x ee) > > > = a " = - P x m m m- w = w = w } IBRARIES SMITHSONIAN INSTITUTION NOILONLILSNI NVINOSHLINS S31IYVUET “” = * ” = wn = < = TL cg = , = z= 4 "Wid fy Ln 4 Nn 7p) on ? fh wm O ; Vi / Life : : 2 Oi teu) 3 Zz = ~S >” S >" | ae 77) ae Zz “” ve —NVINOSHLINS S34 lYvVYdgit LIBRARI ES SMITHSONIAN INSTITUTIC Zz Zz Zz - : = = YG," Z < = < Uff a 2 NO"'s = a M7 fei S O "SN = : Oo = fe) =z oe Zz : ad Ps IBRARIES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS = = z - | z ° “ OD ° wo N& ° = "9 ke aa . = = é > — > “= - GCZY, os - i/o 2 m Q = : 7) = w . £ VOILALILSNI NVINOSHLINS S3INYVYSIT LIBRARIES SMITHSONIAN z . ” ez Roe Y” z << = a = < a = , = at z 7 wy a - wn an < O 4% < O mo r= = = = = 7) z 7%) > 2 on : SMITHSONIAN INSTITUTION NOIJLONLILSNI NVINOSHLINS S3Iiuvdd w > ep) = ep) ul 7 =e o = a. = oc = oc \ 70 > ee > Ie > _ % > - = ms pe = w = w m O = . = Ww INSTITUTION NOILALILSNI NVINOSHLIWS saruva dl N¥INOSHLIWS NVINOSHLIWS SMITHSONIAN NVINOSHLIWS ad > wn S3IYVYRIT LIBRARIES SMITHSONIAN INSTITUT “ya NOD j VOLLALILSNI_ NVINOSHLIWS Aner uo " A ARIES SMITHSONIAN 0, JLILSNI nay pc JLILSNI l, La LILSNI Li» "3 “iy ARIES 0, 4 ayst S3IYVYUEII_LIBRAR ett ws NDC VLILSNI Hy SONIAN ‘TV = Ne SJ m& \ i= li 5 <70n Ae © m Onc z= mal NVINOSHLINS _S31¥Vvud!T LIBRARIES SS rs) = aN N = Pe) SW OE = SSE = Xv SMITHSONIAN : Vell = Z Ls “of he INSTITUTION NOILALI S3J!IYVYSIT LIBRAR ‘ANG y es eh 4 = INSTITUTION NOILAL! ARIES SMITH INSTITUTION NOILALILSNI NVINOSHLIWS Saluvudils 2) a8 w z= a j x = < \. = £ yy =< = Ve = oa z xX + ‘fy zz 4 , \ YM ; ZO NL F § Whi HZ? 5 fei 3 Sie) r MIN 6 HF = O gx NSE E QO" 2.474" = Z = wos i Sa z > 3 ILILSNI_NVINOSHLINS Sa fYuvVYgiT LIBRARI ES SMITHSONIAN _INSTITUTION ss = +3 b Ww & ae yy & Ea -_— oc = oc 4 Yi Ye — [8 a) _< = < Die = < 4 x a = o Vey Cc ow a SS 28) = om.” 4 ni co eee 2 = 2 5 ARIES SMITHSONIAN INSTITUTION NOILNLILSNI S31uYvugly = Me = - z iz = “0 = © oO = a at aA = 3 - ad (ip fg > = > $ > > = ty ‘ a ran ») = = 2 Vif? m a m D ys = nN — 2) . —_ NLILSN!I NVINOSHLINS S31Y¥Vuat SMITHSONIAN z w = 7) > n < = r = = < = i £ fs 4 vy Pa = = z 4 WiC iZH ERS BEDI 3 Yi? = 2 UY it = =: E = ee ee wee 2 - < B E Ww) 7 ‘ARTES SMITHSONIAN INSTITUTION NOILNLILSNI NVINOSHLINS S31YVYusIT_._ ua z tu 2 &. uf Z os a” cas w a (op) =e = = = = = 4 : : : : : = iene 5 S ruse van 2 awd ox} VLILSNI NVWINOSHLINS S3IY¥vVYdIT LIBRARIES SMITHSONIAN — ° 5 a Zz 8 z 3 = ; = Gy x 5 = > bP, = ca = se Ee Jee Zs S Fe = =e Gp 2 = an 7 Z B Z SARIES SMITHSONIAN INSTITUTION NOILOLILSNI NVINOSHLINS S3IYVudt) ” Ze: ” 2 77) : z = x ¢ = < = = = Zz Woe F ayy os z S 2) vy -. N 77) "hh D wn aes 2 O Sa we FAO O WW fh, TX re) aa WN SE E Qo 2Z, ao Z. FE we 5B 5 S 3 ALILSNI_NVINOSHLINS Sa lYvVuail LIBRARI ES SMITHSONIAN INSTITUTION a a = STITUS Te) uJ stiTuy n SOND us tad ~ at — {5 ON s ee uN a MR 2 Ye = Ml EM = = _=—»_ i} ay 7 no . = oD 1 - a ay : MADRONO A WEST AMERICAN JOURNAL OF BOTANY VOLUME XXIII 1975-1976 BOARD OF EDITORS Class of: 1975—ArtTURO GOMEz-PoMPA, Universidad Nacional Autonoma de México Duncan M. Porter, Missouri Botanical Garden, St. Louis 1976—DeEnnis ANDERSON, Humboldt State College, Arcata, California KINGSLEY R. STERN, Chico State College, Chico, California 1977—Date M. SmirH, University of California, Santa Barbara Wit1i1AmM Louis CULBERTSON, Duke University, Durham, North Carolina 1978—SHERWIN CARLQUIST, Claremont Graduate School LesuiE D. Gott iets, University of California, Davis DeENNIs R. PARNELL, California State University, Hayward 1979—Puitiep W. RUNDELL, University of California, Irvine ISABELLE TAVARES, University of California, Berkeley 1980—JaMEs R. GriFFIN, University of California, Hastings Reservation Frank A. Lanc, Southern Oregon College, Ashland 1981—DanIEL J. CRAwrorp, University of Wyoming, Laramie James Henrickson, California State University, Los Angeles Editor — JouHn L. STROTHER Associate Editor — ALAN R. SMITH Department of Botany, University of California, Berkeley 94720 Published quarterly by the California Botanical Society, Inc. Life Sciences Building, University of California, Berkeley 94720 Printed by Gillick Printing, Inc., Berkeley, California 94710 11 Volume 23 of Madrono is dedicated to Lincoln Constance, Professor of Botany, University of California, Berkeley, in recognition of his contri- butions to our knowledge of western American plants, particularly his taxonomic studies on Umbelliferae and Hydrophyllaceae, and for the leadership he has exerted in the botanical community. iil TABLE OF CONTENTS ACKERMAN, JAMES D., Reproductive biology of Goodyera oblongifolia (Orchidaceae), y.a5 25s eo eas tas feces esate a ee ee res eae ALMEDA, FRANK, JrR., A new species of Symplocos (Symplocaceae) from Mexico AsEN, Per Arvin, Upper intertidal algal zonation on Bodega Head, Sonoma (County 7 C alin nascent ee eee Barsour, MicHaer G., THEODORE M. DE Jonc, and Ann F. Jounson, Additions and corrections to a review of North American Pacific Coast beach RODE Lal tel Oita seater Ries Rye thee One ey Se eeeecatese oe poem tae ee LO et ee BENSELER, Rot¥F W., Floral biology of California buckeye Boum, Bruce A. (see ORNDUFF, ROBERT) BREEDLOVE, DENNIS E. and EvizABETH McCtiintock, Thornea (Hypericaceae), a new genus from Mexico and Guatemala BRM Gn I) (Gee Viasek vet alt). coe scctce seston ea ek eet cee ee IB ORKART (Ac (See mlm pSOneteal: )i ceccncare ee tesece es ences eps aeeesae ee ee eee CARMAN. (IN. 2 CSeecolmpson, et: ali): eis ee a ee es ee CarTER, ANNETTA M., The Ynés Mexia collections and N. Floy (Mrs. H. P.) | 5 $0221 6151 LH c uae ee ts eee re SOCIO TOD Ser NTR MmNE APTN Cooma Uren i Wie a Merk ke os Be CAVE: MARTON: (Scex@rmncdluit, RiObert)\\ Ga. ea eee ere ee eee CHAMBERS, KENTON L. (see Fellows, Charles EB.) o0........c....ececeeeeseseeeecccceeeeeeeeeeeeeee Cuuanc, T. I. and L. R. Heckarp, Taxonomic status of Cordylanthus (subg. Dicranostegia) orcuttianus (Scrophulariaceae) ..............2...eeeceeeeeeeeeeeeeeeeeeeeeeee Cuuanc, T. I. and L. R. HEcKArpD, Re-evaluation of bract morphology in taxonomy of Cordylanthus (Scrophulariaceae) -0........2.000.220cceeecceeeeeeeeeeeeeeeeeeeees DavipsE, GErRIT, A study of some intermountain violets (Viola sect. Chamae- TICE ETIZU IVY ce et a ee eR eRe ag ee DE Jonc, THEeopore M.-(see Barbour et al.) (522i kee ee ee DEL Morar, RoGeER (see Lebednik, Gretchen K.) Dempster, LAURAMAY T., A new species of Galium (Rubiaceae) from Coahuila DeMPsTER, LAURAMAY T., An Asian Kelloggia (Rubiaceae) .......0...00000000000.-.00-220---- DempPsTER, LAURAMAY T., A new species of Galiwm (Rubiaceae) from the Sierra VE arcrie vier Cale eae en eae eee as ee ee ee DEMPSTER, LAURAMAY T., Galium mexicanum (Rubiaceae) of Central America ane western INOLtneAIMCTICAs 7. c.ccoe eo eeatecee cee es Sues eg, se DENTON, MELINDA F. (see Elvander, Patrick EF.) 20.............12.ccceeeeceeeeceeeeeeeeeeceneeeedeees Dorn, Rosert D., Cytological and taxonomic notes on North American Salix... Doyle, WILLIAM T., Sphaerocarpus michelii Bell.: a new liverwort for Cali- 1018 11: ee a le me AE ee Nr at eee Oe enero rer ea ern ECKENWALDER, JAMES E., Profiles of California vegetation and The distribution of forest trees in. California (YeVvieWSs)) 2.222. 2 coi acct ee ECKENWALDER, JAMES E., Re-evaluation of Cupressaceae and Taxodiaceae: a PROPOSEC “MEK CEH ons sae ge aca ceekcd oernse eee seeet ede ate weg eee ELVANDER, PATRICK E. and Metinpa F. Denton, Savxifraga tempestiva (Saxi- fragaceae), a new species from the Pacific Northwest .........00000...00..:.:.::eeeeeeee SEINCER OD): El (Sees VAS Ke bral): cic ceaccted foes cates ree eee FELLOws, CHARLES E., Chromosome counts and a new combination in Claytonia SECtmiciiia + (E OFtUlAGACeAe)) i. sre fe opt tee ee ee ee 191 365 114 163 274 386 108 346 FELLOWS, CHARLES E. and Kenton L. CHAMBERS, Nomenclatural notes on Clay- LOVLVO SLD VELOLCEL Cpe ee ne ee ee ee teen eee ee 297 FISHER, JACK Cz Jey. Two new plants for Nevada ...22..220..0 2 12 FRYXELL, PAut A., Mexican species of Abutilon sect. Armata (Malvaceae), in- cluding descriptions of three new SPECIES ......-.....20....22..ceeeeeeeeeeeeeeeceeeeeeeeeeeeeeteeeeees 320 GANDERS, FreED R., Heterostyly, homostyly, and fecundity in Amsinckia specta- Giles BOA RTM AC eA cy ee re ents re seeieae eres eeeerte tert seen eee ere eee 56 GILLESPIE, Ms. Terry S., New distribution records for Montana ~........0202000...... 100 GILLETT, JoHN M., A new species of Trifolium (Leguminosae) from Baja Cali- tO Tet ee MC ACO ee crtccerse Waser sate ad erence ees geet eee 334 Guap, JupITH B., Taxonomy of Mentzelia mollis and allied species (Loasaceae) 283 GoopwIin, Joyce F.,, Spontaneous hybrids in Dodecatheon (Primulaceae)............ 81 Gorpon, THoMAS R. (see Grayum, Michael H.) 00000000 360 Gorpon, THOMAS R. and MicHaEL H. Grayum, Sedum spathulifolium (Crassu- laceae), new to Santa Monica Mountains, California -..0.00000000...00220000....eeeeeeee- 454 GrayumM, Micwaert H. and Kennetu A. Witson, Bacopa monnieri (Scrophu- laviaceac). a new record for @aliformid 22. ee 161 GrayuM, Micuaert H. and Tuomas R. Gorpon, Agrostis thurberiana (Gram- inede )hanew tO mouthern alioriia, 2.22 ct. ee ee eee 360 GrayuM, MicHAEL H. (see Gordon, Thomas R.) @00..2...2..ceee ce eeeeeeeeeceeeeeeeeeeeeeeeeeeeees 454 GrirFIn, J. R. and J. M. Tucker, Range extension for Quercus dunni in GEMUCAls OAM O VT ee Ee etek ant oe Ae Ree, eae eh le ee 295 Hates, Rosert D7, New plants for ‘California 2....2... ee 162 ble CicARD ack, (see CWA me. cbs dh)? coc. ve este tsar. te esto ceecaaed en chet ae epee 88, 169 HENnrRICKSON, JAMES and Barry Pricce, White fir in the mountains of eastern Wiohave 1) eSenrt ton. (Calitanrnia: 2.is ee eee eee ee 164 IENRICKSON,, JANES (See Reveal, Jamies 1.) ssc eee eee eco cee ee 205 Plaine AINA CSCO UIC MIG el al)ise ae cone nore rate: i eo eee gee gee ae 299 Houn, JANET E., Argemone munita (Papaveraceae): range extension and mor- TENO LG CaGall eta OHCs mn meee we nae ee aati Re en oe Neate ose tte, oe hte pane aes 98 WiOHNSON, ANN EF. see Barbour et al.) ..2202..5 sho ccc 130 NOTINSON El Bb. See Vasekiet/ali) o0 ese ese een 1, 114 Jounson, LERoy C., Range extension of three conifers and a dwarf mistletoe in the Panamint Mountains, Death Valley National Monument.......................... 402 KEELEY, Jon E., Morpological evidence of hybridizafion between Arctostaphylos RUOTICOANGUA “PUNGeITS CEIIGACCAC) 1l.c.cqteceee soe eee eye ae 427 Keit, Davin J., Revision of Pectis sect. Heteropectis (Compositae: Tageteae) .... 181 KeiL, Davin J., A new species of Chrysactinia (Compositae: Tageteae) from Sinaloa, Mexico KeiL, Davin J., Chromosome numbers for Euphorbia (Euphorbiaceae) from WESUCTING NOL AIMELIGA: 224.0. oie n upto ee he, ee 405 KeiL, Davip J. (see Lane, Meredith A.) ...o.....2..cecccecccceeccecceeeeececeeeececeeceeseveseveeeceeeeeees 457 LackKscHEw1t1z, Kraus H., Montana mountain flora: new records .......................- 361 Lane, MerepitH A. and Davin J. Keir, Glinus radiatus (Aizoaceae), chromo- some count and range extension to Arizona .......0....22 oo. cece eee eee 457 LarGENT, Davin L., A new species of Claudopus from northern California............ 376 LASSETTER, J. STUART, Taxonomic status of Vicia hassei (Leguminosae) .............. vis LAVEN, NIKKI (see McBride et al.) ............--cccescccecceecceeccccecececeeeccesseeeeceeseeeneeeeseenececeeees: 299 LaysER, EARLE F., Washington wildflowers (review) .............22...--.0c2:cc2eeceeeseeeeeeeees LEBEDNICK, GRETCHEN K. and ROGER DEL Morat, Vegetation surrounding Kings IbakeBog, Washinetons.s 2.525) cviecrces ois Meat ae LESKINEN, PAut H., Occurrence of oaks in late Pleistocene vegetation in the MiojavemDesert of INGVada 22.2 cx erred seca ee LesTER, Gary S., Empetrum hermaphroditum (Empetraceae) in Humboldt COME y, 2 CAIN EI ec eee ce eae aS ck as eee One ta re see eee LitTRELL, Epwarp E. (see McDonald, Philip M.) -....2...-..c.cacceee cece eeeeceeenceneneeeeeee McBripe£, Jor, TANA Hirt, RANDY MILLIKEN, and Nikki LAveEN, Vascular plants of montane coniferous forest in San Bernardino Mountains, California ........ McCtiinTock, ELIZABETH (see Breedlove, Dennis E.) -....00...0...2.0000.22ee tee eeeeeeeeee eee McDonatp, Puitip M. and Epwarp E. Littretzt, The bigcone Douglas-fir— canyon live oak community in Southern California ......0000.20 ue eee eee McLeop, Matcorm G., A new hybrid fleshy-fruited prickly-pear in California Major, Jack, A flora of the Trinity Alps of northern California (review)............ Major, JAck, Die Vegetation Osteuropas, Nord- und Zentralasiens (review)...... Mason, CHARLES T., Jr., Apacheria chiricahuensis: a new genus and species PU OMUATIZ ODAC. tec PO ee oe el 2 aes aye en on a MEACHAM, CHRISTOPHER A. and Top F. Sruessy, Procedural innovations in revisionary studies: computer-assisted citation of representative specimens Mii ETICEN, RANDY i(see WICB ride et: al): <.22.5cce see eee eee MonTGOMERY, KENNETH R. and THoMAs W. Strip, Regeneration of introduced species of Czstus (Cistaceae) after fire in Southern California —.....0..000000000.... Moran, Rein, A field guide to the common and interesting plants of Baja Cali- fornia (review) .............-..-----.-- elu des aac ae tute iac a oe ree ate ae he eee eee oe Morcan, WILLIAM T. (see Spires, David E.) .000.2..........0.ccececceeceeeeeee eee cceeee ee eeeeeeeeeeeee MORRIS, ViaeHArT. 1..)(see Rurier, BUA) 22 ee ee OrNDUFF, RoBerT and Bruce A. Boum, Relationships of Tracyina and Rigio- POD Pas. ((COMPOSIEAC 2c cscec ere ge estonia wes ceeee case eats cee eee eee ORNDUFF, ROBERT and Marton Cave, Geography of ane and chromosomal heteromorphism in Leucocrinum montanum (Liliaceae) ...........220..220...2000020---- PATTERSON, THOMAS F. (see Riskind, David H.) -2.0........02..000cceeettceeeeeeeeeeeeee eee Pati Fins (See Ouiek cC oR ey tcissc.et oe tat rat ole aioe fod erates Sec ane a aye oe, wees eo Piz, "GEORGE, (See Strother, | ONM® lex) geceseeeecntee seems cea ese ee ee PINKAVA, DonaLp J., A new species of cholla (Cactaceae: Opuntia) from Coa- Huntlas WEOXiGO. 2 -x-o 2 ont ecee tee eo oe ee eee PRicce, BARRY (see Henrickson, Jiames): 2.22cc00. Sis. 2a eee Quick, C. R. and F. A. Patty, Seed logevity of the Sierra Gooseberry ................ REEDER, JOHN R., Systematic position of Redfieldia (Gramineae) .......................- REEVES, TrMotTuHy, Range extensions for lower vascular plants in Arizona .......... REVEAL, JAMES L. and JAMEs Henrickson, A new variety of Eriogonum erict- OVI OPGlY COMACEAC) cetera te rast ace cs rence eee ee eee Riskinp, Davin H. and Tuomas F. Patrerson, Distributional and ecological TOTES (OM L271 00S CULLINT ILE COV: ie ene ene ee gf een ee eee ec Rocers, C.M., Relationships of Hesperolinon and Linum (Linaceae) .................- Rost, Tuomas L. and A. Dantet StmPer, The germination lid: a characteristic of thetemma in, the Paniceae 2.25.2 ee 292 164 236 434 454 205 159 13 RUNDEL, Puitie W., Primary succession on granite outcrops in the montane SOU Mel Me SLT ALIN © Vil ge ce Sere eae tee ec ve np eg OP tee se Dn oop ee eeve eae SAucCK, JANE REEsE, Distribution, chromosomes, and taxonomy of Parthenice POLES CG ONIN. SILAC)) gee ace scsi sa oe ea noeseee ne ar cn ace eee Dera eeeet aac este SAUER, JONATHAN D., Remnant seashore vegetation of northwest Costa Rica ........ ScumMip, Rupotr, Fly pollination of Penstemon davidsoni and P. procerus (Senoplhulariaceaey. A cete se eer cect esate coca dao erent tense ee tees sey eee obec eee SEAMAN, F. C., Hymenoclea platyspina (Asteraceae: Ambrosieae), a new species | Hagen ne eye ak ec Ub ie 0 [maa te NG loge une eppeure- (ost oesee cata neal eae aes oO eae DINEPER, A. IDANIEr. (see Rost, Thomas L.)) 2.03... ee Stmpson, B. B., A. BuRKarT, and N. J. CarMAN, Prosopis palmeri: a relict of an ancient North American colonization 2 .2--cccce ose ee SMITH, ALAN R., The California species of Aspidotzs SMITH, JAMES PAYNE, JR., The grasses of Texas (review) SPELLENBERG, RICHARD, Synthetic hybridization and taxonomy of western North American Dichanthelium, group Lanuginosa (Poaceae) SPIRES, Davin E. and Wiii1am T. Morcan, Nebraska flora: new records ............ STERNBERG, LEONEL, Growth forms of Larrea tridentata STEVENSON, DENNIS WM., Taxonomic and morphological observations on Botry- chium multifidum (Ophioglossaceae) _...............2.2..ccqeececeeeeeceeeeeeeeeeeeeeeeeeeeeeeeeeeeeeees STEWART, JOAN G., Development of morphological patterns in three species of Delesseriaceae StrRAw, RicHarp M., A new species of Penstemon (Scrophulariaceae) from Mexico StTRID, THomas W. (see Montgomery, Kenneth R.) STROTHER, JOHN L. and GeorcE Pitz, Taxonomy of Psathyrotes (Compositae: Senecicneae) eee een ee en nn eee eee eee errr re ee STROTHER, Joun L., Aquatic and wetland plants of southwestern United States (review) a ee eee eee ee eee eee eee eee eee eee ee eee eee STROTHER, JOHN L., Chrysanthellum pilzii_ (Compositae), a new species from Oaxaca STUEssy, Top F. (see Meachum, Christopher A.) SUNDBERG, WALTER J. (see Thiers, Harry D.) o0.2...2..22cceecececcceeecceeeeeeeeceeeeeeeeeeeeeeeees Taytor, DEAN Wo., Disjunction of Great Basin plants in the northern Sierra Nevada Tuiers, Harry D. and WALKER J. SUNDBERG, Armillaria (Tricholomataceae, Agaricales) in the western United States including a new species from Cali- fornia See et See ee ee eee eee ee eee ee ee er Tuomas, JoHN H., Salsola soda L. (Chenopodiaceae) in central California ........ Topsen, THomas K., Charles Wright’s “E] Paso” collections and the type locality for Psathyrotes scaposa (Compositae) Tucker, J. M. (see Griffin, J. R.) TurNER, B. L. and Micuaer I. Morris, New taxa of Palafoxia (Asteraceae: Helenieae) SSS nS ESO Se RS SEIS Se IRE gprs. HODES DOO Domo Soon SeSrinicesicserarec UrsatscH, Lower E., Systematics of the Ericameria cuneata complex (Com- positae, Astereae) VaseEKk, F. C., H. B. JoHnson, and D. H. Estincer, Effects of pipeline construc- tion on creosote scrub vegetation of the Mojave Desert Vil VasEK, F. C., H. B. Jounson, and G. D. Brum, Effects of power transmission lines cn vegetation of the Mojave Desert .......0...0.......:.2:2:0cscceecseeeeseeseeeeeeteeeeeees Wiccins, Ira L., Ranunculus flagelliformis discovered in the Galapagos Islands WILKEN, Dieter H., Collomia tinctoria (Polemoniaceae) in Southern California Witson, KennetH A. (see Grayum, Michael H.) ....2..220-..c...2cccc.eeecce eee Younc, Davin A., Correction of the geographic distribution of Rhus microphylla (Anacardiaceae) Pe wre meme ees feces e nee eee cme e cena ewan ad san eee n nate e wee ee ews cece ence scare eens es cccen sense eesees Younc, Davip A., A new subspecies of Rhus chondroloma (Anacardiaceae) from Mexico ERRATUM p. 3. Formula for Community Quality Index should read: CQI= \ % total perennial cover X % of total perennial cover by long-lived species Dates of publication of Madrono, volume 23 No.1,pp. 1-— 64: 2Jan 1975 No. 5, pp. 237-300: 2 Jan 1976 No. 2, pp. 65-104: 1Apr 1975 No. 6, pp. 301-364: 8Apr 1976 No. 3, pp. 105-168: 1Jul 1975 No. 7, pp. 365-404: 1Jul 1976 No. 4, pp. 169-236: 10 Oct 1975 No. 8, pp. 405-460: 1Oct 1976 Vill MAD VOLUME 23, NUMBER 1 JANUARY 1975 A WEST AMERICAN JOURNAL OF BOTANY Contents EFFECTS OF PIPELINE CONSTRUCTION ON CREOSOTE SCRUB VEGETATION OF THE MojavE DEsERT, F. C. Vasek, H. B. Johnson, and D.H. Eslinger A NEw SPECIES OF GALIUM (RUBIACEAE) FROM COAHUILA, Lauramay T. Dempster THE CALIFORNIA SPECIES OF AspripoTis, Alan R. Smith TAXONOMY OF PSATHYROTES (COMPOSITAE: SENECIONEAE), John L. Strother and George Pilz FLorRAL BioLocy OF CALIFORNIA BUCKEYE, Rolf W. Benseler RELATIONSHIPS OF TRACYINA AND RIGIOPAPPUS (COMPOSITAE), Robert Ornduff and Bruce A. Bohm HETEROSTYLY, HOMOSTYLY, AND FECUNDITY IN AMSINCKIA SPECTABILIS (BORAGINACEAE), Fred R. Ganders NOTES AND NEWS ANNOUNCEMENT OF MEETING SPHAEROCARPOS MICHELII BELL.: A NEw LIVERWORT FOR CatrrorniA, William T. Doyle RANUNCULUS FLAGELLIFORMIS DISCOVERED IN THE GALAPAGOS Istanps, Ira L. Wiggins 40 62 UBLISHED QUARTERLY BY THE CALIFORNIA BOTANICAL SOCIETY MaproNo is published quarterly by the California Botanical Society, Inc., and is issued from the office of the Society, Herbarium, Life Sciences Building, University of California, Berkeley. Established 1916. Second-class postage paid at Berkeley. Return requested. Editor — JoHN L. STROTHER Associate Editor — ALAN R. SmitH Department of Botany, University of California, Berkeley 94720 Board of Editors Class of: 1974—-KenrTon L. CHAMBERS, Oregon State University, Corvallis WirtraM A. WEBER, University of Colorado, Boulder 1975—ArTURO GOMEZ-PoMPA, Universidad Nacional Autonoma de México Duncan M. Porter, Smithsonian Institution, Washington, D.C. 1976—DeEnnis ANDERSON, California State University, Arcata Kincs.Ley R. STERN, California State University, Chico 1977—Wi11aM Louis CuLBErson, Duke University, Durham, North Carolina Date M. SmirH, University of California, Santa Barbara 1978—SHERWIN CarLouIsT, Claremont Graduate School LrEstiE D. Gott1ies, University of California, Davis Dennis R. PARNELL, California State University, Hayward 1979—Puitip W. RunpeEt, University of California, Irvine IsABELLE TAVARES, University of California, Berkeley CALIFORNIA BOTANICAL SOCIETY, INC. OFFICERS FOR 1974 President: T. C. Futter, California Department of Agriculture, 1220 “N” Street, Sacramento 95814 First Vice-President: Lestiz D. Gottties, Department of Genetics, University of California, Davis 95616 Second Vice-President: FRANK C. VASEK, Department of Biology, University of California, Riverside 92502 Recording Secretary: CHarLES F. Qurpett, Department of Biological Sciences, Sonoma State College, Rohnert Park, California 94928 Corresponding Secretary: RotF BENSELER, Department of Biological Sciences, Cali- fornia State University, Hayward 94542 Treasurer: MicHAEL F. Baap, Department of Biological Sciences, California State University, Sacramento 95819 The Council of the California Botanical Society consists of the officers listed above plus the immediate Past-President, P. C. Sirva, Department of Botany, University of California, Berkeley 94720; the Editors of Madrofio; and three elected Council Members: GEORGE OBERLANDER, Department of Biological Sciences, California State University, San Francisco 94132 (1972-1974); Jean LancEeNnHEIM, Division of Natural Sciences, University of California, Santa Cruz 95060 (1973-1975); and Joun M. Tucker, Department of Botany, University of California, Davis 95616 (1974-1976). EFFECTS OF PIPELINE CONSTRUCTION ON CREOSOTE BUSH SCRUB VEGETATION OF THE MOJAVE DESERT F. C. VAsSEK, H. B. JOHNSON, and D. H. ESLINGER Department of Biology, University of California, Riverside 92502 Pipeline construction involves trenching, piling, and refilling opera- tions and clearly constitutes a major disturbance of soil and plant cover. The effect on plant cover is one of nearly complete destruction. Revege- tation does occur and its rate probably correlates inversely with the degree of aridity in a disturbed area. However, quantitative estimates of revegetation and recovery rates following pipeline construction have not been made for the Mojave Desert. Accordingly vegetation along a pipeline was surveyed and analyzed to obtain a data base from which to assess the impact of pipeline construction, estimate the indicated recov- ery rate, and suggest the course of secondary succession in creosote bush scrub vegetation. We selected the Southern California Gas Company natural gas pipe- line of 1960 for study because it provides a twelve year recovery refer- ence frame. (Mr. Vlasek of the Southern California Gas Company in Victorville, California, kindly supplied background information on the pipeline). The pipe was laid in a trench 183 cm deep and 152.5 cm wide and then covered over with screened backfill and leveled. The right-of- way includes an access road, the trench line, and the berm. The pipeline segment selected for study travels a utility corridor from Newberry to Lucerne Valley, San Bernardino County, California. We sampled the vegetation in ten areas along a 33.8 km segment of the pipe- line right-of-way. The native vegetation in most of the study area con- sists of creosote bush scrub, which is the characteristic plant community on most of the Mojave Desert except for salt flats and higher mountains. (Plant community and species nomenclature follows Munz, 1959 and 1968.) In each sample area four belt transects were made, each 50 m by 2 m. Two transects served as controls and two transects served to estimate the effects of disturbance. The four transects in each area were located as follows: A) about 50 m east and parallel to the trench line in undis- turbed vegetation, as assessed visually; B) approximately 50 m west of the trench line in representative vegetation and not necessarily parallel to the trench; C) on the berm, that is, along the pipeline right-of-way where trench soil had been piled and then scraped away to refill the trench; and D) directly over the pipeline where trenching had produced the most severe disturbance. (The fourth transect is nearly 48 cm wider than the trench. ) Maprono, Vol. 23, No. 1, pp. 1-64. January 2, 1975. ut 5) MADRONO [Vol. 23 The four transects in each sample area were each read by measuring and recording the distance across each perennial plant. Assuming that, on the average, plants cover a circular ground area, the ground cover area was calculated from the radius for each plant and summed for each species. On the few transects where small perennial herbs were extremely abundant their number was counted in two to four representative 0.5 m? plots and extrapolated to the area of occurrence within transects to esti- mate density and ground cover. The first four areas were sampled in July, 1972, following a relatively dry winter and prior to the full influence of 1972 summer rains. The last six areas were read in November, 1972, following a summer of fairly heavy precipitation and following several early winter storms. The ten sample areas (Table 1) are located sequen- tially southwest from Newberry at distances of 0.0, 4.8, 10.1, 14.6, 19.8, 20.6, 22.9, 26.2, 30.1, and 34.4 km. Direct comparisons among the ten TABLE 1. PHYSICAL CHARACTERISTICS OF SAMPLE AREAS. * Marked change in slope between control transects 1 and 2; ” Evidence of excessively heavy grazing by live- stock; “Evidence of moderate soil salinity; ‘Marked vegetation discontinuities in vicinity. 2 2 € § 5 zs = a. 2 2 a 2 oe S 5 u a) ORS: foe) ou oO oO “4 a 2s g 7 fe ge #8 @ 5 AS as) z 2 |b a eo ® rat - eo §& ae ms A AS AS Ax fa) 1 Broad N.E. Active Alluvium Rocky Moderate 600 wash Gentle erosion 2 Hillside E. Old Bedrock Large to High 670 Moderate erosion small rocks 3. Hillside* N. Stable Residual Small Low 950 Level to rocks moderate 4 Narrow ......... Active Alluvium _......... High 1050 wash* wash 5 Valley Level Stable Alluvium Fine Low 1160 bottom? ° sand 6 Valley S.E. Stable Alluvium Sandy Low 1190 bottom?‘ — gentle 7 Upper N.E. Stable Alluvium Rocky Low 1250 fan” gentle sandy 8 Hillside S.W. Light Residual Rocky Moderate 1310 gentle erosion 9 Lower N.E. Slight Alluvium Sandy Low 1310 fan gentle erosion 10 Middle N.W. Slight Alluvium Gravelly Low 1100 fan moderate erosion sand 1975 | VASEK ET AL.: DESERT VEGETATION 5 sample areas are generally inappropriate because of variation in sub- strate, slope, and aspect (Table 1). Accordingly, primary comparisons are restricted to the four transects within each area. Comparisons among the ten samples are secondary. The four transects were first compared by Jaccard’s coefficient of community similarity (Phillipps, 1959) and then by an index we propose to call a Community Quality Index (CQI). We assume that the eco- logical quality of an area (Suffling et al., 1974) is a product of the com- munity age and the site productivity. Our rationale is based on the premise that consistent, long-term productivity is maximized by stable populations and favorable environmental conditions and that older plant communities are less quickly replaced than younger ones (Suffling et al., 1974). Accordingly, an estimate of productivity, namely ground cover area, and an estimate of relative community age, namely the proportion of ground cover area provided by long-lived species, are integrated by way of their product. As a convenient scaling factor, the square root is extracted to yield: “% Ground covered by % Total perennial CQI = long-lived perennials ground cover The CQI takes into account the successional status of a community and can be applied objectively in assessing ecological quality and poten- tial environmental impacts. Application of the CQI requires assignment of observed species to successional or to functional categories. In the case of creosote bush scrub communities, we recognize four functional groups of perennial plants and assign species to these groups on the basis of our own observations and judgment. The four groups are long- lived perennials, short-lived pioneer shrubs, pioneer perennial herbs, and other perennials. The species observed in our study, together with the number of plants recorded on our study plots, are each assigned to one of the functional groups in Table 2. Long-lived perennials are known or judged on the basis of size or long-term observations (e.g., Shreve and Hinckley, 1937) to live over many decades or perhaps even centuries. In addition, plants of these species generally respond negatively to disturbance. Pioneer shrubs gen- erally are soft shrubs or subshrubs that are known or judged to have relatively short life spans, probably not exceeding one or two' decades. In addition, pioneer shrubs respond positively to disturbance by inva- sion and rapid increase in population size. Pioneer perennial herbs are also active invaders of disturbed sites and develop very large populations. In our area, the occurrence of large populations is irregular perhaps relating to the irregularity of summer rainfall, after which germination typically occurs, or to some other environmental factor. Lastly we recog- nize a group of “other perennials” consisting of perennial herbs and suffrutescent plants whose functional or successional status is not clear. We suspect many are pioneers that grow with winter rains. A MADRONO [Vol. 23 TABLE 2. PERENNIAL SPECIES GROUPED IN Four FUNCTIONAL CATEGORIES AND LISTED IN ORDER OF ABUNDANCE. Nanna ae of Number of individuals individuals recorded recorded LONG-LIVED PERENNIALS PIONEER SHRUBS Ambrosia dumosa (Gray) Payne 113 Hymenoclea salsola T. & G. 5222 Larrea tridentata Gutierrezia microcephala (Sesse & Moc. ex DC.) Cov. 107 (DC.) Gray 9 Hilaria rigida Haplopappus cooperi (Thurb.) Benth. ex Scribn. 95 (Gray) Hall 5 Thamnosma montana Bebbia juncea Greene 4 Torr. & Frem. 73 Eriogonum fasciculatum Ephedra californica Wats. 62 Benth. 2 Opuntia ramosissima Engelm. 22 Viguiera deltoidea Gray 1 Lycium andersonii Gray 22 PIONEER PERENNIAL HERBS Yucca schidigera Euphorbia polycarpa Benth. 5,132 Roezl ex Ortgies 20 Tridens pulchellus Echinocereus engelmannii (HBK) Hitchce. 1,075 (Parry) Ruempl. 7 OTHER PERENNIALS Dalea spinosa Gray 6 Sphaeralcea ambigua Gray 103 Salazaria mexicana Torr. 4 Eriogonum inflatum Krameria parviiclia Benth. 4 Torr. & Frem. 24 Echinocactus polycephalus Dyssodia cooper Gray 16 Engelm. & Bigel. 3 Muhlenbergia porteri Scribn. 13 Opuntia echinocarpa Mirabilis bigelovii Gray 3 Engelm. & Bigel. 3 Allionia incarnata L. 3 Yucca brevifclia Engelm. in Wats. 5 Stephanomeria pauciflora Atriplex canescens (Pursh) Nutt. 2 (Torr.) Nutt. 2 Cassia armata Wats. fd Sarcostemma hirtellum Acacia greggil Gray 1 (Gray) R. Holm. 1 Nicotiana trigonophylla Dunal in A. DC. 1 RESULTS Most of the long-lived plants observed in this study belong to the following species: Ambrosia dumosa, Larrea tridentata, and Hilaria rigida. In addition, Tkamnosma montana and Ephedra californica were commonly observed, but the remaining species of long-lived perennials were uncommon (Table 2). The second category includes short-lived pioneer shrubs that occur sporadically in the creosote bush scrub com- munity usually in washes or other naturally disturbed areas. They also occur as major colonizers in more drastically disturbed areas. Hymeno- clea salsola is the most abundant of these species but other pioneer shrub species contribute slightly to the total number of plants observed (Table 2). The third functional category includes pioneer perennial herbs such as Euphorbia polycarpa and Tridens pulchellus. Both species germinate in great abundance after summer rains. Euphorbia populates sandy, dis- turbed areas and Tridens populates flat, pebbly or rocky areas, particu- larly where run-off might accumulate. The fourth functional category 1975 | VASEK ET AL.: DESERT VEGETATION 5 of perennial plants includes suffrutescent and herbaceous perennials. Most of those observed in our study belonged to Spkaeralcea ambigua and Eriogonum inflatum. In addition, small numbers of Dyssodia coo peri and Muhlenbergia porteri were observed; the remaining species (Table 2) are very sparse. Sphaeralcea and Dyssodia tend to occur on both the disturbed and undisturbed areas so their successional status is unclear. The remaining plants in this category are too few to form a pattern from which a more definitive judgment can be made. The undisturbed vegetation alongside the pipeline right-of-way as well as the regenerating vegetation on the disturbed pipeline right-of-way was recorded in terms of the number and size of its constituent plants. An analysis of those data in terms of total perennial ground cover and the percent of the total cover made up by long-lived perennials, pioneer shrubs, and other perennials is shown in Table 3. In addition, the per- centage of total perennial ground cover contributed by Larrea and Am- brosia is also shown in Table 3 for each of the transects. Plant density generally correlates with ground cover and is not analyzed further. Considerable variation occurs among the ten sample areas. Although the general vegetation in most of the study area can be called creosote bush scrub, the percentage of the perennial ground cover on the control transects varies from O to nearly 94 percent for Larrea and from 0 to nearly 27 percent for Ambrosia. Total ground cover on the control tran- sects ranges from 2.21 to 19.98 m? per transect of 100 m*. Such variations probably reflect local differences in topography, slope, exposure, eleva- tion, substrate, and probably general climate (see Table 1). The area traversed by the pipeline crosses the Newberry and portions of the Ord Mountains. Most of the northern locations are in very rough terrain, and the six northern ones are located in rain shadows of local mountains. The southern four sample areas are located southwest of East Ord Mountain on much gentler terrain and at fairly high elevations suggest- ing better growing conditions or higher site quality. A few more particu- lar characteristics (Table 1) indicate that the vegetation of the ten sample areas cannot be directly compared but rather comparisons should be made among the four transects at each area. First, the two control transects from each area are compared by means of Jaccard’s coefficient of similarity (Table 4). Generally, a similarity coefficient of approxi- mately 0.7 or higher is considered an indication of virtual identity. The two contro] transects in the ten study areas have similarity coefficients ranging from 0.34 to 0.82. In sample areas 1, 4, 7, 8, 9, and 10, the simi- larity coefficients for the two control transects are above 0.64 and we will accept that value as indication of identity. Differences in the physical characteristics within study areas 2, 3, 5, and 6 can account for the dis- similarity between the control transects (Table 1). In sample area 2 the topographic heterogeneity is sufficient to explain that fairly low similarity coefficient. At sample area 3, transect B in the flat to the north of the pipeline should have a higher ground cover than does transect A on the MADRONO [Vol. 23 TABLE 3. ANALYSIS OF PERENNIAL GROUND COVER BY TRANSECT IN TEN SAMPLE AREAS. a =n F © © 7 a 3 3 2 ee Ey, 2 See : 4 Q, Ton ) = ae a= Ke) B= eel ya 3 & es Soo YF S a 5 A, 2 Ov 4 ) known to occur in some species of typical genera of Astereae such as Haplopappus, Petradoria, Bigelowia, Chrysothamnus (Loran Anderson, pers. comm.), and Solidago (Morton, 1968), and in Tracyina there are occasional receptacular bracts interior to the ray florets. One of the most striking differences between the two genera is in the nature of the pap- pus, consisting of 24-40 capillary bristles in Tvacyina and 3-5 awnlike scales in Rigiopappus. The 10-15 pappus bristles of the intergeneric hybrid are intermediate in width. Chromosomal and biochemical similarities between the genera argue for a close relationship. Intergeneric hybrids are vigorous, produce some viable pollen, and show a high degree of chromosomal homology. Both genera have four of six flavonoid compounds in common. The flavonoid produced by one genus but not the other may require only a simple O-methylation of a compound occurring in both species, a chemical trans- formation that could be controlled by a single gene (Harborne, 1967). This evidence from diverse sources leads us to the conclusion that Tvacy- ina and Rigiopappus are very Close relatives that should be placed side by side in Astereae. One could, indeed, use this evidence to argue for the congeneric status of these two taxa. However, the magnitude of the dif- ferences between them in some pappus, achene, and capitular characters is sufficiently great that we believe both taxa should be maintained as distinct monotypic genera. ACKNOWLEDGMENTS This study was supported in part by grants from the National Science Foundation (U.S.A.) and the National Research Council (Canada). We are indebted to D. W. Kyhos for cytological assistance. LITERATURE CITED Brake, S. F. 1937. Tracyina, a new genus of Asteraceae from northern California. Madrono 4:73-77. Boum, B. A., N. A. M. SALEH, and R. Ornpurr. 1974. The flavonoids of Lasthenia (Compositae). Amer. J. Bot. 61:551-561. Cronguist, A. 1955. Phylogeny and taxonomy of the Compositae. Amer. Midl. Naturalist 53:478-511. HARBORNE, J. B. 1967. Comparative biochemistry of the flavonoids. Academic Press, London. HorrMann, O. 1889. Compositae. Jz Engler, A., and K. Prantl, Die natiirlichen Pflanzenfam. IV. 5:87-391. Keck, D. D. 1959. [Treatment of Tracyina and Rigiopappus.| In Munz, P. A., A California flora. Univ. California Press, Berkeley. Morton, G. H. 1968. Chaff on the receptacle of Solidago juncea. Rhodora 70:304—306. Raven, P. H. and D. W. Kyuos. 1961. Chromosome numbers in Compositae. II. Helenieae. Amer. J. Bot. 48:842—850. Ropinson, H. and R. D. Brertrety. 1973. Tribal revisions of Asteraceae. V. The relationships of Rigiopappus. Phytclogia 26:69—70. Van Horn, G. S. 1973. The taxonomic status of Pentachaeta and Chaetopappa with a revision of Pentachaeta. Univ. Calif. Publ. Bot. 65:1-41. HETEROSTYLY, HOMOSTYLY, AND FECUNDITY IN AMSINCKIA SPECTABILIS (BORAGINACEAE) FRED R, GANDERS Department of Botany, University of British Columbia, Vancouver V6T 1W5 Amsinckia is a small genus of Boraginaceae consisting of 10-15 species found primarily in California but occurring naturally north to British Columbia, south to Baja California, east to Utah, and disjunctly in southern South America. Weedy members of the genus have been introduced to Europe, South Africa, central and eastern North America, Alaska and the Yukon, and Australia. Five of the species in the genus are, at least in part, heterostylous, with populations that consist of plants with “‘pin” flowers, with long styles and short stamens, and “thrum”’ flowers, with short styles and long stamens. Heterostylous taxa of Amsinckia are all endemic to California. They appear to be edaphic endemics, restricted to slopes of loose shaly fragments in the South Coast Ranges, except A. spectabilis F. & M., which is restricted to stabilized sand dunes and coastal bluffs. Three of the heterostylous taxa, A. verni- cosa H. & A. var. furcata (Suksd.) Hoov. in Jeps., A. grandiflora Kleeb. ex Gray, and A. douglasiana A. DC., exhibit a pronounced pollen dimorph- ism, with thrum pollen larger than pin pollen, and also possess thrum flowers that are larger than pin flowers. Amsinckia lunaris Macbr. exhibits a peculiar type of heterostyly with a variety of stamen and style length combinations and anthers inserted at two levels in the corolla (Ray and Chisaki, 1957a). All heterostylous taxa in this genus are self- compatible (Ray and Chisaki, 1957a; Ganders, unpublished). The remaining species of Amsinckia, including all the widespread, weedy taxa, are homostylous, with anthers and stigma positioned at the same level in the corolla. Morphological, ecogeographical, and cyto- genetic evidence indicates the homostylous taxa have been derived from heterostylous taxa, which are primitive in this genus. Conspecific homo- stylous and heterostylous populations occur in A. spectabilis and have also been reported in A. vernicosa var. furcata and A. lunaris (Ray and Chisaki, 1957a, 1957b). HOoMOSTYLY IN AMSINCKIA SPECTABILIS The distribution of distylous and homostylous plants in A. spectabilis populations is unusual. Populations may be exclusively distylous, exclu- sively homostylous or contain pins, thrums, and homostyles. Further- more, in mixed populations of distylous and homostylous plants, there is considerable variation in the degree of heterostyly among different plants, that is, the distance separating anthers and stigmas in flowers may vary. There is virtually a complete gradient from pins to homo- styles to thrums (fig. 1). 56 1975] GANDERS: AMSINCKIA ay) 20- NPR 30 ‘ Ms N=100 2 = = ° NIP 30 xe N=75 2 JAL Ne =49 cS LOM | N xe STYLE LENGTH MINUS STAMEN LENGTH (IN mm) Fic. 1. Percentage frequency of stigma-anther separation classes in populations of Amsinckia spectabilis in California. Long styled flowers (pins) are to the left of 0 and short styled flowers (thrums) are to the right of 0. Locations of populations: (distylous populations) NIP—Nipomo, San Luis Obispo Co.; LOM—Lompoc, Santa Barbara Co.; (mixed populations) MBH—Henrietta Street, south side of Morro Bay, San Luis Obispo Co.; BOD—Bodega Bay, Sonoma Co.; CAM—Cambria, San Luis Obispo Co.; MBI—Inyo Street, south side of Morro Bay, San Luis Obispo Co. ; SPR—South Point Reyes Beach, Marin Co.; NPR—North Point Reyes Beach, Marin Co.; JAL—Jalama Beach, Santa Barbara Co.; (homostylous populations) GOR—Goat Rock State Beach, Sonoma Co.; PES—Pescadero State Beach, San Mateo Co. N = sample size; one flower per plant was measured. 58 MADRONO [Vol. 23 Unfortunately the genetics of this situation is not yet known. Ray and Chisaki (1957a) reported that a thrum X homostyle cross gave approxi- mately a 1:1 ratio of thrums and homostyles in the progeny. Such a result is compatible with the interpretation that the homostyle allele rep- resents a crossover of the Primula type, where thrum is dominant to pin. If this is so, it seems likely that other genes and possibly environmental factors are modifying the expression of the heterostyly alleles to produce the variation in degree of heterostyly found in these populations. In mixed populations interbreeding between distylous and homo- stylous plants undoubtedly occurs; progeny from naturally pollinated homostyles contains pins and thrums. It is possible that these popula- tions are evolving toward homostyly, but it is also possible that mixed populations have reached an equilibrium among the forms. Darwin (1877) grew seeds of A. spectabilis after Asa Gray suggested that several species in the genus were heterostylous, but because he found such a range of stamen and style lengths Darwin concluded that A. spectabilis was merely variable. Although prior to the work of Hoover (in Jepson, 1943) and Ray and Chisaki (1957a) the taxonomy of Amsinckia was hopelessly confused, there is no doubt that Darwin’s plants were actu- ally A. spectabilis var. spectabilis, since it is the only taxon in the genus that exhibits this type of variation. Darwin’s record proves that such mixed populations have persisted for at least a century. Homostylous and mixed populations and strictly distylous populations of A. spectabilis differ in several other morphological features and also in geographical range and are best considered infraspecifically distinct. Plants from the distylous populations [A. spectabilis var. microcarpa (Greene) Jeps. & Hoov.]| have all five calyx lobes free to the base, have small mericarps (nutlets) 1.25—1.75 mm long, and the bifurcations of the cotyledons are flat and spreading. Distylous populations are restricted to Late Pleistocene sand dunes (the Orcutt Sand) in southern San Luis Obispo County and northern Santa Barbara County, California. Am- sinckia spectabilis var. spectabilis consists of homostylous and mixed populations, two or three of the adaxial calyx lobes are fused one-half or more of their length, the nutlets are larger (1.75—2.33 mm long), and the bifurcations of the cotyledons are clavate and recurved. Flowers in this variety tend to be smaller and the plants tend to be more spreading in habit, branching from the base. Amsinckia spectabilis var. spectabilts is strictly coastal in distribution and ranges interruptedly from Guada- lupe Island off Baja California to the Queen Charlotte Islands in British Columbia. The two varieties are interfertile and intergrade on the south side of Morro Bay, San Luis Obispo County, California. These popula- tions, arbitrarily referred to var. microcarpa, may contain homostyles (fig. 1) and some plants with fused calyx lobes or a spreading habit. They retain the nutlet and cotyledon characteristics of var. microcarpa. Mixed populations of A. spectabilis var. spectabilis occur only in Marin, Sonoma, San Luis Obispo, and Santa Barbara Counties. Strictly homostylous populations range north and south of the mixed populations 1975 | GANDERS: AMSINCKIA 59 in Marin and Sonoma Counties, California (see fig. 1), and at least for- merly they ranged south of the mixed populations in Santa Barbara County. Only homostyles occur outside of California. The homostylous populations in central California are small and the beaches where they occur have only small areas of suitable sandy habitats. On the other hand, the mixed populations are large and occur at Point Reyes and Bodega Bay where there are large areas of stabilized sand dunes. It might be noted that there are areas of apparently suitable sand dunes, such as at the mouth of Salmon Creek, Sonoma County, where the species has not been found. Geographic distribution of mixed and homostylous populations strongly suggests that (1) homostylous populations were derived by colonization from mixed populations and (2) pins and thrums were selectively elim- inated from the small founder populations. If monomorphic populations were derived solely by genetic drift in founder populations, monomorphic pin or thrum populations would also be expected, but they do not occur. It seems reasonable to hypothesize that large mixed populations are sufficiently conspicuous to attract large numbers of pollinators, so that pins, thrums, and homostyles will all exhibit a high rate of seed set. In very small founding populations pollinator visits to flowers will be erratic and suboptimal for full seed set. If homostyles, because their anthers and stigmas are in close proximity, are capable of significantly higher rates of autogamous seed set than are pins and thrums, they will be at a great selective advantage. In small founding populations, homostyles will replace pins and thrums. MATERIALS AND METHODS To test the validity of this hypothesis, autogamous seed set by the three forms was measured on plants grown in growth chambers. Pollina- tors were excluded, but the ventilation system of the growth chambers provided considerable air circulation, simulating the effect wind might have in nature in shaking pollen from anthers within flowers. Seed set was measured on inflorescences collected from several large and medium sized natural populations where pollinators were presum- ably abundant. Inflorescences were collected from 13 pin and 21 thrum plants at Lompoc; 3 pins, 4 thrums, and 1 homostyle at Morro Bay; 13 pins, 5 thrums, and 28 homostyles at Jalama Beach; 3 pins, 8 thrums, and 7 homostyles at Cambria; 4 pins, 2 thrums, and 9 homostyles at North Point Reyes Beach; and 15 homostyles at Pescadero State Beach. For seed set measurements, homostyles were defined as plants with flowers in which the distance from the center of the anthers to the stigma was 1.5 mm or less. Only one very small mixed population was found during this study, at Atascadero State Beach on Morro Bay in San Luis Obispo County. The population consisted of nine plants in 1971, but seed set data were not obtained that year. In 1972, when seed set was measured, the population consisted of only two plants, one pin and one homostyle. 60 MADRONO [Vol. 23 RESULTS In growth chambers, thrums set 196 out of 1680 possible seeds auto- gamously (11.7 %). Pin flowers set 458 out of 4524 possible seeds (10.1 %). The difference between autogamous seed set in pins and thrums is not significant (y? = 3.09, P > .05). Homostylous flowers set 293 out of 352 possible seeds autogamously (83.2 “). Thus, without pollinator visits, homostylous plants set eight times as many seeds as pins or thrums. Natural seed set was high (80 % or higher) in all forms in large pop- ulations (Table 1). In the distylous population at Lompoc and the mixed populations at Jalama Beach and Morro Bay there was a signifi- cant difference in seed set among the forms. Only at Morro Bay, how- TABLE 1. SEED SET BY DIFFERENT FLOWER FORMS IN NATURAL POPULATIONS OF AMSINCKIA SPECTABILIS. Pin Thrum Homostyle Lompoc, Santa Barbara Co. (var. microcarpa) seed set 1430 2092 seed failed 78 248 total 1508 2340 percent seed set 94.8 89.4 ¥° = 34.12; P < 0.001 Merro Bay, San Luis Obispo Co. (intermediate between var. microcarpa and var. spectabilis) seed set 382 572 226 seed failed 82 144 26 total 464 716 252 percent seed set S26 79.9 89.7 y’ =12.33; P < 0.003 Jalama Beach, Santa Barbara Co. (var. spectabilis) seed set 752 238 1446 seed failed 44 6 46 total 796 244 1492 percent seed set 94.5 97.5 96.9 y= 9.82;P < 0.01 Cambria, San Luis Obispo Co. (var. spectabilis) seed set 221 841 503 seed failed 28 71 33 total 244 912 536 percent seed set 90.6 92.2 93.8 = 2.22;P >01 North Pt. Reyes Beach, Marin Co. (var. speciabilis) seed set 699 354 1270 seed failed 69 30 126 total 768 384 1396 percent seed set 91.0 OFZ 91.0 a0 8 P10) Pescadero State Beach, San Mateo Co. (var. spectabilis) seed set 1226 seed failed 50 total 1276 percent seed set 96.1 1975 | GANDERS: AMSINCKIA 61 ever, were homostyles at an advantage over the other two forms. This population was smaller than other mixed populations studied, occurring in a vacant lot at a housing development in Cuesta-by-the-Sea. It appears that in large natural populations pollinators are sufficiently abundant that all forms have a high rate of seed set. In 1972, the Atascadero State Beach population consisted of one pin and one homostyle. Population size was probably limited by the small area of suitable habitat, by human disturbance, and by abnormally low rainfall in 1972. Nevertheless, the population simulates a small founder population. On inflorescences collected, the homostyle set 30 out of 32 possible seeds (93.8 Yo) whereas the pin plant set 76 out of 136 possible seeds (only 55.9 “). The difference is statistically significant (z = 3.99, Pe 002). DISCUSSIONS AND CONCLUSIONS In large populations where pollinators would be expected to find it advantageous to exploit A. spectabilis, all three forms have very high seed set. Under such conditions homostyles do not have a significant advantage in fecundity over the other two forms. Homostyles, however, are capable of a much higher level of autogamous seed set. Under con- ditions where pollinator activity is erratic or suboptimal, such as may occur in small populations, homostyles wiil be at a strong selective advantage. These conclusions suggest the following model to account for the establishment and geographical distribution of homostyious populations. The model is necessarily a simplification in that it ignores the variation in degree of homostyly. Small founder populations derived from mixed populations, either in nearby habitats of limited carrying capacity (due to the small extent of open, stabilized, sandy areas), or in distant habitats where long dis- tance dispersal results in few colonizers, will evolve quickly toward com- plete homostyly. If, however, a founder population in a habitat of large carrying capacity increases in size quickly enough, pins and thrums may not be eliminated before the selective advantage of homostyly dis- appears. Also, if the founder population is near enough to a mixed source population so that dispersal is more or less continuous, migration will slow the elimination of distylous plants giving the founder population more time to reach the population size at which the selective advantage of homostyly disappears. Geographic distribution of suitable habitats for A. spectabilis is linear and patchy. One would expect populations distant from mixed source populations, where the migration rate is very low (probably close to zero), to be homostylous. Populations established nearby mixed source populations, however, would be expected to remain mixed, if habitat size is large and particularly if migration is also rela- tively high. Nearby founder populations in small habitats, where popu- lation size cannot become large, would be expected to become homo- 62 MADRONO [Vol. 23 stylous unless the migration rate were extremely high, which would be the case only if the habitats were adjacent. The model is consistent with the known distribution of population types. This means that over short periods of a few generations, population size determines the selective value of homostyly. After populations have reached carrying capacity, habitat size, by setting the upper limit to population size, determines the selective value of homostyly. Finally, selection for autogamy in founding populations seems to be a sufficient explanation for the distribution pattern of populations in this species. It is unnecessary to postulate that autogamy is advantageous because it preserves certain closely-adapted genotypes—the “‘fitness” hypothesis first advocated by Mather (1943). This concurs with recent studies of the evolution of autogamy in other annual genera (Lloyd, 1965; Arroyo, 1973). These studies also support the view that autogamy is selected primarily under conditions of unreliable or inefficient pollina- tor availability. LITERATURE CITED Arroyo, M. T. K. 1973. Chiasma frequency evidence on the evolution of autogamy in Limnanthes floccosa (Limnanthaceae). Evolution 27:679-688. Darwin, C. 1877. The different forms of flowers on plants of the same species. John Murray, London. Jepson, W. L. 1943. Amsinckia. Flora of California 3(2) :318-327. Luoyvp, D. G. 1965. Evolution of self-compatibility and racial differentiation in Leavenworthia (Cruciferae). Contr. Gray Herb. 195:3-134. Matu_er, K. 1943. Polygenic inheritance and natural selection. Biol. Rev. Cam- bridge Phil. Soc. 18:32-64. Ray, P. M. and H. F. Cuisaxkr. 1957a. Studies on Amsinckia I. A synopsis of the genus with a study of heterostyly in it. Amer. J. Bot. 44:529-536. . 1957b. Studies on Amsinckia IJ. Relationships among the primitive species. Amer. J. Bot. 44:537-544. NOTES AND NEWS RANUNCULUS FLAGELLIFORMIS DISCOVERED IN THE GALAPAGOS IsLANDS.—No mem- ber of the Ranunculaceae had been reported from the Galapagos Islands by the end of 1970. Then, in June 1971, Paul A. Colvinaux and Charles Racine from Ohio State University discovered plants of a semi-aquatic Ranunculus near the summit of Mount Crocker on Isla Santa Cruz. Upon return to Ohio, Colinvaux sent me a sheet of the material and a slide bearing a preparation of its pollen. An appeal was made to Daniel Weber, who was searching for orchids in the Galapagos Islands at the time, requesting that he look for good specimens of the Ranunculus around the higher flanks of Mount Crocker. He replied that he had already collected buttercups in that area during the previous February and for- warded a duplicate sheet of his specimens. 1975] NOTES AND NEWS 63 Weber’s plants bore several flowers and heads of mature fruits. One flower was soaked and carefully dissected, and mature achenes were examined. A note on Weber’s specimen read “Petals 3” and a check of the flowers available and of a well pressed one on Colinvaux and Racine’s specimen indicated 4 or 5 sepals and 2 or 3 petals. Study of the two collections and comparison of them with material from Mexico and Central and South America showed the plant to be Ranunculus flagelliformis Smith. Many descriptions of this species are brief or inadequate for making a determina- tion of an unfamiliar plant (see Martius, Flora Brasiliensis 13:156-157, pl. 33, fig. IJ, 1864) so the following description and drawings (fig. 1) are presented. RANUNCULUS FLAGELLIFORMIS Smith in Rees, Cycl. 29: no. 13. 1819. Plants peren- nial in shallow water or on muddy margins of pools; primary leaves 10-15 in tufts above slender, white, sparingly branched roots to 10 cm long; petioles erect, 5—20 cm long, 1.5-4.0 mm in diameter, glabrous below, appressed-strigose with upwardly pointing, golden hairs on terminal 2—4 cm of larger leaves; blades deltoid-ovate to suborbicular, 2.0-5.5 cm broad, 2.0-4.5 cm long, basally truncate to shallowly cor- date, appressed-strigose on upper surface near petiole, otherwise glabrous, sub- entire to coarsely but shallowly crenate, 5—9-veined; 1 to several slender stolons spreading 30-50 cm from primary tuft of leaves, at least some of lower nodes rooting and with 1 or 2 small leaves and a single flower at each node; internodes 5-12 cm long; pedicels erect or ascending, 1.5-3.5 cm long, sigmoid, appressed-strigose with golden hairs 0.2-0.5 mm long; sepals 4(5), broadly ovoid, rounded at both ends, shallowly to markedly cucullate, 2.0-2.2 mm wide, 2.8-3.2 mm long, almost mem- branous, pale yellow-green, glabrous; petals 2(3), about 1.0-1.3 mm wide, 2.8-3.0 mm long, pale yellow with tinge of green toward base, obovate, the claw about 0.4 mm wide, 0.5—-0.6 mm long; gland small, covered by a minute transverse scale; stamens few (6-12 in flowers examined), in groups of 2 or 3, filaments 1.0-1.3 mm long, half as thick as broad, strongly incurved; anthers golden yellow, 0.8-1.0 mm long; pollen grains ca 32-38 um long, 21-30 um wide, dark brown, irregularly reticulate-ridged ; fruiting heads 4-6 mm long, 3-5 mm wide, bearing 8-15(20) achenes; mature achenes asymmetrically ovate, slightly flattened laterally, 1.6-2.0 mm long, 0.8-1.0 mm front to back, 0.3-0.5 mm thick, nearly smooth, but with several low, rounded, minute umbos on each side, a blunt apiculation 0.1-0.2 mm long at apex, a narrow ridge along outer, curved margin and a narrower one along the adaxial, nearly straight margin. Specimens examined: Behind Cerro Bandera, Pampa Zone, in a pond, 580 m, (localized) Isla Santa Cruz, Weber 1114, Feb 1971 (DS) ; aquatic, in small pools on saddle between Mt. Crocker and “chimney”, Isla Santa Cruz, Racine and Colin- vaux 60, 26 Jun 1971 (DS). There were minor differences in size and density of pubescence on petioles and blades when the early and late season specimens were compared, but these seemed no more than those shown by many species collected several months apart. Alan B. Franklin reported in June, 1973, that no buttercups could be found in that year. Weber’s characterization of “localized” probably indicates the distribution pattern, but it is also probable that this plant grows only during years of high precipitation. While collecting in anticipation of preparing The flora of the Galdpa- gos Islands (Wiggins and Porter, 1971), I explored the region around Mount Crocker in January-February, 1964, and again in 1967, but found no Ranunculus. Colinvaux (personal communication) reported that studies of sediment cores taken from bottoms of pools on Isla Santa Cruz indicated that pollen from this Ranuncu- lus had been deposited over a considerable period. I thank Colinvaux for telling me of his discovery and for the specimens he gave me. Also, I thank Weber for his generous gift of a beautifully prepared sheet and Alan B. Franklin for searching the higher parts of Isla Santa Cruz in 1973.—Ira L. Wiccins, Department of Biological Sciences, Stanford University, Stanford, Califor- nia 94305. [Vol. 23 ~~ MADRONO 64 (09 xXNDAUYOD PUD aulIDY Wo1y H-F) 9plS Wo1F puv oACqe WOT} ‘suIeIS UdT[Og ‘H pue 5 ‘Jeay ‘A (PII 40q9M Wo} F-W) Jeo] “WA + StayyUR JO sUsISTYap 19}Fe TIMO[Y ‘g {pelo ‘Q {auayoe ainqzeyl ‘gq : qe ‘YW IUWS szmsofijaspy snynounuoy ‘| “SIA Membership in the California Botanical Society is open to individuals ($8.00 per year, regular; $4.00 per year, student). Members of the Society receive Maprono free. 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Subject to approval by the Editors, articles may be published ahead of schedule, as additional pages of an issue, provided the author assumes complete costs of publication. ai Qk M183 VOLUME 23, NUMBER 2 APRIL 1975 Contents GEOGRAPHY OF POLLEN AND CHROMOSOMAL HETEROMORPHISM. IN LEUCOCRINUM MONTANUM (LILIACEAE), Robert Ornduff and Marion Cave 65 THE GERMINATION Lip: A CHARACTERISTIC OF THE LEMMA IN THE PanicEAE, Thomas L, Rost and A. Daniel Simper 68 TAXONOMIC STATUS OF VICIA HASSEI (LEGUMINOSAE), J. Stuart Lassetter 1 New TAXA OF PALAFOXIA (ASTERACEAE: HELENIEAE), B.L. Turner and Michael I, Morris 79 SPONTANEOUS Hysrips IN DODECATHEON (PRIMULACEAE), Joyce F. Goodwin 81 TAXONOMIC STATUS OF CORDYLANTHUS (SUBG. DICRANOSTEGIA) ORCUTTIANUS (SCROPHULARIACEAE), T.I. Chuang and L. R. Heckard 88 A NEw Hyper FLESHY-FRUITED PRICKLY-PEAR IN CALIFORNIA, Malcolm G. McLeod 96 NOTES AND NEWS Two NEw Ptants For NevapA, Jack C. Fisher, Jr. 72 CORRECTION OF THE GEOGRAPHIC DISTRIBUTION OF RHUS MICROPHYLLA (ANACARDIACEAE), David A. Young 78 Continued on back cover Sa Z < = Oo aa fale © pom] = ine scales equal 0.25 mm (Andropogoneae). All 1 Michx nar1us Andropogon ter in 70 MADRONO LVol. 23 Fics. 8 and 9. Germinating florets of Setarza lutescens showing opened germina- tion lid with extended coleorhiza; line scales equal 0.25 mm. TABLE 1. GERMINATION Lip AND FLORET MEASUREMENTS FOR SELECTED MEMBERS or PANICEAE. Size of lid varies with floret size and stage of development. Floret length Lid length Lid width Species (mm) (mm) (mm) Amphicarphum purshii Kunth 2.60 0.37 0.50 Anthaenantia rufa (Ell.) Schult. 2.50 O37 0.37 Anthephora hermaphrodita (L.) Kuntze 3°50 0.25 0.30 Axono pus affinis Chase 1.49 0.24 0.41 A. compressus (Swartz) Beauv. 1.82 0.24 0.33 Brachiaria ophyrodes Chase 2.62 0.37 0.50 B. praetervisa (Domin) Hubbard 2.50 0.50 0.50 B. ramosa (L.) Stapf 2.62 0.50 0.62 B. subquadripara (Trin.) Hitche. 5.00 0.50 0.62 Cenchrus incertus M. A. Curtis 5.00 1.25 0.75 C. myosuroides H.B.K. 2.64 O52 0.39 C. palmeri Vasey 6.25 225 0.75 C. pauciflorus Benth. 6.00 2.00 1.00 Digitaria ischaemum (Schreb.) Schreb. ex Muhl. 1.99 0.24 0.33 D. liniare Kratar 2.00 0.25 0.25 D. pentzii Stent 2.93 0.06 0.16 D. sanguinalis (L.) Scop. 215 O37 O37 Echinochloa colunum (L.) Link 1.08 0.26 0.49 E. crusgalli (L.) Beauv. 215 O25 0.50 E. crus-pavonis (L.) Beauv. 2.76 O23 0.66 E, frumentacea Link 3.00 0.50 0.62 E. haploclada Stapf 2.50 0.50 0.50 E. muricata (Michx.) Fernald 3.50 0.35 0.35 1975 | ROST & SIMPER: PANICEAE Jal TABLE 1. Continued Eriochloa contracta Hitchc. 25a 0.43 0.43 E. gracilis (Fourn.) Hitchc. 2.64 0.39 0.59 E.lemmonii Vasey & Scribn. 2.50 025 0.50 E. montevidensis Griseb. 221 0.59 0.49 E. sericea (Scheele) Munro 2S 0.62 0.62 Leptoloma cognatum (Schult.) Chase 229 0.19 0.16 Neurachne xerophila Domin 5.90 0.50 0.50 Olyra latifolia L. 5.50 1.00 1.00 Panicum agrostoides Spreng. 1:32 0.16 0.26 P. angustifolium Ell. 2.01 O33 O75 P. antidotale Retz. 1.68 0.26 0.56 P. capillare L. 250 0.50 0.62 P. condensum Nash 1.46 0.05 es P. dichotomiflorum Michx. 2.07 O22 0.50 P. hillmanii Chase 212 0.27 0.62 P. lassenianum Schmoll. 0.97 0.45 ON P. miliaceum L. 3.25 0.60 0.90 P. occidentale Scribn. LES) 0:20 0.50 P. pacificum Hitchc. & Chase 1.40 0.25 0.92 P. scribnerianum Nash Snot 0.50 0.87 P. thermale Boland. 1.61 0.26 0.41 Paspalum dilatatum Poir. 2.50 Oly 0.62 P. floridanum Michx. aA) O50 1.50 P.notatum Flugge 2.87 0.25 0.75 P. quadrifarium Lam. VALS O12 OT P. urvillei Steud. Doe, 0.16 0.79 Pennisetum typhoideum (L.) Rich. sa 0.50 0.50 P. villosum R. Br. 10.00 0.49 0.50 Setaria faberi Herrm. 2.08 0.49 0.50 S. geniculata (Lam.) Beauv. 1.90 0.50 0.50 S. lutescens (Weigel) Hubb. Cs 0.50 0.50 S. macrostachya H.B.K. 2.20 0:37 0.50 S. magna Griseb. 1.83 0.33 0.49 S. reverchoni Pilger 2.89 0.37 0.50 S. scheelez (Steud.) Hitchc. 245 OlSi7 0.50 S, Ssphacelata (Schum.) Stapf & C. E. Hubb. 245 0.37 0.50 S. verticillata (L.) Beauv. DENS Ons 0.67 S. viridis (L.) Beauv. 2.38 OAS) O25 Spinifex hirsutus Labill. 10.29 O75) 1.00 Stenotaphrum secundatum (Walt.) Kuntze 4.60 0.39 0.42 Trichachne californica (Benth.) Chase 2.33 0.16 0.33 no structure that could be clearly called a germination lid; Anthephora hermaphrodita (L.) Kuntze on the other hand had a clearly defined lid. On the basis of the presence of a germination lid in the latter species, we would retain Anthephora in the Paniceae, a placement advocated by Stebbins and Crampton (1961). Melinis minutiflora Beauv. does not have a germination lid and should, in our opinion, be once again segre- gated in the Melinideae of Hitchcock (1951). Olyra has been variously placed. Hitchcock (1951) considered Olyra to be in the Paniceae, while Stebbins and Crampton (1961) placed it in 2 MADRONO [Vol. 23 the Oryzoideae, tribe Olyreae. Reeder (1962) examined embryo charac- ters and placed Olyra in the Bambusoideae. We examined O. latifolia L. and observed a large distinct germination lid almost 1 mm square and visible to the naked eye. This observation supports the placement of this genus in the Paniceae of Hitchcock (1951). Twenty-five species of Andropogoneae from 16 genera and one species of Melinideae were examined (e.g., Andropogon ternarius Vasey, fig. 7); none of these Panicoid species has a germination lid. The five remaining subfamilies of the Gramineae were also examined; of 27 tribes and 30 genera none was observed to have a germination lid. The germination lid is, therefore, a feature found in the florets of the Paniceae. This char- acter is suggested as an additional feature useful to determine generic relationships in the grasses. LITERATURE CITED Crampton, B. 1974. Grasses in California. Univ. Calif. Press, Berkeley, CA. Hircucock, A. S. 1951. Manual of the grasses of the United States. 2nd Ed. revised by A. Chase. U.S.D.A. Misc. Publ. No. 200. Keyes, C. E. 1949. Observations on the seed and germination of Setaria italica (L.) Beauv. Trans. Kansas Acad. Sci. 52:474-477. REEDER, J. R. 1957. The embryo in grass systematics. Amer. J. Bot. 44: 756-768. —. 1962. The Bambusoid embryo: A reappraisal. Amer. J. Bot. 49:639-641. Rost, T. L. 1975. The morphology of germination in Setaria lutescens (Gramineae) : The effects of covering structures and chemical inhibitors on dormant and non- dormant florets. Ann. Bot. 39:21-30. STEBBINS, G. L. and B. Crampton. 1961. A suggested revision of the grass genera of temperate North America. Jn Recent advances in botany. Univ. Toronto Press, Toronto, Canada, pp. 133-145. Two New Prants ror NEvADA.—Camissonia cardiophylla (Torr.) Raven subsp. robusta (Raven) Raven is now known from four stations in Clark County, Nevada: Black Hills, crevices in volcanic ridge near crest of mountain (Northwest Boulder City, SE% S16 T23S R63E), 1187 m, 12 Dec 1967, V. Bostick 3225 (Univ. Nevada, Las Vegas) ; south end of Black Hills, steep rocky slopes (Northwest Boulder City, SEY% S31 and SW% S32, T24S R62E), 1000 m, 11 Nov 1970, V. Bostick 5253 (Univ. Nevada, Las Vegas); crevices in basalt cliff, southern exposure (Northwest Sloan, SE% S21 T24S R63E), 1187 m, 15 Feb 1971, V. Bostick 5305 (MO). The range of this plant is hereby extended 129 km east from the Death Valley region, where this subspecies is found in washes at the base of the Panamint, Funeral, Grape- vine, and Argus Mountains. Asplenium resiliens Kunze is now known from one station in Nevada: south- eastern Spring Mountain Range, south fork Pine Creek, north-facing cliff of Navajo sandstone, 1450 m, 16 Jun 1970, J. C. Fisher, Jr.. and G. R, Kennedy sn. (Univ. Nevada, Las Vegas). The range is here extended by 324 km to the west, the nearest location being the mountains about Flagstaff, Arizona. The entire range extends from southern Pennsylvania to Jamaica and Mexico, west through Illinois and Oklahoma to New Mexico and Arizona. It is notable that Pine Creek supports a relictual Pleistocene plant community in southern Nevada.—Jack C. FIsHER, JR., Biology Department, University of California, Riverside 92502. TAXONOMIC STATUS OF VICIA HASSEI (LEGUMINOSAE) J. Stuart LASSETTER Department of Biological Sciences, Eastern Kentucky University, Richmond 40475 The relationship between Vicia hassei and Vicia exigua has been a source of taxonomic confusion. Both taxa are annuals, setting seed by late spring or early summer. Both usually have one or two small incon- spicuous flowers per raceme. Vicia exigua was described by Nuttall in Torrey and Gray in 1838. Vicia exigua of this paper refers only to Cali- fornia and Baja California populations attributed to that species. Vicia hassei was described by Watson in 1890, and Jepson in 1901 reduced it to a variety of V. exigua. More recent floristic workers (e.g., Howell, 1949; Munz, 1959; and Thomas, 1961) have followed Jepson’s concept. MATERIALS AND METHODS Specimens from the following herbaria were examined: ARIZ, BRY, CAS, COLO*, DS, F*, ISC, LA, MEXU*, MICH*, MIN, MO, NY, ORE, PH*, POM*, RSA, SBBG, SBM, SD, SMU, TEX, UARK, UC, US. Asterisks indicate selected material only was studied. Buds collected from field and greenhouse plants were used for anther squashes, and root tips were used for karyotype studies (see Lassetter, 1972, for details). Voucher specimens are deposited at ISC. Field studies included procurement of mass collections as well as observation of pol- linators and habitat characteristics. Plants were not fruiting during the field-work period, but viable seeds were obtained from herbarium specimens. RESULTS AND DISCUSSION Examination of mass collections and herbarium specimens revealed two qualitative characteristics in V. assez that are absent in V. exigua. Stylar pubescence in V. assez is very dense and concentrated on the lower side of the style; this arrangement of hairs is termed the stylar brush. Secondly, the ovaries and legumes are pubescent (fig. 1). Vicia exigua has much sparser pubescence evenly distributed around the style (and, therefore, no stylar brush) and has glabrous ovaries and legumes (fig. 1). In addition, leaflet apices of V. Aassei are mostly truncate or often emarginate with two or three mucros, whereas V. exiguva leaflets are mostly acute or obtuse. These latter characteristics, however, are not wholly consistent, especially in young or stunted specimens of V. /assez. Chromosome numbers in both taxa are 2” =7 IT, and no meiotic irregularities were observed (Lassetter, 1972). All root-tip preparations counted were 2” — 14. These numbers are the first reports for V. exigua Nutt. (sensu stricto) and V. hassei. 73 74 MADRONO [Vol. 23 V. HASSEI V. EXIGUA Fic. 1. Ovary apices, styles, and stigmas of Vicia hassei and V. exigua. Karyotypes of V. Aassei and V. exigua are given in Figure 2, and they obviously are different, as is most often the case in Vicia species (Svesh- nikova, 1927; Shrivastava, 1963; Mettin and Hanelt, 1968). The largest two chromosome pairs in V. assez are metacentric, with secondary con- strictions very near the centromere. In V. exigua, no secondary constric- tions were observed, and the smallest pair of chromosomes is satellited. Chromosomes of V. Aassei are larger (fig. 2). Field study revealed little difference in habitat preference between the two taxa, and both may be found in close proximity. Populations occur on a variety of soils, ranging from very sandy to rocky, but only V. hassei was found on fine-textured clay “‘dry-bog”’ soils. Pollinators were not observed visiting flowers of these taxa. Bagging of greenhouse plants showed both taxa to be self-fertile, and bagging of single flowers indicated that the stigma receives pollen from anthers of the same flower. Because pollination usually occurs just before or some- times at the time the flowers open, pollinators, even if present, probably would have little effect on intrapopulational cross pollination. Prolific fruit and viable seed were produced by several generations of each taxon under greenhouse conditions. VEXIGUA JE Ge Og gy Ba xa i 0pm V. HASSE! Ny} i¢ I (f i KX Fic. 2. Karyotypes of V. exigua (voucher grown from seed collected by J. Marin sm., ISC) and V. hassez (voucher grown from seed from DS 401987). 1975] LASSETTER: VICIA 75 Distributions of these taxa, determined from my own collections and herbarium data, are given in Figure 3. Vicia hassei extends from Baja California to Southern Oregon and is not found farther east than the San Joaquin Valley. Vicia exigua is almost wholly restricted to areas south of Los Angeles except for one collection from Shasta County, California (Eastwood s.n.,in 1912, US!). Vicia hassei is strictly a west coast taxon, while V. exigua is the westernmost extension of a group of interrelated taxa of the Vicia ludoviciana complex (V. ludoviciana Nutt., V. leaven- worthiu T.& G., V. exigua Nutt., and their infraspecific taxa). Relation- ships of this Texas-centered complex will be presented in a later publi- cation. V. HASSE] V. EXIGUA Oo Fic. 3. Distribution of V. hassez and V. exigua. 70 MADRONO LVol. 23 CONCLUSIONS AND TAXONOMIC TREATMENT On the basis of the aforementioned morphological, karyotypic, and distributional differences, I believe it most realistic to consider these taxa as separate species. VICIA HASSEI S. Wats., Proc. Amer. Acad. Arts 25:129. 1890.—Vicia exigua Nutt. var. Hassei (S. Wats.) Jeps., Fl. W. Mid. Calif. 296. 1901. Watson examined and cited several specimens from different collectors to formalize a species concept of V. assez. From his cited material that I examined, I designate as Lectotype: hills about Los Angeles with V. exigua, H. E. Hasse s.n., 1888 (GH!). Other cited specimens that I examined are: Santa Cruz, C. L. Anderson s.n. (Syntype GH! Isosyntype PH!); Benecia, Bigelow s.n. (Syntype GH! Isosyntype NY!). Vicia exigua Nutt. var. ?californica Torr., Pacific Railroad Reports 4:76 1856. Typr: Benicia, Bigelow s.n. (the same collection cited by Wat- son for V. hassei). Holotype GH! Isotype NY!. Herbaceous annual, usually robust, but sometimes flowering when less than 10 cm tall, short and stubby-branched to 60 cm or more tall and climbing. Leaflets (2) 4-8 (12), (1.5) 2.0-6.0 (9.0) mm wide and (8.0) 14.2-27.1 (40.0) mm long, the length-width ratio (1.9) 3.2-7.1 (9.0), the apices usually truncate to retuse or emarginate, often with more than one mucro. Flowers 1-2, arising at separate points on the peduncle, white to faint bluish or lavender, the length from calyx base to the tip of the unreflexed standard (6.2) 6.7—7.9 (8.8) mm, calyx teeth subequal, about 1.0 mm long or less, about half or less the length of the calyx tube, the tube (1.6) 2.0-2.6 (2.9) mm long. Stylar pubescence a dense mass, obvi- ously concentrated on the lower side, the hairs 0.25—0.45 mm long. Ma- ture legumes 24-38 mm long, saber-shaped at the tip, the tip curved upward, the internode between two legumes great, 13.0-26.0 mm long. Ovules up to 10. DISTRIBUTION: Southern Oregon to Baja California, near the coast and on some offshore islands (fig. 3). Lower elevations and near sea level, often among undergrowth and brush, grassy hills and slopes; canyons, arroyos, and ravines; creeks and flood plains; forest margins; only occa- sional on rights-of-way. In sandy to rocky and “dry-bog” clay soils. Flowering in March-April, fruiting in April-May. Victa Ex1cuA Nutt. in T. & G., Fl. N. Amer. 1:272. 1838.—Cracca exigua (Nutt. in T. & G.) Alefeld, Bonplandia 9:119. 1861. Type: Columbia plains, Nuttall s.n., (Isotype PH!). The holotype of V. exzgua (BM) was collected by Nuttall from ‘the Columbia Plains”, but was not available for this study. A photograph was seen, however, and the type sheet contains two specimens. Howell in 1935 expressed some doubt as to the collection site of each specimen by the following note attached to the type sheet: 1975] LASSETTER: VICIA 77 From the type description of Vicia exigua Nutt. (T. & G., Fl. N. A. 1:272) it is obvious Nuttall saw plants from California as well as from the Columbia. I believe that the two specimens on this sheet are from these two regions and that the California label has been lost. From the type description it is easy to determine that the left-hand specimen is the one from “the Oregon,” the one on the right from “Upper California.” 8/26/35 John Thomas Howell. The right-hand specimen is definitely V. exigua, and I here designate it the lectotype. If this specimen were in fact collected from the Colum- bia Plains (Oregon) or even from northern California, as Howell be- lieves, it must rank with Eastwood’s Shasta Co. collection of V. exigua as an outlier from the more common southern California locations. From the type photograph, I cannot positively identify the left-hand specimen. Herbaceous annual, diminutive to robust, 16-110 cm tall, sprawling to erect, climbing if support is available. Leaflets (4) 6-9 (12), (0.9) 1.6-3.4 (6.3) mm wide, and (9.0) 12.0—25.0 (37.0) mm long, the length- width ratio (3.6) 5.8-11.2 (16.4), the apices acute to truncate. Flowers 1-3, rarely 4, occurring singly at points on the peduncle (but almost con- tiguous), or 2 flowers arising from the same point, bluish, the length of the flower from calyx base to the tip of the unreflexed standard (4.3) 4.4-6.5 (7.5) mm. Upper calyx teeth usually slightly shorter than the lowest tooth, the lowest tooth (0.9) 1.1-1.5 (1.7) mm long, usually more than half the length of the calyx tube, the tube (1.3) 1.6-2.0 (2.2) mm long. Stylar pubescence more or less evenly distributed around the style, the hairs 0.08—0.15 mm long. Mature legumes 15—26 mm long, oblique at both ends, the 2 or more legumes contiguous or nearly so. Ovules up to 7. DISTRIBUTION: Southern California and Baja California; one distant collection from northern California seen (fig. 3). Wooded areas; moist slopes, ravines, and canyons; foothills; along creeks; chaparral; beaches; dry wasteland. Sandy to rocky soil. Flowering in (February-March) April-May; fruiting in (February) April-May (June). ACKNOWLEDGMENTS I express sincere gratitude to Dr. Duane Isely for his suggestion of the initial dissertation problem and his guidance throughout this study, which represents a portion of a dissertation submitted as partial fulfillment of the requirements for the Degree of Doctor of Philosophy in Botany, Iowa State University, Ames, 1972. Journal Paper No. J-7850 of the Iowa Agriculture and Home Economics Experiment Station, Ames, Iowa. Proj- ect 1814. The facilities of the Iowa State Herbarium, supported by the Science and Humanities Research Institute, were used in the preparation of this paper. The Iowa State University Agriculture and Home Econom- ics Experiment Station provided funds for laboratory supplies and some field travel. Field work was supported by N. S. F. Grant GB-27935. Ap- preciation is extended to the curators of the herbaria mentioned herein 78 MADRONO [Vol. 23 for their courtesy in arranging loans and in permitting my examination of specimens, and to Clare B. Hardham and the late Ernest C. Twissle- mann for allowing examination of their personal collections. Ingrid Marin collected viable V. exigua seed from Riverside Co., California. LITERATURE CITED Howett, J. T. 1949. Marin flora. Univ. Calif. Press, Berkeley. LASSETTER, J. S. 1972. A biosystematic study of the Vicia ludoviciana complex (Leguminosae). Ph.D. Thesis. Iowa State Univ., Ames. MettTin, D., and P. Hanett. 1968. Bemerkungen zur Karyologie und Systematik einiger Sippen der Gattung Vicia L. Feddes Repert. 77: 11-30. Mownz, P. A. 1959. A California flora. Univ. Calif. Press, Berkeley. SHRIVASTAVA, L. M. 1963. Cytogenetical studies in certain species oi Vicia. Cytologia 28:154-169. SVESHNIKOVA, I. N. 1927. Karyological studies on Vicia. Trudy Prikl. Bot. 17-1, No. 3:37-72. Russian with English summary. Tuomas, J. H. 1961. Flora of the Santa Cruz Mountains of California. Stanford Univ. Press, Stanford, Calif. CORRECTION OF THE GEOGRAPHIC DISTRIBUTION OF RHUS MICROPHYLLA (ANA- CARDIACEAE) —Barkley (Ann. Missouri Bot. Gard. 24:256-500. 1937) in his mono- graph of North American Rhus included a distribution map (pg. 388) of Rhus microphylla Engelm. ex Gray showing its geographic distribution as including Cedros Island and Puerto San Bartolome, Baja California, Mexico. Cedros Island and the adjacent coast were also shown as localitites for R,. microphylla by Shreve and Wiggins (Vegetation and flora of the Sonoran Desert, Stanford Univ. Press. 1964) and Hastings, Turner, and Warren (An atlas of some plant distributions in the Sonoran Desert, Univ. of Arizona. 1972). Hastings et al. pointed out that this rather formidable disjunction is puzzling, since R. microphylla is primarily a Chihuahuan Desert species, and that they had not seen specimens from either locality. In spring 1972 and 1973 I specifically searched for R. microphylla on Cedros Island in areas that seemed suitable for its growth. I was unable to locate it, but in all these areas there was an abundance of Pachycormus discolor var. veatchiana (Kell.) Gentry. Curiously, the disjunct localities reported for R. microphylla in the Sonoran Desert fall within the known distribution of P. discolor var. veatchiana. Because of this and the superficial resemblance of these two species, I borrowed specimens (Cedros Island, 8 Dec 1888, Pond s. n., and Port San Bartolome, 27 Nov 1889, Pond s. n.) from the University of Notre Dame on which this distribution of R. microphylla is based. Although both specimens are sterile branches, they are easily recognized as P. discolor and not R. microphylla, since the leaves are borne in fascicles (Kellogg, Proc. Calif. Acad. Sci. 2:24. 1860). Also, on the specimen from Cedros Island there is a card written by Lt. Pond stating, “low shrubby bush 18 inches high, four feet across, branching at the ground into four stems, two inches each in diameter, bark peeling, single stem from a tree like above six feet high, six inches thick.” The thick stem and peeling bark are both characteristic of P. discolor but not R. microphylla. I have annotated these specimens as Pachycormus discolor var. veatchiana. Oddly, they were not annotated by Barkley as R. microphylla. Rhus microphylla is restricted to the southwestern United States and northern main- land Mexico, but it does not occur in Baja California. I thank Dr. T. Crovello of ND for the loan of the Pachycormus specimens.—Davip A. Younec, Rancho Santa Ana Botanic Garden, Claremont, California 91711. NEW TAXA OF PALAFOXIA (ASTERACEAE: HELENIEAE) B. L. TuRNER and MicHaAeEL I. Morris Department of Botany, The University of Texas, Austin 78712 During preparation of a monograph of Palafoxia, we discovered new taxa that are confined to the deserts of southwestern United States and Mexico, typically in sandy soils at low elevations. Palafoxia arida B. L. Turner & M. I. Morris, sp. nov. Herbae annuae 10-70 cm altae; caules erecti saepe omnino divaricate ramosi scabrosi vel aspero-hispidi raro fere glabri caulibus superis saepe conspice pubescen- tibus trichomatibus glandularibus. Folia mediocaulina linearia vel lancei- linearia 20-100 mm longa 2—8 mm lata apice gradatim angustata utrinque canescenti-scabrosa; petioli mediocaulini 5—20 mm longi. Capitulescentia subcorymbose cymosa. Capitula 5-40 subturbinata vel fere cylindrica 5-10 mm diametro 20—28 mm alta (floribus exsertis inclusis) 9—20-flora; pedunculi 1—5(7) cm longi; phyllaria principales (6) 7—15 linearia 10—20 mm longa 1-2 mm lata scabra vel dense glandulosi-pubescentia saepe dorsaliter aliquantum carinata. Flores centrales actinomorphi; corolla 9-11 mm longa rosei-alba vel rosea tubis 2-4 mm longis faucibus cylin- draceis 6-8 mm longis lobis 1-2 mm longis; rami styli 4-5 mm longi. Achenia linearia 10-15 mm longa faciebus 4 dense vel sparse adpressi- pubescentibus; pappi squamae costis prominentibus; achenia intima supra angulos squamis 4 linearibus acutis 8-12 mm longis supra facies squamis 4 brevioribus abortivis; achenia extima squamis 3—8 inaequales vel nullis. Chromosomatum numerus, 7 = 12. Type: United States, California, San Bernardino Co., ‘““The Needles”, 7 May 1884, M. E. Jones 3849. Holotype: US! Isotypes: ARIZ! CAS! DS! F! NY! UC!. The following voucher specimens, which are deposited in TEX, have a chromosome number of m = 12: Arizona, Mohave Co., Turner 4787; California, Riverside Co., Powell & Sikes 1383; and Mexico, Baja Cali- fornia, Powell & Turner 1705. Palafoxia arida is named for the arid climate in which it grows. The species is widespread in sandy soils throughout the Mojave, Colorado, and Sonoran deserts of southwestern United States and Mexico. Recent taxonomists (Munz, P. A. 1959. 4 California flora.) have re- ferred to this species as Palafoxia linearis. Nevertheless, this name applies to a related, allopatric taxon occupying coastal sand dunes of southern Baja California. Palafoxia arida can be easily distinguished from P. linearis in that plants of the former are erect tap-rooted annuals possess- ing linear-lanceolate leaves with acute apices, whereas plants of the latter are suffruticose, sprawling shrublets having linear leaves with round or obtuse apices. 79 80 MADRONO [Vol. 23 Palafoxia arida var. gigantea (M. E. Jones) B. L. Turner & M. I. Morris, comb. nov. Palafoxia linearis (Cav.) Lag. var. gigantea M. E. Jones, Contr. W. Bot. 18:79. 1933. Tyrer: United States, California, Imperial Co., sand dunes, W of Yuma, 24 Sep 1931, M. E. Jones 28599. Holotype: POM! Isotypes: MO! UC!. Palafoxia arida var. gigantea can be distinguished by its more robust habit (ca 0.9-1.5 m tall), primary stems 0.5—1.0 cm thick, and heads 28-35 mm long. Variety gigantea is endemic to the dunes of southeastern Imperial County, California (just west of Yuma, Arizona). Palafoxia linearis (Cav.) Lag. var. glandulosa B. L. Turner & M. I. Morris, var. nov. A P. linearis var. linearis caulibus foliisque scabro- sissimis glandulari-pubescentibus; pappi squamis abortivis brevioribus differt. Plants perennial, 40-80 cm high, up to 150 cm across; stems suffruti- cose and branched from the base, forming conspicuous clumps, rather evenly pubescent with stiff, appressed white hairs, densely covered with rough glandular pubescence; leaves simple, succulent, alternate; mid- stem leaves lance-linear to nearly obovate, 25-50 mm long, 3-8 mm wide, with petioles 3-8 mm long, blades rather abruptly terminated by an obtuse or rounded apex (very rarely nearly acute), canescent-scabrous on both surfaces, with dense covering of glandular pubescence; inflores- cence a subcorymbose cyme with 3—15(20) heads; heads subturbinate to nearly cylindric, 5-10 mm across, 20-22 mm high (including the project- ing flowers), 10-20 flowered, on peduncles 1.5—5.0 cm long; principal phyllaries 8-14, linear, 10-15 mm long, 1—2 mm wide, scabrous-pubescent, especially below (rarely somewhat glandular); florets “white with faint tinge of magenta-pink” on lobes, regular (outer florets becoming zygo- morphic) ; corolla 7-10 mm long, tube 2-3 mm long, throat cylindric (in outer florets somewhat flaring), 5-7 mm long, the lobes 1-2 mm long; style branches 4-5 mm long; achenes 7-11 mm long, linear, 4-sided, densely appressed-pubescent; pappus scales 4-8, unequal, with pro- nounced midribs, the inner florets normally with 4 linear, acute scales, 6—9 mm long, on the angles, these alternating with 4, much shorter, abor- tive scales. Chromosome number not determined. Type: Mexico, Baja California, mouth of arroyo along beach at Barril, 48 mi E of Pozo Aleman, 2 Mar 1935, /ra L. Wiggins 7825. Holotype: DS! Isotypes: F! GH! UC! US!. Palafoxia linearis var. glandulosa is named for the rough glandular pubescence that densely covers the leaves and mid-stems, a characteristic that distinguishes it from var. linearis, which lacks glandular trichomes on these parts. The taxon is restricted to coastal sand dunes of eastern Baja California from latitude 26°30’ N to 29°30’ N while var. linearis occupies similar but more southern sites. We acknowledge, with thanks, Professor M. C. Johnston for providing the Latin descriptions. This paper was supported by N.S.F. grant 29576X. SPONTANEOUS HYBRIDS IN DODECATHEON (PRIMULACEAE) Joyce F. GoopwIN Department of Life and Physical Sciences Allan Hancock College, Santa Maria, California 93454 When geographically distinct species grow in unusually close proxim- ity, interspecific hybridization may occur and a hybrid swarm may be produced. This appears to be occurring with two species of Dodecatheon on Table Mountain in Butte County, California. One of these, D. cleve- landii Greene ssp. patulum (Greene) H. J. Thomps. is growing out of its usual elevational range, very near a population of D. hendersonu Gray ssp. parvifolium (Kunth) H. J. Thomps. Certain plants in the D. cleve- landii population are of unusual morphology and coloration for this spe- cies and are hypothesized by the author to be hybrids. In his review of Dodecatheon, Thompson (1953) included an analysis of artificial inter- specific hybridization. He demonstrated that a cross producing vigorous hybrids is possible between these two species. A relatively high degree of morphological similarity between two spe- cies, as is exhibited by D. hendersonu and D. clevelandu, may make morphological investigation alone inadequate for delineation of taxa, in- cluding hybrids. This study has utilized chromatography in addition to morphology in analyzing putative hybrids and their parentage. MATERIALS AND METHODS The study site is located on Table Mountain, 5.1 km northeast of Cherokee Road from its intersection with Table Mountain Road in Oro- ville, Butte County, California. Two distinct habitats are represented by an open, grassy pasture adjoining an oak woodland. Dodecatheon cleve- landii (ca 300 plants) grows in the meadow, in full sun. At this site, it is out of its usual elevational range; it is normally confined to grassy mea- dows on the Sacramento Valley floor (ca 67 m), whereas on Table Moun- tain it is found at a much higher elevation (ca 260 m). At this site, D. hendersont (ca 200 plants) is confined to the more shady oak woodland, which is its normal habitat. The putative hybrids are found within the D. clevelandu population and near the ecotone area, also in full sun. Other populations of these two species were also studied in Butte County for morphological and chemical comparison with the Table Mountain plants (see Goodwin, 1973). Morphological analyses were made of approximately 30 plants of each species and of nine of the dozen or so putative hybrids found during the study. Putative hybrids are designated A, B, C, 1a, 1b, 2a, 2b, 3, 4, 6, and 7. Chromosome counts for the two species (four plants each) were made from propiono-carmine squashes of anthers fixed in ethanol:acetic acid 81 82 MADRONO [Vol. 23 (3:1, v:v). These slides were made permanent and have been retained by the author. Fresh pollen from ten plants of each species and six putative hybrids was subjected to aniline blue-lactophenol stain to indicate per- centage of presumably viable pollen; 500 pollen grains per plant were counted. Hauser and Morrison (1964) note that results of this technique tend to give results of slightly higher percentages than the nitro-blue tetrazolium technique they employed, and which they feel to be more accurate. However, the differences in the results of these two techniques were consistently small in their work, and the aniline blue-lactophenol technique was chosen because of its simplicity. Voucher specimens for this study have been deposited in the herbarium of California State Uni- versity, Chico. Thin-layer chromatography was used to separate compounds in leaf extracts. Dried leaves from pressed plants were soaked for 18—24 hours in methanol with 0.5 percent HCl. This extract was spotted on silica gel thin layers (SilicAR TLC-7G, Mallinckrodt). Chromatographs were developed with solvents in two phases: the first phase consisted of t-buta- nol, acetic acid, and water (3:2:2, v:v). The second phase consisted of acetic acid and n-propanol (1:3, v:v). Each phase was run for 10 cm. Plates were dried, and then exposed to ultraviolet light to make non- pigmented spots fluoresce; they were then treated with ammonia vapor and again exposed to ultraviolet light. RESULTS Morphological characteristics of both species and the putative hybrids are summarized in Table 1. Morphologically, all D. clevelandi popula- tions studied were very similar. Individuals with pink-tipped corollas were very rare (only three were found in the entire study) and resembled the individuals with white corollas in all other respects. The D. hender- soni populations were also very similar, with the exception of corolla coloration. Some populations had a high frequency of the darker magenta color, while others had more individuals with the lavender coloration. Only one totally white individual was found during this study; this plant resembled other plants in that population in all other respects. Putative hybrid plants were distinguished by their pink and white corollas, the condition of the maroon band on the corolla tube, and the absence of the yellow anther dot (see Table 1). No individuals in any D. hendersoni population showed the two-toned corolla coloration and maroon band found in the putative hybrids, and no individuals in any D. clevelandi population studied showed the absence of the yellow anther dot. For these reasons, the putative hybrids were rather easy to distinguish once they had flowered. Morphological variation of the Table Mountain plants is summarized in Figure 1. Chromosome counts of both species established that plants with a haploid number of ” = 22 are present in both species at the Table Moun- 32) — if GOODWIN: DODECATHEO 1975] quasqy a[qeHe A YIPIM s[qeuea -S1IY4JUR WOIT ABMP ‘payudpuy yuid Aj[e}0} ysowye 0} paddij-yurg 6'T “BAR ‘e-] Sd asd CTLIEZ AS Of d[qUIasad 0} spud} Safqeiie A 6 IT “BAe ‘WD ¢'gI—-S'g yuasaig opIMm Ayenby dpIM -S1OY4JUR WOT} ABMB ‘poyudpUyT peddi-yurd Ajares ‘ureatd 07 ayIU AA ZZ BAB “PT 9} e[OIDUPTGO 0} a}e[N}eds 16 ‘BAR “UID FI-9 quasqy TIMOLIEN MOAIVU £S19YUP pieMo} Sutjutod ‘pojuspuy dIYM ApIIeI ‘eJUISeCU 0} JOPUIAR’T 6¢ SAe*/-] INdT]IJa 0} a}e[nNyedS Pl ‘sAe ‘UID TE-O1 spriqéy aaneyng mpunjanay9 “qq Uosdapuay — SIoyyUe JO aseq je JOP MOT[AA [JOYM J9y}Ue 0} yoodsor YIM YPIM oqn}-jUIWIE[L 9qn}-e] [0109 uo pueq UOOIeW JO a1N}JeN UOTJVAO[OI JUIWIBIS-BI[OIOD Jequin tad s1aMmop ‘ON Asopoydiow Jeo] 1ystIey edeos AdPIeICY) (SOTPBIOT LO} “C167 ‘UIMpoor) vas) AjJUNOD a}jng ur suoTY[Ndod [e1aAas WoAT VUTeD salads ay} 10y sajdues ‘SATMAAH AATLVLINd ANY NOFHLVOIGOT AO SATOAMS OMY, JO SNOSTYVAINOD TWOIDOTOHANOT, “LT AITAV, 84 MADRONO [Vol. 23 Root bulblets present Degree of pink on petals Yellow anther dot present 2 Number of flowers « Putative hybrids 6 D. hendersonii ~ D. clevelandii ®: + 28 O \ 26 24 20 oO BR Scape Height Ccm) ox Ss S 10 2.0 3.0 40 Average Leaf Length/Width Ratio Fic. 1. Scatter diagram of population of Dodecatheon at the Table Mountain study site. Root bulblet data were not available for the putative hybrids other than plants A, B, and C, since measurements were made on living plants. Plants A, B, and C represent earlier collections, for which chromatographic data are not available. tain site. This is in accordance with Thompson’s (1953) work on this genus. It was not necessary for this study to establish that all plants of each species had m = 22; it was necessary only to establish that such 1975] GOODWIN: DODECATHEON 85 individuals were present at the Table Mountain site, since hybridization producing viable offspring is possible in this genus only between plants with the same chromosome number (Thompson, 1953). Pollen stainabilities indicate that both species are highly fertile. Dode- catheon hendersonii was consistently high, with an average of 98.8 per- cent (SD + 0.66), while D. clevelandii was somewhat lower and more variable (average 90.5 percent; SD + 7.2). Percent stainabilities for the putative hybrids were 58 (plant la), 63 (1b), 8 (2a), 8 (2b), 82 (3), and 87 (7). Plants 4 and 6 were not sampled in this respect. These results correlate well with the observed frequency of collapsed or abortive grains. Drawings of chromatographs are shown in Figure 2. Some spots were consistently present for both species, some consistently present for just one species, and some were inconsistent, both in their presence and their species specificity. Some spots were pigmented, while others were visible only under ultraviolet light. For technical reasons, chromatographic data are not available for putative hybrids A, B, C, 2a, and 2b, nor morpho- logical data for plants 4 and 6. DISCUSSION In general, the putative hybrids tend to resemble D. clevelandii more than D. hendersonii, especially with regard to scape height and leaf shape (aig. 1): Dodecatheon hendersonii and D. clevelandii have high pollen stain- ability and, hence, presumably high viability. They would not, therefore, be expected to have arisen recently as a result of hybridization. Putative hybrids, however, have significantly lower percentages, which supports the hypothesis of hybridization. Plants 1a and 1b, because of their simi- lar morphology and adjacent growing sites, are thought to represent a clone, as do plants 2a and 2b. Pollen stainability results indicate that this is indeed the case. The similar results for each pair would indicate a genetic basis for the sterility, as would be expected in hybrid plants, rather than environmentally induced sterility (frost, nutrition, etc.). Results of chromatography show that these two species have similar chromatographic profiles (fig. 2, A and B). There are a few species-spe- cific compounds, but the majority of the compounds are shared by both, indicating that these species are closely related genetically. Dodecatheon hendersonit was found to be more variable between populations with regard to leaf compounds than was D. clevelandii, which was quite con- sistent chemically between populations. This may indicate a greater degree of genetic heterogeneity among populations of D. hendersonii in Butte County. Chromatographs of all the putative hybrids tested (fig. 2, C—F) showed the presence of at least one species specific compound from each species, in addition to compounds found in common with both spe- cies. The most probable way that these plants could have received such a combination of compounds is through hybridization between the two species. One of the species-specific compounds of D. clevelandii does not 86 MADRONO [Vol. 23 | 7 : s ss ae C—O O CLD d fy | oe QO L G \ 13 AY 1 il e 19) oe © a A NO a) Q S ee © “ohase | ———> . NY A DO hendersoni! ssp. parvifolium B D clevelandii ssp patulum Table Mountain population | | a. ; ean See CB