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MADRONO

A WEST AMERICAN JOURNAL OF BOTANY

VOLUME XXI 1971-1972

BOARD OF EDITORS

Class of:

1971—Marion OwnsBEy, Washington State University, Pullman Joun F. Davipson, University of Nebraska, Lincoln

1972—IrA L. Wiccins, Stanford University, Stanford, California REED C. Roriiins, Harvard University, Cambridge, Massachusetts

1973—MIcHAEL G. BArzsour, University of California, Davis Roy L. Taytor, University of British Columbia, Vancouver

1974—KenTON L. CHAMBERS, Oregon State University, Corvallis Wi11aAM A. WEBER, University of Colorado, Boulder

1975—ArTURO GOMEz-PompPa, Universidad Nacional Autonoma de México Duncan M. Porter, Missouri Botanical Garden, St. Louis

1976—Dennis ANDERSON, Humboldt State College, Arcata, California KINGSLEY R. STERN, Chico State College, Chico, California

1977—Date M. SmirtH, University of California, Santa Barbara WiL1LiAM Louis CuLBErsON, Duke University, Durham, North Carolina

Editor—RoBERT ORNDUFF Managing Editor—Joun L. STROTHER Department of Botany, University of California, Berkeley 94720

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

Printed by Gillick Printing, Inc., Berkeley, California 94710

The California Botanical Society dedicates volume 21 of Madrono to WALLACE Roy Ernst, former member of the Board of Editors and at heart a western botanist. His careful studies of the floral morphology of the Papaveraceae reflect original and fresh interpretations of the rela- tionships among the genera of this difficult and complex family. In the Loasaceae and the genus Lamourouxia (Scrophulariaceae) his correla- tions of variation patterns in floral morphology with different pollination mechanisms represent a novel and pioneering effort in explaining the adaptive significance of these morphological patterns. The importance of his work in these and other groups lies not only in the solutions he offered to perplexing systematic and biological problems, but also in pro- viding models of a broad and eclectic methodology and of a rigor of inter- pretation. His wide store of knowledge extended to a diversity of wild and cultivated plants and was available freely to colleagues and students. His pervasive love of plants was reflected in many ways, including a strong interest in gardening. Whenever the opportunity was available he exhibited a remarkable horticultural expertise and a highly refined aes- thetic sense. Though a native Californian and one who loved the “‘wide open spaces” of California, Wally took a great pride in Washington, D.C. He delighted in showing the city and its numerous sights to visiting bot- anists from all parts of the world. His favorite tour was one of the city at night, including unusual perspectives of illuminated monuments and fountains—a memorable experience indeed. In many respects, he func- tioned as an unofficial host for the Department of Botany at the Smith- sonian Institution and frequently entertained visiting botanists in his home. Wally’s sense of perfection and his highly critical intellect were evident in every task he undertook, botanical or otherwise, and these qualities were particularly evident in the careful and meticulous critiques of manuscripts that he reviewed for others. Those who knew Wally Ernst miss him.

TABLE OF CONTENTS

ALAM, M. T. (See McArthur et al.) = 2 BAKER, HERBERT G., A fen on the northern California coast BAKER, WILLIAM H., Noteworthy records of western plants .

Barsour, M. G., Additions and corrections to the flora of Bodega Head, California

BARLow, Bryan (See Helton et al.) . : BEATTIE, ANDREW J., Itinerant pollinators in a forest

BEAUCHAMP, R. MiTcHEL, New locality for Lavatera venosa S. Wats. (Malvaceae)

BENEDICT, ROBERT G. (See Largent, Dat Ps ie ae Bonar, LEE, A new Mycocalicium on scarred Sequoia in California . Briccs, BARBARA G., Flora of the Australian Capital Territory (review)

Brown, Roy C. and CLark G. SCHAACK, Two new species of Tragopogon for Arizona

CHIMAL, AuRorA (See GOmez-Pompa et ene “ae Conner, H. E., A naturalized Cortaderia (Gramineae) in California

CooKE, WM. BrincE, H.E. Brown and the plants of the “North side of Mount Shasta”

Coulter, Matcorm, A flora of the Farallon Islands, California

CRAWFORD, DantEL J., Morphology, chromosome number, and flavonoid chemistry of Bzdens cordylocarpa (Compositae)

CRUDEN, RosBert W., Information on chemistry and pollination biology sah to the systematics of Nemophila menziesu (Hydrophyllaceae)

Crum, Howarp, Mosses of unusual interest from Baja California

Curtis, Dwayne H., A preliminary report of the myxomycetes from the state i Idaho

Curtis, DWAYNE H., Myxomycetes new to Gee es National Ee wren I

Daty, Howe tt V., L. H. Shinners’ collection of anthophilous Hymenoptera deposited in Dallas Museum of Natural History

DeDEcKER, Mary (See Wiens, Delbert) — a ae DELGADILLO M., CLrAupio and Dare H. Vitr, New moss es from Mexico

DEMPSTER, ee T. and G. LepyArD STEBBINS, The Galium angustifolium complex (Rubiaceae) of California and Baja California

Dunn, E. L. (See Mooney et al.)

EpDIGER, ROBERT and NIcK SANTAMARIA, Ratibida columnifera (Compositae) in California

Evprence, F. A., II (See McArthur et ab ERNST, ne R., Flora of the Galapagos Islands (ica Every, A. Davin and DELBERT WIENS, Triploidy in Utah aspen

EYERDAM, WALTER J., Botanical collecting rambles with Prof. Eric Hulten in the Aleutian Islands .

FERLATTE, WILLIAM J., Haplopappus Iyallii Gray (Composites), a new record from California

Froticu, E. F. (See Wieland et al.) ee FRYXELL, Paut A., A revision of Phymosia (Malvaceae)

lv

GANKIN, Roman, Nomenclature and interpretation of a

California subspecies in Arctostaphylos (Ericaceae). . . . . . . . . 4147 GENTRY, JOHNNIE L., Jr., A new combination and a

new name in Feckenn (Boraginaceae) . ... . oe es le & 4 OG GILLETT, GEORGE W., Annual review of ecology and aces (review) . . . 494

GILLETT, JoHN M., Two new species of Trifolium (Leguminosae)

from @aionns and Nevada. .. . aa Te = Oe . . 451 GLIESSMAN, STEPHEN R. and Cornetius H. oe The iMeetORe BOeeKEEI

of bracken, Ptertdium aquilinum (L.) Kuhn . .. . a See 299 GOMEz-Pompa, ARTURO, RAFAEL VILLALOBOS-PIETRINI, and Naeaen ne

Studies in the Agavaceae. I. Chromosome morphology and

number of seven species . .... . toh) Rl Ga et, Oe een ae GS Gott ies, L. D., A proposal for classification of fite

annual ee of Stephanomeria (Compositae) . . . . . . . . . . 463 GuzMAN, GASTON (See Trappe, James M.) . ........ . . +. +. 128 Hamann, Marcia J., New distributional records for Washington plants . . . 542

HarpHAM, Care B. and Gorpon H. TRUE, JrR., A floristic study of

Point Arena, Mendocino County, California . . . & oot Ao ere a9 HaArDHAM, CLARE B. and Gorpon H. TRUE, JR., Vere torreyt

(Compositae) in California . . . : ae : ens Harpinc, JAMES and C. B. MANKINEN, coene of ore IIT. Evidence for

genetic differentiation and colonization in Lupinus succulentus (Fabaceae) 222

HIARRISON As Lacoee rooney, LAs) oS af sk. teow cs Oe OS ee 439 Harvey, H. THomas and Ronatp J. MASTROGIUSEPPE, Foxtail pine on

Sitnevtambeak. Gali Oriiia ae lah dicee soles mie eect be ener os! HeEcKARD, LAWRENCE R., The biology of parasitic flowering plants Ces . . 496 HeEtton, NANCYE, Tene WIENS, and Bryan Bartow, High polyploidy and

the origin of Balsamorhiza macrophylla (Compositae). . . . . . . . 526

HickMAN, JAMES C., Arenaria, section Eremogone, (Caryophyllaceae) in the Raciic Northwest: a key and discussions <9... . 203 @ S 2 Goa 20d

Hucues, WILLIAM ELryn, Corallorhiza mertensiana Bong. in ivendeeino Cone, ae Cae oy

IsELY, DuaANnE, Legumes of the U.S. VI. Giiieare. Diherclooncis Peer RS) KJELDSEN, Curis K., Pleurophycus gardneri Setchell & Saunders,

a new alga from northern California . . ......... =. =. =. 416 Kowatsk1, DonaALp T., A new name in Licea (Myxomycetes). . . . . . . 455 KUcHLER, A. W., World vegetation types (review) . ...... . . . 495 Kuijt, Jos, Transfer of Phrygilanthus sonorae to

Psittacanthus (Loranthaceae) ...... . Pe oe ee ee Kyuos, Donan W., Evidence of different adaptations of

flower color variants of Encelia farinosa (Compositae) . . . . . . . 49 LANG, FRANK A., The Polypodium vulgare complex in the Pacific Northwest. . 235 LarcENT, Davin L. and RosBert G. BENEDICT, Studies in the

rhodophylloid fungi. I. Generic concepts . . .... ... . . .~ «32 LayYsER, Ear te F., New distributional records for plants

in the Prac Northwest . . . . = 3 : east ee wo LayseER, EARLE F., Notes on the flora of the Pacific Norenest Pew, ew OO LAYSER, EARLE F. oa H. WAYNE PHILtips, Cytisus scoparius (L.)

ihink intnorth central dahos 44 —. ase: 14 ae & ye a 1 ee SE

MAnKINEN, C. B. (See Harding, James) MASTROGIUSEPPE, RonaLp J. (See Harvey, H. Thomas) .

McArtuHur, E. D., M. T. AtaM, F. A. Evprepce, II, W. Tat, and R. K. eines PRY, Chromocone counts in section Simiolus of the genus Mimulus (Scroph- ulariaceae) IX. Polyploid and aneuploid patterns of evolution .

McC teary, JAMES A., The mosses of the Channel Islands, California

Mooney, H. A., E. L. DUNN, FRANCES SHROPSHIRE, and LEO Sone, Jr., Land-use history of California and Chile as related to the structure of the sclerophyll scrub vegetations .

Mooney, H. A. and A. T. Harrison, The Serectiianl aici on niiie fone: slopes of the Sierra San Pedro Martir in northwest Baja California

MUtteEr, Corne ius H. (See Gliessman, Stephen R.) .

OrNDUFF, ROBERT, A new tetraploid subspecies of Lasthenia (Compositae) from Oregon

ORNDUFF, ROBERT, Flora of New Zealand (review)

OrNDUFF, ROBERT, Systematic studies of Limnanthaceae

ORNDUFF, RoBErRT, Wild flowers of the United States (review) PACKARD, Patricia L., Pinus ponderosa in Malheur County, Oregon. PARNELL, DENNIS R., Plant speciation (review) .

Parsons, Davip J., The southern extensions of Tsuga mertensiana (mountain hemlock) in the Sierra Nevada.

PHILBRICK, RALPH N., The plants of Santa Barbara Island, California Puitiips, H. WAYNE (See Layser, Earle F.) . Se

Puitiirs, Lyte L., A new Gossypium from Guerrero, Mexico .

Porter, Duncan M., Geranium potentilloides in California : PowELt, A. Rice New species of Perityle and Amauria (comneue Powe tt, A. MicHAEL, Taxonomy of Amauria (Compositae-Peritylinae) . Roy, Douctass F., Fasciation in redwood .

RuNDEL, Puizie W., An annotated checklist of the groves of Sequoiadendron

giganteum in a Sierra Nevada, California ne RzEpowsKI, J., Manual of the vascular plants of Texas (review) . SANTAMARIA, Nick (See Ediger, Robert)

SAUER, JONATHAN D., The dioecious amaranths: a new species name and major eye extensions

SCHAACK, CLARK G. (See Brown, Roy C.) oe Scott, FLorA Murray, Stellate epidermal hairs, some 10,000 years old SHROPSHIRE, FRANCES (See Mooney et al.)

SMiTH, RicHarp H., Xylem monoterpenes of Pinus ponderosa, P. washoensis, and P. jeffreyi in the Warner Mountains of California .

SoLsric, Otto T., Polyphyletic origin of tetraploid populations of Gutierrezia sarothrae (Compositae)

Sonc, Leo, Jr. (See Mooney et al.) .

SPELLENBERG, RICHARD, A record from the Klamath Se ee ton a San See. monotypic genus, Whitneya (Compositae)

SPELLENBERG, RICHARD, Two species of Panicum (Poaceae) new to Oregon STEBBINS, G. Lepyarp (See Dempster, Lauramay T.) .

STEPHENSON, STEPHEN N., A putative Distichlis X Monanthochloe (ESS) hybrid from Baja Galore Mexico.

vi

aes 152

417 435

305

439 299

96 450 103 492 298 130

536 O29 436 265 449 456 516 462

og 174 12

426 304 458 305

26

20 305

449 102 70

125

STICKNEY, PETER F., Crupina vulgaris (Compositae: Cynareae), new to Idaho fod North America .

SUNDBERG, WALTER J., The genus Chlorophyllum GeuotceES in California

Tar, W. (See McArthur et al.) Sos: Tuers, Harry D., The savory wild mushroom oa

Tuters, Harry D. and Roy WATLING, Secotiaceous fungi from western United States

THomas, JoHN H., Botanico-Periodicum- An uanan (review) . THOMAS, JOHN H., Trifolium hirtum L. (Fabaceae) in California Tuomas, JoHN H., Wallace Roy Ernst, 1928-1971 es Tomes, A. SPENCER, Taxonomy of Chaetadelpha (Compositae: GeioneaS

TRAPPE, JAMES M. and Gaston GuzMAn, A newly determined species of Elaphomyces from Oregon

TRUE, Gorpon H., Jr. (See Hardham, Clare B.) Tucker, J. M., Hermaphroditic flowers in Californian oaks

TurRNER, B. L., A new species of Dyssodia (Compositae) from north central Mexico

VicKErRY, R. K., Jr. (See McArthur et al. ) VILLALOBOS-PIETRINI, RAFAEL (See GOmez-Pompa et al.) Vitt, Date H. (See Delgadillo M., Claudio)

WALKINGTON, Davin L., Cacti of the Southwest—Texas, New Nicene Oklahoma, a eaneas, and Louisiana (review) .

Wattace, A. (See Wieland et al.) WATLING, Roy (See Thiers, Harry D.) . sen , WEBER, WILLIAM A., Mimulus gemmiparus sp. nov. from Calbeee : WELLs, Puitip V., The manzanitas of Baja California, including a Hea species of Arctostaphylos ee ae ‘ WETZEL, CHERIE LALAINE Rivers, The significance of Niles Canyon in ne phytogeography of the Coast Ranges of Central California

WIELAND, P. A. T., E. F. Frovicu, and A. WALLACE, Vegetative rantencon of

woody shrub eneces from the northern Mojave and southern Great Basin deserts

Wiens, DELBERT (See Every, A. David) Wiens, DELBERT (See Helton et al.)

Wiens, DELBERT and Mary DEDECcKER, Rare natural hybridization in Phoradendron (Viscaceae)

WiLzBur, Rosert L., A flora of tropical Florida eae

WILKEN, DIETER H., Seasonal dimorphism in Baccharis glutinosa On,

Wricut, Rosert D., Local photosynthesis ecotypes in Pinus attenuata as related to alitade

YOUNGBERG, ALv Dan, The intrageneric position of Ghee orestera

Vii

ERRATA

p. 70. Second paragraph, third line, should read “. . . subject of the present paper...” p. 71. Map: The small dot under the N of SAN BERNARDINO should be a square. p. 423. Line one should read Mimulus for Minulus.

p. 455. Line one in Note should read Licea deplanata for Licea deplanata.

Dates of publication of Madrono, Volume 21

No.1,pp. 1- 48: 30Sep 1971 No. 5, part one, pp. 265-328: 10 Mar 1972 No.2, pp. 49-112: 9 Nov 1971 No. 5, part two, pp. 329-393: 25 May 1972 No. 3, pp. 113-176: 17 Feb 1972 No. 6, pp. 395-450: 15 Jun 1972 No. 4, pp. 177-264: 3 Mar 1972 No. 7, pp. 451-498: 7 Aug 1972

No. 8, pp. 499-546: 6 Nov 1972

' CK

/ Vi 18 35

a VOLUME 21, NUMBER 1 JANUARY, 1971

Contents

SECOTIACEOUS FUNGI FROM WESTERN UNITED STATES,

Harry D. Thiers and Roy Watling 1 MyxomycetTes NEw To CRATER LAKE NATIONAL Park, OreEcon. I. Dwayne H. Curtis 10

TRANSFER OF PHRYGILANTHUS SONORAE TO PsITTACANTHUS (LORANTHACEAE), Job Kuzjt 13

THE GENUS CHLOROPHYLLUM (LEPIOTACEAE) IN CaLrrorniA, Walter J. Sundberg 15

POLYPHYLETIC ORIGIN OF TETRAPLOID POPULATIONS OF GUTIERREZIA SAROTHRAE (CoMposiTaE), Otto T. Solbrig 20

XYLEM MONOTERPENES OF PINUS PONDEROSA, P. WASHOENSIS, AND P. JEFFREYI IN THE WARNER

Movuntains oF CALirorniA, Richard H. Smith 26 STUDIES IN THE RHODOPHYLLOID FuncI. I. GENERIC

Concepts, David L. Largent and Robert G. Benedict 32 A NATURALIZED CORTADERIA (GRAMINEAE) IN

CatirorniA, H. E. Connor 39

MorPHOLOGY, CHROMOSOME NUMBER, AND FLAVONOID CHEMISTRY OF BIDENS CORDYLOCARPA (COMPOSITAE),

Daniel J. Crawford 4] Notes AND NEws: RATIBIDA COLUMNIFERA (COMPOSITAE) IN CALIFoRNIA, Robert Ediger and Nick Santamaria 2

NOTES ON THE FLORA OF THE PacrFic NORTHWEST, Earle F. Layser 47

A WEST AMERICAN JOURNAL OF BOTANY

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established 1916. Individual subscription price $8.00 per year ($4.00 for students). Institutional subscription price $12.00 per year. Published quarterly in January, April, July, and October by the California Botanical Society, Inc., and issued from the office of Madrofio, Herbarium, Life Sciences Building, University of California, Berkeley, California. Orders for subscriptions, changes in address, and undelivered copies should be sent to the Corresponding Secretary, California Botanical Society, Depart- ment of Botany, University of California, Berkeley, California 94720.

BOARD OF EDITORS CLASS OF:

1971—Marion OwnsBEy, Washington State University, Pullman Joun F. Davinson, University of Nebraska, Lincoln

1972—Ira L. Wiccins, Stanford University, Stanford, California REED C. Rotiins, Harvard University, Cambridge, Massachusetts

1973—WattacE R. Ernst, Smithsonian Institution, Washington, D.C. Roy L. Taytor, University of British Columbia, Vancouver

1974—KeEnTon L. CHAMBERS, Oregon State University, Corvallis EMLEN T. LiTTEL, Simon Frazer University, Burnaby, British Columbia

1975—ArtTuRO GoMEz PomPaA, Universidad Nacional Autonoma de México Duncan M. Porter, Missouri Botanical Garden, St. Louis

1976—Dennis ANDERSON, Humboldt State College, Arcata, California KrincsLey R. STERN, Chico State College, Chico, California

Editor — Joun H. THoMAs Dudley Herbarium, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JuNE McCaskKILy P.O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Lawrence R. Heckard, Department of Botany, University of Califor- nia, Berkeley. First Vice-President: Donald Kyhos, Department of Botany, Uni- versity of California, Davis. Second Vice-President: Dennis Anderson, Department of Botany, Humboldt State College, Arcata. Recording Secretary, John West, De- partment of Botany, University of California, Berkeley. Corresponding Secretary: Dennis R. Parnell, Department of Biological Science, California State College, Hay- ward. Treasurer: June McCaskill, Department of Botany, University of California, Davis.

The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Marion Cave, Department of Botany, University of California, Berkeley; the Editor of Madrofio; and three elected Council Mem- bers: Malcolm Nobs, Carnegie Institution of Washington, Stanford; Elizabeth McClintock, Department of Botany, California Academy of Sciences, San Francisco ; and Thomas Fuller, California Department of Agriculture, Sacramento.

SECOTIACEOUS FUNGI FROM WESTERN UNITED STATES

Harry D. THIERS and Roy WATLING

Department of Biology, San Francisco State College, San Francisco, California 94132 Royal Botanic Garden, Edinburgh, Scotland

It is always pleasing to obtain good material of any unfamiliar fleshy basidiomycete but particularly material of species which are at the center of discussion and conjecture, whose name is based on some ob- scure collection or whose name is based on collections inadequately de- described. These conditions apply for most any secotiaceous fungus, for it appears that collections are so irregularly made that affinities are always in dispute. So much of our knowledge of the bolbitiaceous and strophariaceous gastromycetes is based on single collections that Singer (1963) considers only one species of Galeropsis, G. andina, has been obtained in enough quantity to study it really extensively. When several collections of two species of so-called Galeropsis from widely separated areas came to hand, the opportunity was taken to critically review their taxonomy. The two fungi can be assigned to Galeropsis cucullata and G.. polytrichoides.

A note on recent collections of another secotioid fungus, Setchellio- gaster, appends the discussion.

Smith has recently (1965) described Weraroa coprophila (figs. 1-3) from Payette National Forest, Idaho, which is distinguished primarily by the lack of a spore-print, the color of the gleba, carpophore colora- tion, the spore dimensions and spore attachment to the basidium. He drew attention in his account of the similarity of his fungus to Bolbitius cucullaitus Seaver & Shope, the type of which was examined by one of us (R.W.) during the tenure of a National Science Foundation Grant (G 13282-03779, made available to me while at the University of Mich- igan). The hymenial color of the type approaches “bone brown” of Ridg- way and thus closely resembles that of many dark-spored agarics, i.e., Psilocybe spp. and Stropharia spp. Even under the microscope the spores have a slight purplish flush, and do not approach the gill colors found in either Agrocye or, as indicated by Singer (1963), in Conocybe.

Opportunities have been taken over several years by one of us(R.W.) to examine the types of many so-called bolbitiaceous fungi, both agari- coid and secotioid. Bolbitius cucullatus, it seems, is unique in this group in some of its anatomical details. The cheilocystidial shape (fig. 8), pore characters and gill color lead one to place this species along with Smith’s fungus in the strophariaceous genus Weraroa as it is presently under- stood. Galeropsis (type species—G. destertorum Vel.), the genus to which Bolbitius cucullatus was transferred by Singer (1936), differs

Maprono, Vol. 21, No. 1, pp. 1-48. September 30, 1971.

1

2 MADRONO [Vol. 21

markedly in gill and spore color; moreover it is considered by us (Wat- ling, 1964), as it stands at present, to be an artificial assemblage of secotiaceous fungi, and a revision is being prepared.

The following new combination is proposed:

Werarea cucullata (Seaver & Shope) Thiers & Watling, comb. nov. Bolbitius cuculattus Seaver & Shope, Mycologia 27:649. 1935. Galerop- sis cucullata (Seaver & Shope) Singer, Beih. Bot. Centralb. 56:137. 1936. Secotium long pipes Zeller, Mycologia 33:209. 1941.

Pileus 9-35 mm high, 4.5-15 mm broad at the base, not or hardly expanding, typically conic often ending in a long, relatively sharp point which may curl on drying (fig. 7), wrinkled, fibrillose to fibrillose-scaly from remains of a floccose, “‘mustard yellow” (Color terms defined in Ridgway (1912) are placed in quotes.) to “‘ochraeous tawny” veil, dry, almost “apricot yellow” to “mustard yellow” when young becoming rich “chrome yellow” at the base, finally more ‘old gold” or “buffy citrine” at the base and “tawny” or near “buckthorn brown” toward the apex; margin incurved and tightly fitted around the stipe, joined in the young specimens to the stipe by a distinct, although fugacious, veil which leaves distinct flecks and which can be seen even in the most mature pilei upon careful examination; sometimes becoming free and expanded but retaining a frill of velar flecks. Stipe columella 50-110 « 1—4 mm, equal or slightly swollen at the base in larger specimens, dry with appressed fibrils at the apex or fibrillose toward the base from velar remnants, “pale ivory yellow” at apex and near ‘“‘Saccardo’s umber” to “sepia”? downwards, then “ochraceous buff” to “clay color” at the apex with hint of cinnamon yellow but base unchanging. Gills fusing and anastomosing to form elongate compartments, “‘Verona brown” at first then becoming flushed with “warm sepia,” finally “bone brown” except for white or pale yellow margin. Flesh soft, yellowish in pileus, more buff in stipe, particularly in the base, and drying bright yellow; odor and taste not unpleasant, mild.

Basidia 4-spored, hyaline to slightly colored in KOH, clavate or con- stricted about the middle, 25-35 10-12 yu, pedicellate. Basidiospores (fig. 9) 11.5-13 (14.5) & 6.5-7.5 (8) yu, elliptic to slightly amygdali- form, flattened in side view with a low shoulder over a prominent, al- though not large, apiculus, thick-walled, truncate by a slightly eccentric or apical pore, deep honey with a bister tint in water and KOH. Pleuro- cystidia rare, up to 25 long, langeniform and slightly mucronate, mixed with more vesiclulose cells up to 15 broad, described by some as pseudoparaphyses. Cheilocystidia (fig. 8) numerous, forming a distinct fringe on the gill, filamentous-cylindric, hyaline or slightly yellowish, 35-70 X 5-7.5 uw; caulocystidia lageniform, up to 30 » long and 5-8 pu broad at apex. Hymenophoral trama of irregularly arranged, + enlarged cells some of which have yellow contents. Pileus trama of irregularly ar- ranged, + swollen cells and up to 16 » in width with some colored con-

1971] THIERS & WATLING: FUNGI 3

Fics. 1-9: 1-3, Weraroa coprophila; 1, carpophores, * 1; 2, cheilocystidia, x 400; 3, basidiospores, &K 900; 4-6, Galeropsis polytrichoides; 4, carpophores, xX 1; 5, cheilocystidia, « 400; 6, basidiospores, * 1000; 7-9, Weraroa cucullata; 7, carpophores, X 1; 8 cheilocystidia, * 400; 9, basidiospores, « 800.

4 MADRONO [Vol. 21

tents and covered in a distinct filamentous pellicle of + colored, + faintly colored granulose cells 2-3 » in diam. and up to 35 vp» long. Clamp connections infrequent but present.

On ground among grass and sedges, elev. 9,600 ft, University of Wyo- ming Summer Camp, near Laramie, Wyoming, 2 July 1929, F. J. Seaver a F. Shope (Type: MICH, ex herb. Univ. Colorado No. 249. A perma- nent slide is also in NY).

Gregarious, often in large troops, in boggy or marshy areas; elev. 7,500 ft., Haskner Creek, Sierra Madre, Carbon Co., Wyoming, June, 1950, Smith 36246 (MICH); Hot Lake Area, elev. 6,300 ft., Lassen Volcanic Nat. Park, Shasta Co., California, 8 June, 1965, Thiers 12923 (E, San Francisco State College); Silver Lake, elev. 7,200 ft., Amador Co., California, 6 June, 1966, Thiers 16904 (E, San Francisco State College).

This fungus is very close to Weraroa coprophila as already indicated, and it may be necessary to reassess the position of the Idaho fungus when more material is collected. When the type collections of the two species are compared one gets the impression that W. coprophila (fig. 1) is a much more robust fungus. Smith (1965), however, has indicated three main differences between his taxon and W. cucullata; namely, the darkening stipe, the smoky grey to pale fuscous gills when fresh and the narrow spores (fig. 3). The first difference still applies, however the stipe is very dark at the base from the beginning in W. cucullata; the gills of W.. cucullata are nearer ‘Verona brown” than has been indicated earlier and, after comparing the types of both species directly, one be- sides the other, the colors are indeed very close. The spores are slightly narrower but one must treat this with some caution for the statement is based on only a single collection. Similarities are also seen in cheilo- cystidial shape. Although Smith did not observe cheilocystidia, they are present in the type material of W. coprophila and resemble those of the type of W. cucullata; the contents are much more yellow in KOH in W. coprophila. Now that more and more collections are being made of gastromycetoid fungi, the full range of differences between individuals, which belong to the same taxon, are becoming evident.

Galeropsis polytrichoides was described by Zeller (1941) from collec- tions made by W. B. Cooke on Mount Shasta, and, although not found for several years it has appeared in the last few collecting seasons in sev- eral localities elsewhere in California. A full description based on these fresh collections and subsequent microscopic examination is given.

GALEROPSIS POLYTRICHOIDES (Zeller) Zeller, Mycologia, 35:410. 1943. Figs. 4-6. Secotium polytrichoides Zeller, Mycologia 33:211. 1941.

Pileus 5-15 mm high, 3-10 mm broad at the base, narrowly to acutely conic, subacute or ellipsoid or pinched in at the top, attenuate or be- coming truncate at the base at maturity, moist or slightly lubricous when fresh and immature but soon becoming dry; glabrous to silky

1971] THIERS & WATLING: FUNGI 5

appressed fibrillose, becoming rimose, tan on the disc, darker towards the margin, becoming “‘sudan brown” to “bone brown” at maturity, drying with distinctly paler disc and in herbarium material the disc is ‘‘ochra- ceous tawny” to “yellow ochre” and the margin “date brown” to almost “umber”; margin joined to the stipe when young with white fibrils which darken slightly with age becoming very pale tan to ochraceous and sep- arate at maturity, forming a fimbriate fringe with long dangling “‘ciliae” up to 10 mm long. Veil consisting of parallel hyphae 25-42 (68) » long and 4.5—7 » wide, fused or adhering tightly together. Stipe columella 30-90 mm X 1-2 mm, equal or slightly enlarged at the base, glabrous to silky fibrillose often showing a twisted pattern of fibrillosity, tan to “buckthorn brown”’, finally ““Verona brown” and in dried material with a peculiar steel blue at the very base when seen in certain light. Gills joined by a few thin, anastomosing lamellar partitions, pale brown, on drying becoming ‘“‘buckthorn brown” to ‘“‘Prout’s brown” to “raw umber” in dried material. Flesh pale tan in pileus and upper stipe, darker brown in lower stipe.

Basidia (1-, 2-) 4-spored 25-29 x 7-8 yp, short pedicellate, usually hyaline in KOH although some are distinctly yellow-brown. Basidio- spores (fig. 6) 10-13 « 5-6 X 6-6 pn, ellipsoid, slightly flattened in side view and swollen upwards in face view (obovate of Zeller), slightly tapered, with an apical germ-pore, although fairly thick-walled not strongly pigmented. Pleurocystidia not found; cheilocystidia (fig. 5) lageniform to slightly subcapitate with apex 5-6 (8) » broad and up to 25 uw long. Caulocystidia similar to cheilocystidia, although more vari- able in shape and often with a tapered apex, 20-42 « 18-12 mw (apex 8.5-10 w or 1.5—2 »). Hymenophoral trama consisting of interwoven, swollen hyphae up to 8.5 » broad. Pileus trama filamentous, consisting of interwoven, swollen cells up to 17 » broad with a distinctly filamentum cutrill of hyphae 2-4 » in diameter. Clamp connections present.

Gregarious on moist ground, in boggy areas about streams, and in open seepage areas in conifer woodlands; Lassen Volcanic National Park, elev. 6,000 ft., Shasta Co., California, 3 July, 1965, Thiers 12912 (EF, San Francisco State College); Big Meadows Campground, elev. 7,000 ft., Calaveras Co., California, 30 May, 1966, Thiers 16874, 16875, and 16878 (E, San Francisco State College); Sagehen, California, 29 May, 1966, D. McLaughlin (E).

It must be pointed out that in all the collections examined the basidio- spores are slightly larger than those indicated in the original description. However, authentic material collected on 24 July, 1941, by W. B. Cooke from a site only a few yards from the type locality and now in MICH under No. 136 Mycobiota of North America, Mycobiota of Mt. Shasta, California, is similar to our collections in every way. The type material has not been examined.

A very interesting observation which needs further ecological examina-

6 MADRONO [Vol. 21

Fic. 10. Setchelliogaster tenuipes, X 1.

tion is that on these different occasions Werarora cucullata and Galerop- sis polytrichoides have been found at the same localities, in Lassen Na- tional Park, in Silver Lake area and in the type locality for G. polyiri- choides (from which the type of Secotium longipes, =W. cucullata fide Singer, was also described).

We believe that all Galeropsis-like fungi are xerophytic derivatives, many of which are, perhaps, quite restricted in distribution, of a whole series of familiar agaric groups. It does not follow, however, that we think all secotiaceous fungi exhibit convergance in this way and are such end-products. We envisage these gastromycetes as paralleling the aquatic mammals; although the latter are all adapted to water and live successfully there, they can be quite unrelated, except that they are all mammals.

Several factors convince us that they are end-products: particularly the well-developed stipe, the well-developed cheilocystidia, the well- developed cuticular structure of the stipe and pileus, and the lamellar nature of the “gleba.” It has been shown by one of us (R.W.) that in Conocybe farinacea dye entering a water column maintained in the stipe finally finds its way into the cheilocystidia and then into external, apical droplets. It is postulated that the cystidia which develop on the most exposed parts of the gill have developed in response to water uptake

1971] THIERS & WATLING: FUNGI d

phenomena. This will be expanded in greater detail elsewhere but the question must be posed here, if this is so what would be the part played by the cheilocystidia in such a fungus as Galeropsis cucullata and when would they begin to play a part in the economy of the gill-edge if evolved within the fruit body.

Should, therefore, the Secotiaceae or the majority of its members be placed alongside the units to which they are closest? Martin (1956) has already expressed similar ideas on the subject but from a more general point of view. Also should not Weraroa and Galeropsis be reorganized into genera which show some relationship to the geophiloid agarics and separate out the bolbitiaceous element, if one exists?

There is little doubt that these questions will cause much discussion and possibly disagreement and we sincerely hope it does, so that, when further material is available to us, it can be examined with new thoughts in mind; we are anxious to see more material to support our present approach.

Pouzar (1958) described the genus Setchelliogaster in 1958 basing the taxon on Secotium tenuipes, a fungus originally described by Setch- ell (1907) who collected it on the University of California campus at Berkeley, California. As early as 1950 Heim discussed the possible con- nection between this fungus and the bolbitiaceous agarics mainly on the basis of the cellular cuticle and spore color, but its affinities are still problematic. It is always difficult to make convincing comparative studies from limited herbarium material; we are, for example, ignorant of how the cuticle of this fungus actually develops. The development of such carpophore tissues is considered by Reijnders (1963) and Watling (1963) to be very important in the understanding of relationships be- tween different groups of higher fungi. After examination of several herbarium collections it appears that the cuticular structure, although cellular, does not, in fact, develop in the same way as that of typical members of the Bolbitiaceae, e.g., Conocybe farinacea, Bolbitius vitel- linus, or Agrocybe praecox. The cells of the outer layer are not pyriform with the pedicels developing from an active zone as described by Wat- ling (1964), but more studies are required.

All the studies so far carried out were based on material from the small area embraced by Alameda and Santa Clara counties, California, al- though Singer and Smith (1959) mention a collection in the New York Botanical Garden from Oregon. The material described below is from San Francisco State College Campus and adjacent Marin Co. and are the only recent collections with full field notes made by one of us (H.T.). Grateful appreciation is extended to Mrs. Alix Wennekens for transmit- ting the collections from Marin Co.

SETCHELLIOGASTER TENUIPES (Setchell) Pouzar, Ceska Mykol. 12:34. 1958. Fig. 10. Secotium tenuipes Setchell, J. Mycol. 13:239. 1907, Gastro-

8 MADRONO [Vol. 21

carp 5—25 mm high, 10-20 mm broad at maturity; convex to somewhat cylindric to globose when fully developed, never completely expanding, dry to moist, glabrous, colored near “Hessian brown” to “carob brown” when young, unchanging with age or becoming near “‘hazel” to ‘‘Kaiser brown” to “chestnut brown”; unchanging when bruised; margin typic- ally incurved during all stages; when young typically attached to the stipe columella by white veil fragments, usually breaking free with age, but always remaining incurved to straight, entire to often eroded. Stipe- columella distinct, well-developed and apparent in all gastrocarps, 5—20 mm long, 3—4 mm broad, solid, extending through the entire gleba, more or less concolorous with the gleba or peridium, surface dry, more or less appressed fibrillose; white mycelium at the base. Gleba distinctly lamel- lose to broadly and highly irregular lacunose; lamellae plates very thin and fragile but not becoming pulverulent with age or when dried, typ- icallly folded, convolute and irregular, strongly intervenose with numer- ous, large air spaces interspersed, “ochraceous tawny” to “hazel” dur- ing all stages of development. Peridium less than 1 mm thick, fragile, concolorous with the surface, unchanging when exposed; taste and odor mild.

Basidia 27-38 8-12 p, typically 2-spored with the sterigmata often large and strongly developed. Basidospores 15-17.6 & 9-11.2 p», ovoid to subellipsoid, with well-developed sterigmal appendage, germ-pore typically present but often poorly differentiated and difficult to inter- pret, bright ochraceous to “tawny” in KOH and Melzer’s, surface rough- ened, episporium with numerous small canals extending from inner-wall to surface and sometimes causing an obscure reticulation on the surface. Cystidia not seen, but often large, + cylindric to fusoid cells with obtuse apices present in hymenium (possibly undeveloped basidia). Trama of lamellae plates irregular to subparallel, staining pale ochraceous to yel- low in KOH and Melzer’s. Peridial trama typically loosely interwoven; cutis differentiated as a layer of globose to vesiculose cells, “ochraceous tawny” in KOH and Melzer’s, wall not incrusted, 12-18 » in diam.

Gregarious in soil under Eucalyptus globulus and Cupressus macro- car pa. All collections except one made by Mrs. Alix Wennekens in vicin- ity of Mill Valley, Marin Co., California, Thiers 21893, 21911, 29112, 21913, 21914, and 21924. An additional collection was made on the cam- pus of San Francisco State College, San Francisco, California, Thiers 12413. All collections are deposited in the herbarium of San Francisco State College.

Setchell (1907) noted this fungus growing under Eucalyptus globulus, and Quercus spp. in the vicinity of Eucalyptus spp.. Our more recent collections are from under Eucalyptus globulus and Cupressus macro- carpa; the latter, although native to California, has presumably been planted in Marin Co. Eucalyptus spp. are mainly native of Australia, a few extending into the East Indies, and it is interesting to speculate

1971] THIERS & WATLING: FUNGI 9

as to the habitat of Setchell’s fungus. Various Eucalyptus spp. are widely planted now for ornament in Mexico, South Africa, the southern states of the United States, India, S.E. Asia, etc., and it is possible that native species of fungi have taken up association (mycorrhizal or not) with Eucalyptus when once introduced, e.g., Scleroderma in India (Thayer, et al., 1967), Pisolithus tinctorius in S. Africa (Thayer, et al., 1967). The qeustion arises as to whether Setchelliogaster is an endemic west- ern North American fungus or has it been introduced along with the Eucalyptus trees? It has not been found in Australia yet, but there is every possibility that a fungus can exist for very long periods as a my- corrhizal associate simply reproducing vegetatively or being distributed along with seedlings of mature trees and rarely producing fruit bodies. The climate offered by California may be more favorable to the fungus than its native area, and so regular fruiting can be accomplished in its new environment.

LITERATURE CITED

Herm, R. 1950. Le genre Galeropsis Vel. (= Cytarrophyllum Heim) trait d’union entre agarics and gastrales. Rev. Mycol. (Paris) 15:3-28.

Martin, G. W. 1956. On Lentodium squamulosum. Proc. Iowa Acad. Sci. 63:280—286.

Pouzar, S. 1958. Nove rody vyssich hub. Ceska Mykol. 12:31-36.

ReIJNDERS, A. F. M. 1963. Les problems du development des carpophores des agaricales et de quelques groupes voisins. Hague.

Ripcway, R. 1912. Color Standards and color nomenclature. Washington, D.C.

SETCHELL, W. A. 1907. Two new hypogaeous Secotiaceae. J. Mycol. 13:236-241.

SINGER, R. 1936. Studien zur systematik der basidiomyceten. Beih. Bot. Centralb. 56:137-174.

. 1960. Three new species of Secotiaceae from Patagonia. Persoonia 1:385— 391.

. 1963. Notes on secotiaceous fungi Galeropsis and Brauniellula. Kongl. Ned. Akad. Wetensk. 66:106-117.

. and A. H. Smiru, 1958. Studies on secotiaceous fungi. III. The genus Weraroa. Bull. Torrey Bot. Club 85:324-334.

. 1959. Studies on secotiaceous fungi. VI. Setchelliogaster Pouzar. Madrono 15:73-79.

SmitH, A. H. 1965. New and unusual basidomycetes with comments on hyphae and spore-wall reactions with Melzer’s solution. Mycopathol. Mycol. Appl. 26:385-402.

TuHaver, H. S., S. BALwart, and B. K. Baxksut. 1967. Mycorrhizae in Eucalyptus. Indian Forester 93:756—763.

WartTLinc, R. 1963. In RetyNvErS, A. F. M. Les problems du development des car- pophores des agaricales et de quelques groupes voisins. Hague.

. 1964. Taxonomic characters of the Bolbitiaceae with particular reference to the genus Conocybe. Ph.D. thesis. Edinburgh Univ.

—. 1965. Observations on bolbitiaceae. 2. Conspectus of the family. Notes Roy. Bot. Gard. Edinburgh 10:287-323.

ZELLER, S. N. 1941. Further notes on fungi. Mycologia 33:196-204.

MYXOMYCETES NEW TO CRATER LAKE NATIONAL PARK, OREGON. I.

DWAYNE H. CurrTIs

Department of Biological Sciences, Chico State College, Chico, California 95926

During the winter months an over-abundance of precipitation gener- ally in the form of snow is deposited in the High Cascades of Crater Lake National Park. Each year, the accumulated annual snow depth often exceeds 50 feet from November to May. Early in the spring slimemolds or Myxomycetes are frequently exposed as the snow melts away from forest litter and fallen logs. Decaying, moist organic matter such as bark, wood, and fallen twigs is characteristically noted as the habitat for Myxomycetes. In contrast, the summer is normally quite dry since very little rain falls during the months of July and August. However, during mid-August, 1968, succeeding storms (snow followed by several days of rain) brought 5.34 inches of precipitation which almost doubled any previous weather record set in the park for that month. Following the storms in late August and early September many specimens of slime- molds were collected. Interestingly, some of these are not commonly found in montane areas.

Most of the Myxomycetes reported thus far for Crater Lake National Park were recorded by Peck and Gilbert (1931) as occurring from the High Cascades to the Coast Range in northwestern Oregon. Exceptions include one species reported by Martin (1932), three new species de- scribed by Kowalski (1966, 1968), and I (Curtis, 1968) mentioned Barbeyella minutissima Meylan as well as eight additional species (Curtis, 1969) as being new to the State of Oregon.

All the specimens for this paper were collected on some form of decay- ing wood at elevations from 6,000 to 7,000 feet during the summer of 1968. Previously, this author (Curtis, 1969) indicated the presence of 43 different species in a preliminary report of the slimemolds from the park. The 11 species, included here, bring the total number of Myxomy- cetes found in Crater Lake National Park to 54. At least one collection of each species has been deposted in the University of Iowa Herbarium, Iowa City, Iowa and where possible, duplicate specimens have been given to the Crater Lake National Park Herbarium, Crater Lake, Ore- gon. The numbers for the collections are my own and they indicate only those specimens given to the University of Iowa Herbarium. The names of the organisms are those accepted by Martin (1949).

RETICULARIACEAE

Reticularia splendens Morgan. On a decorticated fallen log, Kerr Val- ley, 6,800 feet, 1522, Sept. 4, 1968. Only two aethalia were found, one 5 mm and the other 13 mm in diameter. Both were brownish-copper col- ored with a white conspicuous margin about the base of the hypothallus. Spores were generally reticulate over two-thirds of their surfaces.

10)

1971] CURTIS: MYXOMYCETES 1a

TRICHIACEAE

Arcyria globosa Schw. On a decorticated fallen coniferous log, west Goodbye Bridge area, 6,100 feet, 1479, Aug. 30, 1968. Several small, short-stalked sporangia about 0.4 to 0.8 mm in diameter were scat- tered throughout the grooves of a decaying log. The peridium, in most cases, was fugacious while the capillitium and spores were the typical drab, ashen-grey color. Several specimens were collected at elevations from 6,000 to 7,000 feet.

A. incarnata (Pers.) Pers. On decayed wood, Kerr Valley, 6,800 feet, 1520, Sept. 4, 196&. The sporangia were generally short stalked with a saucer-like base from which there was a greatly expanded, loose capil- litum. Most were rosaceous to brown in color. This slimemold was found in many areas of the park.

STEMONITACEAE

Enerthenema papillatum (Pers.) Rost. On decayed wood, 2 miles west of Annie Springs, 6,000 feet, 1442, Aug. 27, 1968. The sporangia, for the most part, had a total height ranging from 0.8 to 1.5 mm. The stipe expanded at the tip of the columella to form a disk from 0.05 to 1.5 mm in diameter. The peridium was fugacious and the spores appeared to be black in mass.

Stemonitis axifera (Bull.) Macbr. On decayed wood, west Goodbye Bridge area, 6,100 feet, 1476, Aug. 30, 1968. This collection was obtained on the underside of a decayed stump. The sporangia were densely clus- tered, a bright rusty-brown, and 7-8 mm in height. The spores (about 5uin diameter) were minutely warted.

S. palliada Wing. On decayed wood, northeast of Goodbye Bridge area, 6,100 feet, 1461, Aug. 29, 1968. Small clusters of 5 to 10 sporangia (3-4 mm high) were found scattered over an area of approximately 4 square millimeters. The sporangia were slightly elongate-ovate in shape and lilaceous-brown in color.

S. hyperopia Meylan. On decayed wood, 2.4 miles southeast of Park Headquarters, 6,700 feet, 1423, June 23, 1968. The sporangia were sometimes scattered but generally gregarious in small clusters, lilac- brown in color and 1—2 mm tall.

PHYSARACEAE

Leocarpus fragilis (Dicks.) Rost. On decayed wood, Kerr Valley, 6,800 feet, 1518, Sept. 4, 1968. Sporangia were gregarious with a shiny, smooth, reddish-brown and very brittle peridium. The spores appeared black in mass.

Physarum leucopus Link. On needles and fallen twigs, west side of Munson Ridge, 6,900 feet, 1500, Sept. 1, 1968. Sporangia were some- what scattered. The peridium and stalk were frosty-white with lime deposits.

P. newtont Macbr. On a decayed broken limb, northwest of Goodbye Bridge area, 6,200 feet, 1443, Aug. 27, 1968. The two collections con-

12 MADRONO [Vol. 21

tained hundreds of gregarious sporangia, stalked, bright rose-purple with red lime nodes. Most stalks were as long or longer than the diameter of the sporangium, thereby, differing from the descriptions of Lister (1925) and Martin (1949) who both indicated that the sporangia were short-stalked or sessile. The stalk was not translucent and therefore this Myxomycete could not be Physarum roseum Berk. & Br. Although this Myxomycete has been reported previously from Oregon (Peck and Gil- bert, 1931), it is considered rare.

P. notabile Macbr. On bark and wood of a fallen coniferous tree, west side of Munson Ridge, 6,900 feet, 1498, Sept. 1, 1968. This slime- mold was primarily sessile with a few sporangia merging into short plasmodiocarps while others had short, furrowed stalks. The peridium appeared to be uncrusted with an ashy, bluish-white lime deposit.

This study was supported in part by the Chico State College Founda- tion, Grant GU 2690.

I am indebted to Donald T. Kowalski of Chico State College and to George W. Martin of the University of Iowa for their assistance through- out the course of this investigation.

LITERATURE CITED

Curtis, D. H. 1968. Barbeyella minutissima, a new record for the Western Hemis- phere. Mycologia 60: 708-710. . 1969. New records of Myxomycetes from Oregon. I. Madronio: 20:75-77. . 1969. A preliminary report of the Myxomycetes of Crater Lake National Park, Oregon. Madrono: 20: 278-282. KowatskI, D. T. 1966. A new species of Lamproderma from California. Mycologia 58: 808-810. . 1968. Three new species of Diderma. Mycologia 60: 595-603. . 1968. Observations on the genus Lamproderma. Mycologia 60: 756-768. Lister, A. 1925. A monograph of the Mycetozoa. 3rd ed. by G. Lister, Brit. Mus. Nat. Hist. London. Martin, G. W. 1932. New species of slime molds. J. Wash. Acad. Sci. 22: 88-92. . 1949. North American Flora 1:1-190. Peck, M., and H. Girpert. 1931. Myxomycetes of northwestern Oregon. Amer. J. Bot. 19: 141-147.

NOTES AND NEWS

RATIBIDA COLUMNIFERA (COMPOSITAE) IN CALIFORNIA.—Ratibida columnifera has previously not been known from California, although it is widespread from British Columbia to Minnesota and south to Arizona, Mexico, and Tennessee. Recently we collected specimens of this species on dry rocky soil on the west side of Eagle Lake in Lassen Co. at the Eagle Lake Field Station (Santamaria & Ediger 721, CAS, UC). The local population consists of about 100 plants—Rosert Epicer, Chico State College, Chico 95926, and Nick SantamartiA, Tahoe-Truckee High School, Truckee, California 95734.

TRANSFER OF PHRYGILANTHUS SONORAE TO PSITTACANTHUS (LORANTHACEAE)

Jos Kuljt

Department of Biology, University of Lethbridge, Lethbridge, Alberta, Canada

When Phrygilanthus sonorae (Watson) Standley, a mistletoe from southern Baja California and adjacent Sonora, was first assigned to Phrygilanthus (Standley, 1919) the genus had representatives in both the Old and the New World. More recently, concepts have swung to the exclusion of Old World species (Barlow, 1962). Even the New World remnant is currently under considerable question as a coherent tax- onomic unit. It is no wonder that the appropriateness of P. sonorae in this genus is also at issue. In particular, the possibility of the species belonging in Psittacanthus seemed to warrant exploration.

The major criterion of Psittacanthus at this time is the absence of endosperm in the mature fruit. All other Loranthaceae, sensu stricto, have endosperm. A recent study of Bhatnagar and Chandra (1968) has, however, demonstrated the presence of normal endosperm in Psitta- canthus cuneifolius (Ruiz & Pavon) Blume from Argentina. While the systematic position of this species may be questioned, the reliability of the endosperm—less condition as the major generic criterion is equally insecure. Little doubt exists, nevertheless, that species lacking endosperm are to be included in Psittacanthus. The author, therefore, visited the area of Bahia de los Angeles, Baja California in early January, 1969, in order to study fruits in the fresh condition. Detailed results will be published elsewhere at some future date. The main result, however, is a confirmation of the complete absence of endosperm in mature fruits, the cotyledons having taken over the storage function. There is no alterna- tive, therefore, to transfer this mistletoe to Pszttacanthus. As the internal organization of the genus Psittacanthus has not been adequately worked out it is impossible to place P. sonorae accurately in relation to other species.

PSITTACANTHUS sonorae (Watson) Kuijt, comb. nov. Loranthus sonorae Watson, Proc. Amer. Acad. Arts 24:73. 1889. Phrygilanthus sonorae (Watson) Standley, Contr. U. S. Natl. Herb. 20:212. 1919. Dipodophyllum diguetti Van Tiegham, Bull. Soc. Bot. France 42:177. 1895?

Watson designated this species “Loranthus (Psittacanthus ) Sonorae.” While does this not constitute a nomenclaturally valid publication of Psittacanthus sonorae, it leaves no doubt as to Watson’s ideas of affinity.

A noteworthy fact observed in several instances near Bahia de los Angeles was the hyperparasitism of the common desert mistletoe, Phoradendron californicum Nuttall, on Psittacanthus sonorae (fig. 1). In one case more than a dozen hyperparasitic plants of various sizes grew

1s)

14 MADRONO [Vol. 21

Fic. 1. Hyperparasitism of Phoradendron californicum (Pc) cn Psittacanthus sonorae (Ps), in turn parasitic on Bursera microphylla (B). The chalk-white bark belong to Psittacanthus sonorae.

on a single individual of Psittacanthus sonorae. The haustorial organs of the hyperparasite were surrounded by a crater-like formation of Psztta- canthus wood, and were greatly restricted when compared to their normal development on desert trees. A similar phenotypic variability in haustorial development has been reported in other cases of hyperparasitic mistletoes (Kuijt, 1964). This kind of hyperparasitism in mistletoes is undoubtedly an indication of the fact that local fruit-eating birds eat the berries of both species, the seeds thus being voided together. No evidence was seen of self-parasitism of Psittacanthus sonorae on mem- bers of the same species.

I would like to thank Reid Moran and P. D. Warrington for assistance in the matter of transportation, and F. A. Stafleu for nomenclatural advice.

LITERATURE CITED

Bartow, B. A. 1962. Studies in Australian Loranthaceae. I. Nomenclature and new additions. Proc. Linn. Soc. New South Wales. 87:51-61.

BHATNAGAR, S. P., and S. CHANDRA 1968. Endosperm in Psittacanthus. Curr. Sci. 37:704—-706.

Kurt, J. 1964. Critical observations on the parasitism of New World mistletoes Canad. J. Bot. 42:1243-1278.

STANDLEY, P. C. 1919. Studies in tropical American phanerogams—No. 3. Contr. U.S. Natl. Herb. 20:173-220.

THE GENUS CHLOROPHYLLUM (LEPIOTACEAE) IN CALIFORNIA

WALTER J. SUNDBERG Department of Botany, University of California, Davis 95616

Chorophyllum molybdites (Meyer ex Fr.) Massee, the only species in the genus, was first reported from Southern California by Smith (1936), but without an accompanying description. While conducting a regional taxonomic study of Lepiota S. F. Gray and related genera (Sundberg, 1967), I verified the occurrence of C. molybdites in Southern California. Subsequent field work and herbarium studies indicated that its range extended much further north than Los Angeles, the northern limit reported by Smith (1936). Since C. molybdites is poisonous to some individuals (Singer, 1948; 1962; Smith, 1954), and often appears in cultivated areas such as lawns and gardens, it seems important to report on the characteristics and extended geographical distribution of Cali- fornia material.

A detailed morphological and anatomical description is included be- cause few exist in the literature and published descriptions of California specimens are non-existent. Colors in quotation marks are those of Ridgway (1912). Where Ridgway’s terminology may be unfamiliar to those lacking access to his color standard, approximations of his colors are included in general terms. Thirty-four collections were examined and deposited in either the herbaria of San Francisco State College or Uni- versity of California, Berkeley. The distribution map (fig. 2) is based upon personal observations and data from herbarium specimens.

CHLOROPHYLLUM MOLYBDITES (Meyer ex Fr.) Massee, Bull. Misc. Inform. 1898: 136.1898.

Pileus 4.5—-15.5 cm broad, ovoid to obtuse when young and unopened, becoming convex to broadly convex upon opening, finally plano-convex to uplifted in age, disc sometimes distinctly raised following expansion; margin incurved prior to rupture of the partial veil, remaining incurved upon opening, then becoming decurved, in- frequently plane in age, finely lacerate at first, becoming rimulose to rimose, finally splitting deeply, often faintly striate (more obvious in age) ; surface moist but not viscid when young, becoming dry; cuticle continuous in the young button stage, during enlargement remaining continuous on the disc but becoming concentrically to irregularly diffracted and forming flattened to uplifted scales which often rub off easily toward the margin, infrequently splitting radially, the disc becoming diffracted scaly to areolate scaly in age with the scales composed in part of the flesh; exposed flesh (the remainder of the pileus surface in later stages) appearing appressed fibrillose to appressed fibrillose scaly, frequently shaggy fibrillose scaly to minutely squarrose scaly toward the margin (especially when young), the scales easily rubbing off and formed primarily by aggregation of the upper layers of radially arranged flesh fibrils, rarely with small amounts of cuticle at the tips; cuticle color variable in young unopened carpophores, reddish brown (‘“russet” to “rood’s brown” to “liver brown” to “warm sepia”) to brown (“sayal brown” to “bister”), often variegated and spotted with pale to dark buff shades (“light ochraceous-buff” to “antimony yellow” or “pale pinkish buff” to “cinnamon-buff”’),

15

16 MADRONO [Vol. 21

Fic. 1. Chlorophyllum molybdites, X 13, Sundberg 1228.

occasionally spotted dull pinkish vinaceous (“light cinnamon-drab”’) to pinkish brown (“fawn color”) to blackish purple (“dark livid brown” to “warm blackish brown’’); disc and cuticular scales often assuming somewhat different shades of reddish brown (“natal brown” to “vandyke brown” to “argus brown’) or brown (“brussels brown” to “prout’s brown” to “cinnamon-brown” to “tawny-olive’’) to becoming pinkish brown (“wood brown” to rarely “avellaneous’”’) with age; surface flesh white at first, rarely spotted pale pinkish (“pale cinnamon-pink’’) to dark pinkish vinaceous (‘brownish vinaceous’’) in older but unopened buttons, some- times with pinkish vinaceous (“light grayish vinaceous” to “light russet-vinaceous” or “light cinnamon-drab” to “cinnamon-drab’’) tinges near the margin, becoming brownish pink (“vinaceous-buff’”?) to pinkish brown (“wood brown’’) to light pinkish gray (near “light drab”) in age, usually unchanging or darkening slightly on bruising, rarely staining “cinnamon” at first, then pale reddish brown (“cacao brown’’) and finally becoming brown (‘“‘verona brown”) ; tips of scales composed of flesh becoming brown (dark “wood brown” to “snuff brown” to “saccardo’s um- ber’) in age. Flesh (2—) 5-13 mm thick at the disc, soft but solid; white to off- white (“‘tilleul buff’’) to tinged pinkish gray (“light drab’), cream to buff (‘‘cream- buff” to “pinkish buff” to “cinnamon-buff”) near the stipe apex and lamellae, stain- ing pinkish (“buff-pink’”’) to “light pinkish cinnamon” at first, then darkening to “cinnamon” to “orange-cinnamon,” and finally becoming reddish brown (“vina- ceous-russet” to “rood’s brown” to “verona brown’) when bruised, infrequently staining pale orange (‘‘salmon color” to “orange-buff”) or brown (“sayal brown” to “snuff brown”) at first, then becoming reddish brown. Taste mild. Odor not distinctive.

Lamellae free, remote from the stipe even when young, sometimes forking and more rarely anastomosing near the stipe apex; close; 6-18 mm broad; fragile; thin at the margin, but moderately thick near the pileus flesh; white in mass when young, often even when first expanded, becoming pale green (“‘pale glaucous green” to “yellowish glaucous” to “water green’) to gray-green (“court gray” to “tea green” to “celadine green’’?) when mature, tinged near gold to olive-gold (“honey yellow” to “isabella color’) where beginning to dry out naturally; margin entire and finely white fimbriate at first, becoming irregularly discontinuous and dark brown to almost black in age.Lamellulae in two to three tiers.

1971] SUNDBERG: CHLOROPHYLLUM 17

C

Fic. 2. Distribution of Chlorophyllum molybdites in California.

Stipe 5.7-12.0 cm long, 6-15 mm broad at the apex, equal to slightly enlarged below, rarely clavate; often but not always easily separable from the pileus; surface dry, silky to innately fibrillose throughout at first, sometimes appearing peronate- scaly above, areolate-splitting and shiny below in age, clothed at the base with a tightly appressed white mycelial growth; white to off-white (“tilleul buff”) to rarely pale buff (“pinkish buff”) above the annulus, sometimes streaked to tinted with pinkish vinaceous (“pale vinaceous-fawn” to “light russet vinaceous”) to dark pinkish brown (“fawn color” to “army brown”) to rarely dark pinkish gray (“benzo brown’’) shades, often superficially green (“water green” to “gnaphalium green”) from spore deposits at maturity, staining buff (‘“‘cinnamon-buff”), then darkening to “cinnamon” to “vinaceous-cinnamon” when bruised; below the an- nulus white to near brownish pink (‘“avellaneous”) and sometimes spotted with pinkish vinaceous (“pale brownish vinaceous’’) or purple-gray (“light brownish drab’) at first, becoming pale to dark pinkish brown (“wood brown” to “fawn color” to “natal brown’’) to reddish brown (“cinnamon-brown” to “rood’s brown” to “auburn” to “warm sepia’’) in age, whitish fibrils forming an overtone giving the darker colors a streaked appearance; stuffed, becoming hollow; pith fibrils white to off-white (“tilleul buff’) ; cortex “tilleul buff” to infrequently brownish pink (“vin- aceous-buff” to “avellaneous’’), sometimes concolorous with the surface where ad- jacent to it, staining pale buff (“pinkish buff’), then buff (“cinnamon-buff” to “pinkish cinnamon’’) to “cinnamon,” then pale orange (‘‘orange-buff”) to “orange- cinnamon,” finally becoming pale to dark reddish brown (‘“‘vinaceous-tawny” to “russet” to “walnut brown” to “vandyke brown’’) where bruised.

Annulus superior to infrequently median, attached and sleeved above, flaring below, typically immovable when fresh, often becoming free and movable on drying, thick and solid, complex and having three frequently indistinct flanges;

18 MADRONO [Vol. 21

Fic. 3. Clamp connections from the pileus trama of Chlorophyllum molybdites: a, Sundberg 1251; b, c, Sundberg 1229. All approximately x 2,100.

upper surface white and often spotted with pinkish vinaceous “pale vinaceous- fawn” to “light grayish vinaceous”) to pinkish gray (“ecru drab’) shades at first, becoming pinkish brown (‘“avellaneous” to “wood brown”) in age, sometimes tinged green (“water green” to “gnaphalium green’) from the spore deposit, undersurface brown (“snuff brown” to “prout’s brown”) to reddish brown (warm sepia’’) near the margin and white with infrequent pinkish vinaceous tinges elsewhere.

Spore deposit pale green (“pale glaucous green” to “deep lichen green” to “water green” to “corydalis green’) to gray-green (‘mineral gray” to “vetiver green” to “onaphalium green” to “sage green” to “celadine green”) to almost gray (“storm gray’) when fresh, occasionally as light as “cream-buff,” becoming slightly darker green (“tea green” to “andover green” to “lincoln green”) on drying, fading in age to a shade of dark gold to brownish gold (“yellow ocher” to “mars yellow”).

Spores (7.7—) 10.3-12.2 (-13.5) X& (6.3—-) 7.7-9.0 (-10.9) w, ovoid to very short ellipsoid, apex truncate, infrequently at a slight angle, inequilateral in side view, smooth, hilar appendix present, apical pore present giving some a notched appear- ance, wall thick, aguttulate or uniguttulate, entire spore dark blue in cresyl blue, pale yellowish green to pale yellowish brown in KOH when viewed singly (color most obvious in wall), weakly to strongly dextrinoid (pale yellowish orange to reddish orange to reddish brown) when viewed singly in Melzer’s reagent.

Basidia (24.2—) 29.4-44.8 (-61.5) X& 9.0-12.9 (-14.1) mw, mostly 4-spored, occa- sionally 3- or 2- or 1-spored but not notably different in size, clavate, some with narrow elongate bases, often granulose, hyaline to rarely tinged pale yellow to pale yellowish brown in KOH, yellowish to pale to deep yellowish orange in mass and hyaline to pale yellowish when viewed singly in Melzer’s reagent.

Cheilocystidia (18.0—) 21.9-47.4 (-62.4) & (9.5-) 10.3-25.5 wu, abundant, some- times scattered in fascicles along the lamellae margin, arising as hyphal tips or branches, narrowly to broadly clavate to sphaeropedunculate to subsaccate, infre- quently with angular to rostrate apices, rarely covered with an amorphous material, bases often narrow and elongate, walls thin to unevenly thickened and yellowish when viewed singly in KOH, hyaline to yellowish brown in mass and hyaline to pale yellowish to pale yellowish brown when viewed singly in KOH, concolorous in Melzer’s reagent. Pleurocystidia absent.

Lamellae trama composed of loosely interwoven hyphae, hyaline in KOH, hyaline to pale yellowish in Melzer’s reagent; oleiferous hyphae often present; subhymenium cellular, segments small, compact, concolorous with the trama proper.

Pileus trama composed of tightly or more often loosely interwoven hyphae, seg- ments 5.1-15.4 mw wide, hypodermal segments usually shorter and broader than cuticular elements, sometimes swollen and enlarged, rarely almost globose, entire trama hyaline to infrequently yellowish to yellowish brown in KOH, pale yellow to pale yellowish orange in mass and hyaline to pale yellowish to pale yellowish orange

1971] SUNDBERG: CHLOROPHYLLUM 19

to rarely pale yellowish brown when viewed singly in Melzer’s reagent (segments of the same hyphal strand may differ in color) ; oleiferous hyphae present, hyaline to pale yellowish to rarely pale yellowish green in KOH.

Cuticle scattered in patches due to diffraction caused by pileus expansion, com- posed of tightly and irregularly arranged more or less upright hypae and hyphal tips, appearing somewhat turf-like, longer hyphae or at least the apical portions frequently repent and forming a thin, often loosely arranged, appressed and inter- woven layer above the upright elements; terminal segments usually not well dif- ferentiated, length extremely variable, (18—-) 20.6-86.2 (-—132.4) X& (2.6—) 3.8-9.0 (—15.8) mw, often arising as hyphal branches, rarely ventricose to slightly enlarged above, very seldom clavate, apices usually rounded, infrequently tapered to mucro- nate or rostrate at the apex, walls irregularly flexuous, thin to unevenly thickened, especially in the upright segments, rarely encrusted, some appearing striated in KOH, infrequently with a fine sparse granular content; subterminal segments often with thickened walls; hyaline where thin to yellowish brown to rarely reddish brown where thick in mass and hyaline to pale yellowish to pale yellowish brown when viewed singly in KOH, pale yellowish orange to orange-brown to pale reddish brown in mass and hyaline to pale yellowish to pale to dark yellowish brown when viewed singly in Melzer’s reagent.

Clamp connecticns always present in the pileus trama of both young and old specimens, often rare and never at every septum, seen at the base of some cheilo- cystidia in one collection, apparently jacking elsewhere.

A pale yellowish to pale yellowish brown pigment, which is soluble in water but not alcohol, diffuses from the tissue during sectioning preparation.

Habit and habitat: Scattered to gregarious in laws or more rarely in garden soils, usually arranged in fairy rings. July through September.

Specimens examined. Fresno Co.: Sundberg, 1217, 1220, 1222 1246- 1248, Thiers 20742. Kern Co.: Sundberg 1236, 1237, Thiers 20740. Kings Co.: Sundberg 1230-1232, 1234. Los Angeles Co.: Floyd s. n., Sloan, in 1966, 1967, Brubaker s. n. Riverside Co.: Smith s. n. San Diego Co.: McLean s.n., Miller s. n. Santa Barbara Co.: Walker s. n. Tulare Co.: Sundberg 1225, 1227-1229, 1243-1245, 1249-1251, Thiers 20741.

DISCUSSION

Chlorophyllum molybdites differs from closely related taxa by its colord spores. It appears most similar to Lepiota rachodes (Vitt.) Quel., but lacks the palisade cuticle of pyriform cells foud in the latter. Its summer and early fall fruiting habit is also distinctive.

As indicated in Fig. 2, C. molybdites is apparently restricted to the southern part of California. It may possibly occur further north than present data indicates, especially in the Central Valley, since it has been reported from much more northerly latitudes elsewhere (Groves, 1962; Singer, 1948; 1962; Smith, 1954).

California material studied in the fresh condition appears to be C. molybdites var. marginata A. H. Smith, but the lamellae margins are white in the button stage rather than dark as indicated in the original description of the variety (Smith, 1949). However, the margins do darken to almost black as the carpophores mature.

The presence of clamp connections (fig. 3), also recently noted in

20 MADRONO [Vol. 21

Arizona collections by C. Leathers (personal communication to H. D. Thiers), is of interest since Singer (1948; 1962), Smith (1949), and Smith (1954) all reported their absence. Preliminary anatomical exam- ination of collections from other parts of the United States (Sundberg and H. D. Thiers, in preparation) has verified the presence of clamp connections in this species. This evidence indicates that they were prob- ably overlooked by previous workers and their absence cannot be used, as had been by Singer (1948; 1962), in support of the generic segrega- tion of Chlorophyllum from closely allied species of Lepiota sensu lato.

I wish to thank H. D. Thiers for guidance leading to and during this study and for his critical comments on the manuscript. The criticisms of K. Wells and G. Breckon were most appreciated. Financial support for field work was provided by the Department of Botany, University of California, Davis.

Note added in proof. Recently clamp connections were reported in C. molybdites from Tennessee and Israel (Heinemann, P. Bull. Jard. Bot. Etat 38:195—206. 1968) and from Africa and South America (Singer, R. Beih. Nova Hedwigia 29: 1—405. 1969).

LITERATURE CITED Groves, J. W. 1962. Edible and poisonous mushrooms of Canada. Queen’s Printer, Ottawa. Ripcway, R. 1912. Color standards and color nomenclature. Washington, D. C. SINGER, R. 1948. New and interesting species of basidiomycetes. II. Pap. Michigan Acad. Sci. 32:103-150. . 1962. The Agaricales in modern taxonomy. 2nd ed. J. Cramer, Weinheim. Smirn, A. H. 1949. Mushrooms in their natural habitats. Sawyers, Portland. Smitu, C. O. 1936. Lepiota morgani in Southern California. Mycologia 28:86. SmitH, H. 1954. A revision of the Michigan species of Lepiota. Lloydia 17:307-328. SUNDBERG, W. J. 1967. The family Lepiotaceae in California. Master’s thesis. San Francisco State College.

POLYPHYLETIC ORIGIN OF TETRAPLOID POPULATIONS OF GUTIERREZIA SAROTHRAE (COMPOSITAE)

Otto T. SOLBRIG Gray Herbarium, Harvard University, Cambridge, Massachusetts 02138

The widespread western North American species Gutierrezia sarothrae (Compositae—Astereae) possesses two chromosomal races: a diploid (n = 4) race that is the most common, and a less common tetraploid (n = 8) race, that comprises approximately 20% of populations. Both races are morphologically very similar. Although the polyploids tend to have statsistically slightly larger pollen and stomata, there is no single character by which the two races can be separated (Solbrig, 1964). This is due to a great deal of variability in both the diploid and tetra- ploid forms. Neither can the two races be separated on the basis of geographical distribution since the tetraploid populations occur inter- spersed throughout the distribution of the species.

1971] SOLBRIG: GUTIERREZIA wi

In a previous paper (Solbrig, 1964) I suggested that polploids may have arisen from diploids more than once. This would tend to explain their morphological diversity and geographical distribution. The major reason for making this suggestion, was that at any one locality where polyploids occurred, they tend to show the expected “gigas” characters in relation to the diploids of that area. However, the diploid populations of another region are not necessarily smaller. In the present paper fur- ther evidence in favor of the hypothesis of a polyphyletic origin of the polyploid populations is presented as a result of a numerical analysis of relationships among the populations.

MATERIALS AND METHODS

The data collected in previous studies (Solbrig, 1960; 1964; 1965) were utilized in the present analysis. The OTU is the breeding popula- tion. The characters used are the mean value obtained after measuring 13 characters in 50 individual plants chosen at random in a population. For this study data from 15 diploid (n = 4), 5 tetraploid (n = 8) and 14 populations of unknown chromosome number of G. sarothrae, as well as 3 tetraploid (n = 8), 1 hexaploid (n = 12) and 1 of unknown chromosome number of G. bracteata; 1 diploid (n = 4) population of G. serotina and 1 hexaploid (n = 12) population of G. serotina and 1 hexaploid (n = 12) population of G. californica; and 2 tetraploid popu- lations of G. microphala, were utilized (table 1). The data were pro- cessed on an IBM 7090 Computer at the University of Michigan Com- puting Center.

Two numerical analyses were used. One, the Prim Network (Prim, 1957), is essentially a coefficient of similarity in a multidimensional space that produces a linear clustering of OTU’s. The second is a pro- gram that produces a phylogenetic tree following the principle of parsi- mony, where species are derived from a given ancestor along the most parsimonious paths (Kluge and Farris, 1969; Wagner, 1961).

RESULTS

Figure 1 depicts graphically the Prim Network that was obtained. It can be seen that the variability of the species follows three major lines. First, a line that is characterized by a tendency towards smaller heads with a reduced number of florets per head and that culminates in populations of G. microcephala that have only 1 ligulate and 1 tubular florets. Of the 16 populations in this line, 6 are diploid, 3 are tetraploid (including both populations of G. microcephala) and seven are of un- known chromosome number. Ten of the populations come from the southeast area of the distribution of these species, that is, the area comprised by Colorado, New Mexico, Oklahoma and Texas. The other six populations come from Utah, Wyoming, Kansas and South Dakota.

The second direction exhibited by the pattern of variation is toward an increase in head size with accompanying increase in the number of florets. Only three populations of G. sarothrae are involved, together

22 MADRONO

3256

3150 2769 2801 2765 3215 3286 3246 3250 3243 3280 3248 3264 3231 2805 3224 3255 3254 3258 3151 2758 oF SEROTINA

3265

[Vol. 21

2 MICROCEPHALA 32

3269 327 3238 3262 3292 329]

3284

3273

3263 3259 3252

3266 BRACTEATA

CALIFORNICA

Fic. 1. Prim network of the populations studied. The lines separating any two populations represent the phenetic distances. The angles are arbitrary, and distances between populations other than along the drawn lines non-significant. All popula- tions that are not shaded are Gutierrezia sarothrae, those encircled in a balloon are tetraploid with n = 8; all other G. sarothrae are either diploids or of unknown

chromosome number (see table 1).

1971] SOLBRIG: GUTIERREZIA Us)

TABLE 1. LOCALITIES SAMPLED FOR THIS STUDY AND CHROMOSOME NUMBERS. Collection numbers are those of the author and specimens are in the Gray Herbarium.

Gutierrezia bracteata. California: Cache Creek, 3432; Arroyo del Puerto, 3433, n = 8; Arroyo del Puerto, 2743, n = 8; Pond Ranch, 2748; Temblor Range, 2755.

Gutierrezia californica. California: Point Bonita, 3431,n = 12.

Gutierrezia microcephala. Arizona: Hyde Park, 2804; New Mexico: White City, 3272.

Gutierrezia sarothrae. Arizona: Payson, 2794, n = 8; Ash Fork, 2801, n = 4; Hyde Park, 2805, n = 4. California: near Temecula, 2768, n = 4; near Chula Vista; 2776, n = 4; near Rancho Santa Fe, 2769, n = 4. Colorado: Colorado National Monument, 3261, n = 8; Colorado National Monument, 3262, n = 4; near Salida, 3263; near Fountain, 3264; near Trinidad, 3265. Idaho: near Dubois, 3254,n = 4; near Pocatello, 3255. Kansas: near Medicine Lodge, 3292, n = 4. Montana: Glen- dive, 3248; Sanders, 3250, n = 4. Nebraska: Roscoe, 3224, n = 4; near Chadron, 3231. New Mexico: near jct. highways 285 and 286, 3169, n = 8; near Glenwood, 3215; near Las Vegas, 3266; near Roswell, 3269, n = 4. North Dakota: Dickinson, 3243; Little Missouri R. and highway 85, 3246. Oklahoma: near Bouse Jct., 3291. South Dakota: near Rapid City, 3238. Texas: near Van Horn, 3212, n = 8; near White City, 3271, n = 4; near Van Horn, 3273, n = 8; near Alpine, 3280, n = 4; near San Angelo, 3284; near Abilene State Park, 3286, n = 4. Utah: near Boulder, 3150, n = 4; near jct. highways 24 and 6, 3151, n = 4; near Ogden, 3256; near Vernal, 3258,n = 4; near Talmadge, 3259.

Gutierrezia serotina. Arizona: Tucson, 2777,n = 4.

with all the populations of G. bracteata, G. californica and G. serotina that were analyzed. Two of the three populations of G. sarothrae that exhibit this pattern of variation are tetraploid; in addition all popula- tions of G. bracteata and G. californica are either tetraploid or hexaploid, so that of 10 populations involved, 2 are hexaploid, 5 are tetraploid, 2 are diploid and one is of unknown chromosome number. All but one of the populations grow in either Arizona or California. The exception is population 3261, a tetraploid G. sarothrae from Grand Junction, Colo- rado.

The remaining 21 populations, all of them G. sarothrae, comprise what may be called “typical” sarothrae. Geographically they come from throughout the range of the species, including California and Texas. Chromosomally 8 populations are diploid, 2 tetraploid and eleven are of unknown ploidy level.

Figure 2 depicts the phylogeny produced by the Farris program. For purposes of this analysis, population 3256 of G. sarothrae which exhibits characters at the center of the species variability in the Prim Network was chosen arbitrarily as exhibiting the most primitive characters. Runs were made using other populations as primitive. This produced different shaped trees, but no variation in the basic relations of the populations to each other. According to the phylogenetic tree produced by this pro- gram, there are three evolutionary tendencies within G. sarothrae that follow roughly the lines of variation uncovered by the PRIM diagram: one is a tendency towards smaller heads with fewer flowers that cul- minates in G. microcephala,; the other is a tendency towards larger heads that culminates in G. bracteata and G. californica, and the third is also

24 MADRONO [Vol. 21

CALIFORNICA N=12

3431

SEROTINA N=4

2777 MICROCEPHALA BRACTEATA a N=812 16 : 2804 ar:

Fic. 2. Hypothetical phylogeny following the most parsimonious lines drawn by a computer following a program devised by J. A. Farris (Kluge, 1969). Non-shaded populations are G. sarothrae; those included in a balloon are tetraploids with n = 8; all other G. sarothrae are either diploid or of unknown chromosome number (see table 1).

a tendency towards larger plants with larger heads not connected to the previous one. Gutierrezia californica appears to be the most modified of the species analyzed; the line that culminates in G. microcephala follows. These numerical indices tend to confirm the taxonomic treatment with one exception: population 2794 from Arizona appears to be related both in its pattern of variation and in its hypothetical phylogenetic history to G. bracteata, and probably should be classified as such. Blake in the Flora of Arizona (Kearney and Peebles, 1942) classified the large headed Arizona populations as G. californica (= G. bracteata). This would extend the range of this species, previously considered by me (Solbrig,

1971] SOLBRIG: GUTIERREZIA 25

1965) as restricted to California and Baja California. Of greater interest for our present discussion, is that polyploid populations are found in all three of the phylogenetic lines.

DISCUSSION

The present numerical analysis tends to confirm the hypothesis that tetraploid populations of G. sarothrae have arisen and become estab- lished in more than one instance. This would explain the geographical dispersion of the tetraploids, and also the fact that they cannot be sep- arated as a group from diploids also taken as a whole. However, when tetraploid populations are compared to the diploid populations to which they are most closely related within each of the three major lines of variability, they show the expected “gigas” characteristics (fig. 1). This analysis permits us to obtain a reasonable idea of the evolutionary patterns within a species. From the “typical” or “modal” population it appears that a line with larger heads and more flowers developed in the southwest United States that eventually lead to the formation of three species: G. serotina, G. bracteata and G. californica. On the other hand, in the southeast part of the range, forms with narrower heads and fewer flowers were selected that lead to the formation of G. microcephala, a species that eventually became widespread over all of southern United States and northern Mexico. It is impossible to affirm that polyploidy provided the isolating barrier in these instances and aided the selective trends, but it appears to be a plausible hypothesis.

I wish to thank J. S. Farris and A. K. Kluge for assistance and for the loan of the program decks used.

LITERATURE CITED

Kearney, T. H., and R. H. PEEBLEs. 1942. Flowering plants and ferns of Arizona. U.S. Gvt. Printing Office, Washington. KiuceE, A. K. and J. S. Farris. 1969. Quantitative phyletics and the evolution of Anurans. Syst. Zool. 18:1-32. Prim, R. C., 1957. Shortest connection networks and some generalizations. Bell System Tech. J. 36:1389-1401. Sorpric, O. T. 1960. Cytotaxonomic and evolutionary studies in the North American species of Gutierrezia (Compositae). Contr. Gray Herb. 188:1-63. . 1964. Infra-specific variation in the Gutierrezia sarothrae complex (Com- positae-Astereae). Contr. Gray Herb. 193:67-115. 1965. The California species of Gutierrezia (Compositae-Astereae). Madrofno 18:75-84. Wacner, W. H. 1961. Problems in the classification of ferns. In Recent Advances in Botany 1:841-844.

XYLEM MONOTERPENES OF PINUS PONDEROSA, P. WASHOENSIS, AND P. JEFFREYI IN THE WARNER MOUNTAINS OF CALIFORNIA

RICHARD H. SMITH

Pacific Southwest Forest and Range Experiment Station U.S. Department of Agriculture, Berkeley, California 94701

In the Warner Mountains of northeastern California, the subsection Ponderosae of the genus Pinus is represented by three species: ponderosa pine (Pinus ponderosa Laws.), Washoe pine (P. washoensis Mason & Stockwell), and Jeffrey pine (P. jeffrey Grev. & Balf.). Washoe was initially reported to be confined to a small stand at Patterson Meadow at the Warners’ southern end (Haller, 1961), but it will probably be greatly enlarged by more recent studies. Ponderosa is found quite ex- tensively throughout the area. Jeffrey grows considerably less extensively than ponderosa but much more extensively than Washoe, though limited to the southern end of the Warners (Critchfield and Little, 1966). Bo- tanically, the area is of interest because it lies on the northeastern fringe of the range of Jeffrey pine and contains one of the isolated stands of Washoe.

The monoterpene composition of the xylem resin of ponderosa is ex- tremely variable, while that of the other pines is much less variable and can be somewhat stereotyped (Mirov, 1961; Smith, 1964; 1967). The composition of an average ponderosa pine from low eievation in the Warners is about 7% a-pinene, 25% £-pinene, 46% 3-carene, 10% myrcene, 9% limonene, 1% -phellandrene, and 2% terpinolene. A normalized composition for average Washoe is 5% a-pinene, 10% B- pinene, 65% 3-carene, 15% myrcene, 1% limonene, 1% £-phellandrene, and 3% terpinolene and some trace components; Jeffrey pine is about 95% heptane, 2% nonane, and the remaining 3% divided among several of the monoterpenes.

Individual ponderosa pines have been found whose monoterpene composition falls within the average composition of Washoe. A few such trees have been found in low-elevation stands in the northern Warners and in stands in the central and northern Rocky Mountains. These ponderosa stands in the eastern portion of its range often are classified as P. ponderosa var. scopulorum Englem.

In 1965-66 a study was made of the monoterpene composition of the xylem resin of ponderosa or Washoe or both in three areas of the Warner Mountains (fig. 1) to determine a, range of stands which could be classified as Washoe on the basis of monoterpene composition, b, how often individual trees in ponderosa stands showed the full characteris- tics of Washoe monoterpenes, and c, the possibility of hybridization among the three species. Jeffrey pine was studied in the southern area where it grows with ponderosa or Washoe or both.

26

1971] SMITH: MONTERPENES Dd

Cedarville

N fe) jo) oO

€ Fork Py Likely {NS eS

orporation Mdw.

x jo) jo) fe)

Y) a g ue

Contour Interval 10 MILES 1000 feet

Fic. 1. Location of study plots (0) and their elevation in three areas near Al- turas, Modoc Co., Calif.

28 MADRONO [Vol. 21

TABLE 1. INDIVIDUAL TREES SELECTED FRoM Eacu Pitot To SHOW CHARACTERISTIC Mayor MoNoTERPENE COMPOSITION OF WASHOE PINE IN PERCENT

Plot No. a-pinene B-pinene 3-carene Myrcene Limonene 1 7 1 69 17 Trace 6 1 65 22 1 2 6 20 60 8 Trace 6 14 62 12 1 3 7 4 69 13 1 15 14 60 6 Trace 4 5 16 63 11 1 5 4 1 79 10 Trace 10 2 74 8 Trace 16 1 68 10 Trace 6 8 4 66 15 Trace $ 6 71 15 Trace 7 4 69 12 2 Hi 4 1 78 12 1 6 3 69 15 Trace 6 4 65 19 Trace 8 5 11 64 14 Trace 6 8 65 16 Trace 6 12 60 1 1 9 5 4 73 12 1 5 2 78 9 — 9 1 65 19 Trace

Two to four plots were selected at different elevations in the three areas. The elevation at each plot is as follows: plot 1, 5,500 ft.; plot 2, 5,700 ft.; plot 3, 5,000 ft.; plot 4, 6,000 ft.; plot 5, 5,000 ft.; plot 6, 5,800 ft.; plot 7, 7,000 ft.; plot 8, 6,800 ft.; and plot 9, 7,200 ft. About 20 trees per plot were tapped for resin at plots 1 to 8, inclusive. Except for the Jeffrey pines, no effort was made to describe the trees morpho- logically; they were selected simply on the basis of geography. Previous data gathered for the Patterson Meadow location of Washoe were in- cluded for a reference as the ninth plot.

I used previously described procedures for tapping the trees, pre- paring the samples for analysis, and analyzing the samples by gas chromatography (Smith, 1964). All results of analysis are based on the normalization of the monoterpene fraction of the xylem resin; i.e., each monoterpene is expressed as a percent of the total monoterpenes.

RESULTS

The frequency distribution of each of the five major components— a-pinene, 8-pinene, 3-carene, myrcene, and limonene—in each of the nine plots is given in figure 2. Trees with the typical composition of Washoe pine were found in each location (table 1). The frequency of this type of tree seemed to shift markedly between 6,000 and 7,000 feet; plots below this zone (plots 1-6) can be classified as typical ponderosa stands, containing a scattering of trees which have monoterpene char-

1971] SMITH: MONOTERPENES 29

a-pinene B-pinene 3-carene myrcene limonene Flot le}

Frequency ro) OS s) fo) : an

| 20 0 10 20 30 40 20 30 40 50 60 70 800 Percent

Fic. 2. Frequency distribution of the five major monoterpenes of Washoe and ponderosa pine from nine plots in the Warner Mountains, northeastern California.

acteristics of Washoe, comprising generally about 10% of the stand. Trees at higher elevation are typical Washoe; in these plots (7-9) an infrequent tree has the monoterpene characteristics of ponderosa pine, comprising generally about 10% of the stand. In addition to trees in low-elevation stands with all the monoterpene characteristics of Washoe, individual trees may be found which show at least one of the character- istics of Washoe; i.e., either > 60% 3-carene, << 20% #-pinene, or < 2% limonene. The occurence of these two types of trees suggests the possibility of both geographical and physiological mixing of ponderosa and Washoe.

This study also suggested that the previously defined range of Washoe in this area should be enlarged (Haller, 1961). If a criterion of > 60% 3-carene, < 20% f-pinene, and < 2% limonene is used for Washoe pine, 10% of the trees below the 6,000- to 7,000-foot zone fit this criterion and might be called Washoe; 38% of the trees above this zone do not fit this criterion and might be called either ponderosa or hybrids between ponderosa and Washoe. There is some evidence that plot 6, at about 5,800 feet, is near the transition zone, since the frequency dis- tribution of limonene for the trees in this plot (fig. 1) is typical of Washoe, while the 3-carene and #-pinene distributions are typical of ponderosa.

Limonene was of particular interest in this study. Except in a few instances it is lacking in trees from the typical Washoe plots; in all plots considered typically ponderosa in the Warners or the Sierra Nevada it may be absent or it may occur in an amount of from about 5 to 10%.

30 MADRONO [Vol. 21

limonene

o-carene

Frequency

a aN Lj} A O° |O=-20- 30 +40. 50 60° 7O “BO! 90) GO" WO. 20, 350 340 50 Percent Percent Fic. 3. Frequency distribution of #-pinene, limonene, and 3-carene from four sources: A, Washoe pine, B, low-elevation ponderosa in Warner Mts., C, ponderosa from central Sierra Nevada, D, var. scopulorum of ponderosa from Colorado. Wyoming, and Nebraska.

Only a few trees had between 1 and 5%.

The Jeffrey pines growing at plot 6 were found to be very similar to those in the Sierra Nevada, but the group included one tree that had a monoterpene composition highly suggestive of that of a hybrid between Jeffrey and Washoe. Its composition was 17% heptane, 2% nonane, 2% a-pinene, a trace of a-thujene, 4% -pinene, 55% 3-carene, 6% myrcene, 11% limonene, and 3% terpinolene. The amounts of heptane and nonane are clear signs of Jeffrey pine. The low £-pinene, high 3- carene, and trace of a-thujene are the basis for believing the other parent is Washoe. However, the 11% limonene suggests that ponderosa could be the other parent. There is always the rather remote possibility of a three-way hybrid.

Hybrids have been artificially produced between Jeffrey and both ponderosa and Washoe (Liddicoet and Righter, 1960) but not without

1971] SMITH: MONOTERPENES 31

TABLE 2. DIFFERENCES AMONG MEANS OF LIMONENE AND t-VALUES OF Four SOURCES OF PONDEROSA OR WASHOE PINE

Comparison Xi - Xz df. t Washoe vs. Warner, low elevation Dee Ze 5.10** Washoe vs. Sierra Nevada 16.1 290 i026 Washoe vs. var. scopulorum 2.8 158 2,015" Warner, low elevation vs. Sierra Nevada 10.9 388 4.52** Warner, low elevation vs. var. scopulorum 2.4 258 2.88** Sierra Nevada vs. var. scopulorum 1233 274 5.20n%

*— 95, and ** = 99% level of confidence for rejecting the null hypothesis.

difficulty (Critchfield and Little, 1966). Natural hybrids between Jef- frey and ponderosa have been found (Mirov, 1929), but natural hybrids between Jeffrey and Washoe have not been reported. The Jeffrey pines in the Warners are growing in a stand which chemically might be called ponderosa with a scattering of Washoe.

A frequency distribution of limonene, 8-pinene, and 3-carene was made of four sources of ponderosa or Washoe pine (fig. 3): 1, typical Washoe sources (plots 7—9 plus data from Smith (1967)); 2, ponderosa at low elevations in the Warner Mountains (plots 1—6 plus previous data gathered near plot 1); 3, ponderosa from the central Sierra Nevada (Smith, 1966); and 4, ponderosa from Wyoming, Colorado, and Ne- braska, where it is usually called var. scopulorum (Peloquin, 1964). This last grouping is a collection of trees from widely separate locations; but they do represent the whole region fairly well.

A null hypothesis was established that there was no difference in the limonene percentage among the various sources; a t-test of arc-sin transformed percentages shows that this null hypothesis can be rejected at the 95% or greater level of confidence in all comparisons among the four sources (table 2). However, from a visual inspection of the fre- quency distribution, it does appear that low-elevation ponderosa in the Warners is somewhat intermediate between central Sierra Nevada pon- derosa and var. scopulorum but may be more closely related to var. scopulorum.

CONCLUSIONS

This study of xylem monoterpenes suggests: 1, that the range of Washoe pine in the southern Warner Mountains should be enlarged; 2, that some ponderosa stands have geographically mixed with Washoe stands; 3, that composition of the monoterpenes of the two species may overlap considerably; 4, that the two species hybridize naturally; 5, that Washoe is closely related to ponderosa, particularly to var. scopulorum; and 6, that low-elevation ponderosa in the Warners is intermediate be- tween Washoe pine and ponderosa pine of the central Sierra Nevada of California.

32 MADRONO [Vol. 21

LITERATURE CITED

CRITCHFIELD, W. B., and E. L. Lirtre,, Jr. 1966. Geographic distribution of the pines of the world. U.S. D. A. Misc. Publ. 991.

Hatier, J. R. 1961. Some recent observations on ponderosa, Jeffrey and Washoe pine in northeastern California. Madrono 16:126-132.

LippicorT, A. R., and F. I. RiGHTER. 1960. Trees of the Eddy Arboretum. US. Forest Serv., Pacific SW Forest Range Exp. Sta. Misc. Pap. 43.

Mirov, N. T. 1929. Chemical analysis of the oleoresins as a means of distinguishing Jeffrey pine and western yellow pine. J. Forest. (Washington) 27:176-187.

. 1961. Composition of gum turpentines of pines. Techn. Bull. U.S.D.A. 1239.

PELOGUIN, R. L. 1964. Geographic variation of the monoterpenes of Pinus ponderosa. Master’s thesis, Stanford Univ.

SmitTH, R. H. 1964. Variations in the monoterpenes of Pinus ponderosa Laws. Science 143 :1337-1338.

. 1966. The monoterpene composition of Pinus ponderosa xylem resin and

of Dendroctonus brevicomis pitch tubes. Forest Sci. 12:63-68. . 1967. Variations in the monoterpene composition of the wood resin of

Jeffrey, Washoe, Coulter, and Lodgepole pines. Forest Sci. 13:246-252.

STUDIEDS IN THE RHODOPHYLLOID FUNGI. I, GENERIC CONCEPTS

Davip L. LARGENT and ROBERT G. BENEDICT

Department of Biology, Humboldt State College, Arcata, California 95521 College of Pharmacy, University of Washington, Seattle 98105

This paper is concerned with the Rhodophyllaceae in the sense of Singer (except Clitopilus and Rhodocybe).

In 1821 Elias Fries classified the rhodophylloid fungi into tribes which were distinguished by variations of the following features: 1, consistency of the carpophore, particularly of the stipe, 2, attachment of the lamellae, 3, shape of the pileus, and 4, nature of the pileal surface. In 1838 he further emphasized the nature of the pileal surface as a diagnostic feature, using it as well as other characteristics to divide the tribe Entoloma into three sections, and admitted species with flocculose pilei to tribe Nolanea, which previously had contained only mushrooms with glabrous pilei. In subsequent publications Fries no longer used the pileal surface in diagnostic characterizations but defined his tribes (which he now called subgenera) only on the basis of consistency of stipe, type of pileal margin, and attachment of lamellae. These three features have continued to be used by mycologists who have chosen to maintain the Friesian groupings, whether at the generic or subgeneric level. Unfortunately all three are variable, or hard to assess, or both. The difficulty of accurately defining taxa by such features has led some

1971] LARGENT & BENEDICT: FUNGI 33

contemporary mycologists to place all rhodophylloid fungi in a single genus, Rhodophyllus Quel., or Entoloma (Fr.) Kummer emend. Donk, characterized by pink, angular spores, subparallel lamellar trama, and attached lamellae.

This solution does indeed create a clearly defined genus for the rho- dophylloid fungi, but it avoids the questions of what characteristics would be more satisfactory than those used by Fries to delineate sub- generic taxa. In recent years some efforts have been made to find such characters. Romagnesi (in Kithner and Romagnesi, 1953) and Hesler (1967) have emphasized microscopic characters more than macroscopic ones. Smith and Shaffer (1964) have suggested using diameter rather than consistency of the stipe; they define a fleshy-fibrous stipe as one with a diameter at the apex greater than 5 mm, and a cartilaginous stipe as one with the apex less than 5 mm broad. The probable value of urea as a chemotaxonomic aid in separating certain groups of the rhodo- phylloid fungi was shown by Tyler, et al. (1965).

The present study proposes the use of the anatomy and the general aspect of the pileal surface as important diagnostic characters, and correlates concentration of urea with various microscopic and macro- scopic features.

AFFINITIES AND CHARACTERIZATION

The rhodophylloid fungi have pink to vinaceous spores that are angu- lar in face, side, and end views, lamellar trama composed of subparallel hyphae, and lamellae that are variously attached but never distinctly free. These fungi are closely related to species of Clitopilus and of Rhodocybe which have spores that are angular in end view. In Clito pilus, the spores are longitudinally grooved or striate when seen in side or face view; in Rhodocybe, the spores are roughened to warty when seen in face or side view. Several other genera also have pink spores, but differ from the rhodophylloid genera in other ways. For example, the punctate- roughened spores of Lepista are not angular in any view. Rhodotus has smooth to echinulate spores, free lamellae, and divergent lamellar trama. Volvariella, Chamaeota, and Pluteus all possess smooth spores, converg- ent lamellar trama, and free gills; furthermore, Volvariella has a volva, Chamaeota an annulus.

CHROMATOGRAPHIC METHODS

One hundred mg of finely ground gill, pileus and stipe tissue was placed in a screw cap vial, along with 1.5 ml ethanol and agitated vig- orously on a rotary shaking machine for approximately 24 hours. Fifty vl of clear supernatant of each extract was spotted one inch from the base of 944 X 221%”, oxalic-acid-washed Whatman No. 1 filter paper, together with reference spots of 5, 10 and 25 ug of urea respectively. The chromatogram was developed with a wash liquid composed of n-butanol-acetic acid-water (4:1:1) for approximately 18 hours. Four to five chromatograms were run simultaneously in each chromatographic

34 MADRONO [Vol. 21

TABLE 1. UREA AS A CHEMOTAXONOMIC INDICATOR IN CERTAIN RHODOPHYLLOID FUNGI

Genera (sensu Dennis, et al. Number of Species Urea Concentrations Claudopus 1 @Q)e 5 Nolanea 5 (34) 5 Entoloma ((alp) 0-2 Leptonia 5 (42) 0-0.3

*Figures in parentheses are number of collections.

chamber located in a constant temperature room at 25°C. The chroma- tograms were dried at room temperature, marked to show the location of fluorescent spots and then sprayed with 2% p-dimethylaminoben- zaldehyde in hydrochloric acid (Ehrlichs reagent, hereinafter referred to as PDAB). (Ehrlich’s reagent: p-dimethylaminobenzaldehyde (PDAB), 6 grams; Ethanol, 95%, 229 ml; and Concentrated hydro- chloric acid, 71 ml. If the spray is stored in a colored bottle and kept below room temperature, it should remain stable for extended periods.)

The Rf values were calculated for all unknown compounds and for three urea standards. The sizes and color intensities of yellow spots with the same mobility as the three standards were compared with the latter. Extracts rated O-1 contained none to approximately 5 pg/spot; 2, around 10 pg; 4, about 25 »g and 5, more than 25 ung.

RESULTS

Although 77 compounds were detected in the 279 extracts examined, only those compounds forming a yellow chromophore at the same Rf value as those of the urea standards (0.50-0.06) are of chemotaxo- nomic concern in the present study. Ninety-five of the 279 collections an- alyzed in the above manner were identified to species by classical micro- scopic and macroscopic techniques. The placement of these species into genera (sensu Dennis, et al., 1960), together with data on their urea concentrations, are shown in Table 1.

The data in Table 2 include Pouzaromyces Pilat and a segregate of Leptonia (L. sericella species complex). The data further show that urea levels in all collections placed in Nolanea were consistently high, whereas those in Leptonia and Entoloma were much lower.

TABLE 2. UREA IN 279 COLLECTIONS OF RHODOPHYLLOID FUNGI

Genera Urea Concentrations O-1 2 4 5 Leptonia sericella 15* 15 0 0 0 species complex Pouzaromyces 2 0 1 0 1 Leptonia 86 63 15 8) 5 Entoloma 66 40 2 1 Nolanea 108 0) 0) 1 107 Claudopus 2 0 0) 0 Z Total 279

*Number of collections examined.

1971] LARGENT & BENEDICT: FUNGI 35

KEY TO THE GENERA

Stipe eccentric, lateral, or lacking. . . ..... .. . . . +. Claudopus Stipe central. Pileal surface in radial section a layer of repent, radially oriented hyphae.

Urea concentration low (0 to +1, rarely +2); clamps usually present on 20% or more of the pileal cuticular hyphae; carpophores large and fleshy (pileal trama 3 mm or more at pice of umbo, apex of stipe 5 mm or more in diam. . . . . . . Entoloma

Urea ponecnariien high to very Gent Ga to sys series absent or if present, on less than 10% of the pileal cuticular hyphae; carpophcres usually small or medium sized, and thin-fleshed (pileal trama 2 mm or less, apex of stipe less than 5 mm), but sometimes large and fleshy as in Entoloma . Nolanea

Pileal surface in radial section an entangled to irregularly interwoven tricho- dermium, a palisade trichodermium, an ixotrichodermium, or a hymeniform layer (at least when young).

Urea concentration high to very high (+4 to +5); pileus thin-fleshed, conic to campanulate, often with a papillate or cuspidate umbo, the cuticle often overlain by a colorless superficial layer or veil. . . . . . . Nolanea

Urea concentration low to medium (0 to +3); pileus varying in shape and thickness of flesh, but not conical with papillate or cuspidate umbo.

Pileal cuticle a palisade trichodermium, cr hymenifcrm, at least on the disc . . . . . . . Leptonia Pileal cuticle an Se acal fo an aaah, Pan TOn trichodermium or an ixotrichodermium. Terminal cells of pileal cuticle averaging less than 8 uw wide. . Entoloma Terminal cells averaging 8 u cr more wide. Young, fresh carpophores entirely white to pale cinerous. Alboleptonia Carpophores not entirely white to pale cinereous. Cuticle at apex of stipe similar to pileal cuticle. Cuticular hyphae of the pileal disc 190-600 wu long, 5-10(-15) -septate, average length greater than 250 u.. . . Pouzaromyces Cuticulzr hyphae of the pileal disc up to but not exceeding 300 uw in length, (1-)3-5-septate, average length 225 wor less . Leptonia Stipe and pileal cuticles not similar. Caulocystidia, if present, versiform but not indense clusters . Leptonia Caulocystidia obclavate to aculeate and in rosette-like clusters. Pouzaromyces

GENERA

Ciaupopus (W. G. Smith) Gill., emend Pat, Les Hyménomycetes d’Europe, 113. 1887. Basionym: Agaricus subgenus Claudopus W. G. Smith, Clavis Agaricinorum. Type species: Agaricus byssisedus Pers. per Fr.

In members of this genus the carpophore often lacks a stipe; when one is present, it is always lateral or eccentric. Also, the habit of the carpophores of growing on the underside of logs or of shelving clumps of moss has not been encountered in other fungi with pink, angular spores. Leptonia dichroa and related lignicolous species have a tricholo- matoid carpophore.

Extracts from only two collections of Claudopus were chromato- grammed. Each possessed a high concentration of urea, but otherwise had no distinctive Erlich- positive spots.

36 MADRONO [Vol. 21

ENTOLOMA (Fr.) Kummer, Der Fuhrer in die Pilzkunde, 23, 97. 1871. emend. Basionym: Agaricus tribus Entoloma Fr., Epicrisis, 143. 1838. Synonym: Agaricus subgenus Entoloma (Fr.) Rabenh., Deutschl. Krypt.- Fl. 1:508. 1844. Type species: Agaricus prunuloides Fr.

Pileus: glabrous, at times with a pruina of whitish or grayish fribrils, or with minute colorless squamules; surface dry, lubricous, or viscid; context 3 mm or more thick at the edge of the umbo. Stipe: 5 mm or more broad at the apex. Pileal cuticle: in radial section, either of repent hyphae, or an entangled to interwoven trichodermium, or an ixotricho- dermium; terminal cells long, thin, claviform, averaging less than 8.0 p» wide; hypoderm similar to the pileal cuticle. Clamp connections: on at least 20% or more of the septa of all pileal cuticular hyphae, long, thin, at times rare. Pigmentation: vacuolar, rarely externally incrusted. Urea concentration: low (0 to +1).

Deviations from one or another of these characteristics are encoun- tered. For instance, some specimens have very few clamps on the pileal cuticular hyphae, but are recognizable as Entolomas by their fleshy, tricholomatoid carpophores with glabrous pileus and low urea content. In other specimens the pileus instead of being glabrous is covered with a pruina of whitish or grayish fibrils, or with minute colorless squamules, and in still others, externally incrusting pigment replaces vacuolar pig- mentation. But in all of these instances, however, the specimen can be identified as an Entoloma by its fleshy carpophore, low urea content, and relatively abundant clamp connections.

NoLanEA (Fr.) Kummer, Der Fuhrer in die Pilzkunde, 24, 95. 1871. emend. Basionym: Agaricus tribus Nolanea Fr., Syst. Mycol. 1:204. 1821. Type species: Agaricus pascuus Fr.

Pileus: convex to campanulate or conic, rarely depressed or umbili- cate, often umbonate, papillate, or cuspidate; surface glabrous, at times overlaid with a superficial veil, dry or lubricous; context 2(-5) mm or less thick at the center or at the edge of the umbo. Stipe: 4(-8) mm or less broad at the apex. Pileal cuticle: in radial section either of repent hyphae or an entangled to interwoven trichodermium; hypoderm often differentiated. Clamp connections: absent or if present, on less than 10% of the septa of all pileal cuticular hyphae, rarely more abundant. Pigmentation: vacuolar or etxernally incrusted. Urea Concentration: high (+4 to +5).

In occasional specimens the pileus is depressed to umbilicate, but more often, the carpophore is almost tricholomatoid in stature, and has clamps on more than 10% of the septa in the pileal surface. In both instances the fungus can be included in Nolanea, because the specimens also possess the distinctive combination of high urea content and repent pileal cuticle.

Nolanea can be distinguished from Entoloma by its pileal cuticle whose hyphae are well differentiated from those of the pileal trama, by the absence or rarity of clamp connections, the less fleshy carpophore,

1971] LARGENT & BENEDICT: FUNGI 37

and more reliably by the high urea concentration. Leptonia, Pouzaro- myces, and the Leptonia sericella species complex all differ from Nolanea by virtue of the different structure of their pileal cuticle, and their low urea content.

PouzAroMyces Pilat, Sborn. Narodn. Mus. Praze 9B (2):60. 1953. Type species: Agaricus fumosellus Winter.

The species of this genus are distinguished by their dark brown to gray-brown tomentulose to densely scaly pilei, initially dark brown to gray-brown lamellae, and pruinose to densely scaly stipes that are con- colorous with the pilei. Cheilocystidia are present on the lamellar edge. The pileal cuticle is composed of septate, entangled hyphae with dark brown vacuolar pigment and some externally incrusted pigment. The cuticle at the apex of the stipe is either similar to the pileal cuticle, or bears rosette-like clusters of long, obclavate to aculeate caulocystidia.

Pouzaromyces can be distinguished from Entoloma, Nolanea, the Leptonia sericella species complex, and most other Leptonias on the basis of color, and of the nature of the surface and cuticle of both pileus and stipe. Certain species of Leptonia (e.g., L. subdysthales) resemble Pouzaromyces in general appearance, especially as to color, but differ in the structure of the pileal cuticle, or the cuticle of the stipe, or both.

LeptToniaA (Fr.) Kummer, Der Fuhrer in die Pilzkunde, 24, 96. 1871. emend. Basionym: Agaricus tribus Leptonia Fr., Syst. Mycol. 1:201. 1821. Synonyms: Eccilia (Fr.) Kummer, Der Fuhrer in die Pilzkunde, 23, 94. 1871. Leptoniella Earle, Bull. N. Y. Botan. Gard. 5.424. 1909. Type of species: Agaricus euchrous Pers. per Fries.

Pileus: tomentulose or punctate or squamose to squamulose, at least on the disc, at times silky-, matted-, or appressed-fibrillose. Pileal cuticle (in radial section and at least on the disc): an entangled trichodermium, hymeniform, or a palisade trichodermium; pilocystidia or cystidioid terminal cells 84 or more in diameter. Clamp connections: present or absent. Pigmentation: vacuolar or rarely incrusted. Urea concentration: variable (0 to +3).

The species of Leptonia are characterized primarily by the nature of the pileal surface and the structure of the pileal cuticle. The surface is tomentulose or punctate or squamose to squamulose, at least on the disc, and the cuticle is an entangled trichodermium of agglutinated or non- agglutinated hyphae, or is hymeniform, or a palisade trichodermium. Pilocystidia or cystidioid terminal cells are 8u or more in diameter, with vacuolar (rarely incrusted) pigment. The cuticle at the apex of the stipe is of repent hyphae, or bears scattered clusters of caulocystidia. Clamps may be present or absent. Most of the species have small to medium- sized, rather thin-fleshed carpophores, but there are some exceptions. A fleshy stature, or frequent clamp connections on the pileal cuticular hyphae, or both, relates some species of Leptonia (e.g., L. jubata, L. dichroa, L. griseo-cyanea, and L. chalybaea) to Entoloma. However,

38 MADRONO [Vol. 21

the species of Entoloma differ in their smooth, glabrous pilei, the struc- ture of their pileal cuticle, and the nature of their pilocystidia.

In Leptonia, urea concentration varies from 0 to +3. Of 86 collections examined, 78 had little or no urea (0 to +1). Eight collections, including 3 each of L. dichroa and L. jubata, showed a concentration of +2 to +3. All eight belong to sections of Leptonia that do not have a tomentulose pileal disc, thus there seems to be a correlation between urea concen- tration and the type of pileal surface, which may have significance at the sectional level.

One group of Leptonias, referred to previously as the L. sericella species complex, seems different enough from the remainder of the genus to warrant special attention. This group of species has been placed in a new genus, Alboleptonia (Largent and Benedict, 1970). This group is characterized by the entirely white to pale cinereous carpophore, the silky to appressed-fibrillose or minutely squamulose pileal surface con- sisting of an entangled trichodermium, the presence of a fugacious super- ficial veil, low urea concentration (0 to +1), and a set of unique Ehrlich- positive compounds. The few white Leptonias not included in Aldo- leptonia (Leptonia albinella and related species) have the pileal cuticle hymeniform, or consisting of a palisade trichodermium. Pallid or nearly white species of Entoloma differ in their glabrous, dry to viscid pilei, larger stature, abundant clamp connections, and dissimilar Ehrlich-posi- tive compounds.

ACKNOWLEDGEMENTS

We wish to acknowledge financial support of this study from the Na- tional Institute of Health, for Research Grant GM-07515-07, summer of 1967. One of us (D.L.L.) also received financial help from Sigma Xi (Grant-in-Aid of Research), the New York Botanical Garden (Gertrude S. Burlingham scholarship), and the National Institute of Health (Public Health Service Fellowship, 1-F1-GM-36, 352-01). We wish to thank D. E. Stuntz for his critical reading of the manuscript and for his valu- able comments.

SUMMARY

As suggested by Tyler, et al. (1965), a study of the urea concentration in species of Rhodophyllus proved to be a useful chemotaxonomic aid in classifying these fungi. On the basis primarily of urea concentration, type of pileal surface, and structure of the pileal cuticle, plus the occa- sional use of other structural or macroscopic features, five genera can be delimited, as follows:

Claudopus. Stipe lacking, or present and lateral to eccentric; place of growth, on the underside of logs or on shelving clumps of moss.

Entoloma. Clamps long, thin, relatively abundant; pileal surface glabrous or frosted; pileal cuticle repent, or an interwoven trichoderm- ium; pilocystidia less than 8u broad; urea concentration low.

Nolanea. Clamps absent or rare; pileal surface glabrous, dry or lubri-

1971] CONNOR CORTADERIA 39

cous; pileal cuticle of repent hyphae; urea concentration high.

Pouzaromyces. Clamps absent or rare; pileal surface minutely to- mentose to densely scaly; surface of stipe pruinose to densely scaly; pileal cuticle of septate, entangled hyphae with dark pigment; cheilo- cysidia present; apex of stipe with a distinct cuticle; urea concentration not determined.

Leptonia. Clamps usually absent, if present, thick and rather numer- ous; pileal surface appressed-fibrillose, tomentulose, punctate, or squa- mulose, at least on the disc; pileal cuticle hymeniform, a palisade tricho- dermium, an entangled trichodermium, or an intewoven layer of submoniliform hyphae; pilocystidia broad; urea concentration low to medium.

In addition to the above five genera, a distinctive group of species, the Leptonia sericella species complex (now Alboleptonia Largent & Benedict, 1970), can be characterized as follows: entire corpophore white to pale cinereous; clamps rarely present; pileal surface silky to minutely squamulose; pileal cuticle an entangled trichodermium with a fugacious superficial veil; several unique Ehrlich-positive compounds present.

LITERATURE CITED

Dennis, R. W. G., P. D. Orton, and F. B. Hora. 1960. New check list of British agarics and boleti. Trans. Brit. Mycol. Soc. 43. Suppl.

Fries, E. M. 1821. Systema mycologicum. Vol. I. Ex Officina Berlingiana, Lund.

. 1838. Epicrisis systematis mycologici. Gleerup, Lund.

Hester, L. R. 1967. Entoloma in southeastern North America. Beih. Nova Hed- wigia 23.

KUHNER, R., and H. Romacnesi. 1953. Flore analytique des champignons supérieurs. Mason et Cie., Paris.

LarGEeNT, D. L. and R. G. BENeEpict. 1970. Studies in the rhodophylloid fungi IT: Alboleptonia, a new genus. Mycologia 62:437—452.

SmitH, A. H., and R. L. SHarrer. 1964. Keys to generat of higher fungi. Univ. Michigan Biol. Station, Ann Arbor.

Tyrer, V. E., R. G. BeneEpict, and D. E. Stuntz. 1965. Chemotaxonomic signficance of urea in the higher fungi. Lloydia 28:342-353.

A NATURALIZED CORTADERIA (GRAMINEAE) IN CALIFORNIA

H. E. CONNOR

Botany Division, Department of Scientific and Industrial Research, Christchurch, New Zealand

Munz (1968) recorded Cortaderia selloana (Schult.) Asch. & Graebn. as naturalized in San Francisco and the North Coast Ranges. He further referred to a heavy infestation of this species at Big Lagoon, Humboldt Co. From the Big Lagoon populations D. W. Cooper of Eureka sent me specimens, transplants, and seed from which plants were raised. All have proved identical with plants widely naturalized in northern North

40 MADRONO [Vol. 21

Island, New Zealand. These are referred by Connor (1965) to C. atacamensis (Philippi) Pilger following the treatment of Chilean Cortaderia by Acevedo de Vargas (1959).

The two most recent revisions of Cortaderia for the section of the genus including C. selloana, atacamensis, and rudiuscula (Acevedo, 1959; Conert, 1961) are not in good agreement, and Conert does not cite Acevedo’s paper. Acevedo (1959) truly describing C. atacamensis as ““Hermosa Cortaderia,” also referred to the confusion between it and C. rudiuscula Stapf emend. Acevedo; she distinguishes C. atacamensis from C. rudiuscula by such characters as floret size, awns of lemmata, branching habit and panicle color. Conert (1961) preferred the often used combination C. quila (Nees & Meyen) Stapf to C. rudiuscula. While I cannot be sure that all records of C. rudiuscula in the United States refer to the entity here reported as C. atacamensis (cf. Bailey, 1949; Chase, 1950), it is certainly true of the recent report from Marin Co., California (Howell, 1970; pers. comm.).

Some points of comparison between C. selloana and C. atacamensis are given in Table 1.

TABLE 1. COMPARISON BETWEEN CORTADERIA SELLOANA AND C, ATACAMENSIS.

Leaf color Leaf blades

Leaf sheath Panicle color Panicle branches

Lemma

Sex form

Flowering time

Chromosome no.

atacamensis deep green

hairy towards base on abaxial surface

densely hairy deep violet flexuous

acuminate or shortly awned or mucronate; the small awn is very fragile and breaks off easily

female only late summer

2n = 108

LITERATURE CITED

selloana bluish green

glabrous on abaxial surface

glabrous or faintly hairy white through to light violet stiff

strongly awned

hermaphrodite and female mid to late autumn

In

ACEVEDO DE VarRGAS, R. 1959. Las especies de gramineas del genero Cortaderia en Chile. Bol. Mus. Nac. Hist. Nat. 27:205-246.

BarLey, L. H. 1949. Manual of cultivated plants. Macmillan, New York.

Cuase, A. 1950. Manual of the grasses of the United States. U. S. D. A. Misc. Publ.

200.

Conert, H. J. 1961. Die Systematik und Anatomie der Arundineae. J. Cramer,

Weinheim.

Connor, H. E. 1965. Breeding systems in New Zealand grasses. V. New Zealand J.

Bot.$¢ 17-25.

HowELt, J. T. 1970. Marin flora. Ed. 2. Univ. Calif. Press, Berkeley. Muwz, P. A. 1968. Supplement to a California flora. Univ. Calif. Press, Berkeley.

MORPHOLOGY, CHROMOSOME NUMBER, AND FLAVONOID CHEMISTRY OF BIDENS CORDYLOCARPA (COMPOSITAE)

DANIEL J. CRAWFORD Department of Botany, The University of Wyoming, Laramie 82070

Coreopsis cordylocarpa was described by Gray in 1887. For many years no one seems to have questioned that this species had an appro- priate generic assignment. In his two comprehensive treatments of Coreopsis, Sherff (1936, 1955) does not even suggest that C. cordylo- carpa might have been improperly placed. In his later years, however, he indicated a specimen in the Field Museum (Cronquist 9779) as the type of a new Bidens species which he proposed to name in honor of Arthur Cronquist. The name was never published, and Sherff later recognized the Cronquist specimen as belonging to Coreopsis cordylocarpa, and so anno- tated it.

My first encounter with this species in the field was in the state of Jalisco in the late summer of 1966, where I was collecting with T. Melchert and P. Sorensen. All three of us were somewhat familiar with members of the Coreopsidinae, and it seems significant that our first impression was that we were observing a species of Bidens. In view of this, along with the fact that Sherff had also at one time assigned this taxon to Bidens, a detailed study seemed indicated. The present paper gives the result of this study.

MATERIALS AND METHODS

For the cytological studies, floral buds were fixed in the field in chloro- form: absolute ethyl alcohol: glacial acetic acid (4:3:1). The anthers were squashed in aceto-hematoxylin, and the chromosomes observed in dividing microsporocytes.

Leaves and flowers which were collected in the field and dried, served as one source of flavonoids for chromatographic analysis. Achenes were collected at the same time, and fresh material from plants grown in the greenhouse was also analyzed for flavonoid constituents. The floral tissues (ray floret corollas, disk floret corollas and adnate anthers, disk floret ovaries, and chaff) were analyzed separately and found to be chromatographically identical. The leaf profiles were also determined. The tissues were placed in 0.1% HCl in methanol for 24—48 hours. This extract was applied to Whatman 3MM chromatographic paper (46 57 cm sheets) and run in two dimensions by the descending method. The first solvent system was tertiary butyl alcohol: glacial acetic acid: dis- tilled water (3:1:1 v/v); the second glacial acetic acid: distilled water (15:85 v/v). Drawings of these chromatograms are shown in Fig 5. In these figures each chalcone-aurone pair is designated by a single letter and represented as a single spot because they invariably occur together as a complex mixture.

Individual flavonoids were purified by repeated chromatography, and

41

42 MADRONO [Vol. 21

TABLE 1. IDENTIFICATION AND SPECIAL MAXIMA OF THE FLAVONOIDS OF C. CORDYLOCARPA

Absorption maxima in mu Spot +A1C1;/ +NaOAc/ Designation Identity MeOH +AIC1; HC1 +NaOMe +NaOAc H;BO;

Leaves 6 quercetin-3- 359 432 404 413 381 381 glycoside 295 330 365 330 325 295a 265a 300a 300a 272 271 261 255 274 267 16 naringenin-7- 330 385 385 445 330 330 glycoside 283 306 306 390a 283 283 286 Leaves and floral tissues A coreopsin 383 505a 440 450 480a 510a 295 450 320 380a 385 413 260 318 I) 285 288 345a 245 PES 250 255a 285 250 A sulfurein 403 450 405 488 487 435 340a 342 335a 345 405 Sai 274 292 215 291 340a 288 256 255a 257 278 259 255 F marein 382 520a 420 452 383 393 320a 429 332 340 325a 320 263 3O2 272 285 265 270 213 250 253 F maritimein 416 455 413 497 443 442 330 328 BAS) 348 365 328 213 287 21S 290a 259 282 242 248a 243 261 245

Floral tissues xX butein-sulfure- —-— — ee fae ae ee tin mixture?

a—denotes a shoulder or inflection.

the compounds were finally analyzed spectrally, utilizing a Beckman DB-G Grating Spectrophotometer. Standard methods and diagnostic reagents were employed (Markham and Mabry, 1968; Jurd, 1962). The spectral properties of these compounds, together with their identi- fications, are given in Table 1.

RESULTS AND DISCUSSION

There are a combination of morphological features which serve to distinguish this species from other taxa in the genus Coreopsis found in

1971] CRAWFORD: BIDENS 43

Fic. 1. Distribution of C. cordylocarpa.

Mexico. The (8)10-16 ray florets, club-shaped wingless achenes (fig. 3B), fruticose habit, relatively undifferentiated outer and inner in- volucral bracts (fig. 2B), and large (to 20 cm) pinnatisect, deltoid leaves (fig. 4A) are quite unique. Sherff (1955) treated C. cordylocarpa as a member of sect. Coreopsis. It is clearly a discordant element here, how- ever, for all other species in this section are small annual or perennial herbs with dorsiventrally flattened, winged achenes. In fact, all species of Coreopsis which I have examined (primarily those from North Amer- ica) have achenes which are variously flattened dorsiventrally. That Sherff placed C. cordylocarpa in the type section seems to indicate a lack of understanding of its affinities within the genus Coreopsis.

Certain morphological features of C. cordylocarpa are much more suggestive of some Mexican Bidens species than they are of any member of Coreopsis. Specifically, the outer and inner involucral bracts of C. cordylocarpa are quite similar in shape (fig. 2B), and differ primarily in color, the outer ones being dark green, whereas the inner are pale green to nearly white. These involucral characteristics are very similar to those encountered in many species of Mexican Bidens. In contrast, species of Mexican Coreopsis typically exhibit a highly dimorphic in- volucre with somewhat green and fleshy outer bracts which differ from the inner ones in shape, size, color, and texture (fig. 2A). Moreover, the elongate, club-shaped, terete, striate, and wingless achenes of C. cordy- locarpa are similar in general shape and appearance to those of several species of Bidens from Mexico (fig. 3A, B). Certainly, the achenes of C. cordylocarpa in no way resemble the flat, winged fruits which are typical of all Mexican Coreopsis, and indeed of the genus as a whole (fig. 3B, C).

The chromosome number of C. cordylocarpa offers no clues as to its

44 MADRONO [Vol. 21

Fic. 2. Photographs of flloral heads. A, floral head of typical Mexican Coreopsis showing the highly dimorphic involucre; B, floral head of C. cordylocarpa showing the undifferentiated outer and inner involucral bracts.

generic affinities. A count of 2n = 146 = 2 (fig. 4B) was determined in a large number of cells from Melchert, Sorensen, & Crawford 6347A. It must be emphasized that observations from several other populations (Melchert, Sorensen, & Crawford 6354 & 6371; Carman 68-60) re- vealed a chromosome complement of 2n — 146 + 6-8. From these data, it appears justifiable to conclude that only one ploidy level exists in C. cordylocarpa, and that probably all populations have the same or nearly the same chromosome number. This high number, unique in the Coreopsidinae and one of the highest reported in the Compositae, is particularly interesting from an evolutionary point of view when con- sidered together with the geographic distribution and ecology of the species. Coreopsis cordylocarpa is endemic to Jalisco, Mexico (fig. 1) and occurs only in or along the banks of shallow streams, indicating that it may be an old species, representing the only extant taxon of an otherwise extinct polyploid complex.

The flavonoid chemistry of C. cordylocarpa suggests a closer affinity to other Mexican species of Bidens than to any Coreopsis taxon. As shown in Fig. 5, the leaves and floral tissues are dominated by two chalcone-aurone pairs. Coreopsin-sulfurein (spot A) and marein-mari- timein (spot F) are invariably present in large quantities in both leaves and flowers. In addition, the leaves (fig. 5, left) contain a flavonol (spot 6, quercetin-3-glycoside) and a flavanone (spot 16, naringenin-7- glycoside). The flowers also contain spot X (fig. 5, right), which ap- pears to be a mixture of butein and sulfuretin, these being the aglycones of coreopsin and sulfurein respectively.

Chemical analysis of the leaves of other suffruticose or fruticose Coreopsis species from Mexico (members of sections Electra, Anathy- sana, and Pseudo-Agarista) has revealed the complete absence of coreopsin-sulfurein and marein-maritimein. These compounds are some- times present in the floral tissues of certain of these species, but never

1971] CRAWFORD: BIDENS 45

A B

Fic. 3. Drawing of Coreopsis and Bidens achenes. A, achene of a typical Mexican Bidens; B, achene of C. cordylocarpa; C, achene of a typical Mexican Coreopsis (all x< ca. 5).

in the leaves. It must be admitted that sufficient data are not available to make a meaningful statement concerning the distribution of these substances in the genus Coreopsis as a whole. However, it is instructive to compare the leaf profile of C. cordylocarpa to those of several species of Bidens from the United States and Mexico. The leaves of these taxa contain an unidentified chalcone-aurone pair which is chromatograph- ically very similar (probably identical) to marein-maritimein. This evaluation is based upon conversations with T. E. Melchert and my observations of numerous chromatograms of the leaves of Bidens species. Although the chemical evidence is not conclusive, it certainly suggests that C. coryvlocarpa is much more similar to Bidens in its flavonoid chemistry than it is to Coreopsis.

Since the general morphology, as well as the preliminary chemical data, suggest that the affinities of C. cordylocarpa are with Bidens rather than with Coreopsis, the following new combination is proposed.

46 MADRONO [Vol. 21

é iS “Ge © yal @ |) eS & oO” Bae. 9 6347 46h ae © oe @ 4. a S0%

A B

Fic. 4. A, silhouette of leaf of C. cordylocarpa (X ca. 1/2); B, meiotic chromo- somes of C. cordylocarpa.

Bidens cordylocarpa (A. Gray) Crawford, comb. nov. Coreopsis cordylocarpa A. Gray, Proc. Amer. Acad. Arts 22:428. 1887. Fruticose, 0.5-2m tall, stems several from the base, red, hispid or with appressed hairs, becoming glabrous toward the base; leaves opposite, 10-20 cm long (including petiole), deltoid in general outline, pinnately divided, appressedly-pubescent on both surfaces; heads cymosely disposed, mostly 3-12 aggregated, peduncles 2-15 cm long, becoming densely pubescent near the involucre; heads 4-9 cm wide at anthesis; outer involucral bracts 6-10, lanceolate to narrowly so, hispid, 3-9 mm long; inner involucral bracts 8-12, lanceolate to narrowly ovate, hispid, 4-8 mm long; chaff narrowly lanceolate to linear, glabrous or sparsely hispid, 5-8 mm long at anthesis; ray florets 8-16, sometimes in a double whorl, neutral, ligule oblong to oblanceolate or linear, 0.6—4 cm long, 0.4—1.2 cm wide, entire or shallowly notched at the apex; disk florets 20-60, stigma hispid, shortly caudate; achenes club-shaped, essentially terete in cross section, glabrous, weakly striate, wingless, exaristate, and topped by a bald disk.

Representative specimens: MEXICO. Jalisco, bank of stream, 5200 ft. Sierra de San Estéban, Barnes & Land 155 (F); banks of Rio Blanco near Guadalajara, 5000 ft., Pringle 11506 (F, MICH, MO, MSC, US); gravel along small stream 15 road mi N of Guadalajara, on road to San Crist6ébal de la Barranca, 5100 ft., Cronquist 9817 (F, MICH, MO, MSC, NY, TEX, US); in boulders and sand of stream bed leading into the barranca of the Rio Blanco, ca. 8 mi N of Guadalajara, Mel-

1971] CRAWFORD: BIDENS 47

Fic. 5. Drawings of two dimensional chromatographic patterns of flavonoids in C. cordylocarpa (horizontal axis = tertiary butyl alcohol run; vertical axis = 15% acetic acid run) ; left, profile of leaves; right, profile of floral tissues.

chert, Sorensen, & Crawford 6354 (IA, RM); among boulders of swift stream, ca. 3 mi W of Cuaulta along road to Los Volcanes and Puerto Vallarta, Melchert, Sorensen, & Crawford 6371 (IA, RM); among boulders of rocky stream bed 12.5-13 mi N of Zapopan, along dirt road to San Cristébal de la Barranca, Melchert, Sorensen, & Crawford 6347 A-B (IA, RM); Hwy. 41, 7-8 mi N of Guadalajara, Carman 68-60 (IA, RM).

ACKNOWLEDGEMENTS

The research was supported by NSF Grant GB-3851 to T. E. Mel- chert, University of Iowa, whose guidance during the study is much appreciated. Thanks are also due R. L. Hulbary, University of Iowa, who prepared the photographs of the floral heads, and to J. R. Reeder and Charlotte G. Reeder for helpful suggestions during the preparation of the paper. Ruby S. Quarterman, Laramie, executed the drawings of the achenes.

LITERATURE CITED

Jurp, L. 1962. Spectral properties of flavonoid compounds. Jn T. A. Geissman (Ed.). The chemistry of flavonoid compounds. Pergamon Press, Oxford & New York. MarxuaM, K. R., and T. J. Masry. 1968. A procedure for the ultraviolet spectral detection of ortho-dihydroxy] groups in flavonoids. Phytochemistry 7:1197-1200. SHERFF, E. E. 1936. Revision of the genus Coreopsis. Fieldiana Bot. 11:277-475. . 1955. Coreopsis. In E. E. Sherff and E. J. Alexander. Compositae—Hel- iantheae—Coreopsidinae. North Amer. FI. II. Pt. 2.

NOTES AND NEWS

NOTES ON THE FLORA OF THE PaciFic NoRTHWEST.—Extensive collections from Pend Oreille Co., Washington, were made by the author in connection with a floristic study (Layser, E. F. A floristic study of Pend Oreille County, Washington. M.S. thesis, State Univ. New York, College Forestry. 1969). Among the collections, certain ones seem worth special note.

Berteroa incana (L.) DC., a weedy European crucifer, was collected along the roadside in the northern part of Pend Oreille Co. (Layser 1175, WS) and previously

48 MADRONO [Vol. 21

was known in the Pacific Northwest from British Columbia, Idaho, and Montana.

Carex sychocephala Carey, was reported by C. L. Hitchcock, et al. (Vascular plants of the Pacific Northwest, 5 vols., Univ. Washington Press. 1955-1969) from the Pacific Northwest from Montana, Kamloops, British Columbia, and Okanogan Co., Washington, and was noted as being “Seldom collected in our range.” Since then it was found along a slough of the Pend Orielle River (Layser 1201, WS).

A hitherto undescribed form of Crepis was discovered at two different localities 10 miles apart. With respect to the polyploidy and apomictic induced polymorphic nature of this group, it is being treated as an anomalous form within a heteroploid complex (Babcock, E. B., and G. L. Stebbins. The American species of Crepis. Publ. Carnegie Inst. Wash. 504. 1938). The decision to recognize this ferm is based on its diverse morphology and its distribution.

CREPIS ATRABARBA Heller ssp. ORIGINALIS Babc. & Stebb. forma pend-oreillensis Layser, forma apm. nov. Caulis 3-5 dm alta; basalia folia linearis, 8-20 mm. longa, 2-6 mm lata, non pinnata vel rudimentaria; fclia caulis linearis, dimidia supra; inflorescentia cymosa, axia centralis; capitulis 3-8; involucra 10-12 mm longa, tomentosa; flosculi 12-13; coma 7-8 mm longa; achenia viridula, 8 mm longa.

Material examined. Washington: Pend Oreille Co. Dry, rocky slopes in Dry Canyon, south of Cato Creek, Sec. 23, T37N, R37E, alt. 4000 ft., E. F. Layser 894 (WS-holotype), June 1969; dry, shallow-soiled rocky slopes on Hall Mt., alt. 3500 ft., E. F. Layser 830 (WS), June 8, 1969.

Geum rivale L. was known in Washington from one locality in Okanagon Co. (Thomsen, J. W. Notes on the flora of the state of Washington. Rhodora 36:8-13. 1934). It has now been found in wet meadows in Pend Oreille Co. (Layser 127, WS).

Hieracium aurantiacum L. was noted by Hitchcock, et al. (op. cit.) to occur at Bremerton, Washington. The species is not uncommon along roadsides in the northern part of Pend Oreille Co. (Layser 891, NY, WS). This weedy species is rapidly becoming established in the Pacific Northwest.

Hieracium pratense Tausch, an aggressive European weed naturalized in eastern North America, has become established in the northern part of Pend Oreille Co. (Layser 829, NY, WS). This constitutes the first report of it from the Pacific North- west, where it appears to have been introduced through erosion control seedlings of logging skid roads and road banks.

Mertensia platyphylla Heller was found in Pend Oreille Co. (Layser 134, F) and is the first report of this species east of the Cascades. The significance of this collec- tion may involve more than a range extension, that is, a reassessment of the taxonomic affinities of M. paniculata (Ait.) G. Don. and M. platyphylla may be in order. Formerly these two species were thought to be allopatric. Evidence to the contrary not withstanding, this collection may provide support to the statement by Hitchcock (op. cit.) that M. platyphylla might better be considered a variety of M. paniculata.

Penstemon ellipticus Coult. & Fisch, is not uncommon in the northern part of Pend Oreille Co. and adjacent Stevens Co. on mountain peaks above 6000 ft. in elevation (Layser 503, 867, WS), and is an addition to Washington’s flora.

Ranunculus longirostris Godr. was collected from Pend Oreille Co. (Layser 80, POM, WS) and is new for the state of Washington.

Sorbaria arborea Schneid, the cultivated false spirea, is occasionally found about old abandoned homesteads, where it reproduces and maintains itself by suckering (Layser 1143, WS).

Stellaria calycantha (Ledeb.) Bong. var. calycantha represents a new variety for Washington. It was collected in the northern part of Pend Oreille Co. (Layser 310, NY, WS). This variety is known from high latitudes in North America and was also reported from Oregon (Fernald, M. L., Gray’s manual of botany. American Book Co., New York. 1950).—EarteE F. Layser, Department of Botany, Washing- ton State University, Pullman 99163.

ew

DRONO

VOLUME 21, NUMBER 2 APRIL 1971

Contents

EVIDENCE OF DIFFERENT ADAPTATIONS OF FLOWER COLOR VARIENTS OF ENCELIA FARINOSA (ComposiTAE), Donald W. Kyhos 49

A New MYcocaLicIuM ON SCARRED SEQUOIA IN CALIFORNIA, Lee Bonar 62

THE GALIUM ANGUSTIFOLIUM COMPLEX (RUBIACEAE) OF CALIFORNIA AND BAJA CALIFORNIA, Lauramay

T. Dempster and G.. Ledyard Stebbins 70 A NEw TETRAPLOID SUBSPECIES OF LASTHENIA

(COMPOSITAE) FROM OREGON, Robert Ornduff 96 NEw Moss REcorpDs FROM Mexico, Claudio Delgadillo

M. and Dale H. Viti 99 SYSTEMATIC STUDIES OF LIMNANTHACEAE,

Robert Ornduff 103 NoTES AND NEws: WALLACE Roy ErRwnsT, 1928-1971 69

Two SPECIES OF PANICUM (POACEAE) NEw TO

OrEGON, Richard Spellenberg 102

Reviews: Ira L. Wiggins and Duncan M. Porter, Flora of the Galdpagos Islands (Wallace R. Ernst) ital

A WEST AMERICAN JOURNAL OF BOTANY

PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY

MADRONO A WEST AMERICAN JOURNAL OF BOTANY

Second-class postage paid at Berkeley, California. Return requested. Established 1916. Individual subscription price $8.00 per year ($4.00 for students). Institutional subscription price $12.00 per year. Published quarterly in January, April, July, and October by the California Botanical Society, Inc., and issued from the office of Madrofio, Herbarium, Life Sciences Building, University of California, Berkeley, California. Orders for subscriptions, changes in address, and undelivered copies should be sent to the Corresponding Secretary, California Botanical Society, Depart- ment of Botany, University of California, Berkeley, California 94720. °

BOARD OF EDITORS CLASS OF:

1971—Marion OwnsBEy, Washington State University, Pullman Joun F. Davison, University of Nebraska, Lincoln

1972—Ira L. Wicctns, Stanford University, Stanford, California ReEeEp C. Rots, Harvard University, Cambridge, Massachusetts

1973—WattacE R. Ernst, Smithsonian Institution, Washington, D.C. Roy L. Taytor, University of British Columbia, Vancouver

1974—-KenTon L. CHAMBERS, Oregon State University, Corvallis EMLEN T. LitTELL, Simon Fraser University, Burnaby, British Columbia

1975—ArtTURO GOMEZ PompPa, Universidad Nacional Aut6noma de México Duncan M. Porter, Missouri Botanical Garden, St. Louis

1976—Dennis ANDERSON, Humboldt State College, Arcata, California KINGSLEY R. STERN, Chico State College, Chico, California

Editor — JoHn H. THomas Department of Biological Sciences, Stanford University, Stanford, California 94305

Business Manager and Treasurer — JuNE McCasKILy P.O. Box 23, Davis, California 95616

CALIFORNIA BOTANICAL SOCIETY, INC.

President: Lawrence R. Heckard, Department of Botany, University of Califor- nia, Berkeley. First Vice-President: Donald Kyhos, Department of Botany, Uni- versity of California, Davis. Second Vice-President: Dennis Anderson, Department of Botany, Humboldt State College, Arcata. Recording Secretary, John West, De- partment of Botany, University of California, Berkeley. Corresponding Secretary: Dennis R. Parnell, Department of Biological Science, California State College, Hay- ward. Treasurer: June McCaskill, Department of Botany, University of California, Davis.

The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Marion Cave, Department of Botany, University of California, Berkeley; the Editor of Madrofio; and three elected Council Mem- bers: Malcolm Nobs, Carnegie Institution of Washington, Stanford; Elizabeth McClintock, Department of Botany, California Academy of Sciences, San Francisco ; and Thomas Fuller, California Department of Agriculture, Sacramento.

EVIDENCE OF DIFFERENT ADAPTATIONS OF FLOWER COLOR VARIANTS OF ENCELIA FARINOSA (COMPOSITAE)

By Donatp W. KyHos

It has long been recognized that plants are extremely sensitive to their environments, particularly to moisture and edaphic factors. In natural undisturbed environments one of the most notable ways this remarkable sensitivity is revealed is by the general lack of hybrids be- tween sympatric species which in disturbed habitats do produce hybrids and their various derivatives. A most lucid account of this well known phenomenon is provided by Anderson (1949). In most instances such sympatric species differ by several to many externally visible features and many differ also by a number of less obvious aspects, including physiological attributes.

In such cases where a relatively large number of genetically deter- mined attributes differentiate ecologically distinct taxa, it seems likely that their differing ecologic adaptations will be correspondingly complex and hence difficult to analyze. On the other hand, with taxa that differ by one or at most a few genetically controlled features, the probability is much greater that their ecologic adaptations are simple and relatively easy to analyze.

Moreover, in the case of taxa that differ in many genetically based traits there appears to be little chance of obtaining evidence of the manner in which they differentiated. For example, whether geographic isolation was necessary for their divergence and their sympatry second- ary, or whether it is possible that in some instances ecological differen- tiation initially resulted from a very few, or even a single heritable difference which permitted the occupation of new, closely adjacent sites well within effective pollination range of the original population.

Thus it is apparent that there are at least two substantial advantages in seeking populations to study that are ecologically distinct, but which have few genetic differences. First, it is more likely that their ecologic adaptation can be discerned and successfully analyzed and second, they are rather more likely to provide evidence of their mode of origin.

If a relatively simple heritable difference can bring about an altered ecologic adaptation in natural environments, then it should be possible to find populations at this initial stage of ecologic divergence when they differ by one, or at most, a very few heritable attributes, as appears to be the case in Ricinus in Peru (Harland, 1946) and Spergula in Europe (New, 1958; 1959) and perhaps in Australian Eucalyptus (Barber and Jackson, 1957).

Encelta farinosa Gray also appears to provide an excellent opportunity to study populations at such an early, but very significant stage of

Maprono, Vol. 21, No. 2, pp. 49-112. November 9, 1971. 49

50 MADRONO [Vol. 21

divergence. Many populations of Encelia farinosa in the Sonoran Desert are polymorphic for a single conspicuous feature. In California, southern Nevada, Arizona, and adjacent Mexico, Encelia farinosa occurs as variety farinosa, in which the ray and disk florets are a bright yellow and it also occurs as E. farinosa var. phenicodonta, which differs only in that the disk florets are a rich brownish-purple. Populations may consist entirely of one or the other of these varieties, or may contain both inter- mixed. This seemingly trivial phenetic difference would not ordinarily lead one to suspect that it is an expression of a more fundamental bio- logical difference. However, an examination of what proves to be a rather remarkable distribution pattern of these two varieties of E. farinosa strongly suggests that this apparently simple flower color differ- ence has a high selective value itself, is part of a pleiotropic expression of genetic material which is adaptive in some other less obvious feature, or that this flower color character is genetically very closely linked with some other factor of high selective value that is not readily discernible. Distribution pattern.

Encelia farinosa is one of the dominant species of the Sonoran Desert, to which it is largely confined. It occurs in abundance from near the Cape region of Baja California, northward into the deserts of California, southern Nevada, southern Utah, Arizona, and Sonora, Mexico to Sina- loa (fig. 1). In the Cape region of Baja California, the southern-most populations of E. farinosa belong to var. radians, which occurs mainly in the tropical zone (Shreve and Wiggins, 1964). In the immediately adjacent deserts, radians is replaced by the very similar var. phenico- donta, which extends northward through the major portion of the Baja California peninsula to about San Felipe, Mexico. It is in this general region that the yellow disk flowered variety of E. farinosa farinosa ap- pears and begins to replace phenicodonta in a most unusual pattern. In this more northern area of the Sonoran Desert, populations consisting entirely or predominantly of var. phenicodonta are largely confined to major river valleys. The yellow flowered populations of E. farinosa occupy much of the remainder of the northern Sonoran Desert, and are marginally sympatric with the brown disk flowered populations.

The writer first noticed this most unusual distribution pattern when making an approximately west to east transect across the Colorado River, near Blythe, California. While travelling south-eastward along the road from Rice, California toward Blythe, only yellow flowered E.

Fic. 1. Distribution of Encelia farinosa. The dotted line outlines the limits of the Sonoran Desert in northwest Mexico and the adjacent United States as defined by Shreve and Wiggins (1964). The shading denotes areas where Encelia farinosa phenicodonta occurs either in high concentrations or to the exclusion of E. farinosa farinosa. Five small disjunct phenicodonta localities are shown at about twice their proportionate size for clarity. Encelia farinosa farinosa occupies much of the remain- ing Sonoran Desert that is unshaded, as well as the area to the immediate north, included in this map.

KYHOS: ENCELIA 51

1971]

%e.° e e eo cont ® *. e e e e

52 MADRONO [Vol. 21

farinosa populations were initially encountered until a point approxi- mately 26 miles from Blythe was reached (16 miles due west of the Colorado River). The next mile of /. farinosa populations included phenicodonta individuals at a frequency of 5 percent. In the next 6-mile interval approaching Blythe the frequency of phenicodonta individuals in the populations progressively increased to 38 percent, and upon ap- proaching to within 9.3 miles of Blythe the frequency of phenicodonta plants in the population gradually increased to 70 percent. Within Blythe and its environs, intensive agricultural practices have essentially elimi- nated E. farinosa, however, upon crossing the Colorado River into Ari- zona, EF. farinosa is once again abundant. Proceeding east from the Colo- rado River along U.S. Highway 60-70 for the first two miles only pheni- codonta individuals are encountered and with each mile eastward the frequency of this taxon continually declines, until at 13 to 14 miles east of the Colorado River, E. farinosa populations consist of less than 7 percent phenicodonta plants. Still farther eastward only yellow flow- ered populations are encountered.

The well defined cline revealed by this initial transect suggested that phenicodonta individuals would probably be concentrated in other areas of the Colorado River valley to the north and south of Blythe, Cali- fornia. Much of the Colorado River valley to the south of Blythe is under intensive cultivation, and still farther south the few areas near the Colorado River that are readily accessible lack E. farinosa. This is also the case on both the Arizona and California sides of the Colorado River at Yuma. Somewhat to the north of Yuma, as the Picacho Recre- ation Area is approached from U. S. Hwy. 80, the clinal trend is again repeated, although less dramatically. In the area 8.1 miles north of Hwy. 80 on the road to the Picacho area, var. phenicodonta occurs in a frequency of less than 1.0 percent. At 11.7 miles north of the above locality the frequency climbs to 9.0 percent, and at the edge of the Colorado River the frequency reaches 15.0 percent.

Approximately 30 miles to the southwest of Yuma, near the conflu- ence of the Rio Hardy and the Colorado River, in Mexico, a population sample of over 100 individuals includes only phenicodonta plants. Thirty-five miles to the south, near the mouth of the Rio Hardy, two closely adjacent samples, comprising 465 plants, average 90 percent phenicodonta individuals. Approximately 75 miles east of these three localities, along Mexico Hwy. 2, the frequency of phenicodonta drops to 4.7 percent and continues to decline still farther east from this point. Although the £. favinosa population samples in Mexico are few, they agree with the basic distribution pattern near Blythe. A sampling of E. farinosa along the Colorado River to the immediate north of Blythe, further confirms this clinal distribution pattern. In this area U. S. Hwy. 95 closely follows the course of the Colorado River for approximately 27 miles. An overall frequency of 84.8 percent phenicodonta individuals

1971] KYHOS: ENCELIA ye)

occurs in the 27 mile interval. North of this area Hwy. 95 takes a north- westward course away from the Colorado River and it is at this point that the frequency of pkenicodonta plants begins to decline and con- tinues to decrease rapidly as one travels farther away from the Colorado River. If on the other hand, populations are sampled in an eastward direction, it is found that the frequency of phenicodonta plants increases as the Colorado River is approached and that they reach their highest incidence of 67.0 percent adjacent to the river in the Earp, California- Parker, Arizona area and to the north. Still farther north, with sampling along the road leading to the Chemehuevi Valley Indian Reservation, the pattern is repeated again. As the Colorado River is approached the proportion of pkenicodonta in the populations progressively increases from zero (about 15 miles west of the river) to about 20 percent (within one or two hundred yards of the river). Twenty miles to the north in the Topock, Arizona area, a low incidence of 2.0 percent phenicodonta plants found at the river’s edge quickly drops to zero within less than 0.2 miles from the river. Another 35 miles to the north, where Nevada State Highway 77 crosses the Colorado River, the pattern repeats, with E. farinosa populations within a quarter mile of the river containing ap- proximately 20.0 percent phenicodonta individuals and with the fre- quency of these plants decreasing rapidly to zero at less than a mile from the river either in an eastward or westward direction.

In Arizona, other than along or relatively near the Colorado River valley, this writer has so far found only a single area where var. phenico- donta occurs in appreciable numbers. This locality is in south-central Arizona, in the Granite Reef Dam vicinity, about 10 miles northeast of Mesa, Arizona.

This site is most remarkable, since in a very small area it duplicates the pattern of distribution observed for the two varieties of E. farinosa in and around the much more extensive area of the Colorado River valley. Approaching this Granite Reef Dam locality from the south, one observes that within the interval of 2.2-0.4 miles from the margin of the canyon, the frequency of phenicodonta plants averages between 1.0 to 2.0 percent of the populations. In the interval between 0.4 miles south of the canyon and the canyon’s south edge there is a progressive increase in phenicodonta individuals to 11.0 percent and in the next 0.2 mile interval, as one begins to descend into the canyon toward the Salt River, the frequency sharply increases to 37.0 percent and then to 62.0 percent within the next 0.3 miles. An even higher incidence of 88.4 percent phenicodonta plants is encountered at the base of the north wall of the canyon, essentially on the upper banks of the river.

While the association of var. phenicodonta with water courses is a prominent feature of its regional, as well as its local distribution pattern, it is apparent that relatively local concentrations of the variety are also found in another kind of habitat. The most notable of these occurs at the west base of the Granite Mountains, 0.2 miles south of the inter-

54 MADRONO [Vol. 21

section of the road to Twentynine Palms with the road to Desert Center, California. This area is conspicuous because of the large sand dune that sprawls on the west slope of the Granite Mountains. Within and adjacent to a sandy wash that spills westward from these mountains, E. farinosa phenicodonta attains a frequency of 70.0 percent. Sampling to the immediate south of this locality reveals that this frequency rapidly drops to near zero within less than 5.0 miles, but again increases to about 17.0 percent in the populations at the southeast base of the Coxcomb Mountains. Continuing still farther south toward Desert Center this variety decreases to one percent or less of the populations. Approxi- mately 15 miles north of the Granite Mountains locality, a concentration of phenicodonta attaining a frequency of about 20.0 percent occurs in the Iron Mountains. Another 33 miles to the north, approximately a 20.0 percent concentration of phenicodonta is found on the lower south- east-facing slope of Clipper Mountain, immediately northwest of Danby, California. In Arizona along U. S. Hwy. 80 at the summit of Mohawk Pass, immediately east of the town of Mohawk, var. phenicodonta locally makes up 54.0 percent of the population. Proceeding either east or west from this pass in the Mohawk Mountains, one descends and a rapid decrease in the frequency of pkenicodonta plants occurs.

In Sonora, Mexico, where very little information on populations of E. farinosa is available, an examination of herbarium specimens suggests a most interesting geographic distribution of the two flower color forms of E. farinosa. Variety phenicodonta generally seems to occur abundantly only near the coast as far south as the latitude of Tiburon Island, and the yellow flowered populations appear to occupy largely the inland areas. Population samples are badly needed from this area of Mexico to establish if this suggested distribution pattern is correct.

Thus the data available from populations of E. farinosa indicate that phenicodonta is generally associated with major water courses or is largely confined to higher elevations of the California and Arizona deserts, and may be generally limited to the coastal area of the Sonoran Desert of Mexico.

INHERITANCE OF FLOWER COLOR

While a great deal yet remains to be learned about the inheritance of flower color in E. farinosa, some interesting observations from experi- mental crosses as well as natural populations shed some light on this important aspect. One of the first facts established was that when progeny are grown from F. farinosa farinosa and phenicodonta, where these taxa grow sympatrically in about equal proportions, both varieties can be obtained in the next generation from some of the plants belonging either to var. farinosa or phenicodonta. This progeny test clearly demon- strates that where the two varieties are sympatric they interbreed. This observation also suggests a simple inheritance for flower color. However,

1971] KYHOS: ENCELIA a5

an examination of natural populations reveals that there is the following additional complication. For while disk floret coloration as discussed above appears to be inherited in a relatively simple, perhaps single gene fashion, a survey of over 4000 individuals in natural populations shows a low incidence of individuals possessing brownish-purple anthers with corollas that are yellow or only slightly tinged with reddish-purple. However, this same sampling of plants has failed to reveal the reciprocal combination of yellow anthers and brownish-purple corollas in the disk florets. This observation seems to indicate that anther and corolla color are not determined by two independently segregating genetic factors, one for anther color and another for corolla color. Instead, it appears likely that one gentic factor produces yellow anthers and corollas, a second genetic factor produces brown anthers, with little or no effect on corolla color, and that a third factor produces brownish-purple anthers and corollas in the disk florets. This genetic scheme would explain the ab- sence of plants with brownish-purple disk corollas and yellow anthers.

A similar inheritance pattern is seen in natural intergeneric hybrids between E. farinosa and Geraea canescens. Eleven such hybrids have been reported to date and of these, two had entirely brownish-purple disks. A third hybrid had yellow disk corollas and brownish-purple anthers, whereas the remaining eight hybrids had entirely yellow floral parts (Kyhos, 1967). Geraza canescens invariably has corollas and anthers that range in color from yellow to yellowish-orange, therefore the brownish-purple disk coloration in the intergeneric hybrid had to be inherited from FE. farinosa in a dominant fashion. These observations re- veal that the brownish-purple pigmentation of the disk corollas and anthers is transmitted to the hybrid in the same manner as in progeny of E. farinosa. It has not yet been possible by controlled crosses, however, to produce a large enough progeny in E. farinosa to fully examine the mode of inheritance of the three flower color forms. This is largely due to the difficulty of germinating the fruits of E. farinosa and the reluc- tance of this species to flower under greenhouse conditions.

Interspecific hybrids experimentally produced between E. farinosa phenicodonta and the yellow disk flowered E. frutescens further sup- port an interpretation that flower color is probably inherited in a rela- tively simple way. These F, hybrids have all segregated into two classes: either they have had entirely yellow disk corollas and anthers, or these structures have been brownish-purple. Moreover, they have segregated in essentially equal frequencies with a Chi-square value conforming well to the 1:1 ratio expected in the test cross of a heterozygous domi- nant parent to a homozygous recessive one (see table 1).

Finally, controlled intraspecific crosses between individuals of E. farinosa farinosa and phenicodonta also indicate a relatively simple genetic basis for disk coloration, depending on which individuals are used as parents. So far in these crosses one of two things occurs. Either all the progeny are var. phenicodonta or the progeny segregate for the

56 MADRONO [Vol. 21

TABLE 1. SEGREGATION OF DISK COLORATION IN F,; HyBrRIDS BETWEEN ENCELIA FRUTESCENS AND ENCELIA FARINOSA PHENICODONTA

observed expected d d? Yellow 21 21 6) 0) disks xX? = 0.0 p 99 Brownish- 21 ZA 0 0)

purple disks

two varieties in a frequency that conforms well to a 1:1 ratio with the Chi-square test (table 2).

OBSERVATIONS ON POLLINATORS

Since disk floret color differentiates EL. farinosa farinosa from pheni- codonta, it was important to observe if natural pollinators distinguish between these two color forms and in so doing were perhaps responsible for the remarkable geographic distribution of these plant taxa. In line with this approach twelve populations were studied within and adjacent to the Colorado River valley, between Blythe and Needles, California. Six populations were in the area of sympatry of farinosa and phenico- donta and the remaining six were in areas occupied only by farinosa, between 10 to 14 miles removed from the Coloroda River. In all twelve populations it was found that pollen shed in both farinosa and phenico- donta begins about 9:00 a.m. and ceases in the early afternoon, with some day to day variation apparently depending on the weather. Pollen was shed somewhat earlier on warm sunny days than on cool cloudy days. Nevertheless, on any given day the two forms of E. farinosa always shed essentially synchronously, indicating that temporal factors appar- ently provide no reproductive isolation.

The remaining important aspect was whether different pollinators were attracted to the two flower color types. An examination of the six sympatric populations of farvinosa and phenicodonta demonstrated a variety of flower visiting insects, including diptera, hymenoptera, lepi- doptera, homoptera, and coleoptera. All of the species in each of these insect orders occurred on both varieties of E. farinosa. Morevover, they fly from individuals of each FE. farinosa variety in a random fashion, with individual insects showing no preference for one flower color. The most important pollinator in each of these populations proved to be a small beetle, Tanaops abdominalis Le Conte, of the Malachiidae, which occurred in a frequency over 10 times greater than all of the other insect species combined. This beetle species moved from flower head to flower head, efficiently pushing among the disk florets, inadvertently picking up pollen along the way and then flying quickly and quite accurately to the next flowering head of EF. farinosa, which might be either that of farinosa or phenicodonta.

1971] KYHOS: ENCELIA oF

TABLE 2. SEGREGATION OF DisK COLORATION IN PROGENY OF ENCELIA FARINOSA FARINOSA < ENCELIA FARINOSA PHENICODONTA

observed expected d d? Yellow 19 18 1 1 disks X2 = .111 p= 57/0 Brownish- purple disks 17 18 if 1 DISCUSSION

Encelia farinosa phenicodonta has been recognized as a form or variety of Encelia farinosa Gray, for approximately fifty years (Blake, 1913: Johnston, 1924). Botanists who have dealt with these taxa apparently considered them to be simply flower color variants that differ only in this seemingly trivial aspect (Abrams and Ferris, 1960; Blake, 1913; Johnston, 1924; Munz, 1959; Shreve and Wiggins, 1964). However, closer scrutiny provides compelling evidence that these taxa have a much deeper and most intriguing biological significance. The distribution pattern of these taxa is quite remarkable and seemingly can only be interpreted as resulting from natural selection. It surely cannot be fortui- tous that comparatively high frequencies of phenicodonta plants occur as a narrow, occasionally interrupted strand closely following the course of the Colorado River for approximately 260 miles north of the main body of this taxon (fig. 1). It seems even less likely that chance is a significant determinant in this remarkable distribution, when one notes that this 260 mile strand of pkenicodonta is completely included within the range of Encelia farinosa farinosa and that these two taxa are sym- patric along this entire 260 mile interval, and are known to interbreed with no evidence of reproductive isolation. In seeking an understanding of what selective forces may be operating to produce this striking dis- tribution pattern, we perhaps can gain insight from the observation that the pattern along the Colorado River valley is dramatically repeated on a much smaller scale at Granite Reef Dam in east-central Arizona, near the city of Mesa. Both in the Colorado River valley and the Granite Reef Dam area of Arizona the populations of Encelia farinosa with the greatest proportion of phenicodonta plants occur closest to the water in these drainage systems, and as one travels away from the water course the frequency of phenicodonta individuals progressively declines until individuals of var. farinosa entirely replace phenicodonta. These ob- servations suggest that in some manner water is important in determining the distribution of E. farinosa farinosa and phenicodonta. The apparent predominance of phenicodonta populations near the coast, with farinosa populations characterizing the more inland areas of Sonora, Mexico, again lends support to the idea that effective moisture is a crucial factor in the distribution of these two taxa. The credibility of this interpreta- tion is further enhanced when it is recalled that the comparatively small

58 MADRONO [Vol. 21

local concentrations of phenicodonta individuals, which occur away from water courses and large bodies of water, are found at higher elevations in the deserts of California and Arizona (e.g., at Mohawk Pass, Arizona, near Hwy. 80; and at Lobecks Pass near Hwy. 95, about 15 miles south of Needles, California; and in the Iron, Granite, Coxcomb Mts., and on Clipper Mt., all located in the Mohave Desert of California and its transition to the Colorado Desert to the south). These desert mountain ranges characteristically receive greater amounts of rainfall than the surrounding lower desert and thus in this regard would be similar to the habitats near water courses and large bodies of water.

It might be argued that the water course habitats, higher desert ele- vations, and coastal areas would provide cooler temperatures and thus temperature itself may be a critical factor in the distribution of these taxa. It is obvious that lower temperatures would tend to reduce tran- spiration and hence increase the amount of effective moisture. Thus it is important to separate the action of these two factors. An examination of maximum, minimum, and mean temperatures in the journal, Climato- logic Data, through the ranges of these two taxa reveals no correlation with their distribution. If water stress is physiologically crucial in determining where these two taxa grow, it remains obscure how this factor is invariably associated with a seemingly unrelated characteristic such as disk flower coloration, except perhaps through pleiotropy or an extremely close genetic linkage.

On the other hand, the possibility cannot as yet be totally ruled out that flower color is itself adaptive. For example, it might be imagined that a pollinator preferring brownish-purple disks in Encelia is limited to areas of greater moisture in the desert, however no such evidence has yet been obtained. In fact, field observations support quite the opposite conclusion, namely, that the effective pollinators do not dis- criminate between farinosa and phenicodonta and progeny tests of these two taxa where they are sympatric bear this out.

While the selective factors which produce the remarkable distri- bution pattern of favinosa and phenicodonta are as yet unknown, some insight into the possible origin of these very similar taxa can be gained from what is known about the genetic basis of their differences. The available evidence indicates that these taxa are based on comparatively simple genetic differences. Brownish-purple disk floret coloration ap- parently can be inherited in a dominant, single gene fashion. Individuals of phenicodonta when crossed to var. farinosa either have produced progeny all with brownish-purple disks or in other crosses the progeny have segregated for brownish-purple and yellow disks in essentially equal frequencies. This rather simple situation becomes slightly more com- plicated, since many natural populations include a low frequency of individuals possessing yellow disk corollas associated with brownish- purple anthers. However, even with this additional aspect the genetic difference between these taxa appears to be quite simple, with disk

1971] KYHOS: ENCELIA 59

TABLE 3. LOCALITIES OF POPULATION SAMPLES

After each locality the letter t followed by a number indicates how many Encelia farinosa farinosa individuals occurred in the sample, whereas a number preceded by the letter p indicates how many E&. farinosa phenicodonta individuals were in the sample.

Arizona. Near hwy 60-70, 2 mi E Colorado R., t-0, p-45; 1 mi farther E, t-10, p-240; 1 mi farther E, t-2, p-77; 1 mi farther E, t-17, p-122; 1 mi farther E, t-34, p-76; 1 mi farther E, t-33, p-251; 1 mi farther E, t-6, p-17; 1 mi farther E, t-30, p-14; 1 mi farther E, t-50, p-28; 1 mi farther E, t-28, p-3; 1 mi farther E, t-130, p-32; 1 mi farther E, t-157, p-16; 1 mi farther E, t-55, p-4, near hwy 80, 13.4 mi E jet with San Luis rd, t-96, p-4; 2 mi farther E, t-99, p-1, 1.7 mi farther E, t-337, p-6; 2.5 mi farther E, t-87, p-22, near hwy 80, 0.3 mi W Mohawk Pass summit, t-46, p-54; 1.1 mi E Mohawk Pass summit, t-170, p-30; hwy 80, 14 mi W ject with Theba rd, t-195, p-5; hwy 80, 15.9 mi W Buckeye, t-200, p-0; near hwy 80, 2.8 mi E jct with hwy 72, t-99, p-1; near hwy 80, 21 mi W jct with hwy 72, t-200, p-0; 9.3 mi N Wenden, t-200, p-0, rd to Dome, 6.3 mi N of hwy 80, t-94, p-11; 5.3 mi N of Tucson, t-200, p-O; 3-4 mi S Parker Dam, t-69, p-131; edge S rim Salt R. canyon at Granite Reef Dam, t-89; p-11; 0.8 mi farther S, t-98, p-2; 1.5-2 mi farther S, t-99, p-1; 2.2-2.7 mi farther S, t-152; p-3; 0.2 mi N, S rim Salt R. canyon, t-63, p-37; 0.3 mi farther N, t-38, p-62; N bank Salt R. at Granite Reef Dam, t-35, p-265.

CALIFORNIA. Hwy 60-70, 11.2 mi W Colorado R, t-110, p-24; near hwy 95, 8.4— 33.4 mi N jct with hwy 60-70, t-106, p-384; at 34.6 mi N, above jet, t-48, p-532, and one plant with a light purple disk; at 46.5 mi N, above jet, t-48, p-4; Vidal Jct., t-39, p-1, 1 mi E Vidal Jct., t-16, p-7; 1 mi farther E, t-16, p-4; 0.8 mi farther E, t-172, p-28; from Earp to 7.4 mi N, t-17, p-35; Parker to 5 mi S, t-45, p-135; Vidal Jct. to 4 mi N, t-92, p-8; 1.5 mi farther N, t-99, p-1, 1.7 mi farther N, t-200, p-0O; 1.2 mi farther N, t-290, p-10, 1 mi farther N, t-200, p-0, jct hwy 95 with rd to Chemehuevi Valley Indian Res., t-200, p-0; 4.8 mi farther E, t-100, p-0; 5.2 mi farther E, t-94, p-6; 4 mi farther E, t-49, p-1, 3.4 mi farther E, t-16, p-1, bank Colorado R. at Chemehuevi Indian Res., t-79, p-21; 4.3-5.5 mi W Colorado R. near hwy 66, t-100, p-0, W bank Colorado R. near hwy 66, t-98, p-2, 1.1 mi N Needles, t-99, p-1, S slope Clipper Mt. near Danby, t-182, p-38; hwy 95 at Lobecks Pass near jct with hwy 66, t-95, p-5, near rd from Blythe to Rice, 2 mi N Riverside Co. dump, t-30, p-70; 4.1 mi N, above, t-62, p-38, 3.5 mi farther N, t-190, p-10, 5.9 mi farther N, t-200, p-O; summit at S end Iron Mts., t-64, p-4; 0.4 mi farther W, t-26, p-0; 0.1 mi farther W, t-284, p-16; 0.5 mi N near summit Iron Mts., t-171, p-29; 0.5 mi S jct of rd to Desert Center and Twentynine Palms rd., t-44, p-173, 5.6 mi S of above, t-99, p-1; 8.3 mi farther S, t-83, p-17; 1 mi W of Desert Center, t-99, p-1; 0.1 mi N hwy 60-70, near rd to Twentynine Palms through Joshua Tree Nat. Mon., t-300, p-0; Morongo Wash, 2.6 mi N hwy 60-70, t-300, p-0; White Water Canyon, NW Palm Springs, t-400, p-2; 1 mi S, S border Anza-Borrego State Park near rd S2, t-200, p-0; Panamint Valley, 0.5 mi W Darwin Springs, t-200, p-0; 8.1 mi N hwy 80 near rd to Picacho Resort Area, t-107, p-1; 11.7 mi N, above, t-91, p-9; S bank Colorado R. at Picacho Resort Area, t-85, p-15.

Mexico. 4-5 km § San Felipe, Baja California, t-0, p-100; 180 km N San Felipe, Baja California, t-0, p-100; 122 km N San Felipe, Baja California, t-19, p-142; 115 km N San Felipe, Baja California, t-24, p-280; Mexican hwy 2, 50 mi E San Luis, Sonora, t-100, p-5; Mexican hwy 2, 106 mi E San Luis, Sonora, t-96, p-4; Mexican hwy 2, 16 mi S Sonoyta, Sonora, t-98, p-2; 12.7 mi W Caborca, Sonora, (22 2123)

Nevapa. Near hwy 77, 0.3 mi W of Colorado R., t-90, p-23; 5.9 mi W, above, 153-10

60 MADRONO [Vol. 21

corolla and anther coloration in the great majority of individuals match- ing one another. Despite the relatively simple genetic basis for disk floret coloration, this minor phenetic difference appears to be biologically very important, perhaps in itself or possibly it is inextricably associated with some factor that is crucial in determining where these taxa can survive. In this situation the sympatric origin of one taxon from the other appears quite feasible, with this simple genetic difference providing the means by which a new or different habitat could be occupied, resulting secondarily in allopatry of the two taxa in much of their ranges, rather than allopatry being a necessary prerequisite for the differentiation of these taxa.

The present distribution of E. farinosa farinosa and phenicodonta suggests that the latter was previously more widespread, perhaps within fairly recent times. Phenicodonta in the northern portions of the Sonoran Desert not only occurs in high frequency, as a more or less continuous strand, along the Colorado River valley, but it is found also as isolated populations, some of which are quite small and far removed from the great concentrations of phenicodonta. These scattered, well isolated pop- ulations of phenicodonta may represent remnants of a once more extensive distribution. The converse situation of scattered, well iso- lated populations of farinosa in areas populated predominantly by phenicodonta is unknown. This suggests that perhaps during a period such as the hypsithermal, when comparatively warm, moist climates may have extended farther north than today (Martin, 1963), phenico- donta may also have occurred farther north than at present. With the decline of the hypsithermal, phenicodonta may have retreated to the south, leaving small isolated populations surviving in favorable local sites as evidence of this former distribution, while farinosa came to occupy most of the remainder of the Sonoran Desert, now untenable for phenicodonta.

The relationships of Encelia farinosa farinosa and phenicodonta some- what parallel those described for Linanthus parryae by Epling, Lewis, and Ball (1960). In both Linanthus and Encelia different flower colors, which have a relatively simple inheritance, seem to be indicative of more deep-seated biological differences between individuals which ap- parently interbreed freely. In each of these cases, the importance of these distinct flower color differences as indicators of a crucial biological dif- ference is revealed by the local and regional distribution patterns of these flower color types. The distribution patterns in Linanthus and Encelia appear to be the result of natural selection, but in neither case is the nature of the selective forces known. However, the relatively simple genetic differences of the individuals possessing these distinct adaptations offer hope that their adaptations are commensurately simple and hence ultimately can be understood.

1971] KYHOS: ENCELIA 61

ACKNOWLEDGMENTS

The writer is indebted to H. B. Leech of the California Academy of Sciences, who kindly identified the beetle, Tanaops abdominalis Le Conte, which pollinates Encelia farinosa. I am also grateful to Albert Johnson, Spencer Smith-White, and Keith Jones who generously pro- vided data on several population samples in Mexico. Finally, the sup- port for a portion of this work by National Science Foundation grant GB 6098 is gratefully acknowledged.

SUMMARY

An unusual geographic distribution of Encelia farinosa farinosa and E. farinosa phenicodonta is presented as evidence that these taxa have a much greater biological significance than mere flower color variants. Crossing experiments indicate that the flower color differences of these taxa can be inherited in a single gene, dominant fashion. The association between flower color differences and the survival of these taxa in par- ticular geographic areas remains unresolved.

LITERATURE CITED

ABRAMS, L., and Ferris, R. S. 1960. Illustrated flora of the Pacific States. Stanford Univ. Press.

ANDERSON, E. 1949. Introgressive hybridization. Wiley and Sons., New York.

BarBer, H. N., and Jackson, W. D. 1957. Natural selection in action in Eucalyptus. Nature 179:1267-1269.

Brake, S. F. 1913. A revision of Encelia and some related genera. Proc. Amer. Acad. Arts 49:362-641.

EpLinc, C., Lewis, H., and Barr, F. M. 1960. The breeding group and seed storage: a study in population dynamics. Evolution 14:238-255.

Har.anp, S. C. 1946. An alteration in gene frequency in Ricinus communis L. due to climatic conditions. Heredity 1:121-125.

Jounston, I. M. 1924. Expedition of the California Academy of Sciences to the Gulf of California in 1921. The Botany (The Vascular Plants). Proc. Calif. Acad. Sci. IV. 12:951-1218.

Kyuos, D. W. 1967. Natural hybridization between Encelia and Geraea (Com- positae) and some related experimental investigations. Madrono 19:33-43. Martin, P. S. 1963. The Last 10,000 years. A fossil pollen record of the American

Southwest. Univ. Arizona Press, Tucson.

Muwz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.

New, J. K. 1958. A population study of Spergula arvensis 1. Ann. Bot. (London) 22:457-477.

—-—. 1959. A population study of Spergula arvensis 2. Ann. Bot. (London) Z3223-33%

SHREVE, F., and Wiccrns, I. L. 1964. Vegetation and flora of the Sonoran Desert. Vol. 2. Stanford Univ. Press.

A NEW MYCOCALICIUM ON SCARRED SEQUOIA IN CALIFORNIA

LEE BONAR Department of Botany, University of California, Berkeley 94720

Observations and collections have been made through many years of a discomycetous fungus that grows as a mat on an exudate from the surface of exposed heartwood of living specimens of the big tree, Se- quoiadendron giganteum (Lindl.) Buchh., and the coast redwood, Se- quota sempervirens (Lamb.) Endl., in California. The fungus is never found on unscarred trees. It is present on the charred surface of burns and rarely on other large scars where there has been a flow of exudate from the wood.

The exudate flows down over the charred surface in spreading dark sheets or strands. It is watery in the fresh portion, becoming soft and sticky, then hard and brittle, lustrous Hessian Brown (R) to black on exposure and drying. The dried portion sometimes forms sheets up to 0.5 cm thick or balls up to 3 cm in diameter. In one instance the exudate streamed down over the burn for 5 m with the fungus growing over the lower 3 m of the area. The exudate usually issues from the more recently formed wood adjacent to the callus and not on the deeper parts of the burns.

Limited chemical examination of the dried exudate indicated that it is a very complex mixture of organic compounds, mostly water soluble, and very different from the resinous exudates from conifers such as pines and firs.

Surveys of fire-scarred specimens of the big tree were made in three areas:

Exudate Exudate and and fungus fungus present absent Sequoia National Park (Giant Forest and Swanee Grove) 145 5

Yosemite National Park (Mariposa Grove) 35 5 Calaveras State Park (Calaveras Grove) 28 8 Total ZOS=s 8

Many fire-scarred trees of Pinus, Abies, and Libocedrus were examined in each of the above areas but no growth of the fungus was found.

The fungus has been found less often and less profuse in growth on the coast redwood than on the big tree of the Sierra Nevada. Flow of exudate onto scarred surfaces of the trees was found to be much more limited and growth of the fungus correspondingly less. For example, a survey of 63 fire-scarred trees in Big Basin Redwoods State Park, Santa Cruz Co., California, showed exudate present on 9, visible growth of the fungus on 3 with 1 showing mature apothecia, while 60 of the trees

62

1971] DONAR: MYCOCALICIUM 63

Fics. 1-4. Mycocalicium sequoiae: 1, median section of apothecium (mazaedium lost in processing) ; 2, median section of lobulate apothecial head, & 82; 3, apothecia dissected from pseudostroma; early stages right, proliferating secondary growth left, xX 6; 4, microtome section of pseudostroma showing interlaced hyphae, x 82.

lacked any growth of the fungus. A search in the Cazadero Creek area in Sonoma Co. resulted in finding one tree with exudate and a growth of the fungus 5 & 3 cm in extent.

Approximately 100 fire-scarred trees were examined in Humboldt State Park, Humboldt Co., especially in the vicinity of Weott and Dyerville, and limited growth was found on 3 trees. Two of these occur- rences were not on fire scars: scanty growth without fruting, forming a line of growth at the outer margin of the heartwood, where some exu- date had appeared, was found on the cut end of a log from a recently felled tree. A second tree, at the roadside, had received a relatively large cut to remove the buttress next to the road. Quite copious exudate flowed from the cut and the fungus had formed a matted growth up to 3 cm thick; this showed some mature apothecia.

64 MADRONO [Vol. 21

Besides scarcity of the exudate, another factor limiting growth of the fungus on the redwood is the growth of superficial molds over the sur- face of the discomycete mat. The molds frequently form floccose growths over the surface of the mat and apparently inhibit growth and fruiting of the discomycete. Such mold growths were seen in only a few instances on the fungus on the big tree. Growth of such superficial molds is un- doubtedly favored by the relatively higher humidity surrounding those fire scars near the ground and commonly in deep shade in the fog-ridden belt occupied by the redwood. Corresponding sites on fire scars of big trees are surrounded through the long dry Sierran summer by less favor- able environment for surface mold growth.

DESCRIPTION OF THE FUNGUS

Vegetative growth of the fungus varies from a floccose subiculum (very rare) to a mat-like pseudostroma (Vainio, 1890). The pseudo- stroma is resupinate and adherent to the charred surface. In extent of growth it varies greatly with moisture present. Examples were found where the pseudostromata covered a few cm*, while some could be measured in m?. The maximum development observed, on an extensive burn on a very large tree in Swanee Creek Grove, Sequoia National Park, measured 6 m at the ground perimeter and spread upward over the burn 0.5 to 1.5 m with tongues of growth extending upward for 2.4 m.

The pseudostroma is commonly 0.5 to 2 cm thick but may be up to 4.5 cm. It is composed of interlaced hyphae (fig. 4) which in younger growth near the edge are abundantly encrusted with crystals of calcium oxalate. The hyphae vary greatly in form and pigmentation. Those of younger portions appear hyaline under the microscope with thickened gelantinized walls. The walls do not stain—hence the spaced arrange- ment in the figure. In older portions the hyphae are mostly free of crystals, become amber to deep brown in color, and are frequently made up of torulose cells. The hyphae are 7-15 » in diameter, with narrow lumina.

Stipitate apothecia develop in large numbers within the pseudostroma and grow to project above the surface of the mat (fig. 5). Stipes first appear as compact bundles of interlaced hyphae. As these elongate toward the surface of the pseudostroma, the outer cells of the cylindrical structure become thick-walled and dark in color. The stipes emerge through the surface of the pseudostroma as minute blackish columns with acute tips, giving the surface a distinctly setose character. There may be as many as 100 of these per cm. The stipes are 5-6 (4-8) mm long. Many remain as sterile pointed structures. In others, there is a branching of the hyphae within the tip and development of a palisade of hyphal elements to form the hymenium. The outer layer of blackened cells is reflexed to form the exciple (fig. 1), which surrounds a plane to convex hymenium. The surface of the hymenium is Chrysolite Green

1971] BONAR: MYCOCALICIUM 65

Fics. 5-7. Mycocalicium sequoiae; 5, surface view of apothecia in nature; 6 ma- ture apothecia from culture on wood block, vegetative growth scanty and floccose; 7, side view of apothecia in position in pseudostroma, x 15.

(R) and is farinose when dry due to the numerous green amorphous crystals attached to the surface. The apothecial heads are 0.5-1.5 mm in diameter. Stipes are frequently branched at various levels. Branching most commonly occurs near the tip, resulting in development of 2-20 contiguous convexities forming a lobulate head on a common stipe (fig.

2). Branching may occur at lower levels, each branch forming an apothe- cial head (fig. 7).

66 MADRONO [Vol. 21

The stipitate apothecia are frequently found completely buried in the pseudostroma due to continued surface growth of the pseudostroma. These may show a second stipe developed as a proliferation through the hymenium of the first (fig. 3) or, more commonly, new stipes develop and grow through to the new surface.

Asci appear first near the center of the hymenial area and successively toward the margin as the surface expands and becomes more convex. The asci are inoperculate, cylindric-clavate, short stipitate, 50-70 6-10 pw, 8-spored, with the apical portion thick-walled. The walls of the asci stain blue with iodine. The spores are 1-celled, smooth, brown, uniseriate to biseriate in the ascus and 6-8 * 4-5 yw. Paraphyses are abundant, branched, colorless, with swollen tips 3—5 » in diameter.

The walls and stipes of the asci gelatinize early, leaving the ascospores in linear groups of eight embedded in a gelatinous matrix and then ex- truded to the surface of the disk to form a mazaedium (Acharius, 1817). Acharius defined mazaedium as the superficial amorphous layer over the surface of the hymenium. Smith (1921), Dennis (1960), and Ainsworth (1961) define it as a type of fruit body or apothecium.

The production of asci is long-continued, new ones forming in areas where earlier ones have gelatinized and have been extruded to the surface.

No spermogonia were found as are described for some species of Calicium and related genera.

CULTURAL STUDIES

Germination tests showed a high percentage of viability in spores of material that had been held in a dry condition for as long as a year’s time. Single ascospore cultures were grown on a number of common types of agar media and on extracts of Sequoza wood in agar. Maximum growth was a mycelial colony 2-5 mm in diameter at the end of 1 month at room temperature.

Blocks of heartwood of Sequoiadendron giganteum 15 cm long by 2.5 cm diameter and found by microscopic examination to be free of any fungus hyphae were soaked in water and sterilized by autoclaving in glass culture vessels with water to keep the lower ends of the blocks moist. Young colonies from single ascospores were placed on the end of the blocks. Such cultures were held at 10°C in dark, at room temperature in light and in dark, and in a daylight chamber at outdoor temperature. Growth occurred under all these conditions. It varied from light-brown floccose cushions to a delicate scarcely visible subiculum on the surface of portions of the blocks, the growth becoming darker with age. Least growth was in the chambers held at 10°C. Best growth was at outdoor temperature. Cultures held at room temperature showed an intermediate rate of growth. No marked difference was observed in vegetative growth in light and in darkness. No conidia were observed in any of the cultures.

Initial development of stipitate apothecia appeared in cultures in the outdoor chamber at the end of 4 months and these matured by the end

1971] BONAR: MYCOCALICIUM 67

of 5 to 6 months (fig. 6). Cultures held at room temperature did not develop apothecia in either light or dark. Those held at 10°C for 5 months were sterile. Part of these were then moved to the outdoor cham- ber and apothecia developed after 3 additional months. Apothecia de- veloped in culture chambers showed typical asci and spores. Vegetative growth was not the compact pseudostroma found in nature but at most a loose cottony felt (fig. 6).

MYycELIUM IN Woop

Blocks of heartwood up to 5 cm thick were cut out from under burned areas in standing trees where the surface was covered by an extensive growth of the fungus. The wood appeared to be sound with no macro- scopic evidence of decay. Microscopic examination of this wood showed occasional hyphae in the lumina of the wood cells, but there was no evidence of any appreciable destruction of the cell walls. The blocks of wood used for culturing the fungus were split and examined at intervals up to 1 year from the time the cultures were started, having been main- tained in a moist condition. By the end of 3 months from the time of inoculation, hyphae were found within the cells throughout the length of the blocks. The hyphae ramified through the wood cells both longi- tudinally and transversely. The transverse hyphae commonly passed through pits and occasionally penetrated directly through the walls. Blocks that had been subjected to the action of the fungus for 1 year showed some discoloration, but no other evidence of change. Micro- scopic examination of these showed only slight corrosion of the cell walls. The evidence indicates that this fungus is capable of very slow digestion of wood when compared with the rate of action of other common wood- rotting fungi.

Field studies indicate that the fungus grows primarily on the exudate from the wood, this providing both the water and food material. Wood underlying areas bearing a heavy growth of the surface pseudostroma— which is certainly several years old—is sound in appearance and shows only very limited development of hyphae in the wood tissues. Kimmey and Lytle (1952) concur, saying, “Sterile black fructifications of an ascomycete were occasionally found growing in association with exudate from rift cracks in exposed heartwood but were not associated with decay.”

CLASSIFICATION

The characters of the apothecium place this fungus in the Caliciaceae. This family is usually classified under the lichens and its members usu- ally consist of a fungus-algal complex. Various authors have, however, described species under this family as lacking any algal component. They have frequently placed such species in the same genera with typical lichen species. Vainio (1890) established the genus Mycocalicium for species lacking gonidia and having one-celled, brown ascospores. Reinke

68 MADRONO [Vol. 21

(1895) proposed the Protocaliciaceae as a sub-family under the Cali- ciaceae for those organisms that were non-lichen in nature and cited Mycocalicium and Mycoconiocybe as generic names for non-lichen members of the genera Calicium and Coniocybe. Vainio (1927) lists some seven genera of gonidia-free organisms under the Caliciaceae and gives descriptions for five species of Mycocalicium.

Various other workers have not accepted these proposals. Rehm (1896) lists seven genera and numerous species of this group of Dis- comycetes that do not have algal components. He states that the ma- jority of the species in the family are true lichens and no generic distinc- tion is made between the lichen and non-lichen members. He lists Mycocalicium as a synonym under Calicium. Schneider (1897) used the name Mycocalicium for typical lichens with one-celled ascospores as a distinction from Calicium with two-celled ascospores. The name was used in the same sense by Nearing (1962). Keissler (1938) lists all the gonidia-free segregates as synonyms under the regular lichen genera having gonidia, and discusses the difficulty of being able to know for certain that the lack of gonidia may not be merely a fortuitous circum- stance. Arnaud (1931) presented the Caliciaceae as a complex of diverse elements drawn from the lichens, Sphaeriaceae, Perisporiaceae, Hysteria- ceae, etc. It is very difficult to characterize clearly such a mixture. I prefer to consider the family as a more restricted group.

Clements and Shear (1931) divided the genera of the Caliciaceae into two groups: one saprophytic and non-lichen, the other forming typical lichen thalli with algae.

Studies, including the complete life cycle in pure culture, have proved that our fungus is not a lichen. The form of the apothecia, the manner of maturation, and the characteristics of the ascospores agree with those of the genus Mycocalicium as established by Vainio. Vegetative develop- ment and habitat are highly distinctive and lead to the conclusion that this is an underscribed species of Mycocalicium.

Mycocalicium sequoiae Bonar, sp. nov. Pars vegetativa corallina uvidaque ex subiculo hyphae intertextae vel ex pseudostromate expanso crassitudine usque ad 4.5 cm constans, in sinectute atrata rimosaque; apothecia super superficium extendentia; stipes teres, simplex vel ramosus, 4.8 mm longus; apice acuto demum capita turbinata diam. 0.5-1.5 mm formante; apothecia primaria demum pseudostromate obvo- luta inde evolvent plura; asci inoperculati, cylindro-clavati, 8-spori, evanescentes, 50-70 * 6-10 yp, parietes iodino tingentes; gelatina sporaeque mazaedium in sicco crystalla irridia obtectae formantes; ascosporae 1-cellulae, ellipsoideae, fuscae, 6-8 4—5 yp, in asco uni-vel biseriatae; paraphyses ramosae; sporae asexuales deficientes.

Habitat. On exudate from exposed heartwood of living sequoias in California.

Holotype. On Sequoiadendron giganteum, Crescent Meadow, Sequoia

1971] BONAR: MYCOCALICIUM 69

National Park, Tulare C.o, California, Lee Bonar (UC 1403569-holo- type), July 1, 1935.

Collections studied. In addition to the holotype, 12 collections on S. giganteum from Calaveras, Mariposa, and Tulare counties, and 4 collections on S. sempervirens from Humboldt, Sonoma and Santa Cruz counties (all UC) were studied.

I am indebted to various persons for collections and aid in this study, especially to Doris Brenneman for help in the critical laboratory studies, to Rimo Bacigalupi for the Latin description, and to Victor Duran and A. A. Blaker for photographs.

LITERATURE CITED

AcHartius, E. 1817. Om de cryptogamiske vexter, som komma under nann af Calicioidea III. Kongl. Vetensk. Akad. Handl. 1817:220-244.

AINSworTH, G. C., and G. E. Bispy. 1961. Dictionary of the fungi. Kew, England.

ARNAUD, G. 1931. Les Asterinees V. Etude sur les champignons parasites; Caliciacees, Hemispheriacees, etc. Ann. Epiphyt. 16:235-302.

CLEMENTS, F. C., and C. L. SHear. 1931. The genera of the fungi. New York.

Dennis, R. W. G. 1960. British cup fungi. London.

KEISSLER, K. von. 1938. Die Flechten: Caliciaceae. In Rab., Krypt. Fl. Deutschl., Oesterr. u.d. Schweiz. 9(1, 2) :520-527.

KimMeEy, J. W., and P. C. Lyte. 1952. Fungi associated with cull in redwood. Forest Sci. 1:104-110.

NEARING, G. G. 1962. The lichen book. Ashton, Maryland.

ReuM, H. 1896. Ascomyceteen, Hysteriaceen und Discomyceteen. Jn Rab., Krypt. Fl. Deutschl., Oesterr. u. d. Schweiz. 1(3) :382-414.

SCHNEIDER, A. 1897. A textbook of general lichenology. Binghamton, New York.

SmiTH, A. 1921. Lichens. Cambridge.

REINKE, I. 1895. Abhandlungen tuber Flechten. Jahrb. Wiss. Bot. 28:39-150.

Vartnio, E. A. 1890. Etude sur le classification naturelle et la morphologie des lichens du Bresil. Act. Soc. Fauna FI. Fenn. 7(2) :1-256.

. 1927. Lichenographia Fennica. III. Coniocarpae. Act. Soc. Fauna FI. Fenn.

57:1-138.

NOTES AND NEWS

WALLACE Roy Ernst, 1928-1971.—Wallace Ernst, Curator of Botany in the Smithsonian Institution died of cancer in Washington, D.C., on October 8th, 1971, after an illness of about nine months. In addition to his Smithsonian curatorship, he held a professorship in absentia at the University of Kansas, and was for a number of years a valuable member of the editorial board of Madrono. Dr. Ernst was a recognized authority in the systematics of the Papaveraceae and an accom- plished worker in the field of floral morphology. He received his first two degrees at the University of California at Los Angeles and his doctorate at Stanford Uni- versity. An appreciation and biography will appear in Madrono in the near future.

—J.H.T.

THE GALIUM ANGUSTIFOLIUM COMPLEX (RUBIACEAE) OF CALIFORNIA AND BAJA CALIFORNIA

LAURAMAY T. DEMPSTER AND G. LEDYARD STEBBINS

Department of Botany, University of California, Berkeley 94720 Department of Genetics, University of California, Davis 95616

INTRODUCTION

There are three groups of dioecious Galium species endemic to western North America, of which the G. angustifolium complex is the smallest, both in number of species and in its range. The other two were treated in earlier papers, as follows: the G. multiflorum complex of western United States, Sonora and Baja California (Ehrendorfer, 1956; 1961; Dempster, 1959; Dempster and Ehrendorfer, 1965); the fleshy-fruited polyploid complex of California, Oregon and Baja California (Dempster, 1958; 1962; Dempster and Stebbins, 1965; 1968). The imperfectly dioecious, or polygamous, group exemplified by G. parishii Hilend & Howell and G. wrighti Gray has not been comprehensively dealt with, although its relationship to the G. multiflorum group is obviously close and, quite likely, ancestral.

All four groups have four-leaved whorls, with the exception of G. hardhamae Demp. of the fleshy-fruited group, which has six leaves per whorl. The G. angustifolium complex, subject to the present paper, has important characters in common with the G. multiflorum group and the polygamous G. parishii group, notably the long straight specialized fruit hairs and the three-nerved leaves. Resemblance to the fleshy-fruited group is less marked, but is apparent in the rather extraordinary habit of reaching upward with greatly elongated stems which later become the slender woody scaffolds for subsequent herbaceous growth. Much of G. angustifolium Nutt. of the present group, and especially G. nuttalli Gray of the fleshy-fruited group exemplify this character.

The distribution of the present group, unlike that of the G. multi- florum complex, is not archipelagic, but resembles more that of the fleshy-fruited group in that it is largely continuous. Its occurrence (fig. 1) is chiefly in the southern coastal ranges, from the Sierra San Pedro Martir of Baja California, northward into the Santa Lucia and Gabilan ranges of Monterey and San Benito counties. In Kern Co. it also occurs farther inland in the Tehachapi, Greenhorn, and Piute ranges, and at the southern end of the Sierra Nevada west of the crest. In the Mohave Desert it has been collected as far east as the Providence Mountains.

The G. angustifolium complex consists of one narrowly endemic, dip- loid, uniform species, G. jepsonii, one widely distributed polymorphic species on three ploidy levels, G. angustifolium, and one local hexaploid species, G. johnstonii, which apparently originated by hybridization from the other two. Galium johnstonii expresses its hybrid origin in

70

1971] DEMPSTER & STEBBINS: GALIUM 71

SAN BERNARDINO

jepsonii

johnstonii

[0]

NG. subse fOliosum

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[¥)

NG. subs ONYCENSe

NG. subsp NUGIiCcaule

ANG subs Jacinticum

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NG. subsp Gracillimum

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[vy =)

Gg. ubsp OOTregodense

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NG. ubsp Gabrielense

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O®# O0G@HOP D

o =)

G. ourep ANG USTIFOli UM

Fic. 1. Map showing entire distribution of the G. angustifolium complex. Num- bers indicate actual chromosome counts divided by 11.

great variability within small populations, but seems to be genetically isolated from both parental species and is not taxonomically subdivisible. Galium angustifolum, on the other hand, is divisible into subspecies, of which the number recognized is limited chiefly by practical considera- tions. Unfortunately the diploids are not clearly separated from the tetraploids with respect to morphology and geography. Consequently we must content ourselves with describing as separate subspecies several of the forms which are morphologically uniform and geographically restricted, leaving the large remainder, both diploid and _ tetraploid, under ssp. angustifolium. Throughout this paper, all subspecies men- tioned should be understood as belonging to G. angustifolium.

Were it not for the existence of the intermediate species G. johnstoni, there would scarcely be adequate reason to include G. jepsoni in the G. angustifolium complex. The concentration of leaves near the base of the plant, the truly campanulate corollas, and the short falcate ovary- and fruit-hairs all set G. jepsoni sharply apart. There is furthermore

MADRONO [Vol. 21

Fic. 2. Representative chromosome figures xX 1700: A, G. jepsoniz, meiotic, n = 11 (4120); B, G. johnstonii, mitotic, 2n = 66 (4118); C-J, G. angustifolium; C, ssp. angustifolium, meiotic, n = 11 (4128); D, ssp. angustifolium, mitotic, 2n = 44 (4067); E, ssp. foliosum, mitotic, 2n = 22 (4083); F, ssp. onycense, mitotic, 2n = 22 (4207); G, ssp. gracillimum, mitotic, 2n = 22 (4174); H, ssp. jacinticum, meiotic, 1 pole, n = 33 (4129); I, ssp. nudicaule, mitotic, 2n = 22 (4147); J, ssp. gabrielense, mitotic, 2n = 44 (4295).

no evidence that genes from G. jepsonii are present in any subspecies of G. angustifolium, nor that hybridization between the two species ever occurs on the diploid level. It is possible that G. johnstonii originated only once, perhaps from unreduced gametes of G. jepsoni and tetraploid G.. angustifolium.

Any definition of the complex is therefore brief: plants perennial, dioecious, with 4 leaves to a node (i.e., 2 leaves and 2 stipular append- ages, looking exactly alike); the generally narrow leaves obscurely 3- nerved, the marginal hairs spreading or apically directed; fruits with specialized hairs which are not uncinate.

The present group has gone a little farther with polyploidy than has the G. multiflorum complex, although not nearly so far as the fleshy- fruited group (Dempster and Stebbins, 1965; 1968). Of the species and subspecies here recognized, 5 are diploid, 1 tetraploid, 1 indistinguish- ably diploid and tetraploid, 2 hexaploid, and 1 unknown (fig. 2, table 1). The basic chromosome number is 11. The geographically isolated sub- species foliosum and onycense are both diploid. No comparable degree of isolation occurs within the fleshy-fruited group, although it is usual in the G. multiflorum complex.

The center of evolutionary activity appears to have been in the San Gabriel, San Bernardino, and San Jacinto Mountains, where the great

1971] DEMPSTER & STEBBINS: GALIUM 73

TABLE 1. CHROMOSOME COUNTS IN THE GALIUM ANGUSTIFOLIUM COMPLEX

(An asterisk after a collection number means that the count was about the num- ber reported. All counts are from California, except the first two listed.)

G. angustifolium ssp. angustifolium. 2n = 22. Baja California. Santo Tomas, 4229; Punto Banda, Hardham & Dempster 17,010*. Riverside Co.: 1890*; 4128; 4131; 4134; 4177; 4178; 4239; 4241; 4245. San Bernardino Co.: 4126; 4145*; 4149*, San Diego Co.: 4179; 4181* ;4182* ; 4183; 4330; Bacigalupi 8282; Bacagalupi 8288; Bacigalupi 8371*. 2n = 44, Kern Co.: 4208*; 4218. Los Angeles Co.: 4114*; 4115*; 4116; 4117; 4169*; 4172*; 4190; 4191*; 4216; 4369*; 4731. Monterey Co.: 4067; 4159; 4161*; 4193. Orange Co.: 4352*; 4354*. Riverside Co.: 4137*; 4176* ; 4249; 4258; 4276. San Benito Co.: 4194*. San Bernardino Co.: 4141*; 4142; 4152; 4154*,. San Diego Co.: 4189*; 4333; 4336*; 4340* ; 4341*; 4347*; Hardham & Dempster 17,013. San Luis Obispo Co.: Stebbins s. n.*. Santa Barbara Co.: 4163* ; 4164* ; 4166* ; 4167. Ventura Co.: 4111; 4113*; 4168*.

G. angustifolium ssp. foliosum. 2n = 22. Santa Barbara Co.: 4083.

G. angustifolium ssp. gabrielense. 2n = 44. Los Angeles Co.: 4295. San Bernar- dino Co.: 4294*,

G. angustifolium ssp. gracilimum, 2n = 22. San Bernardino Co.: 4173; 4174.

G. angustifolium ssp. jacinticum. 2n = 66. Riverside Co.: 4129; 4130; 4244*; 4246* ,

G. angustifolium ssp. nudicaule. 2n = 22. Los Angeles Co.: 4123*; 4236. San Bernardino Co.: 4147 ; 4150.

G. angustifolium ssp. onycense. 2n = 22. Kern Co.: 4206* ; 4207.

G. jepsonii. 2n = 22. Los Angeles Co.: 4120; 4122. San Bernardino Co.: 4143.

G. johnstonii. 2n = 66. Los Angeles Co.: 4118, 4121.

majority of forms are to be found (fig. 3). Tetraploidy probably oc- curred repeatedly, involving ssp. nudicaule at least once to produce ssp. gabrielense, and perhaps once again in the production ultimately of hexaploid ssp. jacinticum. Other tetraploids, probably not involving either G. jepsoniu or ssp. nudicaule, produced the taller forms of ssp. angustifolium which spread northward and westward, remaining region- ally sympatric with the diploid form in San Diego Co. and the San Jacinto and San Bernardino Mountains, and occupying alone the north- ern and western remainder of the range, including the lower altitudes of the San Gabriel Mountains. The desert diploid ssp. gracillimum probably originated from the tall southern diploid by selection for the desert habitat. Subspecies onycense, on the other hand, is probably much more ancient in its isolation, and had not obviously contributed to any poly- ploid. The diploid island race ssp. foliosum is probably also very ancient, and may or may not have been involved in the production of the main- land tetraploids.

Again, in this group, for the third time, we find hispid polyploids with no visible hispid diploid progenitors. In the G. multiflorum complex (Dempster and Ehrendorfer, 1965) we observed two very hispid tetra- ploids, G. hilendiae Dempster & Ehrend. and G. munzii Hilend & Howell, the former related to diploid G. multiflorum Kell. and the latter to diploid G. magnifolium Dempster, both lacking any obvious, ade- quately hispid diploid ancestor. In the fleshy-fruited group (Dempster

74 MADRONO [Vol. 21

SAN BERNARDINO

if

Dl : \ if oO < ORANGE RINE RST DiE

=G.ang.nudicaule “Ss =G.ang. jacinticum PAK =G.ang. gracillimum =Gang. gabrielense

Fic. 3. Map showing distribution in greater detail, particularly in the San Gabriel, San Bernardino, and San Jacinto mountains. Numbers indicate actual chromosome counts divided by 11.

and Stebbins, 1965; 1968), the hispidity of some of G. bolanderi Gray and G. andrewsi Gray ssp. gatense (Dempster) Dempster & Stebbins cannot be adequately explained by assuming hybridization between existing diploids. In the present group, the hispid tetraploid forms of ssp. angustifolium, as well as the hispid tetraploid ssp. gabrielense, can- not be explained with reference to existing diploids. One may postulate the extinction of one hairy diploid ancestor, but it strains credulity to suppose that only the hairy diploids have become extinct in all three of our dioecious West American species groups. One is left to wonder whether some additive genetic mechanism may not be at work to cause greater hairiness in some polyploids than was expressed in their diploid ancestors.

MorPHOLOGY

Habit is of some taxonomic importance. Low tufted forms, lacking woody stems above ground (fig. 4A-C), characterize G. jepsonii, ssp. nudicaule, usually ssp. gabrielense, and sometimes ssp. angustifolium. Taller forms with perennial scaffold stems (fig. 4D), as in much of ssp. angustifolium, may grow as high as 100 cm.

Stems in this genus are 4-sided, i.e., basically square in cross section. The angles are, however, more or less thickened and broadened by ex-

1971] DEMPSTER & STEBBINS: GALIUM 75

Fic. 4 Some representative habits « ¥%. In A, B, and C, ground level is indi- cated by dotted lines; D, E, and F are fragments broken off well above ground level. A, G. jepsonii, Cloudburst Summit (4235); B, G. johnstonii, Chilao Recreation Area (4234); C-F, G. angustifolium: C, ssp. nudicaule, Cloudburst Summit (4236); D, ssp. angustifolium, Little Thomas Mountain (4347); E, ssp. foliosum, Santa Rosa Island (Niehaus 459); F, ssp. gracillimum, 49 Palms Canyon (4174).

76 MADRONO [Vole2i

Fic. 5. Stem cross sections, X 24, to show comparative development of the angles. Collenchyma tissue with epidermis shown in black; stippled areas represent xylem, dotted circles endodermis. Size differences, although partially dependent on age of stems, are nonetheless of some taxonomic importance. A, G. jepsoni, San Gabriel Mountains (4235); B, G. johnstonii north side San Gabriel Mountains (4156); C-K, G. angustifolium; C, ssp. nudicaule, San Bernardino Mountains (4147); D, ssp.

1971] DEMPSTER & STEBBINS: GALIUM 77

Fic. 6. Details of epidermis from the sides (not angles) of the stem cross sections in Fic. 5: A, G. jepsonii; B, G. johnstonii; C-K, G. angustifolium: C, ssp. nudicaule ; D, ssp. gracillimum; E, I, ssp. angustifolium; F, ssp. onycense; G, ssp. gabrielense ; H, ssp. jacinticum ; J, ssp. borregoense; K, ssp. foliosum. All < 140.

trusion of the cortex and addition of collenchyma. Such enlargement of the angles occurs in varying degrees in the different species and sub- species of this group (fig. 5). It is least developed in G. jepsoni (fig. 5A), and has been carried to such an extreme in young stems of ssp. borregoense (fig. 5J) and ssp. onycense (fig. 5F) that the angles appear as sides and the nearly covered sides appear as mere longitudinal fissures. Subsequent growth in width of the stems causes the sides to emerge into visibility.

Stems are chiefly glabrous, but in ssp. gabrielense (fig. 5G) and in much of the tetraploid material of ssp. angustifolium (fig. 51) they are well supplied with hairs. Stems of ssp. nudicaule (fig. 5C), and others to a less noticeable degree, are papillose, the papillae consisting of soli- tary enlarged epidermal cells. Figure 6 shows details of the epidermis of the sides (not angles) of the stems illustrated in fig. 5.

All plants of the G. angustifolium complex apparently have stomata on both leaf surfaces. All leaves have 3 nerves, although the 3-nerved condition is not very obvious, and the lateral nerves, even when seen in

gracillimum, Black Rock Spring (Cole 989, POM); E, ssp. angustifolium, diploid form from Rincon Springs, San Diego Co. (4339); F, ssp. onycense, east of Onyx (1432); G, ssp. gabrielense, Ontario Peak (Johnston 1616, UC); H, ssp. jacinticum (4244, type); I, ssp. angustifolium, hairy tetraploid form from Del Mar (4347); J, ssp. borregoense, Palm Canyon (4375); K, ssp. foliosum, Santa Cruz Island (Elli- SON S.1., WiC):

78

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MADRONO

[Vol. 21

sears

\

=

NT ery:

\

BD) \ wenn ' i i) aS sa or a Oa cone a Roan

Wee \

Fic. 7. Leaves < 7, cleared and stained to show principal veins, distribution of secretory cells, and features of the leaf margins. All surface hairs have been omitted. A, G. jepsonii, Cloudburst Summit (4235); B, G. johnstonii, Chilao Recre- ation Area (4234); C-K, G. angustifolium; C, ssp. gabrielense, San Antonio Moun- tains (Raven 11,232, CAS, JEPS) ; D, ssp. foliosum, Anacapa Island (Hoffmann s.n., CAS); E, ssp. angustifolium, Catalina Island (Fosberg $4354, SMU, UC); F, ssp.

1971] DEMPSTER & STEBBINS: GALIUM 79

cleared and stained leaves, are sometimes tenuous, or even interrupted in very small leaves, as in ssp. foliosum.

Leaves (fig. 7), except the lowest, are for the most part strapshaped. Surface hairs are usual, and at least a few marginal hairs are nearly always present. Hairs vary considerably in length and stoutness, but are always either spreading or apically directed (fig. 9), never basally di- rected as in some members of the fleshy-fruited group.

The presence and distribution of secretory cells on the under side of the leaves is an important taxonomic character. These cells, of unknown function, have been observed in Galium species having 4-leaved whorls both in Europe (Nicolas, 1929) and in North America (Dempster and Ehrendorfer, 1965; Dempster and Stebbins, 1968). They are most commonly found clustered just below the leaf apex (fig. 7B-G, I-K) but are sometimes distributed over the entire lower surface, as in G. jepson (fig. 7A) and sometimes ssp. angustifolium. In ssp. gracillimum (fig. 7H) they occur in 2 bands at either side of the midrib. Table 2 summarizes our observations of this feature in the G. angustifolium complex, and shows that there is a considerable degree of taxonomic consistency with respect to this character.

Corollas of G. angustifolium are in general rotate, in conformity with the Linnaean description of the genus (fig. 8C-J). Old corollas es- pecially, however, may be a little cupped at the base, so that they need to be cut before they will lie flat. The occurrence of truly campanulate corollas in G. jepsoni (fig. 8A) and its derivative G. johnston (fig. 8B) adds another to an increasing list of Galium species known to have this contra-diagnostic character (see discussion, Dempster and Ehrendorfer, 1965; Dempster, 1968).

The occurrence of hispid corollas in conjunction with glabrous or relatively glabrous herbage is also of interest. This character occurs not only in the present group, but also in the G. multiflorum complex, where it is diagnostic of the two species G. matthewsi Gray and G. magnifolium Dempster. It is apparent that hispid corollas may be pro- duced in either of two ways: 1, by genes for general hairiness, the com- moner situation, wherein the corolla hairs are commensurate in size and abundance with those on stems and leaves; and 2, by a separate gene or genes, as in G. angustifolum ssp. nudicaule (fig. 8F), onycense (fig. 8D), gabrielense (fig. 8H), borregoense (fig. 8G) and jacinticum (fig. 8J). In the latter circumstance, the corolla hairs are usually long, stout, abundant, and conspicuous, often quite disproportionately to those, if any, on stems and leaves. Since hairs occur only on the outside of the corollas, they are especially noticeable before the buds open. This char- acter 1s apparent even to the naked eye, and is therefore very useful

angustifolium, Castle Peak (4067); G, ssp. borregoense, Palm Canyon (4375); H, ssp. gracillimum, type locality (Cole 989, POM); I, ssp. jacinticum, Fulmor Lake (4129); J, ssp. onycense, Spanish Needle Creek (Twisselmann 10,911, CAS, JEPS) ; K, ssp. nudicaule, Cloudburst Summit (4236).

80 MADRONO [Vol. 21

Fic. 8. Pistillate and staminate flowers, X 6: A, G. jepsonii, Cloudburst Summit (4235). B, G. johnstonii, Big Pines (4156). C-J, G. angustifolium: C, ssp. foliosum,

1971] DEMPSTER & STEBBINS: GALIUM 81

TABLE 2. DISTRIBUTION OF SECRETORY CELLS ON THE LOWER SURFACE OF THE LEAVES

(FIG. 7) Number of individuals with secretory cells Taxon clustered mostly below subapical scattered leaf apex plusafew over entire (subapical) scattered surface none seen G. angustifolium ssp. angustifolium 1G) 8 11 13 ssp. borregoense 1 1 ssp. foliosum 5 (few) 4 ssp. gabrielense 6 2 ssp. gracillimum 9(2ranks) 1 ssp. jacinticum 2 2 1 ssp. nudicaule 3 1 1 ssp. onycense 5 5 G. jepsonii i G. johnstonii 8

in identification. Its presence also provides a valuable clue to the origin of the polyploid sspp. gabrielense and jacinticum.

All races and subspecies of G. angustifolium, like all members of the G. multiflorum complex and of the polygamous G. parishii group, are characterized by long straight spreading hairs on ovaries and fruits (fig. 81’). Galium jepsonii, however, has short, upwardly curved hairs (fig. 8A’), and those of G. johnstonii are variably intermediate with re- spect to both length and curvature (fig. 8B’).

ACKNOWLEDGMENTS

This study was supported by National Science Foundation grant No. GB649. Laboratory space was generously provided by the Jepson Her- barium under the curatorship of Rimo Bacigalupi, who also kindly criticised and improved the Latin diagnoses.

Material from the following herbaria was examined, annotated, and most will not be cited: CAS, JEPS, NO, POM, RSA, SBBG, SBM, SMU, UC, UCSB. All unassigned numbers in this paper refer to collections (JEPS) of Dempster or of Dempster and Stebbins.

Anacapa Island (Howell 3792, CAS, POM); D, ssp. onycense, Spanish Needle Creek (Twisselmann 10,924, 10,911, CAS, JEPS) ; E. ssp. gracillimum, Snow Creek Canyon (Wolf 3648, RSA, UC), Black Rock Spring (Cole 990, POM); F, ssp. nudicaule, Cloudburst Summit (4236); G, ssp. borregoense, Palm Canyon (Munz & Hitchcock 11,339, POM, Dempster 4375); H, ssp. gabrielense, Sunset Trail, Johnston s.n., POM), San Antonio Canyon (Roos 400, POM); I, ssp. angustifolium, Del Mar (4347), Sespe Creek (4111); J, ssp. jacinticum (type, 4244). Position of corolla lobes depends considerably on the developmental stage. Size of corollas, although it varies with the individual, is nevertheless significant taxonomically. A’, B’, I’, ovary hairs x 40, to show relative size, shape and position: A’, G. jepsonii; B’, G. johnstonii; I’, G. angustifolium.

82 MADRONO [Vol. 21

KEY TO THE SPECIES AND SUBSPECIES

Fruiting pedicels 1-4 times as long as fruits; upper nodes much longer than the lower, the leaves often congested toward base of plant; fruits appearing longer than wide, the hairs shorter than the fruit body; corollas usually more or less cupped or campanulate.

Plants low, generally less than 16 cm high; corollas cleft about halfway, the lobes little spreading; secretory cells evenly distributed; diploid G. jepsonii Plants tall, generally over 18 cm high; corollas deeply cleft, spreading; secretory cells echnical: hexaploid . . . . . . G. johnstonii

Fruiting pedicels usually shorter than Pee modes proses equal, the leaves not congested toward base of plant; fruits, with hairs, appearing spheri- cal, the hairs usually as long as fruit body; corollas rotate . .G. angustifolium

Corollas usually hispid (not always in ssp. jacinticum). Stems glabrous or nearly so. Inflorescences narrow, relatively few-flowered, the branching little com-

pounded. Plants 6-16 cm high; leaves mostly 2-10 mm long; San Gabriel and San Bernardino mountains; diploid. . .. . . . . ssp. nudicaule Plants 17-35 cm high; [eae 11-26 mm long; San Jacinto Mountains; hexaploid . . . .. SSp. jacinticum Inflorescences pommel atone fone compodndly branched; plants 35— 50 cm high; Borrego Desert. . . . . . . . . . ssp. borregoense

Stems not glabrous.

Stems merely scabrous, the hairs few and shorter than those on the leaves; angles of stems much thickened, often nearly concealing the faces; north- eastern Kern Co.; diploid . . . . . SSp. onycense

Stems hispid, the Baits usually abundant ona lone fe those on the leaves; angles of stems narrow; vicinity of San Antonio Canyon, Los Angeles and San Bernardino counties; tetraploid . . . . . ssp. gabrielense

Corollas usually glabrous, or not more hispid than stems and leaves.

Internodes of scaffold stems short, often shorter than the leaves; plants gen- erally glabrous, the leaves very slender, often crowded; northern group of Channel Islands . .. . . . . ssp. foliosum

Internodes of scaffold stems fame pencralle) mace fenced than the leaves, the leaves thus never crowded; not in northern Channel Islands.

Plants tall and very slender, essentially glabrous; flowers, fruits and leaves diminutive, the latter early deciduous; deserts, San Bernardino and Riverside counties . . . . . . ssp. gracillimum

Plants stouter, tall or sometimes omen znd need: glabrous to often canes- cent; flowers, fruits, and leaves larger, the latter not soon deciduous; widespread in hills and mountains, but not in the deserts.

ssp. angustifolzum

GALIUM JEPSONII Hilend & Howell, Leafl. W. Bot. 1:135. 1934. Based on G. angustifolium var. subglabrum Jepson, Manual FI. Pl. Calif. 962. 1925. Type from Whitewater Basin, Wilder 1113 (UC), undoubtedly from San Bernardino Co.

Plants low (fig. 4A), the erect glabrous stems commonly 8-16 cm high, arising in small clumps from widely spreading slender rhizomes or small woody base; leaves congested at the lower third or half of the stem, the upper portion having conspicuously elongated nodes and reduced leaves (fig. 4A), internodes of the lower, leafy portion ™% to as long as leaves, internodes of the upper portion 2—5 times as long as leaves; leaves 6—15

1971] DEMPSTER & STEBBINS: GALIUM 83

Fic. 9. Details of margins of the leaves in Fic. 7: A, G. jepsonii; B, G. johnstonii;