of California, Davis,
under the direction of Drs. James A. Doyle, Grady L. Webster, and
Jack A. Wolfe. Dr. Levin uses a large set of characters (in this case leaf
characters) to address questions of systematic relationships and phy-
logeny at higher taxonomic levels, using the results from both phenetic
and cladistic analysis to evaluate a more traditional classification sys-
tem, and to identify genera or groups of genera whose position and
relationship are not clear and, therefore, are in need of additional study.

The Award is named for Jesse More Greenman (1867-1951), who
was Curator of the Missouri Botanical Garden Herbarium from 1919
until 1943. A cash prize of $250 is presented each year by the Garden,
recognizing the paper judged best in vascular plant or bryophyte sys-
tematics based on a doctoral dissertation that was published during the
previous year. Papers published during 1987 are now being considered
for the 20th annual award, which will be presented in the summer of
1988. Reprints of such papers should be sent to: Greenman Award
Committee, Division of Research, Missouri Botanical Garden, P.O. Box
299, St. Louis, MO 63166-0299, U.S.A. In order to be considered for
the 1988 award, reprints must be received by 1 June 1988.

SEEDCROP CHARACTERISTICS AND MINIMUM
REPRODUCTIVE SIZE OF ORGAN PIPE CACTUS
(STENOCEREUS THURBERT) IN SOUTHERN ARIZONA

KATHLEEN C. PARKER
Department of Geography, University of Georgia,
Athens 30602

ABSTRACT

Seedcrop characteristics and the relationship of reproductive activity to size of
organ pipe cactus (Stenocereus thurberi) were examined in Organ Pipe Cactus National
Monument, Arizona. A sample of 19 fruits collected had a mean diameter of 52.9
mm and a mean seed content of 1969 seeds/fruit. Laboratory germination percentage
at 20—25°C and a 12 hr photoperiod was 88%. Height measurements and fruit pres-
ence/absence observations made at two locations in the Monument indicate that most
S. thurberi individuals begin to reproduce when they are 2—2.5 m in height. Plants
typically have 4-10 arms by the onset of reproductive maturity. Large individuals
of S. thurberi may produce more than 50 fruits in a season. These results indicate
that the reproductive potential of mature individuals of this species is high.

The giant cactus forests of the Southwest have captured the interest
of botanists and travelers to that region for over a century. Although
three of the columnar cactus species found in the Sonoran Desert
occur naturally in the United States, the vast majority of scientific
studies on columnar cactus conducted in this country (e.g., Shreve
1910, Niering et al. 1963, Steenbergh and Lowe 1969, 1977, 1983)
have focused on saguaro [Carnegiea gigantea (Engelm.) Britt. &
Rose], the most widespread and conspicuous of the columnar cacti
in the United States. In favorable habitats, organ pipe cactus [Steno-
cereus thurberi (Engelm.) Buxb.] (Fig. 1) also is prominent, but its
range within the United States is restricted to several populations
in southern Arizona (Hastings et al. 1972). It has received much less
attention from scholars than C. gigantea; consequently, we know
less about the basic ecology and population dynamics of S. thurberi
than of C. gigantea. Recent studies have begun to identify ecological
characteristics (Nobel 1980, Smith et al. 1984) and site preferences
(Yeaton and Cody 1979) of S. thurberi in the northern part of its
range, but McDonough’s (1964) study of factors affecting seed ger-
mination is the only work published on the reproductive character-
istics of this species.

The purpose of this study was to determine the size at which
individuals become reproductively active and the fruit size and seed
production and germinability for populations of S. thurberi in south-
ern Arizona.

MADRONO, Vol. 34, No. 4, pp. 294-303, 1987

1987] PARKER: ORGAN PIPE CACTUS 295

-

: - ¥
= Sahil ‘

Wa he
a“ Ck ae

~

Fic. 1. Fruiting individual of Stenocereus thurberi in Organ Pipe Cactus National
Monument.

METHODS

Study area. The study was conducted in Organ Pipe Cactus Na-
tional Monument (OPCNM), which supports one of the most ex-
tensive populations of S. thurberi in the United States. Data were
collected in late June and early July 1976, approximately midway

296 MADRONO [Vol. 34

through the flowering and fruiting season for reproductively active
plants.

Mean annual precipitation at the Monument headquarters is 233
mm (Weather Bureau 1951-1974, NOAA 1975-1980), although
data from a network of backcountry raingauges indicate that rainfall
is generally higher in the Ajo Mountains along the eastern boundary
of the Monument because of orographic uplift (Table 1). For the 17
yr period from 1962-1983 (exclusive of 1967 and 1973-1976 when
backcountry records were incomplete), annual rainfall at a remote
station in the Ajo Mountains exceeded rainfall at the official weather
station in the Monument by a mean value of 101 mm (s.d. = 148
mm, range = —45 to 418 mm). Precipitation is distributed bimodally
throughout the year, with the primary rainfall maximum coinciding
with the time of fruit maturation during summer and the secondary
maximum occurring during winter. Annual precipitation for the year
prior to the study (1975) was well below the mean, and the summer
of that year was the driest of the last 35 yr (1950-1984; Table 1).

Nocturnal freezes occur occasionally in the Monument (X = 19
freezes/year; Table 1), but no subfreezing daily maximum has been
recorded at the Monument headquarters in its 42 yr history as a
weather station. The winter preceding the study (1975-1976) had a
typical number of freezes, with —5°C as the lowest temperature
recorded that winter.

Sample sites. To analyze the relationship between size and repro-
ductive activity, data were collected from two sites within OPCNM
that had similar slope characteristics. Both were on south—southwest-
facing rocky hillsides with slope angles from 15—20° and approxi-
mately 25-50 m above the the adjacent valley floor. Both valleys
sloped gently southward as part of the Sonoyta River drainage net-
work. Soils in both sites were shallow gravelly loams. The Twin
Peaks site (31°57'N, 112°49’W) was located north of the Monument
campground (within 2 km of the weather station) on the lower slopes
of Twin Peaks between 1740-1800 m. The site was sampled with
ten 7 X 50 m rectangular quadrats. The Ajo Mountain site (32°00'N,
112°42'W) was located on the lower slopes of the Ajo Mountains
from 2120-2200 m (within 2 km of the backcountry raingauge in
the Ajo Mountains). This site was sampled with three 7 x 50 m
quadrats. The area sampled was smaller for the Ajo Mountain site
because there was less homogeneous habitat available for quadrat
placement than in the Twin Peaks site. For all individuals within
the sampling quadrats, the height of the tallest arm, the number of
arms, and the presence or absence of buds, flowers, or fruits were
recorded.

Fruit and seed samples. Nineteen ripe fruits were collected from
S. thurberi plants within the two study sites. Each fruit was taken

1987] PARKER: ORGAN PIPE CACTUS 297

TABLE 1. CLIMATIC SUMMARY AND CONDITIONS IN THE YEAR PRIOR TO STUDY IN
ORGAN PIPE CACTUS NATIONAL MONUMENT. Means and standard deviations (s.d.)
for the official station (near Twin Peaks) are calculated for the period 1951-1980
with data published by the Weather Bureau (1951-1974) and NOAA (1975-1980);
mean and s.d. for freeze frequency are based on the 29 yr period excluding 1980
because of missing data. The previous year is defined as 1975 for annual and summer
precipitation and as the winter of 1975-1976 for the number of freezes/winter. Mean
and s.d. for the Ajo Mountains are based on data from a backcountry raingauge
monitored by the Monument staff for the period 1962-1983 (exclusive of 1967 and
1973-1976).

Official station Ajo Mountains
Pre-
vious
X+s.d. Range year xX = Sid: Range
Annual precipitation
(mm) 233 + 78 87-377 111 342 2. 1547132: 657
Summer precipitation
(mm) (June-
September) 107 + 56 17-192 17 — —
Freezes/winter 19+7 5-34 20 — —

from a different individual. After removal of any persistent spines
and dried flower parts, fruit lengths and maximum diameters were
measured with a dial caliper graduated by 0.05 mm. Fruits were
weighed with a triple beam balance immediately after collection.
Seeds were then separated from the flesh of the fruits with a sieve.
Seeds were air dried and weighed, and the number per fruit was
determined. Seeds were stored for about 30 days in the dark at 20—-
25°C before they were used in germination tests.

Seed germinability was determined on a random sample (100
seeds) of those collected. These were placed in covered glass dishes
on moist loam and sand combined in a 1:1 ratio. The dishes were
kept at 20—25°C and exposed to 12 hr of fluorescent light/day (400
wE-m~?’-s~'). Germination was monitored daily for five days.

RESULTS

Size and reproductive activity. Differences were apparent between
the two study sites in the relationship of reproductive activity to
size (Fig. 2, Table 2). The shortest individuals sampled that bore
flowers or fruits were 0.99 m tall in the Twin Peaks site and 1.32
m tall in the Ajo Mountain site. In the Twin Peaks site, only three
of 14 plants that flowered and were less than 1.49 m in height failed
to produce fruits, whereas in the Ajo Mountain site, two of the four
reproductively active individuals in the same size range failed to
fruit. Despite the greater minimum height of reproductive activity
in the Ajo Mountain site, the threshold height above which all plants

298 MADRONO

e
O

Twin Peaks
Ajo Mountains

00
O

zs 9
WF F

pe)
Oo

DWWW °>°’°F"FFBAA

PERCENTAGE OF HEIGHT CLASS
SHOWING REPRODUCTIVE ACTIVITY

oO

oo’ WY

og

HEIGHT CLASS (M)

Fic. 2. The percentage of each height class showing evidence of reproductive
activity of Stenocereus thurberi in the two study plots. Numbers above the bars
indicate sample size for each height class.

in that sample produced flowers or fruits was lower (1.50 m) than
in the Twin Peaks sample (2.50 m). Similarly, in the Ajo Mountain
sample, all plants with more than seven arms produced flowers and
fruits (Table 2) during the year of study, whereas in the Twin Peaks
sample several plants with more than 10 arms did not flower. Three
of the individuals in the Twin Peaks site greater than 2 m tall and
with at least seven arms bore flowers, but not fruits. Despite these
differences between the two samples, the overall relationship of size
to reproductive activity suggests that most individuals of S. thurberi
begin reproductive growth by the time they are 2—2.5 m tall. By this
time, most plants have 4-10 arms.

Seed production per fruit and germinability. Fruits of S. thurberi
are spherical in shape (Fig. 3, Table 3). The mean weight of the ripe
fruits collected was 73.03 g. The significant intercorrelation (p <
0.001; Table 4) between fruit dimensions and weight indicates that
these characteristics all vary proportionally. On average, seeds ac-
counted for only 3.76 g of the total fruit weight. The mean number
of seeds per fruit was 1969, and larger fruits generally produced
more seeds than smaller fruits (Table 4). The mean seed weight per

1987] PARKER: ORGAN PIPE CACTUS 299

TABLE 2. RELATIONSHIP OF REPRODUCTIVE STATUS TO ARM NUMBER IN S. thurberi
AT Two SITES IN ORGAN PIPE CACTUS NATIONAL MONUMENT. Reproducing plants
include individuals bearing buds, flowers, or fruits. RP = the number of reproducing
plants; n = the sample size for each arm-number category.

Twin Peaks (N = 207) Ajo Mountains (N = 60) Total (N = 267)

Nee Reproducing plants Reproducing plants Reproducing plants
arms RP/n % RP/n % RP/n %

l 0/11 (0.0) — _ 0/11 (0.0)

2 0/19 (0.0) 1/5 (20.0) 1/24 (4.2)

3 1/27 (3.7) 0/5 (0.0) 1/32 (3.1)

4 6/23 (26.1) 6/14 (42.9) 12/37 (32.4)

5 7/16 (43.7) 4/5 (80.0) 11/21 (52.4)

6 13/24 (54.2) 2/5 (40.0) 15/29 (51.7)

7 10/16 (62.5) 5/6 (83.3) 15/22 (68.2)

8 7/10 (70.0) 3/3 (100.0) 10/13 (76.9)

9 9/15 (60.0) 5/5 (100.0) 14/20 (70.0)

10 10/10 (100.0) 4/4 (100.0) 14/14 (100.0)

>10 33/36 (91.7) 8/8 (100.0) 41/44 (93.2)

fruit was not correlated significantly with any of the other fruit
characteristics measured (Table 4). Eighty-eight percent of the S.
thurberi seeds planted had germinated after five days.

DISCUSSION

Reproductive characteristics. In Saguaro National Monument, lo-
cated approximately 150 km east of OPCNM, Steenbergh and Lowe
(1977) reported a minimum reproductive height for C. gigantea
similar to that reported here for S. thurberi. They found that all
individuals greater than 2.5 m tall produced reproductive structures
and that the smallest individual that showed evidence of reproduc-
tive activity was between 1.5 and 1.99 m tall. Steenbergh and Lowe
(1977) concluded that healthy C. gigantea individuals typically reach
reproductive maturity at a height of 2.2 m, or an age of about 30
yr.

Important differences exist between the two species in the rela-
tionship of arm number to reproductive activity. Steenbergh and
Lowe (1977) found that individuals of C. gigantea begin to develop
arms after they reach a height of approximately 4.5 m, or more than
twice the size at which they typically begin reproducing. In southern
Arizona, S. thurberi consists of many relatively narrow stems (ca.
15 cm diameter) that emerge from the base of the plant, rather than
a primary stem with arms forming several meters above the base.
Unlike C. gigantea, S. thurberi individuals generally do not begin
reproducing until after they have more than one arm.

Steenbergh and Lowe (1977) hypothesized that the production of

300 MADRONO [Vol. 34

Fic. 3. Fruit of Stenocereus thurberi.

1987] PARKER: ORGAN PIPE CACTUS 301

TABLE 3. DIMENSIONS, WEIGHT, AND SEED NUMBER OF FRUITS OF Stenocereus
thurberi (n = 19). Whole fruits were measured with spines and desiccated flower parts
removed; seeds from each fruit were air dried after removal of pericarp and flesh.
The mean weight/seed by fruit for the sample was calculated by averaging the means
for each fruit.

Measurement Dar al: Range

Whole fruits

Length (mm) dial = 6:5 41.4-63.3

Diameter (mm) 52.9 + 4.9 41.7-63.5

Weight (g) 73.03 + 23.18 32.19-125.22
Seeds per fruit

Total weight (g) 3.76 + 1.43 1.63-6.56

Total number 1969 + 703 688-3373

Weight/seed by fruit (mg) 1.91 + 0.23 1.47-2.37

arms in C. gigantea increases its reproductive potential and that this
is their primary function. Undoubtedly, individuals of S. thurberi
with numerous arms have a greater reproductive potential than those
with few arms, because fruits are only borne on the upper portion
of the arms. Whether enhancement of reproductive capacity serves
as a Selective force in arm production by S. thurberi, or is simply a
fortuitous consequence of a basally-branched form conferred by dif-
ferent selective constraints is debatable.

Steenbergh and Lowe (1977) reported a slightly greater mean seed
number per fruit for C. gigantea than I obtained for S. thurberi,
although results of a t-test (t = 1.51, df = 31) indicate that this
difference is not significant (p < 0.05; data for C. gigantea fruits
were taken from Steenbergh and Lowe 1977). The mean seed weight
for S. thurberi (1.9 mg) is greater than that reported by Steenbergh
and Lowe (1977) for C. gigantea (1.3 mg), but the absence of a
standard deviation value for mean seed weight for C. gigantea pre-
cluded calculation of the t-statistic to determine whether this dif-
ference is statistically significant.

TABLE 4. SPEARMAN CORRELATION COEFFICIENTS (r,) BETWEEN FRUIT CHARACTER-
ISTICS (n = 19). ** = significant at p < 0.0001; * = significant at p < 0.05.

Mean Total
weight/ seed Fruit
seed Seed number weight Fruit weight diameter
Fruit length 0.04 0.45 0.39 0.86** Osa
Fruit diameter =(.03 0.47* 0.42 O:95**
Fruit weight 0.11 0.47* 0.42
Total seed weight 0.10 0.94**

Seed number —0.10

302 MADRONO [Vol. 34

The germination percentage that I found for S. thurberi was similar
to the 91% germination for this species (at 25°C with 8 hr photo-
periods for 6 days) reported by McDonough (1964). Under the same
conditions, he found a slightly higher percentage germination (97%)
for C. gigantea.

Relationships between environment and reproductive activity. The
mean number of fruits borne by individuals of S. thurberi was not
quantified. Many plants observed during the course of data collec-
tion, however, bore at least 50 fruits. With a mean of approximately
2000 seeds per fruit, individuals that bear more than 50 fruits pro-
duce about 100,000 seeds in a single season. The results of the
germination test indicate that a high percentage of the seeds produced
by an individual have the potential of germinating if environmental
conditions are favorable. Thus, the reproductive potential of even
a small population of S. thurberi is great. Field germination per-
centages and the survival of seedlings, however, have not been de-
termined.

In the Twin Peaks site, some large individuals of S. thurberi did
not reproduce during 1976, and some that flowered failed to set
fruit. Flowering and fruiting among large plants were more consistent
in the Ajo Mountain site than in the Twin Peaks site. Variation in
moisture regimes between the two study sites may be responsible,
in part, for the differences in reproductive activity. In a particularly
dry year, such as the one preceding the study, the higher rainfall
characteristic of the Ajo Mountains may foster consistent repro-
ductive activity of S. thurberi occurring there, while reproductive
activity is more sporadic in drier parts of the Monument. Although
Thackery and Leding (1929) and Steenbergh and Lowe (1977) re-
ported that drought had little influence on fruit production in the
closely related C. gigantea, Thackery and Leding (1929) suggested
that reproductive activity in S. thurberi is more sensitive to drought
stress than in C. gigantea. Steenbergh and Lowe (1977) also reported
that severe freezes may reduce greatly the reproductive activity of
C. gigantea the following summer. It is unlikely, however, that spa-
tial variation in the occurrence of severe freezes caused the differ-
ences in reproductive activity between the two S. thurberi sites be-
cause of their similar topographic positions (i.e., susceptibility to
cold air drainage).

The reproductive traits of S. thurberi are well adapted to the
variable environment characteristic of the region of study. Near the
margin of its range, successful establishment of young individuals
of S. thurberi is limited by frequent severe freezes (Nobel 1980) and
by periodic prolonged drought. Most individuals of S. thurberi be-
come reproductively active at heights from 2—2.5 m. The fruiting of
very large individuals indicates that plants are reproductively active

1987] PARKER: ORGAN PIPE CACTUS 303

throughout most of their adult life. By producing a large quantity
of seeds every year for many years, individuals of S. thurberi improve
the chance that an occasional seed will disperse to a site favorable
for germination and growth in a year when climatic factors are
favorable, thereby maintaining a stable population.

ACKNOWLEDGMENTS

I thank the staff of Organ Pipe Cactus National Monument, who provided me with
precipitation data from their remote weather stations, Warren F. Steenbergh, who
encouraged me to examine these questions, Albert J. Parker, who helped collect the
data and made helpful comments on the manuscript, and Thomas R. Vale, who read
an earlier version of the manuscript and made valuable suggestions.

LITERATURE CITED

HASTINGS, J. R., R. M. TURNER, and D. K. WARREN. 1972. An atlas of some plant
distributions in the Sonoran Desert. Tech. Rep. on the Meteorology and Cli-
matology of Arid Regions No. 21. Univ. of Arizona Institute of Atmospheric
Physics, Tucson.

McDonouGuH, W. T. 1964. Germination responses of Carnegiea gigantea and Le-
maireocereus thurberi. Ecology 45:155-159.

NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION. 1975-1980. Climatolog-
ical data: Arizona. Vol. 79-84. U.S. Dept. Commerce, Washington, DC.

NIERING, W. A., R. H. WHITTAKER, and C.H. Lowe. 1963. The saguaro: a population
in relation to environment. Science 142:15-23.

NoBEL, P. S. 1980. Morphology, surface temperatures, and northern limits of co-
lumnar cacti in the Sonoran Desert. Ecology 61:1-7.

SHREVE, F. 1910. The rate of establishment of the giant cactus. Plant World 13:
235-240.

SMITH, S. D., B. DIDDEN-ZopFy, and P.S. NoBEL. 1984. High-temperature responses
of North American cacti. Ecology 65:643-651.

STEENBERGH, W. F. and C. H. Lowe. 1969. Critical factors during the first years of
life of the saguaro (Cereus giganteus) at Saguaro National Monument, Arizona.
Ecology 50:825-834.

and 1977. Ecology of the saguaro: II reproduction, germination,

establishment, growth, and survival of the young plant. Natl. Park Serv. Sci.

Monogr. Ser. No. 8.

and . 1983. Ecology of the saguaro: III growth and demography. Natl.
Park Serv. Sci. Monogr. Ser. No. 17.

THACKERY, F. A. and A. R. LEDING. 1929. The giant cactus of Arizona: the use of
its fruits and other cactus fruits by the Indians. J. Hered. 20:400-414.

WEATHER BUREAU. 1951-1974. Climatological data for the United States by sec-
tions: annual summary. Vols. 57-80. U.S. Dept. Commerce, Washington, DC.

YEATON, R. I. and M. L. Copy. 1979. The distribution of cacti along environmental
gradients in the Sonoran and Mojave deserts. J. Ecol. 67:529-541.

(Received 17 Jun 1986; revision accepted 10 Apr 1987.)

REPRODUCTIVE BIOLOGY OF THE TREE
IPOMOEA WOLCOTTIANA (CONVOLVULACEAE)

STEPHEN H. BULLOCK and RICARDO AYALA
Estacion de Biologia Chamela,
Universidad Nacional Autonoma de México,
Apartado Postal 21,

San Patricio, Jalisco 48980, México

IRENE BAKER and HERBERT G. BAKER
Department of Botany, University of California,
Berkeley 94720

ABSTRACT

Ipomoea wolcottiana is an hermaphroditic tree of the tropical deciduous forest of
México. It flowers after leaf drop, but may be highly asynchronous among trees if
interrupted by rainfall early in the dry season. Anthesis is nocturnal and nectar
secretion is constant until after midday. Insect visitors include sphingid moths and
21 species of bees; few of these have characteristics of effective cross-pollinators. The
fragrant flowers present abundant pollen, but the sucrose-rich nectar differs between
trees by as much as a factor of three. Significant individual variation also occurs in
stamen and style length, and flower weight, but only the latter two are correlated.
Pollination experiments show self-incompatibility and some female-sterile trees; the
latter result is supported by records of up to four years from marked trees, and is
unrelated to floral morphology or nectar volume.

Variation between individuals in female fecundity is conspicuous
in some species of Jpomoea, but the causes have proven difficult to
unravel (Martin 1970, Stucky and Beckmann 1982). Although I[p-
omoea is principally a genus of vines, a few species are trees (Lujan
1974, McPherson 1982), a life form that differs substantially from
vines in the pattern of breeding systems (Bullock 1985). Tree species
of Ipomoea are of broader ecological interest because they are com-
mon pioneers in the seasonally arid tropics of México, and may have
an important role in supporting bee populations in a season when
few other trees are in flower (“keystone mutualists”’ of Gilbert 1980).
In this study, we outline the reproductive biology of J. wolcottiana
Rose, including phenology, nectar production and quality, flower
visitors, morphological variation in the flowers, compatibility, and
sterility.

STUDY AREA

The present study was carried out from 1980-1987 in lowland
Jalisco, México, at the Estacion de Biologia Chamela (19°30'N,

MApDRONO, Vol. 34, No. 4, pp. 304-314, 1987

1987] BULLOCK ET AL.: TPOMOEA REPRODUCTIVE BIOLOGY 305

105°03'W). The summer rainy season lasts about four months, but
rains between November and early February account for 11% of the
average annual precipitation, with a range of 0-30%; no measurable
rain has been recorded from March through late May (Bullock 1986).
The vegetation is tropical deciduous forest (Rzedowski 1978, Lott
et al. 1987). The flora of the field station includes 20 species of
Ipomoea (Lott 1985). Ipomoea wolcottiana is here a tree of 2-9 m
or more, and is scarce except in large areas of disturbance.

MATERIALS AND METHODS

Measurements were made of 10 flowers from each of 17 trees,
including length of the style (from the base of the free filament), and
of the longest and shortest stamen filaments, and flower weight.
Nectar volume was measured on various dates from flowers on 16
trees, with five flowers sampled per tree per hour (at 2330, 0300,
0600, 0900, and 1330 hr local time), although not all trees were
collected at all hours. Nectar analyses were made for six trees and
included methods described previously for sugars and amino acids
(Baker and Baker 1975, Baker and Baker 1982), as well as screening
for alkaloids (Dragendorff test), proteins (brom-phenol blue test),
and phenolics (p-nitro aniline test). Sugar concentration was mea-
sured from nectar spotted on filter paper.

Activity periods of bees were recorded for hourly periods on five
days in February and March 1985. The species were rated for abun-
dance and behavioral observations were made, which comple-
mented collections and observations made yearly since 1980. Bee
sizes were measured as the width of the scutum at the tegula, for
five females of each species (males in three species, see below).
Additional observations were made to detect visits by bats and
moths, and birds were noted during the day.

Compatibility tests followed the protocol of Bullock (1985), with
a total of 17 trees tested in 1982 and 1983. Fruit set was zero on
some of these trees, so a sample was marked for yearly observation
of fruiting (24 trees in 1982, 36 in 1983-1985). A qualitative rating
system with four levels (0-3) was used to accommodate the range
of variation of several orders of magnitude and the differences due
to canopy size.

Statistical analyses were one-way ANOVA, Spearman rank cor-
relations (r,), least-square linear regression (r”) and analysis of co-
variance (Killian 1981), the Student-Newman-Keuls procedure, and
the Mann-Whitney test (Zar 1974). Significance was accepted at the
0.05 level. Plant and animal specimens were deposited in the sta-
tion’s museum or at the Instituto de Biologia, U.N.A.M., in México
City.

306 MADRONO [Vol. 34

RESULTS

Phenology. The trees flowered in the early months of the dry
season, and most were leafless then. In April 1980, the population
contained trees with only flowers or fruits, and a few individuals
with both. This late and asynchronous reproduction may have been
due to rain in late January and early February (46 mm). In 1981,
following mid-January precipitation (124 mm), a similar phenology
was noted. With no rain from November 1981 through May 1982,
flowering occurred in January and February 1982. Leaf fall was well
advanced in November 1982, but heavy rains in late November and
December (160 mm) caused a major leaf flush (sudden, vigorous
growth). Flowering throughout the population was delayed until late
February to early March 1983. The next dry season began in No-
vember 1983, and flowering followed in January and February, with
rain in mid-January (38 mm) causing only a minor flush. The 1984—
1985 dry season started in October, but rain in mid-December (85
mm) and in mid-January (19 mm) delayed flowering. In 1986, flow-
ering was very sparse and fruit production zero, presumably due to
lack of rain in summer 1985 (60% of previous normal). The next
flowering season began in December 1986, and was not interrupted
by mid-January rain (25 mm). Near the station buildings, well-
watered trees retained some leaves year round, whereas unirrigated
trees nearby lost all their leaves in the dry season.

Anthesis. The corolla was furled in bud, and opened gradually
after midnight, until the distal half of the corolla was flared perpen-
dicular to the floral axis. The entrance to the effective corolla tube
was about 11-12 mm in diameter (Fig. 1). At anthesis, the anthers
were dehisced and nectar was present. The flowers closed perma-
nently the following afternoon.

Nectar. Nectar accumulated throughout the night and morning
until at least 1300 hr. The volume-time regressions were all signif-
icant and ranged from r? = 0.41-0.94. Rates of accumulation were
heterogeneous (analysis of covariance, F = 41), varying from 0.90-
1.98 ul/hr. The estimated volume prior to anthesis (2300 hr) differed
more than flow rates between trees, ranging from 0.6—16.2 ul. Be-
cause the nectar accumulation curves were not heterogeneous in
form, the trees can be compared most simply by the volume at a
particular hour. At 0900 hr, somewhat after the peak of pollinator
activity, accumulated volumes ranged from 11-34 ul (Table 1). The
variation was nearly continuous, with no significant differences be-
tween trees differing in rank by less than three (SNK test). No sig-
nificant trend emerged in relating flower weight to nectar volume
(r, = 0.51, Table 1). No significant correlations were found between
nectar volume and length of style or stamen (r, = —0.09, —0.10,
respectively). Furthermore, nectar production was not clearly related

1987] BULLOCK ET AL.: IPOMOEA REPRODUCTIVE BIOLOGY 307

Fic. 1. Flower of Ipomoea wolcottiana in nearly axial (a) and cross-sectional (b)
views (in situ, unfiltered daylight).

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1987] BULLOCK ET AL.: TIPOMOEA REPRODUCTIVE BIOLOGY 309

to fruit production from hand cross-pollination (r, = —0.20). The
volume in trees that are fertile by hand cross-pollination was not
different from that of sterile trees (Mann-Whitney test).

Amino acid concentration in the nectar was low (1-2 on the his-
tidine scale). The sugar component was mostly sucrose, with ratios
of sucrose to glucose plus fructose ranging from 2.0-3.4 (X = 2.42).
Sugar concentration was measured for four trees, giving the following
values: 25.8% (w/w), 29.6%, 27.7%, and 38.8%. None of the samples
showed detectable traces of alkaloids or proteins, but the reaction
for phenolics varied from not detectable to moderately strong. The
flowers also had a strong fragrance. Apparently glandular hairs were
present on the lower part of the filaments, but secretions have not
been obtained in analyzable quantity.

Flower visitors. Twenty-one species of bees were found feeding at
the flowers, but nine of these were rare (Table 2). Most of the species
were represented by females or workers, but only males were noted
for Eulaema polychroma, Melissodes tepaneca, and Melitoma mar-
ginella. The activity of the nocturnal Megalopta sp. remained poorly
defined due to their reaction to light. Ceratina capitosa spent much
time inactive inside the flowers, but this behavior may be limited
to males. The range of body widths was almost an order of magnitude
(Table 2) and included the largest and smallest bees in the Chamela
fauna. The larger species always brushed against the anthers, and
usually the stigma, on entering. All bee species foraged for nectar
by entering the corolla tube. All species can become dusted by fallen
pollen in the corolla tube, but only six species were noted as pollen
collectors, including four of the common species (Table 2).

Other diurnal visitors included Cacicus melanicterus (Bonaparte)
(Aves: Icteridae) and Amazilia rutila (DeLattre) (Aves: Trochilidae).
Nocturnal observations showed some visitation by Erinnyis sp.
(Lepidoptera: Sphingidae) and we have identified pollen collected
from E. ello (L.) as I. wolcottiana. No visits by bats have been
observed.

Style and stamen length. The length of the style and longest stamen
varied significantly between trees (Table 1; ANOVA, F = 39 and
35, respectively). The length of styles but not of stamens was cor-
related with flower weight (respectively, r, = 0.52, p < 0.05; r, =
—Q.02). Stamen and style lengths were not significantly correlated
(r, = 0.28), but showed similar ranges of variation, 8.0 and 7.8 mm,
respectively. Mean length of the shortest stamen varied significantly
(F = 21, range 16.5-24.8 mm) and correlated with length of the
longest stamen (r, = 0.58).

Compatibility and sterility. Of those trees that bore any fruit as a
result of pollination by hand, all except one were self-incompatible
(Table 1). Because all hand pollination was done between 0630 hr

310 MADRONO [Vol. 34

TABLE 2. BEE VISITORS TO FLOWERS OF [pomoea wolcottiana. Size is scutum width
in mm for females, or males (*). Activity period is time of day by hour intervals
(n.d. = species not observed in 1985). Notes are given for species observed collecting
pollen (p), and for those more than rare in abundance.

Family and species Size Activity period and notes

HALICTIDAE

Augochlora smaragdina Friese 1.9 9-13

Augochlora albiceps Friese 2.0 10-12 (p)

Augochlora nigrocyanea Cockerell 2A 8-16 (p, few)

Megalopta sp 2.6 20-7 (?, common)
ANTHOPHORIDAE

Ceratina capitosa F. Sm. 2.4 8-17

Ceratina sp 1 122 10-17

Ceratina sp 2 1.0 10-14

Centris nitida F. Sm. 4.6 n.d.

Centris segregata Crawford 5.4 n.d.

Melissodes tepaneca Cresson 2312 9-15 (common)

Melitoma marginella (Cresson) lis 8-15 (common)

Xylocopa fimbriata Fabricius 8.8 10-11

Xylocopa mexicanorum F. Sm. 6.9 6-12 (p, common)

Xylocopa muscaria (Fabricius) D2 11-12

Xylocopa t. tabaniformis (F. Sm.) 5.8 6-10 (p, common)
APIDAE

Eulaema polychroma (Mocsary) 6.4* 10-11

Melipona beecheii Bennett 3:3 6-11 (few)

Trigona buyssoni Friese 0.9 9-13 (few)

Trigona fulviventris Guerin 1.6 8-12 (few)

Trigona hellwegeri Friese 1.8 n.d. (few)

Trigona orizabaensis Strand 1.7 6-17 (p, common)

and 1300 hr, there was no evidence for the breakdown of incom-
patibility barriers with flower age. No fruit were produced from hand
pollination on seven of 17 trees tested, and 10 of 17 had fruit set of
10% or less. Fruit set was not significantly correlated with either
style or stamen length or flower weight (Table 1; respectively, r, =
0.03, 0.31, and —0.09).

The exceptionally low fertility of I. wolcottiana compared with
other tree species at Chamela (Bullock 1985) led to further obser-
vations in the population at large. Some trees of J. wolcottiana pro-
duced massive numbers of flowers, but produced few fruits or were
completely barren, whereas adjacent trees, sometimes with inter-
laced canopies, produced many fruits. The differences between fruit
production in trees with low and high fecundity were consistent.
Data from four consecutive years showed no tendency to oscillate
between barren and productive states, and the majority of trees of
low or moderate average fecundity had no peak year (Table 3).

1987] BULLOCK ET AL.: TPOMOEA REPRODUCTIVE BIOLOGY 311

TABLE 3. MAXIMUM FRUIT PRODUCTION AND FREQUENCY OF (FRUIT-)BARREN YEARS
FOR INDIVIDUALS OF [pomoea wolcottiana WITH DIFFERENT AVERAGE FRUIT PRO-
DUCTION.

Overall
frequency
EEO Percent of trees attaining a given ee
1982 (or 83)- maximum level of fruiting ee ee
1985 n 0 1 2 3 fruit
0-0.49 8 88 12 0 0 97
0.50—1.39 8 — 62 38 0 10
1.40-—2.19 7 — — 57 43 0)
2.20-3.0 13 — — — 100 4
DISCUSSION

The annual phenology of J. wolcottiana varies considerably and
apparently is conditioned by the timing of rainfall. Thus, flowering
may begin in early December or not until late February, or may split
between early and late starting trees, and some individuals may
flower twice in one season. Drought appears necessary for the onset
of flowering, but rains in December or January can cause flushing
in trees not well advanced in flower development. Desynchronized
or delayed flowering has occurred in four years from 1980-1987.
On a local spatial scale, soil and vegetation conditions affect the
timing of drought experienced by the trees, which probably increases
asynchrony in the population. Whatever the result for plant fitness,
asynchronous and delayed flowering may benefit flower visitors by
prolonging the availability of nectar. Also, the prolonged presence
of immature fruit may benefit pre-dispersal seed predators (Schlising
1980, Augspurger 1981), which include Megacerus cubicus (Mot-
schulsky) (Bruchidae), and unidentified species of Curculionidae,
Diptera, and Lepidoptera. Other trees flowering during the dry sea-
son at Chamela are largely unresponsive to rain in those months.

The population is not limited to pollination by either nocturnal
or diurnal animals. Nectar flow continues from late night anthesis
to midday wilting. Also, the sugar and amino acid analyses are
consistent with those from many species pollinated by sphingid moths
or large bees (Baker and Baker 1982). Despite the observed diversity
of visitors, most of the bees do not reliably contact the anthers or
stigma on account of body size or behavior. Altogether, they must
remove considerable nectar. Xylocopa mexicanorum and_ X. taban-
iformis are the only bee species that pollinate J. wolcottiana consis-
tently, are common, and move frequently between trees. Ceratina
capitosa also might be a significant pollinator, as are its congeners
on flowers of other Ipomoea species. Moreover, the importance of

312 MADRONO [Vol. 34

nocturnal visitors remains to be clarified, especially for Sphingidae
and Megalopta. Bats were not observed visiting J. wolcottiana, and
the nectar sugar composition is contradictory to the trend in species
pollinated by microchiroptera (Baker and Baker 1982). However,
abundant pollen of Jpomoea sp. was found in stomachs of the bats
Leptonycteris yerbabuenae Martinez and Villa and Glossophaga
soricina Pallas in Guerrero (Quiroz et al. 1986).

Nectar quantity differs among individuals, but we presently have
no reason to interpret variation around the linear regressions of
quantity on time as representing time-varying secretion rate. The
latter has been shown for a few trees and was suggested as a mech-
anism to induce cross-pollination (Frankie and Haber 1983).

Variation within and between individuals in style and stamen
length has been reported in few Jpomoea species (Wilson 1977,
Ennos 1981) despite many studies of pollination. In I. wolcottiana,
the difference between longest and shortest stamens is always sub-
stantial (3-8.2 mm), although its distribution was not normal or
unimodal. Greater anther-stigma distance entailed an order of mag-
nitude greater outcrossing frequency in J. purpurea compared with
I. hederacea (Ennos 1981). Also, seed set from autogamy in J. pur-
purea was negatively correlated with anther-stigma distance (Ennos
1981). The style-filament difference in J. wolcottiana ranged from
—3.8 to 5.1 mm, but was not normal or unimodal, and was not
different between trees that were fertile or barren in hand cross-
pollination (Mann-Whitney test).

Thus, Ipomoea wolcottiana varies significantly in morphological
and functional characters that are not sorted into well-defined groups
or correlated in interpretable patterns. If the system is evolving, the
path is unclear as yet. As a further element in the breeding system,
variation in male fertility has not yet been detected. A limited search
for male sterility, using cotton blue in lactophenol (in vitro germi-
nation of Jpomoea pollen is problematic; Martin and Ortiz 1966,
Stucky and Beckmann 1982), did not show any notable differences
among trees. A similar condition was described for Mirabilis froebelii
(Behr) Greene (Nyctaginaceae; Baker 1964), where variability in the
floral organs had no apparent relationship to the breeding system.

In fruit production, most variation was among trees of interme-
diate to low average fecundity, and non-fruiting trees were consis-
tently barren. Thus, this J. wolcottiana population does not consist
of individuals that fruit heavily on a supra-annual basis with inter-
vening years of little or no fruit (Janzen 1978). The proportion of
trees with different bearing levels is unknown, because the sample
population was not taken at random. Barrenness might be due to
nutrient limitation, lack of pollination, lack of compatible pollen,
genetic sterility, or cytogenetic problems. The former two alterna-
tives are not probable because trees with different levels of fecundity

1987] BULLOCK ET AL.: TPOMOEA REPRODUCTIVE BIOLOGY 313

(by natural or hand pollination) were frequently commingled.The
consistent fruiting behavior or barrenness and the failure of polli-
nation by hand to overcome barrenness of some trees lead us to
conclude that some form(s) of female sterility exists in this popu-
lation. As noted above, this is not correlated with a suite of floral
characters. Female sterility is known in other Ipomoea species (Mar-
tin and Cabanillas 1966, Stucky and Beckmann 1982). Chromo-
somal aberrations and imbalance add to the problems in J. batatas
(L.) Lam. (Ting et al. 1957). Thus, the sterility problem may require
cytological study.

Female fecundity of some vine species is limited by the scarcity
of compatible mates (Martin 1968, Stucky and Beckmann 1982).
The potential for incompatible pollination is suggested by the results
of extensive tests with the diploid J. setifera Poir., which revealed
10 incompatibility groups (Martin 1968). In vines, extensive clonal
growth (Penalosa 1984) or establishment from vegetative fragments
may result in patches containing one or very few incompatibility
groups. When plants establish only from seed, as in I. wolcottiana,
lack of compatible mates seems unlikely to account for a wide range
of fecundities. However, when trees are partially or completely self-
incompatible, such a situation may arise. For example, in some
sparse populations of Jnga species (Mimosoideae) fruit set was lim-
ited by the low frequency of more distant and more successful crosses
(Koptur 1984). The effects of population structure and its variation
in the disturbance-following J. wolcottiana remain to be assessed in
relation to both the generalized and individually variable characters
of the breeding system we have described.

ACKNOWLEDGMENTS

Useful comments on earlier drafts were made by T. Atkinson, S. Barrett, S. Koptur,
E. Lott, C. Martinez del Rio, and R. Meinke. The plants were determined by E. Lott,
and insects by R. Ayala, R. Snelling and H. Daly (bees), A. Pescador (sphingids), and
J. Kingsolver (bruchids).

LITERATURE CITED

AUGSPURGER, C.K. 1981. Reproductive synchrony ofa tropical shrub: experimental
studies on effects of pollinators and seed predators on Hybanthus prunifolius
(Violaceae). Ecology 62:775-789.

BAKER, H.G. 1964. Variation in style length in relation to outbreeding in Mirabilis
(Nyctaginaceae). Evolution 18:507—5 12.

and I. BAKER. 1975. Studies of nectar constitution and pollinator-plant

coevolution. Jn L. E. Gilbert and P. H. Raven, eds., Coevolution of animals and

plants, p. 100-140. Univ. Texas Press, Austin.

and 1982. Floral nectar sugar constituents in relation to pollinator
type. In C. E. Jones and R. J. Little, eds., Handbook of experimental pollination
biology, p. 118-141. Van Nostrand Reinhold, New York.

BuLLock, S. H. 1985. Breeding systems in the flora of a tropical deciduous forest
in México. Biotropica 17:287-301.

314 MADRONO [Vol. 34

1986. Climate of Chamela, Jalisco, and trends in the south coastal region
of México. Arch. Met. Geoph. Biocl. Ser. B 36:297-316.

Ennos, R. A. 1981. Quantitative studies of the mating systems in two sympatric
species of Ipomoea (Convolvulaceae). Genetica 57:93-98.

FRANKIE, G. W. and W. A. HABER. 1983. Why bees move among mass-flowering
Neotropical trees. Jn C. E. Jones and R. J. Little, eds., Handbook of experimental
pollination biology, p. 360-372. Van Nostrand Reinhold, New York.

GILBERT, L. E. 1980. Food web organization and conservation of Neotropical di-
versity. Jn M. E. Soulé and B. A. Wilcox, eds., Conservation biology: an evo-
lutionary-ecological perspective, p. 11—33. Sinauer Assoc., Sunderland.

JANZEN, D. H. 1978. Seeding patterns of tropical trees. Jn P. B. Tomlinson and M.
H. Zimmermann, eds., Tropical trees as living systems, p. 615-638. Cambridge
Univ. Press, Cambridge.

KILLIAN, K.C. 1981. Statistics with DAISY. Rainbow Computing Inc., Northridge.

Koptur,S. 1984. Outcrossing and pollinator limitation of fruit set: breeding systems
of Neotropical /nga trees (Fabaceae: Mimosoideae). Evolution 38:1130-1143.

Lott, E. J. 1985. Listados floristicos de México. III. La Estacion de Biologia Cha-
mela, Jalisco. Inst. Biologia, Univ. Nac. Auton. Méx., México.

, S. H. BULLOCK, and J. A. SOLIS-MAGALLANES. 1987. Floristic diversity and
structure of upland and arroyo forests in lowland Jalisco. Biotropica 19:228-
239.

LuJAN, O. E. 1974. Anomalous secondary growth in stems of the arborescent [po-
moeas (Convolvulaceae). Master’s thesis, California State Univ., Los Angeles.

McPHERSON, G. 1982. Studies in Ipomoea (Convolvulaceae). I. The arborescens
group. Ann. Missouri Bot. Gard. 68:527-545.

MarTINn, F. W. 1968. The system of incompatibility in Ipomoea. J. Heredity 59:
263-267.

1970. Self and interspecific incompatibility in the Convolvulaceae. Bot.

Gaz. (Crawfordsville) 131:139-144.

and E. CABANILLAS. 1966. Post pollen-germination barriers to seed set in

the sweet potato. Euphytica 15:404-411.

and S. Ortiz. 1966. Germination of sweet potato pollen in relation to
incompatibility and sterility. Proc. Am. Soc. Hort. Sci. 88:491-497.

PENALOSA, J. 1984. Basal branching and vegetative spread in two tropical rain forest
lianas. Biotropica 16:1-9.

Quiroz G., D. L., M. S. XELHUANTzI L., and M. C. ZAMORA M. 1986. Analisis
palinologico del contenido gastrointestinal de los murciélagos Glossophaga sor-
icina y Leptonycteris yerbabuenae de las Grutas de Juxtlahuaca, Guerrero. Inst.
Nac. Antropol. Hist., México.

RZEDOWSKI, J. 1978. Vegetacion de México. Editorial Limusa, México.

SCHLISING, R. A. 1980. Seed destruction of California morning glory (Convolvu-
laceae, Calystegia) by bruchid beetles. Madrono 27:1-16.

Stucky, J. M. and R. L. BECKMANN. 1982. Pollination biology, self-incompatibility
and sterility in Jpomoea pandurata (L.) G. F. W. Meyer (Convolvulaceae). Amer.
J. Bot. 69:1022-1031.

Tina, Y. C., A. E. KeHrR, and J.C. MULLER. 1957. A cytological study of the potato
plant Ipomoea batatas (L.) Lam. and its related species. Amer. Nat. 91:197-203.

Witson, D. E. 1977. Ecological observations of the tropical strand plants Ipomoea
pes-caprae (L.) R. Br. (Convolvulaceae) and Canavalia martima (Aubl.) Thou.
(Fabaceae). Brenesia 10/11:31-42.

ZAR, J. H. 1974. Biostatistical analysis. Prentice-Hall, Inc., Englewood Cliffs.

(Received 7 Jul 1986; revision accepted 8 May 1987.)

ALPINE ANNUAL PLANT SPECIES IN THE
WHITE MOUNTAINS OF EASTERN CALIFORNIA

TIMOTHY P. SPIRA!
White Mountain Research Station, University of California,
3000 East Line St., Bishop 93514

ABSTRACT

The White Mountains resemble the Sierra Nevada in having an unusually high
concentration of alpine annual plant species. Thirteen species of annuals comprise
8% of the alpine flora of the White Mountains. Most species (69%) have distributions
that extend throughout western North America; however, they are known to occur
above treeline only in the White Mountains and Sierra Nevada. Two of 13 species
of alpine annuals in the White Mountains have distributions from near sea level to
the alpine, three occur from the desert to the alpine, and seven have distributions
from the coniferous forest to the alpine in California. Three additional species (Che-
nopodium rubrum, Gentiana tenella, and G. prostrata), generally known as annuals,
function as biennials in alpine habitats of the White Mountains.

Few species of annual plants occur in alpine environments (Bil-
lings and Mooney 1968, Bliss 1971, Billings 1974). This probably
reflects the inability of most annuals to successfully complete their
life cycle in a short, cold growing season. The Sierra Nevada of
California, however, has an unusually large number of annual plants
at high elevations. Sharsmith (1940), for example, lists about 10
annuals in the alpine zone, and Went (1953) and Jackson (1985)
describe approximately 47 annuals in the high subalpine and alpine
zones.

The ability of annuals to occupy alpine habitats in the Sierra
Nevada may be caused by relatively high levels of solar radiation
and more moderate conditions (due to fewer summer storms) than
are generally found in other alpine areas (Chabot and Billings
1972, Jackson 1985). An abundance of annual plant species at lower
elevations, some of which may have been pre-adapted to conditions
at higher elevations, contribute to the increased number of alpine
annuals in the Sierra Nevada (Went 1953, Chabot and Billings 1972,
Jackson 1985).

The alpine zone of the White Mountains 1s similar to the Sierra
Nevada in having relatively warm dry summers with high levels of
solar radiation and large numbers of annuals at lower elevations
(Lloyd and Mitchell 1973, Major and Taylor 1977). On this basis,

' Present address: Department of Biology, Georgia Southern College, Statesboro
30460.

MADRONO, Vol. 34, No. 4, pp. 315-323, 1987

316 MADRONO [Vol. 34

a relatively high concentration of annuals should occur at high el-
evations in the White Mountains. This study tests this prediction
and provides information on the distribution of alpine annual plant
species in the White Mountains of eastern California.

STUDY AREA

The White Mountains are located in the eastern part of central
California (37°13'’-38°N and 117°55’—118°25'W), with the extreme
northeastern portion extending into Nevada. The range is approx-
imately 90 km long and 32 km wide at its widest point and is about
2278 km? in area. Because the Sierra Nevada occurs immediately
to the west, the White Mountains are in a pronounced rain shadow
and the vegetation is Great Basin in character (Lloyd and Mitchell
1973, Major and Taylor 1977). The four main plant zones described
in Lloyd and Mitchell (1973) are desert scrub (1220-1980 m), pinyon
woodland (1980-2895 m), subalpine forest (2895-3505 m), and al-
pine tundra (3505-4340 m). Within these vegetation zones, they list
811 taxa of vascular plants of which 20% (160 species) occur in the
alpine zone. Recent collections have increased the total number of
known species in the White Mountains to 988 (Morefield 1986).

Climatic records from 1953 through 1973 at the Barcroft Labo-
ratory (elevation 3801 m) of the White Mountain Research Station
indicate a mean July temperature of 7.4°C, a mean January tem-
perature of —9.1°C, a mean annual temperature of —5.8°C, and a
mean annual precipitation of 49.6 cm, of which 18.3% (9.1 cm) falls
during June, July, and August (Pace et al. 1974).

METHODS

The number and proportion of annual plant species in each of the
four major plant zones of the White Mountains were based on species
distributions given in Lloyd and Mitchell (1973). Life cycles for
species not included in this flora were obtained from other floras
(primarily Munz 1968, Hitchcock et al. 1969). Geographic affinities
were categorized as cosmopolitan, western North American (widely
distributed at low and high elevations from the Pacific Coast to the
Rocky Mountains), and endemic (restricted to the White Mountains
and nearby Sierra Nevada and Sweetwater ranges) according to de-
scriptions in Munz (1968), Hitchcock et al. (1969), and Jackson
(1985). Elevational distributions were based on Lloyd and Mitchell
(1973) and Spira (pers. obs.) for the White Mountains and Munz
(1968) for California. Field observations by the author were confined
to the southern part of the range (south of White Mountain Peak).

Voucher specimens were deposited at JEPS. Nomenclature follows
Lloyd and Mitchell (1973); for species not included in this source,
nomenclature follows Munz (1968).

1987] SPIRA: ALPINE ANNUAL PLANTS Su)

RESULTS AND DISCUSSION

The number and proportion of annual plant species in each of the
major plant zones of the White Mountains are shown in Fig. 1.
Annuals become progressively less common with increasing eleva-
tion. For example, an analysis of species distributions as listed in
Lloyd and Mitchell (1973) indicates that the desert scrub (the lowest
zone) has 148 annuals (comprising 35% of the desert flora), whereas
the alpine zone has only five annuals (comprising 3% of the alpine
flora).

Field observations by the author from 1980-1985 revealed eight
additional annuals in the alpine zone (Table 1). This increases the
known number of alpine annuals to 13 species (8% of the alpine
flora) in the White Mountains. Because annuals are easily overlooked
due to their small size and short life cycle, further field studies would
undoubtedly reveal additional species in each plant zone. Although
the data shown in Fig. 1 underestimate the total number of annual
plant species, they do indicate the relative number of annuals in
each of the four major plant zones.

The 13 species of alpine annuals in the White Mountains represent
11 genera in 10 families (Table 1). This list is conservative in that
it does not include species having an annual and/or biennial life
cycle (discussed later) or species described as annuals to perennials
(e.g., Calyptridium umbellatum var. caudiferum, Androsace septen-
trionalis ssp. subumbellata). Also, because my field observations
were limited to alpine areas in the southern part of the range (south
of White Mountain Peak), additional alpine annuals may occur in
the northern part of the range.

Geographical distribution. Alpine annuals in the White Mountains
generally have wide geographical distributions (Table 1). Nine of 13
species (69%) have a western North American distribution, two
(15%) are weedy annuals with a cosmopolitan distribution, and only
two species (15%) are endemic to the White Mountains and nearby
Sierra Nevada-Sweetwater ranges.

None of the nine alpine annuals with a western North American
distribution are known to occur in the alpine zone outside of the
White Mountains and Sierra Nevada. The apparent ability of these
nine species to occur at higher elevations in the White Mountains
and Sierra Nevada relative to other mountain ranges could be ex-
plained by a warmer, drier growing season. In lower radiant energy
areas, there may not be sufficient time for annuals to complete their
life cycle and form viable seed, particularly in unusually short grow-
ing seasons (Jackson 1985).

Jackson (1985) lists the proportion of annuals at or above treeline
in a number of western North American mountain ranges. Of the
19 areas sampled, seven (37%) had less than 1% annuals, nine (47%)

318 MADRONO [Vol. 34

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2 135 90 i
120 BOz
uJ
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DESERT PINYON SUB- ALPINE
SCRUB WD. ALPINE
Fic. 1. Number of annual plant species and percent annuals within four main
plant zones of the White Mountains based on Lloyd and Mitchell (1973). Recent

field observations by author have increased the known number of alpine annuals in
the White Mountains from 5 to 13 species (8% of alpine flora).

had 1—4% annuals, and only three (16%) had more than 4% annuals
at or above treeline. The Wassuk Range, NV (4.3% annuals),
Charleston Mountains, NV (7.6% annuals), and Hall Natural Area,
Sierra Nevada, CA (8.3% annuals) comprise the latter group. The
Wassuk Range and Charleston Mountains were based on samples
of only 70 and 39 species, respectively. Consequently, more com-
prehensive surveys need to be made prior to evaluating these two
areas.

The proportion of annuals in the alpine flora of the White Moun-
tains was comparable to the Hall Natural Area of the Sierra Nevada,
the area with the highest proportion of alpine annuals in Jackson’s
(1985) survey of western North American mountain ranges. Al-
though annuals comprised about 8% of the alpine flora in both
ranges, the number of alpine annuals in the Sierra Nevada was much
greater than in the White Mountains (47 vs. 13 species). In addition
to a larger pool of species, the Sierra Nevada survey included annuals
in both the high subalpine and alpine zones (Jackson 1985), whereas
only annuals known to occur above treeline were included in the
present survey.

1987] SPIRA: ALPINE ANNUAL PLANTS 319

TABLE |. SPECIES, FAMILY, AND GEOGRAPHIC AFFINITY OF ALPINE ANNUAL PLANT
SPECIES IN THE WHITE MOUNTAINS. ! = Species listed in alpine zone of the White
Mountains by Lloyd and Mitchell (1973); 2 = Restricted to White Mountains and
nearby Sierra Nevada and Sweetwater ranges; > = Extends into eastern North America

as a weed.

Species Family Geographic affinity
Cryptantha glomeriflora* Boraginaceae Endemic
Mimulus coccineus? Scrophulariaceae Endemic
M. suksdorfii' Scrophulariaceae w. North American
Nama densum' Hydrophyllaceae w. North American
Juncus bryoides Juncaceae w. North American
Gayophytum racemosum Onagraceae w. North American
Gymnosteris parvula' Polemoniaceae w. North American
Eriogonum cernuum! Polygonaceae w. North American
Calyptridium roseum' Portulacaceae w. North American
Chenopodium atrovirens Chenopodiaceae w. North American
C. leptophyllum? Chenopodiaceae w. North American
Monolepis nuttalliana Chenopodiaceae Cosmopolitan
Capsella bursa-pastoris Brassicaceae Cosmopolitan

Elevational distribution. Upper elevational extremes for alpine
annuals in the White Mountains (Table 2) were consistently higher
than those listed by Munz (1968) for California. Because elevational
distributions in large floras are not always reliable, it is unclear
whether or not alpine annuals in the White Mountains occur at
higher elevations than elsewhere in California.

In the White Mountains, however, alpine annuals generally have
broad elevational distributions (Table 2). Ten of the 13 species (77%)
occur at elevations ranging from less than 1850 m to more than
3500 m. Within California, two of the 13 species (Capsella bursa-
pastoris and Monolepis nuttalliana, 15%) have distributions from
near sea level to the alpine, three species (23%) occur from the
sagebrush (desert) scrub to the alpine, and seven species (54%) have
distributions from the coniferous forest to the alpine (Table 2).

Jackson (1985) found that alpine annuals in the Sierra Nevada
also have broad elevational ranges and suggests they migrated to
higher elevations from lower elevation populations. Went (1948)
and Chabot and Billings (1972) also suggest a low elevation origin
for Sierran alpine annuals and note that conditions in the alpine in
July and August are similar to those at lower elevations in March
and April (e.g., high light levels, large diurnal temperature fluctua-
tions, and limited moisture availability). Consequently, lower ele-
vation annuals may have been pre-adapted, at least to some extent,
to an alpine environment.

Axelrod (1981) suggests that a warmer, drier climate in the xe-

[Vol. 34

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1987] SPIRA: ALPINE ANNUAL PLANTS 5711

rothermic (ca. 83000—4000 yr B.P.) facilitated species migrations from
lower to higher elevations in the Sierra Nevada (and presumably in
the White Mountains). Thus, a number of annuals may have mi-
grated successfully into the alpine zone at this time. As temperatures
dropped and rainfall increased following the xerothermic, some an-
nuals probably were eliminated, whereas others may have persisted
in favorable microsites at high elevations.

Went (1948) and Chabot and Billings (1972) suggest that nearby
deserts were the primary source of alpine annuals in the Sierra Ne-
vada. In a recent floristic study, Jackson (1985) suggests that mon-
tane coniferous forests were probably the primary source of alpine
annuals in the Sierra Nevada. Because 23% of alpine annuals in the
White Mountains extend from the sagebrush (desert) scrub to the
alpine and 54% extend from the coniferous forest to the alpine, both
the desert and forest were probably important source areas for alpine
annuals in the White Mountains.

Habitats. The alpine annuals observed in this study were almost
always found on bare soil, and often on dry south-facing slopes. The
high light levels, comparatively warm temperatures, and reduced
competition for soil moisture in such habitats enhance the ability
of annuals to successfully complete their life cycle in a short growing
season (Jackson and Bliss 1982).

Alpine annuals were observed frequently along roadsides and
grazed areas in the White Mountains, which suggests humans and
domestic animals (e.g., sheep and cattle) have increased available
habitat and may have introduced seeds of some annuals into the
alpine zone. This was probably the case for six weedy annuals re-
stricted to a disturbed roadside area immediately south of the en-
trance gate to the Barcroft Laboratory, where cars occasionally parked
and where horses were tethered during the fall hunting season. Be-
cause populations of five of these species (Sisymbrium irio, S. ori-
entale, Descurainia sophia, Stellaria media, and Senecio vulgaris)
were small (generally fewer than 10 plants), highly localized, and
present in only one or at most two of the six years (1980-1985)
observations were made, they were not included in my list of alpine
annuals in the White Mountains.

Capsella bursa-pastoris was a sixth weedy annual restricted to the
same roadside area. Unlike the other five species, however, a stable
population of 25 to several hundred individuals was maintained
during five successive years, and the area occupied by this species
increased during the period of study. Thus, C. bursa-pastoris appears
to have successfully colonized the alpine zone; but it and several
other weedy alpine annuals (e.g., Monolepis nuttalliana, Chenopo-
dium atrovirens, and C. leptophyllum) probably would not be pres-
ent, or would be much less common in the alpine zone of the White
Mountains, were it not for the presence of disturbed areas.

322 MADRONO [Vol. 34

Life cycle variation. Except for Nama densum and Eriogonum
cernuum, I observed each of the 13 alpine annuals described here
under field conditions. As expected, individuals of each species ger-
minated, flowered, fruited, and died within a single summer. In
contrast, several other species, generally described as annuals, ex-
hibited a biennial life cycle in alpine habitats of the White Moun-
tains. For example, Gentiana tenella is described as an annual and
G. prostrata as an annual and/or biennial in regional floras (e.g.,
Abrams and Ferris 1960, Munz 1968, Hitchcock et al. 1969). In the
White Mountains, however, individuals of both species consistently
form a vegetative rosette during their first summer, overwinter as a
taproot, and then flower, fruit, and senesce during their second sum-
mer. Unlike a number of purported biennials in which plant size
(rather than age) determines when flowering occurs (Gross 1981),
individuals of G. tenella and G. prostrata flowered in their second
year regardless of plant size (Spira 1983).

Chenopodium rubrum is known as an annual that occurs at ele-
vations to 1000 m in Britain, 2000 m in the European Alps, and
3000 m in the United States (Williams 1969). In the White Moun-
tains, however, I have observed C. rubrum in alpine habitats to 3750
m, where individuals grow vegetatively one summer and then flower,
fruit, and die during their second summer.

A biennial rather than annual life cycle in alpine individuals of
C. rubrum may be influenced by several factors. First year plants
may fail to reach some critical size before flowering can be induced,
individuals may require a cold treatment (as overwintering plants
would receive) prior to flowering, or individuals may not receive
the necessary photoperiod to induce flowering during their first sum-
mer’s growth (Harper 1977). It would be interesting to know whether
the shift in life cycle in C. rubrum is a genotypic response (1.e., life
cycle ecotypes over an elevational gradient) or an environmentally
controlled (phenotypic) response, and at what elevation the shift
from an annual to a biennial life cycle occurs.

ACKNOWLEDGMENTS

I thank Jack Major, James Morefield, Oren Pollak, and Lisa Wagner for reviewing
the manuscript; Oren Pollak and Hannah Carey for locating alpine populations of
Juncus bryoides and Mimulus coccineus; Larry Heckard and Jim Hickman for veri-
fying species names; and the staffof the White Mountain Research Station for logistical
support. Financial support was provided by the University of California, Berkeley,
the White Mountain Research Station, and Sigma Xi.

LITERATURE CITED

ABRAMS, L. and R. Ferris. 1960. An illustrated flora of the Pacific States, Wash-
ington, Oregon and California. Stanford Univ. Press, Palo Alto, CA.

AXELROD, D. I. 1981. Holocene climatic changes in relation to vegetation disjunc-
tion and speciation. Am. Nat. 117:847-870.

1987] SPIRA: ALPINE ANNUAL PLANTS 323

BILLINGS, W. D. 1974. Adaptations and origins of alpine plants. Arc. and Alp. Res.
6:129-142.

and H. A. Mooney. 1968. The ecology of arctic and alpine plants. Biol.
Rev. 43:481-529.

Buiss, L. C. 1971. Arctic and alpine life cycles. Ann. Rev. Ecol. Syst. 2:405-438.

CHABOT, B. F. and W. D. BILLINGS. 1972. Origins and ecology of the Sierran alpine
flora and vegetation. Ecol. Monogr. 42:163-199.

Gross, K. L. 1981. Predictions of fate from rosette size in four “biennial” plant
species: Verbascum thapsus, Oenothera biennis, Daucus carota, and Tragopogon
dubius. Oecologia (Berlin) 48:209-213.

HARPER, J. L. 1977. Population biology of plants. Academic Press, San Francisco.
Hitcucock, C. L., A. CRONQUIST, M. OWNBEY, and J. W. THOMPSON. 1969. Vascular
plants of the Pacific Northwest. Vols. 1-5. Univ. Washington Press, Seattle.
JACKSON, L. E. 1985. Floristic analysis of the distribution of ephemeral plants in
treeline areas of the western United States. Arc. and Alp. Res. 17:251-—260.
and L. C. Buiss. 1982. Distribution of ephemeral herbaceous plants near

treeline in the Sierra Nevada, California, U.S.A. Arc. and Alp. Res. 14:33-42.

LLoyp, R. M.and R.S. MITCHELL. 1973. A flora of the White Mountains, California
and Nevada. Univ. California Press, Berkeley.

Mayor, J. and D. W. TAyLor. 1977. Alpine, Jn M. G. Barbour and J. Major, eds.,
Terrestrial vegetation of California, p. 601-665. Wiley and Sons, New York.

MOREFIELD, J. D. 1986. Current status of the White Mountain flora. Jn C. A. Hall,
Jr. and D. J. Young, eds., Natural history of the White-Inyo Range, eastern
California and western Nevada and high altitude physiology. White Mountain
Research Station Symposium 1:51-57. Univ. California, Los Angeles.

Munz, P. A. 1968. A California flora and supplement. Univ. California Press,
Berkeley.

Pace, N., D. W. KIEPERT, and E. M. Nissen. 1974. Climatological data summary
for the Crooked Creek Laboratory, 1949-1973 and the Barcroft Laboratory,
1953-1973. Univ. California, White Mountain Research Station, Special Pub-
lication, 4th Ed.

SHARSMITH, C. W. 1940. A contribution to the history of the alpine flora of the
Sierra Nevada. Ph.D. dissertation, Univ. California, Berkeley.

SpirA, T. P. 1983. Reproductive and demographic characteristics of alpine biennial
and perennial gentians (Gentiana spp.) in the White Mountains, California. Ph.D.
dissertation, Univ. California, Berkeley.

WENT, F. W. 1948. Some parallels between desert and alpine flora in California.
Madrono 9:241-249.

—. 1953. Annual plants at high altitudes in the Sierra Nevada, California.
Madrono 12:109-114.

WILLIAMS, J. T. 1969. Biological flora of the British Isles: Chenopodium rubrum L.
J. Ecol. 57:831-841.

(Received 28 Jul 1986; revision accepted 10 Jun 1987.)

PROSOPIS (MIMOSACEAE) IN THE
SAN JOAQUIN VALLEY, CALIFORNIA:
VANISHING RELICT OR RECENT INVADER?

DAN C. HOLLAND!
Department of Biology,
California State University, Fresno,
Fresno 93740-0001

ABSTRACT

The presence of at least two species of mesquite (Prosopis glandulosa and P. pu-
bescens) in the San Joaquin Valley of California has been explained previously as the
result of the invasion of Mojavean floral elements during the Xerothermal period
some 8000-5000 years B.P. I propose that a number of lines of negative evidence
argue for the establishment and spread of both species within approximately the last
120 years.

The mesquites (Mimosaceae: Prosopis) are a group of woody le-
gumes restricted to the New World. Prosopis contains 10 species
that range from Argentina and Chile north to the west-central United
States. Most species are large shrubs or trees, some of which exceed
12 m. In subtropical regions, Prosopis species are often physiognomic
dominants or codominants that cover extensive areas.

Two of the four species of mesquite that occur in California are
native. Prosopis pubescens Benth. (Screwbean or Tornillo) primarily
inhabits washes and bajadas in the southern Mojave and Sonoran
deserts in California, and ranges widely throughout the southwestern
United States and Mexico. Prosopis glandulosa L. Benson (Honey-
bean mesquite) also ranges widely in the southwest and in the deserts
of California. It is more widespread than P. pubescens in its habitat
preferences, however, and occurs in xeric grasslands, on the fringes
of lake beds, and in flood plains, washes, and other riparian areas.
Prosopis velutina Woot. (Velvet mesquite) 1s native to the Arizona-
Sonora region, and also has a scattered distribution in California,
which suggests that it is naturalized in this area. Benson (1941) and
Munz (1959) have considered P. velutina to be a variety of P. glan-
dulosa. Prosopis strombulifera Lam. (Benth.) is native to South
America and is established near Bard, Imperial Co. (Munz 1959).

The first three species noted have disjunct distributions in Cali-
fornia, with populations present in the San Joaquin Valley (SJV)

' Present address: Department of Biology, University of Southwestern Louisiana,
Lafayette 70504-2451.

MADRONO, Vol. 34, No. 4, pp. 324-333, 1987

1987] HOLLAND: PROSOPIS, RELICT OR INVADER? 325

that are some distance from the main body of their range in the
state. In this disjunct area, P. glandulosa occurs primarily along the
floodplain of the north fork of the Kern River, between Bakersfield
and the former bed of Buena Vista Lake. Isolated populations occur
throughout the valley and adjacent foothills north to Alameda Co.
and east to the Sierra Nevada as far north as Fresno Co. (Fig. 1).
Benson (1941) reported an unvouchered occurrence of P. glandulosa
in the Cuyama River Valley, San Luis Obispo Co. Prosopis pubescens
is known only from Warthan and Los Gatos canyons in western
Fresno Co. and San Emigdio Canyon in southwestern Kern Co. Hilu
et al. (1982) reported P. velutina from the vicinity of Bakersfield,
where it occurs with P. glandulosa.

Barbour and Major (1980) concluded that P. glandulosa became
established in the SJV during the climatic warming trends of the
Xerothermal period, between 8000-5000 yr B.P. At this time and
during the Pleistocene, elements of the Mojavean biota presumably
invaded the Central Valley. Such organisms included Sceloporus
magister Hallowell (desert spiny lizard), Xantusia vigilis Baird (des-
ert night lizard), Gopherus agassizi (Cooper) (desert tortoise), Ephed-
ra viridis Cov., E. californica Wats., and Yucca whipplei Torr. As
the climate cooled, some species were extirpated (e.g., the desert
tortoise), whereas others became restricted to the drier parts of near-
by mountains (e.g., the Diablo and Temblor ranges).

A number of lines of evidence, however, indicate that the presence
and spread of P. glandulosa and P. pubescens in the SJV are due to
human-induced factors, and that prior to the 1870’s neither species
existed there. Support for this hypothesis is based largely upon neg-
ative evidence, such as a lack of documentation of the extended
occurrence of either species where it would be expected to exist. I
summarize the salient points in the present paper.

HISTORICAL EVIDENCE

Available historical accounts of expeditions through areas where
mesquite now occurs or occurred in the recent past lack documen-
tation of the presence of the species. A particularly interesting ac-
count is that of Lt. George Derby, who was commissioned by the
War Department in 1850 to survey the “‘Tulare Valley” (the area
between Tulare and Buena Vista lakes) for the purpose of establishing
a military outpost. Derby’s party traveled along the north bank of
the north fork of the Kern River to the north shore of Buena Vista
Lake. His account of the area (Boyd 1977) reads as follows: ‘Like
other bodies of water in the valley, it is nearly surrounded by tules
[Scirpus], and upon its north and east banks there is a heavy growth
of willows. A slough, some sixty miles in length [Goose Lake Slough],
connects it with the swamps and bodies of standing water in the bed

326 MADRONO [Vol. 34

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ae pee, SS

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SAN LUIS
OBISPO

Prosopis glandulosa
current range ~—= |

historical range --—-—

Prosopis pubescens ©

Prosopis velutina

Fic. 1. Distribution of various species of Prosopis in the San Joaquin Valley,
California. - = isolated populations, usually of less than 10 trees. ? = unvouchered
occurrence in the vicinity of the Cuyama River, San Luis Obispo Co.

1987] HOLLAND: PROSOPIS, RELICT OR INVADER? 327

of the Ton Tache [south of present-day Alpaugh], and through them
with the great northern lake [Tulare Lake]. The surrounding country
is sterile and unproductive when not an absolute swamp... . nothing
can be conceived more inappropriate than its name, for no place
can be imagined more forlorn or desolate in appearance”. Traveling
north by way of the east side of Goose Lake Slough, Derby continued
his appraisal of the region, noting that it was of “‘precisely the same
character throughout—barren, decomposed soil with no trace of
vegetation but a few straggling Artemesias [presumably A/lenrolfea
or Atriplex], except on the margins of the creeks’’. Given Derby’s
attention to details of the flora in areas such as the Tule River and
Poso Creek, it seems curious that he would fail to note the presence
of a plant as conspicuous as mesquite, now one of the largest and
relatively common elements of the flora in this area (Fig. 2).

The account of W. H. Brewer (Farquhar 1966) concerning the area
of Corral Hollow (San Joaquin and Alameda cos.) is fairly detailed,
noting the species and relative abundance of trees in an area where
they are often scarce. Again there is no mention of mesquite or even
anything that might resemble it.

War Department surveys in the 1850’s (Williamson 1853) for a
railroad route through the Central Valley and their botanical col-
lections also failed to document or note the presence of mesquite,
as did the botanical portion of the California Geological Survey of
the area (Gray 1876). In 1983, I retraced most of the route followed
by the railroad survey and noted the occurrence of mesquite in at
least two areas. A vegetational survey (Davy 1898) of the “waste
lands” (alkali sink areas) southwest of Bakersfield in 1896 failed to
note the presence of mesquite. The first published reference con-
firming the presence of P. glandulosa in this area was that of Linton
(1908), noting that along Buena Vista Lake “‘on the north shore for
several miles is an alkaline desert with an occasional patch of mes-
quite and sage.”

I examined over 100 photographs taken from the late 1880’s to
the early 1900’s in the hypothesized area of establishment and was
unable to identify anything that resembled mesquite, although other
native taxa (Populus, Salix, Allenrolfea) were readily observable.

There is a consistent lack of herbarium collections from the pro-
posed areas of origin during the time period of concern. This is a
weak argument, as the number and scope of collections from the
SJV were limited, and the collectors were often more concerned with
annuals and smaller perennials (Eastwood 1893). The earliest known
collection is a specimen of P. glandulosa from the area of Button-
willow, Kern Co., collected in 1914 (CAS #65104).

A paucity of paleobotanical information exists concerning the flora
of the valley floor at the time of the presumed invasion and estab-

328 MADRONO [Vol. 34

re... on

Ps

0 PR leti “ SS de

Fic. 2. Prosopis—Atriplex association near the former site of Buena Vista Lake
and in the approximate vicinity of the route traversed by Derby’s party.

lishment of many Mojavean elements. Available information does
not indicate that mesquite was present in areas near Buena Vista
Lake (Mason 1944). Mesquite did reach the “‘rim”’ of the San Joaquin
Valley in the Pliocene, however, as indicated by fossil material from
the vicinity of Tehachapi, at the current western edge of the Mojave
Desert (Axelrod 1950). Analyses of fossil pollen records in Arizona
have documented the long-term existence of mesquite (Martin 1963),
but I was unable to find any core sample data from the area and
time period of concern for this study.

ETHNOBOTANICAL EVIDENCE

The seed pods of P. glandulosa are and were used widely as a food
source by the aboriginal inhabitants of the southwestern United
States and adjacent Mexico, and the fibers were utilized to some
extent in basketry. The area around Buena Vista and Tulare lakes
was inhabited by various groups of the Valley Floor Yokuts (Latta
1949). Ethnographers, e.g., Alfred Kroeber and Frank Latta, have
gained valuable insights into the ecology of California aboriginals,
which included their uses of the native flora. Latta’s (1949) study
of the Yokut culture made no mention of the use of mesquite despite
an otherwise extensive compendium of food, fiber, and medicinal
plants. Driver (1937) questioned members of Central Valley and

1987] HOLLAND: PROSOPIS, RELICT OR INVADER? 529

Death Valley tribes about their use of mesquite. Responses from the
tribes of Death Valley and adjacent areas indicated extensive use,
whereas there were no such indications from Valley Floor peoples.
Furthermore, use of mesquite fibers in baskets among tribes of the
Mojave Desert was documented but was conspicuously lacking among
Valley Floor tribes (Merrill 1923).

ENVIRONMENTAL DISTURBANCE

The areas in which I suspect P. glandulosa and P. pubescens ini-
tially became established have an extensive history of environmental
disturbance, both human-induced and otherwise (floods, drought).
This situation may have fostered the establishment of mesquite. I
suggest that in the interval from 1870-1890 P. glandulosa became
established in the area between Bakersfield and Buena Vista Lake.
The hypothesized mechanism of establishment was ingestion of seed
pods by cattle in areas where mesquite occurs naturally followed by
transport of those cattle by rail to the SJV, defecation by the cattle,
and subsequent germination of the seeds. Benson (1941) implicated
this mechanism in the establishment of mesquite in Louisiana and
Missouri. I suspect a similar situation led to the establishment of P.
pubescens in the vicinity of Coalinga in western Fresno Co. Darrell
Zwang (pers. comm.), a long-term (70+ yr) resident of the area,
noted that cattle transport was nonstop, increasing the probability
that this mechanism occurred. The seeds of P. velutina are known
to remain viable for extended periods of time (44 yr) (Martin 1948),
which also increases the probability of establishment. If mesquite
was carried as fodder in cattle cars, establishment might be due to
incidental release into the habitat.

The establishment of mesquite 1n these and similar sites was prob-
ably facilitated by at least two types of environmental disturbances.
Many areas in which Prosopis occurs are situated in flood plains or
washes that are periodically inundated or have high water tables.
Prior to the construction of Isabella Dam in 1952, much of the
Buena Vista Lake area was unsuitable for continuous agricultural
use due to recurrent flooding. In certain areas of the southwest,
similar situations have fostered the establishment of extensive mes-
quite-dominated communities (Minckley and Clark 1984).

Heavy grazing has been shown to facilitate the establishment and
spread of mesquite (Glendening 1952, Martin 1975). The number
of cattle in Kern Co. increased over 500% between 1870-1880, and
an additional 350% from 1880-1890. By the 1890’s, approximately
16% of all the cattle in California were being grazed in Kern Co.
(Burcham 1957). Further spread of Prosopis may have been facili-
tated by the ingestion of seed pods from maturing trees and move-
ment of cattle to other areas. Populations in Alameda and Fresno

330 MADRONO [Vol. 34

cos. may have become established as the result of deliberate plant-
ings.

ECOLOGICAL CONSIDERATIONS

The ecology of the two species of Prosopis lends support to this
hypothesis of establishment. Prosopis glandulosa and P. pubescens
are known to invade disturbed areas, particularly in association with
drought and overgrazing (Caraher 1970, Herbel et al. 1972, Cable
1973). The physiognomy of an area such as the Santa Rita Exper-
imental Range in Arizona (Martin 1973) can change to such an extent
in as little as 50 yr that to those unfamiliar with the original ap-
pearance of the site, mesquite might appear to be a “‘normal’’ dom-
inant. Historical (Hastings and Turner 1965) and recent (Minckley
and Clark 1984) evidence of this type has been noted in the ap-
pearance of mesquite-dominated floodplains in Arizona.

I have observed extensive, apparently suitable habitat for mes-
quite in the SJV, and, thus, it is curious that I have found neither
species more widespread. For example, several large canyons with
similar relief and soil conditions occur to the north and south of the
Fresno Co. populations of P. pubescens, but the species is not known
from any of these.

Prosopis glandulosa formerly covered an estimated 25-35 km? in
the Old River area of Kern Co. (Bill Asserson, pers. comm.), where
it coexisted with a mixture of A/lenrolfea occidentalis, Suaeda mo-
quinil, Atriplex polycarpa, and A. lentiformis. Estimates by Wersch-
kull et al. (1983) indicate that the Prosopis—Atriplex association cov-
ered over 57,000 ha at its peak. Based upon my research, I estimate
that at most, only 20,000—25,000 ha in the SJV supported mesquite
in any association, and of this only 8000—12,000 ha supported high-
density stands. Given the extensive areas that have seemingly suit-
able soil and climatic characteristics and the invasive abilities of
this group, I estimate that the potential range for Prosopis spp. in
the SJV and vicinity was or is greater than 75,000 ha. I suggest that
this area has not been occupied due to insufficient time for the species
to spread since their advent in the late 1800’s. Additionally, massive
habitat alteration and concurrent destruction of many populations
by humans has slowed the spread.

CURRENT STATUS

The status of mesquite populations in the SJV is of concern to
conservationists who consider the Prosopis—Atriplex association to
be a threatened plant community (Jack Zaninovich, pers. comm.).
Construction of the Central Valley Water Project and the California
Aqueduct resulted in the conversion of a considerable percentage of
the remaining wildlands of the Central Valley to agricultural use.

1987] HOLLAND: PROSOPIS, RELICT OR INVADER? 331

Mesquite-dominated communities were estimated to cover approx-
imately 20,000 ha in Kern Co. in 1963 (CDFG 1965), but patterns
of land use over the next 20 yr were projected to reduce this cover
to zero. Major reductions in the habitats supporting mesquite did
take place over this time interval: approximately 2000 ha remained
in Kern Co. in 1979 (Bill Asserson, pers. comm.), and 6500 ha
remained in the Tulare basin as a whole (Werschkull et al. 1983). I
estimate that as of summer 1986 about 5000 ha of habitat supporting
P. glandulosa remained in the SJV. Of this, only 15-20% supports
vigorous populations. Lowering of water tables in many areas may
result in the decline of this species.

I estimate the amount of habitat supporting P. pubescens to be
less than 1000 ha (primarily in Warthan Canyon), and the number
of trees probably does not exceed a few hundred. The status of the
populations in Los Gatos Canyon and San Emigdio Canyon is un-
known. Prosopis velutina probably is represented by only a few trees
in the vicinity of Bakersfield (Hilu et al. 1982).

Populations of Prosopis are commonly sympatric with other gen-
era of native plants, including Atriplex, Allenrolfea, Cephalanthus,
Salix, and Suaeda. Destruction of mesquite populations will nec-
essarily entail the alteration of a large percentage of the remaining
areas of native vegetation in the southern (San Joaquin) valley. For
example, the largest remaining population of P. glandulosa occurs
along the lower Kern River southwest of Bakersfield. Approximately
1100 ha of this area is being developed as a ground-water recharge
facility for the City of Bakersfield. This action will eliminate most
of the native habitat containing mesquite (Stetson Engineers 1983).

CONCLUSIONS

Historical, ethnobotanical, and ecological evidence indicate that
one or both species of mesquite may have naturalized recently (< 120
yr) in the SJV. This is equivocal, however, as “. . . | have never seen,
and never shall see, that the cessation of the evidence of existence
is necessarily evidence of the cessation of existence” (de Morgan
1906). The question of status might be resolved through paleobo-
tanical evidence or core sampling that documents the presence of
the species well prior to the hypothesized period of establishment
(1870-1890). Mesquite, however, may deposit several growth rings
per year (Tom Griggs, pers. comm.), which may frustrate efforts to
date individuals.

Monitoring of mesquite populations in the southern SJV is of
primary importance. Valuable scientific opportunities will be lost
with their further decline or elimination. If the species are indeed
native, the chance will be lost to study their associations with floral
elements that do not occur elsewhere. If my recent-invasive hy-

332 MADRONO [Vol. 34

pothesis is correct, the destruction of mesquite-dominated com-
munities represents the loss of an unusual opportunity to better
understand the nature of historical habitat disturbance and invasive
plant ecology.

ACKNOWLEDGMENTS

Inquiries concerning specimens of Prosopis were conducted at the following her-
baria— Bakersfield Junior College, California Department of Food and Agriculture,
California State College—Bakersfield, Merced Junior College, CAS, CSLA, CSPU, DS,
FSC, LA, OBI, POM, RSA, SJSU, UC, and UCSB. I thank Daniel Axelrod, Jim
Bartel (USFWS), Annetta Carter, Elizabeth Coley, W. J. Ferlatte, Wayne Ferren,
Marc Hayes, James Jackson, Robert Jaeger, William Reese, Barry Tanowitz, and an
anonymous reviewer for editorial comments. I also thank Bill Asserson (CDFG),
Lyman Benson, Ben Chichester, Tom Griggs, Khirdir Hilu, Howard Latimer, May-
nard Moe, John Stebbins, Jack Zaninovich, Darrell Zwang, and the curators of the
herbaria noted. This paper is dedicated to the memory of Lt. John Reed (CDFG)
who gave his life in the protection of California’s natural resources.

LITERATURE CITED

AXELROD, D. 1950. Evolution of desert vegetation in western North America. Car-
negie Inst. Wash. Publ. 590:215-360.

BARBOUR, M. and J. MAyor. 1980. Terrestrial vegetation of California. John Wiley
& Sons, New York.

BENSON, L. 1941. The mesquites and screw-beans of the United States. Amer. J.
Bot. 28:748-754.

Boyp, W. 1977. Kern County wayfarers 1844-1881. Kern Co. Hist. Soc. & Kern
Co. Museum, Bakersfield, CA.

BuRCHAM, L. 1957. California range land: a historico-ecological study of the range
resource in California. Div. of For., State of California, Sacramento.

CaBLE, D. 1973. Invasion of semi-desert grassland by Velvet mesquite and asso-
ciated vegetation changes. J. Ariz. Acad. Sci. 8:127-134.

CALIFORNIA DEPARTMENT OF FISH AND GAME. 1965. California Fish & Wildlife
Plan. Vol. III. Supporting data. Sacramento.

CARAHER, D. 1970. Effects of longtime livestock exclusion versus grazing on the
desert grassland of Arizona. M.S. thesis, Univ. Arizona.

Davy, J. 1898. Natural vegetation of alkali lands. Calif. Agr. Exp. St., Berkeley Rrt.
1895-1898:63-75.

Driver, H. 1937. Culture element distributions VI: southern Sierra Nevada. Univ.
Cal. Publ. Am. Arch. and Ethn. 37:53-154.

EAstTwoop, A. 1893. Field notes at San Emigdio. Zoe 4:144-147.

FARQUHAR, F. P., ed. 1966. Up and down California in 1860-1864: the journal of
Dr. William H. Brewer. Univ. California Press, Berkeley.

GLENDENING, G. E. 1952. Some quantitative data on the increase of mesquite and
cactus on a desert grassland range in southern Arizona. Ecology 33:319-328.

Gray, A. 1876. Botany. Vol. I. Gamopetalae. Geological Survey of California.
Welch, Bigelow & Co. Univ. Press, Cambridge, MA.

HAsTINGs, J. R. and R.M. TURNER. 1965. The changing mile: an ecological study
of vegetation change with time in the lower mile of an arid and semiarid region.
Univ. Arizona Press, Tucson.

HERBEL, C., F. Ares, and R. WriGHT. 1972. Drought effects on a semi-desert
grassland range. Ecology 53:1084—-1093.

Hiiu, K., S. Boyp, and P. FELKER. 1982. Morphological diversity and taxonomy
of California mesquites (Prosopis, Leguminosae). Madrono 29:237-254.

1987] HOLLAND: PROSOPIS, RELICT OR INVADER? 333

LaTTA, F. 1949. Handbook of Yokuts Indians. Bear State Books, Oildale, CA.

LINTON, C. 1908. Notes from Buena Vista Lake, May 20 to June 16, 1907. Condor
10:196-198.

Martin, P. S. 1963. The last 10,000 years: a fossil pollen record of the American
southwest. Univ. Arizona Press, Tucson.

Martin, S.C. 1948. Mesquite seeds remain viable after 44 years. Ecology 29:393.

1973. Santa Rita Experimental Range: your facility for research on semi-

desert ecosystems. J. Ariz. Acad. Sci. 8:56-67.

1975. Ecology and management of southwestern semi-desert grass-shrub
ranges: the status of our knowledge. U.S.D.A. Forest Service Res. paper RM-
156.

Mason, H. 1944. A Pleistocene flora from the McKittrick asphalt deposits of Cal-
ifornia. Proc. Cal. Acad. Sci. 25:221-234.

MERRILL, R. 1923. Plants used in basketry by the California Indians. Univ. Cali-
fornia Publ. Am. Arch. and Ethn. 20.

MINCKLEY, W. and T. CLARK. 1984. Formation and destruction of a Gila River
Mesquite Bosque community. Desert Plants 6:23-30.

DE MorGAn, W. 1906. Joseph Vance: an ill-written autobiography. H. Holt and
Co., New York.

Munz, P. 1959. A California flora. Univ. California Press, Berkeley.

STETSON ENGINEERS, INC. 1983. Draft Environmental Impact Report—2800 acre
ground-water recharge facility along the Kern River for the City of Bakersfield.

WERSCHKULL, G., T. GRIGGS, and J. ZANINOVICH. 1983. Tulare Basin protection
plan. Publ. Calif. Nature Conservancy.

WILLIAMSON, R. 1853. Report on explorations in California for railroad routes to
connect with the routes near the 35th and 32nd Parallels of north latitude. U.S.
War Dept.

(Received 27 Nov 1985; revision accepted 16 Jun 1987.)

ANNOUNCEMENT

1987 AWARDS PRESENTED BY ASPT

The George R. Cooley Award for 1987 was presented to Robert Wyatt
of the University of Georgia, Athens, for his paper co-authored with
Ireneusz J. Odrzykoski and Ann Stoneburner entitled “‘Allopolyploidy
in bryophytes: recurring origins of Plagiomnium medium.” The award
is given annually by the American Society of Plant Taxonomists for the

outstanding contributed paper in plant systematics presented at the
annual meeting.

The fourth Asa Gray Award was presented to Reed C. Rollins of
Harvard University, Cambridge, Massachusetts. The Asa Gray Award
is given by the American Society of Plant Taxonomists to honor an
individual “for outstanding accomplishments pertinent to the goals of
the Society.”’ The award has been presented to Rogers McVaugh at the
1984 meeting, Arthur Cronquist at the 1985 meeting, and Lincoln Con-
stance at the 1986 meeting.

SOME NEW AND RECONSIDERED CALIFORNIA
DUDLEYA (CRASSULACEAE)

KEI M. NAKAI
Herbarium, Mildred E. Mathias Botanical Garden,

University of California,
Los Angeles 90024

ABSTRACT

Three new taxa of Dudleya from California are described: Dudleya cymosa subsp.
agourensis, D. cymosa subsp. crebrifolia, and D. abramsii subsp. affinis. Three new
combinations of Dudleya also are proposed from California: D. cymosa subsp. pumila,
D. cymosa subsp. paniculata, and D. abramsii subsp. calcicola. Dudleya gigantea is
reduced to a synonym of D. cymosa subsp. cymosa and D. minor is reduced to a
synonym of D. lanceolata. A neotype is designated for Echeveria cymosa Lem. (=D.
cymosa subsp. cymosa).

Recent collections made during a study of the Dudleya cymosa—
abramsii complex revealed some undescribed taxa and a need to re-
evaluate others. Dudleya species often exhibit considerable pheno-
typic plasticity resulting from the variability of climatic conditions,
soil, and exposure that often alters, sometimes quite dramatically,
the appearance of the same plant from year to year. Although mor-
phological measurements taken from the field are important to ac-
curately identify many plants, field measurements from succulent
plants like Dudleya often have proven to be unreliable. Therefore,
the interpretation of each taxon discussed here will rely heavily on
data obtained from cultivated plants in an effort to reduce variability
in certain morphological characters (e.g., leaf size and shape, floral
stem length, inflorescence shape, pedicel length, etc.) that are used
in determining taxa. This approach makes it possible to evaluate
under a uniform and stable environment those characters that ap-
parently are controlled genetically.

MATERIALS AND METHODS

In collecting live material for study, five plants were selected by
tossing a 10 cm hoop within an area greater than 9 m? where there
were many flowering plants of Dudleya. The first five plants selected
were measured in the field and then collected for cultivation studies.

Herbarium and living specimens were examined. Most of the floral
measurements were obtained from wild or cultivated living material.
Because of the tendency for certain flower parts, in particular the
staminal filaments, of Dudleya to continue to grow even after the

MADRONO, Vol. 34, No. 4, pp. 334-353, 1987

1987] NAKAI: DUDLEYA 335

flower appears to be fully mature, all measurements were taken two
days following anther dehiscence.

Cultivated plants were grown at Hawthorne, California (33°55'N,
118°22'W), in a structure constructed with a clear fiberglass roof and
30% shadecloth for the sides. An artificial soil was used that consisted
of two parts fine redwood shavings, two parts coarse Canadian peat
moss, two parts diatomaceous earth, one part sand, and one part
fine perlite. One kg of milorganite was incorporated for each m? of
medium. The medium was then moistened and aged for a minimum
of one month.

Flower buds used for cytological observations were collected from
cultivated plants between 0930 and 1000 hr and fixed in a modified
Carnoy solution (Uhl and Moran 1953), which consisted of chlo-
roform, EtOH, and glacial acetic acid (3:2:1, v/v/v). Counts were
made from a minimum of five buds.

TAXONOMIC TREATMENT

DUDLEYA CYMOSA (Lemaire) Britton & Rose subsp. CYMOSA— Ech-
everia cymosa Lemaire, Revue Hortic. 7:439. 1858.— Cotyledon
cymosa Baker in Saunders, Refug. Bot. 1:pl. 69. 1869.— Dudleya
cymosa Britton & Rose, Bull. N.Y. Bot. Gard 3:21. 1903.—
Cotyledon laxa var. cymosa Jepson, Man. FI. Pl. Calif. 453.
1925.—Echeveria laxa var. cymosa Jepson, Fl. Calif. 2:114.
1936.—Neotype: Plate in Saunders, Refug. Bot., pl. 69. 1869.

Dudleya gigantea Rose in Britton & Rose, op. cit. p. 23.—Cotyledon
gigantea Fedde, Bot. Jahresber. Just. 31:826. 1904.—Echeveria
amadorana Berger in Engler & Prantl, Nat. Pflanzen fam., ed.
2, 18a:479. 1930 (based on Dudleya gigantea Rose).—TYPE:
USA, CA, Amador Co., New York Falls, ca. 1500 ft (460 m),
15 Jun 1896, G. Hansen 2012 (Holotype: US! (US 338497),
photo LA!; isotype: CAS!, NY!).

Echeveria lanceolata var. incerta Jepson, Fl. Calif., p. 115. 1936.—
Type: USA, CA, Calaveras Co., Calaveritas Creek, near Ken-
tucky House, ca. 900 ft (275 m), 27 May 1923, W. L. Jepson
9919 (Holotype: JEPS!, photo LA!).

Caudex short, usually less than 5 cm long, 1—3.5 cm diam., un-
branched or few branched. Basal rosettes 6—20(—25) cm diam., con-
sisting of 6—25 oblanceolate to rarely spatulate leaves, acute, acu-
minate or, infrequently, cuspidate, 2-17 cm long, 1.5-6 cm wide,
and 1-5 mm diam. Floral stem 0.5—4.5 dm tall, 2-8 mm diam., with
7-20(—30) horizontal to ascending ovate to triangular-lanceolate
leaves, acute to acuminate, the lowermost 0.5—3(—10) cm long and
5-15 mm wide. Inflorescence obpyramidal, infrequently paniculate
or simple, commonly with 2—4 branches that rebranch O-3 times;
cincinnus circinate when young, ascending in age, 1—5(—15) cm long

336 MADRONO [Vol. 34

and (1—)2—10(—20) flowers; pedicels erect, the lowermost 5-15 mm
long, 0.5—2 mm diam. Calyx 3-7 mm long, 2.5—6 mm wide, rounded
to truncate below; lobes triangular to triangular-ovate, acute to +
acuminate, 1.5-—5 mm long, 1.5—4 mm wide. Corolla ovoid in bud,
cylindrical in anthesis, often with the petal apices spreading from
45°-90°; petals yellow, orange, or red, occasionally glaucous along
the midrib, elliptic to narrowly lanceolate, acute, 7-15 mm long, 2-
4 mm wide, connate 1—2.5 mm; filaments 4-8.5 mm long, adnate
for 1-3.5 mm, the epipetalous mostly 0.5 mm shorter and adnate,
mostly 0.5—0.7 mm higher than the antesepalous; anthers yellow,
1-2 mm long. Gynoecium 4—10 mm long, erect when young, slightly
spreading in age, ovaries 3-8 mm long, styles 1-2 mm long. Nectaries
reniform, 1-2 mm wide. Chromosome number: n = 17. Flowering
April to July.

Distribution. USA, California: Coast Ranges from Humboldt Co.
to Santa Clara Co.; Sierra Nevada. Elev. 100-2700 m.

Based on the original description, the type locality is probably in
California (“‘... corolles jaune-pale. Californie? Tres distincle!’’—
Lemaire 1858). According to Moran (1951), no authentic specimen
is available; however, he noted that Lemaire’s plant came from the
horticulturist Louis de Smet of Ledeburg, Belgium, in or before 1858,
and the plant illustrated by Baker in Saunders Refugio Botanicum
came from a horticulturist in nearby Ghent in 1855 and probably
was of the same introduction. Because the plate compares well with
the original description and clearly illustrates most of the plants
presently referred to Dudleya cymosa, this plate serves as the neotype
until an authentic type specimen is found.

Based on its greater average size when compared with D. cymosa
subsp. cymosa, Moran (1951, 1957) recognized Dudleya gigantea as
a subspecies of Dudleya cymosa. Moran (1951) noted, however, that
there was a dwarf specimen on the type sheet. He also mentioned
that subsp. cymosa occurs at nearby localities, with no evident nat-
ural barriers. As a result of additional collections and the data from
cultivation, the size difference between D. gigantea and D. cymosa
is not apparent and, thus, there is not enough evidence to warrant
maintaining D. gigantea as a distinct taxonomic entity.

Moran (1951, 1960) reduced Echeveria lanceolata var. incerta to
a synonym of Dudleya cymosa subsp. gigantea. Variety incerta also
does not appear to be different from D. cymosa subsp. cymosa.

Dudleya cymosa subsp. pumila (Rose) K. Nakai, comb. nov. — Dud-
leya pumila Rose, Bull. N.Y. Bot. Gard. 3:14. 1903.— Cotyledon
pumila Fedde, Bot. Jahresber. Just. 31:826. 1904.—Echeveria
parva Berger in Engler & Prantl, Nat. Pflanzenfam., ed. 2, 18a:
477. 1930 (based on Dudleya pumila Rose).— Type: USA, CA,

1987] NAKAI: DUDLEYA 337

San Bernardino Co., San Bernardino Mtns., between Green Val-
ley and Deep Creek, 7000 ft (2120 m), 19 Jul 1899, H. M. Hall
1350 (Holotype: US!, photo DS!, LA!; isotype: UC’).

Distribution. USA, California: South Coast Ranges from Monterey
Co. south to the San Gabriel and San Bernardino mtns. Elev. 100-
1800 m from Santa Barbara Co. north, 600—2600 m from Ventura
Co. south.

Rose (1903) based D. minor on two collections [Rose 421 (ex
Hasse) (NY, US), 1893 McClatchie (NY, UC)]; both were collected
from San Gabriel Canyon. Moran (1957) reduced D. minor to a
subspecies of D. cymosa. Munz (1959, 1974) lists D. cymosa subsp.
minor from the Santa Lucia Range in Monterey Co. south into the
Transverse Range of southern California.

I was unable to find any dudleya resembling the many herbarium
specimens labeled D. minor or D. cymosa subsp. minor (Rose) Moran
at the type locality. Although the original description suggests a
relationship with D. cymosa based on leaf shape, pedicel length, and
petal shape, I have determined that the holotype of D. minor resem-
bles plants intermediate between D. cymosaand D. lanceolata (Nutt.)
Britt. & Rose.

In southern California, D. cymosa is usually a small plant, mostly
5-8 cm in diameter. In the central portion of the Transverse Ranges,
the rosette leaves are mostly oblanceolate to spatulate. The floral
stem is usually less than 15 cm tall, the inflorescence is rather com-
pact with 4—8 flowers per cincinnus. The lowermost pedicels are
often at least as long as the length of the corolla. Dudleya cymosa
is a diploid (n = 17; Uhl and Moran 1953).

Dudleya lanceolata is often larger, 8-12 cm in diameter. The ro-
sette leaves are typically oblong to lanceolate or infrequently elliptic
to ovate. The floral stem is usually greater than 15 cm tall, the
inflorescence is often lax with 6—15 flowers per cincinnus. The low-
ermost pedicels may be as long as the corolla, but usually they are
much shorter. Dudleya lanceolata is a tetraploid (n = 34; Uhl and
Moran 1953).

The shape of the basal rosette leaves and the length of the pedicels
of the holotype of D. minor suggest a relationship with D. cymosa,
whereas the length of the floral stem, the more lax inflorescence,
and the greater number of flowers per cincinnus is similar to D.
lanceolata. Indeed, most of the dudleyas observed from elevation
500-1000 m were clearly D. lanceolata; however, a few individuals
resemble D. cymosa morphologically even though cytologically they
were tetraploids.

For comparison of Dudleya minor with D. cymosa and D. lan-
ceolata, dudleyas were collected along California State Highway 39,
which travels through San Gabriel Canyon, at elevational increments

338 MADRONO [Vol. 34

of 100 m from 500-1700 m. Morphological measurements were
made in the field. The plants were then cultivated and the same
measurements were repeated the following year and chromosome
counts were made. The measurements were translated into numer-
ical values or scores so that the sum of the total characters from
each population may be compared (Table 1, Fig. 1). Voucher spec-
imens were deposited at LA.

Figure 1 shows there is a distinct break in morphological mea-
surements between 1200-1500 m. Wild plants from 500-1200 m
had an average score of 11.5, whereas those from 1500-1700 m had
an average of 18.7. Cultivated plants displayed a similar break,
although less than that found in wild plants (13.3 vs. 19.0). The
chromosome number also correlated with elevation. Tetraploid plants
occurred below 1200 m and diploid plants occurred above 1500 m.
Based on morphology and chromosome number, D. lanceolata ap-
parently occurs from 500-1200 m and D. cymosa is found above
1500 m in San Gabriel Canyon.

Although the type specimen of D. minor may resemble D. cymosa,
it was collected below 1200 m. Thus, on the basis of morphological
and cytological data obtained from both wild and cultivated plants
I consider D. minor to be conspecific with D. lanceolata. Because
the type specimen of D. pumila Rose represents most of the D.
cymosa in southern California, I propose the combination D. cymosa
subsp. pumila to replace D. cymosa subsp. minor.

Dudleya cymosa subsp. paniculata (Jeps.) K. Nakai, comb. nov.—
Cotyledon caespitosa var. paniculata Jeps., Fl. W. Mid. Calif.
267. 1901.—Dudleya paniculata Britt. & Rose, Bull. N.Y. Bot.
Gard. 3:27. 1903.— Cotyledon paniculata Fedde, Bot. Jahresber.
Gard. 31:826. 1904 (non C. paniculata Thunberg.).—Echeveria
jepsonii Nelson & Macbride, Bot. Gaz. (Crawfordsville) 56:477.
1913 (based on Cotyledon caespitosa var. paniculata Jeps.).—
Cotyledon laxa var. paniculata Jeps., Man. Fl. Pl. Calif. 543.
1925.—Echeveria laxa var. paniculata Jeps. Fl. Calif. 2:114.
1936.—Type: USA, CA, Alameda Co., Morrison Canyon, 20
Jun 1897, W. L. Jepson 13419 (Holotype: JEPS!, photo LA!).

Dudleya humilis Rose, Bull. N.Y. Bot. Gard. 3:27. 1903.— Cotyledon
humilis Fedde, Bot. Jahresber. Just. 31:826. 1904 (non C. hu-
milis Marloth. 1915).—Echeveria diaboli Berger in Engler &
Prantl, Nat. Pflanzenfam., ed. 2, 18a:480. 1930 (based on Dud-
leva humilis Rose.).—TyYPe: USA, CA, Contra Costa Co., sum-
mit of Mt. Diablo, 2 Jun 1903, Alice Eastwood s.n. (Rose 620)
(Holotype: US!, photo LA!; isotype: NY!).

Distribution. Inner South Coast Range from Contra Costa Co.
south to western Fresno and northeastern Monterey cos.

1987] NAKAI: DUDLE YA Bog

Moran (1951, 1960) considered Cotyledon caespitosa var. panicu-
lata a synonym of Dudleya cymosa subsp. setchellii (Jeps.) Moran.
He noted, however, that they might be separated on the basis of the
rosette leaf shape. Based on studying 34 herbarium specimens and
100 live plants from 20 populations of subsp. paniculata and 13
herbarium specimens and 20 live plants from four populations of
subsp. setchellii, I found that subsp. paniculata differs from subsp.
setchellii by having oblong to oblanceolate basal rosette leaves com-
pared with oblong-triangular leaves, an inflorescence of 2—3 branches
that rebranch one or twice rather than 2—3 mostly simple branches,
and pedicels 6—12 mm long versus 4—7 mm long. Subspecies setchellii
is restricted to the serpentine rock outcrops within the Santa Clara
Valley, whereas subsp. paniculata occurs within the Inner South
Coast Range on various rock substrates.

Dudleya humilis Rose is reduced to a synonym of D. cymosa subsp.
paniculata. A similar form [K. Nakai 816 (LA)] was collected near
the summit of Mt. Hamilton, which is south of Mt. Diablo, the type
locality of D. humilis. Dudleya humilis appears to be an edaphic
dwarf of subsp. paniculata because cultivated plants I have grown
from each location did not appear different from cultivated plants
of subsp. paniculata.

Dudleya cymosa subsp. agourensis K. Nakai, subsp. nov.

A subspecie typica caulis ramosus, rosulae foliis 6-10, glaucis,
ellipticis vel oblongis differt. Figs. 2a,b, 5.

Plants simple or with six or more cespitosely branched rosettes
5-10 cm diam., with 6-10 elliptic to oblong glaucous leaves. Basal
rosette leaves 3-10 cm long, 1—1.5 cm wide, acute to acuminate;
cauline leaves lanceolate, glaucous, 1—2.5 cm long, 7-10 mm wide,
acute to acuminate. Floral stem erect, 10-20 cm tall, glaucous, often
tinged with red; inflorescence of 2—3 simple to bifurcate branches;
cincinnus ascending, 1-3 cm long with 3-8 flowers; lowermost ped-
icels 6—12 mm long. Petals bright yellow, occasionally glaucous along
the midrib, petal apex spreading 45—90°. Chromosome number: n =
17. Flowering May to June.

Type: USA, CA, Los Angeles Co., Santa Monica Mtns., ca. 0.5
km s. of the junction of Agoura and Cornell roads on Cornell Road,
34°0814'N, 118°45'4’W, on nw.-facing volcanic rock road embank-
ment, locally abundant, 275 m. Associated with Malosma laurina,
Haplopappus linearis, Dichelostemma pulchella, Delphinium parryi,
Calochortus venustus. 27 May 1980, K. Nakai 606 (Holotype: CAS;
isotype: LA, SD).

PARATYPES: Los Angeles Co., w. of Calabasas Moran 3472 (UC);
Agoura, along Cornell Road, Nakai 436 (LA); n.-facing volcanic

[Vol. 34

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ELEVATION (X|OO METERS)

Fic. 1. Bar graph illustrating index score vs. elevation derived from Table 1 for
Dudleya cymosa and D. lanceolata as collected from along California State Highway
39. O = wild plants; M@ = cultivated plants.

slopes of Ladyface Mountain, 400 m, Nakai 1119 (CAS, LA); rocks
on the e. side of State Hwy. 23 near the jct. of Portrero Road and
Hwy. 23, 365 m, Nakai 512 (LA). Ventura Co., n.-facing volcanic
rock along Hwy. 23 e. of Lake Eleanor, 300 m, Nakai 607 (CAS,
LA).

Distribution. North slope of the western portions of the Santa
Monica Mtns.

Dudleya cymosa subsp. agourensis differs from subsp. cymosa by
having rosettes consisting of 6-10 elliptic to oblong leaves rather
than 10—25 leaves that are oblanceolate to spatulate. Uhl and Moran
(1953) placed a population of subsp. agourensis (Moran 3472) with
D. cymosa subsp. ovatifolia (Britt.) Moran. Nakai (1983) considered
this population a distinct race of subsp. ovatifolia. Subspecies ova-
tifolia also occurs in the Santa Monica Mountains and, like subsp.
agourensis, 1t possesses yellow petals and rosettes of 6—10 leaves.
Subspecies ovatifolia, however, differs from subsp. agourensis by its
unbranched caudex, ovate to elliptic basal rosette leaves, green with

—

Fics. 2-4. Photographs of new taxa in Dudleya. 2. Dudleya cymosa subsp. agou-
rensis K. Nakai. a. Isotype plant (0.4 x). b. Inflorescence in detail (1.25 x). 3. Dudleya
cymosa subsp. crebrifolia Nakai & Verity. a. Holotype plant (0.2 x). b. Inflorescence
in detail (2x). 4. Dudleya abramsii subsp. affinis K. Nakai. a. Inflorescence in detail
(1.25 x). b. Isotype plant (0.5 x).

1987] NAKAI: DUDLEYA 343

344 MADRONO [Vol. 34

I SAN BERNARDINO CO

) - D. ABRAMSII AFFINIS

D. CYMOSA “AGOURENSIS :

!
/
D.CYMOSA 9 |

Fic. 5. Geographic distribution of Dudleya cymosa subsp. agourensis, D. cymosa
subsp. crebrifolia, and D. abramsii subsp. affinis.

a maroon suffusion on the underside, slightly longer pedicels, and
petal apices spreading 90° or more. Subspecies ovatifolia is found
on the southern slopes of the Santa Monica Mountains on shaded
sedimentary rock slopes. In contrast, subsp. agourensis is found on
the northern slopes of the range on drier, exposed west- to northwest-
facing rock outcrops.

Dudleya verityi K. Nakai, also found in the Santa Monica Moun-
tains, has flowers similar to subsp. agourensis. This species differs
from subsp. agourensis by its paler flowers, several to many di-
chotomously branched stems that may elongate to more than 10 cm
long rather than the one or infrequently several cespitosely branched
stems that are mostly less than 5 cm long.

Dudleya cymosa subsp. crebrifolia Nakai & Verity, subsp. nov.

A subspecie typica caulis simplex, foliis caulis floriferi multibus
et crebribus et tardiflorentem differt. Figs. 3a,b, 5.

Caudex 1—2 cm diam. with simple to rarely few branched basal
rosettes, 5-12 cm diam., with 6-15 spreading to ascending leaves.

1987] NAKAI: DUDLE YA 345

Basal rosettes leaves mostly elliptic to spatulate, acute to acuminate,
4—10(—15) cm long, 2—5 cm wide near the middle, olive-green, rarely
glaucous, slightly maroon on the undersurface. Floral stem 10-
30(—n50) cm tall, yellowish-green, with 20-50 close-set, horizontal,
alternate to subopposite leaves. Inflorescence obpyramidal, with 2-
4 branches that branch 1-2 times; cincinnus 2-10 cm long, with 2—
15 flowers, pedicels 3-8 mm long. Petals mustard yellow, midribs
glaucous, elliptic, acute, 9-10 mm long, 3-3.5 mm wide at the mid-
dle, connate 1—-1.5 mm, apices spreading to 45°. Chromosome num-
ber: n = 17. Flowering late June to July (August).

Type: USA, CA, Los Angeles Co., San Gabriel Mtns., Fish Can-
yon, 34°11'N, 117°55'2'W, ca. 1.5 km nw. from the mouth of the
canyon on n.-facing granitic slopes, common, 400 m. Associated
with: Alnus rhombifolia, Umbellularia californica, Toxicodendron
diverilobum, Dudleya lanceolata, and D. densiflora. 25 Jun 1981, K.
Nakai 775 (Holotype: CAS; isotype: LA, MO, RSA, SD, US).

PARATYPES: Los Angeles Co., Fish Canyon, Davidson 3578 (US),
Hood 43-77k (LA), Nakai 361 (CAS, LA), 776 (CAS, LA).

Distribution. San Gabriel Mtns., Fish Canyon, 0.5—4 km from the
mouth of canyon.

Subspecies crebrifolia is distinguished by its mostly solitary basal
rosette with mostly elliptic to spatulate leaves, the large number of
cauline leaves that are often crowded, and a later flowering period.
At the higher elevations (2000 m or more), subsp. crebrifolia may
still be in flower as late as late July, but when cultivated, it flowers
from April to early June. In a letter at US to J. N. Rose (25 Jun
1923), A. Davidson suggested that this plant may be new and noted
that one plant had 13 floral stems and the basal rosette leaves were
6 inches (15 cm) long. Subspecies crebrifolia is known only in Fish
Canyon on vertical granite slopes on both walls of the canyon in
partly shaded areas.

Subspecies crebrifolia apparently is related most closely to D. cy-
mosa subsp. pumila and is well within the range of subsp. pumila.
Although its basal rosettes are often larger than those of subsp.
pumila from the San Gabriel Mountains, the size is within the overall
limit of subsp. pumila when plants from the entire geographic range
of the latter are compared. Subspecies crebrifolia differs from subsp.
pumila by its elliptic leaves, longer floral stem with 2-3 times the
number of cauline leaves that are often crowded together, and a
flowering period that is usually 4-6 weeks later. Subspecies crebri-
folia occurs from 350-600 m, whereas subsp. pumila, in the San
Gabriel Mountains, occurs below 750 m only in the northern portion
of the range [Elizabeth Lake Cyn., elev. 675 m, Nakai 1015 (LA);
elev. 610 m, Nakai 1016 (LA)].

In Fish Canyon, D. lanceolata is common and occasionally grows
sympatrically with subsp. crebrifolia. Dudleya lanceolata is similar

346 MADRONO [Vol. 34

to subsp. crebrifolia in basal rosette size, length of the floral stem,
and petal color. Infrequently, the number of cauline leaves in D.
lanceolata is similar to those of subsp. crebrifolia. Dudleya lanceolata
differs from subsp. crebrifolia in its lanceolate, often glaucous, leaves,
a more lax inflorescence, and an earlier flowering period. Dudleya
lanceolata is tetraploid (n = 34), whereas subsp. crebrifolia is a dip-
loid.

DUDLEYA ABRAMSII Rose subsp. ABRAMSII— Dudleya abramsii Rose
in Britton & Rose, Bull. N.Y. Bot. Gard. 3:14. 1903.—Cotyledon
abramsii Fedde, Bot. Jahresber. Just. 31:826. 1904.—Echeveria
abramsii Berger in Engler & Prantl, Nat. Pflanzenfam., ed. 2,
18a:477. 1930.-TypE: USA, CA, San Diego Co., wet crevices
of rocks west of Jacumba, | Jun 1903, Leroy Abrams 3707
(Holotype: US!, photo LA!; isotype: DS!, GH!, NY!, POM!, UC!,
US!)).

Caudex diameter 1-3 cm thick, mostly less than 5 cm long, un-
branched or 5—20(—75) cespitose branches. Basal rosette 2—8(—15)
cm diam. consisting of 8-25 erect or ascending oblong-lanceolate,
acute to acuminate, glaucous leaves, apiculate, 1-10 cm long, 3—15
mm wide, 1-4 mm diam., plane or concave ventrally, rounded dor-
sally, the base 5—15 mm wide. Floral stem 2—25 cm tall, 1-5 mm
diam. Cauline leaves 3—20, ascending, triangular-lanceolate, acute,
the lowermost 0.5—3 cm long, 2—11 mm wide. Inflorescence of 2-3
simple to once or twice bifurcate ascending branches or obpyramidal.
Cincinnus 3-15 cm long, with 2—10(—20) flowers. Pedicels erect, 0.5—
5(—11) mm long. Calyx 3-5 mm wide, 3-6 mm high, subtruncate to
tapered below, the segments triangular-ovate to lanceolate, acute,
2-5 mm long, |1.5-3 mm wide. Petals pale yellow, often with red
vertical lines particularly along the midrib, elliptic to narrow lan-
ceolate, acute, occasionally erose, erect with petal apices often
spreading to 90°, 8-13 mm long, 2—3 mm wide, connate 1—4.5 mm.
Epipetalous filaments 2.5—6 mm long, adnate 1—3 mm; antesepalous
filaments 4.5-8 mm long, adnate 2-4 mm; anthers yellow, 1-1.5
mm long. Gynoecium erect, slender, attenuate, 4-7 mm high, ovary
3-5.5 mm long, styles 0.5—2 mm long. Nectaries reniform, pale
yellow, 0.5-1 mm long. Chromosome number: n = 17. Flowering
April to June.

Distribution. USA, CA, Riverside Co., San Jacinto Mtns., San
Diego Co., Laguna Mtns.; MEXICO, Baja California Norte, Sierra
Juarez and Sierra San Perdo Martir. Elev. 750-1750 m.

The label data from the type specimens varies, and thus, the exact
type locality is uncertain. The holotype and an isotype specimen at
NY, both of which are a plant cultivated by Rose, are labeled “‘five
miles [6.7 km] west of Jacumba’’. The specimen at POM, however,

1987] NAKAI: DUDLEYA 347

is labeled ‘“‘Walker’s Ranch, near Jacumba’’. Others, including an
isotype at US, are labeled “‘two miles [2.7 km] west of Jacumba’’.
Because Abrams spent most of his academic career at Stanford Uni-
versity, one would suspect that the specimen at DS would have an
accurate label. I was able to collect D. abramsii 2 mi west of Jacumba,
but I was unable to find it 5 mi west of Jacumba.

Moran (1951) listed two populations of D. abramsii from the San
Jacinto Mountains, one from near Kenworthy [Munz 5788 (POM)]
and the other from Taquitz (sic) Ridge, 9000 ft (2770 m) [Jaeger in
1921 (POM)]. The population near Kenworthy is D. abramsii, but
differs from typical D. abramsii by lacking the characteristic red
striations along the petal midrib. The specimen from Tahquitz Ridge
differs considerably from the Kenworthy population in its compar-
atively broad, oblong to ovate leaves and short floral stem. It re-
sembles no specimens of D. abramsii that I have studied. The ‘‘Tah-
quitz’’ specimen, however, does resemble plants of D. cvmosa subsp.
pumila. A recent collection made near Tahquitz Ridge by J. Catlin
[Lily Rock, near Idlywild, 2300 m, Nakai 984 (CAS, LA)] has reddish
flowers and is similar to the ““Tahquitz” specimen. Catlin (pers.
comm.) reports similar plants on Suicide Rock, across Strawberry
Valley from Lily Rock, and along Snow Creek on the north slope
of San Jacinto Peak. The status of these populations is uncertain
until more material can be studied, but it appears they are not D.
abramsii.

Dudleya abramsii subsp. calcicola (Bartel & Shevock) K. Nakai,
comb. nov. — Dudleya calcicola Bartel & Shevock, Madrono 30:
210. 1983.—TypeE: USA, CA, Tulare Co., Kern River at Roads
End, T23S R32E S13, Sequoia National Forest, 1200 m, 11 Jul
1981, Shevock 8802 (Holotype: CAS!, photo LA!; isotype: FSC,
NY!, RSA!, SBBG, SD!, UC).

Distribution. USA, CA, southern Sierra Nevada from the Rincon
area south of Durrwood Creek in Tulare Co. to the southern Piute
Mtns. in Kern Co. Elev. 500-1550 m.

Collections I have made from the Piute Mountains, the vicinity
of Lake Isabella, and along the Kern River suggest that D. calcicola
is closer to D. abramsii than previously believed. Bartel and Shevock
(1983) suggested that D. calcicola was intermediate between D.
abramsii and D. cymosa, but closer to D. abramsii. They noted that
D. calcicola was distinct from D. abramsii in its 1) heavier foliar
bloom, 2) obpyramidal inflorescence with a thicker floral stem and
spreading cincinni versus a comparatively simple inflorescence and
ascending cincinni, 3) slightly longer pedicel, 4) pale yellow petals
unmarked with red, and 5) occurrence predominately on limestone.

The type locality of D. calcicola is the limestone outcrops above

348 MADRONO [Vol. 34

Roads End along the Kern River in Tulare Co., but I have observed
plants resembling D. calcicola at a number of localities along the
Kern River on substrates other than limestone. For example, one
population was found less than 0.5 km north of the type locality on
metamorphic rock. Pedicel length from cultivated plants from 12
populations of D. abramsii and 13 populations of D. calcicola ex-
hibited no differences. They measured 3-7 mm (X = 5 mm) for D.
abramsii and 3—8 mm, (X = 5 mm), for D. calcicola. Measurements
made from herbarium specimens, however, ranged from 2-10 mm
(X = 5 mm), for D. abramsii and from 2-12 mm (X = 6 mm) for
D. calcicola. Although several populations of D. calcicola do have
flowers with plain, pale yellow petals, others have the characteristic
D. abramsii red striations along the petal midrib. Cultivated plants
I collected from the type locality and observed for five flowering
seasons had flowers with petals conspicuously marked with red. In
at least three populations of D. abramsii [MEXICO, Baja California
Norte, Cerra Blanco, Moran 17608 (SD); 2 km w. of Rancho Santa
Cruz, Sierra San Pedro Martir, Moran 23461 (SD), Nakai and Prigge
1136 (CAS, LA); USA, CA, Riverside Co., Kenworthy, Munz 5788
(POM, SD), Nakai 1007 (CAS, LA)] the petals lack red pigment.
The density of the foliar bloom does not appear to differ between
the two taxa.

Typical D. calcicola is a densely-packed plant with up to 50 ro-
settes. Bartel and Shevock (1983), however, cite populations (e.g.,
Long Canyon) that have plants with one to a few rosettes. In the
Laguna Mountains of San Diego Co., the higher elevation popula-
tions of D. abramsii also have plants with 50 or more rosettes, and
a population of a D. abramsii subsp. affinis K. Nakai in Cushenbury
Canyon has densely-packed plants with up to 50 rosettes. Thus, the
number of rosettes per plant may not differ between the taxa.

An important character in which D. calcicola is similar to D.
abramsii is the relative lengths of the antesepalous and epipetalous
stamens. In D. cymosa, the difference in staminal length is often
small (<0.5 mm). In both D. abramsii and D. calcicola, the difference
is usually 1-1.5 mm. This strongly supports a close relationship
between D. calcicola and D. abramsii.

Another character that distinguishes D. calcicola from D. abramsii
is the inflorescence. Although the inflorescence of D. calcicola found
in the wild may be similar to D. abramsii, cultivated plants consis-
tently have an obpyramidal inflorescence consisting of 2—4 branches
that bifurcate once or twice. Cultivated plants of D. abramsii have
a simpler inflorescence of 2—3 mostly simple branches. Cultivated
D. calcicola tends to have more cauline leaves (8-15) in comparison
to D. abramsii (2-8).

Although there are enough differences to warrant taxonomic rec-
ognition, the two taxa have considerable overlap in most of the key

1987] NAKAI: DUDLEYA 349

characters. Of additional interest are two collections from the geo-
graphic range of D. calcicola that were identified by authorities as
D. abramsii: one from the Tehachapi Mtns. [May 1925, Davidson
3599 (US)], annotated by Rose; another collected by J. Zavinowich
from Jawbone Canyon on the east slope of the Piute Mtns. [Moran
24196 (SD)], determined by Moran. These two authorities on Dud-
leya apparently also recognized the resemblance of this material to
D. abramsii. Thus, I proposed the combination D. abramsii subsp.
calcicola.

Dudleya abramsii subsp. affinis K. Nakai, subsp. nov.

A subspecie typica caudice simplicis, foliis rosulae oblanceolatis
vel ellipticis differt. Figs. 4a,b, 5.

Plants simple, rarely cespitosely branched. Basal rosette 3-6 cm
diam., of 10—25 oblanceolate to elliptic, glaucous leaves, 2—4 cm
long, 7-15 mm wide; apices acute to acuminate. Cauline leaves
lanceolate, glaucous, 5-6 mm long, 2-3 mm wide, acute. Floral stem
erect, 5-11 cm long, pale yellow to glaucous. Inflorescence of 2-3
mostly simple branches, cincinnus ascending, 2—8 cm long, with 3-
8 flowers, lowermost pedicels 2.5-8 mm long. Petals pale yellow
with red striations along the midribs, connate 1.5—2.5 mm, apices
spreading to 90°. Epipetalous filaments 2.5—4 mm long, antesepalous
filaments 3.5-6 mm long. Chromosome number: n = 17. Flowering
May to July.

Type: USA, CA, San Bernardino Co., San Bernardino Mtns., Green
Canyon, ca. 0.5 km sw. of National Forest Road 2N93 on trail to
Sugarloaf Mountain where the trail crosses the creek, 34°13'N,
116°48’'W, on e. side of creek on granite; common; 2600 m; with
Juniperus, Pinus, Cercocarpus, Echinocerus. 28 Jun 1984, K. Nakai
1146 (Holotype: CAS; isotype: LA, NY).

PARATYPES: USA, CA, San Bernardino Co., confluence of Deep
and Hook creeks, Nakai 1110 (CAS, LA), 1114 (CAS, LA, RSA);
Holcomb Creek, 8 km sw. of Big Pine Flat, Nakai 1153 (LA); nw.
slope of Gold Mountain, 3.2 km w. of Big Bear refuse dump, Nakai
1151 (CAS, LA); n. shore of Lake Baldwin, D. B. Stark 4992 (RSA),
Nakai 702 (CAS, LA); Johnson Grade, Peirson 8972 (POM, UC),
Peirce s.n. (POM), Moran 2193 (UC), Nakai 1145 (CAS, LA); ridge
e. of Lake Baldwin, Munz 10494 (POM); plateau s. of Lake Baldwin,
Peirson s.n. (RSA); Cushenbury Canyon, Deburg 2608 (RSA), Nakai
1147 (CAS, LA); Cushenbury Spring, Parish 1629 (POM); Green
Canyon, Clausen & Trapido 4770 (CU, NY, US, WTU), Nakai 1113
(CAS, LA).

Distribution. North slopes of the San Bernardino Mtns. from 1800-
2600 m on granite, quartzite, or, rarely, limestone.
Dudleya abramsii subsp. affinis differs from subsp. abramsii by

350 MADRONO [Vol. 34

its mostly unbranched caudex and basal rosette leaves that are ob-
lanceolate to elliptic rather than oblong-lanceolate. Some plants of
subsp. affinis resemble D. cymosa subsp. pumila, particularly in
rosette size, leaf shape, and, in more robust plants, the inflorescence.
The most consistent differences are a shorter pedicel length, pale
yellow petals with red striations, and the difference in length between
the antesepalous and epipetalous stamens.

Plants from Cushenbury Canyon grow on limestone and differ
from typical plants of subsp. affinis by their several to many branched
caudex and smaller rosettes, 1.5—3 cm in diameter. This population
seems similar to subsp. calcicola and subsp. abramsii. It differs from
subsp. calcicola by its more simple inflorescence and from subsp.
abramsii by its rosette leaf shape. Although this population is in-
termediate between these three subspecies of D. abramsii, I presently
consider this population an aberrant form of subsp. a/ffinis.

KEY TO SPECIES AND SUBSPECIES

A. Pedicels 5—20 mm long; petals connate 1—2.5 mm; the difference
between epipetalous and antesepalous staminal length is usually
<0.5 mm.

B. Basal rosette leaves evergreen, 2—17 cm long, 0.5—6 cm wide;
caudex more than | cm diam.
C. Floral stem mostly 1.5—4.5 dm tall.
D. Floral stem with 20—50 close-set leaves; basal rosette
leaves elliptic to spatulate. San Gabriel Mtns., s. Cal-
IORI e eee D. cymosa subsp. crebrifolia
D. Floral stem usually with <20 leaves; basal rosette
leaves oblong-oblanceolate to oblong-triangular, rarely
spatulate. Central and n. California.
E. Petals bright yellow to red; rosette leaves oblong-
oblanceolate, rarely spatulate, 1-6 cm wide. Coast
Range from the Salinas River, Santa Clara Co.,
n. to Humboldt Co.; Sierra Nevada ...........
eee D. cymosa subsp. cymosa
E. Petals pale yellow.
F. Basal rosette leaves oblong to oblanceolate; in-
florescence of 2—3 bifurcate branches; pedicels
6-12 mm long. Inner South Coast Range from
Contra Costa Co., to w. Fresno and ne. Mon-
terey COs., on various rock substrates .......
ee eT D. cymosa subsp. paniculata
F. Basal rosette leaves oblong-triangular; inflo-
rescence of 2—3 simple branches; pedicels 4—7
mm long. Santa Clara Valley on serpentine
eon d ys Oat ene D. cymosa subsp. setchellii

1987] NAKAI: DUDLE YA 351

C. Floral stem mostly <1.5 dm tall. Outer South Coast Range
from the Salinas River s. to s. California.
G. Basal rosette leaves oblanceolate to spatulate, usually
10-25, mostly short acuminate to cuspidate. Outer
South Coast Range to San Gabriel and San Bernar-
Gin@ MtHS: 2af.4200 a, D. cymosa subsp. pumila
G. Basal rosette leaves oblong to elliptic or ovate, usually
6-10, acute to acuminate.
H. Basal rosette leaves ovate, green, often with a
maroon suffusion on the underside; caudex un-
branched. Santa Monica and Santa Ana mtns. .
decd amet nees Gliese te AIL D. cymosa subsp. ovatifolia
H. Basal rosette leaves oblong to elliptic, glaucous;
caudex simple or few to, rarely, several branches.
Santa Monica Mtns. .................0....-.
Ce ye ere eee D. cymosa subsp. agourensis
B. Basal rosette leaves withering in summer, 1.5—4 cm long, 5—
12 mm wide; caudex <1 cm diam. .....................
eat eee ee D. cymosa subsp. marcescens
A. Pedicels 0.5—7 mm long; petals connate 1.5—4.5 mm; the differ-
ence between the epipetalous and antesepalous staminal length
is usually 1-1.5 mm.
I. Basal rosette leaves oblong to oblong-lanceolate; plants with
few to many branches.

J. Inflorescence of 2—3 mostly simple branches; pedicels
mostly <5S mm long; petals usually with red striations
along the midribs.

K. Lower cauline leaves usually <15 mm long; floral stem
2-15 cm tall; petals connate 2—4.5 mm. San Jacinto
Mtns., Riverside Co., and Laguna Mtns., San Diego
Co.; Sierra Juarez and Sierra San Pedro Martir, Baja
California Norte ...... D. abramsii subsp. abramsii

K. Lower cauline leaves 10-30 mm long; floral stem 5-
25 cm tall; petals connate 1.5—3 mm. San Luis Obispo
CC Fae ne ne ee a D. abramsii subsp. murina

J. Inflorescence of 2—3 simple to usually bifurcate branches;
pedicels 3—8 mm long; petals with or without red striations
along the midribs. Southern Sierra Nevada ............

AGT CTP Oe re etn Te re D. abramsii subsp. calcicola

I. Basal rosette leaves elliptic to oblanceolate; plants usually
unbranched. San Bernardino Mtns., San Bernardino Co.

eh eae ee oe D. abramsii subsp. affinis

Additional specimens of Dudleya that were cultivated and examined but not cited
in the text. Collection numbers, unless otherwise noted, are the author’s.
DUDLEYA ABRAMSII: subsp. ABRAMSII— USA, CA, Riverside Co.: San Jacinto Mtns.,

352 MADRONO [Vol. 34

near Kenworthy, 1059. San Diego Co.: Mt. Laguna, 454; Laguna Mtns., Kwaaymit
Pt., 1180; Descanso Junction, 845; Kitchen Creek, 844; Corte Madera Lake, Van
Der Werff s.n., Campo 840; Jacumba, 841; Dubber Spur, 842. MEX, Baja Calif.
Norte: Sierra Juarez, near Laguna Hansen, Prigge 5098; n. of Valle Trinidad, Verity
s.n.; Sierra San Pedro Martir, between Mike Sky Ranch and El Burro, Verity and
Prigge s.n.; w. of Rancho Santa Cruz, Nakai and Prigge 1136. Subsp. CALCICOLA—
USA, CA, Tulare Co.: 2 km n. of Road’s End, 1080; Road’s End, 825; Hospital Flat
Campground, ca. 9.5 km n. of Kernville, 7078. Kern Co.: 2.0 km s. of Kernville,
1077; 8 km s. of Kernville, 827; Long Canyon, 828; near Mountain Mesa, 829;
Bodfish Cyn., 830; s. of Bodfish, 83/7; near Twin Oaks, 678; near Loraine, 679, 680;
Caliente, 832; Cottonwood Creek, near Kelso Valley, 676, 677.

DUDLEYA CYMOSA: subsp. CyMoOsA— USA, CA, Mendocino Co.: Hopland, McCabe
356. Sonoma Co.: near Mt. St. Helena, A/meda s.n.; Cazadero, 973. Marin Co.: Bolinas
Bay, 975; Stinson Beach 398; s. of Stinson Beach, 398; Mt. Talmapais, 399. Solano
Co.: s. of American Cyn. Rd., 827; Mix Cyn., Vaca Mtns., 531. Santa Cruz Co.: Eagle
Rock, McCabe 369. Santa Clara Co.: Lexington Reservoir, 8/4; New Almaden, 8/4;
Loma Prieta, 8/3; Stevens Creek, 1088. Tehama Co.: near Paynes Creek, 96/. Sierra
Co.: w. of Downieville, McCabe 507. Nevada Co.: near Nevada City, 95/. Placer
Co.: Applegate, 950. Amador Co.: near Ione, 532; w. of Volcano, 394; Volcano, 534;
near Mokelumne River, 535. Calaveras Co.: near San Andreas, 536; near Kentucky
House, 393, 537. Tuolumne Co.: Table Mtn., 1087; near Coulterville, 538. Mariposa
Co.: near Yosemite Valley, 1/084, 1085; near Bear Valley, 539, 822. Inyo Co.: Sawmill
Creek, 1081. Tulare Co.: near Springville, 392; w. of Pierpoint, 823. Kern Co.: Bear
Hollow Creek, 1075; Shirley Creek, 1076. Subsp. PANICULATA— Contra Costa Co.: n.
slope of Mt. Diablo, 949; Mt. Diablo, 396; s. slope Mt. Diablo, 948. Alameda Co.:
Palomare Rd., 820; Welch Creek, 8/9. Santa Clara Co.: Coyote Reservoir, 8/8; Alum
Rock, 817; Mt. Hamilton, 8/6; Anderson Reservoir, 977; Coyote Lake, 978. Stan-
islaus Co.: Arroyo del Puerto, 946, 947. Merced Co.: Pacheco Pass, 945. San Benito
Co.: Pinnacles, 809; near Panoche Pass, 944; Clear Creek, 942. Monterey Co.: Lewis
Creek, 808; Lorenzo Creek, 628; Bull Cyn., 1092. Fresno Co.: Coalinga Hot Spr.,
McCabe 501. Subsp. PUMILA— Monterey Co.: Pine Valley, 526; n. of Castro Cyn.,
631; s. of Castro Cyn., 632; n. of Mission San Antonio, 523, 524; Nacimiento-
Fergusson Rd., 100 m, 4/0; Nacimiento-Fergusson Rd., 700 m, 409, 633. Santa
Barbara Co.: Santa Barbara Cyn., 788; Rattlesnake Cyn., 17/00. Ventura Co.: conflu-
ence of Potrero John and Sespe creeks, 544; Sespe Gorge, 545, 789. Los Angeles Co.:
Elizabeth Lake Cyn., 825 m, 1013; 790 m, 1014; 670 m, 1015; 610 m, 1016; Arroyo
Seco, 915 m, 430; 850 m, 553; 1070 m, 555; Hidden Springs, 383, 427; San Gabriel
Cyn., 1500 m, 7/25; 1600 m, 1126; 1700 m, 1/02]. San Bernardino Co.: San Antonio
Falls, 1900 m, 689; Waterman Cyn., 386, 1183; s. of Crestline 1/84; Miller Cyn.,
1185: Little Mill Creek, 700; Keller Mtn., 702, 1143; Skinner Creek, 1148.

ACKNOWLEDGMENTS

I thank Barry Prigge, David Verity, Geoff Levin, and Jim Dice for reviewing an
early draft of the manuscript and for their helpful comments. Wayne Ferren, Jim
Bartel, James Shevock, and an anonymous reviewer also provided many useful com-
ments and helpful criticism.

LITERATURE CITED

BARTEL, J. and J. SHEVocK. 1983. Dudleya calcicola (Crassulaceae), a new species
from the southern Sierra Nevada. Madrono 30:210-216.

LEMAIRE, C. A. 1858. Revue Hortic. 7:439.

Moran, R. V. 1951. A revision of Dudleya. Ph.D. dissertation, Univ. California,
Berkeley.

1957. Innovations in Dudleya. Madrono 14:106-108.

1987] NAKAI: DUDLEYA Se:

. 1960. Dudleya. In H. Jacobsen, A handbook of succulent plants, p. 344—
359. Blandsford Press, U.K.

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

1974. A flora of southern California. Univ. California Press, Berkeley.

NAKAI, K. M. 1983. A new species and hybrid of Dudleya (Crassulaceae) from the
Santa Monica Mountains, California. Cact. Succ. J. 55:196-200.

Rose, J. N. 1903. Jn N. L. Britton and J. N. Rose, New or noteworthy North
American Crassulaceae. Bull. N.Y. Bot. Gard. 3:1-45.

Unt, C. H. and R. V. Moran. 1953. Cytotaxonomy of Dudleya and Hasseanthus.
Amer. J. Bot. 40:492-501.

(Received 6 Jan 1986; revision accepted 5 May 1987.)

ANNOUNCEMENT

BOTANICAL COLLECTIONS IN MICROFICHE FORM AT UCSB

The UCSB Library and the UCSB Herbarium are pleased to announce
the purchase of type and special botanical collections, in microfiche
form, from Meckler Publishing. Three collections were acquired through
the UC Shared Acquisition Program, which is a University-wide library
effort designed to facilitate sharing of unique or expensive materials
among the nine UC campuses and Stanford University. The collections
include: 1) New York Botanical Garden Vascular Plant Type Collections;
2) Vascular Plant Types and Early Authentic Specimens of the Academy
of Natural Sciences of Philadelphia; and 3) United States National Her-
barium Smithsonian Institution Vascular Plant Types. Printed indices
have been purchased for each campus to help users locate individual
specimens. The microfiche are housed in a separate viewing room in
the UCSB Library and can be borrowed through interlibrary loan agree-
ments. In addition to those collections purchased through Shared Ac-
quisitions, the UCSB Library has purchased the California Academy of

Sciences Plant Type Collection in microfiche form. We encourage all
interested parties to make use of these important and accessible re-
sources.

ANNOUNCEMENT

NEw PUBLICATION

Jepson Globe: A Newsletter from the Friends of the Jepson Herbarium,
vol. 1, no. 1, pp. 1-4, 1987, no ISSN, subscription with contribution
of $15.00 or higher (from Friends of the Jepson Herbarium, Dept.
Botany, Univ. of California, Berkeley, CA 94720). [With message by
G. L. Stebbins and 2 articles: J. H. Thomas on history of herbaria, pt.
1; J. C. Hickman on status of Jepson Manual Project (text and illus.
15% complete, some 150 collaborators, 3 paid staff, and many volun-
teers, notably Emily Reid, scientific illustrator). ]

CYMOPHORA (ASTERACEAE: HELIANTHEAE)
RETURNED TO TRIDAX

DAVID J. KEIL
Biological Sciences Department,
California Polytechnic State University,
San Luis Obispo 93407

MELISSA A. LUCKOW
Department of Botany, The University of Texas,
Austin 78713

DONALD J. PINKAVA
Department of Botany and Microbiology,
Arizona State University, Tempe 85287

ABSTRACT

Cytological and morphological evidence supports the merger of Cymophora B. L.
Robinson with Tridax L. (Asteraceae: Heliantheae). Chromosome counts of n = 9
are reported for Tridax accedens Blake and the closely related 7. dubia Rose. Tridax
hintonii (Turner & Powell) Keil, Luckow & Pinkava, comb. nov., is proposed, based
on Cymophora hintonii Turner & Powell.

Our chromosome count of n = 9 for Tridax accedens Blake (As-
teraceae: Heliantheae), the first report for this species, provides key
information in an ongoing taxonomic controversy. During the past
20 years several researchers have discussed the status of Cymophora
and its relationship to 7ridax (Anderson and Beaman 1968, Turner
et al. 1973, Turner and Powell 1977, Canne 1977, 1978, 1983,
Robinson et al. 1981, McVaugh 1984). Anderson and Beaman noted
numerous morphological similarities between C. pringlei B. L. Ro-
bins. (at the time the only species of Cymophora) and two species
of Tridax (T. accedens and T. dubia) and concluded that Cymophora
could not be maintained separate from Tridax. They noted that C.
pringlei differs mainly from the two Tridax species in having smaller,
fewer-flowered heads and epappose achenes. It is particularly similar
to T. accedens. They transferred C. pringlei into Tridax as T. oli-
gantha Anderson & Beaman.

Turner et al. (1973) published a chromosome number of Zn = 16
(counted by Robert Irving) for 7. oligantha and questioned the re-
lationship of this species to Tridax (x = 9, 10). They suggested instead
a relationship of Cymophora to Galinsoga (x = 8) and Sabazia (x =
4). They further suggested that a chromosome count for 7. accedens
would be helpful in evaluating the relationship of 7. oligantha.

MADRONO, Vol. 34, No. 4, pp. 354-358, 1987

1987] KEIL ET AL.: CY MOPHORA RETURNED TO TRIDAX 6p)

Turner and Powell (1977) reinstated Cymophora (as a genus distinct
from Tridax), transferred 7. accedens to it and described a third
species, C. hintonii Turner & Powell. They discounted the purported
relationship of 7. accedens to T. dubia (Blake 1943, Powell 1965,
Anderson and Beaman 1968), but provided neither a key nor dis-
cussed the morphological differences between the two genera.
Canne (1977) noted that Tridax venezuelensis Arist. & Cuatr.,
which bears features of both Galinsoga and Tridax, is morpholog-
ically most similar to species placed by Turner and Powell into
Cymophora, and transferred this species into Cymophora. She fur-
ther noted that the four species of Cymophora fall into two well-
defined morphological species groups characterized by differences
in leaf shape, petiole length and the number of veins in the phyllaries
and pales. Cymophora hintonii and C. venezuelensis form one group,
and C. accedens and C. pringlei the other. Robinson et al. (1981)
again noted the similarity of 7. dubia to Cymophora, but questioned
the transfer of 7. venezuelensis into Cymophora, suggesting that it
represented a different phyletic line than the remainder of Cymo-
phora. They reported an approximate count of 2” = ca. 18 for T.
venezuelensis. Robinson (1981) listed Cymophora distinct from Tri-
dax, but did not discuss its relationships or composition. Canne
(1983) reported n = 9 for C. hintonii which further weakened the
chromosomal basis for distinguishing Cymophora from Tridax.
Two types of evidence have been used to date in studies of the
two genera: morphology and chromosome numbers. Anderson and
Beaman (1968) and McVaugh (1984) used morphological evidence
to support union of Cymophora with Tridax. Turner and Powell
(1977) and Canne (1977, 1978) used a combination of morphological
and cytological data to support their separation of the genera. A
review of the conflicting sources of evidence is presented below.

MORPHOLOGICAL EVIDENCE

Habit and vegetative morphology cannot be used to separate the
two genera. Both genera are composed of opposite leaved herbs. All
of the species that have been assigned to Cymophora are taprooted
annuals. Eleven of the 25 species of 7ridax (s. str.) are annuals,
including 7. dubia, a species considered to be a link between Tridax
and Cymophora by Anderson and Beaman (1968). Turner and Pow-
ell (1977) considered 7. dubia to be “‘a true Tridax’’. Canne (1978)
listed the following features as distinctive of Cymophora: paniculate-
cymose capitulescence, zygomorphic outer disc corollas and white
corollas. In a tabular comparison of several genera of the Galinso-
ginae she described the capitulescences of Tridax as “heads solitary
or in few-headed subcymes”’ and those of Cymophora as “‘heads in
several- to many-headed cymose panicles’’. Although some Tridax

356 MADRONO [Vol. 34

species have solitary heads, 7. dubia and several other species (e.g.,
T. platyphylla) have many-headed cymose panicles. The capitules-
cences of 7. dubia and Cymophora accedens are similar in appear-
ance and in number and distribution of heads and were used as
evidence of the relationship of these taxa and C. pringlei by Anderson
and Beaman (1968).

The heads and the included bracts and florets of Cymophora species
are smaller than those of most 7ridax. However, T. dubia has flowers
and bracts similar in size to those of Cymophora species. The in-
volucral bracts of Tridax are 2-5 seriate and those of Cymophora
are 1-3 seriate. The species of 7ridax most similar to Cymophora
have involucral bracts of similar number and form (Anderson and
Beaman 1968). The outer florets of both Cymophora and Tridax
heads tend to be zygomorphic. In most Tridax species, these flowers
are pistillate and have an evident ligule (anterior lip) and are con-
sidered to be rays even though a small posterior lip is present. Rays
are absent in several species of Tridax and in T. bilabiata the disc
florets are bilabiate. In three Cymophora species, the outer florets
are perfect and bilabiate with a short anterior lip and are considered
to be bilabiate disc florets (Anderson and Beaman 1968, Turner and
Powell 1977, Canne 1978). The fourth species, C. venezuelensis, has
pistillate, bilabiate outer florets that are treated as rays (Canne, 1977).
The remaining disc florets are mostly actinomorphic or nearly so in
both genera. Disc corollas in Tridax species vary from creamy yellow
to bright yellow or yellow-green. Those of Cymophora are creamy
white.

In Cymophora, the achenes may bear short fimbriate or plumose
scales or may be epappose. The pappus of most TJridax species
consists of slender plumose or fimbriate scales or bristles. The pap-
pus of 7. dubia, however, is similar to that of Cymophora, which
consists of short, fimbriate-margined scales.

CYTOLOGICAL EVIDENCE

Our count of m = 9 for C. accedens indicates that two (and possibly
three) of the Cymophora species share a base of x = 9. Both mor-
phological groups recognized by Canne contain species with this
base number, as does Tridax (including 7. dubia). In addition, it is
possible that the single reported count of 2 = 16 may be inaccurate.
Irving, as listed by Turner et al. (1973), reported a chromosome
count of 2” = 16 from mitotic material. Turner (pers. comm.) notes
that a crude penciled camera lucida drawing attached to the voucher
specimen (seeds of which served as the source material) does suggest
a number of 2” = 16, but some of the chromosomes may be un-
resolved and, thus, a count of 2” = 18 might still hold.

Tridax is dibasic with x = 9 and 10. The species of 7ridax most
similar to Cymophora have n = 9.

1987] KEIL ET AL.: CV MOPHORA RETURNED TO TRIDAX So,

&
oO
aed

Fics. 1, 2. Camera lucida drawings of chromosomes at diakinesis. 1. Tridax
accedens. 2. Tridax dubia.

CONCLUSIONS

We propose here that continued recognition of Cymophora as a
genus distinct from 7ridax is not supported by either morphological
or cytological evidence. The closest relatives of Cymophora appear
to be species of 7ridax and the morphological characters that sep-
arate the genera are weak if they exist at all. Of the characters listed
by Canne (1978), only the color of the disc corollas seems to stand
up to scrutiny. The cymose paniculate capitulescence of Cymophora
species 1s fundamentally similar to the capitulescences of several
Tridax species. Rayless heads with bilabiate outer disc florets occur
in T. bilabiata and in Cymophora species. We agree with those who
consider 7. dubia and T. accedens to be closely related, and, there-
fore, arrive at the same conclusion as Anderson and Beaman (1968).
We suggest that all four species of Cymophora be returned to Tridax.

McVaugh (1984) arrived at a similar conclusion. Noting the lack
of morphological differences between Cymophora and Tridax, he
placed C. accedens back into Tridax. He did not propose, however,
a combination in Tridax for C. hintonii, but merely listed it (as
Cymophora hintonii) among the species of 7ridax in Nueva Galicia.
We, therefore, propose the following combination:

Tridax hintonii (Turner & Powell) Keil, Luckow & Pinkava, comb.
nov.—Cymophora hintonii Turner & Powell, Madrono 24:2.
LOTT:

We report the following chromosome counts and the voucher
specimens that document them:

Tridax accedens S. F. Blake. 2n = 9,, (Fig. 1). MEXICO: Colima:
Hwy. 110, 17 mi ne. of jet. with Hwy. 200, Keil and Luckow
15139 (OBI).

Tridax dubia Rose. 2n = 9, (Fig. 2). MEXICO: Jalisco: 8 min. of
El Tuito, Keil and Luckow 15112 (OBI).

358 MADRONO [Vol. 34

ACKNOWLEDGMENTS

We thank Judith M. Canne for verifying determinations of Cymophora accedens
and Tridax dubia, and B. L. Turner and John L. Strother for comments on an earlier
version of our manuscript. Field work was supported by NSF Grant DEB 81-04683.

LITERATURE CITED

ANDERSON, C. E. and J. H. BEAMAN. 1968. Status of the genus Cymophora (Com-
positae). Rhodora 70:241-246.

BLAKE, S. F. 1943. Ten new American Asteraceae. J. Washington Acad. Sci. 33:
265-272.

CANNE, J. M. 1977. A new combination in Cymophora (Compositae: Heliantheae:
Galinsoginae). Madrono 24:190-191.

. 1978. Circumscription and generic relationships of Galinsoga (Compositae:

Heliantheae). Madrono 25:81-93.

. 1983. Cytological and morphological observations in Galinsoga and related
genera (Asteraceae). Rhodora 85:355-366.

McVauau, R. 1984. Flora Novo-Galiciana. Vol. 12. Compositae. Univ. Michigan
Press, Ann Arbor.

PoweELL, A. M. 1965. Taxonomy of 7ridax (Compositae). Brittonia 17:47-96.

ROBINSON, H. 1981. A revision of the tribal and subtribal limits of the Heliantheae
(Asteraceae). Smithsonian Contr. Bot. 51:1-102.

, A. M. PowELL, R. M. KING, and J. F. WEEDIN. 1981. Chromosome numbers
in Compositae, XII: Heliantheae. Smithsonian Contrib. Bot. 52:1-28.

TURNER, B. L. and A. M. PoweLi_. 1977. Taxonomy of the genus Cymophora
(Asteraceae: Heliantheae). Madrono 24:1-6.

, , and T. J. WATSON. 1973. Chromosome numbers in Mexican As-

teraceae. Amer. J. Bot. 60:592-596.

(Received 5 Dec 1986; revision accepted 16 Jul 1987.)

NEW MADRONO EDITOR

The Executive Council of the California Botanical Society is pleased
to announce the appointment of Dr. David J. Keil to the position of
Editor of Madrono. Dr. Keil, Professor of Botany, is the Director of the
Robert F. Hoover Herbarium (OBI). His editorship will commence in
January 1988 with volume 35. All new manuscripts submitted to Ma-
drono and all returned revisions should be mailed to him at the Bio-
logical Sciences Department, California Polytechnic State University,
San Luis Obispo, CA 93407. Mr. Wayne R. Ferren, Jr., who has com-
pleted his tenure as Editor, will be appointed to the Board of Editors
to assist with continuity of journal management.

VASCULAR PLANTS OF EASTERN IMPERIAL
COUNTY, CALIFORNIA

STEVEN P. MCLAUGHLIN
Office of Arid Lands Studies, University of Arizona,
845 N. Park Ave., Tucson 85719

JANICE E. BOWERS
3949 E. Paseo Dorado, Tucson, AZ 85711

KENNETH R. F. HALL
Picacho State Recreation Area, P.O. Box 1207,
Winterhaven, CA 92283

ABSTRACT

An annotated catalogue of the vascular flora was compiled from field collections
and herbarium records from the eastern third of Imperial Co., California, including
wetland habitats along the lower Colorado River. The study area is located in the
Lower Colorado Valley subdivision of the Sonoran Desert, an arid area with little
elevational relief. The flora is correspondingly depauperate, with only 278 native
species recorded from 2050 km7?, including 56 species found only in wetland habitats
along the lower Colorado River. Ecological and floristic factors that contribute to the
low diversity include a relative lack of habitat differentiation among desert species
and the lack of a well developed herbaceous perennial flora.

The Lower Colorado Valley subdivision of the Sonoran Desert
(Shreve 1951) in eastern Imperial Co., California, is one of the driest
parts of the desert region of the southwestern United States and
northwestern Mexico. It is perhaps also the most poorly known area
botanically in California. The objective of this study was to catalogue
the flora of this area, emphasizing the desert habitats and the wetland
habitats along the lower Colorado River. This area was selected for
study because it is centered within a section of the Sonoran Desert
for which no other local floras have been compiled; the data were
used in a broader study of the floristics of the southwestern U.S.
(McLaughlin 1986).

STUDY AREA

The study area includes most of Imperial Co. east of 115°W lon-
gitude (Fig. 1). The Riverside Co. line and the International Bound-
ary form the northern and southern boundaries, respectively; the
eastern boundary is the Colorado River. Wetland habitats along both
river banks are included in the study area. The western boundary
excludes the Algodones Dune field, which has its own distinctive

MADRONO, Vol. 34, No. 4, pp. 359-378, 1987

RIVERSIDE C COUNTY Palo Verde ARIZONA

Study area
boundry

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Fic. 1. Map of study area in eastern Imperial Co., extreme southeastern corner
of California.

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 361

TABLE 1. CLIMATIC DATA FOR STATIONS ADJACENT TO STUDY AREA (WILLMOTT
ET AL. 1981).

Precipitation
(mm)
Mean monthly

temperature Sum-

(°C) Winter mer

Elev. ————-. Oct—__ Apr-

Station Lat. Long. (m) Jan Jul Mar Sep
Amos, CA 33°6' 115°13’ 78 12.4 35.2 40 21
Blythe, CA 33°30 114°36’ 82 10.2. 32.6 63 37
Brawley, CA 32°59’ 115°32’ — 36 11.4 32.3 47 15
Mexicali, B.C. 32°39’ 115°39' 4 11.8 33.0 48 23
Quartzsite, AZ 33°40’ 114°15' 147 10.1 34.0 Ta 70

Yuma, AZ 32°44! 114°37' 42 12.4 32.8 =e) 31

flora (Bowers 1984), and the Chocolate Mountains Naval Gunnery
Range, to which access is restricted.

The study area is approximately 2050 km’. Elevations range from
near sea level to 664 m at Quartz Peak in the Chocolate Mountains.
The major mountain ranges in the area include the Palo Verde
Mountains in the north, the Cargo Muchacho Mountains in the
southwest, and the Chocolate Mountains that form an irregular mass
trending northwest to southeast across the center of the area. Picacho
Peak, a distinctive landmark 29 km north of Yuma, lies in the
Chocolate Mountains.

The climate of the study area is extremely arid (Table 1). Annual
rainfall varies from about 60 mm in the southwest corner to about
100 mm in the higher elevations. Most of the rain occurs in the
winter; late summer storms in September account for most of the
rain that falls from April through October. Mean daily maximum
summer temperatures are 38—41°C; winter temperatures seldom drop
below freezing.

VEGETATION AND HABITATS
Deserts

The vegetation of the Lower Colorado Valley subdivision of the
Sonoran Desert has been described by Shreve (1925, 1951) and
Turner and Brown (1982). Because of the extreme aridity of the area,
the vegetation is remarkably monotonous. Undisturbed sections of
the desert on mountain slopes, bajadas, and sandy flats are all dom-
inated by Larrea tridentata—Ambrosia dumosa microphyllous desert
(Shreve 1951). The low-lying, fine-textured soils of the Colorado
River floodplains that are currently under cultivation—the eastern

362 MADRONO [Vol. 34

half of the Quechan (formerly Ft. Yuma) Indian Reservation and
the Palo Verde Valley—were probably once extensive Atriplex flats
(Turner and Brown 1982).

Mountains, slopes, and flats. A group of about a dozen woody and
succulent species are dominant on mountain tops, steep, rocky slopes
(Fig. 2), gentle slopes (Fig. 3), and sandy and gravelly flats (Fig. 4).
These species are: Larrea tridentata, Ambrosia dumosa, Encelia far-
inosa, Hilaria rigida, Opuntia acanthocarpa, O. basilaris, O. bige-
lovil, Cercidium floridum, Fouquieria splendens, Krameria grayi,
Lycium andersonii, and Fagonia laevis. A few species are confined
mostly to broad, sandy washes (Fig. 5): Justicia californica, Bebbia
jJuncea, Chilopsis linearis, Acacia greggii, Olneya tesota, Psorotham-
nus spinosus, Lycium fremontii, and L. torreyi. Fewer species are
confined to the relatively mesic, steep, north-facing slopes within
the study area: Pleurocoronis pluriseta, Salazaria mexicana, and
Galium stellatum. Several are found only in broad, sandy washes
and north-facing slopes, suggesting that the two habitats are similar
in their moisture availability. Included in this last group are 4m-
brosia ilicifolia, Trixis californica, Hyptis emoryi, Condalia globosa,
and Zizyphus obtusifolia.

The common annuals also display a low degree of habitat differ-
entiation. Many species occur in washes, on sandy and gravelly flats,
on rocky slopes of all aspects, and on mountain tops. Common
species include Oligomeris linifolia, Lupinus arizonicus, Camissonia
brevipes, C. refracta, Chaenactis carphoclinia, Phacelia crenulata,
Mentzelia involucrata, Mohavea confertiflora, Cryptantha angusti-
folia, Plantago fastigiata, and Eriogonum thomasii.

Desert pavement and riverine dunes. Two habitats of special in-
terest are desert pavement and riverine dunes. Desert pavement
refers to unusually barren flats covered by a closely-packed layer of
pebbles, which usually have a well-developed coating of desert var-
nish (Musick 1975). Pavement soils are typically saline and sodic
and have very low infiltration rates. They are consequently the driest
habitats in the desert. Shrubs, mostly Larrea and Ambrosia dumosa
in the study area, occur only along narrow channels or runnels.
Annuals most commonly found on the pavement are Phacelia ne-
glecta, Chorizanthe rigida, Oligomeris linifolia, and Plantago fasti-
giata.

Riverine dunes are low dunefields that occur infrequently on the
floodplain adjacent to the Colorado River. Dunes have high rates
of infiltration and consequently are relatively mesic habitats (Bowers
1982). Riverine dunes in the study area are dominated by Atriplex
canescens subsp. linearis and Tessaria sericea. Some annuals that are
common on dunes elsewhere in the Sonoran Desert region are found
only on riverine dunes in the study area. These include Dicoria

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 363

%

Fics. 2-5. Desert habitats of eastern Imperial Co., California. 2. North-facing
slope in Cargo Muchacho Mountains, with sparse cover of Encelia farinosa and
Ambrosia dumosa. 3. Gentle slopes (bajadas) south of Picacho Peak. The sparse cover
in the foreground is mostly Larrea tridentata; Cercidium floridum and Olneya tesota
line the wash in the midground. 4. Sandy flat south of Cargo Muchacho Mountains
with Encelia farinosa, Larrea tridentata, and Ambrosia dumosa. 5. Ferguson Wash
at southeast end of Chocolate Mountains; Ferguson Lake, Colorado River, and Mar-
tinez Lake in the background. Dense vegetation along wash includes Cercidium florid-
um, Olneya tesota, and Condalia globosa.

canescens, Loeseliastrum schottii, Cryptantha costata, C. micrantha,
and Phacelia pediculoides.

Phytogeography. The study area is of interest to historical phy-
togeography because it is one of the few areas within the Southwest
that remained a desert throughout the Wisconsin glaciation (Cole
1986). During the last ice age the vegetation and flora more closely
resembled that occurring today in the Mohave Desert to the north.
Species recorded from packrat middens found near Picacho Peak
and dated to 13,000 yr B.P. include Coleogyne ramosissima, Chry-
sothamnus teretifolius, Salvia mohavensis, Eriogonum fasciculatum,
Brickellia atractyloides, Yucca brevifolia, and Yucca whipplei, none
of which occur in the modern flora. Species in the modern flora that
were present in the oldest middens include Larrea tridentata, Opun-
tia acanthocarpa, O. basilaris, and Ferocactus acanthodes. Many of
the modern dominants of the area, including Ambrosia dumosa,
Olneya tesota, Fouquieria splendens, Bebbia juncea, Ambrosia ili-

364 MADRONO [Vol. 34

cifolia, Petalonyx linearis, and Fagonia laevis do not appear in the
fossil record until the last few thousand years.

Wetlands

The vegetation along the Colorado River has changed consider-
ably during historical times. Doubtless there also have been undocu-
mented changes in the flora. Extensive gallery forests of Populus
fremontii and Salix gooddingii once lined both banks of the river
(Grinnell 1914, Minckley and Rinne 1985). These gallery forests
have been replaced by thickets of Tamarix chinensis, Tessaria ser-
icea, Salix exigua, Prosopis pubescens, Baccharis emoryi, and B.
glutinosa in the study area (Fig. 6). These thickets have been called
“Sonoran riparian scrublands” by Minckley and Brown (1982).

Other important wetland habitats occurring along the river include
sandbars, marshes, and alkaline depressions. Sandbars are inundated
by the spring floods and exposed when water levels recede in the
summer and fall. Older sandbars are dominated by Salix exigua,
Typha domingensis, Tamarix chinensis, and other perennials; youn-
ger sandbars (Fig. 7) provide habitats for several annual and peren-
nial species of Cyperus, Eleocharis, Juncus, and various Asteraceae.
The marshes (Fig. 8) were described by Minckley and Brown (1982)
and are dominated by several species of emergent perennials, in-
cluding Scirpus californicus, Typha domingensis, and Phragmites
australis, usually in dense, monospecific stands (Fig. 9).

The alkaline depressions form in fine-textured soils at the upper
limit of spring floods (Grinnell 1914); dominant species include
Distichlis spicata, Suaeda moquinii, and Eustoma exaltatum. Grin-
nell (1914) included Allenrolfea occidentalis as a dominant of such
sites, but we have yet to encounter this widespread plant in the study
area.

FLORA

The known vascular plant flora of the study area includes only
322 species (Table 2). Of these, 44 are introduced species, 12 are
“native weeds”’ found mostly on disturbed sites (e.g., Trianthema
portulacastrum, Amaranthus palmeri, Conyza canadensis, Mono-
lepis nuttalliana, Helianthus annuus), and 56 species are found only
in wetland habitats along the Colorado River. The flora of the un-
disturbed desert areas includes only 210 species. The low rainfall
and moderate topographic relief within the study area probably ac-
count for the rather depauperate flora. Because of the extreme aridity,
plant species do not “‘partition” the topographic gradient into several
distinct habitats. Flats, washes, slopes, and mountain tops are uni-
formly unfavorable for much of the growing season for most of the

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 365

Fics. 6-9. Wetland habitats of eastern Imperial Co., California. 6. Gallery forest
remnant in Impezial National Wildlife Refuge. Trees are Populus fremontii and Salix
gooddingii; dense thicket below trees along Colorado River includes Phragmites aus-
tralis, Tessaria sericea, and Tamarix chinensis. 7. Recently formed sandbar in Col-
orado River, densely colonized by Typha domingensis. 8. Marsh at Picacho State
Recreation Area with Typha domingensis. 9. Marsh with dense stand of Scirpus
californicus.

flora. In other words, extreme aridity reduces habitat diversity, with
a concomitant reduction in species diversity.

There are 15 genera within the study area with four or more
species. Camissonia (7 spp.), Cryptantha (7 spp.), Opuntia (6 spp.),
Phacelia (6 spp.), Atriplex (5 spp.), Mentzelia (5 spp.), Astragalus (4
spp.), and Eriogonum (4 spp.) are widespread and common through-
out the Intermountain Southwest. Chamaesyce (5 spp.), Lycium (4
spp.), and Aristida (4 spp.) are well developed in the southern desert
biogeographic provinces and Scirpus (5 spp.), Cyperus (4 spp.), El-
eocharis (4 spp.), and Juncus (4 spp.) are widespread in warm tem-
perate wetlands.

A tabulation of the flora by life-forms shows a high percentage of
annuals and shrubs in the study area (Table 3). The absolute number
of species of annuals and shrubs is probably no greater than would
be expected on the basis of the size and elevational range of the
study area (Bowers and McLaughlin 1982). This flora differs from
more mesophytic southwestern floras in its low number of herba-
ceous perennials; annuals and shrubs are only relatively more abun-

366 MADRONO [Vol. 34

TABLE 2. SUMMARY OF THE VASCULAR PLANT TAXA FOUND IN EASTERN IMPERIAL
Co., CALIFORNIA, INCLUDING THE TEN LARGEST FAMILIES (LISTED IN DECREASING
NUMBER, BY SPECIES).

No. of
No. of No. of No. of non-

Group families genera species natives
Vascular cryptogams l l l 0)
Gymnosperms 1 l 3 0
Dicotyledons 49 167 256 Pag |
Monocotyledons 11 38 62 17
Total all groups 62 207 522 44
Asteraceae 41 48 2
Poaceae 25 52 14
Fabaceae 14 22 2
Boraginaceae 6 17 0
Cyperaceae 4 14 1
Chenopodiaceae 7 12 5
Polygonaceae 4 12 4
Solanaceae 6 12 2
Cactaceae 6 11 0
Euphorbiaceae 5 11 0

dant here due to the poor development of an herbaceous perennial
flora. Only 50 species of perennial herbs occur in the undisturbed
desert habitats, accounting for only 24% of the desert flora. In the
more mesic wetlands of the Colorado River Valley, 35 herbaceous
perennial species account for 63% of the species found only in wet-
land habitats on the study site. In the desert areas perennial herbs
often are restricted to the relatively mesic microsites, 1.e., steeper
north-facing slopes and shaded banks of washes in deep sand.

ANNOTATED CATALOGUE

Botanical nomenclature follows Munz (1974), except where noted.
Each species is annotated with notes on distribution and frequency.
Frequency classes are: rare—known from only one or two localities
or collections; infrequent— known from several localities, but found
in only one or two habitats; occasional—often encountered, but
found mostly in particular habitats; common—often encountered,
occurring in several different habitats; and widespread — usually found
in several different habitats. Collection numbers, except where not-
ed, are for our collections deposited at ARIZ.

The catalogue is based primarily on our collections and field notes
from 1983-1986. In addition, a list of species that are likely to occur
in the study area was drawn up and the herbarium records at ARIZ,
ASU, RSA, and SD were examined for collections of these species.

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 567

TABLE 3. GROWTH FORMS OF SPECIES FOUND IN EASTERN IMPERIAL Co. STUDY
SITE AND THROUGHOUT THE SOUTHWESTERN UNITED STATES. ' Source: McLaughlin
(1986).

Percent of native species in:

Eastern Imperial Co.

a «Southwestern

Growth form Deserts Wetlands United States!
Trees 2.4 7.1 4.3
Shrubs 21.9 5.4 14.8
Herbaceous perennials 23.8 62.5 53.0
Annuals 46.7 25.0 24.6
Succulents ji. 0.0 3.3

Our emphasis was on the native, terrestrial species; aquatic and
wetland habitats along the Colorado River were collected from sev-
eral localities accessible by land and were more extensively examined
by boat during the fall of 1986. Agricultural areas and other disturbed
habitats were less intensively collected. Introduced species are pre-
ceded by an asterisk (*).

VASCULAR PLANTS OF EASTERN IMPERIAL COUNTY, CALIFORNIA
PTEROPHYTA
Pteridaceae
Notholaena parryi D. C. Eaton. Rare; n.-facing slopes, northwest end of Palo Verde
Mtns.; 2897.
CONIFEROPHYTA
Ephedraceae
Ephedra aspera Engelm. Widespread; slopes and washes; 2762, 2893.
Ephedra fasciculata A. Nels. var. clokeyi (Cutler) Clokey. Occasional; slopes, Cargo
Muchacho Mtns.; 2959.
Ephedra trifurca Torr. Rare; Larrea flats east of Midway Well; Wiggins 8552 (ARIZ).
ANTHOPHYTA — DICOTYLEDONEAE
Acanthaceae
Justicia californica (Benth.) D. Gibson [Beloperone c. Benth.]. Common; washes;
2802.
Aizoaceae

*Mesembryanthemum nodiflorum L. [Gasoul n. (L.) Rotm.]. Occasional; alkaline
depressions; 2632, 3242.
Trianthema portulacastrum L. Infrequent; fields and roadsides; 4138.

Amaranthaceae

Amaranthus palmeri S. Wats. Infrequent; fields and roadsides; 3054.
Tidestromia oblongifolia (S. Wats.) Standl. Common; flats and slopes; 26/8, 2718.

368 MADRONO [Vol. 34

Apiaceae

Daucus pusillus Michx. Rare; washes, west of Palo Verde Mtns.; 2880.
Hydrocotyle verticillata Thunb. Occasional; sandbars, banks of irrigation ditches;
3243, 349].

Asclepiadaceae

Asclepias albicans S$. Wats. Occasional; rocky slopes; 2934.

Asclepias erosa Torr. Rare; sandy wash near Midway Well; 3477.

Asclepias subulata Decne. Occasional; sandy flats; 3008.

Sarcostemma cynanchoides Decne. subsp. hartwegii (Vail) R. Holm. Occasional;
washes, climbing in perennials; 2650.

Sarcostemma hirtellum (Gray) R. Holm. Occasional; washes; 2891], 2925.

Asteraceae

Ambrosia dumosa (Gray) Payne. Widespread; flats and all slopes; 3479.

Ambrosia ilicifolia (Gray) Payne. Occasional; washes and on n.-facing slopes about
Picacho Peak; 2803.

Aster exilis Ell. Infrequent; sandbars; 3050.

Aster spinosus Benth. Infrequent; irrigation ditches and field borders; 3485.

Atrichoseris platyphylla Gray. Common; washes and gravelly slopes; 2647, 2811.

Baccharis emoryi Gray. Infrequent; sandbars, thickets, and marshes; 3055.

Baccharis glutinosa Pers. Common, sandbars and thickets; 3024, 3052.

Baileya pauciradiata Harv. & Gray. Infrequent; riverine dunes; 3224.

Baileya pleniradiata Harv. & Gray. Common; sandy flats and riverine dunes; 3026,
3134,

Bebbia juncea (Benth.) Greene. Common; washes; 3469.

Calycoseris wrightii Gray. Rare; rocky slopes and flats, Palo Verde Mtns.; 2902, 2926.

Chaenactis carphoclinia Gray. Common; slopes and washes; 2659, 2805.

Chaenactis stevioides Hook. & Arn. Common; slopes, washes, and riverine dunes;
2862, 2890, 3226.

Conyza canadensis (L.) Cronq. Infrequent; sandbars and disturbed sites; 3051.

Dicoria canescens Torr. & Gray. Rare; riverine dunes; Thornber, 22 Sep 1912 (ARIZ).

Dyssodia porophylloides Gray. Infrequent; slopes and canyons, Palo Verde Mtns.;
2914.

Eclipta alba (L.) Hassk. Rare; Colorado River, open water; Thornber 24 Sep 1912
(ARIZ), Pinkava et al. 10,358 (ASU).

Encelia farinosa Gray. Widespread; flats and mostly s.-facing slopes; 2759.

Encelia frutescens (Gray) Gray. Occasional; washes and on sandy flats near Ogilby
Hills; 2904.

Geraea canescens Torr. & Gray. Widespread; flats, slopes, and desert pavement; 3473.

Gnaphalium purpureum L. Infrequent; sandbars and thickets; 3230.

Helianthus annuus L. subsp. lenticularis (Dougl.) Ckll. Infrequent; fields and road-
sides; 2855, 3492.

Heterotheca sp. Occasional; sandbars; 3023, 4154, 4163, 4168.

Heterotheca subaxillaris (Lam.) Britt. & Rusby. Rare; sandbars north of Yuma; 3044.

Hymenoclea salsola Torr. & Gray. Common; washes; 2745.

Hymenoxys odorata DC. Infrequent; sandbars; Thornber 22 Sep 1912 (ARIZ), Ferris
22 Apr 1928 (RSA).

*Lactuca serriola L. Infrequent; disturbed ground; 3483.

Machaeranthera tephrodes (Gray) Greene. Infrequent; disturbed areas, roadsides;
3465.

Malacothrix glabrata Gray. Rare; rocky slopes, Palo Verde Mtns.; 2903.

Monoptilon bellioides (Gray) Hall. Widespread; washes, slopes, and flats; 2763, 2770.

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 369

Palafoxia arida B. L. Turner & M. I. Morris [P. /inearis (Cav.) Lag.]. Common; sandy
flats and riverine dunes; 264/, 2921, 3225.

Pectis papposa Harv. & Gray. Infrequent; sandy flats near Ogilby; 3060.

Perityle emoryi Torr. Widespread; washes, flats, and rocky slopes; 3470.

Peucephyllum schottii Gray. Common, mostly n.-facing, rocky slopes; 3487.

Pleurocoronis pluriseta (Gray) King & H. E. Robins. Common; n.-facing slopes; 2736.

Pluchea odorata (L.) Cass. [P. purpurascens (Sw.) DC.]. Common; sandbars and
marshes; 2724, 3025.

Porophyllum gracile Benth. Occasional; slopes and flats; 2717, 2932.

Prenanthella exigua (Gray) Rydb. [Stephanomeria e. Nutt.]. Rare; rocky slopes, Palo
Verde Mtns.; 2892.

Psathyrotes ramosissima (Torr.) Gray. Occasional; flats and slopes; 2636.

Rafinesquia neomexicana Gray. Common; washes and flats; rare on riverine dunes;
2863, 3223.

Senecio mohavensis Gray. Infrequent; washes and n.-facing slopes; 2773, 2904.

*Sonchus oleraceus L. Occasional; roadsides, fields, wasteground; 2730, 3229.

Stephanomeria pauciflora (Torr.) Nutt. Occasional; washes; 2642.

Stylocline micropoides Gray. Infrequent; gravelly flats; 2887, 2974.

Tessaria sericea (Nutt.) Shinners [Pluchea s. (Nutt.) Cov.]. Widespread; marshes,
thickets, sandbars, and riverine dunes; 30/1, 3498.

Trichoptilium incisum (Gray) Gray. Occasional; gravelly slopes and flats; 2630, 2912.

Trixis californica Kell. Common, n.-facing slopes; 3478.

Xanthium strumarium L. var. canadense (Mill.) Torr. & Gray. Rare; sandy banks;
4168B.

Bignoniaceae

Chilopsis linearis (Cav.) Sweet var. arcuata Fosb. Infrequent; washes; 2804.

Boraginaceae

Amsinckia intermedia Fisch. & Mey. Infrequent; washes, Palo Verde Mtns.; 2873.

Amsinckia tessellata Gray. Rare; n.-facing slope, Picacho Peak; 2798.

Cryptantha angustifolia (Torr.) Greene. Widespread; slopes, flats, washes, and riverine
dunes; 2616, 2740, 2781.

Cryptantha barbigera (Gray) Greene. Occasional; sandy flats and washes; 2741, 2752,
2937.

Cryptantha costata Brandegee. Infrequent; riverine dunes; 3220.

Cryptantha holoptera (Gray) Macbr. Occasional; n.-facing slopes; 2734, 2737, 2789,
2799.

Cryptantha maritima (Greene) Greene. Common; sandy washes; 2631, 2739.

Cryptantha micrantha (Torr.) Johnst. Infrequent; riverine dunes; 3/40.

Cryptantha pterocarya (Torr.) Greene. Infrequent; n.-facing slopes; 2788, 2962.

Heliotropium curassavicum L. var. oculatum (Heller) Jtn. Infrequent; alkaline depres-
sions; 2818, 2972.

Pectocarya heterocarpa (Johnst.) Johnst. Common; washes, flats, and slopes; 2776.

Pectocarya platycarpa (Munz & Johnst.) Munz & Johnst. Common; rocky flats, wash-
es; 2866.

Pectocarya recurvata Johnst. Rare; slopes; 2928.

Plagiobothrys jonesii Gray. Infrequent; sandy washes; 2782.

Tiquilia canescens (DC.) A. Richards. [Coldenia c. DC.]. Infrequent; volcanic slopes;
2760.

Tiquilia palmeri (Gray) A. Richards. [Coldenia p. Gray]. Occasional; sandy flats;
2640, 2711.

Tiquilia plicata (Torr.) A. Richards. [Coldenia p. (Torr.) Cov.]. Occasional; riverine
dunes and other sandy soils; 2850.

370 MADRONO [Vol. 34

Brassicaceae

*Brassica tournefortii Gouan. Common; sandy flats and washes, roadsides; 2742.

Caulanthus lasiophyllus (Hook. & Arn.) Payson [Thelypodium |. (Hook. & Arn.)
Greene]. Widespread; flats and washes; 2778.

Descurainia pinnata (Walt.) Britt. subsp. glabra (Woot. & Standl) Detl. Rare; wash
near Mitchell’s Camp; 2923.

Dithyrea californica Harv. Common; sandy flats and riverine dunes; 2919, 3141.

Draba cuneifolia Nutt. var. integrifolia S. Wats. Infrequent; washes and _ n.-facing
slopes, Palo Verde Mtns.; 2870.

Lepidium lasiocarpum Nutt. Common; washes; 2779, 2883, 3135.

Lesquerella palmeri S. Wats. Rare; 27 min. of Ogilby; Alexander and Kellogg 1924
(RSA).

*Sisymbrium irio L. Widespread; washes and disturbed areas; 2884.

Cactaceae

Cereus giganteus Engelm. [Carnegiea g. (Engelm.) Britt. & Rose]. Rare; several in-
dividuals vicinity of Senator Wash camp area, one plant near Ferguson Lake.
Grinnell (1914) noted 75 plants on the California side of the Colorado River;
there are many fewer today.

Coryphantha vivipara (Nutt.) Britt. & Rose var. a/versonii (Coulter) L. Benson. Rare;
mapped in study area by Benson (1982).

Echinocactus polycephalus Engelm. & Bigel. Occasional; rocky slopes, Chocolate Mtns.

Ferocactus acanthodes (Lem.) Britt. & Rose. Occasional; rocky slopes.

Mammiillaria tetrancistra Engelm. Widespread; rocky slopes.

Opuntia acanthocarpa Engelm. & Bigel. var. coloradensis L. Benson. Occasional;
gravelly flats and slopes near Colorado River.

Opuntia basilaris Engelm. & Bigel. Widespread; flats and slopes.

Opuntia bigelovii Engelm. Occasional; rocky slopes.

Opuntia echinocarpa Engelm. & Bigel. Common; slopes and flats, Palo Verde Mtns.

Opuntia ramosissima Engelm. Occasional; sandy and gravelly flats.

Opuntia wigginsii L. Benson. Infrequent; flats and slopes, Palo Verde Mtns.

Campanulaceae

Nemacladus glanduliferus Jepson. Common; washes and flats; 2775, 2813, 2910.
Nemacladus rubescens Greene var. tenuis McVaugh. Rare; 2 min. of Cargo Muchacho
Mtns.; Munz and Hitchcock 12145 (RSA).

Caryophyllaceae

Achyronychia cooperi Torr. & Gray. Infrequent; sandy flats and washes; 2939, 3246.
*Spergularia marina (L.) Griseb. Infrequent; alkaline depressions; 2973, 3232.

Ceratophyllaceae

Ceratophyllum demersum L. Infrequent; Colorado River, open water; Ricci 10, Minckley
and Dunfee YLD-32, YLD-54 (ASU).

Chenopodiaceae

Atriplex canescens (Pursh) Nutt. subsp. /inearis (S. Wats.) Hall & Clem. Occasional;
alkaline depressions and riverine dunes; 3027.

Atriplex elegans (Mogq.) D. Dietr. subsp. fasciculata (S. Wats.) Hall & Clem. Occa-
sional; desert pavement and roadsides; 2869, 3059.

Atriplex hymenelytra (Torr.) S. Wats. Occasional; rocky slopes, Chocolate Mtns.;
3481.

Atriplex lentiformis (Torr.) S. Wats. Common; alkaline depressions; 2777.

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 371

Atriplex polycarpa (Torr.) S. Wats. Common; washes; 2635B.

*Bassia hyssopifolia (Pall.) Kuntze. Infrequent; roadsides, sandbars; 3094.

*Chenopodium album L. Occasional; fields and roadsides; 3028.

*Chenopodium murale L. Common; fields and roadsides; 2615, 2845.

*Kochia scoparia (L.) Schrad. Occasional; fields and wasteground; 2949.

Monolepis nuttalliana (Schult.) Greene. Occasional; roadsides; 2954.

*Salsola australis R. Br. [S. iberica Sennen & Pau]. Common; roadsides, wasteground,
and riverine dunes; 346/.

Suaeda moquinii Greene [S. torreyana S. Wats. var. ramosissima (Standl.) Munz].
Common; washes and alkaline depressions; 3480.

Convolvulaceae

Cressa truxillensis HBK. Infrequent; alkaline depressions; McMurray 1396 (ARIZ).

Cucurbitaceae

Brandegea bigelovii (S. Wats.) Cogn. Occasional; climbing in trees and shrubs, washes;
3476.
Cucurbita palmata S. Wats. Rare; near Bard; Thornber 22 Sep 1912 (ARIZ).

Euphorbiaceae

Argythamnia lanceolata (Benth.) Muell.-Arg. [Ditaxis /. (Benth.) Pax & K. Hoffm.].
Common; washes and rocky slopes; 2746, 2755, 2758.

Argythamnia neomexicana Muell.-Arg. [Ditaxis n. (Muell.-Arg.) Heller]. Occasional;
rocky slopes; 2613, 2738, 3061.

Argythamnia serrata (Torr.) Muell.-Arg. [Ditaxis s. (Torr.) Heller]. Infrequent; sandy
flats; Thorne et al. 50920 (RSA), Balls and Everett 22900 (RSA).

Chamaesyce albomarginata (Torr. & Gray) Small. [Euphorbia a. Torr. & Gray]. Rare;
Colorado River Valley; Peirson 7200 (RSA).

Chamaesyce micromera (Boiss.) Woot. & Stand]. [Euphorbia m. Boiss.]. Infrequent;
sandy flats and washes; 27/4, 3061.

Chamaesyce pediculifera (Engelm.) Rose & Standl. [Euphorbia p. Engelm.]. Occa-
sional; rocky slopes, flats; 29/75.

Chamaesyce polycarpa (Benth.) Millsp. var. hirtella (Boiss.) Millsp. [Euphorbia p.
Benth. var. h. (Engelm.) Wheeler]. Widespread; washes and sandy and gravelly
flats; 2614.

Chamaesyce setiloba (Engelm.) Millsp. [Euphorbia s. Engelm.]. Common; sandy wash-
es and flats; 2719.

Croton californicus Muell.-Arg. var. mohavensis Ferg. Occasional; sandy flats; 2941.

Euphorbia eriantha Benth. Infrequent; washes; 2757, 2878.

Stillingia spinulosa Torr. Infrequent; sandy flats near Ogilby; Balls and Everett 22899
(RSA), Jones 8 May 1903 (RSA).

Fabaceae

Acacia greggii Gray. Common; washes; 2806.

Astragalus aridus Gray. Rare; found along the lower Colorado River Valley according
to Barneby (1964).

Astragalus insularis Kell. var. harwoodii Munz & McBurney. Infrequent; sandy flats;
2942, 2946.

Astragalus lentiginosus Dougl. var. borreganus Jones. Rare; near Ogilby; Balls and
Everett 22896 (RSA), Armstrong 1129 (SD).

Astragalus nuttallianus DC. var. imperfectus (Rydb.) Barneby. Rare; washes and sandy
flats; 2886, 2943.

Caesalpinia virgata Fisher [Hoffmannseggia microphylla Torr.]. Infrequent; rocky
slopes; 2909.

B72 MADRONO [Vol. 34

Calliandra eriophylla Benth. Occasional; washes and flats near Cargo Muchacho
Mtns.; 2756, 2931.

Cercidium floridum Benth. Common; washes; 3468.

Dalea mollis Benth. Widespread; sandy and gravelly flats; 2715, 2776.

Dalea mollissima (Rydb.) Munz. Occasional; gravelly flats; 2733B, 2749.

Lotus tomentellus Greene. Occasional; washes and sandy flats; 2653, 2944, 3137.

Lupinus arizonicus (S. Wats.) S. Wats. Common; washes, flats, and slopes; 2645,
2814.

Marina parryi (Torr. & Gray) Barneby [Dalea p. Torr. & Gray]. Infrequent; washes
and flats; 2898.

*Melilotus albus Desr. Infrequent; disturbed ground; 3460.

*Melilotus indicus (L.) All. Infrequent; roadsides, fields, irrigation ditches; 3247.

Olneya tesota Gray. Widespread; washes; 3472.

Prosopis glandulosa Torr. var. torreyana (L. Benson) M. C. Johnst. Occasional; broad
washes; 3489.

Prosopis pubescens Benth. Infrequent; thickets along Colorado River; 2658.

Psorothamnus emoryi (Gray) Barneby [Dalea e. Gray]. Infrequent; sandy soils along
Colorado River and Picacho Wash at All-American Canal; 2957, 3456. We have
yet to find Pilostyles thurberi Gray on plants in our study area.

Psorothamnus schottii (Torr.) Barneby [Dalea s. Torr.]. Infrequent; washes; 2655.

Psorothamnus spinosus (Gray) Barneby [Dalea s. Gray]. Widespread; washes; 3474.

Sesbania exaltata (Raf.) Cory. Infrequent; sandbars, fields and irrigation ditches;
3047.

Fouquieriaceae

Fouquieria splendens Engelm. Widespread; desert pavement, flats, and rocky slopes.

Gentianaceae

Eustoma exaltatum (L.) Griseb. Infrequent; sandbars and alkaline depressions; 3016.

Geraniaceae

*Erodium cicutarium (L.) L’Her. Occasional; roadsides; 2917.
Erodium texanum Gray. Infrequent; gravelly flats and slopes; 2888.

Haloragaceae

Myriophyllum exalbescens Fern. Occasional; open water; 3042.

Hydrophyllaceae

Eucrypta micrantha (Torr.) Heller. Occasional; densely vegetated washes and n.-facing
slopes; 2875, 2958.

Nama demissum Gray. Occasional; sandy washes; 2796.

Nama hispidum Gray var. spathulatum (Torr.) C. L. Hitchc. Occasional; sandy flats,
washes; 2624, 2947.

Phacelia ambigua Jones var. minutiflora (Voss) Atwood [P. minutiflora J. Voss].
Common; washes, flats, and rocky slopes; 2810, 2872.

Phacelia crenulata Torr. Widespread; washes, flats, rocky slopes; 2651, 2868.

Phacelia neglecta Jones. Common; desert pavement, broad washes; 2795, 2868.

Phacelia pachyphylla Gray. Infrequent; slopes and washes; 2664, 3245.

Phacelia pedicellata Gray. Infrequent; rocky slopes; 2801.

Phacelia pediculoides (J. T. Howell) Constance. Rare; riverine dunes; 3/38.

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 375

Krameriaceae

Krameria grayi Rose & Painter. Common; rocky slopes and flats; 2638.
Krameria parviflora Benth. var. imparata J. F. Macbr. Common; rocky slopes; Thorne
et al. 50910 (RSA), Ferris 7172 (RSA).

Lamiaceae

Hyptis emoryi Torr. Common; broad washes and n.-facing slopes; 2743.

Salazaria mexicana Torr. Rare; n.-facing slopes, vicinity Picacho Peak; 2800, 2965.

Teucrium cubense L. Rare; Palo Verde Valley; J. and L. Roos 4201 (RSA), Jepson
5258 (RSA).

Loasaceae

Mentzelia affinis Greene. Rare; vicinity Cargo Muchacho Mtns.; Peirson 9791 (RSA),
Munz and Hitchcock 12149 (RSA).

Mentzelia albicaulis (Hook.) Torr. & Gray. Infrequent; washes; 2874.

Mentzelia californica Thompson & Roberts. Rare; granitic flats east of Ogilby Hills;
2938.

Mentzelia involucrata S. Wats. Common; washes, flats, and rocky slopes; 2628.

Mentzelia longiloba J. Darl. Occasional; sandy flats, riverine dunes; 2955, 3136, 3227.

Petalonyx linearis Greene. Infrequent; rocky slopes; 2535, 2933.

Petalonyx thurberi Gray. Rare; 3 mi w. of Winterhaven; Raven 12909 (RSA), McMinn

1453 (RSA).
Lythraceae
Ammania coccinea Rottb. Rare; sandbars; Irwin 3 (ARIZ), Thornber 22 Sep 1912
(ARIZ).

Lythrum californicum Torr. & Gray. Infrequent; sandbars; 3019, 3045, 4165.

Malvaceae

Eremalche rotundifolia (Gray) Greene. Occasional; washes and gravelly flats; 2560,
2884.

Hibiscus denudatus Benth. Occasional; washes, flats, and slopes; 2716, 2754.

Horsfordia alata (S. Wats.) Gray. Rare; broad washes; 2750.

Horsfordia newberryi (S. Wats.) Gray. Rare; wash, Picacho State Rec. Area; 2732.

*Malva parviflora L. Common, roadsides and fields.

Sida leprosa (Ort.) K. Schumm. var. hederacea K. Schumm. Infrequent; weed in
cultivated areas; 4137.

Sphaeralcea ambigua Gray. Common; rocky slopes; 2720, 2960.

Sphaeralcea emoryi Torr. Occasional; sandy flats; 2968.

Martyniaceae

Proboscidea althaeifolia (Benth.) Decne. Rare; flats; Thorne et al. 50914 (RSA), Stark
1546 (RSA).

Nyctaginaceae

Abronia villosa S. Wats. Common; sandy flats, riverine dunes; 2945, 3142.

Allionia incarnata L. Common; rocky slopes; 3486.

Boerhaavia erecta L. var. intermedia (Jones) Kearney & Peebles. Infrequent; road-
sides; 3058, 4144.

Boerhaavia triquetra S. Wats. Infrequent; washes and flats; 2734.

374 MADRONO [Vol. 34

Boerhaavia wrightii Gray. Infrequent; washes, roadsides; 2725, 3056, 4143.
Mirabilis bigelovii Gray. Common; washes and n.-facing slopes; 2769, 2899.

Onagraceae

Camissonia arenaria (A. Nels.) Raven. Rare; rocky slopes; 2905.

Camissonia boothii (Dougl.) Raven subsp. condensata (Munz) Raven. Occasional;
sandy flats; 2864, 2969.

Camissonia brevipes (Gray) Raven. Common; washes, flats; 2639, 2885, 2924.

Camissonia cardiophylla (Torr.) Raven. Common; rocky slopes, washes; 2619, 2787.

Camissonia chamaenerioides (Gray) Raven. Infrequent; washes and slopes; 2877,
2964.

Camissonia clavaeformis Torr. & Frem. subsp. aurantiaca (S. Wats.) Raven. Com-
mon; gravelly flats; 2876, 2889, 2970.

Camissonia refracta (S. Wats.) Raven. Common; washes and gravelly flats; 2646,
2816.

Oenothera deltoides Torr. & Frem. Infrequent; northeast side of Palo Verde Mtns.;
Klein 144 (RSA).

Orobanchaceae

Orobanche cooperi (Gray) Heller. Infrequent; sandy and gravelly flats; Munz and
Hitchcock 12152 (RSA), Balls and Everett 22927 (RSA).

Papaveraceae

Eschscholzia minutiflora S. Wats. Common; washes and flats; 2768.
Eschscholzia parishii Greene. Infrequent; flats n. of Cargo Muchacho Mtns.; 2927.

Pedaliaceae

*Sesamum indicum L. Infrequent; cultivated in Colorado River Valley and occa-
sionally found along roadsides; 4/4/.

Plantaginaceae

Plantago fastigiata Morris [P. insularis Eastw. var. fastigiata (Morris) Jeps.]. Wide-
spread; slopes, flats, and washes; 2648.

Polemoniaceae

Gilia latifolia S. Wats. Infrequent; sandy washes; 2652.

Gilia stellata Heller. Widespread; washes, slopes, and flats; 2764, 2791, 2815, 2908.

Langloisia setosissima (Torr. & Gray) Greene. Occasional; washes, flats and slopes;
2649, 2817, 2911.

Linanthus jonesii (Gray) Greene. Occasional; washes and sandy flats; 2774, 2929.

Loeseliastrum schottii (Torr.) Timbrook [Langloisia schottii (Torr.) Greene]. Rare;
riverine dunes; 295], 3228.

Polygonaceae

Chorizanthe brevicornu Torr. Common; washes and flats; 2622, 2767.

Chorizanthe corrugata (Torr.) Torr. & Gray. Common; sandy and gravelly flats; 2629,
2871, 2936.

Chorizanthe rigida (Torr.) Torr. & Gray. Common; desert pavement, flats, and wash-
es: 2023.

Eriogonum deflexum Torr. Common; washes and gravelly flats; 262/.

Eriogonum inflatum Torr. & Frem. Common; gravelly flats and slopes; 3467.

Eriogonum thomasii Torr. Widespread; washes, slopes, flats; 2625, 2657, 2857, 2867.

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY 375

Eriogonum trichopes Torr. Infrequent; flats and gentle slopes; 2930.

*Polygonum argyrocoleon Steud. Infrequent; alkaline depressions; 3234.

*Polygonum aviculare L. Occasional; roadsides and wasteground; 2950.

Polygonum fusiforme Greene. Infrequent; marshes, canals, banks of Colorado River;
3236, 3248.

*Rumex conglomeratus Murr. Infrequent; disturbed ground; 3493.

*Rumex persicarioides L. Occasional; alkaline depressions and sandbars; 3015, 3235.

Portulacaceae

Calandrinia ambigua (S. Wats.) Howell. Infrequent; sandy flats; 2922.

Resedaceae

Oligomeris linifolia (Vahl) J. F. Macbr. Widespread; washes, desert pavement, sandy
and gravelly slopes; 3463.

Rhamnaceae

Colubrina californica 1. M. Johnst. Rare; Gavilan Wash e. of Indian Pass, Chocolate
Mtns.; 2766.

Condalia globosa 1. M. Johnst. var. pubescens I. M. Johnst. Common; broad washes
and n.-facing slopes; 2858.

Zizyphus obtusifolia (Hook.) Gray [Condaliopsis lycioides (Gray) Suesseng var. ca-
nescens (Gray) Suesseng]. Occasional; n.-facing slopes; 3488.

Rubiaceae

Galium stellatum Kell. subsp. eremicum (Hilend & Howell) Ehrendf. Infrequent; n.-
facing slopes, Palo Verde and Cargo Muchacho Mtns.; 2894, 2963.

Salicaceae

Populus fremontii S. Wats. Common; thickets and marshes; 3482.
Salix exigua Nutt. Occasional; sandbars; 3029, 3048.
Salix gooddingii Ball. Common; thickets and marshes; 2847.

Scrophulariaceae

*Bacopa monnieri (L.) Wettst. Occasional; sandbars; 3239.

Mohavea confertiflora (Benth.) Heller. Common; washes, slopes, and flats; 2656, 2771.

Penstemon pseudospectabilis Jones. Infrequent; n.-facing slopes, Chocolate and Palo
Verde Mtns.; 2784, 2913, 2969.

Simmondsiaceae

Simmondsia chinensis (Link) Schneider. Infrequent; sandy flats; 2748, 2901.

Solanaceae

Datura discolor Bernh. Occasional; washes; 27 /2.

Lycium andersonii Gray. Common; washes and n.-facing slopes; 2633, 2747, 2786,
2845,

Lycium fremontii Gray. Infrequent; washes; 2807.

Lycium parishii Gray. Infrequent; washes, Palo Verde Mtns.; 2879.

Lycium torreyi Gray. Infrequent; sandy flats and washes near Colorado River; 2643,
2952.

*Nicotiana glauca Grah. Infrequent; along drainage ditches; 3496.

Nicotiana trigonophylla Dunal. Common; washes and n.-facing slopes; 2654.

376 MADRONO [Vol. 34

Petunia parviflora Juss. Infrequent; sandbars; Swingle 260 (ARIZ), Monson 11 (ARIZ),
Parish 8322 (RSA).

Physalis acutifolia (Miers) Sandwith. Infrequent; weed of agricultural areas; 4/39.

Physalis angulata L. var. lanceolata (Nees) Waterfall. Infrequent; weed of agricultural
areas; 4/40.

Physalis crassifolia Benth. var. versicolor (Rydb.) Waterfall. Occasional; washes, flats,
and slopes; 2728, 2751, 2753.

*Solanum elaeagnifolium Cav. Infrequent; roadsides and fields; 3053.

Tamaricaceae

*Tamarix chinensis Lour. Common; marshes and thickets; 2634, 3030.
*Tamarix ramosissima Ledeb. Occasional; washes, ditches, and alkaline depressions;
2794.

Urticaceae

Parietaria floridana Nutt. Rare; rocky, n.-facing slopes, Picacho Peak and vicinity,
Chocolate Mtns.; 2797, 2966.

Viscaceae

Phoradendron californicum Nutt. Common; in Olneya, Prosopis, and Acacia; 3475.

Zygophyllaceae

Fagonia laevis Standl. Common; mostly on rocky slopes; 26/7.

Fagonia pachyacantha Rydb. Infrequent; slopes, Picacho State Rec. Area; 2627.

Larrea tridentata (DC.) Coville. Widespread; sandy and gravelly slopes, washes, and
steep rocky slopes; 2744.

*Tribulus terrestris L. Occasional; roadsides, disturbed areas; 3009.

ANTHOPHYTA— MONOCOTYLEDONEAE
Alismataceae

Echinodorus berteroi (Spreng.) Fassett. Rare; marshes; Monson 2 (ARIZ).

Araceae

*Pistia stratiotes L. Rare; drainage canal, Ft. Yuma; Peebles and Noble, 28 Oct 1941
(ARIZ).

Cyperaceae

*Cyperus alternifolius L. Infrequent; sandbars near Yuma; 30/4.

Cyperus erythrorhizos Muhl. Occasional; sandbars; 3012, 3022.

Cyperus laevigatus L. Infrequent; sandbars; 3241.

Cyperus odoratus L. Common; sandbars and marshes; 2852, 4134.

Eleocharis coloradoensis (Britt.) Gilly. Infrequent; sandbars; 4/50.

Eleocharis geniculata (L.) Roemer & Schultes. Common; sandbars, marshes and
thickets; 2851, 2860, 3497.

Eleocharis macrostachya Britton in Small. Rare; sandbars; Thornber 24 Sep 1912
(ARIZ).

Eleocharis parishii Britt. Infrequent; sandbars; 4/53.

Fimbrystylis vahlii (Lam.) Link. Infrequent; sandbars; Parish 8375 (RSA), Thornber
25 Sep 1912 (ARIZ).

Scirpus acutus Muhl. Rare; marshes; Striegler 20 (ARIZ), Behrends I (ASU).

Scirpus americanus Pers. Occasional; sandbars and marshes; 3240, 3244.

Scirpus californicus (C. A. Mey.) Steud. Common; marshes; 2859, 3020.

1987] McLAUGHLIN ET AL.: EASTERN IMPERIAL COUNTY el

Scirpus robustus Pursh. Infrequent; sandbars and marshes; Goodding 43-1 (ARIZ),
McMurray 1397 (ARIZ).

Scirpus validus Vahl. Infrequent; marshes; Booth A-112 (ARIZ), McMurray 1365
(ARIZ).

Juncaceae

Juncus acutus L. var. sphaerocarpus Engelm. Infrequent; sandbars; 3045, 3237, 3490.

Juncus articulatus L. Rare; sandbars; 4/61.

Juncus bufonius L. Rare; sandbars; Peebles and Harrison 5062 (ARIZ), Griner 10
Mar 1941 (ARIZ).

Juncus torreyi Cov. Occasional; sandbars; 3020, 4147.

Lemnaceae
Lemna gibba L. Rare; drainage ditches; 3233, 3501.

Liliaceae

Hesperocallis undulata Gray. Occasional; sandy and gravelly flats; 2920.

Najadaceae

Najas marina L. Infrequent; Colorado River backwaters; 4159.

Poaceae

Aristida adscensionis L. Widespread; slopes, flats, and washes; 2637.

Aristida californica Thurb. Infrequent; riverine dunes and sandy flats; 3462.

Aristida purpurea Nutt. Occasional; slopes; 2906.

Aristida wrightii Nash. Occasional; slopes; 2846, 2895.

*Avena fatua L. Common; fields and roadsides; 2856.

Bouteloua aristidoides (HBK.) Griseb. Common; sandy flats and washes; 2721].

Bouteloua barbata Lag. Infrequent; sandy and gravelly flats; 2722.

*Bromus catharticus Vahl [B. willdenowii Kunth]. Infrequent; roadsides and waste-
ground; 3238.

*Cynodon dactylon (L.) Pers. Common; fields and roadsides.

Diplachne uninervia (Presl) Parodi [Leptochloa u. (Presl) Hitchc. & Chase]. Occa-
sional; alkaline depressions and sandbars; 2723, 3231.

Distichlis spicata (L.) Greene var. stricta (Torr.) Beetle. Infrequent; alkaline depres-
sions; 297 /.

*Echinochloa colonum (L.) Link. Infrequent; sandbars near Yuma; 30/7.

Eragrostis pectinacea (Michx.) Nees. Infrequent; alkaline depressions; 2854.

Eriochloa aristata Vasey. Rare; near Bard; Reeder 29 Jun 1944 (ARIZ).

Erioneuron pulchellum (HBK.) Tateoko. Occasional; slopes and flats; 2865.

Heteropogon contortus (L.) Beauv. Rare; wash, w. side of Palo Verde Mtns.; Fuller
19014 (RSA).

Hilaria rigida (Thurb.) Benth. Common; washes and n.-facing slopes; 2713.

*Hordeum glaucum Steud. Occasional; roadsides; 2916.

*Hordeum vulgare L. Occasional; roadsides, sandy flats; 29/8.

*Leptochlioa filiformis (Lam.) Beauv. Infrequent; weed in agricultural areas; 4142.

Muhlenbergia microsperma (DC.) Kunth. Common; washes; 2731, 2882.

*Paspalum dilatatum Poir. Rare; disturbed ground; 3503.

Paspalum distichum L. Rare; irrigation ditch; Goodding and Reeder 8 Sep 1943 (ASU).

*Pennisetum setaceum (Forsk.) Chiov. Rare; roadside near Laguna Dam; 2849.

*Phalaris minor Retz. Occasional; fields and roadsides; 2948.

Phragmites australis (Cav.) Trin. Widespread; marshes and sandbars; 2953, 4158.

*Polypogon monspeliensis (L.) Desf. Occasional; irrigation and drainage ditches; 2853,
3494.

378 MADRONO [Vol. 34

*Schismus barbatus (L.) Thell. Common; flats and washes; 276 /.

*Sorghum bicolor Moench. Infrequent; roadsides; Barr 66-39 (ARIZ).
*Sorghum halepense (L.) Pers. Infrequent; fields and wasteground; 4/35.
Tridens muticus (Torr.) Nash. Rare; n.-facing slopes, Palo Verde Mtns.; 2896.
Vulpia octoflora Rydb. [Festuca o. Walt.]. Occasional; washes; 2784.

Potamogetonaceae

*Potamogeton crispus L. Common; shallow water in Colorado River; 4146.
Potamogeton foliosus Raf. Infrequent; Colorado River; 4/60.

Potamogeton nodosus Poir. Rare; Colorado River; Monson 3 (ARIZ).
Potamogeton pectinatus L. Common; Colorado River; 3502, 4145.

Typhaceae

Typha angustifolia L. Infrequent; marshes; Tuttle 14 Sep 1959 (ARIZ).
Typha domingensis Pers. Common; marshes; 3459.

Zannichelliaceae

Zannichellia palustris L. Occasional; drainage ditches and sandbars; 3500, 4148.

LITERATURE CITED

BARNEBY, R. C. 1964. Atlas of North American Astragalus. Mem. New York Bot.
Gard. 13:1-1188.

BENSON, L. 1982. The cacti of the United States and Canada. Stanford Univ. Press,
Stanford, CA.

Bowers, J. E. 1982. The plant ecology of inland dunes in western North America.
J. Arid Environ. 5:199-220.

1984. Plant geography of southwestern sand dunes. Desert Plants 6:3 1-42,

51-54.

and S. P. MCLAUGHLIN. 1982. Plant species diversity in Arizona. Madrono
29:227-233.

Cote, K. L. 1986. The lower Colorado River Valley: a Pleistocene desert. Quat.
Res. 25:392—400.

GRINNELL, J. 1914. An account of the mammals and birds of the lower Colorado
Valley. Univ. Calif. Publ. Zool. 12(4):51-294.

McLAUGHLIN, S. P. 1986. Floristic analysis of the southwestern United States. Great
Basin Nat. 46:46-65.

MINCKLEY, W. L. and D. E. BRown. 1982. Wetlands. Jn D. E. Brown, ed., Biotic
communities of the American Southwest— United States and Mexico. Desert
Plants 4:223-287.

and J. N. RINNE. 1985. Large woody debris in hot-desert streams: an his-
torical review. Desert Plants 7:142-153.

Munz, P. A. 1974. A flora of southern California. Univ. California Press, Berkeley.

Musick, M. B. 1975. Barrenness of desert pavement in Yuma County, Arizona. J.
Ariz. Acad. Sci. 10:24-28.

SHREVE, F. 1925. Ecological aspects of the deserts of California. Ecology 6:93-103.

. 1951. Vegetation of the Sonoran Desert. Carnegie Inst. Wash. Pub. 591,
Washington, DC.

TURNER, R. M. and D. E. BRown. 1982. Sonoran desertscrub. Jn D. E. Brown, ed.,
Biotic communities of the American Southwest— United States and Mexico.
Desert Plants 4:181-—221.

WILLMOTT, C. J., J. R. MATHER, and C. M. Rowe. 1981. Average monthly and
annual surface air temperature and precipitation data for the world. Part 2. The
Western Hemisphere. C. W. Thornthwaite Assoc. Publ. Climat. 34(2):1-378.

(Received 25 Jun 1985; revision accepted 16 Jun 1987.)

NOTES

THE RANGE AND Two New Locations OF Boschniakia strobilacea (OROBANCHA-
CEAE).— The known range of the root parasite Boschniakia strobilacea Gray (Ground
Cone) is from Vancouver Island, B.C., Canada [1885, Macoun s.n. (GH)] south to
the San Jacinto Mountains, Riverside Co., California [Reed 2535 (JEPS)]. A gap in
this range has existed between San Benito Peak, San Benito Co. [Jepson 2718 (JEPS)]
and Mt. Williamson, San Gabriel Mountains, Los Angeles Co. [9 Nov 1968, Thorne
et al. s.n. (RSA)], a distance of 330 km.

Two recent collections have narrowed this gap: one in the Scodie Mountains, Kern
Co. [3 Jul 1984, Shevock 10948 (RSA, CAS)] and another on Dry Lakes Ridge,
Ventura Co. [Magney 195-83, 124-84, 31-86, 32-86 (UCSB)]. The latter collections
were reported recently by Magney (A flora of Dry Lakes Ridge, Ventura Co., UCSB
Herb. Publ. No. 5, 1986). The Scodie Mountains locality is approximately 100 km
north of the Mt. Williamson site and 245 km southeast of the San Benito Peak
population. The Dry Lakes Ridge site is 130 km west of Mt. Williamson, 235 km
south of San Benito Peak, and 170 km southwest of the Scodie Mountains (Fig. 1,
page 380).

Boschniakia strobilacea is a fleshy-stemmed, parasitic herb 15—25 cm tall that arises
from a corm-like thickening at the junction with the root of the host plant (Gilkey,
Oregon St. Monogr., Studies in Botany No. 9, 1945). The leaves are scalelike, mostly
imbricated, and brownish. The flowers are dark reddish-brown and occur on a thick
spike 3.5-—6 cm thick. Flowering occurs from April through July. Boschniakia stro-
bilacea grows in a wide range of plant communities from near sea level to 2277 m
in the northern portion of its range, and from 1450 m at Dry Lakes Ridge (Magney,
op. cit.) to over 3015 m in southern California (Munz, A California fl., 1959; Abrams
and Ferris, Illustr. fl. Pacific States, 1960). Gray (Pacific Railroad Report iv, 118
(1857), 1876) described it from a specimen collected in the foothills of the Sierra
Nevada. Label data from the type specimen reads, “‘dry and rocky hills, South Yuba,
California” [23 May 1854, Bigelow s.n. (NY)].

Specimen label data obtained from many herbaria (A, CAS, CSUC, DAV, F, GH,
HSC, JEPS, K, LA, NY, RSA, SD, SFSU, UC, UCR, UCSB, US) indicate that B.
strobilacea has been collected most frequently in northern California. Approximately
300 collections are from about 150 locations throughout its range (Fig. 1). A list of
collection sites obtained from the herbaria mentioned above is available from the
author upon request.

Arbutus menziesii and Arctostaphylos glauca, A. nevadensis, A. parryana, A. patula,
A. pungens, and A. tomentosa have been suggested as host plants for B. strobilacea.
The Dry Lakes Ridge population adds another host, Arctostaphylos glandulosa.

I appreciate the comments of reviewers L. Heckard, C. Mason, B. Tanowitz, an
anonymous reviewer, and the editor.—DAvip L. MAGNEyY, Dames & Moore, 175
Cremona Dr., Suite A-E, Goleta, CA 93117. (Received 17 Dec 1986; revision ac-
cepted 15 Jul 1987.)

MADRONO, Vol. 34, No. 4, pp. 379-380, 1987

Scodie Mtns.
&

Dry Lakes Ridge

“og,°°

oO 80 km
oO 50 miles

Fic. 1. Distribution of Boschniakia strobilacea Gray in California and Oregon.
It also has been collected from Vancouver Island, B.C., Canada (not shown). @ =
one or more collections of B. strobilacea and roughly represents one population.

NOTEWORTHY COLLECTIONS

CALIFORNIA

SCRIBNERIA BOLANDERI (Thurb.) Hack. (POACEAE). —San Diego Co., Del Mar Mesa,
Caltrans vernal pool preserve, at the edges of vernal pools with Agrostis microphylla,
Festuca myuros, Juncus bufonius, etc. in an area of Adenostoma chaparral, T14S R3W
S23, 128 m, 28 Apr 1987, Moran, Rilling, and Zedler s.n. (SD).

Significance. Fills a gap between California collections from Santa Barbara Co., ca.
340 km nw. (Smith, A Flora of the Santa Barbara Region, CA, 1976), and Kern Co.,
ca. 320 km n. (Twissleman, A Flora of Kern County California, 1967); and the single
Mexican collection from Laguna Hanson, Baja California Norte, ca. 160 km se. (R.
F. Thorne, RSA, pers. comm.).

AGROSTIS AVENACEA Gmel. (POACEAE).—San Diego Co., Kearny Mesa, Miramar
Mounds National Landmark, Miramar Naval Air Station, ca. 100 m w. of Hwy. 163
and 50 mn. of the as yet uncompleted extension of Hwy. 52, 32°50'30’N, 117°8’00’W,
130 m. In an artificially impounded seasonal wetland with Polypogon monspeliensis,
Eleocharis acicularis, E. cf. macrostachya, Aponogeton distachyus, and a variety of
native vernal pool species, 7 Jun 1987, Zedler, Moran, and Rilling s.n. (SD).

Significance. First report of this introduced species from southern California. Like
Aponogeton (Keeley and Keeley, Madrono 26:188, 1979), this species may have
invaded because of the unusually long water duration imposed on a former vernal
pool area by the highway blocking the drainage. Known previously from n. CA
(Crampton, Grasses in CA, 1974) and scattered locations in central U.S.; native to
Australia, New Zealand, and the South Pacific. — PAUL H. ZEDLER, VIRGINIA MORAN,
TRUDY RILLING, Biology Dept., San Diego State Univ., San Diego, CA 92182-0057;
and GEOFFREY A. LEVIN, Botany Dept., San Diego Natural History Museum, P.O.
Box 1390, San Diego, CA 92112-1390.

PETERIA THOMPSONAE S. Wats. (FABACEAE).— Inyo Co., Kingston Range, California
Valley, Mesquite Valley Rd. 8 mi ne. of Smith Talc Mine Rd., sandy bajada, 2700
ft, 4 May 1980, de Nevers 150 (RSA).

Significance. First report of genus from California. Previously known from adjacent
Nye and Clark cos. in NV.—AARON LISTON, Rancho Santa Ana Botanic Garden,
Claremont 91711.

ECUADOR

BUDDLEJA AMERICANA L. (BUDDLEJACEAE). — Ecuador, Galapagos Islands, Floreana,
Cerro de Naranjas, 200 m, 13 Feb 1986, J. E. Lawesson and H. Zederkof 2849 (CDS,
DLF, QCA); e. of Cerro Pajas, near Wittmer’s farm, 13 Feb 1986, Y. Carvajal 162
(CDS). Several hundred plants of several ages observed, associated with common
guava in the first location and with a Scalesia pedunculata forest in the latter.

Previous knowledge. Floreana, near Wittmer’s farm, 330 m, 14 Mar 1970, S. Jtow
3 1400-1 (DS), but unreported previously. Known from Mexico to Bolivia, Cuba, and
Jamaica (Norman, Buddlejaceae, Fl. Ecuador, 1982).

Significance. New family for the Galapagos Islands. Older settlers report (F. Cruz,
pers. comm.) it was present 50 years ago. — ELIANE M. NORMAN, Dept. Biology, Stetson
Univ., DeLand, Fl 32720; and JoNAs E. LAwesson, Estacion Cientifica Charles Dar-
win, Isla Santa Cruz, Galapagos, Ecuador. Field work was supported by Danish

MADRONO, Vol. 34, No. 4, pp. 381-382, 1987

382 MADRONO [Vol. 34

Natural Science Council, grants 11-5471 and 11-5663 to JEL. We are grateful to the
Galapagos National Park Service and Charles Darwin Research Station for their
assistance.

NEVADA

ASTRAGALUS GILMANII Tidest. (FABACEAE). — Lincoln Co., Groom Mountain Range,
ca. 110 km w. of Caliente, occasional on tuff, se. side of basalt cone just n. of Cattle
Spring in scattered Pinyon-Juniper, T6S R55'2E $18, 1830 m, 7 May 1985, Marrs-
Smith and Nachlinger 91 (NY, RENO, UNLV) (determined by R. C. Barneby, NY).

Significance. First record for NV and an e. range extension of ca. 215 km from the
Panamint Mtns., Inyo Co., CA.

ERIGERON OVINUS Cronquist (ASTERACEAE). — Lincoln Co., Groom Mountain Range,
ca. 110 km w. of Caliente, limestone ridge with Cercocarpus ledifolius and Forsellesia
nevadensis, T7S R56E S6, 2260 m, 4 Jun 1985, Marrs-Smith and Nachlinger 47 (NY,
RENO, UNLV).

Significance. A w. range extension of 29 km. Known only from Clark and Lincoln
cos., NV.

POLYGALA SUBSPINOSA S. Wats. var. HETERORHYNCHA Barneby (POLYGALACEAE). —
Lincoln Co., Groom Mountain Range, ca. 110 km w. of Caliente, on volcanic tuff
with scattered Artemisia tridentata, T6S RS55E S13, 1890 m, 7 May 1985, Marrs-
Smith s.n. (NY) (determined by R. C. Barneby, NY).

Significance. First record for Lincoln Co. and a n. extension of ca. 60 km. Previously
known from Clark and Nye cos., NV.—GAYLE MARRS-SMITH, Dept. Biological Sci-
ences, Univ. Nevada, Las Vegas, 89154; and JAN NACHLINGER, Biological Sciences
Center, Desert Research Inst., Reno, NV 89506.

REVIEW

Poisonous Plants of California. By THOMAS C. FULLER and ELIZABETH MCCLINTOCK.
433 pp. + 16 color plates. University of California Press, Berkeley, CA.

This volume is one of the California Natural History Guides (#53) published by
the U.C. Press. This interesting and readable book provides a broad overview of
plant toxicity at an introductory level. The main body consists of brief descriptions
of hundreds of plants and fungi poisonous to humans and animals. It includes brief
descriptions of symptoms of poisonings, brief chemical identification of the toxins,
and interesting anecdotes of poisoning case histories. Included also are several useful
species lists such as the most seriously poisonous plants and fungi, plants most often
toxic to livestock, plants causing dermatitis, plants causing hay fever and plants
accumulating nitrates. Although most of the book deals with flowering plants, there
are also brief chapters on toxic algae, fungi, ferns and horsetails, and gymnosperms.
The flowering plants are treated alphabetically by family. The book is well organized
and cross referenced so as to facilitate finding specific information about plants or
toxins. There are separate indices of common and scientific names, as well as a general
subject index. There are many (but not enough) good line illustrations, and over 60
small but very good color photos that stress diagnostic characteristics.

MADRONO, Vol. 34, No. 4, pp. 382-383, 1987

1987] REVIEW 383

There is a separate chapter on the chemistry of plant toxins and derivative drugs
which is written in non-technical terms understandable to the layperson. The basic
chemistry of allelopathy and photosensitization are described briefly. The reader with
more knowledge of plant chemistry, especially of secondary metabolites, will be
tantalized continually and will want to dig back into the literature for more detailed
information.

This book contains no identification keys, but provides an excellent starting place
to learn about both California native and introduced toxic plants. Although the
descriptions of the plants are brief and non-technical, they are accurate and stress
the diagnostic characteristics. There is no attempt to duplicate the exhaustive, bo-
tanically complete plant descriptions and keys as in Munz’s California Flora and
Bailey’s Manual of Cultivated Plants. Likewise, because the descriptions of the chem-
istry and symptoms of toxicity are very brief, and considering the omission of an-
tidotes and medical treatments, it can be concluded that medical advisement is
prudently beyond the scope of the book. The book will be able to provide a quick,
preliminary identification of plant material, toxins, and symptoms.

Poisonous Plants of California is welcome due to the on-going interest in diet,
health, herbs, and edible and medicinal plants. I will put this book on my shelf next
to the mushroom identification books and alongside Lewis’ Medical Botany and
Lampke’s Plant Toxicity and Dermatitis. As a botany teacher, I am often asked by
students about the toxicity (edibility, caffeine content, medicinal use, etc.) of some
particular plant (mushroom, weed, herb, ornamental, etc.). It will be the first book I
pull off my shelf to answer those questions. This book will be useful to anyone with
an interest in plant edibility, toxicity, or medicinal qualities. It will be especially useful
to naturalists, field biologists, ranchers, emergency room physicians, and veterinar-
ians.— ROBERT CUMMINGS, Dept. Biological Sciences, Santa Barbara City College,
Santa Barbara, CA 93109.

ANNOUNCEMENT

CORRECTIONS TO CBS SPEAKER SCHEDULE* FOR 1987-1988

Date Speaker & Topic

Jan 21 Donald Koehler, Santa Barbara
‘Spectral quality of yellow flowers in relation to pol-
lination”’

Feb 20 Stephen J. Gould
Annual Banquet, Topic to be announced

May 19 Meeting room at UC Botanic Garden, not LSB 2503
* See Madrono 34(3):272.

384 MADRONO [Vol. 34

LETTERS

Dear Editor:

I'd like to give a rather general response to the request for reviewer’s opinions on
the amount of interest each paper might stimulate among the members of the Cali-
fornia Botanical Society. I occasionally review manuscripts for Madrono and I am
interested in new species descriptions. In general, such papers aren’t too exciting.
Nevertheless, they constitute a significant aspect of systematics and I think Madrono
should continue to provide a place for such papers, especially when they deal with
California taxa. Perhaps more important for CBS’s general membership, however, is
the role that new species descriptions have for environmentalists. For example, in
some papers, the new species 1s judged by authorities to have a highly restricted range.
This information may be quite valuable in future efforts to protect land where the
new species grows. Furthermore, briefaccounts on how new species were “‘discovered”’
also can be exciting illustrations of how interested amateurs can be involved in
furthering scientific knowledge of our native flora. Thus, it might be good to actually
encourage authors of new species descriptions to include people-oriented and envi-
ronmental information!

Mark A. Schlessman
Dept. of Biology

Vassar College
Poughkeepsie, NY 12601

COMMENTARY
MESSAGE FROM THE PAST CBS PRESIDENT

It is with great pleasure that I address the membership of the California Botanical
Society in this message column that was initiated at the suggestion of Wayne Ferren
in our October 1986 issue of Madrono.

For decades, our journal has served as the vital organ for disseminating the results
of scientific research to our membership. Historical convention and space limitations
have all too often precluded commentaries that encourage dialogue or summarize
the Society’s activities, accomplishments, and aspirations for the future. Due in large
part to Ferren’s efforts, this is changing in a positive way. In conveying information
on the inner workings of our Society, it is my hope that this column will help to serve
as a catalyst for more communication and constructive input from our members.

The By-Laws of CBS give us a broad mandate—“‘. . . to stimulate interest and to
further advancement in the entire field of botany especially in the western United
States’. This charge has rested largely on the shoulders of a cadre of elected and
appointed volunteers, the Executive Council, who meet at Berkeley on a monthly
basis during the academic year to formulate policy, tend to routine business, and
plan for the future. The Council addressed a heavy agenda during the past year. The
burdensome task of handling the Society’s financial transactions and modest invest-
ments required careful consideration. The heavy demands on the CBS Treasurer led
to the creation of the new position of Financial Officer. Our incumbent Treasurer,
Dr. Cherie L. Wetzel, kindly agreed to continue her long service to the Society by
assuming the duties of this new position, and Dr. Thomas F. Daniel accepted an
appointment to the Council as our new Treasurer. These changes also forced us to
take a closer look at the Society’s financial health. The Society’s major financial
commitment each year is the publication of Madronio. Printing costs, however, have
escalated an estimated 30% during the last six years. During this time, we have held
to a steadfast policy of no dues increases because the Council strongly believed in
providing its members with a quality journal at a bargain basement price. We will

1987] COMMENTARY 385

continue to produce a journal of the quality that we currently enjoy, but we can do
so only by implementing modest dues increases that will go into effect with volume
35 of Madrono. For the short term, this will put us in a sound financial position that
will allow us to continue to provide partial support for worthy scientific endeavors.
We continue to support the annual CBS Graduate Student Meetings and provide
small cash awards for the best papers in several categories. Year before last we made
a much-needed contribution toward the start-up phase of the Jepson Herbarium
revision of the ‘““Manual of the Flora of California’. This past year we provided a
modest no-interest loan that together with financial assistance from the California
Native Plant Society will facilitate the publication of a new edition of “‘Terrestrial
Vegetation of California’. During the past year, the Council directed its attention to
some projects that were deemed long overdue. Council member Barbara Ertter spear-
headed the drive to assemble a computerized questionnaire that will be used to
produce the Society’s first membership directory. We have also explored the possi-
bilities of publishing a comprehensive cumulative index to all published volumes of
Madrojno. The latter project could prove to be more costly than initially anticipated,
but some decision on the most cost-effective method of producing an index should
be reached during the coming year. To comfortably support the above kinds of projects
on an ongoing basis, it will be imperative that the Society enhance its income base.
This could be accomplished by a vigorous membership drive and perhaps the solic-
itation of private and corporate support. This must all be done in a way that will not
jeopardize our non-profit, tax exempt status.

The most onerous task that confronted the Council during the past year was the
selection of a new Editor to succeed Wayne Ferren who will complete his term early
in 1988. Only the job of Editor itself exceeds the task of identifying a talented and
qualified individual willing to give freely of his or her time in the service of science.
I am pleased to announce that Dr. David J. Keil accepted our invitation to become
the new Editor of Madrono. The Council joins me in welcoming him aboard. We
regret that Wayne’s term has passed so swiftly. He assumed his editorial duties with
alacrity and good humor during a difficult transition period. He has maintained a
standard of excellence and introduced new and effective format changes that we will
want to continue. Our thanks to you, Wayne, for a job well done! We look forward
to your continued advice as a member of the Board of Editors.

In reviewing Society activities, I note with concern a total lack of graduate student
response to the financial support offered by the Society in the form of small research
grants to assist with field work, travel expenses, and supplies. I encourage all interested
students and/or their advisors to contact the CBS President for more information on
this potential source of research support.

In closing I would like to thank our Executive Council, our Editor, Associate Editor,
and their Board, and the many reviewers who have graciously given of their time
and expertise during the past year. I would be remiss if I neglected to extend our
appreciation to the diverse group of scientists who continue to share their observations
and research findings with us through the printed pages of Madrofio. —FRANK AL-
MEDA, Dept. Botany, California Academy of Sciences, Golden Gate Park, San Fran-
cisco, CA 94118-9961.

EDITOR’S REPORT FOR VOLUME 34

This annual report provides an opportunity for the Editor to communicate the
status of manuscripts received for publication in Madrofo and to comment on other
aspects of the journal. Between 1 Jul 1986 and 30 Jun 1987, 80 manuscripts were
received (37 articles, five notes, 38 individual noteworthy collections). Since 30 Jun
1987, 18 manuscripts (8, 1, 9) have been received. The current status of the 93

386 MADRONO [Vol. 34

unpublished manuscripts is 26 in review (13, 2, 11), 28 in revision (25, 3, 0), eight
needing a decision by the editors (6, 2, 0), and 31 accepted for publication (9, 1, 21).
There are two unpublished book reviews. Volume 34 included 81 published manu-
scripts (32, 2, 35), nine book reviews, and three commentaries or letters. The period
between submittal and publication averaged 1.5 years for articles. We rejected five
manuscripts in 1987, and one was withdrawn by the author.

With publication of this issue, I complete my term as Editor of Madrono. I am
honored to have served the California Botanical Society in this capacity and I hope
my editorship has contributed to the growth of this important regional journal. The
success of the past year is shared with many individuals who have helped guide
Madrofo and its editor. Marion Cave provided photographs for the dedication; Barry
D. Tanowitz, Associate Editor, provided much editorial and technicai advice and
often sparked healthy debate on many topics; Steven Timbrook, member of the Board
of Editors, contributed the Table of Contents and the Index to volume 34, a job
appreciated by all members and subscribers. Thanks Barry and Steve! I also give
special thanks to Annetta Carter, who has been a mentor during my editorship.

I thank the Executive Council of CBS for its guidance and the Board of Editors
and the numerous reviewers (62) for their assistance with volume 34. Many reviewers,
e.g., Jim Henrickson, Mary Carroll, Bill Critchfield, Susan Conard, and John Strother,
commented on several of the manuscripts published this year and I am grateful for
their dedication to Madrofo. The organization, readability, and style of all papers
reflect ideas provided by our reviewers and the high quality of papers in Madrono is
attributed in part to their sincere efforts. In addition, the excellent style, clarity, and
general appearance of our journal is always enhanced by the exceptional assistance
provided by my friends at Allen Press.

During the past 2.5 years as Editor, I have implemented several policies that I
hope will continue to stimulate interaction among members of the Society and the
readership of our journal. Editorials, commentaries, letters to the editor, and an-
nouncements have extended our journal beyond its role as a vehicle for reporting
science to one also of comment and reflection. With the publication of 35(1), Madronio
also will be a bilingual journal. Members of the CBS have a long-term and increasing
interest in the flora and vegetation of Mexico and Central and South America. Because
of this interest, I believe our new policy to publish Spanish language papers with
English abstracts and, where appropriate, English language papers with Spanish ab-
stracts will open Madrofio to an even wider readership, perhaps increase the mem-
bership in CBS, and extend a gesture of goodwill to our Hispanic colleagues and
neighbors. There has been widespread support for this decision, and I request that
a!l members encourage their Hispanic colleages to join the CBS and contribute to
Madrojno. I thank Roberto Iglesias Prieto (UCSB) and various reviewers for assistance
with these manuscripts, and David J. Keil (the new Editor) for his encouragement.

Most important to Madrovio, however, are the contributions by authors. The strength
of our journal depends on the quality and quantity of manuscripts, and a review of
papers published in volume 34 reveals the diversity of botanical investigations re-
ported by members of the Society. For example, we have published contributions in
historical botany, ecology, systematics, morphology, cytotaxonomy, floristics, phy-
togeography, physiology, fire ecology, reproductive biology, dispersal and germina-
tion, and mycorrhizae. Habitats and plant communities discussed by contributors
included, for example, wetlands, oak woodlands, chaparral, grasslands, the alpine,
many desert types, and coniferous, mixed evergreen, and tropicel deciduous forests.
Geographic areas covered in papers included many regions of western North America
(in Canada, U.S., Mexico) and portions of South America (e.g., Colombia and Ec-
uador). Plant groups investigated by our contributors are many, including myxo-
mycetes, zygomycetes, bryophytes, hepatics, gymnosperms, and angiosperms. In ad-
dition, nine new taxa were described and four new combinations were made in
volume 34.

My goals as Editor have been to guide authors through the review, revision, and

1987] EDITOR’S REPORT FOR VOLUME 34 387

publication processes and to implement new journal policies that could enhance the
editorial and managerial aspects of Madrono. To these ends, I can report a healthy
journal with larger, more diverse issues than ever. Although this editor may not have
been as prompt in responding to author’s needs as some would have liked, and
although a few authors were ‘“‘extremely concerned” at times and found the process
to be “‘very upsetting” or “‘increasingly objectionable’’, neither the reviewers nor the
editors ever intended to be ““venomous’’, “‘inattentive’’, ““incoherent’’, ““antagonistic’’,
or ““unreasonable’’. I recognize the “‘fallibility of editors and reviewers’’, as well as
that of authors; but, hopefully, we strive for “careful and thorough editing” to correct
errors of fact, lack of clarity, poor grammar, and inconsistent style. Our reviewers,
in particular, have volunteered a considerable effort to provide generally “excellent
editorial and technical comments’, “‘valuable additions’’, and “‘helpful critiques”’,
and they are an essential part of a peer-reviewed journal such as Madrono. The review
and revision process is never meant to be “ludicrous’’, “counter productive’’, nor an
“increasingly frustrating experience’’, but should be “‘a valuable learning experience”’
that often “improves the manuscript considerably’. I trust we are all students of
the plant sciences because of our love for the field and, within this, our desire for
communication among our peers. The reviewers and editors, however, would be most
appreciative of fewer hard knocks, and thus, friends, some of us could lighten-up a
bit! A result of thorough preparation, patience, and good humor should be even better
papers and, certainly, lower blood pressure. Best wishes to the new Editor! W. R. F.

Jr. 30 Nov 1987.

REVIEWERS OF MANUSCRIPTS

The editors thank all reviewers for their assistance with manuscripts, and extend
special thanks to those who reviewed several manuscripts published in 1987. We are
grateful for your generous contributions of time and effort toward maintaining and
improving the quality of papers published in Madrono. Reviewers for volume 34
are:

Spencer C. H. Barrett
Jim A. Bartell

Jerry M. Baskin

John H. Beaman
Meredith Blackworth
Robert Boyd

Jack H. Burk

Ragan Callaway
Judith M. Cann-Hilliker
Mary Carroll

Annetta Carter
Kenton L. Chambers
Susan G. Conard
Lincoln Constance
William B. Critchfield
Christopher Davidson
Frank W. Davis
Dorothy Douglas
Barbara Ertter

Marie Farr

William J. Ferlatte

Arthur Gibson

James R. Griffin

J. Robert Haller
Lawrence R. Heckard
James Henrickson
Richard Jensen
Walter Knight
Arthur R. Krucheberg
Jochen Kummerow
Harlan Lewis

Jack Major

Charles T. Mason
Robert J. Meinke
Louis V. Mingrove
Richard A. Minnich
Reid Moran

James D. Morefield
Maynard M. Moseley
Walter H. Muller
Brad Musick

Nalini M. Nadkarni

Arthur M. Phillips III
Donald J. Pinkava
David C. Randall
Philip W. Rundel
John O. Sawyer Jr.
Robert A. Schlising
James Shevock
Bradley G. Smith
James P. Smith Jr.
Ted V. St. John

John L. Strother
Barry D. Tanowitz
Barbara M. Thiers
Harry D. Thiers

John H. Thomas
Robert F. Thorne

B. L. Turner

Thomas R. Van Devender
Frank Vasek

William A. Weber

INDEX TO VOLUME 34

Classified entries: major subjects, key words, and results; botanical names (new
names are in boldface); geographical areas; reviews; letters; commentaries. Incidental
references to taxa (including most lists and tables) are not indexed separately. Species
appearing in Noteworthy Collections are indexed under name, family, and state or
country. Authors and titles are listed alphabetically in the Table of Contents to the

volume.

Acacia smallii, in successional commu-
nities, 250.

Agrostis avenacea, noteworthy collection
in CA, 381.

Alliaceae: Allium shevockii, new species
from CA, 150.

Allium shevockii, new species from CA,
150.

Alloispermum insuetum, new species from
Colombia, 162

Alpine annual plant species in the White
Mountains, CA, 315.

Amaryllidaceae: (see Alliaceae).

Amaurochaete comata, new record for
CA, 51.

Amblystegiaceae: Campylium halleri, new
record for Mexico, 69.

Anacardiaceae: Rhus trilobata var. sim-
plicifolia, new record for CA, 171.

Arcyria magna, new record for CA, 50.

Arecaceae: cold tolerance in Washing-
tonia filifera, 57.

Arizona: Astragalus hypoxylus, notewor-
thy collection, 170; Stenocereus thur-
beri seedcrop characteristics and min-
imum reproductive size in southern AZ,
294.

Asteraceae: Cymophora returned to 7ri-
dax, 354; Erechtites glomerata, my-
corrhizae associated with invasion on
San Miquel Island, CA, 260; Erigeron
ovinus, range extension in NV, 382;
Lessingia tenuis, range extension,
chromosome count, and mephitism,
168; Lygodesmia grandiflora, new
record from NM, 171; Soliva in CA,
228.

New combination: Tridax hintonii,
S57.

New species: Alloispermum insuetum,
from Colombia, 162; Axiniphyllum
durangense, from Mexico, 165.

Astragalus: Astragalus gilmanii, new rec-
ord for Nevada, 382; A. hypoxylus,
noteworthy collection from Arizona,
170;

Athalamia hyalina, new record for Mex-
ico, 69.

Axiniphyllum durangense, new species
from Mexico, 165.

Aytoniaceae: Mannia fragrans, new rec-
ord for CA, 69.

Boschniakia strobilacea, range and new
locations, 379.

Bryaceae: Bryum blindii, new record for
CO; 174.

Bryum blindii, new record for CO, 171.

Buddleja americana, new family for Ga-
lapagos Islands, 381.

Buddlejaceae: Buddleja americana, new
family for Galapagos Islands, 381.

Cactaceae: Stenocereus thurberi seedcrop
characteristics and minimum repro-
ductive size in southern AZ, 294.

California: Agrostis avenacea, notewor-
thy collection, 381; alpine annual plant
species in the White Mountains, 315;
Boschniakia strobilacea, range and new
locations, 379; Ceanothus cuneatus and
C. leucodermis seed dispersal in oak-
woodland savanna, 283; Erechtites
glomerata, mycorrhizae associated with
invasion on San Miguel Island, 260;
Lessingia tenuis, range extension,
chromosome count, and mephitism,
168; Pinus contorta var. murrayana,
germination, establishment, and in-
vasion in mountain meadows of Yo-
semite National Park, 77, 91; plant
communities of Ring Mountain Pre-
serve, 173; Prosopis in the San Joaquin
Valley, CA, 324; Scribneria bolanderi,
noteworthy collection, 381; Sequoia
sempervirens, fire history in old-growth
forest, 128; Silene suksdorfi, S. grayi,
and S. sargentii, systematic study of,
29: Soliva,228; vascular plants of east-
ern Imperial Co., 359; vegetation of

MADRONO, Vol. 34, No. 4, pp. 388-392, 1987

1987]

bald hills oak woodland, Redwood Na-

tional Park, 193; Vina Plains Preserve,

flora, 209; Washingtonia filifera cold

tolerance, 57.

New combinations: Dudleya abramsii
subsp. calcicola, 347; D. cymosa
subsp. paniculata, 338; D. cymosa
subsp. pumila, 336.

New records: Mannia fragrans, Mylia
anomala, Presissia quadrata, 69;
Myxomycetes, 48; Peteria thomp-
sonae, 381; Rhus trilobata var. sim-
plicifolia, 171.

New taxa: Allium shevockii, 150;
Clarkia concinna subsp. automixa,
41; Claytonia palustris, 155. Dud-
leya abramsii subsp. affinis, 349; D.
cymosa subsp. agourensis, 339; D.
cymosa subsp. crebrifolia, 344.

Range extensions: Festuca occidental-
is, Mimulus congdonii, 170.

California Botanical Society, early his-
tory, 1; message from past president,
383.

Campylium halleri, new record for Mex-
ico, 69.

Canada: Salix tweedyi, noteworthy col-
lection in British Columbia, 268.

Caryophyllaceae, systematic study of Si-
lene suksdorfu, S. grayi, S. sargentii,
29.

Ceanothus: C. crassifolius fruit produc-
tion, 273; C. cuneatus and C. leuco-
dermis seed dispersal in oak-woodland
savanna, 283.

Chaparral: Ceanothus crassifolius fruit
production, 273; role of fire in germi-
nation of herbs and suffrutescents, 240.

Chenopodiaceae: Grayia brandegei chro-
mosome races, 142.

New combination: Grayia brandegei
var. plummeri, 148.

Chromosome numbers: Allium she-
vockii, 152; Claytonia nevadensis, C.
palustris, 159; Dudleya abramsii subsp.
affinis, D. abramsii subsp. abramsii, D.
cymosa subsp. agourensis, D. cymosa
subsp. crebrifolia, D. cymosa subsp.
cymosa, D. cymosa subsp. pumila, 334;
Grayia brandegei, 142; Lessingia ten-
uis, 168; Tridax accedens, T. hintonii,
354.

Clarkia concinna subsp. automixa, new
subspecies from CA, 41.

Claytonia palustris, new species from CA,
155.

INDEX 389

Cleveaceae: Athalamia hyalina, new rec-
ord for Mexico, 69.

Cold tolerance in Washingtonia filifera,
Sie

Colloderma oculatum, new record for CA,
SI,

Colombia: New species: Alloispermum
insuetum, 162.

Colorado: New record: Bryum blindii,
171,
Comatricha: C. ellae, C. longipila, C.
penicillata, new records for CA, 52.
Commentary: message from Frank Al-
meda, past president of the California
Botanical Society, 384; Editor’s report
for vol. 34, 385.

Compositae (see Asteraceae).

Convolvulaceae: reproductive biology of
Ipomoea wolcottiana, 304.

Crassulaceae: some new and reconsid-
ered Dudleya from CA, 334.

Cribraria ferruginea, new record from CA,
50.

Cribrariaceae: Cribraria ferruginea, new
record from CA, 50.

Cymophora returned to Tridax, 354.

Desert plants: cold tolerance in Wash-
ingtonia filifera, 57; phosphorus and
pH tolerance in germination of Larrea
tridentata, 63.

Diderma effusum, new record from CA,
54.

Didymiaceae: Diderma effusum, Didum-
ium bahiense, D. verrucosporum, and
Lepidoderma aggregatum, new records
from CA, 54.

Didymium: D. bahiense and D. verruco-
sporum, new records from CA, 55.
Dudleya: New taxa or combinations: D.
abramsii subsp. affinis, 349; D. abram-
sii subsp. calcicola, 347; D. cymosa

subsp. agourensis, 339; D. cymosa
subsp. crebrifolia, 344; D. cymosa
subsp. paniculata, 338; D. cymosa

subsp. pumila, 336.

Ecuador: Buddleja americana, new fam-
ily for Galapagos Islands, 381.

Enteridium minutum, new record for CA,
49.

Entodon schleicheri, new record for Mex-
ico, 70.

Entodontaceae: Entodon schleicheri, new
record for Mexico, 70.

390

Erigeron ovinus, range extension in NV,
382.
Eucnide, inflorescence architecture, 18.

Fabaceae: Astragalus gilmanii, new rec-
ord for Nevada, 382; A. hypoxylus,
noteworthy collection from AZ, 170;
Peteria thompsonae, new record for CA,
381.

Festuca: F. minutiflora, new record for
New Mexico, 269; F. occidentalis, range
extension in CA, 170.

Fire: history in old-growth forest of Se-
quola sempervirens, 128; role in chap-
arral germination, 240.

Flora: eastern Imperial Co., CA, 359; Vina
Plains Reserve, Tehama Co., CA, 209.

Forest plants: distribution of forest trees
in northern Baja California, 98; Se-
quoia sempervirens, fire history in old-
growth forest of, 128.

Galapagos Islands, Buddleja americana,
new family for, 381.

Germination: chaparral, role of fire in,
240; Larrea tridentata, phosphorus and
pH tolerance in, 63; Pinus contorta var.
murrayana in mountain meadows, 77.

Grayia brandegei chromosome races, 142.

Grayia brandegei var. plummeri, new
combination, 148.

Humboldt Redwoods State Park: Se-
quoia sempervirens, fire history in old-
growth forest, 128.

Imperial Co., CA, vascular plants of, 359.
Ipomoea wolcottiana, reproductive bi-
ology of, 304.

Jungermanniaceae: Mylia anomala, new
record for CA, 69.

Larrea tridentata, phosphorus and pH
tolerance in germination of, 63.

Lepidoderma aggregatum, new record
from CA, 55.

Lessingia tenuis, range extension, chro-
mosome count, and mephitism, 168.

Letters: new species descriptions, 384.

Licea: L. lucens and L. operculata, new
records from CA, 48.

Liceaceae: Licea lucens and L. opercu-
lata, new records from CA, 48.

MADRONO

[Vol. 34

Loasaceae: Inflorescence architecture of
Eucnide, 18.

Lycogala exiguum, new record from CA,
49.

Lygodesmia_ grandiflora,
from NM, 171.

new record

Macbrideola: M. argentea and M. mar-

tinil, new records from CA, 51.

Mannia fragrans, new record for CA, 69.
Marchantiaceae: Presissia quadrata, new

record for CA, 69.

Mephitism in Lessingia tenuis, 168.
Mexico: Cymophora returned to Tridax,

354; distribution of forest trees in

northern Baja California, 98; repro-

ductive biology of Ipomoea wolcot-

tiana, 304.

New combination: Tridax hintonii,
357,

New records: Athalamia hyalina,
Campylium halleri, Entodon
schleicheri, Riccia albida, Timmia
megapolitana subsp. bavarica, 69.

New species: Axiniphyllum duran-
gense, 165.

Mimosaceae: Acacia smallii, in succes-
sional communities, 250; Prosopis in
the San Joaquin Valley, CA, 324.

Mimulus congdonii, range extension in
CA, 170.

Montane meadow plants: Pinus contorta
var. murrayana, germination, estab-
lishment, and invasion in mountain
meadows of Yosemite National Park,
7791.

Mycorrhizae, association with invasion
of Erechtites glomerata on San Miguel
Island, CA, 260.

Mylia anomala, new record for CA, 69.

Nevada:
New record: Astragalus gilmanii, 382.
Range extensions: Erigeron ovinus, Po-
lygala subspinosa var. heterorhyn-
cha, 382.
New Mexico: Penstemon ramosus, note-
worthy collection, 171.
New records: Festuca minutiflora, 269;
Lygodesmia grandiflora, 171; Salix
geyeriana, 268.

Oak woodland, bald hills, Redwood Na-
tional Park, CA, 193; Ceanothus cu-

1987]

neatus and C. leucodermis seed dis-
persal in Sierran savanna, 283.
Orbanchaceae: range and new locations
of Boschniakia strobilacea, 379.
Onagraceae: New subspecies: Clarkia
concinna subsp. automixa, 41.

Paradiacheopsis: P. cribrata, P. micro-
carpa, and P. rigida, new records for
CA, 53.

Penstemon ramosus, noteworthy collec-
tion from NM, 171.

Peteria thompsonae, new record for CA,
381.

Pinaceae: Pinus contorta var. murray-
ana, germination, establishment, and
invasion in mountain meadows of Yo-
semite National Park, 77, 91.

Pinus contorta var. murrayana, germi-
nation, establishment, and invasion in
mountain meadows of Yosemite Na-
tional Park, 77, 91.

Poaceae: Agrostis avenacea, noteworthy
collection in CA, 381; Festuca minu-
tiflora, new record for NM, 269; F. oc-
cidentalis, range extension in CA, 170;
Scribneria bolanderi, noteworthy col-
lection in CA, 381.

Polygala subspinosa var. heterorhyncha,
range extension in NV, 382.

Polygalaceae: Polygala subspinosa var.
heterorhyncha, range extension in NV,
382.

Portulacaceae Claytonia palustris, new
species from CA, 155.

Presissia quadrata, new record for CA,
69.

Prosopis in the San Joaquin Valley, CA,
324.

Redwood National Park, CA, bald hills
oak woodland, 193.

Reproductive biology: Ceanothus cras-
sifolius fruit production, 273; Steno-
cereus thurberi seedcrop characteristics
and minimum reproductive size in
southern AZ, 294: Ipomoea wolcot-
tiana, 304.

Reticulariaceae: Enteridium minutum
and Lycogala exiguum,new records
from CA, 49.

Reviews: T. C. Fuller and E. McClintock,
Poisonous plants of California, 382; S.
Goodrich and E. Neese, Uinta Basin

INDEX Syl

flora, 172; D. L. Magney, A flora of Dry
Lakes Ridge, Ventura County, Califor-
nia, 271; V. H. Oswald, Vascular plants
of Upper Bidwell Park, Chico, CA, 169;
P. H. Raven, H. J. Thompson, and B.
A. Prigge, Flora of the Santa Monica
Mountains, California, 70; J. Rze-
dowski and G. C. de Rzedowski, Flora
Fanerogamica del Valle de Mexico.
Volumen II. Dicotyledonae (Euphor-
biaceae-Compositae), 270; A. H.
Zwinger, Xdntus, the letters of John
Xantus to Spencer Fullerton Baird from
San Francisco and Cabo San Lucus,
1854-1861, 269.

Rhamnaceae: Ceanothus crassifolius fruit
production, 273; C. cuneatus and C.
leucodermis seed dispersal in oak-
woodland savanna, 283.

Rhus trilobata var. simplicifolia, new re-
cord for CA, 171.

Riccia albida, new record for Mexico, 69.

Ricciaceae: Riccia albida, new record for
Mexico, 69.

Ring Mountain Preserve, CA, plant com-
munities of, 173.

Salicaceae: Salix geyveriana, noteworthy
collection in NM; S. tweedyi, note-
worthy collection in British Columbia,
268.

Salix: S. geyeriana, noteworthy collec-
tion in NM, S. tweedyi, noteworthy
collection in British Columbia, 268.

San Joaquin Valley, CA, Prosopis in, 324.

San Miguel Island, CA, mycorrhizae as-
sociated with invasion of Erechtites
glomerata, 260.

Scribneria bolanderi, noteworthy collec-
tion in CA, 381.

Scrophulariaceae: Mimulus congdonii,
range extension in CA, 170; Penste-
mon ramosus, noteworthy collection
from NM, 171.

Seed dispersal of Ceanothus cuneatus and
C. leucodermis in oak-woodland sa-
vanna, 283.

Sequoia sempervirens, fire history in old-
growth forest, 128.

Sierra Nevada range: Allium shevockii,
new species from CA, 150; Ceanothus
cuneatus and C. leucodermis seed dis-
persal in oak-woodland savanna, 283;
Pinus contorta var. murrayana, ger-
mination, establishment, and invasion

B92

in mountain meadows of Yosemite
National Park, 77, 91.

Silene grayi, S. sargentii, S. suksdorfii,
systematic study of, 18.

Soliva in CA, 228.

Stemonitaceae: Amaurochaete comata,
Colloderma oculatum, Comatricha el-
lae, C. longipila, C. penicillata, Mac-
brideola argentea, M. martinii, Para-
diacheopsis cribrata, P. microcarpa, and
P. rigida, new records for CA, 51.

Stenocereus thurberi seedcrop character-
istics and minimum reproductive size
in southern AZ, 294.

Successional communities, Acacia smal-
lii in, 250.

Taxodiaceae: Sequoia sempervirens, fire
history in old-growth forest, 128.

Timmia megapolitana subsp. bavarica,
new record for Mexico, 70.

Timmiaceae: 7immia megapolitana

MADRONO

[Vol. 34

subsp. bavarica, new record for Mex-
ico, 70.

Trichia: T. macbridei and T. subfusca,
new records for CA, 50.

Trichiaceae: Arcyria magna, Trichia
macbridei, T. subfusca, new records for
CA, 50.

Tridax hintonii, new combination, 357.

Vina Plains Reserve, Tehama Co., CA,
flora, 209.

White Mountains, CA, alpine annual
plant species in, 315.

Yosemite National Park: Pinus contorta
var. murrayana, germination, estab-
lishment, and invasion in mountain
meadows, 77, 91.

Zygophyllaceae: phosphorus and pH tol-
erance in germination of Larrea tri-
dentata, 63.

ANNOUNCEMENT

NEW PUBLICATION

REGALADO, JR., J. C., R. K. RABELER, and J. H. BEAMAN, LABELS3
user’s manual: Guide to development of a collection database, Beal-
Darlington Herbarium, Dept. Botany and Plant Pathology, Michigan
State Univ., East Lansing, MI 48824-1312, 26 Jun 1987, [vi], 102 pp.
(+ 5.25” computer disk), ISBN 0-9617739-0-1 (paperbound), price un-
known. [““LABELS3 is a program package written in dBASE III Plus to
produce herbarium specimen databases and make labels. With the as-
sociated programs QUERY and CHKLIST, the database can be queried
for relevant information about a series of collections, and checklists can
be made for the entire database or selected segments. A collection note-
book can be generated from the database as an archival record of a set
of collections. .. . LABELS3 is more than a label production program.
Its primary objective is for developing and maintaining databases that
can be queried for relevant information about collections and from
which checklists can be produced.’’— from preface. Requirements: IBM
computer or compatible, 256 RAM, dBASE III Plus, WordStar.]

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MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

VOLUME XXxXV
1988

BOARD OF EDITORS

Class of:

1988—SuSAN G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA

1989— FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley

1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona-Sonora Desert Museum, Tucson

1991—JAMES HENRICKSON, California State University, Los Angeles
WAYNE R. FERREN, University of California, Santa Barbara

Editor—Davib J. KEIL
Biological Sciences Department
California Polytechnic State University
San Luis Obispo, CA 93407

Published quarterly by the
California Botanical Society, Inc.
Life Sciences Building, University of California, Berkeley 94720

Printed by Allen Press, Inc., Lawrence, KS 66044

Cornelius H. Muller, Professor of Botany, Emeritus, University
of California, Santa Barbara, and Adjunct Professor of Botany, Uni-
versity of Texas, Austin, has stamped a broad impression on Cali-
fornia botany. His personal influence has been a major force in
determining the direction of research and thought in ecology, sys-
tematics, and evolutionary biology. Much of the research reported

11

in Madrono was directly influenced by Dr. Muller and his students
or close associates.

Most people know Professor Muller as Ecologist or Systematist,
but few are aware of his international stature in both fields. He has
a record of accomplishment in a dozen or more subdisciplines, any
one of which would be viewed by many of us as a successful lifetime
achievement. Noteworthy examples include, but are not exhausted
by, the following: pioneer plant explorer of Mexico; early advocate
of the study of tropical botany; taxonomic authority in legumes,
Solanaceae, and oaks; guayule specialist; chemical analyst of plant
products; and botanical historian. He produced a definitive meth-
odology for the study and evaluation of allelopathy and demonstrat-
ed the fundamental importance of alleopathy in California vegeta-
tion dynamics, he demonstrated the importance of ecological control
in actual or potentially hybridizing populations, and he enunciated
fundamental principles of the structure and evolution of vegetation.

That the pages of Madrono so thoroughly reflect Dr. Muller’s
influence shows the power of his teaching. Several of his classes and
seminars are legendary at UCSB, but Dr. Muller taught all the time
whether or not he always recognized it. The twenty years that I spent
at his lunch desk provided my most broadening education! Hardly
a lunch hour escaped without some new research subject being enun-
ciated or some established ikon being devastated. The power of his
insight and intellect on these occasions was inspiring but also hum-
bling. A compliment from Dr. Muller was always genuine and stim-
ulating.

The breadth and power of Dr. Muller’s influence will continue to
be felt for generations. Even now, there is a re-awakening of study
of oaks that 1s multi-dimensional, but at the focal point of this
activity remain Dr. Muller’s established guidelines and new insights
continued to flow from this fount of knowledge.

To Cornelius H. Muller this volume of Madrofo is humbly and
respectfully dedicated. — DALE M. SMITH, Professor of Botany, Emer-
itus, University of California at Santa Barbara, Red Maple Farm,
HC-65, Box 100 BB, Windsor, KY 4256S.

il

TABLE OF CONTENTS

ALLEN, GERALDINE A. and JOSEPH A. ANTOos, Morphological and ecological
variation across a hybrid zone between Erythronium oreganum and E.
PEVOUAUTIU (WANA CCAC) oe aaa nce re ee

ANDERSON, BARRETT (See HOLLAND, DAN C.)

ANNABLE, CAROL R. (See PETERSON, PAUL M. and CAROL R. ANNABLE)

ANTOS, JOSEPH A. (see ALLEN, GERALDINE A.)

ARGUS, GEORGE W., Salix scouleriana (Salicaceae) discovered in Mexico .........

ARGUS, GEORGE W. and CARL-ERIC GRANFELT, Noteworthy collection of Salix
DIGNIOUG SSP DIGI OU Ge, eee ee

BOURELL, MONA and THOMAS F. DANIEL, Noteworthy collection of Carlowrigh-
LLAGQTIZOMICG, TROT: Cal iTO Ta eee ee eae eee a

CALDERON DE RZEDOWSKI, GRACIELA, AND JERZY RZEDOWSKI, Nota sobre el
genero Commelina (Commelinaceae) en el Valle de México con cambios
en la nomenclatura de algunas de SUS CSPCCIES oe cic eeeeccneeeeeeeeenneeeeeeennneeee

CARTER, ANNETTA, Review of Atlas Cultural de México. Flora by JERZY RZE-
DOWSKI And MIGUEL EQUIHUA 0ounu-n-sn-s--cn-cceccn-ssessneeoeeseesoessecsoessssoessnesovssssaessessnessessnessussaseseesene

CHAREST, NANCY (see CLARK, CURTIS)

CLARK, CURTIS, DONALD W. KyuHos, and NANCY CHAREST, A new Encelia
(Asteraceae: Heliantheae) from Baja California 20 e ccc

CONSTANCE, LINCOLN (see HARTMAN, RONALD L. and LINCOLN CONSTANCE)

CRITCHFIELD, WILLIAM B. (see MILLAR, CONSTANCE I.)

DANIEL, THOMAS F. (see BOURELL, MONA)

DAvIs, FRANK W., DIANA E. HICKSON, and DENNIS C. ODION, Composition of
maritime chaparral related to fire history and soil, Burton Mesa, Santa
Barbara County, Califomia. == 22 ee

DAvIs, OWEN K. and MICHAEL J. MoRATTO, Evidence for a warm dry early
Holocene in the western Sierra Nevada of California: pollen and plant
macrofossil analysis of Dinkey and Exchequer meadows eee eee

DEL MORAL, ROGER and DAviD M. Woop, The high elevation flora of Mount
St Pieler? W asia Otro ee,

DEMPSTER, LAURAMAY T., Three new species of Galium from the northern
PTE S ec a hgh een Bo

Dorn, RosBerT D., Typification of Chaenactis alpina (Asteraceae) ..................--

Dorn, RoBerT D., Chenopodium simplex, an older name for C. gigantosper-
VIULETIUN © WC TOP OC TACE AC) earner ree ene

DorN, ROBERT D. and RONALD L. HARTMAN, Nomenclature of Lomatium
nuttallii, L. kingii, and L. megarrhizum (Apiaceae) 2... eeeeeeeceeeeeeeeee eee

FANTZ, PAUL R., A new section of Clitoria subgenus Neurocarpum (Legumi-
TOS Ys Saran ia as ai ems IO De, Pre. eee rere

FERLATTE, WILLIAM J. (see STROTHER, JOHN L.)

GRANFELT, CARL-ERIC (see ARGUS, GEORGE W. and CARL-ERIC GRANFELT)

HARTMAN, EmMILy L. and MARY Lou RotTTMAN, Noteworthy collection of An-
tennaria aromatica (Astetaceae) 22a eee

HARTMAN, EmILy L. and MARY Lou ROTTMAN, The vegetation and alpine
vascular flora of the Sawatch Range, Colorado 2. ececcsecseeeeeccecneeeeeeeeeeeees

HARTMAN, RONALD L. (see DORN, ROBERT D. and RONALD L. HARTMAN)

HARTMAN, RONALD L. (see O’KANE, STEVE L., JR., DIETER H. WILKEN, and
RONALD L. HARTMAN)

HARTMAN, RONALD L. and LINCOLN CONSTANCE, A new Lomatium (Apiaceae)
from: the Sierran crestiof California ce oe ee

HENDERSON, DOUGLASS M. (see SCHAACK, CLARK G. and DOoUuGLAss M.
HENDERSON)

HICKSON, DIANA E. (see DAviIs, FRANK W.)

iV

32

pif)

169

162

202

HIRSCHBERG, JERILYN, and GEOFFREY A. LEVIN, Noteworthy collections of As-
pidotis densa and Holodiscus boursieri from California 2...
HOLLAND, DAN C. and BARRETT ANDERSON, Additional support for the recent-
invasive advent of mesquite (Mimosaceae: Prosopis) in the San Joaquin
AYE Cos Zk GNU Coy 9 00 ener eb cree etc ne eee ao ae nT ea
KEELER-WOLF, TODD, The role of Chrysolepis chrysophylla (Fagaceae) in the
Pseudotsuga-hardwood forest of the Klamath Mountains of California ....
KEIL, DAvip J., Editor’s report for volume 35 occ ceecccsseeeecccsseeeeseseneeeeseeneeeseeee
KyYHOoS, DONALD W. (see CLARK, CURTIS)
LACKSCHEWITZ, KLAUS, PETER LESICA, and J. STEPHEN SHELLY, Noteworthy
Collections minoma Wd aN > Aeterna lg ee ae
LANE, MEREDITH A., Generic relationships and taxonomy of Acamptopappus
(Compositae: AStereae)! a2 te se ee
LANE, THOMAS M. and Guy L. Nesom, Scutellaria lutilabia (Labiatae), a new
gypsophile from Nuevo Leon, Mexico 2icceccccccencseeeeeeceeeceeeennnneeeeeeeeecceceecnnnneneneneeeeeseecene
LEON DE LA Luz, JosE Luis, Noteworthy collections of Quercus oblongifolia
and Quercus arizonica from Baja California Sur, Mexico
LESICA, PETER (See LACKSCHEWITZ, KLAUS)
LEVIN, GEOFFREY A., Noteworthy collection of Astragalus pachypus from Cal-
JTL10 oC: Cg een hae INOS ene) Onc im MPN Eo sav ner T retro ee ane pe oven PI eee
LEVIN, GEOFFREY A. (see HIRSCHBERG, JERILYN)
McCarTEN, NIALL F. Review of Serpentine and Its Vegetation: A Multidisci-
plinary Approach by Robert Richard Brooks 2. ccc eeeeeeeeecneeeeeeeetenneeeeee
McCartTENn, NIALL F. and THOMAS R. VAN DEVENDER, Late Wisconsin vege-
tation of Robber’s Roost in the western Mojave Desert, California .............
McNEAL, DALE W., Message from the past CBS president
MEINKE, ROBERT J., Leptodactylon pungens subsp. hazeliae (Polemoniaceae),
a new combination for a Snake River Canyon endemic ee
MILLAR, CONSTANCE I. and WILLIAM B. CRITCHFIELD, Crossability and rela-
tionships Of PINUS MUPICALA (PIMACCAL) aan eesaneeeesnneecvvneeevvveeevveeessveeeesseesessseeessoeeeeee
MoraATTo, MICHAEL J. (see DAVIS, OWEN K.)
MOREFIELD, JAMES D., Noteworthy collections of Stylocline sonorensis from
LGPL 1) Coy 9 0 Fs emer rr oe eet ole openSSL oP SRC OTRO Un RUPE ee a
MOREFIELD, JAMES D. and DEAN WM. TAYLOR, Several noteworthy collections
from the White Mountains of California and Nevada occ
NEELEY, ELIZABETH E. (see O’KANE, STEVE L., JR., ELIZABETH E. NEELY, and
DIETER H. WILKEN)
Nesom, Guy L. (see LANE, THOMAS M.)
ODION, DENNIS C. (see DAvis, FRANK W.)
O’KANE, STEVE L., JR., ELIZABETH E. NEELY, and DIETER H. WILKEN, Note-
worthy collections fromy COlorddO. 2 ee ee
O’ KANE STEVE L., JR., DIETER H. WILKEN, and RONALD L. HARTMAN, Note-
WOU COMCCHOIMS EOI OLO LAC Ogre ee eee ene veer
PARFITT, BRUCE D., Review of Annotated Checklist of Vascular Plants of Grand
Canyon National Park 1987 by Barbara G. Phillips, Arthur M. Phillips,
Di andMarilyn Amn Schinid tfc oe: ee ee ee es
PARFITT, BRUCE D. and DONALD J. PINKAVA, Nomenclatural and systematic
reassessment of Opuntia engelmannii and O. lindheimeri (Cactaceae) .......
PEMBERTON, ROBERT W., The abundance of plants bearing extrafloral nectaries
in Colorado and Mojave Desert communities of southern California ........
PETERSON, PAUL M., Chromosome numbers in the annual Muhlenbergia (Po-
AR CS) Penh lr Rasa pea oe No ae eae IRs hg PN eae EU etre eI
PETERSON, PAUL M. and CAROL R. ANNABLE, Noteworthy collections from
OFT NT To) 3 5 yt: eens amen cle SOC en ee SE Ger ar? Were ree SPREE Ree Neer Seren
PETERSON, PAUL M. and CAROL R. ANNABLE, Noteworthy collections from
INV USED ITN CU) Tiree PR ae ees Smet at tea, eh tea cardio, en hat Speedin ote Pera ate

355

I2

PINKAVA, DONALD J. (see PARFITT, BRUCE D. and DONALD J. PINKAVA)

PRESTON, ROBERT E., Arabis breweri var. austinae (Cruciferae) i eeeeeeeeeooone

REEDER, CHARLOTTE G. (see REEDER, JOHN R.)

REEDER, JOHN R. and CHARLOTTE G. REEDER, Hilaria annua (Gramineae), a
TOW SPECIES Hr Tun IVN Cx CO eee a eer rere ue meses nee ene

RIGGINS, RHONDA, A comparison of North and South American Lupinus group
IVITCHOCAT DI: (UEC R TAT ING SAG) ctr seneeoee ereeeeet neeeeee

ROTTMAN, MAry Lou (see HARTMAN, EMILY L. and MAry Lou ROTTMAN)

RZEDOWSKI, JERZY (see CALDERON DE RZEDOWSKI, GRACIELA)

SANDERSON, MICHAEL J., Astragalus nutriosensis (Fabaceae): a new species from
CASULA TZ O Mek 5 2s ce Pra Neer es ets are es

SAWYER, JOHN O., JR. (see SMITH, JAMES P., JR.)

SCHAACK, CLARK G., Noteworthy collection of Aquilegia triternata x A. chry-
R12 ea EE EE REL OE NE SE aA TY ME OP ror SO Tee nent Peon

SCHAACK, CLARK G. and DouGLAss M. HENDERSON, Noteworthy collection of
Cardamine constancel troim IdalnQyc2. e

SCHEID, GERALD A. (see ZEDLER, PAUL H. and GERALD A. SCHEID)

SCHMID, RUDOLF, Review of The Plant-Book: A Portable Dictionary of the
Hicsher Piants by D.). Mabbetley = ee) ee ee

SCHOOLCRAFT, GARY (see TIEHM, ARNOLD)

SHELLY, J. STEPHEN (see LACKSCHEWITZ, KLAUS)

SILVA, PAUL C., Report on the XIV International Botanical Congress ...........

SKELLY, Rick J., A new species of Saxifraga (Saxifragaceae) from the Olympic
Mountains, Washington, and Vancouver Island, British Columbia ..............

SMITH, JAMES P., JR. and JOHN O. SAWYER, JR., Endemic vascular plants of
northwestern California and southwestern Oregon 2c

STROTHER, JOHN L. and WILLIAM J. FERLATTE, Review of Erigeron eatonii and
allied taxa:(Compositac: ASteme ae) ese recs ee ieee

TAYLOR, DEAN WM. (see MOREFIELD, JAMES D. and DEAN WM. TAYLOR)

TAYLOR, RONALD J. (see WAGSTAFF, STEVEN J.)

TIEHM, ARNOLD and GARY SCHOOLCRAFT, Several noteworthy collections from
INevadaand (Oregon: 2 eee ee re

VAN DEVENDER, THOMAS R. (see MCCARTEN, NIALL F. and THOMAS R. VAN
DEVENDER)

VorA, Rosin S., Species frequency in relation to timber harvest methods and
elevation in the pine type of northeast Califormia 20 eeeeee

WAGSTAFF, STEVEN J. and RONALD J. TAYLOR, Genecology of Cerastium arvense
and C. beeringianum (Caryophyllaceae) in northwest Washington __...........

WEBSTER, GRADY L., A new species of Croton (Euphorbiaceae) from Nicaragua

TSU Of GANGES 11S LACS ere ee eee

WILKEN, DIETER H. (see O’KANE, STEVE L., JR., ELIZABETH E. NEELY, and
DIETER H. WILKEN)

WILKEN, DIETER H. (see O’ KANE, STEVE L., JR., DIETER H. WILKEN, and RONALD
L. HARTMAN)

Woop, DAvIpD M. (see DEL MORAL, ROGER)

ZEDLER, PAUL H., Review of Conservation and Management of Rare and En-
aangered Planis:by Thomas:s. Blase. = ee

ZEDLER, PAUL H. and GERALD A. SCHEID, Invasion of Carpobrotus edulis and
Salix lasiolepis after fire in a coastal chaparral site in Santa Barbara County,
GUT Po) 5 01 ¢- Seeene ene Raieeemer Reet SMC rk Pee mn aN or ARR on ee eee Aer ns

vl

92

325

278

354

167

a9

126

54

gl

166

150

266

117

330

280

/VOLUME 35, NUMBER 1 JANUARY-MARCH 1988

jes
4 =

ADRONO

A WEST AMERICAN JOURNAL OF BOTANY

NS

Contents

THREE NEw SPECIES OF Galium FROM THE NORTHERN AN j
Lauramay T. Dempster i

Hilaria annua (GRAMINEAE), A NEW SPECIES FROM MEXxIcc

John R. Reeder and Charlotte G. Reeder 6
A New Encelia (ASTERACEAE: HELIANTHEAE) FROM BAJA CADE

Curtis Clark, Donald W. Kyhos, and Nancy Charest : 10
NOTA SOBRE EL GENERO Commelina (COMMELINACEAE) EN EL VAEET DE MEXICO CON

CAMBIOS EN LA NOMENCLATURA DE ALGUNAS DE SUS ESPECIES

Graciela Calderon de Rzedowski and Jerzy Rzedowski 16
A New SECTION OF Clitoria SUBGENUS Neurocarpum (LEGUMINOSAE)

Paul R. Fantz abe)
MORPHOLOGICAL AND ECOLOGICAL VARIATION ACROSS A HYBRID ZONE BETWEEN

Erythronium oregonum AND E. revolutum (LILIACEAE)

Geraldine A. Allen and Joseph A. Antos 32
CROSSABILITY AND RELATIONSHIPS OF Pinus muricata (PINACEAE)

Constance I. Millar and William B. Critchfield 39
ENDEMIC VASCULAR PLANTS OF NORTHWESTERN CALIFORNIA AND SOUTHWESTERN

OREGON

James P. Smith, Jr. and John O. Sawyer, Jr. 54
NOTES
NOMENCLATURE OF Lomatium nuttallii, L. kingii, AND L. megarrhizum (APIACEAE)

Robert D. Dorn and Ronald L. Hartman 70
NOTEWORTHY COLLECTIONS

ARIZONA =)

COLORADO 12
REVIEWS 74
ANNOUNCEMENTS OR lay 22609... 6

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
ciety, Inc., and is issued from the office of the Society, Herbarium, Life Sciences
Building, University of California, Berkeley, CA 94720. Subscription rate: $25 per
calendar year. Subscription information on inside back cover. Established 1916.
Second-class postage paid at Berkeley, CA, and additional mailing offices. Return
requested. POSTMASTER: Send address changes to James R. Shevock, Botany Dept.,
California Academy of Sciences, San Francisco, CA 94118.

Editor—Davip J. KEIL

Biological Sciences Department
California Polytechnic State University
San Luis Obispo, CA 93407

Board of Editors
Class of:

1988—SuSAN G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989— FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991 —JAMES HENRICKSON, California State University, Los Angeles
WAYNE R. FERREN, JR., University of California, Santa Barbara

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1987-88

President: DALE MCNEAL, Department of Biological Sciences, University of the
Pacific, Stockton, CA 95211

First Vice President: PEGGY FIEDLER, Department of Biology, San Francisco State
University, 1600 Holloway Ave., San Francisco, CA 94132

Second Vice President: STEVEN TIMBROOK, Ganna Walska Lotusland Foundation,
695 Ashley Rd., Montecito, CA 93108

Recording Secretary: V. THOMAS PARKER, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
Francisco, 50 Phelan Ave., San Francisco, CA 94112

The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, FRANK ALMEDA, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118; the Editor of MADRONO; three elected
Council Members: ANNETTA CARTER, Department of Botany, University of Califor-
nia, Berkeley, CA 94720; JOHN Moorinc, Department of Biology, University of
Santa Clara, Santa Clara, CA 95053; BARBARA ERTTER, University Herbarium, De-
partment of Botany, University of California, Berkeley, CA 94720; and a Graduate
Student Representative, NIALL F. MCCARTEN, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132.

THREE NEW SPECIES OF GALIUM FROM
THE NORTHERN ANDES

LAURAMAY T. DEMPSTER
Jepson Herbarium, Department of Botany,
University of California, Berkeley 94720

ABSTRACT

Three new species of Ga/ium are named and described: G. antuneziae of Peru, with
hairless fruits, shows some similarities to G. mandonii Britton, G. cajamarcense from
Peru, also with hairless fruits, has a general resemblance to G. corymbosum R. & P.
of sect. Relbunium, but lacks the involucre diagnostic of that section; G. fosbergii of
Ecuador seems closest to G. diffusoramosum Dempster & Ehrendorfer, although the
hairs are reduced in number and not uncinate.

RESUMEN

Se nombran y describen tres nuevas especies de Galium: G. antuneziae de Peru,
con frutos sin tricomas, muestra semejanzas con G. mandonii Britton, G. cajamar-
cense de Peru, también con frutos sin tricomas, se asemeja a G. corymbosa R. & P.
de la seccion Re/bunium, pero carece del involucro diagnostico de esa seccion; G.
fosbergii de Ecuador al parecer se relaciona con G. diffusoramosum Dempster &
Ehrendorfer, aunque las tricomas de los frutos son escasos y no uncinulados.

In the course of my work on Galium sect. Relbunium, I have
located three specimens that do not belong to that section nor to
any named species of Galium. This paper is, therefore, a codicil to
my three longer papers on South American Galium (Dempster 1980,
1981, 1982). I describe and name herein three new species based on
these specimens.

Galium antuneziae Dempster, sp. nov. (Fig. 1).

Herba perennis semiprostrata, caulibus lignosis flexibilibus. Foliis
in quoque verticillo quatuor, ellipticis, petiolo brevi. Floribus so-
litariis axillaribus pedunculo brevi; corollis rotatis, profunde divisis,
albis, latitudine 2.5 mm; lobis ovatis apice gracili; ovaris fructib-
usque subtiliter tuberculatis.

Semi-erect perennials with long (to 50 cm) flexible woody stems
ending in tufts of fine-textured herbage; young stems with narrow
angles having short urceolate hairs. Leaves 4 at nodes, less than 8
mm long, elliptical, shortly petiolate with few short stoutish curved
hairs, mostly on reflexed margins, ribs and pulvinate bases. Flowers
perfect, solitary in axils, on short peduncles, scarcely exserted beyond
leaves; corollas white, rotate, 2.5 mm across, deeply divided, the 4

MADRONO, Vol. 35, No. 1, pp. 1-5, 1988

®) MADRONO [Vol. 35

Fic. 1. Galium antuneziae. a. Flowering branchlet. b. Node with solitary flower.
c. Young fruit. All illustrated from type, Antunez de Mayolo 110.

lobes ovate with short attenuated tips, glabrous externally, the in-
terior surfaces with many glandular tack-shaped hairs scattered on
the apical half; ovaries and fruits very finely tuberculate; fruits dry,
3 mm across; mericarps partially spherical, but flattened at juncture.

TYPE: Peru, Huancavelica, near Churcampa, 2500 m, on slopes
bordering cultivated fields, Antunez de Mayolo 110 (holotype: F;
isotype: OBI).

PARATYPE: Near Churcampa at 3100 m, Antunez de Mayolo 248
(UC), erroneously attributed to G. mandonii Britton (Dempster 1982).

Galium antuneziae shows some similarity to G. mandonii in that
the leaves of both are narrowed to a petiole above a hairy pulvinate
base, have narrow stem angles with numerous small retrorse hairs,
and have short-pedicelled flowers solitary in axils. Galium mandonii,
however, differs in having lax herbaceous stems above a fibrous root
system, smaller corollas relative to ovaries, larger, narrower and
more remote leaves, and 8-sulcate ovaries.

1988] DEMPSTER: NEW GALIUM SPECIES 3

a

Fic. 2. Galium cajamarcense. a. Node. b. Flowering branchlet. Illustrated from
type, Beck 7880.

The name is for the collector, whom I met in Peru in 1977, where
she was studying indigenous dye plants, including Galium.

Galium cajamarcense Dempster, sp. nov. (Fig. 2).

Herba perennis caespitosa caulibus foliisque pilis longis patentibus
dispersis. Foliis in quoque verticillo quatuor, usque ad 4 mm, ob-
longis, internodia aequantibus. Floribus paucis, pedicellatis, ramulis
brevibus foliosis insidentibus; corollis rotatis, profunde divisis, lobis
ovatis obtusis; ovarus laevibus. Fructibus ignotis.

Tufted perennials to 6 cm tall, from creeping root system; stems
and leaves with scattered long spreading hairs. Leaves 4 at nodes,
ca. 4 mm long, about as long as internodes, oblong with abruptly
acute unarmed apices; glandular cells in a dense subapical cluster.
Flowers few, pedicellate, on short leafy axillary branchlets; corollas
rotate, 2 mm across, deeply divided, the four lobes ovate, obtuse,
glabrous; ovaries smooth. Fruits unknown.

TYPE: Peru, Dept. Cajamarca, 15 km southwest of Cajamarca at
edge of Cumbe Mayo Park, 3400 m, “‘Cesped abierto pedregoso’’,
Beck 7880 (holotype: UB).

This species bears a general resemblance to forms of G. corym-
bosum Ruiz & Pavon sensu lato of sect. Re/bunium. Unlike that
complex species, however, the flowers of G. cajamarcense are ped-
icellate and without involucres.

Galium fosbergii Dempster, sp. nov. (Fig. 3).

Herba perennis scandens polygama, caulibus gracilibus lignosis;
foliis in quoque verticillo quatuor, usque ad 6 mm, ovatis apicem
tenuem versus angustatis, petiolatis, base tomentosis, margine cos-
taque et pagina superna pilis brevibus antrorsis instructis; floribus
paucis ramulis insidentibus; pedicellis capillaribus usque ad 9 mm,
exsertis; corollis rotatis, profunde divisis, lobis ovatis, obtusis; fruc-
tibus immaturis, pilis rectis sparse instructis.

4 MADRONO [Vol. 35

Fic. 3. Galium fosbergii. a. Node, showing under side of 3 leaves. b. Leaf, upper
side, showing triple venation. c. Staminate flower. d. Same from below. e. Immature
fruit. All illustrated from type, Fosberg and Giler 22945.

Lax, monoecious or polygamous perennials with slender woody
stems, clambering or climbing to 35 cm; stems sparsely set with
short retrorse hairs. Leaves 4 at nodes, up to 6 mm long, ovate,
tapered to a slender apex and narrowed to a petiole above a hairy,
often swollen and sometimes persistent base; leaves more or less
obscurely 3-nerved, the upper surfaces set with short antrorse hairs,
the lower surfaces glabrous except for the midribs. Flowers on few-
flowered well exserted axillary branches; pedicels capillary, 1.5-9
mm long, well exserted beyond bracts; corollas rotate, 2 mm across,
divided to near base, glabrous, the lobes ovate, obtuse. Fruits (im-
mature) sparsely set with very short, straight, slightly antrorse
spreading hairs.

Type: Ecuador, Loja, northeast slope of Cerro Mataperro, 1815
m, steep dry slope of decomposed slaty shale, tangled in bushes,
Fosberg and Giler 22945 (holotype: US).

The often swollen and persistent leaf bases of G. fosbergii suggest
a relationship with G. diffusoramosum Dempster & Ehrendorfer, of
northern Chile. Characters of inflorescence, flowers, and leaves also
are compatible with that species. The fruit hairs of G. fosbergii,

1988] DEMPSTER: NEW GALIUM SPECIES 5

however, are not uncinate but, although sparse, they are straight as
in sect. Lophogalium. Although fruit hairs are typically of taxonomic
importance in Galium, I think that G. fosbergii represents a reduction
in number and shape of hairs, rather than a closer relationship with
sect. Lophogalium. The large geographical distance between G. /fos-
bergii and G. diffusoramosum (ca. 2800 km) precludes the consid-
eration of the former as an aberrant form of the latter.

ACKNOWLEDGMENTS

Thanks are owing to Alan Smith for criticising my Latin, and to Victoria Sosa for
correcting my Spanish.

LITERATURE CITED

Dempster, L. T. 1980. The genus Galium section Lophogalium (Rubiaceae) in South
America. Allertonia 2:247-279.

1981. The genus Galium (Rubiaceae) in South America. II. Allertonia 2:

393-426.

. 1982. The genus Galium (Rubiaceae) in South America. II. Allertonia 3:

211-258.

(Received 10 Nov 1986; revision accepted 4 Oct 1987.)

NOTEWORTHY COLLECTION

ARIZONA

SALIX PLANIFOLIA Pursh. ssp. PLANIFOLIA (SALICACEAE).— Apache Co., Fort Apache
Indian Reservation, White Mts., Smith Cieniga, along Ord Creek, 33°56'N, 109°35'W,
9890-10,000 ft, dominant in Salix thicket along creek, associated with S. arizonica,
wet saturated soils of igneous origin, heavily browsed by elk, 9 Jul 1987, C.-E. Granfelt
87-1 (ARIZ, CAN). White Mts., Bull Cieniga, along Ord Creek, 33°55’30’N,
109°35'30’W, 10,240 ft, only Salix in saturated meadows and on slopes adjacent to
creek, closely browsed by elk, 29 Jul 1987, C.-E. Granfelt 87-24, 87-25, 87-26 (CAN).
Identified by George W. Argus.

Previous knowledge. Trans-subarctic-boreal from Alaska to Newfoundland south
to New Hampshire and British Columbia extending southward in the cordillera to
California and the mountains of northern New Mexico and Utah. Inclusion of Arizona
in range given by Martin and Hutchins (Flora New Mexico, 1980) is undocumented.

Significance. New to the flora of Arizona. In Arizona this species seems to occur
at somewhat lower elevations (10,000—10,240 ft) than it does in nc. New Mexico
where it occurs at 10,500—11,500 ft. The species in the southern cordillera has usually
been referred to var. monica (Bebb) C. Schneider. The status of this taxon is under
study.— GEORGE W. ArGus, National Herbarium, Museum of Natural Sciences, Ot-
tawa, ON K1A 0M8, Canada and CARL-ERIC GRANFELT, Box 630, Pinetop, AZ 85935.

MApbrRONO, Vol. 35, No. 1, p. 5, 1988

HILARIA ANNUA (GRAMINEAE), A NEW
SPECIES FROM MEXICO

JOHN R. REEDER and CHARLOTTE G. REEDER
Herbarium, University of Arizona, Tucson 85721

ABSTRACT

Hilaria annua from the state of Colima, Mexico, is described as new. This, the first
annual species known in the genus, is clearly related to H. ciliata (Scribn.) Nash. It
differs from that species in its annual habit, somewhat smaller spikelets, and chro-
mosome number which is tetraploid (2/ = 36) rather than octoploid (2n = 72).

RESUMEN

Se describe Hilaria annua del estado de Colima, México como especie nueva. Se
trata de la primera especie anual conocida para el género, la cual es similar a H.
ciliata (Scribner) Nash. No solo se distingue por su condicion anual, sino también
por tener espiguillas algo mas pequenas y por el numero cromosomico. La especie
nueva es tetraploide (2n = 36) y H. ciliata es octoploide (2n = 72).

The genus Hi/aria (s.1.) comprises a small group of grasses that
inhabit arid and semi-arid regions and range from southwestern
United States to Guatemala. The inflorescence 1s a narrow terminal
spike, with spikelets borne in groups (fascicles) of three at the nodes,
falling entire from the axis when mature. In each fascicle the two
lateral spikelets are staminate and usually at least 2-flowered; the
central one is 1-flowered, pistillate or perfect. Species of Hilaria fall
quite naturally into two groups which currently are usually treated
as subgenera. The distinctions characterizing these subgenera are
summarized in the following key:

1. Glumes thin, membranous, not fused nor indurate at their bases;
central spikelet 1-flowered, perfect ...... Subgenus Pleuraphis
1. Glumes indurate, fused at their bases; central spikelet pistillate
Pe aA ad Sete a a ee ea ee ae Subgenus Hilaria

In the most recent revision of the genus Hi/aria (Sohns 1956),
nine species and one variety are recognized. Sohns stated that taxa
in both subgenera are remarkably uniform vegetatively: 1.e., all are
strong perennials and most are either stoloniferous or rhizomatous.
It was somewhat of a surprise, therefore, when we encountered what
appeared to be a weedy annual Hilaria growing in abundance in two
different areas a short distance south of Cd. Colima, México. These
plants are members of the subgenus Hi/aria and superficially resem-
ble H. ciliata (Scribn.) Nash, a species also found in that region.

MApDRONO, Vol. 35, No. 1, pp. 6-9, 1988

1988] REEDER AND REEDER: HILARIA ANNUA y

They differ not only in being annual, but in having smaller spikelets
and a chromosome number of 2” = 36. All chromosome counts of
H. ciliata reported to date are 2n = 72.

Our gatherings were made in 1974. Since that time we have seen
no other collections of an annual Hilaria, nor does McVaugh (1983)
mention any annual species in his treatment of this genus. Differ-
ences which separate plants of our collections from others in the
group, however, suggest that they represent a previously unrecog-
nized species which is described below.

Hilaria annua J. & C. Reeder, sp. nov. (Fig. 1).

Gramina annua, caespitosa; culmi 40—S50(—60) cm alti, erecti vel
interdum geniculato-adscendentes, nodis radicantes, papilloso-pi-
losis; culmi gracili, glabri, ramosi. Vaginae glabrae vel plus minusve
papilloso-pilosae, quam internodiis breviores; ligula membranaceo-
hyalina, ciliata, 2-3 mm longa; laminae 10—20(—30) cm longae, usque
ad 3.5-5 mm latae, planae, plerumque glabrae sed supra interdum
sparsim papilloso-pilosae, marginibus scabrae. Spicae usque ad 5
cm longae, densiflorae, articuli rhachis 44.5 mm longi, plani, gra-
cillimi, ca. 0.3 mm lati, marginibus brevi ciliati; fasciculi 4—4.5 mm
longi, pallidi vel niger-purpurascentes. Glumae induratae, papillo-
sae, plus minusve valde nervosae, marginibus scabri vel brevi ciliati;
spiculae laterales masculae, plerumque uniflores, interdum bi- vel
triflores, lemmata hyalina, ca. 3.5 mm longa, 3-nervis. Antheris ca.
2 mm longis; spicula intermedia uniflora, feminina, lemmata am-
pulliforma, 3-nervis, ca. 4.5 mm longa, basi hyalini, apice aliquanti
apicibus membranacibus. Caryopside translucida, ca. 1.8—2 mm lon-
ga, embryo fuscus, caryopsidi fere aequilongus. Chromosomatum
numerus: 2n = 36.

Plants annual, caespitose; culms 40—50(—60) cm tall, slender, gla-
brous, branching, erect to somewhat ascending and rooting at the
papillose-pilose nodes. Sheaths shorter than the internodes, glabrous
or more or less papillose-pilose; ligule membranous-hyaline, ciliate,
2-3 mm long; blades flat, 10—20(-30) cm long, 3.5-5 mm wide,
mostly glabrous but sometimes sparsely papillose-pilose on the adax-
ial surface, the margins scabrous. Spikes (3.5—)4—5 cm long, densely
flowered, the rachis joints flat, slender, 2.5—4 mm long and ca. 0.3
mm wide, the margins ciliolate; fascicles 3.5—4.5 mm long, pale or
becoming somewhat blackish-purple. Glumes indurate, minutely
papillose, the nerves prominent, especially toward the apex, the
margins scabrous to ciliolate; lateral spikelets staminate, mostly
1-flowered, sometimes 2- or 3-flowered, lemma hyaline, 3-nerved,
about as long as the glumes. Anthers ca. 2 mm long; central spikelet
pistillate, 1-flowered, the glumes slightly shorter than the fascicle,
with one (rarely two) lateral, flattened, scabrous to short-ciliate awns,

[Vol. 35

~

MADRONO

9

S

ES ee ee, ee res ere Leer | Aarne Cee

Hilaria annua. Photograph of the holotype (J. R. & C. G. Reeder 6333).

Fia. 1.

1988] REEDER AND REEDER: HILARIA ANNUA 2

these reaching to the apex of the glume; lemma flask-shaped, ca. 4.5
mm long, 3-nerved, basal portion hyaline, the upper part somewhat
membranous. Caryopsis whitish, translucent, 1.8—2 mm long, the
embryo brownish, nearly as long as the grain. Chromosome number:
2n = 36.

Type: MEXICO: Colima, 10 km s. of Cd. Colima, abundant along
roadside with other rank weeds, 300 m, 24 Sep 1974, J. R. & C. G.
Reeder 6333 (holotype: ARIZ; isotypes: MEXU, MICH, RM, US).

PARATYPE: MEXICO: Colima, 5 km s. of Cd. Colima, frequent
with other weeds amongst thorny shrubs, 430 m, 24 Sep 1974, J.
R. & C. G. Reeder 6331 (ARIZ, RM, UC, US). This collection
certainly represents the same species, but exhibits minor differences.
The plants are clearly annual, but more profusely branched, and the
culms tend to be more ascending, some of them rooting at the lower
nodes. The shorter spikes average 2—3 cm long rather than 4—5, and
the fascicles are slightly smaller, averaging 3.5—4 mm long, rather
than 4.0-4.5 mm as in the type. The chromosome number of this
collection also was determined as 2n = 36.

LITERATURE CITED

McVaAuGH, R. 1983. Flora Novo-Galiciana. A descriptive account of the vascular
plants of western Mexico. Vol. 14. Gramineae. Univ. Michigan Press, Ann Arbor.

SouNS, E. R. 1956. The genus Hilaria (Gramineae). J. Wash. Acad. Sci. 46:31 1-
321.

(Received 30 Mar 1987; revision accepted 30 Sep 1987.)

ANNOUNCEMENT

NEw PUBLICATION

GRIFFIN, J. R., P. M. MCDONALD, and P. C. Muick, compilers. 1987.
California oaks: a bibliography. U.S.D.A. Forest Service, Pacific
Southwest Forest and Range Experiment Station, Gen. Tech. Rep.
PSW-96, Berkeley, CA. 38 pp. [California oaks continue to attract
considerable attention among natural resource professionals. This

report provides a comprehensive bibliography of the extensive but
scattered oak literature. The 768 references are organized into two
systems: (a) a topical outline, in which references are displayed under
key word headings and subheadings, and author-date entries that help
to locate items by researcher or date; and (b) a Quercus species index,
in which references contain serial numbers for all species and hybrids.
Single copies are available from Pacific Southwest Forest and Range
Experiment Station, 1960 Addison Street, Berkeley, CA 94704.]

A NEW ENCELIA (ASTERACEAE: HELIANTHEAE)
FROM BAJA CALIFORNIA

CURTIS CLARK
Biological Sciences, California State Polytechnic University,
Pomona 91768

DONALD W. KYHOS
Botany, University of California, Davis 95616

NANCY CHAREST
Biological Sciences, California State Polytechnic University,
Pomona 91768

ABSTRACT

Encelia densifolia Clark & Kyhos, from the isolated Picachos de Santa Clara in
northern Baja California Sur, is morphologically distinctive, having short peduncles,
broad obovate phyllaries, and remotely dentate leaves.

RESUMEN

Encelia densifolia Clark & Kyhos, de los aislados Picachos de Santa Clara en el
norte de Baja California Sur, es distinto morfologicamente, con pedunculos cortos,
filarios largos y obovados, y hojas remotamente dentadas.

In 1947, H. S. Gentry made two collections of a new Encelia from
the Picachos de Santa Clara that he later labeled ‘“‘Encelia densifo-
lia’’, but never formally published. We have recollected the species
and studied it in the field and in cultivation to understand better its
relationships with other members of the genus, and describe it here.

Encelia densifolia Clark & Kyhos, sp. nov.
Encelia densifolia H. S. Gentry, nom. ined. herb., Gentry 7757, 5—
10 Nov 1947.

Frutex erectus vel aliquantum effusus, usque ad 1.5 m altus. Folia
14-65 mm longa, 7-35 mm lata, griseo-prasina, pilosa, rigide di-
vergentia vel ascendentia, ovata, plerumque dentata, petiolis alatis.
Capitula solitaria, floribus 20-30 mm latis trans flores radiantes,
fructibus plerumque pendula. Pedunculi breves (7-30 mm longi),
minute pubentes. Involucra 11-15 mm longa, 9-13 mm lata. Phyl-
laria 3-6 mm lata, 7-10 mm longa, imbricata, obovata, minute
pubentia, marginibus ciliatae. Flores radiantes 7—12 mm longi, flavi,
lucem ultraviolaceum reflectentes. Flores disci flavi, lucem ultra-
violaceum absorbentes. Antherae porphyreae aureaeve. Stigmata fla-
va. Achenia obovata, compressa, in superficie plana glabra pro parte

MADRONO, Vol. 35, No. 1, pp. 10-15, 1988

1988] CLARK ET AL.: NEW ENCELIA 11

Fics. 1, 2. Encelia densifolia (Clark 585). 1. Capitulum in flower. 2. Capitulum
in early fruit, showing pendulous habit and enlarged phyllaries. Photographs taken
in the field in the vicinity of the type specimen (x 2.2).

2 MADRONO [Vol. 35

Lee

Fics. 3, 4. Encelia densifolia. 3. Leaf (transilluminated to emphasize venation),
x2. 4. Achene, x10. Photographs taken of plants grown from seed (Clark 585) in
outdoor cultivation.

maxima, in margine ciliata, sine aristis, ad apicem incisura lata et
non profunda. Chromosomatum gametophytorum numerus 18 (Figs.
1-4).

Erect or occasionally spreading shrub, to 1.5 m tall. Leaves 14—
65 mm long, 7-35 mm wide, gray-green, pilose, rigidly divergent or
ascending, ovate, usually remotely dentate, petioles winged. Capitula
solitary, 20-30 mm wide across the rays in flower, pendulous in
fruit. Peduncles short (7-30 mm long), minutely pubescent. Invo-
lucre 11-15 mm long, 9-13 mm wide. Phyllaries 3-6 mm wide, 7—
10 mm long, imbricate, obovate, minutely pubescent, margins cil-
late. Ray florets 7-12 mm long, yellow, reflecting ultraviolet. Disc
florets yellow, absorbing ultraviolet. Anthers brown or yellow. Stig-
mas yellow. Achenes obovate, flattened, mostly glabrous on the face,
ciliate on the margin, without awns, with a broad shallow apical
cleft. Chromosome number nv = 18.

Type: Mexico, Baja California Sur: Picachos de Santa Clara, 13.6
mi nw. of San Ignacio-Abreojos road at a point 24.7 mi ne. of Punta
Abreojos, 300 m, 24 Mar 1981, C. Clark 585 (holotype: CAS; iso-
types: DAV, CSPU, RSA, UC; all material is from the same plant).

PARATYPES: Mexico, Baja California Sur: n. slope and in arroyo,

1988] CLARK ET AL.: NEW ENCELIA 13

Picachos de Santa Clara, 28 Dec 1975, Moran 22758 (DAV, SD).
Las Tinajas and vicinity in cerros e. of Los Picachos de Santa Clara,
21-23 Mar 1947, H. S. Gentry 7560 (SD). Picachos de Santa Clara,
Arroyo de los Picachos, 5-10 Nov 1947, H. S. Gentry 7757 (SD).

Distribution and habitat. Existing collections of the species may
correspond to only two populations. Clark 585 and Moran 22758
are from the same location. We suspect Gentry 7757 is also from
this location; Arroyo de los Picachos may be Arroyo Tecolote, which
is the major southeast drainage of the range (Moran pers. comm.)
and the location of the other two collections. Gentry 7560 represents
the other known population. It is possible that there are other un-
discovered populations in this rugged mountain range, but the species
does seem to be endemic to it.

The holotype grew along a dry stream-course. Over 100 plants
were found up to 30 m higher on the gravely slopes above it, pri-
marily on the north-facing shoulder of the southeastern-most peak.
Surrounding vegetation was desert scrub, and E. densifolia was co-
dominant with species of Bursera, Ferocactus, and Lycium.

Morphology. Encelia densifolia can easily be distinguished from
other species in the genus. The peduncles are short enough that the
heads are partially submerged in the leaves, a feature otherwise found
only in E. ventorum, which has finely divided leaves (Kyhos et al.
1981). The remote dentation of the leaf margin and the obovate
phyllaries are unique in the genus.

The phyllaries are the broadest in the genus. Werk and Ehleringer
(1983) found that photosynthesis by phyllaries and paleae in E.
farinosa and E. californica did not contribute much to the overall
energy budget of the plant. We suspect the form of the E. densifolia
phyllaries, in conjunction with the generally nodding fruiting heads,
may be an adaptation for increased photosynthesis by exposing these
organs to direct sunlight.

The leaf and phyllary pubescence consists of multicellular uni-
seriate hairs of the sort that are ubiquitous in the Heliantheae. The
junctions between cells are slightly swollen (Fig. 5). Although the
trichomes form a continuous covering over the surface, the leaves
are not as reflective as those of other species such as E. pa/meri and
E. farinosa (Harrington and Clark unpubl.). The trichomes also are
easily wettable, and are less reflective when wet. On foggy summer
days, common in the region, the wet leaves would absorb more light
for photosynthesis without adversely increasing water loss or leaf
temperature.

Relationships. The ultraviolet-reflecting rays (Clark and Sanders
1986) and the possession of a suite of benzopyran and benzofuran
secondary metabolites (Proksch and Clark 1987) are apomorphies

14 MADRONO [Vol. 35

Fic. 5. Scanning electron micrograph of the trichomes of the adaxial leaf surface
of Encelia densifolia (Clark 585).

linking E. densifolia to the clade containing the other Baja California
species EF. farinosa, E. californica, E. palmeri, E. ventorum, E. con-
spersa, and E. asperifolia. Its yellow discs are not found in other
Encelia species of the region, and, along with other features, suggest
that E. densifolia is basal in this clade (although its autapomorphies
argue against any consideration of it as “‘primitive’’).

Hybridization. In cultivation, EF. densifolia forms fertile hybrids
with every other Encelia species to which it is crossed; all other
Encelia species are equally interfertile (Kyhos et al. 1981). In the
natural environment, it is sympatric with E. farinosa and occurs
near E. palmeri. Although we saw no hybrids in the field, two col-
lections [Gentry 7559 (SD, UC), and 7587 (UC)] labeled “‘Encelia
viscainoensis”” appear intermediate between E. densifolia and one
of these species. Progeny testing of several dozen achenes collected
from an E. densifolia plant growing among E. farinosa yielded no
hybrid plants, but we crossed the species in cultivation and obtained
hybrids resembling “‘viscainoensis”’ in leaf morphology, so it is likely
that the “‘viscainoensis”’ collections represent field hybrids between
these species.

ACKNOWLEDGMENTS

This study was supported in part by grants from the Cal Poly Kellogg Unit Foun-
dation, the Cal Poly Institute for Cellular and Molecular Biology, and the Affirmative
Action Faculty Development Program, all to C.C.

1988] CLARK ET AL.: NEW ENCELIA Ne)

LITERATURE CITED

CLARK, C. and D. L. SANDERS. 1986. Floral ultraviolet in the Encelia alliance
(Asteraceae: Heliantheae). Madrono 33:130-135.

KyuHos, D. W., C. CLARK, and W. C. THOMPSON. 1981. The hybrid nature of Encelia
laciniata (Compositae: Heliantheae) and control of population composition by
post-dispersal selection. Syst. Bot. 6:399-411.

PROKSCH, P. and C. CLARK. 1987. Systematic implications of chromenes and ben-
zofurans from Encelia (Asteraceae). Phytochemistry 26:171-174.

WERK, K. S. and J. R. EHLERINGER. 1983. Photosynthesis by flowers in Encelia
farinosa and Encelia californica (Asteraceae). Oecologia 57:311-315.

(Received 17 Nov 1986; revision accepted 9 Nov 1987.)

ANNOUNCEMENT

OREGON ENDANGERED SPECIES PROGRAM

In September, 1987, Oregon Senate Bill 533, popularly known as the
Oregon Endangered Species Act, was passed into law. One of the pro-
visions of this legislation is the establishment of a threatened and en-
dangered species program under the direction of the state Department
of Agriculture. This new program was activated in February of this year,
with early goals being the development of rule-making procedures for
state listing of threatened and endangered plants, the initiation of a
review process to facilitate the ranking of candidate species, and the
establishment of research projects focusing on biological aspects of rarity
in the flora of the Pacific Northwest.

The Oregon Department of Agriculture is anxious to interact with
individuals and organizations with an interest in the sensitive plant
species of Oregon and adjacent states. Inquiries pertaining to the review
process, listing procedures, research, or questions concerning particular
taxa, should be addressed to: R. Meinke, Coordinator, Endangered Species
Program, Plant Division, ODA, 635 Capitol Street NE, Salem, Oregon
97310-0110.

aaa

NOTA SOBRE EL GENERO COMMELINA
(COMMELINACEAE) EN EL VALLE DE MEXICO CON
CAMBIOS EN LA NOMENCLATURA DE
ALGUNAS DE SUS ESPECIES

GRACIELA CALDERON DE RZEDOWSKI y JERZY RZEDOWSKI
Instituto de Ecologia, A. C., Centro Regional del Bajio,
Apartado postal 386, 61600 Patzcuaro, Michoacan, México

RESUMEN

Como resultado de la revision del genero Commelina, efectuada para el volumen
III de la Flora Fanerogamica del Valle de México, se reconoce la existencia de 7
especies, de las cuales 4 (C. coelestis Willd., C. dianthifolia DC., C. diffusa Burm. f.
y C. erecta L.) se aceptan en la circunscripcidn y nomenclatura usualmente empleadas.
En cuanto a las 3 restantes: el nombre C. orchioides Booth ex Lindl. substituye por
prioridad a C. alpestris Standl. & Steyerm.,; el de C. pallida Willd. es necesario restituir
para las plantas de la region generalmente identificadas como C. texcocana Matuda
y Phaeosphaerion leiocarpum (Benth.) Hassk.; el de C. tuberosa L., en cambio, se
aplica a las determinadas como C. coelestis var. bourgeaui Clarke.

ABSTRACT

In the treatment of the genus Commelina for the ‘““Flora Fanerogamica del Valle
de México” seven species are recognized, four of which (C. coelestis Willd., C. dian-
thifolia DC., C. diffusa Burm. f. and C. erecta L.) are accepted in their usual circum-
scription and nomenclature. As to the remaining three: the name C. orchioides Booth
ex Lindl., for reasons of priority, replaces C. a/pestris Standl. & Steyerm.; the name
C. pallida Willd. is rescued for local plants usually identified as C. texcocana Matuda
and Phaeosphaerion leiocarpum (Benth.) Hassk.; and the name C. tuberosa L. applies
to those determined as C. coelestis var. bourgeaui Clarke.

Antes de que llegara el turno de revisar las Commelinaceae para
el volumen III de la Flora Fanerogamica del Valle de México, se
podia prever que ésta no iba a ser una familia sencilla de estudiar.

A traves del tiempo y segun el criterio de diferentes autores, di-
versos miembros de las Comelinaceas han estado cambiando de
nombres y de situacion taxonomica, no solo a nivel de especie sino
también de género. Para el Valle de México resulto ser Commelina
que, conteniendo mayor numero de especies, presentO asimismo
mas problemas.

En general, las plantas correspondientes a Commelina son muy
atractivas, tanto por su follaje como especialmente por sus flores
vistosas, razon por la cual se hallan prodigamente representadas en
los herbarios; en total se examinaron mas de 500 especimenes. Ade-
mas se consultaron microfichas del herbario de Humboldt y Bon-
pland, del herbario Linneano y del herbario de Willdenow.

A pesar del considerable numero de eyjemplares conservados en

MAprRONO, Vol. 35, No. 1, pp. 16-22, 1988

1988] RZEDOWSKI AND RZEDOWSKI: MEXICAN COMMELINA 17

las instituciones, a menudo se nota inseguridad en las identificacio-
nes y una gran proporcion del material revisado cambio de nombre
al aplicar las conclusiones a las que se llego al finalizar este trabajo.

Uno de los problemas serios para el estudio de individuos her-
borizados de Commelina |o constituye la dificultad para observar
las caracteristicas de las flores, ya que éstas son fragiles y delicadas
y presentan el fenoOmeno de delicuescencia, conservandose mal en
los eyjemplares de herbario. Es por ello que ha resultado indispensable
hacer observaciones de cada una de las especies también en el campo.

De manera opuesta a lo que sucede en los herbarios, la bibliografia
sobre este grupo es relativamente escasa y se encuentra bastante
dispersa.

Hemsley (1885) y Reiche (1914), de acuerdo con Clarke (1881),
citan del Valle de Mexico las siguientes especies y variedades: Com-
melina coelestis Willd., C. dianthifolia DC., C. graminifolia HBK..,
C. graminifolia var. stricta (Desf.) Clarke, C. pallida Willd., C. qui-
tensis var. cardiosepala (Kunze) Clarke (var. “cardiophylla’’, segan
Reiche) y C. tuberosa L.; Reiche agrega asimismo C. scabra Benth.

La puesta al dia mas reciente para las Comelinaceas mexicanas
la constituyen los trabajos de Matuda (1956a, b), en los cuales se
intenta depurar algunos de los criterios de Clarke y donde se men-
cionan para nuestra region: C. alpestris Standl. & Steyerm., C. coe-
lestis Willd. (con C. pallida Willd. como sinonimo), C. coelestis var.
bourgeaui Clarke, C. dianthifolia DC., C. diffusa Burm. f., C. scabra
Benth. y C. texcocana Matuda, ademas de Phaeosphaerion leiocar-
pum (Benth.) Hassk. Sanchez (1969) sigue esta misma disposiciOn.

El grupo evidentemente requiere de una revision critica a nivel
continental y el presente estudio no puede resolver todos los pro-
blemas inherentes. Los resultados obtenidos, en ocasiones aun ten-
tativos, llevan a reconocer 7 especies para el Valle de México: C.
coelestis Willd., C. dianthifolia DC., C. diffusa Burm. f., C. erecta
L., C. orchioides Booth, C. pallida Willd. y C. tuberosa L.

1. COMMELINA COELESTIS Willd., Enum. Hort. Berol. 1:69. 1909.

Es una especie vistosa, mas bien alta y tosca, erecta, de hojas
largas y anchas, con la base envainadora; flores de 2 a 3 cm de ancho,
de color azul intenso. Comun en matorrales, pastizales o bosques
de encino y pino, en altitudes entre 2250 y 2750 m.

Coincidimos con la acepcion general de este nombre en la lite-
ratura y los herbarios; diferimos en considerar como sinonimo a C.
pallida Willd., que es especie distinta. Un gran numero de ejemplares
del Valle de México concuerda bien con la imagen del tipo, depo-
sitado en el herbario de Willdenow. Existen, sin embargo, individuos
con caracteristicas intermedias entre C. coelestis y C. orchioides y
también entre C. coelestis y C. tuberosa. A este respecto véase tam-
bien la discusion sobre la ultima especie.

18 MADRONO [Vol. 35

2. COMMELINA DIANTHIFOLIA DC. in Redoute, Lil. 7. Tab. 390. 1812.

Erecta o suberecta, delicada tanto en sus tallos delgados y hojas
angostas, como en sus espatas florales largamente acuminadas; flores
de aproximadamente 2 cm de ancho, de color azul intenso. Habita
en matorrales, pastizales, encinares y bosques de Abies; ampliamente
distribuida en altitudes entre 2250 y 3000 m.

Bastante bien definida en bibliografia y herbarios, aunque no fal-
tan individuos intermedios entre esta especie y C. tuberosa.

3. COMMELINA DIFFUSA Burm. f., Fl. Ind. 18, pl. 7, f. 2. 1768.
C. nudiflora sensu Burm. f., Fl. Ind. 17, non L.
C. longicaulis Jacq., Coll. Bot. 3:334. 1789.

Rastrera a ascendente, ramificada, tallos delgados, radicantes en
los nudos inferiores; flores de unos 2 cm de ancho, de color azul
intenso. Colectada principalmente en matorrales y pastizales; am-
pliamente distribuida entre 2300 y 2750 m.

Hemsley y Reiche no la citan, debido probablemente a que Clarke,
aun cuando hace mencion de “C. nudiflora L.’’, no la refiere espe-
cificamente para México. Por lo general bien determinada en los
herbarios, donde con frecuencia se le encuentra nominada con al-
guno de los sinonimos.

4. COMMELINA ERECTA L., Sp. Pl. 41. 1753.

Mas bien erguida, se distingue de las demas por tener las espatas
connadas en su extremo posterior y las flores con 2 pétalos de color
azul claro, de alrededor de | cm de largo y el tercero unas 10 veces
mas pequeno que los anteriores, de color amarillo con blanco. Planta
de afinidades termOfilas, escasisima (jausente en la actualidad?) en
el Valle de México.

No se ha encontrado citada con anterioridad para esta region, con
excepcion de Matuda, quien, en sus ““Commelinaceas mexicanas”’
(1956a), senala la existencia de C. erecta var. angustifolia (Michx.)
Fern., en base a la colecta de Purpus 14327 de ‘San Carlos, D.F.”’,
de septiembre de 1930; ésta no se menciona en las ““Commelinaceas
del Estado de México” (Matuda 1956b). Posiblemente se trata de
un error, pues la localidad no resulta familiar y casi no se conocen
colectas de Purpus del Distrito Federal.

Sin embargo, en los herbarios ENCB y MEXU se encuentran
ejemplares de Lyonnet 588, colectados en el Pedregal de San Angel
en agosto de 1930, pertenecientes a C. erecta. Ademas se han visto
los siguientes especimenes: 1. ““C. de Azompan, Tequezquinahuac,
Agosto 1°, 1954, en matorral alto bosque secundario a 2860 m de

1988] RZEDOWSKI AND RZEDOWSKI: MEXICAN COMMELINA 19

altitud”, Matuda 31196 (MEXU). Tal altitud resulta excesiva para
esta especie y la localidad dudosamente pertenece al Valle de México.
2. “Valle de México, Distrito Federal, VI 1946’, M. Sanchez R. s.n.
(ENCB). 3. “Al oeste de Los Remedios, cerca de Naucalpan, cerca
del rio, 15 VIII 1966”, M. Rodriguez s.n. (ENCB). Estas dos ultimas
colectas fueron hechas por alumnos y no es imposible que las lo-
calidades estén equivocadas.

5. COMMELINA ORCHIOIDES Booth ex Lindl., Bot. Reg. Misc. 53.
1838.

C. variabilis Schlecht., Ind. Sem. Hort. Hal. 7:13. 1838.

C. scapigera Kunth, Enum. Pl. 4:46. 1843.

C. alpestris Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Bot.
Ser. 23:213. 1947.

Planta mas bien baja, escaposa o subescaposa; flores de 2 a 3 cm
de ancho, de color azul intenso. Presente en pastizales y bosques de
encino o de coniferas de regiones montanosas, en altitudes entre
2800 y 3500 m.

Commelina alpestris es el nombre mas difundido en los herbarios,
pero como lo apunto Rohweder (1956), tal binomio no es el mas
antiguo para esta especie, por cierto bastante variable y comun en
el centro de México. C. orchioides tiene prioridad de varios meses
sobre C. variabilis, ambas descritas a base de materiales cultivados
procedentes de Real del Monte, Hidalgo. C. scapigera, aunque men-
cionada desde 1832, al parecer no fue formalmente publicada sino
hasta 1843. Clarke (1881), desde hace tiempo reconocio que los
ultimos tres binomios corresponden a un solo taxon, pero a su vez
los consider6o como sinonimos de C. e/liptica HBK., descrita a base
de eyjemplares de la costa de Venezuela. Aparentemente ni en el
herbario de Paris ni en el de Berlin existen materiales originales de
C. elliptica, pero su protologo senala tantas diferencias con respecto
a la especie mexicana de alta montana, que el nombre en cuestion
debe rechazarse para nuestra planta.

6. COMMELINA PALLIDA Willd., Hort. Berol. 2:87. 1816.
C. texcocana Matuda, An. Inst. Biol. Méx. 24:60. 1955.

Planta hasta de 2.5 m de alto, por lo comtn muy ramificada y
sarmentosa; flores de alrededor de 1.5 cm de ancho, de color-vio-
laceo claro. Habita en matorrales y a veces en bosque perturbado
de encino, principalmente del oeste y sur del Valle, en altitudes entre
2300 y 2600 m.

El tipo de C. texcocana coincide bastante bien con la fotografia
del material original de C. pallida y es idéntico a plantas reciente-

20 MADRONO [Vol. 35

mente colectadas entre San Juan del Rio y Querétaro, de donde
proviene también tal material. En los herbarios revisados, la mayor
parte de los ejemplares de C. pallida se encontraban identificados
como C. texcocana Matuda 0 equivocadamente como Phaeosphae-
rion leiocarpum (Benth.) Hassk., especie de clima caliente y frutos
carnosos.

En apariencia corresponde aqui igualmente lo referido por Clarke
(1881) y Hemsley (1885) como C. quitensis var. cardiosepala (Kunze)
Clarke y por Reiche (1914) como C. quitensis var. cardiophylla (sic),
ya que la descripcion original de C. cardiosepala Kunze concuerda
de manera aceptable con C. pallida.

7. COMMELINA TUBEROSA L., Sp. Pl. 61. 1753.

C. graminifolia HBK., Nov. Gen. Sp. Pl. 1:258. 1815.

C. graminifolia var. stricta (Desf.) Clarke in A. DC., Monogr. Phaner.
37152-18812

C. coelestis var. bourgeaui Clarke in A. DC., Monogr. Phaner. 3:
153. 1881.

Mas bien delicada, de tallo Unico o poco ramificado, delgado,
hojas angostas, espatas cortas, abruptamente agudas, flores de 2 a 3
cm de ancho, de color azul intenso. Comun, ampliamente distribuida
en matorrales, pastizales, bosques de encino o de coniferas, en al-
titudes entre 2300 y 3100 m.

La mayor parte de los eyjemplares de herbario se encontraron
identificados como C. coelestis var. bourgeaui; sin embargo, al com-
parar las fotografias de tipos de esta ultima y de C. tuberosa L., no
se hallaron mayores diferencias y mas que todo, el estudio de las
poblaciones en el campo indico que, al parecer, no existe en México
mas que una sola entidad biologica que corresponde a los tipos en
cuestion.

A la misma conclusion ha llegado Rohweder (1956), quien ubico
ademas a C. coelestis en la sinonimia de C. tuberosa.

Las observaciones en el Valle de México llevan a la conclusion
que C. tuberosa, C. coelestis, C. dianthifolia y C. orchioides estan
muy emparentadas entre si y al parecer pueden cruzarse con fre-
cuencia. Solo un estudio mas profundo y mas extenso podra deter-
minar si conviene mantenerlas como especies separadas 0 mas bien
considerarlas como variedades de C. tuberosa.

Algunos especimenes han sido erroneamente determinados como
C. scabra Benth. y de ahi tal vez derivan las citas de esta ultima
para el Valle de México (Reiche 1914, Matuda 1956b). Se trata, al
parecer, de una especie independiente, caracterizada por flores de
color rojizo, de cuya presencia en la region no se tiene certidumbre.

A continuaciOn se proporciona una clave para separar las siete
especies involucradas.

1988]

1.

RZEDOWSKI AND RZEDOWSKI: MEXICAN COMMELINA ZA

Espata connada en su extremo posterior, el cual es recto 0 casi
recto; pétalo inferior mucho mas pequeno (1.5 mm de largo por
menos de | mm de ancho) que los dos pétalos superiores; planta
escasa (probablemente ya inexistente) en el Valle de México, solo
conocida con seguridad del Pedregal de San Angel ...........

eo 8 © ee

IE TD BR dim Nel Oo, Os Lh BN Aes 4. CC. erecta

Espata libre en su extremo posterior, el cual por lo general es
redondeado; pétalo inferior semejante en forma y tamano a los
dos superiores.

2. Raices cilindricas, no tuberosas; plantas por lo general rastre-
ras a ascendentes o sarmentosas, rara vez erectas, profusa-
mente ramificadas al menos en la madurez, las ramificaciones
divergentes; pétalos por lo general de menos de | cm de largo
y 1 cm de ancho.

by,

Planta rastrera a ascendente, rara vez erecta, mas bien
delicada, con tallos por lo general de menos de 5 mm de
diametro (en la parte basal, en vivo); pétalos de color azul
intenso, el inferior cortamente unguiculado, con una una
de +1 mm de largo por +1 mm de ancho 3. C. diffusa
Planta a menudo sarmentosa, con tallos robustos, por lo
general de mas de 5 mm de diametro (en la parte basal,
en vivo); pétalos de color azul-violaceo claro, el inferior
subsésil, con una una de +0.5 mm de largo por 0.5 mm
CS eae tee en ee ee ee ee 6. C. pallida

2. Raices tuberosas, fusiformes; plantas erectas, de tallo unico o
poco ramificado, con las ramificaciones ascendentes; pétalos
por lo general de mas de | cm de largo y | cm de ancho.

4.

Plantas escaposas 0 subescaposas, por lo general de menos
de 30 cm de alto, con hojas principalmente basales; pe-
dunculo (uno o varios desde la base) largo, muy recto, con
una sola espata terminal; habita en altitudes de 2800 m o
TVASS ge ie nde ses Ga ee ae ee ee 5. C. orchioides

. Plantas con tallos bien definidos, simples o algo ramifi-

cados, por lo general de mas de 30 cm de alto, con varias
a numerosas espatas terminales o axilares; plantas de al-
titudes por lo general inferiores a 2800 m.
5. Hojas ampliamente lanceoladas, de mas de 2.5 cm de
ancho, con la base amplexicaule ...... 1. C. coelestis
5. Hojas estrechamente lanceoladas a lineares, de 2.5 cm
oO menos de ancho.
6. Espata floral corta, abruptamente aguda; hojas por
lo general lanceoladas, de 0.6 a 1 cm de ancho
neta tetrad atercat teeny Faber a Pees tes nel moi heat 7. C. tuberosa
6. Espata floral largamente acuminado-caudada; hojas
por lo general lineares, de 0.2 a 0.7 cm de ancho .
a ee ee era 2. C. dianthifolia

29 MADRONO [Vol. 35

AGRADECIMIENTOS

Este trabajo ha sido subvencionado por el Consejo Nacional de Ciencia y Tecnologia
y por el Centro de Investigacion y Desarrollo del Estado de Michoacan.

Se agradece a las autoridades de los herbarios CHAPA, ENCB y MEXU sus aten-
ciones en la consulta y préstamo de ejemplares. Al Dr. P. H. Raven, director del
Jardin Botanico de Missouri, por su ayuda en la obtencion de fotocopias de publi-
caciones que han sido de gran utilidad. A las personas responsables de las bibliotecas
del Departamento de Botanica de la Escuela Nacional de Ciencias Biologicas y del
Instituto de Biologia de la Universidad Nacional Autonoma de México y especial-
mente al Bibl. Armando Butanda, por las facilidades y apoyo ofrecidos para la consulta
de material bibliografico. A la Dra. A. Lourteig, del Museo Nacional de Historia
Natural de Paris, por la busqueda de materiales originales de C. elliptica. Al Sr. J.
Flanagan, bibliotecario de los Jardines Botanicos Reales de Kew, por su ayuda en la
determinacion de fechas de publicacion de C. orchioides y C. variabilis.

LITERATURA CITADA

CLARKE, C. B. 1881. Commelinaceae. Jn A. DC., Monogr. Phan. 3:1 13-324.

HEMSLEY, W. B. 1885. Botany. Jn F. D. Godman and O. Salvin, eds., Biologia
Centrali-Americana. R. H. Porter, London. Vol. 3:386-389.

MatTuDA, E. 1956a. Las Commelinaceas mexicanas. Anales Inst. Biol. Univ. Nac.
México 26:303-432.

1956b. Las Commelinaceas del Estado de México. Direccion de Recursos
Naturales del Gobierno del Estado de México. Toluca, México.

REICHE, C. 1914. La vegetacion en los alrededores de la capital de México. México,
D.F.

ROHWEDER, O. 1956. Die Farinosae in der Vegetation von El Salvador. Abh. Aus-
landsk., Reihe C, Naturwiss. 61.

SANCHEZ, O. 1969. La flora del Valle de México. Editorial Herrero, S.A., México,
D.F.

(Received 18 Feb 1987; revision accepted 30 Sep 1987.)

ANNOUNCEMENT

NEw PUBLICATIONS

Nose, W. J, AuTi, G. F. Otro, and I. M. BRopo. 1987. A second
checklist and bibliography of the lichens and allied fungi of British
Columbia. Syllogeus 61. ISSN 0704-576X. [Syllogeus is a publication
of the National Museum of Natural Sciences designed to permit rapid
dessemination of information pertaining to the disciplines and edu-
cational functions for which the Museum is responsible. Articles are
published in English, French, or in both languages, and issues appear
at irregular intervals. Individual copies of no. 61 and a list of other
publications from the Sy//ogeus series are available by mail from the
National Museum of Natural Sciences, Ottawa, Ontario, Canada KIA
OM8.]

RUSHFORTH, K. D. 1987. Conifers. Facts on File Publications, New
York. ISBN 0-8 160-1735-2, $24.95. [Topics covered include, biology
of conifers, conifers in the garden, dwarf conifers, propagation, plant-
ing and replanting, pests and diseases, and a gazetteer of conifers. The
gazetteer comprises over half of the book.]

A NEW SECTION OF CLITORIA SUBGENUS
NEUROCARPUM (LEGUMINOSAE)

PAUL R. FANTZ
Department of Horticultural Science,
North Carolina State University, Raleigh 27695-7609

ABSTRACT

Clitoria subgenus Neurocarpum sect. Mexicana, sect. nov. is described including
ser. nov. Mexicana, Tucumania, and Americana. Section Mexicana is contrasted with
sect. Tanystyloba and sect. Neurocarpum. Series Mexicana includes C. humilus Rose,
C. mexicana Link, C. monticola Brandegee, C. polystachya Benth., and C. triflora
Wats. Series Tucumania includes C. cordobensis Burk. Series Americana includes C.
fragrans Small and C. mariana L. A key to species of sect. Mexicana is provided.

RESUMEN

Se describe Clitoria subgénero Neurocarpum seccion Mexicana, secc. nov., e in-
cluye las ser. nov. Mexicana, Tucumania y Americana. La seccion Mexicana se con-
trasta con la seccion Tanystyloba y con la seccion Neurocarpum. La serie Mexicana
incluye, C. humilus Rose, C. mexicana Link, C. monticola Brandegee, C. polystachya
Benth. y C. triflora Wats. La serie Tucumania incluye C. cordobensis Burk. La serie
Americana incluye C. fragrans Small y C. mariana L. Hay una clave para identificar
las especies en la seccion Mexicana.

Clitoria (Leguminosae) comprises 60 species distributed primarily
within the pantropical-subtropical belt, of which 49 species are na-
tive to the neotropics. The habit of species of Clitoria range from
trees, shrubs, lianas, and subshrubs or perennial herbs with erect or
twining, annual, aerial stems from an underground xylopodium. It
is characterized by its showy, resupinate, papilionaceous flowers,
infundibular calyx with persistent bracteoles, and stalked ovaries
with a geniculate, bearded style.

Clitoria was last revised by Bentham in 1858. Fantz (1979b) rec-
ognized three subgenera circumscribed by fruits and seed morphol-
ogy, supported by leaf, calyx, androecial, and gynoecial features,
germination characteristics, and the presence or absence of cleis-
togamy. Proposed revisions are based on examination of nine thou-
sand specimens, collected worldwide, that included types.

Fantz (1979a) circumscribed Clitoria subg. Neurocarpum by its
turgid fruit with convex values (ecostate or bearing a prominent
medial nerve), thickened viscid seeds, a subcartilaginous 10-nerved
calyx, hypogeal germination, and the presence of cleistogamous flow-
ers in half of its 24 species. Species of subg. Neurocarpum have been
segregated traditionally by growth form and vegetative characters,

MADRONO, Vol. 35, No. 1, pp. 23-31, 1988

[Vol. 35

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1988] FANTZ: NEW SECTION OF CLITORIA 25

Fic. 1. Clitoria laurifolia Poir. (a) aerial stem, <0.5; (b) flower, x 0.5; (c) calyx,
x 0.5; (d) gynoecium, x<0.5; (e) legumes, x 0.5; (f) three views of seed, x 0.5. (Sal/tz-
mann s.n., G, a, Smith 2050, GH, b-f.)

e.g., leaflet number, petiole length, and leaflet shape and apices.
Fruits and floral characters have been used rarely in floristic treat-
ments or Bentham’s (1858) treatise.

There are three sections within subg. Neurocarpum as distin-
guished by a combination of morphological characters and geograph-
ical distribution (Table 1). Section Neurocarpum (lectotype = C.
laurifolia Poir., Fig. 1) isa South American group of 10 species with

26 MADRONO [Vol. 35

Fic. 2. Clitoria macrophylla Wall. ex Benth. (a) portion of stem, x 0.5; (b) flower,
x 1; (c) calyx, x1; (d) gynoecium, x1; (e) legumes, x1. (Wallich 5345, K, a-e.)

three that extend into Central America and the West Indies. Mem-
bers are characterized by fruits that are turgid, not depressed between
the seeds, and often bearing a longitudinal, medial nerve (=costate
fruit). Fruits in some species are ecostate or the costa is imperfectly
formed. Seeds typically are slightly longer than they are wide (seed
length = axis parallel with funiculus). Chasmogamous flowers typ-

1988] FANTZ: NEW SECTION OF CLITORIA Zh

Fic. 3. Clitoria mexicana Link. (a) portion of stem, x 0.5; (b) flower, x 1; (c) calyx,
x 1; (d) gynoecium, x 1; (e) legumes, x 1; (f) three views of seed, x 1. (Breedlove 12025,

F, a; Hinton 11668, NY, b; Molina 18472, NY, c, d; Tucker 784, UC, e, f; Hernandez
1287, F, g.)

ically are larger, usually pigmented in shades of blue to purple, or
occasionally white. Cleistogamy is present in most species.

Section Tanystyloba (type = C. macrophylla Wall. ex Benth., Fig.
2) includes five species of Southeast Asia and one endemic in Aus-
tralia (Fantz, 1979a). Species are unique because they (1) lack cleis-

28 MADRONO [Vol. 35

togamy, (2) bear a calyx with elongated lobes longer than the tube,
and (3) possess a dense indumentum of uncinate hairs on the outer
calyx surface. Short-petiolate to subsessile leaves and occurrence of
unifoliate leaves is similar to some species of sect. Neurocarpum.

A third group includes seven species from North and Central
America and one species native to northern Argentina. These have
turgid, ecostate fruits that are depressed between the seeds. Seeds
are slightly wider than long. Plants are similar to those of species in
sect. Tanystyloba, but differ by the presence of cleistogamous flowers,
narrower fruits, and elongated inflorescences. These eight species
are placed in sect. Mexicana (type = C. mexicana Link, Fig. 3), a
new section of subg. Neurocarpum.

TAXONOMIC TREATMENT

Sect. Mexicana includes three distinct groups distinguished by
morphological differences in fruit and chasmogamous and cleistog-
amous flowers, supported by geographical distribution. These are
treated as Series as described below.

Key to Taxa of sect. Mexicana

Floral characteristics, unless otherwise noted, refer to chasmog-
amous flowers.

1. Gynophore 4-8 mm long, slightly shorter than the ovary; fruits
conspicuously depressed between the seeds; staminal tube of
cleistogamous flowers nearly lacking, ca. 0.1 mm long, filaments
nearly free; flowers blue to lilac to pale purple, (3.5—)4—6 cm long
ee ee ee ee ete eet ae ee Series Americana
2. Calyx tube 7-10 mm long; leaflets narrow, 0.5—1 cm wide,

primary nerves of 6-8 pairs; stipules 2-4 mm long, stipels 1-
3 mm long; stipe 15-21 mm long; cleistogamous flowers with
bracteoles 2-3 mm long, calyx tube 3—4 mm long, stipe 9-14
1046 OG (0) 8 ae ee ete ey Me Se EN ree RRS. ie eet C. fragrans

2. Calyx tube 10-14 mm long; leaflets broad, 1—4(—6.5) cm wide,
primary nerves of 7-12 pairs; stipules 5-10 mm long, stipels
3-8 mm long; stipe 5—17 mm long; cleistogamous flowers with
bracteoles 3—5 mm long, calyx tube 4-5 mm long, stipe 5-10
PUTO ec een eae atien, eae age C. mariana

1. Gynophore 3-4 mm long, much shorter than the ovary; fruits
weakly depressed between seeds; staminal tube of cleistogamous
flowers elongate, 3—5 mm long; flowers white or pale yellow,
occasionally purple, 2.5—4 cm long.

3. Stipe 8-16 mm long, exserted beyond calyx tube; carina with
blade 10-11 mm long, claw 18-22 mm; style 8-10 mm long;
cleistogamous flowers with staminal tube 3—4 mm long. One
SPCCICS: sans en yer test Series Tucumania (C. cordobensis)

1988] FANTZ: NEW SECTION OF CLITORIA 20

3. Stipe 4-8 mm long, enclosed within calyx; carina with blade
5-9 mm long, claw 12-17 mm long; style 10-17 mm long;
cleistogamous flowers with staminal tube 4-5 mm long ....
Ee el re nL EN Ma 8 ah een GLP Series Mexicana
4. Flowers 3—4 cm long; calyx lobes 6-8 mm long; inflores-
cence racemose, 2—4(—6)-flowered; perennial herbs with ae-

rial stems erect to twining.

5. Vine; calyx tube 9-12 mm long, purplish; bracteoles 5—
9 mm; inflorescence 2-11 cm long; leaflets ovate to
lanceolate; stipules 6-9 mm long ...... C. mexicana

5. Erect herb; calyx tube 7-9 mm long, greenish; bracteoles
3-4 mm long; inflorescence 0.2—0.3 cm long; leaflets
oblong; stipules 4-6 mm long ........... C. humilus

4. Flowers 2.5-—3 cm long; calyx lobes 2—5 mm long; inflo-
rescence paniculate (occasionally racemose), 4—8-flowered
or more; shrubs to subshrubs.

6. Flowers lilac to purple; calyx lobes 4-5 mm long; stem
pubescence spreading; bracteoles 7-9 mm long; cleis-
togamous flowers not observed ........... C. triflora

6. Flowers white; calyx lobes 2-4 mm long; stem pubes-
cence ascending, subappressed; bracteoles 4—7(—9 in va-
riety C. polystachya) mm long; cleistogamous flowers
present.

7. Inflorescence racemose, few flowered; carina with
blade 5—7 mm long; wings with blade 8—1 1 mm long,
claw 10-14 mm long; style 10-11 mm long; ovary
moderately uncinate pubescent; leaflets narrower,
1.5-3.5 cm wide, primary nerves of 3-6 pairs; petiole
3-6 cm long, petiolule 2-3 mm long ............

7. Inflorescence paniculate, many-flowered; carina with
blade 7-8 mm long; wings with blade 12-14 mm
long, claw 8-11 mm long; leaflets broader, 2-6 cm
wide, primary nerves of 8—12 pairs; petiole 4—10 cm
long, petiolule 3-5 mm long...... C. polystachya

Clitoria L. subgenus Neurocarpum (Desv.) Baker sect. Mexicana
Fantz, sect. nov.

Sectione nova Clitoria subgenere Neurocarpum cum Tanystyloba
affini optimo distinguitur a inflorescentia solitaro et elongato, calyce
lobis breviores longi tubo et flores cleistogamis praesentia.

Leaves 3-foliolate, petiolate. Inflorescences axillary, solitary, pa-
niculate or racemose, (1—)2-several flowered; peduncles 2-13 cm
long. Chasmogamous flowers showy, white or blue to purple, 2.5—
4(-6) cm long. Calyx tube 7—12(—14) mm long; lobes shorter than

30 MADRONO [Vol. 35

the tube; sparingly to moderately pubescent with uncinate trichomes
and subappressed macroscopic trichomes. Staminal tube 1|.3-—2.2(—3)
cm long. Style 1-2 cm long. Cleistogamous flowers inconspicuous,
0.4-0.9 cm long. Calyx tube 3-6 mm long; lobes 1.5-—3 mm long.
Staminal tube elongate, 3-5 mm long, or ca. 0.1 mm long. Legume
ecostate, valves weakly to strongly depressed between seeds.

Sp. typica: Clitoria mexicana Link.

Members of sect. Mexicana typically occur in dry sandy soils,
often in drier woodlands of lowlands and mountain slopes to about
2700 m. Three distinct series are observed.

1. Series Mexicana

Shrubs, subshrubs to suffrutescent herbs. Chasmogamous flowers
2.5-4 cm long, white fading pale yellow, or occasionally purplish;
gynophore 3—4 mm long, much shorter than the ovary; style 10-17
mm long. Cleistogamous flowers with staminal tube 3—S5 mm long.
Legume weakly depressed between the seeds, short-stipitate; stipe
4—8 mm long; seeds slightly wider than long. Central America.

Series Mexicana includes the following species: C. humilus Rose,
Mexico (holotype: Rose 2251, US!), C. mexicana Link, Mexico to
Nicaragua (neotype: Hinton 11668, NY!), C. monticola Brand., Mex-
ico (holotype: Brandegee s.n., UC 83907!), C. polystachya Benth..,
Mexico to Panama (lectotype: Hartweg 454, K!), and C. triflora
Wats., Mexico (lectotype: Palmer 159, GH!). These species are pos-
tulated to have originated in Mexico with two species migrating into
Central America. Wiggins (1980) omitted C. monticola from the
flora of Baja California.

2. Series Tucumania Fantz, ser. nov.

Serie nova Mexicana optimo distinguitur a stipite elongato, carino
longiores, stylis breviores, et tubo staminali e flores cleistogamis
breviores. Tucumania.

Perennial suffrutescent herbs. Chasmogamous flowers 2.5—3 cm
long, white; gynophore 3—4 mm, much shorter than the ovary; style
5—6 mm long. Cleistogamous flowers with staminal tube 3-4 mm
long. Legume long-stipitate, valves weakly depressed between the
seeds; stipe 10-16 mm long; seeds slightly wider than long. Argen-
tina.

Sp. typica: Clitoria cordobensis Burkart.

Clitoria cordobensis, endemic to northern Argentina (lectotype:
Nicora 1774, SI'), is the only member of this series.

3. Series Americana Fantz, ser. nov.

Serie nova Mexicana optimo distinguitur a gynophoro elongato
fere subequalis ovario, flores statura ampliore, et tubo staminali e

1988] FANTZ: NEW SECTION OF CLITORIA 31

flores cleistogamis brevissimo. Americana cum orientalis disjuncto
varieto.

Perennial suffrutescent herbs. Chasmogamous flowers 3.5—6 cm,
bluish to lilac to pale purple; gynophore 4—8 mm long, slightly shorter
than the ovary; style 13-20 mm long. Cleistogamous flowers with
the staminal tube ca. 0.1 mm long, the stamens appearing to be free.
Legumes conspicuously depressed between the seeds, long-stipitate;
stipe 10—21 mm long; seeds slightly longer than wide. United States
(one variety in se. Asia).

Sp. typica: Clitoria mariana L.

Series Mariana includes two species, C. mariana L., from the U.S.
with a variety in Southeast Asia (type: Hb. Petiver, BM—Hb. Sloane)
and C. fragrans, an endemic to southern Florida (lectotype: Small
and Wherry 12626, NY’).

ACKNOWLEDGMENTS

I thank Tom Davis for the Latin diagnoses and the reviewers for their critical
comments and excellent suggestions that improved the quality of this manuscript. I
am especially grateful to the curators of the following institutions for the loan of
material: A, BA, BM, BR, CAL, CGE, CM, DUKE, E, F, FLAS, G, GH, HAL, K,
LA, M, MG, MICH, MO, MPU, NCSC, NY, P, PENN, PH, PR, RB, S, SI, U, UC,
UMO, UNC, US, USCH, VEN, VSC, W, WIS. Paper No. 10564 of the Journal Series
of the North Carolina Agricultural Research Service, Raleigh, NC 27695-7601.

LITERATURE CITED

BENTHAM, G. 1858. Synopsis of the genus C/itoria. J. Proc. Linn. Soc., Bot. 2:
33-44.

FANTz, P.R. 1979a. Anew section of Clitoria subgenus Neurocarpum (Leguminosae)
and a new species endemic to Thailand. Brittonia 31:115-118.

1979b. Taxonomic notes and new sections of Clitoria subgenus Bractearia
(Leguminosae). Sida 8:90-94.

WiaaIns, I. L. 1980. Flora of Baja California. Stanford Univ. Press, Stanford, CA.

(Received 11 Jul 1986; revision accepted | Oct 1987.)

MORPHOLOGICAL AND ECOLOGICAL VARIATION
ACROSS A HYBRID ZONE BETWEEN
ERYTHRONIUM OREGONUM AND
E. REVOLUTUM (LILIACEAE)

GERALDINE A. ALLEN and JOSEPH A. ANTOS
Department of Biology, University of Victoria,
Victoria, British Columbia V8W 2Y2, Canada

ABSTRACT

Erythronium oregonum and E. revolutum overlap only slightly in geographic dis-
tribution, but can hybridize where they co-occur. Morphological traits of Erythronium
and cover of associated species were assessed in |m? plots in one hybrid and several
pure populations of these two species on Vancouver Island. The species differ in size
of flowers, relative size of scape and leaves, and several flower color characters. They
also differ ecologically, E. oregonum occupying drier microsites than E. revolutum.
Hybrids between them exhibited various recombinations of the parental morpho-
logical traits. Morphological variation in Erythronium at the hybrid locality was
correlated with the ecological characteristics of the plots as indicated by a detrended
correspondence analysis of associated plant species (r = 0.64). Erythronium oregonum
and FE. revolutum apparently are maintained as separate taxa by their ecological
differences.

The genus Erythronium reaches its greatest diversity in western
North America, where approximately 14 species occur (Applegate
1935, Hitchcock et al. 1969, Hammond and Chambers 1985). Al-
though some of these have been studied ecologically (Caldwell 1969,
Antos and Zobel 1984, Thomson and Stratton 1985), little is known
about evolutionary relationships within the genus, or the genetic and
ecological basis for species differences.

Erythronium oregonum Applegate and E. revolutum Smith are
lowland species of the Pacific Northwest. They are morphologically
and ecologically similar, and Applegate (1935) has reported hybrids
between them. Both species have mottled leaves, small saccate ap-
pendages on the inner tepals, dilated anther filaments, and a three-
lobed stigma. Erythronium revolutum has yellow anthers and rose
pink tepals with yellow banding at the base. Erythronium oregonum
is more variable; in many populations (including those in British
Columbia) the anthers are yellow and the tepals are white with yellow
and red banding at the base, but southern populations commonly
have paler anthers and cream-colored tepals lacking the red mark-
ings. In this paper we describe an instance of hybridization between
E. oregonum and E. revolutum in British Columbia, and present
evidence concerning morphological and ecological differences be-
tween the two species.

MADRONO, Vol. 35, No. 1, pp. 32-38, 1988

1988] ALLEN AND ANTOS: VARIATION IN ERYTHRONIUM 33

METHODS

Study plots. Hybrids occurred with E. oregonum and E. revolutum
at Skutz Creek (48°47'N, 123°57’'W), a small tributary of the Co-
wichan River located 9.2 km east of Cowichan Lake on southern
Vancouver Island, British Columbia. We established 29 | m? plots
at this site in areas of high Erythronium density. The plots were
located to encompass the full range of variation in morphology and
habitat of Erythronium at the site. For comparison, we located eight
additional plots in pure populations of FE. oregonum, of which five
were along the Cowichan River (1.0 km downriver from Skutz Creek,
48°47'N, 123°56’W) and three were near Victoria (at Beaver Lake,
48°31'N, 123°23’W, and on the University of Victoria campus,
48°28'N, 123°19'W). We also located five plots in pure populations
of E. revolutum along Sutton Creek (48°49'N, 124°13'’W), 2.0 km
west of Honeymoon Bay on Cowichan Lake.

Morphological characteristics. We measured eight morphological
characters of taxonomic importance (tepal color, style color, stamen
filament color, intensity of red banding on tepals, tepal length, length
of dehisced anthers, scape height, and leaf length) on 25 plants in
each of the 42 plots. Only characters that could be assessed non-
destructively were used. Color intensities (hue and lightness) were
determined using a Munsell color chart (Munsell Color, Baitimore,
MD). We also calculated a derived character, scape-leaf ratio (=scape
length/leaf length).

We used pure populations of &. oregonum and E. revolutum to
establish mean values of the diagnostic characters of each species.
The Cowichan River and Victoria populations of E. oregonum were
not significantly different (t-test for each character, p > 0.05 in all
cases), so the data from these were combined. We constructed a
hybrid index using six characters (tepal color, style color, filament
color, red banding, tepal length, and scape-leaf ratio) that differed
significantly between the two species, but were not highly correlated
with other characters. For all plants examined at the Skutz Creek
hybrid locality, the values of each character were scaled between the
mean values for FE. revolutum (set at zero) and for E. oregonum (set
at 10). Values outside the range of zero to 10 were set to zero or 10
as appropriate. A weighting factor inversely related to the amount
of overlap between the pure populations was then applied to each
character (weights were 1.000 for tepal color, 0.984 for style color,
0.887 for filament color, 0.930 for red markings, 0.720 for tepal
length, and 0.645 for scape-leaf ratio). The hybrid index was cal-
culated as the sum of these weighted values for the individual char-
acters, and this sum was then scaled between zero and 60. We cal-
culated a mean hybrid index for each plot in order to determine

34 MADRONO [Vol. 35

TABLE |. MEAN VALUES OF CHARACTERS USED IN STUDY OF Erythronium oregonum
AND E. revolutum. Color traits are expressed as values on an integer scale; tepal, style,
and filament color vary from 0 (white) to 7 (deep violet-pink), and red banding from
0 (absent) to 5 (broad deep-red bands). All characters differed significantly between
the two species (t-test; p values given below).

E. oregonum E. revolutum
(n = 200) (n = 125)

Character XK s.d. x sd. p
Tepal color 0.0 0.1 4.6 0.5 0.0001
Style color 0.0 0.0 3.4 lS 0.0001
Filament color 0.2 0.9 3.7 1.8 0.0001
Red banding De }.2 0.0 0.0 0.0001
Anther length (mm) 6.6 1.1 6.2 0.9 0.0003
Petal length (mm) 44.7 5.0 37.0 2.9 0.0001
Leaf length (cm) 1639 4.3 16.1 23 0.0253
Scape height (cm) 28.0 il 21.4 2.9 0.0001
Scape-leaf ratio 17 Oe ie OF 0.0001

whether plants in the plot were predominantly “revolutum”, “‘ore-
gonum” or “‘hybrid”’ in character.

Ecological characteristics. For all 1 m? plots except those at Vic-
toria, we recorded the percent cover of each vascular plant and moss
species present. All cover values were subjected to log transformation
before analysis. We carried out detrended correspondence analyses
of these data using the program DECORANA (Hill 1979) in order
to assess ecological resemblance among the plots. The analyses were
performed with all plots and with the Skutz Creek plots only, with
and without tree species included. We then examined the relation-
ship between Erythronium morphology, as indicated by the hybrid
index, and ecological characteristics, as indicated by plot ordination
scores derived from the detrended correspondence analysis.

RESULTS

Morphological characteristics. Populations of F. oregonum and E.
revolutum differed in a number of the morphological characters ex-
amined (Table |). The most striking differences between the two
species are in tepal color and markings. Other differences, which
were observed during this study but not measured, included (1)
length of stigma lobes (shorter in EF. revolutum); (2) curvature of
tepals (much reflexed in FE. revolutum, generally less so in E. ore-
gonum), (3) anther position (connivent around the style in E£. re-
volutum, spreading in E. oregonum), and (4) leaf position (generally
more erect in E. revolutum).

We examined correlations among the characters in Table 1, first
for all of the specimens measured at the Skutz Creek locality, and

1988] ALLEN AND ANTOS: VARIATION IN ERYTHRONIUM 5)

TABLE 2. PERCENT COVER AND (IN PARENTHESES) PERCENT OCCURRENCE OF
VASCULAR PLANT SPECIES IN PLOTS GROUPED BY THE Erythronium HYBRID INDEX.
Included are all species with >5% cover in at least one group. T = <0.5% cover;
nomenclature follows Hitchcock and Cronquist (1973).

Hybrid Index

=10 10-50 50
Species (n = 6) (n = 16) (n = 7)
Marees
Alnus rubra 78 (100) 50 (94) 16 (57)
Abies grandis 2 (17) 8 (50) 0 (0)
Acer macrophyllum 11 (100) 27 (100) 29 (57)
Pseudotsuga menziesii LOL?) 1 (19) 42 (86)
Shrubs
Rubus spectabilis 32 (83) 22 (81) 1 (14)
Oemleria cerasiformis 8 (67) 9 (69) 11 (29)
Symphoricarpos albus 0 (0) 5 (31) 7 (43)
Holodiscus discolor T (17) T (6) 37 (57)
Herbs
Trautvetteria caroliniensis 32 (83) 5 (38) T (14)
Maianthemum dilatatum 13 (33) 10 (38) 1 (14)
Dicentra formosa 7 (33) 6 (50) 1 (14)
Tolmiea menziesii 2 (50) 13 (56) T (14)
Heracleum lanatum 4 (50) 9 (31) 6 (29)
Viola glabella pS Ge) 6 (50) 8 (43)
Cardamine pulcherrima Belay 4 (38) 16 (86)
Perideridia gairdneri 0 (0) 0 (0) 9 (43)

secondly for that subset of specimens with a hybrid index between
20 and 40. In the first comparison some high correlations occurred
between characters found together in the same species, particularly
color characters. The highest correlation coefficient (0.78) was be-
tween tepal color and filament color. In the second comparison,
which was restricted to clearly hybrid specimens, no correlation
coefhcient higher than 0.51 was obtained, and most were much
lower.

Ecological characteristics. Results of the detrended correspon-
dence analyses indicate that E. oregonum and E. revolutum differ
in their ecological characteristics. The ordinations were all similar
and the first axis in each case corresponded to a moisture gradient.
At the hybrid locality typical E. oregonum was usually found on
well-drained microsites underlain with shale, whereas typical EF.
revolutum was usually found on wetter microsites with black, humus-
rich soils. Sample plots grouped on the basis of mean hybrid index
into “oregonum’’, “‘revolutum” or “hybrid” differed substantially in
species composition (Table 2). Erythronium oregonum typically grew
in sites with a Douglas-fir canopy and sparse herbaceous understory;

36 MADRONO [Vol. 35

60

a i ret
3 aad
oe te | é ;
50 *8 H
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é *3* @ ¢ | Se
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$s 8 ° oe ®@ e @
40 [: "e 4, : . ;
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30 : = :
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o: re 1 . §
o8 83 £. : : 3
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10 eo? Z . e
8 r)

0 100 200 300 400

DCA First Axis

Fic. 1. Hybrid index of individual Erythronium in relation to their position on
the first axis of a detrended correspondence analysis (DCA) of the 29 plots at Skutz
Creek. The analysis was based on all associated species. Low scores on the DCA axis
correspond to wetter sites and high scores to drier sites; typical E. revolutum has a
hybrid index of zero, and typical E. oregonum a hybrid index of 60.

in contrast, EF. revolutum was typically associated with luxuriant
herb layers under a red alder canopy.

To examine relationships between morphological and ecological
characteristics of E. oregonum, E. revolutum and their hybrids, we
plotted the hybrid indices of all plants from the Skutz Creek locality
against plot scores from the first axis of the detrended correspon-
dence analysis (Fig. 1). The first DCA axis accounted for 57% of the
variation in the species composition of the sample plots, and cor-
responded roughly to a moisture gradient with low scores indicating
wetter sites and high scores indicating dry sites. Plots with low scores
on the first DCA axis contained mainly E. revolutum, and those
with high scores on this axis contained mainly FE. oregonum (Fig.
1). The large amount of scatter in the central region of the figure
indicates that FE. oregonum and E. revolutum overlapped slightly in

1988] ALLEN AND ANTOS: VARIATION IN ERYTHRONIUM 37

their ecological tolerances, and also that the hybrid plants at Skutz
Creek exhibited many combinations of morphological and ecological
characteristics. Although the morphology of Erythronium plants var-
ied greatly in plots located in intermediate habitat, 41% of the vari-
ation in mean hybrid index was related to position on the first DCA
axls.

DISCUSSION

The western North American species of Erythronium are largely
allopatric in their distribution, and little quantitative information
exists concerning the degree of interfertility and naturally occurring
intergradation among species. Although only a few cases of hybrid-
ization between E. oregonum and E. revolutum have been reported
(Applegate 1935), extensive hybridization can occur under appro-
priate conditions. The variety and abundance of intermediate forms
found at the Skutz Falls study site suggest that many hybrid gen-
erations are present and that there is little or no post-zygotic repro-
ductive isolation between the species, at least at this locality. Ap-
plegate (1935) considered these two species to be related closely on
morphological grounds. Nevertheless, E. oregonum and E. revolu-
tum are consistently different in a number of morphological char-
acters as well as in their ecological requirements.

The evidence presented in this paper suggests that E. oregonum
and £. revolutum maintain their distinct taxonomic characteristics
largely through ecological separation. On Vancouver Island, E. re-
volutum typically occurs under deciduous trees on relatively wet
sites such as stream terraces. It can be abundant, locally dominating
the herb layer, but occurs on a restricted range of sites. Erythronium
oregonum is most common on dry open sites such as forest openings
and rock outcrops, although it occasionally occurs in wetter loca-
tions; it appears to occupy a wider variety of sites than EF. revolutum.
The overall geographic distributions of the two species reflect these
ecological differences. Erythronium revolutum occurs on Vancouver
Island and the adjacent mainland to the north, then south through
the Olympic Peninsula of Washington and coastal regions of Oregon
to northern California (Bierly and Stockhouse 1982, James 1983).
Its occurrence is somewhat sporadic, probably reflecting relatively
specific habitat requirements and the patchy distribution of suitable
sites. Erythronium oregonum extends from the east coast of Van-
couver Island and adjacent mainland British Columbia, southward
through the Puget Trough of Washington and the Willamette Valley
of Oregon (extending into the Coast Range and to the west slope of
the Cascades); it reaches its southern limit in the vicinity of the
Illinois River of southern Oregon (Applegate 1935).

Although the two species exhibit some overlap in range, they

38 MADRONO [Vol. 35

seldom grow in close proximity because of their differences in habitat
preference. For successful hybridization to occur, pollen or seeds
must be transferred between parental populations. In Erythronium,
this seems likely only if the two taxa are established immediately
adjacent to one another. Such a situation exists at Skutz Creek, where
dry slopes with fairly pure EF. oregonum populations occur within
100 m of alluvial flats with populations of E. revolutum. Given the
apparent absence of intrinsic genetic barriers between these two
species, the close juxtaposition of divergent habitats that can support
both taxa is probably the major reason for hybridization.

ACKNOWLEDGMENT

This work was supported by Natural Science and Engineering Council of Canada
Grant A-8087.

LITERATURE CITED

AnTos, J. A. and D. B. ZoBEL. 1984. Ecological implications of belowground mor-
phology of nine coniferous forest herbs. Bot. Gaz. (Crawfordsville) 145:508-517.

APPLEGATE, E. I. 1935. The genus Erythronium: a taxonomic and distributional
study of the western North American species. Madrono 3:58-113.

BiERLY, K. F. and R. E. StockHousg, II. 1982. Coast fawn lily (Erythronium re-
volutum) sensitive species conservation status report. U.S. Forest Service, Suislaw
National Forest, Corvallis, OR.

CALDWELL, M. L. H. 1969. Erythronium: comparative phenology of alpine and
deciduous forest species in relation to environment. Amer. Midl. Naturalist 82:
543-558.

HAMMOND, P. C. and K. L. CHAMBERS. 1985. A new species of Erythronium (Lil-
iaceae) from the Coast Range of Oregon. Madrono 32:49-56.

Hitt, M. O. 1979. DECORANA—a FORTRAN program for detrended corre-
spondence analysis and reciprocal averaging. Cornell University, Ithaca, NY.

Hitcucock, C. L. and A. CRONQuIST. 1973. Flora of the Pacific Northwest. Univ.
Washington Press, Seattle.

, M. Ownsey, and J. W. THompson. 1969. Vascular plants of the
Pacific Northwest. Part 1: Vascular cryptograms, gymnosperms and monocot-
yledons. Univ. Washington Press, Seattle.

JAMES, J. C. 1983. Species management guide for Erythronium revolutum. U.S.
Forest Service, Olympic National Forest (Quinault Ranger District), Quinault,
WA.

THomson, J. D. and D. A. STRATTON. 1985. Floral morphology and cross-polli-
nation in Erythronium grandiflorum (Liliaceae). Amer. J. Bot. 72:433-437.

(Received 11 Dec 1986; revision accepted 6 Oct 1987.)

CROSSABILITY AND RELATIONSHIPS OF
PINUS MURICATA (PINACEAE)

CONSTANCE I. MILLAR!
Wildland Resources Center, University of California,
Berkeley 94720

WILLIAM B. CRITCHFIELD
Pacific Southwest Forest and Range Experiment Station,
USDA Forest Service, Berkeley, CA 94701

ABSTRACT

Crossing relationships were studied within and among the variable populations of
Pinus muricata to test hypotheses about crossing barriers among certain populations.
Crossability was assessed at the level of viable seed production following planned
crosses. Populations north of Sea Ranch, Sonoma Co., California, crossed freely with
parapatric but genetically distinct populations in central Sonoma Co., although some
reduction in seed-set occurred in the F, and backcrosses to F,. The distinctness of
these adjacent populations is most likely not maintained by post-pollination crossing
barriers. Crossability of disjunct P. muricata populations generally decreased with
distance between populations. Populations north of Sea Ranch crossed freely with
the Pt. Reyes population in Marin Co., less readily with the Monterey population,
and not at all with the Purisima (southern California) or Baja California populations.
Mainland and island P. muricata populations south of Monterey were highly inter-
fertile. Test crosses were also attempted between P. muricata and the island popu-
lations of P. radiata, which have been considered closely related to southern P.
muricata populations. Pinus muricata from Baja California did not cross, however,
with either Guadalupe Island pine (P. radiata var. binata) or Cedros Island pine (P.
radiata var. cedrosensis). Together with results from other crossing studies in the
Californian closed-cone pines, the patterns of crossability indicate three crossing units
in P. muricata: 1) northern P. muricata populations from Marin Co. northward,
which are reproductively isolated from, 2) southern P. muricata populations including
mainland and Channel Islands populations from Purisima southward, and 3) Mon-
terey P. muricata, which is intermediate between the first two units.

Crossing patterns within the three Californian species of closed-
cone pines (subsect. Oocarpae, Critchfield and Little 1966) are un-
usual for Pinus. In experimental pollinations, Pinus radiata D. Don
and P. attenuata Lemmon hybridize more readily than most other
combinations in the genus, whereas crosses between certain northern
and southern populations of P. muricata D. Don do not produce
viable seeds (Critchfield 1967). This is the only known instance in
Pinus of complete infertility between populations within a species.

Pinus muricata 1s unique among the California closed-cone pines
because of the distribution of genetic variation within and among

' Present address: Institute of Forest Genetics, Pacific Southwest Forest and Range
Experiment Station, U.S.D.A. Forest Service, Box 245, Berkeley, CA 94701.

MApDRONO, Vol. 35, No. 1, pp. 39-53, 1988

40 MADRONO [Vol. 35

populations. The frequencies of several morphological and bio-
chemical traits in northern populations of P. muricata change abruptly
within continuous stands. Cone morphology in southern populations
also differs markedly within and among nearby disjunct stands. Vari-
ation is especially complex in the southern and island populations
where several traits intergrade between P. muricata and P. radiata.

This variation has led taxonomists since the early 1800’s to apply
many species and varietal names to populations and morphological
types in these taxa (Millar 1986). It also has led them to explain the
origins and relationships of populations with often contradictory
hypotheses. Early botanists focused on, and paleontologists still rely
on, seed-cone observations in making evolutionary inferences about
the closed-cone pines. Observations of other traits often have led to
conflicting interpretations. In this paper, we focus on crossability, a
measure of genetic relatedness that estimates the potential for gene
exchange among taxa. We use this measure to test several hypotheses
about relationships among the populations of P. muricata. In par-
ticular, we consider the hypotheses that genetic distinctness among
parapatric races of P. muricata in northern California is maintained
by crossing barriers, that Monterey P. muricata crosses more readily
with northern than with southern populations, that P. muricata pop-
ulations at Purisima cross more readily with southern than with
northern populations, and that southern P. muricata populations
are isolated reproductively from the island populations of P. radiata.

VARIATION AND HYBRIDIZATION

Variation in Pinus muricata. The three Californian species of sub-
sect. Oocarpae are separated from the four taxa restricted to Mexico
and Central America by a 640 km gap (Critchfield and Little 1966).
The northern group includes P. attenuata, a montane, interior ele-
ment, and P. muricata and P. radiata, which are maritime/insular
elements (Fig. 1; Griffin and Critchfield 1976). Whereas P. attenuata
ranges widely in southern Oregon, California, and Baja California,
P. radiata is limited to three mainland populations in California
(Ano Nuevo, Monterey, and Cambria) and two distinct island pop-
ulations in Mexico (Cedros and Guadalupe Islands).

Pinus muricata comprises nine disjunct populations that extend
from Trinidad in northern California to San Vicente in northern
Baja California and to two of the Channel Islands, Santa Cruz and
Santa Rosa (Fig. 1). Of the three closed-cone pine species, P. mur-
icata has the most interpopulation variability (Fielding 1961, Doran
1974, Millar et al. 1987). Both discontinuous and clinal patterns of
variation occur (Millar 1986, Millar et al. 1987). Populations north
of Monterey (P. muricata var. borealis, Axelrod 1983) are distinct
in growth and form from the highly variable southern populations

1988] MILLAR AND CRITCHFIELD: PINUS MURICATA 4]

: OREGON
ey

Trinidad =
A
Ft. Brago——p? N
ee Ft. Rogs Ac
Mar in—Sye m ; 118
Pt. Ano NUeVO—RE: ah
Monterey." C
“Cambria A
San Luis Obispon *
A i
Purisima : ,
Sta. Barbara :
Sta. Rosa 1s, '
Sikes EhUZe Ss |
e@ * AA ;
‘
A
San Vicente
MEX I GO

Guadalupe Is. §

ff

R
Cedros Is.-@

Ue 100 600 km
Fic. 1. Distribution of P. muricata (shaded areas and areas marked M), P. radiata
(areas marked with R), and P. attenuata (enclosed with dotted lines and areas marked
with A) in California and Baja California.

~

MADRONO

[Vol. 35

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1988] MILLAR AND CRITCHFIELD: PINUS MURICATA 43

(Table 1; Duffield 1951). Pinus muricata var. borealis is further
divided into two discrete genetic races that are separated by a narrow
contact zone at Sea Ranch, Sonoma Co., California. The races are
distinguished by needle anatomy (Duffield 1951), monoterpene com-
position (Mirov et al. 1966), phenology, and isozymes (Table 1;
Millar 1983). The northern race has foliage that appears blue due
to open, wax-filled epistomatal chambers. In this paper we follow
the convention of calling all populations with this stomatal type,
‘blue P. muricata’’. The blue populations also have a distinct terpene
composition (Mirov et al. 1966). The partly closed, wax-free epi-
stomatal chambers of the southern race of var. borealis and all other
P. muricata populations southward give the foliage a yellow-green
color, here called “green P. muricata’’. Although the green needle
anatomy is typical of all populations south of Sea Ranch, the terpene
composition found in the green populations immediately south of
Sea Ranch occurs elsewhere only in the Monterey population of P.
muricata (Mirov et al. 1966). The highly variable southern mainiand
populations (P. muricata var. muricata) form a third distinct terpene
type, whereas the island populations form a fourth terpene type
(Table 1). The island pines, variously designated as P. remorata
(Mason 1930), P. muricata var. remorata (Dufheld 1951), or P.
muricata f. remorata (Hoover 1966), have high frequencies of a
deviant cone type (symmetric cones with smooth apophyses) that
also occurs in the central and southern mainland populations (Ta-
lett):

Previous reports of hybridization. Natural hybrids between P. mu-
ricata and P. radiata have been reported at Monterey, which is the
only place they are sympatric (Mason 1949, Stebbins 1950, Duffield
1951). Lack of intermediates, however, in quantitative traits (Forde
1964) and terpene compositions (Bannister et al. 1962, Forde and
Blight 1964), differences in phenology (Dufheld 1953, Critchfield
1967), and low crossability of P. muricata and P. radiata (Critchfield
1967) make this hybridization seem unlikely.

Brown (1966) and Critchfield (1967) studied interfertility among
certain populations of closed-cone pines. Using relative numbers of
viable seed produced among experimental crosses, they concluded
that P. muricata consists of three crossing groups: 1) a northern,
divergent group, that is isolated from related species and from south-
ern P. muricata populations; 2) a southern group that is able to cross
with related species; and 3) a central group that is intermediate in
crossing relationships. It is noteworthy that all interpopulation cross-
es in P. muricata between northern blue populations and green pop-
ulations south of Monterey failed. The geographically intermediate
population at Monterey crossed poorly with the southern popula-
tions, and more readily with the northern blue populations. Crosses

44 MADRONO [Vol. 35

TABLE 2. PARENT TREES OF Pinus muricata AND P. radiata CROSSES MADE IN
NATIVE POPULATIONS AND IN PLANTATIONS AT THE INSTITUTE OF FOREST GENETICS
AND RUSSELL RESERVATION.

Number of parents
Race or taxon Geographic origin Females Males

1. Native populations

Blue bishop Mendocino Co. — 4
Sonoma Co. 3 2
Green bishop Sonoma Co. 4 10
Marin Co. — 3
Monterey — 2
Purisima — 3
2. Plantations
Russell Reservation (Lafayette, CA)
Blue bishop Mendocino Co. 8 15
Green bishop Sonoma Co. 10 13
Institute of Forest Genetics (Placerville, CA)
Blue bishop Mendocino Co. 3 3
Green bishop Monterey | —
Santa Cruz Is. ] 28
Santa Rosa Is. — l
San Vicente 3 Z
Blue x green bishop Mendocino & Marin 4 4
(F)) cos., Monterey
Monterey pine Guadalupe Is. ~ —
Cedros Is. — l

between trees from contiguous blue and green stands were not in-
cluded in studies by Critchfield (1967) or Brown (1966).

All crosses attempted by Critchfield and Brown failed between
blue P. muricata and either P. attenuata or P. radiata. By contrast,
P. muricata from Monterey set some sound seeds in combinations
with mainland P. radiata and P. attenuata, whereas Channel Islands
P. muricata produced many sound seeds in those combinations.
Pinus muricata from San Vicente produced moderate amounts of
viable seed in combination with P. attenuata, but none with main-
land P. radiata, and few with P. radiata from Guadalupe Island.

MATERIALS AND METHODS

Parent trees. Crosses on P. muricata were made during two periods
and in three places (Table 2). In 1965-66 crosses were made on blue
and green trees in native stands near Sea Ranch. In the same years,
pollen parents from several native stands were crossed with females
of different origins planted at the Institute of Forest Genetics (IFG),
Placerville, California (elev. 825 m). In 1980-81 crosses were made
at IFG on a single blue tree and on four blue x green hybrids. The

1988] MILLAR AND CRITCHFIELD: PINUS MURICATA 45

hybrids had originated from 1965 crosses and were planted in 1968.
Although the trees grew poorly, by 1980 they had been sexually
mature for several years. Crosses in 1980-81 also were made on 13-
and 14-year-old trees growing in a common-garden plantation at the
Russell Reservation, Univ. of California, 16 km east of Berkeley.
Pinus muricata grows well there, and the trees had been sexually
mature for several years prior to pollination.

Pollen for the 1965-66 pollinations was collected from arboretum
and native-grown trees, and stored frozen for a year before use. Fresh
pollen from plantation trees was used for the 1980-81 crosses.

Breeding techniques and terminology. Trees were pollinated and
seeds processed using standard techniques (Cumming and Righter
1948). Seeds from 1965-66 pollinations were sorted using a Clipper
mill. If less than 10 viable seeds per cone remained, they were
x-rayed to determine viability. Viable seed yields from 1980-81
crosses were determined by germination of all harvested seeds.
Crosses that failed to yield cones or had severely insect-damaged
cones were excluded from analyses.

An attempt in this study refers to the pollination in a single season
ofa single female parent with pollen from a particular pollen source.
All but one set of crosses in this study were single-pair matings with
only one male parent contributing pollen. In these matings, the
number of attempts for a given cross also specifies the number of
male and female parents. In the single cross where a pollen mix was
used (San Vicente <X San Vicente, pollen mix of two males), each
attempt involved three parents: one female and these two males.

Crossability refers to the ease with which two taxa can be suc-
cessfully crossed, compared to control crosses within the maternal
taxon. For accurate estimation of crossability, it is essential to com-
pare between-taxon crosses to within-taxon control crosses, since
the amount of viable seed naturally produced within taxa varies.
Controls reported here for each cross combine results from control
crosses involving the same female used in the interpopulation cross-
es, and control crosses on other females of the maternal taxon. We
quantify crossability as the number of viable seeds per cone produced
from between-taxon crosses expressed as a percent of the number
of viable seeds from the within-taxon control crosses. Thus, percents
less than 100 indicate fewer viable seed produced from between-
than within-taxon control crosses, percents equal to 100 indicate an
equal number from each type of cross, and percents greater than
100 indicate more viable seed produced from between- than within-
taxon crosses. Differences in numbers of viable seeds among crosses
were tested by analysis of variance with significant differences re-
ported for p values less than 0.05. Differences among crosses in
germination of seeds were tested by chi-square analyses, with the
same significance level.

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1988] MILLAR AND CRITCHFIELD: PINUS MURICATA 47

RESULTS

Crosses of northern Pinus muricata populations. Crossability be-
tween blue and green trees from all northern sources averaged 100%
(Table 3). In crosses on native trees, the average number of sound
seeds from blue X green crosses was significantly higher than the
blue X blue control crosses. Germination percents of green x blue
and blue X green crosses did not differ significantly from the
blue x blue control crosses. In crosses at Russell Reservation, the
average numbers of sound seeds and germination percents from
green X blue crosses did not differ significantly from the green
green control crosses. The lower seed set for the blue control crosses
at Russell was probably due to premature opening of the unbagged
cones, which permitted some seed to shed before harvesting. The
percent germination of these crosses was only slightly lower than
the other control and blue x green families. Blue x green crosses
on trees at IFG produced numbers of sound seed similar to those
in blue X blue control crosses.

Crosses on F, females produced fewer sound seeds than either the
within-population crosses or crosses that used the F, as male (Table
3). At IFG, the average numbers of viable seeds and percent ger-
mination for crosses involving the F, females were significantly lower
than results from the blue < blue cross. At Russell, however, results
from backcrosses using the F,’s as males did not differ significantly
from the blue x blue and green X green crosses.

Viable seed from all blue x green crosses and crosses involving
the F,’s were sown in a nursery. Heights of seedlings from all crosses
did not differ significantly through their second year. The only dif-
ferences in survival were between crosses on F, females (x = 33%)
and all other crosses (x > 80%).

Crosses of widespread Pinus muricata populations. Crossability
generally decreased with increased distance between populations
(Table 4). The blue populations north of Sea Ranch crossed easily
with the Marin population (crossability > 100%); easily with the
Monterey population in one direction (using a blue female, > 100%),
but less easily in the reciprocal cross (52%); and did not cross at all
with green trees from Purisima or San Vicente (0%).

The green population in southern Sonoma Co. crossed less readily
than the blue population with Monterey trees (crossability = 27%)
and was nearly unsuccessful (<5%) in crosses with San Vicente trees.
Marin trees followed a similar pattern: low success with Monterey
(26%) and no seed from a single attempt with Santa Cruz Island
(0%). Crossability between Monterey and Purisima was very low
(6%).

Populations south of Monterey were highly interfertile. Cross-
ability of the San Vicente < Purisima combination was 82%, and

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48

1988] MILLAR AND CRITCHFIELD: PINUS MURICATA 49

although the control cross was lacking, the Santa Cruz x Santa Rosa
combination set more seed on average than any of the controls. In
two attempts to cross San Vicente P. muricata with Guadalupe and
Cedros Island pines, none of the 39 seeds produced was viable (0%).

First-year nursery heights and survival did not differ significantly
between seedlings from interpopulational crosses and control
crosses.

DISCUSSION

Crossability of Pinus muricata. Crossing data support conclusions
from previous studies of growth, form, and anatomical traits that
P. muricata is a highly polymorphic species, with great intra- and
interpopulation variability, especially in the southern mainland and
island populations. Relationships among the populations of Cali-
fornian closed-cone pines are illustrated in a crossing polygon (Fig.
2). We redefined three distinct breeding units (Critchfield 1967)
within P. muricata: 1) northern P. muricata populations from Marin
Co. northward; 2) southern P. muricata populations including main-
land and Channel Islands populations from Purisima southward;
and 3) Monterey P. muricata, which remains intermediate between
the first two units. Pinus radiata, including Guadalupe and Cedros
Island pines, and P. attenuata remain distinct from northern and
central bishop pine, but appear weakly related to certain southern
P. muricata populations.

The occurrence of barriers to hybridization among widespread P.
muricata populations led Critchfield (1967) to hypothesize that the
genetic differences between blue and green races in northern Cali-
fornia also are maintained by crossing barriers. Contrary to this
speculation, we found no post-pollination barriers in crosses between
contiguous blue (Mendocino and northern Sonoma cos.) and north-
ern green (central Sonoma Co.) populations. These results corrob-
orate prior indications of natural hybridization in Sonoma Co. from
terpene (Mirov et al. 1966) and isozyme evidence (Millar 1983).
Natural introgression may be inhibited, however, by differences in
flowering times and by lower fertility in the hybrid female strobili.

In the limited number of combinations we made, the Marin and
Monterey P. muricata populations responded more like northern P.
muricata than the southern populations of the species. Monterey
(green) trees set only a few viable seeds in combination with Santa
Cruz Island pines, but had moderate crossability in combinations
with Marin and Sonoma-green populations and moderate to high
crossability to Sonoma/Mendocino blue populations. The southern
breeding unit defined by Critchfield now can be expanded to include
Purisima.

Phylogenetic inferences. Patterns of crossability supplement pre-
vious studies of phylogenetic relationships in P. muricata. The ex-

50 MADRONO [Vol. 35

MENDOCINO-
BLUE
6) ESTIMATED CROSSABILITY
citer. IEE HIGH (> 80%)
we ‘ “sts
ii = NeDIUM (6-40%)

== EOWNC<6%)

NO GERMINABLE SEED SE
ONE OR MORE VALID AT

ANO NUEVO, MONTEREY

/
/
an
;
COTETrIK:)
/
/
/

oS. GUADALUPE
“OT SLAND

pootaaser@ 7
eaten ir ei 7

P. RADIATA

MONTEREY -
GREEN

PURISIMA-%

GREEN
ioe
=o Qzs==as ()
VICENTE -
STA. CRUZ STA. ROSA
GREEN ISLAND- TSLAND=
GREEN CRFEN

Fic. 2. Crossing relationships in the California closed-cone pines, including all
available information to date. All populations not labeled with a species name are
P. muricata.

istence and maintenance of abrupt genetic discontinuities in mor-
phological and biochemical traits between blue and green races of
P. muricata at Sea Ranch in northern California are difficult to
interpret. Forests of P. muricata are continuous through a narrow
(2 km) transition, and no environmental or ecological changes co-
incide with the discontinuity. Our crossing studies showed that the
races have remained interfertile. This suggests that, despite genetic
differences in several traits, the Sonoma green population and north-
ern Mendocino populations are closely related. The races appear to
have evolved recently in a mosaic pattern that did not affect inter-
population fertility except possibly at the F, level. Although blue
and green populations can hybridize, other barriers, such as differ-
ences in phenology (Millar 1983) and different responses to soils
(Millar unpubl. data), may contribute to keeping the contiguous races
distinct.

Dufheld’s (1951) proposal that a distinct northern race of P. mu-
ricata exists 1s re-enforced by reproductive barriers between northern
and southern groups. From an analysis of many traits, Dufhield con-
cluded that populations in Humboldt, Mendocino, and Sonoma cos.
are a distinct variety and that Marin and Monterey are intermediate
between this northern variety and the rest of the species. Axelrod

1988] MILLAR AND CRITCHFIELD: PINUS MURICATA 51

(1983) formally published the northern variety as P. muricata var.
borealis. He based the description, however, only on cone shape and
extended var. borealis south to include some “‘relict trees” at Mon-
terey. Axelrod’s designation leaves P. muricata var. borealis unsat-
isfactorily heterogeneous in other traits (Table 1), and suggests that
two sympatric varieties somehow maintain their genetic integrity at
Monterey. Crossing relationships show Marin and Monterey pop-
ulations to be distinct and intermediate, although they have greater
affinities to northern than to southern populations. We suggest that
the varietal designation conservatively be limited to populations
north of Marin Co.

The infertility between the northern populations of P. muricata
(e.g., Mendocino) and P. radiata and P. attenuata suggests that var.
borealis has diverged considerably from common ancestors of the
Californian closed-cone pines, and from closely related species. In
contrast, the southern populations of P. muricata, especially the San
Vicente and Channel Islands populations, are sufficiently similar to
allow successful interspecific hybridization. Apparently evolution in
these taxa has not affected hybridization potential.

Evolutionary interpretations of central and southern P. muricata
populations (summarized in Millar 1986) have also differed de-
pending on the emphasis given cone morphology. Mason (1930,
1949) and Axelrod (1967, 1980, 1983) argue that pines having sym-
metric cones with smooth apophyses represent an independent evo-
lutionary lineage (P. remorata Mason). Mason restricts this desig-
nation to pines with symmetric cones on the Channel Islands, whereas
Axelrod uses the name for all trees with this cone type wherever
they occur on the islands and mainland. Both authors suggest that
the present variation in cone traits, typical of many stands south of
Sonoma Co., resulted from hybridization of P. remorata with P.
muricata. Other authors have concluded that cone shape is just one
of many variable traits in P. muricata (Dufheld 1951, Fielding 1961,
Linhart et al. 1967, Doran 1974). In quantitative analyses, Linhart
et al. (1967) found that distinct variation in resin canals, terpenes,
and several needle anatomy traits did not correlate with cone vari-
ation, and concluded that P. remorata was “primarily a name given
a particular cone type in a variable species”’.

We found no support from crossing studies for the hypothesis that
P. remorata is a distinct taxon from P. muricata. Although most of
our breeding trees were not identified individually by cone type, all
our pollen and seed lots from Marin south contained trees with the
smooth, symmetric cone type. Pines sampled from the Channel
Islands, especially those from Santa Rosa, had high frequencies of
smooth cones. We found no pattern of crossability to suggest that
these trees were distinct taxonomically. The Channel Islands pines
resembled southern populations of P. muricata in crossing behavior

52 MADRONO [Vol. 35

among all populations tested. Furthermore, if P. remorata extends
north to Monterey and Marin (Axelrod 1980), we would expect to
find greater crossability between those populations and Channel Is-
lands populations than was found. Observations on resin canals
indicated that number of canals varied greatly among trees, and that
variation was related to geographical location and not to cone type.
Thus, we found no evidence that smooth, symmetric cones found
on trees throughout the species are indicators of an independent
lineage.

CONCLUSIONS

Crossing results reported here supplemented and corroborated
other studies on P. muricata which indicate that complex patterns
of variation exist in the species. Unique in Pinus is the presence of
intraspecific post-pollination barriers among P. muricata popula-
tions. These barriers, together with distinguishing patterns of vari-
ation in other traits, suggest that the northern and southern popu-
lations have long been isolated and perhaps should be considered
distinct species. By contrast, evolution of genetic differences between
blue and green races within the northern populations has not been
accompanied by evolution of post-reproductive barriers. Genetic
differences between these races must be maintained by other factors.

Since southern P. muricata populations retain crossability to P.
radiata and P. attenuata, the great variation in these populations
may have been imported through prior interspecific hybridization.
Patterns of crossability, coupled with evidence from variation in
other traits, gave no evidence to suggest that the smooth, symmetric
cone type alone is an indicator of a distinct evolutionary lineage
within the species complex. This cone type is found in nearly all P.
muricata populations and appears to be one of many polymorphic
traits in the species.

ACKNOWLEDGMENTS

We thank J. Duffield, J. Griffin, J. Haller, Y. Linhart, K. Rindlaub, S. Strauss, B.
Tanowitz, and the editor for their comments on various drafts of the manuscript.

LITERATURE CITED

AXELROD, D. I. 1967. Evolution of the California closed-cone pine forest. Jn R. W.
Philbrick, ed., Proceedings of the symposium on the biology of the California
islands, p. 93-150. Santa Barbara Botanic Garden.

. 1980. History of the maritime closed-cone pines, Alta and Baja California.

Univ. California Press, Berkeley.

. 1983. New Pleistocene conifer records, coastal California. Univ. Calif. Publ.
Geo. Sci. 127.

BANNISTER, M. H., A. L. WILLIAMS, I. R. C. MCDONALD, and M. B. ForDE. 1962.
Variation of turpentine composition in five population samples of Pinus radiata.
New Zealand J. Sci. 5:486-495.

1988] MILLAR AND CRITCHFIELD: PINUS MURICATA 3

Brown, A. G. 1966. Isolating barriers between the California closed cone pines.
M.S. thesis, Univ. Sydney.

CRITCHFIELD, W. B. 1967. Crossability and relationships of the California closed
cone pines. Silvae Genetica 16:89-97.

and E. L. Litre. 1966. Geographic distribution of pines of the world.
U.S.D.A. Forest Serv. Misc. Publ. 991.

CUMMING, W. C. and F. I. RIGHTER. 1948. Methods used for control pollination of
pines in the Sierra Nevada of California. U.S.D.A. Circular 792.

DorAN, J. C. 1974. Variation in growth of Pinus muricata provenances and com-
parison with Pinus radiata. Austral. Forest Res. 6(3):19-24.

DUFFIELD, J. W. 1951. Interrelationships of the California closed cone pines with
special reference to Pinus muricata D. Don. Ph.D. dissertation, Univ. California,
Berkeley.

. 1953. Pine pollen collection dates—annual and geographic variation.
U.S.D.A. Forest Serv. Forest Res. Notes 85.

FIELDING, J. M. 1961. Provenances of Monterey and bishop pines. Austral. Forest
Timb. Bureau Bull. 38:1-30.

Forbes, M. B. 1964. Variation in natural populations of Pinus radiata in California.
Parts 1-4. New Zealand J. Bot. 2:213-—259, 459-501.

and M. M. BLicHT. 1964. Geographical variation in the turpentine of bishop
pine. New Zealand J. Bot. 2:44—-52.

GRIFFIN, J. R. and W. B. CRITCHFIELD. 1976. The distribution of forest trees in
California. U.S.D.A. Forest Serv. Res. Pap. PSW-82.

Hoover, R. F. 1966. Miscellaneous new names for California plants. Leafl. W. Bot.
10:337-338.

LINHART, Y. B., B. Burr, and M. T. CONKLE. 1967. The closed-cone pines of the
northern Channel Islands. Jn R. B. Philbrick, ed., Proceedings of the symposium
on the biology of the California islands, p. 151-177. Santa Barbara Botanic
Garden.

Mason, H. L. 1930. The Santa Cruz Island pine. Madrono 2:8-10.

1949. Evidence for the genetic submergence of Pinus remorata. In G. L.
Jepsen, G. G. Simpson, and E. Mayr, eds., Genetics, paleontology and evolution,
p. 356—362. Princeton Univ. Press, Princeton, NJ.

MILLAR, C. I. 1983. A steep cline in Pinus muricata. Evolution 37:311-319.

1986. The California closed cone pines (subsection Oocarpae Little and

Critchfield): a taxonomic history and review. Taxon 35:657-670.

, S. H. Strauss, M. T. CONKLE, and R. D. WESTFALL. 1987. Allozyme
differentiation and biosystematics of the closed-cone pines (subsection Oocarpae
Little and Critchfield, genus Pinus). Syst. Bot. 13(3), in press.

Mirov, N. T., E. ZAVARIN, K. SNAJBERK, and K. COSTELLO. 1966. Further studies
of Pinus muricata in relation to its taxonomy. Phytochemistry 5:343-355.
SHELBOURNE, C. J. A. 1974. Recent investigations of wood properties and growth

performance in Pinus muricata. New Zealand J. Forest. 19:13-—45.

, M. H. BANNISTER, and M. D. WiLcox. 1982. Early results of provenance
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STEBBINS, G. L. 1950. Variation and evolution in plants. Columbia Univ. Press,
New York.

(Received 27 Jan 1987; revision accepted 19 Nov 1987.)

ENDEMIC VASCULAR PLANTS OF NORTHWESTERN
CALIFORNIA AND SOUTHWESTERN OREGON

JAMES P. SMITH, JR. and JOHN O. SAWYER, JR.
Department of Biological Sciences, Humboldt State University,
Arcata, CA 95521

ABSTRACT

An account of the endemic vascular plants of northwestern California and south-
western Oregon is presented. This component of the California Floristic Province,
known for its high species richness, was found to have a large number of neoendemics
in a regional flora generally characterized as relictual in nature. A list of endemics
provides distribution by county and formal designations of rarity and endangerment
for 281 taxa in 42 families.

The flora of northwestern California and southwestern Oregon has
long been known for its high floristic richness (Stebbins and Major
1965). We estimate that there are over 3500 taxa of vascular plants,
in about 150 families and 760 genera, in northwestern California
alone (Smith and Sawyer 1987). The region may be viewed as the
last major frontier along the Pacific Coast to be studied in detail.
Intensive collecting began only about 20 years ago, after the pi-
oneering work of Alice Eastwood, Joseph P. Tracy, Thomas Jefferson
Howell, Edward Greene, Milo Baker, and Doris Niles. Our work in
the last two decades, along with our students and colleagues (Muth
1967, Ferlatte 1974, Oettinger 1975, Barker 1979, Nelson 1979,
Stillman 1980, Whipple 1981), has resulted in a more detailed
knowledge of this interesting flora. Recent efforts at determining the
status of rare and endangered plants in both states also has added
greatly to our knowledge (Siddall et al. 1979, Meinke 1981, Smith
and York 1984, Soper et al. 1985, York 1985). We now have a good
account of the endemic vascular flora of this region.

STUDY AREA

Northwestern California and southwestern Oregon, a region of
about 55,000 km7?, are considered part of the California Floristic
Province (Howell 1955, 1956, 1957, Noldenke and Howell 1960,
Stebbens and Major 1965, Raven and Axelrod 1978). Therefore, for
both floristic and geographic reasons, the plants endemic to south-
western Oregon also are included here. Whereas the Klamath Moun-
tains and the North Coast Ranges represent only 15% of the area of
the California Floristic Province, they include some 65% of the 4452
native taxa found growing in the province as a whole (Raven and
Axelrod 1978). The floristic diversity here is exceptional.

MADRONO, Vol. 35, No. 1, pp. 54-69, 1988

1988] SMITH AND SAWYER: ENDEMIC PLANTS

BANDON
43°N
PORT

42°N

CRESCENT
CITY

4I°N

40°N

PACIFIC
OCEAN

39°N

l24°W

SAN
FRANCISC OF

55

Fic. 1. Location of the study area in northwestern California and southwestern

Oregon (shown in white).

56 MADRONO [Vol. 35

The study area (Fig. 1) centers on the Klamath Mountains, a
geologic province of ancient rocks and landforms (Irwin 1960, 1981).
They constitute a poorly defined set of ranges, including the Eddies,
Marbles, Salmons, Scott Bars, Scotts, Siskiyous, Trinities, Trinity
Alps, and the Yolla Bollys. We also include the topographically
continuous North Coast Ranges of California to the west and to the
south of the Klamath Mountains. Lake and Mendocino cos. mark
the southern extent of the region; counties farther south lack the
montane environments present to Snow Mountain (Heckard and
Hickman 1984). Northwestern California is treated by a checklist
of the vascular plants (Smith and Sawyer 1987) and by a key to
families and genera (Smith and Sawyer 1981).

The traditional explanation for the area’s rich flora is that it is a
mixture of California and northern plants. It is not surprising that
plants from the north and from the south occur here. The area is
geographically and environmentally central on the West Coast of
North America (Whittaker 1961), and geologically complicated, with
its many disjunct areas of ultramafic rock (Whittaker 1960, Sawyer
and Thornburgh 1977, Kruckeberg 1984). In addition, the ancient
terrain supports great habitat variety in a moderated, maritime cli-
mate (Richerson and Lum 1980). The area, especially the Klamath
Mountains, is viewed as a refugium of Tertiary plants (Wolfe 1969,
Axelrod 1976).

TYPES OF ENDEMISM

The relictual nature of the flora is seen in many families and
genera, although not all of them appear in the list of endemic taxa
because they also occur outside of the region. Aruncus dioicus, Ca-
lypso bulbosa, Darlingtonia californica, Disporum hookeri, Euony-
mous occidentalis, Mahonia nervosa, Polystichum munitum, Sequoia
sempervirens, and Trautvetteria carolinensis are typical of the many
‘““Arcto-Tertiary” plants that grow in the Klamath Mountains or
along the coast at lower elevations.

In addition, the flora is seen as being enriched by plants of Mexican
origin, such as Arbutus, Garrya, and Gaultheria that now grow with
Sequoia in the redwood forest (Abrams 1925, Axelrod 1977). Many
of these southern elements grow in the woodlands, chaparral, and
grasslands found at lower elevations or near the coast. The events
of the Pleistocene and hypsithermal are also seen as causing further
accumulations of various plants from the north, such as Empetrum
nigrum and Menyanthes trifoliata; of Purshia tridentata and For-
sellesia stipulifera from the Great Basin; and of Pinus sabiniana from
central California.

The relictual nature of the flora can also be evaluated by a review
of a list of endemics (Appendix 1). Plants without close relatives or

1988] SMITH AND SAWYER: ENDEMIC PLANTS a7

those whose close relatives are disjunct are typically considered pa-
leoendemic or relicts (Stebbins 1980). Ka/miopsis leachiana, Picea
breweriana, Quercus sadleriana (Tucker 1983), and Cornus sessilis
are good examples. But the list contains surprisingly few relicts.

In this geologically stable area, with its moderated climate, we
might also expect to find a larger number of endemic species, and
perhaps even endemic genera (Kruckeberg and Rabinowitz 1985).
Only two monotypic genera, Bensoniella and Tracyina, are endemic.
Others, such as Cycladenia humilis, Darlingtonia californica, and
Whitneya dealbata, often thought to be endemic to the region, are
not.

To summarize, the centrally positioned, continuous montane en-
vironment among the North Coast Ranges, the Klamath Mountains,
the Cascades, and the Sierra Nevada accounts for much of the flo-
ristic richness, but not for the degree of endemism. Similarly, the
invoking of paleoendemism, taken by itself, is not adequate.

ANALYSIS OF THE ENDEMIC FLORA

In surveying the list of endemic taxa, we were impressed by the
large number of infraspecific taxa. In this observation lies another
explanation for the local level of endemism. Some of the taxa, such
as Iris tenax subsp. klamathensis, Dicentra formosa subsp. oregana,
and Holodiscus discolor var. delnortensis, represent regional variants
of widespread, western species. Others, such as Juniperus communis
var. jackii and Chlorogalum pomeridianum var. minus, are typical
of serpentine substrates. Some plants, as in 7ri//ium ovatum subsp.
oettingerl, grow at higher elevations than do the typical forms of the
species. Still others, such as Monardella odoratissima subsp. pallida
and Penstemon newberryi subsp. berryi, appear to be local expres-
sions of common Sierran species.

To evaluate further the list of endemics, genera with five or more
taxa were singled out and appear in Table 1. Many of them, such
as Arabis, Penstemon, or Lupinus, are expected, as they are known
for their diversity in the western United States. Other large genera,
such as Aster, Carex, Lotus, or Phlox, are conspicuously absent.

The number of endemic species can be compared to the total taxa
in each genus. For example, Phacelia is a genus of about 200 species,
of which 29 grow in the area, seven of them endemic. A few genera,
such as Arabis, Horkelia, Lilium, and Limnanthes, stand out as being
unusually high in regional endemics. Of all of the taxa tallied, Lew-
isia, Sedum, and Sidalcea have an exceptionally high number of
regional endemics.

Such comparisons might be better judged in a larger geographical
context. Table 1 also shows the number of taxa for California (Munz
1959, 1968). A larger number of species and infraspecific taxa would

58 MADRONO [Vol. 35

TABLE 1. GENERA IN NORTHWESTERN CALIFORNIA AND SOUTHWESTERN OREGON
WITH FIVE OR MORE ENDEMIC TAXA. The fraction represents the number of species/
number of subspecific taxa. Estimates for size in each genus are after Willis (1973);
those with ‘**”’ are from Raven and Axelrod (1978). Taxa in the region itself are from
Peck (1961), Smith and Sawyer (1987), and recent monographs. The values in pa-
rentheses are species : taxa ratios. If all taxa are at the species rank, the ratio equals
1.0.

No.
species Endemics in Taxa in
Genus per genus area Taxa in area California
I. Large genera, <100 species
Arabis 120 LS led) 24/26 (E22) 35/52 (1.5)
Epilobium 215 6/6 (1.0) 18/22 (1.2) 36/22 (1.6)
Eriogonum 250 10/11 (1.1) 33/47 (1.4) 104/158 (1.5)
Lupinus 2007 8/9 (1.1) 36/54 (1.5) 82/144 (1.8)
Penstemon 2IOF 5/5 (1.0) 20/27 (1.4) 49/75 (1.2)
Phacelia 200 7/7 (1.0) 29/31 (1.1) 91/116 (1.3)
Plagiobothrys 100 4/5 (1.2) 17/20 (1.1) 39/50 (1.3)
Sedum 600 5/10 (2.0) 11/20 (1.8) 12/18 (1.5)
II. Moderate-sized genera, 10-80 species
Arctostaphylos 50* 7/7 (1.0) 16/21 (1.3) 32/5307)
Calochortus 60 6/6 (1.0) 16/18 (1.1) 39/52 (1.3)
Horkelia 30 4/5 (1.2) 8/10 (1.2) 16/25 (1.6)
Lewisia 20 3/6: (2:0) 8/11 (1.4) 13/18 (1.4)
Lilium 80 5/5 (1.0) fey te) 15/19 (1.3)
Limnanthes 10* 375-1) 4/9 (2.3) GAS (G6)
Sidalcea 25 3/7243) 9/21 (2.3) 18/33 (1.8)

be expected for this larger area. One way to reduce the effect of area
is to express the numbers as ratios. Lupinus, for example, is a genus
of about 200 species. Munz reports 82 species and 144 subspecies
and varieties in California. There are, then, almost two infraspecific
taxa per species of Lupinus in the state.

When northwest California is compared to the state as a whole,
a predicted pattern is seen, 1.e., the smaller the area, the smaller the
ratio. California includes those taxa of the Sierra Nevada, the Cas-
cades, the Klamaths, and the North Coast in the tally, so that the
ratio would be larger than that for the northwest section of the state
alone. Furthermore, the ratio for endemics would be expected to be
even smaller yet, because they are restricted to a smaller area. The
expected ratio reduction does occur for most of the genera in Table
1. Exceptions are Lewisia, Sedum, and Sidalcea, where the ratio
increases. This is taken as evidence that adaptive radiation is oc-
curring in the region. We conclude, therefore, that northwest Cali-
fornia and southwest Oregon is not only a refugium, but it is also
an area of active diversification today.

An abundance of local varieties and subspecies is expected as
populations adapt to the unique set of environmental controls

1988] SMITH AND SAWYER: ENDEMIC PLANTS 5D

(Kruckeberg and Rabinowitz 1985). The region’s heterogeneity of
topography and parent material offers the setting for this diversifi-
cation. Stebbins and Major (1965), using Lake Co., California, an
area containing volcanic, sedimentary, and ultramafic substrates,
argued that under such settings neoendemics would be developed
during periods of changing climate. Axelrod (1982) makes a similar
argument for the Monterey endemic area. The celebrated patchy
matrix of habitats found in northwest California and southwest Or-
egon supplies a larger stage for the addition of a high number of new
taxa into the region’s flora during the recent period of climatic change.

ACKNOWLEDGMENTS

We thank the Curators of CAS, JEPS, ORE, OSU, and UC for allowing us to
examine specimens in their herbaria; Kenton Chambers, Lawrence Heckard, Veva
Stansell, and David Wagner for their comments on the list of endemic taxa; and the
Rare Plant Program of the California Native Plant Society and the Natural Diversity
Data Base of the California Department of Fish and Game for distribution data on
rare, threatened, and endangered plants.

LITERATURE CITED

ABRAMS, L. 1925. The origin and geographical affinities of the flora of California.
Ecology 6:1-6.

AXELROD, D. I. 1976. History of the coniferous forests, California and Nevada.
Univ. Calif. Publ. Bot. 70:1-62.

1977. Outline history of California vegetation. Jn M. G. Barbour and J.

Major, eds., Terrestrial vegetation of California, p. 140-193. Wiley-Interscience,

New York.

. 1982. Age and origin of the Monterey endemic area. Madrono 29:127-147.

BARKER, L. M. 1979. A flora of the Old Gasquet Toll Road, Del Norte County,
California. M.A. thesis, Humboldt State Univ., Arcata, CA.

FERLATTE, W. J. 1974. A flora of the Trinity Alps of northern California. Univ.
California Press, Berkeley.

HECKARD, L. R. and J. C. HICKMAN. 1984. The phytogeographical significance of
Snow Mountain, North Coast Ranges, California. Madrono 31:30-47.

Howe LL, J. T. 1955. A tabulation of California endemics. Leafl. W. Bot. 7:257-
205.

. 1956. The numerical summary of California plants and endemism. Leafl.

W. Bot. 8:59-60.

. 1957. The California floral province and its endemic genera. Leafl. W. Bot.
8:138-141.

IRwINn, W. P. 1960. Geological reconnaissance of the northern Coast Ranges and
Klamath Mountains, California, with a summary of mineralogical resources.
Calif. Mines and Geol. Bull. 179.

1981. Tectonic accretion of the Klamath Mountains. /n W. G. Ernst, ed.,
The geotectonic development of California, p. 29-49. Prentice Hall, Englewood
Cliffs, NJ.

KRUCKEBERG, A. R. 1984. California serpentines: flora, vegetation, geology, soils
and management problems. Univ. Calif. Publ. Bot. 78:1—180.

and D. RABINOWITZ. 1985. Biological aspects of endemism in higher plants.
Ann. Rev. Ecol. Syst. 16:447-479.

MEINKE, R. J. 1981. Threatened and endangered vascular plants of Oregon: an
illustrated guide. United States Fish and Wildlife Service, Office of Endangered
Species, Portland, OR.

60 MADRONO [Vol. 35

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

. 1968. Supplement to a California flora. Univ. California Press, Berkeley.

Muth, G. J. 1967. A flora of Marble Valley, Siskiyou County, California. M.S.

thesis, Pacific Union College, Angwin, CA.

NELson, T. W. 1979. A flora of the Lassics, Humboldt and Trinity counties, Cal-

ifornia. M.A. thesis, Humboldt State Univ., Arcata, CA.

NOLDEKE, A. M. and J. T. HOWELL. 1960. Endemism and a California flora. Leaf.
W. Bot. 9:124—-127.

OETTINGER, F. W. 1975. The vascular plants of the High Lake Basins in the vicinity
of English Peak, Siskiyou County, California. M.A. thesis, Claremont Graduate
School, Claremont, CA.

Peck, M. E. 1961. A manual of the higher plants of Oregon. 2nd ed. Binsford &
Mort, Portland, OR.

RAVEN, P. H. and D. I. AXELRop. 1978. Origin and relationships of the California
flora. Univ. Calif. Publ. Bot. 72:1-134.

RICHERSON, P. J. and K. L. Lum. 1980. Patterns of plant species diversity in Cal-
ifornia: relation to weather and topography. Amer. Nat. 116:504—536.

SAWYER, J. O. and D. A. THORNBURGH. 1977. Montane and subalpine vegetation
of the Klamath Mountains. 7n M. G. Barbour and J. Major, eds., Terrestrial
vegetation of California, p. 669-732. Wiley-Interscience, New York.

SIDDALL, J. L., K. L. CHAMBERS, and D. H. WAGNER. 1979. Rare, threatened and
endangered vascular plants in Oregon—an interim report. Oregon Natural Area
Preserves Advisory Committee, Division of State Lands, Salem, OR.

SMITH, J. P. and J. O. SAwyeR. 1981. Keys to the families and genera of vascular
plants of northwest California. 4th ed. Mad River Press, Eureka, CA.

and 1987. Achecklist of the vascular plants in northwest California.

9th ed. Humboldt State Univ. Herbarium Misc. Publ. No. 2, Arcata, CA.

and R. York. 1984. Inventory of rare and endangered vascular plants of
California. Special Publication No. |. (3rd ed.) California Native Plant Society,
Berkeley, CA.

SoOPER, C., J. KAGAN, S. YAMAMOTO, C. LEVESQUE, and J. GAMON. 1985. Rare,
threatened and endangered plants and animals of Oregon. Oregon Natural Her-
itage Data Base, Portland.

STEBBINS, G. L. 1980. Rarity of plant species: a synthetic viewpoint. Rhodora 82:
77-86.

and J. MAyor. 1965. Endemism and speciation in the California flora. Ecol.
Monogr. 35:1-35.

STILLMAN, K. T. 1980. Meadow vegetation on metasedimentary and metavolcanic
parent materials in the north central Marble Mountains, California. M.A. thesis,
Humboldt State Univ., Arcata, CA.

Tucker, J. M. 1983. California’s native oaks. Fremontia 1 1(3):3-12.

WHIPPLE, J. 1981. A flora of Mt. Eddy, Klamath Mountains, California. M.A. thesis,
Humboldt State Univ., Arcata, CA.

WHITTAKER, R. H. 1960. Vegetation of the Siskiyou Mountains, Oregon and Cal-
ifornia. Ecol. Monogr. 30:279-338.

1961. Vegetation history of the Pacific coast states and the central signifi-
cance of the Klamath Region. Madrono 16:5-23.

WILLIS, J. C. 1973. A dictionary of the flowering plants and ferns. 8th ed. revised
by H. K. Airy Shaw. Cambridge Univ. Press, Cambridge.

Wo LF, J. A. 1969. Neogene floristic and vegetational history of the Pacific north-
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(Received 21 Jul 1986; revision accepted 11 Aug 1987.)

1988] SMITH AND SAWYER: ENDEMIC PLANTS 61

APPENDIX |. ANNOTATED CATALOGUE OF ENDEMIC VASCULAR PLANTS

County abbreviations: CALIFORNIA. COL = Colusa; DNT = Del Norte; GLE =
Glenn; HUM = Humboldt; LAK = Lake; MEN = Mendocino; SHA = Shasta;
SIS = Siskiyou; TEH = Tehama; TRI = Trinity. OREGON. COS = Coos; CUR =
Curry; DOU = Douglas; JAC = Jackson; JOS = Josephine.

A-1 to A-4 indicates that the plant is listed in Smith and York (1984).
A-| = Plants of highest priority
A-2 = Plants rare or endangered in California, but more common elsewhere
A-3 = Plants about which we need more information
A-4 = Plants of limited distribution

B-1 to B-3 indicates that the plant is listed in Siddall et al. (1979).
B-la = Very local endemic

B-1b = Regional endemic

B-2a = Plants thinly scattered over a wide range

B-2b = Known only from a few widely disjunct populations

B-3 = Of concern in Oregon, more abundant elsewhere
C indicates that the plant is listed in Meinke (1981).
D-1 to D-3 indicates that the plant is listed in Soper et al. (1985).

D-1 = Taxa endangered or threatened throughout range, including those possibly
extinct

D-2 = Taxa endangered or threatened in Oregon, but more common or stable
elsewhere

D-3 = Taxa limited in abundance throughout range, but currently stable

PINOPHYTA
Cupressaceae

Juniperus communis Linnaeus var. jackii Rehder. DNT, SIS; CUR.

Pinaceae
Picea breweriana Watson. DNT, HUM, TRI, SIS; COS, CUR, JAC, JOS.

MAGNOLIOPHYTA: MAGNOLIOPSIDA
Berberidaceae

Mahonia nervosa (Pursh) Nuttall var. mendocinensis (Roof) Roof. MEN; A-3.
Vancouveria chrysantha Greene. DNT, SIS; CUR, JOS; A-4, B-1b.

Boraginaceae

Cryptantha clevelandii Greene var. dissita (1. M. Johnston) Jepson & Hoover. LAK.

Cryptantha crinita Greene. SHA, TEH; A-1.

Hackelia amethystina J. T. Howell. GLE, LAK, MEN, TEH, TRI; A-4.

Hackelia bella (Macbride) I. M. Johnston. HUM, MEN, SIS, TEH, TRI.

Plagiobothrys hirtus (Greene) I. M. Johnston var. corallicarpa (Piper) I. M. Johnston.
JAC, JOS; B-1b, C, D-1.

Plagiobothrys hirtus (Greene) I. M. Johnston var. hirtus. DOU; B-1b, C.

Plagiobothrys lamprocarpus (Piper) I. M. Johnston. JOS; B-la, C, D-1.

Plagiobothrys lithocaryus (Greene ex A. Gray) I. M. Johnston. LAK, MEN; A-4.

Plagiobothrys tener (Greene) I. M. Johnston var. subglaber I. M. Johnston. LAK.

Campanulaceae
Campanula shetleri Heckard. SHA, SIS; A-1.

62 MADRONO [Vol. 35

Caryophyllaceae

Minuartia decumbens T. W. & J. P. Nelson. SHA, TEH, TRI; A-1.

Minuartia howellii (Watson) Mattfeld. DNT; CUR, JOS; A-4.

Minuartia nuttallii (Pax) Briquet subsp. gregaria (Heller) Maguire. COL, HUM, LAK,
MEN, SIS, TEH, TRI; CUR, JAC, JOS.

Minuartia rosei (Maguire & Barneby) McNeill. SHA, TEH, TRI; A-4.

Silene campanulata Watson subsp. campanulata. MEN; A-1.

Silene hookeri Nuttall ex Torrey & A. Gray subsp. bolanderi (A. Gray) Abrams.
HUM, MEN, TRI; CUR, JOS; B-3, D-2.

Silene hookeri Nuttall ex Torrey & A. Gray subsp. pulverulenta (Peck) Hitchcock &
Maguire. JAC, JOS; B-1b.

Silene marmorensis Kruckeberg. SIS; A-1.

Compositae

Antennaria suffrutescens Greene. DNT, HUM; CUR, JOS; A-4.

Arnica cernua T. J. Howell. DNT, HUM, SIS, SHA, TRI; COS, CUR, JOS; A-4.

Arnica spathulata Greene subsp. eastwoodiae (Rydberg) Ediger & Barkley. DNT,
HUM, SIS; CUR, DOU, JAC, JOS; A-4.

Arnica venosa Hall. SHA, TRI; A-4.

Aster brickellioides Greene var. brickellioides. DNT, SIS; CUR, JAC, JOS; A-4.

Aster siskiyouense Nelson & Macbride. GLE, SIS, TEH, TRI; JAC, JOS.

Balsamorhiza sericea Weber. TRI, SIS; JOS; A-4, B-1b, C, D-1.

Chaenactis suffrutescens A. Gray. SIS, TRI; A-4.

Cirsium acanthodontum Blake. DNT, HUM; COS, CUR, DOU.

Cirsium ciliolatum (Henderson) J. T. Howell. SIS; JAC; A-1, B-1b, D-3.

Erigeron bloomeri A. Gray var. nudatus (A. Gray) Cronquist. DNT, SIS; JAC, JOS;
A-2.

Erigeron bloomeri A. Gray var. pubens Keck. SIS, TEH, TRI.

Erigeron delicatus Cronquist. DNT; CUR, JOS; A-3, B-1b, C.

Erigeron flexuosus Cronquist. DNT, SHA, TRI; A-1.

Eriophyllum lanatum (Pursh) Forbes var. aphanactis J. T. Howell. GLE, COL, LAK.

Eriophyllum lanatum var. lanceolatum(T. J. Howell) Jepson. DNT, HUM, SIS, TEH,
TRI; CUR, JAC, JOS.

Eupatorium shastense Taylor & Stebbins. SHA; A-4.

Grindelia stricta De Candolle subsp. blakei (Steyermark) Keck. HUM; A-1.

Haplopappus ophitidis (J. T. Howell) Keck. SHA, TEH, TRI; A-4.

Haplopappus racemosus (Nuttall) Torrey subsp. congestus (Greene) Hall. DNT; CUR,
DOU, JAC, JOS; A-4, C.

Haplopappus racemosus subsp. pinetorum Keck. SIS, TRI.

Hazardia whitneyi (A. Gray) Greene var. discoideus (J. T. Howell) D. Clark. GLE,
HUM, LAK, SIS, TRI; DOU, JOS; B-3, D-2.

Helianthella californica A. Gray var. shastensis W. Weber. SHA, SIS, TRI.

Hemizonia calyculata (Babcock & Hall) Keck. LAK, MEN; A-4.

Hemizonia tracyi (Babcock & Hall) Keck. HUM, MEN, TRI; A-4.

Heterotheca breweri (A. Gray) Shinners var. multibracteata Jepson. SIS, TEH, TRI.

Lasthenia macrantha (A. Gray) Greene subsp. prisca Ornduff. CUR; B-1b, C, D-3.

Madia doris-nilesiae T. W. Nelson & J. P. Nelson. TRI.

Madia gracilis (Small) Keck subsp. pi/osa Keck. HUM.

Madia stebbinsii T. W. & J. P. Nelson. TEH, TRI; A-1!.

Microseris detlingii Chambers. JAC; B-1la, C.

Microseris howellii A. Gray. DNT; CUR, JAC, JOS; B-1b, C, D-1.

Microseris laciniata (Hooker) Schulz-Bipontinus subsp. siskiyouensis Chambers. DNT,
HUM, SIS; CUR, JOS.

Raillardella pringlei Greene. SIS, TRI; A-1.

Rudbeckia californica A. Gray var. glauca Blake. DNT, TRI; CUR, DOU, JOS.

1988] SMITH AND SAWYER: ENDEMIC PLANTS 63

Rudbeckia californica A. Gray var. intermedia Perdue. SIS, TRI.
Senecio greenei A. Gray. GLE, LAK, MEN, TRI.

Senecio hesperius Greene. CUR, JOS; B-1b, C, D-1.

Tracyina rostrata Blake. HUM, LAK; A-1.

Wyethia longicaulis A. Gray. HUM, MEN, TRI; A-4.

Convolvulaceae

Calystegia collina (Greene) Brummitt subsp. tridactylosa (Eastwood) Brummitt. MEN.

Crassulaceae

Parvisedum leiocarpum (H. K. Sharsmith) Clausen. LAK; A-1.

Sedum laxum (Britton) Berger subsp. eastwoodiae (Britton) Clausen. MEN; A-1.

Sedum laxum (Britton) Berger subsp. flavidum Denton. DNT, HUM, SIS, TRI;
A-1.

Sedum laxum (Britton) Berger subsp. heckneri (Peck) Clausen. DNT, HUM, SIS,
TRI; CUR, JAC, JOS; A-4, D-3.

Sedum laxum (Britton) Berger subsp. /atifolium Clausen. DNT.

Sedum laxum (Britton) Berger subsp. /axum. DNT, SIS; CUR, JAC, JOS.

Sedum moranii Clausen. JOS; C, D-1.

Sedum oblanceolatum Clausen. SIS; JAC; C.

Sedum obtusatum A. Gray subsp. paradisum Denton. SHA, TRI; A-1.

Sedum obtusatum A. Gray subsp. retusum (Rose) Clausen. LAK, MEN, SIS, TRI;
CUR, JAC.

Sedum radiatum Watson subsp. depauperatum Clausen. SIS; JOS; A-3.

Cruciferae

Arabis aculeolata Greene. DNT, SIS; CUR, JOS; A-1, C.

Arabis koehleri T. J. Howell var. koehleri. DOU, JOS; B-1b, C, D-1.

Arabis koehleri T. J. Howell var. stipitata Rollins. CUR, JOS; C, D-3.

Arabis macdonaldiana Eastwood. DNT, MEN; CUR, JOS; A-2, C, D-1.

Arabis oregona Rollins. MEN, SIS, TRI; JAC, JOS; A-3.

Arabis rigidissima Rollins. HUM, SIS, TRI, A-4.

Arabis serpentinicola Rollins. SIS; CUR; A-1, B-1b, C.

Arabis subpinnatifida Watson. GLE, HUM, MEN, SIS; DOU, JAC, JOS.
Cardamine gemmata Greene. DNT, SIS; CUR, JAC, JOS; A-1, B-1b, D-2.
Draba carnosula O. E. Schulz. SIS, TRI; A-1.

Draba howellii Watson. DNT, HUM, SHA, SIS, TRI; JOS; A-4, B-1b, D-1.
Draba pterosperma Payson. SIS; A-4.

Streptanthus barbatus Watson. SIS, TEH, TRI.

Streptanthus howellii Watson. DNT; CUR, JOS; A-2, B-1b, C, D-1.
Streptanthus tortuosus Keller var. pallidus Jepson. HUM, SIS, TRI.

Thlaspi montanum Linnaeus var. californicum (Watson) P. Holmgren. HUM; A-1.
Thlaspi montanum Linnaeus var. siskiyouense P. Holmgren. CUR, JOS; C, D-3.

Cuscutaceae

Cuscuta salina Engelmann var. papillata Yuncken. MEN.

Ericaceae

Arctostaphylos x cinerea T. J. Howell. DNT; CUR, DOU, JOS.

Arctostaphylos hispidula T. J. Howell. DNT, HUM; CUR, JOS; A-4, B-1b, C, D-3.
Arctostaphylos klamathensis Edwards, Keeler-Wolf, & Knight. SIS; A-1.
Arctostaphylos knightii Gankin & Hildreth. DNT, HUM.

Arctostaphylos manzanita Parry subsp. roofii (Gankin) P. V. Wells. LAK, TEH.

64 MADRONO [Vol. 35

Arctostaphylos stanfordiana Parry subsp. raichei Knight. LAK, MEN.
Arctostaphylos tracyi Eastwood. DNT, HUM, MEN.

Kalmiopsis leachiana (Henderson) Rehder. CUR, DOU, JOS; C.

Rhododendron occidentale (Torrey & A. Gray) var. paludosum Jepson. HUM, DNT.

Euphorbiaceae

Chamaesyce ocellata (Durand & Hilgard) Millspaugh var. rattanii (Watson) Koutnik.
GLE, TEH; A-4.

Fagaceae

Quercus garryana Douglas var. breweri (Engelmann in Watson) Jepson. LAK, HUM,
MEN, SIS, TRI; CUR, JAC, JOS.

Quercus sadleriana R. Brown of Campster. DNT, SIS, TEH; COS, CUR, DOU, JAC,
JOS.

Fumariaceae

Dicentra formosa (Haworth) Walpers subsp. oregana (Eastwood) Munz. DNT, HUM,
SIS, TRI; CUR, JOS; A-4, B-1b, C.

Garryaceae
Garrya buxifolia A. Gray. DNT, HUM, MEN, SIS; CUR, JOS.

Gentianaceae

Gentiana bisetaea T. J. Howell. CUR, JOS; C, D-1.
Gentiana setigera A. Gray. HUM, MEN, SIS, TRI; JAC, JOS; A-3.

Grossulariaceae

Ribes inerme Rydberg var. subarmatum Peck. JAC.
Ribes marshallii Greene. HUM, SIS; JAC, JOS; A-4, B-3, D-2.

Hydrophyllaceae

Phacelia argentea Nelson & Macbride. DNT; COS, CUR; A-1, B-1b, C, D-1.
Phacelia capitata Kruckeberg. COS, DOU, JAC; B-1b, C.

Phacelia cookei Constance & Heckard. SIS; A-1.

Phacelia dalesiana J. T. Howell. SIS, TRI; A-1.

Phacelia greenei J. T. Howell. SIS, TRI; A-1.

Phacelia leonis J. T. Howell. SIS, TRI; JOS; A-3, B-1b, D-2.

Phacelia pringlei A. Gray. SIS, TRI; JAC; A-1.

Labiatae

Monardella purpurea T. J. Howell. DNT, HUM, SIS; CUR, JOS; A-4, B-1b, D-2.
Stachys rigida Nuttall ex Bentham subsp. /anata Epling. DNT, HUM.

Leguminosae

Astragalus agnicidus Barneby. HUM; A-1.

Astragalus rattanii A. Gray var. rattanii. COL, HUM, MEN, LAK, TRI.
Lathyrus biflorus T. W. Nelson & J. P. Nelson. HUM; A-1.

Lathyrus delnorticus C. L. Hitchcock. DNT; COS, CUR, JOS; A-4, B-1b, D-2.
Lathyrus glandulosus Broich. HUM, MEN.

Lathyrus sulfureus Brewer ex A. Gray var. argillaceus Jepson. SHA, TEH.
Lathyrus tracyi Bradshaw. GLE, MEN, SIS, TRI.

1988] SMITH AND SAWYER: ENDEMIC PLANTS 65

Lotus yollabolliensis Munz. HUM, TRI; A-4.

Lupinus antoninus Eastwood. MEN, TEH, TRI; A-1.

Lupinus aridus Douglas ex Lindley subsp. ashlandensis Cox. JAC; B-la, C, D-1.

Lupinus constancei T. W. Nelson & J. P. Nelson. HUM, TRI; A-1.

Lupinus croceus Eastwood var. croceus. SIS, TRI.

Lupinus croceus Eastwood var. pilosellus (Eastwood) Munz. SHA, SIS, TRI, A-4.

Lupinus lapidicola Heller. DNT, SIS; A-4.

Lupinus milo-bakeri C. P. Smith. MEN; A-1.

Lupinus mucronulatus T. J. Howell var. mucronulatus. JOS; B-1b.

Lupinus tracyi Eastwood. DNT, HUM, SIS, TRI; JOS; A-4, B-2b, C, D-2.

Sophora leachiana Peck. CUR, JOS; B-1b, C, D-3.

Trifolium longipes Nuttall subsp. oreganum (T. J. Howell) J. Gillett. HUM, SHA,
ERE JOS:

Trifolium longipes Nuttall subsp. shastense (House) J. Gillett. DNT, SHA, SIS.

Limnanthaceae

Limnanthes bakeri T. J. Howell. MEN; A-1.

Limnanthes floccosa T. J. Howell subsp. bellingeriana (Peck) Arroyo. SHA; JAC;
A-1, B-2b, C, D-1.

Limnanthes floccosa T. J. Howell subsp. grandiflora Arroyo. JAC; B-la, C, D-1.

Limnanthes floccosa T. J. Howell subsp. pumila (T. J. Howell) Arroyo. JAC; B-la,
C, D-1.

Limnanthes gracilis T. J. Howell var. gracilis. DOU, JAC, JOS; B-1b, C, D-1.

Linaceae

Hesperolinon adenophyllum (A. Gray) Small. HUM, LAK, MEN; A-4.
Hesperolinon tehamense H. K. Sharsmith. GLE, TEH.

Malvaceae

Malacothamnus mendocinensis (Eastwood) Kearney. MEN; A-1.

Sidalcea malvaeflora (De Candolle) A. Gray ex Bentham subsp. ce/ata (Jepson) C.
L. Hitchcock. SHA, SIS, TRI.

Sidalcea malvaeflora (De Candolle) A. Gray ex Bentham subsp. e/egans (Greene)
C. L. Hitchcock. DNT, SIS; CUR, JAC, JOS; A-4.

Sidalcea malvaeflora (De Candolle) A. Gray ex Bentham subsp. nana (Jepson) C. L.
Hitchcock. SIS, TEH; JAC, JOS.

Sidalcea malvaeflora (De Candolle) A. Gray ex Bentham subsp. patula C. L. Hitch-
cock. CUR; B-1b, D-2.

Sidalcea oregana (Nuttall ex Torrey & A. Gray) A. Gray subsp. eximia (Greene) C.
L. Hitchcock. HUM, MEN, SIS, TRI; CUR, JAC, JOS.

Sidalcea setosa C. L. Hitchcock subsp. guerceta C. L. Hitchcock. CUR; B-1la, D-1.

Sidalcea setosa C. L. Hitchcock subsp. setosa. SIS; CUR, DOU, JAC, JOS; A-4, C,
D-3.

Nyctaginaceae
Mirabilis greenei Watson. COL, GLE, SHA, SIS, TEH; JAC; D-2.

Onagraceae

Clarkia amoena (Lehmann) Nelson & Macbride subsp. whitneyi (A. Gray) Lewis &
Lewis. HUM, MEN; A-4.

Clarkia borealis E. Small subsp. borealis. SHA, TRI; A-4.

Epilobium canum (Greene) Raven subsp. septentrionale (Keck) Raven. HUM, MEN,
TRI; A-4.

66 MADRONO [Vol. 35

Epilobium nivium Brandegee. COL, GLE, LAK, MEN, TRI; A-1.

Epilobium oreganum Greene. DNT, HUM, SIS, TEH, TRI; DOU, JOS; A-4, B-1b,
C, D-1.

Epilobium rigidum Haussknecht. DNT, SIS; CUR, JAC, JOS; A-4, B-1b, D-2.

Epilobium siskiyouense (Munz) Hoch & Raven. SIS, TRI; JAC; A-1, C, D-2.

Polemoniaceae

Eriastrum brandegeae Mason. COL, GLE, LAK; A-1.

Linanthus harknesii (Curran) Greene subsp. condensatus Mason. GLE; A-1.
Linanthus nuttallii Milliken subsp. howellii Nelson & Patterson. TEH.
Linanthus rattanii (A. Gray) Greene. COL, GLE, LAK, MEN, TEH; A-4.
Navarretia pauciflora Mason. LAK; A-1.

Phlox azurea G. L. Smith. MEN.

Phlox hirsuta E. Nelson. SIS; A-1.

Polygonaceae

Chorizanthe howellii Goodman. MEN; A-1.

Eriogonum alpinum Engelmann. SIS, TRI; A-1.

Eriogonum congdonii (S. Stokes) Reveal. SHA, SIS, TRI; A-4.

Eriogonum diclinum Reveal. SIS; JOS; A-4, B-1b, D-2.

Eriogonum hirtellum J. T. Howell & Bacigalupi. DNT, SIS; A-4.
Eriogonum kelloggti A. Gray. MEN; A-1.

Eriogonum libertini Reveal. SHA, TEH, TRI; A-4.

Eriogonum pendulum Watson. DNT; CUR, JOS; A-2, B-1b, C.
Eriogonum siskiyouense Small. SIS, TRI; A-4.

Eriogonum ternatum T. J. Howell. DNT, SIS, TEH; CUR, JOS; A-4, D-3.
Eriogonum umbellatum Torrey var. speciosum (Drew) S. Stokes. DNT, SIS.

Portulacaceae

Claytonia saxosa Brandegee. HUM, LAK, MEN, SIS.

Lewisia cotyledon (Watson) Robinson in A. Gray subsp. cotyledon. DNT, SIS, TRI;
JAC, JOS.

Lewisia cotyledon (Watson) Robinson in A. Gray subsp. heckneri (Morton) Munz.
SIS, TRI; A-1.

Lewisia cotyledon (Watson) Robinson in A. Gray var. howellii (Watson) Jepson. DNT,
HUM, SIS, TRI; CUR, DOU, JAC, JOS; A-3, C.

Lewisia cotyledon (Watson) Robinson in A. Gray var. purdyi Jepson. CUR, JOS; C,
D-1.

Lewisia oppositifolia (Watson) Robinson in A. Gray. DNT; CUR, JAC, JOS; A-1,
B-1b, C, D-2.

Lewisia stebbinsii Gankin & Hildreth. MEN, TRI; A-1.

Ranunculaceae

Delphinium decorum Fischer & Meyer subsp. tracyi Ewan. COL, GLE, HUM, MEN,
LAK, TEH, TRI; CUR, JAC, JOS.
Ranunculus austro-oreganus Benson. JAC; C, D-3.

Rhamnaceae
Ceanothus pumilus Greene. DNT, HUM, MEN, SIS, TRI; CUR, JAC, JOS.

Rosaceae

Holodiscus discolor (Pursh) Maximowicz var. delnortensis Ley. DNT, SIS, TRI; JOS.
Horkelia bolanderi A. Gray subsp. bolanderi. COL, LAK.

1988] SMITH AND SAWYER: ENDEMIC PLANTS 67

Horkelia daucifolia (Greene) Rydberg subsp. daucifolia. SIS, TEH, TRI; JAC.

Horkelia daucifolia (Greene) Rydberg subsp. /atior Keck. TRI.

Horkelia hendersonii T. J. Howell. JAC; B-1b, C, D-1.

Horkelia sericata Watson. DNT, HUM; CUR, JOS; A-2.

Ivesia pickeringii Torrey ex A. Gray. SIS, TRI; A-1.

Potentilla glandulosa Lindley subsp. globosa Keck. DNT, HUM, SIS; CUR, JAC,
JOS; D-2.

Rubus leucodermis Douglas ex Torrey & A. Gray var. trinitatis Berger. TRI.

Rubiaceae

Galium ambiguum Wight var. siskiyouense Ferris. DNT, HUM, MEN, SIS, TRI;
CUR, DOU, JOS.

Galium glabrescens (Ehrendorfer) Dempster & Ehrendorfer subsp. glabrescens. DNT,
SIS, TRI.

Galium glabrescens (Ehrendorfer) Dempster & Ehrendorfer subsp. josephinense
Dempster & Ehrendorfer. JOS.

Galium serpenticum Dempster subsp. scotticum Dempster & Ehrendorfer. SIS, TRI;
A-1.

Salicaceae

Salix delnorticus C. K. Schneider. DNT; JOS; A-4, B-1b, D-2.
Salix tracyi Ball. DNT, HUM; CUR, JAC, JOS; A-4, B-1b, D-2.

Saxifragaceae

Bensoniella oregona (Abrams & Bacigalupi) Morton. HUM; CUR, DOU, JOS; A-1,
C, D-3.

Heuchera pringlei Rydberg. DNT, SIS.

Saxifraga fragarioides Greene. DNT, HUM, SIS, TRI; CUR, JAC, JOS.

Scrophulariaceae

Antirrhinum subcordatum A. Gray. COL, GLE, LAK, TEH; A-4.

Castilleja brevilobata Piper. DNT, SIS; JOS; A-4.

Castilleja elata Piper. DNT, SIS; CUR, JOS; A-2.

Castilleja mendocinensis (Eastwood) Pennell. MEN; A-1.

Castilleja schizotricha Greenman. SIS; JAC; A-4.

Collinsia linearis A. Gray. DNT, HUM, SIS, TRI; JAC, JOS.

Cordylanthus tenuis A. Gray subsp. pallescens (Pennell) Chuang & Heckard. SIS;
A-l.

Mimulus brachiatus Pennell. LAK; A-3.

Mimulus primuloides Bentham var. linearifolius Grant. SHA, SIS, TRI.

Orthocarpus castillejoides Bentham var. humboldtiensis Keck. HUM; A-1.

Orthocarpus erianthus Bentham var. gratiosus Jepson & Tracy. DNT, HUM, MEN;
CUR, JOS.

Orthocarpus pachystachys A. Gray. SIS; A-1.

Pedicularis howellii A. Gray. SIS; JOS; A-4, B-1b, C, D-3.

Penstemon anguineus Eastwood. DNT, GLE, HUM, MEN, SIS, TRI; CUR, DOU,
JAC, JOS.

Penstemon filiformis (Keck) Keck. SHA, TRI; A-1.

Penstemon newberryi A. Gray subsp. berryi (Eastwood) Keck. DNT, GLE, HUM,
SIS, TRI; CUR, JOS.

Penstemon purpusii Brandegee. COL, GLE, HUM, LAK, MEN, TEH, TRI; A-4.

Penstemon tracyi Keck. SIS, TRI; A-1.

Synthyris missurica (Rafinesque) Pennell subsp. hirsuta Pennell. DOU; B-la, C,
D-1.

Veronica copelandii Eastwood. SIS, TRI; A-4.

68 MADRONO [Vol. 35

Umbelliferae

Eryngium constancei Sheikh. LAK; A-1.

Ligusticum californicum Coulter & Rose. DNT, GLE, HUM, MEN, SIS, TRI.

Lomatium cookii J. S. Kagan. JAC.

Lomatium engelmannii Mathias. MEN, SIS, TRI; CUR, JOS; A-4, B-1b, D-2.

Lomatium howellii (Watson) Jepson. DNT, SIS; CUR, JOS; A-4, B-1b.

Lomatium tracyi Mathias & Constance. HUM, SHA, SIS, TEH, TRI; A-4, B-1b,
D-2.

Perideridia leptocarpa Chuang & Constance. SIS; A-4.

Sanicula peckiana Macbride. DNT; CUR, JOS; A-4.

Sanicula tracyi Shan & Constance. HUM, TEH, TRI; JOS; A-1, B-1b, C.

Tauschia glauca (Coulter & Rose) Mathias & Constance. DNT, HUM, TRI; CUR,
DOU, JAC, JOS; A-4.

Tauschia howellii (Coulter & Rose) Macbride. SIS; CUR, JAC; A-1, B-1b, C, D-1.

Violaceae

Viola lanceolata Linnaeus subsp. occidentalis (A. Gray) Russell. DNT; CUR, DOU,
JOS; A-1, B-1b, C, D-2.
Viola macloskeyi Lloyd subsp. pallens (Banks ex De Candolle) M. S. Baker. SIS.

MAGNOLIOPHYTA: LILIOPSIDA
Gramineae

Calamagrostis foliosa Kearney. DNT, HUM, MEN; A-1.
Lophochlaena californicus Nees var. davyi (L. Benson) Love. LAK, MEN.
Poa piperi Hitchcock. DNT, SIS; CUR, JOS; A-4, B-1b, C, D-2.

Iridaceae

Tris bracteata Watson. DNT; CUR, JOS; A-2.
Tris innominata Henderson. DNT: COS, CUR, DOU, JOS; A-2.
Tris tenax Douglas subsp. Alamathensis Lenz. HUM, SIS; A-4.

Liliaceae

Allium fimbriatum Watson var. purdyi (Eastwood) Ownbey & Aase. COL, LAK;
A-4.

Allium hoffmanii Ownbey. HUM, SHA, TEH, TRI; A-4.

Allium mirabile Henderson. DOU.

Allium siskiyouense Ownbey. DNT, HUM, SIS, TRI; CUR, DOU, JAC, JOS; A-4.

Brodiaea coronaria (Salisbury) Engler subsp. rosea (Greene) Niehaus. LAK, TEH;
A-1.

Calochortus greenei Watson. SHA, SIS; JAC; A-1, B-1b, C, D-1.

Calochortus howellii Watson. DOU, JOS; C, D-1.

Calochortus indecorus Ownbey & Peck. JOS; C, D-1.

Calochortus monanthus Ownbey. SIS; A-1.

Calochortus nudus Watson var. shastensis (Purdy) Jepson. SIS; A-3.

Calochortus persistens Ownbey. SIS; A-1.

Camassia howellii Watson. CUR, JAC, JOS.

Chlorogalum pomeridianum (De Candolle) Kunth var. minus Hoover. TEH.

Dichelostemma ida-maia (Wood) Greene. DNT, HUM, LAK, MEN, SHA, TRI;
CUR, DOU, JOS.

Dichelostemma venustum (Greene) Hoover. DNT, HUM, MEN, SHA, SIS, TRI;
DOU; A-4.

Disporum parvifolium (Watson) Britton. DNT; CUR, JOS.

Erythronium citrinum Watson. DNT, SIS; CUR, JOS; A-4, B-1b.

1988] SMITH AND SAWYER: ENDEMIC PLANTS 69

Erythronium hendersonii Watson. DNT, SIS; JAC, JOS; A-3.

Erythronium howellii Watson. DNT; CUR, JOS; A-4, B-1b, D-2.

Fritillaria adamantina Peck. DOU, JAC; B-1b.

Fritillaria gentneri Gilkey. JAC, JOS; B-1b, C, D-1.

Fritillaria glauca Greene. DNT,. GLE, HUM, LAK, TRI; CUR, DOU, JAC, JOS;
D-2.

Hastingsia atropurpurea Becking. JOS.

Hastingsia bracteosa Watson. JOS; B-1b, C, D-1.

Lilium bolanderi Watson. DNT, HUM, MEN, SIS; CUR, JAC, JOS; A-4, B-1b.

Lilium kelloggii Purdy. DNT, HUM; JOS.

Lilium occidentale Purdy. HUM; COS, CUR; A-1, B-1b, C, D-1.

Lilium vollmeri Eastwood. DNT, HUM, SIS; CUR, JAC, JOS; A-3, C.

Lilium wigginsii Beane & Vollmer. DNT, SIS; JAC; A-3, B-1b, C.

Trillium ovatum Pursh subsp. oettingeri Munz & Thorne. SHA, SIS, TRI; A-4.

Trillium rivale Watson. DNT, SIS; CUR, DOU, JOS; A-4.

Triteleia crocea Greene var. crocea. DNT, SHA, SIS, TRI; CUR, JAC, JOS; A-4.

Triteleia crocea Greene var. modesta (Hall) Hoover. SHA, SIS, TRI; A-4.

Triteleia hendersonii Greene var. leachiae (Peck) Hoover. CUR; D-2.

Smilacaceae
Smilax jamesii Wallace. DNT, SHA, SIS, TRI; A-4.

ANNOUNCEMENT

NEw PUBLICATION

WALTERS, D. R. and D. J. Kett. 1988. Vascular plant taxonomy, 3rd
ed., Kendall/Hunt Publishing Co., Dubuque, Iowa, 1988, 488 pp.,
illus., ISBN 0-8403-4614-X, $39.95 (paperbound). [Text for intro-
ductory level taxonomy classes, completely rewritten and expanded
from 2nd edition. Organized in four sections: Part I, Basics of Intro-
ductory Taxonomy, includes nomenclature, vegetative terminology,
key construction, introduction to manuals and floras, and plant col-
lecting. Part II, Survey of Vascular Plant Families, includes chapters
on ferns and fern allies, gymnosperms, and eleven chapters on an-
glosperms organized according to Cronquist’s 1981 system of clas-
sification. Families receiving greatest emphasis are illustrated with
original line drawings, floral diagrams, and floral formulas. Part III,
Approaches to Classification, briefly surveys character variation, ar-
tificial and phenetic systems, traditional phylogenetic systems, and
cladistics. Part IV, Gathering and Analysis of Data, examines exper-
imental taxonomy and the preparation of revisions and monographs.
The book includes a key to many but not all plant families and a
detailed glossary.]

NOTES

NOMENCLATURE OF Lomatium nuttallii, L. kingii, AND L. megarrhizum (API-
ACEAE).— The recent new combination of Lomatium kingii by Cronquist (Great Basin
Nat. 46:254, 1986) prompted a routine review of the nomenclature of Lomatium
nuttallii (Gray) Macbr., L. megarrhizum (A. Nels.) Mathias, and L. kingii (Wats.)
Cronq. in preparation by the senior author for an upcoming revision of Manual of
the Vascular Plants of Wyoming. The oldest name in the group is Seseli nuttallii Gray
(Proc. Amer. Acad. Arts 8:287, 1870). Gray cited two collections (syntypes) in his
original description, one from the Rocky Mountains (Nuttall s.n.), and the other from
the Huerfano Mountains, New Mexico (actually Colorado—see Rhodora 60:265-
271, 1958) (Parry 83). The Nuttall collection is what has recently been called L.
megarrhizum. The Parry collection is the holotype of Neoparrya lithophila Mathias.

It is first necessary to typify the name Seseli nuttallii, because current usage is
incorrect. Gray’s original description included both the Nuttall and Parry collections
so both have equal standing for a lectotype. Watson (Proc. Amer. Acad. Arts 22:474,
1887) was the first to deal with Gray’s name when he placed the Nuttall collection
cited by Gray into synonymy with Peucedanum kingii Wats., a new name for P.
graveolens Wats. that was reputedly a later homonym. (P. nuttallii was preoccupied
so a transfer could not be made.) The type of P. graveolens, and therefore of P. kingii,
is not the same taxon as the Nuttall collection, however. Watson was not sure what
the Parry collection was. We will return to the Watson names later.

Coulter and Rose (Revision of North American Umbelliferae 71, 1888) were next
to deal with the problem. They used the name Peucedanum kingii Wats., with P.
graveolens Wats. and Seseli nuttallii Gray “in part” in synonymy. Seseli nuttallii was
not treated elsewhere in their paper. This is no change from Watson’s treatment. In
their 1900 revision (Contr. U.S. Natl. Herb. 7:245) they used the name Cynoma-
rathrum nuttallii (Gray) C. & R. with the following in synonymy: Seseli nuttailii Gray,
Peucedanum graveolens Wats., P. kingii Wats., and P. megarrhiza A. Nels. The epithet
“nuttalli’’” was not preoccupied in Cynomarathrum as it was in Peucedanum. The
type locality was given as “‘Rocky Mountains; collected by Nuttall.’ Here is the
first designation of a lectotype. Subsequent workers (Mathias, Ann. Missouri Bot.
Gard. 25:225-297, 1938; Mathias and Constance, North Amer. Fl. 28B(2):161-295,
1945) have followed this designation. Furthermore, Mathias’ use (Ann. Missouri Bot.
Gard. 16:393-398, 1929) of the Parry specimen as the holotype of Neoparrya lith-
ophila left the Nuttall specimen the only remaining element of Gray’s Seseli nuttallii.
The Nuttall specimen then typifies Sese/i nuttallii Gray, and Lomatium nuttallii (Gray)
Macbr. becomes the correct name for the species on barren clay hills and flats of
Nebraska, Wyoming, and Colorado that previously was known as L. megarrhizum
(A. Nels.) Mathias.

Now we must deal with Watson’s names in Peucedanum. Watson first used the
name Peucedanum graveolens (Watson in King, Rep. Geol. Explor. 40th Parallel 5:
128, 1871), the holotype being Watson 463 from the Wasatch Mountains of Utah at
around 9000 feet. He later discovered that Bentham and Hooker had apparently used
the same name in 1867 (Genera Plantarum 1:919) fora different taxon, so he proposed
the new name of P. kingii (Proc. Amer. Acad. Arts 22:474, 1887) for his species. In
examining the Bentham and Hooker publication, we discovered that the name Peu-
cedanum graveolens had in fact not been used by them. They simply listed A. gra-
veolens (Anethum) under the genus Peucedanum, which is not a valid transfer. The
International Code of Botanical Nomenclature is very clear on this point as it has an
example (Article 33.1, Ex. 2) from this very same publication. Peucedanum graveolens
Wats., therefore, is legitimate, and P. kingii Wats. is superfluous. Peucedanum gra-

MApbrRONO, Vol. 35, No. 1, pp. 70-71, 1988

1988] NOTES 71

veolens Wats. is the oldest name for the species that has been called Lomatium nuttallii
and, therefore, must be transferred.
The nomenclature of the entire group follows.

LOMATIUM NUTTALLI (Gray) Macbr., Contr. Gray Herb. 56:35. 1918.—Seseli nuttallii
Gray, Proc. Amer. Acad. Arts 8:287. 1870.—Cynomarathrum nuttallii (Gray)
C. & R., Contr. U.S. Natl. Herb. 7:245. 1900.— Cogswellia nuttallii (Gray) Jones,
Contr. W. Bot. 12:32. 1908.—Aletes nuttallii (Gray) Weber, Phytologia 55:6.
1984.—LEcTOTYPE by Coulter and Rose (Contr. U.S. Natl. Herb. 7:245. 1900):
Rocky Mountains, Nuttall s.n. (GH!, photo UC!; isolectotype: NY!, photo RM!).

Peucedanum megarrhiza A. Nels., Bull. Torrey Bot. Club 26:130. 1899.—Cynoma-
rathrum megarrhizum (A. Nels.) Rydb., Flora Rocky Mountains 629, 1064.
1917.—Lomatium megarrhizum (A. Nels.) Mathias, Ann. Missouri Bot. Gard.
25: 282. 1938, issued 1937.— Neoparrya megarrhiza(A. Nels.) Weber, Phytologia
41:487. 1979.—Aletes megarrhiza (A. Nels.) Weber, Phytologia 55:6. 1984.—
LECTOTYPE by Mathias (Ann. Missouri Bot. Gard. 25:282. 1938, issued 1937):
Wyoming, Point of Rocks, Nelson 4769 (RM!; isolectotype: GH!, MO!, NY,
photo RM!, US).

Lomatium graveolens (Wats.) Dorn & Hartman, comb. nov. — Peucedanum graveolens
Wats. in King, Rep. Geol. Explor. 40th Parallel 5:128. 1871.—Peucedanum kingii
Wats., Proc. Amer. Acad. Arts 22:474. 1887 (nomen superfl.).—Lomatium kingii
(Wats.) Cronq. [in Welsh], Great Basin Nat. 46:254. 1986.—HOLOoTYPE: Utah,
Wasatch (as Wahsatch) Mountains, Watson 463 (US, photo RM!; isotype: NY!,
photo RM!).

LOMATIUM GRAVEOLENS var. alpinum (Wats.) Dorn & Hartman, comb. nov. —Peu-
cedanum graveolens var. alpinum Wats. in King, Rep. Geol. Explor. 40th Parallel
5:129. 1871.—Peucedanum kingii var. alpina (Wats.) C. & R., Revision of North
American Umbelliferae 71. 1888.—Cynomarathrum alpinum (Wats.) C. & R.,
Contr. U.S. Natl. Herb. 7:245. 1900.—Cogswellia nuttallii var. alpina (Wats.)
Jones, Contr. W. Bot. 12:32. 1908.—Lomatium alpinum (Wats.) Macbr., Contr.
Gray Herb. 56:35. 1918.—Lomatium nuttallii var. alpinum (Wats.) Mathias,
Ann. Missouri Bot. Gard. 25:279. 1938, issued 1937.—Lomatium kingii var.
alpinum (Wats.) Cronq. [in Welsh], Great Basin Nat. 46:255. 1986.— HOLOTYPE:
Nevada, East Humboldt Mountains, Watson 464 (US, photo RM!; isotypes:
GH!, NY!, photo RM!).

NEOPARRYA LITHOPHILA Mathias, Ann. Missouri Bot. Gard. 16:393. 1929.—Aletes
lithophila (Mathias) Weber, Phytologia 55:5. 1984.—HoLotypPe: Colorado (as
New Mexico), Huerfano (as Huefano) Mountains, Parry 83 (GH, photo Ann.
Missouri Bot. Gard. 16:pl. 33, after p. 398. 1929!; isotype: MO!).

—RosBeERT D. Dorn, Box 1471, Cheyenne, WY 82003 and RONALD L. HARTMAN,
Rocky Mountain Herbarium, Dept. Botany, Univ. Wyoming, Laramie 82071-3165.
(Received 27 Feb 1987; revision accepted 5 Oct 1987.)

NOTEWORTHY COLLECTIONS

COLORADO

ANTENNARIA AROMATICA Evert (ASTERACEAE).— Gunnison Co.: Cottonwood Pass,
about 0.5 mis. of hwy., rocky slopes, mostly nw.-facing fell fields, T14S R81W S14,
3877 m, 1 Aug 1984, Bayer et al. CO-449 (RM); Cumberland Pass, rt. 765, T51N
R4E, 1 Aug 1984, Bayer et al. CO-441 (RM); sw. summit of Galena Peak, T12S
R87W S11, 3749 m, 31 Jul 1984, Bayer et al. CO-435 (RM). Summit Co.: Hoosier
Pass area, 0.25 mis. of Hoosier Pass, T8S R78W S10, 2 Aug 1984, Bayer et al. CO-
458 (RM), (verified by R. Bayer). Pitkin Co.: Taylor Pass, Jul 1986, K. Matthews s.n.
(verified by W. A. Weber). Lake Co.: Sawatch Mts., Mt. Champion Basin, fell field
adj. to old mine, granitic substrate, 3751 m, 20 Aug 1986, Hartman and Rottman
6671/2942 (COLO, CU—Denver) (verified by W. A. Weber). In CO, the species
occurs 1n fell fields of calcareous rocks in the alpine.

Significance. First records for CO and a range extension of ca. 610 km sse. of the
nearest localities at 21 km sw. and 8 km w. of Cody, Park Co., WY (Evert, Madrono
31:109-112, 1984). The species also is known from mountainous areas of w. central
(Cascade Co.) and sw. (Gallatin and Carbon cos.) MT (verified by W. A. Weber).—
STEVE L. O’KANE, JR., Colorado Natural Areas Program, 1313 Sherman St., Room
718, Denver, CO 80203; EmMity L. HARTMAN and Mary Lou ROTTMAN, Biology
Dept., Univ. Colorado, Denver 80202.

ARALIA RACEMOSA L. (ARALIACEAE).—La Plata Co.: Elbert Ck, 1.2 mi sw. of con-
fluence of Sawmill Creek and the Animas River, T38N R8W S31, 2290 m, 26 Jul
1986, W. Baker and D. Paulson s.n. (COLO, CS).

Significance. First record for CO, a range extension of ca. 150 km n. from the
nearest locality in Rio Arriba Co., NM.

ASTRAGALUS HUMILLIMUS A. Gray ex Brand. (FABACEAE). — Montezuma Co.: n. rim
of unnamed mesa between Tanner Mesa and Short Mesa, on exfoliating Point Lookout
sandstone, T33N R17W S22, 1859 m, 14 May 1986, O’Kane 2342 (COLO).

Significance. First CO record of this federally endangered species (Fed. Reg. 50:
26568-26572, 1985) since the type collection was made in 1875. This collection came
from a population growing under conditions similar to those reported by Brandegee
(Bull. U.S. Geol. Surv. Terr. 2:235, 1876) as “growing upon sandstone rock of the
Mesa Verde, near the edge of Mancos Canyon.” The nearest known locality is 31 km
se. in San Juan Co., NM.

ASTRAGALUS SERICOLEUCUS A. Gray (FABACEAE). — Chaffee Co.: Harrington Gulch,
w. of Salida and above and s. of Adobe Park, barren hills of alluvium with Pinus
and Cercocarpus, TSON R8E $35, 2225 m, | Aug 1985, O’Kane and Anderson 2217
(NY) (verified by R. C. Barneby).

Significance. Range extension of 260 km w. of the nearest locality on the plains in
Lincoln Co. (Barneby, Memoirs N.Y. Bot. Gard. 13:1144-1146, 1964). Two rare
endemics restricted to sedimentary strata in the Canon City area also occur at this
site. These species, Eriogonum brandegei Rydb. (Reveal, A revision of the genus
Eriogonum. Ph.D. diss., Brigham Young Univ., 1969) and Parthenium tetraneuris
Barneby (Rollins, Contr. Gray Herbarium 172:1—72, 1950), grow at least 440 m
higher than at locations below the mouth of the Canyon of the Arkansas River. Also,
Neoparrya lithophila Mathias grows here on a sedimentary, rather than igneous sub-
strate.

MADRONO, Vol. 35, No. 1, pp. 72-74, 1988

1988] NOTEWORTHY COLLECTIONS Us

ATRIPLEX PLEIANTHA W. A. Weber (CHENOPODIACEAE).— Montezuma Co.: 2.2 mi
due e. of jct. of Hwys. 160 and 41 in gray clays derived from Mancos shale, T32N
R19W S8, 1493 m, 8 May 1985, O’Kane, Anderson and Fleming 2022 (CS).

Significance. Re-collection of type locality and first collection from CO since 1949
(Weber 4788, COLO). This species is a candidate for federal listing as endangered or
threatened (Fed. Reg. 50:39534, 1985).

CREPIS CAPILLARIS (L.) Wallr. (ASTERACEAE). — Larimer Co.: Dixon Reservoir, | mi
se. of Dixon Canyon Dam, T7N R69W S829, 1585 m, 8 Aug 1986, D. Wilken 14676
(COLO, CS, RM).

Significance. First record for CO, adventive and presumably a range extension w.
from the eastern Great Plains (Barkley, Asteraceae, Jn Flora of the Great Plains,
Univ. Kansas Press, 1986).

CRYPTANTHA WEBERI I. M. Johnston (BORAGINACEAE).— Conejos Co.: “Flat Top”
Mountain in San Luis Hills, sides of mesa in dark volcanic rocks with Artemisia,
Bouteloua, and Pinus, T34N R1I1E S8, 2682 m, 10 Jul 1986, O’Kane and Anderson
2503 (CS).

Significance. Range extension of 111 km se. from a small area of endemism in
Saguache and Hinsdale cos. (Higgins, Brigham Young Univ. Science Bull. 13:1-63,
197):

DITHYREA WIZLIZENII Engelm. (BRASSICACEAE). — Montezuma Co.: Along Cowboy
Wash, 2 mie. of Utah State Line, T32N R20W, 27 Apr 1985, Fleming s.n. (SJNM).

Significance. First modern collection of the species from CO. Some doubt exists
whether Brandegee’s report from the “‘valley of the San Juan” (Bull. U.S. Geol. Surv.
Terr. 2:233, 1876) was actually from CO or UT (Weber, Univ. Colo. Studies, Biology
2377, 1966).

IPOMOPSIS CONGESTA (Hook.) V. Grant subsp. CREBRIFOLIA (Nutt.) Day (POLEMONIA-
CEAE).— La Plata Co.: 0.3 mi ne. of La Boca on Shellhammer Ridge, T32N R7W S15,
1 May 1985, O’Kane 85-39 (CS).

Significance. First record for CO. Previously known from sw. MT and n. WY to
NM and UT (Cronquist et al., Intermountain Flora 4:128, 1985) and recently reported
from NV (Tiehm, Madrono 33:228, 1986). Range extension of ca. 200 km n. of
Sandoval Co., NM and ca. 180 km e. of San Juan Co., UT.

LOMATIUM BICOLOR (S. Wats.) Coult. & Rose var. BICOLOR (APIACEAE). — Gunnison
Co.: Snowshoe Mesa, abundant on clay loam with Wyethia sp., sagebrush, and oak,
T13-T14S R89W, 2590 m, 18 Jul 1938, F. E. Read R-406 (USFS) (verified by M.
Schlessman).

Significance. First record for CO; a range extension of ca. 430 km to se. from Salt
Lake Co., UT and Lincoln Co., WY. Variety /eptocarpum (Nutt. ex Torr. & Gray)
Schlessman is known from Gunnison Co. (Schlessman, Syst. Bot. Monogr. 4:26-28,
1984).

MENTZELIA DENSA Greene (LOASACEAE).— Fremont Co.: 2.0 road mie. of Cotopaxi
on Hwy. 50, Arkansas River Canyon, T48N R12ES29, 1951 m, 31 Jul 1985, O’Kane
and Anderson 2204 (CS).

Significance. Relocation of the most recent collection of the species (H. Thompson
1684, LA, US) made in 1955. The species is limited to the Arkansas River Canyon
from Canon City to Cotopaxi. Darlington (Annals Mo. Bot. Gard. 21:157—158, 1934)
enigmatically reports the species from “‘southern Colorado” with specimens from
**Mesa County.” Specimens examined by Darlington are probably best ascribed to
M. multiflora (Nutt.) Gray. Darlington does not indicate that the species is found in
the Arkansas Canyon, although Greene (Pittonia 3:99, 1896) states that it is ““com-
mon” here.

74 MADRONO [Vol. 35

NEOPARRYA LITHOPHILA Mathias (APIACEAE). — Chaffee Co.: on county road 111A,
ca. 0.8 km s. of Salida, T49N R9E S7, 2217 m, 2 Aug 1985, O’Kane and Anderson
2218 (COLO, CS), Anderson 85-110 (RM). Conejos Co.: Flat Top, San Luis Hills,
T34N RIIE S8, 2682 m, 10 Jul 1986, O’Kane, Anderson, and Dixon 2500 (COLO,
CS). Rio Grande Co.: Elephant Rocks, 8 km ne. of Del Norte, T40N R6E S3, 2423
m, 25 Jul 1984, J. Anderson s.n. (RM), 24 Jul 1985, B. C. Johnston 3038 (RM), 3041
(COLO, RM); T40N ROE S4, 24 Jul 1985, Johnston 3051 (COLO, RM). Saguache
Co.: Middle Creek, 3048 m, 10 Jul 1922, C. E. Taylor 475 (USFS); ca. 0.8 air km
w. of Upper Saguache Forest Service Station, T45N RSE S19, 2621 m, 18 Sep 1983,
Hartman 17350 (COLO, CS, RM); Upper Saguache Station, 2621 m, 6 Jul 1922,
Taylor 476 (USFS); road to Jacks Creek, 0.8 km nw. of jet. of Hwy. 114, T45N R6E
S10, 25 Jul 1985, Johnston 3062 (RM); road to Middle Creek, 2.4 km nnw. of Hwy.
114, T45N R6E S4, 25 Jul 1985, Johnston 3063 (RM); Forest road 660, ca. 2.4 km
w. of jet. with Del Norte-—La Garita road, T41N R6E S21, 2500 m, 8 Aug 1985,
O’Kane and Anderson 2241 (CS); hill along Cottonwood Creek, ca. 3.2 km w. of Rio
Grande Canal and ca. 8.9 km sw. of Swede Corners, T43N R7E $32, 2408 m, 9 Aug
1985, O’Kane 2245 (BRY, CS). Locally occasional to abundant on hills, benches, cliff
faces, and boulder fields of Tertiary volcanics with Artemisia, Bouteloua, Chryso-
thamnus, Eriogonum, Hymenoxys, Muhlenbergia, Oryzopsis, Pinus ponderosa, Pseu-
dotsuga, Ribes, and Symphoricarpos. O’Kane and Anderson 2218 is unusual in being
abundant on barren, near-white, silt-loam alluvium of the Dry Union Formation.

Significance. Range extension of 82 km sw. or 110-130 km w. to nw. of the only
previously published locality (type locality?) at Silver Mt. (as Dike Mt., Weber, Rho-
dora 60:265-271, 1958), in w. Huerfano Co., CO. The two collections by Taylor,
both in late flowering and early fruiting condition, were filed under Pseudocymopterus
anisatus (A. Gray) Coult. & Rose when discovered by RLH in 1983. Neoparrya
lithophila is a candidate for federal listing as endangered or threatened (Fed. Reg. 50:
39584, 1985), because repeated visits by several workers to Silver Mt. and adjacent
areas indicated that it was restricted in distribution. The new records show it scattered
along the eastern margin of the San Juan Volcanic Area, a region ca. 15,000 km? in
extent and composed of basalts and other volcanics deposited during the early Ter-
tiary. The known altitudinal range is now from ca. 2130 m (Silver Mt. locality) to
ca. 3048 m. Theobald, Tseng, and Mathias (Brittonia 16:296-315, 1964) note in the
species description: “rays... reflexed in flower and fruit... pedicels. . . reflexed in
fruit.” A study of material from all known localities shows that the compound umbels
are rounded in early flower with erect to spreading rays (and pedicels), the outer of
which become reflexed only with age, thereby often leading to a spherical infructes-
cence.

RUMEX VERTICILLATUS L. (POLYGONACEAE).— Weld Co.: near county road 114, 0.5
mie. of U.S. Hwy. 85, 7 min. of Nunn, 1650 m, 22 Aug 1986, D. Hazlett 7527
(CS).

Significance. First record for CO; naturalized and presumably a range extension
from e. Kansas (R. Kaul, Polygonaceae, /n Flora of the Great Plains, Univ. Kansas
Press, 1986).—STEVE L. O’KANE, JR., Colorado Natural Areas Program, 1313 Sher-
man St., Room 718, Denver 80203; DIETER H. WILKEN, Dept. Botany, Colorado
State Univ., Ft. Collins 80523; and RONALD L. HARTMAN, Rocky Mountain Her-
barium, Univ. Wyoming, Laramie 82071-3165.

REVIEWS

Serpentine and Its Vegetation: A Multidisciplinary Approach. By ROBERT RICHARD
Brooks. 449 pp. Ecology, Phytogeography & Physiology Series Volume 1. T. R.

MApDRONO, Vol. 35, No. 1, pp. 74-76, 1988

1988] REVIEWS ie:

DuDLEY, General Editor. Dioscorides Press, Portland, OR. 1987. Hardbound. $47.50.
ISBN 0-931146-04-6.

For the dedicated student of plant “‘serpentine’’ soil endemism, as well as the casual
observer, a book such as this one has long been awaited. Because most botanists lack
a strong background in geochemistry and geology, few take the time to wade through
the appropriate literature in those fields to develop the needed understanding of the
ultramafic environment. Robert Brooks has provided an excellent account on the
nature of ultramafic rocks and their derived serpentine soils. The first three chapters
outline the geochemistry of ultramafic minerals and their derived soils. Chapters 4—
6 review the major works on various aspects of plant endemism on serpentine soils
including heavy metal accumulation and nutrient imbalances such as calcium and
magnesium. The chapter on plant evolution and serpentine is brief and primarily
discusses the evolution of plant groups from a global perspective with little infor-
mation at the species or population level. Dr. Brooks’ expertise and personal bias
appears in Chapter 8 with an extensive discussion on plant hyperaccumulation of
nickel.

The remaining three quarters of the book are dedicated to the serpentine vegetation
of the world. There are eleven vegetation chapters beginning with North America
and also including tropical America, northwest Europe, central and southern Europe,
continental Asia, Japan, Africa, the Malay Archipelago, New Caledonia, Australia,
and New Zealand. Each chapter includes the region’s geology, vegetation, and im-
portant botanical studies. The book 1s well-illustrated with maps, tables, graphs, and
black-and-white as well as color photographs. The photographs have been reproduced
very well and clearly illustrate many interesting plants and places. There are three
indices: a subject index, a geographical index, and a botanical index. The botanical
index is excellent and includes 2,219 species, subspecies, and varieties of vascular
plants, mosses, and lichens. This book has brought together a considerable amount
of information; most notable are the many international journal articles. This work
will undoubtedly be the main reference source for the serpentine plant literature for
some time to come. — NIALL F. MCCARTEN, Dept. Biology, San Francisco State Uni-
versity, San Francisco, CA 94132.

Atlas Cultural de México. Flora. By JERZY RZEDOWSKI and MIGUEL EQUIHUA. 223
pp. Secretaria de Educacion Publica, Instituto Nacional de Antropologia e Histoira,
Grupo Editorial Planeta. 1987. $8400 (pesos).

This volume comprises part of an Atlas Cultural series. Three other volumes have
been published to date: Archeology, Tourism, Handicrafts.

The history of Mexico shows that the Mayans, Toltecans, and other early civili-
zations had a tremendous understanding and appreciation of plants and were suc-
cessful in cultivating many of them. Urbanization in modern times has resulted in
loss of much of this early knowledge and appreciation. It is the authors’ hope that
this volume will stimulate local peoples’ interest in the plants occurring in the many
varied habitats of Mexico as well as to enable visitors to become familiar with many
of the plants.

The 621 colored photographs presented in the volume constitute less than 2 percent
of the flora of Mexico, but they give an excellent idea of the diversity of the native
plants in Mexico. Chapters are arranged under nine vegetational categories: 1) Bosque
tropical perennifolio, 2) Bosque tropical subcaducifolio, 3) Bosque tropical caduci-
folio, 4) Bosque espinoso, 5) Matorral xerofilo, 6) Pastizal, 7) Bosque de coniferas y
de encinos, 8) Bosque mesofila de montana, and 9) VegetaciOn aquatica y subaquatica.
Two maps serve to illustrate this classification. In addition, five more general groups
are presented in separate chapters: 10) Algunos otros tipos de vegetacion, como la
costera y los palmares, 11) Las malezas, 12) Las plantas del hombre, su historia en
México, 13) Plantas que caracterizan espicificamente la flora mexicana, and 14) La
flora patrimonio de México y del mundo. For each plant illustrated there is a brief
description, general distribution, common name, flowering period, uses, and the

16 MADRONO [Vol. 35

highway routes (as shown on an introductory map) where one might expect to see it.
Each of these chapters is prefaced with a brief discussion characterizing the vegeta-
tional type. Where appropriate, mention is included of man’s impact on the area.
That preceding Las Plantas del Hombre tells of the beginning of agriculture, domes-
tication of vegetables and precolumbian agriculture in Mexico. The final two chapters
stress the distinctiveness and beauty of the Mexican flora and the importance of
conservation and of rational use of land rather than its despoilment. Within each
chapter there is neither taxonomic nor alphabetical arrangement of the entries, but
rather there are pleasing groupings of the many photographs presented. In such a
book, no formal taxonomic arrangement would be practical. The volume ends with
a two page glossary and an incomplete index to common names with their scientific
equivalents.

The problem of common names, as discussed in the Introduction, is well illustrated
in the chapter “El Matorral XerOofilo,” the most abundant vegetation type in Baja
California. The common name given in the Flora for Olneya tesota (p. 75) is palo
fierro, a name applied to that tree in parts of Sonora, but not in southern Baja
California where it is aptly called u/fia de gato (cat’s claw), and where palo fierro is
applied to the southern peninsular endemic Prosopis palmeri. However, in California
and Arizona, the translation “‘ironwood” refers to Olneya tesota. To further confuse
the matter, in northern Baja California, California and Arizona, ufia de gato refers
to Acacia greggii. Other examples of common name problems in this chapter are:
colorin (p. 74) which in Baja California refers to Erythrina flabelliformis; pitaya agria
(p. 80) always refers to Machaerocereus (Stenocereus) gummosus in Baja California
and never to Lophocereus schottii; Palo verde (p. 82) might be considered a “‘generic”’
common name for Cercidium, but in Baja California there are four taxa in Cercidium,
each with its own name: dipua for C. microphyllum, palo brea for C. praecox, palo
estribo for C. sonorae, and palo verde for C. floridum subsp. peninsulare. Torote (p.
82) is usually applied to species of Bursera whereas Jatropha cuneata is known as
matacora. Space limitations in the F/ora make it impossible, however, to detail such
geographic variation in application of common names.

This Flora presents an excellent “overview” of the vegetation of Mexico. It merits
wide distribution within Mexico and should be readily available to those visiting our
neighbor to the south. At the present value of the peso, it is practically a “‘give-away”’.
It is to be hoped that some adjustment can be made. — ANNETTA CARTER, Herbarium,
Department of Botany, University of California, Berkeley 94720.

Volume 35, Number 1, pages 1-76, published 12 April 1988

ANNOUNCEMENT

FOURTH ANNUAL SOUTHWESTERN BOTANICAL SYSTEMATICS SYMPOSIUM

‘*Adaptation and Evolution in Arid Areas”

For information write to: Rancho Santa Ana Botanic Garden, Bo-
tanical Systematics Symposium, 1500 N. College Ave., Claremont, CA
91711; phone (714) 625-8767. Date: 20-21 May 1988.

SUBSCRIPTIONS — MEMBERSHIP

Membership in the California Botanical Society is open to individuals ($22 per year;
students $12 per year fora maximum of seven years). Members of the Society receive
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STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION
(Required by Title 39, U.S.C. 3685)

MADRONO, A West American Journal of Botany, is published quarterly at Berkeley,
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The Publisher is the California Botanical Society, Inc., Life Sciences Building,
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The editor is David J. Keil, Biological Sciences Department, California Polytechnic
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17 February 1988 DAVID J. KEIL, Editor

PF ve 35, NUMBER 2 | APRIL-JUNE 1988
99 MiG

Bot -
NT” p

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Contents

| Review oF Erigeron eatonii AND ALLIED TAXA (COMPOSITAE A
John L. Strother and William J. Ferlatte fa

_A COMPARISON OF NORTH AND SOUTH AMERICAN Lupinonour Microchrpiy | (989 7
| (LEGUMINOSAE) }
| Rhonda Riggins aN - 92

_Leptodactylon pungens sussp. hazeliae (POLEMONIACEAE), A “NEW _ COMBINATION FOR A A

nc elie

SNAKE RIVER CANYON ENDEMIC Dreamer

Robert J. Meinke 105
| Scutellaria lutilabia (LABIATAE), A NEW GYPSOPHILE FROM NUEVO LEON, MEXICO

Thomas M. Lane and Guy L. Nesom 112
_A NEw SPECIES OF Croton (EUPHORBIACEAE) FROM NICARAGUA

Grady L. Webster 7

A New Lomatium (APIACEAE) FROM THE SIERRAN CREST OF CALIFORNIA
Ronald L. Hartman and Lincoln Constance 121

A New SPEcIES OF Saxifraga (SAXIFRAGACEAE) FROM THE OLYMPIC MOUNTAINS,
WASHINGTON, AND VANCOUVER ISLAND, BRITISH COLUMBIA
Rick J. Skelly 126

| EVIDENCE FOR A WARM Dry EARLY HOLOCENE IN THE WESTERN SIERRA NEVADA OF
CALIFORNIA: POLLEN AND PLANT MACROFOSSIL ANALYSIS OF DINKEY AND
EXCHEQUER MEADOWS

_ Owen K. Davis and Michael J. Moratto 132
SPECIES FREQUENCY IN RELATION TO TIMBER HARVEST METHODS AND ELEVATION IN

| THE PINE TYPE OF NORTHEAST CALIFORNIA

Robin S. Vora 150
NOTES
| REPORT ON THE XIV INTERNATIONAL BOTANICAL CONGRESS
_ Paul C. Silva 159
TYPIFICATION OF Chaenactis alpina (ASTERACEAE)

Robert D. Dorn 161
Chenopodium simplex, AN OLDER NAME FOR C. gigantospermum (CHENOPODIACEAE)
| Robert D. Dorn 162
Arabis breweri S. WATS. VAR. austinae (GREENE) ROLL. (CRUCIFERAE)
_ Robert E. Preston 162

|
NOTEWORTHY COLLECTIONS

CALIFORNIA 164
_ NeEvADA 166
| OREGON 167
‘ANNOUNCEMENTS 111, 125, 149, 158
REVIEWS 167

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
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Editor—Davip J. KEIL

Biological Sciences Department
California Polytechnic State University
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Board of Editors
Class of:

1988—SusAn G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989— FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991—JAMES HENRICKSON, California State University, Los Angeles
WAYNE R. FERREN, JR., University of California, Santa Barbara

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1987-88

President: DALE MCNEAL, Department of Biological Sciences, University of the
Pacific, Stockton, CA 95211

First Vice President: PEGGY FIEDLER, Department of Biology, San Francisco State
University, 1600 Holloway Ave., San Francisco, CA 94132

Second Vice President: STEVEN TIMBROOK, Ganna Walska Lotusland Foundation,
695 Ashley Rd., Montecito, CA 93108

Recording Secretary: V.THOMAS PARKER, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
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The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, FRANK ALMEDA, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118; the Editor of MADRONO; three elected
Council Members: ANNETTA CARTER, Department of Botany, University of Califor-
nia, Berkeley, CA 94720; JoHN MoorInc, Department of Biology, University of
Santa Clara, Santa Clara, CA 95053; BARBARA ERTTER, University Herbarium, De-
partment of Botany, University of California, Berkeley, CA 94720; and a Graduate
Student Representative, NIALL F. MCCARTEN, Department of Biological Sciences, San
Francisco State University, San Francisco, CA 94132.

REVIEW OF ERIGERON EATONIT AND ALLIED TAXA
(COMPOSITAE: ASTEREAE)

JOHN L. STROTHER
Herbarium, University of California, Berkeley 94720

WILLIAM J. FERLATTE
Siskiyou County Department of Agriculture,
525 South Foothill Drive, Yreka, CA 96097

ABSTRACT

Review of morphological attributes and geographic distributions has led to revised
circumscriptions for Erigeron eatonii and allied taxa, including recognition of one
new taxon (Erigeron eatonii var. lavandulus) and re-establishment of an older one
(E. sonnel).

‘““Crazy-quilt’’ aptly describes the confounding geographic pattern
of sometimes marked and, more often, subtle morphological vari-
ants that are subsumed by the phrase “‘Erigeron eatonii and allied
taxa’ (Figs. 1 and 2). Distributions of some of the morphs seem to
be strictly determined by particular, often discontinuous substrates;
other morphs are more eclectic in choice of habitat. Although pollen
grains show quite a range of volumes (6371 to 15,448 cubic mi-
crometers), all chromosome counts for the group are from diploid
(2n = 18) plants (Solbrig et al. 1969, Keil and Pinkava 1976, Nesom
1978, and counts reported here). [A report of 2n = 36, as n = 18,
for E. eatonii var. eatonii came from a plant referable to E. tener
A. Gray (NV, White Pine Co., Breedlove 5815, DS, DUKE).] Because
local populations often show considerable internal uniformity, but
vary from one to another, we suspect that apomixis may be con-
tributing to the complexity in a manner similar to that documented
by Beaman (1957) for some species of Townsendia and by Noyes et
al. (1987) in Erigeron compositus. All of the taxa treated here seem
to be very closely related; they variously intergrade morphologically
and may constitute a single, polymorphic species. Overall, the com-
plex pattern of morphology and geography is similar in some ways
to that of Galium multiflorum Nutt. (cf. Dempster and Ehrendorfer
1965, Cronquist 1984).

There may be other taxa that belong in this rather loosely defined
“alliance”. Precise circumscription of the “‘alliance”’ is beyond the
scope of this paper and probably cannot be satisfactorily determined
without detailed field-studies, at least, and should benefit from trans-
plant and breeding experiments. For the present we offer a taxonomic

MADRONO, Vol. 35, No. 2, pp. 77-91, 1988

78 MADRONO [Vol. 35

decumbens var. decumbens
decumbens var. robustior
jonesii

lassenianus

nevadincola

sonnei

a

o
N2FaUOnU0
uw wou i
ahaa

es ees

Fic. 1. Distribution of Erigeron spp.

*‘tidying up”’ based almost wholly on herbarium specimens (ca. 1700
sheets, 25 herbaria).

Treatments of Erigeron eatonii and allied taxa in virtually all
recent floras are either by Cronquist or derive directly from his
excellent revision (Cronquist 1947). Since 1947, however, the num-
bers of collections of these taxa available for study have perhaps
doubled and many of the newer collections fill in gaps in geographic
and/or morphologic ranges. Attempts to identify some of these re-
cent collections with Cronquist’s keys and descriptions have led us
to review the taxonomy of the alliance. As a result, we offer revision
of some of his circumscriptions and characterize a previously un-
recognized taxon. In assigning ranks to taxa, we have taken a con-
servative, utilitarian view in order to avoid changes in established
nomenclature so far as practicable.

1988] STROTHER AND FERLATTE: ERIGERON TS

eatonii var. eatonii
eatonii var. lavandulus

eatonii var. plantagineus

eatonii var. villosus

<utre
nou uw
Fal aaa

Fic. 2. Distribution of Erigeron eatonii subspp.

Key TO Erigeron eatonii AND ALLIED TAXA

a. Hairs of stems closely appressed or strongly ascending (except
peduncles near the heads).

b. Flowering stems prostrate, decumbent, or erect, 4—15(—30)
cm long and bearing 1—2(—7) heads; basal leaves evident
at anthesis.

c. Phyllaries 4—-5(—7) mm long; disc corollas 2.5—4(—-5) mm
long; pappus of 12—18(—28) fragile, barbellulate bris-

tles 2.5—4 mm long plus 6-12 fine setae 0.1-0.5 mm

OTN ee pres ae ee eee res es 2. E. eatonii

d. Phyllaries moderately to densely glandular-puber-
ulent (may be pilo-hirtellous to villous as well).

e. Heads 1—2(rarely to 7) per flowering stem, held

80 MADRONO [Vol. 35

well beyond the basal leaves; shorter phyl-
laries '2—*/4 as long as the longer; phyllaries
sparsely pilo-hirtellous with hairs mostly
0.3-0.8 mm long; Rocky Mts. (AZ, CO, se.
WED MT i WN re ae re
Cee On ara eer 2a. E. eatonii var. eatonil
ee. Heads 2—4(1-6) per flowering stem, held just
beyond the basal leaves; shorter phyllaries
ca. %4+ as long as the longer; phyllaries
sparsely to moderately villous with hairs
mostly 0.5—1.5+ long; w.-cen. ID, adj. OR.
Bette ah 2b. E. eatonii var. lavandulus

dd. Phyllaries sparsely, if at all, glandular-puberulent,
variously pilo-hirtellous or villous.

f. Phyllaries sparsely to moderately pilo-hirtellous
with hairs mostly 0.3—0.8 mm long; s.-sw.
OURS nese CA ue ha, ne a ee
See 2c. E. eatonii var. plantagineus
ff. Phyllaries moderately to densely villous with
hairs mostly 1-2 mm long; ne. OR, WA,
cen. 1D. 22 2d. E. eatonii var. villosus
cc. Phyllaries 4.5—10.5 mm long; disc corollas (3.5—)4.5-6.8
mm long; pappus of 20-30 barbellate bristles 4—5
mm long plus 12-20 setae to 1 mm long.

g. Phyllaries 7-10.5 mm long, length 4.7-6.6 times
width; ray floret lamina 7-11 mm long; disc
corollas 4.4—6.8 mm long; ne., w.-cen. NV, adj.
OARS 2 Rarer ear trie 5. E. nevadincola

gg. Phyllaries 4.5-8 mm long, length 3.8—5.5 times width;
ray floret lamina 4.5—6.6(—8.5) mm long; disc
corollas 3.5—5 mm long; border cos. CA-NV and
SCO IN Va yous aes ne toe 6. E. sonnei
bb. Flowering stems mostly erect, (15—)30—60 cm long and bear-
ing 2—5(1—18+) heads; basal leaves often withered prior
tO -ANtNESIS.9 = 35 eee 1. E. decumbens
h. Phyllaries 4-6 mm long; achenes 1.2—1.6 mm long; w.
ORS pitti et la. E. decumbens var. decumbens
hh. Phyllaries 6-8.5 mm long; achenes 1.8—2.5 mm long;
nw. CA. ....... lb. E. decumbens var. robustior

aa. Hairs of stems patent or retrorse.
i. Phyllaries 21-32, mostly 4.2-5.6 mm long, length 3.5-5.3
times width; ray florets none or 12-24; CA. .........
sete Sea ne eee Ben Ee Henao 4. E. lassenianus
u. Phyllaries 36-62, mostly 5.5—7 mm long, length 5—9 times
width; ray florets 20-52; e.-cen. NV, sw. UT. [See also

1988] STROTHER AND FERLATTE: ERIGERON 81

NOMENCLATURE, DESCRIPTION, AND DISCUSSION

1. ERIGERON DECUMBENS Nutt., Trans. Amer. Philos. Soc. ser. 2. 7:
309. 1840 [not Erigeron decumbens Eastwood 1906].—TyYPE:
Nuttall indicated, ““Rocky Mountains, towards the Oregon’’, but
Pennell (1936), in reviewing the meager evidence of Nuttall’s
itinerary in Oregon, reported for late summer of 1835, “‘a record
from ‘prairies of the Wahlamet’, suggesting that Nuttall had
gone... up the Willamette valley”, Nuttall s.n. (Holotype: PH!;
isotypes: frag ex BM in DS!; GH fide Cronquist).

la. ERIGERON DECUMBENS Nutt. var. DECUMBENS—“‘Erigeron de-
cumbens Nutt. subsp. typicus” Cronquist, Brittonia 6:174. 1947.

Stems erect or decumbent only at base, 15-60 cm long, stramin-
eous to reddish at base, sparsely appressed-pubescent; basal leaves
often withered before anthesis, linear to very narrowly oblanceolate,
the longest 5—12(-18) cm long, 3—4(—9) mm wide; mid-stem leaves
mostly linear, 3—9 cm long, 2—4(-8) mm wide; all leaves sparsely to
moderately pilo-hirtellous with patent or ascending hairs; flowering
stems bearing 2—5(1—18+) heads, these held well beyond the basal
leaves, stem/basal-leaf length = 1.8—4.2; involucres broadly hemi-
spheric to rotate, 9-12 mm diam. (pressed); phyllaries 34—47, sub-
equal, linear, length mostly 7—-8(6-10) times width, the longest
(4.3-)5—6 mm long, often with a strong, orange nerve, sparsely to
moderately villous, not glandular; ray florets ca. 35(26—57), corollas
white or fading to white from pinkish or pale blue, lamina 7—12 mm
long, 1-1.5 mm wide; disc florets ca. 200+, corollas yellow, 3—3.5
mm long; achenes (seldom collected) tan with orange ribs, 1.2—1.6
mm long, sparsely strigillose; pappus of ca. 16 whitish, very fragile
bristles ca. 3 mm long plus 8—10+ fine setae ca. 0.1 mm long.

Open grasslands; Oregon: Willamette River drainage (Benton,
Clackamas, Lane, Linn, Marion, Polk, and Yamhill cos.); below 300
m; late May to mid-July.

1b. ERIGERON DECUMBENS Nutt. var. ROBUSTIOR (Cronq.) Cronq. in
C. Hitche. et al., Vasc. Pl. Pacific Northw. 5:175. 1955.—Erig-
eron decumbens Nutt. subsp. robustior Crongq., Brittonia 6:174.
1947.—TypeE: California, Humboldt Co., valley of South Yager
Creek, 26 Jun 1932, J. P. Tracy 10252 (Holotype: UC!; isotype:
JEPS!).

Stems erect or decumbent only at the base, 25—55 cm long, stra-
mineous or purplish at base, finely and sparsely appressed-strigillose

82 MADRONO [Vol. 35

or with weakly spreading hairs 0.3—1 mm long, sometimes minutely
glandular as well; basal leaves often withered before anthesis, linear
to very narrowly oblanceolate, the longest 9-17 cm long, 3—6(—1 1)
mm wide; mid-stem leaves mostly linear, sometimes oblanceolate,
3-6 cm long, 2—5(—11) mm wide; all leaves sparsely pilo-hirtellous
with patent or ascending hairs; flowering stems bearing (1—)2—4+
heads, these held well beyond the basal leaves at peak anthesis, stem/
basal-leaf length = 1.7—3.1; involucres broadly hemispheric to ro-
tate, 12-18 mm diam. (pressed); phyllaries 30-45, subequal, nar-
rowly lanceolate to linear, length 6-8 times width, the longest 6—8.5
mm long, usually with a strong orange nerve, all moderately to
densely villous, little if at all glandular; ray florets 21-36, corollas
lilac to deep blue or bluish violet (often fading to white), lamina 7—
12(-17) mm long, 1.5—3 mm wide; disc florets 150—200+, corollas
yellow, 3.5—4.5 mm long; achenes (seldom collected) tan to greenish
with orange ribs, 1.8—2.5 mm long, minutely and sparsely strigillose;
pappus of 14—20+ whitish, fragile, barbellulate bristles ca. 4 mm
long plus 8-12+ minute setae ca. 0.1 mm long.

Glades and meadows, sometimes associated with serpentine sub-
strates; California (Humboldt and western Trinity cos.; upper drain-
ages of the Eel, Mad, and Van Duzen rivers and headwaters of
Redwood Creek); mostly 700-1500 m; early June to late July.

Reports of E. decumbens var. robustior from Plumas Co., CA,
and from Klamath Co., OR, are based on robust specimens of E.
eatonii var. plantagineus.

2. ERIGERON EATONII A. Gray, Proc. Amer. Acad. Arts 16:91. 1880.—
LECTOTYPE (Cronquist 1947): Utah, Uintah Mountains, on a
divide west of Duchesne River and on a ridge above Bear River
Canyon, 10,000 feet, Jul 1869, S. Watson “546” (Lectotype:
GH!; isolectotypes: NY!, US!).

2a. ERIGERON EATONII A. Gray var. EATONII.—‘“‘Erigeron eatonii A.
Gray subsp. typicus” Cronquist, Brittonia 6:172. 1947.

Erigeron microlonchus E. Greene, Pittonia 3:293. 1898.— LECTOTYPE
(here designated): ““Common on grassy plains and hills of south-
ern Wyoming; collected by the writer plentifully in meadows of
Dale Creek .. .”, 30 Jun 1896, EF. Greene s.n. (Lectotype: ND-
G, 057127!; isolectotype: ND-G, 057128!).

Erigeron eatonii A. Gray f. molestus Cronq., Brittonia 6:172. 1947.—
Type: Utah, Tooele Co., Stansbury Range, east base of Mt.
Deseret Peak, 10,000 feet, 23 Jun 1943, Maguire and Holmgren
21773 (Holotype: NY!; isotype: GH!).

Erigeron canaani Welsh, Great Basin Naturalist 43:366. 1983.—
Type: Utah, Washington Co., Canaan Mountain, 11 Jun 1980,
J. Anderson s.n. (Holotype: BRY!; isotype: NY).

Stems prostrate, decumbent, or erect, 9-20(-32) cm long, green,

1988] STROTHER AND FERLATTE: ERIGERON 83

stramineous, or purplish at base, very sparsely to moderately strigil-
lose with appressed or ascending hairs 0.1—0.8 mm long (rarely with
patent hairs); basal leaves linear or very narrowly oblanceolate, the
longest 5—10(—18) cm long, 2—5(1-9) mm wide; mid-stem leaves
linear, 1-4 cm long, 1—3(—5) mm wide; all leaves moderately strigil-
lose to subglabrous (especially adaxially), hairs appressed to as-
cending (sometimes erect adaxially), mostly 0.3-0.8 mm long, some-
times longer on basal margins; flowering stems bearing 1—2(—7) heads,
these mostly held well beyond the basal leaves, stem/basal-leaf
length = (1.1—)1.7—2.8; involucres hemispheric, 10—13(—16) mm diam.
(pressed); phyllaries 29-56, linear-oblanceolate, length (4.2—)5—6
times width, the longest 4.5-6 mm long, the outermost 3-6 about
half as long as the rest, all moderately to densely glandular-puber-
ulent and sparsely pilo-villous with hairs mostly 0.3—0.8 mm long;
ray florets 21-42, corollas white or variously pink, lilac, or lavender,
sometimes the pigment more pronounced abaxially and distally,
lamina 5—7 mm long; disc florets 80—150+, corollas yellowish to
ochroleucous, often tipped with red or purple; achenes grayish tan,
1.7-2.4 mm long, sparsely hairy; pappus of 16—20 fragile, barbel-
lulate bristles 3—3.5 mm long plus 12-16 fine setae 0.2—0.5 mm long;
2n = 18 (Keil and Pinkava 1976, Nesom 1978).

Open or exposed, often meadowy places, with sagebrush, pinyon-
juniper, aspen-conifer forests; western Rocky Mts., Arizona (Co-
conino Co.), Colorado (Eagle, Grand, Gunnison, Mesa, Moffat,
Montezuma, Montrose, Rio Blanco, and Saguache cos.), Idaho (Ban-
nock, Bear Lake, Bonneville, Caribou, and Franklin cos.), Montana
(Park and Stillwater cos.), Utah (all but Box Elder and Morgan cos.),
and Wyoming (Albany, Carbon, Fremont, Lincoln, Natrona, Park,
Sweetwater, Sublette, Teton, and Uinta cos.); 1900-2700 (1650—-
3600) m; mid-May to mid-August.

Some plants from the Stansbury Range in western Utah differ
from typical E. eatonii in having patent hairs on the stems; in this
regard they approach E. jonesii (q.v.). Cronquist (1947) called such
plants E. eatonii f. molestus. Other plants from the same area closely
match typical E. eatonii.

Welsh (1983) based E. canaani on plants from southwestern Utah
that differ from typical E. eatonii primarily in having very narrow
leaves with rather long hairs along the proximal margins. Although
the habit of these plants is striking, they seem to represent an extreme
form of E. eatonii var. eatonii rather than a distinct taxon. Specimens
quite similar to the type of E. canaani have been collected elsewhere
in the range of var. eatonii (e.g., Duchesne Co., Utah, Neese 9341,
BRY, NY).

2b. Erigeron eatonii A. Gray var. lavandulus Strother & Ferlatte,
var. nov.— TYPE: Idaho, Idaho Co., Seven Devils Mountains,
plateau area of Dry Diggins, dry, barren flat, 1 Aug 1952, A. R.

84 MADRONO [Vol. 35

Kruckeberg 3207 (Holotype: UC!; isotypes: NY!, ORE!, RM!,
RSA!, UTC).

Herbae perennes caulibus floriferis foliis basalibus 1—1.5-plo lon-
gioribus, phyllariis 26—40 plerumque modice glandulosis tantum
parce vel modice villosis (piliis 0.5—1.5 mm longis) raro apicibus
rubris, corollis radiorum lavandulis vel purpureis haud subroseis
vel erubescentibus raro niveis vel albescentibus.

Stems prostrate to decumbent, 5—12(—22) cm long, closely or loose-
ly strigillose with appressed to weakly spreading hairs 0.1—0.8 mm
long, usually minutely glandular as well, at least distally; basal leaves
linear to narrowly oblanceolate, the longest 6—9(—17) cm long, 2—4
(—7) mm wide; mid-stem leaves linear, 2—4(—8) cm long, 1—2(—4) mm
wide; all leaves loosely strigillose; flowering stems bearing 2—4(1-—6)
heads, these held barely if at all above the basal leaves at peak
anthesis, stem/basal-leaf length 1—1.5(—1.9); involucres hemispher-
ic, 9-13 mm diam. (pressed); phyllaries 26-40, subequal or weakly
graduate, linear to lanceolate, length mostly 5—6 times width, the
longest (4.5—)5—7 mm long, little if at all carinate, usually glandular
and sparsely to moderately villous with hairs 0.5—-1.5+ mm long,
rarely reddish on distal margins and in the minutely erose, attenuate
tip; ray florets 25(16-36), corollas deep lavender or bluish-purple,
rarely white or fading to white, not pink or reddish, lamina 5.5—-8
(-—9) mm long, 1-2 mm wide; disc florets 100-—150+, corollas ochro-
leucous to yellowish, often tipped with purple, 2.5—3.5(—4) mm long;
achenes tan with tan to orange ribs, 1.8—2.2 mm long, very sparsely
strigillose; pappus of 12—20 white, fragile, barbellulate bristles 3—3.5
mm long plus 6—9+ minute setae 0.1—0.3 mm long; 2n = 18 (Solbrig
et al. 1969; as E. eatonii var. villosus).

Open grasslands and meadows, mostly in sagebrush scrub com-
munities on scablands; Idaho (Adams, Idaho, Valley, and Wash-
ington cos.) and adjacent Oregon (Union and Wallowa cos.); 900-
1850(—2300) m; mid-May to mid-August.

The plants treated here as var. lavandulus were included in var.
villosus by Cronquist. In addition to differences evident in the key,
var. lavandulus differs from var. vi/losus in having lavender to bluish
ray corollas (rarely white or fading to white) vs. usually white (or
tinged with pink, not bluish), flowering stems usually bear 2—4+
heads vs. | (rarely 2), stem length to basal-leaf length ratio equals
1-1.5(-1.9) vs. 1.7—2.8, and habitat preference is mostly for sage-
brush associations vs. lodgepole pine associations.

Note: Three collections from west of the Cascade crest in northern
Oregon, some 400 km disjunct from typical var. /Javandulus, may
belong within this circumscription.

1) T. J. Howell s.n. [Clackamas Co., “‘on grassy slopes of the
Cascade mountains near Table Rock”’ (with description), ““Rooster

1988] STROTHER AND FERLATTE: ERIGERON 85

Rock” (on label), 22 Aug 1899, ORE!] is the type of Evrigeron pa-
cificus Howell [Fl. Northw. Amer. 1(3):319. 1900]. The specimens
have suffered considerable insect damage and are difficult to inter-
pret; they are somewhat hairier than is typical of var. /avandulus.

2) L. M. Kemp s.n. [Clackamas Co., Fish Creek Mtn., 3600 ft,
27 Jul 1979, ORE!, OSC!] has proportionately wider basal leaves
and longer flowering stems and has absolutely longer ray corollas
(lamina 12-15 mm) than in typical var. /avandulus.

3) G. Whitehead 1905 [Marion Co., Scorpion Mtn., 4800 ft, 28
Jul 1982, ORE!, OSC!] has flowering stems up to twice as long as
basal leaves but otherwise fits well within our circumscription of
var. lavandulus.

Alternative treatments of these collections are: 1) resurrection of
E. pacificus as a distinct taxon or 2) treatment of E. pacificus as
taxonomic synonym of E. eatonii var. lavandulus. A reasonable
choice cannot be made from information at hand and must await
further studies of plants from eastern Clackamas and Marion coun-
ties. It seems certain that the type of FE. pacificus does not fall within
the circumscription of E. eatonii var. villosus as here drawn.

2c. ERIGERON EATONII A. Gray var. PLANTAGINEUS (E. Greene) Cronq.
in C. Hitche. et al., Vasc. Pl. Pacific Northw. 5:175. 1955.—
Erigeron plantagineus E. Greene, Pittonia 3:292. 1898.—Erig-
eron eatonii A. Gray subsp. plantagineus (E. Greene) Crongq.,
Brittonia 6:173. 1947.— LECTOTYPE (here designated): Califor-
nia, Modoc Co., Lava Beds, Jun 1894, Mrs. R. M. Austin s.n.
[268 in UC] (Lectotype: ND-G, 057228!; isolectotypes: ND-G,
057224!, UC!). Note: Cronquist (1947) cited Austin s.n. (ND-G)
as type but did not annotate either of the specimens now in
ND-G.

Erigeron robertianus E. Greene, Pittonia 3:293. 1898.—TyYPE: Or-
egon, ““Roberts’ Ranch”’ (in “‘southeast Oregon’’), 1893, Mrs.
R. M. Austin s.n. (Holotype: ND-G!).

Stems prostrate to decumbent, 10—23+ cm long, stramineous or
somewhat purplish at base, sparsely to moderately strigillose with
appressed or slightly spreading hairs 0.2—0.5 mm long, usually some-
what glandular as well; basal leaves linear to narrowly oblanceolate,
the longest 5—11(—16) cm long, 3—8(—13) mm wide; mid-stem leaves
narrowly oblong to linear, 1.5—3(—5) cm long, 2—4(—8) mm wide; all
leaves sparsely to moderately strigillose with closely appressed to
weakly spreading hairs on both faces, usually minutely glandular as
well; flowering stems bearing 1—4 heads, these held just beyond to
well beyond the basal leaves, stem/basal-leaf length = (1.1-)1.5-
2.7; involucres broadly hemispheric to subrotate, 1 1—12(9-14) mm
diam. (pressed); phyllaries 31—49, subequal to weakly graduate, lin-
ear to lanceolate, length 5—7(4.5—8.5) times width, the longest 5—7

86 MADRONO [Vol. 35

mm long, usually somewhat navicular, little, if at all, glandular,
usually sparsely to moderately hirsute or villous with hairs 0.3-0.8
mm long; ray florets 23-39, corollas white or pink to bluish or purple,
then often fading to white, lamina 6—8(—13) mm long, 1.5-2 mm
wide; disc florets 100-—150+, corollas ochroleucous to yellow, often
tipped with red in age, 3-4 mm long; achenes (seldom collected)
stramineous to orange-tan or puce with stramineous or orange ribs,
1.8—2.3 mm long, sparsely strigillose; pappus of 16—20+ white, fra-
gile, finely barbellulate bristles 3—3.5 mm long plus 6-12 very fine
setae 0.1-0.3 mm long; 2” = 18 (determined from Strother 1359,
CA, Plumas Co.).

Exposed, often meadowy places, often with sagebrush scrub; usu-
ally associated with volcanic/lava substrates, rarely with serpentine
(e.g., Chambers 4358, Oregon, Jackson Co., Red Mtn.); California
(Lassen, Modoc, Plumas, Shasta, and Siskiyou cos.) and Oregon
(Jackson, Klamath, and Lake cos.); 1500—2300(1050—2500) m; late
May to mid-August.

Robust specimens of E. eatonii var. plantagineus have been con-
fused with EF. decumbens var. robustior. In Plumas Co., California,
E. eatonii var. plantagineus co-occurs with E. lassenianus (cf. Kear-
ney 153, US) and specimens intermediate between the two may be
found (see discussion of EF. lassenianus).

2d. ERIGERON EATONII A. Gray var. VILLOSUS (Crongq.) Cronq. in C.
Hitchce. et al., Vasc. Pl. Pacific Northw. 5:175. 1955.—Erigeron
eatonil A. Gray subsp. villosus Cronq., Brittonia 6:172. 1947.—
TYPE: Oregon, “‘open summit of the Wallowa Mountains’’, 2300
m, 29 Jul 1907, Cusick 3186 [Holotype: NY!; isotypes: GH!,
MIN, MO!, ORE(2)!, RM!, UC!, US (2)!, WS].

Stems weakly decumbent to erect, 10—15(5—30) cm long, stramin-
eous or reddish at base, moderately to densely appressed-strigillose
with hairs 0.3-0.7 mm long, sometimes minutely glandular as well;
basal leaves linear to narrowly oblanceolate, the longest 4—8(-19)
cm long, 4—7(2—11) mm wide; mid-stem leaves linear, 2—3(—5) cm
long, 1—2(—3) mm wide; all leaves appressed-strigillose, rarely mi-
nutely glandular as well; flowering stems bearing 1(—2) heads, these
held well beyond the basal leaves at peak anthesis, stem/basal-leaf
length = 1.7—2.8; involucres broadly hemispheric to nearly rotate,
12—14(10—-17) mm diam. (pressed); phyllaries 28-60, linear to very
narrowly oblanceolate, length mostly 5—9.3 times width, the longest
4.5-6.5 mm long, subequal, little if at all carinate, moderately to
densely shaggy-villous with white hairs to 0.8—1.5 mm long, usually
reddish on distal borders, tips attenuate and minutely erose; ray
florets 35(20-—50), corollas white, sometimes pinkish, never bluish
(rarely bicolored—abaxially pink/adaxially white; e.g., Maguire 2683 1
from OR, Grant Co., Strawberry Mtn.), lamina (5—)6—8 mm long,

1988] STROTHER AND FERLATTE: ERIGERON 87

1-2.5 mm wide; disc florets (60—)90-—160, corollas ochroleucous to
yellowish, 2.5-3.5 mm long, lobes sometimes with bristle-like hairs;
achenes (seldom collected) tan with tan ribs, 1.7—2.2 mm long,
sparsely strigillose; pappus of 16—26 white, fragile, barbellulate bris-
tles 2.5—3.5 mm long plus 5-12 very fine setae 0.3-0.5 mm long.

Meadows and open places, often in lodgepole pine forests, some-
times in sagebrush; Idaho (Adams, Blaine, Custer, Idaho, Lemhi,
Valley, and Washington cos.), Oregon (Baker, Crook, Grant, Harney,
Union, Wallowa, and Wheeler cos.), and Washington (Asotin, Gar-
field, and Kittitas cos.); 1850—2500(1 500-2950) m; mid-June to mid-
August.

3. ERIGERON JONESII Cronq., Brittonia 6:166. 1947.—TyYPE: Nevada,
White Pine Co., Aurum, 9 Jul 1891, M. E. Jones s.n. (Holotype:
POM; isotypes: MO!, NY, ORE!, PH, RM, UC!, US).

Erigeron wahwahensis S. Welsh, Great Basin Naturalist 43:368.
1983.—TyYpe: Utah, Beaver Co., Wah Wah Mtns., Pine Grove
Pass, ca. 12 miles SSW of Wah Wah Spring, 2450 m, 12 Jun
1982, 8. L. Welsh 21229 (Holotype: BRY!; isotypes: CAS!, GH,
MO, NY, POM, RM, US, UT, UTC).

Stems decumbent to ascending or nearly erect, 8—20(—38) cm long,
sparsely to densely hirtellous with patent or retrorse hairs 0.1—0.8+
mm long (rarely subglabrous or with ascending hairs); basal leaves
spatulate to oblanceolate or linear, the longest 5—8(—17) cm long, 3-
8(-19) mm wide, rarely obscurely toothed; mid-stem leaves linear
to oblong, often somewhat clasping at base, 1—3(—4) cm long, 4—6
(—9) mm wide; all leaves sparsely to moderately pilo-hirtellous with
patent or ascending hairs; flowering stems bearing (1—)2—4 heads,
these held well beyond the basal leaves at peak anthesis, stem/basal-
leaf length = 1.7—3.5; involucres hemispheric to turbinate or sub-
rotate, 12-18 mm diam. (pressed); phyllaries 36-62, subequal or
weakly graduate, linear to lance-linear, length mostly 5—7(—9) times
width, the longest 5.5—7 mm long, little if at all carinate, somewhat
indurate basally, minutely glandular and sparsely to moderately vil-
lous; ray florets 34(20—-52), corollas whitish, usually with blush of
blue or pink distally on abaxial face, rarely colored on adaxial face,
lamina 5—7 mm long, 1—2 mm wide; disc florets 150—200+, corollas
ochroleucous to yellowish, sometimes tipped with pink or purple,
3-—3.5 mm long; achenes grayish tan with pale ribs, 2.2—2.5 mm long,
sparsely strigo-hirtellous; pappus of 20-24 whitish, fragile, barbel-
lulate bristles ca. 3 mm long plus ca. 12 fine setae 0.1-0.3 mm long.

Open places, mostly in sagebrush, pinyon-juniper, ponderosa pine,
oak-maple, and white fir communities on granitic, limestone, or
volcanic substrates; eastern Nevada (Elko, Lincoln, Nye, and White
Pine cos.) and adjacent Utah (Beaver, Iron, Juab, Millard, Tooele,
and Washington cos.); 2000—2700(—3400) m; late May to mid-Au-
gust.

88 MADRONO [Vol. 35

Many specimens from various genera and families from south-
western Utah, especially Washington Co., do not “‘conform”’ or are
unusual within their groups. Welsh (1983) noted that E. wahwah-
ensis 1s “‘more or less intermediate between phases of E. eatonii and
E. jonesii’’. We agree that in and near the area where both E. eatonii
and E. jonesii may be found, some specimens are intermediate be-
tween the two (e.g., Atwood 1484 from Washington Co. and Cottam
3977 from Iron Co.). In the type of E. wahwahensis, however, not
only is the indument like that of EF. jonesii but so is the “general
aspect”’.

Pollen volumes in cubic micrometers (cum) were determined for
five samples of ““wahwahensis” from Beaver and Washington cos.,
UT (6371-7795 cum) and for four samples of E. jonesii s. str. from
Elko and White Pine cos., NV (6710-9097 cum). For five samples
from E. eatonii var. eatonii from Colorado, Utah, and Wyoming,
the volumes are 6813-7832 cum. These ranges of volumes suggest
that polyploidy may occur within species or varieties of this group
of erigerons and do not help resolve placement of “‘wahwahensis’’.

4. ERIGERON LASSENIANUS E. Greene, Fl. Franciscana 389. 1897.—
LECTOTYPE (Cronquist 1947): California, Plumas Co., Mt. Dyer,
1880, Mrs. R. M. Austin s.n. (Cronquist cited GH and ND-G;
we have seen both and take the ND-G specimen to be the
lectotype).

Erigeron lassenianus E. Greene var. deficiens Cronquist, Brittonia
6:171. 1947.—TyYPpe: California, Plumas Co., United States For-
est Reserve, on trail to Long Valley, 15 Jun 1927, A. Eastwood
14613 (Holotype: CAS!).

Erigeron flexuosus Cronq., Brittonia 6:174. 1947.—TypeE: Califor-
nia, Trinity Co., Trinity Alps Resort, 25 Jun 1937, A. Eastwood
and J. T. Howell 4903 (Holotype: CAS!).

Stems prostrate to ascending or erect, 9—20(—35) cm long, often
somewhat flexuous, mostly stramineous at base, sparsely to densely
strigillose to hirtellous with ascending to patent hairs 0.2-0.7 mm
long, usually glandular as well, at least on the distal 1-3 internodes
(peduncles); basal leaves very narrowly oblanceolate to spatulate or
linear, the longest 5—12(—15) cm long, 3-—6(—9) mm wide; mid-stem
leaves linear to narrowly spatulate, 1-3(—5) cm long, 1.5—3(-5) mm
wide; all leaves sparsely to moderately pilo-hirtellous to villous with
erect or ascending hairs, often minutely glandular as well; flowering
stems bearing (1—)3-7+ heads, these held well beyond the basal
leaves at peak anthesis, stem/basal-leaf length = 1.5—-3.5; involucres
mostly hemispheric, 6—10(—12) mm diam. (pressed); phyllaries 21-
32, weakly graduate, oblanceolate, length mostly 3.5—5.3 times width,
the longest 4.2—-5.6 mm long, all somewhat carinate, minutely gran-
ular-glandular and moderately to densely pilo-villous; ray florets
none or 21(12—24), corollas pink to lavender or white, lamina 4.5—

1988] STROTHER AND FERLATTE: ERIGERON 89

8 mm long, 1.5—2.5 mm wide; disc florets 60—90(—120+), corollas
ochroleucous to yellowish, sometimes reddish at tip, 2.5—3.5 mm
long, lobes often bearing 1—2 bristle-like hairs; achenes tan to green-
ish brown with stramineous ribs, 1.5-2.3 mm long, very sparsely
strigillose; pappus of 12—24 whitish, fragile, barbellulate bristles 1.5—
3 mm long plus 8-15 very fine setae 0.1-0.4 mm long; 2n = 18
(determined from Strother 1360, CA, Plumas Co.).

Open places, usually associated with ultramafic (serpentine) or
glacial moraine substrates; California (Butte, Eldorado, Lassen,
Mendocino, Plumas, Shasta, Sierra, Tehama, and Trinity cos.); 700—
2300 m; early June to early September.

Until fairly recently, only a few collections referable to E. flexuosus
s. str. were available. Thus, their characteristic rosettes of basal
leaves (1—3-nerved) and their habital similarity to E. lassenianus
were not evident. Hairs on the stems of some plants, especially those
from Trinity Co., are only weakly spreading, unlike those of typical
E. lassenianus. In number of heads per stem, numbers and sizes of
phyllaries and florets, and glandular hairs of the phyllaries, however,
these plants correspond very well with typical E. lassenianus. Con-
sequently, we choose to treat the two as one.

Note: Plants from Mendocino Co. (all from near Hell’s Half Acre
north of Hull Mt.) have phyllaries eglandular or nearly so and in
this regard are similar to E. eatonii var. plantagineus. Plants with
sparsely glandular phyllaries and hairs of the stems spreading to
appressed are found near Red Clover Valley, Plumas Co., e.g., A.
A. Heller 8704 and J. L. Strother 1360. In Strother 1360, 2n = 9 II
with regular meiosis. Although these collections are somewhat atyp-
ical and approach E. eatonii var. plantagineus (q.v.), we include
them here in E. lassenianus.

5. ERIGERON NEVADINCOLA S. F. Blake, Proc. Biol. Soc. Wash. 35:
78. 1922.— Erigeron nevadense A. Gray, Proc. Amer. Acad. Arts
8:649. 1873, non Wedd. 1857.— LECTOTYPE (Cronquist 1947):
Nevada, Storey Co., ““Cedar Hill and Mt. Davidson, near Vir-
ginia City” (label), 1863-64, H. G. Bloomer s.n. (Lectotype:
GH)!).

Stems decumbent to erect, 14—22(—33) mm long, usually green or
stramineous at base, sparsely to moderately appressed-strigillose
(sometimes with patent hairs, see discussion), the hairs 0.2-0.7 mm
long; basal leaves linear to oblanceolate or spatulate, the longest 6—
12(-19) cm long, 3—6(—13) mm wide; mid-stem leaves linear to nar-
rowly oblong, 2—4(—7) cm long, 1—3(—7) mm wide, often with wavy
margins; all leaves sparsely to moderately pilo-hirtellous to ascend-
ing-strigillose, sometimes minutely glandular as well; flowering stems
bearing | head (rarely 2), heads held well beyond the basal leaves
at peak anthesis, stem/basal-leaf length = 1.5—3.3; involucres broad-
ly hemispheric to turbinate, (14—)17—23 mm diam. (pressed); phyl-

90 MADRONO [Vol. 35

laries 31-68, weakly graduate, linear to lanceolate, length 4.7—6.6
times width, the longest 7-10.5 mm long, villous with hairs 1-2 mm
long, little if at all glandular; ray florets 22—39, corollas white, often
flushed with pale lavender or pink on abaxial (sometimes adaxial)
face, rarely suffused with lavender, lamina 7-11 mm long, 1.5-3
mm wide; disc florets 150—250+, corollas ochroleucous or yellowish,
4.4—6.8 mm long; achenes stramineous to whitish with pale or orange
ribs, 4—4.5 mm long, strigo-hirtellous to subsericeous; pappus of 20—
24+ stramineous, rather coarse, barbellate bristles 4-5 mm long
plus 12-20 setae 0.3—1 mm long.

Meadowy places and gravelly slopes, mostly in sagebrush and
pinyon-juniper communities; northeastern to west-central Nevada
(Elko, Humboldt, Lander, Lyon, Ormsby, Pershing, Storey, and
Washoe cos.) and adjacent California (Lassen and Sierra cos.); 1450—
2850 m; mid-May to late July.

Occasional plants (e.g., Schoolcraft 984 from east of Skedaddle
Creek, Washoe Co., NV) are intermediate between FE. nevadincola
and E. eatonii var. plantagineus. Populations from the Virginia Range,
Lyon Co., NV (e.g., Tiehm 7728 and 7814) that match typical E.
nevadincola in most traits include some plants with typical, ap-
pressed-strigillose indument on stems and other plants with strongly
spreading to quite erect hairs on stems. This dimorphism is also
encountered in E. eatonii var. eatonii (q.v.).

6. ERIGERON SONNEI E. Greene, Pittonia 1:218. 1888.—Ervrigeron
nevadensis (sic) A. Gray var. sonnei (E. Greene) Smiley, Univ.
Calif. Publ. Bot. 9:373. 1921.—TyYPE: protologue: ““Western slope
of the Washoe Mountains, Nevada’’; label: ““Nevada and Placer
Counties, Cal., W. slope Washoe Mts.”’, 22 Jul 1888, C. F. Sonne
“2” (Holotype: ND-G!).

Stems prostrate to decumbent or erect, 4—12(—21) cm long, stra-
mineous or somewhat reddish at base, sparsely to moderately strigil-
lose with appressed or ascending hairs 0.1—0.7 mm long, not glan-
dular; basal leaves linear to narrowly oblanceolate, the longest 3-
10 cm long, (1.5—)4—6 mm wide; mid-stem leaves linear, 1-3 cm
long, (0.7-)2—4 mm wide; all leaves sparsely to moderately pilo-
hirtellous to ascending-strigillose, not glandular; flowering stems
bearing | head (rarely 2), the heads held just beyond to well beyond
the basal leaves at peak anthesis, stem/basal-leaf length = 1.2-3;
involucres broadly hemispheric to subrotate, (8—)12—16 mm diam.
(pressed); phyllaries 22—50, subequal or weakly graduate, linear to
lanceolate, length 3.8—5.5 times width, the longest 4.5-8 mm long,
flat to weakly navicular, noticeably scarious-margined, hirsutulous
to villous with hairs 0.5—1.5 mm long, not glandular; ray florets 15-
36, corollas white, usually flushed abaxially with pink, lavender, or
violet, lamina 4.5—6.6(—8.5) mm long, 1.5—2.5 mm wide; disc florets
80-150+, corollas ochroleucous to yellow, sometimes tipped with

1988] STROTHER AND FERLATTE: ERIGERON 2

red, 3.5—5 mm long; achenes tan to reddish brown with stramineous
to orange ribs, 2.8-3.5 mm long, strigo-hirtellous, more so distally;
pappus of 18-30 barbellate bristles 3.5—5 mm long plus 8-12 fine
setae 0.5—1.5 mm long; 2” = 18 (determined from Strother 1357,
CA, Alpine Co.).

Meadowy places, seeps, or rocky flats, often with sagebrush; Cal-
ifornia (Alpine, Eldorado, Mono, Nevada, Placer?, Plumas?, and
Sierra cos.) and Nevada (Lyon, Nye, and Washoe cos.); 1800-2800
m; late May to early September.

Many specimens from along the border of California and Nevada
near Lake Tahoe cannot be assigned confidently to either E. sonnei
or E.. nevadincola; elsewhere the two morphs are easily distinguished.
These intergrading taxa are probably conspecific; we maintain them
at specific rank for the present in order to preserve established no-
menclature. Some plants from north of Lake Tahoe (e.g., old Lem-
mon collections labeled “‘Sierra Valley’’) are intermediate between
E. sonnei and E. eatonii var. plantagineus.

ACKNOWLEDGMENTS

We thank curators and staff of the following herbaria for loans, for searching for
specimens, or for arranging for access to collections during visits: A, BRY, CAS,
DAV, DS, DUKE, GH, HSC, JEPS, LL, MO, ND, ND-G, NY, ORE, OS, OSC, PH,
POM, RM, RSA, TEX, UC, US, and UTC. For help in other ways, we thank A.
Cronquist, B. Ertter, R. Hartman, D. Haskell, G. Nesom, A. Smith, E. Voss, S. Welsh,
and B. Williams.

LITERATURE CITED

BEAMAN, J. H. 1957. The systematics and evolution of Townsendia (Compositae).
Contr. Gray Herbarium 183:1-151.

CRONQUIST, A. 1947. Revision of the North American species of Erigeron, north
of Mexico. Brittonia 6:121-302.

1984. Rubiaceae. Jn A. Cronquist et al., eds., Intermountain flora. Vol. 4.
N.Y. Botanical Garden, New York.

DempPsTER, L. and F. EHRENDORFER. 1965. Evolution of the Galium multiflorum
complex in western North America. II. Critical taxonomic revision. Brittonia
17:289-334.

KEIL, D. J. and D. J. PINKAVA. 1976. Chromosome counts and taxonomic notes
for Compositae from the United States and Mexico. Amer. J. Bot. 63:1393-
1403.

Nesom, G. 1978. Chromosome numbers in Erigeron and Conyza (Compositae).
Sida 7:375-381.

Noyes, R. D., D. E. SOLtis, and J. H. BEAMAN. 1987. Cytological and electrophoretic
investigations in Erigeron compositus Pursh (Asteraceae). Amer. J. Bot. 74(5):
748. [Abstract.]

PENNELL, F. W. 1936. Travels and scientific collections of Thomas Nuttall. Bartonia
18:1-51.

SOLBRIG, O. T., L. C. ANDERSON, D. W. KyuHos, and P. H. RAVEN. 1969. Chro-
mosome numbers in Compositae VII: Astereae III. Amer. J. Bot. 56:348-353.

WELSH, S. L. 1983. A bouquet of daisies (Erigeron, Compositae). Great Basin
Naturalist 43:365-368.

(Received 3 Aug 1987; revision accepted 22 Dec 1987.)

A COMPARISON OF NORTH AND SOUTH AMERICAN
LUPINUS GROUP MICROCARPI (LEGUMINOSAE)

RHONDA RIGGINS
Biological Sciences Department,
California Polytechnic State University,
San Luis Obispo 93407

ABSTRACT

Lupinus group Microcarpi occurs disjunctly in North and South America, primarily
in central California and Chile. This study addresses the question of whether the
disjunct representatives, variously referred to L. microcarpus, L. densiflorus, L. ruber
or L. subvexus, are distinct. Data for vegetative and floral features were taken from
South American specimens and were compared to those from population samples
from California. The analyses show that some South American plants are smaller,
but in all features have a range of variation within that of California plants. Affinities
of the South American specimens were assessed by multigroup discriminant analysis
and an a posteriori classification procedure whereby each one was assigned to a
population sample from California. The South American specimens were assigned
to a few California populations identified as L. densiflorus, L. subvexus and L. ruber,
or intermediates between them. Neither floral nor vegetative features can be used to
distinguish the South American representatives of group Microcarpi from some North
American representatives.

RESUMEN

Lupinus del grupo Microcarpi ocurre descontinuadamente en América del Norte y
del Sur, principalmente en California Central y en Chile. Este estudio se dirige a tratar
de resolver la pregunta que si las especies llamadas L. microcarpus, L. densiflorus, L.
ruber y L. subvexus son distintas. Los datos de las caracteristicas vegetativas y florales
fueron tomados de ejemplares sudamericanos y comparados con ejemplares obtenidos
de poblaciones en California. El analisis de los datos indica que algunas de las plantas
sudamericanas son mas pequenas que las de California, pero en todas las otras ca-
racteristicas, el rango de varaciOn esta dentro del que se obtiene de los ejemplares
obtenidos en California. Las afinidades de los ejemplares sudamericanos fueron va-
lorados por medio de un analisis discriminativo multigrupo y un método de clasi-
ficacion de posterioridad en el cual cada uno de los ejemplares sudamericanos fueron
asignados a una muestra de la poblacidn de California. Los ejemplares
sudamericanos se asignaron a unas poblaciones californianas identificadas como L.
densiflorus, L. ruber y L. subvexus 0 especies intermedias. Las caracteristicas vege-
tativas y florales no se pueden usar para distinguir entre las especies obtenidas en
America del Sur y aquellas obtenidas en California.

The informal group Microcarpi is easily delimited from the var-
ious assemblages of Lupinus summarized by Charles Piper Smith
(1944). It is a group of annuals with sessile perfoliate cotyledons,
ovoid two-seeded fruits and verticillate flowers. Members of the
group occur disjunctly in North and South America, primarily in
central California and Chile.

MADRONO, Vol. 35, No. 2, pp. 92-104, 1988

1988] RIGGINS: LUPINUS GROUP MICROCARPI 95

In a series of papers Smith (1917, 1918a,b, 1919) treated the
Microcarpi as consisting of five species including 35 new or newly
combined varieties. The group has not been studied in its entirety
since. Authors of regional floras have treated it only in part, and
their various interpretations largely have resulted in a more con-
fusing taxonomy. No two subsequent treatments are in complete
agreement as to the disposition of the taxa or the names to be used
for them. The disagreement centers around a fundamental taxo-
nomic question: are populations from North and South America
distinct?

In this paper I compare morphological data obtained from her-
barium specimens from South America with those obtained from
population samples from California. The aim of the comparison is
to document the range of morphological variation in the disjunct
representatives, and to determine if the South American plants are
morphologically distinct from their North American counterparts.
I also compare distributional and ecological information obtained
from the specimens and from my collection data.

HISTORICAL PERSPECTIVE

Smith’s (1917, 1918a,b, 1919) group Microcarpi included L. mi-
crocarpus Sims, described from plants grown from seed originally
collected in Chile, and L. densiflorus Benth., described from plants
grown from seed collected by Douglas in California. Smith also
recognized L. subvexus C. P. Smith, L. horizontalis Heller and L.
luteolus Kell. Three of these (L. microcarpus, L. densiflorus and L.
subvexus) were described as occurring in both North and South
America. These three and L. horizontalis, of California desert hab-
itats (Smith 1918a), form a problematical complex. The fifth species,
L. luteolus, was described as occurring in California and Oregon. It
can be separated from the complex by several features (Smith 1919a),
and has been treated as a distinct species by subsequent authors.

Jepson (1936) commented on the close resemblance of North and
South American specimens. He wrote (p. 278), “‘In certain cases, if
the labels were removed, it would seem impossible, on the basis of
the material itself, to say whether a given sheet were Californian or
Chilean.” Jepson placed all California representatives of the complex
into L. microcarpus. He considered a portion of the California ma-
terial, including L. subvexus, to be typical of the species. He rec-
ognized three additional varieties, L. m. var. densiflorus Jeps., L.
m. var. horizontalis Jeps., and L. m. var. ruber (Heller) C. P. Smith
(=L. ruber Heller).

The only other work in which North American members of the
complex are treated as L. microcarpus is that of Hitchcock et al.
(1961). They recognized L. m. var. scopulorum C. P. Smith from

94 MADRONO [Vol. 35

Vancouver Island and adjacent islands of Washington, and L. m.
var. microcarpus for all other populations from Washington to Baja
California, and South America.

Munz’s (1959) treatment of the California complex is diametri-
cally opposed to Jepson’s. Munz recognized L. horizontalis, L. ruber,
L. subvexus with four varieties, and L. densiflorus with six varieties.
From this treatment it might be concluded that the name, L. mi-
crocarpus, does not apply to North American plants.

Dunn and Gillett (1966) stated that L. microcarpus is a southern
hemisphere relative of the L. densiflorus complex of the northern
hemisphere. They concluded that North American taxa could not
be interpreted as L. microcarpus because its original description
referred to blue flowers and torulose pods, but did not refer to keel
ciliation.

In a recent dissertation Planchuelo (1978) placed all Argentinean
specimens of the group into L. microcarpus. She treated the Argen-
tinean taxa described by Smith (1943) as synonyms, but did not
study the Chilean taxa described by Smith (1918a,b, 1940).

To avoid confusion in the following discussion, I refer to North
American representatives as the L. densiflorus complex, and follow
Munz’s (1959) treatment. Lupinus luteolus is excluded from the
study.

METHODS AND MATERIALS

To document geographic distribution of South American mem-
bers of group Microcarpi, | examined approximately 125 collections
from BM, CAS, DS, GH, K, MO, RSA, UC and US. Collection data
for those used in the analyses are given in Table 1. A total of 74
specimens for 56 collections were measured, and are identified by
numbers as given in Table 1. The South American specimens include
representatives of the nine taxa recognized by Smith (1918a,b, 1940,
1943) as occurring in Chile and Argentina. Six collections are type
specimens.

To document distribution and variation of North American taxa,
I collected extensively in California and consulted herbarium spec-
imens from outside the state. The 41 samples used here (Table 2)
are part of a larger study of the L. densiflorus complex in California.
Each sample consisted of 20 plants, so data from 820 specimens
form the data base. Most of the samples are from San Luis Obispo
Co., near the center of the range of the complex and where all four
species (L. densiflorus, L. ruber, L. subvexus, L. horizontalis) are
known to occur (Munz 1959, Hoover 1970). The populations are
from localities along west to east climatic gradients characterized by
decreasing winter rainfall and increasing summer temperatures. Al-
though the majority of populations can be identified as belonging

1988] RIGGINS: LUPINUS GROUP MICROCARPI 95

TABLE 1. COLLECTION DATA FOR SOUTH AMERICAN SPECIMENS OF Lupinus. Num-
bers in parentheses represent identification numbers for purposes of analysis. An
asterisk (*) designates specimens for which data for all variables were obtained and
included in the diagnosis.

Without collection data (1 GH*). Argentina, Chubut: Pampette s. of Lago Colhué
Huapi, Riggs 56 (type of L. verticillatus C. P. Smith, 2 GH*). Neuquen: Chos Malal
y Agrio, 600-1200 m, Comber 188 (type of L. comberanus C. P. Smith, 3 K*). Rio
Negro: Vicinity of General Roca, 250-360 m, Fischer 280 (type of L. fischerianus C.
P. Smith, 4 BM, GH*, K). Chile, without collection data (5 BM*); without locality,
Cuming s.n. (6 BM*); Cuming 567 (7 BM); Cruckshanks 135 (8 K*); 1832, Bridges
s.n. (9 K*); 1864-65, Reed s.n. (10a,b K* 2 sheets); Feb 1888, Philippi s.n. (11 K*).
Province undetermined, Andes, Reynolds s.n. (12 GH*); Salto de Conchali, Nov
1883, Philippi s.n. (13 BM*); San Pedro Nolasco, collector unknown (14 BM*); Cor-
dillero de Curico, Ruiz P. s.n. (15 GH*). Aconcagua: Uspallata Pass, Juncal, 2300
m, Buchtien 1180 (16a BM*, 16b GH*). Antofagasta: Taltal, 600 m, Werderman 856
(17a BM*, 17b DS*, 17c GH*, 17d K*, 17e UC*); ca. 10 km e. of Taltal, 75 m,
Worth and Morrison 15807 (18 UC). Arauco: Arauco, Pennell 1297 (19 GH*). Ata-
cama: Cerro Campana, 15 Nov 1884, Philippi and Borchers s.n. (20 BM); Rio San-
carron below Rucas, ca. 3200 m, Johnston 6204 (21a GH*, 21b K*). Biobio: Paila-
hueque, Pirion 203 (22 GH*). Cautin: Between Temuco and Rio Quepe, Dec 1905,
collector unknown (23 BM*); Temuco, Elliott 218 (24 BM*). Colchagua: San Fer-
nando near Tinguiririca Bridge, Montero 15 (25 GH*). Coquimbo: Banos del Toro,
3500 m, Werdermann 197 (26a BM*, 26b CAS*, 26c GH*, 26d UC*); Coquimbo,
July—Aug 1856, Harvey s.n. (27 GH*, K); 14 km e. of Nueva Elqui, 3200 m, Wa-
genknecht 18122 (28 GH*, UC). Concepcion: Concepcion, Reed s.n. (type of L.
densiflorus var. reedii C. P. Smith, 29 GH*); Concepcion, Elliott 78 (30 BM*); Con-
cepcion, Nov 1926, Gunther and Buchtien s.n. (31 BM*); Lota, 7 Nov 1868, Cun-
ningham s.n. (32 K*); Lota, 20 Dec 1902, Elwes s.n. (33 K*). O’Higgins: Rancagua,
Bertero s.n. (34 K*); Cachapual, Rancagua, Bertero 393 (35 GH*); Cachapual, Ran-
cagua, Bertero 393 et 1116 (36a BM*, 36b GH*). Santiago: Bath of Colina, 1825,
Macrae s.n. (type of L. densiflorus var. barbatissimus C. P. Smith, 37 GH*, K); Colina,
1825, Macrae s.n. (38 K*); 3 km n. of El Tabo, 20 m, 30 Nov 1970, Simon s.n. (39
DS*, RSA); Rio Teso Romeral, Biere 57 (40 GH*). Talca: Talca, Nov 1925, Gunckel
s.n. (41 GH*). Valparaiso: Valparaiso, Cuming 567 (42a BM*, 42b K*); Valparaiso,
1844, Bridges s.n. (43 BM*); rd from Valparaiso to Quillota, Bridges s.n. (type of L.
densiflorus var. decumbens C. P. Smith, 44 K*); Valparaiso, Cumming s.n. (45 K*);
Valparaiso, Robinson s.n. (46 K*); Valparaiso, Mathews 363 (47a BM*, 47b GH*,
47c K*); Valparaiso, 1914, Calvert s.n. (48 BM); ca. 4 km from Valparaiso on rd to
Quebrada Verde, 290 m, Morrison 16713 (49a GH*, 49b K*, UC); Renaca ca. 18
km from Valparaiso, 10 m, Morrison 16847 (50 GH*, K, UC); between Vina del
Mar and Concon, 60 m, Landeman 193 (S1a BM*, 51b K*); Vina del Mar a Concon,
Pirion 268 (52 GH*); Concon, collector unknown (53 BM*); 21 m wege nach Concon,
Nov 1928, Gunther and Buchtien s.n. (54a CAS*, 54b DS*); 6.2 km n. of Puchuncavia,
30 m, Simon 134 (55 CAS*, RSA); Limache, Camino al Paugal, Looser 135 (56 GH*).

to one of the four species of Munz and Hoover, several are mor-
phologically intermediate and cannot be identified with certainty.
Morphological data consisted of seven vegetative and 12 floral
variables. Leaf measurements were taken from the largest leaf of the
specimen, and floral measurements from flowers at anthesis. Sev-
enteen quantitative variables are listed in Table 3. The other two

96 MADRONO [Vol. 35

TABLE 2. COLLECTION DATA FOR CALIFORNIA POPULATION SAMPLES OF THE Lu-
pinus densiflorus COMPLEX. Collection numbers are those of the author.

Kern Co.: 1.0 min. of Reyes Sta., 7134; Crocker Cyn., 2.9 mie. of San Luis Obispo
Co. line, 1/54; Gypsum Mine Rd., 1.0 mi e. of Simmler—Bitterwater Rd., 1186.
Monterey Co.: county rd G19, | mi w. of US 101, n. of Bradley, 1/179. San Luis
Obispo Co.: CA 166, 3.1 mi w. of Sierra Madre Rd., 1/133; county rd 285, 0.9 mi
se. of CA 58, 1137; Hurricane Rd., 0.9 mi ne. of county rd 285, 1138; base of Crocker
Grade at county rd 285, 1/39; 1140; slope w. of San Juan R. at CA 58, 1/41]; Shell
Cr. Rd. at CA 58, 1142; Atascadero—Creston Rd., 1.9 mie. of Templeton Rd., 1/43;
Huerhuero—LaPanza Rd., 0.2 mi nw. of CA 58, 1144; CA 41, 0.7 mie. of Cripple
Cr. Rd., 1145; El Camino Real, Santa Margarita, 0.3 mie. of US 101, 1146; Pozo
Rd., 0.3 mie. of CA 58, 1147; Pozo Rd., 0.7 mi w. of Salinas R. Bridge, 1148; CA
166, 11.9 mie. of US 101, 1/49; CA 166, 3.2 mie. of Sierra Madre Rd., 1150; Hi
Mt. Rd., 8.8 mi ne. of Lopez Lake Rd., //5/7; Klau Mine Rd. just e. of Cypress Mt.
Rd., 1152; county rd 285, 6.9 mis. of CA 58, 1153; Elkhorn Trail Rd., 4.8 mi se.
of Hurricane Rd., 7755; 1156; county rd 285, 2.3 min. of Coachoro Camp Rd., 1/157;
1158; Avenales Ranch Rd., 3.3 mi e. of American Cyn. Rd., 1160; Avenales
Ranch Rd., 0.3 mi nw. of Avenales Guard Sta., 1/61; Avenales Ranch Rd., 2.0 mi
se. of Avenales Guard Sta., 1/62; 1163; USFS Rd., 2.1 mie. of Los Machos Cr.,
1171; 1172; Thirty-Five Cyn. Rd., 2.7 mis. of Branch Mt. Rd., 1173; Cable Corral
Rd. at CA 166, 7174; Almond Spring Ranch, Adelaida—Nacimiento Rd., 1175; Na-
cimiento Rd., 1.4 mi e. Chimney Rock Rd., 1/76; Nacimiento Lake Rd., 0.6 mi n.
of Nacimiento Dam, //77; Wellsona Rd. at River Rd., //80; El Pomar Rd., 0.1 mi
s. of Vaquero Drive, /18/; Eagle Ranch, 0.3 mi s. of Santa Barbara Rd. n. of Santa
Margarita, 7/782; Camatti Cyn. Rd., 1.2 mis. of Gillis Cyn. Rd., 1/85.

were appraisals of wing and keel ciliation. Wing ciliation was re-
corded as either of three states: 0, absent; 1, present above; 2, present
above and below. Keel ciliation was recorded as either of two states:
0, absent below; 1, present below. I selected the 19 variables to
summarize size and shape of vegetative and floral structures, and
to include features used in previous treatments.

Some taxonomic or field characters are not easily assessed for
numerical analyses and were excluded for this reason. Smith (1918b
et seq.), Munz (1959) and Hoover (1970) distinguished L. densiflorus
as having spreading or arching racemes with secund flowers and
fruits. This feature is related to the degree of branching on any one
plant, and where the plants are growing. Often flowers on primary
racemes do not become secund. Plants otherwise identifiable as L.
subvexus or L. horizontalis may develop secund flowers and fruits.
The feature is neither consistent within populations, nor unique to
those identifiable by other features as L. densiflorus. Furthermore,
it is extremely difficult to assess in pressed specimens if fruits are
not present. Lupinus horizontalis, L. ruber and L. subvexus have
been described as having ascending, erect or suberect flowers and
fruits. Erectness is related to flower size; small flowers are ascending
to erect whereas larger ones are suberect to spreading. The original
figure of L. microcarpus showed ascending flowers, but the South

1988] RIGGINS: LUPINUS GROUP MICROCARPI i)

TABLE 3. RANGE AND MEAN VALUES FOR SOUTH AMERICAN AND CALIFORNIAN
SPECIMENS OF Lupinus GROUP Microcarpi. All measures are in mm. n = number of
specimens.

South American Californian

Variable n Range Mean Range Mean
Leaflet number 72 6-11 8.3 5-12 9.3
Leaflet width 71 2.0-12.0 aya 2.0-12.5 6.9
Leaflet length 7p 8.0-40.0 Dba 9.5—59%5 29.6
Petiole length Ti W3.0=120:0> 61.0 15.0-221.0 106.5
Peduncle length 72 15.0-150.0 65.7 19.0-320.0 140.1
Length between verticils | and 2 72 7.0-40.0 17.8 8.0-50.5 22.4
Bract length ie. 2.5-8.5 5.0 3.5-12.5 6.3
Pedicel length 73 05-325 1.4 0.5-5.0 1g
Upper calyx lobe length 73 2:2—5.0 37 1.1-7.8 4.0
Lower calyx lobe length 73 5.4-10.0 of: »2- 10:7, 7.9
Banner length, base to flexion 74 4.5-8.0 3.8 4.2-10.6 6.8
Banner length, flexion to apex 74 4.5-8.2 ae 3.7-9.7 6.5
Banner width, flexion to margin 74 1.4-3.5 ZS 1.3-6.2 oo,
Wing width 74 2.7-5.3 3.9 2.2-7.8 4.9
Wing length 74 9.0-14.4 11.7 8.8-17.7 13.6
Keel length 73 9.2-14.0 ree 8.4-17.7 29
Keel width WS 0.9-1.5 Il 0.9-2.9 ai

American specimens exhibit as much variation in this feature as
California plants.

Flower color in L. densiflorus varies from white to yellow, to pink
and rose, and to lavender and purple. Often the amount of pink or
purple varies in the wing and banner petals so that overall flower
color is not easily described. Yellow and white flowers are generally
restricted to populations of L. densiflorus, but all degrees of pink to
purple are found in other members of the complex. Yellow, pink,
and purple are generally intensified in dried specimens, but retention
of original color is related to duration and method of drying. Some-
times flowers fade to a straw color on drying. The South American
specimens do not appear to have flower colors different from Cal-
ifornia plants. Although the original description of L. microcarpus
referred to blue flowers, all subsequent authors have described them
as rose or lavender. I have not seen any specimen of the group that
appears to have blue flowers typical of other lupine species.

Data analyses included a tabulation of minimum, maximum, and
mean values for each variable. All variables could not be measured
from some South American specimens, so the mean values were
based on a varying number of observations (n) as given in Table 3.
In addition, multigroup discriminant analysis and diagnosis were
carried out as described in BIOSTAT II (Pimentel and Smith 1985).
With these methods discriminant analysis is first performed on pop-

98 MADRONO [Vol. 35

ulation samples, and then each individual of uncertain affinity is
assigned to a population of the discriminant analysis by an a pos-
teriori Geisser classification procedure. In this study discriminant
analysis was performed on the samples from California, and the
diagnosis on the South American specimens. Each South American
specimen was assigned to a population sample from California. Be-
cause missing data are not allowed for these analyses, four South
American collections (numbers 7, 18, 20 and 48) were excluded and
the diagnosis was performed on 69 of the specimens indicated in
Table 1. For these analyses the data were log transformed.

RESULTS

Geographic distribution. All South American specimens I exam-
ined are from Argentina and Chile. Smith’s (1941) report of L.
microcarpus from Peru was based on Weberbauer 148 (Dpto. Lima,
inter Matucana et Chanpothio, 26 Dec 1901, B) a specimen pre-
sumably destroyed.

Chilean plants occur along the coast from Taltal (25°26’S, Prov.
Antofagasta) to Valdivia (39°49’S, Prov. Valdivia), and inland from
Rio Sancarron (29°33'S, Prov. Atacama) to Temuco (38°44'S, Prov.
Cautin). Approximately one-third of the specimens I examined were
collected before 1900, many from areas near ports. Precise locality
and habitat data are often scanty but are sufficient for the following
ecological characterization. The Chilean plants grow in sandy soils,
rocky places and grasslands from the coast to the Andes at elevations
from near sea level to 600 m. A few specimens were collected along
the western slope of the Andes at reported elevations of 2300 to
3500 m.

Argentinean plants occur from latitude 33°S in Prov. Mendoza to
latitude 46°S near the southern border of Prov. Chubut. They grow
in the same kinds of habitats as in Chile, but are regarded as rare
and introduced (Planchuelo 1978).

In North America members of the L. densiflorus complex occur
near the coast from San Diego Co. (32°N) to Humboldt Co., Cali-
fornia (40-41°N), and disjunctly near Victoria, British Columbia
(48°N). Inland localities extend from Sierra de Juarez, Baja Califor-
nia Norte (31°N) to central Washington (45°N). Within this range
they are most abundant in California between latitudes 34°N and
38°N, from the coast eastward to the Sierra Nevada foothills. In
central California these lupines grow primarily in sandy soils of
valleys and low hills at elevations from near sea level to 1500 m.
They are most abundant in roadside and intermittent streamside
habitats, but also occur in grasslands and desert washes. They do
not occur at elevations above 1550 m, nor east of the Sierra Nevada.

These distribution records, my field observations, and informa-

1988] RIGGINS: LUPINUS GROUP MICROCARPI 09

TABLE 4. ACTUAL AND PERCENTAGE OCCURRENCE OF WING AND KEEL CILIATION
STATES IN SOUTH AMERICAN AND CALIFORNIAN SPECIMENS OF Lupinus GROUP Micro-
carpl.

South American Californian
n % n %
Wing ciliation:
0, absent 21 28 175 Zi
1, present above 46 67 539 66
2, present above and below 2 5 105 13
Keel ciliation:
0, absent below 67 90 542 66
1, present below 7 10 217 34

tion from the literature, indicate that plants from both hemispheres
occur generally within the same latitudes and elevations, and in
similar habitats. Both areas of distribution have Mediterranean cli-
mates and are well-known for their disjunct ranges of closely related
species (Raven 1963).

Morphological comparisons. Table 3 shows that the South Amer-
ican specimens are often smaller, particularly in vegetative features,
than the California specimens. South American specimens have a
narrower range of variation than those from California, but generally
exhibit a range of variation within that of the California specimens.
Minimum values for five vegetative measurements were recorded
from South American plants, but all maximum values were from
California plants.

For all variables, South American specimens have smaller mean
values than California specimens. Differences in mean values are
particularly striking for the petiole and peduncle measurements.
Differences in the mean values for the floral variables are less ap-
parent. Except for wing and keel petal lengths, the differences be-
tween the two groups is <1 mm.

Results for wing and keel ciliation features are given in Table 4.
Fewer South American specimens have cilia present on both margins
of the wing and keel petals.

Although the aim of this paper is to determine if the South Amer-
ican representatives are distinct, some understanding of variation
and discrimination of the California samples is necessary to clarify
the relationships. Figure | portrays the results of the discriminant
analysis of the California specimens on canonical axes | and 2 that
respectively represent 47% and 16% of the differences between the
samples. Vectors of variables contributing to ordination of the sam-
ples indicate that separation on axis 1 is primarily due to floral

[Vol. 35

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1988] RIGGINS: LUPINUS GROUP MICROCARPI 101

variables, measurements of the calyx, and of the banner and keel
petals. Samples on the right side of the graph have longer and wider
keel petals, wider banner petals and longer upper calyx lobes than
those on the left side of the graph. Separation on axis 2 is mostly
due to vegetative features of peduncle, petiole and verticil lengths.
Samples on the upper half of the graph are taller, whereas those on
the lower half have smaller leaves. Ordination along axis two roughly
parallels an east-to-west climatic gradient of arid-to-mesic habitats;
samples on the lower half were collected in the most arid habitats.

Clear or tight clusters of the California samples are not detectable
in Fig. 1. I initially identified the majority on the right side as L.
densiflorus, and those on the lower right as L. horizontalis. The
remote samples on the lower left were initially identified as L. ruber,
and those on the upper left as L. subvexus. Several samples near the
middle of the graph were identified as intermediate between L. den-
siflorus and L. subvexus, or intermediate between L. subvexus and
L. ruber, and were collected in areas of sympatric distribution (Hoo-
ver 1970).

South American specimens were assigned to 15 samples desig-
nated by stars in Fig. 1. All except sample //63 are on the left side
of the graph, and are samples that were identified as L. ruber, L.
subvexus or intermediates between them. Sample //63 was initially
identified as L. densiflorus. The 15 samples are from interior local-
ities and more arid habitats than those not involved in the assign-
ments.

Results of probability assignments for the South American spec-
imens are summarized in Table 5. The probabilities ranged from
19% to 97% and averaged 51.4%. For 61 of 69 South American
specimens, assignment to a specific California sample was evident;
1.e., resemblance to any other sample was remote. Eight South Amer-
ican specimens (17c, 22, 26a, 31, 33, 36b, 38, 51b as identified in
Table 1) had close affinities (<2%) to two different California sam-
ples; in each case the two samples were from nearby localities and
like habitats. Forty (58%) of the South American specimens were
assigned to just three California samples: 1/42, 1162 and 1172.

The California samples show a clinal pattern of geographic vari-
ation (Fig. 1), but there is no evidence of a similar pattern among
the South American specimens. This could be a reflection of inad-
equate sampling, although the specimens are from localities that
represent the geographic range and ecological zones where they occur

—

Fic. 1. Plot of California samples of Lupinus group Microcarpi on canonical axes
1 and 2. South American specimens were assigned to those designated by stars. Vectors
of variables contributing to the ordination are also plotted.

102 MADRONO [Vol. 35

TABLE 5. DIAGNOSIS ASSIGNMENTS OF SOUTH AMERICAN SPECIMENS (S. Am.) TO
CALIFORNIA (Calif.) POPULATION SAMPLES OF Lupinus GROUP Microcarpi. Column I
refers to diagnosis based on 19 variables; Column II to that based on 17 variables.

Calif. Calif. Calif.

S.Am. I (%) I S.Am. I (%) I S.Am I (%) UJI

1 1162 (65) 2la 1158 (84) 1156 39 1186 (57)

2 1172 (95) 21b 1158 (78) 1152 40 1150 (46)

3 1185 (29) 1140 22 1163 (23) 41 1172 (36) 1162

4 1172 (54) 23-1154 (33) 42a 1141 (19)

5 1154 (48) 24 1152 (36) 42b 1142 (57)

6 1142 (26) 1150 25 1141 (54) 43 1142 (54) 1145

8 1156 (38) 1162 26a 1172 (51) 44 1162 (53) 1141

9 1162 (57) 26b 1140 (25) 1154 45 1142 (56) 1145
10a «1154 (31) 1145 26c 1162 (66) 46 1142 (49) 1145
10b 1156 (76) 26d 1172 (75) 47a 1142 (38) 1145
11 1150 (65) 1154 27 1162 (67) 1141 47b 1172 (63) 1145
12 1162 (55) 28 1162 (76) 47c 1172 (85)

13 1162 (97) 29 1185 (43) 1186 49a 1172 (41) 1141
14 1162 (80) 30 1142 (66) 49b 1172 (79)

15 1162 (34) at 4145023) 50 =: 1150 (58) 1145
16a: 1162 (67) 32-1172 (69) Sla 1142 (42) 1145
16b 1156 (58) aa ea 51b 1185 (21) 1141

17a 1158 (44) 1162 34 1142 (26) 1143 aD 1150 (58)
17b 1162 (35) 1141 35 T172 (92) 1145 53 1162 (57)

17c 1158 (46) 1141 36a 1162 (84) 54a 1154 (27) 1150
17d 1172 (24) 1141 = 36b-s 1145 (23) 54b 1172 (48) 1141
17e 1172 (30) 1141 37 1144 (38) 55 1142 (80)

19 1162 (87) 38 1145 (25) 56 1162 (80) 1141

in South America. Geographical variation among the South Amer-
ican plants would be detected by a differential affinity to the Cali-
fornia plants; i.e., specimens would be assigned to populations from
similar climatic and ecological zones in California. The South Amer-
ican specimens, however, were identified with a few samples from
arid interior localities, the majority to three samples. Comparison
of the 15 South American specimens assigned to sample //72 il-
lustrates that they are from localities of latitudinal and elevational
extremes. They include specimen 2 from Prov. Chubut, Argentina
at latitude 46°S, specimens /7d and /7e from Taltal, Chile at latitude
25°S, specimen 48 from Prov. Valparaiso, Chile at elevation 10 m,
and specimens 26a and 26d from Prov. Coquimbo, Chile at elevation
3500 m. These results suggest that the South American plants exhibit
a more mosaic pattern of variation than the California plants.

As shown in Table 3 South American specimens have shorter
peduncles and petioles than California plants. Because these two
variables were involved in the discriminant analysis (Fig. 1), as-
signment of the South American specimens could be influenced by
the discrepant values. To test this hypothesis, a second diagnosis

1988] RIGGINS: LUPINUS GROUP MICROCARPI 103

was performed with these variables deleted. Assignment of 36 South
American specimens was to the same sample as the previous analysis
(Table 5). The assignments were to 14 samples, 13 in common with
the previous diagnosis and an additional one (/ 1/43). There was some
variation in the number of South American specimens assigned to
the particular California samples, but the overall pattern of assign-
ment did not change. These results show that the widely varying
vegetative features do not influence the assignments of the South
American specimens.

DISCUSSION AND CONCLUSIONS

Comparison of the disjunct representatives of Lupinus group Mi-
crocarpi reveals that vegetative structures are smaller in South Amer-
ican plants. As shown by the discrepant values for peduncle length,
this size difference is ascribable to plant height. Two explanations
can be advanced for the difference; one concerns environment and
growing conditions, and the other collecting practices and sampling
methodology.

Smith (1918a) pointed out that size and degree of branching of
these lupines are a reflection of the plant’s environment. Short,
unbranched plants are generally found in arid habitats whereas tall,
well-branched plants are generally found in more mesic environ-
ments. I have observed that plant size at any given locality can vary
from year to year depending on relative amount and periodicity of
precipitation and temperature extremes. The samples of California
populations are from a variety of habitats and were made during
favorable years, but collection of individual plants was by random
sampling. There is no reason to assume that the South American
specimens were collected from less favorable habitats or during less
favorable years, but herbarium specimens must be viewed, in an
analytical sense, as representing biased samples.

I think the small size of the South American specimens is most
likely attributable to past collecting practices. The majority of spec-
imens were collected before 1900, and several during early botanical
expeditions to South America. It is reasonable to assume that early
collectors were concerned with obtaining as many specimens as pos-
sible with limited equipment and facilities, and consequently col-
lected mostly small individuals.

The comparison reveals only slight differences in floral features
between North and South American plants. The range of variation
observed in South American specimens is within that of the Cali-
fornia samples, but the mean values of the South American plants
are slightly smaller. The difference in mean values is attributable to
relative abundance of large flowered L. densiflorus and L. horizon-
talis among the California samples. Fewer South American speci-
mens have cilia present on both margins of the wing and keel petals.

104 MADRONO [Vol. 35

These features are observed more frequently in L. densiflorus and
L. horizontalis.

Although some South American specimens can be readily iden-
tified as L. densiflorus, the majority are more similar to California
populations of L. subvexus, L. ruber or intermediates between them.
The results clearly demonstrate that South American representatives
of group Microcarpi are not distinct from some North American
representatives. The implications of these results will be addressed
in a forthcoming revision of group Microcarpi.

ACKNOWLEDGMENTS

I thank the curators and personnel of BM, CAS, DS, GH, K, MO, RSA, UC and
US for providing loans and/or space for study. Nancy Arnold, Rosemary Bowker,
Melissa Luckow, and Julie Vanderwier assisted in data collection. I thank Raul Cano
for translating the abstract into Spanish. I especially thank Richard Pimentel for his
assistance in the field and encouragement throughout the study. I also thank Duane
Isely, William Weber, Teresa Sholars, Wayne Ferren, and Dave Keil for their helpful
reviews. Part of the research was funded by CARE grant C81-11, California Poly-
technic State University.

LITERATURE CITED

Dunn, D. B. and J. M. GILLeTr. 1966. The lupines of Canada and Alaska. Canada
Dept. Agric. Monograph No. 2.

HitcHcock, C. L., A. CRONQUIST, M. OWNBEY, and J. W. THOMPSON. 1961. Vascular
plants of the Pacific Northwest. Part 3: Saxifragaceae to Ericaeae. Univ. Wash-
ington Press, Seattle.

Hoover, R. L. 1970. The vascular plants of San Luis Obispo County, California.
Univ. California Press, Berkeley.

JEPSON, W. L. 1936. A flora of California. Vol. 2. Associated Students Store, Univ.
California, Berkeley.

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

PIMENTEL, R. A. and J. D. SmitH. 1985. Biostat II. Sigma Soft, Placentia, CA.

PLANCHUELO, A. M. 1978. A monograph of Lupinus for Argentina. Ph.D. disser-
tation, Univ. Missouri, Columbia.

RAVEN, P. H. 1963. Amphitropical relationships in the flora of North and South
America. Quart. Rev. Biol. 38:151-177.

SmiTH, C. P. 1917. Studies in the genus Lupinus—I. A new species of the subgenus
Platycarpos. Bull. Torrey Bot. Club 44:405—406.

1918a. Studies in the genus Lupinus—II. The Microcarpi, exclusive of

Lupinus densiflorus. Bull. Torrey Bot. Club 45:1-22.

1918b. Studies in the genus Lupinus—III. Lupinus densiflorus. Bull. Torrey

Bot. Club 45:167—202.

. 1919. Studies in the genus Lupinus—IV. The Pusilli. Bull. Torrey Bot. Club

46:389-410.

1940. The genus Lupinus in Chile. Species Lupinorum, Signature 13.

1941. A first report on the genus Lupinus in Peru. Species Lupinorum,

Signature 17.

1943. The genus Lupinus in Argentina. Species Lupinorum, Signature 21.

1944. Lupinus. In L. Abrams, Illustrated flora of the Pacific States. Vol. II.

Polygonaceae to Krameriaceae, p. 483-519. Stanford Univ. Press, Stanford, CA.

(Received 10 Mar 1987; revision accepted 25 Jan 1988.)

LEPTODACTYLON PUNGENS SUBSP. HAZELIAE
(POLEMONIACEAE), A NEW COMBINATION FOR
A SNAKE RIVER CANYON ENDEMIC

ROBERT J. MEINKE
Department of Botany & Plant Pathology,
Oregon State University,
Corvallis 97331-2902

ABSTRACT

Morphological study of herbarium and living specimens of Leptodactylon from the
Pacific Northwest and northern Great Basin indicates that the little-known species
Leptodactylon hazeliae Peck is more appropriately treated as a subspecies of L.
pungens (Torr.) Rydb. This rare taxon occurs only in the Snake River Canyon of
Oregon and Idaho, where it inhabits sheer rock outcrops in Poa-Agropyron-Purshia
communities. The relationship of subsp. hazeliae to other species of Leptodactylon
is not clear. The subspecies is morphologically intermediate between L. pungens and
L. watsonii (A. Gray) Greene, a trait it shares with the recently described L. glabrum
Patterson and Yoder-Williams, an epilithic species of northern Nevada and adjacent
Idaho. It is speculated that L. glabrum and subsp. hazeliae may have originated from
past hybridization events involving L. pungens and L. watsonii.

Leptodactylon H. & A. is a small genus of suffrutescent perennials
and low subshrubs endemic to western North America (Grant 1959,
Cronquist 1984). It is primarily distributed in arid regions east of
the Cascade-Sierran axis, but also occurs in the coastal mountains
and maritime areas of southern California and adjacent Mexico. Its
members are reminiscent of perennial species of Phlox and Li-
nanthus, but are distinguished from those genera by the combination
of prickly leaves, prominent membranes in the calyx sinuses, and
equally inserted stamens.

The generic limits of Leptodactylon are relatively well-marked
morphologically, but the delineation of species and infraspecific taxa
has been historically a source of frustration for students of Pole-
moniaceae. Several species exhibit considerable phenotypic plastic-
ity, often with patterns of regional variation that are difficult to
circumscribe. In the only comprehensive treatment of the genus,
Wherry (1945) recognized 10 species and several varieties and for-
mae. Unfortunately, the relationships between morphology and geo-
graphic range presented in his account are vague, and the taxonomy
has not been readily adaptable for use in state or local floras. Later
workers (Davis 1950, Mason 1951, Harrington 1954, Cronquist
1959, 1984, Munz 1959, Kearney and Peebles 1960) reduced the
number of accepted species to six or seven and elected to disregard

MADRONO, Vol. 35, No. 2, pp. 105-111, 1988

106 MADRONO [Vol. 35

those proposed by Wherry (1945) that intergrade extensively across
broad geographic zones. Some authors advocate the use of infra-
specific categories to accommodate these variants. However, in lieu
of detailed phytogeographic studies, the application of these names
in any given area Is largely conjectural.

Most of the variation in Leptodactylon is represented in a complex
centered around L. pungens (Torr.) Rydb., one of three widespread,
polymorphic species that range throughout all or much of the In-
termountain Region and American Southwest. Fourteen of the twen-
ty Leptodactylon taxa recognized by Wherry (1945) are segregates
of L. pungens, and the species has a lengthy synonymy of over forty
nomenclatural combinations dating to the early nineteenth century
(Cronquist 1959, 1984). The other two intermountain species, L.
watsonii (A. Gray) Greene and L. caespitosum Nutt., share a number
of traits with L. pungens, and in some respects the three entities
constitute a morphological continuum. Populations of the three
species also overlap along an ecological gradient, and may coincide
geographically, particularly in the eastern Great Basin and Wyoming.
Despite the morphological similarities that imply common ancestry,
and the high potential for sympatry, there are no available data to
suggest that these species are interfertile. The two characters that
typically are used to identify them, 1.e., phyllotaxy and the number
of flower parts, show remarkable consistency considering the overall
variability of the groups. Leptodactylon caespitosum and L. watsonli
are opposite-leaved and have 4-merous and 6-merous flowers, re-
spectively, whereas L. pungens has 5-merous flowers and variable
leaf insertion, usually with the upper alternate and the lower op-
posite. In the few cases where 5-merous flowers occur in L. watsonii
(Cronquist 1984, Meinke pers. observ.), the plants are distinguished
from the woodier L. pungens on the basis of flexible, subherbaceous
flowering stems and opposite leaves throughout, including the in-
florescence bracts.

Because of the consistency of the aforementioned differences, it
is noteworthy that populations of Leptodactylon have been discov-
ered recently that do not fit patterns of variation previously described
for L. pungens or L. watsonii. The plants are located in the Snake
River Canyon of northeastern Oregon and adjoining Idaho, a locality
rich in disjunct and endemic species (Peck 1961). The most striking
features of living specimens of the riverine populations are the short
inflorescence branches, the pliable, bright green leaves, and the diur-
nal flowering that contrasts with the mostly vespertine corolla ex-
pansion of other Pacific Northwest Leptodactylon taxa.

My preliminary study showed that the plants correspond mor-
phologically with the only known collection of L. hazeliae Peck, a
taxon described in 1936 from three immature branches gathered

1988] MEINKE: LEPTODACTYLON PUNGENS SUBSP. HAZELIAE 107

near the Snake River. Subsequent examination of several hundred
herbarium specimens of L. pungens and L. watsonii supports the
taxonomic recognition of the Snake River populations based on
several minor but constant morphologic traits (Table 1). These plants
are apparently ecologically specialized as well, being restricted to the
uniquely mild (for the region) climate of the Snake River Canyon.
The number of unambiguous characters separating the Snake River
plants from other populations of L. pungens are fewer than the
number distinguishing L. pungens from other species in the genus.
I propose that these populations be recognized at the level of sub-
species, under L. pungens. The spelling of the subspecific epithet in
the following new combination reflects an orthographic correction
from Peck’s (1936) original ‘‘Hazelae’, after Recommendation
73C.1b of the ICBN.

Leptodactylon pungens (Torr.) Rydb. subsp. hazeliae (Peck) Meinke,
stat. et comb. nov. (Fig. 1)—Leptodactylon Hazelae Peck, Proc.
Biol. Soc. Wash. 49:111. 1936; L. pungens subsp. hookeri (Doug.
ex Hook.) Wherry forma hazelae (Peck) Wherry, Amer. Mid].
Naturalist 34:383. 1945.—Type: USA, OR, Wallowa Co., dry
rocky slope, Snake River Canyon near mouth of Battle Creek,
13 Apr 1934, Barton s.n. (Holotype: WILLU 184]5!).

Additional specimens. USA, ID, Idaho Co.: Snake River Canyon,
4 mi downstream from Granite Creek, local on cliffs, 22 May 1974,
Henderson, Wellner, and Bingham 1306 (ID!, two sheets); Snake
River Canyon, Suicide Point, on trail near U.S. Forest Service sign,
15 Jun 1978, Mattson and Bishoff s.n. (IDF!). Adams Co.: Snake
River Canyon, ca. 5 kms. of Hell’s Canyon Dam, along Idaho Power
Company right-of-way, 20 Apr 1977, Meinke 1545 (OSC)).

Habitat. Leptodactylon pungens subsp. hazeliae occurs below 650
m, inhabiting rock walls and talus covered slopes. It has only been
recorded from the deepest part of the Snake River Canyon, between
latitudes 45° and 46°N, and is not known to be sympatric with any
other species or subspecies of Leptodactylon. The vegetation in this
area 1s dominated by Poa sandbergii Vasey, Agropyron spicatum
(Pursh) Scribn. & Smith, Purshia tridentata (Pursh) DC., and Celtis
reticulata Torr. Other endemic taxa occurring with subsp. hazeliae
include Rubus bartonianus Peck, Ribes cereum Dougl. var. colu-
brinum Hitchc., Phlox colubrina Wherry & Const., Astragalus cu-
sickii Gray, A. vallaris Jones, Nemophila kirtleyi Hend., and Hack-
elia hispida (Gray) Johnst.

Floral phenology. Branch development is initiated in late February
or early March, with flowering occurring from April through June.
Inflorescences consist of one to three flowers [not strictly single-

[Vol. 35

~

MADRONO

108

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Fic. 1. Leptodactylon pungens subsp. hazeliae. A. Flowering stem. B. Enlargement
u ubescence.

illustrating the glandular stem, leaf, and calyx p

110 MADRONO [Vol. 35

flowered as reported by Peck (1936, 1961)]. Corollas generally re-
main open throughout the day and produce copious nectar. Insect
visitors are primarily pierid and lycaenid butterflies and (occasion-
ally) sphingid moths. Capsules are tardily dehiscent and persistent,
with seed dispersal occurring throughout the summer.

Relationships. The taxonomic relationship between subsp. haze-
liae and its potential relatives is difficult to assess. By tradition, the
possession of pentamerous flowers and at least some clearly alternate
leaves implies afhliation with L. pungens. However, the sprawling
habit, herbaceous flowering stems, and sparsely flowered inflores-
cences are suggestive of L. watsonii (Table 1). Moreover, the presence
of 5-merous flowers is not unprecedented in L. watsonii, being known
from a few scattered localities such as the Quinn Canyon Range of
central Nevada (Cronquist 1984). The habit and stem traits also are
shared by L. glabrum Patterson and Yoder-Williams, a recently
described species that occurs at two sites in northwestern Nevada
and southwestern Idaho. Leptodactylon glabrum is considered closely
allied with L. watsonii because of its completely opposite phyllotaxy
and strictly 6-merous flowers (Patterson and Yoder-Williams 1984).
It is possible that subsp. hazeliae has a close affinity with L. glabrum
because it is the only other intermountain congener with soft, fili-
form-linear leaflets less than 0.5 mm broad. Furthermore, subsp.
hazeliae plants also, on occasion, possess a few 6-merous flowers.
Both entities are apparently restricted to rocky habitats isolated in
steep canyons, and neither are known to be sympatric with other
members of the L. pungens or L. watsonii complexes. Although leaf
insertion and the number of flower parts will generally discriminate
the two taxa, there are distinctive pubescence differences as well.
Leptodactylon glabrum is eglandular and often glabrous (Patterson
and Yoder-Williams 1984), whereas L. pungens subsp. hazeliae is
stipitate-glandular on the stems, leaves, and calyces (Fig. 1B). Many
of the morphological characters that distinguish L. glabrum and
subsp. hazeliae within the genus are intermediate between L. wat-
sonii and L. pungens. This, coupled with their narrow geographic
distributions, suggests that the two endemics could be remnants of
past intergradations between L. watsonii and L. pungens in areas
where these widespread species no longer coexist.

ACKNOWLEDGMENTS

I thank Kenton Chambers, Teresa Magee, Joseph Antos, Robert Frenkel, and J.
Stephen Shelly for helpful comments and advice, and Knut Noraas for his excellent
job on the line illustrations. I am also grateful to the curators and staff of CIC, HSC,
ID, IDF, LAGO, ORE, OSC, RSA, UNLV, WILLU, and WS for the loan of specimens
and the use of facilities. The Oregon State University Herbarium provided technical
and financial support. This paper represents Technical Contribution No. 1 of the
Endangered Species Program, Oregon Department of Agriculture, Salem.

1988] MEINKE: LEPTODACTYLON PUNGENS SUBSP. HAZELIAE Jit

LITERATURE CITED

CRONQUIST, A. 1959. Polemoniaceae. Jn C. L. Hitchcock, A. Cronquist, M. Ownbey,
and J. W. Thompson, Vascular plants of the Pacific Northwest 4:95-145. Univ.
Washington Press, Seattle.

1984. Polemoniaceae. Jn A. Cronquist, A. Holmgren, N. Holmgren, J. L.
Reveal, and P. K. Holmgren, Intermountain flora 4:86—155. New York Botanical
Garden, Bronx.

Davis, R. J. 1950. Flora of Idaho. W. C. Brown Company, Dubuque, IA.

GRANT, V. 1959. The natural history of the phlox family. Martinus Niyhoff, The
Hague.

HARRINGTON, H. D. 1954. Manual of the plants of Colorado. Sage Books, Denver.

KEARNEY, T. H. and R. H. PEEBLES. 1960. Flora of Arizona. Univ. California Press,
Berkeley.

Mason, H.L. 1951. Polemoniaceae. Jn L. R. Abrams, Illustrated flora of the Pacific
States 3:396-474. Stanford Univ. Press, Stanford, CA.

Munz, P. A. 1959. A California flora. Univ. California Press, Berkeley.

PATTERSON, R. and M. YODER-WILLIAMS. 1984. Leptodactylon glabrum, a new
intermountain species of Polemoniaceae. Syst. Bot. 9:261-—262.

Peck, M. E. 1936. Six new plants from Oregon. Proc. Biol. Soc. Wash. 49:111.

1961. A manual of the higher plants of Oregon, 2nd ed. Oregon State Univ.
Press, Corvallis.

Wuerry, E. T. 1945. Two linanthoid genera. Amer. Midl. Naturalist 34:38 1-385.

(Received 9 Mar 1987; revision accepted 26 Jan 1988.)

ANNOUNCEMENT

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REVIEWS SOLICITED

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reviewed journal specializing in the publication of papers with a regional
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Beginning with Volume 88, the Bulletin will include solicited review
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Persons interested 1n writing a review should send an outline of the
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topics in need of review. Send topic suggestions and names of potential
authors to the technical editor, Dr. Jon E. Keeley, Editor, Department
of Biology, Occidental College, Los Angeles, CA 90041.

SCUTELLARIA LUTILABIA (LABIATAE),
A NEW GYPSOPHILE FROM NUEVO LEON, MEXICO

THOMAS M. LANE
P.O. Box 327, Philo, CA 95466

Guy L. NESOM
Department of Botany, University of Texas,
Austin 78713

ABSTRACT

Scutellaria lutilabia, an endemic skullcap from an area of gypsum outcrops on the
west side of Cerro Pena Nevada, Nuevo Leon, Mexico, is described. It appears to be
a member of sect. Resinosae Epling but differs strongly from all species of that section
by its vestiture, flower color, and mericarp morphology.

RESUMEN

Scutellaria lutilabia, una capa calavera de una afloramiento de rocas yesosas ex-
puestas en el parte Oeste del Cerro Pena Nevada, Nuevo Leon, México, esta descrita.
Parece ser un miembro de la secc. Resinosae Epling, pero se diferencia marcadamente
de todos las especies de esa secciOn por su indumento, el color de sus flores, y su
morfologia mericarpica.

Intensive collecting on gypsum outcrops in northeastern Mexico
has brought to attention a new species of Scutellaria.

Scutellaria lutilabia Lane & Nesom, sp. nov.

Inter species sect. Resinosae Epling sed caulibus, foliis, pedicellis,
et calycibus argenteo-virides cum pilis brevi-sericeis densis vestita.
Corolla atrosanguinea, labio inferiore cum macula lutea, et meri-
carplis tuberculatis et brevi-aculeatis distingueda (Fig. 1).

Rounded to ascending, strongly taprooted, subshrubs 7—30 cm tall
and to 70 cm wide, branched especially in the lower half; stems
many, arising from an expanded and branched woody crown; stems,
leaves, pedicels and calyces silver-green, densely clothed in a seri-
ceous, antrorse vestiture of short, appressed white hairs to 0.25
(-—0.5) mm long (Figs. 1-3); sessile glands present but obscured by
the hairs. Leaves opposite, elliptic to ovate, 5—15 mm long, 3-7 mm
wide, twice as long as wide, basally attenuate to a short petiole 1-2
mm on upper leaves and up to 5 mm on lower leaves, midrib and
several pairs of lateral veins raised and prominent beneath, apex
obtuse, margins slightly revolute, entire. Flowers solitary in axils,
without subtending bracteoles; pedicels 2-4 mm long. Calyces

MaprONo, Vol. 35, No. 2, pp. 112-116, 1988

LANE AND NESOM: NEW SCUTELLARIA 113

1988]

and flower.

’

Scutellaria lutilabia. A. Habit. B. Leaves, stem

Fic. 1.

114 MADRONO [Vol. 35

2-lipped, 4-5 mm long, accrescent and closing in fruit; upper lip
expanded upwardly 3 mm into a dorsally impressed, shield-like
scutellum and falling at maturity with the mericarps; lower lip dish-
like, persistent. Corolla sigmoid in outline, 15—20 mm long, maroon,
and densely villous on the outer surface with hairs to 1 mm long,
bilabiate; lower lip 3-lobed, ca. 8 mm long and 8 mm wide with a
median yellow-gold blotch starting just behind the tip and running
into the throat; upper lip galeate, ca. 7 mm long, flaring and revolute
at both axial edges, with some scattered villosity within; tube 10
mm long, ca. 2 mm wide, densely villous inside at base; stamens
didynamous, included within the galea, basal anterior pair adnate
ca. two-thirds their length and ultimately longer, basal posterior pair
adnate ca. three-fourths their length; anthers maroon-purple, densely
short-ciliate, with sessile glands along the line of attachment to the
filament; gynoecium with an elevated 4-lobed ovary, free and gyno-
basic style, and a linear, 2-parted stigma inserted between the 2
anther pairs. Fruit (Figs. 4—5) 4 brownish-black, ovoid mericarps
1.5 mm long, | mm wide, surface with tubercules ca. 90 x 90 wm
with linear-lanceolate, outwardly elongated apical cells, giving the
whole mericarp a densely and minutely hispidulous appearance.

TYPE: México, Nuevo Leon: ca. 30 km ene. of Doctor Arroyo,
21.5 km ene. of San Antonio de Pena Nevada, 23°46'N, 99°55’W,
2015 m, 3-5 Aug 1981 (fl, fr), Nesom 4273 (Holotype: MEXU;
isotypes: ARIZ, ASU, ENCB, NY, OS, US).

PARATYPES: México, Nuevo Leon: [type locality], 10 Jul 1984,
Michener 4428 with Prigge (GH, to be distributed); ca. 7 km ne. of
San Antonio de Pena Nevada, [1.3 km n. of the type locality], Jul
1977 (fl, fr), Wells and Nesom 513 (LL, MO, NCU).

Habitat and distribution. Scutellaria lutilabia is known only from
gypsum outcrops at an elevation of about 2000-2050 meters on the
northwest slope and west base of Cerro Pena Nevada. It is associated
there with Agave, Bauhinia, Cowania, Euphorbia, Krameria, Leu-
cophyllum, Lindleya, Mortonia, Nolina, and Opuntia, as well as
numerous herbaceous species.

Relationships. A suite of characteristics of the new species refer it
to the southwestern U.S.-Mexican sect. Resinosae Epling (Epling
1942): a xeric habitat; a taproot topped by a woody crown from
which arise numerous leafy stems; small, entire leaves; solitary flow-
ers in the axils of the upper leaves; a broad, impressed scutellum;
stamens attached near the center of the corolla tube; and mericarp
epidermal cells generally with a circular outline at their bases (Fig.
5) (Lane 1983). Features that clearly separate this new species from
other members of the section are its sericeous vestiture, densely
“‘*hispidulous’”’ mericarps, and maroon corollas with a golden-yellow
blotch on the lower corolla lip (from which the specific epithet is

1988] LANE AND NESOM: NEW SCUTELLARIA 1B ies)

Fics. 2-5. Scutellaria lutilabia. 2. Young leaf. Note nature of vestiture, scale =
500 um. 3. Detail of leaf vestiture, scale = 50 wm. 4. Mericarp lateral view, scale =
250 wm. 5. Detail of mericarp surface. Note elongated epidermal cells with hair-like
extensions, scale = 50 um. All from Wells and Nesom 513. Specimens processed via
standard techniques (cf. Lane 1983).

derived). Other species of sect. Resinosae have more sparse or at
least non-sericeous vestiture, tuberculate mericarps without apical
extensions on the tuberculae, and blue or violet-blue corollas with
a white blotch on the lower lip.

Scutellaria lutilabia stands apart morphologically from all other
described taxa of sect. Resinosae. Perhaps its closest relative is an
undescribed species from eastern Chihuahua (James Henrickson pers.

116 MADRONO [Vol. 35

comm.), which is similar in habit (though more condensed and
intricately branched), leaf morphology, and stem vestiture. In con-
trast, its flowers are blue with white markings and the mericarps are
granular-papillate, lacking the prominent hispidulous surface of S.
lutilabia. The papillae, however, are rough in appearance apparently
because of unevenly protruding epidermal cells and need to be ex-
amined in closer detail for possible homologies with those of S.
lutilabia.

Despite Epling’s reservation about the taxonomic value of meri-
carp morphology, this character has been used previously in the
delineation of species and species groups in Scutellaria (Lane 1983
and unpubl.). Many of Epling’s groups display a high degree of
homogeneity in the nature of their mericarp surface. Mericarp sur-
faces in American species of the genus have been described as smooth,
roughened, rugose, granulate, squamellate, mammillate, lacerate-
dentate angled, coarsely toothed, with peg-like processes, verrucose,
muricate, tuberculate, lamellate, and papillate (see Epling 1942 and
various floristic manuals). Epling described those of S. horridula
Epling (sect. Resinosae) as “‘horridulis”’; they have slender, strongly
projecting tuberculae but lack the hairlike extensions as in S. /uti-
labia. Almost all of these descriptions, however, are based on ob-
servations made with only low magnification. We predict that SEM
studies of mericarp surfaces will reveal similarities indicative of
recent common ancestry.

ACKNOWLEDGMENTS

Weare grateful to Maura McNulty for the illustrations, Stella Muller for the Spanish
translation, and the first author thanks Laurie Stull for affording him writing time.
SEM facilities were provided by the Department of Botany of the Ohio State Uni-
versity. James Henrickson made comments on the manuscript and generously pro-
vided information on undescribed species of Scutellaria. We thank David Michener
and Barry Prigge for information on their recent collection.

LITERATURE CITED

EpLING, C. 1942. The American species of Scutellaria. Univ. Calif. Publ. Bot. 20:
1-146.

LANE, T. M. 1983. Mericarp micromorphology of Great Plains Scutellaria (La-
biatae). Southw. Naturalist 28:71-79.

(Received 5 May 1987; revision accepted 28 Oct 1987.)

A NEW SPECIES OF CROTON (EUPHORBIACEAE)
FROM NICARAGUA

GRADY L. WEBSTER
Department of Botany, University of California,
Davis 95616

ABSTRACT

Croton nubigenus, a new species of cloud forests in northern Nicaragua, belongs
to sect. Tiglium and appears most closely related to Mexican species such as C.
adspersus, C. tremulifolius, and C. ynesae, as well as C. wilsonii of Jamaica.

RESUMEN

Croton nubigenus, una nueva especie de los bosques nublados del norte de Nica-
ragua, pertence a la secc. 7iglium y parece tener afinidades estrechas con algumas
especies Mexicanas como C. adspersus, C. tremulifolius, y C. ynesae, incluyendo a
C. wilsonii de Jamaica.

During the preparation of the treatment of the Euphorbiaceae for
the “Flora of Nicaragua” project at the Missouri Botanical Garden,
I encountered a number of specimens of a Croton from cloud forests
on peaks in Nicaragua. These specimens represent a species not
formerly described.

Croton nubigenus Webster, sp. nov. (Figs. 1-4)

Arbor dioica, foliis penninervuis subglabris, limbo basi subsessili-
biglanduloso, stipulis deltoidis, floribus receptaculo parce villosulo,
staminibus 13-15, ovario stellato-hirtello, stylis bifidis, caruncula
seminis obsoleta.

Dioecious shrubs or trees 3-5 m high; twigs and leaves nearly
glabrous (with appressed pauciradiate stellate hairs mainly on flow-
ers and young growth). Leaves with petioles 1.5—7 cm long; paired
glands at apex of petiole subsessile, 0.4—0.6 mm high and broad;
stipules deltate, 0.2-0.3 mm; blade chartaceous, oblong-lanceolate,
bluntly acuminate at tip, cuneate at base, glabrous on both faces;
venation pinnate, major lateral veins about 8-10 on a side, slightly
arching, veinlet reticulum distinct but tenuous beneath; margins
entire. Inflorescences terminal (and sometimes also at uppermost
adjacent nodes), racemose, unisexual, 5—18 cm long; pistillate flow-
ers solitary at nodes, 4—6 per raceme; staminate flowers 2—5 per
node; bracts deltate, entire, about 0.5 mm long. Staminate flower:
pedicel 2.5—4.5 mm long; sepals 5, valvate or slightly imbricate,

MADRONO, Vol. 35, No. 2, pp. 117-120, 1988

118 MADRONO [Vol. 35

Fics. 1-4. Photographs of Croton nubigenus. Fic. 1. Flowering branch of sta-
minate plant (Pipoly 6038). Fic. 2. Apical portion of pistillate inflorescence (Grijalva
313). Fic. 3. Base of leaf blade showing petiolar glands (Grijalva 313). Fic. 4. Adaxial
view of seed (Pipoly 6052). Bar equals 5 cm in Fig. 1, 1 cm in Fig. 2, 2.5 cm in Figs.
3 and 4.

1988] WEBSTER: NEW SPECIES OF CROTON 119

deltate-ovate, acute, glabrous or with very sparse appressed stellate
hairs, 1.8—2 mm long, 1.2—1.5 mm broad; receptacle sparsely villose;
petals 5, elliptic-spatulate, glabrous on both faces, short-villosulous
at tip and margins near base, 1.7—2 mm long, 0.5—0.8 mm broad;
stamens 13-15; filaments glabrous, anthers 0.6—0.8 mm long. Pis-
tillate flower: pedicel becoming 5—8.5 mm long, 0.8—1 mm thick,
subglabrous; sepals 5, lanceolate, entire, acute, basally connate, |.2—
1.5 mm long, subglabrous or sparsely appressed-stellate abaxially,
copiously hirsutulous (with simple hairs) adaxially; disk patelliform,
thickish, shallowly 5-lobed, subglabrous, 1.5—1.8 mm across; petals
obsolete (represented by tufted hairs); ovary ellipsoidal, densely stel-
late with more or less appressed 8—12-radiate hairs about 0.2—0.3
mm across; styles bifid, 2.5—3.5 mm long, branches slender. Capsule
not seen intact; columella slender, 8—8.5 mm long; seeds plump,
ellipsoidal, brownish, smooth, 7.3—8 mm long, 5.5—5.6 mm broad;
caruncle reduced or obsolete.

TyPeE: Nicaragua, Zelaya, primary cloud forest on summit of Cerro
La Piminenta, 900-980 m, 13°45'N, 84°59’W, 13 Apr 1979, Pipoly
5113 (Holotype: MO; isotype, DAV).

PARATYPES: Nicaragua, Zelaya, Cerro La Piminenta, Pipoly 6038,
6052 (DAV, MO); Cerro El Hormiguero, Grijalva 313, 462 (DAV,
MO); Cano El Hormiguero, Pipoly 6102 (DAV, MO).

This new species appears to be rather narrowly restricted to cloud
forests on peaks of a small area of the Cordillera Isabella near the
boundaries of the departments of Jinotega and Zelaya. In the last
complete revision of Croton by Mueller Argoviensis (1866), it would
key down near Croton wilsonii Griseb., a Jamaican species referred
by Mueller to series III of sect. Croton. According to the revision of
his supraspecific taxa of Croton in the “‘Flora Brasiliensis”’ (Mueller
1873), the position of Croton wilsonii would fall in sect. Croton,
subsect. Cleodora, ser. Medea. As noted by Bentham (1880) and
others, however, the sectional and subsectional taxa of Mueller are
defined arbitrarily and often are highly unnatural. It appears on the
basis of a number of common characters (discussed below) that C.
wilsonii and C. nubigenus should be referred to sect. Tiglium (K1.)
Baillon. Although Baillon (1858) included in his section the single
species C. tiglium L., a medicinal plant native to India, Mueller
(1866) recognized several related Asiatic and African species (with-
out, however, granting 7ig/ium any formal taxonomic recognition).

In the absence of any thorough revision of the genus Croton during
the century subsequent to Mueller’s monographic work, description
of new species has taken place with little appreciation of possible
biogeographic relationships. The relationships among species of sect.
Tiglium have been almost totally obscured by the complications of
fragmentation of effort and failure of the classical 19th century clas-
sifications to adequately reflect phylogeny.

120 MADRONO [Vol. 35

Croton nubigenus clearly belongs in the same section as C. wilsonii
because of its sparse indumentum of appressed stellate hairs, pen-
ninerved leaves biglandular at base, small entire sepals of pistillate
flowers and bifid styles. It differs from the Jamaican species, how-
ever, in its broader entire leaves, sexual condition (dioecious instead
of monoecious), larger stamen number, and distinctly pedicellate
pistillate flowers. Among species of mainland North America, C.
nubigenus shows some similarity to C. ynesae Croizat from western
Mexico. Croton ynesae nevertheless differs in many ways, including
coarsely serrate leaves, monoecious inflorescences, reduced pistillate
calyx, and carunculate seeds. In South America, there are a few
species that are suggestively similar to C. nubigenus, including C.
fraseri Muell. Arg. from Ecuador and C. sapiifolius Muell. Arg. from
Brazil. Mueller (1865, 1866) made the latter species the type of sect.
Quadrilobus Muell. Arg. because of the 4-merous flowers, but in
both vegetative and floral characters it somewhat resembles C. nu-
bigenus and C. wilsonii. If the American species of the ““Tiglium”’
alliance are treated as a section distinct from the Old World species,
then sect. Quadrilobus would be the correct name. However, al-
though our knowledge of these plants is still rather fragmentary, I
believe that it is better to recognize sect. Tig/ium in an inclusive
sense to include not only sect. Quadrilobus but also sect. Gymno-
croton Baillon (1858), based on the Australian C. verreauxii Baillon.
The widespread but fragmentary nature of the distribution of sect.
Tiglium (s. lat.) raises interesting biogeographical questions that can
only be answered by a revision of the sections of Croton, and a more
thorough study of the species putatively related to C. nubigenus.

ACKNOWLEDGMENTS

This new species of Croton was discovered during a winter interlude at the Missouri
Botanical Garden, where studies were made in collaboration with the Flora of Nic-
aragua project, directed by Dr. W. Douglas Stevens. I wish to thank Ms. Lynn Gillespie
for preparing the photographs.

LITERATURE CITED

BAILLON, H. 1858. Etude général du groupe des Euphorbiacées. Victor Masson,
Paris.

BENTHAM, G. 1880. Euphorbiaceae. Jn G. Bentham and J. D. Hooker, Genera
Plantarum 3:239-—340. L. Reeve & Co., London.

MUELLER, J. 1865. Vorlaufige Mitteilungen aus dem fur De Candolle’s Prodromus
bestimmten Manuscript. Linnaea 34:1—224.

1866. Euphorbiaceae Jn A. De Candolle, ed., Prodromus systematis na-

turalis regni vegetabilis 15(2):189-1261. Victor Masson, Paris.

. 1873. Euphorbiaceae [part 1]. Jn A. W. Eichler, ed., Flora Brasiliensis 1 1(2):

1-292. Munich.

(Received 27 May 1987; revision accepted 2 Dec 1987.)

A NEW LOMATIUM (APIACEAE) FROM THE
SIERRAN CREST OF CALIFORNIA

RONALD L. HARTMAN
Rocky Mountain Herbarium, Department of Botany,
University of Wyoming, Laramie 82071-3165

LINCOLN CONSTANCE
Department of Botany, University of California,
Berkeley 94720

ABSTRACT

Lomatium shevockii, a low, tufted, acaulescent species, 1s described from the south-
ern Sierra Nevada, Kern County, California. Although the growth habit is reminiscent
of some species of Oreonana and Cymopterus, the new species is clearly a Lomatium,
based on morphology and ecology. It appears closely related to Lomatium rigidum,
but differs strikingly by its flower color, subcapitate inflorescence, nearly prostrate
peduncles, smaller leaves, and subsessile fruit.

An exceedingly rare species of Lomatium was discovered by Mr.
Shevock on 7 April 1984 during a cursory field survey of Owens
Peak via the newly constructed section of the Pacific Crest Trail
north of Walker Pass. Although at that time only a few young leaves
had emerged, the collector correctly identified the plants as “‘new”
for the southern Sierra Nevada. At first glance the young leaves
resemble those of Oreonana clementis. Closer observation of the
blue-green, glabrous, white spinule-tipped leaves, however, resulted
in a tentative assignment to Cymopterus—a genus that has several
species with white-spinulose leaves. Subsequent collection of mature
fruit in June 1986, conclusively placed the species in Lomatium.

Lomatium shevockii Hartman & Constance, sp. nov. (Fig. 1)

Plantae perennes glabrae et glaucae acaulescentes 4-12 cm altae
e radice palari elongato cum caudice fibrilloso. Folia rosulata ovato-
deltoidea 1.5—4 cm longa, 2—5 cm lata, 2—3-pinnato-pinnatifida
divisionibus ultimis oblongis ovatisve saepe confluentibus acerosis;
petioli 1.5—5.5 cm longis scarioso-vaginantes. Pedunculi folia ae-
quantes excedentesve 4—12 cm longi; involucrum plerumque nul-
lum; radii 5-9 inaequales divergentes vel reflexi; umbellulae
andromonoeciae ex floribus perfectis 5—10+ et floribus staminatis
1—4 constantes; involucellum dimidiatum bracteolis 3—6 lanceolatis
vel ovatis integris distinctis 1—3.5 mm longis. Flores purpurei, sepalis

MADRONO, Vol. 35, No. 2, pp. 121-125, 1988

122 MADRONO [Vol. 35

Fic. 1. Lomatium shevockii. a. Habit. b. Foliage leaf. c. Leaflet. d. Fruiting umbel.
e. Flower. f. Fruit, dorsal view. g. Carpophore. h. Transaction of mericarp. (a, c, e
from Shevock et al. 11197; b, d, fh from the type collection.)

evidentibus triangularibus vel lanceolatis, petalis obovatis, antheris
flavis, stylis 2—-2.5 mm longis; stylopodium nullum, disco praesenti;
pedicelli usque ad | mm longi; carpophorum bipartitum. Fructus
dorsaliter compressus ellipticus orbicularisve 8-10 mm longus 7-9
mm latus apice rotundatis basi emarginatis, alis tenuibus distinctis
quam corpore parum angustioribus; vittae in intervallis 3—5, in com-
missuris 8—10.

1988] HARTMAN AND CONSTANCE: NEW LOMATIUM 123

Low, tufted, herbaceous perennial 4—12 cm tall, acaulescent, ar-
omatic, with a primary root 20-30 cm long or more, 0.3—1.2 cm in
diameter at summit, the crown unbranched or with few to several
branches arising 4-15 cm below ground, the crown or branches
enveloped their distal 1-3 cm by persistent leaf bases or their frayed
remains, which may double the apparent diameter. Leaves broadly
ovate to deltoid, 1.5—4 cm long, 2—5 cm wide, 2—3-pinnate-pinnati-
fid, pale green, glaucous, glabrous; primary leaf divisions 3-5 usually
in opposite pairs, the terminal one deeply pinnate-1—2-pinnatifid,
usually confluent with the upper pair, the lower distinct, more widely
spread proximally, the lowest often remote, the lateral primary di-
visions asymmetrically pinnate-1—2-pinnatifid into 40-50 or more
often confluent oblong to ovate segments, the lobes and teeth acerose;
petiole subterete to grooved adaxially, 1.5-5.5 cm long, expanded
at the base into a scarious sheath. Inflorescence a compact compound
umbel 10—25 mm in diameter, or enlarging to 40 mm in fruit; pe-
duncles ascending to erect in flower, or decumbent in fruit, 4-12 cm
long, equaling to much exceeding the leaves, glabrous; involucre
none, or rarely of an ovate to lanceolate bract 2-6 mm long; rays
5-9, 1-6 mm long in flower and up to 11 mm in fruit, subterete to
flattened, unequal, those bearing fruit becoming divergent to reflexed
and markedly enlarged at the base; umbellets andromonoecious, of
1-4 staminate and 5-10 (or more) perfect flowers or of all perfect
or all staminate flowers; involucels dimidiate, of 3-6 mostly lan-
ceolate to ovate bractlets that are entire, acute to acuminate, 1—3.5
mm long, and about equaling to somewhat exceeding the flowers,
usually distinct or essentially so, with a thin, white or purplish,
scarious margin, glabrous; pedicels 0.1—2 mm long on staminate
flowers, 0.1-1 mm on hermaphroditic flowers, those bearing fruit
becoming markedly enlarged at base. Flowers purple; sepals trian-
gular to lanceolate, 0.2-0.6 mm long, often unequal, enlarging little
in fruit, greenish; petals 1.6-1.9 mm long; anthers yellow, 0.7-0.8
mm long; filaments 1—-1.3 mm long; styles subterete, 2—2.4 mm long,
enlarging little in fruit, spreading to recurved; stylopodium none;
disc present; ovary glabrous, glaucous; carpophore bipartite. Fruit
dorsally flattened, broadly elliptic to orbicular, 8-10 mm long, 7-9
mm broad, glabrous, the wings distinct, narrower than the body;
vittae 3-5 in intervals, 8-10 on commissure.

TYPE: CA, Kern Co., se. slope of Owens Peak, eastern crest of the
southern Sierra Nevada, T25S R37E S21 ne.’4 MDB-+M, 8200 ft
(2500 m), 11 Jun 1986, James R. Shevock 11681 (Holotype: UC;
isotypes: CAS, MO, NY, RM, RSA, US).

PARATYPES: CA, Kern Co., type locality, 7 Apr 1984, Shevock
10812 (CAS: leaf only), 27 May 1985, Shevock, Norris & Rose 11197
(CAS, MO, NY, RM, RSA, UC); se. slope of Mt. Jenkins above
Pacific Crest Trail, T25S R37E $34 nw.'’4 MDB+M, 7300 ft (2225
m), 21 Apr 1986, Shevock & Ertter 11439 (CAS, RSA, UC).

124 MADRONO [Vol. 35

Distribution, ecology, and phenology. Lomatium shevockii occurs
on colluvial slopes and talus, usually along contact zones of meta-
morphic and granitic rock in open mixed coniferous forest or pinyon
pine/canyon live oak woodland. Populations are restricted to the
eastern side of the crest, generally between 2225 and 2500 m. Plants
occurring at the lower of these elevations are in canyon bottoms,
where the seeds presumably washed downslope primarily from late
summer thunderstorms. Flowers appear from late April to mid-May,
with fruit developing by mid-June. All populations are on federal
lands administered by the California Desert Conservation Area,
Bureau of Land Management.

This Lomatium is apparently a very restricted endemic. Habitats
with the combination of slope, aspect, geology, and elevation re-
quired by L. shevockii are limited along the rugged crest and are
believed to comprise less than 5 air km. The population near the
summit of Owens Peak is within an open, park-like mixed coniferous
forest of Pinus jeffreyi, P. flexilis, P. monophylla, P. lambertiana,
Abies concolor, and Juniperus occidentalis subsp. australis. On steep-
er slopes below this forest, the species extends into openings in the
pinyon pine/canyon live oak woodland. The site of the small pop-
ulation on Mt. Jenkins lacks limber pines, but is otherwise similar.
No single set of associated species accompany all populations of
Lomatium shevockii, but Allium burlewii, Eriogonum wrightii var.
subscaposum, Mimulus sp., Monardella spp., Orochaenactis thys-
anocarpha, Salvia pachyphylla, and Zauschneria latifolia were found
in proximity.

Despite the rarity of the new species, the rugged terrain it inhabits
should protect it from human disturbance. Other rare taxa being
inventoried by the California Natural Diversity Data Base and the
California Native Plant Society that are known to occur on Owens
Peak include Eriogonum breedlovei var. shevockii, Monardella sp..,
Phacelia novenmillensis, and Raillardella muirii.

Lomatium shevockii belongs to the Euryptera group, which com-
prises seven species and extends along the Pacific Coast from south-
ern Oregon to Baja California and Guadalupe Island. Coulter and
Rose (1900) said of Euryptera, which they treated as a genus distinct
from Lomatium: “‘... differs from typical Lomatium especially in
its foliage, which is much more simple, with broad often orbicular
leaflets, and sharp mucronate teeth. The wings of the fruit are in-
clined to be distinct, while in Lomatium the wings are united and
project below the seed.”

ACKNOWLEDGMENTS

The authors are grateful to Mark A. Schlessman, Robert Meinke, and the Editor
for numerous helpful comments. Charlotte Mentges Hannan provided the illustration.

1988] HARTMAN AND CONSTANCE: NEW LOMATIUM [25

LITERATURE CITED

CouLTER, J. M. and J. N. Rose. 1900. Monograph of the North American Um-
belliferae. Contr. U.S. Natl. Herb. 7:240, fig. 61.

(Received 8 Mar 1987; revision accepted 30 Nov 1987.)

ANNOUNCEMENT

NEw PUBLICATION

Dorn, RoBERT D. 1988. Vascular Plants of Wyoming, illustrated by
JANE L. DorRN. Mountain West Publishing, Cheyenne, WY. vi + 340
pp., paperbound. [Keys to 120 families, 650 genera, 2369 species, 39
subspecies, and 690 varieties; 93 new combinations, | new species, 4
new varieties, and | new name; section of taxonomic notes. Available
postpaid for $13.00 from Mountain West Publishing, Cheyenne, WY
82003.]

ANNOUNCEMENTS

NEw PUBLICATIONS

Hunter, S. C. and T. E. PAYSEN, Vegetation classification system for
California: User’s guide, U.S.D.A. For. Serv., Gen. Tech. Rep. PSW-94:
i-ll, 1-12, 1986. [A system of classifying plant communities in Cali-
fornia, with guidelines for recognizing such in the field. ]

KoutTNIk, D. L., A taxonomic revision of the Hawaiian species of the
genus Chamaesyce (Euphorbiaceae), Al/ertonia, vol. 4, no. 6, pp. 331-
388, Sep 1987, ISSN 0735-8032, $9.50 (from Publications Secretary,
Pacific Tropical Botanical Garden, P.O. Box 340, Lawai, Kauai, HI
96765). [Treatment of 14 spp., 17 vars.]

A NEW SPECIES OF SAXIFRAGA (SAXIFRAGACEAE)
FROM THE OLYMPIC MOUNTAINS, WASHINGTON, AND
VANCOUVER ISLAND, BRITISH COLUMBIA

RIck J. SKELLY
930 W. 10th, Port Angeles, WA 98362

ABSTRACT

Saxifraga tischii, a new species from the Olympic Mountains of Washington, and
Vancouver Island, British Columbia, is described and illustrated. Distinctive features
include its dwarf size, persistent chlorophyllous petals, and leaves with relatively
aporous spongy mesophyll.

Recent surveys of Olympic Peninsula flora (Buckingham and Tisch
1979, 1983) have revealed the existence of a new species, herein
described.

Saxifraga tischii Skelly, sp. nov.

Herba rosulata, perennis. Rhizoma breve. Laminae 5.5—17(—22)
mm longae, 3.5—10(—17) mm latae, ovatae, ellipticae vel orbiculatae,
supra glabratae, infra fusco-tomentosae, apicibus acutis vel obtusis,
basibus cuneatis vel acutis, marginibus crenato-serratis; petioli
(3—)4—10(—20) mm longi, marginibus pilosis. Caulis florifer (2—)3.5—
7.5 cm altus, purpureus, glandulosis pubescentibus. Flores (3—)5—10
(—-18); calyx 2-4 mm longus, rotatus vel campanulatus, purpureus,
dentibus ovatis vel lanceolatis (0.7—)1—2(—2.5) mm longis; petales
5-6, lanceolati vel spathulati, 1.2—2.2(—2.6) mm longi, 0.3—0.8(-1.2)
mm lati, virentes vel purpureli, parce ciliati; filamenta 0.6—1.9(—2.3)
mm longa; antherae roseae; styli divaricati; pedicelli 2-6 mm longi
cum 0-2 bracteolis lanceolatis. Fructus purpureus, glaber, 2.3-—3.5
mm longus (Fig. 1).

Rosulate perennial herb arising from short rhizome. Leaf blades
5.5—17(—22) mm long, 3.5—10(—17) mm wide, ovate, elliptical, round
or rhomboidal, glabrous (rarely puberulent) above, brownish-to-
mentose and anthocyanic beneath, with acute to obtuse apex, cu-
neate to acute (rarely obtuse) base, and crenate-serrate margins, teeth
7-17; petioles (3-)4-10(-20) mm long, 1.5—3(-—5) mm wide, with
pilose margins. Flowering stem (2—)3.5—7.5 cm tall (in fruiting con-
dition to 12.5 cm), purplish, glandular-pubescent, scapiform, brac-
teate; primary bracts lanceolate, 3—5.5(—7) mm long. Flowers (3—)5-—
10(—18) in small cymes; calyx 2-4 mm long, rotate to campanulate,
purplish, puberulent, lobes 3-nerved, ovate to lanceolate, (0.7—)1-

MADRONO, Vol. 35, No. 2, pp. 126-131, 1988

1988] SKELLY: NEW SPECIES OF SAXIF RAGA 27

500 “wm

(<p)
SSS SSS ee
150 ~.m
i, Ly,
ee
Se .
X&
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3cm

Fic. 1. Saxifraga tischii. A. Habit. B. Flower and pedicel. C. Cellular details of
petal margin and trichome. D. Trichomes from pedicels. E. Comparable trichomes
from S. aequidentata.

2(—2.5) mm long, with acute to rounded apices; petals 5—6, lanceolate
to spatulate or oblanceolate, clawless, persistent, 1.2—2(—2.6) mm
long, 0.3-0.8(—1.2) mm wide, greenish to purple, often with 1-7
cilia-like trichomes; stamens 10-12, filaments 0.6—1.9(—2.3) mm long,
usually white (rarely purplish), widest at base; anthers pinkish to
orange, ca. 0.3-0.5 mm wide; styles divaricate, ca. 1 mm long; ped-
icels 2-6 mm long with 0-2 bracteoles 1.5—3.5 mm long. Fruit pur-
plish, glabrous, 2.3—3.5 mm long. Seeds tawny-colored, fusiform,
ca. 0.6 mm long.

128 MADRONO [Vol. 35

Type: USA, WA, Clallam Co.: Mt. Baldy, Olympic Mountains,
T28N R4W S22, 6300 ft (1920 m), 24 Jul 1976, Tisch 2307 (Ho-
lotype: UC).

PARATYPES: WA, Jefferson Co.: Buckhorn Mt., Olympic Mts.,
T27N R4W S13, 6800 ft (2070 m), 3 Aug 1984, Tisch 2817 (WTU)
and Tisch 2818 (WS); Goat Lake area, Olympic Mts., T27N R4W
S9, 6300 ft (1920 m), 18 Aug 1984, Tisch 2897 (OSC); ‘‘Petunia
Peak”, Olympic Mts., T27N R4W S39, 6500 ft (1980 m), 18 Aug
1984, Tisch 2905 (ORE); Mt. Olympus, Olympic Mts., T27N R8W
S33, “8150 ft.’ (probably ca. 7900 ft), 13 Aug 1907, Flett 3015
(WTU); Clallam Co.: Mt. Angeles, Olympic Mts., T29N R6W, 5
Sept 1909, Webster 1879 (WTU); Mt. Angeles, 5500 ft (1676 m), 2
Aug 1930, Thompson 5556 (WTU); Hurricane Ridge, Olympic Mts.,
4500 ft (1372 m), 9 Jun 1934, Thompson 10589A (WTU). Canada,
British Columbia, Vancouver Island, Castlecrag Mt., Strathcona Park,
1500 m, 9 Aug 1979, Ogilvie and Beguin 798911 (V).

Distribution, habitat and phenology. Saxifraga tischii grows on
ledges and in rock crevices at subalpine and alpine levels, ca. 1372-
2400 m, in the Olympic Mountains, Clallam and Jefferson cos.,
Washington, and the interior of Vancouver Island, British Colum-
bia. Substrates include basalt, breccia, limestone and sandstone. The
plants occupy shallow (ca. 1-3 cm), well-drained soil pockets on
north to northeasterly aspects, often in cirques near persistent snow
patches. They flower from June to August. The chlorophyllous petals
may reflect extended adaptation to the short growing seasons of these
shady, microthermal environments.

Associated species include: Anemone drummondii Wats. var.
drummondii, Carex nardina Fries, Cystopteris fragilis (L.) Bernh.,
Douglasia laevigata A. Gray var. ciliolata Const., Draba lonchocarpa
Rydb. var. lonchocarpa, Luetkea pectinata (Pursh) Kuntze, Luzula
spicata (L.) DC., Poa paucispicula Scribn. & Merr., Ranunculus
eschscholtzii Schlecht. var. eschscholtzii, Romanzoffia_ sitchensis
Bong., Saxifraga caespitosa L. var. emarginata (Small) Rosend.,
Senecio flettii Wieg., Veronica cusickii A. Gray, and Viola flettii
Piper.

Relationships. Saxifraga tischii does not key well to known taxa
in floras covering the Olympic Peninsula (Jones 1936, Hitchcock
and Cronquist 1974), nor is it found in treatments for adjacent
regions to the north (Hulten 1968, Scoggan 1978-79) and east (Davis
1952). Hitchcock (in Hitchcock and Cronquist 1961) discusses
apetalous and purple-petaled Saxifraga from Mt. Olympus and vi-
cinity, referring them to S. occidentalis Wats. var. rufidula (Small)
Hitchc. Perkins (1978), who monographed the S. occidentalis species
complex, accepted Hitchcock’s interpretation and merely reassigned

1988] SKELLY: NEW SPECIES OF SAXIF RAGA 129

the specimens to S. aequidentata (Small) Rosend., a synonym of the
former taxon. I examined these early collections, listed here as para-
types, and found them to match the S. tischii holotype in all pertinent
features.

The type specimens of S. tischii, without exception, possess tiny
chlorophyllous petals with anthocyanic colorings on their margins
and apices. The petals do not wither following anthesis, but remain
alive and, apparently, photosynthetic through advanced fruiting.
They form no visible abscission layer. Petals of S. tischii, in contrast
with those of S. aequidentata, have a mesophyll-like core, stomata,
and often bear multicellular trichomes (ca. 1-7) along their margins
(Fig. 1). Although plants of S. tischii are generally smaller than those
of S. aequidentata, developmental differences and environmental
extremes produce some quantitative overlap. There are significant
differences (p < 0.001) in petal width, filament length, number of
leaf dentations, and petiole length/width ratio (Table 1). The mea-
surements of S. aequidentata are largely from specimens at WTU
that were annotated by Perkins and represent the range of that species.

Similarities in pistil structure, scape branching, and the general
shape and dentation of basal leaves suggest that S. tischii and S.
aequidentata share common ancestry within the SS. occidentalis species
complex. Saxifraga tischii may have evolved during the Pleistocene
in mountain refugia of coastal British Columbia and/or in the Olym-
pic Peninsula. The current restriction of S. tischii to cool, shady rock
crevices at high elevations suggests it has limited ecological ampli-
tude and is a highly specialized, narrowly restricted species. In the
Olympics and elsewhere, S. aequidentata has wider distribution and
occupies a variety of moderately sunny to shady environments from
near Sea level (e.g., near Portland, Oregon) to above 2000 m. Perkins
(1978) stated that it prefers thin-soiled rock outcrops “‘in vernally
moist, often dripping seeps”. Such environments in the Olympics
(e.g., cliffs at Lake Crescent cited by Perkins 1978) usually dry out
by late June, leaving the plants in a shriveled condition. Saxifraga
tischii is semi-evergreen and carries some leaves through two grow-
ing seasons. In its normal habitats it does not shrivel after anthesis.
Moreover, it grows poorly when transplanted to lower elevations.
Ten cold-frame plants of S. tischii, kept at 650 m near Port Angeles,
Washington, produced only one 2-cm (5-flowered) scape in two
growing seasons. Three S. aequidentata, similarly located, produced
eight scapes with up to 40 flowers. Under cultivation S. tischii re-
tained its dwarf stature and produced typical flowers with green
petals, short filaments, etc., indicating that these traits are genetically
fixed. Scapes of S. aequidentata grew to over 12 cm and produced
their normal white flowers.

Floral structure is usually more conservative and less subject to

[Vol. 35

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1988] SKELLY: NEW SPECIES OF SAXIF RAGA 131

evolutionary change than vegetative structure (Benson 1957, Jones
and Luchsinger 1986). The minimum of 12 floral differences that
separate these two species (Table 1) indicate much greater evolu-
tionary distance than their superficial vegetative similarities may
have suggested to earlier investigators.

This species is named after E. L. Tisch, my biology instructor and
advisor at Peninsula College, Port Angeles, Washington.

ACKNOWLEDGMENTS

I am grateful to A. Cronquist and an anonymous reviewer for their comments, to
Karen Lull-Butler for doing the illustration, and to Vince Murray for assisting with
the Latin. I also thank the B. C. Provincial Museum, WTU herbarium, and Peninsula
College for use of their facilities, and E. Tisch for taking me to collection sites and
for making available his field notes and living and mounted specimens.

LITERATURE CITED

BENSON, L. 1957. Plant classification. D. C. Heath & Co., Boston.

BUCKINGHAM, N. M and E. L. Tiscu. 1979. Vascular plants of the Olympic Pen-
insula, Washington (a catalog). Natl. Park Serv., Univ. Wash. Coop. Park Studies
Unit Rep. B-79-2, Seattle.

and 1983. Additions to the native vascular flora of the Olympic
Peninsula, Washington. Madrono 30:67-78.

Davis, R. J. 1952. Flora of Idaho. Wm. C. Brown Co., Dubuque, IA.

Hitcucock, C. L. and A. CRONQUIST. 1974. Flora of the Pacific Northwest. Second
printing with corrections. Univ. Washington Press, Seattle.

and 1961. Part 3: Saxifragaceae to Ericaceae. In C. L. Hitchcock,
A. Cronquist, M. Ownbey, and J. W. Thompson, Vascular plants of the Pacific
Northwest. Univ. Washington Press, Seattle.

HULTEN, E. 1968. Flora of Alaska and neighboring territories. Stanford Univ. Press,
Stanford, CA.

JONES, G. N. 1936. A botanical survey of the Olympic Peninsula, Washington.
Univ. Wash. Publ. Biol. 5, Seattle.

Jones, S. B. and A. E. LUCHSINGER. 1986. Plant systematics. McGraw-Hill Book
Co., N.Y.

PERKINS, W. E. 1978. Systematics of Saxifraga rufidula and related species from
the Columbia River Gorge to southwestern British Columbia. Ph.D. thesis, Univ.
British Columbia, Vancouver.

SCOGGAN, H. J. 1978-79. The Flora of Canada. Parts 1-4. National Museums of
Canada, Ottawa, Ontario.

(Received 15 Sep 1986; revision accepted 7 Feb 1988.)

EVIDENCE FOR A WARM DRY EARLY HOLOCENE
IN THE WESTERN SIERRA NEVADA OF CALIFORNIA:
POLLEN AND PLANT MACROFOSSIL ANALYSIS OF
DINKEY AND EXCHEQUER MEADOWS

OwEN K. DAVIS
Department of Geosciences, University of Arizona,
Tucson 85721

MICHAEL J. MORATTO
INFOTEC Research Incorporated, 19524 Hillsdale Drive,
Sonora, CA 95370

ABSTRACT

Pollen and plant macrofossil analysis and ten radiocarbon dates for the sediments
of Dinkey and Exchequer Meadows provide a detailed record of environmental
change in the western Sierra Nevada. The Dinkey Meadow sedimentary record is
nearly 5000 yr long, and the Exchequer Meadow record reaches 13,500 yr B.P. The
Exchequer Meadow pollen diagram is divided into an upper Abies zone (O-1870 yr
B.P.), a Pinus zone (1870-7070 yr B.P.), and a basal Artemisia zone (7070—13,500
yr B.P.), which is subdivided into upper Quercus and lower Gramineae subzones at
ca. 10,680 yr B.P. The Artemisia zone records more xeric vegetation than occurs west
of the Sierra today, and it contains Sequoiadendron pollen, indicating temperatures
little colder than today. The presence of spores of the dung fungus Sporormiella
indicates that grazing animals were abundant during the Gramineae subzone. A period
of maximum Abies percentages at Dinkey, Exchequer, and other Sierra Nevada sites
may result from warm dry climate shortly after 1900 yr B.P.

California’s remarkable diversity of vegetation types results in
large part from its topographic and climatic heterogeneity. The for-
ests of the Coast Ranges and Sierra Nevada are separated by the
grasslands of the Central Valley, and deserts occupy the rainshadow
of eastern California (Major 1977). An unexpected finding of pa-
leoenvironmental research in California is that the coastal, interior,
and rainshadow areas may have had different climatic histories.
During the late-Pleistocene and early Holocene, coastal California
and eastern California appear to have been moister than today,
whereas the western Sierra was drier (Davis et al. 1985, Davis and
Sellers 1987).

At the end of the Pleistocene, mesic pine forests of coastal Cali-
fornia were replaced by oak woodlands. Adam and West (1983)
interpret the higher Pinus/Quercus pollen ratios from Tule Lake
(West 1982) and Clear Lake (Adam et al. 1981) as indicating greater
moisture and cooler temperature before 7000 yr B.P. Plant macro-
fossil deposits also indicate increased moisture during the early Ho-

MADRONO, Vol. 35, No. 2, pp. 132-149, 1988

1988] DAVIS AND MORATTO: WARM DRY EARLY HOLOCENE 133

locene (Johnson 1977), and Bergquist (1977) reports Picea pollen,
evidence for a cool-moist climate, in sediments older than 8400 +
100 yr B.P. at Bolinas Lagoon, near San Francisco.

In contrast, vegetation of the western Sierra during the early Ho-
locene resembled that found east of the Sierra today. Artemisia pollen
is abundant in early Holocene sediments from Balsam Meadow
(Davis et al. 1985), Swamp Lake (Batchelder 1980), Tulare Lake in
the San Joaquin Valley of California (Atwater et al. 1986), and in
lower King’s Canyon (Cole 1983). Comparable Artemisia percent-
ages have not been duplicated in any modern pollen samples from
west of the Sierran crest (e.g., Adam 1967, West 1982, Davis et al.
1985, Anderson 1987). Such high Artemisia percentages have been
found only in vegetation east of the crest, in the Sierran rain shadow
(Adam 1967, Mehringer 1967, Anderson 1987). For such vegetation
to occur on the west slope of the Sierra, the climate must have been
drier than today. A glacial-age expansion of the Great Basin vege-
tation west of the Sierra is also documented by the abundance of
Sarcobatus pollen in the Tulare Lake core (Atwater et al. 1986).

Early-Holocene aridity is documented at other sites of interior
California by low lake levels and the expansion of xeric vegetation.
At Gabbott Meadow Lake (1900 m, Mackey and Sullivan 1986)
near the Sierra crest, oak percentages rise from 10,500 + 140 to
7570 + 100 yr B.P. and decline until 2270 + 80 yr B.P., probably
indicating an expansion of xeric oak woodland into pine forest. At
Cedar Lake, Siskiyou Co., California (1743 m, West 1986), increased
aridity during the early Holocene is indicated by elevated “TCT”
(Taxaceae, Cupressaceae, Taxodiaceae) percentages from 7910 +
120 to 10,180 + 150 yr B.P. Most of the TCT pollen is probably
derived from Chamaecyparis lawsoniana, which must have become
established on the moraines surrounding the lake during a period
of xeric climate. Because the lake is near the coast, this aridity
conflicts with interpretations for other coastal sites; alternatively,
the climatic history of the northern California coast may differ from
that of more southern sites. However, Cedar Lake is leeward of the
Siskiyou Mountains, which reach elevations over 2100 m. If ocean
fogs are responsible for the early Holocene moisture in coastal sites,
the Siskiyou rain shadow may have produced a climate like that of
interior sites.

East of the Sierra Nevada in the Mojave Desert, the climate was
wetter at the end of the last glaciation. Glacial Lake Mojave over-
flowed from ca. 15,500 to 10,500 yr B.P. (Wells et al. 1987), and
Searles Lake overflowed ca. 11,000 yr B.P. (Smith and Street-Perrott
1983). Packrat middens from west of Las Vegas, Nevada, contain
elevated percentages of mesic shrubs and succulents, indicators of
increased summer precipitation from 12,000 to 8000 years ago
(Spaulding and Graumlich 1986).

134 MADRONO [Vol. 35

Because the Sierra Nevada was a major mountain range by 3
million yr B.P. (Chase and Wallace 1986), the trans-Sierra climatic
contrast has existed throughout the Pleistocene. During the last gla-
ciation the direction of prevailing winds was probably the same as
today’s. Coastal sand dunes of late glacial age record wind directions
equivalent to modern (Johnson 1977). Pleistocene snowlines were
600 m higher in the rainshadow of the Sierra Nevada than in adjacent
mountains. This greater elevation indicates moisture patterns sim-
ilar to today’s patterns (Porter et al. 1983).

Purpose of the study. Pollen analysis of Dinkey and Exchequer
meadows, Fresno County, California, was undertaken to confirm
the early Holocene aridity of the western Sierra Nevada. This ver-
ification is particularly important because areas to the west (coastal
sites) and east (desert sites) of the western Sierra record greater
moisture during the early Holocene. These regional differences in
climatic change are in marked contrast to traditional climatic scenar-
ios (Antevs 1948) that call for uniform climatic change throughout
western North America: cool and moist before 7000 yr B.P., hot
and dry from 7000 to 4500 yr B.P. (the Altithermal), and near
modern climate from 4500 yr B.P. to present.

STUDY AREA

Regional climate. Precipitation in the western Sierra is dominated
by the Aleutian low, which sends cyclonic storms eastward from the
Pacific Ocean during winter months (Pyke 1972). Precipitation is
greatest in January and February, with a distinct period of drought
in June, July, and August when dry descending air from the Pacific
high covers the Pacific coast and the juxtaposition of cold, dense
oceanic air and warm continental air produce the stable Pacific air
mass boundary (Mitchell 1976). As cyclonic storms move westward
in winter, they cross the Sierra crest, losing most of their moisture
before they enter the western Great Basin.

Present vegetation. The vegetation near Dinkey Meadow (37°N,
119°10'W, 1683 m) and Exchequer Meadow (37°N, 119°5’W, 2219
m) is characteristic of the upper (Exchequer) and lower (Dinkey)
Sierran Montane Forest. Pinus murrayana is scattered over Dinkey
Meadow, which is surrounded by a mixed stand of Abies concolor,
Pinus jeffreyi, Pinus lambertiana, and Calocedrus decurrens. The
coring site is covered with forbs, grasses, and scattered shrubs in-
cluding Vaccinium occidentale, Polygonum amphibium, Scirpus sp.,
and Poa spp. in wet places; and Ribes roezlii, Ribes nevadense, Sym-
phoricarpos parishii, Ceanothus leucodermis, Viola macloskeyi, and
Apocynum pumilum on the uplands.

The coring site at Exchequer meadow is wetter and is dominated

1988] DAVIS AND MORATTO: WARM DRY EARLY HOLOCENE 135

by Carex spp. and Scirpus spp. Scattered Pinus murrayana grow on
the meadow, which is surrounded by Pinus jeffreyi and occasional
Abies magnifica. Ribes roezlii and Ribes nevadense are common
understory plants in the forest.

Glacial deposits. The western Sierra was glaciated extensively dur-
ing the late Pleistocene, and Matthes (1960) mapped glacial deposits
nearly down to the elevation of Dinkey Meadow ca. 1700 m. Al-
though we know of no detailed maps of glacial deposits for the area,
Matthes’ (1960) maps indicate that Dinkey Meadow was beyond the
terminal moraine, and Exchequer Meadow was adjacent to, but not
covered by, the Dinkey Creek glacial lobe (Fig. 1).

METHODS

The sediments of Dinkey and Exchequer Meadows were cored on
September 1, 1985. The wettest portions of the meadows were cho-
sen for coring to avoid oxidation or loss of sediment due to drying.

Sampling techniques. At Exchequer Meadow the upper 124 cm
of sediment was collected with a 5 cm diameter square rod piston
sampler (Wright 1967), and the lower sediment was cored with a
2.5 cm diameter Dachnowsky corer (Faegri and Iversen 1975) to
399 cm depth. The entire 300 cm Dinkey Meadow core was re-
covered with the piston sampler except for the interval from 210 to
230 cm, which was recovered with the Dachnowsky.

The cores were wrapped in plastic film and aluminum foil, and
were stored at 1°C prior to sampling. Volcanic ash layers were sub-
mitted to Andrei Sarna-Wojcicki, U.S. Geological Survey, Menlo
Park, California, for identification. Radiocarbon samples were sub-
mitted to Beta Analytic, Coral Gables, Florida.

Radiocarbon dates. All ten radiocarbon samples (Figs. 2 and 3)
were adjusted for '3C/!?C fractionation, and samples Beta-16113
and Beta-17185 from Exchequer Meadow were given extended
counting times because they contained small amounts (0.20 and 0.35
g, respectively) of carbon. For Dinkey Meadow the sedimentation
rate changed from 0.08 cm yr“! to a slower 0.05 cm yr7! below 150
cm (Fig. 2). For Exchequer Meadow the sedimentation rate was
nearly constant (0.03 cm yr~') from the surface to the base of the
core (Fig. 3).

Pollen extraction. Pollen extraction followed standard procedures
(Faegri and Iversen 1975). The samples (volume | cm?) were placed
in 10% HCl and Lycopodium tracers were added to permit calcu-
lation of pollen concentration. After screening, the samples were
treated with concentrated HCl and left overnight in 40% HF to
remove carbonates and silicates. The samples were acetolyzed to

136 MADRONO [Vol. 35

QUILLWORT @
POND

EXCHEQUER
MEADOW

@ TULE LAKE

McKinley

e
GLEAR'EAKE GABBOTT LAKE

® Sacramento
©@ SWAMP LAKE 118°

STUDY AREAL_]
®@KING’S CANYON

@ TULARE LAKE

Fic. 1. Map of Dinkey and Exchequer Meadows area showing extent of late-
Pleistocene Glaciation (stippled pattern). Inset shows location of California sites

mentioned in text.

remove cellulose and similar organic compounds, and treated with
10% KOH to remove humates. After staining, the samples were

transferred to glycerin.
Pollen identifications were based on the reference collection and

1988] DAVIS AND MORATTO: WARM DRY EARLY HOLOCENE 137

DINKEY MEADOW

100

1980 + 160 a
BETA 16109

(2920 + 90
BETA 17186

2660 + 100 []
BETA 17187

DEPTH (cm)

200

0.05 cm yr

4850 + 110

300 BETA 16110 ©
) 1 2 3 4 5
RADIOCARBON YEARS B.P.
(THOUSANDS)

Fic. 2. Graph of Dinkey Meadow radiocarbon dates versus sediment depth. Solid
line shows least squares regression of dates on sediment depth below 150 cm; above
this depth it is connected directly to the surface. Height of squares is sample interval,
width is date + one standard deviation.

library at the University of Arizona Palynology Laboratory. The
pollen sum (divisor for percentage calculations) does not include
aquatics; e.g., Salix or Cyperaceae, or spores. The following notes
apply to the types shown on the pollen diagrams (Figs. 4 and 5):
Cupressaceae may include some Taxaceae and Taxodiaceae except
Sequoiadendron. Ericaceae includes mostly Arctostaphylos but at
least one other type was seen. “Other Compositae”’ includes all
pollen of that family excluding Ambrosia, Artemisia, Cirsium, and
Liguliflorae.

Plant macrofossils. The sediment from both sites was suspended
in water and screened to remove fine inorganic particles. Identifiable
remains (seeds, needles, and large pieces of wood) were removed
from the matrix under 7-45 x magnification. Conifer needles were
sectioned to permit species identification, and the abundance of

138 MADRONO [Vol. 35

EXCHEQUER MEADOW

qo 1870 + 160
BETA 16111

~ 2980 + 80

100 0 BETA 17183

~~

E 4540 + 90 0

ce BETA 17184

120

zi 7070 + 70 |

+ BETA 16112

a) 1

Cc
11,490 + 270
BETA 17185

300

10,330 + 380
BETA 16113

400
to) 2 4 6 8 10 12
RADIOCARBON YEARS B.P.
(THOUSANDS)

Fic. 3. Graph of Exchequer Meadow radiocarbon dates versus sediment depth.
Solid line shows least squares regression of dates on sediment depth. Height of squares
is sample interval, width is date + one standard deviation.

charcoal was noted on a scale of zero (absence) to four (very abun-
dant).

RESULTS

The Dinkey Meadow sediments are yellowish brown (1OYR 5/4,
Munsell color) to very dark gray (SYR 3/1) peat down to 69 cm;
and dark gray (1OYR 4/1) medium sand to the base of the core. A
volcanic ash layer is present at 39-40 cm. Exchequer Meadow sed-
iments are less homogeneous. They are primarily peat down to 101

1988] DAVIS AND MORATTO: WARM DRY EARLY HOLOCENE hoi

DINKEY MEADOW OOS 2)
POLLEN PERCENT oo a z me 2 aves R BS KS Ko ue
or” or? Gey ie of eee ES ec or

G
() 200 400 600
@ 1620 @ 18 20 30 40 50 6B 78 88 98 B il 18 B 182i 18 28 » 18 8188 \ B i SS Ve 1828 B i 18 @ 18 2838 48

DEPTH (cm)

328

Fic. 4. Dinkey Meadow pollen diagram for abundant pollen types plotted versus
sample depth. Dots are for percentages less than 2%. Cyperaceae and other aquatics
are outside the pollen sum, Pinus is in the sum. Pollen zones and ages of boundaries
shown on right. Vertical width of charcoal histograms shows width of sediment sample
interval.

cm, interbedded sand and peat to 290 cm, and alternating fine and
coarse sand down to 399 cm. Two volcanic ashes are present at
Exchequer Meadow: one at 50-54 cm; the other at 169-170 cm; The
54 cm Exchequer Meadow ash, the only one suitable for identifi-
cation, was identified as a “‘young Inyo Crater Ash.”’

Dinkey Meadow. The Dinkey Meadow pollen diagram (Fig. 4)
shows relatively little change, which is not surprising given its rel-
atively brief (<5000 yr) record. Pinus is the most abundant pollen
type (50-83%), followed by Abies (4-19%) and Quercus (O-13%).
The percentages of Abies are lower and the percentages of Quercus
are higher than at Balsam Meadow, which is consistent with the
lower elevation (2040 vs. 1683 m) of Dinkey Meadow. The age of
maximum Abies pollen percentages (Fig. 4) at Dinkey Meadow
(1980 + 160) is very close to estimated age of maximum Abies
percentages (1710 yr B.P.) at Balsam Meadow.

The plant macrofossils from Dinkey Meadow (Table 1) provide
additional data for paleoenvironmental reconstructions. All of the
conifer species now at the site are present as plant macrofossils.
Pinus murrayana needles are the most abundant macrofossil, but
other conifers are absent above 100 cm. The sediments from 100-
170 cm, which contain maximum Abies pollen percentages, contain

140 MADRONO [Vol. 35

EXCHEQUER MEADOW oe) : e
POLLEN PERCENT oF eX Oe oe
\e? v co OV WON eee AS oes es \°

10% \a e) oe% Fao. Qe Fe \ Se SAO Cas Raa Coo

@ 18 2038 8 18 2030 485860 708898 8 1828 8 10 8 188 188 188 1828 10 a 1828 @ 1828 @ 18 20 30 4050
Q —— a ae t 4 7 a oy — — — jo — — — a 4 444

28

48

62

82
128
120
148
162
182
202
220
240
262
280
302
3204) --
340
362
388
402

(cm)

DEPTH

ee ee ee ee ee

SS SS oe Se iS eS ees

Fic. 5. Exchequer Meadow pollen diagram for abundant pollen types plotted
versus sample depth. Dots are for percentages less than 2%. Cyperaceae and other
aquatics are outside the pollen sum, Pinus is in the sum. Pollen zones and ages of
boundaries shown on right. Vertical width of charcoal histograms shows width of
sediment sample interval.

both Abies concolor and Abies magnifica needles, so both species
could have contributed to the Abies pollen maximum. The Abies
magnifica needles in the 110-120 cm sample, however, are the only
remains of this species in the core. Charcoal abundance (Fig. 4)
shows two peaks: one at 170-180 cm (2365 yr B.P.) just before
maximum Abies pollen percentages, and the other at 260-270 cm
(ca. 4180 yr B.P.). This trend in charcoal concentration is similar
to that at Balsam Meadow (Davis et al. 1985).

Exchequer Meadow. The pollen zones for Exchequer Meadow (Fig.
5) are the same as those for Balsam Meadow (Davis et al. 1985)
demonstrating the regional nature of the vegetation change, but the
beginning of the Abies zone is later (1870 + 60 yr B.P.) than at
Balsam Meadow (3000 yr B.P.). The age of maximum fir percentages,
however, 1s nearly the same at Exchequer (1870 + 70), Dinkey
Meadow (1980 + 160) and Balsam Meadow (1710 yr B.P.). The
boundary between the Pinus and Artemisia zones at 7070 + 70 yr
B.P. is synchronous with this transition at Balsam Meadow, but the
basal Artemisia zone is longer, and is subdivided into upper Quercus
and lower Gramineae subzones at ca. 10,680 yr B.P. In the Gra-
mineae subzone percentages of Seqguoiadendron, Gramineae, Cas-
tilleja, and Cruciferae are greater than any at Balsam Meadow.

The diversity and concentration of plant macrofossils at Exche-
quer Meadow is less than at Dinkey Meadow. Sedge (Carex) remains

1988] DAVIS AND MORATTO: WARM DRY EARLY HOLOCENE 141

‘ S °
OD ae
on 4 & CS oo OK 2 es ee <® A o°
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coe ‘cove Q YE ob Yi* We < Ly?

8 1820 8 1828 8 18 8 10 B 182838 8 18 B 1820308405060 8 18 @ 243 4878 800

1840

Fic. 5. Continued.

are the most abundant type. The Abies needles were preserved too
poorly to identify to species, but Abies sp. and Pinus murrayana
remains were present in surface sediment (Table 2). Charcoal frag-
ments reach peak abundances at 80-90 cm (2740 yr B.P.), 124-129
cm (4140 yr B.P.), and 199-244 cm (7350 yr B.P.), a sequence similar
to that at Balsam Meadow.

DISCUSSION

Paleoclimatology. An Abies pollen maximum shortly after 1900
yr B.P. is present at all three sites we have studied in the area. This
event follows the general initiation of meadow development in the
western Sierra (Wood 1975) that accompanied the beginning of
Neoglacial cooling ca. 3000 years ago. Nineteen hundred yr B.P. is
recognized as an interval of glacier retreat within the Neoglacial in
western North America (Porter and Denton 1967), but Scuderi (1984)
states that glaciers in the high Sierra Nevada may have advanced at
this time (1850 yr B.P.). Nineteen hundred yr B.P. falls between
periods of cool moist climate recorded in the growth of bristlecone
pine in the White Mountains of eastern California (LaMarche 1978).

At Balsam and Dinkey Meadows macrofossils of Abies concolor
and A. magnifica are present during the Abies pollen maximum
(Table 2, Fig. 5). Today, these species do not grow on the bogs. Both
white and red fir are characteristic of relatively dry slopes (Munz
1959), so the invasion of the meadows 1900 yr B.P. may indicate a
period of warm, dry climate when the bogs dried. The Exchequer
Meadow core was taken near the edge of the meadow; consequently,

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144 MADRONO [Vol. 35

its surface sediment contains fir macrofossils. During the fir pollen
maximum, sedge pollen percentages are low at Dinkey Meadow (Fig.
4), and sedge macrofossils are absent in Exchequer Meadow sedi-
ments (Table 2), perhaps indicating meadow desiccation.

A date of 1920 + 50 yr B.P. (A-4428) at 64-70 cm marks a
transition from inorganic to organic sediments at Quillwort Pond
(Fig. 1). Jsoetes megaspores are present throughout the | m core,
but below 70 cm pebbles are abundant and plant macrofossils are
absent. Above 70 cm Potamogeton and Carex are common, and
conifer needles and seeds are abundant (up to 12 Abies needles cm).
Thus, the date marks the beginning of encroachment of trees and
emergent aquatics into the Quillwort Pond basin.

Sites at higher elevation do not consistently show the Abies pollen
maximum (Anderson 1987), so its importance may be limited to
low elevation sites (below 2400 m?) where precipitation is less.

An “early Holocene Xerothermic.’’ The classical climatic sequence
for western North America calls for greatest temperatures and lowest
moisture 7000-4500 yr B.P. during the “‘Altithermal’’ of Ernst An-
tevs (1948). This chronology was never accepted universally (Asch-
man 1957, Martin 1963), and many authors (Kearney and Luckman
1983, Ritchie et al. 1983, Hebda and Mathewes 1984, Davis et al.
1985, 1986, Elias 1985, Vance 1985) have found evidence for a
much earlier thermal maximum centered ca. 10,000—8000 yr B.P.,
before the beginning of the classical Altithermal.

Many of the studies documenting the “‘early Holocene Xerother-
mic’ (a phrase proposed by Hebda and Mathewes 1984), have been
of sites at high elevation (Kearney and Luckman 1983, Elias 1985)
or at high latitudes (Ritchie et al. 1983) where summer temperature
controls the position of tree line. Due to the changing relationship
of the perihelion and the summer solstice, insolation during summer
months (June, July, August) was greatest in the northern hemisphere
prior to 7000 yr (Davis et al. 1986). Although the actual change in
insolation is small, atmospheric circulation is very sensitive to even
small changes in insolation. General Circulation Models (e.g., Sellers
1984, Kutzbach and Guetter 1986) indicate summer temperature
9000 yr B.P. were 1—2° C warmer than today.

Previous paleoenvironmental studies in the western Sierra Nevada
have demonstrated maximum aridity in the early Holocene ca. 7000—
10,000 yr B.P. (Davis et al. 1985). The data from Exchequer Meadow
corroborate this finding, providing support for the contrast between
western Sierra climate and that of regions to the west and east. The
early-Holocene sediments from Exchequer Meadow also contain the
pollen of species that are today near their upper-elevational limits,
indicating that temperatures were not much colder than today, there-
by supporting the findings of Kearney and Luckman (1983), Ritchie
et al. (1983), and Elias (1985) based on the position of upper treeline.

1988] DAVIS AND MORATTO: WARM DRY EARLY HOLOCENE 145

A B C D: .. . | Ee
ei%¢°%

Fic. 6. A-—D. Pollen from Exchequer Meadow, 312 cm Artemisia (A), Cupres-
saceae (B), Sequoiadendron (C—D); E. Sporormiella spore (Exchequer Meadow, 345
cm); F—G. terminal bud of Calocedrus or Juniperus (Exchequer Meadow, 274-299
cm).

At Exchequer Meadow, Sequoiadendron pollen is present from
312-340 cm and acf. Calocedrus or Juniperus macrofossil is present
at 274-299 cm (Fig. 6). Both Sequoiadendron and Calocedrus (now
at 1400-2560 m and 730-2500 m respectively, Munz 1959, 1968)
would be near their upper elevational limit at Exchequer Meadow
(2219 m), and neither are present there today. The nearest Sequoia-
dendron grove to Exchequer Meadow today is the McKinley grove,
5 km south at 1951 m elevation (Fig. 1).

A series of environmental factors influence the distributions of
plant taxa. Soil moisture and fire frequency are particularly impor-
tant for the regeneration of Sequoiadendron. But in general, the upper
elevational limits of plants are set by temperature (Daubenmire
1943). For these species to have been present near their current
upper-elevation limits during the early Holocene, temperatures could
not have been much colder than today.

Cole (1983) has documented the presence of Sequoiadendron pol-
len and Calocedrus macrofossils from 14,190 to over 45,000 yr B.P.
in packrat middens from 980 to 1280 m elevation in Kings Canyon
(Fig. 1). It appears that these species also were more widespread at
low elevation during the late-glacial and early Holocene.

Prior to the expansion of Sequoiadendron, the vegetation near
Exchequer meadow probably resembled Crucifereae-dominated al-

146 MADRONO [Vol. 35

pine grassland that today occurs on dry open areas with raw soil
(Major and Taylor 1977, p. 629). The dearth of tree pollen and
abundance of herb pollen, particularly Castilleja and Cruciferae (Fig.
5) indicate alpine vegetation. We infer a rapid climatic warming ca.
11,000 yr B.P.

The absence of spores of the dung fungus Sporormiella in sedi-
ments younger than ca. 11,000 yr B.P. at Exchequer Meadow may
date the extinction of the Rancholabrean megafauna in the western
Sierra. The spores are not present in Holocene sediments, but are
abundant (2.3%, 32 grains cm~ yr~') in sediments below 340 cm
(Fig. 5) equivalent to an age of 11,600 yr B.P. (Fig. 3). Sporormiella
spores (Fig. 6) are abundant in modern sediments only where in-
troduced grazing animals are plentiful, and they are even more pro-
fuse in sediments older than 11,000 yr B.P. in several sites (Davis
1987). The spores are linked directly to extinct animals by their
presence in mammoth dung (Davis et al. 1984). Although Sporor-
miella spores are not restricted to extinct animals, their presence in
late-Pleistocene sediments at Exchequer Meadow and other sites
appears to record a declining abundance of grazing animals at the
end of the Pleistocene.

CONCLUSIONS

The pollen and macrofossil records from Exchequer Meadow in-
dicate vegetation during the early Holocene resembling that found
east of the Sierra Nevada today. These records corroborate earlier
findings of aridity in the western Sierra at a time when areas to the
west and east were relatively moist, concurrent with the extinction
of Pleistocene megafauna in the area. Modern climatic contrasts
between coastal, interior, and rainshadow regions play an important
role in the vegetational differences among these areas. Paleovege-
tation data from Exchequer Meadow and other sites indicate dif-
ferent climatic and vegetational histories for these areas during the
late Quaternary. These paleoclimatic differences also may have played
a role in the differentiation of these vegetation types.

ACKNOWLEDGMENTS

Financial support for the analysis of Dinkey Meadow and Exchequer Meadow
sediments was provided by Kings River Conservation District through Clinton Blount,
Theodoratus Cultural Research Inc., Fair Oaks, California. Pollen from Dinkey and
Exchequer Meadows was counted by R. S. Anderson, O. K. Davis, K. L. Moore, and
D. S. Shafer. R. S. Anderson analyzed plant macrofossils from Dinkey Meadow. O.
K. Davis and J. A. Kailey analyzed plant macrofossils from Exchequer Meadow. We
thank C. T. Mason, curator, University of Arizona Herbarium for specimens used
in the identification of plant macrofossils.

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Department, San Diego State Univ.

1986. Late Pleistocene and Holocene vegetation and climate. Jn M. Basgall
and W. R. Hildebrandt, principal investigators, Data recovery excavations at
sites CA SHA 1176, CA SHA 1175, CA SHA 1169, and CA SHA 476, Shasta
County, California, p. 2.15-2.35. Draft Report, Calif. Dept. Transportation,
Redding District.

Woop, S. H. 1975. Holocene stratigraphy and chronology of mountain meadows,
Sierra Nevada, California. Ph.D. dissertation, California Inst. Technology, Pas-
adena.

WRIGHT, H. E. 1967. A square-rod piston sampler for lake sediments. J. Sediment.
Pet. 37:975-976.

(Received 9 Apr 1987; revision accepted 25 Jan 1988.)

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STIEBER, M. T., A. L. KARG, M. WALKER, G. D. R. BRIDSON, H. M.
BURDET, M. M. CHAUTEMpS, and T. Moruzzi-BAYo (compilers), Cat-
alogue of portraits of naturalists, mostly botanists, in the collections of
the Hunt Institute, The Linnean Society of London, and the Conser-
vatoire et Jardin Botaniques de la Ville de Genéve, pt. 1, Group portraits,
Hunt Institute for Botanical Documentation, Carnegie-Mellon Univ.,
Pittsburgh, PA 15213, 1987, xi, 93 pp., unillus., ISBN 0-913196-50-9
(paperbound), $9.00 (for U.S. and Canada, from preceding address; from
elsewhere order from Wheldon & Wesley, Lytton Lodge, Codicote,
Hitchin, Herts. SG4 8TE, England).

ANNOUNCEMENT

TWELFTH GRADUATE STUDENT MEETINGS

The California Botanical Society will sponsor the Twelfth Graduate
Student Meetings on 29 October 1988 at San Jose State University.

The presentation categories (proposed research, research in progress,
and finished research) allow the sharing of ideas and knowledge within
the graduate student community. Awards for each of these categories
will be presented at the banquet on 29 October.

For information contact Valerie Haley, Graduate Student Represen-
tative, Department of Biological Sciences, San Jose State University,
San Jose, CA 95192.

SPECIES FREQUENCY IN RELATION TO TIMBER
HARVEST METHODS AND ELEVATION IN THE
PINE TYPE OF NORTHEAST CALIFORNIA

ROBIN S. VORA
U.S. Fish & Wildlife Service, Room 334, Federal Building,
Brunswick, GA 31520

ABSTRACT

Few trends in species presence and frequency due to different timber harvest treat-
ments were apparent 40 years after logging in the pine type of northeastern California.
Ceanothus prostratus was generally absent from units where all trees larger than 29.5
cm dbh had been removed 40 years ago. Calocedrus decurrens was more frequent in
uncut control plots. Species richness was constant across the area. Differences in
species frequencies among experimental blocks located at five elevations suggested
at least two major plant communities within the forested zone, with elevation a major
environmental influence. A list of common plant species and their frequencies are
provided for the Blacks Mountain Experimental Forest in northeastern California.

The pine type of northeastern California has received minimal
botanical study. It has been described as the eastside phase of mixed
conifer forest (Griffin and Critchfield 1972), yellow pine forest by
Munz (1973), interior ponderosa pine by the Society of American
Foresters (Barrett et al. 1980), and eastside pine type by McDonald
(1983). Vasek (1978) studied forests dominated by Pinus jeffreyi
between 1280 and 1950 m elevation. He noted that these forests
ranged along moisture and temperature gradients across the region,
with local variation influenced by elevation, aspect, surface rock and
soil depth. In higher montane forests, Rundel et al. (1977) suggested
that Calocedrus decurrens and Abies concolor have both increased
their relative and absolute densities since the turn of the century
when fire control programs went into effect. Logging and widespread
grazing may have caused compositional changes.

At the Blacks Mountain Experimental Forest, the U.S. Forest
Service has undertaken several long-term studies on the growth and
harvest of the major tree species: Pinus ponderosa, Pinus jeffreyi,
Abies concolor, and Calocedrus decurrens (Hallin 1959, Robson and
Standiford 1983). Roy (1946) studied the sagebrush flats within the
Experimental Forest. A list of the most abundant range plant species
has been compiled for nearby Harvey Valley Range Allotment (Hor-
may 1959). Similar lists are available for the California State Uni-
versity at Chico field station at Eagle Lake (R. Schlising pers. comm.).
Hormay (1940) documented some impacts of logging on forage

MADRONO, Vol. 35, No. 2, pp. 150-158, 1988

1988] VORA: SPECIES FREQUENCY IN PINE TYPE 151

Oregon 121

eas erry

UU4pptts

BLACKS MOUNTAIN
EXPERIMENTAL FOREST

ay 4

//
Redding®@® Cf,
Mt. Lassen® Meg

Susanville
vA

California Oe,

Fic. 1. Location of the Blacks Mountain Experimental Forest and pine type of
northeast California (diagonal lines) (Hallin 1959).

plants. The impacts of cattle grazing on Purshia tridentata have been
studied extensively (Hormay 1943, Neal 1982).

Hallin (1959) stated that the Blacks Mountain Experimental For-
est 1s representative of the pine type of northeastern California. As
a study site, 1t provided the opportunity to examine the presence of
understory species in relation to well-planned and documented tim-
ber harvest research. I undertook exploratory analyses to identify
differences in species frequency due to timber harvest methods 40
years after logging, and to note changes in species presence with
elevation.

STUDY SITE

The Blacks Mountain Experimental Forest is located in the Lassen
National Forest, Lassen Co., California (Fig. 1). Approximately half
of the study site lies in a gently rolling basin; the remainder extends
up moderate slopes to the north and east. Elevations range from
1700 to 2100 m. The Forest is composed of a mosaic of small, even-
aged groups of trees of various ages that vary in size from a fraction
of a hectare to 4 ha, and by scattered older trees in younger stands
(Hallin 1959). Forest gaps, or openings, are characteristic of these
forests (Franklin and Dyrness 1973).

Hallin (1959) reported that the mean annual precipitation varied
from 23 to 74 cm, and averaged 46 cm during the period 1935-53.
About 90% of the precipitation falls in the months of October through
May. McDonald (1983) and Franklin and Dyrness (1973) state that

152 MADRONO [Vol. 35

soil moisture becomes depleted rapidly and warm summer temper-
atures and low humidity increase evapotranspiration to the point of
creating moisture-deficient soil. Consequently, the growing season
for an average ponderosa pine (Pinus ponderosa), for example, lasts
only from | May to about 15 June in a typical year. Seasonal brows-
ing or grazing of plants by herbivores occurs between May and
October.

METHODS

Hallin (1959) described timber harvest studies initiated in 1938.
A randomized block design with six treatments was used. Four to
six treatments were randomly allocated to 8-ha macroplots within
each block (Fig. 2). Treatments (listed by increasing intensity of cut)
were: (1) control (C)—no cutting; (2) sanitation salvage (IS)—re-
moval of diseased, dead and dying trees; (3) unit area control (UAC)—
group selection removal of aggregations of mature trees combined
with sanitation salvage in the stand; (4) moderate selection (MFS)—
approximately 55% of the volume cut; (5) heavy selection (HFS)—
approximately 75% of the volume cut; and (6) “‘clear-cut”? (CC)—
removal of trees 29.5 cm dbh and greater and natural regrowth.
Timber harvesting discriminated against Abies concolor and Calo-
cedrus decurrens (a greater percentage of trees of these species were
removed compared to pine species).

I sampled five blocks that represented site elevations (Fig. 2).
Experimental design, however, did not control for variables such as
slope, aspect, soil depth and stoniness, and seed source. No major
soil differences were apparent among the blocks based on profile
descriptions in a soil survey (Storie et al. 1940). Three blocks [39,
40, and 42 (the number represents the year of logging)] included all
six treatments, whereas the IS and CC treatments were absent from
two blocks (43 and 47).

Ten 25-m? microplots were located randomly along a transect
within each macroplot. A species list was recorded for each microplot
and from this frequency per 25 m7? (frequency) estimated for each
macroplot. Plant nomenclature was based on Munz (1973). It was
not feasible to compile a complete list in the field. I was not able to
identify several species found without flowers. Individual plants
often lacked inflorescences when I visited the microplots. Cattle
grazing contributed to this problem in several areas. Therefore, I
was unable to gather data on frequency of occurrences of grasses,
penstemons, and a few other species. Pinus ponderosa and P. jeffreyi
were sampled and analyzed as one species because of potential hy-
bridization and difficulty in separating the two in field identification
(Vasek 1978). Sampling was done between June and August 1983.
This period followed an unusually wet winter for the region (167%

1988] VORA: SPECIES FREQUENCY IN PINE TYPE 153

1950
1900

a

yd iM
ay

an

Patterson
Flat

scale

Fic. 2. Elevation contours (m), cutting blocks, and treatment units (C = control;
IS = sanitation salvage; UAC = unit area control or group selection; MFS = moderate
selection; HFS = heavy selection; CC = clear cut).

of normal precipitation occurred at a U.S. Forest Service weather
Station at 1390 m, and located 58 km to the southeast).

There was a total of 26 macroplots within the five blocks, and
frequency studies were based on these twenty-six 8-ha units (Fig. 2).
I performed analyses of variance (ANOVA) to identify differences
(p < 0.05) among harvest treatments and among blocks.

RESULTS

Ninety-five species were identified within the 330 microplots. The
woody plants, especially the trees, dominated cover or biomass. In
terms of numbers of species, the dominant plant families were As-
teraceae (17 species, 18%), Poaceae (11 species, 12%), and Scroph-
ulariaceae (12 species, 13%). Consideration also of species frequency

154 MADRONO [Vol. 35

would have added Pinaceae, Rhamnaceae, Laminaceae, Rubiaceae,
Ericaceae, Cupressaceae, and Rosaceae to the list of major plant
families. The overall (all 26 macroplots) standard deviation of mean
frequency of major species ranged from 5 to 20%; variances were
greater within each treatment or block.

Few trends in species presence and frequency due to timber harvest
treatments were apparent 40 years after logging (Table 1). Pine species
frequency was lower in the CC treatment, but not significantly dif-
ferent (p = 0.11) from the control. Ceanothus prostratus was gen-
erally absent from the CC treatment (p = 0.02). Calocedrus decurrens
was more frequent in control plots, but frequencies were not statis-
tically different from other treatments (p = 0.19).

Species richness ranged from 27 to 37 per block. Frequency of
species of Poaceae lumped together was higher in the lower elevation
blocks (ANOVA p = 0.02).

Field observation and examination of the block data in Table 2
suggest that Blacks Mountain Experimental Forest is located in a
transition zone between two major plant communities. Table 2 lists
the plant species that characterize the two communities, including
those that were significantly different among blocks. The lower el-
evation community (below approximately 1800 m) is typical of what
is commonly referred to as eastside pine or interior ponderosa pine,
with the overstory dominated almost exclusively by Pinus ponderosa
and P. jeffreyi, and the understory dominated by Purshia tridentata
and Artemisia tridentata. Frequency of finding one plant of the Po-
aceae in a 25-m? plot was 100%. The overstory in the higher ele-
vation community is characterized by the same two pine species
and two additional conifers, Abies concolor and Calocedrus decur-
rens. Typical understory species are Symphoricarpos vaccinoides and
Monardella odoratissima. Frequency of finding one plant of the Fam-
ily Poaceae in a 25-m? plot was 70%.

DISCUSSION

Vasek (1978) observed that despite logging, the forests of the
southern Modoc Forest had considerable resemblance to pristine
vegetation. Few differences in species frequency were observed on
Blacks Mountain 40 years after logging. Perhaps the plants of this
forest are adapted to a disclimax state. Prior to 1900, insects and
fire maintained forests in a variety of successional stages. Pine mor-
tality caused by western bark beetle (Dendroctonus brevicomis) is
still heavy (Hallin 1959, Hart 1983). Fire frequencies of 6-36 years
are reported in the Lava Beds National Monument [110 km to the
north by Johnson and Smathers (1976) and Martin and Johnson
(1979)], and eastern Oregon by Soeriaatmadja (1966). Logging and
grazing by cattle and sheep have influenced succession over the past

1988] VORA: SPECIES FREQUENCY IN PINE TYPE Ess)

TABLE |. PLANT SPECIES FREQUENCY IN 40-YEAR-OLD CUT UNITS ON BLACKS
MOUNTAIN EXPERIMENTAL FOREST. Means are presented for common species (> 5%
overall mean frequency) for each treatment and block, and are estimated from 10
25-m~? plots in each 8-ha unit. A dash (—) indicates frequency < 1%. Treatment codes:
C = control; IS = sanitation salvage; UAC = unit area control; MFS = moderate
selection; HFS = heavy selection; CC = “clear-cut”. IS and CC treatments were
absent irom blocks 43 and 47. Block numbers refer to the year they were cut, and
are listed in ascending elevation (39— 1750 m, 43—1800 m, 47-1830 m, 42—1875
m, and 40—1920 m).

Treatment Block

C IS UAC MFS HFS CC 39 43 47 42 40
5

No. of cutting units 3 5 5 5 3 6 4 4 6 6

Abies concolor 44 53 46 54 52 63 — 10 72 85 82
Arabis holboellii 6 “17> 22 220° 10% 30 — — 28 29 25
Arctostaphylos patula 26 7 10 22 = «34 7 10 45 65 21 33
Artemisia tridentata 2 — 8 6 12 33 400 -—- —- — =
Calocedrus decurrens 50 Mal 30" 722° < 32° (13 — 20 80 28 32
Castilleja applegatei 8 7 20 = 18 16 10 5 13 25° 42 “18
Ceanothus prostratus 68 57 78 68 £54 3 47 85 90 52 52
Ceanothus velutinus 6 — 14 16 4 7 — — 25 18 2
Chrysothamnus

nNauSeosus 8 6 10 22 16 20 40 10 5 3 7
Collinsia torreyi 18 13 30 32 30 40 38 43 15 23 #18
Crepis acuminata 2 4 2 8 100 — 2 23 — 5 —
Cryptantha affinis 8 10 20 18 10 30 2 — 3 3 15
Eriophyllum lanatum 16 “20° 22. 34 °28 27 8 27 33 20 38
Fritillaria atropurpurea 6 17 22 18 10 3 - -— 5 17 2
Gayophytum humile 8 — 8 14 10 33 20 28 — 5 7
Hieraceum albiflorum — 10 7 24 #18 #17 | — — 25 21 15
Lupinus caudatus 24 20 18 24 10 — 20 18 13 £6 26
Microseris nutans 16 — 16 8 12 20 7. ls <5 (ae
Microsteris gracilis 10 7 8 8 2 3 8 5 — 10 6
Monardella odoratis-

sima 44 43 56 60 36 43 13 53 45 68 60
Pedicularis semibar-

bata 4 13 2 8 6 6 8 — 8 10 3
Pinus ponderosa and

Pinus jeffreyi 100 80 88 86 92 70 88 88 83 87 93
Purshia tridentata 26 33 22 28 26 33 90 38 — — 3

Senecio integerrimus 26 37 30 34 36 = 33 12 28 28 55 37
Symphoricarpos vac-

cinoides 42 53 40 54 32 60 — 8 43 82 82
Viola purpurea 16 16 12 24 24 20 £48 25 5 3 10
Wyethia mollis 36 © 3606-332.—C— 332 3384S 17 56 78 2 10 18

100 years. Plant reestablishment is slow, especially in xeric envi-
ronments (Franklin and Dyrness 1973), yet 40 years is perhaps suf-
ficient time for many species to at least partially recover. This is
aided greatly by the heterogeneous nature of the forest, especially
the “‘gap”’ openings.

156 MADRONO [Vol. 35

TABLE 2. PLANT SPECIES THAT CHARACTERIZE THE TWO COMMUNITIES IN THE FOREST
ZONE OF BLACKS MOUNTAIN EXPERIMENTAL FOREST. The numbers in parentheses are
the probability of type I error estimated from Analysis of Variance (e.g., p < 0.01
for Purshia tridentata). Frequency data are estimated from 10 25-m? microplots in
each 8-ha macroplot. Species are listed in declining order of overall mean frequency.
Species of the Family Poaceae are not included. Pinus ponderosa, Pinus jeffreyi, and
Ceanothus prostratus are common in both communities and therefore not listed.
' Elevation 1700-1830, m, 10 macroplots sampled. ? Elevation 1800-1950 m, 16
macroplots sampled. * Not present on plots in other community. * More frequent at
lower elevations.

Community

Pinus ponderosa/Purshia
tridentata (PIPO/PUTR)!

Purshia tridentata (<0.01)
Wyethia mollis (<0.01)

Viola purpurea (<0.01)

Artemisia tridentata®
Chrysothamnus nauseosus (<0.01)
Collinsia torreyi (0.12)*

Achillea millefolium>

Cercocarpus ledifolius*

Pinus ponderosa/Symphoricarpos
vaccinoides (PIPO/SY VA)

Abies concolor (<0.01)
Symphoricarpos vaccinoides (0.06)
Monardella odoratissima (<0.01)
Senecio integerrimus (0.04)
Arabis holboellii (<0.01)
Calocedrus decurrens (<0.01)
Collinsia torreyi (0.12)
Hieraceum albiflorum (0.06)

Arctostaphylos patula (0.06)
Eriophyllum lanatum (0.08)
Ceanothus velutinus (<0.01)

The results of this study, however, do not imply that timber har-
vest has no effect on the plant community 40 years after logging in
the eastside pine type. Differences in vegetation structure were ap-
parent between the control and more heavily-cut units (e.g., more
large snags in the control). The results merely state that long-term
effects of timber harvest were not observed in frequency or presence
data.

Greater ground disturbance in the CC treatment may have reduced
populations of Ceanothus prostratus (a prostrate shrub). Table 1
shows that several other species were absent on plots in the CC and
IS treatments, or had higher or lower frequencies in the CC treat-
ment; more than three replications of these treatments, and perhaps
more than 10 microplots in each unit (replication) were probably
necessary to confirm significance.

I estimated that there were 110 to 140 plant species in the forested
zone. The total observable flora was noticeably greater during the
first summer (1983) following record precipitation, and richness of
the flora was probably greater than during a year of average precip-
itation.

Elevation appeared to be the single most important environmental
variable, with a change of as little as 60 m influencing species pres-

1988] VORA: SPECIES FREQUENCY IN PINE TYPE 157

ence. Higher frequency of grasses at lower elevations was probably
due to decreased canopy cover and increased light filtration in the
lower elevation pine forest. Precipitation and soil moisture appeared
to increase with elevation. Measurement of these physical influences,
as well as aspect and other soil characteristics, is needed to identify
which of them may have also contributed to differences in plant
communities associated with elevation.

ACKNOWLEDGMENTS

I thank Robert Schlising, Wayne Dakan, Gary Schoolcraft, Tom Janecke, and Steve
Brunsfeld for assistance with plant identification. Karen Toor helped greatly with
field collection of plants and other data. Dale Everson, Peter Mika, James Norris,
Robert Brewster, and Karen Falke assisted me with statistical analyses and use of the
computer. William Laudenslayer, Jr., Robert Powers, Douglas Roy, Fred Johnson,
Paul Opler, Brenda Smith, Robert Schlising, Wayne Ferren, and William Critchfield
assisted with editing this paper.

LITERATURE CITED

BARRETT, J. W., P. M. MCDONALD, F. RONCO, JR., and R. A. RYKER. 1980. Interior
ponderosa pine. /n F. H. Eyre, ed., Forest cover types of the United States and
Canada, p. 114-115. Soc. Amer. Foresters, Washington, DC.

FRANKLIN, J. F. and C. T. DyrngEss. 1973. Natural vegetation of Oregon and Wash-
ington. U.S.D.A. For. Serv. Gen. Tech. Rep. PNW-8.

GRIFFIN, J. W. and W. B. CRITCHFIELD. 1972. The distribution of forest trees in
California. U.S.D.A. For. Serv. Res. Paper PSW-82.

HALLIN, W. E. 1959. The application of unit area control in the management of
ponderosa-Jeffrey pine at Blacks Mountain Experimental Forest. U.S.D.A. For.
Serv. Tech. Bull. 1191.

Hart, D. 1983. Tree mortality on the method-of-cutting treatment plots at Blacks
Mountain Experimental Forest for the years 1980 through 1981. U.S.D.A. For.
Serv. R5 For. Pest Managem. Staff, San Francisco, CA, unpubl.

Hormay, A. L. 1940. The effect of logging on forage. Chron. Bot. 6:6-7.

1943. Bitterbrush in California. U.S.D.A. For. Serv. Calif. For. Range Exp.

Sta. Res. Note 34.

. 1959. Most abundant range plant species by vegetation types, Harvey Valley
Range Allotment, Lassen National Forest. U.S.D.A. For. Serv. PSW, Berkeley,
CA, unpubl.

JOHNSON, A. H. and G. A. SMATHERS. 1976. Fire history and ecology of Lava Beds
National Monument. Proc. Ann. Tall Timbers Fire Ecol. Conf. 15:103-115.

Martin, R. E. and A. H. JOHNSON. 1979. Fire management of Lava Beds National
Monument. Jn Proc. First Conf. Sci. Res. Natl. Parks, Vol. II (9-12 Nov. 1976,
New Orleans, LA), p. 1209-1217. U.S.D.I. Natl. Park Serv. Trans. and Proc.
Ser...

McDONALD, P. M. 1983. Climate, history, and vegetation of the eastside pine type
in California. Jn T. F. Robson and R. B. Standiford, eds., Management of the
eastside pine type in northeastern California, proceedings of a symposium, p. 1l—
16. N. Calif. Soc. Amer. For. 83-06. Arcata, CA.

Munz, P. A. 1973. A California flora and supplement. Univ. California Press,
Berkeley.

NEAL, D. L. 1982. Improvement of Great Basin deer winter range with livestock
grazing. Jn Proc. wildlife-livestock relationships symposium, p. 61-73. Coeur
d’Alene, ID.

158 MADRONO [Vol. 35

Rosson, T. F. and R. B. STANDIFORD. 1983. Management of the eastside pine type
in northeastern California, proceedings of a symposium. N. Calif. Soc. Amer.
For., SAF 83-06.

Roy, D. F. 1946. Some factors affecting establishment of pine reproduction on
sagebrush-flat edges in northeastern California. M.S. thesis, Univ. California,
Berkeley.

RUNDEL, P. W., D. J. PARSONS, and D. T. GORDON. 1977. Montane and subalpine
vegetation of the Sierra Nevada and Cascade Ranges. Jn M. G. Barbour and J.
Major, eds., Terrestrial vegetation of California, p. 559-599. John Wiley & Sons,
NY.

SOERIAATMADJA, R. E. 1966. Fire history of the ponderosa pine forests of the Warm
Springs Indian Reservation, Oregon. Ph.D. dissertation, Oregon State Univ.,
Corvallis. Abstract in Diss. Abstr. Int. 27:2612-B.

STORIE, R. E., W. W. WEIR, and R. C. Cote. 1940. Soil survey of Blacks Mountain
Experimental Forest. Division of Soils, Univ. California, Berkeley. On file at
U.S.D.A., For. Serv., PSW, Redding, CA.

VASEK, F. C. 1978. Jeffrey pine and vegetation of the southern Modoc National
Forest. Madrono 25:9-30.

(Received 25 Feb 1987; revision accepted 30 Nov 1987.)

ANNOUNCEMENT
NEw PUBLICATIONS

GABRIELSON, P. W., R. F. SCAGEL, and T. B. Wippowson, Keys to the
benthic marine algae of British Columbia, northern Washington and
southeast Alaska, Phycological Contribution, no. 2, pp. i-ili, 1-197, Mar
1987, ISSN 0831-4861, ISBN 0-88865-461-8, Can $6.50 (from R. F.
Scagel, Dept. Botany, Univ. of British Columbia, 3529-6270 University
Blvd., Vancouver, BC V6T 2B1). [Reproduced from camera-ready copy.

For the related “‘A synopsis of the benthic marine algae of British Co-
lumbia, northern Washington and southeast Alaska,’ PC, no. 1, see
review by I. A. Abbott, Taxon 36:670-671.]

GONZALEZ VILLARREAL, L. M., Contribucion al conocimiento del genera
Quercus (Fagaceae) en el Estado de Jalisco, Instituto de Botanica, Uni-
versidad de Guadalajara, Apartado 139, Zapopan 45110, Jalisco, 7 Jan
1987, 240, [1] pp., illus., ISBN 968-895-027-0 (paperbound), US $14.00.
[The first volume in Colleci6n flora de Jalisco—editorship not indicated.
Treatment of the species of Quercus, with maps for species, glossary,
lists of exsiccatae.]

NOTES

REPORT ON THE XIV INTERNATIONAL BOTANICAL CONGRESS.—The XIV Interna-
tional Botanical Congress was held in West Berlin during the period 20 July through
1 August 1987. There were 14 pre-Congress and 17 post-Congress botanical excur-
sions, extending to the far reaches of Western Europe. Before the Congress, for ex-
ample, Cherie Wetzel and Larry Heckard joined a small group led by Prof. Arne Strid
of the University of Copenhagen in a botanical tour of northern Greece, especially
Mount Olympus, while Elizabeth McClintock explored the Canary Islands and Wil-
liam Sanders, a graduate student at Berkeley, collected lichens in Sardinia.

After the Congress, Larry Heckard and I, along with Tom and Mary Fuller and
their son Ken, participated in a memorable tour of Czechoslovakia under the guidance
of numerous Czech and Slovak botanists, especially Dr. Jan Jenik and Dr. Viera
Ferakova. Unlike the excursions sponsored by the XII International Botanical Con-
gress in the Soviet Union in 1975, over which the government tourist bureau had
rigid control, our program was mainly in the hands of botanists. The politics of the
excursion were obvious and amusing. Czechoslovakia comprises two ethnic and
political entities, the Czech Socialist Republic with its capital at Prague and the Slovak
Socialist Republic with its capital at Bratislava. Everything was counterbalanced. We
visited high mountains, lowlands, and limestone outcrops in Bohemia, and high
mountains, lowlands, and limestone outcrops in Slovakia. Czechoslovakia has a long
and strong botanical tradition; its flora and vegetation have been studied so intensively
that only minutiae remain. At times we were exposed to more details than we could
absorb, but it was a small price to pay for the privilege of seeing the spectacular plants
and scenery of Czechoslovakia and sharing the knowledge and warmth of her won-
derfully hospitable people.

The Congress itself was held in the monumental International Congress Centre,
conveniently served by Berlin’s efficient subway and bus systems. The main building
staggers the imagination. It is 300 m long, 80 m wide, and 40 m high. Inside this
enormous shell are facilities sufficient to meet the ordinary and special needs of a far
larger assemblage than the 4130 botanists and companions of this Congress. Foremost
is the main theater, its grand proportions recalling the Radio City Music Hall in New
York City. Here is where the opening and closing ceremonies were held. The opening
ceremony will be remembered by many of us for the truly outstanding performance
by the Westphalian Symphony Orchestra conducted by Walter Gillessen. After an
opening fanfare, the ceremony began with vigor and excitement by the playing of
Brahms’s Academic Festival Overture. Brief addresses were given by the President of
the Congress (Karl Esser), the Senator for Science and Research for West Berlin
(George Turner), the Vice-President of the International Union of Biological Sciences
(David Ride), the President of the German Botanical Society (Wilhelm Nultsch), and
the Honorary President of the Congress (Frans Stafleu). The ceremony closed with a
magnificent concert of German music, running the gamut from Wagner’s Prelude to
Die Meistersinger to Johann Strauss’s Overture to Die Fledermaus, and including
major works by Beethoven, Schumann, and Richard Strauss. Immediately after the
opening ceremony, there was a reception, where many members of the California
Botanical Society crossed paths and welcomed one another. Besides those already
mentioned, I happened to meet Herbert and Irene Baker, Doug Barbie, Tania Beliz,
Winslow Briggs, Heidi Dobson, Kent Holsinger, Donald Kaplan, David Keil,
Stephanie Mayer, Jeanine Olsen, Ledyard Stebbins, John Thomas, Nancy Vivrette,
and Grady Webster.

In addition to the main theater, there are two small theaters with banks of seats
that can be retracted to the ceiling, leaving a vast open space. Here is where the

MADRONO, Vol. 35, No. 2, pp. 159-163, 1988

160 MADRONO [Vol. 35

Congress banquet was held, with more than a thousand persons served nearly si-
multaneously. The smaller meeting rooms varied greatly in their desirability. Topping
the list was the roof garden, where the Nomenclature Section met during the week
preceding the regular Congress. From the roof of the Congress Centre a panoramic
vista of West Berlin and the edge of East Berlin could be obtained, although there
was Often interference from ominous clouds that brought some rain nearly every day.
At the bottom of the list were small meeting spaces cut off from main passageways
only by portable partitions. The consensus seemed to be that while nearly everyone
was initially turned off by the sheer size and impersonal nature of the Congress
Centre, within a few days the dissidents had been converted and looked forward each
morning to rejoining their spaceship, which offered comfortable, spacious, and im-
maculate facilities to accommodate all conceivable daily needs. Next to spaciousness,
the most obvious hallmark of the center is the extremely high caliber of materials
and workmanship. There are miles of chrome and acres of black synthetic leather.
The seats in the main theater are undoubtedly the most comfortable and most ex-
pensive I have ever experienced, offering special lumbar support and a reading light.

As for the scientific program, the Congress was very well organized, but there were
too many events, even more so than is usually the case. There were 24 general lectures,
224 symposia, 123 poster sessions, 36 special interest meetings, and 25 society meet-
ings. One of my days began with a poster session at 8:30 in the morning and ended
at the close of a symposium at 10 in the evening! In my field (phycology), the symposia
were generally disappointing, some of the presentations being rehashes of papers
given two years earlier at an international phycological congress in Copenhagen. I
thought that the contributed papers were more interesting, with some of the best
work being presented as posters. Unfortunately, the poster area was extremely crowd-
ed. Poster presentations were formalized, being grouped into sessions, each chaired
by a moderator. One of the best posters, incidentally, was by Bob Haller of the
University of California at Santa Barbara, on the distribution, evolution, and sys-
tematics of western American yellow pines.

Despite the fact that most papers dealt with aspects of botany other than those that
would be of greatest interest to members of the California Botanical Society, the
leadership of the Congress was largely in the hands of taxonomists, reflecting the great
strength and importance of the Berlin Botanical Museum and Garden. Its director,
Werner Greuter, was chairman of the organizing committee. We were often reminded
of the remarkable contributions of a long succession of eminent botanists in Berlin,
especially Adolf Engler. During the nomenclature sessions, a magnificent new wing
to the botanical museum and library was dedicated. At the closing ceremony, the
International Association of Plant Taxonomists awarded the first Engler gold medal
to Frans Stafleu in recognition of his enormous contributions to taxonomic botany.
The next award of the Engler medal will be made at the X Vth Congress in Tokyo in
1993.

What was accomplished at the Congress? As usual the greatest benefit came from
personal contacts rather than from formal presentations. The motto of the Congress
was “‘Forests of the World” and attention was focused on the serious plight of our
rain forests as well as the equally serious plight of European woodlands. In some
parts of Europe a quarter of the trees are dead or dying as a result of atmospheric
pollution.

With regard to nomenclature, we were faced with a record number of proposals—
334—almost all of which were defeated, either by a preliminary mail vote or by a
vote on the floor. Stafleu skillfully but autocratically kept the sessions moving, often
so rapidly that confusion ensued. Towards the end of the sessions, fatigue set in, and
numerous proposals were referred either to the Editorial Committee or to a series of
ad hoc committees charged with making their reports prior to the Tokyo Congress.

In December 1987, the Editorial Committee received a compilation of the proposals
accepted by the Congress or referred to them for discretionary action, and we met in
Berlin during the first week of January to write the new Code. Most changes are of

1988] NOTES 161

the nature of clarification or elimination of conflicting rules. The only fundamental
change concerns lectotypification. Implicit lectotypification, that is, lectotypification
expressed by a taxonomic treatment rather than by an explicit statement, has been
outlawed, both in the past and in the future. Thousands of lectotypifications may be
affected, especially at the level of species and infraspecific taxa, but the full effect of
the new ruling will not be known for many years. I should mention that the official
Berlin Code will be in English only, resulting in a prompter, smaller, and less expensive
publication compared to previous versions.

Finally, I want to say that all the congressists enjoyed Berlin and Berliners. It is an
open-minded, cosmopolitan city, making up for its lack of beauty by its tremendous
energy and excitement.— PAUL C. SILvA, Herbarium, Department of Botany, Uni-
versity of California, Berkeley 94720. (Received 9 Nov 1987; revision accepted 22
Feb 1988.)

TYPIFICATION OF Chaenactis alpina (ASTERACEAE).— Asa Gray cited no specimens
when he described Chaenactis douglasii var. alpina (Synoptical Fl. N. Amer. 17:341,
1884). The range was given as “Alpine region of the Rocky and Cascade Mountains
in Colorado and Wyoming, of the Sierra Nevada, California, and north to Washington
Terr.”’ Stockwell (Contr. Dudley Herb. 3:113, 1940) designated a type, ““Alta, Wasatch
Mountains, Utah, M. E. Jones 1232. (NY) and stated ““Type of A. Gray not known.”
There is no indication on the sheet that Gray ever saw this specimen. In GH there
are at least five collections prior to 1884 with the name “‘var. alpina” and “‘Syn. FI.”
on the sheets, including two collections each from California (Hooker and Gray s.n.
in 1877, Brewer 1901) and Colorado (Parry 55, Hall and Harbour 283) and one
collection possibly from Wyoming (not labeled but next to label for C. douglasii
specimen from Wyoming). These account for Gray’s distribution except for Wash-
ington Territory. One sheet contains a fragment collected by Geyer apparently at
Spokane Falls but belongs to another species. It can be safely concluded that this
material represents some or all of that which Gray used to describe var. alpina, and
the lectotype must therefore be chosen from among these specimens [Art. T.4.(a),
ICBN].

There are two elements represented in the specimens I take to be type material, a
glandular or viscid-hirsute element and a tomentose or lanate element. Only the Hall
and Harbour specimen fits Gray’s description completely because it is the only spec-
imen with complete rootstocks. It is not, however, the typical variety of Stockwell
and others (Harrington, Manual Pl. Colorado 588, 1964; Welsh et al., A Utah FI.
163, 1987). Another specimen (Parry 55), which fits Gray’s protologue except for
lacking complete rootstocks, is therefore chosen as the lectotype in order to preserve
current usage [Art. T.4.(e), ICBN].

Stockwell’s varieties rubella and leucopsis appear to be the same taxon. Var. leu-
copsis 1s taken up here to be consistent with Harrington (Manual Pl. Colorado 588,
1964) and Welsh (Great Basin Naturalist 43:235, 1983). The nomenclature is sum-
marized below.

CHAENACTIS ALPINA (Gray) Jones, Proc. Calif. Acad. Sci. II, 5:699. 1895.— Chaenactis
Douglasii Hook. & Arn. var. alpina Gray, Synoptical Fl. N. Amer. 17:341. 1884.—
Lectotype: CO, headwaters of Clear Creek and alpine ridges e. of Middle Park,
1861, Parry 55 (GH)!).

Chaenactis pedicularia Greene, Pittonia 4:98. 1899. Holotype: CO, La Plata Mts.,
Little Kate Mine, 11,500 ft, Baker, Earle, and Tracy 536, 16 Jul 1898 (ND-G;
isotype: RM!, US).

Chaenactis pumila Greene, Leafl. Bot. Observ. Crit. 2:221. 1912. Holotype: CA, peak
near Sonora Pass, 11,500 ft, Brewer 1901 (US; isotype: GH!).

CHAENACTIS ALPINA (Gray) Jones var. LEUCOPSIS (Greene) Cock. ex. Stockw., Contr.
Dudley Herb. 3:114. 1940.—Chaenactis leucopsis Greene, Leafl. Bot. Observ.

162 MADRONO [Vol. 35

Crit. 2:221. 1912.—Chaenactis alpina leucopsis (Greene) Cock., Univ. Colorado
Stud. 11:218. 1915. Holotype: CO, Needle Mountains, 14 Jul 1901, Whitman
Cross 61 (US!).

Chaenactis rubella Greene, Leafl. Bot. Observ. Crit. 2:222. 1912.—Chaenactis alpina
var. rubella (Greene) Stockw., Contr. Dudley Herb. 3:114. 1940. Holotype: north-
west Wyoming, 31 Aug 1893, J. N. Rose 298 (US!).

Variety alpina has peduncles and involucres glandular to densely viscid-hirsute
with occasionally two to several heads per scape. Variety /eucopsis has peduncles and
involucres tomentose or lanate with usually one head per scape.

Loan of type specimens by US and GH and use of facilities at RM are gratefully
acknowledged. Barbara Hellenthal checked for type material at ND-G.— RosBErT D.
Dorn, Box 1471, Cheyenne, WY 82003. (Received 31 March 1987; revision accepted
30 Nov 1987.)

Chenopodium simplex, AN OLDER NAME FOR C. gigantospermum (CHENOPODI-
ACEAE).— Edwin James with the Long Expedition to the Rocky Mountains collected
a species of Chenopodium in 1820 that John Torrey described in 1827 as a new
variety of the European C. hybridum L. Torrey thought that it might be a new species.
Rafinesque raised it to a species in 1832. These names, C. hybridum var. simplex
Torrey and C. simplex (Torrey) Raf., apparently have been largely overlooked ever
since. Standley [N. Amer. Flora 21(1):13, 1916] and Wahl (Bartonia 27:30, 1954) do
not include them in their treatments, but the Rafinesque combination does appear
in Merrill (Index Rafinesquianus, The Arnold Arboretum, p. 118, 1949), and Torrey’s
variety appears in the Gray Herbarium Index. The holotype is the North American
plant that has been called C. gigantospermum or C. hybridum var. gigantospermum.
These names must be replaced by C. simplex or C. hybridum var. simplex, respec-
tively.

Bassett and Crompton (Canad. J. Bot. 60:600, 1982) selected a Macoun specimen
at CAN for the lectotype of C. gigantospermum Aellen. Article T. 4. (c) of the
International Code states: “If no holotype was designated by the original author and
if syntypes (Art. 7.7) exist, one of them must be chosen as the lectotype.” The CAN
specimen, therefore, would be a duplicate of the lectotype and the Macoun specimen
at US would be the lectotype because Aellen cited only specimens from US (except
for one in his own herbarium). Wahl (Bartonia 27:16, 30, 1954), Baronov (Rhodora
66:168—-171, 1964), and Bassett and Crompton (Canad. J. Bot. 60:600, 1982) discuss
the differences between the European C. hybridum and the North American C. simplex
(as C. gigantospermum). The nomenclature is summarized below.

CHENOPODIUM SIMPLEX (Torrey) Raf., Atlantic J. 1:146. 1832.—Chenopodium hy-
bridum, B? simplex Torrey, Ann. Lyceum Nat. Hist. New York 2:239. 1827.—
Holotype: ‘““Near Council Bluff, on the Missouri,” Edwin James s.n. in 1820
(NY!).

Chenopodium gigantospermum Aellen, Feddes Repert. Spec. Nov. Regni Veg. 26:
144. 1929.—Chenopodium hybridum var. gigantospermum (Aellen) Rouleau,
Naturaliste Canad. 71:268. 1944.—Lectotype by Bassett and Crompton (Canad.
J. Bot. 60:600, 1982): British Columbia, Vernon, 9 Jul 1889, Macoun s.n. (US,
photo RM!; isolectotype CAN, photo DAO).

Loan of type material by NY and use of facilities at RM are gratefully acknowl-
edged. — RoBERT D. Dorn, Box 1471, Cheyenne, WY 82003. (Received 30 Mar 1987;
revision accepted 30 Nov 1987.)

Arabis breweri S. WATS. VAR. austinae (GREENE) ROLL. (CRUCIFERAE)— Ventura
Co.: Rose Valley Falls, Sespe Valley, 7 Mar 1947, Pollard s.n. (CAS). Monterey Co.:
Santa Lucia Range, n. slope of Twin Peak, trail between Goat Camp and Trail Spring

1988] NOTES 163

Camp, T21S R4E S34, 1130 m, 2 Feb 1984, Haller 3600 (UCSB). Colusa Co.: ridge
above Red Bridge Camp and confluence of n. and middle forks of Stony Creek, on
serpentine, 580-640 m, 16 Apr 1950, Bacigalupi 3106 (JEPS, UC). Tehama Co.: ca.
10.9 km sw. of Paskenta, on metavolcanic outcrops, T23N R7W sw.'4 828, 793 m,
25 Apr 1986, Preston 534 (CAS, DAV). Trinity Co.: dry rocky hillside above Deer
Lick Springs Rd., 4.8 km from junction with Hwy. 3, 12 May 1979, York 271 (HSC).
Siskiyou Co.: near Yreka, along Shasta River, 20 Apr 1934, Eastwood and Howell
1762 (DS, RSA, UC). Shasta Co.: canyon of Low Pass Creek, base of limestone cliffs,
460 m, 30 Jun 1959, Bacigalupi 7176 (JEPS); n. side of McLoud arm of Lake Shasta,
off Gilman Rd., 24 km from U.S. 99, 490 m, 25 Jun 1969, Heckard 2319 (UC);
Shasta-Trinity National Forest, T36N R2W nw.'4 of nw. 4 S32, limestone outcrop,
13 May 1980, Williams 359 (UC). Plumas Co.: 4.8 km w. of Belden, moist cliffs, 5
Jun 1942, Heller 16500 (UC).

Previous knowledge. Known from the foothill canyons of Butte Co. and from the
canyon of the South Fork of the Yuba River near Washington, Nevada Co. [Howell,
Notes on Arabis in the Sierra Nevada, Fremontia 1(2):13-16, 1973]. Previously, the
populations appeared to be confined to volcanic or metavolcanic outcrops.

Significance. The discovery of a disjunct population in Tehama Co. prompted a
search for additional locations among specimens of Arabis breweri in the major
California herbaria. The above collections indicate that var. austinae ranges from
Ventura Co. north through the Coast Ranges to near the Oregon border, then south
into the northern Sierra Nevada. In addition, the list of substrates has increased to
include serpentine and limestone. This distribution and substrate preference parallels
that of the typical variety, although var. austinae generally occurs within the lower
range of elevations for Arabis breweri.

Arabis breweri var. austinae is currently on the watch list of the inventory of
California’s rare and endangered plants (Smith and York, CNPS Spec. Publ. No. 1,
3rd ed., 1984). This range extension indicates that var. austinae is too widespread to
be a plant of concern. In addition, the parallel distribution of this variety and the
typical variety reenforces the position that var. austinae is a weakly segregated taxon.
Although var. austinae is generally more robust and has larger leaves, flowers, and
fruiting pedicels than the typical variety, there were many intermediates among the
specimens examined, and populations of var. austinae that I have visited have a
mixture of robust and typical individuals. The basis for this intrapopulational vari-
ation is unknown. On the other hand, the size variation observed between populations
clearly has a genetic component. Progeny of robust individuals from a population of
var. austinae and of typical individuals from a population of var. breweri maintained
the characteristics of their respective parents when grown together under identical
controlled conditions (pers. obs.).

I thank curators of the cited institutions for loans or access to specimens and Rick
York for information from the CNPS rare plant file on var. austinae.— ROBERT E.
PRESTON, Department of Botany, University of California, Davis 95616. (Received
20 Oct 1986; revision accepted 9 Dec 1987.)

NOTEWORTHY COLLECTIONS

CALIFORNIA

Renewed field and herbarium work on the White Mts. of e. CA and w. NV during
recent years has revealed the following CA collections of note:

CAREX PARRYANA C. Dewey var. HALLII (Olney) Kukenthal (Cyperaceae).— Mono
Co.: Inyo Natl. For., White Mts.: subalpine meadow at Deep Springs Cow Camp,
0.65 mi [1.05 km] s. 10° w. of Station Peak, Deep Springs Valley drainage, 9490 ft
[2890 m], 13 Aug 1983, Morefield 1698 (NY); head of e. branch of s. fork of upper
Middle Cr., Fishlake Valley drainage, T2S R33E S4, 11,100 ft [3380 m], 23 Aug
1986, Taylor 8851 (RSA).

Significance. First CA reports of the species and variety, and an extension for both
ca. 190 km sw. from the nearest known site in Nye Co., NV.

DRYOPTERIS FILIX-MAS (L.) H. Schott (Aspleniaceae).— Mono Co.: Inyo Natl. For.,
White Mts., Cottonwood Cr., Fishlake Valley drainage: 1.4 mi [2.3 km] s. 70° e. of
Eva Belle Mine site in Granite Meadow, deep vertical crevices in granite, 9850 ft
[3000 m], 2 Aug 1984, Morefield 2446 (NY, RSA); 1.4 mi [2.3 km] n. 65° e. of Eva
Belle Mine site in deep granite crevices, 10,080 ft [3070 m], 10 Jul 1984, Morefield
2320 (NY); 1.5 mi [2.4 km] nnw. of Station Pk. summit at wall meadow, T5S R35E,
se.'4 of Sect. 10, 8900 ft [2710 m], 31 Jul 1987, Taylor 9189 (RSA, UC).

Previous knowledge. In CA, known only from a single collection (small scrap) in
1882 from Holcomb Valley in the San Bernardino Mts. (Parish Bros. 1513, POM).

Significance. Second known station for CA. The nearest known station of this
circumboreal species outside CA is the Flagstaff area of n.-cen. AZ, ca. 560 km to
the ese.

MALCOLMIA AFRICANA (L.) R. Brown (Brassicaceae).—Inyo Co.: Inyo Natl. For.
White Mts. in Deep Springs Valley drainage, 1.8 mi [2.9 km] due n. of Antelope
Spgs. in bed of an old mining road, 6450 ft [1970 m], 25 May 1984, Morefield 1929
(NY). San Bernardino Co.: naturalized around microwave relay station on nw. side
of Kelso Mts., ca. 23 mi [37 km] s. of Baker on Kelbaker Road, 4100 ft [1250 m],
26 May 1983, Barbe 4087 (RSA).

Significance. First CA reports of this introduced weed. Undoubtedly brought in
from NV, where it and several other introductions [such as Halogeton glomeratus
(Steph. ex Bieb.) C. A. Meyer and Cardaria pubescens (C. A. Meyer) Jarm.] are
spreading largely unchecked.

MENTZELIA REFLEXA Coville (Loasaceae).— Mono Co.: BLM land, White Mts., Ow-
ens Valley drainage at mouth of Coldwater Canyon on calcareous shale talus and
scree, T5S R33E 826, 5000 ft [1520 m], 28 May 1986, Morefield 3700 and McCarty
(BRY, NY, RSA, UC, UCR, and others, to be distributed).

Significance. First report for Mono Co., a disjunct extension ca. 100 km nnw. for
a plant otherwise endemic to the Death Valley region.

PENSTEMON BARNEBYI N. Holmgren (Scrophulariaceae).—Mono Co.: Inyo Natl.
For., White Mts., Fishlake Valley drainage, moist calcareous gravel along Busher Cr.
0.25 mi [0.4 km] w. of the CA border, T3S R35E, ne.% of nw.'4 of sw.'4 of Sect. 17,
5910 ft [1800 m], 3 May 1987, Morefield 4380 and Turner-Jones (RSA).

Previous knowledge. Segregated from the P. miser complex by N. Holmgren (Brit-

MADRONO, Vol. 35, No. 2, pp. 164-167, 1988

1988] NOTEWORTHY COLLECTIONS 165

tonia 31:226, 1979) as an endemic of e. and cen. NV, known as far w. as Esmeralda
Co. (A. Tiehm 7705, RSA).

Significance. First CA collection, and an extension 33 km wnw. from the Silver
Peak Range in NV. Also collected in the White Mts. farther e. down the same drainage
on the NV side (Morefield 4014, RSA). This species should be considered rare and
endangered in CA. It is rare in the Busher Cr. drainage, where it is reestablishing
after a scouring flash flood on 18 Jul 1984. More plants are likely present farther up
the drainage into CA. This canyon apparently is open to livestock grazing, though
no recent evidence thereof can be seen. In CA, one also should look for Penstemon
barnebyi just s. of Busher Cr. in the McAfee Cr. and adjacent drainages, where
extensive decaying carbonates provide similar habitats.

POA PATTERSONII Vasey (Poaceae).— Mono Co.: Inyo Natl. For., White Mts., Fish-
lake Valley drainage, n.-facing s. wall of cirque heading the North Fork of Perry Aiken
Cr., ca. | mi [1.6 km] ese. of White Mountain Pk., T4S R34E, nw. of Sect. 4, 12,000
ft [3660 m], 24 Jul 1987, Morefield 4695.1 and Ross (RSA, and others to be distrib-
uted).

Significance. First report for CA, an extension of at least 320 km sw. from NV and
the Rocky Mts. It was growing intimately with P. /ettermannii Vasey and P. suksdorfii
(Beal) Vasey ex Piper, with the latter of which it appears to hybridize, and for both
of which it easily can be mistaken in the field.

POTENTILLA CONCINNA J. Richardson var. DIvISA Rydberg (Rosaceae). — Mono Co.:
Inyo Natl. For., White Mts., Fishlake Valley drainage, protected granitic grus on the
ridge ne. of Tres Plumas Meadow, 1.3 mi[2.1 km]s. 55°e. of Tres Plumas benchmark
11,107, 10,400 ft [3170 m], 3 Jul 1984, Morefield 2239 (NY, RSA).

Significance. First CA report for the species and variety, and an extension for both
ca. 150 km sw. from NV.

RIBES VELUTINUM Greene var. GOODDINGII (Peck) C. L. Hitchc. (Grossulariaceae). —
Mono Co.: Inyo Natl. For., White Mts., Owens Valley drainage, steep protected
marble talus at the mouth of Pellisier Cr., T3S R33E S5, 5900 ft [1800 m], 19 Apr
1986, Morefield 3453 and McCarty (BRY, GH, MO, NY, RSA, UC, and others).
Siskiyou Co.: Klamath Mts. 0.6 mi [1.0 km] n. of Callahan, 50 yds from road in a
dry gulch, 3400 ft [1040 m], 28 Jun 1955, Barbe 018 (RSA); Lava Beds Natl. Mon.
near Fleener Chimneys, ca. 5000 ft [1520 m], 3 Sep 1969, Thorne et al. 39015 (RSA);
dry hillside along Klamath R. between Shovel Cr. and Fall Cr., ca. 2700 ft [820 ml],
15 May 1898, Applegate 2126 (RSA).

Significance. First CA reports of this taxon, extending its range ca. 600 km s. from
Malheur Co., OR.

SENECIO PATTERSONENSIS Hoover (Asteraceae).— Mono Co.: Inyo Natl. For., White
Mts., Fishlake Valley drainage, s.-facing n. wall of cirque heading the North Fork of
Perry Aiken Cr., ca. 1 mi (1.6 km) ene. of White Mountain Pk., T3S R34E, se.”
Sect. 32, 12,200 ft [3720 m], 25 Jul 1987, Morefield 4703 and Ross (RSA, and others
to be distributed).

Previous knowledge. Endemic to the Sweetwater Mts. and adjacent Sierra Nevada
(Hoover, Leafl. W. Bot. 3:256, 1943 and 5:60, 1947), and reported as rare from the
Wassuk Range of w. NV (Bell and Johnson, Madrono 27:30, 1980).

Significance. Disjunct extension ca. 120 km se. for this rare Senecio.

STYLOCLINE PSILOCARPHOIDES Peck (Asteraceae). — Representative collections: Inyo
Co.: Inyo Natl. For., White Mts., Owens Valley drainage, n. wall of canyon 4 mi [6.4
km] ese. of Laws, T6S R34E S31, 5650 ft [1720 m], 11 Apr 1986, Morefield 3389.1
and McCarty (RSA and others to be distributed); Panamint Mts.: Pleasant Canyon,
7400 ft [2260 m], 10 May 1906, Hall and Chandler 6957 (ARIZ, JEPS, POM, UC,

166 MADRONO [Vol. 35

mixed with and det. as S. micropoides A. Gray); Surprise Canyon, 1625 m, 15 Apr
1891, Coville and Funston 640 (US, det. as S. micropoides). Los Angeles Co.: San
Gabriel Mts., Mojave Desert slope, % mi [0.4 km] n. of Bob’s Gap, 3.5 mi [5.6 km]
s. of Llano, n. slope of Holcomb Ridge, ca. 3850 ft [1170 m], 25 Apr 1973, Thorne
43384 and Wallace (RSA, det. as Filago depressa A. Gray). San Bernardino Co.: Salt
Wells Valley, 0.8 mi [1.3 km] w. of CA hwy. 178, 8 air mi [13 km] e. of Ridgecrest,
T26S R41E S35, 2300 ft [700 m], 12 Apr 1974, Holmgren 7749 and Holmgren (BRY,
NY, WTU, det. as Filago arizonica A. Gray). Morefield thanks the curators of the
herbaria above for loans of material in their care.

Previous knowledge. Widespread in se. OR, sw. ID, w. and s. NV, and sw. UT.

Significance. First reports for CA, where it is frequent throughout the Mojave desert,
having passed previously for several other taxa. Use Abrams and Ferris, I//. Fl. Pacific
States IV, 1960, to distinguish these and all CA taxa of subtribe Filagininae.

TRIFOLIUM DEDECKERAE J. Gillett (Fabaceae).— Mono Co.: Inyo Natl. For., White
Mts., Fishlake Valley drainage, 1.5 mi [2.4 km] nnw. of Station Pk. summit at wall
meadow, T5S R35E, se.'4 of Sect. 10, 8900 ft [2710 m], 31 Jul 1987, Taylor 9190
(RSA, UC). Inyo Co.: Inyo Natl. For., White Mts., Deep Springs Valley drainage, 3.2
mi [5.1 km] s. 40° e. of Sage Hen Pk. just below Dead Horse Meadow, ne.-sloping
granite crevices above Crooked Cr., 7700 ft [2350 m], 26 Jun 1984, Morefield 2191
(ASC, MICH, MNA, NY, RENO, RSA, UNLV, VDB).

Previous knowledge. Known from one site in Wyman Canyon of the White Mts.,
and from 6 or 7 isolated sites in the Sierra Nevada farther s.

Significance. First record for Mono Co., an extension 14 km nnw. from Wyman
Canyon.— JAMES D. MOREFIELD, Rancho Santa Ana Botanic Garden, 1500 N. College
Ave., Claremont, CA 91711-3101; and DEAN Wm. TAyLor, Biosystems Analysis
Inc., 303 Potrero St. Suite 29-203, Santa Cruz, CA 95060. (Received 10 Jul 1987;
revision accepted 10 Nov 1987.)

NEVADA

ELATINE CALIFORNICA A. Gray (Elatinaceae).— Washoe Co., Pilgrim Lake at the
California state line on the Buckhorn Rd. from Duck Flat to Ravendale, T35N R18E
S29, ca. 1828 m, 14 Aug 1984, Tiehm and Schoolcraft 9241 (CAS, NSMC, NY).
Locally common on mud flats at the edge of the lake.

Significance. First record for Nevada. Previously known from northern Mexico n.
to WA and e. to OR, MT, and UT.

ERYNGIUM ALISMAEFOLIUM E. L. Greene (Apiaceae). — Washoe Co., center of Macy
Flat, T47N R21E S832, 1759 m, 22 Jul 1986, Schoolcraft 1660 (NY), Rye Creek
Reservoir, T46N R22E S6, 1654 m, 22 Jul 1986, Schoolcraft 1661 (UC). Growing
on seasonally inundated flats with Artemisia cana and near the edge of a reservoir.

Significance. First records for Nevada. Previously known from n. CA to OR and ID.

HACKELIA CUSICKII (Piper) A. Brand (Boraginaceae).— Washoe Co., near California
state line at s. end of the Coppersmith Hills, T37N R18E S29, 1889 m, 23 Jun 1986,
Schoolcraft 1636 (NY). Growing under Juniperus.

Significance. First record for Nevada. Previously known from Crook and Harney
cos. OR s. to Lassen and Siskiyou cos. CA.

LATHYRUS LAETIVIRENS E. L. Greene ex Rydb. (Fabaceae).— Lincoln Co., Clover
Mts., Sawmill Canyon, 4 road mi ssw. of the Ella Mt. rd. from Caliente, T5S R67E,
1828 m, 14 May 1987, Tiehm and Williams 11010 (CAS, NY, RM, RSA). Growing
with Pinus on talus slopes of rhyolitic rock.

Significance. First record for Nevada. Previously known from n. AZ, s. UT, and
w. CO.

1988] NOTEWORTHY COLLECTIONS 167

OREGON

CRYPTANTHA MICRANTHA (Torr.) I. M. Johnston (Boraginaceae).— Harney Co., Pueblo
Valley, 2.6 road mi n. of the state line on highway from Denio to Field and Burns,
then 1.6 road mi ne. along a fence line road, T41S R35E S10, 1274 m, 22 May 1987,
Tiehm 11059 (CAS, NY, ORE, OSC, RSA). Growing with Sarcobatus in areas of
sand on the valley floor.

Significance. First record for Oregon. Previously known from s. CA n. through w.
NV to Humboldt Co.

ERIOGONUM BRACHYANTHUM Coville (Polygonaceae).— Harney Co., Pueblo Valley,
0.7 road mi n. of the state line at Denio then 1.8 road mie. on a rural road, T41S
R35E 822, 1274 m, 4 Aug 1987, Tiehm 11499 (CAS, MARY, NY, ORE, OSC, RSA).
Growing with Sarcobatus on sand dunes on the valley floor.

Significance. First record for Oregon. Previously known from s. CA n. through w.
NV to Humboldt Co.— ARNOLD TIEHM, New York Botanical Garden, Bronx 10458;
and GARY SCHOOLCRAFT, BLM, 2545 Riverside, Susanville, CA 96130. (Received
27 Oct 1987; accepted 7 Dec 1987.)

REVIEWS

The Plant-Book: A Portable Dictionary of the Higher Plants. By D. J. MABBERLEY
[further subtitle — Utilising Cronquist’s An Integrated System of Classification of Flow-
ering Plants (1981) and Current Botanical Literature, Arranged Largely on the Prin-
ciples of Editions 1-6 (1896/97-—193 1) of Willis’s A Dictionary of the Flowering Plants
and Ferns], Cambridge University Press, Trumpington St., Cambridge CB2 IRP,
England, 1987, xu, 706 pp., ISBN 0-521-34060-8 (hardbound). $34.50.

Although Mabberley’s work is not specifically on Western North American botany,
it seems worthy of notice in Madro/fio because it lists almost all the taxa of the region.
The book is essentially the real seventh edition of J. C. Willis’s A Dictionary of the
Flowering Plants and Ferns (6 eds. 1897-1931). As is well known, when H. K. Airy
Shaw revised Willis’s Dictionary in 1966 and 1973 (as the 7th and 8th eds. of Willis),
to save space he dispensed with much of the general information, for example,
common names, definitions of botanical terms, and most of the accounts of economic
products and ornamental plants, and in effect turned the work into a nomenclatural
dictionary that has, of course, proven to be indispensable. In 1974 F. N. Howes’s A
Dictionary of Useful and Everyday Plants and Their Common Names appeared. This
was based on the information expurgated from the 6th, 1931 edition of Willis. Mab-
berley thoroughly updated this Willis in a comparable-size work that “‘attempts to
present all currently accepted generic and family names and commonly used English
names” of extant vascular plants. Economically important plants get very good treat-
ment, for instance, 55 lines for Eucalyptus. Most of the families get attention, although
there are likely to be omissions for some monotypic or trivial families. Many ref-
erences are included. Unfortunately, it was not feasible to include a glossary of tech-
nical terms, as did Willis (1931). Back matter includes a synopsis of Cronquist’s
classification system for angiosperms, a bibliography, and lists of abbreviations, in-
cluding an excellent 46-page list of names of authors. Overall, this is an incredibly
valuable effort and should prove to be one of the most useful books published in
recent years.—RUDOLF SCHMID, Department of Botany, University of California,
Berkeley 94720.

MADRONO, Vol. 35, No. 2, pp. 167-168, 1988

168 MADRONO [Vol. 35

Annotated Checklist of Vascular Plants of Grand Canyon National Park 1987. By
BARBARA G. PHILLIPS, ARTHUR M. PHILLIPS, II], and MARILYN ANN SCHMIDT BERNZOTT.
79 pp., soft cover. Grand Canyon Natural History Association, P.O. Box 399, Grand
Canyon, AZ 86023-0399; Monograph No. 7. 1987. $10.00 (free to researchers making
request on official letterhead).

This is an attractive and well-organized book on the botany of the Grand Canyon.
The large format (8'2 x 11 inches) and the small, but easy to read, font allow for
much information per page. The first checklist of the Canyon, by Patraw in 1932,
listed only 450 species of plants. The last checklist, by McDougall in 1947, listed
about 900. The present effort includes some 1400 species. The authors point out that
much of the Inner Canyon remains botanically unexplored and that the list will
continue to grow.

Part One includes a brief survey of the history of botany in the Canyon and sources
of information (recent field work, herbaria, publications) for the checklist, followed
by an in-depth discussion of the vegetation and climate of the North Rim, South
Rim, and Inner Canyon. This is followed by a discussion of Grand Canyon paleo-
ecology. This section includes the only figure, an interesting drawing that compares
Late Pleistocene and present-day vegetation in the vicinity of Rampart Cave.

Part Two is the annotated checklist. In preparing the list the authors used, but did
not blindly adhere to, Lehr (1978, A Catalogue of the Arizona Flora) and the sub-
sequent supplements (Lehr and Pinkava, 1980 and 1982, J. Ariz.-Nev. Acad. Sci.).
All taxa are alphabetically arranged within major groups: ferns and fern allies, gym-
nosperms, monocotyledons, and dicotyledons. Annotation information for each species
and infraspecific taxon includes “‘scientific authority” (=author of scientific name),
common name, growth form, notation if exotic or introduced, habitat, and (within
the Park) distribution, elevational range, and flowering and fruiting times. Only six
easy-to-remember abbreviations are used (CRM = Colorado River Mile, a standard
designation; L = left side of river; etc.), making the annotations very readable.

A useful appendix cross-references the nomenclature from Kearney and Peebles
(1960, Arizona Flora) to that used in the checklist. A rather generalized map of
important localities within the Park is printed on the end covers.

I find it difficult to find faults with this well-written book. My only major criticism
is that a large percentage of the species are cited as “reported from” one or more
particular localities: ““Fimbristylis thermalis Wats. Perennial herb; reported only from
foot of Bright Angel Trail, Inner Canyon. 2400 feet. Fl. & Fr. Aug. (Thornber 8237).”
Initially I was confused by the use of the phrase “reported from” in combination
with a specimen citation because neither an herbarium nor a publication is cited.
The authors do not make clear that in each of these cases they did not examine a
specimen but instead found the species, and sometimes a specimen, cited in a pub-
lished or unpublished list for the Park (A. Phillips, pers. comm.).

Another minor peculiarity of the list is that the epithets johnsoni, watsoni, eatoni,
etc. are sometimes corrected according to the International Code of Botanical No-
menclature (Art. 73.10: johnsonii, watsonii, eatonii, etc.) but often they are not. After
much searching for typographical errors I was able to find only one (p. 67, correct
spelling: Forsellesia nevadensis). In the bibliography McClintock’s single reference
(1952) occurs between McDougall 1964 and McDougall 1973. I include these cor-
rigenda for the serious user; this book’s errors are rare and do not significantly detract
from its high quality.

This checklist is indicative of the remarkable diversity of the Grand Canyon. Any
student of the plants of Arizona, particularly northern Arizona, will find Part One to
be interesting reading and Part Two a useful reference on the plants of this vast and
largely inaccessible region of the state. —BRucE D. PARFiTT, Department of Botany,
Arizona State University, Tempe 85287-1601.

Volume 35, Number 2, pages 77-168, published 9 June 1988

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CALIFORNIA BOTANICAL SOCIETY

| VOLUME 35, NUMBER 3 j JULY-SEPTEMBER 1988

‘A WEST AMERICAN JOURNAL ORS BOM AN ¥

ee
gees” ao OY a, eR “typ, .
AF %
Re sntents / SULT & 9 ;
i
| COMPOSITION OF MARITIME CHAPARRAL RELATED TO Fie History AND SOIL, BUREQN ;
MESA, SANTA BARBARA COUNTY, CALIFORNIA ~ qi Fay fa, Ric Dill
ro eet”
| Frank W. Davis, Diana E. Hickson, and Dennis C. Odion™ ameail noxinnscet 169
| INVASION OF Carpobrotus edulis AND Salix lasiolepis AFTER Fire IN A ores
CHAPARRAL SITE IN SANTA BARBARA COUNTY, CALIFORNIA
Paul H. Zedler and Gerald A. Scheid 196
_ THE VEGETATION AND ALPINE VASCULAR FLORA OF THE SAWATCH RANGE, COLORADO
| Emily L. Hartman and Mary Lou Rottman 202
| LATE WISCONSIN VEGETATION OF ROBBER’S ROOST IN THE WESTERN MOJAVE DESERT,
: CALIFORNIA
Niall McCarten and Thomas R. Van Devender 226

| THE ABUNDANCE OF PLANTS BEARING EXTRAFLORAL NECTARIES IN COLORADO AND
MOJAVE DESERT COMMUNITIES OF SOUTHERN CALIFORNIA

| Robert W. Pemberton 238
| GENERIC RELATIONSHIPS AND TAXONOMY OF Acamptopappus (COMPOSITAE: ASTEREAE)
_ Meredith A. Lane 2477
| GENECOLOGY OF Cerastium arvense AND C. beeringianum (CARYOPHYLLACEAE) IN

NORTHWEST WASHINGTON

Steven J. Wagstaff and Ronald J. Taylor 266
REVIEW 280
NOTEWORTHY COLLECTIONS
| ARIZONA 278

CALIFORNIA 279
| BAJA CALIFORNIA SUR 280
ANNOUNCEMENTS 195, 225, 237, 246, 265, 282, 283, 284

_ ERRATUM 225

| PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
ciety, Inc., and is issued from the office of the Society, Herbarium, Life Sciences
Building, University of California, Berkeley, CA 94720. Subscription rate: $30 per
calendar year. Subscription information on inside back cover. Established 1916.
Second-class postage paid at Berkeley, CA, and additional mailing offices. Return
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California Academy of Sciences, San Francisco, CA 94118.

Editor—DaAvip J. KEIL

Biological Sciences Department
California Polytechnic State University
San Luis Obispo, CA 93407

Board of Editors
Class of:

1988—SusANn G. CONARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989— FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991—JAMES HENRICKSON, California State University, Los Angeles
WAYNE R. FERREN, JR., University of California, Santa Barbara

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1988-89

President: JOHN L. STROTHER, Botany Department, University of California, Berke-
ley, CA 94720

First Vice President: JAMES AFFOLTER, Botanical Garden, University of California,
Berkeley, CA 94720

Second Vice President: JAMES HENRICKSON, California State University, Los An-
geles, CA 90032

Recording Secretary: RODNEY G. MyAatTT, Department of Biological Sciences, San
Jose State University, San Jose, CA 95192

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
Francisco, 50 Phelan Ave., San Francisco, CA 94112

The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, DALE MCNEAL, Department of Biological Sciences,
University of the Pacific, Stockton, CA 95211; the Editor of MADRONO; three elected
Council Members: JOHN MOORING, Department of Biology, University of Santa Clara,
Santa Clara, CA 95053; BARBARA ERTTER, Herbarium, Botany Department, Uni-
versity of California, Berkeley, CA 94720; ELIZABETH MCCLINTOCK, Herbarium, Bot-
any Department, University of California, Berkeley, CA 94720; and a Graduate
Student Representative, VALERIE HALEY, Department of Biological Sciences, San Jose
State University, San Jose, CA 95192.

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

COMPOSITION OF MARITIME CHAPARRAL RELATED ‘1O
FIRE HISTORY AND SOIL, BURTON MESA,
SANTA BARBARA COUNTY, CALIFORNIA

FRANK W. DAVIS, DIANA E. HICKSON,
and DENNIS C. ODION
Department of Geography,
University of California,

Santa Barbara 93106

ABSTRACT

Maritime chaparral of Burton Mesa, California was sampled to determine trends
in species richness, vegetation structure, and composition after fire. Species cover is
estimated in 75 100 m? plots distributed among 28 sites that range from 1 to 50+
years since burning. Twenty-eight plots were located under coast live oaks (Quercus
agrifolia) that are scattered among the chaparral shrubs, and 47 plots located in
surrounding chaparral. Vegetation data were analyzed and related to stand age, soil
depth, texture, and pH, and distance from the coast using Detrended Correspondence
Analysis (DCA) and Canonical Correlation Analysis (CCA).

Species richness is highest during the first 3-5 years after fire, but is more strongly
related to total evergreen shrub cover than stand age. Physiognomic trends in chap-
arral samples are similar to those reported for other chaparral types. Oak understories
differ from chaparral in that annuals decline more rapidly during the first 10 years
after fire, and subshrubs are not as important. The composition of the herb layer
under oaks is associated with stand age and distance from the coast. In chaparral
samples, composition of perennials is related to stand age, distance from the coast,
depth to a subsoil pan, soil pH, and soil texture. The composition of annuals and
biennials in chaparral is related to stand age, canopy coverage by evergreen shrubs,
depth to a subsoil pan, and distance from coast.

After fire in California chaparral, regeneration of shrubs by sprouts
and seeds produces a rapid return of the vegetation present before
the burn (e.g., Sampson 1944, Horton and Kraebel 1955, Sweeney
1956, Hanes 1971, Keeley et al. 1981; reviews in Hanes 1977 and
Vogl 1981). A diverse flora of herbs and subshrubs also flourishes
for several years after fire, declining or disappearing with closure of
the shrub canopy (Christensen and Muller 1975, Hanes 1971, J.
Keeley et al. 1985, S. Keeley et al. 1981, Schlesinger et al. 1982).
Patterns in post-fire vegetation development vary depending on
chaparral composition, fire timing and intensity, and the physical
attributes and disturbance history of the site (Keeley and Zedler
1978, Malanson and O’Leary 1985). Most studies of chaparral
succession have been located on the steep slopes and shallow soils
of the Transverse and Peninsular ranges of California. Although
general vegetational trends are known, more research is needed to

MADRONO, Vol. 35, No. 3, pp. 169-195, 1988

170 MADRONO [Vol. 35

understand how physical and biotic factors interact to control local
and regional variation in post-burn vegetation recovery.

We have studied post-fire succession in maritime chaparral on
Burton Mesa, near Lompoc, California. We chose this area for two
reasons. Since 1938, at least 27 fires have occurred in a small area
(ca. 4000 ha) of uniform climate, geology, and topography, providing
a relatively large sample for analyzing patterns of chaparral devel-
opment after fire. Moreover, there is a practical reason for analyzing
this chaparral community. Burton Mesa supports a rich chaparral
flora with many endemic taxa (Ferren et al. 1984). As in other
maritime chaparral communities, much acreage has been converted
to residential, agricultural, and military uses, and most remaining
areas are under development pressure and are experiencing invasion
by exotic weeds such as Carpobrotus edulis and Cortaderia jubata
(Griffin 1978, Jacks et al. 1984). Of the approximately 9000 hectares
of original upland habitat, we estimate that only 5890 ha of Burton
Mesa chaparral existed in 1938 and less than 3500 ha remain today.
An understanding of vegetation ecology and regeneration after fire
is needed to provide a basis for adequate protection and management
of the remainder of this threatened, endemic-rich chaparral. In this
paper we describe the modern fire history of Burton Mesa, document
the importance of subsoil morphology, distance from the coast and
stand age in determining stand composition, and compare post-fire
development with that occurring in other chaparral types.

STUDY AREA

Location and climate. Burton Mesa is located north of Lompoc
in northern Santa Barbara County (Fig. 1). The western half of
Burton Mesa is within Vandenberg Air Force Base.

The local climate is Mediterranean, having a strong maritime
influence, cool summers, and mild winters. Over 90% of the 36 cm
average annual precipitation falls between November and April.
Prevailing winds from the northwest deliver salt spray up to 50 km
into the Santa Ynez Valley (Ogden 1975). Coastal fogs, especially
prevalent during late spring and summer, greatly reduce potential
evapotranspiration. We sampled vegetation during the winter and
spring of 1985/86, which were warmer than average (Fig. 2). De-
cember and April were relatively dry, but November and March
were wet, and total precipitation was near average (Fig. 2).

Geology and soils. Burton Mesa is underlain by marine sedimen-
tary rocks and gravels which are covered with Orcutt sandstone, 0.5
to 40 m of weakly cemented Quaternary aeolian sand (Dibblee 1950,
Johnson 1983). The topography comprises level to gently rolling
uplands 100 to 120 m above sea level.

In general, topsoil on Burton Mesa is uniformly medium sand.

1988] DAVIS ET AL.: MARITIME CHAPARRAL Fal

Fic. 1. Map of study area showing approximate extent of Burton Mesa uplands
(unshaded) and documented fires occurring between 1938 and 1985 (dark shading).
Numbers designate historic fires, dots (@) represent plots in chaparral not burned
within the last 50+ years (see Table 2 for explanation).

However, soils vary in depth to bedrock, or to a clay, iron-, or silica-
indurated pan, and several soil series are recognizable based on soil
depth and subsoil properties, the most widespread being the Marina,
Tangair, and Narlon sands (Shipman 1972). The Marina series cov-
ers most of the eastern mesa and consists of deep (> 1.5 m), exces-
sively drained and infertile loamy sand. Tangair and Narlon sands
cover large portions of the central and western mesa. The Tangair
series is characterized by one to several meters of nutrient-poor light
gray sand over an impermeable or slowly draining subsoil. The sandy
layer often contains fresh iron nodules. The Narlon series is similar
to the Tangair series but distinguished by a clay subsoil and very
poor drainage. In our experience, the color, depth, and drainage
characteristics of all three series can vary considerably within Soil
Conservation Service mapping units.

Vegetation. Our analysis 1s restricted to the level uplands of Bur-
ton Mesa. The native vegetation is fragmented by roads, residential
areas, agriculture, and other developments. Vegetated areas are cov-
ered by chaparral shrubs including Adenostoma fasciculatum and
the local endemics Arctostaphylos rudis Jeps. & Wies., A. purissima
P. V. Wells, Ceanothus ramulosus var. fascicularis McMinn, and C.
impressus var. impressus. Multi-stemmed coast live oaks (Quercus
agrifolia) 3-6 m in height are interspersed throughout the chaparral,
attaining >20% crown cover in some areas not recently disturbed

172 MADRONO [Vol. 35

140

120

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80

60

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8 oes oes ote ie
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precipitation in mm.

month

Fic. 2. Climate data for Lompoc, California. Open bars are monthly average
precipitation for the period 1951-1985; shaded bars are monthly means for July 1985
to June 1986. Solid line is monthly average temperatures for the period 1951-1985,
and the dashed line for July 1985 to June 1986.

by burning or clearing. Annual grassland and coastal sage scrub
characterized by Ericameria ericoides, Artemisia californica, and
Baccharis pilularis occur on formerly cleared sites and on xeric slopes.
Some poorly drained upland sites in the central and western portions
of Burton Mesa form seasonal wetlands characterized by native
perennial grasses such as Elymus glaucus and vernal pool species
including Eryngium armatum.

The vegetation of Burton Mesa has received little systematic study.
Wells (1962) and Cole (1980) have described the strong association
of plant species and geology in the region. Ferren et al. (1984) provide
a thorough description and floristic analysis of vegetation of La
Purisima Mission State Historic Park, at the eastern end of Burton
Mesa. Halligan (1973) studied stands of Artemisia californica in the
park to determine the factors suppressing understory growth in coast-
al sage scrub, and Ogden (1975) investigated the role of salt aerosols
in limiting the local distributions of Quercus lobata and Quercus
agrifolia.

1988] DAVIS ET AL.: MARITIME CHAPARRAL sess)

METHODS

Fire history. We reconstructed the fire history of Burton Mesa
since 1938 from aerial photography that provided coverage at 5- to
10-year intervals. Past issues of the Lompoc Record were studied
for references to the date, size, location, and cause of fires. In ad-
dition, Santa Barbara County records were checked, and fire de-
partment personnel at Vandenberg Air Force Base were interviewed.
Burn scars were located on air photos and mapped onto USGS 7',
minute topographic maps using a Bausch and Lomb Zoom Transfer
Scope. Burned areas in the chaparral were detectable on air photos
for up to 10 years after the fire, allowing almost continuous coverage
for the 50-year time period.

Vegetation and soil sampling. To study vegetation development
after fire, we sampled level to moderately sloping uplands that had
not been grazed or cleared since at least 1945, locating plots away
from roads or trails to avoid edge effects. We sampled 47 stands on
33 sites that spanned nearly the entire length of Burton Mesa and
ranged in age from 1 to 50+ years. The Braun-Blanquet relevé
approach (Mueller-Dombois and Ellenberg 1974) was used to char-
acterize vegetation of a burn. Each burn area was first reconnoitered,
and sample plots were subjectively located in large distinctive stands
that best represented the post-fire vegetation. Time constraints lim-
ited the number of plots to a maximum of four for the most het-
erogeneous burn sites. Because of limited historical aerial photog-
raphy for Vandenberg Air Force Base, and also because of apparently
lower fire frequency on western Burton Mesa, we sampled the central
and eastern portions of Burton Mesa more intensively than western
Burton Mesa. We subsequently have discovered that much of the
western mesa was cultivated prior to 1938.

The vegetation at most sites included chaparral with scattered
coast live oaks. We sampled oak understories and chaparral sepa-
rately to compare post-fire vegetation development in the two mi-
croenvironments. Depending on the size of the burn and the vari-
ation in vegetation, we placed one to three sample plots in chaparral
(hereafter referred to as “‘chaparral plots’’), and one under oak can-
opy (“‘oak plots’’), for a total of 47 chaparral plots and 28 oak plots.

Chaparral plots were circular and 100 m7, an adequate size based
on species-area curves at four sites from 1 to 50+ years. The shape
and size of oak plots varied among individual oak canopies, which
were sampled in their entirety and which ranged in size from roughly
50 to 150 m?. We sampled stands by visually estimating cover (Braun-
Blanquet cover classes) of all vascular plant species in the plots. The
same observers, at least two and usually three of us, estimated species
cover for all samples, helping to reduce effects of observer bias on
species cover estimates (Gotfryd and Hansell 1985). We visited all

174 MADRONO [Vol. 35

plots in 1986 in early, mid-, and late spring (22 March through 20
May), and in August, as needed, so that their entire flora and the
maximum cover for each species could be observed.

Oak canopy cover in a stand was measured photogrammetrically
by centering a 10 x 10 grid on the stand in 1:24,000 1983 aerial
photographs. The grid corresponded to 120 x 120 m on the ground.
The fraction of each 12 x 12 m grid cell covered by oak canopy
was visually estimated and values summed to obtain the percent
oak cover for the stand.

For each chaparral plot we used a 2.5 cm diameter soil probe to
observe soil texture, color, and stratigraphy and to measure the depth
(up to 1.8 m) to a sub-surface pan or to bedrock. For all plots, soil
samples composited from the top 20-30 cm of topsoil were dry-
sieved in the lab to determine particle size distribution. Soil pH in
both water and a KCI solution was measured. Values reported here
are for measurements made in water.

Nomenclature follows Munz (1959, 1968) except where author is
noted. Voucher specimens are deposited at UCSB and SBBG.

Data analysis. We analyzed species cover data using Detrended
Correspondence Analysis (DCA) (Hill and Gauch 1980), canonical
correlation analysis, and direct ordination. DCA is an indirect or-
dination method for representing the relative similarity of samples
or species along a few principal axes of variation, which can be
studied for their relationship to environmental variables. The meth-
od is especially useful for detecting linear or unimodal gaussian
relationships between species and environmental gradients. Thus
the method works best for sampling vegetation along one to several
environmental gradients within a narrow range of environmental
variation over which species responses may exhibit single optima
(Austin 1976, Noy-Meir and Whittaker 1977). We restricted our
sampling to chaparral on level uplands of a single geologic substrate
in an effort to produce data that would be amenable to indirect
ordination analysis. We performed separate ordinations for four
species groups, because we found that the stratification produced
clearer relationships between ordination scores and environmental
variables. These groups included annual and biennial species in
chaparral plots, perennial species in chaparral plots, annual and
biennial species in oak plots, and perennial species in oak plots.

Ordination results can be sensitive to data quality, data transfor-
mations, weighting of rare species, and standardization and nor-
malization procedures (Noy-Meir et al. 1975). We tested the stability
of DCA ordinations obtained using species presence-absence versus
cover class data, and those with and without downweighting of rare
species. Downweighting in the DECORANA program was accom-
plished by reducing species abundance values in proportion to their

1988] DAVIS ET AL.: MARITIME CHAPARRAL 175

frequencies of occurrence, for species with frequencies less than 20%
of the most frequent species (Hill 1979). The ordinations were rel-
atively insensitive to the weighting of rare species. However, ordi-
nation scores were sensitive to data quality, with poor agreement
between results obtained for presence-absence data versus cover
class data. The ordinations based on perennial species cover pro-
vided a better reconstruction of plot similarity than the presence-
absence ordinations. This is expected, given the importance of dom-
inant canopy species in chaparral stands. On the other hand, the
ordinations based on the presence or absence of annual species gave
more interpretable axes than those based on cover data. Annual
species rarely exceeded 1-—5% ground cover in a plot; their cover
varied over the season and was typically patchier than perennial
cover. The cover estimates for annuals were thus not as meaningful
nor reliable as those for perennials, and species presence was a more
appropriate measure. Here we present plot ordination results based
on perennial cover and on annual species presence-absence data,
with no downweighting of rare species.

Variables analyzed for their correlation with the first and second
DCA axes included years since burning, depth of soil over a clay or
iron pan or over bedrock, distance from the coast along the pre-
dominant wind direction (northwest), soil pH, and percent fine frac-
tion (<0.1 mm) in the upper 30 cm of the soil. Also, correlation
was measured between sample ordination scores based on herb species
composition and total canopy cover of evergreen shrubs.

Some environmental parameters are highly correlated, making it
difficult to interpret correlations between ordination axes and in-
dividual environmental variables. Soil depth and soil fine fraction
are both significantly negatively correlated with distance from the
coast (Table 1). Soil fine fraction is positively correlated with soil
pH, and evergreen shrub canopy cover is positively associated with
stand age. To help account for the interrelationships between en-
vironmental variables, we performed canonical correlation analyses
using the first and second axis scores for plot ordinations and selected
environmental factors (Dillon and Goldstein 1984). Environmental
variables were first standardized to mean O and standard deviation
of 1. Log transforms of plot age and distance from the coast were
used because the transformation increased the linear relationship
between these variables and the ordination axes.

Canonical correlation analysis is a generalization of multiple
regression analysis that identifies canonical axes that maximize the
correlation between two groups of descriptors, in this case DCA axis
scores and environmental factors. The technique is applicable only
when descriptors are linearly related, limiting its use in ecological
analyses (Legendre and Legendre 1983). Some alternative techniques
have been developed recently, such as canonical correspondence

~

MADRONO

[Vol. 35

176

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1988] DAVIS ET AL.: MARITIME CHAPARRAL eg

TABLE 2. DATES AND SAMPLE ALLOCATION FOR FIRES OCCURRING ON THE BURTON
MESA BETWEEN 1938 AND 1985. Fire numbers correspond to areas in Fig. 1.

Fire Number of plots Year of fire
1 l 1984
2 2 1983
3 2 1981
4 3 1981
Ss pp 1985
6 0 1983
a 4 1982
8 2 1983
9 3 1982

10 l 1976
11 0 ca. 1973
2 4 1967
13 4 1976
14 2 1967
15 4 1962
16 3 1976
17 5 1985
18 1 1972
19 2 1974

20 0) ca. 1973

21 0) ca. 1962

22, 0) ca. 1960

23 3 1974

24 4 1974

25 6 1961

26 0 1971

Deh 0) ca. 1970

analysis (Ter Braak 1987), which use constrained ordination to en-
sure linear relationships between ordination axes and environmental
factors. We tested several constrained ordination methods, including
detrended canonical correspondence analysis and redundancy anal-
ysis (Ter Braak 1987), and obtained results similar to those obtained
using canonical correlation analysis. We present the latter results
here because canonical correlation analysis is a more widely known
method.

RESULTS

Fire history analysis. We located 27 fires larger than 1 ha that
occurred between 1938 and 1986 (Fig. 1, Table 2). Several other
large fires occurred on VAFB during the period, but these could not
be accurately mapped due to inaccessible or incomplete photo cov-
erage and records for the base. Most fires spread in the direction of
prevailing northwesterly winds and were extinguished at or arrested
by roads, fields, or fuel breaks. All 27 fires were started by humans;

178 MADRONO [Vol. 35

the eight fires on Vandenberg AFB were controlled burns, whereas
the others were either accidental or arson fires. The Lompoc Record
reported only one possibly lightning-caused fire in the region during
this period. That fire originated on Tranquillon Mountain, a 650 m
peak 15 km s. of Burton Mesa, but did not reach the mesa.

Flora and physiognomy. We encountered 41 families, 110 genera
and 152 species of vascular plants in the sample plots (Appendix
1). The largest families include Asteraceae (36 species), Poaceae (16),
and Scrophulariaceae (8). Two species, Arctostaphylos rudis and A.
purissima, are endemic to the Burton Mesa; five others are endemic
subspecies or varieties, including Ceanothus ramulosus var. fasci-
cularis, C. impressus var. impressus, Mimulus aurantiacus subsp.
lompocensis, Amsinckia spectabilis var. microcarpa, and Erysimum
suffrutescens var. lompocense. Thirty species are exotic.

The flora includes one tree, 17 shrubs, 17 subshrubs, 31 perennial
herbs, and 86 annual or biennial species. The most frequent shrubs
are Adenostoma fasciculatum and Ceanothus ramulosus var. fasci-
cularis, whereas the most frequent subshrubs are Horkelia cuneata
and Lotus scoparius. Melica imperfecta is the most frequent peren-
nial herb, and Camissonia micrantha and Vulpia octoflora the most
frequent annuals. Of the annuals and biennials 39 of 86 species (45%)
occur in three or fewer plots, and 18 species (21%) occur in only one
plot. Twelve of 31 perennial herb species (39%) occur in three or
fewer plots, and, of these, 10 (32%) occur only once. In contrast,
only one of 35 (3%) woody perennial species (Solanum douglasii)
occurs in fewer than three plots.

In general, species richness declines during the first 15-20 years
after burning, but there is considerable variation in richness between
plots of the same age (Fig. 3). Recently-burned chaparral plots have
consistently high richness; S = 36 species for 14 plots <10 years of
age. Species richness averages 20 in 35 plots older than 20 years,
and does not differ significantly between plots 20-30 years of age
and those at least 50 years old. Species richness is generally lower
in oak plots, and for the first 25 years after fire, species richness
appears to decline more rapidly (p < 0.10) in oak plots than in
chaparral plots. The slope of the regression line relating species
richness to plot age is —0.93 for chaparral (n = 38, s.d. = 0.17) and
— 1.26 under oak canopies (n = 26, s.d. = 0.17). When all chaparral
plots are considered, species richness is negatively correlated with
total evergreen shrub cover (r = —0.76, p < 0.01), less so with years
since burning (r = —0.56, p < 0.01) or the logarithm of years since
burning (r = —0.65, p < 0.01). We tested for possible effects of other
environmental variables on species richness by including them with
plot age and shrub cover in a stepwise multiple regression analysis.
Neither soil variables nor distance from the coast were significantly

1988] DAVIS ET AL.: MARITIME CHAPARRAL 179

i¢p)
dp)
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i

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—

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Y) * *

O P|

8 ‘ :

@) *

O O O

® O 8

O x

x

O

| | a] | Elbe

0 10 20 30 40 50 60
Age

Fic. 3. Species richness versus years since fire for plots in chaparral (*) and oak
understories (©).

correlated with species richness, and none of them significantly 1m-
proved regression model r? over the model based solely on total
shrub cover.

The decline in species richness in chaparral plots is due to pro-
portional declines in number of species of all growth forms, whereas
under oaks it is due mainly to a reduction in the number of annual
species (Table 3). The decline in understory richness under oaks,
unlike that in the chaparral plots, is not accompanied by a decline
in understory cover. The understories of oak plots in the older age
classes varied from a cover of 25-50% Toxicodendron diversilobum
to less than 5% cover of one to several perennial species. Trees with
low understory richness and cover were frequently associated with
nests of the wood rat (Neotoma sp.).

For chaparral plots, physiognomic trends during recovery from
fire are similar to those described for other chaparral communities
(Table 3). Herbs and subshrubs account for most of the cover in
chaparral plots less than 5 years old. Dominant herb species include
Vulpia octoflora, Chorizanthe diffusa, Camissonia micrantha, Cras-

[Vol. 35

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MADRONO

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1988] DAVIS ET AL.: MARITIME CHAPARRAL 181

sula erecta, Filago californica, and Cryptantha clevelandii. Cover of
subshrubs such as Helianthemum scoparium, Lotus scoparius, and
Eriophyllum confertiflorum peaks 3-5 years after fire, after which
shrub cover dominates. Total cover is not as high in the oldest plots
as in the 18-25 year range, due mainly to Ceanothus ramulosus
mortality. Dead individuals of this species are common in chaparral
over 20 years old on deep soils.

Coast live oaks attain highest density and canopy cover on sites
with no recent fires, particularly on deep sands in the eastern portion
of the mesa. Oak canopy cover is significantly related to time since
last fire (r = 0.67, n = 49, p < 0.001), although cover ranges widely
on sites of similar age depending on microtopography, soil, coastal
influence, and fire intensity.

The physiognomic trend in plant cover under oaks differs from
that under the chaparral shrubs, in that annuals continue to dominate
plant cover for 3-5 years after fire, and subshrubs like Helianthemum
scoparium and Lotus scoparius are never a major component of the
understory (Table 3).

Vegetation ordinations. The vegetation under oak canopies differs
considerably from that in chaparral plots, as shown by relative fre-
quencies of many species in oak versus chaparral plots (Appendix
1), and based on ordination scores for oak versus chaparral plots in
a sample ordination including both plot types (Fig. 4). Characteristic
species under oak canopies include the perennials, Toxicodendron
diversilobum, Baccharis pilularis, Galium nuttallii, and Marah fa-
baceus (Appendix 1). The only annual that occurs with much greater
frequency under oaks is Claytonia perfoliata. On the other hand,
many annuals occur exclusively or preferentially in chaparral, for
example, Calyptridium monandrum, Camissonia strigulosa, Linaria
canadensis var. texana, and Vulpia myuros (Appendix 1).

The first DCA axis for the ordination based on annuals and bien-
nials in oak understory plots is significantly related to the logarithm
of time since burning (r = 0.45, n = 21, p < 0.01, Table 4). For
perennial species under oaks, the first DCA axis is related to the
logarithm of distance from the coast (r = —0.51, n = 28, p < 0.05),
and plot age is not significantly associated with either DCA axis. We
do not know whether distance from the coast or soil depth is the
more important factor associated with perennial composition in oak
understories.

The first DCA axis for the chaparral plot ordination based on
woody perennial species is related to percent evergreen shrub cover
(r = —0.67, p < 0.001), stand age (r = —0.43, p < 0.01), and soil
pH (r = 0.28, p < 0.05) (Table 4). Vegetation changes are most
rapid during the years immediately following fire, so that an im-
proved linear relationship with DCA axis | is obtained by taking

182 MADRONO [Vol. 35

DCA axis 2

0 100 200 300 400 500

DCA axis 1

Fic. 4. DCA first versus second axis scores for plot ordination based on both chap-
arral (*) and oak plots (O).

the logarithm of stand age (r = —0.53). Two 25-year plots are severe
outliers, one with unusually high cover of Horkelia cuneata and the
other with low shrub diversity and dominance by Ceanothus ra-
mulosus (although these plots had not been disturbed since last
burning, they were mechanically cleared 40-50 years ago). Elimi-
nation of these outliers increases correlation of DCA axis | with the
log of stand age to —0.69. The second DCA axis for woody perennials
is significantly related to soil depth (r = 0.63, p < 0.001), the log-
arithm of distance from the coast (r = 0.61, p < 0.001), and soil
texture (r = —0.39, p < 0.01). The joint effects of these environ-
mental variables are displayed by plotting their loadings on two
canonical correlation axes, which have canonical correlations of 0.65
and 0.63 (Fig. 5). Plot age has the highest loading on axis 1, whereas
plot depth has the highest loading on the second canonical axes.
Distance and soil texture have moderate loadings; soil pH has the
lowest score.

We have observed a strong association of soil depth and vegetation
composition, with distance from the coast held constant, at some

DAVIS ET AL.: MARITIME CHAPARRAL 183

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184 MADRONO [Vol. 35

1.0

-1.0

Fic. 5. Loadings of environmental variables on canonical correlation axes ex-
tracted jointly for environmental variables and chaparral plot scores in the first two
DCA ordination axes. Environmental variables are standardized to N(0, 1). The length
of the arrow is proportional to the weighting of the variable in the canonical correlation
function. a) Plot scores based on perennial species ordination. Canonical correlations
are 0.65 and 0.63 for axes | and 2. DCA axis | loadings: —0.48, —0.71; DCA axis
2 loadings: 0.88, —0.69. Environmental variables are log(years since fire) (AGE),
log(distance from coast) (DIS), soil depth (DEP), percent fine fraction (FIN), and pH.
b) Plot scores based on ordination of annual and biennial species. Canonical corre-
lations are 0.77 and 0.58 for axes | and 2. DCA axis | loadings: 0.89, —0.16; DCA
axis 2 loadings: 0.44, 0.99. CAN is total canopy cover by evergreen shrubs.

sites where soil grades from deep sand to shallow sand over a clay
pan or bedrock. In older stands, we observed a gradient from Ad-
enostoma-dominated to Arctostaphylos-dominated chaparral when
moving from deep to shallow sand. On recently-burned sites, one
can observe a decline in Lotus scoparius and an increase in Helian-
themum scoparium on increasingly shallow soil.

We were unable to sample 10-25 year stands on very shallow
soils, and also could not locate any 25—30-year-old stands. Never-

1988] DAVIS ET AL.: MARITIME CHAPARRAL 185

theless, the data indicate some strong patterns in the distribution of
the dominant shrubs of Burton Mesa chaparral (Fig. 6). Canopy
cover of Ceanothus ramulosus var. fascicularis peaked in stands
between 10 and 25 years of age on deep sand. We encountered C.
impressus only in plots that burned in the past 10 years on shallow
soils. The highest cover values for Arctostaphylos purissima occurred
in stands greater than 50 years of age on very shallow soil. Arc-
tostaphylos rudis reached maximum cover in the oldest plots, but
was not strongly associated with soil depth.

Chaparral plot ordinations based on annual species show many
of the same patterns as those based on perennials, although the
relationships are not as strong (Table 4). The first DCA axis is
significantly related to stand age (r = 0.43, p < 0.01) and its logarithm
(r = 0.52, p < 0.01), but is more clearly related to total cover of
evergreen sclerophyllous shrubs (r = —0.67, p < 0.001). The second
DCA axis based on annual species is significantly related to soil
depth (r = —0.55, p < 0.01) and to the log of distance from the
coast (r = —0.42, p < 0.05). Other soil variables are not strong
predictors of vegetation composition. Evergreen shrub canopy cover
and depth have the highest loadings in the canonical correlation
axes, which have canonical correlations of 0.77 and 0.58 (Fig. 5).
As with the canonical correlation analysis for perennial species, the
relative importance of the different environmental variables is not
especially sensitive to variable transformations.

Most annuals and biennials did not occur in very many plots and
thus we cannot describe their distributions with much certainty.
Also, micro-scale pattern in distribution of annual herbs, particularly
in relation to shrub canopies and canopy gaps, makes it difficult to
draw strong conclusions about their habitat associations based on
their occurrence in 100 m? sample plots. Analysis of direct ordi-
nations (not shown) indicates that some of the most frequent species
are nearly ubiquitous, and their presence is not associated with plot
age, soil depth or distance from the coast. These include Camissonia
micrantha, Chorizanthe diffusa var. nivea, Crassula erecta, and Vul-
pia octoflora. Frequent species characteristic of younger, more open
stands include Hypochoeris glabra, which occurs on all soils, and
Daucus pusillus and Pterostegia drymarioides, which occur on shal-
low and deep soils, respectively. Perezia microcephala is a perennial
herb species characteristic of older closed stands, and it occurs ex-
clusively on deep, well-drained soils.

DISCUSSION

Fire regime. The present fire regime of the Burton Mesa is entirely
anthropogenic. Modern land use is increasingly fragmenting the re-
maining chaparral into isolated patches, reducing potential fire size
and probably increasing the average time between fires on a site.

186 MADRONO [Vol. 35

100 100

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Fic. 6. Three-dimensional graphs of canopy cover versus stand age and soil depth
for the 5 dominant canopy shrubs in Burton Mesa chaparral (n = 47): a) Ceanothus
ramulosus, b) C. impressus, c) Adenostoma fasciculatum, da) Arctostaphylos rudis, e)
A. purissima. Cover values shown on the Z axis are midpoints of Braun-Blanquet
cover classes. Shaded squares indicate species absence in a sample. Numbers along

1988] DAVIS ET AL.: MARITIME CHAPARRAL 187

With a long fire-recurrence interval coast live oak should increase
on all but the driest sites in sandy coastal chaparral environments
(Wells 1962, Griffin 1978). Our data support this interpretation for
the central and eastern mesa. However, on very poorly drained soils
and sites near the coast, oak cover is low even in the oldest plots
sampled, implying that these areas persist as chaparral for long time
periods.

Flora. Ferren et al. (1984) collected 342 taxa (252 native, 90
naturalized) from the range of upland and wetland habitats present
in the eastern Burton Mesa. Our list of 152 species is not compre-
hensive but includes nearly all species occurring on unplowed and
ungrazed uplands of the Burton Mesa. Numerous other species doc-
umented on the Burton Mesa, such as the common Hordeum le-
porinum, Cnicus benedictus, Sisyrinchium bellum, and Juncus bu-
fonius, typically occur on previously disturbed sites, where the present
vegetation is mostly introduced grassland or coastal scrub species.

The presence of 152 species within a total sample area of only
0.79 ha of level uplands is indicative of moderately high total rich-
ness of the Burton Mesa chaparral flora compared to other shrub-
lands in Mediterranean climates (cf. Parsons and Moldenke 1975,
Trabaud and Lepart 1980, Shmida 1981, Cowling 1983). Species
richness is increased somewhat by addition of exotic species, but
may be more related to the sandy oligotrophic substrate, which has
been associated with exceptional richness in other coastal chaparral
communities (Griffin 1978) and in similar areas of the Mediterra-
nean region (Naveh and Whittaker 1979), Australia (George et al.
1979), and South Africa (Cowling 1983). High richness has also been
attributed to the location of the Burton Mesa in a transitional region
combining northern and southern California floras (Ferren et al.
1984).

Spatial and temporal variation in species richness for 100 m7? plots
makes it difficult to compare richness at this scale to other values
reported in the literature. We note that average species richness on
the Burton Mesa is close to the 29 spp./100 m? reported by Shmida
and Whittaker (1981) for other California chaparral and is lower
than in Mediterranean shrublands recognized for their high richness,
such as South Africa renosteveld, a coastal shrubland on sandy soils
[42.1 spp./100 m? (Cowling 1983)], and heathland on western Aus-
tralian sand plains [46 spp./100 m? (George et al. 1979)].

Factors that have been analyzed to explain variation in species

—
the top margin of the figure base in figure b) are the number of replicates for that

combination of stand age and soil depth >180 cm (bars are sample average for
replicates).

188 MADRONO [Vol. 35

richness in Mediterranean shrublands include climate, topography,
soil texture, soil fertility, soil pH, time since burning, and grazing
intensity. Our analysis is somewhat different in that samples are not
recently cleared or grazed and share the same climate, topography,
and geologic substrate. Under these conditions, time since burning
is a significant but weak predictor of species richness in maritime
chaparral. Radtke (1981) obtained a similar result for other chaparral
types. Also, soil depth, texture, and pH are not significantly related
to sample richness. However, total evergreen shrub cover is a strong
predictor, accounting for 58% of total sample variance, perhaps due
to differences in levels of allelopathic chemicals or herbivory (e.g.,
McPherson and Muller 1969, Halligan 1973, Christensen and Muller
1975, Davis and Mooney 1985).

Vegetation ordinations. Although we sampled a narrow range of
topographic conditions on a single geologic substrate, there are im-
portant differences between samples in soil depth and distance from
the coast, so that they cannot be arrayed as a simple chronosequence
to reconstruct successional patterns of maritime chaparral after fire.
For oak understory vegetation, we have identified one or two major
axes of variation, time since burning and distance from the coast,
and for chaparral plots we have identified two or three major axes
of variation, stand age or shrub canopy cover, soil depth, and dis-
tance from the coast, that must be considered jointly in relation to
vegetation composition. More replicates of the same age and en-
vironmental conditions are needed to document conclusively the
vegetation dynamics in these different environments.

Chaparral composition has been associated with geologic substrate
and soil properties such as fertility, texture, and chemistry (e.g., Wells
1962, O’Leary 1984, Beatty 1987). The edaphic variable most af-
fecting chaparral composition on Burton Mesa is the depth of sand
overlying bedrock or a subsoil pan. There are probably large differ-
ences in soil water regime between shallow and deep sands. We
expect that shallow soils have low water holding capacity, perhaps
remaining waterlogged during the winter and early spring, and then
losing most of their water by late spring or early summer. The high
water holding capacity of deep soils means that waterlogging does
not occur, but that soil water remains available at depth in the profile
later in the year (Nixon and Lawless 1960). Differences in soil fertility
would also be expected due to differences in the extent of leaching.
Soil fertility was not measured here, except indirectly by pH deter-
mination, and deserves more careful study (Christensen and Muller
1975, Parker 1977, O’Leary 1984).

Soil variation may partly explain the differentiation of the endemic
congeners, Ceanothus impressus var. impressus versus C. ramulosus
var. fascicularis, and Arctostaphylos purissima versus A. rudis. The

1988] DAVIS ET AL.: MARITIME CHAPARRAL 189

importance of edaphic variation in promoting narrow endemics was
noted by Mason (1946). Wells (1969) has described rapid speciation
by obligate seeding Ceanothus and Arctostaphylos on different sub-
strates.

The association of many herb species with either chaparral or oak
understories produces abrupt changes in herb layer composition at
the edge of oak canopies, a pattern that is evident on all sites where
oaks occur and at all stages of vegetation development after fire. We
observed but did not measure additional within-stand pattern in the
chaparral herb layer at the scale of canopy gaps and shrub under-
stories (e.g., Shmida and Whittaker 1981). We are now analyzing
local and microscale patterns in fire behavior, soil chemistry, and
seed banks of chaparral and oak understories to help understand the
origin of this microscale floristic pattern in maritime chaparral.

ACKNOWLEDGMENTS

We thank Joe McCummins and the other members of the La Purisima Mission
State Historic Park staff, the Environmental Task Force Office of Vandenberg Air
Force Base, and Unocal. We also extend our appreciation to Dean Capralis, Laura
Haston, and Robert Paul for assistance in the field; to Clifton Smith, Wayne Ferren,
Steve Junak, Holly Forbes, and Dale Smith for identifying many of the taxa; and to
Nicholas Graham, Judy Paddon, and Dave Lawson for their assistance in data han-
dling and graphics. Waldo Tobler provided software for 3-dimensional plots. Three
anonymous reviewers provided very helpful criticisms of the draft manuscript. This
study was funded in part by the California Parks and Recreation Department, contract
4252-609.

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1988] DAVIS ET AL.: MARITIME CHAPARRAL 11

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192 MADRONO [Vol. 35

APPENDIX |. SPECIES LIST FROM SAMPLE PLots. *Vulpia megalura and V. myuros
were treated as a single entity by Lonard and Gould (1974). We treated them as
separate taxa because they were readily distinguishable in the field, and there was
some evidence of ecological differentiation between them.

Frequency
Shrub Oak
Species Family plots plots
Trees and shrubs
Adenostoma fasciculatum Rosaceae 87 39
Arctostaphylos purissima P. V. Wells Ericaceae 47 14
Arctostaphylos rudis Ericaceae 55 29
Artemisia californica Asteraceae Gl 4
Baccharis pilularis subsp. consanguinea Asteraceae 38 54
Ceanothus impressus var. impressus Rhamnaceae 21 11
Ceanothus ramulosus var. fascicularis Rhamnaceae 72 36
Cercocarpus betuloides Rosaceae 6 4
Dendromecon rigida Papaveraceae 11 7
Ericameria ericoides (Less.) Jeps.

subsp. ericoides Asteraceae 51 29
Leptodactylon californicum Polemoniaceae 19 a
Mimulus aurantiacus subsp. lompocensis Scrophulariaceae 60 54
Quercus agrifolia Fagaceae 13 100
Rhamuus californica Rhamnaceae 21 29
Rhamnus crocea Rhamnaceae 21 4
Salvia mellifera Lamiaceae 57 14
Sambucus mexicana Caprifoliaceae 0 18

Toxicodendron diversilobum (Torr. &
A. Gray) Greene Anacardiaceae 36 75

Subshrubs
Carpobrotus edulis (L.) L. Bolus Aizoaceae 15 11
Corethrogyne filaginifolia Asteraceae 32 14
Croton californicus var. californicus Euphorbiaceae 40 11
Eriastrum densifolium subsp. elongatum Polemoniaceae 23 4
Eriogonum parvifolium Polygonaceae 9 0
Eriophyllum confertiflorum Asteraceae 25) 50
Erysimum suffrutescens var. lompocense Brassicaceae 9 7
Galium andrewsii Rubiaceae 42 54
Galium nuttallii Rubiaceae 62 79
Helianthemum scoparium Cistaceae 66 21
Horkelia cuneata Rosaceae 68 32
Lotus scoparius (and L. junceus) Fabaceae 64 39
Phacelia ramosissima var. suffrutescens Hydrophyllaceae 11 11
Senecio douglasii Asteraceae 11 11
Solanum douglasii Solanaceae 0 11
Solanum umbelliferum Solanaceae 13 21
Solanum xanti Solanaceae 2 11
Perennial herbs

Bloomeria crocea Amaryllidaceae 2 0
Calochortus albus Liliaceae 2 0
Calystegia macrostegia subsp. cyclostegia Convolvulaceae 2 4
Carex globosa Cyperaceae 30 21
Carex triquetra Cyperaceae 6 0

1988]

DAVIS ET AL.: MARITIME CHAPARRAL
APPENDIX 1. CONTINUED.

Species Family
Chenopodium californicum Chenopodiaceae
Chlorogalum pomeridianum Liliaceae
Conicosia pugioniformis (L.) N. E. Br. Aizoaceae
Cortaderia jubata (Lem.) Stapf. Poaceae
Cuscuta californica Convolvulaceae
Dichelostemma pulchellum (Salisb.) Heller Amaryllidaceae
Dudleya lanceolata Crassulaceae
Erigeron sanctarum Asteraceae
Fritillaria biflora Liliaceae
Gnaphalium beneolens Asteraceae
Gnaphalium bicolor Asteraceae
Juncus textilis Juncaceae
Marah fabaceus var. agrestis Cucurbitaceae
Melica imperfecta Poaceae
Paeonia californica Paeoniaceae
Pedicularis densiflora Scrophulariaceae
Penstemon centranthifolius Scrophulariaceae
Perezia microcephala Asteraceae
Pityrogramma triangularis Pteridaceae
Pteridium aquilinum var. pubescens Pteridaceae
Rumex angiocarpus Polygonaceae
Sanicula bipinnatifida Apiaceae
Sanicula crassicaulus Apiaceae
Silene laciniata subsp. major Caryophyllaceae
Stipa cernua Poaceae
Zigadenus fremontii Liliaceae

Annual and biennial herbs

Amsinckia spectabilis var. microcarpa

Anagallis arvensis

Apiastrum angustifolium

Arenaria douglasii

Avena barbata

Bromus diandrus

Bromus mollis

Bromus rubens

Calandrinia breweri

Calyptridium monandrum

Camissonia micrantha

Camissonia strigulosa

Centaurea melitensis

Centaurium davyi

Chorizanthe californica

Chorizanthe coriacea

Chorizanthe diffusa Benth. var. nivea
(Curran) Hoover

Cirsium californicum

Cirsium occidentale var. occidentale

Claytonia perfoliata (Donn) Howell

Conium maculatum

Boraginaceae
Primulaceae
Apiaceae
Caryophyllaceae
Poaceae
Poaceae
Poaceae
Poaceae
Portulacaceae
Portulacaceae
Onagraceae
Onagraceae
Asteraceae
Gentianaceae
Polygonaceae
Polygonaceae

Polygonaceae
Asteraceae
Asteraceae
Portulacaceae
Apiaceae

193
Frequency
Shrub Oak
plots plots
0 7
6 0
2 4
0 4
2 0
23 0)
19 0
34 28
4 0)
17 4
11 0
0 4
23 ful
32 29
19 18
4 0
4 0
19 11
2 0
13 14
Z 0
Z 4
0 4
17 14
2) 0
15 29
ys 4
28 18
17 21
2 0)
4 7
6 11
9 0
43 32
2 4
15 4
68 32
19 4
0 4
pe. 0)
15 4
y 4
74 25
2 4
13 14
15 54
0 4

194 MADRONO [Vol. 35

APPENDIX 1. CONTINUED.

Frequency

Shrub Oak

Species Family plots plots
Conyza canadensis Asteraceae 17 25

Cordylanthus rigidus (Benth.) Jeps.

subsp. /ittoralis Chuang & Heckard Scrophulariaceae 4 0
Crassula erecta (H. & A.) Berger Crassulaceae 70 Zt
Cryptantha clevelandii Boraginaceae 53 43
Daucus pusillus Apiaceae 32 36
Descurainia pinnata ssp. menziesii Brassicaceae 11 21
Eriophyllum multicaule Asteraceae 4 0
Erodium botrys Geraniaceae 4 0
Erodium cicutarium Geraniaceae 21 11
Filago californica Asteraceae 70 18
Filago gallica Asteraceae 40 18
Galium aparine Rubiaceae 2 7
Gastridium ventricosum Poaceae 0 4
Gilia austrooccidentalis Polemoniaceae 30 7
Gnaphalium californicum Asteraceae 6 11
Gnaphalium luteo-album Asteraceae 2 0
Gnaphalium purpureum Asteraceae 13 14
Gnaphalium ramosissimum Asteraceae 17 25
Hesperocnide tenella Urticaceae 2 4
Heterotheca grandiflora Asteraceae 6 0
Hypochoeris glabra Asteraceae 51 25
Koeleria phleoides Poaceae 4 0
Layia glandulosa Asteraceae 9 7
Layia paniculata Asteraceae 2 0
Linaria canadensis var. texana Scrophulariaceae 26 4
Loeflingia squarrosa Caryophyllaceae 9 0
Lotus hamatus Fabaceae 4 0
Lotus strigosis Fabaceae 32 18
Lupinus bicolor Fabaceae 2 4
Lupinus truncatus Fabaceae 6 0
Madia exigua Asteraceae 2 4
Malacothrix californica Asteraceae 9 qi
Malacothrix clevelandii Asteraceae 4 4
Melilotus indica Fabaceae 0 7
Microseris heterocarpa Asteraceae 2 4
Microseris linearifolia Asteraceae 9 4
Mimulus subsecundus Scrophulariaceae 2 0
Navarretia atractyloides Polemoniaceae 51 18
Orthocarpus purpurascens Scrophulariaceae 2 0
Pectocarya penicillata Boraginaceae 2 0)
Phacelia douglasii Hydrophyllaceae 2 4
Plagiobothrys sp. Boraginaceae 2 0
Plantago erecta Plantaginaceae 2 0
Polycarpon depressum Caryophyllaceae 28 4
Polypogon monspeliensis Poaceae 2 0
Pterostegia drymarioides Polygonaceae 36 29
Rafinesquia californica Asteraceae 15 21
Salvia columbariae Lamiaceae 9 0

1988] DAVIS ET AL.: MARITIME CHAPARRAL 195
CONTINUED.

Frequency
Shrub Oak

Species Family plots plots
Schismus barbatus Poaceae 2 0
Scrophularia sp. Scrophulariaceae 2 0
Senecio californicus Asteraceae 9 11
Senecio vulgaris Asteraceae 2 7
Silene gallica Caryophyllaceae 2 0
Sonchus asper Asteraceae 13 18
Sonchus oleraceus Asteraceae pe 11
Stellaria media Caryophyllaceae 0 qi
Stylocline gnaphalioides Asteraceae 11 4
Thelypodium lasiophyllum Brassicaceae 0 4
Trifolium microcephalum Fabaceae 2 0
Trifolium sp. Fabaceae 0 4
Vulpia bromoides (L.) S. F. Gray Poaceae 15 4
*Vulpia megalura Nutt. Poaceae 23 18

Vulpia microstachys (Nutt.) Benth.

var. ciliata (Beal) Lonard & Gould Poaceae 0 4
*Vulpia myuros (L.) C. C. Gmelin Poaceae 15 0
Vulpia octoflora (Walt.) Rydb. Poaceae 66 39

ANNOUNCEMENT

SIXTH WILDLAND SHRUB SYMPOSIUM

The Shrub Research Consortium is sponsoring the Sixth Wildland
Shrub Symposium, 5-7 April 1989, at the Holiday Inn, Las Vegas, NV.
The symposium will address topics in wildland shrub biology and man-
agement including cheatgrass invasion, shrub die-off, and other aspects
of shrub biology and management. A field trip is planned to the Nevada
Test Site of the U.S. Department of Energy. Both contributed and in-
vited papers will be presented. Contributed presentations will be 20
minutes. The proceedings will be published by the USDA Forest Service,
Intermountain Research Station.

If you would like to present a paper, send a title and abstract by 15
December 1988, to Dr. E. M. Romney, Laboratory of Biomedical and
Environmental Sciences, University of California, Los Angeles, 900
Veteran Avenue, Los Angeles, CA 90024. To receive preregistration
materials and information please contact Keith McNeil, Division of
Continuing Education, University of Nevada, Las Vegas, 4505 Uni-
versity Parkway, Las Vegas, NV 89154; (702) 739-3707.

INVASION OF CARPOBROTUS EDULIS AND
SALIX LASTIOLEPIS AFTER FIRE IN A
COASTAL CHAPARRAL SITE IN
SANTA BARBARA COUNTY,
CALIFORNIA

PAUL H. ZEDLER and GERALD A. SCHEID
Biology Department,
San Diego State University,
San Diego, CA 92182-0057

ABSTRACT

Observations in permanent plots after a 1982 controlled fire in chaparral vegetation
in coastal Santa Barbara County, California demonstrate that Carpobrotus edulis, the
common introduced ice plant, increased substantially along with other native plants
capable of invading disturbed sites such as Salix lasiolepis. Although fire is a natural
disturbance, it can favor the spread of invasive exotics when a seed source is available.
Controlled burning programs must consider the possibility and risks of invasion by
exotics.

The importance of human disturbances such as grazing, agricul-
ture, and road construction in promoting the invasion of exotics 1s
well known (Elton 1958). It is less clear if natural disturbance factors
such as fire hinder or assist invasion (Johnstone 1986). A well-
accepted explanation for weed invasion is that human disturbance
creates a new environment in which the native plants are at a dis-
advantage with respect to invaders. Thus, the argument can be made
that fire in a landscape where it has a long history should not give
an advantage to exotics. On the contrary, the native plants, which
are presumably fitted to the special local characteristics of the fires,
might be favored. This theoretical reasoning is given practical sup-
port by burning experiments that have shown a decrease in exotics
(e.g., Hillyard 1985). It is the purpose of this paper to demonstrate
that burning by no means inevitably favors natives and may, in
some instances, promote the spread of exotics.

The exotic studied here, Carpobrotus edulis (L.) Bolus (“‘ice plant’),
has been widely planted in California and is now viewed as a weed
(McClintock 1985) that should be eradicated in sensitive natural
habitats (e.g., Libby 1979). It is particularly aggressive in sandy
coastal sites (Griffin 1978), where it can become the dominant plant
over large areas. Populations of an exotic succulent plant such as C.
edulis might be expected to decrease with burning, and fire might
be expected to serve as a means of controlling this species. Our
results show, on the contrary, that fire can favor its expansion.

MApDRONO, Vol. 35, No. 3, pp. 196-201, 1988

1988] ZEDLER AND SCHEID: INVASION AFTER FIRE 197

STUDY AREA

The study was conducted on Burton Mesa in Santa Barbara Coun-
ty, California (34°42'30”N, 120°43’W) about 2.6 km from the ocean
to the west of the railroad tracks near the intersection of 35th Street
and California Boulevard on Vandenberg Air Force Base. The soil
type at the site is mapped as Tangair with inclusions of the poorly-
drained Narlon series (Shipman 1972). Both soils have coarse sandy
loam textures, are derived from marine deposits, and are low in
fertility.

The vegetation at the site is a distinctive central-coast phase of
chaparral. It is characterized by low, sometimes salt-spray trimmed
canopies of evergreen species with an admixture of drought-decid-
uous coastal sage scrub elements. The site also includes other species
of limited or disjunct distribution, such as Arctostaphylos rudis and
Eriodictyon capitatum.

METHODS

The data reported here were collected in conjunction with an
experimental burn of approximately 40 ha conducted in the summer
of 1982 to determine the effect of prescribed fire on E. capitatum
(Jacks, Zedler, and Scheidlinger unpubl. report). Before the fire, a
sample area of approximately 0.6 ha, delimited by clearing along a
paved road, a railroad track, and an old unpaved track, was selected
and divided into two plots of about 2500 and 3600 sq m, the larger
of which was left unburned. A 100 m transect, crossing both the
burned and unburned vegetation, was established in June 1982 be-
fore the fire and was sampled for crown cover. In addition to marking
individual E. capitatum to follow in survivorship studies, we estab-
lished four 3 x 3 m plots in the burn area centered on E. capitatum
clumps. These plots were therefore not random with respect to E.
capitatum but were not selected with reference to C. edulis. The
cover of all shrub species within these plots was recorded and the
locations of all E. capitatum were mapped before the fire. After the
fire, seedlings and sprouts were mapped.

We estimated seed production of C. edulis in 1985 by counting
the number of fruits in 40 regularly spaced meter-square quadrats,
collecting 3 fruits from each quadrat in which they were present,
and counting the seeds in a randomly selected sub-sample of 12
fruits.

RESULTS

In 1982, before the fire, C. edulis was present along the road and
the railway that bordered the site. No C. edulis plants were recorded
within the experimental area, however, which had a nearly complete

198 MADRONO [Vol. 35

TABLE 1. PRE- AND POST-BURN COVER OF SHRUBS, SUB-SHRUBS, AND Carpobrotus
edulis ON A CHAPARRAL SITE ON VANDENBERG AIR FORCE BASE, SANTA BARBARA
COUNTY, CALIFORNIA. Transect lengths were 60 m for 1982 and 100 m for 1985.
Live cover values include overlap. Bare ground is area not covered by live or dead
plant canopies. Nomenclature after Smith (1976).

Pre-burn 1982 cover Post-burn 1985 cover

(%) (%)
Species Live Dead Live Dead
Adenostoma fasciculatum 45.3 1.2 4.3 0.0
Arctostaphylos purissima 39.3 0.3 1.1 0.0
Arctostaphylos rudis 15.9 1EF 1.0 0.0
Carex sp. 0.0 0.0 0.3 0.0
Carpobrotus edulis 0.0 0.0 26.2 0.3
Ceanothus impressus 0.0 0.0 0.1 0.0
Ceanothus ramulosus 12 2.6 0.5 0.0
Eriodictyon capitatum 3.2 |fe? 1.7 0.0
Haplopappus ericoides 1.0 0.1 0.0 0.0
Helianthemum scoparium 0.0 0.0 30.4 1.9
Lotus scoparius 0.0 0.0 3.0 0.9
Salvia mellifera 3.9 0.4 3.6 0.0
Bare ground 1.8 — 33.1 —

cover of living or dead shrub canopies of primarily evergreen species
(Table 1). Because of this dense cover, we cannot assert that C. edulis
was not present somewhere in the experimental area, but there is
no doubt that its total density was negligible. In contrast, in 1985,
three years after the fire, the cover of C. edulis was 26%, making it
the second-most prevalent post-fire perennial plant (Table 1).

Observations in the permanent plots confirm that seedling estab-
lishment is responsible for the increase in C. edulis. These plots were
resampled in 1983, and the location of C. edulis and shrub seedlings
was recorded (Table 2). Seedlings of C. edulis were recorded at an
average density of over 7000/ha. A 1985 resample relocated 70%
of these, indicating a high survivorship. Three new plants of C. edulis
were found in 1985 that may have been established in the second
season of recovery but more probably were missed in the initial
survey.

Although C. edulis has been reported to reproduce only vegeta-
tively (McClintock 1985), the observed seedling establishment ob-
viously contradicts this. We collected fruits and found an average
of 5.3 (s.d. = 12.1, n = 40) fruits/m? and an average of 1004 seeds/
fruit (s.d. = 431, n = 12). This indicates a 1985 seed production of
over 5.3 million seeds/ha. This figure can be expected to vary from
year to year and place to place, but the numbers serve to show that
C. edulis can have prodigious seed production.

Coastal sage scrub communities are particularly vulnerable to
changes in species composition (i.e., invasion) when, as in this case,

1988] ZEDLER AND SCHEID: INVASION AFTER FIRE eae

TABLE 2. ABUNDANCE OF SEEDLINGS IN PERMANENT QUADRATS NOTED IN JULY
1983 AFTER THE SUMMER 1982 PRESCRIBED BURN. Values are based on averages of
four 3 x 3 m plots. s.e. represents the standard error of the mean for the sample of
four plots.

Species Number/ha S.e.
Arctostaphylos rudis 2230 4450
A. purissima 41,075 18,000
Adenostoma fasciculatum 1650 1650
Salvia mellifera 6930 2700
Ceanothus ramulosus 825 830
C. impressus 1375 1380
Salix lasiolepis 6400 715
Baccharis pilularis 4425 1211
Lotus scoparius 3600 1895
Carpobrotus edulis 7780 2100
Solanum xantii 4700 1470

the vegetation is composed mainly of species with no ability (e.g.,
Ceanothus ramulosus, Arctostaphylos purissima) or only a weak abil-
ity (e.g., Salvia mellifera) to resprout after fire (Westman and O’Leary
1986). The establishment of several other species is evidence of this
susceptibility of burned coastal sage chaparral to invasion. The pres-
ence of seedlings of Salix lasiolepis in the burned area was very
unexpected (Table 2). No mature individuals of this species were
observed anywhere within a kilometer of the site before or after the
fire. The identity of the species was confirmed, however, by com-
parison with seedlings found along the Santa Ynez River, where the
species is very abundant. We assume that the seeds were blown onto
the site from these large stands along the Santa Ynez River which
lies about 2 km to the south.

It is not surprising that willow seeds dispersed to the site and
germinated there. What is more remarkable is that they established
and survived to early July 1983, and that a few were still present
and alive in the area the following summer. The mortality in the
permanent plots was, however, complete by the second summer.
The initial survival of the willows probably was aided by the fact
that the 1982-83 hydrologic year for the area was one of above-
normal precipitation (81.9 cm; mean rainfall is 35.2 cm), and it may
have been enhanced by the presence of a heavy clay layer overlying
shale bedrock that underlies the sandy surface soil at a depth of a
meter or more. The clay layer may have allowed high moisture
conditions to persist in the first summer. This wet year was followed
by 2 years of below-average precipitation (1983-84, 21.6 cm; 1984—
85, 26.5 cm) which, in part, may explain the lack of willow estab-
lishment.

Other exotic species besides C. edulis were observed in the burn
area. A number of Eucalyptus sp. seedlings, whose seeds evidently

200 MADRONO [Vol. 35

dispersed from a nearby windbreak, were present as were the exotics
Cortaderia sp. and Herrea elongata.

Two readily dispersed native species, Baccharis pilularis (wind-
dispersed) and Solanum xantii (animal-dispersed) were common as
seedlings in the post-fire vegetation (Table 2) even though they were
minor elements as mature shrubs before the fire. These species are
frequent in chaparral and coastal sage scrub, and it is questionable
whether their presence constitutes “invasion’’.

DISCUSSION

The substantial cover of C. edulis after the 1982 fire is evidence
that the invasion of exotic species into native vegetation can be
advanced, rather than retarded by burning. The success of C. edulis
as an invader is probably explained by its evolution with fire. Ob-
servations on Carpobrotus spp. in South Africa and Australia, where
they are native, show that they often establish from seed after fire.
Eugene Moll (pers. comm. 1985) of the University of Cape Town
has noted post-fire seedling establishment of C. dinidiata in the sand
plain and mesic mountain fynbos communities in South Africa,
although he notes that the species is most abundant in communities
that are seldom burned. Judith Brown of the Western Australian
Wildlife Research Centre (pers. comm. 1985) reports that C. edulis,
also introduced into W. Australia, establishes by seed after hot fires
in coastal locations near Perth, although in her opinion it is “‘not an
aggressive colonizer’’. Native species of Carpobrotus, however, can
invade woodlands after fire. In one case on Middle Island off the
coast of W. Australia a thick carpet of Carpobrotus developed from
seedlings after fire in a Eucalyptus angulosa—E. platypus forest un-
burned for 170 years. This evidence suggests that invasion of C.
edulis into burned chaparral at Vandenberg AFB may not be as
anomalous as it appears.

Although fire provided the “temporary invasion window” (John-
stone 1986) there must also be propagules to exploit it. We do not
know how and when the seeds of C. edulis dispersed to the site.
Fruits of C. edulis are eaten by small mammals (W. Ferren pers.
comm.) and the seeds are small and hard-coated. We suspect that
most of the seeds were deposited at the site in small mammal feces.
Therefore, the majority of the seeds probably were in the soil for
some time before the fire.

It is known that fire can be used to decrease exotics in coastal
settings. W. James Berry of the State Department of Parks and
Recreation (pers. comm. 1985) reports several successes in con-
trolling introduced species— Avena at Pt. Mugu and Malibu, Bromus
diandrus at Montana de Oro, and Brassica at Pt. Lobos. Timing was
a key element in these efforts. The burns were conducted when they

1988] ZEDLER AND SCHEID: INVASION AFTER FIRE 201

would kill most of the seed crop of the exotics without seriously
harming the desirable species, mostly perennial natives.

Our observations make it clear that these successes must not be
taken as an indication that fire will inevitably act to the favor of
natives over exotics. A case in point comes from South Africa where
the native vegetation is well adapted to survive fire, but invasion of
exotics, including pines from the Northern Hemisphere and Hakea
from Australia, has become a serious problem. Fire can be used to
reduce the abundance of some of these invaders, but others (e.g.,
Acacia) cannot be eliminated with burning (Kruger and Bigalke 1984).

It is also apparent that edge effects were important in the situation
we describe. Human disturbance along the margins of the experi-
mental plot allowed the populations of C. edulis to establish and
maintain themselves. Fire provided the opportunity for the seedlings
to establish. These results underline the importance of minimizing
edge to area ratios in retarding the expansion of exotics. They also
suggest that longer fire rotations should be favored over shorter
rotations when undesirable exotics that require open conditions for
establishment are present.

ACKNOWLEDGMENTS

We thank Clay Gautier and Paul Jacks for help in sampling and data analysis, and
Jim Johnston and others at Vandenberg AFB for funding and assistance. F. Davis,
M. Carroll, W. Ferren, and D. Keil made helpful comments on the manuscript. This
project was funded by AFOSR grant #84-0284, the U.S. Fish and Wildlife Service
Office of Endangered Species, and NSF Grant BSR-8507699.

LITERATURE CITED

ELTon, C. S. 1958. The ecology of invasions by animals and plants. Chapman and
Hall, London.

GRIFFIN, J. R. 1978. Maritime chaparral and endemic shrubs of the Monterey Bay
region, California. Madrono 25:65—112.

HILLYARD, D. S. 1985. Weed management in California’s State Park system. Fre-
montia 13:18-19.

JOHNSTONE, I. M. 1986. Plant invasion windows: a time-based classification of
invasion potential. Biol. Rev. 61:369-394.

KRUGER, F. J. and R. C. BIGALKE. 1984. Fire in fynbos. Jn P. de V. Booysen and
N. M. Tainton, eds., Ecological effects of fire in South African ecosystems, Chap-
ter 5, p. 67-114. Springer-Verlag, Berlin.

Lipsy, J. 1979. Chapter weed reports: Acacia and pampas grass in Santa Cruz.
Fremontia 6:19-20.

McCLinTock, E. 1985. Escaped exotic weeds in California. Fremontia 12:3-6.

SHIPMAN, G. E. 1972. Soil survey of the northern Santa Barbara area. U.S.D.A.
Soil Conservation Service, Washington, DC.

SMITH, C. F. 1976. A flora of the Santa Barbara region, California. Santa Barbara
Museum of Natural History, Santa Barbara, CA.

WESTMAN, W. E. and J. F. O’LEARY. 1986. Measures of resilience: the response of
coastal sage scrub to fire. Vegetatio 65:179-189.

(Received 29 May 1987; revision accepted 10 Feb 1988.)

THE VEGETATION AND ALPINE VASCULAR FLORA OF
THE SAWATCH RANGE, COLORADO

EMILY L. HARTMAN and MARY LOU ROTTMAN
Department of Biology, University of Colorado at Denver,
Denver 80204

ABSTRACT

The Sawatch Range, located in central Colorado, is part of the Southern Rocky
Mountains. Extending over 130 km in a north-south direction and 65 km in an east-
west direction, the range is the highest and one of the most extensive in the state.
No previous floristic work has been done on its tundra. Sixteen study areas distributed
throughout the entire range were analyzed over four field seasons. A vascular flora
of 289 taxa in 118 genera and 35 families is reported. Two taxa are recent new records
for the state. Eight taxa are Colorado endemics. Sixteen taxa are limited to calcareous
substrates within the range. The phytogeographic distribution of the flora is primarily
alpine (38.1%) and western North American (31.5%). According to Sorensen’s Index
of Similarity, the floristic inventory of the Sawatch Range shows an overall consistency
among the tundra vascular floras of the Mosquito and West Elk Ranges, Indian Peaks
area of the Front Range, and the San Juan Mountains.

The Sawatch Range of central Colorado (Fig. 1), a segment of the
Southern Rocky Mountains, extends over 130 km in a north-south
direction between the valleys of the Eagle River on the north and
Tomichi Creek on the south. The Arkansas River valley forms the
eastern boundary of the 65 km wide range (Chronic and Chronic
1972); the Gunnison Basin, Taylor Park, and the Elk Mountains
form the boundary on the west. Much of the 8450 sq. km area
included within the Sawatch Range lies above timberline. The range
lies between 38°30’ and 39°40'N latitude and 106°10’ and 106°50’W
longitude. The Continental Divide follows the range for more than
two-thirds of its length (Stark 1934). The Sawatch Range is the
highest range in the state. It contains four of the state’s five highest
peaks and 15 of the state’s 54 peaks that are over 4270 m.

The topography of the range is rugged and variable with a max-
imum relief of over 2135 m from the summits of the highest peaks
to the floor of the Arkansas River valley (Stark and Barnes 1935).
Extensive Pleistocene alpine and valley glaciers carved numerous
cirques and scoured broad glacial troughs on both sides of the range.
However, the rounded summit of Mt. Elbert, Colorado’s highest
mountain at 4401 m, extended beyond the upper limit of glaciation
(Stark and Barnes 1935). Glacial erosional forms in the present
alpine landscape include cirques, tarns, basins, hanging tributary
valleys, and broad U-shaped valleys. Depositional landforms of gla-

MApDRONO, Vol. 35, No. 3, pp. 202-225, 1988

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 203

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Fic. 1. Map showing the location of the Sawatch Range in the Southern Rocky
Mountains of Colorado. (Courtesy of U.S.G:S.)

cial origin consist of both terminal and lateral moraines, some of
which reach heights up to 305 m (Westgate 1905).

The Sawatch Range is composed primarily of coarse schists and
gneisses intruded by pre-Cambrian granites, and of metamorphosed
limestone and quartzite. Paleozoic sediments, including limestones,
sandstones, and shales, dip away from the crystalline core on both
sides of the range. Tertiary igneous intrusive rocks include stocks
of quartz monzonite porphyry and various porphyritic dikes and
sills. Extrusive rocks are highly localized: rhyolite occurs in the
Independence Pass area in the north, and volcanic breccia occurs in

204 MADRONO [Vol. 35

the southern part of the range near Tomichi Creek (Stark and Barnes
1932, 1935, Stark 1934, Brock and Barker 1972, Tweto 1974, Van
Loenen 1985).

Climatic data from the Sawatch Range are non-existent. However,
one can extrapolate data from a climograph for timberline elevation
of 3446 m at Climax, Colorado, located 16 km east of the Sawatch
Range (Arno and Hammerly 1984). Mean average temperature is
—1.0°C and mean average precipitation is 149 cm. The climate of
the Sawatch Range is typical of high mountainous areas in Colorado.
Summer thunderstorms, often accompanied by hail, occur almost
daily. Snowfall begins in September with major accumulations by
mid-October and continuing until late May or early June. Daytime
temperatures in the summer are temperate but frost may occur any
night (Dings and Robinson 1957).

Our investigation is the first floristic study of the alpine tundra of
the Sawatch Range. An ecological study by Loder (1964) was limited
to an area on Cottonwood Pass. The primary objective of this study
was to inventory the vascular flora of the alpine tundra of the Sa-
watch Range, thus filling in a noticeable gap in the tundra flora of
the central mountainous area of Colorado. The importance of this
range in expanding our knowledge of the Colorado tundra is am-
plified by the pivotal position the range occupies between the drier
north-south oriented ranges to the east and the more moist east-
west oriented ranges to the west. This study is part of our compre-
hensive floristic inventory of all of the tundra areas of the major
mountain ranges in Colorado.

METHODS

This study was conducted over four field seasons: 1982, 1984,
1985, and 1986. The entire 1986 season, from 10 June through 10
October, was devoted to intensive field work in the range. The au-
thors follow the definition of alpine tundra as the area above the
subalpine forest on all peaks, ridges, and in basins that rise above
the general level of tree-limit at an elevation of about 3477 m in
Colorado; however, scattered krummholz conifers may extend as
isolated patches or cushions into the alpine tundra zone (Marr 1961,
Zwinger and Willard 1972, Arno and Hammerly 1984). In the Sa-
watch Range tree-limit is found to vary from 3629 m in the lower
portions of the basins to 3782 m on the highest slopes. Sixteen alpine
areas including eight passes and eight cirque basins distributed
throughout the length of the Sawatch Range were selected for study
(Fig. 2). The passes are basically saddles flanked by convex slopes
and are relatively homogeneous in habitats and associated com-
munity types. Patterned ground forms, including sorted stripes, poly-
gons, and frost boils, are common. Cirque basins which include basin

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 205

106°50' 106°40°

106°30" 106°20° 106°10"

SAWATCH MOUNTAINS
39°40"

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Drawn by: Dale Boody & Associates, Canon City, Colorado 3-30-87

Fic. 2. Map ofthe Sawatch Range, Colorado, showing study areas. Letters indicate

passes; numbers indicate basins.

206

TABLE |.

WATCH RANGE, COLORADO.

Map
reference Location— pass Orientation Elevation (m)
a Hagerman e./w. 3637
b Independence e./w. 3689
Cc Cottonwood nnw./sse. 3698
d Tincup wnw./ese. 3707
e Cumberland n./s. 3663
f Altman nne./ssw. 3642
g Hancock n./s. 3691
h Monarch n./s. 3629
Map
reference Location— basin Orientation Elevational range (m)
] Linkins Lake se. 3660-3721
2 Mt. Champion e. 3614-3798
3 Peekaboo ne. 3538-3752
4 Mineral se. 3733-3904
5 Emma Burr e. 3736-3813
6 Fairview Peak ne. 3599-3691
7 Billings Lake se. 3569-3736
8 Island Lake n. 3630-3691

MADRONO

[Vol. 35

LOCATION, ORIENTATION, AND ELEVATION OF STUDY AREAS IN THE SA-

floors, adjacent slopes, headwalls, and ridge tops, on the other hand,
contain a greater diversity of habitats and community types. Cirque
basins in the Sawatch Range tend to have a broader and more shallow
morphology resulting in a more gradual drainage gradient than
cirque basins in other mountain ranges of the state (Hartman and
Rottman 1985a,b, 1987). Table 1 gives the orientation and elevation
of the passes and the orientation and elevational ranges of the basin
study areas.

A total of 825 communities representative of nine community
types found in the Sawatch Range were inventoried in the study.
The community types include dry, moist, and wet meadows, shrub
tundra, krummbholz, fellfield, rock-predominating (ledges, rock crev-
ices, and talus), rivulet, and snowbank. Care was taken to ensure
that the number of communities inventoried was proportionate to
the predominance of the community type in each study area.

Identification of questionable taxa were verified by R. D. Dorn,
R. L. Hartman, R. A. Price, R. C. Rollins, and W. A. Weber. A
complete voucher set for all taxa inventoried in the Sawatch Range
is deposited in the herbarium of the University of Colorado—Denver,
where the authors are associated. In addition duplicates of most
specimens are deposited in COLO. Nomenclature, for the most part,
follows Nelson and Hartman (1987). The Colorado endemics, al-
ternative names for taxa in the checklist, and 22 taxa not found in

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 207

Nelson and Hartman (1987) and indicated by an asterisk after the
authority follow Weber (1987) and Wittmann et al. (1988).

VEGETATION

Meadow communities. The dry meadow community type is most
common in the tundra of the Sawatch Range, unlike the moist mead-
ow type which predominates in the ranges to the west and southwest
in Colorado (Langenheim 1962, Webber et al. 1976, Rottman and
Hartman 1985, Hartman and Rottman 1987). Dry meadows pre-
dominate on passes, on convex mountain slopes, and on the upper
slopes of cirque basins. The predominance of dry meadows in the
Sawatch correlates with the Mosquito Range adjacent to the Arkan-
sas River valley on the east (Hartman and Rottman 1985a). We
anticipated finding different dominants in the dry meadows on passes
than in those in basins; however, Kobresia bellardii, Dryas octopetala
var. hookeriana, Geum rossii var. turbinatum, Carex elynoides, and
Salix reticulata subsp. nivalis occur in both. The only slight differ-
ence found is that Kobresia bellardii, the most frequent dominant
on the passes, is replaced by Dryas octopetala var. hookeriana in
the basins. The listed dominants appear to vary with local microen-
vironments which are not exposure correlated. Among the grami-
noid taxa, grasses exceed sedges only in dry meadows. Closed turf
communities dominated by Kobresia bellardii or Carex elynoides
have a low species richness.

Moist meadow dominants occur in three size classes: an erect tall
form such as Deschampsia caespitosa, a short caespitose form rep-
resented by Carex nigricans, and a prostrate semi-shrub form typical
of Sibbaldia procumbens and Salix reticulata subsp. nivalis. Moist
meadows on the passes are restricted to swales where Deschampsia
caespitosa 1s dominant and on the concave bases of pass slopes where
either Sibbaldia procumbens or Salix reticulata subsp. nivalis pre-
dominate. In the basins this community type occupies the mid- to
lower slopes and borders the basin floor where it may interdigitate
with shrub tundra or wet meadow. Deschampsia-dominated moist
meadow communities in the basins are associated with a wide va-
riety of tall forb taxa including Erigeron peregrinus subsp. callian-
themus, Potentilla diversifolia, and Senecio crassulus. Carex ni-
gricans and Sibbaldia procumbens are often co-dominant where basin
slope and floor interface.

Wet meadows, although relatively infrequent on passes, occur on
flat areas underlain by sporadic permafrost (Ives 1974). These areas
are often adjacent to small ponds. Dominants include Caltha lep-
tosepala, Carex scopulorum, C. nigricans, and Juncus drummondii.
This same community type in basins usually is adjacent to ponds,
lakes, or drainages. Carex scopulorum, Caltha leptosepala, Carex

208 MADRONO [Vol. 35

nigricans, and Deschampsia caespitosa are the most frequent dom-
inants. There is a conspicuous difference in species richness between
wet meadow hummock communities. Because of the persistence of
old leaf bases in hummocks dominated by Carex scopulorum, vir-
tually no other taxa can compete. Hummocks dominated by Carex
nigricans are moderately species-rich.

Shrub tundra. Shrub tundra is far more extensive in the Sawatch
Range, than in other mountain ranges studied (Rottman and Hart-
man 1985, Hartman and Rottman 1985a, 1987), often to the extent
that almost the entire basin floor is covered with this community
type. Salix brachycarpa and S. glauca subsp. glauca var. villosa
dominate in drier or well-drained areas on both passes and in basins
where they are associated with dry or moist meadow herbaceous
understory taxa. Salix planifolia is dominant primarily in hydric
sites, especially in basins where it may be co-dominant with Betula
glandulosa. Wet and moist meadow herbaceous taxa are associated
with these stands. Basin elevations for shrub tundra were observed
from 3538-3904 m.

Krummholz community. The krummholz community type occurs
as isolated patches or cushions primarily on slopes of both passes
and basins. The overwhelming dominant is Picea engelmannii; how-
ever, Pinus aristata is found on Cumberland Pass and Pinus contorta
var. latifolia occurs in Peekaboo Basin. The herbaceous understory
component of this community is represented by both alpine and
subalpine taxa. The latter are able to extend into the tundra in this
community type because of the greater snow accumulation partic-
ularly on the leeward side of the cushion and the lower evapotrans-
piration rate as a result of overstory shading during the growing
season.

Fellfield community. Fellfield community sites are characterized
by a high proportion of finely-weathered rock material (up to 80%),
coarse-textured soils, and little organic material (Willard 1963). Fell-
field communities are far more prevalent on the passes where Pa-
ronychia pulvinata, Trifolium nanum, Minuartia obtusiloba, and
Silene acaulis var. subacaulescens form the dominants. This com-
munity type in the basins is found primarily on wind-swept ridges.
The dominants listed for the passes also occur in the basins; however,
Dryas octopetala var. hookeriana is the most frequent dominant in
the latter. The typical cushion plant-dominated fellfield communi-
ties found in the Sawatch Range are similar to those of the Front
Range (Komarkova 1976, Eddleman and Ward 1984). In the Mos-
quito Range this community type appears to be restricted to moun-
tain tops and wind-swept ridges (Hartman and Rottman 1985a). In
the east-west-trending mountains of the state, the Elk Mountains

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 209

(Langenheim 1962), West Elk Mountains (Hartman and Rottman
1987), and San Juan Mountains (Webber et al. 1976, Hartman and
Rottman 1985b), this community type is poorly represented.

Rock-predominating community types. Bedrock ledges, rock crev-
ices, and talus habitats are combined into one category of minor
importance in terms of community development. Composed of either
fractured bedrock outcrops or talus, these habitats are characterized
by minimal soil accumulation available for the growth of vascular
plant taxa. As a result, community structure is replaced by a diverse
assortment of various opportunistic taxa that can tolerate the lim-
itations of the habitat substrates. In some instances, however, bed-
rock outcrops in the basins are capped by krummholz or dry meadow
communities.

Rivulet community. Small rivulets converge to form the major
drainages of most basins. Taxa bordering the rivulets are primarily
subalpine plants capable of extending upward to high elevations in
the tundra because of enhanced moisture availability. Some of the
subalpine taxa that can dominate rivulet communities are Senecio
triangularis, Aconitum columbianum, Cardamine cordifolia, Mimu-
lus guttatus, and Saxifraga odontoloma. Shrub tundra communities
often border the rivulets.

Snowbank community. Snowbanks may persist late into August
in the basins. These areas are subject to shortened growing seasons,
often permitting only vegetative growth of taxa found in the area in
front of the receding snowbank. Carex pyrenaica is a good indicator
dominant of this community type. Other taxa worthy of note are
Ranunculus adoneus, Sibbaldia procumbens, and Salix arctica. Car-
ex incurviformis appears to be restricted to this community. A wide-
spread early-melting snowbank community dominated by Vaccin-
ium caespitosum, Carex pyrenaica, and Stereocaulon sp. assumes a
deceivingly dry appearance as the season progresses.

FLORA

Comparative floristics. The alpine flora of the Sawatch Range con-
sists of 289 taxa representing 109 genera of angiosperms, four genera
and five species of gymnosperms, and five genera and five species
of pteridophytes. Our total of 289 taxa compared to the estimated
300 vascular plant taxa in the alpine tundra of Colorado (Bliss 1962)
attests to the thoroughness of this study in the Sawatch Range. The
largest family is Asteraceae with 47 taxa, followed by Cyperaceae,
Poaceae, Brassicaceae, Ranunculaceae, Caryophyllaceae, Rosaceae,
and Scrophulariaceae with 34, 23, 21, 16, 15, 14, and 14 taxa,
respectively. Comparing these leading families with those of other
Colorado alpine floras, the greatest similarity is to the Mosquito

210 MADRONO [Vol. 35

Range to the east (Hartman and Rottman 1985a), the only difference
being the addition of Ranunculaceae which exceeds Rosaceae and
Scrophulariaceae in the Sawatch alpine. In the Ruby Range of the
West Elk Mountains (Hartman and Rottman 1987), San Juan Moun-
tains (Hartman and Rottman 1985b), and Indian Peaks area of the
Front Range (Komarkova 1976), Saxifragaceae is among the leading
families. Similarities between alpine floristic inventories of the Sa-
watch Range and the above ranges were analyzed by using Sorensen’s
Index of Similarity (Mueller-Dombois and Ellenberg 1974). The
greatest similarity, 73.3%, again occurs between the Sawatch and
Mosquito ranges; however, similarities to the West Elk (73.1%),
Indian Peaks area of the Front Range (72.6%), and San Juan Moun-
tains (72.5%) show an overall consistency among the tundra floras
of the various ranges. Differences between the inventories at the
individual taxon level are in many cases striking, and relate primarily
to apparent distributional ranges of the plants or to substrate and
microenvironmental dissimilarities.

Rare taxa with restricted occurrences in the Colorado tundra may
reflect local environmental conditions or distributional ranges. The
following taxa in the Sawatch Range are considered rare or infre-
quent by Weber (1987): Anemone parviflora, Asplenium viride, As-
tragalus aboriginum, Carex arctogena, Draba streptobrachia, Gen-
tianella tenella, Physaria alpina, Ranunculus pygmaeus var.
pygmaeus, Saxifraga adscendens subsp. oregonensis, and Ligularia
taraxacoides. From our experiences in the various tundras of the
state we would add Arabis lemmonii, Astragalus molybdenus, Draba
incerta, Erigeron vagus, and Ligularia porteri to this list. Carex
vernacula, Erigeron grandiflorus, and Ligularia soldanella, all of
which Weber (1987) considers rare, are infrequent in the Sawatch
Range. Two new state records recently were reported for the Sawatch
Range from Mt. Champion basin. These are Draba apiculata var.
apiculata (Price et al. 1985) and Antennaria aromatica (O’Kane et
al. 1988).

Phenology. Our 1986 field season extended from 10 June through
10 October, and permitted some interesting phenological observa-
tions. Complete snow release in a fellfield on Independence Pass
was observed 11 June with 14 taxa in prime anthesis while all other
communities on the pass were snow-covered. By 2 July the pass was
completely snow-free and vegetative growth was in progress. Thlaspi
montanum var. montanum and Smelowskia calycina var. ameri-
cana (fellfield constituents) were in fruit on 10 July. By 5 September
this pass and all others were showing the typical reddish-brown
coloration of autumn. In the basins, however, a number of extremely
dwarfed moist meadows, dominated by Geum rossii var. turbinatum,
Sibbaldia procumbens, and Erigeron melanocephalus, often found

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE AU

adjacent to talus or at the interface of slopes and basin floor were
in prime anthesis in late August and early September. Ronning (1968)
and Owen (1976) have suggested that both stature and late anthesis
may relate to either cold soil temperatures or late snow release. On
10 September the entire range was blanketed by the first snow.
Seasonal snow accumulations were in evidence by 3 October. At
this time some snow depth measurements were made on Indepen-
dence Pass in communities studied earlier in the season. Even at
this early date it was possible to correlate relative snow accumula-
tion, wind patterns of snow distribution, and community types.
Snow depths ranged from 2.4—12.6 cm in fellfield communities to
a maximum of 10.7—30.0 cm in moist meadows. Shrub tundra com-
munities were intermediate with 12.6 cm on the windward side and
20.8 cm on the leeward side. Obviously these values cannot reflect
the total snow accumulation that will occur; however, we feel that
the relative ratios of snow depth to community type are maintained
throughout the winter and spring seasons.

Calcareous substrates. Although the Sawatch Range is primarily
metamorphic in composition, three of our study areas contain lo-
calized occurrences of limestone and dolomite. The following taxa
were found only on these calcareous substrates: Anemone parviflora,
Asplenium viride, Astragalus aboriginum, A. molybdenus, Juncus
albescens, J. castaneus, Ligularia taraxacoides, Oreochrysum parryi,
Oxytropis podocarpa, O. viscida, Physaria alpina, Potentilla ovina,
Pyrrocoma uniflora, and Senecio canus. Some of these taxa were
reported also for limestone substrates in the Mosquito Range (Hart-
man and Rottman 1985a), Elk Mountains (Langenheim 1962), and
Montana (Bamberg and Major 1968). Schroeter (1926), Nordhagen
(1955), Curry (1962), and Bamberg and Major (1968) consider Dryas
octopetala var. hookeriana to be an indicator of calcareous rock or
soil, but Murdock (1951), Johnson and Billings (1962), and Willard
(1963) fail to substantiate this substrate specificity. In the Sawatch
Range the occurrence of Dryas octopetala var. hookeriana does not
appear to be substrate specific and, as suggested by Willard (1963),
the var. hookeriana may be a Southern Rocky Mountain ecotype
with low calcium requirements. As previously mentioned, shrub
tundra forms extensive stands both on passes and in basins in the
range; however, in our calcareous study sites this community type
is extremely limited, perhaps reflecting the porosity and downward
percolation of water characteristic of limestone-dolomite areas.

Phytogeography. Table 2 shows the phytogeographic distribution
of the alpine flora of the Sawatch Range compared to other ranges
in Colorado. Four elements are recognized, each of which may be
combined with more specific geographical subelements (Komarkova

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[Vol. 35

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1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 213

1976). The Rocky Mountains subelement includes the Northern
Rocky Mountain province south to the Laramie Basin in Wyoming.
The Southern Rocky Mountains subelement includes southern Wy-
oming, Colorado, New Mexico, and Arizona. The Colorado subele-
ment contains taxa endemic to Colorado. Phytogeographic deter-
minations for taxa are based upon distributions given by Porsild
(1957), Weber (1965), Munz and Keck (1970), Komarkova (1976),
Cronquist et al. (1977), Porsild and Cody (1980), and Moss (1983).

The largest part of the vascular flora of the Sawatch Range is made
up of Alpine (38.1%) and Western North American (31.5%) taxa.
The Circumpolar subelement (19.4%), which is primarily linked with
the Arctic-Alpine element, is another important component of the
flora. The North American-Asiatic subelement (6.6%), although
smaller in the Sawatch Range than in the other ranges compared,
still indicates a stronger affinity to the Asiatic alpine flora than to
the European alpine flora. The Alpine element is consistently the
highest in all ranges compared, with the Arctic-Alpine element sec-
ond in all but the Sawatch Range and West Elk Mountains where
the Boreal-Montane is better represented. The proximity of the Mos-
quito Range to the Sawatch Range makes comparisons of these two
particularly significant. For the most part there is a greater corre-
lation between subelements of the two than between elements.

Colorado endemics found in this study include A/sinanthe mac-
rantha, Draba streptobrachia, Ligularia soldanella, Luzula subcap-
itata, Penstemon hallii, P. harbourii, Physaria alpina, and Potentilla
subjuga var. subjuga. Fewer Colorado endemics occur in the Sawatch
Range than in the more northerly Indian Peaks area, and in the
Mosquito Range, which is due east of the northern part of the Sa-
watch.

ANNOTATED CATALOGUE OF VASCULAR PLANT TAXA

The terms used in the annotated catalogue to describe occurrence/
abundance are a combination of abundance classes and constancy
values. In estimating abundance, standard abundance classes were
used: very abundant, abundant, frequent, occasional, and rare (Dau-
benmire 1968). As data from multiple samples of a particular com-
munity type accumulated, it was possible to analyze the inventoried
taxa and their abundance ratings on the basis of constancy of oc-
currence between samples of communities. The percentage values
of the constancy classes used were taken from Mueller-Dombois and
Ellenberg (1974): very abundant (81—100% constancy), abundant
(60.1-80% constancy), frequent (40.1-60% constancy), occasional
(20.1-40% constancy), and rare (1.5—20% constancy). The abbre-
viations used for community types for each taxon are: dry meadow
(dm); moist meadow (mm); wet meadow (wm); shrub tundra (st);

214 MADRONO [Vol. 35

krummbholz (kr); fellfield (ff); rock-predominating including bedrock
ledges, rock crevices, and talus (rp); rivulet (rv); and snowbank (sn).
In the phytogeographic citation for each taxon, the element precedes
the subelement, the two being separated by a slash (Table 2). Brack-
eted nomenclature follows Weber (1987) and Wittmann et al. (1988).
Twenty-two taxa that are indicated by an asterisk (*) after the au-
thority are not found in Nelson and Hartman (1987) and follow the
nomenclature of W. A. Weber (1987) and Wittmann et al. (1988).

PTEROPHYTA
Selaginellaceae

Selaginella densa Rydb. var. densa. Abundant; dm, mm, st, kr, ff, rp, sn; A/WNA.

Adiantaceae

Cryptogramma acrostichoides R. Br. Rare; rp; BM/NAA.

Aspleniaceae

Asplenium viride Huds. [A. trichomanes-ramosum L.]. Rare; rp; AA/C.
Cystopteris fragilis (L.) Bernh. var. fragilis. Occasional; st, rp; AA/C.
Woodsia oregana D. C. Eat. var. oregana. Occasional; dm, rp; BM/NA.

CONIFEROPHYTA
Cupressaceae

Juniperus communis L. var. depressa Pursh [J. communis L. subsp. alpina (Neilr.)
Celak.]. Rare; kr; BM/C.

Pinaceae

Abies lasiocarpa (Hook.) Nutt. var. /Jasiocarpa. Rare; kr; BM/WNA.

Picea engelmannii Parry ex Engelm. Occasional; kr; BM/WNA.

Pinus aristata Engelm.* Rare; kr; BM/WNA.

Pinus contorta Dougl. ex Loud. var. /atifolia Engelm. ex Wats. [P. contorta Dougl.
subsp. /atifolia (Engelm.) Critch.]. Rare; kr; BM/WNA.

ANTHOPHYTA — DICOTYLEDONEAE
Adoxaceae

Adoxa moschatellina L. Rare; mm; BM/C.

Apiaceae

Angelica grayi (Coult. & Rose) Coult. & Rose. Frequent; dm, mm, st, rp; A/SRM.

Cymopterus lemmonii (Coult. & Rose) Dorn [Pseudocymopterus montanus (A. Gray)
Coult. & Rose]. Occasional; dm, mm, st, kr; M/SRM.

Ligusticum tenuifolium Wats. [L. filicinum Wats. var. tenuifolium (Wats.) Math. &
Const.]. Rare; mm; BM/WNA.

Oreoxis alpina (A. Gray) Coult. & Rose. Very abundant; dm, mm, wm, st, kr, ff, rp,
sn; A/SRM.

O. bakeri Coult. & Rose.* Frequent; dm, mm, wm, st, ff, rp; A/SRM.

Oxypolis fendleri (A. Gray) Heller. Rare; rv; M/SRM.

Podistera eastwoodiae (Coult. & Rose) Math. & Const.* Rare; dm; A/SRM.

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 21S

Asteraceae

Achillea millefolium L. var. lanulosa (Nutt.) Piper [A. /anulosa Nutt.]. Frequent; dm,
mm, wm, st, kr, ff, rp; A/WNA.

Agoseris aurantiaca (Hook.) Greene var. aurantiaca. Occasional; dm, mm, rp; BM/
WNA.

A. glauca (Pursh) Raf. var. dasycephala (Torr. & A. Gray) Jeps. Rare; dm; M/WNA.

A. glauca (Pursh) Raf. var. laciniata (D. C. Eat.) Smiley. Occasional; dm, mm, kr;
BM/NA.

Antennaria aromatica Evert. Rare; ff; M/RM. #6671/2942 COLO.

A. media Greene. Abundant; dm, mm, wm, st, kr, ff, rp, sn; AA/NAE.

A. rosea Greene. Occasional; mm, st, kr; BM/NA.

A. umbrinella Rydb. Rare; dm; A/WNA.

Arnica cordifolia Hook. Rare; rp; BM/WNA.

A. mollis Hook. Rare; mm, rp; BM/NA.

A. rydbergii Greene. Occasional; dm, kr; BM/WNA.

Artemisia campestris L. subsp. borealis (Pall.) Hall & Clem. var. borealis. Occasional;
dm, mm, kr, ff, rp; AA/C.

A. scopulorum A. Gray. Very abundant; dm, mm, wm, st, kr, ff, rp, sn; A/RM.

Chaenactis alpina (A. Gray) Jones. Occasional; ff, rp; M/WNA.

Cirsium scopulorum (Greene) Ckll. Occasional; dm, rp; A/RM.

C. tweedyi (Rydb.) Petr. Occasional; mm, wm, rp; BM/NA.

Erigeron compositus Pursh. Occasional; dm, rp; BM/NA.

E. coulteri Porter. Rare; dm; BM/WNA.

E. grandiflorus Hook. Occasional; dm, mm, st, sn; AA/WNA.

E. leiomerus A. Gray. Occasional; dm, mm, ff, rp; M/RM.

E. melanocephalus (A. Nels.) A. Nels. Abundant; dm, mm, wm, rp, sn; A/SRM.

E. peregrinus (Banks ex Pursh) Greene subsp. callianthemus (Greene) Cronq. var.
callianthemus [E. peregrinus (Banks ex Pursh) Greene subsp. callianthemus
(Greene) Cronq.]. Occasional; dm, mm, wm, st, kr; BM/WNA.

E. pinnatisectus (Gray) A. Nels. Abundant; dm, mm, st, ff, rp; A/SRM.

E. simplex Greene. Very Abundant; dm, mm, wm, st, kr, ff, rp, sn; A/WNA.

E. ursinus D. C. Eat. Rare; dm; M/WNA.

E. vagus Pays.* Rare; dm; A/WNA.

Haplopappus pygmaeus (Torr. & A. Gray) A. Gray [Tonestus pygmaeus (Torr. & A.
Gray) A. Nels.]. Frequent; dm, kr, ff, rp; A/RM.

Heterotheca fulcrata (Greene) Shinners. Occasional; dm, rp; M/RM.

Hieracium gracile Hook. var. gracile [Chlorocrepis tristis (Willd. ex Spreng.) A. Love
& D. Love subsp. gracile (Hook.) W. A. Weber]. Occasional; dm, mm, st, rp;
A/WNA.

Hymenoxys grandiflora (Torr. & A. Gray ex A. Gray) K. Parker [Rydbergia grandiflora
(Torr. & A. Gray) Greene]. Frequent; dm, mm, st, kr, ff, rp; A/RM.

Ligularia amplectens (A. Gray) W. A. Weber.* Occasional; mm, wm, rp; M/RM.

L. holmii (Greene) W. A. Weber.* Abundant; dm, mm, ff, rp, sn; A/RM.

L. porteri (Greene) W. A. Weber.* Rare; rp; A/RM.

L. soldanella (A. Gray) W. A. Weber. Occasional; ff, rp; A/CO.

L. taraxacoides (A. Gray) W. A. Weber.* Rare; rp; A/SRM.

Oreochrysum parryi (A. Gray) Rydb.* Rare; dm; M/SRM.

Pyrrocoma uniflora (Hook.) Greene. Occasional; dm, mm, st, sn; M/WNA.

Senecio atratus Greene. Rare; rp; A/SRM.

S. canus Hook. [Packera cana (Hook.) W. A. Weber & A. L6ve]. Occasional; dm,
rp; M/WNA.

S. crassulus A. Gray. Occasional; dm, mm, wm, st, rp; BM/WNA.

S. dimorphophyllus Greene var. dimorphophyllus [Packera dimorphophylla (Greene)
W. A. Weber & A. Love]. Frequent; mm, wm, st, rp, rv, sn; M/RM.

S. fremontii Torr. & A. Gray var. blitoides (Greene) Cronq. [S. fremontii Torr. & A.
Gray subsp. blitoides (Greene) Cronq.]. Occasional; rp; A/SRM.

216 MADRONO [Vol. 35

S. integerrimus Nutt. var. integerrimus. Occasional; mm; M/WNA.

S. triangularis Hook. Occasional; mm, wm, st, rv; BM/WNA.

S. werneriaefolius (A. Gray) A. Gray [Packera werneriifolia (A. Gray) W. A. Weber
& A. Léve]. Abundant; dm, mm, kr, ff, rp, sn; A/WNA.

Solidago spathulata DC. var. nana (A. Gray) Cronq. Frequent; dm, mm, st, kr, ff,
rp; A/WNA.

Taraxacum ceratophorum (Ledeb.) DC. [T. ovinum Greene]. Occasional; kr, ff, rp,
sn; AA/C.

Betulaceae

Betula glandulosa Michx. Occasional; wm, st, kr; BM/NA.

Boraginaceae

Eritrichium nanum (Vill.) Schrad. var. elongatum (Rydb.) Cronq. [Eritrichum are-
tioides (Cham.) DC.]. Frequent; dm, mm, st, ff, rp; AA/NAA.

Mertensia ciliata (James ex Torr.) G. Don. Occasional; mm, wm, st, rp, rv; BM/
WNA.

M. lanceolata (Pursh) A. DC.* Occasional; dm, mm, st; A/SRM.

M. viridis (A. Nels.) A. Nels. [M. lanceolata (Pursh) A. DC. var. viridis A. Nels.].
Very abundant; dm, mm, wm, st, kr, ff, rp, rv; A/WNA.

Brassicaceae

Arabis drummondii A. Gray [Boechera drummondii (A. Gray) A. Love & D. Léve].
Rare; st, kr; BM/NA.

A. lemmonii Wats. [Boechera lemmonii (Wats.) W. A. Weber]. Rare; rp; A/WNA.

Cardamine cordifolia A. Gray var. cordifolia. Occasional; mm, st, rp, rv; BM/WNA.

Descurainia richardsonii (Sweet) O. E. Schulz var. richardsonii. Rare; st, rp; BM/NA.

Draba apiculata C. L. Hitche. var. apiculata. Rare; rp; A/RM. #6025/2296 COLO.

. aurea Vahl ex Hornem. var. aurea. Abundant; dm, mm, st, kr, ff, rp; AA/C.

. cana Rydb. Occasional; dm, mm, st, rp; AA/C.

. crassa Rydb. Occasional; rp; A/RM.

. crassifolia Grah. var. crassifolia. Very abundant; dm, mm, wm, st, kr, ff, rp, sn;

AA/NAE.

fladnizensis Wulf. Rare; dm, st; AA/C.

. incerta Pays. Rare; rp; A/RM.

. lonchocarpa Rydb. var. lonchocarpa. Frequent; dm, mm, rp; AA/NA.

oligosperma Hook. Occasional; dm, mm, rp; AA/WNA.

. spectabilis Greene var. spectabilis. Rare; dm, kr; M/RM.

. streptobrachia Price. Rare; dm, ff; A/CO.

. streptocarpa A. Gray var. streptocarpa. Occasional; ff, rp; A/SRM.

Erysimum nivale (Greene) Rydb. [E. capitatum (Dougl.) Greene]. Frequent; dm, mm,
st, ff, rp; A/SRM.

Physaria alpina Roll. Rare; rp; A/CO.

Rorippa curvipes Greene var. alpina (Wats.) Stuckey. Rare; wm, rv; A/RM.

Smelowskia calycina (Steph. ex Willd.) C. A. Mey. var. americana (Regel & Herd.)
Drury & Roll. [S. calycina (Steph. ex Willd.) C. A. Mey.]. Abundant; dm, mm,
st, kr, ff, rp; AA/NAA.

Thlaspi montanum L. var. montanum [Noccaea montana (L.) F. K. Mey.]. Very
abundant; dm, mm, wm, st, kr, ff, rp, sn; AA/C.

Bo Sooo. UV eg

Campanulaceae

Campanula rotundifolia L. Rare; dm, rp; BM/C.
C. uniflora L. Frequent; dm, mm, st, ff, rp; A/C.

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 217

Caryophyllaceae

Alsinanthe macrantha (Rydb.) W. A. Weber. Rare; rp; A/CO.

Arenaria congesta Nutt. var. congesta [Eremogone congesta (Nutt. ex Torr. & A.
Gray) Ikonn.]. Occasional; dm, rp; M/WNA.

A. fendleri A. Gray var. fendleri [Eremogone fendleri (A. Gray) Ikonn.]. Abundant;
dm, mm, st, kr, ff, rp; A/SRM.

Cerastium arvense L. [Alsine media L.]. Occasional; dm, mm, st, ff, rp; BM/C.

C. beeringianum Cham. & Schlecht. [C. beeringianum Cham. & Schlecht. subsp.
earlei (Rydb.) Hulten]. Occasional; dm, rp; A/RM.

Gastrolychnis kingii (Wats.) W. A. Weber.* Rare; dm, rp; A/SRM.

Minuartia austromontana S. J. Wolf & Packer [Alsinanthe stricta (Sw.) Reichenb.].
Rare; mm, ff, rp; AA/NA.

M. obtusiloba (Rydb.) House [Lidia obtusiloba (Rydb.) A. Léve & D. L6ve]. Very
abundant; dm, mm, st, kr, ff, rp, sn; AA/NAA.

M. rubella (Wahlenb.) Hiern [7ryphane rubella (Wahlenb.) Reichenb.]. Frequent; dm,
mm, st, ff, rp; AA/C.

Paronychia pulvinata A. Gray. Occasional; dm, ff, rp; A/SRM.

Sagina saginoides (L.) Karst. Frequent; dm, mm, wm, rp, sn; AA/C.

Silene acaulis (L.) Jacq. var. subacaulescens (F. N. Williams) Fern. & St. John [S.
acaulis subsp. subacaulescens (F. N. Will.) C. L. Hitchce. & Maguire]. Very abun-
dant; dm, mm, st, kr, rp, sn; AA/NAA.

S. dr'ummondii Hook. var. drummondii [Gastrolychnis drummondii (Hook.) A. Love
& D. Léve]. Occasional; dm, wm, kr; BM/NA.

Stellaria longipes Goldie var. longipes. Frequent; dm, mm, st, kr, rp; BM/NA.

S. umbellata Turcz. ex Kar. & Kir. Frequent; dm, mm, st, rp, sn; A/NAA.

Crassulaceae

Sedum integrifolium (Raf.) A. Nels. subsp. integrifolium [Rhodiola integrifolia Raf.].
Abundant; dm, mm, wm, rp, rv; AA/NAA.

S. lanceolatum Torr. var. lanceolatum [Amerosedum lanceolatum (Torr.) A. Love &
D. Love]. Frequent; dm, mm, st, kr, ff, rp; A/RM.

S. rhodanthum A. Gray [Clementsia rhodantha (A. Gray) Rose]. Occasional; mm,
wm, st, rp, rv; A/RM.

Ericaceae

Arctostaphylos uva-ursi (L.) Spreng. subsp. uva-ursi [A. adenotricha (Fern. & Macbr.)
A. Léve, D. Love & Kapoor]. Rare; dm, st; BM/NA.

Gaultheria humifusa (Grah.) Rydb. Rare; mm; BM/WNA.

Kalmia microphylla (Hook.) Heller var. microphylla. Rare; wm; BM/WNA.

Vaccinium caespitosum Michx. [V. cespitosum Michx.]. Very abundant; dm, mm,
wm, st, kr, rp, sn; BM/NA.

V. myrtillus L. subsp. oreophilum (Rydb.) A. L6ve, D. Love & Kapoor. Rare; dm,
mm, rp; BM/C.

V. scoparium Leib. ex Cov. Rare; kr; BM/WNA.

Fabaceae

Astragalus aboriginum Richards. Rare; dm; BM/WNA.

A. molybdenus Barneby.* Rare; dm; AA/C.

A. tenellus Pursh. Rare; dm, mm; BM/WNA.

Oxytropis deflexa (Pall.) DC. var. sericea Torr. & A. Gray. Occasional; dm, mm, st,
rp; BM/C.

O. parryi A. Gray. Rare; st; A/WNA.

O. podocarpa A. Gray. Occasional; dm, ff, rp; AA/C.

O. sericea Nutt. var. sericea. Rare; dm, st; BM/NA.

O. viscida Nutt. var. viscida. Rare; mm; A/WNA.

218 MADRONO [Vol. 35

Trifolium dasyphyllum Torr. & A. Gray var. dasyphyllum. Abundant; dm, mn, st,
kr, ff, rp; A/RM.

T. nanum Torr. Abundant; dm, mm, st, kr, ff, rp; A/RM.

T. parryi A. Gray var. parryi. Frequent; dm, mm, wm, st, rp; A/RM.

Gentianaceae

Frasera speciosa Dougl. ex Griseb. Rare; dm; BM/WNA.

Gentiana algida Pall. [Gentianodes algida (Pall.) A. Love & D. Love]. Occasional;
dm, mm, st; AA/NAA.

G. calycosa Griseb. [Pneumonanthe parryi (Engelm.) Greene]. Rare; dm, mm; A/WNA.

G. prostrata Haenke ex Jacq. [Chondrophylla prostrata (Haenke ex Jacq.) J. P. An-
ders.]. Occasional; dm, mm, st; AA/NAA.

Gentianella amarella (L.) Borner [G. acuta (Michx.) Hiit.]. Occasional; dm, mm, st,
ff, rp; BM/C.

G. tenella (Rottb.) Borner [Comastoma tenellum (Rottb.) Toyokuni]. Rare; mm;
AA/C.

Gentianopsis barbellata (Engelm.) IItis. Rare; dm, ff; A/SRM.

G. detonsa (Rottb.) Ma var. elegans (A. Nels.) N. Holmgren [G. thermalis (Kuntze)
Iltis]. Occasional; mm; A/RM.

Swertia perennis L. Occasional; mm, wm, st; A/C.

Grossulariaceae

Ribes montigenum McClat. Occasional; st, kr, rp; BM/WNA.

Hydrophyllaceae

Phacelia glandulosa Nutt. Rare; rp; BM/WNA.
P. hastata Dougl. ex Lehm. var. hastata. Rare; dm, rp; M/NA.
P. sericea (Grah. ex Hook.) A. Gray var. sericea. Frequent; dm, mm, st, kr, ff A/WNA.

Onagraceae

Epilobium anagallidifolium Lam. Frequent; wm, st, rp, rv, sn; AA/C.
E. hornemannii Reichenb. subsp. hornemannii. Rare; wm, st; AA/C.

Polemoniaceae

Phlox pulvinata (Wherry) Cronq. [P. condensata (A. Gray) E. Nels.]. Abundant; dm,
st, ff, rp; A/SRM.

Polemonium pulcherrimum Hook. var. pulcherrimum [P. pulcherrimum Hook. subsp.
delicatum (Rydb.) Brand]. Rare; st, kr; M/SRM.

P. viscosum Nutt. Abundant; dm, mm, st, kr, ff, rp; A/WNA.

Polygonaceae

Eriogonum jamesii Benth. var. xanthum (Stokes) Reveal.* Rare; dm, st; A/WNA.

Oxyria digyna (L.) Hill. Frequent; mm, rp, rv; AA/C.

Polygonum bistortoides Pursh [Bistorta bistortoides (Pursh) Small]. Very abundant;
dm, mm, wm, st, kr, ff, rp; A/WNA.

P. douglasii Greene var. douglasii. Rare; ff; BM/NA.

P. viviparum L. var. viviparum [Bistorta vivipara (L.) S. Gray]. Abundant; dm, mm,
wm, st, kr, ff, rp; AA/C.

Portulacaceae

Claytonia megarhiza (A. Gray) Parry ex Wats. var. megarhiza [C. megarhiza (Parry
ex A. Gray) Wats.]. Frequent; ff, rp; A/RM.

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE 219

Lewisia pygmaea (A. Gray) B. L. Robins. [Oreobroma pygmaea (A. Gray) Howell].
Frequent; dm, mm, st, kr, rp, sn; A/WNA.

Primulaceae

Androsaceae septentrionalis L. Abundant; dm, mm, wm, st, kr, ff, rp; AA/C.
Primula angustifolia Torr.* Frequent; dm, mm, ff, rp, sn; A/SRM.
P. parryi A. Gray. Frequent; mm, wm, rp, rv, sn; A/RM.

Ranunculaceae

Aconitum columbianum Nutt. var. columbianum. Rare; mm, rv; BM/WNA.

Anemone multifida Poir. var. multifida [A. multifida Poir. var. globosa (Nutt.) Torr.
& A. Gray]. Occasional; dm, st, kr, rp; BM/NA.

A. narcissiflora L. subsp. zephyra (A. Nels.) A. Love, D. Love & Kapoor [Anemo-
nastrum narcissiflorum (L.) Holub. subsp. zephyrum (A. Nels.) W. A. Weber].
Frequent; dm, mm, wm, st, kr, ff, sn; A/SRM.

A. parviflora Michx. Rare; mm, rp; BM/NAA.

A. patens L. [Pulsatilla patens (L.) Mill. subsp. multifida (Pritz.) Zamels]. Rare; rp;
BM/NA.

Aquilegia caerulea James. Frequent; dm, mm, st, kr, rp; M/RM.

Caltha leptosepala DC. subsp. leptosepala var. leptosepala [Psychrophila leptosepala
(DC.) W. A. Weber]. Occasional; mm, wm, st, rv, sn; A/WNA.

Delphinium barbeyi (Huth) Huth. Rare; mm; M/SRM.

Ranunculus adoneus A. Gray. Infrequent; mm, sn; A/WNA.

R. eschscholtzii Schlecht. var. eschscholtzii. Frequent; dm, mm, wm, rp, rv, sn; AA/
NAA.

R. inamoenus Greene var. alpeophilus (A. Nels.) Benson [R. inamoenus Greene].
Rare; mm; M/WNA.

R. macauleyi A. Gray.* Rare; mm; A/SRM.

R. pedatifidus J. E. Sm. var. affinis (R. Br.) Benson [R. peditifidus J. E. Sm.]. Rare;
dm; AA/C.

R. pygmaeus Wahlenb. var. pygmaeus. Rare; wm; AA/C.

Thalictrum alpinum L. var. hebetum Boivin [T. alpinum L.]. Frequent; dm, mm,
wm, st, rp, sn; A/WNA.

Trollius laxus Salisb. [T. albiflorus (A. Gray) Rydb.]. Occasional; mm, st; BM/WNA.

Rosaceae

Dryas octopetala L. var. hookeriana (Juz.) Breit. [D. octopetala L. subsp. hookeriana
(Juz.) Hulten]. Frequent; dm, mm, st, kr, ff, rp; A/RM.

Fragaria vesca L. var. bracteata (Heller) R. J. Davis [F. vesca L. subsp. bracteata
(Heller) R. J. Davis]. Rare; dm, kr; BM/NA.

Geum rossii (R. Br.) Ser. var. turbinatum (Rydb.) C. L. Hitche. [Acomastylis rossii
(R. Br.) Greene subsp. turbinata (Rydb.) W. A. Weber]. Very abundant; dm,
mm, wm, st, kr, ff, rp, rv, sn; AA/NA.

Potentilla concinna Richards. var. concinna. Rare; dm, st; M/WNA.

P. diversifolia Lehm. var. diversifolia. Very abundant; dm, mm, wm, st, kr, ff, rp, sn;
A/WNA.

P. fruticosa L. [Pentaphylloides floribunda (Pursh) A. Love]. Occasional; dm, st, rp;
BM/C.

P. hookeriana Lehm. subsp. hookeriana var. hookeriana [P. hookeriana Lehm.].
Frequent; dm, st, ff, rp; AA/NAA.

P. nivea L. Occasional; dm, ff, rp; AA/C.

P. ovina Macoun var. decurrens (Wats.) Welsh & B. C. Johnston. Rare; dm; M/RM.

P. ovina Macoun var. ovina. Rare; dm; M/WNA.

P. pulcherrima Lehm. Occasional; dm, mm, st, ff, rp; BM/WNA.

220 MADRONO [Vol. 35

P. rubricaulis Lehm. Occasional; dm, kr, ff, rp; AA/NA.
P. subjuga Rydb. var. subjuga. Frequent; dm, ff, rp; A/CO.
Sibbaldia procumbens L. Abundant; dm, mm, wm, st, rp, sn; AA/C.

Salicaceae

Salix arctica Pall. [S. arctica Pall. subsp. petraea (Anderss.) A. Love, D. Love &
Kapoor]. Abundant; dm, mm, wm, st, ff, rp, sn; A/WNA.

S. brachycarpa Nutt. subsp. brachycarpa var. brachycarpa [S. brachycarpa Nutt.].
Frequent; dm, mm, st, kr, rp; BM/NA.

S. glauca L. subsp. glauca var. villosa (Hook.) Anderss. Occasional; mm, st, kr; BM/
WNA.

S. planifolia Pursh subsp. planifolia var. planifolia [S. planifolia Pursh]. Occasional;
mm, wm, st, ff; BM/NA.

S. reticulata L. subsp. nivalis (Hook.) A. Love, D. Love & Kapoor. Very abundant;
dm, mm, wm, st, kr, ff, rp, sn; A/WNA.

Saxifragaceae

Heuchera parvifolia Nutt. ex Torr. & A. Gray [H. parvifolia Nutt. ex Torr. & A. Gray
var. nivalis (Rosend.) A. Love, D. Love & Kapoor]. Abundant; dm, mm, st, kr,
ff, rp; M/RM.

Micranthes oregana (Howell) Small.* Occasional; mm, wm, st, rv; M/WNA.

Saxifraga adscendens L. subsp. oregonensis (Raf.) Breit. [Muscaria adscendens (L.)
Small]. Occasional; rp; AA/NAE.

S. bronchialis L. subsp. austromontana (Wieg.) G. N. Jones [Ciliaria austromontana
(Wieg.) W. A. Weber]. Frequent; dm, mm, st, kr, ff, rp; A/WNA.

S. caespitosa L. var. minima Blank. [Muscaria delicatula Small]. Rare; rp; AA/C.

S. cernua L. Occasional; rp; AA/C.

S. debilis Engelm. ex A. Gray [S. Ayperborea R. Br. subsp. debilis (Engelm. ex A.
Gray) A. Love, D. Love & Kapoor]. Frequent; dm, rp; AA/NAA.

S. flagellaris Willd. ex Sternb. subsp. flagellaris [Hirculus platysepalus (Trautv.) W.
A. Weber subsp. crandallii (Gand.) W. A. Weber]. Frequent; dm, mm, st, kr, ff,
rp; A/SRM.

S. odontoloma Piper [Micranthes odontoloma (Piper) Heller]. Occasional; wm, st, rv;
BM/WNA.

S. rhomboidea Greene var. rhomboidea [Micranthes rhomboidea (Greene) Small].
Very abundant; dm, mm, st, kr, ff, rp, rv, sn; A/WNA.

S. rivularis L. var. flexuosa (Sternb.) Engl. & Irmsch. [S. rivularis L.]. Rare; rp; AA/C.

Scrophulariaceae

Besseya alpina (A. Gray) Rydb. Frequent; dm, mm, ff, rp; A/SRM.

Castilleja miniata Dougl. ex Hook. Rare; dm, rp; BM/WNA.

C. occidentalis Torr.* Very abundant; dm, mm, wm, st, kr, ff, rp, rv, sn; A/RM.

C. rhexifolia Rydb. Occasional; mm, wm, st, rv; BM/WNA.

Chionophila jamesii Benth. Frequent; dm, mm, ff, rp, sn; A/SRM.

Mimulus guttatus DC. subsp. guttatus. Rare; rv; BM/NA.

Pedicularis bracteosa Benth. ex Hook. var. paysoniana (Penn.) Cronq. [P. bracteosa
Benth. subsp. paysoniana (Penn.) W. A. Weber]. Rare; st; M/RM.

P. groenlandica Retz. var. surrecta (Benth. ex Hook.) A. Gray [P. groenlandica Retz.].
Frequent; mm, wm, st, rp, rv, sn; AA/NA.

P. parryi A. Gray subsp. parryi. Occasional; dm, mm, st, kr; A/RM.

P. scopulorum A. Gray.* Occasional; mm, wm, st; A/RM.

Penstemon hallii A. Gray. Rare; dm, ff; A/CO.

P. harbourii A. Gray. Rare; rp; A/CO.

P. whippleanus A. Gray. Occasional; st, kr, rp; M/RM.

Veronica nutans Bong.* Frequent; dm, mm, wm, st, kr, rp, rv; AA/NA.

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE pips)|

Valerianaceae

Valeriana acutiloba Rydb. var. acutiloba [V. capitata Pall. ex Link subsp. acutiloba
(Rydb.) F. G. Mey.]. Rare; dm; AA/NAA.
V. edulis Nutt. ex Torr. & A. Gray var. edulis. Rare; dm, rp; BM/WNA.

Violaceae

Viola adunca Sm. var. bellidifolia (Greene) Harrington [V. labradorica Schrank].
Occasional; dm, mm, wm, st, kr, rp; BM/NA.

ANTHOPHYTA — MONOCOTYLEDONEAE
Cyperaceae

Carex albonigra Mack. [C. albo-nigra Mack. in Rydb.]. Frequent; dm, mm, wm, st,
kr, ff, rp; AA/WNA.
C. aquatilis Wahlenb. var. aquatilis [C. aquatilis Wahlenb. subsp. stans (Drejer)
Hulten]. Occasional; mm, wm, rv; AA/C.
C. arapahoensis Clokey.* Rare; dm, mm; A/SRM.
C. arctogena H. Sm. [C. capitata L. subsp. arctogena (H. Smith) Bocher]. Rare; mm;
BM/NA.
aurea Nutt. Rare; mm; BM/NA.
. bipartita Bellardi ex All. var. bipartita [C. lachenalii Schkuhr]. Rare; mm; AA/C.
brevipes W. Boott. Rare; dm, kr; BM/NA.
capillaris L. subsp. capillaris. Rare; mm; AA/C.
ebenea Rydb. Abundant; dm, mm, wm, st, kr, rp, rv, sn; A/RM.
elynoides Holm. Very abundant; dm, mm, st, kr, ff, rp, sn; A/WNA.
foenea Willd. Occasional; dm, mm, st, kr, ff; BM/NA.
haydeniana Olney. Occasional; dm, mm, wm, rp; A/WNA.
heteroneura W. Boott var. chalciolepis (Holm) F. J. Herm. [C. chalciolepis Holm].
Abundant; dm, mm, wm, st, kr, ff, rp, sn; A/WNA.
heteroneura W. Boott var. epapillosa (Mack.) F. J. Herm. [C. epapillosa Mack. in
Rydb.]. Rare; dm, mm; M/WNA.
. illota Bailey. Rare; mm, wm; A/WNA.
incurviformis Mack. [C. maritima Gunn.]. Rare; rp, sn; A/WNA.
misandra R. Br. Rare; mm, st; AA/C.
nardina Fries var. hepburnii (Boott) Kukenth. [C. nardina Fries subsp. hepburnii
(Boott) A. Love, D. Love & Kapoor]. Rare; mm, wm; A/SRM.
nelsonii Mack. Rare; mm; A/SRM.
nigricans C. A. Mey. Frequent; mm, wm, st, rp, rv; AA/NA.
norvegica Retz. Rare; dm, mm, rp; AA/NAE.
nova Bailey. Frequent; mm, wm, st, rp, rv; BM/WNA.
obtusata Lilj. Rare; mm, st; BM/C.
perglobosa Mack.* Rare; rp, sn; A/SRM.
phaeocephala Piper. Frequent; dm, mm, st, kr, rp, sn; A/WNA.
praeceptorum Mack. Rare; mm, wm; A/WNA.
pseudoscirpoidea Rydb. Frequent; dm, mm, st, kr, ff, rp; A/WNA.
pyrenaica Wahlenb. [C. praegracilis Boott]. Occasional; mm, rp, sn; A/C.
rupestris All. var. drummondii (Dewey) Bailey [C. rupestris All. subsp. drummon-
diana (Dewey) Holub]. Frequent; dm, mm, st, kr, ff, rp; A/RM.
saxatilis L. [C. saxatilis L. subsp. laxa (Trautv.) Kalela]. Rare; wm; AA/C.
scopulorum Holm var. scopulorum. Frequent; mm, wm, st, rp, rv; A/WNA.
vernacula Bailey. Occasional; mm, wm, rp, rv; A/WNA.
Eleocharis pauciflora (Lightf.) Link LE. quinquefolia (F. X. Hartm.) Schwartz]. Rare;
wm; A/C.
Kobresia bellardii (All.) Degl. ex Loisel. [K. myosuroides (Vill.) Fiori & Paoli]. Fre-
quent; dm, mm, st, kr, ff, rp; AA/C.

OGY Ok SG Gy Cy) OG Gy Gr Gye GY GY GiGGy “SG Gy GG a GG

229 MADRONO [Vol. 35

Juncaceae

Juncus albescens (Lange) Fern. Rare; mm; AA/NA.

J. balticus Willd. var. montanus Engelm. [J. arcticus Willd. subsp. ater (Rydb.) Hul-
ten]. Rare; wm; M/WNA.

J. biglumis L. Rare; mm, wm, rp; AA/C.

J. castaneus Sm. subsp. castaneus var. castaneus. Rare; mm; AA/C.

J. drummondii E. Mey. var. drummondii. Very abundant; dm, mm, wm, st, kr, rp,
rv, sn; A/WNA.

J. mertensianus Bong. Occasional; mm, wm, rv; A/NAA.

J. parryi Engelm. Rare; mm; M/WNA.

J. triglumis L. Rare; mm; AA/C.

Luzula parviflora (Ehrh.) Desv. [L. parviflora (Ehrh. ex Hoffm.) Lejeune]. Rare; mm,
rv; BM/C.

L. spicata (L.) DC. Very abundant; dm, mm, wn, st, kr, ff, rp, sn; A/RM.

L. subcapitata (Rydb.) Harr. Rare; mm, wm; A/CO.

Liliaceae

Lloydia serotina (L.) Sweet subsp. serotina. Abundant; dm, mm, wm, st, kr, ff, rp,
sn; AA/C.

Zigadenus elegans Pursh subsp. elegans [Anticlea elegans (Pursh) Rydb.]. Occasional;
dm, mm, st, rp; AA/NA.

Poaceae

Agropyron latiglume (Scribn. & Sm.) Rydb. [Elymus trachycaulus (Link) Gould ex
Shinners subsp. andinus (Scribn. & Sm.) A. Love & D. Loéve]. Occasional; dm,
st, rp; AA/NA.

A. scribneri Vasey [Elymus scribneri (Vasey) Jones]. Frequent; dm, kr, ff, rp; A/WNA.

Agrostis borealis Hartm. [A. mertensii Trin.]. Rare; dm, mm; AA/C.

A. humilis Vasey. Rare; mm, st; A/WNA.

A. variabilis Rydb. Rare; dm; A/WNA.

Calamagrostis canadensis (Michx.) Beauv. Rare; st; BM/NAA.

C. purpurascens R. Br. var. purpurascens. Occasional; dm, kr, ff, rp; AA/NAA.

Danthonia intermedia Vasey. Occasional; dm, st, rp; BM/NAA.

Deschampsia caespitosa (L.) Beauv. var. caespitosa [D. cespitosa (L.) P. Beauv. subsp.
alpicola (Rydb.) A. Léve, D. Love & Kapoor]. Abundant; dm, mm, wm, st, rp,
rv, sn; BM/C.

Festuca ovina L. var. brevifolia (R. Br.) Wats. LF. minutiflora Rydb.]. Very abundant;
dm, mm, wn, st, kr, ff, rp, sn; AA/C.

F. ovina L. var. rydbergii St. Yves LF. brachyphylla subsp. coloradensis Fred.]. Oc-
casional; dm, st, kr, ff, M/NA.

Helictotrichon mortonianum (Scribn.) Henr. Occasional; dm, mm, ff, rp; A/SRM.

Phleum alpinum L. [P. commutatum Gaud.]. Frequent; dm, mm, wm, st, rp, sn;
AA/C.

Poa alpina L. var. alpina. Abundant; dm, mm, wm, st, kr, rp, sn; AA/C.

P. cusickii Vasey. Rare; dm; M/RM.

P. epilis Scribn. [P. cusickii Vasey subsp. epilis (Scribn.) W. A. Weber]. Frequent;

dm, mm, wn, st, kr, rp; BM/WNA.

. fendleriana (Steud.) Vasey. Occasional; dm, mm, st; BM/NA.

. grayana Vasey [P. arctica R. Br.]. Occasional; dm, mm, st, rp; A/RM.

. interior Rydb. [P. nemoralis L. subsp. interior (Rydb.) W. A. Weber]. Rare; dm;

BM/NA.
reflexa Vasey & Scribn. ex Vasey. Occasional; mm, st, kr; A/WNA.
. rupicola Nash ex Rydb. [P. glauca M. Vahl subsp. rupicola (Nash) W. A. Weber].
Abundant; dm, mm, st, kr, ff, rp; A/WNA.

> a= Da a= |

1988] HARTMAN AND ROTTMAN: SAWATCH RANGE PISS

P. sandbergii Vasey [P. secunda C. Presl]. Rare; dm, mm; BM/NA.
Trisetum spicatum (L.) Richt. var. spicatum [T. spicatum (L.) Richt. subsp. congdonii
(Scribn. & Merr.) Hulten]. Very abundant; dm, mm, wm, st, kr, ff, rp, sn; AA/C.

ACKNOWLEDGMENT

Research conducted during the 1986 field season was supported by a Senior Faculty
Grant Award from the University of Colorado at Denver.

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224 MADRONO [Vol. 35

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Murpbock, J. R. 1951. Alpine plant succession near Mt. Emmon, Uinta Mountains,
Utah. M.S. thesis, Brigham Young Univ., Provo, UT.

NELSON, B. E. and R. L. HARTMAN. 1987. Rocky Mountain Herbarium checklist—
flora of Wyoming. Univ. of Wyoming, Laramie.

NORDHAGEN, R. 1955. Kobresieto-Dryadion in northern Scandinavia. Svensk. Bot.
Tidskr. 49:63-87.

O’ KANE, S., E. L. HARTMAN, and M. L. ROTTMAN. 1988. Noteworthy collections:
Colorado. Madrono 35:72.

Owen, H. E. 1976. Phenological development of herbaceous plants in relationship
to snowmelt date. Jn H. W. Steinhoff and J. D. Ives, eds., Ecological impacts of
snowpack augmentation in the San Juan Mountains, Colorado, p. 323-342. Final
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PorsILp, A. E. 1957. Illustrated flora of the Canadian Arctic Archipelago. Nat. Mus.
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and W. J. Copy. 1980. Vascular plants of continental Northwest Territories,
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Price, R. A., M. L. ROTTMAN, and E. L. HARTMAN. 1985. Noteworthy collections:
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County, Colorado. Misc. Field Stud. Map MF-1792-A.

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1988] HARTMAN AND ROTTMAN: SAWATCH RANGE Pip he)

WITTMANN, R. C., W. A. WEBER, and B. C. JOHNSTON. 1988. Flora of Colorado:
computer-generated catalog. Univ. Colorado, Boulder.

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New York.

(Received 22 Apr 1987; revision accepted 4 Feb 1988.)

ERRATUM

Part of the address of Mountain West Publishing was inadver-
tently omitted when the announcement below appeared in Madrofio
35(2):125. The announcement is repeated here with the complete
address.

ANNOUNCEMENT

New Publication

Dorn, RoBERT D. 1988. Vascular Plants of Wyoming, illustrated by
JANE L. Dorn. Mountain West Publishing, Cheyenne, WY. vi + 340
pp., paperbound. [Keys to 120 families, 650 genera, 2369 species, 39
subspecies, and 690 varieties; 93 new combinations, 1 new species,
4 new varieties, and 1 new name; section of taxonomic notes. Avail-
able postpaid for $13.00 from Mountain West Publishing, Box 1471,
Cheyenne, WY 82003.]

ANNOUNCEMENT

NEw PUBLICATION

PowELL, A. M. 1988. Trees and Shrubs of Trans-Pecos Texas, Big
Bend Natural History Association, P.O. Box 68, Big Bend National
Park, TX 79834, 536 pp., illus., ISBN 0-912001-14-3, $19.95 (paper-
bound). Includes keys, descriptions, distributions, and illustrations of
about 450 species of woody plants native to the Texas mountain and
desert region west of the Pecos River. (Also available, Chihuahuan
Desert Research Institute, P.O. Box 1334, Alpine, TX 79831, $17.95
to members, $19.95 to non-members.)

LATE WISCONSIN VEGETATION OF ROBBER’S ROOST IN
THE WESTERN MOJAVE DESERT, CALIFORNIA

NIALL MCCARTEN
Department of Biology, San Francisco State University,
1600 Holloway Ave., San Francisco, CA 94132

THOMAS R. VAN DEVENDER
Arizona-Sonora Desert Museum, 2021 N. Kinney Rd.,
Tucson, AZ 85743

ABSTRACT

A total of 22 plant taxa were identified from three packrat (Neotoma sp.) midden
assemblages radiocarbon dated at 12,870-13,330 yr B.P. at 1215 m elevation in
Robber’s Roost in the Scodie Mountains of Kern Co., California. The Late Wisconsin
vegetation was a pinyon-juniper woodland dominated by Pinus monophylla, Junip-
erus californica, and Ceanothus greggil. Excellent modern analogs are in Cushenberry
Canyon and similar areas on the desert slopes of the San Bernardino and San Gabriel
mountains. These woodlands are probably relicts of a pinyon-juniper-Joshua tree
woodland that was widespread across the southern Mojave Desert in the Late Wis-
consin.

Plant macrofossils from ancient packrat (Neotoma sp.) middens
have provided a detailed record of the invasion of the present deserts
of North America by woodland or forest trees (Van Devender and
Spaulding 1979). Woodlands dominated by Pinus monophylla, Ju-
niperus osteosperma, and Yucca brevifolia were present in many parts
of the present Mojave Desert in southern Nevada (Wells and Berger
1967, Leskinen 1975, Spaulding 1981) and southeastern California
(Mehringer 1965, Wells and Berger 1967, King 1976, Wells and
Woodcock 1985). Characteristic dominants of the modern Mojave
desertscrub such as Larrea divaricata and Ambrosia dumosa were
noticeably absent from these areas. In this paper, we present the first
analyses of Late Wisconsin packrat midden plant assemblages from
the base of the Sierra Nevada on the western edge of the Mojave
Desert and discuss their local and regional implications.

STUDY AREA

The Scodie Mountains are on the southeastern end of the Sierra
Nevada in Kern Co., California (Fig. 1). They range from 1160 m
elevation on the southeastern base to 2075 m on Pinyon Peak and
2170 mon Skinner Peak. The upper portion of the range is in Sequoia
National Forest. Walker Pass at 1905 m in Freeman Canyon marks
the northeastern edge of the range. The Scodie Mountains divide

MApDRONO, Vol. 35, No. 3, pp. 226-237, 1988

1988] McCARTEN & VAN DEVENDER: WISCONSIN VEGETATION 227,

A
40° Winnemucca

Lake

KERN CO.

StRETK oinyokern Searles
PFreeman uct. Lake

NEVADA

ie Tehachapi
36°-[&

Mts.
2 y Great Basin
uf Desert
37° z -
he y Ped.
- ‘M@ Specter: -
7. Range -.
36° -—36°
350 —35°
Ket eens Cerne - “Mts Whipple
oD ey Set : : A : ;
See, SON ONS eh, ee oY
Bernardino . Ag a Mis, a
Mts. (6 ee a :
Eh Jie Pes 7
fo rs eee
Woodland and Forest nat

Mojave Desert

@ San Diego
Sonoran Desert

\
115°

a I ,

Fic. 1. Map of southern California showing the location of Robber’s Roost in the

Mojave Desert and in relation to the Sierra Nevada and other fossil packrat midden
(solid triangles) and tar pit (open triangle) sites discussed in text. Mojave and Sonoran
deserts stippled after Brown and Lowe (1978) and Kiichler (1977).

228 MADRONO [Vol. 35

the internal drainage basins of the western Mojave Desert from the
Kern River drainage and the San Joaquin Valley. Robber’s Roost
is a series of rhyolitic plugs at 1190-1230 m elevation on the south-
eastern edge of the Scodie Mountains (35°35'45’N, 117°57'W), 4.3
km ene. of Freeman Junction. The packrat midden rockshelters are
at ca. 1215 m.

The rainshadows of the Scodie Mountains, the Sierra Nevada, and
the Transverse Ranges to the south are responsible for the general
aridity of the interior Mojave and Great Basin deserts. The clima-
tological means for Inyokern at 590 m, 14 km ne. of Robber’s Roost,
are 5.8°C for January, 29.2°C for July, and 106 mm/yr precipitation
with 7.5% in the summer (June-August; NOAA 1986). Estimates
of lapse rates for these climatic variables vary considerably. Major
(1977) reported temperature lapse rates of —0.45°C/100 m for Jan-
uary means and —0.60°C/100 m for July means for Owens Valley
north of Inyokern. Rowlands (1978) estimated —0.5°C to —0.6°C/
100 m for January means in the northern Mojave Desert in general.
A lapse rate of 9.4 mm/100 for annual precipitation was found for
the western Mojave Desert in general (Rowlands 1978), for Death
Valley from 1220 to 1830 m, and for the Kern River Canyon in the
southwestern Sierra Nevada (Major 1977). Using these lapse rates
the estimated climatic means for Robber’s Roost at 1215 m are
2.5°C for January, 25.3°C for July, and 168 mm/yr precipitation
with ca. 8% in the summer. If these lapse rates apply to the gradient
above Robber’s Roost, Skinner Peak at 2170 m would have —2.1°C
January mean, 19.7°C July mean, and 255 mm/yr precipitation.

Three vegetation zones occur from 1000 to 2100 m on the east
slope of the Scodie Mountains above Robber’s Roost (Fig. 2, Table
1). Above 1900 m, all slopes support a mixed woodland of Pinus
monophylla, P. sabiniana, Quercus chrysolepis, and occasional Yucca
brevifolia. Important shrubs include Artemisia tridentata, Eriogo-
num fasciculatum, and Purshia glandulosa. An open woodland
dominated by Pinus monophylla and P. sabiniana extends down to
1700 m on north slopes in Cow Heaven Canyon.

A mixed desertscrub with low shrubs grows from 1600 to 1900
m on hot, rocky east-facing slopes. Between 1300 and 1600 ma
sparse desertscrub includes low shrubs of Ericameria cooperi, E.
cuneata, Purshia glandulosa, and Eriastrum densifolium subsp. mo-
havense. On alluvial fans at these elevations a Yucca brevifolia des-
ertscrub community with associated bunchgrasses (e.g., E/ymus ely-
moides, Oryzopsis hymenoides, and Stipa speciosa) and shrubs (e.g.,
Ephedra californica, Ericameria arborescens, Hymenoclea salsola,
and Salvia dorrii) is well developed.

Below 1300 m is a desertscrub dominated by Larrea divaricata
and Ambrosia dumosa associated with other desert shrubs and
bunchgrasses (Table 1). The Robber’s Roost packrat midden site at

1988] McCARTEN & VAN DEVENDER: WISCONSIN VEGETATION 229

2200

Pinyon-
Oak
Woodland

1900

Joshua Tree

Desertscrub 1600

(Ww) U014DAe13

Robber's Roost
acer ee ee een bee eae ae 1300

(0)
Creosote Bush Desertscrub -
Kilometers

1000

Fic. 2. Generalized vegetation zonation along an east-west transect above Rob-
ber’s Roost (1215 m elev.) in the Scodie Mountains, Kern Co., California. Scale
approximate.

1215 m is in this community (Fig. 3). Ceratoides lanata, Encelia
virginensis, Hymenoclea salsola, Salazaria mexicana, and Sphaer-
alcea ambigua are scattered in the area. Chrysothamnus nauseosus,
Ericameria arborescens, and Ephedra californica are restricted to
relatively mesic microhabitats in the rhyolitic outcrops and along
shallow washes. A small (ca. 1 m) plant of Yucca brevifolia was
found at the edge of a small wash within 50 m of the midden rock-
shelter.

METHODS AND RESULTS

Packrat middens are hard, dark organic deposits that can be pre-
served in dry rockshelters for thousands of years. Middens are readily
disaggregated by soaking in water, screened through soil sieves, oven
dried, and hand sorted. The plant remains provide excellent samples
of the local floras on rocky slopes and are excellent for radiocarbon
dating. They contain well preserved plant remains collected within
ca. 30 m that can often be identified to species.

Three packrat midden samples from Robber’s Roost yielded a
total of 22 plant taxa with 9-21 per sample. The specimens were
identified by comparison with reference collections in the Laboratory
for Paleoenvironmental Studies at the University of Arizona (Table
2). This number of taxa is adequate to describe the Late Wisconsin
plant community at the site considering that 14—22 species were
observed in modern Scodie Mountains communities (Table 1). Rel-
ative abundance classes in the fossil assemblages and modern com-
munities were assigned from the most common (abundant = 5) to
single specimens (rare = 1). Ranks of the intermediate classes (very
common to uncommon, 4—2) varied depending on the total number
of specimens identified. Plant nomenclature mostly follows Munz
(1974); authorities for exceptions are included in Tables | and 2.

230 MADRONO

[Vol. 35

TABLE 1. PLANTS OBSERVED IN THE SCODIE MOUNTAINS, KERN Co., AND
CUSHENBERRY CANYON, SAN BERNARDINO CoO., CALIFORNIA. Authorities cited for
names differing from Munz (1974). Relative abundance: | = rare, 2 = uncommon,
3 = common, 4 = very common, 5 = abundant. * = identified in Robber’s Roost

packrat middens.

Species

Ambrosia dumosa
*Artemisia tridentata

Atriplex canescens
*Ceanothus greggli

Ceratoides lanata (Pursh)

J. T. Howell
Chrysothamnus nauseosus
Echinocereus engelmannii
Elymus elymoides (Raf.) Swezey
Encelia virginensis
Ephedra californica
Ephedra viridis
Ericameria cooperi (A. Gray) Hall

*Ericameria cuneata (A. Gray)

McClat.

Ericameria laricifolia (DC.)

Urbatsch & Wussow
Ericameria linearifolia (T. & G.)

Urbatsch & Wussow
Eriodictyon crassifolium

*Eriogonum fasciculatum
Fremontodendron californicum
Gutierrezia microcephala
Hymenoclea salsola

* Juniperus californica
Juniperus osteosperma
Larrea divaricata Cov.

*TLepidium fremontii

*TLupinus excubitus
Lycium andersonli
Machaeranthera tortifolia
Mirabilis bigelovii

*Opuntia basilaris

*Opuntia echinocarpa
Opuntia phaeacantha
Oryzopsis hymenoides

*Penstemon incertus

*Pinus monophylla
Pinus sabiniana
Poa secunda Presl.
Prunus andersonii
Prunus fasciculata

*Purshia glandulosa

Scodie Mountains
1300-1600 >1600

<1300

lwol | pl

le lunan!

Leong | kort oo Goa: |

wlwluw!l

Cushen-

N

NNNN WN

ASSO nt Stoel Ss Eel Al

1988] McCARTEN & VAN DEVENDER: WISCONSIN VEGETATION 231

TABLE |. CONTINUED.

Cushen-
berry
; ; Canyon
Sen De CUI te ise
<1300 1300-1600 >1600 1830
Species m m m m)
Quercus chrysolepis _ — 4 —
*Quercus turbinella _ _ — 2
Salazaria mexicana 3 3 _ 2
Salvia dorrii — _ 3 1
Sphaeralcea sp. 4 l — 1
Stipa speciosa 3 5 _ 2
Tetradymia spinosa _ 3 —_ —
*Yucca brevifolia 1 4 1 3
Total = 19 22 14 31

Radiocarbon dates on twigs and seeds of Juniperus californica
from the samples yielded ages of 12,870 + 400 (A-1762, RR#1D),
12,960 + 270 (A-1761, RR#2A), and 13,330 + 360 (A-1763, RR#1C)
yr B.P. (radiocarbon years before 1950). Attempts were not made

Fic. 3. View of packrat midden site at Robber’s Roost, Kern Co., California. A
pinyon-juniper woodland grew in the area from 12,870 to 13,330 yr B.P. in the Late
Wisconsin.

232 MADRONO [Vol. 35

to expand the three Late Wisconsin samples into a local chronology
because of the rarity of fossil middens in the area.

Although the Neotoma teeth from the samples were not identifi-
able to species, they had rounded lophs characteristic of N. albigula,
N. fuscipes, and N. lepida rather than the prismatic teeth of N. cinerea
or N. mexicana. Neotoma albigula inhabits various communities in
summer rainfall areas from southeasternmost California east to cen-
tral Texas and the Mexican Plateau; it was probably not the builder
of the Robber’s Roost middens. More likely candidates are N. lepida
of the Mojave Desert and nearby woodlands, or N. fuscipes of Sierran
and coastal chaparral.

DISCUSSION AND CONCLUSIONS

The plant assemblages from the Robber’s Roost packrat middens
record a Late Wisconsin pinyon-juniper woodland dominated by
Juniperus californica and Pinus monophylla in association with Ce-
anothus greggii, Purshia glandulosa, Artemisia tridentata, Eriogo-
num fasciculatum, Yucca brevifolia, Quercus turbinella, and Erica-
meria cuneata (Table 2). Twelve species (54.5%) identified from the
samples no longer occur in the Robber’s Roost area. Preliminary
electrophoretic studies of phenolics including flavonoids suggest that
the fossil junipers represent J. californica rather than J. osteosperma.
The modern woodland above 1900 m is not a good analog of the
Robber’s Roost paleowoodland because J. californica, C. greggii,
and Q. turbinella are rare or absent and P. sabiniana and Q. chry-
solepis are associated with P. monophylla. The nearest populations
of J. californica and C. greggii are 9 km nw. of Robber’s Roost in
Walker Pass at 1750 m elevation. Farther west, the vegetation begins
a gradual transition to chaparral in South Fork Valley north of the
Piute Mountains. The nearest populations of Quercus turbinella are
in Kern Co. at Erskine Canyon in the northwestern Piute Mountains
(ca. 40 km w.) and in the northeastern Tehachapi Mountains (47
km ssw.; Twisselman 1967; Fig. 1).

Estimates of climatic parameters for Robber’s Roost in the Late
Wisconsin using lapse rates discussed above for a 685 m minimum
lowering of Pinus monophylla in the Scodie Mountains are —0.75°C
mean for January, 21.3°C for July, and 218 mm/yr precipitation
with little in summer. As mentioned above the Scodie Mountains
woodland is a poor analog, suggesting the modern climates may also
be different. Different lapse rates in the Late Wisconsin would yield
different estimates. The paleoclimatic estimates are within the cli-
matic limits estimated for modern pinyon-juniper woodlands in the
mountains of the eastern Mojave Desert by Thorne et al. (1981):
e.g., January mean less than 4°C, July mean less than 25.5°C, and
annual precipitation greater than 250 mm/yr with 165-185 mm in

233

1988] McCARTEN & VAN DEVENDER: WISCONSIN VEGETATION

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234 MADRONO [Vol. 35

the cool seasons (September to May). This would represent an in-
crease in annual and winter precipitation at Robber’s Roost of about
36%.

An excellent modern analog to the Robber’s Roost paleocom-
munity is on the north slopes of the San Bernardino Mountains
above the Lucerne Valley (Fig. 1) in San Bernardino Co. An
extensive pinyon-juniper woodland in Cushenberry Canyon (1830
m) is dominated by Pinus monophylla, Juniperus osteosperma, and
Yucca brevifolia and also includes Atriplex canescens, Artemisia tri-
dentata, Ceanothus greggii, Chrysothamnus nauseosus, Ephedra vir-
idis, Ericameria linearifolia, Eriogonum fasciculatum, Opuntia ba-
silaris, O. echinocarpa, O. phaeacantha, Purshia glandulosa, Quercus
turbinella, and Salvia dorrii. Encelia virginensis and Lepidium fre-
montii were seen just below at 1585 m. Juniperus californica is
present in the lower portion of the woodland. This area is 500-700
m higher than Robber’s Roost and 170 km to the southeast. Similar
woodlands can be found in other areas along the desert slopes of the
San Bernardino and San Gabriel mountains.

The Robber’s Roost paleowoodland is also similar to modern
woodlands isolated in the mountains of the eastern Mojave Desert
(Thorne et al. 1981). Late Wisconsin pinyon-juniper woodlands with
Pinus monophylla have been recorded in the Mojave Desert from
Ord Mountain (King 1976), the Turtle Mountains (Wells and Berger
1967), and Clark Mountain (Mehringer and Ferguson 1969) in Cal-
ifornia and the Newberry Mountains (Leskinen 1975), Spring Range
(Van Devender and Spaulding 1979), Sheep Range (Spaulding 1981),
and Specter Range (Spaulding 1985) in Nevada. Xeric woodland
assemblages dominated by Juniperus californica with low levels of
Pinus monophylla, Yucca brevifolia, and Y. whipplei have been found
as low as 510 m in the Whipple Mountains just above the Colorado
River in eastern San Bernardino Co. (Van Devender and Spaulding
1979, Van Devender et al. 1987). These relict distributions of mod-
ern woodlands and their fossil records suggest that pinyon-juniper
woodland was widespread across the central and southern Mojave
Desert in the Late Wisconsin (Wells 1986, Betancourt 1986).

Full-glacial packrat midden records from King’s Canyon in the
Sierra Nevada (170 km nnw. of Robber’s Roost) record the expan-
sion of Pinus monophylla even farther to the west (Cole 1983).
Fossils of P. monophylla and J. californica (including J. osteosperma)
were reported from the Late Pleistocene McKittrick asphalt deposits
at 320 m elevation in the San Joaquin Valley 170 km wsw. of
Robber’s Roost (Mason 1944). The plant fossils were associated with
extinct mammals characteristic of the Rancholabrean Land Mam-
mal Age including Bison, Camelops, Equus, Hemiauchenia, Mam-
mut, and Mammuthus (Harris 1985). A middle Wisconsin radiocar-
bon age (38,000 + 2500 yr B.P.) was obtained on plant materials

1988] McCARTEN & VAN DEVENDER: WISCONSIN VEGETATION 235

from the site (Kurtén and Anderson 1980). These records suggest
that many of the disjunct populations of Great Basin plants did not
disperse into the Sierra Nevada during the “warm, dry Xerothermic”’
dispersal (=middle Holocene; Taylor 1976, Axelrod 1981) but in
glacial climates in the Wisconsin if not earlier.

The Robber’s Roost paleoflora is especially interesting because of
its location near the eastern base of the Sierra Nevada. A number
of trees that potentially could have dispersed southward along the
main axis or eastward from the mesic west slopes of the Sierra or
northwest from the Transverse Ranges were not found; e.g., Abies
concolor, A. magnifica, Calocedrus decurrens, Juniperus occidentalis,
Pinus coulteri, P. lambertiana, P. jeffreyi, P. ponderosa, Pseudotsuga
macrocarpa, Quercus agrifolia, O. douglasii, QO. lobata, or Sequoia-
dendron giganteum. Thompson et al. (1985) reported an eastward
expansion of J. occidentalis from the Sierra Nevada into the Win-
nemuca Lake Basin of Nevada in the Great Basin. Plants indicative
of the Adenostoma fasciculatum-dominated chaparral of the Los
Angeles Basin have not been found in middens.

Packrat middens from the Mojave and Sonoran deserts have yield-
ed records of some plants now found in the chaparral and woodlands
of the interior slopes of the mountains of southern California with
more extensive eastern distributions in the Late Wisconsin and early
Holocene. Most of them presently have disjunct populations in
woodland, chaparral, or desertscrub communities in Arizona or So-
nora (e.g., Arctostaphylos pungens, Ceanothus greggii, Eriogonum
fasciculatum, Juniperus californica, Nolina bigelovii, Quercus chry-
solepis, O. dunnii, O. turbinella, Yucca brevifolia, and Y. whipplei).
Most Late Wisconsin dispersals of woodland, chaparral, or desert-
scrub species into the Mojave Desert recorded by packrat midden
fossils were from the north or northeast.

Plant remains in packrat middens from Robber’s Roost and other
areas in the Mojave Desert provide insight into the historical com-
ponents in modern communities. The modern vegetation of Rob-
ber’s Roost is a mixture of species that were present in the Late
Wisconsin woodland and warm desert species that dispersed into
the area in the Holocene. The regional vegetation of the western
Mojave Desert reflects similar historical processes with pinyon-ju-
niper woodlands contracted to isolated mountaintops or the lower
elevational zones of larger mountain ranges. In the Late Wisconsin
a Mojave desertscrub with Larrea divaricata and Yucca brevifolia
was present below 310 m in the Picacho Peak area in Imperial Co..,
California (Van Devender et al. 1985, Cole 1986). The area is just
north of Yuma in the modern Lower Colorado River Valley sub-
division of the Sonoran Desert. The Larrea divaricata—Ambrosia
dumosa desertscrub of much of the Mojave Desert developed in the
last 11,000 years as these species and their associates migrated from

236 MADRONO [Vol. 35

their glacial refugium in the Lower Colorado River Valley and the
Gran Desierto surrounding the head of the Gulf of California in
Sonora and Baja California. Great Basin elements of the paleo-
woodlands survive today in the region as elevational zones domi-
nated by Artemisia tridentata and in mixed desertscrub communities
in the western Mojave Desert (e.g., Artemisia spinescens, Atriplex
confertifolia, and Ceratoides lanata).

ACKNOWLEDGMENTS

We thank Austin Long for the radiocarbon dates, Don Koehler for identification
of the Purshia glandulosa fossils, Ron Lanner for anatomical analysis of the fossil
Pinus monophylla needles, and Roxanne Bittman, Jim Yates, Alan Davis, Andy
Sanders, and Dave Morafka for their help in the field. The careful editing of Wayne
R. Ferren, David J. Keil, and two anonymous reviewers greatly improved the paper.
This research was supported in part by National Science Foundation grant DEB 75-
13944 to Paul S. Martin. Dana Dorner and Helen Wilson drafted the diagrams. Jean
Morgan typed the manuscript.

LITERATURE CITED

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BETANCOURT, J. L. 1986. Paleoecology of pinyon-juniper woodlands: summary. Jn
R. L. Everett, compiler, Proceedings—pinyon-juniper conference, p. 129-139.
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Brown, D. E.andC. H. Lowe. 1978. Biotic communities of the Southwest. U.S.D.A.
Forest Serv. Gen. Tech. Rept. RM-41. Rocky Mountain Forest and Range Exp.
Sta., Fort Collins, CO.

Cote, K. L. 1983. Late Pleistocene vegetation of King’s Canyon, Sierra Nevada,
California. Quarternary Res. 19:117-129.

1986. The Lower Colorado Valley: a Pleistocene desert. Quarternary Res.
25:392-400.

Harris, A. H. 1985. Late Pleistocene vertebrate paleoecology of the West. Univ.
Texas Press, Austin.

Kina, T. J. 1976. Late Pleistocene—Early Holocene history of coniferous woodlands
in the Lucerne Valley region, Mohave Desert, California. Great Basin Naturalist,
36:227-238.

KUCHLER, A. W. 1977. Natural vegetation of California. Jn M. G. Barbour and J.
Major, eds., Terrestrial vegetation of California, map. John Wiley and Sons, New
York.

KurTEN, B. and E. ANDERSON. 1980. Pleistocene mammals of North America.
Columbia Univ. Press, New York.

LESKINEN, P. H. 1975. Occurrence of oaks in late Pleistocene vegetation in the
Mohave Desert of Nevada. Madrono 23:234-235.

Mayor, J. 1977. California climate in relation to vegetation. Jn M. G. Barbour and
J. Major, eds., Terrestrial vegetation of California, p. 11-74. John Wiley and
Sons, New York.

Mason, H. L. 1944. A Pleistocene flora from the McKittrick asphalt deposits of
California. Proc. Calif. Acad. Sci. 25:221-233.

MEHRINGER, P. J. 1965. Late Pleistocene vegetation in the Mohave Desert of south-
ern Nevada. J. Arizona Acad. Sci. 3:172-188.

and C. W. FERGUSON. 1969. Pluvial occurrence of bristlecone pine, Pinus

aristata, in a Mohave Desert mountain range. J. Arizona Acad. Sci. 5:284—-292.

1988] McCARTEN & VAN DEVENDER: WISCONSIN VEGETATION 2a1

Muwnz, P. A. 1974. A flora of southern California. Univ. California Press, Berkeley.

NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION. 1986. Climatological data—
California: annual summary. 90:1-57.

ROWLANDs, P.G. 1978. The vegetation dynamics of the Joshua tree (Yucca brevifolia
Engelm.) in the southwestern United States of America. Ph.D. dissertation, Univ.
California, Riverside.

SPAULDING, W. G. 1981. The late Quarternary vegetation of a southern Nevada
mountain range. Ph.D. dissertation, Univ. Arizona, Tucson.

1985. Vegetation and climates of the last 45,000 years of the vicinity of
the Nevada Test Site, south-central Nevada. U.S. Geol. Survey Prof. Pap. 1329:
1-83.

TAYLoR, D. M. 1976. Disjunction of Great Basin plants in the northern Sierra
Nevada. Madrono 23:301-364.

THOMPSON, R. S., L. BENSON, and E. M. HATTori. 1985. A revised chronology for
the Last Pleistocene lake cycle in the central Lahontan Basin. Quarternary Res.
25:1-9.

THORNE, R. F., B. A. PRIGGE, and J. HENRICKSON. 1981. A flora of the higher range
and the Kelso Dunes of the eastern Mojave Desert in California. Aliso 10:71-
186.

TWISSELMAN, E. C. 1967. A flora of Kern County, California. Univ. San Francisco,
San Francisco, CA.

VAN DEVENDER, T. R., P. S. MARTIN, R. S. THompson, K. L. Coe, A. J. T. JULL,
A. Lona, L. J. Too.in, and D. J. DONAHUE. 1985. Fossil packrat middens and
the tandem accelerator mass spectrometer. Nature 317:610-613.

and W. G. SPAULDING. 1979. Development of vegetation and climate in the

southwestern United States. Science 204:701-710.

, R. S. THOMPSON, and J. L. BETANCOURT. 1987. Vegetation history of the
Southwest: the nature and timing of the Late Wisconsin-Holocene transition. Jn
W. F. Ruddiman and H. E. Wright, Jr., eds., North America and adjacent oceans
during the last deglaciation, p. 323-352. Geol. Soc. America, Boulder, CO.

WELLS, P. V. 1986. Systematics and distribution of pinyons in the late Quarternary.
InR. L. Everett, compiler, Proceedings— pinyon-juniper conference, p. 104-108.
Intermountain Res. Station, U.S.D.A. Forest Serv., Ogden, UT.

and R. BERGER. 1967. Late Pleistocene history of coniferous woodland in

the Mohave Desert. Science 155:1640-1647.

and D. Woopcock. 1985. Full-glacial vegetation of Death Valley, California:

juniper woodland opening to Yucca semidesert. Madrono 32:1 1-23.

(Received 17 Apr 1987; revision accepted 11 Feb 1988.)

ANNOUNCEMENT

NEw PUBLICATION
BARBOUR, M. G. and W. D. BILLINGS, eds. 1988. North American

terrestrial vegetation. Cambridge Univ. Press, New York. 434 pp.
ISBN 0-521-26198-8 (clothbound), $49.50. [Vegetation of North
America including tropical areas. Thirteen chapters, each separately
authored and with its own references. Illustrated with black and white
photographs, maps, and diagrams. ]

THE ABUNDANCE OF PLANTS BEARING EXTRAFLORAL
NECTARIES IN COLORADO AND MOJAVE DESERT
COMMUNITIES OF SOUTHERN CALIFORNIA

ROBERT W. PEMBERTON
Agricultural Research Service,
United States Department of Agriculture,
Rangeland Insect Laboratory, Montana State University,
Bozeman 59717

ABSTRACT

Measurements of the cover and frequency of EFN-bearing plants in seven warm
desert communities in California revealed some of the highest levels of abundance
of EFN-bearing plants that have been recorded for the temperate zone. The desert
wash communities of both deserts had the highest covers (28 and 24%) and fre-
quencies (0.27, 0.27) of EFN-bearing plants, whereas the sand dune communities
had the lowest levels of abundance of EFN-bearing plants with covers of 2 and 0.0%
and frequencies of 0.01 and 0.0. Colorado Desert communities had higher covers,
frequencies, and numbers of EFN-bearing plants than Mojave Desert communities.
The EFN antiherbivore defense system is predicted to be also common in other
warm-dry communities of the world because those environments have an abundance
of ants and plant groups, such as mimosoid legumes and cacti, known to have many
EFN-bearing species. The EFN defense system may be particularly well suited to
plants growing in warm-dry zones.

Extrafloral nectaries are nectar-secreting glands occurring most
commonly on the vegetative parts of plants, but also at other sites
such as developing fruit and the external parts of flowers. Instead
of attracting pollinators, extrafloral nectaries (EFN’s) have been shown
to promote mutualistic interactions between plants and the insects,
especially ants, that visit the EFN’s. The insect participants gain
sugars, amino acids, and water from the EFN’s and benefit the plants
by reducing the damage caused by the plant’s herbivores (Janzen
1966, Bentley 1977a, Tilman 1978, Pickett and Clark 1979, Keeler
1980, Schemske 1980). At least 73 angiosperm families with almost
1000 species, and a few ferns have EFN’s (Keeler 1979b). Plants
with EFN’s occur in most parts of the world (Zimmermann 1932,
Schnell et al. 1963) and appear to be most common in the tropics
(Bentley 1977b).

The abundance of EFN plants in plant communities has been
examined in Costa Rica (Bentley 1976), Jamaica (Keeler 1979a),
Nebraska (Keeler 1979b), Northern California (Keeler 1981a), Ar-
izona (Keeler 1981b), and Hawaii (Keeler 1985). The cover of EFN
plants has been found to be highest in the communities in Costa
Rica (40-80%) and in the aspen (Populus tremuloides Michx.) dom-

MAbDRONO, Vol. 35, No. 3, pp. 238-246, 1988

1988] PEMBERTON: EXTRAFLORAL NECTARIES IN DESERT PLANTS 239

inated mountain forests of Arizona (39%). The lowest covers of EFN
plants were in the Nebraska communities (0.0-8%) and in northern
California where no EFN plants were found in the four communities
sampled.

Zimmermann (1932) thought that xerophytes, as a rule, lacked
EFN’s and for this reason believed the dry floras of California to
have practically no EFN plants. Except for Helianthella californica
Gray (Keeler 1981a), no native EFN plants have been reported from
California (Buckley 1982). After observing EFN’s on cacti growing
in California’s deserts, I suspected that plants with EFN’s were more
abundant in California than was previously known. A greater abun-
dance of plants with EFN’s in California’s deserts was also suggested
by the abundance of ants (Wheeler and Wheeler 1973), which has
been correlated with the abundance of EFN plants in other com-
munities (Bentley 1976). The object of this study was to learn how
abundant EFN plants might be in some California desert commu-
nities.

METHODS

The abundance of EFN plants was determined by measuring their
frequency and cover in four Colorado Desert and three Mojave
Desert communities in southern California during March 1986. Fre-
quency was determined by scoring the presence or absence of EFN
plants at 1 m intervals along three 100 m transects through each
community. Cover was determined by measuring to the nearest cm
the linear distance occupied by EFN plants along each of the tran-
sects. Detection of EFN plants was made by direct observation of
secreting EFN’s on the plants, which was often aided by the presence
of ants and other insects tending the nectaries. Locating EFN plants
was made easier by examining species (and their relatives) previously
reported to bear EFN’s. The percentages of the floras with EFN
plants in the areas studied was made by identifying the species,
known to me, to have EFN in “Plants of Deep Canyon” (Zabriskie
1980), the area of the Colorado Desert transects, and in “‘A Flora
of the Higher Ranges and Kelso Dunes of the Eastern Mojave Desert
in California” (Thorne et al. 1981), the area of the Mojave Desert
transects.

Colorado Desert Transects

The Colorado Desert transects were taken at the Phillip L. Boyd
Deep Canyon Desert Research Center of the University of Califor-
nia. This area lies on the northeast slopes of the Santa Rosa Moun-
tains and the adjacent southwest slopes of Coachella Valley, Riv-
erside Co., California between 116°%-117°W and 33°-34°N. The

240 MADRONO [Vol. 35

Colorado Desert is the northwestern subsection of the Sonoran Des-
ert, and is lower in altitude and more arboreal in character than the
Mojave. Creosote bush scrub occupies the largest areas in both the
Colorado and Mojave deserts (Munz and Keck 1959).

1. Creosote bush scrub—on rocky alluvial fan, west of the Chan-
nel of Deep Canyon Creek, 300 m elev. Common plants: Encelia
farinosa A. Gray (Compositae), Fouquieria splendens Engelm. (Fou-
quieriaceae), Larrea divaricata Cav. (Zygophylaceae), and Opuntia
spp. (Cactaceae).

2. Desert wash—sand and pebbles, Deep Canyon creek wash, 265
m elev. Common plants: Acacia greggii A. Gray and Cercidium
floridum Benth. (Leguminosae), Chilopsis linearis (Cav.) Sweet (Big-
noniaceae), and Hyptis emoryi Torr. (Labiatae).

3. Yucca-galleta grass—sand and rock hillside, adjacent to Hwy.
74 overlooking Deep Canyon, 820 m elev. Common plants: Agave
deserti Engelm. and Yucca schidigera Roezl. ex Ortgies (Agavaceae),
Fouquieria splendens and Hilaria rigida (Thurb.) Benth. ex Scribn.
(Gramineae).

4. Sand dunes—Coachella Valley floor east of Thousand Palms,
40 m elev. Common plants: Atriplex spp. and Salsola australis R.
Br. (Chenopodiaceae), Larrea divaricata and Prosopis juliflora (Sw.)
DC. (Leguminosae).

Mojave Desert Transects

The Mojave Desert transects were located on the northern side of
the Granite Mountains and at Kelso Dunes in San Bernardino Co.,
California at approximately 116°W and 35°N. The Mojave Desert
is intermediate between the cold-temperate Great Basin Desert and
the subtropical Colorado Desert (Turner 1982) and has a lower
diversity of perennial plants than the Colorado Desert (Vasek and
Barbour 1977). The average annual rainfall for the Mojave study
areas is less than 200 mm (estimated from Thorne et al. 1981) and
90-150 mm for the Colorado Desert sites (estimated from I. P.
Tinginan, unpublished booklet, ‘““Natural History of Deep Canyon’’).
The average annual temperature for the Mojave sites is estimated
to be around 26°C (Thorne et al. 1981) and higher for Deep Canyon,
where it rarely freezes.

5. Sand dunes—eastern slope of Kelso Dunes, 900-1000 m elev.
Common plants: Astragalus sp. (Leguminosae), Croton californicus
Muell. Arg. (Euphorbiaceae), and various grasses.

6. Creosote bush scrub—sand and rock, alluvial fan, northern
slope of the Granite Mts., 1250 m elev. Common plants: Coleogyne
ramosissima Torr. (Rosaceae), Eriogonum spp. (Polygonaceae), Lar-
rea divaricata and Salazaria mexicana Torr. (Labiatae), and Yucca
schidigera.

7. Desert wash—boulders and sand, northern slope of Granite

1988] PEMBERTON: EXTRAFLORAL NECTARIES IN DESERT PLANTS 241

Mts., 1350 m elev. Common plants: Acacia greggii, Ephedra sp.
(Ephedraceae), Jsomeris arborea Nutt. (Capparidaceae), Prunus fas-
ciculata (Torr.) Gray (Rosaceae), and Rhus trilobata Nutt. ex T. &
G. (Anacardiaceae).

RESULTS

The plants observed to bear EFN’s are listed in Table 1. All 11
species had active secreting EFN’s in either the Colorado or Mojave
Desert study areas, or both. The largest number of the species found
to possess EFN’s were cacti. The four Opuntia species had EFN’s
located on the areoles of the newly formed pads, flower buds, and
flowers. The EFN’s of Ferocactus were tubercules located above the
areoles on the inside of the ring of flowers on top of the cacti. The
EFN’s of all cacti, except O. acanthocarpa Engelm. and Bigel., were
tended by ants. The EFN’s of Chilopsis were located on the leaf
blades and were variable in their occurrence both within and between
trees. The EFN’s of ocotillo (Fouquieria splendens) were located on
the flower buds, where relatively large 5 mm diameter drops of sweet
tasting viscous nectar accumulated. Acacia greggii had small EFN’s
located on the leaves along the primary rachis between the branching
secondary rachis bearing the leaflets. Prosopis juliflora bore EFN’s
on the rachis between the leaflets and also on the leaf petioles. Ants
were tending its EFN’s. The Prunus species had EFN’s at the bases
of their leaf blades. Prunus fasciculata had large numbers of small
parasitic wasps (mainly Chalcidoidea) visiting its EFN’s. In addition
to the hymenoptera (ants and wasps) visiting the EFN’s, lady beetles
(Hippodamia convergens Guerin-Meneville) were observed on the
EFN’s of Opuntia echinocarpa Engelm. & Bigel. and small uniden-
tified flies were observed feeding on the nectaries of Chilopsis.

The abundance of EFN plants in the different communities of the
Colorado and Mojave deserts is shown in Table 2. The desert wash
communities of both deserts had the highest covers (27.74%, 23.89%)
and frequencies (0.277, 0.266) of EFN plants. The sand dune com-
munities, with 1.36% and 0.0% covers, and 0.016 and 0.0 frequen-
cies, had the lowest abundance of species with EFN’s. The creosote
bush scrub communities were intermediate in both deserts (cover
6.58%, 0.07%; frequency 0.120, 0.003). The communities of the
Colorado Desert had, on average, a higher EFN plant cover (x =
9.8%) and frequency (KX = 0.118) than those of the Mojave cover (x
= 8.0%) and frequency (KX = 0.090). The x number of EFN plant
species was also higher in the Colorado communities with 3 vs. 1.66
species for the Mojave communities.

The percentages of species with EFN’s in the native flora were
0.95% (1/105) for Kelso Dunes and 2.61% (10/382) for the Granite
Mountains of the Mojave, and 3.20% (18/562) for Deep Canyon of
the Colorado.

[Vol. 35

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1988] PEMBERTON: EXTRAFLORAL NECTARIES IN DESERT PLANTS 243

TABLE 2. ABUNDANCE OF PLANTS WITH EXTRAFLORAL NECTARIES IN THE COLORADO
AND MOJAVE DESERTS. Combined data for three 100 meter transects per community.

Frequency % cover Number of
Community location n/300 points n/300 meters EFN species
Colorado Desert
1. Creosote bush scrub 0.120 6.58 6
Deep Canyon 36/300 19.7/300
2. Desert wash 0.277 27.74 2
Deep Canyon 83/300 83.2/300
3. Yucca agave galeta grass 0.060 3.65 3
Deep Canyon 18/300 10.9/300
4. Sand dunes 0.016 1.36 1
Cocachella Valley 5/300 4.1/300
Mojave Desert
5. Sand dunes 0.000 0.00 0
Kelso 0/300 0/300
6. Creosote bush scrub 0.003 0.07 1
Granite Mt. 1/300 0.2/300
7. Desert wash 0.266 23.89 4
Granite Mts. 80/300 71.7/300
DISCUSSION

The detection of cacti previously unreported to bear EFN was
predicted by their occurrence in other cacti (Lloyd 1908, Pickett and
Clark 1979). Similarly, many Prunus (Dorsey and Weiss 1920) and
Acacia species (Delpino 1886) are known to bear EFN’s. Chilopsis
was suspected to have EFN’s because most members of the Big-
noniaceae have them (Elias 1983). Less expected were the EFN’s in
ocotillo (Fouquieria splendens) since few members of the Fouquieria-
ceae have them (Elias 1983).

Although the abundance of EFN plants in some of the desert
communities of this study was quite high (24 and 28% cover), none
approached the high levels (40-80%) measured in three dry tropical
forest habitats in Costa Rica (Bentley 1976). More similar were the
Jamaican lowland (Keeler 1979a) and Hawaiian Acacia koa Grey
(Keeler 1985) communities with covers by EFN plants of 28 and
21%. Most temperate communities that have been measured have
much lower abundances of EFN plants than found in this study.
The exceptions are Arizona aspen forest (39%) and an Arizona So-
noran Desert community, found to have a cover of 22% (Keeler
1981b). The cover for that Arizona desert community is similar to
the cover (24 and 28%) of desert washes measured in this study.

The only published accounts of the frequency of EFN species in
floras are for Hawaii and Nebraska. Keeler (1979b, 1985) found
2.5% of the indigenous species in Nebraska to have EFN’s and 1.5%
of Hawaiian natives in Hawaii Volcano National Park to bear EFN’s.

244 MADRONO [Vol. 35

The figures for this study (0.95, 2.61, and 3.20%) are similar and
noteworthy because most of the desert community EFN plant covers
are much greater than in Nebraska and most of the Hawaiian com-
munities sampled. These differences are explained by the presence
of EFN’s in species that are both abundant and of large stature, such
as Chilopsis, Acacia, Prosopis, Fouquieria, and Prunus. High plant
covers of EFN plants have been measured in other communities
having few or single or large statured EFN plants, such as Acacia
koa in Hawaii and Populus tremuloides in Arizona.

The desert plants in this study comprised some of the highest EFN
plant covers that have been measured in the temperate zone com-
munities. I predict that EFN plants are also common in many of
the world’s warm deserts and other hot dry biomes such as savanna
and tropical scrub.

A number of the taxa found to bear EFN’s in this study have
dryland relatives that are known to bear EFN’s. Delpino (1886)
found that 172 of the 258 Acacia and 11 of the 15 Prosopis species
he examined bore EFN’s. Broughton (1981) found EFN’s in all 42
species of Australian Acacia that she studied, including those from
the interior desert areas which had formerly been thought to lack
EFN’s. EFN’s also occur in Acacia species that are native to Central
America (Janzen 1966), the Caribbean and South America (Keeler
pers. comm.), Africa (Ross and Gordon-Gray 1966), and India
(Bhattacharyya and Maheshwari 1971). EFN’s are also common in
species of other mimosoid genera such as Mimosa, Albizia, and
Leucaena (Bhattacharyya and Maheshwari 1971) that are prominent
members of the world’s warm-dry floras. The prevalence of EFN’s
in species of cacti in the genera Opuntia and Echinocactus (Lloyd
1908), Ferocactus (Blom and Clark 1980, Ruffner and Clark 1986),
and others native to both North America and South America, further
support the probability of an abundance of EFN plants in the New
World warm deserts and tropical scrub communities.

The general richness and abundance of ants in the world’s desert
and warm-dry communities also supports the prediction of high
levels of abundance of EFN-bearing plants in those regions, as they
did in the deserts of southern California.

The use of a water based antiherbivore defense system may appear
to be an inappropriate strategy for arid land plants; however, growth
and reproduction in warm desert plants usually occur only in periods
of increased water availability. Since secretion in EFN’s is most
active in expanding foliage and reproductive structures (Bentley
1977b, and this study), EFN defense is used during periods of water
availability. The greatest abundance of EFN-bearing plants in this
study was desert washes, areas where plants have greater access to
water.

Protection of new growth and reproductive tissues may be rela-

1988] PEMBERTON: EXTRAFLORAL NECTARIES IN DESERT PLANTS 245

tively more important in desert plants, since the possibilities of
regrowth of tissues lost to herbivores is restricted by limited water.
The EFN defense may be particularly well suited to these arid land
plants because the vulnerable tissues are protected as they are being
produced. EFN defenses also have the advantage of being effective
against both specialist and generalist insect herbivores, which is
usually not the case for specific chemical defenses (Beattie 1985).

ACKNOWLEDGMENTS

I wish to thank the University of California for permission to work at Deep Canyon
Desert Research Center; K. Kang (Berkeley, CA) and J. B. Pemberton (Pleasanton,
CA) for technical assistance; and K. H. Keeler (University of Nebraska) and H. G.
Baker (University of California, Berkeley) for helpful reviews of the manuscript.

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BENTLEY, B. L. 1976. Plants bearing extrafloral nectaries and the associated ant
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1977a. The protective function of ants visiting the extrafloral nectaries of

Bixa orellana L. (Bixaceae). J. Ecol. 65:27-38.

. 1977b. Extrafloral nectaries and protection by pugnacious bodyguards. Ann.
Rev. Ecol. Syst. 8:407-427.

BHATTACHARYYA, B. and J. K. MAHESHWARI. 1971. Studies on extrafloral nectaries
of the Leguminales. Proc. Indian National Acad. Sci. 37B:11-30.

BLom, P. E. and W. H. CLARK. 1980. Observations of ants (Hymenoptera: For-
micidae) visiting extrafloral nectaries of the barrel cactus Ferocactus gracilis
Gates (Cactaceae), in Baja California, Mexico. Southw. Naturalist 25:181-196.

BROUGHTON, V. H. 1981. Extrafloral nectaries of some Australian phyllodineous
acacias. Austral. J. Bot. 29:653-664.

BUCKLEY, R. C. 1982. Ant-plant interactions: a world review. /n R. C. Buckley,
ed., Ant-plant interactions in Australia, p. 111-141. W. Junk, The Hague, Neth-
erlands.

DELPINO, F. 1886. Funzione mirmecofile nel regno vegetale. Mem. Reale. Accad.
Sci Ist. Bologna. Ser. 4. VII. 21:5-392; VII:602-659; X:115-147.

Dorsey, M. J. and F. Weiss. 1920. Petiolar glands in the plum. Bot. Gaz. (Craw-
fordsville) 69:39 1—406.

Euias, T.S. 1983. Extrafloral nectaries: their structure and distribution. /n B. Bentley
and T. Elias, eds., The biology of nectaries, p. 174-203. Columbia Univ. Press,
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JANZEN, D.H. 1966. Coevolution of mutualism between ants and acacias in Central
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KEELER, K. H. 1979a. Frequency of extrafloral nectaries and ants at two elevations
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246 MADRONO [Vol. 35

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(Received 9 Dec 1987; revision accepted 9 May 1988.)

ANNOUNCEMENT

PUBLICATION AVAILABLE

Biology of the California Islands — Proceedings of the First Symposium

(R. W. Philbrick, ed.). 1967. 363 pp. Hard cover, $3.75. [We have a
large number of copies of the first Symposium in mint condition that
we would like to get into the hands of interested persons.] Available
from the Santa Barbara Botanic Garden, 1212 Mission Canyon Road,
Santa Barbara, CA 93105. Price includes shipping and California sales
tax.

GENERIC RELATIONSHIPS AND TAXONOMY OF
ACAMPTOPAPPUS (COMPOSITAE: ASTEREAE)

MEREDITH A. LANE
Department of Environmental, Population and Organismic Biology,
University of Colorado, Boulder 80309-0334

ABSTRACT

Acamptopappus comprises A. sphaerocephalus var. sphaerocephalus and var. hir-
tellus, and A. shockleyi, taxa of the southwestern deserts of the United States. Char-
acteristics shared by these taxa include extremely long-villous achenes, a pappus of
long, somewhat erose scales with lanceolate to spatulate apices, deeply alveolate
receptacles, short, funnelform disk corollas with deep sinuses and reflexed lobes, very
broad phyllaries with very broad, scarious margins, and nearly globose capitula.
Chromosome numbers of all taxa are n = 9. The probable closest relatives of this
outlying genus are to be found among larger genera of Astereae having x = 9, par-
ticularly the Chrysothamnus— Ericameria— Macronema alliance.

Acamptopappus (A. Gray) A. Gray comprises three taxa of the
Mojave and Sonoran deserts of the southwestern United States (Figs.
1, 2). The taxa are distinctive in appearance, with very light green
foliage, whitish stems, nearly globose capitula, phyllaries nearly as
broad as long, and the most villous achenes of all North American
Astereae. No treatment encompassing all three taxa has previously
been published, except in floras. This paper circumscribes these taxa,
and discusses the possible relationships of Acamptopappus with oth-
er genera of Astereae.

TAXONOMIC HISTORY

Gray (1849) named sect. Acamptopappus of Haplopappus Cass.
to accommodate H. sphaerocephalus Harvey & A. Gray in A. Gray
(1849). It was based on a specimen collected by Thomas Coulter in
1832 that had been forwarded to Gray between 1846 and 1848 by
W. H. Harvey, Coulter’s successor as curator of the herbarium of
Trinity College, Dublin (Coville 1895). Later, Gray (1873) accorded
generic status to Acamptopappus, a move with which Hall (1928, p.
365) concurred. In 1882, Gray described A. shockleyi. Jones (1898)
established A. microcephalus, which was placed in synonymy with
Ericameria cooperi (A. Gray) H. M. Hall by Blake (1929), when he
named A. sphaerocephalus var. hirtellus. No additions have been
made to the genus as a result of the present study, although questions
about the types are clarified in comments following the descriptions
of taxa.

MADRONO, Vol. 35, No. 3, pp. 247-265, 1988

248 MADRONO [Vol. 35

Fic. 1. Distribution of Acamptopappus shockleyi. Each symbol may represent one
or more collections. Type locality indicated by star.

METHODS

Standard herbarium techniques were used to study 1182 speci-
mens of the three taxa, including types, borrowed from or observed
at ARIZ, ASU, CAS, COLO, F, GH, JEPS, K, LL, MO, NY, POM,
RM, RSA, SD, TCD, TEX, UC, US, and UTC. For comparison of
generic features, specimens of Chrysothamnus, Eastwoodia, Erica-
meria, Petradoria, Stenotus, and Vanclevea belonging to COLO, LL,
RM, and TEX were studied. Acamptopappus taxa were observed in
the field during trips taken in 1984 and 1986.

Achenes, style-branches, corollas and pappus members used for
scanning electron microscopy were carefully removed from herbar-
lum specimens (COLO, LL, RM, or TEX). Achenes and pappus
were mounted on stubs with double-stick tape; style-branches and
corollas were rehydrated by soaking in Wetter’s solution (Wetter
1983), rinsed, and then mounted with double-stick tape. Stubs were
sputter-coated with ca. 400 nm gold, and observed on an AMR
1O00A at 20 kV.

DISCUSSION

Morphology. The two speices of Acamptopappus are very similar
to one another (Table 1, Fig. 3), except that A. sphaerocephalus plants

1988] LANE: ACAMPTOPAPPUS 249

ape °
“no, © epse
; e if. e e

oe M
® fe) e ¢ ee
aye é Conger 0
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@ SPHAEROCEPHALU . °°
ca e
= e = e 4
O HIRTELLUS

oe —

COULTER'S ROUTE ee

Fic. 2. Distribution of Acamptopappus sphaerocephalus var. hirtellus and var.
sphaerocephalus. Each symbol may represent one or more collections. Type locality
for var. hirtellus indicated by star. Route of Thomas Coulter (Coville 1895) indicated
by double line; see comments under description of var. sphaerocephalus for expla-
nation.

are usually slightly larger (to 4 dm) than those of A. shockleyi (to
3.3 dm), are more highly ramified, have narrower leaves, and have
smaller, more numerous and occasionally clustered capitula that are
eradiate and have fewer disk florets.

The pappus of 4. shockleyi usually consists of 17-30 moderately
erose, white scales. Ray floret pappus apices usually are lanceolate
to acute, whereas those of the disk florets are more spatulate (Fig.
4a); the pappus elements of A. sphaerocephalus disk florets are spat-
ulate.

Both species have narrowly triangular-lanceolate disk style-branch
appendages that are acute and flattened on the adaxial face, and have
collecting hairs that are of moderate length (Fig. 4a). Disk corolla
epidermes (Fig. 5a) of the two species of Acamptopappus are iden-
tical. Ray-corolla epidermis of A. shockleyi is shown in Fig. 5b.

Achenes of Acamptopappus species are actually cylindric, but ap-
pear obconic in outline because they are covered with the longest
and densest zwillingshaares that I have seen in any taxon of Astereae.
Anderson and Weberg (1974) noted that there are “long, isotropic,
non-glandular (shag) hairs’? present with the “‘anisotropic duplex
hairs” on Acamptopappus achenes. However, I found that all tri-
chomes are of the normal “‘anisotropic duplex’ type exemplified by
those of Stenotus acaulis (Fig. 5c), except that some are twisted and
contorted (Fig. 5d). The latter are usually concealed by a layer of

~

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[Vol. 35

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252 MADRONO [Vol. 35

+ SY diy!
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Fic. 3. Line drawings of habits (left) and capitula (right): a. Acamptopappus
shockleyi (Henrickson 9584, TEX); b. A. sphaerocephalus var. sphaerocephalus (M.
E. Jones s.n., TEX); c. Amphipappus fremontii var. fremontii (Cronquist 10649, TEX).
Scale bars = 2 cm.

straight trichomes, much as the downy underhairs of animal fur are
covered by long, straight guard hairs. The adaptive significance of
this arangement is obscure, but may protect the achene from des-
iccation or overheating, or enhance dispersal (see discussion of ecol-
ogy, below).

The two varieties of A. sphaerocephalus differ only in that stems
and leaves of var. sphaerocephalus are typically glabrous, or some

1988] LANE: ACAMPTOPAPPUS pf)

\
| i in d i
" / VW
) | y
a b f
&

Fic. 4. Line drawings (traced from scanning electron micrographs) of pappus-
member apices (above; r = ray, d = disk) and disk style-branch appendages (below):
a. Acamptopappus shockleyi (style-branch appendage: Henrickson 9584, TEX; pappus
member: Clokey 8157, TEX); b. Amphipappus fremontii var. fremontii (Cronquist
10649, TEX); c. Eastwoodia elegans (Eastwood and Howell 5791, TEX); d. Vanclevea
stylosa (Shultz and Shultz 7393, COLO); e. Ericameria cooperi subsp. cooperi(Gierisch
and Esplin 3460, COLO); f. Stenotus acaulis (Weber and Salamun 12568, COLO).
Scale bar = 1 mm.

plants may have a very few, scattered trichomes on the leaf margins,
whereas herbage of var. hirtellus is scabro-hirtellous (Blake 1929).
This is a minor difference, although there is a geographic component
to the variation (Fig. 2). Some populations in Los Angeles and San
Bernardino cos., California, which lie in the area of overlap of the
ranges of the varieties, have individuals with and without the ves-
titure. There are no intermediate individuals, either in these pop-
ulations or elsewhere. This situation is parallel to that for the two
varieties of Amphipappus fremontii (Porter 1943, Lane unpubl. data),
in which var. fremontii is glabrous, and var. spinosus is scabro-
hirtellous.

Ecology. Acamptopappus is well-adapted to the arid climate of the
Mojave Desert. The leaves are drought-deciduous, and the white
stems reflect sunlight. In favorable years, the plants are in leaf by
February, have flowered by late March, and are in fruit by late April
to early June (Ackerman et al. 1980, pers. obs.). In unfavorable
years, plants often do not bloom at all, or the capitula wither before
achenes are matured or even set.

Achenes are dispersed by wind and/or rain, being blown “‘tum-

254 MADRONO [Vol. 35

Fic. 5.
trichomes: a. disk corolla epidermis of Acamptopappus sphaerocephalus var. sphaer-
ocephalus (M. E. Jones s.n., TEX); b. ray corolla epidermis of A. shockleyi (Henrickson
9584, TEX); c. achene trichomes of Stenotus acaulis (Weber and Salamun 12568,
COLO); d. achene trichomes of A. shockleyi (Clokey 8157, TEX). Scale bar in a, b
= 25 um; inc, d = 100 um.

bleweed style’’ or carried in runoff across the soil surface. The achenes
are moved either individually or clustered by the interdigitation of
their pappus members and achene trichomes. This dispersal syn-
drome, together with a seedling mortality of only 28% over eight
years of study (Wallace and Romney 1980) may account for the
finding of Wallace and co-workers (1980d) that A. shockleyiis usually
found growing in clumps of several individuals, and of Wallace and
Romney (1980) that the species is a pioneer that initiates new “fertile
islands” in bare desert areas.

Acamptopappus shockleyi has been one among several subjects of
a number of ecological and ecophysiological studies conducted on
shrubby taxa of the Atomic Energy Commission (Nuclear Regulatory
Commission) test site in southern Nevada (unfortunately, compa-
rable data are not available for A. sphaerocephalus). Wallace et al.
(1980d) found 28-101 plants per hectare, although the relative den-

1988] LANE: ACAMPTOPAPPUS 259

sity of the species was less than 1% (El-Ghonemy et al. 1980c). The
mean stem weight per plant in these studies ranged from 35.2 to
68.0 g (Bamberg et al. 1980, El-Ghonemy et al. 1980a, Wallace et
al. 1980a). Plants are not particularly salt-tolerant (Romney and
Wallace 1980). Seventy-five percent of the root mass usually lies
within 20 cm of the surface (Wallace et al. 1980b) of soils that have
relatively low cation exchange capacity, low exchangeable sodium,
a medium amount of moisture retention, and low potassium content
(El-Ghonemy et al. 1980b).

Phytochemistry. Acamptopappus produces sesquiterpenoids (C-15
compounds) and labdane diterpenoids (C-20 compounds). Eight
compounds of the latter type that are new to science were isolated
and characterized from 4. sphaerocephalus by Jolad et al. (1988);
these compounds were also found in 4. shockleyi. The terpenoids
of Acamptopappus (Jolad et al. 1988) are similar to those produced
by other genera of the tribe Astereae, such as Chrysothamnus and
Ericameria (B. Timmermann pers. comm.).

Generic relationships. Distinguishing features of the two Acamp-
topappus species, Amphipappus, Eastwoodia, and Vanclevea (all
monotypic) are presented in Table 1 and Fig. 4. I include Amphi-
Dappus in this discussion of the relationships of Acamptopappus
because the two genera are placed near one another in floras (e.g.,
Keck in Munz 1959, Kearney and Peebles 1969), Vanclevea for the
same reason and because of Steyermark’s (1937) suggestion, and
Eastwoodia because of its gross morphological similarity to these
other genera. However, I suggest, based on the evidence presented
below, that each of these genera is derived independently from an
ancestral complex that also gave rise to Chrysothamnus, Ericameria,
and Macronema (Haplopappus sects. Ericameria and Macronema
sensu Hall), which in turn have been considered related by Hall
(Hall and Clements 1923, Hall 1928), and other authors.

Steyermark (1937) placed Acamptopappus near Xanthisma and
distinguished it from Grindelia in his discussion of the relationships
of the latter genus. I agree that Acamptapappus and Grindelia are
very dissimilar, but Acamptopappus also differs from Xanthisma in
many features, such as disk-corolla shape and epidermis pattern,
style-branch appendage shape, leaf shape and vestiture, habit, hab-
itat, and distribution. These three genera also differ in base chro-
mosome number. All Acamptopappus taxa have n = 9 (Raven et al.
1960, Keil and Pinkava 1976, Pinkava and Keil 1977, Kovanda
1978, Schaak et al. 1982), Grindelia has x = 6, and Xanthisma has
x = 4 (or possibly x = 5, see Semple 1976). All of the genera listed
in Table 1, and Chrysothamnus, Ericameria, and Macronema, have
x = 9. Although chromosome number alone is insufficient evidence
of relationship or lack thereof, the consistent correlation of mor-

256 MADRONO [Vol. 35

phological characters with base chromosome number that is found
in the Astereae supports its use as a character in a discussion such
as this one.

Acamptopappus and Amphipappus are found in the Mojave and
Sonoran deserts (Figs. 1, 2, and Porter 1943), and Vanclevea in the
southeastern extension of the Great Basin desert in Utah and Arizona
(Anderson and Weberg 1974). Eastwoodia occurs along the xeric
western and southern rim of the San Joaquin Valley (Brandegee
1894) on the eastern slopes of the South Coast Ranges and northern
slopes of the Transverse Ranges of California (Lane in Hickman in
prep.). All share certain features such as low, shrubby habit, greenish-
or yellowish-white new growth, and white, sometimes varnished
stems that become gray and shreddy with age.

Among the genera detailed in Table 1, Acamptopappus is most
similar to Amphipappus (Fig. 3). Shared characters include those of
the foliage, phyllary shape, color, and texture, disk-corolla shape,
and the tortuous nature of the achene trichomes (Fig. 5d). These
genera differ in capitulum size and shape, floret number, receptacle
features, sexuality of the disk florets, style-branch appendage shape,
pappus type, and degree of achene pubescence.

Steyermark (1937) saw a resemblance between Eastwoodia and
Acamptopappus on the basis of receptacular projections between the
florets. However, those of Eastwoodia are true paleae, whereas those
of Acamptopappus are merely the extended rims of the alveolae.
Both Eastwoodia and Acamptopappus have hermaphroditic disk flo-
rets and prominent ray florets in at least one taxon, but these are
plesiomorphic conditions and therefore do not necessarily indicate
relationship. Differences are found in the leaves, capitulum shape,
phyllary shape and texture, disk-corolla shape, style-branch ap-
pendages and pappus type (Fig. 4), and degree of achene pubescence
(Table 1).

Vanclevea differs from Acamptopappus in characters of the foliage,
phyllaries, disk-corolla shape, style-branch appendages and pappus
(Fig. 4), and degree of achene pubescence (Table 1). Very few sim-
ilarities with Acamptopappus, except for those listed above for all
four genera, can be found.

I suggest that although the four genera probably share a common
heritage, they are independently derived because there are so few
synapomorphies among them. Because each genus is so distinctive,
indications of the nature of their common heritage must be sought
in a large grouping of extant taxa. Of the Astereae genera that might
be considered, the genera having x = 9 and comprising shrubby taxa
of the southwestern deserts include Chrysothamnus (sensu Anderson
1984), Ericameria (sensu Urbatsch and Wussow 1979), and Mac-
ronema (=Haplopappus sect. Macronema sensu Hall 1928). Petra-
doria (sensu Anderson 1963) and Stenotus (Haplopappus sect. Steno-

1988] LANE: ACAMPTOPAPPUS ye

tus sensu Hall 1928), although herbaceous, are nonetheless perennial
with woody caudices and share general habitat preference and dis-
tribution with the other members of this alliance.

Many members of this group have greenish-white young stems
that become white and then gray with shredding bark in age, as do
all the genera of Table 1. Some members have the light green, non-
resinous leaves of Acamptopappus and Amphipappus, and others
have the dark green, resinous leaves of Eastwoodia and Vanclevea.
A complete intergradation between the short, broadly funnelform
disk-corolla shape of Acamptopappus and Amphipappus and the
tubular-funnelform one of Eastwoodia and Vanclevea is found in
this alliance as well.

The disk style-branch appendages of the Chrysothamnus—Erica-
meria—Macronema alliance, represented in Fig. 4e by that of Eri-
cameria cooperi, are generally lanceolate-acute, although some species
have more lanceolate ones. Those of Acamptopappus (Fig. 4a) are
similar, whereas Eastwoodia (Fig. 4c) has broader ones, and Van-
clevea (Fig. 4d) has the largest style-branch appendages of any taxon
of Astereae that I have studied. Amphipappus (Fig. 4b) has lost
female fertility in its disk florets, and this is reflected in absence of
stigmatic lines on its style branches, which also have obtuse ap-
pendages. A reasonable interpretation of these data is that the style
branches of Amphipappus, Eastwoodia, and Vanclevea are each, but
separately, apomorphic with respect to those of the Chrysothamnus—
Ericameria—Macronema alliance.

The pappus of members of this phylad is generally composed of
barbellate bristles that are more or less round in cross section as are
those of Stenotus acaulis (Fig. 4f), although Ericameria cooperi (Fig.
4e) and other members have flattened bristles. Porter (1943) sug-
gested that the ray pappus of Amphipappus (Fig. 4b) is formed by
fusion of bristles like those of the disk pappus (Fig. 4b) into scales.
The same process, extended over evolutionary time, may account
for the origin of the pappus scales of Acamptopappus, Eastwoodia,
and Vanclevea (Fig. 4a, c, d) from those of ancestor(s) with broad,
flat bristles such as those found in Ericameria cooperi (Fig. 4e) today.
The pappus of Amphipappus (Fig. 4b) is always tortuous, but this
may result from compression within the tightly imbricate involucre,
much as the pappus of some florets within an Acamptopappus head
may become twisted because it is compressed by surrounding florets
during development. Thus, the similarity between these two genera
with respect to tortuous pappus may be a parallelism rather than a
synapomorphy.

Corolla epidermis patterns have been found to be useful characters
at the generic and infrageneric levels in the Astereae (Lane 1982,
1985). Acamptopappus, Amphipappus, Eastwoodia, Vanclevea,
Chrysothamnus, Macronema, and Petradoria have the same disk

258 MADRONO [Vol. 35

corolla epidermis pattern (Fig. 5a). Acamptopappus, Amphipappus,
and Macronema also share the same ray corolla epidermis pattern
(Fig. 5b), whereas Petradoria, Ericameria, and Stenotus have a dif-
ferent one (Lane unpubl. data). It is difficult to polarize the epidermal
pattern characters, but it would seem that one or the other of these
two groups of three genera is synapomorphic in this respect.

It would be desirable to have a cladistic analysis of the relation-
ships of the genera discussed above. However, to present a cladogram
at this time would be premature because such an analysis requires
that all taxa belonging to a lineage be included in the analysis. The
scope of the current study has not ensured that this is the case;
neither has it yet been possible to determine an appropriate out-
group.

Future studies leading to thorough phylogenetic analysis have been
designed to test the hypothesis that Acamptopappus has been derived
from the ancestral complex that gave rise more directly to the Chry-
sothamnus—Ericameria—Macronema phylad. The single taxon of this
group to which Acamptopappus is most similar is E.. parrasana. With
this species, Acamptopappus shares capitulum shape, phyllary fea-
tures, and reflexing involucres in addition to the overall similarities
of the genera given above. The similar but separate derivation of
Amphipappus, Eastwoodia, and Vanclevea is another hypothesis to
be tested in future. The distinctiveness of Acamptopappus and each
of these genera may be accounted for by elapsed time since sepa-
ration of the lineages, and the strong selection pressures of their
desert habitats.

TAXONOMIC TREATMENT

Acamptopappus (A. Gray) A. Gray. Proc. Amer. Acad. Arts 8:634.
1873.—Aplopappus Cass. sect. Acamptopappus A. Gray, Mem.
Amer. Acad. Arts (ser. 2) 4:76 [Pl. Fendler. 76]. 1849.—TyYPE:
Acamptopappus sphaerocephalus (Harv. & A. Gray in A. Gray)
A. Gray.

Shrubs to 4 dm high, scraggly or rounded (Fig. 3); taproots woody,
vertical or usually laterally spreading; stems decumbent, divergent
or erect, striate; young stems greenish-white or -yellow, becoming
white; old stems gray, usually with shredding bark; leaves pale green
to light gray-green, borne singly, rarely in axillary fascicles below,
spreading-ascendent to appressed-erect, linear to lanceolate or nar-
rowly obovate or spatulate, 1-nervate, entire, glabrous or scabro-
hirtellous at margins, generally minutely spinulose at apices, gla-
brous or scabro-hirtellous on both surfaces; capitula borne singly or
occasionally in cymose clusters; buds expanding rapidly just prior
to anthesis; involucres broadly campanulate-hemispheric to nearly
spheric; phyllaries in 2—3 series, broadly ovate to ovate-elliptic, char-
taceous, brittle, cream-yellow at bases, green at apices, with broad,

1988] LANE: ACAMPTOPAPPUS 29

scarious, erose margins, all distinctly reflexing at maturity to release
achenes; receptacle deeply alveolate, with projections between florets
but not chaffy; heads radiate or eradiate, corollas yellow; disk corollas
broadly funnelform, sinuses deep, lobes spreading to reflexed; style-
branch-appendages narrowly triangular-lanceolate (Fig. 4a), some-
what exceeding the stigmatic portion; achenes obconic, extremely
densely long-villous; trichomes white, bronze, rufous, or brownish,
outer straight, inner contorted or tortuous (Fig. 5d); pappus of
l-seriate, white, scarcely erose scales with acute-lanceolate to nar-
rowly spatulate apices (Fig. 4a), slightly exceeding achenes; base
chromosome number: x = 9. Flowering (Mar—)Apr—May(—Jun) (Ack-
erman et al. 1980, Lane hoc. loc.).

KEY TO TAXA OF Acamptopappus

1. Heads radiate, involucres campanulate to hemispheric .......
PAR ne SORT ee ee ee ae ee A. shockleyi

1. Heads eradiate, involucres hemispheric to globose.
2. Stems and leaves scabro-hirtellous ........................
NE hee ee ee A. sphaerocephalus var. hirtellus
2. Stems and leaves glabrous, or only leaf margins scabro-hirtel-
LOU Sar Oe ee ye A. sphaerocephalus var. sphaerocephalus

Acamptopappus shockleyi A. Gray, Proc. Amer. Acad. Arts 17:208.
1882.—Type: USA, Nevada, Esmeralda Co., Candelaria, 1881,
Shockley 34 (GH!).

Stems decumbent to ascendent, (1.5—)2—3(—3.3) dm, usually spi-
nescent with age, surfaces usually scabro-hirtellous; leaves spread-
ing-ascendent, narrowly obovate to narrowly spatulate, (0.7—)1—1.6
(—2) cm long, (2—)3—4(—5) mm wide, scabro-hirtellous; capitula borne
singly; involucres campanulate to hemispheric, 7—11(—13) mm high,
(10-)13-19 mm wide; phyllaries 13-18(—23), (3.5-—)5—9(-11) mm
long, (1.8—)2—4(-6) mm wide; ray florets 5-14, corollas (3.5—)6—-
17(-19.5) mm long, (1.5-)2.5—6.5 mm wide; disk florets 30-80, co-
rollas (2.3—)3.2—5(—5.5) mm high; achenes (1—)1.5—3.5(-4.7) mm long,
(0.4—-)0.8-1.8(—2.9) mm wide; pappus scales (15—)18—30(-38), less
spatulate in rays than disks, (2.7—)3-—4.5(—5) mm high.

Distribution and habitat. Mojave Desert areas of southeastern Cal-
ifornia and southern Nevada (Fig. 1); 500-2000 m. Mesas, slopes,
ravines, and washes in Larrea and Yucca brevifolia communities, in
association with Atriplex, Amphipappus, Artemisia, Lycium, Grayia,
Encelia, Psilostrophe, Thamnosma, Hymenoclea, Eurotia, and/or
Hilaria.

Comments. Gray’s (1882) description of A. shockleyi was based
on a specimen numbered Shockley 34, collected in 1881. Other

260 MADRONO [Vol. 35

specimens with this number are at CAS and UC (but these are from
Rhyolite or Tonopah and dated 1883 or 1907), and another at NY
(dated 1886). Clearly these specimens cannot be considered isotypes,
though the NY specimen presumably is a topotype. Isotypic status
for a sheet at RSA is doubtful because it gives only ‘““Apr—May”’ for
collection date, even though it bears the correct locality and number.
Given Shockley’s re-use of the number 34, I doubt that it was col-
lected in 1881.

Representative specimens. USA, California, Inyo Co., Payson Can-
yon, White Mts., 14 Jun 1932, Duran 3295 (CAS, NY, GH, F, MO,
NY, RM, RSA, UC, US, UTC). San Bernardino Co., 4 mi e. of
Horse Spring, Mojave Desert, Kensington Mts., 15 May 1935, Wolf
6848 (CAS, COLO, MO, NY, RSA, UC). Nevada, Clark Co., Old
Kyle Canyon fan, 11 May 1938, Clokey 8157 (ARIZ, CAS, F, GH,
K, MO, NY, RM, RSA, SD, TEX, UC, UTC). Esmeralda Co., Can-
delaria, 22 Jun 1882, Jones 3895 (CAS, F, MO, NY, POM, RSA,
NY, UC, UTC). Lincoln Co., 11 mis. of Alamo, 6 Apr 1934,
Maguire et al. 5033 (GH, MO, RM, UC, UTC). Mineral Co., near
Mina, 5 Jun 1906, Heller 8368 (CAS, F, GH, MO, NY, US). Nye
Co., Smokey Valley, 9 Jun 1945, Maguire and Holmgren 25362
(ARIZ, GH, NY, US, UTC).

Acamptopappus sphaerocephalus (Harvey & A. Gray in A. Gray) A.
Gray.

Stems usually many, much-branched, ascendent to erect, (1.5—)2-
3.5(—4) dm, with surfaces scabro-hirtellous or glabrous; leaves as-
cendent-appressed, linear to narrowly oblanceolate, 0.5—2(—2.8) cm
long, (1-)1.5—3(—4) mm wide, scabro-hirtellous or glabrous; capitula
very numerous, borne singly or in clusters; involucres hemispheric
to spheric, 4-7 mm high, (1.5—)6-11 mm wide; phyllaries 11-18
20), (2.5—)3.2—5.5(—6.5) mm long, (1.5—)1.9-3(—3.6) mm wide; heads
eradiate; disk florets (13—-)14—24(—27), corollas (2.1—)2.5—4.3(—4.7)
mm high; achenes (1.2—)1.7—3.3(—3.7) mm high, (0.6—)0.8—1.9(—2.2)
mm wide; pappus bristles (15-)17—26(—28), (1.7—)2.1-3.7(-4.4) mm
high, apices narrowly spatulate.

Distribution and habitat. Mojave and Sonoran desert areas of
southern California, southern Nevada and Utah, and south-central
Arizona (Fig. 2); 5-2000 m. Gravelly, rocky soils on slopes and flat
areas in grasslands, deserts, and Juniperus woodlands; in association
with Larrea, Yucca, Viguiera, Eriogonum, Salsola, Ambrosia, Ar-
temisia, Chrysothamnus, Coleogyne, Ephedra, Canotia, Hymeno-
clea, Cercidium, Fouquieria, Carnegia, Opuntia, and/or Ferocactus.

Comments. Some populations in the area of overlap of the two
varieties of this species (Fig. 2) are mixed with respect to the dis-
tinguishing pubescence character. Representative collections from

1988] LANE: ACAMPTOPAPPUS 261

this area include: California, Los Angeles Co., Lancaster, Jun 1902,
Elmer 3621 (CAS, F, GH, K, MO, NY, RSA, US). San Bernardino
Co., Mojave Desert, May 1882, Parish and Parish 139 (CAS, F,
GH, MO, NY, SD, UC, US); Cima, Mojave Desert, Jun 1915, Bran-
degee s.n. (F, GH, MO, NY, RM, UC); Mojave Desert, Spring 1927,
Hutchinson s.n. (LL, TEX).

Populations of A. sphaerocephalus may very rarely contain indi-
viduals with vestigial ray florets (D. Keil pers. comm., M. Lane pers.
obs.). This condition might result from one of two phenomena: 1)
hybridization with the radiate A. shockleyi or 2) partial expression
of ray-floret genes that were suppressed during the evolution of 4.
sphaerocephalus from a radiate ancestor. Either explanation is
plausible, but neither is more strongly supported than the other by
evidence available at this time. There are infrequent cases of co-
occurrence of the two Acamptopappus species where their ranges
overlap (Figs. 1, 2) and hybridization may occur, although I have
seen no specimens that I would suspect to be of hybrid origin. Ves-
tigial rays occasionally occur in eradiate taxa of other genera of
Astereae that I have studied (for example, /socoma), indicating that
loss of rays is an apomorphic condition but that their suppression
is not absolute.

Acamptopappus sphaerocephalus (Harv. & A. Gray in A. Gray) var.
hirtellus S. F. Blake, J. Wash. Acad. Sci. 19:270. 1929.—TyYpPE:
USA, California, Inyo Co., near Lone Pine, 7 Jun 1891, Coville
and Funston 890 (US)).

Stems (1.5—)2.5—3.3(—3.8) dm high, scabro-hirtellous; leaves
(0.6-)0.8-1.3 cm long, 1.5—3(—4) mm wide, scabro-hirtellous; in-
volucres 6—7 mm high, (1.5—)8—11 mm wide; phyllaries (14—)15—-
18(—20), (2.8-)3.7-5.4(—5.7) mm long, (1.5)2—2.8(—3.1) mm wide;
disk florets (13-)16-24(-27), corollas 2.8-3.6(—4.5) mm _ high;
achenes 1.7—-3.2(-3.7) mm long, 0.7—1.7(—2.2) mm wide; pappus
bristles 15-24, (2.1-)2.8—3.3(—3.7) mm high. Gravelly soils in deserts
and Juniperus woodlands; 5—1600 m.

Representative specimens. USA, Arizona, Mohave Co., Fort Mo-
have, Apr 1884, Lemmon s.n. (UC, US). California, Inyo Co., Al-
abama Hills, 3 mi w. of Lone Pine, 23 May 1958, Rose 58061 (CAS,
COLO, GH, JEPS, NY, RSA, US). Kern Co., near Searles, 28 May
1932, Duran 3224 (CAS, COLO, F, GH, MO, NY, RM, RSA, UC,
UTC, US). Los Angeles Co., near Lancaster, 11 Jun 1906, Hall and
Chandler 7388 (ARIZ, F, K, MO, NY, RM, RSA). San Bernardino
Co., Mojave River district, Apr [or 23 May or 1 Jun?] 1876, Palmer
219 (F, MO, NY, US). Nevada, Clark Co., 15 mie. of Glendale, 19
May 1933, Maguire and Blood 4487 (MO, RM, RSA, UC, UTC).
Lincoln Co., Moapa, 12 May 1905, Kennedy 1077 (F).

262 MADRONO [Vol. 35

Acamptopappus sphaerocephalus (Harv. & A. Gray in A. Gray) A.
Gray var. sphaerocephalus, Proc. Amer. Acad. Arts 8:634.
1873.—Haplopappus sphaerocephalus Harvey & A. Gray in A.
Gray, Mem. Amer. Acad. Arts, ser. 2, 4:76 [Pl. Fendler. 76].
1849.—Type: USA, California, [San Diego Co.?], [without lo-
cality], [1832], Coulter s.n. exsic. no. 281 (GH!; isotypes K!,
TCD)).

Stems (1.8—)2.4—3.5(—3.8) dm high, glabrous; leaves (0.5—)1—2(—2.8)
cm long, (1—)1.5—3(—4) mm wide, glabrous; involucres 4—7 mm high,
(1.5-)6-10 mm wide; phyllaries 1 1—18(—19), (2.5—)3.2—5.3(—6.4) mm
long, 1.9-3(—3.6) mm wide; disk florets (13—)14—22(—26), corollas
(2.1-)2.5—4.3(-4.7) mm high; achenes (1.2—)1.7—3.2(-3.7) mm long,
(0.6—-)0.8—1.9(—2.2) mm wide; pappus scales (15—)17—28, (1.7-)2.1-
3.7(-4.4) mm high. Gravelly, rocky soils in grasslands, deserts, and
woodlands; 60—2000 m.

Comments. Coulter’s specimen bears no date, but it probably was
collected sometime during his excursion from Monterey to Yuma,
Arizona, by way of San Diego, between 20 March and 19 July 1832
(Coville 1895, McKelvey 1955). The route followed by Coulter’s
party (Fig. 2) passed through or near three possible collecton areas.
1) Northeastern Los Angeles Co. According to Coville (1895), the
route was on the southwestern side of the San Gabriel Mountains,
between San Fernando and San Gabriel. The only known localities
for A. sphaerocephalus in or near the San Gabriels (Pallett Creek,
Little Rock Creek, and Bob’s Gap) are on the northeastern side of
the mountains. 2) East-central San Diego Co. Coulter passed through
either the San Felipe Valley or the next valley south between 30
April and 8 May, and again between 17 and 27 May on the return
trip (Coville 1895). There are a number of specimens from this area,
and the dates of Coulter’s visit during the outbound trip coincide
with those for collections that are in the same state of maturity as
the type. 3) Area of Yuma, Arizona. Coulter was in this area 8
through 17 May (Coville 1895); however, specimens from that por-
tion of the range of the species have completely mature achenes by
that date, and the type specimen has only partially mature achenes.
This information suggests that the type locality is one of the valleys
of east-central San Diego Co.

Representative specimens. USA, Arizona, Coconino Co., Glen
Canyon National Recreation Area, Glen Canyon, ca. 1 mi due s. of
Wahweap Marina, 26 May 1983, Welsh 22066 (BRY, RM). Gila
Co., between Roosevelt Dam and Tonto Basin, 15 May 1935, Nelson
and Nelson 1933 (GH, K, MO, NY, RM, UC, US, UTC). Graham
Co., 2 mi below San Juan Mine, Gila Mts., 8 Apr 1935, Maguire
and Maguire 10546 (ARIZ, GH, MO, NY, UTC). La Paz Co., 1 mi
s. of Alamo State Park boundary on road to Wenden, 10 Mar 1973,

1988] LANE: ACAMPTOPAPPUS 263

McLeod and Pinkava 10326 (ARIZ, ASU, LL, NY, SD, TEX). Mar-
icopa Co., roadside s. of Canon, 21 Apr 1938, Foster and Arnold
338 (CAS, GH, UC, US). Mohave Co., Yucca, 15 May 1884, Jones
3911 (ARIZ, CAS, F, GH, POM, RM, UC, US, UTC). Pima Co..,
Walls Well, Organ Pipe Cactus National Monument, 28 Apr 1939,
Nichol s.n. (ARIZ). Pinal Co., Camp Grant, 14 May 1867, Palmer
114 (GH, MO). Yavapai Co., Black Canyon P.O., 14 Apr 1960,
Demaree 42241 (ARIZ, NY, TEX). Yuma Co., Yuma, 21 Apr 1938,
Crooks and Darrow s.n. (ARIZ, NY). California, Imperial Co.,
Mountain Springs Grade, 17 Apr [without year], Orcutt s.n. (UC).
Inyo Co., Dante’s View above Death Valley, 11 Jun 1930, Peebles
302 (ARIZ, NY). Kern Co., 8 mi ne. of Mojave, 12 May 1930,
Howell 4913 (CAS). Los Angeles Co., Pallett Creek, San Gabriel
Mts., 27 May 1923, Munz 6896 (CAS, NY, RSA). Riverside Co.,
San Jacinto Mts., e. base along the borders of the Colorado Desert,
Jun 1901, Hall 2108 (CAS, K, MO, NY, POM, UC, US). San Ber-
nardino Co., Baker, Mohave Desert, 2 May 1933, Jones s.n. (GH,
K, RM, RSA, UC, UTC). San Diego Co., Yaqui Wells, Colorado
Desert, 14 Apr 1913, Eastwood 2800 (CAS, GH, K, NY, US). Ne-
vada, Clark Co., St. Thomas Gap area sw. of Whitney Ridge, 0.9
road mi n. of Grand Gulch Rd on Reservoir Rd, 14 May 1982,
Tiehm 6867 (MO, RSA, UTC). Lincoln Co., Moapa, 5 May 1909,
Kennedy 1808 (F). Utah, Kane Co., ca. 2 mis. of Nun Butte, ca. 20
mie. of Glen Canyon City, 24 May 1972, Atwood 4062 (MO, US).
Washington Co., e. slope of Black Hill, St. George, 24 May 1942,
Gould 1772 (CAS, COLO, F, GH, NY, POM, UC, UTC).

EXCLUDED TAXON

Acamptopappus microcephalus M. E. Jones, Contr. W. Bot. 8:33.
1898. =Ericameria cooperi (A. Gray) H. M. Hall subsp. cooperi.

ACKNOWLEDGMENTS

This study was supported in part by NSF grant BSR-850631. I thank Robyn Tierney
for the illustrations in Fig. 3. Barbara Ertter, Leila Shultz, Barbara Timmermann,
and especially John Strother provided information on specimens, chemistry, and
chromosome counts; Ron Hartman, John Semple, and Greg Brown discussed generic
relationships with me. The comments of three reviewers improved the manuscript.
Loan of specimens by the several herbaria listed in the methods section is acknowl-
edged, and I am particularly grateful for the hospitality of RM during most of this
study.

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Romney, E. M. and A. WALLACE. 1980. Ecotonal distribution of salt-tolerant shrubs
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SCHAAK, C. G., R. HEVLy, and M. L. RuScHE. 1982. IOPB chromosome number
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SEMPLE, J.C. 1976. The cytogenetics of Xanthisma texanum DC. (Asteraceae) and
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GENECOLOGY OF CERASTIUM ARVENSE AND
C. BEERINGIANUM (CARYOPHYLLACEAE)
IN NORTHWEST WASHINGTON

STEVEN J. WAGSTAFF
Department of Botany, Ohio University,
Athens 45701

RONALD J. TAYLOR
Department of Biology, Western Washington University,
Bellingham 98225

ABSTRACT

From principal components analyses, patterns of morphological variation were
determined in and among three populations of Cerastium arvense and one population
of C. beeringianum, under both field (natural) and garden conditions. The three
populations of C. arvense occurred at near sea level, mid-montane and alpine ele-
vations, respectively. The C. beeringianum population was also alpine and occurred
on a serpentine substrate, as did the mid-montane population of C. arvense. The
analyses showed considerable phenotypic plasticity in non-serpentine populations,
much less in serpentine populations. There appeared to be little genetic differentiation
in C. arvense along the elevational gradient, except for a more or less persistent
cushion habit in the alpine population. The two closely related species overlapped
morphologically but could be separated by variables used, especially when grown
under garden conditions.

Cerastium arvense was described as “‘one of the most perplexing
species in our range”’ by Hitchcock et al. (1964). They noted further
that C. beeringianum cannot be satisfactorily separated from C.
arvense in the high Cascades where the characteristics of the two
taxa tend to merge. According to Hultén (1956) these species hy-
bridize in Newfoundland and Labrador and are part of a large poly-
ploid complex united by introgressive hybridization. Chromosome
numbers of most members of the complex are known (Sollner 1954,
Brett 1955, Ugborogho 1973, 1977). Ploidy of C. arvense varies but
appears to be diploid (2” = 36) throughout the Pacific Northwest,
whereas C. beeringianum 1s a tetraploid (2n = 72). Meiotic regularity
and high pollen viability provide evidence that C. beeringianum is
a stable allotetraploid.

Interspecific hybridization in Hultén’s complex has undoubtedly
been facilitated by reproductive biology. At least C. arvense is an
obligate outcrosser that requires insect pollination for successful seed
set (Ugborogho 1977); and both taxa have open, bowl-shaped flowers
that provide easy access to pollen and nectar rewards by insects. The

MApRONO, Vol. 35, No. 3, pp. 266-277, 1988

1988] WAGSTAFF AND TAYLOR: GENECOLOGY OF CERASTIUM — 267

generalist pollination strategy of the two species has been substan-
tiated (Ugborogho 1977, Arroyo et al. 1982, Shaw and Taylor 1986).

Cerastium arvense is widely distributed, both geographically and
altitudinally. In the Pacific Northwest it ranges from rocky, exposed
coastlines to alpine ridges. Cerastium beeringianum, on the other
hand, has a restricted distribution in the Northwest, occurring in a
few alpine locations. The purpose of our study, then, was two-fold:
(1) to examine the patterns of variation within and among geograph-
ically and elevationally disjunct populations of Cerastium and to
ascertain the extent to which observed variation was the result of
phenotypic plasticity and to what extent it is genetically fixed; (2)
to confirm the taxonomic distinction of an alpine population thought
to be C. beeringianum and to compare patterns of variation. of this
population to those of C. arvense.

METHODS

Four sites varying in climate, elevation, and edaphic conditions
were chosen for study. These sites (Deception Pass, Sumas Moun-
tain, the Twin Sisters, and Chowder Ridge) are shown in Fig. | and
general descriptions are given in Table 1. Detailed descriptions of
the climate and geology relating to the four sites are available from
Moen (1962), Phillips (1966), and McKee (1972). Taylor and Doug-
las (1978) described the natural history of Chowder Ridge, and
Kruckeberg (1969) published a detailed account of vegetation oc-
curring on serpentine soils in the northwest, including Sumas Moun-
tain and the Twin Sisters.

Sampling and collection. At Deception Pass, Chowder Ridge, and
the Twin Sisters Cerastium populations were large and the plants
occurred in diverse habitats. To effectively sample the variation of
these populations, we positioned four widely separated, 55 m tran-
sects parallel to the slope; the sum of the transect length was therefore
220 m. Ten specimens were collected at 20 m intervals along the
total transect length. Because the Cerastium population at Sumas
Mountain was smaller and more limited in distribution, seven short-
er (20-55 m) transects were established, again with a total length of
220 m. Ten specimens were also collected at 20 m intervals along
this total transect length. From each of the 40 specimens collected,
three mature flowers with non-dehisced anthers were selected for
pollen analysis, and plants were pressed for subsequent morpholog-
ical analyses. These collections were made in the summer of 1983.

Also during the summer of 1983, 40 cuttings were collected at 5
m intervals along the transects at each study site. These were prop-
agated in a potting mixture of one part each mineral soil (taken
from respective study sites), peat, and perlite. In 1984 an additional
20 cuttings were similarly collected from each study site and prop-

268 MADRONO [Vol. 35

BRITISH COLUMBIA
WASHINGTON

GLACIER

CHOWDER RIDGE
ASTUDY SITE
Qu. BAKER

TWIN SISTERS
STUDY SITE
a

WHATCOM CO.
SKAGIT CO.

122°

49°
SKAGIT RIVER

fo
‘ISLAND.CO.

&
YWSITE
~
Via
| . i141

Fic. 1. Location of the four populations (triangles) of Cerastium studied in North-
west Washington. The Twin Sisters population = C. beeringianum.

agated in a mixture of one part each sand, peat, and perlite. All
cuttings were protected from direct sunlight until new growth was
observed, then transferred to an experimental garden at Western
Washington University. In June 1985, 10 mature specimens were
randomly collected from each of the four transplanted populations
and pressed.

Pollen analysis. The flowers sampled for pollen analysis were stored
at 5°C until slides could be prepared using a technique adapted from
Ugborogho (1973). Two anthers from each of the three flowers were
squashed in a 1% acetocarmine and glycerine solution. The stained
mounts were allowed to set for 24 hours before being examined.
This enabled the pollen grains to absorb the stain and stabilize in
size. Using a calibrated stage, each slide was then divided into quad-
rats and 25 pollen grains were observed from each quadrat. Pollen
grains that were swollen and deeply stained were scored as viable.
In addition, every tenth pollen grain was measured using a calibrated
occular.

Morphometric analysis. Twenty-three morphologic and pollen
variables were chosen for analyses. These are listed in Tables 2 and

269

1988] WAGSTAFF AND TAYLOR: GENECOLOGY OF CERASTIUM

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272 MADRONO [Vol. 35

3 and were used in principal components analyses available through
Nie et al. (1983). In addition, a t-test of independent means was
used to compare the intrapopulation variation between garden and
field collections from each of the four locations. The significance
level was established at p < 0.05.

RESULTS

Transplant establishment. Between 25 and 41% of the cuttings
survived transplantation. Survival rates during the several months
that the plants were maintained in the pots depended on the pop-
ulation source and potting mixture. Plants from Chowder Ridge and
Deception Pass exhibited apparent preference for soil mixtures con-
taining sand rather than native soil. In contrast, plants from the
serpentine populations fared significantly better when propagated in
mixtures containing native soils.

Following transplantation from the pots into the garden, the plants
readily became established and formed mats up to 83 cm in diameter
by the end of the second year. During the second year the garden
transplants flowered and successfully set seed. As shown in Fig. 2,
the low elevation field populations flowered earlier in the season
than did the alpine populations. However, garden transplants flow-
ered simultaneously regardless of their source.

The requirements for seed germination were less uniform. Seeds
from Deception Pass and Sumas Mountain transplants germinated
within ca. 14 days on moistened filter paper at 5°C, whereas low
germination rates were observed for alpine transplants.

Factor analyses. Analyses involving variables listed in Table 2
were used to quantify intra- and interpopulational variation, to es-
tablish relationships among populations, and to determine the extent
to which observed variation was due to phenotypic plasticity. Figure
3 is a two-dimensional ordination produced from a principal com-
ponents analysis (PCA) of the 40 field specimens. This analysis
revealed variation within each population and overlap among pop-
ulations. Forty-one percent of the variance was accounted for by the
first two components. Characters that received high loading on the
first component were those involving overall size, inflorescence, and
pollen diameter. Characters that described bract shape, and width
of the scarious bract margin received high loadings on the first and
second components. Factor loadings for the 23 variables in this and
subsequent PCA’s are given in Table 2.

As reflected in the PCA ordination of Fig. 4, interpopulational
variation was reduced in garden transplants, especially between the
Deception Pass and Sumas Mountain populations. The distinction
between the Twin Sisters population, which ultimately proved to be
C. beeringianum, and other, C. arvense, populations was rather sharp.

1988] WAGSTAFF AND TAYLOR: GENECOLOGY OF CERASTIUM — 273

JUNE JULY AUG. SEPT.
zr)

APRIL MAY
a

LD cn 0 Nn 40 oN 0 en
ple 4A0® FieX yak pie ade pir aoe

DECEPTION SUMAS TWIN CHOWDER
PASS MOUNTAIN SISTERS RIDGE

Fic. 2. Flowering period for field populations of Cerastium and garden transplants.

In this analysis 35% of the variation was accounted for by the first
and second components. Characters that described the overall height
of the plants, the number of sterile shoots, and the width of the
scarious bract margin had high loadings on the first component and
therefore differed between species. Characters describing the shape
of the bracts and the width of the leaves received high loadings on
the second component (Table 2).

A PCA ordination of the combined field and garden data sets (Fig.
5) indicates considerable overlap among the populations. There is
little differentiation between field and garden populations from the
Twin Sisters and from Sumas Mountain. However, field and garden

[Vol. 35

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MADRONO

274

1988] WAGSTAFF AND TAYLOR: GENECOLOGY OF CERASTIUM — 275

populations from the non-serpentine sites, Chowder Ridge and De-
ception Pass, clustered separately. This suggests that plants of the
latter sites exhibited greater phenotypic plasticity. In Fig. 5 the first
and second principal components accounted for 42% of the variance.
Morphological complexes that described plant height, leaf width,
and the inflorescence received high loading on the first component
and were therefore important in separation of the Twin Sisters pop-
ulation and, to a lesser extent, the Chowder Ridge field population
from other populations. Plant height, the width of the scarious mar-
gin on the bracts and the number of sterile shoots at the leaf nodes
received high scores on the second component and thus were 1m-
portant in separating the field from the garden populations from
Chowder Ridge and, especially, Deception Pass. Factor loadings are
given in Table 2.

To determine extent of variation due to phenotypic plasticity, the
independent means of variables of field and garden specimens were
compared (Table 3). The number of variables that differed signifi-
cantly between field and garden specimens was: 17 from the Chowder
Ridge population, 14 from Deception Pass, 6 from the Twin Sisters,
and 5 from Sumas Mountain. This strongly suggests that the Chow-
der Ridge and Deception Pass populations expressed more pheno-
typic plasticity than did the populations occurring on serpentine
substrates. The most plastic variables were length of internodes,
distance between bracts and first leaves, number of nodes, length of
pubescence, and pollen diameter. These variables varied signifi-
cantly between field and garden specimens from at least three of the
four populations (Table 3). The least plastic variables were those
describing flowers, width of the scarious margin of bracts, leaf width,
number of sterile shoots, and pollen stainability.

DISCUSSION

In spite of the morphological similarity and overlap between al-
pine populations of Cerastium arvense and C. beeringianum, the two
taxa were separable on the basis of characters used in this study.
This was especially true of garden specimens (see Fig. 4). As pre-
dicted from taxonomic treatments (Fernald and Wiegand 1920, Hul-
tén 1956, Hitchcock et al. 1964), vegetative characters were more
useful in distinguishing the taxa than were the conservative floral

_

Fics. 3-5. Principal components ordinations. 3. Forty field specimens from four
populations: | = Deception Pass, 2 = Sumas Mountain, 3 = Twin Sisters, 4 = Chowder
Ridge. 4. Forty garden specimens originally from four populations: 5 = Deception
Pass, 6 = Sumas Mountain, 7 = Twin Sisters, 8 = Chowder Ridge. 5. Eighty field
and garden specimens (populations numbered as above); populations 5, 6, 8 = Ce-
rastium arvense, 7 = C. beeringianum.

276 MADRONO [Vol. 35

characters. In general, bracts subtending the inflorescence were re-
duced and scarious-margined in C. arvense, foliaceous and non-
scarious in C. beeringianum. Leaves were narrower and longer in
C. arvense, and axillary fascicles were restricted to the lower leaf
nodes in C. beeringianum. However, the separation of the taxa by
principal components analyses was the result of the correlation of
characters and not absolute differences between the taxa, thus the
difficulty of field identification of alpine forms, as noted by Hitchcock
et al. (1964).

The PCA’s suggest that there is little genetic distinction between
the alpine (Chowder Ridge) population and low elevation popula-
tions of C. arvense. However, the dwarf, mat-like growth habit of
the former was apparently genetically fixed. In this respect the Chow-
der Ridge and Twin Sisters populations were similar. The two alpine
populations also exhibited more variability in terms of seed ger-
mination requirements, a common adaptive characteristic of alpine
plants. Flowering times were not similarly fixed since all populations
flowered simultaneously in the garden. The large amount of phe-
notypic plasticity in non-serpentine populations is undoubtedly
adaptive and helps to explain the broad ecological tolerance of Ce-
rastium arvense. It also masks genetic distinctions among popula-
tions and species. This can be seen by comparing Figs. 3 and 4; the
two species were much more distinct when grown under similar
conditions. The restricted plasticity of the Twin Sisters and Sumas
Mountain populations was perhaps due to selective pressures and
specialization associated with the peculiarities of serpentine soils.
Specialization is reflected not only by low plasticity, but also by the
lower rooting success in non-serpentine substrates. Similar obser-
vations were recorded by Kruckeberg (1967) in his work with ser-
pentine plants.

As previously noted, one of the objectives of this study was to
confirm the taxonomic distinction of Cerastium arvense and C. beer-
ingianum. The selection of the Twin Sisters population was made
for two reasons: it was a representative alpine population, and it
was suspected to be C. beeringianum. As noted above, this popu-
lation was morphologically distinguishable, especially under uni-
form (garden) conditions (Fig. 4) and it proved to be tetraploid in
contrast with the diploid C. arvense populations (Wagstaff 1986).
Our study confirms, however, that because of phenotypic plasticity,
the taxa cannot easily be distinguished in the field. From our limited
study, it would seem that the keys and descriptions of Hitchcock et
al. (1964) and Hultén (1956) are satisfactory but that the non-plastic
and correlated characters, plant height, width of scarious margins
of bracts, leaf width, and number of sterile shoots, should be em-
phasized.

1988] WAGSTAFF AND TAYLOR: GENECOLOGY OF CERASTIUM — 277

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McKeEE, B. 1972. Cascadia: the geologic evolution of the Pacific Northwest. McGraw-
Hill, New York.

Moen, W.S. 1962. Geology and mineral deposits of the north half of the Van Zandt
Quadrangle. U.S.D.I. Div. Mines Geol. Bull. 50.

Nig, N. H., C. H. Hutt, J. G. JENKINS, K. STEINBRENNER, and D. L. BENT. 1983.
Statistical packages for the social sciences. 2nd ed. McGraw-Hill, New York.

PHILLIPS, E. L. 1966. Washington climate. Cooperative Extension Service, Wash-
ington State Univ., Pullman.

SHAW, D. and R. J. TAYLOR. 1986. Pollination biology of a fell-field community
on Chowder Ridge. Northw. Sci. 60:21-31.

SOLLNER, R. 1954. Recherches cytotaxinomiques sur le genre Cerastium. Ber. Schweiz.
Bot. Ges. 64:221-354.

TAYLOR, R. J. and G. W. DouGLas. 1978. Plant ecology and natural history of
Chowder Ridge, Mt. Baker: a potential alpine research natural area in the western
North Cascades. Northw. Sci. 52:35—SO.

UGBOROGHO, R. E. 1973. North American Cerastium arvense. Cytologia 38:559-
566.

1977. North American Cerastium arvense L.: taxonomy, reproductive sys-
tem and evolution. Phyton 35:169-187.

WAGSTAFF, S. J. 1986. Genecology of Cerastium arvense and C. beeringianum in
Northwest Washington. Unpublished M.S. thesis, Western Washington Univ.
Bellingham. 62 p.

(Received 30 Jul 1987; revision accepted 16 Mar 1988.)

NOTEWORTHY COLLECTIONS

ARIZONA

AQUILEGIA TRITERNATA Payson X A. CHRYSANTHA A. Gray (RANUNCULACEAE). —
Coconino Co., Mogollon Rim, Dane Spring Canyon, at the base of a damp, shaded,
sandstone cliff face, T13N R11E S35, 7 Jul 1987, Schaack 2115 and Goodwin (ASC).

Significance. The first known report of natural hybridization between A. chrysantha
[A. caerulea group] with erect, yellow, long spurred [(40-)45—70 mm] flowers and A.
triternata [A. canadensis group] with nodding, primarily red, short spurred [(12-)16—
22 mm] flowers. Miller (Southw. Naturalist 30(1):69-76, 1985) did not see evidence
of this hybridization during his research in the mountains of southern Arizona and
indicated that the degree of genetic isolation between these species, due to habitat
and pollination system, has not been studied. Flowers of the vouchered hybrid were
intermediate in flower coloration (a curious mix of red and yellow) and spur length
(32-35 mm). Hybrid flowers drooped on their peduncle ends. Two putative F, hybrids
were the only hybridization products noticed at this site. The vouchered hybrid was
rooted 0.46 m above ground level in a recess on a sheer, shaded, damp sandstone
cliff face that housed the red flowered parent, A. triternata, above. The other hybrid
was discovered growing in the understory, at streamside, among A. chrysantha just
down canyon from the cliff face that held A. triternata. Although apparently a new
Aquilegia hybridization report, this is but one in a series of known hybridizations,
either artificial [Cockerell, Bot. Gaz. (Crawfordsville) 62:413-414, 1916; Anderson
and Schafer, Ann. Bot. (London) 45:639-646, 1931; and Taylor, Brittonia 19:374—
390, 1967] or natural (Grant, Aliso 2:341-360, 1952; and Miller, Amer. J. Bot. 65:
406-414, 1978) between members of the A. canadensis and A. caerulea species groups.
Seed was collected from the vouchered F, later in 1987. Future research will include
observation of pollinator activity between the parents and among parents and the
putative reciprocal F,’s and a search for the factors that apparently limit the estab-
lishment of backcross progeny at the Dane Spring Canyon population.— CLARK G.
SCHAACK, Department of Botany, University of Wisconsin, Madison 53706 and
GreGory A. Goopwin, Coconino National Forest, 2323 E. Greenlaw Lane, Flagstaff,
AZ 86004.

STYLOCLINE SONORENSIS Wiggins (ASTERACEAE). — Representative collections: Gra-
ham Co.: Hawk Hollow, 26 Apr 1935, Maguire s.n. (ARIZ, det. as Evax multicaulis
DC. or Stylocline gnaphaloides Nutt.; NY, mixed with and det. as Filago californica
Nutt.). Pima Co.: Tucson, Desert Research Laboratory, sandy plain w. of Tumamoc
Hill, 26 Apr 1968, Turner 68-146 and Mason (ARIZ, miixed with Filago depressa A.
Gray; det. as Stylocline micropoides A. Gray); mesas near Camp Lowell, 15 Apr 1881,
Pringle s.n. (F, MICH, both mixed with S. gnaphaloides and S. micropoides and det.
as the latter). Pinal Co.: Big Wash 0.5 mi [0.8 km] nw. of Oracle Junction on route
89, 29 Apr 1965, Hermann 19770 (RM, det. as S. micropoides). Santa Cruz Co.: ca.
16 mi [26 km] n. of Nogales along Hwy. 89, 27 Mar 1970, Higgins 2813 (BRY,
mixed with Filago californica; det. as F. depressa A. Gray). Also known from ca. 10
other collections in the above counties. Morefield thanks the curators of the herbaria
above for loans of material in their care.

Previous knowledge. See CA Noteworthy Collections, below.

Significance. First reports for the United States. —JAMES D. MOREFIELD, see note
below.

MADRONO, Vol. 35, No. 3, pp. 278-280, 1988

1988] NOTEWORTHY COLLECTIONS 279

CALIFORNIA

STYLOCLINE SONORENSIS Wiggins (ASTERACEAE).— Riverside Co.: Hayfields, n. of
Chuckwalla Mts., Colorado Desert, 2 Apr 1930, M. E. Jones 25845 (POM, originally
determined as Evax multicaulis DC., then as S. micropoides A. Gray).

Previous knowledge. Based only on the holotype (Mexico, n. Sonora, “One mile
north of Cumeral, on road to Nogales”, 9 Apr 1932, Abrams 13199, DS!) and on the
original description (Contrib. Dudley Herb. 4:26, 1950).

Significance. First report for the United States. The species is an inconspicuous
gray-woolly spring annual, and probably is more widespread in the Colorado Desert
of CA than the single extant CA specimen would indicate. It should be considered
rare and endangered in CA, however, until more sites can be located. The Hayfields
population may well have been extirpated after 1930 by development activities. The
species is more widespread in s. AZ (see AZ Noteworthy Collections, this issue).-—
JAMES D. MOREFIELD, Rancho Santa Ana Botanic Garden, 1500 N. College Ave.,
Claremont, CA 91711-3101.

ASPIDOTIS DENSA (Brackenr. in Wilkes) Lellinger (SINOPTERIDACEAE).—San Diego
Co., Cuyamaca Ranco State Park (CRSP), n. slope of Cherry Flat near Conejos Hiking
Trail, 32°57'34”N, 116°36'35”W, 1800 m, moist rocky areas in gabbro outcrops, 8
Jul 1987, Hirshberg s.n. (SD).

Significance. A range extension of ca. 350 km s. from the Greenhorn Mts., Kern
Co. Known previously from BC, Canada, s. to San Luis Obispo and Kern cos., CA,
e. to ID, MT, and UT, always on serpentine-derived soils (Smith, Madrono 23:15,
1974; Lellinger, Field Manual of Ferns and Fern-Allies of U.S. and Canada, p. 149,
1985). Gabbro is an ultramafic rock chemically similar to serpentine.

HOLODISCUS BOURSIERI (Carriére) Rehder in Bailey (ROSACEAE).—San Diego Co.,
CRSP, n. side of Cuyamaca Peak on rocky cliff above Cherry Flat, 32°56'49’N,
116°36'22”W, 1920 m, 28 Jul 1987, Hirshberg s.n. (SD, UC) (det. R. Lis).

Significance. A range extension of ca. 115 km se. from the Santa Ana Mts., Orange
Co., CA. Known previously from Trinity Co. s. to Orange Co., CA, and e. to w. NV.
This genus is currently under revision and species determination is necessarily ten-
tative (R. Lis pers. comm.).— JERILYN HIRSHBERG, P.O. Box 2, Julian, CA 92036 and
GEOFFREY A. LEVIN, see note below.

ASTRAGALUS PACHYPUS E. Greene var. PACHYPUS (FABACEAE).—San Diego Co.,
Anza-Borrego Desert State Park, Bighorn Cyn., T13S R6E n.'2 $2, 730 m, 14 Feb
1987, A. Morley s.n. (SD); same, 26 Feb 1987, Morley s.n. (SD) (det. R. C. Barneby).
Few plants in sandy wash.

Significance. A range extension of ca. 250 km se. from Antelope Valley, Los Angeles
Co. Known previously from Santa Barbara and Kern cos. se. to Los Angeles Co.,
and in San Benito Co.— GEOFFREY A. LEVIN, Botany Department, San Diego Natural
History Museum, P.O. Box 1390, San Diego, CA 92112.

CARLOWRIGHTIA ARIZONICA A. Gray (ACANTHACEAE).—San Diego Co.: Anza Bor-
rego Desert State Park, Borrego Palm Canyon Nature Trail ca. 2 km nw. of Borrego
Springs, rocky slope in Sonoran desert scrub with Larrea, Fouquieria, Justicia, Hyptis,
and Encelia, Borrego Palm Canyon 7.5’ ser. T10S R5E 826 se.%, ca. 300 m, 27 Mar
1988, M. Bourell 3509 (CAS).

Previous knowledge. Northwestern Baja California, central Arizona, and west Texas
southward throughout dry regions of Mexico, Honduras, and Nicaragua to north-
western Costa Rica (Guanacaste).

Significance. First report for this genus in California, doubling the number of taxa

280 MADRONO [Vol. 35

of Acanthaceae in the state. Range extension of 220 km nw. of the nearest Mexican
locality (Baja California, 9.5 km s. of La Ventana, Daniel 1545, ASU, CAS) and 230
km w. of the nearest known locality in the United States (Arizona, Yuma Co.: Kofa
Mountains, various collectons cited in Daniel, Fl. Neotrop. 34:1-116, 1983). The
California population marks the western limit of the distribution of both species and
genus. Of the numerous forms discussed by Daniel (ibid. and Desert Pl. 5:162-179,
1984) in this morphologically diverse species, Bourell 3509 most closely resembles
plants originally described as C. californica var. pallida 1. M. Johnst. from Baja
California, indicating a link with plants from that region rather than with those from
Arizona. — MONA BouRELL and THOMAS F. DANIEL, Department of Botany, California
Academy of Sciences, Golden Gate Park, San Francisco 94118.

BAJA CALIFORNIA SUR

QUERCUS OBLONGIFOLIA Torr. (FAGACEAE).— Mpio. de La Paz, Sierra de la Victoria,
oak woodland community, road to San Antonio de la Sierra Ranch, 6 km se. of El
Triunfo, 900 m, 23°43’N, 110°03’W; small population; José L. Leén 1132 (CIB).
Additional trees have been seen in exposed sites at middle elevations in the Sierra
de la Laguna, where they are called “‘encino laurel’. (Det. by comparison with spec-
imens at CAS.)

Significance. A range extension of 420 km se. from the Sierra de la Giganta where
it was reported by Carter (1955; ““Observaciones sobre los encinos de Baja California”’,
Bol. Soc. Bot. Mex. 18:39-42).

QUERCUS ARIZONICA Sargent (FAGACEAE).— Mpio. de La Paz, Sierra de la Laguna,
oak-pine woodland community, 2 km n. of La Laguna meadow, 1790 m, 23°36’N,
109°58'W, José L. Leén 1887, 2331 (CIB). A unique population of about 50 trees.
Acorn production is uncertain in this area. (Det. by D. E. Breedlove.)

Significance. Known previously in mountains of Arizona, and in the Sierra Madre
Occidental of Sonora and Chihuahua. Near the collection site is a deep brook where
Quercus reticulata H. & B. grows; the main distribution of this species is also in the
Sierra Madre Occidental.—JosE Luis LEON DE LA Luz, Centro de Investigaciones
Biologicas de Baja California Sur, Apdo. postal 128, La Paz, Baja California Sur,
México.

REVIEW

Conservation and Management of Rare and Endangered Plants. Edited by THOMAS
S. ELtAs. 630 pp. California Native Plant Society. Sacramento, CA. 1987. ISBN 0-
943460-11-5 (cloth), $45.00, ISBN 0-943460-12-3 (paper), $24.95.

This significant volume is the proceedings of a well-attended conference on rare
and endangered plants held under the auspices of the California Native Plant Society
in November of 1986. The objective of the conference was to bring together persons
interested in the biology, management, and preservation of rare plants for exchange
of ideas and information. The editors also sought early publication of the proceedings,
a goal which they achieved in good style. The resulting volume inlcudes 92 papers
by 106 authors. Both the picture and typography on the cover are very attractive—
suitable for display on the coffee table. The back cover, somewhat less striking,
provides a picture key for the identification of the editor and conference coordinator
(J. R. Nelson).

MADRONO, Vol. 35, No. 3, pp. 280-281, 1988

1988] REVIEW 281

Twenty papers addressed the social, legal, and institutional aspects of rare plant
protection and management. Though not the most riveting part of the volume, they
provide a valuable summary of the current laws and programs relevant to rare plant
conservation. The articles by Bartel and Cochrane will probably be especially useful
as references. About 15 papers deal with research needs and general methodology.
Among the topics addressed are reserve design, methods for sampling rare plants,
the role of artificial propagation, and computerized systems for storing and analyzing
data on rare plant habitat. The remaining papers are mostly case studies, though
many also attempt to generalize. Genetic and evolutionary questions are addressed
in a number of papers (e.g., Ledig, Conkle, Palmer), but the primary emphasis is on
population and community ecology. The work reported ranges from sophisticated
long-term studies (e.g., Kruckeberg on serpentine, Palmer’s study of Holocarpha) to
brief reports on obviously still-incomplete work in progress. The papers mostly deal
with California though Arizona work is reported in three papers and other studies
describe situations in Oregon, New Mexico, Alberta, Minnesota, and South Africa.

Though the editor and conference organizers are to be commended for getting the
volume out promptly, this no doubt contributed to the major problem with the book—
the uneven quality of the contributions. A few papers have major typographical and
stylistic blunders that more leisurely editing might have caught. Peer review would
also have shortened and improved the papers, and justified the exclusion of some.
Summary papers prepared after the conference that integrated the contributions and
guided the reader through them would also have been valuable.

A curious sidelight, of uncertain significance, is the greater than random frequency
of occurrence of some authors. One author appears on five papers (only four of which
are listed in the General Index), two are involved in three papers, and nine others
appear on two papers. Is this evidence that rare plant biologists, like some rare plants,
are highly aggregated and have a tendency to inbreeding?

The heterogeneity of the contributions makes it difficult to summarize the major
conclusions. It is apparent, as Messick and others point out, that we need more status
data on population sizes, locations, and degree of protection and threat. We also need
more information on the population ecology and life histories of rare plants, especially
as these are affected by events of low frequency like fire and severe drought. The
papers, as is appropriate for rare taxa, stress minimally intrusive observational meth-
ods. But experimental studies, where possible, are also necessary and should be
supported by those agencies with the resources to do so.

The heterogeneity is also probably an accurate reflection of the diversity of persons
and approaches that deal with rare plant problems. Research on rare plants is relatively
new, geographically dispersed, and in large part low-tech and labor intensive. These
circumstances favor decentralization and a release of variability. No doubt there will
be evolution in the direction of greater uniformity as the most useful research methods
and management techniques come to be recognized. This volume, by putting a large
sample of the current activity before us, should speed the selection process. It provides
a good review of the state of our knowledge about endangered plants. There is no
doubt that these proceedings will have an influence on the direction that rare plant
research will take in the future. It is a book anyone with an interest in rare plants,
especially in California, will want to have.— PAUL H. ZEDLER, Department of Biology,
San Diego State University, San Diego, CA 92182-0057.

Volume 35, Number 3, pages 169-281, published 2 November 1988

ANNOUNCEMENT

CALIFORNIA BOTANICAL SOCIETY
Schedule of Speakers
1988-1989

8:00 p.m. University of California, Berkeley LSB 2503*
Speaker and Topic

Date
15 Sep

Linda Newstrom
Dept. Botany, University of California, Berkeley
“Botany and ethnobotany of Chayote
(Sechium edule)”

James Shevock
U.S. Forest Service
““Phytogeography of the southern Sierra Nevada”

John Thomas
Dept. Biological Sciences, Stanford University
‘“‘A history of botany in northern California’

Thomas Duncan, Christopher Meacham
Dept. Botany, University of California, Berkeley
(1) Computerized image analysis and
(2) Floristic data bases”

SPEAKER AND LOCATION FOR
ANNUAL BANQUET TO BE ANNOUNCED

Roger Raiche
University of California Botanical Garden, Berkeley
“The native plant collection at the
U.C. Botanical Garden”

Thomas Rosatti
Jepson Herbarium, University of California, Berkeley
‘Systematics and ecology of Arctostaphylos uva-ursi
in North America”

Linda McMahan
Center for Plant Conservation,
Arnold Arboretum, Harvard University
‘Helping the Ark land: the role of
botanical gardens in plant conservation”’
[joint meeting with the California
Native Plant Society]

* Except 18 May, which will be at the U.C. Berkeley Botanical Garden

For a monthly reminder of the meetings, please notify Dr. James
Affolter, Botanical Garden, University of California, Berkeley, CA 94720.

ANNOUNCEMENT

NEw PUBLICATION

KEELEY, J. E. 1988. Bibliographies on chaparral and the fire ecology
of other Mediterranean systems, 2nd ed. California Water Resources
Center, Univ. California, Rept. 69. 328 pp. ISSN 0575-4968 (paper-
bound). [A series of lengthy bibliographies dealing with chaparral and
other Mediterranean ecosystems, much expanded from Ist edition
(1984). Bibliographies for (mostly California) chaparral are (1) evo-
lution and systematics, (2) community structure, (3) fire and demog-
raphy, (4) seed germination and allelopathy, (5) morphology and phys-
iology, (6) soils and management, and (7) animals. Bibliographies are
also included for (8) California grasslands and (9) California forests—
fire and demography. For each Mediterranean system outside of Cal-
ifornia (Australia, Chile, Mediterranean Europe [including western
Asia and northern Africa], and South Africa) there is a bibliography
on fire and demography and a bibliography on morphology and phys-
iology. Also included is a bibliography on fire and demography of
miscellaneous (mostly non-Mediterranean) regions. English-language
publications predominate. Because duplicate citations are not in-
cluded under different headings, a search of more than one bibliog-
raphy may be necessary to insure that all references on a particular
topic are found.] Copies can be obtained from: Water Resources
Center, University of California, Rubidoux Hall, Riverside, CA 92521;
(714) 787-4327. The bibliographies can also be obtained for a nominal
fee on IBM compatible floppy disks from Dr. Jon E. Keeley, Occi-
dental College, Los Angeles, CA 90041.

ANNOUNCEMENT

SYMPOSIUM ON CALIFORNIA’S OAK WOODLANDS:
ATTITUDES AND RESPONSIBILITIES

The Range Management Advisory Committee to the California Board
of Forestry, and the California Department of Forestry and Fire Pro-
tection are sponsoring a symposium on California’s Oak Woodlands:
Attitudes and Responsibilities, 22-24 January 1989, at the Red Lion
Inn, 2001 Point West Way, Sacramento, CA. Landowners, policy mak-
ers, planners, resource managers, realtors, investors, development con-
sultants, engineers, and citizens will discuss key issues and viable options
for conserving oak woodlands and hardwood range. Panel discussions
will include: (1) Hardwood rangelands as investment property: impli-
cations for the future of the resource; (2) Development and planning in
oak woodlands: opportunities and constraints; (3) County-level planning
for hardwood resources; (4) Oak woodland and hardwood rangeland
residents: attitudes and objectives; (5) Effects of hardwood policy; and
(6) Is multiple use management possible? A complete program with
registration information can be obtained by contacting the Natural Re-
sources Institute, Humboldt State University, Arcata, CA 95521; (707)
826-4172.

ANNOUNCEMENT

ASSOCIATION OF CALIFORNIA HERBARIA

The Association of California Herbaria met on 30 July 1988 and
voted to adopt a set of bylaws and to officially incorporate as a non-
profit organization. The Association is organized for the purposes of (1)
promoting the development and use of California herbaria; (2) effecting
cooperation among California herbaria; and (3) increasing the awareness
of the value of these herbaria for (a) maintenance and management of
botanical diversity, (b) research in taxonomy and evolution of plants,

and (c) training and education concerning plant resources.

All public and privately owned herbaria and working collections in
California are eligible for membership. Dues are based upon the size of
the herbarium or collection as follows: <5000 specimens—$5.00/year;
between 5000 and 50,000 specimens— $10.00/year; between 50,000 and
100,000 specimens—$25.00/year; between 100,000 and 500,000 spec-
imens—$50.00/year; and >500,000 specimens—$250.00/year. Each
herbarium or working collection desiring to become a member of the
Association should submit a request in writing to Dr. Thomas Duncan,
Botany Department, University of California, Berkeley, CA 94720. Ad-
ditional information and a request for payment of dues will be sent in

reply.

SUBSCRIPTIONS — MEMBERSHIP

Membership in the California Botanical Society is open to individuals ($22 per year;
students $12 per year for a maximum of seven years). Members of the Society receive
MApDRONO free. Family memberships ($25) include one five-page publishing allotment
and one journal. Emeritus rates are available from the Corresponding Secretary.
Institutional subscriptions to MADRONO are available ($30). Membership is based on
a calendar year only. Applications for membership (including dues), orders for sub-
scriptions, and renewal payments should be sent to the Treasurer. Requests and rates
for back issues, changes of address, and undelivered copies of MADRONO should be
sent to the Corresponding Secretary.

INFORMATION FOR CONTRIBUTORS

Manuscripts submitted for publication in MADRONO should be sent to the editor.
All authors must be members, and membership is prerequisite for review.

Manuscripts and review copies of illustrations must be submitted in triplicate for
all articles and short items intended for NOTES AND NEWS. Follow the format
used in recent issues for the type of item submitted and allow ample margins all
around. All manuscripts MUST BE DOUBLE SPACED THROUGHOUT. For ar-
ticles this includes title (all caps, centered), author names (all caps, centered), addresses
(caps and lower case, centered), abstract, text, acknowledgments, literature cited,
tables (caption on same page), and figure captions (grouped as consecutive paragraphs
on one page). Order parts in the sequence listed ending with figures, and number each
page. Do not use a separate cover page, “erasable’’ paper, or footnotes. Manuscripts
prepared on dot matrix printers may not be considered. Table captions should include
all information relevant to tables. All measurements should be in metric units.

Line copy illustrations should be clean and legible, proportioned (including cap-
tions) to the MADRONO page, and designed for reduction to 7% original size. Scales
should be included in figures, as should explanation of symbols, including graph
coordinates. Symbols smaller than 1 mm after reduction are not acceptable. Maps
must include latitude and longitude references. Halftone copy should be designed for
reproduction at actual size. In no case should original illustrations be sent prior to
the acceptance of a manuscript. When needed they should be mounted on stiff card-
board and sent flat. No illustrations larger than 22 x 28 cm will be accepted.

Presentation of nomenclatural matter (accepted names, synonyms, typification)
should follow the format used for Rhus integrifolia in MADRONO 22:288. 1974.
Institutional abbreviations in specimen citations should follow Holmgren, Keuken,
and Schofield, Index Herbariorum, 7th edition. Abbreviations of serial titles should
be those in Botanico-Periodicum-Huntianum (Lawrence et al., 1968, Hunt Botanical
Library, Pittsburgh). If the correct abbreviation cannot be determined, the full serial
title should be used. Titles of books should be given in full, together with the place
and date of publication, publisher, and edition, if other than the first.

All members of the California Botanical Society are allotted five free pages per
volume in MADRONO. Joint authors may split the full page number. Beyond that
number of pages a required editorial fee of $65.00 per page will be assessed. The
purpose of this fee is not to pay directly for the costs of publishing any particular
paper, but rather to allow the Society to continue publishing MADRONO on a reasonable
schedule, with equity among all members for access to its pages. Printer’s fees for
illustrations and typographically difficult material @ $35.00 per page (if their sum
exceeds 30 percent of the paper) and for author’s changes after typesetting @ $4.50
per line will be charged to authors.

At the time of submission, authors must provide information describing the extent
to which data in the manuscript have been used in other papers that are published,
in press, submitted, or soon to be submitted elsewhere.

CALIFORNIA BOTANICAL SOCIETY

, VOLUME 35, NUMBER 4 OCTOBER-DECEMBER 1988

MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

Contents

THE ROLE OF CHRYSOLEPIS CHRYSOPHYLLA (FAGACEAE) IN THE Pseudotsuga-
HARDWOOD FOREST OF THE KLAMATH MOUNTAINS OF CALIFORNIA

Todd Keeler-Wolf 285
THE HIGH ELEVATION FLORA OF MOUNT ST. HELENS, WASHINGTON

Roger del Moral and David M. Wood 309
CHROMOSOME NUMBERS IN THE ANNUAL Muhlenbergia (POACEAE)

Paul M. Peterson 320
Astragalus nutriosensis (FABACEAE): A NEW SPECIES FROM EASTERN ARIZONA

Michael J. Sanderson B25
NEw COMBINATIONS IN Arctostaphylos (ERICACEAE): ANNOTATED LIST OF CHANGES IN

STATUS

Philip V. Wells 330

NOMENCLATURAL AND SYSTEMATIC REASSESSMENT OF Opuntia engelmannli AND O.
lindheimeri (CACTACEAE)

Bruce D. Parfitt and Donald J. Pinkava 342
Salix scouleriana (SALICACEAE) DISCOVERED IN MEXICO

| George W. Argus 350
NOTES

ADDITIONAL SUPPORT FOR THE RECENT-INVASIVE ADVENT OF MESQUITE (MIMOSACEAE:
Prosopis) IN THE SAN JOAQUIN VALLEY, CALIFORNIA
Dan C. Holland and Barrett Anderson 329

| NOTEWORTHY COLLECTIONS

| CALIFORNIA aoe
COLORADO 353
IDAHO 354
MONTANA 355
WASHINGTON 358
- ANNOUNCEMENTS 308, 319, 341, 349, 359, 360, 362, 364, 365
LETTERS AND COMMENTARY ff CRT SON A grey,
MESSAGE FROM PAST CBS PRESIDENT ra %, 361
EDITOR’S REPORT FOR VOLUME 35 # AN aK 363
_REVIEWERS OF MANUSCRIPTS Nan: 1770 Bea
INDEX TO VOLUME 35 bt at ea 21
DEDICATION | | Ne IERARIES Fi
TABLE OF CONTENTS FOR VOLUME 35 eras. iv

_DATES OF PUBLICATION 370

‘PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

}
}
if

MADRONO (ISSN 0024-9637) is published quarterly by the California Botanical So-
ciety, Inc., and is issued from the office of the Society, Herbarium, Life Sciences
Building, University of California, Berkeley, CA 94720. Subscription rate: $30 per
calendar year. Subscription information on inside back cover. Established 1916.
Second-class postage paid at Berkeley, CA, and additional mailing offices. Return
requested. POSTMASTER: Send address changes to James R. Shevock, Botany Dept.,
California Academy of Sciences, San Francisco, CA 94118.

Editor—Davip J. KEIL

Biological Sciences Department
California Polytechnic State University
San Luis Obispo, CA 93407

Board of Editors
Class of:

1988—SusAN G. CoNARD, USDA Forest Service, Riverside, CA
WILLIAM B. CRITCHFIELD, USDA Forest Service, Berkeley, CA
1989— FRANK VASEK, University of California, Riverside
BARBARA ERTTER, University of California, Berkeley
1990—STEVEN TIMBROOK, Ganna Walska Lotusland Foundation, Montecito, CA
THOMAS R. VAN DEVENDER, Arizona—Sonora Desert Museum, Tucson
1991—JAMEs HENRICKSON, California State University, Los Angeles
WAYNE R. FERREN, JR., University of California, Santa Barbara

CALIFORNIA BOTANICAL SOCIETY, INC.

OFFICERS FOR 1988-89

President: JOHN L. STROTHER, University Herbarium, University of California,
Berkeley, CA 94720

First Vice President: JAMES AFFOLTER, Botanical Garden, University of California,
Berkeley, CA 94720

Second Vice President: JAMES HENRICKSON, California State University, Los An-
geles, CA 90032

Recording Secretary: RODNEY G. Myatt, Department of Biological Sciences, San
Jose State University, San Jose, CA 95192

Corresponding Secretary: JAMES R. SHEVOCK, Department of Botany, California
Academy of Sciences, San Francisco, CA 94118

Treasurer: THOMAS F. DANIEL, Department of Botany, California Academy of Sci-
ences, San Francisco, CA 94118

Financial Officer: CHERIE L. WETZEL, Department of Biology, City College of San
Francisco, 50 Phelan Ave., San Francisco, CA 94112

The Council of the California Botanical Society consists of the officers listed above
plus the immediate Past President, DALE MCNEAL, Department of Biological Sciences,
University of the Pacific, Stockton, CA 95211; the Editor of MADRONO; three elected
Council Members: JOHN MOORING, Department of Biology, University of Santa Clara,
Santa Clara, CA 95053; BARBARA ERTTER, Herbarium, Botany Department, Uni-
versity of California, Berkeley, CA 94720; ELIZABETH MCCLINTOCK, Herbarium, Bot-
any Department, University of California, Berkeley, CA 94720; and a Graduate
Student Representative, VALERIE HALEY, Department of Biological Sciences, San Jose
State University, San Jose, CA 95192.

THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.

THE ROLE OF CHRYSOLEPIS CHRYSOPHYLLA
(FAGACEAE) IN THE PSEUDOTSUGA-HARDWOOD
FOREST OF THE KLAMATH MOUNTAINS
OF CALIFORNIA

TODD KEELER-WOLF
943 Aquarius Way, Oakland, CA 94611

ABSTRACT

The tree form of Chrysolepis chrysophylla (giant chinquapin) is the most restricted
and uncommon of the major sclerophyllous hardwoods in the Pseudotsuga-hardwood
zone of the Klamath Mountains Geologic Province. The vegetation of three areas
containing the species under different climatic conditions was examined to better
understand its restriction. Chrysolepis chrysophylla was found to have the highest
importance value of any hardwood tree averaged over all the sampling transects and
was second in importance only to Pseudotsuga menziesii, the major canopy species.
At all sites C. chrysophylla showed a distinct preference for mesic conditions with
highest basal area cover occurring on northerly-facing slopes, or in mesic canyon
bottoms. Seed-originated saplings were densest at the most mesic sites, whereas
asexual sprouts were most prevalent at xeric sites. Areas of highest ground cover
(>52%) appeared to have lower densities of sexual reproduction. Lithocarpus den-
siflora the predominant hardwood associate with C. chrysophylla had higher impor-
tance values at relatively low elevations, becoming insignificant at elevations above
ca. 1000 m, whereas C. chrysophylla continued as an important forest component to
above 1400 m. Comparative research from other areas in the Klamath Province
indicate C. chrysophylla requires mesic northerly-facing exposures and an annual
rainfall of 1524 mm or greater to attain subdominance. Its architecture pre-adapts it
for subdominance of the highest, snowiest portions of the Pseudotsuga-hardwood
zone at the interface with the Abies concolor zone. In contrast to previous assumptions,
C. chrysophylla is a climax, as well as a successional species in its small zone of
occurrence in the Klamath Mountains.

Giant chinquapin, Chrysolepis chrysophylla (Dougl.) Hjelmquist,
is an evergreen sclerophyllous species that ranges from San Luis
Obispo County in southwestern California to Madison County in
west-central Washington. The main body of its distribution les in
the coastal areas of northern California and southern and central
Oregon (Little 1971). Within this zone the species occurs in a variety
of vegetation types ranging from mesic coastal forests dominated by
western hemlock and redwood to montane mixed conifer, white fir,
red fir, and chaparral associations (Franklin and Dyrness 1973, Tay-
lor 1982, McKee ms.).

Whittaker (1960, 1961) lists giant chinquapin as one of the prin-
cipal components of the mixed evergreen forest formation. Whit-
taker chaiacterizes mixed evergreen forest as occurring in its most
highly developed state in the Klamath Mountains Province and as
the central prevailing climax for that area. According to Atzet and

MADRONO, Vol. 35, No. 4, pp. 285-308, 1988

286 MADRONO [Vol. 35

Wheeler (1982, 1984) giant chinquapin occurs within the Douglas-
fir, Port Orford cedar, and tanoak plant series in the Siskiyou Moun-
tains, within the Klamath Mountains Province. These three series
and the mixed evergreen forest of Whittaker are included within the
broad classification known as the Pseudotsuga-hardwood forest of
Sawyer et al. (1977). Pseudotsuga-hardwood forests occur in both
the North Coast Range and Klamath Mountains provinces of Cal-
ifornia and adjacent Oregon.

The ecological role of Chrysolepis chrysophylla within the Pseu-
dotsuga-hardwood zone is poorly understood. It is typically listed
as one of the major hardwoods of the zone (Munz and Keck 1959,
Sawyer et al. 1977, Holland 1986). However, it is not nearly as
ubiquitous a species as the other principal sclerophylls; Lithocarpus
densiflora (tanoak), Quercus chrysolepis (canyon live oak), or Ar-
butus menziesii (madrone). Only five out of ten candidate or estab-
lished Forest Service Research Natural Areas within the Douglas-
fir-hardwood zone of the Klamath province contain any significant
giant chinquapin component, whereas seven contain tanoak and ten
contain madrone and canyon live oak (Keeler-Wolf ms.). This re-
strictiveness is also indicated by the disjunct west slope Sierra Ne-
vada distribution of the same broad-leafed sclerophyll species. Here
giant chinquapin is the most restricted followed in order by tanoak,
madrone, and canyon live oak (Griffin and Critchfield 1972).

Within a given area Chrysolepis chrysophylla is typically a species
of lower crown cover than either madrone, canyon live oak, or
tanoak. This point may be inferred from the maps of Griffin and
Critchfield (1972), which rely in large part on the Vegetation Type
Map and Soil-Vegetation surveys produced by the U.S. Forest Ser-
vice. These surveys only list a tree species if it covers 20% or more
(VTM) or 5% or more (S-V) of a given area. The areas of extensive
stands of Lithocarpus, Arbutus, and Quercus chrysolepis shown in
Griffin and Critchfield are much larger than those of Chrysolepis.
Giant chinquapin’s low density and cover is also indicated in Whit-
taker’s extensive 1960 study of vegetation in the Siskiyou Moun-
tains. There C. chrysophylla is in its greatest densities at elevations
between 610 and 915 m on quartz diorite where it is between two
and nine times less dense than the other sclerophylls. Whittaker’s
study indicates giant chinquapin is substantially reduced in density
compared to most other evergreen sclerophylls on gabbro, and is
only represented by an uncommon shrubby form on serpentine. In
the Klamath Province the tree forms occupy mesic forest from near
sea level to ca. 1600 m elevation, higher than any other broad-leafed
sclerophyll except QO. chrysolepis.

The rather broad ecological amplitude of giant chinquapin alluded
to in the first paragraph has much to do with the inclusion of shrubby
ecotypes of C. chrysophylla (including C. c. var. minor, the golden

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA 287

chinquapin) in the description of its range. These shrub forms may
inhabit relatively xeric, poor soils, and/or high-elevation sites, and
comprise most, if not all, of the individuals of the species south of
Marin County, California (Whittaker 1960, Griffin and Critchfield
1972, Franklin and Dyrness 1973, McKee ms., D. Keil pers. comm.).
The shrubby forms appear to be ecologically very different than the
tree form. The tree form is usually restricted to mesic forest where
it occurs as a subcanopy tree beneath a Pseudotsuga-dominated
canopy.

Why should Chrysolepis be the least common of the major broad-
leafed sclerophylls in the Pseudotsuga-hardwood forest? What con-
ditions are optimum for growth of the species, and are these pre-
dictable, unifying conditions under which this species may form a
major part of acommunity? I will attempt to answer these questions
by investigating three different sites where giant chinquapin forms
a significant part of the tree layer.

METHODS

The three sites selected for study are all in and adjacent to can-
didate and proposed Forest Service Research Areas (RNA’s) in the
Klamath Mountains Province of California (Fig. 1). All of these
areas contain unlogged and otherwise humanly undisturbed exam-
ples of natural vegetation. The three sites contain principally mature
or late successional forest with dominant canopy trees typically rang-
ing in age from 250 to 600 years.

The Rough Gulch and Chinquapin Ridge proposed RNA’s (re-
ferred to together as the Rough-Chinquapin site) lie adjacent to one
another in southern Trinity County on the northern slopes of South
Fork Mountain (ca. 40°15'N latitude, 123°15'W longitude). They
include the most extensive study area (ca. 2500 ha) and range in
elevation from ca. 800 to 1800 m. Slope aspects are primarily nne.,
although significant areas of e., n., and some w. and s. exposures
also occur. The parent material is entirely South Fork Mountain
Schist (Irwin 1966, 1981), an early Cretaceous metasediment prone
to mass-wasting (Scott et al. 1980). Precipitation is estimated be-
tween 1524 and 1778 mm annually (Rantz 1972). This area was
originally surveyed in 1984 (Keeler-Wolf 1984), but was re-visited
and more thoroughly surveyed in 1987.

The second site occurs in and near the Bridge Creek RNA located
in the southwestern Marble Mountains in western Siskiyou County
(ca. 41°28’N latitude, 123°21’W longitude). This RNA covers ca.
730 ha, lies ca. 120 km north of the first site, and ranges in elevation
from ca. 670 to 1280 m. It is underlain entirely by granitic rock of
the Wooley Creek Pluton (Donato et al. 1982). Precipitation is es-
timated at between 1651 and 1778 mm annually (Rantz 1972). Slope

288 MADRONO [Vol. 35

fo) fo)
42 124 123 122

41

40

=

ea
50km
Fic. 1. Map of Northwestern California showing the boundary of the Klamath
Province (dashed line) and the location of the three study sites; R = Rough-Chin-
quapin, B = Bridge Creek, and G = Goose Creek.

aspects are also varied, with all cardinal directions represented with-
in the study area. Preliminary vegetation surveys were conducted
in 1984 (Keeler-Wolf 1985) with additional work in 1987 both in
and adjacent to the RNA.

The third site is centered within the Upper Goose Creek candidate
RNA located in southern Del Norte County (ca. 41°33’N latitude,
123°52’W longitude). This site lies ca. 43 km nw. of Bridge Creek
and is broken up into two units (total area of ca. 200 ha) that are
separated by a small area of disturbed forest. Elevations at this site
range from ca. 548 to 1097 m. The area is underlain by metasedi-
mentary rocks of the Galice Formation (Irwin 1966). Precipitation
is estimated at being between 2794 and 3048 mm annually (Rantz

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA 289

1972). Slopes are predominantly north and south facing with smaller
areas of west- and east-facing slopes. An ecological survey of the
Upper Goose Creek candidate RNA was performed in 1986 (Keeler-
Wolf 1987a) and additional work was conducted in 1987 both in the
candidate RNA and in adjacent undisturbed forest on Lems Ridge.

Each area was first surveyed to determine major vegetation types,
and coverage and distribution of giant chinquapin. Following this,
representative stands of forest containing chinquapin were sampled
using transects laid out along contours or compass bearings. Each
transect was intended to represent a relatively homogeneous cover
type, and an example of each type occurring at various slope ex-
posures and aspects was sampled so that the broad spectrum of
Chrysolepis occurrence in the area would be represented. Transects
were terminated upon entering a visibly different vegetation type.
The transects were punctuated every 50 or 75 m witha 10 X 10m
quadrat. In each quadrat the following information was obtained:
1) identity, density, and breast height cover of stems on all trees
over 2 m, 2) density and identity of all tree saplings between 0.3
and 2 m in height, 3) density and identity of all seedlings of trees
under 0.3 m in height, 4) presence and estimated percent cover of
all shrub and herb species, 5) average slope aspect, 6) average ele-
vation, and 7) average slope steepness. In this way 197 quadrats (ca.
2 ha) were sampled at the three sites. Chinquapin saplings were
further broken down into sexual (seed-originated) and asexual (re-
sprouts from stumps and trunk bases).

RESULTS

Forest composition. Table 1 shows importance values (relative
density, relative frequency, and relative basal area cover summed
and multiplied by 100) of all trees for all 11 transects. The mean
value of these over the three sites indicates forests with highest
importance values for Pseudotsuga with Abies concolor and Chry-
solepis as the principal associated species. Of the 19 species of trees
encountered, only these three had mean importance values over 16
and frequencies greater than 64%. However, eight additional species
occur on 46% or more of the transects.

Table 2 indicates similarity values (Bray and Curtis 1957) based
on importance values of trees for all stands on the three sites. The
similarity values are generally high (x = 0.642; where 0 = no sim-
ilarity and | = total similarity) and indicate the relatively uniform
frequency, density and cover of the major tree species. Note that
similarity values varied within and between the three sites so that
there were at least some pairs that had lower values within the same
sites than between the three sites. The distribution of species at the

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290

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA Poe iM|

TABLE 2. SIMILARITY MATRIX BASED ON IMPORTANCE VALUES OF TREES ON 11
TRANSECTS AT THE THREE SITES IN THE KLAMATH MOUNTAINS OF CALIFORNIA.

RCN RCR RRN REM RUN BLV BMN BLN BUS’ GM

RCR 0.662

RRN 0.822 0.789

REM ~ 0.653 0.765 0.745

RUN 0.555 0.879 0.718 0.824

BLV 0.742 0.698 0.810 0.780 0.627

BMN _ 0.694 0.624 0.706 0.638 0.568 0.784

BLN 0.762 0.637 0.809 0.678 0.567 0.794 0.680

BUS 0.610 0.613 0.670 0.551 0.522 0.588 0.561 0.618

GM 0.630 0.419 0.564 0.474 0.363 0.650 0.686 0.604 0.461

GU 0.778 0.434 0.622 0.463 0.350 0.603 0.612 0.621 0.535 0.743

three sites indicates the Rough-Chinquapin area as being the most
xeric inland location, lacking Lithocarpus and Acer circinatum, as
well as the mesic coastal species such as Tsuga heterophylla and
Chamaecyparis lawsoniana, which are restricted to the Goose Creek
site.

Evidence for a moisture gradient from the Rough-Chinquapin to
the Goose Creek sites is further indicated in the understory (shrub
and herb) importance values (relative frequency summed with rel-
ative cover, multiplied by 100) on the sites (Table 3). Although a
core of shrubs and herbs including Chimaphila umbellata, Berberis
nervosa, Vaccinium parvifolium, Rubus ursinus, Chimaphila men-
ziesii, Pteridium aquilinum, Iris spp., Goodyera oblongifolia, and
Viola sempervirens occur regularly within all sites, several xero-
morphic species such as Arctostaphylos nevadensis, Ceanothus cor-
dulatus, Quercus vaccinifolia, and Galium bolanderi occur only at
the most inland and xeric Rough-Chinquapin sites. Conversely, sev-
eral mesomorphic species such as Vaccinium ovatum, Gaultheria
shallon, Coptis lacinata, and Blechnum spicant occur only at the
most mesic and coastal Goose Creek site. The largest number of
species (58) occur at the Bridge Creek site, which is intermediate in
moisture relations between the other two sites.

Similarity values (Table 4) between the importance values of shrub
and herbaceous species on all transects are relatively low (X = 0.286),
but show the greatest dissimilarity between the Goose Creek and
other sites. This is largely because of the high importance values of
species unique, or nearly so, to that sample area such as Vaccinium
ovatum, Gautheria shallon, and Rhododendron macrophyllum. The
similarity values between the Rough-Chinquapin and the Bridge
Creek transects tend to be higher and there are only a few species
that are not shared between the two areas. The slightly more mesic
conditions at Bridge Creek are indicated best by the presence of
Quercus sadleriana and Achlys triphylla, as well as by the relatively

[Vol. 35

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MADRONO

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pydydp pjoid vjosdd
sdyjidoddy vdosjouop
DIOfIUDSY DION
DYIJUDANIUU DAIYINILT
40]021g UO]NDIOUapY
DSOWIIDA DUIIDIIWS
pIpUIID] S11d0)
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DAOYJIUN DIUOJUI]D

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IQAOD % X

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA 295

TABLE 4. SIMILARITY MATRIX BASED ON IMPORTANCE VALUES OF SHRUBS AND
HERBS ON 10 TRANSECTS ON THE THREE STUDY SITES IN THE KLAMATH MOUNTAINS
OF CALIFORNIA.

RCN RCR RRN REM RUN BMN- BLN - BUS’ GM

RCR 0.281

RRN 0.615 02273

REM 0.362 0.187 0.176

RUN 0.545 0.378 0.693 0.142

BMN 0.486 0.239 0.448 0.371 0.321

BLN 0.496 0.184 0.449 0.364 0.326 0.707

BUS O73. 03320 “O174 O191 (0.164 0.339 0.321

GM 0.152 0.062 0.102 0.151 (0.093 0.165 O0:102 0.026

GU 0.299 0.155 0.218 0.148 0.267 0.253 0.220 0.189 0.575

unimportant species, Paxistima myrsinites and Rhododendron mac-
rophyllum.

Tree regeneration. The importance values (relative density + rel-
ative frequency x 100) of saplings on the 11 transects indicate that
the principal saplings throughout the three study areas are Pseudo-
tsuga, Abies concolor, Lithocarpus, Chrysolepis (including both sex-
ual and resprout saplings), and Pinus lambertiana (Table 5). How-
ever, A. concolor does not occur at the Goose Creek site and
Lithocarpus does not occur at the Rough-Chinquapin site. The data
for seedlings (Table 6) indicate that Pseudotsuga, A. concolor, Chry-
solepis, Lithocarpus, Quercus chrysolepis, and Pinus lambertiana are
the most important species.

The combined importance values of both resprout and seed-orig-
inated saplings of Chrysolepis exceeds all other hardwood species at
the Rough-Chinquapin sites (Table 5). However, Lithocarpus sap-
lings may exceed total Chrysolepis saplings at both Bridge Creek
(BMN and BUS) and at the Goose Creek site (GM).

When importance values of seed-originated saplings of giant chin-
quapin are compared with densities of basal or stump sprout-orig-
inated saplings it is clear that at the Rough-Chinquapin site asexual
reproduction is prevalent. There, ratios of sexual to asexual saplings
ranged from total asexual representation on transect REM to 0.731
at RCN with an average of 0.294 for the five transects. In contrast,
both the Bridge Creek and the Goose Creek sites had ratios of sexual
to asexual sapings all greater than 1.0. At Goose Creek these were
1.09 and 1.35 whereas at Bridge Creek these ranged from total sexual
reproduction on BLE to 2.58 at BMN with an average of 8.06.

The densities of seedlings on the 11 transects were not lowest at
the Rough-Chinquapin site. Seedling densities were consistently low
at all sites, with the highest densities occurring at the RCN (16/ha~“')
and RUN (9/ha‘') transects at the Rough-Chinquapin site. All other
transects ranged between O and 3 seedlings per 0.1 ha.

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298 MADRONO [Vol. 35

TABLE 7. DENSITY AND BASAL AREA OF Chrysolepis, TOTAL BASAL AREA AND
DENSITY OF ALL TREES, DENSITY OF SEXUAL AND ASEXUAL Chrysolepis REPRODUCTION,
AND SLOPE DIRECTION AND ELEVATION OF ALL TRANSECTS AT THREE STUDY SITES IN
THE KLAMATH MOUNTAINS OF CALIFORNIA. *Density figures are per ha~'!. "Cover figures
are m’/ha_!. ‘Not applicable due to inner valley location.

Chry-

solepis

sapling

and
Chry- seed- Chry-
solepis Total ling solepis Chryso-

Tran- den- tree density sprout — /epis Total x slope <x elev.
sect sity? density (sexual) density cover? cover _ direction (m)
RCN 42 118 19 4 2.44 13.90 n. 1020
RCR 16 179 4 19 aol 15.80 nnw. 1390
RRN 37 147 4 26 4.16 1270 n. 1180
REM i 106 1 1 1.20 15:87 ene. 1220
RUN 11 126 9 27 0.59 11.01 n. 1432
BLV 54 175 49 3 0.76 16.33 —< 1158
BMN oH 154 20 5 1.94 10.94 nnw. 1036
BLN 62 156 70 8 1.45 13.89 n. 975
BUS 42 118 0 3 3.07 7.93 ssw. 1220
BLE 10 155 20 0 1.28 5.72 e: 1073
GM 28 110 3 3 1.98 jy | nnw. 716
GU 36 90 23 12 1.28 9.42 n. 1082

Dominance and slope aspect. The total basal area cover (m?/ha™')
for tree size chinquapin on all transects is shown in Table 7. The
very highest cover is at RRN on a north-facing slope that averages
ca. 1180 m elevation. The lowest cover is at RUN, which is an upper
north-facing slope averaging ca. 1432 m elevation. When just the
seven mid-slope transects (modal positions, as opposed to obviously
mesic lower slope and xeric ridgetop locations) are considered, the
highest cover consistently occurs on slope aspects which average
between n. and nnw. The lowest cover in these seven modal transects
occurs at REM, with an average slope aspect of ene. In a thorough
survey of slope aspects at all three study sites no extensive modal
slope areas were found where Chrysolepis occurred on aspects to the
south of due e. or due w. This was true even at the wettest Goose
Creek site. The only locations south of e. and w. where Chrysolepis
formed a significant cover were in ravines and valley bottoms (e.g.,
BLV, BLE) or at summit ridges (RCR, BUS). Table 7 indicates that
there is no correlation between total tree cover and chinquapin cover.

Moisture and reproduction. The highest densities of sexually re-
producing giant chinquapin (saplings and seedlings combined) gen-
erally occur at the most mesic sites. These sites (BLV, BLN, see
Table 7) are in valley bottoms, which, although not typically within

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA 299

eh Shy

“ae 2 AY eae ates) MESS ae = APT et

Fic. 2, Testes mesic north-facing understory exhibiting good Chrysolepis el
regeneration. Rough-Chinquapin site ca. 200 m from South Fork of Trinity River,
elevation 800 m.

the influence of permanent stream moisture, are at least experiencing
the effects of the relatively cool, shady environment (equivalent to
moisture gradient steps | and 2 of Whittaker 1960). Concomitantly,
the areas with lowest densities of seed-generated reproduction occur
on relatively xeric upper slopes (steps 6—8 of Whittaker 1960). These
include RCR, RRN, REM, and BUS transects (Table 7). These xeric
exposures are either near ridgetops (RCR and BUS) or are on rel-
atively open upper slopes. This situation is the rule throughout the
Bridge Creek and Rough-Chinquapin sites. Despite the fact that no
lower slope sites were sampled at Rough-Chinquapin, observation
(Fig. 2) indicates highest densities of sexual reproduction on the
lowest, mesic slopes in that area. The only exception to this rule is
at the Goose Creek site where the upper slope GU transect has higher
densities of saplings and seedlings than the mid-slope GM site.
Ground cover and moisture relations. As can be seen from Table
3 the average percent ground cover ranges from a low of ca. 6 at the
largely south-facing BUS ridgetop transect to a high of 83 at the
upper slope Goose Creek site (GU). Although there is a trend toward
increasing cover at wetter sites, this is confounded by the presence
of high densities of Xerophyllum tenax (beargrass) at RRN and
RUN, which otherwise have relatively xeric understories (its pres-
ence at these sites may relate to a history of slope instability). Al-

300 MADRONO [Vol. 35

hen + f “oh, a PPS SS , 4 -$ ° a >
, es ” i ae a ‘ a4 ee, 6i, ; j Rae * a eds ' oe: AS. ¢, Pg inca ed pe
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Fic. 3. Typical dense undergrowth of Rhododendron macrophyllum, Vaccinium
ovatum, and Gaultheria shallon at the wet-coastal Goose Creek site.

though the most mesic lower slope sites at BLV and BLN were shown
to have the highest densities of sexual reproduction, the effect of
understory density on sexual reproduction of chinquapin is negative
at high values (above ca. 52% cover). The five transects between 52
and 83% cover show a negative correlation (r = —0.85, d.f. = 3, t
= 3.09, p = 0.05) with sexual sapling and seedling density. This
suggests that the shade cast by high shrub and herb cover is inhibiting
germination and/or growth of Chrysolepis seedlings (Fig. 3). Thus,
chinquapin appears to reproduce best sexually in areas where mesic
conditions prevail, but where conditions are not conducive to dense
layers of understory species.

In contrast asexual reproduction by basal and stump sprouting
appears to be more prevalent at relatively xeric sites. The highest
values occur on the ridgetop and upper slope sites at Rough-Chin-
quapin.

Elevation and tree distribution. The importance of Lithocarpus
densiflora in the forests at Bridge Creek and Goose Creek establish
that it is the principal hardwood and subcanopy associate of Chry-
solepis. When Lithocarpus and Chrysolepis importance values are
compared at sites of co-occurrence, it appears that Lithocarpus 1s
most important at the lowest elevation site (GM) and least important
at the highest elevation sites (BLV, BUS). Elevation and importance
of Lithocarpus are negatively correlated with increasing elevation (r
= —0.89, t = 3.38, d.f. = 3, p= 0.05). On the other hand, Chrysolepis

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA 301

dominance and importance within areas of Lithocarpus presence are
not correlated with altitude (716-1220 m). However, if the area with
the greatest altitudinal variation, Rough-Chinquapin, is analyzed
there is a strong negative correlation between altitude and chin-
quapin importance on the four transects with similar slope exposure
(r = —0.99, t = 98.0, d.f. = 2, p > 0.001). Importance values (I.V.)
drop off sharply above 1200 m. Observation in higher elevation
areas adjacent to the other two study areas containing Lithocarpus
confirm that giant chinquapin cover drops at a much higher elevation
than Lithocarpus cover.

DISCUSSION

Comparative data confirm requirements for Chrysolepis subdom-
inance. A review of the other surveyed Forest Service candidate
RNAV’s within the Pseudotsuga-hardwood zone of the Klamath Prov-
ince reaffirms the data from this report and indicates the regular
requirements of the species. Giant Chinquapin is unimportant or
absent at the Adorni (Sawyer 1981), Ruth (Thornburgh 1981), Hen-
nessy Ridge (Thornburgh 1987), Pearch Creek (Keeler-Wolf 1987b),
and Williams Point (Sawyer and Stillman 1977a) candidate RNA’s
and is present in relatively low density and cover at Specimen Creek
(Sawyer and Stillman 1977b) and South Fork Mountain (Taylor
1975). What emerges is that Chrysolepis requires cool, mesic north-
erly-facing exposures and an annual rainfall regime of ca. 1524 mm
or greater to attain subdominance in the Pseudotsuga-hardwood
forest of Klamath Province.

Chrysolepis compared with Lithocarpus and other broad-leafed
sclerophylls. Chrysolepis predominates on mesic northerly-facing
slopes, where it co-dominates the subcanopy with Lithocarpus at
lower elevations, but comes to dominate the subcanopy above ca.
1000 m elevation (or at lower elevations of sites inland from the
distribution of Lithocarpus). Although Lithocarpus is tolerant of
shade and high moisture (Roy 1957) Chrysolepis is apparently more
tolerant of moisture. At the BLN site Chrysolepis was found at its
greatest importance values with Lithocarpus totally lacking from the
transect. This was surprising because Lithocarpus was common on
surrounding adjacent higher elevation, more xeric and exposed slopes,
suggesting that the shady, mesic BLN location with its high impor-
tance of Jaxus, an indicator of very mesic conditions (Whittaker
1960, Atzet and Wheeler 1982), was too mesic to support Litho-
carpus (Fig. 4). However, the relatively high I.V. of Lithocarpus
saplings at the shady, high understory cover GM transect indicates
that tanoak may be slightly more shade tolerant than Chrysolepis
saplings.

In addition to its shade and moisture tolerance, Chrysolepis is

302 MADRONO [Vol. 35

© he

Fic. 4. Lower eT jee Pseudotsuga— Chrysolepis forest with dense cover of
Chrysolepis saplings and Taxus brevifolia, BLN site.

considerably more cold- and/or snow-tolerant than Lithocarpus, or
Arbutus. This is supported in the data on elevational distribution
and clearly indicated by the presence of tree-sized Chrysolepis on
north-facing slopes up to 1600 m in elevation, whereas Lithocarpus
and Arbutus both lose their tree form at ca. 1220 m. The archi-
tecture of both Lithocarpus and Arbutus is very different from Chry-
solepis and provides a clue to their low densities or absence in the
upper elevations of this zone. Both Arbutus and Lithocarpus have
relatively broad, plane or concave leaves with typically ascending
branches of mature trees. The outlines of uncrowded mature Litho-
carpus and Arbutus tend to be relatively broad at the top and at best
cylindrical. Conversely, Chrysolepis has typically longitudinally re-
flexed, convex leaves that are relatively narrow. In addition, mature
trees of Chrysolepis typically have principal branches deflexed, and
uncrowded mature trees tend to have a conical shape, similar to
middle aged Pseudotsuga or Abies concolor. The net result is that
the snow shedding abilities of the tree form of C. chrysophylla are
superior to those of all other sclerophyllous hardwoods in California
(Fig. 5). Although shrubby Quercus chrysolepis may occur at higher
elevations (over 1800 m), these are always southerly exposures where
snow tends not to accumulate and melts quickly. Thus, Chrysolepis
appears to be pre-adapted as the predominant sclerophyll of the
mesic upper elevations of the Pseudotsuga-hardwood zone.
Asexual versus sexual reproduction. The giant chinquapin has been

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA 303

< 3 } ae Va ; “Pw MLS is

Fic. 5. A 17m tall Chrysolepis at ca. 1550 m surrounded by Abies concolor forest.
Snow depth ca. 45 cm, Rough Gulch, 13 November 1984.

considered a poor and irregular seeder (mast seeding reported at 2—
5 year intervals) with resprouting its main mode of regeneration
(Roof 1969, 1970, Atzet and Wheeler 1982, McDonald et al. 1983,
McKee ms.). In fact, McDonald et al. (1983) report giant chinquapin
as having the poorest germination rate of all hardwoods in the Klam-
ath Province (14-53%). However, it is able to sexually establish itself
under mesic conditions in shade or partial shade at all three of the
sample areas in this report. Within the three sites it appears to require
a more mesic seed-bed than does Lithocarpus and does not establish
well from seed on upper slopes, ridges, or southerly exposures except
perhaps during an abnormally moist series of years. The density of

304 MADRONO [Vol. 35

seedlings in some areas without numerous saplings (e.g., RCN) sug-
gests that the survivorship of seedlings at those sites is low. The site
BLN indicates the affinity the species has for shady, mesic nursery
conditions, suggesting that under more widespread mesic condi-
tions in the Miocene (e.g., Axelrod 1977) it was a more common
tree, more tolerant of extremely mesic conditions than Lithocarpus,
which today appears to prefer summer fog or maritime influence,
but is able to tolerate drier slope aspects. The widespread restriction
to northerly-facing slopes is probably best explained by the long-
lasting and year-round moisture requirements for seedling estab-
lishment.

Once established, however, giant chinquapin is able to tolerate
relatively high moisture stress, evidenced by the presence of stands
on ridgetops and upper slopes. Its drought tolerance as a mature tree
and its ability to resprout vigorously have added greatly to its present
day ecological amplitude.

Chrysolepis reproduces asexually by adventitious budding from
stumps and basal burls after fires or other disturbance (e.g., wind
and snow damage, logging, and browsing by herbivores). Transect
BUS is a good example of a resprout xeric type which probably
originated from seed dispersed from adjacent north slope stands
long ago during a wet regime and is maintained through vigorous
post-fire resprouting (the last fire at BUS was ca. 1936). Resprout
stems averaged 2-3 per clump (with a maximum of 7) and ca. 28
cm dbh. The largest reprouts were ca. 46 cm dbh and ca. 24 m tall.
This contrasts sharply with typical seed-originated stems on n. slopes
which average ca. 140 years for 46 cm dbh.

Longevity versus reproductive ability. Trees average more stems
per ha on mesic sites, but the longest-living trees and the largest
individuals occur at more xeric sites such as mid-slopes at Rough-
Chinquapin. Ages of mature chinquapins are often difficult to obtain
due to heart rot and indistinct annual rings. However, the few I
successfully aged indicated specimens on north slopes grow relatively
slowly and uniformly, attaining diameters of ca. 60 cm in ca. 210-
260 years. The oldest Chrysolepis stems (those at Rough-Chinquapin
ca. 122 cm dbh) had estimated ages of 400-500 years. The Bridge
Creek and Goose Creek sites had fewer trees in the large size classes
than Rough-Chinquapin, the largest measured there were ca. 81 cm
dbh (Bridge, Fig. 6) and 70 cm dbh (Goose). Heart rot was more
prevalent at Bridge and Goose creeks and may be the reason for
fewer large trees at both sites. It appears, from the lack of young
trees on many of the upper slope and xeric sites, that sexual repro-
duction there is sporadic with major bursts perhaps once every few
hundred years. Yet, the more xeric conditions may be less conducive
to fungal damage to heartwood, apparently the main cause of death
in mature trees of undisturbed forest.

1988] KEELER-WOLF: CHRYSOLEPIS CHRYSOPHYLLA 305

4 ‘-

Fic. 6. One of the largest Chrysolepis individuals noted at the Bridge Creek study
site (transect BMN). This tree (in center) is ca. 35 m in height with a dbh of 75 cm.
Pseudotsuga to right is 86 cm dbh.

Successional versus climax species? Atzet and Wheeler (1982) and
McKee (ms.) consider C. chrysophylla to be an early successional
species only. However, contrary to these reports, giant chinquapin
does not necessarily decline in importance in late stages of succes-
sion. For example, the Bridge Creek lower valley (BLV) and lower
north-facing (BLN) sites were last burned by ground fires some 150-
165 years ago (evidence from aging scarred and singed trees) and
many of the canopy Pseudotsuga are over 500 years old at both sites,
yet Chrysolepis has the highest density of seed-originated saplings
and small trees in the understory and is represented at both sites by

306 MADRONO [Vol. 35

the highest densities for trees of any transect. No correlation exists
between total cover of all species and chinquapin cover, suggesting
that successional stage of the forest has nothing to do with Chry-
solepis density or cover.

On the three sites stems of giant chinquapins over 35 cm dbh
appear to tolerate ground fires better than many similar-sized hard-
woods (Quercus spp., Acer macrophyllum, Lithocarpus). At the
Rough-Chinquapin site, where the largest chinquapin trees were
noted, the last major fire occurred ca. 75-80 years ago and virtually
all Chrysolepis over 45 cm dbh had at least small fire scars which
were all or partially healed. The largest specimens (between 91 and
122 cm dbh) had cat-face scars and large sections of rotten core,
indicating repeated fire damage. Most of the fire-scarred trees in
mesic areas had single trunks, even those that were repeatedly dam-
aged.

The successional reputation of giant chinquapin appears to come
primarily from its ability to stump sprout following logging or crown
fires (McKee ms.). Under these conditions it may outstrip young
conifer growth for several years.

Atzet and Wheeler (1982, 1984) consider Lithocarpus as the major
climax species where both soil and atmospheric moisture is plentiful
at mid-slopes and elevations in the Siskiyou Mountains. Average
elevation for their tanoak series is 945 m. Above the tanoak and
below the white fir series (defined by Atzet and Wheeler as the area
where Abies concolor dominates the sapling and seedling layers) lies
the zone where Chrysolepis predominates. This zone is limited, but
under the appropriate mesic conditions such as at transects BMN,
BLN, and north slopes between 750 and 950 m at Rough-Chin-
quapin it may be considered as the hypothetical climax species based
on its ability to reproduce and come to dominate in undisturbed
situations.

ACKNOWLEDGMENTS

Many thanks to P. McDonald, D. Thornburgh, J. Griffin, and D. Keil for improving
the manuscript. I thank A. McKee and J. Franklin for unpublished information. This
research was funded jointly by the U.S. Forest Service Pacific Southwest Forest and
Range Experiment Station, Berkeley, CA, and the U.S. Forest Service Regional Office,
San Francisco, CA, and also by the Shasta-Trinity National Forest, and by Friends
of the Chinquapin.

LITERATURE CITED

ATzET, T. and D. L. WHEELER. 1982. Historical and ecological perspectives on fire
activity in the Klamath Geological Province of the Rogue River and Siskiyou
National Forests. U.S.D.A. For. Serv. Pacific Northwest Region, Portland, OR.

and . 1984. Preliminary plant associations of the Siskiyou Mountains
Province. U.S.D.A. For. Serv. Pacific Northwest Region, Portland, OR.

AXELROD, D. 1977. Outline history of California vegetation. Jn M. Barbour and J.
Major, eds., Terrestrial vegetation of California. Wiley-Interscience, New York.

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BRAY, J. R.and J. T. Curtis. 1957. An ordination of the upland forest communities
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Donato, M., C. BARNES, R. COLMAN, W. ERnsT, and M. KAys. 1982. Geologic
map of the Marble Mountain Wilderness, Siskiyou County, California. U.S.G:S.,
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FRANKLIN, J. and C. DyRNEss. 1973. Natural vegetation of Oregon and Washington.
U.S.D.A. For. Serv. Gen. Tech. Rep. PNW-8.

GRIFFIN, J. R. and W. B. CRITCHFIELD. 1972. The distribution of forest trees in
California. U.S.D.A. For. Serv. Res. Paper PSW-82.

HOLLAND, R. F. 1986. Preliminary descriptions of the terrestrial natural commu-
nities of California. Calif. Dept. Fish and Game, Sacramento, unpublished
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IRwIN, W. P. 1966. Geology of the Klamath Mountains Province. Jn E. Bailey, ed.,
Geology of Northern California. Calif. Div. Mines and Geol. Bull. 190.

1981. Tectonic accretion of the Klamath Mountains. Jn W. G. Ernst, ed.,
The geotectonic development of California. Prentice-Hall, Englewood Cliffs, NJ.

KEELER-WOLF, T. 1984. Ecological evaluation of the Rough Gulch drainage with
comparisons to the adjacent Chinquapin and Yolla Bolla Candidate Research
Natural Areas, Shasta-Trinity National Forests, California. Unpublished report
on file at Shasta-Trinity National Forest, Redding, CA.

1985. Ecological survey of the proposed Bridge Creek Research Natural

Area, Klamath National Forest, Siskiyou County, California. Unpublished report

on file at U.S.D.A. For. Serv. Pacific Southwest For. and Range Exp. Sta., Berke-

ley, CA.

1987a. An ecological survey of the proposed Upper Goose Creek Research

Natural Area, Six Rivers National Forest, Del Norte County, California. Un-

published report on file at U.S.D.A. For. Serv. Pacific Southwest For. and Range

Exp. Sta., Berkeley, CA.

. 1987b. An ecological survey of the proposed Pearch Creek Research Natural

Area, Six Rivers National Forest, Humboldt County, California. Unpublished

report on file at U.S.D.A. For. Serv. Pacific Southwest For. and Range Exp. Sta.,

Berkeley, CA.

. m.s. Asummary of ecological surveys conducted on Forest Service Research
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LITTLE, E. L., JR. 1971. Atlas of United States trees. Vol. 1. Conifers and important
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308 MADRONO [Vol. 35

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and 1977b. An ecological survey of the proposed Specimen Creek
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TAYLOR, D. 1975. Site evaluation: Yolla Bolla Research Natural Area. Unpublished
report on file at U.S.D.A. For. Serv. Pacific Southwest For. and Range Exp. Sta.,
Berkeley, CA.

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THORNBURGH, D. A. 1981. An ecological survey of the proposed Ruth Research
Natural Area, Six Rivers National Forest, California. Unpublished report on file
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. 1987. An ecological survey of the proposed Hennessy Ridge Research
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ANNOUNCEMENTS

NEw PuBLICATIONS

BARBOUR, M. G. and J. MAJor (eds.), Zerrestrial vegetation of Califor-
nia, revised ed., California Native Plant Society, 909 12th St., Suite
116, Sacramento, CA 95814, Jan 1988, 1036 pp., with map (A. W.
Kichler, ‘““Vegetation map of California’) in pocket, illus., ISBN

0-943460- 13-1 (hardbound), $50.00. [With 26 chapters by 30 authors,
a limited revision of the 1977 book.]

BEETLE, A. ‘A., and collaborators, Las Gramineas de México, vol. 2,
Secretaria de Agricultura y Recursos Hidraulicos [México, D.F.], 1987,
344 pp., illus. (some color), no ISBN, paperbound, gratis (from CO-
TECOCA, S.A.R.H. Manzanillo #83, Desp. 802, C.P. 06760, México,
D.F.). [Arrangement of genera and species alphabetical; with ca. 40
genera: Bambusa through Distichlis (vol. 1 on the “A” genera ap-
peared in 1983).]

THE HIGH ELEVATION FLORA OF
MOUNT ST. HELENS, WASHINGTON

ROGER DEL MORAL and DAvID M. Woop!
Department of Botany (KB-15),
University of Washington, Seattle 98195

ABSTRACT

The subalpine and alpine flora of Mount St. Helens is estimated to consist of about
95 species of vascular plants of which over 20 may have become locally extinct in
1980. In contrast, three nearby volcanoes contain two to three times as many species.
The causes for this limited flora include intense and frequent volcanism that has
locally extirpated some species and prevented others from reinvading. Mount St.
Helens is very young and may not have existed when dispersal for alpine species was
facilitated by full glacial conditions. It is isolated by 50 to 80 km of inhospitable
lowlands from seed sources on other volcanoes. The area above treeline is small and
lacks mesic and hydric habitats, thereby excluding many species capable of dispersing,
but incapable of finding a suitable habitat. Mount St. Helens is biogeographically
analogous to an oceanic island recently emerged from the sea, distant from potential
seed source pools, and suffering the trauma of repeated catastrophes.

The eruptive history of Mount St. Helens is rich and well docu-
mented (Yamaguchi 1986) with at least eight major events occurring
during the last 400 years. Prior to the eruption of 18 May 1980, this
young volcano was the smallest and lowest (2950 m) of the volcanoes
of the Pacific Northwest. The treeline of Mount St. Helens was
abnormally low, ranging from about 1150 to 1350 m, depending on
aspect (Lawrence 1938). Its elevation was reduced to 2550 m in
1980 by a huge debris avalanche and explosive lateral blast (see
Rosenfeld 1980 for a description of events). In contrast, timberlines
on neighboring volcanoes are at about 1800 m. Mount St. Helens
is located at 46°12'N, 122°11'W and is 70 km north of Mount Hood
(3428 m), 80 km west of Mount Adams (3752 m) and 52 km south-
southwest of Mount Rainier (4393 m).

Prior to 1980, studies of subalpine vegetation on Mount St. Helens
included only qualitative descriptions of xeric parkland and meadow
communities developed on lava flows, pyroclastic flows, and mud-
flows (lahars) less than 500 years old (Crandell et al. 1975, Hoblitt
et al. 1980). Unfortunately, there are no detailed floristic or vege-
tational studies that describe vegetation above treeline prior to the
1980 eruption.

This study evaluates what was known of the flora of Mount St.

' Present address: Wheaton College, Norton, MA 02766.

MADRONO, Vol. 35, No. 4, pp. 309-319, 1988

310 MADRONO [Vol. 35

Helens prior to 1980, compares it to relevant floras of surrounding
volcanoes, discusses reasons for its depauperate nature, and suggests
that the most recent eruption has produced numerous local extinc-
tions.

METHODS

The subalpine and alpine flora and meadow vegetation on the
slopes of Mount St. Helens were poorly known prior to 1980. We
used Hitchcock and Cronquist (1973) and Kruckeberg (1987) to
develop an estimate of the flora prior to the massive eruptions that
devastated the north flank of the volcano. The general characteristics
of pre-eruption high elevation communities on Mount St. Helens
can be inferred from the qualitative descriptions and checklists found
in Piper (1906), St. John (1976, describing a collecting trip in 1925),
and Kruckeberg (1987, describing a 3-day trip in 1979). In order to
compare this subalpine flora with those of surrounding volcanoes,
the following sources were consulted: for Mount Rainier, Dunwiddie
(1983); for Mount Adams, Riley (1986); and for Mount Hood, Bur-
nett (1986). Only species from these checklists likely to occur in
subalpine or alpine habitats are included. Although Riley (1986) and
Burnett (1986) excluded graminoids, we have included these families
based on habitat descriptions in Hitchcock and Cronquist (1973)
and personal observations.

Between 1980 and 1987 the first author investigated the flora
above 1200 m for an average of 19 days per year. The second author
averaged 21 days on the mountain each year between 1983 and 1987.
Voucher specimens were prepared for all species encountered and
are deposited in WTU.

Species were tabulated in several ways. All species encountered
by any primary source on Mount St. Helens are listed by family.
Species immigration and potential local extinction are thus indi-
cated. Dispersal mechanisms are inferred from morphology (see
Wood and del Moral 1987). Likely reasons for exclusion from the
flora are estimated from knowledge of dispersal and habitat require-
ments.

In order to estimate the degree of disharmony in the Mount St.
Helens flora, species were aggregated by plant family for each of the
four volcanoes compared. Disharmony refers to an unbalanced dis-
tribution of species per family, relative to the flora of the region. A
harmonious flora on Mount St. Helens would have about the same
proportion of its species distributed among the families as on the
other volcanoes.

RESULTS

The flora. The high elevation flora of Mount St. Helens is ex-
tremely poor, consisting of no endemics (St. John 1976, Kruckeberg
1987) and dominated by common species found at most high ele-

1988] DEL MORAL AND WOOD: MOUNT ST. HELENS 311

vations in the Pacific Northwest. Table 1 lists species reported for
the higher elevation vegetation (Piper 1906, St. John 1976, del Moral
1983, del Moral and Wood pers. obs. 1980-87, Kruckeberg 1987).
Species noted by Kruckeberg and not by St. John may be species
that have invaded since 1925. The flora is very poorly-represented
in those families dominated by mesophytes and well-represented in
taxa with good dispersal mechanisms. Nomenclature follows that of
Hitchcock and Cronquist (1973).

The 23 species noted by St. John or Kruckeberg, but not by del
Moral, may be species that have been eliminated as a consequence
of the 1980 eruption. (Negative evidence, in this case the non-ob-
servation of species, is always problematic.) These are indicated by
an asterisk in Table 1 and include Botrychium lanceolatum, Lewisia
columbiana, Caltha biflora, Trautvetteria caroliniensis, Heuchera
micrantha, Saxifraga arguta, Phyllodoce glanduliflora, Dodecatheon
Jeffreyi, Collomia debilis, Penstemon davidsonii, Valeriana sitchen-
sis, Aster alpigenus, Erigeron peregrinus, and Luzula divaricata.

The pre-eruptive subalpine and alpine flora on Mount St. Helens
was dominated by species common to other Northwest volcanoes.
Dominants included Lupinus lepidus, L. latifolius, Eriogonum pyro-
lifolium, Polygonum newberryi, Luetkea pectinata, Saxifraga tol-
miei, Phyllodoce empetriformis, Arctostaphylos uva-ursi, Juncus par-
ryi, Spraguea umbellata, and Castilleja miniata. Today, on the
southern slopes of Mount St. Helens, these species are common
meadow plants.

Impoverishment and disharmony. We are unaware of any biogeo-
graphic studies of non-equilibrium conditions on terrestrial islands.
Table 2 summarizes the depauperate and disharmonious nature of
the Mount St. Helens flora compared to the floras of Mount Rainier,
Mount Adams, and Mount Hood. Subalpine and alpine species rich-
ness 1S estimated to be as follows: for Mount St. Helens, 95 species
(including 23 species, 24%, now possibly extinct on the cone); for
Mount Hood, 185 (Burnett 1986); for Mount Adams, 198 (Riley
1986); and for Mount Rainier, 261 (Dunwiddie 1983).

Prior to 1980, Mount St. Helens’ flora was only about '4 of that
of Mount Rainier. Families with fewer than '4 as many species are
under-represented, whereas those with more than 3 as many are
over-represented. Based on this criterion, the following families are
under-represented: Polypodiaceae, Salicaceae, Caryophyllaceae, Ra-
nunculaceae, Brassicaceae, Saxifragaceae, Fabaceae, Onagraceae,
Apiaceae, Boraginaceae, Scrophulariaceae, and Liliaceae. The Ly-
copodiaceae, Polygonaceae, Juncaceae, Cyperaceae, and Poaceae are
Over-represented.

DISCUSSION AND CONCLUSIONS

We postulate four reasons for low species richness and dishar-
mony: 1) frequent disturbance causing relatively high local extinc-

312 MADRONO [Vol. 35

TABLE 1. PLANT SPECIES COLLECTED OR OBSERVED ABOVE TIMBERLINE ON THE
FLANKS OF Mount St. HELENS. (Nomenclature has been modified to conform to
Hitchcock and Cronquist, 1973. Piper’s observations are from a table of “arctic-
alpine species” plus higher elevation species from a table of ‘““Hudsonian”’ species.)
X = Species observed. * = Species not observed by authors or other field workers
since 1980. '“‘Characteristic species”; Piper’s list was not exhaustive. *Species listed
by these authors that are clearly not from subalpine habitats and tree species are
omitted.

Plant family Piper’, St. John’, Krucke- del Moral,
Species ca. 1900 1925 berg”, 1979 1980-1986

PTERIDOPHYTES

Lycopodiaceae

Lycopodium sitchense 4 XxX 4 x

L. annotinum — xX
Ophioglossaceae

Botrychium lanceolatum — x xX *
Polypodiaceae

Cryptogramma crispa — xX xX xX
Selaginellaceae

Selaginella wallacei — 4 — *

GYMNOSPERMS
Cupressaceae
Juniperus communis X xX xX xX

ANGIOSPERMS— DICOTS
Aplaceae
Lomatium martindalei xX

~*~
~<
~

Asteraceae

Achillea millefolium
Agoseris aurantiaca

A. glauca

Anaphalis margaritacea
Antennaria microphylla
A. lanata

Arnica latifolia

A. cordifolia

Aster alpigenus

A. ledophyllus

Erigeron peregrinus
Eriophyllum lanatum
Hieracium albiflorum
H. gracile

Luina hypoleuca
Microseris alpestris

x |
|

|
| XK KX |

1 | xx 1 KI XK
| * |

KK mM KK KL KI KE
KKK KK EK EK KK KKM KM

xxKK XK |

x |

Campanulaceae
Campanula rotundifolia —

Sd
|
*

Caryophyllaceae
Silene parryi _ XxX ~ XxX

1988] DEL MORAL AND WOOD: MOUNT ST. HELENS 313

TABLE |. CONTINUED.

Plant family Piper’, St. John’, Krucke- del Moral,
Species ca. 1900 1925 berg”, 1979 1980-1986

Crassulaceae
Sedum oreganum

|
x
*

Ericaceae

Arctostaphylos uva-ursi
Phyllodoce empetriformis
Phyllodoce glanduliflora
Vaccinium scoparium
Vaccinium membranaceum

| xxx |
| Xxx

| xxx
xx KM

Fabaceae

Lupinus latifolius — xX

L. lepidus var. lobbii xX 4
Gentianaceae

Gentiana calycosa — — _ x
Hydrophyllaceae

Phacelia hastata _ xX 4 *
Onagraceae

Epilobium angustifolium — XxX xX
E. alpinum — xX

xx

Polemoniaceae
Collomia debilis — x =
Linanthastrum nuttallii = = _
Phlox diffusa xX x x
Polygonaceae

Eriogonum pyrolifolium 4
E. ovalifolium nivale _
X

x xX *

Polygonum newberryi
P. minimum

KKK x

Portulacaceae

Lewisia columbiana —

Spraguea umbellata 4 xX xX xX
Primulaceae

Dodecatheon jeffreyi — 4 _ *
Ranunculaceae

Aquilegia formosa —
Caltha biflora
Trautvetteria caroliniensis

~
|
x * XX

|
x |

Rosaceae
Fragaria virginiana —
Luetkea pectinata —
Potentilla arguta
Rubus lasiococcus
Sibbaldia procumbens
Sorbus sitchensis
Spiraea densiflora

xxx mK | KM
xx KK | KM
KKK KKK

xx | XX |

314

Plant family
Species
Salicaceae
Salix barclayi

Saxifragaceae
Heuchera micrantha
Saxifraga arguta
S. tolmiei

Scrophulariaceae

Castilleja miniata
Penstemon cardwellii
P. confertus

MADRONO

TABLE 1.

Piper’,
ca. 1900

P. davidsonii var. menziesii 4

P. serrulatus

Valerianaceae
Valeriana sitchensis

Violaceae
Viola adunca

Cyperaceae

Carex mertensii
C. pachystachya
C. phaeocephala
C. rossii
C. spectabilis
Juncaceae
Juncus parryi
J. drummondii
Luzula divaricata
L. piperi
Liliaceae
Smilacina racemosa
S. stellata
Xerophyllum tenax
Poaceae
Agrostis diegoensis
A. exarata
A. variabilis
Bromus carinatus

CONTINUED.

St. John’,
1925

x

xxx

x1 x x

[Vol. 35

Krucke- del Moral,
berg’, 1979 1980-1986

ANGIOSPERMS— MONOCOTS

Calamagrostis sesquiflora —

Danthonia intermedia
D. spicata

Festuca occidentalis
Festuca viridula
Phleum alpinum

Poa incurva

Sitanion hystrix

S. jubatum

Stipa occidentalis
Trisetum spicatum

xxx | XM

| x | *

xx x

KK 1 KKK KK KK KM |

xX xX
xX
= *
xX xX
xX xX
xX xX
_ xX
= *
— *
xX *
— xX
—_— x
_— xX
— X
> & xX
x xX
xX xX
_— xX
= *
» xX
xX xX
x xX
— xX
xX xX
— xX
= *
— xX
= *
xX
= *
_ xX
xX *
xX xX
— xX
xX xX
x xX
— xX
— xX

1988] DEL MORAL AND WOOD: MOUNT ST. HELENS 315

TABLE 2. NUMBER OF SPECIES IN PLANT FAMILIES ON FOUR VOLCANOES: MOUNT
St. HELENS, MOUNT RAINIER (Dunwiddie 1983), MoUNT ADAMS (Riley 1986), AND
Mount Hoop (Burnett 1986). Parenthetical values are post-1980 estimates, where
different. 'Lycopodiaceae and Selaginellaceae. ?Ophioglossaceae and Polypodiaceae.

Mount Mount Mount Mount
Family St. Helens Rainier Adams Hood
Fern allies! 3 (2) 4 3 3
Ferns? 2 (1) 6 5 3
Cupressaceae I l 1 l
ANGIOSPERMS-— DICOTS
Apiaceae l 5 5 6
Asteraceae 16 (14) 50 35 27
Boraginaceae 0 2 0) 2
Brassicaceae 0 14 8 2)
Campanulaceae 1 l l 1
Caryophyllaceae 1 9 5 4
Crassulaceae 1 (O) 3 2 l
Droseraceae 0) l l l
Ericaceae 5 (4) et 9 7
Fabaceae 2 ui 4 5
Gentianaceae 1 l l l
Hippuridaceae 0 1 0 0
Hydrophyllaceae 1 (0) 3 2 2
Hypericaceae 0 1 l 1
Onagraceae 2 11 6 8
Polemoniaceae 3 (2) 9 fs a
Polygonaceae 4 9 7 8
Portulacaceae 2 (1) 5 3 2
Primulaceae 1 (O) 3 2 2
Ranunculaceae 32) 13 8 9
Rosaceae 7 1) 12 10
Salicaceae 1 4 2 3
Saxifragaceae 3 (1) 15 9 8
Scrophulariaceae 5 (3) 20 18 17
Valerianaceae 1 (0) 1 | l
Violaceae l 1 l l
ANGIOSPERMS—MONOCOTS

Cyperaceae 5 13 10 9
Juncaceae 4 (3) 6 4 3
Liliaceae 3 9 5 7
Poaceae 15 (11) 20 19 19
Orchidaceae 0 D. l l

Totals 95 (72) 276 198 185

tion rates; 2) low immigration rates due to isolation and ineffective
dispersal mechanisms (e.g., mammals, ants, water, and gravity); 3)
the immaturity of the volcano resulting in poor soils; and 4) a lack
of mesic and hydric habitats.

The flora may have increased during the interval between major
eruptions (1852 to 1980). For example, Piper (1906) listed only 45
species for the upper slopes of Mount St. Helens. Lawrence (1938,

316 MADRONO [Vol. 35

1939) found 80 vascular plant species on pumice between 1200 and
2200 m. Lawrence stated that at least some were recent immigrants
and that new species should be expected to invade each year. Krucke-
berg (1987) stated that 70 vascular plants species common in other
subalpine and alpine regions of the Cascades do not occur on Mount
St. Helens. In contrast, species in groups with effective long-distance
dispersal mechanisms, such as lycopods, composites, grasses, sedges
and rushes, are well represented. The vegetation of subalpine mead-
ows on Mount St. Helens is characterized by widely distributed
species common to Northwestern volcanoes, whereas many other
common species are lacking.

Species strangely absent from Mount St. Helens are noted in Table
3, along with potential reasons for their absence, related to those
factors stated above. In addition, several species may have suc-
cumbed to the most recent disturbance. The list is meant to illustrate
reasons potentially excluding a species from a habitat and should
not be construed as definitive. Species from the list of Kruckeberg
(1987) (except for three known to exist in 1979) have been catego-
rized as follows: 18 of 67 species appear to be absent primarily
because their dispersal mechanisms are effective for only short dis-
tances. The bulk of the species (27 of 67) appear limited both by
inefhicient dispersal and the absence or small size of suitable habitats.
The true alpine zone is small on this volcano, especially since 1980.
Lush meadow habitats, such as the extensive meadows near Paradise
on Mount Rainier, have never been common and today are virtually
absent. Glaciers that existed on steep slopes were nearly completely
removed. Their remnants apparently did not support any distinct
wetland vegetation. Bogs do not exist on or near the cone. Therefore
lush meadow species (Anemone occidentalis) and bog dwellers (Kal-
mia microphylla) do not occur. A few species, such as Allium cer-
nuum, may tolerate drought, but require better developed soils than
occur. Twenty species have effective dispersal mechanisms, but are
absent primarily for reasons related to their special habitat require-
ments. The lack ofa sizable, stable alpine habitat may exclude genera
like Empetrum and Dryas. Wet meadow species are absent because
of insufficient moisture. The absence of stable, dry soil and very acid
soils may limit other species. Species such as Vaccinium deliciosum
are expected; their absence may be due only to chance. Species other
than those mentioned by Kruckeberg (1987) might be expected, but
are lacking for one or more of the reasons discussed below.

Although none of the studies cited above is complete, each implies
a depauperate flora. Incomplete collecting may account for some
gaps in earlier studies since these concentrated on the north slopes.
However, species confined to the north slope may indeed now be
absent from the volcano.

It is likely that low richness and disharmony of the flora above

1988] DEL MORAL AND WOOD: MOUNT ST. HELENS 37

TABLE 3. SUMMARY OF PROBABLE CAUSES FOR THE ABSENCE OF COMMON SPECIES
FROM MOounrtT ST. HELENS VOLCANO.

Reason Total Examples

Primarily dispersal 18
No special mechanism 12 Arabis spp., Draba aureola, Arenaria spp., Saxi-
fraga spp., Smelowskia spp., Thlaspi fendleri,
Phacelia sericea, Polemonium elegans

Mammals 6 Agropyron spp., Deschampsia atropurpurea, Erig-
eron spp.
Dispersal and habitat 27
Lack of alpine habitat 1 Oxyria digyna
Too dry 24 Anemone spp., Kalmia microphylla, Erythronium

montanum, Ranunculus spp., Dodecatheon jef-
freyi, Douglasia laevigata, Solidago multiradia-
ta, Veronica cusickii, Silene spp., Thalictrum

spp.
Lack of suitable soil 2 Allium cernuum, Spiraea betulifolia
Primarily habitat 20
Lack of alpine habitat 6 Artemisia norvegica, Silene acaulis, Salix spp..,
Dryas octopetala, Empetrum nigrum
Too dry 10 = Epilobium latifolium, Polypodium hesperium, Ha-
benaria dilatata, Aster alpigenus, Pedicularis
spp., Senecio triangularis, Saussurea americana
Lack of suitable soil 4 Sedum spp., Cheilanthes gracillima, Cassiope
mertensiana
Chance 2 Vaccinium deliciosum, Haplopappus lyallii

timberline on Mount St. Helens result from a combination of several
factors. Frequent eruptive disturbances probably caused the local
extinction of many species. Caltha biflora, Dodecatheon jeffreyi, and
Festuca viridula are among species probably extirpated from the
cone in 1980. These are species not known from the south half of
the cone, and they generally occur in mesic habitats.

The youth and small size of the volcano have several important
consequences. Some species may have reached other alpine areas
during glacial maxima at a time when Mount St. Helens either did
not exist or lacked high elevation habitats. Poorly developed sub-
strates preclude the development of habitats for mesophytic species.
Over time, soil development will occur so that species common, for
example, at Sunrise on Mount Rainier, may be able to grow. Such
species include Potentilla flabellifolia and Veronica cusickii. Many
families under-represented on Mount St. Helens are primarily me-
sophytes.

Restricted habitats subject the flora to several processes likely to
reduce richness. 1) Disturbances may totally eliminate certain hab-
itats; 2) existing habitats are smaller and less diverse and therefore
less likely to have as many species due to simple species-area effects;

318 MADRONO [Vol. 35

3) populations of rare species will be small and of limited distribution
and therefore more susceptible to elimination; and 4) small habitats
are less likely to intercept invading seeds than large habitats. In
addition, species unable to withstand burial by tephra and other
mild impacts may succumb whereas more tolerant species survive.
Long distances from seed sources also contribute to limited recol-
onization from surrounding areas, despite the 123-year dormant
period preceding the 1980 eruption. Nearby subalpine and alpine
communities on the Old Cascades nonvolcanic landscape, notably
on Mount Margaret and Strawberry Mountain, probably were de-
pauperate because they were decimated by thick tephra deposited
in A.D. 1480 (Yamaguchi 1986). As a result, many seed sources of
the pre-1980 landscape may have been as distant as the nearest
volcanoes.

The frequency of major disturbances on Mount St. Helens is such
that the flora appears to be far below its equilibrium richness (see
Malanson 1984). Studies of isolated young woodlands (less than 350
years old) in Britain (Peterken and Game 1984) suggest first that
such woodlands are depauperate and disharmonious and second that
they acquire most of their species within 20 years. Even minor
isolation serves to restrict colonization rates dramatically.

The balance between frequent episodic local extinction and pre-
sumably gradual colonization on Mount St. Helens, combined with
small area and an immature landscape, has produced a monotonous
flora remarkably limited in richness. The 1980 eruptions may have
contributed to this impoverishment, with as many as 23 of 95 species
being eliminated in 1980.

ACKNOWLEDGMENTS

This paper is dedicated to the memory of Warren Tanaka, an ardent collector of
high elevation plants, who contributed in many ways to the studies of the regional
flora. We also are indebted to the many people who collected plants on Northwestern
volcanoes during the last century. A. R. Kruckeberg and M. F. Denton made valuable
comments on our manuscript; A. R. Kruckeberg also provided crucial reference data.
David Yamaguchi stimulated this undertaking. Funds for this study were provided
by N.S.F. grants DEB-81-07042 and BSR-84-07213.

LITERATURE CITED

BurRNETT, R. E. 1986. Flowering plants of the Mt. Hood Area. Mazama 67:1-12.

CRANDELL, D. R., D. R. MULLINEAUX, and R. MEYER. 1975. Mount St. Helens
volcano; recent and future behavior. Science 187:438-441.

DEL MoRAL, R. 1983. Initial recovery of subalpine vegetation on Mount St. Helens,
Washington. Amer. Midl. Naturalist 109:72-80.

and D. M. Woop. 1988. Dynamics of herbaceous vegetation recovery on
Mount St. Helens. Vegetatio 74:11-27.

DunwippiE, P. W. 1983. Checklist of vascular plants: Mount Rainier National
Park, PNW Nat. Parks & Forests Association.

1988] DEL MORAL AND WOOD: MOUNT ST. HELENS Slt

Hitcucock, C. L. and A. CRONQUIST. 1973. Flora of the Pacific Northwest. Univ.
Washington Press, Seattle.

Hos tT, R. P., D. R. CRANDELL, and D. R. MULLINEAUX. 1980. Mount St. Helens
eruptive behavior during the past 1500 years. Geol. Mag. 8:555—560.

KRUCKEBERG, A. R. 1987. Plant life on Mount St. Helens before 1980. Jn D. E.
Bilderback, ed., Mount St. Helens 1980, botanical consequences of the explosive
eruptions, p. 3-21. Univ. California Press, Berkeley.

LAWRENCE, D. B. 1938. Trees on the march. Mazama 20:49-54.

. 1939. Continuing research on the flora of Mount St. Helens. Mazama
21(12):56-60.

MALANSON, G. P. 1984. Intensity as a third factor of disturbance regime and its
effect on species diversity. Oikos 43:41 1-413.

PETERKEN, G. F. and M. GAME. 1984. Historical factors affecting the number and
distribution of vascular plant species in the woodlands of central Lincolnshire.
J. Ecol. 72:155-182.

Pirer, C. V. 1906. Flora of the state of Washington. Contr. U.S. Nat. Herb. Govt.
Printing Office, Washington, D.C.

RILEY, J. 1986. Mount Adams Wilderness plant list. Mimeographed report, U.S.
Forest Service.

ROSENFELD, C. L. 1980. Observations on the Mount St. Helens eruption. Amer.
Sci. 68:494—509.

St. JOHN, H. 1976. The flora of Mt. St. Helens, Washington. The Mountaineer
70(7):65-77.

Woop, D. M. and R. DEL MorRAL. 1987. Mechanisms of early primary succession
in subalpine habitats on Mount St. Helens. Ecology 68:780-790.

YAMAGUCHI, D. K. 1986. The development of old-growth Douglas-fir forests North-
east of Mount St. Helens, Washington, following an A.D. 1480 eruption. Ph.D.
dissertation, Univ. Washington, Seattle.

(Received 12 May 1987; revision accepted 27 Jul 1988.)

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What do we know?, International Union for Conservation of Nature
and Natural Resources, Avenue du Mont-Blanc, CH-1196 Gland,
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everywhere else. Arrangement by country, including for each data on
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GRIFFIN, J. R., P. M. MCDONALD, and P. C. Muick (compilers), Cali-
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CHROMOSOME NUMBERS IN THE ANNUAL
MUHLENBERGIA (POACEAE)

PAUL M. PETERSON!
Department of Botany and Ownbey Herbarium,
Washington State University, Pullman 99164

ABSTRACT

Chromosome numbers were obtained from field-collected microsporocytes and
root-tip preparations from greenhouse-grown specimens for 25 species of annual
Muhlenbergia sensu lato. Nine of these counts are first reports: M. annua (2n=20),
M. biloba (n=8), M. eludens (n=20), M. fragilis (n=10), M. pectinata (n=10), M.
schmitzii (n=20), M. shepherdii (n=8), M. sinuosa (n=10, 12), and M. tenuissima
(n=10). Besides sharing a haploid number of n=8, M. biloba and M. shepherdii also
have large chromosomes unlike those of the other species. Both appear to be misplaced
in Muhlenbergia: M. biloba is better treated as Bealia mexicana and M. shepherdii
probably should be transferred to Blepharoneuron.

RESUMEN

El numero de cromosomas de 25 especies de Muhlenbergia sensu lato se obtuvieron
de muestras obtenidas de esporocitos recogidos en el campo o de preparaciones de
apices de la raices de ejemplares cultivados en invernaderos. Se reporta por primera
ves el numero de cromosomas de 9 especies diferentes. Estas son las siguientes: M.
annua (2n=20), M. biloba (n=8), M. eludens (2n=20), M. fragilis (n=10), M. pectinata
(n=10), M. schmitzii (2n=20), M. shepherdii (n=8), M. sinuosa (n=10, 12), y M.
tenuissima (n=10). Muhlenbergia biloba y M. shepherdii se destacan de otras especies
al tener un numero haploide de 8 cromosomas y al tener cromosomas de tamano
mayor. Parece ser que ninguna de estas dos ultimas especies pertenecen en el género
Muhlenbergia: M. biloba se debe condiderar como Bealia mexicana y M. shepherdii
se debe transferir al genero Blepharoneuron.

Muhlenbergia Schreb. (Chloridoideae: Eragrostideae) comprises
over 160 species (Clayton and Renvoize 1986), most of which occur
in arid lands of the New World. Of the 32 annual species, 31 taxa
occur in Mexico, 14 of which also occur in the southwestern United
States, and a single species is restricted to Guatemala. Although
several alliances within the annuals are apparent, some groups ap-
pear to be paraphyletic and closely related to several perennial Muh-
lenbergia.

Chromosome counts reported here are part of a long-range sys-
tematic study on the annual species of Muhlenbergia (Peterson and
Rieseberg 1987, Peterson et al. 1988, Peterson 1989). Chromosome
counts for only 16 species had been reported (Myers 1947, Tateoka

' Present address: Department of Botany, National Museum of Natural History,
Smithsonian Institution, Washington, DC 20560.

MADRONO, Vol. 35, No. 4, pp. 320-324, 1988

1988] PETERSON: MUHLENBERGIA CHROMOSOMES 321

1962, Gould 1965, 1966, Reeder 1967, 1968, Gould and Soderstrom
1970, Pohl and Davidse 1971, Davidse and Pohl 1974, Hatch 1980).
I determined additional chromosome counts to help provide base-
line data for a systematic study of the group. Chromosome data may
also be valuable for inferring phylogeny and interpreting patterns of
evolution (Clausen et al. 1945).

MATERIALS AND METHODS

Floral buds were field collected and fixed in ethanol-acetic acid
(3:1, V:V) prior to storage under refrigeration in 70% ethanol. Meiot-
ic chromosome counts were obtained from aceto-carmine squashes
of pollen mother cells. Root tips were collected from greenhouse-
grown specimens and subsequently treated in 0.002 M 8-hydroxy-
quinoline (4 hr), ethanol-acetic acid fixative (2 hr), glusulase (45
min), and 0.2 N hydrochloric acid (2 min) before being squashed in
aceto-carmine (Soltis 1980). Representative cells were recorded with
sketches and photographed using a Nikon Biophot, phase-contrast
microscope, using 35 mm Kodak technical pan 2415 film. Chromo-
some number determinations were based on observations of 15 or
more cells from at least three individuals per population. Vouchers
of the plants counted are deposited in WS.

RESULTS

A total of 67 chromosome counts were made, representing 25
species of annual Muhlenbergia sensu lato. Chromosome number,
locations, and collection numbers are listed in Table | for each of
the 25 species. Chromosome numbers for the following nine species
are first reports: M@. annua (2n=20), M. biloba (n=8), M. eludens
(n=20), M. fragilis (n=10), M. pectinata (n=10), M. schmitzii (n=20),
M. shepherdii (n=8), M. sinuosa (n=10, 12), and M. tenuissima
(n=10). The chromosome numbers of M. brevis (n=10), M. ciliata
(n=10), M. minutissima (n=30), M. crispiseta (n=10), M. depau-
perata (n=10), M. diversiglumis (n=10), M. filiformis (n=9), M.
implicata (n=10), M. microsperma (n=10), M. ramulosa (n=10),
M. strictior (n=10), M. tenella (n=10), M. tenuifolia (n=20), M.
texana (n=20), and M. vaginata (n=9) are concordant with previous
reported counts (Meyers 1947, Tateoka 1962, Gould 1965, 1966,
Reeder 1967, 1968, Gould and Soderstrom 1970, Pohl and Davidse
1971, Davidse and Pohl 1974, Hatch 1980). Muhlenbergia sinuosa
has two different chromosome races with two n=12 populations
from Cochise and Santa Cruz counties, Arizona and two n=10 pop-
ulations from Grant County, New Mexico and Chihuahua, Mexico.
The meiotic chromosome count for M. pusilla (n=15) substantiated
a previous somatic count of 2n=30 for the same species (Reeder
1968). Some microsporocytes of three populations of M. strictior

322 MADRONO [Vol. 35

TABLE |. CHROMOSOME COUNTS IN THE ANNUAL Muhlenbergia SENSU LATO. Pop-
ulations are arranged alphabetically by species and locality. All collections are those
of the author and C. R. Annable. Vouchers are deposited in WS. Unless otherwise
noted, all counts were obtained from microsporocytes.

M. annua (Vasey) Swallen 2n=20. Mexico, Chihuahua, nw. of Hernandez Javales,
4053; Durango, w. of Navios, 4582 (root tip count).

M. biloba A. S. Hitche. n=8. Mexico, Durango, sw. of El Ojito, 4570.

M. brevis C. Goodding n=10. Mexico, Chihuahua, s. of Hernandez Javales, 4041; n.
of Cuesta Blanca, 4047; sw. of Madera, 405/; ne. of El Vergel, 4061; Durango, w.
of Rio Chico Crossing, 4094.

M. ciliata (H.B.K.) Kunth n=10. Mexico, Chihuahua, ne. of El Vergel, 4080 (some
cells at n=9); Durango, w. of Rio Chico Crossing, 4093; e. of El Salto, 4/19;
Michoacan, s. of Uruapan, 46/9; Sinaloa, nw. of Surutato, 4165.

M. crispiseta A. S. Hitchc. n=10. Mexico, Durango, sw. of El Ojito, 457].

M. depauperata Scribn. n=10. Mexico, Chihuahua, n. of Villa Matamoros, 4082; s.
of Villa Matamoros, 4083; Zacatecas, nw. of Fresnillo, 4596.

M. diversiglumis Trin. n=10. Mexico, Sinaloa, e. of Santa Lucia, 4/47.

M. eludens C. Reeder n=20. Mexico, Durango, w. of Rio Chico Crossing, 4096. USA,
AZ, Santa Cruz Co., sw. of Canelo, 40/8.

M. filiformis (Thurb.) Rydb. n=9. USA, AZ, Apache Co., e. of McNary, 3994; Wash-
ington, Klickitat Co., Wash. St. Salmon Hatchery, 3987.

M. fragilis Swallen n=10. Mexico, Chihuahua, w. of Parral, 4554. USA, AZ, Santa
Cruz Co., Sycamore Canyon, 4024.

M. implicata (H.B.K.) Kunth n=10. Mexico, Chihuahua, ne. of El Vergel, 4079;
Durango, s. of El Ojito, 4566; Oaxaca, se. of Sinaxtla, 4670.

M. microsperma (DC.) Kunth. n=10. USA, AZ, Santa Cruz Co., Sycamore Canyon,
4023, NV, Clark Co., Lake Mead, 3067.

M. minutissima (Steud.) Swallen [incl. M. confusa (Fourn.) Swallen] n=30. Mexico,
Chihuahua, sw. of Colonia Juarez, 4037; Durango, w. of Rio Chico Crossing, 4097;
Mexico, w. of Toluca, 4634. USA, AZ, Apache Co., n. of Sunrise Lake, 3998.

M. pectinata C. Goodding n=10. Mexico, Durango, s. of Durango, 4089; w. of El
Salto, 4/32: w. of La Ciudad, 4735, 4139, 4141; Sinaloa, s. of Surutato, 4/5/,
4152.

M. pusilla Steud. [incl. M. pulcherrima Scribn.] n=15. Mexico, Chiapas, nw. of Mo-
tozintla de Mendoza, 47/2; Chihuahua, sw. of El Vergel, 4073.

M. ramulosa (H.B.K.) Swallen [incl. MM. wolfi (Vasey) Rydb.] n=10. Mexico, Chi-
huahua, ne. of El Vergel, 4064. USA, AZ, Cochise Co., Rustler Campground, 40/ /.

M. schmitzii Hack. n=20. Mexico, Michoacan, w. of Ciudad Hidalgo, 4631.

M. shepherdi (Vasey) Swallen n=8. Mexico, Durango, w. of El Salto, 4/22, s. of El
Ojito, 4561.

M. sinuosa Swallen n=10, 12. Mexico, Chihuahua, w. of Tomochic, 4540 (n=10).
USA, AZ, Cochise Co., Rucker Lake, 40/3 (n=12); Santa Cruz Co., sw. of Canelo,
4020 (n=12); NM, Grant Co., San Lorenzo, 4008 (n=10).

M. strictior Scribn. n=10. Mexico, Chihuahua, s. of Hernandez Javales, 4039 (some
cells at n=9), 4043: w. of Tomochic, 4553; w. of La Junta, 4054 (some cells at
n=9); Durango, s. of El Ojito, 4563 (some cells at n=9); w. of Navios, 4584.

M. tenella (H.B.K.) Trin. n=10. Mexico, Chiapas, s. of Frontera Comalapa, 4704.

M. tenuifolia (H.B.K.) Trin. n=20. Mexico, Chihuahua, ne. of Parral, 4059; s. of Villa
Matamoros, 4085.

M. tenuissima (Presl) Kunth n=10. Mexico, Jalisco, s. of Yahualica, 4062.

M. texana Buckl. n=20. Mexico, Durango, w. of Navios, 4/08. USA, AZ, Santa Cruz
Co., sw. of Canelo, 4019; Sycamore Canyon, 4028.

M. vaginata Swallen n=9. Mexico, Durango, w. of El Salto, 4124, 4591; w. of Navios,
4587.

1988] PETERSON: MUHLENBERGIA CHROMOSOMES 325

and one population of M. ciliata exhibited irregularity (n=9) sug-
gesting facultative aneuploidy. However, most microsporocytes of
these populations had normal bivalents during meiosis (n=10). The
chromosomes of M. biloba and M. shepherdii were significantly
larger than those of any other species in this survey.

DISCUSSION

Twenty of the species investigated have a basic number of x=10
which is in agreement with the previously reported base number for
the genus (Darlington and Wylie 1956, Pohl and Mitchell 1965).
Muhlenbergia filiformis and M. vaginata have a base number of x=9
which suggests a close relationship with Sporobolus, also x=9. This
base number is perhaps a result of stabilized aneuploidy or dysploi-
dy. The haploid number of n=15 for M. pusilla is unusual in the
genus. Individuals of M. pusilla show tremendous morphological
variation within and among natural populations. Lemmas and awns
vary from 2.0—4.2 mm and 0-22 m long, respectively. The chro-
mosome number and varying morphological forms suggest that these
plants may be triploid and completely apomictic (Reeder 1968).

The two counts of n=8 for M. biloba and M. shepherdii, coupled
with their large chromosome size compare with other Muhlenbergia,
suggest that these taxa are misplaced in this genus. Muhlenbergia
biloba, originally described as Bealia mexicana by Scribner in Hack-
el (1890), is perhaps best retained in its own monotypic genus. Cy-
tologically, M. biloba shows close affinities with Dasyochloa Rydb.
and Erioneuron Nash, both x=8. Lemma morphology among these
taxa 1s very similar. All possess three-nerved, emarginate to bilobed,
and often awned lemmas with pilose hairs associated with either the
nerves, margins, and/or lower *3 of the lemma. However, M. biloba
[Bealia mexicana] differs from Dasyochloa and Erioneuron by being
single-flowered and annual in nature.

Muhlenbergia shepherdii seems more closely related to the mono-
typic genus Blepharoneuron Nash than to other members of Muhlen-
bergia. These two taxa share the following features: chromosome
number of n=8, strongly ribbed leaf blades, indistinguishable leaf
anatomy, 1-flowered spikelets, subequal glumes almost as long as
the floret, rounded lemmas with pilose margins and midnerve, and
paleas that are densely (appressed in M. shepherdii) villous on and
between the keel. I plan to transfer this species into Blepharoneuron.

The evidence from cytology, coupled with anatomy and mor-
phology suggests that the annual species of Muhlenbergia sensu lato
form at least three phylogenetically distinct lines. Muhlenbergia bi-
loba |Bealia mexicana] and M. [Blepharoneuron] shepherdii each
seem to form two distant but related lines, whereas all other members
with x=9, 10 form a possible third lineage.

324 MADRONO [Vol. 35

ACKNOWLEDGMENTS

This study was supported by grants from the National Science Foundation to Amy
Jean Gilmartin and PMP (BSR-8612611), Sigma Xi, and WSU. Special thanks are
given to Carol R. Annable for assistance in the field and discussions pertinent to this
manuscript. Amy Jean Gilmartin, David J. Keil, Askell Love, Douglas E. Soltis, and
an anonymous reviewer are greatfully acknowledged for significantly improving this
manuscript. I thank Raul Cano for preparing the Spanish abstract.

LITERATURE CITED

CLAUSEN, J., D. D. KEck, and W. M. HigseEy. 1945. Experimental studies on the
nature of species. II. Plant evolution through amphiploidy and autoploidy, with
examples from Madiinae. Carnegie Inst. Wash. Publ. 564:1-174.

CLAYTON, W. D. and S. A. RENvoIzE. 1986. Genera Graminum. Grasses of the
world. Her Majesty’s Stationery Office, London.

DARLINGTON, C. D. and A. P. WyLiz. 1956. Chromosome atlas of flowering plants.
George Allen and Unwin, London.

DAvIbsE, G. and R. W. POHL. 1974. Chromosome numbers, meiotic behavior, and
notes on tropical American grasses (Gramineae). Can. J. Bot. 52:317-328.
GOULD, F. W. 1965. Chromosome numbers in some Mexican grasses. Bol. Soc.

Bot. Mex. 29:49-62.

. 1966. Chromosome numbers in some Mexican grasses. Can. J. Bot. 44:

1683-1696.

and T. R. SODERSTROM. 1970. Chromosome numbers of some Mexican and
Canadian grasses. Can. J. Bot. 48:1633-1639.

HACKEL, E. 1890. The true grasses. Henry Holt and Company, New York.

Hatcu, S. L. 1980. Chromosome numbers of some grasses from the southwestern
United States and Mexico. Southw. Naturalist 25:278-280.

Myers, W.M. 1947. Cytology and genetics of forage grasses. Bot. Rev. (Crawfords-
ville) 13:319-421.

PETERSON, P. M. 1989. Lemma micromorphology in the annual Muhlenbergia
(Poaceae). Southw. Naturalist 34: in press.

, C. R. ANNABLE, and V. R. FRANCESCHI. 1988. Comparative leaf anatomy

of the annual Muhlenbergia (Poaceae). Nordic J. Bot. 8: in press.

and L. H. RIESEBERG. 1987. Flavonoids of the annual Muhlenbergia (Po-
aceae). Biochem. Syst. Ecol. 15:647-652.

POHL, R. W. and G. DAvipsE. 1971. Chromosome numbers of Costa Rican grasses.
Brittonia 23:293-324.

and W. W. MITCHELL. 1965. Cytogeography of the rhizomatous American
species of Muhlenbergia. Brittonia 17:107-112.

REEDER, J. R. 1967. Notes on Mexican grasses VI. Miscellaneous chromosome
numbers. Bull. Torrey Bot. Club 94:1-17.

. 1968. Notes on Mexican grasses VIII. Miscellaneous chromosome numbers.
Bull. Torrey Bot. Club 95:69-86.

SoLTis, D. E. 1980. Karyotypic relationships among species of Boykinia, Heuchera,
Mitella, Sullivantia, Tiarella, and Tolmiea (Saxifragaceae). Syst. Bot. 5:17-29.

TATEOKA, T. 1962. A cytological study of some Mexican grasses. Bull. Torrey Bot.
Club 89:77-82.

(Received 22 Dec 1987; revision accepted 10 Aug 1988.)

ASTRAGALUS NUTRIOSENSIS (FABACEAE):
A NEW SPECIES FROM EASTERN ARIZONA

MICHAEL J. SANDERSON
Department of Ecology and Evolutionary Biology,
University of Arizona, Tucson 85721

ABSTRACT

A new species of Astragalus from the White Mountains of Arizona is described
and its possible relationships discussed. Its unusual morphology combines features
found in two western North American sections of the genus. Morphological evidence
suggests it is most closely related to species in section Mollissimi, although it possesses
several character states that are unusual in the context of that section.

A distinctive new species of Astragalus has recently been discov-
ered among the mesas along Nutrioso Creek in the White Mountains
of eastern Arizona. Easily overlooked because of its dwarf habit and
cryptic flowers, it remains a difficult species to study in the field.
Two relatively large populations several kilometers apart were found
following the wet spring of 1987. The spring of 1988 was unusually
dry; population sizes were lower, and the plants were evidently suf-
fering from severe water stress, producing fewer leaves and flowers
than in the previous year. Preliminary indications are that the species
is a narrow endemic, but a complete survey of its distribution must
await a future, more favorable season.

Astragalus nutriosensis Sanderson, sp. nov. (Fig. 1)

Habitu A. mollissimus var. matthewsii (Wats.) Barneby, sed pe-
dunculis brevissimus; floribus perangustis longis, niveis, apicibus
subroseis; fructibus parvis (8-10 mm longis), bilocularibus perfecte,
in duo dimidia inaperta findentibus.

Dwarf, tufted, acaulescent, perennial herbs from a well-developed,
short but broad, knotty caudex on a simple taproot. Upper surface
of leaflets glabrescent-green; herbage otherwise cinereous and cov-
ered with spreading, basifixed hairs up to 2 mm long. Leaves 3-1 1
cmi long; petiole usually at least half that length; leaflets 4-8 mm
long, 1.5—4 mm wide, in 5-9 pairs, obovate or elliptic, acute or
obtuse at apex; stipules 3—9 mm long, deltoid or lanceolate, with
prominent green midrib and scarious or hyaline margins. Peduncles
(rachises) 0O—2(—5) cm long, almost always much shorter than the
leaves, and often appearing obsolete, not elongating in fruit; racemes
very short, up to 0.6 cm, (1—)2—5(—7)-flowered, the flowers strongly
ascending; bracts 2-5 mm long, greenish, narrowly lanceolate; ped-

MADRONO, Vol. 35, No. 4, pp. 325-329, 1988

326 MADRONO [Vol. 35

B
Cc
D
ess
—S
~ N 5
Sy iN
OSS Wad.
A

Fic. 1. Astragalus nutriosensis Sanderson (Sanderson 551). A. Habit. B. Flower.
C. Gynoecium. D. Wing petal. E. Longitudinal view of “dehiscent”? pod showing
complete septum with funicular scars. F. Pod, external view. G. Pod, transverse
section.

icels 2-4 mm long. Calyx 11-14 mm long, 2—3 mm wide at base of
teeth, narrowly cylindric, pilose with spreading hairs, subsymmetri-
cally attached at base, the receptacle not oblique. Petals white with
pink-purple veins and/or tips; banner 20-23 mm long, about 6 mm
wide, oblanceolate and emarginate at apex, little recurved (less than
30 degrees); wings 19-22 mm long, blade about 10 mm long, of
uniform, narrow width (2 mm), rounded at apex and not at all
incurved; keel significantly shorter than the wings, about 14-17 mm
long, blade of keel 5-6 mm long, obtuse and longer than wide.
Anthers about 0.5 mm long, filaments incurved only very distally.

1988] SANDERSON: NEW ASTRAGALUS FROM ARIZONA 321

Ovary unusually short at anthesis (about one-fifth the entire length
of the gynoecium), the septal intrusion only partially developed at
this time; ovules about 20. Pods small, 8-10 mm long, 6-8 mm
wide, almost beakless, broadly half-elliptic in profile, subterete in
cross-section, the ventral suture slightly depressed proximally, ses-
sile on receptacle (estipitate), on the very short peduncles, close to
the caudex, readily deciduous; valves papery, thin, brownish-gray
and covered with spreading hairs, septum complete throughout; de-
hiscence of fruit by fracture of pod into two closed, carpel-like halves,
each half retaining one side of the folded endocarp, the seeds prob-
ably eventually escaping through weathered septum walls.

Type: USA, Arizona, Apache Co., on hillside ne. of intersection
of AZ hwy. 666 and Apache Co. hwy. 116, about 9.5 km n. of
Nutrioso, 2270 m (7460 ft), 25 May 1987, Sanderson 551 (Holotype:
ARIZ; isotypes: NY, others to be distributed).

PARATYPE: USA, Arizona, Apache Co., nw. of intersection of AZ
hwy. 666 and Apache Co. hwy. 130, 2310 m (7600 ft), 6 km se. of
Springerville, and about 8 km from the type locality, 28 May 1988,
Sanderson 706 (ARIZ).

Ecology and distribution. Known only from two localities along
Nutrioso Creek in the White Mountains of eastern Arizona. Rare at
the type locality, but fairly common at the second locality. Found
on volcanic silty-clay soils on gently sloping hillsides; associated
with Bouteloua gracilis (H.B.K.) Lag. and Gutierrezia sarothrae
(Pursh) Britt. & Rusby in open grassland, sometimes among pinyon
and juniper on slopes. Sympatric with Oxytropis lambertii Greene,
Astragalus humistratus A. Gray, A. brandegei Porter, and A. mol-
lissimus var. matthewsii (S. Wats.) Barneby. Flowering in May, fruit-
ing by late May or June.

Phylogenetic relationships. Several morphological characters of A.
nutriosensis are shared by certain members of sections Argophylli
A. Gray and Mollissimi A. Gray. These include an acaulescent growth
habit, free stipules, basifixed pubescence, long, narrow flowers with
barely incurved petals, and a bilocular legume. The finer details of
the pod provide the best clues to the relatives of this species. Within
section Argophylli is a group of four species (A. waterfallii Barneby,
A. feensis M. E. Jones, A. holmgreniorum Barneby, and A. eurylobus
Barneby), all of which have bilocular pods technically similar to
those of A. nutriosensis. However, the shape, texture, and dehiscence
of the bilocular pod in these four species is fundamentally different.
Each is marked by a long, narrow, incurved pod, with trigonous or
cordate cross-section (and sometimes incomplete septum), fleshy or
ligneous valves, and dehiscence which is first apical, then through
ventral and dorsal sutures. The pod of A. nutriosensis 1s short, broad,
little incurved, terete in cross-section, completely bilocular, and pa-

328 MADRONO [Vol. 35

pery in texture. Its dehiscence is unusual, involving separation of
the pod into two closed halves, the seeds ultimately spilling through
the septum walls.

The shape and texture of the pod are more similar to species in
section Mollissimi A. Gray, particularly to Astragalus mollissimus
vars. thompsonae (S. Wats.) Barneby and matthewsii (S. Wats.) Bar-
neby, which are difficult to distinguish in the area of Nutrioso Creek.
The dehiscence is similar to that of A. anisus, a narrow endemic of
the Gunnison Valley, Colorado, of somewhat uncertain phylogenetic
position [placed by Barneby in section Argophylli (Barneby 1964:
718), but by Jones in section Mollissimi (Jones 1923:230)]. Mature
plants of A. mollissimus are typically much more robust than those
of A. nutriosensis, but seedlings of the former may easily be confused
with specimens of the latter if flowers or fruits are not present. The
copious pubescence of 4. nutriosensis, matched in A. mollissimus,
contributes to the overall impression of similarity. The pod of var.
matthewsii 1s completely bilocular, which, along with a tendency
towards a dwarf habit, suggests that it may be the closest extant
relative to A. nutriosensis.

Astragalus nutriosensis may be distinguished from A. mollissimus
var. matthewsii by several characters: a much smaller pod, barely
half the length of typical pods of species in the section; an almost
sessile inflorescence with few flowers, which is unique in the section
and rare among North American groups in general; and white flowers
with barely recurved banners. These differ from the purple flowers
and more reflexed banners characteristic of all other members of
the section, except perhaps the recently rediscovered Astragalus hart-
manil Rydb. from northern Chihuahua (Spellenberg pers. comm.;
see e.g., Hartman 678, at NY) which is a robust, strongly caulescent,
leafy plant otherwise quite dissimilar to A. nutriosensis.

Floral morphology. The flowers of A. nutriosensis are remarkably
straight and narrow, even in the context of the long, narrow flowers
typical of sections Argophylli and Mollissimi. In size and shape the
white flowers resemble those of the distantly related section Oro-
phaca (Torrey & A. Gray) Barneby and of A. wittmannii Barneby,
of section Humillimi (M. E. Jones) Barneby. The flowers in both
those cases are found on very short peduncles crowded together at
the bases of the leaves (Barneby 1979). Perhaps this unusual syn-
drome represents convergent evolution for increasing flower number
given the constraints of an inflorescence crowded among leaf-bases.

ACKNOWLEDGMENTS

I thank the American Society of Plant Taxonomists for providing funds to study
at the New York Botanical Garden. Rupert Barneby kindly pointed out the interesting
flowers of A. wittmannii and made useful comments on an early draft of this manu-
script. Renee Rusler provided indefatigable help while collecting, and Anne Gondor
drew the illustration.

1988] SANDERSON: NEW ASTRAGALUS FROM ARIZONA 329

LITERATURE CITED

BARNEBY, R. C. 1964. Atlas of North American Astragalus. Mem. New York Bot.
Gard. 13:1-1188.

. 1979. Dragma Hippomanicum IV: new taxa of Astragalus sect. Humillimi.
Brittonia 31:459-463.

Jones, M. E. 1923. Revision of North American species of Astragalus. Published
by the author.

(Received 27 Jan 1988; revision accepted | Jul 1988.)

NOTES

ADDITIONAL SUPPORT FOR THE RECENT-INVASIVE ADVENT OF MESQUITE (MIMOSA-
CEAE: Prosopis) IN THE SAN JOAQUIN VALLEY, CALIFORNIA.—In a recent article (Hol-
land, Madrono 34:324—333, 1987), I speculated that one or both species of mesquite
present in the San Joaquin Valley (Prosopis glandulosa and P. pubescens) were nat-
uralized sometime between 1870 and 1890. Additional support for the hypothesis of
human-mediated establishment of P. glandulosa has since been found in an article
by Mackie (Nemophila 8:30, 1920) from which we quote in part: ““Some years ago,
Mr. J. A. Jastro, a well-known cattleman, introduced the large-podded mesquite
(Prosopis glandulosa) into the head of the San Joaquin Valley in Kern County to
improve the cattle ranges. The bushes grew from seeds producing less than a dozen
individuals. When these specimens fruited the cattle at once began to feed on the
pods and in this manner the mesquite was spread over a large area lying between
Buena Vista lake reservoir and Button Willow.” Haas et al. (Texas Agric. Exp. Sta.
Monogr. 1:10—19, 1973) have shown that P. glandulosa can produce flowers within
three years of germination, although other information (Mooney et al., 7n Simpson,
ed., Mesquite: its biology in two desert scrub ecosystems, 1977) indicates that longer
periods may be normal. Given Linton’s observation of “‘an occasional patch of mes-
quite and sage”’ on the north shore of Buena Vista Lake in 1907 (Condor 10:196-
198, 1908), the introduction may have occurred as late as 1900.

Additionally, the statement in the first paragraph of my recent article (Holland,
op. cit.) concerning the number and distribution of species of Prosopis is in error.
According to Burkart and Simpson (Jn Simpson, op. cit.), the genus contains 44
species, 40 of which occur in the New World.— DAN C. HOLLAND, Dept. of Biology,
Univ. Southwestern Louisiana, Lafayette 70504-2451 and BARRETT ANDERSON, Dept.
of Botany, California Academy of Sciences, Golden Gate Park, San Francisco 94118.
(Received 29 Mar 1988; revision accepted 12 Sep 1988.)

MADRONO, Vol. 35, No. 4, p. 329, 1988

NEW COMBINATIONS IN ARCTOSTAPHYLOS
(ERICACEAE): ANNOTATED LIST OF
CHANGES IN STATUS

PHILIP V. WELLS
Department of Botany, University of Kansas, Lawrence 66045

ABSTRACT

A total of 53 names in Arctostaphylos are reviewed, 16 being synonyms. Of the
remaining 37, designated hybrids account for 13 names: A. x benitoensis Roof, A.
x bracteata T. J. Howell, A. x cinerea T. J. Howell, A. <x coloradensis Rollins, A.
x helleri Eastw., A. x jepsonii Eastw., A. x laxiflora Heller, A. x oblongifolia T. J.
Howell, A. x pacifica Roof, A. x parvifolia T. J. Howell, A. x strigosa T. J. Howell
(and also A. x campbellae Eastw. and A. x media Greene that had earlier achieved
this status). Reductions from species to subspecies include 3 names: A. sonomensis
Eastw., A. montaraensis Roof, and A. knightii Gankin & Hildreth; reductions from
species to form comprise 5 names: 4. acutifolia Eastw., A. adenotricha Love, Love
& Kapoor, A. candidissima Eastw., A. setosissima Eastw., and A. tracyi Eastw.; and
the remaining 16 names are downward shifts in infraspecific rank, mainly from variety
to form.

Having studied the genus 4rctostaphylos over a period of 30 years,
I perceive difficulties for the non-specialist, but none greater than
the numerous names existing at more than one, often vaguely de-
fined, rank, mostly proposed at the species level, many of them
synonyms or based on local forms or hybrid individuals. This paper
deals mainly with reassignments in rank for validly published names
that have escaped evaluation. A brief exposition of the logic guiding
these taxonomic dispositions is essential. The taxonomic category
variety has been used ambiguously in Arctostaphylos. It has been
applied indiscriminately, on the one hand, to major geographic taxa
with substantially allopatric distributions (that zoologists have long
recognized as subspecies) and, on the other hand, to minor forms
with locally sympatric or largely intrapopulational distributions
(known to zoologists as morphs). There is a strong traditional usage
of varietal rank in botany in lieu of subspecies, but the International
Code recognizes variety as a rank intermediate between subspecies
and form. Within Arctostaphylos, however, subspecies serves well
as the category for variants of the species with more or less discrete
geographic distributions, whereas form is the obvious choice for
intrapopulational morphs (many species are dimorphic; e.g., hairy
form/smooth form). Variety is then a superfluous category in the
genus.

Examples of geographically discrete subspecies are well delineated
in A. hookeri, A. manzanita, and A. viscida (Wells 1968) and more

MApDRONO, Vol. 35, No. 4, pp. 330-341, 1988

1988] WELLS: ARCTOSTAPHYLOS oo

complexly so in A. glandulosa and A. tomentosa (Wells 1987). In-
trapopulational forms, based on minute indument characters, have
received an inordinate amount of attention in the wide-ranging
species, A. uva-ursi, predominantly in the northern part of North
America and not in Eurasia. Comparable variation exists in the
center of diversity for the genus in California, but the plethora of
species and subspecies there inhibits the indulgence of naming all
of the individual variation within populations; a beginning has been
made in the relatively wide-ranging species, A. glandulosa (Wells
1987). The sympatric forms of A. uva-ursi include the following taxa
named at the varietal or even subspecific level: var. adenotricha
Fern. & Macbr., var. coactilis Fern. & Macbr., subsp. /ongipilosa
Packer & Denford, subsp. stipitata Packer & Denford, and the nom-
inate subsp. and var. uva-ursi in part of its North American range.
In part because the pubescence forms serve as markers for ploidy
level, many of them coexist in the same populations; adenotricha is
diploid, stipitata and uva-ursi tetraploid, and coactilis and longipi-
losa variable (Packer and Denford 1974). On the other hand, the
pubescence types intergrade and some phenotypic plasticity of in-
dument in response to ecological factors has been observed (Rosatti
1987). The medley of intrapopulational variation within North
American A. uva-ursi is most appropriately treated at the taxonomic
level of form.

Another problem in 4rctostaphylos is the occurrence of localized
hybrid swarms or individuals at some points of sympatry for certain
species. Unfortunately, some botanists with an eye for differences
have collected hybrid individuals that later became the types of new
taxa, invariably at the species level. A well known example is the
group of taxa named at Waldo, Oregon, by T. J. Howell (1901).
Waldo was a mining district of diverse lithology (serpentinite, con-
glomerate, sands, basalt-gabbro), much disturbed and open to in-
vasion by manzanitas, chiefly Arctostaphylos viscida ssp. pulchella
(on the serpentine) and 4. canescens (on non-serpentine soils). The
circumstantial evidence suggests the possibility of hybridization be-
tween these two species. Of the four Waldo taxa named as species
by Howell, A. bracteata and A. strigosa are quite similar to A. ca-
nescens but differ slightly in the direction of A. viscida; the other
two, A. cinerea and A. oblongifolia, are intermediate between A.
canescens and A. viscida. Obvious hybrid swarms still existed at the
Waldo site in the period 1960-1967, when I visited the area in search
of Howell’s taxa, but I was unable to discern any valid populations
corresponding to his descriptions or to his specimens at Eugene
(ORE). Fortunately, the populations of manzanitas at Waldo, as well
as others in an area of several hundred square miles in southwestern
Oregon, were analyzed biosystematically by Gottlieb (1968). Using
hybrid index and scattered diagrams, Gottlieb quantitatively defined

332 MADRONO [Vol. 35

the existence of hybrid swarms between 4. canescens and A. viscida
at Waldo and elsewhere in the surrounding region. He concluded
that Howell’s names were based on individual variants selected from
hybrid swarms (Gottlieb 1968). In order to avoid arbitrary assign-
ments to synonymy, as has been done with A. bracteata and A.
strigosa under A. canescens by Adams (1940), reduction to hybrid
status (e.g., 4. x bracteata T. J. Howell) should suffice to neutralize
the four superfluous names from the Waldo type locality. Prolifer-
ation of names for hybrid individuals (as done to excess in Quercus)
should be avoided in Arctostaphylos, however. Only published names
that already clutter the literature deserve this fate. Species of hybrid
origin with substantial, stable populations (often ecologically iso-
lated from the putative parental species) ought to be sustained as
valid species; arbitrary use of hybrid designations for such well de-
fined entities would wreak extensive havoc on the established tax-
onomy of the genus, inasmuch as the pattern of variation throughout
Arctostaphylos is reticulate in nature.

ANNOTATED LIST OF PROPOSED CHANGES IN STATUS FOR
ARCTOSTAPHYLOS TAXA

A. acutifolia Eastw. See A. patula forma acutifolia.

A. adenotricha (Fern. & Macbr.) Love, Love & Kapoor. See A. uva-
ursi forma adenotricha.

A. X benitoensis Roof (pro sp.), stat. nov. Basionym: A. benitoensis
Roof, Four Seasons 5(4):5-8, 1978. This taxon appears to be
A. pungens H.B.K.; introgressed with few traits of A. glauca
Lindl. If A. x benitoensis constituted a coherent entity, it would
be extremely close to A. parrvana Lemmon, and might be placed
in synonymy with that species.

A. bowermanae Roof, Four Seasons 5(4):15—18, 1978, from the north
side of Mt. Diablo, is certainly A. manzanita Parry and possibly
synonymous with subsp. manzanita.

A. X bracteata T. J. Howell (pro sp.), stat. nov. Basionym: A. brac-
teata T. J. Howell, Fl. N.W. Amer., 417, 1901. This is one of
several hybrid intergrades between A. canescens Eastw. and A.
viscida Parry (closer to the former), named as species by T. J.
Howell, as elucidated by Gottlieb (1968) at the type locality
near the site of Waldo, Oregon.

A. X campbellae Eastw. (pro sp.). Based on A. campbellae Eastw..,
Leafl. W. Bot. 1:75, 1933. Probably 4. tomentosa (Pursh) Lindl.
subsp. crustacea (Eastw.) Wells, slightly introgressed with few
traits of A. glauca Lindl. (Wells 1987).

A. candidissima Eastw. See A. canescens forma candidissima.

A. canescens Eastw. subsp. canescens forma candidissima (Eastw.)
Wells, comb. et stat. nov. Basionym: A. candidissima Eastw.,

1988] WELLS: ARCTOSTAPHYLOS 335

Leafl. W. Bot. 3:124, 1942. A variably white-downy extreme
form of A. canescens subsp. canescens.

A. canescens Eastw. subsp. sonomensis (Eastw.) Wells, comb. et stat.
nov. Basionym: A. sonomensis Eastw., Leafl. W. Bot. 1:78, 1933.
A consistently different glandular race of A. canescens with a
wide but segregated (allopatric) distribution relative to the nom-
inate subspecies (Knight 1985). Although subsp. sonomensis
occurs on volcanic and other rocks, it appears to be restricted
to serpentinite at the northern limits of its known range, as on
the summit of Horse Mountain, Humboldt Co. (unpublished
collection). Perhaps both glandulosity of pedicels and fruit and
serpentine tolerance derive from some genes of A. viscida subsp.
pulchella having introgressed into 4. canescens subsp. canescens
at some point in time and place.

A. chaloneorum Roof, Four Seasons 5(4):2—5, 1978, falls within the
range of variation of A. pungens H.B.K. as does A. benitoensis
Roof and A. pseudopungens Roof, all published in 1978. In this
interlude of critical splitting, Roof departed from his prior course
of lumping even distinct species such as A. manzanita Parry
under A. pungens in an extraordinarily broad conception of the
A. pungens “‘alliance” (Roof 1976). Later, he reduced A. chal-
oneorum as a subspecies under A. pungens (Roof 1979), a con-
sistency that he did not extend to A. benitoensis and A. pseu-
dopungens. Pending biosystematic elucidation of these
populations, synonymy under 4. pungens H.B.K. 1s appropriate
for A. chaloneorum and A. pseudopungens.

A. X cinerea T. J. Howell (pro sp.), stat. nov. Basionym: A. cinerea
T. J. Howell, Fl. N.W. Amer. 416, 1901. Another, more inter-
mediate, individual variant in the well-known Waldo hybrid
swarm, A. canescens X A. viscida (cf. Gottlieb 1968).

A. X coloradensis Rollins (pro sp.), stat. nov. Basionym: A. colo-
radensis Rollins, Rhodora 39:463, 1937. This name is based on
intermediate individuals in the hybrid swarm A. uva-ursi X A.
patula on the Uncompahgre Plateau of western Colorado. Re-
markably, the same cross is taking place in northwestern Mon-
tana (ridge north of Lake Mary Ronan, Lake Co.; Lesica and
Wells 1986) with some individuals matching 4. x coloradensis
(A. patula was previously unknown there, but A. uva-ursi is
sympatric, being widespread in the Rocky Mountains). Other
instances of this polytopic hybridization may come to light by
surveying the wide distribution of A. patula forma platyphylla.

A. columbiana Piper forma setosissima (Eastw.) Wells, comb. et stat.
nov. Basionym: A. setosissima Eastw., Leafl. W. Bot. 1:78, 1933.
An intensely setose form of the variably hairy species that occurs
locally with the nominate form, especially in southern Men-
docino Co.

334 MADRONO [Vol. 35

A. columbiana Piper forma tracyi (Eastw.) Wells, comb. et stat. nov.
Basionym: A. tracyi Eastw., Leafl. W. Bot. 1:79, 1933. A local
form lacking setose hairs, except on the bracts (as on the type);
all degrees of setosity can be found around Big Lagoon, Hum-
boldt Co., the type locality of A. tracyi. Eastwood named it on
the basis of the smooth-form specimens collected by Tracy. Both
forma tracyi and forma setosissima occur as intrapopulational
variants and should be treated as forms.

A. edmundsii J. T. Howell forma parvifolia (Roof) Wells, stat. nov.
Basionym: A. edmundsii var. parvifolia Roof, Leafl. W. Bot. 9:
191, 1961. Alocalized and intrapopulational, small-leaved form
of possible horticultural value.

A. glauca Lindl. forma eremicola (Jeps.) Wells, stat. nov. Basionym:
A. glauca var. eremicola Jeps., Madrono 1:78, 1922. This ep-
ithet is available for a decumbent form of A. glauca that layers;
the spreading form occurs on the desert slopes of the Transverse
and Peninsular Ranges. Layering from lower branches is a wide-
spread trait in Arctostaphylos, however, and there is no need to
formalize these vegetative forms by naming them unless, per-
haps, there is horticultural potential.

A. glauca Lindl. forma puberula (J. T. Howell) Wells, stat. nov.
Basionym: A. glauca var. puberula J. T. Howell, Leafl. W. Bot.
2:70, 1938. This local variant in indument deserves a rank no
higher than form.

A. X helleri Eastw. (pro sp.), stat. nov. Basionym: A. helleri Eastw.,
Leafl. W. Bot. 4:148, 1945. A putative hybrid, sympatric with
both parents: A. viscida Parry < A. myrtifolia Parry on the Ione
formation, a substratum to which the latter is narrowly endemic.
Eastwood named it from an individual specimen collected by
Heller (in Arctostaphylos, a treacherous undertaking). Surpris-
ingly, this cross has escaped biosystematic attention, whereas
the unnamed analogous cross, A. viscida xX A. nissenana Mer-
riam, has been well analyzed (Schmid et al. 1968).

A. imbricata Eastw. subsp. montaraensis (Roof) Wells, comb. et stat.
nov. Basionym: A. montaraensis Roof, Four Seasons 2(3):6—16,
1967. Aside from its tall, erect habit, this taxon is similar to
the creeping or mound-forming A. imbricata Eastw.; also, the
nascent bracts differ subtly in shape, subsp. montaraensis having
more acuminate tips. Although the differences are relatively
minor, the main populations of the two taxa are segregated on
two different mountains south of San Francisco, subsp. imbri-
cata on San Bruno and subsp. montaraensis on Montara Moun-
tain. At one spot on San Bruno, the two taxa coexist, indicating
that they are genetically distinct (also shown in common gar-
dens, as at Tilden); at San Bruno there are only a few individuals
of subsp. montaraensis growing with a large population of subsp.

1988] WELLS: ARCTOSTAPHYLOS 335

imbricata, whereas on Montara Mountain, closer to the Pacific
coast, there are large populations of erect subsp. montaraensis
but none of prostrate subsp. imbricata. The mainly allopatric
distribution argues for a rank of subspecies.

A. intricata T. J. Howell, Fl. N.W. Amer., 416, 1901, is a later
synonym for A. glandulosa Eastw. (1897); cf. Wells (1987).

A. X jepsonii Eastw. (pro sp.), stat. nov. Basionym: A. jepsonii
Eastw., Leafl. W. Bot. 1:110, 1934. The existence of local hybrid
zones between the elevationally segregated A. patula Greene
and A. viscida subsp. mariposa (Dudley) Wells has been well
documented (Epling 1947, Dobzhansky 1953). The earliest for-
mal recognition of the intergradation was described as A. mar-
iposa Dudley var. bivisa Jepson, Madrono 1:79, 1922. An ap-
propriate name for the hybrid A. patula x A. viscida subsp.
mariposa would be A. X jepsonii Eastw. because it honors the
prior author and was proposed at the species level.

A. knightii Gankin & Hildreth. See A. nevadensis subsp. knightii.

A. X laxiflora Heller (pro sp.), stat. nov. Basionym: A. laxiflora
Heller, Leafl. W. Bot. 4:148, 1945. This rare hybrid with very
showy panicles stems from the putative cross A. manzanita
Parry xX A. truei W. Knight, the two most plausible parents
near the type locality in Butte Co. on the lower slope of the
Sierra Nevada.

A. manzanita Parry subsp. bakeri (Eastw.) Wells. Synonym for A.
bakeri Eastw., which is now upheld as a distinct species.

A. X media Greene (pro sp.). Basionym: A. media Greene, Pittonia
2:171, 1891. The well known hybrid A. uva-ursi (L.) Spreng. x
A. columbiana Piper has been studied most recently by Krucke-
berg (1977). He has uncovered a parallel cross (A. nevadensis
A. Gray X A. columbiana) that produces a phenotype similar
to A. X media (as might be expected from the similarity of A.
nevadensis and A. uva-ursi). Fortunately, no formal name has
been proposed for this very similar hybrid.

A. montaraensis Roof. See A. imbricata subsp. montaraensis.

A. nevadensis A. Gray subsp. knightii (Gankin & Hildreth) Wells,
comb. et stat. nov. Basionym: A. knightii Gankin & Hildreth,
Four Seasons 3(3):23-24, 1970. My observations of this taxon
in the field indicate a very close similarity to A. nevadensis A.
Gray (and this is also apparent on the type specimen), except
for variably developed lignotubers that are most consistently
present at the type locality and nearby areas of Humboldt Co.
In Del Norte Co., in the vicinity of Gasquet, there are extensive
populations of A. nevadensis that mostly lack lignotubers; at
Humboldt Flat in the hills above Gasquet, I noted as long ago
as 1967 that some A. nevadensis had small basal burls, but
attributed this to hybridization with sympatric A. glandulosa

MADRONO [Vol. 35

Eastw. forma cushingiana (the latter as abundant as burl-free
A. nevadensis). Since lack of consistency as to the presence of
lignotubers is well known within other species of Arctostaphylos
(e.g., A. patula Greene), an infraspecific rank is indicated. In
view of the substantial allopatric populations in Humboldt Co.,
a rank of subspecies seems appropriate.

A. nitens Eastw., Leafl. W. Bot. 4:149, 1945, appears from the type

to be part of the A. glandulosa complex (Wells 1987), but along
with other collections from southwestern Oregon deserves pop-
ulational analysis in the field to determine consistency of the
described traits, presence or absence of burls (uncertain), etc.
Previous experience in this region indicates an extremely low
probability of taxonomic significance for this name.

A. X oblongifolia T. J. Howell (pro sp.), stat. nov. Basionym: A.

oblongifolia T. J. Howell, Fl. N.W. Amer. 416, 1901. Another
name based on the hybrid swarm at Waldo, Oregon: A. canes-
cens X A. viscida, and morphologically intermediate between
the two parental species.

A. obtusifolia Piper, Bull. Torrey Bot. Club 29:642, 1902, is syn-

onymous with A. patula forma platyphylla (A. Gray) Wells, q.v.

A. X pacifica Roof (pro sp.), stat. nov. Basionym: A. pacifica Roof,

Leafl. W. Bot. 9:217, 1962. Although it bears the stamp of an
A. uva-ursi lineage, this tiny population on San Bruno Mountain
has isofacial stomatal distribution and crown-sprouts from lig-
notubers. Past hybridization between A. uva-ursi (L.) Spreng.
and A. glandulosa Eastw. is the putative ancestry, both parental
species extant on San Bruno; non-sprouting forms of A. uva-
ursi formerly grew near the putative hybrid but were locally
eliminated by a relatively recent fire, while A. x pacifica re-
sprouted under the observation of Knight, Raiche, Roof and
others (see also the sprouting A. uva-ursi forma suborbiculata).

A. parryana Lemmon var. pinetorum (Rollins) Wiesl. & Schreib.

A.

See A. pinetorum.

x parvifolia T. J. Howell (pro sp.), stat. nov. Basionym: A.
parvifolia T. J. Howell, Fl. N.W. Amer. 416, 1901. Unlike the
group of taxa named from the hybrid swarms at Waldo, A.
parvifolia was based on collections from mountains west of
Andersons, Oregon, a considerable distance north and west of
Waldo. The type specimen has rather small green leaves, not
gray as in the Waldo taxa, which are derived from the cross A.
viscida (glaucous leaves) X A. canescens (gray, strigose-canes-
cent leaves). The simple, racemose inflorescence and small, green
leaves suggest that one parent was A. nevadensis A. Gray; the
white-hairy ovary and pedicels could be derived either from A.
glandulosa forma cushingiana (Eastw.) Wells or A. canescens.

1988] WELLS: ARCTOSTAPHYLOS 337

Hybrid swarms of A. nevadensis x A. glandulosa and individ-
uals resembling descriptions and type of A. parvifolia (with or
without a burl) are still being generated at Humboldt Flat, Del
Norte Co. On the other hand, Gottlieb (1968) decided that A.
parvifolia stems from the same A. viscida cross as the Waldo
hybrids.

A. patula Greene forma acutifolia (Eastw.) Wells, comb. et stat. nov.
Basionym: A. acutifolia Eastw., Leafl. W. Bot. 3:125, 1942. A
poorly known taxon, apparently collected only near the type
locality, Log Springs Ridge in southwestern Tehama Co. Pos-
sibly, Eastwood named it as a counterpoint in leaf shape to A.
obtusifolia Piper, a taxon that she accepted as a species (East-
wood 1934), even though the latter is indistinguishable mor-
phologically from A. patula forma platyphylla. Examination of
the type of A. acutifolia at CAS shows that it, too, is very close
to A. patula, but differs in having glandular-hairy pedicels and
stipitate-glandular fruit; the coalesced nutlets are seen also in
A. patula forma coalescens, as described next.

A. patula Greene forma coalescens (W. Knight) Wells, stat. nov.
Basionym: A. patula var. coalescens W. Knight, Four Seasons
7(1):20-21, 1984. The only distinguishing character is a ten-
dency toward partial coalescence of the normally separable nut-
lets; coherence of nutlets occurs sporadically in the North Coast
Range sector of the range of A. patula and may be expected
elsewhere. Inasmuch as it has been formally named, it is retained
as a form, but an occasional tendency toward coalescence of
nutlets is a commonly observed variation in the genus, and
ought not to be named; consistent fusion as a solid, indehiscent
stone, on the other hand, is an excellent character.

A. patula Greene forma platyphylla (A. Gray) Wells, stat. nov. Basi-
onym: A. pungens var. platyphylla A. Gray, Syn. Fl. N. Amer.
2:28, 1878; A. patula subsp. platyphylla (A. Gray) Wells, Ma-
drono 19:203, 1968. Recent field studies indicate that many
populations of A. patula commonly lack basal burls (lignotu-
bers) in the northern part of the Sierra Nevada and in many
parts of the North Coast Range, thus greatly reducing the al-
lopatry of burl-forming A. patula. Since the greater part of the
range of A. patula, from the northern Sierras and Cascades
eastward disjunctly to the Rockies of Montana, Utah and Col-
orado and southward in Nevada, Arizona and Baja California,
is occupied by populations that seem to /ack the burl (forma
platyphylla), attention should be focused on the actual extent
of burl-forming populations (forma patul/a) in the Sierra Nevada
and North Coast Range and whether there is segregation for the
burl trait there. Considering that presence or absence of the burl

338 MADRONO [Vol. 35

is the only distinguishing character and that this trait has not
been well documented in the putative burl-forming populations,
it seems best to recognize this difference as a form.

A. pinetorum Rollins, Rhodora 39:462, 1937, and A. parryana var.
pinetorum (Rollins) Wiesl. & Schreiber, Madrono 5:46, 1939,
are synonyms for A. patula forma platyphylla.

A. pseudopungens Roof, Four Seasons 5(4):9-11, 1978, is a mis-
nomer because, like 4. chaloneorum Roof, it is merely an out-
lying population of A. pungens H.B.K. It is apparent from the
late James Roof’s extensive writings (1978) that he miscon-
ceived A. pungens as being tetraploid on the basis of the somatic
count (2n=26) reported in Munz (1959), when, in fact, it is
mostly diploid, as is further confirmed by Niehaus’ counts on
A. pseudopungens (n=13), reported by Roof (1978). Neither A.
pseudopungens nor A. chaloneorum are sufficiently different from
A, pungens to require a formal name, though Roof is undoubt-
edly correct in his astute observation that both are introgressed
(limitedly) by certain traits of A. glauca Lindl. The name A. X
benitoensis Roof suffices to designate this introgression for-
mally.

A. pulchella T. J. Howell, Fl. N.W. Amer. 416, 1901, is synonymous
with A. viscida Parry subsp. pulchella (T. J. Howell) Wells,
Madrono 19:204, 1968.

A. serpentinicola Roof, Four Seasons 5(4):12-15, 1978, is synony-
mous with A. viscida subsp. pulchella (T. J. Howell) Wells. In
publishing this name, Roof (1978) neither justified the status
of full species, distinct from A. viscida Parry, nor in any way
distinguished 4. serpentinicola from the prior name, A. viscida
subsp. pulchella. In examining Howell’s type of A. pulchella,
Roof (1978:12) apparently did not observe that the pedicels are
glandular-hispidulous and the ovaries stipitate-glandular, as was
noted by me when I visited at ORE in 1967. Roof correctly
noted that there are two fragments, one a sterile branch of A.
viscida, the other a flowering branch that can be diagnosed, both
obtained in mountains west of Andersons, Josephine Co., Or-
egon, April 1886 (T. J. Howell’s handwriting). Finally, an af-
finity for serpentinite bedrock is also shown by the smooth-
fruited A. viscida subsp. viscida of the Sierra Nevada, but it is
not restricted to serpentine, being widespread on the primarily
granitic terrane. Thus, the correct name for the viscid-fruited,
serpentinicolous race of the North Coast Range and Siskiyou
Mountains, north into southwestern Oregon, is A. viscida Parry
subsp. pulchella (T. J. Howell) Wells.

A. setosissima Eastw. See A. columbiana forma setosissima.

A. sonomensis Eastw. See A. canescens subsp. sonomensis.

A. stanfordiana Parry forma decumbens Wells, stat. et nom. nov.

1988] WELLS: ARCTOSTAPHYLOS 309

Basionym and holotype: as in 4. stanfordiana var. repens Roof,
Four Seasons 4(2):16-17, 1972. Because of the horticultural
possibilities of this exceptionally beautiful species, this decum-
bent form deserves recognition. It should be noted that wherever
manzanitas branch to the base, the lower branches layer (take
root) if they contact the ground, so that there may be no end
to the naming of decumbent forms in the genus. In this instance,
the shrub is not repent or prostrate. Furthermore, the epithet
repens should be avoided in this genus, as it has been used
previously to designate another taxon, A. x repens (J. T. Howell)
Wells, based on A. cushingiana Eastw. forma repens J.T. Howell
(Leafl. W. Bot. 4:161, 1945).

A. X strigosa T. J. Howell (pro sp.), stat. nov. Basionym: A. strigosa
T. J. Howell, Fl. N.W. Amer. 417, 1901. Yet another name
proposed by Howell for a variant closer to A. canescens in the
hybrid swarm between the latter and A. viscida at Waldo, Or-
egon (cf. Gottlieb 1968).

A. tracyi Eastw. See A. columbiana forma tracyi.

A. uva-ursi (L.) Spreng. forma adenotricha (Fern. & Macbr.) Wells,
stat. nov. Basionym: A. uva-ursi var. adenotricha Fern. & Macbr.,
Rhodora 16:213, 1914. A largely intrapopulational, minutely
glandular form, widely sympatric in northern North America
and in the Rocky Mountains with nominate forma uva-ursi.
The latter, eglandular form extends farthest north in the Arctic
and has become circumboreal through Eurasia, where the species
is relatively uniform and tetraploid (n=26). Counts on forma
adenotricha have been consistently diploid (Packer and Denford
1974).

A. uva-ursi (L.) Spreng. forma coactilis (Fern. & Macbr.) Wells, stat.
nov. Basionym: 4. uva-ursi var. coactilis Fern. & Macbr., Rho-
dora 16:212, 1914. Another intrapopulational form commonly
present with forma adenotricha and forma uva-ursi in North
America, differing from the former in being eglandular and from
the latter in having the twigs and rachises minutely tomentulose;
ploidy level is variable, mostly diploid. Forma coactilis alone
extends south along the Pacific coast to California, where it
encounters a number of other species of the genus, possibly
giving rise to several local forms through hybridization. The
named forms are listed below.

A. uva-ursi (L.) Spreng. forma leobreweri (Roof) Wells, stat. nov.
Basionym: A. uva-ursi var. leobreweri Roof, Changing Seasons
1(2):26, 1980. This is one of several slightly differing forms (two
have been named) that occur as separate populations on San
Bruno Mountain, just south of San Francisco; all but /eobreweri
(glabrescent twigs) have indument similar to forma coactilis,
and are scarcely distinguished by leaf shape and habit. Forma

340 MADRONO [Vol. 35

leobreweri has incipient lignotubers; it propagates clonally by
suckering.

A. uva-ursi (L.) Spreng. forma longipilosa (Packer & Denford) Wells,
stat. nov. Basionym: A. uva-ursi subsp. /ongipilosa Packer &
Denford, Canad. J. Bot. 52:751, 1974. Yet another widely dis-
tributed intrapopulational form in North America, often sym-
patric with a number of other forms, especially forma coactilis,
forma adenotricha, and forma stipitata. Both diploid and tet-
raploid counts were reported by the authors.

A. uva-ursi (L.) Spreng. forma marinensis (Roof) Wells, stat. nov.
Basionym: A. uva-ursi var. marinensis Roof, Changing Seasons
1(2):19-21, 1980. A narrowly endemic, tetraploid form from
Pt. Reyes (n=26, unpublished meiotic count). Reportedly has
basal burl; may be sympatric with forma coactilis which is very
similar, but forma coactilis lacks lignotubers.

A, uva-ursi (L.) Spreng. subsp. monoensis Roof, Changing Seasons
1(3):7—9, 1980, from the Sierra Nevada, is not significantly dif-
ferent in its minutely glandular indument from forma adeno-
tricha and has a similar somatic number of 2n=26 (diploid level,
unpublished count by Wells on material from Tilden Botanical
Garden, Berkeley). Closely resembles Rocky Mountain material
of forma adenotricha (also diploid) and should be reduced to
synonymy with it.

A. uva-ursi (L.) Spreng. forma stipitata (Packer & Denford) Wells,
stat. nov. Basionym: A. uva-ursi subsp. stipitata Packer & Den-
ford, Canad. J. Bot. 52:750, 1974. A consistently tetraploid form
with indument solely of stipitate glands, but occurs as intra-
populational morph with forma /ongipilosa, forma coactilis, for-
ma adenotricha, etc., only in the far west. None of this intra-
populational variation deserves recognition at a rank higher
than form.

A. uva-ursi (L.) Spreng. forma suborbiculata (W. Knight) Wells, stat.
nov. Basionym: A. uva-ursi var. suborbiculata W. Knight, Four
Seasons 7(2):31—32, 1984. Another population from San Bruno
Mountain, San Francisco that is known to horticulturists in the
Bay Area by the sobriquet “‘miniature’’, distinguished mainly
by the rather round leaves and incipient lignotubers (docu-
mented crown-sprouter after recent fire; cf. A. X pacifica).

LITERATURE CITED

ADAMS, J. E. 1940. A systematic study of the genus Arctostaphylos Adans. J. Elisha
Mitchell Sci. Soc. 56:1-62.

DoOBZHANSKY, T. 1953. Natural hybrids of two species of Arctostaphylos in the
Yosemite region of California. Heredity 7:73-79.

Eastwoop, A. 1934. A revision of Arctostaphylos with key and descriptions. Leafl.
W. Bot. 1:105-127.

1988] WELLS: ARCTOSTAPHYLOS 341

EPLING, C. 1947. The genetic aspects of natural populations. Amer. Naturalist 81:
104-113.

GOTTLIEB, L. 1968. Hybridization between Arctostaphylos viscida and A. canescens
in Oregon. Brittonia 20:83-93.

HowELL, T. J. 1901. Flora of Northwest America. Published privately by author;
p. 415-417, re Arctostaphylos.

KNIGHT, W. 1985. A reappraisal of Arctostaphylos canescens Eastwood var. sono-
mensis (Eastw.) Adams. Four Seasons 7(3):42—-56.

KRUCKEBERG, A. R. 1977. Manzanita (Arctostaphylos) hybrids in the Pacific North-
west: effects of human and natural disturbance. Syst. Bot. 2:233-250.

LesicA, P. and P. V. WELLS. 1986. Arctostaphylos patula ssp. platyphylla (Gray)
Wells. Madrono 33:227-228.

Munz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley, p. 423.

PACKER, J. G. and K. E. DENFoRD. 1974. A contribution to the taxonomy of
Arctostaphylos uva-ursi. Canad. J. Bot. 52:743-753.

Roor, J. 1976. A fresh approach to the genus Arctostaphylos in California. Four
Seasons 5(2):2-24.

1978. Studies in Arctostaphylos (Ericaceae). Four Seasons 5(4):1-11.

1979. California’s Arctostaphylos pungens alliance. Changing Seasons 1(1):

20.

RosatTTI, T. J. 1987. Field and garden studies of Arctostaphylos uva-ursi (Ericaceae)
in North America. Syst. Bot. 12:61-77.

SCHMID, R., T. E. MALLORY, and J. M. TUCKER. 1968. Biosystematic evidence for
hybridization between Arctostaphylos nissenana and A. viscida. Brittonia 20:34-
43.

WELLS, P. V. 1968. New taxa, combinations and chromosome numbers in Arc-
tostaphylos (Ericaceae). Madrono 19:193-—210.

1987. The leafy-bracted, crown-sprouting manzanitas, an ancestral group

in Arctostaphylos. Four Seasons 7(4):4—27.

(Received 12 Feb 1988; revision accepted 10 Aug 1988.)

ANNOUNCEMENTS

NEw PUBLICATIONS

HumpnHreey, R. R., 90 years and 535 miles: Vegetation changes along
the Mexican border, University of New Mexico Press, Albuquerque,
New Mexico 87131, 1987, v, [i], 448 pp., illus., ISBN 0-8263-0945-3
(hardbound), price unknown. [A fascinating then-and-now photographic
comparison of the 535 changing miles of vegetation along the 205 mark-

ers designating the U.S.-Mexican boundary from El Paso, Texas, to San
Luis (by Yuma), Arizona; changes esp. evident in the Chihuahuan Des-
ert, the semi-desert grassland, and the Sonoran Desert, with “‘no life-
form or appreciable taxonomic changes along the largely ungrazed 60%
of the boundary” east of El Paso (p. 430).]

Mason, C. T., JR. and P. B. MAson, A handbook of Mexican roadside
flora, University of Arizona Press, 1615 E. Speedway, Tucson, Arizona
85719, 30 Oct 1987, [iv], 380 pp., illus., ISBN 0-8165-0997-2 (pap-
erbound), $19.95. [Some 200 taxa included.]

NOMENCLATURAL AND SYSTEMATIC REASSESSMENT OF
OPUNTIA ENGELMANNII AND O. LINDHEIMERI
(CACTACEAE)

BRUCE D. PARFITT and DONALD J. PINKAVA
Department of Botany, Arizona State University,
Tempe 85287-1601

ABSTRACT

For more than 100 years the conspicuous, large prickly-pear of the Southwest was
known as Engelmann Prickly-pear, Opuntia engelmannii Salm-Dyck. In 1965 Benson
and Walkington placed the name in synonymy under O. ficus-indica (L.) Miller and
proposed O. phaeacantha Engelm. var. discata (Griffiths) L. Benson and Walkington
for the Engelmann Prickly-pear. Newly discovered morphological characters, espe-
cially a unique glochid arrangement within areoles, and restudy of the publication
dates shows the correct name for the Engelmann Prickly-pear to be Opuntia engel-
mannii Salm-Dyck ex Engelm. Certain other taxa were found to share the glochid
arrangement, in addition to other characters, warranting revised synonymy and the
following new combinations: O. engelmannii var. lindheimeri; O. engelmannii var.
linguiformis; O. engelmannii var. flavispina; and O. engelmannii var. flexospina.

RESUMEN

Por mas de cien anos el nopal grande y conspicuo del suroeste de los Estados
Unidos ha sido conocido como “Engelmann Prickly-pear,” Opuntia engelmannii
Salm-Dyck. En el ano 1965 Benson y Walkington reconocieron este nombre como
sinonimo de O. ficus-indica (L.) Miller y propusieron para el ““Engelmann Prickly-
pear” el nombre O. phaeacantha Engelm. var. discata (Griffiths) L. Benson y Walking-
ton. Caracteres morfoldgicos recientemente descubiertos, especialmente una distri-
bucion unica de gloquidas adentro de las areolas, y un nuevo estudio de las fechas
de publicacion, muestra que el nombre correcto para el “‘Engelmann Prickly-pear”’
es O. engelmannii Salm-Dyck ex Engelm. Algunos otros taxa tienen en comun con
O. engelmannii la distribucion de gloquidas y otros caracteres obligando a una revision
de sinonimia y a las combinaciones nuevas siguientes: O. engelmannii var. lindhei-
meri; O. engelmannii var. linguiformis; O. engelmannii var. flavispina; and O. en-
gelmannii var. flexospina.

The Engelmann Prickly-pear was discovered by Wislizenus on his
expedition through northern Chihuahua in 1846. Duplicate speci-
mens were made available to Salm-Dyck and Engelmann. Engel-
mann’s notes packaged on the lectotype read in part, ‘Original spec-
imen...A shoot of this specimen was sent to Pr. Salm and described
by him.”’ Engelmann (1850) published his own description which
differs significantly from that of Salm-Dyck (1850), under the name
Opuntia engelmannii Salm-Dyck. Although Engelmann considered
Salm-Dyck to be the author of the name (as “Salm. Mss.’’), his
publication appeared in January 1850 (Stafleu and Cowan 1976),
four months before Salm-Dyck’s publication date (Salm-Dyck 1850,

MADRONO, Vol. 35, No. 4, pp. 342-349, 1988

1988] PARFITT AND PINKAVA: OPUNTIA 343

Stafleu and Cowan 1985). Because Engelmann’s paper has priority
and because he attributed the name to Salm-Dyck, the correct full
author citation of the name according to ICBN Art. 46.1, and Rec.
46E.1 (Voss et al. 1983) is Opuntia engelmannii Salm-Dyck ex En-
gelm.

The name, O. enge/mannii, remained in wide usage for the con-
spicuous, large prickly-pear of the Southwest until 1965 when Ben-
son and Walkington stated that the type represents a spiny individual
(““O. megacantha’’) of the cultivated, usually spineless species, O.
ficus-indica (L.) Miller. They placed O. engelmannii and O. mega-
cantha Salm-Dyck in synonymy under O. ficus-indica and applied
O. phaeacantha Engelm. var. discata (Griffiths) Benson and Walk-
ington to the wild material, which continued to bear the common
name, Engelmann Prickly-pear.

DISCUSSION

Upon examination of the morphological characters of O. phaea-
cantha var. discata, spiny and non-spiny forms of O. ficus-indica,
and the lectotype of O. engelmannii, we discovered several char-
acters apparently overlooked by Benson and Walkington. We found
no significant differentiating morphological characters between the
lectotype of O. engelmannii and O. phaeacantha var. discata; we
consider them conspecific.

However, the following characters distinguish O. engelmannii from
O. ficus-indica: areoles on pericarp 12—35 vs. 40-70; shrub habit vs.
tree habit; glochids (within each areole on the middle of a stem-
segment) conspicuous, 3—5 or more mm long, stout, of unequal
lengths and widely spaced, generally throughout the areole (Fig. 1),
vs. cryptic (even in spiny forms), less than 1.5 mm long, and in a
tiny crescent near the apical margin of the areole (sometimes a few
subapical, hidden in the wool) (e.g., McLeod 1255, OBI; Millspaugh
4523, NY; Stover 195, SD; Vanderwier s.n., OBI); and 2n=66 (2n=22
for var. cuija) vs. 2n=88 (Pinkava and McLeod 1971, Pinkava et
al. 1973, Pinkava and Parfitt 1982). They cannot be distinguished
on the basis of presence or absence of spines. Although spineless
individuals of both species might have glochids reduced and incon-
spicuous, usually by removing the areole’s tomentum the charac-
teristic glochid pattern may be seen.

We do not agree with Benson and Walkington’s interpretation that
the type of O. enge/mannii is from a cultivated plant. The label on
lectotype reads: “Shab. north of Chihuahua, common as high up as
El Paso. 5-6 feet high. August 1846. A. Wislizenus, leg.”’ A fruit
packet reads: “largest species, North of Chihuahua, 5-6’ high, also
cultivated. Dr. Wislizenus, August 1846.”

Britton and Rose (1919:148) wrote: ‘““Salm-Dyck, who first studied

344 MADRONO [Vol. 35

the species, doubtless had but a single specimen before him, and
this or a duplicate is now in the herbarium of the Missouri Botanical
Garden. This type specimen came from near Chihuahua City, from
which place Dr. Rose has collected identical material. Dr. Engel-
mann, who published Salm-Dyck’s name, described the plant as
erect and 5-6 feet high, giving its range from Chihuahua City to
Texas. These remarks of his were doubtless based on notes of Dr.
Wislizenus, who collected the type and must have included more
than one species; as Engelmann says it is both cultivated and wild,
the cultivated plants doubtless referring to some of the many forms
grown about towns and ranches.” The lectotype specimen (MO)
includes only a fruit and a half-stem-segment; there is no evidence
that two species are included.

In regard to O. ficus-indica, Britton and Rose (1919:177-178)
stated: ‘‘Dr. Griffiths has recently figured a reversion which appeared
on the common spineless form which points very definitely to O.
megacantha as the origin of this form.” Benson (1982:517-518)
wrote: ““These two plants [O. ficus-indica and O. megacantha] were
quite similar, and the evolutionary origin of those known as O. ficus-
indica from those called O. megacantha (as the wild type) was pos-
tulated by David Griffiths (Journal of Heredity 5:222. 1914).” In
fact, Griffiths (1914) did not mention O. megacantha. Benson (1982:
932) further stated: ‘““This name [O. enge/mannii] has been applied
erroneously to the large conspicuous prickly-pear occurring from the
deserts of California to those of Texas but the type is from a culti-
vated individual of the spiny ‘Opuntia megacantha’ type.” We in-
terpret Wislizenus’ note, “‘also cultivated,’’ as saying that although
he collected from a native (wild) plant of common distribution be-
tween northern Chihuahua and Texas, he also saw cultivated plants
that, in his opinion, were of the same species.

Specimens collected in 1976 northeast of Ciudad Chihuahua (e.g.,
Mexico, Chihuahua, Rte 10, 2.4 mi e. of Buenaventura, 4950 ft
elev., Pinkava et al. 13223, ASU 86565, 86566) very closely match
the lectotype of O. engelmannii. These plants were neither under
cultivation nor appear to have escaped; habit and young fruit were
that of the wild Engelmann Prickly-pear, not that of O. ficus-indica.
From photos, these plants are estimated to be from 4—5 feet tall and
more or less erect; in cultivation, one might easily expect the native
plants to grow to 5-6 feet. If the lectotype were from the cultivated
spiny form of O. ficus-indica as proposed by Benson (1965, 1982),
we would expect it to reach 10-12 or more feet in cultivation.

Therefore, we do not consider O. enge/mannii as conspecific with
O. ficus-indica. We are recognizing O. ficus-indica in the sense of
Britton and Rose (1919) but including forms that differ only in the
presence of spines.

Opuntia ficus-indica and O. megacantha have not been typified.
Opuntia megacantha, known only from an inadequate description,

1988] PARFITT AND PINKAVA: OPUNTIA 345

would require the selection of a neotype before this name could be
applied to O. ficus-indica, which has priority. Opuntia megacantha
predates O. engelmannii but until typified, O. megacantha plays no
role in our decisions.

Opuntia lindheimeri Engelm. shares with O. engelmannii the glo-
chid arrangement, fruit color, habit, ““hairy’’ seedlings, and lack of
red bases on yellow inner perianth segments (though some flowers
have the perianth completely red, orange, etc.). There are relatively
subtle differences between these taxa, warranting no more than vari-
etal status for O. lindheimeri. In variety lindheimeri the chalky-white
outer layer of the spines (as found in var. enge/mannii) is absent or
nearly so, allowing the translucent yellow core to be conspicuous,
and the fruit is pyriform or rather abruptly narrowed at the base
instead of barrel-shaped.

Opuntia engelmannii is distinguished from O. phaeacantha sensu
stricto by the following characters: fruit internally red-purple vs.
green; inner perianth segments completely yellow vs. yellow with
red bases; taller habit; spines of seedlings long and hair-like; and
glochids (within each areole on the middle of a stem-segment) stout-
er, of unequal lengths and widely spaced, generally throughout the
areole (Fig. 1), vs. compacted into a dense crescent-shaped tuft at
the apex of the areole and, at least in more mature areoles, a prom-
inent columnar tuft near the center of the areole (Fig. 1). Although
intermediate specimens occur, especially in some populations of
Arizona, Nevada, and California, we recognize O. phaeacantha as
distinct from O. engelmannii. We do not recognize varieties of the
former.

The discovery of these suites of distinguishing characters neces-
sitates recognition of O. engelmannii as a species distinct from O.
phaeacantha and from O. ficus-indica but not specifically distinct
from O. lindheimeri.

TAXONOMIC TREATMENT

OPUNTIA ENGELMANNII Salm-Dyck ex Engelm. in Engelm. & A. Gray,
Boston J. Nat. Hist. 6:208. Jan 1850. Type: Mexico, Chihuahua,
n. of Chihuahua, common as high up as El Paso. Aug 1846,
Wislizenus 223 (Lectotype: MO 2015202! designated by Benson
and Walkington [1965], photos ASU!, NY, POM).

Opuntia engelmannii Salm-Dyck, Cact. hort. dyck. 1849. Apr 1850.
Illegitimate, later homonym (ICBN Art. 64). Type: duplicate
material of type of O. engelmannii Salm-Dyck ex Engelm. None
preserved (see Stafleu and Cowan, 1985).

1. OPUNTIA ENGELMANNII var. ENGELMANNII. Autonym created with
the publication of O. engelmannii var. cyclodes Engelm. & Big-
elow, Proc. Amer. Acad. Arts 3:291. 1856.

346 MADRONO [Vol. 35

Fic. 1. Comparison of glochid arrangements of areoles midway between the base
and apex of stem segments in four Opuntia taxa. A. O. engelmannii var. engelmannii
(lectotype, Wislizenus 223, MO). B. O. ficus-indica (Stover 165, SD). C. O. engel-
mannii var. lindheimeri (topotype, Reeves 6333c sheet 2, ASU). D. O. phaeacantha
(Worthington 10587, ASU). Scale line = 2 mm; placed at apical end of areole.

Opuntia discata Griffiths, Annual Rep. Missouri Bot. Gard. 19:265
+ pl. 27 (upper fig.). 1908.—O. engelmannii Salm-Dyck var.
discata C. Z. Nelson, Galesburg Republican Register, 20 Jul
1915; Trans. Illinois State Acad. Sci. 12:124. 1919.—O. phaea-
cantha Engelm. var. discata (Griffiths) L. Benson & Walkington,
Ann. Missouri Bot. Gard. 52:265. 1965. Type: USA, AZ, foot-
hills of the Santa Rita Mts, Apr 1905, Griffiths 7790 (Holotype:
US 2572028A!, 2572029A!, 25720308A!, photos ASU!; iso-
type: POM 287144 [2 sheets]!, photos ASU!).

Opuntia dillei Griffiths, Annual Rep. Missouri Bot. Gard. 20:82-83
+ pls. 2 (fig. 10), 4 (ower fig.), 13 (fig. 7). 1909. Type: USA,
NM, San Andreas canyon of the Sacramento Mts, about 15
miles s. of Alamogordo, 3 Aug 1908, Griffiths 9460 (Holotype:
US 2576308A!, 2576309A!, photos ASU!; isotype: POM
288128).

The names O. engelmannii and O. lindheimeri were simulta-
neously published (Engelmann 1850). Engelmann (1856) first treated
the two as conspecific and placed O. lindheimeri into synonymy of
O. engelmannii.

2. Opuntia engelmannii var. lindheimeri (Engelm.) Parfitt & Pin-
kava, comb. nov.—O. lindheimeri Engelm. in Engelm. & A.

1988] PARFITT AND PINKAVA: OPUNTIA 347

Gray, Boston J. Nat. Hist. 6:207. Jan 1850.—O. lindheimeri
Engelm. var. /indheimeri. Autonym created by Coulter (1896)
with publication of O. lindheimeri var. dulcis (Engelm.) J. Coul-
ter, var. occidentalis (Engelm. & Bigelow) J. Coulter, var. cy-
clodes (Engelm.) J. Coulter, and var. /ittoralis (Engelm.) J. Coul-
ter. Type: USA, TX, New Braunfels, 1845, Lindheimer s.n.
(Lectotype: MO 2016376! designated by Benson [1982], photo
ASU!).

Opuntia lindheimeri Engelm. var. lehmannii L. Benson, Cact. Succ.
J. (Los Angeles) 41:125. 1969. Type: USA, TX, Kleberg Co.,
King Ranch, 10 miles s. of ranch headquarters at Kingsville,
19 Apr 1965, Lehman and Benson 16557 (Holotype: POM
317076! [4 sheets], photos ASU!).

Opuntia tricolor Griffiths, Annual Rep. Missouri Bot. Gard. 20:85-—
86 + pl. 4 (upper fig.). 1909.—Opuntia lindheimeri Engelm.
var. tricolor (Griffiths) L. Benson, Cact. Succ. J. (Los Angeles)
41:125. 1969. Type: USA, TX, “‘prepared October 2,1908, from
cultivated specimens collected March 29, 1907, near Laredo,”’
Griffiths 8651 (Holotype: US 2571220A!, photo ASU!; isotype:
POM 287271; clonotypes: US 2571219A!, 2571221A!, photos
ASU!).

Opuntia subarmata Griffiths, Ann. Rep. Missouri Bot. Gard. 20:94
+ pls. 2 (fig. 1), 11, 13 (fig. 4). 1909.—[O. engelmannii Salm-
Dyck var. subarmata Weniger, Cacti of the Southwest 180. 1970
(invalid name, ICBN Art. 33.2)].—O. lindheimeri Engelm. var.
subarmata (Griffiths) Elizondo & Wehbe, Cact. Suc. Mex. 32:
16-18. 1987. Type: USA, TX, near Devils River, 22 Jul 1908,
Griffiths 9422 (Holotype: US 2572063A!, photo ASU!; isotype:
POM 288607, 288608; clonotype: US 2572064A!, photo ASU!).

3. Opuntia engelmannii var. linguiformis (Griffiths) Parfitt & Pin-
kava, comb. nov.—0O. linguiformis Griffiths, Annual Rep. Mis-
souri Bot. Gard. 19:270. 1908.—Opuntia lindheimeri Engelm.
var. /inguiformis (Griffiths) L. Benson, Cact. Succ. J. (Los An-
geles) 41:125. 1969.—[Opuntia engelmannii Salm-Dyck var.
linguiformis (Griffiths) Weniger, Cacti of the Southwest 181.
1970 (invalid name, ICBN Art. 33.2)]. Type: USA, TX, near
San Antonio, Aug 1906, Griffiths 8377 (Holotype: US 2571222:
isotypes: ASU 140761!, POM 317780).

[Opuntia lindheimeri Engelm. var. brava E. Schulz & Runyon, Trans.
Texas Acad. Sci. 14:57. 1930 (invalid name, ICBN Art. 34. 1a)].
For discussion of the name “var. brava,”’ see Benson (1969).

Variety /inguiformis was considered by Schulz and Runyon (1930)
to be a sterile mutant form. It is readily distinguishable by the
lanceolate stem segments. In other characters it resembles var. /ind-
heimeri. No native populations are known.

348 MADRONO [Vol. 35

4. Opuntia engelmannii var. flavispina (L. Benson) Parfitt & Pin-
kava, comb. nov.—O. phaeacantha Engelm. var. flavispina L.
Benson, Cact. Succ. J. (Los Angeles) 46:79. 1969. Type: USA,
AZ, Pima Co., Organ Pipe Cactus National Monument, Ajo
Mts, Alamo Canyon, 2300 ft, 27 Apr 1939, Nichol s.n. (Ho-
lotype: POM 306987; isotypes: ARIZ 64930, 83680; Herb.
Organ Pipe Cactus Natl. Mon.!; Herb. U.S. Natl. Park Service,
Santa Fe).

5. OPUNTIA ENGELMANNII var. CUIJA Griffiths & Hare, New Mexico
Agric. Exp. Sta. Bull. 60:44. 1906.— Opuntia cuija (Griffiths &
Hare) Rose in Britton & Rose, Smithsonian Misc. Coll. 50:529.
1908.— Opuntia lindheimeri Engelm. var. cuija (Griffiths & Hare)
L. Benson, Cact. Succ. J. (Los Angeles) 41:125. 1969. Type:
Mexico, San Luis Potosi, Griffiths 7596 (=7636) (Holotype: US
2576155A!, 2576156A!, photos ASU!; isotype: POM 287125).

Pinkava and Parfitt (1982) reported this taxon as diploid (2n=22).
The remainder of O. engelmannii is hexaploid (2n=66). Although
this taxon may be better treated as a species, we have chosen to treat
it here pending resolution of the uncertainty regarding the applica-
tion of O. cantabrigiensis Lynch, which has priority at the species
level.

6. Opuntia engelmannii var. flexospina (Griffiths) Parfitt & Pinkava,
comb. nov.—O. flexospina Griffiths, Bull. Torrey Bot. Club 43:
87. 1916.—[O. engelmannii Salm-Dyck var. flexispina [sic]
(Grifiths) Weniger, Cacti of the Southwest 178. 1970 (invalid
name, ICBN Art. 33.2)].—O. strigil Engelm. var. flexospina
(Griffiths) L. Benson, Cact. Succ. J. (Los Angeles) 46:79. 1974.
Type: USA, TX, vicinity of Laredo, dry gravelly hills, Jun 1911,
Griffiths 10301 (Holotype: US 2571224A!, 2571225A!, photos
ASU!; isotypes: POM 299916, 290308).

ACKNOWLEDGMENTS

Reviews of the nomenclature by Edward G. Voss and Dan H. Nicolson are greatly
appreciated. Allan D. Zimmerman provided helpful comments and discussion, es-
pecially regarding the Texas taxa. Sonia and Leslie Landrum kindly translated the
abstract to Spanish. We thank the curators of ARIZ, ASU, MO, NY, OBI, POM,
SD, US, and the herbarium of Organ Pipe Cactus National Monument for making
specimens available for study.

LITERATURE CITED

BENSON, L. 1969. The cacti of the United States and Canada—new names and
nomenclatural combinations—I. Cact. Succ. J. (Los Angeles) 41:124-128.

. 1982. The cacti of the United States and Canada. Stanford Univ. Press,

Stanford, CA.

1988] PARFITT AND PINKAVA: OPUNTIA 349

and D. L. WALKINGTON. 1965. The southern California prickly pears—
invasion, adulteration, and trial-by-fire. Ann. Missouri Bot. Gard. 52:262-273.

BRITTON, N. L. and J. N. Rose. 1919. The Cactaceae. Vol. 1. Publ. Carnegie Inst.
Wash. 248.

COULTER, J. M. 1896. Preliminary revision of the North American species of Echi-
nocactus, Cereus, and Opuntia. Contr. U.S. Natl. Herb. 3:355—462.

ENGELMANN, G. 1850. Cactaceae. Jn G. Engelmann and A. Gray, Plantae Lind-
heimerianae. II. Boston J. Nat. Hist. 6:195-209.

1856. Synopsis of the Cactaceae of the Territory of the United States and
adjacent regions. Proc. Amer. Acad. Arts 3:259-346.

GRIFFITHS, D. 1914. “Reversion” in prickly pears. J. Heredity 5:222-225.

PINKAVA, D. J. and M. G. McLEop. 1971. Chromosome numbers in some cacti of
western North America. Brittonia 23:171-176.

; , L. A. MCGILL, and R. C. BRown. 1973. Chromosome numbers in

some cacti of western North America. II. Brittonia 25:2-9.

and B. D. PARFITT. 1982. Chromosome numbers in some cacti of western
North America. IV. Bull. Torrey Bot. Club 109:121-128.

SALM-Dyck, J. zU. 1850. Cacteae in Horto Dyckensi Cultae Anno 1849. Bonn.

SCHULZ, E. D. and R. RUNYON. 1930. Texas cacti. Trans. Texas Acad. Sci. 14:1-
181.

STAFLEU, F. and R. S. COWAN. 1976. Taxonomic literature, 2nd ed. Vol. 1: A-—G.
Regnum Veg. 94.

and 1985. Taxonomic literature, 2nd ed. Vol. 5: Sal-Ste. Regnum
Veo, fi:
Voss, E. G., et al. 1983. International code of botanical nomenclature. Regnum
Veg. 111.

(Received 13 Jun 1988; revision accepted 25 Aug 1988.)

ANNOUNCEMENT

NEw PUBLICATIONS

FERREN, W. R., D. G. CAPRALIS, and D. Hickson, 1987, University of
California, Santa Barbara, Campus Wetlands Management Plan. Part
I, Technical Report on the Botanical Resources of West and Storke
Campuses. A Report to the UCSB Wetlands Committee. Environ-
mental Research Team, The Herbarium, Department of Biological
Sciences, University of California, Santa Barbara. Environmental Re-
port No. 12. 198 pp. $10.00. (Including Devereux Slough and portions
of Goleta Slough.)

ApT, K., C. D’ANTONIO, J. Crisp, and J. GAUVAIN. 1988. Intertidal
Macrophytes of Santa Cruz Island, California. The Herbarium, De-
partment of Biological Sciences, University of California, Santa Bar-
bara. Publication No. 6. 87 pp. $8.00.

Orders should be sent to The Herbarium, Department of Biological
Sciences, University of California, Santa Barbara 93106. Checks should
be payable to The UCSB Foundation.

SALIX SCOULERIANA (SALICACEAE)
DISCOVERED IN MEXICO

GEORGE W. ARGUS
National Museum of Natural Sciences,
Ottawa, K1A OM8, Canada

ABSTRACT

The boreal species Salix scouleriana has been found to occur in Mexico. A study
of the type material of Salix pattersonii and S. wendtii revealed that both are S.
scouleriana. The original identification of S. wendtii as new was due, in part, to a
misunderstanding of the change that occurs in ament morphology when a plant flowers
twice in the same year.

RESUMEN

La especie boreal Salix scouleriana se ha encontrado en México. Un estudio de
Salix pattersonii y de S. wendtii ha descubierto que ambas especies son S. scouleriana.
La identificaciOn original de S. wendtii como especie nueva se debe, en parte, a un
mal entendimiento del cambio que ocurre en la morfologia del amento cuando una
planta produce sus flores dos veces durante el mismo ano.

The genus Salix is less well known in Mexico than it is in other
parts of North America. Since C. Schneider’s 1918 conspectus of
the genus in Mexico, West Indies, and Central and South America
only five papers on the genus in this region are known to me (Standley
1920, Johnston 1944, Espinosa 1979, Johnston 1981, Nee 1984).
In the southwestern United States there are about eight willows of
arctic-alpine (S. arctica Pall., S. brachycarpa Nutt., S. glauca L., S.
reticulata ssp. nivalis (Hook.) A. Love, D. Love & Kapoor) or boreal
GS. bebbiana Sarg., S. dr'ummondiana J. Barratt ex Hook., S. plani-
folia Pursh, S. scouleriana J. Barratt ex Hook.) affinities that occur
at relatively high elevations. None of these species have been re-
ported to occur in Mexico, although suitable habitats, at least for
some of the boreal species, are to be found there. It did not come
as a complete surprise, therefore, to find one of them present in
Mexico under a different name.

In the course of studying the holotypes of two Mexican Salix, S.
pattersonii M. C. Johnston and S. wendtii M. C. Johnston, I dis-
covered that both are the widespread boreal species S. scouleriana,
a species not previously known to occur in Mexico. Both of John-
ston’s new species, which were described in 1981, were based on
single collections: S. pattersonii on Riskind and Patterson 1809 (LL)
and S. wendtii on Wendt and Adamcewicz 518 (TEX). In addition

MADRONO, Vol. 35, No. 4, pp. 350-352, 1988

1988] ARGUS: SALIX SCOULERIANA IN MEXICO 351

to the holotypes two specimens tentatively identified as S. patter-
sonii, namely, Wendt 124a (TEX), and Riskind et al. 1720 (LL) and
two specimens listed as “Salix sp. nov.?”, Wendt and Lott P29
(CAN, TEX) and Mueller 3242 (LL), have also been identified here
as S. scouleriana.

The Mexican specimens have the following characteristics that
are typical of S. scouleriana (Argus 1973): (1) Large floral winter
buds; (2) leaves with two kinds of indumentum, (a) leaves sparsely
pubescent beneath with a mixture of white and ferruginous hairs,
and (b) leaves densely tomentose beneath with white hairs, but with
a few ferruginous hairs on the upper surface; (3) aments usually
sessile, or borne on a short spur shoot and usually flowering pre-
cociously (see below); (4) pistils sericeous with a mixture of ferru-
ginous and white hairs; (5) pistils borne on long stipes (1.2—2.4 mm);
and (6) black or dark brown floral bracts.

The holotype of S. wendtii is an apparent exception to character
(3) in that its aments are borne on long, leafy shoots and are flowering
in early August. Precocious species flower before the leaves appear
and have sessile aments, whereas coetaneous and serotinous species
flower with or after leaf emergence and have aments borne on prom-
inent leafy spur shoots. This distinction between precocious and
coetaneous and serotinous species is usually given high taxonomic
value in the identification of Sa/ix. The presence of an exceptionally
long spur shoot in this holotype is most likely what directed John-
ston’s attention away from S. scouleriana or any of the other pre-
cociously flowering species. A close examination, however, reveals
that the aments on this plant emerged from buds formed that year
and therefore represent a second flowering. In some precociously
flowering species aments that emerge from buds in the same year in
which they were formed are borne on long, leafy shoots rather than
being sessile. This condition was observed previously in S. planifolia
in northern Saskatchewan (Argus 1979). In that particular case, as
well, the plants were described as a new species, S. tyrrellii Raup,
and incorrectly aligned with S. glauca before being correctly rec-
ognized as S. planifolia. In addition to having aments borne on long,
leafy spur shoots, plants that flower twice in the same year also tend
to have more prominent stipules— possibly a reflection of their vig-
orous growth. During field work in Arizona in 1986 I found a spec-
imen of S. scouleriana exhibiting second flowering and very vigorous
growth (Argus and Argus 12383 CAN), so the condition displayed
by the type of S. wendtii was familiar to me.

The Mexican specimens of S. scouleriana tend to differ from
northern specimens in having more villous or shaggy branchlets.
The species usually has short, velutinous hairs on young branchlets,
but some variation in branchlet indumentum does occur in the
species. It is not surprising, in any case, that these isolated, disjunct

352 MADRONO [Vol. 35

populations in the Southwest diverge somewhat from the main species
populations.

Salix scouleriana occurs in western boreal North America (Little
1976, map 179) from Alaska eastward to Manitoba and southward
in the cordillera to southern California, Arizona, and New Mexico.
The Mexican specimens of S. scouleriana known so far are all from
Coahuila, two from Sierra Maderas del Carmen, Mpio. de Ocampo
and two from Sierra de la Madera, Canon de Agua.

The habitat of S. scouleriana in the southwestern United States,
where it usually occurs as scattered individuals, is in mixed conifer
forests on steep, relatively dry slopes at elevations of 2350 to
3000 m.

Salix scouleriana in Mexico is similar morphologically and eco-
logically to the populations of the species in adjacent New Mexico
and Arizona. Its Mexican occurrence is significantly disjunct from
known localities in northern Arizona and New Mexico, but not
unexpected. A thorough field study of Mexican Salix could be ex-
pected to extend the range of this boreal-cordilleran species even
further and perhaps even to find other species of northern affinities.

ACKNOWLEDGMENTS

I thank Kathleen Pryer for critically reading the manuscript. The Spanish abstract
was prepared by Raul Cano.

LITERATURE CITED

Arcus, G. W. 1973. The genus Sa/ix in Alaska and the Yukon. Natl. Mus. Nat.
Sci., Publ. in Botany 2. 279 pp.

. 1979. Areevaluation of the taxonomy of Salix tyrrellii, a sand dune endemic.
Systematic Botany 4:163-177.

ESPINOSA DE G. RUL, J. 1979. Salicaceae. Flora fanerogamica del Valle de México
1:95-99.

JOHNSTON, I. M. 1944. Plants of Coahuila, eastern Chihuahua, and adjoining Za-
catecas and Durango, IV. J. Arnold Arbor. 25:431-434.

JOHNSTON, M. C. 1981. The diandrous, hypostomatous willows (Salicaceae) of the
Chihuahuan Desert Region. Madrono 28:148-158.

LitTLeE, E. L., Jk. 1976. Atlas of United States trees. Vol. 3. Minor western hard-
woods. U.S. Dept. Agric., Forest Serv., Misc. Publ. 1314.

NEE, M. 1984. Salicaceae. Flora de Veracruz. Fasc. 34. Instituto Nacional de In-
vestigaciones sobre Recursos Bioticos. Xalapa, Ver. 24 pp.

STANDLEY, P. C. 1920. Salix. In Trees and shrubs of Mexico. Contr. U.S. Natl.
Herb. 23:160-163.

(Received 2 Dec 1987; revision accepted 22 Jun 1988.)

NOTEWORTHY COLLECTIONS

CALIFORNIA

MUHLENBERGIA APPRESSA C. O. Goodding (POACEAE).— Los Angeles Co., San Cle-
mente Island: 26 Apr 1912, Wooton s.n. (US); canyon below the Tomb, w. side of
island, 148 m, 12 Apr 1962, Raven 17324 (RSA); second canyon s. of Seal Cove, 33
m, 8 May 1962, Raven 17603 (RSA); road just n. of Guds, 270 m, 9 May 1962,
Raven 17651 (RSA); canyon just n. of Gray, 460 m, 9 May 1962, Raven 17708 (RSA);
middle portion of Norton Canyon, 295 m, 11 Apr 1973, Thorne 42857 (RSA, NY).
San Bernardino Co., Providence Mts.: rocky slopes, 1476 m, 17 May 1930, Hoffman
18 (US); canyon above Bonanza King Mine, 1509 m, 29 Oct 1977, Thorne, Tilforth,
and Prigge 50701 (RSA).

Previous knowledge. Known only from s.c. AZ and Baja California, Mexico where
it generally occurs above 1300 m. Closest known station to San Clemente Island is
280 km se. near Santa Catarina, Sierra de Juarez, Baja California (1962, Broder 708,
ARIZ, MEXU). Closest known station to the Providence Mts. is 350 km e. at Camp
Creek Divide, Maricopa Co., AZ (1970, Parker s.n., ARIZ).

Significance. New records for CA and the California Channel Islands.

This species is closely related to M. microsperma (DC.) Kunth and can be distin-
guished by its basally broadened and longer (4.0-6.2 mm) lemmas, longer glumes
(1.0-2.0 mm), and by its narrow, ascending, loosely flowered panicle with closely
appressed branches. The specimens from San Clemente Island tend to have slightly
more open panicles than the type (Harrison and Kearney 1493, US) but match in all
other characteristics.

MUHLENBERGIA FRAGILIS Swallen (POACEAE). —San Bernardino Co., Clark Mt. Range,
gravelly limestone soil in wash, below corral near Pacific Mine, T17N RS7E, 1570
m, 9 Oct 1977, Prigge 2214 (RSA).

Significance. First record for CA, a range extension of ca. 475 km n. and nw. from
previously known locations in Sierra Juarez, Baja California, Mexico, and Gila Co.,
AZ.

MUHLENBERGIA PAUCIFLORA Buckl. (POACEAE).—San Bernardino Co., New York
Mts., upper Caruthers Canyon, sandy loam soils with Pinus monophylla Torr. &
Frem., Quercus turbinella Greene, and Garrya flavescens Wats., T14N R16E S31
se.44, 1755 m, 11 Sep 1983, Peterson, Annable, and Barkworth 1622 (UNLV, WS).

Significance. First record for CA, a range extension of ca. 200 km w. from previously
known locations in Yavapai Co., AZ.— PAUL M. PETERSON and CAROL R. ANNABLE,
Dept. of Botany and Ownbey Herbarium, Washington State Univ., Pullman, 99164.

COLORADO

ASPLENIUM SEPTENTRIONALE (L.) Hoffm. (ASPLENIACEAE).— Moffat Co., Dinosaur
National Monument, Douglas Mountain, 1.6 air km ne. of Zenobia Peak, on red
sandstone outcrops and knolls of the Uinta Mountain Group, with Pinus ponderosa
and Cercocarpus ledifolius, T3N R102W S28, 2560 m, 28 Jun 1987, Neely 4455
(COLO, CS, Dinosaur N. M.).

Significance. This collection represents a range extension of ca. 52 km from the
nearest location in Uintah Co., UT. Known elsewhere in w. CO only from Box
Canyon, Ouray Co., 305 km to the se. (Weber, Colorado Flora: Western Slope, CO
Assoc. Univ. Press, 1987).

ASTRAGALUS HAMILTONII C. L. Porter (FABACEAE). — Moffat Co., Dinosaur National
Monument, hills west of Deerlodge Park, in sandy interstices of small barren outcrops

MADRONO, Vol. 35, No. 4, pp. 353-359, 1988

354 MADRONO [Vol. 35

of the Morrison Formation with Juniperus and Cercocarpus intricatus, T6N R99W
$28, 1756 m, 17 May 1987, O’Kane 2768 (COLO, Dinosaur N. M.).

Significance. First CO record of this candidate for Threatened or Endangered status.
This collection represents a range extension of ca. 62 km ne. of the nearest population
16 km north of Bonanza in Uintah Co., UT. The few other populations occur in nw.
Uintah Co. at 1580 to 1935 m on soil derived from the Duchesne River Formation
(Welsh & Chatterly, Great Basin Naturalist 45:173-236, 1985).

ENCELIOPSIS NUDICAULIS (Gray) A. Nels. (ASTERACEAE).— Moffat Co., Dinosaur
National Monument, Yampa Bench, between the Billiard Table and Mantle Ranch
Road, 2.4 km n. of Red Rock Ranch, Park City Formation, on heavy clay hills with
Atriplex confertifolia, Tetradymia spinosa, and Machaeranthera grindelioides, T6N
R103W S10, 1720 m, 30 May 1987, Neely 4131 (CS, COLO, Dinosaur N. M.);
Dinosaur National Monument, Mantle Ranch Road where it crosses Red Rock Can-
yon, Park City Formation, T6N R103W S11, 1691 m, 2 Jun 1987, O’Kane 3035
(COLO, CS, Dinosaur N. M.).

Significance. First CO records and representing a range extension of ca. 32 km
from near the Green River in Uintah Co., UT. The species is scattered across the
Great Basin, Mohave Desert, Uinta Basin and Colorado Plateau in UT, NV, AZ and
CA (Welsh, Jn Welsh et al., Great Basin Nat. Mem. 9:1-894, 1987) and is disjunct
along the Salmon and Lemhi rivers in ID (Hitchcock and Cronquist, Flora of the
Pacific Northwest, Univ. Wash. Press, 1973).

ZIGADENUS VAGINATUS (Rydb.) Macbr. (LILIACEAE).— Moffat Co.: Dinosaur Na-
tional Monument, Blind Canyon, s. tributary of Yampa River, ca. 0.8 km n. of Mantle
Ranch Road, in deep overhanging alcove with Aquilegia micrantha and Cirsium
ownbeyi, T6N R102W S823, 1762 m, 2 Jun 1987, O’Kane 3037 (COLO, CS, Dinosaur
N. M.).

Significance. These collections are the first reports of this species from Colorado.
It is infrequently found in Dinosaur National Monument where it grows in hanging
gardens of alcoves with Aquilegia micrantha and, occasionally, with the narrowly
endemic Cirsium ownbeyi. Populations in CO are disjunct by ca. 210 km from the
nearest location in Grand Co., UT. A few other populations are found in Kane and
San Pete Cos., UT (Welsh, Jn Welsh et al., Great Basin Nat. Mem. 9:1-894, 1987).
A collection from e. of Echo Park Campground in the Monument (N. Holmgren in
1961, UTC) also is referable to this species (L. Schultz pers. comm.).

The field work leading to this report was supported by the National Park Service
under a contract with the Colorado Department of Natural Resources.—STEVE L.
O’KANE, JR., Dept. Biology, Box 1137, Washington Univ., St. Louis, MO 63130;
ELIZABETH E. NEELY, The Nature Conservancy, 1244 Pine St., Boulder, CO 80302;
and DIETER H. WILKEN, Dept. Biology, Colorado State Univ., Ft. Collins, CO 80523.

IDAHO

CARDAMINE CONSTANCE! Detling (BRASSICACEAE). — Shoshone Co., hillsides, 1.6 km
w. of Kellogg, 15 Jun 1929, Geo. G. Hedgcock s.n. (WIS); on slope, 6.4 km se. of
Kellogg, 15 Jun 1929, Geo. G. Hedgcock s.n. (WIS).

Previous knowledge. Cardamine constancei was first applied to specimens collected
2 Jun 1935 from Three Devils Creek, a tributary of the Middle Fork of the Clearwater
River, Selway National Forest (Clearwater National Forest), Idaho County, Idaho.
Greatest numbers of this formerly threatened, rare, endemic species occur under
Thuja plicata Donn ex D. Don forests at lower elevations within the coast-like
refugium, lower Selway River and nearby portions of the Middle Fork of the Clear-
water River, that house other Idaho endemics and disjunct coastal vegetation. Smaller
populations exist outside this optimum environment along the lower St. Joe, South
Fork of the Coeur d’Alene (Pine Cr.), and the North and South Forks of the Clearwater

1988] NOTEWORTHY COLLECTIONS 3)5)5)

River in Clearwater, Idaho, Nez Perce, and Shoshone counties (Henderson et al.,
For., Wildlife, Range Exp. Sta. Bull. 21. Univ. Idaho, Moscow, 1977; and Crawford,
M.S. thesis, Univ. of Idaho, Moscow, 1980). A collection from along Jackass Cr., a
tributary of the w. fork of Pine Cr., and from the w. fork of Pine Cr., 9.7 and 6.4 km
s. of Pinehurst made 31 May 1950 by J. H. and C. B. Christ, 19295 and 19279
respectively, are the only previous records of this species from northern Shoshone
County.

Significance. The Hedgcock collections, located 10.3 and 15.3 km by air northeast
of the northernmost Pinehurst collection, document a second location for the rare
‘“*State Watch” species, Cardamine constancei, in northern Shoshone County, Idaho
and extend its northern limit 7.1 km. Additionally, Hedgcock’s collections, discovered
among the exsiccatae purchased by WIS in 1985 from LCU, predate the type collection
and may be the first gatherings of this rare Idaho endemic. Mining and road building
activities have modified the landscape in and around Kellogg since Hedgcock’s visit
and his collection sites may no longer exist. Tributaries of the South Fork of the
Coeur d’Alene River in the vicinity of Kellogg could house as yet undiscovered
populations of this rare Cardamine. —CLARK G. SCHAACK, Dept. of Botany, Univ.
of Wisconsin, Madison 53706 and DoOuGLAss M. HENDERSON, Dept. of Biological
Sciences, Univ. of Idaho, Moscow 83843.

MONTANA

ASTRAGALUS CICER L. (FABACEAE).— Lake Co., foothills of the Swan Range ca. 5
km se. of the town of Swan Lake, T25N R17W S30, abundant in disturbed soil along
a logging road, 1160 m, 1 Sep 1987, P. Lesica 4505 (MONTU, NY) (verified by R.
Barneby, NY).

Significance. First report of this Eurasian species for MT.

BARBAREA VULGARIS R. Br. (BRASSICACEAE). — Flathead Co., Glacier National Park,
horse pasture near the Quarter Circle Bridge on McDonald Creek ca. 2 km w. of Park
Headquarters, common in disturbed soil, 945 m, 3 Jun 1986, P. Lesica and A. DeBolt
3759 (MONTU); Lincoln Co., less than 1 km se. of Idaho border and 25 km nw. of
Troy, T34N R34W S32 nw.'4, many flowering plants in lush, moist meadow on the
bank of Curley Creek, 755 m, 3 Jun 1987, K. H. Lackschewitz 11232 (ID, MONTU,
NY); Missoula Co., vicinity of Ninemile Ranger Station, 10 km ne. of Alberton,
TI5N R22W S8, frequent in a cleared pasture, 965 m, 7 Jun 1966, J. Christensen
and D. Owen s.n. (RM); Kelly Island Fishing Access, 8 km w. of Missoula, T13N
R20W S22 s.'2, moist meadow along Clark Fork River, 975 m, 22 Jun 1987, K. H.
Lackschewitz 11256 (MONTU, NY) (Lackschewitz 11232 and 11256 verified by A.
Cronquist, NY).

Significance. First records of this Eurasian species for MT.

CAREX EBURNEA Boott (CYPERACEAE).— Dawson Co., head of North Fork of Burns
Creek ca. 40 km nw. of Glendive, T21N R55E S19, locally common in hardwood
forest on a moderate n.-facing slope, 900 m, 11 Jun 1987, P. Lesica 4290 (MONTU,
NY); Fergus Co., foothills of the Big Snowy Mtns. along Rock Creek ca. 24 km sw.
of Lewistown, T13N R17E S25 se.%4, locally common in moist spruce forest on
gravelly limestone soil, 1555 m, 17 Jun 1987, P. Lesica 4321 (MONTU); Flathead
Co., foothills of the Whitefish Range ca. 2 km ne. of Whitefish, T31N R21W S29
nw.'4, uncommon in moist spruce forest, 915 m, 21 Jul 1987, P. Lesica 4429 (MON-
TU, NY) (Lesica 4290 and 4429 verified by A. Cronquist, NY).

Significance. Although reported for MT by Kolstadt (Great Plains Flora Associa-
tion, Flora of the Great Plains, 1986), these are the first confirmed records. The
Flathead Co. collection is the first record for the Pacific Northwest and a range
extension of ca. 400 km s. from Alberta. The Dawson Co. collection is a range
extension of ca. 120 km nw. from Billings Co., ND.

356 MADRONO [Vol. 35

CAREX TENUIFLORA Wahl. (CYPERACEAE).— Flathead Co., Glacier National Park,
Numa Ridge ca. 3 km ne. of the foot of Bowman Lake ca. 10 km ne. of Polebridge,
common on hummocks of a Sphagnum bog, 1525 m, 24 Aug 1987, P. Lesica and
A. DeBolt 4106 (MONTU, NY) (verified by A. Cronquist, NY).

Significance. First report for MT and the Pacific Northwest, a range extension of
ca. 400 km s. from Alberta.

CYTISUS SCOPARIUS (L.) Link (FABACEAE). — Sanders Co., n. above Hwy 200 between
Hwy 56 and the Heron turnoff, T26N R33W S30, many plants for ca. 300 m in the
roadcut and on the partially disturbed slope along the highway, 705 m, 4 Jun 1987,
K. H. Lackschewitz 11245 (ID, MONTU, NY) (verified by A. Cronquist, NY and
D. Henderson, ID).

Significance. First report of this Eurasian species for MT.

GENTIANOPSIS SIMPLEX (A. Gray) Iltis [=Gentiana simplex A. Gray] (GENTIANA-
CEAE).— Missoula Co., Granite Creek, 4.8 km wsw. of Lolo Hot Springs, 0.16 km e.
of Lolo National Forest Rd. 9942, ca. 3 km s. of ject. with Rd. 4209, TLIN R24W
S15 ne.%4 of se.4, ca. 80-100 plants, growing in small clusters on hummocks in a
spring seep area along the creek, 1360 m, 19 Jul 1986, J. S. Shelly and G. V. King
1231 (WIS); same location ca. 100-150 plants, 20 Jul 1987, J. S. Shelly, K. H.
Lackschewitz, and J. H. Rumely 1377 (MONTU); same location, locally abundant,
on raised margins and interfluves of braided creek meanderings, substrate muddy,
appearing marly, 20 Jul 1987, J. H. Rumely, J. S. Shelly and K. H. Lackschewitz 20/
VII/87-04 (MONT).

Significance. First records for MT, a range extension of ca. 280 km from central
ID. Also known from CA, OR, and NV.

HELENIUM HOoPESII A. Gray [=DUGALDIA HOOPESII (A. Gray) Rydb.] (ASTER-
ACEAE).— Beaverhead Co., ca. 0.5 km e. of Monida, T14S R6W S35 sw.'4, in boggy
area below the road to Red Rock Lakes, 2040 m, 1 Aug 1986, K. H. Lackschewitz
11061 (MONTU, NY) (verified by A. Cronquist, NY).

Significance. Although previously reported for MT by Davis (Flora of Idaho, 1952),
no MT records were cited by Bierner (Brittonia 26:385-392, 1974). This is the first
confirmed report. This species becomes more common just south in adjacent Clark
Co., ID.

HUTCHINSIA PROCUMBENS (L.) Desv. (BRASSICACEAE). — Beaverhead Co., Armstead,
dry saline soil among grass, 1675 m., 20 Jun 1920, E. B. Payson and L. B. Payson
1729 (RM); Tendoy Mtns., n. side of upper Big Sheep Creek Canyon 20 km sw. of
Lima, T15S R10W S10, locally common beneath sagebrush on a steep, w.-facing,
limestone talus slope, 2090 m, 13 Jun 1986, P. Lesica 3834 (GH, MONTU) (Lesica
3834 verified by R. Rollins, GH).

Significance. First records for MT. Reed Rollins (personal communication) believes
that the correct name for this species is Hymenolobus procumbens (L.) Nutt. ex Torr.
and A. Gray.

LINARIA CANADENSIS (L.) Dum. var. TEXANA (Scheele) Penn. (SCROPHULARI-
ACEAE).— Dawson Co., Makoshika State Park just e. of Glendive, TISN R56E S32,
disturbed grassland, 635 m, 30 Jun 1982, K. Scow, D. Culwell and L. Larson s.n.
(MONTU); Carter Co., low ridge ca. 8 km sw. of Alzada, T9S R59E S31, common
in open, sandy soil of a pine-oak woodland, 1125 m, 15 Jun 1986, P. Lesica 4124
(MONTU, NY) (Lesica 4124 verified by A. Cronquist, NY).

Significance. First records for MT. The Carter Co. collection is a range extension
of ca. 32 km nw. from Crook Co., WY; the Dawson Co. record is a range extension
of ca. 280 km nw. from Grant Co., ND.

1988] NOTEWORTHY COLLECTIONS Sy

LYCHNIS FLOS-CUCULI L. (CARYOPHYLLACEAE). — Lincoln Co., less than | km se. of
Idaho border and 25 km nw. of Troy, T34N R34W S32 nw.'%4, 50-60 plants in a
moist meadow on the w. bank of Curley Creek, 755 m, 3 Jun 1987, K. H. Lackschewitz
11233 (ID, MONTU, NY) (verified by A. Cronquist, NY and D. Henderson, ID).

Significance. First report of this Eurasian species for MT.

MYOSOTIS DISCOLOR Pers. (BORAGINACEAE). — Sanders Co., Beaver Gulch Rd., 5 km
w. of Heron, T26N R34W S5 e.!4, in drying vernal pools, 680 m, 4 Jun 1987, K. H.
Lackschewitz 11239 (ID, MONTU, NY) (verified by A. Cronquist, NY and D. Hen-
derson, ID).

Significance. First report of this Eurasian species for MT.

POLYSTICHUM SCOPULINUM (D. C. Eat.) Maxon (POLYPODIACEAE).—Sanders Co.,
Cabinet Gorge, w. bank of the Clark Fork River 4.8 km nw. of Noxon, T26N R33W
S23 n.'2, one large plant in lichen-covered rocks of old riprap just above the water
line, 665 m, 24 Jun 1986, K. H. Lackschewitz 10915 (MONTU, NY) (verified by A.
Cronquist, NY).

Significance. Second report for MT. The occurrence of this species in a man-made,
low elevation habitat is unusual.

RIBES COGNATUM E. Greene (GROSSULARIACEAE). — Sanders Co., Cabinet Gorge, w.
bank of Clark Fork River 3 km nw. of Noxon, T26N R33W S23 n.'4, in a relatively
dry site at the foot of exposed cliffs, 675 m, 24 Jun 1986, K. H. Lackschewitz 10919
(MONTU, NY) (verified by A. Cronquist, NY); Lincoln Co., above Yaak River Falls
20 km n. of Troy, T33N R33W S5 se.%, in dry cliffs, 730 m, 26 Jun 1986, K. H.
Lackschewitz 10934 (MONTU, NY); along Kootenai River above Kootenai Falls 10
km e. of Troy, T31N R32W S13, one plant in cliffs, 620 m, 2 Jun 1987, K. H.
Lackschewitz 11220 (ID, MONTU) (verified by D. Henderson, ID).

Significance. First records for MT, an extension of 40 km east from Bonner
Co., ID.

SATUREJA VULGARIS (L.) Fritsch [=Clinopodium vulgare L.] (LAMIACEAE). — Flat-
head Co., Glacier National Park, along Going-to-the-Sun Road 10 km ne. of West
Glacier, locally common in gravelly soil of road shoulder, 960 m, 15 Jul 1987, P.
Lesica and R. Potter 4400 (MONTU, NY) (verified by A. Cronquist, NY).

Significance. First report of this Eurasian species for MT and the Pacific Northwest.

SCLERANTHUS ANNUUS L. (CARYOPHYLLACEAE). — Missoula Co., road to Montana
Power Station at the Rattlesnake Creek Dam ca. 2 km n. of Missoula, T13N R19W
S2 ne.'4, large colonies along the fence in a horse corral, 1080 m, 20 May 1986, K.
H. Lackschewitz 10835 (MONTU, NY); along the n. bank of the Clark Fork River
ca. 8 km nw. of Missoula, T13N R20W S1, common in sandy soil among river
cobbles, 945 m, 27 Jul 1986, P. Lesica and A. Bradley 3994 (MONTU, NY); Sanders
Co., Beaver Gulch Road 5 km w. of Heron, T26N R34W S5 e.'4, in drying vernal
pools, 680 m, 4 Jun 1987, K. H. Lackschewitz 11238 (ID, MONTU); Hutchins’ house
at the s. edge of Heron, T27N R34W S34, locally common in driveway gravel, 670
m, 15 Jul 1987, P. Lesica 4399 (MONTU) (Lackschewitz 10835 and Lesica and
Bradley 3994 verified by A. Cronquist, NY; Lackschewitz 11238 verified by D.
Henderson, ID).

Significance. First records of this Eurasian species for MT.

SPIRAEA X PYRAMIDATA E. Greene (ROSACEAE).— Lincoln Co., along Pete Creek 3
km w. of Yaak, T35N R32W S4, 915 m, 21 Jul 1965, Mooar 753 (MONTU); Missoula
Co., along East Fork Lolo Creek ca. 40 km sw. of Missoula, T11N R23W S28, marshy
and mossy area, 1295 m, 19 Jul 1970, Mooar 12743 (MONTU); Sanders Co., Bull

358 MADRONO [Vol. 35

River Forest Service Campground 7 km nw. of Noxon, T26N R33W S10 n.'42, under
forest canopy, 700 m, 24 Jun 1986, K. H. Lackschewitz 10909 (MONTU, NY)
(verified by A. Cronquist, NY).

Significance. First records for MT. At all sites, this species occurs with S. betulifolia
and S. douglasii.

STELLARIA GRAMINEA L. (CARYOPHYLLACEAE). —Sanders Co., Beaver Gulch Road
4 km w. of Heron, T26N R34W S4, common in grassy places along the road, 680
m, 4 Jun 1987, K. H. Lackschewitz 11243 (ID, MONTU, NY) (verified by A. Cron-
quist, NY and D. Henderson, ID).

Significance. First report of this Eurasian species for MT.

THALICTRUM ALPINUM L. (RANUNCULACEAE). — Beaverhead Co., w. edge of Monida,
T15S R6W S3, common on hummocks in a moist alkaline meadow, 2070 m, 27 Jun
1986, P. Lesica 3918 (MONTU, NY) (verified by A. Cronquist, NY).

Significance. Although reported for MT by Hitchcock and Cronquist (Flora of the
Pacific Northwest, 1973), this is the first confirmed record, a range extension of ca.
110 km w. from nw. WY.

VALERIANELLA LOCUSTA (L.) Betcke (VALERIANACEAE).— Lincoln Co., Slee Lake 3.2
km ne. of Troy, T31N R33W S6, formerly disturbed ground in a moist, acidic
meadow, 770 m, 2 Jun 1987, K. H. Lackschewitz 11226 (ID, MONTU, NY) (verified
by A. Cronquist, NY and D. Henderson, ID).

Significance. — First report of this Eurasian species for MT.

WALDSTEINIA IDAHOENSIS Piper (ROSACEAE). — Missoula Co., e. of Fish Creek Road
ca. 100 m past the crossing of Granite Creek ca. 40 km sw. of Missoula, T11N R23W
S7 nw.'4s, large colonies in open ponderosa pine forest, 1280 m, 9 Jun 1987, K. H.
Lackschewitz 11246 (ID, MONTU, NY) (verified by A. Cronquist, NY and D. Hen-
derson, ID).

Significance. First report for MT. Previously known only from central ID, a range
extension of 29 km e. from Idaho Co.—KLAuUs LACKSCHEWITZ and PETER LESICA,
Dept. Botany, University of Montana, Missoula 59812 and J. STEPHEN SHELLY, Mon-
tana Natural Heritage Program, State Library Bldg., 1515 E. 6th Ave., Helena 59620.
We are grateful to Ronald Hartman for providing records from the Rocky Mountain
Herbarium.

WASHINGTON

ERIOGONUM DOUGLASII Benth var. DOUGLASII (POLYGONACEAE). — Ferry Co., Kettle
Range, Colville National Forest, 2 km w. of Thirteen Mile Mt., T35N R33E S27,
1220 m, 25 May 1986, Peterson and Annable 4230 (WS); 0.5 km s. of Thirteen Mile
Mt., T35N R33E S33 s.'2, 1070 m, 26 May 1986, Peterson and Annable 4267 (WS).
At both sites, common to dominant on shallow, loamy soils derived from andesite
bedrock, associated with Poa secunda Presl and Selaginella wallacei Hieron.

Significance. First record for Ferry Co., a range extension of ca. 140 km n. from a
previously reported site near Badger Mt., Douglas Co.—PAUL M. PETERSON and
CAROL R. ANNABLE, Dept. of Botany and Ownbey Herbarium, Washington State
Univ., Pullman 99164.

FESTUCA CALIFORNICA Vasey (POACEAE).—Ferry Co., Kettle Range, Colville Na-
tional Forest, open grassy slopes with Festuca idahoensis Elmer, Calamagrostis ru-
bescens Buckl., and Pseudotsuga menziesii (Mirbel) Franco., e. slopes of Mt. Leona,
T38N R34E S25 n.2, 1800 m, 3 Jul 1985, Peterson and Annable 3629 (WS).

Significance. First record for Ferry Co., previously known only from w. of the

1988] NOTEWORTHY COLLECTIONS Spe]

Cascades in Skagit Co., a range extension of ca. 300 km e. of known locations on
Fidalgo and Hat Islands.

LISTERA BOREALIS Morong. (ORCHIDACEAE). — Ferry Co., Kettle Range, Colville Na-
tional Forest, needle duff below closed forest of Picea engelmannii Parry and Abies
lasiocarpa (Hook.) Nutt., 1 km n. of Jungle Hill along Kettle Crest Trail, T36N R34E
S1 sw.4, 1830 m, 6 Jul 1985, Peterson and Annable 3784 (WS).

Significance. First record for Ferry Co., a sensitive plant in WA previously known
only from Okanogan Co., a range extension of ca. 50 km e. of the previously reported
site near Mt. Bonaparte.

PHACELIA FRANKLINII (R.Br.) Gray (HYDROPHYLLACEAE).— Ferry Co., Kettle Range,
Colville National Forest, 1 km nw. of Lambert Mt., T37N R34E S2 n.%2, 1890 m, 3
Jul 1985, Peterson and Annable 3651, 3652 (WS); trail junction between Midnight
Mt. and Copper Butte, T37N R34E S11 se.4, 1860 m, 19 Jul 1986, Peterson and
Annable 4482 (WS); e. of Copper Butte, T37N R34E S13 e.'2, 1645 m, 19 Jul 1986,
Peterson and Annable 4502 (WS). Small populations were found on grassy slopes,
along trails, and road berms beneath open forests of Picea engelmannii Parry, Pseu-
dotsuga menziesii (Mirbel) Franco., and occasionally Pinus contorta Dougl.

Significance. First record for Ferry Co., a sensitive plant in WA previously known
only from Okanogan Co., a range extension of ca. 45 km e. of the previously known
reported site near Cayuse Mt.

ANNOUNCEMENT

WHITEBARK PINE SYMPOSIUM

A symposium entitled ““Whitebark Pine Ecosystems: Ecology and
Management of a High Mountain Resource” will be held 29-31 March
1989 at Montana State University. This symposium will explore the
ecology and management of whitebark pine forests and associated re-
sources of the high mountains in western North America—a subject of
rapidly emerging recognition and importance. The symposium will con-
clude with a field trip into whitebark pine forests near Yellowstone
National Park. The subject matter is intended for natural resource man-
agers, research scientists, educators, specialists in wildlife, hydrology,
soils, fire, recreation, range, silviculture, and others interested in high
mountain resources. The symposium is co-sponsored by the U.S. Forest
Service, U.S. National Park Service, Montana State University, and the
Society of American Foresters who have designated it as a Regional
Technical Conference.

For registration information or to be added to the mailing list, write
to: University Scheduling and Conference Center, Room 280 F, Strand
Union Building, Montana State University, Bozeman 59717, or call
(406) 994-3333.

360 MADRONO [Vol. 35

ANNOUNCEMENTS

NEw PUBLICATIONS

ALLEN, B. H., Ecological type classification for California: The Forest
Service approach, U.S.D.A. Pacific Southwest Forest and Range Ex-
periment Station, General Technical Report PSW-98, pp. [i], 1-8, Nov
1987, no ISSN, gratis (from Pacific Southwest Forest and Range Ex-
periment Station, P.O. Box 245, Berkeley, California 94701).

CONRAD, C. E., Common shrubs of chaparral and associated ecosystems
of southern California, U.S.D.A. Pacific Southwest Forest and Range
Experiment Station, General Technical Report PSW-99, pp. [i], 1-
86, July 1987, no ISSN, gratis (for address see entry for Allen). [De-
scriptions of and keys to 132 taxa; detailed glossary; appendix covers
all genera including chaparral shrub and subshrub species recognized
in P. A. Munz’s 4 flora of southern California (1974).]

CONSTANCE, L., Versatile Berkeley botanists: Plant taxonomy and uni-
versity governance, Regional Oral History Office, Bancroft Library,
University of California, Berkeley, California 94720, 1988, 362-page
interview, $67.00 (or available for examination at Bancroft Library,
UC Berkeley, or Dept. of Special Collections, UCLA). [An account
of the ‘“‘unending synthesis” of more than 50 years as a former UC
Berkeley graduate student, professor, chair of the department, director
of UC (the herbarium), dean of the College of Letters and Sciences,
and umbellologist.]

CULLMANN, W., E. GOTz, and G. GRONER, The encyclopedia of cacti,
trans. by K. M. Thomas, Alphabooks, Sherborne, Dorset DT9 3LN,
1986, 340 pp., illus. (most color), ISBN 0-906670-37-3 (hardbound),
price unknown. [Translation of Kakteen, 2. Aufl., Eugen Ulmer GmbH
& Co., 1984. A revision of a 1963 book by Cullmann by Groner (esp.
on cultivation and photos) and Gotz (systematics). Sections on: struc-
ture, mode of living and classification of cacti; cactus culture; prop-
agation of cacti; building up and accommodating a cactus collection;
special cultural problems; genera and species of cacti (with keys); idem
alphabetically arranged. Superb photos, with useful keys to taxa and
clear descriptions. ]

FULLER, T. C. and E. McCLINTocK, Poisonous plants of California,
University of California Press, 2120 Berkeley Way, Berkeley, Cali-
fornia 94720, 1986, [11], vii, 433 pp., 16 pls. (color), illus. (B&W),
ISBN 0-520-05568-3 (hardbound), $25.00, ISBN 0-520-05569-1
(paperbound), price unknown. [=California Natural History Guides,
no. 53. Contents: detailed descriptions of various taxa (294 pp.); plant
toxins and derivative drugs (63 pp.); appendices (plants causing der-
matitis, plants causing hay fever and asthma, plants accumulating
nitrates); biblio.; indices. An immensely useful book, obviously not
just for Californians—chock-full of fascinating information, e.g., tea
mistakenly made of Digitalis purpurea instead of Symphytum x
uplandicum leaves killed an elderly Washington couple in 1977. For
review see D. Cheatham, Fremontia 16(1):30-31.]

ee

1988] COMMENTARY 361

COMMENTARY
MESSAGE FROM THE PAST CBS PRESIDENT

It is an honor and a pleasure for me to address the members of the society in this
third report from the past President since the inception of the practice in 1986. It
was particularly gratifying to note the response to a comment in last year’s report in
which Frank Almeda expressed concern at the lack of graduate student response to
financial support offered by the society to assist with their research. Several inquiries
were received. It is nice to know that the membership reads these messages and
responds to them.

As seems to be the case each year, the Society, through its Executive Council, faced
a number of challenges in the last year. Not the least of these came with the realization
that the costs associated with producing and distributing Madrono were once again
taxing our resources. The Financial Officer estimated that costs associated with the
journal would exceed the total income of the Society for the year. Clearly we could
not continue on that basis. In an attempt to bring these costs under control and restore
the Society to a sound financial status, the Council voted to reduce the number of
free pages allotted to members to five (5) per volume. We realized that this would
make it difficult for some authors but hope that members will be able use grant funds
as well as funds from other sources to continue to publish in the journal. I should
point out that for papers having joint authorship, the five page allotment applies for
each author who is a member of the society.

One further step taken by the Executive Council this year was to reduce the max-
imum number of years of eligibility for student membership to five years. This is in
line with the student membership period of other organizations and will help reduce
the Society’s deficit in producing Madrojno.

By the time you read this the CBS Graduate Student Meetings will have concluded
at San Jose State University. These meetings are sponsored by the Society and rein-
force our commitment to the research of our graduate student members, recognize
their outstanding work, and encourage communication between participating insti-
tutions. It is a time-consuming undertaking on the part of the graduate student
representative and the Society owes a debt of gratitude to Valerie Haley of San Jose
State for her efforts in organizing this year’s meeting. These meetings take place at
eighteen-month intervals. I encourage members to consider having their institutions
host future meetings and, perhaps more important, I urge you to keep the meetings
in mind and encourage your graduate students to participate. Papers are presented
in catagories of Proposed Research, Research in Progress, and Completed Research.

At the November 1987 meeting of the Council, proposals from graduate students
for the Society’s annual Graduate Student Research Award were reviewed. After
consideration of the excellent proposals that had been submitted it was decided to
make the award to Samuel Hammer of San Francisco State University for his project
entitled A Taxonomic Survey of the Lichen Genus Cladonia in California. Mr. Ham-
mer is a Student of Dr. Harry Thiers. A check for $250 was forwarded to Mr. Hammer
to help support his investigation. I know that the membership of the Society joins
me in offering our congratulations to Mr. Hammer. The procedures for applying for
the Graduate Student Research Award are quite simple and I encourage all of the
Society’s graduate student members to consider submitting proposals to the Council.
Requests for information and proposals should be sent to the current President of
the Society. Proposals should reach the Society by 1 October each year. Now is the
time to be thinking about a proposal for next year.

One final item, our annual banquet in February had the largest turnout in recent
memory, thanks in no small measure to the speaker, Dr. Stephen J. Gould. It was a
gala evening and enjoyed by all those in attendance. Our thanks to Dr. Peggy Fiedler,
First Vice President of the Society for arranging an exciting program. By contrast the
interesting monthly programs that Peggy arranged were attended by relatively few

362 MADRONO [Vol. 35

members. This continues a trend of poor attendance that has been noted in recent
years. I hope that members in the Bay Area will check the schedule of speakers for
the coming year and plan to attend one or more of the regular meetings. It is hard
to justify continuing monthly meetings with the present attendance.

It has been an honor for me to serve as President of the California Botanical Society
for the past year and I thank you all for your support and advice. I must say, however,
that the office of President is not what makes our society work. With the exception
of the occasional “‘fire’’ the President has to put out, it is the volunteer members of
the Executive Council that do all of the day-to-day chores that make things run
smoothly. The names of those people are listed inside the front cover. I think that
the membership of the Society owes those individuals a debt of gratitude for their
hard work, and I know that I do. Thanks to all of you.k—DALE W. MCNEAL, De-
partment of Biological Sciences, University of the Pacific, Stockton, CA 95211.

ANNOUNCEMENT

NEw PUBLICATIONS

MICKEL, J. T., with contributions by R. MCVAUGH, S. KARELL, and H.
BALSLEV, Liebmann’s Mexican ferns: His itinerary, a translation of
his “‘Mexicos Bregner,” and a reprinting of the original work, Con-
tributions from the New York Botanical Garden, vol. 19, pp. [i-v], 1-
173-[175-350], 27 Oct 1987, ISSN 0736-0509, ISBN 0-89327-324-
4, $30.50 U.S., $31.75 foreign, postpaid (for address see entry for
Austin et al.). [On Frederik Michael Liebmann’s (1813-1856) rare
1849 work describing 308 spp., 95 new. Contents: introduction (Mick-
el); itinerary and gazetteer (McVaugh); the translation (Karell and
Balslev); index to the translation; the reprint on pp. 175-end.]

Mozinco, H., Shrubs of the Great Basin: A natural history, University
of Nevada Press, Reno, Nevada, 1987, xvii, [1i], 342 pp., 24 pls.
(color), illus. (B&W), ISBN 0-87417-111-3 (hardbound), ISBN
0-87417-112-1 (paperbound), prices unknown. [Lengthy, somewhat
rambling, occasionally botanically inaccurate (e.g., Ephedra ‘“‘flow-
ers”), but still interesting and quite useful descriptions of 66 spp. in
25 fam.]

WARREN, B., Wilderness walkers: Naturalists in early Texas, Hendrick-
Long Publishing Co., P.O. Box 25123, Dallas, Texas 75225, 1987,
112 pp., illus., ISBN 0-937460-26-5 (hardbound), $12.95. [Excellent
children’s books on seven ‘“‘walkers,’’ horseback riders, coachmates,
and just wanderers across Texas: Berlandier, Drummond, Lindhei-
mer, Wright, Audubon, Roemer, and Lincecum. ]

1988] COMMENTARY 363

EDITOR’S REPORT FOR VOLUME 35

This annual report provides an opportunity for the editor to communicate the
status of manuscripts received for publication in Madrofio and to comment on
the other aspects of the journal. Between | July 1987 and 30 June 1988, 52 manu-
scripts were received (29 articles, 6 notes, and 17 noteworthy collections contributions
totalling 57 individual taxa). Since 30 June 1988, 26 manuscripts have been received
(14, 7, 5). The current status of the 51 unpublished manuscripts is 8 in review (7, 1,
0), 20 in revision (16, 4, 0), 8 needing decision by the editor (6, 2, 0), and 15
accepted for publication (8, 3, 4). There are 4 unpublished book reviews. Volume 35
included 62 published manuscripts (31, 6, 16), 5 book reviews, and 2 commentaries
or letters. The period between submittal and publication averaged 14 months for
articles. Five manuscripts were rejected, and 3 were withdrawn by the author. At the
beginning of my editorship I inherited several manuscripts that had been returned
to authors more than one year previously; the authors of 11 of these did not respond
to my inquiries about the status of the manuscripts, and I consider these manuscripts
to have been withdrawn.

Several individuals have been of much help to me as I took on the responsibilities
of Editor. Wayne Ferren, the outgoing Editor presented me with a healthy journal
with manuscripts in various stages of processing. Wayne gave me the opportunity to
gradually assume the editorial reins during a transition period in the fall of 1987.
Wayne’s help was invaluable in getting my term as Editor off to a smooth start. He
has continued to be an excellent source of advice when I have felt the need for counsel
on journal matters. In addition he made arrangements to provide the excellent pho-
tograph of C. H. Muller to whom this volume is dedicated and arranged to have Dale
Smith write the dedication. Steven Timbrook has continued to serve the Society both
as a member of the Editorial Board and by preparing the annual Index and Table of
Contents.

My thanks also go to the authors whose papers I have edited for their patience and
tolerance as I have learned the job of Editor. I have, on occasion, taken longer than
authors would have liked to process the manuscripts. Sometimes the fault is mine;
sometimes it is out of my hands. I thank authors for their frank comments when I
have made mistakes.

With volume 35 Madrono began the publication of Spanish language manu-
scripts and abstracts. To date we have published only one article in Spanish. An
additional 10 articles in English that pertain to Latin American topics have Spanish
abstracts. Raul Cano, a colleague here at Cal Poly, has been of much assistance to
me and to several authors in the preparation of Spanish abstracts. On behalf of the
Editorial Board I encourage Latin American members of the California Botanical
Society to submit Spanish-language manuscripts to Madrono.

Articles and notes in Madrofo reflect the wide-ranging interests of the members
of the California Botanical Society. Topics discussed in volume 35 include cytology,
ecology, floristics, hybridization, nomenclature, paleobotany, phytogeography, and
systematics. The papers represent research in widely dispersed geographical areas
including Canada, various parts of the United States, Mexico, Nicaragua, Ecuador,
and Peru. Madrofo continues to serve as a source of additional information as
well through reviews and announcements that inform members of the Society of
publications, meetings, and other topics of interest. As Editor, I encourage potential
authors to continue to submit manuscripts that maintain the broad cross-section of
botanical topics.—D.J.K. 5 Dec 1988.

364 MADRONO [Vol. 35

REVIEWERS OF MANUSCRIPTS

As Editor, I thank all reviewers for their assistance with manuscripts. Special thanks
are extended to those who reviewed several manuscripts published in 1988. I am
very grateful to each reviewer for his or her generous contribution of time and effort.
The quality of papers published in Madrofo reflects this contribution. Reviewers for
volume 35 are:

George W. Argus Amy Jean Gilmartin Rhonda Riggins
Daniel I. Axelrod Lisa Graumlich Reed C. Rollins
Mary E. Barkworth James R. Griffin Jerzy Rzedowski
Rupert C. Barneby James W. Grimes John O. Sawyer
Mark Borchert William L. Halvorson Mark A. Schlessman
Mary C. Carroll Emily L. Hartman Theresa Scholars
Annetta Carter Ronald L. Hartman James Shevock
Kenton L. Chambers James Henrickson Paul C. Silva

Curtis Clark Duane Isley James P. Smith
Martin L. Cody David Keil John Stechman
Susan Conard Leslie R. Landrum John Strother
Lincoln Constance Malcolm G. McLeod Barry Tanowitz
William B. Critchfield Dale McNeal Dale Thornburgh
Arthur Cronquist Robert Meinke Steven Timbrook
Thomas Daniel James D. Morefield Thomas Van Devender
Carla D’Antonio Erik T. Nilsen Frank Vasek

Frank W. Davis Kevin Nixon Nancy Vivrette

D. Douglas Robert W. Patterson William A. Weber
Patrick E. Elvander Arthur M. Phillips, HI Dieter H. Wilken
Barbara Ertter Donald J. Pinkava

ANNOUNCEMENT

NEw PUBLICATION

Hewson, H. J., Plant indumentum: A handbook of terminology, Aus-
tralian Flora and Fauna Series, no. 9, pp. i-vil, 1-27, 1988, ISSN
0813-6726, ISBN 0-644-50456-0, Aust $4.95 (from Australian Gov-
ernment Publishing Service, Box 84, Canberra, A.C.T. 2601). [In-

tended esp. for writers of the Flora of Australia to standardize ter-
minology. Contents: intro; basic concepts; description formula; glossary
and key to terms; biblio.; index. Each term is clearly described and
figured, usually with diminutive and adjectival forms of the term,
derivation of the term, examples. “‘The use of plain English descriptive
terminology is encouraged, e.g., ‘bifid T-shaped trichome’ is preferred
to ‘Malpighiaceous’ (p. 1).]

INDEX TO VOLUME 35

Classified entries: major subjects, key words, and results; botanical names (new
names are in boldface); geographical areas; reviews, commentaries. Incidental ref-
erences to taxa (including most lists and tables) are not indexed separately. Species
appearing in Noteworthy Collections are indexed under name, family, and state or
country. Authors and titles are listed alphabetically in the Table of Contents to the

volume.

Acamptopappus, generic relationships and
taxonomy, 247.

Acanthaceae (see Carolowrightia arizo-
nica).

Aizoaceae (see Carpobrotus edulis).
Alpine vascular flora: Mount St. Helens,
WA, 309; Sawatch Range, CO, 202.
Andes, northern, three new species of

Galium from, 1.

Antennaria aromatica, noteworthy col-
lection from CO, 72.

Apiaceae: New combinations: Lomatium
graveolens, L. graveolens var. alpinum,
rae
New species: Lomatium shevockii, from

CA, 121.

Noteworthy collections: Eryngium al-
ismaefolium from NV, 166; Loma-
tium bicolor var. bicolor and Neo-
parrya lithophila from CO, 73.

Aquilegia triternata x A. chrysantha,
noteworthy collection from AZ, 278.

Arabis breweri var. austinae range, 162.

Aralia racemosa, noteworthy collection
from CO, 72.

Araliaceae (see Aralia racemosa).

Arctostaphylos: New combinations: A. x
benitoensis, A. xX bracteata, A. x
campbellae, A. canescens subsp. ca-
nescens forma candidissima, A. canes-
cens subsp. sonomensis, A. x cinerea,
A. columbiana forma setosissima, A.
columbiana forma tracyi, A. edmundsii
forma parvifolia, A. glauca forma ere-
micola, A. glauca forma puberula, A.
x helleri, A. imbricata subsp. monta-
raensis, A. < jepsonii, A. x laxiflora,
A. X media, A. nevadensis subsp.
knightii, A. < oblongifolia, A. < pa-
cifica, A. x parvifolia, A. patula forma
acutifolia, A. patula forma coalescens,
A. patula forma platyphylla, A. stan-
fordiana forma decumbens, A. stri-
gosa, A. uva-ursi forma adenotricha,
A. uva-ursi forma coactilis, A. uva-ursi
forma leobreweri, A. uva-ursi forma
longipilosa, A. uva-ursi forma mari-

nensis, A. uva-ursi forma stipitata, A.

uva-ursi forma suborbiculata, 330.

Arizona: New species: Astragalus nu-
triosensis, 325.

Noteworthy collections: Aquilegia tri-
ternata x A. chrysantha, 278; Salix
planifolia subsp. planifolia, 5; Sty-
locline sonorensis, 278.

Aspleniaceae: Noteworthy collections:
Asplenium septentrionale from CO,
353; Dryopteris filix-mas from CA, 164.

Aspidotis densa, noteworthy collection
from CA, 279.

Asplenium septentrionale, noteworthy
collection from CO, 353.

Asteraceae: Acamptopappus generic re-
lationships and taxonomy, 247;
Chaenactis alpina, typification, 161;
review of Erigeron eatonii and allied
taxa, 77.

New taxa: Encelia densifolia, from
Mexico, 10; Erigeron eatonii var. la-
vandulus, from ID and OR, 83.

Noteworthy collections: Antennaria
aromatica and Crepis capillaris from
CO, 72; Enceliopsis nudicaulis from
CO, 354; Helenium hoopesii from
MT, 356; Senecio pattersonensis and
Stylocline psilocarphoides from CA,
165; S. sonorensis from AZ and CA,
278,219,

Astragalus: New species: A. nutriosensis,
from AZ, 325.

Noteworthy collections: A. cicer from
MT, 355; A. humillimus from CO,
72; A. pachypus var. pachypus from
CA, 279; A. sericoleucus from CO,
i2e

Atriplex pleiantha, noteworthy collection
from CO, 73.

Barbarea vulgaris, noteworthy collection
from MT, 355.

Boraginaceae: Noteworthy collections:
Cryptantha micrantha from OR, 167;
C. weberi from CO, 73; Hackelia cu-

MADRONO, Vol. 35, No. 4, pp. 365-370, 1988

366

sickii from NV, 166; Myosotis discolor

from MT, 357.

Brassicaceae: Arabis breweri var. austin-

ae range, 162.

Noteworthy collections: Barbarea vul-
garis from MT, 355; Cardamine
constancei from ID, 354; Dithyrea
wizlizenii from CO, 73; Hutchinsia
procumbens from MT, 356; Malcol-
mia africana from CA, 164.

Cactaceae (see Opuntia).

California: Abundance of plants with
extrafloral nectaries in desert com-
munities, 238; analysis of pollen and
plant macrofossils from western Sierra
Nevada, 132; Arabis breweri var. aus-
tinae range, 162; Chrysolepis chryso-
phyllain Pseudotsuga-hardwood forest
of the Klamath Mountains of CA, 285;
crossability and relationships of Pinus
muricata, 39; endemic vascular plants
of northwestern CA and southwestern
OR, 54; invasion of chaparral by Car-
pobrotus edulis and Salix lasiolepis af-
ter fire, 196; late Wisconsin vegetation
in western Mojave Desert, 226; Pro-
sopis, recent-invasive in San Joaquin
Valley, 329; relation of fire history and
soil to composition of maritime chap-
arral, 169; species frequency in relation
to timber harvest methods and eleva-
tion in the pine type of northeastern
CA, 150.

New taxa: (also see Arctostaphylos);
Lomatium shevockii, 121.

Noteworthy collections: Aspidotis den-
sa, Astragalus pachypus var. pachy-
pus, 279; Carex parryana var. hallii,
164; Carlowrightia arizonica, 279;
Dryopteris filix-mas, 164; Holodis-
cus boursieri, 279; Malcolmia afri-
cana, Mentzelia reflexa, 164, Muh-
lenbergia appressa, M. fragilis, and
M. pauciflora, 353; Penstemon bar-
nebyi, Poa pattersonii, Potentilla
concinna var. divisa, Ribes veluti-
num var. gooddingii, Senecio patter-
sonensis, Stylocline psilocarphoides,
164; S. sonorensis, 279; Trifolium
dedeckerae, 164.

Canada: Morphological and ecological
variation across a hybrid zone between
Erythronium oreganum and E. revo-
lutum on Vancouver Island, British
Columbia, 32; Saxifraga tischii, new
species from Olympic Mountains of

MADRONO

[Vol. 35

WA and Vancouver Island, British Co-
lumbia, 126.

Cardamine constancei, noteworthy col-
lection from ID, 354.

Carex: Noteworthy collections: C. ebur-
nea from MT, 355; C. parryana var.
hallii, from CA, 164; C. tenuiflora from
MT, 356.

Carlowrightia arizonica, noteworthy col-
lection from CA, 279.

Carpobrotus edulis, invasion in maritime
chaparral after fire, 196.

Caryophyllaceae: Genecology of Ceras-
tium arvense and C. beeringianum in
northwest Washington, 266.
Noteworthy collections from MT:

Lychnis flos-cuculi, Scleranthus an-
nuus, and Stellaria graminea, 357.

Cerastium arvense and C. beeringianum,
genecology of, in northwest Washing-
ton, 266.

Chaenactis alpina, typification, 161.

Chaparral, invasion by Carpobrotus ed-
ulis and Salix lasiolepis after fire, 196.

Chaparral, maritime, relation to fire his-
tory and soil, Burton Mesa, Santa Bar-
bara County, CA, 169.

Chenopodiaceae (see Chenopodium sim-
plex).

Chenopodium simplex, synonymy, 162.

Chromosome numbers: 25 annual species
of Muhlenbergia, 309.

Chrysolepis chrysophylla in Pseudotsuga-
hardwood forest of the Klamath
Mountains of CA, 285.

Clitoria: New taxa: C. subg. Neurocar-
pum sect. Mexicana including new se-
ries Mexicana, Tucumania, and Amer-
icana, 29.

Colorado: Alpine vascular flora and
vegetation of Sawatch Range, 202.
Noteworthy collections: Antennaria

aromatica, Aralia racemosa, 712; As-
plenium septentrionale, 353; Astrag-
alus humillimus, A. hamiltonii, 353;
A. sericoleucus, 72; Atriplex pleian-
tha, Crepis capillaris, Cryptantha
weberi, Dithyrea wizlizenii, 73; En-
celiopsis nudicaulis, 354; Ipomopsis
congesta subsp. crebrifolia, Loma-
tium bicolor var. bicolor, Mentzelia
densa, 73; Neoparrya lithophila,
Rumex verticillatus, 74; Zigadenus
vaginatus, 354.

Commelina, notes and changes in no-
menclature for Valley of Mexico taxa,
16.

1988]

Commelinaceae (see Commelina).

Commentary: Message from Dale W.
McNeal, past president of the Califor-
nia Botanical Society, 364; Editor’s re-
port for Vol. 35, 365.

Compositae (see Asteraceae).

Crepis capillaris, noteworthy collection
from CO, 73.

Croton nubigenus, new species from Nic-
aragua, 117.

Cruciferae (see Brassicaceae).

Cryptantha: Noteworthy collections: C.
micrantha from OR, 167; C. weberi
from CO, 73.

Cyperaceae: Noteworthy collections:
Carex eburnea from MT, 355; C. par-
ryana var. hallii from CA, 164; C. ten-
uiflora from MT, 356.

Cytisus scoparius, noteworthy collection
from MT, 356.

Desert communities, abundance of plants
with extrafloral nectaries, 238.

Dithyrea wizlizenii, noteworthy collec-
tion from CO, 73.

Dryopteris filix-mas, noteworthy collec-
tion from CA, 164.

Ecuador: New species: Galium fosber-
gii, 3.

Elatinaceae (see Elatine californica).

Elatine californica, noteworthy collec-
tion from NV, 166.

Encelia densifolia, new species from
Mexico, 10.

Enceliopsis nudicaulis, noteworthy col-
lection from CO, 354.

Endemic vascular plants of northwestern
CA and southwestern OR, 54.

Ericaceae (see Arctostaphylos).

Erigeron: E. eatonii and allied taxa, re-
view of, 77; E. eatonii var. lavandulus,
83.

Eriogonum: Noteworthy collections: E.
brachyanthum from OR, 167; E. doug-
lasii var. douglasii from WA, 358.

Eryngium alismaefolium, noteworthy
collection from NV, 166.

Erythronium: Morphological and ecolog-
ical variation across a hybrid zone be-
tween E. oreganum and E. revolutum,
S2.

Euphorbiaceae (See Croton nubigenus).

Extrafloral nectaries, abundance in desert
communities, 238.

INDEX

367

Fabaceae: Comparison of North and
South American Lupinus group Micro-
carpi, 92.

New taxa: Astragalus nutriosensis from
AZ, 325; Clitoria subg. Neurocar-
pum sect. Mexicana including new
series Mexicana, Tucumania, and
Americana, 29.

Noteworthy collections: Astragalus ci-
cer from MT, 355; A. humillimus
from CO, 72; A. pachypus var.
pachypus from CA, 279; A. serico-
leucus from CO, 72; Cytisus sco-
parius from MT, 356; Lathyrus lae-
tivirens from NV, 166; Trifolium
dedeckerae from CA, 166.

Fagaceae: Chrysolepis chrysophylla in
Pseudotsuga-hardwood forest of the
Klamath Mountains of CA, 285; Quer-
cus arizonica and Q. oblongifolia, note-
worthy collections from Baja Califor-
nia Sur, Mexico, 280.

Festuca californica, noteworthy collec-
tion from WA, 358.

Fire history and soil, relation to com-
position of maritime chaparral, Burton
Mesa, Santa Barbara County, CA, 169.

Fire, invasion of chaparral by Carpo-
brotus edulis and Salix lasiolepis after,
196.

Floras: Alpine vascular plants of Sawatch
Range, CO, 202; endemic vascular
plants of northwestern CA and south-
western OR, 54; high elevation vas-
cular plants of Mount St. Helens, WA,
309.

Galium: New species from northern An-
des: G. antuneziae, |; G. cajamarcense,
G. fosbergii, 3.

Genecology of Cerastium arvense and C.
beeringianum in northwest Washing-
ton, 266.

Gentianaceae (see Gentianopsis simplex).

Gentianopsis simplex, noteworthy collec-
tion from MT, 356.

Gramineae (see Poaceae).

Grossulariaceae: Noteworthy collec-
tions: Ribes cognatum from MT, 357;
R. velutinum var. gooddingii from CA,
165.

Hackelia cusickii, noteworthy collection
from NV, 166.

Helenium hoopesii, noteworthy collec-
tion from MT, 356.

368

Hilaria annua, new species from Mexi-
COm/.

Holocene, early: Analysis of pollen and
plant macrofossils from western Sierra
Nevada, CA, 132.

Holodiscus boursieri, noteworthy collec-
tion from CA, 279.

Hutchinsia procumbens, noteworthy col-
lection from MT, 356.

Hydrophyllaceae (see Phacelia frank-
linii).

Idaho: Cardamine constancei, notewor-
thy collection, 354; Erigeron eatonii
var. lavandulus, new variety, 83; Lep-
todactylon pungens subsp. hazeliae,
new combination from Snake River
Canyon, 107.

International Botanical Congress, XIV,
report on, 159.

Ipomopsis congesta subsp. crebrifolia,
noteworthy collection from CO, 73.

Klamath Mountains, CA, Chrysolepis
chrysophylla in Pseudotsuga-hard-
wood forest, 285.

Labiatae (see Lamiaceae).

Lamiaceae: Satureja vulgaris, notewor-
thy collection from MT, 357; Scutel-
laria lutilabia, new species from Mex-
ico, 112,

Lathyrus laetivirens, noteworthy collec-
tion from NV, 166.

Leguminosae (see Fabaceae and Mimo-
saceae).

Leptodactylon pungens subsp. hazeliae,
new combination from Snake River
Canyon of ID and OR, 107.

Liliaceae: Morphological and ecological
variation across a hybrid zone between
Erythronium oreganum and E. revo-
lutum, 32; Zigadenus vaginatus, note-
worthy collection from CO, 354.

Linaria canadensis var. texana, note-
worthy collection from MT, 356.

Listera borealis, noteworthy collection
from WA, 359.

Loasaceae (see Mentzelia).

Lomatium: L. bicolor var. bicolor, note-
worthy collection from CO, 73; no-
menclature of L. kingii, L. megarrhi-
zum, L. nuttallii, 70.

New combinations: L. graveolens, L.
graveolens var. alpinum, 71.

MADRONO

[Vol. 35

New species: L. shevockii, from CA,
12h.

Lupinus: Comparison of North and South
American L. group Microcarpi, 92.
Lychnis flos-cuculi, noteworthy collec-

tion from MT, 357.

Malcolmia africana, noteworthy collec-
tion from CA, 164.

Mentzelia: Noteworthy collections: M.
densa from CO, 73; M. reflexa from
CA, 164.

Mexico: Notes and changes in nomen-
clature for Valley of Mexico taxa of
Commelina, 16; Salix scouleriana dis-
covered in, 350.

New species: Encelia densifolia, 10;
Hilaria annua, 7; Scutellaria lutila-
bia, 112.

Noteworthy collections: Quercus ari-
zonica and Q. oblongifolia, from Baja
California Sur, 280.

Mimosaceae (see Prosopis).

Mojave Desert, late Wisconsin vegeta-
tion, 226.

Montana: Noteworthy collections of As-
tragalus cicer, Barbarea vulgaris, Car-
ex eburnea, C. tenuiflora, Cytisus sco-
parius, Gentianopsis simplex, Helenium
hoopesii, Hutchinsia procumbens, Li-
naria canadensis var. texana, Lychnis
flos-cuculi, Myosotis discolor, Polysti-
chum scopulinum, Ribes cognatum,
Satureja vulgaris, Scleranthus annuus,
Spiraea x pyramidata, Stellaria gra-
minea, Thalictrum alpinum, Valeri-
anella locusta, and Waldsteinia ida-
hoensis, 355.

Mount St. Helens, WA, high elevation
flora, 309.

Muhlenbergia: Chromosome numbers in
25 annual species, 320; noteworthy
collections of M. appressa, M. fragilis,
and M. pauciflora from CA, 353.

Myosotis discolor, noteworthy collection
from MT, 357.

Neoparrya lithophila, noteworthy collec-
tion from CO, 74.

Nevada: Noteworthy collections of E/a-
tine californica, Eryngium alismaefo-
lium, Hackelia cusickii, and Lathyrus
laetivirens, 166.

Nicaragua: Croton nubigenus, new
species, 117.

1988]

Opuntia: Nomenclatural and systematic
reassessment of O. engelmanniiand O.
lindheimeri, 342.

New combinations: O. engelmannii var.
lindheimeri, 346; O. e. var. lingui-
formis, 347; O. e. var. flavispina, O.
e. var. flexospina, 348.

Orchidaceae (see Listera borealis).

Oregon: Endemic vascular plants of
northwestern CA and southwestern
OR, 54; Leptodactylon pungens subsp.
hazeliae, new combination from Snake
River Canyon, 107; noteworthy col-
lections of Cryptantha micrantha and
Eriogonum brachyanthum, 167.

Packrat midden plant macrofossils, 226.
Paleobotany: Analysis of pollen and plant
macrofossils from western Sierra Ne-
vada, CA, 132; Late Wisconsin vege-
tation in western Mojave Desert, CA,

226.

Penstemon barnebyi, noteworthy collec-
tion from CA, 164.

Peru: New species: Galium antuneziae, |;
G. cajamarcense, 3.

Phacelia franklinii, noteworthy collec-
tion from WA, 359.

Pinaceae (see Pinus muricata).

Pine type of northeastern CA, species fre-
quency in relation to timber harvest
methods and elevation, 150.

Pinus muricata, crossability and rela-
tionships of, 39.

Poa pattersonii, noteworthy collection
from CA, 16S.

Poaceae: Hilaria annua, new species from
Mexico, 7; Muhlenbergia, chromo-
some numbers in 25 annual species,
320.

Noteworthy collections: Festuca cali-
fornica from WA, 358; Muhlenber-
gia appressa, M. fragilis, and M.
pauciflora from CA, 353; Poa pat-
tersonii from CA, 165.

Polemoniaceae: [pomopsis congesta
subsp. crebrifolia, noteworthy collec-
tion from CO, 73.

New combination, Leptodactylon pun-
gens subsp. hazeliae, from Snake
River Canyon of ID and OR, 107.

Pollen and plant macrofossils from west-
ern Sierra Nevada, CA, analysis of, 132.

Polygonaceae: Noteworthy collections:
Eriogonum brachyanthum from OR,

INDEX

369

167; E. douglasii var. douglasii from
WA, 358; Rumex verticillatus from CO,
74.

Polypodiaceae (see Polystichum scopu-
linum).

Polystichum scopulinum, noteworthy
collection from MT, 357.

Potentilla concinna var. divisa, notewor-
thy collection from CA, 165.

Prosopis, recent-invasive in San Joaquin
Valley, CA, 329.

Quercus arizonica and Q. oblongifolia,
noteworthy collections from Baja Cal-
ifornia Sur, Mexico, 280.

Ranunculaceae: Noteworthy collections:
Aquilegia triternata x A. chrysantha
from AZ, 278; Thalictrum alpinum
from MT, 358.

Reviews: Robert Richard Brooks, Ser-
pentine and Its Vegetation: A Multi-
disciplinary Approach, 74, Thomas S.
Elias, Conservation and Management
of Rare and Endangered Plants, 280;
D. J. Mabberley, The Plant-Book: A
Portable Dictionary of the Higher
Plants, 167; Barbara G. Phillips et al.,
Annotated Checklist of Vascular Plants
of Grand Canyon National Park 1987,
168; Jerzy Rzedowski and Miguel
Equihua, Atlas Cultural de México.
Flora, 75.

Ribes: Noteworthy collections: R. cog-
natum from MT, 357; R. velutinum var.
gooddingii from CA, 165.

Rosaceae: Noteworthy collections: Hol-
odiscus boursieri from CA, 279; Poten-
tilla concinna var. divisa from CA, 165;
Spiraea X pyramidata and Waldstein-
ia idahoensis from MT, 357.

Rubiaceae (see Galium).

Rumex verticillatus, noteworthy collec-
tion from CO, 74.

Salicaceae (see Salix).

Salix: S. lasiolepis, invasion in maritime
chaparral after fire, 196; S. planifolia
subsp. planifolia, noteworthy collec-
tion from AZ, 5; S. scouleriana dis-
covered in Mexico, 350.

San Joaquin Valley, CA, Prosopis, re-
cent-invasive in, 329.

Satureja vulgaris, noteworthy collection
from MT, 357.

370

Sawatch Range, CO, vegetation and al-
pine vascular flora, 202.

Saxifraga tischii, new species from
Olympic Mountains of WA and Van-
couver Island, British Columbia, 126.

Saxifragaceae (see Saxifraga tischii).

Scleranthus annuus, noteworthy collec-
tion from MT, 357.

Scutellaria lutilabia, new species from
Mexico, 112.

Scrophulariaceae: Noteworthy collec-
tions: Linaria canadensis var. texana
from MT, 356; Penstemon barnebyi
from CA, 164.

Senecio pattersonensis, noteworthy col-
lection from CA, 165.

Sierra Nevada range: Analysis of pollen
and plant macrofossils from Dinkey
and Exchequer meadows, 132; Loma-
tium shevockii, new species from CA,
121.

Sinopteridaceae (see Aspidotis densa).

Spiraea x pyramidata, noteworthy col-
lection from MT, 357.

Stellaria graminea, noteworthy collec-
tion from MT, 358.

Stylocline: Noteworthy collections: S.
psilocarphoides from CA, 165; S. son-
orensis from AZ and CA, 278.

Thalictrum alpinum, noteworthy collec-
tion from MT, 358.

Timber harvest methods and elevation
in the pine type of northeastern CA,
species frequency in relation to, 150.

Trifolium dedeckerae, noteworthy collec-
tion from CA, 166.

MADRONO

[Vol. 35

Valerianaceae (see Valerianella locusta).

Valerianella locusta, noteworthy collec-
tion from MT, 358.

Vancouver Island: Morphological and
ecological variation across a hybrid
zone between Erythronium oreganum
and E. revolutum, 32; Saxifraga tisch-
ii, new species from Olympic Moun-
tains of WA and Vancouver Island,
British Columbia, 126.

Vegetation of Sawatch Range, CO, 202.

Waldsteinia idahoensis, noteworthy col-
lection from MT, 358.

Washington: Genecology of Cerastium
arvense and C. beeringianum, 266; high
elevation flora of Mount St. Helens,
309; Saxifraga tischii, new species from
Olympic Mountains and Vancouver
Island, British Columbia, 126.
Noteworthy collections: Eriogonum

douglasii var. douglasii, Festuca cal-
ifornica, Listera borealis, and Pha-
celia franklinii, 358.

White Mountains of CA and NV: Note-
worthy collections of Carex parryana
var. hallii, Dryopteris filix-mas, Mal-
colmia africana, Mentzelia reflexa,
Penstemon barnebyi, Poa pattersonii,
Potentilla concinna var. divisa, Ribes
velutinum var. gooddingii, Senecio pat-
tersonensis, Stylocline psilocarphoides,
and Trifolium dedeckerae, 164.

Zigadenus vaginatus, noteworthy collec-
tion from CO, 354.

DATES OF PUBLICATION OF MADRONO, VOLUME 35

Number 1, pages

1-76, published 12 April 1988

Number 2, pages 77-168, published 9 June 1988
Number 3, pages 169-284, published 2 November 1988
Number 4, pages 285-370, published 2 March 1989

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