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MADRONO
A WEST AMERICAN JOURNAL OF BOTANY VOLUME XVIII
1965-1966
BOARD OF EDITORS
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LyMAN Benson, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis HERBERT F. COPELAND, Sacramento College, Sacramento, California Joun F. Davipson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California MiuprepD E. MATHIAS, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley MARION OWNBEY, Washington State University, Pullman REED C. RoLiins, Gray Herbarium, Harvard University
Ira L. Wiccrns, Stanford University, Stanford, California
Editor—JoHN H. THOMAS Dudley Herbarium, Stanford University, Stanford, California 94305
BUSINESS MANAGER AND TREASURER Douglas M. Post
Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
Published quarterly by the California Botanical Society, Inc.
Life Sciences Building, University of California, Berkeley 94720
Printed by Gillick Printing, Inc., Berkeley, California 94710
To you. ANNETTA MARY CARTER—‘Senorita Anita” south of the Border—recent President of the California Botanical Society, long- time Secretary of the Editorial Board of Madrono, and during three administrations the leavening spirit of the University of California Her- barium, we affectionately dedicate this volume.
During your long association with the University Herbarium, from student assistant to Principal Museum Scientist, you have been the trusted advisor of faculty and administrative officers, a generous coun- selor and confidante of successive generations of grateful students, and an esteemed friend to your associates and herbarium visitors.
Champion of human rights and friend of the friendless; interpid field botanist and indefatigible collector and interpreter of the plants of the remote ranges of Baja California, especially of the Sierra de la Giganta; gracious ambassadress to our Mexican botanical friends—you have shown in all your broad and varied responsibilities over the years an unfailing skill and competence which is overshadowed only by your personal warmth and outstanding human spirit.
CONTENTS
PAGE Frontispiece: Annetta Mary Carter The inter-relationships of the Alaskan lupines, David B. Dumnn.............0.000000.0...... 1 The tribe Orcuttieae and the subtribes of the Pappophoreae (Gramineae), OV IUGR MIO OUL CT We Mii geno De cesd oan ca ate an Je Aaa ofa ae eee oon x ect oes pain e cates Pext ca ee De 18 1 RRCER TA GN 1ST a a re LE ete Oe Pe 28, 62, 126, 246 IN@beSiaiinG IN GW Stes, see cece ee esac oe cete tees neta eae cds cca 17, 31, 63, 104, 127, 160, 173,
184, 192, 213, 224, 250
Allolepis: a new segregate of Distichlis (Gramineae), Thomas R. Soderstrom
LIN CUE IEI- LID CONC, etter crac. etn cer ae ee tesa, Me tae, eee caster need 33 Natural hybridization between Cucurbita digitata and C. palmata, W. P. Bemis
VINA AOMILOS VY VV EUEGIC CF Pras atc crite sa det sd hase Sept aincd aces seats ac temas, geo 39 Manzanita chaparral in the Santa Ana Mountains, California, R. C. Wilson and
BES eee) fea EIU anna Re eet Ree, 2 IPE FO SEIN Zeer 2 GU nen ere nero 47 Clinal patterns in front anatomy of Polypodium, Robert M. Llovyd...................... 65
The California species of Gutierrezia (Compositae-Astereae), Otto T. Solbrig 75
Taxonomy of Polygonum, section Polygonum (Avicularia) in North America,
Homes ertens and Peter Lo WOv CH... i ee ice 85 A reevaluation of Buddleia corrugata, Eliane M. Norman................00.0000000000.000022--- 92 Dr. LeRoy Abrams’ collecting trip of 1908, Ernest Alexander McGregor.............. 97
The fleshy-fruited Galium species of California (Rubiaceae). I. Cytological find- ings and some taxonomic conclusions, Lauramay T. Dempster and G.
ECCS OIG PSUCUOIIUS a eens tee tae Se DOR MEN ac eat ech Rec te, cok 2s Chena ey ea ee 105 A further description of Gossypium trilobum, Paul A. Fryxell......00000000000ce eee. 113 New taxa in the genus Monarda (Labiatae), Rainier W. Scora............0..02....00.222--- 119 Documented chromosome numbers of plants.....000020000000000 cece eee 122, 245
A taxonomic revision of section Sonoricola of the genus Verbesina L. (Com- POsitae) a OM C\ma CO) CIA ore sate 5 ae, oe eee ene 129
Escobaria leei Bodeker rediscovered in New Mexico, Edward F. Castetter and
WUTC CTRL OTC Tree ti Unt! se a BI esd. Ease acc een: oes esta een a 13)/ New records of Myxomycetes from California I., Donald T. Kowalski................ 140 Contribution to the embryology of Muilla, with a remark on the taxonmic
position of the genus, Rolf Y. Berg and Jack R. Maze.................00..000000000022.-- 143 The genus Tiarella in western North America, Patricia Kern...........22..2.02200000----000--- 152 Varietal transfers in Cupressus and Chamaecyparis, Elbert L. Little, Jr............... 161
On the specific distinction of Eriogonum nutans and collinum, James L. Reveal 167 RORY Gs 8 ICY 0 Oe meng a eR Om ene Oe een TE 174
Notes on marine algae of Washington and southern British Columbia, Richard Be IOV ESMANNCS SGIED: VILES br cee ccf enece eet eee nce tet cours aetna ego ee 176
Some new chromosome numbers in Linum, C. Marvin Rogers and Betty D.
FL QUE 1S ota roe ek ee a es DN huastnd tee! co 19 Sclerolinon, a new genus in the Linaceae, C. Marvin Rogers...........2...2...c000c00000000000+ 181 Notes on Mexican grasses V. Two gypsophilous species of Muhlenbergia, Char-
lotte-G, Reedertand. JonneR Reed er een ae es 0 eo ee 185 Influences of fire and sodium-calcium borate on chaparral vegetation, Stephen
I ASCOCRING: cE ae eect A ee een nt VRS | ne ee 193
The taxonomic status of Stemmatella (Compositae-Heliantheae), B. L. Turner 203
Volatile growth inhibitors produced by Eucalyptus globulus, Herbert G. Baker 207
The chromosomsomes of Scoliopus (Liliaceae), Marion S. Cave............2...2..:2000000--- 211 Two new species of Verbesina (Compositae) with taxonomic notes on other species,.J ames I. iC leman 2 xc. :7, ee eee eee 214 Observations on witches’-broom formation, autoparasitism, and new hosts in Phoradendron, Frank G. Hawksworth and Delbert Wiens............20..220.200020-.- 218 Chrysothamnus bolanderi, an intergeneric hybrid, Loran C. Anderson and Jack BO ReveGl:. fone ise ied Rea ee 225 A new species of Brodiaea from San Clemente Island, California, Theodore F. IN GOW GUS. cS ec 2e osat Deaiescgp ok ccf SEs 233 Chromosome numbers in the Eriophyllum lanatum (Compositae, Helenieae) COmDleXS SON SM OOviIG ere ck, ce acps crs eee 236 Propagation of some woody desert perennials by stem cuttings, Valerie C. Chase:and (Bos SU Git cncaoka ce ee 240 Notes on the Hordeum jubatum complex, Tibor Rajhathy...............0..00...0:.0000-0-- 243 1 1 cc (<>, qt ae aos See Ren Dee RE OPOTY URC ON IRNny OMe ey ter er WMO lek. Mer tree aa we RA 252 ERRATA
Page 26, delete 3rd line from the bottom and replace by Cottea pappophoroides Kunth. Mexico. Sonora, Palmer 339, in 1887 (YU).
Page 93, delete first US and replace by UC
Page 124, line 26, E. should be Eriogonum
Page 124, under Eriogonum correct J. L. Reveal & J. L. Reveal to J. L. Reveal & J. A. Reveal
Page 124, under Eriogonum correct A. H. Holmgren to N. H. Holmgren
Page 135, line 13, correct spelling is encelioides
Page 135, line 14, correct spelling is PENINSULARIS
Page 149, note 2, line 2, correct spelling is carznatum
‘Pes | Mies 1-19 (727 od yes ges VOLUME 18, NUMBER 1 JANUARY, 1965 Contents PAGE THE INTER-RELATIONSHIP OF THE ALASKAN LUPINES, David B. Dunn 1
THE TRIBE ORCUTTIEAE AND THE SUBTRIBES OF THE PAPPOPHOREAE (GRAMINEAE), John R. Reeder 18
REvIEwS: Julian A. Steyermark, Flora of Missouri (Ed- gar Anderson); Herbert Parkes Riley, Families of Flowering Planis of Southern Africa (Robert J. Rodin) ; C. Leo Hitchcock, Arthur Cronquist, Marion Ownbey, and J. W. Thompson, Vascular Plants of the Pacific Northwest (Robert Ornduff); Richard Evans Schultes and Arthur Stanley Pease, Generic Names of Orchids (Donovan S. Correll) 28
NOTES AND News: THE MICHIGAN BOTANIST ily
TRANSLOCATION HETEROZYGOSITY IN CROTON CALI- FORNICUS, Jerzy Szweykowski; UNITED STATES NA- TIONAL HERBARIUM, William L. Stern; CROCIDIUM IN SAN Luis OBtsPo Co., CALIFORNIA, Peter H. Raven and Thomas R. Mertens; A REED COLLEGE HERBA- RIuM, Robert Ornduff 31
A WEST AMERICAN JOURNAL OF BOTANY
>UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
Entered as second-class matter at the post office at Berkeley, California, January 29,
1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price
$6.00 per year. Published quarterly and issued from the office of Madronio, Herbarium, Life Sciences Building, University of California, Berkeley, California 94720.
BOARD OF EDITORS
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LymMAN BENSON, Pomona College, Claremont, California HERBERT F. COPELAND, Sacramento College, Sacramento, California
Joun F. Davipson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California Mivtprep E. Matuias, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley Marion OwNBEY, Washington State University, Pullman REED C. RoLitins, Gray Harbarium, Harvard University Ira L. Wiccins, Stanford University, Stanford, California
Editor—JoHn H. THomas Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Annetta Carter, Department of Botany, University of California, Berke- ley. First Vice-President: Winslow R. Briggs, Department of Biological Sciences, Stanford University. Second Vice-President: Arthur R. Kruckeberg, Department of Botany, University of Washington. Recording Secretary: Mary L. Bowerman, De- partment of Botany, University of California, Berkeley. Corresponding Secretary, Margaret Bergseng, Department of Botany, University of California, Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State College.
THE INTER-RELATIONSHIPS OF THE ALASKAN LUPINES Davip B. DUNN
This paper is an attempt to clarify the complex relationships of Lupinus arcticus, L. nootkatensis, and L. polyphyllus. These are three of the north- ernmost species of a genus which has fascinated both professional and amateur botanists as well as horticulturalists and agronomists. Specimens upon which this study is based are deposited in the following herbaria: CAS, DAO, DS, GH, ISC, MO, PH, RENO, RM, UBC, UC, UMO, V, and WIS.
Recently Phillips (1955) reduced L. arcticus to a subspecies of L. poly- phyllus and L. nootkatensis to a subspecies of L. perennis. I find both of these transfers of rank untenable. Since Phillips cited no material other than type specimens and annotated a limited number of specimens, it is difficult to understand on what he based these changes.
I believe that Anderson’s methods (1949) offer the most sound means of interpreting species which have abundant morphological diversity but still permit occasional hybrids to form. In fact, if the presence of fer- tile hybrids is the criterion used in reducing the taxa referred to above to subspecific status, it would be necessary to reduce an entire series of very different morphological entities of the northwestern United States, as shown by the following list of putative hybrids (morphological inter- mediates). I have been hybridizing both annual and perennial lupines for the past 20 years and the hybrids have been almost invariably mor- phologically intermediate between the parents. It is also possible to dem- onstrate beyond reasonable doubt that these specimens are of hybrid origin by the use of pictorialized scatter diagrams. Six of the putative hybrids listed below have been plotted in Figs. 1-3.
It would be necessary to make additional diagrams for the species not treated in the present paper to demonstrate that the specimens listed actually do fall into place between the two parents. Some of the putative hybrids have been collected within the last 20 years, while others were collected much earlier. Hence, only part of the specimens in the list may be attributed to man’s activities in the transport of seeds and the devel- opment of habitats suitable to hybridization.
TABLE 1. List OF PUTATIVE HYBRIDS OR INTERMEDIATES IN HERBARIA
L. polyphyllus (fig. 4) one parent X:
L. arbustus ssp. pseudo parviflorus. Vegetatively mostly pseudoparviflorus but the flower structure can be accounted for by a blend of the characters of both. Montana. Lolo Valley, Kirkwood 1199 (MO); Hamilton, Blankinship 772 (MO).
L. arcticus (fig. 4). Alaska. 60 mi S of Fairbanks, Barkalow P21 (ISC, figs. 1, 3) ; Richardson Highway, mi 350, Anderson 2764 (ISC, figs. 1, 3). British Columbia. near Barkerville, Calder & Savile 14224 (DAO).
Maprono, Vol. 8, No. 1, pp. 1-32. February 10, 1965.
2 MADRONO [Vol. 18
L. nootkatensis (fig. 4). Alaska. Hope, Anderson 6589 (ISC, figs. 2, 3); 6589b (ISC) ; Kenai Peninsula, Calder 6207 (DAO); 6907 (DAO); 5728 (DAO). British Columbia. Queen Charlotte I., Calder, Savile, & Taylor 21382 (DAO, UMO, figs. 2, 3); Skeena Station, Calder, Savile, & Ferguson 13287 (DAO, UMO, figs. 2, 3) ; Prince Rupert, Taylor & Lewis 710 (DAO). Prince Edward I., Fernald & St.John 11104 (GH).
L. perennis. In areas where both are probably introduced. Nova Scotia. Col- chester Co., Zinck 371 (DAO). Victoria Co., Bassett 1778 (DAO). Prince Edward L., Bassett 1622 (DAO); probably backcrossed to L. perennis; Groh, 1926 (DAO, primarily L. perennis, a few characters show introgression).
L. arboreus. Probably a garden hybrid or field hybrid in an area where L. arbo- reus was introduced. British Columbia. Vancouver I., Newcombe, 1931 (V).
Unknown or uncertain second parent: a. Some parent in the L. lepidus-sellulus- lyalli group. Oregon. Klamath, Peck 9551 (MO). b. Possibly L. arbustus ssp. neo- laxiflorus Dunn. Washington. Seattle, Frezberg, 1915 (MO). c. Unknown or tran- sitional material to L. prunophilus, which has been treated as a variety of L. wyethii, L. polyphyllus and L. arcticus and as a species. The spreading sparse, pilose to hirsute hairs on the stems, petioles and midvein of the lower side of the leaflets, as well as cilia along the acumen of the keel are characters found in prunophilus; plus the habitat, on dry mountain slopes in open forests, eastern Oregon and Wash. Oregon. Cusick 1926 (MO). d. Unknown sources or mutant. Very narrow linear leaflets, a narrow banner and the acumen slants forward instead of the usual falcate condition. Washington. Chehalis Co., A. & G. Heller 3878 (MO); Snohomish Co., Thompson 6805 (GH, MO); Thurston Co., Meyer 943 (MO). e. L. (albicaulis ?). This is the material that Heller named L. pallidipes and Smith reduced to a variety of L. poly- phyllus. It is characterized by rusty strigose hairs above and below on the leaflets, in age; also ecologically distinct, growing in a dry rocky site, and flower structure intermediate. (Washington. Chelan Co., Thompson 6429 (MO, upper leaflet surface g'abrous). Oregon. Benton Co., Searing, May 22, 1925 (MO); Klamath Co., Rose 1241b (MO); Lane Co., Heller 10041 (RENO, type of L. pallidipes).
L. nootkatensis one parent x:
L.arcticus ssp. arcticus. Alaska. King Salmon, Schofield 2033 (DAO, fig. 3).
L. arcticus ssp. subalpinus. British Columbia. Bennett, Mitchell 155 (DAO)
L. (kuschei ?). Alaska. Adak I., Dowart 32 (ISC); without locality, Glassberg 980 (ISC). Both of these specimens would be explainable if they came from the upper end of Lynn Canal toward Skagway since L.kuschei was described frcm Carcross and Whitehorse, Yukon.
L. polyphyllus. See above.
L.arcticus one parent Xx: L. kuschei. British Columbia. Mt. Apex, Calder & Savile 10759 (DAO). Yukon. Near Whitehorse, Gillett & Mitchel 3040 (DAO); 3431 (DAO); Gillett & Ca'der 3179 (DAO); Kusche (CAS 62470), head of Dease L., McCabe 8819 (UC).
Lupinus polyphyllus has a natural range extending from central Cali- fornia into British Columbia in sites of a rather wet habitat (fig. 5). The material in Alaska appears to have been introduced since it does not show a natural continuous distribution and since the material from the Kenai Peninsula had very few specimens which could be considered as true L. polyphyllus. Most of the population suggested a hybrid swarm such as the /ris colonies reviewed in Anderson (1949). The Kenai Peninsula area near Hope and Anchorage has had considerable disturbance by man. In addition, the coastal stretch of British Columbia would appear either to be so poorly collected that none were found or else it has a long stretch
1965 | DUNN: ALASKAN LUPINES 3
IN CM.
RE RIOUE -EENG iH
8 oF On lt he W314 lS 167-18 19720721 WING LENGTH IN MM.
SPREADING APRESSED HEIGHT LOWER-LIP VERTICILS HAIRS HAIRS caYLx 6-204 6=ABSENT O=ABSENT 0-10 DM+ O=SHORT 0- 15+ 6=SPARSE O=SPARSE O- 5 DM+ O-INTER. O- 10+ O=MANY O=MANY 0+ 3DM- O“ LONG 07-7
Fic. 1. Scatter diagram comparing of L. polyphyllus, open glyphs, and L. arciicus, solid glyphs. Putative hybrids are indicated with a cross in the glyph. Petiole length is that of the lowest fully developed leaf (also in figs. 2, 3).
devoid of L. polyphyllus. In the southern part of the range of L. poly- phyllus two of the populations have become moderately distinct and have been treated as subspecies by Munz (1959). Of these two L. poly- phyllus ssp. superbus is apparently a relatively stabilized taxon derived by introgression from L. latifolius. The blend of characters has permitted the occupation of wet meadow habitats within a geographic area other- wise primarily suited to L. latifolius. These southern taxa will be
4 MADRONO [Vol. 18
treated in a latter paper. In western Oregon L. polyphyllus var. pallidipes appears to be of hybrid origin, combining traits from L. polyphyllus and apparently L. albicaulis. In eastern Oregon, Washington, and Utah L. prunophilus appears to have had some of its genetic traits derived from L. polyphyllus but the other sources are still obscure. In northern Wash- ington, southern British Columbia, and western Montana, L. polyphyllus forms hybrids with L. arbustus ssp. pseudoparviflorus as cited above, which by backcrossing appear to be the source of the material referred to as L. laxispicatus var. withamii, as well as, introducing a glabrous keel in a portion of the population (Dunn, 1956), where these latter two names were treated as synonyms. Specimens within the range of L. ar- bustus ssp. neolaxiflorus were obvious hybrids between that taxon and L. polyphyllus. In British Columbia a finely, sparsely sericeus condition of the upper surface of the leaflets and ciliation on the keel were mute evidence of introgression from yet another undetermined source. Hybrids of L. polyphyllus with L. nootkatensis were observed from British Co- lumbia, Alaska, and Nova Scotia (fig. 2). The introduced populations of L. polyphyllus on Prince Edward I. and in Nova Scotia now contain hybrids with the eastern L. perennis. Only a few hybrids with L. arcticus ssp. arcticus were found, but the flower structure suggests that both L. arcticus and L. polyphyllus contributed to L. nootkatensis. The latter has many distinct characters of its own as well as others derived from other sources. In short everywhere that L. polyphyllus has come into contact with another lupine, there has been some hybridization and subsequent introgression. In spite of this, it continues to maintain its distinctness. Several putative hybrids in which L. polyphyllus characters dominate have been included in the list, even though the second source of genes cannot be determined.
Key To TAXA TREATED AND OTHERS HYBRIDIZING WITH THEM OR IN CONTACT GEOGRAPHICALLY
Banner reflexed above midpoint, sulcus deep, clasping the wings; calyx spurred or gibbous at base; wings pubescent or glabrous; leaflets glabrous or pubescent @DOVE 225): anc rgtet At cee he ee tae me Beaten nes et eae L. arbustus (Dunn ,1956)
Banner reflexed approximately at midpoint, tip well separated from tips of the wings; calyx gibbous or not, not spurred; wings glabrous.
Longest petioles basal or on radical shoots. Leaflets glabrous above, occasionally with a few marginal hairs.
Bracts subpersistent; flowers small, 10 mm long or less; pedicels 1-3 mm long; Wyoming, Montana, and Idaho...............................- L. burkei (not treated)
Bracts caducous; flowers 10-20 mm long; pedicels 4-17 mm long. Stems with numerous hirsute, spreading hairs, 3-4 mm long; keel ciliate toward the acumen; Washington, Oregon, Idaho, Utah, and Nevada. L. prunophilus (Dunn, 1956a, not treated) Stems with appressed pubescence and some spreading pilose hairs, generally less than 2 mm long (execpt occasionally in L. polyphyllus ssp. poly-
1965 ] DUNN: ALASKAN LUPINES 5
phyllus, which introgresses with the last); keel glabrous (sometimes
ciliate in L. arcticus).
Stems generally slender, 3-4 mm in diameter, with 2-4 cauline leaves; plants usually 2—3 dm tall; racemes of 2—8 well spaced verticils.
3a. L. arcticus ssp. arcticus
Stems fistulose, 4-8 mm or more in diameter; plants generally 6—9 or 15 dm tall; racemes elongate, 20 cm or more long at maturity.
Leaflets generally 10-17, elliptic-oblanceolate, acute to acuminate; keel glabrous (except for hybrids or hybrid colonies) ; streamsides and wet meadows, primariy in areas of high rainfall of British Columbia, Washington, Oregon, and California
1. L. polyphyllus ssp. polyphyllus var. polyphyllus
Leaflets generally 6-9, oblanceolate, rounded or acute; keel with minute papillae above the claws; endemic in the Sierra Nevada of Cali- fornia and Nevada........ L. polyphyllus ssp. suberbus (Munz, 1959)
Leaflets pubescent above. Plants 4-10 dm tall; stems fistulose; keel slender, falcate, angle less than a 90° angle......L. polyphyllus ssp. polyphyllus var. pallidipes (Munz,. 1959) Plants 2-5 dm tall; keel angle about 90°-120°.
Flowers slender, viewed laterally; banner elliptic to obovate; bracts per- yoy =) 0 Fe cee oe L. lepidus complex (not treated)
Flowers orbicular, viewed laterally; banner orbicular to broadly obovate; bracts persistent in L. kuschei, tardy or caducous in L. cottonii, L. orna- DUS IL UHLTOOLG. exe TO NCUINIL <2 et eee eee yy ee (none treated)
Longest petioles cauline, several leaves above base.
All petioles short, about as long as leaflets to slightly longer; leaflets linear to linear-oblanceolate..................200.....2.0--- L. rivularis, L. arboreus (not treated)
Longer petioles 114-4 times as long as the leaflets. Foliage finely sericeus; keel glabrous, strongly arcuate-falcate, angle about 80°; stems solid; banner tip acute...................... L. albicaulis (not treated) Foliage finely strigose to sericeus or glabrate; leaflets generally glabrous above (note a % in L. nootkatensis and in L. arcticus ssp. subalpinus) ; keel
usually ciliate; stews hollow, often fistulose.
Plants rhizomatous with slender stems arising singly or several close to- gether; keel ciliate toward the acumen; eastern North America, west to Minnesota and Iowa...................... L. perennis and allies (not treated)
Plants from an underground caudex, offshoots may arise a few inches out (occasionally rhizomatous in L. arcticus, but then with long basal petioles, see above).
Banner obovate, often obtuse; stems slender, branching above; racemes primarily south of the Columbia R. ............ L. latifolius (not treated)
Banner orbicular, wider than long, emarginate; stems hollow, fistulose to slender; plants 2-3 dm tall.
Stems fistulose; lower calyx lip enlarged, boat shaped; secondary branches often slender; pubescence shaggy-lanate (sericeus in L. nootkatensis var. fruticosus), upper surface of leaflets glabrous or strigose; wings 15-22 mm long; keel ciliate above the claws; mari- time or insular Alaska and British Columbia......2. L. nootkatensis
Stems hollow but usually not fistulose; lower calyx lip relatively slender (as in L. arcticus); leaflets various but not lanate; wings 12-15 mm long; keel glabrous or ciliate toward the acumen; near timber- line, Vancouver I., British Columbia, to montane Washington and Je E13) 6 ene Re ee eee ee er 3b. L.arcticus ssp. subalpinus
6 MADRONO [Vol. 18
1. LUPINUS POLYPHYLLUs Lindl., Bot. Reg. pl. 1096. 1827, ssp. POLY- PHYLLUS var. POLYPHYLLUS. Type: Douglas 116 (CGE, Dunn, 1956b). L. grandifolius Lindl. ex Agardh, Syn. Gen. Lup. 18. 1835. Type: Cali- fornia?, Douglas (Lindley Herb., CGE). L. polyphyllus var. grandifolius (Lindl. ex Agardh) Torr. & Gray, Fl. N. Am. 1:375. 1840. L. magnus Greene, Pittonia 3:160. 1897. Type: not seen, but specimens labeled by Green seen (JEPS, UC). LZ. matanuskensis Smith, Spec. Lup. 651. 1949. Matanuska, Alaska, Anderson 945 (ISC). L. pseudopolyphyllus Smith, Spec. Lup. 650. 1949. Type: Moose Pass, Alaska, Anderson 6825 (ISC, shows traits of introgression from L. nootkatensis). L. stationis Smith, Spec. Lup. 651, 1949. Type: Anchorage, Alaska, Purer 7539 (DS, hybrid to L. nootkatensis). L. polyphyllus ssp. polyphyllus var. polyphyllus, sensu Phill., in part, Res. Stud. St. Col. Wash. 23: 180. 1955.
Plants perennial, from a branched caudex, new shoots sometimes aris- ing several inches away from the old stem, subrhizomatous; stems gener- ally unbranched, fistulose, 5-15 dm high, glabrate but sometimes scat- tered pilose to hirsute hairs present, with a sparse undercoat of minute appressed puberulent hairs, generally only 3—5 cauline nodes, several basal leaves with long petioles; petioles of the lowest normal leaves vary from 1.4—6.2 dm long, gradated upward, with the upper petioles only 6-10 cm long; stipules also gradated, as much as 3.5 cm long basally and only 1 cm long above, with 3—5 mm free at the tip of the lower ones and 5—7 mm free on the upper ones, the remainder of each stipule connate to the petiole; leaflets 11-14 (17) elliptic-oblanceolate, the tip acute, some- times accuminate, the largest 5.5—-12 cm long, 10-22 mm wide, glabrous above (strigose in L. polyphyllus var. pallidipes), sparsely strigose be- low; peduncles 4-14 cm long; racemes 18—40 cm long at maturity, gen- erally verticillate but sometimes scattered, usually dense, the verticils 10-22 mm distant at anthesis; bracts subulate to lanceolate, caducous, 10-13 mm long; pedicels spreading to ascending at anthesis, 5.4—9.4 mm long, filiform, densely to sparsely spreading hispidulose; calyx sericeus to puberulent, the two lips nearly equal, 3.6—7.5 mm long, both entire or with minute teeth, the upper lip with a notch averaging 0.3 mm, the lips connate 0.7-1.5 mm with a flat bracteole 0.4-1.5 mm long at the sinus; banner 11.6-14.8 mm long, 9.6-14.0 mm wide, broadest below the mid- dle, reflexed at the midpoint (reflexed/appressed ratio 1.05), the angle of reflection 128°-154° (average 137°), the deep sulcus averages 2.1 mm deep at the mid-point along the appressed part of the banner; wings 12.0-15.7 mm long (average 14.1), 6.7-9.8 mm wide (average 8.2); keel 3.7-4.8 mm wide at the middle with a slender arcing acumen, the angle 68°—90° (average 82°); pods arching, 28-50 mm long, 7-9 mm wide, densely lanate with hairs 2-3.5 mm long; ovules 6-10; mature seeds 2.2—2.8 mm wide, 3.0—3.8 mm long.
The relationships of several characteristics are illustrated (figs. 1-3) showing a random sample of 25 members of each of the three species treated. The characteristics conformation of the floral parts of L. poly-
1965] DUNN: ALASKAN LUPINES 7
62 45) Ne v HEIGHT 36;
‘0 10 DM+ 34 v
0 5 DMt
2 WY ef 0 3 DM-
IN CM
PEN IOLE “LENGTH
6229" 10 MW l2 1314 5 1617. 8 19:20<2) WING LENGTH IN MM
SPREADING APRESSED LEAFLET LOWER-LIP VERTICILS HAIRS HAIRS TIP CAYLX 0-204
6=ABSENT O=ABSENT O=ACUTE O=SLENDER O= 15+
O=SPARSE O=SPARSE O=INTER. O= INTER. OF 10+
O-MANY O=MANY O*ROUND O= BROAD ON 7
Fic. 2. Scatter diagram comparing L. polyphyllus, open glyphs, with L. nootka- tensis, solid glyphs. Putative hybrids are indicated with a cross in the glyph. Both varieties of L. nootkatensis are included among the plants plotted.
phyllus is lilustrated in Fig. 4-1. The size of each part of the flower was drawn to scale for the mean value of a sample of 25. The range of L. poly- phyllus (in the strict sense) is shown in Fig. 5. Ecologically the species prefers a wet habitat, commonly occurring in seepage areas or on stream- banks; areas of high rainfall, cool nights, cold winters and the fog belt from northern California to British Columbia. Those specimens from drier sites show introgression or intergradation into subspecific taxa as indicated.
8 MADRONO [Vol. 18
2a. LUPINUS NOOTKATENSIS Donn ex Sims, Bot. Mag. 32: pl. 1311. 1810, var. NOOTKATENSIS. L. nootkatensis var. henry-looffii Smith, Sp. Lup. 653. 1949. Type: Olga Bay, Alaska, E. & H. Looff 667 (DS). L. nootkatensts var. ethel-loo fiae Smith, Spec. Lup. 653. 1949. Type: Alitak Mts., Alaska, EZ. & H. Looff 125 (DS, isotypes at GH, MO, intergrades to L. nootkatensis var. fruticosus). L. nootkatensis var. perlanatus Smith, Spec. Lup. 655. 1949. Type: St. Paul’s I., Alaska, Anderson 4035 (ISC). L. nootkatensis {. leucanthus Lepage, Am. Midl. Nat. 46: 758. 1951. Type: Kodiak, Alaska, Lepage 25223 (LCU, not seen). L. perensis ssp. nootkatensis (Donn ex Sims) Phill., in part, Res. Stud. St. Coll. Wash. 23.2 V75, 1955:
Perennial plants, dying back annually to a subterranean woody cau- dex; stems 4-10 dm tall in favorable locations, barely 10 cm tall in loca- tions of severe exposure such as windswept bluffs, hollow, the larger quite fistulose, 1—-1.5 cm in diameter, with abundant pilose hairs, becom- ing sparser by sloughing off with age; petioles all cauline, all short, 2-10 cm long, seldom twice as long as the leaflets, the longest on the primary stem about midway in height; stipules on lower abnormal leaves up to 6 cm long, connate, except for 6-10 mm of subulate to filiform free tip, upper stipules 1.5—2.5 cm long, connate for only 2—4 mm, woolly with pilose hairs up to 4 mm long; leaflets 5-8, broadly oblanceolate, the tip round, mucronate, shaggy-villous below, generally glabrous above, but sparsely villous near the margins in about 30% of those seen, about 5% were pubescent over the entire upper surface, the largest leaflets from 1.8—6.5 cm long (average 4.17), 4-16 mm wide (average 10.7), the small- er from sites of severe exposure; peduncles 1—9 cm long, longest in fruit; racemes from 2—10 cm, sometimes to 30 cm in favorable sites, 1-15 ver- ticils, sometimes 20, generally less than 10, verticils 15-35 mm distant at anthesis; pedicels 7-13 mm long, pilose; bracts 16-19 mm long, filiform, convolute, caducous; lower lip of calyx 7.6-11.5 mm long, broad, boat shaped, bent backwards, deeply toothed at the apex, woolly without and sericeus Over most of the inside; upper calyx lip deeply two lobed the notch 0.4—2.7 mm deep, lobes 6—9 mm long, woolly outside, sericeus in- side, the lips connate 1-1.4 mm. bracteole at the lip of the sinus attached midway or basal, filiform, 1.7—5.0 mm long, woolly with long pilose hairs; banner 12.5-19 mm long, 14.0—-21.4 mm wide (length/width radio 0.85) broadest below the middle, reflexed part 7.4-10 mm long, appressed part 6-9 mm, the sulcus 1.5—2.7 mm deep midway along the appressed part, angle of reflection 112°-141° (average 129°); wings 14-20 mm long, 8.3-11.0 mm wide (average 10); keel 4.0-6.0 mm wide at middle, ciliate along upper edges, the angle curvature 82°—95° (average 90.4° ) ; ovules 7-11 (average 9.4); mature pods 5-6 cm long, 10-12 mm wide, arcing up and outward, lanate when young but most sloughing off the sides, thinly sericeus when seeds are shed, black in age; seeds 4.6—5.0 mm long, 2.5—3.0 mm wide, scar of the funiculus large and subterminal, leaving a deep hole in seed coat, from black to light, mottled.
1965 | DUNN: ALASKAN LUPINES 9
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A= LUPINUS” POLYPHYLEUS;: s=L.ARCTICUS: O=L. NOOTKATENSIS; 4,0, = HYBRIDS ?
Fic. 3. Scatter diagram of all three species and their putative hybrids. Values on the abscissa represent the sum of the index values plotted in Figs. 1, 2. Dotted line arrows indicate how the putative hybrids fall in between the taxa being compared. Position of the glyph for one putative hybrid between L. arcticus and L. polyphyllus suggests that it represents backcrossing to L. arcticus.
The habitat of this species is essentially maritime (fig. 6). It occupies coastal areas in Alaska and British Columbia and the Aleutian I. chain. Its southern limit appears to be Vancouver I. It has been introduced into Newfoundland, Nova Scotia, and Scotland. It has the most westerly nat- ural distribution of any species in the genus.
In addition to the relationships discussed in the introduction of this paper pertaining to L. polyphyllus and L. arcticus, L. nootkatensis com- bines or shares characteristics from several taxa with which it currently does not share any geographic range. Lupinus arboreus has a maritime
10 MADRONO [Vol. 18
habitat, the boatshaped lower calyx lip, large flowers of similar confor- mation, ciliate keel toward the acumen and short petioles in common. Lupinus latifolius has fistulose, herbaceous stems, the same shaped leaf- lets and a ciliate keel above, near the claws, in common. These character- istics in common may be interpreted as sources of the genetic traits in the present day material recognized as L. nootkatensis, or that these sev- eral taxa had ancestors in common, in part. Natural hybrids occur be- tween L. nootkatensis and several other species as discussed above (figs. 2,3) and it has characteristics from several of these. It has a rather well defined geographic range and a genome which has become adequately established to consider this taxon as a species.
Two unusual specimens were noted Dorwart 32 (ISC) from Adak I. and Glassberg 980 (ISC) labeled “‘Alaska.”” These two specimens are vir- tually identical in their morphological traits and if they did not come from the same population they constitute a variant which could require a name. The difference between these specimens and typical L. nootka- tensis is greater than that between var. fruticosus and var. nootkatensis and there have been several names applied to each of these varieties. Lupinus kiskensis Smith may be similar material. The stems are more slender and covered with hirsute straight spreading hairs about 1-2 mm long, instead of soft lanate hairs 2-4 mm long; the leaflets are narrowly oblanceolate, obtuse, mucronate, 2-3.5 cm long, 5—7.5 mm wide, sericeus beneath and appressed strigose to pilose hairy above; the peduncle is 2—5 cm long; the raceme is dense with scattered flowers; bracts caducous; pedicels 6-10 mm long, densely villous; the lower calyx lip is slightly reflexed, boat shaped but not as large as typical nootkatensis; the flowers are 13-15 mm long, which is small for nootkatensis and the veins are dark and darken further with age; the petals are glabrous including the keel, except for minute papillae above and below the claws as found in L. polyphyllus ssp. superbus and occasionally in typical polyphyllus. The appearance of these plants could be accounted for if they were hybrids between L. kuschei from northern British Columbia and southern Yukon and L. nootkatensis. However, Adak I. is so far from the nearest stations known for L. kuschei that this hypothesis seems unlikely.
2b. LUPINUS NOOTKATENSIS var. FRUTICOSUS Sims, Bot. Mag. 47: pl. 2136. 1820. L. nootkatensis var. glaber Hook., Fl. Bor. Am. 1:163. 1834. L. nootkatensis var. unalaskensis Wats., Proc. Am. Acad. 8:524. 1873. Lectotype: designated by previous authors, Harrington, 1871 (GH). L. perensis ssp. nootkatensis (Donn ex Sims) Phill., in part, Res. Stud. St. Coll. Wash. 23: 175. 1955.
Stems slender, the primary hollow, to 6—7 mm in diameter, the upper branches 1.5—2.5 mm in diameter, finely appressed silky sericeus through- out, except the upper surface of the leaflets generally glabrous, about 13 were sparsely strigose above near the margin; stipules shorter than in L. nootkatensis var. nootkatensis and sericeus; pedicels hispidulose, bracts 7-13 mm long, sericeus and generally flat and subulate; lower
1965 ] DUNN: ALASKAN LUPINES 11
calyx lip the same shape but with smaller teeth and sericeus inside and outside; the upper lip the same as the species but sericeus on both sides; bracteole shorter and sericeus; the remaining parts of the flower are not distinct from the species.
The original description stated a sub-shrubby plant, which has led most authors to question the identity. The plants indeed appear sub-shrubby in vigorous specimens, particularly in locations mild enough to permit secondary and tertiary branches to flower. These later flowering branches often have only a few verticils. The floral structures in nature are clearly conspecific. The sericeus nature of the pubescence and the shape of the pods, and the number and shape of the seeds, as well as the short petioles are characteristics which occur in both L. littoralis and L. arboreus. Lupinus littoralis is the northernmost representative of the subshrubby L. arboreus complex and it occurs on the north end of the Queen Char- lotte Islands in proximity with L. nootkatensis. The above traits are dif- fused throughout the species, so that var. fruitcosus is completely sym- patric with var. nootkatensis (fig. 6) and the genome for the sericeus condition and the more slender stems appears intermittently. Late season growth of var. nootkatensis may resemble var. fruticosus in general ab- sence of lanate hairs by the sloughing of the long hairs; however, there are usually some areas on the stem with residual patches of long hair.
Watson (1873) placed var. fruticosus in L. littoralis. The original illus- tration does show some characteristics of L. littoralis. As noted above L. nootkatensis has several characteristics which are in common with or derived from L. arboreus. In the original description of var. fruticosus, Sims wrote: ““Communicated by Mr. ANDERSON from the Chelsea garden, where it was introduced by Mr. LEE, of the Hammersmith Nursery, who informs us it was first raised in Somersetshire from seed of LUPINUS perensis or nootkatensis, with the latter of which particularly it appears to have a very near affinity.”’ Since the name was applied to a plant of garden origin, which quite obviously had L. nootkatensts as the female parent (both taxa mentioned are herbaceous in nature) I would suggest that the nursery had material of L. arboreus growing nearby, available as the male parent. Lupinus arboreus had already been introduced and described in England. I have personally succeeded in crossing L. arboreus with several other shrubby lupines from California, using it as the male parent, but all attempts using L. arboreus as the female parent failed.
3a. LupINus ArctTicus Wats., Proc. Am. Acad. 8:526. 1873, ssp. ARC- Ticus. Lectotype: Bear Lake, Richardson (GH, specimen formerly in herb. Hookerianum), designated by Eggleston. L. nootkaiensis var. kjcll- mannu Ostf., Vidensk.-Selsk Skr., Math.-Naturv. Kl. 8. 1910. Type: King Pt., Yukon, Ostenfeld, 1908 (not seen, location excludes all other taxa). L. borealis Heller, Muhlenbergia 8:82. 1912. Type: Klondike R.. Yukon, Macoun 58426 (RENO). L. yukonensis Greene, Leafl. Bot. Obs. 2:233. 1912. Type: Klondike R., Yukon, Macoun 58426 (ND). L. gako- nensis Smith, Spec. Lup. 649. 1949. Type: Gakona, Alaska, Anderson
12 MADRONO [Vol. 18
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Fic. 4. Floral and leaflet characteristics of: 1, L. polyphyllus, 2, L. nootkatensis, and 3, L. arctcius ssp. arcticus. Drawings are based on the mean values of a random sample of 25 plants. Floral measurements were made as described by Dunn (1954). Floral parts from top to bottom are: ventral view of the unfolded calyx cut along the left lateral sinus; side view of keel; side view of wing; banner flattened; and side view of intact flower. Dissected calyx in the right row illustrates the different pubes-
1965 | DUNN: ALASKAN LUPINES 13
8532 (ISC). L. multicaulis Smith, Spec. Lup. 649. 1949. Type: Glenn Highway, Alaska, Anderson, 1944 (ISC). L. donnellyensis Smith, Spec. Lup. 654. 1949. Type: Richardson Highway, Alaska, Anderson 2281 (ISC). L. multifolius Smith, Spec. Lup. 660. 1949. Type: Lake Kluane, Yukon, Anderson 9449 (ISC). L. relictus Hult., Fl. Alaska and Yukon 7:1074. 1947, in synon. L. polyphyllus ssp. arcticus (Wats.) Phill., in part, Res. Stud. St. Coll. Wash. 23:181. 1955.
Plants perennial, dying back to an underground branched, woody caudex; stems clustered 2—4 dm tall at anthesis elongating more as the fruit develops, pubescence thinly silky sericeus, appressed or with few to numerous spreading pilose hairs; petioles of the early leaves very stunted at northern latitudes on overwintering radical shoots of the previous season, while new radical shoots develop the longest petioles at the base of the plants, radical shoot with petioles 9-15 cm long, apparently devel- oping flowering culms after one or more years growth as a radical shoot; stipules variable, subulate to setaceous in the free portion, the connate portion up to 3 cm long on basal leaves; leaves all dying back to ground level annually; leaflets 6—10, elliptic-oblanceolate with an acute, mucro- nulate tip, glabrous above, thinly strigose below, 2.5—5 cm long and 7-9 mm wide at anthesis, becoming 10-15 mm wide and up to 9 cm long at fruiting time or in favorable habitats; peduncles 4—7 cm long; racemes generally 5—8 cm long occasionally to 15 cm long; 3-7 verticils, 12-18 mm distant at anthesis; bracts, subpersistent to caducous, setaceous. 8-15 mm long; pedicels filiform, 4-6 mm long at anthesis, fine pilose spreading hairs 0.5-1.0 mm long; calyx strigose to villous, the lower-lip 6.5-10.8 mm long, slender lanceolate arcing down, the upper-lip quite gibbous at the base, 4.7—-8.2 mm long, the notch 0.3—2.7 mm deep, the lateral sinus-bracteole 0.4—3.0 mm long attached at the base, midway up or at the lip of the sinus notch between the lips of the calyx; banner orbicular 13.8-19 mm wide (average 15.8), 13.8-17.3 mm long (average 15.2), length width ratio 0.96, reflexed at about the midpoint, average 7.8 mm reflexed and 7.6 mm appressed, reflexed appressed ratio 1.026, the angle of reflection 107°-142° (average 124.5°); wings 14.2—-18.7 mm long, 7.2-9.0 mm wide; keel 3.5—5.0 mm wide at the mid-point, the angle 82°-96°, glabrous or sparsely ciliate above toward the acumen; ovules normally 5-8 (average 7); mature pods 43 mm long, 9 mm wide, silky- lanate.
Of the four specimens available to Watson when he described this species two were formerly in Herbarium Hookerianum and apparently were exchange material obtained from a British voyage along the arctic
cence and the bract of L. nootkatensis var. fruticosus. Large leaflet outline is of typical L. polyphyllus. Lower surface of the typical leaflet of L. arc icus is drawn within the outline of a L. polyphyllus leaflet. Typical leaflet of L. nootkatensis is drawn in the right row, with the lower half as the bottom surface. Variation in the pubescence of the two varieties is illustrated for both surfaces, with L. nootkatensis var. fruticosus as the right half.
14 MADRONO | Vol. 18
A4-LUPINUS POLYPHYLLUS O=LUPINUS ARCTICUS
@=LUPINUS ARCTICUS ssp. SUBALPINUS
Fic. 5. Distribution of L. polyphyllus and L. arcticus in western North America. Lupinus polyphyllus has been introduced in to eastern Canada as is shown by the insert. Half shaded circles indicate L. arcticus intermediates, see text.
coast of the Northwest Territories. Present day material most closely matching the Polar Sea specimen of Dease and Simpson is from the vicin- ity of Bathhurst Inlet, south of Dease Strait. Both of these specimens, used by Watson, were from the vicinity of the Arctic Circle. The other two specimens included: Vancouver I., Wood; and Washington Territory, Lyll, both from the southern end of the range of ssp. subalpinus. Wat-
1965 ] DUNN: ALASKAN LUPINES 15
aes
@=LUPINUS NOOTKATENS!S
O=LUPINUS NOOTKATENSIS vaR. FRUTICOSUS
Fic. 6. Distribution of L. nootkatensis in western North America. It has been introduced into eastern Canada as is shown by the insert.
son’s statement ‘“‘and probably all the L. perennis from the Arctic Coast,”’ clearly suggests that Eggleston’s selection of the arctic specimen as the type complied with Watson’s intention.
The typical material is thus on the arctic slopes extending upward into the montane areas in Alaska and Yukon but primarily on the mounds in the relatively flat tundra regions. The montane material in both areas has smaller flowers, is more erect, and has more cauline leaves. The latter character may have been acquired from L. nootkatensis but the habitat is generally not allied, nor is the floral conformation. The smaller flowers
16 MADRONO [Vol. 18
could have come from L. kuschei in the Carcross area in southern Yukon and several hybrids are cited but L. kuschei is itself a “stabilized” entity derived from a few L. arcticus characters, blended with primarily L. seri- ceus characters. The smaller flowered ssp. subalpinus with cauline leaves extends southward in British Columbia and into Washington, in areas near permanent snow. Subspecies subalpinus extends far southward into the range of L. polyphyllus but occurs at higher altitudes in the south and in a very different habitat. I have not been able to find any indication of a transition between the two, suggesting that free gene exchange does not occur. Since only occasional isolated hybrids have been found (fig. 1) and each has a distinct distribution pattern (fig. 5), I consider L. arcticus and L. polyphyllus to be ecospecies, which introgress only occasionally. The greatest affinity of L. arcticus is with L. perennis and from which it is isolated by a distance of over 1000 miles.
3b. LUPINUS ARCTICUS ssp. subalpinus (Piper & Robins.) Dunn, comb. nov. L. subalpinus Piper & Robins., Contr. U.S. Nat. Herb. 11:356. 1906. Type: Cascade Mountains to Fort Colville, Washington, Lyall, 1860 (GH). L. latifolius var. subalpinus (Piper & Robins.) Smith, Bull. Torrey Club 51:308. 1924. L. arcticus var. subalpinus (Piper & Robins.) Smith, in Abrams, Il. Fl. Pac. States 2:518. 1944. L. glacialis Smith, Spec. Lup. 236. 1940. Type: Snohomish Co., Washington, Thompson 14705 (DS). L. volcanicus var. rupestricola Smith, Spec. Lup. 236. 1940. Not L. volcanicus Greene. Type: Lewis Co., Washington, Thompson 15210 (DS). L. latifolius var. canadensis Smith, in part, Bull. Torrey Clubs: 307, 1924.
The leaves all cauline; the longest petioles 4-8 cm long, lower and mid- dle; leaflets narrowly oblanceolate, acute to obtuse or rounded and mu- cronate at the tip, generally glabrous above and strigose beneath but often also sparsely strigose above; racemes commonly with 4—6 verticils but occasionally more; flowers generally smaller than the species but gen- erally with the same conformation, smallest in the areas of montane Washington; ciliation may occur on the lower edge of the wings, as well as along the entire length of the upper edge of the keel or only along parts of the upper edge, often reduced to minute papillae above the claws of the keel.
This subspecies combines the cauline leaf trait and the rounded tips of the leaflets in a percentage of the population, from L. nootkatensis, as well as ciliation characters of the keel, with floral and leaf character and habitat preference of L. arcticus. In addition the smaller flowers may have been acquired by introgression from L. kuschei of the L. sericeus complex. The more southerly range suggests that daylength factors have also been acquired, probably through introgression. I consider the speci- men from Needles, British Columbia, Calder & Savile 10100 (DAQ) to be a hybrid between L. argenteus and L. arcticus ssp. subalpinus, while a second specimen from Edgewood, British Columbia, is primarily L. arcticus ssp. subalpinus with introgression from L. argenteus. Hybrids
1965 ] DUNN: ALASKAN LUPINES 1
with L. polyphyllus or L. nootkatensis are obvious but the transition into L. arcticus appears complete in the montane areas in Alaska, Yukon and British Columbia. Hence, the contention that this taxon is primarily a derivative of L. arcticus. To the south, this taxon does not form a bridge or connecting link with L. polyphyllus as Phillips’ (1955) treatment would suggest but instead it appears to be more closely connected with L. latifolius and L. rivularis of Washington, Oregon, and California. The best break in the maze of inter-acting taxa appears to be in the Columbia River area. The glacier margin or arctic-alpine habitat of ssp. subalpinus suggests that both it and LZ. arcticus form a northern ecospecies, while L. nootkatensis forms a maritime ecospecies, and L. latifolius a related and connected southern ecospecies, which also has multiple derivatives, as evidenced by the numerous names that have been applied to the latter complex.
Department of Botany, University of Missouri, Columbia
LITERATURE CITED
ANDERSON, E. 1949. Introgressive Hybridization. John Wiley & Sons, N. Y.
Dunn, D.B. 1954. A method of mounting pressed flowers for study and preserva tion. Rhodora 56:258—260.
. 1956. A revision of the Lupinus arbustus complex of the Laxiflori. Ma- drofo 14:54-73.
. 1956a. Leguminosae of Nevada, Part IJ]—Lupinus. Contr. Fl. Nevada 39:1-64.
————-, 1956b. The lupines collected by David Douglas from 1825 to 1827: their type localities and nomenclature. Leafl. West. Bot. 8:47-54.
Mowz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
Puiiies, L. L. 1955. A revision of the perennial species of Lupinus of North Amer- ica exclusive of southwestern United States and Mexico. Res. Stud. St. Coll. Wash. 23:161-201.
Watson, S. 1873. Revision of the extra-tropical North American species of the genus Lupinus. Proc. Am. Acad. 8:517-548.
NOTES AND NEWS
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Published four times per year by the Michigan Botanical Club, this journal is devoted to all phases of botany in the Great Lakes region, and has included major articles on vascular plants, fungi, bryophytes and lichens, and exploration. Articles have been included on ecology, phytogeography, taxonomy, morphology, and tech- nique, as well as annotated bibliographies and lists of new literature and maps.
THE TRIBE ORCUTTIEAE AND THE SUBTRIBES OF THE PAPPOPHOREAE (GRAMINEAE)
JOHN R. REEDER
In the days when grasses were classified on the basis of gross morpho- logical features alone, the placement of the annual California endemics, Orcuttia and Neostapfia, presented few problems to the agrostologist. Vasey (1886), who described Orcuttia, assigned it confidently to the tribe Festuceae, subtribe Seslerieae. Neostapfia likewise was considered a mem- ber of this tribe by its discoverer (Davy, 1898), but he assigned it to a different subtribe, the Meliceae. The genus was originally described as Stapfia, but this name was “stillborn” since it is a homonym of Stapfia Chodat (1897). In the following year, Davy (1899) renamed his new genus Neostapfia. The structure of the spikelet in these genera would suggest that their alliances are, indeed, with the Festuceae. In both, the spikelets are more or less laterally compressed, consist of sev- eral to many florets, and the glumes and lemmas are of a similar texture. The most recent floras which treat the California grasses (Hitchcock, 1950; Mason, 1957; Munz, 1959) still follow this placement.
Neostapfia is monotypic, whereas Orcuttia is known from 5 species and two varieties. Both genera are restricted to a special type of vernal pool or “hog wallow” within the Great Valley of California. Thanks to the work of Hoover (1941) and more recently Crampton (1959), a great deal is now known about the habitats and ecology of these rare but inter- esting grasses. Hoover’s work on Orcuttia is largely taxonomic, and he includes a key to the four species and two varieties then known. Although he explored no further than gross morphology, he allowed himself the luxury of speculation regarding relationships. He states that the phylog- eny of Orcuttia is obscure, but that it is obviously related to Neostapfa. He indicates no dissatisfaction with the placement of these genera in the l’estuceae, but believes that it is a fallacy to relate them to Pappophorum. He makes the rather remarkable statement: “I am inclined to believe that Pleuropogon is involved in the ancestry of Orcuttia and Neostapfia.”
Crampton (1959) also believes that Orcuttia and Neostapfia are close- ly related, but he states correctly that they do not belong with the Festu- ceae. He believes that they are without a doubt the most unusual and rarest of the California grasses. He indicates that: “Their relationship to other grasses is not apparent and quite likely they represent a relict group, the ancestors of which are unknown.” He feels that they should be re- moved from the Festuceae and might well be considered as a separate tribe. Stebbins and Crampton (1961) include these two genera in the tribe Pappophoreae, associated with Pappophorum, Enneapogon, and Cottea.
The present paper is a review of the characteristics of these two un- usual genera along with new information, particularly from the caryopsis
1965 | REEDER: ORCUTTIEAE AND PAPPOPHOREAE 19
and embryo.! Comparison is made with typical members of the Pappo- phoreae, their apparent closest allies.
As Crampton (1959) has pointed out, the caryopses are more or less strongly laterally flattened in both Orcuttia and Neostapfia, and the hilum is rather conspicuous. Grains of Neostapfia are obovate and dark brown, whereas those of Orcuttza are oblong and somewhat translucent. In both genera the embryo is narrow and nearly as long as the seed. A noteworthy feature is that the base of the caryopsis is largely endosperm, the slender lower portion of the embryo occupying only a very small area (figs. 1-3). In members of the Pappophoreae, the caryopses are dorsally compressed or nearly circular in cross-section, the hilum is punctiform, and the lower part of the grain is made up almost entirely of the embryo (figs. 4-6).
Embryos of Orcuttia, Neostapfia, and members of the Pappophoreae are, in most respects, rather similar (figs. 7-16). In longitudinal sagittal section it can be seen that 1. the coleoptile is inserted at some distance above the point of divergence of the scutellar trace, and that there is a distinct “internode” between coleoptile and scutellum, 2. the lower part of the scutellum is free from the coleorhiza, there being a distinct cleft between the two, and 3. usually an epiblast is present, although in some species of Orcuttia it may be very small or lacking altogether. Evidence from the embryo thus supports strongly removal of all of these genera from the Festuceae, and even from the subfamily Festucoideae (Reeder, 1957). .
A more critical study of the embryos reveals that those of Orcuttia and Neostapfia share certain features which are not found among members of the Pappophoreae. In the former, the embryos are rather slender, the “internode” is often unusually long, and the tip of the coleoptile is greatly elongated and composed of solid tissue (figs. 7-11). It may also be of significance that the shape of the scutellum, as seen in transection, in embryos of Orcuttia and Neostapfia is rather different than in members of the Pappophoreae. |
The leaf epidermis in grasses often is a source of characters which are of significance in systematics. In connection with the present study, leaves of Neostapfia, all species of Orcuttia, and representatives of Pap- pophorum, Cottea, Enneapogon, Schmidtia, and Kaokochloa were ex- amined. It has been pointed out by Metcalfe (1960) and Stebbins and Crampton (1961) that both Neostapfia and Orcuttia have characteristic bicellular microhairs which apparently are not found among members of any other grass genus. In this investigation, these ‘““mushroom-button” microhairs (figs. 17, 18) were found to be present in all species of Orcut-
1 This research was supported in part by Grant G—9070 from the National Science Foundation. I am indebted to Beecher Crampton for seeds and specimens of Orcuttia and Neostapfia, to Margaret E. Speer for technical assistance, to Chester F. Natune- wicz for help with the Latin diagnoses, and to my wife, who prepared the illustra- tions and assisted in numerous other ways.
20 MADRONO [Vol. 18
tia and also in Neostapfia. These unusual hairs are correctly illustrated by Metcalfe, but the figures in Stebbins and Crampton are quite inac- curate. In the latter work, the terminal cell is shown much too large, and the hair appears to have no base, but to emerge from between two regular epidermal cells. Actually, each hair arises from a more or less cubical cell which is only slightly narrower than others with which it is associated. In addition to the bicellular hairs discussed above, the epidermis of Neo- stapfia bears distinctive ‘“‘crozier hairs” which seem to be unique. The stomata in both Neostapfia and Orcuttia are very similar and appear somewhat unusual. Rather than the lozenge shape (figs. 19, 20) which is common among non-festucoid grasses, these are often as broad, or even broader, than long. In Orcuttia siliceous cells are irregular in outline, some of them being rather distinctly dumbbell-shaped. No siliceous cells were found in Neostapfia.
The evidence presented above leaves little doubt that Neostapfia and Orcuttia are very closely related. Moreover, it is equally apparent that their affinities are not with members of the Festucoidea, but rather with the Eragrostoideae. As to a more precise placement, the only sug- gestion made thus far is the one by Stebbins and Crampton (1961), who include them with the Pappophoreae. In view of the differences already discussed, the assignment of Orcuttia and Neostapfia to the tribe Pappo- phoreae would not appear to constitute a particularly natural alliance. In Pappophoreae the caryopses are dorsally compressed, the hilum is punctiform, and the lower part of the broader embryo constitutes the base of the seed. We may add that the testa in Orcuttia and Neostapfia is rather loose, a characteristic which has not been noted among mem- bers of the Pappophoreae. The embryo itself shows distinctive features, the most noteworthy being the elongated solid coleoptilar tip in mem- bers of the former two genera. The epidermis of the leaf has rather dis- tinctive stomata, and microhairs which appear to be unique.
Not only do Orcuttia and Neostapfia differ rather sharply from mem- bers of the Pappophoreae in the features discussed above, but they are quite different morphologically as well. Plants of both genera are short- lived annuals of very restricted ranges, and they occupy distinctive habi- tats. The leaves are liguleless, there being no sharp distinction between sheath and blade. Although the lemmas are many-nerved, the nerves ord-narily do not extend into prominent awns, as in members of the Pappophoreae. In view of these differences, it appears to the writer that a more natural arrangement would be achieved if Orcuttza and Neostapfia were treated as constituting a distinct tribe. This is described below:
Orcuttieae Reeder, tribus nova. Gramina annua, viscida; foliis eligu- latis, sine distinctione in laminas et vaginas; spiculis plurifloris; lem- matibus multinerviis (plerumque 9-15); caryopsibus e latere compressis, hilo in basi amplo, embryone 34 vel plus longitudine caryopsidis; epi- dermide foliorum cellulis siliceis nullis vel irregulariter- vel halteriformis; mic:op:.is biceilulariis parvis, immersis, fungiculiformis.
1965] REEDER: ORCUTTIEAE AND PAPPOPHOREAE 21
Fics. 1-6. Caryopses in three views: left, face view; right, median sagittal sec- tion; above, transection through the coleoptile region: 1, Orcuttia pilosa, * 20; 2, O. mucronata, X 20; 3 Neostapfia colusana, X 15; 4, Pappophorum mucronu- latum, X 20; 5, Enneapogon desvauxii, * 30; 6, Cottea tappophoroides, X 20. Note in Figs. 1-3 (Orcuttieae) that the caryopses are laterally flattened, the lower part of the embryo barely reaches the base of the seed, and that there is a large basal hilum; in Figs. 4-6 (Pappophoreae) the caryopses are dorsally flattened or nearly circular in transection, and the lower part of the embryo constitutes the base of the seed.
Plants annual, viscid, leaves liguleless, without distinction into sheath and blade, spikelets several- to many-flowered, lemmas many-nerved (mostly 9-15), caryopses laterally flattened, hilum large, basal, embryo three-fourths or more the length of the caryopsis; siliceous cells of the epidermis absent or irregular to dumbbell-shaped, bicellular microhairs small, sunken, ‘“‘mushroom-button”’ shaped.
Included genera: Orcuttia Vasey; Neostapfia Davy.
Neostapfia thus far is known from a single species, V. colusana. Hitch- cock (1950) recognizes 4 species of Orcuttia, one of which, O. califor- nica, consists of 2 varieties in addition to the typical form. The recently described O. mucronata (Crampton, 1959) brings the total number of commonly recognized species to 5. Two of these, O. pilosa and O. tenuis, are so similar to O. californica in spikelet morphology that it might be more realistic to treat them as varieties of that species. Until more is known concerning these taxa, both cytologically and genetically, how- ever, it is perhaps most prudent to accept the standard treatments.
WD) MADRONO [Vol. 18
The literature on Orcuttia usually states that the rachilla is continuous or that the spikelet does not disarticulate between the florets. In this study it was found that when dry, the spikelets, indeed, do not break up. However, when fully mature, the florets readily break apart when soaked in water. This may be an adaptation of considerable importance to these species. During the time that the pools or “hog wallows” are dry, the florets remain on the plants and are thus not blown away by the wind. When the area becomes flooded, the spikelets readily disarticulate and the seeds are thus free for dispersal.
Two genera of grasses which appear to be similar ecologically to the Orcuttieae are Crypsis and Heleochloa. Lorch (1962), as the result of a critical study, has reunited these under Crypsis. He states: “Crypsis shows a very striking preference for localities swamped or at least tem- porarily inundated during winter, and its adaptation to such habitats where it often becomes a dominant component of the summer vegeta- tion.”’ In California, where they have been introduced, members of this genus are often found growing with the Orcuttieae. It is rather curious, moreover, that the caryopses and embryos of Orcuttia and Crypsis are very similar. We have examined C. niliaca and C. schoenoides (Heleo- chloa schoenoides) and find the caryopses in both to be laterally flat- tened, and to have an embryo nearly as long as the seed. These embryos have the same form as those of the Orcuttieae, even to the elongate solid coleoptilar tip. The leaf epidermis of Crypsis, however, bears normal eragrostoid hairs. Since the gross morphology is also quite different in Crypsis, the similar caryopses and embryos are probably best construed as evidences of parallelism, perhaps conditioned by the environment, rather than as suggesting close genetic affinity.
The tribe Pappophoreae is generally recognized by present-day agro- stologists as comprising the genera Pappophorum, Cottea, Enneapogon, and Schmidtia. In the older systems, the group was treated as a subtribe of the Festuceae, and this placement is followed in many manuals even today. Prat (1935) studied the leaf epidermis of representatives of the first two genera and concluded that they should be removed from the Festuceae and transferred to the subfamily Panicoideae. He was much impressed with elongate bulbous-tipped hairs which he found in Cofttea, and indicates that they are similar to those seen by Lohauss (1905) in Pappophorum commune F. Muell. Prat states that this indicates a close relationship between Cottea and Pappophorum. Actually, the plant which Lohauss studied was a species of Enneapogon, since the genus Pappo- phorum is confned to the Americas (Chase, 1946). A note under P. commune F. Muell. in Chase and Niles (1962) states: ‘““A group name for all the Australian species, the 4 of R. Br., 3 of Lindley, and P. coerules- cens Gaud. = Enneapogon.” Prat indicates that this unusual type of hair is pluricellular, but this is certainly incorrect. Caceres (1958) studied the Pappophoreae of Argentina, and reported similar hairs in both Enne- apogon and Cottea. He states emphatically, however, that these hairs are
REEDER: ORCUTTIEAE AND PAPPOPHOREAE oS
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LS I4
Fics. 7-16. Embryos in median sagitta! section and transection through the co!e- optile region: 7, Orcuttia californica var. inequalis; 8, O. pilosa; 9, O. greenez; 10, O. mucronata; 11, Neostapfia colusana; 12, Pappophorum pappiferum; 13, P. mu- cronulatum; 14, Cottea pappophoroides; 15, Enneapogon desvauxii: 16, Schmid ‘ta pappohoroides. Note in Figs. 7-11 (Orcuttieae) that the embryos are very similar except that 7 and 8 have no epiblast. Note particularly the elongated solid coleoptilar tip. In Figs. 12-16 (Pappophoreae) the embryos are very similar in longisection, but in transection 12 and 13 (subtribe Pappophorinae) have an embryonic leaf in which the margins meet but do not overlap, whereas in 14-16 (subtribe Coitinae)
the embryonic leaves have overlapping margins. These figures are not drawn to the same scale.
24 MADRONO [Vol. 18
not pluricellular, but are clearly bicellular. Jacques-Félix (1962) like- wise figures bicellular hairs of this type in Enneapogon and Schmidtia from Africa.
Tateoka et al (1959) discuss and figure these elongated bulbous- tipped bicellular hairs, which they state they found in Enneapogon, Schmidtia, and Pappophorum elegans. Stebbins and Crampton (1961) also report these distinctive hairs in the latter species, as well as in Cottea and Enneapogon. Actually, Tateoka et al, and also Stebbins and Cramp- ton are the victims of bad taxonomy. Pappophorum elegans Nees is, in fact, a species of Enneapogon, and was transferred to that genus by Stapf in 1907! Stebbins and Crampton have also made a faulty observation, since they state (p. 143): “The present writers have determined that in Cottea the stalk itself is composed of two cells, so that the hair is actually tricellular rather than bicellular.”” We have examined the epidermis of Cottea and find the hairs to be clearly bicellular as reported by Caceres and Tateoka et al. The reports by Prat (1935), Caceres (1958), and Jacques-Félix (1962), that siliceous cells in Cottea, Enneapogon, and Schmidtia are dumbbell-shaped, have been confirmed by the present study.
In the work referred to above, Caceres examined the epidermis of 5 species of Pappophorum. In all he found that the bicellular hairs were short and of the usual eragrostoid type. The siliceous cells of the lower epidermis were found to be saddle- (or double-axe) shaped, although on the upper epidermis they are cross- or dumbbell-shaped. These observa- tions have been verified in the present study (fig. 19). Other workers who have examined the epidermis of correctly identified Pappophorum species have reported these same characteristics. It is seen, therefore, that on the basis of epidermal characters, Pappophorum, is somewhat different from other members of the Pappophoreae.
A further distinction between Pappophorum, on the one hand, and Cottea, Enneapogon, and Schmidtia on the other, is revealed by a study of the embryo. In longisection these are almost identical and are much like those of other grasses of the chloridoid-eragrostoid alliance. A tran- section through the coleoptile region of Pappophorum, however, shows a festucoid leaf—one in which the margins meet but do not overlap (figs. 12, 13), whilst in the others, this leaf is of the panicoid type, having distinctly overlapping margins (figs. 14-16).
In view of the differences found in both epidermis and embryo be- tween Pappophorum and the other genera, it might seem appropriate to consider that they represent two distinct tribes. Although there is, in- deed, some justification for such a treatment, in the writer’s opinion these genera are so similar morphologically that they are better kept together. Such an alliance is not an unnatural one since all have a similar basic chromosome number and in other respects clearly have eragrostoid affinities. The one gross morphological character which appears to sep- arate Pappophorum readily from the others is its 1-nerved glumes; in the others the glumes are several-nerved. Although these differences do not
1965] REEDER: ORCUTTIEAE AND PAPPOPHOREAE 25
Fics. 17-20. Portion of the lower epidermis from a leaf, * 225. 17, Neostapfia colusana; 18, Orcuttia mucronata; 19, Pappophorum bicolor; 20, Cottea pappo- phoroides. Note distinctive stomata and “mushroom-button” microhairs in Neostapfa and Orcuttia and the crozier hairs in the former. Also that Pappophorum has double- axe shaped siliceous cells and typical chloridoid microhairs, in contrast to Cottea in which the siliceous cells are dumbbell-shaped and the microhair has a slender elon- gated basal cell and a bulbous apical cell. appear to be sufficient for tribal separation, subtribes seem clearly to be
indicated.
Tribe PAPPOPHOREAE Kunth, Rév. Gram. 1:82. 1829.
Subtribe PAPPOPHORINAE. Pappophorum Schreb. ex Vahl, Symbol. Bot. 3:10. 1794.
Subtribe Cottinae Reeder, subtribus nova. Gramina perennia vel rare annua; spiculis plurifloris, glumis et lemmatibus multinerviis, aliquot vel omnibus nervis lemmatum in aristas extendentibus; embryone cum formula P + P P; epidermide foliorum cellulis siliceis halteriformis, pilis bicellulariis cum cella apicali glandulosa et plusminusve bulbosa, cella basali angusta, plerumque elongata.
Plants perennial or rarely annual; spikelets several-flowered, glumes and lemmas many-nerved, some or all of the nerves extending into awns; embryo with the formula P + P P; leaf epidermis with dumbbell-shaped siliceous cells and bicellular hairs with a glandular, usually bulbous, apical cell and a slender, usually much elongated, basal cell.
Included genera: Cottea Kunth, Rév. Gram. 1:84. 1829; Enneapo- gon Desv. ex Beauv., Ess. Agrost. 81, pl. 16, fig. 11. 1812, Jour. de
26 MADRONO [Vol. 18
Bot. 1:70. 1813; Schmidtia Steud. in J. A. Schmidt, Beitr. Fl. Capver- dischen Ins. 144. 1852 (nom. conserv.); Kaokochloa deWinter, Bothalia 7:479, 1961.
This latter genus, which was recently described from Africa, clearly belongs to the subtribe Cottinae also. Through the kindness of Dr. de- Winter we have recently received specimens-and mature seeds of this grass for our studies. The leaf epidermis has dumbbell-shaped siliceous cells and the same type of elongated bicellular hairs which are character- istic of Cottea, Enneapogon, and Schmidtia. Moreover, the embryo, as in those genera, is of the formula P + P P, the embryonic leaf having over- lapping margins. Dr. deWinter (in personal communication) states that he considers Kaokochloa to be a member of the Pappophoreae, closely allied to Schmidtia.
Pommereulla L.f., of India and Ceylon, is sometimes included in the Pappophoreae. Pilger (1954), however, treated it as a subtribe of the Festuceae, separating it from the Pappophoreae, which he considered a distinct tribe, but still assigned to the Festucoideae. Metcalfe (1960) reports chloridoid anatomy, and Bor (1960) has created for the genus a separate tribe, the Pommereulleae. Despite the anatomical similarity between Pappophorum and Pommereulla, in view of the very different gross morphology, the disposition by Bor seems fully justified. The tribe clearly belongs with the Eragrostoideae, however, and not with the Pooideae, as indicated by Bor.
Until recently it was thought that the elongate bulbous-tipped bicellular hair, shown here to be characteristic of the subtribe Cottinae, did not occur outside the Pappophoreae. Nicora (1962), however, has found hairs of a similar type in Neeragrostis reptans (Michx.) Nicora (Era- grostis reptans Michx.). Such unusual hairs immediately suggest a rela- tionship between Neeragrostis and Cottinae, but in gross morphology they are very different. In the former the plants are dioecious annuals with 1-nerved glumes and 3-nerved awnless lemmas. Although future study may reveal some close affinity, the evidence now available does not appear to warrant assignment of Neeragrostis to Pappophoreae, subtribe Cottinae.
List OF SPECIES STUDIED AND SOURCES OF MATERIAL
The embryos were secured from mature seeds. These were embedded in paraffin, sectioned at 10 microns with a rotary microtome, and stained with saffranin and fast green. Leaf epidermis material was prepared by first boiling the leaf, then placing it on a slide and scraping away the tissue down to the epidermis. Portions of epidermal tissue thus obtained were stained in methylene blue and ruthenium red, dehydrated, and mounted in Permount.
Crypsisniliaca Fig. & DeNot. California, Belluet & Henson, Oct. 1944 (NY). Crypsis niliaca Fig. & DeNot. California, Belluet & Henson, Oct. 1944 (NY). C. schoenoides (L.) Lam. Pennsylvania, Small, Sept. 1897 (YU).
1965] REEDER: ORCUTTIEAE AND PAPPOPHOREAE HG
Enneapogon desvauxit Beauv. Mexico, Hidalgo, Reeder & Reeder 1629 (YU); Durango, Reeder & Reeder 2603 (YU).
E. flavescens (Lindl.) Burb. Australia, Queensland, Blake 8054 (US).
E. glumosus (Hochst.) Marie & Weiller. West Africa, French Sudan, Hagerup 441a (US).
Kaokochloa nigrirostis deWinter. South West Africa, Kaokoveld, deWinter & Leistner 5679A (YU).
Neostapfia colusana (Davy) Davy. California, Solano Co., Crampton 5421 (YU); 5406 (YU); Stanislaus Co., Crampton 5026 (YU).
Orcuttia californica Vasey var. inaequalis (Hoover) Hoover. California, Merced Co., Crampton 5433 (YU); 5040 (YU).
O. california var. viscida Hoover. California, Sacramento Co., Crampton 5125 (YU).
O. greenei Vasey. California, Butte Co., Crampton 5069 (YU).
O. mucronata Crampton. California, Solano Co., Crampton 5113 (YU); 5057 CU):
O. pilosa Hoover. California, Stanislaus Co., Crampton 5431 (YU).
O. tenuis Hitchc. California, Shasta Co., Crampton 5078 (YU); 5079 (YU).
Pappophorum bicolor Fourn. Texas, Reeder & Reeder 2899 (YU).
P. mucronulatum Nees. Arizona, Goodding 124-47 (YU).
P. pappiferum (Lam.) Kuntze. Paraguay, Puerto Casado, Hartley SH121 (US).
Schmidtia pappophoroides Steud. Africa, Rhodesia, Hitchcock 24170 (US).
Yale University, New Haven, Connecticut
LITERATURE CITED
Bor, N. L. 1960. The grasses of Burma, Ceylon, India and Pakistan. Pergamon Press, New York.
Caceres, M.R. 1958. La anatomia foliar de las ‘““Pappophoreae” de Mendoza y su valor taxonomico. Revista Argent. Agron. 25:1-11.
CuasE, A. 1946. Enneapogon desvauxii and Pappophorum wrightii, an agrostological detective story. Madrono 8:187-189.
, and C. D. Nixes. 1962. Index to Grass Species. 3 vols. G. K. Hall and Co.,
Boston, Mass.
Crampton, B. 1959. The grass genera Orcuttia and Neostapfia: A study of habitat and morphological specialization. Madrono 15:97-110.
Davy, J. B. 1898. Stapfia, a new genus of Meliceae, and other noteworthy grasses. Erythea 6:109-113.
. 1899. Concerning Stapfia Davy. Erythea 7:43.
Hitcucock, A. S. 1950. Manual of the grasses of the United States. Ed. 2, revised by Agnes Chase. U.S. Dept. Agr. Misc. Publ. 200.
Hoover, R. F. 1941. The genus Orcuttia. Bull. Torrey Club 68:149-156.
JacquEs-FEitx, H. 1962. Les Graminées d’Afriaue Tropicale. I. Généralités, Classi- fication, Description des Genres. Institut de Recherches Agronomiques Tropicales. Paris.
Lonwauss, K. 1905. Der anatomische Bau der Laubblatter der Festucaceen und dessen Bedeutung fur die Systematik. Bibliot. Bot. 13(63) :1-114.
Lorcu, J. 1962. A revision of Crypsis Ait. s. 1. (Gramineae). Bull. Res. Counc. Israel 11D:91-116.
Mason, H.L. 1957. A flora of the marshes of California. Univ. Calif. Press, Berkeley
Metcatre, C. R. 1960. Anatomy of the Monocotyledons. I. Gramineae. Oxfcrd Univ. Press, London.
Mouwz, P. A. 1959. A California Flora. Univ. Calif. Press, Berkeley.
Nicora, E. G. 1962. Revalidacion del genero de Gramineas “Neeragrostis” de la flora Norteamericana. Revista Argent. Agron. 29:1-11.
28 MADRONO [Vol. 18
PILGER, R. 1954. Das System der Gramineae unter Ausschluss der Bambusoideae. Bot. Jahrb. 76:281-384.
Prat, H. 1935. Contribution a 1’ étude systématique et histologique des Festucées. Bull. Soc. Bot. Fr. 82:498-506.
REEDER, J. R. 1957. The embryo in grass systematics. Am. Jour. Bot. 44:756-768.
STEBBINS, G. L. and B. Crampton. 1961. A suggested revision of grass genera of temperate North America. Recent Advances in Botany (IX Int. Bot. Congress, Montreal, 1959) 1:133-145.
TATEOKA, T., S. INOUE, and S. Kawano. 1959. Notes on some grasses. IX. Systematic significance of bicellular microhairs of leaf epidermis. Bot. Gaz. 121:80-91.
VAsEY, G. 1886. A new genus of grasses. Bull. Torrey Club 13:219.
REVIEWS ] Flora of Missouri. By JuLtian A. STEYERMARK. lxxxiv + 1725 pp., 390 plates (line drawings), 2409 distribution maps, 1 text figure. Iowa State University Press, Ames. 1963. $18.50.
The appearance of this large, detailed flora (it weighs over five and one-half pounds) is a tribute to the author’s persistence and industry. During the quarter century that it has been under way he has been living most of the time outside the state. Except at rare intervals it has been a project for evenings, weekends, and holidays.
In addition to keys, the volume presents the 2438 species now reported as occurring, or having once occurred, within the state of Missouri. Of these, 60 are ferns or fern allies, 6 are gymnosperms, and 2,372 are flowering plants. Between one-fourth and one-fifth of the species are introduced. These go all the way from species so well established that they might almost be taken for natives, to waifs collected in St. Louis freight yards by Viktor Muehlenbachs. (A minor but consistent error is the citation of this name without the terminal ‘s’.) The degree to which they are becoming established is usually indicated; attention is called to those few which are proving troublesome.
Whether we like it or not these introduced species are playing an ever increasing role in our vegetation. The completeness and precision of Steyermark’s record will make it a baseline for future investigations of this basic biological problem.
Another of the author’s special interests has been minor variations in flower color, pubescence, and growth habit; the flora lists 297 forms, many of them previously named by him. As basic data about variation, these are a valuable record, particularly when effort has been made to make it so complete.
In citing forms and varieties Steyermark follows the machinery set up by the International Code. If a form or variety is cited then the specific name is repeated as a pigeonhole for the specimens not otherwise catalogued at that level. This makes for a complicated text and takes up space. Steyermark reports it as a scheme easily dealt with editorially.
An introductory chapter gives expositions of the thinking behind his editorial decisions. It includes a chronological sketch of the principal Missouri collectors and the areas in which they were active. He estimates their total output as over 200,000 numbered collections.
A chapter on “Flora and Vegetation” has a summary of the basic physiography of the state followed by his delimitation of its plant regions: Southeastern Lowlands, Ozark, Prairie. He describes characteristic habitats in each with further details about salt licks, saline springs, upland sinkhole ponds and loess hills. He lists those species known from bordering states for which there is a sporting chance they might be found in Missouri.
1965 | REVIEWS 29
There is a large detailed county map arranged for ready reference, an index of common and scientific names, a supplement of 17 additions while the book was in the press, and tables totaling genera, species, varieties, forms, and hybrids, family by family, with various summaries.
Line drawings are provided for all but a few species. These are from a variety of sources and are of uneven merit. For virtually all the species there is a map of distribution by counties. In addition to herbarium specimens these are based on careful field surveys. There is the scientific name recognized by the author and when this differs from commonly accepted ones, these synonyms are given. Flowering seasons are summarized by months from herbarium records. The range outside the state is indicated.
For many of the species there is further information about a variety of matters; details of the habitat; the uses made of certain p!ants by wild animals, pioneers, Indians and modern man; poisonous properties; plants causing hay fever and dermatitis. Those he has himself used for food are indicated and there is even an occasional recipe. For several aquatics he describes the morphological effects of changing water levels. For various species he has interesting comments about their behavior in the wild-flower garden he long maintained at his home in northern Illinois. — EpcAr ANDERSON, Missouri Botanical Garden, St. Louis.
Families of Flowering Plants of Southern Africa. By HERBERT PARKES RILEY. xviii + 269 pp. University of Kentucky Press, Lexington. 1963. $14.00.
An unusual taxonomic publication has been prepared by Dr. Riley, Professor and Head of the Botany Department at the University of Kentucky. It is one of the first regional floras to contain rather extensive listings of chromosome numbers, many published here for the first time. Many of the chromosome counts were made by the author from collections of African plants, or from chromosome counts made during one year spent in South Africa on a Fullbright Fellowship.
The person unfamiliar with South African plants gains much insight into the floristics of that part of the world through the introductory chapter which includes 3 maps showing respectively the political, physiographic and vegetative features of the area. The first 7 photographs in the book show different South African plant associations. Two excellent photographs of cone bearing male and female plants of Encephalartos kosiensis are included as illustrations of one type of gymnosperm from the coastal subtropical forest near the Indian Ocean (on page 8, not 81, as indexed). The last 6 photographs in the book are almost superfluous, representing introduced plants around the settings of dwellings, including introduced plants such as Bougainvillea, Carica papaya, Pinus, and Jacaranda. The remainder of the 144 photographs are illustrations of plant families in Southern Africa, some in cultiva- tion, but many in their native habitats. Unfortunately the photographs are not indexed at the back of the book and are only found within the text where specific genera are described in each family. No indication of the family is given by the photograph. If all plates illustrating one family had been listed just below the title of each family, or some similar means of relating the illustrations to families, the illustrations would prove more useful to persons unfamiliar with African families. Since all the photographs are in color, arranged 6 to a page, generally with high quality photography and color reproduction, the color photographs alone are worth the price of the book.
Most American botanists are familiar with the Englerian system of classification, and some have become acquainted with the Bentham and Hooker system, the latter more widely used by the British. Dr. Riley has used a modified system, based on John Hutchinson’s herbaceae-lignosaea system. Although Hutchinson’s system gives insight to some relationships that other systems have ignored with its much-branched trees from two sources, it is not well suited to a linear sequence. It is difficult to
30 MADRONO [Vol. 18
see that new understanding o} .elationships is gained, for instance, by the following sequence of families (pp. 38-44): Escalloniaceae, Corncaeae, Araliaceae, Hamameli- daceae, Myrothamnaceae, Buxaceae, Bruniaceae, and Salicaceae !
It is unfortunate that a book of plant families is not indexed for the scientific names of all the families. The extensive index is 15 pages, 5 columns wide, but contains only generic or for the most part common and Afrikaans names of plant families. Thus Ginger and Ginger family are listed, but not Zingiberaceae. Barberry family is listed, but not Berberidaceae. Where does one look for Hamamelidaceae (not indexed), if one does not know the common name?
Much useful information is contained about genera, such as their uses by natives and Europeans living there, and the ornamental value of the plants. The book uses generic concepts very close to those of Phillips in his Genera of South African Plants. A check shows that the families are very closely matched, and Riley has added at least one family: Taccaceae——RoBErT J. Ropin, California State Polytechnic Col- lege, San Luis Obispo.
Vascular Plants of the Pacific Northwest. By C. LEo HitcHcock, ARTHUR CRON- QUIST, MARION OwnBEy, and J. W. THompson. Part 2, vi+ 597 pp., illustrated. University of Washington Press. 1964. $15.00.
Publication of this second part of the series marks the completion of the treat- ment of the dicotyledons. My earlier comments on the work (Madrono 15:74-76; 17:90-91) are still applicable to the present part, which covers Salicaceae through Crassulaceae. Preceding the main text are two keys to the families of dicotyledonous plants in the region covered. One is a synoptical key intended to aid in an under- standing of Cronquist’s phylogenetic arrangement of families (which is not followed in this work) and the other is an artificial key which almost everyone will use for identifying families. All families in this part but Salicaceae were treated by Hitchcock. The same high standards of previous contributions are maintained. The only new taxa described are several varieties but numerous changes in taxonomic rank are made. In spite of ever-increasing inflation, the University Press has managed to keep the price per page of the series remarkably stable—RoOBERT OrNpDUFF, Department of Botany, University of California, Berkeley.
Generic Names of Orchids. By RicHARD Evans SCHULTES and ARTHUR STANLEY PEASE. xvi + 331 pp. Academic Press, New York, London. 1963. $12.00.
This work represents a valuable addition to the literature relating to the Orchidaceae. It will be useful to anyone who is interested in this vast family, whether professional or amateur. The book is beautifully printed, ornamented by numercus illustrations and vignettes, and will be an embellishment to any library.
The production is exactly what it claims to be —a dictionary that explains tne etymological history of the 1250 generic names of Orchidaceae and the morpnoiogical or other characteristics of each plant that led to the choice of the generic name. Also included are the original Greek or Latin components of the names, a.ong with the geographic distribution and tribal or subtribal classification of the genus shown together with the place of first publication of the epithet. A lengthy introduction reviews the Orchid Famiiy taxonomicaliy, his.oricaliy, and ethnoboianically, aad explains briefly the importance of and rules governing nomenclature.
Since the authors state that it was not their purpose to make this dictionary in any way a taxonomic treatment, they have precluded any criticism that poe be {crchcoming due to this deletion.
Both authors are outstanding scholars in their respective fields at Harvard University: Schultes in economic and systematic botany, and Pease, an emeritus
1965] NOTES AND NEWS 31
professor of Latin who also excels in botany as an avocation. These two formed a superior association to produce this volume. Fortunately, to supplement their manu- script, the authors were able to make use of numerous illustrations that have accrued through the years to the Oakes Ames Orchid Herbarium of Harvard, some of which have previously been used to delineate the various orchid floras that have been produced in the Ames Herbarium.
The scholarship evidenced by this work makes it a very worthwhile contribution, and it should be available to all who are interested in orchids. A leisurely perusal! through this attractive book will be an enlightening experience for anyone.— Donovan S. CoRRELL, Texas Research Foundation, Renner.
NOTES AND NEWS
TRANSLOCATION HETEROZYGOSITY IN CROTON CALIFORNICUS.—A morphologically relatively uniform population of Croton californicus var. tenuis (Wats.) Ferg. from Point Dume, Los Angeles Co., California, was examined cytologically from buds of male plants growing on the sand dunes near the edge of the bluff or in the rock crevices just below (LA 33482). Buds were fixed in Carnoy solution and examined after squashing in acetocarmine.
Meiosis of seven plants could be examined and gave the following results: one plant consistently formed 14 pairs and was apparently structurally homozygous; five plants regularly showed 12 pairs and a ring of 4 chromosomes, indicating struc- tural heterozygosity for one reciprocal translocation; one plant showed 10 pairs, a ring of 4, and a chain of 4 chromosomes, indicating that it was heterozygous for two translocations.
The chromosome number (2n = 28) is new for the genus Croton. Previous counts, for C. glandulosa, C. monanthogynis, and C. palmeri by Perry (see Darlington and Wylie, Chromosome Atlas of Flowering Plants, 1956), have all been 2n = 16. The present count suggests, therefore, a second basic number of 7.
Of special interest, however, is the high frequency of translocation he‘erozygotes. Although the present sample is small, it suggests that translocation heterozygosity may be an integral part of the adaptive genetic system of the population. It is also interesting to note that pollen fertility is apparently extremely variable. Some indi- viduals had as low as 30 percent visibly good pollen when stained with cotton blue in lactophenol, whereas others had as high as 96 percent stainable poilen. The vari- ation did not appear, however, to be directly correlated with structural heterozy- gosity. Further study of the cytology and genecology of this polytypic species should be rewarding. Such a study has been initiated by S. Smith-White and Albert W. Johnson.
These observations were made in the Botany Department of the University of Caifornia, Los Angeles, during my tenure as a Rockefeller Foundation Fellow. I am indebted to Harlan Lewis and Kunyama Mathew for their helpful advice and assistance. —JERZY SZWEYKOWSKI, Adam Mickiewicz Univ., Poznan, Poland.
Unitep States NATIONAL HERBARIUM.—During the month of June 1965, the Department of Botany of the U.S. National Museum, which includes the U.S. Na- tional Herbarium, will be moving from its present quarters in the Smithsonian Insti- tution Building into the new west wing of the Museum of Natural History Bui'ding. The new space will provide the department with two complete floors for specimen storage, laboratories, libraries, offices, rooms for visiting botanists, and herbarium maintenance facilities.
It is estimated that preparation for the move will occupy a 3-month pericd, and an equivalent amount of time will be needed to arrange our property and resume activities after the move has taken place. Regretfully, it will be necessary for the best interests of the Herbarium and our correspondents throughout the world to request a moratorium on the receipt and shipment of herbarium specimens during this period.
32 MADRONO [Vol. 18
We ask that between April 1, 1965 and October 31, 1965, no specimens be sent to us, nor that we be requested to ship specimens on loan. Specimens received during this period will be retained for processing until after October 31, 1965; requests for specimens also will be held for action until after this rate——WILLIAM L. STERN.
Crocip1uM IN SAN Luts Opispo Co., CALIFORNIA—Crocidium multicaule Hook. (Compositae-Senecioneae), an annual herb of uncertain affinities which is the only species of its genus, is reported to range from Vancouver Island southward to the Mount Hamilton Range of central California. Now it can be reported some 130 mi to the south, on the basis of the following collections: damp sandy soil on north slopes of narrow canyons above semi-permanent streams, the Bear Trap, foot of Castle Mountain, on the headwaters of San Juan Creek, San Luis Obispo Co., Hardham 11001 (DS), Feb. 8, 1964; Hardham 11007 (Cas, DS, OSC, RSA, US), March 8, 1964. Since Ornduff (Leafl. West. Bot. 9:84. 1960) recently pointed out that forms of this species with epappose ray florets occur throughout its range, with the possible exception of the populations in the central Sierra Nevada foot- hills of California, we asked Mrs. Hardham to obtain a sample from her isolated population on which to score this character. Her no. 11007 thus consisted of 214 individuals obtained at intervals along a transect of about a mile in length. All 214 of these individuals, as well as all 13 in no. 11001, had pappose ray florets. Whether this is the result of the origin of this colony from a few original plants with pappose ray flowers, or the genes controlling this characteristic are linked with others physiologically favorable for survival at this southern station, cannot be determined at present. — PETER H. RAvEN and Tuomas R. MERTENS, Division of Systematic Biclogy, Stanford University.
A REED COLLEGE HERBARIUM. — Plant taxonomists working on the flora of the Pacific Northwest are generally unaware of the existence of a small herbarium housed in the Department of Biology at Reed College, Portland, Oregon. The purpose of this note is to describe its contents and to bring it to the attention of interested botanists. The herbarium contains an extensive collection of vascular plants of Multnomah Co. made by Richard van Dersal in the late 1920’s; plants of the Mount Hood region collected by Kate C. Rogers in 1938; specimens of the native, weedy, and cultivated plants of the college campus collected by Una V. Davies in 1937-1938; and a number of species from Ocracoke Island, North Carolina, deposited by Katherine Rondthaler in 1951. There are also small collections of Oregon mosses gathered by Carra Horsfall and by Margery Washburn, and miscel- laneous vascular plants collected in various parts of Oregon by Donald B. Lawrence. The majority of these collections was made in connection with the preparation of baccalaureate theses, none of which has been published. In addition to these col- lections by former students there are numerous plants collected in Oregon and Washington by two pioneer northwestern botanists—Thomas Howell and William Cusick. Finally, there are several sheets bearing collections made elsewhere in the United States by such botanists as Canby, Chickering, Hasse, Hapeman, Hyams, Kellerman, and Vasey. This herbarium is, therefore, of moderate historical interest. Because it houses a number of unicates from northwestern Oregon it should certainly be consulted by botanists working on the floristics of this portion of the state. — RoBERT ORNDUFF, Department of Botany, University of California, Berkeley.
DA pri. 7
Ta 7
MADRONO
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VOLUME 18, NUMBER 2 APRIL, 1965
Contents
PAGE
ALLOLEPIS: A NEW SEGREGATE OF DISTICHLIS (GRAMI- NEAE), Thomas R. Soderstrom and Henry F. Decker 33
NATURAL HYBRIDIZATION BETWEEN CUCURBITA DIGI- TATA AND C, PALMATA, W. P. Bemis and Thomas W.
W hitaker 39 | MANZANITA CHAPARRAL IN THE SANTA ANA MOoun- TAINS, CALIFORNIA, R. C. Wilson and R. J. Vogl 47 | | Reviews: Flora Europaea (Wallace R. Ernst) 62
Notes AND NEws: ARCEUTHOBIUM DOUGLASII IN NE- VADA AND WYOMING, Frank G. Hawksworth; WASHOE PINE ON THE BALD MOUNTAIN RANGE, CALIFORNIA,
W. B. Critchfield and G. L. Allenbaugh; NEw PuBLti- CATIONS 63
A WEST AMERICAN JOURNAL OF BOTANY
| ?>UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
MADRONO-
A WEST AMERICAN JOURNAL OF BOTANY
Entered as second-class matter at the post office at Berkeley, California, January 29,
1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price
$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, Life Sciences Building, University of California, Berkeley, California 94720.
BOARD OF EDITORS
EpGAR ANDERSON, Missouri Botanical Garden, St. Louis LyMAN BENSON, Pomona College, Claremont, California HERBERT F, COPELAND, Sacramento College, Sacramento, California Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California Mivprep E. Matutas, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley Marion OwneBEY, Washington State University, Pullman REED C. RoLiins, Gray Harbarium, Harvard University Ira L. Wiccins, Stanford University, Stanford, California
Editor—JoHn H. THomas Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Annetta Carter, Department of Botany, University of California, Berke- ley. First Vice-President: Winslow R. Briggs, Department of Biological Sciences, — Stanford University. Second Vice-President: Arthur R. Kruckeberg, Department of Botany, University of Washington. Recording Secretary: Mary L. Bowerman, De- partment of Botany, University of California, Berkeley. Corresponding Secretary, Margaret Bergseng, Department of Botany, University of California, Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State College.
SODERSTROM & DECKER: ALLOLEPIS 33
ALLOLEPIS: A NEW SEGREGATE OF DISTICHLIS (GRAMINEAE)
THOMAS R. SODERSTROM AND HENRY F. DECKER
In a recent study to establish the relationships of the grass Reeder- ochloa (Soderstrom and Decker, 1964), the authors studied the morphol- ogy and anatomy of species of the similarly dioecious genus Distichlis. Attention was drawn in particular to D. texana, a rather robust species which has been collected only rarely from the Big Bend country of Texas to El Paso and south into Coahuila, Chihuahua, and Durango, Mexico. The species was first described by Vasey (in Coulter, 1890) as Poa texana, based on a collection made by G. C. Nealley in 1887, apparently in Pre- sidio Co., Texas. In the description Vasey states that all of the specimens are male and questions whether or not the species might be dioecious. Later (1893) when both male and female plants were available to him he redescribed it as Sieglingia wrightii, pointing out his earlier misinter- pretation. The name Sieglingia is now valid only for the single species, S. decumbens. What Vasey evidently had in mind when he used the name was a group of species which now are included in the genera Tvidens and Erioneuron, in which no dioecious members are found. Nine years after being first described, the species found its way into yet another genus, this time Dzstichlis, to which it was transferred by Scribner (1899). Scribner felt it belonged in Distichlis because of “‘the dioecious habit, the character of the inflorescence, the rigid subcoriaceous glumes, long- exserted styles protruding from the apex of the floret, and grains en- closed in the coriaceous base of the palea.” This disposition was accepted by Hitchcock in the first edition of Manual of the Grasses of the United States and in the 1950 edition revised by Agnes Chase.
Since the time of Scribner and Hitchcock, characters of leaf anatomy have come to play an increasingly important role in elucidating the true relationships of grasses. Studies of Reederochloa, for example, showed that the leaves of that genus possess peculiar bicellular microhairs in which the enlarged bases are actually sunken in the epidermis. Such hairs are found also in Monanthochloé and, with the exception of D. texana, in all species of Distichlis. The genera with this type of microhair share many other anatomical as well as morphological characters, such as the dioecious habit and coriaceous many-nerved lemmas and glumes, which unite them into a close, natural association. Distichlis texana (for the genus and indeed for this whole alliance) is striking because its bicellular microhairs are not sunken but are of the normal eragrostoid type, similar to those found in Eragrostis, Muhlenbergia, Sporobolus, etc. This dis- similarity of bicellular microhairs prompted a more thorough look at D. texana and the other members of the genus. Our studies revealed that this species differs remarkably in fundamental anatomical and morpho-
Mapbrono, Vol. 18, No. 2, pp. 33-64. May 26, 1965.
34 MADRONO [Vol. 18
logical characters and has indeed been misplaced even in Distichlis. Now, after 65 years in the latter genus, we choose to remove it and recognize it separately as a new genus, Allolepis, the fourth generic name to be associated with this interesting grass.
Allclepis Soderstrom & Decker, gen. nov. Perennis dioica stolonifera; culmi ascendentes vel decumbentes, vaginae forte costatae, glabrae; ligula brevis, ciliata; rami paniculae appressi vel ascendentes, plerumque ad basin floriferi. Planta masculina: Spiculae ovato-lanceolatae vel lineares, stramineae, illustres; glumae ovatae, hyalinae, glabrae; gluma prima 1l-nervia, quam secunda paulo brevior; gluma secunda 1(-3)- nervia; lemmata forte 3-nervia, glabra, illustria; palea lemma aequans vel quam lemma paulo longior, linearis, glabra; antherae 3, flavae. Planta feminea: Spiculae ovato-lanceolatae, teretes vel paulo compressae, strami- neae; glumae ovatae, coriaceae, glabrae, marginibus membranaceis; gluma prima quam secunda paulo brevior et angustior, forte 1-nervia (interdum cum ampliis nerviis indistinctis); gluma secunda 3-nervia (interdum cum ampliis nerviis indistinctis); lemmata forte 3-nervia (interdum cum ampliis nerviis indistinctis), glabra, ovata, coriacea, mar- ginibus scariis; palea quam lemma paulo brevior, a latere visa supra versus angusta et linearis, basin versus curvata, glabra; lodiculae 2, cuneatae.
Type species: Allolepis texana (Vasey) Soderstrom & Decker, comb. nov. Poa texana Vasey, Contr. U.S. Natl. Herb. 1:60. 1890. Sieglingia wrightit Vasey, Contr. U.S. Natl. Herb. 1:269. 1893. Distichlis texana (Vasey) Scribn., U.S. Dept. Agri. Div. Agrost. Cir. 16:2. 1899, based on Poa texana Vasey.
The following description of A. ¢exana is intended to supplement pre- vious descriptions of this taxon which have been scanty and based on fewer specimens than those available at this date: Perennial, dioecious, stoloniferous, the stolons glabrous, strongly ribbed, 1-4 mm broad, to 25 cm long; culms loosely ascending or spreading as stolons, 10-70 (25-65) cm long (from point of rooting to base of inflorescence), gla- brous; nodes 3-several, glabrous; lower sheaths shorter than the inter- nodes, strongly ribbed, glabrous; blades flat when dry, sometimes in- volute toward the tip, to 30 or more cm long, 2.5-6 (4-5) mm wide, glabrous above and below, becoming scaberulous toward the tip, the edges scabrous; ligule short-ciliate from an inconspicuous membranous base, 0.5-1.4 (0.5-1) mm long; inflorescence a panicle, 3-23 (10-17) cm long, 1-6 (1-3) cm wide, consisting of stiffly appressed or ascending branches 3—6 cm long, usually floriferous to the base, the panicles con- taining as many as 70 spikelets; lodicules 2, cuneate in outline. Male plant: Spikelets ovate-lanceolate to linear, stramineous, lustrous, 9-23 (10-15) mm long, 3-8 (3-6) mm wide, up to 20 florets per spikelet, apparently not disarticulating; glumes broadly ovate, hyaline, glabrous, the midnerve scabrous; first glume a little shorter than the second, ca. 4—5 mm long, 1-nerved; second glume ca. 1 mm longer than the first,
:
1965 ] SODERSTROM & DECKER: ALLOLEPIS 35
1 (—3)-nerved; lemmas of lower florets 5—5.5 mm long, strongly 3-nerved, glabrous, lustrous; palea equal to or a little longer than the lemma, linear, glabrous, the keels minutely ciliolate; anthers 3, yellow, 3—3.5 mm long. Female plant: Spikelets ovate-lanceolate, terete or only slightly com- pressed, stramineous, 1—2 cm long, 2.5—3.5(3) mm wide, with up to 8 or 9 closely imbricate florets per spikelet, apparently not disarticulating; glumes broadly ovate, coriaceous with broad scarious margins, glabrous, the midnerve scabrous; first glume a little shorter and narrower than the second, ca. 7-9 mm long, strongly l-nerved (up to 4 or 5 additional faint nerves may be present) ; second glume 3-nerved, with an additional 1 or 2 pairs of faint nerves sometimes present; lemmas of the lower florets 7.5-10 mm long, strongly 3-nerved, glabrous, midnerve scabrous above, broadly ovate, coriaceous with irregular scarious margins; palea a little shorter than the lemma, narrow above and strongly bowed out below, the margins overlapping, glabrous, the keels ciliolate.
Specimens examined. UniTEp States, Texas. Without precise locality (Presidio Co. ?): Nealley s.n., in 1887 (Isotype, US: ¢ plants, basis of Poa texana Vasey). Brewster Co.: Castalon, Silueus 648 (TAES, TEX, US). Jeff Davis Co.: Brown 53-240 (TEX: cultivated in Univ. Texas grass garden); Brown s.n., 25 July 1949 (TEX); Brown s.n., 1 Aug. 1950 (TEX: grown in Univ. Texas grass garden, cyto- logical voucher) ; Hinckley 4571 (US); Hinckley & Hinckley 316 (US); Limpia Canyon, Nealley s.n., in 1892 (US); Warnock 7942 (LL, SMU, TEX); Warnock 8039 (LL, SMU); Warnock 10187 (LL, SMU); Valley of the Limpio, Wright 2038 (US: @ plant, basis of Szeglingia wrightii Vasey). Presidio Co.: Nealley 136 (TAKES) ; Nealley 137 (NY, US); near Presidio, Nealley s.n., in 1892 (US) ; Silveus 737 (US) ; Warnock 158 (TEX, US).
Mexico. Chihuahua: 5 km west of Cd. Camargo, Harvey 1412 (US). Coahuila: Saltillo, Palmer 507 (NY, TAES, US). Durango: Torreon, Hitchcock 7541 (US).
A photograph of a single plant of Allolepis clearly showing its strong stoloniferous habit is given by Silveus (1933). There is also an excellent illustration of the habit and of the male and female spikelets (Scribner, 1901), and a sketch of the female panicle and spikelet (Hitchcock, 1950).
Allolepis can at once be distinguished from Distichlis by the absence of scaly rhizomes and presence of long, thick stolons. Distichlis rarely produces narrow stolons but, when found, they are in addition to the scaly rhizomes which are so characteristic of the genus. Beetle (1943) has even recognized as a variety of D. spicata some California plants with stolons, D. spicata (L.) Greene var. stolonifera Beetle, but these are apparently produced only under certain conditions and do not seem to be correlated to any degree with other characters. The length of the stolons in Allolepis is hardly apparent on herbarium specimens which represent only segments of the whole. Silveus (1933) states that they are about 10 ft long and Barton Warnock (pers. comm.) of Sul Ross State College, Alpine, Texas, has observed in the field that some of them are easily 50 ft long. The blades of Distichlis are narrow, involute, pungent and arranged on the culm in an obvious distichous manner, hence the generic name. In contrast, those of Allolepis are flat, longer and broader, and not arranged in such a distinct distichous fashion.
36 MADRONO [Vol. 18
The male and female spikelets of Distichlis are about equal in size within the same species: each is composed of several florets, the lemmas of which are coriaceous and many-nerved. In Allolepis, the female spike- lets are about twice as large as the male spikelets and the texture and nerves of the lemmas in each are distinct. Lemmas of the male spikelets are thin in texture and strongly 3-nerved whereas those of the female spikelets are coriaceous and 3-nerved but occasionally with an additional pair of faint nerves. The 3-nerved condition of the lemmas is quite un- like that of the Distichlis-Monanthochloé-Jouvea-Reederochloa alliance in which the lemmas of both sexes are consistently coriaceous and many- nerved. The palea offers a further distinction between the two genera, for the keels of the palea in Distichlis are winged but in Allole pis they are not.
The dissimilarity of lemmas between the sexes of Allolepis has sug- gested its new name, a synthesis of the Greek words allo (different) and lepis (scale, or lemma).
The most prominent difference in the leaf epidermis between Allole pis and Distichlis is the microhair—normal eragrostoid in the former (fig. 1A), bulbous-based and sunken in the latter (fig. 1B). The shape of the siliceous cells is irregular (“‘potatolike”) in Distichlis (fig. 1D), pre- dominantly dumbbell-shaped in Allolepis (fig. 1C). The epidermal sur- face of Distichlis is rough and irregular due to the presence of many papillae (fig. 1B) while that of Allolepis has few or no papillae and hence is relatively smooth. A few minor differences are also encountered in the leaf cross section. Arm cells (fig. 1A) are conspicuous in each bundle of an Allolepis leaf but not obvious, when present, in Distichlis. The bulli- form cells of Adlolepis (fig. 1A) are relatively large in tall girders and the outer bundle sheath is elongate in outline and complete on most of the bundles. The bulliform cells of Distichlis (fig. 1B) are relatively small and in short girders and the outer bundle sheath is generally round in outline and complete only on the smaller bundles.
From published chromosome reports, no cytological distinction can be made between the two genera. Brown (1951) has listed a count of 2n—=40 for Distichlis texana (Allolepis). Similar counts have also been given for D. spicata from California material (Stebbins & Love, 1941) and Argentine material (Rahn, 1960), and for D. stricta from Canadian ma- terial (Bowden, 1960) and California material (Stebbins & Love, 1941). The basic number of x10 indicates these genera do not belong in the traditional tribe Festuceae but rather to the eragrostoid complex, as sub- stantiated by the leaf anatomy and other studies (Soderstrom and Deck- er, 1964).
Distichlis, Reederochloa, Monanthochloé, and Jouvea are plants of saline habitats. The latter two are strictly littoral or maritime; of the former two, Reederochioa occurs in alkaline inland flats, and Déstichlis occurs both along the coasts and in saline areas of the interior. Informa- tion regarding the habitat of Allolepis has been difficult to obtain, espe- cially since so few botanists have collected it. To our knowledge the only
1965 ] SODERSTROM & DECKER: ALLOLEPIS Sif
838 | Jo |
Fic. 1. A, B, Transverse sections of grass leaves; C, D, portions of the bands of siliceous cells over the costal regions on the epidermis of grass leaves; A, C, Allolepis texana (Silveus 648); B, D, Distichlis spicata (Reeder & Reeder 3074). The illus- trations were done with the aid of a microprojector and are at various magnifica- tions: ar, arm cell; bu, bulliform cell; mi, bicellular microhair; obs, outer bundle sheath; pa, papilla.
living botanists in Texas who have collected the genus are Barton War- nock and W. V. Brown. Warnock remarks (pers. comm.)! that he found it growing in igneous sandy soil in a broad flat among Hymenoclea mono- gyra, and also in silty soil at the mouth of Cibolo Creek between the town
1 We wish to acknowledge the information supplied to us by Barton H. Warnock.
38 MADRONO [Vol. 18
of Presidio and the international bridge where it was rather abundant. Silveus (1933) gives cultivated and waste meadowlands or sand flats as the habitat. It has even been cultured successfully in ordinary loam in the University of Texas grass garden. Such information, plus the scanty data available from herbarium labels, does not indicate that Allolepis is a plant of alkaline areas. If this is the case, it presents us with yet an- other major feature to distinguish it from Distichlis and related genera.
Voucher specimens. Leaf cross sections and epidermis of the following specimens were examined in this study. A taxonomic revision of Distichlis is much needed and such a study would probably reduce the number of valid species; nevertheless the material which we examined does at least represent the genus as it occurs in North and South America.
Allolepis texana (Vasey) Soderstrom & Decker. UNITED STATES. New Mexico (or western Texas): Wright 2038, ¢ (US). Texas: Nealley, in 1889, ¢ (US); Nealley, in 1892, 2 (US); Castalon, Silveus 648, 9 (US). Mexico. Coahuila: Saltillo, Palmer 507, in 1898, 9 (US).
Distichlis hirta Phil. CurLe. Batuco, Philippi 377, 2 (US).
D. humilis Phil. Peru. Chuquibambillo, Hitchcock 22450, 2 (US).
D. marginata Phil. CHILE. Atacama, Werdermann 997, 2 (US).
D. palmeri (Vasey) Fassett ex I. M. Johnston. Mexico. Sonora, Bacigalupi 2873, d (US).
D. scoparia (Kunth) Arech. ARGENTINA. Neuquen, Senn 4329, 9 (US).
D. spicata (L.) Greene. UNITED StTATEs. Connecticut: Lighthouse Pt., Reeder & Reeder 2756 (YU). Massachusetts: Cape Cod, Reeder & Reeder 252, 6 & 2 (YU). Mexico. Michoacan, Reeder & Reeder 3074, 6 & 9 (YU).
D. stricta (Torr.) Rydb. UnitEp States. Utah: Reeder & Reeder 1858, 6 & @ (YU).
D. thalassica (H.B.K.) Desv. ARGENTINA. Mendoza, Bartlett 19530, 6 (US).
D. viridis Phil. Cu1Le. Atacama, Werdermann 431, & (US)
Department of Botany, Smithsonian Institution, Washington, D.C., and Depart- ment of Botany and Bacteriology, Ohio Wesleyan University, Delaware, Ohio
LITERATURE CITED
BEETLE, A. A. 1943. The North American variations of Distichlis spicata. Bull. Torrey Club 70:638-650.
Bowpen, W. M. 1960. Chromosome numbers and taxonomic notes on northern grasses. II. Tribe Festuceae. Canad. Jour. Bot. 38:117-131.
Brown, W.V. 1951. Chromosome numbers of some Texas grasses. Bull. Torrey Club 78:292-299.
Coutter, J. M. 1890. Upon a collection of plants made by Mr. G. C. Nealley, in the region of the Rio Grande, in Texas, from Brazos Santiago to El Paso County. Contr. U.S. Natl. Herb. 1:29-65.
Hircucock, A.S. 1950. Manual of the grasses of the United States. Ed. 2, revised by Agnes Chase. U.S. Dept. Agr. Misc. Publ. 200.
Raun, K. 1960. Danish scientific investigations in the Argentine under the auspices of the Fundacién Williams, Buenos Aires Chromosome numbers in some South American angiosperms. Bot. Tidsskr. 56:117-127.
ScRIBNER, F. Lamson. 1899a. American grasses II. U.S. Dept. Agr. Div. Agrost. Bull. 17: June. Rev. ed. May 22, 1901.
. 1899b. New species of North American grasses. U.S. Dept. Agr. Div. Agrost. Circ. 16.
1965] BEMIS & WHITAKER: CUCURBITA 39
Sitveus, W. A. 1933. Texas grasses. San Antonio, Texas.
SopERSTROM, T.R. & H.F. DECKER. 1964. Reederochloa, a new genus of dioecious grasses from Mexico. Brittonia 16:334-339.
STEBBINS, G. L. & R. M. Love. 1941. A cytological study of California forage grasses. Amer. Jour. Bot. 28:371-382.
Vasey, G. 1893. Descriptions of new or noteworthy grasses from the United States. Contr. U.S. Natl. Herb. 1:267-280.
NATURAL HYBRIDIZATION BETWEEN CUCURBITA DIGITATA AND C. PALMATA
W. P. BEMIS AND THOMAS W. WHITAKER
Cucurbita digitata Gray and C. palmata Wats. are xerophytic, per- ennial species indigenous to the desert regions of southwestern United States and northwestern Mexico.! According to Bailey (1943) C. digitata is found in New Mexico, Arizona, sparingly in southeastern southern California (eastern Riverside County) and in Sonora, Mexico. It is doubt- ful that C. digitata is indigenous west of the Colorado River. In Cali- fornia it is found in limited roadside areas, suggesting that it was prob- ably introduced by camper-tourists. Cucurbita palmata is found in south- ern California from the Mohave Desert southwards in interior valleys and at low elevations. It also occurs in western Arizona and northern Baja California, Mexico. In the area adjacent to the Arizona-California border and directly east of the Colorado River the species are sympatric.
Each species has 20 pairs of chromosomes (McKay, 1931). They also have similar growth patterns. The root systems have large fleshy storage roots which enable the plants to survive prolonged periods of high tem- peratures and extremely arid conditions. Vine growth is rapid under favorable conditions of moisture and temperature. Adventitious roots are produced at intervals on the nodes of the runners, and under favorable moisture conditions these roots are able to develop into tuberous roots which when separated from the mother vine will establish another plant. This asexual method of propagation produces a clone with the individual plants grouped in colonies. Cucurbita digitata and C. palmata are prolific producers of fruit and seed. The fruit is a relatively hard-shelled, round pepo, two to three inches in diameter, and contains from 200 to 600 seeds. Reproduction by seed, however, is probably limited because of low seed and seedling survival in the environments to which these species are adapted.
1 We wish to acknowledge with thanks the assistance of F. D. Cole, University of Arizona, and Paul D. Hurd, Jr., University of California, Berkeley, for directing our attention to localities where hybrids were likely to occur. Dr. Hurd and Dr. Edgar Anderson have read and made suggestions for improvement of the manuscript.
40 MADRONO [Vol. 18
METHODS AND RESULTS
The most conspicuous morphological difference between the two species is leaf shape. The leaf of C. digitata has five lanceolate lobes with sinuses developed to the base of the leaf veins. The leaf of C. palmata has five pal- mate lobes with sinuses developed to approximately one-third the length of the leaf veins. The first four to six seedling leaves of both species are similar to C. palmata.
Since leaf shape is the most striking morphological difference between these species a method of measurement was developed to describe the leaves numerically. The measurement that was found to reflect the leaf diffrences was the ratio of the length to the width of the central palmate or lanceolate lobe and the two adjacent lobes. The length of the lobes was measured from the petiole attachment to the tip of the leaf blade. The widths of the lobes were measured at their widest point not including appendages which were present on some leaves. The widest point on the lobes of C. palmata leaves was at the base of the sinuses.
Figure 1 shows three diagrammatic Cucurbita leaves superimposed. The solid leaf represents C. digitata and has a length to width ratio of ten. The open leaf represents C. palmata, with a length to width ratio of two. The shaded leaf is intermediate between C. digitata and C. pal- mata with a length to width ratio of five and represents an F, hybrid between the two species.
GREENHOUSE OBSERVATIONS. As a part of a general study of the genus Cucurbita, reciprocal crosses between C. digitata and C. palmata were made. The F, and F. generations were fertile, indicating these species are cross compatible.
Plants from self-pollinated seed of two C. digitata lines and one C. pal- mata line, along with the reciprocal F, hybrids were grown in the green- house at the University of Arizona in the fall of 1963. The C. digitata lines originated from fleshy roots collected 5 miles and 22 miles N of Tucson, Arizona. The C. palmata line came from a fleshy root collected in the Yuma Valley, Arizona, adjacent to the Colorado River and 5 miles N of the Mexican border. Leaf ratio measurements for these seven plants are given in Table 1. Leaf ratios for the two C. digitata lines were 9.42 + 1.66 and 8.34 + .72 respectively, while for C. palmata the ratio was 2.28 + .27. For the F; hybrids of C. digitata 2 x C. palmata ¢ the leaf ratio were 4.62 + .39 and 4.89 + .39; for the reciprocal cross the ratios were 3.87 + .79 and 3.65 + .73 (fig. 2). It is evident that the F, hybrids are intermediate between the parent species. Data from the hybrids sug- gest a possible cytoplasmic effect on leaf ratios as they tend to show an effect of the female parent. More information on this question is neces- sary before a statement can be made. The point is that both kinds of hybrids do have intermediate leaf ratios.
FIELD OBSERVATIONS. The range of C. digitata and C. palmata over- lap in the vicinity of the Arizona-California border roughly parallel to the Colorado River from near the vicinity of Davis Dam in southern
1965] BEMIS & WHITAKER: CUCURBITA 4\
Fic. 1. Diagrammatic sketch representing a leaf of C. digitata (solid), C. palmata (open), and an intermediate (shaded).
Nevada, on the north to San Luis, Sonora, Mexico, on the south. In this general area one might expect natural hybridization between the two species to occur. Our attention was directed to some plants which ap- peared to be suspiciously like hybrids in the area north of Dateland, Ari- zona, by F. D. Cole and to the area between Quartzite, Arizona, and Yuma, Arizona paralleling Arizona Highway 95 by P. D. Hurd, Jr. Col- lections were made in the above two areas, and in a third area where Highway 95 crosses the Gila River, about 12 miles NW of Yuma and 6 miles E of the Colorado River. At each location herbarium specimens (ARIZ) were collected and leaf ratios were obtained.
Dateland locality. This locality is in the bed of the Gila River north of Dateland, Arizona. It is approximately 70 miles E of the Colorado River (Arizona-California border). The Gila River at this point forms a flood plain about two miles wide. The soil of the plain is a silty loam and it supports a dense growth of mesquite. The samples were taken from the south edge of this flood plain on Nov. 21, 1963. The soil of the flood plain had at one time been disturbed by heavy equipment operating at
MADRONO [Vol. 18
St a A ees RE Oe
as NS)
ii ES
Fic. 2. Two representative leaves of C. ding (left) ; Fi (C. palmata X C. digi-
tata) center; and C. digitata (right). Hybrids were produced from controlled pol- linations.
a nearby sand and gravel quarry. The action of this equipment may have created favorable conditions for seed germination and seedling establish- ment of Cucurbita seed produced locally or from fruit transported down the Gila River during periods of flooding.
A particularly dense population of Cucurbita occupied an area approxi- mately 90 * 150 feet (fig. 3). From this area fourteen individual col- onies were sampled. The leaf ratio data are given in Table 1 under “Dateland.” Two plants from this area had leaf ratios indicative of “pure” C. digitata, and one plant was identified as C. palmata based on leaf ratio data. The remaining eleven plants had intermediate leaf ratios varying from 5.60 + .72 to 2.97 + .38 (fig. 4). The range in leaf ratios of intermediates suggests that they include segregates beyond the Fy generation or perhaps backcrosses, or if they are representative of F, hybrids, that the parental species were not homozygous for factors con- trolling leaf shape. Plants with intermediate leaf ratios are undoubtedly the result of natural interspecific hybridization.
Yuma locality. A sample was taken where U.S. Highway 95 crosses the Gila River. This area is approximately 6 miles E of the Colorado River and 64 miles down river from the Dateland sampling area. The Cucurbita at this location were not particularly dense and only seven colonies were sampled on Dec. 19, 1963, in approximately a quarter mile length of the Gila River. Leaf ratio data for the seven plants from this area are given in Table 1 under “Yuma.”
1965] BEMIS & WHITAKER: CUCURBITA 43
Fic. 3. Field photograph from Dateland, Arizona, showing runners from plants with digitata-like and palmata-like colonies intertwined. The pole is marked in dms.
Three of the plants had leaf ratios indicative of C. palmata, a fourth was like C. palmata but showed some indication of C. digitata contamina- tion since it had a leaf ratio of 2.85 + .27. Two plants were intermediate, and one plant was principally C. digitata but showed some evidence of C. palmata contamination with a leaf ratio of 7.14 + .87. These limited data suggest that the dominant species in this area is C. palmata and that fruit and seed of C. digitata segregates were probably transported down the Gila River during periods of flooding.
Five locations on Highway 95. Eighteen colonies were sampled on Dec. 3, 1963, at 5 locations extending from 18 to 55 miles S of Quartzite, Ari- zona, on Highway 95. The colonies were located either in the stream bed or along the banks of dry washes that drain westward into the Colorado River. Abundant late summer and fall rains had produced excellent vege- tative growth, but flowers and fruits were absent from most colonies.
Of the 18 colonies sampled, one appeared to be “pure” C. palmata, and 5 were C. digitata or digitata-like (Table 1). The remainder were inter-
[ Vol. 18
~
MADRONO
44
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1965] BEMIS & WHITAKER: CUCURBITA 45
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Fic. 4. Leaves from natural hybrids, Dateland; 1 (upper left) is “pure” digitata; 6 (upper right) is “pure” palmata; the remainder, 2-5, are intermediates.
mediates, but a majority of these plants were more like C. digitata than C. palmata. There was no noticeable trend in favor of one or another of the two species from north to south, nor did the individual sampling areas appear to be homogeneous for leaf type.
DISCUSSION
Two perennial, xerophytic species of Cucurbita, C. digitata and C. pal- mata, were successfully hybridized under controlled conditions. They produced fertile F; and F, progenies, indicating a marked degree of com- patibility. Reasoning from this information, it was considered probable that natural hybridization might occur in locations where the ranges of the two species overlap. The area immediately east of the Colorado River, adjacent to the Arizona-California border and stretching north to south for a distance of 200 miles seems the most likely place for hybridization to occur. Before the search for natural hybrids could be profitably pur- sued, it was necessary to have a reliable morphological marker to indi- cate putative hybrids. The two species are nearly similar in vegetative, fruit and seed morphology, but there is a striking difference in one vege- tative character, leaf shape. A ratio of the lobe length to lobe width was devised to describe numerically the leaves of the parent species and the hybrid between them.
Since these species are cross compatible and monoecious, we would anticipate considerable hybridization in nature if a suitable pollinator
46 MADRONO [Vol. 18
were present. That such a pollinator is present was established by Hurd and Linsley (1964) in their scholarly studies of the squash and gourd bee of the genera Xenoglossa and Peponapis. The authors report that one species, P. timberlake, is limited to C. digitata and C. palmata for its pollen and nectar supplies. Apparently P. timberlakei cannot use the pollen of other Cucurbita species even when available within its range. The infrequent, but severe rainstorms that cause flooding and erosion in this area could produce niches free from competition where young Cucur- bita seedlings would have an excellent chance to become established. Thus four elements for successful natural hybridization are present: 1, compatible species; 2, overlapping ranges; 3, an efficient pollinator; and 4, a niche for the survival of the hybrids.
Using the leaf ratio index as a criterion it is apparent from field col- lections that natural hybridization exists along a narrow front roughly parallel to the Colorado River in western Arizona. We are suggesting the two species migrated into the area and subsequently hybridized. This area of the Colorado River basin is traversed by the Gila River and nu- merous dry washes all flowing westward toward the Colorado River. Violent summer and fall rains, although relatively infrequent, produce much flooding and erosion and would be capable of transporting fruits and seeds from south and east central Arizona toward the Colorado.
SUMMARY
1. F, and Fy, hybrid populations of C. digitata « C. palmata were produced in the greenhouse. These hybrids were fertile, indicating the two species are compatible.
2. Cucurbita digitata and C. palmata are sympatric along a narrow front parallel to and directly east of the Colorado River, extending from the vicinity of Davis Dam in southern Nevada to San Luis in northern Sonora, Mexico.
3. The most conspicuous morphological difference between the two species is the structure of the leaf. Cucurbita digitata has five lanceolate lobes with the sinuses extending to the base of the leaf. Cucurbita pal- mata has five palmate lobes with sinuses extending about one-third the distance to the base. A ratio was devised to reflect this difference: thus C. digitata has a length width ratio of 10; C. palmata—2; and the Fy hybrid—5.
4. In the areas where the two species are sympatric we have sampled colonies in three locations and determined by leaf length to width ratios that natural hybridization has occurred.
5. We have shown that the elements for successful natural hybridiza- tion are present, i.e. compatible species, overlapping ranges, a specific pollinating insect, and niches for the survival of the hybrid seedlings.
University of Arizona, Tucson, Arizona and U.S. Department of Agriculture, La Jolla, Calif.
1965 | WILSON & VOGL: CHAPARRAL 47
LITERATURE CITED
BariLey, L.H. 1943. Species of Cucurbita. Gentes Herb. 6:267-315.
Hurp, P. D., Jr., and E. G. Linstey. 1964. The squash and ground bees—Genera Peponapis Robertson and Xenoglossa Smith—inhabiting America north of Mexico (Hymenoptera: Apoidea). Hilgardia 35:375-477.
McKay, J. W. 1931. Chromosome studies in the Cucurbitaceae. Univ. Calif. Publ. Bot. 16:339-350.
MANZANITA CHAPARRAL IN THE SANTA ANA MOUNTAINS, CALIFORNIA
R. C. WILSON AND R. J. VOGL
A quantitative study of Arctostaphylos glandulosa Eastw. chaparral associations in the Santa Ana Mountains was undertaken during the years 1962 and 1963. This phytosociological study correlates numerical data with information on climate, soils, slope exposure, and elevation to estimate the stability, site preferences, and species associations for Arcto- staphylos glandulosa.
Little consideration has been given to the quantitative aspects of chaparral associations dominated by Arctostaphylos species. Major at- tention has been given to Adenostoma fasciculatum H. & A. chaparral, a more widespread and lower elevation community. Extensive studies of Arctostaphylos chaparral have possibly been neglected because it is more restricted than Adenostoma chaparral, occurs at higher elevations which are often more difficult to reach, and is usually considered im- penetrable. Since Arctostaphylos chaparral generally occurs at higher elevations, it does not present as many erosion, watershed, or fire prob- lems and is considered of less economic, agricultural, or wildlife im- portance than chaparral types bordering grazing land, cultivation, or settlement.
The classic work of Cooper (1922) in central California is a floristic study of California chaparral (Delting, 1961). Cooper observed and commented on slope, exposure preferences, behavioral characteristics, humus accumulation, and evaporation ratios of Arctostaphylos associa- tions as compared with other chaparral associations. Cooper cited 19 species of Arctostaphylos as components of chaparral and classified Arctostaphylos site requirements as intermediate between Adenostoma fasciculatum and coniferous forest.
Following Cooper’s work, a number of studies were made of chaparral ecology, again mainly in 4denostoma communities. These include reports by Bauer (1936), Miller (1944; 1947), Sampson (1944), and Wells (1962). Sampson quantitatively studied chaparral succession in central and northern California. He indicated that pine suffered when associated with Arctostaphylos due to the greater potential of Arctostaphylos to recover after fire by resprouting or by seed germination. Some Arcto-
48 MADRONO [Vol. 18
staphylos species survive fire by possessing fire-resistant basal burls while other species rely entirely on seeds that depend upon high tem- peratures for germination (Jepson, 1916). More recently, Wells (1962) correlated mosaic chaparral patterns with geological mosaics and dis- turbances around San Luis Obispo, California. He noted that the Arcto- stapyhlos species A. obispoensis Eastw., A. rudis Jeps. & Wiesl., and A. morroensis Weisl. & Schreib. had definite soil preferences for serpen- tinite and unconsolidated siliceous sand and succeeded on soils where other plants were unable to survive.
Specific reference to the chaparral in the Santa Ana Mountains was made by Pequegnat (1951) who noted the plant species present, plant- animal associations, and weather conditions in the area. Three Arcto- staphylos species are listed as present; A. glauca Lindl., A. glandulosa, and A. pringlei Parry var. drupacea Parry. He reported that Garrya flavescens Wats., G. fremonti Torr., and A. glandulosa form an associa- tion above 4000 ft elevations.
Abrams (1951), McMinn (1939), and Wieslander and Schreiber (1939) have presented taxonomic descriptions of Arctostapyhlos. These descriptions define the distribution of species and give clues to possible site preferences.
THE Stupy AREA
The Santa Ana Mountains are located in the southern California Peninsular Ranges, a complex of inland and coastal ranges, which extend through Riverside, Orange, and San Diego counties (Jahn, 1954). The Santa Anas are located 10 miles east of the City of Santa Ana, and are about 40 miles long extending southeasterly from the Santa Ana River. The entire range covers approximately 400 square miles varying in width from 4 to 13 miles. The crest with an average height of 3500 ft, is located some 20 to 25 miles from and parallel to the Pacific Ocean. Santiago Peak is the highest point (5860 ft) and Modjeska Peak is the only peak over 5000 ft in the Santa Ana chain.
The Santa Ana Mountains were elevated along the Great Elsinore Fault during the Pleistocene Period. The range has been broken into secondary blocks by further faulting processes, contributing to their rugged character. This range is tilted southwestward with the crest lying near the eastern limits. The east face is a battered fault scarp, drained by short streams. Santiago Creek is the main stream draining the moun- tains to the west (Hinds, 1952).
Pequegnat (1951) reports that the basement core is partly metamor- phosed sediments and granitic intrusives. Along the north and south edges of the range, the ends of the basement core are surrounded by a complex of sedimentary clays, shales, and sandstones.
Close proximity to the sea moderates any climatic extremes in this winter-wet, summer-dry region. Pequegnat (1951) reports that 90% of the precipitation occurs from December to April. Snowfall is moder-
1965 ] WILSON & VOGL: CHAPARRAL 49
ate with the 3500 ft level as its lower limit. Most precipitation occurs within a belt between the elevations of 2000 and 4000 ft. Fog is common on the Pacific coast slopes in the spring and early summer. The prevail- ing winds that emanate from the southwest and west are interrupted only by hot, dry “Santa Ana”, foehn winds which occasionally sweep down from the high desert.
The vegetational composition is as varied as the undulating topog- raphy. Major forms include coniferous forest of Pseudotsuga macro- carpa (Vasey) Mayr. and Pinus coulteri D. Don., and chaparral. The main vegetational type is chaparral, dominated by Adenostoma fascicu- latum at lower elevations and by Arctostaphylos glandulosa at higher elevations.
The sample stands for this study were scattered over a 12 mile range within the boundaries of the Cleveland National Forest in the central part of the Santa Ana Mountains, and confined to areas of Arctostaphy- los chaparral. The U.S. Forest Service, Santa Ana Ranger Station fire records show numerous fires for the Santa Ana Mountains since 1922, but fire disturbance has been absent from the study areas for at least 25 years.
METHODS
Sample sites were selected using the following criteria: 1. a lack of unnatural disturbances such as bulldozing, road building, and cutting, 2. no natural catastrophe such as fire within the last 25 years, 3. relative- ly homogenous areas of vegetation, 4. each site uniform in its elevation, slope, exposure, and soils, and, 5. a presence of mature, reproducing plants on established sites. Specimens of all species were collected and classied according to Munz (1959).
Each site was sampled quantitatively with a 100 ft line-intercept and a one-fortieth acre quadrat. The total number of feet intercepting the line was recorded for each species, including overlapping species. From this, relative per cent cover on the 100 ft line was calculated.
Each one-fortieth acre quadrat was placed adjacent to the line-inter- cept in a rectangle, 100 « 10.9 ft. This quadrat was broken into four contiguous sections each 25 *« 10.9 ft. The number of individuals of each species was counted in each quadrat. An individual plant was any plant possessing a burl or trunk distinct from other burls or trunks. Relative per cent frequency, relative per cent dominance, and density of individuals per acre were calculated from the quadrat data. Values of relative per cent cover, relative per cent density, and relative per cent frequency were totaled to obtain importance values (I.V.) (Bray and Curtis, 1957). I.V.’s were derived for each species in each stand. Stands with similar floristic contents, as reflected by species I.V.’s, were classed together in association groups.
Additional measurements included height and number of stems per Arctostaphylos individual, elevation, exposure, degree of slope, and soil horizon depths. These measurements, in addition to observations on dis-
50 MADRONO [Vol. 18
TABLE 1. AVERAGE IMPORTANCE VALUES (I.V.) FOR THE SPECIES IN EACH ARCTOSTAPHYLOS ASSOCIATION
Association Groups
ak Manzanita Manzanita “Pure”
Species Manzanita Chamise Coulter Pine Manzanita Garrya fremontii 12 Lonicera johnstonii 3 Rhus diversiloba 13 Rhus ovata 4 Rhus laurina 0.4 Ribes indecorum 9 Prunus tlicifolia 0.1 Yucca whipplei 5 Pinus coultert 63 Cercocarpus betuloides 15 1 Quercus dumosa, wislizenit 153 3 3 Heteromeles arbutifolia Z 2 4 Adenostoma fasciculatum 14 104 20 24 Arctostaphylos glandulosa 13 174 200 265 Ceanothus crassifolius,
leucodermis, tomentosus 7 13 12 8
turbance, fire, growth habits, climate, and other meteorological data (Pe- quegnat, 1951), were used to determine factors influencing the distribu- tions of plant species and associations.
The data were further analyzed by a mathematical summary which utilizes a coefficient of similarity (Bray and Curtis, 1957). With this co- efficient, I.V.’s of the species from each stand were compared to those in all of the other stand sampled. In this way, stands were arranged ac- cording to their similarity or dissimilarity in species composition.
RESULTS
The results obtained are based on the data summarized from 40 one- fortieth acre quadrats and 40 one-hundred ft line-intercepts.
Importance values (I.V.) were obtained for each species. These are summarized in Table 1 for stands classified into four Arctostaphylos chaparral associations. The associations; oak-manzanita, manzanita- chamise, manzanita-Coulter pine, and “pure” manzanita are named for the two leading plant dominants. Manzanita stands dominated only by Arctostaphylos glandulosa were called “pure” manzanita, since they were practically pure stands.
Table 1 lists the relative importance of each species in its association group and serves as a species presence list.
The two oak species in the oak-manzanita association are Quercus dumosa Nutt. and Q. wislizentt A. DC. This association group is the most heterogeneous in composition with 13 species represented. When compared to the two dominants, Quercus and Arctostaphylos, the other species are of minor importance. Among the minor species are Cercocar- pus betuloides Nutt., Adenostoma fasciculation, Rhus diversiloba T.& G., and Garrya fremonti Torr.
1965] WILSON & VOGL: CHAPARRAL oH
TABLE 2. AVERAGE NUMBER OF INDIVIDUALS PER ACRE OF THE IMPORTANT SPECIES IN EACH ARCTOSTAPHYLOS ASSOCIATION
Association Groups O
ak Manzanita Manzanita “Pure”’ Species Manzanita Chamise Coulter Pine Manzanita Garrya fremontit 160 Lonicera johnstoni 20 Rhus diversiloba 80 Rhus ovata 16 Rhus laurina 4 Yucca whip plei 7 Pinus coulteri 320 Cercocarpus betuloides 80 6 Quercus dumosa, wislizenit 1040 5 19 Heteromeles arbutifolia 4 2 28 Adenostoma fasciculatum 16 1308 24 160 Arctostaphylos glandulosa 800 2142 2443 3520 Ceanothus crassifolius,
leucodermis, tomentosus 12 255 118 28
Totals L232 3717 2730 3736
In the manzanita-chamise association, few other species are present other than occasional individuals of Ceanothus spp. (including C. crassifolus Torr., C. leucodermis Greene, and C. tomentosus var. oliva- ceus Jeps.), and Yucca whipple: Torr. This association is the most xeric and occupies the lowest elevations studied.
The manzanita-Coulter pine association is dominated by Arctostaphy- los glandulosa followed to a lesser extent by Pinus coulteri and Adeno- stoma fasciculatum. Again, a few individuals of Ceanothus are present.
The “pure” manzanita association is represented by the lowest number of species (four). The Arctostaphylos I.V. is high when compared with those of the other three species, of which Adenostoma is the most com- mon.
The I.V.’s of Arctostaphylos increase as it is associated with Quercus, Adenostoma, Pinus coulteri, and “pure” stands respectively.
The number of individuals per acre for each species is summarized in Table 2 for the four associations. The total number of individuals of all species per acre ranges from 2232 in oak-manzanita to 3736 in “pure” manzanita. Manzanita-chamise and “pure” manzanita have approxi- mately the same total number of individuals per acre. The density of Arctostaphylos, like the I.V.’s, increases as it is associated with Quercus, Adenostoma, Pinus coulteri, and “pure” stands. When the average num- ber of stems per Arctostaphylos glandulosa individual (five) is multiplied by the number of A. glandulosa individuals per acre for each association, the range is from 4,000 stems per acre in oak-manzanita to 17,610 stems per acre in “pure” manzanita. The numbers per acre among the codomi- nants is higher for Quercus in the oak-manzanita than it is for Arcto- staphylos. With the exception of Adenostoma in the manzanita-chamise association, the other codominants are low when compared to Arcto-
§2 MADRONO [Vol. 18
TABLE 3. AVERAGE ABSOLUTE PER CENT COVER FOR THE SPECIES IN EACH ARCTOSTAPHYLOS ASSOCIATION
Association Groups
Oak Manzanita Manzanita “Pure”’ Species Manzanita Chamise Coulter Pine Manzanita Garrya fremontii 3 Lonicera johnstonii 0.3 Rhus diversiloba 2 Rhus ovata Z Rhus laurina 0.4 Ribes indecorum 1 Prunus ilicifolia 0.1 Yucca whipplei 0.1 Pinus coulteri 17 Cercocarpus betuloides 7 0.1 Quercus dumosa, wislizenii 59 2 1 Heteromeles arbutifolia 0.4 Adenostoma fasciculatum 30 3 4.6 Arctostaphylos glandulosa 26 67 80 94 Ceanothus crassifolius, 1 5 2 0.6 leucodermtis, tomentosus Bare ground 6.6 7 9.3 2.6
staphylos. Subdominant species such as Ceanothus attained their great- est density in manzanita-chamise stands and to a lesser degree in man- zanita-Coulter pine stands. Garrya fremonti, another minor species, was fairly dense in the oak-manzanita association.
Table 3 is a summary of absolute per cent cover for the species in the four associations. This is calculated by dividing the total feet of cover (intercept on the 100 ft line) for each individual by the total length of the transect taken and is expressed as per cent. Absolute per cent cover can be used as an indicator of dominance for each species. These figures are proportionately comparable to the density and I.V. data. An addi- tional figure is the per cent bare ground or percentage of the transect devoid of plant cover. The manzanita-Coulter pine association has the greatest percentage of bare ground and the “pure” manzanita association has the least with a 94% cover of Arctorstaphylos glandulosa and a 2.6% intercept of bare ground. The bare ground figures for all four associa- tions are low (less than 10%) verifying that plant growth in dense in all four associations.
The 40 stands, each designated as an association, were arranged graph- ically on a target diagram to summarize and compare slope and exposure data (fig. 1). The preferences for slope show that Arctostaphylos glandu- losa formed solid stands on gentle slopes (10-30°), mixed with Adeno- stoma fasciculatum on a wide variety of slopes, and occurred with Pinus coulteri on steep slopes (28-35°). Quercus species and Pinus coulteri locate most successfully on the steeper slopes (30-40°) (figs. 2 and 3).
Slope is related to soil depth and the presence of organic matter. The gentle slopes of ‘pure’? manzanita had the deepest and richest soils, the intermediate slopes of manzanita-chamise had shallow soils often lacking
1965 | WILSON & VOGL: CHAPARRAL 5)
@ OAK- MANZANITA
O MANZANITA—CHAMISE @ "PURE" MANZANITA
« MANZANITA— COULTER
180°
Fic. 1. Slopes and exposures of Arctostaphylos association stands. Each circle represents 10 degrees of slope with the smallest circle representing 0-10 degree slopes. Compass directions indicate exposure.
in organic matter, and the steep slopes of manzanita-pine had a general absence of soils.
A gradient of vegetation often follows an elevational gradient, but in the Santa Ana Range, this gradient is highly modified by exposure. Con- sidering exposure preferences, “pure” manzanita was most often located on the north and northeasterly-facing slopes. Manzanita-pine and oak- manzanita were also found on north-facing slopes. Manzanita-chamise was scattered on both north and south-facing slopes, but occurred as the dominant cover on the southerly faces on the seaward side of the main divide. The west faces are warm-xeric with chamise going to the moun-
54 MADRONO [Vol. 18
tain crest. On the colder north exposures, Adenostoma does not succeed to the top but generally terminates in a chamise-manzanita association. The east or inland faces support Cercocarpus betuloides and Quercus species which form a solid cover mixing with Arctostaphylos only at the crest.
Stands situated on southwesterly slopes were generally on decomposed granite (quartzite and potash feldspar), while northerly and easterly slope stands were usually on a sedimentary complex of shale and fine clay.
In each plant association depths of soil horizons were measured. Lit- ter (A,) of oak-manzanita stands averaged 1 inch in depth which is less than in any other association. This may be due to a rapid rate of decom- position since the humus layer (A,) averages 6 inches in depth. Chamise- manzanita stands had an average A, of 1.5 inches. Slow decay resulted in a shallow average A, of 0.6 inches. These stands are on drier slopes with Adenostoma providing little cover or soil protection. Pine-manzani- ta stands had an average A, depth of 2.5 inches and an A, of 1.5 inches. “Pure” manzanita stands produced the deepest average Ay of 3.7 inches, with a rich average A, horizon of 5 inches.
These results suggest that “pure” manzanita produces and accumu- lates more litter and organic matter than when mixed with Adenostoma or Pinus coulteri, and produces excellent humus when associated with Quercus on moist sites. As a member of broadleaf chaparral Arctostaphy- los is a good soil producer, especially when compared to needleleaf Adeno- stoma.
Arctostaphylos glandulosa assumes different physiognomic character- istics growing under various conditions or associated with different spe- cles.
In manzanita-chamise stands Arctostaphylos glandulosa grows to a maximum height of about 6 ft. Branches are thin with numerous laterals and dead twigs remaining on the main stems beneath the surface canopy.
In oak-manzanita chaparral the Quercus species and Arctostaphylos do not mix but grow as separate aggregations (fig. 3). In other respects the growth character of A. glandulosa is much like that in manzanita- chamise stands. Small open areas within Quercus clumps are often filled by Arctostaphylos suggesting that it can grow in oak litter, but perhaps not in oak shade or deep oak litter.
Beneath Pinus coulteri, Arctostaphylos glandulosa stems grow pros- trate and parallel to the ground. At the edge of the pine canopy, the stems respond phototropically and become vertically orientated. The tallest A. glandulosa individuals (14 ft) were measured in this associa- tion.
Arctostaphylos in pure stands forms a dense mass of branch mesh- work (fig. 4). This type of growth produces a uniform canopy. Individu- als grow from 10 to 12 ft in height, with stout, twisted stems lacking twigs beneath the canopy.
1965] WILSON & VOGL: CHAPARRAL 55
Fic. 2. A manzanita-Coulter pine association on a northeast slope. Arctostaphylos glandulosa occupies the gentler slopes and the Pinus coulteri occupies the steeper slopes of the drainage pattern.
In all associations, Arctostaphylos stems were observed to grow at right angles to the slope or to exhibit a down-hill tendency.
56 MADRONO [Vol. 18
Heteromeles arbutifolia M. Roem., like Quercus, grows adjacent to Arctostaphylos with few individuals intermixing. Most H. arbutifolia plants grow on steeper sites in isolated clusters away from Arctostaphy- los. The Ceanothus species appear to be rapidly declining. Most of the Ceanothus plants encountered were either dead or failing. Surviving plants were usually taller than the surrounding Arctostaphylos indicating that the shading competition of Arctostaphylos may hasten the death of the relative short-lived Ceanothus. Garrya observed in association with Arctostaphylos exhibited growth habits similar to those of Arctostaphy- los.
Herbaceous plants were generally absent in the understory of Arcto- staphylos glandulosa chaparral. Annuals, perennials, and grasses were found only on disturbed sites, along road cuts, or on fire breaks in this chaparral.
DISCUSSION
For ease of comparison, the community approach categorizes the na- tural vegetation in the mind of the investigator. In reality, the vegeta- tion is not always as neatly segmented or defined by sharp borders and actually varies gradually and continuously, both in time and space. The vegetation in this study is summarized into four associations. These as- sociations become convenient tools in isolating segments and lumping floristic similarities for the purpose of summarizing trends.
The oak-manzanita association has the greatest variety of species, perhaps because it is found on steep and rocky terrain which provides a great variety of environmental conditions and thus a number of ecologic- al niches.
In comparing the role of competition between Pinus coulteri and Arctostaphylos glandulosa it appears as though P. coultert cannot suc- cessfully compete with the fire-resprouting A. glandulosa in the face of repeated burning. After fire P. coulteri establishes itself in rivulets, channels, and erosion patterns where the burls of A. glandulosa have been washed out or in areas too steep or unstable for A. glandulosa to persist. Manzanita-Coulter pine associations contained little Adenostoma and even less Ceanothus. The lack of living Ceanothus is attributed to the lack of disturbance in the area for at least the last 25 years (Hadley, 1961; Patric and Hanes, 1964). Ceanothus is known to be relatively short-lived and to require fire or other disturbances to reproduce (Quick and Quick, 1961). Accounts of competition between Pinus coulteri and Arctostaphylos species indicate that often with fire, the association may give way to a Ceanothus-Arctostaphylos association or “pure” man- zanita (Cooper, 1922; Sampson, 1944; Zobel, 1953).
The “‘pure” manzanita association occupies the most gentle slopes, has the highest number of individuals per acre, and provides the most com- plete cover (fig. 4). The cover produces a canopy equal to that of a forest (Cooper, 1922) and demonstrates the greatest control of the ground surface.
1965 | WILSON & VOGL: CHAPARRAL oe
*
iO Bae 3
Fic. 3. A gentle north slope supporting “pure’’ manzanita. The steeper sur- rounding slopes were covered mainly with Quercus dumosa and Q. wislizenii.
The dense, tight growth and closed canopy of Arctostaphylos glandu- losa have several marked affects on the microenvironment of the stands. Direct soil insolation is prevented and subsequent drying is retarded. Evaporation from the soil surface is reduced producing semi-mesic con- ditions. The closed canopy tends to eliminate competition of other species as evidenced by the lack of seedlings or herbaceous plants in the understory. Strips of A. glandulosa, in which canopy cover was mechanically removed by cutting, contained many shade intolerant
58 MADRONO [Vol. 18
shrub seedlings and herbaceous plants. The umbrella-like canopy affords protection against raindrop compaction and erosion. However, erosion by rain is not as important in the Santa Ana Arctostaphylos belt as it may be elsewhere. The major forms of precipitation in this “cold” chaparral are fog-drip, freezing moisture (hoar frost), and snow.
The high density of stems retards slope erosion, aids water infiltra- tion, and adds large quantities of litter in the form of twigs and leaves, thus influencing soil genesis. This litter is not readily washed, wind- blown, or lost by desiccation as it is in Adenostoma chaparral. More organic matter is accumulated and retained in Arctostaphylos chaparral than in any of the other Santa Ana chaparral types. The mesic nature of the soil aids in rapid decay and incorporation of organic material with mineral soil (by chaparral comparisons), producing a pronounced A, soil horizon (Wilson, 1963). Arctostaphylos glandulosa, then, is a good soil builder and watershed cover as compared to other chaparral types.
Cooper (1922) noted that Arctostaphylos species showed a preference for degree of slope and became established as a dominant on the most gentle slopes available in a favorable site. He also noted that Arcto- staphylos gave way to oak and pine on steeper slopes. Similarly, A. glandulosa in the Santa Ana Mountains formed solid stands on the gentlest north and northeast slopes above 3200 ft elevation in the presence of marine air and grew with Pinus coulteri and Quercus species on the steeper slopes (fig. 1). Of the steepest slopes (40°-100°), the xeric, unstable types with running tallus were occupied by Quercus while the more mesic, stable precipices were covered by Pseudotsuga macro- carpa. Occasional Arctostaphylos plants grew in rocky crags or on steep cliffs but did not form solid stands. The germination, seedling, and root- ing requirements of Pinus coultert, Pseudotsuga macrocarpa, and Quer- cus species may enable them to dominate pioneer and unstable sites of steeper slopes, rocky outcrops, or slides. Pinus coulteri is also found on gentle slopes but is limited, possibly because it cannot successfully com- pete with the burl-sprouting Arctostaphylos glandulosa. In many areas, A. glandulosa with the aid of fire, appears to be a potential successor of pine. Barring disturbance, established P. coulteri might gradually replace the apparently long-lived Arctostaphylos stage of fire succes- sion. However, succession is complicated by periodic fires and subse- quent erosion, as well as by fungus and insect damage or kill to the P. coulteri (Pequegnat, 1951). Further research is necessary to deter- mine precisely the relative successional positions of A. glandulosa and P. coultert.
The exposure of slope is another important factor which affects the establishment of Arctostaphylos. The most dense stands are on north- facing slopes, particularly those slopes reached by the late winter, spring, and early summer fogs. These fogs strike the upper ocean-facing slopes and crests. As they billow over the crest, through openings, and across lower ridges, they afford moisture to some upper and northern slopes not
1965] WILSON & VOGL: CHAPARRAL 59
Fic. 4. The interior of a “pure” manzanita stand with the typical high density of stems and heavy cover of solid Arctostapylos glandulosa.
directly exposed to the sea. In addition, these northern slopes receive less solar insolation. Since the north faces are not directly exposed to the prevailing westerly winds, they are also subjected to less transpiration stress. Coulter pine-manzanita is correlated with the presence of fog belt areas on northern exposures.
East-facing slopes are in a fog and rain shadow and are subjected to descending winds with greater evaporation-transpiration stress for
60 MADRONO [Vol. 18
plants. These east slopes support Quercus-Cercocarpus chaparral with only scattered individuals of Arctostaphylos near the crest.
Amplitudes of tolerance and successional relationships are implied from the results of the comparisons of similarity and dissimilarity of stands outlined in the Methods. Each species associated with Arcto- staphylos has its own amplitude of tolerance or range of ecological re- quirements and no two amplitudes are the same for any of the species. Stands with a predominance of Quercus are the most dissimilar from those dominated by Arctostaphylos. Therefore, Quercus and Arcto- staphy are considered the most dissimilar in their ecological require- ments. Arctostaphylos and Pinus coulteri are the most similar. Within the Santa Ana Mountains, Quercus species find optimum growth condi- tions in hot, dry environments while Arctostaphylos glandulosa and Pinus coulteri prefer cooler and semi-mesic environments. Adenostoma differs from Arctostaphylos in that it reaches greatest importance on hot, xeric sites exposed to marine air. Ceanothus, although suffering from old age, demonstrated the widest amplitude of tolerance of any species in the Arctostaphylos chaparral associations, occurring with Adeno- stoma, Pinus, and Quercus.
All of the species vary gradually and continuously on an elevational gradient. That is, none of the species start or end abruptly at a certain elevation. This gradient is greatly modified by exposure, slope, soils, fire, and air currents. However, optimum conditions as reflected by the great- est numbers are found for Adenostoma at lower elevations, and for Arctostaphylos and Pinus at the highest elevations.
SUMMARY
Little quantitative or phytosociological study has been made of Arctostaphylos chaparral in the southern California Peninsular Ranges. In this study, it is proposed that Arctostaphylos glandulosa is a signifi- cant unit in the Santa Ana Mountain ‘‘cold” chaparral as; 1. a forest (Pinus coultert) competitor, 2. a potential factor in watershed manage- ment, 3. an important contributor in soil genesis, and 4. a possible agent of erosion control.
Importance values derived from the line-intercept and quadrat data are used to classify the 40 stands sampled into four associations; 1. oak- manzanita, 2. manzanita-chamise, 3. manzanita-Coulter pine, and 4. “pure” manzanita. These Arctostaphylos associations are designated by dominant and codominant species present and are all found above 3200 ft elevation in the Santa Ana Mountains. The results indicate that Arctostaphylos; 1. becomes a significant dominant when associated with Adenostoma fasciculatum at upper elevations on south- and west- facing slopes and at lower elevations on north- and northeast- facing slopes, 2. with fire is a potential successor of Pinus coulteri on the north and northeast slopes and ridges, 3. is associated with Quercus aggregations but demonstrates mutual exclusion, and 4. forms solid stands on gentle north and northeast slopes above 3200 ft in the presence of marine air.
1965] WILSON & VOGL: CHAPARRAL 61
Plant growth is dense in all four associations with less than 10% bare ground in any association.
A comparison of the species composition of the 40 stands suggests that Quercus and Arctostaphylos are the most dissimilar in ecological tol- erance and that A. glandulosa and Pinus coulteri have similar require- ments. Quercus species attain their greatest growth and density in hot, dry environments and Arctostaphylos and Pinus coulteri find optimum conditions in cooler and semi-mesic environments.
Factors of altitude, exposure, slope, fire, and marine air determine the extent to which Arctostaphylos glandulosa successfully competes with associated species to become the dominant plant in the “cold” chaparral of the upper elevations in the Santa Ana Mountains.
Botany Department, California State College at Los Angeles
LITERATURE CITED
ABRAMS, L. 1951. Illustrated flora of the Pacific States. Vol. 3. Stanford Univ. Press.
BavEr, H. L. 1936. Moisture relations in the chaparral of the Santa Monica Moun- tains, California. Ecol. Monogr. 6:409-454.
Bray, J. R. and J. T. Curtis. 1957. An ordination of the upland forest communities of southern Wisconsin. Ecol. Monogr. 27:325—349.
Cooper, W. S. 1922. The broad-sclerophyll vegetation of California. Carnegie Inst. Publ. 319.
DeTLinc, L. E. 1961. The chaparral formation of southeastern Oregon, with consid- eration of its postglacial history. Ecology 42:348-357.
Haptey, E. B. 1961. Influence of temperature and other factors on Ceanothus megacarpus seed germination, Madrofo 16:132-137.
Hinps, N. E. 1952. Evolution of the California landscape. Calif. Dept. Nat. Res. Div. Mines Bull. 158.
Jauns, R. H. 1954. Geology of the Peninsular Range Province, southern California and Baja California. In R. H. Jahns, Geology of southern California. Calif. Dept. Nat. Res. Bull. 170.
Jepson, W. L. 1916. Regeneration in manzanita. Madrono 1:3-11.
McMiny, H. E. 1939. An illustrated manual of California shrubs. Univ. Calif. Press, Berkeley.
Mitter, E. H. 1944. The effect of altitude and slope exposure on environmental conditions and the growth of chaparral species in the San Gabriel Mountains of southern California. Ph.D. thesis (unpublished). Univ. of Southern Calif., Los Angeles.
. 1947. Growth and environmental conditions in southern California chap- arral. Am. Midl. Nat. 37:379-420.
Mownz, P. A. 1959. A California flora. Univ. Calif Press, Berkeley.
Patric, J. H. and T. L. Hanes. 1964. Chaparral succession in a San Gabriel Moun- tain area of California. Ecology 45:353-361.
PEQuEGNAT, W. E. 1951. The biota of the Santa Ana Mountains. Jour. Entomol. and Zool. 42:1-84.
Quick, C. R. and A. S. Quick. 1961. Germination of Ceanothus seeds. Madrono 16:23-30.
Sampson, A. W. 1944. Plant succession on burned chaparral lands in northern Cali- fornia. Univ. Calif. Bull. 685. Berkeley.
WELILs, P. V. 1962. Vegetation in relation to geological substratum and fire in the San Luis Obispo Quadrangle, California. Ecol. Monogr. 32:79-103.
62 MADRONO [Vol. 18
WIESLANDER, A. C. and B. O. SCHREIBER. 1939. Notes on the genus Arctostapyhlos. Madrono. 5:38-47.
Witson, R C. 1963. Phytosociology of manzanita (Arctostapyhlos) chaparral in a southern California Peninsular Range. Master’s degree thesis (unpublished). Calif. State College at Los Angeles.
ZOBEL, B. 1953. Geographic range and intraspecific variation of coulter pine. Madrono 12:1-7.
REVIEWS
Flora Europaea. Edited by T. G. Tutin, V. H. HeEywoop, N. A. Burces, D. H. VALENTINE, S. M. WALTERS, AND D. A. WEBB. Vol. 1. Lycopodiaceae to Platanaceae, xxxil + 464 pp., 5 maps. Cambridge Univ. Press. 1964. $16.00.
From the beginning in 1956, the Flora Europaea project has been broadly co- operative, highly organized, and based on a premise that much value lies in having a flora completed in the shortest possible time. Volume I, contributed by 51 authors representing 14 nationalities, provides in part the first synthesis of the European flora on a continental scale and, much to the benefit of most American botanists, in a single language. The remaining three volumes are planned for publication within the next eight years. The area covered, shown by maps, extends from Spitzbergen to the Azores and eastward through the Mediterranean (including Sicily, Crete, Euro- pean Turkey, Crimea, and northern shore of Caspian Sea) to the Ural Mountains.
Readers should be able to name to subspecies any fern (or fern ally), conifer, or flowering plant which grows wild in Europe and also those commonly cultivated. The descriptions are brief but adequate and often are followed by a cogent state- ment regarding variability and relationships to other taxa—a strong feature. Author or editor credit is provided for each generic treatment. Chromosome numbers sensi- bly are given only if determined from materials of known wild European origin. A check-list of references will be published separately which ought to explain some new reports such as 2n = 22 for Meconopsis cambrica. Sections, subgenera, and subfamilies appear where appropriate in the text. All families to be treated in the four volumes are keyed in Volume I. The sequence is Englerian except that mono- cots will come last. Filicopsida are divided into 21 families. Molluginaceae, Tetra- goniaceae, and Parnassiaceae will seem strange to Americans. Paeoniaceae follows Ranunculaceae, Fumariaceae is submerged. Mahonia is separated from Berberis. Platanus acerifolia is considered a synonym of P. hybrida Brot. Raphanus raphanis- trum is divided into five subspecies; Brassica rapa L. is favored over B. campestris, showing that California botanists have something still to learn. Of very minor mo- ment, Capparaceae is conserved over Capparidaceae and the perigynous condition of Eschscholzia was overlooked.
The format should satisfy nearly everyone and there are useful dividends such as the list of basic or standard floras of the region, the list of families in Volume I, and the lists of titles of books and periodicals cited in the text. An index to signs, abbreviations, and explanatory notes, as well as a glossary of technical terms and some English-Latin equivalents are printed on blue paper for quick finding. The only suggestion that I dare make is that type species or lectotype species for genera ought to have been indicated at least when a member of the flora.
Many years will pass before the full significance of the Flora Europaea project can be known. Undoubtedly many European botanists, besides the contributors, will be stimulated to look once again at the plants in their own backyards. The numerous statements of compromise in this text point out many interesting prob- lems which call for more detailed study. Instead of bringing floristic researches to a close, the Flora surely will be the starting point for an entirely new look at the plants of Europe. With this major effort recently completed, the floristic gaps in the northern hemisphere stand almost as a challenge. Here is a model, which ought
1965] NOTES AND NEWS 63
to be in every practicing taxonomist’s office, and perhaps it is time to think seri- ously about a Flora North America—WaALLAcE R. ERnsT, Smithsonian Institution, Washington, D.C.
NOTES AND NEWS
ARCEUTHOBIUM DOUGLASII IN NEVADA AND WYOMING.—The Douglas-fir dwarf- mistletoe, Arceuthobium douglasti Engelm., is one of the most widely distributed members of the genus in North America, ranging from southern British Columbia (J. Kuijt, Natl Mus. Canada Bull. 186:134-148. 1963) south to Durango and Nuevo Léon in Mexico (F. G. Hawksworth and D. Wiens, Brittonia, in press). Gill (Trans. Conn. Acad. 32:111-245.1935) recorded this species in all of the eleven Western States except Nevada and Wyoming. Recently A. douglasii was discovered on Pseudotsuga menziesii (Mirb.) Franco in these two states by R. S. Peterson of the U. S. Forest Service. Nevada. White Pine Co.: NE slope of Wheeler Peak, W of Baker, Peterson 63-337. Wyoming. Lincoln Co.: Wolf Creek Campground, Snake River, Targhee National Forest, Peterson 62-64 (RM); Teton Co.: 4 miles E of Idaho border on Teton Pass Road, Peterson 62-12 (RM). Specimens of the above are in the herbarium of the Rocky Mountain Forest and Range Experiment Station. Arceuthobium douglasii does not occur in many areas where Pseudotsuga menziesii, its principal host, is found. It has not been observed in northern Colorado, central Wyoming, central Montana, Washington or Oregon west of the Cascades except in southern Oregon, or in California south of Siskiyou and Shasta counties (J. Kuijt, Madrofio 15:129-139. 1960).—FRaAnK G. HAwxksworTH, U. S. Forest Service, Rocky Mountain Forest and Range Experiment Station, Colorado State University, Fort Collins.
WASHOE PINE ON THE BALD MountTAIn RANGE, CALIFORNIA.—Until recently Washoe pine, Pinus washoensis Mason & Stock., was known only from its type local- ity, a small area on the east slope of Mount Rose, southwest of Reno, Nevada (Ma- drono 8:61-63. 1965). In 1961 Haller (Madrono 16:126-132) reported an exten- sive uncut stand in the southern Warner Mountains, in the northeastern corner of California. Both stands have certain features in common: they occur at higher eleva- tions (7000-8000 ft) than the closely related P. ponderosa Laws., and they front on the sagebrush plains of the Great Basin at the eastern edge of the Sierra Nevada mixed-conifer forest.
During the summer of 1964 we encoutnered Washoe pine in the Bald Mountain Range, about 20 miles northwest of the Mount Rose stand and 100 miles south of the Warner Mountains stand (UC, and conifer herbarium of the Institute of Forest Genetics, Placerville.) We looked for it there because this range is the most extensive area of high-elevation forest fronting the sagebrush plains between Mount Rose and the Warner Mountains. The 5-mile-long crest of the Bald Mountain Range is from 8000 to 8760 ft in elevation, is oriented in a NNW-SSE direction, and forms part of the easternmost ridge of the Sierra Nevada. Its high point, Babbitt Peak, is at 39° 36.1’ N. latitude and 120° 06.2’ W. longitude.
We examined only the northern part of the range, from Babbitt Peak to a point about 3 miles northwest of the peak where the ridge drops off sharply to the north. Washoe pine is mostly confined to the top of the ridge, but is occasional in the Cercocar pus ledifolius Nutt. scrub on the west slope and in the P. monticola Dougl. forest on the east slope. The other common trees on the ridge are Abies concolor (Gord. & Glend.) Lindl., Juniperus occidentalis Hook., and Populus tremuloides Michx. Pinus yeffreyi Grev. & Balf. and A. magnifica A. Murr. are present but rare.
This Washoe pine stand consists of only a few hundred trees. The pines on the ridge-top are mostly wind-deformed, but occasional trees on the east slope range up to about 3 ft in diameter at breast height and 90 ft in height.
Flowering had just begun when we visited the area on July 4. The species can
64 MADRONO [Vol. 18
be easily distinguished from P. ponderosa by the color of its pollen cones and devel- oping second-season seed cones. In Washoe pine both kinds of cones are reddish- purple to purplish-black, in contrast to the red pollen cones and green seed cones that characterize P. ponderosa throughout most of the central Sierra Nevada region.
This stand occurs at higher elevations than the other known stands of Washoe pine. All trees that we encountered were at elevations of 8300 to 8600 ft, and it is doubtful if either the Mount Rose or the Warner Mountains stand extends much above 8000 ft. Unlike the other two stands, the Bald Mountain stand appears to be remote from P. ponderosa, and may represent P. washoensis in a relatively un- diluted form.—W. B. CRITCHFIELD, Pacific Southwest Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture, Berkeley, and G. L. ALLEN- BAUGH, Berkeley.
The following publications are of interest:
Secondary Phloem in the Pinaceae. By Lait M. Srivastava. University of Cali- fornia Publications in Botany 36:1—142. University of California Press, Berkeley and Los Angeles. 1963. $3.00.
Plan's of Arches National Monument. By BERTRAND F. HarRISsON, STANLEY L. WELSH, AND GLEN Moore. Brigham Young University Science Bulletin, Biological Series 5:1—23. Provo, Utah. 1964.
An Annotated Checklist of Cultivated Palms. By HaArotp E. Moore, Jr. Prin- cipes 7(4):119-182. Available from the Palm Society, 7229 S.W. 54th Ave., Miami, Florida 33143. 1963. $2.00.
100 Desert Wildflowers in Natural Color. By Natt N. Donce. 64 pp., 100 colored photographs. Southwest Monuments Association, Globe, Arizona, 1963.
Common Edible and Useful Plants of the West. By MuRIEL SWEET. 66 pp., 116 line drawings. Naturegraph Co., Healdsburg, California. 1962. $1.00.
Flowers and Ferns of Muir Woods. By GLApys L. SMITH AND DRAWINGS BY JEANNE R. JANISH. 32 pp., 1 map, 1 black and white photograph, 12 colored photographs, 60 line drawings. Muir Woods Natural History Association, Muir Woods National Mon- ument, Marin Co., California. 1963.
The Conifer Walk. By P. H. Brypon. 32 pp., illustrated. Strybing Arboretum Society, Golden Gate Park, San Francisco. 1964. This is a guide to the conifers in the Strybing Arboretum.
Flora of West Virginia. Introductory Section and Part IV. By P. D. StrausBAUGH AND Earu L. Core. West Virginia University Bulletin 65(3-1) :xxxi; 65(3-2):vi + 861-1075. Morgantown, West Virginia. 1964. These two parts complete the Flora of West Virginia, the first part of which was published in 1952.
An Annotated Catalogue of Plants from Window Rock, Arizona. By V. L. BOHRER AND M. BercseEnc. Navajoland Publications, Navajo Tribal Museum, Window Rock, Arizona. li + 29 pp. 1963. $0.25.
ADRONO
VOLUME 18, NUMBER 3 JULY, 1965
Contents
PAGE
CLINAL PATTERNS IN FROND ANATOMY OF POLYPODIUM, Robert M. Lloyd 65
THE CALIFORNIA SPECIES OF GUTIERREZIA (COMPOSI- TAE-ASTEREAE), Otto T. Solbrig 75
TAXONOMY OF POLYGONUM, SECTION POLYGONUM (AVICULARIA) IN NortH America, Thomas R. Mer- tens and Peter H. Raven 85
A REEVALUATION OF BUDDLEIA CORRUGATA, Eliane M. Norman 92
A WEST AMERICAN JOURNAL OF BOTANY
‘UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
Entered as second-class matter at the post office at Berkeley, California, January 29,
1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price
$6.00 per year. Published quarterly and issued from the office of Madrofio, Herbarium, Life Sciences Building, University of California, Berkeley, California 94720.
BOARD OF EDITORS
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LymANn Benson, Pomona College, Claremont, California HERBERT F. COPELAND, Sacramento College, Sacramento, California JouN F. Davipson, University of Nebraska, Lincoln WALLACE R. Ernst, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California Mitprep E. Marurtas, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley Marion OwnBeEY, Washington State University, Pullman REED C. Rotiins, Gray Harbarium, Harvard University Ira L. Wiccins, Stanford University, Stanford, California
Editor—JoHN H. THomAs Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Annetta Carter, Department of Botany, University of California, Berke- ley. First Vice-President: Winslow R. Briggs, Department of Biological Sciences, Stanford University. Second Vice-President: Arthur R. Kruckeberg, Department of Botany, University of Washington. Recording Secretary: Mary L. Bowerman, De- partment of Botany, University of California, Berkeley. Corresponding Secretary, Margaret Bergseng, Department of Botany, University of California, Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State College.
1965] LLOYD: POLYPODIUM 65
CLINAL PATTERNS IN FROND ANATOMY OF POLYPODIUM
RoBERT M. LLoyp
Variability in frond anatomy of Polypodium californicum Kaulf. has been known as early as Eaton’s (1879) monograph delineating two vari- eties, var. Raulfussii D. C. Eaton with coriaceous fronds and var. inter- medium (H. & A.) D. C. Eaton, a more herbaceous inland form. He re- ported the former growing usually on rocks near the sea and stated: “The texture of fronds from inland localities is rather thinner than in P. vulgare, and the veins are more easily seen; but plants from the sea- coast .. . have a firmer frond, and less conspicuous veins,”’ the latter, var. kaulfussu, “runs by gradation into the inland form.” Kendall (1923) presented evidence for retention of the varieties but stated that the two forms vary within the same morphological limits except for the extremely leathery texture of var. kaulfussii. Lloyd (1962) found frond texture to change from herbaceous to coriaceous as a function of distance from the ocean.
Although the phenomenon of anatomical plasticity in relation to the environment has been frequently dealt with, few studies have shown it to occur in a clinal pattern and even fewer studies have offered quantita- tive anatomical data to demonstrate the actual nature of this change. Shields (1950) summarizes much of the work which has been done re- lating specific influences of the habitat to morphology. This paper pre- sents an analysis of five populations of P. californicum growing at Pt. Reyes, California.
The author extends his grateful thanks to Sherwin Carlquist, H. G. Baker, R. W. Cruden, F. Rickson, and D. M. Hutt for their critical appraisal of the manuscript and to Rancho Santa Ana Botanic Garden and Duke University for the use of their facilities.
METHODS AND MATERIALS. Fronds were collected in the field at five locations (A-E. Lloyd 567, 568, 569, 570, 571, RSA) during January 1961, and the longest pair of pinnae were killed and fixed in formalin- aceto-alcohol (Johansen, 1940). Material was infiltrated, embedded, and sectioned at 12—15 uw. according to the usual paraffin techniques. Sec- tions were stained with Northen’s modification of Foster’s method (Johansen, 1940) and mounted in Canada Balsam. Surface characters were measured on preparations made by infiltrating and mounting 10 mm squares of dried material in water. Observations were made from transverse sections of pinnae on both cells and tissues to determine variation in each and provide evidence for the nature of anatomical dis- tinctions.
Maprono, Vol. 18, No. 3, pp. 65-96. August 25, 1965.
66 MADRONO [Vol. 18
PACIFIC OCEAN
7 Noon ™ « Bd :
ORAKES BAY A.
PT. REYES — _~ ! MILES
Fic. 1. Map of the Pt. Reyes region showing locations of populations A to E.
Ecotocy. Pt. Reyes Peninsula, extending 12 miles from the mainland of Marin Co., California, is directly exposed to ocean influences on two sides (fig. 1). Forested areas in more protected regions are gradually replaced by coastal scrub and grassland communities closer to the shore. This change in vegetation is accompanied by higher wind velocity and relative humidity. The topographic relief in relation to the prevailing wind thus defines a mosaic of micro-habitats, but one that changes to greater exposure as the wind assumes dominance over the topography near the coast.
Fronds of P. californicum are drought-deciduous unless ground water is available. The summer period, therefore, during which the wind velocity and relative humidity are at their highest on the coast, will have little or no affect on frond characteristics.
The five populations dealt with in this paper are summarized below:
Population A. Fronds of P. californicum were collected 2.2 miles east of the entrance to Samuel P. Taylor State Park on a protected road- bank under deep shade of Sequoia sempervirens. This location is char- acteristic of many forested areas of the region in which large herbaceous fronds develop.
Population B. Plants from this location, 1.4 miles west of Pt. Reyes Station on California highway 1, were restricted mainly to moss-covered tree trunks in the moderate shade of Umbellularia californica and Pseudotsuga menziesii and were in close proximity to Tomales Bay.
1965 | LLOYD: POLYPODIUM 67
Population C. This population, 0.9 miles west of Inverness, is from a habitat similar to but more open than that of population B. The same dominant trees are present. Polypodium californicum was found growing both epiphytically and in soil in a predominantly fern understory in moderate shade.
Population D. Plants were collected 5.3 miles west of Inverness by the side of the road to the Pt. Reyes lighthouse. In this area there are no trees and the ferns were found on moss-covered rocks of a north- facing road bank, shaded only part of the day. Associated species were Mimulus guttatus, Scrophularia californica, Polystichum munitum, Stachys, and Rubus.
Population E. Just north of the lighthouse gate at the southern tip of Pt. Reyes Peninsula, P. californicum is found on top of the bluffs above the ocean growing in full sun, only around and under small shrubs of Vaccinium ovatum and Gaultheria shallon. These species, along with Lupinus arboreus, are the dominant shrubs in the coastal grassland. Ferns were usually found on the leeward side of the shrubs where they were partially protected from ocean winds.
A related species, P. scoulert Hook. & Grev., is very rare at this local- ity but is found in abundance at lower elevations on the bluffs. There is some evidence of hybridization between this species and P. californt- cum in these latter localities (Lloyd, 1962; 1964).
Resutts. Data from this study are summarized in Tables 1 and 2.
Fronds of Polypodium are bifacial but there is no great difference in cell size and shape in the palisade and spongy tissues as is the case in many mesomorphic dicotyledonous plants. For this reason it is some- times difficult to demarcate tissues. In transverse section, the abaxial cells of the spongy layer are arranged with their long axes parallel with the frond surface. Many air spaces are present. In contrast, the adaxial cells or the palisade layer are very compactly arranged. In both layers there are fewer intercellular spaces near veins, producing a centric ar- rangement of cells in the chlorenchyma. Virtually all of the cells of the epidermis and mesophyll, except those of the vascular system, contain chlorophyll. Stomata are confined to the abaxial surface of the frond. Trichomes occur abundantly on the latter surface and are bicellular, un- branched, and glandular.
Investigation of fronds has demonstrated a gradient in thickness from a minimum of 128 » in population A to a maximum of 260 p» in popula- tion E. The change in proportion of the various tissues due to increased exposure follows a well established pattern reported in higher plants (Turner, 1923; Weaver and Clements, 1929).
Computation of relative proportions of the tissues of an entire frond shows an increase in the palisade layer as the coast is approached (table 1). There is a corresponding increase in the spongy layer until a maximum is reached at population C (fig. 2) although the mean size of the spongy tissue is larger than the palisade in all populations except
[Vol. 18
MADRONO
80
O 80 60
O Co)
68
1965] LLOYD: POLYPODIUM 69
E. Epidermal tissue show a range (A—E) from 20 to 13% on the adaxial cells and 14 to 10% on the abaxial side accompanying the grades of the mean thickness of the palisade. However, investigation of individual cell size of epidermal layers shows that both increase in depth and diameter in direct relation to coastal influences (table 2). This is most apparent on the adaxial side which is usually more directly exposed to sun and wind. In both cases the most striking increase is in the diameter of the cells.
Depth of individual cells in the palisade mesophyll gradually increases from a mean of 25 » in population A to 32 » in E. The diameters of these cells show a corresponding but greater increase from 31 » at A to 51 » at E. In both instances, however, the largest cells observed were in population D. Cell depth of the spongy mesophyll increases from a mean of 20 » in population A to 28 » in E. Diameters indicate a much more marked change with an average increase of 26 » from population A to E. In both layers, but especially in the spongy mesophyll, an obvi- ous decrease in the proportion of intercellular spaces characterize plants from near the coast. Related to this is the percentage of mesophyll cells contiguous with the epidermis. Palisade cells show an increase from 86% to 99.7% from population A to E whereas spongy mesophyll in- creases more markedly from 56% to 84%.
The number of tiers in the palisade tissue increases from two in the fronds of population A to four or more in population E, and the aver- age thickness of the tissue increases from 37 p» to 120 p (figs. 3-7). The spongy mesophyll increases in average thickness from 46 » in population A to 107 » at C. However, in contrast to palisade mesophyll the spongy tissue decreases in depth to 79 » in the most xeric population.
Stomatal size remains relatively constant in all populations. The small- est occur in population E. Number of stomata per mm’, however, varies considerably with a decrease from 33 to 16 in populations A to D and a sharp increase in E to 55. The number of trichomes per mm? shows a corresponding decrease in number to population D with a sharp increase at E.
DIscUSSION AND CoNcLuSIONS. The populations examined in this paper were chosen for their location in relation to the coast. Habitat characteristics of each relate to the topography of Pt. Reyes, which has a central spine of mountains sloping to low plains at the bluffs above the ocean. Anatomical data can be correlated with these habitats. The basic environmental factors influencing frond histology of the popula- tions studied are probably sunlight and wind exposure. The effect of salt-spray on population E is more pronounced than at the other stations and may help explain the anomalies of increased density of stomata
Fic. 2. Composition of the four tissues of the blade; clear = adaxial epidermis; stippled = palisade layer; solid = spongy layer; clear (lower) = abaxial epidermis; abscissa = population; ordinate = number of u in 20 » separations.
[Vol. 18
MADRONO
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LLOYD: POLYPODIUM
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72 MADRONO [Vol. 18
Fics. 3-7. Transverse section of frond pinnae from populations A to E at Pt. Reyes; 3. Pop. A; 4. Pop. B; 5. Pop. C; 6. Pop. D; 7. Pop. E. Figs. 3, 5-7 X 150; fig. 4 « 450.
1965] LLOYD: POLYPODIUM 73
and trichomes in that population. However, in the latter case there were more trichomes in population A than E.
In other parts of the range of P. californicum plants with herbaceous to semi-coriaceous fronds can be found growing in shaded forests directly above the ocean. Coriaceous fronds have also been found on plants in exposed situations completely removed from coastal influences. Fronds from these latter areas, however, do not reach the extremes in morphol- ogy demonstrated by Pt. Reyes plants.
Population A grows in the most mesic environment of all those studied here, a fact clearly reflected in the relatively thin fronds which have thick spongy mesophyll and numerous air spaces. Habitats of popula- tions B and C were open on the north and had less canopy cover. Al- though they appeared somewhat similar in exposure frond thickness of plants from population C differs from that of B almost to the same extent as B does from A. Contrastingly, differences between plants of C, D, and E are less extreme with a gradual increase in thickness toward the latter. Fronds of population E are more than twice as thick as those of A, a difference attributable to the palisade mesophyll.
Angiosperms may exhibit elongation of palisade cells (Clements, 1905) but this does not seem to be the case in Polypodium where the isodiametric cells only become larger. However, in both an increasing amount of palisade tissue forms at the expense of air space and spongy mesophyll. Although individual cells become larger in Polypodium with relation to more xeric habitats the increase of tissue thickness results from an increase in cell and tier number. This parallels the situation in Rumex acetosella (Transeau, 1904) which exhibits one cell layer of palisade mesophyll in mesic habitats and 2 to 3 layers in xeric environ- ments.
The increase in stomata number in angiosperms inhabiting xeric en- vironments has been noted by Salisbury (1927) and Philpott (1956). This increase in number is usually attributed to spacing and change of epidermal size. It has also been shown that numerous stomata usually have a higher transpiration rate. The sudden increase in stomata from population D to E indicates a drastic change in environment, perhaps regulated by the concentration of salt-spray in the atmosphere. Kunkel (1961) reported larger stomata in fronds of P. lastopus from shaded areas, smaller ones in plants of sunny areas. In addition, fronds more exposed to sun were much more fleshy and approached xerotypes in suc- culence. He stated that all possible transitional forms may be encoun- tered in individual habitats and feels that both of the forms (shaded and sunny) could be described as ecotypes. Transplant experiments with P. californicum have shown that in some instances the coriaceous char- acter of fronds is genetically controlled.
It is evident at Pt. Reyes that there is a clinal gradient in frond struc- ture correlated with increased exposure to sun and wind. Generally, the trends which were discovered may be summarized as follows:
74 MADRONO [Vol. 18
1. There is a direct relationship between frond thickness and degree of habitat modification from mesic to xeric environments.
2. This thickness is due mainly to a greater size of the palisade tissue although corresponding increases accompanied it in other tissues.
3. Individual cell sizes within each tissue was greatest in population D, somewhat smaller in E. The feature responsible for the greater thick-
ness of fronds in the latter was the additional tiers in the palisade meso- phyll.
4. Stoma number gradually decreases from mesic to more xeric habi- tats but sharply increases at the coast.
Department of Botany, University of California, Berkeley
LITERATURE CITED
CLEMENTS, E. S. 1905. The relation of leaf structure to physical factors. Trans. Am. Micr. Soc. 26:19-102.
Eaton, D. C. 1879. The ferns of North America. Vol. 1. S.E. Cassino Publ. Salem.
JoHANSEN, D. A. 1940. Plant microtechnique. McGraw-Hill, New York.
KENDALL, M. L. 1923. A study of variation in Polypodium californicum Kaulf. Am. Fern Jour. 13:75-87.
KUNKEL, G. 1961. Uber die morpho-dkologische Variabilitat von Polypodium lasi- opus Kze. Nova Hedwigia 3:375-381.
Lioyp, R. M. 1962. Studies in Californian species of Polypodium (Polypodiaceae). Master’s degree thesis (unpublished). Claremont Graduate School, Calif.
and F. A. Lane, 1964. The Polypodium vulgare complex in North Amer- ica. Brit. Fern Gaz. 9:168-177.
PHILPoTT, J. 1956. Blade tissue organization of foliage leaves of some Carolina shrub-bog species as compared with their Appalachian mountain affinities. Bot. Gaz. 118:88-105.
SALISBURY, E. J. 1927. On the causes and ecological significance of stomatal fre- quency, with special reference to the woodland flora. Phil. Trans. Roy. Soc. London B216:1-65.
Suretps, L. M. 1950. Leaf xeromorphy as related to physiological and structural influences. Bot. Rev. 16:399-447.
TranseAu, E. N. 1604. On the development of palisade tissue and resinous deposits in leaves. Science 19:866-867.
TuRNER, T. W. 1923. The ecology of Rhus toxicodendron.
Weaver, J. E. and F. E. Crements. 1938. Plant ecology, 2nd ed. McGraw-Hill, New York.
1965] SOLBRIG: CALIFORNIA GUTIERREZIA 75
THE CALIFORNIA SPECIES OF GUTIERREZIA (COMPOSITAE-ASTEREAE)
OtTTo T. SOLBRIG
The genus Gutierrezia (Compositae-Astereae) is represented in Cali- fornia by four species: G. californica Douglas, G. bracteata Abrams, G. sarothrae (Pursh) Britt. & Rusby, and G. microcephala (DC.) T. & G. (Solbrig, 1960). Of these four species, G. microcephala is fairly distinct and has been recognized in all the major floristic treatments. The other three species are closely related and their limits and identities have been considered differently in each of the major floristic treatments of Cali- fornia (Jepson, 1925; Abrams and Ferris, 1960; Munz, 1959). Since the characters distinguishing G. bracteata from G. sarothrae are not recog- nized easily in herbarium specimens and since there has been some doubt expressed in the past (Munz, 1935) as to the validity of these species, it was believed necessary to pursue the matter further by means of intensive field studies and population analyses. This report is an account of the results.
I am grateful to the Evolutionary Biology Program of Harvard Uni- versity for assistance in field studies and to the National Science Foun- dation for a research grant. I also wish to acknowledge Miss Karen Niecke for laboratory assistance and Peter H. Raven for seeds. Lorin I. Nevling, Jr. kindly has read the manuscript and made valuable editorial suggestions. I am also very grateful to G. L. Stebbins for a stimulating discussion on this subject.
MATERIALS AND METHODS
This study is based on statistical analyses of wild populations, chro- mosome counts, garden investigations, hybridization experiments and herbarium specimens. Up to nine characters (height of plant, length and width of the involucre, number of ligulate and tubular flowers, length and width of achenes, length of pappus and pollen diameter) were meas- ured in 50 plants chosen at random in each population studied. In this way, the value of different taxonomic characters could be evaluated in detail. Twenty populations were analyzed: two of G. microcephala, three each of G. sarothrae and G. californica and 12 of G. bracteata. Chromo- some numbers were determined in a total of 25 populations. Two hun- dred pollen grains were measured in 16 populations of known chromo- some number. Several collections were grown under uniform conditions at the Experimental Gardens of the University of California Botanical Gardens in Berkeley and later at the Harvard University Greenhouses in Cambridge, Massachusetts.
76 MADRONO [Vol. 18
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Fic. 1. Distribution of populations of Gutzerrezia in California. (Base map Copy- righted by Ginn & Co., used by permissicn.)
RESULTS
Distribution and ecological conditions. The most xerophytic of the California species of the genus is G. microcephala. It grows on the fringes of the Colorado and Mohave deserts on dry mountain slopes and in the valleys, but not in true desert areas (fig. 1). The more mesophytic G. bracteata and G. sarothrae have similar requirements. Both species are found in dry washes, rocky slopes and also occasionally in roadside : ditches and in fields. Gutierrezia bracteata grows from Yolo Co. south along the drier parts of the Coast Ranges to northern Baja California; G. sarothrae is found only south of the Tehachapi Mts. all the way to southern Baja California, Cedros I. and west into the Great Basin and northern Mexico. Gutierrezia californica on the other hand is a species with narrow ecological requirements. It is found only on three serpentine
1965 | SOLBRIG: CALIFORNIA GUTIERREZIA 77
outcrops in San Francisco Bay: one population on the Oakland Hills, one population on Angel Island, and one population on Point Bonita, Marin Co. Each of these three populations consists of less than 500 plants and all of them, but particularly the ones in Oakland and Marin Co., are threatened by encroaching civilization. Although G. californica is found in nature only on serpentine, it can be grown on non-serpentine soil without adverse effects. The same is true for progeny of the popula- tions of G. bracteata which grow on serpentine soil.
Mor phological analysis. a. Habit. The four species are similar in habit. All four are slightly woody subshrubs, semi- or completely globose, with small leaves and medium to small inflorescences borne at the end of the branches. Gutierrezia californica, which is significantly smaller than the rest, also conforms less to the habit description given, having a tendency to form only a few undivided and spreading branches which give rise to a rather open shrub. It also produces fewer inflorescences which tend to be borne solitary or on long stalks at the end of the branches. The other species form rather tight globose shrubs which are covered in the fall by small yellow heads which offer a rather showy display. Gutier- rezia bracteata is probably the largest species, but the differences are not significant (table 1). When grown in the experimental garden, all four species maintained their characteristics but increased in size.
b. Inflorescence. It already has been stated that the capitula of G. californica are borne solitary or in loose groupings at the end of the branches. Those of G. microcephala, on the contrary, are found in tight glomerules of 3 to 8 sessile inflorescences. The disposition of the heads of G. sarothrae is similar to that of G. microcephala but there is often a short pedicel to each inflorescence while in G. bracteata as a rule the heads are borne on pedicels several millimeters long. There is, neverthe- less, quite a bit of variability in this character.
There are differences in addition in the size of the capitula (table 1). The widest capitula are those of G. californica which vary from a popu- lation mean of 3.6 mm—4.20 mm; the narrowest are those of G. micro- cephala which average less than 1 mm wide. The capitula of G. sarothrae vary from an average value of 1.5 mm—2.5 mm per population while those of G. bracteata vary from 2.0 mm—3.4 mm. Although there is a slight overlap, it is clear that this last species tends to have wider heads. Gutierrezia californica also has the highest involucres (mean 5.70—7.50 mm average per population), but there is a sizable overlap with G. brac- teata (mean 5.1-6.4 mm). Gutierrezia sarothrae (mean 3.4—4.4 mm) and G. microcephala (mean less than 3 mm) have shorter heads.
Populations of these four species can be separated on the basis of mean population values of these two capitulum parameters (figs. 2, 3). Not so individual heads which can deviate up to two times from the mean population size. Nevertheless, if an effort is made to sample aver- age plants and to measure several heads, the capitulum furnishes one of the most valuable diagnostic characters.
78 MADRONO. [Vol. 18
TABLE 1. MEAN VALUES FOR PLANT HEIGHT, HEIGHT AND WIDTH OF INVOLUCRE, NUMBERS OF FLOWERS PER CAPITULUM, Pappus LENGTH AND POLLEN DIAMETER IN POPULATIONS OF CALIFORNIA GUTIERREZIA
All values in mm with the exception of plant height which is given in inches. Numbers indicate Solbrig vouchers at GH and UC except two G. californica samples which represent mean values taken from herbarium specimens.
Plant Involucre No.of Flowers Pappus Pollen Height i. W. Lig. Tub. Lig. Tub. Diam. G. californica.
Point Bonita, 3431 10.86 (ey) 3.63 15 12.54 0.97 1.74 18.57 Angel Island
herb.sample) —sC_i..... 5.70 4.20 1:3¢ 102: 332. 09022 Oakland (herb.sample) —__..... Sal ei wl “VISE Ry Ee
G. bracteata Cache Creek, 3432 23.44 5.58 FEA? 6.4 6.9 0.63 1.40 Arroyo del Puerto,
3433 17.31 6.13 2530 lie 8.6 0.56 140 14.23 Arroyo del Puerto,
2743 15.03 S12 3.39 7.6 9.0 0.91 1.80 13.83 New Idria Rd., 3434 17.85 6.19 2.19 5.9 6:5° 9° whe © 2a eee New Idria, 3435 16.48 5.86 DAS cee gto OO tes, ee Newldria 28302 26 © Soy. tee ES Bea” Pee 16.18 Los Gatos Canyon,
3436 22.58 Set Z0S:N 282° .° 2 ie 15.99 Kettleman Hills,
3437 24.34 6.34 2.15.) “at Qeees . 228. 6 =e Pond Ranch, 2748 23.7 5.16 3.30 6.5 8.9 1.24 202 “3 McKitrick, 3438 18.89 5.88 224% 2808 0 See? « (es 142 ee Temblor Range,
i payey 18.53 D257 2.61 4.9 4.3 0.98 eS Cuyamas Valley,
3439 19.94 6.38 ZO9w 6 Soon) ~Satteeceeee ee eee 16.51 Los Angeles, 3440 32.25 nines) 204s Sete Weeey dae eee 13.97 Tahquitz Canyon,
Bile, - —* ghee Agi LA Shae oe, eee 15.63 G. sarothrae Waviwild2773° te | Dee ee oe 13.48 Temecula, 2758 22.4 4.36 2.48 5:5 523 0.58 146 == AgWanea 2/60. °- ‘asan8 . S22. ‘feed oe). eee 13.90 Santa Vsabel 2/09 sccus dneee eee. atten, "2 ie a 13.32 Santa Vsabelecs09: = gic, ‘ee Bele: A 2 ee 13.98 Rancho Santa Fe,
2769 20.3 3.43 2.15 5.8 5.9 0.34 0.93 13.06 ChulavVistas2/68 > seca eee ee ee ee eee 13.84
Chula Vista, 2766 Ia Stel | 1.85 525 4.6 0.59 1.14 13.30 G. microcephala Hyde Park, Ariz.,
2504 1256 3.0 1.00 1.0 LO ootym oe White City, N.M., S22 10.4 220 0.80 0.9 PO: nisy §) 222 ee
St. David, Ariz., 2790 eS. bee fac “Aree! “ql * 5 2h cave eee) rn
1965] SOLBRIG: CALIFORNIA GUTIERREZIA 79
c. Flowers. Two floral characters are of taxonomic significance: num- ber of flowers per inflorescence and size of pappus, both for the ligulate and tubular flowers.
The largest number of flowers per capitulum is found in G. californica. The mean values per population are 7.2—7.5 ligulate flowers and 10.2— 12.5 tubular flowers per capitulum. Gutierrezia bracteata follows with 4.9-7.6 ligulate and 4.3—9.0 tubular flowers; G. sarothrae has 5.5—5.8 ligulate and 4.6—5.9 tubular ones; while G. microcephala has almost consistently one ligulate and one tubular flower per head. As with the characteristics of the capitulum, G. californica and G. microcephala are fairly distinct, while G. bracteata and G. sarothrae, although differing, have an area of overlap (fig. 2). Also it has to be remembered that these are mean values. Individual heads may have an excess or deficiency of the numbers given and at least 10 capitula should be analyzed in order to obtain a fairly reliable value.
The pappus length follows the same pattern as the number of flowers in a head: G. californica has the largest pappus, G. microcephala the smallest; G. bracteata and G. sarothrae have similar values, although the figures for the former are slightly larger. The pappus furnishes a valuable additional character.
d. Pollen size. The pollen diameter of populations of G. sarothrae, G. bracteata and G. californica of known chromosome number was meas- ured. The results are represented graphically in Fig. 2. Although there is a correlation between chromosome number and pollen diameter, the distribution is a continuous one. The variability exhibited by G. brac- teata in this character is worth noting.
Chromosome number. The California populations of Gutierrezia counted to date are listed in Table 2. All populations of G. sarothrae had n = 4, those of G. microcephala n = 8 and those of G. californica n = 12. So far only one number has been found in each population, but not more than five plants have been sampled per population. All the populations of G. bracteata with n = 12 are found on the south Coast Ranges while populations with n = 8 are found throughout the range of the species. No obvious morphological difference could be found in the chromosomes of the various species (Riidenberg and Solbrig, 1963).
Hybridization attempts. Plants of G. sarothrae, G. californica and G. bracteata (n = 8 and = 12) were intercrossed in Berkeley in 1957 and 1958 (Solbrig, 1960) and again in Cambridge in 1960 and 1961. All these attempts except one were unsuccessful. An interspecific hybird was obtained between G. californica and G. bracteata (n = 8), but was com- pletely sterile (loc. cit.). Although normally outbreeding, the plants of Gutierrezia are self-compatible; this hinders artificial hybridization at- tempts considerably. The results therefore are not as conclusive as could be wished, but apparently the species are fairly intersterile. In view of the differences in chromosome number this result is not surprising.
80 MADRONO [Vol. 18
CALIFORNICA
12 ———— Oo
BRACTEATA (mM)
=8 5
HEIGHT
SAROTHRAE
—_—__}+—_
INVOLUCRE
ne eee
10 15 20 25 1 2 3 4 POLLEN WIDTH INVOLUCRE WIDTH (mm)
—<—<—<——
e686 eo e @
an
a
FLOWERS O O
@ CALIFORNICA @ BRACTEATA @
SAROTHRAE QO) ad
MICROCEPHALA
OF RAY rs
a
NUMBER Nn
it} 12 13
4 5 6 7 8 9 10 NUMBER OF DISK FLOWERS
Fic. 2. Graphs showing correlation of pollen diameter (in micra) and chromo- some number (horizontal bar = range; vertical bar = mean); mean population width and height of the involucre (in mm) ; and mean number of disk and ligulate flowers in a head.
DISCUSSION
The taxonomic problem. Viewed from the vantage point of good knowledge of the species in the field and after carefully analyzing a number of breeding populations, the species appear fairly distinct. The same cannot be said, unfortunately, when the problem at hand is the
1965] SOLBRIG: CALIFORNIA GUTIERREZIA 81
TABLE 2. CHROMOSOME NUMBERS IN CALIFORNIA SPECIES OF GUTIERREZIA
G. californica. n = 12. Contra Costa Co., Oakland Hills, Solbrig 2154. Marin Co., Angel Island, Solbrig 2681; Point Bonita, Solbrig3431.
G. bracteata. n = 8. Yolo Co., Cache Creek, Rancho S. Ana Bot. Garden seed. Alameda Co., Corral Hollow, Solbrig 2159. Stanislaus Co., Patterson, Solbrig 2743, 3433. San Benito Co., Los Gatos Canyon, Solbrig 3436. San Luis Obispo Co., La Panza, Solbrig 2751. Los Angeles Co., Tujunga, Wash., Solbrig 3440. n = 12. San Luis Obispo Co., Temblor Range, Solbrig 2753. Santa Barbara Co., Padres Nat. Forest, Solbrig 2167; Cuyama Valley, Solbrig 2166, 3439. Riverside Co., Idylwild, Solbrig 2774. San Benito Co., New Idria, Solbrig 2830. San Bernardino Co., Tah- quitz Canyon, Solbrig 2775.
G. sarothrae. n = 4. Riverside Co., Idylwild, Solbrig 2773. San Diego Co., Agul anga, Solbrig 2760; Temecula, Solbrig 2758; Santa Ysabel, Solbrig 2763, 2765; Rancho Santa Fe, Solbrig 2769; Chula Vista, Solbrig 2766, 2768.
G. microcephala. n = 8. Inyo Co., White Mts., Rancho S. Ana Bot. Garden seed 9389, San Bernardino Co., Morongo, Rancho S. Ana Bot. Garden seed 8382.
identification of herbarium specimens, particularly incomplete or “scrappy” ones with poor or no label annotations. Gutierrezia californica and G. microcephala can be separated easily on the basis of involucral size and number of flowers per capitulum. On the other hand, large specimens of G. sarothrae and small ones of G. bracteata are likely to be confused. The characters used here offer the highest probability of correct identification, but the only absolute character is the chromo- some number which obviously cannot be obtained from a herbarium specimen. A key to identify the California species of Gutierrezia follows.
KEY TO CALIFORNIA SPECIES OF GUTIERREZIA
Involucre 3.5 mm wide or more; tubular flowers more than 10; ligulate flowers more
than 72 Plants of the San Francisco Bay area_...--.5...2.0.2 an. 1. G. californica Involucre less than 3.5 mm wide; tubular flowers less than 9; ligulate flowers less than 7.
Involucre 1.5 mm wide or less; 2.5 mm high or shorter; ligulate flower 1; tubular flower 1. Plants of the fringes of the Mohave and Colorado deserts and the IVAN SIVA Soe eee Serco ese eee cere eee ee eecseee es eee ee een Gr ICY OCE DUO
Involucre 1.5 mm wide or more; 3.0 mm high or more; ligulate flowers more than 4; tubular flowers 3 or more.
Involucre 2-3.5 wide; 5-7 mm high; ligulate flowers 5-7; tubular ones 4-9.
Plants of the Coast Ranges and southern California.................. 3. G. bracteata Involucre 1.5—2.5 mm wide; 3.5—4.5 mm high; ligulate flowers 4-6; tubular one 60, Plants of southern California 2.222 cee cies ee eee. 4. G. sarothrae
A second taxonomic question is if populations of G. bracteata with n = 8 andn = 12 merit recognition as distinct taxa. The morphological analysis does not show any appreciable difference but the crossing attempts seem to indicate that they are intersterile as expected. The situation of G. bracteata is not unique in the genus; polyploid popula- tions of G. sarothrae (Solbrig 1960; 1964) and of G. microcephala (Sol- brig, 1960) are also known. Even though there might be some merit in giving taxonomic recognition to these genetically isolated populations
82 MADRONO [Vol. 18
‘“ "4 7h ey ‘Wl Ne EN VAAN
pad
Mh KA Aa j
.
Fic. 3. Capitula of the four species of Gutierrezia: A, G. sarothrae; B, G. califor- nica; C, G. bracteata; and D, G. microcephala.
(in reality sibling species), I prefer not to. In the absence of any abso- lute morphological difference (no matter how small), it serves, in my opinion, no practical purpose.
Evolutionary history. Any statement about the evolutionary history of these species is highly speculative since there is no direct paleobotani- cal evidence. Nevertheless this much seems clear. The genus Gutierrezia
1965] SOLBRIG: CALIFORNIA GUTIERREZIA 83
represents a polyploid complex with a base number of x = 4 and specia- tion has taken place by increases in chromosome number together with ecological specialization. Of the living species, G. sarothrae is the most primitive one and is probably not much differffent from the ancestral stock (Solbrig, 1964, and in prep.), which was probably native to the lower central basin or to northern Mexico. It is clear from the morpho- logical analysis and geographic distribution that there were at least two invasions of Gutierrezia into California. One was by G. sarothrae and the other by G. microcephala. This latter species is probably of more recent arrival, both on account of its highly specialized type and also because it occupies a habitat which is relatively recent in California (Axelrod, 1950; 1956). The range of G. sarothrae might have once been continuous between the central basin and its present California distribu- tion, continuity which was interrupted by the formation of the Mohave and Colorado deserts in late Pliocene age. The presence of isolated popu- lations of G. sarothrae in the New York Mts. seems to indicate so. The species might have also extended once north along the coast ranges, although I have no indication of this. But it is tempting to speculate that this hypothetical continuous range was broken by the invasion of the sea into the Cuyama and Central Valley in the late Miocene and early Pliocene (Axelrod, 1956; James, 1963). The isolated populations to the north could have then evolved into n = 8 G. bracteata. Secondary contact after the retreat of the water in late Pliocene or Pleistocene might have produced n = 12 G. bracteata through amphiploidy. The tetraploid populations of G. bracteata might be the result of autopoly- ploidy followed by selection in the manner postulated by Darlington (1956), or they may be the result of amphiploidy with a now extinct species. This last pattern would be more in accordance with what is observed in other polyploid complexes (Stebbins 1950; and _ pers. comm.), nevertheless the close morphological similarity between G. sarothrae and G. bracteata does not rule out the first explanation.
The origin of G. californica is equally obscure, but it probably diverged from G. bracteata, by rapid evolution after moving into the slightlv more humid climate of the San Francisco Bay area. If the evolution of G. cali- fornica was the result of isolation or of hybridization of a tetraploid G. bracteata with an extinct diploid species cannot be determined. Nev- ertheless, the absence of known hexaploid G. bracteata north of San Benito Co., the morphological distinctness of G. californica and the pattern of behavior of other polyploid complexes appear to favor the hybridization hypotheses.
A point worth mentioning is the adaptation of populations of G. brac- teata and G. californica to serpentine soils. The first of these species is not quite a true serpentine species. In every instance, the populations are found on soil fairly mixed with materials derived from other types of
rocks. This can be observed best in the large serpentine outcrops in New Idria.
84 MADRONO [Vol. 18
On the other hand G. californica is a true serpentine endemic. Krucke- berg (1954) has discussed some genetic aspects of serpentine adaptation in particular reference to the genus Streptanthus. Gutierrezia californica can be grown on regular soils, but apparently cannot withstand com- petition in such soils in a natural situation. Plants of G. bracteata from non-serpentine localities can grow on serpentine, but they do better on other soils. They can, nevertheless, be considered preadapted to serpen- tine. In the San Francisco Bay area, the only suitable habitats available for G. bracteata plants were probably the serpentine outcrops. After that, isolation together with selection and genetic drift can be accounted to have brought forth a faster rate of evolution.
Gray Herbarium, Harvard University
LITERATURE CITED
ApRAMs, L. and R. S. Ferris. 1960. Illustrated flora of the Pacific States. Vol. 4. Stanford Univ. Press.
AXELROD, D. I. 1950. Evolution of desert vegetation in western North America. Car- negie Inst. Publ. 590.
———. 1956. Mio-Pliocene floras from west-central Nevada. Univ. Calif. Publ. Geol. Sci. 33:1-322.
DarincTon, C. D. 1956. Chromosome botany. Allen and Unwin, London.
James, G. T. 1963. Paleontology and nonmarine stratigraphy of the Cuyama Valley Badlands, California. Univ. Calif. Publ. Geol. Sci. 45:1-154.
Jepson, W. L. 1925. A manual of the flowering plants of California. Assoc. Students, Berkeley.
KRUCKEBERG, A. 1954. The ecology of serpentine soils: a symposium. III. Plant species in relation to serpentine soils. Ecology 35:267-274.
Muwnz, P. A. 1935. A manual of southern California botany. Claremont Colleges, Claremont.
——. 1959. A California Flora. Univ. Calif. Press, Berkeley.
RUDENBERG, L. and O. T. Sorpric, 1963. Chromosome number and morphology in the genus Gutierrezia (Compositae-Astereae). Phyton 20:199-204.
SOLBRIG, 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.
STEBBINS, G. L. 1950. Variation and evolution in plants. Columbia Univ. Press.
1965] MERTENS & RAVEN: POLYGONUM 85
TAXONOMY OF POLYGONUM, SECTION POLYGONUM (AVICULARIA) IN NORTH AMERICA
THOMAS R. MERTENS AND PETER H. RAVEN
The limits of Polygonum sect. Polygonum (Avicularia) have been de- fined by Hedberg (1946) in an illuminating study of pollen morphol- ogy in Polygonum and related genera. Excluded are most of the native species of North America, now regarded as belonging to Polygonum sect. Duravia. Within sect. Polygonum, however, there has been little agreement as to specific limits in North America (Fernald, 1950; Glea- son, 1952; Love and Love, 1956) or elsewhere.
Styles’ (1962) excellent biosystematic treatment of this group in the British Isles appeared to offer some orientation for the possible resolu- tion of its complexities in North America. We have addressed ourselves specifically to two problems: 1. The identity of certain populations found in Marin Co., California, and heretofore referred to P. fowleri Robins. (Howell, 1949); and 2. The resolution of the complex referred to P. aviculare L. in North America, especially in the western United States. The solution of these problems has demanded a consideration of all species of the group in North America, and certain critical observa- tions made during the course of this work will be reported here. Thus the present paper is a preliminary survey of the North American species’.
Four species of Polygonum sect. Polvgonum found in North America have inflorescences that appear terminal, the flowers being more or less clustered at the summits of the stems among very reduced leaves or bracts. Polygonum tenue Michx., morphologically similar to this group, has pollen of the Duravia-type, and is presumably referable to that sec- tion. The four species that do belong to sect. Polygonum, occur in North America, and appear to have terminal inflorescences are P. arenarium Waldst. & Kit., P. argyrocoleon Steud. ex Kunth, P. patulum Bieb. (P. bellardu auct., non All.) and P. vamosissimum Michx. The first three of these species are introduced from Eurasia, the last a native of North America. We concur with Gleason (1952) in considering F. exsertum Small (type, US!) a synonym of P. ramosissimum. In this species, as in some of those that will be discussed below, enlarged, oliva- ceous achenes are sometimes formed, particularly late in the season (Styles, 1962). Such fruits are usually flattened and wrinkled but per- fectly viable. They are very different in form from the normal fruits borne by the same plant. In this species, normal fruits are mostly 2.75- 3.25 mm long and 1.75—2 mm wide, whereas late-season fruits are
1 This study was carried out at Stanford University while the senior author held
National Science Foundation Faculty Fellowship 63115. The authors are most grate- ful to B. T. Styles and to J. T. Howell for their suggestions concerning this paper.
86 MADRONO [Vol. 18
3.75—5.4 mm long and 1.5—2.4 mm wide. Plants that have been referred to P. exsertum have a high proportion of distended fruits. We are un- able to account for the report by Love and Love (1956) of 2n — 20 in P. ramosissimum and 2n = 60 in P. exsertum and suggest that chro- mosome counts be made of a wide range of material before these num- bers be associated with morphologically defined entities. Some of the material named P. exsertum in herbaria is not P. ramosissimum but rather the analogous late-season form of P. prolificum (Small) Robins., a species discussed below.
The remaining North American species of sect. Polygonum, in which the flowers are scattered along the stem, may be separated by the fol- lowing key.
Achenes shining. Fresh foliage conspicuously glaucous; ocreae of lower nodes very conspicuous,
710mm long Mtlamtic COaSt- 2. oe ee 1. P. glaucum
Fresh foliage not conspicuously glaucous; ocreae less than 8 mm long, not conspicuous.
Fruits sublustrous; undistended fruits about 2 mm long................ 2. P. prolificum
Fruits glossy ; undistended fruits 2.8-4.7 mm long. Undistended fruits 4-4.7 mm long; distended fruits to 5.75 mm long. 3. P. oxyspermum Undistended fruits 2.8-3.4 mm long; distended fruits to 4.6 (-5) mm long. 4, P. marinense Achenes dull. Perianth bottle-shaped, constricted just below the apex; leaves blunt, oblong;
ACHENES AaDOWL SMT TOMS. st, cates See eee 5. P. erectum Perianth not constricted below the apex. Achenes abruptly beaked, granular.<.....22...222.:.42 3.2 6. P. fowleri
Achenes not abruptly beaked, not notably granular. Branch leaves much smaller than the stem leaves; fruiting perianth divided nearly to the base; fruit with 3 equal concave sides. Stem leaves subsessile or with petioles included in the ocreae; fruits 2.5—
5 ero 6) 02a ene enna re eran a Agi Nn tater Menem ore DO Saker 7. P. aviculare s. str. Stem leaves with petioles 4-8 mm long projecting from the ocreae; fruits Wt’ Scie 0010 0 gl (6) 0| een nen enn une Heme T RRR Savana aay: 9. P. boreale
Branch leaves and stem leaves subequal; fruiting perianth divided about half its length; fruit with 2 convex and one shorter concave side. 8. P. arenastrum
1. P. glaucum Nutt. A very distinctive species of the sands of the Atlantic Coast from Massachusetts to Georgia, with very large ocreae, P. glaucum has occasionally been recorded as the west Eurasian and North African P. maritimum L., which is a perennial and differs in sev- eral other characters. It has not been confused with other species in this treatment. Love and Love (1956) have reported 2n = 40 in P. glancum.
2. P. prolificum (Small) Robins. This species has sometimes been confused with P. ramosissimum, but can be distinguished by its short pedicels (included within, rather than exserted from, the ocreae) and less shiny fruits. Its flowers are not markedly clustered at the apex of
1965 | MERTENS & RAVEN: POLYGONUM 87
the stems. It is widespread across the United States and southern Can- ada, but commoner in the central and eastern portions of the continent. It can be reported, probably for the first time, from California, on the basis of the following collection: Cuttings Wharf on Napa River, Napa Co., 8 Oct. 1932, Howell 10800 (CAS). At this locality, and over much of the remainder of its range, it is doubtless introduced; but it is known only from North America and must therefore be native there.
3. P. oxyspermum Meyer & Bunge ex Ledeb. In North America this species occurs in a very small area in eastern Canada (Newfoundland, New Brunswick, and Nova Scotia). It is widely distributed on the coasts of northwestern Europe and may be introduced in North Amer- ica, as suggested by Hultén (1963) for a number of species with sim- ilar distribution patterns. Styles (1962) pointed out that the differ- ences between P. oxyvspermum and P. rai Babington were not constant, and reported a chromosome number of 2n — 40 for representatives of both entities from Europe. Later, Webb and Chater (1963; 1964) con- sidered P. rait a subspecies of P. oxyspermum, making the combination P. oxyspermum ssp. rait (Babington) Webb and Chater (1963). Styles described the fruits of P. rai as being 5-6 mm long and 3-3.5 mm wide, but his illustration (Styles, 1962, pl. 9) shows them as ranging from 4—5 mm in length and from 2.4—3.2 mm in width. On the other hand, his illustration of P. oxyspermum shows enlarged fruits that are apparently comparable to the late season fruits of P. ramosissimum dis- cussed above. These fruits are 5—6.1 mm long, and paler than the normal ones. Webb and Chater (1964) consider P. oxyspermum ssp. oxysper- mum to have fruits 5-6.5 mm long, and ssp. vam fruits 2.5-5.5 mm long. Thus the differences between the two subspecies may depend on the proportion with which they produce distended fruits late in the season (or earlier?). In the small amount of European material named P. raii that we have examined, the fruits are mostly 4.5—-5 mm long. North American specimens we examined had normal fruits 4—-4.7 mm long and 2.5-3.2 mm wide, and occasionally abnormal ones (as in the collection that is the type of P. acadiense Fernald) to 5.75 mm long. Thus they seem to fall in the same range of the European material. We have de- determined a very large series of this species (DAO, GH, NMC, US) as P. oxyspermum, since subspecific lines in the European material may or may not be applicable in North America. Love and Love’s counts of 2n = 80 (perhaps from a cell with doubled chromosome number?) in P. oxyspermum and 2n — 60 in P. raii, both from North American material, need to be confirmed.
4. P. marinense Mertens & Raven, sp. nov. A P. oxyspermo differt: achaeniis minoribus, 2.8-3.4 mm longis, 1.9-2.3 mm latis, interdum ad 4.6 (-5) mm longis, 2.5 mm latis, refertis, subplanis, subrugosis; sepalis ex comparatione angustiore hyalinomarginatis; pedicellis filiformis,
2—5.5 mm longis, floribus fructibusque saepe longe ab ocreis exsertis; chromosomata 2n = 60.
88 MADRONO [Vol. 18
Type: Marin Co., Calif., common in salt marshes at the head of Drake’s Estero, Point Reyes Peninsula, 22 Aug. 1961, Raven 16568 (DS). Growing up among mats of Salicornia virginica and Distichlis spicata.
Additional specimens examined: California. Marin Co.: salt marshes, Burdell, Howell 21252 (CAS, DS); Escalle salt marsh, Howell 19396 (CAS); salt marsh, Inverness, Howell, 1945 (CAS); saline marsh, north of Inverness, Nobs & Smith 912 (CAS, DS) ; salt marsh at head of Drake’s Estero, Howell 25326 (CAS).
This distinctive new species is entirely different from P. fowleri, with which it has been confused, but very similar to P. oxyspermum. It dif- fers from the latter in its smaller fruits, normally 2.8-3.4 mm long, and even when distended only up to 4.6 (—5) mm long; its longer, more slender pedicels in fruit; its more narrowly margined sepals; and its less conspicuous ocreae. In addition, its chromosome number is 2n = 60 (progeny of the type collection), not 2n = 40.
Polygonum roberti Lois. is a name that has been associated with a similar group of plants found on the shores of the western Mediter- ranean. These plants are said often to be perennial and to have shining achenes 2~3 mm long. Styles (1962) excluded them from his concept of P. rai, but Webb and Chater (1964) include them in their concept of P. oxyspermum ssp. rai. Consequently, Webb and Chater considered this taxon to have achenes 2.5—5.5 mm long. Whether these Mediter- ranean plants are in fact conspecific with P. oxyspermum or with P. marinense (geographically there is more in favor of the latter hy- pothesis than the former), there is considerable doubt about the correct application of the name P. roberti Lois. (Styles, 1962; Webb and Chater, 1964). Polygonum marinense is certainly not conspecific with P. oxyspermum, and even if it is found to include the plants of the western Mediterranean, there is apparently no earlier, valid, certain name for these. The native status of P. marinense in North America must for the present remain doubtful.
5. P. erectum L. This very distinctive species, which occurs from the region of the Rocky Mountains east to the Atlantic seaboard, should not be confused with the taxon called P. aviculare L. var. erectum (Roth) Koch, which is generally considered a synonym of P. aviculare s. str. Love and Love (1956) reported a somatic chromosome number of 2n = 40 for this species and 2n = 20 for P. achoreum Blake, but as we are unable to find any character to distinguish these two entities sat- isfactorily, we believe that it would be desirable to reinvestigate this complex cytologically.
6. P. fowleri Robins. This species ranges from Maine north to New- foundland, across the continent to Alaska, and south at least to the vicinity of Puget Sound, Washington. It is perhaps the most distinctive entity in the entire group. Its achenes are 2.75—3.5 mm long and 1.75- 2.25 mm wide. Enlarged late season achenes are relatively rare, and are up to 4.25 mm long and 2.25 mm wide. A high incidence of biconvex fruits is also characteristic. Plants from Marin Co., California, hitherto referred to this species (Howell, 1949; Munz, 1959) are totally different
1965] MERTENS & RAVEN: POLYGONUM 89
and have been described above as P. marinense. We have examined the type of P. fowleri (GH) and a large series of specimens from throughout its range. We have also examined the type of P. allocarpum Blake (US) and a series of specimens referred to that species. Like Gleason (1952) we are unable to distinguish this entity consistently from P. fowleri, and are thus unable to account for the somatic chromosome numbers 2n = 40 for P. fowlerti and 2n = 60 for P. allocarpum that have been reported by Love and Love (1956).
7. P. aviculare L. s. str. and 8. P. arenastrum Jord. ex Bor. With P. boreale (no. 9), these species constitute the North American mem- bers of the Polygonum aviculare complex. Thev are characterized by their dull, striate fruits and flowers in axillary fascicles. Styles (1962) divided this complex in Britain into 4 species: P. arenastrum Jord. ex Bor., P. aviculare L. s. str., P. boreale (Lange) Small, and P. rurivagum Jord. ex Bor. The last-mentioned entity is not known from North Amer- ica, the plants described by Love and Love (1956) as this species not agreeing with those treated under this name in Europe (Styles, 1962).
In Britain, the distinction between P. arenastrum and P. aviculare is rather sharp. Styles (1962) characterized P. arenastrum as having branch leaves about the same size as the stem leaves; persistent perianth divided about half its length; and fruits 1.5—2.78 mm long, with two convex and one narrowly concave side (rarely with two concave and one convex side). The chromsome number in this species was deter- mined as 2n = 40. In P. aviculare s. str. the branch leaves are smaller than those of the main stem (this is most evident in young plants); the persistent perianth is divided almost to the base; and the fruits are 2.5-3.5 mm long, with three more or less equal concave sides. The chromsome number is 2n = 60. It should be pointed out that Lindman (1912) in an early paper on this group called P. aviculare s. str. P. heterophyllum Lindm. and P. arenastrum P. aequale Lindm., and that Love and Love (1956) called P. aviculare s. str. P. heterophyllum and P. arenastrum P. aviculare.
It became evident early in the course of our study that most of the North American collections referred to “P. aviculare” were P. arenas- trum, and that plants referrable to this species occurred throughout the temperate United States and southern Canada, and were occasional farther north. We determined the chromosome number 2n = 40 in two collections typical of this species: one from the Stanford University campus, Santa Clara Co., Calif., Mertens 3 (DS), the second from Muncie, Indiana, Mertens 10 (DS). In addition, this chromosome num- ber was found in the root tips of germinating seeds from a collection from the serpentine area at Magalia, Butte Co., Calif., Howell 37564 (CAS). In this collection, the achenes were nearly triangular in tran- section and ranged from 2.5—2.75 mm long. The perianth likewise was divided about 24 of its length, and the plant appeared to be relatively heterophyllous. As Styles (1962) points out, however, accurate determi-
90 MADRONO [ Vol. 18
nations of heterophylly can be made only in young, well grown plants. We encountered numerous collections which we referred to P. arena- strum, largely on the basis of fruit size, which did not have the charac- teristic single narrow concave side of this species; but we have likewise seen similar collections from southern Europe.
We have determined the chromsome number characteristic of P. aviculare, 2n — 60, in the following two collections: about 5 miles north of Santa Cruz, Santa Cruz Co., Calif., along edge of an irrigated brus- sels sprout field, Mertens 5 (DS); State Highway 1 about 2 miles north of Santa Cruz Co. line, San Mateo Co., Calif., Mertens 6 (DS). Although the perianth in these collections was deeply divided and the fruits typical of P. aviculare in transection, the fruits, very small for this species, were 2.25—2.5 mm long. Moreover, the plants were not appreciably heterophyllous. Both the small fruit size and the lack of heterophylly of these collections may have been attributable to the fact that both were late season plants which, growing in irrigated ground, had been flowering for a very long time before they were collected in late September. Nevertheless, they would have been most difficult to determine as P. aviculare had their chromsome number not been deter- mined. Because we have seen relatively few collections from North America that were P. aviculare, it appears to be worthwhile to cite these. Doubtless it is much commoner than would appear here, as these weedy plants are poorly collected.
Alaska. Skagway, Eastwood 734 (CAS). Yukon Territory. Moosehide Mountain, Calder & Billard 4549 (CAS). British Columbia. Atlin, Setchell & Parks, 1930 (CAS). Alberta. Near Fort Saskatchewan, Turner 33 (CAS). Idaho. Kootenai Co.: valley of Lake Pend d’Oreille, Sandberg et al. 966 (CAS). Nez Perces Co.: near Forest, Brown 18 (DS). Nevada. Mineral Co.: Wassuk Range, Archer 7165 (DS). Washce Co.: Reno, Eastwood 14795 (CAS). California. Del Norte Co.: Crescent City, Van Deventer, 1934 (CAS); Kildale 1030 (DS). Humboldt Co.: near Cape Mendocino, Wolf 1208 (DS). Shasta Co.: Mt. Shasta, Cook 15403 (DS). Yolo Co.: Davis, Goodwin, 1932 (DS). Madera Co.: Red’s Meadows Pack Station, Raven 3666 (CAS) Mono Co.: Hilton Creek, Howell 27389 (CAS, immature). Sonoma Co.: Petaluma, Eastwood 10476 (CAS). Marin Co.: Almonte salt marsh, Howell 14814 (CAS); Greenbrae, Howell 19428 (CAS). San Francisco: Howell 32667 (CAS). San Mateo Co.: Redwood City, Thomas 8784A (DS; with P. arenastrum). Santa Clara Co.: near Mountain View, Thomas 7239 (DS). Santa Cruz Co.: Pajaro River, Thomas 3222 (DS). San Benito Co.: Hollister, Raven 2150 (CAS). Tulare Co.: Terra Bella, Howell & Barneby 29200 (CAS). Ventura Co.: Ojai Valley, Pettibone & Hubby, 1894 (CAS). San Bernardino/Riverside Co.: Jurupa Mountains west of Colton, Kamb 763A (CAS).
In concluding this section it can be stated that P. arenastrum is much commoner than P. aviculare in North America and that both are more variable than would be suspected from the descriptions of Styles (1962). In general certain identifications can be made of plants with the charac- teristic fruit shape of P. arenastrum or with fruits averaging shorter than 2.25 mm; or of plants with fruits longer than 2.75 mm. Well grown plants with fruits between 2.25 and 2.75 mm in length are likely to be
1965] MERTENS & RAVEN: POLYGONUM 91
P. arenastrum particularly if they are not heterophyllous and have the perianth divided about half of its length. The identification of late sea- son plants with fruits falling in this range is likely to be problematical in the absence of chromosome counts; on statistical grounds they are most likely to be P. arenastrum. The identity of the plants discussed by Love (1956) as P. buxiforme Small with a reported chromosome count of 2n = 20, must remain problematical for the time being. Their P. neg- lectum Bess. and P. aviculare L., both with 2n = 40, are probably P. arenastrum, and their P. keterophyllum Lindm. and P. littorale Link, 2n = 60, P. aviculare L. Both of these species are introduced in North America.
9. P. boreale (Lange) Small. We have not seen specimens of this spe- cies from North America, but it has been reported from Arctic regions in this continent and is probably present.
Doubtful and excluded species. On the basis of the material we have seen, we are unable at present to evaluate the status of P. autumnale Brenkle (said by Love and Love, 1956, to have 2n = 20), P. latum Small (said by Love and Love, 1956, to have 2n = 40), and P. mon- tereyense Brenkle (sect. Duravia?). A biosystematic study conducted along the lines of that carried out on British species by Styles (1962) would be necessary to evaluate the badly oversplit entities recognized in this group by some authors, notably Love and Love (1956), on the basis of very little evidence. Since these authors cite no specimens to document their chromosome number determinations, these reports must, in such a critical group, be disregarded.
One additional species of this group, P. polvcnemoides Jaub. & Spach, has been reported by Hitchcock (1964, p. 163) from Idaho and New York; it is said to have shining achenes and to be distinguished from P. aviculare (sens. lat.) by its nonstriate, scaberulous-papillate and strongly angled stems, larger, less deeply lacerate stipules, and papillate perianth. It is a native of southern Asia, and, as suggested by Hitchcock. may occur more widely in North America than reported.
Biology Department, Ball State University, Muncie, Indiana, and Division of Systematic Biology, Stanford University, Stanford, California
LITERATURE CITED
FERNALD, M. L. 1950. Gray’s Manual of Botany. Ed. 8. American Book Co., New York.
GLeason, H. A. 1952. Illustrated Flora of the Northeastern United States and adjacent Canada. Vol. 2. New York Botanical Garden.
HEDBERG, O. 1946. Pollen morphology in the genus Polygonum L. s. lat. and its taxonomical significance. Sv. Bot. Tidskr. 40:371-404.
Hircucock, C. L. 1964. Polygonum. Jn Vascular Plants of the Pacific Northwest 2:139-168.
Howe tt, J. T. 1949. Marin Flora. Univ. Calif. Press, Berkeley.
Hu ten, E. 1963. Phytogeographical connections of the North Atlantic. In Love, A., and D. Love (eds.), North Atlantic Biota and their History, pp. 45-72.
92 MADRONO [Vol. 18
LinpMaAn, C. A. M. 1912. Wie ist die Kollektivart Polygonum aviculare zu spalten? Sv. Bot. Tidskr. 6:673-696.
Love, A., and D. Love. 1956. Chromosomes and taxonomy of eastern North Ameri- can Polygonum. Canad. Jour. Bot. 34:501—-521.
Muwz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
STYLEs, B. T. 1962. The taxonomy of Polygonum aviculare and its allies in Britain. Watsonia 5:177-214.
Wess, D. A. and A. O. CuHaTER. 1963. Polygonaceae. Repert. Sp. Nov. 68:187-189.
—. 1964. Polygonum. Jn FI]. Europaea 1:76-80.
A REEVALUATION OF BUDDLEIA CORRUGATA ELIANE M. NoRMAN
As a thesis at the L. H. Bailey Hortorium, Cornell University, I pre- pared a revision of the North American species of Buddleia (Logania- ceae). This revision has not yet been published. However additional material from Baja California, Mexico, has recently become available which has made it necessary to alter the interpretation of two species treated in the thesis.
Buddleia corrugata was first collected by Marcus Jones at Arroyo Hondo (he spelled it Undo), Loreto, in October 1930. This locality lies at an altitude of approximately 2000 ft. on the north side of Cerro Gi- ganta. The type collection is quite depauperate. Now with additional collections from several areas of Baja California, the species can be much better defined and the variability better evaluated. Material from the Cape Region which had been interpreted in my thesis as a different, if closely allied species, should be regarded as a subspecies of corrugata. Two recent collections from the area of Volcan Las Tres Virgenes are very similar to the type in morphology and it seems best to treat them also as a subspecies.
The pattern of variation in B. corrugata may be a reflection of the geo- logic history of Baja California. During pre-Pleistocene time the Cape Region was cut off repeatedly from the northern part of the peninsula (Durham and Allison, 1960) thus isolating the two areas and allowing divergent evolution to take place. The volcanic peaks of Las Tres Vir- genes which arose in late Miocene (Savage, 1960) similarly provided an area where ecotypic variation could develop.
KEY TO THE SUBSPECIES
Leaves ovate, 0.7-3.0 cm broad. Lower surface of the leaves with erect pubescence, the stellate hairs candelabra; calyx 2.0-2.5 mm long; corolla 2.5-3.7 mm long; capsule 1.5-2.0 mm long B. corrugata ssp. corrugata Lower surface of the leaves with appressed pubescence, the stellate hairs mostly simple; calyx 3.0-4.0 mm long; corolla 4.0-5.5 mm long; capsule 2.5-3.0 mm ONG ott eae ti aa eaten tee Maret dae! B. corrugata ssp. gentryt Leaves linear, 0.2-0.3.mm: broad)... 4.0. ee B. corrugata ssp. moranii
1965] NORMAN: BUDDLEIA CORRUGATA 93
BUDDLEIA CORRUGATA Jones, Contr. West. Bot. 18:56.1933, ssp. cor- RUGATA. Shrub dioecious, 0.5-1.0 m high, with diffuse growth, the young twigs stellate-tomentose, the older branches greyish with rimose bark, the old naked branchlets persisting; leaves greyish-green with petiole 0.3-0.5 cm long, the stipular line inconspicuous, the blade ovate 1.0-3.0 cm long, 0.7—1.5 cm broad, thick erect stellate tomentum on both surfaces, underlain by glandular trichomes, the apex acute or ob- tuse, the base cuneate, somewhat decurrent, the margin strongly crenate or dentate; inflorescences sometimes branching at the base, with 3-15 pairs of sessile or short-pedunculate heads subtended by bracts, each head 0.5 cm in diameter usually with 5 flowers; calyx tubular, stellate outside, the tube 1.2—1.5 mm long, the lobes acute 0.5—1.0 mm long; corolla orange at maturity, campanulate, stellate-tomentose outside on upper half, the tube 1.7—2.5 mm long, with pitted hairs within, the lobes orbicular, spreading, 0.8-1.2 mm long; stamens subsessile, inserted on upper third of tube, the anthers 0.5 mm long, ovary oblong, 1.0—-1.5 mm long, the style 0.5—0.8 mm long, the stigma 0.4—0.6 mm long, shallowly 2-lipped; capsule ovoid, 1.5—2.0 mm long, puberulent, opening septi- cidally and loculicidally at the apex, the valves separating broadly; seeds ovoid, wingless, approximately 0.5 mm long.
Northern portion of Sierra Giganta, at 2000-3000 ft. and limestone mesa of Monserrate Island (fig. 1). Flowering in the spring.
Type: Arroyo Hondo (Undo) Ranch, Loreto, Sierra Giganta, Oct. 26, 1930, Jones 27361 (POM !, isotype at GH!).
Specimens examined. Rocky N facing slope, Valle de los Encinos, S side of Cerro Giganta, 750 m, Carter & Reese 4570 (UC); Scandent on nearly vertical N facing cliffs of peak S of Portezuela de Peloteado, S of La Victoria, 1050 m Carter &
Medellin-Lean 4675 (UC); Locally common at N edge of limestone mesa, Mon- serrate I., 200 m, Moran 9309 (SD).
BUDDLEIA CORRUGATA ssp. gentryi Norman, ssp. nov. Frutex 0.5—1.0 m altus; folia ovato-oblonga 2.0-6.0 cm longa, 1.0—-3.0 cm lata primum utrinque appressa stellato-tomentulosa et glandulosa deinde superne glabrata, marginis inequale sinuatis; inflorescentia aliquando basi ra- mosa, axe principe 4-10 juga cymarum capitarum sessilium breviterve pedunculatarum ferente, capite quoque 0.7-1.0 cm lato, 3—5 floro; calyx tubiformis 3.0-4.0 mm longus; corolla aurantiaca campanulata tubo 2.8-3.8 mm longo intus piloso, extus stellato-tomentello, lobis orbiculatis patentibus, 1.0-1.5 mm longis; capsula ovata 2.5-3.0 mm longa dehis- cens; semina ovata, non alata 0.5 mm longa.
Differing from ssp. corrugata by the larger leaves with appressed tomentum on the underside (fig. 2) and larger flowers and fruits.
Southernmost part of the Magdalena Plain (fig. 1). Flowering in the spring.
Type: Arroyo Seco about 25 mi W of La Paz, arroyo margin at base cf limy cliff, March 29, 1939, Gentry 4446 (UC!, isotypes at ARIZ!, DS!, GH!, K!, US!).
Specimen examined. Arroyo Conejo near the mouth, Moran 7455 (BH, CAS, DS, GH, RSA, SD, US, US).
94 MADRONO [Vol. 18
NS x 2 bé f Jones 27361)
x1 Gentry LLY 6 | 14 mL
Fic. 1. Distribution of Buddleia corrugata; ssp. corrugata, closed circle; ssp. gentryi, open circle; ssp. moranii, open square.
BUDDLEIA CORRUGATA ssp. moranii Norman, ssp. nov. Frutex 0.5-1.0 m altus: folia lineares 0.5-2.0 cm longa, 0.2—-0.3 cm lata, subsessiles, utrinque erecta stellato-tomentulosa, marginis aliquantum involutis, apice ovata, basi attenuata; inflorescentia 3-6 juga cymarum capitarum
; |
1965 | NORMAN: BUDDLEIA CORRUGATA )5
Fic. 2. Trichomes of underside of leaves of Buddleia corrugata; A, ssp. corrugata (Carter 4570) ; B, ssp. gentryi (Gentry 4446) ; C, ssp. moranii (Moran 11695).
sessilium breviterve pedunculatarum ferente, capite quoque 0.5—0.7 cm lato, 3-5 floro; calyx tubiformis 3.0 mm longus, corolla aurantiaca, cam- panulata tubo 3.0-3.5 mm longo intus piloso, extus stellato-tomentello, lobis orbiculatis patentibus 1.0 mm longis; capsula ovata 2.5-3.0 mm longa dehiscens; semina ovata, non alata 0.5 mm longa.
Differing from ssp. corrugata and genryi by its much narrower linear leaves and shorter inflorescences.
Las Tres Virgenes Volcano, central Gulf Coast Region (fig. 1). Flower- ing in the spring.
96 MADRONO [Vol. 18
Type: Volcan las Tres Virgenes, on cliff, steep arroyo on east slope from 1600- 1900 m, Lat. 27° 28’ N, Long. 112° 36’ W, Feb. 12, 1964, Moran 11695 (SD!).
Specimen examined. Crater of Volcan las Tres Virgenes, 1850 m, Moran 11690 (SD).
It is very interesting to note the parallelism in leaf shape that has taken place in corrugata ssp. morani and B. utahensis. Both have evolved in similar habitat—the latter is limited to the Majave Desert at elevations of 800-1800 m, in an area where the mean annual rainfall rarely exceeds 15 cm (Clokey, 1951); it grows mostly on limestone and volcanic cliffs. The former, as already noted is known only from the Volcan las Tres Virgenes at elevations of 1600-1900 m, an area which lies in the driest subdivision of the Sonoran Desert with an annual mean rainfall of 13.8 cm (Shreve and Wiggins, 1964). I believe that this de- velopment in leaf structure is an instance of parallel evolution as it is very likely that B. utahensis and B. corrugata were both derived from B. marrubufolia stock.
Buddleia corrugata closely resembles B. marrubtfolia of Texas and northern Mexico, differing in the size of the seed and the shape of the capsular valve. The westernmost element of that species, which seems to be a relict, possesses an inflorescence of several heads as does B. cor- rugata. This type of inflorescence is thought to be more primitive than the solitary head characteristic of B. marrubifolia ssp. marrubiufolia.
Judging from their great similarity, it would seem likely that B. cor- rugata and B. marrubufolia had a common ancestry. It is not known when Buddleia migrated to Baja California; it may have been in early Miocene or earlier when it is believed that the Gulf of California ex- tended north only to the latitude of Tiburon Island. At that time direct interchange would have been possible between the Sonoran-Chihuahuan Desert and the peninsula (Wiggins, 1960).
I am indebted to Annetta Carter and Reid Moran for making their collections available to me and for helpful suggestions.
Department of Botany, Rutgers—The State University, New Brunswick, New Jersey
LITERATURE CITED
CLoxey, I. W. 1951. Flora of the Charleston Mountains, Clark Co., Nevada. Univ. Calif. Publ. Bot. 24:1-274.
DuruamM, J. W. and E. C. ALrtson. 1960. The biogeography of Baja California and adjacent seas. The geologic history cf Baja California and its marine faunas. Syst. Zool. 9:47-91.
SAVAGE, J]. M. 1960. The biogeography of Baja California and adjacent seas. Evolu- tion of a peninsular herpetofauna. Syst. Zool. 9:184—232.
SHREVE, F. and J. L. Wiccrins. 1964. Vegetation and flora of the Sororan Desert. Vol. 1. Stanford Univ. Press.
Wiccins, I. L. 1960. The biogeography of Baja California and adjacent seas. The origin and relationships of the land flora. Syst. Zool. 9:148-165.
ADRONO
VOLUME 18, NUMBER 4 OCTOBER, 1965
Contents
Dr. LE Roy AsraAms’ COLLECTING TRIP OF 1908, Ernest Alexander McGregor 97
THE FLESHY-FRUITED GALIUM SPECIES OF CALIFORNIA (RUBIACEAE). I. CYTOLOGICAL FINDINGS AND
SOME TAXONOMIC CONCLUSIONS
Lauramay T. Dempster and G. Ledyard Stebbins 105
A FURTHER DESCRIPTION OF GOSSYPIUM TRILOBUM,
Paul A. Fryxell 113 NEw TAXA IN THE GENUS MONARDA (LABIATAE),
Rainer W. Scora 119 DocUMENTED CHROMOSOME NUMBERS OF PLANTS 122
Reviews: Ida Kaplan Langman, A Selected Guide to the Literature of the Flowering Plants of Mexico (Annetta Carter) ; Beatrice F. Howitt and John Thomas Howell,
The Vascular Plants of Monterey County, California (John H. Thomas) | 126
NOTES AND NEws CALIFORNIA NATIVE PLANT Society; 104 LABELS FOR HERBARIUM SPECIMENS, John H. Thomas ; THE HERBARIUM OF Howarp E. McMinn, Baki Kasa- pligil; NEw PUBLICATIONS 127
A WEST AMERICAN JOURNAL OF BOTANY
HBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
Entered as second-class matter at the post office at Berkeley, California, January 29,
1954, under the Act of Congress of March 3, 1879. Established 1916. Subscription price $6.00 per year. Published quarterly and issued from the office of Madronio, Herbarium, Life Sciences Building, University of California, Berkeley, California 94720.
BOARD OF EDITORS
EpcArR ANDERSON, Missouri Botanical Garden, St. Louis LyMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis HERBERT F, CopELAND, Sacramento College, Sacramento, California Joun F. Davipson, University of Nebraska, Lincoln WALLACE R. Ernst, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California MILpreD E. Maturitas, University of California, Los Angeles RoBERT ORNDUFF, University of California, Berkeley MARION OWNBEY, Washington State University, Pullman REED C. Rotiins, Gray Harbarium, Harvard University IrA L. Wiccins, Stanford University, Stanford, California
Editor—JouHN H. THOMAS Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC. President: Annetta Carter, Department of Botany, University of California, Berke- ley. First Vice-President: Winslow R. Briggs, Department of Biological Sciences, Stanford University. Second Vice-President: Arthur R. Kruckeberg, Department of
Botany, University of Washington. Recording Secretary: Mary L. Bowerman, De- ©
partment of Botany, University of California, Berkeley. Corresponding Secretary, Margaret Bergseng, Department of Botany, University of California, Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State College.
|
1965] McGREGOR: ABRAMS’ COLLECTING TRIP 97
DR. LE ROY ABRAMS’ COLLECTING TRIP OF 1908
ERNEST ALEXANDER MCGREGOR
In spite of the fact that I was registered as a Zoology major at Stan- ford University, I had by 1908 taken Botany courses every semester since matriculating. Incidental to receiving an appointment as assistant curator of the Herbarium, Professor Dudley informed me that I had taken more botanical courses than any previous student, to that time. This may have had something to do with the invitation of Dr. Abrams for me to accompany him on a botanical collecting trip during the summer of 1908 (fig. 1) to the mountains and Mojave Desert of Southern California.
Our trip commenced soon after the end of the spring semester and we made our first collection, Calochortus clavatus, on May 31 near Santa Paula. Our equipment was as frugal as consistent with the needs of successful collecting. Each of us had made a simple sleeping bag. A supply of provisions, such as desiccated soups, potatoes, rice, tea, coffee, and a few other needed supplies were shipped ahead to Hot Springs Camp, a short distance above Santa Paula. Naturally, a good supply of newspapers and heavy blotters were included. At the Springs we purchased six burros, two for riding and four for transporting equipment and specimens.
The camp wrangler gave us a hasty lesson in the tying of the diamond cinch, for securing the burden on the burros. The camp supplied a quantity of rolled oats, intended for inducing the untethered burros to stay near camp at night.
Our first day we headed toward Hines Peak and the Topatopa Moun- tains. As we were picking our way along a narrow mountain trail in the Topatopa Mountains (fig. 2), we were met by a typical old pros- pector who was laughing heartily. Said he: “‘Who showed you how to tie the diamond cinch?” A glance revealed that the packs of three of the burros had swung down, and were suspended under their bellies. The kind old fellow unpacked and repacked the loads, taking care to show us the secret of a good diamond cinch. The main idea is to pull the final tug of the rope so tight that the belly-band sinks into the animal’s belly. When you hear the burro groan, the cinch is then just right.
On June 12 we camped near the base of Mt. Pinos. Being a hot day, we climbed to the summit of the mountain. Dr. Abrams and I stripped naked and bathed with handfuls of snow. We spent parts of two days at this location and found the collecting very good. We saw
Mapron 0, Vol. 18, No. 4, pp. 97-128. November 23, 1965.
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1965] McGREGOR: ABRAMS’ COLLECTING TRIP 99
Fic. 2. McGregor leading the pack burros through the Topatopa Mountains.
trout in the pools, but had no fishhooks. We bent common pins, and attached them to stout thread and used grasshopper bait. To our great surprise we caught enough fish for a fine meal!
As we traveled through the domain of the limber pine, Pinus flexilis, we were impressed by their stunted and distorted condition. Little did we then suspect that the students of this tree would discover, many years later, that some of the limber pines are older than the big trees of the Sierra Nevada. It is claimed that some of them are over 4000 years old.
Continuing northeastward we collected at Fort Tejon and Gorman and marvelled at the profusion of noble valley oaks on the high plateau near the Fort.
Entering the desert (Antelope Valley), we inquired of an old pros- pector as to where we could find water at the eastern border of this arid area. He assured us that there was a spring at the eastern end of the desert trail. So we pressed on, collecting a good supply of desert species. As we were about to pass Neenach, a teacher, seeing our six burros, came out of the little one-room schoolhouse and she inquired if we would sell her a burro. She had a small boy of her own, and wanted the animal as a playmate for him. Our “Outlaw” was well named.
Fic. 1. Route of Abrams and McGregor through Southern California in 1908.
100 MADRONO [Vol. 18
Fic. 3. Tehachapi Pass.
He had a mind of his own, and was a cause of some trouble and worry. He had a good physique, however. After some inspection—and with no discouragement from us—the lady agreed to purchase, and we sold him at cost.
At the end of our hot trek across the Antelope Valley, we reached the base of a foothill, where the old prospector had assured us we would find the locale for a good camp, and water. We soon spotted a spring, but at its bottom was a collection of drowned and putrid desert rats. Dr. Abrams and I discussed the situation, and we decided there was nothing to do but use the water. So we boiled potatoes and made coffee with the ill-smelling liquid! Next morning we again used the water for coffee and cereal. As we started toward Mojave via the Tehachapi Pass (fig. 3) and about 100 yards from where we had camped, we came upon a beautiful spring of crystal pure water. Many a time Abrams and I joked about this incident.
The hike to Mojave was on a very hot, trying day and, as I recall, we collected a minimum of plants. As we were struggling along, with parched throats, two men on a buck-board overtook us and stopped to pass the time of day. Probably noting our discomfort, one of them asked: “How would you like a cool bottle of beer?” Not waiting for an answer, he reached into an ice container and handed each of us a bottle. That was another incident that Abrams and I often mentioned in later years.
In 1908 Mojave was little more than a section-point, where the Santa
1965] McGREGOR: ABRAMS’ COLLECTING TRIP 101
Fe trains took on water and changed crews. As we and our five burros trudged into town, we wondered where to make camp. We were a dirty, unshaven, doubtful looking pair, and dreaded the reception we might receive, even at the questionable looking tavern. We came upon a horse corral, deep with straw and manure and supporting a million flies. In desperation, we pitched camp there.
Since the start of our trip at Santa Paula, we had subsisted entirely on our crudely prepared camp meals and here at Mojave was a Harvey Cafe! Why not buy a real meal? So, in hobo attire, into Harvey’s we tromped. A group of properly dressed gents eyed us, and one of them said: ‘‘Look, see what’s coming!”’ That, of course, was another incident that Abrams and I recalled.
Next day we headed for Lancaster, across the Mojave Desert, abound- ing with Joshua trees, Yucca brevifolia. This was to be a two-day hike with a night camp at a small ranch. After a broiling-hot day, it seemed good to crawl into our sleeping bags and in the cool of the night gaze up at the millions of crystal-clear stars.
As we were about to fall asleep, I heard Abrams say: ‘“‘What’s that noise I hear?”’ Then I too heard the somewhat alarming noise. We arose and with a flashlight crept toward the point of the sound. There was a domestic cat holding its paw as if in self-protection. Suddenly our light revealed the cause of the noise—a rattlesnake coiled to strike. We found a suitable stick and killed the rattler. In the morning we saw that the coiled snake had been about two feet from our heads—in the sleeping bags. There was no trace of the cat.
That morning we noted that our food supply was getting scarce and we planned to stock up at Lancaster. We found botanical material much more abundant than one would expect in a desert and we secured a gratifying number of specimens. As we toiled along toward the western border of the desert, we were a sweating but jolly pair. Dr. Abrams could hardly be called the athletic type and he rode nearly the full distance of the summer trip. Having run the mile on the Stanford track team, with many an hour of training, it was perhaps natural that I preferred walking to sitting astride a groaning little burro.
As the hours passed in our trek toward Lancaster, the continuing sameness drugged us into a state of semi-hypnosis. Early that day I had asked Abrams for the technical name of a composite that abounded as we traversed the Mojave Desert. He informed that it was Hymen- oclea salsola, burrobrush, and in my drowsy condition, I began chant- ing: “Hy-men-o-clea sal-so-la.” I must have chanted these names inter- mitently for hours and finally Abrams joined me in the chanting. If anyone had come upon us as we chanted, they might have questioned our sanity.
As we came to the outskirts of Lancaster, we made camp near a factory of some sort. Upon finishing our desert supper, a man approached us and presented us with a bottle of cool claret. He was an Italian worker
102 MADRONO [Vol. 18
Fic. 4. Abrams examining Pinus flexilis at the summit of Mt. San Gorgonio.
in the factory, a modest fellow, and when we thanked him feelingly for the very welcome gift, he bowed and said it was not much. Though Abrams might have been classified as an abstainer, I recall that we finished the bottle and slept the sleep of weary but contented travelers.
One of our problems was to be able to ship our accumulation of plants often enough to adjust to our active collecting. The hot dry climate of many of localities we visited, favored rather speedy drying of the plants. It often was a considerable distance between express offices and at times the little burros became heavily laden by the time we reached a shipping point. Thus we were glad to unload at Palmdale. It is well to explain here that arrangements had been made with Miss Josephine Randall, then a zoology major, to care for the plants as they arrived at Stanford University. Luckily, she was a very careful and dependable person.
From Palmdale we travelled through Cajon Pass to Lake Arrowhead and Big Bear Lake, in both of which localities the collecting was good. We next visited Dry and Dollar lakes. The latter was a very small, circular body of water near the base of Mt. San Gorgonio at an eleva- tion of about 9300 feet. It had a very attractive setting and we could not resist making it our camp for parts of two days. At such an elevation we found a limited but very interesting flora and we collected with enthusiasm.
While at Dollar Lake, we had a sudden inspiration to hike to the summit of Mt. San Gorgonio. At best, the trail was rather poor, and
1965] McGREGOR: ABRAMS’ COLLECTING TRIP 103
Fic. 5. Matilija poppy, Romneya trichocalyx, along Sespee Creek, Ventura Co.
gradually deteriorated as we climbed until, at about the 10000-foot level, the trail vanished and we had to pick our way the remaining distance. The view from the summit (fig. 4) of the highest mountain in Southern California (11485 feet) was inspiring! We could see the Imperial Valley and the northern zone of Baja California. We were well above timber line and with the exception of stunted Pinus flexilis, the mountain was practically devoid of flora.
On return to Dollar Lake, a look at our map indicated that it was about 30 miles to Redlands, our final point of travel. Planning to take parts of two days to reach our destination, we followed Mill Creek Canyon at an easy pace for man and animal, collecting as we went. Reaching Redlands, our purpose had been accomplished and the final shipment of specimens was dispatched. At Redlands we found a tourist outfitter who purchased our five burros, “Ari”, “Zona’’, “Razor”, “Maybe”, and “Groaner”. They had served us patiently and effectively. We received nearly as much for them as they had cost us originally. Considering the net outlay and the total mileage covered (estimated 400 miles), the burros furnished us very cheap transportation. During the six weeks of our intimate association with the peaceful, friendly animals, Abrams and I could not but help forming a sort of fellowship with them. For my part, I felt that I owed them a debt for the con- tribution they made toward obtaining the botanical specimens.
104 MADRONO LVol. 18
Upon returning to the Stanford Campus, we found that our many days of rugged toil had been made more valuable by the final attention given to the material by Miss Randall upon receipt.
As we gathered plants on the trip, we tried to collect four or five specimens of each species at each locality. Thus there were several more or less complete sets in the entire collection. Stanford retained a complete set and the others were sold to the herbaria of the New York Botanical Garden, the Smithsonian Institution, the Royal Botanical Garden of Edinburgh, the Jardin botaniques in Genéve, Harvard University, and the University of Zurich. Dr. Abrams very generously split the balance with me after all expenses were deducted. My half, as I recall, amounted to roughly $75.
During the entire trip, no accidents occurred and neither Dr. Abrams nor I incurred any physical ailments. I don’t seem to recall any rainfall that caused any worry. All that can be said for our food and cooking is that it didn’t poison us and it did maintain our bodily functions. With the exception of a week’s supply of rolled oats, the burros kept in seemingly good condition by grazing upon the grass and other burro eatables they could find in and around camp. What plant impressed me the most during the entire trip? I am inclined to select the matilija poppy, Romneva trichocalyx (fig. 5).
I would be very remiss were I not to record that I cannot recall any unkind attitude or remark ever occurring throughout the entire duration of the collecting trip. To characterize Dr. Abrams in the vernacular, he was a “darn good scout,” and an ever pleasant companion. I have thought and spoken of the trip many times. It was one of the high spots in my life!
128 S. Washington Ave., Whittier, California
NOTES AND NEWS
CALIFORNIA NATIVE PLant Society.—The rapid population growth in Cali- fornia will inevitably result in the reduction or even extinction of endemic plants in many parts of the State. The California Native Plant Society was established in July to counteract this threat to the native flora. This is a statewide, non-profit, educational, and charitable organization specifically concerned with the conservation of the California flora. It will cooperate with other conservation groups in this task. The organization consists of members at large and a program involving the forma- tion of local chapters is being initiated. The sponsors of the California Native Plant Society include a number of distinguished California citizens. Information on the programs of the Society can be obtained from P. O. Box 397, Berkeley, California 94701 or from the Jepson Herbarium, University of California, Berkeley, California 94720. Membership dues include a monthly newsletter and a quarterly publication.
i \
THE FLESHY-FRUITED GALIUM SPECIES OF CALIFORNIA (RUBIACEAE).
I. CYTOLOGICAL FINDINGS AND SOME TAXONOMIC CONCLUSIONS
LAURAMAY T. DEMPSTER AND G. LEDYARD STEBBINS
Preliminary work by the senior author, done under the auspices of the Jepson Herbarium, demonstrated the presence of morphological, cytological and taxonomic complexity in the fleshy-fruited species of Galium of California. It was apparent that only by further study of the chromosomes would it be possible to determine the interrelationships between the many existing forms. This study was undertaken in 1963 and 1964, supported by the National Science Foundation, grant number GBo649. It has involved an intensive and extensive field investigation, including visits to a number of areas where the plants had not been previously collected. Chromosome counts were made chiefly from somatic cells of leaf buds, either preserved in the field or taken later from transplanted individuals. Material was pretreated with a saturated aqueous solution of paradichlorbenzene before fixing, as a result of which pretreatment the shape of the chromosomes as seen in the figures has been considerably altered.
We wish to thank Rimo Bacigalupi, Curator of the Jepson Herbarium, for generous provision of laboratory space, and for critical reading of the manuscript.
The plants under consideration are dioecious, have four “leaves” to a node (with the exception of G. hardhamiae and sometimes G. cle- mentis), and bear fleshy fruits, a character unusual in this genus. The species that comprise the group are G. californicum H. & A., G. nut- talii Gray, G. bolanderi Gray (including G. pubens Gray), G. andrewsii Gray, G. ambiguum Wight, G. sparsiflorum Wight, G. miguelense Greene, G. muricatum Wight, G. clementis Eastw., G. grande McClatch., and G. hardhamiae Dempst.
The group is restricted to California, southern Oregon and northern Baja California. Within this area it occupies most of the cismontane hilly and mountainous areas, from ocean bluffs at near sea level to almost 8000 feet in the Sierra Nevada.
Considered from the traditional morphological and geographic points of view, the species have in the past proven difficult to delimit. Many specific names have been published to cover the various forms, and all attempts to evaluate these by classical taxonomic means have been un- satisfactory, owing to the great diversity of forms within what appeared to be the chief divisions or species, and to marginal resemblances be-
105
106 MADRONO [Vol. 18
tween these species. Preliminary chromosome counts showed the exist- ence of polyploidy, and an attempt has therefore been made by further cytological study to arrive at a clearer understanding of the taxonomic divisions, and eventually of the evolutionary pattern within the group.
The present paper has as its purpose the publication of chromosome numbers (table 1, figs. 1, 2) together with some taxonomic conclusions to be derived therefrom. Our new knowledge of ploidy levels within the group has clarified a number of previously obscure taxonomic lines, such as those between G. californicum and G. nattallii, and between G. bolan- deri and G. sparsiflorum or G. nuttalli. It is hoped, therefore, that a sys- tematic revision based on these new data will shortly follow. Voucher specimens for all chromosome counts will be deposited in the herbarium of the University of California at Berkeley, except those already in the Jepson Herbarium.
The fleshy-fruited Galium complex is now known to consist of one widely distributed and polymorphic diploid species, G. nuttalli, at least six other more restricted and more uniform diploid species, and at least ten polyploids, most of which are apparently hybrids. All of the diploids can be related readily to one or more polyploids and vice versa, the result being a network of polyploids and diploids in which most, or perhaps all, of the taxa are connected to each other, directly or indirectly.
In the following paragraphs, some specific taxonomic conclusions are set forth, and two new taxa are described.
The epithet pubens, derived from Galium pubens Gray, the applica- tion of which has always been vague and uncertain, is now clearly seen to be unnecessary. Field observation has repeatedly shown us that very pubescent forms referable to G. pubens grow in very close association or even in contact with glabrous or glabrate forms of G. bolanderi, and that highly diverse populations are common, containing plants of all degrees of pubescence. Since we now know that all of these varying elements of such populations are hexaploid, there seems no good reason to doubt that they are merely genetic segregates in freely interbreeding populations of G. bolanderi.
Galium grande McClatch., which has sometimes been confused with G. pubens in the herbarium and in the literature, is not only quite dis- tinct in the field, owing to its more fleshy leaves and its lavishly sprawl- ing habit, but is now known to be a very high polyploid, perhaps 20x. It must therefore stand as a species. It was subordinated to G. pubens as a variety by Jepson (1925), Munz (1959) and Ferris (1960), and in the latter instance was defined so as to include much pubescent material of G. bolanderi from the southern Sierra Nevada.
Galium sparsiflorum Wight is now known to be a diploid species, a fact which greatly eases the matter of differentiating it from some forms of G. bolanderi. The latter species is always hexaploid, although some individuals resemble the diploid taxa G. sparsiflorum or G. nuttallii. The inference is very strong that one genome from each of the latter two
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1965 | DEMPSTER & STEBBINS: GALIUM 107
species is present in G. bolanderi, together with a third genome from some pubescent species. True G. sparsiflorum can be distinguished from sparsifloroid individuals of G. bolanderi by the narrow leaf insertion, the very slender pedicels, and often by the leaf texture, which may be thin or somewhat coriaceous, but is never soft and thick as is common with G. bolandert.
Galium muricatum Wight, which was included under G. californicum by Jepson but was recognized as a species by Munz and by Ferris, is clearly a separate species. It has been observed repeatedly by us in the field, growing in closest association with G. californicum without inter- grading. Furthermore, cytological study shows the two species to be far apart in their chromosome numbers, G. californicum being, except for the very local ssp. luciense, highly polyploid, and G. muricatum of low ploidy. The latter is apparently always diploid south of Albion, Mendocino Co., and tetraploid north of Fort Bragg. No consistent dif- ference between the diploid and tetraploid series of populations has yet been found.
Galium cliftonsmithii (Dempster) Dempster & Stebbins, comb. nov. Galium nuttalli Gray var. cliftonsmithi Dempster, Brittonia 10:183. 1958. Studies since 1958 indicate G. cliftonsmithu to be a high polyploid, while G. nuttallu is consistently diploid throughout its many forms. The closest relative of G. cliftonsmithu is probably G. californicum, especially the typical form from the Monterey coast, but the large, shiny, coriaceous and acerose leaves, vigorous habit and higher chromosome number of G. cliftonsmithi entitle it to specific rank.
GALIUM CALIFORNICUM H. & A. ssp. luciense Dempster & Stebbins, ssp. nov. Plantae perennes dioicae humiles (usque ad 16 cm), caulibus pluribus e rhizomatibus tenuibus diffusis orientibus, ubique molliter pubescentes pallide virides; folia pusilla, 4-6 (10) mm longa, plana vel leviter revoluta, carnosula, elliptica vel leviter obovata, apicibus inermi- bus; corollae rotatae, galbinae; ovaria dense pubescentes; fructus albi carnosi; chromosomata 44.
Type. Cone Peak, Monterey Co., California, T. 21 S., R. 4 E., Oct. 12, 1960, at about 4000 feet, Dempster & Hardham 1403 (JEPS 24912). Named for the Santa Lucia Mountains.
Additional collections Monterey Co.: Cone Peak trail ca. 34 mile north of road- head, ca. 4500 feet, Dempster & Stebbins 3921. Specimens from the ridge between Mill Creek and Alder Creek, Monterey Co., Hardham 3785, with very small stomata, appear to be of this subspecies.
The plants are not conspicuously tufted, the many annual stems arising separately from extensive underground rhizomes, clones often covering many square feet. Apparently of very local distribution, ssp. luciense grows in forest duff or gravelly talus in partial shade, with G. clementis on Cone Peak, Monterey Co. It was supposedly derived as an allotetraploid from G. clementis with some other diploid. It shares with G. clementis the distinctive light color and dense, soft long hairs. It is readily distinguished from that species, however, by its habit, which
108 MADRONO
Fr SNe = 2 1 LG 244 stn Te Eo SAR sauze ins a 4a WS 4b ~ Pa) Kas i, } 7 Ps oe nN g ee Re” ott 6a oe? oye o) ends ayo 9% o e é 60 y & ee
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[Vol. 18
Fic. 1. Camera lucida drawings showing chromosome numbers in species of Galium (xX ca. 2000). No. 1 is meiotic from P.M.C., the rest mitotic from shoot ter- minals. 1, G. sparsiflorum, 3807, n = 11. 2, G. clementis, 1402, 2n = 22. 3, G. nut- tallii, 3968, 2n = 22. 4a, G. muricatum, 3944, 2n = 22; 4b, G. muricatum, 3957, 2n = 44. 5, G. martirense, 3987, 2n = 44. 6a, G. andrewsii, 6151, 2n = 223 6b, G. andrewsii, 3983, 2n = 44; 6c, G. andrewsii var. gatense, 4079, 2n = 88. 7, G. cliftonsmithi, 3970, 2n = ca. 187. All mitotic chromosomes were pretreated with
saturated aqueous solution of paradichlorbenzene.
1965 | DEMPSTER & STEBBINS: GALIUM 109
“4 P o oe a@. Nr» fi la SS, AYO = Lx ay, Ava co 2 048 SSI EAE ares Nene FG A oF cl Ve 10 1a ge xy oe 2 WV by PAE fe i -— AS eed | Def Zi Pe 35° se SS : ~ é i 2a c 2b = 82s ta os as Jrfeo% Rar & & we si v., 3s, 39st, “Zr Se 509 sSaSe 2a, O° WE, we Langs were” g5:5'Ce <i ms ad 3 ae a Pd see oat 2g, = 0 ~ y ~~ & . Q \ ar: {= d wD zoe ° gn 207% As PIE Sre hae cP Gad a> @ ry Mee bacysee tgs 2,76 \ a, ) SS, «Paes ~ise oly Ao Ws ~
Fic. 2. Camera lucida drawings showing chromosome numbers of species of Galium (x ca. 2000). No. 4 is meiotic from P.M.C., the rest mitotic from shoot terminals. la, G. bolanderi, 3806, 2n = 66; 1b, G. bolanderi (“G. pubens”), 3797, 2n = 66. 2a, G. ambiguum, 4034, 2n = 22; 2b, G. ambiguum var. siskiyouense, 3913, 2n = 66. 3a, G. californicum subsp. luciense, 3921, 2n = 44; 3b, G. californi- cum, 3979, 2n = 88; 3c, G. californicum, 3898, 2n = 132. 4, G. grande, 3975, Anaphase I, 1 pole, n = ca. 110. All mitotic chromosomes were pretreated with saturated aqueous solution of paradichlorbenzene.
110 MADRONO [ Vol. 18
is generally lower and much less tufted; by the shape of the leaves, which are considerably broader and not notably revolute; and by the number of leaves, which is invariably four, whereas G. clementis often bears five or six to a node. Subspecies /uciense is, however, less easily distinguished from some forms of octoploid G. californicum in the Cone Peak area. Although ssp. /uciense is relatively uniform in appearance, the octoploid has many forms throughout its range. On the lower slopes of Cone Peak it sometimes resembles ssp. /wciense rather closely.
On the top of Cone Peak, G. clementis is found growing alone. A little lower down, it grows with G. californicum ssp. luctense. Still lower, G. clementis drops out and octoploid G. californicum makes its appear- ance. Lower still ssp. /uciense drops out, leaving only the octoploid in diverse forms, some very like ssp. luciense, a few with narrow leaves suggestive of the deep-shade form of G. clementis, but the greater part broad-leaved, flaccid and easily distinguished from both the diploid (clementis) and the tetraploid (duciense). The new tetraploid taxon may be supposed from its very close morphological resemblance to many forms of G. californicum to have been one of the ancestors of that species, and to give us a clue as to the way in which the diploid species G. clementis may have contributed to the composition of the modern polymorphic species known as G. californicum. It is for this reason, and because of the practical difficulty of differentiating it from some forms of octoploid G. californicum, that we have described it as a sub- species of this complex polyploid, rather than as a species.
Galium martirense Dempster & Stebbins, sp. nov. Plantae perennes dioicae 12—50 cm altae, e radicibus robustis caespitosae, caulibus ple- rumque tenuibus lignosis vix scandentibus, ubique molliter pubescentes pallide virides; folia 5-12 mm longa, apicibus inermibus, plerumque elliptica vel anguste ovata revolutaque, sed nonnumquam latiores vix revoluta; corollae rotatae albo-flavidae extus hispidae; fructus carnosi; chromosomata 44.
Type. Oaks Pasture, 15 miles by crooked road northeast of the Meling Ranch (Rancho San José), about 20 air miles east of San Telmo, Baja California, Demp- ster & Cory 3987 (UC 1199709). Named for the San Pedro Martir Mountains.
Additional collections. Sierra de Juarez: 1 mile south of Rancho Viejo, Moran 9845 (UC), Ensenada (certainly east of), Jones 3692 (UC), Howell (CAS). Sierra de San Pedro Martir: northeast of the Meling Ranch (Rancho San José), east of San Telmo, Dempster 3989 (UC), 3990 (UC), 3991 (UC), Robertson 12 (UC), Moran 10950 (UC).
Habitat: Sierra de Juarez and Sierra de San Pedro Martir, Baja California, at 3000 to 5600 feet; in shelter of trees, shrubs or boulders, with Quercus, Adenostoma, Artemisia, Arctostaphylos, Chrysothamnus.
In growth habit, G. martirense somewhat resembles G. nuttalli, or even more one of its polyploid derivatives, such as G. bolanderi, or coastal forms of G. californicum. The primary stems are usually elongated, with nodes sometimes as long as 8 cm, becoming woody but remaining slender and serving as a scaffold for subsequent herbaceous and more congested
1965] DEMPSTER & STEBBINS: GALIUM 111
fertile growth. The species is most similar to G. grande McClatchie, which it resembles more in the herbarium than in the field. Both species are hairy, and often become very dark when dry, the hairs then appear- ing brownish or white against the blackish chlorenchyma. In habit G. martirense is more tufted and the stems are straighter than in G. grande, which is decidedly weak and sprawling as well as larger. The leaves of G. martirense are generally (although not always) narrower than those of G. grande and are noticeably revolute, in contrast with the broader and fleshier, relatively plane leaves of G. grande. Neverthe- less, a small minority of plants of G. martirense with broader leaves and less obviously tufted habit resemble rather closely, at least in the herbarium, a small minority of plants of G. grande with narrower leaves or stiffer habit. What at first appears to be a very close relationship is thus not supported by closer inspection, since in addition to the dif- ferences in habit and leaves and the wide geographic separation, we find a great difference in chromosome number, G. martirense being of low ploidy (4x) as contrasted with the very high ploidy (possibly 20x) of G. grande.
Galium martirense seems very closely akin to the diploid species G. clementis Eastwood, with which it shares, in most individuals, the narrow, revolute leaves and the pale softly hispid character. Galium martirense is, however, a much larger and stiffer plant in every way, with a fair range of morphological variation. Galium clementis is appar- ently restricted in present times to the northern Santa Lucia Mountains in Monterey County, California, and if it was parental to G. martirense, it must be supposed that it had formerly a much greater range. If G. martirense is an allotetraploid between a clementis-like ancestor and some other diploid, the identity of the second ancestor is not apparent. Resemblances to G. nuttalla Gray, which in some places grows with it are, except for habit, slight.
TABLE I. CHROMOSOME COUNTS IN GALIUM SPECIES OF THE FLESHY-FRUITED GROUP
Collection numbers are those of Dempster, or Dempster and Stebbins, unless otherwise indicated.
G. nuttalla Gray var. nuttalliz, 2n = 22. San Diego Co., Mt. Helix, 3911; Baja California, Socorro Mine, Dempster 3991A ; Santa Cruz I., 4085.
G. nuttalli Gray (other varieties). 2n = 22. Contra Costa Co., Orinda, 3805; Humboldt Co., Bull Creek, 3856, Briceland, 3887, Shelter Cove, 3888; Los Angeles Co., Dalton Canyon, 3978; Madera Co., North Fork, 3813; Marin Co., Tiburon, Stebbins s.n.; Mendocino Co., Albion, 3881, Anchor Bay, 3948, Newport Hutchi- son 887 (JEPS); Monterey Co., Arroyo Seco Public Camp, Bacigalupi 7561 (JEPS); Napa Co., Saint Helena, 3780B, 3780E; San Benito Co., Clear Creek, 4077; San Luis Obispo Co., San Luis Obispo, 3960; Santa Barbara Co., Buellton, 3962, Nojoqui Falls, 3965, Refugio Pass, 3968; Santa Cruz I., 4082, 4088; Trinity Co., Mill Creek, 3841; Ventura Co., Ojai, Dempster s. n. (JEPS); Josephine Co. Oregon, Merlin, Hutchison 2065 (JEPS).
G. sparsiflorum Wight. n = 11. Fresno Co., Shaver Lake, 3818; Madera Co., Whiskey Creek, 3807; Tuolumne Co., Crane Flat, 4035. 2n = 22. Placer Co., Baxter, 1443 (JEPS); Tehama Co., Potato Patch, 4050.
ny MADRONO [Vol. 18
G. hardhamiae Dempst. 2n = 22. San Luis Obispo Co., Cypress Mountain, 5703 (JEPS).
G. clementis Eastwood. 2n = 22. Monterey Co., Cone Peak, 1402 (JEPS), Dempster s. n.
G. andrewsii Gray. 2n = 22. Fresno Co., Los Gatos Creek, 4074, 4075; Kern Co., Bodfish, 1437 (JEPS) ; Monterey Co., Castle Peak, 4066, 4065, 4069; Lake Co., Butts Canyon, Stebbins 6151; Napa Co., Corona Mine, Stebbins 6160; San Benito Co., Clear Creek, 4078; San Luis Obispo Co., Morro Bay, Stebbins s. n.; Tehama Co., west of Paskenta, 3995, Patton Mills, 3997, Colyear Springs, 4006. 2n = 44. Los Angeles Co., Gorman, 3983; San Luis Obispo Co., Kiler Canyon, 4073.
G. andrewsii Gray var. gatense Dempst. 2n = 88. Monterey Co., Parkfield- Coalinga summit, 4072, Table Mountain, 4064 (D1 & D3); San Benito Co., Clear Creek, 4076, 4079, Idria Summit, 4080; Santa Clara Co., Mt. Hamilton, 4062.
G. andrewsii x nuttalli (natural hybrid). 2n = 22. Tehama Co., Round Moun- tain, 4002.
G. ambiguum Wight. 2n = 22. Humboldt Co., Low Gap, 3854; Shasta Co., Harrison Gulch, 4009, Regan Meadow, 4010; Tehama Co., Riley Ridge, 4003, Ball Rock, 4004, Round Mountain, 4001; Trinity Co., southeast of Peanut, 4072, west of Peanut, 4013, South Fork Mountain, 4022, Ruth to Zenia, 4026, Ruth, 4019.
G. ambiguum Wight var. siskiyouense Ferris. 2n = 66. Del Norte Co., Gasquet, 3912, 3913, Happy Camp Road, 3917.
G. muricatum Wight. 2n = 22. Humboldt Co., Manchester, 3896; Mendocino Co., Hendy Redwoods, 3878, Navarro, 3880; Sonoma Co., Stewart’s Point, 3944, 3945. 2n = 44. Humboldt Co., Willow Creek, 3845 (A & B), Larrabee Valley, 3851, Dinsmore’s, 3852, Bull Creek, 3858, Weott, 3860, Orleans, 3919, Boise Creek, 3920, Richardson Grove, 3957 (1 & 2) ; Mendocino Co., Laytonville, 3871, Branscomb, 3950 to 3954, Leggett, 3956 (1 & 3), Willits, 3958.
G. martirense Dempst. & Stebb. 2n = 44. Baja California, above Meling Ranch, 3987.
G. bolanderi Gray (including G. pubens Gray). n = 33. Butte Co., Jonesville, 4048; Humboldt Co., Orleans, Hutchison 2031 (JEPS); Madera Co., Oakhurst, 3806 (11 & 12), South Fork, 3809, 3810; Placer Co., Baxter, 1442 (JEPS); Napa Co., Angwin, 3782A; Plumas Co, Quincy, 1458 (JEPS). 2n = 66. Humboldt Co., Dinsmore’s, 3853, Berry Summit, Hutchison 2010 (JEPS); Lake Co., Glenbrook Resort, 3797B; Madera Co., Oakhurst, 3805™%, 3806 (2, 3, 4, 5 & 7), Whiskey Creek, 3808 (1 & 2), South Fork, 3811, North Fork, 3814; Mariposa Co., Usona, 3820; Plumas Co., Graeagle, 1456 (JEPS); Shasta Co., Hat Creek, 4053; Siskiyou Co., Happy Camp, 3918, Hutchison 2039 (JEPS) ; Trinity Co., Burnt Ranch, 3842.
G. californicum H. & A. subsp. luciense Dempst. & Stebb. 2n = 44. Monterey Co., Cone Peak, 1403 (JEPS), 3921.
G. californicum H. & A. sensu lato. n = 44. Monterey Co., Kirk Creek, 3939. 2n = 88. Los Angeles Co., Glendora Ridge, 3979; Monterey Co., Cone Peak, 3922 to 3928, 3933 to 3935, Nacimiento, 3936, 3937; Santa Barbara Co., Nojoqui Falls, 3963, 3964, Refugio Pass, 3969. 2n = ca. 88. Monterey Co., Cypress Mountain, 1187 (JEPS); Santa Barbara Co., Buellton, 3961. n = 66. Marin Co., Mt. Tamal- pais, Bacigalupi 8880 (JEPS); Trinity Co., Ruth Lake, 4017. 2n = 132. Humboldt Co., Manchester 3898; Mendocino Co., Stewart’s Point, 3907; Monterey Co., Point Lobos, 4081; Santa Cruz Co., Boulder Creek, Stebbins s. n. 2n = ca. 1382. Hum- boldt Co., Bridgeville, 3848, Manchester, 3897; Mendocino Co., Anchor Bay, 3909, Laytonville, 3872, 3955, Bell Springs, 3866, Hendy Redwoods, 3877; Santa Cruz Co., Santa Cruz, Stebbins s.n.
G. cliftonsmithii (Demps.) Dempst. & Stebb. 2n = c. 182-189. Santa Barbara Co., Refugio Pass, 3970.
G. grande McClatchie. n = more than 110. Los Angeles Co., Chantry’s Flat, 3975,
1965 | FRYXELL: GOSSYPIUM 113
SUMMARY
Chromosome counts are published for all mainland species of the fleshy-fruited species of Galzum of California and environs, showing the complex polyploid nature of the group. Some specific taxonomic conclusions are drawn, and two new taxa are described: G. californicum ssp. luciense of the Santa Lucia Mountains and G. martirense of Baja California. A former variety is raised to specific rank as G. cliftonsmithit.
Department of Botany, University of California, Berkeley Department of Genetics, University of California, Davis
LITERATURE CITED
Ferris, R. S. 1960. Galium. Jum L. R. Abrams and R. S. Ferris, Illustrated flora of the Pacific States. Vol. 4. Stanford Univ. Press.
Jepson, W. L. 1925. A manual of the flowering plants of California. Associated Stu- dents Store, Berkeley.
Mowz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
A FURTHER DESCRIPTION OF GOSSYPIUM TRILOBUM
PAuL A. FRYXELL
The exact nature of the Mexican plant originally described by De Candolle (1824) as Ingenhouzta triloba has never been clear, and considerable doubt has been expressed that it is, in fact, distinct from Thurberia thespesioides A. Gray. It is now possible to provide a more complete description and on such a basis to resolve the doubt by assert- ing that these two species are indeed distinct.
De Candolle’s generic name was noted by Tidestrom (in Dayton, 1927) as a variant spelling of Jngenhoussia Dennst. 1818 and therefore to be illegitimate under Article 64. Irrespective of this rejection, how- ever, it is generally accepted that both De Candolle’s and Gray’s plants belong in Gossypium L.
A third name, G. lanceiforme Miers, was subsequently published that is probably based on an isotype of De Candolle’s species (Kearney, 1952). A historical summary of the taxonomic disposition of these three names is presented in Fig. 1.
Mauer’s publication (1954) of three varietal names under G. trilobum contravenes Articles 26 and 36; the varietal names therefore are not validly published.
Gossypium thurberi Tod. (= Thurberia thespesioides A. Gray) is a plant, well represented in herbaria and in living collections, that occurs in southern Arizona, northern Sonora, and parts of western Chihuahua. It has been described and illustrated many times, notably among the
114 MADRONO [Vol. 18
Schumann 1890
(AB) Kuntze 1891 (AB) Watt 1907 (BC ) Ulbrich 1913 (AB) Standley 1923 (A )(B) Tidestrom 1927 (AB) Skovsted 1935 (ABC) Kearney 1937 (A )(B) Hutchinson 1947 Prokhanov 1947 (AC)(B) (A) (BC) Roberty 1950 (ABC) Mauer 1954 (ABC) Fryxell 1965 (AC)(B)
Fic. 1. Historical summary of views on the taxonomic disposition of: A. Ingen- houzia triloba, B. Thurberia thespesioides, and C. Gossypium lanceiforme.
fine drawings published by Saunders (1961). Gossypium trilobum (= Ingenhouzia triloba DC.), on the other hand, is poorly known; its existence is doubted by some and its distribution has been unclear to those who accept the species. It has never been available in culture.
1965] FRYXELL: GOSSYPIUM 115
Gossypium thurberi as
Gossypium yi trilobum
Mexice City
Gossypium gossypioides ——
Fic. 2. Geographic distribution of three species of Gossypium.
°o
Gossypium trilobum has, moreover, been confused to some extent with still a third, relatively dissimilar species, G. gossypioides (Ulbr.) Standl. (= Selera gossypioides Ulbr), because of certain superficial similarities, although the distinction was made sufficiently and explicitly clear by Ulbrich (1913) in his description of Selera gossypioides. Roberty (1950), in fact, placed all three species in synonymy under G. trilobum. Standley (1923) recognized the distinctiveness of G. gossypioides and its position in Gossypium. Kearney (1937) underscored Standley’s view and, except for Roberty (1950), has been followed by subsequent authors, including Hutchinson (1947) who, however, unfortunately cited two specimens under G. gossypioides that in fact represent G. trilobum. Hutchinson’s citation of these specimens was quoted by Mauer (1954). Consequently, detailed comparisons involving G. gos- syptoides are also needed.
Gossypium gossypioides would be as poorly known as G. trilobum were it not that it has become available in culture from the seed col- lections of T. R. Richmond and C. W. Manning in 1946 and C. W. Manning and J. O. Ware in 1948. Brown and Menzel (1952a; 1952b)
116 MADRONO [Vol. 18
TABLE 1. PRINCIPAL SIMILARITIES OF G. TRILOBUM AND G. THURBERI TOGETHER WITH PRINCIPAL FEATURES DISTINGUISHING THEM FROM G. GOSSYPIOIDES.
G. trilobum or G. thurberi G. gossyptoides Investiture of young Stellate, clusters <.1 mm, Soft-tomentose; hairs predom- twigs and petioles appearing lepidote, sparse, inantly single, whitish, up becoming glabrate to 0.5 mm Twigs Angled or ridged Terete Foliar nectaries Present and functional Vestigial Involucral nectaries Present and functional Lacking Bracts 1-2 cm long, distinct 3 cm long, connate prior to anthesis Petal size 2-3 cm long 5 cm long Petal spot Small to vestigial Large, covering half of petal Filament color Cream-colored Dark red Capsules Non-flaring, persistent, with Flaring, abscissing at base of copious hairs on sutures peduncles, lacking suture hairs Seeds 6-10 per loculus, 3-4 mm _ 1-4 per loculus, 5—7 mm long, long, with sparse, minute with tan seed hairs up to pubescence, appearing black 1 cm
TABLE 2. PRINCIPAL FEATURES DISTINGUISHING G. TRILOBUM FROM G. THURBERI.
G. trilobum G. thurberi Leaf dissection About % dissected; sinuses About 9/10 dissected; sinuses acute; lobes ovate-acumi- rounded; lobes lanceolate- nate, 2-31%4 cm broad acuminate, 1—2 cm broad Bracts Cordate-acuminate, entire, 1O— Subulate to cuneate, entire to 12 mm broad, 15-20 mm (sometimes) toothed, 2-4 long mm broad, 8-12 mm long Calyx With few to many (up to 12) Subtruncate, becoming undu- irregular acuminate to aris- late in fruit tate tips up to 4 mm long Petal Pale yellow Cream-colored Fruit 15-18 mm long, 10-12 mm 10-15 mm long, 8-12 mm
broad, 8-10 seeds per loculus broad, 6-8 seeds per loculus
presented genetic and cytological data bearing on the position of this species in Gossypium. Subsequent studies (recently summarized by Saunders, 1961) indicate G. thurberi and G. gossypioides not to be closely related genetically.
Published knowledge of G. trilobum beyond the original description is nearly confined to the two plates and accompanying description pub- lished by Hutchinson (1947) and the few descriptive comments made by Kearney (1937). However, additional specimens of G. trilobum are now known that permit a full (and comparative) description of the species and an indication of its geographical distribution.
The following three species are compared in Tables 1 and 2 and their distribution is shown in Fig. 2. It is worth noting that the speci- mens of G. trilobum that are cited below fall within the relatively narrow altitudinal range of approximately 2500-5000 feet.
1965] FRYXELL: GOSSYPIUM aly
NOMENCLATURE
GOSSYPIUM THURBERI Todaro, Relaz. 120. 1877. Thespesia thurberi Alefeld, Bot. Zeit. 19:301. 1861, name illegitimate. Thurberia thespe- stoides A. Gray, Mem. Am. Acad. 5:308. 1855, not Gossypium thes pe- sioides F. Muell. ex Tod. Relaz. 103. 1877. Hibiscus ingenhousti Kuntze, Rev. Gen. Pl. 1:69. 1891, pro parte. Thurberia triloba Tid. ex Dayt. Proc. Biol. Soc. Wash. 40:120. 1927, pro parte. Gossypium trilobum Skov. Jour. Genet. 31:288.1935, pro parte.
Type. Cocospera-Barbasaqui, Sonora, Mexico, Thurber 914 (US).
Representative specimens. CHIHUAHUA. SW Chihuahua, Palmer 269 (BM); Rio Bonito, LeSueur 792 (LL, TEX, UC, US). SONORA. 18 mi SE of Magdalena, Wiggins 7183 (ARIZ, DS, UC, US); Municipio de Nacore Chico, Muller 3653 (GH, LL, UC, US); Rio de Bavispe, White 4718 (ARIZ, GH, LL, MICH). ARIZONA. Santa Rita Mts., Thornber 186 (ARIZ, UC); Mule Mts., Goodding 3 (ARIZ); Rincon Mts., Blumer 3323 (ARIZ); near Bisbee, Jones 308 (ARIZ) ; Santa Catalina Mts. Ginter s. n. (ARIZ, UC).
GOSSYPIUM TRILOBUM (Mog. & Sess. ex DC.) Skov. Jour. Genet. 31:288. 1935, pro parte; emend. Kearn. Am. Jour. Bot. 24:299. 1937. Ingenhouzta triloba Moc. & Sess. ex DC. Prodr. 1:474. 1824. Hibiscus ingenhousu Kuntze, Rev. Gen. Pl. 1:69. 1891, pro parte. Gossypium lanceiforme Miers ex Britt. Jour. Bot. 31:331. 1893. Thurberia triloba (Moc. & Sess. ex DC.) Tid. ex Dayt. Proc. Biol. Soc. Wash. 40:120. 1927, pro parte.
Type. Sessé and Mocino’s specimen, kept in the Madrid herbarium and illus- trated by Hutchinson (1947). Kearney (1952) notes that the type of G. lancei- forme may be a duplicate of the type of Ingenhouzza triloba DC.
Specimens examined. Guadalajara, Jalisco, Rose & Hay 6284 (GH, US); Con- cordia, Sinaloa, de Ortega 6033 (US); Acatitlan, Mexico, Hinton 5133 (GH, K); Zihuagio, Mexico, Hinton 9701 (MO, UC, US); Zitacuaro, Michoacan, Hinton 13253 (UC, US); km 15 Yautepec-Cuernavaca Road, Morelos, Lundell & Lundell 12498 (LL, UC, US) ; Sinaloa, Mexia 432 (MO).
In addition to the above specimens, Hutchinson’s plate of the type and of the type of G. lanceiforme Miers ex Britt. (Hutchinson, 1947) have been consulted. On this basis the following description is presented.
Shrub, up to 4 m high, nigro-punctate, minutely stellate-puberulent becoming glabrate; twigs distinctly ridged. Leaves 3— (sometimes 5—) lobed, glabrate; lobes ovate-acuminate, up to 3% cm broad; margin minutely ciliate. Petiole glabrate, ridged, nearly equaling lamina. Foliar nectary single, narrowly elongate, 1-2 mm. Stipules subulate to acumi- nate, ciliate, caducous. Fruiting branches many-jointed. Pedicels upright, ridged, glabrate, 1-1'4 cm, surmounted by 3 nectaries. Bracts distinct, cordate, entire, acuminate, 10-12 mm broad, 15-20 mm long. Calyx prominently nigro-punctate with variable number (up to 12) of irregu- lar acuminate to aristate tips up to 4 mm long. Petals punctate, pale yellow with small red basal spot. Androecial column pale-colored, punc- tate; filaments 2-3 mm. Style slender, exceeding androecium; stigma
118 MADRONO [Vol. 18
clavate. Fruit glabrous, nigro-punctate, oblong, beaked, 15-18 mm long, 10-12 mm broad, generally 3-loculed; 8-10 seeds per loculus. Hairs on inner suture of capsule dense, tan, 2 mm long. Seeds angularly turbin- ate, 3-4 mm, black appearing striped because of minute tan pubescence.
GOSSYPIUM GOSSYPIOIDES (Ulbr.) Stand]. Contr. U. S. Nat. Herb. 23:783. 1923. Selera gossypioides Ulbr. Verh. Bot. Ver. Prov. Branden- burg 55:51. 1913. Gossypium trilobum (Moc. & Sess. ex DC.) Skov. emend. Rob. Candollea 13:30. 1950, pro parte.
Type. Im Gebtisch, San Bartolo Yautepec, Oaxaca, C. & E. Seler 1700 (location of specimen not ascertained).
Specimens examined. Cuicatlan, Oaxaca, Nelson 1704 (GH, US); Conzatti 3874 (US); cultivated plants grown from collections from 83 miles S of Oaxaca, Rich- mond & Manning s.n. (ARIZ, TAES).
U.S. Department of Agriculture, Soil and Crop Sciences Department, Texas A & M University, College Station
LITERATURE CITED
Brown, M. S. and M. Y. Menzev. 1952a. The cytology and crossing behavior of Gossypium gossypioides. Bull. Torrey Club 79:110-125.
. 1952b. Additional evidence on the crossing behavior of Gossypium gos- sypioides. Bull. Torrey Club 79:285-292.
Dayton, W. A. 1927. New names for some western montane plants. Proc. Biol. Soc. Wash. 40:119-122.
Hutcuinson, J. B. 1947. The classification of the genus Gossypium. Jn Hutchinson, J. B., R. A. Silow, and S. G. Stephens. The Evolution of Gossypium. Oxford Univ. Press, London.
KEARNEY, T. H. 1937. Ingenhouzia and Thurberia. Am. Jour. Bot 24:298—300.
. 1952. Notes on Malvaceae II. Leafl. West. Bot. 6:165-172.
KuntTzeE, O. 1891. Revisio generum plantarum. Vol. I.
Mauer, F. M. 1954. Origin and systematics of cotton. [In Russian.] Tashkent.
PROKHANOV, Y. I. 1947. Conspectus of a new system of Gossypium L. [In Russian. ] Bot. Zhur. 32:61-78.
RosBerty, G. 1950. Gossypiorum revisionis tentamen. Candollea 13:9-165.
SAUNDERS, J. H. 1961. The wild species of Gossvpium. Oxford Univ. Press, London.
SKOVSTED, A. 1935. Chromosome numbers in the Malvaceae. I. Jour. Genet. 31:263- 296.
SCHUMANN, K. 1890. Malvaceae. Jn Die Nattirlichen Pflanzenfamilien 3(6): 30-53.
STANDLEY, P. C. 1923. Trees and shrubs of Mexico. Contr. U. S. Natl. Herb. Vol. 23.
UrpricyH, E. 1913. Selera, eine neue Malvaceen-Gattung aus der Verwandtschaft von Gossypium L. Verh. Bot. Ver. Prov. Brandenburg 55:50—54.
Watt, G. 1907. The wild and cultivated cotton plants of the world. Longmans, Green, & Co., London.
NEW TAXA IN THE GENUS MONARDA (LABIATAE)
RAINER W. SCORA
During a biosystematic study of the genus Monarda (Scora, 1964) it became necessary to describe the following new taxa and to make one new combination.
Monarda dressleri Scora, sp. nov. Herba perennis, gracilis erecta, altitudine 60—80 cm, pilis minutis puberulis, simplex vel ramosa; foliorum laminis lanceolatis, 23-45 mm longis, 11—21 mm latis, in apicem acumi- natis, in basem obtusis vel rotundis, marginibus subserratis, supra pubes- centibus pilis minutis appressis, subtus pubescentibus pilis minutis canescentibus, petiolis 4-8 mm longis; bracteis paucis foliaceis, ovatis- lanceolatis, puniceis, glomerulis 20-30 mm latis, paucifloris, monocepha- lis; calycum tubis 8-13 mm longis, 13 venis, pubescentibus pilis minutis canescentibus, in ore dense hirsutis pilis 0.5 mm longis, lobis acutissimis 1.5-2.0 mm longis, glandulis elevatis; collorarum coccinearum purpu- rearum saepius immaculatarum, tubis 25-30 mm longis, labia superiore 9-12 mm longa; staminibus inter fauces corollae 3-5 mm insertis; stylo glabro.
Type. 40 km W of Aldama, Tamaulipas, Mexico, Dressler 1851 (MEXU 47528— holotype, MICH, MO).
Monarda dressleri is a member of the subgenus Monarda and is named for R. L. Dressler who has collected extensively in Tamaulipas.
Monarda dresslert has the habit of the M. fistulosa L. complex, but its corolla resembles more closely those of the larger flowered Mexican species of M. pringle: Fernald and M. bartlettu Standley. It differs from all the other Mexican species in the subgenus Monarda in its corolla color and its larger pollen size of about 20 per cent in diameter. It differs from its near relative MM. pringlei in that it inhabits oak forests, has an evenly expanded corolla with a very pubescent upper lip and a stamen attachment of 3-5 mm depth in the corolla throat, while M. pringlei occurs on calcareous soil, has an abruptly expanded corolla with a slightly pubescent upper lip and a stamen attachment 6—7 mm deep. Monarda dressleri differs from the related M. bartlettii in leaf pubes- cence, in the narrower and shorter leaves, its stamen attachment and in the indument of the calyx lobes. Monarda dressleri is distinguished from M. eplingiana Standley in that the leaves are not membranaceous and the calyx lobes are twice as long and beset with stalked glands, while in M. eplingiana the calyx lobes are non-glandular. From M. mal- loryi Gilly it differs in that the latter grows near the borders of tropical forests, is of a more robust habit, has shorter calyx lobes which are non-glandular.
119
120 MADRONO [Vol. 18
MONARDA CITRIODORA Cerv. ex Lag. The varieties and subspecies of M. citriodora may be identified with the aid of the following key.
Bracts subtending glomerules lanceolate to linear-lanceolate, gradually acuminate to a spinose bristle, not more than 4 mm wide, strongly reflexed from base. ssp. austromontana Bracts subtending glomerules oblong, abruptly acuminate to a spine-like bristle, or attenuate, commonly more than 4 mm wide, spreading horizontally, or forming
a cup-like invulcre, but not reflexed from base.
Foliage light green; inner bracts subtending glomerules densely pubescent with short white or purple hairs; the apex abruptly acuminate, bracts reflexed from middle and thus forming a cup-shaped involucre.
Leaves 25-60 mm long, 8-12 mm wide; bracts mostly wider than 5 mm; calyx tubes 7-14 mm long; unexpanded portion of corolla tube 7-10 mm long, the expanded portion 4-6 mm long; apex of upper corolla lip with 1 min One: Whales eee ee ee ssp. citrtodora var. citriodora
Leaves (12)—20—35—(40) mm long, (3)—5—-10-(14) mm wide; bracts not wider than 5 mm; calyx tubes 57-7 mm long; calyx lobes about 2—3 mm long; unexpanded portion of corolla tube 6-7 mm long, expanded portion 2-4 mm long; apex of upper corolla lip mainly glabrate.
ssp. citriodora var. parva
Foliage dark green; inner bracts subtending glomerules glabrate or sparingly pubescent with short hairs; the apex attenuate, spreading horizontally and not forming a cup-like involucre; calyx lobes rarely ciliate; upper half of corolla lip pubescent with 1-2 mm long hairs.
ssp. citriodora var. attenuata
MONARDA CITRIODORA Cerv. ex Lag. ssp. austromontana (FEpling) Scora, com. nov. M. austomontana Epling, Madrono 3:29. 1935.
MONARDA CITRIODORA Cerv. ex Lag. ssp. CITRIODORA var. parva Scora, var. nov. Herba annua, erecta, altitudine 20-50 cm, caulis simplicis, pills minutibus puberulis; foliorum laminis lanceolatis vel oblongis, maximis (12)—20-35—(40) mm longis, (3)—5—10—-(14) mm latis, in apicem abrupte acutis, in basem acuminatis vel obtusis, marginibus serratis vel subserratis, supra scabratis, subtus hirsutis pilis minutis; petiolis 2-8 mm longis; glomerulis verticillastris; bracteis exterioribus foliaceis, mediis oblongis, erectis, in apicem abrupte acuminatis et ari- statis, pagina adaxilia dense pubescente pilis minutis, basis marginibus ciliatis, calycum tubis 5—7 mm longis, 15 venis, lobis aristatis 2-3 mm longis; corollis 14-19 mm longis, abrupte dilatatis; labia superiore 5—7 mm longa, incisa 0.5—1.0 mm, glabrata, rarissime puberula pilis 0.5—1.0 mm longis; labia inferiore 7-8 mm longa, lobo mediano 1—3 mm longo; staminibus glabris, inter fauces 1-3 mm insertis; stylo glabro; chromo- somis n = 9; oleis e-pineno, y-terpineno, heptanone, nonanale, thymole et carvacrole satis inhaerentibus.
Type. Near Sinton, San Patricio Co., Texas, C. M. Rowell 4977 (WWF).
Specimens examined. Texas. Cameron Co.: Brownsville, Tharp 1196 (US); Ferris & Duncan 3131 (MO); Los Fresnos, Wolff 2368 (TAES). Hidalgo Co.: San Juan, Clover 5 (NY). San Patricio Co.: Welder Wildlife Refuge, Scora 2222 (MICH); E of Sinton, Powell 4977 (WWF); Box 148 (WWF).
1965] SCORA: MONARDA 121
SS a Nae ee i —¢ Y &7 Se SF i “Oh MK
ig y ei f am
\
var. attenuata
/ var. parva se var. citriodora
Fic. 1. Varieties of M. citriodora ssp. citriodora.
This variety is the “fairly well-defined but unnamed form in southern Texas” of McClintock and Epling (1942)
MONARDA CITRIODORA Cerv. ex Lag. ssp. CITRIODORA var. attenuata Scora, var. nov. Herba annua, erecta, altitudine 30-60 cm, caulibus virgatis, pilis minutibus puberulis; foliorum laminis lanceolatis interdum oblongis, maximis 30-65 mm longis, 4-13 mm latis, in apicem acutis vel obtusis, in basem angustatis, marginibus serratis, supra scabratis, subtus puberulis pilis minutis; petiolis 4-28 mm longis; glomerulis ver- ticillastris; bracteis exterioribus foliaceis longioribus, mediis oblongis
122 MADRONO [Vol. 18
rarius lanceolatis, in apicem attenuatis, pagina adaxilia glabrata, raris- sime puberula pilis minutis albidis, basis marginibus ciliatis, calycum tubis 9-13 mm longis, 15 venis, lobis aristatis 3-7 mm longis; corollis roseis saepius immaculatis, 21-30 mm longis, abrupte dilatatis; labia superiore 9-12 mm longa, incisa 1.0-1.5 mm, pubescentium pilis 1-2 mm longis; labia inferiore 12-17 mm longa, lobo mediano 4-6 mm longo; staminibus glabris, inter fauces 2-4 mm insertis; stylo glabro; chromosomis n = 9; oleis a-pineno, y-terpineno, heptanone, nonanale, thymole et carvacrole satis inhaerentibus.
Type. Near Musquiz, Coahuila, Mexico, R. W. Scora 2340 (MICH—holotype, WCR).
Specimens examined. Mexico. Coahuila: Musquiz, Marsh 155 (F); Sierra del Carmen, Canon de Sentenela, Wyend & Mueller 524 (MO).
Monarda citriodora var. attenuata (fig. 1) is distinguished from var. citriodora (fig. 1) mainly by its more horizontally spreading bracts subtending the glomerules, their nearly glabrous adaxialbract surfaces, green coloration and the more attenuate bract apices. The calyx lobes have few or no lateral hairs and the corolla dimensions are larger than those of var. citriodora. Variety parva (fig. 1.) differs from var. attenuata in having the bracts subtending the glomerules cupped and not horizon- tally spreading, in their pubescence, whitish coloration, abrupt acumi- nate bract apices, shorter calyx lobes and smaller overall dimensions of stem, foliage, calyx tubes and corollas. It differs from var. citriodora in the almost always glabrous upper corolla lip, and the reduced corol- la size, calyx tube, calyx lobe, number of glomerules, leaf size and plant height. Living cultures which were grown for several generations in the greenhouses and growth chambers at the University of Michigan Botanical Gardens to find if these differences were genetically fixed or mere environmental variations showed, that all these differences re- mained although internode length increased in var. parva when grown under greenhouse conditions.
University of California, Riverside
LITERATURE CITED
McCurintock, E. and C. Eprinc. 1942. A review of the genus Monarda (Labiatae). Univ. Calif. Publ. Bot. 20:147-194.
Scora, R. W. 1964. Interspecific relationships in the genus Monarda (Labiatae). Ph.D. dissertation (unpublished). Univ. of Michigan, Ann Arbor.
DOCUMENTED CHROMOSOME NUMBERS OF PLANTS (See Madrono 9:257-258. 1949; 17:255. 1964)
Aesculus californica (Spach) Nutt. n = 20. California, Alameda Co., Univ. Calif. campus. R. Lloyd 2633, UC. Counted by R. Ornduff and R. Lloyd.
1965] DOCUMENTED CHROMOSOME NUMBERS 123
A. sylvatica Bartr. n = 20. North Carolina, Durham Co., Duke Forest. R. Ornduff 6377, DUKE. Counted by R. Ornduff and R. Lloyd.
Allotropa virgata T. & G. ex Gray. 2n = 26. California, Humboldt Co., near Hawkins Bar. D. E. Anderson 2447, UC.
Alopecurus aequalis Sobol. 2n = 71. Utah, Summit Co. D. E. Anderson 2790, UC.
Anthoxanthum odoratum L. 2n = 20. California, Humboldt Co., Arcata. D. L. Gray 64001, UC.
Aristolochia californica Torr. 2n = 16,1. California, Yuba Co. C. B. Wolf 1741 (progeny), RSA. Counted by P. H. Raven. Gregory (Am. Jour. Bot. 43:111. 1956) reported 2n = 28 in this species from material “from Rancho Santa Ana Botanic Garden” (Gregory, pers. comm.) and therefore this is possibly the same strain.
Bidens pilosa L. var. radiata Sch.-Bip. n = 24. Florida, Charlotte Co. E. B. Smith 530, KANU.
Bocconia frutescens L. n = 20. California, San Francisco Co., Strybing Arboretum, W. R. Ernst 709, DS. Cultivated; Alameda Co., Univ. Calif. W. R. Ernst 732, DS (progeny from Peru, Machu Picchu, West 6472, UC).
Brodiaea appendiculata Hoover. n = 6. California, Amador Co., Ione. T. F. Niehaus OO WIC,
B. californica var. leptandra (Greene) Hoover. 2 = 12. California, Napa Co., Mt. St. Helena. R. F. Hoover 5904, UC. Counted by T. F. Niehaus. Cultivated UCBG 55.926.
B. clementina (Hoover) Munz. 2n = 16. California, Los Angeles Co., San Clemente I. 7.F. Niehaus 376, UC.
B. coronaria (Salisb.) Engler. 2n = 24. California, Sacramento Co., near Ione. 7. F. Niehaus 512, UC.
B. coronaria var. macropoda (Torr.) Hoover. 2n = 36. California, Marin Co., Drakes Beach, 7. F. Niehaus 535, UC.
B. coronaria var. rosea (Greene) Hoover. 2n = 12. California, Lake Co., Indian Valley, T. F. Niehaus 556, UC.
B. dudleyi (Hoover) Munz. n = 8. California, Tulare Co., Summit Lake, 7. F. Niehaus 611, UC.
B. elegans Hoover. n = 8.California, Nevada Co., Grass Valley. T. F. Niehaus 128, We:
B. filifolia Wats. 2n = 32. California, Los Angeles Co., San Clemente I. 7. F. Niehaus 397, UC.
B. gracilis Wats. n = 8. California, Plumas Co., Spanish Peak, 7. F. Niehaus 587, We:
B. hyacinthina (Lindl.) Baker. n = 28. California, Nevada Co., Bear Valley. 7. F. Niehaus 549, UC; n = 35. California, Yuba Co., near Marysville, T. F. Niehaus 22OR KIC.
B. hyacinthina var. greenei (Hoover) Munz. 2n = 16. California, Butte Co., Table Mt., T. F. Niehaus 372, UC.
B. jolonensis Eastw. 2n = 12. California, Monterey Co., Jolon. T. F. Niehaus 511, Wie.
B. lutea var. anilina (Greene) Munz. n = 20. California, Sierra Co., Gold Lake. T. F. Niehaus 586, UC.
B. pallida Hoover. n = 6. California, Tuolumne Co., Chinese Camp. 7. F. Niehaus & W. Savage 526. Counted by T. F. Niehaus.
Carpentaria californica Torr. n = 10. California, Santa Clara Co., Carnegie Insti- tution. W. R. Ernst 733, DS. Cultivated.
Cathcartia villosa Hook. f. n= 16. California, Alameda Co., Univ. Calif. W. R. Ernst 731, DS. Cultivated.
Chelidonium majus L. 2n = 12. Connecticut, New Haven. K. Chambers 1151, DS.
124 MADRONO [Vol. 18
Counted by W. R. Ernst. Mitosis in anther.
Chrysopsis aspera Shuttlew. n = 18. Alabama, Coosa Co. E. B. Smith 475, KANU.
Chrysosplenium glechomifolium Nutt. 2n = 91. California, Humboldt Co., near Arcata, D. FE. Anderson 2975, UC.
Clematis crispa L. n = 8. South Carolina, Berkeley Co., near Monck’s Corner. R. Ornduff 6412, UC.
Clethra lanata Mart. & Gal. 2 = 81. Mexico, Chiapas, Municipio of Tenejapa. D. E. Breedlove 6947, DS, F. Counted by D. W. Kyhos.
Corydalis lutea DC. n = 28. California. Santa Clara Co., Palo Alto. W. R. Ernst 575, DS. Cultivated.
Dicentra chrysantha (H. & A.) Walp. n= 12. California, San Benito Co., near Idria. W. R. Ernst 521, DS; Fresno Co., Los Gatos Creek road near San Benito Co. line. W. R. Ernst 530, DS. Some cells with an extra fragment.
D. formosa (Haw.) Walp. n = 8. California, Tuolumne Co., Crocker Ridge. W. R. Ernst 764, DS.
D. nevadensis Eastw. n = 8. California. Tulare Co., near Big Meadow Guard Sta- tion. W. R. Ernst 570, DS.
D. ochroleuca Engelm. n = 16. California, Los Angeles Co., Mulholland Drive. P. H. Raven 14859, DS. Counted by W. R. Ernst.
D. speciabilis (L.) Lem. n = 8. California, Santa Clara Co., Stanford Univ. W. R: Ernst 488, DS. Cultivated.
Drymaria debilis Brandegee. 2n = 221. Mexico, Baja California, Bahia de la Con- cepcion. J. L. Wiggins 19001, DS. Counted by D. W. Kyhos.
Erigeron quercifolius Lam. n = 9. South Carolina, Berkeley Co., near Monck’s Cor- near. R. Ornduff 6415, UC.
E. inflatum T. & F. var. inflatum.n = 16. Clark Co., near Las Vegas. A. Cronquist 9919, NY, UTC. Counted by J. L. Reveal.
E. maculatum Heller. n = 20. California, Mono Co., near Old Benton. J. L. Reveal & J. L. Reveal 551, UTC. Nevada, Clark Co., near Searchlight. A. Cronquist 9974, NY, UTC; Humboldt Co., Pine Forest Range, A. H. Holmgren & J. L. Reveal 1325, NY, UTC. Counted by J. L. Reveal.
E. nudum Dougl. ex Benth. var. pubiflorum Benth. n = 20. California, Modoc Co., Warner Range. A. H. Holmgren & J. L. Reveal 1717, NY, UTC. Counted by J. L. Reveal.
E. ovalifolium Nutt. var. nivale (Canby) Jones. n= 20. California, Mono Co., Sonora Pass. J. L. Reveal & J. L. Reveal 492, UTC. Nevada, Nye Co., Toquima Range, J. L. Reveal 629, NY, UTC. Counted by J. L. Reveal.
E. ovalifolium Nutt. var. ochroleucum (Small) Peck. n = 20. Nevada, Humboldt Co., Pine Forest Range. A. H. Holmgren & J. L. Reveal 1263, NY, UTC. Counted by J. L. Reveal.
E. ovalifolium Nutt. var. orthocaulon (Small) Hitchc. n = 20. Idaho, Butte Co., near Big Lost River. A. H. Holmgren & J. L. Reveal 825, NY, UTC. Nevada, Humboldt Co., Pine Forest Range. A. H. Holmgren & J. L. Reveal 896, NY, UTC. Counted by J. L. Reveal.
E. pusillum T. & G.n = 16. California, San Bernardino Co., near Needles. A. Cron- quist 9912, NY, UTC. Counted by J. L. Reveal.
E. reniforme T. & F.n = 16. Arizona, Mohave Co., near Kingman. A. Cronquist 9945, NY, UTC. Counted by J. L. Reveal.
E. sphaerocephalum Dougl. ex Benth. var. halimioides (Gand.) Stokes. n = 20. Nevada, Humboldt Co., Pine Forest Range. A. H. Holmgren & J. L. Reveal 1237, NY, UTC. Counted by J. L. Reveal.
E. trichopes Torr. ssp. minor (Benth.) Stokes. n = 16. Nevada, Nye Co., Mercury. A. Cronquist 9917, NY, UTC. Counted by J. L. Reveal.
Eriogonum umbellatum Torr. ssp. polyanthum (Benth.) Stokes. n = 40. Nevada, Humboldt Co., Pine Forest Range. A. H. Holmgren & J. L. Reveal 1225, NY, UTC. Counted by J. L. Reveal.
1965] DOCUMENTED CHROMOSOME NUMBERS 125
Eupatorium album L. n = 10. South Carolina, Richland Co. E. B. Smith 504, KANU.
Fritillaria atropurpurea Nutt. n = 12. Utah, Cache Co., Green Canyon. J. L. Reveal & P. J. Bottino 711, UTC. Counted by P. J. Bottino.
Helenium decurrens (Macbr.) Moldenke. n = 20. South Carolina, Jasper Co. E. B. Smith 523, KANU.
Helianthus angustifolius L. n= 17. Louisiana, Vernon Parish. FE. B. Smith 545, KANU.
Heterotheca subaxillaris (Lam.) Britt. & Rusby. n=9Q. Florida, Okeechobee Co. E. B. Smith 531, KANU.
Hierochloe occidentalis Buckl. 2n = 21n. California, Humboldt Co., near Arcata. D. E. Anderson 2925, UC.
Hilaria jamesii (Torr.) Benth. 2n = 18;;. Colorado, Mesa Co., Colorado National Monument. D. E. Anderson 7288, UC.
Houstonia caerulea L. n = 16. North Carolina, Durham Co., Duke University. W. R. Anderson s.n., UC. Counted by R. Ornduff.
H. serpyllifolia Michx. n = 8. South Carolina, Oconee Co., Camp Jocassee. R. Orn- duff 6382, UC.
Illictum floridanum Ellis. n = 13. Louisiana, Washington Parish. D. E. Stone 1357, NO. 2n = 26. Mississippi, Hancock Co. D. E. Stone 1522, DUKE.
I. parviflorum Michx. ex Vent. n= 14. Florida, Gainesville. D. E. Stone 1422, DUKE. Cultivated.
Marshallia graminifolia (Walt.) Small. n = 9. South Carolina, Georgetown Co. E. B. Smith 516, KANU.
Meconopsis betonicifolia Franch. n = 40. California, Mendocino Co., Fort Bragg. W. R. Ernst 738, DS. This is “var. bazley2”? with pubescent fruits. Cultivated.
M. cambrica (L.) Vig. n = 28. California, Berkeley, Univ. Calif. W. R. Ernst 730, DS. Cultivated
M. ? paniculata (D. Don) Prain. n = 28. California, San Francisco Co., Strybing Arboretum. W. R. Ernst 734, DS. Corollas yellow. Cultivated.
Melianthus major. 2n = 18. California, Humboldt Co., Arcata. D. E. Anderson 2917, UC. Cultivated.
Orthocarpus castillejoides Benth. var. humboldtiensis Keck. 2n = 121. California, Humboldt Co., salt marshes of Humboldt Bay. D. E. Anderson 2973, UC.
Palafoxia integrifolia T. & G.n = 10. Texas, Fayette Co. E. B. Smith 552, KANU.
P. sphacelata (Nutt.) Cory. n = 12. Texas, Bastrop Co. E. B. Smith 555, KANU.
Papaver alpinum L.n = 7. California, Santa Clara Co., Stanford Univ. W. R. Ernst 551, DS. Pubescent and glabrous plants studied. Cultivated.
P. atlanticum Ball. n = 7. California, Santa Clara Co., Stanford Univ. W. R. Ernst 553, DS. Some cells with a tetravelant figure. Cultivated.
P. bracteatum Lindl. n = 21. California, Santa Clara Co., Stanford Univ. W. R. Ernst 729B, DS. Most flowers subtended by a pair of large foliose bracts. Culti- vated.
P. nudicaule L. n= 14. California, Santa Clara Co., Stanford Univ. W. R. Ernst 552, DS. Common Iceland Poppy of nursery trade. Cultivated.
P. rhoeas L. n = 7. California, Santa Clara Co., Stanford Univ. W. R. Ernst 599, DS. Common single and double forms of Shirley poppy. Cultivated.
P. somniferum L. n = 11. California, Santa Clara Co., Stanford Univ. W. R. Ernst 566A, DS. Common opium poppy, occasionally with a few hairs on peduncles and petioles. Cultivated; W. R. Ernst 566B, DS. Common multiple-petaled or double opium poppy. Cultivated.
Plantago erecta Morris. 2n = 10;. California, Ventura Co., Santa Monica Mts. P. H. Raven 14866, LA. 2n = 20. California, Humboldt Co., near Samoa. D. E. Breedlove 3046 (progeny), DAV; Los Angeles Co., Santa Catalina I. P. H. Raven (progeny), RSA; Santa Catalina I., Middle Ranch, P. H. Raven 17841 (progeny), DAV; San Clemente I. P. H. Raven 17227 (progeny), DAV, RSA. (P. erecta ssp.
126 MADRONO [Vol. 18
vigidior Pilger). 2n = 42. California, Santa Barbara Co., near Lompoc, P. H. Raven 15511 (progeny), DAV. No evident correlation was found between chromo- some number and either morphology or pollen size in these collections, which appear to span the range of variation in this species. All collections studied were autogamous, in contrast to the allogamous but similar P. znsularis Eastw. Counted by D. M. Moore.
P. insularis Eastw. 2n = 8. California, San Bernardino Co., near Barstow. D. M. Moore 422, DAV.
Pleuropogon refractus (Gray) Benth. ex Vasey. 2n = 18. California, Humboldt Co., near Arcata. D. E. Anderson 2404b, UC.
Porophyllum tridentatum Wats. var. crassifolium Johnston. n = 15. Mexico, Baja California, San Bruno. A. Carter & J. Reese 4538, UC, KANU. Counted by R. R. Johnson.
Primula maguirez Williams n = 22. Utah, Cache Co., Logan Canyon. J. L. Re- veal © P. J. Botino 710, UTC:
Schisandra glabra (Brickell) Rehder. 2n = 28. Florida, Liberty Co., D. Demaree, April 6, 1964, DUKE. Counted by D. E. Stone.
Sidalcea malachroides (H. & A.) Gray. 2n = 10;. California, Humboldt Co., near Arcata. D. E. Anderson 2502, UC.
Solanum brachystachys Mart. & Gal. 2n = 121. Mexico, Chiapas, Municipio of Chamula. D. E. Breedlove 7060, DS. Counted by D. W. Kyhos.
Tolmiea menziesti (Pursh) T. & G. 2n = 147. California, Humboldt Co., near Ar- cata. D. E. Anderson & B.D. Rogers 1564, UC. Counted by B. D. Rogers.
REVIEWS
A Selected Guide to the Literature on the Flowering Plants of Mexico. By Ipa Kaptan LANGMAN. 1015 pp. University of Pennsylvania Press, Philadelphia, 1964. $25.00.
The publication of this annotated bibliography culminates more than 20 years of dedicated labor. The breadth of Mrs. Langman’s research on the literature of Mexi- can botany is indicated to some extent by the journals and the bibliographic titles cited in the preliminary materials and by the list of libraries which she consulted. In order to obtain references to local material that might not have found its way to the larger libraries, Mrs. Langman visited libraries in 27 states of Mexico, so that here we have an invaluable tool for all having an interest in any aspect of Mexican botany from the time of the Conquistadores to the present. The viscissitudes of publishing evidently made impossible the last minute inclusions of references to the most important taxonomic contribution to our knowledge of the flora of northwest- ern Mexico, I. L. Wiggins, Flora of the Sonoran Desert (Stanford University Press, 1964).
Of the 1015 pages, 792 are devoted to an alphabetical compilation of author cita- tions. The user should refer to Mrs. Langman’s remarks (p. 12) regarding the diffi- culties encountered in citing authors’ names, especially the Latin American “tri- nomials.”’
The 156 pages of Index (four columns to a page) immeasurably increase the value of the Bibliography, for the author’s labor has forestalled ours. Familiarity with its organization will enable knowledgeable researchers to by-pass several of the reference steps. Otherwise, a step-by-step search for information about j:cama would lead the reader from jicama to the genera Cacara, Dolichos, and Pachyrrhi- zus, each of which in turn refers him to “Leguminosae.” Under this family the genera are listed in alphabetical order. Under the genera are various subject head- ings, and, finally, the desired author references to the main part of the Bibliography, q.v. A typographically understandable but unfortunate structural difficulty in the Index is the minimal indentation of the first subheadings. This, coupled with the fact that the main heading (when a subject is carried from column to column) is
1965 | NOTES AND NEWS 127
included only on the first of the four columns of each page, sometimes makes it difficult to find the desired entry in the Index. This problem is amply illustrated by perusal of the several pages of references to the Cactaceae.
Ida Langman, the institutions and grantors which supported her work, and the University of Pennsylvania Press deserve our thanks and congratulations for bring- ing this work to fruition. It is a “must” for all libraries having any interest in Mexico, as well as for all people concerned with any aspect of the plants of the area. Anyone using the volume should first turn to p. 9 and lighten his day by reading “Random Thoughts on Bibliographies.”—ANNETTA CARTER, Department of Botany, University of California, Berkeley.
The Vascular Plants of Monterey County, California. By BEATRICE F. Howitt and JoHN THomMaAs Howe tr. The Wasmann Journal of Biology 22(1) :1i+1—184. 1964. Published by the University of San Francisco. Available from the California Academy of Sciences, Golden Gate Park, San Francisco, California 94118. $2.75.
This work is a welcome addition to the floras and annotated checklists of the vascular plants of central California. Monterey County, the area covered by this annotated catalogue, is about 3324 square miles in extent and is located along the California coast south of San Francisco. The vascular flora is made up of 119 families, 539 genera, 1713 specific and infraspecific categories. Of these, 291 are in- troduced and 37 are endemic. Some 302 species have either their northern or south- ern limits of distribution in the Coast Ranges in Monterey County. For the Santa Cruz Mountains immediately to the north, the corresponding figure is 242 and for Marin County to the north of San Francisco the figure is 123. Monterey County then would appear to have special significance in the phytogeography of the Cali- fornia Coast Ranges.
The County has been divided into eight major sections based on terrain, and these are discussed, outlined on a map, and the conspicuous plants in them listed. These sections and specific localities within them are used in the catalogue to indi- cate distribution within the County. There is no discussion of the plant communities, characteristic assemblages of plants, as such. The short section by Oliver E. Bowen outlines the main features of the geology. The arrangement of families follows that of A California Flora; common names are listed; specific names are capitalized; and neither keys nor descriptions are included. There are 22 text figures and a frontispiece which depict various vegetation types. Undue emphasis has been given to type localities. Information of this sort for each species adds little to local floristic works except length. More often than not, the geographical location of a type locality is a matter of historical accident rather than of biological significance. It does not follow from the fact that one fifth of the plants in Monterey County have their type locality there that it is (p. 24) “. .. one of the richest and most important research areas for field studies in systematic botany in western America.”
Hopefully this work will stimulate local groups and individuals to guard and preserve the diversity of species in this area, an area known to many throughout the world because of the Monterey Cypress and the Monterey Pine——Jonn H. Tuomas, Dudley Herbarium, Stanford University.
NOTES AND NEWS
LABELS FOR HERBARIUM SPECIMENS.—With relatively few exceptions labels ac- companying herbarium specimens are unsatisfactory. Many include only the sketchi- est data as to locality; most indicate nothing about habitat, soil type, slope, expo- sure, associated plants, flower color, etc.; and many are of a very poor quality paper. Often this is understandable. It takes time and/or money to have adequate labels typed or printed. Advances in the art of photo-offset printing have, however, made it possible to produce labels at a reasonable cost, less in fact than it costs to have them typed. At Stanford University, we now routinely have labels made by the offset method even when only 20-25 copies of any particular one are needed.
128 MADRONO [Vol. 18
Sample labels are typed, and six of them pasted on a standard size sheet of typing paper. These are then photographed by the University photographic service with an Itek camera on an Itek photodirect master. Up to 1000 copies can be made from one master. Should more be required, a metal offset plate can be made with the capacity of producing over 5000 copies. Any kind of paper, including 100% rag, and even 90 lb. card stock, can be used with either the photodirect master or the metal offset plate. The cost of the photodirect master (in our experience) is $1.00. A press run of 100 copies is $1.10: that for 1000 copies is down to $4.20. A good quality, 100% rag paper is about $7.50 per ream. Assuming that one needs 100 copies of each of six different labels, the cost per individual label is slightly over one-half a cent. The advantages of this method are obvious—JoHn H. Tuomas, Dudley Herbarium, Stanford University.
THE HERBARIUM OF Howarp E. McMinn.—During his teaching and research career from 1918 to 1957, the late Professor Howard E. McMinn built up, in the Biology Department of Mills College, a significant herbarium of native woody plants of California as well as collections of introduced woody ornamentals and allergenic plants of California. A few months ago, in order to facilitate the avail- ability of McMinn’s voucher specimens to interested botanists, his native plant collections were transferred to the Herbarium of the University of California at Berkeley. These specimens (approximately 4000) served as the basis for McMinn’s following important books and monographs: H. E. McMinn and E. Maino, 1937, An Illustrated Manual of Pacific Coast Trees, Univ. Calif. Press, Berkeley; H. E. McMinn, 1939, An Illustrated Manual of California Shrubs, Univ. of Calif. Press; M. van Rensselaer and H. E. McMinn, 1942, Ceanothus, Santa Barbara Botanic Garden, Santa Barbara, Calif.; H. E. McMinn, 1951, Studies in the Genus Diplacus, Scrophulariaceae, Madrono 11:33-128. McMinn’s specimens will be undergoing processing at the University of California shortly and should be available for study by the end of 1966. Thereafter, requests for borrowing them should be addressed directly to the Director of the University of California Herbarium, University of California, Berkeley, California.
McMinn’s collections of “Introduced Woody Plants of California” and “Wind Pollinated Plants of California” are kept in the Biology Department of Mills College as reference and teaching material in Botany classes. This portion of McMinn’s herbarium consists of approximately 2000 specimens of ornamental plants and over 1000 allergenic plants. Most of the voucher specimens that have been utilized in the publication of the following two books are maintained in this herbarium: H. E. McMinn, 1919, A Manual of Trees, Shrubs and Vines of Mills College Campus, H. S. Crocker Co., Inc., San Francisco and Oakland; E. Graham and H. E. McMinn, 1941, Ornamental Shrubs and Woody Vines of the Pacific Coast, Gillick Press, Berkeley, Calif. These specimens are filed according to generic names among which Acacia, Acer, Berberis, Callistemon, Cotoneaster, Erica, Escal- lonia, Eucalyptus, Fraxinus, Ilex, Leptospermum, Melaleuca, Pittosporum, Pyra- cantha and Tilia are particularly well-represented. Previous arrangement is neces- sary for those interested in studying these specimens——BAkI Kasaptiicit, Depart- ment of Biology, Mills College, Oakland, Calif.
The following publications are of interest:
The Genus Lithophragma. By Roy L. Taytor. University of California Publica- tions in Botany 37:1—122. University of California Press, Berkeley and Los Angeles. 1965. $3.00.
Orchids, Care and Growth. By Micuet Pau. Universe Books Inc., 381 Park Ave. S., New York 10016. 135 pp. 1965. Paper, $1.95; hardcover, $2.75.
Drawings of British Plants. By STELLA Ross-Craic. Part XVIII. Boraginaceae, Convolvulaceae, Solanaceae. 32 plates. G. Bell & Sons, London. 1965. 10/6d.
Illustrated Dictionary of Botanical Terms (excerpt from). By JoHNn LINDLEY. With an introduction by AtickE EAstwoop and a foreword by JoseEPH J. GRAHAM. 48 pp. School of Earth Sciences, Stanford University. 1964.
sae
ADRONO
VOLUME 18, NUMBER 5 JANUARY, 1966
Contents
A TaxONOMIC REVISION OF SECTION SONORICOLA OF THE GENUS VERBESINA L. (COMPOSITAE), James R. Coleman 129
ESCOBARIA LEEI BODEKER REDISCOVERED IN New Mexico, Edward F. Castetter and Prince Pierce 137
New ReEcorps OF MYXOMYCETES FROM CALIFORNIA I., Donald T. Kowalski 140
CONTRIBUTION TO THE EMBRYOLOGY OF MUILLA, WITH A REMARK ON THE TAXONOMIC POSITION OF THE GENusS, Rolf Y. Berg and Jack R. Maze 143
THE GENUS TIARELLA IN WESTERN NorTH America, Patricia Kern e152
NoTES AND NEws: OCCURENCE OF THE GENUS EUPHRASIA IN THE PactFic NORTHWEST, Fred R. Ganders ; HERBARIUM SUPPLIES 160
A WEST AMERICAN JOURNAL OF BOTANY
UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
Second-class postage paid at Berkeley, California. Return requested. Established 1916. Subscription price $6.00 per year ($4.00 for students). Published quarterly in January, April, July, and October by the California Botanical Society, Inc., and issued from the office of Madrofo, Herbarium ,Life Sciences Building, University of California, Berkeley, California. Orders for subscriptions, changes in address, and un- delivered copies should be sent to the Corresponding Secretary, California Botanical Society, Department of Botany, University of California, Berkeley, California 94720.
BOARD OF EDITORS
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LyMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis HERBERT F. CopELAND, Sacramento College, Sacramento, California JoHN F. Davipson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California MILpreD E. MATHIAS, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley Marion OwnBEY, Washington State University, Pullman REED C. RoLiins, Gray Harbarium, Harvard University IrA L. Wiccins, Stanford University, Stanford, California
Editor—JoHN H. THomMAS Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Malcolm A. Nobs, Carnegie Institution of Washington, Stanford. First Vice-President: W. B. Critchfield, Pacific Southwest Forest and Range Experiment Station, Forest Service, U. S. Department of Agriculture, Berkeley. Second Vice- President; William S. Boyle, Department of Botany, Utah State University, Logan. Recording Secretary: Mary L. Bowerman, Department of Botany, University of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Department of Bot- any, University of California, Berkeley. Treasurer: Douglas M. Post, Biology De- partment, San Francisco State College.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Annetta Carter, Department of Botany, University of California, Berkeley ; the Editor of Madrono, and three elected Council Members: William M. Hiesey, Carnegie Institution of Washington, Stanford; Herbert L. Ma- son, Department of Botany, University of California, Berkeley; and Virginia M. Page, Department of Biological Sciences, Stanford University.
A TAXONOMIC REVISION OF SECTION SONORICOLA OF THE GENUS VERBESINA L. (COMPOSITAE)
JAMES R. COLEMAN
Verbesina is the largest genus in the subtribe Verbesineae of the tribe Heliantheae, the number of species being about 250. The genus is en- tirely new world with the species occurring from the United States to Argentina but being best developed in the tropics.
This paper is part of a Ph. D. thesis submitted to Indiana University. The work was supported by fellowships from the National Science Foundation and Indiana University. I wish to express gratitude to Charles B. Heiser for suggesting this investigation and to Annetta Carter for generously supplying valuable materials and information. My current address is: Institute de Botanica, Sao Paulo, Brazil.
The genus was last treated in its entirety by Robinson and Greenman (1899) in which treatment they recognized 109 species in 12 sections. During this century one section, Lipactinia, was revised by Blake (1925) and recently three others, Pterophyton, Sonoricola and Ximenesia, were revised by Coleman (1964). This paper is based on the revision of one of these sections, Sonoricola.
Section Sonoricola is a natural group of Baja California and northern mainland Mexico, being best developed in Baja California where seven of its nine species occur. Endemism appears to be the rule in this section with only one species, V. chihuahuensis Gray, occupying any consider- able range. Additional collecting in Baja California may show that this endemism more apparent than real. The species occur in regions of coastal sand dunes, semi-desert plateaus and, more commonly, moun- tains. This includes altitudes of from near sea level to approximately 10000 feet.
Robinson and Greenman (1899) treated section Sonoricola as com- posed of six species. It is here expanded to nine species.
Section Sonoricola is most closely related to sections Pterophyton and Ximenesia and is separable from them as shown in the following key:
Erect perennials; stems winged or wingless; petiole bases mostly exauriculate; rays yellow, rarely lacking; chaff not particularly slender; awns, if present, not filiform.
Herbaceous; stems winged or wingless; ray achenes, if present, not tuberculate; awns lacking or quite short, rarely to 3 mm long. .....................- Sec. Pterophyton Fruticose or suffruticose; stems wingless; ray achenes, if present, tuberculate; awns mostly 3-7 mm long, rarely JeSS. ..............0....-:ee2ecceeeeeeeeeeeeeeeees Sec. Sonoricola Annuals or prostrate perennials; stems wingless; petiole bases mostly conspicuously auriculate; rays orange or yellow; chaff quite slender, even filiform; awns fili- Onan OCCASIONS ly) LACKING: ooccz cc. cceecececcseeeceotecdccclavedet cae dedectecassctbeectease Sec. Ximenesia
MaproNo, Vol. 18, No. 5, pp. 129-160. February 16, 1966.
130 MADRONO [Vol. 18
Unfortunately the amount of material available for crossing and cyto- logical studies was quite limited. However chromosome counts were ob- tained for two species, V. peninsularis Blake and V. oligocephala John- ston (table 1). The counts were made by the aceto-carmine squash method on buds which had previously been fixed in a solution of 95% ethyl alcohol, chloroform and proprionic acid (6:2:2). Meiosis of V. peninsularis was normal with 17 pairs being observed. Of the two plants of V. oligocephala examined, however, one showed 17 pairs plus one B chromosome whereas the second showed 17 pairs plus two B chrom- osomes. This is the first report of B chromosomes in Verbesina. These two species agree with the majority of the species of Verbesina counted thus far in having a haploid number of 17. However, counts of n = 16 and 18 have been reported (De Jong and Longpre, 1963; Turner, Elli- son, and King, 1961; Turner, Beaman, and Rock, 1961) and tetraploid species are known (Heiser and Smith, 1955; Coleman, 1964). The counts presented in this paper are the first reports for section Sonoricola.
The section has no economic importance.
TABLE 1. CHROMOSOME NUMBERS
V. peninsularis Blake. n= 17. Mexico, Baja California, near Rancho San Matias. A. Carter 4619 (IND, UC). Counted by J. R. Coleman.
V. oligocephala Johnston. n = 17 + 1b and 17 + 2b. Mexico, Baja California, near Puerto Escondido. J. R. Coleman V6233 (IND). Progeny of A. Carter 4356 (UC).
Section SonorIcoLaA Robinson and Greenman, Proc. Am. Acad. 16:542. 1899. Type species, Verbesina dissita Gray.
Fruticose or suffruticose; stems wingless; leaves all opposite or the uppermost alternate, petiolate (except V. dissita), the petioles narrowly to broadly winged and slightly decurrent or, more commonly, not so, quite lustrous to not at all so, mostly deltoid-ovate to lanceolate, spar- ingly to quite pubescent; phyllaries ca. 3—4—-seriate, most narrowly to broadly oblong, at times spatulate; heads medium to large (ca. 1.7—4.5 cm wide); rays yellow, neutral or styliferous, sterile or fertile; disc florets yellow; ray achenes triangular in cross section, mostly awnless; disc ahenes provided with well-developed lateral wings and 2 long (ca. 3.0-7.0 mm) awns.
KEY TO THE SPECIES
Leaves sessile; disc achenes ca. 7—9 mm long and essentially glabrous........ 1. V. dissita Leaves petiolate; disc achenes less than 7 mm long or densely hirsute. Leaves quite lustrous; petioles often broadly winged, some usually slightly de- current. Older stems waxy white, younger stems mostly white tomentoes; inflorescences clustered amongst: terminal, follidces 22a see 2. V. oligocephala Stems not white; inflorescences borne well above terminal foliage. Leaves quite scabrous; ray florets neutral.....................----.------se-200+ 3. V. palmert Leaves hispid: ray florets: styliterous:4224.0 2a t ee eeceeee se 4. V. hastata
1966] COLEMAN: VERBESINA 131
Leaves not particularly lustrous; petioles mostly narrowly winged and _ never decurrent. Outer phyllaries lax; species of northern mainland Mexico. Principal leaves often over 10 cm long; ray florets sterile; achenes oblong. 5. V. leptochaeta Principal leaves rarely over 10 cm long; ray florets fertile; achenes cuneate. 6. V. chihuahuensis Outer phyllaries erect; species of Baja California. Leaves not densely pubescent, mostly quite membranaceous. 7. V. peninsularis Leaves densely hispid-scabrous. Leaves coarsely serrate; ray flowers awnless..............00.....000...00--.--- 8. V. erosa Leaves finely serrate; ray flowers awned...................0.....-20-.------ 9. V. pustulata
1. VERBESINA DISSITA Gray, Proc. Am. Acad. 20:299. 1885. Type, near All Saints Bay, Lower California, Orcutt s.n., Sept. 1884, GH.
Suffruticose, ca. 1 m tall; stems sparingly hispidulous near the in- florescences, otherwise glabrous; leaves mostly opposite, becoming alter- nate above, sessile, occasionally clasping the stem, mostly ovate, serrate to entire, acute or obtuse, to ca. 11 cm long, mostly 2—5 cm wide, hispid to smoothish, lustrous; inflorescences 3-16—headed, cymose-paniculate, pedicels ca. 2-10 cm long and stout, heads mostly 4.0-4.5 cm wide, the discs mostly 1.5—2.0 cm wide; phyllaries ca. 3-seriate, mostly broadly oblong, at times spatulate, the outer series 5—8 mm long, the inner series ca. 8-12 mm long, mostly acute, hirsute, black specked; ray florets ca. 10, neutral, ca. 2.0—2.5 cm long, the ligule oblong, the tube papillate, the ovary ca. 2.5-3.2 mm long, glabrous, awnless; disc florets mostly 13-14 mm long, the tube papillate or hirsute-papillate, the ovary ca. 5 mm long, sparingly hirtellous, the awns ca. 3.2—-3.6 mm long, the style tips acute; receptacle conical; chaff mostly 10-13 mm tall, hirtellous, acute; disc achenes obovate, mostly 8-9 mm long, 5.5—7.0 mm wide, essentially glabrous, the wings to 3 mm wide, the margins often erose or wavy, the awns mostly 3.2—4.2 mm long, erect or bent inward so as to cross each other.
Distribution. Sand dunes, coastal regions of extreme southern Cali- fornia and northern Baja California.
Specimens examined: CALIFORNIA. Orange Co.: near Arch Beach, Bradshaw s. n., May 1903 (UC). BAJA CALIFORNIA. La Grulla, Orcutt 355 (UC); Sanzal, Orcutt s. n., May 21, 1886 (F, NY, UC); near Descanso Point, Wiggins 11904B (TEX, UC); Todos Santos., Brandegee s. n., May 10, 1897 (UC); Todos Santos Bay, Orcutt 1233 (UC).
Verbesina dissita appears somewhat intermediate between sections Sonoricola and Pterophyton. It possesses the long pappus awns char- acteristic of section Sonoricola and is geographically associated with that section. However the sessile leaves, large heads and the achenes (except the awns) are somewhat suggestive of V. lindheimeri Rob. & Greenman of section Pterophyton.
In this treatment V. dissita is considered closest to V. oligocephala, V. palmeri and V. hastata, but is rather distinct from each by virtue of its
132 MADRONO [Vol. 18
sessile leaves and glabrous achenes. It shares lustrous foliage with each, neutral ray florets with the first two species and subglabrous or merely hispid foliage with the latter.
2. VERBESINA OLIGOCEPHALA. Johnston, Proc. Calif. Acad. 12:1200. 1924. Type, on a rocky slope in the mountains back of Aqua Verde Bay, Lower California, Johnston 3899, UC.
Spreading shrub ca. 1.5—2.0 m tall; older stems waxy white with bases of old hairs, younger stems often white tomentose; leaves opposite or the uppermost alternate, petioles ca. 1.5—3.0 cm long, cuneately winged, often auriculate, clasping the stem and decurrent for ca. 4-6 cm, mostly ovate but to lance-ovate or lanceolate, coarsely crenate-dentate to sub- entire, acute or obtuse, rarely subacuminate, mostly 6—14 cm long, 2.0- 7.5 cm wide, scabrous, lustrous; inflorescences corymbose, mostly 2-8 headed, scarcely or not at all extending beyond the upper foliage, heads ca. 2-3 cm wide, discs ca. 1.8—2.2 cm wide; phyllaries ca. 3-seriate, ovate to oblong, the distal half often greatly expanded, subequal, mostly 6-8 mm long, mostly obtuse, the margins ciliate; ray florets 8-12, neutral, ca. 1.3-2.0 cm long, the ligule oblong or elliptic-oblong with the tube and base hirsute, the ovary ca. 2.0-2.5 mm long, papillate, awnless; disc florets ca. 10-12 mm long, the lower half of the corolla hirsute, the ovary ca. 3.5-4.5 mm long, densely upwardly hirsute, the awns ca. 5-7 mm long, the style tips acute; receptacle low convex; chaff mostly 9-12 mm tall, acute, the margins ciliate, subscarious; disc achenes cuneate, ca. 7 mm long, 4-5 mm wide, densely upwardly hirsute, the wings to ca. 1.5 mm wide, the margins ciliate, the awns ca. 5—7 mm long, slightly un- equal, mostly spreading.
Distribution. Northern sector of the Sierra de La Giganta, Baja Cali- fornia.
Specimens examined. BAJA CALIFORNIA. Near Puerto Escondido, Carter 4356 (UC) ; Carter & Kellogg 2855 (UC) ; Carter 4345 (UC); near Loreto, Carter & Kel-
logg 2834 (UC) Carter 3705 (UC) ; Arroyo Gabilan, Carter & Ferris 4079 (UC) ; near San Javier, Carter & Ferris 3807 (UC).
Verbesina oligocephala has its closest affinities with V. palmeri, differ- ing most strikingly from that species by its whitish stems and short inflorescence.
3. VERBESINA PALMERI Wats., Proc. Am. Acad. 24:56. 1889. Type, in canons above Los Angeles Bay, Palmer 528, UC!, isotype, NY!
Suffruticose, ca. 1.0-1.3 m tall; stems hispid-scabrous; leaves mostly opposite, alternate above, petioles to ca. 2 cm long, winged (wings 3-16 mm wide), often subauriculate, clasping the stem and decurrent for up to ca. 1 cm, deltoid, ovate or lance-ovate, repandly serrate-dentate to entire, acute or acuminate, to ca. 8.5 cm long, mostly ca. 5 cm long, to ca. 6.5 cm wide, commonly 2.5—4.0 cm wide, lustrous, scabrous; inflores- cences corymbose, ca. 5—7-headed, pedicels mostly 2—5 cm long, heads ca. 2.5 cm wide, discs ca 8-10 mm wide; phyllaries ca. 3—4-seriate, the
1966] COLEMAN: VERBESINA 133
outer series obovate to oblong, 2.5—-5.0 mm long, obtuse, the inner series oblong to lance oblong, 6-8 mm long, acute; ray florets ca. 10, stylifer- ous, sterile, ca. 1.0-1.5 cm long, the ligule oblong to elliptic-oblong, the tube hirtellous, the ovary ca. 1.8-2.3 mm long, awnless; disc florets mostly 9-12 mm long, the tube hirtellous, the ovary 3-5 mm long, densely upwardly hirsute, the awns ca. 4-5 mm long, the style tips acute; receptacle low convex; chaff mostly 9-10 mm tall, the margins ciliate, the tips acute or acuminate, some recurved; disc achenes obovate, ca. 6-7 mm long, 4-5 mm wide, upwardly hirsute, the wings to ca. 1.3 mm wide, the margins ciliate, the awns ca. 4-5 mm long, spreading to vary- ing degrees. Distribution. Northeastern Baja California.
Specimens examined. BAJA CALIFORNIA. Canyon del Diablo, E bank of Sierra San Pedro Martir, Chambers 563 (UC).
Verbesina palmeri appears quite close to V. hastata, differing from that species by possessing scabrous leaves, chaff yellowish throughout and pappus awns ca. 4-5 mm long. Verbesina hastata has smooth or merely hispid leaves, chaff often black apically and pappus awns 6-7 mm long.
4. VERBESINA HASTATA (Kellogg) Curran, Bull. Calif. Acad. 1:140. 1885. Liptochaeta hastata Kellogg, Proc. Cal. Acad. 2:106, fig. 31. 1861. Type, illustrated cited! V. venosa Greene, Bull. Torrey Club 9:110. 1882. Type, Cedros I., Veatch?, UC. Encelia cedrocensis Rose, Contr. U.S. Natl. Herb. 1:17. 1890. Type, Cedros I., Palmer 741, UC!
Shrub to ca. 1 m tall; stems subglabrous; leaves all opposite or the upper alternate, petioles to ca. 1.5 cm long, winged (the wings to ca. 5 mm wide), often clasping the stem and decurrent for ca. 3-5 cm, mostly ovate but to lanceolate, repandly dentate to entire, acute or acuminate, rarely obtuse, mostly 5-7 cm long, 2—4 cm wide, subglabrous to hispid, lustrous; inflorescences loosely corymbose, to ca. 10-headed, the pedicels mostly under 5 cm long, the heads ca. 2.0—2.5 cm wide, the discs ca. 1.0— 1.5 cm wide; phyllaries 3—5—-seriate, the outer series obovate or spatu- late, mostly 3-5 mm long, obtuse, the inner series lanceolate, ca. 6.7—7.5 mm long, acute; ray florets ca. 12, styliferous, sterile, ca. 1 cm long, the ligules broadly oblong, the tube hirsute, the ovary 1.5—2.2 mm long, awnless; disc florets mostly 9-12 mm long, the tube hirsute, the ovary ca. 3.0-4.2 mm long, densely upwardly hirsute, the awns to ca. 6.5 mm long, the style tips acute; receptacle low convex; chaff ca. 6.5-8.5 mm tall, sparingly hirtellous, the apical portion black, acute; disc achenes obovate, ca. 5—6 mm long, 2.8-4.2 mm wide, the margins ciliate, the awns to ca. 7 mm long, mostly slightly unequal, erect or somewhat spreading.
Distribution: Cedros I., Baja California.
Specimens examined. BAJA CALIFORNIA. Cedros I., Rose 16117 (NY); Anthony 63 (NY, UC); Anthony 296 (F, UC).
See discussion under V. palmeri for relationship to that species.
134 MADRONO [Vol. 18
5. VERBESINA LEPTOCHAETA Gray, Proc. Am. Acad. 21:389. 1886. Type, near Batopilas, Chihuahua, Palmer 170, GH!
Bushy shrub ca. 1.5—-2.5 m tall; stems mostly glabrous, hispid near the heads; leaves opposite, petioles to ca. 4 cm long, very narrowly winged, ovate-deltoid to broadly elliptical, serrate, acute, to ca. 13 cm long, to ca. 8 cm wide, scabrous; heads subsolitary, discs ca. 2 cm wide; phyllaries 2—3-seriate, narrowly oblong, subequal, ca. 7 mm long, acute, the outer series lax herbaceous; ray florets styliferous, sterile, the ligule linear, the ovary ca. 5 mm long, sparingly hirsute; chaff ca. 10 mm long, subacute; disc achenes oblong, ca. 6-7 mm long, 2.5—3.0 wide, hirsute, the wings ca. 0.5-1.0 mm wide, the margins ciliate, the awns subequal, ca. 4-5 mm long, suberect.
Verbesima leptochaeta is known only from its type specimen which is unfortunately in rather poor condition. The species fits well into sec- tion Sonoricola, but its relationships are obscure. Blake (1924) considers it to have an affinity with V. peninsularis, but certainly this relationship would not be a very close one. It resembles the only other mainland species of the section, V. chihuahuensis, in having lax, spreading outer phyllaries.
6. VERBESINA CHIHUAHUENSIS Gray, Proc. Am. Acad. 21:389. 1886. Type, Santa Eulalia Mountains, near Chihuahua, on limestone, in canons, Pringle 657, GH, isotypes, F!, NY! V. parrasana Brandegee, Univ. Calif. Publ. Bot. 4:191. 1911. Type, Sierra de Parras, Purpus 4635, UC!
Suffruticose, to ca. 0.75 m tall; stems puberulent to densely pilose, occasionally reddish; leaves opposite below, alternate above, the petiloes to ca. 4% the length of the leaf, narrowly winged, ovate-deltoid to lanceo- late, coarsely serrate or dentate to subentire, acute, rarely obtuse, to ca. 11 cm long, commonly 4—6 cm long, mostly 1.5—3.0 cm wide, hispid, at times densely pilose; inflorescences mostly 1—4-headed, the pedicels quite slender, mostly 5—6 cm long, the heads ca. 2.5—3.5 cm wide, the discs ca. 1.0—-1.5 cm wide; phyllaries ca. 3-seriate, linear-oblong to lanceolate, sub- equal, to ca. 15 mm long, mostly 5-10 mm long, mostly acute, the outer series lax; ray florets ca. 12-14, commonly styliferous and fertile, rarely neutral, mostly 1.8—2.4 cm long, the ligule linear to oblong, the tube hirsute, the ovary 2.5-3.5 mm long, subglabrous, awnless; disc florets mostly 7.5-10.0 mm long, the corolla hirsute throughout, the ovary ca. 3.0-4.5 mm long, densely upwardly hirsute, the awns slender, to ca 4—5 mm long, the style tips attenuate; receptacle conical; chaff 5.5-8.5 mm tall, sparingly hirtellous, subscarious, often black-specked, acute-attenu- ate; ray achenes ca. 3-4 mm long, awns usually lacking, occasionally to ca. 2.5 mm long; disc achenes cuneate, mostly 4—7 mm long, 2.5-3.5 mm wide, densely upwardly hirsute, the wings to ca. 1 mm wide, the awns equal, to ca. 4-5 mm long, erect.
Distribution. Northern regions of the Mexican Plateau.
1966] COLEMAN: VERBESINA L3o
Specimens examined. CHIHUAHUA. Santa Eulalia Plains, Wilkinson s. n., 1885 (F); mountains NW of Chihuahua, Le Sueur 1020 (F, TEX); near Piramide, Johnston & Muller 1421 (GH); near Chihuahua, Pringle 1058 (NY, UC). COA- HUILA. Near Saltillo, Palmer 407 (F, NY); Carneros Pass, Pringle 2782 (F); Sierra de la Paila, Purpus 4694 (UC); near Parras on Sierra Negras, Stanford, Reth- erford, & Northcraft 288 (NY). DURANGO. Durango, Palmer 322, (F, NY). ZA- CATECAS. Near Concepcion del Oro, Stanford, Retherford, & Northcraft 515 (UC).
Verbesina chihuahuensis shows a strong similarity to V. encelioides (Cav.) Benth. & Hook. of section Ximenesia. This similarity includes the ray florets which in both cases are styliferous and fertile, the achenes, both ray and disc, and the spreading outer phyllaries. The species con- trast most notably in regards to habit, V. chihuahuensis being perennial
and V. enceliodides being annual.
7, VERBESINA PENINSULARIES Blake, Proc. Biol. Soc. Wash. 37:58. 1924. Type, 32 km E of San Ignacio, Baja California, Wilson & Gold- man 7227, US. V. cayucensis Jones, Contr. West. Bot. 18:76. 1933. Type, Cayuca Ranch, Loreto, Baja California, Jones 27718, UC!
Suffruticose or fruticose, ca. 0.5—1.5 m tall; stems hispidulous-puberu- lent, occasionally reddish; leaves mostly opposite, alternate above, the petioles ca. 2 cm long, narrowly cuneately winged, ovate-deltoid to lanceolate, rather coarsely serrate, often doubly serrate, acute or acumi- nate, to ca. 13.5 cm long, mostly 2.5-5.0 cm wide, scabrous or sub- scabrous, membranaceous; inflorescences subsolitary or cymose, to ca. 6—headed, the heads mostly 1.7—2.5 cm wide, the discs ca. 10-12 mm wide; phyllaries subequal, mostly 5.5—7.5 mm long, acute or obtuse, the margins ciliate; ray florets ca. 10, styliferous, fertile, mostly 1.0-1.5 cm long, the ligule linear-oblong, the tube hirsute, the ovary ca. 2 mm long, glabrous, awnless; disc florets ca. 7-8 mm long, the tube hirsute, the ovary ca. 2-3 mm long, the style tips attenuate; receptacle low convex; chaff ca. 7-8 mm tall, subglabrous, obtuse or acute, scarious; ray achenes ca. 4 mm long, awnless; disc achenes obovate, ca. 6—7 mm long, ca. 4 mm wide, sparingly hirsute, the wings to ca. 3 mm wide, the mar- gins erose and sparingly ciliate, the awns subequal, mostly 2.5-3.0 mm long, erect.
Distribution. Northern region of the Sierra de La Giganta, Baja Calif.
Specimens examined. BAJA CALIFORNIA. Near Canipole, Wiggins 11455 (UC); Cerro de La Giganta, Carter & Kellogg 3120 (UC); Carter & Ferris 3989 (UC); near San Javier, Carter & Ferris 3851 (UC); Chisaki & Carter 130 (UC); Rancho San Matias, Carter 4619 (UC); La Esperanza, Carter 4373 (UC); near Llanos de San Juan, Carter 4499 (IND, UC).
Verbesina peninsularis differs from V. erosa to which it is mostly closely related by having membranaceous leaves which are not densely pubescent.
The rather meager distributional data available for this section indi- cates that sympatry is not common. A definite exception is however pre- sented by V. peninsularis and V. oligocephala; however, the specimens examined give no reason to suspect that hybridization is occurring.
136 MADRONO [Vol. 18
8. VERBESINA EROSA Brandegee, Proc. Calif. Acad. 3:146. 1891. Type, Sierra de San Francisquito, Brandegee 316, UC!
Suffruticose, ca. 1-2 m tall; stems hirsute-hispid, often reddish; leaves mostly opposite, the uppermost alternate, the petioles mostly 1-2 cm long, cuneately winged, mostly ovate to lance-ovate, coarsely serrate or doubly serrate to subentire, acute or acuminate, to ca. 15 cm long, mostly 3-6 cm wide, densely hispid-scabrous; inflorescences subcorym- bose, 3—14-headed, pedicels mostly 1-5 cm long, the heads mostly 2-3 cm wide, the discs ca. 8-10 mm wide; phyllaries 2—3-seriate, oblong or lanceolate, subequal, mostly 5-9 mm long, acute or subacute, margins of the inner series ciliate; ray florets ca. 8-10, styliferous, fertile, ca. 0.8—-1.3 mm long, the ligule oblong or elliptic-oblong, the tube and base densely hirsute, the ovary ca. 1.5—2.2 mm long, papillate, awnless; disc florets mostly 5-8 mm long, the tube densely hirsute, the ovary ca. 2.0- 2.5 mm long, hirsute, the awns 2—3 mm long, the style tips acute; recep- tacle conical; chaff ca. 4.5—-7.8 mm tall, hirsute-hirtellous, acute, the mar- gins ciliate; ray achenes 2.2—4.2 mm long, awnless; disc achenes obovate, 4—6 mm long, 4.0-5.5 mm wide, subglabrous, often reddish, wings to ca. 1.8 mm wide, the margins entire or, more often, lacerate, the awns sub- equal, 2.8-4.0 mm long, slightly spreading.
Distribution. Mountains of the Cape region of Baja California.
Specimens examined. BAJA CALIFORNIA. Pescadero, Brandegee s. n., Sept. 20, 1893 (UC); La Chuparosa, Brandegee s. n., Oct. 2, 1899 (UC) ; Sierra de San Fran- cisquito, Brandege s. n., Sept. 29, 1899 (F); Sierra San Laguna, Brandegee s. n., Sept. 10, 1899 (NY); W side of Cape Region, Brandegee s. n., Nov. 1902 (UC); San Filipe, Purpus s. n., Jan-Mar. 1901 (UC); Sierra El Taste, Carter & Chisaki 3309 (UOC).
Verbesina erosa has its closest affinity with V. pustulata, differing from that species by its more closely serrate leaf margins and awnless ray ovaries. These two species are so similar that it is quite possible that further collecting will show them to be conspecific.
9, VERBESINA PUSTULATA Jones, Contr. West. Bot. 18:77. 1933. Type, The Laguna, Laguna Mountains, Baja California, Jones 22711, UC!
Suffruticose, ca. 1-2 m tall; stems hispid, red; leaves opposite, the petioles ca. 1 cm long, narrowly winged, lance—ovate, closely serrate, acute or acuminate, to ca. 7 cm long, 2.5—3.0 cm wide, densely hispid- scabrous; inflorescences subcorymbose, ca. 4-headed, pedicels ca. 3 cm long; discs ca. 1 cm wide; phyllaries ca. 4-seriate, oblong or lanceolate, subequal, 5—7 mm long, acute, the margins ciliate; ray florets styliferous, the tube densely hirsute, the ovary ca. 1.5 mm long, glabrous, the awns ca. 1.7 mm long; disc florets ca. 9 mm long, the tube and throat hirsute, the ovary ca. 2.8 mm long, glabrous, the awns ca. 3.0—4.8 mm long, the style tips attenuate; receptacle conical; chaff ca. 7-8 mm tall, acuminate, subscarious, the margins ciliate; disc achenes (immature) obovate, 4—5 mm long, subglabrous, the wings narrow, the margins ciliate, the awns 2.8-3.5 mm long, one ca. % the length of the other.
1966] CASTETTER AND PIERCE: ESCOBARIA 137
Verbesina pustulata, known only from its type material, is most closely related to V. erosa. The reader is referred to the discussion under that species.
Department of Botany, Indiana University, Bloomington
LITERATURE CITED
BuakgE, S. F. 1924a. New South American Verbesinas. Bull. Torrey Club 51:421-436. . 1924b. Eight new Asteraceae from Mexico, Guatemala, and Hispaniola. Proc. Biol. Soc. Wash. 37:55-61.
CoLeMaAN, J. R. 1964. A taxonomic revision of sections Pterophyton, Sonoricola and Ximensia of the genus Vergesina (Compositae). Ph. D. thesis (unpublished). Indiana University, Bloomington.
De Jone, D. C. D. and E. K. Loncpre. 1963. Chromosome studies in Mexican Com- positae. Rhodora 65:225-239.
Heiser, C. B. and D. M. SmirH. 1955. New chromosome numbers in Helianthus and related genera (Compositae). Proc. Indiana Acad. 64:250-253.
Rosinson, B. L. and J. M. GreenMAN. 1899, A synopsis of the genus Verbesina, with an analytical key to the species. Proc. Am. Acad. 34:534-564.
Turner, B. L., W. L. Ettison, and R. M. Kinc. 1961. Chromosome numbers in the Compositae IV. North American species with phyletic interpretations. Am. Jour. Bot. 48:216-223.
, J. H. Beaman and H. F. L. Rock. 1961. Chromosome numbers in the Compositae V. Mexican and Guatemalan species. Rhodora 63:121-129.
ESCOBARIA LEEI BODEKER REDISCOVERED IN NEW MEXICO
EDWARD F. CASTETTER AND PRINCE PIERCE
This species was briefly described by Bodeker in 1933 as follows: “Escobaria Leet (Rose as Mammillaria 1924 No. 282) Bod. From New Mexico. Plants proliferating at the ground, only 15 mm. high and 5 mm. thick; spines numerous, white, up to 3 mm. long.”’ Although Bodeker refers to a Mammillaria specimen No. 282 of J. N. Rose, we have been unable to locate any reference or description of this plant by Rose in the literature. Despite the obvious shortcomings of Bodeker’s descrip- tion, the name E&. /eez holds priority and we have decided to publish a detailed, meaningful description of the species on the basis of field and laboratory studies of numerous specimens.
In the U. S. National Herbarium is an herbarium sheet (US 72134) with the following information on the label: Rattlesnake Canyon, 30 miles southwest of Carlsbad, altitude 5500 ft., W. T. Lee, 1924. Attached to the herbarium sheet is a piece of notepaper which reads: ‘‘Found in Rattlesnake Canyon, 30 miles southwest of Carlsbad, New Mexico, at 5500 ft. elevation, growing on limestone ledges on north facing slopes. It is new. I have never seen it before. W. T. Lee.”
138 MADRONO [Vol. 18
Fic. 1. Escobaria leei
This sheet bears, in Rose’s handwriting, the number 24,282 which was his greenhouse number and represents specimen 282 collected in the year 1924. Conrad Morton of the U.S. National Herbarium is of the opinion that, although there is no record of it, Bodeker may have visited the National Herbarium at some time while Rose was there and may very well have studied this specimen; he is convinced that this specimen is the basis for Escobaria leei (from a letter by S. G. Shetler to L. Ben- son, Feb. 9, 1965).
Also in the National Herbarium is a carton containing a whole plant consisting of a cluster of numerous small stems together with a slip of paper bearing the following annotation: ‘“Neom. Leev”’, Carlsbad, N. Mex., 24,282, see N. lasiacantha, April 1925, Lee, 24,282. Thus, both the slip of paper and the carton plant tie in with the herbarium sheet by the same greenhouse number. The fact that Rose had marked this specimen as Neomammillaria leei indicates that he had considered describing it as a new species which, however, he never did.
Plants on the herbarium sheet (US 72134) and the carton specimen are designated as the lectotype of E. leez.
We recently found this species near the original collections site in Rattlesnake Canyon (fig. 1) about 30 miles southwest of Carlsbad, in the tributaries of this canyon and in Slaughter Canyon, at elevations of 4200 to 4800 feet, in the Guadalupe Mountains of southwestern New Mexico. We have studied the plants in situ and have grown them in our gardens (Castetter & Pierce 2397 through 2717, UNM).
1966] CASTETTER AND PIERCE: ESCOBARIA 139
EscoBARIA LEEII Bodeker, Ein Mammillarien Vergleichs, Schlussel 17, Neudam, Germany, 1933. Low growing plants usually forming densely spreading clumps bearing as many as 250 stems (usually not more than 100) formed by proliferation from the bases of stems of older plants and thus forming flat tufts or mats up to 15 cm in diameter; with most clumps, all stems remaining immature for a considerable period of time; mature stems as much as 7.5 cm long and 3 cm in diameter but usually not exceeding 3.5 by 1.7 cm; stems typically clavate, less frequently cylindric, the smallest ones subglobose and as little as 3 cm in diameter; aerial spiny portion of stem extending as much as 5 cm above ground level, densely white spiny, the lower part of the stem subter- ranean, bearing dry knobby tubercles although many spines on these tubercles remain intact; flesh of stem ranging from green to pale or deep dull magenta, firm, tough; root system mainly fibrous, but usually with a single moderately slender taproot which is connected to the original stem of the clump. Tubercles cylindric or nearly so, small, as much as 0.5 cm long by 2—3 mm wide, almost completely obscured by the dense spine clusters; some of the older tubercles grooved on the upper surface, the groove extending from the tip of the tubercle to % or more of its entire length, short white woolly, with a dense woolly tuft at the base; aeroles circular to broadly elliptic in outline, densely white woolly becoming dark in age. Central spines irregular in length, but as much as 1 cm long, stout, straight, white to yellowish-brown tipped, varying in number from 6-22, 6 or 7 of them stouter than the others, and 1 of these, not necessarily the longest of the group, occupying a truly central position, porrect; most of the central spines depressed and subcentral in appearance; radial spines 40-85, white, very slender, irreg- ular in length, some of them curved or twisted at the tip, both radial and subcentral spines disposed in an eccentric whorl; all spines on very young plants densely short white pubescent, those on older stems naked. Flowers funnelform, 1.7—2.0 cm long, 1.2—1.8 cm in diameter, overall color dull medium brownish-pink; outer perianth segments terminating in a short awn, the broad midstripe brown to greenish-brown, ciliate on the margins, bearing copious long white translucent, curled, twisted, kinked hairs on the upper 12—34 of the segment, some of them branched near the bases; inner perianth segments each ending in an abrupt short pointed tip, white, the broad midstripe rose to dull orchid with a brownish infusion; stigma lobes 4—6, short oblong, pure white. Fruit pale brownish-green to green with a pinkish cast, 1.2 cm long, about 0.5 cm in diameter, most of them bearing 1—several pubescent scales; perianth persistent; dry seeds nearly obovoid, more specifically asym- metrically pear-shaped on one side to somewhat kidney shaped on the other, alveolate, lustrous reddish-brown, about 1.25-i1.5 mm long by 1 mm wide, hilum subbasal, flat.
140 MADRONO [Vol. 18
Plants usually begin to bloom in their native habitat in mid-April to early May and continue to do so for several weeks, closing at night and opening about noon.
This species differs from all others of the genus in New Mexico in that the aerial portion of many of the medium-sized and smaller stems has a unique shape which may be described as blockish-spherical, a characteristic attributable to the fact that the central part of the aerole projects beyond the spine cluster, owing to most spines being depressed.
University of New Mexico, Albuquerque
NEW RECORDS OF MYXOMYCETES FROM CALIFORNIA I. DONALD T. KOWALSKI
A search of the pertinent literature, which consists of some eight papers dealing with slime molds of California, reveals that 154 species of Myxomycetes have been definitely recorded as occurring in California. Hagelstein (1944) listed additional species that have been reported, but he considered their authenticity questionable for various reasons. The largest collection was that of Plunkett (1934). He gave a list of 87 species, most of which were collected within a 50 mile radius of Los Angeles. The only papers dealing with Myxomycetes of Northern Cali- fornia were those of Cooke (1949) and Pratt and Pratt (1944). All of Cooke’s collections were made on the slopes of Mt. Shasta and the Pratts collected their specimens within a 25 mile radius of San Francisco. Thus, essentially no Myxomycetes have been reported to date from the Sacramento Valley.
In the fall of 1964 I initiated a study of the Myxomycetes of Cali- fornia, and since have collected exclusively in Butte Co. Most of the collections listed below have come from Lower Bidwell Park, Chico. This is one of the few remaining areas in the Sacramento Valley that has not been disturbed for reasons of agriculture. Thus, it represents a particu- larly fruitful area and the species found here may have been abundant at one time throughout the Great Valley. In this first of a series of papers, 11 species of Myxomycetes are listed as new to the state in the sense that no report of their occurrence in California has been previously published. This brings the total number of slime molds found in Cali- fornia to 165 species.
All collections have been deposited in the Herbarium of the Univer- sity of Michigan. The names of the organisms are those accepted by Martin (1949) and the numbers are my own.
1966] KOWALSKI: CALIFORNIA MYXOMYCETES 141
LICEACEAE
Licea pusilla Schrad. On badly decayed coniferous wood approximately 5 miles N of Forest Ranch, 3500 ft., 1220, Mar. 19, 1965. The sporangia are purplish-brown, sessile, and dehisce by preformed lobes, appearing stellate at maturity. This exceedingly tiny species, the maxi- mum diameter of the sporangia being only 1.0 mm, has been found previously only as far west as Iowa, and is considered rare. This, how- ever, is undoubtedly due to its inconspicuousness and it may occur throughout the United States.
TRICHIACEAE
Perichaena vermicularis (Schw.) Rost. No. On a dead herbaceous stem in Lower Bidwell Park, 1079, Jan. 16, 1965. This collection con- sists of a few scattered plasmodiocarps, but can be clearly identified by its light brown color and large spores, reaching 14.0 » in diameter.
Arcyodes incarnata (Alb. & Schw.) O. F. Cook. On a decorticated log near the Sacramento River approximately 1.7 miles S of Sacramento Road, 972, Dec. 12, 1964. The fructifications consist of sessile, heaped, pinkish sporangia, each about 1 mm in diameter. They are quite dis- tinctive by reason of their persistent peridia. This is considered a rare species and is known from scattered collections across the United States. It is not new to the west coast, having been reported from Oregon.
Arcyria occidentalis (Macbr.) G. Lister. On decayed wood, 5 miles N of Forest Ranch, about 3500 ft., 1161, Mar. 6, 1965. This collection has spores averaging 9-10 » in diameter. All published descriptions list the spores as 7—8 » in diameter. In all other respects, however, it fits per- fectly with the description given by Martin (1949), In my opinion, this discrepancy is not large enough to warrant the establishment of a new species or variety.
Arcyria insignis Kalchbr. & Cooke. Three collections, all on decayed wood in Lower Bidwell Park: 1049, Dec. 31, 1964; 1268, Apr. 17, 1965; and 1302, Apr. 24, 1965. This common species has previously somehow been overlooked in California. It can be distinguished from other mem- bers of the genus by its small size, firmly attached capillitium, and pinkish color.
STEMONITACEAE
Enerthenema papillatum (Pers.) Rost. Two collections, both on de- cayed wood in Lower Bidwell Park: 1086, Jan. 16, 1965 and 1307, Apr. 24, 1965. This common species is characterized by having the columella expanding into a disk at the apex of the sporangium, and nonclustered spores which are only minutely warted.
Stemonitis hyperopta Meylan. On decayed wood in Lower Bidwell Park, 1111, Jan. 24, 1965. This collection consists of just a few spo- rangia, but it is easily recognizable by the reticulate spores, which are inconspicuously and irregularly banded-reticulate. It is another rare species which is known from scattered collections from Maine to Wash- ington.
142 MADRONO [Vol. 18
Stemonitis pallida Wing. Two collections, both on decayed wood in Lower Bidwell Park on Apr. 24, 1965: 1301 and 1310. This species is characterized by having spores that are nearly smooth and over 7.0 p» in diameter, sporangia which are pallid in color, and a surface net which is poorly developed towards the apex. It is an uncommon species, pre- viously being found only as far west as Iowa.
Comatricha cornea Lister & Cran. Three collections: 939 on bark of a fallen tree in Lower Bidwell Park, Nov. 26, 1964; 986 and 1002 which developed in moist-chamber culture on bark taken from living trees along the Sacramento River 1.7 miles S of Sacramento Road, Dec. 12, 1964. This species has been reported only from Michigan, Kansas, and Iowa and then only from moist-chamber culture. I have thus collected this species in the field for the first time (939). The small collar at the base of the columella and the translucent stalk are the outstanding characteristics of this minute species.
DIDYMIACEAE
Didymium serpula Fries. Two collections, both from Lower Bidwell Park on decaying leaves: 1009, Dec. 31, 1964 and 1183, Mar. 14, 1965. The grayish plasmodiocarps with vesiculose bodies attached to the capil- litium separate this species from others in the genus. It is a very rare taxon, previously found only as far west as Iowa.
Didymium minus (Lister) Morgan. On a decaying leaf in Lower Bid- well Park, 1115, Jan. 24, 1965. This is a cosmopolitan species, somehow overlooked or misidentified in California until now. Its major character- istics are: stalked subglobose sporangia, the stalks being dark-brown or black, and spores 8-11 » in diameter. Didymium melanospermum (Pers. ) Macbr. is closely allied with this species but has spores 10-14 » in diam- eter. It is almost a certainty that many California collections labeled D. melanospermum are in reality D. minus, since many authors consider D. minus a variety of D. melanospermum. This probably accounts for its exclusion from California in the published reports.
Department of Biology, Chico State College, Chico, California
LITERATURE CITED
Cooke, W. B. 1949. Myxomycetes of Mount Shasta, Madrono 10:55-62. HAGELSTEIN, R. 1944. Mycetozoa of North America. Mineola. Martin, G. W. 1948. Two new species of Physarum. Jour. Wash. Acad. 38:238-240. . 1949. North American Flora 1:1-190. . 1961. The genus Schenella. Mycologia 53:25-30. Puitiies, W. 1877. Fungi of California and the Sierra Nevada Mountains. Grevillea 5:113-118. PrunketTT, O. A. 1934. Contributions to the knowledge of Southern California fungi. I. Myxomycetes. Univ. Calif. Publ. Biol. Sci. 1:35-48. Pratt, R. and J. Pratt, 1944. Myxomycetes of the San Francisco region. Amer. Jour. Bot. 31:559-561.
CONTRIBUTION TO THE EMBRYOLOGY OF MUILLA, WITH A REMARK ON THE TAXONOMIC POSITION OF THE GENUS
RotF Y. BERG AND JACK R. MAZE
This paper is the first of an intended series on the tribe Allieae, sub- family Allioideae, of the family Liliaceae (sensu Krause, 1930) or Ama- ryllidaceae (sensu Hutchinson, 1959). This series will be a continuation of previous studies within the Liliaceae (Berg, 1958; 1959; 1960; 1962a; 1962b).
Muilla (anagram of Allium) is a small genus of five species native to the southwestern United States and Mexico (Ingram, 1953). Only one species, MM. maritima (Torr.) Wats., was studied. Megasporogenesis, embryo sac development, and early endosperm development is described, as well as the accompanying changes in the ovule. The taxonomic signifi- cance of this new information is considered and inconsistencies with the present taxonomic position of Muilla are pointed out.
No information about the embryology of Mulla is available in the literature.
Material was collected from plants growing in an alkaline flat by the Pole Line Road, 1.1 miles north of Davis city limit, Yolo Co., California (DAV). Ovaries were fixed in Belling’s modified Navashin fluid and dehydrated with tertiary butyl alcohol. Sections were cut at 10 and 12 microns and stained with safranin and fast green.
We are grateful to Marion S. Cave for critical review of the manu- script.
OBSERVATIONS
OvuLe. The ovule is anatropous, bitegmic, and crassinucellate (fig 3). At the early megaspore mother cell stage, it is short and nearly straight (fig. 1); at the megaspore tetrad stage it has become curved back ap- proximately 90° (fig. 2); the anatropous stage is attained at ca. the four-nucleate embryo sac stage. There is no distinct funiculus. Raphides were not observed in the ovule, but may develop in the seed, since they are common in the ovary wall.
The inner integument develops near the apex at approximately the stage when the archesporial cell divides (fig. 1). Throughout develop- ment to the proembryo stage, the inner integument is two cells thick, except at the apex where it becomes three to five cells thick. The cells at the apex are more or less isodiametric whereas the remainder are elongate, being two to three times as long as broad at the time of fertili- zation (fig. 13). The outer integument begins to develop at the young megaspore mother cell stage and by the time of meiosis is well defined. It is three cells thick early in its development but soon develops into a structure that is four to five cells thick, the cells being approximately
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Fics. 1-12. 1, Ovule from a bud which was still underground, showing megaspore mother cell, primary parietal cell, and young annular inner integument; 2, Ovule from a bud located at the soil surface, showing a T-shaped tetrad and short integu- ments; 3, Ovule from a young bud, its pedicel still short, showing embryo sac before fusion of polar nuclei with inner integument closed above nucellus; 4, Young seed, showing helobial endosperm of four plus eight nuclei (all nuclei projected into one plane). Note characteristic elongation of outer integument; 5, young ovule, show- ing the arrangement of nucellar cells and an early stage of integument develop- ment; 6, Nucellus at a somewhat later stage with two parietal cells and megaspore mother cell in diakinesis; 7, Nucellus with two tiers of parietal cells and megaspore mother cell nucleus in metaphase I; 8, Linear tetrad in which division in upper dyad
1966] BERG AND MAZE: MUILLA 145
Fics. 13-17. 13, Integuments shortly before anthesis. 7: inner integument, o: outer integument; 14, Nucellus from a medium-sized bud, showing numerous periclinal walls in upper part of nucellar epidermis, characteristic radial rows of cells in lower part, and a central core of narrow cells directly below the embryo sac; 15, Mature embryo sac as seen at anthesis; 16, Young helobial endosperm, both chambers four-nucleate. One synergid destroyed and antipodals degenerating; 17, Endosperm of 128 plus 8 nuclei surrounded by greatly enlarged nucellus. a: rem- nants of antipodal cells, c: central core of nucellar cells, e: one-layered nucellar epidermis, ee: two-layered nucellar epidermis, p: proembryo. Figs 13-15 360, fig. 16 X90, fig. 17 X56.
cell is slightly delayed; 9, T-shaped tetrad; 10, Nucellus showing first periclinal wall in epidermis, three tiers of parietal cells above the megaspores, and germination of lower megaspore; 11, Two-nucleate embryo sac stage; 12, Four-nucleate embryo sac stage. All parietal cells are absorbed. Note characteristic radial arrangement of cells in nucellus, s: stigmatoid tissue. Figs. 1-3 90, fig. 4 «56, figs. 5-12 360.
146 MADRONO [Vol. 18
isodiametric at the time of fertilization (fig. 13). The inner integument closes above the nucellus approximately when the embryo sac has reached the four-nucleate stage (figs. 11-12). The outer integument remains short for a long time so that the micropyle is formed by the inner integument only (fig. 3). However, after fertilization the outer integument grows considerably to form an unusual elongation towards the placenta (fig. 4).
The nucellus is medium-sized and completely surrounds the embryo sac at the time of fertilization (fig. 3). By this time the nucellus is ap- proximately twice as long as the embryo sac and nearly as broad as long. The nucellar epidermis undergoes periclinal divisions, the first of which are found shortly after the germination of the functioning megaspore (fig. 10). These periclinal divisions in the nucellar epidermis continue until the time of fertilization but are rstricted to the micropylar half of the nucellus only (figs. 11-12, 14), where a multiple epidermis of two, rarely three, layers eventually forms. Most of the body of the nucellus develops from the cells immediately beneath the epidermis. As the ovule grows, these subepidermal nucellar cells divide repeatedly, both pe- riclinally and anticlinally, to form a characteristic pattern of radiating rows of cells (figs. 11-12, 14). After fertilization, more cells are added in this region and, as the ovule grows, most nucellar cells enlarge greatly. However, those of the central core region at the chalazal end of the embryo sac and those of the multiple epidermis at the micropylar end of the embryo sac remain small, and contribute only insignificantly towards the size of the post-fertilization nucellus (fig. 17).
ARCHESPORIUM AND MEGASPOROGENESIS. Only one archesporial cell is present in the ovule of Mulla maritima. Before the bud appears above eround in spring, the archesporial cell has divided to form the megaspore mother cell and a primary parietal cell (fig. 5). The primary parietal cell divides anticlinally, or periclinally, while the ovule is still in the mega- spore mother cell stage (fig. 6). The two cells resulting from this division divide further to give rise to several parietal cells arranged in three, sometimes four, tiers above the megaspores (figs. 7-10). Some of the parietal cells remain until the two-nucleate stage in embryo sac develop- ment (fig. 11), after which all are absorbed by the developing embryo sac (fig. 12).
Megasporogenesis begins with the elongation of the megaspore mother cell (figs. 5-6). The spindle of the first meiotic division is longitudinally oriented and located a little above the middle of the cell (fig. 7). Nuclear division is followed by wall formation resulting in a dyad, the micro- pylar cell of which is slightly smaller than the chalazal one. Each dyad cell divides, the chalazal one normally by a cross-wall, the micropylar one by a wall which is sometimes horizontal but more often longitudinal or oblique (figs. 8-9). The division in the upper dyad cell may be slightly delayed (fig. 8). The tetrad of megaspores will be linear or T-shaped. At the time of formation, the two lowermost megaspores are
1966] BERG AND MAZE: MUILLA 147
about equal in size and a little larger than the uppermost two (fig. 8). Embryo sac development is monosporic. The upper three megaspores degenerate, normally starting with the micropylar one (fig. 10) and proceeding progressively downward. In all ovules observed, with one possible exception, the chalazal megaspore functioned.
MEGAGAMETOGENESIS AND MEGAGAMETOPHYTE. By enlargement and vacuolization, the functioning megaspore becomes the embry sac mother cell (fig. 10). Normally two vacuoles are formed, one above and one below the nucleus. As the embryo sac grows, the upper vacuole en- larges to become the central vacuole (figs. 11-12) while the lower one remains small and finally disappears (figs. 11-12, 14-15). The eight- nucleate stage is reached by three successive divisions of the embryo sac mother cell nucleus (figs. 10-12, 14). The micropylar part of the embryo sac increases more in size than the chalazal part (figs. 11-12), the latter eventually forming a small, narrow portion where the antipodals lie (figs. 14-15).
The mature embryo sac (fig. 15) is approximately one-and-a-half times as long as broad and of typically ovoid shape. The synergids are similar in size and appearance and are smaller than the egg. They are pear-shaped, each with a vacuole in the chalazal end, a nucleus located immediately above the vacuole, and a filiform apparatus radiating out from the point of attachment. The large egg cell has the nucleus in its lower end and is highly vacuolized. The three antipodal cells are ar- ranged differently in different embryo sacs; most often two are found at the same level with the third one above (fig. 15) or below (fig. 14). Occasionally they are all in a row (fig. 16). The antipodal cells are as large as, or somewhat larger than, the synergids. They begin to degen- erate a little before or during fertilization, but they are still obvious in early endosperm stages (figs. 16-17). The polar nuclei fuse before fertilization to form a large secondary nucleus, which lies close to the antipodals (figs. 14-15).
FERTILIZATION AND ENDOSPERM DEVELOPMENT. After pollination the pollen tube grows down the open stylar canal, which is lined with papil- late stigmatoid tissue. The stigmatoid tissue of the style connects with a similar tissue on the placentae, some of which is apparent in figs. 2 and 3. One of the synergids is destroyed when the pollen tube enters the embryo sac. The other remains until more than a hundred nuclei have formed in the endosperm. Double fertilization occurs. Fusion between the second- ary nucleus and one of the male nuclei takes place before the fusion in the egg nucleus. Apomictic phenomena were not observed.
The primary endosperm nucleus starts to divide before the zygote. Endosperm is helobial. The first nuclear division takes place in the chalazal end of the endosperm cell. It is followed by the formation of a transverse wall, which cuts off a smaller chalazal chamber from the larger micropylar one. In both chambers free nuclear divisions occur, and in the bginning they are synchronous. The following stages were
148 MADRONO [Vol. 18
observed (nuclei of micropylar chamber/nuclei of chalazal chamber): 1/1, 2/2, 4/4, 8/4, 16/4, 32/4, 8/8, 16/8, 128/8. To judge from this series, nuclear divisions occur simultaneously in the two chambers until four nuclei are present in each (fig. 16). Then development slows down in the chalazal chamber: after eight, or possibly sometimes only four, nuclei have been formed in this chamber, nuclear divisions apparently cease. The nuclei already present become hypertrophied to some extent and the chamber shows signs of degeneration. Since later stages were not present in our material, the ultimate fate of the chalazal endosperm chamber could not be ascertained. In the micropylar chamber, nuclear divisions continue (figs. 4, 17), and the endosperm proper is wholly or largely developed from this chamber.
DISCUSSION
A full discussion of embryologic similarities and dissimilarities be- tween the genera of the tribe Allieae must wait until data, comparable to those presented above for Muilla, are available for the remaining genera of the tribe. Such a discussion will constitute the final paper of this series. However, in a preliminary way, a few interesting facts can be pointed out.
The tribe Allieae, as conceived by Krause (1930), consists of 16 genera, some of which later have been united. Embryologic information has, so far, been produced only for Gagea, Allium, Nothoscordum, Bro- diaea, and Leucocoryne. Gagea, furthermore, is now generally considered to be a member of the subfamily Lilioideae (Berg, 1962b). The informa- tion on Brodiaea (which is at present under investigation by the senior author) is limited to a record of helobial endosperm in B. peduncularis (Stenar, 1949). We are left, therefore, with only three genera for our preliminary survey, viz. Allium, Nothoscordum, and Leucocoryne (table ily
Of special interest is the lack of a parietal cell in all these three genera, as opposed to the presence of a multicellular parietal tissue in Muilla. Stenar (1932, p. 39) points to the absence of a parietal cell as a typical feature of the Allieae. The bisporic type of embryo sac develop- ment in Allium and Leucocoryne and the nuclear type of endosperm in Allium constitute additional important deviations from the Muilla pat- tern. Allium differs from Muilla also in the shape of the embryo sac, in the number of cell layers in the integuments, in the position of the secondary nucleus, and in other minor features. Nothoscordum is most similar to Muilla, but this genus is also set somewhat apart from Muilla by the nature of its nucellus and the nature of its synergids, as well as by the absence of a parietal cell, and the occurrence of an Allium type embryo sac in one of its species.
The “key combination” (Berg, 1962b) of “+ N He” (parietal cell present, normal- or Polygonum type embryo sac, and helobial endo- sperm) that characterizes Muilla is not found in any other genus of the
1966] BERG AND MAZE: MUILLA 149
TABLE 1. EMBRYOLOGICAL COMPARISON OF SOME GENERA OF THE TRIBE ALLIEAE
(+ = yes or present, 0 = no or absent)
Allium Nothoscordum Leucocoryne Muilla
No. of cell layers
in inner integument 3-4 2 2 No. of cell layers in
outer integument 4-10 2-4 3-5 Periclinal walls in
nucellar epidermis + - +6 + Numerous such walls 0 0 06 + Bulk of nucellus
produced from
nucellar epidermis 0 0 06 0 Nucellar apex pene-
trated by embryo sac 01 0 06 0 Parietal cell 0 0 0 ao Type of embryo
sac development Allium? Polygonum? Allium Polygonum Lower spore functions + 4. + + Length/ width of
embryo sac 4/2 Sie 3y2 3/2 Synergid filiform
apparatus O&+ oi + Synergids vacuolated O&+ 0 + Polar nuclei fuse
prior to fertilization 0&-+ +5 ot + Endosperm type Nu Het He References Hakansson, 1951 Eckles, 1941 Cave, 1939
Jones and Hakansson, 1953
Emsweller, 1936 Messeri, 1931 Messeri, 1931 Schnarf, 1931 Murphy, 1946 Stenar, 1932 Rao, 1940
Schnarf, 1931
Schiirhoff, 1922
Weber, 1929
1Weber (1929), Hakansson (1951), and Hasitschka-Jenschke (1958) state that the nucellus is partly destroyed when the embryo sac is at the four-nucleate stage.
~Weber (1929) reported both mono- and tetrasporic embryo sac development for Allium carinautm and monosporic embryo sac development for A. paradoxum However, the significance of these variants is questinable because both of these species reproduce by bulblets and either the embryo sac does not develop or, if it does, the embryo does not survive. All normally reproducing, amphimictic species show Allium type embryo sac development.
3 One of the three species of Nothoscordum included in this table, VN. fragrans, is different from the others in that the embryo sac development is of the Allium type (Messeri, 1931; Stenar, 1932). However, it is of interest to note that this species shows adventive polyembryony and may present a similar case as Allium carinatum and A. paradoxum, i.e. an abnormal deviation within its genus.
4Schnarf (1931, p. 242) states that nuclear endosperm has been encountered in Gagea, Allium, and Nothoscordum, a statement that as far as Nothoscordum is con- cerned is contrary to Stenar’s (1932) observations.
> Polar nuclei fuse after fertilization, according to Hakansson (1953).
6 Marion Cave, personal communication.
150 MADRONO [Vol. 18
tribe Allieae. On the other hand, Muilla’s key combination occurs in several genera of four other subfamilies of the Liliaceae sensu Krause, namely the Melanthioideae (Schnarf, 1929), the Asphodeloideae (Schnarf and Wunderlich, 1939; Cave, 1953), the Scilloideae (Wunderlich, 1937; Buchner, 1948), and the Aletroideae (Schnarf, 1929), as well as in some genera of the family Amaryllidaceae (Schnarf, 1931; Stenar, 1951), to which the Allioideae is referred by Hutchinson (1959).
From a taxonomic point of view, the difference in embryological char- acters that exists between Muilla and other genera of the Allieae, par- ticularly Allium, is important since it indicates that Muilla certainly is more distantly related to Allium and the other Allieae here discussed than is now generally assumed. For lack of information about the embryo- logical variation within the Allieae as a whole, this line of thought can- not possibly be pursued any further at this time. For the same reason a detailed elaboration on the embryologic similarity and possible rela- tionship of Muilla to one or more genera of the Melanthioideae, Aspho- deloideae, Scilloideae, Aletroideae, or Amaryllidaceae sensu Krause will have to wait.
SUMMARY
The ovule is anatropous, bitegmic, and crassinucellate. The inner in- tegument consists of two, the outer of four to five, layers of cells. The micropyle is formed by the inner integument only, but after fertilization the outer integument elongates considerably and forms a characteristic protrusion toward the placenta. Periclinal walls occur in the nucellar epidermis, but the bulk of the nucellus is produced from the original subepidermal cells. At the time of fertilization, the massive nucellus is one-and-a-half times as long as the embryo sac and completely sur- rounds it. A parietal tissue of several cells is present. Embryo sac devel- opment is according to the Polygonum type. The endosperm is helobial.
Numerous differences in embryologic characters exist between Muilla and the other genera of the Allieae that have been investigated in this respect. Probably Muilla is more:distantly related to these members of the Allieae than is realized at present. Embryologically, Muilla shows similarity with several genera of the subfamilies Melanthioideae, Aspho- deloideae, Scilloideae, and Aletroideae of the Liliaceae, and with genera of the family Amaryllidaceae sensu Krause.
Department of Botany, University of California, Davis
LITERATURE CITED
BerG, R. Y. 1958. Seed dispersal, morphology, and phylogeny of Trillium. Skr. Vid.- Akad. Oslo I. Mat.-Nat. K] 1959(1) :1-36. . 1959. Seed dispersal, morphology, and taxonomic position of Scoliopus, Liliaceae. Skr. Vid.-Akad. Oslo I. Mat.-Nat. KI. 1959(4) :1-56.
1966] BERG AND MAZE: MUILLA 151
. 1960. Ovary, ovule, and endosperm of Calochortus amabilis with remarks on the taxonomic position of Calochortus. Nytt Mag. Bot. 8:189-206.
. 1962a. Morphology and taxonomic position of Medeola, Liliaceae. Skr. Vid.-Akad. Oslo I. Mat.-Nat. KI. 1962 (3) :1-55.
. 1962b. Contribution to the comparative embryology of the Liliaceae: Scoliopus, Trillium, Paris, and Medeola. Skr. Vid.-Akad. Oslo I. Mat.-Nat. KI. 1962 (4) :1-64.
BucuNeER, L. 1948. Vergleichende embryologische Studien an Scilloideae. Osterr. Bot. Zeitschr. 95:428-450. Cave, M. S. 1939. Macrosporogenesis in Leucocoryne ixioides Lindl. Cytologia
9:407-411. . 1953. Cytology and embryology in the delimitation of genera. Chron. Bot. 14:140-153.
Ecxies, M. L. 1941. Megasporogenesis and development of the megagametophyte in Nothoscordum bivalve. Bot. Gaz. 102:806-809.
HAxansson, A. 1951. Parthenogenesis in Allium. Bot. Not. 1951:143-179.
. 1953. Die Samenbildung bei Nothoscordum fragrans. Bot. Not. 1953:129- 139.
HASITSCHKA-JENSCHKE, G. 1958. Die Entwicklung der Samenanlage von Allium ursinum mit besonderer Bericksichtigung der endopolyploiden Kerne in Syner- giden und Antipoden. Osterr. Bot. Zeitschr. 104:1-24.
HvutTcuHInson, J. 1959. The families of flowering plants. Vol. IJ. Monocotyledons. Second ed., Oxford.
IncrRAM, J. 1953. A monograph of the genera Bloomeria and Muilla. Madrono 12:19-27.
Jones, H. A. and S. L. EMSwELter. 1936. Development of the flower and macro- gametophyte of Allium cepa. Hilgardia 10:415-428.
Krause, K. 1930. Liliaceae. Jn Die nattrlichen Pflanzenfamilien 15a:227-386.
Messer, A. 1931. Ricerche embriologiche e cariologiche sopra i generi “Allium” e “Nothoscordon’”’. Nuovo Gior. Bot. Ital. 38:409-441.
Murpnuy, J. B. 1946. Megasporogenesis and development of the embryo sac of Allium cernuum. Bot. Gaz. 108:129-136.
Rao, Y. 1940. Structure and development of the embry sac of Drimiopsis kirki Baker and Allium govanianum Wall. Jour. Indian Bot. Soc. 19:273-282.
ScHNnaRF, K. 1929. Die Embryologie der Liliaceae und ihre systematische Bedeutung. Sitz.-ber. Akad. Wien math.-nat. KI]. Abt. I, 138:69-92.
. 1931. Vergleichende Embryologie der Angiospermen. Berlin.
. and R. Wunderlich. 1939. Zur vergleichenden Embryologie der Liliaceae- Asphodeloideae. Flora 133:297-327.
SCHURHOFF, P. N. 1922. Zur Polyembryonie von Allium odorum. Ber. Deutch. Bot. Ges. 40:374-381.
STENAR, H. 1932. Studien iiber die Entwicklungsgeschichte von Nothoscordum fragrans Kunth und N. striatum Kunth. Sv. Bot. Tidskr. 26:25—-44.
. 1949. Zur Kenntnis der Embryologie und der Raphiden-Zellen bei Bowiea volubilis Harvey und anderen Liliazeen. Acta Horti Berg. 15:45-63.
. 1951. Zur Embryologie von Haemanthus katharinae Bak., nebst Erorter- ungen uber das helobial Endosperm in den Amaryllidaceae und Liliaceae. Acta Horti Berg. 16(1954) :57-72.
WesER, E. 1929. Entwicklungsgeschichtliche Untersuchungen tiber die Gattung Al- lium. Bot. Arch. 25:1-44.
WunperRLicH, R. 1937. Zur vergleichenden Embryologie der Liliaceae-Scilloideae. Flora 132:48-90.
THE GENUS TIARELLA IN WESTERN NORTH AMERICA
PATRICIA KERN
This paper treats the three western North American taxa of Tiarella, a genus of the Saxifragaceae which is distinguished by two unequal valves of the dehiscent fruits and panicles of white flowers. These taxa have previously been treated as distinct species, recognizable by the basal leaf conformation. Study of the complex by analysis of population samples, field observations, and experimental breeding clearly indicates that there is continuous variation among the three. Their recognition as elements of one species complex, T. trifoliata L. is proposed. Within this species, two subspecies, ssp. wnifoliata and ssp. trifoliata with two vari- eties, var. trifoliata and var. laciniata, are recognized.
DISTRIBUTION. The most widely distributed of the three, ssp. unifoli- ata has its eastern boundary in western Alberta and western Montana. From Alaska it extends southward, along both sides of the Cascade range, to the Santa Cruz Mountains in California. In California, it grows as low as at 150 feet elevation, whereas in most other areas it is found growing in moist forests in the mountains above 2000 feet.
Subspecies tvifoliata is centered primarily west of the Cascade Moun- tains in Washington, but extends from Alaska to southern Oregon and to eastern British Columbia, and is found in scattered localities in western Montana, Idaho, and eastern Washington. It occurs in unmixed stands in western Washington at elevations less than 1500 feet.
Subspecies trifoliata and unifoliata both occur, mostly at altitudes that are mutually exclusive, along the western slopes of the Cascade range from Skamania Co. to Whatcom Co. in Washington and also in south- western and eastern British Columbia. The ranges of the two, however, often overlap above 1500 feet elevation. Between 1500-2000 feet, ssp. trifoliata seems to be the more frequent, whereas ssp. unifoliata is the more frequent above 2500 feet. In their area of overlap, intergradient plants are evident.
Variety Jaciniata is narrowly limited to Vancouver Island, and to the San Juan and other adjacent islands of Puget Sound. Plants which closely approximate var. laciniata occur sporadically with mixed populations of ssp. trifoliata and unifoliata on the mainland. Variety Jaciniata appears always to be sympatric with var. trifoliata.
CyTOLOGY AND HypripizATION. All three taxa were found to have a gametic chromosome number of 7.
Controlled pollinations gave evidence that all were self-compatible, although none was spontaneously self-pollinating. All possible crosses resulted in fruits in percentages ranging from 40-65% (in 1963) or 45—88% (in 1964). Reciprocal crosses between var. érifoliata and var. laciniata yielded especially high percentages of fruit set (Kern, 1964).
152
1966 | KERN: TIARELLA 153
No differences in the embryo and endosperm, or in seed-size, could be detected between seeds collected in the field and those produced by arti- ficial pollinations. However, neither the seed produced from artificial pollinations nor that collected in the field germinated, possibly because. it was stored for five months before planting. Numerous seedlings were observed in the field, so apparently germination readily occurs in the natural populations.
MORPHOLOGICAL STUDY OF THE LEAVES. Local population samples of 30-100 mature plants were taken from fifteen different locations in the Northwest. The plants were pressed in the usual manner but were left unmounted. The collections were used in the following ways: 1, for a series of simple leaf measurements; 2, for the portrayal of variability within a single population, 10-15 extreme and intermediate forms from each population were photographed; 3, for the construction of polygonal graphs showing intra- and inter-populational variation of leaf pattern; and 4, for the construction of a bar graph designed to detect hybridiza- tion between the extreme forms (Anderson, 1941).
1. Leaf-sinus measurements. In order to obtain a precise assessment of morphological differences, a series of linear (fig. 1) and vein-angled measurements was made on the three largest basal leaves from each dried plant. The method of measuring the depth of lobation was an in- direct one of measuring the length of the line forming a right angle with the midrib and extending straightway to the base of the sinus be- tween the lobes. Approximately fifty plants per population (150 leaves) were measured.
The measurements of depth of lobation (numbers 10, 11, and 12 in fig. 1) were the only ones that might be interpreted to indicate a dis- continuity between taxa. Measurement number 12, the depth of the sinus which splits the single leaf into three leaflets with distinct petiolules, readily separates the unifoliate from the trifoliolate plants even though there is a continuous gradation in the depth of the sinus (Kern, 1964), No true discontinuity exists in the increasing depth of the sinuses be- tween the lobes of the unifoliate leaves and the spaces which these sinuses ultimately become in those plants with trifoliolately compound leaves. The critical mean leaf-sinus measurements, nos. 10-12 (fig. 1), are given in Table 1. All measurements are given in centimeters.
2. Variability within populations. Although unmixed populations of either ssp. trifoliata or unifoliata showed some variation from leaf to leaf, any one of the leaves in each could be readily identified as belong- ing to one or the other of these two taxa.
Populations with mixtures of plants with unifoliate, trifoliolate, and laciniately incised leaves presented a more complex problem. Nine mass collections of 30-100 plants were made along the western slopes of the Cascade Range from Mt. Rainier National Park north to Mt. Baker National Forest. Four of these will be reviewed below (Kern, 1964).
154 MADRONO | Voi. 18
Fic. 1. Tiarella leaf showing measurements and enumerations used.
Plants of Denny Creek, about five miles west of Snoqualmie Pass, Washington, were representative of a population where the unifoliate phase was the most frequent. In the mass collection, 24 plants could definitely be called ssp. unifoliata, 12 ssp. trifoliata, 2 var. laciniata, and 13 intermediate between ssp. trifoliata and unifoliata. The plants that were considered intermediate all had some indentation toward the mid- vein. The range was from a slight indentation to a lobation as deep as that seen in plants misidentified as JT. trifoliata, even though the apical and lateral lobes do not have distinct petiolules. Plants with such deeply dissected leaves are comparable to those that Lakela (1937) refers to as T. unifoliata f. trisecta, since they are very deeply dissected toward the midvein, but without distinct petiolules. All plants without distinct petiolules are considered here as unifoliate rather than trifoliolate phases.
There were a few plants in this population that could be called var. laciniata, although they had larger and less-deeply dissected leaves than those characteristic of var. laciniata of the San Juan Islands.
Mixed populations of ssp. unifoliata and ssp. trifoliata which were linked by intergradient forms were found in numerous locations, only three of which are pictured (fig. 2 A-C).
1966] KERN: TIARELLA 155
& fe, DENNY CREEK at Se NOOKSACK RIVER ake
a tet fat te we he Se abe
he Be Se Me Ne. Se ORCAS ISLAND
maga
sd “ te Me
he fe & %
we we &
Je Se : see a te We Ps we we
Fic. 2. Leaf outlines showing the range of variation within the population. The leaves in the upper two corners represent the extreme phases with intergradient
forms between. The bottom two rows represent a portion of the variability within the population: A, Denny Creek, King Co., Wash.; B, Silverton, Snohomish Co., Wash.; C, Nootsack River, Skagit Co., Wash.; D, Orcas Island, Island Co., Wash.
B
The variation in leaf character becomes greatly complicated in the San Juan Island populations. On Whidbey I. and Orcas I., var. laciniata occurs in almost equal numbers with var. trifoliata in the moist wood- land. On Orcas I. (fig. 2D), there are so many intergradient forms that the majority do not fit in either category.
3. Intra- and inter-populational variation of leaf pattern. The varia- tion within a population may be portrayed by the polygonal graph method (Davidson, 1947). As stated by Davidson, the graph “consists of a circle, with as many radii as there are characters to be compared. The characters, measured along each radius, are assigned absolute, rela- tive, or arbitrary values. The characters possessed by each specimen are plotted along each radius, and the points are joined” (fig. 3A). By the overlap, or non-overlap, of the separate polygons on each graph, one can determine continuous or non-continuous variation. This would indicate whether a population contains two or more segregates or merely one variable entity. Eight characters, either absolute or relative, are graphed in fig. 3B—D.
Figure 3B, as indicated by the normal distribution (Davidson, 1947) on all radii, shows very little variability in an unmixed population of subsp. unifoliata. Polygons of a mixed population of var. trifoliata and
156 MADRONO [Vol. 18
Width of lateral tebe Length of lateral lebe 4
Length of lateral lobe
Length 2 =
| , | { LNSy CAP
2 Depth of incision . ef primary sinus |
Width
IL
A Depthixetiilaclsion|iefsapicalilebe. Width of apical lebe
SS, SS S&S
Fic. 3. Polygonal graphs for comparison of patterns of several characters: A, form used for the polygonal graphs, showing the basis for the graphing of eight characters; B, unmixed populations of ssp. wnifoliata; C, mixed population of var. trifoliata and var. laciniata; D, mixed populations of ssp. wnifoliata; ssp. trifoliata, var. trifoliata, and var. laciniata.
var. laciniata from Orcas I. (fig. 3C) show normal variability on all radii, indicating that there are numerous intergradient forms from one extreme (the trifoliolate phase) to the other (the laciniate phase).
A single mixed population from Denny Creek is graphed in fig. 3D. The variability on the radii for depth of primary sinus/width (12/18) and depth of incision of apical lobe/width of apical lobe (11/13) indi- cates the total range of variation within the population. From this and other patterns of continuous variation, it is evident that the mixed popu- lations form one variable entity.
4. Hybrid index. The hybrid index presents the tabular data in a single figure. To each of three leaf measurements an index number was assigned (table 2). The numbers were then totaled to obtain the index value. The bar graph (fig. 4) was constructed with these values plotted against their frequency within the populations.
1966] KERN: TIARELLA 157
RYBRID INDEX VALUES
Fic. 4. Bar graph of hybrid index values for the taxa of Tzarella.
The index value of 4 is typically that of var. Jaciniata, 8 that of var. trifoliata, and 14 that of ssp. unifoliata. It can be seen that three peaks, at 4, 8, and 12—14 occur. There is a distinct decrease of frequencies at index value 11; intermediates at this value are not as numerous as those more closely resembling either the trifoliolate extreme (8) or the uni- foliate extreme (12-14).
No sharp decrease of frequencies between index values 4 and 7 can be noted; the range of variation from var. laciniata to var. trifoliata appears to be continuous.
TAXONOMIC CONSIDERATIONS. As shown by controlled breeding experi- ments and by strong circumstantial field evidence, the taxa of Tiarella are freely interbreeding. The flower and fruit structures of the taxa are indistinguishable. The various and variant attributes of leaf morphology remain as the only bases for assessing discontinuities. From the evidence presented above, no sharply discontinuous variation can be found. There- fore, I consider the western populations of Tiarella to constitute a single polymorphic species—T. trifoliata L.
The partial discontinuity in geographical distribution and morphology is sufficient, however, to justify the recognition of infraspecific elements. The unifoliate and trifoliolate taxa of Tiarella are here considered to be of subspecific rank. Each occupies a major geographical area of the species range, and the two have an area of overlap, at elevations ranging between 2000-4500 feet, where they intergrade more or less completely.
The laciniate-leaved phase, var. Jaciniata, occurs sporadically with mixed populations on the mainland but occurs in approximately equal numbers with the trifoliolate phase in insular populations. It, as found on the mainland, is possibly simply the result of recurrent and sporadic
158 MADRONO [Vol. 18
TABLE 1. LEAF-SINUS MEASUREMENTS FOR THE TAXA OF TIARELLA
—$__—
Measurement Number
No. of 10 i 12 Taxa plants mean range mean range mean range ssp. unifoliata 24 2 13276 1.1 0.5-1.6 1.5 0.8-21 (Denny Cr.) 24 1S) e122 0.9 0.6-1.2 1.3 0.9-1.8 (Silverton) 10 1.8 1.0-2.2 10 0.5-1.3 1.3 0.8-1.6 (Nooksack R.) Intermediate 13 2.2 1.9-3.0 1.0 0.7-1.2 0.4 0.1-0.9 (DC) 9 2 al =2e5 1.1 0.8-1.2 0.4 0.1-0.9 (Sil.) 9 13° 1,1-1.7 0.6 0.6-0.7 0.4 0.2-0.6 (Nook.) var. trifoliata 12 1.0 0.6-2.6 0.6 0.4-0.9 0 (Dic) 15 10 0.6-1.5 0.7 0.4-0.9 0 (Sil.) 36 1.2 0.4-2.4 0.7 0.5-1.2 0 (Nook.) 25 1.3 0.7-1.7 0.5 0.4-0.8 0
var. laciniata 2 0.1 0.3 0 (Dic) 2 0.8 0.4-1.3 0.4 0.3-0.6 0 (Sil.) 2 0.6 0.3-0.8 0.4 0.3-0.5 0) (Nook.) 25 0.1 0.1-0.5 Ol 30-02 6) (O.1.)
recombination of recessive genes. These individuals are of limited occur- rence in a population and apparently cannot establish themselves as a major segment of the species. ,
Therefore, in T. trifoliata L., two subspecies are recognized: ssp. trifoliata, with two varieties, tvifoliata and laciniata (Hooker) Wheelock, and ssp. unifoliata (Hooker) Kern. The appropriate new combination is hereby proposed. 7
SYSTEMATICS. TIARELLA TRIFOLIATA L. Sp. Pl. 406. 1753.
Blondia trifoliata Raf. Fl. Tell. 2:75. 1836. T. rhombifalia Nutt. in Torr. & Gray, Fl. N. Am. 1:588. 1840. (G. Demidoff, “Habitat in Asia boreali”.) According to Hultén (FI. of Alas. and Yukon 5:943. 1945) the actual type was: Steller, Cape St. Elias, Kayak I., Alas.
T. stenopetala Presl, Rel. Haenk. 2:55. 1831. (Haenke, “Nootka- Sund”’.)
T. trifoliata var. laciniata (Hook.) Wheelock, Bull. Torrey Club 23:72. 1896. T. laciniata Hook. Fl. Bor. Am. 1:239. 1832. Petalosteira laciniata Raf. Fl. Tell. 2:74. 1836. (Menzies, ‘North-West coast of America’”’. )
1966] KERN: TIARELLA 159
TABLE 2. HYBRID INDEX VALUES
Character Range Index Number Depth of incision of apical 0.1-0.2 1 lobe (measurement #10) 0.3-0.9 g 1.0-1.6 3 1.7-3.0 4 Depth of incision of lateral 0.1-0.2 1 lobe (measurement #11) 0.3-0.4 2 0.5-0.9 5 1.0-1.7 4 Depth of incision between 0 2 primary and lateral lobes 0.1-0.9 4 (measurement #12) 1.0-2.0 6
T. trifoliata ssp. unifoliata (Hook.) Kern, com. nov. T. unifoliata Hook. Fl. Bor. Am. 1:238. 1832. Petalosteira unifoliata Raf. Fl. Tell. 2:74. 1836. Heuchera californica Kell. Proc. Calif. Acad. 5:53. 1873. T. unifoliata var. procera A. Gray, Bot. Calif. 1:199. 1876. T. trifoliata var. unifoliata Kurtz, Bot. Jahrb. 19:378. 1894. T. californica Rydb. N. Am. FI. 22:118. 1905. (Drummond, “Height of land in the Rocky Moun- tains, near the source of the Columbia, and at Portage River’’)
T. unifoliata f. trisecta Lakela, Am. Jour. Bot. 24:350. 1937. (Butters & Holway 216c, Beaver Valley, Alta.) = ssp. unifoliata.
Perennial, rhizomatous herbs; flowering stems 10—60 cm high; cauline leaves 2—3, short-petioled; basal leaves simple to compound; inflores- cence paniculate; flowers white, sometimes with a pinkish tinge; sepals 5, oblong to triangular, 2-5 mm long, somewhat irregular, the hypathium short-campanulate, '4—'% as long as the lobes, almost free of the ovary; petals 2-5 mm long, subulate, almost indistinguishable from the fila- ments; stamens 10, 2-6 mm long, those opposite the petals the shorter; pistil 1-celled, with two parietal and nearly basal placentae; valves of the capsule unequal, the upper one % as long as the lower; seeds 1.5—2.0 mm, black, shining and smooth, the raphe prominent. The species con- sists of two subspecies and two varieties as follows:
Leaves simple, 4-12 cm broad, 2.5-7.0 cm long (including those plants with leaves dissected to within 0.1 cm of the midvein but without distinct petiolules)
ssp. unifoliata (Hook.) Kern
Leaves compound, 2-12 cm broad, 2.5—7.0 cm long.........000222.e eee ssp. trifoliata
Leaflets laciniately cleft, the lateral pair deeply dissected to within 0.1-0.4 cm of
the midvein, the middle one dissected to within 0.1-0.9 cm of the midvein;
occurring as a sporadic variant with ssp. wnifoliata and ssp. trifoliata and
sympatrically with ssp. trifoliata on Vancouver Island and adjacent islands
Olmbucret SOUNU. <.2 A. ee var. laciniata (Hook.) Wheelock
Leaflets not laciniately cleft, the lateral pair shallowly lobed to lobed within 0.5
cm of the midvein, lobes of the middle leaflet dissected to within 1.0 cm of
the midvein; occurring primarily west of the Cascade Range in Washington,
but ranging from Alaska to Oregon and east to eastern B.C......... var. trifoliata
Department of Botany, University of Washington, Seattle
160 MADRONO [Vol. 18
LITERATURE CITED
ANDERSON, E. 1941. The technique and use of mass collections in plant taxonomy. Ann. Missouri Bot. Gard. 28:287-292.
Davipson, J. F. 1947. The polygonal graph for simultaneous portrayal of several variables in population analysis. Madrofio 9:105-110.
Davis, P. H., and V. H. HEywoop. 1963. Principles of angiosperm taxonomy. Oliver and Boyd, Edinburgh.
Kern, P. 1964. A taxonomic revision of the genus Tiarella. Masters degree thesis (unpublished). Univ. of Washington, Seattle.
LaKEILA, O. 1937. A monograph of the genus Tiarella L. in North America. Am. Jour. Bot. 24:344-351.
NOTES AND NEWS
OCCURRENCE OF THE GENUS EUPHRASIA IN THE PaciFic NorTHWEST.—The genus Euphrasia has not previously been reported from Washington. Now, E. canadensis Townsend has been collected in Parkland, near Tacoma, Pierce Co., Washington, where it is locally abundant in a moist grassy area adjacent to a small swamp one- half mile east of the intersection of state highways 5 and 512 (Ganders 135, WS). Euphrasia canadensis is almost certainly introduced at Parkland, and the fact that another localized species, Viola lanceolata L., occurs with it suggests the possibility of the same mode of introduction. V. lanceolata was introduced in the early twentieth century with importations of cranberry plants from New England and Wisconsin (Schultz, J. H., Madrofio. 8:191-193. 1946), and since E. canadensis occurs in New England this same thing could have occurred. However, if this is the case, it is sur- prising that the plant has not been collected before in Washington, as V. lanceolata was detected in 1936 (Jones, G. N., Madrofio. 4:34-37. 1937). Another record for Euphrasia that should noted is a collection of E. americana Wettst. from Qualicum, Vancouver I., British Columbia (on golf links, Redfern, WS). This species was not included in Vascular Plants of the Pacific Northwest —Frep R. GANDERS, Washing- ton State University, Pullman.
HERBARIUM SUPPLIES.—Bonestell & Co., of San Francisco, for many years a sup- plier of herbarium supplies, has recently been forced to go out of business. The line of herbarium supplies developed by Mr. Cyril A. Cross, in consultation initially with Dr. LeRoy Abrams and Dr. W. L. Jepson, is now available through the Commercial Paper Corp., 300 Brannon St., San Francisco, California 94107. Mr. Cross will handle these supplies at the Commercial Paper Corp. and he has indicated that the same high standards will prevail and that the prices will be the same as they were as of September 15, 1965.
MADRONO
VOLUME 18, NUMBER 6 APRIL, 1966
Contents
VARIETAL TRANSFERS IN CUPRESSUS AND
CHAMAECYPARIS, Elbert L. Litile, Jr. 161 ON THE SPECIFIC DISTINCTION OF ERIOGONUM
NUTANS AND COLLINUM, James L. Reveal 167 JENS C. CLAUSEN | 174
NOTES ON MARINE ALGAE OF WASHINGTON AND SOUTHERN BritisH CoLumMBIA, Richard E. Norris and John West 176
SoME NEw CHROMOSOME NUMBERS IN LINUM, C. Marvin Rogers and Betty D. Harris 179
SCLEROLINON, A NEw GENUS IN THE LINACEAE, C. Marvin Rogers 181
NoTES ON MEXICAN GRASSES V. Two GypPpso- PHILOUS SPECIES OF MUHLENBERGIA, Charlotte G. Reeder and John R. Reeder 185
NoTES AND NEws: NEw PUBLICATIONS 173, 184, 192
A WEST AMERICAN JOURNAL OF BOTANY
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BOARD OF EDITORS
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LymMAN BENsoN, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis HERBERT F, CopELAND, Sacramento College, Sacramento, California JoHN F. Davipson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California Miuprep E. Maturas, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley Marion OwnsBEy, Washington State University, Pullman REeEpD C. Rotiins, Gray Herbarium, Harvard University Ira L. Wiccins, Stanford University, Stanford, California
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VARIETAL TRANSFERS IN CUPRESSUS AND CHAMAECYPARIS
ELBERT L. LITTLE, JR.
Four species of Cupressus L. (cypress) are transferred here as vari- eties of C. arizonica Greene (Arizona cypress), and a recently described species of Chamaecyparis Spach (white-cedar) is reduced to a variety. A review of the New World species of these two genera of Pinaceae has been made while distribution maps were being revised.
CUPRESSUS
In a comprehensive monograph of the New World species of Cupressus based on studies of wild and cultivated plants, Wolf (1948) accepted 15 species (12 in the United States), one with two subspecies. Two species and one subspecies were proposed as new.
Discussing the specific concepts, Wolf (1948) agreed with W. L. Jep- son that the species rest on rather insecure morphological features. His key to species illustrated the scarcity of well-defined qualitative mor- phological characters. Many characters used, such as color of foliage, size of parts, and bark, would not be of specific value in related genera. As Wolf observed, there has been confusion in taxonomic literature of Cupressus and disagreements on ranks and synonymy. And the same author has changed his concepts.
Wolf concluded that this very old group of species once occupied larger areas but in later times, etiher through climatic changes, diseases, or competition, had been reduced to the present minor role in the veg- etation. The range of each species is discontinuous, consisting usually of a few isolated groves or stations which can be counted.
Twisselman (1962) has suggested that C. nevadensis Abrams, of Kern Co., California, is a remnant of a once widespread complex in the arid woodland occurring across the Mojave Desert until late Pliocene. This species has been progressively restricted by increasing aridity and asso- ciated factors and now survives precariously. He noted that C. mohav- ensis Axelrod (Carnegie Inst. Wash. Pub. 516:86. 1939), a Miocene species related to C. arizonica, was found nearby. It seemed tenable that geographic isolation could account for the differences between C. nev- adensis, C. arizonica, C. forbesu, and C. montana.
Earlier, Sargent (1926) described and figured for the United States six species and one variety of Cupressus. My conservative compilation accepted six species without varieties (Little, 1953), to which should be added C. sargenti Jeps., of California, and the five varieties below. The other New World species, C. lusitanica mill. (C. benthamu Endl., C. lindleyi Klotzch), is native from Mexico to Honduras.
Maprono, Vol. 18, No. 6, pp. 161-192. May 10, 1966. 161
162 MADRONO [ Vol. 18
CUPRESSUS ARIZONICA Greene. Three species cited also as synonyms (Little, 1953) and another from Baja California are transferred here as varieties. The summary key below to the five varieties is adapted from Wolf (1948). The combined range extends from Chisos Mountains, Trans-Pecos Texas, to central Arizona and southern California, and also to northern Mexico (Baja California and from Sonora to Tamauli- pas, south to San Luis Potosi, Zacatecas, and Durango).
CUPRESSUS ARIZONICA var. ARIZONICA. C. arizonica Greene, Bull. Torrey Club 9:64. 1882. C. arizonica var. bonita Lemm., Handb. W.Amer. Cone-Bearers. Ed. 3. 76. 1895. C. arizonica var. bonito Lemm., Sierra Club Bull. 4:118. 1902. C. benthamu Endl. var. arizonica (Greene) Mast., Jour. Linn. Soc. Bot. 31:340. 1896. C. arizonica f. glomerata Martinez, Anal. Inst. Biol. Mexico 18:129. 1947. C. arizonica f. minor Martinez, Anal. Inst. Biol. Mexico 18:133. 1947.
The type of C. arizonica was collected by Edward L. Greene “‘on the mountains back of Clifton,’ Greenlee Co., Arizona (isotype, NA!). Lemmon (1895) distinguished also var. bonita, with “trunk and limbs not the least decorticated,” abundant and of large size in Bonita Can- yon, Chiricahua Mountains, now in Chiricahua National Monument. This variety was afterwards united with C. glabra Sudw. but was re- duced to synonymy of C. arizonica by Kearney and Peebles (1942). Variety arizonica is native from Chisos Mountains, Texas, to south- eastern Arizona and in northern Mexico.
In his monograph of the Mexican species, Martinez (1947) named two forms as new, C. arizonica {. glomerata and f. minor.
CUPRESSUS ARIZONICA var. glabra (Sudw.) Little, comb. nov. C. glabra Sudw., Am. Forestry 16:88, fig. 1910 Feb.
This variety is native in central Arizona from Oak Creek Canyon, Coconino Co., to Mazatzal Mountains, Gila Co. George B. Sudworth (1910), dendrologist of the Forest Service and predecessor agencies of the U. S. Department of Agriculture from 1886 until his death in 1927, published the original description with a photograph of twig and cones but without citation of specimens. In December 1909, he studied this tree at Verde River Canyon in Yavapai Co., Arizona. The specific epithet presumably referred to the “very thin smooth, dark purple-red bark.” Also, the “perfectly smooth bark” was contrasted with the rough bark of C. arizonica, ‘characterized by a rather thin, somewhat stringy, anastomosely furrowed bark of dark red-brown color.” Earlier, Toumey (1895) had observed at Natural Bridge the trees with different reddish, peeling bark.
Recently I have found Sudworth’s type specimen among several black cardboard boxes of bulky unmounted conifers in storage. One lot, con- sidered the holotype, consists of several unpressed twigs with mature cones and male strobili and two seedlings, mostly bearing tags ‘“‘Cupres- sus glabra GBS” and negative numbers. A note contains the name Cupressus n. sp., changed to Cupressus glabra, “Verde Canyon, Pres-
1966 | LITTLE: CUPRESSUS AND CHAMAECYPARIS 163
cott N. Forest 12/29/09 GBS.” The exact locality, apparently added later and shown also on specially prepared maps in the map file, was “From Tinsley’s Ranch approx. T. 11 N., R. 6 E., Sec. 17 (NW % of NW %).’ A second lot of undated specimens with two pieces of reddish brown smoothish thin bark is labelled ‘‘from W. A. Tinsley, Prescott Natl. Forest, Verde River Canyon (Type locality). Another lot, with two pieces of slightly rough bark, from Natural Bridge, Arizona, re- ceived Sept. 17, 1910, had the name C. glabra changed to C. arizonica. The first lot, the one element used by the author, is the holotype. It has been divided and distributed as indicated by the citation: George B. Sudworth Dec. 29, 1909, William A. Tinsley’s Ranch, Sec. 17, T. 11 N., R. 6 E., Verde Canyon, 16 mi. SE. of Camp Verde, Yavapai Co., Ari- zona (US—holotypus, A, ARIZ, RSA, USFS). The type locality is now within the Tonto National Forest.
Sudworth (1915) described, illustrated, and mapped both C. glabra and C. arizonica. However, the maps (maps 1, 2) showed both occupying nearly the same ranges in Arizona. His plate 3, a line drawing copied from a photograph, illustrated two large trunks touching at the base and with this legend: ‘Cupressus arizonica (right) with characteristic fur- rowed bark, Cupressus glabra (left) with smooth scaly bark.” It was reported that a forester had submitted this photograph to show that only one species was represented. As now interpreted, the two taxonomic groups native in Arizona have separate ranges, and both trunks would be within the variation of var. glabra, which sometimes develops fur- rowed bark on very large trunks.
While working in Arizona from 1935 to 1941, I learned of the un- certainty among local foresters whether there were one or two species of Cupressus. I reduced C. glabra to synonymy (Little, 1944), stating that the two were separated only by bark characters and different range. Wolf (1948) noted that the foliage of C. glabra was usually much grayer. Variety glabra is more ornamental than the typical variety be- cause of its smoothish bark and grayish foliage and is much commoner in cultivation under the names C. arizonica and var. bonita.
Two varietal epithets were given to horticultural varieties, in Europe, but their correlation with wild trees is uncertain. Cupressus arizonica var. compacta C. K. Schneid. (Silva Tarouca, Uns. Freil.-Nadelh. 174. 1913) was a low roundish bush. Cupressus arizonica glauca Woodall (Garden 80:16. 1916) was a juvenile form of silvery gray foliage. Both were accepted as varieties by Rehder (Bailey, 1923; 1933; Rehder, 1940) but were reduced to forms later by Rehder (1949). Kriissmann (1960) reduced them to horticultural varieties (cultivars).
Bannan (1954) found C. arizonica and C. glabra to have similar wood structure but did not study the other three species of this group.
CUPRESSUS ARIZONICA var. montana (Wiggins) Little, comb. nov. C. montana Wiggins, Contr. Dudley Herb. 1:161. 1933.
An isotype of this variety from Sierra San Pedro Martir, northern
164 MADRONO [Vol. 18
Baja California, Wiggins & Demaree 4990 (US), has been examined. Wolf (1948) noted a relationship with C. arizonica.
CUPRESSUS ARIZONICA var. nevadensis (Abrams) Little, comb. nov. C. nevadensis Abrams, Torreya 19:92. 1919. C. macnabiana nevadensis (Abrams) Abrams, Illus. Fl. Pacif. States 1:73. 1923.
Wolf (1948) observed that C. nevadensis has its closest affinities with C. arizonica. This variety is known from a few groves in Piute Moun- tains, Kern Co., California.
CUPRESSUS ARIZONICA var. stephensonii (Wolf) Little, comb. nov. C. stephensonii Wolf, Aliso 1:125. 1948.
This variety, the rarest of the named cypresses, is known from a single grove in Cuyamaca Mountains, Cleveland National Forest, San Diego Co., California. It was noted there by the late Ranger J. Bert Stephen- son, whose specimen is deposited in the Forest Service Herbarium. Ac- cording to Wolf (1948), C. stephensonii is somewhat intermediate be- tween C. glabra, which it resembles in its smooth, exfoliating bark, and C. arizonica, in its much less glandular leaves.
Trunks of mature trees with rough, furrowed, gray or dark brown bark. Cones opening at maturity; leaves with active gland; San Pedro Martire Baja” California... 222 eee ee nee var. montana Cones usually remaining closed for several seasons. Leaves with inactive or occasionally active gland or sometimes none; southeastern Arizona to Chisos Mountains, Texas,
and northern Wie os yee eee var. arizonica Leaves with conspicuous, active gland; Piute Mountains, Kern Cos) Calihopnia cre eee ete leg ee cea var. nevadensis
Trunks of mature trees with smooth, cherry-red or mahogany-brown bark, peeling in thin, curling plates. Leaves with comparatively inactive gland; seeds mostly more than 5 mm. long, not glaucous; Cuyamaca Mountains, San Diego
Co; Califormla:.:4..42:10 sneer ee var. stephensonii Leaves with very active gland; seeds mostly 4-5 mm. long, glaucous; central Anzona kA ee ee ee var. glauca CHAMAECYPARIS
Three species of Chamaecyparis Spach have long been accepted for North America, two western and one eastern. Chamaecyparis nootkat- ensis (D. Don) Spach is native in the Pacific Coast Region from south- eastern Alaska to northwestern California. Chamaecyparis lawsoniana (A. Murr.) Parl. has a limited range in southwestern Oregon and north- western California.
Chamaecyparis thyoides (L.) B. S. P. is found in the Atlantic and Gulf Coastal Plains, especially in swamps, from southern Maine to northern Florida and west to southern Mississippi. This Linnaean species without named geographical varieties is the lectotype of the genus (Britton & Shafer, 1908).
1966] LITTLE: CUPRESSUS AND CHAMAECYPARIS 165
A segregate species proposed in 1962 is reduced here to a variety. Both geographic varieties summarized below, have separate, discontin- uous ranges. A few horticultural varieties named in Europe apparently were derived from the typical, accessible, northern variety, which was introduced into Europe first. Rehder (1940; 1949) reduced four of these to forms.
CHAMAECYPARIS THYOIDES (L.) B.S. P., Presl. Cat. Anth. Pter. N. Y. 71. 1888, var. THYOIDES. Cupressus thyoides L., Sp. Pl. 1003. 1753.
Range. Atlantic Coastal Plain from Waldo Co. in southern Maine to Lexington Co. in central South Carolina and extinct along the Savanna River near Augusta, Georgia.
CHAMAECYPARIS THYOIDES var. henryae (Li) Little, Sorin nov. C. henrvae Li, Morris Arb. Bull. 13:43. 1962.
Range. Gulf Coastal Plain from Liberty Co. in northwestern Florida west through southern Alabama to Pearl River Co. in southern Missis- sippi. Also local in Taylor, Talbot, and Marion counties in western Georgia and in Putnam and Marion counties in northeastern Florida.
Li (1962) has published a detailed description of this segregate. The differences from the northern type include bark, color of foliage, size and position of leaves, glands of leaves, lower surface of juvenile leaves, color of male strobili, size, number of scales, and color of cones, size of seeds and wings, and time and appearance of male strobili. Li noted that Bartram (1791) first observed in 1777 that the trees in southern Ala- bama differed a little from those of New Jersey and Pennsylvania.
Like the differences between some geographically separated popula- tions of Cupressus, these characters seem to me to represent two geo- graphical varieties of one species. The original description suggested somewhat closer relationship of the new species with the western species C. nootkatensis than with C. thyoides. Ward (1963) remarked that “such speculations merit further investigation.” Chamaecyparis noot- katensis differs from typical C. thyoides and its segregate in having stout 4-angled twigs, closely appressed larger obtuse leaves, usually without glands, cones almost twice the size and scales with more prominent points, and larger seeds with broader wings. In sterile specimens the stout foliage sprays of C. nootkatensis are readily distinguished from the fine foliage sprays of both typical C. thyoides and its segreate.
Mrs. Mary Gibson Henry, President of the Henry Foundation for Botanical Research, Gladwyne, Pennsylvania, has made intensive field studies of the var. henryae. When first collecting specimens in 1939, she noted the differences from the northern trees. She has made several special field trips to make detailed collections throughout the geographic range.
I am indebted to Mrs. Henry for a duplicate set of her extensive col- lections of 1962-63 totaling 56 numbers (NA) of var. Henryae including an isotype (IM. G. Henry 7108 (23), NA, US, Perdido River, Barrineau Park, Escambia Co., Florida) and collections from eight counties of
166 MADRONO [Vol. 18
Florida, three counties of Georgia, four counties of Alabama, and two counties of Mississippi.
The trees are found in privately-owned forests along streams and in swamps. Li reported that large trees are now extremely rare and that extensive logging has greatly reduced the number of stands and large specimens. However, small trees are fairly plentiful along streams and in less accessible swamps, accordling to Mrs. Henry’s labels.
Specimens from two isolated stations in northeastern Florida are re- ferred to var. henryae, having mostly five cone scales instead of six and bearing male strobili in the autumn instead of the spring. Ward (1963) has reported on the well-known station four miles south of Interlachen, Putnam Co., and the second stand about 30 miles east of Ocala on Juniper Creek along the boundary of Marion and Lake counties. The name Juniper Creek apparently refers to these trees, known locally as junipers. This stand at the southeastern limit of the species is within Ocala National Forest and is protected from cutting. It was brought to the attention of Ward in 1962 by Paul E. Bielling of the Florida Forest Service and formerly of the U. S. Forest Service. Bielling sent me speci- mens collected in Dec. 1963 (NA, US, USFS). Mrs. Henry collected specimens at both localities also.
Forest Service, U.S. Department of Agriculture, Washington, D. C.
LITERATURE CITED
Bartey, L. H. 1923. The cultivated evergreens. Macmillan, New York.
. 1933. The cultivated conifers in North America. Macmillan, New York.
Bannan, M. W. 1954. The wood structure of some Arizonan and Californian species of Cupressus. Canad. Jour. Bot. 32:285-307.
BartraM, W. 1791. Travels through North and South Carolina, Georgia, East and West Florida, the Cherokee country, the extensive territory of the Muscogulges or Creek confederacy, and the country of the Chactaws. James and Johnson, Philadelphia.
Britton, N. L., and J. A. SHAFER. 1908. North American trees. Henry Holt, New York.
Kearney, T. H. and R. H. Peesres. 1942. Flowering plants and ferns of Arizona U.S. Dept. Agr. Misc. Publ. 423.
KrussMan, G. 1960. Die Nadelgeholze. Ed. 2. Paul Parey, Berlin and Hamburg.
Lemmon, J. G. 1895. Handbook of West-American cone-bearers. Ed. 3. Oakland. Calif.
Li, H. 1962. A new species of Chamaecyparis. Morris Arb. Bull. 13:43-46.
Littte, E. L., Jr. 1944. Notes on nomenclature in Pinaceae. Am. Jour. Bot. 31:587- 596.
. 1953. Check list of native and naturalized trees of the United States (in- cluding Alaska). U.S. Dept. Agr. Handb. 41.
Martinez, M. 1947. Los Cupressus de Mexico. Anal. Inst. Biol. Mexico 18:71-149.
Renper, A. 1940. Manual of cultivated trees and shrubs hardy in North America. Ed. 2. Macmillan, New York.
1966 | REVEAL: ERIOGONUM 167
. 1949. Bibliography of cultivated trees and shrubs. Arnold Arboretum of Harvard University, Jamaica Plain, Mass.
SARGENT, C. S. 1926. Manual of the trees of North America (exclusive of Mexico). Ed. 2. Houghton Mifflin, Boston and New York.
SupworTH, G. B. 1910. A new cypress for Arizona. Am. Forestry 16:88-90.
. 1915. The cypress and juniper trees of the Rocky Mountain region. U. S. Dept. Agr. Bull. 207.
ToumEy, J. W. 1895. Notes on the tree flora of the Chiricahua Mountains—I!. Garden Forest 8:22-23.
TWISSELMANN, E. C. 1962. The Piute cypress. Leafl. West. Bot. 9:248-253.
Warp, D. B. 1963. Southeastern limit of Chamaecyparis thyoides. Rhodora 65:359- 363.
Wotr, C. B. 1948. Taxonomic and distributional studies of the New World cypresses. Aliso 1:1-250.
ON THE SPECIFIC DISTINCTION OF ERIOGONUM NUTANS AND COLLINUM
JAMES L. REVEAL
While preparing a taxonomic revision of the Eriogonum deflexum Torr. complex, it became necessary to investigate the relationship of E. nutans Torr. & Gray to the complex. From the morphological stand- point, E. nutans somewhat resembles EF. parryi A. Gray and E. brachy- podum Torr. & Gray because of its glandular pubescence, and is similar to EL. hookeri S. Wats. in its wide campanulate involucre and the general plant stature of the immature specimens of FE. hookeri. However, E. nutans lacks the cordate base of the outer perianth segments typically found in these species, and is an octoploid rather than tetraploid like all of the members of the E. deflexum complex. The historical and morpho- logical confusion, however, has been with the unrelated taxon, E. col- linum S. Stokes ex M. E. Jones.
The type of E. nutans was collected by Beckwith from Mud Lake Valley, Washoe County, Nevada in 1854, and was originally described by Torrey and Gray (1855) as E. cernuum Nutt. var. purpurascens. In their revision of the genus (1870), they elevated the taxon to the species rank.
The inclusion of E. nutans into the California flora has been based on Watson’s treatment of Eriogonum in Botany of California (1880), at which time he erronously cited the Beckwith collection as coming from Lassen County. Jepson (1914), Abrams (1944), and Munz (1959) have all considered this species to be in California on the basis of this error. A recent review of herbarium material has revealed that E. nutans is not in California.
E. collinum was described in 1903 from a series of collections made by Stokes, Jones, and Hillman from the vicinity of Reno, Nevada. Until
168 MADRONO [Vol. 18
now, four taxa described by M. Gandoger (1906) have been thought, by this writer and others, to be referrable to E. collinum: E. praebens Gand., E. praebens var. divaricatum Gand., E. reniforme Torr. & Frem. var. asarifolium Gand., and E. thurberi Torr. var. acutangulum Gand. Jep- son placed the California collection of EZ. praebens under the southern California and adjacent Nevada species, E. reniforme Torr. & Frem., without comment. Tidestrom (1925), possibly following Jepson’s treat- ment, placed E. praebens under E. reniforme, and questionably referred E. collinum to that taxon. While Abrams did not handle E. praebens in any manner, Munz modified the problem slightly by placing the Gan- doger species under E. nutans. When Stokes monographed the genus (1936), she referred EL. praebens and the var. divaricatum to her species, but considered EL. reniforme var. asarifolium as a synonym of E. reni- forme, and made a new combination, FE. cernuum Nutt. ssp. acutangulum for the last Gandoger taxon. The main reason for this assumption is that all of the Hillman specimens deposited at Reno, which bear the same location data as that given by Gandoger, are E. collinum, but the holo- types at Lyon, France, are not that species. Evriogonum praebens and var. divaricatum are E. baileyi S. Wats. var. tomentosum S. Wats., var. asarifolium is E. pusillum Torr. & Gray, and var. acutangulum is E. maculatum Heller. One Gandoger species not previously associated with EL. collinum is E. restioides. The description was based mainly upon two plants of E. baileyi, but the leaf description is that of E. collinum, the left hand specimen on this mixed sheet.
The relationship of E. collinum to the other members of the Section Pedunculata Benth. in DC. is still obscure. The closest species to E. collinum, at least from a morphological and cytological standpoint, seems to be a taxon Stokes described as EF. demissum S. Stokes. However this taxon, by our present subgeneric concept, is a member of the subgenus Oregonium S. Wats., rather than Ganysma S. Wats., and thus more work is necessary before the relationship of either taxa can be established.
During the summer of 1964, Noel H. Holmgren of the New York Bo- tanical Garden and this writer made field observations on the two species, and obtained cytological material. Buds were collected from living speci- mens, and placed in Newcomer’s Solution (1953). Anther smears were made using the iron-acetocarmine techniques, and counts were taken from temporary mounts of Telophase IJ. A tabular arrangement of the chromosome numbers and citation of specimens follows. All collections were made by N. H. Holmgren and J. L. Reveal.
E. collinum S$. Stokes ex M. E. Jones. n = 18. Nevada, Humboldt Co., Leonard Cr., Pine Forest Range, 887; Correl Cr., Pine Forest Range, 1347; California, Lassen Co., Ravendale, 1723. Counted by J. L. Reveal.
E. nutans Torr. & Gray var. nutans. n = 40. Nevada, Humboldt Co., Quinn River Crossing, 1256. Counted by J. L. Reveal.
E. nutans var. glabratum Reveal. n = 40. Nevada, Elko Co., Deeth, 1037; Halleck, 1468. Counted by J. L. Reveal.
1966] REVEAL: ERIOGONUM 169
The results of this study are given in the following key and synopsis. To the curators of herbaria that were visited, or who have sent loan material for this study, the author is grateful.
KEY
Peduncles curving upward, 1-3 (—5) cm long; involucres turbinate, greenish-yellow ; perianth white with a yellowish cast, or yellow, strongly pustulose at the base, or rarely hirsutulous, the calyx-segments similar; achene glabrous; southwestern Idaho, northwestern Nevada, and adjacent northeastern California. E. collinum
Peduncles curving downward, 3-10 mm long; involucres campanulate, reddish or ashy-gray; perianth white with a reddish cast, becoming red, or yellow, weakly pustulose or glabrous at the base, the calyx-segments dissimilar; achene pubes- cent; west-central Nevada through central and northern Nevada to southeast- eEneOreson, ang, cemonal Witan, costes 2 yee ees eee E. nutans
Involucres and peduncles glandular, reddish; range of the species. E. nutans var. nutans
Involucres and peduncles glabrous, ashy-gray; Elko Co., Nevada. E. nutans var. glabratum
ERIOGONUM COLLINUM S. Stokes ex M. E. Jones, Contr. West. Bot. 11:15. 1903. Type: S. Stokes s.n., Reno, Washoe Co., Nevada, June 19, 1900. (UC, isotypes NY, US). £. restioides Gand., Bull. Soc. Bot. Belg. 42:199. 1906. Type: Reno, Hillman s.n. (LY), pro parte. E. nutans of Watson, Bot. Calif. 2:23. 1880; Jepson, Fl. Calif. 1:409. 1914; Abrams, Ill. Fl. Pac. States 2:34. 1944; and Munz, Calif. Fl. 343. 1959, as to the name, not as to the type. E. reniforme of Tidestrom, Contr. U. S. Natl. Herb. 25:154. 1925, as to synonymy in part.
Annual; scapes one to several from or slightly above the caudex, (0.5—) 1-5 (-7) dm high, trichotomous at the first node, floccose at the base becoming glabrous at maturity; leaves basal, 1-2.5 (—3) cm long, 1-3 (—3.5) cm wide, round, cordate, elliptic, to obovate, the apex rounded, the base subcordate to reniform, sparsely hirsute to densely white tomentose below, subglabrous to glabrate, or glabrous above, leaf margins plane or crenulate; petiole 1-5 cm long, tomentose, becoming sparsely floccose to glabrous at maturity; bracts (1—) 1.5-3 (-4) mm long, subulate to lanceolate, ternate and connate at the base, glabrous to hispid within and without; branches dichotomous, glabrous, rather open and divaricated, often with an involucral—bearing peduncle in the node; peduncles curving or ascending upwards, the one or two peduncles in the upper forks occasionally straight, stoutish or slender, 1-3 (—5) cm long, glabrous; involucres turbinate (1.5—) 2-3 mm long, (1-) 1.5—2.5 mm wide, glabrous within and without, 5-lobed, the lobes acute, triangular and erect, dividing the tube 4% to ™% its length, bractlets white to yellowish, numerous, linear-oblanceolate, 1-2.5 mm long, spar- sely hirsutulous with pointed marginal cells, pedicels glabrous, 2-4 mm long, 5 to 15 flowered; perianth white with a yellowish cast, to pinkish-
170 MADRONO [Vol. 18
yellow, or yellow, the midrib pink to red-brown, pale green below the union of the outer calyx-segments, 1—2.5 mm long in anthesis, glabrous except for the pustulose or rarely hirsutulous basal portion of the perianth tube, calyx-segments similar, the lobes ovate, lanceolate to spatulate, margins of the segments often crispate, becoming somewhat revolute; stamens glabrous, included, 1—1.5 mm long, anthers greenish to yellow, or reddish; calyx-segments elongated in fruit; achene glabrous, fusiform, 2—2.5 mm long, tapering to a short 3-angled beak. n = 18
Distribution. In the sandy valley bottoms and clayey foothills of the Reno and Carson City area north to Humboldt Co., Nevada, and adja- cent Owyhee Co., Idaho, and eastern California from Sierra Co. north to Lassen Co. Flowering from June to September.
Representative specimens. CALIFORNIA. Lassen Co.: near Secret Valley, Kelly in 1937 (CAS); near Termo, True 912 (CAS); Bull Flat near Honey Lake, True 784 (CAS); between Mapes and Secret Valley, Heller 15208 (CAS, NY, POM, US, UTC) ; between Shaffer Mountain and Warren Peak, Hoover 4645 (CAS, NY, US) ; Viewland, Ripley & Barneby 5758 (CAS, NY); 0.4 mi S of Ravendale, Raven & Solbrig 13293 (CAS); % mi S of road junction to Smoke Creek, along U. S. 395, Bacigalupi & Constance 7136 (CAS); 1 mi S of Ravendale, Cronquist & Holmgren 8478 (CAS, NY, UTC); 4 mi S of Ravendale, VN. Holmgren & Reveal 1723 (ARIZ, BRY, CAS, DS, GH, MO, NY, RENO, RM, RSA, SMU, UC, US, UT, UTC). IpaHo. Owyhee Co.: 13 mi SE of Bruneau, near Hot Creek, Christ & Christ 16721 (NY). Nevapa. Humboldt Co.: Leonard Creek, Pine Forest Range, NV. Holmgren & Reveal 887 (ARIZ, BRY, CAS, DS, GH, KSC, MO, NY, RENO, RM, RSA, SMU, UC, US, UT, UTC); Corral Creek, Pine Forest Range, N. Holmgren & Reveal 1347 (ARIZ, BRY, CAS, DS, GH, KSC, MO, NY, RENO, RM, RSA, SMU; UC, US; UT, UTC). Lyon Co.: 2.3 mi SE of Dayton, Stackhouse 64 (UC). Washoe Co.: Reno, M. E. Jones in 1897 (US); Reno, Heller 9711a (CAS, RENO); Reno, East- wood 14775 (CAS); 2.3 mi NE of Sparks, Raven 14286 (RSA); Peavine foothills, J.T. Howell 38100 (CAS).
ERIOGONUM NUTANS Torr. & Gray, Proc. Am. Acad. 8:181. 1870. Annual; scapes one to several from or slightly above the caudex, 5—30 cm high, trichotomus at the first node, glabrous; leaves basal, 5-20 mm long, 5—25 mm wide, rounded to broadly reniform, the apex rounded, the base somewhat truncate, subcordate, or cordate, densely white tomentose below, floccose to glabrate above; petiole 5-25 mm long, tomentose, occasionally sheathing up the stem for as much as 1 cm; bracts 1-2 mm long, triangular, 2- or 3-parted, glandular or glabrous without, arachnoid pubescent or glabrous within; branches dichotomous, glandular or glabrous, open to diffuse, often with an involucral-bearing peduncle in the nodes; peduncles curving downward, or somewhat cernuous, slender, 3-10 mm long, glandular or glabrous; involucres campanulate, 2-3 mm long, 2—3.5 mm wide, glandular or glabrous within and without, 5-lobed, the lobes rounded, erect, dividing the tube to about /% its length, reddish- brown with a white membraneous margin, or ashy-gray, bractlets 1—1.5 mm long, linear-oblanceolate, minutely hirsutulous with capitate margi- nal cells; pedicels glabrous or sparsely glandular at the base, 2.5-3.5
1966 | REVEAL: ERIOGONUM 171
Fic. 1. Distribution of E. collinum (squares), E. nutans var. nutans (circles), and E. nutans var. glabratum (triangles).
mm long, 5 to 10 flowered; perianth white, rose, deep red, or yellow, 2—3 mm long in anthesis, weakly pustulose or glabrous on the basal portion of the perianth tube, calyx-segments dissimilar, the outer whorl oblong or oval, emarginate at the broad apex, obtuse at the base, the inner whorl narrower and shorter; stamens glabrous, 2-3 mm long, anthers green or red; calyx-segments elongated in fruit; achene pubescent with short appressed brownish hairs, 1.7-2 mm long, the subglobose base tapering to a long 3-angled beak. n = 40.
ERIOGONUM NUTANS Torr. & Gray var. NUTANS. Type: Beckwith, Mud Lake Valley, Washoe Co., Nevada, June 16, 1854, (NY, isotypes GH, photograph of this isotype at DS, UTC, additional isotype at MO). E. cernuum Nutt. var. purpurascens Torr. & Gray, Pacif. Railroad Re- port 2:124. 1855. E. rubiflorum M. E. Jones, Zoe 4:281. 1895. Type: Dugway, Tooele Co., Utah, May 28, 1891, M. E. Jones sn. (POM,
172 MADRONO [Vol. 18
isotypes, A, ARIZ, DS, MO, NY, UC, US). E. nutans var. brevipedicel- latum S. Stokes, Gen. Eriog. 43. 1936. Type: 30 mi W of Eureka, Eureka Co., Nevada, August 25, 1931, J. T. Howell 7974 (CAS, isotype GH). E. deflexum Torr. in Ives ssp. ultrum S. Stokes, Gen. Eriog. 45. 1936.
Type: Sevier Valley, Sevier Co., Utah, June 18, 1933, Eastwood & Howell 623 (CAS).
Peduncles and involucres glandular, reddish; plants 5—20 cm high: range of the species.
Distribution. In the sandy valley bottoms and lower foothills of west- ern Nevada eastward through central and northern Nevada into south- eastern Oregon, and central Utah (fig. 1). Flowering June to September.
Representative specimens. NEVADA without definite location: desert E of walker Lake, Bryan in 1859 (MO). Churchill Co.: Lahontan, Headley 8 (RENO); Burnt Cabin Canyon, Beach 1005 (UC); W of Eastgate, Ripley & Barneby 5940 (CAS, NY). Elko Co.: Sprucemont, M. E. Jones in 1896 (A, UC); Cave Creek, Ruby Valley, Mason 4719 (UC) ; 6 mi NE of Elko, Holmgren 1103 (MO, OKL, UC, UTC). Esmeralda Co.: Goldfield, Shockley in 1881 (RENO). Eureka Co.: 12 mi from Cor- tez, Goodner & Henning 963b (RENO) ; 2 mi N of Beowawe, Holmgren 1035 (OKL, UC, UTC). Humboldt Co.: W of Valmy, Ripley & Barneby 5598 (NY); 3 mi E of Quinn River Crossing, VN. Holmgren & Reveal 1256 (ARIZ, BRY, CAS, DS, GH, KSC, MO, NY, RENO, RM, RSA, SMU, UC, US, UT, UTC). Lander Co.: Battle Mountain, Kennedy 3084 (DS); Battle Mountain, A. E. Hitchcock 638 (US); 16 mi E of Battle Mountain, Eastwood & Howell 157 (CAS); Kingston Canyon, Goodner & Henning 222 (RENO); 25 mi N of Austin, Ripley & Barneby 6195 (CAS, NY). Mineral Co.: Candelaria, Shockley in 1881 (GH); Candelaria, Shock- ley 293 (DS, JEPS, GH, NY, UC). Nye Co.: Currant, Bentley in 1916 (DS, MO, NY, RM, US); North Twin River, Linsdale & Linsdale 992 (CAS); 1.3 mi N of Belmont, NV. Holmgren & Reveal 1542 (UTC). Pershing Co.: Unionville Valley, Watson 1034 (GH, NY, US). White Pine Co.: 30 mi W of Ely, Delameter in 1947 (DS). Orecon. Harney Co.: White Horse Ranch, Peck 25637 (CAS, NY). UTaH. Beaver Co.: Milford, M. E. Jones 1793 (A, CAS, DS, MO, NY, OKL, RM, UC, UTC). Carbon Co.: E of Wellington, Ripley & Barneby 8640 (CAS, NY). Millard Co.: Tule Valley, Maguire & Becraft 2554 (UTC). Piute Co.: Marysvale, M. E. Jones 5338ab (US). Sevier Co.: Sevier Valley, near Glenwood, Ripley & Barneby 4760 (CAS, NY).
ERIOGONUM NUTANS Torr. & Gray var. glabratum Reveal, var. nov. A var. nutans differt pedunculis et involucris glabris. Peduncles and involucres glabrous, ashy-gray; plant 1-3 dm high; Elko Co., Nevada. Type: 1 mi W of the Deeth turnoff, along the sandy roadside of U.S. 40, Elko Co., Nevada, June 20, and July 14, 1964, NV. H. Holmgren & J. L. Reveal 1037 (UTC, isotypes ARIZ, BRY, CAS, DS, GH, KSC, MO, NY, RENO, RM, RSA, SMU, UC, US, UT, WIS).
Distribution. Known only from the dry valley floor from Wells to Halleck, Elko Co., Nevada (fig. 1). Flowering from June to September.
Representative specimens. NEVADA. Elko Co.: Halleck, Heller 9015 (NY, RENO, US) ; between Wells and Humboldt Wells, Heller 9194 (NY, RENO, US); Deeth, Heller 10561 (DS, NY, RENO, US); Wells, M. E. Jones 25220 (CAS, MO, UC) ; 3.6 mi W of Wells, Stahmann 8 (BRY); 3 mi W of Deeth, Magwre et al 5833 (UTC); 2 mi W of Halleck, N. Holmgren & Reveal 1468 (ARIZ, BRY, CAS, DS, GH, KSC, MO, NY, RENO, RM, RSA, SMU, UC, US, UT, UTC).
1966] REVEAL: ERIOGONUM 173
Most of the specimens cited above have been referred to EL. cernuum, from which var. glabratum may be separated by its wide campanulate involucre, oblong non-undulated calyx-segments, and pubescent achene.
ACKNOWLEDGMENT
The author wishes to express his appreciation to Arthur H. Holmgren and W. S. Boyle for their many helpful suggestions. The assistance of Madame Antonia Gilles of the Herbiers de la Faculte des Sciences de Lyon in obtaining the Gandoger types is greatfully acknowledged. The collecting trip of N. H. Holmgren and this writer was largely supported by an NSF grant under the direction of Arthur Cronquist and Arthur H. Holmgren.
Intermountain Herbarium, Utah State University, Logan (Present address: Department of Botany, Brigham Young University, Provo, Utah)
LITERATURE CITED
ABRAMS, L. R. 1944. Illustrated Flora of the Pacific States. Vol. II. Stanford Univ.
Press.
Jepson, W. L. 1914. Flora of California. Vol. I, in part. Associated Student Store, Berkeley.
GanpoceEr, M. 1906. Le Genre Eriogonum (Polygonaceae). Bull. Soc. Bot. Belg. 42: 183-200.
Muwz, P. A. 1959. A California Flora. Univ. California Press, Berkeley.
NEwcoMER, E. H. 1953. A new cytological and histological fixing fluid. Science 118: 161.
STOKES, S. G. 1936. The genus Eriogonum, a preliminary study based on geographic distribution. San Francisco.
TiDEsTROM, I. 1925. Flora of Utah and Nevada. Contr. U. S. Natl. Herb. Vol. 25.
Torrey, J. and A. Gray. 1855. Pacific Railroad Report. Vol. II. Washington, D. C.
. 1870. A revision of Eriogoneae. Proc. Am. Acad. 8:145-—200. Watson, S. 1880. Botany of California. Vol. II. John Wilson & Son, Cambridge.
NOTES AND NEWS
Trees of the Panhandle, Golden Gate Park, San Francisco. By ELizABETH McCuintock and VircintA Moore. Miscellaneous Paper No. 1. vi + 58 pp. Cali- fornia Academy of Sciences, Golden Gate Park, San Francisco 94118. 1965. $1.00. This pamphlet describes the 50 kinds of trees to be found in the eight block-long extension or Panhandle of Golden Gate Park. Photographs of most of the trees are included and there is a map of the Panhandle showing the location of each tree. Interesting bits of horticultural, botanical, and historical information are also pre- sented. This booklet should be in the hands of every one interested in the trees of San Francisco.
ene
JENS C. CLAUSEN
We salute Dr. Jens C. Clausen, long-time member of the California Botanical Society. Jens, the son of a Danish farmer and house-builder. developed a strong interest in plants at an early age. An uncle introduced him to the flora of Denmark at the age of 9, and through extensive read- ing he absorbed much of the classic botanical literature while still a teen-ager. Jens’ early interests also included geology and paleoecology, his curiousity in these fields having been aroused by the prominent gla- cial features of his native countryside.
His education was unorthodox as judged by current practices. He was taught at home until the age of 8, and by the time he was 13 had at- tended two upper grades of formal schooling. Later he studied at a private secondary school, but at 14 his education was interrupted by a stint at farming although his evenings were devoted to study. At 22 he entered the University of Copenhagen after passing a comprehensive entrance examination. While attending university he supported himself by teaching part-time at a private secondary school. His education was again interrupted during World War I by two years of military service in the Danish army. He received his Ph.D. degree from the University of Copenhagen in 1926 at the age of 34. During both his undergraduate and graduate years he was closely associated with Professor QMjvind Winge with whom he worked intensively on the cytogenetic exploration of various plant groups.
Dr. Clausen’s thesis investigations on the cytogenetics of selected sec- tions of the genus Viola attracted the attention of Dr. Harvey Monroe Hall of the Carnegie Institution of Washington. In 1931 Dr. Clausen became associated with Hall as a staff member of the Department of Plant Biology of the Carnegie Institution at Stanford.
Dr. Clausen’s contributions as a member of a Carnegie team are too well known to require elaboration. He has received many honors, in- cluding election to the National Academy of Sciences, the Royal Swedish Academy of Sciences, the Royal Danish Academy of Sciences and Let- ters, and to honorary fellowship in the Botanical Society of Edinburgh. In 1949 he was awarded the honorary degree of Doctor of Agronomy at the Royal Agricultural College of Sweden, and in 1961 was knighted by King Frederik [X of Denmark to the Order of Dannebrog. Those who know him think of Jens Clausen as a keen, kindly scientist with un- bounded enthusiasm and apparently limitless energy which he directs unstintingly to the enhancement of knowledge in the plant sciences.
VES)
NOTES ON MARINE ALGAE OF WASHINGTON AND SOUTHERN BRITISH COLUMBIA
RICHARD E. Norris and JOHN WEST
Marine algae on the Pacific Coast of North America often show inter- esting distribution patterns because typically southern species are ab- sent or rarely found in northern regions and northern species usually do not occur in southern waters. In recent years the number of species known only in southern or northern parts of the coast has become smaller, primarily through the records of Scagel (1957) and Dawson (1961). The following list of algae supplements the lists of Dawson and Scagel by recording species that have not previously been collected in Washington or southern British Columbia. In addition, further records for three rare genera, Thuretellopsis, Porphyropsis, and Platysiphonia are provided for this region. All specimens are in the herbarium of the University of Washington and are from the state of Washington unless otherwise indicated.
CHLOROPHYTA
Endophyton ramosum Gardner was collected in thalli of /ridaea sp. in drift on a beach 2 miles W of the Sekiu R., Straits of Juan de Fuca, Clallam Co., Oct. 25, 1964 (Norris 5074). This endophyte has not been collected previously north of southern Oregon.
PHAEOPHYTA
Punctaria chartacea Setchell & Gardner was collected by dredging in approximately 7 fathoms at Dot Rock, E of Decatur I., San Juan Co., June 18, 1964 (Norris 4618). It was also found intertidally at Whiffen Spit, Sooke, Vancouver I., British Columbia, July 10, 1964 (Norris 4747, West 133). This species was previously known only from Sitka, Alaska.
RHODOPHYTA
Porphyropsis coccinea (J. Ag.) Rosenv. was observed by Kylin grow- ing on hydrozoa in the vicinity of Friday Harbor. As far as we can determine, Porphyropsis has not been recorded on the North American west coast since that time. We found the same species on hydrozoa and Rhodymenia dredged from 5—7 fathoms at Hein Bank, SW of San Juan I., June 13, 1964 (Norris 4696 ).
Thuretellopsis peggiana Kylin. Previously known only from the type collection, cystocarpic specimens of this species were dredged in 5-7 fathoms at Salmon and Hein Banks, SW of San Juan I., on June 13, 1964 (Norris 4584, West 76).
Gelidium sinicola Gardner was collected on the reef at Mukkaw Bay, Calallam Co., on Oct. 24, 1964 (Norris 5087). This species has not been reported north of central Oregon before this time.
176
1966] NORRIS AND WEST: MARINE ALGAE 177
Rhodophysema elegans (J. Ag.) Dixon was found commonly in dredged material at Hein Bank, SW of San Juan I., June 13, 1964 (Norris 4695); Salmon Bank, SW of San Juan I., July 16, 1964 (Norris 4792); Partridge Bank, W of Whidbey I., July 27, 1964 (Norris 5010). This species has not been previously recorded north of central California.
Rhodophysema georgiu Batt. was collected intertidally several times on leaves of Phyllospadix and Zostera at Whiffen Spit, Sooke, Vancouver I., British Columbia, July 12, 1964 (Norris 4752); N of False Bay, San Juan I., July 24, 1964 (Norris 4842); Cattle Pt., San Juan I., Aug. 8, 1964 (Norris 4970). The only previous record of this species from the northeastern Pacific is a collection from Cape Arago, Oregon, reported by Gardner (Doty, 1947). Doty was unable to find Gardner’s specimens and the species has not been collected since that time. It is possible that the specimens reported here are a stage in the lif history of Smithora naiadum, but the structure of the thalli seems to be identical with speci- mens from Great Britain.
Dermatolithon pustulatum f{. ascripticium (Foslie) DeToni was col- lected at Mukkaw Bay, Clallam Co., Oct. 24, 1964 (Norris 5082). It was growing on Botryoglossum ruprechtianum. This corralline has been recorded previously no farther north than central Oregon.
Rhodoglossum roseum (Kylin) G. M. Smith was collected intertidally at Deadman Bay, San Juan I., June 25, 1964 (Norris 4680) and N of False Bay, San Juan I., July 24, 1964 (Norris 4828). This species for- merly was known only from central California and Baja California.
Gymnogongrus linearis (Turner) J. Ag. was collected at Glacier Pt. (Point No. Pt.), S. Vancouver I., British Columbia (Norris 5136, West 150), on July 11, 1964. Previous to this time, this species was known only as far north as central Oregon.
Besa papilliformis Setchell was collected by dredging in 5—7 fathoms at Partridge Bank, W of Whidbey I., July 27, 1964 (Norris 5018). This species has been collected only twice before, in San Francisco and on the Monterey Peninsula.
Callithamnion lejolisea Farlow was collected on Calliarthron regen- erans at American Camp Beach, San Juan I., July 9, 1964 (Norris 4858, West 120). Formerly this epiphyte was reported only as far north as central Oregon.
Callithamnion endovagum Setchell & Gardner was collected by dredg- ing at Partridge Bank, W of Whidbey I., July 27, 1964 (Norris 4925), growing in thalli of Opuntiella. Previously, this species was reported from Mexico and southern California.
Spermothamnion snyderae Farlow was collected by dredging in 5—7 fathoms at Salmon Bank, SW of San Juan I., June 13, 1964 (Norris 4586). It was also collected intertidally at Whiffen Spit, Sooke, Van- couver I., British Columbia, July 12, 1964 (Norris 4750), and at Mitchell Bay, San Juan I., July 7, 1964 (West 112). Collected specimens are sterile, but polysporangia are produced in culture.
178 MADRONO [Vol. 18
Ceramium eatonianum (Farlow) DeToni was collected intertidally at Whiffen Spit, Sooke, Vancouver I., British Columbia, July 12, 1964 (Norris 4751, West 134) and at Meadow Point (Golden Gardens) Seattle, Nov. 21, 1964 (Norris 5130, West 220). Formerly this species was recorded as far north as central Oregon.
Platysiphonia clevelandi (Farlow) Papenfuss, previously known in this region only from Whidbey I., was collected by dredging in 5-7 fathoms at Salmon Bank, SW of San Juan I., June 13, 1964 (Norris 4599); by dredging at Smith I., Island Co., July 27, 1964 (Norris 4890 ) ; and by dredging at Hein Bank, SW of San Juan I., July 6, 1964 (Norris 5014). This species was isolated in unialgal culture from 4599.
Memobranoptera multiramosa Gardner was collected by dredging in 5—7 fathoms at Salmon Bank, SW of San Juan I., July 16, 1964 (Norris 4758). This species formerly was recorded as far north as the central Oregon coast.
Myriogramme hollenbergi Kylin has been found in both dredged material and in intertidal collections: Whiffen Spit, Sooke, Vancouver I., British Columbia, July 12, 1964, (Norris 4754, West 140); dredged in 5—7 fathoms, W of Smith I., Island Co., July 27, 1964 (Norris 4591) ; on intertidal rocks, N of False Bay, San Juan I., July 24, 1964 (Norris 4839); dredged at Partridge Bank, W of Whidbey I., July 27, 1964 (Norris 4914). Unialgal cultures were isolated from 4914. This species has not been reported north of Monterey, California.
Hymenena multiloba (J. Ag.) Kylin was collected at Mukkaw Bay, Clallam Co., Oct. 24, 1964 (Norris 4103). Previously this species was reported only as far north as central Oregon.
Cryptopleura lobulifera (J. Ag.) Kylin was collected intertidally at Deadman Bay, San Juan I., June 25, 1964 (Norris 4631). This species has not been recorded farther north than central California.
Polysiphonia brodiaei (Dillwyn) Greville was collected on a wood float at Lloyd’s Boat House, Duwamish Head, West Seattle, Jan. 14. 1965 (West 225, 226, 228). The previous known Pacific range of this well known Atlantic species was from Santa Monica to San Francisco Bay, California.
Department of Botany, University of Washington, Seattle
LITERATURE CITED
Dawson, E. Y. 1961. A guide to the literature and distributions of Pacific benthic algae from Alaska to the Galapagos Islands. Pacific Sci. 15:370-461.
Doty, M. S. 1949. The marine algae of Oregon. Part II. Rhodophyta. Farlowia 3:159-215.
ScaceL, R. F. 1957. An annotated list of the marine algae of British Columbia and northern Washington. Natl. Mus. Canada Bull. 150.
SOME NEW CHROMOSOME NUMBERS IN LINUM
C. MARVIN RoceErs and Betty D. HARRIS
Chromosome complements in the North American species of Linum have been the subject of several papers (Dillman, 1933; Ray, 1944; Raven, 1959; Osborne and Lewis, 1962) and, among the species studied, there is evidence of considerable correlation between chromosome num- ber and taxonomic groupings within the genus based on gross morphol- ogy. It is clear that a continuation of such studies may provide further substantiation and perhaps clarify the relationships among a number of species, the systematic position of which is poorly known. For this reason we have studied the following additional taxa: L. bahamense, L. stria- tum, L. medium var. medium, L. australe, L. aristatum, L. alatum and L. pringlet. We have also substantiated earlier counts of chromosome numbers in L. lewtsit (13171, Catron Co., New Mexico) in which n = 9; L. sulcatum (13191, Newaygo Co., Michigan) in which 2n — 30; and L. medium var. texanum (12517, Long Point, Ontario, and Harris 200, Wayne Co., Michigan) in which n = 18. Of the species for which new counts are presented, some are of particular interest in that they possess unusual chromosome numbers and may point to areas where further systematic and cytogenetic investigations would be profitalbe.
Young flower buds and root tips were fixed in Carnoy’s fluid and stained with either aceto-carmine or aceto-orcein. Specimens of all of the collections examined are in the herbarium of Wayne State University. Collection numbers are those of C. M. Rogers except as otherwise indi- cated. This paper is contribution No. 130 from the Department of Biology, Wayne State University.
Linum bahamense Northrop var. bahamense (12138, Grand Bahama I.). The single collection so far examined has n = 34, unlike any other species in the genus thus far studied. This species, endemic to the Ba- hama Islands, is a member of a widespread North American complex which includes L. arenicola from Florida, as well as L. schiedeanum, L. rupestre and several other species which are predominantly Mexican. Of these only L. schiedeanum (under the name of L. greggii) and L. arenicola have been counted and each has n = 18 (Osborne and Lewis, 1962). Linum bahamense may conceivably be an aneuploid variant of a tetraploid derivative of this series. The various populations of L. ba- hamense, distributed through several of the islands, have been, on the basis of morphological studies, treated as four species (Small, 1907) or as three varieties of one species (Rogers, 1963). A study of them now in progress may provide some data of interest concerning speciation in this island complex.
Linum striatum Walter (12166, Leon Co., Florida). The chromosome number is 2n = 36, which is the same as that of L. virginianum, L.
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180 MADRONO [Vol. 18
medium var. texanum and L. floridanum var. chrysocarpum (Osborne and Lewis, 1962), all of which are members of a well defined eastern North American complex.
Linum medium (Planchon) Britton var. medium (12458, Bruce Penin- sula, Ontario). This northernmost representative of the L. medium- striatum-virginianum group is thus far the only taxon to diverge from n = 18. While an exact count of the chromosomes has not been made, the variety appears to have a diploid number of approximately 72. It may have been derived as an allotetraploid from L. medium var. texanum and L. virginianum, both of which it resembles in a number of features. The morphological distinctions which exist between L. medium var. medium and var. texanum (Rogers, 1963), together with their different chromosome numbers, may indicate that the taxa would be better con- sidered as two separate species.
Linum australe Heller var. australe (13169, Catron Co., New Mexico, and 13176, Coconino Co., Arizona) ; L. aristatum Engelm. (13177, Luna Co., New Mexico, and 13152, Culberson Co., Texas); and L. alatum (Small) Winkler (13131, Refugio Co., Texas). These three taxa, part of the L. rigidum complex, have, like L. rigidum (Dillman, 1933) been found to haven = 15.
Linum pringlet S. Watson (13181, 12 miles west of Durango, Mexico, and 13182, 18 miles west of Durango). On the basis of relatively little material seen, this species appears to be closely related to, but distinct from, L. neomexicanum. It may also be allied to L. kingu of Utah and such Mexican species as L. mexicanum and L. hypericifolium, but the relationship of these plants to each other and to other species in the genus is not clear. Linum pringlei has n = 26. This is the first count for the complex and the study of other species may help in clarifying the significance of the unusual chromosome number.
Department of Biology, Wayne State University, Detroit, Michigan
LITERATURE CITED
Dittman, A. C. 1933. Chromosome numbers in flax (Linum). Science 78:409.
OsBorne, W. P. and W. H. Lewis. 1962. Chromosome numbers of Linum from south- ern United States and Mexico. Sida 1:63—68.
RAVEN, P. 1959. In Documented chromosome numbers of plants. Madrono 15:49.
Ray, C. 1944. Cytological studies on the flax genus (Linum). Am. Jour. Bot. 31:241- 248.
Rocers, C. M. 1963. Yellow flowered species of Linum in eastern North America. Brittonia 15:97-122.
SMALL, J. K. 1907. Linaceae. Jn North American Flora 25:67-87.
SCLEROLINON, A NEW GENUS IN THE LINACEAE
C. MarvIN ROGERS
Linum digynum is a diminutive yellow-flowered annual ranging from east central California north to Washington and Idaho. Since it was originally described, L. digynum has been regularly placed in Linum or, occasionally, along with the other North American yellow-flowered species of flax, in the segregate genus Cathartolinum. At first glance, however, the species appears to resemble equally closely several other western annuals which were first set apart in the genus Linum as the section Hesperolinon (Gray, 1865) and subsequently (Small, 1907) as the genus Hesperolinon. The resemblance of L. digynum to Hesperolinon was noted by Brewer and Watson (1876) and Trelease (1887; 1897) who placed it in the section Hesperolinon. The recent inclusion of the western annual species, exclusive of L. digynum, under the genus Hesperolinon (Sharsmith, 1961) pointed up the need for a reexamina- tion of this species to determine more precisely to which genus it properly belonged.
The plant is similar to many species of Linum in its yellow, un- appendaged petals, attached near the base of the stamen tube, and in its capitate stigmas, while it resembles Hesperolinon in its annual habit, bicarpellate fruit (though this is not common in that genus) and three- angled seeds.
Further examination, however, shows that L. digynum possesses an impressive array of features which are not shared with either of the other genera and that, rather than being a possible link between the two genera, it appears to stand on quite firm ground as a separate genus. Several of these unique features concern the fruit and include its shape, the rugulose surface, the thick, hard walls which have accounted for the selection of Sclerolinon as the new generic name, the false septa which are quite complete and similar to the true septa, and the separation of the fruit into four indehiscent nutlets. In addition, the very short style, comparatively large capitate stigmas, the diminutive stamens with anthers sometimes containing as few as 8-12 pollen grains per anther sac are distinctive, as is the shape and lacerate margin of the sepals and upper floral bracts. The haploid number of chromosomes is eight (Raven, 1959) and though some Old World species of Linum (including L. catharticum, which may be a relatively recent introduction into north- eastern North America) have this number, it has not been found among other North American species of the family. Furthermore, the geo- graphical range of L. digynum is distinct from the ranges of species of Hesperolinon and the yellow-flowered species of Linum.
In that the primitive members of the genus Linum are thought to have five carpels with partially developed false septa and freely dehiscent
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fruit, it is probable, on the basis of some of the characteristics mentioned above, plus such features as annual habit, the absence of staminodia and stipular glands, that L. digynum is one of the most highly specialized species of the family in North America. Its relationship to the other flaxes is not clear, but the features which it possesses in common with Linum and Hes perolinon may indicate that it has been derived from some ancestor along a line leading from the former to the latter, since Hesper- olinon, too, appears to be more highly evolved than Linum. But if the species did originate in this fashion, it has since undergone considerable modification, and warrants its segregation into the monotypic genus Sclerolinon.
Sclerolinon Rogers, gen. nov. Annuum, glabrum; foliis oppositis ob- longis; glandulis stipularibus nullis; sepalis oblongo-lanceolatis obtusis, margine lacerato-denticulatis glandulosisque; petalis flavis; dentibus inter stamina nullis; stylis 2 liberis; stigmatibus capitatis; capsula quadrilocellata. There is but one known species.
Sclerolinon digynum (Gray) Rogers, comb. nov. Linum digynum Gray, Proc. Am. Acad. 7:334. 1868. Cathartolinum digynum (Gray) Small, N. Am. FI. 25:78. 1907.
Annual, glabrous herb (6—) 13-25 (—42) cm. tall; stems terete or some- what striate above, simple or occasionally branched below, few-branched in the inflorescence; leaves opposite throughout or alternate above, 1-3 nerved, lanceolate to oblanceolate or elliptic, acute or obtuse, (6—) 9-17 (—21) mm long, (1.6—)2.9-5.3(—8) mm wide, entire; stipular glands none; floral bracts narrowly lanceolate with few gland-tipped lacera- tions; sepals 5, unequal, the outer triangular-ovate to panduriform, ob- tuse, irregularly few-lacerate along the margin mostly below the middle, the teeth gland-tipped, outer sepals (1.7—)2.3-2.8(—4.5) mm long, the inner shorter, lance-ovate, with glandular teeth, but scarcely incised; petals 5 attached near base of stamen tube, obovate or obcordate, yel- low, 3-4 mm long, without a well defined gland or appendages at the base; stamens 5, 1-2 mm long, glabrous, filaments hyaline, united at the somewhat broadened base; staminodia none; anthers elliptic, 0.1—0.3 mm long; fruit pale or occasionally lightly purple-tinged above, bicarpel- late, 4-celled, 4-seeded, thick-walled, pyriform, broadly 4-angled, con- spicuously mucronate at the otherwise truncate or retuse, clearly 4- angled summit, surface irregularly rugulose, (1.6—)1.7-1.8(-2) mm long, (1.4—)1.6—1.8(—1.9) mm in diameter, prominently jointed at the pedicel attachment point, ultimately falling intact, or very tardily de- hiscent, at first along the true septa, then along the false septa, and falling as 4 indehiscent 1-seeded nutlets; false septa complete and simi- lar to the true septa; styles separate or barely united at the base, 0.3-0.7 mm long; stigmas capitate, conspicuous; seeds filling locules, narrowly ovate, 3-angled, rather sharp pointed above, dark brown to nearly black, not glossy (figs. 2-6).
1966] ROGERS: SCLEROLINON 183
J i]
* Sandiego.
| Mexico Fa
Fics. 1-6. Sclerolinon digynum: 1, range; 2, fruit; enclosed in calyx, X 15; 3, fruit, X 15; 4, one segment of the fruit, X 15; 5, seed, X 15; 6, habit k 34.
In dry or more often moist meadows, at 3000-5000 feet, from east central California to southwestern Oregon and east central Washington and adjacent Idaho (fig. 1); flowering mainly from June to August.
Type: Bolander 4900, Yosemite Valley, Mariposa Co., California (US, isotype GH). The US specimen has been selected as the holotype, al- though Bolander’s original collections are at GH. Both specimens were seen and annotated by Gray, but only the US sheet bears the data ‘Mariposa Trail” and what appears to be the original field note “valley, rare,” both of which entered into the original description. In addition
184 MADRONO [Vol. 18
the US specimen is the more ample one, while the GH sheet may be the source of some possible confusion in that it bears three additional col- lections.
Approximately 100 collections were examined, but only a single one from each county is cited.
CaLiForniA. Amador: Hansen 591 (US); Calaveras: Eggleston 9713 (US) ; Fres- no: Raven 6125 (CAS); Madera: Jenney, June 15, 1914 (UC); Mariposa: Bolan- der 4900 (GH, US); Plumas: Hall 9382 (UC, US); Shasta: Eastwood 710 (CAS, NY, POM, US); Siskiyou: Brown 548% (F. NY, PH, RM, US); Tuolumne: Ma- son 2130 (CAS, DS, GH, MICH, NY, POM, UC, US). Ipano. Kootenai: Sandberg, July 1888 (IA, KANU, NY); Nez Perce: Sandberg et al. 311 (GH, NY, POM, US). Orecon. Jackson: Copeland 3472 (CAS, GH, NY, POM, RM, UC, US); Josephine: Peck 7931 (GH); Klamath: Peck 9475 (DS, GH, NY, WTU); Linn: Lloyd, July 11, 1894 (CM, NY); Wasco: Howell, June 1881 (CM, DS, F, KSC, MICH, NY, OSC, PH). Wasuincton. Spokane: Suksdorf 8901 (CAS, DS, NY, RSA, UC, WTU) ; Whitman: Piper 1803 (GH, WIS).
This paper is Contribution No. 134 from the Department of Biology, Wayne State University.
Department of Biology, Wayne State University, Detroit, Michigan
LITERATURE CITED
BrewWER, W. H. and S. Watson. 1876. Botany of California, Vol. 1.
Gray, A. 1865. Characters of some new plants of California and Nevada. Proc. Am. Acad. 6:519-556.
Raven, P. H. 1959. In Documented chromosome numbers of plants. Madronfo 15:49.
SHARSMITH, H. K. 1961. The genus Hesperolinon (Linaceae). Univ. Calif. Publ. Bot. 32:235-314.
SMALL, J. K. 1907. Linaceae. In N. Am. FI. 25:84-86.
TRELEASE, W. 1887. A revision of North American Linaceae. Trans. St. Louis Acad. 5:7-20.
. 1897. Linaceae. Jn A. Gray, Synoptical Fl. N. Am. 1:344-349.
NOTES AND NEWS
Naming the Living World. By THEODORE Savory. xiv + 128 pp. John Wiley & Sons Inc., New York. 1962. $3.95. This book is divided into three parts: Principles of Nomenclature, Codes of Nomenclature, and Practice of Nomenclature. The first chapter, Follies and Foibles, of the last part is delightful reading with such sub- headings as: Light-hearted Names, Foolish Names, Worship of Priority, etc. In addition to providing a source of a good deal of information, the author has man- aged to get the idea across very nicely that “nomenclature is a servant of biology” and not an end in itself.
NOTES ON MEXICAN GRASSES V. TWO GYPSOPHILOUS SPECIES OF MUHLENBERGIA
CHARLOTTE G, REEDER AND JOHN R. REEDER
This research was supported, in part, by Grants G-9070 and GB-2272 from the National Science Foundation. We are indebted to the curators of the following herbaria for making their facilities available: New York Botanical Garden (NY); U. S. National Herbarium (US); Escuela Nacional de Ciencias Bioldgicas, México (ENCB); Instituto de Investi- gacion de Zonas Desérticas, San Luis Potosi, México (SLP). Chester F. Natunewicz, of the Classics Department at Yale, has kindly checked the Latin diagnosis.
In the spring of 1962, Alan A. Beetle, of the University of Wyoming, sent us a number of specimens of Muhlenbergia for determination. Among them was a single plant collected in Nuevo Leon, Mexico, by A. Cuevas, agronomist at the Instituto Tecnologico in Monterrey. Although this grass differed in several respects from any species with which we were familiar, and appeared to be a novelty, we were reluctant to describe it from only a single specimen.
The following autumn, in connection with attendance at the Segundo Congreso Mexicano de Botanica, we were able to visit the locality from which the Cuevas plant had come. Along the edges of a sizeable barranca about 3 miles southeast of Galeana, we found a few plants which were similar to the original collection. These were growing in gypsum soil, and associated with Bouleloua chasei, a marked gypsophile. The area is, in fact, the type locality of this latter species. Searching in a number of other gypsiferous areas in the states of Coahuila, Nuevo Leon, and San Luis Potosi, we encountered our plant in two additional localities. Al- ways it appeared to be relatively rare, and usually was growing on the sides of sink-holes, which are a characteristic feature of these gypsum flats. For an excellent description of the physical characteristics of gyp- sum flats, see Johnston (1941).
In late October of the following year we were again in Mexico. Return- ing to the area near Galeana, we found our “new species” to be rather abundant, and a quite conspicuous element of the grass flora there. It was also apparent that it is by no means confined to sink-holes, but may be scattered about amongst the common and dominant elements of these areas: Bouteloua chasei Swallen and Muhlenbergia villiflora Hitchce.
In one of their papers dealing with the ecology and vegetation of the state of San Luis Potosi, Rzedowski & Rzedowski (1957) discussed the plants new to the State. Unreported for San Luis Potosi at that time was Bouteloua chasei, which the Rzedowskis had found in several gypsiferous areas. They indicated (p. 203): ‘“... . otra graminea a veces dominante es una especie al parecer aun no descrita de Muhlenbergia.” Even though
185
186 MADRONO [Vol. 18
we had never found our “new species” abundant enough in any area to be considered dominant, we reasoned that perhaps it was the grass to which they had referred. In a conversation with Rzedowski, however, we learned that his supposedly unknown Muhlenbergia had been determined later at M/. purpusii Mez. This little known species was originally collected by Purpus (#5011) in 1910, near the Minas de San Rafael in the state of San Luis Potosi. Hitchcock (1935) states that it is “known only from the type collection.”
Through the courtesy of T. R. Soderstrom, of the U. S. National Herb- arium, we were able to examine a type fragment of M. purpusii, along with two other collections (McVaugh et al. 18204 and Rzedowski s. n.). These latter two specimens were gathered in essentially the same locality, some 36 miles south of Matehuala in the state of San Luis Potosi. Later, while visiting the New York Botanical Garden Herbarium, we encoun- tered a complete isotype of this species amongst the Muhlenbergia “in- dets.”’
There seems little doubt that MW. purpusi is the species most closely related to the plant originally collected near Galeana by Ing. Cuevas. The two differ in a number of respects, however, and these appear to be sufficiently great to justify their segregation as quite distinct species.
Muhlenbergia gypsophila C. & J. Reeder, sp. nov. Gramen perenne circiter 35 (raro ad 50) cm altum; culmis caespitosis teretibus erectis, sub nodis et sub inflorescentia dense pubescentibus, ceterum glabris; va- ginis scabris vel apicem versus breviter pubescentibus, quam internodi longioribus; ligula 2-3 (1.5-6) mm longa, membranacea firma, apice truncata et breviter ciliata, marginibus puberulentibus et plusminusve decurrentibus; laminis falcatis 2-10 (raro ad 25) cm longis, involutis, rigidis, supra breviter pubescentibus, subtus plerumque glabris et levibus sed raro cum spiculis paucis praebitis; panicula exserta confertiflorata. plerumque 7.5—-15 cm longa, contracta; ramis arte appressis, ad basin paniculae plerumque 20 mm vel minus longis, in parte superiore brevi- oribus gradatim, rhachi pedicellisque brevibus plusminusve dense pubes- centibus; spiculis stramineis vel interdum leviter purpuratis, glumis subaequalibus acutis 1.5—2 mm longis, obscure 1-nervalibus, puberulen- tibus praecipue apicem versus; lemmate 3-nervali pubescente 3.5—4 mm longa aristata, arista flexuosa 10-15 mm longa ex apice lemmatis bifida; palea lemmati amplitudine paene aequali, 2-nervali, inter nervos pubes- cente, ceterum glabra; antheris pallido-luteis 1.5—2 mm longis; stigmati- bus penicilliformis purpureis; caryopside late fusiforma fusca circiter 2 mm longa; numero chromosomae 2n — 20.
Type: MEXICO: Nuevo Leon, 3 miles east of junction of Linares- Galeana road with Hwy. 85, gypsum flat with widely scattered shrubs
and tree yuccas, occasional in depression along roadside, 6400 ft, 30 Oct 1964, J. R. & C. G. Reeder 3963 (holotype, YU: isotype, US).
1966] REEDER AND REEDER: MUHLENBERGIA 187
™A —<T
: Saas ae SSS
SS
4 hy / (; )
ff a ws ( ieee rey oe x y SANG 1 as
Wy
ap
SRE
Y
WE
A G
Fic. 1. Habit silhouettes and details of various structures. Muhlenbergia pur pusii on the left; M. gypsophila on the right. A, A,, habit, x ca. 1g; B, B,, ligule, x 5; C, Ci, glumes; D, Du, floret; E, E:, stamen; F, Fi, caryopsis in two views (C-F,, xX 10); G, G,, lodicules, X 50; H, H,, metaphase I of PMC division, <x 900 (2n—20). A-F from Purpus 5011; G & H from J.& C. Reeder 3669; A,-K, & G, from J. & C. Reeder 3963; Fi from Beetle M-481; H, from J. & C. Reeder 3622.
Additional collections:
MEXICO: Coahuila, 35-36 mi S of Saltillo along road to Concepcidn del Oro, Zacatecas, 6200 ft, J. R. & C. G. Reeder 3622 (YU), 6000 ft, J. R. & C. G. Reeder 3987 (YU). Nuevo Leon: Galeana y cercanias, A. Cuevas 107 (YU), Beetle M-481 (US) ; 3-4 mi SE of Galeana, 5200 ft, J. R. & C. G. Reeder 3659 (YU); J. R. & C. G. Reeder 3965 (YU). San Luis Potosi: ca 12 mi NW of Matehuala on road to Cedral, 5500 ft, J. R. & C. G. Reeder 3668 (YU).
188 MADRONO [Vol. 18
Fic. 2. Details of leaf anatomy and epidermis. Muhlenbergia purpusi (J. & C. Reeder 3669) on the left; M. gypsophila (J. & C. Reeder 3963) on the right. A, A,, diagram of transection of entire leaf blade, x ca. 35 (the black areas represent par- enchyma sheaths); B, B, transection through region of a major vein, B, X 200; Bi, X 150; C, Ci, portion of epidermis from abaxial surface, * 225; D, Du, epi- dermal bicellular microhair, & 225. All drawings made from tracings of photo- micrographs.
1966] REEDER AND REEDER: MUHLENBERGIA 189
As indicated above, M. gypsophila appears to be related most closely to M. purpusit. These two species occupy similar habitats, and on one occasion we collected both in the same gypsum flat in San Luis Potosi. The most obvious difference between the two is that the new species is a somewhat smaller plant with a contracted spike-like panicle, whereas its ally is taller and the panicle is open. Both species have membranous ligules, but that of M. gypsophila is shorter and more firm. The leaf blades of the two appear somewhat alike but those of the new species are closely involute and essentially smooth on the abaxial surface, where- as in M. purpusi they are flat or folded and scabrous to hispidulous be- low. In spikelet morphology the two species are quite similar, but in M. gypsophila the glumes and florets are somewhat larger. This is true also in the case of lodicules, stamens, and caryopses (fig. 1).
Both species are diploid with a chromosome number of 2n = 20, as determined from study of meiosis in pollen mother cells. It is noteworthy, however, that M. gypsophila has somewhat larger chromosomes (fig. 1, H&H,)).
It has been noted already that the leaves of M. gvpsophila are closely involute, whereas in M. purpusi they are flat or folded. More striking and important differences are revealed, however, when leaves of these two species are examined in transection. In those of M. gvpsophia the lower epidermis is essentially smooth, whereas the upper surface bears numerous spines and papillae. The leaf has commonly 5 major bundles which are raised prominently above the remainder of the tissue. Two smaller bundles which are partially submerged occur one on either side of the midrib. Alternating with the units already mentioned are small bundles which are almost completely submerged in thick-walled paren- chyma. Above the smooth lower epidermis is a layer of sclerenchyma 2 to 4 cells in thickness, and above this, thick-walled parenchyma which is in contact with the mesophyll. The major units show the familiar era- grostoid-chloridoid pattern, the bundle being surrounded by a double sheath, the inner of which consists of thick-walled sclerenchyma, and the outer of large parenchyma cells which contain chloroplasts. The paren- chyma sheath is continuous in all except the 5 largest bundles. Outside this sheath, the cells of the mesophyll are arranged in a radial pattern. Bulliform cells in this leaf are rather poorly developed (fig. 2, A; & B,).
The contrast between the leaf described above and that of VM. purpusii is striking. In the latter, spines and papillae are to be found in about equal numbers on both the upper and lower epidermis. It will be noted also that the units of the leaf differ only slightly in size, except for the larger midrib, and that they appear rather like the links of a chain. Each of these units is separated from its neighbors by a row of bulliform cells which extend from the upper to the lower epidermis. The bundles are of the typical eragrostoid-chloridoid type, as described above, but here the parenchyma sheath is continuous in all except the main vein. The radiat-
190 MADRONO [Vol. 18
ing mesophyll extends to the bulliform cells on either side, and above and below is in contact with small patches of sclerenchyma which are just below the epidermis (fig. 2, A & B).
Not only the leaf transection, but also the epidermis is strikingly dif- ferent in these two species. Both bear rather typical eragrostoid-chlori- doid bicellular microhairs, but these have a different shape and arrange- ment. In those of M. purpusi, which are shorter in overall length, the apical cell is proportionately somewhat larger. They are found in the intercostal regions of both the upper and lower epidermis, where the stomata are also located. The numerous siliceous cells, which are in the costal region, are saddle-shaped (fig. 2, C & D). On the lower epidermis of M. gypsophila, which is essentially smooth, there are no stomata nor microhairs. The most striking feature is, perhaps, its uniformity, there being little or no difference between the costal and intercostal areas. The entire epidermis consists of essentially three types of cells — long cells, short cells, and siliceous cells. These latter which are abundant and rounded in outline, quite unlike those of M. purpusti, occur in rows of alternating short and siliceous cells. These are separated from similar series by one or two rows of elongated cells. The long and short cells both have very wavy margins. The upper epidermis, as mentioned previously, bears numerous spines and papillae. Stomata are frequent in the inter- costal regions, and bicellular microhairs occur in these areas as well. Neither stomata nor microhairs are to be found elsewhere on the leaves of this species (fig. 2, C; & Dj).
To encounter such striking differences in the leaf anatomy and epider- mis of presumably closely related species is certainly not to be expected. Perhaps the relationship here is, in fact, rather remote. The similarities in gross morphology may reflect parallelism or convergence rather than close genetic affinity.
MUHLENBERGIA PURPUSII Mez, in Repert. Sp. Nov. 17: 214. 1921.
Specimens examined: Mexico. San Luis Potosi: Minas de San Rafael, Nov. 1910, Purpus 5011 (US, Type fragment; NY, isotype). Near the Minas de San Rafael in the Sierra de Guadalcazar, 1900-2100 m, Sohns 1515 (US, YU); 3 mi E of junction of Guadalcazar road and Hwy. 57, 5500 ft, J. R. & C. G. Reeder 4075 (YU); Near Km 549, 36 mi S of Matehuala, 1250 m, McVaugh, Loveland & Pippen 18204 (ENCB, US); Km 556, Carretera México-Piedras Negras, 1400 m, Rzedowski s.n. (US); San José del Refugio, carretera central, Km 550, Medellin L. 1130 (ENCB, SLP, YU); 12 mi NW of Matehuala on road to Cedral, 5500 ft, J. R. & C. G. Reeder 3669 (YU); 4 km al norte de Rioverde, sobre el camino a Pastora, 1000 m, Rzedowski 9574 (ENCB, SLP, YU).
Although M. gypsophila and M. purpusii occupy similar habitats, their ranges appear to be rather distinct. The new species has been collected in the states of Coahuila, Nuevo Leon, and northern San Luis Potosi. In the last locality it was found growing with M. purpusi and this is, in- deed, the only area in which we have found these two species associated. All collections of M. purpusii, of which we are aware, have come from the
1966 | REEDER AND REEDER: MUHLENBERGIA 191
COAHUILA NUEVO LEON
TAMAULIPAS
ZACATECAS
role)
° °
SAN LUIS POTOSI
°
Fic. 3. Map showing the range of Muhlenbergia gypsophila (closed circles) and M. purpusi (open circles).
state of San Luis Potosi. The ranges of the two species, as far as we have been able to determine them, are shown on the map (fig. 3).
In his comprehensive thesis on the “Vegetacidn del Estado de San Luis Potosi,” Rzedowski (1961) describes the gypsiferous areas in some detail. He refers to the vegetation of these habitats as ‘“‘zacatal ralo,” in- dicating that only about 30 per cent of the soil is occupied by plants, the dominant elements of which form circular colonies with bare centers [fairy-rings|. Bouteloua chasei and M. purpusii are given as the domi- nant grass species. In Fig. 26 of the publication cited above, which is a photograph of one of these habitats near Vallejo, the “fairy-rings” are clearly shown.
Rzedowski’s description of the habitat applies well to most of the areas in which we have encountered both M. purpusiu and M. gypsophila. Except for the region near Vallejo, however, in the areas in which we have observed VM. purpusii it has not appeared to be a dominant element. For the most part, we have found it as scattered clumps amongst Bou- teloua chasei and M. villiflora which in our experience are usually the dominants in these gypsum flats.
SUMMARY
Muhlenbergia gypsophila, a species apparently restricted to gypsum soils of north central Mexico, is described as new. It is compared with its presumed closest ally, M. purpusii Mez, a poorly known Mexican
192 MADRONO [Vol. 18
endemic. Although both species are diploid (2n = 20) and occupy simi- lar habitats, detailed studies of the leaf anatomy and epidermis reveal striking differences between these two grasses. A range map showing the known collecting sites for both species is provided.
Yale University, New Haven, Connecticut
LITERATURE CITED
Hitcucock, A. S. 1935. Muhlenbergia. Jn N. Am. FI. 17:431-476.
Jounston, I. M. 1941. Gypsophily among Mexican desert plants. Jour. Arnold Arb. 22:145-170.
RzEDOWSKI, J. & GRACIELA C. DE RZEDOWSKI 1957. Notas sobre la flora y la vegeta- cion del estado de San Luis Potosi. VI. Algunas plantas nuevas para la flora del Estado. Acta Ci. Potos. 1:199-218.
. 1961. Vegetacion del estado de San Luis Potosi. Tesis: Universidad Naci- onal Autonoma de México, D. F., Facultad de Ciencas.
NOTES AND NEWS
Selenium, Geobotany, Biochemistry, Toxicity and Nutrition. By IRENE ROSEN- FELD and OrvILLE A. BEATH. xii + 411 pp. Academic Press, New York. 1964. $15.00.
Wild Flowers in South Carolina. By Wave T. Batson. vi + 146 pp. University of South Carolina Press, Columbia. 1964. This is an attractive, popular book with glossary, keys, descriptions, and colored photographs of the more common woody and herbaceous flowering plants of South Carolina.
Great Smoky Mountains Wildflowers. By Cartos C. CAMPBELL, WILLIAM F. Hutson, HersHat L. Macon, and Aaron J. SHARP. 88 pp. University of Tennessee Press, Knoxville. Enlarged edition. 1964. $1.50. This spiral-bound booklet contains very good colored photographs and notes on about 125 kinds of flowering plants of the Great Smoky Mountains.
Trees, Shrubs, and Woody Vines of Great Smoky Mountains National Park. By ARTHUR STUPKA. x + 186 pp. University of Tennessee Press, Knoxville. 1964.
Algae and Man. Edited by Danie F. JAcKson. x + 434 pp. Plenum Press, New York. 1964. $14.50.
Aquatic Plants of the Pacific Northwest with Vegetative Keys. By A. N. STEW- ARD, LA REA J. DENNIS, and HELEN M. GILkey. Oregon State Monographs, Studies in Botany No. 11. x + 261 pp. Oregon State University Press, Corvallis. Second edition. 1963. $4.50.
The Developmental Anatomy of Isoetes. By Dominick J. PAOLILLO, JR. Illinois Biological Monographs No. 31. vi + 130 pp. University of Illinois Press, Urbana. 1963.
A Journal of Explorations Northward along the Coast from Monterey in the Year 1775. By FaTHER MIGUEL DE LA CaMPA. Edited by JoHN GALVIN. 67 pp. John Howell Books, 434 Post St., San Francisco. 1964. $7.50.
Journal of José Longinos Martinez, Notes and Observations of the Botanical Expedition in Old and New California and the South Coast 1791-1792. Translated and edited by LEstEy Byrp SIMPSON. xviii + 114 pp. John Howell Books, 434 Post Street, San Francisco. 1961.
The Oregon Desert. By E. R. Jackson and R. A. Lone. xvi + 407 pp. The Caxton Printers, Ltd., Caldwell, Idaho. 1964. $6.50. Although primarily historical, this book contains considerable information about the natural history of the arid part of Oregon. In particular there are two chapters, The Desert-Wise Plants and Grass Grows by Inches—It is Destroyed by Feet, dealing with the vegetation of the region.
B; +4 AY » OHA
ff VOLUME 18, NUMBER 7 JULY, 1966
Contents
INFLUENCES OF FIRE AND SODIUM-CALCIUM BORATE ON CHAPARRAL VEGETATION, Stephen K. Stocking 193
THE TAXONOMIC STATUS OF STEMMATELLA (CoMPOSITAE-HELIANTHEAE), B. L. Turner 203
VOLATILE GROWTH INHIBITORS PRODUCED BY EUCALYPTUS GLOBULUS, Herbert G. Baker 207
THE CHROMOSOMES OF SCOLIOPUS (LILIACEAE), Marion S. Cave 211
Two NEw SPECIES OF VERBESINA (COMPOSITAE) WITH TAXONOMIC NOTES ON OTHER SPECIES, James R. Coleman 214
OBSERVATIONS ON WITCHES’-BROOM FORMATION, AUTOPARASITISM, AND NEw Hosts IN PHORADENDRON, Frank G. Hawksworth and Delbert Wiens 218
NoTEs AND NEws: THE OCCURRENCE OF A NEW ZEALAND PLANT, GERANIUM MICROPHYLLUM IN CALIFORNIA, Margaret S. Bergseng 213 NEw PUBLICATIONS 224
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INFLUENCES OF FIRE AND SODIUM-CALCIUM BORATE ON CHAPARRAL VEGETATION
STEPHEN K. STOCKING
Fire is commonly recognized as an important factor in the determina- tion of the composition of the extensive chaparral areas of California. Environmental conditions, physiognomy of the vegetation, and physio- logical responses of the individual components have led many authors to conclude that the chaparral is a firetype vegetation, the composition of which is perpetuated through the action of fire. Cooper (1922), Sampson (1944), Horton and Kraebel (1955), Patric and Hanes (1964) and Sweeney (1956) have made detailed ecological studies of chaparral areas of various parts of California. The literature on vegetation and fire has been reviewed by the above authors and by Ahlgren and Ahlgren (1960).
Seeds may germinate readily or require various conditions to break dormancy caused by embroyo or seed coat conditions. Amen (1963) asserts that seed germination, germination mechanics, and the adaptive significance of the dormant state have not been adequately investigated. Quick (1935), Stone and Juhren (1951), Sweeney (1956), and Hadley (1961) have investigated the germination mechanics of some species which appear following chaparral fire. Sweeney (1956) found that both scarification and experimental burning of excelsior above planted seeds of “burn” species increased germination. Hadley (1961) increased germi- nation of Ceanothus megacarpus both by scarification and by heating the seeds to 100° C for five minutes.
In recent years, chemicals such as sodium-calcium borate, bentonite clay, and diammonium phosphate have found increasing use as fire re- tardant chemicals. Experiments have shown that borate compounds inhibit germination and are toxic to mature plants when used in high concentrations (Brenchley, 1914; Crafts, 1956; Johansen, 1958). Field investigation of the effects of sodium-calcium borate on chaparral vege- tation has not been reported.
The purpose of this investigation is mainly to record the nature of vegetation which appears following fire in a chamise chaparral area. Consideration is also given to germination responses of some of the herbaceous species which appeared on the burned area immediately fol- lowing the fire and to the effect of sodium-calcium borate on growth in an affected area.
The study area is located in the southern Sierra Nevada foothills with- in Sequoia National Park. The study plots are in the lower Elk Creek drainage at an elevation of approximately 2100 feet. Soil is of the Vista
Maprono, Vol. 18, No. 7, pp. 193-224. September 23, 1966. 193
194 MADRONO [Vol. 18
series (Storie and Weir, 1962), and ranges from two to five feet in depth. Weather and fire fighting data were taken from National Park Service records at the nearby Ash Mountain headquarters. The Tunnel Rock fire began June 25, 1960 and burned 4673 acres of chaparral and woodland on the south and east slopes of Ash Peaks. It was estimated that 25 to 30 500 gallon drops of sodium-calcium borate were made before the fire was brought under control.
Seventeen field plots, each four meters square, were located in the lower edge of the burned chaparral. Eleven of these plots are on the south-facing slopes which predominate in the burned area. Five plots are in the burned area, three on the margin of the borated and burned area, and three in the borated and burned area. The three control plots are in one of the few limited areas of unburned chaparral. All plots lie within a radius of 300 yards.
Ring counts were made of specimens of the woody perennial Adeno- stoma fasciculatum which had burned near the study plots. The number of annual rings in three complete sections varied from 45 to 51. The center of some sections was rotten or hollow, but 30 to 48 annual rings were visible even in the six incomplete specimens. Because aerial portions of this plant are killed by the heat of chaparral fire, it is assumed that there had not been a major fire in the immediate area for at least 50 vears.
Principal observations and studies of the vegetation were made during the first growing season following the fire, the spring of 1961. Plant collections were made to determine the floristic composition of the area, and to find the frequency of occurrence of these plants on burned and adjacent unburned areas. The collections included 148 species, of which 118 were herbaceous, 25 annuals, 20 perennials, and seedlings of three woody perennials were found on the burned chaparral areas. Those species occurring on the study plots are listed in Table 1. More detailed information regarding distribution of vegetation in all areas may be found in Stocking (1962).
Marked differences in occurrence and abundance of certain species was evident when the burned area and the unburned control areas were compared. The chaparral cover of unburned south-facing slopes is pre- dominately Adenostoma fasciculatum, Ceanothus cuneatus, Arctosta- phylos viscida, Fraxinus dipetala, and Quercus wislizenit occur near margins of the dense chaparral and on the more mesic north-facing slopes. No seedlings of woody perennials were encountered either under the unburned brush or in openings in the brush. Only 12 herbaceous species were found in the control area. Failure of seedling establishment in these areas may be due to any of many factors including limited temperature range, root distribution, low light intensity, presence of inhibitors, and necessity of heat of fire for germination.
Seedlings of Eviodictyon californicum and Ceanothus cuneatus were occasionally encountered, particularly on the north-facing slopes (table
1966] STOCKING: CHAPARRAL 195
1). On burned chaparral areas, seedlings of chamise were abundant: 372 seedlings appeared on one 4 m study plot. Only three of these seedlings survived the hot dry summer. Survival of these seedlings varied on the different plots, but was low in all cases. Low survival of chamise is thought to be due to the shallow nature of the root system during the first year of growth.
Presence of herbaceous vegetation in open areas indicates that light intensity may play an important part in the establishment of some species in unburned chaparral areas. Species found most often in open areas in the brush of control plots are shown in Table 1. Of these species only Lotus scoparius, Daucus pusillus, and Gastridium ventricosum were found on adjacent burned areas. Occurrence of these species in un- burned brush areas indicates that conditions caused by fire are not requisite for their germination and growth. If invasion from unburned chaparral areas is to account for vegetation on the adjacent burned areas it would be expected that these species would predominate. Of the species which occurred on the burned area only the above mentioned species could have originated in neighboring unburned areas. The bulbous perennials, Brodiaea lutea var. scabra, and Chloragalum pomeridianum developed from underground bulbs on both burned and unburned areas. The other species of abundance on the burn were not present in the control area.
All of the scattered individuals of Quercus wislizeni and Fraxinus dipetala were in mesic locations, and all crown sprouted following the fire. Less than one-half of the burned chamise sprouted from their con- spicuous basal burls. Most of these crown sprouts survived the first hot summer following the fire. No seedlings of any species were noted im- mediately adjacent to the bases of mature chamise. Lack of herbaceous vegetation near chamise may be due to the action of phytoxic agent found in the leaves of the species by Naveh (1960). Herbaceous plants were present only where open areas existed in the otherwise dense cover of the control plots. The thinning out of the cover appared to be due both to the advanced age of the stand and to the long period of low rainfall.
Observations were made of: 1, grassy areas surrounded by chaparral: 2, disturbed areas formed by an improved trail; and 3, disturbed areas formed by a road and utility right of way. The herbaceous flora of these burned areas differed from that of burned chaparral areas. With the exception of D. pusillus, and Gastridium ventricosum, no species oc- curred on the burned chaparral area which also occurred on the dis- turbed areas. Presence of these species on the above mentioned areas may be due to many factors. It is possible that no mature plants, and therefore no seeds, existed on chaparral areas prior to the fire. It is also possible that the seeds of these species have low heat tolerance, and could survive fire only in open areas where the heat would not be so intense. The relation of heat generated by chaparral fire to seed survival and germination has been discussed by Sweeney (1956).
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In addition to collections made in burned chaparral areas, many species were found in woodland and riparian areas burned in the same fire. No species were found in these areas which occurred in abundance on the chaparral area. This eliminated the likelihood of invasion from these areas during the first season following the fire.
Dendromecon rigida and Emmenanthe penduliflora were found in great abundance on localized areas of burned chaparral. These species were not found on unburned areas but have been located on other dis- turbed chaparral areas. They did not occur on the study plots. The most abundant herbaceous plant which appeared on burned chaparral areas was the perennial, Astragalus congdonti. Malacothamnus fremontii was locally abundant but only scattered in the area of the study plots. Both these plants are very rare in unburned chaparral areas. Only a few plants have been located by the author in disturbed areas of unburned chaparral in the southern Sierra Nevada.
Other seedlings found in abundance on burned chaparral areas, but rarely elsewhere, were Crypantha micromeres, Trifolium ciliolatum, T. microcephalum, Lotus subpinnatus, L. micranthus, and Astragalus di- dymocarpus. Distribution patterns of species restricted to the burn varied suggesting differences in abundance of these plants prior to the fire. Emmenanthe penduliflora was most common in rocky areas follow- ing the fire. This suggested that there were focal areas in which a few plants occurred in the interval between fires. Horton (1960) postulated similar origins of firetype plants in southern California.
Ceanothus cuneatus was nowhere abundant prior to fire, but seedlings appeared following fire on north-facing slopes (table 1). This finding concurs with those of Horton (1945), Hadley (1961), and Patric and Hanes (1964) who found that in areas unburned for over 50 years, Ceanothus spp. were replaced by species with a longer life span. Horton and Kraebel (1955) found that the seeds of Ceanothus spp. were stimu- lated to germinate through the action of fire. Although chamise seedlings were not found on the control areas they were abundant on the burned areas. Stone and Juhren (1953) found that chamise produced seeds with different germination responses at different times in their lives. They believed that an old stand of brush would contain a large number of viable seeds which would germinate in response to fire.
One possible explanation for the widespread abundance of Astragalus congdonu, Trifolium ciliolatum, and some other species on burned areas but not elsewhere is long term seed survival. It is possible that an abund- ance of seeds of these species had survived in the soil since a period of abundance following the last fire. These two species were found to have two types of seed, one which germinated readily and another more abundant type which required scarification for germination (table 2). The readily germinable seed would make possible scattered occurrence in the absence of fire. The other seed type would remain in the soil in the period between fires. Both species are legumes, a group which is
200 MADRONO [Vol. 18
known to contain many species with long lived seeds (Quick, 1961). This condition could explain the widespread occurrence of these two plants following fire. Emmenanthe penduliflora and Malacothamnus fremontu were found to have seeds which would germinate only when scarified (table 2). It is postulated that these species occur in the ab- sence of fire only where disturbances would break the seed coat of an occasional seed. This would explain their local abundance following fire. Amen (1963) states that the mechanisms of such dormancy are elusive, but that delayed germination represents a special adaptation which favors the continued propagation and survival of the species.
Burned north-facing slopes had a somewhat different cover of both annuals and perennials than did the plots on the drier south-facing slopes (table 1). In these areas vegetative activity was observed to begin earlier and reproductive activity to end later than on the south-facing slopes. Seedling survival of perennial species was also greater.
Observations made during the second spring following the fire indi- cated that the vegetational composition had changed. Annual species which were unique to the burned area during the first season had, for the most part, disappeared. Only a few individuals of Emmenanthe penduli- flora were found at this time. The transient occurrence of such species suggests a direct relationship between fire and germination. Perennial species such as Astragalus congdonii and Brodiaea lutea var. scabra re- mained in abundance on the burned areas during the second season.
In the course of field investigations during the first growing season it was found that a section of the study area lacked the annual and peren- nial vegetation characteristics of other burned areas. It was learned that this area was one of 25 to 30 locations hit by aerial drops of the fire retardant chemical, sodium-calcium borate.
Complete lack of perennial and herbaceous seedlings in this area indi- cated that boron was present in highly toxic concentrations in the surface layers of the soil. At a depth of 0 to 7.6 cm, any toxic materials would be in contact with young absorbing roots. The one herbaceous perennial present, Chloragalum pomeridianum, showed typical boron toxicity symptoms of leaf burn and marginal chlorosis. These symptoms in Chlorogalum pomeridianum suggested that boron was also present in the root zone of this plant, 7.6 to 30.5 cm beneath the surface. No con- ditions attributable to boron were exhibited by crown sprouts of chamise which has a deeper and more extensive root system than do the her- baceous species.
Vegetation of plots adjacent to the above mentioned area was more sparse than on neighboring unaffected areas. The species here were the same as on areas not affected by “borate.” Sparcity of vegetation in this area suggested that boron was present in relatively low concentrations. The species present of the affected areas are shown in Table 1.
Colorimetric determination of boron in soil collected from the affected area was done according to the quantitative method of Hatcher and
1966] STOCKING: CHAPARRAL 201
TABLE 2. EFFECT OF MECHANICAL RUPTURING OF SEED COAT ON GERMINATION OF SOME BurRN SPECIES INCLUDING TWo WITH SEED Coat DIMORPHISM
Percentage germination!
Sepcies Not Scarified Scarified
Malacothamnus fremontii @) 30
Emmenanthe penduliflora 0 41
Trifolium ciliolatum 65 — (buff seed color)
Trifolium ciliolatum 0 82 (mottled seed color)
Astragalus congdonii 74 — (buff seed color)
Astragalus congdonii 0 78
(mottled seed color)
1 Each treatment consisted of three replications using 25 seeds per dish. Germi- nation was defined as emergence of the radicle by the end of 30 days.
Wilcox (1950). Analysis was made to determine the residual water soluble boron in the soil following aerial application. The manufacturer stated that concentrations of two to five ppm of boron in soil is toxic to most plants (U.S. Borax and Chemical Corp., 1957).
Boron is slowly removed from soil both by leaching and fixation. Crafts (1956) stated that a pattern of concentrated rainfall is much more effective in leaching soil sterilants of this type than is a comparable amount of intermittent rainfall. During the first winter following the application of “borate,” rainfall was intermittent in occurrence and in small quantities. 5.66 cm of rain fell between the application and the collection of the first soil samples. Progressive leaching of the soil by low, intermittent rainfall is presumed to have reduced the boron con- centration present immediately following application. At the end of the growing season, boron remaining in the top 7.6 cm of the soil in the affected area was reduced to 25 to 30 ppm. Fifteen ppm boron were found at depths of both 7.6 to 20.3 cm and 20.3 to 30.5 cm. All these concentrations are well above the minimum toxic concentrations. It is assumed that a somewhat higher boron concentration was present during the period of seed germination.
Analysis of soil collected in the same area during the second spring following the fire showed that 6.5 ppm of boron remained in the top 7.6 cm. This concentration remained after a total rainfall of 101 cm. It was again observed that no seedlings occurred in the area and that Chlorogalum pomeridianum continued to exhibit toxicity symptoms. Vegetation of the plots near the margin of the “borate” drop was much more abundant than during the first year. Vigorous chamise crown sprouts evidenced no toxicity in any areas.
These data indicate that concentrations of boron between 30 ppm and 6.5 ppm made seedling development impossible. Soil in this area remained sterlie for twenty months after the aerial application of
202 MADRONO [Vol. 18
“borate.” At least 23.5 ppm of boron was removed by leaching or be- come unavailable by absorption during the second year following appli- cation. It is likely that 6.5 ppm of boron remained at the onset of the third winter (1962-1963). Rainfall during the winter of 1962-1963 was 75.11 cm.
Most seeds, both with or without seed coat dormancy, would germi- nate within the first two seasons following fire. Once these seeds began to germinate they would be exposed to the toxic concentrations of “borate.” Not until the third growing season following fire did Lactuca serriola, a species with wind borne seeds, and a few grasses begin to appear at the margin of the area directly affected by the aerial drop. This would indicate that the rate of depletion continued during the third year, and that boron was reduced to below toxic levels.
Observations made during the springs of 1963 through 1965 showed that species appearing on the area which had been affected by “borate” were not the same as those which appeared on the surrounding unaffected chaparral area immediately following fire. Species found on the “‘borated”’ areas were predominately those of neighboring disturbed areas. Further study of such areas may lead to a better understanding of the compara- tive roles of invasion and seed survival in the establishment of plants on burned chaparral areas.
This paper is adapted from a portion of a masters thesis prepared at San Francisco State College, San Francisco, California. I wish to ac- knowledge the assistance of James Sweeney, Eva Estermann, Arthur Nelson, and Harry Thiers of San Francisco State College. I am indebted to J. T. Howell for aid in plant identification, and to the staff of Sequoia- Kings Canyon National Park for their cooperation.
2217 Polk Way, Stockton, California
LITERATURE CITED
AHLGCREN, I. F., and C. E. AHLGREN. 1960. Ecological effects of forest fires. Bot. Rev. 26:483—-533.
AMEN, R. D. 1963. The concept of seed dormancy. Am. Sci. 51:408—-424.
BRENCHLEY, W. E. 1914. On the action of certain compounds of zinc, arsenic, and boron on the growth of plants. Ann. Bot. 28:283—301.
Cooper, W. S. 1922. The broad sclerophyll vegetation of California. Carnegie Inst. Wash. Pub. 319.
Crarts, W. S. 1956. Weed control: applied botany. Am. Jour. Bot. 43:548—-556.
Haptey, E. E. 1961. Influence of temperature and other factors on Ceanothus megacarpus seed germination. Madrono 16:132-138.
Harcuer, J. T., and L. V. Witcox. 1950. Colorimetric determination of boron using carmine. Anal. Chem. 22:567—569.
Horton, J. S. 1945. Vegetation of the San Bernardino Forest. (As cited in Shantz, H.L. 1947. The use of fire as a tool in the management of the brush ranges of California. Div. of Forestry, Calif. State Board of Foresty. Sacramento.)
, and C. I. Kraepev. 1955. Development of vegetation after fire in the chamise chaparral of Southern California. Ecology 36:244—263.
Jouansen, R. W. 1958. Final report: toxicity phase of study FS 2-f2-11 (S-1). US.
Department of Agriculture Forest Service. Southern Forest Fire Lab.
1966] TURNER: STEMMATELLA 203
Muwz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
NaveEH, Z. 1960. The ecology of chamise as effected by its toxic leachates. Bull. Ecol. Soc. Am. 41:56—57.
Patric, J. H. and T. H. Hanes, 1964. Chaparral succession in a San Gabriel Moun- tain area of California. Ecology 45:353—360.
Quick, C. R. 1935. Notes on the germination of Ceanothus seeds. Madrono. 3:135-
139. . 1961. How long can a seed remain alive. Jn U.S. Department of Agricul-
ture Yearbook.
Sampson, A. W. 1944. Plant succession on burned chaparral lands in northern California. Calif. Agr. Exp. Sta. Bull. 635.
StocKI1nG, S. K. 1962. Some aspects of the influence of fire and sodium-calcium borate on chaparral vegetation. Master’s degree thesis (unpublished). San Francisco State College, California.
Stone, E. C., and G. JuHreN. 1953. Fire stimulated germination. Calif. Agric. 7(9) 213-14.
Storie, R. E., and W. W. WErr. 1962. Generalized soil map of California. Univ. Calif.
SwEENEY, J. R. 1956. Responses of vegetation to fire. Univ. Calif. Publ. Bot.
28:143-250. U.S. Borax and Chemical Corp. 1957. Firebreak chemical fire retardant. Tech. Bull,
FB/1.
THE TAXONOMIC STATUS OF STEMMATELLA (COM POSITAE-HELIANTHEAE)
B. L. TURNER
The genus Stemmatella was first proposed by Weddell (Bull. Soc. Bot. Fr, 12:82. 1865) in a list of plants collected in Bolivia by M. G. Mandon and identified by Schultz-Bipontinus. Specifically, the name Stemmatella congesta Wedd. preceded by the collector’s No. 293, was listed without description or comment. As such Stemmatella was a nomen nudum until subsequently published by Hoffmann (1894) in his treatment of the Compositae. Bentham (1873), however, drew up a rather complete generic description, although he neglected to mention S. congesta, or any other species name, nor did he make reference to a specimen, but pre- sumably he had access to one of the Mandon collections (see below).
I became interested in the taxonomy of Stemmatella while attempting to identify epappose forms of Galinsoga parviflora (Turner et al., 1962; Turner and King, 1964), for such specimens, by Hoffmann’s treatment would key to Stemmatella instead of Galinsoga; indeed, collections filed under the two generic names were almost identical, in spite of the fact that they were placed by both Bentham and Hoffmann in the subtribes Verbesininae and Galinsoginae respectively.
Bentham had a single sheet with 2 attached specimens of the Mandon 293 collection at his disposal (K). One of the specimens contains heads with epappose achenes; the other has heads with epappose ray achenes but the disk achenes possess a well-developed pappus. One must assume
204 MADRONO [Vol. 18
that Bentham examined only the former in drawing up his otherwise excellent generic description.' Hoffmann, apparently relying on Ben- tham’s treatment, also assumed that the disk achenes were epappose, for this is the character which he uses to separate Stemmatella from the subtribe Galinsoginae. In any case, I have examined in detail nine iso- types of S. congesta (K, 2; NY, 6; US, 1; the plants are all quite dwarf as if closely browsed or trodden on) and in each (except for one of the specimens at Kew) the ray achenes are epappose while the disk achenes support a well-developed pappus, very similar to that of Galinsoga parviflora.
Hoffmann, after his 1894 treatment of Stemmatella in which but a single species was acknowledged, must have recognized additional species, for in 1901 we find Hieronymus publishing a manuscript name of Hoff- mann’s, Stemmatella urticifoia (H.B.K.) O. Hoffm. ex Hieron. (Bot. Jahb. 28:601). In this same publication Hieronymus described two addi- tional species, S. sodiroi and S. lehmannii. Both of the latter were recog- nized as closely related to S. urticifolia. Oddly enough, Hieronymus himself must have recognized the illogical separation of the two genera, for after his description of S. sodiroi he comments, “An igitur melius genus Stemmatella genere Galinsoga conjungendum sit?” He was, in fact, as indicated below, describing nothing but a form (with glabrate achenes) of Galinsoga urticifolia (H.B.K.) Benth.
Having now examined type material of all of the species-names pro- posed for Stemmatella, it is my opinion that the genus should be sub- merged in Galinsoga.- My nomenclatural and morphological observations follow:
STEMMATELLA CONGESTA Wedd. ex Hoffm., Naturl. Pflanzenfam. 4(5): 231. 1894. Bolivia, Prov. Larecaga “Viciniis sorata,”’ Mandon 293, (w/o date), (isotypes, GH!, K!, NY!, US!).
The nature of the types on which this name is based is discussed above. Stemmatella congesta appears to be closest to Galinsoga calva Rusby (non G. calva Sch.-Bip., 1865; nomen nudum ). The Mandon 293
1 Jronically, in the original version of the present paper, after my initial examina- tion of the material at Kew, I implied that Bentham had been guilty of faulty observation. In fact, I had looked at only the specimen which Bentham had not examined—lI am grateful to the reviewer of this paper for suggesting a second look.
2 While re-examining specimens at Kew filed under the generic name Stemmatella, I came across an unpublished manuscript (inserted in one of the species-folders) entitled “Notes by P. G. Wilson, 1957.” Mr. Wilson makes the following observa- tions which substantiate my own, “Stemmatella Wedd. has been delimited in the Genera Plantarum by having the ray florets enclosed within an outer involucral bract and two palae, as well as having naked achenes. This description applies to Wiborgia urticifolia Kunth which has been transfered to Stemmatella by Hoffm. ex Hieronymous. However, the character of the ray florets being enclosed within three bracts is also held by Galinsoga which normally has achenes with a pappus. W. urticifolia matches Galinsoga hispida Benth. exactly, except for the pappus character (present in G. hispida) and they come from the same locality.”
1966] TURNER: STEMMATELLA 205
collections are a good match for isotypic material of G. calva (Bolivia, Talca Chugiaguillo, A. Miguel Bang 809, Apr. 1890) which was ex- amined at Kew. The latter is from the same general area, has similar, congested heads, ray achenes without pappus and disc achenes with pappus, etc. A critical evaluation of the nomenclatural status of the numerous specific names proposed for Galinsoga must await a more thorough biological study of that genus, but in my opinion there can be no question but what the type species of Stemmatella is in fact a Galinsoga.
GALINSOGA URTICIFOLIA (H.B.K.) Benth., Oesterr. Vid. Middel. 102. 1852. Wiborgia urticifola H.B.K., Nov. Gen. Sp. 4:201 (257) t. 389. 1820. “Crescit juxta villam Marchionis de Miraflores, inter Mulalo et Pansache, alt. 1700 hex. (Regno Quitensi.)” June. Sabazia urticifolia (H.B.K.) DC., Prod. 5:497. 1836. Stemmatella urticifolia (H.B.K.) O. Hoffm. ex Hieron., Bot. Jahrb. 28: 601 1901. Stemmatella sodiroi O. Hoffm. ex Hieron., Bot. Jahrb. 28: 601-602. 1901. Ecuador, Crescit in regione interandina, Sodiro 31/1, (fragment of holotype, US!; photo- types, GH!, NY!, US!). Stemmatella urticifolia var. eglandulosa Hieron., Bot. Jahrb. 36:487. 1905, as Stematella. Peru, near Cutervo, April 1879, Jelski 609. Sabazia urticifolia var. venezuelensis Steyer., Fieldiana Bot. 28: 672. 1953. Venezuela, Lara, “‘Pastured open slopes of mountain between Santo Domingo and Los Quebraditos,” J. A. Stever- mark 55379, Feb. 8, 1944, (holotype, F! ).
Galinsoga urticifolia is, except for its epappose disk florets, very simi- lar to G. parviflora (as noted by Wilson, cf. footnote 3, above). How- ever, its natural distribution and ecology appears to differ from the latter in that it occupies more tropical, wetter areas at presumably some- what higher elevations. G. parviflora is more widespread, occurring in both tropical and subtropical areas, usually in dryer, somewhat lower habitats. These two species unquestionably come in close contact in cer- tain areas so that hybridization must be reckoned as a factor in their variation. Additional cyto-morphological observations can be found in Turner et al. (1962) and Turner & King (1964).
Stemmatella urticifolia var. eglandulosa and Sabazia urticifolia var. venezuelensis are apparently only forms of the species, the former lack- ing glandular trichomes on the stems and the latter possessing more densely pubescent achenes than is typical for the species. Both of these conditions in varying combinations are found in individuals from throughout the range of Galinsoga urticifolia.
STEMMATELLA LEHMANII Hieron., Bot. Jahrb. 28:602. 1902. Colum- bia, Vicinity of Popayan, 1600-2200 m, Lehmann 5667 (phototypes, NY!, TEX!, US!).
This species appears to belong to the genus Sadazia. It is apparently a perennial with thin rhizones (Core 1015); the heads are relatively large on long peduncles which arise singly from the leaf axils; the ligules are quite well developed and, in proportion to their length, shortly 3-
206 MADRONO [Vol. 18
lobed. To my knowledge, these characters are not found in Galinsoga. Except for the types, the only collection of this taxon which I have ex- amined has been that of Core 1015 (US), from a rodaside between Valencia and San Sebastian, Colombia, 21 July, 1944: ‘Weed in thick- ets; disk flower yellow, ray flowers white above, pink beneath, rather showy.”
Finally it should be noted that the epappose Galinsoga mandonii Sch.- Bip. (Linnaea 34:529. 1866), which is distinguished from the more widespread G. urticifolia by its smaller, congested (sessile) heads and deep purple involucral bracts, is sympatric with G. urticifolia in south- ern Bolivia and Peru. I have examined isotypic material of G. mandonii (Bolivia, Prov. Larecaja, Mandon 76, F, K) and the following collec- tions, all from Peru: Cook & Gilbert 210 (US); Killip & Smith 21741 (F); Macbride & Featherstone 573, 1582 (F). While closely related to G. urticifolia, it seems distinct; at least collections of a similar nature were not seen from the relatively well collected Colombia and Ecuador. However, a tendency toward intergradation with G. mandonii may be inferred from some of the collections of G. urticifolia from Peru (e.g., Macbride & Featherstone 2105, F ).
As may be surmised, Galinsoga is in much need of experimental study, both in the garden and in the field. Several of its “species” have become widespread weeds in the wetter temperate regions of both hemispheres and because it grows well in such climates it ought to lend itself to experimental work in those areas. If the amplitude of morphological variation of the several tropical species of Galinsoga which I haxe ex- amined is paralleled by a similar amplitude in physiological variation, then its success as an aggressive (or perhaps more aptly, “delightful” ) weed outside of its native habitat becomes explicable (cf. Baker’s com- ments, 1962, on tropical weeds).
John Beaman and an anonymous reviewer provided helpful sugges- tions and L. H. Shinners assisted with certain bibliographic problems. Financial support was provided in part by NSF GB-1216. I am grateful for all these benefits.
Department of Botany, The University of Texas, Austin LITERATURE CITED
BAKER, H. G. 1962. Evolution—the need for research in the tropics. Bull. Assoc. Trop. Biol. 1:102-104.
BENTHAM, G. 1873. Stemmatella. Jn Genera plantarum 2:359-360.
Hor¥Mann, O. 1894. Compositae. Jn Die Natiirlichen Pflanzenfamilien 4(5):87-391.
TurNER, B. L. and R. M. Kine. 1964. Chromosome numbers in the Compositae. VIII. Mexican and Central American species. Southw. Nat. 9:27-39.
, M. Powe 1, and R. M. Kine. 1962. Chromosome numbers in the Com-
positae. VI. Additional Mexican and Guatemalan species. Rhodora 64:251-269.
VOLATILE GROWTH INHIBITORS PRODUCED BY ECUALYPTUS GLOBULUS
HERBERT G. BAKER
The possibility that plants of some species gain advantage in natural competition by the excretion of substances (exocrines or ectocrines) in- hibitory to the growth of their potential neighbors is under investigation in a number of laboratories. Among these investigations, the studies by Muller and his co-workers on volatile substances produced by aromatic shrubs have aroused considerable interest. Working particularly with Salvia and Artemisia they have shown that volatile terpenes from the leaves may inhibit root-growth of Cucumis seedlings placed in a closed container with them (Muller, Muller, and Haines, 1964). They have also demonstrated the presence of these terpenes in the air surrounding these shrubs in nature (Muller and Muller, 1964; Muller, 1965) and have postulated how they may enter the cells of victimized seedlings through solution in cuticular lipids (Muller, 1965).
The simplicity of the experimental set-up used by these workers in their demonstration of root-growth inhibition by volatile substances from shredded Salvia leaves suggested that this might form a suitable class exercise in an ecology course. Consequently, it was tried with the Evolutionary Plant Ecology course at the University of California, Berkeley, in the spring semester of 1964, with subsequent additional experiments which have been facilitated by National Science Foundation Research Grant No. G 21821. I am indebted to Charles Quibell who carried out the extra experiments, meantime making very valuable sug- gestions regarding technique.
Results obtained by the class were variable until the experimental design was fully standardized. The chambers and contents (fig. 1) were modeled after the setup described in Muller, Muller and Haines (1964). The seeds of Cucumis sativus L. were soaked in distilled water for 2 hours and then placed in position on the moist filter paper in the cham- bers. The glass lid to each chamber was fixed in place with petrolatum. The chambers were then kept together in the dark at 26.7°C (80°F) usually for 48 hours before measurements of root-growth were made. Only main roots were measured, laterals being ignored. The very few ungerminated seeds were also left out of account.
The availability of a small introduced population of Salvia apiana Jeps. on Grizzly Peak, Berkeley, enabled the results in Table 1A to be obtained. The differences between controls and test growths are all signifi- cant at the 1% level, confirming the findings of Muller et al.
However, it seemed likely that some other, more abundant local source of exocrines could be found. In the vicinity of Berkeley there are numer- ous plantations of Eucalyptus globulus Labill. under which very few
207
208 MADRONO [Vol. 18
TABLE 1. VOLATILE GROWTH INHIBITORS PRODUCED BY EUCALYPTUS GLOBULUS
Duration Mean root-Standard %of of Test No. of length devia- growth of (hours) Seedlings (cms.) tion control
A. Salvia leaves — Cucumis roots
Control (3 chambers) 48 142 2.20 0.56 Salvia 1 42 1.62 0.54 74 Salvia 2 48 tea 0.48 67 Salvia 3 42 1.48 0.48 67 Control (2 chambers) 48 90 1.93 0.64 Salvia (2 chambers) 94 1.31 0.53 68 Control (2 chambers) 60 83 2.86 1.04 Salvia (2 chambers) 70 2.09 0.60 73 Control (4 chambers) 70 185 3.04 0.95 Salvia (3 chambers) 126 123 0.48 41
B. Eucalyptus leaves — Cucumis roots
Control (3 chambers) 48 142 2.20 0.56 Eucalyptus 1 46 1.68 0.57 76 Eucalyptus 2 48 1.34 0.46 61 Eucalyptus 3 44 1.61 0.56 73 Control (2 chambers) 48 90 1.93 0.64 Eucalyptus (2 chambers) 73 0.38 0.24 20 Control (2 chambers 60 83 2.86 1.04 Eucalyptus (2 chambers 7 0.71 0.35 25
C. Eucalyptus leaves — Eucalyptus roots
Control (a) 24 50 0.67 0.27 Juvenile Eucal. (a) 51 0.56 0.17 83 Mature Eucal. (a) 50 O57 0.16 85 Control (b) 24 48 0.91 0.11 Juvenile Eucal. (b) 50 0.83 0.25 91 Mature Eucal. (b) 50 1.06 0.32 117 Control (c) 24 10 1.35 0.32 Juvenile Eucal. (c) 10 1.33 0.30 99 Mature Eucal. (c) 12 125 0.37 93
D. Eucalyptus leaves — Eucalyptus hypocotyls
Control (a) 24 50 0.33 0.07 Juvenile Eucal. (a) Sik 0.25 0.05 76 Mature Eucal. (a) 50 0.21 0.04 64 Control (b) 24 48 0.43 0.15 Juvenile Eucal. (b) 50 0.28 0.06 66 Mature Eucal. (b) 50 0.27 0.06 62 Control (c) 24 10 0.48 0.14 Juvenile Eucal. (c) 10 0.25 0.04 53
Mature Eucal. (c) 12 0.31 0.05 68
1966] BAKER: GROWTH INHIBITORS 209
Fic. 1. Diagrammatic section through test chamber: a, glass lid; b, petrolatum seal; c, chamber; d, perforated container for test leaves; e, test seeds; f, filter paper ; g, sponge, moistened with distilled water.
other plants grow. Although this forest tree has many natural associates in its native Australia, the contemporary California flora (a mixture of native species and mostly European introductions) appears generally to be unadapted to it as yet. The leaves of E. globulus are intensely aro- matic, releasing a number of terpenes, including cineole (eucalyptole) (Baker and Smith, 1902). Cineole has been shown to be strongly in- hibitory to root-growth (Muller and Muller, 1964) and to be one of the volatile toxins released by species of Salvia (Muller, 1965). Conse- quently, it seemed likely that terpenes from EL. globulus might be respon- sible for the paucity of accompanying vegetation in the plantations.
Shredded Eucalyptus leaves proved to be just as efficacious as Salvia leaves in the inhibition of Cucumis seedling root-growth in class experi- ments in May, 1964 and in subsequent experiments in July 1964 (table IB).
If volatile exocrines from Eucalyptus trees can be inhibitory to the root-growth of seedlings of other species, it is of interest to know their effects on the growth of their own seedlings. A modification of the experi- mental procedure was necessary in testing this, however, because Eu- calyptus seed does not have the “simultaneous” germination pattern shown by Cucumis. It was necessary to germinate rather a large quantity of Eucalyptus seed after soaking for 48 hours in distilled water and then pick out seedlings with radicles of comparable length with which to begin the tests in the chambers. Table 1C shows the results obtained with shredded juvenile and adult leaves of E. globulus in the test cham- bers (the three sets of experiments being begun with germinated Eucalyptus seed having radicles of approximately (a) 1 mm, (b) 2 mm and (c) 3 mm, respectively).
210 MADRONO [Vol. 18
Although exact comparisons between experiments carried out at separate times are not completely valid (and amounts of leaf were not weighed exactly), it seems that the leaves tested in July 1964 were more effective in inhibiting root-growth than those sampled earlier in the year, in May. This might reflect a difference in terpene production at these times.
For E. globulus root-growth, the test-results vary around 100% and none is significantly different at the 1% level from the controls, suggest- ing that the roots of this species are little affected by the volatile sub- stances produced by the leaves of the same species. This might be ad- vantageous in allowing more than one plant of the species to establish itself from seed in a newly-invaded area. On the other hand, there was a consistent effect (P< 0.01) on the elongation of the hypocotyls of the Eucalyptus seedlings (table 1D). Several other experiments than those reported gave similar results with hypocotyl elongation. The ecological import of this finding has yet to be estimated.
The lack of effect of the volatile exocrines from Eucalyptus on the root-growth of its own seedlings recalls the apparent lack of sensitivity of the cells of Colchicum plants to the polyploidizing action of colchicine and the insensitivity to mutagenic allyl compounds shown by plants which produce them.
That plants will not always prove to be insensitive to the chemical products of their own species, however, is suggested by the observations of Went (1955) on creosote bushes (Larrea divaricata Cav.) which are often spaced remarkably evenly on the desert floors in California. Went attributes this to the susceptibility of Larrea seedlings to exocrines pro- duced from the roots of existing creosote bushes, as a result of which seedlings only become established outside the radius of the circle of roots of existing shrubs.
Department of Botany, University of California, Berkeley. LITERATURE CITED
Baker, R. T. and H. G. SmitrH. 1902. A research on the eucalyptus, especially in regard to their essential oils. Tech. Mus. New South Wales, Educ. series, no. 13. Sydney.
Mutter, C. H. 1965. Inhibitory terpenes volatilized from Salvia shrubs. Bull. Torrey Club 92:38-45.
, W. H. Mutter and B. L. Haines. 1964. Volatile growth inhibitors pro- duced by aromatic shrubs. Science 143:471-473.
Mutter, W. H. and C. H. Mutter. 1964. Volatile growth inhibitors produced by Salvia species. Bull. Torrey Club 91:327-330.
WEnT, F. W. 1955. The ecology of desert plants. Sci. Amer. Offprint no. 114. W. H. Freeman, San Francisco.
THE CHROMOSOMES OF SCOLIOPUS (LILIACEAE)
Marion S. CAVE
Two species of Scoliopus have been described: 1, S. dbigelova Torr., occurring in the coast ranges of California from Santa Cruz Co. north to Humboldt Co. and 2, S. hallit Wats., from the west slope of the Cascades and Coast Mountains of Oregon.
In 1932 Johansen reported 7 pairs of chromosomes in root tip mate- rial of Scoliopus bigelovii, originally obtained from the “native habitat”’ and “grown by the sand or water-culture method.” He described the karyotype as: 1 long pair of chromosomes with medium constrictions, 1 long U-shaped pair, 1 medium-long straight pair, 1 medium-long curved pair, 2 short broad pairs, 1 short narrow pair.
Recently a number of collections of S. bigelovii (60.135, Sonoma Co., Cave 62150; 60.220, Humboldt Co., Cave 6530; 61.015, San Mateo Co., Cave 6514; 61.019, San Mateo Co., Cave 6506; 63.120, Sonoma Co., Cave 6515) made by Helen Mar Beard and Wayne Roderick, and now growing at the University of California Botanical Garden in Berkeley, have been examined for chromosome number, by means of the acetocar- mine smear technique. By the time the tip of the first leaf is visible above the ground the youngest flower is already at the stage of mitosis in the microspore. Metaphase figures of this division (fig. 1) showed 8 chromosomes: 1 large (a), 2 medium-large (b and c), and 5 small. This discovery led the writer to examine some slides made in 1948 from ma- terial from Muir Woods, Marin Co., California. In all slides there were numerous examples of pollen grain metaphases with 8 chromosomes.
Pollen grains in all collections, save one, now growing at the Botanical Garden were of uniform size, with only a few scattered abortive cells, and the plants set good seed. Smears from ovules showed 16 mitotic chromosomes (fig. 2). That meiosis was normal is at least suggested. In the collections from Humboldt Co. there were also 16 mitotic chromo- somes in the root tips, but the pollen grains were of all different sizes, and many were aborted. Two grains were found showing metaphases with 9 and 10 chromosomes respectively, including the 3 large ones, but varying numbers of small ones (figs. 3, 4). Pollen grains with deviating chromosome numbers may be the result of structural hybridity leading to asynapsis or desynapsis and consequent maldistribution of chromo- somes at meiosis.
Root tips of the other species, Scoliopus hallii, received from Stanton Cook from Fall Creek, Lane Co., Oregon, had only 14 mitotic chromo- somes (fig. 5). The 3 largest chromosome pairs resembled those in S. bigelovii, but there were only 4 small pairs.
The three largest chromosomes are probably homologous with Johan- sen’s long U-shaped pair, medium-long straight pair, and medium-long
Za
210 MADRONO [Vol. 18
Fics. 1-5 Chromosomes of Scoliopus: 1-4, S. bigeloviz; 1, (6515) metaphase of division in microspore (n=8); 2 (6506) metaphase from cell of ovule (2n=16) ; 3, 4 (6530) metaphases of microspores showing extra chromosomes; 5, (65137) S. hallii, metaphase from root tip cell (2n=14). & 1660.
curved pair. His long pair with median constrictions, represented in his drawing by a thin line connecting the arms, is probably homologous with two of the short chromosome pairs found in the present collections of S. bigelovit. It would have been of interest to know the origin of his plants, because of the suggestion of structural hybridity in our collection of S. bigelovu from Humboldt Co. It is also possible that there are some popu- lations in which translocation has given rise to a long chromosome from two short ones.
The taxonomic position of Scoliopus has recently been discussed by Berg (1959; 1962). He states that it should be removed from the Pari- deae of Krause (1930) and “placed in the Melanthoideae, probably as a tribe of its own near the Tricyrteae and the Uvularieae (and the Calo- chorteae)”’. Berg accepted Johansen’s count for S. bigelovii of 2n — 14,
1966] NOTES AND NEWS 203
but stated that the difference in basic chromosome number between this genus and other members of the Parideae cannot rightly be evaluated at present. It may be pointed out here that Sat6’s (1942) figure 54 of Oake- sia sessilifolia of the Uvularieae shows 3 fairly large pairs and 5 small pairs as do our examples of Scoliopus bigelovii. This similarity of karyo- types thus substantiates Berg’s view of the close relationship of Scoliopus to the Uvularieae.
It will be of interest to see what is the number of chromosomes found in plants of both species of Scoliopus from other localities. The present study suggests that 8 is the normal haploid number for S. bigelovii and 7 for S. hallit.
Botanical Garden, Department of Botany, University of California, Berkeley
LITERATURE CITED
Berc, R. Y. 1959. Seed dispersal, morphology, and taxonomic position of Scoliopus, Liliaceae. Skr. Vid.-Akad. Oslo. I. Mat.-Nat. Kl. 1959 (4):1-56.
, 1962. Contribution to the comparative embryology of the Liliaceae: Sco- liopus, Trilljum, Paris, and Medeola. Skr. Vid.-Akad. Oslo. I. Mat.-Nat. KI. 1962 (4) :1-64.
JOHANSEN, D. A. 1932. The chromosomes of the California Liliaceae I. Am. Jour. Bot. 19:779-783.
Krause, K. 1930. Liliaceae. Jn Die natiirlichen Pflanzenfamilien. Vol. 15a.
SaTo, D. 1942. Karyotype alteration and phylogeny in Liliaceae and allied families. Jap. Jour. Bot. 12:57-191.
NOTES AND NEWS
THE OCCURRENCE OF A NEW ZEALAND PLANT, GERANIUM MICROPHYLLUM IN CALIFORNIA.—This small-flowered geranium native to New Zealand was first col- lected near Olema, Marin Co., in 1898 by Davy (4342, UC). Eastwood (Erythea 6:117. 1898) identified it as G. szbiricum L. and indicated that it was “abundant on the Abbott Ranch, at the entrance to Bear Valley.”” Howell (Marin Flora. 1949) annotated it as “perhaps a shade form of G. pilosum Forst. f.”. A more recent col- lection was made in this same area (McHoul, May 3, 1965, UC) and has been identified as G. microphyllum Hook. f. Plants in this population numbered in the hundreds. Geranium microphyllum differs from G. pilosum and G. retrorsum L’Her. in having 1-flowered pedicels, white flowers with edges faintly tinged with pink, short, thickened roots, not tap roots, almost tuberous in character, and the caudex not branched. Two kinds of pubescence are present, long, white, slightly flattened, trichomes to 1 mm long and shorter hairs. The long trichomes are present on the edges and nerves of the subulate-tipped sepals and the shorter hairs are between the nerves. The upper sides of the leaves are dark green, sparsely covered with short trichomes. The under sides of the leaves are sparsely long and short pubescent and are grayish in color from a thick covering of minutely mealy granules—granu- lose rather than glandular. The pedicels below the flowers, the stems at nodes, and the petioles are thickly covered with long trichomes. A specimen was sent to Dr. G. Brownlie, University of Canterbury, New Zealand, who confirmed the identifi- cation. He indicated that although our specimens are larger in general than New Zealand plants, some in shady sites in scrub do approach the Olema specimens in size—MarGareET S. BERGSENG, University of California Herbarium, Berkeley.
TWO NEW SPECIES OF VERBESINA (COMPOSITAE) WITH TAXONOMIC NOTES ON OTHER SPECIES
James R. COLEMAN
While working on a revision of the genus Verbesina, I encountered two previously undescribed species of section Pterophyton which are described herein. In addition. a confused taxonomic situation was dis- covered and is reported. This is part of a Ph.D. thesis submitted to Indiana University. This work was supported by fellowships from the National Science Foundation and Indiana University and I wish to ex- press my gratitude to Charles B. Heiser who suggested this study and read the manuscript.
Verbesina apiculata Coleman, sp. nov. Type: Ocurachui, Sierra Suro- tato, Sinaloa, Mexico, Gentry 6263 (NY! ).
Herb erecta perennis ca. 0.7 m alta; caulis anguste alatus; folia alterna sessilia lanceolate-oblonga vel linearia acuta vel acuminata usque ad 8 cm longa scabra; inflorescentia corymbosa capitulis 5-6 ca. 2 cm latis praedita; phyllaria ca. 2-seriata suboblonga ca. 3.5—4.0 mm longa; radii ca. 8 styliferi; receptacula convexa; paleae acutae vel apiculatae; achenia immatura obovata ca. 3 mm longa anguste alata pappi aristis 2 ca. 0.8 mm longis praedita.
Erect herbaceous perennial ca. 70 cm tall; stems to 3 mm in diameter, puberulent, winged to near the inflorescence (wings to 1.5 mm wide) ; leaves alternate, sessile, lance—oblong to linear, serrate to serrulate, acute to acuminate, to ca. 8 cm long, ca. 1.0—-1.7 cm wide, scabrous above, hispid beneath, pinnately veined; peduncles to ca. 10 cm long; in- florescences corymbose, heads 5-6, ca. 2 cm wide; phyllaries ca. 2—seri- ate, oblong to subspatulate, ca. 3.5—4.0 mm long, obtuse or acute, black, some with tips recurved; ray florets ca. 8, styliferous, linear, ca. 1.7 cm long, hirtellous, awnless; disc florets ca. 7 mm long, corolla sparingly hirsute, ovary ca. 2.5 mm long, upwardly hirsute, awns ca. 0.8 mm long, style tips attenuate; receptacle convex; chaff ca. 5.5 mm long, hirsute, acute or apiculate, black apically; disc achenes (immature) obovate, ca. 3 mm long, 1.5 mm wide, the crown with a slightly thickened ridge, the wings ca. 0.2 mm wide, the margins ciliate, the awns 2 ca. 0.8 mm long, (fig. 1).
Verbesina apiculata is known only from the type material which the collector states as having been collected at about 6000-7000 feet in a pine forest area in a steep, moist, shady canyon with mixed dominants. The closest affinities of V. apiculata are with V. roset from which it differs by its alternate, scabrous leaves and black phyllaries as well as by its styliferous ray florets. The leaves of V. rosei are opposite and strigillose. Verbesina rosei occurs about 350 miles SSE of the type locality of V. apiculata in the state of Nayrait.
214
1966] COLEMAN: VERBESINA JANG)
fl , jl bed QS, of > tay G
4a, Ee e _ ‘
ey Ay. Q
itd hay Nh -
pint pr)
R
Fic. 1. Verbesina apiculata: A, habit sketch, X ca. ¥%3; B, ray floret, x 4.5; C, chaff, < 7; D, disc floret, < 7.5.
Verbesina tequilana Coleman, sp. nov. Type: rocky hillsides, Te- quila, Jalisco, Mexico, Palmer 377 (NY!, isotype, NY!).
Herba erecta perennis; caulis alatus usque ad capitulum; folia plerumque opposita sessilia elliptica acuta ca. 5-8 cm longa penninervia scabra; capitulum ca. 4.5 cm latum; phyllaria 2—3—-seriata oblonga ca.
216 MADRONO [Vol. 18
Fic. 2. Verbesina tequilana: A, habit sketch, x ca. ¥%3; B, disc floret, & 5.5; C, chaff, < 5.
5-8 mm longa marginibus revolutis praedita; radii ca. 10 styliferi; ovaria ca. 4 mm longa pappi aristis 2 ca. 1.5 mm longis praedita.
Erect herbaceous perennial exceeding 1 m. tall; stems to 3 mm in diameter, winged to the heads (wings ca. 2.5 mm wide), sparingly
1966] COLEMAN: VERBESINA AN)
hirsute; leaves mostly opposite, becoming alternate above, sessile, ellipti- cal, serrate, acute, ca. 5-8 cm long, ca. 2.5—4.0 cm wide, quite scabrous, pinnately veined; peduncles ca. 2 cm long; inflorescences 1—2—headed, heads ca. 4.5 cm wide, discs ca. 2 cm wide; phyllaries 2—3—-seriate, ob- long, obtuse, margins revolute, the outer series ca. 5—7 mm long, the inner series ca. 8 mm long; ray florets ca. 10-12, styliferous, ca. 1.7 cm long, corolla sparingly hirsute, oblong, ovary ca. 2.5 mm long, awnless; disc florets ca 11 mm long, style tips attenuate, ovary ca. 4 mm long with marginal wings developing, the awns 2 ca. 1.5 mm long; chaff ca. 7—8 mm tall, acute (fig. 2).
Verbesina tequilana is known only from its type material which un- fortunately lacks achenes. It is quite closely allied to V. tetraptera (Orteg.) Gray from which it is distinguishable by its sessile, elliptical, pinnately veined leaves and its fewer (ca. 10-12 vs. 15-25) and stylifer- ous ray florets. The generally rhomboid-hastate leaves of V. tetraptera, although varying greatly in shape, never approach being elliptical and are always distinctly petiolate and strongly triple nerved. The type locality of V. tequilana is approximately 200 miles NW of the known range of V. tetraptera.
The type of V. tequilana was referred by Robinson and Greenman (1899) to V. scabra Benth. which they placed in section Alatipes. They considered the specimen as being “‘surely distinct from V. tetraptera” and identified it with Bentham’s description and a drawing by Klatt. However an examination of an isotype and a photograph of the holo- type permits no doubt of that species’ being identical with V. tetraptera. Blake (1930), commenting on this situation, said of Palmer 377, “it may represent a distinct species.” However he failed to describe it as such.
In their introduction to section Pterophyton Robinson and Greenman (1899) refer to V. ovata, but do not mention the name further except as a synonym of V. pterocaula (Moc. & Sess.) DC. to which they refer Delessert t. 597 and Seaton 340. A comparison of a photograph of Deles- sert’s drawing of Mocino and Sessé 30728 with Seaton 340 revealed that the two are certainly not of the same species. Mocifio and Sessé’s speci- men is depicted as having styliferous ray florets, the ovaries bearing two well developed awns. Also, the leaves are linear-lanceolate with acuminate tips. Seaton 340 contrasts with this by having neutral ray florets, the ovaries of which are awnless, as well as by having acute or obtuse, oblong or elliptic-oblong leaves. Seaton 340 fits well enough the description and illustration of Coreopsis ovata Cav. which Gray later correctly trans- ferred to Verbesina. Robinson and Greenman erred in applying the name V. pterocaula (Moc. & Sess.) DC. to Seaton 340 rather than V. ovata (Cav.) Gray and in citing the two specimens together.
Verbesina pterocaula is however a legitimate species which fits well into section Verbesinaria. A comparison of Mocino and Sessé 30728 with the type of V. stenophylla Greenman. (Pringle 6503), which Robinson
218 MADRONO [Vol. 18
and Greenman placed in section Verbesinaria, revealed no significant differences. Since V. pterocaula has priority, it is necessary that V. stenophylla be reduced to a synonym of that species.
Department of Botany, Indiana University, Bloomington (Present address: Instituto de Botanica, C. P. 4005, Sao Paulo, Brazil)
LITERATURE CITED
BLAKE, S. F. 1930. Notes on certain type specimens of American Asteraceae in Euro- pean Herbaria. Contr. U.S. Natl. Herb. 26:227-263.
Rosinson, B. L. and J. M. GrEENMAN. 1899. A synopsis of the genus Verbesina, with an analytical key to the species. Proc. Am. Acad. 34:534—564.
OBSERVATIONS ON WITCHES’-BROOM FORMATION, AUTOPARASITISM, AND NEW HOSTS IN PHORADENDRON
FRANK G. HAWKSWORTH AND DELBERT WIENS
During the course of field studies on Arceuthobium in the southwestern United States and Mexico, we also had the opportunity to make a num- ber of collections of the related genus, Pkoradendron. This report is a compilation of our observations on Phoradendron, including witches’- broom formation, autoparasitism, and some new hosts.
This research was supported, in part, by a cooperative agreement be- tween the Rocky Mountain Forest and Range Experiment Station and the University of Coolrado where the junior author was located during the conduct of the study. Herbaria abbreviations used here but not occurring in Index Herbariorum include: FPF—Forest Pathology Herbarium, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado, and INI F—Instituto Nacional de Investigaciones Forestales, Mexico, D.F.
WITCHES’-BROOMS
Witches’-broom formation is a common symptom of conifers infected by Arceuthobium, but has not been reported previously to be associated with parasitism by Phoradendron. The only other reported instance of witches’-brooms caused by a member of the Loranthaceae is in Chile where brooms are induced by Phrygilanthus tetrandrus Eichler on Popu- lus (Reiche, 1907; Kuijt, 1964). This report is verified (fig. 1) from collections made in Santiago, Chile (Wiens 3833 UT). Brooms in Populus consisted of masses of vertical branches that rose 5 to 10 ft. No broom formation was observed on native hosts attacked by this mistletoe.
1966] HAWKSWORTH & WIENS: ARCEUTHOBIUM 219
Fic. 1. Witches’-broom on Populus induced by Phrygilanthus tetrandrus, Santi- ago, Chile.
In an area two miles south of La Cuesta along Highway 45, Durango, Mexico, we found P. tomentosum (DC.) Engelm. ex Gray ssp. tomen- tosum on several ocotillo plants, Fouquieria fasciculata (Roem. & Schult.) Nash (Wiens 3294, COLO; Hawksworth 324, FPF). The mistletoe was common nearby on Forestiera, and within a few miles on Prosopis and Celtis. The mistletoe plants on ocotillo were small (2 to 3 inches high), pale yellow, and generally of low vigor compared with those on the other hosts. Symptoms of infection included markedly swollen branches and formation of witches’-brooms (fig. 2).
During a recent trip to southeastern Utah, R. S. Peterson (pers. comm.) noted several massive witches’-brooms in Utah juniper, Juni- perus osteosperma (Torr.) Little. His examinations showed that they were not of the type caused by rust fungi but, surprisingly, were associ- ated with P. juniperinum Engelm. (fig. 3). The foliage in the broomed part of the tree was denser and darker green than in the non-broomed parts of the tree. Plants of P. juniperinum were distributed systemically throughout the brooms (Peterson 65-80, FPF) in a manner somewhat similar to parasitism of pines by certain dwarf-mistletoes, Arceuthobium
220 MADRONO [Vol. 18
(Hawksworth, 1961; Kuijt, 1960). Non-systemic infections of P. juni- perinum occurred both within the broomed and non-broomed parts of the tree. This is in contrast to Arceuthobium where non-systemic infec- tions rarely occur on systemic brooms. Whether P. juniperinum actually stimulated development of latent or adventitious buds was not deter- mined. The broom portions where the parasite developed systemically, however, had a dense, intricately branched live twig system. The non- broomed parts, conversely, possessed a sparsely branched, open branch system. Peterson has also observed similar brooms associated with P. juniperinum on J. oteosperma and J. scopulorum Sarg. at Bryce Canyon, Utah.
INTRA- AND INTER-SPECIFIC PARASITISM
Autoparasitism, i.e., the occurrence of one mistletoe on a different individual of the same species, has been reported for P. serotinum (Raf.) Johnston in Florida (Curtiss, 1878) and also for several tropical Ameri- can mistletoes (Kuijt, 1964; Wellman, 1964). We have observed auto- parasitism in P. californicum Nutt. near Tucson, Arizona, on the Mount Lemmon highway (Hawksworth & Lightle 174, FPF). We have also noticed autoparasitism in P. tomentosum ssp. tomentosum on Highway 40 about 50 miles east of Torreon, Chihuahua, Mexico (Wiens 2616, RSA) and in P. villosum (Nutt.) Nutt. ssp. villosum on Highway 30 3 miles south of Running Springs in the San Bernardino Mountains, California Adawksworth 737, FPF).
The occurrence of mistletoes on other species and genera of mistletoe is common in the tropics (Kuijt, 1964; Wellman, 1964), but not in more temperate regions. The only previous instance that we are aware of in the United States is Brown’s (1918) report of P. californicum on P. tomentosum (DC.) Engelm. ex Gray ssp. macrophyllum (Engelm.) Wiens in Arizona. In an area 3 miles east of the Mimbres River on Highway 180 in Grant Co., New Mexico, P. juniperinum was collected on Juniperus deppeana Steud. A closer examination of the collection (Hawksworth & Lightle 124a, FPF) revealed that an individual of the P. juniperinum was, in turn, parasitized by P. villosum ssp. coryae (Trel.) Wiens. The latter plant was young and had shoots only about one inch high. Typical individuals of P. villosum ssp. coryae were com- mon in the vicinity on various oaks.
NEw AND UNUSUAL HostT-PARASITE COMBINATIONS
Phoradendron californicum Nutt. This mistletoe is a characteristic plant of the deserts of the southwestern United States and northwestern Mexico. Its hosts are principally leguminous shrubs (Wiens, 1964).
Harris et al. (1930) reported that the possible explanation for the “exceedingly rare’ occurrence of P. californicum on creosote bush, Larrea divaricata (DC.) Coville, was that this shrub has a much higher osmotic concentration than the usual hosts for this mistletoe. The only case of parasitism of this shrub by P. californicum that they found was
1966 | HAWKSWORTH & WIENS: ARCEUTHOBIUM 221
Fic. 2. Witches’-broom on Fouquieria fasciculata caused by Phoradendron tomentosum subsp. tomentosum. Note hypertrophy and change in tropic response of branches on the broom. La Cuesta, Durango, Mexico.
at Sacaton, Arizona, near a pumping station where the soil moisture was abnormally high.
Our observations, and the listing of Larrea divaricata as an ‘“‘oc- casional” host by Munz (1959), indicate that parasitism of this shrub is not as rare as the earlier reports suggested. We have found this host- parasite combination north of Amboy, San Bernardino Co., California, (Wiens 967, RSA) and in several localities in Arizona: Pinal Co., 10 miles north of Oracle Junction (Hawksworth 706, FPF) and near Apache Junction (Guill in 1965, FPF); Gila Co., 24 miles northwest of Globe (Lighile & Lampi 65-12, FPF) and Mohave Co., Black Range (Hawksworth & Peterson 723, FPF).
Similarly, L. S. Gill (pers. comm.) informs us that P. californicum is common on creosote bush in an area 11 to 15 miles south of San Felipe on the road to Puertecitos, Baja California, Mexico. He noted that wherever legumes were heavily infected, the associated creosote bushes were similarly infected (Gill, 1965, FPF). Creosote bush was not in- fected where it grew in pure stands or in association with legumes that were not attacked by the mistletoe.
Phoradendron bolleanum (Seem.) Eichler ssp. bolleanum. This mistle- toe of northern Mexico exhibits “double specificity” that is unusual among phanerogamic parasites. Its principal hosts include a gymnosperm (Juniperus) and an angiosperm (Arbutus). We know of no other in-
222 MADRONO [Vol. 18
stance where a mistletoe is essentially restricted to two such diverse genera. The occurrence of a Mexican mistletoe on Arbutus has been known for decades. Trelease (1916) questioned whether the same mistle- toe occurred on such unrelated genera; however, Fosberg (1941) and Wiens (1964) stated that they were morphologically the same taxon. Our more recent observations in the Sierra Madre Occidental of Chi- huahua and Durango confirm this. Usually Arbutus and Juniperus are infested with about equal frequency, although an occasional individual of Arctostaphylos is also attacked. Its occurrence on Arctostaphylos is perhaps not unexpected because of the close relationship of this genus to Arbutus. Cross inoculation experiments might be desirable to preclude the possibility that the populations on Arbutus and Juniperus are not morphologically indistinguishable physiological races.
Phoradendron bolleanum ssp. pauciflorum (Torr.) Wiens. This mistle- toe is a common parasite of Abies concolor (Gord. & Glend.) Lindl. in Baja California, California, and in the Santa Catalina Mountains in Arizona (Wiens, 1964). In the Sierra San Pedro Martir of Baja Cali- fornia we found it rarely parasitizing Cupressus arizonica var. montana (Wiggins) Little (Hawksworth & Scharpf 763, DS, FPF, INIF, UT). The mistletoe was present in the vicinity on Abies.
Phoradendron bolleanum ssp. densum (Torr.) Wiens, a mistletoe found on Cupressus and Juniperus in Arizona and California, also occurs on the lower parts of the Sierra San Pedro Martir on Juniperus cali- fornica Carr. However, our observations indicate that here P. bolleanum ssp. densum is not sympatric with Cupressus, which occurs about 2,000 to 3,000 feet higher.
Phoradendron villosum (Nutt.) Nutt. ssp. villosum. This taxon is the common oak-infesting mistletoe of the Pacific Coast States, ranging from northern Baja California to central Oregon west of the Cascades (Wiens, 1964). In addition to oaks, it sometimes parasitizes Umbellu- laria, Aesculus, Arctostaphylos, Castanopsis, Fraxinus, Adenostoma, (McMinn, 1951) and Rius when these grow in association with infected oaks.
In Baja California, this parasite occurs on two host species apparently not previously recorded in the literature, although both genera are in- fected in California. Along the road east of Rancho San Jose, on the approach to the Sierra San Pedro Martir, we collected this mistletoe on Adenostoma sparsifolium Torr. (Hawksworth & Scharpf 754, DS, FPF, INIF) and on Rhus ovata Wats. (Hawksworth & Scharpf 757, DS, FPF, INIF). This mistletoe also occurs on the latter host 72 miles south of Ensenada on Mexico Route 1 (Lightle & Gill 64-36, DS, FPF, INIF). Several infected shrubs of both species were observed, sometimes at considerable distances from the closest infected oaks.
Phoradendron villosum ssp. coryae (Trel.) Wiens. This is the com- mon mistletoe on oak in the southwestern United States and adjacent northern Mexico (Wiens, 1964). Along Highway 666, 16 miles north
1966] HAWKSWORTH & WIENS: ARCEUTHOBIUM 223
Fic. 3. Witches’-broom on Juniperus osteosperma associated with Phoradendron juniperinum. The left branch is a massive broom about 8 feet across. Twenty-six miles west of Blanding, San Juan Co., Utah. Photograph by R. S. Peterson, U. S. Forest Service.
of Clifton, Greenlee Co., Arizona, P. villosum ssp. coryvae was found on several bushes of Condalia globosa Johnston var. pubescens Johnston (Hawksworth & Lightle 138, FPF). The parasite was common nearby on associated Quercus turbinella Greene.
Phoradendron juniperinum Engelm. ssp. juniperinum. This mistletoe is a common parasite of Juniperus from Oregon and Colorado south to northern Mexico (Wiens, 1964). Hedgcock (1915) noted that it had been “reported on Cupressus arizonica,” although he had not seen col- lections of it on this host. We have examined the following specimens of P. juniperinum ssp. juniperinum on Cupressus arizonica: Cochise Co., Chiricahua National Monument (Gill FP 89611, FPF) and Chiricahua Mountains, Rucker Canyon (Hawksworth & Peterson 803 FPF). Mexico: Chihuahua, 38 miles southwest of Matachic on Ocampo Road (Wiens 3439, COLO and Hawksworth 482, FPF, INIF).
This mistletoe also occurs occasionally on Cupressus bakeri Jeps. in California: Plumas Co., 1.5 miles west of Wheeler Peak, east of Green- ville (Quick 62-93, CAS); Shasta Co., 2 miles east of Burney Springs
224 MADRONO [Vol. 18
(Peterson 65-458, FPF). In the latter locality, P. juniperinum was asso- ciated with P. bolleanum ssp. densum.
Rocky Mountain Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture, Colorado State University,
Fort Collins, Colorado
Department of Botany, University of Utah, Salt Lake City
LITERATURE CITED
Brown, J. G. 1918. Mistletoe vs. mistletoe. Bot. Gaz. 65:193.
Curtiss, A. H. 1878. Mistletoe parasitic upon itself. Bot. Gaz. 3:36-37.
FosBeErG, F. R. 1941. Notes on Mexican plants. Lloydia 4:274-290.
Harris, J. A., Harrison, G. J. and Pascog, J. A. 1930. Osmotic concentration and water relations in the mistletoes, with special reference to the occurrence of Phoradendron californicum on Covillea tridentata. Ecology 11:687—702.
HAwxkswortTH, F. G. 1961. Dwarf mistletoe of ponderosa pine in the Southwest. U.S. Dept. Agr. Tech. Bull. 1246.
Hepccock, G. G. 1915. Notes on some diseases of trees in our national forests. V. Phytopathology 5:175—181.
Kurt, J. 1960. Morphological aspects of parasitism in the dwarf mistletoes (Ar- ceuthobium). Univ. Calif. Publ. Bot. 30:337-436.
. 1964. Critical observations on the parasitism of new world mistletoes. Canad. Jour. Bot. 42:1243-1278.
McMiny, H. E. 1951. An illustrated manual of California shrubs. Univ. Calif. Press, Berkeley.
Mowz, P. A. 1959. A California Flora. Univ. Calif. Press, Berkeley.
REIcHE, K. F. 1907. Bau und Leben der hemiparisitchen Phrygilanthus-Arten Chiles. Flora 97:375—400.
TRELEASE, W. 1916. The genus Phoradendron. Univ. Illinois Press, Urbana.
WeE.timan, F. L. 1964. Parasitism among neotropical phanerogams. Ann. Rev. Phytopath. 2:43-56.
Wiens, D. 1964. Revision of the acataphyllous species of Phoradendron. Brittonia 16:11-54.
NOTES AND NEWS
The Hafner Publishing Company, 31 East 10th Street, New York, New York 10003, has recently reprinted several books of interest to botanists. The photographic reproductions are of a very good quality and the cloth bindings are stout. Among the facsimile editions are: Flora of Bermuda, by N. L. Britton, $12.00; Flora of the Hawaiian Islands, by WILLIAM HILLEBRAND, $20.00; Liberty Hyde Bailey, by A. D. Ropcers, $10.00; Flora of the Prairies and Plains of Central North America, by P. A. Rypperc, $12.00; Life of Mendel, by Huco Ittts, $5.75; Taxonomic Terminology of the Higher Plants, by H. I. FEATHERLY, $4.25; and Plant Sociology, the Study of Plant Communities, by J. BRAUN-BLANQUET, $8.00.
MADRONO
VOLUME 18, NUMBER 8 OCTOBER, 1966
As TAN
y Fay mene
Contents
CHRYSOTHAMUS BOLANDERI, AN INTERGENERIC HYBRID, Loran C. Anderson and Jack L. Reveal
A NEw SPECIES OF BRODIAEA FROM SAN CLEMENTE _ Istanp, Catirornia, Theodore F. Niehaus CHROMOSOME NUMBERS IN THE ERIOPHYLLUM LANATUM (COMPOSITAE, HELENIEAE) CompLex, John S. Mooring
PROPAGATION OF SOME Woopy DESERT PERENNIALS BY STEM CuTTINGS, Valerie C. Chase and B.R. Strain
NOTES ON THE HORDEUM JUBATUM COMPLEX, Tibor Rajhathy
DOCUMENTED CHROMOSOME NUMBERS OF PLANTS
Reviews: The Hunt Botanical Library and Its Publi- cations (John H. Thomas); Franz Schwanitz, The Origin of Cultivated Plants (Thomas W. Whitaker) ; Edmund C. Jaeger, The California Deserts (Robert M. Lloyd)
NoTEs AND News: NOTES ON SOME GRASSES OF THE Paciric Coast, Kenton L. Chambers
INDEX
A WEST AMERICAN JOURNAL OF BOTANY
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250 252
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wy MADRONO A WEST AMERICAN JOURNAL OF BOTANY
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BOARD OF EDITORS
EpcAR ANDERSON, Missouri Botanical Garden, St. Louis LymANn Benson, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis JouHN F. Davinson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California Mivprep E. Matutas, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley Marion OwnBEY, Washington State University, Pullman REED C. Roiiins, Gray Herbarium, Harvard University Ira L. Wiccins, Stanford University, Stanford, California
Editor—JoHNn H. THoMAS Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Malcolm A. Nobs, Carnegie Institution of Washington, Stanford. First Vice-President: W. B. Critchfield, Pacific Southwest Forest and Range Experiment Station, Forest Service, U. S. Department of Agriculture, Berkeley. Second Vice- President; William S. Boyle, Department of Botany, Utah State University, Logan. Recording Secretary: Mary L. Bowerman, Department of Botany, University of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Department of Bot- any, University of California, Berkeley. Treasurer: Douglas M. Post, Biology De- partment, San Francisco State College.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Annetta Carter, Department of Botany, University of California, Berkeley; the Editor of Madrofio, and three elected Council Members: William M. Hiesey, Carnegie Institution of Washington, Stanford; Herbert L. Ma- son, Department of Botany, University of California, Berkeley; and Virginia M. Page, Department of Biological Sciences, Stanford University.
CHRYSOTHAMNUS BOLANDERI, AN INTERGENERIC HYBRID
LorAN C. ANDERSON AND JACK L. REVEAL
The genus Chrysothamnus (Astereae, Compositae) is closely related to Haplopappus, and, particularly, C. bolanderi (Gray) Greene has been noted for its similarity to H. macronema Gray (section Macronema). Gray (1873) and Greene (1895) both suggested C. bolanderi could easily be placed in the other genus, and later Greene (1904) did so as Macronema bolanderi. This close resemblance to a species of Haplopap- pus makes the identification of C. bolanderi essential to an understand- ing of the systematics and evolution of Chrysothamnus. This plant has been, unfortunately, rarely collected. In addition to the earlier collec- tions: Bolander 6137 in 1866 (GH—holotype, UC!, US!) and Rattan s.n.in 1867 (DS!, mixed, H. macronema in part), it is now represented by Reveal 1057 (KSC) and Anderson & Fish 2923 and 2926 (KSC).
Bolander’s type collection is labeled: “At Mono Pass, California, elevation 9000-10000 ft.” The only plants we found after extensive in- dependent search (Reveal in 1964, Anderson in 1965) were located east of Mono Pass (elevation 10604 ft.) in Bloody Canyon where they were restricted to a small oasis surrounded by barren rock. A small population was found at 10000 ft. on a steep talus-filled crevasse which rises abruptly from the northwest edge of Lower Sardine Lake (ca. 8 air miles southwest of Lee Vining). A few more plants were found in talus along the trail just below the lake at 9800 ft. A total of 25-50 plants, then, represents the extent of C. bolanderi.
Hall (Hall and Clements, 1923) described this taxon as a relict sub- species derived from C. parryi (Gray) Greene ssp. parryi. However, recent studies (Sharp and Birman, 1963) show that five advances of upper Pleistocene glaciation can be recognized in Bloody Canyon; these are Tioga, Tenaya, and Tahoe (of the Wisconsin), Mono Basin (lIlli- noian?), and Sherwin (Kansan?). This history of recent repeated glaciation does not support the designation of C. bolanderi as a relict. An alternative, as indicated by the extremely limited range and the fact that the plants are growing in very close association with C. nauseosus (Pallas) Britt. ssp. albicaulis (Nutt.) Hall & Clem. and H. macronema, is that it is of recent hybrid origin.
To elucidate the relationship of C. bolanderi, comparative studies on gross morphology, cytology, and anatomy were undertaken. Since C. nauseosus and H. macronema both vary considerably throughout their ranges, only collections associated with C. bolanderi are dealt with here. Data collected for C. parryi ssp. monocephalus (Nels. & Kenn.) Hall & Clem. and H. suffruticosus (Nutt.) Gray, also found in Bloody Canyon, indicate they are not related to the problem.
Maprono, Vol. 18, No. 8, pp. 225-256. December 2, 1966.
229
226 MADRONO [Vol. 18
MATERIALS AND METHODS
Methods for gathering data and preparing materials have been pre- viously outlined (Anderson, 1963). Permanent slides and herbarium specimens are filed at Kansas State University. Before detailed studies were begun, Reveal’s collection, being a sampling of the entire C. bolanderi population, was segregated according to superficial resemblance into three categories by the senior author: Reveal 1057A, those plants most similar to C. nauseosus; Reveal 1057B, those most similar to the type collection (Bolander 6137); and Reveal 1057C, those most like H. macronema.
DISCUSSION
Morphology. On the talus slopes below Mono Pass, C. nauseosus is a woody shrub, 2.5—4 dm tall; C. bolanderi is less shrubby and intermedi- ate in size (2-3 dm), and H. macronema is suffruticose from a woody caudex and 1.5—2 dm tall.
Several aspects of leaf morphology are compiled in Table 1. The arbitrary trichome frequency classes are easily determined under 10 X magnification; they range in series from absent, through occasional, sparse, moderate, and abundant to very abundant. For illustration of frequency classes, see Eiten (1963). Specificity of trichome types and structure permits their use in analysis of hybrids; sometimes they may be used as the primary basis for determination of hybrids (Carlquist, 1961). With C. nauseosus having essentially only nonglandular trichomes and leaves of H. macronema glandular ones, the trichome complement in specimens of C. bolanderi gives an excellent index to degree of inter- gradation. Leaves of C. nauseosus do have a few glandular hairs, but they are not visible under 25 magnification because of the tomentum. Nonglandular trichomes on leaves of Bolander 6136 (H. macronema) are distributed evenly over the leaf surface, wheras in Reveal 1052 they are restricted to the midribs.
Leaf size and shape are also useful as indicators of the intermediacy of C. bolanderi. Leaves of plants most like H. macronema (Reveal 1057C and Anderson & Fish 2926) are larger than those found in either puta- tive parental populations. Leaves of H. macronema are unique in having undulate margins.
Several characteristics of floral morphology in Chrysothamnus have been demonstrated to be of diagnostic value (Anderson, 1946). Many which apply here are tabulated in Table 2 (omissions due to immaturity of specimens). Other features are discussed in the text.
The inflorescence of C. nauseosus is densely corymbose; that of C. bolanderi contains fewer heads, and H. macronema has a few or some- times only a single head in the inflorescence. Heads are illustrated for the three taxa in Fig. 1. Involucre height and especially bract number indicate intergradation; further indications are found in bract align- ment and structure. Bracts of C. mauseosus are in pronounced vertical ranks (a key character for the genus), keeled, and chartaceous. Those of
ANDERSON & REVEAL: CHRYSOTHAMNUS ITT]
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228 MADRONO [Vol. 18
C. bolanderi are in obscure vertical ranks, less prominently keeled, and more membranous. Bracts of H. macronema are not vertically aligned; they lack keels, and the outermost are green, leaf-like, and covered with glandular trichomes (not shown in Fig. 1).
Elsewhere throughout its range C. nauseosus is consistently 5-flow- ered; therefore, in the case of Anderson & Fish 2924, there is evidence of introgression. In addition to flower number, shape of the corolla tubes also suggests intergradation; they gradually flare out in C. nauseosus, are somewhat dilated in C. bolanderi, and more noticeably so in H. macronema. The collections most like H. macronema in gross appear- ance and trichome frequencies are, interestingly, more like C. nauseosus in floral morphology.
Cytology. Chromosome numbers as determined from pollen mother cells in meiosis are listed in Table 2; in each taxon, n = 9. Chromosomal pairing and other aspects of meiosis appear to be regular in C. bolandert. Pollen fertility as determined by stainability in analine blue in lacto- phenol is also listed in Table 2. Apparently normal meiosis and notice- ably reduced pollen fertility also characterize the only other presumed hybrid known in Chrysothamnus (Anderson, 1966). The relatively high pollen fertility in Reveal 1057A is possibly due to back-crossing with C. nauseosus ; this conclusion is supported by many similarities in mor- phology as reported here. Backcrossing with H. macronema is also indi- cated; the reduced pollen fertility of Bolander 6136 suggests less compatibility between this parent and C. bolanderi. However, these vari- ations in fertility and morphology could also be due to genic recombi- nations in the F».
The extent of viable seed formation in C. bolanderi has not been determined. Only one seed of Reveal 1057C successfully germinated, and the seedling died a few days later.
Anatomy. Vegetative anatomy, excluding features of the wood, is similar in most respects among the three taxa. The amounts of primary phloem sclerenchyma tend to vary according to topography and age of the sample; nevertheless, phloem fibers appear to be most abundant in stems of H. macronema and least abundant in C. nauseosus, with C. bolanderi appearing intermediate. Quantative differences in trichome distribution have already been noted; in addition, there are differnces in trichome size and structure. On stems of all samples there are uniseri- ate nonglandular hairs that are anisotropic under polarized light and biseriate glandular hairs that are isotropic. On stems of C. nauseosus the long uniseriate trichomes that form the tomentum cover the less frequent biseriate glandular hairs, the longest glandular hairs being only 9.8 » long. In C. bolanderi, the longest glandular trichomes are 12.7 and 13.2 uw (Anderson & Fish 2923 and 2926 respectively); whereas on stems of H. macronema the longest glandular trichomes are 32.2 w.
A similar correlation exists for foliar trichomes with one interesting modification. On leaves of C. nauseosus the longest glandular trichomes
ANDERSON & REVEAL: CHRYSOTHAMNUS 229
1966]
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21746) £86
% pollen fertility
gametic chromosome number
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0°6 IES | v'0Z O71 12-01 St (OSM) 2262 YStq GD UOsSsapuy O'OT ¢7-OI 6 +1 OTT 6I-¢1 VST (OSM) ZS0I Jva0ay OT 61-8 SET wal SI-II 61 (OW) 9E19 4apunjog DUIUOAIDWU ‘H €I-Ol Sa EE O'OT Hao L Yl (OSM) £262 YSty D Uossapup O'OI Ota OTT O'OT 6I-IT eal (SQ) ZEl9 sapunjog 98 jee? O'OT O'+rT Glee Sel (OSM) ILSOL 12900y CTS Z'8 vI-IT UL (OSM) 9262 YStY BD Uossapuy Z'OT 6-9 vl O71 stash Col (OSM) ILS0I Jvan2y C8 6-9 Li, S'6 I-21 9°71 (OSM) VZS0I Jbaaay 1AdPUDIOG *) L-9 a) ¢I-I1 Cor (OSM) £262 YSlY | Uossapuy OTT S S vO1 Cio OTT (SQ) 8EL9 szapunjog TOT S S C8 II-6 O'OT (OSM) OS0L 10000y SNSOIASHDU no) uw uw e| w uM | a g 3 = 8 g UOTIITJOI pue UOXR], e = & ae S = ao =) =} = oO = = = q q os q q fe a ce a 3 ~ ae) o = = nie = x 5 g : SE : : 2 oa a a S| Z i S a a S a ee Loum o ra i oA s Be © a = > a a (50) a pan ss (5°)
YAMOTY GNV AMON TOANT AHL JO SAYNLVAY AAILVAVdWO,) °C ATAV EE
230 MADRONO [Vol. 18
Fic. 1. Representative heads: a, C. nauseosus; b, C. bolanderi; c, H. macronema.
measure 7.3 » and on C. bolanderi, 10.4 and 12.0 uw. The anisotropic uniseriate, nonglandular trichomes characteristic of leaves of the pre- ceding are converted on H. macronema leaves into uniseriate glandular trichomes which are isotropic. These uniseriate glandular hairs are equally as long as the biseriates, the longest measuring 25.0 p.
Selected characteristics of the wood anatomy for the three taxa are summarized in Table 3. The plus symbol indicates the feature is abun- dantly present, and the minus, present but not abundant. In nearly all features listed in Table 3, C. bolanderi is intermediate. Additional xylem features either indicate the possible hybridity of C. bolandert or demon- strate it is more like Chrysothamnus than Haplopappus.
Vessels in the Chrysothamni are angular as seen in transection and appear in large groups of 10 or more; whereas in H. macronema, they are roughly circular in transection and average 4.09 per group. Terminal elements of a year’s growth are vascular tracheids in each case, but in H. macronema they are less abundant; this coupled with other cellular patterns makes age determination of H. macronema wood difficult. Uniseriate rays are not frequent in woods of these two genera (Carlquist, 1960); in our samples they are occasional in H. macronema, less fre- quent in C. bolanderi, and extremely rare in C. nauseosus. Axial wood parenchyma is mostly vasicentric and scanty, being most abundant in C. nauseosus and least abundant in C. bolanderi.
Most features of floral anatomy are similar in the taxa; however, two are of significance. These are relative abundance and distribution of sclerenchyma in the phyllaries and distribution of secretory canals in the achenes and corollas. Sclerenchyma is less abundant in the phyllaries
1966] ANDERSON & REVEAL: CHRYSOTHAMNUS 2311
average number of rays per mm of tangential me = = N A 1ve) section average number of cells 6 ae of maximum multiseriate = oOo + ray width ray cells isodiametric l Tn ts to erect ray cells isodiametric Seeee Tee to procumbent 8 S$ average height of Oe = multiseriate rays, mm | sn) & fo ) ty st m~ O e B average length eg eee: SI of vessels, u SY oS sk fj Gy [<a] > i= & < . si average diameter N WO = of vessels, a ee S) O 6 e = diameter of widest A nA A H vessel, / ee ec re
Anderson & Fish 2924 (KSC) Anderson & Fish 2922 (KSC)
C. bolanderi Anderson & Fish 2923 (KSC)
Anderson & Fish 2926 (KSC)
H. macronema
Taxon and collection
C. nauseosus
232 MADRONO [Vol. 18
of H. macronema as compared to the Chrysothamni, C. bolanderi again being intermediate. Secretory canals are absent in achenes and flowers of H. macronema. They are equally abundant in achenes and style branches in our Chrysothamnus samples, but in the corollas they are more extensive in C. nauseosus, where they are in the distal portion of the corolla tube as well as in the corolla lobes as in C. bolanderi.
CONCLUSIONS
Absolute proof of hybridity, the successful synthesis of C. bolanderi from artificially crossing C. nauseosus and H. macronema, is lacking. Nevertheless, sufficient circumstantial evidence has been amassed from field observations and studies on morphology, cytology, and anatomy to strongly indicate that C. bolanderi is indeed a hybrid swarm. Back- crossing, particularly with C. nauseosus, is also indicated. In nearly all features, C. bolanderi is intermediate; however, in some collections, fea- tures such as taller wood rays and larger leaves and involucre suggest a complementary genic action in the hybrids.
Secondly, floral anatomy and especially the wood anatomy of the hybrids is more like that of Chrysothamnus than of Haplopappus. There- fore, if the genera are treated as distinct, the hybrids should be placed in Chrysothamnus. The overall appearance of C. bolanderi is that of a subspecies of C. parryi. In fact, it is currently treated as C. parryi ssp. bolanderi (Gray) Hall & Clem. in all floras. Nevertheless, since infra- specific taxa of C. parryi are not involved in the origin of this taxon, it is better referred to as C. bolanderi.
The hybrid origin of C. bolanderi was discovered in preparation of a monograph on Chrysothamnus by the senior author. The taxonomic status of the genus in light of this intergeneric hybridization will be considered in the moongraph.
Contribution no. 685, Department of Botany and Plant Pathology, Kansas Agr. Expt. Station, Manhattan, Botany serial no. 849. This study supported by National Science Foundation grant GB3058 to the senior author.
Department of Botany, Kansas State University, Manhattan U.S. Dept. Agriculture, Forest Service, Lee Vining, California
LITERATURE CITED
ANDERSON, L. C. 1963. Studies on Petradoria (Compositae): anatomy, cytology, taxonomy. Trans. Kan. Acad. 66:632—684. . 1964. Taxonomic notes on the Chrysothamnus viscidiflorus complex (Astereae, Compositae). Madrono 17:222—227. . 1966. Cytotaxonomic studies on Chrysothamnus (Astereae, Compositae). Am. Jour. Bot. 53:204—212. CarLouist, S. 1960. Wood anatomy of Astereae (Compositae). Trop. Woods 113:54-84. . 1961. Comparative plant anatomy. Holt, Rinehart, and Winston, New York.
1966] NIEHAUS: BRODIAEA 238
E1Ten, G. 1963. Taxonomy and regional variation of Oxalis section Corniculatae. I. Introduction, keys and synopsis of species. Am. Midl. Nat. 69:257—309. Gray, A. 1873. Notes on Compositae and characters of certain genera and species,
etc. Proc. Am. Acad. 8:631-661. GREENE, E. L. 1895. Observations on Compositae. X. Erythea 3:107-115. . 1904. New plants from middle California. Leafl. Bot. Obs. Crit. 1:73-81. Hatt, H. M. and F. E. Crements. 1923. The phylogenetic method in taxonomy. The North American species of Artemisia, Chrysothamnus, and Atriplex. Carnegie Inst. Publ. 326. SHarP, R. P. and J. H. Birman. 1963. Additions to classical sequences of Pleistocene glaciation, Sierra Nevada, California. Geol. Soc. Am. Bull. 74:1079-1086.
A NEW SPECIES OF BRODIAEA FROM SAN CLEMENTE ISLAND, CALIFORNIA
THEODORE F. NIEHAUS
The island of San Clemente off the southern California coast is noted for its high number of endemics (Raven, 1963). A new species here reported raises the total number of endemics on San Clemente I. to eleven.
Brodiaea kinkiensis Niehaus, sp. nov. Planta ex cormo tunica fibrosa instructo; perianthii tubo 12 mm longo, 4-5 mm lato, basi rotundato, ejus segmentis patentibus 13-17 mm longis, eis exterioribus oblongis, interioribus obovatis; staminodia paulo involuta, 3 mm lata, 7 mm longa, apice cuspidata, erecta, a staminibus distantia; capsula oblonga; perianthii tubo demum haud fisso.
Corm with heavy fibrous outer coat; leaves linear, 2-4 dm long; scape 2-3 dm tall; pedicels 3-8 cm long; perianth-tube whitish with brown- purple midribs extending to tips of the segments, rounded at the base, 12 mm long and 4-5 mm wide; perianth-segments violet, 13-17 mm long, spreading at right angles to perianth-tube; outer segments oblong, inner ones obovate; staminodia slightly involute, 3 mm wide, 7 mm long, apically cuspidate, standing erect and well apart from stamens; fila- ments 1 mm long, anthers retuse, channeled on back, 4-5 mm long; capsule oblong; perianth-tube not splitting as capsule matures; chro- mosome number 2n = 32.
Holotype. 0.5 mi W of Stone at junction of road to Stone with main island axis road, San Clemente I., Los Angeles Co., 1395 ft., April 3, 1965,T. F. Niehaus 407 (UC, 1200400).
Brodiaea kinkiensis is known at present only from San Clemente I. One previous and fragmentary collection was cited as B. filifolia Wats. by Hoover (1939) (Murbarger 219, UC). It is probable that B. kinkien-
234 MADRONO [Vol. 18
Fic. 1. Brodiaea kinkiensis: a, habit, x ¥%; b, individual flower, X 2; c, overhead view, X 2; d, interior arrangement, X 2.
1966] NIEHAUS: BRODIAEA 235
sis is most closely related to B. filifolia. Brodiaea kinkiensis may be distinguished from B. filifolia by the very coarse and thick fibrous outer coat of its corm, its longer perianth-tube (that of B. filifolia being 6—7 mm in length), and its erect cuspidate much wider staminodia (those of B. filifolia being much shorter and virtually obsolescent and capillary). Continued research on the probable relationships and on other questions is under way for the entire genus.
Brodiaea kinkiensis was collected by the author and Wayne Roderick, April 1—4, 1965, at which time it was in bud. Corms were grown at the University of California Botanic Garden where I ultimately obtained flowering specimens. This species was observed to be common in the clay flats on the mesa of the island from near Lemon Tank south to Boulder, a distance of about 4 miles. It undoubtedly occurs farther to the north and south along the mesa-top. A chromosome count of 2n = 32 (Niehaus, 1965) was reported (as B. filifolia) for B. kinkiensis. This was previous to my having seen living material of the mainland species, B. filifolia, and having compared it with that growing on San Clemente Island. As of this date, B. filifolia now remains the only species of Brodiaea (as defined by Hoover) for which a chromosome count still remains to be made.
The specific epithet is based on the word ‘“‘kinki,”’ the Gabrielino Indian name for San Clemente I. (Kroeber, 1925).
Department of Botany, University of California, Berkeley
LITERATURE CITED
Hoover, R. F. 1939. A revision of the genus Brodiaea. Am. Midl. Nat. 22:551—574.
KroesBeEr, A. L. 1925. Handbook of the Indians of California. Bureau Am. Ethnology, Smithsonian Inst., Bull. 78.
NieHAus, T. F. 1965. In Documented chromosome numbers of plants. Madrono Set 23.
Raven, P. H. 1963. A flora of San Clemente Island, California. Aliso 5:289-347.
CHROMOSOME NUMBERS IN THE ERIOPHYLLUM LANATUM (COMPOSITAE, HELENIEAE) COMPLEX
JoHN S. Moorinc
Eriophyllum lanatum (Pursh) Forbes is a species-complex consisting of at least 10 ill-defined regional varieties that range from southern California to British Columbia, Montana and Wyoming. Constance (1937) noted that of the 157 binomials and trinomials which had been applied to members of Eviophyllum, 75 were referable to E. lanatum. Carlquist (1956) recorded chromosome counts of n = 8 and n = 16 in 2 of the varieties of E. /anatum, and observed that additional studies should prove interesting. My interest in the LE. lanatum complex stems partly from my studies of the Chaenactis douglasii complex (Mooring, 1965; and unpublished). Both of these Helenieae have approximately the same geographic distribution, ecologic amplitude, life span and degree of taxonomic complexity. Among other items, I wished to com- pare the geographic distribution of diploids and polyploids in EL. lanatum with that known for C. douglasii, in which a remarkable correlation exists between ploidy level and geographic distribution.
The purpose of this paper is to present the chromosome counts ob- tained in the first part of this study, and to suggest the direction of further research.
I appreciate the support afforded by National Science Foundation Re- search Grant GB-3338, and the stimulating accounts of Eriophyllum by Constance (1937) and Carlquist (1956).
MATERIALS AND METHODS
Young heads from wild or garden plants were fixed either in 1:3 acetic-alcohol or in 1:3:6 acetic-chloroform-alcohol. Chromosome counts were obtained from pollen mother cells squashed in acetocarmine. Speci- mens have been deposited in the University of Santa Clara Herbarium and replicates of most of them will be distributed elsewhere.
RESULTS
Table 1 shows my chromosome counts. Figure 1 shows my counts, those of Carlquist (1956) and one provided by Peter Raven (letter). The counts of 2n = c. 48 in E. lanatum var. achillaeoides, all the counts in vars. aphanactis, croceum, cuneatum, integrifolium, lanatum, lanceo- latum and leucophyllum, and those for the putative hybrid between var. arachnoideum and E. confertiflorum are first reports. Meiosis was some- times unclear in PMC’s from wild plants, especially at the tetraploid and hexaploid level. Bivalents seemed the rule, although 1028 and 1032 had an occasional association of 4 chromosomes, 1455, 1590, 1642 and Raven 20294 each had a small supernumerary chromosome, and 1024 had a small pair of extra chromosomes.
236
1966] MOORING: ERIOPHYLLUM LANATUM ERT
TABLE 1. CHROMOSOME COUNTS IN ERIOPHYLLUM
Collection numbers are my own, unless otherwise stated.
E. lanatum (Pursh) Forbes var. lanatum. 2n = 16. Idaho. Idaho Co., Kamiah, 1122. Oregon. Wallowa Co., Grand Ronde Canyon, 1107. Washington. Clark Co., Battleground, 1586; Yakima Co., American River, 1600.
var. achillaeoides (DC.) Jepson. 2n = 16. California. Contra Costa Co., Mt. Diablo, 1028; Glenn Co., Stonyford, 1532; Humboldt Co., Garberville, 1574; Santa Clara Co., Loma Prieta, 1375; Siskiyou Co., Grass Lake, 1676; Trinity Co., Douglas City, 1495. Oregon. Klamath Co., Modoc Point, 1665. 2n = 32. California. Colusa Co., Leesville, 1535; Tehama Co., Beegum, 1517. 2n = c. 48. California. Mendocino Co., Hopland, 1032.
var. aphanactis Howell. 2n = 32. California. Glenn Co., Elk Creek, 1520; McGrew Springs, 1525.
var. arachnoideum (Fisch. & Ave.-Lall.) Jepson. 2n = c. 16. California. San Mateo Co., Bear Gulch Road, 1557.
var. croceum (Greene) Jepson. 2n = 32. California. Nevada Co., Banner Hill, 1002, 1005.
var. cuneatum (Kell.) Jepson. 2n = c. 32. California. Nevada Co., Cisco Grove, 1150, 1151.
var. grandiflorum (Gray) Jepson. 2n = 16. California. Amador Co., Jackson, 1467; Calaveras Co., Murphys, 1455; Del Norte Co., Klamath Glen, 1039, 1040, 1043; Idlewild, 1048; El Dorado Co., Pollock Pines, 1471; Mariposa Co., Bagby, 1430; Nevada Co., Banner Hill, 1010; Placer Co., Auburn, 1021; Shasta Co., Whiskeytown Creek, 1490; Platina, 1510; Siskiyou Co., Black Butte Summit, 1681; Trinity Co., Douglas City, 1502; 1506; Tuolumne Co., Stent, 1445; Twainharte, 1450. 2n = 32. California. Butte Co., Oroville, 1476; Shasta Co., Millville, 1487.
var. integrifoltum (Hook.) Smiley. 2n = 16. California. Tuolumne Co., Ken- nedy Meadows, Raven 20294 (DS). Idaho. Owyhee Co., Jump Creek Canyon, 13517, 1352. Oregon. Baker Co., Dooley Mountain, 1094; Deschutes Co., Sisters, 1059; La Pine, 1642; Grant Co., Canyon City, 1079; Klamath Co., Collier State Park, 1650. Wyoming. Fremont Co., Togwotee Pass, 1301.
var. lanceolatum (Howell) Jepson. 2n = 16. California. Trinity Co., Mountain Meadows Ranch, 1712. Oregon. Josephine Co., Oregon Caves, 1053.
var. leucophyllum (DC.) W.R. Carter. 2n = 32. Washington. Lewis Co., Toledo, 1590.
var. arachnoideum (Fisch. & Ave.-Lall.) Jepson & E. confertifiorum (DC.) Gray. California. San Mateo Co., Black Mountain, 1024 (2n = c. 32-40), 1027 (2n= 50).
E. confertiflorum (DC.) Gray. 2n = 16. California. Los Angeles Co., La Canada, 5525 Alta Canada, 1365; San Bernardino Co., Fredalba, 1360.
DISCUSSION
At this time my results merely extend the investigations begun by Constance (1937) and Carlquist (1956). Evriophyllum lanatum var. grandiflorum includes diploids and tetraploids, and var. achillaeoides diploids, tetraploids and hexaploids. The other varieties of the E. lanatum complex for which counts are available appear to consist of diploids (arachnoideum, integrifolium, lanatum, lanceolatum) or of tetraploids (aphanactis, croceum, cuneatum, leucophyllum). My 2 counts for E. confertiflorum are in line with Carlquist’s (1956) results: n — 8 for southern California mainland material. Further counts may well change these impressions.
238 MADRONO [Vol. 18
Fic. 1. Distribution of diploids (circles), tetraploids (squares) and hexaploids (triangles) in the E. lanatum complex. Symbols marked “1” derived from Carlquist (1956), that marked “2” of Raven 20294, all other counts my own. Range of the complex shown in outline.
The paucity of chromosome counts provides small basis for speculation about the evolutionary role of polyploidy in the EZ. lanatum complex. In most instances only one plant per population has been examined cytologically. So far, however, only one polyploid has been found out- side California, and no evidence exists that polyploidy has been mainly responsible for the development of the complex as a whole. Available data do suggest, however, that polyploidy has contributed occasionally, sometimes by promoting interbreeding and elsewhere by inhibiting it. For example, var. cuneatum may be of alloploid origin. It is in many ways intermediate between vars. grandiflorum and integrifolium, and occurs in a relatively narrow zone between them. Chromosome counts of Cisco Grove plants show them to be tetraploids, whereas counts in grandiflorum and integrifolium reveal only diploids, at least near the range of cuneatum. Polyploidy also may have promoted morphologic
1966] MOORING: ERIOPHYLLUM LANATUM 239
divergence by decreasing interbreeding where varieties exist sym- patrically. At Banner Hill, where they grow in a mixed colony, vars. croceum and grandiflorum differ sharply morphologically, phenologically and chromosomally, and, to a lesser extent, ecologically. Counts show croceum to be tetraploid and grandiflorum diploid. Varieties achil- laeoides and aphanactis may represent a similar situation. The latter apparently is essentially a rayless expression of the former, but counts show it to be tetraploid, whereas, achillaeoides may be diploid, tetraploid or hexaploid. Elsewhere, however, varieties may exist sympatrically or nearly so, maintain their integrity, and yet have, apparently, the same chromosome number. This appears to be the situation near Weaverville and near Weed for vars. achillaeoides and grandiflorum. It is quite clear that much additional work is needed before much can be said about either the role of polyploidy in the complex or the taxonomic status of the “varieties.” The foregoing remarks merely represent possible conclusions to be drawn from the scanty data, and suggest the direction of future research. Obviously, much remains to be done, especially in north- western California-southwestern Oregon, the central Sierra Nevada, and in the synonomy-ridden and wide-ranging var. integrifolium.
Eriophyllum lanatum apparently hybridizes with E. confertiflorum (Constance, 1937; Thomas, 1961), and E. jepsonii Greene and E. lati- lobum Rydb. possibly arose, independently, from these sources (Con- stance, 1937; Munz, 1959). I have found 2 localities where E. lanatum apparently hybridizes with LE. confertiflorum. In one it is via var. arachnoideum, in the other via var. achillaeoides. The former popula- tion is the source of the 2n = 32-40 and 2n = 50 counts in Table 1. These populations will receive further study.
Eriophyllum seems to be a rapidly evolving genus. The E. lanatum complex appears to be ancestral or near-ancestral to the rest of the genus, and perhaps to Monolopia and other closely related genera as well, as Constance (1937), without benefit of chromosome number in- formation, speculated a generation ago. Hopefully, further study will clarify the biologic and taxonomic situation in the E. lanatum complex, and perhaps lead to a better understanding of Eriophyllum and of the subtribe Eriophyllanae.
Biology Department, University of Santa Clara, California
LITERATURE CITED
CarLouisT, S. S. 1956. On the generic limits of Eriophyllum (Compositae) and related genera. Madrofio 13:226—239.
ConsTANce, L. 1937. A systematic study of the genus Eriophyllum Lag. Univ. Calif. Publ. Bot. 18:69-136.
Moorinc, J. S. 1965. Chromosome studies in Chaenactis and Chamaechaenactis (Compositae, Helenieae). Brittonia 17:17—25.
Mowz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
THomas, J. H. 1961. Flora of the Santa Cruz Mountains of California. Stanford Univ. Press.
PROPAGATION OF SOME WOODY DESERT PERENNIALS BY STEM CUTTINGS
VALERIE C. CHASE AND B. R. STRAIN
Vegetatively propagated material provides an excellent control param- eter in investigations of the effect of various environmental factors on plant growth. Clonal plants from a single ortet, grown under various natural or controlled environments, may be compared with some assur- ance that the elicited morphological or physiological differences are environmentally induced and not caused by genetic variation (Heslop- Harrison, 1964).
It is well known that desert succulents may be propagated by stem cuttings (many members of Cactaceae), rhizomes (certain members of the genus Yucca), or plantlets (Agave and Kalanchoae ). Little has been published, however, about vegetative propagation of woody desert peren- nials. Although a relatively recent report (Shreve, 1951) stated that Fouquieria splendens was the only woody perennial of the Sonoran Desert which could be propagated by stem cuttings, more recent work with Chilopsis linearis (Everett, 1957) and Simmondsia chinensis (Gentry, 1958) indicates that other desert shrubs may be propagated by stem cuttings.
The general lack of information concerning vegetative propagation of woody desert perennials prompted the testing of some predominant desert shrubs and trees to determine which propagating techniques could be used to obtain clonal material for a proposed comparative physiologi- cal-ecological analysis of desert perennials.
Stem cuttings from 11 woody perennial species (table 1) were col- lected near Palm Springs, California on March 7, 1965 and from nine species (table 2) on June 9, 1965. The cuttings were transported to the laboratory in plastic bags containing a little water.
The cuttings of each species of the first collection were divided into four equal lots, a fresh cut made, and the basipetal 1 inch placed in solu- tions of indole-3-acetic acid (IAA) for 24 hours (Audus, 1959). Two lots were treated with 200 ppm IAA, and one lot each with 100 and 300 ppm IAA.
The cuttings of one of the 200 ppm lots were suspended in an aerated nutrient solution (half-strength Hoagland’s). The solution was changed weekly.
The remaining three lots were planted in vermiculite in a rooting box. All the cuttings were maintained on a 24 hour photoperiod (Lanphear and Meahl, 1961; Baker and Link, 1963), the natural light being supple- mented by fluorescent bulbs. The intensity of the artificial light ranged from 150 to 200 ft-c at plant level. The rooting box was provided with bottom heat of 70° F by a thermistated heating tape (Audus, 1959).
240
1966] CHASE & STRAIN: DESERT PERENNIALS 241
TABLE 1. PERCENTAGE ROOTING OF CUTTINGS TAKEN Marcu 7. THE NUMBERS IN PARENTHESIS INDICATE THE NUMBER OF CUTTINGS IN EACH LOT
TAA concentration
Species 100 ppm 200 ppm 300 ppm Acacia greggii Gray OEcis) O< (13) 0 (13) Cercidium floridum Benth. O (20) O (20) O (20) Dalea schottu Torr. O (20) O (20) O (20) Encelia farinosa Gray #1 0 (10) 10 (10) O (11)
#2 82 (11) 72 ly) 70 (10) Ephedra californica Wats. OmC20) 0 (20) 07(20) Franseria dumosa Gray 33 46) O (6) 33 (6) Hymenoclea salsola T. & G. 78 (23) 95 (20) 95 (20) Isomeris arborea Nutt. O (20) O (20) O (20) Larrea divaricata Cav. O (20) 0) (20) O (20) Prosopis juliflora D.C. ORGS) O (10) O (14) Simmondsia chinensis 30 (20) 24 (20) 24 (20)
C. K. Schneid.
Each species of the second group of cuttings was divided into three lots. Two of the lots were placed in water for 24 hours while the remain- ing lot was soaked for 24 hours in IAA of 200 ppm. One of the water treated lots of each species was dipped in Rootone (Naphthylacetamide, 0.06% ; 2-methyl-1-naphthylacetic acid, 0.033%; 2-methyl-1-naphthy- lacetamide, 0.013%; indole-3-butyric acid, 0.057%; inert ingredients, 99.85% ), a commercial rooting powder, and the other lot was used as an untreated control. The cuttings were placed in the rooting box as out- lined above. After rooting (48 days), the cuttings were transplanted into 4 inch pots in light greenhouse soil and returned to the rooting box to facilitate establishment.
The cuttings placed in the nutrient solution died within 30 days. No roots formed on any of the plants treated in this manner.
Daily maximum and minimum temperatures in the rooting box aver- aged 82 and 87 and 64 and 62 F during the first and second treatment periods respectively. The average maximum relative humidity was 86 and 85 per cent, and the average minimum relative humidity was 59 and 60 per cent during the first and second periods respectively.
In the first test, the percentage of rooting was high in Hymenoclea salsola and Encelia farinosa and fairly high in Franseria dumosa and Simmondsia chinensis (table 1). In this test, all of the cuttings of a species came from a single individual with the exception of Prosopis juliflora and Encelia farinosa. The cuttings of Prosopis were taken from eight plants, those from Encelia from two plants in the same stage of flowering, growing less than 10 ft apart. Only 1 of 31 cuttings of one shrub rooted while 24 of 32 cuttings of the neighboring bush formed roots, indicating the possibility of intraspecific physiological or genetic variation in the rooting response. Consequently, a random sampling technique, in which cuttings were taken from several plants, was em- ployed in the second test.
242 MADRONO [Vol. 18
TABLE 2. PERCENTAGE ROOTING OF CUTTINGS TAKEN JUNE 9. THE NUMBERS IN PARENTHESIS INDICATE THE NUMBER OF CUTTINGS IN THE LOT
Species control Rootone 200 ppm IAA Acacia greggit Gray O= G) O (4) 0 (4) Chilo psis linearis Sweet. 83 (6) 100 (5) 0 (5) Dalea schottu Torr. O (10) O (10) 20 (10) Franseria dumosa Gray 50 (10) 70 (10) 50 (10) Hymenoclea salsola T. & G. 20 (10) 10 (10) 80 (10) Isomeris arborea Nutt. O (10) 50 (10) O (10) Peucephyllum schottii Gray ASG) 100 (8) 33) 706) Pluchea sericea Gove. 20 (10) O (10) 20 (10) Simmondsia chinensis 10 (10) 0 (10) 10 (10) C. K. Schneid.
The second group of cuttings was taken to determine the relative values of IAA and Rootone as rooting hormones. In this test, Chilopsis linearis, Franseria dumosa, Hymenoclea salsola, and Peucephyllum schottiu rooted well. Dalea schotiu, Isomeris arborea, Pluchea serica and Simmondsia chinensis showed some degree of root formation (table 2). I. arborea and D. schottiu had not rooted in the previous test.
The success with Rootone as a rooting substance and the relative ease of its application has precluded further work with IAA. Hymenoclea salsola is the only species that responded significantly better to IAA than to the commercial rooting powder.
CONCLUSIONS
1. Stem cuttings of many woody desert perennials can be induced to form roots, contrary to an earlier report (Shreve, 1951).
2. For most of the shrubs which rooted, maintenance of Rootone treated cuttings in a rooting box under continuous light with moderate bottom heat provides an acceptable percentage of rooting.
3. Treatment with 200 ppm indole-3-acetic acid may enhance rooting in cuttings which respond poorly to Rootone.
4. Some species, however, (Acacia greggit, Cercidium floridum, Ephedra californica, Larrea divaricata and Prosopis juliflora) did not root under any of our treatments.
We are grateful to Mr. Percy C. Everett of the Rancho Santa Ana Botanic Garden for his suggestions and to the National Science Founda- tion for an Undergraduate Student Education Program grant to the senior author.
Department of Life Science, University of California, Riverside
LITERATURE CITED
Aupus, L. J. 1959. Plant growth substances. Intersci. Publ., New York.
Baker, R. L. and C. B. Linx. 1963. The influence of photoperiod on rooting cut- tings of some woody ornamental plants. Proc. Am. Soc. Hort. Sci. 82:596-601.
EveretTT, P. C. 1957. A summary of the culture of California plants at the Rancho Santa Ana Botanic Gardens 1927-1950. Abbey Garden Press, Pasadena.
Gentry, H. S. 1958. Natural history of jojoba (Simmondsia chinensis) and its cultural aspects. Econ. Bot. 12:261-295.
1966 | RAJHATHY: HORDEUM JUBATUM 243
Hestop-Harrison, J. 1964. Forty years of genecology. Jn Advances in ecological research. (J. B. Gragg, ed.) 2:159-247.
LAMPHEAR, F. O. and R. P. MEAHL. 1961. The effects of various photoperiods on the rooting and subsequent growth of selected woody ornamental plants. Proc. Am. Soc. Hort. Sci. 77:620-634.
SHREVE, F. 1951. Vegetation and flora of the Sonoran Desert. Carnegie Inst. Wash. Publ. 591.
NOTES ON THE HORDEUM JUBATUM COMPLEX Trpor RAJHATHY
Mitchell and Wilton (1964) examined the jubatum-caespitosum- brachyantherum complex in Alaska. Their report contains interesting information on the behaviour of the complex at the northern border of the North American population. They concluded that there is “little support for combining the Alaskan populations of H. brachyantherum with H. jubatum.” If the word “Alaskan” is omitted then the conclusion is at variance with the one suggested by Rajhathy and Morrison (1959; 1961) and with the taxonomic treatment of this group by Bowden (1962). Although Mitchell and Wilton dealt with Alaskan populations which are only a small peripheral fraction of the whole North American population, they generalize from their data and propose to maintain species status for both jubatum and brachyantherum. Since in their generalization they disregard a mass of evidence obtained on the popu- lations of the Canadian and American West and Midwest as well as in the experimental garden, a brief discussion of the problem seems to be warranted. This paper is contribution No. 139 from the Research Sta- tion, Canada Department of Agriculture, Central Experimental Farm, Ottawa.
Species relationships should be assessed in the light of the biological species concept. Thus, interbreeding should be recognized as the main criterion of conspecific status. Mitchell and Wilton stated that H. brachy- antherum and H. jubatum fail to “hybridize on many sites where they are in contact.” This appears to be the case in Alaska. Although they located a few hybrid populations, their future remains doubtful. The authors may be right because the extreme environment of Alaska may not provide the ecological niche for successful competition or introgres- sion. However, even if this were the case, generalizing from a special- ized peripheral situation cannot be considered valid. When interbreed- ing between two species populations is examined surely greater signifi- cance should be assigned to those sites where hybridization does occur than to those where it does not. Large hybrid swarms exist in the Canadian and American West and Midwest. These became well estab- lished also in areas such as Saskatchewan and Manitoba where only H. jubatum occurs and brachyantherum is absent (Bowden, 1962). Thus, they not only maintain themselves but also migrate from the original sites.
244 MADRONO [Vol. 18
Hybridization is a prerequisite of interbreeding, the fertility of the hybrids being a more sensitive indicator of the degree of relationship than crossability alone. Using these in evaluating relationships the breed- ing system of the parental species and isolation mechanisms should be considered. Both jubatum and brachyantherum are inbreeders, hence it is not surprising that they do not hybridize at some sites of contact. Inhibited embryo development in interspecific hybrids is a very char- acteristic isolation barrier in the Hordeum genus (Rajhathy et al., 1964). In view of this, the absence of this barrier in the jubatum- brachyantherum group must be considered very important. Some ad- vanced generation hybrid strains are fertile to the extent that they are just as troublesome weeds as H. jubatum. Failure of seed set is not likely to be an isolation barrier even in Alaska where one plant in a relatively small sample had 47 per cent seed set. The chances for superior and fertile combinations are much better in nature than in the nursery where usually only a few genotypes are involved. A wide array of types seg- regate in F, and F, generations for awn length which is the best char- acter differentiating the parental taxa. The natural hybrid swarms are more similar morphologically to jubatum than to brachyantherum be- cause jubatum is usually the recurrent parent.
The abundance of natural hybrids and the partial fertility of the F, hybrids in the nursery are indicative of a relatively regular gametogene- sis. Some irregularities in chromosome pairings were noted in the hybrids (Rajhathy and Morrison, 1959). However, a sufficient amount of good pollen grains was produced to secure good seed set. Thus, it is surpris- ing that Mitchell and Wilton were unable to record chromosome pair- ing at first metaphase because of the incidence of irregularities.
The taxonomic treatment of the jubatum group by Bowden was based on experimental results obtained by cytogenetical methods and reflected biological relationships. The interesting data of Mitchell and Wilton make the information more complete but do not change the overall pattern of species relationships in this important group.
Ottawa Research Station, Research Branch, Canada Department of Agriculture, Ottawa, Ontario
LITERATURE CITED
Bownpen, W. M. 1962. Cytotaxonomy of the native and adventive species of Hor- deum, Eremopyrum, Secale, Sitanion and Triticum in Canada. Canad. Jour. Bot. 40:1675-1711.
MitcuHe 1, W. W. and A. C. Witton. 1964. The Hordeum jubatum-caespitosum- brachyantherum complex in Alaska. Madronio 17:269-280.
Rayuatuy, T. and J. W. Morrison. 1959. Cytogenetic studies in the genus Hordeum. IV. Hybrids of H. jubatum, H. brachyantherum, H. vulgare and a hexaploid Hordeum sp. Canad. Jour. Genet. Cytol. 1:124-132.
. 1961. Cytogenetic studies in the genus Hordeum. V. H. jubatum and the New World species. Canad. Journ. Genet. Cytol. 3:378-390.
, and S. Symxko. 1964. Interspecific and Intergeneric hybrids in Hordeum. Barley Genetics I. Proc. First Intern. Barley Genet. Symp. Wageningen.
DOCUMENTED CHROMOSOME NUMBERS OF PLANTS (See Madrono 9:257-258. 1949; 17:255. 1964)
Astranthium condimentum De Jong. n= 3. Mexico, Jalisco, Mazamitla. D. C. D. De Jong 1675, MU; Michoacan, near Quiroga, D. C. D. De Jong 1656, MU; Michoacan, near Ario de Rosales. D. C. D. De Jong 1657, MU.
A. orthopodum (Robins. & Fern.) Larsen. n = 3. Mexico, Chihuahua, near Madera. D.C.D. De Jong 1769, MU; Durango, near Hacienda Coyotes. D. C. D. De Jong 1382, MU.
A. purpurascens (Robins.) Larsen. n = 8. Mexico, Chiapas, between Comitan and San Nicolas. D. C. D. De Jong 1419, MU; Chiapas, between Teopisca and San Cristobal las Casas. D. C. D. De Jong 1438, MU; Hidalgo, Puerto de Horcones. D.C. D. De Jong 1584, MU; Hidalgo, near Jacala. D. C. D. De Jong 1585, MU.
A. xanthocomoides (Less.) Larsen. n = 8. Mexico, Hidalgo, Elvado. D. C. D. De Jong 1535, MU; Hidalgo, Real del Monte. D. C. D. De Jong 1550, MU; Puebla, Laguna de Atezca. D. C. D. De Jong 1594, MU; Veracruz, near Magueytos. D.C. D. De Jong 1607, MU.
A. xylopodum Larsen. n = 5. Mexico, Nayarit, near Compostela. D. C. D. De Jong 1690, MU.
Baileya multiradiata Torr. n= 16. Mexico, Chihuahua, near Ciudad Chihuahua. D.C.D. De Jong 1377, MU.
Buddleia skutchii Morton. 2n = ca. 38n. Mexico, Chiapas, Chamula. D. E. Breed- love 6654, DS, F. Counted by D. W. Kyhos.
Chlorogalum pomeridianum (DC.) Kunth. n = 15. California, Fresno Co. H. M. Beard & W. Roderick, 1961, JEPS. Counted by M. S. Cave.
Cleome spinosa Jacq. 2n = 101. Mexico. Chiapas, Tuxtla Gutierrez. D. E. Breed- love 9193, DS, F. One pair was twice the size of the others as in the tetraploid Cleome isomeris (Nutt.) Greene (Isomeris arborea Nutt.). Counted by D. W. Kyhos.
Coriaria thymifolia Hum. & Bonpl. ex Willd. 2n = 20n. Mexico, Chiapas, Zinacan- tan. P. H. Raven & D. E. Breedlove 20142, DS. Fig. 1. Counted by D. W. Kyhos.
Fic. 1. Coriaria thymifolia, * 1470.
Erythronium tuolumnense Appleg. n = 12. California, Tuolumne Co. W. Roderick, 1962, JEPS. Counted by M. S. Cave.
Galphimia glauca Cav. n = 6. Mexico, Chiapas, Pinola. D. E. Breedlove 7072, DS, F. Very large chromosomes. Adds a third basic number to the two (x = 9,10) already known for the Malphigiaceae and suggests that the counts of n = 12 in this family are tetraploid. Counted by D. W. Kyhos.
Microspermum debile Benth. n = 12. Mexico, Oaxaca, Ixtlan de Juarez, D. C. D. De Jong 1481, MU.
245
246 MADRONO [Vol. 18
Narthecium californicum Baker. n= 18. Oregon, Josephine Co. W. Roderick, 1962, JEPS; W. Roderick, 1965, JEPS. Counted by M. S. Cave.
Olivaea leptocarpa De Jong & Beaman. n= 6. Mexico, Durango, near J. G. Agui- lera. D.C. D. De Jong 1754, MU; D.C. D. De Jong 1760, MU.
O. tricuspis Sch.-Bip. ex Benth. n = 6. Mexico, Jalisco, near Acatlan. D. C. D. De Jong 1687, MU; Jalisco, near Guadalajara. D. C. D. De Jong 1688, MU; State of Mexico, near Toluca. D. C. D. De Jong 1619, MU; D.C. D. De Jong 1650, MU; State of Mexico, Del Rio. D. C. D. De Jong 1647, MU; Nayarit, Tepic. D. C. D. De Jong 1693, MU.
Schoenolirion album Dur. n= 26. California, Plumas Co. G. E. See, 1957, JEPS. Counted by M. S. Cave.
Spiranthes gracilis (Bigel.) Beck. n= 15. Florida, Hillsborough Co. R. W. Long 1168, USF. Counted by M. S. Cave.
Stenanthium occidentale Gray. n = 8. California, Siskiyou Co. W. Roderick, 1962, JEPS; W. Roderick, 1965, JEPS. Counted by M. S. Cave.
Tofieldia glutinosa (Michx.) Pers. ssp. occidentalis (Wats.) Hitch. n = 15. Oregon, Josephine Co. W. Roderick, 1962, JEPS. Counted by M.S. Cave.
Veratrum californicum Dur. n = 16. California, Alpine Co. W. Roderick, 1965, JEPS; Eldorado Co. H. G. Baker, 1965, JEPS. Counted by M. S. Cave.
V. fimbriatum Gray. n= 16. Califoria, Mendocino Co. W. Roderick, 1965, JEPS. Counted by M. S. Cave.
V. insolitum Jeps. n= 16. California, Siskiyou Co. W. Roderick, 1965, JEPS. Counted by M.S. Cave.
Xerophyllum tenax (Pursh) Nutt. n = 15. California, Mendocino Co. W. Roder- ick, 1965, JEPS. Counted by M.S. Cave.
REVIEWS
THe Hunt BoranicaLt Liprary AND Its Pusrications.—The institution known as the Rachel McMasters Miller Hunt Botanical Library is the lengthened shadow of not only Rachel McMasters Miller Hunt but also of its director, George H. M. Lawrence. The history of the Library has been told well in The Rachel McMasters Miller Hunt Botanical Library (1961). The Library is a relatively new institution and has been housed in its present location in the penthouse of the Hunt Library on the campus of the Carnegie Institute of Technology in Pittsburgh since 1961. The setting and decor of the penthouse are truly beautiful and are a fitting place for the care and study of Mrs. Hunt’s collection of incunabula, rare botanical and horticultural books, botanical prints, letters, manuscripts, and fine bindings.
Mrs. Hunt’s interest in botany stemmed from her early girlhood—at the age of six she was given a copy of Mrs. Dana’s How to Know Wildflowers. With the pass- ing years as Mrs. Hunt acquired more and more rare works she realized that her collection was in many ways unique in North America. Her first steps toward sharing her collection with others resulted in the magnificent two volume Catalogue of Botanical Books in the Collection of Rachel McMasters Miller Hunt (1958, 1961). These two volumes deal with 764 works dating from about 1150 through 1800. The Catalogue was issued in a limited edition of 750 copies. Volume I was compiled by Jane Quinby, Mrs. Hunt’s friend and librarian, and contains chapters by Harold W. Rickett, John F. Fulton, Paul B. Sears, Wilfrid Blunt, and Margaret B. Stillwell: Volume II, which is in two parts, was compiled by Allan Stevenson and contains introductory material prepared by Gordon Dunthorne, John S. Gilmour, and Wil- liam T. Stearn. The volumes of the Catalogue are beautifully done with respect to printing, composition, paper, and binding.
1966] REVIEWS 247
In 1961 Mrs. Hunt’s books were moved from the library at the Hunt’s residence in Pittsburgh, Elmhurst, to the newly completed Hunt Library. The previous year, Dr. Lawrence had been appointed Director of the Hunt Botanical Library and since that time the Library has expanded its scope and activities to become a most important botanical research center.
The main activity of the Library at present is the Bibliography Huntiana, which may be characterized as a new Pritzel. When completed it will be a major biblio- graphical source in systematic botany, covering the literature from 1730 through 1840 and will comprise 16 quarto volumes. Title pages of books will be reproduced in facsimile. The preparation of Bzbliographia Huntiana is being done, in keeping with the times, with the aid of computers.
The Library has already published two volumes of Huntia, A Yearbook of Botanical and Horticultural Bibliography (1964, 1965). Huntia contains an inter- esting series of papers by members of the Library staff and others. Of particular interest to western botanists is a paper entitled Some Recollections of Percy Train, by Chester A. Arnold (Huntia 2:111-116. 1965).
The collection of portraits of botanists, now numbering several thousand, is the special concern of Theodore W. Bossert. The collection is growing and in time should be the most complete in North America.
The Library has as another of its important functions the periodic exhibition of botanical illustrations. Notable among these have been the exhibition of Redoutéana during August, 1963. The catalogue of the exhibition, A Catalogue of Redoutéana (1963), follows the same high standards as the other Library publications.
Another function of the Library is periodic symposia. Those who attended the Adanson Symposium in 1963 will remember it with pleasure. The papers delivered by Jean-Paul Nicolas, Frans Stafleu, William Margadant, Mme. G. Duprat, Peter Sneath, and Theodore Monod were published in two parts entitled, Adanson, The Bicentennial of Michel Adanson’s “Familles des plantes” (1963, 1965). Adanson is the first of the Hunt Monographic Series.
Additional collections in the Library consist of bookplates, examples of fine bindings, letters, and manuscripts. Eventually the Library will be a major deposit- ory for letters and thus an important source of material for those interested in historical aspects of botany.
Two works have so far been reproduced in the Hunt Facsimile Series. The first (1963) was L’Héritier de Brutelle’s Sertum Anglicum. It is indeed a rare book in that only 35 complete copies of the original are known. The book is beautifully executed and includes introductory chapters by Frans Stafleu, John S. Gilmour, C. J. King, L. H. J. Williams, and Wilfrid Blunt. The second (1966) in the Facsimile Series is Walahfrid Strabo’s Hortulus or “The Little Garden.” The Hortulus is of less interest to professional botanists than Sertum Anglicum as it predates Linnean nomenclature by nearly a thousand years. The Hortulus was reproduced because “Of all the gardening literature of this and ages past, nothing has survived longer, and little has been extolled more ... .It has survived this long, and should live for more than as long again, not because of gardening innovations, accounts of new plants, or pharmacological discoveries reported in it, but because it is beautiful poetry, because it is full of man’s love for the earth and for the plants he grows in it.” (forward, p. v).
Perhaps one of the most interesting aspects of the Hortulus facsimile is the series of prints by Henry Evans of San Francisco. Twenty-eight of his prints are used to decorate the English translation done by Raef Payne. Henry Evans is cer- tainly among the foremost of contemporary botanical print makers and this is con- firmed by the use of specially commissioned prints in the Hortulus.
Long before Mr. Evans started making botanical prints he was well known in San Francisco as a bookman, as a printer, and as the proprietor of the Peregrin Press. His botanical prints are not scientific illustrations in the sense of those that det our technical floras and manuals, but are designed to give an impression of a
248 MADRONO [Vol. 18
Fic. 1. Reproduction of a print of Scoliopus bigeloviit Torrey, California Fetid Adder’s Tongue, made by Henry Evans and reproduced by permission.
particular plant. He is interested in making botanical prints for decorative purposes. Whereas a scientific illustration puts in as many details as possible, Mr. Evans’ prints avoid many details to achieve a clear, clean representation. His prints have an open, graceful, and airy character. There is no doubt, however, about their identity, as the reproduction (fig. 1) of Scoliopus bigelovii Torrey or California Fetid Adder’s Tongue will demonstrate.
All of Mr. Evans’ plants are drawn in their natural settings. He believes that the best composition exists in nature and that it should not be tampered with. For him printing is work and as he is more interested in finding a plant, drawing it, and cutting the linoleum block, he limits the number of prints from any one block to 100 and many to less than that number.
A complete set of Henry Evans’ prints are among the recent acquisitions of the Hunt Botanical Library and an exhibit of his prints will be held at the Library in November of this year.
1966] REVIEWS 249
The Hunt Botanical Library is a specialized institution performing scholarly functions with both efficiency and taste. As our libraries become more and more crowded we will have to rely increasingly on specialized libraries, for much as we may hate to contemplate the notion, many libraries will not be able to house material as they have been accustomed to in the past. For this purpose, the Hunt Botanical Library serves as an admirable model. It deserves the full support of the botanical community and we should be grateful to its sponsors, director, and staff for their efforts —Joun H. THomas, Dudley Herbarium, Stanford University.
The Origin of Cultivated Plants. By FrRANz SCHWANITZ. pp. 175. Harvard Uni- versity Press, Cambridge. 1966. $4.75.
There is urgent need for a modern treatment of the origin of cultivated plants, patterned after De Candolle’s classical work, “The origin of cultivated plants’ pub- lished in 1886. In spite of the title, Schwanitz’ little book of 175 pages evidently was never intended to fulfill this need. Indeed, the accumulated knowledge of the origin of cultivated plants is now so voluminous that it could not be compressed within the covers of a single volume of reasonable length. Furthermore, it is probably beyond the ability or capacity of one individual to treat the subject adequately.
Schwanitz’ book was first published in 1955. The present version was translated from German by Gerd von Wahlert and published by Harvard University Press in 1966. Unfortunately, there is no indication that the opportunity for up-dating the text was used to advantage. As a result statements about corn, wheat and perhaps others do not reflect new and significant information about these crops.
The audience for which the book was prepared is not indicated, but it would be useful as supplemental reading for advanced undergraduate and graduate courses in economic botany, origin of cultivated plants, and other courses of like nature. Scientists or laymen whose work is with cultivated crops should find much of inter- est in the text.
A large portion of the book is devoted to specific examples of the ways in which wild plants differ from their corresponding cultivated forms. This is the most inter- esting and useful part of the book. Also, the well-known story of the transition of lupine from an essentially wild plant to a cultivated one under the skillful guidance of the German plant breeders, von Sengbusch and his colleagues, is reviewed in detail. The concluding chapter is a good short sketch of some of the highlights in the history of plant breeding.
The unwary reader should be cautioned against some rather serious errors that have crept into the text, most likely as the result of faulty translation. For example on page 5, Correns, de Vries and Tschermak are credited with the “discovery” of the laws of heredity. Obviously it should have been “rediscovery.” This error is partially corrected on page 161 (156 pages later) with the statement: ‘The ‘redis- covery’ of Mendel’s laws of heredity around the turn of the century, etc.” On page 7, it is stated that wild rice, Zizania aquatica “was one of the most important food- stuffs of North American Indians.” True, it was an important food for a few tribes in northern Minnesota where conditions are suitable for its culture. But to imply its use was wide-spread among North American Indians is completely misleading.
I doubt that those familiar with the taxonomy and systematics of maize would agree with the statement on page 146, “Maize, Zea maize is closely related to millet.” Since this is an undocumented assertion it is difficult to guess what millet the author or the translator had in mind. These are samples of the errors that could be cor- rected should the popularity of the book call for a reprinting or a new edition.
It is apparent that the translator is not at home with agricultural terms and practices. Unfamiliarity has led to a number of awkward statements and phrases, and several redundancies. Under Literature about 20 general works are cited, none more recent than 1955. An index of about 4%4 pages, with a few minor errors, com- pletes the book—Tuomas W. WHITAKER, United States Department of Agriculture, Agricultural Research Service, La Jolla, California.
250 MADRONO [Vol. 18
The California Deserts. By EDMUND C. JAEGER. xii + 208 pp. Stanford Uni- versity Press, 4th edition, 1965. $4.95.
E. C. Jaeger’s new edition of The California Deserts illuminates for the reader only a limited portion of the vast phenomena which occur in this extensive area of southeastern California. The book is primarily intended as a guide whose purpose is to acquaint the amateur naturalist or traveler with the general array of life in this region. It fulfills this function admirably as it includes chapters on history, climate, physiography, travel, preservation, and nearly all biological groups found in the desert including insects, fish, amphibians, reptiles, birds, mammals, aborigines, fungi, lichens, ferns, and higher vascular plants. However, the elucida- tion of each species is very short, which, as the author states has “. . . left it for the reader to detect where the nuggets lie ensconced.” The new edition is essentially similar to the third except for several new photographs and the addition of chap- ters on aborigines and desert conservation.
Dr. Jaeger captures the very essence of the region with 16 magnificent plates and the simple statement that “. . . incoming rain clouds are generally for me the signal to start desertward. Let me have the delicious odors of the creosote bush and saltbush when they are wetted with gentle rains, look upon the endless variety and beauty of the clouds’ far-flung forms, have the silence of the uninhabited mesas, and I am in a land enchanted.”
For a person not acquainted with the desert this book offers an opportunity to learn some of the interesting salient features about the more common plant and animal inhabitants. The descriptions and drawings of the organisms, although suffi- cient for the uninitiated, are for the most part inadequate to provide exact identifica- tion. At times these become more of a check-list of species and their habitats leaving the reader with a crying desire for more information. Dr. Jaeger also describes many marvelous adaptations for survival in a desert environment and endows the reader with an appreciation for the country and its inhabitants.
Unfortunately, the desert region is rapidly being depleted as the heavily popu- lated areas of California expand. Dr. Jaeger cites several examples of this encroach- ment and states: “Most unfortunate it is that, perhaps more than any portion of our land heritage, our deserts are thought by many as the most expendable lands we possess, mere wastelands that should if possible be utilized for gain. Even for many ‘conservationists’, conservation means only efficient exploitation of the land, getting everything possible out of it while perpetuating enough of what is left to serve as ‘seed’ for successive exploitations. . . . They would completely make over nature, leaving the ‘man-did-it-look’ upon almost every part of it.”
Dr. Jaeger has combined his sensitive appreciation for the desert with his vast knowledge and has produced a well-written, pleasant book, which will be enjoyable reading for anyone interested in this region—RoBert M. Lioyp, Department of Botany, University of California, Berkeley.
NOTES AND NEWS
Notes ON SOME GRASSES OF THE Pacrric Coast. — Brachypodium sylvaticum (Huds.) Beauv. was discovered growing in large colonies and apparently thoroughly naturalized at two localities in Benton Co., Oregon. This common woodland grass of Europe, Asia, and North Africa has been mentioned as introduced into North Ameri- ca for cultivation in ornamental grass gardens (Hitchcock, Man. Grasses U. S., ed. 2,57, 1950; Weintraub, U. S. Dept. Agri. Handb. 58:16. 1953). However, it has not previously been reported as well established in nature, at least on the Pacific Coast. At the Benton Co. sites, it grows in partial shade at the edges of woods, associated mainly with Acer macrophyllum and Quercus garryana. The following collections will be deposited at the Oregon State University herbarium (OSC): John Mellott
1966] NOTES AND NEWS 251
14, slopes E of Oak Creek, southern border of McDonald State Forest, R. 5 W., T. 11 S., Section 20; K. L. Chambers 2478, 3 miles W of the Willamette River bridge at Albany, R. 4 W., T. 11 S., Section 3. An earlier Oregon collection (OSC) of this species was submitted in December, 1939, by O. S. Fletcher, the locality given as “near Eugene,” but without further habitat data. This grass probably is more com- mon in the Willamette Valley than these few collections would indicate and has sim- ply been overlooked. Although tall and gregarious, it could easily be mistaken for Agropyron, a genus whose inflorescence is quite similar to that of Brachypodium.
Agrostis californica Trin. is the name given in current manuals for a low-grow- ing, dense-panicled coastal species which ranges from Monterey County, California, to about Lincoln County, Oregon, and which represents an element of the wide- spread and variable A. exarata-complex. In his early monograph of Agrostis (U. S. Dept. Agr., Bur. Pl. Indus. Bull. 68:30. 1905), Hitchcock referred this plant to A. glomerata (Presl) Kunth, based on Vilfa glomerata Pres] (Rel. Haenk. 1:239. 1830), whose type locality is “in montanis Peruviae huanoccensibus.” In subsequent pub- lications Hitchcock segregated the North American material as A. californica Trin., described in 1841. However, the bibliographical citation given in the Hitchcock (op. cit., 804) leads one to suspect that this name may be illegitimate. It states that in the description of A. californica, Vilfa glomerata Pres] was “erroneously cited as synonym.” I have examined a reprint of the original publication by Trinius, from the Hitchcock and Chase agrostology library at the U. S. National Herbarium. It is titled ‘“Agrostidea,” and is a separate from the Memoires of the Royal Academy of St. Petersburg bearing somewhat different pagination (according to Hitchcock’s notes). Following his Latin description of A. californica, Trinius gives the citation, “Vilfa glomerata Presl! Rel. Haenk. fasc. IV, p. 239. California. In montanis Peru- viae huanoccensibus.” Through this reference to an earlier available epithet, the name as published is superfluous, and under the present rules of nomenclature it must be rejected. The correct name for this plant, assuming it is not conspecific with A. glo- merata of Peru, is A. densiflora Vasey (Contr. U. S. Natl. Herb. 3:72. 1892). The type specimen is from Santa Cruz, California.
Agrostis alaskana Hultén, found in coastal Alaska and the Aleutian Islands, is a member of the species group that includes A. scabra Willd. and A. borealis Hartm. Hultén proposed the name (FI. Aleut. Isl., ed. 1, 71, 1937) as a substitute for A. aenea (Trin.) Trin., an illegitimate homonym. However, Hitchcock, in his 1905 mono- graph of Agrostis (op. cit.,51) had published the name A. melaleuca (Trin.) Hitchce. for this plant, and a reference to this is cited by Hultén in his synonymy of A. alas- kana. Hultén argued that A. melaleuca could not be used because it was based on A. canina var. melaleuca Trin., which was described from two specimens representing different species (as was admitted later by Trinius, and by Hitchcock). It is clear from Hitchcock’s publication, however, that A. melaleuca is a legitimate name; it is an avowed new combination of the epithet melaleuca of Trinius, and Hitchcock specifies as a lectotype the specimen he believes is conspecific with A. aenea, exclud- ing the second specimen (which was probably a plant of A. aequivalvis, a very dif- ferent species). It must be admitted that Hitchcock made the lectotypification by inference from the description, since on his visit to the Trinius herbarium at St. Petersburg he could not locate the type specimen of A. canina var. melaleuca. None- theless, he did not, as Hultén believed, base the name A. melaleuca on the type specimen of A. canina var. aeneéa; he merely stated that he saw the latter type at St. Petersburg. I conclude, therefore, that A. alaskana Hult. is superfluous and illegiti- mate, and the name A. melaleuca (Trin.) Hitchc. should be used if this plant is con- sidered a distinct species. It is worthwhile to note that Hitchcock’s own later dispo- sition of this taxon (FI. N. Am. 17:530. 1937) was as a strict synonym of A. borealis. The status of these taxa in Alaska very much needs clarification.
Thanks are extended to LaRea D. Johnston and Curt G. Carlbom for their help in the preparation of these notes. — Kenton L. CHAMBers, Department of Botany, Oregon State University, Corvallis.
INDEX TO VOLUME XVIII
Classified entries: Chromcesome numbers of plants, documented; Reviews. New botanical names are in boldface type. Unannotated taxa in floral lists are ommitted
from the Index.
Abrams’ collecting trip of 1908, Dr. Le- Roy, 97
Acacia greggii, 241
Agrostis: aenae, 251; aequivalis, 251, alaskana, 251; borealis, 251; califor- nica, 251; canina var. aenea, 251; c. var. melaleuca, 251; densiflora, 251; glomerata, 251; melaleuca, 251; scab- tan 5 1
Alatipes, 217
Algae of Washington and southern Brit- ish Columbia, Notes on marine, 176
Allium: carinatum, 148; paradoxum, 149
Allolepsis: A new segregate of Distich- lis (Gramineae), 33; gen. nov., 34; texana, comb. nov., 34
Anderson, E.: Review, Flora of Mis- sour, 28
Anderson, L. C. and J. L. Reveal: Chrysothamnus bolanderi, an inter- generic hybrid, 225
Arceuthobium: douglasii in Nevada and Wyoming, 63; 218
Arctostaphylos: glandulosa 47; glauca, 48; morroensis, 48; obispoensis, 48; pringlei var. drupacea, 48; rudis, 48
Arcyodes: incarnata, 141; insignis, 141; occidentalis, 141
Avicularia, 85
Baker, H. G.: Volatile growth inhibitors produced by Eucalyptus globulus, 207
Bemis, W. P. and T. W. Whitaker: Na- tural hybridization between Cucurbita digitata and C. palmeri, 39
Berg, R. Y. and R. J. Maze: Contribu- tion to the embryology of Muilla, with a remark on the taxonomic po- sition of the genus, 143
Bergseng, M. S.: The occurrence of a New Zealand plant, Geranium micro- phyllum in California, 213
Besa papilliformis, 177
Blondia trifoliata, 158
Bouteloua chasei, 185
Brachypodium sylvaticum, 250
Brodiaea from San Clemente Island, California, A new species from, 233; filifolia, 233; kinkiensis, sp. nov., 233; peduncularis, 148
Buddleia: corrugata, A reevaluation of, 92; corrugata ssp. gentryi, ssp. nov., 93; c. ssp. Moranii, ssp. nov., 94; mar- rubiifolia, 96; utahensis, 96
California Native Plant Society, 104 Callithamnion: endovagum, 177; lejo- lisea, 177
252
Carter, A.: Review, A selected guide to the literature on the flowering plants of Mexico, 127
Castetter, E. F. and P. Pierce: Escobaria leei Bodeker rediscovered in New Mexico, 137
Cathartolinum digynum, 182
Cave, M. S.: The chromosomes of Sco- liopus (Liliaceae), 211
Ceramium eatonianum, 178
Cercidium floridum, 241
Chaenactis douglasii, 236
Chamaecyparis: Varietal transfers in Cupressus and, 161; henryae, 165; lawsoniana, 164; nootkatensis, 164;
thyoides, 165; t. var. henryae, comb. nov., 165
Chambers, K. L.: Notes on some grasses of the Pacific Coast, 250
Chaparral: in the Santa Ana Moun- tains, California, Manzanita, 47; veg- etation, Influences of fire and sodium- calcium borate on, 193
Chase, V. C. and B. R. Strain: Propaga- tion of some woody desert perennials by stem cuttings, 240
Chromosome numbers of plants, docu- mented: Aesculus californica, 122; sylvatica, 123; Allotropa virgata, 123; Alopecurus aequalis, 123; Anthoxan- thum odoratum, 123; Aristolochia cal- ifornica, 123; Astranthium condimen- tum, 245; orthopodum, 245; purpu- rascens, 245; xanthocomoides, 245; xylopodum, 245; Baileya multiradiata, 245; Bidens pilosa var. radiata, 123; Bocconia frutescens, 123; Brodiaea appendiculata 123; californica var. leptandra, 123; clementina, 123; cor- onaria, 123; c. var. macropoda, 123; c. var. rosea, 123; dudleyi, 123; ele- gans, 123; filifolia, 123; gracilis, 123; hyacinthina, 123; h. var. greenei, 123; jolonensis, 123; lutea var. anilina, 123; pallida, 123; Buddleia skutchii, 245; Carpentaria californica, 123; Cath- cartia villosa, 123; Chelidonium ma- jus, 123; Chlorogalum pomeridianum, 245; Chrysopsis asper, 124; Chryso- splenium glechomifolium, 124; Cle- matis crispa, 124; Cleome spinosa, 245; Clethra Janata, 124; Coriaria thymifolia, 245; Corydalis lutea 124; Dicentra chrysantha, 124; formosa, 124; nevadensis, 124; ochroleuca, 124; spectabilis, 124; Drymaria debilis, 124; Erigeron quercifolius, 124; Erio- gonum inflatum var. inflatum, 124;
1966]
maculatum, 124; nudum var. pubi- florum, 124; ovalifolium var. nivale, 124; o. var. ochroleucum, 124; o. var. orthocaulon, 124; pusillum, 124; reni- forme, 124; sphaerocephalum var. halimioides, 124; trichopes ssp. minor, 124; umbellatum ssp. polyanthum, 124; Erythronium tuolumense, 245; Eupatorium album, 125; Fritillaria atropurpurea, 125; Galphimia glauca, 245; Helenium decurrens, 125; Heli- anthus angustifolius, 125; Hetero- theca subaxillaris, 125; Hierochloe oc- cidentalis, 125; Hilaria jamesii, 125; Houstonia caerulea, 125; serpyllifolia, 125; Illicium floridanum, 125; parvi- florum, 125; Marshallia graminifolia, 125; Meconopsis betonicifolia, 125; cambrica, 125; paniculata, 125; Meli- anthus major, 125; Microspermum de- bile, 245; Narthecium californicum, 246; Olivaea leptocarpa, 246, tricus- pis, 246; Orthocarpus castillejoides var. humboldtiensis, 125; Palafoxia integrifolia, 125; sphacelata, 125; Papaver alpinum, 125; atlanticum, 125; bracteatum, 125; nudicaule, 125; rhoeas, 125; somniferum, 125; Plan- tago erecta, 125; insularis, 126; Pleu- ropogon refractus, 126; Porophyllum tridentatum var. crassifolium, 126; Primula maguieri, 126; Schizandra glabra, 126; Schoenolirion album, 246; Sidalcea malachroides, 126; Solanum brachystachys, 126; Spiranthes graci- lis, 246; Stenanthium occidentale, 246; Tofieldia glutinosa ssp. occidentalis 246; Tolmeia menziesii, 126; Veratrum californicum, 246; fimbriatum, 246; insolitum, 246, Xerophyllum tenax, 246
Chrysothamnus: bolanderi, an _ inter- generic hybrid, 225; nauseosus ssp. al- bicaulis, 225; parryi ssp. monoceph- alus, 225
Clausen, Jens C., 175
Coleman, J. R. A taxonomic revision of section Sonoricola of the genus Ver- besina L. (Compositae), 129; Two new species of Verbesina (Composi- tae) with taxonomic notes on other species, 214
Comatricha cornea, 142
Coreopsis ovata, 217
Correll, D. S.: Review, Generic names of orchids, 30
Cottea pappophoroides, 26
Cottinae, subtr. nov., 25
Critchfield, W. B. and G. L. Allenbaugh: Washoe pine on the Bald Mountain Range, California, 63
Crocidium in San Luis Obispo Co., California, 32
Croton: californicus, Translocation het- erozygosity in, 31; californicus var. tenuis, 31; glandulosa, 31; monantho-
INDEX 258
gynis, 31; palmeri, 31
Crypsis: niliaca, 26; schoenoides, 26
Cryptopleura lobulifera, 178
Cucurbita digitata and C. palmata, Na- tural hybridization between, 39
Cupressus: and Chamaecyparis, Varietal transfers in, 161; arizonica var. a., 162; a. var. bonita, 162; a. var. bo- nito,; 162; as. var: compacta, 163° Fa, var. glabra, comb. nov., 162; a. glau- ca, 163; a. f. glomerata, 162; a. f. minor, 162; a. var. montana, comb. nov., 163; a. var. nevadensis, comb. nov., 164; a. var. stepensonii, comb.
nov., 164; benthamii, 161; b. var. arizonica, 162; forbesii, 161; glabra, 162; lindleyi, 161; lusitanica, 161;
macnabiana nevadensis, 164, montana, 163; nevadensis, 164; sargentii, 161; stephensonii, 164; thyoides, 165
Dalea schottii, 241
Dempster, L. T. and G. L. Stebbins: The fleshy-fruited Galium species of Cali- fornia (Rubiaceae). I., 105
Dermatolithon pustulatum f. ascripti- cum, 177
Desert perennials by stem cutting, Prop- agation of some woody, 240
Didymium: melanospermum, 142 minus, 142; serpula, 142
Distichlis: (Gramineae), Allolepis a new segregate of, 33; hirta, 38; marginata, 38; palmeri, 38; scoparia, 38; spicata, 38; texana, 34; thalassica, 38: viridis, 38
Dunn, D. B.: The inter-relationships of the Alaskan lupines, 1
Duravia, 85
Encelia: cedrocensis, 131; farinosa, 241
Endophyton ramosum, 176
Enerthenema papillatum, 141
Enneapogon: desvauxii, 27; flavescens, 27; glumosus, 27
Ephedra californica, 241
Eragrostis: 33; reptans, 26
Eriogonum: nutans and collinum, On the specific distinction of, 167; baileyi var. tomentosum, 168; brachypodum, 167; cernuum ssp. acutangulum, 168; c. var. purpurascens, 171; collium, 169; de- flexum, 167; d. ssp. ultrum, 172; de- missum, 168; hookeri, 167; macu- latum, 172; nutans, 169; n. var. brevipedicelatum, 172; n. var. glabra- tum, var; nov., 1/25 ny varon.,, 1/1; parryi, 167; praebens, 168; p. var. divaricatum, 168; reniforme, 169; r. var. asarifolium, 168; restioides, 169, rubiflorum, 171; thurberi var. acutan-
gulum, 168 Erioneuron, 33 Eriophyllum: lanatum (Compositae, Helenieae) complex, Chromosome
254 MADRONO
numbers in the, 236; confertiflorum, 237; lanatum var. achillaeoides, 237; l. var. aphanactis, 237; 1. var. arach- noideum, 237; 1. var. croceum, 237; l. var. cuneatum, 237; 1. var. grandi- florum, 237; l. var. integrifolium, 237; ]. var. 1., 237; 1. var. lanceolatum, 237; l. var. leucophyllum, 237
Ernst, W. R.: Review, Flora Europaea, 62
Escobaria leei Bodeker rediscovered in New Mexico, 137
Eucalyptus globulus, Volatile growth in- hibitors produced by, 207
Euphrasia: in the Pacific Northwest, Oc- currence of the genus, 160; americana, 160; canadensis, 160
Fire and sodium-calcium borate on chaparral vegetation, Influences of, 193
Franseria dumosa, 241
Fryxell, P. A.: A further description of Gossypium trilobum, 113
Gagae, 148
Galinsoga: calva, 204; hispida, 204; mandonii, 206; parviflora, 205; urtici- folia, 205
Galium: species of California (Rubia- ceae), The fleshy-fruited, 105; ambi- guum, 112; a. var. siskiyouense, 112; andrewsii, 112; a. var. gatense, 112; bo- landeri, 112; californicum, 112; c. ssp. luciense, ssp. nov., 107; clementis, 112; cliftonsmithii, comb. nov., 112; grande, 112; hardhamiae, 112; marti- rense, sp. nov., 110; miguelense, 105; muricatum, 112; nuttallii, 111; n. var. cliftonsmithii, 107 ; pubens, 112; sparsi- florum, 111
Ganders, F. R.: Occurrence of the genus Euphrasia in the Pacific Northwest, 160
Ganysma, 168
Gelidium sinicola, 176
Geranium: microphyllum in California, The occurrence of a New Zealand plant, 213; pilosum, 213; retrorsum, 213; sibiricum, 213
Gossypium: trilobum, A further descrip- tion of, 113; gossypioides, 118; lancei- forme, 117; thespesioides, 117
Grasses of the Pacific Coast, Notes on some, 250
Gutierrezia: |(Compositae — Astereae), The California species of, 75; brac- teata, 81; californica, 81; microce- phala 81; sarothrae, 81
Haplopappus: macronema, 225; suffruti- cosus, 225
Hawksworth, F. G.: Arceuthobium dou- glasii in Nevada and Wyoming, 63; and D. Wiens: Observations on witches’-broom formation, autopara-
[Vol. 18
sitism, and new hosts in Phoraden- dron, 218
Helecochloa schoenoides, 22
Herbarium: A Reed College, 31; of Howard E. McMinn, The of, 128; specimens, Labels for, 127; supplies, 160
Hesperolinon, 181
Heuchera californica, 159
Hibiscus ingenhousii, 117
Hordeum: jubatum complex, Notes on the, 243; brachyantherum, 243
Hymenena multiloba, 178
Hymenoclea salsola, 241
Isomeris arborea, 241 Jouvea, 36
Kasapligil, B.: The herbarium of Howard E. McMinn, 128
Kaokochloa nigrirostis, 27
Kern, P.: The genus Tiarella in western North America, 152
Kowalski, D. T.: New records of Myxo- mycetes from California I, 140
Larrea divaricata, 241
Leucocoryne, 148
Licea pusilla, 141
Linum: Some new chromosome numbers in, 179; alatum, 179; arenicola, 179; aristatum, 179; australe, 179; baha- mense, 179; catharticum, 181, digy- num, 182; floridanum var. chrysocar- pum, 180; greggii, 179; hypericifolium, 180; kingii, 180; lewisii, 179; medium var. m., 179; m. var. texanum, 179; mexicanum, 180; neomexicanum, 180; pringlei, 179; rigidum, 180; rupestre, 179; schiedeanum, 179; striatum, 179; sulcatum, 179; virginianum, 179
Lipactinia, 129
Liptochaeta hastata, 133
Little, E. L.: Varietal transfers in Cu- pressus and Chamaecyparis, 161
Lloyd, R. M.: Cinal patterns in frond anatomy of Polypodium, 65; Review, The California deserts, 250
Lupines, The inter-relationships of the Alaskan, 1
Lupinus: albicaulis, 5; arboreus, 5; ar- bustus ssp. neolaxiflorus, 4; a. ssp. pseudoparviflorus, 4; arcticus ssp. subalpinus, comb. nov., 16; a. ssp. a., 11; argenteus, 16; burkei, 11; cottonii, 5; donnellyensis, 13; gakonensis, 11; glacialis, 16; grandifolius, 6; humicola, 5; kushei, 2; latifolius, 5; 1. var. canadensis, 16; |. var. subalpinus, 16; lepidus, 5; littoralis, 11; lyallii, 2; magnus, 6; matanuskensis, 6; multi- caulis, 13; multiflorus, 13; nootkaten- sis var. ethel-loofiae, 8; n. var. fruti- cosus, 10; n. var. glaber, 10; n. var. henry-looffii, 8; n. var. kjellmannii,
1966]
11; n. f. leucanthus, 8; n. var. n., 8; n. var. perlanatus, 8; n. var. unalas- kensis, 10; ornatus, 5; pallidipes, 2; perennis, 5; p. ssp. nootkatensis, 8; polyphyllus ssp. arcticus, 13; p. var. grandifolius, 6; p. var. pallidipes, 4; p. ssp. p., 6; p. ssp. suberbus, 5; prunophilus, 4; pseudopolyphyllus, 6; relictus, 13; rivularis, 5; sellulus, 2; stationis, 6; subalpinus, 16; volcanicus var. rupestricola, 16; wyethii, 5; yukonensis, 11
McGregor, E. A.: Dr. La Roy Abrams’ collecting trip of 1908, 97
McMinn, The herbarium of Howard E., 128
Membranoptera multiramosa, 178
Mertens, T. R. and P. H. Raven: Taxon- omy of Polygonum, section Polygo- num (Avicularia) in North America, 85
Michigan Botanist, 17
Monanthochloe, 33
Monarda: (Labiatae), New taxa in the genus, 119; austromontana, 120; bart- letti, 119; citriodora ssp. c. var. parva, var. nov., 120; c. ssp. c. var. attenuata, var. nov., 121; c. ssp. austromontana, comb. nov., 120; dressleri, sp. nov., 119; eplingiana, 119; fistulosa, 119; malloryi, 119; pringlei, 119
Monolopia, 239
Mooring, J. S.: Chromosome numbers in the Eriophyllum lanatum (Compo- sitae, Helenieae) complex, 236
Muhlengergia: Notes on Mexican Grasses V. Two gypophilous species of, 185; 33; gypsophila, sp. nov., 186; pur- pusii, 190; villiflora, 185
Muilla: with a remark on the taxonomic position of the genus, Contribution to the embryology of, 143; maritima, 143
Myriogramme hollenbergii, 178
Myxomycetes from California I, New records of, 140
Neeragrostis reptans, 26
Neomammillaria leei, 138
Neostapfia colusana, 27
Niehaus, T. F.: A new species of Bro- diaea from San Clemente Island, Cali- fornia, 233
Norman, E. M.: A reevaluation of Bud- dleia corrugata, 92
Norris, R. E. and J. West: Notes on Marine algae of Washington and southern British Columbia, 176
Notes and news: 17, 31, 63, 104, 127, 160, 173, 184, 192, 213, 224, 250
Norhoscordum fragrans, 149
Oakesia sessilifolia, 213 Orcuttia: californica, 21; c. aequalis, 27; c. var.
var. in- viscida, 27;
INDEX 250
greenei, 27; mucronata, 27; pilosa, 27; tenius, 27
Orcuttieae: and the subtribes of the Pappophoreae (Gramineae), The tribe, 18; tribus nov., 20
Oregonium, 168
Ornduff, R.: Review, Vascular plants of the Pacific Northwest, Part 2, 30; A Reed College herbarium, 32
Pappophoreae (Gramineae), the tribe Orcuttieae and the subtribes of the, 18
Pappophorum: bicolor, 27; commune; 22; elegans 24; mucronulatum 27; pappiferum, 27
Pedunculata, 168
Peponapis timberlakei, 46
Perichaena vermicularis, 141
Petalosteira: laciniata, 158; unifoliata, 159
Phoradendron: Observations on witches’- broom formation, autoparasitism, and new hosts in, 218; bolleanum ssp. b., 221; b. ssp. densum, 222; b. ssp. pauci- florum, 221; californicum, 220; juni- perinum ssp. j., 223; serotinum, 220; tomentosum ssp. macrophyllum, 220; t. ssp. t., 219; villosum ssp. coryae, DLL NSS a Via OU
Phrygilanthus tetrandrus, 218
Pinus washoensis, 63
Platysiphonia clevelandii, 178
Poa texana, 34
Polygonum: section Polygonum (Avicu- laria) in North America, Taxonomy of, 85; acadiense, 87; achoreum, 88; aequale, 89; arenarium, 85; arena- strum, 89; argyrocoleon, 85; allocar- pum, 89; autumnale, 91; aviculare, 89; a. var. erectum, 88; bellardii, 85; boreale, 91; buxiforme, 91; erectum, 88; exsertum, 85; flowleri, 88; glau- cum, 86; heterophyllum, 89; lanatum, 91; littorale, 91; maritimum, 86; marinense, sp. nov., 87; monterey- ense, 91; neglectum, 91; oxyspermum, 87; ©. ssp. rail, 87; patulum, 85; polycnemoides, 91; prolificum, 86; raii, 87; ramosissimum, 85; robertii, 88; rurivagum, 89; tenue, 85
Polypodium: Clinal patters in frond anatomy of, 65; califoricum var. inter- medium, 65; c. var. kaulfussii, 65
Polysiphonia brodiaei, 178
Pommereulla, 66
Porphyropsis, 176
Prosopis juliflora, 241
Pterophyton: 129; 214
Punctaria chartacea 176
Rajhathy, T.: Notes on the Hordeum jubatum complex, 243
Raven, P. H. and T. R. Mertens: Cro- cidium in San Luis Obispo Co., Cali- fornia, 32
256 MADRONO
Reed College Herbarium, 32
Reeder, C. G. and J. R. Reeder: Notes on Mexican Grasses V. Two gypso- philous species of Muhlenbergia, 185
Reeder, J. R.: The tribe Orcuttieae and the subtribes of the Pappophoreae (Gramineae), 18
Reederochloa, 33
Reveal, J. L.: On the specific distinction of Eriogonum nutans and collinum, 167
Reviews: Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson, Vascular plants of the Pacific North- west, Part 2, 30; Howitt, B. F. and J. T. Howell, The vascular plants of Monterey County, California, 127; Hunt Botanical Library and its publi- caitons, 246; Jaeger, E. C., The Cali- fornia deserts, 250; Langman, I. K., A selected guide to the literature on the flowering plants of Mexico, 126; McClintock, E. and V. Moore, Trees of the Panhandle, Golden Gate Park, San Francisco, 173; Riley, H.. P; Families of Flowering plants of south- ern Africa, 29; Savi, T., Naming the living world, 184; Schultes, R. E. and A. S. Pease, Generic names of orchids, 30; Schwanitz, F., The origin of culti- vated plants, 249; Steyermark, J. A., Flora of Missouri, 28; Tutin, T. G., V. H. Heywood, N. A. Burges, D. H. Valentine, S. M. Walters, and D. A. Webb, Flora Europaea, Vol. 1, 62
Rhodoglossum reseum, 177
Rhodophysema: elegans, 177; georgi, 177
Rodin, R. J.: Review, Families of flow- ering plants of southern Africa, 29
Rogers, C. M.: Sclerolinon, a new genus in the Linaceae, 181; and B. D. Harris, Some new chromosome num- bers in Linum, 179
Sabazia urticifolia, 205
Schmidtia pappophoroides, 27
Sclerolinon: a new genus in the Lina- ceae, 181; gen. nov., 182; digynum, comb. nov., 182
Scoliopus: (Liliaceae), The chromosomes of, 211; bigelovii, 211; hallii, 211
Scora, R. W.: New taxa in the genus Monarda (Labiatae), 119
Selera gossypioides, 118
Sielingia wrightii, 34
Simmondsia chinensis, 241
Soderstrom, T. R. and H. F. Decker; Allolepis: a new segregate of Distichlis (Gramineae), 33
Solbrig, O. T.: The California species of Gutierrezia (Compositate—Astereae), 15)
Sonoricola, 129 Spermothamnion snyderae, 177
[Vol. 18 Sporobolus, 33 Stapfia, 18 Stemmatella (Compositae — Helian-
theae), The taxonomic status of, 203; congests, 204; lehmanii, 205; sodiroi; urticifolia, 205; u. var. eglandulosa, 206; u. var. venezuelensis, 205
Stemonitis: hyperopta, 141; pallida, 142
Stocking, S. K.: Influences of fire and sodium-calcium borate on chaparral vegetation, 193
Szweykowski, J.: Transolocation hetero- sygosity in Croton californicus, 31
Thespesia thurberi, 117
Thomas, J. H.: Review, The vascular plants of Monterey County, Califor- nia, 127; Labels for herbarium speci- mens, 127; Review, The Hunt Bo- tanical Library and its publications, 246
Thurberia: 17
Thuretellopsis peggiana, 176
Tiarella: in western North America, The genus, 152; californica, 159; rhombi- folia, 158; stenopetala, 158; trifoliata var. laciniata, 158; t. ssp. unifoliata, comb. nov., 159; t. var. unifoliata, 159; unifoliata, 159; u. var. procera, 159; u. f. trisecta, 159
thespesioides, 117; triloba,
Tridens, 33
Turner, M. L.: The taxonomic status of Stemmatella (Compositae — Helian- theae), 203
United States National Herbarium, 31
Verbesinaria, 217
Vilfa glomerata, 251
Verbesina: L. (Compositae), A taxo- nomic revision of section Sonoricola of the genus, 129; (Compositae with taxonomic notes on other species, Two new species of, 214; apiculata, sp. nov., 214; cayucensis, 135; chihua- huensis, 134; dissita, 131; encelioides, 135; erosa, 136; hastata, 133; lepto- chaeta, 134; lindheimeri, 131; oligo- cephala, 132; ovata, 217; palmeri, 132; parrasana, 134; peninsularis, 135; pustulata, 136; pterocaulon, 217; rosei, 214; scabra, 217; stenophylla, 217; tequilana, sp. nov., 215; tetrap- tera, 217; venosa, 133
Whitaker, T. W.: Review, The origin of cultivated plants, 249
Wiborgia urticifolia, 205
Wilson, R. C. and R. J. Vogl: Man- zanita chaparral in the Santa Ana Mountains, California, 47
Xenoglossa 46 Ximenesia, 129
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MADRONO
VOLUME 19, NUMBER 1 JANUARY, 1967
Contents
A REVISION OF PETALONYX (LOASACEAE) WITH A CON- SIDERATION OF AFFINITIES IN SUBFAMILY GRONOVI- OIDEAE, William S. Davis and Henry J. Thompson 1
MAcNA4AsB Cypress IN NORTHERN CALIFORNIA: A GEOGRAPHICAL REvIEW, J. R. Griffin and C. O. Stone 19
A NEw SPECIES OF PRIMROSE FROM NEVADA, Noel H. Holmgren 27
REviEws: Olga Reifschneider, Biographies of Nevada Botanists (Joseph Ewan) 30
A WEST AMERICAN JOURNAL OF BOTANY
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A REVISION OF PETALONYX (LOASACEAE) WITH A CONSIDERATION OF AFFINITIES IN SUBFAMILY GRONOVIOIDEAE
WILLIAM S. Davis and HENRY J. THOMPSON
Petalonyx is a genus of shrubs and sub-shrubs of the desert and semi- arid regions of the southwestern United States and Northern Mexico. Although a few modern floras give good accounts of some of the species, no monograph treating the entire genus has been prepared. Apparently Petalonyx has not invited the attention of monographers because the species are relatively distinct and hence new collections are not difficult to identify. Nevertheless, it is remarkable that taxonomists have been content merely to describe the flowers of Petalonyx as having the stamens located outside the corolla tube without attempting to under- stand such unorthodox morphology. Our attention was first drawn to Petalonyx during the general survey of the family to determine generic limits preparatory to a more detailed study of Mentzelia. Although we encountered no difficulty in delimiting Mentzelia from Petalonyx we did raise some questions concerning the generic affinities of Petalonyx with the other members of subfam. Gronovioideae. In addition to the five species of Petalonyx, subfam. Gronovioideae accommodates the one species of Cevallia, the two species of Gronovia, and the one species of Fuertesia (Urban and Gilg, 1900). Although our interest has been centered on Petalonyx we have extended our observations to the other genera of this subfamily whenever possible.
Four of the five species of Petalonyx have been studied as living plants in natural populations and only P. crenatus of north central Mexico remains unknown to us as living material. Cevallia has been studied in two natural populations and one species of Gronovia has been grown in our greenhouse. Thus we have had living material of six of the nine species of subfam. Gronovioideae, making cytological and ana- tomical studies possible. Chromosomes were examined by fixing anthers in 3:1 ethanol-acetic acid, squashing in acetocarmine, and observing with a phase microscope. Anatomical studies were made from material either embedded in paraffin and sectioned or simply cleared in lactic acid. Voucher specimens for our chromosome number determinations are on file in the herbarium of the University of California, Los Angeles (LA). We have examined all the specimens of Petalonyx from the fol- lowing herbaria: BAYLU, DAS, DS, GH, LA, LL, POM, RSA, SMU, TAES, TEX, US. The manuscript benefited greatly from the critical comments of Peter H. Raven and Wallace R. Ernst. This study was sup- ported by Grant GB-1337 from the National Science Foundation.
Maprono, Vol. 19, No. 1, pp. 1-32, January 31, 1967. 1
2 MADRONO [Vol. 19
PETALONYX
Some of the features of Petalonyx are of particular interest and will be singled out here for detailed consideration while a later section of this paper will present a formal taxonomic treatment with descriptions and keys. The flowers of the five species are borne in naked or bracted racemes and are generally similar in their inferior, unilocular ovary with a single, pendulous ovule. No matrix tissue proliferates in the lumen between the developing seed and the ovary wall and there is no disc at the apex of the ovary. The fruit is indehiscent and the mature seed is pyriform, lacks endosperm, and has a smooth to finely striated testa. The cotyledons are ovate and the inner surfaces are smooth. The simple style and stigma and the calyx, which is deeply divided into five lanceo- late lobes, also are quite uniform throughout the genus. In contrast, the petals and stamens exhibit some unusual features that distinguish some of the species. Petalonyx nitidus, P. thurberi, and P. parryi are distinctive by their apparently sympetalous corollas with the anthers outside the corolla tube (fig. 1). Petalonyx crenatus is unique in having only two functional stamens and the flower is thus bilaterally symmetri- cal (fig. 1). Petalonyx linearis remains distinct by its orthodoxy for it lacks these unusual characters. Certain morphological features, whose modifications provide these unusual floral differences, are present throughout the genus. The petals in all species are clawed and each side of the claw has two narrow flaps of tissue, one inner and one outer, that extend laterally toward the adjacent petals (fig. 2e). The stamens are not in themselves unusual, being composed of a slender filament with a terminal, 4-chambered anther, however the filaments arise from the short hypanthium opposite the sepals and between, but apparently in the same whorl with, the petals (fig. 2a). In bud the low portions of the filaments are held between the inner and outer flaps of adjacent petal claws (fig. 2b-f) while the distal portion of the filaments and the anthers are enclosed by the petal limbs (fig. 2g). In P. thurberi, P. nitidus, and P. parryi the inner flaps of tissue of adjacent petals remain free near the base (fig. 2b) but become firmly connivent along the upper half of the claw thus forming a corolla tube (fig. 2c-e). As the buds open, the staminal filaments are freed to incline outward while the petals remain held together—hence the unusual con- dition of the stamens outside the corolla tube. Although the claws become strongly connivent and sometimes can be pulled apart only by tearing tissue, it is clear that they are secondarily adhered together because the cuticle that covers the epidermis is continuous through the point of connection of adjacent claws. Although the petal claws in P. linearis and P. crenatus have the flaps of tissue, they do not become connivent in these species, no tube is formed, and the stamens are not held outside of the corolla.
The cross sections of the flowers also revealed a difference in the vascularization of the style. In P. thurberi, P. nitidus, and P. parryi
1967] DAVIS & THOMPSON: PETALONYX 5
nitidus
thurberi
crenatus linearis
Fic. 1. Drawings of flowers and bractlets of the five species of Petalonyx. The petals in P. nztidus are just opening, all others with petals fully expanded, 4.
there are three vascular traces while in P. linearis and P. crenatus there are only two. These differences were noted in the examination of about 15 flowers of the five species with only one flower of P. parryi being anomolous in having four traces in the style.
Plants of the five species of Petalonyx are perennial but the growth form characteristic of each species ranges from woody only at the very base, as in P. thurberi, to a well developed but small shrub, as in
4 MADRONO [Vol. 19
P. parryi. The leaves are simple but have a number of consistent dif- ferences of shape and margin, which distinguish the species.
Like most members of the Loasaceae, the plants of Petalonyx are covered with a harsh pubescence. Petalonyx has a type of trichome that also is found in all plants of the family, but unlike many genera, Petalonyx only has this “basic” type (fig. 6a, b). These trichomes are found in somewhat modified forms on different parts of the same indi- vidual and all of the taxa can be recognized by at least slight dif- ferences in the pubescence. Petalonyx linearis is most distinct in its pubescence in having short, truncate trichomes on the stems which contrast sharply with the long slender trichomes of the bracts and ovary. The pubescence of P. parryi and P. nitidus is quite similar but the stem pubescence in P. parryi is straight while in P. nitidus it is reflexed. The two subspecies of P. thurberi are distinguished primarily by a dif- ference in pubescence with ssp. gilmani having a finer, longer, and more dense pubescence than ssp. thurbert.
Meiosis has been observed in plants from six populations representing four of the five species of Petalonyx, including both ssp. of P. thurbert, with only P. crenatus remaining unknown chromosomally. All of the plants examined showed 23 pairs of chromosomes. Metaphase bivalents are approximately 3 micra long although in every cell examined there were slight but obvious differences in the size of the bivalents so that a continuous series from smaller to larger bivalents could be noted. Chromosome observations of Petalonyx, were made on the following collections: P. linearis, n = 23, N of San Felipe, Baja California, Mexico, Raven 14791; P. nitidus, n = 23, Panamint Springs, Inyo Co., Cali- fornia, Thompson 3273; P. parryi, n = 23, near Glendale, Clark Co., Nevada, Thompson 3274; P. thurberi ssp. thurberi, n = 23, 10 miles W of Kingman, Mohave Co., Arizona, Thompson 3294; 9 miles W of Randsberg, Kern Co., California, Kyhos, May 12, 1962; P. thurberi ssp. gilmanit, n = 23, Ryan Wash, Death Valley National Monument, California, Davis 170.
Information about the breeding system in Petalonyx is limited to that suggested by flower structure and seed set in natural populations. In P. linearis the stigma is located among the anthers at the time pollen is shed (fig. 1) and self-pollination in each flower seems inescapable. In the other four species the anthers are positioned well away from the stigma when the pollen is shed (fig. 1) and some vector must be necessary for pollination. In the herbarium specimens of P. linearis over 90 percent of the ovaries contain a seed while in the other four species seed set frequently is about 50 percent. This information, to- gether with the floral morphology, suggests that P. linearis is highly self-pollinated, self-compatible, and hence an inbreeding species while the other four species are more highly outbreeding. Timberlake (1962, pp. 14, 36-38) suggests that the bees Perdita exilis Timberlake and P. crandalli Timberlake probably are oligolectic on Petalonyx thurbert
1967] DAVIS & THOMPSON: PETALONYX 5
Fic. 2. Transverse sections through a flower of Petalonyx thurberi before anthesis (filaments stippled): a, section through the calyx tube and upper portion of the ovary; b, section through an area just above (a); c, section through the calyx lobes, petal claws, and style; d, section through an area above (c); e, section through an area above (qd); f, section through an area above (e); g, section through an area above (f) showing the calyx lobes, petal limbs, filaments, and style.
and they may be important pollinators. The flowers of P. thurberi, P. nitidus, and P. parryi are lightly fragrant and attract a variety of insects, many of which could effect pollination.
Although all five species of Petalonyx are restricted to the arid regions of the southwestern United States and northern Mexico we have been unable to find two species growing in mixed or adjacent populations. This is because of the very limited ranges of two of the
6 MADRONO [Vol. 19
P crenatus
Fic. 3. Distribution of Petalonyx crenatus.
species, their different ecological requirements, and the small size of local populations. Petalonyx crenatus, morphologically very distinct with its slightly irregular flowers, narrow petals, and distinctive crenate leaves also is geographically isolated, growing only in western Coahuila, Mexico, where it is disjunct by 500 miles from the other species of the genus (fig. 3). Petalonyx linearis, also morphologically very distinct, occurs in sandy areas of the lower Colorado Valley and Vizcaino regions of the Sonoran Desert (fig. 4) usually below 2500 feet elevation. The autogamous breeding habit and restriction to the geologically new desert habitats suggest a derivative status for P. linearis. Petalonyx thurberi also ranges into the Sonoran Desert and seems so similar to P. linearis in habitat preference that we are forced to ascribe our in- ability to find them growing together merely to chance. Petalonyx thurberi, P. nitidus, and P. parryi are not only morphologically similar to each other but they occur in the same general area (fig. 5). Petalonyx thurberi occurs in sandy areas and washes of the Mojave Desert and extends into the upper regions of the Sonoran Desert. It usually occurs at elevations between 1000 and 3000 feet although stations above and below these limits are not exceptional. Petalonyx thurberi ssp. gilmanu is restricted to the Death Valley region. Petalonyx nitidus occurs on more rocky soils in the Mojave Desert, usually above 3000 feet eleva- tion and extending as high as 7000 feet. Where P. thurberi and P. nitidus occur in the same area we have found P. nitidus at higher elevations than P. thurberi. Petalonyx parryi is restricted to a local area near the Virgin and Colorado rivers in southern Nevada and adjacent Utah and Arizona (fig. 5). It is found at elevations below 3000 feet and is prob- ably restricted to white to grey clay soils.
1967] DAVIS & THOMPSON: PETALONYX 7
P linearis
Fic. 4. Distribution of Petalonyx linearis.
The species of Petalonyx obviously are similar in many respects and the differences between them can be understood as variations on a basic theme. The question of which other group of populations in the Loasa- ceae is most similar to Petalonyx is not so readily answered. Similarity to Cevallia, Gronovia, and Fuertesia is suggested by the grouping of these genera with Petalonyx to form subfam. Gronovioideae on the basis of the single, pendulous ovule and the reduced number of stamens common to all four genera.
CEVALLIA
The single species of this genus, C. sinuata Lag., occurs in northern and eastern Mexico and the southwestern United States. The plants are low herbaceous perennials with only the basal stems becoming woody. The leaves are sinuate-pinnatifid and, like all members of Loasaceae, the plants are covered with a harsh pubescence. There are three dif- ferent types of trichomes, all one-celled, which may occur side by side on a single organ. Some of the trichomes are like the basic type found
MADRONO
[Vol. 19 So ! | ~ . \ N E V A OD nN me: oN O Up ae a AGRA | Oo’. O | earth, ores ee ae | _ OO 1@0; O eo Stole pee ee a xO '@ wae 8 : O 7. 56 @N fe) P| | Oo”; Ooms 6 ‘s xe O g | tC) eo a | * Vek pla: ke ke A) ORe WeZe 40" CN PA | * ee * rs Oo | i | —s * | ae mae * | eae n : * \ A 5 | * x aoe | ee ae at " ! — + xy | fri ey ee ** | BAJA Seek | is | * se ee *« Ch P nitidus
P parryl P thurberi
*
subsp. thurberl
subsp. gilmanil
Fic. 5. Distribution of Petalonyx nitidus, P. parryi, and P. thurberi.
in Petalonyx and throughout Loasaceae (fig. 6a, b). A second trichome type is dendritic, 0.1 mm long, and forms a dense tomentum over much of the plant (fig. of). A third trichome type is up to 3 mm long, straight, without barbs but with a very acute apex (fig. 6e). The base of these trichomes is buttressed by a ring of numerous columnar epidermal cells. These trichomes probably are the ones that so readily penetrate human skin and account for the painful, stinging sensation when the plant is touched. The first two types of trichomes lack the basal, columnar cells that surround the base of the long trichome.
The flowers are clustered in tight heads at the ends of peduncles that are borne opposite the leaves. The ovary is inferior and produces one, pendulous seed that lacks endosperm. The cotyledons are ovate and smooth on the inner surface. The fruit is indehiscent with the sepals and petals persistent. There is no matrix tissue that develops about the
1967] DAVIS & THOMPSON: PETALONYX 9 dq b ae P ceo am as!
Fic. 6. Diagrams of longitudinal sections of trichomes in subfam. Gronovioideae: a, b, Petalonyx, 50X; c, Fuertesia, 20x; d, Gronovia, 20X; e, Cevallia, 20; f, Cevallia, 15X.
ovule as in Gronovia. Atop the ovary is a short hypanthium from the outer edge of which arise ten similar segments, but there is no disc within the hypanthium. The five segments opposite the stamens usually are considered calyx-lobes while the other five are considered petals. Both series are erect, linear, very pubescent, and plumose by virtue of the long trichomes along the margins. When the flower opens and pollen is shed the petals are held inward about the stamens while the sepals are turned slightly outward. Both petals and sepals are yellow on the inner surface but the petals are somewhat brighter in color and are slightly less plumose. The five stamens arise from the top of the hypanthium with short, thick filaments and the anther is borne on the adaxial surface of the filament with the basal portion of the anther free. The connective extends well beyond the anther as a hollow, thin- walled, somewhat inflated appendage. The style is short and topped by a conical stigma that is positioned opposite the base of the anthers. Meiotic chromosomes have been observed in two plants from two dif- ferent populations and each had 13 pairs of chromosomes. The chromo- somes are all the same size and the metaphase bivalents are nearly five micra long. Chromosome observations of Cevallia were made on the following collections: C. sinuata, n = 13, 14 miles S of Mammoth, Pinal Co., Arizona, Thompson 3221; 33 miles N of Van Horn, Culbert- son Co., Texas, Thompson 3286. In the Van Horn population the flowers were noted to have a faint fragrance and were visited by a great variety of insects, particularly bees and butterflies. Flower struc- ture makes self-pollination inevitable but outcrossing might still obtain if the plants are self-incompatible.
GRONOVIA
The two species of this genus, G. scandens L. and G. grandiflora Urban & Gilg, are readily distinguished by differences in the size and proportions of the flowers but otherwise are so similar that for our purpose of characterizing the genus we need not distinguish between
10 MADRONO [Vol. 19
them. Gronovia occurs throughout much of Tropical America from Venezuela and Ecuador to northern Mexico. The plants are herbaceous perennial vines with long petioled, simple leaves. The blades are broadly ovate with cordate bases and coarsely toothed, often 3—5 lobed margins. In addition to having the typical trichomes found throughout Loasaceae, (fig. 6a, b) the plants have a very characteristic trichome type that is about 3 mm long and is unarmed except that the apex is formed into a double hook (fig. 6d). These hairs are supported by a mass of columnar cells that surround the basal portion of the trichome cell and raise the base of this cell well above the level of the plane of the epidermis. This type of trichome occurs only on the stems and the ad- joining portion of the petioles. The inflorescence is open and borne on peduncles that are opposite the leaves. The ovary is inferior, is topped by a disc with a crenate margin, and has one pendulous ovule. The single ovule is surrounded by a matrix of spongy tissue which is compacted as the seed enlarges and fills the ovary cavity after the seed is formed. The fruit develops five prominent, longitudinal ridges and is in- deshiscent. The seed lacks endosperm and the seed coat is membranous and unornamented. The cotyledons are ovate and their inner surfaces are strongly rugose. The ovary is topped by a short hypanthium from the rim of which extend five calyx-lobes. The petals are free, narrow, and rather inconspicuous. The stamens have slender filaments and a terminal, 4-celled anther. The style is slender and has a small, capitate stigma that is situated among the anthers. Meiotic chromosomes of one plant of Gronovia scandens have been examined. The 37 pairs of chromosomes were of equal size and somewhat smaller than those of Cevallia but larger than the chromosomes of Petalonyx. Chromosome observations were made from the following collection: G. scandens, n — 37, cultivated strain grown from seed obtained from the Botanical Garden, Copenhagen, culture number at Los Angeles, Thompson 3234. A mitotic chromosome count of 2n = 76 was reported by Hamel (1938) for G. scandens growing in the Jardin des Plantes at Paris. We have readily cultured G. scandens in our greenhouse. The position of the anthers and stigma insure self-pollination and the formation of many viable seeds on the plants cultured in our greenhouse indicates that the plants are self-compatible.
FUERTESIA
This genus contains a single species, F. domingensis Urban, which is restricted to the island of Hispaniola in the West Indies. It differs from Gronovia in having lobed petals, an entire margined disc on the apex of the ovary, and a simple stigma, but in general appearance it is like Gronovia. The pubescence of Fuertesia consists of two different types of trichomes in addition to the basic type found in other genera of Loasaceae. The first type is like the double-hooked trichome found on the stem and basal portions of the petioles of Gronovia (fig. 6d) but
1967] DAVIS & THOMPSON: PETALONYX 11
is shorter with a few barbs along its length. The second type of trichome is unique to Fuertesia and has been found only on the leaves (fig. 6c).
SUBFAMILY GRONOVIOIDEAE
Although our considerations of Cevallia, Gronovia, and Fuertsia were only peripheral to our monograph of Petalonyx, we feel some authority to comment on the affinities of these genera within the Loasaceae.
Gronovia and Fuertesia obviously are more similar to each other than to any other genera. They have similar forms, similar floral morphology, and share the unique bifid trichome which occurs nowhere else in the family. Although these two genera may be considered closely related, no such comparable relationship is indicated among the other genera. Petalonyx, Cevallia, and Gronovia were grouped by Urban and Gilg. in subfam. Gronovioideae by giving great weight to two characters: stamens 5 in number and ovary with a single seed. Emphasis of these two characters gives a distorted representation of the affinities among these genera. For example, while Petalonyx and Cevallia are similar in having 5 stamens, and thus are unlike all species of subfam. Ment- zelioideae and subfam. Loasoideae, a different view is reached when the individual stamens are compared. In stamen morphology Petalonyx and Cevallia are more different from each other than either of these two genera are from any other genera in the family. Similar evaluations may be made about the number of seeds per capsule. Several species (e.g., Mentzelia oligosperma, Fissenia sp.) have the seeds reduced to 1-3 per capsule. Reduction in seed number occurs in conjunction with indehiscent capsules and small flowers in many otherwise distinct groups in Loasaceae. Furthermore, the information from chromosome numbers does not indicate any relationships among any of the genera currently grouped in subfam. Gronovioideae.
We consider Petalonyx, Cevallia, and the Gronovia-Fuertesia alli- ance, which now comprise subfam. Gronovioideae, to be three divergent groups whose affinities lie with other genera in the family and not with each other. However, until a more thorough study is made of all of the genera in this family none of the details of these similarities can be recorded in a formal taxonomic form and we must be content merely to have pointed out the very artificial basis of subfam. Gronovoideae.
TAXONOMY OF PETALONYX
PETALONYX A. Gray, Mem. Am. Acad. 5:319. 1855. Shrubs or sub- shrubs; leaves alternate, petioled or sessile, entire, toothed or crenate; inflorescence a naked or bracted raceme, or spicate raceme; each flower involucrate with three green bractlets at the summit of the pedicel, two of which are smaller and lateral; sepals joined to form a tube adnate to the ovary, cleft almost to the base, the lobes linear to lance-linear, inner surface glabrous, outer hispid, persistent or irregularly deciduous; petals cream to white, clawed, inner surface glabrous, outer hispid;
12 MADRONO [Vol. 19
claws free or connivent to form a tube; limbs usually reflexed; stamens free, originating between, and apparently in the same whorl as the petals, outside the petals at anthesis and in bud curving inside and covered by the overlapping petals; anthers 4-celled; pollen elliptic, tricolpate, unornamented; style filamentous; stigma simple; mature fruit a uniloculate utricle, cylindrical; constricted near the apex, then flaring above, 3-5 nerved, with brittle, usually ribbed walls; ovule 1, without endosperm, pendulous; seed pointed at the hilum, unornamented or with fine striations on the testa; embryo with thick, plano-convex cotyledons and a short, acute, radicle. Type Species: Petalonyx thurbert A. Gray.
KEY TO THE SPECIES Petal distinct.
Anther bearing stamens 2, about twice as long as the petals, sterile filaments 3, about as long as the petals; leaf margins revolutely crenate; bractlets oblong, truncate at the base; petal limb less than twice a wide as the claw; Coahuila, Mexico. . . we Se aPorenaints,
Anther bearing stamens 5, equaling the asl in ienethe leaf margins entire or remotely dentate; bractlets ovate, the base cordate; petal limbs at least twice as wide as the claw: deserts of California, western Arizona, Baja California, Sonora, and the Islands of the Gulf of California
2. P. linearis Petals connivent, the claws forming a tube.
Leaves sessile, reduced upward along the stems; petals shorter than 5 mm; widely distributed in the desert areas of southern California, Baja California and Arizona. . . . 2 etna iunoent
Leaves petiolate, similar in size along the peme: pe rare longer than 5 mm.
Sub-shrubs, predominately herbaceous, current seasons growth longer than 10 cm; leaf margins coarsely few-toothed; inflorescence a bracted raceme; petals shorter than 11 mm; Inyo and San Bernardino of California to southern Nevada and northern Arizona above 3000 feet elevation. . . ey ee Ae Bie us
Shrubs, predominately woody, caren seasons ion less than 10 cm long; leaf margins entire to crenate; inflorescence a naked, terminal raceme; petals longer than 11 mm; Coconino and Mohave coun- ties of Arizona to southern Utah and Nevada, usually on white to grey clay soils and below 3000 feet elevation. . . 5. P. parryi
1. PETALONYX CRENATUS A. Gray ex Watson, Proc. Am. Acad. 17:358. 1882. Suffruticose, with short, erect, brittle, herbaceous branchlets, up to 10 dm high; stems of the current season 10-20 cm long, to 6 mm wide, surface smooth, grey-green or yellow-green, densely hispid with long, pointed, antrorsely barbed, retrorsely appressed hair; older stems straw-colored to dark grey; leaves sessile, similar in size along the stems, 0.6-1 cm long, 0.2-0.4 cm wide, oblong-lanceolate, obtuse, cordate, revolutely crenate, green, hispid with fine, long, antrorsely barbed hairs on both surfaces, and short, truncate retrorsely barbed hair along the margins; inflorescence a terminal, naked raceme, 20-40 flowered, to 5 cm long, elongated in fruit, the lower fruit fallen; pedicels
1967] DAVIS & THOMPSON: PETALONYX 13
1.0-2.5 mm long; large bractlet 3-5 mm long, 1.5—2.5 mm wide, triangu- lar-ovate to oblong with a prominent midrib before maturity, acute, truncate to sub-cordate, crenate basally, hispid with long, pointed, antrorsely barbed hair; lateral bractlets 2-3 mm long, 1.5-—2 mm wide, ovate to triangular-ovate, pinnately cleft, hispid; sepal lobes 1.5—2.5 mm long, 0.5—0.9 mm wide, recurved, persistent; petals white, linear to narrowly spatulate, acute, claw not sharply delimited from the limb, 4-7 mm long, 1-1.5 mm wide, not connivent; claw 0.7—-0.9 mm wide, with short, epidermal papillae; two fertile stamens, well exserted, 10-11 mm long, three sterile stamens 5—9 mm long; filaments with basal epi- dermal papillae; anthers 0.5-0.6 mm wide; pollen 24.7—28.2 micra in diameter; style 5-10 mm long, shorter than fertile stamens; mature fruit 1.5-2.5 mm long, 1.1—-1.5 mm wide at the base, 0.8-1 mm wide at the apex, hispid with pointed or truncate, retrorsely barbed hair to 0.4 mm long; seed 1.8—2 mm long, 0.8—-1 mm wide. Chromosome number unknown.
Type. San Lorenzo de Laguna, 75 miles SW of Parras, Coahuila, Mexico, Palmer 833, in May of 1880 (GH, isotype US).
Distribution (fig. 3). Sandy plains, gypsum mesas, and dry trough- like valleys, Coahuila, Mexico.
Specimens examined. MEXICO. COAHUILA: About 30 miles S of Sierra Mojada, Wynd 769; Sierra del Rey, Purpus 4466; W base of Picacho del Fuste, Johnston 8353; 16 miles S of Laguna del Rey, Johnston 7808; near Mohovano, Shreve 8831, between La Vibora and Matrimonia Viejo, Johnston 9348; Rancho Parritas, E margin of the Valle de Acatita, Stewart 2762; Las Delicias, Stewart 2959; W of El Oro, White 2006; S end of Canada Oscuro near Tanque La Luz, John- ston 8487.
2. PETALONYX LINEARIS Greene, Bull. Calif. Acad. 1:188. 1885. Suf- frutescent, almost globose with numerous erect branches, 1.5-10 dm high; current seasons growth 10-38 cm long, to 6 mm wide, grooved, grey-green to dull green, scabrous with short, tapered, truncate, re- trorsely barbed hair, perpendicular to the stem; older stems white to grey; leaves sessile to very short petioled, similar in size along the stems, 1—-4.2 cm long, 0.3-1.5 cm wide, linear to oblanceolate, acute or occasionally obtuse, narrowed basally becoming cuneate to obtuse, entire or rarely remotely dentate, green and somewhat shiny, muricate- hispid with long, pointed, antrorsely barbed hair on the abaxial surface and truncate, retrorsely barbed hair on the adaxial surface, and mixed truncate and pointed hair along the margins; inflorescence a short, capitate, terminal spike, 30-60 flowered, 4-10 cm long, elongated in fruit to 21 cm, the lower fruit fallen; pedicels 1-2 mm long; large bractlet 5-8 mm long, 5-6 mm wide, ovate to round-ovate, acute to obtuse or rarely retuse, cordate to deeply cordate, entire, sinuate or minutely crenulate at the base, hispid on both surfaces and the margins
14 MADRONO [Vol. 19
with long, fine, pointed, antrorsely barbed hair; lateral bractlets 3—3.5 mm long, 2—2.2 mm wide, ovate, acute to obtuse, sub-cordate, lobed, hispid; sepal lobes 1-2 mm long, 0.7—0.8 mm wide, erect, irregularly deciduous; petals white, 2—5.5 mm long; claw 1.4—-3.1 mm long, linear, with no epidermal papellae, few hairs, and not connivent; limb 1.7—2.4 mm long, 1.3—1.4 mm wide, ovate, acute, hispid; stamens barely exserted, 3—7 mm long; filaments without epidermal papillae; anthers 0.3-0.5 mm wide; pollen 24.2—28.2 micra in diameter; style 2.8-6 mm long, nearly equal in length to the stamens, without epidermal papillae; mature fruit 5 nerved, usually 3 ribbed, 2-4 mm long, 1.2—-1.6 mm wide at the base and 0.5—0.6 mm wide at the apex, pubescent with dense, long, pointed, retrorsely barbed, antrorsely appressed, easily deciduous hair, to 0.7 mm long; seed 2.3-2.5 mm long, 0.9-1 mm wide. Chromosome number, M25)
Type. Cedros Island, common in the canyons of the middle of the island, on the E side, Baja California, Mexico, Greene, May 1, 1885 (CAS):
Distribution (fig. 4). Occasional in rocky places in canyons but com- mon in sandy soil below 3,000 ft.; Lower Sonoran Zone, Creosote Bush Scrub, California to southwestern Arizona, southward to Baja California, Sonora and the islands of the Gulf of California.
Specimens examined. Petalonyx linearis is well represented in U.S. herbaria and over 100 collections have been annotated.
3. PETALONYX THURBERI A. Gray, Mem. Am. Acad. 5:319. 1855. Suffrutescent, often broader than tall, to 10 dm high; stems of the current season 12-45 cm long, to 6 mm wide, striate or smooth, grey- green, hispid; older stems white or yellowish; leaves sessile, reducing in size along the branches, 0.4-4.5 cm long, 0.2—1.4 cm wide, variable in shape, deltoid-ovate to lanceolate, acute to acuminate, cuneate to cordate-clasping, entire to few toothed (teeth 0.3—-1.8 mm long and 0-4 on each margin), grey-green, dull to somewhat shiny, hispid to densely hispid with long pointed antrorsely barbed hair; inflorescence a short, dense, naked, terminal, spicate raceme, 10-40 flowered, 1-4 cm long, slightly elongated in fruit; pedicels 0.2-0.7 mm long; large bractlet 3.2-7.5 mm long, 2.1-5.9 mm wide, deltoid-ovate to triangular-ovate, acute to somewhat acuminate, obtuse to sub-cordate, entire or crenate at the base (O—8 teeth per margin), hispid with dense, antrorsely barbed, pointed hair; lateral bractlets 2-3 mm long, 0.8—1.4 mm wide, lanceolate to ovate, unequally crenate to lobed, hispid; sepal lobes 1-2.2 mm long, 0.3-0.5 mm wide, erect, irregularly deciduous; petals cream colored, 2.6—6.5 mm long; claw linear, 1.4—4.1 mm long. 0.3—0.5 mm wide, hispid, without epidermal papillae, the upper one-fifth irregularly connivent; limb 1.2—2.5 mm long, 0.8-1.5 mm wide, ovate, acute, hispid; stamens well exserted, 5—10 mm long, coiled in bud; filaments without epidermal papillae; anthers 0.3-0.5 mm wide; pollen 23.5—29.4 micra in diameter;
1967] DAVIS & THOMPSON: PETALONYX 15
style 3.7-11 mm long; mature fruit 1.6-3.1 mm long, 0.7—1.6 mm wide at the base, and 0.7-0.9 mm wide at the apex, obscurely five-ribbed or smooth, hispid to densely hispid with pointed, antrorsely barbed hair to 0.17 mm long; seed 1.6—2.5 mm long, 0.7—1.3 mm wide. Chromosome number, n = 23.
Type. Valley of the Rio Gila, Arizona, Thurber, June, 1850 (GH).
Distribution (fig. 5). Frequent in dry sandy places, sandy plains, desert washes, dry stream beds, or gravelly places below 4,000 ft., from Inyo Co., California, to Nevada, Arizona, Sonora, and Lower California.
Although this species is the most widely distributed of the five, it is morphologically uniform and only two distinct population groups have been noted and only one of these is given subspecific recognition in this study. Subspecies gi/manii is restricted to Inyo Co., California and will be discussed below. The other morphologically distinct populations are distributed in Imperial Co., California, and Baja California. The leaves on plants from these areas are on the average slightly smaller than P. thurberi ssp. thurberi and are closely appressed to the stems. The remainder of the vegetative and floral characters are within the range of plants of P. thurbert ssp. thurberi from other areas and for this reason we have not accorded subspecific recognition to these populations.
Key TO SUBSPECIES
Leaf and stem pubescence harsh, retrorsely appressed on the stems, stamens longer than 6 mm, widely distributed and common throughout the deserts of California, Arizona, Nevada, Baja California, and Sonora 3a. ssp. thurberi
Leaf and stem pubescence very soft, perpendicular to the stems, satmens less than 6 mm long, restricted to Inyo Co., California. . . . . 3b. ssp. gélmanii
3a, PETALONYX THURBERI ssp. THURBERI. Plants 3-8 dm high; cur- rent seasons growth 22-45 cm long; stems grooved, hispid, with retrorsely appressed hair; leaves 0.6-4.5 cm long, 0.2—-1.4 cm wide, lanceolate to lance-ovate, acute to acuminate, cuneate to truncate, some- what shiny, hispid on both surfaces; petals pale cream to white, 2.6—6.5 mm long; stamens 6.0-10.0 mm long; style 5.0-11.0 mm long. Chromo- some number, n — 23.
Distribution (fig. 5). Frequent in dry sandy places, Lower Sonoran Zone, Creosote Bush Scrub, Covillea and Artemisia belts, Mojave and Colorado deserts, from Inyo Co., California, to Nevada, Arizona, Sonora, and Baja California.
Specimens examined. Petalonyx thurberi ssp. thurberi frequently is collected and is well represented in U.S. herbaria. Over 150 collections have been annotated.
3b. PETALONYX THURBERI ssp. gilmanii (Munz) Davis & Thompson, n. comb. Petalonyx gilmanii Munz, Leafl. West. Bot. 2:69. 1938. Plants up to 10 dm high; current seasons growth 12—21 cm long; stems not grooved, densely hispid with long hair, perpendicular to the stems; leaves 0.4—2 cm long, 0.4-1.3 cm wide, deltoid-ovate, acute to abruptly
16 MADRONO [Vol. 19
acuminate, truncate to cordate-clasping, dull, never shiny, densely hispid on both surfaces; petals pale cream, 2.9-4.1 mm long; stamens 5—7.5 mm long; style 3.7-6 mm long. Chromosome number, n = 23.
Type. Ryan Wash, Death Valley, at 1500 ft. elevation, Inyo Co., California, M. French Gilman 1568, May 20, 1937 (POM 228696; Iso- type frag. CAS).
Distribution (fig. 5). Rare in washes and canyons, restricted to Inyo, California.
Specimens examined. CALIFORNIA. Inyo Co.: along road to Ryan, Death Valley, Davis 170; Cottonwood Canyon, Panamint Mountains, Coville & Gilman 308; 20 miles N of Stovepipe Wells Hotel, Coville & Gilman 156; Furnace Creek Canyon, Funeral Mountains, Coville & Funston 356; 5 miles NW of Saratoga Springs, Coville & Funston 249; Furnace Creek, Death Valley, Parrish 10047; Ubehebe Crater, Death Valley, Kerr in 1940; Ubehebe Crater Wash, Gilman 2681, 3374, and 3377; Slopes of Ubehebe Crater, Gilman 2101; Trona highway, Pana- mint Valley, Twisselman 5843.
In general, ssp. gilmanit has smaller flowers, broader leaves, and longer finer pubescence than ssp. thurberi. In addition, the stem hair is erect in gilmani instead of appressed as in thurberi, and the current seasons growth generally is shorter than in thurbert.
4, PETALONYX NITIDUS Watson, Am. Nat. 7:300. 1873. Petalonyx thurberi var. nitidus (Watson) M. E. Jones, Contr. West. Bot. 12:16. 1908. Suffructicose with many erect branches, 1.5—4.5 dm high; current seasons growth 13-37 cm long, to 8 mm wide, not grooved, grey-green to dull green, scabrous with short, pointed antrorsely barbed hair, mostly antrorsely appressed; older stems straw-colored to dark gray; leaves petiolate, similar in size along the branches, 1.5—-4.5 cm long, 1-3 cm wide, ovate to broadly ovate, acute to shortly acuminate, obtuse to rarely truncate, serrate to coarsely few-toothed (teeth 0.7-3 mm long and 2—8 on each margin), dark green, vernicose, muricate-scabrous on both surfaces with short, pointed, antrorsely barbed hair and on the margins with short, truncate, antrorsely barbed hair; petiole 0.8—5 mm long; inflorescence a terminal bracted raceme, 10-30 flowered, 3—4.5 cm long, not particularly elongated in fruit; pedicels 1-2 mm long; large bractlet 4.8-13.2 mm long, 2.1-7.9 mm wide, lance-ovate, acuminate, truncate, crenate with 3—7 teeth on each margin, scabrous with short, pointed, antrorsely barbed, appressed hair on both surfaces, and short, stout, truncate, retrorsely barbed hair on the margins; lateral bractlet 1-5 mm long, 1—2.4 mm wide, elliptic to ovate, acute to shortly acumi- nate, crenate along the basal margins, hispid; sepal lobes 1.4-3 mm long, 0.5-0.8 mm wide, irregularly deciduous; petals cream colored, 5-11 mm long; claws linear, 3.9-7 mm long, 0.3-0.4 mm wide, hispid along the upper one third, the upper one fourth connivent, the lower portion without epidermal papillae; limbs 1.5-3 mm long, 1.7-1.9 mm
1967] DAVIS & THOMPSON: PETALONYX 17
wide, ovate, acute; stamens well exserted, 7-14 mm long, filaments with scattered epidermal papillae at the base; anthers 0.5-0.7 mm wide; pollen 24—29 micra in diameter; styles 8-15 mm long, usually shorter than the stamens; mature fruit five nerved and five ribbed, 1.3-3.1 mm long, 1-1.5 mm wide at the base, 0.7—0.8 mm wide at the apex, hispid with short, slightly tapered, truncate, retrorsely barbed hairs to 0.08 mm long; seed 2—2.2 mm long, 1.2-1.3 mm wide. Chromosome number, m= -25.
Type. Southern Nevada, Wheeler in 1871 (GH).
Distribution (fig. 5). Common in rocky canyons, washes of open desert, and sandy road banks above 3500 ft. Mojave Desert; Creosote Bush Scrub; Joshua Tree Woodland; and Pinyon-Juniper Woodlands. Inyo and San Bernardino counties of California, to southern Nevada, southwestern Utah, and northwestern Arizona.
Specimens examined. Petalonyx nitidus is well represented in U.S. herbaria and we have annotated over 50 collections.
We have encountered one collection of P. nitidus that falls well out- side of the normal distribution of this species. This collection by Marcus Jones from Yucca, Arizona, is labeled Jones 4453, May 15, 1884. Although Jones’ itinerary indicates that he was collecting around Yucca on May 15, 1884, it is also recorded that number 4483 was collected on Sept. 11, 1884 at El Paso, Texas. The uncertainty concerning the collection is unfortunate since some of the specimens appear to be vegetatively intermediate between P. thurberi, which Jones did collect from Yucca, Arizona (Jones 3936, May 20, 1884) and P. nitidus. The presence of Jones 4453 probably accounts for the inclusion of P. nitidus in the Flora of the Sonoran Desert (Shreve and Wiggins, 1964).
5. PETALONYX PARRYI A. Gray, Proc. Am. Acad. 10:72. 1874. Peta- lonyx nitidus ssp. parryi (A. Gray) Urban & Gilg, Nova Acta Acad. Leop.-Carol. 76:20. 1900. Frutiscose, 8-15 dm high; stems of the cur- rent season usually less than 13 cm long, to 10 mm wide, not grooved, pale green to straw-colored, scabrous with scattered, short pointed, retrorsely barbed hairs, perpendicular or antrorsely appressed; older stems white-grey to dark grey; leaves petiolate, similar in size along the stems, 1.5—4 cm long, 1.2-3 mm wide, oblong-ovate to broadly elliptic, acute to obtuse, obtuse, entire or crenate (teeth 0.3—1.2 mm long and 5-12 on each margin), grey-green to yellow-green, usually not shiny, muricate-scabrous with short, pointed, strongly appressed hairs on both surfaces and short, pointed, strongly curved hairs on the margins; petiole 0.5—2.3 mm long; inflorescence a naked, spikelike, raceme, con- gested apically, up to 65 flowered, 4-8 cm long, elongated somewhat in fruit, the lower fruit fallen; pedicels 1-2.5 mm long; large bractlets 6.3-14 mm long, 2.4-10.1 mm wide, lanceolate to lance-ovate, acumi- nate, obtuse to truncate, crenate to dentate basally with 3-13 teeth per margin, hispid to scabrous with short, pointed, antrorsely barbed, ap-
18 MADRONO [Vol. 19
pressed hair on both surfaces and short, truncate, retrorsely barbed hair on the margins; lateral bractlets 2.5—-2.7 mm long, 1.3-1.6 mm wide, ovate, acuminate, dentate-crenate basally, hispid; sepal lobes 2.5-4 mm long, 0.5—-0.9 mm wide, erect, irregularly deciduous; petals cream colored, 10-15 mm long; claws linear, 6.3—9.8 mm long, 0.3-0.4 mm wide, the upper one-half connivent, no epidermal papillae basally; limb 2.9-5 mm long, 1.7—2.3 mm wide, ovate, acute to obtuse, hispid; stamens well exserted, 11-17 mm long, filaments without epidermal papillae; anthers 0.6—0.8 mm wide; pollen 24—28.1 micra in diameter; style 11-15 mm long, generally shorter than the stamens; mature fruit five nerved and five ribbed, 2-4 mm long, 1.5—-1.6 mm wide at the base, 0.2-0.9 mm wide at the apex, hispid with short, conical, truncate or pointed hairs, antrorsely barbed, to 0.15 mm long; seed 2.5—2.6 mm long, 1-1.2 mm wide. Chromosome number, n = 23.
Type. St. George, southern Utah, C. C. Parry 75, 1874 (GH). Accord- ing to Parry (1875, p. 144) the type collection was taken from a single plant “within a stone’s throw of the great Mormon Temple.”
Distribution (fig. 5). In dry washes and on low slopes, usually on white to grey clay soils and below 3000 ft.; from Coconino and Mohave counties, Arizona, to southern Utah and Nevada.
Specimens examined. Petalonyx parryi is well represented in U.S. herbaria and over 25 collections have been annotated.
Although very similar to P. nitidus in floral morphology, P. parry is vegetatively distinct. For example, P. parryi is woodier, and the leaf margins are crenate rather than dentate as in P. nitidus. The morpho- logical distinctions between the two are quite sharp, and these along with ecological differences support the maintenance of these two as separate species.
Department of Biology, University of Louisville, Kentucky Department of Botanical Sciences, University of California, Los Angeles
LITERATURE CITED
Hamel, J. L. 1938. Etude de la mitose somatique et numération chromosomique chez quelques Loasacées. Revue Cyt. Cytophys. Vég. 3:153.
Parry, C. C. 1875. Botanical observations in southern Utah. II. Am. Nat. 9:144-146.
SHREVE F., and I. L. Wiccrns. 1964. Vegetation and flora of the Sonoran Desert. Vol. 2. Stanford Univ. Press.
TIMBERLAKE, P. H. 1962. A revisional study of the bees of the genus Perdita F. Smith, with special reference to the fauna of the Pacific Coast. Univ. Calif. Publ. Ent. 28:1-124.
Urpan, I., and E. Grrc. 1900. Monographia Loasacearum. Nova Acta Ocad. Leop.- Carol. 76:10-22.
MACNAB CYPRESS IN NORTHERN CALIFORNIA: A GEOGRAPHIC REVIEW
J. R. GRIFFIN AND C. O. STONE
Wolf’s (1948) monograph on the American cypresses is an excellent taxonomic and geographic study. Recent activities of the State Coopera- tive Soil-Vegetation Survey, however, have uncovered some distributional data about cypresses that were not available to Wolf. Much of the infor- mation concerns the little-known northern groves of MacNab cypress, Cupressus macnabiana A. Murr., in Shasta and Tehama counties.
MacNab cypress was probably discovered in Shasta Co. in 1854. The exact locality of the type collection cannot be determined. Published material still does not adequately describe the distribution and ecology of this cypress which is scattered as small groves or as larger clusters of thickets around the Sacramento Valley (fig. 1, table 1). Ironically, the only well publicized and easily accessible stand of MacNab cypress in this region was destroyed recently by the construction of Whiskey- town reservoir.
In Lake Co. and adjacent areas of the Coast Range, MacNab cypress populations geographically overlap those of C. sargenti Jeps. Within these areas C. sargenti often dominates in lower canyon slopes and creek bottoms. Morphological “intermediates” between these two species have been obsrved by several collectors. Wolf (1948), however, discounted suggestions of introgression between C. sargenti and C. macnabiana in the Napa-Lake Co. area. Our observations support this suggestion.
MacNab cypress is geographically isolated from C. bakeri Jeps. in the Sierra Nevada-Cascade and Siskiyou Mountains. Jepson (1909; 1931) and Wolf (1948) have clearly outlined the morphological and ecological differences between these two species.
REVIEW OF SHASTA-TEHAMA CYPRESS REPORTS
Since Sudworth (1908) included C. bakeri in C. macnabiana, we should first consider these C. bakari reports. In northeastern Shasta Co., a large cypress stand occurs near Timbered Crater (Stone, 1965). Al- though discovered in 1898, it was apparently unknown to Sudworth. In 1909 Jepson described this Timbered Crater cypress as C. bakert.
Sudworth (1908) specifically mentioned seven C. macnabiana local- ities in Shasta and Tehama counties. Wolf transferred one of these seven to C. bakeri. Sudworth’s ‘‘west of Burney Creek” grove is the west- ern outpost of the large Burney Springs C. bakeri population. Another locality vaguely cited by Sudworth as “small grove at base of Lassen buttes” was probably based on another ambiguous report of the same Burney Springs population. Sudworth also mentioned a cypress locality at “South Fork of Cow Creek,” nine miles southwest of the Burney
19
20 MADRONO [Vol. 19
) j aed es) Se ——
' -f Ay —) a SHASTA CO. : _ oN? nea ; : 5 : ! \ bf os | - 4, 6 | | OX a Redding | ( ! - ae | ! | F eee Ae Co a a =; = | i Z 9 1 fon 7 | > ; i f | ¢ Xx cm ‘J ar : \ n 1 | ‘ fs = Eee, 2 TEHAMA CO, Of : | \ -4 ji uy rd 4 ; Gerning ee 1 ! -—:—-—— Se es at : 1 = Chico x TL an J MENDOCINO CO. | : OE ee aa) 2 al 2 p49 ( BUTTE co. [+14 cae 2@ one ae f j { ae : f 14 “ 1 a ie Seen ee eM ¢ e Uae ee rs a { y CoulsAy © Si a Neer ata | J a ‘ CO. ate Siar ae 4+& ae i ae a XX Ay ‘ : YUB KXSrass eit H Ukiah | t Williams 2 \ A | palley r “a x, LAKE CO.> a Y {Co} ‘ Ae Se eee ae G 7 a a ‘ 1B x *.>—-— a ~ Wee ae . a ws’ pe ea | \ 5 . ’ N | y ‘sey YOLO Co, PF r ‘ at he | SONOMA CO. re XG a ‘\ Placerville , \ ~ : a APA eee ° Poo ie eae Whe Cee ae 124 » | 2 | a v ! ¢ |AMADOR Lae ° i [aye | CO. | 126 § } Pe eee 2 ee Nee ae | | O 25 ESS eee eee iB Miles |
Fic. 1. Diagrammatic map of Cupressus macnabiana distribution in northern California. Shaded areas represent stands mapped by the State Cooperative Soil- Vegetation Survey. Crosses indicate unmapped stands listed in Tables 1 or 2. Num- bered stands are described in Tables 2 and 3.
Creek grove. Through the efforts of E. L. Little, Jr. we examined many unpublished notes and maps and found that Sudworth’s report was based on a letter dated Jan. 19, 1907, from J. C. La Plant. The letter men- tioned two thrifty trees in the area. We checked this region but found no cypress. The trees that were there were most likely the montane C. bakeri and not the foothill C. macnabiana.
Wolf became skeptical of Sudworth’s notes after he found Juniperus californica Carr. at the ‘“‘Butte Creek” (Battle Creek) locality. There is no cypress now in the section cited (table 2). Sudworth had estab- lished the locality from another report forwarded by La Plant on Feb. 13, 1907. La Plant’s report may have been prompted by a rumor about the Ash Creek cypress grove three miles north of the juniper locality.
1967]
GRIFFIN & STONE: MACNAB CYPRESS 21
TABLE 1. GENERAL LOCALITY AND ASSOCIATED Sort PARENT MATERIAL OF MacnaB Cypress POPULATIONS. SHASTA AND TEHAMA Co. STANDS
Locality
AMADOR CO. E of River Pines
NEVADA CO. NE of Grass Valley SW of Grass Valley
Dominant Soil Parent Material!
ARE DESCRIBED IN TABLES 2 AND 3
Remarks
Ultrabasics (serpentinite)
Ultrabasics (serpentinite) Metavolcanics (greenstone with serpentinite veins)
In T.8N. not T.9N. as listed by Wolf, 1948.
Hall 10182 UC) and Wieslander, 1928.
and basic intrusives (gabbro)
Probably basic intrusives or utrabasics
NE of Nevada City Burk 10 (UC) YUBA CO.
All mapped stands Basic intrusives (gabbro) Not “granitic” as
listed in Wolf, 1948.
E of Forbestown Ultrabasics (serpentinite) Szas (UC) BUTTE CO.
Magalia Ultrabasics (serpentinite) SONOMA CO.
Hooker Canyon Pliocene pyroclastics (tuff) Small groves 4 miles N. and 2 miles SE of main grove reported in 1905. NAPA, LAKE, COLUSA, MENDOCINO CO.S All major populations mapped by SV Survey and most small un-
mapped stands.
Ultrabasics (serpentinite)
{ Zinke 196 (UC) Example
High Valley Ridge Metasediments { of minor nonserpentine | grove in Lake Co. YOLO CO. Cedar Basin Sandstone
1 Calif. Div. Mines. Chico sheet (1962), Santa Rosa sheet (1963), Ukiah sheet (1960), Geologic map of California, San Francisco. All rocks were mapped as Mezo- zoic in age except for Sonoma Co. tuff.
Another area of confusion in Sudworth’s published list is near Paynes Creek, Tehama Co. The ‘‘Payne Creek” report was probably based on the same trees as the “‘Inskip”’ report.
Kildale’s ‘Trinity River” collection (table 2) is confusing because no other Trinity Co. cypress report has come to our attention. Sargent’s (1922) mention of “Trinity County between Shasta and Whiskeytown” obviously referred to the Shasta Co., Whiskeytown grove. Kildale’s specimen label stated that it was collected on the “Trinity River, at Betty May Mine.” Unfortunately, no Betty May mine has ever been recorded in the legal files of Trinity Co. A mine with a similar name did exist in the Clear Creek mining district near Whiskeytown.
22
MADRONO
[Vol. 19
TABLE 2. SUMMARY OF MACNAB CYPRESS STANDS REPORTED IN SHASTA AND TEHAMA COUNTIES
General Area
“Butte Creek”
Paynes Creek
Trinity River
Raglin Ridge (1)
Rainbow Lake (2)
Whiskeytown (3)
Kennett (4)
Montgomery Creek (5)
Round Mountain (6) Lack Creek (7)
Ash Creek (8)
Inskip Butte (9)
Mill Creek Rim (10)
Specific Locality
Status in Wolf (1948)
“Sec. 25, T. 30 N., R.1.W.” Discredited.
(Sudworth, 1908)
“Near Payne post office
and on Payne Creek
Hill... .”’- (Sudworth,
1908)
“..at Betty May Mine,
Trinity County.” Kildale 10276 (DS)
Sec. 20, 21, 22, 27, 28, 29,
T. 25N., R. 7 W.
SW-%, Sec. 31, T 31 N.,R.7W,
See. 16, 21, P32 N;
R. 6 W.
“Along road off U.S. 99, leading to Kennett...” Bacigalupi 2386 (DS)
DSEC.35,00 Oo IN. LOW
R.1W.
Sec. 26, T. 34 N., Reo We
Sec. 36, T. 31 N., R.2 W. Sec. 31, T. 31 N., R.1 W.
Sec. 11, 12, T. 30N., R.1W.
oT. 29 Ne oRo dW? (Sudworth, 1908)
“Sec, 5, T. 27 N., R.3 E.”
Discredited.
This one 1930 collection mentioned.
Not known to Wolf
Not known to Wolf
Described in detail.
This one 1934 collection mentioned.
Not known to Wolf.
Not known to Wolf.
Not known to Wolf.
Not known to Wolf.
Discredited.
Mentioned by Jepson (1931), not discussed by Wolf.
Remarks
Juniperus californica present at this locality.
Same as Inskip Butte stand.
Probably a Whiskeytown collection. Shown on S-V map 31 D-3, 42 A-2. Portions on public lands. Shown on S-V map 31 A-2. Private land. Entire stand destroyed by construction of Whiskeytown Lake. These trees can no longer be found.
Shown on S-V map 22 B-3. Private land. Shown on S-V map 22 B-3 Private land. Shown on S-V map 32 A-1. Private land. Shown on S-V map 33 B-2. Portions on public lands. Shown on S-V map 33 B-3. In Sec. 2223..27, Private land. This stand has not been relocated.
1967] GRIFFIN & STONE: MACNAB CYPRESS 23
We probably have not located the exact point of the Kennett collec- tion (table 2). Owing to the many road relocations and widespread hab- itat disturbance since Shasta Dam was built, it is not surprising that these trees cannot be found. Some cypress might still survive in the brush above Shasta Lake, but no significant population exists in the area today.
The six previously unreported cypress stands, in Shasta and Tehama counties (table 2), have probably been known to a variety of non- botanists for a long time. But no published references to these groves or herbarium specimens from them before the Soil-Vegetation Survey have come to our attention. Sudworth had limited information about at least one of these groves. C. Hart Merriam mentioned MacNab cypress near Round Mountain in a 1908 note.
Population samples from all localities (except Whiskeytown) listed in Table 3 are available at the Pacific Southwest Forest and Range Exper- iment Station, Redding.
GrRovE DESCRIPTIONS
The Shasta-Tehama cypress groves have physiographic and broad climatic similarities. All are above the valley terraces (table 3). Sum- mers are dry and hot, and the abundant fuel creates extreme fire hazards in all of them. Vegetationally, the groves are part of a complex mosaic of chaparral and woodland communities. They grow above the oak savannas and just within the lower fringes of the coniferous forest. Three widely distributed shrubs—Arctostaphylos viscida Parry, Ceanothus cuneatus (Hook.) Nutt., and Eriodictyon californicum (H. & A.) Torr. —are associated with all the groves. Pinus sabiniana Dougl. grows in, or adjacent to, all the groves. Salvia sonomensis Greene carpets the ground around many of the groves. Yet, these MacNab cypress habitats reveal considerable diversity.
Climate. While emphasizing the dry nature of C. macnabiana habitats, Wolf (1948) underestimated their mean annual precipitation. He gen- eralized that most localities had about 20 inches of rainfall, noting Whiskeytown as an exception with 35 inches. Current data raise this Whiskeytown estimate nearer to 50 inches (table 3). The Montgomery Creek grove is the wettest in the Shasta-Tehama area, with 60 inches or more. Other groves to the south in the Sierra Nevada also receive heavy precipitation. The Inskip locality has half as much rainfall as the wetter areas, and it may be as dry as any of the Coast Range local- ities.
No long-term temperature data are available for any of these local- ities. Although their heat regimes appear superficially similar, aspect and elevational differences are great enough so that some temperature variation should be present. Edaphic factors appear more helpful in explaining the distribution of these disjunct populations than climatic patterns.
Soils and Vegetation. Although many of the Sierra Nevada and Coast Range populations are associated with ultrabasic rocks (table 1),
24 MADRONO [Vol. 19 TABLE 3. GEOGRAPHIC COMPARISON OF SHASTA AND TEHAMA COUNTY MAcnaB CyprEsS POPULATIONS Mean Soil Population Elevational Annual Parent Soil (B Horizon) Locality Size Range Precipitation! Material? Characteristics acres feet inches Raglin 1,500 2300-3200 45-50 Mesozoic Shallow, rocky, Ridge (1) utrabasics clay loam, (serpentinized) neutral. Rainbow 22 1700-1900 45-50 Mesozoic Shallow, rocky, Lake (2) ultrabasics clay loam, (serpentinized) neutral. Whiskey- 300 1000-1200 45-50 Devonian Shallow, town (3) Meta-rhyolite clay loam, strongly acid. Montgomery 75 1800-2200 60-65 Eocene non- Moderately deep, Creek (5) marine rocky, clay, sediments strongly acid. Round 23 2500-2600 55-60 Eocene non- Moderately deep, Mountain (6) marine rocky, clay, sediments strongly acid. Lack 350 1400-1550 30-35 Pleistocene Moderately deep, Creek (7) basalt rocky, clay, strongly acid. Ash 200 2800-2900 30-35 Pleistocene Moderately deep, Creek (8) basalt very rocky, silty clay, moderately acid. Inskip 100 1850-2250 25-30 Recent Extremely rocky, Butte (9) basalt silt loam.
1 Calif. Dept. Water Resources. Shasta County Investigation Bulletin 22, Sacra- mento. 1964.
* Calif. Div. Mines. Redding sheet (1962), Westwood sheet (1960), Geologic map of California. San Francisco.
only two of these populations grow on serpentinite (table 3). The widely scattered thickets on Raglin Ridge are on part of an extensive ultrabasic tract resembling the Lake Co. cypress areas. The little Rainbow Lake grove is within a smaller, perhaps less serpentinized ultrabasic mass 30 miles to the north of Raglin Ridge.
The chaparral associated with these two groves has a similar aspect, for the serpentine endemic scrub oak, Quercus durata Jeps., is abundant in both. Shrubs of Rkamnus californica ssp. crassifolia (Jeps.) Wolf occur in both. The Raglin Ridge cypress area appears to have a richer shrub flora, including Ceanothus jepsonii Greene and Garrya congdonit Eastw. The higher elevation cypress groves on Raglin Ridge are inter- spersed with dense Pinus attenuata Lemm. thickets.
The edaphic situation at Whiskeytown was unique in relation to all other MacNab cypress localities. The cypress was concentrated on a bench of meta-rhyolite. Portions of the area had fine textured, mottled
1967 | GRIFFIN & STONE: MACNAB CYPRESS 25
soil horizons that restricted deep rooting. Wolf described the area as having “‘sterile nearly white rocky soil.” McMillan (1956) character- ized it as “‘fine whitish alluvium” from granite. He sampled soils at six different cypress localities, and Whiskeytown had the lowest pH (4.98). The chaparral in the basin was relatively poor in species. The cypress here was also mixed with Pinus attenuata thickets.
The Round Mountain and Montgomery Creek groves are only five miles apart. Both are on part of an extensive formation of loosely con- solidated sandstone and gravel. These soils are mostly covered with woodland at all elevations.
The Lack Creek and Ash Creek groves are also about five miles apart. Both are on reddish soils derived from basalt. Although deep in spots, these soils are liberally mixed with large boulders. Such zonal wood- land soils occur on tens of thousands of acres of volcanic flows to the east of the Sacramento Valley. The chaparral around these two popula- tions has a great diversity of species present. Both have dense clumps of shrubby Quercus garryana Dougl. The Lack Creek community in- cludes such species as Fraxinus dipetala H. & A., Fremontodendron californicum (Todd.) Cov., and Ptelea crenulata Greene which are not present at the other Shasta-Tehama cypress localities. A disjunct Aden- ostoma fasciculatum H. & A. population (Wiselander 338, UC) also occurs at Lack Creek. Although it is the dominant shrub on millions of acres in the hills west of the Sacramento Valley, Adenostoma is rare in the volcanic region to the east of the Valley.
Inskip Butte contrasts with all the other C. macnabiana localities. The “soil” on this small, recently formed volcano consists of silty material in the fractures of the porous basalt. Annual grassland and oak savanna grow on older volcanic materials which surround the butte. Northeast slopes of the butte support a poor pine forest, while the cyp- resses occur on the lower west slopes in depauperate chaparral com- munities. Associated with the cypresses on Inskip Butte are thickets of a disjunct Pinus attenuata population.
Intensive study of aerial photographs and limited field work have proved insufficient to relocate the Mill Creek stand. There is no reason to doubt that the specimen sent to Jepson (1931) was C. macnabiana and that it came from the vicinity of the Mill Creek Canyon in Tehama Co. A small grove may still be there.
DISCUSSION
The relatively little-known stands discussed here do not significantly extend the total distribution of C. macnabiana. The rejection of the Trin- ity River locality even restricts the northwestern corner of the range. But these data do supplement our knowledge of how this species is scat- tered about the Sacramento Valley. The Whiskeytown stand was most inadequate to illustrate the range of edaphic and climatic conditions that this cypress can tolerate in its northern distribution.
26 MADRONO [Vol. 19
The Raglin Ridge and Rainbow Lake stands appear to be a northern extension of the common Coast Range distribution pattern. More inten- sive study of the ultrabasic zone across Glenn Co. may yet reveal some small MacNab cypress groves in the gap between the Colusa and Tehama County populations (fig. 1). One difference in these two northern Coast Range MacNab cypress communities is the absence of any other cypress species. In the Lake Co. and adjacent serpentinite areas, C. sargenti grows in or near many C. macnabiana stands.
The eastern Shasta-Tehama stands may be viewed as an extension of the Yuba and Butte Co. populations. Again, closer study of the most inaccessible brushy canyons might reveal a few isolated trees or small groves that have escaped botanical notice. In contrast to the Coast Range situation, no edaphic trend is apparent along this Sierra Nevada- Cacsade distribution.
From the standpoint of base exchange properties, there might be some similarities between the basic extrusive, basic intrusive, and ultrabasic soils represented here. The Kennett grove which must have been on greenstone also fits into this pattern. Even the Montgomery Creek and Round Mountain sediments were derived largely from basic igneous materials. But if there are any soil chemical similarities, they seem to be obscured by the gross differences in physical properties. The contrast between the reddish clay loam over clay at Lack Creek, which may be more than five feet deep, with the greyish silty loam at Inskip, which may be only a few inches deep, is startling.
Gankin and Major (1964) discussed the distributional patterns of many disjunct and endemic species. They noted that disjunct species frequently exploited non-zonal sites “where competition is decreased by some kind of extraordinary soil parent material or other continuously effective disturbance of climax vegetation development.” Most of these MacNab cypress groves fit conveniently into the context of their discus- sion. The Inskip Butte grove is a good example. The zonal vegetation appears to be excluded from the recent volcanics because of insufficient soil development. In the absence of savanna-woodland species, the cypress can survive on the lava.
Perhaps of greater interest are the Lack Creek and Ash Creek popula- tions, that seem to depart from this general pattern. They grow on widely distributed geologic materials with normal degrees of soil develop- ment for this area. Although these cypresses compete with zonal chapar- ral species on zonal soils, they seem to be in no imminent danger of losing out. Edaphic factors are not obviously involved with the particular location of these cypress groves. Other environmental factors that may contribute to the competitive advantage of cypress in these localities are not apparent. Since it is difficult to explain the presence of the cypresses ecologically, they must grow on these zonal soils because of historical reasons. The coincidental presence of the isolated Adenostoma
1967] HOLMGREN: PRIMULA at
colony at Lack Creek suggests that some peculiar set of circumstances has influenced this vegetation in the past.
Three of the Shasta-Tehama cypress groves are closely associated with Pinus attenuata populations. Two other groves are not far from P. attenuata areas. Several ecological similarities between these two different fire-adapted conifers are apparent. The historical events that have contributed to the widely scattered distribution of P. attenuata probably have also influenced the distribution of MacNab cypress. The pine, however, has a broader range of ecological tolerances. Its range closely approximates the combined range of the three cypresses men- tioned here: C. macnabiana, C. sargentu, and C. bakeri.
Pacific Southwest Forest and Range Experiment Station, U.S. Department of Agriculture, Redding, California
California Division of Forestry, Sonora
LITERATURE CITED
GANKIN, R. and J. Major. 1964. Arctostaphylos myrtifolia, its biology and relation- ship to the problem of endemism. Ecology 45:792-808.
Jepson ,W.L. 1909. A flora of California. Vol. 1. Associated Students Store, Berkeley.
Jepson, W. L. 1931. Open letter on Cupressus macnabiana. Madronfio 2:31.
McMirtan, C. 1956. The edaphic restriction of Cupressus and Pinus in the Coast Ranges of central California. Ecological Monographs 26:177-212.
SARGENT, C. S. 1922. Manual of the trees of North America... Houghton Mifflin Co., Boston and New York.
STONE, C. O. 1965. Modoc cypress, Cupressus bakerii, does occur in Modoc County. Aliso 6:77-87.
SupworTH, G. B. 1908. Forest trees of the Pacific slope. U.S. Dept. Agr., Forest Service.
WIESLANDER, A. E. 1928. (Note on Cupressus macnabiana). Madrono 1:186.
Wotr, C. B. 1948. Taxonomic and distributional studies of the New World cypresses. Aliso 1:1-250.
A NEW SPECIES OF PRIMROSE FROM NEVADA
NorEL H. HOLMGREN
Primula nevadensis N. Holmgren, sp. nov. Folia erecta, scapos plerumque superantes, oblanceolata vel lineari-oblanceolata, (2.5)—5— 10-(12) cm longa, 0.6-1-(1.5) cm lata, distaliter grosse dentata vel enterdum subtiliter dentata vel etiam integra, ad apicem obtusa vel
28 MADRONO [Vol. 19
acuta, raro rotundata; scapa 3.5-9-(11) cm alta, pauciflora, bracteis involucri (3)—4—8—(11) mm longis; flores heterostyli, calyce 6-10 mm longo, tubo lobos superanti, corolla lobis violaceis patentibus emargi- natis, tubo in flore brachystylo (7)—8—-10-(11) mm longo, limbo (4)— 6—10-—(11) mm lato, tubo in flore dolichostylo (6)—7—9-(10) mm longo, limbo (5)—7—11—(12) mm lato.
Herbage, minutely glandular puberulent; leaves erect, usually sur- passing the scapes, oblanceolate to linear oblanceolate, gradually taper- ing into a winged petiole, (2.5)—5—10—-(12) cm long, 0.6-1—(1.5) cm wide, coarsely toothed beyond the broadest point to sometimes entire, obtuse or acute, rarely rounded at apex; scapes 3.5-9-(11) cm high, bearing an umbel of (1)—2—3-—(8) flowers; involucral bracts as many as the flowers and opposite the pedicels, the largest one (3)—4—8—(11) mm long, lanceolate, the smallest rudimentary and up to 3 mm long, triangu- lar to lanceolate; pedicels 5—15—(23) mm long; calyx pale green and densely farinose, especially in a rhomboid patch below each cleft, to purplish and scarcely farinose, tube (3.5)—4—7—(8) mm long, the teeth 2—4 mm long, triangular; corolla purple in bud, lobes violet at anthesis with a dark purple ring around the throat, throat yellow to yellow- orange inside, yellow or brown with a violet tinge outside, tube yellow to white; flowers heterostylous, the measurements of the tube and limb varying accordingly, the lobes alike in all flowers; in thrum flowers (anthers above the stigma), tube (7)—8-10-(11) mm long, limb (4)- 6—-10-(11) mm wide, in pin flowers (anthers below the stigma), tube (6)-—7-9-(10) mm long, limb (5)—7-11—(12) mm wide; corolla lobes (4)-5—9-(10) mm long and (3)—5—8-(10) mm broad, emarginate to obcordate; anthers yellow to yellow orange, about 2 mm long; pollen grains tetrahedral with convex walls, measurements in pin flowers about 15.6 and in thrum flowers about 21.84; capsules 5-6 mm long, 4 mm thick, elliptical, within a persistent and slightly accrescent calvx.
Type. Holmgren, Reveal & LaFrance 2216, south slope of Mt. Wash- ington, Snake Range, White Pine Co., Nevada, at about 11,500 feet elevation, July 20, 1965 (NY-holotype, 34 isotypes distributed).
Additional specimens. Nevada. Nye Co.: Troy Preak, Grant Range, Worth & Priest 601 (NY); Holmgren, Reveal & La France 2213 (34 sheets distributed). White Pine Co.: Mt. Washington, Snake Range, Holmgren & Reveal 1653 (14 sheets distributed); Snake Range, Lin- coln Peak, Holmgren, Reveal, & LaFrance 2219 (35 sheets distributed).
Primula nevadensis is known from only the Grant Range and the Snake Range of east central Nevada, where it seems to be restricted to limestone outcrops at elevations above 11,000 feet. Similar habitats exist in the two intervening ranges, the Egan and Schell Creek ranges, and also in two other nearby ranges, the White Pine Range just north of the Grant Range and the Mt. Moriah region of the northern Snake Range. With the assistance of James L. Reveal and Charles LaFrance, we explored these areas and failed to find it.
1967 | HOLMGREN: PRIMULA 29
In its known localities P. nevadensis is very common in crevices of limestone rock and on open gravelly slopes and ridges with scattered Pinus aristata Engelm., Ribes montigenum McClat., Eriogonum holm- grenit Reveal, Phlox hoodiw Richards., Aquilegia caerulea James, Cas- tilleja nana Eastw. and krumholz of Picea engelmannu Engelm.
The nearest relatives of P. nevadensis are P. cusickiana Gray, P. maguire: L. Williams and P. angustifolia Torr. These four species are the New World representatives of the section Nivales Pax. Primula nevadensis is probably more closely allied to P. maguieret and P. cu- sickiana than to P. angustifolia. The distinguishing characteristics are shown in the following key:
Inflorescence with 1, rarely 2 or 3 flowers; bracts 1.5-3-(4) mm long; alpine areas
of the eolicrn Rocky Mts. . . . ee . P. angustifolia Inflorescence usually with more than 1 flower; Thee etl 3-9 mm long.
Corolla tube 1.5—2 times as long as the calyx; leaves (7)—10-25 mm wide; calci-
phile; known only from Logan Canyon of the Bear River Range in
northern Utah, at lower elevations. . . : . . P.maguirei
Corolla tube 1-1.5 times as long as the calyx; leaves 4— 10- (15) mm wide.
Scape longer than the spreading leaves; leaves mostly 2.5-5 cm long; foot- hills and montaine slopes in central Idaho and _ northeastern
Oregon, below timberline. . . . . . . . P. cusickiana Scape usually overtopped by the erect leaves; eaves mostly s- 10 cm long; calciphile; alpine areas of east central Nevada. . P. nevadensis
Our discovery of this plant was not entirely by accident. Both James L. Reveal and I had heard independently of an undescribed primrose from eastern Nevada. So while collecting in the Snake Range in the summer of 1964 we were both consciously looking for it, but after a week of collecting on the quartzite and granite slopes of Wheeler Peak and upper Snake Creek drainage we had given it up as not being a Snake Range inhabitant. Because of this it came as a real thrill when we stumbled across it a few days later on the limestone slopes of Mt. Washington only a couple of miles south of Snake Creek.
That winter in the New York Botanical Garden herbarium I dis- covered a C. R. Worth & Amel Priest collection in the unidentified primroses. Worth responded to my inquiry with a letter giving me the history of its discovery in the Grant Range. In the summer of 1945 Rogers McVaugh showed him a specimen he had collected on Troy Peak, which Worth recognized as being distinct from anything he knew. He collected it himself in 1947 and again in 1958. He sent a specimen to William Wright Smith at Edinburgh who did not want to commit himself without more material to study from. To my knowledge these are the only collections of it prior to my own.
I am very grateful to Arthur Cronquist for the opportunity to do field work supported by his NSF grant and for his assistance in pre- paring this note, C. R. Worth for his help, Arthur H. Holmgren for loaning materials, and James L. Reveal for his suggestions and assist- ance in the field.
New York Botanical Garden, Bronx
30 MADRONO [Vol. 19
REVIEWS
Biographies of Nevada Botanists, 1844-1963. By OLGA REIFSCHNEIDER. 165 pp. University of Nevada Press, Reno. 1964. $4.95.
A little more than a hundred years ago William Henry Brewer wrote from Virginia City that “Nevada Territory embraces over eighty thousand square miles, but is nearly all desert. It has just been made a state, but I see no elements here to make a state. It has mines of marvelous richness but it has nothing else . . . The climate is bad, water bad, land a desert, and the population floating.” Nevada, the “land between,” the well-nigh country, was more often sampled by the naturalist on the road than as a destination in itself. Nevada’s explorers seldom lingered. Exhausted, short of food and funds, wearying of the desert, they were often desper- ate to reach their Eldorado. For that reason they sometimes passed by some remark- able plants such as the composite Hecastocleis. On the other hand that Fremont should stumble upon Arctomecon, a papaveraceous genus of restricted range, is a wonder of botanical history. Nevada specimens were often fragmentary and it was not until Sereno Watson threw his energies into the King Expedition that the content of the Nevada flora out of the conestoga tracks came to be known, and even then specimens were sparse by modern standards.
After eight pages of the “physical background” of Nevada and a general account of the history of botanical exploration in the state there begins a chronological album of forty-seven portraits usually with a page or two of biographical matter opposite. About half of the portraits are heretofore unpublished and comprise a notable biohistorical record. A smaller cut would usually have improved the pub- lished photograph. Contemporaries will be the historic figures of tomorrow and it is important to record information and portraits of them, a successful facet of the author’s book. The source of the portrait, the photographer’s name, and the date, are more to be valued than the credit line. The bibliography as it stands is a mixture of technical papers on Nevada botany written by persons included in the book, and biographical accounts of the authors; these two aims should have been divorced with attention given primarily to biographical references. For periodicals pagination as well as volume numbers should have been cited. Separate maps for the collectors, or a map for two or more collectors where their areas of activity will not be easily confused, in the manner of Geiser’s Naturalists of the Frontier, would have been preferable to the “Botanical collection areas” (p. 17). There is need for a complete index of persons. All of these comments are directed toward the preparation of the second edition which we hope will be forthcoming.
Fred Hebard Hillman introduced his Early Flora of the Truckee Valley (1895) by saying it was “issued as a preliminary step toward a future and more complete Flora of Nevada.” Comparably Mrs. Reifschneider’s album of botanical explorers may be described as a pioneer botanical history of Nevada. Corrigenda or addenda are arranged in chronological order of the year of the collector’s visit to Nevada:
1841. William Gambel (1821-1849), Nuttall’s protegé, journeyed across the Virgin River country but doubtfully made any plant collections there. A few of Gambel’s specimens are encountered in the British Museum (Natural History) being originally a part of Nuttall’s herbarium.
1846. Joseph Burke (fl. 1841-1853) may have collected plants along his route from Fort Hall to Willamette Valley. Historian Dale Morgan of the Bancroft Library will presently publish additional information on Burke.
1854. James Aitken Snyder (d. 1900) assisted Lt. E. G. Beckwith in the explora- tion of the western Great Basin. Lt. Beckwith assumed command of Gunnison’s Expedition after the massacre of Capt. Gunnison in October, 1853. Beckwith listed Snyder’s collections in the second volume of the Pacific Railway Reports. These Snyder specimens have been cited frequently as Beckwith’s (for example, King Report, pp. 36 and 109). Susan G. Stokes, Genus Eriogonum, p. 43, 1936, credits the type of Eriogonum nutans T.&G. as being Beckwith’s specimen though it was
1967 ] REVIEWS ot
more accurately Snyder’s. Snyder’s field localities are few: Agate Pass of the Quartz Mountains (Pac. R.R. Rep. 2:121) and “Nevada: canyon at the eastern base of the Sierra Nevada,” the type locality for Eriogonum nutans. Snyder collected Dode- catheon integrifolium and the type of Viola beckwithii “in a canyon between Great Salt Lake and the Sierra Nevada”! but Viola Brainerd Baird, in Wild Violets of North America, p. 212, 1942, fixes the locality as “Diamond Mt., Eureka Co., Nevada, 1854, Snyder.”’ The well known botanical roster in volume two of the Botany of California (1880) mentions Dr. J. A. Snyder but J. H. Barnhart doubts the appellation “Dr.”
1885. Ezechiel Jules Remy (1826-1893), French natural history collector, trav- elled from San Francisco to Salt Lake City in the company of Julius Lucius Brenchley, ornithological collector and author, and recorded his impressions of Nevada in chapters one and two of his Journey to Great-Salt-Lake-City (London, 1861). Remy took a particular interest in the genus Eriogonum and his specimens, never numerous, are in the Natural History Museum, Paris.
186_? Hiram G. Bloomer (1821-1874) is frequently mentioned in the King Report (pp. 27, 140, 153, 166, etc.). His collections made about Mr. Davidson, Steamboat Springs, Washoe Valley, Cedar Hill, and Virginia City, and preserved in the California Academy were mostly lost in the San Francisco fire of 1906.
186__? C. Herbert Dorr collected the type of Audibertia dorrit named by Albert Kellogg in 1863 and presumably taken near Virginia City, but the specimen is lost. The King Report (p. 44) refers to Dorr’s collection of Spraguea paniculata from the western base of Mt. Davidson. Was he related to Sullivan Dorr (1778-1858) of Boston and Providence? Papers of the Dorr family are preserved at the Essex Institute, Salem, Mass.
1862. Charles Lewis Anderson. W. H. Brewer and C. L. Anderson are mistakenly cited in Mrs. Reifschneider’s bibliography as the authors of Asa Gray’s paper on Anderson’s and Brewer’s collections. Interesting biographical facts on Dr. Anderson will be found in “The History of Botanical Collecting in the Santa Cruz Mountains of central California” by J. H. Thomas (Contr. Dudley Herb. 5:147-148. 1961).
1863. William Henry Brewer (1828-1910) collected on the Truckee River, at Aurora, and in the Esmeralda district, and in 1864 about Virginia City. Farquhar published an interesting letter from this period of Brewer’s life in Up and Down California, pp. 551-559. 1930.
1863-64. John Allen Veatch (1808-1870), mining engineer, associate of Dr. Albert Kellogg, and active in the early years of the California Academy, was a resi- dent of Virginia City in 1863-64 where he was evidently engaged in medical prac- tice (as well as mining activities) while his son Andrew Allen Veatch was super- intendent of the reduction works of the central mill. The King Report (p. 133) cites a Veatch collection from the Washoe Mts.
1865. John Torrey (1796-1873) collected at Carson City, Washoe Lake (his no. 123), and about Steamboat Springs, Washoe Valley (no. 372), as cited in the King Report. This visit of 1865 was a stopover on a confidential mission to Cali- fornia as an assayer for the Treasury Department.
1866. John Gill Lemmon (1832-1908) is said (p. 39) to have sent 50 plant speci- ments to Henry N. Bolander, ‘“‘the only botanist on the Pacific Coast at that time.” Lemmon was himself the source of that erroneous statement, which certainly dis- regarded Dr. Kellogg at the Academy! Here we encounter the gentle rivalry that existed between Kellogg and Asa Gray for the description of novelties that were being discovered every year, and Lemmon’s allegiance to Gray. The history of the portrait of Lemmon, whose middle name was Gill not Gibbs, is interestingly told in Madrono 5:77. 1939, where it was first published.
1867. William More Gabb (1839-1878), paleontologist, geologist, assistant to J. D. Whitney on the California State Geological Survey, collected insects as well as plants. Gabb is mentioned in the King Report as having collected on the “sage plains of western Nevada” (p. 117), south of Walker’s Lake (p. 118), and along the
32 MADRONO [Vol. 19
state boundary. For references to Gabb and others in this list see my roster in the Century of Progress in the Natural Sciences, 1853-1953, p. 49. 1955.
1867. William Whiman Bailey (1843-1914), son of Jacob Whitman Bailey, was the first botanist of the King Expedition; he was evidently not a Doctor of Medi- cine as cited by Reifschneider (p. 47). “Entering Nevada from California,” wrote Sereno Watson in the Botany of the Fortieth Parallel, xxii, 1871, “collection was commenced on the 16th of July, 1867, in the valley of the Truckee River by Mr. W. W. Bailey, botanist of the expedition, and was continued through the season but confined wholly to the district between the Virginia and the Pah-Ute Moun- tains. The summer was already so far advanced that the earlier vegetation had dis- appeared and little was left to reward the collector. Some gleanings were however mace, especially in the West Humboldt Mountains [type of Jvesia baileyi was taken in Wright’s Canyon, 7,000 ft., in Sept., no. 346] and in the bottoms and sinks of the Truckee, Carson, and Humboldt Rivers.” Bailey’s health continued to decline dur- ing the winter of 1867 and Watson succeeded him in the spring of 1868.
1868-69. Richard Harper Stretch (1837-1923), entomologist, is mentioned sev- eral times in the King Report (pp. 13, 63, 98, 177, 216, etc.) as having botanized at Carson City and Washoe Valley. He spent the winter of 1868-69 studying the Comstock Lode for the U.S. Geological Survey. Stretch named the Nevada buck- moth (Hemileuca nevadensis ), moths being his particular interest.
1877. When Joseph Dalton Hooker toured the West with Asa Gray by rail they “left the direct route at Reno,” wrote Gray to Engelmann, “went to Carson City, with detour to Virginia City, —queer place; first got hold of Pinus monophylla, but there no fruit. Hired conveyance to take us from Carson right across the Sierra Nevada via Silver Mt. to Calaveras Big Trees.” (J. L. Gray, Letters of Asa Gray, 2:674. 1893).
1878. Charles Frederick Sonne. Portrait published in Madrono 2:115. 1934.
1880. William Hillman Shockley (1855-1925), mining engineer, who as partner in the firm of Zabriskie and Shockley, lived in Candelaria, Nevada, from 1880 to 1893, botanized ‘‘on the side” in Esmeralda County and the White Mts. He took the type of Lupinus shockleyi at Soda Springs and discovered two composites, Acampto- pappus shockleyi Gray, and the endemic monotypic mutisiaceous genus Hecastoclets Gray. In 1878 Shockley was collecting ferns for D. C. Eaton in the limestone sinks of Florida. Jepson published a biographical sketch and portrait in Madrono (2:26-28. 1931).
1890. Vernon (Orlando) Bailey (1864-1942), biologist, started his career with the Death Valley Expedition. On May Ist, 1890, Bailey collected two new species, Abronia orbiculata Stand]. and Arctomecon merriami Cov. between Cotttonwood Springs and Las Vegas Ranch.
1898. Carl Albert Purpus (1853-1941), one of a triumvirate of Mexican botan- ical explorers—Palmer, Pringle, and Purpus—, crossed the Charleston Mountains from Pahrump Ranch to Indian Springs, visiting Clark Canyon on the way, and collecting the types of Arabis pendulina Greene and Frasera induta Tidestr.
1927. Charles Leo Htchcock (1902- ) botanized in the Charleston Moun- tains where he took the type of Lesquerella hitchcockii Munz and Erigeron uncialis var. conjugens Blake. He returned to botanize in the Charleston Mountains with George Jones Goodman (1904— ) in July, 1930.—JosrpH Ewan, Tulane Uni- versity, New Orleans.
Botany
MADRONO
VOLUME 19, NUMBER 2 APRIL, 1967
Contents
NATURAL HYBRIDIZATION BETWEEN ENCELIA AND GERAEA (COMPOSITAE) AND SOME RELATED EXPERI-
MENTAL INVESTIGATIONS, Donald W. Kyhos 33 New REeEcorpDs oF MyXoMyYceETES From CAtirornia I, Donald T, Kowalski 43 Cypress RUSTS IN CALIFORNIA AND BAJA CALIFORNIA, Roger S. Peterson 47 Two New ERIOGONUMS FROM BAJA CALIFORNIA, Mexico, James L. Reveal and Craig A. Hanson 55
Reviews: R. K. Brummitt, Index to European Tax- onomic Literature for 1965 (Wallace R. Ernst) ; Wil- liam T. Stearn, Botanical Latin (Rimo Bacigalup!) ;
H. L. Blomquist, The Grasses of North Carolina and Frank W. Gould and Thadis W. Box, Grasses of the Texas Coastal Bend (Dennis Anderson) ; George J. Hollenberg and Isabella A. Abbott, Supplement to Smith’s Marine Algae of the Monterey Peninsula (John A. West) ; George Neville Jones, An Annotated Bibliography of Mexican Ferns (Duncan M. Porter) 58
NoTES AND NEws: ELwoop WENDELL MOLSEED 46 THE GENUS KIcKxIA IN OREGON, La Rea J. Dennis ; CALAMAGROSTIS RUBESCENS ON SANTA Cruz ISLAND, CALIFORNIA, Peter H. Raven; Two NEw ApDITIONS TO THE FiLora oF Montana, Dennis W. Woodland; FtorA NortH AMERICA, Stanwyn G. Shetler; HETERANTHERA LIMOSA IN CALIFORNIA, John M. Tucker and June McCaskill 62
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. Subscription price $6.00 per year ($4.00 for students). 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 un- delivered copies should be sent to the Corresponding Secretary, California Botanical Society, Department of Botany, University of California, Berkeley, California 94720.
BOARD OF EDITORS
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LyMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. Ernst, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California Mitprep E. Matuias, University of California, Los Angeles RoBERT ORNDUvuF?, University of California, Berkeley Marion OwnBeEy, Washington State University, Pullman REED C. Rotiins, Harvard University, Cambridge, Massachusetts Tra L. Wiccrns, Stanford University, Stanford, California
Editor—JoHn H. THomas Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Carl W. Sharsmith, Department of Biological Sciences, San Jose State College, San Jose. First Vice-President: Robert Ornduff, De- partment of Botany, University of California, Berkeley. Second Vice- President: Robert F. Thorne, Rancho Santa Ana Botanic Garden, Clare- mont. Recording Secretary: Mary L. Bowerman, Department of Botany, University of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Department of Botany, University of California, Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State College.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Malcolm A. Nobs, Car- negie Institution of Washington, Stanford; the Editor of Madrono; and three elected Council Members: Annetta Carter, Department of Botany, University of California, Berkeley; Virginia M. Page, Department of Biological Sciences, Stanford University; and Herbert L. Mason, De- partment of Botany, University of California, Berkeley.
NATURAL HYBRIDIZATION BETWEEN ENCELIA AND GERAEA (COMPOSITAE) AND SOME RELATED EXPERIMENTAL INVESTIGATIONS
DONALD W. KyHOos
During the spring of 1965, while collecting in the California and Arizona deserts, I was surprised to encounter what appeared to be natural hybrids between an annual, Geraea canescens Torr. & Gray, and a shrub, Encelia farinosa Gray. Both of these species are excep- tionally common over large areas and often create some of the most spectacular floral displays on the lower deserts of California, Arizona, and Mexico. Despite their great numbers and frequent sympatry, hybrids between these two rather different taxa have not heretofore been reported. It would appear, however, that such hybrids are not rare, inasmuch as eleven were found within a two-week period, in four well separated areas of the Colorado River valley (table 1).
It would be of unusual interest to learn that an “‘intergeneric” hybrid could be experimentally produced between a woody shrub and an herbaceous annual, as has been done in Parthenium by Rollins (1946). The potential significance of such a hybrid, however, is much greater when it occurs naturally, as in the case of Encelia and Geraea, since if such an unusual hybrid has any fertility at all, it may serve to exchange genetic material between taxa that ordinarily would be assumed to have taken independent courses of evolution. With this possibility in mind, an investigation was undertaken to determine the capacity for gene exchange in these hybrids, as well as examine other aspects of their hybridity.
MATERIALS AND METHODS
All chromosome materials were fixed in a solution of one part glacial acetic acid, three parts absolute alcohol and the cytological investiga- tions were done by the conventional aceto-carmine squash technique. For the paper chromatography studies one gram of leaves from a given plant were placed in 10 ml of one percent HCL in methyl alcohol. The leaves were then thoroughly ground up and allowed to stand in the dark for 12 hours before 25 drops of this solution were applied to #1 Whatman filter paper. After a thorough drying the filter paper was placed in a chromatography chamber where the first dimension was developed in a solution of one part acetic acid, one part water, and three parts tertiary butyl alcohol, for 22 hours. Then the filter paper was thoroughly dried and the second dimension development in the cham- ber was completed in four hours, in a solution of 15 percent glacial acetic acid in water. After drying, the filter paper was examined in
Maprono, VoL. 19, No. 2, pp. 33-64. May 16, 1967. 33
34 MADRONO [Vol. 19
TABLE 1. Hysrip LOCALITIES
65-130 Along road to Earp, Calif., 2.8 mi E of Vidal Junction, San Bernardino Co.
65-158 Dome Valley, Ariz., 5.4 mi N of U.S. Hwy 80, Yuma Co.
65-159 same as above.
65-221 Near road to Lake Havasu, Calif., 18.5 mi E of U.S. Hwy 95, San Bernar- dino Co.
65-222 same as above.
65-225 Near road to Lake Havasu, Calif., 18.2 mi E of U.S. Hwy 95, San Ber- nardino Co.
65-226 same as above.
65-229 Near U.S. Hwy 95 2.6 mi S of San Bernardino Co. line, Riverside Co.
65-230 same as above.
65-231 same as above.
65-232 same as above.
sunlight, in UV light, in UV light with ammonia fumes, and finally after spraying with a 2% ferric chloride solution.
In all experimental crosses discussed in this paper, each plant neces- sarily served as both a maternal and paternal parent. Because E. fari- nosa, G. canescens, and their hybrid are all strongly self-incompatible, no emasculation was necessary and therefore each experimental cross was done by simply rubbing flowering heads gently together. This action exchanges pollen between capitula of different plants, thus simultane- ously effecting reciprocal pollination.
Voucher specimens are deposited in the University of California, Davis, herbarium.
NATURE OF THE SPECIES AND THEIR HYBRIDS
Habitats of the hybrids. Nine of the eleven hybrids grew in obviously disturbed sites at road margins; a frequently observed phenomenon in plants. The remaining two hybrids from Dome Valley, Arizona are of greater interest, since they occurred on small hills that bore no signs of disturbance. The occurrence of these plants in undisturbed sites, suggested that such hybrids might have existed in sufficient numbers to permit the exchange of genes between Geraea and Encelia, even prior to man’s disturbing influence. Such hybrids might also be expected to survive in naturally disturbed areas, although no examples of this kind were observed in the course of this investigation.
Characteristics of the parent species and hybrids. Encelia farinosa and G. canescens are very easily distinguished. The former is a grayish, rounded shrub that is intricately branched and moderately woody. In the more open desert it usually attains a height of two to three feet, but in more protected canyons and washes it may exceed five feet. Geraea canescens, in contrast, is a distinctly green, often sparsely branched ,erect annual, which usually attains a height of only one to two feet. A closer inspection reveals a number of additional features, both vegetative and reproductive, which further differentiate these spe-
1967] KYHOS: ENCELIA AND GERAEA 35
A Cae | Cas
XN 2 3 b be Meat wok SSI NOP
D F M 4 [ f t Ell A Co (ig Ge \ J WV G HI I) eK L P Q R
Fic. 1. Various organs and morphological details of Geraea canescens, Encelia farinosa, and their F, hybrid. In each series of three organs or structures, G. cane- scens is positioned at the left, the F, hybrid in the center, and E. farinosa is on the right: A-C, view of the upper side of leaves, X 34; D-F, pubescence on the under leaf surface, X 12; G-I, involucral bracts, X 144; J-L receptacular bracts, x 1%; M-O, achenes, x 2; P-R, distal portion of the peduncle, x 2.
cis (fig. 1). Although the hybrids between E. farinosa and G. canescens exhibit some gigas features, their morphology is for the most part predictably intermediate, with two notable exceptions. One of the ex- ceptional plants is hybrid 65-130, which has a leaf pubescence remark- ably similar to Encelia farinosa, whereas all of the other hybrids are intermediate for this character. Since all other characteristics of 65-130
36 MADRONO [Vol. 19
are those expected of an F, hybrid, it appears most probable that its Encelia-like leaf pubescence has not resulted from genetic recombina- tion beyond the F, level, but instead represents part of the variation that can occur in this particular F,; combination. The other exception to intermediacy involves disk floret color. In this instance two of the hybrids, 65-130 and 65-229, have brownish-purple disks and another, 65-231, has brownish anthers with yellow disk corollas. All of the other hybrids have yellow florets throughout their flowering heads. The dark color of the disk florets in these hybrids was undoubtedly inherited from E. farinosa, because G. canescens uniformly has yellow florets, whereas some individuals of FE. farinosa have brownish-purple and others have yellow disks in the areas where these hybrids occurred. Dark colored disks in Encelia are inherited in a dominant fashion, with a relatively simple genetic basis (Kyhos, unpublished). It seems likely that disk color is inherited in the same manner in the Encelia X Geraea F, hybrid. Thus the presence of brownish-purple and yellow disks in different hybrid individuals cannot in itself be considered evidence of hybridization past the F, stage.
When discovered, the eleven hybrids were of widely different ages, ranging from very small seedlings to vigorous plants, up to three feet in height and in full flower. From the three largest hybrids, 65-130, 65-158, and 65-159, cuttings were made which slowly took root and developed into robust plants. The remaining hybrids were small enough to be removed in their entirety and brought successfully into cultivation. Under the conditions of cultivation these hybrids have maintained a perennial growth habit. It is not known whether they can exist as per- ennials in nature, although there is a suggestion that this is possible. One of the hybrids (65-130) had earlier grown to a height of about three feet, but when discovered it had already died back to its base. A renewed growth and flowering had occurred from this basal portion, sug- gesting that this individual had been established for more than one season.
Chromatographic studies provided additional evidence that these hybrid individuals are of the F; generation. By means of paper chro- matography over 30 chemical constituents, i.e. spots, were identified from each of the parent species and from among these, four occurred exclusively in EZ. farinosa and another five were found solely in G. canescens. In each of the putative hybrids a complementation of these spots invariably occurred, providing strong evidence for the F, nature of these plants (fig. 2).
Meiosis in the parent species and hybrids. Both E. farinosa and G. canescens consistently form 18 pairs of chromosomes. Pairing of the chromosomes is readily observed in these species and no meiotic irregu- larities have been observed in the more than ten individuals examined of each of these species from widely scattered areas of California, Ari- zona, and Mexico.
1967] KYHOS: ENCELIA AND GERAEA 37
@ +
Fic. 2. Diagrammatic representation of a paper chromatogram of the Geraea canescens X Encelia farinosa F, hybrid. The dark spots are those specific to G. canescens and the unshaded spots are those peculiar to E. farinosa. In the putative F, hybrids all of these spots are complimentarily present as shown in this illustra- tion. The cross in the upper right hand corner of the chromatogram is the origin.
In sharp contrast, the pachytene and subsequent stages of meiosis in the putative hybrids consistently reveal a general lack of chromosome pairing. Moreover, when pairs were formed, they were only very loosely held together, as opposed to the very definite bivalent connections in the parent species. The few chromosomes pairs that occur appear to result from very localized synapsis. A considerable variation exists in the metaphase I-anaphase I chromosome behavior of the hybrids. These plants displayed two distinct types of meiotic behavior, which varied greatly in frequency from one floret to another, even in the same flower- ing head. In one case only one to three chromosome pairs are usually formed, while the numerous univalents seem to segregate to the ana- phase poles at random. In the other case the chromosomes fail to pair at all, and at what would normally be meiotic metaphase, all of the chromosomes divide equationally to produce two equivalent daughter nuclei, each with 36 chromosomes. The resulting telophase I nuclei are subsequently separated by cytokinesis; a phenomenon which normally does not take place in members of the Compositae until after the second meiotic division. There is no evidence that a second meiotic division
38 MADRONO [Vol. 19
occurs in these cells. Each of the two cells continue to develop inside the original PMC wall where normally four cells would be found. It is apparent that not all of these cells develop into mature pollen grains, as indicated by the generally low frequency of filled, presumably functional pollen grains (table 2). In the PMCs where the chromosomes appear to segregate at random to the anaphase I poles, cytokinesis occurs in a normal fashion, i.e. after the second meiotic division. The great majority of these cells possessed obviously unbalanced chromosome numbers and as would be expected seldom produce mature pollen grains. Usually only one of these two types of meiotic behavior predominates in a given floret, although some florets contained approximately equal numbers of both kinds of PMCs. This variation in meiotic behavior was observed in chromosome materials collected from the three largest hybrid plants as they grew in the wild state and thus represents the variation in meiosis that existed under natural conditions.
Fertility of the hybrids. An estimate of male fertility in the hybrids was obtained by staining the pollen grains with cotton blue in lactophenol and determining the proportion of empty or partially empty grains to those completely filled with cytoplasm. The number of filled pollen grains in most of the hybrids ranged around one per cent, with two outstanding exceptions (table 2). The F, hybrids 65-221 and 65-222 had apparently good pollen at frequencies of 12% and 67.5%, respectively. In the hybrid 65-221 the completely filled pollen grains were relatively large (33-37 microns) and four-pored, in contrast to the smaller (24-27 microns), three-pored grains of both parent species. The larger size and the occur- rence of four-pores suggest that these pollen grains have twice the usual chromosome number; a conclusion which is further supported by the occurrence of large (34-37 microns), often four-pored pollen grains in two of the polyploids produced experimentally from these F, hybrids. The relatively high incidence of large, four-pored, apparently functional pollen grains in hybrid 65-221 could be explained by assuming that a greater proportion of the cells, which form unreduced nuclei, ultimately produce functional pollen grains. In the case of the hybrid 65-222 all of the apparently normal pollen grains are consistently three-pored like the parent species, but these grains are the size of the pollen produced by experimental Geraea X Encelia polyploids, and thus presumably contain twice the normal number of chromosomes. This finding indicates a close correlation of ploid-level with pollen size, but gives no indication of a correlation of pore number with the degree of ploidy; a conclusion which is confirmed by the occurrence of large pollen grains (33-37 microns) that are invariably three-pored in one of the experimentally produced Geraea X Encelia polyploids. An attempt was made to test the validity of the relatively high pollen fertility estimate obtained for hybrids 65-222 by using this hybrid in a series of controlled crosses. Twenty controlled crosses of hybrid 65-222 to each of the parent species produced no filled seeds, which is the same result that was obtained with
1967] KYHOS: ENCELIA AND GERAEA 39
TABLE 2. HysripD FERTILITIES
Male Fertility Female Fertility Collection Sample Number of Percent Number of Number of Fo number size filled pollen filled seeds with _ seeds that grains grains embryos germinated 65-130 500 if 1.4 2 Z 65-158 500 q 1.4 65-159 500 4 0.8 65-221 500 60 12.0 oe 65-222 1000 675 67.5 2 _ 1 65-225 pollen sterile 4 died as a seedling 65—226 500 i, 3.4 2 Bs: 65—229 500 3° 0.6 4 1 65-330 500 6 1-2 65-231 500 3 0.6 1 65-232 died as seedling
all of the other F, hybrids. Another 30 controlled crosses of 65-222 to the other F, hybrids produced only two filled seeds, neither of which germinated. Apparently 65-222 has no greater female fertility than the other F, hybrids. It is possible that 65-222 is as pollen fertile as the cotton blue staining method indicates, but this can not be established by crosses to the other F, hybrids, since female fertility, which is con- sistently low in these hybrids, would be expected to be the important limiting factor in producing an F, progeny.
A total of 92 seeds was collected from the three largest hybrids (65- 130, 65-158, and 65-159) when they were first encountered in the field, however, none of these fruits contained embryos. Twenty controlled crosses of the ten surviving hybrids were made to each of the parent species, but these likewise failed to produce any fruits with normal embryos. Fifty controlled crosses of the F; hybrids in various combina- tions among themselves, however, produced a total of 1,186 apparently fully developed seeds, of which 13 contained embryos (table 2). Four of these 13 seeds germinated and three of these F; plants survived to maturity.
The F,. generation. All three surviving F, plants originated from two maternal parents, each with brownish-purple disk florets. Two of the three F, plants came from the only filled seeds produced by hybrid 65-130 and like their maternal parent, both had brownish-purple disk florets. These F, plants were extremely vigorous, under greenhouse con- ditions, and were nearly double the size of their parents in stature as well as the size of their individual organs. Morphologically both of these F, individuals closely resembled the majority of the F, hybrids. Their
40 MADRONO [Vol. 19
maternal parent, it will be recalled, differed from all of the other F, hybrids in that its leaf pubescence was strikingly similar to E. farinosa. Both of these F, individuals, however, had a leaf pubescence that was most like the intermediate condition of the other F, hybrids.
The exaggerated size of these F, individuals suggested that they were polyploid. A cytological examination confirmed this supposition. Each of these plants formed 36 pairs of chromosomes at meiosis and their esti- mated pollen fertility was at a comparatively high level (86.0% and 71.3%) relative to the F, hybrids. In attempting to reconstruct the origin of these F., individuals, it is probably impossible to completely rule out apomixis accompanied by spontaneous chromosome doubling as one possibility. A more likely mode of origin, however, is from the union of unreduced gametes. This interpretation is supported by the frequent occurrence of anaphase I and II meiotic nuclei with 36 chromosomes in the F, hybrids, as well as by the occurrence of large, often pour-pored, pollen grains, that presumably have double the normal chromosome number.
A third mode of origin, which requires the union of two haploid gametes, each with a balanced complement of 18 chromosomes, followed by chromosome doubling, is so improbable that it need not be considered.
The third F, individual is essentially like the other two, both in its morphology and exaggerated size. It differs conspicuously only in that its disk florets are a comparatively light brownish-purple. The cytology of this individual is more interesting, since it provides evidence of the origin of this plant. Instead of possessing a 2n chromosome number of 72 and forming 36 pairs of chromosomes at meiosis, this individual had a 2n chromosome number of 65 and consistently formed 29 pairs and 7 single chromosomes in the 40 PMCs observed. Seemingly, such a cyto- logical condition could only arise by the union of an unreduced gamete with another gamete having an unbalanced chromosome number of 29. At meiosis, in an individual possessing this combination of chromosomes, the 29 chromosomes from the unbalanced gamete would all have homol- oges with which to pair, while the remaining seven chromosomes, that were contributed by the unreduced gamete, would be left without pairing partners and thus should consistently occur as univalents, which appar- ently they do. Assuming that the 36 univalents in the F, hybrids ran- domly segregate at anaphase I, then the chance of an unbalanced gamete arising that contained 29 chromosomes would be scarcely more than one in 5000. An examination of anaphase I PMCs in the two F, hybrids that were the maternal parents of the F, plants, revealed that while the equal and near equal segregations of chromosomes, e.g. 18:18, 19:17, and 20:16, approximated a random distribution, the very unequal chromo- some segregations, such as 25:11, 26:10, and 27:9, were clearly not random. The latter categories consistently occurred in much higher fre- quencies than chance would predict. This nonrandomness probably ac- counts for the recovery of a 2n = 65 plant in a sample of only 3 F,
1967] KYHOS: ENCELIA AND GERAEA 41
plants. The cytology of this F, individual demonstrates the participation of an unreduced gamete in its origin and thus supports an interpretation of the origin of the other two tetraploid F. plants by the union of un- reduced gametes.
DISCUSSION
From this investigation it has been shown that previously undetected F, hybrids are produced, perhaps yearly, between a moderately woody shrub, E. favinosa, and an herbaceous annual, G. canescens. The occur- rence of at least some of these F, hybrids in apparently undisturbed sites suggests that appreciable numbers of these hybrids may also have existed in the long period before man’s activities created the numerous disturbed areas that are so wide-spread today. Thus, it seemed possible that such hybrids may have permitted an exchange of genetic material between these rather different taxa for a very long time. A most important factor influencing such a genetic exchange would be the fertility of the F, hybrid. Evidence of pollen fertility and crossing experiments of the Fs seems to indicate that an exchange of genes between Encelia and Geraea is extremely limited, if not essentially impossible. Not only is the fertility of these hybrids extremely low, due to a general lack of chromosome pairing in conjunction with a relatively high chromosome number, but all attempts to backcross these F, hybrids to their parent species failed to produce any progeny. On the other hand, 3 polyploid individuals were produced with comparative ease by crossing the F, hybrids among them- selves. Two of these F, individuals were extremely vigorous tetraploids, with the expected increase in fertility, whereas the third F, plant ap- proached the tetraploid state, but was deficient for 7 chromosomes and hence, not unexpectedly, had a comparatively low pollen fertility (17.6% ). This F; individual exhibited the same vigor and gigas features, in greenhouse culture, as the other F.s despite its chromosome imbalance.
The ease with which these polyploids were obtained, from the other- wise rather sterile Encelia « Geraea F, hybrids, suggests that such tetraploid derivatives are also generated in nature, but apparently have not become established. Such a possibility seems likely, considering the fact that in a two week period two clusters of 4 such hybrids were found. The 4 individuals in each of the clusters were growing within pollination distance of each other, but far enough apart to make it very unlikely that they were from the same parent plants. The frequency and close proximity of these hybrids appears to be more than adequate for the natural production of tetraploid derivatives, particularly considering the long span of time available for the necessary events to occur in. The absence of these tetraploids in nature would seem to result from their elimination by natural selection. The behavior of these hybrids in greenhouse and out-of-doors culture, perhaps provides a clue to their biological failure. As noted earlier, the tetraploids grew very vigorously in greenhouse culture, but when moved outside into: mild April weather
42 MADRONO [Vol. 19
and maintained there for the next 5 months, all three of these tetraploids progressively declined in vigor, nearly to the point of death. During this culture regime most of the flowering heads on these plants were extremely deformed and seldom produced any pollen. Upon returning the tetra- ploids to the greenhouse, they promptly resumed extremely vigorous growth. On the other hand, all of the F, hybrids also placed out-of-doors during the same time interval showed no floral deformation or decline in vigor. The inability to withstand even mild environments outside of the greenhouse was thus established as a phenomenon peculiar to the tetraploids, not effecting the very parent individuals from which these tetraploids were derived. The apparent inability of the tetraploids to grow normally outside, may well be the basis for their failure to become established in nature, however, it will be necessary to study many more such polyploids, produced from parents over a much wider geographic range, in order to properly test this explanation.
The few experimental tetraploids produced in this investigation did, however, resolve one question, namely, whether the poor meiotic chromosome pairing in the F, hybrids has a genetic or chromosomal basis. The formation of 36 normal meiotic chromosome pairs in the tetra- ploids, provides strong evidence that very different chromosome arrange- ments of the parent species, and not genetic factors, are responsible for the greatly reduced meiotic pairing in the F, hybrid.
Finally one might ask, does the finding that G. canescens and E. farinosa hybridize, necessitate any taxonomic revision? In my opinion, the answer to this question is no. In more recent taxonomic treatments Geraea and Encelia have not been considered congeneric (Abrams and Ferris, 1960; Munz, 1963; Shreve and Wiggins, 1964), however, earlier treatments (Gray, 1873; Hall, 1907) did consider them to be congeners. The mere fact that highly sterile hybrids occur naturally between these taxa, provides no compelling evidence in favor of either of these taxon- omic viewpoints. The significant criteria in this case, are to be derived, rather, from classical morphology and perhaps chromatographic evi- dence. Much additional work, particularly in the latter area is required before a taxonomic revision would be justified.
ACKNOWLEDGMENTS
A portion of this work was conducted at Stanford University as a postdoctoral trainee with the support of Grant 2G-365-R1 from the U. S. Public Health Service, National Institutes of Health. Laboratory space and equipment were generously provided by Peter H. Raven. Sincere appreciation is also expressed to William Heisey, who kindly made avail- able greenhouse and field space at the Department of Plant Biology, Carnegie Institution of Washington, Stanford, California.
1967] KOWALSKI: MYXOMYCETES 43
SUMMARY
Natural hybridization between Encelia farinosa and Geraea canescens is reported and the hybrids are described in terms of their morphology, chromatographic features, and their capacity for gene exchange. From the F, hybrids, spontaneous tetraploids were derived, which had normal chromosome pairing and restored fertility, indicating that the parent species have fundamentally different chromosome arrangements. The ease with which polyploid derivatives were experimentally obtained from the F, hybrids, suggests that they are produced in nature also, but are at a selective disadvantage, and hence have not become established. The taxonomic implications of these findings are considered.
Department of Botany, University of California, Davis
LITERATURE CITED
ABRAMS, L. and Ferris, R. S. 1960. Illustrated flora of the Pacific States. Stanford
Univ. Press.
Gray, A. 1873. Notes on Compositae. .. . Proc. Am. Acad. 8:631-661.
Harr, H. M. 1907. Compositae of southern California. Univ. Calif. Publ. Botany. 3:136.
Mowz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
Ro.iins, R. C. 1946. Interspecific hybridization in Parthenium. II. Crosses involv- ing P. argentatum, P. incanum, P. stramonium, P. tomentosum and P. hystero- phorus. Am. Jour. Bot. 33:21-30.
SHREVE F., and Wiccrns, I. L. 1964. Vegetation and flora of the Sonoran Desert. Vol. 2. Stanford Univ. Press.
NEW RECORDS OF MYXOMYCETES FROM CALIFORNIA II.
DONALD T. KOWALSKI
The new records of slime molds listed in my first paper (Kowalski, 1966) brought the total number of Myxomycetes recorded in print for California to 165. Since then, two new species have been described from the state (Brooks and Kowalski, 1966; Kowalski, 1966a). Ten new records are reported in this paper. This brings the total number of slime molds found in California to 177 species. This investigation was supported by the National Science Foundation (Grant GB-3865).
The species listed in my first paper were collected exclusively in Butte Co. Since that time, I have collected extensively in Northern California on a line north of Chico. Although a few of the species to be reported are from the Sacramento Valley, and a few from the North Coast, the majority are from alpine regions. The great majority of montane Myxo- mycetes are found only near the melting snow, but, if no rain has fallen,
44 MADRONO [Vol. 19
they may be found a few hundred yards from any snowbanks. It appears that most species actually form their fruiting bodies under the snow and that as the snow melts back, they are then exposed. This belief is based on the fact that the discovery of plasmodia in the mountains is very rare. They may be found inside or under rotten logs, but practically never out in the open where most of the fruiting bodies are found. The fact that many collections are found part way up shrubs and trees and on top of fallen twigs also reinforces this view.
All collections have been deposited in the Herbarium of the Univer- sity of Michigan. The names of the organisms are those accepted by Martin (1949) and the numbers are my own.
LICEACEAE
Licea kleistobolus G. W. Martin. On decayed wood, Lower Bidwell Park, Chico, Butte Co., 2156, Nov. 21, 1965. The sporangia are sessile, bright coppery-brown, and dehisce by a circumscissile performed lid. They are exceedingly tiny, averaging about 0.1 mm in diameter. This species has been found as far west as Colorado and is considered rare. This, however, is undoubtedly due to its inconspicuousness and it may occur throughout the United States.
Licea variabilis Schrad. Three collections, all on decayed wood: 5 miles east of Stirling City, 4000 ft., Butte Co., 2063, Aug. 19, 1965; Inskip, 4000 ft., Butte Co., 2082, Oct. 22, 1965; Eagle Lake, 5000 ft., Lassen Co., 2662, April 8, 1966. This taxon is very easy to identify because it is the only truly plasmodiocarpus form in the genus. Most of the plasmodiocarps are black, but a few are light brown in color. It is fairly common at higher elevations and has been reported pre- viously from Washington and Oregon.
CRIBRARIACEAE
Cribraria purpurea Schrad. On decayed wood, MacKerricher Beach State Park, Mendocino Co., 2355, Jan. 25, 1966. This is a handsome species being bright-reddish-purple in color. This collection is somewhat atypical in that the fruiting bodies are extremely robust, some reaching 5 mm in total height. Most descriptions give 2.5 mm as the maximum total height. However, it is easily determined by its color and the plasmodic granules which are soluble in water. It is not common, but is known from scattered localities across the United States.
Lindbladia effusa (Ehrenb.) Rost. On decayed wood, Well’s Cabin Campground, 6300 ft., Tehama Co., 1607, July 2, 1965. The fructifica- tion is an aethalium in which the individual sporangia are quite distinct. No other member of the family is aethalioid. It is probably closely re- lated to Cribraria argillacea (Pers.) Pers., which is practically identical to L. effusa except that it does not form an aethalium. It may even be that L. effusa is simply a variation of C. argillacea. L. effusa is rela- tively common throughout North America in coniferous regions.
1967 | KOWALSKI: MYXOMYCETES 45
DIANEMACEAE
Dianema depressum (List.) List. Three collections, two on decaying bark, 5 miles west of Childs’ Meadows, 5100 ft., Tehama Co., 2852, 2856, April 30, 1966, and one on duff, 4 miles east of Stirling City, 4000 ft., Butte Co., 2687, April 9, 1966. This species is distinct in the genus because it is the only member with reticulate spores. These col- lections vary from most descriptions because the capillitial threads are not penicillate. It is uncommon but has been reported from Washington and Oregon.
STEMONITACEAE
Lamproderma cribrarioides (Fries) R. E. Fries. Four collections, all on decaying bark, 5 miles east of Mineral, 5800 ft., Tehama Co., 2946, 2947, 2950, 2963, May 15, 1966. The spores of this taxon separate it from every other member in the genus. They are distinctly reticulate, the reticulations consisting of raised bands up to 1.5 » high. The spores in these collections are larger than what is given in most descriptions, often being up to 18 » in diameter. It is an exceedingly rare species, being known previously in the United States only from Colorado.
Lamproderma gulielmae Meylan. Two collections, both found 4 miles E. of Inskip, 5500 ft., Butte Co., 2806 on decaying leaf and 28/3 on decayed twig, April 23, 1966. This species has a silvery-blue peridum with black depressed spots which gives it a netted appearance. Unlike other montane Lamproderma’s, which tend to form massive collections, collections of this form usually consist of just a few scattered sporangia. Until now, it also was only known in this country from Colorado.
PHYSARACEAE
Physarum crateriforme Petch. Five collections, four on bark of live oak, Lower Bidwell Park, Chico, Butte Co.: 2089, Nov. 19, 1965; 2/71, Nov. 20, 1965; 2152, 2163, Nov. 21, 1965; and on persimmon bark, Lower Bidwell Park, Chico, Butte Co., 2230, Dec. 4, 1965. Typically, the sporangium has a large columella which can vary from white to dark brown, and a dark stripe. These collections are frequently sessile and the columella may be absent. It is thought to be rare, being re- ported only from Iowa and Kansas in this country. This, however, may be due to the fact that it grows on the bark of living trees and, thus, simply is not found often because few collectors search this type of substrate. During the winter in the Sacramento Valley it was abundant. When conditions were right, practically every rough-barked tree that I searched had this slime mold in great abundance.
Physarum diderma Rost. Four collections, all on decaying bark, Lower Bidwell Park, Chico, Butte Co.: 2320, Jan. 2, 1966; 2338, Jan. 8, 1966; 2351, Jan. 15, 1966; 2465, Feb. 7, 1966. The sporangia are sessile, crowded, pure white and have a double wall. Perhaps its most out- standing characteristic, which is not mentioned in most species descrip- tions, is the presence of a large, globose, hollow, calcareous columella,
46 MADRONO [Vol. 19
situated directly in the center of the sporangium. It has been reported from scattered localities across the United States and on the west coast from Oregon.
DIDYMIACEAE
Diderma ochraceum Hoffm. On decaying wood, Jedediah Smith Red- woods State Park, Del Norte Co., 2442, Jan. 30, 1966. The sporangia are scattered, sessile, yellowish in color, and the peridium is cartilage- nous. This species is apparently quite rare, being found previously only as far west as Tennessee.
Department of Biology, Chico State College, Chico, California
LITERATURE CITED
Brooks, T. E. and D. T. Kowatskr1. 1966. A new species of Didymium from Califor- nia. Mycologia 58:169-173. KowatskI, D. T. 1966. New records of Myxomycetes from California I. Madrono 18:140-142. . 1966a. A new species of Lamproderma from California. Mycologia 58: (In press). Martin, G. W. 1949. North American Flora 1:-190.
Notes AND NEws
Dr. ELwoop WENDELL MOtLsEED (1938-1967), Assistant Professor of Botany at the University of San Francisco, San Francisco, California passed away on April 4, 1967, after an illness of several months. He was a keen field botanist and one of those rare modern taxonomists with a green thumb. During his all too-short career, he made significant contributions to our knowledge of Mexican and Central Amer- ican Iridaceae. Plans are under way for the posthumous publication of his manu- scripts on Tigridia and related genera. His untimely death is mourned not only by his friends and associates in California, but also by those in Mexico where he spent so much time since he began his field studies on Tigridia in 1962.
CYPRESS RUSTS IN CALIFORNIA AND BAJA CALIFORNIA ROGER S. PETERSON
This study concerns two rust fungi parasitizing cypress (Cupressus ) in California (United States) and Baja California (Mexico), and the aecial stage of one of them on Amelanchier.
The only “rust” of cypress previously known in California was caused by an alga (Wagener, 1948), while no cypress rusts have been recorded in Mexico. Six rust fungi, all in Gymnosporangium, have been described from cypress found elsewhere, two of them in North America (Kern, 1964). No sori other than telia have been recorded for rust fungi on Cupressus, although a few species of Gymnosporangium produce uredinia as well as telia on other genera of Cupressaceae.
Lee Bonar, F. D. Kern, and L. N. Goodding kindly contributed advice and information to this study.
Uredo cupressicola R. Peterson, sp. nov. Urediniis planis vel cristatis, exiguis v. ad 4 X 7 mm. magnitudine, cinnamomeo-brunneis; peridio nullo. Urediniosporis late ellipsoideis v. globosis, (13-) 22-24 (-28) (21-) 24-28 (-35) uw (sextantibus maximis minimisque separatim in- clusis) ; membrana 1.5—2.0 (-3.0) » crassa, dense induta verrucis 0.3—0.4 p diam. minusque 0.5 » alt, brunnea, v. sporarum immaturarum crassiore et hyalina; poris germinationis 5-8, plerumque 5-6, papillatis, dispersis; pedicello hyalino, saltem usque 145 longo.
Habitat in caulibus Cupressi tempore tumores fusiformes faciens.
Specimens examined. On Cupressus pygmaea (Lemm.) Sarg. California. Mendo- cino Co.; E of Noyo, Peterson 64-21; Peterson 641-21; Peterson 65-120 (BPI- holotype, PUR, UC, Forest Service Herb., Logan); E of Anchor Bay, Peterson 64-33. On Cupressus arizonica var. montana (Wiggins) Little: Baja California:
Campo de Viejas between Vallecitos and La Encantada, Sierra de San Pedro Martir, Peterson 65-30.
Although U. cupressicola is presumably related to Gymnosporangium, careful search failed to reveal any sure teliospores. Urediniospores from both northern California and Baja California germinate to produce long tubes, typically with several short, near-apical branches (fig. 1). Average dimensions of 50-spore samples in water from the five collections, in the order listed above, were 22.8 &K 25.8, 22.4 & 25.4, 22.6 X 25.7, 22.9 & 25.6, and 23.2 & 28.2 w. These are remarkably similar except for the last, which was from Mexico. Somewhat distinctive dimensions might well be expected for a collection from a different cypress in a different ecologic situation at a distance of 700 miles from the other collections, and seem not to warrant taxonomic recognition in the ab- sence of additional evidence. The intersextile range of dimensions (the largest and smallest one-sixth of the 250 measured spores having been omitted) is narrow, 22-24 & 24-28 pm.
These urediniospores, except for their pedicels, seem to be indis- tinguishable from aeciospores of Gymnosporangium in general, although
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48 MADRONO [Vol. 19
Fics. 1-3. Uredo cupressicola urediniospores; 1, with typical germ tubes after 10 hours at 18° C. on 2% water agar, from type specimen, X 310; 2, including two spores that arose from a single pedicel, from type specimen, 650; 3, spores from a single stalk separated by a length of pedicel, from Baja California specimen, X 650.
1967] PETERSON: CYPRESS RUSTS 49
Rak eee RC LOONIE:
ROR IN Fa
| MET —
LAS RN ROR
Fics. 4-6. Uredo cupressicola on Cupressus pygmaea from Noyo, California, scale in millimeters: 4, uredinium between leaves on two-year-old stem; 5, fusiform gall with uredinia near both ends; 6, dead primary gall with pitted xylem exposed and larger, live secondary gall.
50 MADRONO [Vol. 19
not precisely like the aeciospores of any one species. Urediniospores are occasionally borne in tandem on a single pedicel and then appear some- what like teliospores (fig. 2) but with scattered germ pores and almost no modification of the separate walls along their common border. The two spores of such pairs cannot easily be teased apart; in fact they seem more firmly joined than are the two cells of teliospores of G. clavarii- forme (Pers.) DC. The spores from a single stalk rarely are separated by a length of pedicel (fig. 3). Erikkson (1919) pictures teliospores of G. tremelloides Hartig with separable cells that are similar to the paired spores of U. cupressicola. Perhaps the similarity suggests a homology between teliospores and paired urediniospores.
Uredinia of U. cupressicola first appear between the scale leaves of young twigs (2-year-old twigs being the youngest seen) without percept- ible hypertrophy (fig. 4). Fusiform swellings (fig. 5) that result largely from increases in xylem tissue are evident on twigs 4 years old and older. The xylem surface becomes pitted or irregular as the bark dies (fig. 6). Cankers usually kill the twigs within a few years but occasionally occur on branches up to 25 years old. Probably older infections resulted from invasion by the fungus from secondary branches into stems already too large to be killed quickly.
Uredo cupressicola was not abundant in any of the three areas where it was studied although a few trees bore hundreds of infections. At Noyo, infections were found on twigs from 2 to about 10 years old; at Anchor Bay, only infections older than 10 years were found; and in Baja Cali- fornia, infections were found on twigs only 6 to 8 years old (more than 95% of these had already been killed). Old galls also occur on Cupressus pygmaea inland from the towns of Caspar and Mendocino, but the rust fungus, if present at all on those galls, had been overgrown by dis- comycetes.
Gymnosporangium cupressi Long & Goodding, Bot. Gaz. 72:39, 1921. This fungus has previously been reported on Cupressus arizonica Greene in two areas in Arizona (Long & Goodding, 1940). Telia scarcely dis- tinguishable from those of G. cupressii are locally abundant on Cupressus bakeri Jeps. near Burney Springs in the Lassen National Forest, Shasta Co., California (Peterson 65-457 and 65-459). A few microscopic char- acteristics of the California specimens differ somewhat from Long and Goodding’s description: most notably, the thick-walled teliospores occa- sionally have warts to 2 « high near their apices, unmentioned for this species. But my one collection of G. cupressi from Arizona (62-167, from Sedona) also has teliospores with small verrucose areas. The telio- spores from California are, on the average, somewhat wider than those from Arizona, and the colorless-walled spores have even thinner walls than their Arizona counterparts.
Although most common on green twigs, all stages of gall and canker development to ages greater than a century are present. The old cankers enlarge very slowly. Where infection spreads from a branch into a trunk
|
SS ees
1967] PETERSON: CYPRESS RUSTS 51
Fic. 7. Cankers caused by Gymnosporangium cupressi var. cascadense on Cu- pressas gakeri, near Burney Springs, Shasta Co., California. Hand-axe is 18 inches long.
already several years old, host growth just about keeps pace with fungus growth and the tree may never be girdled. Old cankers (fig. 7), not previously described for G. cupressi resemble the “hip cankers” of Peridermium harknessti on pine (Peterson, 1961) with flaring sides (“hips”) of mostly uninfected reaction tissue.
Closely associated with the rust on Cupressus in one area were aecia on Amelanchier pallida Greene. About 15 bushes were heavily infected. Where Amelanchier grew within 1-2 meters of Cupressus branches, both Amelanchier and Cupressus bore hundreds or thousands of rust infec- tions per plant. Where the two hosts were several meters apart, onlv light infection was found on both. In Cupressus stands where no Ame- lanchier was found, G. cupressi was lacking (e.g., south of Cypress
oy MADRONO [Vol. 19
Camp) or was found only in cankers several decades old (e.g., in the easternmost part of the Cupressus stand along the road to Hat Creek). No Cupressaceae other than C. bakeri could be found during an intensive search covering about two acres around the heavily infected A melanchier, nor in a less intensive search in a larger area. There being a very high correlation between proximity and abundance of the aecial and telial stages, and no other Cupressaceae present to complicate the picture, it seems inescapable that the fungus on Amelanchier is the aecial form of G. cupressi as this species is represented in the Burney Springs stand.
The aecia on Amelanchier at Burney Springs differ markedly from the aecial stage of G. cupressi as it is described from eastern Arizona by Long and Goodding (1940). Height of aecia, manner of dehiscence, length of peridial cells, and especially the size of aeciospores appeared, by the published description, to distinguish the two Amelanchier para- sites. The only known Arizona collection, Goodding and Mallery 10-6- 1939 (BPI-Herb. W. H. Long No. 8439), was borrowed and some of the seeming differences were easily resolved. Long’s figures on aeciospore size, for instance, must have been based on immature spores, which are found at bases of aecia; measurements of mature spores from upper parts of the aecia do not even overlap Long’s measurements but match closely those of the California specimen. Some other differences between the two fungi were real. Because characteristics of the sporophytes from Arizona and California are so similar, and because in the aecial stage these two fungi, although distinct, are more similar to one another than to other Gymnosporangia, they can best be regarded as varieties of a single species, described as follows.
Gymnosporangium cupressi var. cupressi. Spermogonia epiphyllous, conspicuous, on discolored spots. Aecia hypophyllous, on thickened and discolored spots, cylindric, to 6 mm. high by 0.2-0.3 mm. in diameter. Dehiscent along sides and from apex by longitudinal slits. Peridial cells not hygroscopic, linear to lanceolate or oblong in face view, 11-28 X 54-93 «; in side view roughly oblique parallelograms in outline, 20-31 u thick; inner and side walls 4-11 » thick, verrucose- or tuberculate- rugose; outer walls 1-2 » thick, smooth. Aeciospores angular to globoid, 19-27 & 22-33 pw, walls 2-3 » thick, cinnamon brown, very minutely and closely verrucose; germ pores conspicuous, 5-6, scattered.
Telia as described by Long and Goodding (1940), or some teliospores with apical warts. On Amelanchier utahensis Koehne sensu lato and Cupressus arizonica Greene var. arizonica and var. glabra (Sudw.) Little in Arizona.
Gymnosporangium cupressi var. cascadense R. Peterson, var. nov. Ac G. cupressi var. cupressi simile, sed aeciis usque 3 mm. alt., cellulis ex medio peridio visis de fronte 16-36 54-130 p», plerumque 85-115 pu longis, visis de latere oblongis, 17-30 cr.; aeciospores (17-) 21-24 (-29) & (22-) 24-29 (-33) mw (sextantibus maximis minimisque separa- tim inclusis) cum poris germinationis 6-11, dispersis; teliosporiis cum
1967] PETERSON: CYPRESS RUSTS 58)
Fics. 8-11. Gymnosporangium cupressi var. cascadense on Amelanchier pallida: 8, aeclum showing initial apical dehiscence; 9 aecia lacerate to below the middle; 10, aeciospores and peridial cell in face view, X 500. 11, aeciospores and peridial cell in side view, X 750.
membranis coloratis 25-30 & 41-53 u, membranis 0.8-3.0 wu cr.; telio- sporlis cum membranis hyalinis 20-22 43-50 », membranis 0.8—1.0 pcr.
_ On leaves of Amelanchier pallida (causing swellings to 0. 6 mm. high and necrosis) and Cupressus bakert.
54 MADRONO [Vol. 19
Specimens examined. California. Shasta Co.: Lassen National Forest, W of Hat Cr. Ranger Station, Peterson 65-461 (on Amelanchier) ; S of Hat Cr.-Burney Springs road and north of Cypress Camp, Peterson 65-459 (BPI-holotype, PUR, UC, Forest Service Herb., Logan (on Cupressus); along Hat Cr.-Burney Springs road, 8 miles W of Ranger Station, Peterson 65-457 (on Cupressus).
Average dimensions of 150 aeciospores of var. cascadense mounted in water were 22.4 & 26.5 pw, versus 22.7 X 25.9 u for 50 spores of var. cupressit. Germ pores are conspicuous in aeciospores of both varieties, and their number provides the best single characteristic by which the varieties may be distinguished: 5-6 in var. cupressi and 6-11 (usually 7-8 in var. cascadense. Other distinguishing characteristics, such as heights of aecia and lengths of peridial cells, will be less reliable at least until more specimens are available of var. cupressi: the Goodding and Mallery collection consists of only two aecium-bearing infections bearing a total of four overmature aecia.
Aecia of G. cupressi (figs. 8-9) are similar to those of G. clavariuforme (Pers.) DC., which was reported (as Roestelia lacerata Fr.) on Ame- lanchier from the Sierra Nevada of California by Harkness (1887). Unfortunately, according to Bonar, no supporting specimens are avail- able, and no further aecial collections of G. clavariiforme are recorded from the state. It may be that Harkness had an early collection of G. cupressit, which may very well occur in Cupressus stands of the Sierra Nevada. Wall markings of peridial cells of the two species are distin- guishable, those of G. cupressi (figs. 10-11) tend to be coarser and more ridge-like (rather than round) on the side walls than those of G. clavartiforme. Peridial cells of the latter species also have elongate markings, but mainly on inside walls. Probably the most clearcut dis- tinction is in the aeciospore ornamentation, which is much finer in G. cupresst than in G. clavariuforme; in fact most spores of the former appear smooth in silhouette except under very high magnification. The warts of G cupressi aeciospores are generally less than 0.2 » high and occur at an average density (excluding germ pore areas) of 1.5—2 per square micron; those of G. clavariiforme are more than 0.2 » high and occur at an average density of 0.5—0.7 per square micron.
Intermountain Forest and Range Experiment Station, Forest Service, U.S. Department of Agriculture, Logan, Utah
LITERATURE CITED
ERIKKSON, J. 1919 Die schwedischen Gymnosporangeen. Sv. Vet-akad. Hand. 59:1-82.
Harkness, H. W. 1887. Fungi of the Pacific coast. V. Bull. Calif. Acad. 2:438-477.
Kern, F. D. 1964. Lists and keys of the cedar rusts of the world. Mem. N. Y. Bot. Gard. 10:305-326.
Lonc, W. H. and L. N. Goopprnc. 1940. Notes on Gymnosporangium cupressi. Mycologia 32:489-492.
PeTerSON, R. S. 1961. Western gall rust cankers in lodgepole pine. Jour. Forest. 59:194-196.
WacEnNeER, W. W. 1948. The New World cypresses. Part II. Diseases of American cypresses. Aliso 1:253-321.
TWO NEW ERIOGONUMS FROM BAJA CALIFORNIA, MEXICO
JaMEs L. REVEAL AND Craic A. HANSON
During recent herbarium studies on Eviogonum conducted inde- pendently by both authors we discovered two undescribed buckwheats collected by H. S. Gentry from the Sierra Vizcaino region of west-cen- tral Baja California, Mexico. They are as follows:
Eriogonum encelioides Reveal & Hanson, spec. nov. Frutex basi valde ramosus, 3-5 dm altus, lanatus; foliis basalibus, oblongis vel ovatis, (2—)2.5-4.5(—5) cm longis, 1.5-2.5(—3) cm latis; inflorescentia ramemosa cum bracteis 2-4 mm longis; involucris turbinato-cylindricis, 2-3.5 mm longis, 1.5-2 mm latis; perianthiis (2.5—)3—3.5 mm _ longis, albis, extus glabris, intus pilosis, segmentis oblongis vel ellipticis, (2.5—) 3-3.5 (—4) mm longis; staminibus exsertis, filamentis basi minute pilosis, antheris oblongis; acheniis exsertis, 4 mm longis, glabris. E. elongato et E. vollmeri affinis sed habitu frutescente differt et E. wrighti var. pringlei et var. nodoso foliis latioribus differt.
Low spreading loosely branched perennial shrubs 3-5 dm high with the ascending dichotomous branches densely lante-tomentose through- out, leafy in the lower portions, sheathing up the stem 1-5 cm; leaves oblong to ovate, the blades (2—)2.5-4.5(—5) cm long, 1.5-—2.5(—3) cm wide, leathery, the apices acute to obtuse, the bases acute to truncate, densely white-lanate on both surfaces, prominently veined, the petioles long, 1-2 cm long, semi-terete, lanate, tapering to short bases, these ex- tending completely, or nearly so, around the stem; inflorescence of sev- eral erect slender primary branches, with several shorter lateral spread- ing secondary branches each with 3-15 racemose involucre-bearing nodes, the internodes up to 15 cm long, these also often with several shorter tertiary branches with 3-8 nodes, these internodes up to 5 cm long; bracts triangular-deltoid, 2-4 mm long, brownish, thinly lanate, the acute linear apices usually curving inwardly, ternate, connate; in- volucres turbinate-cylindrical, 2-3.5 mm long, 1.5—2 mm wide, densely lanate between the 5 disinct, sparsely brownish lanate ribs without, glabrous within except for the lobes, the bractlets linear-oblanceolate, nu- merous, 2—3 mm long, 0.2 mm wide, hirsutulous with long marginal cells, the pedicels glabrous, 2.5—4 mm long, stoutish, 8—15-flowered; perianth (2.5—)3-3.5(—4) mm long at anthesis, white to brownish-white with a reddish-brown midrib, glabrous without, pilose within on the wide upper part of the segments, the calyx-segments similar, oblong to narrowly elliptical, 3-4 mm long, 1-1.2 mm wide, the apices rounded, the bases tapering, united only at the base; stamens exserted, the filaments up to 4 mm long, pilose at the bases, the anthers oblong, 0.6—0.8 mm long, 0.4 mm wide, white to pink; perianth 4 mm long in fruit; achenes exserted, 4 mm long, ovate, the bases tapering to a point, the apices tapering to 3-angled beaks.
55
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Type. Coarse rocky soil in a mountain pass at Portozuelo near Aguaje San Jose, Sierra Vizcaino, Baja California, Mexico, 13 Nov. 1947, Gentry 7777 (UC-holotype, AHFH, DS, SD, US).
Additional specimens examined. At Portoguelo near Aguaje San Jose, Gentry 7777a (AHFH, DS, UC) 77776 (AHFH, SD, UC, US).
Eriogonum encelioides is a member of the subgenus Eucycla (Nutt.) Kuntze and is a member of an undescribed subsection of the section Racemosa Rydb. which includes EF. elongatum Benth. and E. vollmeri Wiggins. The species name was suggested by Gentry’s label which noted that the plant resembled a “low spreading Encelia-like bush . . .”
This new species is related to EF. elongatum and E. vollmeri, but it differs from both in being more woody and shrubby with a decidedly more spreading branched paniculate inflorescence with the involucres race- mosely arranged on secondary and tertiary branches. The involucres of the new species are shorter (2—3.5 mm long) than those of E. elongatum and EF. vollmeri (5-7 mm long), but the flowers of E. encelioides are larger (3-3.5(-—4) mm long) than those of its two relatives (less than 3.5 mm long). The leaf characters vary in E. elongatum, but the leaves are never as wide as in E. encelioides and E. vollmeri, and these two species differ mainly in that the petiole of EL. encelioides is less than 2 cm long whereas £. vollmeri has petioles rarely under 3 cm long.
From £. wrightu Torr. ex Benth. the new species differs in having much larger, oblong to ovate leaves rather than the lance-elliptic leaves of EL. wrightti var. pringlei (Coult. & Fish.) Reveal and var. nodosum (Small) Reveal which the new species approaches in its woody habit.
ERIOGONUM PONDII Greene, Pittonia 1:267. 1889. Low densely branched shrubs 2.5—5 dm high with several ascending di- or trichotom- ous leafy, lanate branches; leaves 3-25 mm long, spathulate or elliptic to obovate, the blades 2-20 mm long, 2—9 mm wide, leathery, the apices acute to rounded, gradually narrowing to short petioles 1-5 mm long, densely tomentose above and below, often less so above, present the whole length of the stem except on the short flowering branches; inflor- escence of a few spreading branches, the flowering stems up to 1 cm long; bracts lacking; involucres campanulate, 3-5 mm long, 3-4.5 mm wide, densely lanate between the 5 distinct lanate ribs, the bractlets short, 1-2 mm long, linear, short-hirsutulous, the pedicels glabrous, 3-7 mm long, 10—20-flowered; perianth 2-6 mm long, white to pink, calyx-seg- ments dissimilar, the outer whorl orbicular, 2.5—5 mm long and wide, the inner whorl spathulate, 2-6 mm long, 1—2.5 mm wide, glabrous without, minutely glandular-puberulent within toward the base of each segment; stamens included, the filaments 1-4 mm long, densely yellowish-white pilose at the bases, the anthers oblong, 0.3-0.5 mm long; achenes brown, 2.5—4 mm long, ovate, the apices tapering to 3-angled beaks.
ERIOGONUM PONDII var. PONDII. Shrubs 2—3 dm high; leaves densely tomentose on both surfaces, less than 12 mm long; perianth pink, 2.5-3 mm long, the outer whorl of segments up to 3 mm long and wide, the
1967] REVEAL & HANSON: ERIOGONUM a7,
inner whorl of segments up to 3.5 mm long and 1.5 mm wide; involucres 2.5-3.5 mm long and wide, the pedicels up to 5 mm long, the bractlets 1.3—2 mm long.
Type. Cedros I., Baja California, Mexico, Dec. 1888—Feb. 1889, C. F. Pond 83 (US).
Additional specimens examined. Cedros I.: Anthony 310A (GH, US); Haines & Hale s.n. (NY, UC, US, UTC); Moran 10599 (DS, SD, US); Palmer 706 (GH, NY, US); Rose 16113 (GH, US); Rose 16114 (US). Natividad I.: Moran 10803 (GH, SD, US). San Bartolome Bay: Howell 10659a, 10708 (CAS, GH); Mason 1960 (GH, UC, US); Pond sn. (US); Rose 16203 (US).
ERIOGONUM PONDII var. gentryi Reveal & Hanson, var. nov. A var. pondii differt planta 3-5 dm alta; foliis 10-25 mm longis; involucriis 4.5-5 mm longis et 4—4.5 mm latis, pedicellis ad 7 mm longis, bract- eolis 1-1.3 mm longis; peranthiis albis, 4.5-6 mm longis, exterioribus seg- mentis ad 5 mm longis et latis, interioribus segmentis ad 6 mm longis et 2.5 mm latis.
Shrubs 3-5 dm high; leaves densely tomentose below, less so above, 1—2.5 cm long; involucres 4.5-5 mm long, 4-4.5 mm wide, the pedicles up to 7 mm long, the bractlets 1-1.3 mm long; perianth white, 4.5-6 mm long, the outer whorl of segments up to 5 mm long and wide, the inner whorl of segments up to 6 mm long and 2.5 mm wide.
Type. Near eastern bajada in gravelly arroyo, Sierra Calvario, south- eastern flank of the Sierra Vizcaino, Vizcaino Desert, Baja California, Mexico, 10—15 Mar. 1947, Gentry 7510 (UC-holotype, AHFH, DS, SD).
Additional specimens examined. Near Cerro Tordillo, Vizcaino Desert, Gentry 7441 (AHFH, DS, SD, UC).
This species belongs to the subgenus Eucycla (Nutt.) Kuntze and is a member of an undescribed section.
Variety gentry: is an inland form apparently well isolated from var. pondii which grows along the immediate coast and adjacent islands.
It is a pleasure for us to name this variety in honor of Dr. Howard S. Gentry of the Crop Research Division, U.S. Department of Agriculture, and an authority on Yucca and Agave, who not only collected the plants, but gave us additional information on both.
We are grateful to the various curators who kindly allowed us the use of their herbarium material or facilities. We wish to thank C. V. Morton for his help with the Latin diagnoses. The senior author wishes to thank the Smithsonian Institution and the U.S. National Herbarium which sponsored his Predoctoral Internship program in Washington, D.C., from September 1966 to February 1967 where this study was concluded.
Department of Botany, Brigham Young University, Provo, Utah 1228 Stratford Avenue, South Pasadena, California
58 MADRONO [Vol. 19
REVIEWS
Index to European Taxonomic Literature for 1965. Compiled by R. K. Brummitt. 166 pp. Published under the auspices of the Flora Europaea Organization by the International Bureau for Plant Taxonomy and Nomenclature, Utrecht, 1966, as Regnum Vegetabile Vol. 45. About $6.33; members of IAPT $3.00.
The importance of an annual index to botanical literature for an area the size of Europe can now be appreciated with this surprisingly vast first volume, described by the compiler as “experimental.” The geographical coverage is extended from that of the Flora Europaea project to include also the Mediterranean margin of Africa down to the Sahara, and eastward through Jordan and Syria to the Caucasus. This is a handy guide to the periodic literature on north temperate plants and it goes quite far beyond the limitations of a strictly taxonomic index. It is patterned after and geographically complements the A. E. T. F. A. T. Index (Association pour Etude Taxonomique de la Flore d’Afrique Tropicale) which has appeared since 1953, covering similar information for Tropical and Southern Africa.
The table of contents directs one to the following categories: General; Biography; Bibliography; Herbaria, Botanical Institutes, Botanic Gardens; Phytogeography; Floras, Floristic Studies; Chromosome Surveys—these taking up 20 pages. The next 114 pages include the references to Pteridophyta, Gymnospermae, and Angiospermae. The angiosperms are arranged alphabetically by family and then by genus. The appendix, 15 pages contributed by I. K. Ferguson, is a listing of the 408 abbreviated journal names cited in the Jndex followed by the complete title— and even better coverage is promised for the future!
Most Americans, I should think, would believe that the plants of Europe are quite well known, at least in comparison to other parts of the world, and will be startled to find that this Index lists 1009 new names published in 1965 for Europe and adjacent areas. This includes five new family names, six new genera, and 263 new specific names. New names are presented in bold face if they are judged to be simultaneously validly published and legitimate. Other names appear in ordinary roman type. The introduction points out that some of the names may not be legitimate or valid for reasons not readily apparent in the place cited since it was not possible to confirm all details for this essentially one man production.
The format is good and easy to use and it would not seem possible to create a volume of this size and scope without the introduction of some judgment. This is seen in the selection of family names. The use of Compositae, Cruciferae, Gramineae, Labiatae, and Leguminosae shows pleasing conservatism. Names such as Bud- dlejaceae, Cleomaceae, Leonticaceae, and Visaceae remind one that systematic arrangement of even some common plants still is in a state of confusion with many interesting and unresolved problems. Use of the name ‘“‘Capparidaceae” is a con- tradiction of the Code since only the name Capparaceae is conserved. The name Cleomaceae shows the consequences of splitting traditional groups since the genus Polanisia (herbaceous and intimately alled to Cleome) is indexed under “Cappari- daceae” while Cleome appears only under Cleomaceae. There are cross references and it is quite clear that the purpose of this Index never was to arbitrate taxonomic controversy. The placement of Ebenus under Ebenaceae and Eremurus under Gramineae are rare but quite unimportant inadvertances.
Selected indices for specific fields or areas make very convenient reference re- sources for special audiences and there is the advantage that much information can be left out resulting in more rapid publication of the Index. This Index for Europe, undoubtedly stimulated by the Flora Europea project, also will be useful to Americans. A similar kind of index would be desirable for North America now that a Flora North America project is underway. As in Europe and Great Britain, there is a limit to the number of institutions which receive enough journals with botanical inclusions. A world index to botanical literature, to include also other topics ap-
1967 | REVIEWS 59
plicable to taxonomy, already is a vacant necessity, and its realization might bring botanists closer to the much sought after stability that they all want. With the combined efforts of I. A. P. T., A. S. P. T., and other large groups of botanists it would seem feasible to begin an annual review of the literature on systematic botany for rapid publication. The Bibliography of Agriculture comes closest to serving this need.
In this age of mechanical enlightenment it is too much to expect narrowly sup- ported one-man or one-institution productions to continue indefinitely to turn out reference tools such as the Gray Herbarium Card Index, Index Kewensis, Card Index to American Botanical Literature, Index Nominum Genericorum, or Index to Plant Chromosome Numbers. There also is a need for the continuation of completed works such as Langman’s A selected guide to the literature on the flowering plants of Mexico, Merrill & Walker’s A bibliography of eastern Asiatic botany, Pfeifer’s Nomenclator, Rehder’s Bibliography of cultivated trees and shrubs, to name only a few. These noble contributions are of lasting international importance and should receive international support for continuance. With such a fine example at hand it may seem unreasonable to ask for even more but I believe that Brummitt’s Index reaffirms the need for an annual world index to botanical literature and it would be most useful to me if it were in card form or rapidly available printouts. The be- ginning of such a long term enterprise could be one of the botanical goals of the forthcoming International Biological Years and mechanical sorting methods could satisfy almost everyone with just the right references——WALLACE R. Ernst, Smith- sonian Institution, Washington, D.C.
Botanical Latin. By W1Li1AM T. STEARN. xiv + 566 pp., many illustrations. Haf- ner Publishing Company, 31 East 10th Street, New York, N. Y. 10003. 1966. $16.75.
William T. Stearn’s admirable work is not only a guide to the writing and de- ciphering of Latin descriptions and diagnoses but a valuable compendium of im- mensely varied botanical information as well. As such, it has earned an indispensable place on every systematic botanist’s workshelf. Here is no mere “dry as dust’ expo- sition of paradigms, conjugations, rules of syntax and all the other bases for correct Latin writing but a most entertainingly presented and eminently readable work re- plete with many topics of absorbing general interest. Among such, for example, is the history of the development of botanical Latin, both pre- and post-Linnaean, as an autonomous and fairly simplified and conventionalized form of expression based, however tenuously, on classical Latin. Another is an exposition of the influence of ancient and classical authors, culminating in the works of Ray and Tournefort, in supplying a concise descriptive vocabulary which was considerably elaborated and crystalized by Linnaeus — to mention only two of the many topics so felicitously touched upon. [One is indirectly admonished as to pitfalls to be avoided by the au- thor’s citation of somewhat amusing examples of bad “modern” botanical Latin.] Even as a reference compendium for such diverse matters as latinized geographical names of cities and regions, the formation of epithets based on the names of persons, standard botanical symbols and abbreviations, well illustrated examples of descrip- tive adjectives, a list of Latin color terms based on Lindley’s and J. W. Jackson’s works, pronunciation of botanical terms — again choosing at random subjects dealt with — Stearn’s excellent and often discursive though nicely organized labor of love will readily recommend itself as a work the systematic botanical worker will feel he cannot afford to deny himself.
The book is divided into four main sections: Introductory chapters on the histori- cal development of botanical Latin and a preliminary chapter on “How to Use This Book”; Grammar; Syntax and Other Matters; and Vocabulary and Bibliography.
60 MADRONO [Vol. 19
In the chapter on syntax, numerous approved examples of diagnoses and descriptions of plants from all the main groups, from Algae, Fungi and Lichens to the Angio- sperms, are proferred as models to emulate.
An unusual feature is the extensive vocabulary which devotes some 170 pages to a listing of both Latin-English and English-Latin terms under a single caption. A section which has already proved immensely instructive to the present reviewer is a chapter on the use of Greek words in botanical Latin which contains much infor- mation under such subheadings as the indebtedness of botanical Latin to Greek, the nature of word-endings, the gender of Greek nouns and adjectives, the formation of compounds, consonant-changes, a trans-literation of the Greek alphabet, and a vo- cabulary of Greek word-elements.
It is truly amazing how much information has been compressed by the author into something like 550 pages. A perusal of these pages is very likely to fill in a num- ber of gaps in one’s knowledge of classical botany and, more than incidentally, to provide several hours of most pleasurable reading. And it DOES provide, within the confines of a single cover, perhaps for the first time in English at least, all one needs to know about writing, and to a slightly lesser extent, about deciphering Latin botanical descriptions. |
Regrettable, in the first printing at least, is the misspelling of the author’s name on the spine of the work. — Rrmo BacicaLupI, Jepson Herbarium, Dept. of Botany, University of California, Berkeley.
The Grasses of North Carolina. By H. L. Bromaguist. Duke University Press, Durham, N. C., vi + 276 pp. 1948. $7.50. Grasses of the Texas Coastal Bend. By Frank W. GouLp and Tuapis W. Box. Texas A & M University Press, College Sta- tion, Texas. 189 pp. 1965. $3.25.
In a large and taxonomically difficult family such as the Gramineae, regional treat- ments are especially valuable in that they treat fewer species than a manual such as Chase (Man. Gr. U. S., rev. ed., 1950) where identification may become difficult simply because of the sheer weight of numbers of species that must be dealt with. Thus, the appearance of state and local grass floras, such as the two reviewed here, is welcome for they serve to make identification easier, thereby stimulating interest in the family and providing more information of grass species and their distribution.
The keys, generic order and, to a lesser extent, descriptions, in Blomquist’s treat- ment of North Carolina grasses all follow quite closely the pattern of Chase. As such the keys have time proven advantages (and errors) and will “run” reasonably well for most genera. A few genera, such as Aegilops, will not key in either Chase or Blomquist. The line drawings are clear and, except in a few cases where they have been too reduced, will be valuable in checking identifications. The accompanying detailed distribution maps, one for each species, clearly show the need for additional collections in North Carolina. Notes on the economic importance of various species are interesting and valuable. The chapter on distribution of grasses in North Caro- lina will be helpful both to the floristic collector and the monographer.
Gould and Box’s Grasses of the Texas Coastal Bend has been given a new and fresh approach throughout, including keys, descriptions and illustrations. The keys are well written and make use of characters that are relatively easily seen. Even sexual dimorphic species such as Buchloe dactyloides may be keyed on the basis of either male or female plants. Notes on geographic distribution, ecological relation- ships, economic importance as well as a description are provided for each species. Many of the descriptions are too short and general to be of value in checking identi- fications but in most cases a good illustration is provided. — DrENNIs ANDERSON, Division of Biological Sciences, Humboldt State College, Arcata, California
1967] REVIEWS 61
Supplement to Smith’s Marine Algae of the Monterey Peninsula. By GrorcE J. HOLLENBERG and ISABELLA A. ABBOTT. II]lustrated. Stanford University Press, Stan- ford, California. 1966. ix + 130 pp. $3.50.
As the title indicates this book is a supplement to G. M. Smith’s volume on the Monterey algae published in 1944. Because both authors have studied the Monterey algal flora extensively during the past twenty years they are well qualified to treat this subject.
The supplement essentially serves four purposes: to add 55 species to the flora, to update the nomenclature where necessary, to include revisions of descriptions of species or other taxa and revisions of keys, and finally to add new information on distribution, life histories, etc. The format of the supplement is closely similar to that of the original work by Smith. Continuity is carefully maintained through duplication of style, method of treatment, literature and taxonomic citations, and typography. Each entry is cross referenced to pages of the main volume by a system of marginal symbols.
Two departures from the coverage and format of the main work will prove most helpful. First, the range of coverage has been extended, both north and south along the peninsula. Second, all new illustrations are included as text figures with each entry rather than in plates segregated from the text. This simplifies usage con- siderably.
The authors’ failure to revise the main generic key to include the newly recorded genera is unfortunate, however. Because of this, students often will find it necessary to cross-check the supplement when identifying specimens to genus— a very time consuming process.
The illustrations are of a quality in keeping with those in Smith. Identification of various foliose species of the Cryptonemiales has been troublesome to many students; the inclusion of illustrations of thallus cross sections will be very helpful.
Lobocolax, previously considered to be a parasite on Prionitis, is excluded from the flora because the authors regard these structures as bacterial galls. This may be true but until definitive experimental evidence is available concerning their origin and development the authors’ conclusion must be accepted with reservations.
Three informational errors, although of minor significance, should be pointed out. The illustration (p. 116) said to be of Pterosiphonia gracilis Kylin actually appears to be of P. dendroidea (Montagne) Falkenberg. Comparison with an illus- tration of the type specimen of P. gracilis (Kylin, H. 1925. The marine red algae in the vicinity of Friday Harbor, Wash. Lunds Univ. Arsskr. N.F. Avd. 2. 21:1-87) will indicate clearly the distinction. An error in Smith’s book also should be pointed out at this time. The gland cells of Antithamnion subulatum (Harvey) J. Agardh are borne on a single branch cell (see Kylin, 1925) and not on two adjacent branch cells as illustrated in plate 78, figure 3 of Smith. This mistake could lead to misidentification because the manner of gland cell support is an important taxonomic criterion for some species of Antithamnion. The Pacific coast distribu- tion for Platysiphonia clevelandii (Farlow) Papenfuss is much greater than indi- cated by the authors; Scagel (1957. An annotated list of the marine algae of British Columbia and northern Washington. Nat. Mus. Canada. Bull. no. 150) records P. clevelandii for northern Washington.
The supplement has very few shortcomings and will prove to be a most useful and important addition to knowledge of the local marine algae. The authors are to be commended for a job well done—JouHn A. West, Department of Botany, University of California, Berkeley
62 MADRONO [Vol. 19
An Annotated Bibliography of Mexican Ferns. By GrorcE NEVILLE JONES. xxxill + 297 pp. University of Illinois Press, Urbana and London, 1966. $5.00.
This volume is a welcome addition to the literature dealing with the flora of our sister republic and is a convenient supplement to Ida K. Langman’s recent A Selected Guide to the Literature on the Flowering Plants of Mexico (University of Pennsylvania Press, 1964). While a certain amount of overlap between the two publications is to be expected, as they both cite a number of general works per- taining to the Mexican flora, this overlap does not appear to be excessive.
Following a short Introduction and a list of Abbreviations of Periodicals Cited is the meat of Jones’ book, 237 pages of Authors and Titles, arranged alphabetic- ally. Comments on each publication are short and to the point: Mexican taxa men- tioned are often listed, along with any pertinent notes on the nature of the publica- tion and the information contained therein. The author uses the term “ferns” broadly, including both the true ferns and the so-called “fern allies” as well. He covers not only publications dealing directly with Mexico, but also many in which plants that range into Mexico are discussed. While the bibliography is slanted toward taxonomy and phytogeography, other disciplines are not slighted. Finding Indexes (pp. 238-282) are included under four categories: General [with sub- categories of Bibliographies; Collections, Excursions, Expeditions, Explorations; Economic Botany, Horticulture and Botanical Gardens, Medicinal Plants, Useful Ferns, Symbolic Uses of Ferns, Folklore; Geographical Distribution, Phytogeog- raphy (or Biogeography), Floristics; Herbaria; Annotated Catalogues, Lists and Enumerations; Manuals, Floras, Monographs, Revisions, Keys; Nomenclators, Nomenclature, Indexes, Illustrations, Gazetteers; Vegetation: Ecology; and Mor- phology], Geographical Index (arranged by states), Biographical Index (contain- ing information on and references to authors cited), and Systematic Index (ar- ranged by genus). Truly something for everyone!
The Finding Indexes are followed by a short Supplement listing a few publica- tions and additional comments apparently compiled after the book had gone to press. Included here are only two entries published since 1962, the date through which the bibliography is intended to cover. Lastly, there is an Index of Personal Names, not including names mentioned only in the Finding Indexes.
Although this is not nearly so exhausting a bibliography for the ferns as Mrs. Langman has provided for the flowering plants, it is not to be criticized for its omission of obscure Mexican references unavailable to the author. The latter will be available for a work of this kind only when someone is willing to scour the libraries of Mexico as did Mrs. Langman.
This bibliography will prove useful not only to those interested in Mexican ferns, but also to anyone intrigued either by Mexico’s bountiful flora, or by pteri- dophytes in general. The type, while not the most aesthetically appealing, being reproduced through photo-offset from the pica of the typed original, undoubtedly has held down the price of the book to a reasonable sum.—Duncan M. Porter, Dudley Herbarium, Stanford University.
NOTES AND NEWS
Tue Genus Kickxia IN OreGon.—Kickxia spuria (L.) Dumort. apparently has not been reported for Oregon although it was found in the southwestern portion of the state as early as 1948 (Jackson Co., Jossy s.n., OSC). The increasing frequency of its collection would seem to indicate that this European species is now well es- tablished in at least Jackson, Josephine, and Douglas counties, and it has been found as far north as Marion Co.
1967] NOTES AND NEWS 63
Kickxia elatine (L.) Dumort. was introduced earlier and is more wide-spread in Oregon than has previously been recorded (C. L. Hitchcock et al., Vasc. Pl. Paczif. N. W. 4:333. 1959; R. Ornduff, Leafl. West. Bot. 9:224. 1962). These two publica- tions cite reports from Washington Co. and Multonmah Co. Specimens in the herbaria of Oregon State University (OSC) and University of Oregon (ORE) date back to 1934 and indicate a range for the species throughout western Oregon (Wash- ington and Multnomah counties to Coos, Josephine, and Jackson counties).
I am grateful to Kenton L. Chambers and Leroy Detling for assistance in the preparation of this report—La Rea J. Dennis, Department of Botany and Plant Pathology, Oregon State University, Corvallis.
CALAMAGROSTIS RUBESCENS ON SANTA Cruz ISLAND, CALIFORNIA.—The intensive program of field work being carried out by the Santa Barbara Botanic Garden has resulted in one of the most notable additions to the flora of the California Channel Islands obtained in recent years—Calamagrostis rubescens Buckl. This tall attractive rhizomatous grass was collected on Santa Cruz Island, July 22, 1960, Blakley & Muller 3726 (SBBG, DS). It grew on sandy clay soil, and was scattered in sunny spots of the pine forest on the ridge between the north and south forks of Cafion Cervada, elevation 1300 feet. It is not surprising that the species had not been col- lected in this relatively inaccessible locality previously, especially considering that it was just coming into bloom in late July. This record not only represents a range extension southward from maritime San Luis Obispo Co. (Leafl. West. Bot. 10:78. 1964), but it is the first report of any species of Calamagrostis from the California Channel Islands. As such, it fits in well with the other plants of the northern tier of islands which reach the mainland only at some distance to the north of these islands PETER H. Raven, Stanford University.
Two New ADDITIONS TO THE FLORA OF Montana.—Grindelia howelli Steyer- mark was first collected by L. F. Henderson (2791, GH-type, RM, US) on dry arid bluff tops, St. Maries River, Kootenai Co., Idaho, and has been known only from the type collection. Recent collections in 1966 (Woodland 827, 906, MONTU, DS) of G. howellit have been made in Missoula Co. at Placid Lake, in dry roadside where the road bridge crosses the stream outlet. Reconnaissance throughout the region has revealed only the presence of this Placid Lake population. G. howellii differs from the more common G. nana Nutt. and G. squarrosa (Pursh) Dunal in having a glandular and somewhat villous stem at least in the inflorescence, leaf margins which are dentate to denticulate or entire, and the middle and lower leaves are clasping and not narrowed at the base. The stems of G. nana and G. squarrosa are essentially glabrous, and the leaves are scarcely clasping and more narrowed to the base. The identification of G. howellii was confirmed by Arthur Cronquist.
Saussurea americana Eaton has been known in the Pacific Northwest only as far east as central and northern Idaho. Booth and Wright (Flora of Montana, Part II, 1966) do not report the genus as occurring in Montana. Four different collec- tions and three county records are now available for western Montana: Lincoln Co.; Leigh Lake, Cabinet Mts. Wild Area, 5500 ft., Woodland 854 (MONTU, DS). Mineral Co.; near Hoodo Pass, Stickney 1401 (Intermountain Forest & Range Ex- periment Station Herbarium, Missoula). Ravalli Co.; above big bend of Fred Burr Canyon 6000 ft., McDonald 1797 (IFRES) ; Lost Horse Creek, Bitterroot Mts., 5400 ft.. Thomas 11365 (MONTU, DS). All collections were made in early August.— Dennis W. Wooptanp, Department of Botany, University of Montana, Missoula.
64 MADRONO [Vol. 19
FLora NortH AMERICA.—The newly formed Editorial Committee of Flora North America, as the project will be called, met for the first time on January 30, 1967, at the Smithsonian Institution in Washington, D.C. This three-day meeting, con- vened by William L. Stern (Smithsonian), Chairman pro tem. of the Steering Com- mittee, was attended by all members of the Editorial Committee: Peter H. Raven, Chairman, Stanford University; Stanwyn G. Shetler, Secretary, Smithsonian Insti- tution; John H. Beaman, Michigan State University; Kenton L. Chambers, Oregon State University; Robert Kral, Vanderbilt University; Walter H. Lewis, Missouri Botanical Garden; John T. Mickel, Iowa State University; Roy L. Taylor, Canada Department of Agriculture, Ottawa; John H. Thomas, Stanford University; also attending were Robert F. Thorne (Rancho Santa Ana Botanic Gardan), Chairman of the Advisory Council, and Vernon H. Heywood (University of Liverpool), Sec- retary of Flora Europaea, who served as a consultant in the discussions and gave a concluding public lecture, “Flora Europaea, Its Conception and History,” on 1 February. The purpose of the project is to prepare a concise diagnostic manual to the vascular plants of the continental United States, Canada, and Greenland, and the Editorial Committee dealt at least in a preliminary way with a large range of questions concerning the roles of the respective committees, the functioning of the Editorial Committee and its secretariat, the solicitation of authors and advisors, and the format, arrangement, timetable, and funding for Flora North America. It is expected that the first 12 to 18 months will be occupied getting the project fully or- ganized and the working procedures implemented. This will be followed by the second phase of intensive writing and editing for the first volume. Tentatively, four volumes, followed by a fifth comprising a theoretical symposium on the North American flora, are anticipated. The whole effort is expected to last 12-15 years. A full progress report on Flora North America will be published at an early date. The Editorial Committee will convene its next meeting at College Station, Texas, in August 1967, when the American Institute of Biological Sciences holds its annual meetings at the Agricultural and Mechanical College of Texas. The first meeting was financed by the Smithsonian Office of Systematics (Richard S. Cowan, Direc- tor) —STANWYN G. SHETLER, Smithsonian Institution, Washington, D.C.
HETERANTHERA LIMOSA IN CALIFORNIA. — In June, 1966, Donald E. Seaman (at that time, Assistant Research Botanist, University of California, Davis) noted in a rice field in Glenn County a water weed, H. limosa (Sw.) Willd. (“duck salad’, or “mud plaintain’’), of the family Pontederiaceae, which had not previously been re- ported in California. On June 30, we visited the area with Seaman to observe the plant and to make a large collection (McCaskill 713 and Tucker, DAV).
This location is on the farm of Harvell Hansen, Sr., about three miles southwest of Codora, Glenn Co. According to Carl Wick, Butte Co. Farm Advisor, about 100 acres are infested here. We observed it in a number of individual plots between County Road U and Willow Creek, 1.2 miles south of County Road 61 (‘Riz Road”). It was most abundant along the west side of several of the low levees be- tween plots, in water 8-10 inches in depth, the leaf blades and flowering peduncles projecting above the surface. Of the plants in flower that we examined closely, most were also maturing seed in abundance. Flower color was consistently white, although it is often described in manuals as blue. Wick has subsequently reported finding a second infestation in Glenn Co. (ca. 3 miles southeast of Willows), and two in Butte Co. (one at Biggs, the other just north of Richvale).
Current manuals record H. limosa for the southern and midwestern states, and indicate a range extending as far west as Colorado, New Mexico, southeastern Ari- zona, and Sonora. Although apparently not previously reported in the literature in California, Thomas C. Fuller, California State Department of Agriculture, Sacra- mento, informs us that two sheets of this species, collected in Glenn Co. in 1948, are filed in the State Department of Agriculture Herbarium in Sacramento. Thus, since the plant has existed in this rice-growing area for at least eighteen years, it probably does not pose a very serious weed threat—JoHn M. TucKeEr and JuNE McCAsKILL, Department of Botany, University of California, Davis.
pst
MADRONO
VOLUME 19, NUMBER 3 JULY, 1967 Contents CHECK LIST OF CALIFORNIA PTERIDOPHYTES, Lawrence L. Kiefer and Barbara Joe 65 A New HESPERALOE FROM SonorA, Mexico, Howard Scott Gentry 74
STUDIES OF JUNIPER RUSTS IN THE WEST, Roger S. Peterson 79
A NEw SPECIES OF PTERIGONEURUM FROM CALIFORNIA, Howard Crum 92
Reviews: Wilmere Jordan Neitzel, The Flora and Fauna of Solano County (John H. Thomas); Rupert C. Barneby, Atlas of North American Astragalus (John H. Thomas); Gwenda L. Davis, Systematic Embryology of the Angiosperms (Marion S. Cave) 94
Notes AND NEws: LASTHENIA GLABRATA REMAINS DI- PLoID, Robert Ornduff 78
STEENS MounrTaIn, Kenton L. Chambers 91
TERRESTRIAL VASCULAR PLANTS OF ANo Nuevo I., SAN Mateo Co., CALirorniA, John H. Thomas ; NEw PUBLICATIONS 95
A WEST AMERICAN JOURNAL OF BOTANY
‘UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
Second-class postage paid at Berkeley, California. Return requested. Established 1916. Subscription price $6.00 per year ($4.00 for students). Published quarterly in January, April, July, and October by the California Botanical Society, Inc., and issued from the office of Madrono, Herbarium, Life Sciences Building, University of California, Berkeley, California. Orders for subscriptions, changes in address, and un- delivered copies should be sent to the Corresponding Secretary, California Botanical Society, Department of Botany, University of California, Berkeley, California 94720.
BOARD OF EDITORS
EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LyMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. EMLEN T. LITTELL, San Francisco, California Mitprep E. Marutas, University of California, Los Angeles RoBERT ORNDUFF, University of California, Berkeley Marion OwnsBey, Washington State University, Pullman REED C. Rotiins, Harvard University, Cambridge, Massachusetts Tra L. Wiccrns, Stanford University, Stanford, California
Editor—JoHn H. THomAs Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—Douglas M. Post Biology Department, San Francisco State College 1600 Holloway Avenue, San Francisco, California 94132
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Carl W. Sharsmith, Department of Biological Sciences, San Jose State College, San Jose. First Vice-President: Robert Ornduff, De- partment of Botany, University of California, Berkeley. Second Vice- President: Robert F. Thorne, Rancho Santa Ana Botanic Garden, Clare- mont. Recording Secretary: Mary L. Bowerman, Department of Botany, University of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Department of Botany, University of California, Berkeley. Treasurer: Douglas M. Post, Biology Department, San Francisco State College.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Malcolm A. Nobs, Car- negie Institution of Washington, Stanford; the Editor of Madrofo; and three elected Council Members: Annetta Carter, Department of Botany, University of California, Berkeley; Virginia M. Page, Department of Biological Sciences, Stanford University; and Herbert L. Mason, De- partment of Botany, University of California, Berkeley.
CHECK LIST OF CALIFORNIA PTERIDOPHYTES
LAWRENCE L. KIEFER and BARBARA JOE
Those engaged in identifying California plants are well aware of the different toxonomic treatments given pteriodophytes in the standard manuals and floras. These treatments are confusing, but it must be re- membered that they were prepared at different times and by different people. In addition, specialists and nonspecialists alike must reckon with recent name changes, newly established populations, and occa- sional new records of pteridophytes not previously known from Cali- fornia. Thus, there is no readily available single source which can be used to ascertain the correct name or to determine the number and kinds of California pteridophytes. It follows that a check list giving in- formation of this nature is in order and is especially needed since ferns are increasingly being used for general research and are receiving more public interest.
In this check list we have endeavored to select the most widely ac- cepted name. So users of this check list may have a broader scope of in- formation, we thought that it would be helpful to list after each entry the corresponding name and page on which these names may be found in three standard works. These are: Illustrated Flora of the Pacific States, Vol. I (1940) by LeRoy Abrams, however, the treatment of the pteridophytes was done by William R. Maxon except for the Isoetaceae which were done by Norma E. Pfeiffer; A Manual of the Flowering Plants of California (1925) by Willis Linn Jepson; and A California Flora (1959) by Philip A. Munz. Where the check list entry is absent in all three books, a reference to a bibliography or herbarium record which first reports the occurrence of the plant in California is given. All other bibliographies have been omitted. Synonymous generic names used in the three standard works are listed with cross references. Synonymous names will be found in the text following the accepted name. Plants fall- ing into the categories of subspecies or variety are entered but forms have been omitted. Taxa deemed in need of further study or are of doubtful occurrence in California are marked with an asterisk. Many of these problem plants are being studied currently.
It is inevitable that changes will be required in this check list as the many gaps in our knowledge of California pteridophytes are filled. Until then, however, we hope that it will be helpful.
We wish to acknowledge the kind help and careful attention given to checking the manuscript by Ira L. Wiggins of Stanford University, Rolla Tryon of Harvard University, Warren H. Wagner, Jr. of the University of Michigan, Thomas R. Pray of the University of Southern California, and C. V. Morton and David B. Lellinger of the Smithsonian Institution.
Maprono, Vol. 19, No. 3, pp. 65-96. Aug. 31, 1967. 65
66 MADRONO [Vol. 19
OPHIOGLOSSACEAE. Adder’s-tongue Family
*BOTRYCHIUM LUNARIA (L.) Swartz, Moonwort, Abrams, 3; Jepson, 25; Munz, 30, misinterpreted as B. lunaria var. minganense (Victorin) Dole, a variety not native to California.
*BOTRYCHIUM MULTIFIDUM (Gmel.) Rupr. ssp. COULTERI (Underw.) Clausen, Leather Grape Fern. Abrams, not listed; Jepson, 26, as B. silai- folium var. coulteri (Underw.) Jepson; Munz, 29. Infraspecific status not recognize by W. H. Wagner (pers. comm., 1966).
*BOTRYCHIUM MULTIFIDUM ssp. SILAIFOLIUM (Presl) Clausen. Abrams, 4, as £ silaifolium Presl and B. californicum Underw.; Jepson, 26, as B. silaifolium var. californicum (Underw.) Jepson; Munz, 29. (Wagner, W. H., pers. comm., 1966).
BOTRYCHIUM SIMPLEX E. Hitchc., Least Grape Fern. Abrams, 3; Jep- son, 25; Munz, 29.
*BOTRYCHIUM SIMPLEX var. COMPOSITUM (Lasch) Milde. Abrams, 3, included with the species; Jepson, 25, included with the species; Munz, 29, as B. multifidum var. compositum, an erroneous combination.
*OPHIOGLOSSUM CALIFORNICUM Prantl, California Adder’s-tongue. Abrams, 2; Jepson, 25; Munz, 29. Regarded as a variety of O. lusitan- icum L. by W. H. Wagner (pers. comm., 1966).
OPHIOGLOSSUM VULGATUM L., Adder’s-tongue. Abrams, 2, but not in- cluded for California; Jepson, 25; Munz, 29.
PoLYPODIACEAE. Fern Family
ADIANTUM CAPILLUS-VENERIS L., Southern Maidenhair. Abrams, 24; Jepson, 29; Munz, 38.
ADIANTUM JORDANII C. Muell., California Maidenhair. Abrams, 24; Jepson, 29, as A. emarginatum D. C. Eaton; Munz, 38.
ADIANTUM PEDATUM L., Northern Maidenhair. Abrams, 24, included with A. pedatum var. aleuticum Rupr.; Jepson, 29; Munz, 38, included with A. pedatum var. aleuticum.
* ADIANTUM PEDATUM var. ALEUTICUM Rupr. Abrams, 24; Jepson, 30; Munz, 38.
ADIANTUM X TRACYI C. C. Hall ex Wagner.Not in California refer- ences (Wagner, W. H., Madrofo 13:198, 1956).
ALEURITOPTERIS: see Notholaena californica.
ASPIDIUM: see Dryopteris and Thelypterts.
AspipoTis: see Cheilanthes californica.
ASPLENIUM SEPTENTRIONALE (L.) Hoffm., Forked Spleenwort. Not in California references (Howell, J. T., Base Camp Botany. Sierra Club of California. p. 3. Sept. 10, 1942; Ewan, J., Am. Fern Journ. 33:29. 1943).
ASPLENIUM VESPERTINUM Maxon, Western Spleenwort. Abrams, 18; Jepson, 35, as A. trichomanes L. var. vespertinum (Maxon) Jepson; Munz, 44.
ASPLENIUM VIRIDE Huds., Green Spleenwort. Abrams, 19, but not in- cluded for California; Jepson, not listed; Munz, 45.
1967] KIEFER & JOE: CALIFORNIA PTERIDOPHYTES 67
*ATHYRIUM ALPESTRE (Hoppe) Rylands var. AMERICANUM Butters, Alpine Lady Fern. Abrams, 20, as A. americanum Maxon; Jepson, 29, as Phegopteris alpestris Hoppe var. americanum (Butters) Jepson; Munz, 43.
*ATHYRIUM FILIX-FEMINA (L.) Roth var. CALIFoRNICUM Butters, California Lady Fern. Abrams, 19, included with the species; Jepson, 35, as var. californicum but some plants misinterpreted as var. angustum (Willd.) Farwell, a variety not native to California; Munz, 43.
*ATHYRIUM FILIX-FEMINA var. SITCHENSE Rupr. Abrams, 19, in- cluded with the species; Jepson, 35; Munz, 43.
BLECHNUM SPICANT (L.) J. Smith, Deer Fern. Abrams, 21, as Stru- thiopteris spicant Weiss; Jepson, 34, as Lomaria spicant Desv.; Munz, 44.
CHEILANTHES CALIFORNICA (Hook.) Mett., California Lace Fern. Abrams, 25; Jepson, 30; Munz, 35, as Aspidotis californica Nutt. ex Copel.
CHEILANTHES X CARLOTTA-HALLIAE Wagner and Gilbert. Not in Cali- fornia references (Wagner, W. H. and E. F. Gilbert, Am. Jour. Bot. 44: 738. 1957).
CHEILANTHES CLEVELANDII D. C. Eaton, Cleveland’s Lip Fern. Abrams, 29; Jepson, 32; Munz, 34.
CHEILANTHES COOPERAE D. C. Eaton, Cooper’s Lip Fern. Abrams, 26; Jepson, 31; Munz, 33.
CHEILANTHES COVILLEI Maxon, Coville’s Lip Fern. Abrams, 28; Jep- son, 32; Munz, 34.
CHEILANTHES FEEI Moore, Slender Lip Fern. Abrams, 27; Jepson, 31; Munz, 33.
*CHEILANTHES FIBRILLOSA Davenp., Fibrillose Lip Fern. Abrams, 27; Jepson, 31; Munz, 33.
CHEILANTHES GRACILLIMA D. C. Easton, Lace Fern. Abrams, 28; Jep- son, 31; Munz, 33.
CHEILANTHES INTERTEXTA Maxon, Coastal Lip Fern. Abrams, 28; Jepson, 32, as C. covillei var. intertexta Maxon; Munz, 34.
*CHEILANTHES PARISHII Davenp., Parish’s Lip Fern. Abrams, 27; Jepson, 31; Munz, 33.
CHEILANTHES SILIQUOSA Maxon, Claw Fern; Indian’s Dream. Abrams, 26; Jepson, 33-34, as Pellaea densa Hook.; Munz, 27, as Ony- chium densum Brack.
CHEILANTHES VISCIDA Davenp., Viscid Lip Fern. Abrams, 26; Jepson, 31; Munz, 33.
CHEILANTHES wooTonit Maxon, Wooton’s Lip Fern. Abrams, not listed; Jepson, not listed; Munz, 34.
*CRYPTOGRAMMA ACROSTICHOIDES R. Br., American Parsley Fern. Abrams, 22; Jepson, 34; Munz, 37. Possibly a variety of C. crispa (L.) R. Br. according to W. H. Wagner (pers. comm., 1966).
68 MADRONO [Vol. 19
CYRTOMIUM FALCATUM (L.f.) Presl, Holly Fern. Not in California references. An escape from cultivation at La Jolla, San Diego Co.: Kiefer 1479 (LA); and Big Dalton Canyon, Los Angeles Co.: Kiefer 2111 (LA).
CYSTOPTERIS FRAGILIS (L.) Bernh., Fragile Fern; Brittle Fern. Abrams, 7; Jepson, 38; Munz, 43.
DRYOPTERIS ARGUTA (Kaulf.) Watt, Coastal Wood Fern. Abrams, 16; Jepson, 37, as Aspidium rigidum Swartz var. argutum D. C. Eaton; Munz, 42.
*DRYOPTERIS DILATATA (Hoffm.) A. Gray, Spreading Wood Fern. Abrams, 17; Jepson, 38, as Aspidium spinulosum (Muell.) Swartz var. dilatatum Hoffm.; Munz, 41.
DRYOPTERIS FILIX-MAS (L.) Schott, Male Fern. Abrams, 16; Jepson, 37; as Aspidium filix-mas (L.) Swartz; Munz, 42.
GYMNOGRAMME: see Pitryogramma.
LastTrREA: see Thelypteris.
Lomartia: see Blechnum.
NOTHOLAENA CALIFORNICA D. C. Eaton, California Cloak Fern. Abrams, 33; Jepson, 28, as NV. californica but also as N. candida Hook. var. acessita Jepson; Munz, 34, misinterpreted as Aleuritopteris cretacea (Liebm.) Fourn., a synonym of NV. sulphurea (Cav.) J. Smith, a species not native to California. Notholaena californica ssp. nigrescans Ewan (listed by Munz as Aleuritopteris cretacea ssp. nigrescens) is not con- sidered distinct from the typical variety (Tryon, R., Contr. Gray Herb. 179:73. 1956).
NOTHOLAENA JONESII Maxon, Jones’ Cloak Fern. Abrams, 33; Jep- son, 28; Munz, 32, as Cheilanthes jonesu (Maxon) Munz.
NOTHOLAENA NEWBERRYI D. C. Eaton, Cotton Fern. Abrams, 32; Jepson, 27; Munz, 33, as Cheilanthes newberrvi (D. C. Eaton) Domin.
NOTHOLAENA PARRYI D. C. Eaton, Parry’s Cloak Fern. Abrams, 32; Jepson, 27; Munz, 33, as Cheilanthes parryi (D.C. Eaton) Domin.
*NOTHOLAENA SINUATA (Lagasca ex Swartz) Kaulf. var. CHOCHIS- ENSIS (Goodd.) Weath. Abrams, not listed; Jepson, 27, as the species; Munz, 33, as Cheilanthes sinuata (Lagasca ex Swartz) Domin. var. cochis- ensis (Goodd.) Munz.
OnycuHium: see Cheilanthes siliquosa.
PELLAEA ANDROMEDIFOLIA (Kaulf.) Fee, Coffee Fern. Abrams, 30; Jepson, 33; Munz, 36.
*PELLAEA ANDROMEDIFOLIA var. PUBESCENS D. C. Eaton. Abrams, not listed; Jepson, not listed; Munz, 36. This variety is not recognized by A. Tryon (Ann. Missouri Bot. Gard. 44:179. 1957). Pray intends to publish on this taxon giving it subspecies status (pers. comm., 1966).
PELLAEA BRACHYPTERA (Moore) Baker, Sierra Cliff Brake. Abrams, 31; Jepson, 33; Munz, 36.
PELLAEA BREWERI D. C. Eaton, Brewer’s Cliff Brake. Abrams, 29; Jepson, 32; Munz, 36.
1967 | KIEFER & JOE: CALIFORNIA PTERIDOPHYTES 69
PELLAEA BRIDGESII Hook., Bridges’ Cliff Brake. Abrams, 30; Jepson, 32; Munz, 36.
PELLAEA LONGIMUCRONATA Hook., Spiny Cliff Brake. Abrams, 31, misinterpreted in part as P. compacta (Davenp.) Maxon, a syn of P. mucronata (D.C. Eaton) D. C. Eaton var. californica (Lemmon) Munz and Johnston; Jepson, 33, misinterpreted in part as P. compacta and P. ornithopus Hook., the latter a syn. of P. mucronata; Munz, 35, misin- terpreted in part as P. compacta and P. mucronata var. californica (Pray, T. R., Phytomorhpology 16: in press. 1966).
PELLAEA MUCRONATA (D.C. Eaton) D. C. Eaton, Bird’s Foot Cliff Brake. Abrams, 31; Jepson, 33, as P. ornithopus Hook.; Munz, 36.
*PELLAEA MUCRONATA var. CALIFORNICA (Lemmon) Munz and John- ston. Abrams, 31, as P. compacta (Davenp.) Maxon; Jepson, 33, as P. compacta; Munz, 35, as var. californica but also partly as P. compacta. The Pellaea compacta entries in Abrams, Jepson, and Munz apparently include P. longimucronata Hook. in error as well as some forms of P. mucronata.
PHEGOPTERIS: see Athyrium alpestre.
*PITYROGRAMMA PALLIDA (Weath.) Alt and Grant, White Fern. Abrams, 21, as P. triangularis var. pallida Weath.; Jepson, 26, not listed, such plants apparently included with Gymnogramme triangularis Kaulf., a syn. of Pityrogramma triangularis (Kaulf.) Maxon; Munz, 38, as P. triangularis var. pallida (Alt., C. and V. Grant, Brittonia 12:168. 1960).
*PITYROGRAMMA SEMIPALLIDA Howell. Not in California references (Howell, J. T., Leafl. West. Bot. 9:223. 1962).
PITYROGRAMMA TRIANGULARIS (Kaulf.) Maxon, Goldback Fern; Tri- angle Fern; Stamp Fern. Abrams, 20; Jepson, 26, as Gymnogramme tri- angularis Kaulf.; Munz. 37.
*PITYROGRAMMA TRIANGULARIS var. MAXONII Weath. Abrams, 21; Jepson, not listed, such plants apparently included with Gymnogramme triangularis Kaulf., a syn. of Pityrogramma triangularis (Kaulf.) Maxon; Munz, 38.
*PITYROGRAMMA TRIANGULARIS var. VIRIDIS Hoover. Not in Cali- fornia references (Hoover, R. F., Am. Fern Jour. 56:19. 1966). Prob- ably best placed under the category of forma.
*PITYROGRAMMA VISCOSA (D. C. Eaton) Maxon, Silverback Fern. Abrams, 21, as P. triangularis var. viscosa (D. C. Eaton) Weath.; Jep- son, 27, as Gymnogramme triangularis var. viscosa Nutt. ex D. C. Eaton; Munz, 30, as P. triangularis var. viscosa.
POLYPODIUM CALIFORNICUM Kaulf., California Polypody. Abrams, 8; Jepson, 28, as P. vulgare L. var. intermedium Fern; Munz, 45.
*POLYPODIUM CALIFORNICUM var. KAULFuUssII D. C. Eaton. Abrams, not listed, such plants apparently included with the species; Jepson, 28, as P. vulgare L. var kaulfussii (D. C. Eaton) Fern.; Munz, 46.
POLYPODIUM GLYCYRRHIZA D. C. Eaton, Licorice Fern. Abrams, 8; Jepson, 28, as P. vulgare L. var. occidentale Hook.; Munz, 46.
70 MADRONO [Vol. 19
POLYPODIUM HESPERIUM Maxon, Western Polypody. Abrams, 8; Jep- son, 28, misinterpreted as P. virginianum L., a species not native to Cali- fornia; Munz, 45, as syn. P. vulgare L. var. columbianum Gilbert.
POLYPODIUM SCOULERI Hook. and Grev., Coast Polypody; Leathery Polypody. Abrams, 7; Jepson, 29; Munz, 45.
POLYSTICHUM XX CALIFORNICUM (D. C. Eaton) Underw. pro. spec., California Shield Fern. Abrams, 11, as species; Jepson, 36, as P. aculea- tum (Swartz) Roth var. californicum Jepson (a misinterpretation, how- ever, as P. aculeatum is not native to California) ; Munz. 41, as species. (Wagner, W. H., Am. Fern Jour. 53:8. 1963).
POLYSTICHUM DUDLEYI Maxon, Dudley’s Shield Fern. Abrams, 12; Jepson, 36, as P. aculeatum var. dudleyi Jepson (a misinterpretation, however, as P. aculeatum is not native to California) ; Munz, 41.
POLYSTICHUM KRUCKEBERGII Wagner. Not in California references (Wagner, W. H., Am. Fern Jour. 56:4. 1966).
*POLYSTICHUM LEMMONII Underw., Lemmon’s Shield Fern. Abrams, 10; Jepson, 37; Munz, 40. Regarded as synonymous with P. mohrioides Pres] (Wagner, W. H., Am. Fern Jour. 56:7. 1966).
POLYSTICHUM LONCHITIS (L.) Roth, Holly Fern; Narrow Holly Fern; Northern Holly Fern. Abrams, 9; Jepson, 35; Munz, 40.
POLYSTICHUM MUNITUM (Kaulf.) Presl, Western Sword Fern. Abrams, 10; Jepson, 36; Munz, 40.
*POLYSTICHUM MUNITUM ssp. CURTUM Ewan. Abrams, not listed; Jepson, not listed; Munz, 40.
*POLYSTICHUM MUNITUM var. IMBRICANS (D. C. Eaton) Maxon. Abrams, 10; Jepson, 36; Munz, 40.
*POLYSTICHUM MUNITUM ssp. NUDATUM (D. C. Eaton) Ewan. Abrams, not listed; Jepson, 36, as P. munitum var. nudatum (D. C. Eaton) Gilbert; Munz, 40.
POLYSTICHUM X SCOPULINUM (D. C. Eaton) Maxon, pro. spec., West- ern Holly Fern; Mountain Holly Fern. Abrams, 11, as species; Jepson, 36, as species; Munz, 41, as species. (Wagner, W. H., pers. comm., 1966).
PTERIDIUM AQUILINUM (L.) Kuhn var. PUBESCENS Underw., West- ern Bracken. Abrams, 23; Jepson, 30, as Pteris aquilina var. lanugin- osa (Bong.) Hook.; Munz, 32, as Pteridium aquilinum var. languinosum (Bong.) Fern.
PTERIS AQUILINA: see Pteridium aquilinum.
PTERIS CRETICA L., Cretan Brake. Not in California references. An escape from cultivation. Santa Cruz Co., Collett s.n. (DS).
PTERIS VITTATA L., Chinese Brake; Rusty Brake. Not in California references. An escape from cultivation in canyons of the San Gabriel Mountains, Los Angeles Co.: Beach 1900, 1901 (UC), Hutt 599 (UC), Kiefer 432, 1070 (UC).
STRUTHIOPTERIS: see Blechnum.
THELYPTERIS NEVADENSIS (Baker) Clute ex Morton, Sierra Water
1967] KIEFER & JOE: CALIFORNIA PTERIDOPHYTES 71
Fern. Abrams, 15, as Dryopteris oregana C. Chr.; Jepson, 37, as Aspid- ium nevadense D. C. Eaton; Munz, 42, as Lastrea oregana (C. Chr.) Copel. (Morton, C. V., Am. Fern Jour. 48:139. 1958).
THELYPTERIS PUBERULA (Baker) Morton. Abrams, 14, as Dryopteris feez C. Chr.; Jepson, 37, misinterpreted as Aspidium patens Swartz, a syn. of Thelypteris patens (Swartz) Small, a species not native to Cali- fornia; Munz, 42, misinterpreted as Lastrea augescens (Link) J. Smith, a syn. of Thelypteris augescens (Link) Munz and Johnson, a species not native to California. (Morton, C. V., Am. Fern. Jour. 48:139 1958.)
WoobsIA OREGANA D. C. Eaton, Oregon Woodsia; Western Cliff Fern. Abrams, 6; Jepson, 38; Munz, 39.
WoopsiA SCOPULINA D. C. Eaton, Mountain Woodsia. Abrams, 6; Jepson, 38; Munz, 39.
WoopWARDIA FIMBRIATA J. Smith, Giant Chain Fern. Abrams, 17, as W. chamissoi Brack.; Jepson, 34, misinterpreted as W. radicans (L.) J. E. Sm., a species not native to California; Munz, 44.
MARSILEACEAE. Pepperwort Family
* MARSILEA OLIGOSPORA Goodd, Abrams, 34; Jepson, not listed; Munz, 47. Doubtfully distinct from M. vestita Hook. and Grev., generally con- sidered to be merely a dry form of the latter (Flowers, S., Ferns of Utah, p. 67. 1944).
*MARSILEA VESTITA Hook. and Grev., Hairy Pepperwort; Water Clover. Abrams, 34; Jepson, 39; Munz, 47. Considered by some authors to be synonymous with M. mucronata A. Dr.; however, we have fol- lowed C. A. Weatherby (Jour. Arnold Arb. 24:325. 1943) in maintain- ing the distinctiveness of M. vestita.
PILULARIA AMERICANA A, Br., American Pillwort. Abrams, 34; Jep- son, 39; Munz, 47.
SALVINIACEAE, Salvinia Family
AZOLLA FILICULOIDES Lam., Duckweed Fern; Mosquito Fern. Abrams, 35; Jepson, 39; Munz, 47.
*AZOLLA MEXICANA Presl. Abrams, not listed; Jepson, not listed; Munz, 47, mentioned under the text of A. filiculoides as a possible oc- currence in California.
ISsOETACEAE. Quillwort Family
ISOETES BOLANDERI Engelm., Bolander’s Quillwort. Abrams, 37; Jep- son, 43, as J. bolanderi and as I. bolanderi var. sonnei Henderson; Munz, 25.
ISOETES BOLANDERI var. PYGMAEA (Engelm.) Clute. Abrams, 37; Jep- son, 43, as species /. pygmaea Engelm.; Munz, 25.
*ISOETES ECHINOSPORA Dur. var. BRAUNII (Dur.) Engelm. ex A. Gray. Abrams, 37, as J. braunii Dur. but not included for California; Jepson, 43; Munz, 25, as J. muricata var. hesperia Reed. This variety
1 MADRONO [Vol. 19
is listed as J. echinospora ssp. muricata var. braunti by B. Boivin (Am. Fern Jour. 51:84. 1961).
ISOETES HOWELLII Engelm., Howell’s Quillwort. Abrams, 37, Jepson, 44; Munz, 26.
*ISOETES HOWELLII var. MINIMA Pfeiffer. Not in California references (Pfeiffer, N. E., Ann. Missouri Bot. Gard. 9:142. 1922).
IsOETES NUTTALLII A. Br., Nuttall’s Quillwort. Abrams, 38; Jepson, 44: Munz, 26.
ISOETES OCCIDENTALIS Henderson, Western Quillwort. Abrams, 36; Jepson, 43, as J. lacustris var. paupercula Engelm.; Munz, 25.
*ISOETES ORCUTTII A. A. Eaton, Orcutt’s Quillwort. Abrams, 38; Jep- son, 44; Munz, 26.
EQUISETACEAE. Horsetail Family
EQUISETUM ARVENSE L., Field Horsetail. Abrams, 39; Jepson, 41; Munz, 28.
*EQUISETUM ARVENSE var. BOREALE (Bong.) A. A. Eaton. Abrams, not listed; Jepson, 41; Munz, not listed. Indications are that var. boreale is merely a shade form of the species and should not be recognized taxo- nomically (Hauke, R. L., pers. comm., 1965).
EQUISETUM X FERRISSII Clute, pro spec. Abrams, 42, misinterpreted as E. laevigatum A. Br., another species; Jepson, 40, misinterpreted in part as E. laevigatum and as E. hyemale var. intermedium A. A. Eaton, the latter a synonym of FE. X ferrissii; Munz, 27, misinterpreted as E. laevigatum, another species. (Hauke, R. L., Am. Fern Jour. 50:193. 1960).
EQUISETUM HYEMALE L. var. AFFINE (Englm.) A. A. Eaton, Scouring Rush. Abrams, 42, as E. prealtum Raf. and E. hiemale var. californicum Milde; Jepson, 40, as EF. hiemale var. californicum Milde, var. herba- ceum A. A. Eaton and var. robustum (A. Br.) A. A. Eaton; Munz, 27 and 29, as E. hyemale var. robustum (A. Br.) A. A. Eaton and var. californicum Milde. (Hauke, R. L., Am. Fern Jour. 50:191. 1960).
EQUISETUM LAEVIGATUM A. Br., Smooth Scouring Rush. Abrams, 40, as E. funstonit A. A. Eaton and E. kansanum J. H. Schaffner; Jepson, 40, as E. laevigatum but also as E. funstonu, E. funstonii vars. caespitosum (A. A. Eaton) Jepson, nudum (A. A. Eaton) Jepson and ramosum (A. A. Eaton) Jepson; Munz, 27, as E. laevigatum but also as E. funstonit and EF. kansanum. (Hauke, R. L., Am. Fern Jour. 50:192. 1960).
*EQUISETUM PALUSTRE L., Marsh Horsetail. Abrams, not listed; Jep- son, 41; Munz, not listed. Doubtful if found in California; no confirming herbarium specimens found.
EQUISETUM TELMATEIA Ehrh. var. BRAUNII (Milde) Ehrh. Giant Horsetail. Abrams, 39, misinterpreted as E. telmateia Ehrh., only the variety native to California; Jepson, 41; Munz, 27.
1967 | KIEFER & JOE: CALIFORNIA PTERIDOPHYTES US:
LycopopiAcEAE. Club-moss Family
LYCOPODIUM CLAVATUM L., Running Club-moss. Abrams, 45, but not included for California; Jepson, 41; Munz, 22.
LYCOPODIUM INUNDATUM L., Bog Club-moss. Not in California ref- erences. Dorothea Luhr (Am. Fern Jour. 48:61. 1959) reported finding this plant at Big Lagoon State Park, Humboldt Co.; however, the plant has been known from this area since 1929, Kildale 8397 (DS). |
SELAGINELLACEAE. Selaginella Family
SELAGINELLA ASPRELLA Maxon, Bluish Selaginella. Abrams, 50; Jep- son, not listed; Munz, 24.
SELAGINELLA BIGELOVII Underw., Bushy Selaginella. Abrams, 47; Jep- son, 42; Munz, 23.
SELAGINELLA CINERASCENS A. A. Eaton, Pygmy Selaginella; Gray Selaginella. Abrams, 47; Jepson, 42, as S. bryoides Underw.; Munz, 24.
*SELAGINELLA DOUGLASII (Hook. and Grev.) Spr., Douglas’ Selagi- nella. Abrams, 46; Jepson, 42; Munz, 23. Of doubtful occurrence in California (Jones, G. N., Am. Fern Jour. 54:83. 1964).
SELAGINELLA ENGELMANNII Hieron. var. SCOPULORIUM (Maxon) Reed, Rocky Mountain Selaginella. Abrams, as S. scopulorum Maxon but not included for California; Jepson, not listed; Munz, 24, as S. densa var. scopulorum (Maxon) Tryon (Reed, Phytologia 9:499. 1964).
SELAGINELLA EREMOPHILA Maxon, Desert Selaginella. Abrams, 47; Jepson, 42, misinterpreted as S. parishii Underw., a species not native to California; Munz, 24.
SELAGINELLA HANSENII Hieron., Hansen’s Selaginella. Abrams, 48; Jepson, 42, as S. rupestris (L.) Spr. var. kensenu Jepson and S. rupes- tris (L.) Spr. var. bolanderi (Hieron.) Jepson; Munz, 24.
SELAGINELLA LEUCOBRYOIDES Maxon, Mojave Selaginella. Abrams, 49; Jepson, not listed; Munz, 24.
SELAGINELLA OREGANA D. C. Eaton, Oregon Selaginella. Abrams, 48; Jepson, 42, as S. struthioloides Underw.; Munz, 24.
SELAGINELLA WALLACE! Hieron., Wallace’s Selaginella. Abrams, 49; Jepson, 42, misinterpreted as S. rupestris var. fendleri Underw. which is a synonym of S. underwoodi Hieron., a plant not native to California; Munz, 24.
SELAGINELLA WATSONII Underw., Alpine Selaginella. Abrams, 49; Jep- son, 43; Munz, 24.
University of California, Los Angeles, and Los Angeles City College, Los Angeles
A NEW HESPERALOE FROM SONORA, MEXICO
HowaArbD Scott GENTRY
Hesperalce nocturna Gentry, sp. nov. Planta perennis, acaulis, multi-cespitosa, 1—-1.4 m alta, ad 1—2 m late; foliis 100—150 cm longis, 1-2 cm latis ad basim, linearis longiapiculatis supra plana et crassa ad basim, canaliculata ad apice, viridibus striatis, margine angusto brunneo sparse-filifero, apice pungens vel asperigilliformis aetate pro- vectal inflorescentia racemosa, gracila, 1.2—2 m longa, simplice vel 1-3- ramosus; flores pedicellates, fasciculates, noctiflorus, 25-30 mm longes; tepala pruinosa-violacea, adpressa vel reflexa anthesis nocturna, 15-18 mm longa, 5 mm lata; filamenta glabra 8-9 mm longa ad basim tepala inserta; antheris sagitatis 8-9 mm longis in corolla inclusis; ovario 8-9 mm longo, oblongo; styli 8 mm longi crassi; stigmata capitati papillosi; capsulae 3-4 cm longae, 3.5—4.5 cm latae, depresso-ovoidae vel oblongae, rugosae apiculatae; semina grandii nigra, 8 & 11 mm.
Acaulescent, densely cespitose rosettes forming large clumps about 1.5 m tall and 1-2 m broad; leaves narrowly linear, 1—-1.5 m long, 1-2 cm broad near the base, striate, plane towards the base and deeply concave upwards on the upper side, deeply rounded below, tip acicular and pungent, fraying in age; margin narrow, brown, irregularly and finely filiferous; infloresecence slender, 1.5—2 m tall, a simple or 2-3- branched arching raceme on long peduncles; bracts of the peduncle dry, lanceolate, 6—4 cm or less long; bracteoles chartaceous, 3-5 mm long, obtuse to acute; flowers 24-30 mm long, fasciculate on unequal pedicels 5-18 mm long, 4 to 6 or 8 at the node, nocturnal, pruinose pink to lavender in the bud, greenish below, stipitate on a pedicellar joint; tube 2-3 mm long; tepals 15-22 mm long, reflexing at nocturnal anthesis, other-times appressed, whitish within, the outer tepals pink or lavender on the back and narrower than the inner, the inner with a broad flat keel and greenish pink or greenish lavender on the back; filaments equaling the pistil, 8-9 mm long, attached to base of tepals; anthers sagitate, versatile, 8-9 mm long; ovary 8—9 mm long above tube base, green, trigonous, oblong, roundly angled at the apex, ringed at the base with nectary; style 8 mm long, stout, stigma capitate, papil- late; capsules depressed ovoid or oblong, 3-4 cm long, 2.5-4.5 cm broad, apiculate, rugose; seeds large, black, 8 & 11 mm.
Holotype: Gentry & Felger 19985 (US), 15 miles southeast of Mag- dalena along road to Curcurpe, by Sierra Baviso, Sonora, Mexico, alt. 3200-3500 feet, May 21, 1963.
Also at the type locality: Gentry 19890 (US), April 1 (sterile) ; Felger 3458 (US), July 16, 1960; Wiggins 7132 (US), talus slope of basaltic cliff toward south end of Baviso Mts., 17 miles southeast of Magdalena, Sept. 11, 1934. The description of the flowers is drawn from preserved material collected at the time of anthesis. The dried speci- mens are much smaller.
74
1967] GENTRY: HESPERALOE aS
Fic. 1. Hesperaloe nocturna with flower, N, compared with flower and inflor- escence section of H. parviflora, P.
The two other species known in this genus occur east of the con- tinental divide, H. parviflora (Torr.) Coult. in Texas and H. funifera (Koch) Trel. in northeastern Mexico. Trelease (1902) gave an his- torical account of them. He observed A. parviflora in cultivation and
76 MADRONO [Vol. 19
Fic. 2. Hesperaloe nocturna southeast of Magdalena. The clumps are like “bear grass.”
noted considerable variation in the size and color of flowers on the same plant during the several months of flowering. In both H. parviflora and H. nocturna the flowers appear one by one at any given node, so there is little apparent progression up or down the raceme. First ex- amination of the Sonoran collections tentatively assigned them to var. engelmannit under H. parviflora, but with subsequent consideration of the morphology and biology I evaluate the differences at the specific level. The strictly nocturnal flowering of H. nocturna is of particular importance in separating it from the other two species with diurnal flowers. Speculatively, the reflexing of the tepals with their whitish interior are correlative to pollination by night-flying insects, while the red, tubular form of the diurnal flower is indicative of bird pollina- tors. Other structures, such as the large nocturnal anthers and the small diurnal anthers, may also reflect biotic adaptations. The following synoptical key shows the more contrasting characters of the two rela- tives.
Leaves large and course, 3-6 cm wide near the base and to 2 m long, nearly straight, with coarse white marginal filiferae; flowers greenish tinged with purple. . . . ee aa Leaves smaller, 1-3 cm wide near the base and 1-1.5 m ones arcuately spreading, with very fine marginal filiferae Flowers nocturnal, greenish lavender ,with a short tube; tepals reflexed at anthesis; filaments short (8-9 mm) with large anthers (8-9 mm); ovary large (8-9 mm long), oblong, imbedded in receptacle; style short (8 mm), stout. . . . . nocturna Flowers diurnal, dark red to light red, tubeless; tepals appressed at anthesis; filaments elongate (13-14 mm) with small anthers (4 mm); ovary small (4-5 mm long), ovoid, free of receptacle; style elongate (12-13 mm), slender . ..... . . . parviflora
1967 | GENTRY: HESPERALOE 77
se e,
Fic. 3. Hesperaloe nocturna southeast of Magdalena. The capsules show the gen- eric difference between it and “bear grass.”
Figure 1 compares the floral structures of H. nocturna and H. parvt- flora, drawn respectively from the type and Gentry 19966 from Richard- son, Texas. The development of a short but definite tube is unusual in the Yucceae, but has been observed in other members, as Yucca car- nerosana (Trel.) McKelvey. Of special significance is the ovarian tissue imbedded in the fleshy receptacle of H. nocturna, or a tendency of the tube to grow up around the ovary, leading towards an inferior position of the ovary exhibited in other members of the Agavaceae, where it is not always completely inferior. Manfreda guttata (Jacobi & Bouché) Rose (A. protuberans Engelm.) and Agave striata Zucc. both have Ovaries that protrude into the tube and hence are incompletely inferior.
78 MADRONO [Vol. 19
Further observations may discover still other agavoid taxa to bridge the gap between the superior ovary of the Liliaceae and the inferior ovary of the Amarillidaceae, the principal character conservatively aligning the Agaves with the Amaryllids.
Hesperaloe nocturna is patently rare, as it is not known from other collections or places. However, as much of northeastern Sonora has not been botanized, it may be expected in other Sonoran localities. This occurrence records the genus from west of the continental divide. In appearance the plants closely resemble the clumped ‘‘bear grass,” Nolina microcarpa S. Wats., which occurs in the same region (figs. 2, 3). Doyle Noel of the USDA Plant Quarantine Station in Nogales, Arizona, first called the plant to my attention. Plants are presently growing in his garden and he can be credited with making the first introduction.
Transplants are also responding well in the Desert Botanical Gar- den, Phoenix, Arizona. No native names or uses were obtained for the plant.
US. Department of Agriculture, Agricultural Research Service, Crops Research Division, Beltsville, Maryland
LITERATURE CITED
TRELEASE, W. 1902. The Yucceae. Missouri Bot. Gard. Rep. 13:27-31, pls. 1-4.
NOTES AND NEWS
LASTHENIA GLABRATA REMAINS DIPLOID.—Recently Mehra et al. (Caryologia 18:35-68. 1965) reported n= 14 for the helenioid composite Lasthenia glabrata Lindl. This count was based on plants of unspecified origin cultivated in Chandigarh, India. In view of my determination of n = 7 for plants in several populations of this species in its native California (Univ. Calif. Publ. Bot. 40:1-92. 1966), this record of an apparent tetraploid is of unusual interest. In response to my request, achenes of the Indian plants were sent through the courtesy of Prof. Mehra, and were subsequently planted in Berkeley. Upon flowering, it was clear that the plants belonged to L. chry- sostoma (F. & M.) Greene, a species known to have n= 8 and n= 16 but not n = 14. Although hypotetraploids in this species would be of unusual cytophyletic interest, examination of microsporogenesis in them indicated that they have n= 16 rather than n= I4. Meiosis was characterised by sticky chromosomes in the first meiotic division and this may have led Mehra and his co-workers to an impression of fewer chromosome pairs than exist. In fact, it is possible to reinterpret fig. 33 pub- lished by them as n = 16, if it is assumed that one of the alleged bivalents is a pair of loosely associated bivalents, and that what appear to be two sets of dissociated bivalents are each trivalents, or that perhaps the large ringlike figure is a quadriv- alent. I am indebted to T. F. Niehaus for his assistance with this problem—ROBERT ORNDUFF, Dept. of Botany, University of California, Berkeley.
STUDIES OF JUNIPER RUSTS IN THE WEST ROGER S. PETERSON
Included here are a review of Gymnosporangium species that occur in the Great Basin and a description of a new species of rust fungus on Juniperus deppeana in the Southwest. No taxonomic changes are proposed for the 11 species known in the Great Basin (which includes most of Nevada, half of Utah, and parts of California and other adja- cent States), but amendments to published descriptions are pointed out, new data on ecology, hosts, and distribution are given, and the life cycle of G. kernianum is established on the basis of inoculations. One species that does not infect juniper, G. zbocedri, is included to com- plete the roster of Gymnosporangium for the Great Basin.
Hyphenated numbers are those of the author’s collections, the first digits representing the year. Specimens are housed in the Forest Service herbarium at Logan, Utah, and in the National Fungus Collections at Beltsville, Maryland.
GYMNOSPORANGIUM IN THE GREAT BASIN Previous keys to telial stages have included as primary criteria some— but not all—of the symptoms caused by these fungi, and only some of the host organs on which telia are found. As a result, many specimens could not be reached in the keys. The following artificial key empha- sizes fungus morphology, with some attention to host range where this is constant. Descriptions of shapes apply to telia before they gelatinize.
Many teliospores with 1 or 3 cells. Telia round; brown or red-brown.
Apical germ pore usually present; on Heyderia. . . . . . . .G. libocedri Apical germ pores lacking; on ie ea ~ oe ew ww ew the le GG. RErnianum Telia elongate; orange . . ws e s ws « « G.SpEectosum
Teliospores with other than 2 cells rare or ising Many teliospore germ pores apical or scattered.
Germ pores several per cell, scattered. . . . . 2 . . . G.multiporum Germ pores 1 or 2 per cell, near crosswall or apical Telia more than 2 mm diameter; on J. sect. Juniperus. . .G. tremelloides Telia less than 2 mm diameter; on J. sect. Sabina. Most pedicel-tops swelling greatly in water. . . . . .G.inconspicuum Pedicels remaining cylindric in water. . . . . . . . . G.nidus-avis
Nearly all teliospore germ pores near crosswalls. Telia mostly broader than high: hemispherical or indefinite mounds.
Teliospores 12-26 & 39-614; a few pores apical. . . . . . G.nidus-avis
Teliospores 20-30 & 41—804; no apical pores. . . . . . G.kernianum Telia higher than broad; cylindric, conic, frustum- or tongue-shaped.
Telia cylindric or club-shaped, tan to yellow-orange . . . G.clavariiforme
Telia tapering upward; chestnut- to chocolate-brown. Telia less than 2 mm high; not on galls.
Spore tips often protuberant; pores not papillate . . G. harknessianum
Spore tips seldom protuberant; pores usually papillate . .G. kernianum Telia often more than 2 mm high; usually on galls.
Average length brown-wall spores less than 504. . . . . G. bethelii
Average length brown-wall spores more than 504. . . . G.nelsonii
80 MADRONO [Vol. 19
Fics. 1-4. 1, Aecia of Gymnosporangium harknessianum on Amelanchier; 2, aecia of G. kernianum (left) and G. inconspicuum (right) in the same collection on Amelanchier utahensis; 3, teliospores of G. harknessianum from Juniperus occiden- talis, X 515; 4, teliospores of G. kernianum from Juniperus osteosperma, X 515.
Discussion that expands upon treatments by Arthur (1934) or Kern (1911, 1964), rather than repetitive description, follows for the above species, alphabetically arranged. New western records from outside the Great Basin are listed in separate paragraphs.
GYMNOSPORANGIUM BETHELII Kern. Rhoads’ discussion (1946) sup- plements previous inadequate descriptions of symptoms. Gymnospo- rangium bethelit is abundant on Juniperus scopulorum and Crataegus rivularis in the parts of Utah, Idaho, and Wyomng that are in the Great Basin. C. rivularis in the Basin is almost restricted to riverbanks and moist canyons, and the rust is similarly restricted. Gymnosporangium
1967] PETERSON: JUNIPER RUSTS 81
bethelu has not been collected in Nevada, where Crataegus may be re- stricted to part of Elko Co. (McVaugh, 1942), and occurs mostly at lower elevations than J. scopulorum.
Gymnosporangium betheli has not previously been reported from Arizona but is common there on J. scopulorum in Oak Creek Canyon, Coconino Co. (66-37 ).
The key characteristic given above to separate G. bethelii from G. nel- nelsonti may not prove satisfactory when more collections are examined, but on account of their distinct aecial states the two fungi should be maintained as separate. The matter is discussed further under G. nelsoni, below. I did not study whether G. betheli is distinct at the spe- cies level from eastern G. globosum Farl., which is somewhat similar in all spore stages.
GYMNOSPORANGIUM CLAVARIIFORME (Pers.) DC. Krebill (1967) showed variations in morphology and effects of this species, which can cause witches’-brooms as well as woody swellings on juniper. Gymno- sporangium clavariiforme is not abundant in the Great Basin, but Krebill found that it is occasional on Juniperus communis in Utah, and he has collected it on that host just outside the Great Basin in western Wyo- ming and southeastern Idaho. It also appears to be only occasional— or at least rarely collected—on Amelanchier alnifolia and A. utahensts in Utah. It is not yet recorded from the western half of the Basin.
Gymnosporangium clavariforme occurs in Oregon on Crataegus columbiana (63-232 from North Burnt River, Baker Co.) and has previously been recorded in the other Pacific Coast States.
Aeciospores of this species are often larger than recorded in American descriptions. Sixty spores from a collection obtained from artificial in- fection on A. alnifolia (R. G. Krebill 388) measured 23-30 * 28-37 with an average of 27 & 32u. A collection on A. utahensis from north- ern Utah (Krebill 186) averaged 26 * 33, with a few spores as large as 33 xX 42yu. Specimens in which the aecia remain firm and finally dehisce apically are common but cannot be reached in present keys.
GYMNOSPORANGIUM HARKNESSIANUM Kern ex Arth. This species has not previously been recorded from the Great Basin. It occurs there not only in the eastern Sierra Nevada on the known telial host, Juniperus occidentalis (63-174 from Fallen Leaf Lake, Eldorado Co., California, and W. W. Wagener F. P. 97510 from Cascade Creek in Alpine Co., California) but also in the hills of western Nevada on new hosts: J. osteosperma and Amelanchier utahensis (65-131 and 65-132 from Flowery Ridge, Storey Co.). The latter collections were within two meters of one another. The occurrence of this fungus in relatively dry pinyon-juniper woodland establishes a newly recorded ecologic habitat as well as a new State and host record. In A. utahensis the fungus is perennial and causes cankers up to 30 cm long. Aecia range up to 15 mm high (fig. 1), exceeding those of any other species in the Great Basin and exceeding previous measurements for G. harknessianaum.
82 MADRONO [Vol. 19
Fics. 5-6. Peridial cell of Gymnosporangium kernianum: 5, in face view, < 600; 6, in side view, X 600. The concave, smooth outside wall appears to be within the cell.
Aeciospores in a 100-spore sample averaged 25 & 30, and their range of dimensions, 20-29 24-37, is much greater than previously re- corded. However, similar dimensions are present in my collections on A. pallida from California and Oregon, so the large size is not an effect of A. utahensis. A parallel situation exists for teliospores: on J. osteo- sperma the largest seen were 23 X 96n, to be compared with the 22 X 72 maxima given for G. harknessianum by Arthur (1934). How- ever, three collections on J. occidentalis all contained teliospores as long as 87, with a maximum of 98 in F. P. 97510 (cited above). The ten- dency of this species to have part of the teliospore wall concave (fig. 3) distinguishes it from some related species, for instance G. kernianum (fig. 4).
GYMNOSPORANGIUM INCONSPICUUM Kern. This is by far the most abundant Gymnosporangium on juniper and one of the most abundant species on Amelanchier in the Great Basin. McVaugh (1942) wrote that ‘“‘.. . in some areas... of the Great Basin the Amelanchier fruits may be so heavily infested that it is almost impossible to find one maturing normally; the infected fruits are often colored bright yellow or orange by the rust, so that whole bushes may appear at first glance to bear yellow or orange fruit.” It is almost entirely G. inconspicuum
1967 | PETERSON: JUNIPER RUSTS 83
that causes these color effects; other species are much less common on fruits in the Basin and are much less bright in color.
Telia can be found in almost any stand of J. osteosperma throughout the millions of acres of this species in Utah and Nevada. G. incon- spicuum is as common on older branches and larger stems as on the green twigs from which it has been reported before. Swelling of host tissue is slight or lacking in most stem infections, but the bark becomes abnormally rough. Although telia usually fall off green twigs after gela- tinization, leaving no obvious sign of infection, on woody bark they are often retained in the cracks so that the disease can be diagnosed at any time of year. Sometimes telia are associated with ill-defined witches’- brooms on J. osteosperma, but no systemic mycelium was found and a causal relation was not established. The fungus does cause woody swell- ings, but only rarely.
Gymnosporangium inconspicuum is newly recorded from Nevada (many collections, including 63-333 on A. utahensis and 63-334 on J. osteosperma from the Grant Range, Nye Co.); Idaho (66-11 on J. osteosperma from the Bear River Range, Franklin Co.); and Wyo- ming (66-7 on J. osteosperma from the Crawford Mountains, Lin- coln Co.).
Outside the Great Basin, newly recorded hosts of G. inconspicuum are J. deppeana (65-55 from the Chiricahua Mountains, Cochise Co., Arizona, and 66-36 from Oak Creek Canyon, Coconino Co., Arizona) and J. monosperma (65-58 from Bear Mountain, Grant Co., New Mexico, and 66-41 from Showlow, Navajo Co., Arizona). On J. mono- sperma the fungus appears to cause release of a few dormant buds but no well-developed witches’-brooms. On J. deppeana the telia collected were on woody twigs and, in one instance, on a large gall caused by G. speciosum. Gymnosporangium inconspicuum is also common on J. os- teosperma in the Southwest (61-44 and 66-31 from the Kaibab Plateau, Coconino Co., Arizona) and is often the only Gymnosporangium to be seen on this host under arid conditions, for instance in Petrified Forest National Park.
Although later manuals list only J. osteosperma as telial host of G. inconspicuum and have Arizona and Utah as its western limits, Blasdale (1919) recorded the species on J. occidentalis from Big Bear Valley in San Bernardino Co., California. Two of E. Bethel’s collections on which Blasdale’s record was based appear to me to have been cor- rectly identified as G. inconspicuum.
GYMNOSPORANGIUM KERNIANUM Bethel. This fungus is locally com- mon on J. osteosperma and occurs on Amelanchier utahensis virtually throughout the Great Basin. Unlike G. inconspicuum and G. nelsonii, however, it is absent from many stands where these alternate hosts occur together. It was found in Wyoming (66-5 on J. osteosperma from the Crawford Mountains, Lincoln Co.) in addition to States previously recorded.
84 MADRONO [Vol. 19
Fics. 7-10. 7-8, Galls caused by Gymnosporangium nelsoni on Juniperus sp. near Zitacuaro, Michoacan; 9, galls caused by G. speciosum on Juniperus deppeana in the Chiricahua Mountains, Arizona; 10, cross-section of a G. speciosum gall like that in fig. 9.
Arthur’s description (1934) of the aecia on Amelanchier was based on a wrong concept of the species, so Kern (1964) omitted Amelanchier as a host and substituted a corrected concept based on specimens on Pyrus, Cydonia, and Crataegus. However, in many of the Great Basin mountain ranges where G. kernianum is common, Amelanchier is the only one of these rosaceous genera present. Scores of collections of rust fungi on A. utahensis were examined; one (65-466 from the Pequop Mountains, Elko Co., Nevada) was found to include G. kernianum. The aecla on Amelanchier fruits are inconspicuous among the larger, brighter, and more abundant aecia of G. inconspicuum (fig. 2).
1967 | PETERSON: JUNIPER RUSTS 85
Previous inoculations to establish the life cycle of G. kernianum re- sulted in the production of spermogonia but not aecia on Amelanchier (Arthur, 1934). Production of aecia would have averted the 30-year misunderstanding of the species. Because Kern’s new description of the aecia, too, was not confirmed by inoculation results, on 12 May 1966, I inoculated two shoots of a potted Amelanchier alnifolia in the green- house by suspending over them telia on J. osteosperma (66—4 from Rich Co., Utah). Abundant spermogonia appeared on leaves (both surfaces) on 19 May and continued to be produced for several weeks on the inoculated shoots. Other equally succulent shoots on the same plant remained free of infection, as did inoculated seedlings of Malus sylves- tris cv. Red Delicious and cv. Winesap. Aecial initials were first observed on Amelanchier on 6 June but no aecia were mature until 20 June. Between then and 12 July when the leaves were harvested (66-16), about 50 aecia were produced from about 10 infections, and most of them began to dehisce; scores of other infections did not go beyond the spermogonial stage in spite of attempts to crossfertilize them. Aecia were mostly on lower leaf surfaces.
Aecia of this ‘“pedigreed”’ specimen are roestelioid and 1-3 mm high. Peridial cells vary from linear rhomboidal to (mostly) boat shaped: the outer surface is concave, forming the interior of the “boat,” and the sides curve inward to the rounded “bottom” (figs. 5, 6). Peridial cells from halfway up an aecium are 24-33 & 61-110, in face view and 24-41, thick. Inner and side walls are moderately verrucose for the most part, but there are usually some cylindrical tubercles 3 or 4p high, especially near the ‘“‘gunwales.”’ Sidewalls are about 4p thick and inner walls are 4-13, thick. Outer walls are smooth and usually about 1.5u thick, or occasionally as much as 3u. Aeciospores in a 50-spore sample measured 22-28 & 23-33 with an average of 25 & 28. Aecio- spore walls are yellow-brown, 1.5—2.5u thick and finely verrucose; they bear six to nine conspicuous germ pores.
Although the cultured aecia did not seem to contain any very long peridial cells, I have no doubt that they represent the same taxon in which cells more than 200p long were found (Peterson, 1963). Although mature, the greenhouse-grown aecia as well as peridial cells are smaller than in natural collections. Perhaps Amelanchier leaves are an unfavor- able substrate for the fungus in comparison with fruits of this or other genera.
Telia of G. kernianum have been described as caulicolous (Arthur, 1934). Microtome sections of telia in four specimens showed them to be of subhypodermal origin on leaves rather than on stems. Mycelium occurs throughout both leaves and stems.
Gymnosporangium kernianum is much more widely distributed than Arthur (1934) recorded. Cummins (1964) found it in Texas. It is also on J. californica in California (65-35 near Jacumba, San Diego Co., and R. G. Krebill 249 from Phelan, San Bernardino Co.) and in Baja
86 MADRONO [Vol. 19
California (65-27 from the western foothills of the Sierra San Pedro Martir)—a first record from Mexico. In Arizona it infects J. osteo- sperma (62-137 from the South Rim, Grand Canyon National Park) as well as J. deppeana.
On J. californica, a newly recorded host species, some teliospore walls are much thicker than previously recorded for G. kernianum, as much as 3 or rarely 5» thick including the occasional apical ornamentation. Most spores, however, have smooth walls 0.8-1.5u thick as on other Junipers.
GYMNOSPORANGIUM LIBOCEDRI (P. Henn.) Kern. This species with telia on Heyderia decurrens and aecia on Amelanchier and other genera is the only Gymnosporangium in the Great Basin that does not inhabit juniper. It is abundant on Heyderia in the Sierra Nevada, including the western Nevada portion of this mountain range. The common, conspicu- ous witches’-brooms caused by this fungus represent only a minute per- centage of infections, most of which do not become systemic.
This species is surely the most abundant Gymnosporangium in north- ern California, at least in recent years. It is surprising that it is quite uncommon south of Yosemite National Park, and I could find none of it on Heyderia in Baja California.
GYMNOSPORANGIUM MULTIPORUM Kern. This seldom-collected species is the only Gyvmnosporangium of unknown life cycle in the Great Basin. It occurs on Juniperus occidentalis in Eldorado Co., California, but has not yet been found in typical Great Basin habitats.
Kern and Keener (1960) hypothesized that Peridermium ephedrae is the aecial state of G. multiporum. My inoculations with P. ephedrae aeciospores, which were kindly supplied by Professor Keener, failed to produce infection on J. osteosperma, J. virginiana, or Ephedra viridis. I do not believe that any Ephedra grows near Fallen Leaf Lake, where G. multiporum was collected in Eldorado Co.
GYMNOSPORANGIUM NELSONII Arth. This species is occasional to abundant throughout the Great Basin, and is the only Gymnosporan- gium known on both principal junipers of the area, J. osteosperma and J. scopulorum. It is also abundant in some years on Amelanchier alni- folia, A. utahensis, and, very locally, on Peraphyllum ramosissimum in the Basin.
Gymnosporangium nelsonu infects J. californica in the vicinity of Jacumba, San Diego Co., California—both a new host and a new State for the fungus (65-34).
Gymnosporangium nelsoni and G. bethelu are very similar in the telial state. In my collections the overall size ranges for teliospores are 17-28 & 41-74 in G. nelsoni 15-26 & 32-65p in G. betheli. How- ever, many G. nelsoni collections lack distinctively long spores. Inter- sextile lengths (the sixth largest and sixth smallest omitted) of spores with colored walls are more helpful: 50-59» in G. nelsoni and 46—53y in G. betheli. Spores with walls that are hyaline or nearly so are pres- ent in most collections; in G. bethelii they average longer than spores
1967 | PETERSON: JUNIPER RUSTS 87
with definitely brown walls. Teliospores of G. nelsoni tend to have nar- rower ends than the rounded spores of G. bethelii. Usually the two can be distinguished because G. nelsoni in the Great Basin produces a single, subglobose gall from each infection whereas G. bethelii pro- duces a canker and a series of irregular galls. But these symptoms inter- grade and in any event are useless for identification of young infections.
Specimens tentatively identified as G. nelsonii were collected from Juniperus flaccida in the State of San Luis Potosi, Mexico (63-46, from Barranca de los Marmoles) and on Juniperus sp., sect. Sabina, in the state of Michoacan (63-46, from near Zitacuaro). These specimens, and also those from Michoacan and Texas mentioned by Cummins (1964) and one on J. deppeana from Hidalgo illustrated by Martinez (1963), display symptoms unlike any described for G. nelsoniw. The cankers with their series of lobed, stalked galls from a single infection (figs. 7, 8) resemble those of G. bethelli more than typical infections of G. nelsoni. The teliospores, however, are longer even than typical G. nelsoni and quite unlike G. bethelit.
Another doubtful assignment to G. nelsoni is collection 64-187 on leaves of Amelanchier utahensis from a near-desert woodland east of Escalante, Utah. Aeciospore walls are unusually dark chocolate brown, perhaps an example of desert melanism. Aeciospore dimensions in a 50-spore sample are 23-33 & 27-37» with an average of 27 &K 32u— larger than previously described for G. nelsoni or indeed, for any rust fungus on Amelanchier leaves except G. clavariiforme as it is described above. Peridial cells were like those of G. nelsoni, and galls of this spe- cles as well as G. inconspicuum infections were nearby on J. osteo- sperma.
GYMNOSPORANGIUM NIbuS-AVis Thaxter. This species is locally com- mon to abundant on J. scopulorum and Amelanchier spp. in the three eastern States of the Great Basin—Wyoming, Idaho, and Utah. I have not yet seen specimens from Nevada.
Gymnosporangium nidus-avis causes witches’-brooms on J. scopu- lorum similar to those of G. kernianum on J. osteosperma; also, the spore descriptions of the two species overlap to a confusing extent. Nevertheless, they are distinct fungi. Probably in all telial collections of G. nidus-avis a substantial number of spores have apical germ pores, which is not true of G. kernianum. Also they are host specific—even where J. scopulorum and J. osteosperma are closely mingled it is com- mon to see one of them heavily infected by its own peculiar broom-rust parasite, whereas the other juniper is free of rust.
GYMNOSPORANGIUM SPECIOSUM Peck. This species is locally com- mon on J. osteosperma and Philadelphus in southern and eastern Utah. Its symptoms on J. osteosperma include witches’-brooms as well as fusi- form galls (Rhoads, 1946). Sometimes whole trees appear to be in- fected, with telia in small patches scattered over the rough-barked trunks and branches.
88 MADRONO [Vol. 19
Swellings caused by G. speciosum are described as “gradually fusi- form” (Kern, 1964). The distinctive galls that it causes on J. deppeana (figs. 9, 10) are often rather abrupt, and large ones are 35 cm or more in diameter, some of them almost spherical. Others are elongate and may resemble those on J. mexicana, illustrated by Cummins (1943).
Although most witches’-brooms on J. deppeana are caused by G. kernianum, in the Santa Rita Mountains of southern Arizona I ob- served on this host occasional brooms associated with long-fusiform swellings caused by G. speciosum.
A NEw SPECIES OF Rust FUNGUS ON JUNIPERUS
Uredo apacheca R. Peterson, sp. nov. Urediniis forma irregulari, exiguis vel usque ad 3 & 10 mm, plerumque occultatis sub cortice, albidis vel pallide flavidis; peridio crassitudine cellula unica, frag- ile; cellulis peridii oblongis, plerumque 24-35 « 35-45, ca. 15-20u cr., membrana verrucosa; urediniosporis in catenis, ellipsoideis vel piriformibus, 16-33 & 25-54u, plerumque 21-26 x 27-33 mu, mem- brana inaequabiliter (1.0-) 1.5-2.0 (-—3.5)m cr., verruculosa; poris germinationis non visis. In caulibus Juniperi deppeanae, tumores fusi- formes faciens.
Specimens examined, all collected by the author in the San Fran- cisco Mountains in Catron Co., New Mexico: 66-50 (BPI-holotype), Gila National Forest about 8 road-miles south of Luna along U.S. Highway 180, Sec. 31, T. 6 S. R. 20W; 65-60, same place as the pre- ceding; 66-51, Apache National Forest, Blue River-Brushy Mountain road, Sec. 7 or 8, T. 8S. R. 21 W.
The specific epithet honors a major ethnic group of the area.
Uredo apacheca causes conspicuous, fusiform galls on alligator juni- per. Old galls become very rough-surfaced, like those described above for Gymnosporangium speciosum on the same host.
Uredo apacheca was not abundant in either locality where it was studied. One or two trees were heavily infected in each area and re- sembled the juniper pictured by Cummins (1943) that was infected by G. ?speciosum. Other trees in each locality bore only one or two galls. No young infections could be found: the youngest one (fig. 11) is at least 10 years old. The clustering of galls in the heavily infected trees suggests direct juniper-to-juniper transmission of the fungus.
Woody vegetation in both collection localities consisted largely of Pinus edulis, Juniperus deppeana, Quercus gambelu, and Cercocarpus montanus. No Pomoideae—the usual aecial hosts of rust fungi on Juniperus—were seen.
Uredo apacheca is the fifth described fungus with uredinia on Cup- ressaceae and the first on Juniperus sect. Sabina, the others being on J. sect. Juniperus, Chamaecyparis, Heyderia, and Cupressus (Kern, 1964; Peterson, 1967). It is the first species on Cupressaceae reported to have uredinia with peridia or urediniospores borne in chains (figs.
1967] PETERSON: JUNIPER RUSTS 89
Fics. 11-14. 11, Gall caused by Uredo apacheca on Juniperus deppeana; 12, peridial cells of U. apacheca, — 700; 13, a small uredinium and spores of U. apa- checa, X 140, overlying bark was cut away in preparing the section; 14, pyriform spores of U. apacheca, X 720.
12, 13). Its urediniospores (fig. 14) scarcely resemble those of any of the other four species. Uredo apacheca is more similar to another fun- gus on Cupressaceae, the Japanese species Caeoma deformans (Berk. & Br.) Tub. Host reactions to the two differ, but general morphology— large, irregularly shaped spores borne in chains under a weak peridium —are similar. Of all the rust fungi on Cupressaceae, only U. apacheca, C. deformans, and C. espinosae Syd. have light-colored spores that lack obvious germ pores. The biggest difference between U. apacheca and C. deformans is that the latter regularly produces spermogonia and is regarded as an aecial state.
90 MADRONO [Vol. 19
There was a possibility that the sori of U. apacheca, too, might be aecia rather than uredinia, although no spermogonia could be found in the infections. Nuclear constitution of the sorus initials and of the mycelium which produces them provides the critical means of dis- tinguishing the two spore stages. Sections of collection 65-60 were cut at 15» in paraffin and stained by Heidenhain’s iron alum-haematoxylon technique. Most rust hyphae in the vicinity of the sori and elsewhere were entirely composed of binucleate cells. In hyphae where apparently uninucleate cells were seen there were also binucleate cells, suggesting that the uninucleate appearance was due to artifacts of sectioning and staining. As the mycelium appeared to be dikaryotic, and there were no spermogonia, it was concluded that the sori are uredinia.
The same tissue sections showed that most urediniospores are binu- cleate, with a minority of trinucleate spores, a few quadrinucleate ones, and possibly a few uninucleate ones.
The same sections plus others from the same specimen, stained in safranin, showed that the hymenium and underlying mass of fungus tissue totaled only 40—90y thick except for occasional “‘pockets” between host cells—these are up to 160 thick. Some basal cells appeared to produce more than one chain of urediniospores by lateral budding or branching. Mycelium of the fungus is intercellular throughout the host cortex and in much of the phloem, rarely reaching the cambium, and in the one specimen sectioned mycelium was never observed in the xylem. Hyphae are thick—often as much as 7»—and somewhat con- torted, therein resembling some species of Gymnosporangium. Haus- toria are simple, thick, and usually obovoid.
Individual chains of urediniospores in some instances exceeded 50 spores in length. Narrow ends of pyriform spores and “handles” on occasional other spores pointed toward the hymenium, unlike the ar- rangement in some other species, as Cronartium comandrae Pk., with pointed spores. No clear intercalary cells were seen, but thin-walled fungus tissue in the uredinium probably represented intercalary ma- terial. The spore mass was usually covered by host bark, with only narrow openings through cracks.
Coloration, a distinctive characteristic for most rust fungi on Cu- pressaceae, is a puzzling feature in U. apacheca. Most sori appear to be nearly white, but some are yellowish. Both walls and protoplasts of most spores are hyaline, but there are carotenoid pigments in the proto- plasm of some. Spore mounts from the sori that are deepest yellow reveal a different sort of coloration: spore walls and probably spore contents are of a uniform, dingy yellow that seems unlike carotenoid pigment. I did not determine whether this color was native to the fungus or the result of leaching of tannins from the host bark.
Germination of spores from collection 66-50 was tested on 2% water agar. Germ tubes were mostly 100-150, long after 24 hours at 18° C. Tubes were 3—4 wide and were unbranched or had a few short branches
1967] PETERSON: JUNIPER RUSTS 91
near their bases. It happened that only colorless spores were included in the test, and their tubes were also colorless.
On the surface of collection 65—60 were a few scattered teliospores that probably belong to a Gymnosporangium. None could be found clearly associated with the uredinia, which are therefore described in Uredo, with the hope that this description will stimulate more thorough study of the life cycle of this remarkable fungus.
Intermountain Forest and Range Experiment Station Forest Service, U.S. Department of Agriculture, Logan, Utah Present address: St. John’s College, Santa Fe, New Mexico
LITERATURE CITED
ARTHUR, J. C. 1934. Manual of the rusts in United States and Canada. Purdue Res. Found., Lafayette. BLASDALE, W. C. 1919. A preliminary list of the Uredinales of California. Univ. Calif. Publ. Bot. 7:101-157. Cummins, G. B. 1943. Descriptions of tropical rusts—V. Bull. Torrey Club 70:68-81. . 1964. Uredinales of the Big Bend National Park and adjacent areas of Texas. Southw. Nat. 8:181—195. Kern, F. D. 1911. A biologic and taxonomic study of the genus Gymnosporangium, Bull. N. Y. Bot. Gard. 7:391—483. . 1964. Lists and keys of the cedar rusts of the world. Mem. N. Y. Bot. Gard. 10:305-326. , and P. D. KrENner. 1960. Identity of a rust on Ephedra. Science 131:298. KReEBILL, R. G. 1967. Gymnosporangium clavariiforme in witches’-brooms of Junip- erus communis. U.S. Dep. Agr., Plant Dis. Rep. 51:143-147. McVaucu, R. 1942. Rosaceae of Nevada. Contr. Fl. Nev. 22. Martinez, M. 1963. Las Pinaceas mexicanas. Ed. 3. Univ. Nac. Auton. Mexico, Mexico. PETERSON, R. S. 1962. Notes on western rust fungi. II. Pucciniaceae. Mycologia 54:389_-394. . 1967. Cypress rusts in California and Baja California. Madrono 19:47-54. Ruoaps, A. S. 1946. A contribution to the fungus flora of Utah and Nevada. U‘S. Dep. Agr., Plant Dis. Rep. Suppl. 162:67-99.
NOTES AND NEWS
STEENS Mountain.—A paper entitled, ‘A botanical excursion to Steens Moun- tain, SE Oregon, U.S.A.,” appeared in the Scandinavian journal Blyttia (24: 177-181. 1966). The author is Prof. Knut Faegri, University of Bergen, who spent a portion of the 1965 school year in residence at Oregon State University. Prof. Faegri compares the plant communities observed on Steens Mounatin with cor- responding ones in the mountains of northern Europe, and outlines his ideas on the major climatic factors affecting alpine plant distributions. Because this journal may not be readily available to botanists in western North America, reprints will be provided on request by the author (Botanisk Museum, Postboks, 2637, Bergen, Norway).—Kenton L. Cuamsers, Botany Department, Oregon State University, Corvallis.
A NEW SPECIES OF PTERIGONEURUM FROM CALIFORNIA
HowARp CRUM
Recently I discovered in the herbarium of the University of Michigan a California collection of a curious Pterigoneurum which had been er- roneously assigned to P. subsessile (Brid.) Jur. Although related to P. subsessile, it differs from that species in its compact, bulbiform habit of growth (especially before complete maturity), smooth spores, and cucullate calyptrae. In these and other less tangible characteristics which result in a distinctive aspect, the plants clearly warrant description as a new species.
Pterigoneurum californicum n. sp. Plantae minutae, ca. 1.5 mm altitudine, plus minus bulbiformes, gregariae. Folia incurva et imbricata vel laxe erecta, concava, usque ad 2 mm longitudine, in pilo hyalino usque ad 1 mm terminata, marginibus integris vel superne serrulatis, costa ex- currente, cellulis rhomboideis laevibusque. Capsula immersa, subhemis- phaerica, sicca et madida irregulariter rugosa, gymnostoma, usque ad 0.8 mm altitudine, operculo conico-rostrato. Sporae 22-29 mu, sphaericae vel breviter ellipsoideae, brunneae, laeves. Calyptra cucullata et laevis, operculum non exedens.
Plants very small, about 1.5 mm high, bulbiform, becoming more loosely foliate with maturity, dull, yellow-brown, loosely gregarious. Leaves few, incurved and imbricate or loosely erect, keeled-concave, small below, larger above, the upper leaves 1-2 mm long (exclusive of the awn), oblong-ovate, oblong, and oblong-obovate, obtuse, abruptly narrowed to a hyaline, flexuose, somewhat serrulate awn up to about 1 mm long; margins erect, subentire to serrulate near the apex, especially at the base of the awn; costa strong, brown, excurrent into and filling the awn; lamellae on the upper surface of the costa 4, somewhat irregu- lar at the margin; upper cells smooth, irregularly hexagonal to oblong- rhomboidal, about 10-15 » wide, 1-2, sometimes 3:1, with thin to rather firm walls not thickened at the corners; basal cells laxly rectangular. Presumably autoicous (antheridia not seen). Setae erect, 0.35-0.5 mm long; capsules deeply immersed, 0.55-—0.8 mm long, subhemispheric, yellow-brown, strongly wrinkled (especially when dry); annulus none; operculum stoutly conic-rostrate from a low, convex base, with cells in straight vertical rows; exothecial cells thin-walled; stomata in a single row at the extreme base of the capsule; peristome none. Spores spherical or slightly ellipsoidal, 22-29 », brown, smooth. Calyptra cucullate, smooth, naked, covering only the operculum (figs. 1-9).
Holotype. Alkali flats along Panama Lane 5 miles W of Highway 99 just S of Bakersfield, Kern Co., California, Leo F. Koch 4026, January 24,1952 (MICH.)
The meager specimen at the University of Michigan was received from Tulane University, where Koch once held a teaching appointment.
92
1967] CRUM: PTERIGONEURUM 93
Fics. 1-9. Pterigoneurum californicum: 1, a rather young plant of bulbiform as- pect, X 29; 2, a fully mature plant with leaves somewhat spreading (wet or dry), xX 29; 3, three leaves, & 55; 4, cells at base of the awn, X 247; 5, cells somewhat below the awn, X 247; 6, lamella (at its upper end), x 247; 7, capsule, xX 29; 8, operculum, 29; 9, calyptra, X 29. I am indebted to Dale H. Vitt for prepara- tion of the drawings.
94 MADRONO [Vol. 19
It is presumed that duplicate specimens may be found there, under the name of P. subsessile, and also in Koch’s herbarium, which is deposited at the University of Illinois.
The distinctive features of this new species are the deeply immersed, strongly wrinkled capsules, completely lacking a peristome, smooth spores, cucullate calyptrae, and leaf cells without collenchymatous
corner thickenings. Herbarium, University of Michigan, Ann Arbor, Michigan
REVIEWS
The Flora and Fauna of Solano County. Edited by WILMERE JorDAN NEITZEL. viii + 160 pp., illustrated in color. Solano County Office of Education, Fairfield, California. 1965. $5.00.
The intent of this book is admirable; unfortunately its content leaves much to be desired. There are seven sections dealing with the wild flowers, trees, insects, fishes, amphibians and reptiles, birds, and mammals. Each kind of plant or animal is illustrated by a colored drawing; some bear only a vague resemblance to what they are supposed to represent. Inaccuracies appear in practically every one of the 151 accounts dealing with plants. These range from misspellings, such as “Eschscholtzia”’ (p. 28), to statements that Heteromeles arbutifolia has two ovaries (p. 30), to comments that Collinsia heterophylla has bulbs which “when boiled yielded a molasses-like syrup which was very popular with the early Indians of this county who used it as a syrup on meal” (p. 14), to the inclusion of Aconitum columbianum, a plant which does not occurs in Solano County (p. 7), to the manu- facture of such nonexistent names as “Dodecatheon majus” (p. 28). One can only wish, as Katharine Brandegee did (Zoe 1:224. 1890), “that future Boards of Education who would probably employ a carpenter if they wished to build a house, will show their common sense by employing a botanist when they want a local flora.”—JoHn H. THomas, Dudley Herbarium, Stanford University.
Atlas of North American Astragalus. By RUPERT C. BARNEBY. Memoirs of the New York Botanical Garden 13:lv + 1-1188, illustrated, in two parts. 1964. Cloth, $35.00; paperbound, $30.00. Available from Stechert-Hafner, 31 East 10th St., New York, New York 10003.
This monograph is the culmination of many years of work on a particularly difficult genus of higher plants. Many earlier papers on Astragalus by Mr. Barneby appeared under the title of “Pugillus Astragalorum,” indicating in an indirect way some of the difficulties encountered.
In all, 368 species of Astragalus are native in North America: only half a dozen have been introduced. Each species is fully described, synonymy is given, range is stated, good distribution maps are included, representative specimens are cited, and pertinent notes are presented. Illustrations of individual species are not included, but in a work of this length that is probably more than one can expect. Rather than prepare one very long key to all the taxa, Mr. Barneby has written 11 regional keys, a feature that local botanists will find most convenient.
This monograph is based on field and herbarium studies and as such indicates the kind of work that is still necessary in many, many groups before detailed biosystematic investigations can be carried out. Mr. Barneby is to be congratulated for his perseverance and patience in bringing forth this fine treatment—Joun H. Tuomas, Dudley Herbarium, Stanford University.
1967] NOTES AND NEWS 95
Systematic Embryology of the Angiosperms. By Gwenpa L. Davis. 528 pp. John Wiley and Sons, N.Y., London, Sydney, 1966. $19.75.
Professor Davis uses embryology in the broad sense of including structures in- volved in reproduction by seed, such as the entire ovule and anther, the male and female gametophytes, and the embryo and endosperm.
It is pointed out that “far from the descriptive phase of angiosperm embryology being over, it is hardly begun. . . . In the Compositae, for example, although there are over 300 publications, these concern species in only 15% of the genera... .” In nearly 20% of the families recognized by Hutchinson there could be traced no infor- mation on any embryological structure or process.
Embryological characters useful in systematics and taxonomy are described in the introduction. Considerably more attention is paid to the development of the anther than is usually found in embrvological discussions. In the treatment of the ovule, apomixis, adventive embryony, and polyembryony are included.
The second part of the book lists alphabetically the angiosperm families as clas- sified by Hutchinson (1959). Brief references to important embryological works are given for each family, together with a list of the authors. It is unfortunate that, for the larger families at least, the genera on which these authors worked are not men- tioned, so that a long search in the bibliography is required.
By far the greatest part of the book consists of a bibliography of some 5000 items dealing with embryological literature.
It is this last feature that makes the book so valuable, not only to plant embry- ologists and plant breeders, but also to all botanists who have occasion to know what information is available on the embryological aspects of a particular species.
The last comprehensive similar treatment of plant embryology was Schnarf’s “Vergleichende Embryologie der Angiospermen” published in 1931. The “System- atic Embryology of the Angiosperms” can now supplant this valuable reference work. Professor Davis is to be heartily congratulated by embryologists for stating the case for the use of embryology in taxonomy, and by all botanists in making available to them such a comprehensive embryological bibliography—Marron S. Cave, Department of Botany, University of California, Berkeley.
NOTES AND NEWS
TERRESTRIAL VASCULAR PLANTS OF ANO NuEvo I., SAN MaTEO Co., CALIFORNIA.— This small, 12 acre island (Orr, R. T., and T. C. Poulter, Pacific Discovery 15(1): 13-19. 1962) lies about one-fourth mile off the coast at Afo Nuevo Pt. about Lat N 37° 06.5’, Long 122° 20’ W. It is of interest for several reasons. An historic light- house was maintained on the island from 1873 to 1948. Afio Nuevo I. serves as a rookery for California sea lions, Steller sea lions, elephant seals, and harbor seals (Orr, R. T., and T. C. Poulter, Proc. Calif. Acad: 32:377-404. 1965) and studies in the population dynamics of a rabbit population introduced between 1948 and 1953 are being carried out by David C. Regnery of Stanford University. The island is now administered through the California Division of Beaches and Parks.
Because of the effect that the animal populations will have, and indeed have had, upon the vegetation, I think it worth while to publish a list of the plants I col- lected during three visits (May 20, 1963, July 19, 1963, and June 19, 1964) which were made possible through the courtesy of Thomas C. Poulter. With the exception of Rhus diversiloba and Mesembryanthemum chilense, which I did not collect, my specimens are in the Dudley Herbarium. Nomenclature follows that in my Flora of the Santa Cruz Mountains of California (Stanford Univ. Press, 1961) except for Onagraceae.
96 MADRONO [Vol. 19
List of plants: Achillea millefolium var. californica, Agrostis californica, Am- sinckia spectabilis, Anagallis arvensis, Atriplex californica, Bromus marginatus, B. maritimus, B. rigidus, Cakile maritima, Camissonia cheiranthifolia (Spreng.) Rai- mann, Cardamine oligosperma, Centaurea melitensis, Chenopodium californicum, Chlorogalum pomeridianum, Cirsium occidentale, C. vulgare, Cotula coronopifolia, Daucus pusillus, Distichlis spicata, Dudleya farinosa, Elymus mollis, E. triticoides Eriogonum latifolium, Eriophyllum staechadifolium, Eschscholzia californica, Fes- tuca dertonensis, Franseria chamissonis ssp. bipinnatisecta, Gnaphalium chilense, Hordeum brachyantherum, H. leporinum, Linaria texana, Lupinus arboreus, Malva parviflora, Marah fabaceus, Mesembryanthemum chilense, Microseris bigelovit, Monta perfoliata, Parapholis incurva, Plantago juncoides var. californica, Polypo- gon monspeliensis, Rhus diversiloba, Rubus ursinus, Rumex acetosella, R. conglom- eratus, R. crassus, Senecio vulgaris, Sagina occidentalis, Silene gallica, Solanum furcatum, Sonchus asper, S. oleraceus, Spergularia macrotheca, Stachys rigida ssp. quercetorum, Stellaria media, Trifolium gracilentum, T. microcephalum, and T. tridentatum.
A number of coastal strand plants occur on the adjacent mainland but are absent from Ano Nuevo I. Among them are: Abronia latifolia, A. umbellata, Cas- tilleja latifolia, Convolvulus soldanella, Fragaria chiloensis, Lathyrus littoralis, and Poa douglasiz. Coastal strand vegetation is fragile and does not survive repeated or continual disturbance. The pinnipeds move freely over much of the island and have perhaps either exterminated some strand plants or have prevented them from becoming established —Jonun H. Tuomas, Dudley Herbarium, Stanford University.
NEW PUBLICATIONS
Handbook of the Vascular Plants of the Northern Wasatch. By ArTHUR H. HOLMGREN. v + 202 pp. National Press, Palo Alto, California. Third Edition. 1965.
Processes of Organic Evolution. By G. LEDYARD STEBBINS. xiv + 191 pp. Pren- tice-Hall Inc., Engelwood Cliffs, New Jersey. 1966.
Trees of Northern Florida. By HERMAN Kurz and ROBERT K. GODFREY. xxxiv + 309 pp. University of Florida Press, Gainesville. 1962.
Flora of Southeastern Washington and of Adjacent Idaho. By Harotp St. Joun. xxix + 583 pp. Outdoor Pictures, Escondido, California. Third Edition. 1963.
Drawings of British Plants. By Stet~a Ross-Craic. Part XXII. Scrophulari- aceae (1). 39 plates. G. Bell & Sons, London. 1966. 11/6d.
A Checklist of the Vascular Plants of Tonto National Monument, Arizona. By Rospert L. Burcess. Journal of the Arizona Academy of Sciences 3:213-223. 1965.
A Glossary of Terms Frequently Used by Alpine Ecologists and Others. By James R. HaBeck and Ernest Hart Ley. 29 pp. Available from the authors, De- partment of Botany, University of Montana, Missoula. 1965.
A Flora of Wyoming. By C. L. Porter. Part III (Cyperaceae-Orchidaceae), 81 pp., Part IV (Poaceae), 88 pp. Bulletins 418 and 434, Agricultural Experiment Sta- tion, University of Wyoming, Laramie. 1964, 1965.
A Revision of the Genus Bowlesia Ruiz & Pav. (Umbelliferae-H ydrocotyloideae ) and Its Relatives. By MitpreD E. Matuias and LINCOLN CONSTANCE. University of California Publications in Botany 38:vi + 1-73. University of California Press, Berkeley and Los Angeles. 1965.
Geographical Distribution of the Pines of the World. By Wit1t1AM B. CritTcH- FIELD and Expert L. LitTLe, Jr. vi + 97 pp. Miscellaneous Publication 991, Forest Service, U.S. Department of Agriculture, Superintendent of Documents, U.S. Gov- ernment Printing Office, Washington, D.C. 20402. 1966.
Wildflowers of the Monterey Area, California. By BEatricE F. Howitt. 50 pp., illustrated with colored photographs. Wheelwright Press, 975 S. West Temple, Salt Lake City, Utah. Available from the Museum of Natural History, Forest & Central Avenues, Pacific Grove, California. 1965. This spiral bound booklet con- tains photographs and notes about 63 of the more common kinds of flowering plants of Monterey County.
MADRONO
VOLUME 19, NUMBER 4 OCTOBER, 1967
Contents
THE SCIENTIFIC WRITINGS OF WILLIS LINN JEPSON, Lawrence R. Heckard, John Thomas Howell, and Rimo Bacigalupi 97
ON THE HORDEUM JUBATUM — H. BRACHYANTHERUM QuEsTION, Wm. W. Mitchell 108
Notes ON MARINE ALGAE OF WASHINGTON AND SOUTH- ERN British Cotumpszia, II, Richard E. Norris and
John A. West 111 GOSSYPIUM TRILOBUM: AN ADDENDUM, Paul A. Fryxell
and Clifford R. Parks 117 LEE BONAR 124 A Hysrip HAWKWEED FROM THE OLtympPic Moun-
TAINS OF WASHINGTON, Arthur R. Kruckeberg 126 A SURVEY OF THE VASCULAR PLANTS ABOVE TIMBER-
LINE ON MrT. Hoop, OrEcOoN, Lurel Hall 130 DOCUMENTED CHROMOSOME NUMBERS OF PLANTS 134
VARIATION IN THE DiPLoip GOSSYPIUM SPECIES OF Baja CAirorniA, Lyle L. Phillips and Duncan Clement 137
CALIFORNIA BoLeETES III. THE GENus SuUILLus, Harry D. Thiers 148
Notes AND News: XI INTERNATIONAL BOTANICAL CONGRESS 136
A WEST AMERICAN JOURNAL OF BOTANY
PUBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
/ ;
en ad rae
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
Second-class postage paid at Berkeley, California. Return requested. Established 1916. Subscription price $6.00 per year ($4.00 for students). 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 un- delivered copies should be sent to the Corresponding Secretary, California Botanical Society, Department of Botany, University of California, Berkeley, California 94720.
BOARD OF EDITORS
LyMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. Ernst, Smithsonian Institution, Washington, D.C. ARTURO GOMEZ PompPa, Universidad Nacional Autonoma de México EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia MILpReED E. Matus, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley MARION OwNBEY, Washington State University, Pullman REED C. Ro.tiins, Harvard University, Cambridge, Massachusetts IrA L. Wiccrns, Stanford University, Stanford, California
Editor—JoHn H. THomAs Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—J UNE McCasKILL P.O. Box 23, Davis, California 95616
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Carl W. Sharsmith, Department of Biological Sciences, San Jose State College, San Jose. First Vice-President: Robert Ornduff, De- partment of Botany, University of California, Berkeley. Second Vice- President: Robert F. Thorne, Rancho Santa Ana Botanic Garden, Clare- mont. Recording Secretary: Mary L. Bowerman, Department of Botany, University of California, Berkeley. Corresponding Secretary: Mary E. Ashton, Department of Botany, University of California, Berkeley. Treasurer: June McCaskill, University of California, Davis.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Malcolm A. Nobs, Car- negie Institution of Washington, Stanford; the Editor of Madrofio; and three elected Council Members: Annetta Carter, Department of Botany, University of California, Berkeley; Virginia M. Page, Department of Biological Sciences, Stanford University; and Herbert L. Mason, De- partment of Botany, University of California, Berkeley.
THE SCIENTIFIC WRITINGS OF WILLIS LINN JEPSON (1867-1946)
LAWRENCE R. HECKARD, JOHN THOMAS HOWELL, and RIMO BACIGALUPI
This year marks the centenary of the birth of Willis Linn Jepson and it seems appropriate to commemorate this event by presenting a bibliog- raphy of the writings by and about this prolific botanist of California. Although Jepson’s publications have never been catalogued, their sig- nificance is fully recognized. Keck (1948) states that the “writings of Jepson are both extensive and profound, and they have exerted a lasting influence upon our knowledge of the botany of California.”’ Jepson’s botanical writings, spanning 55 years, were exceedingly diverse, ranging from comprehensive scientific volumes to articles in farm journals and newspapers which were aimed at increasing the awareness of the public in matters of conservation, horticulture, or botany. We have in- cluded all scientific and biographical writings traceable to Jepson’s authorship, many of which are on file in the Jepson Library in the Botany Department at the University of California, Berkeley. Non- botanical writings have not been included here but a listing of these has been placed in the Jepson archives. Other items not included here, such as newspaper articles concerning Jepson or his public addresses, are also in the Jepson archives.
It is timely to announce the availability for scholarly study at Berke- ley of the Jepson Correspondence, recently bound in 62 volumes. The letters, covering the period 1887-1946, were arranged and indexed an- nually under Jepson’s supervision. Entries in the indices include per- sonal names, plant names (both scientific and common), place names, and various other categories. Such correspondents are included as: LeRoy Abrams, L. H. Bailey, Katharine Brandegee, Luther Burbank, F. V. Coville, Alice Eastwood, A. Engler, E. L. Greene, John Muir, B. L. Robinson, C. S. Sargent, J. K. Small, as well as many other contempo- raries of Jepson.
We wish to thank Joseph and Nesta Ewan and Helen-Mar Beard for their helpful suggestions and for supplying entries for the bibliography.
OBITUARIES AND HISTORICAL APPRAISALS OF WILLIS LINN JEPSON
1928. Carew, H. D. The high priest of flora; a glance at the life and works of California’s foremost botanist. Touring Topics 20(12): 32-34, portrait. [Automobile Club of Southern Calif., Los Angeles].
1933. ANon. The work of Professor Willis Linn Jepson. Science 78:597— 598.
1938. [Portrait of Jepson. Frontispiece to Madrono Vol. 4].
Maprono, Vol. 19, No. 4, pp. 97-160. December 15, 1967. 97
98 MADRONO [Vol. 19
1946. Davis, A. R., L. CoNsTANnceE, G. D. LOUDERBACK. In Memoriam, University of California, Berkeley.
1947. CONSTANCE, L. Willis Linn Jepson. Science 195:614.
1947. Fritz, E. Willis Linn Jepson, a eulogy and a bit of California for- estry history, Calif. Forester 14:6—8, portrait.
1947. Ewan, J. | Willis Linn Jepson]|. Jour. Wash. Acad. 37:414—416.
1947. Mason, H. L. Willis Linn Jepson. Madrono 9:61—64.
1947. Parsons, M. R. Willis Linn Jepson. Sierra Club Bull. 32: 104-107.
1947. ANON. Willis Linn Jepson. Des. Pl. Life 19:43.
1947. WHEELER, H. W. Writings of W. L. Jepson. Des. Pl. Life 19:44— 45.
1947. ANon. Dr. W. L. Jepson. Forestry News 2:7.
1948. Fritz, E. Willis Linn Jepson, lover of California’s trees and flowers. Jour. Calif. Hort. Soc. 9:23-26.
1948. Keck, D. D. The place of Willis Linn Jepson in California botany. Madrono 9:223-228.
1950. Constance, L. Disposition of the Willis Linn Jepson estate. Ma- drono 10:128.
MEMORIALS TO WILLIS LINN JEPSON 1923. The Jepson Laurel [Umbellularia californica|. San Francisco Water 2:2-6. 1951. Memorial to Willis Linn Jepson [at Tomales Bay State Park]. Sierra Club Bull. 36:7. 1957. Vacaville school named in honor of Willis Linn Jepson, famed ‘botanist. Vacaville Reporter, Oct. 18.
I. Books By WILLIS LINN JEPSON
1901. A Flora of Western Middle California. Encina Publishing Co., Berkeley, 625 pp. Second Edition, Cunningham, Curtiss & Welch, San Francisco, 515 pp., 1911.
1902. A School Flora for the Pacific Coast. D. Appleton & Co., New York, 96 pp.
1909. The Trees of California. Cunningham, Curtis & Welch, San Fran- cisco, 228 pp., 117 figs. Second Edition, Assoc. Students Store, Uni- versity of California, Berkeley, 240 pp., 124 figs., 1923.
1909-43. A Flora of California [incomplete]. 1909: Vol. 1, Part I, pp. 33-64: Part II, pp. 337-368. 1912: Part III, pp. 65-192. 1914: Part IV, pp. 369-464; Part V, pp. 465-528. 1922: Part VI, pp. 193-336; Part VII, pp. 529-578. 1936: Vol. 2, Part I, pp. 1-16, 337-684; Part II, pp. 17-176. Part ITI, pp. 177-336. 1939: Vol. 3, Part I, pp. 17-128. 1943: Part II, pp. 129-464. [Tllus. with 451 figs. ].
1910. The Silva of California; Univ. Calif. Mem. 2:1—480, 85 pls., 10 text-figs., 3 maps. Berkeley.
1923-25. A Manual of the Flowering Plants of California, pp. 33-384 (1923); pp. 1-32, 385-1238 (1925), 1023 figs., 1 map. Assoc. Stu-
1967 | HECKARD, HOWELL, & BACIGALUPI: JEPSON 99
dents Store, University of California, Berkeley. Second Printing, 1938. Third and subsequent printings, Univ. Calif. Press, Berkeley, 1951 et seq.
1924. A Flora of the Economic Plants of California for Agricultural Stu- dents. Assoc. Students Store, University of California, Berkeley, 223 pp., 16 figs.
1935. A High School Flora for California. Associated Students Store, University of California, Berkeley [a verbatim reprinting of the 1924 Flora of the Economic Plants of California for Agricultural Students |.
II. BOTANICAL ARTICLES AND BIOGRAPHIES BY WILLIS LINN JEPSON (with indices to taxa and to biographies, see pg. 107)
1891. Botany of the Marysville Buttes. Bull. Torrey Club 18:317-327.
1892. New Californian atriplices. Pittonia 2: 303-306.
1893. a. Studies in the Californian Umbelliferae. I, Erythea 1:8—10. IT, 62-63.
b. The mountain region of Clear Lake. Erythea 1: 10-16.
c. [Review of] Sudworth, G. B., On legitimate authorship of certain binomials, with other notes on nomenclature. Erythea 1:123-124.
d. On a variety of the western sumach | Rhus trilobata Nutt. var. quinata|. Erythea 1:140-141.
e. Alien plants in California. Erythea 1:141-143.
f. Early scientific expeditions to California. I. [La Pérouse Ex- pedition|]. Erythea 1:185-190.
g. The riparian botany of the lower Sacramento. Erythea 1:238- 246.
1894. a. Teratological notes. Erythea 2:14—-15. [Dendromecon, Esch- scholzia, Boisduvalia |.
b. Professor Greene’s mission in Europe. Erythea 2:153-155.
c. Odontostomum hartwegt. Erythea 2(9):157-158.
d. Chamisso Botanical Club. Erythea 2:171-172.
1895. |Note on|Arctostaphylos elegans Jepson. Erythea 3:178. 1896. a. The chaparral of the Coast Range. Univ. Calif. Mag. 2:98— 105, 1 pl. (frontispiece).
b. [Review of] Gray, A., S. Watson, B. L. Robinson, Synoptical Flora of North America, Vol. I, Part I, Fasc. I. Erythea 4: 16— 19,
Distribution of the coast redwood. Erythea 4:125.
d. [Review of] Bailey, L. H., Plant Breeding, Erythea 4:143-— 144.
e. [Review of] Rusby, H. H., and S. E. Jeliffe, Essentials of Vegeta- ble Pharmacognosy. Erythea 4:144—145.
1897. a. A new west American Peucedanum. Erythea 5:1. b. Botanical exploration in California in 1896. Erythea 5: 7-9.
@
100
1898.
1899.
MADRONO [Vol. 19
. The new series of Proceedings of the California Academy of
Sciences. Erythea 5:25-26.
d. [Note on occurrence of | Velezia rigida L. Erythea 5:28-—29.
Ean er OQ rh
ed .
. The explorations of Hartweg in America. Erythea 5:31-35; 51-
56.
. Boschniakia strobilacea Gray. Erythea 5:63—66, 2 pls. . [Review of] Howell, T., A Flora of Northwest America.
Erythea 5:66-68.
. Biographical sketch of Dr. W. P. Gibbons. Erythea 5: 74-76.
[Notice of] Rose, J. N., The North American species of Chryso- splenium. Erythea 5:99.
[ Review of] Gray, A., The Synoptical Flora of North America, Vol. I, Part 2, Fasc. 2[B. L. Robinson, Ed.|. Erythea 5:106- 108.
. Carpenteria californica. Erythea 5:124. . Seed plants of western middle California. Key to the orders and
anomalous genera. 10 pp. The University Press, Berkeley.
b. [Note on locality of] Picea breweriana. Erythea 6:12.
. [Review of] Bergen, J. Y., Elements of Botany. Erythea 6:16. . [Review of] Rattan, V., Exercises in Botany for the Pacific
States. Erythea 6:16-17.
. [Editorial comment on] Platt, R. H., Fritillaria pluriflora in
Solano County. Erythea 6:27.
. Baron Ferdinand von Mueller. Erythea 6:32—35. [Abstract
from Victorian Naturalist with editorial comment |.
. [Editorial comment on] Eastwood, A., Is Xerophyllum tenax
a septennial? Erythea 6:75—76.
. Beckwithia, a new genus of Ranunculaceae. Erythea 6:97—99,
1 pl.
Dr. Henry N. Bolander, botanical explorer. Erythea 6:100- 107, 1 portrait.
[Review of] Bailey, L. H., Lessons with Plants. Erythea 6: 107-108.
. [Review of] Barnes, C. R., Plant Life, considered with special
reference to form and function. Erythea 6:113. [Review of] Jones, M. E., Contributions to western botany, No. 8, Erythea 6:119-120.
. Vegetation of the summit of Mt. St. Helena. Erythea 7:105-
113, 1 pl.
. Teratology of Scrophularia californica. Erythea 7:123-127, 1
pl.
. Early scientific expeditions to California. II. [Malaspina Ex-
pedition |]. Erythea 7: 129-134.
. [Review of] Campbell, D. H., The Evolution of Plants.
Erythea 7:147-148.
. [Review of] Bessey, C. E., The phylogeny and taxonomy of
1967 |
1903.
1904.
1905.
1906. 1907.
1908.
1909.
b.
GC:
HECKARD, HOWELL, & BACIGALUPI: JEPSON 101
the Angiosperms. Erythea 7: 148-149. Biographical sketch of H. G. Bloomer. Erythea 7: 163-166, 1 portrait.
. Epitaph of David Douglas. Erythea 7:174—-175. . [Review of] Davidson, A. M., California Plants in their
Homes. Erythea 7:176—-177.
[Review of] Coulter, J. M., Plant Relations. Erythea 7:177- 178.
[Review of| Bailey, L. H., Sketch of the Evolution of our Na- tive Fruits. Erythea 7:178—-179.
. The live oaks of the University of California campus. Univer-
sity Chronicle 6:179-185.
Mount Whitney, Whitney Creek, and the Poison Meadow trail. Sierra Club Bull. 4:207—215, 2 pls.
[Letter] Largest bay laurel [Umbellularia|. The Reveille, Dec. 26 [Cloverdale, Calif. newspaper |.
Suggestions in city forestry. Oakland Enquirer 44:18, Jan. 2
a.
b.
C.
[Oakland, Calif. newspaper |.
Street trees in California. Hand Book series, The California Promotion Committee, San Francisco, 16 pp.
International horticultural exhibition at Edinburgh, Scotland. Pacif. Rural Press 70:228.
Where ducks dine. Sunset Mag. 14:409-411.
Bret Harte and the Argonauts. Sunset Mag. 18:85.
a. b.
a.
The deer oak [Quercus sadleriana|. Muhlenbergia 3:130. The tan oak of the Pacific coast |Lithocarpus densiflorus |. Jour. Am. Leather Chem. Assn. 2:197—201.
. A synopsis of the North American godetias. Univ. Calif. Publ.
Bot. 2:319-354, 1 pl.
. A world of timber. Sunset Mag. 20:97. . The Washington palms |Washingtonia|. Muhlenbergia 4:41. . The distribution of Juglans californica Wats. Bull. So. Calif.
Acad. 7:23-24.
. The name of white sage [Salvia apiana|. Muhlenbergia 3:144. . Northernmost stations for two common Californian trees | Popu-
lus fremontu, Platanus racemosa|. Muhlenbergia 3: 144.
. Gardens and their kinds, [a review of] Robinson, W., The Gar-
den Beautiful. Pacif. Rural Press 75:70. | Letter on] The California walnut. Pacif. Rural Press 75:357.
. The California balloon plant [Astragalus]. Pacif. Rural Press
156 56:
. [Letter on] Big tree and redwood. San Francisco Call 103:27,
Jan. 26 [newspaper |. Spontaneous hybrids of native Californian trees [in Cupressus, Juglans, Quercus, Salix|. Am. Breeder’s Assn. Rep. 5:259-261.
. Forestral barrenness in the California foothills. Trans. Com-
102
1910.
1911.
1912.
1913.
1914.
1915.
MADRONO [Vol. 19
monwealth Club Calif. 4:50. Reprinted in Pacif. Rural Press 77:244—-245.
. [Remarks on forest fire laws in California]. Trans. Common-
wealth Club Calif. 4: 75-77.
d. Singular Californian oaks. Pacif. Rural Press 77:9.
. Plan for highway trees for California. Pacif. Rural Press 77:
30S: The acorn crop of 1908 and the pine seed crop of 1909. Pacif. Rural Press 78:309.
. Suggestions for tree planting. Pacif. Rural Press 78:408. . Engelmann spruce near the California line. Muhlenbergia 5:64.
Mule ears |W yethia|. Muhlenbergia 5:153. Hard and soft woods. Pacif. Rural Press 79:244.
. Ornamental trees commended. Pacif. Rural Press 79:476. alifornia tanbark oak | Lithocarpus densiflorus|. 1. Tanbark oak
and the tanning industry. U.S. Dep. Agr. Forest Serv. Bull. 75:5-23, 5 pls.
. How tall and how old are the great trees. Pacif. Rural Press
84:660—-661.
. The steer’s head flower of the Sierra Nevada [Dicentra uni-
flora|. Sierra Club Bull. 8:266—269, 2 figs.
. Trees, shrubs, and flowers of Napa County. /n Gregory, T., et
al., History of Solano and Napa Counties. Historic Record Co., Los Angeles, pp. 150-153.
d. The native vegetation of Solano County. /bid., pp. 136-139.
a.
. Freeman Grove of big trees. Porterville Recorder, Aug. 3. | Por-
terville, Calif. newspaper |. A lightning struck tree in Kern Canyon. Sierra Club Bull. 9: 25-27, 2 pls.
. Mountain misery [Chamaebatia foliolosa|. Sierra Club Bull.
9:40-42, 2 figs.
. [Comment on] The Ballota oak [Quercus ilex var. ballota|.
Pacif. Rural Press 85:197.
d. The oldest big tree. Pacif. Rural Press 85:259.
a.
. Excusion program of the fourth section. International Phyto-
geographic Excursion (I.P.E.) in America, pp. 3-15. First across the Sierra Nevada | Joseph Reddeford Walker]. Sierra Club Bull. 9:165—-169.
. Madrona | Arbutus menziesii|. Tamalpais Mag. 3(6):5. |Tam-
alpais Conservation Club, Marin Co., Calif. ].
. The stump-sprout habit of the redwoods. Tamalpais Mag.
4(2):5. [not seen].
d. The early history of the redwood. Tamalpais Mag. 4(2):7.
a.
. When fir is pine |Pseudotsuga menziesii|. Pacif. Rural Press
87:425. [ Remarks on] Common names of native flowers in the National
1967]
1916.
1917.
1918.
1919.
1920.
eu ieu, (@y lor
QA0
HECKARD, HOWELL, & BACIGALUPI: JEPSON 103
Parks. Proc. |Third] National Park Conference, Washington, D.C., pp. 126-128.
. Godetia. In Bailey, L. H., Standard Cyclopedia of Horticulture
3:1353-1354.
. [Note on] Principal trees between the summit of the Sierras
and San Francisco Bay. Jn Guidebook of the Western United States. U.S. Geol. Survey Bull. 612, pp. 202-203.
. Forests of the western United States. Jn Grinnell, J. (Ed.), Na-
ture and Science on the Pacific Coast. Paul Elder and Co., San Francisco, pp. 159-167, 2 pls.
. Regeneration in manzanita. Madrono 1:3-11, 2 pls., 5 figs. . The International Phytogeographic Excursion in California.
Madrono 1:13-14, 1 pl.
. Phellopterus littoralis Schmidt. Madrono 1:27. . The Mohave Desert. Gardening Illustrated, June 9, p. 308. (A
nearly verbatim reprinting of 1917b).
. Springtime in the desert. Univ. Calif. Jour. Agr. 4:214-215.
. The native walnuts of California. Madrono 1:55—57, 2 figs.
. Taxonomic notes on California plants. Madrono 1:61-62.
. Protection and preservation of California flora. Jn Kennedy,
P. B., Annotated List of the Wild Flowers of California. Third Annual Calif. Wild Flower Fete (privately printed), San Fran- cisco, Calif., p. 11.
. Edward Lee Greene, the man and botanist. Newman Hall Rev.
1:24—29, portrait.
. The White Mountains of California. Sierra Club Bull. 10:282-—
286, 2 pls.
. Collection and preparation of specimens of flowering plants.
Nemophila, {Meeting and Field Guide, Calif. Bot. Soc.| 3:11- We
. The botanical range of Christmas berry |Heteromeles arbuti-
folia|. Nemophila 4: 15-16.
. Trees for the Sacramento Valley. Pacif. Rural Press 97:146. . Selection of memorial trees. Pacif. Rural Press 98:713. . Save the redwoods. Northern Crown 7:1 | Published at Ukiah,
Calif. ].
. Plea for preservation of redwoods. San Francisco Chronicle
106:14, Jul. 15 [newspaper |.
. The vitality of redwood. Nemophila 7:27. . Quercus agrifolia on Warner Ranch. Nemophila 9:35. . Yellow skunk cabbage [Lysichiton| in the Santa Cruz Moun-
tains. Nemophila 9:35.
. Rare or little known species in need of collection. Nemophila
9:36.
. Water hyacinth | Eichhornia crassipes| in California. Nemo-
phila 11:43.
104
1922—23.
1923
1923-29:
1924.
1926 1927
1928.
MADRONO [Vol. 19
. Pleuricospora fimbriolata in the Santa Cruz Mts. Nemophila
11:43-44. Variation in Hemitomes congestum. Nemophila 11:44. The Ayars oak [Quercus lobata|. Nemophila 13:52.
. The long-lost Carpenteria. Sierra Club Bull. 11:151—-153, 2 pls. . The fire-type forest of the Sierra Nevada. Intercollegiate For-
estry Club Annual 1:7—10.
. The giant Sequoia. Jn Hall, A. F. [Ed.], Yosemite National
Park Handbook, G. P. Putnam’s Sons, New York, pp. 235-— 246, 2 pls.
. The flowers of Yosemite National Park. Jbid. pp. 247-267, 4
pls.
. The Hudson oak [Quercus chrysolepis|. Republican Press,
June 15 | Ukiah, Calif. newspaper |. Historical notes on Carex collecting in California. Erythea 8: 5-6.
b. A new species of cypress [Cupressus forbesit|. Madrono 1:75.
. a.
b.
Q
a.
b.
. Notes and news. [Visit of J. H. Patton, Cornell botanical ex-
pedition; visit of Olof Arrhenius]. Madrono 1:75—76.
. The forests of California. San Francisco Chronicle 120: p. 12E,
June 12 [newspaper |.
Revision of the California species of the genus Arctostaphylos. Madrono 1:76—86 (1922); 87-96, 3 figs. (1923).
Revision of the California species of the genus, Downingia Torr. Madrono 1:98—102, 4 figs.
[Comment on] Quercus morehus in the San Bernardino Mts. Madrono 1:98.
. [Comment on] Cercis occidentalis. Madrono 1:99. . A new species of Evriogonum |bidentatum| from California.
Madrono 1:115-116.
. A conifer new to California | Picea engelmanniu|. Madrono 1:
116. |Comment on| Physalis ixocarpa Brot. Madrofo 1:116.
. The sequoias of California, their life history and geographic
distribution. Proc. [second] Pan-Pacific Sci. Congress 1:307- SZ,
A revision of Californian Umbelliferae. I-III, Madrono 1:101- 114; 117-130; 133-146 (1923). IV, 1:149-162 (1924). V, 1: 281-285 (1929). 38 figs.
Specific status of Claytonia nevadensis Wats. Madrofto 1:147- 148.
{Comment on] Kumlienia hystricula Greene. Madrono 1:147.
. [Letter on] The Lebanon cedars in Syria. Calif. Cultivator 67:31. . The unique cypress trees of Cedar Mt. The Livermore Herald,
a.
Feb. 4, 1 photo. [Livermore, Calif. newspaper |. The botanical explorers of California I, Volney Rattan. Ma-
1967]
1929.
1930.
OST.
HECKARD, HOWELL, & BACIGALUPI: JEPSON 105
drono 1:167—170, portrait. I1, Joseph Whipple Congdon. Ma- drono 1:175-177, portrait. III, George Hansen. Madrono 1: 183-185, portrait. [V, George D. Butler. Madrono 1:188—190, portrait.
. The annual dinner for 1927. [California Botanical Society].
Madrono 1:177.
. Biological peculiarities of Californian flowering plants. Longi-
tudinal fission in the plant body of Ceanothus cuneatus Nutt. Madrono 1: 190-192, 2 figs.
d. California Park Commission. Madrono 1:192.
eh
AS = Eo
1.
. [Notice of] Grant, A. L., A monograph of the genus Mimulus.
Madrono 1:194. Samuel Bonsall Parish. Proc. Twenty-Third Annual Meeting, Bot. Soc. Am. Publ. 99: 26-28.
. Samuel Bonsall Parish. Science 69:63. . Physiological bilabiation in the flowers of Californian angio-
sperms [in Clarkia pulchella|. Madrono 1:198-199.
. Variation in the pappus of Layia pentachaeta Gray. Madrono
1: 200-201.
. [Note on] The exploitation of Darlingtonia californica. Ma-
drono 1:201.
. The botanical explorers of California —V, Charles Lewis An-
derson. Madrono 1:214—216, portrait. VI, Archibald Menzies, Daniel Cleveland, Harley Pierce Chandler. Madrono 1:262-270, 3 portraits.
[Note on] Pepoon, H. S., Annotated Flora of the Chicago Area. Madrono 1:226.
. Sequoia sempervirens in Granada. Madrono 1:242. . Johann Friederich Eschscholtz. Madronfo 1:253.
Sertulum Californiense. I. Madrono 1:254—260.
. Harriet A. Walker. Madrono 1:261. . The annual dinner for 1928 and for 1929 [California Botani-
cal Society]. Madrofio 1:270—273. Charles Russell Orcutt, natural history collector. Madrono 1: 273-274.
m. A new Erythronium. Calif. Cultivator 72:475. [Notes on the plant life in Death Valley] in Chalfant, W. A.,
a.
C.
Death Valley—the Facts, pp. 85-87; Ed. 3, pp. 89-91, Stan- ford University Press.
Patrick Beveridge Kennedy. Proc. Twenty-fifth Annual Meet- ing, Bot. Soc. Am. Publ. 105: 19-20.
. The botanical explorers of California —VII, Joseph Cook
Nevin, William Hillman Shockley, Charles Austin Stivers, Lucia A. Summers. Madrono 2:25-29, 3 portraits [Summers lacking]. [Comment on] Cupressus macnabiana and C. bakeri. Madrono Leo le
106
1932.
ISS
1934.
MADRONO [Vol. 19
d. Bolander’s Mendocino and Humboldt trips. Madrono 2:33—-34.
-" a IQ mH
. Biographical sketch of Patrick Beveridge Kennedy. Madrono
2:34-35, portrait. [Comment on] Joshua forests about Cima. Madrofo 2:43.
. Wilderness areas in California. Madrono 2:45—47. . Edward Lee Greene. Dict. Am. Biog. 7:564—565.
The role of fire in relation to the differentiation of species in the chaparral. Proc. Fifth International Bot. Congress, pp. 114-116.
. Weed control in California. Madrono 1:57—59.
b. Death of Dr. Harvey Monroe Hall. Madrono 2:63.
fay)
mo O&O oP Bog
po
ao
. Rainfall predictions for California, season of 1931-32. Ma-
drono 2: 71-72.
. Note on the genus Kvameria in California. Madrono 2:73. . Samuel Bonsall Parish. Univ. Calif. Publ. Bot. 16:427-444,
portrait. George Hansen. Dict. Am. Biog. 8:229-—230.
. Charles Frederick Holder. Dict. Am. Biog. 9: 140. . Thomas Jefferson Howell. Dict. Am. Biog. 9:305—306.
Albert Kellogg. Dict. Am. Biog. 10: 300-301.
. Frank Hall Knowlton. Dict. Am. Biog. 10:472—473.
Adam Kuhn. Dict. Am. Biog. 10:510—511.
. Richard Ernest Kunze. Dict. Am. Biog. 10:513-514. . John Gill Lemmon. Dict. Am. Biog. 11:162-163.
The botanical explorers of California. —VIII, William G. W. Harford, Thomas Bridges. Madrono 2:83—88, 1 portrait [of Harford |.
. David Douglas in California. Madrono 2:97—100. . In memoriam [A. Scouvert, C. A. Hunt, H. M. Hall, A. Head].
Madrono 2: 102-103.
The Monterey cypress grove of Point Lobos. Madrono 2:108. Scientific and vernacular names for the species of Sequoza. Sierra Club Bull. 18:116-118.
. The cypresses of Monterey. Sierra Club Bull. 18:64—67, 2 pls.
Phytogeography of the Coniferae of western North America. Proc. Fifth Pacif. Sci. Cong. 4: 3255-3264.
. The botanical explorers of California —IX, Charles Frederick
Sonne, Nancy Jane Davis, John Boardman Trask. Madrofto 2:115-118, 1 portrait [of Sonne]. X, Rachel [i.e. Rebecca] Merritt Austin, Lorenzo Dow Jared. Madrono 2:130-133, 2 portraits. XI, George W. Dunn, Rebecca Merritt Austin [name correction |. Madrono 2:156—-157, 1 portrait [of Dunn].
. The overland journey of Thomas Nuttall. Madrono 2:143-147,
portrait as frontispiece to Vol. 2 of Madrono.
. Marcus Eugene Jones. Madrofio 2:152-154. . [Comment on] Pinus insignis and Pinus radiata. Madrono 2:
159.
1967] HECKARD, HOWELL, & BACIGALUPI: JEPSON 107
f. Charles Christopher Parry. Dict. Am. Biog. 14:261-262.
1935. a. Trees, shrubs, and flowers of the redwood region. Save the Red- woods League, 15 pp., illust. Revised, 1965.
b. Centers of plant endemism in California in relation to geolog- ical history. Proc. Sixth Internat. Bot. Congress 2:82—83. [ Ab- stract |.
1936. a. [Review of] Parish, S. B., Cactaceae of California. Cact. Succ. Jour. 8:93. b. Edward James Wickson. Dict. Am. Biog. 20:184. 1938. a. Viae felicitatis: the beginning years of the California Botanical Society. Madrono 4:276—286. b. Embryonic panicles in Arctostaphylos. Erythea 8:97-99. Three new Californian Arctostaphyli Erythea 8:99-102.
d. California botanical explorers —XII, John Milton Bigelow. Erythea 8:102—104 [incomplete|. Reprinted with conclusion (Madrono 16:179-183. 1962).
1941. The joy of days afield in the redwood belt with its native flowers and shrubs. Twelfth Annual Calif. Spring Garden Show Year- book, pp. 27-28.
1942. a. Early botanical ascents of Mount Shasta. Sierra Club Bull. 27: 23-30.
b. The Smyrna fig in California. Science 95:19.
1943. a. Edward Lee Greene the individual. Am. Midl. Nat. 30:3—5.
b. [Remarks on] the names of the giant sequoia. Leafl. West. Bot. 3:214.
1944. Common names of plants. Word Study 19:6. (G. & C. Merriam Co., Springfield, Mass. )
1946. Concerning a coastal Habenaria in California. Leafl. West. Bot. 4:260.
1947. Carl Purdy, lover of the lilies. Jour. Calif. Hort. Soc. 8:2—4.
1962. California botanical explorers —XII, John Milton Bigelow. A re- printing of 1938d with concluding portion. Madrono 16:179— 133). >”
©
Index to taxa: -
Arbutus 1914b. Arctostaphylos 1895, 1916a, 1922-23, 1938b,c. Astragalus 1908g. Atriplex 1892. Beckwithia 1898h. Boisduvalia 1894a. Boschniakia 1897{. Cactaceae 1936a. Carex 1922a. Carpenteria 1897k, 1921b. Ceanothus 1928c. Cedrus 1926. Cercis 1923c. Chamaebatia 1913b. Chrysosplenium 18971. Clarkia 1929b. Claytonia 1924a. Coniferae 1934a. Cupressus 1909a, 1922b, 1927, 1931c, 1933i,k. Darlingtonia 1929d. Dendromecon 1894a. Dicentra 1912b. Downingia 1923a. Eichhornia 1920g. Eriog- onum 1923d. Erythronium 1929m. Eschscholzia 1894a. Ficus 1942b. Fritillaria 1898e. Godetia 1907c, 1915b. Habenaria 1946. Hemitomes 1920i. Heteromeles 1919b. Juglans 1908b,f, 1909a, 1917c. Krameria 1932d. Kumlienia 1924b. Layia 1929c. Lithocarpus 1907b, 1911. Lysichiton 1920e. Mimulus 1928e. Odontostomum 1894c. Peucedanum 1897a. Phellopterus 1916c. Physalis 1923f. Picea 1898b, 1909h, 1923e. Pinus 1909f, 1934e. Platanus 1908d. Pleuricospora 1920h. Populus 1908d. Pseudotsuga 1914e.
108 MADRONO [Vol. 19
Quercus 1903a, 1907a, 1909a,d,f, 1913c, 1920d, 1921a,f, 1923b. Rhus 1893d. Salix 1909a. Salvia 1908c. Scrophularia 1899b. Sequoia (including Sequoiadendron) 1896c, 1908h, 1912a,e, 1913d, 1914c,d, 1920a,b,c, 1921d, 1923g, 1929g, 1933j, 1935a, 1941, 1943b. Umbelliferae 1893a, 1923-29. Umbellularia 1903c. Velezia 1897d. Washingtonia 1908a. Wyethia 19091. Xerophyllum 1898g. Yucca 1931f.
Index to the biographical writings
Anderson, C. L. 1929e. Austin, R. M. 1934b. Bigelow, J. M. 1938d, 1962. Bloomer, H. G. 1899f. Bolander, H. N. 1898i, 1931d. Bridges, T. 1933f. Butler, G. D. 1928a. Chandler, H. 1929e. Cleveland, D. 1929e. Collignon, J. N. 1893f. Congdon, J. W. 1928a. Davis, N. J. 1934b. Douglas, D. 1899g, 1933g. Dunn, G. W. 1934b. Esch- scholtz, J. F. 1929h. Gibbons, W. P. 1897h. Greene, E. L. 1894b, 1918a, 1931h, 1943a. Haenke, T. 1899c. Hall, H. M. 1932b, 1933h. Hansen, G. 1928a, 1932f. Harford, W. G. W. 1933f. Harte, B. 1906. Hartweg, T. 1897e. Head, A. 1933h. Holder, C. F. 1932g. Howell, T. J. 1932h. Hunt, C. A. 1933h. Jared, L. D. 1934b. Jones, M. E. 1934d. Kellogg, A. 1933a. Kennedy, P. B. 1931a,e. Knowlton, F. H. 1933b. Kuhn, A. 1933c. Kunze, R. E. 1933d. La Pérouse, J. F. G. 1893f. Lemmon, J. G. 1933e. Mala- spina, A. 1899c. De la Martiniére 1893f. Menzies, A. 1929e. Née, L. 1899c. Nevin, J. C. 1931b. Nuttall, T. 1934c. Orcutt, C. R. 1929]. Parish, S. B. 1928f, 1929a, 1932e. Parry, C. C. 1934f. Purdy, C. 1947. Rattan, V. 1928a. Scouvert, A. 1933h. Shockley, W.H. 1931b. Sonne, C. F. 1934b. Stivers, C. A. 1931b. Summers, L. A. 1931b. Trask, J. B. 1934b. von Mueller, (B.) F. 1898f. Walker, Harriet 1929}. Walker, J. R. 1914a. Wickson, E. J. 1936b.
Jepson Herbarium, University of California, Berkeley
California Academy of Sciences, San Francisco
ON THE HORDEUM JUBATUM—H. BRACHYANTHERUM QUESTION
Wo. W. MITCHELL
Rajhathy (1966) disagrees with the separate species status recom- mended by Mitchell and Wilton (1964) for Hordeum jubatum L. and H. brachyantherum Nevski. He accepts proof of interbreeding as the overriding criterion for making species determinations; he considers that the above disposition ignores a ‘‘mass of evidence” obtained on the interbreeding of these taxa in the Canadian and American West and Midwest. Rajhathy also asserts that the Alaskan work is further inval- idated by being based on a “specialized peripheral situation.”
Alaska is on the northwestern periphery of the North American range
1967 | MITCHELL: HORDEUM 109
of H. jubatum. The species is nonetheless well adapted to subarctic conditions, occurring frequently and abundantly on disturbances throughout the Interior south of the Brooks Range and in the south- central region transitional to the coast. The propensity of jubatum for hybridization appears not in the least affected by Alaskan conditions since it hybridizes frequently with various species (Mitchell and Hodg- son, 1965). Alaska hardly can be considered peripheral with regard to the range and adaptability of H. brachyantherum. The Alaskan occur- rences of brachyantherum include much of _ its principal range along the coastal arc of the Pacific Ocean. The species frequents eastern Asia, is common through the Aleutian Islands and along the coast of Alaska to the panhandle, and extends southward to California. It occurs in well- developed beach meadows, invades disturbed ground, and is a very abundant weed in many Alaskan coastal communities. The species ranges inland along the highway and railway systems of southcentral Alaska. Hordeum jubatum and H. brachyantherum meet and hybridize in this southcentral region.
Bowden (1962) examined about 160 specimens of brachyantherum in the study which led to his combining jubatum and brachyantherum. Of a total of 82 accessions listed, 50 originated in British Columbia and Alaska, 8 in Alberta, and 2 in Saskatchewan. Moss (1959) stated brachyantherum to be “very rare” in Alberta, a plant of saline situa- tions. It is absent in Manitoba (Scoggan 1957) and in areas of Sas- katchewan and neighboring United States where the hybrid occurs in abundance (Bowden, 1962; Rajhathy, 1966). Obviously brachyan- therum prefers the summer-cool and moist coastal region of the North.
Apparently the hybrid is most successful in areas where bracyhan- therum is rare or absent and preserves its identity after migration from its site of origin. In controlled crosses (Rajhathy and Morrison, 1959) even the F, plants maintained this identity, remaining ‘“‘indistinguish- able from the F, plants,” although the F; and F, plants segregated for awn length (Rajhathy, 1966). Bowden (1962) reported little segrega- tion in Fy generations that he examined. The hybrid has been recog- nized by many taxonomists as a variety of jubatum (Covas, 1949; Hitchcock, 1950; Scoggins, 1957; Love, 1959).
In all probability jubatum and brachyantherum have had a common ancestry (Rajhathy and Morrison, 1959; 1961). But the large amount of morphologic and chorologic differentiation now manifest between the two reflects a successful divergence and reproductive separation effec- tive over a long period of time. The fact that the two produce hybrids on disturbances today is not unusual since jubatum hybridizes with numerous species in three or four genera. The production of fertile hybrids is of significance, but it occurs principally in an area that must be considered marginal for brachyantherum. Moreover, the effects on the integrity of the brachyantherum populations prevalent in this area cannot be ascertained from the data now available.
110 MADRONO [Vol. 19
The increase of the hybrid in the areas where brachyantherum is absent must depend on the compatibility of the hybrid with its own kind or with jubatum. Hordeum brachyantherum cannot be involved in this proliferation. In fact, data from analyses of naturally-occurring, interbreeding populations have not been published from this northern interior area to show that intergrading is indeed taking place. We do not know if backcrossing is occurring, in particular with brachyan- therum,; nor do we know if the ecologically successful hybrid has had any competitive impact on the sparsely-occurring brachyantherum. Hor- deum brachyantherum quite possibly is being eliminated without inter- grading with the hybrid.
Our observations and analyses of interbreeding populations (Mitchell and Wilton, 1964) indicate brachyantherum is maintaining a separate identity in a substantial portion of its principal range. Such hybridiza- tion as occurs in Alaska is not, according to the evidence, producing an incorporation of the gene complexes of jubatum and brachyantherum. The two species are morphologically distinctive, ecologically at variance except where they meet in disturbances, and for the most part geo- graphically separate as well.
These interpretative differences with the research workers of Ottawa are in no way intended to detract from their excellent cytogenetic and taxonomic work on taxa of the Triticeae.
University of Alaska Agricultural Experiment Station, Palmer, Alaska
LITERATURE CITED
Bownen, W. M. 1962. Cytotaxonomy of the native and adventive species of Hor- deum, Eremopyrum, Secale, Sitanion, and Triticum in Canada. Canad. Jour. Bot. 40:1675-1711.
Covas, G. 1949. Taxonomic observations on the North American species of Hor- deum. Madrono 1:1-21.
Hitcucock, A. S. 1950. Manual of the grasses of the United States. Ed. 2, revised by Agnes Chase. U.S. Dep. Agr. Misc. Publ. 200.
Love, D. 1959. The postglacial development of the flora of Manitoba: A discussion. Canad. Jour. Bot. 37:547-585.
MitTcH_ELL, W. W., and H. J. Hopcson. 1965. A new X Argohordeum from Alaska. Bull. Torrey Club. 92:403-407.
., and A. C. Wirton. 1964. The Hordeum jubatum—caespitosum—brachy- antherum complex in Alaska. Madrono 17:269-280.
Moss, E. H. 1959. Flora of Alberta. Univ. Toronto Press.
RayJHATHY, T. 1966. Notes on the Hordeum jubatum complex. Madrono 18: 243-244.
., and J. W. Morrison. 1959. Cytogenetic studies in the genus Hordeum. IV. Hybrids of H. jubatum, H. brachyantherum, H. vulgare and hexaploid Hordeum sp. Canad. Jour. Genet. Cytol. 1:124-132.
. 1961. Cytogenetic studies in the genus Hordeum. V. H. jubatum and the New World species. Canad. Jour. Genet. Cytol. 3:378-390. Scoccan, H. J. 1957. Flora of Manitoba. Natl. Mus. Canada Bull. 140.
NOTES ON MARINE ALGAE OF WASHINGTON AND SOUTHERN BRITISH COLUMBIA, II
RICHARD E. Norris and JOHN A. WEST
The following list of algae continues the report on marine algae that have not previously been reported in Washington and southern British Columbia (Norris and West, 1966). Our records supplement the lists of Scagel (1957) and Dawson (1961). All specimens are in the herbarium of the University of Washington and are from the State of Washington unless otherwise indicated.
This investigation was supported by the State of Washington Initia- tive 171 Funds for research in biology and medicine. The junior author thanks the Friday Harbor Laboratories for support during the summer of 1964 on their National Science Foundation Grant 747 for the Synop- tic Collections of specimens on the San Juan Islands.
CYANOPHYTA
Aphanocapsa litoralis Hansging was collected on the piles of the new wharf at the Friday Harbor Laboratories, San Juan I., July 27, 1965 (Norris 5307). This alga was growing in large gelatinous masses on posts that contained large amounts of creosote. The algae showed no deleterious affects from the creosote. This species has not been pre- viously reported from the Pacific Coast of North America. It was orig- inally described from the Adriatic Sea. (See Geitler, 1930-1932).
Chlorogloea conferta (Kiitzing) Setchell and Gardner was collected on filaments of Rkodochorton purpureum growing on rocks in a shaded area of the upper littoral region at Mt. Dallas Beach, San Juan I., July 2, 1965 (West 312). This species has not been recorded north of San Mateo Co., California on the Pacific Coast of North America (Setchell and Gardner, 1919).
Dermocarpa hemisphaerica Setchell and Gardner was found growing with Chlorogloea conferta on Rhodochorton purpureum Mt. Dallas Beach, San Juan Island, July 2, 1965 (West 336). The previous known range for this species was the coast of central California (Setchell and Gardner, 1919). D. hemisphaerica is not clearly distinct from D. suf- fulta which also often grows on Rhodochorton purpureum.
Pleurocapsa entophysaloides Setchell and Gardner was collected grow- ing on a boulder cliff at the high littoral level just south of Deadman Bay, San Juan I., June 11, 1964 (Norris 4986). Previous records of this species do not list it north of central California (Setchell and Gardner, 1919).
PHAEOPHYTA
Myrionema phyllophilum Setchell and Gardner was found at Mt. Dallas Beach, San Juan I., June 26, 1965, (Norris 5186). This species, which grows on leaves of Phyllospadix, has previously been known only from Sitka, Alaska (Setchell and Gardner, 1925).
111
112 MADRONO [Vol. 19
RHODOPHYTA
Goniotrichopsis sublittoralis G. M. Smith is very similar to Goniotri- chum elegans in its morphology, but it has numerous small irregularly discoidal chromatophores instead of a single stellate chromatophore with a prominent pyrenoid as in Goniotrichum. Previously found only on the Monterey Peninsula, California this species was dredged as an epiphyte on various larger algae at Hein Bank, south of San Juan I., July 13, 1965. Field collected plants were quite scarce in the collection, but it was isolated into unialgal culture and herbarium specimens (West 397) have been prepared from the cultured material. Smith (1943) did not observe reproductive structures on his specimens but we have found the species readily reproducing by monospores in our cultures. Monospores appear to be transformed vegetative cells produced in both uniseriate and multiseriate regions of the thalli. When mature, the spores are ex- truded through the thick sheath.
Erythrotrichia carnea (Dillwyn) J. Agardh was collected as an epi- phyte on Ceramium gardneri from concrete floats at Shilshole Bay Ma- rina, Seattle on October 27, 1964. The species was isolated into unialgal culture and herbarium specimens have been prepared from these cul- tures (West 398). Relatively common as an epiphyte in larger benthic algae in the inland waters of Washington, it is surprising that this spe- cies has not been recorded here before this time. Field collected plants and the specimens in culture are morphologically the same as the speci- mens described from other regions. Erythrotrichia kylini, described by Gardner (1927) as having band-shaped plastids, may be conspecific with E. carnea because Kylin (1941) described the plastids in former species as stellate, like those of E. carnea. If this characteristic is the same in both species, there appears to be no reason to separate them except for a slight difference in diameter of the filaments. Herbarium specimens of E. kylinii, unfortunatey, do not seem to be abailable for comparison with our specimens.
Acrochaetium densum (Drew) Papenfuss was originally described by Drew (1928) as an epiphyte on Spongomorpha mertensu from San Francisco. Since that time it has not been observed on the Pacific coast of North America, although Nakamura (1944) discovered it as an epi- phyte on Polysiphonia in Japan. In our collection it is epiphytic on Polysiphonia brodiaei (West 245, 345) from Lloyd’s Boat House, Du- wamish Head, Seattle, January 11, 1965 and March 15, 1966. This species with its host may have been transported by marine shipping from Cailfornia since both are known only from the one location in Wash- ington.
Kylinia arcuata (Drew) Kylin was previously reported from Duxbury Reef, Moss Beach (Drew, 1928) and Bodega Bay (Johansen, 1966) in California and only once from Washington (Cape Flattery; Drew, 1928). We have observed K. arcuata as an epiphyte on Polysiphonia brodiaei from Lloyd’s Boat House, Duwamish Head, Seattle, March 15,
1967] NORRIS & WEST: MARINE ALGAE 113
1966 (West 347). Acrochaetium densum and Kylinia arcuata grow on the same host and have certain morphological features in common. Both species were isolated into unialgal culture and monosporangia of both are similar in position and in general morphology. The two species usu- ally can be distinguished by differences in their basal systems, however. K. arcuata has a single basal cell whereas Acrochaetium densum ordi- narily has a system of prostrate filaments arising from two hemispherical cells derived from the original monospore. Occasionally, however, in cul- ture the spores of A. densum do not divide to form two hemispherical cells but instead produce erect filaments directly. In these cases the plants are indistinguishable from Kylinia arcuata. The basal system of K. arcuata shows no modifications, remaining unicellular in all culture con- ditions tested.
The variation exhibited by Acrochaetium densum indicates that use of the basal system morphology may not be entirely reliable as a major taxonomic criterion. Further culture work with other species will dis- close more information on this problem.
Audouinella pectinata (Kylin) Papenfuss was previously known only from Denmark (Rosenvinge, 1909) and Sweden (Kylin, 1944). Our Identificaiton was based on specimens from culture. One clone now in unialgal culture was obtained from a mixed species culture of plants dredged from Hein Bank, south of San Juan I., Washington, June 10, 1964 (West 399). A second clone was obtained from a specimen of Ptilota hypnoides (West 293) collected in the intertidal zone at Glacier Point, Vancouver I., British Columbia, Canada, June 30, 1965. The tetraspor- ophytes of the clones in culture produce monosporangia and tetraspor- angia. Monospores produce new tetrasporophytes. The dioecious game- tophytes which develop from the tetraspores have a unicellular base in contrast to the multicellular base of the tetrasporophyte. The gameto- phytes form monosporangia and sexual structures.
Rhodophysema georgi Batters. Uncertainty was expressed in our pre- vious report on this species in our region (Norris and West, 1966) be- cause it often is especially difficult to distinguish R. georgi from basal cushions of Smithora naiadum. Both species grow on the margins of Zostera and Phyllospadix leaves, but with close examination one can see a distinct difference between the two species. Smithora cushions are large, thick and purplish-black, whereas the thalli of Rhodophysema georgit are smaller, thinner and are a more distinct purplish-red color. The most reliable method for distinguishing between these two species is by examination of the chromatophores in the cells. Smithora cells contain a single chromatophore that is stellate and may be axial or peri- pheral in position. Rhodophysema chromatophores, on the other hand, are numerous in each cell, peripheral in position and disc-shaped.
Iridea lineare (Setchell and Gardner) Kylin has been reported as far north as Cape Mears, Oregon (Doty, 1947). In our collection it was found growing in the intertidal region at the site of the Minnesota Sea-
114 MADRONO [Vol. 19
side Station near Port Renfrew, Vancouver I., British Columbia, July 29, 1964 (Norris 5239). This species is perhaps the most distinctive in the genus as it occurs on the Pacific coast of North America in that sev- eral very narrowly cuneate thalli grow from a common holdfast. Most of the other species have a much broader basal part of the blade.
Antithamnion pygmaeum Gardner, a species closely allied to Anti- thamnion defectum and occupying a similar habitat, previously had a known range extending from Pacific Baja California (Dawson, 1962) to Cape Arago, Oregon (Doty, 1947). We now record A. pygmaeum for two locations in northern Washington: in the sublittoral zone at Hein Bank, Southwest of San Juan I., June 10, 1964, and in the littoral zone at Larabee State Park, South of Bellingham, October 15, 1965 (collected by Maurice Dube). In both cases the plants were discovered growing upon larger benthic algae collected from these sites and placed in cul- ture. Both clones in culture lack the acutely pointed terminal cells of determinate branches found on plants growing in the field. Because of this the cultured specimens appear very much like A. defectum which has blunt branch tips in the field and in culture. Several features indicate that the two species are not identical, however. First, hybridization at- tempts between A. pygmaeum and A. defectum in culture have not been successful. Second, gland cells of A. defectum regularly contain a dis- tinct crystalline body extending longitudinally but the gland cells of A. pygmaeum apparently never include this structure. Finally the basal cell of each determinate branch in A. pygmaeum has the potential to form an indeterminate branch and often does so. A. defectum does not exhibit this tendency in either field collected or laboratory cultured material.
In general the dimensions for vegetative and reproductive structures are smaller in the cultured clones than other authors’ data indicate for field collected specimens. Herbarium specimens of male, female and tetra- sporic material have been prepared from cultured plants. The Hein Bank clone was assigned the number West 406 and the Larabee State Park clone the number West 407.
Ceramium gardneri Kylin is isolated and growing in culture from two collections at Shilshole Bay Marina, Seattle, October 27, 1964 (West 215), and September 8, 1965 (West 334). Previously this species was not known north of Coos Bay, Oregon (Doty, 1947).
Hymenena kylinu Gardner was found growing as one of the dominant intertidal species on the reef at the site of the Minnesota Seaside Sta- tion near Port Renfrew, Vancouver I., July 29, 1965 (Norris 5254). Previously this species was collected no farther north than Cape Blanco, Oregon (Doty, 1947). This species seems to be closely allied with Hymenena flabelligera and may be only a variant of that species caused by different environmental conditions.
Cryptopleura corallinara (Nott) Gardner was known only from Mexico and southern California until 1965 when Dawson reported it in his collections in northern California (Dawson, 1965). In our region
1967 | NORRIS & WEST: MARINE ALGAE mS
fruiting specimens were found in several localities: dredged at 5-7 fathoms, Partridge Bank, west of Whidbey I., July 6, 1965 (Norris 5196); intertidal at American Camp Beach, San Juan I., July 2, 1965 (Norris 5217); intertidal on the reef at the side of the Minnesota Sea- side Station, near Port Renfrew, Vancouver I., British Columbia, July 29, 1965 (Norris 5256).
CHLOROPHYTA
Pseudodictyon geniculatum Gardner. Blades of Cymathere triplicata collected in the drift at Cattle Point, San Juan I., June 15, 1965 (Norris 5181) have extensive infections of a partially endophytic green alga that seems to be Pseudodictyon because of the tendency of the branches to unite to form a net. Unlike the plants described by Gardner, the endo- phyte in our collection forms a pseudoparenchymatous tissue mostly on the surface of the host. Sporangia may be formed in the pseudopar- enchyma as well as by the endophytic filaments. Sporangia in our speci- mens are from 2 to 4 times larger than the vegetative cells. Biflagellate zoospores were released from the sporangia two days after the collection was made. The zoospores are approximately 10 » by 5 p, ovoid, with an acute flagellar end, with a single band-shaped chloroplast onto which a conspicuous stigma is attached. The flagella are approximately 1% times the cell length. The chloroplasts of most of the vegetative cells con- tain more than one pyrenoid. Gardner (1909) described a single pyre- noid in his specimens of P. geniculatum and it may be that our specimens represent a new species in producing pseudodoparenchyma. Previously P. geniculatum has been collected no farther north than Trinidad Bay, California (Dawson, 1965), although Doty (1947) records a doubtful collection by Gardner at Sunset Bay, Oregon.
Pseudulvella consociata Setchell and Gardner was found on shells of Calliostoma sp. dredged at Mosquito Pass, Northwest of San Juan I., Oc- tober 29, 1966 (Norris 5308 ). The filaments of this alga form a solid en- crusting layer on the shells, and the basal system does not penetrate the host shells. One or two pyrenoids may be present in the chloroplasts of apical cells of erect filaments, although most chloroplasts have one. This species has not been previously recorded north of central Oregon (Doty 1947).
Department of Botany, University of Washington, Seattle Department of Botany, University of California, Berkeley
LITERATURE CITED
Dawson, E. Y. 1961. A guide to the literature and distributions of Pacific benthic algae from Alaska to the Galapagos Islands. Pacific Sci. 15.370-461. . 1962. The marine red algae of Pacific Mexico. Part 7. Ceramiales: Cera- miaceae, Delesseriaceae. Allan Hancock Pacif. Exped. 26:1—-207. . 1965. Marine algae in the vicinity of Humboldt State College. Humboldt State College, Arcata, California (mimeographed). Dory, M.S. 1947. The marine algae of Oregon. Part I. Chlorophyta and Phaeophyta. Farlowia 3: 1-65.
116 MADRONO [Vol. 19
. 1947. The marine algae of Oregon. Part II. Rhodophyta. Farlowia 3:159- 2155
Drew, K. M. 1928. A revision of the genera Chantransia, Rhodochorton, and Acro- chaetium with descriptions of the marine species of Rhodochorton (Naeg.) gen. emend. on the Pacific coast of North America. Univ. Calif. Publ. Bot. 14: 139-224.
GARDNER, N. L. 1909. New Chlorophyceae from California. Univ. Calif. Publ. Bot. 3:371-375.
. 1927. New Rhodophyceae from the Pacific Coast of North America. II. Univ. Calif. Publ. Bot. 13:235-272.
GEITLER, L. 1930-1932. Cyanophyceae. Jn Rabenhorst’s Kryptogamen-Flora von Deutschland, Osterreich und der Schweiz, 2nd ed. Bd. 14 (issued by R. Kolk- witz) Akad. Verlagsges. Leipzig.
JOHANSEN, H. W. 1966. The benthic marine algae of Bodega Head, p. 35-112. In Hand, Cadet (ed.) A marine ecological survey of the Bodega Head region. University of California, Berkeley, California (mimeographed).
Kyun, H. 1941. Californische Rhodophyceen. Lunds Univ. Arsskr. II. Sect. 2. 37(2): 1-51.
. 1944. Die Rhodophyceen der Schwedischen Westkiste. Lunds Univ. Arsskr. IT. Sect. 2. 40(2) :1—104.
NAKAMURA, Y. 1944. The species of Rhodochorton from Japan II. Sci. Pap. Inst. Algol. Res. Fac. Sci. Hokkaido Imp. Univ. 2:273-291.
Norris, R. S., and J. A. West. 1966. Notes on marine algae of Washington and southern British Columbia. Madrono 18:176-178.
ROSENVINGE, L. K. 1909. The marine algae of Denmark. Par I. Introduction. Rho- dophyceae I. (Bangiales and Nemalionales). Danske Vid. Selsk. Skr. Ser. 7, Nat. Math. Afd. 7(1):1-149.
ScaGEL, R. F. 1957. An annotated list of the marine algae of British Columbia and northern Washington. Nat. Mus. Canada. Bull. 150.
SETCHELL, W. A., and N. L. GARDNER. 1919. The marine algae of the Pacific Coast of North America. Part I. Myxophyceae. Univ. Calif. Publ. Bot. 8:1-138.
. 1925. The marine algae of the Pacific coast of North America. Part III. Melanophyceae. Univ. Calif. Publ. Bot. 8:383—-898.
SmiTH, G. M., and G. J. HoLiLensBerc. 1943. On some Rhodophyceae from the
Monterey Peninsula, California. Am. Jour. Bot. 30:211-222.
GOSSYPIUM TRILOBUM: AN ADDENDUM
PAuL A. FRYXELL AND CLIFFORD R. PARKS
Existing knowledge concerning Gossypium trilobum (Moc. & Sess. ex DC.) Skov. emend. Kearn., has been summarized by Fryxell (1965a). He concluded that this species is distinct from G. thurberi Tod. in spite of much nomenclatural confusion, but is more closely allied to the latter than to G. gossypioides (Ulbr.) Standl., as was suggested by Phillips (1966). It was noted that the plant unfortunately was not available in any of the living collections of Gossypium upon which the extensive experimental work with this genus has been and continues to be based. Some recent attempts to collect living material of this species have been unsuccessful.
On the basis of the distributional data summarized in the previous study, we planned to recollect this species, with the special intention of adding living material to existing Gossypium gardens. Accordingly, we visited the Mexican states of Sinaloa, Jalisco, Michoacan, México, Guerrero, and Morelos, where the plant is known to occur. Established populations of G. trilobum were eventually found in Michoacan, and both living and preserved specimens were collected. The present com- munication reports observations on these collections.
This trip was supported by the junior author’s National Science Foundation Grant GB3900. The trip was undertaken to collect fresh petal tissues of Gossypium spp. for chemotaxonomic analysis from the winter garden at Iguala, Guerrero, Mexico, maintained jointly by the U.S. Department of Agriculture and the National Cotton Council, in cooperation with the Mexican government, and to introduce G. trilobum into cultivation. The authors were aided in these collections by Kenneth Montgomery and David L. Dreyer.
Ecological observations. The city of Zitacuaro, Michoacan, is situa- ated at an elevation of 1,993 m. A road extends south from Zitacuaro down the valley of the Rio Zitacuaro and across to the valley of its tributary, the Rio Temascaltepec, where it reaches the town of Tuzantla. The Zitacuaro-Tuzantla road descends steadily as it progresses south- ward. We found populations of G. trilobum at km 9 and at km 21, at elevations of approximately 1,800 m and 1,600 m respectively. The km 9 site represents the upper altitudinal limit from which plants of this taxon have been collected. Previous collections (Fryxell, 1965a) have been recorded at altitudes of 800 m to approximately 1,500 m.
The most well-developed specimens grew on a steep rocky slope fully exposed to the sun, in what appeared to be undisturbed vegetation. Although other large bushes and small trees grew on the slopes with the Gossypium, they were too scattered to compete directly for light. The hillside also had a good if not dense covering of grasses and forbs, compared to habitats typical for Gossypium, which generally are more arid and open.
117
118 MADRONO [Vol. 19
Other plants of G. trilobum were observed growing in the fence row, along the roadside, and at the edges of cultivated fields (i.e., in dis- turbed situations in dense, weedy, and shrubby vegetation). Some of these specimens had been cut (or grazed) in the past, and thus had a more shrubby aspect. Plants of G. trilobum, apparently established at the time of disturbance, appeared able to compete successfully with their weedy neighbors; but whether seedlings of G. trilobum can succeed in establishing themselves in such dense vegetation is doubtful. The only seedling found was in bare rocky soil near the edge of the road with no other plants in shading distance. It will be necessary, during the right season, to observe germination and seedling establishment in the field. Comparisons of the disturbed and undisturbed habitats in this regard will permit definite statements concerning the competitive abil- ity of G. trilobum seedlings.
The plants of G. trilobum described above were growing in a habitat apparently typical for the species, one fully exposed to strong insolation. However, a few plants were found in quite a different habitat. At one side of the road near km 21 there is a canyon several hundred feet deep, with nearly vertical sides that drop down to a stream at the bottom. A trail cut into the side of the canyon wall enables one to descend to the bottom. A few plants of G. trilobum were observed on this canyon wall, growing in the shade resulting from a heavy growth of brush and scattered trees, and from the curtailed daylength of the nar- row canyon. The original canyon inhabitant, a large and possibly old plant, may have reached its position in the course of a minor land- slide. A few young plants were found below the older plant, some nearly to the canyon bottom. Apparently G. trilobum has a greater tolerance than most species of Gossypium for shading and more ability to with- stand crowding.
The plants observed were in the full flush of flowering and had an ample crop of immature fruit in mid-October. Evidently the flowering and fruiting pattern of this species is similar to that of the closely allied G. thurberi. The latter species begins flowering abruptly in early autumn and then flowers and fruits heavily and rapidly, maturing a large crop of fruits within two months of the onset of flowering. It is vegetative (or dormant) for the balance of the season. This conclusion agrees with collection dates of flowering specimens cited previously (Fryxell, 1965a).
The origin of the boll weevil (Anthonomus grandis Boh.) has puz- zled entomologists for some time. It has been suggested that one of the wild cottons of Mexico might prove to be the original host from which these insects later transferred to cultivated cotton. No convincing evi- dence has yet been put forward in support of this thesis. However, observations have never previously been made on G. trilobum, a species that in geographical terms might have played such a role. The plants here discussed were carefully examined for signs of weevil infestation,
1967] FRYXELL & PARKS: GOSSYPIUM TRILOBUM 119
Fic. 1. Leaves of Gossypium trilobum, showing the five-lobed form (a, b) typical of the vegetative stalks, and the three-lobed form (c, d) typical of the sympodial flowering branches, x 4.
but none could be detected. This evidence suggests that Gossypium trilobum has not played a role as primary host of the boll weevil. More- Over, recent evidence (Fryxell & Lukefahr, 1967) indicates that the primary host may have been found in species of the genus Hampea Schlecht.
120 MADRONO [Vol. 19
Fic. 2. Calyx of Gossypium trilobum with very young fruit showing distinctive form of margin, x 5.
Morphological observations. We were able to observe certain features in living plants that are difficult or impossible to observe in herbarium specimens, but which significantly extend our understanding of the species. The plants are large shrubs or small trees, 3 to 4 m in height, with a rounded crown. The trunks of the specimens on the undisturbed slope did not branch for 12-1 m above the ground, above which they
1967] FRYXELL & PARKS: GOSSYPIUM TRILOBUM 121
branched freely (in contrast to specimens of G. thurberi which branch freely at ground level and thus lack a single trunk).
The fruits of G. trilobum are predominantly three-loculed, but are rarely two-loculed. No fruits were seen with four locules. The fruits of G. thurberi are also predominantly three-loculed, but are occasion- ally four-loculed.
The involucral bracteoles have been described as cordate-acuminate and entire. This is essentially correct, except that the bracts on one plant (Fryxell 562) were weakly laciniate, rather than strictly entire. Bracts on other specimens showed minute, widely scattered teeth that were not superficially evident.
The leaves are a little more than % cut (fig. 1) on the average. A few leaves were nearly as deeply lobed as is typical of G. thurberi, but on the average the leaf dissection in this latter species is much greater than in G. trilobum. In both species leaf dissection is variable among the leaves of a given plant, depending on the position and exposure of the individual leaf, but the leaf spectra (Melville, 1953) of the two species are quite distinct. In young plants of the same age grown in culture, leaves of G. trilobum are less dissected than those of G. thur- beri. In G. trilobum, climax leaves along the main stalk are typically five-lobed, whereas those occurring on the lateral fruiting branches are generally three-lobed. The specific epithet is derived from the latter type of leaf, since it is more commonly gathered and preserved in herbarium material because of its association with flowers or fruits. Some of the reduced, upper leaves are unlobed and ovate in shape. The foliar nectaries are only sometimes present on climax leaves. They are often but not always present on the trilobed leaves of the flowering branches.
The morphology of the calyx is a distinctive feature of the species (fig. 2). The calyx has been described as having a variable number of irregular divisions (Fryxell, 1965a). Examination of living material has clarified the nature of this structure. The calyx is basically five- lobed, a condition that is characteristic of the balance of the genus (although several species have calyces that are truncate by reduction). During the development of the bud of G. trilobum, each calyx lobe splits, generally into two parts, and thus gives rise to a calyx which usually has ten divisions. There is sufficient irregularity in this process, how- ever, that both the number and the size of these divisions are variable. A similar development occurs in no other species of Gossypium.
Gossypium trilobum has been compared several times with the mor- phologically similar G. thurberi (Kearney, 1937; Fryxell, 1965a). Field observation of the plants confirms this similarity, both authors being already familiar with G. thurberi in its native habitat in the Sonoran desert. Specifically, the present observations confirm the hypothesis that these are a pair of vicarious species (Fryxell, 1965b).
Cytology. Buds were collected from two plants (Fryxell 562, 563)
122 MADRONO [Vol. 19
ete ie Aguas-, ‘ Vin oie 3 "ss . -” calientess*-._ ,! ¢ ee ~
ae
ss
Mm vPuebld
sh : ths a s..“> iMorelos: = vs be ay 4
Fic. 3. Map showing known collection sites for Gossypium trilobum.
and examined cytologically by Meta S. Brown, Texas AXM University, whose collaboration is gratefully acknowledged. The gametic chromo- some number was n — 13, and the chiasma frequency 1.8 chiasma per bivalent, as determined from the analysis of 926 bivalents from 74 PMC’s, A low frequency of quadrivalent association was observed.
Additional Comments. Local residents were questioned about the plant, which was well-known to them. The people of the area do not use the plant but call it “algodoncillo” (meaning “little cotton”). They thus recognize its relation to cultivated cotton, even though G. trilobum produces no fiber.
By collating information about the distribution of G. trilobum (Fryx- ell, 1965a), the flowering pattern of the species (described above), and the itinerary of the botanists, Sessé and Mocino, who collected the type specimen of this species (Rickett, 1947), one can deduce that the type was probably collected in the autumn of 1790 in central Michoa- can, as these collectors traveled from Patzcuaro to Apatzingan.
The distribution map of G. trilobum (Fryxell, 1965a) indicates that the species is fairly widespread, extending more than 500 miles from southern Sinaloa to western Morelos. Yet the species has generally been regarded as rare. A plot of the known collection sites (fig. 3) indicates a concentration in the southern parts of the states of Michoacan and México, in those areas where the elevation provides a suitable habitat
1967] FRYXELL & PARKS: GOSSYPIUM TRILOBUM 123
for this species. The region concerned, possibly including the north- western part of Guerrero as well, is one that is not readily accessible by good roads. Perhaps the apparent rarity of this species is consequent on botanists only rarely having penetrated to its native environs. This conclusion is borne out by local informants in the area south of Zitacu- aro, where the present collections were made, being quite familiar with “algoedoncillo,’ and by the specific comment made by one of them that there were many more of the plants to be found in the surround- ing hills. Gossypium trilobum has perhaps never been rare, only poorly known to botanists.
Herbarium material of G. trilobum obtained on the trip described in this paper (Fryxell 562, 563, 564, 565, 567, 568; and Parks 257, 261, 264, 270) will be deposited at the Tracy Herbarium of Texas A&M University (TAES) and at the Herbarium of the Los Angeles State and County Arboretum; duplicates will be distributed elsewhere. The collec- tions were made under a permit granted by the Secretaria de Agricultura y Ganaderia, Instituto Nacional de Investigaciones Forestales, México D.F. It is a pleasure to acknowledge the cooperative attitude of that office. Plants of G. trilobum have been established at the winter garden at Iguala, Guerrero.
Crops Research Division, U.S. Department of Agriculture, Texas A&M University, College Station
University of North Carolina, Chapel Hill
LITERATURE CITED
FrYXELL, P. A. 1965a. A further description of Gossypium trilobum. Madrofio 18:113-118. . 1965b. Stages in the evolution of Gossypium L. Adv. Fron. Plant Sci. 10:31-56. . and M. J. LUKEFAHR. 1967. Hampea Schlecht.: Possible primary host of the boll weevil. Science 155:1568-1569. Kearney, T. H. 1937. Ingenhouzia and Thurberia. Am. Jour. Bot. 24:298-300. MELVILLE, R. 1953. Growth and plant systematics. Proc. Linn. Soc. 164:173-181. Puiiips, L. L. 1966. The crytology and phylogenetics of the diploid species of Gossypium. Am. Jour. Bot. 53:328-335. RickeTt, H. W. 1947. The Royal Botanical Expedition to New Spain 1788-1820. Chron. Bot. 11:1-86.
LEE BONAR
A tribute to Lee Bonar. We honor your many years of dedicated teach- ing and your service as Chairman in the Botany Department at Berkeley, your varied and continuous studies on the fungi of California and your long friendship for our Society, which in the past you served as president.
Colleagues who were associated with you in the teaching of general botany, recall with great admiration your ability to interest undergrad- uates not only in the attractions of the local flora but particularly in a balanced knowledge of all aspects of plant life. Your mycological re- searches extend over more than forty years and have been carried on in both the herbarium and the field. These studies have ranged from ring- worm to wood rot, from rusts to sooty molds, from sausage yeasts to poisonous mushrooms and from cankers to witches brooms. Your com- prehensive knowledge of the fungi has often seemed to have no end and has always been a source of wonder to your students and colleagues, as well as to the public that has depended upon you so frequently for expert assistance.
Our tribute brings affectionate wishes for your continued good health and for many more years of the quiet but deep satisfaction which you derive from your mycological studies.
125
A HYBRID HAWKWEED FROM THE OLYMPIC MOUNTAINS OF WASHINGTON
ARTHUR R. KRUCKEBERG
The subalpine forest type of the Olympic Mountains is not a likely habitat in which to encounter interspecific hybrids. The vegetation typically presents the aspect of a closed community: Open park-like stands of Abies lasiocarpa (Hook.) Nutt. (Subalpine Fir) and 7J'suga mertensiana (Bong.) Sarg. (Mountain Hemlock) form the arboreal setting for the dense growth of grasses and forbs that clothe the inter- vening subalpine meadows so typical of the upper montane in the Pacific Northwest. As every botanist knows, a species hybrid in such a habitat would be most unexpected. But equally well-known to the field botanist is the effect of disturbance in creating the conditions for crossing.
Thus I was not particularly surprised to encounter a hybrid hawk- weed in a disturbed habitat of the upper montane. Along the northeast- ern flank of the Olympic Mountains in northwestern Washington, U.S.A., the range is buttressed by Hurricane Ridge, a high subalpine hogsback running east and west at approximately 5000 feet elevation. Vegetation patterns are complex along the ridge; exposure and slope change continuously—and abruptly—throughout its length. Basically three vegetation patterns occur here: (1) subalpine conifer forest, (2) upper montane (Hudsonian) meadow on gentle slopes, and (3) the flora of rock outcrops and talus. Midway between the U.S. Park Service lodge on Hurricane Ridge and Obstruction Point to the east, there is an old burn in the meadow-and-forest habitat. A dense cover of grasses and forbs has invaded the site. It was in this situation that on August 1 of 1963 I found two plants which I took to be hybrids between the two very different hawkweeds, Hieracium albiflorum Hook. and H. gracile Hook. The plants were growing in an open grass—forb swale of what had been a subalpine forest habitat before the fire. The putative parents were in close association with the hybrids (Kruckeberg 5689 a—e, WTU). In fact, individuals of both species were included in the sods containing the hybrids; the sods were transplanted to pots and have been in greenhouse and outdoor culture at Seattle ever since.
The probable parents of the hybrids can be easily distinguished both morphologically and by their typical ecological preferences (table 1). Hieracium albiflorum is a tall plant of open to deep woods; it has densely pilose basal leaves and tends to be nearly glabrous in the inflorescence. Its ligules are dull white in numerous paniculately disposed heads. In the Olympics, H. albiflorum probably reaches its altitudinal limit nor- mally well below the 5000 foot level, and is not usually found on Hurri- cane Ridge; the species certainly is an unexpected visitor to this habi- tat. It is not improbable that the plants are adventive on the ridge, as they often occur in abundance after fire or other disturbance at lower elevations.
126
1967]
TABLE 1.
KRUCKEBERG: HYBRID HAWKWEED
127
MorPHOLOGY, CHROMOSOME NUMBER, AND ECOLOGICAL PREFERENCES
OF THE SPECIES AND THEIR HyBrip
H. albiflorum
Basal Leaves
Stem
Inflor- escence
Involucre
Ligules
Achenes
Pappus
Chromo- some Number
Habitat Pref- erence
Hybrid plants
H. gracile
10-12 cm long, 2.0—5 cm wide; often re- motely denticulate; pilose (copious above, sparingly below)
22-97 (120) cm tall, long, straight tawny- hued hairs below, be- coming spare upward to glabrous in inflores- cence
Branches with short, greenish, gland-tipped hairs
Long stiff hairs usual- ly sparse or absent; copious short, green, gland-tipped hairs
pale white; well ex- serted
2.5 mm long, pale brown
tawny or sordid
n= 9, Kruckeberg 5618d, 5411 (Chelan Co.), 4393 (Clallam Co.)
Open to dense conifer- ous forests; low to middle altitudes; also in disturbed sites
12-15 cm long, 1.5—2 cm wide; remotely den- ticulate ; shaggy hirsute, esp. at base, hair mostly long hyaline, erect, some short, green and gland- tipped along margin
36-64 cm tall; long hairs copious below, sparse upward; hyaline short and matted hairs few to none below, more dense upward to copious in inflorescence (appearing stellate)
Pubescence as on stem
Black hairs moderately dense, of two types: gland-tipped, short and in + regular rows; scat- tered long hairs; hyaline matted hairs mostly at base; pubescence less dense than in H. gracile
ochroleucous to pale yellow
n=9, 5681le (two of 186 meiocytes with lagging chromosomes)
Open grass-forb swales of old burn, Hurricane Ridge, Olympic Moun- tains, Washington
2-5 cm long, 0.5—1.0 cm wide; usually entire, nearly glabrous, some pale gland-tipped hairs on margin and midrib
5=20" (27)? cm: high ; black, gland-tipped, long hairs mixed with short, hyaline and matted hairs, both more dense upward, esp. long black setose hairs
Densely hairy: mixture of matted hyaline hairs and blackish, gland-tip- ped setose hairs
Pubescence as in inflor- escence but black hairs more dense (gland-tip- ped when young) ; long and short black hairs, or all long and gland-tip- ped
bright yellow; moder- ately exserted
2 mm long; blackish brown to black
dull white
n = 9, 3985and 5633 (Skamania Co.), s.n. (Jefferson Co.)
Subalpine parks and meadows; commonly in closed communities of other grasses and forbs
128 MADRONO [Vol. 19
TABLE 2. POLLEN STAINABILITY AND SEED SET IN SPECIES AND HyYBrIDs
Percent Stainable Number of Pollen* Seed set H. albiflorum Kruckeberg 5681d 97 high H. gracile 5681a, c 94, 100 high (2 plants) Hybrids 5681a 27 13 (from 50 heads) 5681e 40-64 12 (from 46 (3 heads) heads) 2nd Generation hybrid progeny from 5681la G2e782 heads all (2 plants) sterile 5681e 82 30 (from 35 heads)
* Counts of 200 grains in aniline blue-lactophenol.
The other species, Hieracium gracile, is truly at home in the sub- alpine meadow habitat. It is surely as much an indicator of the sub- alpine (Hudsonian) zone in the Pacific Northwest as such ubiquitous grasses and forbs like Deschampsia atropurpurea (Wahlenb.) Scheele, Phleum alpinum L., Erigeron peregrinus (Pursh) Greene, Veronica cusicku Gray, and Luetkea pectinata (Pursh) Kuntze, to name a few. Typically, H. gracile is a much shorter plant (3-35 cm) than H. albi- florum. It has nearly glabrous basal leaves; the pubesence of the cauline portions increases upward to a dense mixture of long and short hairs in the inflorescence. The peduncles and involucral bracts are coated with both short, matted hyaline hairs and long, blackish (often gland-tipped) setae; the heads in bud have a distinct grizzled and blackish aspect. In- cidentally, though the regional manuals (Abrams and Ferris, 1960; Hitchcock, et al, 1955) describe the short hoary portion of the indu- mentum as stellate, I could not confirm this observation; rather, the matted aspect of the hyaline hairs often give the hairs a criss-cross pat- tern which simulates the stellate character. H. gracile has bright yellow ligules in fewer, more congested heads, as compared to H. albiflorum.
The hybrids are clearly intermediate in the features that distinguish the two putative parents (table 1). Stature, foliage, vesture, number of heads, ligule color, and achenes all manifest the influence of both
A. gracile and H. albiflorum.
Both species have the chromosome number, n = 9; and from sev- eral samples taken in Washington, I have yet to detect any chromo- some irregularities during microsporogenesis. Moreover, both species have ‘“‘good” (well stained) pollen (table 2). Thus, there is nothing to suggest that the two species are apomictic, a condition commonly en-
1967] KRUCKEBERG: HYBRID HAWKWEED 129
countered in other species of Hieracium (Stebbins, 1950). The pre- sumed sexualty and the close sympatry of the two species provide the opportunity for interspecific hybridization. The expression of inter- mediate morphology in the suspected hybrids reinforces the judgment of hybridity.
The plants taken from the wild in 1963 have flowered and produced seed in the greenhouse now through 1966. Both species produce copious and viable seed in cultivation. The presumed hybrid plants produce only about 27-64% stainable pollen (table 2). Their heads mature but yield only a few plump seed (table 2). Healthy progeny have been raised from the few good seed harvested from the original wild hybrids and from the second and third generation hybrid derivatives. The later gen- eration hybrid progeny tend more to perpetuate the intermediate fea- tures of the original hybrid than to segregate for the parental charac- ters.
Examination of meiosis in microsporogenesis of hybrids reveals hardly any chromosome irregularity. Of 186 sporads examined, ranging from diakinesis to telophase II, only two cells were observed to have any irregular (lagging) chromosomes.
It would appear that Hieracium albiflorum and H. gracile, though two distinctly different species, can hybridize in the wild. As the hybrids are not wholly sterile, the gene flow could increase by introgression or be restricted to hybrid swarms. Botanists should expect to find the hybrids and their derivative progeny repeatedly throughout the ranges of the two species, presumably most often under the ecological conditions de- scribed above. I have yet to encounter reports of other hawkweed hy- brids for western North America. H. albiflorum overlaps in range with several other western species; other crosses will undoubtedly be en- countered.
Note added in proof. A natural hybrid between H. albiflorum and H. ar- gutum in Monterey Co., California, has been reported by Anderson and Stebbins (1954).
Funds for this study were provided by State of Washington Initiative No. 171. Specimens of species and hybrids will be deposited in the Her- barium of the University of Washington.
Department of Botany, University of Washington, Seattle.
LITERATURE CITED
ABRAMS, L., and R. S. Ferris. 1960. Illustrated flora of the Pacific States. Vol. IV. Stanford Univ. Press.
ANDERSON, E., and G. L. Stepsins, Jr. 1954. Hybridization as an evolutionary stimu- lus. Evolution 8:378-388.
Hircucock, C. L., A. Cronouist, M. Ownsey, and J. W. Tuompson. 1955. Vascular plants of the Pacific Northwest. Part 5: Compositae (A. Cronquist). Univ. Washington Press.
STEBBINS, G. L., Jr. 1950. Variation and Evolution in Plants. Columbia Univ. Press.
A SURVEY OF THE VASCULAR PLANTS ABOVE TIMBERLINE ON MT. HOOD, OREGON
LUREL HALL
INTRODUCTION. To my knowledge, a survey of the vascular plants above Timberline on Mt. Hood, Oregon, has not been made previously. The colorful old Oregon botanist, Thomas Howell, published the Flora of Mt. Hood in the Mazama’s Magazine in 1896 (Howell, 1896) but in this article no differentiation was made between the plants above and below the Timberline. Mr. Howell’s article came as a result of previous work done in the Mt. Hood area with Suksdorf, another famous north- west botanist, and subsequently appeared in the records of congress in 1930 as the only available list, although incomplete, of the flora of the Mt. Hood area (Lange, 1953).
The nearly 70 years since Mr. Howell’s study have wrought changes as shown by the differences in some of the species found.
GrEocRAPHY. Mt. Hood is situated a little east of the middle of the Cascade range of mountains. It is approximately 22 airline miles south of the Columbia River, 50 miles ESE of the city of Portland, 29 miles SW of The Dalles and 70 miles NW of Bend. Highway access is U.S. 26 from Portland or Bend and by State Highway 35 from Hood River. These two roads form a loop highway on the south and east side of the mountain at an average of 4000 ft. elevation. There is no heavy duty road on the west or north sides but a maze of all weather roads traverses these areas to about 4000 ft. Many improved forest service trails make walking access to many outstanding scenic locations easy and enjoy- able.
GroLocy. The mountain is situated in a belt of volcanic activity (Wise, 1964) which has been progressing for many years beginning with the extrusion of the Miocene Columbia River basalt over a wide plateau encompassing northern Oregon and southern Washington. Later (about 15 million years ago) olivine basalt and some andesite erupted from many centers throughout the length of the Cascade range building cones up to 6500 feet in elevation. Some of this activity has continued until the present. In recent times (ending about 10 to 15 thousand years ago) the Mount Hood Volcano, through a series of eruptions and extru- sions, built the cone. Subsequently, a large glacier formed in the crater, spilled over the south rim, and eroded the south face of the mountain including the major part of the south and southwest rim. Other large glaciers formed at various positions on the mountain eroding the ash and detritus and in places leaving remnants of thicker lava flows. These glaciers have been a major factor in reducing the original cone’s height by nearly a thousand feet and creating varied scenery which is unex- celled.
SURVEY Limits. A topographic map of the mountain is included with this paper showing general areas and limits of the survey. The exact
130
1967 | HALL: MT. HOOD 131
limits of Timberline are difficult to pinpoint on a large area such as Mt. Hood. The actual line of demarcation of moderate timber is approx- imately 6500 feet. This is probably 1000 feet lower than the normal timberline for this climate and general edaphic factors would dictate. The major cause of this discrepancy is the rapid erosion taking place on the surface. Glaciation and running water are moving the mountain away faster than the slowly developing trees can establish themselves. Evidence for this is the fact that the trees are growing at higher eleva- tions on the ridges than in the valleys where major erosion factors are progressing.
Timberline is defined here as the approximate route of the Timberline Trail excepting the White River area and the entire west side where it dips considerably below Timberline to facilitate crossing rugged and dangerous canyons and cliffs. In many specific localities the trail traverses timberlands. Where this occurs the survey includes only the plants above the actual timberline. Three trees, Tsuga mertensiana, Abies lasiocarpa and Pinus albicaulis, are included because their range extended, on many ridges, to over 8000 feet.
METHODS AND RESULTS. Several trips were made to six different areas on Mt. Hood (fig. 1) from July, 1965 through Sept. 1966. Specimens on which this report is based are in the Dudley Herbarium of Stanford University with the exception of that of Polygonum minimum which is at Washington State University and was kindly determined by Marion Ownbey. Subsequent collections will probably increase the number of species on Mt. Hood.
Two ascents were made to the summit. These are not included in the general areas because no specimens were taken. These trips were made to find the extremes that any plants were growing, and to search for and collect any that were higher in elevation than the regular areas covered. Both ascents were made up the south side. One of the highest plants found on these trips was a small clump of Sedum divergens growing from a crevice in a large andesitic rock at approximately 9500 feet, west of the White River glacier, just below Crater Rock. Interest- ingly enough, this same species was found blossoming beneath an inch of clear ice on the edge of Eliot glacier, east of Pulpit Rock on the north side of the mountain.
Spraguea umbellata was also found at about the 9500 ft. elevation. At this extreme altitude the plant was small and the blossoms were growing right on the ground. S. umbellata was not found on the north side of the mountain; however, Hydrophyllum capitatum seemed to fill the approximate niche there. Sitanion hystrix, Trisetum spicatum, Pen- stemon menziesu, Juncus parryi, Solidago spathulata, Anemone drum- mondu and Collomia larsenii were all found growing at high altitudes. Lupinus lyallu, Phlox diffusa and Polygonum newberryi were some of the most abundant and were distributed over a wide range. The other extreme was Collomia larsenii, of which only a few plants were found
132 MADRONO [Vol. 19
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Fic. 1. Mt. Hood, Oregon. Hatching indicates areas collected.
and those were in a very restricted habitat.
Many ecotonal species are included which do not occur at any appre- ciable distance above the actual timberline. However, if one plant was found on the upper side of the trail, when the trail was distinctly out of the timber, it was included.
In all, the flora consists of 28 families, 56 genera and 82 species. The following list uses the nomenclature and arrangement to be found in Peck’s Manual (1961).
Pinaceae. Pinus albicaulis Engelm., Tsuga mertensiana (Bong.) Sarg., Abies lasiocarpa (Hook.) Nutt.
Cupressaceae. Juniperus sibirica Burgsd.
1967] HALL: MT. HOOD 133
Gramineae. Bromus breviaristatus Buckl., Festuca viridula Vas., F. ovina L., Sitanion hystrix J. G. Sm., Trisetum spicatum (L.) Richt., Deschampsia atropurpurea (Wahl.) Scheele, Phleum alpinum L., Stipa occidentalis Thurb., Agrostis humilis Vas., A. diegoensis Vas.
Cyperaceae. Elyna bellardii (All.) Degl., Carex straminiformis Bail., C. ablata Bail.
Juncaceae. Juncus parryi Engelm., J. drummondu E. May., J. mer- tensianus Bong., Luzula spicata (L.) DC.
Liliaceae. Erythronium montanum Wats., Calochortus elegans Pursh.
Orchidaceae. Spiranthes romanzo fiana Cham.
Salicaceae. Salix comutata Bebb, S. pennata Ball.
Polygonaceae. Polygonum bistortoides Pursh, P. newberryi Small, P. minimum Wats., Eriogonum marifolium T. & G., E. ovalifolium Nutt.
Portulacaceae. Spraguea umbellata Torr.
Caryophyllaceae. Stellaria umbellata Turcz., Arenaria obtusiloba (Rydb.) Fern., Silene suksdorfii Robins.
Ranunculaceae. Anemone occidentalis Wats., A. drummondu Wats., Ranunculus eschscholzu Schlecht.
Crassulaceae. Sedum divergens Wats., S. debile Wats.
Saxifragaceae. Saxifraga caespitosa L.
Grossulariaceae. Ribes acerifolium How.
Rosaceae. Sorbus occidentalis (Wats.) Greene.
Leguminosae. Lupinus lyallii Gray, L. suksdorfii Robins., L. alpicola Hend., L. arcticus Wats. var. subalpinus (Robins.) C. P. Sm.
Hypericaceae. Hypericum anagalloides C. &S.
Onagraceae. Epilobium hornemannii Reich.
Umbelliferae. Lomatium angustatum (C. & R.) St. John.
Ericaceae. Phyllodoce empetriformis (J. M. Sm.) D. Don, P. glandu- liflora (Hook.) Cov., Cassiope mertensiana (Bong.) G. Don, Vaccinium membranaceum Dougl.
Gentianaceae. Gentiana calycosa Griseb.
Polemoniaceae. Phlox diffusa Benth., Collomia larseniu (Gray) Pays., Polemonium californicum Eastw.
Hydrophllaceae. Hydrophyllum capitatum Dougl.
Scrophulariaceae. Penstemon menziesi Hook., P. euglaucus English, Veronica arvensis L., V. alpina L., Mimulus lewistt Pursh, M. tilingu Reg., Castilleja suksdorfu Gray, C. hispida Benth., C. oreopola Greenm., Pedicularis racemosa Dougl.
Caprifoliaceae. Lonicera involucrata (Rich.) Banks.
Valerianaceae. Valeriana sitchensis Bong.
Compositae. Haplopappus bloomeri (Hook.) Gray, Solidago spathu- lata DC., Aster alpigenus (T. & G.) Gray, A. campestris Nutt., Erigeron poliospermus Gray, Anaphalis margaritaceae (L.) B. & H., Antennaria media Greene, Achillea lanulosa Nutt,, Raillardella scaposa Gray, Arnica latifolia Bong., Senecio triangularis Hook., Agoseris alpestris (Gray) Greene.
134 MADRONO [Vol. 19
General Science Department, Portland State College, Oregon
LITERATURE CITED
HowE Lt, T. J. 1896. The flora of Mt. Hood. Mazama 1:28-48.
Lance, E. F. 1953. Thomas J. Howell, pioneer Oregon botanist. Sci. Month. 77:199-204.
Peck, M. E. 1961. A manual of the higher plants of Oregon. Binfords & Mort, Portland.
Wise, W. S. 1964. The geologic history of Mt. Hood, Oregon. Mazama 46:13-22.
DOCUMENTED CHROMOSOME NUMBERS OF PLANTS (See Madrono 9:257-258; 17:255. 1964.)
Astragalus sclerocarpus Gray. Nm = 11. Washington, Adams Co., Sand Dunes State Park. R. Spellenberg 1420, WTU.
Bomarea hirtella (HBK.) Herb. 2n = 18. Mexico, Chiapas, Municipio of Pueblo Nuevo Solistahuacan. S. S. Tillett 636-3, D.S. Counted by Marion S. Cave. Ap- proximately 5% of the pollen mother cells had 1 bridge with associated fragment, and 1 cell had 2 bridges and fragments, suggesting the presence of two paracen- tric inversions in this plant. This is apparently the first report of a tetraploid in the genus.
Cicendia quadrangularis (Lam.) Griseb. (Microcala quadrangularis (Lam.) Griseb.) n = 13. California, Contra Costa Co., Richmond, D. M. Post 125, UC; near Giant. D. M. Post 126, UC. The more familiar generic name Microcala is appar- ently invalid due to its being superfluous when published. Fig. 1 shows a micro- sporocyte at metaphase II of meiosis.
a e . of ; : aes 2 oy 5% race r,' reat .
Fics. 1, 2. 1, Cicendia quadrangularis, left; 2, Exacum affine, right.
Cleomella macbrideana Payson. 2n = 17;;. Idaho, Lehmi Co., 12 mi SE of Salmon. G. A. Mulligan and C. Crompton 2961, DAO. Counted by G. A. Mulligan.
Eriocaulon compressum Lam. n = 20. Florida, Levy Co., near Bronson, J. L. Wig- gins 19220, DS. Counted by Marion S. Cave.
Eriogonum alatum Torr. var. alatum. n = 20. Utah, Carbon Co., Tavaputs Pla- teau. N. H. Holmgren et al. 1961, NY, UTC; Kane Co., Kaiparowits Plateau. N.H. Holmgren et al. 2070, NY, UTC. Counted by J. L. Reveal.
E. anemophilum Greene. n = 20. California, Mono Co., Sonora Pass. J. L. Reveal & J. A. Reveal 491, UTC. Counted by J. L. Reveal.
E. arborescens Greene. n = 20. California, San Bernardino Co., Rancho Santa Ana Botanic Gardens, #1560. L. L. Kistler s.n., UTC. Cultivated. Counted by J. L. Reveal.
E. caninum (Greene) Munz. n = 12. California, Marin Co., Tiburon. D. L. Breed- love 4965, UTC. Counted by J. L. Reveal.
E. cinereum Benth. n = 40. California, Los Angeles Co., near Malibu. V. H. Holm- gren & J. L. Reveal 2616, NY, UTC. Counted by J. L. Reveal.
E. compositum Dougl. ex Benth. var. compositum. n = 20. Oregon, Wasco Co., Tygh Valley. P. H. Raven 18395, UTC. Counted by J. L. Reveal.
1967] DOCUMENTED CHROMOSOME NUMBERS 135
E.
ope Meola Sole les
covilleanum Eastw. n = 17. California, Kern Co., Temblor Range. E. C. Twissel- mann 13020, UTC; Fresno Co., Warthan Canyon. £. C. Twisselmann 13040, UTC; Fresno Co., Los Gatos Canyon, E. C. Twisselmann 13311, UTC; San Benito Co., Clear Creek, E. C. Twisselmann 13320 UTC; San Benito Co. near Idria. E. C. Twisselmann 13350, UTC. Counted by J. L. Reveal.
. eastwoodianum J. T. Howell. n = 17. California, Fresno Co., Coalinga-Park-
field Grade. E. C. Twisselmann 13109, UTC. Counted by J. L. Reveal.
. elatum Dougl. ex Benth. n = 20. Oregon, Wasco Co., near The Dalles. N. H.
Holmgren & J. L. Reveal 2116, NY, UTC. Counted by J. L. Reveal.
. fasciculatum Benth. var. fasciculatum. n = 20. California, Ventura Co., near
Malibu. N. H. Holmgren & J. L. Reveal 2117, NY, UTC. Counted by J. L. Reveal.
. flecum M. E. Jones. n = 20. Utah, Emery Co., near Ferron. N. H. Holmgren &
J. L. Reveal 1966, NY, UTC; Garfield Co., near Escalante. VN. H. Holmgren et al. 2045, NY, UTC; Garfield Co., near Boulder. VN. H. Holmgren et al. 2122, NY, UTC. Counted by J. L. Reveal.
. giganteum S. Wats. n = 20. California, San Bernardino Co., Rancho Santa Ana
Botanic Gardens, #697. L. L. Kistler s.n., UTC. Cultivated. Counted by J. L. Reveal.
. gossypinum Curran. n = 20. California, Kern Co., near Hart Park. E. C. Twissel-
mann 13307, UTC. Counted by J. L. Reveal.
. gracillimum S. Wats. n = 20. California, San Bernardino Co., Lucerne Valley.
N.H. Holmgren & J. L. Reveal 2594, NY, UTC. Counted by J. L. Reveal.
. inflatum T. & F. var. deflatum Johnst. n = 16. Mexico, Baja California, Angel
de la Guarda Island. R. Moran 12961. Counted by J. L. Reveal.
. jamesii Benth. in DC. var. undulatum (Benth. in DC.) S. Stokes ex M. E. Jones.
n = 20. Mexico, San Luis Potosi, near San Luis Potosi. D. B. Breedlove 15468, UTC. Counted by J. L. Reveal.
. latifolium Sm. n = 20. California, Marin Co., Pt. Reyes. VN. H. Holmgren & J.
L. Reveal 2641, NY, UTC; Sonoma Co., Ocean View, N. H. Holmgren & J. L. Reveal 2648, NY, UTC. Counted by J. L. Reveal.
. parishiti S. Wats. n = 20. Mexico, Baja California, near Rancho Marcos. R.
Moran 13478, UTC. Counted by J. L. Reveal.
. racemosum Nutt. n = 18. Nevada, White Pine Co., Lehman Creek, Snake Range.
D. B. Breedlove 5794, UTC; Nye Co., Scofield Canyon, Grant Range. NV. H. Holmgren et al. 2202, NY, UTC. Counted by J. L. Reveal.
_roeum Dur. & Hilg. n = 9. California, Tulare Co., Sequoia N. P., NV. H. Holm-
gren & J. L. Reveal 2941, NY, UTC; Kern Co., Piute Mts., N. H. Holmgren & J.L. Reveal 2943, NY, UTC. Counted by J. L. Reveal.
. salicornoides Gandg. n = 18. Idaho, Owyhee Co., near Marsing. N. H. Holmgren
& J.L. Reveal 866, NY, UTC. Counted by J. L. Reveal.
. saxatile S. Wats. n = 20. California, Los Angeles Co., Tujunga Creek. N. H.
Holmgren & J. L. Reveal 2613, NY, UTC. Counted by J. L. Reveal.
. Sphaerocephalum Dougl. ex Benth. var. sphaerocephalum. n = 20. Washington,
Grant Co., near Coulee City. P. H. Raven 18493, UTC. Counted by J. L. Reveal.
. temblorense Howell & Twisselmann. n = 17. California, Kern Co., Temblor
Range. E. C. Twisselmann 13110, UTC; San Luis Obispo Co., Cottonwood Pass. E. C. Twisselmann 13104, UTC. Counted by J. L. Reveal.
. thomasii Torr. n = 20. Mexico, Baja California, Los Angeles Bay. R. Moran
12423, UTC; Angel de la Guarda Island. R. Moran 12911, UTC. Counted by J. L. Reveal.
. umbellatum Torr. var. umbellatum. n = 40. California, Alpine Co., near Ebbets
Pass. D. B. Breedlove 5615, UTC; near Markleeville. D. B. Breedlove 5625, UTC; near Monitor Pass. D B. Breedlove 5627, UTC; Idaho, Boise Co., near Banks. P. H. Raven 18521, UTC. Counted by J. L. Reveal.
136 MADRONO [Vol. 19
E. vestitum J. T. Howell. n = 17. California, Fresno Co., near Mercey Hot Springs. E. C. Twisselmann 13359, UTC; San Benito Co., near Idria. E. C. Twisselmann 13349, UTC. Counted by J. L. Reveal.
Exacum affine Balf. f. n = 18. Cultivated plant from Missouri Botanical Garden. D.M. Post 208, UC. Fig. 2 shows a microsporocyte at metaphase II of meiosis. Fagonia laevis Standl. n = 10. Mexico, Baja California, near Cerro Blanco. J. L.
Wiggins & J. H. Thomas 142, DS. Counted by D. M. Porter.
Gentiana calycosa Griseb. n = 13. California, Tulare Co., Mineral King. D. M. Post 250, WC:
Hackelia cinerea (Piper) Johnst. 2n = 48, 2n = 241. Washington, Chelan Co., near Cashmere. R. Spellenberg 1405, WTU.
Holocantha emoryi Gray. 2n = 26. Arizona, Maricopa Co., Progeny of D. Verity in 1966, LA. Counted by P. H. Raven.
Hypericum formosum HBK. var. scouleri (Hook.) Coult. 2n = 8). California, Tuolumne Co., near Pinecrest. P. H. Raven 20807, DS. Counted by D. W. Kyhos.
Idahoa scapigera (Hook.) Nels. & Macbr. n = 8. Washington, Kittitas Co., near Ellensburg. R. Spellenberg 1637, WTU.
Orontium aquaticum L. n = 13. Florida, Alachua Co., near Gainesville, J. L. Wig- gins 19457, DS. Counted by Marion S. Cave.
Panicum capillare L. var. occidentale Rydb. 2n = 91. Idaho, Custer Co., Sunbeam Hot Springs, R. Spellenberg 317, WTU; California, Shasta Co., near Castle Crags. R. Spellenberg 1603, WTU.
Penstemon gairdneri Hook. n = 8. Washington, Douglas Co., Moses Coulee. R. Spellenberg 1400, WTU.
Schoenocaulon dubium (Mich.) Small. 2 = 8. Florida, Alachua Co., Gainesville, I. L. Wiggins 19866, DS. Counted by Marion S. Cave.
Smilacina flexuosa Bertol. 2n = 36. Mexico, Chiapas, Municipio of Pueblo Nuevo Solistachuacan. S. S. Tillett 636-5, DS. Counted by Marion S. Cave.
Taegetes filifolia Lag. n = 12. Ecuador, Quito. C. B. Heiser 6123, IND. Counted
by R. T. Neher. . micrantha Cav. n = 12. Mexico, Jalisco, 70 mi N of Guadalajara. R. T. Neher 1272, IND.
. multiflora HBK. n = 12. Ecuador, Prov. Pichincha, near Pifo. C. B. Heiser 5099, IND. Counted by R. T. Neher.
. subulata Cerv. 2n = 24. Mexico, Durango, 90 mi W of Durango. R. T. Neher 1262, IND:
. terniflora HBK. 2n = 48. Peru, Cuzco, C. B. Heiser 6241, IND. Counted by R. T. Neher.
Tribulus cistoides L. n = 6. Jamaica, St. Catherine, near Caymanas Park. D. M. Porter 1024, DS, GH, IJ; St. Andrew, St. Benedict’s School. D. M. Porter 1037, DS, GH, IJ; Columbia, Atlantica, Barranquilla. D. M. Porter 1117, COL, DS, GH; Mexico, Colima, near Tecoman, D. M. Porter 1483, GH, MEXU.
Se oa Sy. 2S
NOTES AND NEWS
XI INTERNATIONAL BOTANICAL ConGRESS—The XI International Botanical Congress will meet in Seattle, at the University of Washington from August 24 to September 2, 1969. Meetings of the Nomenclature Section will be held on August 21-24. Papers and symposia will be given in nine sections: molecular, metabolic, structural, devel- opmental, genetical and cytogenetical, environmental and evolutionary, systematic, ethnobotany, and historical botany. Pre- and post-Congress scientific field trips are planned. The first circular has already been issued and may be obtained from Dr. Richard S. Cowan, Secretary, XI International Botanical Congress, Smithsonian Institution, Washington, D.C. 20560.
VARIATION IN THE DIPLOID GOSSYPIUM SPECIES OF BAJA CALIFORNIA
LYLE L. PHILLIPS AND DUNCAN CLEMENT
The approximately 30 wild diploid (2n = 26) species of Gossypium occur in desert and steppe regions of the tropics and subtropics. Most of them are represented in experimental cultures and collectively have been the subject of considerable cytogenetic resarch. However, the ulti- mate source of seed for each of the diploid taxa can be traced to only one or two original collections and very little information is available on their morphological and genetic variability.
One of the two largest groups in the genus, in terms of number of taxa, occurs in the Americas from Peru and the Galapagos Islands to Mexico and Arizona. The center of known diversity for this geograph- ically and cytologically defined (D genome) assemblage of species is in western Mexico, where seven of the nine taxa are found. Three taxa, G. klotzschianum Anderss. var. davidsoni (Kell.) Hutch., G. armourianum Kearney, and G. harknessii Brandeg., are found in Baja California del Sur (fig. 1). Variety davidsoni also occurs in the vicin- ity of Guaymas, Sonora, Mexico.
In January and February of 1963 we spent approximately one month in the southern half of Baja California del Sur, making field studies and seed collections of G. klotzschianum var. davidsonu and G. harknessit. Seeds of G. armourianum, a species of northeastern Baja California del Sur, were collected by Duncan M. Porter of Harvard University. Plants from these seed collections were established and studied at the Uni- versity of the West Indies, Kingston, Jamaica, and at Raleigh, North Carolina. Herbarium specimens from the University of California, Stanford University, the California Academy of Sciences, and the U.S. National Herbarium were also studied. Recent collections of herbarium material of var. davidsonii from the northern part of its range (an area we did not bontanize) were made available to us by Miss Annetta Carter of the University of California. Contribution from the Crop Science Department, North Carolina Agricultural Experiment Station, Raleigh, North Carolina. Published with the approval of the Director of Research as Paper No. 2322 of the Journal Series. Aided by grant G-14203 of the National Science Foundation.
The variation listed and dscussed below for each taxon involved in this study is subdivided into “continuous” and “discontinuous” group- ings. To some extent this categorization is arbitrary and intuitive, e.g., variability for characteristics such as capsule loculus number, which varies within as well as between individuals and presumably is poly- genically controlled, is classed as continuous variation. Variability that can be graded on an absolute scale, e.g., gland red vs. gland colorless, is assumed to be simply inherited and is classed as discontinuous varia- tion. However, a majority of the variation classed as discontinuous has
137
138 MADRONO [ Vol. 19
Fic. 1. Distribution of G. armourianum (x), G. harknessi (closed circles), and G. klotzschianum var. davidsonii (open circles) in Baja California.
been thus categorized on the basis of monofactorial F. or testcross segregation frequencies of appropriate F; hybrids, and most of the variation described as continuous is so categorized because of blurred segregation patterns resulting from these breeding tests.
1967 | PHILLIPS & CLEMENT: GOSSYPIUM 139
TAXONOMY AND VARIATION
GOSSYPIUM KLOTZSCHIANUM Andersson var. DAVIDSONII (Kellogg) Hutchinson, Evol. Goss. 22. 1947. Gossypium davidsoni Kellogg, Proc. Calif. Acad. 5:82. 1873 Type: Cape San Lucas, Baja California, Xantus 12 (US!). Gossypium davidsonii var. sonoricum Mauer, Origin and Sys. Cot. 256. 1954. Type: Guaymas, Sonora, Mexico, Azerbaijan Cot. Res. Inst. 1562. Gossypium davidsonii var. lobulatum Mauer, Origin and Sys. Cot. 256. 1954. Type: San José del Cabo, Baja California, Mexico, Azerbaijan Cot. Res. Inst. 3552.
Diffusely branched shrub 1-3 m high; young stems punctate with reddish glands, strigose to tomentose with simple and stellate hairs, rarely glabrate; old stems sparsely strigose to glabrous; sympodia 2—5 jointed, rarely 6-12 jointed; leaves entire to 3-lobed, sub-oricular to ovate in outline, cordate at base, acuminate at tip, 5-10 cm long, stri- gose to tomentose on both surfaces, rarely glabrous, with nectary on abaxial surface of mid-vein, lamina and petiole punctate with reddish glands; stipules small, linear, early caducous; bracteoles cordate, 2—4 cm long, with 6—13 acuminate to obtuse teeth on upper margin; corolla cup-shaped, the petals yellow, usually with small dark-red spot near claw, occasionally spotless; androecium 15-20 mm long, the filaments 2-4 mm long; styles 5-15 mm long, stigmatic from tip to juncture with androecium; capsules 3—5 locular, spherical to fusiform, 15-20 mm long, 10-20 mm wide, usually mucronate, punctate with reddish glands, the sutures ciliate; seeds brown to black, angularly turbinate, with a sparse coating of appressed hairs except on conspicuous raphe.
Distribution. Sea level to ca. 400 m, in open situations, or as an ele- ment of thorn-scrub vegetation from extreme tip of Baja California north to vicinity of Bahia de la Concepcion (Lat 26.5°), including off- shore islands in the Gulf of California (Isla Cerralvo) and in the Pacific Ocean (Isla Magdalena) (fig. 1).
Hutchinson (1947) included the Revillagigedos Islands in the distri- butional range of var. davidsonii. For reasons previously detailed by Moran (1961), this is an error.
The type variety of G. klotzschianum is endemic to several islands of the Galapagos Archipelago. Variety klotzschianum and var. david- soni are morphologically similar, and their hybrid is easily synthesized and fully fertile. The distinguishing features of the two varieties are quantitative rather than qualitative, i.e., var. klotzschianum has larger leaves, bracteoles, flowers, etc., than var. davidsonii and var. klotzschi- anum develops into an open shrub whereas var. davidsonii is usually a compact bush.
The widely disjunct distributions of var. klotzschianum and var. davidsonii must be explained on the basis of relatively recent trans- Oceanic dispersal and Phillips (1966) has argued that the closer phyletic affinity of the two G. klotzschianum varieties to Mexican species than to the South American species (Peru) indicates that the dispersal was from north to south.
140 MADRONO [Vol. 19
® a he %
4 4 & ‘ a
Fic. 2. Bracteole, capsule, and leaf outlines for representative accessions of G. klotsschianum var. davidsonii (1 through 4738) and one accession of G. k. klotzschianum (S 33). Bracteole and capsule, X %; leaf, x 14%.
Mauer (1954) recognized G. davidsonii as a species and described the varieties sonoricum and lobulatum. Variety sonoricum is based on a specimen with entire leaves and 4-locular capsules; var. lobulatum is based on a specimen with lobed leaves and 5-locular capsules. Both leaf shape and capsule loculus number are variable throughout the range of var. davidsoni, and we regard these two epithets as synonyms of var. davidsonit.
Major variation for G. klotzschianum var. davidsonii is as follows:
1967 | PHILLIPS & CLEMENT: GOSSYPIUM 141
Continuous Variation
Pubescence. The entire plant is typically covered with simple and stel- late hairs, though these are semi-caducous on the stems. Our collections ranged from sparingly strigose to canescent. A collection by M. E. Jones (24210; US, CAS) is remarkable in being completely glabrous.
Leaf, Capsule, and Bracteole. Variation for leaf, capsule, and brac- teole characteristics in var. davidsonii is shown in Fig. 2. A majority of those accessions with lobed climax leaves were collected along the south- eastern coast of the Cape region.
Capsule loculus number for var. davidsonii ranges from 3 to 5. The average loculus number for the accessions of our study collection is 4.13 per capsule (computed on 10 capsules per accession), the extremes being 3.8 and 4.8.
Sympodial nodes. The number of sympodial nodes for a majority of our accessions ranges from 2 to 5 and averages about 4. The sym- podial node number for two accessions is outside this range; in one collection it ranges from 4 to 8, in the other from 8 to 12.
Discontinuous Variation
Anthocyanin coloration. The stems and petioles of var. davidsonii are typically “sun-red” owing to a moderate accumulation of antho- cyanin; this coloration is most intense at the juncture of the petiole and leaf lamina (pulvinus). In approximately one fourth of the plants grown from one of our accessions the stems and petioles are completely green; the F. progenies of crosses between red and green individuals segregated approximately 3 red : 1 green. Apparently the green indi- viduals are conditioned by recessive alleles at a single locus, and pre- sumably the plant from which this seed accession was obtained was heterozygous for this locus.
Foliar nectary. Variety davidsonii, and most other Gossypium di- ploids, characteristically have leaf nectaries near the pulvinus on the mid-vein. All of the plants grown from two seed collections lack the nectaries. F2 progenies of hybrids between plants of these accessions and plants with normal foliar nectaries segregate for presence and absence of foliar nectaries in proportions that provide a satisfactory fit to a 3 nectary : 1 nectariless ratio; it is assumed that nectariless is conditioned by recessive alleles at one locus.
Petal spot. The petals of var. davidsonii typically have a red spot, 3-7 mm in diameter, near the claw. One collection lacks this petal spot, and F. progeny tests of appropriate hybrids indicate that spotless is simply inherited and recessive.
Pigment glands. The aerial portions of all species of Gossypium pos- sess conspicuous glands which contain, among other constituents, gossy- pol. The capsules of var. davidsonii normally have 250-300 glands and the dry weight leaf content of gossypol is 1.8-2%. One of our accessions has a markedly reduced number of capsule glands (60-75) and the
142 MADRONO [Vol. 19
leaf gossypol level is reduced to 0.25%. Fe progeny tests involving hybrids between this accession and normally glanded individuals indi- cate a simple recessive genetic model to explain gland and gossypol reduction, though only one locus of a probable two-locus control of gland production is involved.
GOSSYPIUM HARKNESSII Brandegee, Proc. Calif. Acad. 2:136. 1889. Type: Santa Margarita Island, Baja California, Mexico, Brandegee s.n., March 1, 1889 (US! ). Gossypium californicum Mauer, Proc. Cen- tral State Univ. Tashkent, 18:21. 1950. Type: Cerro Colorado to Rod- riguez, Baja California, Mexico, Nelson & Goldman 7328 (US!).
Spreading to erect shrub 1-3 m tall with branches semi-decumbent and crooked to ascending and nearly straight; young stems puberulent, puctate with dark-red glands; old stems with conspicuous transversly elongated lenticels; sympodia usually 1-jointed, occasionally 2-jointed; leaves entire to shallowly 3-lobed, orbicular in outline, cordate at base, obtuse at tip, 5-7 cm long, glabrate on adaxial surface, puberulent on abaxial surface, the nectary on abaxial surface of mid-vein often ves- tigial, occasionally absent, lamina and petiole punctate with dark-red glands; stipules small, linear, caducous; bracteoles 1-3 cm long, entire or with 1-4 acuminate teeth at apex, caducous at anthesis; corolla cup- shaped, the petals yellow with small dark-red spot near claw; andro- ecium 10-15 mm long, the filaments 4-6 mm long, the lowermost pig- mented red; styles 3-4 cm long, sparsely ciliate, stigmatic only near tip; pollen usually yellow, occasionally cream; capsules nearly spherical, 15-20 mm in diameter, 3-4(—5) locular, punctate with dark-red glands, the sutures ciliate; seeds brown, 8-10 mm long, with a sparse coating of appressed hairs except on conspicuous raphe.
Distribution. From Lat 23.8° north to Lat 26.1°, including Isla Santa Margarita in the Pacific Ocean and Isla del Carmen, Isla Cor- onados, and Isla Monserrate in the Gulf of California (fig. 1).
Mauer (1950) distinguished G. californicum (the type specimen is from Isla Santa Margarita) from G. harknessii on the basis of its lobed leaves and beaked capsule. These two characteristics vary greatly throughout the range of G. harknessii, and we consider G. californicum to be a synonym of G. harknessit.
Major G. harknessi variability is summarized below:
Continuous Variation
Leaf, Capsule, and Bracteole (fig. 3). The G. harknessii leaf shape ranges from entire to moderately lobed; the former condition predom- inates in the northern part of the distributional range and the latter in the south.
Variation for capsule size and shape is slight; however, variability in locule number is evident and geographically correlated. Locule num- ber for those accessions from the southern part of the range is rather consistently 3, rarely 4. Accessions from Loreto and the offshore islands
PHILLIPS & CLEMENT: GOSSYPIUM
4
Fic. 3. Bracteole, capsule, and leaf outlines for representative accessions of G. harknessii (40 through 96) and one accession of G. armourianum (108). Brac- teoles and capsules, x 78; leaves, X ¥.
generally range from 3.2 to 3.5 (based on 10 capsules per accession) ; the capsules of an accession from Isla del Carmen average 3.9, and this accession has the only 5-locular capsules observed in the species (ot 110):
Foliar Nectary. The foliar nectary is vestigial (non-flowing) to absent in G. harknessu and variation for these two extremes is observed between accessions, between plants of the same accession and even between leaves of the same plant.
Sympodial nodes. The sympodium of G. harknessii is typically single- jointed; one accession, however, produced some 2-noded sympodia.
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Discontinuous Variation
Pollen color. Among a preponderance of accessions characterized by yellow pollen, there are 5 accessions, from all mainland and insular segments of the range, with cream pollen. F2 progeny tests involving three of these cream pollen mutants indicate that cream pollen is con- trolled by a monofactorial recessive.
Plant habit. As has been noted previously by Miss Annetta Carter (personal communication), there is an ecotype on Isla Coronados with a combination of characters not present in other populations. This eco- type has a low semi-spreading habit, thick, crooked branches and entire leaves. This complex of characters is suggestive of G. armourianum but other morphological features are those of G. harknessii (puberulent stems and petioles, long bracteoles, relatively large, deeply cordate leaves).
Non-flaring Capsule. The carpels of the G. harknessi capsule are normally fully reflexted, and the seeds are dspersed within a few days following capsule dehiscence. A frequent variant found on Isla Coro- nados is a type in which the carpels of dehiscent capsules are only mod- erately reflexed and from which the seeds escape over an extended period of time.
It is of interest to note that G. harknessit and G. armourianum are the only American diploids that do not have hard, impermeable seed coats; “hard” seed is presumably a device that tends to prevent seed germi- nation when the soil moisture level is insufficient for seedling estab- lishment. A mechanism that would ‘‘meter” the dispersal of water- permeable seeds to the substrate, and thereby protract the committment of seeds to possible germination over an extended period, would have an adaptive significance simlar to that of “hard”’ seed.
GOSSYPIUM ARMOURIANUM Kearney, Jour. Wash. Acad. 23:558. 1933. Type: Seeds from San Marcos Island, Baja California, Mexico; grown at Palm Springs, California, Kearney, August 25, 1933 (US!).
Spreading shrub 1-—2(-—3) m tall with decumbent to semi-erect branches; young stems glabrous, punctate with dark-red glands; old stems slightly furrowed, with obscure, circular lenticels; sympodia usually 1-jointed, occasionally 2-jointed; leaves entire, 2—4(—5) cm long, cordate to round-ovate, glabrous, puctate with dark-red glands, with nectary on abaxal surface of mid vein; stipules minute, linear, early caducous; bracteoles 2-5 mm long, entire or rarely with minute teeth at apex, caducous long before anthesis; corolla cup-shaped, the petals yellow with small dark-red spot near claw, 3-5 cm long; androecium 10-15 mm long, the filaments 3-5 mm long, pigmented red; styles 3-4 cm long, stigmatic only at tip; pollen yellow; capsules nearly spherical, 10-15 mm in diameter, 3(—4)-locular, punctate with dark-red glands, the sutures ciliate; seeds brown 7-9 mm long, with a sparse coating of appressed hairs except on conspicuous raphe.
Distribution. Isla San Marcos, Gulf of California, and near Punta
1967 | PHILLIPS & CLEMENT: GOSSYPIUM
Fic. 4. Leaf-shape variation in G. armourianum, X 1.
Trinidad (Lat 27.8°) and Bahia San Francisquito (Lat 28.4°) on main- land Baja California (fig. 1). Major G. armourianum variation is summarized below:
Continuous Variaton
Plant habit. Variants include low, decumbent, diffusely branched bushes 1 to 1.5 m tall to erect, open shrubs 2 to 3 m tall.
Anthocyanin coloration. Variation includes individuals that have essentially green vegetative parts to those that have a considerable accumulation of anthocyanin in stems, petioles, and leaf margins.
Leaf characteristics. Leaf shape in ths material ranges from broadly (and rather deeply) cordate to orbicular (and non-cordate), and leaf length ranges from | to 4 cm (fig. 4).
Discontinuous Variation
A single plant of one accession has colorless glands rather than the normal dark-red color. The dry weight leaf gossypol assay for this ex- ceptional plant is 0.05%, much less than the 0.65% determined for plants with typically colored glands.
G. armourianum and G. harknessii are generally similar in vegetative and floral morphology and are obviously more closely related to each other than to other species. Our study collections show each of these taxa to have considerable variability, some of which (for habit and leaf shape characteristics) blur traditionally cited distinctions between them (Kearney, 1933; Hutchnson, 1947). The two taxa do, however, have discontinuous ranges of variation for style ciliation, bracteole size, gen- eral plant pubescence, and lenticel size, shape, and number; we follow tradition in recognizing these taxa at the specific level.
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NOTES ON DISTRIBUTION AND VARIATION
The present distribution of the three Baja California wild cottons (fig. 1) idicates that the evolutionary lines that differentiated into these taxa were once sympatric over much of Baja California del Sur. As far as we can ascertain, var. davidsonii and G. harknessii are now sympatric only in a few square miles southeast of Bahia de la Paz. No evidence of hybridization involving these two taxa was observed in this area, and in fact, none was expected; it has been shown that var. davidsonii is cross-incompatible with most species of Gossypium, including all of the American diploids (Brown, 1951; Phillips, 1963).
The major distributional area of var. davidsonii is in the Cape region and is comprised of numerous, usually small (several to several hun- dred plants) populations that are often isolated from one another by considerable distances. However, we did encounter several large popu- lations, the largest of which includes an area of 75-100 square miles at the southwest tip of the Cape region. Other rather extensive popula- tions were found between San Bartolo and Loss Barriles and in the vicinity of Las Cruces.
Reflecting this population structure, as well as the self-compatibility and close juxtaposition of androecium and stigma in this species, the intra-population variation in var. davidsonii is very slight, whereas the inter-population variation is considerable. The only noteworthy intra- population heterozygosity detected by progeny testing of our var. davidsonii accessions was for anthocyanin coloration, and this collec- tion was taken from the large population in the Cape region. It should be stated that individual plants and populations of var. davidsonu were in a very depauperate state (apparently due to browsing by cat- tle and goats), making field analysis of variation next to impossible. None of the variation listed above for this taxon was noted in the field. Populations of G. hkarknessiu, in contrast, were unbrowsed and were amenable to field analysis of variation. The reason that G. harknessi is not browsed is not known, but one might speculate that an aromatic oil present in the leaves of G. harknessii (and G. armourianum), but not found in var. davidsoniu, renders the foliage of G. harknessi un- palatable to livestock.
In contrast to the population structure observed for var. davidsonit, small, spatially isolated populations seem to be the exception rather than the rule for G. harknessti. Northwest of Bahia de la Paz G. hark- nessit is a co-dominant element of a vegetation type that ranges over several hundred square miles. Immediately north of Loreto and on Isla del Carmen and Isla Coronados, there are large populations of G. harknessti; on Isla Coronados, which is composed primarily of broken lava, G. harknessiu is the dominant woody species.
Wide variation in vegetative and capsule characteristics (flowers were seldom seen) was evident in all populations of G. harknessii, though only
1967 | PHILLIPS & CLEMENT: GOSSYPIUM 147
on Isla Coronados was variation so canalized and non-random that ecotypes could be recognized. Progeny testing of our G. harknessii accessions revealed a high degree of heterozygosity for each of our single-plant seed collections. Such heterozygosity doubtless is correlated with the spatial separation of androecium and stigma in the species, which though self-compatible, will not set seed in the absence of pollen vectors.
Monographers of Gossypium (Hutchinson, 1947; Mauer, 1954) have cited G. armourianum as restricted to Isla San Marcos. A G. armouri- anum collection (J. N. Rose 16736) taken from the peninsula ca. 100 miles north of Isla San Marcos has apparently been overlooked (the collector identified the specimen as G. karknessit), and recently another peninsular collection of G. armourianum has been made by Reid Moran, this ca. 60 miles north of Isla San Marcos.
Our study material of G. armourianum was limited to three single- plant seed collections from a large population of Isla San Marcos. The progeny from each of these collections included a wide array of varia- tion for several characteristics, indicating that the individuals and pop- ulations of G. armourianum, like those of G. karknessii, contain consid- erable genetic heterozygosity.
Department of Crop Science, North Carolina State University, Raleigh National Science Foundation, San Jose, Costa Rica
LITERATURE CITED
Brown, M. S. 1951. The spontaneous occurrence of amphiploidy in species of Gos- sypium. Evolution 5:25-41.
HvurTcuinson, J. B. 1947. The classification of the genus Gossypium. In Hutchinson, J. B., R. A. Silow, and S. G. Stephens. The Evolution of Gossypium. Oxford Univ. Press, London.
KearNEY, T. H. 1933. A new species of Gossypium from Baja California. Jour. Wash. Acad. 23:558-560.
Mauer, F. M. 1950. Origin of cultivated and wild species of cotton and a basis for their classification. [In Russian.] Proc. Central State Univ. 18:7—21.
. 1954. Origin and systematics of cotton. [In Russian.] Tashkent.
Moran, R. 1961. Stegnosperma cubense and Gossypium davidsonii not known from Revillagigedos. Madrono 16:108.
Puitiips, L. L. 1963. The cytogenetics of Gossypium and the origin of New World Cottons. Evolution 17:460—469.
. 1966. The cytology and phylogenetics of the diploid species of Gossypium. Am. Jour. Bot. 53:328-335.
CALIFORNIA BOLETES III. THE GENUS SUILLUS
Harry D. THIERS
This genus of boletes is characterized by having carpophores with a viscid pileus (rarely dry, but if so with a gelatinous hypodermis), fascicled cystidia that stain dark brown to lavender or black when mounted in potassium hydroxide, relatively small, smooth-walled spores and divergent tube trama. An annulus or veil may or may not be present. A preliminary study of the North American species of Suillus by Smith and Thiers (1964) indicated that the genus was well repre- sented in California. Since that time intensive collecting has shown that the genus is among the larger, if not the largest, of the Boletaceae in the state.
Presented here are detailed descriptions of new and rare species, along with comments on distribution and other characteristics of species previously described in Smith and Thiers. All colors cited in quotes are those of Ridgway (1912), and all collections are deposited in the cryptogamic herbarium of San Francisco State College, San Francisco, California. This study was made possible by a grant from the National Science Foundation (GB 2760). It is a pleasure to express sincere appreciation for collecting permits issued by Burgess Heacox of the State Parks Department and by John O’Marie of the San Francisco Water Department.
KEY TO SUBGENERA
Pileus dry, or if viscid, then stipe annulate and lacking
glandular dots on the surface. . . . . . . . Subg. Boletinus Pileus subviscid to viscid to glutinous; if Shine is tla then glandular dots on the surface . . ....... .. . . Subg. Suillus
KEY TO SPECIES OF SUBGENUS BOLETINUS
Surface of pileus dry to moist, noticeably fibrillose to fibrillose-scaly (these char- acters not always apparent in very old pilei). Pileus reddish to reddish brown to orange-buff; fibrillose scales somewhat ap- pressed; viscid layer below scales well developed ._ . 10. S. lakei var. lakez Pileus brick red to vinaceous tawny; scales prominent and more or less erect; viscid layer be’ow scales often poorly deve'oped. 11. S. lakei var. pseudo pictus Surface of pileus visci to glutinous or at least with viscid scales or spots; glabrous to streaked or appressed fibrillose. Context of pileus changing to blue then fuscous when exposed; pores 1 mm or less in width . . . i eee 12. S. lithocar pi-sequoiae Context of pileus not changing to Wine ‘but may chomee to some other color, and flesh of stipe may change to blue); pores usually greater than 1 mm
broad. Annulus moist to subviscid, not glutinous, more or less fibrillose; pileus streaked to fibrillose. . . s-s = 6. S. caerulescens Annulus viscid to glutinous, thick, Gunes to picilone -orange; pileus glabrous to streaked 4.-. as 205 ~4e8) Gel Bee SIS MS pondernosus
148
1967] THIERS: SUILLUS 149 Key TO SPECIES OF SUBGENUS SUILLUS
Flesh changing to blue when exposed. .. . . . . . . 19. S. tomentosus Flesh unchanging when exposed or changing to some feolor other than blue. Veil or false veil present, either forming as annulus or existing as a roll of tissue on the pileus margin; (check young carpophores). Annulus typically present. Pores up to 5 mm broad, intervenose and often lamellate; annulus evanes- cent; pileus yellow to dark brown. Pileus bright yellow; stipe short, often eccentric. . . 13. S. megaporinus Pileus yellowish brown; stipe well developed, not eccentric. 18. S. riparius Pores not more than 2 mm broad; annulus not evanescent; pileus tan to pale brown or olivaceous. . .... . . . . 20. S. umbonatus Annulus typically absent. Pileus when young intensely yellow, often with red fibrils; glabrous to fibrillose; pores up to 15 mm broad. .. . . . 3. S. americanus Not as above. Pores large, often intervenose and lamellate; annulus evanescent. Pileus bright yellow; stipe short, often eccentric . . 13. S. megaporinus Pileus yellowish brown; stipe well developed, not eccentric. 18. S. riparius Not as above. Pileus white, gray or olive when young, becoming + ocharceous at maturity ; associated with Monterey and knobcone pines. 17. S. pungens Not as above. Pileus cinnamon brown to vinaceous cinnamon; stipe 2—4 cm. long, white when young, becoming ochraceous with age. 4.8. borealis Not as above. Glandulae prominent on stipe during all stages; associated with bishop or beach pine. . . .. . . . 8. 8S. glandulosipes Not as above. Pileus yellow to “ochraceous tawny” when young, often spotted or
mottled; spores 8-10 K 3-44... . 21. S. volcanalis Pileus white to pallid to pale vinaceous when young; spores 6.6—8.8 OUD S =O oer ek “aie, ee Gh Gee Oe te ee 1S ed ee LDA BES: Veil or false veil absent or rudimentary. Pileus conspicuously fibrillose scaly tosquamulose. . . .7.S. fuscotomentosus
Not as above. Stipe conspicuously glandular dotted. Context white during all stages, age ap when exposed; pores up to 2
mm broad. . . . . . 16. S. punctatipes Context yellow; pores ocally, ieee chan 2 mm broad.
Taste unpleasant; spores 9.3-12u long. . .. . . .1. S. acerbus
Taste mild; spores 7-9u long. . . . . . . . . . 9. S. granulatus
Stipe not conspicuously glandular dotted. Stipe clavate to venticose; yellow to olive buff to pale brown. 14. S. monticolus Stipe + equal, often short; white to pallid. . . . . .5. S. brevipes
1, SUILLUS ACERBUS Smith & Thiers, Contr. Monogr. N. Am. Species Suillus. 103. 1964.
Originally described from collections made in the Presidio, San Fran- cisco Co., Calif., this species has been found only in the greater San
150 MADRONO [ Vol. 19
Francisco Bay area. It apparently forms mycorrhizal associations with Monterey pine (Pinus radiata D. Don.) as does Suillus pungens Thiers & Smith with which it is frequently associated. However, S. pungens is distinguished by a cottony roll of veil tissue on the pileus margin and a pallid to gray color on the young pileus. Suillus acerbus seems most closely related to Suillus granulatus (Fr.) Kuntze but has larger spores and an unpleasant taste. The range of the two species apparently does not overlap since, so far as known, S. granulatus has not been reported from the Bay area.
2. SUILLUS ALBIDIPES (Peck) Singer, Farlowia 2:45. 1945.
As far as can be determined this species has not been reported pre- viously from the state. Collections have been made only in the vicinity of Huntington Lake in Fresno Co. As indicated by Smith & Thiers there has been considerable confusion concerning this fungus due to the fact that Peck apparently applied this name to two distinctly different fungi—one with an annulus and one without. In our concept S. albidipes should be restricted to those collections lacking an annulus but possess- ing a roll of veil tissue on the pileus margin. In so doing this distin- guishes it from S. granulatus with which it appears to be most closely related.
3. SUILLUS AMERICANUS (Peck) Snell in Slipp & Snell, Lloydia 7:39. 1944.
This well known species, although commonly found in the Pacific Northwest is apparently rare in California for it has been found only in the vicinity of Dinkey Creek near Huntington Lake in Fresno Co. The carpophores were typical and showed the distinctive characters associated with that species, including the yellow pileus with reddish streaks of fibrils, the distinct roll of veil tissue on the margin, the yellow pores and small stipe. They were growing in a coniferous forest composed of pines and firs.
4. SUILLUS BOREALIS Smith, Thiers & Miller, Lloydia 28:123. 1965. _ The type collection of this species was made in Idaho but collections have since been made throughout the Pacific Northwest. A single col- lection has been found in California in the vicinity of Willow Creek in Humboldt Co. The collection was made in a mixed forest which included pines, Douglas fir and oaks. This species is best characterized by the presence of a veil, dark colored pileus, a white stipe that slowly turns yellow, and obscure glandulae on the surface. It lacks a well de- veloped annulus.
5. SUILLUS BREVIPES (Peck) Kuntze, Rev. Generum Plantarum 3:535. 1898.
This species occurs throughout the state wherever two and three needle pines are found, and is one of the most abundant species. Most frequently it is associated with beach pine (Pinus contorta Doug.) in the coastal areas and lodgepole pine (P. murrayana Grev. & Balf.)-in alpine areas. Collec-
1967 | THIERS: SUILLUS 151
tions have also been made, however, where only ponderosa pine (P. pon- derosa Doug.) was present. Unforunately its name is somewhat misleading since carpophores with stipes better than 6 cm. in length have been col- lected; more commonly, however, the length is 2—4 cm. It is most easily recognized by the rather dark colored pileus with a glabrous margin, and the typically short stipe that is practically devoid of glandulae.
6. SUILLUS CAERULESCENS Smith & Thiers, Contr. Monogr. N. Am. Species Suillus. 36. 1964.
This suillus appears to be associated with Douglas fir in California and has been found wherever this conifer grows. It characteristically fruits early in the fall and is often very abundant. It is one of the few species with a viscid pileus in which the flesh of the stipe, but not of the pileus, turns blue when exposed. This reaction is sometimes spotty and relatively slow and ample time must be allowed for the change to occur. Some difficulty might be encountered in distinguishing it from S. ponderosus Smith & Thiers, but the latter is usually much more brightly colored and has a heavy annulus which is glutinous and bright orange, yellow-orange or tawny in color. S. caerulescens occurs throughout the Pacific Northwest.
7. SUILLUS FUSCOTOMENTOSUS Thiers & Smith, Contr.. Monogr. N. Am. Species Suillus. 65. 1964.
This very distinctive species was described from collections made in the sand hills area near Felton, Santa Cruz Co. It was commonly found in places where ponderosa pine and several different hardwoods were growing. Since that time, however, two additional localities have been established. One of these is in the Mount Shasta area at an altitude of 5—6,000 ft., and the other in the vicinity of Twain Harte, Tuolumne Co., at 3-4000 ft. elevation. In both localities the carpophores were obviously associated with ponderosa pine.
8. SUILLUS GLANDULOSIPES Thiers & Smith, Contr. Monogr. N. Am. Species Suillus. 86. 1964.
This is a relatively rare species in California. With the exception of a single collection made in Patricks Point State Park north of Eureka, Calif., it has only been found in Jackson State Forest near Mendocino, Calif. Although Smith & Thiers indicated that it was probably associ- ated with beach pine, additional observations seem to indicate that it is usually associated with bishop pine (Pinus muricata D. Don.), in California. Since collections have been made in such widely scattered places as Michigan, Oregon and Idaho it is obvious that it has a much wider range of mycorrhizal associates than indicated here in California. It is likely to be confused with S. granulatus since both species fruit simultaneously, but S. glandulosipes is distinguished by the distinct roll of veil tissue on the pileus margin, and paler, more vinaceous color of the pileus.
9. SUILLUS GRANULATUS (Fr.) Kuntze, Rev. Generum Plantarum 3:535. 1898. | :
152 MADRONO L Vol. 19
This is a very cosmopolitan species, having been reported from most of the pine forested regions of the United States, and needs little addi- tional comment. In California it has been found in the coastal forests as well as in the mountains, and is apparently associated with two and three needle pines.
10. SUILLUS LAKEI var. LAKEI (Murr.) Smith & Thiers, Contr. Monogr. N. Am. Species Suillus. 34. 1964.
This species is relatively common in the coastal forests of the state, but rare in the mountains where only one collection has been made. Like S. caerulescens it is associated with Douglas fir, and has been found throughout its range of distribution. Singer (1966, 1967) has indicated that this species is synonymous with S. amabilis (Peck) Singer, but there appears little justification for such an assumption as will be indicated by Smith and Thiers in a forthcoming paper. This is one of the few suilli with a typically dry pileus; however, care should be taken to examine young specimens since frequently in very old pilei the fibrillose scales will have disappeared and the glatinous or viscid layer will have become very pronuonced.
11. SUILLUS LAKEI var. PSEUDOPICTUS Smith & Thiers, Contr. Monogr. N. Am. Species Suillus. 33. 1964.
Only one collection of this variety has been made in California. It was growing in mixed woods in Jackson State Forest near Mendocino, Calif. As indicated by Smith & Thiers it differs from the typical variety by being darker red in color and having a more pronounced scaly pileus.
12. SUILLUS LITHOCARPI-SEQUOIAE Singer, Mycologia 51:859. 1959.
This is the only species, so far as known, that has been reported from the state that has not been seen by either Smith or Thiers. It was de- scribed by Singer from collections made in Muir Woods National Mon- ument in Marin Co., where it is apparently very rare since year around collecting has failed to discover it. That there is some confusion regard- ing its relationship with other suilli and other boletes is seen in the fact that Singer has recently transferred it to the genus Pulveroboletus (Singer, 1962). If, however, it has the characters elaborated by him, then it should remain in Swillus since it has fasciculate cystidia, small pores in connection with a viscid pileus and annulus. It appears to be the only species in the section Boletinus in which the flesh of the pileus consistently changes to blue when exposed.
13. SUILLUS MEGAPORINUS Snell & Dick, Mycologia 48:302. 1956.
Pileus 2-5 cm broad at maturity; globose to convex when young, becoming broadly convex to plano-convex to highly irregular with age; surface subvscid to viscid; streaked to fibrillose to appressed fibrillose scaly, scales occasionally becoming free when very old; background colored “warm buff” to “ochraceous buff”, scales colored near ‘‘cinna- mon buff” to “avellaneous”, sometimes becoming near ‘ochraceous tawny” when older; margin incurved, with obscure partial veil frag-
1967] THIERS: SUILLUS 153
ments when young, glabrous at maturity. Flesh 0.5—1 cm thick, colored “warm buff” to “ochraceous buff” to near “cinnamon buff,” unchang- ing when exposed; taste and odor not distinctive. Tubes depressed be- coming arcuate decurrent to strongly decurrent with age; up to 1 cm long; colored “antimony yellow” to “chamois,” sometimes assuming “tawny” to “russet” tints with age; unchanging when exposed; often appearing similar in arrangement to that of Gastroboletus tur- binatus ; pores up to 5 mm in length and 3 mm broad, intervenose, almost lamellate, with thick, wrinkled, irregular margins, concolorous with tubes, unchanging when bruised. Stipe 1-3 cm long, 4-7 mm broad at apex; often appearing short and reduced; sometimes somewhat ec- centric; pallid to more or less concolorous with the pileus; surface be- coming conspicuously glandular dotted at maturity; no annulus but obscure fibrillose annular zone sometimes apparent; flesh concolorous with that of the pileus, unchanging. Spores ellipsoid to subfusoid to subcylindric, thin-walled, hyaline in KOH, pale yellow in Melzer’s reagent, 8.5-10.5 & 4.5-6 mw; basidia hyaline in KOH, cylindric, 4-spored, 25-30 & 5-8 p; cystidia rare on sides of tubes, abundant to crowded on pores, fascicled, no solitary cystidia seen; dark brown in KOH, chocolate brown in Melzer’s reagent, cylindric to clavate, thin- walled, heavily incrusted, 30-48 * 7-11 pw; tube trama hyaline, ap- pearing subgelatinous in KOH, parallel to regular to obscurely diver- gent; hyphae 3—4 » in diam., pileus trama loosely interwoven, homogenous hyaline to pale brown in KOH;; cuticle differentiated as a la, er of appressed to interwoven hyphae, gelatinous to subgelatinous in KOH, hyaline to pale buff to pale brown in KOH, “ochraceous tawny” in Melzer’s reagent; hypoderm compactly interwoven, dark brown in KOH and Melzer’s re- agent; cuticle of stipe gelatinous in KOH, interwoven, with large clusters of caulocystidia staining dark brown in KOH, often with strong lavender tints, similar in shape and size to the cystidia on pores.
Habit, habitat and distribution. Gregarious to scattered in soil under conifers at edge of seepage areas. Carpophores usvally occurring in large numbers. Known only from the general vicinity of Huntington Lake, Fresno Co., Calif., Thiers 13423, 13440.
Discussion. At the time the monograph by Smith and Thiers was published fresh material of this species had not been seen. Since that time, however, collections have been made in the type locality. This is a highly distinctive suillus. The arrangement of the tubes is such that it appears either diseased or gastroid, but a spore deposit was obtained. The pores are larger than in any other suillus and are so long radially that they have the appearance of lamellae. The stipe is very reduced, short and somewhat eccentric. Like S. americanus it sometimes has reddish streaks or fibrils on the surface but it does not resemble that species otherwise. In Snell and Dick’s description (1956) they noted that an ample annulus was sometimes present. This was not apparent on any of the carpophores seen although a fibrillose annular
154 MADRONO [Vol. 19
zone could be detected on some. This species may be somewhat closely related to S. riparius Thiers since both species have rather large, inter- venose pores, but the latter has a well developed stipe, much darker col- ored pileus, with a rough, scaly surface, larger cystidia and a hymeni- form type of cuticle on the stipe.
14. Suillus monticolus Thiers, sp. nov. Pileus 4-7 cm latus, con- vexus demum plano-convexus, viscidus, appresso-fibrillosus, olivaceous vel cinnamomeus, in margine glaber, evelatus; tubuli 2—5 mm longi, straminei vel luridi; pori 1-2 mm lati; stipes 3-5 cm longus, 2-4 cm crassus, ventricosus basi bulbosus, siccus, glaber vel glandulosus, olivaceus demum cinnamomeus; sporae 8.5-11.2 « 3-4 p, cylindricae vel subellipsoideae; cystidia cylindrica vel subclavata, 25-45 « 4-9 p; cuticula subinnexa.
Holotype. Donner Summit, Nevada Co., California, elev. 7,200 ft., Sept. 22, 1965, Thiers 13248 (Herbarium of San Francisco State Col- lege).
Pileus 4-7 cm broad when mature; convex when young but becom- ing plane to plano-convex to broadly convex to highly irregular with age; surface viscid to subviscid; streaked to fibrillose to appressed fibrillose scaly; when young colored ‘‘clay color” to “pale olive buff” to “olive buff,” with age changing to “buffy brown” to dark “avellaneous” to near “cinnamon brown,” more or less evenly colored although the streaks or fibrils may be slightly darker; margin incurved becoming decurved with age, glabrous during all stages of development with no evidence of a partial or false veil. Flesh 0.5-1 cm thick, white, un- changing when exposed; taste and odor not distinctive. Tubes 2-5 mm long, adnate to shallowly depressed becoming decurrent to subdecur- rent with age; strongly radiately arranged; when young colored ‘straw yellow” to “amber yellow,” darkening with age to “antimony yellow” to “old gold”; pores 1-2 mm broad, often compound, concolorous with the tubes, unchanging when bruised, often staining vinaceous in KOH. Stipe 3-5 cm long, 2-4 cm broad at the apex; sometimes eccentric, distinctly ventricose to bulbous at the base and abruptly pinched below, usually bent, sometimes almost at right angles; surface dry, glabrous when young becoming glandulose to punctate to occasionally obscurely fibrillose with age; glandulae apparent only in the apical region; con- colorous with the tubes at the apex usually becoming colored near “olive buff” to “cinnamon brown” toward the base; frequently reticu- late at the apex; white mycelium at the base; no evidence of an annu- lus or partial veil; stuffed; flesh white, unchanging when exposed. Spores cylindric to subellipsoid, hyaline in KOH, pale yellow in Melzer’s reagent, smooth, thin-walled, 8.5-11.2 « 3-4 uw; basidia clavate to subcylindric, 2— and 4—spored, hyaline in KOH, 30-35 X 7-10 4p; cystidia clustered, often massed, particularly abundant on the pores, scattered to numerous on the sides of the tubes, no solitary cystidia seen, staining dark brown in KOH and Melzer’s reagent, usually stan-
1967 | THIERS: SUILLUS 15S
ing more darkly on the pores, cylindric to subclavate, 25-45 & 4-9 p; tube trama divergent from a distinct mediostratum, hyaline in KOH, appearing gelatinous to subgelatinous in KOH, hyphae 3-6 pw broad; pileus trama loosely interwoven, homogeneous, + hyaline in KOH; cuticle clearly differentiated as a trichodermum of + upright to some- what tangled hyphae with free tips, walls appearing finely incrusted, subgelatinous in KOH, staining chocolate brown in KOH, ochraceous in Melzer’s reagent; hypodermis compact, closely interwoven, staining brown to ochraceous in KOH, “ochraceous tawny” in Melzer’s reagent; surface of stipe hymeniform, with scattered clusters of caulocystidia similar to those on the pores; no clamp connections seen.
Discussion. The most distinctive feature of this suillus is the highly bulbous to ventricose stipe that is frequently bent and often somewhat eccentric. The glandulae, while always present, are never strongly de- veloped. The hymeniform cuticle of the stipe also appears highly dis- tinctive. It is gregarious in soil under lodgepole pines and is known only from the type locality.
15. SUILLUS PONDEROSUS Smith & Thiers, Contr. Monogr. N. Am. Species Suillus. 38. 1964.
This appears to be the largest species of suillus known from the United States and pilei up to 25 cm have been collected in California. So far it has been found only in the north coastal forests under Douglas fir where it frequently fruits with S. caerulescens. Its most distinctive features are the typically bright yellow or orange color of the rather heavy, glutinous annulus and the bright color of the often very large pileus.
16. SUILLUS PUNCTATIPES (Snell & Dick) Smith & Thiers, Contr. Monogr. N. Am. Species Suillus. 94. 1964.
A single collection of this species has been made in the vicinity of Donner Pass in Nevada Co. and, as far as known, this is the first report of its occurrence within the state. It is most probably associated with lodgepole pine but the carpophores were collected in a mixed conifer forest including other species of pines and firs. The relatively large pores that are radially arranged and the white flesh of the pileus dis- tinguish it from S. granulatus with which it might be confused.
17. SUILLUS PUNGENS Thiers & Smith, Contr. Monogr. N. Am. Species Suillus. 92. 1964.
This is the most common suillus in the San Francisco Bay area and typically fruits under Monterey pine, but collections have also been made under knobcone pine (Pinus attenuata Lemmon). The type col- lection was made on the campus of San Francisco State College where it occurs in abundance during the fall season. It is distinguished by the white roll of veil tissue on the margin, the unpleasant taste and the white to gray or olive color of the pileus when young. It has been mis- identified as S. placidus (Bonorden) Singer because in some instances
156 MADRONO [Vol. 19
the entire carpophore is almost pure white when young and has con- spicuous drops of milky exudate on the pores. However, it soon becomes colored and does not show any additional characteristics of S. placidus.
18. Suillus riparius Thiers, sp. nov. Pileus 5-12 cm latus, convexus vel plano-convexus vel planus, viscidus, glaber vel fibrillosus, brunneus vel ochraceus, velo luteo, floccoso in margine; tubuli decurrentes, 1.5 cm longi, lutei vel brunneoli; pori 4 mm longi, 3 mm lati; stipes 3-8 cm longus, 0.5—-1.5 cm crassus, aequalis, subviscidus, glaber, flavus; sporae cinnamomeae, subcylindricae vel subfusoideae, 8—-11.5 *& 3.2—-5 uw; cys- tidia in fasciculis, cylindrica vel subfusoidea, 35-75 & 5-10»; cuticula innexa.
Holotype. Pinecrest, Tuolumne Co., California, elev. 5,600 ft., Sept. 28, 1965. Thiers 13283 (Herbarium of San Francisco State College).
Pileus 5-12 cm broad at maturity; convex to obtusely conic when young becoming plano-convex to plane to broadly convex to shallowly depressed with age, often highly irregular and undulating; surface viscid, when young smooth to glabrous to subglabrous, occasionally somewhat streaked to appressed fibrillose, typically becoming somewhat rimose_ to areolate to fibrillose-scaly, occasionally in very old pilei appearing somewhat squarrose scaly; when young colored “buckthorn brown” to “dresden brown”’ to occasionally “prouts brown,” except for a cottony roll on the margin colored “antimony yellow” to “yellow ocher,” scales colored “‘prouts brown” to “cinnamon brown”’ to occa- sionally ‘“‘ochraceous tawny’; with age pileus “‘yellow ocher’’ to “old gold” to sometimes “naples yellow’” to “straw yellow” to occasionally “ochraceous tawny” to ‘‘ochraceous buff” when very old; scales typically unchanging or sometimes becoming pale vinaceous when bruised; margin when young with a pronounced, large roll of veil tissue which fades to “pale olive buff” with age, either appendiculate or glabrous to eroded with age. Flesh up to 1 cm thick, soft, colored “light ochraceous buff” to “massicot yellow” to ‘naples yellow,’ changing to ‘“‘avellaneous”’ to “drab” after exposure; taste acid, odor not distinctive. Tubes bluntly adnate when young becoming arcuate decurrent to decurrent with age; up to 1.5 cm in length; when young colored ‘mustard yellow” to “amber yellow,” changing to “antimony yellow” to “yellow ocher” with age, sometimes becoming brownish with age, unchanging upon exposure; pores distnctly compound, somewhat intervenose, up to 4 mm in length, 3 mm in width; margin highly uneven, concolorous with tubes, unchanging when bruised. Stipe 3-8 cm long, 0.5-1.5 cm broad at the apex; more or less equal; surface moist to subviscid, glabrous except for strongly developed, elongated, irregularly shaped glandulae which are colored ‘sudan brown” to “amber brown’’; surface colored “amber yellow” to “‘massicot yellow” to “mustard yellow,” often dark- ening slightly toward the base, frequently with a slight vinaceous tint at the base; stuffed to hollow; flesh concolorous with that of the pileus, unchanging when exposed; no annulus but fibrillose zone sometimes
1967 ] THIERS: SUILLUS 157
apparent. Spores in mass colored “cinnamon brown” to near “mars brown” to “dresden brown.” Spores subcylindric to subfusoid, hyaline to pale ochraceous in KOH, pale ochraceous in Melzer’s reagent, smooth, thin-walled, 8-11.5 & 3.2—5 mw; basidia clavate to subcylindric, 2— and 4-spored, hyaline in KOH, 21-29 X 8-10 py; cystidia clustered, scat- tered to numerous, more abundant on the pores, staining dark brown in KOH and Melzer’s reagent; sometimes discolored only in base, in- crusted with incrustations floating free in mounting medium, cylindric to subclavate to subfusoid, thin-walled, occasionally hyaline in KOH, 35-75 & 5-10 pw; tube trama hyaline in KOH, divergent from an in- distinct mediostratum, subgelatinous in KOH, hyphae 3-5 yp in diam.; pileus trama interwoven, homogeneous, hyaline in KOH; hypodermis compactly interwoven, well differentiated, brown in KOH, “ochraceous tawny” in Melzer’s reagent; cuticle differentiated as a layer of inter- woven to tangled hyphae partially gelatinizing in KOH; staining brown in KOH and + “ochraceous tawny” in Melzer’s reagent, finely in- crusted, hyphae 3—4 » in diam.; cuticle of stipe hymeniform with large clusters of fascicled cystidia similar to those on the tubes, dark red- dish brown in KOH; no clamp connections seen.
Habit, habitat and distributon. Gregarous to cespitose in vicinity of dead logs and stumps near edge of streams with ponderosa and sugar pines in general area. Pinecrest and Dodge Ridge Ski area, Tuolumne Co., California, Thiers 13283, 13350, 13351, 13883, 13885, 13889.
Discussion. This species occurs in large numbers along the edges of mountain streams, usually on or very near rotten stumps and logs. It is somewhat suggestive of S. americanus but differs in that the pileus is dark brown, and the stipe has dark brown glandulae. In addition the stipe is characteristically stouter than that reported for S. amer- tcanus, and the pores are yellow-brown. It might also be confused with S. megaporinus but the dark color of the pileus and well developed stipe make it distinct.
19, SUILLUS TOMENTOSUS (Kauff.) Singer, Snell & Dick, Mycologia 51:570. 1960.
This very abundant and easily recognized species occurs throughout the state in association with either beach or lodgepole pine. In a collec- tion made in Nevada Co. the young pilei were densely covered with gray to olive colored fibrils which in older pilei changed to very bright red. Some fibrils commonly become disclosed, but it is unusual to find carpophores in which the fibrils are so intensely colored that the pileus is distinctly red overall.
20. SUILLUS UMBONATUS Dick & Snell, Mycologia 52:446. 1960.
As has been previously indicated in Smith & Thiers (1964) and Smith, Thiers and Miller (1965), there is some confusion surrounding this species and its relationship with S. sibiricus and S. americanus. Close comparisons between the three have been made, and it is now generally agreed that S. umbonatus characteristically possesses a dis-
158 MADRONO [ Vol. 19
tinct annulus, is colored pale watery brown to tan, lacks any reddish discoloration or fibrils on the pileus and shows no vinaceous discolora- tion in the flesh of the base of the stipe. S. szbiricus is characterized by lacking an annulus, having a much brighter and intensely yellow pileus and vinaceous discoloration in the flesh in the base of the stipe. S. americanus, although similar in color to S. sibiricus, characteristically has reddish streaks or fibrils on the pileus and a slender stipe.
Collections of S. umbonatus have been made annually in Jackson State Forest near Mendocino. The carpophores have always been found in mixed forests where both conifers and hardwoods were present, making it impossible to determine a specific mycorrhizal associate. A large collection was also made at Yuba Pass, elevation 6,700 ft., where the fruiting bodies appeared most intimately associated with alder but firs and pines were also in the vicinity.
21. Suillus volcanalis Thiers, sp. nov. Pileus 9-12 cm latus, con- vexus, appresso-fibrillosus, viscidus, luteus vel luridus, velo albo floccoso, conspicuo in margine; tubuli depressi, lutei, 1—1.5 cm longi; pori angulati minus quam 1 mm lati; stipes 3-6 cm longus, apice 1.5—3 cm crassus, aequalis vel ventricosus basi, solidus, non annulatus, albus, siccus, glaber, demum glandulosus, fulvus; sporae 6.6-10 «3-4 uw; cylindricae vel sub- ellipsoideae; cystidia in fasciculis, cylindrica vel subclavata, 24-47 « 5-8 mw; cuticula innexa.
Holotype. Butte Lake Campground, Lassen Volcanic National Park, Lassen Co., California., elev. 6,000 ft., June 27, 1965, Thiers 12800 (Herbarium San Francisco State College).
Pileus 9-12 cm broad when mature; convex to irregular when young becoming plano-convex to highly irregular with an undulating margin to plane with an uplifted margin when older; surface often irregularly pitted; appearing closely appressed fibrillose to streaked; viscid; when young colored “primrose yellow” with numerous areas colored “ochra- ceous buff” to “ochraceous tawny,” when older frequently with some areas colored “‘picric yellow” and others ‘“‘vinaceous rufous,” in some the background colored ‘antimony yellow” to “warm buff” with irregu- lar areas colored ‘“‘testaceous”, very old carpophores colored “russet”’ to “tawny” with paler yellowish areas intermixed; margin strongly in- curved when young, usually with a white to pallid, conspicuous roll of partial veil tissue attached, becoming decurved and glabrous with age. Flesh 1-2 cm thick, white to ‘“‘pale ivory yellow’ when young typically becoming ‘‘massicot yellow” in older pilei, unchanging when exposed; taste and odor not distinctive. Tubes shallowly to deeply depressed during all stages of development; 1-1.5 cm long; when young colored “straw yellow” becoming ‘aniline yellow” when older; unchanging when exposed; pores angular, less than 1 mm broad, con- colorous with tubes; unchanging when bruised. Stipe 3-6 cm long, 1.5—3 cm broad at apex; equal to occasionally somewhat ventricose to flared at the base; surface dry, appearing glabrous to smooth; when
1967 | THIERS: SUILLUS 159
very old, glandulae are apparent at least in the apex; when young white except pale yellow at the apex, when older becoming ‘‘massicot yellow”’ at the apex and “buckthorn brown” to “ochraceous tawny” to “tawny” toward the base; dry; no annulus; solid; flesh white when young, yellowish with age; unchanging when exposed. Spores in mass colored “sudan brown” to “sayal brown.” Spores cylindric to subellipsoid, hyaline in KOH, pale yellow in Melzer’s reagent, smooth, thin-walled, 6.6-10 & 3-4 m»; basidia hyaline in KOH, clavate, 4-spored, 25-31 x 7-11 »; cystidia rare to absent along sides of tubes, scattered to relatively abundant on pores, fascicled, no solitary cystidia seen, dark brown in KOH, thin-walled, bases of many appearing incrusted, cylin- dric to subclavate, “ochraceous tawny” in Melzer’s reagent, 24-47 x 5-8 »; tube trama hyaline, divergent from a distinct mediostratum, appearing subgelatinous in KOH, hyphae 3-5 p» in diam.; pileus trama with a compactly interwoven layer near tubes which stains dark brown in KOH, surmounted by a layer of loosely interwoven, more or less homogeneous hyphae; cuticle composed of a narrow layer of closely interwoven hyphae that appear to be heavily incrusted and stain dark brown in KOH surmounting a broad layer of interwoven hyphae, 5-6 p» in diam., which appear gelatinous in KOH, dark “ochracaeous tawny” in KOH; no clamp connections seen; cuticle of stipe interwoven, subgelatinous in KOH, occasional clusters of cystidia similar to those on the pores.
Habit, habitat and distribution. Gregarious to scattered (often buried) in cinders and humus under Jeffrey pines (Pinus jeffreyi Grev. & Balf.), Butte Lake Campground, Lassen Volcanic National Park, Lassen Co., California, elev. 6,000 ft., Thiers 12800, 12777, 12915, W930, 12937.
Discussion. This species was most often found by uncovering “humps” in the soil under Jeffrey pine. Without exception it was found in dry cinder cone areas. It is somewhat remindful of S. brevipes but is read- ily distinguished by the distinct cottony margin of fibrillose tissue, the yellow to tawny color of the pileus which often appears distinctly fibrillose rather than glabrous and the much darker colored stipe. The spores are also longer and broader than those of S. brevipes.
DISCUSSION
An analysis of the state flora shows that twenty species and one vari- ety have been collected. Seven of these are more or less cosmopolitan, having been found throughout most of the United States; six others appear to be limited to the Pacific Coast, while eight of them seem to be restricted to California. Thus more than a third of the species may be endemic, which follows the pattern of a high percentage of endemism observed in other plants within the state.
The total of twenty species equals or exceeds the number of species reported from other regions except for the Pacific Northwest (Smith and Thiers, 1964), which has a known flora of 23 species. The absence
160 MADRONO [Vol. 19
of Larix in California may be a major factor in this difference in num- bers. Michigan and the northeastern United States also have approxi- mately twenty species (Smith and Thiers, 1964). The California flora is far more varied than that of the southeastern United States since only twelve species have been reported from North Carolina (Coker and Beers, 1943), nine from Texas (Thiers, 1959), and five from Florida (Singer, 1945).
As is generally known, most, if not all, species of Suillus are associ- ated with conifers. If S. lithocarpi-sequoiae is associated with Litho- carpus it will constitute the only known exception in the state. Suillus caerulescens, S. laket, and S. ponderosus are associated with Douglas fir and all of the other species are apparently associated with pines. Along the coast beach pine and Monterey pine are most commonly in- volved. In the mountains, lodgepole pines are most frequently involved in mycorrhizal associations. No suilli have been found associated with any of the white pines, but such associates are known in other parts of the United States and probably occur here. Likewise no species have been found associated with either of the redwoods or with Sitka spruce.
Since most of the conifers occur either along the coast or in the mountains, species of Suzllus occupy a similar distribution. So far eight species appear to be confined to the coastal forests, six to the moun- tains and six occur in both areas. It is interesting to note that in both areas the fruiting period occurs in the fall with practically no carpo- phores appearing in the spring. In the mountains if late summer rains occur, species of Suillus frequently appear in abundance in late August or early September. Generally the fruiting in the coastal areas occurs within a week or two after the onset of the rainy season. Most all species fruit at approximately the same time but lakez, caerulescens, and pungens have a considerably longer fruiting period than the others.
Department of Biology, San Francisco State College
LITERATURE CITED
Coxer, W. C., and A. H. Beers. 1943. The Boletaceae of North Carolina. Univ. of North Carolina Press, Chapel Hill. Ripeway, R. 1912. Color standards and nomenclature. Washington, D.C. SINGER, R. 1945. The Boletinae of Florida with notes on extralimital speceis. II. The Boletaceae (Gyroporoideae). Farlowia 2:223-303. . 1962. The Agaricales in modern taxonomy. Ed. 2. Weinheim. . 1966. The type of Boletus amabilis. Mycologia 58:157-159. ————. 1967. Die Rohrlinge, Teil II. Klinkhardt, Bad Heilbrunn. SmitH, A. H., and H. D. Turers. 1964. A contribution toward a monograph of North American species of Suillus. Ann Arbor. .. and O. Mirtrer. 1965. The species of Suillus and Fuscoboletinus of the Preist River Experimental Forest and vicinity, Priest River, Idaho. Lloydia 28:120-138. SNELL, W. H., and E. A. Dick. 1956. Notes on Boletes. IX. Mycologia 48:302—310. TureErs, H. D. 1959. Notes on the genus Boletus in Texas. Tex. Jour. Sci 11:314- 319.
ADRONO
VOLUME 19, NUMBER 5 JANUARY, 1968
Contents
CHROMOSOME VARIATION IN ECOTYPES OF LARREA DIVARICATA IN THE NorTH AMERICAN DESERT, Tien Wei Yang and Charles H. Lowe 161
CYTOLOGICAL AND EVOLUTIONARY STUDIES ON PALUSTRES VIOLETS, Marja Sorsa 165
A SECOND SPECIES OF AMMOBROMA (LENNOACEAE), IN SmnALoA, Mexico, Robert L. Dressler and Job Kuijt 179
CHROMOSOME NUMBERS IN THE TRIBE PAPPOPHOREAE (GRAMINEAE), John R. Reeder and D. N. Singh 183
SOME ADDITIONS TO THE FLORA oF TExAS—IV,
boa Donovan S. Correll 187 NOTES AND NEws: NOTES ON THE FLORA OF ARIZONA, IV, Charles T,. Mason, Jr. 164 LIBRARY MAILING RATES FOR HERBARIUM SPECIMENS, John H, Thomas 192
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BOARD OF EDITORS
LYMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. Ernst, Smithsonian Institution, Washington, D.C. ARTURO GOMEZ PompPa, Universidad Nacional Autonoma de México EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia Mitprep E. Matuias, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley Marion OwnsBEy, Washington State University, Pullman REED C. Ro.tiins, Harvard University, Cambridge, Massachusetts Ira L. WiccIns, Stanford University, Stanford, California
Editor—JoHN H. THoMAsS Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—J UNE McCAsKILL P.O. Box 23, Davis, California 95616
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Elizabeth McClintock, California Academy of Sciences, San Fran- cisco. First Vice-President: Peter H. Raven, Department of Biological Sciences, Stanford University. Second Vice-President: Cornelius H. Muller, Department of Biological Sciences, University of California, Santa Barbara. Recording Secretary: John West, Department of Botany, University of California, Berkeley. Correspond- ing Secretary: Mary E. Ashton, Department of Botany, University of California, Berkeley. Treasurer: June McCaskill, Department of Botany, University of Cali- fornia, Davis.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Carl W. Sharsmith, Department of Bio- logical Sciences, San Jose State College, San Jose; the Editor or Madrofio; and three elected Council Members: Virginia M. Page, Department of Biological Sci- ences, Stanford University; Annetta Carter, Department of Botany, University of California, Berkeley; and Robert Ornduff, Department of Botany, University of California, Berkeley.
CHROMOSOME VARIATION IN ECOTYPES OF LARREA DIVARICATA IN THE NORTH AMERICAN DESERT
TIEN WEI YANG and CHARLES H. LOWE
A recent report has suggested the presence of a difference in chromo- some number between populations of creosotebush, Larrea divaricata Cav., in the Chihuahuan Desert and in the Sonoran Desert (Yang, 1967). Chromosome counts from additional populations further confirm the hypothesis that 2n = 52 is associated with the western Larrea of the Sonoran Desert subdivision and 2n = 26 characterizes the eastern Larrea of the Chihuahuan Desert subdivision of the North American Desert. Chromosomes of flower buds and root tip cells were examined using aceto-carmine and aceto.orcein squashes.
Table 1 gives a summary of the chromosome numbers of seven new populations recently examined, as well as the earlier reports on the chromosome counts of six other North American creosotebush popula- tions (Covas, 1949; Powell, 1966 (pers. comm.); Yang, 1967). Figure 1 shows that all 13 populations which have provided this chromosome information are located within a distance of 100 miles either north or south of an 800-mile transect between Val Verde Co. Texas (east), and Imperial Co., California (west). This transect traverses essentially the entire longitudinal extent of the Chihuahuan Desert and the Sonoran Desert along the Mexican boundary. The transect also constitutes the only continuous east-west span of Larrea distribution in North America (Shreve, 1940; Garcia, Soto, and Miranda, 1960).
Nine of the populations have counts of n = 26 or ca. 26 and all are located well within the boundary of the Sonoran Desert (Shreve, 1942, 1951; Benson and Darrow, 1954; Lowe, 1964). Three of the four popu- lations with n = 13 or ca. 13 occur in Trans-Pecos Texas and are within the Chihuahuan Desert proper. The remaining population with the same chromosome number (n = 13) is located immediately west of the San Pedro Valley in southeastern Arizona, where the westernmost extension of the Chihuahuan Desert flora occurs (Benson and Darrow, 1954; Lowe, 1964). Between this Pima Co. population of Arizona and the Val Verde Co. population of Texas, covering a distance of 550 miles along the 800-mile transect, all the known chromosome counts have been in the order of n = 13 or ca. 13 and 2n = ca. 26. Between the easternmost Sonoran Desert population at 15 miles southeast of Tucson and the west- ernmost of the Imperial Co. populations of California, a span of 250 miles, all the known chromosome counts have been in the order of n = 26 or ca. 26 and 2n = 52 orca. 52 (Table 1).
The difference in chromosome number between the eastern creosote- bush and the western creosotebush appears to be associated with the major
MaprONO, Vol. 19, No. 5, pp. 161-192. April 5, 1968.
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162 MADRONO [Vol. 19
TABLE 1. SUMMARY OF CHROMOSOME NUMBERS OF 13 LARREA POPULATIONS ALONG AN 800-MILE TRANSECT IN THE SONORAN AND CHIHUAHUAN SUBDIVISIONS OF THE NortH AMERICAN DESERT (see Fig. 1).
Population Locality Chromosome Count 1. California 30 mi (US 80) W Colorado n = ca. 26 Imperial Co. R. In = ca, 52 2. California 20 mi (US 80) W Colorado n = 26 (Yang, 1967) Imperial Co. R. 2n = ca. 52 (Yang, 1967) 3. Arizona 10.8 mi (Ariz 84) W Pinal n = ca. 26 Maricopa Co. Co. line Qne== "cape 4. Arizona 0.1 mi (Ariz 84) W Maricopa n= ca. 26 Pinal Co. Rd. In = ca. 52 5. Arizona 0.4 mi (Ariz 86) E Sells n = 26 (Yang, 1967) Pima Co. 2n = ca. 52 (Yang, 1967) 6. Arizona Boyce Thompson SW Pinal Co. Arboretum 2n = 52 (Covas, 1949) 7. Arizona Tucson n = ca. 26 Pima Co. 2n = ca. 52 8. Arizona 15 mi (US 80) SE Tucson n = ca. 26 Pima Co. 2n = ca. 52 8. Arizona Saguaro Nat. Monument, n = ca. 26 Pima Co. Rincon Mts. Inca on 10. Arizona 16.6 mi (US 80) W San n= 13 Pima o. Pedro R. 2n = ca. 26 11. Texas Alpine n = ca. 13 (Powell, 1966, pers. Brewster Co. comm.) 12. Texas 65.1 mi (Tex 118) S Alpine n = 13 (Powell, 1966, pers. comm. ; Brewster Co. Yang, 1967) 2n = ca. 26 (Yang, 1967) 13. Texas 1.0 mi (US 90) W Pecos R. n = 13 (Yang, 1967) Val Verde Co. 2n = ca. 26 (Yang, 1967)
ecotypic differences in morphological and physiological traits reported earlier (Yang, 1967a). The Chihuahuan Desert Larrea, the eastern eco- type, is distinguished by a bushy and more compact growth form, dis- tinctly more incurved leaflets, more rapid germination rate, faster initial growth rate of the seedling, greater tolerance for low temperature, and lesser tolerance for low moisture. The Sonoran Desert Larrea, the west- ern ecotype, on the other hand, has a more erect growth form, less in- curved leaflets, slower germination rate, slower initial growth rate of the seedling, lesser tolerance for low temperature, and greater tolerance for low moisture.
The morphological and physiological differences between the eastern and the western Larrea reflect the diverse climatic regimes under which the two major populations have been evolving. The Chihuahuan ecotype is adapted to a climate of greater precipitation and relatively low winter temperature; the Sonoran ecotype had developed under a macroclimate of lower rainfall and higher winter temperature. The cytological differ- ence in chromosome number between these two ecotypes appears to be the result of the interaction of the same genetic and ecological processes
1968] YANG & LOWE: LARREA DIVARICATA 163
i i 4 i 1 fase ' ' ] i) | (
Oy a a a a O 6,
Nf dao yall
Fic. 1. Creosotebush populations studied along an 800-mile transect. Popula- tions 1-9 are situated in the Sonoran Desert and are tetraploid. Populations 10-13 are located in the Chihuahuan Desert and are diploid (see text). Shaded area shows the generalized distribution of Larrea divaricata in North America, adapted from Shreve (1940), Garcia, Soto, and Miranda (1960), and modified by unpublished data from the authors.
which have brought about the series of morphological and physiological variations in the evolution of the North American creosotebush.
The authors are grateful to Walter V. Brown for suggestions, to James L. Patton for assistance in the preparation of the illustration, and to A. M. Powell for providing unpublished data. Voucher specimens are de- posited at University of Texas Herbarium, University of Arizona Herbarium, and the Ecology Herbarium in the Department of Biologi- cal Sciences, The University of Arizona.
Department of Botany, The University of Texas, Austin Department of Biological Sciences, The University of Arizona, Tucson
164 MADRONO [Vol. 19
LITERATURE CITED
Benson, L., and R. A. Darrow. 1954. The trees and shrubs of the southwestern deserts. Univ. Ariz. Press, Tucson, and Univ. New Mex. Press, Albuquerque.
Covas, G. 1949. Estudios cariologicos en antofitas. III Darwiniana 9:158-162.
Garcia, E., C. Soro, and F. Miranpa. 1960. Larrea y clima. Anal. Inst. Biol. México. 31:133-190.
Lowe, C. H. 1964. Arizona landscapes and habitats. Jn The vertebrates of Arizona. Univ. Ariz. Press, Tucson.
SHREVE, F. 1940. The edge of the desert. Year. Assoc. Pacif. Coast Geogr. 6:6-11.
. 1942. The desert vegetation of North America. Bot. Rev. 8:195—246. . 1951. Vegetation and flora of the Sonoran Desert. Carnegie Inst. Wash. Publ. 591.
Yanc, T. W. 1967. Chromosome numbers in populations of creosotebush (Larrea divaricata) in the Chihuahuan and Sonoran subdivisions of the North American Desert. Jour. Arizona Acad. 4:183-184.
. 1967a. Ecotypic variation in Larrea divaricata. Am. Jour. Bot. 54:1041- 1044.
NOTES AND NEWS
NOTES OF THE FLORA OF ARIZONA. IV—Five collections worthy cf special rote have been added to the University of Arizona Herbarium. Three represent new species for the state, including one new genus, and two are range extensions.
Urtica urens L. represents a new weed in the Arizona flora. Our collections, made by Dr. K. C. Hamilton, came from the John Jacobs Citrus Grove on the northeast corner of the Black Canyon Highway and Peoria Avenue, in the Salt River Valley, Maricopa Co. The Dwarf or Slender Nettle is an annual from Europe which has become naturalized in the eastern United States and California.
Chaerophyllum tainturieri Hook. is a genus and species new for the state. Our specimen was collected by Stephen Bingham (871), eight miles north of the Safford airport, at an elevation of 4800 feet in the Gila Mountains of Graham Co. Mathias and Constance (North American Fora 28b:117-119. 1944) co net record any species of this genus west of Texas. In this work our material is determined as C. texanum Coult. & Rose; however, in their more recent publication on the Umbelliferae (Flora of Texas 3:288-290. 1951), they consider C. texanum a synonym of C. tainturieri.
Erigeron lonchophyllus Hook. was collected in deep grass near Aspen Spring, Lukachukai Mountains, Apache Co. (Niles & Mason 2486). This collection adds a new species to the flora of Arizona, but it is not an unexpected addition. Cronquist in his monograph of the genus (Brittonia 6:127-302. 1947) cites a specimen from the Tunitcha Mountains of San Juan Co., New Mexico, a mountain range which is adjacent to the Lukachukai-Chuska chain.
Paronychia sessiliflora Nutt. was first recorded for Arizona by McDougall (Pla- teau 37:107. 1965) from a specimen collected at Meteor Crater. An earlier collection by Eastwood and Howell (6918) from the same area had been misdetermined as P. Jamesii. More recently a third collection (Reese & Niles 850) from the East Side Game Road, Kaibab Plateau, Coconino Co. extends the range approximately 125 air miles northward and closer to the center of distribution of the species.
Hippuris vulgaris L., previously reported from lakes in the White Mountain area of Apache Co., has now been collected from lakes in the vicinity of Wide Lake, Luka- chukai Mountains, Apache Co. (Niles & Mason 2487). This new locality is about 175 aerial miles north of the previous collection. —CHARLES T. MAson, Jr., Her- barium, University of Arizona, Tucson, and WEsLrEy E. Nites, New York Botanical Garden.
CYTOLOGICAL AND EVOLUTIONARY STUDIES ON PALUSTRES VIOLETS
MarjJA SORSA
The genus Viola is well known for its many taxonomically difficult complexes. The classification is based on extensive studies carried out by different workers in Europe, North America, and Asia. However, more recent studies, specializing on smaller groups of the genus, have shown that re-evaluation of the classification is often necessary (Baker, 1949; 1953; 1957; 1960; Cain and Dansereau, 1952; Clausen, 1926; 1929; 1931; 1951; 1964; Russell, 1954; 1955; Russell and Cross- white, 1963; Schmidt, 1961; Valentine, 1941; 1950). Within the four sections of the genus there are smaller subgroups which are in many cases connected together by hybrids only partially intersterile (Clausen, 1951). Within such subgroups a gradual chromosomal and morpho- logical stabilization often takes place in a hybrid population, leading to intogressive establishment of several independent types, whose hybrid nature is often difficult to recognize without cytological or populational studies (Russell, 1954; Schofer, 1954; Sorsa, 1965).
The Palustres group belonging to the subsection Stolonosae Kupffer, section Plagiostigma Godr. likewise presents taxonomic problems. No intensive cytological studies have hitherto been carried out on this circumpolar group of species as a whole (Clausen, 1964; Hulten, 1958).
The author had an opportunity of spending a year 1963-64 at the University of California, Berkeley, as an ASLA-Fulbright grantee. Dur- ing that time herbarium material of the subsection Stolonosae was ex- amined from the Herbarium of University of California, Berkeley, the Gray Herbarium, and the U.S. National Herbarium. Material for cytological studies was collected by the author on field trips from Alaska, Washington, Wyoming and California (see later for further explanation). Small flower buds were fixed in acetic-alcohol (3:1) either in the field or from plants grown first in the University of Cali- fornia Botanical Garden, Berkeley, and later at the Department of Genetics and the Botanical Garden of the University of Helsinki. The preparations were mostly made to catch PMC meiosis. Chasmogamous buds were preferred, because the anthers of the cleistogamous buds con- tain very few PMC’s.
Excellent staining of squashes was obtained by the Feulgen method combined with post-staining with acetic-iron-hematoxylin adapted from Wittmann (1962).
The herbarium material of the species studied is deposited in the Botanical Museum, University of Helsinki, Finland.
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THE PALUSTRES GROUP IN EUROPE
The central species of the Palustres group is Viola epipsila Led., which is a diploid 2n = 24, x = 12 being considered to be the basic number of the section (Clausen, 1927; 1929; Love and Love, 1961; Sorsa, 1965). It hybridizes commonly in nature with its tetraploid counterpart Viola palustris L., which has the chromsome number 2n = 48 (Clausen, 1927; 1929; 1964; Gershoy, 1932; 1934; Jorgensen et al., 1958; Love and Love, 1956; 1961; Schmidt, 1960; Sorsa, 1965), wherever their distribution overlaps in Fennoscandia, Northern Russia, and East Prussia (Hiitonen, 1934). It has been shown that the hybrids frequently resemble the V. palustris parent in morphological respects and can only be recognized as hybrids by cytological investigation or thorough analysis of a large population sample (Sorsa, 1965). The hybrid populations obviously mostly represent later hybrid generations and backcrosses with V. palustris (Sorsa, 1965).
VIOLA PALUSTRIS ssp. JURESSII (K. Wein.) Fourn. grows as dis- junctive from V. palustris in France and Spain (Hulten, 1958), where- as both of the types are reported to occur in Britain (Clapham and Tutin, 1957). The taxonomic position of ssp. juressii has been uncertain: it has been reported as V. epipsila as well as a species in its own right, V. juressu Link. (Hulten, 1958). Its possible identity with the hybrid V. epipsila * palustris has also been suspected (Eklund, 1958; Hiitonen, 1934). Morphologically, it is intermediate between V. palustris and V. epipsila (fig. 1). Fernandes (1950) reports the chromosome number 2n = 48 from NE Portugal for V. palustris ssp. herminiu Wein. This type may in fact be identical with V. palustris ssp. juressit.
Cytological investigations have now been carried out on plants of V. palustris ssp. juressii grown from seed which was obtained in 1962 from Portugal, near Ponte de Lima, through the kind offices of P. Silva and J. Jalas. The chromosome number proves to be the same as in V. palustris, 2n = 48, and the meiosis is normal, showing 24 bivalents, all of about the same size (fig. 2). As far as the chromosomes are con- cerned, there thus seems to be no objection to granting this taxon sub- specific status as V. palustris ssp. juressit. Possibly the proper morpho- logical species characters need reconsidering, which might result in merging ssp. juressi with V. palustris.
The juressii type is probably derived from the plants which were cut off in the south by the last Quaternary glaciation. It seems plausible that the tetraploid palustris had already spread to the new and more oceanic areas in Western Europe before the glacial period, while the diploid epipsila was more restricted to the continental and northern cli- mate. The great glaciation thus left Viola palustris as disjunctive popu- lations in the Central and South European mountain areas by restricting gene exchange with its more northern populations. Such a situation
1968] SORSA: PALUSTRES VIOLETS 167
Fic. 1. Viola palustris ssp. juressii, an individual grown in greenhouse from seed obtained from Portugal; morphologically an intermediate between V. palustris and V. epipsila. It is tall (10-12 cm), having bluntly acute leaves, pubescent on the underside, and bracteoles above the middle of the pedicel. The flowers are of the same size and color as in V. palustris, only the spur being longer. In cultivation ssp. juresst is vigorous, spreading cleistogamously by seed ard vegetatively by stolons.
Fic. 2. PMC-meiosis in V. palustris ssp. juressii: a, Diakinesis showing 24 bi-
valents all about the same size; b, Anaphase I showing regular separation of paired chromosomes; c, Metaphase II showing n = 24 in both plates, & ca. 1200.
would explain the slightly different morphological evolution of the south- ern isolates through genetic drift.
THE PALUSTRES GROUP IN NORTH AMERICA
Viola palustris has been used as a kind of general expression for Palustres violets in North America by many American taxonomists. Apart from wrong annotations on the herbarium specimens, V. palustris
168 MADRONO [Vol. 19
Ss
Fic. 3. A plant of Viola epipsila ssp. repens from Moose Creek, Alaska; glab- rous, acute leaves, larger flowers and smaller plant size (8-10 cm) than in V. epipsila.
is often confused with V. epipsila ssp. repens, V. macloskew (n = 12), V. pallens (n = 12), V. renifola (n= 12), V. incognita (n = 22) on even V. blanda (n = 22), the North American V. palustris is by no means a uniform taxonomic entity.
Cytological investigations have now been carried out on four types belonging to the North American palustris complex as well as on one closely related member of the group.
VIOLA EPIPSILA Ledebour ssp. REPENS (Turczaninov) Becker, Bot. Centralbl. Beiheft 34(2): 406. 1917, has a wide distribution, from Eastern Asia to Alaska and western Canada. (Anderson, 1959; Hulten, 1947; 1958). It is reported from British Columbia and Washington (Hitchcock et al., 1961; Hamet-Ahti, 1965) but the specimens seen from that area differ slightly from the Alaskan type in morphological features. Moreover, the chromosomal condition seems to be quite dif- ferent (see later for V. palustris in Washington).
Morphologically, the subspecies repens differs from the main type V. epipsila by a smaller size, glabrous but distinctly acute leaves, and larger flowers (fig. 3).
The chromosome number of the western subspecies has not been re- ported from North America (Clausen, 1964). However, in 1930 Miyaji determined the chromosome number n = 12 of a plant listed as V. repens Turcz. from the moors of Hokkaido. Obviously, this is the Japanese form of V. epipsila ssp. repens. The same chromosome number was now determined on specimens of three populations collected in Central Alaska, at Kantishna, Camp Denali and Moose Creek. The meiotic division is regular (fig. 4). No differences could be detected in chromo-
1968 | SORSA: PALUSTRES VIOLETS 169
—_—
=e
Fic. 4. PMC-meiosis of V. epipsila ssp. repens from Moose Creek, Alaska: a, Anaphase I showing regular separation of bivalent halves; b, Metaphase II show- ing n = 12 in both metaphase plates, x ca. 1800.
some size or morphology between the Fennoscandian V. epipsila and the Alaskan ssp. repens (see Sorsa, 1965). The mitotic division accord- antly showed 2n = 24.
VIOLA ACHYROPHORA Greene, Pittonia 5: 33. 1902, has been reported from both continents in the Beringean area (Hulten, 1947; 1958), though not listed in the Flora of USSR (15:402-404. 1949). Hulten (1947) and Anderson (1959) were hesitant to rank this type as a sep- arate species, merely because of slight morphological differences and a more northerly distribution. In the author’s opinion, as well, the dif- ferences (smaller size, rounder and thicker leaves) between “V. achyro- phora” and V. epipsila ssp. repens are mainly due to modification in the poorer and more northern habitats. No differences between the two are observable in style shape, either.
170 MADRONO [Vol. 19
ey. 06 ee
oS
Fic. 5. PMC metaphase I of “V. achyrophora” from Chatanika River, Alaska, showing 12 normal bivalents all about the same size, x ca. 1800.
The chromosomal picture of the V. achyrophora type is regular, show- ing 12 bivalents in meiosis, and no differences from V. epzpsila ssp. repens can be noticed on the chromosome level (fig. 5). The same chro- mosome number has been found in both populations sampled north of Fairbanks, Alaska, namely Chatanika and Miller House.
It would seem more logical to regard “V. achyrophora” as just a northern variant of V. epipsila ssp. repens keeping it under the same specific name.
VIOLA PALUSTRIS Linnaeus, Sp. Pl. 934. 1753. As mentioned earlier, the name V. palustris has quite often been used as a general denomina- tion for the whole palustris complex in North America. The author’s acquaintance with the palustris type in the Eastern United States and Canada is restricted to herbarium material collected mostly around the turn of the century. Morphologically, this material is very near to the North European V. palustris, which is known also from Iceland, from where the chromosome number 2n = 48 has been reported by Love and Love (1956), and Southern Greenland, from where 2n = 48 has been confirmed by Jorgensen et al. (1958). In eastern North America the species occurs in New Foundland, Gaspé Peninsula, and Quebec on the Canadian side. From eastern U.S. it is reported around the White Mountain area only (Russell, 1965). However, no chromosome counts are available.
The palustris material seen from western North America, from Brit- ish Columbia, Washington and the adjacent areas is morphologically quite near to what in Europe is considered the epipsila type (Hulten, 1958) in size, shape of leaves, flower characters and ecological require- ments (fig. 6).
1968 | SORSA: PALUSTRES VIOLETS iA
Fic. 6. Plants of the collective species of Vzola palustris from Mt. Rainier Nat. Park, Washington. Morphologically this plant appears nearer the epipsila type: the flowers are large, pale blue, and have a long spur. Bracteoles are above the middle of the pedicel. Only two leaves are present at the time of flowering (V. palustris having three or more). Only the shape and glabrousness of the leaves resemble the palustris type.
Gershoy (1934) reported the chromosome number of 2n = 48 from the Pacific coastal form of V. palustris, collected in the vicinity of Ocean Park, British Columbia. This plant was cytologically regular, having 24 bivalents in metaphase I. According to the morphological descrip- tion of the plant given by Bold and Gershoy (1934) there are slight differences with the North European palustris e.g. in shape of leaves and color of seeds.
The herbarium specimens labelled as V. palustris from the Pacific coastal areas show a confusing variation of types not typical at all if compared with the North European V. palustris. Unfortunately no material was obtained of the Pacific coastal types in spite of several field trips to the sites of growth in California, given on the herbarium sheets (Humboldt Co., Mendocino Co.).
The writer has studied cytologically two types of the western palustris complex from Washington and Wyoming.
Plants belonging to the Viola palustris complex were fixed from an abundant population in Frog Heaven, Mt. Rainier National Park, Washington. Though the plants were morphologically more of the Euro- pean epipsila type (fig. 6), they chromosomally belonged to the tetra- ploid level. The meiotic metaphase I showed minor irregularities in the first division (fig. 7). The meiotic picture very much resembles that of the previously studied V. palustris * epipsila hybrids in Finland (Sorsa,
172 MADRONO [Vol. 19
Fic. 7. Two PMC’s in metaphase I of the V. palustris plant from Mt. Rainier Nat. Park, Washington. The orientation of the chromosomes is slightly irregular three small univalents (arrows) dividing not completely synchronous with the bivalent separation. 2n = 22-231 + 2-31, & ca. 1200.
1965). Both seem to be well stabilized hybrid derivatives with little hybrid unbalance, the somatic chromosome number maintaining around 2n = 48. The occurrence of the epzpsila type characters in the tetra- ploid level such as the position of bracteoles and the number of leaves suggests the emerging of the epzpsila genome. This can either be due to actual hybridization of the palustris type with the diploid V. epipsila repens taken place relatively recently. One can also think of the epipsila type as one of the progenitors of palustris, which would then lead into the emergence of various combinations of parental characters in different environments. It seems to me highly probable that some of the obscure western palustris types are of hybrid origin. The possible partners in the hybridization that obviously took place quite long ago will be discussed later.
VIOLA PALUSTRIS ssp. BREVIPES Baker, Madrono 6: 235. 1936, has been described from several sites along the Colorado River at altitudes of 7500-10000 ft (Baker, 1936; probably also Russell, 1963). This white-flowered violet has obviously quite frequently been confused with V. pallens and V. blanda. The plant was found in July 1964 near Lewis Lake, Yellowstone National Park, Wyoming, though stated to be V. blanda in the local flora (McDougall and Baggley, 1956). All char- acters, e.g. shape of style, beardless petals, size, shape of leaves and flowers (fig. 8) fit the description of Baker (1936) for ssp. brevipes.
1968 | SORSA: PALUSTRES VIOLETS 173
Fic. 8. A plant of Vzola palustris ssp. brevipes from Yellowstone Nat. Park, Wyoming. It has white flowers, a short spur, beardless petals, and has bracteoles above the middle of the pedicel. It is small, 5-8 cm, and the leaves are orbicular- cordate with regularly crenate margins.
The meiotic picture is suggestive of some hybridity in the popula- tion, revealed by somewhat irregular pairing and the presence of 2—5 univalents (fig. 9). Clausen (1964) also reports V. palustris ssp. brev- ipes, a type specimen from Colorado, Estes Park, as a hybrid deriva- tive having ‘36-38 pairs or singles.” If about 12 are singles, the chro- mosome number might have arisen as a result of hybridization of a tetraploid and a d’plo’d counterpart. Clausen’s plant from Baker’s type collection possibly represents a young hybrid generation, in which chromosomal stabilization has not yet taken place. The Yellowstone population, in turn, may represent an older, balanced hybrid deriva- tive, showing a rise of chromosome number and moderate stabiliza- tion, which is by no means uncommon among the violet hybrids (Brain- erd, 1921; 1924; Clausen, 1922; 1926; 1931; Schmidt, 1961; Sorsa, 1965).
Without thorough experimental evidence one can only hazard a guess at the most probable history of this hybrid derivative. On morpholog- ical grounds (white flowers, round leaves, small size) the likeliest pos- sibility would appear to be hybridization of the tetraploid western palustris type with the diploid V. macloskeyi, which occurs in similar habitats and areas. This continental and montane to subalpine form is possibly still evolving.
The chromosome number of VIOLA MACLOSKEYI Lloyd, Erythea 3: 74. 1895, was verified as n = 12 from specimens collected from four
174 MADRONO [ Vol. 19
Fic. 9. Diakinesis of PMC meiosis in V. palustris ssp. brevipes from Yellowstone Nat. Park, Wyoming; irregular pairing of the chromosomes, 24 bivalents (one loosely paired) and 3 fragments can be seen, X ca. 1800.
populations, three from Yosemite National Park, California and one from a transplant No. 62.1302 in the Botanical Garden at Berkeley originally collected from Lake Tahoe (fig. 10). The same chromosome number has been reported earlier (Baker, 1953; Clausen, 1964).
REGIONAL AREAS OF SPECIES
The phylogenetic outlines of the genus Viola have been discussed most thoroughly by Clausen (1929). Although deductions of this type are always somewhat speculative, a combination of cytological, genet- ical and geographical information can provide some clue to what hap- pened in the past. The Palustres group probably originated in north- eastern Asia, similarly to many circumpolar groups of species. The di- ploid level is represented as Viola epipsila, and the tetraploid as V. palustris. The ready hybridization of the two suggests that the origin of the tetraploid palustris was probably through hybridization of two diploid types, one of which was the epipsila type (fig. 11).
The tetraploid type has expanded its range into more oceanic envir- onments, while the diploid epipsila type covers the more continental areas of Northern Europe, and as a slightly differentiated subspecies
1968] SORSA: PALUSTRES VIOLETS u/s)
SRS
Fic. 10. Diakinesis of PMC meiosis in Viola macloskeyi grown in Botanical Garden, Berkeley (No. 62.1302); n= 12, the bivalents appear to be mostly ring- shaped, which was found to be frequent in all the macloskeyi samples studied, x ca. 1200.
repens extends to Siberia and the Beringean area as far as Alaska and possibly western Canada. The tetraploid palustris type is widespread in Europe and is connected to Eastern North America over Faeroe Islands, Iceland and Greenland. In North America it tends to have a bi-coastal distribution, although the author is inclined to believe that the original “Linnean” species V. palustris is confined to the eastern coast, i.e. Newfoundland and Gaspé Peninsula in northeastern Canada, and the White Mountains area in the United States, while the western types of palustris are less typical and often show variation in character combinations.
It seems probable that much of the variation of the palustris type is caused by hybridization with ecologically and karyologically suit- able related violets, such as species of the diploid group (n= 12) V. macloskeyi, V. pallens and possibly also V. lanceolata, V. primulifolia and V. renifolia. Evidence from the extensive crossing experiments of Gershoy and his co-workers (Bold and Gershoy, 1934; Gershoy, 1934) drops out the more distant relatives of the Palustres group such as V. blanda and V. incognita (both n = 22) and V. selkirkit (n = 12). The possibility of hybridization between the eastern palustris with the northwestern epipsila repens cannot be excluded, either.
The cluster of species forming the Palustres group thus seems to con- sist of one tetraploid, V. palustris, and at least three diploids, namely V. epipsila and the species pair V. macloskeyi-V. pallens. Experimental evidence is needed to confirm their evolutionary possibilities in creating the variety of forms that constitute the taxonomically difficult Palustres group.
176 MADRONO [Vol. 19
V SHIKOKIANA eine ” WW ae
¢ DIPLOID
7 aay Ele ~ V PALUSTRIS if (| vas MOUPINEN SIS
P REPENS
VV MACLOSKEY! ? ae
V. PALLENS —S
—
pIPSiLA SS V PRIMULIFOLIA ?|
V RENIFOLIA ?
¢ Sa V palustris 7 = 4
SSp aioe 2 Q 2 : \ -9 iy
Fic. 11. Schematic representation of the speculative evolutionary outlines in the Palustres group. The full lines represent diploid species, the broken lines tetraploid species. Heavy arrows in North America suggest the possible contributors to the variability of the palustris complex. V. shikokiana (n = 12, Miyaji, 1929) is the East Asiatic counterpart of the macloskeyi-pallens species pair, V. palustris var. moupinensis is probably a separate species reported from Southern China. See further in the text.
ACKNOWLEDGMENTS
I wish to express my sincere gratitude to the great authority on the genus Viola, Jens Clausen, Carnegie Institution of Washington, Stan- ford, for reading the manuscript and making numerous valuable com- ments both to amend the factual information and to improve my English text. I also want to thank Esko Suomalainen, University of Helsinki, Finland, for advice and encouragement during the studies, and Jaakko Jalas, University of Helsinki for help and criticism during the work. Herbert G. Baker, Margaret Bergseng and Annetta Carter, University of California, Berkeley, were always kindly helping me in many ways during my stay at Berkeley.
The field trip to Alaska in June 1964 was sponsored by U.S. Govern- ment Grant No. 33511-215009 given to me by the International Insti- tute of Education.
1968 | SORSA: PALUSTRES VIOLETS 177
SUMMARY
Chromosome numbers of violets belonging to the Palustres group, subsection Stolonosae Kupff. are reported:
Viola palustris ssp. juressii (K. Wein.) Fourn. n = 24, from culti- vated plants grown from seed obtained from Portugal, near Ponte de Lima. Meiosis normal.
Viola epipsila ssp. repens (Turcz.) Becker, n = 12, from three dif- ferent localities in Central Alaska (Kantishna, Camp Denali, Moose Creek). Meiosis normal.
“Viola achyrophora” Greene, n = 12, from two localities in Northern Alaska (Chatanika, Miller House). Meiosis normal. It is suggested that this type should be treated rather as belonging to V. epipsila ssp. repens.
Viola palustris (coll.), n = 22—23n + 2-31, slightly irregular meiosis, from Frog Heaven, Mt. Rainier National Park, Washington. The type obviously represents a stabilized hybrid derivative probably quite fre- quent among the North American palustris complex.
Viola palustris ssp. brevipes Baker, n= 22—24n + 3-51, slightly irregular meiosis, from Lewis Lake, Yellowstone National Park, Wyo- ming, represents a type of stabilized hybrid derivative. The most prob- able progenitors are the western palustris type and the white-flowered diploid V. macloskey.
Viola macloskeyi Lloyd (=V. macloskew ssp. macloskeyw Baker), n= 12, normal meiosis, from a Lake Tahoe transplant and three sites in Yosemite National Park, Calif.
The emergence of some key morphological characters of V. epipsila on the tetraploid level in V. palustris ssp. juressii and especially in the western North American V. palustris has been observed and discussed.
The distribution of the diploid and the tetraploid types of the Palus- tres group has been presented. The group seems to consist of an evo- lutionary active complex, able to balance itself in different environments by forming various types due to selection and genetic drift.
Department of Genetics, University of Helsinki, Finland
LITERATURE CITED
ANDERSON, J. P. 1959. Flora of Alaska and adjacent parts of Canada. Iowa State Univ. Press.
BAKER, M. S. 1936. Studies in western violets I. Madrofio 6: 232-239.
. 1949. Studies in western violets VI. Madrono 3:110-128. . 1953. Studies in western violets VII. Madronfio 12:8-18. . 1957. Studies in western violets VIII. Brittonia 9:217—230.
Botp, H. C. and A. GrersHoy. 1934. Studies in North American violets IV. Chromo- some relations and fertility in diploid and tetraploid species hybrids. Vt. Agr. Exp. Sta. Bull. 378.
BRAINERD, E. 1921. Violets of North America. Vt. Agr. Exp. Sta. Bull. 224.
. 1924. Some natural violet hybrids of North America. Vt. Agr. Exp. Sta. Bull. 239.
Cain, S. A. and P. Danserau. 1952. A study of the stemmed yellow violets of
eastern North America (four species) on the basis of mass collections. Philos.
178 MADRONO [Vol. 19
Soc. Yearb. 1951: 166. CriapHAM, A. R., T. G. TuTIn and E. F. Warsurc. 1952. Flora of the British Isles. Cambridge Univ. Press. CLAUSEN, J. 1922. Studies on the collective species Viola tricolor L. II. Bot. Tidskr. 37:363-416. . 1926. Genetical and cytological investigations on Viola tricolor L. and V. arvensis Murr. Hereditas 8:1-156. . 1927. Chromosome number and relationship of species in the genus Viola. Ann. Bot. 41:677-714. . 1929. Chromosome number and relationship of some North American species of Viola. Ann. Bot. 43:741-764. . 1931. Cyto-genetic and taxonomic investigations in Melanium violets. Hereditas 15:279-304. . 1951. Stages in the evolution of plant species. Cornell Univ. Press, Ithaca. . 1964. Cytotaxonomy and distributional ecology of western North Amer- ican violets. Madrono 17:173-197. ExKLiunpD, O. 1958. Die Gefasspflanzenflora beiderseits Skiftet im Scharen-Archipel Sudwestfinnlands. Finska Vetensk. Soc. Handl. 101:1-324. FERNANDES, A. 1950. Sobre a cariologia de algumas plantas da Serra do Gerés. Agron. Lusit. 12:551-600. GersHoy, A. 1932. Descriptive notes for Viola exhibit. Sixth Int. Congr. Genet. ithaca; Noy. . 1934. Studies in North American violets III. Chromosome numbers and species characters. Vt. Agr. Exp. Sta. Bull. 367. Huronen, I. 1934. Uber einige sog. intermediare Arten und deren Ursprung. Mem. Soc. Fauna FI. Fenn. 10:59-69. Hitcucock, C. L., A. Cronquist, M. Ownsey, and J. W. THompson. 1964. Vas. cular plants of the Pacific Northwest. Vol. 2. Univ. Wash. Publ. Biol. 17:1-597. Hutrten, E. 1947. Flora of Alaska and Yukon, VII. Lunds Univ. Arsskr. II. Sect. 2. 43. . 1958. The amphi-atlantic plants. Sv. Vet-akad. Handl. III. 7:1-340. HAmet-AntmI, L. 1965. Vascular plants of Wells Gray Provincial Park and its vicin- ity, in eastern British Columbia. Ann. Bot. Fenn. 2:138-164. JORGENSEN, C. A., TH. SORENSEN, and M. WESTERGAARD. 1958. The flowering plants of Greenland. Biol. Skr. Dan. Vid. Selsk. 9(4) :1-172. Komarov, V. L. (ed). 1949. Flora URSS. Acad. Scient. URSS, Mosqua and Lenin- grad. 15:402—-404. Love, A. and D. Love. 1956. Cytotaxonomical conspectus of the Icelandic flora. Acta Horti Gotob. 20:65-291. . 1961. Chromosome numbers of central and northwest European plant species. Op. Bot. 5:1-581. Mc Doveatt, W. B. and H. A. BAccLey. 1956. The plants of Yellowstone National Park, U.S. Gvt. Printing Office, Washington, D.C. Mrvyaji, Y. 1929. Studien tiber die Zahlenverhaltnisse der Chromosomen bei der Gattung Viola. Cytologia 1:28-58. . 1930. Betrachtungen tiber die Chromosomenzahlen von Viola, Violaceen und verwandten Familien. Planta 11:631-649. RussEL1, N. H. 1954. Three field studies of hybridization in stemless white violets. Am. Jour. Bot. 41:679-686. . 1955. Local introgression between Viola cucullata Alt. and V. septentrion- alis Greene. Evolution 9:436—440. . 1963. Viola palustris L. in Arizona. Rhodora 65:49. . 1965. Violets (Viola) of Central and Eastern United States: an intro- ductory survey. Sida 2:1-113. . and F. S. Crosswuirte. 1963. An analysis of variation in Viola nephro- phylla. Madronio 2:56-65.
1968 | DRESSLER & KUIJT: AMMOBROMA 179
ScumipT, A. 1960. Zytotaxonomische Studien an europaischen Veilchen. Diss. Univ. Miinchen. . 1961. Zytotaxonomische Untersuchungen an europaischen Viola-Arten der Sektion Nominum. Oesterr. Bot. Zeitschr. 108: 20-88. ScHorer, G. 1954. Untersuchungen iiber die Polymorphie einheimischer Veilchen. Planta 43:537-565. Sorsa, M. 1965. Hybridization of Palustres violets in Finland. Ann. Acad. Sci. Fenn. A, IV, 86:1-18. VALENTINE, D. C. 1941. Variation in Viola riviniana Rchb. New Phytol. 40:189-209. . 1950. The experimental taxonomy of two species of Viola. New Phytol. 49:193-212. Wittmann, D. 1962. Aceto-iron-haematoxylin for staining chromosomes in squashes of plant material. Stain Tech. 37:27-30.
A SECOND SPECIES OF AMMOBROMA (LENNOACEAE), IN SINALOA, MEXICO
Rosert L. DRESSLER AND JoB KUIJT
The genus Ammobroma has stood as a monotypic taxon since its ini- tial description more than a hundred years ago (Gray, 1854). Its single species, A. sonorae, has through the years attracted much casual atten- tion because of the curious, mushroom-like appearance of the plant, and because of its early use as an edible plant by some of the indigenous Indian tribes of the Sonoran region. It is of some interest to note that Dr. Jerry Rzedowski has observed the use of Lennoa also as a veg- etable in the State of San Luis Potosi (personal communication). Not- withstanding the obvious interest in these bizarre parasites very little botanical work has been done in the family.
It is not our intention here to enter into a discussion of the generic distinctions between the three known genera of Lennoaceae (Ammo- broma, Pholisma, and Lennoa). Even a casual acquaintance with Pho- lisma and Lennoa leaves one with an uneasy feeling about their valid- ity as genera. It could even be argued that the main generic feature of Ammobroma, i.e., the compact, laterally extended head, is insufficient by itself as a generic distinction.
Whatever generic delimitations may be used in the future, it is of great interest that we can now add a new species to this small family. At present the species must undoubtedly be placed in Ammobroma. The plant was first observed by the senior author in December, 1949, in the region of Culiacan, Sinaloa. Only dried and decomposed remains were found at that time. It could be ascertained, nevertheless, that the plants were attached to the roots of Euphorbia (Agaloma) californica. More recently (August, 1957), a second visit was made to the same locality, and many flowering plants were found on the roots of Euphorbia cali- fornica, Jatropha curcas, and J. cordata. As now known, the parasite is
180 MADRONO [Vol. 19
Ammobroma culiacana
Fic. 1. Ammobroma culiacana: a, complete plant on root of host plant, showing you shoot and the roots of the parasite; b, calyx, split open; c. corolla, split open to show pistil and anthers; d, a clump of plants as seen in nature (from a koda- chrome).
quite local, having been found only on a rocky ridge about 15 km south of Culiacan, from which we derive the specific epithet.
Ammobroma culiacana Dressler & Kuijt, n. sp. Caules carnosi, 8-10 cm longi, 1.5—2.0 cm crassi, orientes e radice nutricia ipsa vel paene; folia squamiformia, parva, basalia deltoideo-ovata glabraque, media longiora, superiora circa basim inflorescentiae involucrum effor- mantia. Inflorescentia capitulum compactum, 3-10 cm diam., hemi-
1968 | DRESSLER & KUIJT: AMMOBROMA 181
sphaericum, suborbiculare sed demum circumscriptione irregulari. Flores sessiles vel breviter pedicellati; calyx tubulosus, extra dense pilosus, praecipue superne, 5—5.5 mm longus, incisus ad 1.5—2.5 mm, lobis 6-8, obtusis, irregulariter dentatis, 1-2 mm latis. Tubus corollae 6—6.5 mm longus, glaber; lobi corollae 6-8, patentes, bifidi, plicati, 2.5-3 mm longi, 1.5—2 mm lati, intra glabri, extra sparse p'losi; stamina isomera, lobis corollae alternantia. Filamenta 0.3 mm longa, autherae ca. 0.7 mm longae; ovarium turbinatum, ca. 1.8 mm latum. Stylus ca. 4 mm longus, stigma integrum, capitatum. Fructus semiglobosus, 3-3.5 mm diam.; nuculae 14-18, reniformi-ovoideae, applanatae, ca. 1 mm longae; testa cellulis magnis, papillosis.
Stems fleshy, 8-10 cm long, 1.5—2.0 cm in diameter, arising directly from the infected host root or nearly so; leaf scales small, the basal ones deltoid-ovate and glabrous, the middle ones longer, the upper ones form- ing an envelope of densely pilose, linguiform to ovate, imbricating scales around the base of the florescence. Inflorescence a compound head 3—10 cm in diameter, hemispheric, more or less circular, but becoming irreg- ular in outline. Flowers sessile or shortly pedicellate, calyx tubular, densely pilose without, especially on upper parts, 5—5.5 mm long, divided for 1.5—2.5 mm into 6-8 obtuse, irregularly dentate lobes 1-2 mm in width; corolla tube 6—6.5 mm long, glabrous; corolla lobes 6—8, spread- ing, bifid and plicate, 2.5-3 mm long, 1.5—2 mm wide, glaborus within, somewhat pilose without, stamens as many as corolla lobes and alter- nating with them. Filaments 0.3 mm long, anthers ca. 0.7 mm long, ovary turbinate, ca. 1.8 mm wide; style ca. 4 mm long, stigma entire, capitate. Fruit subhemispheric, 3-3.5 mm in diameter, nutlets 14-18, reniform-ovoid, flattened, about 1 mm long, seed coat consisting of large, papillose cells.
Specimens examined. Mexico. Sinaloa: about 15 km south of Culiacan (km 1410), rocky soil in thorn forest, perasitic on roo's of Euphorbia californica, Jatro- pha curcas and J. cordata, indument and exposed bracts pink, flowers cream with pink shading, Dressler 2175 (F, GH, K, MEXU, MO, NY, UBC, UC-holotype, US, VEN), Aug. 11, 1957; 10 miles south of Culiacan, rocky thorn forest, parasitic on roots of Euphorbia, Dressler 984 (MEXU, MO, UC, US), Dec. 22, 1949.
Ammobroma culiacana differs from all other known Lennoaceae in its gamosepalous clyx. The remaining members of the family, including A. sonorae, have very narrow sepals which are free to the base or nearly so. The matted indument of the sepals gives the heads a flesh-pink color in life. The corollas are cream or tinged with pink and, unlike those of other Lennoaceae, do not contrast markedly with the pale indument of the head. A faint, sweet odor was detected, and the flowers were visited by flies, beetles, and small butterflies. As the inflorescence matures and expands laterally the (earliest) flowers in the central portion become dark brown. The entire perianth persists even when the fruit is mature. The latter, in agreement with other Lennoaceae, is a capsule-like fruit which appears to open by means of an irregular, circumscissile rupture.
182 MADRONO [Vol. 19
It seems that mechanical destruction of old inflorescences is necessary before the seeds are set free.
The inflorescence of Ammobroma culiacana may be said to be some- what intermediate between those of A. sonorae and the Pholisma-Lennoa complex. In the latter two genera we find a panicle-like inflorescence in which the balance between lateral and terminal expansion has been variously struck. Lennoa and Pholisma depressum (P. paniculatum Templeton) show a great deal of lateral branching (Suessenguth, 1927; Templeton, 1962), but in P. arenarium the inflorescence approaches a short, dense spike. In Ammobroma culiacana a further contraction has taken place, and we can speak of a determinate, disk-like inflorescence axis and an involucre of bracts. The extreme is reached in A. sonorae with a very dense capitulum the margin of which unrolls well above the upper leaves. Reduction of floral bracts seems to run parallel to this series: in Pholisma and Lennoa each flower has a subtending bract but such organs are no longer recognizable in Ammobroma, at least not in the central area of the disk.
Although details of root morphology cannot be ascertained at this time, one interesting feature is clearly shown in the collections cited, and is illustrated in Fig. 1A. The stem originates directly from the host root, or nearly so. On the basis of what is known for Pholisma (Kuijt, 1968) and the collections cited, we can tentatively reconstruct the fol- lowing course of events in Ammobroma culiacana. Long roots traverse the soil. When the host rhizosphere is invaded, small haustorial roots are formed nearest the host root and grow towards the latter, finally invading it. A dark, irregular, tubercular mass of variable size develops in the haustorial region. It is from this callus-like cushion that the stem is developed. This is in direct contrast to the situation in Pholisma, where stems arse from the roots of the parasite between successive haustorial attachments, but not at the point of host contact. The para- sitism of A. culiacana would seem to represent an evolutionary advance over that of Pholisma in that it involves a more immediate dependence of the inflorescence on the roots of the host.
We wish to express our indebtedness to Dr. Karel U. Kramer for the Latin diagnosis.
Smithsonian Tropical Research Institute, Balboa, C.Z. University of British Columbia, Vancouver
LITERATURE CITED
Kuyt, J. 1968. The biology of parasitic flowering plants. In preparation.
Gray, A. 1854. Plantae novae Thurberianae. Mem. Am. Acad. 5:297-328.
SUESSENGUTH, K. 1927. Uber die Gattung Lennoa. Ein Beitrag zur Kenntnis exoti- scher Parasiten. Flora 122:264-305.
TEMPLETON, B. C. 1962. A morphological comparison of Pholisma arenarium Nutt- all and Pholisma paniculatum Templeton (Lennoaceae). Los Angeles Mus. Contr. Sci. 57:3-29.
CHROMOSOME NUMBERS IN THE TRIBE PAPPOPHOREAE (GRAMINEAE)
JoHN R. REEDER AND D. N. SINGH
In a recent paper, Reeder (1965) pointed out that on the basis of anatomical characters of the leaf and embryo, the tribe Pappo- phoreae can be divided into two rather distinctive subgroups. One of these, which includes only the genus Pappophorum, was designated sub- tribe Pappophorinae. The other, comprising the genera Cottea, Ennea- pogon, Kaokochloa, and Schmidtia, was named subtribe Cottinae. In the paper referred to above, the cytology of members of the tribe Pap- pophoreae was not discussed.
Cytological information on members of this tribe published to date suggests that basic chromosome numbers of both x = 9 and x = 10 occur. Apparently the only species of Pappophorum which have been examined cytologically are P. bicolor Fourn. and P. mucronulatum Nees. For both of these, Brown (1950) and Gould (1958; 1966) have in- dicated that the chromosomes are in multiples of ten. The lowest num- ber reported in either species is 2n = 40.
The only published record for the monotypic genus Cottea is that of Covas (1945). He examined Argentinian material of C. pappophoroides Kunth, and reported 2n = 20.
One Indian, four African, and one American species of Enneapogon have been subjected to cytological study. Covas (1945) reported 2n = 20 in Argentinian material of E. desvauxit Beauv. (as Pappo- phorum wright S. Wats.). This number has been confirmed by Gould (1960, 1966) and Reeder (1967) in collections from Mexico and the United States. A diploid number of 2n = 20 was also determined for E. elegans (Nees) Stapf by Janaki-Ammal (in Darlington & Wylie, 1956). For E. scoparius Stapf, de Wet (1954) listed 2n = 36. Although basic numbers of x = 9 and x = 10 are common in grasses related to Enneapogon, curiously de Wet concluded that in this genus the basic number is x — 12. This would imply, of course, that his plant was triploid. In a later paper, de Wet & Anderson (1956) record the same number (2n — 36) for E. scoparius, and also for E. cenchroides (Licht.) C. E. Hubbard, and E. brachystachys (Jaub. & Spach) Stapf. For E. pretoriensis Stent, they report a chromsome number of 2n = 18. On the basis of this latter count, the authors suggest that the basic number in Enneapogon is probably x = 9. Thomas (in Darlington & Wylie, 1956) also gives 2n = 36 as the chromsome number for E. cen- chroides.
The two published chromsome counts for the genus Schmidtia are by de Wet & Anderson (1956) and de Wet (1958). These authors report 2n = 36 in S. glabra Pilger, and also in S. bulbosa Stapf. Laurent (1965), in a careful revision of the genus Schmidtia, treats both of the above taxa as synonyms of S. pappophoroides Steud.
183
184 MADRONO [Vol. 19
TABLE I. CHROMOSOME NUMBERS IN PAPPOPHOREAE*
Root-tip PMC Authority PAPPOPHORINAE Pappophorum bicolor 40, 60 Brown (1950) te 100 Gould (1958, 1966) mucronulatum 60 Brown (1950) Ss 60 Gould (1966) COTTINAE Cottea pappophoroides 20 Covas (1945) 20 Enneapogon brachystachys 36 de Wet & Anderson (1956) cenchroides 36 de Wet & Anderson (1956) es 36 Thomas (in Darlington & Wylie, 1956) - 40 40 desvauxit 20 Covas (1945) i. 20 Gould (1960, 1966) - 20 Reeder (1967) elegans 20t Janaki-Ammal (in Darling- ton & Wylie, 1956) pretoriensrs 18 de Wet & Anderson (1956) =< 18 de Wet (1958) scoparius 36 de Wet (1954, 1958) “ 36 de Wet & Anderson (1956) Kaokochloa nigrirostris 22 22 Schmidtia pappophoroides 36 de Wet & Anderson (1956) ze 36 de Wet (1958)
on 40 40
* All listed as 2n. Those in boldface are new counts by the authors. + No information as to part of plant examined.
No cytological studies have been reported thus far for the mono- typic genus Kaokochloa. This distinctive species is apparently confined to the Kaokoveld district of South West Africa, and was described only in 1961.
In view of the fact that within the subtribe Cottinae there appeared to be two basic chromsome numbers, sometimes both within the same genus, additional cytological studies seemed desirable. Through the courtesy of B. de Winter of Pretoria, South Africa, we have recently received caryopses of Enneapogon cenchroides, Schmidtia pappopho- roides, and Kaokochloa nigrirostris de Winter. Plants of all three species have been grown successfully in our greenhouse and experimental gar-
1968] REEDER & SINGH: PAPPOPHOREAE 185
CS & c “S oe ry ae o @¢@ o° 6 @ ~ Lv & 2 od = >} eo 6 @ \ \ ae ote e se Os C4 ® Pa | = 5 ~ me, = SCG a z -* r :~ be a4! p' Ft ve me Tee Nay) 4 ‘ ‘ NY e + 4 vy ? ~ 0 af Y ‘= od ° N 6 2 4
Fics. 1-6. Chromosome configurations; 1, 3, 5, 6, meiotic chromosomes from Division I of microsporocytes; 2, 4, somatic chromosomes from mitosis in root-tips; 1, Enneapogon cenchroides, diakinesis (2n = 40); 2, E. cenchroides, metaphase (2n = 40); 3, Schmidtia pappophoroides, anaphase (2n = 40); S. pappophoroides, metaphase (2n = 40); 5. Kaokochloa nigrirostris, diakenesis (2n = 22); 6. Cottea pappophoroides, anaphase (2n = 20), all X 900.
den. Chromsomes were studied in squashes of both pollen mother cells and root-tips.
Our investigations indicate that in Enneapogon cenchroides the chro- mosome number is 2n — 40 (figs. 1 & 2), and not 2n = 36, as re- ported by previous workers. Some irregularity was noted in meisosis, and univalents and multivalents were sometimes seen at diakinesis. The unexpected number of 2n = 22 was determined for Kaokochloa. Despite this, meiosis appears to be quite normal and eleven bivalents form regularly at diakinesis (fig. 5). Schmidtia proved to be tetraploid with 2n = 40 (figs. 3 & 4). This count is at variance with previous reports of 2n = 36 for this genus. As indicated above, although de Wet & Anderson (1956) and de Wet (1958) used the names S. bulbosa and S. glabra, Laurent (1965), in his monograph, considers both of these to be synonyms of S. pappophoroides.
In addition to the cytological studies reported above, we have also examined meiosis in microsporocytes of Cottea pappophoroides from Mexico. Our count of 2n = 20 (fig. 6) agrees with that given by Covas (1945), whose material came from Argentina. No irregularities were observed in meiosis.
186 MADRONO [Vol. 19
Our cytological studies of representatives of all four genera of the sub- tribe Cottinae, therefore, indicate that in this group the basic chromo- some number is x = 10. In one species each of Cottea and Enneapogon we found 2n — 20. Tetrapoid numbers of 2n = 40 were determined in Schmidtia, and a second species of Enneapogon. In Kaokochloa, it is true, we found 2n = 22, but it seems reasonable to interpret this as a case of aneuploidy, and to consider that the basic chromsome number in this genus is x = 10 also. In view of the close morphological and anatomical similarities between this genus and Schmidtia, a basic num- ber of x = 11 in Kaokochloa appears unlikely. A cytological reexami- nation of those species of Enneapogon in which counts of 2n = 18 and 2n — 36 have been reported seems desirable. The information on chro-
mosome numbers in members of the Pappophoreae is summarized in Table I.
List OF SPECIES STUDIED AND SOURCES OF MATERIAL
Cottea pappophoroides Kunth. Mexico: Chihuahua, 23 miles NW of Zavalza, J.& C. Reeder 4587, 3 October 1966, YU.
Enneapogon cenchroides (Licht.) C. E. Hubbard. South Africa: Pretoria, Pyramid. Seeds from above locality supplied by B. de Winter, and plants grown in ex- perimental garden at Yale University, J. & C. Reeder 4551, 16 July 1966, YU. Determination verified at Kew.
Kaokochloa nigrirostris de Winter. South West Africa: Kaokoveld. Seeds from above area supplied by B. de Winter, and plants grown in experimental garden at Yale University, J. & C. Reeder 4244, 5 July 1965, YU.
Schmidtia pappophoroides Steud. South West Africa: Gobabis. Seeds from above area [collected by H. Tolken] supplied by B. de Winter, and plants grown in greenhouse at Yale University, J. & C. Reeder 4821, April 1967, YU.
Yale University, New Haven, Connecticut
LITERATURE CITED
Brown, W. V. 1950. A cytological study of some Texas Gramineae. Bull. Torrey Club 77:63-76. Covas, G. 1945. Numero de cromosomas de algunas gramineas Argentinas. Revista Argent. Agron. 12:315-317. DarLINcTON, C. D., and A. P. Wytte. 1956. Chromosome Atlas of Flowering Plants. Macmillan Co., New York. pe Wet, J. M. J. 1954. Chromosome numbers of a few South African grasses. Cyto- logia 19:97-103. . 1958. Additional chromosome numbers in Transvaal grasses. Cytologia Zo 11S=118. . and L. J. ANpEerSOoN. 1956. Chromosome numbers in Transvaal grasses. Cytologia 21:1-10. GouLp, F. W. 1958. Chromosome numbers in Southwestern grasses. Am. Jour. Bot. 45:757-767.
1968] CORRELL: FLORA OF TEXAS 187
. 1960. Chromsome numbers in Southwestern grasses IJ. Am. Jour. Bot. 47:873-877. . 1966. Chromsome numbers of some Mexican grasses. Canad. Jour. Bot. 44:1683-1696. LaunerT, E. 1965. The genus Schmidtia Steudel (Gramineae). Bol. Soc. Brot. 39 (Ser. 2a) :303-322. REEDER, J. R. 1965. The tribe Orcuttieae and the subtribes of the Pappophoreae (Gramineae). Madrono 18:18-28. . 1967. Notes on Mexican grasses VI. Miscellaneous chromosome numbers. Bull. Torrey Club 94:1-17.
SOME ADDITIONS TO THE FLORA OF TEXAS—IV
DONOVAN S,. CORRELL
As work progresses on a Manual of the vascular plants of Texas, which Marshall C. Johnston and I are engaged in preparing, additions to the flora of the state are being continually found as evidenced by those reported here and which have been reported elsewhere. (Some Additions and Corrections to the Flora of Texas. Wrightia 3:126—140. 1965; Some Additions and Corrections to the Flora of Texas—II. Brit- tonia 18:306-310. 1966; and —III. Rhodora 68:420—428. 1966). This project is being supported, in part, by a grant from National Science Foundation (GB-3138). All of the specimens cited, unless otherwise noted, are in the Lundell Herbarium (LL) of Texas Research Founda- tion. I wish to acknowledge the help of my wife, Helen B. Correll, in the preparation of this paper.
SETCREASEA LEIANDRA (Torr.) Pilg. var. glandulosa Correll, var. nov. Planta var. leiandrae similis; pedicelli cum pilis brevibus glanduliferis vice villorum nitidorum.
Those plants that are found about Capote Falls and along Capote Creek in Presidio Co. and have short glandular hairs on their pedicels instead of the characteristic long silky hairs are referred to this variety.
Presidio Co., on ledges of cliffs above Capote Falls, in clumps, peren- nial, Nov. 3, 1966, D. S. Correll 34128 (holotype, LL); Capote Creek, Sept. (Oct.) 1883, V. Havard 79 (GH, US).
Nolina arenicola Correll, sp. nov. Caulis florifer ad 1 m altus (inflor- escentia includenti); folia numerosa caespitem magnum formantia, circa 1.3 m longa, 5 mm lata, complanata vel cancavo-convexa, margin- ibus rasilibus; panicula composita, aliquantum aperta, ramis grossis effusis vel patulo-ascendentibus; rami cum ramulis curtis; bracteae non conspicuae, plerumque ramos primarios fere aequantes vel leviter super- antes; perianthium 2.5-3.5 mm longum, segmentis ovato-ellipticis; fructus 4-7 mm lati, cum incisura profunda ad apicem; stylus prom- inens; pedicelli fructiferi 5-7 mm longi, prope basim articulati in fruc-
188 MADRONO [Vol. 19
tus dilatati, cum cristis prominentibus sub fructibus; semina 4-6 mm diametro.
Flowering stem up to 1 m high (including the inflorescence) ; leaves numerous to form a large clump, about 1.3 m long and 5 mm wide, flat or concave-convex, with smooth margins; panicle compound, rather open with coarse spreading or spreading-ascending branches, the branches with short stubby branchlets; bracts not conspicuous, mostly about equal to or only slightly exceeding the primary branches; perianth 2.5-3.5 mm long, the segments ovate-elliptic; fruits 4-7 mm wide, deeply notched at apex, the style prominent; fruiting pedicels 5-7 mm long, articulate near base, dilated into the fruit, with prominent ridges just under the fruits; seeds 4-6 mm. in diameter.
Culberson Co., in dunes, 18 miles east of Van Horn, on route USS. #80, July 24, 1957, D. S. Correll & I. M. Johnston 18445 (holotype, LL); sandy plain, scrub land, 17 miles east of Van Horn, scape 3 ft. tall, plants forming a large bunch, July 30, 1958, D. S. Correll & I. M. Johnston 20360.
This species is most closely allied to N. texana S. Wats. It is, how- ever, a much coarser plant than that species. Its coarsely branched open panicle resembles that of V. erumpens (Torr.) S. Wats. but the long pedicels and smooth-margined leaves readily separate it from that species. This species seems to occur only in sandy instead of the usually rocky or clayey soils of most of our species.
NOLINA MICRANTHA I, M. Johnst., Jour. Arn. Arb. 24: 91. 1943. Culberson Co., on rocky slopes near Pine Springs, Guadalupe Mts., inflorescence shorter than the leaves, July 25, 1957, D. S. Correll & I. M. Johnston 18473; same locality and date, male plant, D. S. Correll & I, M. Johnston 18518; on limestone hill, off highway #62, near Frijole, Guadalupe foothills, acaulescent, inflorescence pinkish, Aug. 8, 1945, C. L. Lundell & Amelia A. Lundell 14372. Hudspeth Co., on Keeling’s Ranch, Sierra Diablo, 12 miles north of Allamore, in clumps, grasslands, June 9, 1964, D. S. Correll, H. S. Gentry, & Craig Hanson 29769, 29774.
This northern Mexican species is new to Texas and the United States. The slenderly dissected and somewhat twiggy inflorescence, slender short pedicels, and the typically purplish or reddish purple tinged main rachis, branches and branchlets, and bracts are distinguishing character- istics of this species. It is quite different from any other species that we have in Texas.
YUCCA CARNEROSANA (Trel.) McKelvey, Yuccas Southw. U. S. 1:24, pls. 6 and 7. 1938.
Culberson Co., McKittrick Canyon, Guadalupe Mts., a few plants scattered on sides of canyon, distinguished by their trunks covered with reflexed leaves, no flowers seen, July 25, 1957, D. S. Correll & I. M. Johnston 18497; south fork of McKittrick Canyon, Guadalupe Mts.,
1968 | CORRELL: FLORA OF TEXAS 189
tall multi-trunked plant with showy heads of white flowers, up to 15 ft. tall, Apr. 20, 1962, D. S. Correll & E. C. Ogden 25057.
In 1938, McKelvey gave the area of distribution of this species in the United States as, “Confined to Brewster Co., Texas, where it extends from the mountainous regions about Boquillas on the Rio Grande north- ward to slightly beyond Persimmon Gap in the Santiago Mts.”’ Webber (in Yuccas of the S. W., U.S.D.A. Agr. Mon. No. 17:18. 1953) did not enlarge the range of this species given by McKelvey. Apparently it has never been reported outside of Brewster County in the United States. Its locality about and just below the mouth of McKittrick Canyon is only a few miles from New Mexico. It is surprising that so spectacular a plant as this escaped notice before now in the Guadalupe Mountains which have been botanized by numerous botanists through the years. Several old plants, some 15 feet high or more, are scattered in the open scrub forest and chaparral.
ANENOME EDWARDSIANA Tharp var. petraea Correll, var. nov. Vari- etas petraea habitu cum var. edwardsiana congruens sed receptaculis et acheniis omino glabris et vernicosis.
In habit similar to var. edwardsiana but with completely glabrous and vernicose achenes and receptacle. The plants grow so tightly embedded in rock crevices that it is almost impossible to obtain rootstocks.
Kendall Co., in crevices of boulders in canyon below falls, Edge Falls, between Kendalia and Bergheim, Mar. 31, 1965, D. C. Correll & Helen B. Correll 30743 (holotype, LL); in crevices of ledges near Edge Falls, along Curry Creek, 5 miles south of Kendalia, April 14, 1964, D. S. Correll 29141.
Anemone edwardsiana is apparently an endemic species on the Ed- wards Plateau and is not referable to the Coloradan A. tuberosa Rydb. where some recent authors have tended to place it.
RANUNCULUS SARDOUS Crantz, Stirp. Austr. ed. 1 fasc. 2. 84. 1763.
Panola Co., grassy soil, small lake on Alexander Ranch about 4 miles north of Deadwood, flowers yellow, May 9, 1962, D. S. Correll & E. C. Ogden 25188.
This is apparently the first report of this Old World species in Texas. Lyman Benson kindly identified this collection.
CRATAEGUS PEARSONII Ashe, Jour. Elisha Mitchell Sci. Soc. 17:10. 1900.
Jasper Co., south of Jasper off highway 96 between road to Magnolia Springs and road to Roganville, east side of road in woodlands, small trees, Apr. 14, 1963. D. S. Correll & Helen B. Correll 27231; about 7 miles south of Jasper, pine-hardwood forest on Kirbyville road, small tree about 15 ft. tall, flowers white, Apr. 10, 1964, D. S. Correll & Helen B. Correll 29109 ; in mixed forest south of Jasper, tree 15 ft. high, 3 in. in diameter, Apr. 15, 1942, C. L. Lundell & Amelia A. Lundell 11191, in high forest south of Jasper, armed shrub 10 ft. tall, Sept. 9, 1942,C. L. Lundell & S.W. Geiser 11819, 11823.
190 MADRONO [Vol. 19
This species of hawthorn is new to Texas. It has heretofore been known to occur in Mississippi and Louisiana. It is one of the most dis- tinctive species in this extremely complex genus, and it is the only rep- resentative of Series Flavae in our region. This Series is characterized by the conspicuous glands of the inflorescence and on the leaf-teeth and petioles. The mature leaves are also characteristic in having a broad subtruncated apex.
CORNUS RACEMOSA Lam., Encycl. 2:116. 1786.
Bowie Co., in wooded seepage, along McKiney Bayou, on Summer- hill Road #1397, 6.5 miles north of Texarkana, fruits green, pith brown, Aug. 14, 1966, D. S. Correll 33420; same locality, fruits white, pith white, D. S. Correll 33419; same locality, small trees to 12 ft. high, fruits white, pith white, Aug. 30, 1966, D. S. Correll 33587. Cass Co., edge of low forest, about 1.5 miles east of McLeod, small tree to 9 ft. tall, pith dull white, Sept. 14, 1964, D. S. Correll 30097.
This species of dogwood is new to Texas. There are several small colonies in the northeast counties noted above.
NAMA DICHOTOMUM (R. & P.) Choisy in DC., Prodr. 10:182. 1846.
Jeff Davis Co., in gravelly bare soil near summit of Mt. Livermore, Davis Mts., flowers pale lavender, Sept. 20, 1966, D. S. Correll 33744.
According to Lincoln Constance, who kindly contributed a treatment of the Hydrophyllaceae to our MANUAL, this is apparently the first report of this species from Texas.
Proboscidea crassibracteata Correll, sp. nov. Herba annua, ampla, procumbens, ad 1.5 dm lata et ad 5 dm alta, omino glanduloso-pubes- cens; caulis crassus et gravis; folia petiolis gracilibus ad 2.5 dm vel pluribus longis, late triangulo-ovata vel suborbiculato-ovata, cordata, rotundata vel ad apicem obtusa, ad 1.5 dm vel plura longa, longitudo foliorum latitudinem aequans, margine grosse sinuato; flores plures, in racemo brevi gracili ad 10 cm longo prodientes; racemus foliis brevior; pedicelli aliquantum crassi, 2—3 cm longi; bracteae 2, ovato-ellipticae in circumstriptione, obtusae, conspicue carnoso-crassae, circa 8 mm longae et 3.5 mm latae, 2 mm crassae, concavae; calyx tenui-herbaceus, circa 1.5 cm longus, ad apicem irregulatim 5-lobatus, ad basim ventra- liter fissus; corolla tubulari-campanulata, non nisi leviter ventricosa; corollae tubus roseus, circa 4 mm supra basim constrictus, extra sparsim pilosus, circa 2.5 cm longus, orificio 1-1.5 cm lato; faux secus medium lineis paucis croceis et ad ostium pilis brevibus articulatis; corollae lobi 5, subaequales, semiorbiculares, late rotundati, sinibus angustis 4-5 mm profundis, lobo medio luteolo circa 2 cm late ubi expanso; capsulae corpus ellipsoideum, ventrali valde carinatum, 8—9 cm longum, circa 2 cm latum, supra in cornu validum arcuatum, longi- tudine corporis duplo longius protractum.
Plants large, annual, sprawling, as much as 1.5 m across and up to 5 dm high, the herbage glandular-pubescent throughout; stem thick and
1968] CORRELL: FLORA OF TEXAS 191
heavy; leaves with slender petioles up to 2.5 dm long or more, broadly triangular-ovate to suborbicular-ovate, cordate, rounded to obtuse at apex, up to 1.5 dm long or more, about as wide as long, the margins coarsely sinuate; flowers several, borne in a short slender raceme up to 10 cm long that is overtopped by the leaves; pedicels rather stout, 2-3 cm long; bracts 2, ovate-elliptic in outline, obtuse, conspicuously fleshy-thickened, about 8 mm long and 3.5 mm wide, 2 mm thick, con- cave so as to conform with the calyx; calyx thin-herbaceous, about 1.5 cm long, irregularly 5-lobed at apex, ventrally split to the base; corolla tubular-campanulate, only slightly ventricose; corolla-tube noticeably constricted about 4 mm. above base, pink, sparsely pilose without, about 2.5 cm long, 1—-1.5 cm wide across orifice, the throat with a few yellow lines along the middle and some short articulate hairs at its mouth; corolla-lobes 5, subequal, semiorbicular, broadly rounded, the narrow sinuses 4-5 mm deep, the middle lower lobe yellowish, about 2 cm across the lobes when spread out; frut-body ellipsoid, strongly keeled on the ventral side, 8-9 cm long, about 2 cm thick, tapered above into a stout curved horn that is about twice as long as the body.
Presidio Co., on clay flats near Adobes, Rte. No. 170, flowers pink, with yellow line in throat, Sept. 25, 1966, D. S. Correll 33900 (holo- type, LL).
Galium frankliniensis Correll, sp. nov. Planta omnino dense hispi- dula; caules principales e base densa lignea et saepe cum caule prostrato ligneo cortice brunneola; caules aerii manifeste 4-angulati, plerumque porcato-erecti vel ascendentes, ad 3 dm alti; folia 4 in verticillum, ses- silia, elliptica vel elliptico-oblanceolata, obtusa vel acuta et ad apicem cuspidata, ad 8 mm longa et 3 mm lata, marginibus aliquantum revo- lutis, costa aliquantum carinata infra medium infra; flores perfecti, quasi sessiles, raro pedunculo brevi nudo, plerumque in ramis brevissi- mis inflorescentiam ramorum compactorum supra partem foliaceam caulis extensorum facientes; corolla plerumque brunneo-violacea, circa 1.5 mm lata; fructus pilis rectis albis dense velati diametrum fructus aequantibus.
Plant densely hispidulous throughout; main stems from a heavy woody base and often with a prostrate woody stem with brownish bark; aerial stems prominently 4-angled, usually rigidly erect or ascending, up to 3 dm high; leaves in whorls of 4, sessile, elliptic to elliptic-oblance- olate, obtuse to acute and cuspidate at apex, up to 8 mm long and 3 mm wide, the margins somewhat revolute, the central vein somewhat keeled below the middle on the lower surface; flowers perfect, sessile or essen- tially so, rarely on a short naked peduncle, usually on very short branches to form an inflorescence of compact branches that stand rigidly above the leafy portion of stem; corolla mostly brownish purple, about 1.5 mm across; fruits densely covered with straight whitish hairs that are about as long as the diameter of the fruit.
El Paso Co., rock crevices, McKelligon Canyon, Franklin Mts., near
192 MADRONO [ Vol. 19
E] Paso, flowers brown-purple, Aug. 13, 1946, D. S. Correll 13837 (holo- type, SMU); rare perennial in sandstone ledges of McKelligon Canyon, Franklin Mts., July 16, 1949, B. L. Turner 1256 (SMU); infrequent perennial on east lower limestone slopes of Franklin Mts., E] Paso, Apr. 19, 1952, B. H. Warnock 10422; infrequent low herb, limestone soil, east lower slope along the arroyos of the Franklin Mts., about 7 miles from El Paso toward the White Sands, Sept. 29, 1956, B. H. Warnock 14291.
Lauramay T. Dempster, of the University of California, was kind enough to examine these specimens, and, as she pointed out, they are most closely related to G. parishii H. & H., of California and Nevada. The hispidulous indument and elliptic to elliptic-oblanceolate leaves of G. frankliniensis are quite different from that species.
SPERMACOCE FLORIDANA Urban, Symb. Antill. 7:550. 1913. Sperma- coce portoricensis A. Gray, Syn. FI. 1, pt. 2. 34. 1884; Small, Fl. South- east. U.S. 1117. 1903, non Balb.
Cameron Co., in Palm Grove, Mar. 9, 1942, C. L. Lundell 10645 (LL, TEX); July 1942, A. M. Davis s.n. (TEX).
This species, which heretofore has been known from Florida and the West Indies, is new to Texas. It differs from S. tenuior L., its nearest ally in Texas, in that its corolla is essentially glabrous within instead of being noticeably pubescent, and its subglobose glabrous fruit is 1—-1.5 mm long in contrast to the didymous-obovoid mostly puberulent fruit of S. tenuior which is 2—2.5 mm long.
Texas Research Foundation, Renner
NOTES AND NEWS
LIBRARY MAILING RATES FOR HERBARIUM SPECIMENS.— Congress, in its recently passed postal bill, enlarged the concept of library mailing rates to include herbarium specimens as well as other kinds of systematic collections (Sect. 135.14, Postal Man- ual). Specimens sent at library rates must be addressed to an institution, not to an individual, but the package may be marked to the attention of an individual at an institution. Packages cannot exceed 70 pounds each nor a length plus girth of more than 72 inches. In view of the low cost of sending herbarium specimens at this rate, I suggest that we send domestic loans prepaid. It will save us all money in the long run.—JoHN H. Tuomas, Dudley Herbarium, Stanford University.
DRONO
VOLUME 19, NUMBER 6 APRIL, 1968
Contents
NEw Taxa, COMBINATIONS, AND CHROMOSOME NUMBERS IN ARCTOSTAPHYLOS (ERICACEAE), Philip V. Wells 193
CHROMOSOME COUNTS IN SECTION SIMIOLUS OF THE GENUS MIMULUS (SCROPHULARIACEAE). VII. NEw NUMBERS FOR M. GuTTATUS, M. CUPREUS, AND M. Titincu, R. K. Vickery, Jr. K.W. Crook, D.W. Lindsay, M. M. Mia, and W. Tai 211
A Rust on ANEMOPSIS CALIFORNICA, Puul D. Keener and Jack E. Hampton 218
REviEws: Josiah L. Lowe, Polyporaceae of North America. The Genus Poria (Harry D. Theirs) ; Stanwyn G. Shelter, The Komarov Botanical Institute, 250 Years of Russian Research (Robert Ornduff) ; John C. Williams and Howard C. Monroe, A Field Guide to the Natural History of the San Francisco Peninsula (John H. Thomas) 220
Notes AND News: Notes oN MADIA SATIVA AND RELATED SPECIES, V.F. Hesse 210 New PvusLicaTIons; VALIDITY OF NOMENCLATURE CHANGES IN THE FLora oF ALASKA AND YUKON, Eric Hultén; THREE ROADSIDE GRASSES AS RECORDS FOR SACRAMENTO, EL Dorapo, AND PLACER COUNTIES, CALIFORNIA, Beecher Crampton; A NEw FAMILY OF VASCULAR PLANTS (PSILOTACEAE) FOR Arizona, Charles T. Mason, Jr. 222
A WEST AMERICAN JOURNAL OF BOTANY > Pe
IJBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
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MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
Second-class postage paid at Berkeley, California. Return requested. Established 1916. Subscription price $6.00 per year ($4.00 for students). 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 un- delivered copies should be sent to the Corresponding Secretary, California Botanical Society, Department of Botany, University of California, Berkeley, California 94720.
BOARD OF EDITORS
LyMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. ARTURO GOMEZ PomPaA, Universidad Nacional Autonoma de México EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia Mitprep E. Matuias, University of California, Los Angeles ROBERT ORNDUF?, University of California, Berkeley Marion OwnBEY, Washington State University, Pullman REED C. Rotiins, Harvard University, Cambridge, Massachusetts Tra L. Wiccins, Stanford University, Stanford, California
Editor—JoHN H. THomAs Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—J UNE McCAsKILL P.O. Box 23, Davis, California 95616
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Elizabeth McClintock, California Academy of Sciences, San Fran- cisco. First Vice-President: Peter H. Raven, Department of Biological Sciences, Stanford University. Second Vice-President: Cornelius H. Muller, Department of Biological Sciences, University of California, Santa Barbara. Recording Secretary: John West, Department of Botany, University of California, Berkeley. Correspond- ing Secretary: Mary E. Ashton, Department of Botany, University of California, Berkeley. Treasurer: June McCaskill, Department of Botany, University of Cali- fornia, Davis.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Carl W. Sharsmith, Department of Bio- logical Sciences, San Jose State College, San Jose; the Editor or Madrofio; and three elected Council Members: Virginia M. Page, Department of Biological Sci- ences, Stanford University; Annetta Carter, Department of Botany, University of California, Berkeley; and Robert Ornduff, Department of Botany, University of California, Berkeley.
NEW TAXA, COMBINATIONS, AND CHROMOSOME NUMBERS IN ARCTOSTAPHYLOS (ERICACEAE)
Puitire V. WELLS
Arctostaphylos is a predominantly Californian genus of woody plants with more than 100 described taxa. About half of these are based on questionable or trivial differences or do not exist as discrete natural populations. The remainder are well defined entities distinguished by numerous morphological and ecological criteria. It must be admitted that the mixture of well and ill defined taxa, which were practically all pro- posed at the specific level, and the prevalence of hybridization, have made Arctostaphylos a béte noire for many botanists. This has not, how- ever, discouraged work on the genus (Abrams, 1914; 1951; Adams, 1940; Eastwood, 1934; Howell, 1945; Jepson, 1922; 1923; 1925; 1939; McMinn, 1939; Munz, 1959; Schreiber, 1940). Each student has con- tributed to the character analysis, yet much remained, and still remains, to be learned about phenetic patterns in Arctostaphylos.
After a period of nomenclatorial quiescence, there has been a renewed flurry of activity in recent years (Gankin, 1966; 1967; Hoover, 1964; Hoover and Roof, 1966; Roof, 1962; 1964; 1967; Wells, 1961; 1965). The law of diminishing returns seems to be taking effect, but two new species, a number of infraspecific taxa, and a number of new combina- tions seem logical.
I carried out a cytotaxonomic survey (Table I) of many of the species of Arctostaphylos during the academic year, 1966-1967, thanks to re- search facilities and funds in the Department of Botany, University of California. My thanks are also due to G. Ledyard Stebbins for his in- formation and encouragement which served as a stimulus in the present survey, to Marion Cave for her help with cytological preparations, to my industrious assistant, Pat Watters, to T. F. Niehaus for fixed mate- rial of some species, and to H. G. Baker, James B. Roof, and Walter Knight for the many discussions of Arctostaphylos that I inflicted upon them.
A NEw MANZANITA FROM SANTA CATALINA I.
Arctostaphylos catalinae P, V. Wells, sp. nov. Frutex erectus, arbores- cens, 2-6 m altus; caudex etumescens; cortex levis ruber; ramuli glandu- losi, plerumque setosi pilis albis; folia virida, sparseque glandulosa ad bas- im, ovata oblongave, apice acuta obtusave, stomatifera supra et infra, basi truncata cordatave, interdum hastata auriculatave; petioli 2-6 mm longi: inflorescentia paniculata, ramuli saepe 5-6; bracteae foliaceae triangu- lato-lanceolatae, hispidociliatae, dense imbricatae in nascentis inflores- centiis; pedicelli glandulosi; corolla 7-10 mm longa; ovarium dense pilo- sum et sparse glandulosum; drupa glabrata, cinnamomea.
MavbrON0O, Vol. 19, No. 6, pp. 193-224. August 15, 1968. 193
194 MADRONO [Vol. 19
Holotype: eroded andesitic bedrock slopes at head of Bulrush Canyon, Santa Catalina I., Los Angeles Co., California, P. V. Wells 6867, UC.
Tall, erect, arborescent shrub, 2-6 m high, with a well defined trunk and without basal burl. Rare individuals near Haypress show a few weak sprouts from the base of the trunk, but no burl, which may reflect a crown-sprouting ancestry. However, at present A. catalinae is a species of the obligately seeding type. Bark smooth, red-brown. Branchlets dense- ly short hispid and with longer setose hairs, most of the hairs at first gland-tipped. Leaves ovate or oblong, acute; base of leaf polymorphic, varying from subcordate to strongly cordate or sometimes with vestigial auriculation or hastation, to truncate, obtuse or tapering in the same pop- ulation, on petioles 2-6 mm long; blades light green, glandular-ciliate towards the base, along the margin and on the midrib, otherwise glabrous; stomatal distribution isofacial, with stomata above and below. Leaf morphology suggests a common ancestry with a member of the auriculate- leaved group, such as A. viridissima of Santa Cruz I. Inflorescence large, openly paniculate with as many as 5—6 branches from the main rachis; bracts leafy, with a characteristic isoceles-deltoid to lanceolate shape, densely imbricated in the nascent inflorescence, much longer than the very short pedicels which average 2—3 mm in length; rachis and pedicels glan- dular-hispidulous, bracts glandular-ciliate. Corolla 7-10 mm long, white or suffused with pink; ovary densely white-hairy and with sparse gland- tipped hairs; fruit oblate-spheroidal, glabrescent, orange-brown, on straight pedicels; nutlets separable.
This species has been identified as 4. glandulosa Eastw., A. subcordata Eastw., 4. viridissima (Eastw.) McMinn, or A. insularis Greene; in other words, it has been confused with nearly all of the species which occur on the Channel Islands. It combines the characters of the last three of the above mentioned species, none of which presently occur on Santa Cata- lina I. It differs from A. tomentosa ssp. subcordata of Santa Cruz I. and A. glandulosa of the mainland in being an erect, arborescent shrub, lack- ing a basal burl. Arctostaphylos tomentosa ssp. subcordata and A. glan- dulosa are burl-forming, crown-sprouting species which usually produce shrubby colonies; also Arctostaphylos catalinae has a larger, more branched inflorescence than the burl-formers. Some individuals of A. catalinae show vestigial auriculation or hastation of the leaf-base, many others have cordate leaves and most have the leaves more or less imbri- cated, as in A. viridissima of Santa Cruz I. The ample inflorescence and orange-brown, glossy fruits of A. catalinae are traits shared with A. in- sularis of Santa Cruz I. It is A. confertiflora Eastw. of Santa Rosa I. which resembles A. catalinae more closely than any other known species. Contrary to published descriptions (Howell, 1945; Munz, 1959), A. con- fertiflora has isofacial leaves and appears to lack a basal burl. Howell and I established the stomatal distribution on the type specimen (CAS), which was previously thought to have bifacial leaves. Also, a chromosome count indicates that 4. confertiflora is diploid (n = 13). Hence it is
1968] WELLS: ARCTOSTAPHYLOS 195
asserted that A. confertiflora of Santa Rosa I. is specifically distinct from A. tomentosa ssp. subcordata of Santa Cruz I. with which it usually has been placed as a variety or in subjective synonymy; A. tomentosa ssp. subcordata is a bifacial, burl-forming tetraploid (n = 26). Arctostaphy- los catalinae differs from A. confertiflora in at least eight characters given in the following key.
KEY TO THE MANZANITAS OF THE CHANNEL ISLANDS
Bracts of the inflorescence subulate, not leafy; nascent inflorescence with very slen- der, thread-like minutely-bracted branches; stomata restricted to lower surface of leaves; basal burl lacking; fruit bright orange-brown; endemic to Santa Cruz I., on volcanic and metamorphic rocks. . . . . . . . A. insularis Greene
Bracts of inflorescence leafy, lanceolate, deltoid or ovate; nascent inflorescence not filiform, its rachis-branches thick with imbricated foliaceous bracts.
Basal burl present, shrubs crown-sprouting; leaves bifacial, stomata and tomen- tum restricted to lower surface of blade. Branchlets, petioles, rachis, bracts, and pedicels glandular-hairy ; bodenvag, Santa Cruz 1. ates . . A. tomentosa ssp. subcordata (Eastw.) Wells Branchlets, petioles, al nfleretcence tomentose, not glandular; Santa Cruz and Santa Rosa islands, chiefly on volcanic roel A. tomentosa ssp. insulicola Wells Basal burl absent, not crown-sprouting; leaves isofacial, stomata on both surfaces. Leaves auriculate-clasping, short-petioled to nearly sessile; branchlets, petioles, rachis, and bracts densely hispidulous and setose with long white hairs, not glandular; ovary and pedicels tomentose, not glandular; Santa Cruz I, bodenstet on siliceous shale. . . . . A. viridissima (Eastw.) McMinn Leaves cordate to truncate, distinctly petioled, not auriculate- clasping; branch- lets, petioles, rachis, and bracts densely glandular-hairy; ovary and pedicels tomentose and sparsely glandular.
Bracts broadly ovate; inflorescence compact, crowded, with 3-4 branches from main rachis; sepals glandular-ciliate; corolla 5-6 mm long; indu- ment of branchlets and inflorescence glandular-hispid, not long-setose; leaves broadly round-ovate; fruit dull reddish brown; endemic to Santa Rosal.. . . . . . A. confertiflora Eastw.
Bracts narrowly icoscsles deltoid ie Onecohate. inflorescence more open, with 5-6 branches from main rachis; sepals not glandular-ciliate; corolla 7-10 mm long; indument of pence: and inflorescence glandular-hispid and also long-setose with white hairs; leaves oblong-ovate; fruit bright orange-brown; endemic to Santa Catalina I., chiefly on volcanic rocks.
A. catalinae Wells
A NEw AURICULATE-LEAVED MANZANITA FROM SANTA BARBARA CO.
Arctostaphylos purissima P, V. Wells, sp. nov. Frutex erectus, arbores- cens, 1-5 m altus, interdum prostratus in situ extremo; caudex etumes- cens, cortex levis, ruber; ramuli hispiduli plerumque setosi pilis albis; folia virida, circinata-ovata, 1-2 cm in diametro, = sessilia, basi auricu- lata, apice obtusa acutave, stomatifera supra et infra; inflorescentia densa, ramuli 0-3; bracteae foliaceae ovatae, marginibus hispido-ciliatis; pedi- celli glabri, 3 mm longi; corolla globosa, 5-6 mm longa; ovarium gla- brum; drupa 5—8 mm in diametro.
Holotype: on Pleistocene sands, west side of La Purisima ridge, Santa Barbara Co., California; Wells 610671, UC.
196 MADRONO [Vol. 19
Tall, erect, arborescent shrub with a well defined trunk, to over 5 m high in canyons, but sometimes nearly prostrate or forming low mounds on windswept mesa tops near the coast; without basal burl; bark smooth, red-brown; branchlets, rachis and bracts densely hispidulous and setose with white hairs, not glandular; leaves isofacial, closely imbricated, round or round-ovate, relatively small, about 1—2 cm in diameter, nearly sessile or on very short petioles, auriculate- clasping, obtuse or rounded to acute at the apex; leaves bright green with red margins, or on new growth deep crimson-red throughout; inflorescence very compact, simple or with 1-3 branches; lower bracts broadly ovate, foliaceous, hispidociliate on the margins, densely imbricated in the nascent inflorescence; pedicels much shorter than the subtending bracts, ca. 3 mm long; corolla globose, 5—6 mm long; pedicels, ovary and fruit glabrous; fruit oblate-spheroidal, 5— 8 mm in diameter, the nutlets separable.
Arctostaphylos purissima has suffered a fate similar to that of A. luci- ana: lack of recognition due to weighing of the auriculation character. It seems that in a difficult genus there is a tendency to achieve order by weighting conspicuous characters. All entities possessing the trait are made conspecific, as subspecies or even synonyms. This approach to Arctostaphylos taxonomy was pioneered by Jepson (1922; 1923; 1925; 1939), who regarded all auriculate-leaved manzanitas as varieties of A. andersoniu Gray. The distinguishing characters of A. purissima have been consistently overlooked by botanists, including Wells (1965). After ob- serving A. viridissima on Santa Cruz I. in 1967, it became obvious that it is distinct from A. pechoensis and from A. purissima, which for years has been called A. viridissima. The three species share some striking char- acters, but they are readily distinguished by a number of other characters, and have allopatric distributions. Chromosome counts of these and other species of the auriculate-leaved group indicate that all are diploid (n = 13). The following key includes six species occupying disjunct areas from the southern extremity of the Santa Lucia range south to Santa Cruz I.
Key TO AURICULATE-LEAVED MANZANITAS OF SOUTHERN CALIFORNIA
Fruit oblate-spheroidal, not apiculate; nutlets separable, not coalesced as a single stone; branchlets and pedicels not glandular. Branchlets and rachis canescent or tomentose, without setose hairs; inflorescence paniculate.
Leaves and bracts gray-canescent, the bracts not hispid or ciliate; ovary gla- brous; flowering pedicels glabrous, 6-7 mm long, strongly recurved in fruit; nascent inflorescence campanulate-cernuous; flowering panicle ample, openly branched; San Luis Obispo Co., on siliceous Monterey shale in the Santa Lucia Mountains, chiefly southeast of Cuesta Pass. . . A. luciana Wells
Leaves and bracts green, not canescent; bracts hispidociliate on the margin; ovary densely pubescent; flowering pedicels pubescent, 4-5 mm _ long, straight or recurved fruit; nascent inflorescence not as above; flowering panicle dense, compactly branched; San Luis Obispo Co. north to s. Mon- terey Co., Pleistocene sands along the coast. . . . . A. cruzensis Roof
Branchlets and rachis hispidulous and densely setose with long white hairs; bracts hispidociliate ; inflorescence compact, simple or with one or two branches.
1968] WELLS: ARCTOSTAPHYLOS 197
Lower bracts broadly ovate; leaves green, not glaucous, round to round-ovate, 1-2 cm in diameter; fruit 5-8 mm in diameter; pedicels very short, 3 mm long, not recurved in fruit; ovary and pedicels glabrous; corolla globose, 5-6 mm long; Santa Barbara Co., La Purisima area and Burton Mesa north to Pt. Sal, on diatomite, siliceous shale and Pleistocene sands.
A. purissima Wells
Lower bracts lanceolate to linear; leaves ovate to oblong-ovate, 2-4 cm long; fruit 10-15 mm in diameter.
Ovary and pedicels densely pubescent; corolla globose, 5-6 mm long; pedicels very short, 3 mm long, not recurved in fruit; leaves green, not glaucous;
Santa Cruz I., on siliceous shale . . A. viridissima (Eastw.) McMinn Ovary and pedicels glabrous; corolla conical-urceolate, 7-8 mm long; pedicels 8-9 mm long, varying from straight to recurved in fruit; leaves green,
but thinly glaucous with a whitish wax which is readily erased; San Luis Obispo Co., Pecho Hills sector of the San Luis Range, on Monterey shale.
A. pechoensis Dudley
Fruit round or ovoid, apiculate, 10-15 mm in diameter; nutlets coalesced as a solid stone, not separable; branchlets glandular-hispid and setose with gland-tipped hairs; pedicels glandular-puberulent; corolla 8-10 mm long; inflorescence an ample, openly-branched panicle; Santa Barbara Co., vicinity of Refugio Pass, on sandstone. . . ....... +... . . . A. refugioensis Gankin
INFRASPECIFIC TAXA OF ARCTOSTAPHYLOS TOMENTOSA
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. insulicola P, V. Wells, ssp. nov. A ssp. tomentosa differt: cortex levis ruber, non-lacin- i0SuUS.
Holotype: terrace on basaltic rocks above Pelican Bay, Santa Cruz I., Santa Barbara Co., California, P.V. Wells & J. B. Roof 5467, UC.
Tall shrubs to more than 3 m high, taller than ssp. tomentosa; bark smooth, bright red-brown; inflorescence an ample, open panicle.
This subspecies is restricted to Santa Cruz and Santa Rosa islands. The largest known population is on basalt at Pelican Bay; on other types of rocks on Santa Cruz I., ssp. subcordata predominates. Exten- sive intergradation between the two subspecies occurs in some parts of the island. However, on Santa Rosa I., ssp. insulicola is allopatric from other subspecies of A. tomentosa.
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. eastwoodiana P. V. Wells, ssp. nov. A ssp. tomentosa differt: cortex levis ruber, non-la- ciniosus; ovarium glabrum; pedicelli glabri; folia glabra.
Holotype: on diatomite, summit of La Purisima ridge, Santa Barbara Co., California, P. V. Wells 610672, UC.
Eastwood was one of the first to comment on this entity (Roof, 1964), which she correctly perceived as a close relative of A. rosei Eastw. It is fitting to honor her name in Arctostaphylos in which she described so many taxa, in fact, threefold more than any contemporary author.
In Pine Canyon, on Burton Mesa, some individuals of ssp. eastwoodi- ana have the leaves tomentulose below, and/or with sparsely pubescent ovary and pedicels; hence these individuals are phenetic intergrades with ssp. insulicola of the Channel Islands or with ssp. vosei of the central California coast.
198 MADRONO [Vol. 19
The description of two new subspecies of A. tomentosa requires a change in rank of several similar and intergrading taxa.
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. rosei (Eastw.) P. V. Wells, comb. nov. A. rosez Eastw., Leafl. West. Bot. 1:77. 1933; A. crustacea var. roset (Eastw.) McMinn, Ill. Man. Calif. Shrubs 413. 1939.
Subspecies vosez differs from ssp. tomentosa in having glabrous leaves and reddish, not gray bark. It is allopatric from ssp. imsulicola and ssp. eastwoodiana, but contacts and intergrades with ssp. tomentosa and ssp. crustacea on the Monterey Peninsula and at Lake Merced, respectively (Roof, 1964).
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. crustacea (Eastw.) P. V. Wells, comb. nov. A. crustacea Eastw., Leafl. West. Bot. 1:74. 1933.
Subspecies crustacea intergrades with ssp. rosez at Lake Merced, with ssp. eastwoodiana at Pt. Sal, and grades clinally into ssp. tomentost- formis in the Santa Cruz Mountains.
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. tomentosiformis (Adams) P. V. Wells, comb. nov. A. tomentosa var. crinata McMinn, Ill. Man. Calif. Shrubs 412. 1939 (without Latin diagnosis); A. crustacea var. tomentosiformis Adams, Jour. Elisha Mitchel Soc. 56:54. 1940; A. tomentosa var. tomentosiformis (Adams) Munz, Aliso 4:95. 1958.
The nomenclatural history of this subspecies reflects its taxonomic position: intermediate between ssp. crustacea and ssp. tomentosa. It oc- cupies a geographic position between the other two subspecies in the Santa Cruz Mountains and intergrades extensively there with ssp. crus- tacea.
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. subcordata (Eastw.) P.V. Wells, comb. nov. A. subcordata Eastw. Leafl. West. Bot. 1:61. 1933.
Some of the intermediates between ssp. insulicola and ssp. subcordata in the polymorphic populations on Santa Cruz I. have non-glandular, setose hairs and therefore present a combination of characters resembl- ing the disjunct ssp. tomentosiformis. Except for the lack of gray, shred- dy persistent bark, typical ssp. swbcordata closely resembles ssp. tomen- tosa f. trichoclada.
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. TOMENTOSA f. trichoclada (DC.) P. V. Wells, comb. nov. Andromeda bracteosa B trichoclada D.C., Prod. 7(2):607. 1839; Arctostaphylos bracteosa (DC.) Abrams, Leafl. West. Bot. 1:84. 1934; A. tomentosa ssp. bracteosa (DC.) Adams, Jour. Elisha Mitchell Soc. 56:57. 1940; A. tomentosa var. trichoclada (DC.) Munz, Aliso 4:96. 1958.
A glandular form, f. trichoclada, occurs within populations of ssp. tomentosa on the Monterey Peninsula and is distinguished by the glan- dular-setose pubescence of the inflorescence and branchlets. Various degrees of intermediacy exist between f. trichoclada and f. tomentosa,
1968] WELLS: ARCTOSTAPHYLOS 199
but the prevalence of both extreme forms in most populations is sug- gestive of balanced polymorphism. Furthermore, the intensely glandu- lar species, A. montereyensis Hoover, which is sympatric with A. tomen- tosa near Monterey, is a likely source of genes for glandulosity.
ARCTOSTAPHYLOS TOMENTOSA (Pursh) Lindley ssp. TOMENTOSA f. hebeclada (DC.) P. V. Wells, comb. nov. Andromeda bracteosa B hebe- clada DC., Prod. 7(2):607. 1839; Arctostaphylos bracteosa var. hebe- clada (DC.) Eastw., Leafl. West. Bot. 1:122. 1934; A. tomentosa var. hebeclada (DC.) McMinn, Ill. Man. Calif. Shrubs 412. 1939.
This is another intrapopulational forma of ssp. tomentosa on the Mon- terey Peninsula; it has leaves which are glabrescent on the lower sur- face, as in ssp. rose?. However, the gray bark distinguishes it from the latter subspecies.
KEY TO THE INFRASPECIFIC TAXA OF ARCTOSTAPHYLOS TOMENTOSA
Surficial bark gray and persistent as flat shreds, underbark reddish-brown. Pubescence of branchlets and inflorescence tomentose, not glandular. Leaves tomentose on lower surface; Monterey Peninsula and disjunctly south to Los Osos Valley, San Luis Obispo Co. A. tomentosa ssp. tomentosa {. tomentosa Leaves glabrescent; Monterey Peninsula A. tomentosa ssp. tomentosa {. hebeclada (DC.) Wells Branchlets and inflorescence with glandular-setose hairs; Monterey Peninsula. A. tomentosa ssp. tomentosa f. trichoclada (DC.) Wells Surficial bark bright reddish-brown, smooth or shreddy. Branchlets and inflorescence with glandular-setose hairs; bark smooth; Santa Cruz Io... 6A. tomentosa ssp. subcordata (Eastw.) Wells Indumentum not glandular. Branchlets and inflorescence setose with long white hairs; bark smooth. Leaves tomentose below; Santa Cruz Mountains. A. tomentosa ssp. tomentosiformis (Adams) Wells Leaves glabrate below; San Francisco Bay area south to Santa Barbara Co. A. tomentosa ssp. crustacea (Eastw.) Wells Indument not setose. Leaves tomentose below; bark smooth; Santa Cruz and Santa Rosa islands. A. tomentosa ssp. insulicola Wells Leaves glabrous. Ovary and pedicels tomentose; bark more or less shreddy-persistent, but reddish-brown, not gray; San Francisco Bay area south to s. Monterey Co... ... . . . . . A. tomentosa ssp. rosei (Eastw.) Wells Ovary and pedicels glabrous or glabrescent; bark smooth; La Purisima ridge n. to Pt. Sal, Santa Barbara Co. A. tomentosa ssp. eastwoodiana Wells
A NEw SUBSPECIES OF ARCTOSTAPHYLOS PILOSULA
ARCTOSTAPHYLOS PILOSULA Jeps. & Wiesl. ssp. pismoensis P. V. Wells. ssp. nov. A ssp. pilosula differt: frutex arborescens, 2-5 m altus; folia virida, elliptica lanceolatave, basi acuta circinatave; pedicelli 3-6 mm longi.
Holotype: on Pismo sandstone, 3 miles southwest of Edna, San Luis Range, San Luis Obispo Co., California, P. V. Wells 23, UC.
200 MADRONO [Vol. 19
Tall arborescent shrub to more than 5 m high; leaves green, elliptic to lanceolate, with acute to rounded base; pedicels 3-6 mm long. Sub- species pilosula differs in having the leaves slightly to markerly glau- cous, round-ovate and distinctly subcordate to truncate at the base; pedicels 6-7 mm long; and the stature more shrubby, 1-2 m high. Sub- species pilosula is restricted to the interior side of the Santa Lucia Range divide, northwest of Santa Margarita and in the La Panza Range, while ssp. pismoensis is confined to the coastal side of the divide, chiefly in the Pismo sandstone country which forms the eastern sector of the San Luis Range, north of Pismo Beach. Introgression from 4A. glauca Lindley, another species from the interior side of the divide, could account for the main differentiating characters which distinguish the interior ssp. pilosula from the coastal ssp. pismoensis.
SUBSPECIES OF ARCTOSTAPHYLOS HOOKERI
ARCTOSTAPHYLOS HOOKERI D. Don ssp. ravenii P. V. Wells, ssp. nov. A ssp. hookeri differt: frutex repens; folia circinata.
Holotype: on serpentinite, Presidio of San Francisco, San Francisco Co., California, P. V. Wells 2767, UC.
Low shrublet, creeping and the branchlets rooting on contact with the serpentine soil; flowers and fruits small as in ssp. hookeri, but smaller than in ssp. franciscana; leaves round or broadly elliptic, tetraploid (n = 26). This subspecies is only one step from the extinction in na- ture which befell ssp. franciscana. It has been identified as ssp. francis- cana, however, it has the smaller flowers and fruits of the diploid (n = 13) ssp. hookert combined with the creeping habit of ssp. franciscana. The small round leaves are distinctive.
Because of the resemblance of the tetraploid (n = 26) A. montana Eastw. to ssp. ravenii in morphology, ecology, and chromosome number, it is now logical to treat A. montana as another serpentine adapted geo- graphic subspecies of A. hookeri.
ARCTOSTAPHYLOS HOOKERI D. Don ssp. montana (Eastw.) P. V. Wells, comb. nov. A. montana Eastw., Proc. Calif. Acad. Sci. 1:83. 1897.
Subspecies montana differs from the other subspecies of A. hookeri in having densely canescent branchlets, and pedicels varying from straight to strongly recurved in fruit; it differs further from the other serpentine taxa, ssp. ravenii and ssp. franciscana, in being erect to low and mound- form, but not prostrate and creeping. Subspecies montana occupies a re- stricted geographical range to the north of the Golden Gate in Marin Co. where it is allopatric from other subspecies of A. hookert. The canes- cent branchlets and recurved pedicels of ssp. montana suggest introgres- sion from the sympatric A. canescens Eastw. (Stebbins and Major, 1965). Both occur on Mt. Tamalpais, but ssp. montana is restricted to serpentinite bedrock and A. canescens to outcrops of sandstone.
ARCTOSTAPHYLOS HOOKERI D. Don ssp. hearstiorum (Hoover & Roof) P. V. Wells, comb. nov. A. hearstiorum Hoover & Roof, Four Seasons
1968] WELLS: ARCTOSTAPHYLOS 201
2:2. 1966.
This diploid (n = 13) subspecies occurs far to the south near the mouth of Arroyo de la Cruz in San Luis Obispo Co., but recombines some of the characters of the northern subspecies of A. hookert. It differs from ssp. Aookeri in having somewhat smaller leaves on shorter petioles. a smaller inflorescence, and a consistently creeping growth habit.
KEY TO THE SUBSPECIES OF ARCTOSTAPHYLOS HOOKERI
Fruit 4-5 mm in diameter; corolla 4-5 mm long.
Erect or forming low mounds, but not prostrate and creeping; leaves narrowly oblanceolate, lanceolate, elliptic or broadly round-elliptic; inflorescence with about 6-10 flowers; Monterey Peninsula north to the Santa Cruz Mountains. on siliceous substrata. . . . . . . . A. hookeri ssp. hookeri
Prostrate and rooting along the Sean pranenices occurring north or south of the above range.
Leaves broadly round-elliptic; inflorescence with about 6-10 flowers; Presidio of San Francisco, restricted to serpentinite bedrock. A. hookeri ssp. ravenii Wells Leaves elliptic to lanceolate; inflorescence with about 3-6 flowers; San Luis Obispo Co., restricted to sandy soil on bluffs near mouth of Arroyo de la Cruz... . . . . A. hookeri ssp. hearstiorum (Hoover & Roof) Wells Fruit 6-8 mm in diameter; corolla 5—7 mm long.
Erect or forming low mounds, but not prostrate and creeping; leaves broadly elliptic; branchlets densely canescent; Marin Co., on serpentinite at Mt. Tam- alpais, on granite at Inverness Ridge. A. hookeri ssp. montana (Eastw.) Wells
Prostrate and creeping; leaves narrowly elliptic or oblanceolate; branchlets pu- berulent, not canescent; San Francisco, now extinct in nature, formerly from Laurel Hill and Masonic Cemeteries, growing on serpentinite.
A. hookeri ssp. franciscana (Eastw.) Munz
SUBSPECIES OF ARCTOSTAPHYLOS MANZANITA
ARCTOSTAPHYLOS MANZANITA Parry ssp. wieslanderi P. V. Wells, ssp. nov. A ssp. manzanita differt: ramuli et petioli minute glanduolsi: ramuli inflorescentiarium glandulosi; bracteae glandulosae.
Holotype: black oak forest at 1,950 ft. elevation, Luck Creek, Shasta Co., California, J. B. Griffin 1267, UC.
Subspecies wieslandert was known to A. E. Wieslander and members of the Vegetation Type Map Survey as “A. mx.” and it is fitting to name it after a keen student of Arctostaphylos. This subspecies occupies an elevation belt above ssp. manzanita, with which it intergrades clin- ally around the northern perimeter of the Sacramento Valley.
It is now necessary to make a number of new combinations to estab- lish a logical ordering of other taxa which closely resemble A. manzanita.
ARCTOSTAPHYLOS MANZANITA Parry ssp. elegans (Jeps.) P. V. Wells. comb. nov. A. elegans Jeps., Erythea 1:15. 1893; A. manzanita var. ele- gans (Jeps.) Benson, Am. Jour. Bot. 27:189. 1940.
This subspecies grows in an elevation zone above ssp. manzanita in the southern sector of the inner North Coast Range.
ARCTOSTAPHYLOS MANZANITA Parry ssp. roofii (Gankin) P. V. Wells. comb. nov. A. roofii Gankin, Leafl. West. Bot. 10:329. 1966.
202 MADRONO [Vol. 19
The exserted stigma cited in the type description is a trait commonly found in ssp. manzanita. Gankin (1966) writes: “The striking similarity of this species to A. manzanita is a curious deception. The upper limit of A. manzanita in this region seems to be between 1,000 and 1,500 feet elevation. Arctostaphylos elegans appears to replace A. manzanita in the intervening areas, but only sparsely.” As with ssp. wieslanderi and ssp. elegans, ssp. roofit occupies an elevation belt above ssp. manzanita occurring between 3,200 and 4,300 feet. However, it is more restricted in distribution than the other taxa, being confined to a small area in the North Coast Range of Glenn Co. The intrapopulational variability of the ovary indument from glandular-pubescent to pubescent to glabrous suggests the possibility of hybridity involving A. glandulosa Eastw., a burl-forming species with pubescent to glandular-pubescent ovaries, and ssp. manzanita, a non-sprouting subspecies with a glabrous ovary. The non-sprouting ssp. elegans may represent a different recombination of characters stemming from a similar cross. Chromosome counts of all three taxa indicate that they are tetraploid (n = 26).
ARCTOSTAPHYLOS MANZANITA Parry ssp. bakeri (Eastw.) P. V. Wells, comb. nov. A. bakert Eastw. Leafl. West. Bot. 1:115, 1934; A. stan- fordiana Parry ssp. bakeri (Eastw.) Adams, Jour. Elisha Mitchell Soc. 56:19. 1940.
The taxonomic position of this highly localized serpentinite endemic of Sonoma Co. has long been uncertain. It has little in common with 4. stanfordiana with which it was placed as a subspecies. The apparent reason for this disposition is that in terms of a key character, the glan- dulosity of branchlets and inflorescences, it is similar to A. hispidula Howell, a distinct species of serpentinite habitat in Del Norte Co., which is closely related to A. stanfordiana. It is clear that with respect to nas- cent inflorescence, flowers, fruit, and chromosome number (n = 26), the relationship of ssp. bakervi lies with A. manzanita. Populations of ssp. manzanita on serpentinite bedrock in Sonoma Co., as at The Cedars, north of Cazadero, bear a striking resemblance to ssp. bakeri in stature, foliage, and compactness of the inflorescence, but lack the glandulosity. Subspecies baker differs from other subspecies of A. manzanita in com- bining glandular-hispid pubescence of branchlets, rachis, and bracts with somewhat smaller leaves and more compact inflorescence.
KEY TO THE SUBSPECIES OF ARCTOSTAPHYLOS MANZANITA
Basal burl lacking. Branchlets, rachis, and bracts puberulent or glabrescent, but not glandular. Ovary and fruit glabrous; branchlets and rachis puberulent.
Branches of nascent inflorescence elongate, slender, not clavate-thickened; widespread at lower elevations in the North Coast Ranges, around the head of the Sacramento Valley, and southward in the Sierra Nevada.
A. manzanita ssp. manzanita
Branches of nascent inflorescence short, clavate-thickened as in A. pungens HBK.; Contra Costa Co., endemic to Mt. Diablo and vicinity.
A. manzanita ssp. laevigata (Eastw.) Munz
1968] WELLS: ARCTOSTAPHYLOS 203
Ovary and fruit with glandular, deciduous hairs; branchlets and rachis glabres-
cent; occupying a zone above ssp. manzanita in the North Coast Ranges. A. manzanita ssp. elegans (Jeps.) Wells Branchlets, rachis, and bracts minutely glandular-pubescent.
Inflorescence a large, ample, openly branched panicle; leaves broadly ovate; oc- cupying a zone above ssp. manzanita around the head of the Sacramento Valley. . . . : . A. manzanita ssp. wieslanderi Wells
Inflorescence ae more neonipac: (eaves elliptic to ovate; Sonoma Co., en- demic to serpentinite outcrops near Occidental.
A. manzanita ssp. bakeri (Eastw.) Wells Basal burl present; occupying a zone above ssp. manzanita on the eastern slope of the North Coast Range in Glenn Co. A. manzanita ssp. roofii (Gankin) Wells
TAXONOMIC CONFUSION CONCERNING THE NON-SPROUTING SUBSPECIES oF ARCTOSTAPHYLOS PATULA
ARCTOSTAPHYLOS PATULA Greene ssp. platyphylla (Gray) P. V. Wells, comb. nov. A. pungens var. platyphylla Gray, Syn. Fl. N. Am. 2:28. 1878; A. platyphylla (Gray) Kuntze, Rev. Gen. 2:385. 1891; A. obtusifolia Piper, Bull. Torrey Club 26:642. 1902; A. pinetorum Rollins, Rhodora 39:462. 1937; A. parryana Lemmon var. pinetorum (Rollins) Wiesl. & Schreib., Madrono 5:46. 1939.
Away from the Sierra Nevada, Cascade Mountains, and North Coast Ranges, where the typical crown-sprouting form occurs, most popula- tions of A. patula lack the burl at the base of the stem. Morphologically similar non-sprouting forms of the conifer/chaparral zone extend east- ward to Utah and Colorado, northward into Oregon on the dry eastern side of the Cascade Mountains, and disjunctly southward in the San Gabriel, San Bernardino, and Santa Rosa mountains of southern Cali- fornia.
The geographically diverse, non-sprouting populations seem to resem- ble typical A. patula very closely, aside from absence of the burl, though careful examination of living populations may yield additional differ- entiae. However, at present, Gray’s epithet, platyphylla, has priority for all non-sprouting populations of A. patula.
On the northeastern side of the Sacramento Valley, an erect form of A. patula ssp. platyphylla occupies a zone of elevation below that of the burl-forming ssp. patula and above that of the erect, non-sprouting A. manzanita ssp. wieslanderi. The zonation, which was called to my atten- tion by James Griffin, is as follows: 1, A. manzanita ssp. manzanita, low- est zone, down to the valley floor; 2, A. manzanita ssp. wieslanderi; 3, A. patula ssp. playphylla; and 4, A. patula ssp. patula, highest zone. Intermediates occur in populations of adjacent zones, but available in- formation is insufficient to establish the existence of a cline along the entire gradient. Both A. manzanita ssp. wieslandert and A. patula ssp. platyphylla may have originated in this area by hybridization of A. manzanita ssp. manzanita and A. patula ssp. patula, with subsequent backcrossing to one or the other parent, but it would be unwarranted to extend this speculation to account for loss of the burl in other popu-
204 MADRONO [Vol. 19
lations of A. patula. It is likely that loss of the burl has occurred inde- pendently in the geographically disjunct populations of A. patula ssp. platyphylla.
SUBSPECIES OF ARCTOSTAPHYLOS VISCIDA
ARCTOSTAPHYLOS VISCIDA Parry ssp. pulchella (Howell) P. V. Wells. comb. nov. A. pulchella Howell, Fl. N. W. Am. 1:146. 1901.
Ranging through the North Coast Ranges, north to the Siskiyou Mountains of southern Oregon, ssp. pulchella differs from the glabrous- fruited ssp. viscida of the Sierra Nevada in having the ovary and fruit glandular-viscid.
ARCTOSTAPHYLOS VISCIDA Parry ssp. mariposa (Dudley) P. V. Wells, comb. nov. A. mariposa Dudley, Publ. Sierra Club 27:52. 1902.
This subspecies is more glandular than ssp. pulchella; not only the fruit, but also the branchlets, petioles, rachis, and bracts are densely glandular-pubescent. It occurs in the same elevation zone in the Sierra Nevada as ssp. viscida, but largely replaces the latter in the southern sector from Amador Co. south to Kern Co. Even at the southern extrem- ity of the range of the species, variations to the glabrous condition of ssp. viscida occur, and very extensive intergradation between the glandu- lar and non-glandular condition exists in the central Sierra Nevada, where ssp. mariposa merges with ssp. viscida. Also, as far north as south- ern Oregon, far from the range of ssp. mariposa, individuals with glan- dular-pubescent branchlets occur sporadically in glabrous-branched populations of ssp. pulchella.
KEY TO THE SUBSPECIES OF ARCTOSTAPHYLOS VISCIDA
Ovary and fruit glabrous, not viscid; branchlets and leaves usually glabrous; Sierra Nevada from Kern Co. north to Shasta Co. . . . . A. viscida ssp. viscida Ovary and fruit densely glandular-viscid. Branchlets, leaves, and inflorescence densely glandular-pubescent; Sierra Nevada from Kern Co. north to Amador Co. A. viscida ssp. mariposa (Dudley) Wells Branchlets and leaves mostly glabrous; inflorescence varying from glabrescent to sparingly glandular on rachis and bracts; North Coast Ranges from Napa Co. north to southern Oregon. . . A. viscida ssp. pulchella (Howell) Wells
INFRASPECIFIC TAXA OF ARCTOSTAPHYLOS GLANDULOSA
ARCTOSTAPHYLOS GLANDULOSA Eastw. ssp. GLANDULOSA f. cushingiana (Eastw.) P. V. Wells, comb. nov. A. cushingiana Eastw., Leafl. West. Bot. 1:175. 1933; A. glandulosa var. cushingiana (Eastw.) McMinn, Il. Man. Calif. Shrubs 417. 1939.
The eglandular “A. cushingiana” and the glandular A. glandulosa occur as alternate intrapopulational forms (morphs) over much of the range of ssp. glandulosa. The parallel to the more localized polymorph- ism of A. tomentosa and its glandular form, f. ¢richoclada, is striking. Other morphological variants of the A. glandulosa complex have definite and largely allopatric geographic distributions and are recognized as subspecies.
1968] WELLS: ARCTOSTAPHYLOS 205
ARCTOSTAPHYLOS GLANDULOSA Eastw. ssp. howellii (Eastw.) P. V. Wells, comb. nov. A. Aowellit Eastw., Leafl. West. Bot. 1:123. 1934; A. glandulosa var. howellu (Eastw.) McMinn, Ill. Man. Calif. Shrubs 417. 1939.
ARCTOSTAPHYLOS GLANDULOSA Eastw. ssp. zacaensis (Eastw.) P. V. Wells, comb. nov. A. zacaensis Eastw., Leafl. West. Bot. 1:79. 1933; A. glandulosa var. zacaensis (Eastw.) McMinn, Ill. Man. Calif. Shrubs 417. 1939,
ARCTOSTAPHYLOS GLANDULOSA Eastw. ssp. mollis (Adams) P. V. Wells, comb. nov. A. glandulosa var. mollis Adams, Jour. Elisha Mitchell Soc. 56:50. 1940.
ARCTOSTAPHYLOS GLANDULOSA Eastw. ssp. crassifolia (Jeps.) P. V. Wells, comb. nov. A. glandulosa var. crassifolia Jeps., Madrono 1:86. 1922; A. tomentosa var. crassifolia (Jeps.) Jeps., Man. FI. Pl. Calif. 749, 1925.
A WIDELY DISJUNCT SUBSPECIES OF ARCTOSTAPHYLOS UVA-URSI IN GUATEMALA
ARCTOSTAPHYLOS CRATERICOLA (J. D. Smith) J. D. Smith is the name applied to a creeping manzanita found on the summits of lofty volcanos and other mountain peaks over 10,000 ft. high in Guatemala. It has been misunderstood, probably because it was originally described as a variety of A. pungens HBK., which is the only other species of Arctostaphylos ranging south through Mexico. It extends as far south as Oaxaca, Mex- ico. The many Mexican species of Comarostaphylis with synonyms in Arctostaphylos have added to the confusion.
An examination of a series of flowering and fruiting specimens of A. cratericola now available at US and NY shows that it has little in com- mon with A. pungens, but is very similar to A. uva-ursi, differing mainly in the indument of the pedicels.
ARCTOSTAPHYLOS UVA-URSI (L.) Spreng. ssp. cratericola (J. D. Smith) P. V. Wells, comb. nov. A. pungens HBK. var. cratericola J. D. Smith, Bot. Gaz. 16:13. 1891; A. cratericola (J. D. Smith) J. D. Smith, Bot. Gaz. 55:437. 1913.
Prostrate, creeping and rooting along the branchlets, forming mats; branches with rough shreddy bark, the branchlets sparsely puberulent; leaves often resupinate on the prostrate branches, bifacial, the stomata restricted to the lower surface, spatulate or oblanceolate to elliptic, acute or apiculate to obtuse or rounded at the tip, cuneate at the base; petiole and leaf margins sparsely puberulent; inflorescence a dense, short simple cluster; rachis sparsely puberulent; bracts sparsely puberulent with glandular or eglandular hairs; pedicels varying from glandular-puberu- lent to glabrous; fruit red, shining, glabrous, oblate-spheroidal, the nut- lets separable (previously, the fruit was unknown. )
Subspecies cratericola differs from ssp. uva-ursi in having pedicels varying from minutely glandular-puberulent to glabrous, instead of
206 MADRONO [Vol. 19
consistently eglandular and usually glabrous; also the leaves vary from spatulate to oblanceolate or elliptic and from apiculate to obtuse at the tip, while ssp. wva-ursi shows leaf shape variation from spatulate to obovate, rarely elliptic, with the leaf tip consistently obtuse, rarely apic- ulate. The holotype of ssp. cratericola has markedly elliptic leaves, an unusual variation, and this may have been enough to obscure its rela- tionship to A. uva-urst.
Subspecies cratericola is known from the summits of three high moun- tain peaks in three different Departamentos of Guatemala: Volcan de Agua, Dept. Zacatepequez, at elevations from 8,000 to 12,300 ft. (holo- type at 12,100 ft.); Volcan Zunil, Dept. Quezaltenango, from 9,850 to 12,500 ft.; and Sierra de los Cuchumatanes, Dept. Huehuetenango, from 10,800 to 11,500 ft., on limestone. These are indeed remarkably dis- junct occurrences of A. wva-ursi, which is unknown from all of Mexico, the southernmost stations being in New Mexico. It is suggestive to note that a collection from Hermit’s Peak, San Miguel Co., N. M., Wells 42962, KANU, has the glandular branchlets and bracts of A. uva-ursi var. adenotricha Fern. & Macbr.; also the leaves are apiculate and oblanceo- late, but the pedicels are glabrous.
A CYTOTAXONOMIC SURVEY OF ARCTOSTAPHYLOS
Chromosome counts have been published for only a few species and all have the base number 13. This number has been observed in other genera of Ericaceae, including the closely related Arctous and X ylococ- cus. During the year 1966-1967, I was able to fix meiotic material of about 58 taxa, nearly all of which yielded good cytological preparations. The data are summarized in Table I.
Although the size of the genus prohibited a series of counts for each taxonomic unit, enough information has been gleaned to show consistent patterns, which have predictive value. In brief, two levels of ploidy exist: n — 13 and n = 26. A leading generalization is that the many races of the leafy-bracted burl-forming species, A. tomentosa and A. glandulosa, are tetraploid (n = 26); they are confined to the coastal half of California, chiefly within the fog belt. On morphological and eco- logical grounds they appear to be relatively less specialized than the non- sprouting species with fire-stimulated germination, which occupy a wider range of habitats, and have a vastly greater geographic distribu- tion. The fact that two burl-forming species with reduced bracts are diploid (n = 13), namely A. patula and A. rudis, shows that tetraploidy is not constantly associated with burl-formation, although this may be a predisposing condition. The much greater life-span of the burl-form- ers under the conditions of frequent fire prevalent in chaparral would also tend to favor the preservation of individuals having undergone a change in chromosome number, thus affording a longer time period in which selection could operate on their progeny.
1968] WELLS: ARCTOSTAPHYLOS 207
TABLE I, CHROMOSOME NUMBERS IN ARCTOSTAPHYLOS. RPBG = PLANTS PROPA-
GATED IN THE REGIONAL PARKS BOTANIC GARDEN, BERKELEY. UCBG = FROM PLANTS PROPAGATED IN THE UNIVERSITY OF CALIFORNIA BOTANICAL GARDEN. PREPARED SLIDES, CAMERA LUCIDA DRAWINGS, AND VOUCHER SPECIMENS ARE AVAILABLE (KANU).
-S RRRR BR BRab
-_ ae Ro bp RB BR Bb
andersoni Gray. n = 13. Santa Cruz Co., Big Basin. Wells 25672.
auriculata Eastw. n = 13. Contra Costa Co., Mt. Diablo, RPBG. Wells 2867. canescens Eastw. n = 138. Marin Co., Mt. Tamalpais. Wells 11560. columbiana Piper. n = 13. Del Norte Co., Darlingtonia Wells 21267 ; UCBG. Wells 223671.
confertiflora Eastw. n = 13. Santa Barbara Co., Santa Rosa I., UCBG. Wells 2250/2.
cruzensis Roof. n = 13. San Luis Obispo Co., Los Osos Valley. Wells 1867. densiflora Baker. n = 13. Sonoma Co., Vine Hill, UCBG. Wells s.n.
. edmundsii Howell. n = 13. Monterey Co., Little Sur River. Wells 919661. . edmundsii Howell X A. tomentosa ssp. rosei (Eastw.) Wells. n = 13. Monterey
Co., Garrapata Creek. Wells 919662.
. glandulosa Eastw. n = 26. Marin Co., Mt. Tamalpais Wells 24671; Wells 24672. . glandulosa f. cushingiana (Eastw.) Wells. n = 26. Mendocino Co., Trinity Grade.,
UCBG. Wells 419671; Marin Co., Mt. Tamalpais. Wells 24673; Wells 24674; San Luis Obispo Co., La Panza Range, UCBG. Wells 419672.
glandulosa ssp. zacaensis (Eastw.) Wells. n = 26. Santa Barbara Co., Figueroa Mt. Wells 43067.
glauca Lindley. n = 13. San Luis Obispo Co., Pozo, UCBG. Niehaus 271; San Diego Co., Dulzura, UCBG. Niehaus 272.
glutinosa Schreib. n = 13. Santa Cruz Co., Eagle Rock, RPBG. Roof s.n.
hookeri D. Don ssp. hookeri. n= 13. Monterey Co., Monterey Bay, UCBG. Wells 84671.
hookeri ssp. hearstiorum (Hoover & Roof) Wells. n = 13. San Luis Obispo Co., Arroyo de la Cruz, RPBG. Roof s.n.
. hookeri ssp. montana (Eastw.) Wells. n = 26. Marin Co., Mt. Tamalpais. Wells
42467 ; UCBG. Wells s.n.
. hookeri ssp. raveni? Wells. n = 26. San Francisco Co., San Francisco, Presidio.
Wells 2767.
. imbricata Eastw. n = 13. San Mateo Co., San Bruno Mt., UCBG. Wells 223673. . insularis Greene f. insularis.n = 13. Santa Barbara Co., Santa Cruz I., UCBG.
Wells 124671.
insularis Greene f. pubescens (Eastw.) P. V. Wells, comb. nov. A. insularis Greene var. pubescens Eastw. Leafl. West. Bot. 1:62. 1933. n = 13. Santa Barbara Co., Santa Cruz I., Christie’s. Niehaus 484. This form with glandular-pubescent branchlets and tomentose ovary is the prevalent one on the island. The glabrous nominate form occurs sporadically as an intra-populational morph, chiefly on the interior ridges. Introgression of genes from the sympatric, glandular species, A. tomentosa ssp. subcordata (Eastw.) Wells, is a suspected cause of the poly- morphism.
. luciana Wells. n = 13. San Luis Obispo Co., Santa Lucia Mts. Wells 1767; UCBG.
Wells 223674.
. manzanita Parry ssp. manzanita. n = 26. Lake Co., N. of Mt. St. Helena, RPBG.
Roof, s.n.
. manzanita ssp. bakeri (Eastw.) Wells. n = 26. Sonoma Co., Occidental, UCBG.
Wells 12567. manzanita ssp. elegans (Jeps.) Wells. n = 26. Lake Co., Lower Lake, UCBG. Niehaus 255.
. mewukka Merriam. n = 26. El Dorado Co., Peavine Ridge. Wells 326673; Wells
326674; Wells 326675.
208 MADRONO [Vol. 19
pln Him jhe ple ple
phe
TABLE I. (Continued).
. morroensis Wiesl. & Schreib. n = 13. San Luis Obispo Co., Morro Bay. Wells
22667 ; Hazard Canyon, UCBG. Wells s.n.
. nevadensis Gray. n = 26. Siskiyou Co., Mt. Shasta. Wells 7167. . nissenana Merriam. n = 13. El Dorado Co., Diamond Springs, UCBG. Wells
124672.
. nummularia Gray ssp. nummularia. n = 13. Mendocino Co., Mendocino Plains,
UCBG. Wells 84673.
.nummularia Gray ssp. sensitiva (Jeps.) P. V. Wells, comb. nov. A. sensitiva Jeps.
Madrono 1:85. 1922. A. nummularia var. sensitiva (Jeps.) McMinn. Ill. Man. Calif. Shrubs 389. 1939. n = 13. Marin Co., Mt. Tamalpais. Wells 51367. Sub- species nummularia differs from ssp. sensitiva chiefly in growth habit. However, ssp. nummularia is quite variable in growth habit, ranging from prostate to al- most as erect as ssp. sensitiva. Hybridization with the prostrate A. uva-ursi (L.) Spreng. is evident near the coast, and probably accounts for the divergence of the maritime ssp. nummularia from the montane ssp. sensitiva.
. obispoensis Eastw. n = 13. San Luis Obispo Co., Santa Lucia Mts., RPBG. Roof
s.n.
. otayensis Wiesl. & Schreib. n = 13. San Diego Co., Otay Mt., RPBG. Wells 2667. . pajaroensis Adams. n = 13. Monterey Co., Pajaro Hills. Wells 919663. . pajaroensis Adams X A. tomentosa (Pursh) Lindley. n = 13. Monterey Co.,
Pajaro Hills, RPBG. Wells 92566.
. pallida Eastw. n = 13. Alameda Co., Huckleberry Ridge. Wells 1124661 ; Manza-
nita Drive. Wells 1124662.
. parryana Lemmon. n = 26. Kern Co., Mt. Abel. Wells 6767; Ventura Co., Mt.
Pinos, RPBG. Roof s.n.
. patula Greene. nn = 13. El] Dorado Co., Peavine Ridge. Wells 326672. . patula Greene X A. viscida Parry. n = 13. El Dorado Co., Peavine Ridge. Wells
320071.
. pechoensis Dudley. n = 18. San Luis Obispo Co., See Canyon. Wells 1023662. . pilosula ssp. pismoensis Wells. n = 13. San Luis Obispo Co., Edna, UCBG. Wells
12567.
. pumila Nutt. n = 13. Monterey Co., Fort Ord, UCBG. Wells 84672.
. purissima Wells. n = 13. Santa Barbara Co., Lompoc, UCBG. Wells 223673.
. refugioensis Gankin. n = 13. Santa Barbara Co., Refugio Pass. Wells 5667.
. regismontana Eastw. n = 13. San Mateo Co., King’s Mt., RPBG. Roof s.n.
. X repens (Howell) P. V. Wells, comb. nov. A. cushingiana f. repens Howell,
Leafl. West. Bot. 4:161. 1945. n = 13. Marin Co., Mt. Vision, UCBG. Wells 41967. Howell’s name is based on hybrid individuals produced in a small area on Inverness Ridge where A. uva-ursi (L.) Spreng., A. virgata Eastw. and A. glan- dulosa Eastw. are sympatric.
. rudis Jeps. & Wiesl. n = 13. San Luis Obispo Co., Nipomo Mesa. Wells 1023661. . stanfordiana Parry. n = 13. North Coast Range, RPBG. Wells 223674. . tomentosa (Pursh) Lindley ssp. tomentosa. n = 26. Monterey Co., Monterey
Bay dunes, RPBG. Roof s.n.; San Luis Obispo Co., Arroyo de la Cruz. Wells 42667 ; Los Osos Valley. Wells 1867.
. tomentosa ssp. crustacea (Eastw.) Wells. n = 26. Contra Costa Co., Moraga
Ridge UCBG. Wells 84674.
. tomentosa ssp. insulicola Wells. n = 26. Santa Barbara Co., Santa Cruz I, Peli-
can Bay. Wells 5467 ; Laguna Canyon. Wells 53671.
. tomentosa ssp. rosei (Eastw.) Wells. n = 26. Monterey Co., Bixby Cr., UCBG.
Wells 84675.
. tomentosa ssp. subcordata (Eastw.) Wells. n = 26. Santa Barbara Co., Santa
Cruz I., China Harbor. Wells 5267 ; Christie’s. Wells 53672.
1968] WELLS: ARCTOSTAPHYLOS 209
TABLE I. (Continued).
A. tomentosa ssp. tomentosiformis (Adams) Wells. n = 26. Santa Cruz Co., Felton. Wells 25671. A. virgata Eastw. n = 138. Marin Co., Mt. Tamalpais. Wells 2467. A. viridissima (Eastw.) McMinn. n = 13. Santa Barbara Co., Santa Cruz I., China Harbor. Niehaus 501. . viscida Parry ssp. viscida. n = 13. Amador Co., Ione, UCBG. Wells 84677. . viscida ssp. mariposa (Dudley) Wells. n = 13. Sierra Nevada, RPBG. Roof s.n.
poe pee
For some tetraploids, there is morphological evidence of hybridity in- volving existing diploid species, and the distributional and ecological facts provide strong support for their origin through amphidiploidy. One example is the tetraploid Arctostaphylos newukka Merriam. It oc- curs as sporadic colonies which are restricted to a zone of elevation where the distributions of two diploid species, A. patula Greene and A. viscida Parry, overlap in the Sierra Nevada. A character analysis em- ploying 50 morphological traits shows that A. mewukka combines the characters of A. patula and A. viscida in the following way: characters shared with A. viscida exclusively, 12; characters shared with A. patula exclusively, 7; characters shared with both A. viscida and A. patula, 30; transgressive characters, 1 (fruit size). The morphology indicates hy- bridization of A. viscida with A. patula, followed by stabilization of the A. mewukka character combination; the cytology suggests that stabiliza- tion was achieved through amphidiploidy.
Department of Botany, University of Kansas, Lawrence
LITERATURE CITED
ABRAMS, L. 1914. Uva-ursi. Jn North American Flora 29:92-102. —. 1951. Illustrated Flora of the Pacific States, Vol. III. Stanford University Press. ApaMs, J. E. 1940. A systematic study of the genus Arctostaphylos Adans. Jour. Elisha Mitchell Soc. 56:1-62. Eastwoop, A. 1934. A revision of Arctostaphylos. Leafl. West. Bot. 1:105-127. GANKIN, R. 1966. A new species of Arctostaphylos from Glenn County, California. Leafl. West. Bot. 10:329-331. -————.. 1967. A new species of Arctostaphylos from Santa Barbara County, Cali- fornia. Four Seasons 2(2):13-14. Hoover, R. F. 1965. A new manzanita from Monterey. Leafl. West Bot. 10:87-90. .. and J. B. Roor. 1966. Two new shrubs from San Luis Obispo County, California. Four Seasons 2(1) : 2-4. Howe tt, J. T. 1945. Concerning stomata on leaves of Arctostaphylos. Wasmann Jour. Biol. 6:57-63. Jepson, W. L. 1922; 1923. Revision of the California species of the genus Arcto- staphylos. Madrofio 1:67—86; 87-96. . 1925. A manual of the flowering plants of California. Associated Students Store, Berkeley. . 1939. A flora of California, Vol. III. Associated Students Store, Berkeley.
210 MADRONO [Vol. 19
McMiny, H. E. 1939. An illustrated manual of California shrubs. Univ. Calif. Press, Berkeley. Mowz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley. Roor, J. B. 1962. Two new species of Arctostaphylos from California. Leafl. West. Bot. 9:217-222. . 1964. The nature and distribution of Arctostaphylos rosei Eastwood. Four Seasons 1(1):1-15. . 1967. Arctostaphylos montaraensis, a new species of manzanita from San Mateo County, California. Four Seasons 2 (3) :6—-16. SCHREIBER, B. O. 1940. The Arctostaphylos canescens complex. Am. Midl. Nat. 23: 617-632. STEBBINS, G. L., and J. Mayor. 1965. Endemism and speciation in the California flora. Ecol. Monogr. 35:1-35. WELLS, P. V. 1961. A new manzanita from the Santa Lucia Range, California. Leafl. West Bot. 9:152-153. . 1965. A misplaced manzanita from the Santa Lucia Range, California. Leafl. West. Bot. 10:176-178.
NOTES AND NEWS
Notes ON MapIA SATIVA AND RELATED SPECIES.—M adia capitata Nutt., M. sativa Molina and M. gracilis (Smith) Keck are closely related and often difficult to dis- tinguish. They tend to intergrade and diagnoses in current manuals lack clearcut characteristics by which to delimit the species. In 1965, at the suggestion of John Thomas Howell, I checked the odors of these plants in the field in the Santa Cruz Mountains, central California. Odor is one of the distinguishing characteristics given, although not a very satsfactory one, for two reasons: 1, it applies only to fresh material, and 2, there are no names for odors as such. Names for odors are always given as similar to other, supposedly universally known ones, or in terms of subjective reaction, which necessarily varies from person to person.
The first discovery of this investigation was rather surprising. The troublesome forms intermediate between M. sativa and M. gracilis smelled different from either of these species, being basically similar but lacking both the spicy fragrance of gracilis and the heavy unpleasant odor of sativa. These considerations led me to speculate that another distinct entity probably was involved and, with the encour- agement of Rimo Bacigalupi, I attempted a diagnosis of this “new” variety (as I would have evaluated it) under an appropriately descriptive epithet, based on Hesse 3318 (JEPS). This analysis proved inconclusive because morphological char- acters proved to be varyingly intermediate, with the single exception of the achenes, which were found to be consistently small (only a little longer than those of gracilis) and slender; in shape most nearly like those of capitata. Howell had further commented that probably sometime someone would conclude that all these species were in reality one variable species. If so, the earliest name, M. sativa, should be used. There is much to be said in favor of this idea. For instance, my specimens, 3315 (CAS), 3317, 3237, 3316, 3318, 3234, 3321, 3319, and 3232 (all JEPS) form an almost continuous series, morphologically, from gracilis to capitata. Moreover, further investigation showed that even the odors of this com- plex graded into one another, i.e., all the intermediate forms did not smell exactly the same but tended toward one or another of the species heretofore recognized. It is my hope that these investigations will be of help to those who may wish to delve more deeply into the Madia sativa complex.—V. F. HessE, Boulder Creek, California.
CHROMOSOME COUNTS IN SECTION SIMIOLUS OF THE GENUS MIMULUS (SCROPHULARIACEAE). VII. NEW NUMBERS FOR M. GUTTATUS, M. CUPREUS,
AND M. TILINGII
R. K. Vickery, JR., K. W. Crook, D. W. Linpsay, M. M. M1, and W. Tar
The purpose of this report is to gather together and summarize the chromosome numbers found in M. guttatus DC. in the course of a series of experimental evolution studies (Vickery, 1966; Vickery and Lind- say, 1961; 1966; Vickery and Mia, 1966; and unpublished) and to pre- sent new numbers for M. cupreus Dombrain and M. tiling Regel. Our findings, particularly those concerning irregular chromosome segrega- tion in M. guttatus, significantly broaden the cytogenetic concept of each of these three species.
The cytological techniques employed were essentially the same as those previously used (Mia, Mukherjee, and Vickery, 1964) except that the first half of the slides were made permanent according to the method of Beeks (1955). His technique proved satisfactory, but was not superior to our usual method. Representative chromosome configurations were recorded either photographically or by drawings made with the aid of a camera lucida. Counts for a culture of a population or for a hybrid were based on a study of an average of over thirty cells representing an aver- age of three plants for each. Vouchers of each culture have been deposited in the University of Utah herbarium. Many of the counts here reported are included in the theses of the junior authors submitted to the Uni- versity of Utah in partial fulfillment of the requirements for the Master’s (K. W. Crook) and Ph.D. degrees. The work was supported in part by grants G 10603, GB 562, and GB 2996 from the National Science Foun- dation and in part by grants from the University of Utah Research Fund.
The normal n=14 chromosome number of Mimulus guttatus (Mia, Mukherjee, and Vickery, 1964) was confirmed for 20 additional popula- tions (table 1). Nine of these populations (table 2), all from northern Utah, exhibited irregular chromosome segregations in meiosis like the two cases previously reported for populations from the Sierra Nevada of California (Mukherjee and Vickery, 1959). Irregular segregations were detected also in culture 5839 (n=14) from the Spruces picnic area in the Big Cottonwood Canyon of northern Utah. The aberrant micro- sporocytes produced by the irregular segregations were functional, at least occasionally, and hence significant. One aneuploid parental plant and one F, hybrid lacking a chromosome were detected.
In all ten cultures the irregular segregations typically produced a few microsporocytes with either n=13 or n=15 chromosomes instead of the normal n=14 number. In two of the populations the irregularities were more pronounced. Culture 6649 of the Brighton population was
211
219 MADRONO [Vol. 19
TABLE 1. CHROMOSOME Counts IN MIMULWUS, SECTION SIMIOLUS
M. cupreus Dombrain. n=31: Chile, Nuble. Termas de Chillan, 7,000 ft, Moore 420, 63181. n=32: Cultivated under the names Bee Dazzler and M. beeslanus Bergm. in the Botanic Garden, Uppsala, Sweden, 5684.
M. guttatus DC.n=14: Utah. Cache Co. Logan River, 5,050 ft, 61762 and 8,500 ft, 6173. Juab Co. Thomas Creek, 4,775 ft, 6154; 7,200 ft, 6172; and 9,000 ft, 6170. Salt Lake Co. Brighton, 8,760 ft, 6649 and 9,250 ft, 7102; East Millcreek, 7,680 ft, 6648. Tooele Co. South Willow Creek, 5,800 ft, 6158 and 8,000 ft, 6178. Utah Co. Utah Lake, 4,500 ft, 6145; Provo River, 5,100 ft, 6164; 5,800 ft, 6167; 7,600 it, 6168; Payson Creek, 7,800 ft, 6658 and 7,800 ft, 6661. Calif. Mono Co. Mt. Dana, 8,000 ft, 6667. New Mexico. Catron Co. Aragon, 6,800 ft, 6611. Colo. Grand Co. Grand Lake, 9,560 ft, 7139. n=15: Utah. Cache Co. Providence, 4,500 ft, 6177. n= 16: Colo. Grand Co. Grand Lake, 9,500 ft, 7141. Mexico, Durango. Santiago Papasquiaro, 8,400 ft, Wiens 3,521, 6646. n=28: New Mexico. Catron Co. Reserve, 5,770 ft, 6613; Grant Co. Gila River, 4,400 ft, 6675; Rio Arriba Co. Rio Chama, 7,500 ft, 6622. Colo. La Plata Co. Los Pinos River, 6,750 ft, 6627. Alaska. Admiralty Island. Gambier Bay, sea level to 1,000 ft, Miller, Aug. 1958, 6152. Mexico, Du- rango. Santiago Papasquiaro, 8,400 ft, Wiens 3,522, 6647.
M. luteus L. n=31: Chile, Nuble. Las Cabras, 5,000 ft, Pena, Feb. 6, 1963, 6685.
M. tilingit Regel. n=28: Calif. El Dorado Co. US 89, 7,200 ft, 6664.
1 Author’s culture number. 2 Populations for which no collector is given were collected by Vickery and/or co-workers and any that were distributed included the indicated culture number.
observed to have one microsporocyte with n=12 chromosomes as well as one with n=15 and 30 with the normal n=14. Culture 6648 from the Big Water Gulch population carried the degree of irregularity further yet. One cell was observed to have n—10 chromosomes in addition to one with n=15 and 10 with n=14. The highest percentage of irregu- larity observed was 60%. This amount was shown by a population from the Utah Valley (table 2). However, the general average was much lower. Irregular chromosome segregation may be a far more widespread phenomenon in M. guttatus than we had supposed because it often occurs at such low frequencies (table 2) that we easily might have missed it previously.
In order to provide additional information to help explain the nature of the irregular chromosome segregations, each of a series of cultures of northern Utah populations including some that exhibited irregular seg- regations and some that did not were crossed to a culture (5052) of a normal California population and to a culture (6082) of a Utah popu- lation that displayed irregular segregation (table 2). Examination of the results suggests that an explanation based on simple gene control is unlikely. Observation of occasional univalents in the F, hybrids, indi- cates that lack of homology may be a valid explanation for some of the observed irregular segregations in the hybrids, particularly those from geographically isolated parents. A few, but widely distributed, observa- tions of chromatin bridges in anaphase coupled with pervasive regular bivalent pairing suggests to us the occurrence of different inversions in
1968 | VICKERY, ET AL.: MIMULUS 213
TABLE 2. SUMMARY OF CHROMOSOME COUNTS OBSERVED IN VARIOUS STAGES OF MICROSPOROGENESIS FROM THE PARENTAL CULTURES AND THE INTER-CULTURE F Hysrips oF NorTHERN UtaH MIMULUs GuTTATUS PoPuULATIONS. See Table 1 for
origins of these cultures or for references for those not listed in this report.
Parental populations: Fi hybrids with 5052: Fi hybrids with 6082:
n=13 n=14 n=15 Gir. n=13 n=14 n=15 Yir. n=13 n=14 n=15) Gir. Controls 5052 Oy. ) O10i- “actu ee ee ee ee res 6082 2 8 0,5 20.0 0 87 0 Oil < cae, ee ¢ ee Populations
5834 QO 10 ) 0.0 0 9 1 10.0 1 18 ? 14.3 5835 O13 0 0.0 O 35 0 0.0 I 27 1 6.9 5839 50-28 0 9.7 OPaess 0 0.0 1 34 0 2.8 6090 a Tale 0 143 1 18 0 55 O 12 0 0.0 6127 O 54 0 0.0 O25 1 3.8 0: 21 0) 0.0 6145 20a toe F112 1603 OF 11 0 0.0 (failed to flower) 6154 OF 33 0 0.0 3 «OF 4 6.7 O 10 0) 0.0 6158 2 34 1 8.1 O 24 1 4.0 1 34 4 12.8 6164 0 27 ) 0.0 0 6 0 0.0 132 0 on 6167 jee Bs) 0 6.2 O 10 1 9.1 Re 6) 2 2G 6168 Oma @) 0.0 (failed to flower) OF 25 0 0.0 6170 1 61 0 1.6 O 15 @) 0.0 ry 10 2) 23 6172 O 29 ¢) 0.0 2 60 0 3.2, 1 45 @) 22 6173 On e21 0 0.0 1 70 ¢) 1.4 O 24 @) 0.0 6175 1 25 0 3.8 3 34 1 10.5 0 81 0 0.0 6176 O 28 0) 0.0 ) 7 0) 0.0 0 46 0 0.0 6178 O 16 0 0.0 0 50 @) 0.0 Fos 0 7.4
the various populations. They would contribute to the explanation of the irregular segregations also. Further study is needed, because knowledge of the real causes of the observed irregular chromosome segregations is fundamental to our understanding of the evolutionary mechanisms op- erating within M. guttatus and particularly of the observed aneuploid changes in the species.
The first example of aneuploidy at the n=15 level in M. guttatus was found in a population (culture 6177) from the Cache Valley, Utah (table 1). The population is relatively uniform morphologically although its leaves are rounder and its calyces are blunter than is usual in M. guttatus. These traits suggest to us that this population may be show- ing the residual effects of previous introgression from M. glabratus var. utahensis Pennell. The latter species generally has n=15 chromosomes as well as the traits mentioned. Presently it occurs to the south and west of the Cache Valley throughout the Great Basin and central Mexico. but it may have spread into the Cache Valley some 7,500 to 4,000 years ago during the altithermal period (Antevs, 1955) when other more southernly distributed species such as Quercus turbinella Greene appar- ently migrated hundreds of miles northward in Utah (Cottam, Tucker. and Drobnick, 1955). Hybridization of M. guttatus with M. glabratus could have occurred at that time. Subsequently, with the return of cooler temperatures, the most northern M. glabratus populations such as the
214 MADRONO [Vol. 19
putative ones in the Cache Valley, would be expected to have died out. Natural selection, in turn, would be expected to have favored the best adapted members of the remaining hybrid swarms and in time to have produced the present populations.
The F, hybrids of this Cache Valley aneuploid population (culture 6177) exhibited irregular chromosome segregation even though no irreg- ularities were observed in the 28 cells studied of the parental culture itself. Its F, hybrids with culture 5052 revealed one microsporocyte with n—12 chromosomes, fourteen with n=13, twenty-one with n—14, twelve with n=15, and one with n=16. In contrast, its F, hybrids with culture 6082 exhibited two microsporocytes with n=13 chromosomes and fifty-five with n=14. Figured on the assumption that a regular segregation of chromosomes of the F, hybrids of this culture (n=15) with each of the controls (both are n=14) should have yielded n=14 and n=15 microsporocytes in equal numbers, the irregular segrega- tion observed occurred at a rate of approximately 50%. The results are a poor fit for this assumption. Perhaps a better hypothesis would be that the fifteenth chromosome of culture 6177 is a special type of B chromo- some that is regularly lost in the culture’s F, hybrids. Then the culture’s F, hybrids would be expected to have n=14 chromosomes which occurs in the cross 6082 X 6177 in over 96% of the cells examined. In the cross 5052 X 6177 there are many 15 to 13 irregular segregations. How- ever, this type of irregularity clearly suggests that n—14 is actually the number rather than n=15. The behavior of the chromosomes of the F, hybrids of aneuploid M. platycalyx Pennell (culture 5752, n=15) could be interpreted in the same way (Mukherjee and Vickery, 1962). However, until clearer proof of the accessory nature of culture 6177’s fifteenth chromosome is obtained it seems best to consider the popula- tion as a straight n=15 aneuploid, although noting the apparent B- chromosome-like loss of the fifteenth chromosome in the hybrids of the culture.
Aneuploidy at the n=16 level occurs in two distinctly different races of M. guttatus (fig. 1). The first race, as represented by culture 6646, consists of a small-flowered, small-leafed type of plant from near the crest of the Sierra Madre Occidental west of Durango, Mexico. Our prev- ious n—16 counts are for closely similar populations from the same general area (Mia, Mukherjee, and Vickery, 1964). The results of hy- bridization experiments now in progress may warrant segregating this race from M. guttatus as a new species. The other n=16 count is for an unusual race, represented by culture 7141, that reproduces by axillary propagules. It has been collected from several localities in the vicinity of Grand Lake, Colorado by Mrs. Margaret Douglas to whom we are indebted for the material studied.
Six different M. guttatus populations were found to be tetraploid (n=28). The five from northern Mexico, New Mexico, and southern Colorado appear to be autotetraploids. However, their allopolyploid origin
1968 ] VICKERY, ET AL.: MIMULUS 245
7102 7139 6646
J\A4l 6664 6622
micra
6627 6318
Fic. 1. Meiotic chromosomes of Mimulus. Cultures 7102 and 7139 are typical n=14, M. guttatus in anaphase II and anaphase I, respectively; cultures 6646 and 7141 are n=16 aneuploid races of M. guttatus in metaphase II and metaphase I, respectively; culture 6664 is tetraploid, n=28 M. tilingiz in metaphase I; cultures 6622 and 6627 are tetraploid, n=28 M. guttatus in diakinesis and metaphase I, respectively (note, the regular pairing in the former and the presence of 19,, and 18, associations in the latter), and culture 6318 is aneuploid, n=31 M. cupreus in diakinesis. The camera lucida drawings were at an original magnification of 2,850, reduced to approximately 950 in reproduction.
from different M. guttatus populations cannot be excluded, particularly in view of a high frequency of univalents in the Colorado population, culture 6627 (fig. 1). Morphologically these populations are indistin- guishable from the diploids of their area as in the previous, single ex-
216 MADRONO [Vol. 19
ample of tetraploidy (fig. 2) in M. guttatus, the population (6152) from the Verde Valley in central Arizona (Mia, Mukherjee, and Vickery, 1964).
The sixth n=28 population is from Admiralty Island towards the other end of the range of M. guttatus. This population is typical of a race that is widespread in coastal Alaska and the Aleutian Islands ac- cording to the herbarium specimens in the British Museum. A photo- graph of the De Candolle herbarium specimen of M. guttatus suggests to us that De Candolle’s type for the species may belong to this race. Plants of a culture (6250) of this population cross readily with plants of culture 5052 from Mt. Diablo, California, our standard test popula- tion. Observations of microsporogensis in the resulting vigorous, but sterile F, hybrids revealed 30 cells with 14 II and 14 I chromosome as- sociations. Fifteen cells exhibited slightly less pairing down to one with a low of 5 II and 32 I associations. In view of the numerous quadrival- ent chromosome associations observed in the culture itself (fig. 2), it is not clear whether autosyndesis of the chromosomes of culture 6250 was being detected or whether the pairing of the basic genome, thought to be present throughout all species of section Simiolus (Mukherjee and Vickery, 1962), was being observed. Plants of culture 6250 will not hy- bridize with those of the tetraploid population (culture 6152) from the Verde Valley which suggests that the former hypothesis is more likely. Clearly, if the Alaskan race is an autotetraploid it comes from a rather different race than the other autotetraploids. Possibly it is derived from the coastal ecotype. However, the short height and relatively large flowers of the plants may reflect an allopolyploid origin for this race from closely related forms (whose chromosomes pair) involving perhaps some combination of the inland race, the coastal ecotype, or alpine M. tilingu or one of its relatives.
As a result of these investigations of M. guttatus we find that aneu- ploidy is part of the cytogenetic picture of the species as is irregular chromosome segregation in meiosis. The latter phenomenon is now known to occur in 12 populations and probably is more widespread yet. Tetraploidy also is more common than we had previously thought. Seven tetraploid populations now have been found among the 76 popu- lations of M. guttatus analyzed to date (Vickery, 1955; Mukherjee, Wiens and Vickery, 1957; Mukherjee and Vickery, 1959 and 1960; Mia, Mukherjee, and Vickery, 1964; and table 1).
Mimulus cupreus Dombrain was found to have n=31 and n=32 pop- ulations (table 1). These results are in line with earlier crossing studies (unpublished) which showed M. cupreus to cross readily in most cases with J. luteus Ls, n=30, 31,°32,-and Wo -evinus hort, nn 3032 (Mukherjee and Vickery, 1959, 1960; Mia, Mukherjee, and Vickery, 1964; and table 1). The latter two species cross successfully with each other in some combinations, but not in others.
A single tetraploid population of M. tilingit from the Sierra Nevada
1968 ] VICKERY, ET AL.: MIMULUS 217
QO 10
6152 6250 nie
Fic. 2. Chromosome configurations of two tetraploid, n=28, races of M. guttatus. Culture 6152 is from the Verde Valley, Arizona population and culture 6250 is from the Admiralty Island, Alaska population. Both cells are in metaphase I. Note how frequently the bivalents are paired in loose quadrivalent associations. Drawn with the aid of a camera lucida at an original magnification of «1800 reduced to 900 in reproduction.
of California was found. This discovery is of interest in explaining the origin of M. tilingw var. corallinus with its n=24 chromosomes (Muk- herjee and Vickery, 1959). Perhaps the latter entity arose from a cross between diploid and tetraploid forms of M. tilingi or between diploid M. gutiatus (no tetraploids are known in the area) and tetraploid M. tilingu. The more tilingi-like than guttatus-like morphology of the variety suggests that the former hypothesis is more probable.
Our cytological findings suggests to us the possible presence in section Simiolus of parallel chromosome races in related species like the parallel series of morphologic races found by Vavilov (1923) in cereals and other plants. For example, M. tilingit has n=14, 15, 24, and 28 populations (table 1) and M. guttatus has n=14, 15, 16, and 28 populations (table 1). Another series occurs in three closely related species of the M. luteus complex. Mimulus tigrinus has n=30 and 32; M. luteus has n=30, 31, and 32; and M. cupreus has n=31 and 32 chromosomes. Perhaps fur- ther counts will complete these series of chromosome races and reveal the presence of other such parallel series in section Simiolus.
Department of Biological Sciences, University of Utah, Salt Lake City
LITERATURE CITED
Antevs, E. 1955. Geologic-climatic dating in the west. American Antiquity 20:317-335.
Beeks, R. M. 1955. Improvements in the squash technique for plant chromosomes. Aliso 3:131-134.
Cottam, W. P., J. M. Tucker, and R. Drosnick. 1959. Some clues to Great Basin postpluvial climates provided by oak distribution. Ecology 40:361-377.
M1, M. M.,B. B. MUKHERJEE, and R. K. Vickery, JR. 1964. Chromosome counts in the section Simiolus of the genus Mimulus (Scrophulariaceae). VI. New numbers in M. guttatus, M. tigrinus, and M. glabratus. Madrono 17:156- 160.
MukueryjeEE, B. B., and R. K. Vickery, Jr. 1959. Chromosome counts in the section Simiolus of the genus Mimulus (Scrophulariaceae). III. Madrono 15:57-62.
218 MADRONO [Vol. 19
. 1960. Chromosome counts in the section Simiolus of the genus Mimulus
(Scrophulariaceae). IV. Madrofio 15:239-245.
. 1962. Chromosome counts in the section Simiolus of the genus Mimulus
(Scrophulariaceae). V. Madrono 16:141-155.
.. D. Wiens, and R. K. Vickery, Jr. 1957. Chromosome counts in the sec- tion Simiolus of the genus Mimulus (Scrophulariaceae). II. Madrofio 14: 128-131.
VaviLov, N. I. 1951. The law of homologous series in the inheritance of variability. Chronica Botanica 13:55-94.
Vickery, R. K., JR. 1955. Chromosome counts in the section Simiolus of the genus Mimulus (Scrophulariaceae). Madronio 13:107-110.
. 1966. Speciation and isolation in Section Simiolus of the genus Mimulus.
Taxon 15:55-63.
.. and D. W. Linpsay. 1961. Crossing barriers in Mimulus guttatus in relation
to recent geologic changes in the Great Basin. Recent Advances in Botany,
pp. 903-906. Univ. Toronto Press, Toronto, Canada.
. and D. W. LinpsAy. 1966. Experimental hybridization in the genus Mimulus
(Scrophulariaceae). II. Proc. Utah Acad. 43:93-104.
,.and M. M. Mia. 1966. Experimental hybridization in the genus Mimulus
(Scrophulariaceae). III. Proc. Utah Acad. 43:105-114.
A RUST ON ANEMOPSIS CALIFORNICA
PAuL D. KEENER and JACK E. HAMPTON
The discovery of aecia on leaves of Anemopsis californica (Nutt.) H. & A., at three locations in Lincoln Co., Nevada (fig. 1) represents the first record of any stage of a rust fungus on plants of the Saurur- aceae in North America. Uredo houttuyniae Sawada has been reported on Houttuynia cordata Thunb., a saururaceous species, in Formosa (Sawada, 1944). Uromyces saururi P. Henn. is known to occur on Sau- rurus chinensis Hort., in China (Cummins and Ling, 1950) and on S. loureirt Decne, in Japan (Hiratsuka, 1955).
Collections of infected Anemopsis (Yerba-mansa) were made at Hiko, Crystal Springs, and Ash Springs on September 20, 1965. The average numbers of aecial sori per leaf from 10 randomly selected samples from each location were: Hiko, 0.3; Crystal Springs, 0.5; and Ash Springs, 1.34. At the two latter locations, infected plants appeared as if burned. Many of the sori and aeciospores were partially or completely destroyed by Tuberculina persicina (Ditm.) Sacc., a hyperparasite. Aeciospores were obtained for morphological comparisons; germination was poor, and inoculations on associated hosts, chiefly Gramineae, were unsuccess- ful.
Intermixed with rusted Anemopsis were plants of saltgrass, Distichlis stricta (Torr.) Rydb., bearing uredia and telia. This rust was most prev- alent on saltgrass leaves adjacent to or in contact with heavily rusted Yerba-mansa. The rust on Distichlis was identified tentatively as Puc-
1968 | KEENER & HAMPTON: RUST ON ANEMOPSIS 219
Fic. 1. Aecial sori of a rust on leaves of Anemopsis californica: A, Numerous grouped sori some of which are altered (at left) by presence of Tuberculina persi- cina, a fungus attacking rusts; B, Enlarged views of aecial sori. Lower right, nor- mal appearing cylindrical shaped structures. Upper left, a group of sori bearing dark-colored hyphae and spores of Tuberculina persicina.
TABLE 1. COMPARISON OF AECIAL CHARACTERS OF A RUST ON ANEMOPSIS CALIFORNICA FROM 3 LOCATIONS IN NEVADA WITH THOSE OF PUCCINIA ARISTIDAE.
Aeciospores Wall Peridial Position of Size Thickness Ornamen- cells Rust aecial sori (Microns) (Microns) tation (Microns)
From Finely 16-24 X 21-2& Hiko Hypophyllous 14-19 « 17-23 1-3 Verrucose Verrucose From Crystal Hypophyllous 14-19 X 17-24 1-3 Finely 12-23 & 22-31 Springs Verrucose Verrucose From Generally Ash Hypophyllous but 16-20 X 17-23 1-3 _ Finely 16-24 & 21-26 Springs some amphigenous Verrucose Verrucose Puccinia Chiefly Finely aristidae hypophyllous 13-21 * 15-23 1-3 Verrucose not given
cinia aristidae Tracy, although Cummins and Husain (1966) consider P. subnitens Diet. on Distichlis to be distinct from P. aristidae on Aris- tida. Comparisons of aecia-aeciospore morphology from the 3 Nevada localities with date reported for P. aristidae (Arthur, 1962) are given in Table 1.
The spermagonia and aecia of Puccinia aristidae have been reported as occurring both naturally and experimentally on numerous species of both Monocotyledoneae and Dicotyledoneae. Arthur (1962) lists 104
220 MADRONO [Vol. 19
species in 66 genera of 22 families as being susceptible to the rust, (but see Cummins & Husain). In Nevada, 12 species in 11 genera of 7 fam- ilies are listed as suscepts.
This paper has been approved as Journal Paper No. 1191, Arizona Agricultural Experiment Station. The late Dr. Keener was Professor of Plant Pathology at the University of Arizona.
University of Arizona, Tucson Nevada Department of Agriculture, Reno
LITERATURE CITED
ARTHUR, J. C. 1962. Manual of the rusts in United States and Canada. With Sup- plement by G. B. Cummins.
Cummins, G. B., and S. M. Husain. 1966. The rust fungi on the genus Aristida. Bull. Torrey Club 93:56-67.
,and Lee Ling. 1950. An index of the plant rusts recorded for Continental
China and Manchuria. Plant Dis. Reptr. Suppl. 196:520-556. Nov. 16.
HiraTsuKA, N. 1955. Uredinological studies. Kasai Publ. Co., Tokyo.
SawabA, K. 1944. Descriptive catalogue of Formosan fungi, Pt. X. Taiwan Agr. Inst. Rept. 87:45.
REVIEWS
Polyporaceae of North America. The Genus Poria. By Jos1AH L. Lowe. 183 pp., 159 line drawings. Technical Bulletin No. 90. State University College of For- estry at Syracuse University, New York. 1966.
The genus Porza is generally considered one of the most difficult of the genera in the Polyporales, and a modern, thorough treatment such as this constitutes a most welcome addition to the literature. Taxonomic keys and descriptions of approxi- mately 130 species are presented plus descriptions of several species belonging to related genera that might be confused with those of Poria. By use of color of the pore surface and type of hyphal system the genus is divided into five sections. Because this arrangement is apparently still somewhat tentative the sections were left unnamed. The species dscriptieons contain sufficient details of both macro- scopic and microscopic characters to enable one to feel relatively confident of his determinations. No new species or combinations are presented. A short introduc- tion and discussion of the morphology, distribution and economic importance of the genus precedes the taxonomic treatment.
Attempts at using the keys were usually successful, but it seems however, that some improvement in their construction could have been made. For example, opposing choices could have been kept adjacent to one another thus eliminating the necessity of searching through several pages for the second choice. The use of such terms in the choices as “frequently,” “usually,” etc. tends to destroy one’s confidence in the keys and thus increase the difficulty of determinations. In this same category, but somewhat more displeasing, is the use of “large” or “small” in referring to size of various structures or cells without giving more precise meas- urements. One final observation is that perhaps if a few photographs had been included, the use of the manual would have been enhanced, at least for use in advanced classes. Typographical errors are few and of minor significance. None of these criticisms are intended to seriously detract from the real value of this publi- cation, and it will be of considerable value to all working with these fungi— Harry D. Tuiers, Department of Biology, San Francisco State College.
1968] REVIEWS 221
The Komarov Botanical Institute, 250 Years of Russian Research. By STANWYN G. SHETLER. xiv + 240 pp., illus. Smithsonian Institution Press, Washington, D.C. 1967. $5.95.
In recent decades biologists have largely relegated the art of biological historiog- raphy to historians. The result is that accounts of the development of biological ideas and institutions show insights into the cultural and political milieu, but often lack perspective when considering the biological Zeitgeist of the period or events covered. Therefore it is refreshing that a botanist has written a balanced and inter- esting account of the evolution of one of the world’s largest and most prominent botanical institutions—the Komarov Botanical Institute in Leningrad. Most Ameri- can systematists are aware that such an institute exists, know that it probably has the largest herbarium in the world, but would be hard-pressed to continue discus- sion beyond this point. Stanwyn Shetler’s comprehensive treatment of the 250-year history of the institute will alleviate our collective gaps in knowledge of this impor- tant center of taxonomic and floristic research.
The first part of the book deals with the establishment of St. Petersburg and its transition into modern Leningrad, a city which provides frustration as well as de- light to a tourist. The second part of the book concerns the development of the Imperial Botanic Garden in St. Petersburg and the Botanical Museum of the Acad- emy of Sciences. The botanic garden eventually came under the aegis of the Acad- emy of Sciences and in 1931 it was decided that the two organizations should be united as the Botanical Institute of the Academy of Sciences of the U.S.S.R. In 1940, the Botanical Institute was named after V. L. Komarov, a distinguished botanist who organized the monumental Flora SSSR project which was completed in 1964. At present, the institute has a “staff of 700, two dozen major laboratories, a large greenhouse and outdoor garden complex, an arboretum-park, several ex- perimental farms, a 450,000-volume library, and combined herbaria of nearly 6 million specimens”! Administratively, the Institute consists of departments of Vas- cular Plants (headed by Armen Takhtajan); Cryptogamic Plants (M. M. Holler- bach) ; Geobotany (B. A. Tikhomirov—whom some readers may know personally since he was a relatively communicative attendee at two recent International Bo- tanical congresses) ; Plant Resources (Economic Plants [Al. A. Federov]) ; Evolu- tionary Morphology (M.S. Yakovlev) ; Botanic Garden (N. A. Avrorin) ; Botanical Museum (F. Kh. Bakhteev) ; and two physiological-biochemical departments: Pho- tosynthesis and Microelements. Thus, the massive staff and facilities of the Institute are largely devoted to systematic studies; such concentrated resources are unparal- leled in the western world. The third and final part of the book is a summary of previous chapters and also discusses the future of the Institute. The latest Five-Year Plan for the Institute calls for continuation of present efforts in floristics, system- atics, and geobotany (which in current Russian usage “refers to the general study of vegetation—its origin, development, distribution, and utilization (especially) .” With the revival of Mendelian genetics in the Soviet Union it seems likely that bio- systematic studies will resume as well. In the international field, the Institute antici- pates a strong role in the International Biological Program, and Shetler suggests that the Russians may initiate intensive studies in the tropics. He adds that “be- cause much exploratory research on the tropics must necessarily proceed along classical lines, Soviet botany may actually be in a better posture today for tropical studies than American botany, owing to the stronger persistence of classical method- ology and greater sympathy for herbaria in the USSR.”
Shetler’s book is lucid and straightforward; it is well illustrated with scenes of Leningrad, of the Institute, and most valuably, of the principal present and past staff members of the Institute. Its parallel usage of Cyrillic and Roman characters for Russian words and names will enhance the utility of the book for librarians and bibliographers. I cannot predict how this study will be received by the Russians, but it is a dispassionate and comprehensive presentation which is most welcome to us in the West.— ROBERT ORNDUFF, University of California, Berkeley.
22) MADRONO [Vol. 19
A Field Guide to the Natural History of the San Francisco Peninsula. By JoHN C. Witiiams and Howarp C. Monroe. xvi + 285 pp., illustrated with numerous line drawings and photographs. McCutchan Publishing Corp., 2526 Grove St., Berkeley, Calif. 94704. Spiral bound, $5.50; hardback, $7.50.
In the preface to this book the authors state that: “The idea behind this book is to foster an understanding of the natural history of our surroundings so that we may better understand the potentials and cope with the problems of our environ- ment.” This is certainly a good idea and we need many more interpretive manuals. The book is divided into nine parts: introduction, bay and salt marsh, the open coast, the chaparral, the grasslands, the broadleaf forests, the cone forest, appendix, and index. The grouping of organisms by community (parts 2-7) is a good way of arranging the material.
Unfortunately, though, this guide falls far short of its goals because of the nu- merous factual errors. A few examples from the botanical contents of the guide, alone, will indicate why this is so.
One page 27, Typha latifolia is in the checklist of coastal and bay salt marsh plants and animals.
Pinus radiata (p. 48) does not “grow wild along much of the coast.”
“Weeds of the genera Brassica ssp., Camelina spp., Capsella spp., are found all through the coastal strand.” This statement from page 51 is true for Brassica and Capsella, but not for Camelina.
The picture labelled Morning Glory (p. 52) is actually one of Senecio mikanioides not of Convolvulus soldanella and that of Spearmint (p. 109) is Stachys, not Men- tha spicata.
One page 109 we find: “Spanish Bayonet (Yucca spp.) is common in the south- ern or high chaparral.” Yucca whipplei is known from Monterey County, but that county is not part of the San Francisco Peninsula by any stretch of the imagination. And on page 123 the Spanish Bayonet is called Yucca schidigera!
On page 131 we learn that Black Acacia is Acacia melanoxia, on p. 137 that Soft Cheat is Bromus hordeaceus, on page 134 that California Needle Grass is Stipa pulchia, on page 135 that Indian Thistle is Cirstum edule, and on page 141 that the California Golden Poppy is Eschscholtz californica. Errors of this kind are found on nearly every page.
On page 173, we find the myth perpetuated that “The California Laurel or Bay (Umbellularia californica) is the ‘Myrtle’ of the Bible.”
Vitis californica (p. 175) does not grow natively on the San Francisco Peninsula and certainly is not “common.”
The drawings on page 266 which are supposed to shew differences between su- perior and inferior ovaries indicate that the authors do not understand the arrange- ment of parts in the flower.
A thorough revision of this book is necessary before it can be used.—JouN H. Tuomas, Dudley Herbarium, Stanford University.
NOTES AND NEWS
NEW PUBLICATIONS
Checklist of Vascular Plants of the Intermountain Region By ARTHUR H. HOLMGREN and JAMEs L. REVEAL. iv + 160 pp. U.S. Forest Service Research Paper INT-32. 1966. Intermountain Forest and Range Experiment Station, Ogden, Utah.
Interspecific Relationships in the Genus Monarda (Labiatae). By RAINER W. Scora. University of California Publications in Botany 41:1—71. University of California Press, Berkeley and Los Angeles. 1967. $2.50.
Memorials of John Bartram and Humphry Marshall. By Wi1LL1AM DartLinctTon. In- troduction by JosEPH Ewav. lii + 585 pp. (Facsimile of the edition of 1849). Hafner Publishing Co., 31 E. 10th St., New York. 1967. $20.00
1968 ] NOTES AND NEWS 223
Bibliography of Utah Botany and Wildlife Conservation. By EArt M. CHRISTENSEN. Brigham Young University Science Bulletin, Biological Series 9(1):1—136. Provo, Utah. 1967.
Guide to the Woody Plants of Utah. By Mi1cHAEL TresHow, STANLEY L. WELSH, and GLEN Moore. 161 pp. illus. Pruett Press, Inc., Boulder, Colorado. 1964. $3.00 Mountain Plants of Northeastern Utah. By Brerniece A. ANDERSON and ARTHUR H. Hoimcren. 148 pp., illus. Circular 319, Utah State University Extension Services,
Logan. 1966. $1.25.
Trees and Flowering Shrubs of Yellowstone and Grand Teton National Parks. By RICHARD J. SHAw. 50 pp., illus. in color. Wheelright Press, 975 SW Temple, Salt Lake City, Utah. 1964. $1.00.
World Vegetation. By Denis RILEY and ANTHONY YOUNG. 96 pp. Cambridge Uni- versity Press. 1966.
Native Trees of Canada. xviii + 291 pp., illus. Bull. 61, Department of Forestry. Queen’s Printer, Ottawa. 6th edition, 1966 reprint. $3.00.
Wild Plants of the Canadian Prairies. By ARCHIBALD C. Bupp and KeirH F. Best. viii + 519 pp., illus. Publication 983, Research Branch, Canada Department of Agriculture. Queen’s Printer, Ottawa. 1964. $3.00, paper; $4.00, cloth.
VALIDITY OF NOMENCLATURE CHANGES UNDERTAKEN IN THE FLorA OF ALASKA AND Yuxon.—In Arkiv for Botanik, Ser. 2., 7(1):1-147. 1967, edited by the Swedish Academy of Sciences, a numberof nomenclatorial changes in the flora of Alaska and Yukon were made. Unfortunately some of them were not validly published in ac- cordance with the Code of International Botanical Nomenclature. They are valided below:
Page 23. Eriophorum callitrix var. pallidus Hultén, var. nov. Type in S. Accord- ing to Marcel Raymond, Montreal, it is the same as var. moravium Raymond (Bull. Soc. Bot. Fr. 98:7. 1951).
Page 35. Platanthera saccata (Greene) Hultén, comb. nov. Add: Habenaria sac- cata Greene (Erythea 3:49. 1895).
Page 52. Minuartia yukonensis Hultén, sp. nov. Type is S.
Page 59. Ranunculus gelidus ssp. grayi var. shumaginensis Hultén, var. nov. Type in S.
Page 60. Papaver hulténii var. salmonicolor Hultén, var. nov. Type in S.
Page 62. Cardamine bellidifolia var. pinatifida Hultén, var. nov. Type in S. Car- damine purpurea var. albiflos Hultén, var. nov. Type: Alaska, McKinley Park High- way, mile 63.5, Throughfare Pass, July 1964, Hultén, S.
Page 64. Draba stenopetala var. purpurea Hulten, var. nov. Type in S.
Page 66. Braya bartlettiana var. vestita Hultén, var. nov. Type in S.
Page 67. Parrya nudicaulis ssp. septentrionalis Hultén, ssp. nov. Type in S.
Page 68. Saxifraga foliolosa var. multiflora Hultén, var. nov. Type in S.
Page 69. Saxifraga nivalis var. rufopilosa Hultén, var. nov. Type in S.
Page 71. Rubus arcticus ssp. arcticus var. pentaphylloides Hultén, var. nov. Type in S.
Page 101. Diapensia lapponica var. rosea Hultén, var. nov. Type in S.
Page 113. Polemonium boreale var. villosissimum Hultén, var. nov. Type in S.
Page 122. Pedicularis labradorica var. sulphurea Hultén, var. nov. Type in S. Pedicularis langsdorffii ssp. arctica (R. Br.) Pennell should be cited as P. langs- dorffii ssp. arctica (R. Br.) Pennell ex Hultén.
Page 134. Antennaria friesiana ssp. alaskana var. beringensis Hultén, var. nov. Type in S.
Page 135. Antennaria monocephala ssp. philonipha var. latisquamea Hultén, var. nov. Type in S.
Page 139. Artemisia glomerata var. subglabra Hultén, var. nov. Type: Alaska, Kagati Lake, Sept. 12, 1965, Hultén, S—-Eric Hutten, Naturhistoriska Riksmuseet,
Stockholm, Sweden.
224 MADRONO [Vol. 19
THREE ROADSIDE GRASSES AS RECORDS FOR SACRAMENTO, EL DoraDo, AND PLACER CounrTIES, CALIFORNIA.—The following grasses occur on gravelly to sandy road- shoulders or on roadbanks of similar texture. Seldom, if at all, do they penetrate be- yond into areas where competition is greater and summer moisture is wanting. Two major highways, Interstate Highway 80 (U.S. Highway 40) and U.S. Highway 50 in the area east of Sacramento are the subject of this study. Collections were made during July and August, 1967 unless otherwise noted. The two perennial species at all locations are in the second, third, or more year’s growth. All collections cited are by the author and are deposited in AHUC.
Panicum hillmani Chase. This grass readily spreads by the large panicles which break away from the stem and are carried by the wind as “tumbleweeds.” The plant extends through Sacramento County along both highways to the Placer and El Dorado county lines (8135, 8136, 8143, 8169, 8173). Currently, the best stand occurs along Interstate Highway 80 between Arden Way and Marconi Avenue in Sacramento.
Sporobolus cryptandrus (Torr.) A. Gray. The commonest grass of the three occur- ring along both highways into the Sierra Nevada to about 3000 feet elevation (8142, 8144, 8145, 8146, 8148, 8156, 8157, 8158, 8163, 8165, 8166, 8171). The grass was first noted in Sacramento County near the Nimbus Road exit, U.S. Highway 50, June 3, 1965 (7363).
Chloris verticillata Nutt. The unique inflorescence of this grass readily disartic- ulates as a unit from the stem below the radiating spikes and behaves as a tumble- weed facilitating its spread along roadshoulders. It occurs along Interstate Highway 80 from east of Sacramento about the 1500 ft. elevation in the Sierra Nevada (8164, 8170, 8172, 8177, 8178). A single collection was made along U.S. Highway 50, 2.5 miles west of Shingle Springs in El] Dorado County (8147). At nearly all locations the grass occurs in association with Sporobolus cryptandrus and Eremocarpus setigerus Benth. At one location between Sacramento and Roseville the grass ap- pears to be in at least the third season of growth—BErEECHER CRAMPTON, Depart- ment of Agronomy, University of California, Davis.
A New Famity oF VASCULAR PLANTS (PSILOTACEAE) FOR ARIZONA.—Sycamore Canyon located about twenty miles west of Nogales in Santa Cruz County, Arizona, has long been noted for its interesting and unusual flora (Goodding, Jour. N. Y. Bot. Gard. 47:86-97. 1946; Jour. Ariz. Acad. 1:113-115. 1961). It is the type lo- cality for Fraxinus gooddingi Little which is found outside the canyon only in the adjacent Pena Blanca drainage area. Several Mexican species Dalea lagopus (Cav.) Willd., Lotus alamosanus (Rose) Gentry, and Passiflora bryonioides H.B.K., are found in the United States only in Sycamore Canyon; likewise, Asplentum exiguum Bedd, known from Asia, and Northern Mexico, occurs in Sycamore and Garden Canyons in Arizona. Several other species, including Dichondra sericea Sw., Aloysia lycioides Cham., Tilandsia recurvata L., and Aeschynomene villosa Poir. having irregular, disjunct distributions, grow in this rugged and picturesque area.
A recent find in Sycamore Canyon adds a new family and order to the known flora of Arizona. Psilotum nudum (L.) Griseb. was collected by Mr. Jack Kaiser of the U.S. Department of Agriculture, from a rock crevasse on a protected west ex- posure. This locality is about 300 miles north of the closest known population in Sonora, Mexico, as reported by Gentry (Carnegie Inst. Pub. 527:328. 1942), and is many miles north and west of the areas near Chihuahua, in the State of Chihuahua, Mexico (Knoblock and Correll, Conr. Texas Res. Found. 3:198. 1962). The closest recorded locality for Pszlotum in the United States is Hardin County in eastern Texas (Knoblock, Wrightia 2:163-165. 1961).
The specimen is on file in the University of Arizona Herbarium. This paper is Arizona Agricultural Experiment Station Journal Article number 1329.—CuHar tes T. Mason, Jr., Herbarium, The University of Arizona, Tucson.
wei
| 4 VOLUME 19, NUMBER 7 JULY, 1968
Contents
THE VEGETATIONAL COMPOSITION OF THE SOUTH
SLOPE oF MrT. Prnos, CALIFORNIA
Richard J. Vogl and Brian C. Miller 225 TAXONOMY, VARIATION, AND CHOROLOGY OF THREE
CHROMOSOME RACES OF THE CALAMAGROSTIS CANADENSIS
CoMPLEX IN ALASKA, Wm. W. Mitchell 235 New ReEcorDs oF MyXOMYCETES FROM CALIFORNIA. IIT, Donald T. Kowalski and Dwayne H. Curtis 246
CHROMOSOME COUNTS IN SECTION SIMIOLUS OF THE GENUS
MIMULUS (SCROPHULARIACEAE). VIII. CHROMOSOMAL
HoMOLOGIES OF M. GLABRATUS AND Its ALLIED SPECIES AND
VarreTies, M. M. Mia and Robert K. Vickery, Jr. 250 IDENTITY OF THE OAK TREE AT LIVE OAK Tank,
JOSHUA TREE NATIONAL MONUMENT, CALIFORNIA,
John M. Tucker 256 SEED EcoLocy OF SEQUOIADENDRON GIGANTEUM, NV. Stark 267 A CyTOTAXONOMIC STUDY OF A NATURAL HYBRID BETWEEN
AGROPYRON CRISTATUM AND A. SUBSECUNDUM,
W.S. Boyle and A. H. Holmgren Dae THE WATER HYACINTH IN CALrForNIA, Jane H. Bock 281 NoTEs ON ORYZOPSIS HENDERSONII (GRAMINEAE),
Richard Spellenberg 283
Reviews: P. H. Davis, Flora of Turkey and the East Aegean Islands, Vol. II (Peter H. Raven) ; W. Dallimore and A. B. Jackson, A Handbook of the Coniferae and
A WEST AMERICAN JOURNAL OF BOTANY
Ginkgoaceae, Fourth Edition (Ira L. Wiggins) 287 | NoTEs AND News: THE OCCURRENCE OF HEMITOMES CONGESTUM IN THE SOUTHERN SIERRA NEVADA, Barbara Rice 234 CONFUSION IN AUTHORSHIP OF STEMODIA VERTICILLATA
Hasster, Ira L. Wiggins 266
JepPson’s FLora OF CALIFORNIA, Robert Ornduff 288
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. Subscription price $6.00 per year ($4.00 for students). 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 un- delivered copies should be sent to the Corresponding Secretary, California Botanical Society, Department of Botany, University of California, Berkeley, California 94720.
BOARD OF EDITORS
LYMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. ERnsT, Smithsonian Institution, Washington, D.C. ARTURO GOMEZ PompPa, Universidad Nacional Autonoma de México EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia Mitprep E. Matuias, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley MARION OwNnBEY, Washington State University, Pullman REED C. Rottiins, Harvard University, Cambridge, Massachusetts Ira L. WiccIns, Stanford University, Stanford, California
Editor—JoHNn H. THomaAs Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—J UNE McCAsKILL P.O. Box 23, Davis, California 95616
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Elizabeth McClintock, California Academy of Sciences, San Fran- cisco. First Vice-President: Peter H. Raven, Department of Biological Sciences, Stanford University. Second Vice-President: Cornelius H. Muller, Department of Biological Sciences, University of California, Santa Barbara. Recording Secretary: John West, Department of Botany, University of California, Berkeley. Correspond- ing Secretary: Mary E. Ashton, Department of Botany, University of California, Berkeley. Treasurer: June McCaskill, Department of Botany, University of Cali- fornia, Davis.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Carl W. Sharsmith, Department of Bio- logical Sciences, San Jose State College, San Jose; the Editor or Madrofio; and three elected Council Members: Virginia M. Page, Department of Biological Sci- ences, Stanford University; Annetta Carter, Department of Botany, University of California, Berkeley; and Robert Ornduff, Department of Botany, University of California, Berkeley.
THE VEGETATIONAL COMPOSITION OF THE SOUTH SLOPE OF MT. PINOS, CALIFORNIA
RIcHARD J. VoGL and BRIAN C. MILLER
An ecological study of the vegetational composition of the coniferous forest on the south half of Mt. Pinos was conducted from 1962 to 1966. The mountain is located on the Kern-Ventura County line at the ex- treme south end of the San Joaquin Valley. The south face rises sharply from Lockwood Valley (5500 ft. elevation) up a steep grade, interrupted by minor canyons, valleys, and ridges, to the summit at 8831 ft. eleva- tion. The slope drains by way of Lockwood and Piru Creeks and the Santa Clara River into the ocean, about 40 miles away. The area was selected because of its accessibility and interesting flora, and because the forest was relatively undisturbed.
Geologically Mt. Pinos is bounded on the north by the San Andreas Fault and on the south by the Big Pines Fault. Formation of the granite core of the mountain is believed to have occurred sometime during the Jurassic Period (Jenkins and Kundert, 1955). In most areas the granite bedrock has decomposed to form soils, but on actively eroding sites and rises, the bedrock is still evident. The soils are composed of a mixture of coarse-grained and extremely fine particles of granitic material, are slightly acid, and are covered by pine duff up to 4 inches in depth.
The climate of the mountain is relatively dry. Precipitation data was unavailable for the mountain, but an average rainfall of 12.5 inches per year based on a 22 year average (1941-1963) was recorded at Chuchu- pate Ranger Station in Lockwood Valley. Major precipitation falls there from December through March, with snow predominant in February and March. Precipitation on the mountain is undoubtedly higher than that of the valley. These higher amounts are perhaps illustrated by the seven-year mountain summit snow depth records kept by the Forest Service. Snow depths averaged 32.4 inches on March 1 and 18.2 inches on April 1. It is generally accepted that 10 inches of snow roughly cor- responds to 1 inch of rain (Oosting, 1958). Rainfall during the remain- der of the year probably averages about one-half inch per month, al- though the summer months are commonly without precipitation except for small local thundershowers. A rain shadow produced by the coast ranges affects the mountain as it does the San Joaquin Valley. For ex- ample, Ojai, 12 miles from the coast at 750 ft. elevation, has an average annual rainfall of 21.6 inches, whereas Maricopa, on the San Joaquin Valley edge about 45 miles inland and 854 ft. above sea level, receives an average of only 5.7 inches of rain (U.S. Dept. Agr., 1941). The cli- mate is typically Mediterranean, but at the same time somewhat conti- nental and more characteristic of arid inland mountains than of adjacent coast ranges with maritime influence.
Maprono, Vol. 19, No. 7, pp. 225-288. December 12, 1968. 225
226 MADRONO [Vol. 19
METHODS
A species list was compiled. Eight sample sites or stands ranging from the base to the summit were selected. Although the entire mountain had been grazed by cattle, each stand was free from logging and road build- ing disturbances and was at least an 8 acre uniform representation of a vegetational association.
In each stand 80 trees and 80 saplings were sampled with the point- centered quarter method (Cottam and Curtis, 1956). Individuals were classed according to forestry standard sizes with those over 4 inches in diameter grouped as trees, those 1—4 inches as saplings, and those less than 1 inch as seedlings. Twenty-five foot line-intercepts were placed systematically at every other point to sample the understory shrub and herb cover. Sapling and tree increment borings were taken to obtain age and growth rates. Stump ring counts were made on trees removed in a partial logging-cut completed near the termination of the study. Counts of lightning-damaged trees were made on the upper one-third of the slope. An Importance Value (I.V.) was obtained for each tree species by summing its relative frequency, density, and dominance. Since dom- inance (basal area) of saplings was not taken, the relative frequency and density of each sapling species were totaled and multiplied by a factor of 1.5 to obtain values comparable to tree I.V.’s.
Nomenclature of species follows Munz (1959). Voucher specimens are in the herbarium at California State College at Los Angeles. A problem of species identification existed between Pinus ponderosa and P. jeffreyi at lower elevations where trees were found with intermediate character- istics. Because of this variability, trees were classified using the criteria suggested by Haller (1962). Based upon these criteria, the study trees were considered to be P. jeffreyi.
RESULTS AND DISCUSSION
Pinus monophylla was the most common tree at lower elevations (5500-5700 ft.). Table 1 lists the average sizes, numbers per acre, and I.V.’s of the trees and saplings. Pinus jeffreyi began to appear in stand two at 5700 ft. elevation, along with a few tree-sized Quercus turbinella ssp. californica. From about 6000 ft. to the summit, P. jeffreyi was the most important tree. In stand four at 6900 ft., P. jeffreyvt appeared savanna-like with 22 trees per acre scattered across a light grassy under- story. P. jeffreyt was associated with Quercus kelloggi along canyon bottoms and on southeastern exposures. Occasional Quercus chrysole pis were also found at mid-elevations. Starting about 7750 ft. (stand six), P. jeffreyi was associated with an occasional A dies concolor, particularly on slopes with northerly exposures. The P. jeffreyi forest on the upper slopes (8750 ft. elevation) consisted of large (48 inch dbh) and medium- sized pines with densities of about 105 trees per acre (fig. 1).
The sapling layer was related and similar to the tree layer in compo-
1968] VOGL & MILLER: MT. PINOS Did
TABLE 1. S1zE, DENSITY, AND IMPORTANCE VALUES OF TREES AND SAPLINGS.
Avg. Basal Stand Species Area per Trees-Saplings Trees-Saplings No. tree (in.*) per acre LVe
1 Pinus monophylla 64 85 61 288 152 Quercus turbinella 18 Z 61 12 148
2 P. jeffreyi 367 11 1 138 31 P. monophylla 13 20 10 132 175
Q. turbinella 19 4 5 30 94
3 JAlbzes concolor ieee OZR ee ah P. jeffreyi 391 27 10 191 237 P.monophylla 32 Z i 2, ~* «seer O.chrysolepis (§ ———aaasss ates OQ,” ee 7
QO. kelloggi 153 18 1 97 48
4 P. jeffreyi 667 2 4 287 266 P. monophylla 12 1 0.1 7 16
QO. chrysolepis Halak 0.3 0.2 6 18 CE IAIANE 309 63 ig 287 250 P.monophylla 32 2 0.2 7 8
Q. chrysolepis 12 1 1 6 42
6 <A.concolor 855 3 1 44 30 P. jeffreyi 297 33 13 256 270
7 A.concolor 349 if 1 12 17 P. jeffreyi 534 50 26 288 283
8 A.concolor ee hae OU = ee 9 P. jeffreyi 697 105 | 300 291
sition, but contained much lower densities. In stands one and two, tree reproduction consisted of about equal amounts of P. monophylla and Quercus turbinella (table 1). A few sapling-sized P. jeffrevi were also present in stand two. In stand three (6500 ft. elevation), P. jeffreyvi and Q. kelloggii dominated the sapling as well as the tree layer. A few Abies and Q. chrysole pis saplings were also found. In stand four tree reproduc- tion was represented by a limited number (four per acre) of P. jeffrey with lesser numbers of Q. chrysolepis and P. monophylla. There were five times more trees than saplings in stand five at 7400 ft. elevation. The sapling layer in the upper elevational stands was dominated by low densities of P. jeffreyi. Abies saplings were most common on northern exposures. Tree reproduction was minimal throughout the mountain- side. Pinus jeffreyi was the most important species, except for P. mono- phylla at the lowest elevations and the scattered Abies saplings at the highest elevations.
Shrub cover was sparse over most of the area. Common lower eleva- tional shrubs. were Quercus turbinella, Artemisia tridentata and Yucca whipplei (table 2). Symphoricarpos parishit was most common at mid- elevations (1.2% cover). Some of the steeper slopes at these elevations were treeless and occupied entirely by an impenetrable cover of Ceano-
228 MADRONO [Vol. 19
Fic. 1. Pinus jeffreyi forest at 8800 ft. elevation. The large trees, sparce number of saplings, and almost complete lack of herbaceous layer give the forest a park-like appearance. The large tree on the right has been damaged by lightning.
thus cordulatus which grew to an average height of 3 ft. Other local areas were occupied by Chrysothamnus nauseosus ssp. mohavensis. The upper forest was dominated by Symphoricarpos which reached its great- est abundance of 23% cover in stand seven (table 2). Ribes cereum was locally important in openings and around rocky outcrops at upper elevations.
Herbaceous vegetation was sparse and discontinuous (table 2). In stand one, 54.3% of the sample area was devoid of plant cover (table 2). Sitanion jubatum was most common in stands one and two and was the most widespread herb since it was found in all stands. Bromus tectorum dominated the forest floor in stand two with 28.8% cover which was slightly more than all of the 20 other herbaceous species combined. Cover in stand three was dominated by Siztanion, B. tectorum, Poa nervosa, and Hesperochloa kingu giving it a somewhat grassy appear- ance. Small forbs mixed with the grasses included Gayvophytum diffusum, Eriogonum latifolium ssp. pauciflorum, Penstemon labrosus, Erysimum capitatum, and Galium aparine. In stands four through eight (6900- 8750 ft. elevation) the sparse herbaceous layer consisted of infrequent grasses and forbs such as Gayophytum, Eriastrum sapphirinum ssp. ambiguum, Collinsia torreyi var. wrightii, Gilia modocensis type tetra- breccia, Penstemon labrosus, Lupinus confertus, and L. elatus, which provided less than 1% cover per species. Stand eight, near the mountain summit, contained the greatest herb variety, probably because of the proximity of forest openings and the summit meadow.
1968] VOGL & MILLER: MT. PINOS 229
TABLE 2. PER CENT COVER OF DOMINANT AND SUBDOMINANT UNDERSTORY SPECIES
Stands Species 1 2 3 4 5 6 fi 8
Artemisia tridentata 01% 68% 05% % % % % % Quercus turbinella (shrubs) 42.1
Sitanion jubatum 2.1 OWS A oS. a0 0.7
Pinus mono phylla (seedlings) 0.6
Bromus tectorum 23.5 16 5.4
Hesperochloa kingit 0.9 3.0
Eriogonum latifolium 0.7
E. kennedyi 1.9
Erigeron foliosus iby
Gayophytum diffusum 0.5 0.1 Eriastrum sapphirinum 0.1 Symphoricar pos parishii 10:2-°23:3— 1455 Collinsia torreyi 1.6 OZ Penstemon labrosus On Lupinus confertus 4.0 Achillea lanulosa 0.8
A rolling meadow occurred around the mountain summit. This open- ing contained many of the same species found in the upper forest, but was dominated by dwarf Chrysothamnus. About 50% of the area was covered by prostrate, cushion, or mat-forming species such as Astra- galus purshiu var. longilobus, Phlox diffusa ssp. subcarinata, Calyptri- dium umbellatum, Pedicularis semibarbata, and Allophyllum violaceum. These plants and the treelessness of the meadow gave it an almost al- pine aspect. The meadow was surrounded by Pinus jeffreyi with occa- sional trees growing in the meadow, except on the north rim where old P. flexilis and Abies concolor occurred. Some of these trees were without leaders, or were wind-topped or flagged, indicating that the summit ap- proaches climatic timberline.
In addition to the summit meadow, hydric associations dominated by [ris missourtensis occurred at upper elevations around seeps and springs. Other plants found in these wet pockets included the more common Carex athrostachya, the conspicuous Veratrum californicum, and oc- casional individuals of Barbarea orthoceras, Equisetum funstoni, Hele- nium bigelovit, Juncus balticus, Linum perenne ssp. lewisii, Mimulus guttatus, M. primuloides, Perideridia parishi, Potentilla glandulosa, Ranunculus cymbalaria var. saximontanus, Sisyrinchium bellum, Smil- acina stellata, Sphenosciadium capitellatum, Stachys albens, and Ver- onica americana.
A total of 46 families and 178 species were encountered. The largest percentages of species belonged to the Compositae (12.4%), Scrophu- lariaceae (8.4%), Leguminosae (7.5%), Onagraceae (3.9% ), Polygona- ceae (3.4%), and Gramineae (3.4%).
As in many California mountains and mountain ranges (Stebbins and Major, 1965), Mt. Pinos has a few endemic species, including Giulia
230 MADRONO [Vol. 19
modocensis type tetrabreccia which occurred in stands five through seven, Monardella linoides ssp. oblonga found in stand five, and Les- singia germanorum var. tenuis which was not sampled, but was present on the south slope.
Because of its central position, the Mt. Pinos flora contained plants also found in the San Joaquin Valley, Sierra Nevada, coast and trans- verse ranges, and the nearby desert (Munz, 1959). The Antelope Valley portion of the Mojave Desert is within 25 miles of Mt. Pinos and the desert flora was represented by such species as Anisocoma acaulis, Cal- ochortus kennedyi, both uncommon, and Chrysothamnus nauseosus ssp. mohavensis. The adjacent Central Valley was represented by Convol- vulus malacophyllus ssp. pedicellatus which was common at mid-eleva- tions, and Haplopappus acradenius ssp. bracteosus which only occurred at the slope base. The Sierra Nevada range is connected to Mt. Pinos by the Tehachapi Mountains and was represented by two uncommon species, Ribes velutinum and Astragalus whitneyi.
Tree species common to adjacent ranges, such as Pinus lambertiana, P. coulteri, P. sabiniana, P. murrayana, and Libocedrus decurrens, were not encountered on the south slope.
The wet iris meadows species have particularly wide distributions since most are found throughout western North America and some are common throughout the United States. Many are northern species, typical of the Sierra Nevada, Rockies, and parts of Canada which have apparently persisted in the semi-arid Southwest by occupying hydric habitats. Some of these species reach their southern limits in the vicin- ity of Mt. Pinos (Munz, 1959).
Abies concolor reproduction occurred both above and below the exist- ing range of Abies trees and grew under Pinus jeffreyi, indicating that the present P. jeffreyi forest could possibly be replaced by Adzes (table 1). Merkle (1954) made similar observations in ponderosa forests near Grand Canyon, Arizona. Abies might be able to replace P. jeffreyi be- cause it is more shade tolerant and can thus become established under P. jeffreyi canopies. Abies seedlings can survive lower temperatures than yellow pine, and can grow on accumulated litter and organic matter (Toumey and Korstian, 1962). Adzes is considered the most mesic conifer in southern California mountains, and, when it dominates a stand, other conifers, including P. jeffrewi, do not appear to become successfully established or grow under its dense canopies.
Pearson (1930) found that mature Pinus ponderosa, which is similar to P. jeffreyi, does not utilize moisture in the top 1 ft. of soil because of its deep tap root system. Abies’ moderately shallow, plate root system (Davis, 1959) could enable it to utilize this surface moisture, and thus become well established before having to compete directly with the mature P. jeffreyi for deeper soil moisture. Observations of Abies grow- ing immediately under and next to P. jeffreyi tend to support this hy- pothesis.
Ecologically, Abies appears to be the successional stage to replace
1968] VOGL & MILLER: MT. PINOS 231
much of the P. jeffreyi forest since it is the most shade tolerant, but apparently on Mt. Pinos, at least in the past, succession has been con- tinually retarded by disturbance factors.
Although there were no ungrazed areas available for comparison, it is probable that grazing has affected succession. Cattle, deer, and rabbits destroy pine and fir seedlings by severe browsing and girdling. Stunted Abies retarded by continual browsing were observed. One seed- ling-classed Abies suppressed by constant deer browsing was 1 ft. high, one-half inch in diameter, and 65 years old. Cattle grazing and concen- trations of burrowing rodents have undoubtedly caused the invasion of Artemisia tridentata and Chrysothamnus nauseosus by the elimination of the more palatable grasses and forbs and the creation of disturbed sites as they have done elsewhere (McKell and Chilcote, 1957).
Sparse understory is unique to southern California (fig. 1) because early descriptions of similar pine forests of other areas indicate that a grass cover was the most striking feature of the understory (Pearson, 1951; Weaver, 1951; Cooper, 1960). If grasses were originally abundant on Mt. Pinos, overgrazing would be the most plausable explanation for their disappearance, but this appears to be unlikely. Grazing probably reduced native perennial grass and forb cover on areas presently oc- cupied by Chrysothamnus, Artemisia, and Bromus tectorum. Savanna areas were probably more extensive before the advent of fire protection and offered more acreage than now present for the establishment of grasses. However, it is believed that a dense or continuous understory of grasses and forbs did not exist in the presettlement forest of Mt. Pinos, and that the understory cover was not much denser than it is today. This conspicuous absence, typical of Mt. Pinos and other conifer forests in southern California mountains, is probably produced more by the lack of summer precipitation during the growing season than by disturbance factors. This assumption is supported by Whittaker and Niering’s (1965) statement that, although the annual precipitation on southern California Mt. San Jacinto was sufficient to support trees, the shade and root competition of the overstory combined with summer drought severely limited understory development. Forests of inland mountain ranges, such as those in Arizona, characteristically produce more understory cover because of additional summer rainfall (Whittaker and Niering, 1965). The Mt. Pinos forest supported 10-50% (average 34%) grass and forb cover, while pine forests in the Arizona Grand Canyon region supported 40-60% ground cover (Merkle, 1962).
Grass species found on Mt. Pinos were Bromus carinatus, B. tectorum, Elymus condensatus, Hesperochloa kingii, Poa nervosa, and Sitanion jubatum. Bromus tectorum was the only introduced annual. Lack of abundant introduced annual grasses which have replaced native peren- nial grasses eliminated by grazing elsewhere in California (Biswell, 1956; Heady, 1958), also supports the hypothesis that Mt. Pinos has always had sparse understory cover.
Although data are lacking on the past vegetation of Mt. Pinos, nat-
232 MADRONO [Vol. 19
ural fires probably had the same retrogressive influence on the vegeta- tion as they had in other places (Weaver, 1951; Cooper, 1960), and were partially responsible for the present park-like appearance of the forest (fig. 1). United States Forest Service records indicated that 17 fires occurred on the south slope between 1919 and 1950. The majority were lightning-caused and resulted in fires of less than four acres. This low number of lightning-caused fires does not adequately reflect the high incidence of lightning occurring on Mt. Pinos. The upper mountain re- ceives as many as 600 lightning strikes during the summer months, with single storms often producing over 100 strikes (D. Roberts, fire control officer, personal communication). Some of these bolts strike open ground or rock; some hit trees, causing damage or death but not fire, or causing fire which is subsequently extinguished by rain (fig. 1). Occasionally, trees which have previously been injured by lightning, fire, wind, or fungi are ignited, producing “‘sleepers” that burn out the tree’s heart- wood. With drying conditions, the “sleepers”? may subsequently spread fire to the surrounding area. However, because of fire protection, these fires are usually detected and extinguished before they spread to other vegetation. Other strikes produce fires that burn individual trees or the immediate duff and understory, or fires that crown and burn a small group of trees. The lack of continuous understory and the scattered savanna growth tends to discourage widespread surface or crown fires.
Mature P. jeffreyi with evidence of having survived lightning strikes and recurring surface fires were common. A random sample of 277 trees on the upper slope contained 32.5% of the P. jeffrew: with lightning damage apparent in the form of spiral trunk or bark splits. Trees with damaged tops were not counted because wind and lightning damage could not be definitely differentiated. P. jeffreyi often survives surface fires because of its thick bark and moderately high, open tree crowns. Fifty trees with a 21 inch average dbh had a 2.1 inch average bark thickness. Though mature Abies concolor possess thick bark, they are usually less fire resistant that P. jeffreyi because the crowns are low and dense and individuals occur close together (Davis, 1959). Twenty A dies with an average 23.1 inch dbh had an average bark thickness of 2.5 inches. Young Pinus and Abies have thin bark and are fire susceptible.
Increment borings and stump ring counts were used to age a total of 75 trees. Data obtained indicated that trees and saplings occurred in small even-aged groups. Small groves of P. jeffreyi, consisting of three or more trees of the same age, were found to be 38, 63, 79, 85, 110, 205, 235, 253, 305, 350, and 358 years old. Each age group occurred over less than one-quarter to one acre. Recently cut P. jeffrey near the sum- mit were all about 250 years old. Two larger Abies (65 and 70 inch dbh) were of the same age. The pines occurred over about one acre, and probably originated after a lighting fire, since one larger P. jeffrey (56 inch dbh) was over 400 years old and had a deep basal fire scar en- larged by subsequent fires that dated back approximately 250 years.
1968] VOGL & MILLER: MT. PINOS 232
Apparently, presettlement fires were also local spot fires.
Although lightning fires were usually small, they had a pronounced effect on the vegetation because of their high frequencies. Recent light- ning strikes that were observed produced ground fires that consumed forest litter, thus creating pioneer soil conditions. Surface fires tempo- rarily reduced existing tree reproduction and shrubs, possibly enabling the overstory trees to obtain more of the limited moisture (Billings, 1950). Meyer (1931) found that when understory competition was re- duced by fire, released growth of overstory trees occurred within two years. Lightning strikes and fires were also observed to eliminate over- mature trees and insect or disease-infected trees. After each local fire, P. jeffreyi appears to replace itself, since it is apparently the best adapted species to re-colonize the fire-altered community (Biswell, 1963). Abies cannot return immediately because of its more mesophytic requirements. Fire on Mt. Pinos has produced a mosaic of small even- aged groups of trees or blocks of vegetation at the same successional stage (Cooper, 1961). The P. jeffreyi forest above 6000 ft. elevation has existed for centuries under continual bombardment from lightning and a subsequent patchwork of small fires.
With fire protection and the presently effective lightning fire suppres- sion, local pockets of the forest are tending to become overstocked with spindly, sapling-size P. jeffreyiz. A denser growth of saplings may pro- duce stunting because of competition, and the saplings may hamper overstory tree growth by competing directly with the trees for mois- ture (Arnold, 1963). This could reduce stand vigor and the trees might become susceptible to insects and disease. The increase in understory stocking may ultimately lead to the destruction of the entire forest by contributing to widespread crown fires (Leopold e¢ al., 1963).
Department of Botany, California State College at Los Angeles
LITERATURE CITED
ARNOLD, J. F. 1963. Uses of fire in the management of Arizona watersheds. Proc. Second Annual Tall Timbers Fire Ecol. Conf. 99-111.
BIswELL, H. H. 1956. Ecology of California grasslands. J. Range Management. 9:19-
24.
. 1963. Research in wildland fire ecology n California. Proc. Second Annual
Tall Timbers Fire Ecol. Conf. 62-97.
Birurncs, W. D. 1950. Vegetation and plant growth as affected by chemically al- tered rocks in the western Great Basin. Ecology 31:62-74.
Cooper, C. F. 1960. Changes in vegetation, structure and growth of southwestern forests since white settlement. Ecol. Monogr. 30:129-164.
. 1961. Pattern in ponderosa pine forests. Ecology 42:493-499.
Cottam, G., and J. T. Curtis. 1956. The use of distance measures in phytosocio- logical sampling. Ecology 37:451—460.
Davis, K. P. 1959. Forest fire: control and use. McGraw-Hill Book Co., New York.
Hatter, J. R. 1962. Variation and hybridization in ponderosa and Jeffrey pines. Univ. Calif. Publ. Bot. 34:123-166.
234 MADRONO [Vol. 19
Heapy, H. F. 1958. Vegetational changes in the California annual type. Ecology 39: 402-416.
Jenkins, O. P., and C. J. Kunpert. 1955. Geologic map of California. Los Angeles sheet No. N 3400. State of Calif. Div. of Mines.
Leopotp, A. S., S. A. Carn, C. H. Cottam, I. N. GAsrietson, and T. L. KImMBALt. 1963. Wildlife management in the national parks. Amer. Forests 69:32-35, 61-63.
McKett, C. M., and W. W. Cuitcore. 1957. Response of rabbitbrush following re- moval of competing vegetation. J. Range Management. 10:228-230.
MERELE, J. 1954. An analysis of the spruce-fir community on the Kaibab Plateau.
Ecology 35:316-321.
. 1962. Plant communities of the Grand Canyon Area, Arizona. Ecology 43:
698-711.
Meyer, W. H. 1931. Effect of release upon the form and volume of western yellow pine. J. Forest. (Washington) 29:1127-1133.
Mouwz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
OostTinc, H. J. 1958. The study of plant communities. W. H. Freeman and Co., San Francisco.
Pearson, G. A. 1930. Light and moisture in forestry. Ecology 11:145-160.
. 1951. A comparison of the climate in four ponderosa pine regions. J.
Forest. (Washington) 49:256-258.
STEBBINS, G. L., and J. Mayor. 1965. Endemism and speciation in the California flora. Ecol. Monogr. 35:1-35.
Toumey, J. W., and C. F. KorstiAn. 1962. Foundations of silviculture upon an eco- logical basis. Wiley and Sons, New York.
U.S. DEPARTMENT AGRICULTURE. 1941. Climate and man. U.S. Gov. Printing Office, Wash. D.C.
WEeaAvER, H. 1951. Fire as an ecological factor in the southwestern ponderosa pine forests. J. Forest. (Washington) 49:93—98.
WHitTTAKER, R. H., and W. A. Nierinc. 1965. Vegetation of the Santa Catalina Mountains, Arizona: a gradient analysis of the south slope. Ecology 46:429- 452.
NOTES AND NEWS
THE OCCURRENCE OF HEMITOMES CONGESTUM IN THE SOUTHERN SIERRA NEVADA, CaLIFoRNIA—Hemitomes congestum Gray has recently been collected from Sequoia National Park and Mineral King Game Refuge in Tulare Co. This, the second record of the species in the Sierra Nevada, is a range extension of approximately 250 miles south from an early collection (E. V. Schurr, May 18, 1934, UC) at Camptonville in Yuba Co.
We first collected this species during the summer of 1966 (Rice & Merryman 478, CAS, DAV) between Evelyn Lake and Hockett Meadow. Additional collections were made in Sept., 1967, when two fruiting specimens were obtained from widely separated locations along the Mineral King—Hockett Meadow trail (Rice 642a, DAV; Rice 642b, DAV). Several other small colonies of Hemitomes from which no specimens were taken were observed in the area.
Hemitomes congestum is best known from the redwood forest in northern Cali- fornia where it grows on humum below 2500 feet. Our collections are especially interesting in that they were made between 8,400 and 8,700 feet on the relatively dry, open floor of a red fir forest on soils of granitic origin. There was very little litter and few herbaceous plants in the vicinity of the colonies—BarBara RICE, University of California, Davis.
TAXONOMY, VARIATION, AND CHOROLOGY OF THREE CHROMOSOME RACES OF THE CALAMAGROSTIS CANADENSIS COMPLEX IN ALASKA
Wo. W. MITCHELL
Calamagrostis canadensis (Michx.) Nutt. (Bluejoint) is one of the most important native grasses in the herbland economy of subarctic Alaska. It predominates in tall grasslands of southcentral and south- western Alaska and is prominent in other mixed forb and grass com- munities. Calamagrostis canadensis demonstrates a wide ecological ampli- tude, occurring in lowland wet areas to windswept alpine ridges and in all major geographic regions from southeastern to arctic Alaska. It is rare or infrequent, however, in the Arctic (Wiggins and Thomas, 1962; Johnson, et al., 1966) and absent from the high Arctic (Porsild, 1957). The grass, considered in the wide sense, is circumpolar in distribution (Hultén, 1962).
Taxonomic treatments of this complex generally recognize two (Hultén, 1942; Porsild, 1951; Gjaerovoll, 1958; Anderson, 1959) or three (Steb- bins, 1930; Raup, 1947) subspecific entities in the far Northwest with some disagreement about nomenclature. Love and Love (1965) have attempted to resolve the differences and relate the taxonomy of the group to reported findings of two principal cytological races, 2n — 56 and 2n = 42, in North America. They concluded that C. canadensis s. str. should be restricted to the hexaploid level and that the octoploids should be referred to C. dangsdorfu (Link) Trin. Later these authors (Love and Love, 1966), concurring with Tzvelev (1965), combined C. langsdor fu with C. purpurea (Trin.) Trin. of Eurasia, it therefore becoming C. purpurea ssp. langsdorfi (Link) Tzvelev. But this treatment did not encompass the tetraploids (2n = 28) reported in eastern Asia as C. langsdorfi (Tateoka, 1954; 1967; Sokolovskaja, 1963). Love and Love (1965) considered these tetraploids to be C. angustifolia Kom.
However, the report of tetraploid C. langsdorfii merits further consid- eration. Seed for the type material of C. langsdorfii presumably originated on Kamchatka Peninsula in eastern Asia. Subalpine meadows in which bluejoint is an important component occur there that are very similar to those in portions of southwestern Alaska (Hultén, 1937a). My investi- gations reveal that Alaskan bluejoint is a complex consisting of three euploid races, the most abundant being tetraploid, new to the records for North America. These findings justify a taxonomic review of the group. Further, the chorology of the three Alaskan races lends itself to some interesting interpretations regarding the glacial history of the region and its effects on plant survival and migrations.
METHODS
Chromosome counts were obtained on root tips taken from vegetative material collected in the field and grown in the greenhouse. The tips
235
236 MADRONO [Vol. 19
were soaked in ice water for 16-20 hours, fixed in 3:1 alcohol and glacial acetic acid solution, hydrolyzed in 1N HC1 at c. 60°C for 8-12 minutes, and stained in Feulgen prior to squashing in aceto-carmine. A few counts were made on pollen mother cells collected in the field, fixed in 3:1 solution and smeared in aceto-carmine.
Specimens were analyzed morphologically for the following characters: 1, length of inflorescence; 2, length of spikelet; 3, length of lemma; 4, ratio: point of awn attachment measured from base of lemma/length of lemma; 5, length by which awn exceeds lemma; 6, length of an anther; 7, length of ligule; and 8, width of leaf.
RESULTS AND DISCUSSION
CyTOLOGyY and DISTRIBUTION. Tetraploid and hexaploid races of blue- joint dominate in Alaska. Octoploids appear to be rare except in particu- lar circumstances. Hybrids between the races also are rare, only a single presumed hybrid (2n = 49) having been found.
Counts were obtained on the following Alaskan specimens (mainly AES) with all but one of the collections made by the author.
2n = 28: W and SW Alaska: Galena, 727A33, 727A38; Bishop Rock, W of Galena, 727A4; Poorman, S of Ruby, 621D44, 621D30, 621D31; Lake Minchumina, N of McKinley Park, 619D29, 619D30, 619D31, 619D32; Farewell, SE of McGrath, 619D16; Medfra, E of McGrath, 621D39, 621D41; McGrath, 622D2; Takotna, W of McGrath, 621D5; Colorado Creek, NW of McGrath, 621D6, 621D7, 621D8; Flat, E of Holy Cross, 621D20, 621D21, 621D22, 621D23, 621D24; Stony River, on Kuskokwim R., 620D1; Sleetmute, on Kuskokwim R., 620D15; Red Devil, NW of Sleetmute, 620D9; Aniak, lower Kuskokwim R., 620D53; Anvik, on Yukon R., 620D27 ; St. Michael, S coast of Norton Sound, 620D24, 620D44, 620D46; Stebbins, W of St. Michael, 620D32; Togiak W of Dillingham, 822D2, 822D6; Dillingham, 822D14; Ekwok, NE of Dillingham, 822D15, 822D17, 822D18; King Salmon, 823D7, 823D10, 823D29; Iliamna, 823D17, 823D19.
Cook Inlet drainage region: meadow N of Homer, 712C26; subalpine grassland E of Anchor Point, 521B1, 521B2, 521B3; between Kenai and Soldotna, 712C15; Funny River road E of Soldotna, 105C1, 105C4; Alyeska, ski lodge SE of Anchor- age, 86D1; cleared areas vicinity of Palmer, 526A1, 526A2, 423B1, 423B2, 816B2, 816B3, 816B4; Little Susitna Valley N of Palmer, 715B1, 715B2, 98D9, 927D2; homestead W of Wasilla 921C5; Skwentna airstrip, Susitna Valley, 619D24, 619D25; Chelatna Lake W of Talkeetna, 622D7, 622D11, 622D14; Peters Creek W of Tal- keetna, 71A25, 71A26; Montana Creek S of Talkeetna, 72410.
Gulkana and Copper River basin: Mile 15.5 Tok Hwy. N of Gakona, 817D12; N of Paxson, 922C6; Tangle R. campground W of Paxson, 926C6; along Denali Hwy. about 54 miles NW of Paxson, 723B1.
Tanana River drainage: Mile 17 Taylor Hwy., 818D48, 818D34; Mile 4 Taylor Hwy., 818D9; Alaska Hwy. near Tok Juncton 818D3; Mile 115 Tok Hwy. S of Tok Junction 818D42; Mile 295 Richardson Hwy. NW of Delta Junction, O2Z3C 2: Mile 310 Richardson Hwy., 923C3; Mile 312 Richardson Hwy., 710A1; Fairbanks, 730A1; Mile 87 Steese Hwy. N of Fairbanks, 924C17.
Qn = 42: Western and southwetern Alaska: McGrath, 622D4; Dillingham, §22D 11.
Cook Inlet drainage region, alpine tundra: Lazy Mt. E of Palmer, 725D10; Eklutna Vally SW of Palmer, 728D17, 728D16; Eagle R. valley SW of Palmer, 97D9.
1968] MITCHELL: CALAMAGROSTIS 23
Gulkana and Copper River Basin: Eureka Lodge area, Glenn Hwy. SW of Glennallen, alpine tundra, 819D10; Mendeltna Creek, SW of Glennallen, 82C3; Mile 180 W of Glennallen, 82C19, 82C20, 817D4, 817D8; Mile 181.5 W of Glen- nallen, 719B16; along Richardson Hwy. 1 mle S of junction with Glenn Hwy., 731B7; Copper Center, 731B10, 819D1, 819D2, 819D6; McCarthy, 815C21; along road between McCarthy and Kennecott, 815C7; Tangle R. campground W of Paxson, 926C5; 5 mile N of Paxson, 624B12, 922C5, 922C7, 922C8; Mile 45 Tok Hwy. N of Gakona, 817D16; Mile 68.5 Tok Hwy., 817D26; Slana, near junction Tok Hwy. and Slana cutoff, Dickson & Taylor Aug. 10, 1961.
Tanana and Yukon R. drainages: Mile 30 Taylor Hwy. N of Tok Junction,, 818D23; Mile 312.7 Richardson Hwy. NW of Delta Junction, 925C5; Fairbanks, near international airport, 818B19, 818B21; Manley Hot Springs, 626B12; Birch Creek S of Circle along Steese Hwy., 924C11; 1 mile S of Circle, 924C16; Circle, terminus of Steese Hwy., 924C22.
2n = 49: Copper Center Lodge area Copper River Valley, 83C11.
2n = 56: Western Alaska: Bishop Rock, along Yukon R. W of Galena, 727A3, VETA 21:
Upper Cook Inlet region, alpine tundra, southcentral Alaska: Lazy Mt. E of Palmer, 725D2, 725D4, 725D5, 725D18, 725D20; Little Susitna Valley N of Palmer, 98D3.
Upper Copper River Drainage: Mile 68.5 Tok Hwy. S of Mentasta Pass, 817D27.
The tetraploid race predominates in Alaska’s southwestern and south- central coastal regions and probably northward along the coast at least to the Seward Peninsula (fig. 1). The extent of its occurrence beyond the Tanana River drainage in the drier Interior of Alaska is not known, but only hexaploids and octoploids have been reported so far across Canada (Bowden, 1960; Love and Ritchie, 1966; Nygren, 1954). Hexaploids are present in the upper Yukon River drainage of Alaska and may dominate there, but further sampling is needed. Both hexaploids and tetraploids are well represented in the Tanana Valley from the Canadian border to Fairbanks and in the relatively dry Copper River and upper Susitna River drainages south of the Alaska Range. A few hexaploids were found in alpine landscapes of the coastal southcentral area. The octoploid has been collected only at four locations — in westcentral Alaska, in eastcentral interior, and at two locations on alpine slopes of the upper Cook Inlet area. Occurrence of a possible hybrid between hexaploid and octoploid races at Copper Center implies the presence of octoploids there. The Arctic, Aleutian Islands, and southeastern panhandle have not been investigated cytologically as yet.
Of particular interest is the chorology of the three races in the upper Cook Inlet area. Dense stands of tall-growing bluejoint occur on burned- over timberland and in the subalpine zone above timberline throughout coastal southcentral Alaska. The bluejoint of these stands is tetraploid as determined by a large number of counts. However, octoploids and hexaploids were found in the alpine zone of the Chugach and Talkeetna Mountains near Palmer. Octoploids and hexaploids appear restricted to alpine sites in this area while tetraploids clearly have the advantage in occupying disturbances and dominating certain herbland communities at lower altitudes.
238 MADRONO [Vol. 19
es eo as e2n=42 aA2n = 56 - f o n= 9 aye
& x4, @
ef. Fic. 1. Distribution of specimens of the Calamagrostis canadensis complex in Alaska on which chromosome counts were obtained.
MorpHo.ocy. Calamagrostis canadensis is distinguished by its rela- tively stout rhizomes; generally tall robust culms, occasionally branched, with 3—6 nodes; relatively long, wide leaves; more or less open panicle; glumes with scabrous hairs throughout, often tending to diverge when empty; and thin membranous lemma with delicate awn and long, gen- erally widely spreading callous hairs.
Representative plants of the different races are shown in Fig. 2. Inflorescence form varies a great deal within each race (fig. 3), some of the variation obviously being related to environmental conditions. Clonal material collected at Galena, when grown in a fertilized experimental garden at Palmer, differed tremendously from its depauperate counter- part collected in the field (fig. 3, e and f), as did to a lesser extent ma- terial from the vicinity of the Fairbanks International Airport (fig. 3, n and 0). Some of these abnormally depauperate individuals resemble C.
1968] MITCHELL: CALAMAGROSTIS 239
#. # *
.
/ ) 71986, 2n=42
816B2, 2n=28 |
725D20, 2n= 56
Fic. 2. Representative vouchers of four chromosome races of Calamagrostis canadensis complex in Alaska.
angustifolia in Komaroyv (1934), but the spikelet characteristics are definitely those of C. canadensis. Circumstances appear to require their assignment to C. canadensis.
Other workers have had varying success in separating the different
[Vol. 19
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240
1968] MITCHELL: CALAMAGROSTIS 241
chromosome races. Bowden (1960) concluded that his hexaploid and octoploid plants of northern Canada could not be separated morphologi- cally. Love and Love (1965), on the other hand, found the octoploids in the alpine zone of Mt. Washington, New Hampshire, clearly distinguish- able from the hexaploids lower on the mountain. Similarly, in the current study spikelet measurements of the octoploids from alpine regions near Palmer were distinctly longer than the bulk of the tetraploids and the few hexaploids found in the same area.
No good qualitative differences were found, however, for distinguish- ing the races and all overlapped in their measurements (fig. 4 and table 1). Overlapping between the tetraploids and hexaploids is so great as to preclude considering separating these two. The apparent separation of the octoploid is confounded by the occurrence of an ecotypic trend in sizes of plant parts. Those in the interior tend to have smaller parts than those in the coastal regions; for instance, spikelet length of 14 tetra- ploids of the Cook Inlet region averaged 4.13 mm compared with 3.60 mm for 15 tetraploids of the more northern interior. The 3 octoploid specimens from the interior averaged 4.6 mm, 3.5 mm, and 2.0 mm for spikelet-, lemma-, and anther-length, respectively. Except for anther length these values are within a standard deviation of the average values determined for the hexaploids (table 1). A number of plants analyzed from the Southeast and the Aleutian Islands, cytology unknown, had particularly long spikelets, those from the Aleutian Islands averaging well above the measurements obtained on the octoploid vouchers. A teraploid composition is indicated for the Aleutian Island population (fig. 1), although it may be otherwise. Thus, any effort to differentiate the cytological races morphologically must take into consideration geographic occurrence.
TAXONOMIC DISPOSITION. There is an obvious contradiction in the various interpretations of the langsdorfu type. Hultén (1942; 1962) differentiates ssp. langsdorfiu from ssp. canadensis by its longer spikelets, larger panicle, broader leaves, and taller growth. He considers it repre- sented across North America but in Alaska more coastal in its occur- rence than ssp. canadensis. Love and Love (1965; 1966), and others, concur with Hultén in associating /angsdor fii with the more robust growth habit; further, these authors assign it to the octoploid level.
It appears, however, that the tetraploid material of eastern Asia and that of Alaska are conspecific and relate to the type material of langsdor fii. Moreover, the tetraploids as a rule have smaller spikelet measurements
Fic. 3. Variation in inflorescence form and size in the different races of Calama- grostis canadensis in Alaska: 2n = 28: a, Mitchell 922C8; b, 921C5; c, 712C15; d, 96D1; e, 727A38; and {, clonal material of 727A38 grown in experimental garden at Palmer. 2ni= 56: 2; /27A21; h, 98D3; and j, 725D20. 2n = 42: k, 292C7; 1, 82C20; m, 922C5; n, 818B21; and o, clonal material of 818B21 grown in experimental garden at Palmer.
242 MADRONO [Vol. 19
leaf width 2n = 28 — — 2n = 42 ——— — 2n = 56 1. of ligule Oe he j _— ~~ 1. of infl. : pis ree of in NZ : ; A oo oe * Yr. 7 ar: VON 7 N a2 ji Nae ns ye / Woes ry —— : ~~ ~ 1. of anther oo / 1. of spklt. za y. i Ff} / ra S \ NN oe eR > a Bf Ho H.
1. of awn beyond lemma
awn attachment
Fic. 4. Polygonal representation of morphological analysis of three chromosome races of Calamagrostis canadensis in Alaska based on information contained in Table 1. Total range, mean, and range of two standard deviations represented for each character. Means are interconnected by lines for each race.
than the octoploids and apparently are restricted to eastern Asia, Alaska, and possibly the Yukon Territory and coastal regions to the south of Alaska.
Biologically there may be reason to recognize all three races as specific entities. But morphological considerations preclude the separation of the tetraploid and hexaploid races. In some local areas floral parts may serve, to a certain extent, to separate the octoploid from the other two races. This is much less certain, however, as collections from different areas become involved. In view of the small sample of octoploids so far con- sidered I believe formal recognition of the three races should be deferred until their ranges and morphological distinctions are better delineated.
1968] MITCHELL: CALAMAGROSTIS 243 TABLE I. QUANTITATIVE ANALYSIS OF THREE CHROMOSOME RACES OF THE CALAMAGROSTIS CANADENSIS COMPLEX.
Number of specimens of each race analyzed: 37 of 2n = 28, 33 of 2n = 42, and 8 of 2n = 56.
Chromosome Range of Character Race, 2n = Range Mean ey By. Two S.D. Leaf width, mm 28 3.0- 9.0 4.9 1.39 oie 0? 42 2.5- 8.0 50 1.42 3.6 -— 64 56 3.2- 6.0 4.8 OL 40 -— 5.6 Length of inflorescence, cm 28 6.0 — 18.0 11.9 Zea 9.1 -—14.7 42 8.5 —21.5 13.5 3.63 9.9 -17.1 56 9.5-—19.5 14.0 3.14 10.9 -17.1 Length of spikelet mm 28 3.0- 5.7 3.9 56 oot = An 42 3.1- 5.3 4.3 59 3.7 — 4.9 56 43-— 5.8 52 55 47 — 5.8 Length of lemma, mm 28 ZO 31 2.9 34 2.6 — 3.2 42 2.5- 3.9 Sed 36 2.6 =. 3.6 56 3.5— 4.5 3.8 96 3.5 = 42 Awn attachment/ length 28 16- 88 A1 17 24- .58 of lemma 42 03- .71 OS 13 20- 46 56 O9-— 47 26 is 13- 39 Length by which awn exceeds 28 0.0- 1.8 .60 38 22- 98 lemma, mm 42 0.0- 1.7 45 40 05-85 56 0.3- 0.9 .60 24 36- 84 Length of anthers, mm 28 11-— 2.2 Ss) af 1.2 - 1.8 42 Del ee 1.6 24 14 -— 1.8 56 16-— 2.3 20 27 1.7 -— 2.3 Length of ligule mm 28 3.5 —10.0 6.9 1.61 5.3 - 8.5 42 4.5-—11.0 ee 1.79 5.4 -— 9.0 56 6.0 — 12.0 Se 2.10 6.1 —10.3
PHYTOGEOGRAPHIC CONSIDERATIONS. Theories on plant survival and migration in glacial and post glacial times propose the revegetation of glaciated southcentral Alaska by plants migrating from the unglaciated interior and the western coastal and exposed Bering Strait region (Hultén, 1937; Heusser, 1967). Heusser also has proposed the survival of plants in small refugia along the southern coast of Alaska. Migration of the octoploids from the interior to their current position in the Cook Inlet area would require their traversing expanses that today are occupied by the obviously more aggressive hexaploids and tetraploids. Some evi- dence of a migrational history was found in the occurrence of an octo- ploid plant (Mitchell 817D27) on deglaciated terrain at the edge of a ponded kettle near the Mentasta Pass of the Alaska Range. A hexaploid (Mitchell §17D26) was found at the same site.
Certainly with a more comprehensive effort the known distribution of the octoploid would become less disjunct than represented in Fig. 1, but it obviously is rare to infrequent through much of Alaska. Octoploid bluejoint would appear to fit Hultén’s (1937b, p. 20) description of a taxon losing its variability through periodic reductions by glacial ad-
244 MADRONO [Vol. 19
vances, thereby becoming rigid and unable to spread. I think it unlikely that the current distribution of the octoploid represents a range reduc- tion following a postglacial migration from interior Alaska. Rather, its presence in the alpine regions of the upper Cook Inlet area appears to be that of a relic surviving a more extensive distribution attained prior to the late glacial period. (Also see Love (1959) concerning the origin and distribution of high polyploids in the pre-Pleistocene and early Pleisto- cene periods.) I hold a similar view for the octoploids found in west- central Alaska on Bishop Rock, a conspicuous knob in the flats of the Yukon River west of Galena, where a possible relic brome also was dis- covered (Wilton, 1965; Mitchell and Wilton, 1966). Bishop Rock and other heights in this unglaciated area contain some relatively dry, shaly bluffs that are not subject to the disturbances of the adjacent lowlands and which support plants that do not occur in the more moist, wooded uplands of the area. These are suitable sites for the persistence of relics adapted to their particular conditions.
In the upper Cook Inlet area tall herbaceous and shrubby subalpine communities extend above timberline to the limits of well marked glacial morainic features (Mitchell, 1968a). Karlstrom (1964) cited 2,000 ft. as the elevation of the lateral moraines of the late Wisconsin ice advances on the upper Kenai Peninsula. Moraines occur at about 2,800 ft. on the Chugach and Talkeetna Mountains north of Anchorage. Well developed alpine communities obtain above these morainic features to over 5,000 ft. on the less precipitous mountain slopes. I think extensive areas of these alpine reaches very likely served as refugia for plants during the last ice advances. Finding hexaploid and octoploid races only on what appear to be unglaciated alpine sites in a region where the tetraploid is eminently successful lends support to the theory. Further evidence was found in the occurrence of a tetraploid colony (Mitchell 927D2) on an alpine site, about 3,700 ft., that obviously had been glaciated. This sug- gests a cytological distinction between populations in the upper Cook Inlet region in relation to duration, those of relatively recent origin on glaciated terrain being tetraploid and those of longer duration on un- glaciated terrain being hexaploid or octoploid. This theme concerning the existence of nunataks in southcentral Alaska (Mitchell, 1968b) will be developed more thoroughly in a future article.
ACKNOWLEDGMENT
I am grateful to The Rockefeller Foundation for its support of this study through grants RF-58108 and RF-61036, to the U. S. Department of Agriculture NC-7 New Plants Committee for travel support, and to Alaska Experiment Station staff members A. H. Mick, L. J. Klebesadel and R. L. Taylor for their constructive criticism of the manuscript.
University of Alaska Agricultural Experiment Station, Palmer
1968] MITCHELL: CALAMAGROSTIS 245
LITERATURE CITED
ANDERSON, J. P. 1959. Flora of Alaska and adjacent parts of Canada. Iowa State Univ. Press, Ames.
Bowpven, W. M. 1960. Chromosome numbers and taxonomic notes on northern grasses. III. Twenty-five genera. Canad. J. Bot. 38:541-557.
GJAEREVOLL, O. 1958. Botanical investigations in central Alaska, especially in the White Mountains, Part I, Pteridophytes and Monocotyledones. Kongel. Norske Vidensk. Selsk. Skr. (Trondheim), No. 5.
HevusseEr, C. J. 1967. Pleistocene and postglacial vegetation of Alaska and the Yukon Territory. Jn H. P. Hansen (ed.), Arctic Biology, 2nd ed. Oregon State Univ. Press, Corvallis.
Hutten, E. 1937a. Flora of the Aleutian Islands. Bokforlags Aktiebolaget Thule, Stockholm.
. 1937b. Outline of the history of arctic and boreal biota during the Quaternary Period. Bokforlags Aktiebolaget Thule, Stockholm.
. 1942. Flora of Alaska and Yukon. II. Monocotyledoneae. Acta Univ. Lund 38:131-412.
. 1962. The circumpolar plants. I. Vascular cryptogams, conifers, monoco- tyledons. Almqvist & Wiksell, Stockholm.
Inman, O. L. 1922. Calamagrostis canadensis and some related species. Rhodora 24:142-144.
Jouwnson, A. W., and L. A. Vrrreck, R. E. Jonnson, and H. Metcuor. 1966. Vege- tation and flora of the Cape Thompson-Ogotoruk Creek area, Alaska. In N. J. Wilimovsky and J. N. Wolfe (eds.), Environment of the Cape Thompson region, Alaska. U.S. Atomic Energy Commission.
Komaroy, V. L. (ed) 1934. Flora of the U.S.S.R. (Transl. from Russian, 1963). U.S. Dep. Com.
Love, A. 1959. Origin of the arctic flora. Publ. McGill Univ. Mus. 1.
., and D. Love. 1965. Taxonomic remarks on some American alpine plants. Univ. Colo. Stud., Ser. D, Phys. Sci. 17.
. 1966. Cytotaxonomy of the alpine vascular plants of Mount Washington. Univ. Colo. Stud., Ser. D, Phys. Sci. 24.
Love, A., and J. C. Ritcure. 1966. Chromosome numbers from central northern Canada. Canad. J. Bot. 44:429-439.
MircHetrt, W. W. 1968a. On the ecology of sitka alder in the subalpine zone of southcentral Alaska. In J. M. Trappe, et al. (eds.), Biology of alder. U. S. Forest Servce, Pacific Northw. Forest and Range Exp. Sta., Portland, Oregon. In press.
. 1968b. Local refugia in southcentral Alaska during the Naptowne glacia- tion. Proc. 18 Alaskan Sci. Conf. In press.
. and A. C. Witton. 1966. A new tetraploid brome, section Bromopsis, of Alaska. Brittonia 18:162—166.
Nycren, A. 1954. Investigations on North American Calamagrostis. I. Hereditas 40:377-397.
Porsitp, A. E. 1951. Botany of southeastern Yukon adjacent to the Canol Road. Nat. Mus. Canada Bull. 121.
. 1957. Illustrated flora of the Canadian arctic archipelago. Nat. Mus. Canada Bull. 146.
Raup, H. M. 1947. The botany of southwestern Mackenzie. Sargentia 6.
SokoLovskKaya, A. P. 1963. Geograficheskoe rasprostranenie poliploidnykh vidov rasteny (issledovanie flora polustrova Kamchatki). Vestn. Leningradsk. Univ. Ser. Biol. 15:38-52.
STEBBINS, G. L. 1930. A revision of some North American species of Calamagrostis. Rhodora 32:35-57.
TaTEOKA, T. 1954. Karyotaxonomic studies in Poaceae I. Nat. Inst. Genet. Ann. Rep. 4:45-47.
246 MADRONO [Vol. 19
. 1967. In IOPB chromosome number reports. Taxon 16:562. TzvELEv, N. 1965. De genere Calamgrostis Adans. In URSS notulate systematicae. Novit. Syst. Plant Vasc. 1965: 5-50. Wiccrns, I. L., and J. H. THomas. 1962. A flora of the Alaskan arctic slope. Univ. Toronto Press. Witton, A. C. 1965. Phylogenetic relationships of an unknown tetraploid Bromus, section Bromopsis. Canad. J. Bot. 43:723-730.
NEW RECORDS OF MYXOMYCETES FROM CALIFORNIA. ITI. DoNALD T. KOWALSKI and DWAYNE H. CurTIS
The number of species of slime molds reported to date in California is 180. Ten new records are listed in this paper. This brings the total number of Myxomycetes reported in print as occurring in California to 190 species. All collections cited here (MICH) are those of the senior author. The names of the organisms are those accepted by Martin (1949) and the numbers are those of the senior author. This investigation was supported by National Science Foundation Grant GB-5799.
We wish to express our appreciation to Travis E. Brooks of the Uni- versity of Kansas for verifying the identifications of some of the more difficult collections.
LICEACEAE
Licea biforis Morgan. On decaying bark, 2 miles west of Child’s Meadows, 4,400 feet elevation, Tehama Co., 2747, Apr. 16, 1966. This species is distinctive in that it is the only member of the genus with sporangia which are repent, fusiform and open by a single longtitudinal slit. Most descriptions give the color as yellow-brown, in this collection the sporangia are black and the depressed longitudinal slit is light tan. Its favorite habitat is stated as being the inner layers of dead bark. The fruiting bodies in this collection are on the outer surface of coniferous bark. This taxon has a very scattered distribution pattern, being known in this country only as far west as Kansas. However, it is probably fairly common. Its rareness is undoubtedly due to its inconspicuousness and may occur throughout the United States.
Licea tenera Jahn. On sheep dung, 3 miles south of Chico, Butte Co., 6238, May, 1967, developed in a damp chamber. This species is mainly known from moist chamber developments. The type collection consists of approximately a dozen sporangia. The California material fits the type material in that the sporangia are sesile, globose, orange and inde-
1968] KOWALSKI & CURTIS: MYXOMYCETES 247
hiscent. It differs from the type, however, in size and in spore character- istics. The type sporangia are 0.4-0.5 mm in diameter while the Cali- fornia material averages about 0.2 mm in diameter. The spores in the type collection have short spines and a pale area on one side, the Cali- fornia material has spores which are smooth and lack the pale area. In regards to the size of the sporangia this collection resembles two collec- tions from Kansas which are 0.1 mm in diameter. The Kansas material, however, consists of sporangia which are thickly coated with refuse material which makes them appear black. Because of these discrepancies the exact taxonomic standing of this taxon is in doubt. It is extremely rare, being known from only a few collections throughout the United States. It has, however, been reported from Oregon.
CRIBRARIACEAE
Cribraria splendens (Schrad.) Pers. Three collections, all on decay- ing coniferous bark, MacKerricher Beach State Park, Mendocino Co., 4800, 4814, 4528, Jan. 25, 1967. These collections are both abundant and fit the published descriptions perfectly. This species is distinct in the genus because it is one of the few members that lacks a distinct cup. The cup is replaced by firm ribs, usually 8-15, which radiate from the tip of the stalk. Occasionally these ribs are connected by a delicate, hyaline membrane. It is an uncommon taxon, but is known from widely scattered regions across the United States, like most species in the genus, it is found mainly in coniferous forests.
DIANEMACEAE
Dianema andersoni Morgan. Two collections, both on decaying co- niferous wood, Well’s Cabin Campground, 6,300 feet elevation, Tehama Co., 1556, 1613. This is one of the true snowline Myxomycetes, i.e., it fruits only under or near the melting snow and is exposed when the snow melts back. This species and D. harveyi Rex are very difficult to separate. Traditionally they are distinguished as follows: D. andersonii is sporangi- ate, sessile or substipitate, brown and has spores which are pinkish by transmitted light, warted and 10-12 » in diameter. Dianema harveyi is sporangiate to plasmodiocarpous, always sessile, dull red or brown, and the spores are pale yellowish by transmitted light, spiny and 8-10 yw in diameter. As can be seen, these differences are slight and it is possible that both of these species simply represent different forms of the same taxon. The major problem is that they are both known from just a few collections and until sufficient material is available, the exact taxonomic ranking of these species will not be resolved. Dianema andersonii has been reported in the Western Hemisphere only from British Columbia and Washington.
STEMONITACEAE
Amaurochaete fuliginosa (Sow.) Macbr. Three collections, all on de- caying bark, Woodson Bridge State Park, Tehama Co., 6041, 6060, 6062.
248 MADRONO [Vol. 19
This aethaloid member of the Stemonitaceae is characterized by having spinulose spores 12—15 » in diameter and a capillitium that is rigid and irregular. It was found in abundance on the bark of a large oak, Quercus lobata Neé, that was recently felled by a storm. Most of the aethalia are small for this species, being only 2—3 cm in diameter. Some descriptions state that the hypothallus is a slender stalk-like strand. In the California collections the hypothalli are extensive, procumbent and attached to the substrate along their entire lower surfaces. It is an uncommon taxon, being reported from scattered localities across the United States and on the west coast from Washington.
Comatricha rubens List. Three collections, two from Lower Bidwell Park, Chico, Butte Co., 2323 on decaying bark, Jan. 2, 1966, 4046 on a decaying leaf, Dec. 9, 1966, and one on decaying Eucalyptus bark, Point Reyes National Seashore, Marin Co., 5164, Jan. 29, 1967. This species is distinct in the genus by having light-colored lilaceous or ferruginous spores and sporangia with a distinct basal cup which is attached to the capillitium. It is extremely common in California and on all types of substrates. We have found it on dead wood, dead leaves, dead bark and on various other types of plant debris. For the sake of brevity, we only listed three collections, but we have made over 50 col- lections. It is not a rare taxon, being known from many scattered locali- ties across the United States.
PHYSARACEAE
Badhamia affinis Rost. Two collections, both on decayed leaves, Butte Creek and Skyway, Butte Co., 3960, 3982, Nov. 25, 1966. The sporangia are white, rugulose, rounded to distinctly flattened. Both collections con- tain sporangia which are either stalked or sessile. When stalked, the stipe is very short, about 1 mm in height, black, and furrowed. The only atypical characteristic about these two collections is the habitat. Both Hagelstein (1944) and Martin (1949) give the substrate as bark, either living or dead. Thus, the occurrence of this species on decayed leaves appears to be unsual. This species is not rare. It is cosmopolitan and is known from many collections in the United States.
Badhamia ainoae Yamashiro. Four collections, all on the bark of living oak trees, Moores Ranch, 12 miles south of Chico, Butte Co., 2337, Jan. 7, 1966, 2461, Feb. 6, 1966; Lower Bidwell Park, Chico, Butte Co., 2341, Jan. 8, 1966, 4389, Dec. 22, 1966. This species is unique in the genus in that the capillitium is represented by tubular calcareous col- umns, simple or forked, which extend from the base to the peridium. This is somewhat reminiscent of the tubes in Didymium sturgsiu Hagelst. The fruiting bodies are sessile, pulvinate sporangia or short plasmodi- ocarps and are widely scattered. They vary from brownish, when lime is lacking, to white in color when the peridium is covered with a thin calcareous crust. These collections mark the first reported occurrence of this species in the Western Hemisphere. Travis E. Brooks of the Uni-
1968] KOWALSKI & CURTIS: MYXOMYCETES 249
versity of Kansas, Lawrence, Kansas has made several collections in the Eastern United States and all of them were also on the bark of living trees (personal communication). It is previously only known in print from Japan. It is apparently a very rare taxon, all known collections being on the bark of living trees. It is possible, however, that it may be much more common than its distribution would indicate. This could be due to the fact that very few collectors look for Myxomycetes on the bark of living trees.
Badhamia dearnessii Hagelst. Two collections, both found 12 miles south of Hamilton City, Glenn Co., Dec. 29. 1966, 4528 on plant debris and 4547 on decaying bark. Previously, this species was only known from a few collections in Quebec and Maine. It is distinct because it is the only species in the genus that has spores which are encicled by a pale band. All descriptions of this taxon state that the sporangia are sessile. In 4547 many of the sporangia are stalked. The stalks are short, 1-2 mm in length, and of a pale yellow color. The descriptions also indicate that the capillitium is a delicate network scantily charged with lime or nearly limeless and then appearing pale yellow. In both of the California collections the capillitium is robust, pure white, and densely filled with lime, there being no limeless threads whatsoever.
Physarum gilkeyanum H, C. Gilbert. On decayed leaf, along the Sacramento River, 1.7 miles south of Sacramento Road, Butte Co., 988, Dec. 14, 1964. The description of this species states that the sporangia are sessile, white, single-walled and covered with limy ridges which give the appearance of a reticulum. The California collection agrees with this description except that only a few of the sporangia have the limy reticu- lum. Since this is an extremely rare species, being known previously only from the type locality in Lebanon, Oregon, we do not believe that this slight discrepancy should be regarded as important enough to rele- gate the California material to a different taxon. Hagelstein (1944) believed that this taxon was probably a sessile form of Physarum mutabile (Rost.) List. We do not concur with this interpretation. We have made several collections of P. mutabile in California and all of the collections contain sporangia which are stalked and have a large cylin- drical columella, which almost attains the apex of the sporangium. Some of the sporangia of P. gilkeyanum have a slight aggregation of lime knots at the base of the sporangia which slightly resembles a pseudocolu- mella. However, they do not remotely approach the large cylindrical columella of P. mutabile. Thus, we believe that these two species are distinct from one another and are not synonymous.
Department of Biology, Chico State College, California
LITERATURE CITED
HAGELSTEIN, R. 1944. The Mycetozoa of North America. Mineola. Martin, G. W. 1949. North American Flora 1:1—190.
CHROMOSOME COUNTS IN SECTION SIMIOLUS OF THE GENUS MIMULUS (SCROPHULARIACEAE). VIII. CHROMOSOMAL HOMOLOGIES OF M. GLABRATUS AND ITS ALLIED SPECIES AND VARIETIES.
M. M. MiA and Robert K. VICKERY, JR.
The purpose of this study is to extend our analysis of the chromosomal homologies of the Mimulus glabratus complex and its allied species and varieties previously initiated by Mukherjee and Vickery (1962) in the course of their investigation of the M. guttatus complex. Essentially the same cytological methods and crossing techniques were employed as be- fore (Vickery, et al, 1968).
The M. glabratus complex was represented by cultures of 10 popula- tions. One was of M. pilosiusculus HBK., a near relative of M. glabratus HBK.., and the others were of three of the varieties of MW. glabratus, M. glabratus var. utahensis Pennell, M. glabratus var. fremontu (Benth.) Grant, and M. glabratus var. parviflorus (Lindl.) Grant (table 1). The populations represent the diploid, tetraploid, and hexaploid chromosome levels of the complex, n = 15, n = 30, and n — 45 respectively. They also include representatives of two aneuploid levels, n = 14 and n = 46,
For comparisons, cultures of eight additional populations, representing the three other main complexes of species in section Simiolus were also included in the investigation. The M. guttatus complex was represented by diploid, n = 14, and tetraploid, n = 28 populations of VW. guttatus DC. and by an aneuploid, n = 13, population of MW. nasutus Greene. The M. luteus complex was represented by two tetraploid, n = 32, populations of M. luteus L. The M. tilingii complex was represented by a single di- ploid, n = 14, culture of MW. tilingii Regel.
Cultures of the 18 selected populations were established in the green- house for the crossing program. Over a hundred interpopulation hybridi- zations were made. Only 48 of these produced hybrids and of these only 33 developed far enough to permit cytological study. The pairing behavior of the chromosomes was analyzed in an average of over 14 microsporo- cytes for each different hybrid. Typical figures were recorded either photographically or with the aid of a camera lucida.
At the diploid, n = 14 level, WM. glabratus var. utahensis hybridized with the main species of section Simiolus, M. guttatus, n = 14, and the chromosomes of the resulting F, hybrids revealed 6 to 14 bivalent associ- ations (table 2). The observed minimum pairing of six suggests that only these six are fully homologous or so nearly so as to consistently synapse. The other eight chromosomes probably are not fully homologus as evi- denced by their occasional failure to pair. On the average, 12 of the 14 chromosomes pair. The basic genome of 14 chromosomes apparently has become significantly differentiated in M. glabratus var. utahensis as com- pared to that of M. guttatus, representing the diverse M. guttatus complex.
250
1968] MIA & VICKERY: MIMULUS 251
TABLE 1. ORIGIN OF CULTURES USED IN THE CYTOGENETIC INVESTIGATION OF THE RELATIONSHIPS OF MIMULUS GLABRATUS AND ITs ALLIED SPECIES
M. glabratus var. fremonti (Benth.) Grant. 5063, n = 30, San Bernardino Co., Calif., ca. 1,200 ft, coll. - (UC); 5373, n = 30, Brewster Co., Texas, Cory 53,186 (UT).
M. glabratus var. parviflorus (Lindl.) Grant. 5041, n = 45, Illapel, Chile, 4,000 ft, U.S.D.A. Plant Introduction number 144,534 (UT); 6162, n = 46, Auetrihue, Argentina, Diem in 1959; 6163, n = 46, Lumaco-Puente del Cina, Chile, 600 ft, Kunkel, Nov. 29, 1958; 6184, n = 46, Tucuman, Argentina, 7,300 ft, de la Sota, Feb. 7, 1959.
M. glabratus var. utahensis Pennell. 5048, n = 14, Mono Co., Calif., 6,440 ft, Stebbins 714 (UT); 5747, m = 14, Mineral Co., Nev., 5,500 ft, J. Figg-Hoblein, July 4, 1950; 5265,n = 15, Wayne Co., Utah, 7,100 ft, Vickery 600.
M. guttatus DC. 6152, mM = 28, Admiralty Island, Alaska, sea level to ca. 1,000 ft, G. L. Miller, Aug. 1958; 5052, n = 14, Contra Costa Co., Calif., 1,000 ft, Stebbins 703 (UT); 6250, n = 28, Yavapi Co., Arizona, 3,010 ft, Vickery 2593; 5017, n = 14, Inyo Co., Calif., 2,500 ft, Alexander and Kellogg 2844 (UC).
M. luteus L. 5042, n = 32, Illapel, Chile, 6,200 ft, U.S.D.A. Plant Introduction number 144,535 (UT) ; 6161, n = 32, Auetrihue, Argentina, Diem in 1959.
M. asutus Greene, 5327, n = 13, Tuolumne Co., Calif. 475 ft, Vickery 168 CUM.
M. pilosiusculus HBK. 5320, n = 46, Botanic Garden, Copenhagen, Denmark (wild in Argentina, Chile, and Peru), U.S.D.A. Plant Introduction number 181,130 CUE):
M. tilingit Regel. 5012, n = 14, Mono Co., Calif., 10,000 ft, Clausen 2075 (UT).
In the hybrids of M. glabratus var. utahensis * M. tilingii, represent- ing the M. tilingit complex of North American alpine species, the chromo- somes exhibited 13 or 14 bivalent associations (table 2). Apparently the glabratus genome is almost fully homologous with the ¢ilingi genome.
The fourth complex of section Simiolus, the M. luteus complex of South America, as represented by M. luteus (n = 32) crossed readily with VM. glabratus var. utahensis but produced sterile hybrids. The chromosomes of its F, hybrids exhibited 6 to 14 bivalent associations (table 2). Ap- parently the first genome of M. luteus has about the same degree of homology with the glabratus genome as does the guttatus genome. The guttatus and luteus genomes are not identical but have differentiated from each other. The chromosomes of the F, hybrids of M. guttatus M. luteus exhibit 11 to 13 bivalent associations (table 2) which suggests that they are about as different from each other as they each are from the chromosomes of the genome of M. glabratus var. utahensis but that different genes are involved in the differences.
At the tetraploid level, the 7. glabratus complex was represented by two, n = 30 cultures of M. glabratus var. fremontii. The two cultures, 5063 and 5373, are from widely different areas (table 1). The cultures cross readily in either direction, 7.e., 5063 & 5373 or 5373 « 5063, but the F, hybrids produced were sterile. Cytological examination of meiosis in these hybrids revealed perdominantly 14 bivalent chromosome associ- ations and only occasionally more pairs (table 2). The first genomes of
252 MADRONO [Vol. 19
TABLE 2. PAIRING BEHAVIOR OF THE CHROMOSOMES IN THE 1s HYBRIDS OF MIMULUS GLABRATUS AND ITS RELATED SPECIES (FIGS. 1, 2).
Combinations, including Q Parent $ Parent Number Number of reciprocals. culture culture of cells bivalents observed (g. = glabratus) number n= number n= studied \ range
M. g. var. utahensis X 5747 14 5048 14 8 13 9-14 M. g. var. utahensis 5048 14 5265 15 14 13 11-14 5747 14 5205 15 14 12 8-14 . g. var. utahensis X 5063 30 5747 14 16 14 9-16 M. g. var. fremonttu 5373 30 5747 14 14 14 9-15 . g. var. utahensis X 6184 46 5747 14 14 14 14-15 M. g. var. parviflorus M. g. var. utahensis X 5048 14 5052 14 LT 13 11-14 M. guttatus 5747 14 5052 14 8 i} 6-14 6162 28 5048 14 14 14 all 14
S
S
M. g. var. utahensis X* 5747 14 5012 14 15 14 13-14 M. tilingi
M. g. var. utahensis X 5042 32 5048 14 12 13 6-14 M. luteus
M. g. var. fremontii X 5373, 330,—'—s«sS5063—S— 380'—i2stidSsts«d2‘-DD M. g. var. fremont
M. g. var. fremonti X 5041 45 5373 30 19 14 12-17 M.g.var. parviflorus 6184 46 9373 30 30 14 11-21
M. g. var. fremonti xX 5320 46 o/s 30 30 16 14-20 M. pilosiusculus
M. g. var. fremontiu X 5063 30 5052 14 7 12 2-15 M. guttatus D373 30 3052 14 9 12 6—14
M. g. var. parvifloras X 5041 45 6162 46 1S 15 11-26 M.g. var. parviflorus 5041 45 6163 46 il) 18 12—22
M. g. var. parviflorus X 6162 46 6163 46 20 19 11-27 M. pilostusculus 6184 46 5320 46 14 15 14-16
M. g. var. parviflorus K 5041 45 5052 14 4 16 all 16 M. guttatus 6162 46 5052 14 6 13 12-14
6163 46 5052 14 4 13 9-14 6184 46 5052 14 24 14 12-14 M. g. var. parviflorus X 6184 46 6161 32 16 15 11-19
M. luteus
M. pilosiusculus X 5320 46 5052 14 5 14 14-15 M. guttatus M. guttatus X 6152 28 5052 14 39 13 5-14 M. guttatus 5052 14 6152 28 5 14 all 14 6250 28 5052 14 31 13 9-14 M. guttatus X 6152 28 5042 32 10 15 9-18 M. luteus 5017 14 5042 32 4 12 11-13 M. guttatus X 5017 14 SEA 13 1 13 13 M. nasutus
these two populations appears to be almost fully homologous. The chrom- osomes of the second genomes exhibit modest homology under the most favorable conditions but the homologies are not sufficiently strong to cause consistent synapsis of any of the chromosomes of the second genomes. Apparently there are two distinctly different second genomes in MM.
1968] MIA & VICKERY: MIMULUS 253
glabtratus var. fremonti. The results bring into question the circumscrip- tion of this entity, but more material and study will be required to clarify the proper limits of M. glabratus var. fremontiu. Unfortunately, tetra- ploid M. glabratus var. fremontii was not crossed with teraploid repre- sentatives of the M. guttatus and M. luteus complexes so the pairing be- havior of the chromosomes of these second genomes was not observed.
The hexaploid level of the M. glabratus complex comprises M. pilo- suisculus and M. glabratus var. parviflorus. In the fertile F, hybrid, M. glabratus var. parviflorus * M. pilosiusculus (6154 5320), all the microsporocytes showed 14 to 16 pairs (table 2). Apparently these two species have only the basic genome in common. Neither the second nor third genomes display any significant amounts of homology. When the F, hybrid, M. pilosiusculus M. glabratus var. fremonti, (5320 X 5373) was studied pairing was not significantly better and the plants failed to set seed. The cross of M. pilosiusculus with the California cul- ture of M. glabratus var. fremonti, 5320 * 5063, failed to yield any hybrids which emphasizes anew the difference between the two forms of M. glabratus var. fremontit.
Chromosome pairing in the F, hybrids of combinations of n = 46 cultures of VM. glabtratus var. parviflorus, specifically, 6184 * 6162 and 6154 X 6163, ranged from 19 to 28 pairs. Apparently all three popula- tions have two almost fully homologous genomes. The results for the F, hybrid 6162 6163 confirm this suggestion (table 2). All these F, hybrids were fertile.
Chromosome pairing in the F, hybrids of combinations of the n = 45 culture, 5041, of M. glabtratus var. parviflorus with n = 46 cultures 6162 and 6163 yielded considerably less pairing than was observed in the intra n — 46 F, hybrids (table 2). Essentially most cells showed pairing of one genome and only a small proportion showed additional pairs. Ap- parently there is a modest amount of homology between either the second or third genomes of culture 5041 and the corresponding chromosomes of the rest of MW. glabratus var. parviflorus. These hybrids were sterile which underlines the difference between this n = 45 culture and the rest of M. glabratus var. parviflorus.
In summary, this investigation reveals that the common genome thought to be present throughout section Szmiolus (Mukherjee and Vick- ery, 1962) has become significantly modified to a greater or lesser extent in each of the four species complexes. The results suggest the presence of at least five distinctly different second genomes, one in M. luteus, the second in tetraploid M. guttatus, the third in the Texas race of MW. glab- ratus var. fremonti, the fourth in the California race of M. glabratus var. fremonti, and the fifth in the various South American populations of M. pilosiusculus and M. glabratus var. parviflorus. Occasionally a few of the chromosomes of the various parviflorus second genomes will synapse, but not consistently in any of the hybrids. The chromosomes of the second genomes of the hexaploids, M. pilosiusculus * M. glabratus var.
254 MADRONO [Vol. 19
AT = eT fy ty 70 LEI ee eine 5048 x 5052 5747 x 5052 :
14 0, 161
09
e 4 e J ner § |
— Se ee Sa \ a e cas Se eA o
ee
5063 x 5747 5373 x 5747 5042 x 5048
14-0,.311 14 0, 321
i 7X - =>
6184 X 5747
22 0, 471
5373 x 5063 5041 X 5373 5041 x 6162
140, 141 140, 141 18 I, 241
6152 x 5048 6250 x 5052 6152 x 5042 Oy O10
1968] MIA & VICKERY: MIMULUS 255
16 0. 60]
27 0. 38]
6184 x 5320 O 5 10 micra
Fic. 2. The original camera lucida drawings were at a magnification of x 2,400. They were reduced to approximately x 1200 in this figure. See Fig. 1 for explanation.
fremontii show the most pairing. Curiously, none of the five hexaploid populations showed any pairing of the chromosomes of the third genomes in their F, hybrids (table 2) yet most of these hybrids were fertile. Possibly these genomes are homologous but the chromosomes separate precociously or, possibly, at this polyploid level the loss or gain of a few chromosomes is not critical to fertility. Apparently, the first genomes of members of section Szmiolus are beginning to diverge significantly, the second genomes have often diverged almost completely, and the third genomes, at least of those tested, seem to have become ntirely distinct.
This investigation was supported by grant no. G 10603 from the Na- tional Science Foundation and by grants from the University of Utah Research Fund. Most of the results form a portion of the dissertation of the senior author submitted to the faculty of the University of Utah in partial fulfillment of the Ph. D. requirements.
The address of the senior author is now: Atomic Energy Center, Dacca, Pakistan.
Department of Biological Sciences, University of Utah, Salt Lake City
Fic. 1. Pairing behavior of the chromosomes in F, hybrids of intra-Mimulus glabratus crosses and of combinations between M. glabratus and its related species. All figures are in or near first metaphase division. Camera lucida drawings were made at an original magnification of * 2,850 reduced to approximately x 945 in this figure.
256 MADRONO [Vol. 19
LITERATURE CITED
MUKHERJEE, B. B., and R. K. VicKEry, Jr. 1962. Chromosome counts in section Simiolus of the genus Mimulus (Scrophulariaceae). V. The chromosomal ho- mologies of M. guttatus and its allied species and varieties. Madrofo 16:141- 155.
VickerY, R. K., Jr., K. W. Croox, D. W. Linnsay, M. M. Mra, and W. Tat. 1968. Chromosome counts in section Simiolus of the genus Mimulus (Scrophularia- ceae). VII. New numbers for M. guttatus, M. cupreus, and M. tillingii. Madrofio 19:211-218.
IDENTITY OF THE OAK TREE AT LIVE OAK TANKS, JOSHUA TREE NATIONAL MONUMENT, CALIFORNIA
JoHN M. TUCKER
Oaks are no rarity in Joshua Tree National Monument. The desert scrub oak, Quercus turbinella Greene ssp. californica Tucker, a shrub or small shrubby tree, is quite common and locally abundant. Two other species, O. dunnii Kell. (Q. palmeri Engelm.) and Q. chrysolepis Liebm., also are found in the Monument, although at only a few locations (Adams, 1957). The former is a shrub; the latter also may be a shrub, but more commonly is arborescent. In the entire Monument, however, there is only one oak tree of any size—a symmetrical 30-foot tree (fig. 1) at Live Oak Tank (fig. 2).
The identity of this tree has long been a matter of conjecture and dis- agreement among Californian botanists. It is a white oak (a member of the subgenus Lepidobalanus), but clearly is not referable to any species recorded for California. This, plus the fact that it is a single, isolated individual of its kind, raises the possibility of its being a hybrid of some sort. In size, degree of lobing, and dentation of lobes, the leaves are very similar to certain hybrids between species with smail, spinose- or mucro- nate-dentate leaves (as one parent), and species with moderately large, deeply lobed leaves (as the other parent); Q. turbinella X Q. gambelu (Tucker, 1961), and Q. dumosa * Q. garryana (Tucker, 1953) come to mind. The leaves of the suspected hybrid have mucronate lobes and teeth. Logically, one parent could have smaller leaves with spinose-dentate margins. This suggests Q. turbinella californica, which occurs with the putative hybrid at Live Oak Tank. As for the other parent, it should logically be another white oak, for no authentic case is thus far known of natural hybridization between different subgenera. If we attempt to extrapolate the characters of the other parent in the manner of Anderson (1949), the detailed reasoning of such an analysis would be that pre- sented in Table I.
1968]
TUCKER: QUERCUS
257,
TABLE I. EXTFAPOLATION OF THE CHARACTERS OF THE SECOND PARENTAL SPECIES
i]
Since Q. turbinella californica
has (is):
Characteristically a shrub
Small leaves (commonly 15- 35 mm long) with very short
petioles
Dentate margins with spinose
teeth
Grayish-green upper leaf surfaces
Thin acorn cups that are scarcely, or not at all,
OF THE PUTATIVE HYBRID.
but the putative hybrid has (is):
the probable second parent has (is):
A fair-sized tree
Larger leaves (40-60 mm long) with longer petioles Moderately lobed leaves, the lobes mucronate Moderate green to vellow-green upper surfaces Thicker cups that are moderately
A large, or moderately large tree
Even larger leaves
(over 60 mm long) with longer petioles
Deeply lobed leaves
with rounded or, at most, obtusely pointed lobes Dark green upper surfaces
Thick, strongly tuberculate cups, that
are hemispheric to deeply bowl-shaped
tuberculate, and are turbinate to sub-hemispheric in shape
tuberculate, and are hemispheric in shape Nuts that are commonly on the Nuts that are mostly Nuts 40 mm or more in order of 20-25 mm in length 30-35 mm in length length
In all the western states only one species fits such a description— Quercus lobata. The leaves, acorns, and cups of the hybrid are compared with those of the putative parents in Figs. 3, 4, and 5, respectively. The nearest trees of Q. lobata, however, are approximately 150 miles to the northwest, along the south side of Antelope Valley in northwestern Los Angeles Co. This poses an obvious problem, which will be discussed later.
In addition to the characters listed in Table I, other significant but less obvious features of the hybrid could be mentioned: 1, the canescence of the acorn cups is very similar to that in Q. lobata, and 2, the stellate hairs of the lower leaf surface are intermediate, most clearly in the size of the lumen of the individual rays (fig. 6).
Another character in which the putative parents differ is the degree of leaf persistence. In Q. turbinella californica the leaves ordinarily persist through the winter. In Q. lobata they are completely deciduous, dropping in the fall or early winter. The hybrid is clearly evergreen and is thus more similar to Q. turbinella in this respect. (James R. Youse and Alan D. Eliason of the Monument staff kindly made observations on this character for me.) Aside from this one conspicuous physiological trait, therefore, the oak at Live Oak Tank is essentially intermediate between the two postulated parental species.
I take pleasure in naming this distinctive hybrid for Philip A. Munz, who in years past has shown considerable interest in its identity.
Quercus munzii Tucker, hybr. nov. Q. lobata Née X OQ. turbinella ssp. californica Tucker. Arbor ca. 9 m alta; ramuli diametro 1.5—2.5 mm griseo- vel brunneo-bubalini, plus minusve tomentosi; gemmae ovoideae,
258 MADRONO [Vol. 19
Fic. 1. The hybrid oak, Quercus * munzii (Q. lobata < Q. turbinella ssp. cali- fornica) at Live Oak Tank, Joshua Tree National Monument, Riverside Co., Cali- fornia.
sparse adpresso-pubescentes vere vel laeve brunneae, 2.5—4 mm longae; folia sempervirentia, 4-6 cm longa, 1.5—3.5 cm lata, ambitu obovata, basi late cuneata vel rotundata vel subtruncata, saepe inaequalia margine irruglariter leviterque lobata, supra vere vel luteovirdia, stellato-pu-
1968] TUCKER: QUERCUS 259
ee ee
San Bernardino Co. | Riverside Co.
TWENTYNINE PALMS
Fic 2.. Joshua Tree National Monument, showing Live Oak Tank, the location of the hybrid.
berulenta, subtus pallide viridia hebitiaque, dense sed minute adpresse stellato-puberulenta, lobis terminaliter 2-pluritaliter mucronate brevi- dentatis, venis utrinsecus 3—5; petioli puberulenti 5—8 mm longi; fructus annuus, cupula hemisphaerica, 13-19 mm lata, 6-9 mm alta, squamae plus minusve tuberculatae pallide brunneo-griseae, canescentes, apice sparsissime canescentes ac pallide brunneae; glans ovoidea demum atro- brunnea glabrataque, 30-35 mm longa, 12—17 mm lata.
Thee ca. 9 m tall with round, symmetrical crown; trunk ca. 7.5 dm in diameter at 6 dm above ground. Trunk bark light brownish-gray, flaky, rather soft and corky. Twigs of the current year’s growth 1.5-2.5 mm in diameter, grayish- or brownish-buff, moderately tomentose, with small, inconspicuous, light-colored lenticels. Buds 2.5-4 mm long, ovoid, sparsely appressed-pubescent, medium to light brown in color. Stipules not seen. Leaves evergreen, obovate (sometimes narrowly so) in outline, 4—6 (or 8) cm long, 1.5—-3.5 (4) cm broad, base broadly cuneate to rounded or sub-truncate, often unequal, margin irregularly and shallowly lobed, the lobes 2- several-toothed, the lobes or teeth mucronate; upper surface medium to yellow-green, stellate-puberulent, lower surface pale green and dull, densely but minutely appressed stellate-puberulent; prin- cipal secondary veins 3-5 on a side; petioles 5-8 mm long, puberulent. Staminate catkins 2 cm or more long, rachis tomentulose, perianth glabrate, the margins of the narrow lobes irregularly fimbriate, the small, glabrous, oval anthers slightly exserted. Fruit annual; acorn cups hemis- pheric to bowl-shaped, 13-19 mm broad, 6-9 mm high, scales deltoid, tips of the upper ones tending to be ligulate, bases of the scales moder- ately tuberculate, light brownish-gray, canescent, the tips sparsely so and
260 MADRONO [Vol. 19
Fic. 3. Leaves of Quercus X munzii (middle row: Tucker 3886 and Cavagnaro). Q. lobata (top row: Tucker 3905), and Q. turbinella ssp. californica (bottom row: Tucker 3895 and Cavagnaro). Scale is in cm.
medium brown in color; acorn ovoid, 30-35 mm long, and 12-17 mm broad, dark brown at maturity, glabrate except for the minutely canescent apex.
Specimens examined. Live Oak Tank, 4,000 feet, Joshua Tree National Monument, Riverside Co., California: Tucker 3886 and Cavagnaro (holotype-DAV), Oct. 2, 1965; Bowerman 4100; Munz 11590 (RSA) ;
1968] TUCKER: QUERCUS 261
I HA a
2 3 4 & 8 "i
LO
Fic. 4. Acorns of Quercus X munzi (middle row: Tucker 3886 and Cavagnaro), Q. lobata (top row: Tucker 3905), and Q. turbinella ssp. californica (bottom row: Tucker 3895 and Cavagnaro). Scale is in cm.
Eliason and Youse s. n. (DAV), Jan. 17, 1966; Eliason s. n. (DAV), June 12, 1966.
The parental species are, for the most part, well-separated ecologically. Quercus turbinella californica is the shrubby, gray-leaved oak of the pinyon-juniper association along the western edge of the Mojave Desert,
262 MADRONO [Vol. 19
jmp
KK
Fic. 5. Acorn cups of Quercus X munzii (middle: Tucker 3886 and Cavagnaro), Q. lobata (left: Tucker 3905), and Q. turbinella ssp. californica (right: Tucker 3895 and Cavagnaro). Scale is in cm.
and of the xeric chaparral in the more arid parts of the inner South Coast Ranges. Quercus lobata is the most characteristic tree of interior valleys in California. Old trees with massive trunks and enormously broad, rounded crowns are a familiar sight in many interior parts of the state. In some areas, however, the two species may occur in close proximity to one another. For example, Q. lobata occurs along U. S. Highway 99 about 2 miles northwest of Lebec, Kern Co., and Q. turbinella californica is abundant on the adjacent hillsides. They also occur together along Oak Creek, 14 miles west of Mojave, Kern Co., on the desert slopes of the Tehachapi Mountains. The tree at Live Oak Tank, however, is the only hybrid between them known to me.
The botanist concerned with experimentally testing a hypothesis of hybridity, may either (1) attempt to synthesize the hybrid by crossing the putative parents, or (2) make a progeny test, if seeds are available from the putative hybrid. In the latter procedure, if the seedlings segre- gate morphologically such that some of them resemble one putative parent and some the other, the hypothesis is confirmed.
To the best of my knowledge, no attempt has yet been made to cross the putative parents of 0. & munzi. Progeny tests, however, have been attempted at least twice. In Oct., 1946, Munz collected acorns from this tree. From these, 105 seedlings were eventually planted at the Rancho Santa Ana Botanic Garden (Percy Everett, pers. commun.). Munz ob- served subsequently that the young oaks differed markedly from one an- other. “Some began to grow fast and upright like young trees, others branched at the base and were shorter and more bushy; some were more deciduous than others” (Vanderspek, pers. commun.). Everett stated, however, that only one small plant was still alive at the date of his writing in 1963. On Nov. 22, 1956, ten years after Munz’s collection, acorns were collected from the hybrid by Ralph D. Cornell (Landscape Architect, Los Angeles), and seedlings from these were planted at the Garden Oct. 17, 1957, according to Everett. Specimens were collected for me from the 11 plants still surviving in 1963. These were all more turbinella-like than the parent tree, having generally smaller and more conspicuously spinose leaves than the latter. Thus, they probably repre- sented backcrosses to Q. turbinella californica which, as stated previously,
1968] TUCKER: QUERCUS 263
Fic. 6. Representative stellate hairs of the lower leaf surface of (B) Quercus munsit (Munz 11590), (A) Q. lobata (Tucker 3442), and (C) Q. turbinella ssp. californica (Tucker 1881-11, 2631, 2763). Drawings by Miss Jean Addicott; all x ca. 50.
occurs in the immediate vicinity of the hybrid. Therefore, since none of them showed any more obvious evidence of Q. lobata ancestry than did the hybrid tree itself, the hypothesis that Q. lobata is one of the hybrid’s parents was neither confirmed nor refuted by this progeny test.
In considering the origin of the hybrid, the most likely hypothesis, in my opinion, is that it was produced zm situ from a direct cross between Q. lobata, probably as the pistillate parent, and Q. turbinella californica, as the pollen parent. This sympatric occurrence of the 2 at Live Oak Tank could have been as recent as 150 to 175 years ago, for the hybrid probably is best regarded as an F,, and does not appear to be an excep- tionally old tree.
Another possible, but much less probable, explanation is that the hybrid resulted from long-range pollination. Such an occurrence would have required a series of interrelated conditions including (Cottam, Tucker, and Drobnick, 1959; Tucker and Maze, 1966): 1, an overlap in the
264 MADRONO [Vol. 19
flowering times of Q. lobata at its southeastern limits and Q. turbinella californica at Live Oak Tank; 2, the occurrence, during the flowering period, of wind from the northwest or west sufficiently sustained to carry pollen the 150 or more miles between the two areas; 3, pollen sufficiently resistant to solar radiation to be capable of germinating after a wind- borne journey of several hours (Jaeger, 1961); 4, atmospheric phenomena which would cause some of the pollen to drift down at Live Oak Tank and settle precisely upon the minute stigmas of the flowers of QO. turbinella californica at a time of receptiveness. The probability of all these require- ments having been met appears very low indeed. But even granting the possibility, it seems that such long-range pollination would stand much less chance of effecting fertilization than the pollen of QO. turbinella cali- fornica, which also would have been present, and in far greater abun- dance. Apparently, fertilization of a pistillate flower is better effected by pollen of its own species than by that of a foreign species, in cases where pollen of both is present. Whatever the basic physiological reason, the validity of this assumption may be inferred from the fact that hybrids between distantly related species of Quercus, as in the present instance, are rarely found in areas where both species are abundant. On the con- trary (Palmer, 1948), they usually are found in areas where one species is abundant but the other is scarce, occurring only as infrequent, isolated individuals. Under such circumstances, it can be assumed that the rare species most probably is the female parent. Its chances of being polli- nated by pollen of its own species would be extremely low, except for the limited possibility of self-pollination, whereas its chances of being pol- linated by the abundant species would be very great.
In seeking an explanation to account for the occurrence of Q. lobata so far south and east of its main, continuous distribution, one cannot ignore the possibility of long-range transport, and inadvertent “planting” of an acorn of Q. lobata by Indians. The possibility appears rather re- mote, however. There seems to be no evidence that acorns were an item of barter between tribes of the Great Valley, or adjacent areas were Q. lobata flourished, and those as far east as Joshua Tree Monument (Sample, 1950). The area with which we are concerned, at least the vicinity of Twenty-nine Palms, was evidently territory of the Serrano (Kroeber, 1908, p. 33), a Shoshonean group centered in the San Bernar- dino Mountains. Although the statement has been made that ‘acorns were fairly abundant in the western part of Serrano territory, but the eastern bands got their supply from the western ones, or substituted other foods” (Kroeber, 1925, p. 618), such acorns would hardly have been those of Q. lobata, for this species probably did not occur in Serrano territory.
Kroeber’s statement that “Quercus lobata was the species that the Cahuilla [a tribe of San Gorgonio Pass and the Colorado Desert] had most frequently accessible to them” (1925, p. 695) is evidently erron- eous. From Barrows’ very detailed survey of the plants utilized by the
1968] TUCKER: QUERCUS 265
Cahuilla, it is evident that they relied on species indigenous to their area (Barrows, 1900).
The most plausible hypothesis, in my judgment, is that Q. lobata had migrated southeastward at some period in the past, when climatic condi- tions were moister than at present. Subsequently, with a shift to drier conditions, the species was reduced in numbers, and finally eliminated altogether. First, however, it hybridized with the abundant and more drought-adapted Q. turbinella californica to produce the hybrid tree at Live Oak Tank.
The pluvial periods of the Pleistocene could have provided the oppor- tunity for such a southeastward migration. However, it should be kept in mind that the last such period, the Wisconsin, was followed by a period of several thousand years that was warmer, and possibly drier, than the present. It seems likely that during this Post-Wisconsin Alti- thermal (Antevs, 1955) or Hypsithermal Interval (Deevey and Flint, 1957), the range of Q. lobata would have extended no farther southeastward than it does today, and probably not so far. Thus, its southeastward migration to the area of Live Oak Tank probably occurred after the Hypsithermal, during a period when the climate of southern California was wetter than at present.
Evidence from tree-ring analyses leaves little doubt that there have been marked climatic fluctuations in the western United States within the last several millennia. Schulman (1956) has noted that long-term shifts to wetter or to drier conditions were typical in southern California for a number of centuries preceding the mid-1600’s (although these gave way to swings of much shorter duration after that time). In our present phytogeographic problem, tree-ring evidence may well provide answers to some of the questions. Certainly, a tree-ring analysis of the hybrid would give the approximate date of the initial hybridization. Al- though a sizeable body of dendrochonological data is at hand for the Southwest, comparable information for southern California is, as yet, very scanty. In future years, however, when many more tree-ring chro- nologies have been worked out for southern California such as that by Schulman (1947) for big-cone spruce, we will be nearer a solution to what, for the present, remains an intriguing riddle.
University of California, Davis
LITERATURE CITED
ApaAMs, C. F. 1957. A checklist of plants of Joshua Tree National Monument. Southwestern Monuments Assoc., Globe, Arizona.
ANDERSON, E. 1949. Introgressive hybridization. John Wiley, New York.
ANTEVS, E. 1955. Geologic-climatic dating in the west. Amer. Antiquity 20:317-335.
Barrows, D. P. 1900. The ethno-botany of the Coahuilla Indians of southern Cali- fornia. Univ. Chicago Press.
266 MADRONO [Vol. 19
Cottam, W. P., J. M. Tucker, and R. Drosnick. 1959. Some clues to Great Basin postpluvial climates provided by oak distributions. Ecology 40:361-377. DeEveEY, E. S., and R. F. Fiint. 1957. Postglacial hypsithermal interval. Science 125:182-184. JAEGER, P. 1961. The wonderful life of flowers. E. P. Dutton, New York. Kroeber, A. L. 1908. Ethnography of the Cahuilla Indians. Univ. Calif. Publ. Amer. Archaeol. Ethnol. 8:29-68. . 1925. Handbook of the Indians of California. Smithsonian Bur. Amer. Ethnol. Bull. 78. PALMER, E. J. 1948. Hybrid oaks of North America. J. Arnold Arbor. 29:1—-48. SAMPLE, L. L. 1950. Trade and trails in aboriginal California. Univ. Calif. Archaeol. Surv. Rep. No. 8. SCHULMAN, E. 1947. Tree-ring hydrology in southern California. Univ. Arizona Bull. 18: No. 3. . 1956. Dendroclimatic changes in semiarid America. Univ. Arizona Press. Tucker, J. M. 1953. Two new oak hybrids from California. Madrofio 12:119-127. . 1961. Studies in the Quercus undulata complex. I. A preliminary state- ment. Amer. J. Bot. 48:202-—208. , and J. R. Maze. 1966. Bur Oak (Quercus macrocarpa) in New Mexico. Southw. Naturalist 11:402—405.
NOTES AND NEWS
CONFUSION IN AUTHORSHIP OF STEMODIA VERTICILLATA HASSLER—In 1964 I collected a small, weedy member of the Scrophulariaceae in a pasture at Horneman’s Ranch (“Bella Vista”) about 4 miles north of Bahia Academy, Isla Santa Cruz (Indifatigalbe I.), E] Archipielago de Colon (Galapagos I.), Ecuador.
While trying to identify this material, I found that it keyed to Stemodia verti- cillata (Mill.) Sprague, in Standley’s Flora of Costa Rica (Fieldiana, Bot. 18:1111. 1938). A check of the Gray Herbarium Card Index revealed that this combination had been made twice prior to the publication of Sprague’s paper (Bull. Misc. Inform. 1921:205-212. 1921). In each of the three cases, Erinus verticillatus Mill. (Gard. Dict. ed. 8, no. 5. 1768) was the basionym, so there is no doubt that each author was dealing with the same entity.
In 1909 Hassler (Trab. Mus. Farm. Fac. Ci. Med. Buenos Aires 21:110. 1909) published the transfer of verticillatus from Erinus to Stemodia, displacing the later S. parviflora Ait. Owing to the restricted circulation of this publication, Hassler’s combination was overlooked and Boldringh again made the transfer (Zakfl. Land- bouwstr. Java 165. 1916). In 1921 Sprague, failing to find the earlier papers, pub- lished Stemodia verticillata as a new combination for the third time.
Therefore, this weedy herb, which is wdely distributed in the American tropics and has been introduced elsewhere, together with its applicable synonymy, should be listed as follows:
STEMODIA VERTICILLATA (Mill.) Hassler, Trab. Mus. Farm. Fac. Ci. Med. Buenos Aires 21:110. 1909. Erinus verticillatus Mill., Gard. ed. 8, no. 5. 1768. Capraria humilis Ait., Hort. Kew. ed. 1. 2:354. 1789. Stemodia parviflora Ait., Hort. Kew. ed. 2. 4:52. 1812. Stemodia arenaria H. B. K., Nov. Gen. Spec. 2:357. t. 175. 1817. Conobea pumila Spreng. Nov. Prov. 13. 1819. Stemodia verticillata (Mill.) Bolcr., Zakfl. Landbouwstr. Java 165. 1916. Stemodia verticillata (Mill.) Sprague, Bull. Misc. Inform. 1921:211. 1921—Ira L. Wiccins, Dudley Herbarium, Stanford University.
SEED ECOLOGY OF SEQUOIADENDRON GIGANTEUM N. STARK
Since the discovery of giant sequoia (Sequoiadendron giganteum (Lindl.) Buchh.) over 100 years ago, people have been concerned about ‘“perserving these forest monarchs.” Yet little is known about seed germi- nation in nature. Many visitors to the California groves ask why there are so few young giant sequoias visible. The answers to this question are most important to the survival of the species. Since we cannot preserve indefinitely the mature trees, we must learn more about seed and seedling ecology to encourage reproduction of the species in nature. This study is one of three and is aimed at a better understanding of the reaction of seeds of the species to the environment both in nature and under labora- tory conditions. The other two studies include a field inventory of the rate of reproduction, and the response of seedlings to environmental extremes.
The proper scientific name of the species and its generic designation are still controversial subjects (Buchholz, 1939; Doyle, 1945). The author prefers to use Sequoiadendron giganteum.
DESCRIPTION OF THE HABITAT
Eighty or more groves of S. giganteum are located on the west slope of the central Sierra Nevada of California at elevations of 4,430—7,540 feet (Anon, 1952). A few trees grow naturally above these elevations.
The range of groves extends a distance of 250 miles from the middle fork of the American River in Placer Co. south to Deer Creek in Tulare Co. Although its natural distribution is limited, this species is planted successfully in at least 14 other countries (Stanford, 1958).
Summers in the natural range of the species are warm and dry with almost no precipitation except for occasional light thunderstorms. Most of the annual precipitation (ranging between 24.8 and 35.8 inches) falls as snow beginning in October or November and continuing through April or May. Annual snow accumulation varies from 45-185 inches.
The warmest month of the year is August when daily air temperatures may rise to 40°C, but often drop to 2°C at night. Intense summer solar radiation on exposed sites may heat the soil surface to 65°C while tem- peratures on shaded sites may be as much as 16°C lower.
The winters are cold; air temperatures seldom exceed 10°C during the day and sometimes drop to minus 23°C at night. The average an- nual frost free period is 124 days.
Mountainous terrain with deep valleys and grassy meadows character- ize the range. The main rock formations are granite, schist, diorite, and andesite. Lava caps are common on the higher peaks.
Most of the groves are on unglaciated terrain but some can be found on glacial moraine. The soils are mainly shallow and rocky but include some deep sandy loams. They are weakly to moderately acid and are
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generally well-drained (Zinke and Crocker, 1962).
The groves are not pure stands but contain these main tree species: Abies concolor (Gord. & Glend.) Lindl., Libocedrus decurrens Torr., Pinus jeffreyi Grev. & Balf., P. lambertiana Dougl., P. murrayana Grev. & Balf., P. ponderosa Laws., Sequoiadendron giganteum (Lindl.) Buchh., and Taxus brevifolia Nutt. A checklist of plants found in and around the groves provides a more complete picture of the flora (Stagner, 1951).
Ferns, mosses, herbs, grasses, and brush are common in some groves. Continuous mats of herbaceous and low shrubby vegetation once grew in most groves, but heavy recreational use of some areas has severely reduced the ground cover (Hartesveldt, 1962).
DESCRIPTION, RESULTS, AND DISCUSSION OF TESTS
Each germination experiment used 10 sets of 100 seeds each placed on moist filter paper in Petri dishes, unless otherwise noted. Seeds used in each of the tests came from five different areas, representing the natural range of the species. Since no significant differences occurred among sources by area, data represent averages of the five sources for each test. Light, moisture, and temperature were standardized with only one factor vary- ing at a time as described below. The ability of the seeds to extend the radicle and free the cotyledons in 32 days served as a criterion of success- ful germination. In preliminary tests, the highest total germination oc- curred in reduced light (5,000 f.c.) at temperatures between 10° and 20°C. The tests and results are as follows:
1. Germination of Controls (20,000 seeds). Germination percent in reduced light (3,000 to 5,000 f.c. day, dark night) on moist filter paper at temperatures of 15°C + 1°C) was used as a germination standard for all other tests. Control germination varied between 21.0 and 55.5%. Each test was compared to its companion control run of 10 sets of 100 seeds from the same seed source and year.
2. Germination and Temperatures (27,000 seeds).
A. Fixed Temperatures. One thousand seeds were germinated in an incubator at each of these temperatures: 5°, 10°, 20°, 30°, and 40°C (= 0.5°C) under 7,000 f.c. light day and night. At 10°C, 39.5% of the seeds germinated while at 20°C, 40.9% of the seeds germinated. All other temperatures produced germination below 39% (table 1).
B. Fluctuating Temperatures. One thousand seeds were placed in Petri dishes in the weather station of the Experimental Plot (used for the seedling studies) every 32 days for 3 years to test germination under natural fluctuating temperatures. When the soil was not frozen, 100 seeds sewn in cheesecloth packets were planted in soil at 2 cm depth every 32 days to check germination in the soil environment. Moisture was added to half of the seeds daily except during the winter months when soil moisture was adequate for germination. Soil and air tempera- ture records were checked against germination. Since fixed temperatures do not occur in nature for any appreciable length of time, germination
1968] STARK: SEQUOIADENDRON 269
TABLE 1. GERMINATION PERCENT OF SEED AT FIXED TEMPERATURES— 5,000 SEEDS ToTAL, 500 PER TEST
52¢ 10°C 20°C 302°C 40°C Percent germination 4.1 39.5 40.9 5.6 0 Percent of control germination 7.4 91.6 94.9 12.9 0
under the influence of naturally fluctuating temperatures is the best indicator of the response of seeds to temperatures.
No one month is dependably better for germination since the climate varies slightly from year to year. In general, field moisture and tempera- ture conditions favor germination in April and May, and sometimes in September in the central Sierra Nevada (table 2). October and Novem- ber seeding usually gives good spring germination. Many seeds planted in the open in June, July, or August failed to germinate because of heat damage and rapid drying even when water was added daily. Lack of germination during the winter months is the result of low temperatures and frozen soil.
Absolute temperature limits on germination were not set, but seeds germinated poorly if air temperatures exceeded 33-34°C or fell below —2°C for any appreciable length of time (table 2).
3. Germination and Water (12,000 seeds). To test the influence of water depth and indirectly of oxygen deficiency on germination, one thousand seeds were germinated under each of the following depths of calm water: 0.5, 1.0, 2.0, 3.0, 4.0, 5.0, 10.0, 20.0 cm and under + 100 cm of fllowing water. Seeds were sewn into weighted cheesecloth packets. A similar test was run using one thousand seeds under 30 cm of water aerated by a pump. Water temperatures varied between 11° and 16°C.
The moisture tests were designed to indicate the ability of seeds to germinate under flooding conditions which often occur in the groves in the spring.
Seeds placed under different depths of water were swollen and few began to extend the radicle. None of the seeds under water for 32 or 64 days was able to completely extend the radicle or epicotyl. After 3 months under water, the seeds were covered by fungi and still showed no signs of germinating. Low oxygen tension in the water tests might explain germination failure. But it was not possible to measure directly oxygen availability to seeds under water. Seeds aerated in 30 cm water did not germinate either, although some were able to extend the radicle. No further germination occurred after 3 months of aeration. Eight per- cent of the seeds removed from aerated water after one month and placed on moist filter paper did germinate proving that these seeds were still alive. Comparable seeds from unaerated water tests were unable to germi- nate when placed on moist filter paper. Seeds in aerated water appeared to receive enough oxygen to maintain life for a month, but not enough to germinate.
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TABLE 2. GERMINATION OF SEEDS UNDER NATURAL FLUCTUATING AIR TEMPERATURES (°C)—22,000 SEEDs.
1959 1960 1961
7o : Yo : %o F Germi- Temp. C Germi- Temp. C Germi- Temp. C
nation Max. Min. nation Max. Min. nation Max. Min.
January — — 0.0 13 —20 —- —- -— February 0.0 19 0 0.0 21 —18 — — — March 9.8 24 —7. 6.7 21 —10 -- — — April 12 207 > eee VO EOS er 10 gs ae May 48.5 34 he 2522 34 —l11 13.0 Sih —1 June Got 37 +2. 19.3 Su — 2 10.0 37 +3 July 6.8 35 0 14.0. 35% © Fao eee Smee August 15.4 31 es 14.3 34 — 3 — — — September 38.2 31 —3,. 36.7 26 — 6 — — —- October — a — -- —— = —- = November 20.0 25 —10. 0.0 27 — 3 — — — December 0.0 18 —22. 0.0 19 — 3 —— -- —
1 Failed because of drying. All other tests were kept moist.
There is little chance for S. giganteum to become established in areas inundated for more than a month. In the groves, widespread flooding rarely lasts more than a few days, but swamps, creek margins and de- pressions may remain under water for months.
4. Germination and Light (10,000 seeds). Germination tests used full light (7,000 f.c.), one-half full light (3,500 f.c.), and total darkness from the time of moistening. Light intensities were checked with a G.E. light meter. Temperatures ranged between 14°-16°C. Full light was cre- ated by placing the Petri dishes 1 m from a fluorescent light source. For half full light the same source was shaded to half intensity. For total darkness, the seeds were moistened after being placed in a black box in total darkness. Another test exposed the seed in covered dishes to full sunlight (10,000 f.c.) during the day.
Seeds showed 98 to 100% of control germination under half light (artificial) (table 3). When seeds were placed in full sunlight out-of- doors during the summer, the compound effects of heat to 60°C and rapid drying appeared to prevent germination. The influence of high light intensity in the absence of high heat was not studied. Seed germination is retarded and seedlings are etiolated in dark tests, but germination ranged between 64.6 and 75.7% of control germination.
In nature, S. giganteum seeds germinate on the litter surface or on mineral soil. Tests show that the litter surface resists wetting and dries quickly producing a poor seed bed. Rarely do the light seeds penetrate to the deeper, moist layers of the litter where germination can occur, but seedling emergence is unlikely because of limited reserve food. Mineral soil in full sunlight often dries too quickly for germination unless the seed is partly buried. Field tests and observations show that the best conditions for germination are disturbed mineral soil in moderate shade.
1968] STARK: SEQUOIADENDRON 271
TABLE 3. PERCENT OF CONTROL GERMINATION UNDER VARIED LIGHT CONDITIONS— 10,000 SEEDS
Test No. Total Darkness Half Full Light Full Light 0. fic. 3,500 f.c. 7,000 f.c. Percent of Control 1 SET 100.0 0.0 2 64.6 100.0 0.0 3 foul 98.0 0.0
Etiolation may enable seedlings arising deep in the soil or litter to reach the surface and begin photosynthesis, although etiolated seedlings 10 cm long often fail to recover in full sunlight. The precise location of seeds on the forest floor determines whether germination and growth can occur.
5. Germination and pH (16,000 seeds). Two thousand seeds were germinated in Petri dishes on filter paper moistened with H.SO,; and NaOH solutions of the following pH levels: 2, 3, 4, 5, 6, 7, 8, and 9. The acidity of the solution was checked before moistening the filter paper and after 32 days using a Hellige pH kit. No measurable changes in pH occurred during the 32 days, but minor fluctuations probably did occur.
Soil acidity influences seedlings after germination more than seed germination itself. Acid solutions may be beneficial in scarifying the seeds, although scarification is not necessary to germination. At pH 6, germination was 82.6% of that of the controls while germination dropped to 63.3% of controls at pH 9, and 35.7% at pH 2 (table 4, 10°C). Germination was better at all pH levels at 20°C than at 10°C. Seedlings developed under pH 2, 3, and 4 were dwarfed with abnormal, stunted, reddened roots. Swelling developed in the root and root crown regions. Seedlings from seed germinated at pH 5 were slightly abnormal. All other treatments produced normal seedlings except for pH 9 where seedlings were dwarfed and intensely blue-green. Although pH 6 and 7 favor the best germination, soil pH should not limit germination in the groves which range from pH 5.6 to 7.9. The influence of sulfate ions in the solution cannot be overlooked in this test. It was not possible to separate the influence of the H+ ions from that of the sulfate ions.
Studies by Zinke and Crocker (1962) show that soil pH varies from acid at the base of a mature tree to progressively less acid away from the tree. The influence of pH on germination was tested in this study to determine whether variability in soil pH within the groves could limit germination.
6. Age of Seeds and Germination (4,100 seeds). The seeds tested had been stored for 1, 2, 3, 4, and 20 years, at —10° to +10°C. The 20-year-old seed was from a different source than the younger seeds, but its average germination at l-year was almost identical with that of the 1-year-old seed used in this test. All seeds appeared to be sound, based on cutting a sample lot of seeds and examining the embryo. In these limited tests, seeds decreased in average viability by 32% in 20 years.
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TABLE 4. PERCENT OF CONTROL GERMINATION OF SEEDS MOISTENED BY SOLUTIONS OF DIFFERENT pH VALUES—16,000 SEEDS
Percent of Control
pH Temperature 2 &) 4 5 6 iI 8 9 107€ S007, 61.2 60.8 70.9 82.6 80.6 78.0 63:3 20°C — 90.3 92.8 87.2 101.3 102.6 90.5 80.9
7. Depth of Planting and Germination (1,400 seeds). Seeds were planted from 0.2 to 6.2 cm deep at 0.2 cm intervals in mineral soil in a glassfaced frame, and the maximum seed depth allowing successful emer- gence over a 60-day period was determined.
In mineral soil, seeds can germinate at depths of 6 cm, but beyond about 2.4 to 3.6 cm, few seedlings can reach the surface.
In litter, seeds grow each day into a microenvironment with more light so that they can emerge from greater depths than in soil. Although seedlings endure etiolation to over 10 cm in total darkness, few of these severely etiolated seedlings survive in full light. At 10 cm depth, the litter is usually sufficiently moist for seeds to germinate, but few S. giganteum seeds can penetrate to this depth, and chances of seedling emergence are slim.
Tests with seeds of competing species show that larger seeds can penetrate the litter more effectively and produce seedlings which emerge from depths greater than 3.6 cm. Sequoiadendron giganteum seeds store a limited amount of food. The seeds of competing species store more reserve food than can S. giganteum seeds and can emerge from greater depths in the soil. This inability to germinate on dry surface litter, or to push through deep litter or soil is seriously hampering reproduction of the species which was formerly adapted to periodic fires that reduced litter depths. Abzes concolor and Libocedrus decurrens which have less strict germination requirements are taking over gradually.
Many groves in undisturbed areas have local spots of good S. gigan- teum reproduction usually occurring in openings on slopes and often sites of old windthrows. Microerosion on these slopes tends to wash the seeds into the litter or soil. Many seedlings of all species grow here, but once established, S. giganteum can hold its own. Logged and burned groves normally have abundant reproduction (average of 7.3 young S. giganteum to each parent stump).
8. Chemical Inhibitors (6,000 seeds). The possible existence of chem- ical inhibitors in the litter of S. giganteum and mixed litter of Pinus spp. and Abies concolor was tested using water extracts (litter soaked for 24 or 48 hours) on seeds. No evidence of inhibition was found with the concentrations used.
9. Rodent Damage to Seeds and Food Preference. Six major food preference tests were set in the fall of 1960 and in the spring of 1961 in an area in the northern range of the species to determine whether repro-
1968] STARK: SEQUOIADENDRON 2S
duction failure is the result of seed-eaters destroying S. giganteum seeds. Twenty-four daily trials consisted of groups of 50 seeds, fruits, or cones of common trees. In the early trials, foods were placed in separate piles so that any disturbance was obvious. Later, the foods were mixed in piles so that seed-eaters had to sort through the seeds to locate those preferred. Isolated tests using only S. giganteum seeds were also made. The number of seeds removed were counted each day and the supply renewed.
These trials were not conducted in an area where seed-eaters com- monly have access to S. giganteum seeds. Animals in the groves might react differently from those of adjacent areas. Seed-eaters were identified by periodic observation of feeding stes and trapping. The results were:
Species % of food parcels removed Libocedrus decurrens On Abies magnifica 0.3 Juniperus occidentalis 20 Sequoiadendron giganteum 0) Abies concolor 4.0 Pinus sabiniana 7.0 Quercus kelloggii 351.0 Quercus chrysolepis 36.0 Quercus wislizenit 40.0 Quercus douglasi 62.0 Pinus ponderosa 76.0 Pinus jeffreyi 82.0 Quercus lobata 84.0 Pinus lambertiana 86.0
Not all seeds, cones, and fruits were sound and their removal does not prove that they were eaten. The animals frequently left unsound food parcels of even the most sought-after species.
These results suggest that the small size and low food return of S. giganteum seeds or cones makes them undesirable to seed-eaters who pre- fer pine seed or acorns. In isolated tests using only sound S. giganteum seeds, Peromyscus maniculatus and Eutamias speciosus destroyed a few seeds. Preference for larger food parcels was noted during the fall in the caching period. Cyanocitta stelleri (Steller Jays) remove large quantities of acorns and pine seed, but whether these were eaten or not is unknown.
The most undesirable foods are the resinous seeds of Libocedrus decur- rens, Abies concolor, Abies magnifica, and the cones of Juniperus occi- dentalis. The most desirable food was Pinus lambertiana.
The unattractive nature of S. giganteum seed argues in favor of a rapid spread of this adaptable species, but deep litter in the absence of fire is preventing its spread.
A. Seed Size (2,000 seeds). Exceptionally large sound seeds (8 mm
274 MADRONO [Vol. 19
average length) germinated 153% better than sound seeds of mixed sizes. Soundness of seed was determined by cutting through a sample lot of seeds and determining if the seed appeared to be normal. Small appar- ently sound seeds (under 4 mm average length) germinated only 6.9% as well as the controls.
B. Germination percent of seeds from 20-year-old trrees (194 seeds). None of the mature seeds collected from 20-year-old trees in a plantation near Pinecrest, California, germinated, even after stratification. Some of these seeds were not sound. It appears that the species does not produce viable seed at 20 years of age.
C. Sequoiadendron giganteum cone extract and seed germination (3,000 seeds). The red material containing gallic acid, pyrogallol, and sugars from the cones is credited with improving germination (Fry and White, 1938). Pigment solutions of 10, 25, 50, 75, and 85% by weight were used to moisten seeds in germination tests.
Cone extract in concentrated dosage (85%) retarded seed germination by one month. This concentration produced smaller seedlings than did the others. The slowing of germination and reduced seedling size may result from osmotic effects since the solution used was supersaturated. No concentration of the extract tested increased seedling size or vigor.
D. The influence of fire on litter and germination (6,000 seeds). In burned groves, young seedlings are often abundant. How does litter in- fluence germination? Several hundred grams of litter from the McKinley grove were burned in an oven. Half of the litter was partly burned and the other half was reduced to a fine ash. Germination was tested using partly burned litter and ash dampened with water in Petri dishes. Half of the tests were covered to prevent drying and half were uncovered to the drying effect of air at 10°C. A related experiment using the gravimetric method determined the changes in dry weight and water holding capacity brought about by burning S. giganteum litter.
Sequoiadendron giganteum seeds placed on unburned litter exposed to the drying effect of the air did not germinate. The litter resists wetting and will float for some time on water, but the strong tendency for surface drying of the litter is unfavorable to germination. Litter floated on water but covered to reduce surface drying proved to be a good seedbed with 98.3% of control germination. Results on mineral soil indicate that germination parallels that of the filter paper controls as long as the sur- face is kept moist. Germination on ash was fair (51%) while germination on partially burned litter (covered) was good (98% of control).
Partially burned litter averaged 68% weight loss over unburned litter, but retained an average of 273% more moisture (by weight) than was retained by unburned litter. Burned litter is reduced in volume so that seeds do not have to penetrate deep layers of dry litter. Partly burned litter holds more moisture than unburned litter and seeds landing on the former have a good chance to germinate before it dries. Partial burning appears to increase the water holding capacity of litter by increasing the
1968] STARK: SEQUOIADENDRON 219
number of spaces which can retain and give up water easily. Small hygro- scopic cells which hold water tenaciously are broken into larger spaces which can take up and give up water more easily than can cellulose walls.
The black, heat absorbent, partly-burned litter should increase sur- face temperatures in the shaded, cool groves to levels well-suited to germination. Fire in the groves in years past appears to have aided natural reproduction in several ways (Biswell, 1961).
E. Germination percent of the seeds from 42 groves (12,000 seeds). Cones containing seeds on the ground were collected from 42 of the groves in 1959. One hundred seeds from each grove were tested for germination in Petri dishes, and another 200 from the same groves were planted in March at 1 cm depth in the soil and covered by litter.
The seeds from these 42 groves averaged 22.5% germination proving that the forest floor has the potential of producing many seedlings, but this potential is not being realized.
DISCUSSION
Sequoiadendron giganteum under natural conditions without fire pro- tection is a long-cycle fire-climax species. Under protection from fire, it appears to be a subclimax species leading to a climax stand dominated by Abies concolor. The absence of natural wildfires is threatening the future of the giant sequoia.
The undisturbed S. giganteum groves before the advent of white man were adapted to periodic light fires and interims of 7-50 or more years for starting young growth. These light fires cleaned away dead or dis- eased trees and removed excess litter and debris so that reproduction was possible. Today there are an average of only 1.7 young trees (recent reproduction not yet producing cones) for each mature parent tree in the groves frequented by tourists. This scarcity of reproduction is traceable directly to the accumulation of litter on the forest floor. Where logging has disturbed the litter and exposed bare soil, there are 7.3 young trees to each old parent stump.
The reproductive potential for the species was based on an average of 2,000 cones per tree per year (figuring good and bad cone years) 210 seeds per cone, and an estimated average life span of 3,000 years.
The 1.2 billion seeds estimated per tree in a lifetime is corrected for 66% which are abnormal or are not released from the cones, 25% rodent damage, and 20% average germination. This leaves an estimated 60,000 potential offspring from one tree, more than enough to increase the natural range of the species. But the tree is not increasing its natural range in most places.
We cannot allow destructive wildfires in the Sierra Nevada today be- cause of the threat to human life and property. We must find some means of producing the same conditions which fire used to create if S. giganteum is to flourish in nature. Perhaps gentle surface scarification of the forest floor will help to create new seedbeds. Where litter has
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accumulated to depts of 0.5 to 0.7 m, the only hope appears to be me- chanical removal. Any disturbance of the soil must take into account the shallow root system.
SUMMARY
Seed germination tests covered the influence of fluctuating and fixed temperatures, water, light, pH, age, planting depth, and seed-eaters on seed germination of S. giganteum. Studies aimed at understanding the failure of the species to reproduce abundantly in nature included the influence of seed size, cone extract, and seedbed conditions on the germination of S. giganteum seeds. Results show that:
1. Under fixed temperatures, S. giganteum seeds germinate well at 10° and 20°C (39.5 to 40.9%).
2. Germination under natural fluctuating air temperatures is best between —2° and +34°C, provided soil moisture is adequate.
3. Seeds will germinate on moist surfaces but not under water.
4. Seeds germinate best in reduced light (5,000 f.c.).
5. Germination is influenced slightly by pH, but seedling growth from seeds germinated below pH 5 is abnormal. Healthy seedlings and good growth resulted from seeds germinated at pH 6, 7, and 8.
6. Sequoiadendron giganteum seeds decreased in viability by 32% in 20 years (limited data).
7. Seedlings seldom reach the surface if seeds are germinated below 2.4 to 3.6 cm in the soil, depending on the degree of compaction.
8. No chemical inhibition of germination by litter extracts or cone extract was found.
9. Larger seeds germinate better than small seeds by as much as 146%.
10. Partially burned litter can hold up to 273% more available water by weight than unburned litter and forms a good seedbed.
11. Seed-eaters prefer the seeds of Pinus lambertiana (sugar pine) and the acorns of many oaks to S. giganteum seed.
12. The failure of the trees to spread and expand their present range is the result of deep layers of surface-dry litter which make germination and early survival difficult for trees with such small seeds.
Desert Research Institute, University of Nevada, Reno
LITERATURE CITED
ANON. 1952. The status of Sequoia gigantea in the Sierra Nevada. Calif. Dept. Nat. Resources. Sacramento.
BISWELL, H. H. 1961. The big trees and fire. Natl. Parks Mag. (Apr.)
Bucuuotiz, J. T. 1939. The generic segregation of the Sequoias. Amer. J. Bot. 25:535-538.
Dayton, W. A. 1943. The names of the giant sequoia, a discussion. Leafl. W. Bot. 3:209-219.
Doytz, J. 1945. Naming the redwoods. Nature 155:254-257.
Fry, W., and J. R. Wuirte. 1938. Big Trees. Stanford Univ. Press.
1968] BOYLE & HOLMGREN: AGROPYRON oT
HarTESVELDT, R. J. 1962. The effect of human impact upon Sequoia gigantea and its environment in the Mariposa Grove, Yosemite National Park, California. Ph.D. dissertation (unpublished). Univ. Michigan.
STAGNER, S. 1951. Checklist of plants of Sequoia and Kings Canyon National Parks. Sequoia Nat. Hist. Assoc., Calif.
STANFORD, E. E. 1958. Redwoods away. College of the Pacific, Stockton, Calif.
ZINKE, P. H., and R. L. Crocker, 1962. The influence of giant sequoia on soil properties. Forest Sci. 8:2-11.
A CYTOTAXONOMIC STUDY OF A NATURAL HYBRID BETWEEN AGROPYRON CRISTATUM AND A. SUBSECUNDUM
W.S. Boyle and A. H. HOLMGREN
Reports of interspecific hybrids in the Tribe Triticeae of the Gramineae have become commonplace except for those involving diploid A. cristatum (L). Gaertn. Grass hybrids involving New World and Old World A gro- pyron are few in number (Dewey, 1967; 1964a; 1964b; (1961). The present paper reports an A. cristatum hybrid found by the authors near the United States Forest Service boundary above Mendon, Utah in 1962. Two species of Agropyron were closely associated with the hybrid: A. cristatum and A. subsecundum (Link) Hitchc. Bowden (1965) and some other investigators have accepted the combination A. trachycaulum (Link) Malte for this entity and regard A. subsecundum as an awned variety of A. trachycaulum. No other species of Agropyron was found in the area after a careful search. Elymus glaucus Buckl. and Hordeum jubatum L. were present in the area but not in abundance and not near the hybrid. Specimens of the species and hybrid discussed in this paper are deposited at the Intermountain Herbarium at Utah State University.
Comparative Morphology of Putative Parents and Hybrid. The diploid A. cristatum growing within inches of the hybrid had glumes distorted near the base, curved awns, and blades strongly pilose on the ventral surface. These morpholological characters will usually separate the diploid A. cristatum from the tetraploid, A. desertorum, where the glumes and awns are straight and the ventral surface of the blades glabrous or only slightly pilose. Agropyron subsecundum was typical of the plants found in northern Utah. The single bunch of the hybrid was conspicuous, as it ap- peared to be intermediate in most characters between the suspected parents.
The hybrid plant as it was found on the mountain produced spikes with solitary spikelets at each node of the rachis but clonal material grown in the field nursery had a tendency to produce two spikelets at a node on the lower part of the spike. Agropyron subsecundum and A. trachycaulum often do this when grown under optimal ecological condi- tions. Agropyron cristatum produces a single spikelet at each node under all conditions.
278 MADRONO [Vol. 19
The spikelets of the sterile hybrid were not as crowded and strongly divergent as those of diploid A. cristatum but not as distant and appressed to the rachis as those of A. subsecundum. In these characters and the length of the spike the hybrid was intermediate between the putative parents (fig. 1). The glumes of the hybrid were only slightly distorted at the base and the awns were nearly straight, and the leaf blades were only slightly pilose on the ventral surface.
Cytological Studies. Both presumed parents had normal meiotic divi- sions. A gropyron cristatum, 2n = 14, regularly formed 7; at metaphase I, in general confirming the observations of Dewey (1964a). A. sub- secundum, 2n = 28, is an allotetraploid regularly forming 14, at meta- phase I and the meiotic behavior did not depart significantly from that described by Dewey (1966).
Tbe sporophytic chromosome number of the hybrid is 21, as expected in view of the virtually certain progenitors described above.
It is questionable whether any gene exchange occurs between chromo- somes in the hybrid. Of the 185 plates interpreted at diakinesis and metaphase I, 39 percent had 21 perfectly clear univalents (fig. 1). The remainder possessed a puzzling kind of chromosomal association involving one or two “pairs” of chromosomes. In 23 percent of the plates there were 19 univalents plus two chromosomes connected terminally by a slender matrix-like thread. In 14 percent there were 17 univalents and two ‘‘matrix pairs” present. In 11 percent there were one or two “pairs” of chromosomes associated simply end-to-end with no indication of an unterminalized region that is generally characteristic of rod bivalents in the Triticeae. The possibility of relatively rare formation of a chiasma in one arm cannot, however, be dismissed. The association observed may be of a homoeologous nature.
At telophase I, The univalents were distributed approximately equally to the poles. Frequently the chromatids were not well defined in contrast to normal telophase dyads. Equational division of univalents was rare. Lagging chromosomes were frequent but less numerous than in many hybrids in this tribe. Characteristically, telophase I chromosomes in the Triticeae do not return to the interphasic condition; only a partial relaxa- tion of the coils occurs prior to re-orientation on the metaphase IT plate. Telophase I chromosomes in the present hybrid were an exception to this. Increase in length and decrease in stainability was pronounced.
In most grass hybrids the lagging univalents that fail to become in- cluded within the nucleus at telophase I become spherical, very darkly staining micronuclei in the cytoplasm. This occurs to a limited extent in the present hybrid but in addition there exists another and more striking kind of micronuclei: one or many lagging chromosomes remain outside the nucleus, uncoil, and apparently become surrounded by a membrane. In addition, partially uncoiled chromosomes may be present outside the nucleus not surrounded by a membrane.
1968] BOYLE & HOLMGREN: AGROPYRON 279
\
Me
Fic. 1. Spikes of Agropyron cristatum (left), A. subsecundum (right), and the hybrid X 2/3. Metaphase I in the hybrid, PMC. 211, * 2400.
In the second division the chromosomes divided equationally in an essentially normal fashion and were distributed in approximately equal numbers to the poles. Laggards were often present and these formed typical micronuclei at the conclusion of the division. The unusual type of micronuclei described above for telophase I were not generally found at telophase IT.
A substantial portion of the pollen mother cells never completed meiosis. Most mature anthers contained approximately half the normal number of pollen grains; some were virtually empty.
Many thousands of pollen grains have been observed in control plants during the course of radiobiological studies on this hybrid. All were judged to be sterile on the basis of staining reaction and abnormal, de- formed structure. The hybrid is completely sterile.
Root tip squash preparations disclosed 21 chromosomes. The divisions were entirely normal and the plants of hybrid clones were unusually vigorous and hardy.
Braegger (1966), in a karyotype analysis of this hybrid, reported chromosome lengths ranging from 6.5—11.5 u in a closely graduated series. All chromosomes possess median or sub-median centromeres. Bottino (1965) reported cyto-genetic effects of gamma irradiation on this hybrid.
280 MADRONO [Vol. 19
Hybridization Attempts. Extensive attempts to produce a controlled A. cristatum—A. subsecundum hybrid were made in 1965 and 1966. Two special difficulties attend crossing these species: A. subsecundum flowers in early June and the florets open in the morning hours. A. cristatum flowers in late June and the florets open in the afternoon. The overlap time is short indeed.
In view of the reported self sterility of A. cristatum (Dewey, 1963) synthesis of the hybrid was attempted the first year without emasculation, with this species serving as the female parent. Both hand pollination, and pollinator culms of A. subsecundum sustained in bottles of water were utilized. The spikes were subsequently bagged. In all, 412 spikes of A. cristatum were used, and 212 seeds later obtained. Of these, 125 germi- nated and produced seedlings all of which proved to be selfed A. crista- tum. Obviously, under the conditions of this experiment, some strains of A. cristatum are not self sterile.
In the second year approximately 20 florets in each of 95 spikes of A. subsecundum were emasculated and utilized as female parents. Thirty- three of the emasculated spikes were hand pollinated with pollen ob- tained from early maturing plants of A. cristatum grown in the green- house, and bagged. The remainder were bagged with pollinater culms of A. cristatum sustained in bottles of water in the conventional fashion. Ninety-five seeds were obtained, 51 of which germinated to produce vig- orous seedlings. Root tip squash preparations disclosed that all 51 pos- sessed 28 chromocomes and were not hybrids. It is difficult to see how such a high percentage of selfed A. subsecundum seeds could be obtained with the precautions taken. The possibility of apomictic development is therefore being investigated.
Diploid agropyrons in the Intermountain region are known only in the native A. spicatum and the introduced A. cristatum. This gives strong support that A. cristatum must be one of the parents of the triploid hybrid. Spike length, straight awns and proximity of the A. subsecundum to the hybrid suggests that this is in all probability the second parent.
We were unable to distinguish, with confidence, between chromosomes of A. cristatum and A. subsecundum in pollen mother cell divisions of the hybrid (fig. 1). Braegger (1967) was also unable to do this with mitotic chromosomes in root tip studies. However, he did find that the largest, satellited, and readily identifiable chromosome in the hybrid had a pre- cise counterpart in A. subsecundum. In addition his total chromosome length studies in the hybrid and the presumed parents are closely consist- ent with the suggested origin of the hybrid.
The authors are pleased to acknowledge the many kindnesses of D. R. Dewey in allowing us to use the facilities of his field nursery, and to David Braegger and Keith Archibald for assistance in the field and laboratory.
Department of Botany, Utah State University, Logan
1968] BOCK: WATER HYACINTH 281
LITERATURE CITED
Bortino, P. J. 1965. Radiosensitivity studies on an interspecific grass hybrid. J. Heredity 56:225—228.
Bownpen, W. M. 1965 Cytotaxonomy of the species and interspecific hybrids of the genus Agropyron in Canada and neighboring areas. Canad. J. Bot. 43:1421- 1448.
BraEGGER, D. R. 1967. Karyotype analysis as an index of radiosensitivity of a grass hybrid. M.S. thesis (unpublished). Utah State University.
Dewey, D. R. 1961. Hybrids between Agropyron repens and Agropyron desertorum. J. Heredity. 52:13-21.
. 1963. Self fertility in crested wheat. Crop Sci. (Madison) 3:351-354.
. 1964a. Synthetic hybrids of new world and old world Agropyrons. I. Tetraploid Agropyron spicatum x diploid Agropyron cristatum. Amer. J. Bot. 51:763—769.
. 1964b. Genome analysis of Agropyron repens xX Agropyron cristatum synthetic hybrids. Amer. J. Bot. 51:1062—1065.
. 1966. Synthetic Agropyron-Elymus hybrids. I. Elymus canadensis X Agropyron subsecundum. Amer. J. Bot. 53:87-94.
. 1967. Synthetic hybrids of new world and old world Agropyrons: III. Agropyron repens X tetraploid Agropyron spicatum. Amer. J. Bot. 54:93-98.
THE WATER HYACINTH IN CALIFORNIA JANE H. Bock
Eichhornia crassipes (Mart.) Solms. (Pontederiaceae) commonly is called water hyacinth. The first record of /. crasstpes in California was from a slough near Clarksburg, Yolo Co. (Smith, Sept. 30, 1904, UC). The first published account (Johnson, 1920) recorded it from the Center- ville Bottoms, King River System, Fresno Co. I have found E. crassipes to be naturalized in California in the Kings, Tuolumne, San Joaquin, and Sacramento rivers systems (fig. 1). The northern limit of naturalization for this species in California was found in Babel Slough, about 10 miles northwest of Sacramento. This may be a world-wide northern limit for the weedy distribution of this species. The southern limit in California was in a pond draining from Lake Hodges, Clevenger Canyon, Ramona, San Diego Co. (Franklin & Buckner, Nov. 29, 1967, SD). The distri- bution of water hyacinths in California appears to be expanding since its early sightings in Yolo and Fresno counties. It still is found in these locations as well as in several new places (fig. 1). The distribution map is based upon personal observation and upon herbarium specimens (CAS, DAV, DS, Fresno State College, GH, LA, ND, NY, POM, RSA, SD, UC, UCR, UCSB, and US).
Unfortunately, no record exists of the progressive spread of E. crassipes in California, so that we must surmise how it occurred. Almost certainly, the species was introduced into the state by man as an ornamental. Subse- quently, plants were put into California waterways. Plants from the Kings River system could have spread to the San Joaquin because of their proximity north of Fresno. And it is not difficult to imagine plants moving
282 MADRONO | Vol. 19
DISTRIBUTION MAP
MUSEUM OF VERTEBRATE ZOOLOGY 9 = ay ae SS aaa | \ ZO | | ae
UNIVERSITY OF CALIFORNIA Ss
SCALE 5 5
) 0 1 ie a a) ' > Ta | 0 25 ) 5 o Li J et ————t . pees ] (es | | |
MILES
Fic. 1. Distribution of Ezchhornia crassipes in California; present, solid circles; past, open circles.
between the San Joaquin and Tuolumne systems, since they meet nine miles west of Modesto. In addition, the Sacramento and San Joaquin systems meet north of Stockton. There is still more room for expansion available. For example, water hyacinths may move into the Stanislaus River and into new regions of the San Joaquin-Sacramento Delta in future years. In some areas, such as the Santa Ana River System, San Bernardino Co., water hyacinths are no longer found because of drought and drastic changes in the natural water supply which have accompanied the rapid increase in human population.
The water hyacinth, a native of the American tropics, owes its success in California to its extremely rapid rate of vegetative growth and repro- duction and its wide range of environmental tolerances (Bock, 1966).
1968] SPELLENBERG: ORYZOPSIS 283
Each winter, thousands of plants are killed, especially in northern Cali- fornia, by the winter cold. However, enough survive through each winter to the following spring to serve as colonizers for the summer’s growth, and to increase distribution. In spite of this natural seasonal control, water hyacinths should be watched carefully both by governmental water weed experts and by interested botanists.
This paper is a portion of a thesis for the Ph. D. degree at the Uni- versity of California, Berkeley. The guidance of H. G. Baker and the assistance of my husband, Carl, are gratefully acknowledged.
Biology Department, University of Colorado, Boulder
LITERATURE CITED
Bock, J. 1966. An ecological study Eichhornia crassipes with special emphasis on its reproductive biology. Ph.D. thesis, University of California, Berkeley. Jounson, E. 1920. Water hyacinth a menace to navigation in the South. Monthly
Bull. Calif. State Dept. Agr. 9:76—80.
NOTES ON ORYZOPSIS HENDERSONIT (GRAMINEAE) RICHARD SPELLENBERG
Prior to 1966 the grass Oryzopsis hendersonu Vasey was known from only four collections. Three of these were from the vicinity of Mt. Clements in Yakima Co., Washington (Mt. Clements, Ff. L. Henderson 2249, 1892; Oak Ridge Rd., D. B. Gray, 1949; Oak Creek Game Range, D. B. Gray & C. F. Martinsen, 1950), the other from the Ochoco Na- tional Forest, Oregon (Trout Creek Basin, D. C. Ingram, 1916). In June, 1966, I discovered a large population north of Colockum Pass in NE Kittitas Co., Washington, extending the known range of the species northward 40 miles. Three more populations were found later, two in the early spring of 1967, along the north side of the Kittitas Valley in the vicinity of Ellensburg. These four new records, listed by my collec- tion numbers, are as follows: 1450, on the Ellensburg—Wenatchee Rd. N of Clockum Pass, ca. 5 miles S of the Kittitas Co. line; 1465, 1466, 1467, stations one half mile apart along Trail Creek ca. 15 miles NE of Ellensburg; 1642, 1662, along Naneum Creek 9 miles N of Ellensburg; 1645, ridge top N of Hwy. 131, 3 miles E of Virden.
The first of these populations grew at 4,300 ft. elev., on a broad ridge top, in a nearly barren opening in mixed ponderosa pine—Douglas fir forest. The oryzopsis formed about 50% of the sparse vegetation in some portions of this area; the most common species with it was Poa secunda Presl. The other collections were made in similar habitats along the lower edge of the pine zone at about 3,000 ft. elev. Each population was found on rocky basaltic soil and occupied what appeared to be the driest site in the area.
The plants form low, dense, gray-green tufts up to 20 cm across and
284 MADRONO [Vol. 19
are easily recognized in the field by this habit, even in the vegetative state. Previous descriptions indicate that the panicle branches are ascending or appressed. This is true of the upper ones, but as the culm continues to elongate during and after anthesis the lower branches may spread to an angle of ca. 45—80° to the main axis. This was observed in two populations in the field and in all plants that were taken from the field and grown at the University of Washington.
Cytological material was collected in the spring of 1967 from plants grown in Seattle. Root tips were treated and sectioned by the method Johnson (1945) used so that the karyotypes obtained would be com- parable to those he studied in his cytotaxonomic treatment of the genus. Also, emerging panicles were fixed in Newcomer’s fixative (Newcomer, 1953), rinsed in 70% alcohol, and stained with Snow’s carmine (Snow, 1963). Permanent slides are filed with the voucher specimens at the University of Washington. From the root tips of plants from collections 1450 and 1465 I obtained counts of 2n = 34, a number previously un- reported for the genus; two of the chromosomes bear satellites (fig. 1). Counts of n = 17, made at various stages of meiosis in developing anthers, were determined in plants from each of the populations I found. No meiotic irregularities were observed and pollen from plants grown in Seattle was about 95% stainable with cotton blue in lactophenol.
It is generally recognized that Oryzopsis and Stipa are closely related, for each genus has some species with features that are used by taxonom- ists to characterize the other. In O. hendersonii, the presence of three, rather than two, lodicules, and the moderately differentiated callus may be considered stipoid features, as might also the convolute lemma, com- mon in Stipa, but known elsewhere in Oryzopsis only in O. asperifolia Michx. Because of these morphological similarities to Stzpa, and the chromosome number which is found in several species of that genus, the embryo was examined to see with which genus it correlated. Reeder (1957) pointed out that the embryo of Oryzopsis has a small epiblast, that of Stipa a large one that usually reaches the tip of the coleoptile, and that “the primary root is often bent at a rather sharp angle to the main axis of the embryo,” a feature especially conspicuous in Sézpa. I was able to confirm these conditions for S. lemmoni (Vasey) Scribn. and O. exigua Thurb. following the methods Reeder described. The embryo of O. hendersonii (fig. 1) compares well with that of the former species but not that of the latter, appearing distinctly stipoid. This might be considered as an indication of affinity to Stipa, but it should be noted that these alleged generic differences in embryology also break down in other species of Oryzopsis. For example, I observed the primary root of O. miliacea (L.) Benth. & Hook. to arise at nearly 90° to the axis; Bruns (1892) illustrated a large epiblast on Piptatherum (—Ory- zopis) paradoxum L.; and Kennedy (1899) illustrated the embryo of Eriocoma cuspidata Nutt. (O. hymenoides (Roem. & Schult.) Ricker) as having an epiblast extending nearly to the tip of the coleoptile, the
1968] SPELLENBERG: ORYZOPSIS 285
epiblast
Fic. 1. Metaphase chromosomes from a root tip cell drawn with the aid of a camera lucida (left). Median longitudinal, section of embryo (right).
primary root arising at a sharp angle to the axis.
In his cytotaxonomic study of Oryzopsis, Johnson (1945) followed the conventional treatment in recognizing three sections, Euoryzopsis, Piptatherum, and Eriocoma. He found Euoryzopsis and Piptatherum to consist mostly of diploid species, those of the first section having 2n = 22 chromosomes, those of the second, 2n = 24. Two species, O. asperifolia and O. racemosa (J. E. Smith) Ricker, were found to have a somatic complement of 46 chromosomes, and since they combine morphological features of both sections, he suggested that they were intersectional alloploids. In absence of cytological data, he discussed O. hendersonii with these two species because it has a similar combination of character- istics, indicating possible intermediacy between the two sections. AI- though I have found that its chromosomes are similar in length to those of the other two species and to the stipoid species in Euoryzopsis, it is unlikely that its complement (34) originated in the same way, followed necessarily by a loss of 12 chromosomes. However, if a basic number of 6 chromsomes, along with 11 and 12, is assumed in the early history of
286 MADRONO [Vol. 19
Stipa and Oryzopsis, as has been suggested by Stebbins and Love (1941) and by Johnson, this complement may have arisen by alloploidy in one of the several ways they proposed for other members of these genera.
While having some features found in Stipa, O. hendersonii (sect. Euoryzopsis) shows possible affinities to section Piptatherum by the partly spreading panicle branches with distally placed spikelets, the wide, several-nerved glumes, and the dark, glabrous, dorsiventrally compressed, indurate lemma. Furthermore, the style branches, which are reflexed out of the side of the open floret at anthesis, resemble those of species in Piptatherum and also those of some species of Stipa, e.g., S. lemmonit. Johnson said that the form of the style branches (reflexed in Prptatherum but exserted from the tip of the lemma in Euoryzopsis) separates the two sections. Lacking critical material of O. hendersonii, he placed it in Euoryzopsis. Such disposition now appears to make that section some- what unnatural, but the species does not fit well in any other subdivision of the genus either. Although it seems to occupy a fairly isolated position, I believe that removing it from Euoryzopsis would necessitate other sub- generic rearrangements. However, before such changes are made, further study of relationships within Oryzopsis is needed.
In summary, the information presented allows some speculation on the ancestry of O. hendersoni?. Several morphological features indicate affini- ties to Stipa, but others reflect a relationship to Oryzopsis (sect. Pipta- therum). Although native species in that section are not presently found in western U. S., Elias (1942) reported a fossil species of Paleoeriocoma from the Tertiary flora of Colorado which bears some resemblance to species in Piptatherum. The possibility that O. hendersonii is an allploid that originated from an ancient hybridization between Stipa-like and Piptatherum-like grasses is plausible if a basic number of 6, in addition to 11 and 12, is assumed for one or the other of these genera.
Department of Botany, University of Washington, Seattle Present address: Department of Botany, New Mexico State University, Las Cruces
LITERATURE CITED
Bruns, E. 1892. Der Grasembryo. Flora 76:1-33.
Exias, M. K. 1942. Tertiary prairie grasses and other herbs from the high plains. Special Pap. Geol. Soc. Amer. No. 41.
Jounson, B. L. 1945. Cyto-taxonomic studies in Oryzopsis. Bot. Gaz. (Crawfords- ville) 107:1-32.
Kennepy, P. P. 1899. The structure of the caryopsis of grasses with reference to their morphology and classification. U.'S.D.A. Div. Agrostol. Bull. 19:1-44.
Newcomer, E. H. 1953. A new cytological and histological fixing fluid. Science 118:161.
REEDER, J. R. 1957. The embryo in grass systematics. Amer. J. Bot. 44:756-768.
Snow, R. 1963. Alcoholic hydrochloric acid-carmine as a stain for chromosomes in squash preparations. Stain Technol. 38:9-13.
Sreppins, G. L., and R. M. Love. 1941. A cytological study of California forage grasses. Amer. J. Bot. 28:371-382.
1968] REVIEWS 287
REVIEWS
Flora of Turkey and the East Aegean Islands, Vol. II. By P. H. Davis. xii + 581 pp. Edinburgh University Press; Aldine Publishing Co., Chicago. 1967. $33.50.
Turkey, a vast country of some 300,000 square miles, has an exceptionally rich flora of vascular plants amounting to perhaps 8,000 species. This figure is consistent with the physiographic diversity of the area and with its crucial position at the meeting place of three vast phytogeographical regions, the Euro-Siberian, Mediter- ranean, and Irano-Turanian. About a fifth of the species of vascular plants found in Turkey appear to be endemic, with exceptionally rich concentrations in southern and eastern Anatolia. The only earlier comprehensive flora which includes the plants of Turkey is Boissier’s six-volume Flora Orientalis (1867-1888). Vastly more informa- tion is of course available about the plants of Turkey at present, with the extensive collections of Peter Davis and his collaborators being of special importance.
The first volume of the present flora appeared in 1965, and thus the second con- forms to the biennial schedule that is contemplated for the eight volumes that will be necessary to complete the flora. The flora includes keys and descriptions, essen- tial synonymy and references, critical taxonomic notes and illustrations, selected distribution maps, citation of types, ecological and phenological information, and general distribution in Turkey.
Volume II includes a list of the major Turkish collections since the completion of the Flora Orientalis (i.e., 1888). In this volume are treated such familiar groups as Centrospermae (Caryophyllales, Chenopodiales), Malvaceae, Linaceae, Geran- iaaceae, Rhamnaceae, and Anacardiaceae. This volume includes families 12-44 of the overall work, and includes such large genera as Silene (119 spp), Hypericum (69 spp.), and Dianthus (67 spp.). The Caryophyllaceae are the largest family treated, occupying 230 pages—well over a third of the volume. The treatment of genera in this family is of particular interest, since what has traditionally been re- garded as Arenaria by North American botanists is broken up on persuasive grounds into Arenaria s. str., Minuartia, Lepyrodiclis, and Moehringia. Melandrium is re- garded as a synonym of Silene but Lychnis is recognized as a distinct genus.
California botanists also will be interested in the division of the Salsola kali L. complex by Paul Aellen into S. kali and S. ruthenica Iljin (including var. tenuzfolia Tausch.), but applicability of this taxonomic treatment to introduced populations in North America remains to be seen. Arthrocnemum is regarded as a genus of peren- nial plants distinct from the wholly annual Salicornia. Frangula is recognized as dis- tinct from Rhajiinus, and the relatively narrow generic standards thus conform closely to those of Flora Europaea. Perhaps it is now possible to achieve some meas- ure of uniformity on the often arbitrary problems of the subdivision of genera, and it is to be hoped that the authors of subsequent floras will depart from the reason- able standards of Flora Europaea only when the preponderance of evidence is over- whelming.—PETER H. Raven, Department of Biological Sciences, Stanford Uni- versity.
A Handbook of Coniferae and Ginkgoaceae. By W. DALLIMORE and A. B. JACK- son. Revised by S. G. Harrison. Fourth Edition. xix + 729 pp. William Clowes and Sons Ltd., London ard Beccles; and St. Martin’s Press, New York, 1967. $35.00.
The fourth edition of this book, which has been a standard reference work since 1923, exhibits a number of departures from the arrangement and treatment in the previous editions. Some parts are the same as those in the 1954 third edition; for example, the key to the sections is essentially the same, the only difference being the use of the term “male strobili” instead of “staminate flowers”; the keys to the genera are virtually the same and are trichotomous in several places; most of the figures are the same as those used in earlier editions.
In contrast, improvements have been made which clearly show the influence of Dr. Harrison. The order of the listing of taxonomic entities is strictly alphabetical
288 MADRONO [Vol. 19
by genera, regardless of the family affiliation. This makes it easier to find an ac- count covering a particular genus without referring to the index, and since the species recognized within genera are also listed alphabetically, it is now quite easy to find the textual coverage of any species. A possible fault in such an order may be that it precludes grouping closely related species or genera together, and one must skip around through many pages to consult the full description of the members of any of the larger families.
Another innovation is apparent in the location of the date of publication appli- cable to the references in the section on “Principal Works Consulted” immediately following the name of the author(s) instead of at the end of the citation.
In several cases the arrangement of back-to-back plates now brings related photo- graphs together instead of having them widely separated as was the case in earlier editions. Thus, plates 37 and 38, following p. 524, bear two photographs of Pseudo- tsuga menziesit on opposite faces of the inserted plate, and the plates between 556 and 557 (plates 39 and 40) represent Sequoia sempervirens and Sequoiadendron gigantea respectively.
The number of named cultivars recognized and included in the text is increased over that of earlier editions, although doubt is sometimes expressed as to the correct systematic alignment of some of them.
The descriptions, as in earlier issues, is reasonably full, and largely non-technical. Economic importance of various species is well covered. Notes about climatic re- quirements, methods of propagation, characteristics of the wood, soil preferences, susceptibility to attacks by insect and fungous pests, and general suitability for garden or park plantings will be helpful to gardeners, whether they be professionals or amateurs. Added notes on age at which various species normally begin to pro- duce seed, approximate number of seeds required to weigh one pound, reactions of the young and mature plants to smog and other industrial wastes often contain information found in few other places or only in scattered reports and papers.
The 4th edition contains 46 plates, 131 line drawings, and 118 references in the “Principal Works Consulted” as compared with 39 plates, 120 figures, and &3 ref- erences in the 3rd edition. The new line drawings are credited to Mr. H. R. W. Herbert, but there is no identifying mark or initials to indicate his work.
Typographical errors seem to be very few. Probably the most noticeable one occurs on page 15 where both the ordinal and family names are correctly spelled, but the genus Ginkgo in the generic heading is spelled “Gingko.”
The index is a model of completeness and very careful work. Generic, specific, varietal, and common names are listed, as well as a huge number of cultivars.
The quality of the paper used in good, the printing excellent and clean-cut. The green buckram binding appears to be a bit light weight for a book of its size and may fail to hold up if the book is handled extensively—as many copies most surely will be. The authors’ names, the title, and the American publisher’s name appear on the spine in silver letters and give an attractive appearance to a shelved volume. This edition beyond doubt will be a standard reference work for years to come and should be received with enthusiasm and praise—IrA L. Wiccrns, Dudley Herbarium, Stanford University.
NOTES AND NEWS
Jepson’s Fiora OF CALIFORNIA.—AIIl published numbers of W. L. Jepson’s “Flora of California” are still available at the original prices from the ASUC Store, University of California, Berkeley, California 94720. Such volumes rarely are listed in booksellers’ catalogues, suggesting that it is not generally known that they are still available new. Although the Flora is incomplete, the parts published are very useful for amateur or professional botanists working in the western United States — ROBERT ORNDUFF, Director, Jepson Herbarium and Library.
VOLUME 19, NUMBER 8 OCTOBER, 1968
Contents
NEw SPECIES OF ERIOGONUM FROM UTAH, James L. Reveal . 289
THE EFFECT OF MISSING DATA AND OF Two SOURCES OF CHARACTER VALUES ON A PHENETIC STUDY OF THE WILLOWS OF CALIFORNIA, Theodore J. Crovello 301
NOTES AND NEws: THE SOUTHERN LIMIT OF TAXUS BREVIFOLIA IN THE SIERRA NEVADA, CALIFORNIA P.W. Rundel 300
INDEX 315
A WEST AMERICAN JOURNAL OF BOTANY
UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY
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BOARD OF EDITORS
Lyman BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis Joun F. Davinson, University of Nebraska, Lincoln WALLACE R. ERNST, Smithsonian Institution, Washington, D.C. ARrTURO GOMEZ PomPA, Universidad Nacional Autonoma de México EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia Mivprep E. Maruras, University of California, Los Angeles ROBERT OrRNDUFF, University of California, Berkeley Marion OwnBEY, Washington State University, Pullman REED C. Rotiins, Harvard University, Cambridge, Massachusetts Ira L. Wiccins, Stanford University, Stanford, California
Editor—JoHN H. THOMAS Dudley Herbarium, Stanford University, Stanford, California 94305
Business Manager and Treasurer—J UNE McCAsKILL P.O. Box 23, Davis, California 95616
CALIFORNIA BOTANICAL SOCIETY, INC.
President: Elizabeth McClintock, California Academy of Sciences, San Fran- cisco. First Vice-President: Peter H. Raven, Department of Biological Sciences, Stanford University. Second Vice-President: Cornelius H. Muller, Department of Biological Sciences, University of California, Santa Barbara. Recording Secretary: John West, Department of Botany, University of California, Berkeley. Correspond- ing Secretary: Mary E. Ashton, Department of Botany, University of California, Berkeley. Treasurer: June McCaskill, Department of Botany, University of Cali- fornia, Davis.
The Council of the California Botanical Society consists of the officers listed above plus the immediate past President, Carl W. Sharsmith, Department of Bio- logical Sciences, San Jose State College, San Jose; the Editor or Madrofio; and three elected Council Members: Virginia M. Page, Department of Biological Sci- ences, Stanford University; Annetta Carter, Department of Botany, University of California, Berkeley; and Robert Ornduff, Department of Botany, University of California, Berkeley.
NEW SPECIES OF ERIOGONUM FROM UTAH JAMES L. REVEAL
During the summer months since 1959 extensive field trips have been taken into the Intermountain Region in connection with the Intermoun- tain Flora Project under the direction of Arthur Cronquist of the New York Botanical Garden. Several new and interesting plants have been found (Cronquist 1963a; b; 1964; Holmgren, 1967; Reveal, 1965; 1966a; b; c; 1967; 1968; a; b). This report is another in the series of contributions in which the various undescribed species are proposed.
The author is grateful to Arthur Cronquist for the opportunity to carry out field and herbarium studies on the Intermountain Region and adjacent areas. This work was largely supported by his National Science Foundation grants from 1964 to 1967, and lead to the discovery of the following new species. The assistance of Noel H. Holmgren of the New York Botanical Garden in the field during those years is appreciated. C. V. Morton assisted the author with the preparation of the Latin descriptions. The United States National Herbarium and the Smithson- ian Institution sponsored the author’s Predoctoral Internship program in Washington, D.C., from September 1966 to February 1967 where this paper was initially prepared. The illustrations were prepared by Mrs. Twila Davis Bird, formerly a student at Utah State University, under the direction of Arthur H. Holmgren, Curator of the Intermountain Herbarium. Up to thirty-five isotypes of the proposed taxa have been distributed from Utah State University.
Eriogonum cronquistii Reveal, sp. nov. Planta perennis herbacea im- plexa, 1-3 dm diam.; caules (0.7)1-1.5(2) dm alti, basi per 1-4 cm foliosi; laminae foliorum ellipticae, 0.5-2 cm longae, 4-10 mm latae, margine crenulatae et crispatae, subtus albo-tomentosae, supra subglabrae et virides, petiolis 3-8 mm longis, tomentosis, basi expansa petioli 1—1.5 mm lati; caules glabri, basi excepta, ca. 1 dm longi, bracteis 1 mm longis, extra glabris, intus tomentosis; inflorescentiae subcapitata vel cymosa, 1—7 cm longae, glabrae; involucra turbinata, 3 mm longa, sessilia, extra et intus glabra, 5-lobata, bracteolis filiformibus, 2 mm longis, pedicellis 2.5-3 mm longis, glabris; perianthia alba, (1.5)2-3 mm longa, extra glabra, intus minute glandulosa, segmentis similibus, oblanceolatis; stamina 2.5—4 mm longa, filamentis basi pilosis, antheris 0.3 mm longis; achaenia brunnea, 2—2.5 mm longa.
Herbaceous perennials from matted, highly branched and spreading, woody caudices, plants (0.7) 1—1.5(2) dm high; leaves basal and sheath- ing up the stems 1—4 cm, leaf-blades elliptical with crenulate margins, 0.5-2 cm long, 4-10 mm wide, densely white-tomentose below, sub- glabrate and green above, the petioles 3-8 mm long, tomentose, the ex-
MaproNno, Vol. 19, No. 8, pp. 289-320. January 8, 1969. 289
290 MADRONO LVol. 19
Fic. 1. Eriogonum cronquisti. Habit sketch showing the erect stems and leafy bases with an enlargement of the inflorescence showing the clustered involucres on short branches and the arrangement of the leaves on the base of the stem.
panded petiole bases 1-1.5 mm wide; stems glabrous and glaucous except for the tomentose bases among the leaves, up to 1 dm long; bracts scale- like, ternate, 1 mm long, glabrous without, tomentose within, the acute apices widening to connate bases; inflorescences open cymes up to 7 cm long, glabrous, dichotomously or trichotomously branched, subcapitate when immature; involucres sessile, 3 mm long, turbinate, glabrous within and without, angled with 5 ridges running down from the 5 acute teeth which form lobes 1% the length of the tubes, the teeth with membranace- ous margins, the bractlets linear and thread-like, 2 mm long, the pedicels
1968 | REVEAL: ERIOGONUM 291
2.5-3 mm long, glabrous; perianth white, (1.5)2—3 mm long, glabrous except for minute glands within along the midribs, calyx-segments = equal, the outer segments oblanceolate with rounded apices, the inner segments slightly narrower and shorter; stamens 2.5—4 mm long, the filaments pilose at the base, the anthers 0.3 mm long, reddish; achenes brown, 2—2.5 mm long, the subglobose bases tapering to smooth 3-angled beaks. n = 20.
Specimens examined. Utah, Garfield Co.: Loose decomposed granite talus slopes on the west side of Bull Mt., Henry Mts., Sec 6, T31S, R11E, 8,300 ft., Holmgren & Reveal 3010 (holotype-UTC), Aug. 14, 1967; Bull Mt., Cronquist & Holmgren 9442 (NY). Fig. 1.
This new species resembles FL. batemaniu M. E. Jones in several of its features and especially so in its glabrous cymose inflorescences, elliptic basal leaves and in flower and involucre size. Yet the two species may be distinguished by the crenulate leaf-margins of E. cronquisti and the shorter leaves than those of E. batemanii which tend to be 2—3 cm long. Also E. cronquistu differs in its loosely matted and spreading habitat which is unlike the more compact and not at all spreading habit of E. batemanii. In addition, EF. batemanii is 2-4 dm high while the new species is rarely 2 dm high. In their ecology, the two species differ greatly. £. cronquisti is the only species in the subgenus Fucycla (Nutt.) Kuntze that grows strictly on talus slopes. The type area is a rock slide where the angle of repose is so great that the slightest disturbance can cause the talus to slide. In collecting specimens, the plants were found to have long creeping caudices—apparently a result of the habitat—and an entire plant is often up to two feet down slope from the point of primary root contact with the soil. On the other hand, F. batemanii is restricted to the heavy gumbo clay hills throughout much of the Uinta Basin and Grand Valley south onto the San Rafael Swells and the lower slopes of the Henry Mountains. Eviogonum batemanii never attains the eleva- tions of E. cronquisti which grows in the rock slide from about 8100 feet to about 9250 feet.
It is a pleasure to name this new species in honor of Arthur Cronquist, Senior Curator of the New York Botanical Garden, who brought this plant to my attention.
Eriogonum humivagans Reveal, sp. nov. Herba perennis, 2-3 dm alta; laminae foliorum oblanceolatae, 1.5-3 cm longae, 2—5(7) mm latae, basiles, subtus albo-tomentosae, supra subglabrae et virides, petiolis 5—10 (12) mm longis, tomentosis, basi expansa petioli, 1.5—2 mm lati; caules glabri, ca. 1 dm longi, bracteis 2-3 mm longis, extra glabris, intus to- mentosis; inflorescentiae cymosa, stricta; involucra turbinata, 3-4 mm longae, 2—2.5 mm latae, sessilia, extra glabri, intus sparsi tomentosa, 5-lobata, bracteolis linearis, 3 mm longis, pedicellis 3.5—-5 mm longis, glabris; periantha alba, costa brunnea, 3—-3.5 mm longa, extra glabra, intus minute glandulosa, segmentis similibus, obovatis; stamina 3—5 mm
292 MADRONO [Vol. 19
Fic. 2. Eriogonum humivagans. Habit sketch showing the general aspect of the species and a single enlarged involucre with exserted flowers. longa, filamentis basi pilosis, antheris 0.4—0.5 mm longis; achaenia brun- nea, 2.5—3 mm longa.
Herbaceous perennials from branched woody caudices, plants 2—3 dm high; leaves strictly basal, leaf-blades 1.5-3 cm long, 2—-5(7) mm wide, oblanceolate, densely white-tomentose below, sparsely tomentose and green above, the petioles 5—10(12) mm long, tomentose, the expanding petiole-bases 1.5-2 mm wide, tomentose within and without; stems glabrous except at the base among the leaves, the stems spreading out- ward from the 5 acute teeth which form lobes '% the length of the tubes, bracts scale-like, ternate, 2-3 mm long, tannish-green to brown and glabrous without, arachnoid-pubescent within, the acute apices widening to connate bases; inflorescences open, cymose, somewhat narrow and strict, trichotomously branched, glabrous; involucres sessile, turbinate, 3-4 mm long, 2—2.5 mm wide, glabrous without, sparsely pubescent in the center of the lobes within, angled with 5 distinct ribs running down-
1968 | REVEAL: ERIOGONUM 293
ward from the 5 acute teeth which form lobes % the length of the tubes, the teeth with membranaceous margins, the bractlets linear, 3 mm long, the pedicels 2.5—4 mm long, glabrous; perianth white with reddish-brown midribs and greenish-brown bases, 3-3.5 mm long, glabrous except for minute tack-shaped glands along the midribs within, calyx-segments similar, obovate, connate less than 1/5 its length; stamens 3-5 mm long, the filaments pilose at the base, the anthers 0.4—0.5 mm long, white to red; achenes light brown, 2.5—3 mm long, the subglobose bases tapering to 3-angled, slightly roughened beaks. n = 20.
Specimens examined. Utah, San Juan Co.: On low clay hills and banks, 13.5 mi E of Monticello, Sec 7, T34s, R26E, 6,800 ft, Holmgren & Reveal 3001 (holotype-UTC); 10 mi E of Monticello, Goodman 5999 (UC); 13 mi E of Monticello, Waterfall 15115 (UC, US). Fig. 2.
Eriogonum humivagans is a member of a difficult group of species that has suffered considerable misinterpretation in various manuals and floras, and to some degree, also in Stokes (1936). This species, as well as the following one, are closely related to E. scoparium Small of western Colo- rado. These three species all have similarly shaped leaves, but E. humz- vagans differs in its inflorescence which is more strict, and the branching pattern of the lower stems which are more spreading, and in various technical characteristics and features of the involucres and perianth parts. In many respects, the new species approaches E. nudicaule (Torr.) Small on northern New Mexico, but E. nudicaule has leaves that are considerably longer and narrower than in this group of species. Of the more technical features, E. humivagans may be distinguished by its larger flowers and in its more narrow involucres. The basal spreading pattern which is designated in the species name is not unique, but it is not found in the following new species.
Eriogonum intermontanum Reveal, sp. nov. Herba perennis, 1.5—3 dm alta; lam‘nae foliorum anguste lanceolatae, (2)3—5 cm longae, 2—4 mm latae, subtus albo-tomentosae, supra subglabrae vel floccosae et virides, petiolis 1-2 cm longis, tomentosis, basi expansa petioli 1-1.5 mm lati; caules glabri, 10-15 cm longi, bracteis 1-3 mm longis, extra glabris, intus arachnoido-pubescentis; inflorescentiae cymosa, 2-10 cm longae; involucra turbinato-campanulata, 2.5-3.5(4) mm longae, 2—3(4) mm latae, sessilia, extra glabri, intus sparsi tomentosa, 5—6-lobata, bracteolis linearis, 3-4.5 mm longis, pedicellis 2.5—4 mm longis, glabris; perianthia alba, costa virides vel rubella, 2—3 mm longa, extra glabra, intus minute glandulosa, segmentis similibus, obovatis; stamina 2.5-4 mm _ longa, filament basi pilosis, antheris 0.4-0.5 mm longis; achaenia brunnea, 2.5—3 mm longa.
Herbaceous perennials from branched woody caudices, plants 1.5—-3 dm high; leaves basal or sheathing up the stems less than 1 cm, leaf-blades (2)3-5 cm long, 2-4 mm wide, narrowly lanceolate, densely white- tomentose below, floccose to subglabrous and green above, the petioles 1-2 cm long, tomentose, the expanding petiole-bases 1-1.5 mm wide, tomentose within and without; stems slender, green and glabrous, erect,
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Fic. 3. Eriogonum intermontanum. Habit sketch showing the erect stems and arrangement of the basal leaves with an enlargement of an involucral cluster and exserted flowers, and a single involucre in the forks of the branches subtended by a ternate bract.
10-15 cm long; bracts scale-like, ternate, 1-3 mm long, narrowly tri- angular to linear, glabrous without, arachnoid-pubescent within, widening from acute apices to connate bases; inflorescences open, cymose, 2—10 cm long, trichotomously branched throughout, glabrous; involucres turbi- nate-campanulate, 2.5-3.5(4) mm long, 2—3(4) mm wide, sessile, 2—5 involucres per cluster, rarely only one, glabrous and reddish without, sparsely pubescent along the lobes within, angled with 5 or 6 distinct ribs running down from the acute teeth which form lobes ca. % of the length of the tubes, the bractlets linear, 3-4 mm long, densely hirsutulous with long marginal cells above, becoming short and capitate below, the pedi- cels 2.5—-4 mm long, glabrous; perianth white with greenish, reddish, or reddish-brown midribs and perianth tubes, 2-3 mm long, glabrous except for minute gland-tipped hairs along the midribs within, calyx-segments similar, obovate, connate less than 4 their length; stamens excluded, 2.5-4 mm long, the filaments pilose at the bases, the anthers reddish- purple, 0.4—0.5 mm long; achenes brown, 2.5—3 mm long, the subglobose bases tapering to 3-angled beaks.
1968 | REVEAL: ERIOGONUM 295
Specimens examined. Utah, Grand Co.: About 1.5 mi S of Uintah Co. line at head of Middle Canyon of West Water Creek drainage in Roan Cliffs, near a cattle pen, Sec 33, T1514S, R24E, 8,400 ft, Holm- gren, Reveal, & LaFrance (holotype-UTC), July 27, 1965, distributed from UTC as E. tristichum Small; Roan Cliffs, Despain s. n. (BRY). Fig. 3.
Eriogonum intermontanum is probably most closely related to E. scoparium Small, but keys out to E. tristichum in Harrington’s Manual (1954). However in reviewing type material, EL. tristichum seems to be a synonym of &. nudicaule (Torr.) Small, but considerable field work is necessary to prove this. Nevertheless, the leaves of the new species are shorter than either. From E. scoparium, the new buckwheat differs in its larger stature and size of its various parts, in its more open inflorescence, and by having its involucres clustered, and only rarely singular as is the typical situation in related taxa. Ecologically E. intermontanum differs from these species. Normally members of this section occur on gumbo clay hills at elevations mostly below 7000 feet. However, this species occurs on gravelly loam soils above 8000 feet elevation where none of the related species occur. It is of some interest to note that the perianth and involucres are exceedingly similar to those of EF. panguicense (M. E. Jones) Reveal of the section Capitata Torr. & Gray, of Southwestern Utah. However as the two do not belong to the same section, they are not believed to be closely related.
Eriogonum ephedroides Reveal, sp. nov. Herba erecta, 2—3.5 dm alta; laminae foliorum lanceolatae, 1.5—2.5 cm longae, 2-3 mm latae, basiles, subtus albo-tomentosae, supra subglabrae vel glabrae et virides, petiolis 5-10 mm longis, basi expansa petioli 3—3.5 mm lati, extra glabra, intus tomentosa; caules glabri, basi excepta, ca. 2 dm longi, bracteis 1-4(7) mm longis; infllorescentiae cymosa, 1.5—2.5 dm longae, stricta, glabrae; pedunculi erecti, inferiores 5-15 mm longi, superiores perbreves; involucra turbinata, 2—2.5 mm longa, extra glabra, intus -_ tomentosa, 5-lobata, bracteolis oblanceolatis, 2 mm longis, pedicellis 2-3 mm longis, glabris; perianthia citrea vel flava, 2—2.5 longa, extra glabra, intus minute glandulosa, segmentis similibus, lanceolatis; stamina 1.5—2 mm longa, filamentis basi pilosis, antheris 0.4—0.5 mm longis, oblongis, flavis; achaenia brunnea, 2 mm longa.
Erect herbaceous perennials from gnarled woody branching caudices, 2-3.5 dm high; leaves strictly basal, 1.5-2.5 cm long, 2-3 mm wide, lanceolate, not revolute, densely white-tomentose below, subglabrous to glabrous and green above, the petioles 5—10 mm long, expanding into membranceous petiole-bases 3—3.5 mm wide, glabrous and tan to light brown without, tomentose within; stems glabrous and bright green, up to 2 dm long; bracts scale-like, ternate, 1-4(7) mm long, linear, the acute apices widening to connate bases; inflorescences narrowly cymose, strict, 1.5—2.5 dm long, trichotomously branched at the first node, dicho- tomous or trichotomous above; peduncles erect, stout, 5-15 mm long
296 MADRONO [Vol. 19
Fic. 4. Eriogonum ephedroides. Habit sketch showing the narrowly erect stems and arrangement of the basal leaves with an enlargement of a single involucre with exserted flowers and an enlarged section of the stem at the node showing the ternate bract.
below, becoming subsessile to sessile above; involucres turbinate, 2—2.5 mm long, glabrous without, sparsely pubescent within, the 5 acute lobes 0.5 mm long, the bractlets 2 mm long, oblanceolate, minutely gland- tipped along the margins, the pedicels 2-3 mm long, glabrous; perianth lemon-yellow to yellow, 2—2.5 mm long, glabrous except for minute glands along the midribs within, calyx-segments similar, lanceolate, con-
1968 ] REVEAL: ERIOGONUM 297
nate ca. % their length; stamens 1.5—2 mm long, the filaments pilose at the bases, the anthers 0.4—0.5 mm long, yellow; achenes brown, 2 mm long, the subglobose bases tapering to 3-angled beaks.
Specimens examined. Utah, Uintah Co.: About 10 mi S of Bonanza along Utah highway 45, on broken white shale slopes, S of White River, Sec 20, T10S, R25E, 5,600 ft, Holmgren, Reveal, & LaFrance 2265 (holotype-UTC), July 25, 1965, distributed from UTC as E. brevicaule Nutt. var. leptothecum (S. Stokes) Reveal; White River, Ripley & Barneby 8738 (CAS, NY); N of Dragon, Ripley & Barneby 13161 (CAS, NY, RSA). Colorado, Rio Blanco Co.: 17 mi NW of Rangely, Ripley & Barneby 8741 (NY). Fig. 4.
This new and distinctive species is a member of the EL. brevicaule Nutt. complex, and is probably most closely related to E. viridulum Reveal. The narrow, strict inflorescence led the author (1966b) to con- sider this plant to be E. brevicaule var. leptothecum. However, during the summer of 1966 detailed herbarium studies were carried out on this species which were completed during an Internship at the United States National Herbarium. This new study allowed the author to review con- siderable material in the entire E. brevicaule complex, and this revealed the distinctiveness of E. ephedroides. While the new species does re- semble var. leptothecum which is restricted to the Front Ranges of Colo- rado in stature, it differs in leaf size and shape, and in the arrangement appear racemose, and in the wider and nonrevolute leaves. Both species differs in its unique inflorescence which, upon a casual observation would appear racemose, and in the wider and nonrevolute leaves. Both of species grow in the same general area. However, E. viridulum is found north and west of the known sites of E. ephedroides.
The bright green and glabrous stems are not unique for the genus, yet they make EL. ephedroides stand out against the light gray to white shale slopes upon which it grows. In the field, one notes the loosely compacted, woody, and gnarled root system. From this root, numerous branches arise, and this gives the plant the appearance of an inverted haystack, with the rounded part at the root, and the flat-topped part above.
The name for this new plant is selected for the striking resemblance it has with an associated species, Ephedra torreyana S. Wats., in its bright green stems and lemon-yellow flowers.
ERIOGONUM THOMPSONAE S. Wats., Amer. Naturalist 7:302. 1873. Herbaceous perennials from branched woody caudices, plants 2-4 dm high; leaves basal or sheathing up the stems less than 2 cm, leaf-blades (2)3-4.5(5) cm long, 8-15 mm wide, oblong to oblanceolate or elliptic, densely white-tomentose below, glabrous and green above, the petioles 3-7 cm long, tomentose, the expanding petiole-bases 3-4 mm wide, densely tomentose on both surfaces; stems green and glabrous, erect, 12-25 cm long; bracts scale-like, ternate, 2-5(7) mm _ long, linear, glabrous, the acute apices widening to connate bases; inflorescences open cymes, trichotomously divided at the first node, dichotomous or trichoto-
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1968 ] REVEAL: ERIOGONUM 299
mous above; involucres sessile, turbinate, 2-3(3.5) mm long, 1—1.5(2) mm wide, green and glabrous without, white-glaucous and glabrous with- in, the 5 acute lobes ca. 0.5 mm long, the bractlets narrowly-oblanceolate, 2-3 mm long, the pedicels 3-4 mm long, glabrous; perianth yellow or white, 3-3.5 mm long, glabrous except for microscopic protuberences along the midribs within, calyx-segments similar, oblong to obovate, connate only at the base; stamens 2—4 mm long, the filaments sparsely pilose at the bases, the anthers 0.3-0.5 mm long, yellow; achenes light brown to brown, 2.5—3 mm long, the subglobose bases tapering to rough- ened 3-angled beaks. Fig. 5.
ERIOGONUM THOMPSONAE var. THOMSONAE. Calyx yellow; involucres (2.5)3—-3.5 mm long, 1—1.5 mm wide. n — 20.
Specimens examined. Utah, Kane Co.: Sandstone cliffs near Kanab, A. P. Thompson s. n., 1872 (BRY-photographs and illustrations, GH- holotype, NY!, UC-fragment!, US-fragment!); Kanab, Holmgren & Reveal 2993 (BRY, NY, UTC). Arizona, Mohave Co.: 9 mi W of Pipe Springs, Parker et al. 6242 (CAS, US).
ERIOGONUM THOMPSONAE var. albiflorum Reveal, var. nov. A var. thompsonae periathiis albis differt. Calyx white; involucres 2-3 mm long, 1.5—2 mm wide. n = 20.
Specimens examined. Utah, Washington Co.: 3 mi W of Virgin, on basaltic gravelly soils associated with Larrea, Sec 19, T41S, R12W, 3,700 ft, Holmgren & Reveal 2991 (holotype-UTC), Aug. 11, 1966; 3.7 mi W of Virgin, Dress 4764 (GH).
The new collection of E. thompsonae var. thompsonae from Utah is the first one to have been made since the type was collected by Mrs. Thompson near Kanab in 1872. Except for the Arizona collection cited above, the species has been virtually unknown. With the discovery of what appeared to be a new variety of E. thompsonae, it became important that an attempt be made to relocate this seemingly rare plant. Following the label data, a brief search was made along the sandstone cliffs north of Kanab, but the species was not seen. However, along the hills north- east of Kanab, and near the local baseball diamond, E. thompsonae was found growing abundantly along the lower slopes of the red sandstone cliffs.
The minor distinction of flower color and involucre size which sepa- rates the two varieties are augmented by the distinct edaphic differences seen in the two type localities. At Kanab, var. thompsonae is restricted to the red sandstone, while the var. albiflorum is found on basaltic gravels near Virgin. In the field, the growth habit of var. albiflorum is more spreading than in var. thompsonae which tends to be more erect and not as dense in the crown.
Fic. 5. Eriogonum thompsonae. Habit sketch showing the general aspect of the species.
300 MADRONO [Vol. 19
Department of Botany, Brigham Young University, Provo, Utah
LITERATURE CITED
CronoguistT, A. 1963a. Two new varieties in Machaeranthera. Leafl. W. Bot. 10:11-13. . 1963b. A new variety of Hackelia patens from Oregon. Leafl. W. Bot. 10:39—-40. . 1964. A new variety of Penstemon davidsonii from Nevada and Oregon. Leafl. W. Bot. 10:129-133. Harrincton, H. D. 1954. Manual of the plants of Colorado. Sage Books, Denver. Hortmcren, N. H. 1967. A new species of Primrose from Nevada. Madrono 19:27-79. ReEvEAL, J. L. 1965. A new alpine Eriogonum from Nevada. Leafl. W. Bot. 10:183- 186. . 1966a. On the specific distinction of Eriogonum nutans and collinum. Madrono 18:167-183. . 1966b. Notes on three Utah Eriogonums. Proc. Utah Acad. Sci. 42:287— 292. . 1966c. Notes on Eriogonum—I. Leafl. W. Bot. 10:334-335. . 1967. A new name for a Utah Lepidium. Great Basin Naturalist 27:177- 181. . 1968a. Notes on Eriogonum IV. A revision of the Eriogonum deflexum complex. Brittonia 20:13-33. . 1968b. Notes on Eriogonum V. A revision of the Eriogonum corymbosum complex. Great Basin Naturalist 27:183-229. STOKES, S. G. 1936. The genus Eriogonum. San Francisco.
NOTES AND NEWS
THE SOUTHERN LIMIT OF TAXUS BREVIFOLIA IN THE SIERRA NEVADA, CALIFORNIA —Current manuals describe the southern limit of distribution of Taxus brevifolia Nutt. in the Sierra Nevada as Tulare County. A recent search of herbaria (A, CAS, DS, GH, Herbarium of Fresno State College, JEPS, POM, RSA, UC, US, USFS, and USNA) and field studies have failed to turn up any locality south of Calaveras Co., 150 miles to the north. A collection from Yosemite Valley in 1874 (Lemmon, DS, US) appears to represent an erroneous record in view of the lack of more recent collections from Yosemite and the careless manner in which the Lemmon Herbarium was handled, and thus this record has been disregarded.
Earliest mention of Tulare Co. as the southern limit of 7. brevifolia in the Sierra Nevada goes back to C. S. Sargent’s Silva of North America (10:65. 1896). Twelve years earlier in his Report on the Forests of North America. (Department of the Interior. Washington, D. C. 1884), Sargent had described ths distribution as “Sierra Nevada to about 37°N,” or central Fresno Co. In view of the evidence presently available it seems plausible that these early reports of Taxus from southern Sierran counties are attributable to errors in collection records or field misidentifica- tions, possibly of Torreya californica Torr. The southernmost occurrence of 7. brevfolia in the Sierra Nevada is now considered to be Calaveras Co. (North Grove, Calaveras Big Trees State Park, 4800 ft., Rundel 1887, DUKE) .—P. W. RUNDEL, Department of Botany, Duke University, Durham, North Carolina.
THE EFFECT OF MISSING DATA AND OF TWO SOURCES OF CHARACTER VALUES ON A PHENETIC STUDY OF THE WILLOWS OF CALIFORNIA
THEODORE J. CROVELLO
Numerical taxonomy is still in a phase of expansion and evaluation. New methods continue to appear and more and more taxonomists are using it to estimate phenetic relationships among plant taxa. Gilmartin (1967) provided an extensive bibliography of recent numerical taxo- nomic applications in botany. Sokal and Sneath (1963) review earlier works.
Concurrent with the development of numerical taxonomy, the use of computers in biological collections and floras has increased. Crovello (1967) lists the uses of electronic data processing in biological collec- tions. Some uses do not involve measurements of characters on specimens, but for other purposes the measurements given in floras would be very helpful. Before information in floras can be employed, the reliability of the measurements provided in a floristic treatment and the problem of missing data must be investigated. By reliability of measurements I mean how accurate they are in estimating the parametric values (e.g., the mean and standard deviation) of a certain character in a given taxon. The problem of missing data arises because many floras are written with- cut the prior preparation of a complete taxon by character table. Many floristic taxonomists include values of a character only in those taxa where the characters are diagnostic of it. This is brought about partly because preserved specimens rarely have all stages of the life cycle present.
The purpose of the present study is twofold: 1, to test the effect of missing data on a phenetic study of a group of plants, and 2, to ascertain the reliability of two sources of information about characters in taxo- species, one arising from a floristic treatment (Munz, 1959) and the other from a study of monographic proportions (Crovello, 1966). The latter is assumed to involve more measurements per character per taxo- species than the former. The results should be of value in estimating the reliability of information from floristic studies for estimation of phenetic relationships. This is especially timely in view of the proposed Flora North Amerca Project. A natural byproduct will be further comprehen- sion of the pattern of variation among the willows of California. By this T mean the phenetic relationships among (not within) the taxospecies in the context of the characters used. For example, Fig. 1 indicates relationships among the willows based on 43 characters. This figure is a reflection of the pattern of variation among the taxospecies based on these 43 characters.
301
MADRONO [ Vol. 19
OTU SECTION COPHENETIC VALUE CODE CODE
0:4 0:6 0:8 lOO.
LASAND barat LAEVIG
CAUDAT
GOODIG - 2 HINDSI
EXIGUA MELANO PARKSI
GEYER! > 13 PSCORD > 4 COMUTA EASTWD 5 ORESTR LEMMON > 13 pRuMSB + 9 JEPSON > 10 BREWER > II SITCHS > 10 DELNRT > II SCOULR > 12 ANGLOR > 7 PLANIF > 9 NIVALL > 8 LUTEA
LIGULI
MACKEN 4 TRACY!
LASLEP
HOOKER PIPERI
Fic. 1. Phenogram of analysis using data from Crovello (1966) for 43 characters.
1968 | CROVELLO: SALIX 303
MATERIALS AND METHODS
In his floristic treatment of Salix, Munz (1959) used 57 morphological characters, but eight were invariant (did not distinguish any willow spe- cies) and were omitted. Of the remaining 49 characters, Crovello (1966) could not obtain information on six of them, so both studies had only 43 characters in common.
Six analyses were made. These differed only in the input data that were used in each. They are the following: 1, data from Crovello (1966) on the 43 available characters of the 57 characters used by Munz (1959); 2, data from Crovello (1966) as the previous analysis, but with the same pattern of missing information as found in Munz (1959); 3, data from Munz (1959) on 43 characters that Crovello (1966) also scored; 4, data from Munz (1959) on 49 of the 57 characters used by him; 5, data from Crovello (1966) on 131 characters; 6, data from Crovello (1966) on 202 characters. Analyses 5 and 6 were included to serve as standards with which to compare the results of the first four analyses. Two standards are considered more desirable, since no one result that uses a large num- ber of characters can be taken as depicting the true, overall, phenetic relationships better than another analysis using many characters. With two “‘standards” before him, the reader has some idea of the variability of results even when information is available on over 100 characters. Table 1 lists the characters employed in the first four analyses. Crovello (1966) gives the characters used in the last two analyses. In it he treated floral characters common to both sexes, e.g., ament length, as one char- acter, but in the present study these are treated as two characters.
For analyses 3 and 4, the information contained in the treatment of the genus by Munz (1959) was used exclusively. This includes the in- formation located both in the keys and in the description of each species. The information under each variety was incorporated into the descrip- tion of its species. The other four analyses used information gathered by Crovello (1966). He used herbarium specimens to reinforce his per- sonal collections. From 7 to 15 plants were chosen for each of the 31 taxospecies recognized by Munz (1959) as native to California. Most plants were represented by several herbarium sheets. Crovello (1966) provides a list of specimens used. This list and copies of the taxospecies by character tables, or Basic Data Matrices (BDM/’s) are on file on punched cards in The Herbarium, University of Notre Dame.
Crovello (1966) concluded that Salix coultert is a synonym of Salix sitchensis. As a result, the number of taxospecies analyzed in the present study is 30, one less than the number recognized by Munz. Table 2 lists the 30 taxospecies and their codes. They are grouped into sections ac- cording to the ideas of Schneider (1921).
There exists no one method of numerical taxonomy. The present study used only one because we are interested in the effect of different sources of data and not in the effect of different taximetric methods. Sokal and Sneath (1963) discuss a number of different methods.
304 MADRONO [Vol. 19 TABLE 1. CHARACTERS USED IN THE PRESENT STUDY. THE First 43 WERE USED BY CROVELLO.
1. Plant habit 30. Stigma lobe length
2. Plant height 31. Style length
3. Last year’s twig color 32. Capsule length
4. This year’s twig color 33. Capsule pedicel length
5. Stipules present or absent 34. Stamen number
6. Stipule length 35. Stamen filament divided
7. Stipule shape 36. Stamen filament pubescent
8. Stipule margin 37. Presence or absence of pubescence
9. Petiole length on last year’s trig 10. Petiole glandular 38. Presence or absence of pubescence 11. Blade length on this year’s twig 12. Blade shape 39. Presence or absence of pubescence 13. Blade margin entire on abaxial leaf surface 14. Blade margin glandular 40. Presence or absence of pubescence 15. Blade margin revolute on adaxial leaf surface 16. Blade base shape 41. Presence or absence of pubescence 17. Blade apex shape on adaxial side of floral scale 18. Blade veins prominent below 42. Presence or absence of pubescence 19. Blade abaxial side glaucous on adaxial side of floral scale 20. Blade adaxial side lustre 43. Presence or absence of pubescence 21. Blade width on capsule surface 22. Female ament length 44. Bark texture 23. Male ament length 45. Bark color 24. Female peduncle length 46. Blade color 25. Female peduncle leaf number 47. Time of flowering compared to 26. Female ament dense or lax time of leaf break 27. Female floral scale length 48. Ovary shape 28. Female floral scale shape 49. Flowering period 29. Female floral scale color
The procedure used here was the same for each of the six analyses. For each analysis the raw data appears in the form of a taxospecies by character table. or Basic Data Matrix (BDM). Each character was trans- formed by condensation to remove the effect of weighting due to measure- ment of different characters in different units. For example, leaf length was measured in millimeters and leaf base shape was measured in angles. To give each character equal weight, each value of a certain character in the tables was condensed, i.e., the value of a character in a certain taxo- species was replaced by a value X,;,
Xo; — X; = Ximin Xo a Xo
where X,j; is the condensed value of character X in taxospecies j, Xj; is the original value of character X in taxospecies j and Ximin and Xmax are the minimum and maximum observed values of character X in the BDM.
Next, I calculated the similarity between each pair of Operational Taxonomic Units (OTU’s), here the taxospecies. I used a modification of the distance coefficient introduced by Sokal (1961). Call this modifi-
MADRONO
A WEST AMERICAN JOURNAL OF BOTANY VOLUME XIX
1967-1968
BOARD OF EDITORS EDGAR ANDERSON, Missouri Botanical Garden, St. Louis LYMAN BENSON, Pomona College, Claremont, California KENTON L. CHAMBERS, Oregon State University, Corvallis JOHN F. Davipson, University of Nebraska, Lincoln WALLACE R. ERNsT, Smithsonian Institution, Washington, D.C. ARTURO GOMEZ PomPaA, Universidad Nacional Autonoma de México EMLEN T. LITTELL, Simon Fraser University, Burnaby, British Columbia MivpreD E. MATHIAS, University of California, Los Angeles ROBERT ORNDUFF, University of California, Berkeley MARION OWNBEY, Washington State University, Pullman REED C. ROLLINS, Harvard University, Cambridge, Massachusetts
IrA L. WiccINs, Stanford University, Stanford, California
Editor—JoHN H. THoMAs Dudley Herbarium, Stanford University, Stanford, California 94305
BUSINESS MANAGER AND TREASURER JuNE MCCASKILL P. O. Box 23, Davis, California 95616
Published quarterly by the
California Botanical Society, Inc. Life Sciences Building, University of California, Berkeley 94720
Printed by Gillick Printing, Inc., Berkeley, California 94710
To JoHN THomas HowE Lt, indefatigable collector and student of the plants of California and the Galapagos Islands, and long time member of the California Botanical Society, this volume of Madrono is dedicated.
Tom Howell was born in Merced where he lived until he graduated from high school. It was as a student at the University of California in Berkeley, where he came under the influence of Professor W. L. Jepson, that he developed his interest in botany. Following his university gradu- ation and subsequent completion of a master’s degree, he went to south- ern California, where he spent the first two years of his professional life. In 1929 at the invitation of Miss Alice Eastwood he came to the Cali- fornia Academy of Sciences in San Francisco where he has remained.
As a collector of botanical specimens, Tom Howell has few equals, and his collection numbers have surpassed 40,000. Moreover, his collections have never lain idle to collect dust but he has determined them and in- corporated them in the Herbarium of the Academy. Although most of his collecting has been done in California and the Galapagos Islands he has made important collections in several western states. In addition to his own collections which have added so extensively to the Academy’s herbarium he has encouraged many others, both amateur and _ profes- sional, to make collections which also have come to the Academy.
His numerous botanical papers have contributed greatly to our knowl- edge of the plarits of the areas where he has collected and particularly his floras of several regions in northern and central California have con- tributed to a greater botanical understanding of local areas.
Leaflets of Western Botany, which he and Miss Eastwood began in 1932 and which he edited and published until the end of 1966, provided a vehicle for significant contributions to the botany of the western United States.
His many years of field work, his keen observations, his great enthu- siasm and unlimited energy, have brought to him an unsurpassed knowl- edge of California’s native and weedy plants. And he is always a source of botanical information, willingly, thoughtfully and kindly given, to all who seek him out.
iy
CONTENTS
PAGE Frontispiece: John Thomas Howell A revision of Petalonyx (Loasaceae) with a consideration of affinities in sub-
family Gronovioideae, William S. Davis and Henry J. Thompson ................ 1 MacNab cypress in Northern California: A geographical review, J. R. Griffin MUN OA SIL OIL Ee ca eae cote Faas shat ae a ee ea tes ce tae ee pean ee Meas ON ones, tee 19 A new species of primrose from Nevada, Noel H. Holmgren .......0.......0.0.20000000200---+ 27 TRSCGNATS) Ge: anes A Bees eA eNY erence Inert eran ae een eer en 30, 58, 94, 220, 287 Natural hybridization between Encelia and Geraea (Compositae) and some re- lated experimental investigations, Donald W. KyhoS .......222000....222222.0222222000----+- 33 New records of Myxomycetes from California. II, Donald T. Kowalski................ 43 INGOs pain te INC WS occ cat cece oot ssestee neve sass ovata seces vac entereys 46, 62, 78, 91, 95, 136, 164, 192, 210, 222, 234, 266, 288, 300 Cypress rusts in California and Baja California, Roger S. Peterson ..............2....... 47 Two new eriogonums from Baja California, Mexico, James L. Reveal and Craig Zilog ts EH OSG) {Thee ie ee Baler ee la OPa VOODOO Ae gr et PO ROR ee a 55 Check list of California Pteridophytes, Lawrence L. Kiefer and Barbara Joe...... 65 A new Hesperaloe from Sonora, Mexico, Howard Scott Gentry ...0.000000000000000000... 74 Studies of juniper rusts in the West, Roger S. Peterson ........0.0...00000..220000c0c0vceeeeee- 79 A new species of Pterigoneurum from California, Howard Crum .............0.000..222... 92 The scientific writings of Willis Linn Jepson, Lawrence R. Heckard, John i Nomas dowel, and R20 BOCtgGli pl 22.2 ee De 97 On the Hordeum jubatum-—H. brachyantherum question, Wm. W. Mitchell ........ 108 Notes on marine algae of Washington and southern British Columbia. II, hicnard Norns and J onn Aja est . 22.5. 2o a ee ee stint Gossypium trilobum: An addendum, Paul A. Fryxell and Clifford R. ParkRs........ 117 bofSree of] 8.0) 011 Re perenne ia aieet to r mee oe Rrra NE apie rer eee Pee OR AA CR See coe ee eer te 124 A hybrid hawkweed from the Olympic Mountains of Washington, Arthur R. DSS TUGIC CO CH gu tesco c ee Sate ot cao te Sec Ss ASIA Sg cee eee EES eee eee 126 A survey of the vascular plants above timberline on Mt. Hood, Oregon, Lurel TE Wtf A ea Oa RL ee ee ee Bari Ee EO 130 Documented chromosome numbers of plants -.........22..222220...220-cceee0eeeeeeceeeeeeeeeeeeeeeee 134 Variation in the diploid Gossypium species of Baja California, Lyle L. Phillips EWING ee) TUFEC OIL a LEME IUD 0 teres Tater, Faeroe Sh Ne GE eg a was Mee eee 137 California boletes. III. The genus Suillus, Harry D. Thiers _...000.000000000000000000200-- 148 Chromosome variation in ecotypes of Larrea divaricata in the North American desert, Tien Wei Yang and Charles H. Lowe ........200....2200. 0.220002 ccceeccceeeeeee eee 161 Cytological and evolutionary studies in palustres violets, Marja Sorsa ................ 165 A second species of Ammobroma (Lennoaceae), in Sinaloa, Mexico, Robert L. HO GESSLC TAINLY OO NUH nian se a oe Sieh A aaa Bod eee oe 179 Chromosome numbers in the tribe Pappophoreae (Gramineae), John R. Reeder A OUGIRS DERI ASS 1 (2-4 dilate lene et eles pier a ce RE Se ee AS eee see eee 183 Some additions to the Flora of Texas. IV, Donovan S. Correll .....0.0.0000.00200.0000020--- 187 New taxa, combinations, and chromosome numbers in Arctostaphylos (Eric- ESE AC) A ILILE PW VY CLUS tse peae eee events Se ean Sa ee Pearce gee ate ace Sea ae 193
Chromosome counts in section Simiolus of the genus Mimulos (Scrophulari- aceae). VII. New numbers for M. guttatus, M. cupreus, and M. tilingii, R. K. Vickery, Jr.. K. W. Crook, D. W. Lindsay, M. M. Mia, and W. Tai.... 211
A rust on Anemopsis californica, Paul D. Keener and Jack E. Hampton................ 218
The vegetational composition of the south slope of Mt. Pinos, California, HChOyd » .) Of) and bran C Vier oe ee ae 225
Taxonomy, variation, and chorology of three chromosome races of the Calamag- rostis canadensis complex in Alaska, Wm. W. Mitchell ..........00.0000000.2............ 235
New records of Myxomycetes from California. III, Donald T. Kowalski and I OY ER TY BA @ (Bad 6771 2 1 5 a eT ee Se RO eI PR ae ee OE OR ere ERTS 246
Chromosome counts in section Simiolus of the genus Mimulus (Scrophulari- aceae). VIII. Chromosomal homologies of M. glabratus and its allied spe-
cies and varieties, M. M. Mia and Robert K. Vickery, Jr. .....2....22.00000000000000-- 250 Identity of the oak tree at Live Oak Tank, Joshua Tree National Monument,
Californias. J O01. . LUGRER soe ete ee ae eee 256 Seed ecology of Sequoiadendron giganteum, N. Stark -..................1ccceccceeeeeee eee 267 A crytotaxonomic study of a natural hybrid between Agropyron cristatum and
A. subsecundum, W.S. Boyle and A. Hl. Holmgren 2.3... nse 270i, The water hyacinth! in’ California, Wave a Bock. en ee 281 Notes on Oryzopsis hendersonii (Gramineae), Richard Spellenberg ..............2....... 283 New species of Eriogonum from Utah, James L. Reveal ............000.0.2200cc20cccceeeceeeee ee 289 The effect of missing data and of two sources of character values on a phenetic
study of the willows of California, Theodore J. Crovello......0...00.0.00.000000000000-- 301 TVG sak oes ace 5c ts egg arte es as naa cD SHI)
STATEMENT OF OWNERSHIP, MANAGEMENT, AND CIRCULATION (Act of Oct. 23, 1962; Section 4369, Title 39, United States Code)
Maprono, A West American Journal of Botany, is published quarterly at Berke- ley, California.
The publisher is the California Botanical Society, Inc., Life Sciences Building, University of California, Berkeley, California 94720.
The editor is John H. Thomas, Division of Systematic Biology, Stanford Univer- sity, Stanford, California 94305.
The owner is the California Botanical Society, Inc., Life Sciences Building, Uni- versity of California, Berkeley, California 94720. There are no bondholders, mort- gagees, or other security holders.
The average number of copies distributed of each issue during the preceding 12 months is 650; the numbers of copies of the single issue closest to the filing date is 650.
I certify that the statements made by me above are correct and complete.
Joun H. Tuomas, Editor November 15, 1968
1968 |
TABLE 2. THE 30 TAXOSPECIES OF SALIX IN CALIFORNIA RECOGNIZED BY CROVELLO (1966). ARRANGEMENT BY SECTIONS FOLLOWS SCHNEIDER (1921)!.
Section Code
ite
Pentrandrae Dumort.
. Nigrae Loudon . Longifoliae
Anderss.
. Cordatae Barr.
. Andeno phyllae
Schneid.
. Chrysanthae Koch . Ovalifoliae Rydb.
. Reticulatae Fries . Phylicifoliae
Dumort.
. Sitchenses Bebb
. Brewerianae
Schneid.
. Discolores Barr. . Fulvae Barr.
CROVELLO: SALIX
Taxospecies Taxospecies
Symbol
LASAND CAUDAT LAEVIG GOODIG HINDSI EXIGUA MELANO PARKSI LUTEA LIGULI MACKEN PSCORD LASLEP TRACYI COMUTA EASTWD ORESTR PIPERI HOOKER ANGLOR
NIVALI PLANIF
DRUMSB
SITCHS JEPSON BREWER DELNRT SCOULR LEMMON GEYERI
nn NANNNNNUNUUUANNWHNURUUHUUHHUHAUYN
i)
S S S S. S S
. lasiandra Benth. . caudata (Nutt.) Hell. . laevigata Bebb.
. gooddingii Ball
. hindsiana Benth. . exigua Nutt.
. melano psis Nutt. . parksiana Ball
. lutea Nutt.
. ligulifolia (Ball) Ball
. mackenziana (Hook.) Barr. . pseudocordata Anderss.
. lasiolepis Benth.
. tracyz Ball .commutata Bebb . eastwoodiae Ckll. . orestera Schneid. . piperi Bebb
. hookeriana Barr. . anglorum Cham. var.
antiplasta Schneid.
. nivalis Hook. . planifolia Pursh var. monica (Bebb) Schneid. . drummondiana Barr. var. subcoerulea (Piper) Ball . Sitchensis Sans.
. jepsonti Schneid. . breweri Bebb delnortensis Schneid. . scouleriana Barr. . lemmonii Bebb Ss
geyeriana Anders.
305
1 The only exception to Schneider’s assignments is S. jepsonii. He placed it in section Phylicifoliae.
cation the similarity coefficient, sj. Then,
Njk
>, (Xi — Xin)? | Y
Sj. — 1 — al
Nyx
where X;; and X;, are the values of character i in OTU’s j and k, re- spectively, and nj, is the total number of characters used in the particular comparison. If there were no missing data, i.e., if the OTU by OTU relevance (Sokal and Sneath, 1963) were always 1.0, then n;, would be the same for all combinations of pairs of OTU’s.
306 MADRONO [Vol. 19
TABLE 3. SUMMARY STATISTICS OF THE SIX ANALYSES.
Analysis Cophenetic number and Similarity OTU by OTU correlation descripion coefficient relevance coefficient iY; S Y S 1. Crovello 43 593 .086 .972 1037, 146 2. Crovello 43 with Munz’s missing data pattern 578 .098 482 082 678 3. Munz 43 590 085 495 .079 741 4. Munz 49 587 082 483 075 719 5. Crovello 131 617 077 908 — 032 858 6. Crovello 202 615 072 829 .080 871
Each set of similarity coefficients from one BDM forms a 30 by 30 OTU by OTU table, or Basic Similarity Matrix (BSM). Each cell in it indicates how similar two taxospecies are in the context of the characters used and based on the source of the data for the BDM being analyzed. The BSM was then used to group OTU’s by the unweighted pair-group method (Sokal and Sneath, 1963). This results in a phenogram, a hier- archic presentation of phenetic relationships among the OTU’s, in the context of the characters analyzed.
RESULTS
Figures 1 to 4 and Tables 3 and 4 present the results of the four analyses. Figures 5 and 6 are standards with which to compare the phenograms of the analyses using Munz’s characters. The last two repre- sent the maximum information on California willows available to the author at the present time. Figure 5 is based on the morphological char- acters studied by Crovello (1966) but without the six (or seven) unit pubescence characters scored for each organ. Figure 6 includes the 62 unit pubescence characters. Two standards were used: 1, to increase comprehension of the results of the present paper; and 2, to emphasize that in numerical taxonomy any one result is not the ultimate truth. For ease of presentation of results, we shall compare Figs. 1 through 6 with the latest nonnumerical monograph, which I summarize in Table 2.
Figure 1 gives the analysis of 43 characters using full data from Cro- vello (1966) which produced seven clusters. Beginning at the top, the first cluster contains four taxospecies that are the only representatives in California of the subgenus Pleiandrae. The next four OTP’s are the repre- sentatives of section Longifoliae appearing in California. GEYERI then joins this cluster. The next cluster of five taxospecies includes members of sections Cordatae, Adenophyllae and Fulvae, while the subsequent cluster includes members of sections Phylicifoliae, Sitchenses, Breweri- anae and Discolores in a mixed pattern. This is followed by a cluster consisting of three high-altitude willows, ANGLOR, PLANIF and NI- VALI. The next to last cluster includes four members of section Cordatae, while the last cluster contains LASLEP, a polymorphic member of sec-
1968 ] CROVELLO: SALIX 307
TABLE 4. CORRELATION BETWEEN ALL PAIRS OF THE SIX SIMILARITY MATRICES (LOWER LEFT), AND CORRELATION BETWEEN ALL Pairs OF THE SIX PHENOGRAMS (UPPER RIGHT). IN ALL Cases n = 435.
Analysis number and description Analysis number 1 Z 6) 4 5 6
1. Crovello 43 1.000 852 .606 643 ALS ONT 2. Crovello 43 with Munz’s
mising data pattern 887 1.000 455 437 460 ASS 3. Munz 43 583 450 1.000 947 658 noes: 4. Munz 49 .604 460 974 1.000 739 653 5. Crovello 131 835 .669 655 .670 1.000 889 6. Crovello 202 ies 585 595 .614 889 1.000
tion Cordatae, and the two taxospecies of section Chrysanthae that ap- pear in California.
Turning to Fig. 2, which is based on Crovello’s data but uses the pattern of missing data present in the Munz data, seven clusters also are seen here. The first is similar to that of Fig. 1. The next two clusters of three and four taxospecies resemble two from Fig. 1, but TRACYI is out of place. The next cluster includes section Longifoliae, but here GEYERI has split the four members of that section. The next cluster contains six taxospecies. Except for ANGLOR, it is similar to a cluster in Fig. 1. The final two clusters also have their counterparts in Fig. 1, with the exception of ANGLOR mentioned above.
Figure 3 is based on Munz’s data on the 43 characters that are com- parable to Crovello’s. The first cluster of four taxospecies is the same as in previous figures. But then HINDSI appears as deviant from all other OTU’s. The other three members of section Longifoliae are far re- moved from it. The next two clusters, the first with six taxospecies, bring together quite different OTU’s as suggested by conventional taxonomy and by Figs. 1 and 2. ANGLOR is a dwarf alpine form, whereas SCOULR is a polymorphic shrub or small tree more common at lower altitudes. By inspection of the rest of Fig. 3 the reader can ascertain the similarities and differences of it when compared to the previous figures.
Figure 4 is based on the 49 characters used by Munz. Seven more or less distinct clusters emerge. The four OTU’s of section Longifoliae are closer together now than in Fig. 3, but as in Fig. 2, GEYERI splits them. Here EASTWD also is among this cluster. Note at the bottom of Fig. 4 that ANGLOR is still grouped with SCOULR.
Figure 5 consists of eight clusters, the first three of which agree exactly with Fig. 1. The remainder of Figs. 1 to 4 is in less agreement with Fig. 5 and Fig. 6. Note, however, that Figs. 5 and 6 are not identical.
Table 3 gives summary statistics for the six analyses. Columns 1 and 2 list the mean and standard deviation of each of the Basic Similarity
308 MADRONO [Vol. 19
OTU SECTION COPHENETIC VALUE CODE CODE 0-4 0-6 0:8 1-00 St a. a Ce ts | eee ae | aaa
LASAND |
LUTEA —— ee 4 MACKEN
LASLEP HOOKER > 6 TRACY! & 4 PIPER) > 6 HINDS| =& 3
GEYERI > 13 EXIGUA PARKSI 3 MELANO Pscorp © 4
COMUTA EASTWD :
ORESTR |
ANGLOR & 7 prumsB & 9g JEPSON > 10 BREWER > I! sitcHs + 10 DELNRT + 11 scour & 12 NivaL! & 8
Fic. 2. Phenogram of analysis using data from Crovello (1966) for 43 characters with Munz’s missing data pattern.
1968 |
CROVELLO: SALIX
COPHENETIC VALUE 0-4 0:6 0:8 1-00 Bois: Wee ad
OTU CODE
LASAND CAUDAT LAEVIG GOODIG HINDSI
Oo P) m ” + ps)
LEMMON DRUMSB SITCHS JEPSON GEYERI ANGLOR SCOULR PLANIF BREWER DELNRT
‘LUTEA
MACKEN COMUTA PIPERI HOOKER LIGULI PSCORD LASLEP TRACYI EXIGUA PARKSI MELANO EASTWD NIVALI
SECTION CODE
b 2 eS
A on
ae a ee
ee
Vad
309
" Fic. 3. Phenogram of analysis using data from Munz (1959) for 43 characters.
310 MADRONO [Vol. 19
Matrices (BSM’s). The closeness of the values is expected when each analysis is based on a high OTU by OTU relevance. But as seen from the next two columns the relevance values fluctuate considerably.
The last column of the table gives the cophenetic correlation coeffici- ent for the six phenograms. It is the correlation coefficient obtained from a comparison of the BSM with the cophenetic values of the phenogram resulting from it (Sokal and Sneath, 1963). Since it compares a pheno- gram with the BSM from which it is derived, it provides an accurate estimate of how well the phenogram reflects the relationships given in the BSM. In unpublished studies the author has noted that, in general, as the number of characters increase, the value of the cophenetic cor- relation coefficient increases.
The lower left half of Table 4 contains the Pearson product moment correlation coefficients computed between all pairs of the similarity matrices used in the study. All of the BSM’s are rather highly positively correlated, but note the extremely high values among close pairs, such as between sets 1 and 2 and sets 3 and 4. Examination of a family of such statistics in a group under study allows one to understand values characteristic of similar analyses and those characteristic of more un- related analyses. These correlation coefficients are not to be confused with the cophenetic correlation coefficient, the values of which are found in Table 3.
Comparison of two phenograms can be made more efficient and ob- jective by computing the coefficient of correlation between the cophenetic values obtained from the two analyses. This was done in the present study and results appear in the upper right half of Table 4. We can tell at a glance how similar are the relationships among OTU’s in any two pheno- grams.
DISCUSSION
The first purpose of this study was to test the effect of missing data on a phenetic study of a group of plants. This test is achieved by com- paring the results of analyses 1 and 2, in which the data are obtained from the same source, but the latter analysis purposely does not use all of it that is available. From Tables 3 and 4, the surprising fact is that even though the average OTU by OTU relevance is halved from 0.972 to 0.482, still the correlation between the two BSM’s is 0.887. This strongly suggests that a phenetic analysis can tolerate a good amount of missing data and still give reasonably consistent results.
The second purpose of the study was to ascertain the reliability of two sources of estimation of values of characters in taxospecies. This is accomplished by comparing results of the second and third analyses. From Table 3, the average similarity values are approximately equal, as are the average relevance values. The average relevance value of analysis 3 is slightly higher than that of anlysis 2 because there was a certain amount of missing data in Crovello’s original BDM. From Table 4 the
1968 ]
CROVELLO: SALIX
COPHENE TIC VALUE
0:4 0:6 0-8 1-00 aT | ue aa la Cia a= OS
OTU CODE
LASAND CAUDAT LAEVIG GOODIG HINDSI GEYERI EXIGUA PARKSI MELANO EASTWD LUTEA COMUTA MACKEN PIPERI HOOKER LIGULI PSCORD LASLEP TRACYI ORESTR LEMMON PLANIF DRUMSB JEPSON SITCHS BREWER DELNRT ANGLOR
SCOULR NIVALI
Ne se gp a en ee ete eee hy
311
SECTION CODE
o ©
th Oo p O
P 8
Fic. 4. Phenogram of analysis using data from Munz (1959) for 49 characters.
MADRONO [Vol. 19
OTU SECTION COPHENETIC VALUE CODE CODE
0:4 0-6 0:8 00 marl Fe = aes ee | een | aaa
LASAND CAUDAT LAEVIG GOODIG HINDSI EXIGUA PARKSI MELANO
= |
2
LIGULI MACKEN TRACY|
PSCORD COMUTA EASTWD LEMMON 13 oresTR |b 5 SCOULR > 12 JEPSON SITCHS
BREWER DELNRT LASLEP HOOKER PIPERI
ANGLOR PLANIF
NIVALI
DRUMSB + 9 GEYERI }& 13
Fic. 5. Phenogram of analysis using data from Crovello (1966) for 131 characters.
3 4
7 9 8
Ky Ys
1968 | CROVELLO: SALIX Sil)
correlations between BSM’s 2 and 3 and between phenograms 2 and 3 are extremely low in comparison to such pairs of analyses as 1 and 2, 3 and 4, and 5 and 6.
Now compare in Table 4 the results of analysis 2 with the standards (5 and 6) and of analysis 3 with the standards. Even though 2 and 3 show low similarity with each other, both show medium level similarity to the two standards. Furthermore, the correlations of the phenogram of analysis 3 with the two standards is higher than those of analysis 2 with the two standards. The reason for this is not clear at present, but it does indicate that data from floristic studies may be much more reliable than previously thought. However, for maximum reliability the missing data will have to be supplied from outside sources before such information is processed.
In summary, when we consider all the analyses, what emerges is not a qualitative yes-no agreement or disagreement, but rather a continuum. Munz’s information does not agree well with Crovello’s information on the same characters. Crovello’s analysis of 43 characters using full data agrees more with the standards than do those of Munz. (This is to be expected, since the BDM of Crovello’s 43 characters is a subset of that of the standards). Low OTU by OTU relevance had to cause the drop in correlation between analyses 2 and 5 and 2 and 6, as compared to analyses 1 and 5 and 1 and 6.
We may conclude that missing data does make a difference and that distortion of phenetic relationships may result when data from floras are used directly without supplementary information, both to supply the missing data on characters used and to supply information on characters not used at all. But although distortion is present, a rough idea of rela- tionships can be obtained, as shown by the phenogram.
SUMMARY
The effect of missing data and of two methods of estimation of char- acter values on a phenetic study of the willows of California was in- vestigated. This was accomplished by comparison of the data from the floristic study of Munz (1959) with the study of Crovello (1966). Tech- niques of numerical taxonomy compared six analyses of data obtained from the two studies. Since Crovello had almost no missing data for 43 of the characters used by Munz while Munz had a considerable amount of missing data, the effect of missing data could be determined. Missing data was found to distort the relationships among willows. Next, a com- parison of Munz’s data for 43 characters with data from Crovello but with Munz’s missing data pattern served to estimate the effect of the reliability of the data from these two sources. As expected, the agree- ment between the two was low. But both still agreed moderately well with the standard analyses. The overall conclusion is that direct data from floras can be valuable in a numerical taxonomic study. But its value is enhanced when missing data is supplied and when the number
314 MADRONO [Vol. 19
OTU SECTION COPHENETIC VALUE CODE CODE
0:4 0:6 0:8 1-00
LASAND CAUDAT LAEVIG GOODIG HINDSI EXIGUA PARKSI MELANO LUTEA MACKEN LIGULI TRACY! NIVALI GEYERI PSCORD EASTWD COMUTA ORESTR LEMMON > 13 ANGLOR > PLANIF > SCOULR > 12 LASLEP > 4 HOOKER PIPERI + DRUMSB > 9 JEPSON SITCHS : = ao ‘ DELNRT
Fic. 6. Phenogram of analysis using data from Crovello (1966) for 202 characters.
N
Ol
uo @
Wi, JL Ju JeE — —_ji__— Ji JH erp)
ON
1968 | INDEX S15
of characters is increased. The present study also suggests that much information on phenetic relationships present in floras is not used. Such information could be revealed by the regular use of taximetric methods. Finally, use of the techniques described above now permits the taxono- mist to compare, in a reliable and consistent manner, relationships sug-
gested in a floristic study with those presented in a monograph.
Department of Biology, University of Notre Dame, Indiana
Costs of manuscript preparation were defrayed by NIH Corollory Grant GM
11935 held by Robert R. Sokal.
LITERATURE CITED
CrovELLO, T. J. 1966. Quantitative taxonomic studies in the genus Salix. Ph.D. dissertation (unpublished). Univ. Calif., Berkeley. . 1967. Problems in the use of electronic data processing in biological col-
lections. Taxon 16:481-494.
GILMARTIN, A. J. 1967. Numerical taxonomy—an eclectic viewpoint. Taxon 16:8-12.
Muwz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley.
SCHNEIDER, C. 1921. Systematic enumeration of the sections species, varieties and forms of American willows. J. Arnold. Arbor. 3:61-125.
SOKAL, R. R. 1961. Distance as a measure of taxonomic similarity. Syst. Zool. 10:
70-79.
SoKAL, R. R., and P. H. A. SNEaTH. 1963. Principles of numerical taxonomy. W. H.
Freeman, San Francisco.
INDEX TO VOLUME XIX
Classified entries: Chromosome numbers of plants, documented; Reviews. New botanical names are in boldface type. Unannotated taxa in floristic lists are omitted
from the Index.
Acrochaetium densum, 112
Adiantum: capillus-veneris, 66; jordanii, 66; pedatum, 66; aleuticum, 66; xX traceyl, 66
Agropyron: cristatum and A. subsecun- dum, A. cytotaxonomic study of a natural hybrid between, 227; deser- torum, 277; trachycaulum, 227
Alaska: and Yukon, Validity of nomen- clature changes in, 223
Aleuritopteris, 66
Algae: of Washington and _ southern British Columbia. II, Notes on Ma- rine, 111
Amaurochaete fuliginosa, 247
Ammobroma: (Lennoaceae), in Sinaloa, Mexico, A second species of, 179; culi- acana, 180; sonorae, 180
Anderson, C. L., 31
Anderson, D.: Review, The grasses of North Carolina, 60
Anemone edwardsiana petraea, 189
Anemopsis californica, 218
Ano Nuevo I.: San Mateo Co., Cali-
fornia, Terrestrial vascular plants of, 95
Antithamnion pygmaeum, 114
Aphanocapsa litoralis, 111
Arctostaphylos: (Ericaceae), New taxa, combinations, and chromosome num- bers in, 193; andersonii, 207; auricu- lata, 207; canescens, 207; catalinae, 193; columbiana, 207; confertiflora, 207; cratericola, 205; cruzensis, 207; densiflora, 207; edmundsii, 207; glan- dulosa, 207; g. crassifolia, 205; g. cushingiana, 204; g. howellii, 205; g. mollis, 205; g. zacaensis, 205; glauca, 207; glutinosa, 207; hookeri hearsti- orum, 200; h. hookeri, 207; h. mon- tana, 200; h. ravenii, 200; imbricata, 207; insularis insularis, 207; i. pubes- cens, 207; luciana, 207; manzanita bakeri, 202; m. elegans, 201; m. man- zanita, 207; m. roofii, 201; m. wies- landeri, 201; mewukka, 207; morro- ensis, 208; nevadensis, 208; nissenana, 208; nummularia sensitiva 208;
316 MADRONO
obispoensis, 208; otayensis, 208; pajaroensis, 208; pallida, 208; parry- ana, 208; patula, 208; p. platyphylla, 203; pechoensis, 208; pilosula pismo- ensis, 199; pumila, 208; purissima, 195; refugioensis, 208; regismontana, 208; X repens, 208; rudis, 208; stan- fordiana, 208; tomentosa crustacea, 198; t. eastwoodiana, 197; t. hebe- clada, 198; t. insulicola, 197; t. rosei, 198: t. subcordata, 198; t. tomentosa, 208; t. tomentosiformis, 198; t. trichoclada, 198; virgata, 209; viridissima, 209: viscida mariposa, 204; v. pulchella, 204; v. viscada, 209; uva-ursi cratericola
Arizona: IV. Notes on the flora of, 164
Aspidium, 66
Aspidotis, 66
Asplenium: septentrionale, 66; tinum, 66; viride, 66
Athyrium: alpestre, 67; filix-femina, 67; f. angustum, 67; f. californicum, 67; f. sitchense, 67
Audouinella pectinata, 113
Azolla: filiculoides, 71; mexicana, 71
vesper-
Bacigalupi, R.: Review, Botanical Latin, 59 Badhamia: affinis, dearnesii, 249 Bailey, V.O., 32 Bailey, W. W,, 32 Blechnum spicant, 67 Bloomer, H. G., 31 Bock, J. H.: The water hyacinth in Cali- fornia, 281
248; ainoae, 248;
Bonar, L., 125
Botanical Congress: XI International, 136
Botrychium: lunaria, 66; multifidum
coulteri, 66; m. silaifolium, 66; sim- plex, 66; s. compositum, 66
Boyl, W. S., and A. H. Holmgren: A cytotaxonomic study of a natural hy- brid between Agropyron cristatum and A. subsecundum, 277
Brewer, W. H., 31
Burke, J., 30 Calamagrostis: canadensis complex in Alaska, Taxonomy, variation, and
chorology of three chromosome races of the, 235; angustifolia, 235; lan- gsdorfii, 235; purpurea Jangsdorfiii, 235; rubescens on Santa Cruz Island, California, 63
Capraria humilis, 266
Cave, M. S.: Review, Systematic em- bryology of the Angiosperms, 95
Ceramium gardneri, 114
Cevallia sinuata, 7
Chaerophyllum tainturieri, 164
[Vol. 19
Chambers, K. L.: Steens Mountain, 91
Cheilanthes: californica, 67; X carlotta- halliae, 67; clevelandii, 67; cooperae, 67; covillei, 67; feei, 67; fibrillosa, 67; gracillima, 67; intertexta, 67; parishii, 67; siliquosa, 67; viscida, 67; woo- tonli, 67
Chlorogloea conferta, 111
Comatricha rubens, 248
Conobea pumila, 266
Cornus racemosa, 190
Correll, D. S.: Some additions to the flora of Texas. IV, 187
Cottea pappophoroides, 184
Crampton, B.: Three roadside grasses as records for Sacramento, El] Dorado, and Placer Counties, California, 224
Crataegus pearsonii, 189
Cribraria: argillacea, 44; purpurea, 44; splendens, 247
Crovello, T. J.: The effect of missing data and of two sources of character values on a phenetic study of the wil- lows of California, 301
Crum, H.: A new species of Pterigo- neurum from California, 92
Cypress: in Northern California: A geo- graphical review, 19
Cupressus: arizonica arizonica, 52; a. glabra, 52; a. montana, 47; bakeri, 19, 52: macnabiana, 19; pygmaea, 47; sargentii, 19
Cryptogramma acrostichoides, 67
Cryptopleura corallinara, 114
Crytomium falcatum, 68
Cystopteris fragilis, 68
Davis, W. S., and H. J. Thompson: A revision of Petalonvx (Loasaceae) with a consideration of affinities in subfamily Gronovioideae, 18
Dennis, L. R. J.: The genus Kickxia in Oregon, 62
Dermocarpa hemisphaerica, 111
Diarema: andersonii, 247; depressum, 45
Diderma ochraceum, 46
D:dymium sturgsil, 248
Chromosome numbers of plants, docu- mented: Astragalus sclerocarpus, 134; Bomarea hirtella, 134; Cicendia quad- rangularis, 134; Cleomella macbride- ana, 134; Eriocaulon compressum, 134; Eriogonum alatum a. 134; anemophilum, 134; arborescens, 134; caninum, 134; cinereum, 134; com- positum, 134; covilleanum, 135; east- woodianum, 135; elatum, 135; fas- ciculatum, 135; flexum, 135; gigan- teum, 135; gossypinum, 135; gracil- limum, 135; inflatum deflatum, 135; jamesii undulatum, 135; latifolium, 135; parishii, 135; racemosum, 135;
1968 |
roseum, 135; salicornoides, 135; saxa- tile, 135; sphaerocephalum, 135; temblorense, 135; thomasii, 135; um- bellatum, 135; vestitum, 136; Ex- acum affine, 136; Fagonia laevis, 136; Gentiana calycosa, 136; Hackelia cin- erea, 136; Holocantha emoryi, 136; Hypericum formosum scouleri, 136; Idahoa_ scapigera, 136; Orontium aquticum, 136; Panicum capillare oc- cidentale, 136; Penstemon gairdneri, 136; Schoenocaulon dubium, 136; Smilacina flexuosa, 136; Taegetes fili- folia, 136; micrantha, 136; multiflora, 136; subulata, 136; terniflora, 136; Tribulus cistoides, 136
Dorr, C. H-. 31
Dressler, R. L., and J. Kuijt: A second species of Ammobroma (Lennoaceae), in Sinaloa, Mexica, 179
Dryopteris: arguta, 68; dilata, 68; filix- mas, 68
Eichhornia crassipes, 281
Encelia farinosa, 34
Enneapogon: brachystachys, 184; cen- chroides, 184; desvauxii, 184; elegans, 184; pretoriensis, 184; scoparius, 184
Equisteum: arvense, 72; a. boreale, 72; x ferrissii, 72; hyemale affne, 72; laevigatum, 72; palustre, 72; telma- tia braunii, 72
Erigeron lonchophyllus, 164
Erinus verticillatus, 266
Eriocoma cuspidatum, 284
Eriogonum: New species from Utah, 289; batemanii, 291; brevicaule lepto- thecum, 297; cronquistii, 289; elonga- tum, 56; ephedroides, 295; encelio- ides, 55; humivagans, 291; inter- montanum, 293; nudicaule, 293; pan- guicense, 295; pondii p., 56; p. gen- tryi, 57; scoparium, 293; thompsonae t., 299; t. albiflorum, 299; torreyana, 297; tristichum, 295; volmeri, 56; wrightii nodosum, 56; w. pringlei, 56
Eriogonums: from Baja California, Mex- Mexico, Two new, 55
Ernst, W. R.: Review, Index to Euro- pean taxonomic literature for 1965, 58
Erythrotrichia carnea, 112
Euoryzopsis, 285
Ewan, J.: Review, Bibliography of Ne- vada Botanists, 30
Faegri, K.: Steens Mountain, 91
Fissenia, 11
Flora Europaea, 64
Flora North America, 64
Fryxell, P. A., and C. R. Parks: Gos- sypium trilobum: an addendum, 117
Fuertesia domingensis, 10
INDEX 317
Gabb, W. M., 31
Galium frankliniensis, 191
Gambel, W., 30
Gentry, H. S.: A new Hesperaloe from Sonora, Mexico, 74
Geraea cranescens, 34
Goniotrichopsis sublittoralis, 112
Gossypium: species of Baja California, Variation in the diploid, 137; armour- ianum, 144; californicum, 142; da- vidsonii lobulatum, 139; d. sonoricum, 139; gossypioides, 117; harknessian- um, 142; klotzchianum davidsonii, 139; thurberi, 118; trilobum: an ad- dendum, 117
Gramineae: Chromosome numbers in the tribe Pappophoreae, 183
Grasses: as records for Sacramento, El Dorado, and Placer Counties, Cali- fornia, Roadside, 264
Griffin, J. R., and C. O. Stone: Macnab cypress in northern California: A geo- graphic review, 19
Grindelia: howellii, 63; nana, 63; squar- rosa, 63
Gronovia: grandiflora, 9; scandens, 9
Gymnogramme, 68
Gymnosporangium: bethellii, 80; cla- variiforme, 50, 81; cupressi cas- cadense, 52; c. cupressi, 52; hark- nessianum, 81; inconspicuum, 82; kernianum, 83; libocedri, 86; multi- porum, 86; nelsonii, 86; nidus—avis, 87; speciosum, 87; tremelloides, 50
Hall, L.: A survey of the vascular plants above timberline on Mt. Hood, Ore- gon, 130
Hampea, 119
Hawkweed: from the Olympic Moun- tains of Washington, A hybrid, 126
Heckard, L. R., J. T. Howell, and R. Bacigalupi: The scientific writings of Willis Linn Jepson (1867-1946), 97
Hemitomes: congestum in the southern Sierra Nevada, The occurrence of, 234
Herbarium: specimens, Library mailing rates for, 192
Hesperaloe: from Sonora, Mexico, A new, 74; funifera, 76; nocturna, 74; parviflora, 76
Hesse, V. F.: Notes on Madia sativa and related species, 210
Heteranthea limosa in California, 63
Hieracium: albiflorum, 126; gracile, 126
Hippuris vulgaris, 164
Hitchcock, C. L., 32
Holmgren, N. H.: A new species of primrose from Nevada, 27
Hooker, J. D.: 32
Hordeum: jubatum-H. brachyantherum Question, On the, 108
Howell, J. T., frontispiece
318 MADRONO
Hulten, E.: Validity of nomenclature changes undertaken in the flora of Alaska and Yukon, 223
Hybridization: between Encelia and Geraea (Compositae) and some re- lated experimental investigations, Nat- ural, 33
Hymenena kylinii, 114
Iridea lineare, 113
Isoetes: bolanderi, 71; b. pygmaea, 71; echinospora braunii, 71; howellii, 72; h. miniata, 72; nuttallii, 72; occi- dentalis, 72; orcuttii, 72
Jepson: The scientific writing of, 97; Flora of California, 288 Juniperus californica, 20
Kaokochloa nigrirostris, 184
Keener, P. D., and J. E. Hampton: A rust on Anemopsis californica, 218
Kickxia: in Oregon, The genus, 62; ela- tine, 63; spuria, 62
Kiefer, L. I., and B. Joe: Check list of California pteridophytes, 65
Kowalski, D. T.: New records of Myxo- mycetes from California. II, 43; and D. H. Curtis: New records of Myxo- mycetes from California. III, 246
Kruckeberg, A. R.: A hybrid hawkweed from the Olympic Mountains of Wash- ington, 126
Kyhos, D.: Natural hybridication be- tween Encelia and Geraea (Compo- sitae) and some related experimental investigations, 33
Kylinia arcuata, 112
Lamproderma: cribrarioides, 45; guliel- mae, 45
Larrea: divaricata in the North Ameri- can Desert, Chromosome variation in ecotypes of, 161
Lasthenia: glabrata remains diploid, 78; chrysostoma, 78
Lastrea, 68 Lemmon, T. G., 31 Licea: biforis, 246; kleistobolus, 44;
tenera, 246; variabliis, 44
Lindbladia effusa, 44
Lomaria, 68
Lycopodium: clavatum, 73; inundatum, As
Madia: sativa and related species, Notes on, 210; capitata, 210; gracilis, 210
Marsilea: oligospora, 71; vestita, 71
Mason, C. T.: A new family of vascular plants (Psilotaceae) for Arizona, 224; and W. E. Niles: Notes on the flora of Arizona. IV, 164
Mentzelia oligosperma, 11
[Vol. 19
Mia, M. M., and R. K. Vickery: Chrom- osome counts in section Simiolus of the genus Mimulus (Scrophularia- ceae). VIII. Chromosomal homologies of M. glabratus and its allied species and varieties, 250
Mimulus: (Scrophulariaceae). VII. Chromosome counts in section Simio- lus of the genus, 211; VIII., 250; cupreus, 211; glabratus, 250; g. fre- montil, 250; g. parviflorus, 251; g. utahensis, 250; guttatus, 211, 250; luteus, 212, 250; nasutus, 250; pilosi- usculus, 250; tiningii, 211, 250; t. cor- allinus, 217
Mitchell, W. W.: On the hordeum ju- batum-H. brachyantherum question, 108; Taxonomy, variation, and chorol- ogy of three chromosome races of the Calamagrostis canadensis complex in Alaska, 235
Molseed, E. W., 46
Montana: Two new additions to the flora of, 63
Mt. Hook: Oregon, A survey of the vas- cular plants above timberline on, 130
Mt. Pinos: California, The vegetational composition of the south slope of, 225
Myrionema phyllophilum, 111
Myxomycetes: from California. II., New records of, 43; III., 246
Nama dichotomum, 190
New publications, 96, 222
Nolina: arenicola, 187; micrantha, 188
Norris, R. E., and J. A. West: Notes on marine algae of Washington and southern British Columbia. II., 111
Notes and news, 46, 62, 78, 91, 95, 136, 164, 192, 210, 222, 234, 266, 288, 300
Notholaena: californica, 86; jonesii, 68; newberryi, 68; parryi, 68, sinuata, 68
Oak Tree: at Live Oak Tank, Joshua Tree National Monument, California, Identity of the, 256
Onychium, 68
Ophioglossum: californicum, 66; vulga- tum, 66
Ornduff, R.: Lasthenia glabrata remains diploid, 78; Review, The Komarov Botanical Institute, 221; Jepson’s Flora of California, 288
Oryzopsis: hendersonii (Gramineae), Notes on, 283; asperifolia, 284; ex- igua, 284; hymenoides, 284; miliacea, 284; racemosa, 285
Paleoeriocoma, 286
Pappophorum: bicolor, 184; mucronu- latum, 184
Paronychia sessiliflora, 164
1968 |
Parthenium, 33
Pellaea: andromedifolia, 68; a. pubes- cens, 68; brachyptera, 68; breweril, 68; bridgesii, 68; longimucronata, 69; mucronata, 69; m. californica, 69
Peridermium harknessii, 51
Petalonyx: (Loasaceae) with a consid- eration of affinities in subfamily Gro- novioideae, A revision of, 1; crenatus, 12; linearis, 13; nitidus, 16; palmeri, 17; parryi, 17; thurberi t., 15; t. gil- manii, 15
Peterson, R. S.: Studies of Juniper rusts in the west, 79; Cypress rusts in Cali- fornia and Baja California, 47
Phegopteris, 69
Phillips, L. L., and D. Clement: Vari- ation in the diploid Gossypium species of Baja California, 137
Physarum: crateriforme, 45; diderma, 45; gilkeyanum, 249; mutabile, 249
Pilularia americana, 71
Pinus jeffreyi, 228
Piptatherum paradoxum, 284
Pityrogramma: pallida, 69; semipallida, 69; triangularis, 69; t. maxonii, 69; t. viridis, 69; viscosa, 69
Pleurocapsa entophysaloides, 111
Polypodium: californicum, 69; c. kaul- fussil, 69; glycyrrhiza, 69; hesperium, 70; scouleri, 70
Polystichum: dudleyi, 70; kruckebergii, 70; lemmonii, 70; lonchitis, 70; muni- tum, 70; m. curtum, 70; m. imbricans, 70; m. nudatum, 70; X scopulinum, 70
Porter, D. M.: Review, An annotated bibliography of Mexican ferns, 62
Primrose: from Nevada, A new species Oi 27
Primula: angustifolia, 29; cusickiana, 29; maguierei, 29; nevadensis, 27
Proboscidea crassibracteata, 190
Pseudodictyon geniculatum, 115
Pseudulvella consociata, 115
Psilotaceae: for Arizona, A new family of vascular plants, 222
Psilotum nudum, 224
Pteridophytes: Check list of California, 65
Pteridium aquilinum pubescens, 70
Pterigoneurum: from California, A new species, 92; californicum, 93; sub- sessile, 93
Pteris: aquilina, 70; cretica, 70; vittata, 70
Puccinia aristidae, 218
Purpus, C. A., 32
Quercus: californica, 256; dumosa, 256; dunii, 256; gambelii, 256; garryana, 256; lobata, 257; & munzii, 257; tur- binella californica, 256
INDEX 319
Ranunculus sardotus, 189
Raven, P. H.: Calamagrastis rubescens on Santa Cruz Island, California, 63; Review, Flora of Turkey and the East Aegean Islands, Vol. II, 287
Reeder, J. R., and D. N. Singh: Chromo- some numbers in the tribe Pappo- phoreae (Gramineae), 183
Remy, E. J., 31
Reveal, J. L.: New species of Eriogonum from Utah, 289; and C. A. Hanson: Two new Eriogonums from Baja Cali- fornia, Mexico, 55
Rhodophysema georgii, 113
Reviews: Reifschneider, O., Biographies of Nevada botanists, 30; Brummitt, R. K., Index to European taxonomic literature for 1965, 58; Stern, W. T., Botanical Latin, 59; Blomquist, H. L., The grasses of North Carolina, 60; Hollenberg, G. J., and I. A. Abbott, Supplement to Smith’s marine algae of the Monterey Peninsula, 61; Jones, G. N., An annotated bibliography of Mexican ferns, 62; Neitzel, W. J., The flora and fauna of Solano County, 94; Barneby, R. C., Atlas of North American Astragalus, 94; Davis, G. L., Systematic embryology of the an- giosperms, 95; Lowe, J. L., Polypo- raceae of North America. The genus Poria, 220; Shetler, S. G., The Ko- marov Botanical Institute, 250; Wil- liams, J. C., and H. C. Monroe, A field guide to the natural history of the San Francisco Peninsula, 222; Davis, P. H., Flora of Turkey and the East Aegean Islands, Vol. II, 287; Dallimore, W., and A. B. Jackson, A handbook of the Coniferae and Gink- goaceae, 287
Rice, B.: The occurrence of Hemitomes congestum in the southern Sierra Ne- vada, 234
Roestelia lacerata, 54
Rundel, P. W.: The southern limit of Texus brevifolia in the Sierra Nevada, California, 300
Rust: on Anemopsis californica, A, 218; in California and Baja California, Cy- press, 47; in the west, Studies of Juniper, 79
Salix: anglorum antiplasta, 305; caudata, 305; commutata, 305; coulteri, 303; delnortensis, 305; drummondiana sub- coerulea, 305; eastwoodiae, 305; ex- igua, 305; geyeriana, 305; hindsiana, 305; hookeriana, 305; jepsonii, 305; laevigata, 305; lasiandra, 305; lasio- lepis, 305; lemmonii, 305; linguifolia, 305; lutea, 305; mackenziana, 305; melanopsis, 305; nivalis, 305; orestra,
320 MADRONO
305; parksiana, 305; piperi, 305; plan- ifolia monica, 305; psuedocordata, 305; scouleriana, 305; sitchensis, 305; traceyl, 305
Saussurea americana, 63
Schmidtia papppophoroides, 184
Selaginella: asprella, 73; bigelovii, 73; cinerascens, 73; douglasii, 73; engel- manii scopulorum, 73; eremophila, 73; hansenii, 73; leucobryoides, 73; oregana, 73; wallacei, 73; watsonil, 73
Sequoiadendron: giganteum, Seed ecol- ogy of, 267
Setcreasea leiandra glandulosa, 187
Shetler, S. G.: Flora North America, 64
Schockley, W. H., 32
Snyder, J. A., 30
Sonne, C. F., 32
Sorsa, M.: Cytological and evolutionary studies on palustres violets, 165
Spellenberg, R.: Notes on Oryzopsis hen- dersonii (Gramineae), 283
Spermacoce floridiana, 192
Stark, N.: Seed ecology of Sequoiaden- dron giganteum, 267
Steens Mountain, 91
Stemodia: verticillata Hassler, Confu- sion in authorship of, 266; arenaria, 266; parviflora, 266; verticillata, 266
Stipa, lemmonii, 284
Stretch, R. H., 32
Struthiopteris, 70
Suillus: California Boletes. III. The genus, 148; acerbus, 149; allidepes, 150; americanus, 150; borealis, 150; brevipes, 150; caerulescens, 150; fus- cotomentosus, 151; glandulosipes, 151; granulatus, 151; lakie 1, 152; 1. pseudopictus, 151; lithocarpi-sequoiae, 152; megaporinus, 152; monticolus, 154; ponderosus, 154; punctatipes, 154; pungens, 154; riparius, 155; to- mentosus, 157; umbronatus, 157; vol- canalis, 158
Taxus: brevifolia in the Sierra Nevada, alifornia, The southern limit of, 300 Texas: Some additions to the flora of, 187
Thelypteris: 71
Thiers, H. D.: California boletes. III. The genus Suillus, 148: Review: Poly- poraceae of North America. The genus Poria, 220
Thomas, J. H.: Review: The flora and fauna of Solano County, 94; Review: Atlas of North American Astragalus, 94; Terrestrial vascular plants of Ano Nuevo I., San Mateo Co., California, 95; Library mailing rates for herbar- ium specimens, 192; Review: A field
nevadensis, 70; puberula,
[Vol. 19
guide to the natural history of the San Francisco Peninsula, 222
Tigridia, 46
Torreya california, 300
Torrey, J., 31
Tuberculina persicina, 218
Tucker, J. M.: Identity of the oak tree at Live Oak Tank, Joshua Tree Na- tional Monument, California, 256; and J. McCaskill: Heteranthera limosa in California, 64
Uredo: apacheca, 88; cuperssicola, 47; houttuyniae, 218 Uromyces saururi, 218
Veatch, J. A., 31
Vickery, R. K., K. W. Crook, D. W. Lindsay, M. M. Mia, and W. Tai: Chromosome counts in section Simi- olus of the genus Mimulus (Scro- phulariaceae). VII. New numbers for M. guttatus, M. cupreus, and M. til- ingii, 211
Viola: achyrophora, 170; blanda, 168; epipsila, 166; e. repens, 168; incognita, 168; juressii, 166; lanceolata, 175; macloskeyi, 173; pallens, 168; palus- tris, 170; p. brevipes, 172; p. herminii, 166; p. juressii, 166; primulifolia, 175; renifolia, 168
Violets: Cytological and evolutionary studies on palustres, 165
Vogl, R. J., and B. C. Miller: The veg- etable composition of the south slope of Mt. Pinos, California, 225
Water Hyacinth: 281
Wells, P. V.: New taxa, combinations, and chromosome numbers in Arcto- staphylos (Ericaceae), 193
West, J. A.: Review, Supplement to Smith’s marine algae of the Monterey Peninsula, 61
Wiggins, I. L.: Confusion in authorship of Stemodia verticillata Hassler, 266; Review, A handbook of the Coniferae and Ginkgoaceae, 287
Willows: of California, The effect of missing data and of two sources of character values on the phenetic study of the, 301
Woodland, D. W.: Two new additions to the flora of Montana, 63
Woodsia: oregana, 71; scopulina, 71
Woodwardia fimbriata, 71
in California, The,
Yang, T. W., and C. H. Lowe: Chromo- some variation in ecotypes of Larrea divaricata in the North American desert, 161
Yucca carnerosana, 188
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