Pa a , 0 Se ivad Can ge dat an AVINo ae Ug, tu a ¥. oti y \ “ rt + Mn ay en ’ ’. mt = , $ on # i ‘ - ‘ : a 5 = \ . ; ! o ~ - . . A fo + Y « ais | - , 1 ‘et | 7 i : i ¥ h n ¢ fi 1 _ . —_ i 7 » 7 v4 ! Ps ty i = a i j . = — * - aa _ Rog —— ‘ f , 7 ' _ . : 2 a ' ! = 7 : 5 - : : : : ter ' E i . * - 4 : ar . z ? cha = ‘ . - rt : \ : ’ x . f : . x : ' 7 z ee. f : ; : F ° a ' 7 é . im ' bee, u 7 e . 7 7 ie fe i p : Wey : ' = 7 = i . i 1 , re - ‘ 1 A ‘ ‘ a > . i . = ! s 4 © ' 3 % - nl é : @ - 5 - . - ' * ' , y *. 1 . Fi i ° “ 7 - e ul ) 5 rar As ; ; he =i . i A ti = “ - i ms f - _ a i" a ’ ' , . Ci - i 28 iy = 4 : a : ‘ : - i : ? jE ‘ he ' ; : ib by - 1 iw i y ‘ Th Louis Fornraurt HENDERSON Frontispiece ta Volume 5, Manrono. MADRONO A WEST AMERICAN JOURNAL OF BOTANY VOLUME V 1939 - - 1940 Published by the California Botanical Society Inc. 4004 Life Sciences Building, University of California, Berkeley North Queen Street and McGovern Avenue, Lancaster, Pennsylvania Board of Editors H. L. Mason, University of California, Berkeley, Chairman. L. R. Aprams, Stanford University, California. Lincotn Constance, University of California, Berkeley. H. F. Coretann, Sacramento Junior College, Sacramento, California. A. W. Haupt, University of California, Los Angeles. Secretary, Editorial Board Ernet Crum, Department of Botany, University of California, Berkeley. Business Manager Davin D. Kecx, Carnegie Institution of Washington, Stanford University, California. To Louis Forniquet Henderson, who celebrated his eighty- seventh birthday on September 17, 1940, this, the fifth, volume of Madrofio is respectfully dedicated. He is the only living representative of that small coterie of western botanists who contributed so largely to the work of Asa Gray and Sereno Watson. He enjoyed the close friendship of Joseph and Thomas Howell, Martin W. Gorman, and Charles V. Piper. His many reminiscences of his botanical associates of another day are a delight to younger ears. His botanical explorations embrace a large portion of the states of Idaho, Washington, and Oregon. He was the first American to collect in the Olympic Mountains of Washington, and his various trips into the Salmon River country of Idaho opened up a region hitherto entirely unknown. In his honor, a host of species has been named, of which the following is only a partial list: Allium Hendersoni Rob. & Seat., Allocarya Hendersoni A. Nels., Amsinckia Hendersoni Suksd., Angelica Hendersoni C. & R., Aster Hendersonii Fernald, Astragalus Hendersoni Wats., Brodiaea Hend- ersoni Wats., Carex Hendersoni Bailey, Dodecatheon Hendersoni Gray, Eriogynia Hendersoni Canby, Erythronium Hendersoni Wats., Lappula Hendersoni Piper, and Lomatium Hendersoni C. & R. His long and fruitful career and his fine personal qualities may well serve as a model to younger generations of botanists. We wish him continued good health, and the enjoyment of many more such anniversaries. i . 433 a De . t at ‘ ‘ CONTENTS PAGE Batpwin, J. T., JR., Cytophyletic Analysis of Certain Annual and Biennial (CEASSIIACC AC Hare Mies fs, PRN pias ihardl cela ean born 4 Fre eee ed a nae Be 184 BALL, CARLETON R., Dr. Setchell and Alaska Willows .................... 231 Baker, Mito S., Studies in Western Violets—III ........................ 218 @ARTER, ANNETIA: Ma, REVIEW 22424 os 6a chip ase san ep oe dwacless Haw sees 237 Cave NUARION. Priscilla AVERY ..82 4.06 suenn ba sadeuss bemoan ¢oeuweddes 4 196 ConniT, Ira J., Mistletoe on Persimmon ................. 0.0000 cece eee 272 ConsraNce, Lincotn, The Genera of the Tribe Hydrophylleae of the Hydro- ONCE ACM Rae emt ary tre oahu ea ee eee eee ere eee ee ee 28 ConsTANCE, LINCOLN, Reviews ............. 00.000 cece eee eee 134, .170, 270 CORETAND: Ahan, FRCVICW. (oa. i te on dee Gens 1 de eda eiel vues Saas 170 CopELAND, Herspert F., A Portrait of John Gill Lemmon ................. ue CoreLAnD, Hersert F., The Structure of Monotropsis and the Classification Ofathe AViOnNOLEOPOldeae. 26.7 iia cite shone ee ea Aes nd ooo eee eae ee 105 CopeLannD, HerzBert F., The Phylogeny of the Angiosperms .......... ee 209 Cory, V. L., Six Thistles Recently Introduced into Texas ................. 200 Crum, Ernet, Alice Eastwood’s Eightieth Birthday .................... TA Crum, Eruet, A Revision of the Genus Monolopia ..................... 250 Dariinc, Louise, Protocoronospora on Phoradendron flavescens in Cali- iA) TN CU ea tart BAe PO bed x OO A fbn ced Oui Pcs Seu aS. Maal 241 Epiinc, Cart, Two Mexican Species of Hyptis ......................... 15 Eriine, Cart, A Note on the Occurrence of Salvia in the New World ... 34 Eriinc, Cari, Notes on the Scutellariae of Western North America ..... 49 Epuine, Cart, Teucrium glandulosum in California and Arizona ......... 135 Epiinc, Cart, A New Locality for Salvia eremostachya Jepson ......... 272 Eriine, Cart, and Witiiam Rosison, Pinus muricata and Cupressus Horbesit in Baja California: 2.20 0... 646 66 ees e ole sass ses wbvese ea 247 PECIRIR USED) A Mase reas ere ee Mn Ale Set mh tain ny Aes, Uva 8 Bey el give Bag pet Mee 280 Wea mil yy Cm VCVle Went «ae a a reas), Rae. cha Aicy nshinurek Oe WAG ae ee 202 GANDER, Franx F., New Records of Alien Plants in San Diego County, © UI onerael ch gee ee ews te ores Meee hehe a let bile thet ng 79 Gentry, Howarp Scott, A New Species of Viscainoa from Baja California 161 Haasis, FErpINaAND W., Transparent Mounts for Field Herbaria ......... 121 Henpverson, L. F., A New Thistle from Oregon .......................... 97 Herre, Atsert W.C. T., A New Species of Lecidea from Brazil .......... 235 Hircucock, C. Leo, Two Interesting Species of Draba from the Charleston NVLOUMULAMGMOIeRN VAG tacit att Gate dicl nail ab eee vat Stabe were eke ees 127 Hoert, Rutn A., A New Species of Arthrobotryum ..................... 75 Hoover, Rosert F., A Monograph of the Genus Chlorogalum ............. 137 FS DEX TON VOLUME, Vow. 2655 vag ls Bierce 0k os alec Dea ab 24 a apa de Pe oe week 274 Jounson, A. M., Some Abnormal Inflorescences ...........:............. 177 Keck, Davi D., Notes on Orthocarpus ................... 0.000000 0 eee 164 Keck, Daviw D., New Subspecies in Haplopappus ...................... 166 Keck, Davw D., The Identity of Madia dissitiflora (Nutt.) Torr. & Gray 169 KECK WiAvin. 1D.; REVIEWS co). «wee a ney She ed Sema oo 9S RE bees 170, 237 Mason, Herserr L., Elodea densa Casp. in California .................... 103 Mason, Herzert L., A Pleistocene Record of Pseudotsuga macrocarpa ... 233 Mason, Herpert L., Reviews ............ 0.000 cee ec eee 101, 170, 237 McCvuttocn, Ernesr C., Nutlets of Amsinckia intermedia Toxic to Swine, FVOESCS@anGiuG abGler see cree ngtes Sinica eR id be san ce 4 dus wage ae Ree ee 202 McMinn, H. E., Notes on the Genus Ceanothus in California ............. 13 MEMBERS OF THE CALIFORNIA BOTANICAL SOCIETY AND SuBSCRIBERS TO Ma- DRON OGM MEHL MONO) hg See Ss MPa do hase htm aced Kona ee io ene Ga eee 205 Mvenscuer, W. C., Additions to Our Knowledge of the Flora of Mount Baker, WV AS HINO LOMS oes cans ct ee eae ae eee a bata be ee ene es 236 Mutter, Cornetius H., New and Otherwise Noteworthy Plants in the SOMME be pegerte conor ae Pt ets tte eat pe ays Sut us teen ee Oe 152 PAGE Nores- AND NEWS 2) #3 sé ¢35%.00 09 $scee oe AT, 103, 135, 175, 202, 240, 272 Peck, Morton EK., Two New Compositae from the Wallowa Mountains of Oregon: Foo gs kn eh le De ee 247 PROCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY ........ AT, 175, 204, 273 Quick, CLiareNcE R., Ribes tularense (Cov.) Fedde in Sequoia National Park: 4ac56 be ee chal BS 4 8 eRe OLS OO ee ee 103 REED, FH. S., Fred. M: Reed 2.030225 3) ee ee eee 163 FUBVIEWS® 2.6... foe «BGG oa Pe ne ee ee 134, 170, 202, 237, 270 Rosison, Witiiam, and Cart Eprinc, Pinus muricata and Cupressus Korbesii in Baja California, 22....... es.) ee ee ee 247 Ro.uins, REED C., Notes on Certain Cruciferae of Mexico and Southwestern United’ States: 20 vih¢ees we ceecoe A ee eee ee ee ee 129 RossspacH, Georce B., and Rutru P., Southern Occurrences of Allium crenu- latum and Meconella:oregana: 2. 12735 eee ee 240 ScuHMo.i, Hazet M., A Realignment of the Panicum thermale Group ..... 90 ScurerBer, Beryt O., The Genus Helianthemum in California ............ 81 ScurersER, Beryt O., and A. E. Wiestanver, Notes on the Genus Arcto- StAP VIO. 5 eicccew kos tw chin ate na spree a eee ee ne ac 38 SHARSMITH, Cart W., Notes on Draba in the Sierra Nevada .............. 147 SuHarsmMiTH, Heten K., A New Species of Cirsium from California ........ 85 SHarsMITH, HELEN K., Further Notes on the Genus Sedella ............. 192 SurevE, Forrest, Observations on the Vegetation of Chihuahua ........... 1 Smirnu, CLayton O., Observations on the Elongation of the Stipe of Mon- PASMITES =. G2 a. als a Ee aa Whee Ae ee ete ee 119 Srepsins, G. Lepyarp, Jr., Notes on Lactuca in Western North America .. 123 Steners, G. Lepyvarp, Jk. Review .2:.55 34-6 oeu de io ses leds on eee 78 STOCKWELL, W. PALMER, Cone Variation in Digger Pine ................. 72 Spotts, Atice Maria, The Violets of Colorado .......................4.- 16 THomPsON,.J «- WibLIAM, Ira CAQUS: . 2.5 4525 ooeed te 2595 ee es ee 98 Wiestanper, A. E., and Beryzt O. Scureiser, Notes on the Genus Arcto- Staphy loss i.4.5 02:05 40's Baw 2a course a og Ge ee oe 38 Wiccins, Ira L., A New Species of Eriogonum from Baja California ...... 158 Witson, ALBERT, Pinus Torreyana in Cultivation ........................ 198 vi VOLUME V _-NUMBER 1 Ohi ao” Had he RP, Re fy me) Lis 16 i\ PAKS = QO ® 7 mS, MADRONO ~ A WEST AMERICAN JOURNAL OF BOTANY \ (rene? wh Contents OBSERVATIONS ON THE VEGETATION OF CuinvUAHUA, Forrest Shreve........ 1 Notes ON THE GENUS CEANOTHUS IN CaLirorniA, H. H. McMinw.......... 13 Two Mexican Species oF Hyptis, Carl Hpling..............0. 000000 eae 15 THE VioLtEets oF Cotorano, Alice Marial Spotts.............. 000 cece ee 16 THe GENERA OF THE TRIBE HYDROPHYLLEAE OF THE HYDROPHYLLACEAE, MOC OUT GIS CMICCE NNN hia) S/S Ps uas a svoue 14 Valevakatepa wcrc ee Mike eiabdeae eats 28 A Norte oN THE OCCURRENCE OF SALVIA IN THE NEw Wor tp, Carl Epling... 34 Notes oN THE GENUs ARcTosTaPHYLos, 4. H. Wieslander and Beryl O. PORT AET CANN WUD cURL GH PAULUS Stamey OF iL oe CRU TRN LNs ea Se a) 38 ROTRERS CAE OSES UES SCM a a Aa a aE eR AT PRocerDINGs OF THE CALIFORNIA BoranicaL SOCIETY.................-.0-- Aq Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania January, 1939 MADRONO A WEST AMERICAN JOURNAL OF BOTANY Board of Editors Dr. H. L. Mason, University of California, Berkeley, Chairman. Dr. L. R. Asrams, Stanford University, California. Dr. Lincotn Constance, University of California, Berkeley. Dr. H. F. Corpetanp, Sacramento Junior College, Sacramento, California. Dr. A. W. Havrt, University of California at Los Angeles. Business Manager—Dr. Davm D. Kreck North Queen Street and McGovern Avenue, Lancaster, Pennsylvania or Carnegie Institution of Washington Stanford University, California Entered as second-class matter October 1, 1935, at the post office at Lancaster, Pa., under the act of March 3, 1879. Established 1916. Published quarterly. Subscription Price $2.50 per year. Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17, complete, $5.00. Volume III, Numbers 1 to 8, complete, $5.00. Single num- bers $0.75. Papers up to 15 or 20 pages are acceptable. Longer contributions may be accepted if the excess costs of printing and illustration are borne by the contributor. Range extensions and similar notes will be published in con- densed form with a suitable title under the general heading “Notes and News.” Articles may be submitted to any member of the editorial board. Manuscripts may be included in the forthcoming issue provided that the contributor pay the cost of the pages added to the issue to accommodate his article. Reprints of any article are furnished at a cost of 4 pages, 50 copies $3.70; 100 copies $4.10; additional 100’s .75¢; 8 pages, 50 copies $5.40; 100 copies $6.00, additional 100’s $1.20. Covers, 50 for $2.50; additional covers at $1.50 per hundred. Reprints should be ordered when page proofs are returned. Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania, for the CALIFORNIA BOTANICAL SOCIETY, INC. President: Professor H. E. McMinn, Mills College, California. First Vice- President: Mr. Maunsell Van Rensselaer, Blaksley Botanic Garden, Santa Barbara, California. Second Vice-President: Mrs. Viola Brainerd Baird, Berkeley, California. Treasurer, Dr. David D. Keck, Carnegie Institution of Washington, Stanford University, California. Secretary: Miss Ethel Crum, 4004 Life Sciences Building, University of California, Berkeley. Annual membership dues of the California Botanincal Society are $2.50, $2.00 of which is for a year’s subscription to Madrofio. Dues should be remitted to the Treasurer. General correspondence and applications for membership should be addressed to the Secretary. 1939 ] SHREVE: VEGETATION OF CHIHUAHUA 1 OBSERVATIONS ON THE VEGETATION OF CHIHUAHUA Forrest SHREVE The state of Chihuahua lies in the northwestern corner of the central table land of Mexico. Its location, size and physical features make it an area of importance in connection with the plant and animal geography of the southwestern United States, as well as in the investigation of the relations between the Sono- ran and Chihuahuan deserts. Knowledge of the flora of Chihuahua is based mainly upon collections made by some fourteen men,’ and information regard- ing it is scattered through scores of publications over a period of sixty years. There is no check list or manual covering any part of the state. On the character and distribution of the vege- tation only a few pages have been written and these with refer- ence to the northern border. In many publications on other topics the plants have received occasional mention. Several men have mapped the vegetation of North America or of Mexico with little to guide them in reference to the Chihuahuan region. Knowledge of the fauna and ecological features of the animal life is in a very similar condition. Brand (1) has published a map of the vegetation of north- western Mexico, recognizing five types in Chihuahua. He makes no reference to the grassland and publishes no descriptions of the regions indicated on his map. In a later paper Brand (2) has described the land forms of the highlands of Chihuahua north of Ciudad Guerrero, with brief notes on the vegetation and some helpful illustrations and climatic tables. The dearth of botanical information about Chihuahua has prompted the publication of the following brief sketch of the state, based on a trip made in the summer of 1937, on which all parts were visited except the oak forest and barrancas of the extreme southwestern corner. The writer was accompanied by Dr. L. R. Dice, of the University of Michigan, and Dr. T. D. Mal- lery, of the Desert Laboratory of the Carnegie Institution, to both of whom he is indebted for helpful discussion of the condi- tions encountered. PHyYSIOGRAPHY Five rather clearly defined physiographic regions may be recognized in Chihuahua. These are the bajadas and low moun- tains of the northeast and east, the enclosed basins of the north- west and southeast, the elevated plains of the central district, the Sierra Madre Occidental, and the barrancas, or canyons of the Pacific slope of the Sierra Madre. 1 Wislizenius, Gregg, Schaffner, Wilkinson, Pringle, Palmer, Nelson, Rose, Hartman and Lloyd, Townsend and Barber, Hitchcock, and Le Seuer. Maprono, vol. 5, pp. 1-48. January 17, 1939. 2 MADRONO [Vol. 5 Tue Basapa Recion. This part of the state borders the Rio Grande and the lower course of its only important Chihuahuan tributary, the Rio Conchos. The elevation varies from 800 to 1450 meters (2700 to 4750 feet). Gently falling bajadas and broad intermont plains occupy a high percentage of the region. Extensive limestone areas present the usual surface features of that material, being either hilly and broken or else rolling or nearly level, with little evidence of erosion and transport. The mountains of this region probably occupy from 5 per cent to 8 per cent of the total area. Tue Enctosep Basin Recion. The largest enclosed basins, or bolsons, lie in the north and the extreme southeast, at elevations from 1175 meters (3850 feet) in the former area to 1400 meters (4600 feet) in the latter. There are also numerous smaller basins in the midst of the Bajada Region as well as in the Ele- vated Plains Region. On the north three large lake beds, or playas, Laguna de Guzman, Laguna de Santa Maria and Laguna de Patos, receive the flood waters of rivers which drain the north end of the Elevated Plains Region. These rivers are fed by heavy summer rains and fall at a steep gradient, conditions which have enabled them to cut into the old deposits of mountain out- wash and to carry sand through to their terminal playas. Dur- ing the dry seasons water sometimes disappears from the lake beds and the deposits of sand are blown for distances of as much as 80 to 100 kilometers (50 to 60 miles). Part of northern Chi- huahua is a vast dune complex and has a surface shaped almost wholly by the wind. The failure of the three contributaries of the largest playas to deliver their flood waters to the Rio Grande is apparently due to the accumulation of their loads between the playas and the Rio Grande. The enclosed basins in the southeast, as well as the smaller ones in the central districts, are of structural origin. They are fed only by short streamways and floods from the torrential rains of the desert lowlands. Their playas are therefore built of fine alluvium rather than sand and have a high salt content. Under these conditions the action of the wind is confined to the daily agitation of the superficial alkaline dust through local spiral movements. Groups of white “tornillos” rise constantly during the daytime in slowly coiling columns to a height of 80 to 100 meters and circle majestically about the playa and its edges. The effects of wind are therefore slight, leaving the surface essentially as determined by the agency of water. The drainage of extreme southeastern Chihuahua and the neighboring part of Coahuila runs into a large number of sepa- rate basins, which have either a central playa, if their catchment area is large, or a grass-covered floor, if their drainage area is small or wholly in the arid lowlands. Tue Exvevatep Priains Recion. This feature of Chihuahua forms a belt along the eastern base of the Sierra Madre which 1939] SHREVE: VEGETATION OF CHIHUAHUA 3 varies from 25 to 110 kilometers (15 to 65 miles) in width. Ele- vations in the Plains Region range from 1600 to 2100 meters (5250 to 7000 feet). Their surface is dotted with numerous small mountains and their eastern edge is supported by a series of barrier ranges which have a basal elevation about 600 meters (2000 feet) higher on the west than on the east. The Plains are composed of thoroughly disintegrated outwash material de- rived from the ancient degradation of the Sierra Madre. On the east the present level of the Plains is being effectively maintained by the barrier ranges. In the north, however, the surface is be- ing attacked by the three rivers which feed the northern basins, and on the west the Rio Papigochic, a tributary of the Yaqui, has eaten back into the Plains far east of the continental divide. Tue Sierra Mapre Recion. The Sierra Madre Occidental lies in a north northwest-south southeast position and forms the con- tinental divide, as well as the boundary between Chihuahua and Sonora. It consists of a broad series of ridges and mesas with summit elevations of 2500 to 2900 meters (8200 to 9500 feet), but at least three unexplored summits are believed to exceed 3500 meters (10,000 feet). The ridges are maturely worn and there are innumerable parks, meadows and flood plains with heavy accumulations of alluvium. There are large areas in the midst of which are no abrupt changes of elevation of more than 100 to 150 meters. The northernmost forested elevation is the Sierra de la Espejuela, 80 kilometers (50 miles) south of the International Boundary. With a single break at Carretas Pass, immediately south of this mountain, the Sierra Madre lies con- tinuously above 2150 meters (7000 feet) for 800 kilometers (500 miles) to the south. Tue Barranca Recion. A small part of the Pacific slope of the Sierra Madre is comprised in this region, which lies partly in the states of Sonora and Sinaloa. Here are the evidences of the active cutting which has taken place since the last uplift of the Sierra Madre. Deep well-watered canyons with steep slopes have eaten far back into the mountains and in many places have isolated former portions of the Sierra or its subsidiary ranges. The Barranca Region is thus given an extremely irregular con- figuration and is confined to narrow bands and small areas. The fall from the western edge'of the mature summit of the Sierra at 2150 meters (7000 feet) to the foothills of Sonora at 200 to 500 meters (650 to 1650 feet) takes place within a distance of 45 to 80 milometers (28 to 50 miles). VEGETATION The distribution of the types of vegetation which clothe Chi- huahua is found to run closely parallel to that of the physio- graphic provinces which have just been described. On the accompanying map (text fig. 1) are shown the approximate boun- A MADRONO © . [Vol. 5 OESERT OAK FOREST & |! GRASSLAND I PINE FOREST BARRANCA Fic. 1. Distribution of types of vegetation in Chihuahua. daries of desert, coinciding with the Bajada Region and the Enclosed Basin Region; of grassland and oak forest, covering the Elevated Plains Region and the lesser mountains; of pine forest, clothing the Sierra Madre Region; and of oak and broad- leaf forest filling the Barranca Region. These types of vegeta- tion are clearly distinct except for the existence of intermediate areas between the desert and grassland and some intermingling of oaks and pines. Desert. In the north and east Chihuahua is occupied by an unbroken stretch of desert which covers about 60 per cent of the state. It is a region of broad plains, long bajadas and large basins with widely spaced ranges of low mountains. There are many opportunities to travel for 150 to 300 kilometers (100 to 1939 | SHREVE: VEGETATION OF CHIHUAHUA 5 200 miles) without encountering substantial differences in the vegetation. Nevertheless there are marked differences between the widely separated parts of the area. The most distinct types of desert vegetation are those found in the sandy basins and dune region of the north, the bajadas and outwash plains of the north- east, the volcanic and limestone mountains of the northeast and east, the limestone plains of the south, and the bolsons of the southeast. In the dune region a high percentage of the surface is wholly or partly stabilized but is covered by a very open stand of plants. There are local areas of the light cream colored sand which are in active movement, as well as numerous massive dunes almost wholly without plants. All of the features of dune development and plant succession which have been so adequately described by many workers are here exemplified on a large scale. The stabilized areas have a very open stand of Prosopis glandulosa, Artemisia filifolia, Poliomintha incana, Ephedra Torreyana, Dalea scoparia, Hymenoclea monogyra, Atriplex canescens, Leiostemon Thur- beri and Yucca elata. Cacti are very uncommon and represented almost wholly by low platyopuntias. The most characteristic small perennials are: Croton neomexicanus Coldenia hispidissima Riddelia Cooperi Melampodium leucanthum Heliotropium Greggii Sporobolus flexuosus Bahia absinthifolia Wislizenia refracta Sphaeralcea incana Palafoxia linearis Euphorbia Parryi The bajadas and outwash plains occupy a larger area in Chi- huahua than any other type of desert, covering the sector which lies between lines drawn north and east from Chihuahua City. Their vegetation is very uniform in physiognomy and very simple in composition. It is strongly dominated by Larrea, which is often nearly pure, or elsewhere associated with Flourensia cernua where the soil is thin or with Prosopis glandulosa where the soil is deep. In the lowest part of each basin or valley Prosopis is dominant as a shrub of 1 to 2 meters with its trunk buried and its branches radiating from a hummock of soil. On the bajadas there are only a few large perennials asso- ciated with the three that have been mentioned, and none that are abundant. The important ones are: Koeberlinia spinosa Opuntia macrocentra Acacia Greggii Coldenia Greggii Condalia spathulata Opuntia imbricata Celtis pallida Acacia vernicosa Condalia lycioides Microrhamnus ericoides On the higher parts of the bajadas, on the pediments sur- rounding the mountains and on the hills and mountain slopes the 6 MADRONO [Vol. 5 vegetation is much more diversified (pl. 1, fig. 1). Larrea loses its dominance although it does not disappear, and all of the asso- ciates just listed become more abundant. The most conspicuous ' plants are Dasylirion Wheeleri, Fouquieria splendens, Opuntia imbri- cata and the massive Yucca macrocarpa. The most abundant plants are Agave Lechuguilla, Coldenia Greggiti, Euphorbia anti- syphilitica, Parthenium incanum and Jatropha spathulata. ‘The con- spicuous plants are particularly abundant at the foot of mountain slopes and on the pediments. On gentler slopes, especially on limestone, the larger plants give way to the small but abundant ones. The only conspicuous cacti are Opuntia imbricata, reaching a height of 1 to 2 meters and O. macrocentra, which is here an . erect plant but rarely exceeds 1 meter. Smaller cacti are, how- ever, an important element in the vegetation, being chiefly repre- sented by Echinocactus horizonthalonius, Echinocereus dasyacanthus, E. stramineus, Coryphantha macromeris, Escobaria tuberculosa, and Thelocactus bicolor. Along the small drainageways of the desert there is a slight increase in density on the part of Prosopis, Acacia, Celtis and Con- dalia spathulata, but no considerable increase in height. Only along the largest arroyos are there any trees in the desert sense of woody plants with one or several trunks, branching from the ground level, and reaching a height of 5 to 6 meters. In addi- tion to larger individuals of Prosopis will be found Rhus chorio- phylla, Berberis trifoliata, Porlieria angustifolia, Chilopsis linearis, Celtis pallida, and Acacia paucispina. The enclosed basins, or bolsons, of southeastern Chihuahua are part of a large region of similar character which extends into the neighboring states of Coahuila and Durango. Here the mountains are low and very widely spaced and the drainage is shallow and vacillating. Nearly level plains from 30 to 80 kilo- meters (20 to 50 miles) in width present very uniform conditions. The elevation is from 1200 to 1500 meters (4000 to 5000 feet). The soil is light gray and of fine texture. The vegetation is low, sparse and monotonous. In the higher basins the floor is covered by pure stands of Hilaria, in the lower ones the dominant plants are Larrea, Acacia vernicosa and Flourensia cernua, constantly vary- ing from place to place in their relative abundance. Prosopis and Atriplex canescens are also locally abundant as well as spar- ingly represented throughout the plains. Less common are Celtis pallida, Condalia lycioides, Koeberlinia spinosa, Buddleia scor- dioides, and Lippia Wrightii. Cacti are uncommon and repre- sented only by Opuntia Kleiniae, O. macrocentra, O. leptocaulis and Echinocereus conglomeratus. South of Saucillo the Rio Conchos and the Rio Florido form the western boundary of the bolson region. West of this boun- dary is a rolling limestone plain which extends south nearly as far as Parral and rises toward the west. The elevation of the Ficure 1 Figure 2 Puate 1. Vecerarion or Cuinvanva. Fig. 1. Vegetation on outwash slopes of Sierra Tasajera in northeastern Chihuahua at 1600 meters elevation: dominant plants, Yucca macrocarpa, Dasylirion Wheeleri, Agave Lechuguilla, Coldenia Greggii, Lippia Wrightii and Hilaria mutica. Fig. 2. Limestone piains west of Boquillas, in southern Chihuahua at 1400 meters elevation: vegetation a low closed stand of Acacia vernicosa and Larrea tridentata. In the depression: Celtis reticulata, Condalia spathulata and Condalia lycioides. 1939 | SHREVE: VEGETATION OF CHIHUAHUA 7 plain varies from 1150 to 1375 meters (3800 to 4500 feet). The surface has both shallow and deeply cut drainageways, as well as many large and small depressions which have no surface out- let. The shallow drainageways and the large depressions have a deep clay soil which is commonly a very dark gray. A high percentage of the region has a soil which is very shallow or con- fined to irregular pockets in the surface of the limestone. The vegetation of the thin soil of the limestone plains is a dense stand of shrubbery which rarely exceeds 1 meter in height (pl. 1, fig. 2). Acacia vernicosa is strongly dominant, with Lar- rea and Flourensia forming less than 15 per cent of the stand. Plants of secondary importance are few and infrequent, includ- ing: Celtis pallida Rhus microphylla Condalia spathulata Leucophyllum laevigatum Agave Lechuguilla Opuntia macrocentra Opuntia Engelmannii Koeberlinia spinosa On the slightly deeper soil surrounding the depressions Larrea and Flourensia become more abundant than Acacia vernicosa, and nearly pure stands of Flourensia are frequently seen. In the de- pressions, which are in some cases from 10 to 15 kilometers in width, there are no shrubs and the prevailing vegetation is a very open sod of Hilaria mutica. The western edge of the limestone plains was crossed at only one locality and the configuration of the area can not be stated. Without essential change of elevation the Acacia thickets give way on leaving the limestone, and at 1275 to 1375 meters (4200 to 4500 feet) on a deep loam soil a wholly different type of vegetation is encountered. This is a very open stand of grasses and root perennials with widely spaced shrubs 1.5 to 3 meters high. The three characteristic shrubs of the limestone, Larrea, Flourensia and Acacia vernicosa, are uncommon. The principal large shrubs are Celtis pallida, Prosopis glutinosa, Condalia lycioides, Cassia Wislizeni, Tecoma Stans, Mimosa biuncifera and Ephedra antisyphilitica. This is the uppermost type of desert vegetation and merges gradually into a desert-grassland transition similar to that which covers the plains of southern New Mexico. In Chihuahua the desert-grassland transition occupies the plains and bajadas which extend from Moctezuma on the north to the southern boundary of the state; lying above 1525 meters (5000 feet). It also covers the foothills and lower slopes of the barrier ranges, but the conditions there are diversified and the distribu- tion of the vegetation is complex (pl. 2, fig. 1). Along stream- ways of mountain origin Quercus Emoryi descends to 1450 meters (4800 feet) and Q. chihuahuensis is sparingly represented on the slopes of the foothills at 1600 to 1775 meters (5250 to 5825 feet). Cacti and yuccas are uncommon but are seen much more fre- quently than in the upper edge of the desert. On ascending 8 MADRONO [Vol. 5 through the desert-grassland transition there is a gradual in- crease in the percentage of the surface covered by grasses, par- ticularly on nearly level ground with soil 8 or 4 decimeters or more in depth. The elevation of the lower edge of the grassland is close to 1700 meters (5500 feet) at all localities visited. GrassLanp. The grassland of Chihuahua resembles closely that of southeastern Arizona and southwestern New Mexico in its local distribution with respect to topography, in physiognomy, in its dominant grasses and to some extent in the associated non- gramineous plants (pl. 2, fig. 2). At all elevations the prevail- ing grasses are species of Bouteloua (B. gracilis, B. hirsuta, B. radi- cosa, B. chondrosioides) ; which serve to give a striking uniformity to the texture of the plant covering over large areas. In certain places taller grasses form bunches scattered over the sod of Bouteloua. In many parts of the desert-grassland transition there is evi- dence that grazing has altered the natural vegetation, with a re- duction in the amount of grass and an increase in the native weeds Gutierrezia, Aplopappus and Verbesina. Above 1700 meters (5500 feet), however, the continuity of the grass cover, the absence of gullying and excessive erosion, and the scarcity of unpalatable weeds indicates that the vegetation is in a con- dition close to the natural one. Although the higher grass- lands are grazed, it appears obvious that the drain upon them has been light enough for the maintenance of a good self-per- petuating cover under the existing rainfall conditions. On the upper margins of the grassland valleys a very open oak forest has, in many places, been superposed on the grass- land, with little modification of the composition of the grass- land except in the shade of the oaks. In such situations a few shrubs are found, being species of more abundant occurrence at somewhat higher elevations. Where extrusions of lava or dykes of other rock occur in the grassland and now lie nearly at accordant level with the surface of the outwash material surrounding them, there is a light and broken cover of grasses (Andropogon barbinodis, A. hirtiflorus, Eragrostis mexicana, Aris- tida adscensionis, A. divaricata, Muhlenbergia Emersleyi, Oryzopsis fimbriata). In such situations there is commonly a very open stand of woody perennials, including stunted individuals of Quercus Emoryi, Q. santaclarensis, Juniperus mexicana, Mimosa biuncifera, Cowania Stansburiana and Mimosa dysocarpa. When the grassland region is considered as a whole the total number of root perennials associated with the grasses is large. The presence of a particular group of species in a given spot appears to be determined to some extent by the tex- ture and prevailing moisture of the soil, and to a very con- siderable degree by the chances of seed dispersal and other more obscure conditions. Certain of the associated perennials Ficure 1 Figure 2 Piate 2. Vecerarion or Cuinvuanua. Fig. 1. Hills west of Santa Isabel in desert-grassland transition at 1700 meters elevation. The tree is Quercus chihuahuensis. Fig. 2. Looking east across grassland in upper drainage of Rio Papigochic at San Isidro in western Chihuahua at 2100 meters elevation. \% ith 3 . 7 de 1939 ] SHREVE: VEGETATION OF CHIHUAHUA 9 are of general occurrence while others are abundant in areas of 1, 10 or 100 hectares and rarely seen elsewhere. Many of the perennials are prostrate in habit or low in their mature height, while a number of others reach a height of 6 to 8 decimeters at the time of flowering. About half of the tall species are slender and have leaves either so small in size or so few in number that the plants are inconspicuous. Very few native annuals are found in the region. The great majority of the herbaceous perennials have enlarged roots or rhizomes. The number of species and individuals of bulbous plants is much greater than in the grasslands of Arizona and New Mexico. The following grassland associates are low or prostrate plants little exceeding the turf of grasses in height: Brayulinea densa Tragia nepetifolia Houstonia Wrightii Acalypha Lindheimeri Dyschoriste decumbens Verbena canescens Plantago mexicana Vincetoxicum nummularium Calliandra humilis Sisyrinchium tenuifolium Potentilla Mexiae Nothoscordum fragrans Phlox mesoleuca Oenothera serrulata Helianthemum glomeratum The taller perennials of the grassland include: Hymenopappus mexicanus Scutellaria sp. Petalostemon oligophyllum Eryngium Wrightii Thelesperma gracile Castilleia mexicana Dalea Grayi Lupinus Shrevei Tagetes lucida Gaura gracilis Lithospermum cobrense Erigeron divergens Linum australe Ratibida columnaris Some of the grasses and many of the herbaceous perennials of the grassland are found in the oak forest and even in the lower and more open parts of the pine forest. The upper edge of the grassland is determined by the fact that there is no soil above 2200 meters (7200 feet) which is rock-free, level and not ex- tremely moist. Oak Forest (Encinat). The oak forest of Chihuahua is found through nearly the same range of altitude as the grassland, which indicates that the climatic requirements of the two are closely similar. The oaks of lowest range, Quercus Emoryi and Q. chihuahuensis, descend locally into the upper edge of the desert- grassland transition. The oaks of highest range, Q. hypoleuca, Q. pennivenia, Q. Sipuraca and Q. epileuca, are associated with the pines at 2150 to 2300 meters (7000 to 7550 feet) and become minor constituents of the pine forest above that elevation. The distribution of oak forest and grassland throughout their common range of altitude is determined by soil conditions, as 10 MADRONO [Vol. 5 already indicated. Below 2000 meters (6560 feet) the oak forest | is very open and many of the trees are small. On the eastern edge of the grassland plateau there are ridges and hills of 2150 meters (7000 feet) with very poor stands of Quercus Emoryi, but at the same elevation nearer the base of the Sierra Madre the stands are heavier and the trees are larger. The finest groves of oaks, below the elevations at which they mingle with the pines, are on the coarse soil of broad flood plains, bordering streams that have just emerged from mountain canyons at 2000 to 2150 meters (6560 to 7000 feet). Rarely are such groves dense enough for the full grown trees to shade more than half of the surface of the ground. On stony, level soil with poorer moisture conditions there are dense thickets of small oaks. On the hills and low mountain ranges that interrupt the continuity of the great grass- land valleys the most abundant oaks are Quercus Emoryi, Q. santa- clarensis, Q. chuichupensis and Q. hypoleuca. Other oaks occur- ring locally or in very scattered stands are Quercus chihuahuensis, Q. arizonica and Q. durangensis. Juniperus mexicana and Pinus cembroides often occur with the oaks and on arid ridges out- number them. In the latter part of July 1987 all of the species of oak were in leaf in favorable localities. The foliage of Quercus santa- clarensis and Q. chuichupensis appeared to be young at that time, indicating that these species are deciduous. On some of the small ranges of hills well to the east of the Sierra Madre the dominant oaks are Q. santaclarensis and the evergreen Q. Emoryi, both of which were conspicuously bare of foliage. This condi- tion was probably due to the unusually late commencement of the summer rains in 19387. Many grasses and a large number of shrubby and herbaceous perennials are characteristic of the oak forest. Fully one-fourth of the number are equally abundant in the grassland. The pres- ence of many of the others is favored by the shade of the oaks, but a far larger number are present as a result of the stony sur- face and diversified soil structure and are to be found outside the shade of the oaks. The most abundant shrubs and herbaceous perennials in the oak forest are: Rhus trilobata (shrub) Cologania Lemmoni (a9 Ceanothus buxifolius Mimosa dysocarpa Cowania Stansburiana Rhamnus pinetorum Bouvardia triphylla Houstonia Wrightii Erigeron divergens Gnaphalium chilense Cyperus seslerioides Muhlenbergia gracilis Tradescantia pinetorum Dyschoriste decumbens Penstemon pulchellus Thalictrum pinnatum Lithospermum cobrense Cologania longifolia Eriogonum atrorubens Ipomoea costellata 1939 ] SHREVE: VEGETATION OF CHIHUAHUA 11 Indigofera ornithopodioides Bidens heterosperma Tradescantia crassifolia Geranium niveum Eragrostis mexicana Hymenopappus mexicanus Phaseolus Metcalfei Drymaria tenella Ipomoea madrensis Desmodium neomexi- Oxalis albicans canum (annual) Tephrosia leucantha Schkuhria Pringlei . Helianthemum glomeratum Crotalaria sagittalis i Stipa Pringlei Pine Forest. On the eastern slopes of the Sierra Madre the lower edge of the pine forest is approximately at 2150 meters (7000 feet) elevation but in the valleys draining into the Pacific it descends to 1800 meters (5900 feet), while open stands mingled with oaks occur below 1525 meters (5000 feet). About 18 per cent of the area of Chihuahua is blanketed by a nearly continuous forest of pines, more than half of which is in the Pacific drainage. Like all other great mountain ranges the Sierra Madre pre- sents a wide diversity of habitat conditions, with its stony ridges, gentle slopes, parks, meadows and constant streams. In general aspect the pine forest and its accompanying communities of plants resemble the mountain forests of Arizona and New Mexico. The varying density of the stands of pine, the fringe of broad-leaved trees along the water courses, the thickets of Populus tremuloides on the cool slopes, or of shrubby oaks and Arctostaphylos pungens on the sunny ones, the parks, meadows and streambed gravel all bear vegetation with the familiar physi- ognomy of the north (pl. 3, figs. 1, 2). The similarity is found to be due to plants of the same genera, or the same life forms, very many of which belong to species not found north of Chihua- hua. Many trees, a few shrubs and a large number of aquatic and palustrine plants are common to the pine forests of the south- western states. The number of species of trees found above 2150 meters (7000 feet) is not large. On the lower edge of the pine forest and in its outlying areas Pinus apacheca and P. chihuahuana are the domi- nant trees, with P. cembroides in the least favorable situations. The principal pine on the highest ridges and slopes is Pinus ari- zonica. On the Pacific slopes the dominance is held in some val- leys by P. Lumholtzii and in others by P. ayacahuite. The only other abundant conifers are Pseudotsuga mucronata, Pinus pinceana, P. strobiformis, Cupressus arizonica and Juniperus mexicana. Other trees of scattered occurrence or of distinctive habitat require- ments are: Quercus pennivenia Quercus durifolia Alnus acuminata Quercus diversicolor Populus tremuloides Arbutus xalapensis Quercus epileuca Quercus incarnata 12 MADRONO [Vol. 5 In small shrubs and herbaceous perennials the flora of the pine forest of the Sierra Madre is extremely rich. Not only new species but many new genera confront the visitor from the north. The Leguminosae and Compositae are very richly represented by plants of both wet and dry situations. Many of the important genera of mountain plants of the western United States are pres- ent here, as Rubus, Potentilla, Penstemon, Lithospermum, Erigeron, Gnaphalium, Ranunculus, Stellaria, Dalea and Desmodium. A large number of genera are either endemic to the Sierra Madre or else find their northern limits in the forests of Chihuahua. Affinity with the characteristic elements of the flora of the Sierra Madre is found in the United States only in the Chiricahua, Huachuca, Santa Catalina and adjacent mountains of southern Arizona. The number of collectors who have penetrated the Sierra Madre is small and the percentage of new species found in the most recent collections indicates that the wealth of its flora can not yet be estimated. The distribution of many of the common- est plants is very poorly known, and the character, distribution and composition of the plant communities remain uninvestigated. Barranca Forest. The writer has seen nothing of the higher elevations of the Barranca Region in Chihuahua and only a few areas on its lower edge in Sonora. Lumholtz (4) gave some brief descriptions of this vegetation in the narrative of his expedi- tion across the Sierra Madre from Granados, Sonora, to Casas Grandes, Chihuahua, through the valley of Rio Bavispe. Hovey (3) published a few botanical observations made on a geological excursion from Minaca to Dolores, in the drainage of Rio Aros, a tributary of Rio Yaqui. The forest in the valley of Rio Mayo and its tributaries has been explored recently by Gentry, and the writer is indebted to him for an opportunity to examine the manuscript of a paper which he is preparing on the flora and vegetation of that region. Knowledge of the barranca forest is nevertheless sufficient to indicate that it is one of the distinctive vegetational regions of Chihuahua. The collections made by Gentry in the Mayo Valley give ample evidence of the richness and sub-tropical affinities of the flora. SuMMARY Chihuahua occupies the eastern slope of the continental divide immediately south of the Rio Grande. The surface of the state is highly diversified in elevation and physiographic development. The distribution of the types of vegetation conforms closely to the physiographic provinces. The broad detrital valleys, with their pediments, low mountain ranges and enclosed basins, are occupied by desert. The elevated plains and their included mountains are occupied by grassland where the soil is deep and nearly level and by evergreen oak forest on mountain slopes or Ficure 1 FIcure 2 Pirate 3. Vecreratrion or Cutruanva. Fig. 1. Mountain slopes twelve kilometers north of San Juanito in southwestern Chihuahua at 2400 meters elevation. The open forest of Pinus arizonica, Pinus apacheca, and Populus tremuloides faces a slope covered by a thicket of Quercus bocoyensis. Fig. 2. Park in pine forest near San Juanito at 2500 meters elevation with grasses, Ranunculus pentodontus and Achaetogeron affnis. a = 1939 ] McMINN: CEANOTHUS 13 shallow soil. The slopes and summits of the Sierra Madre Occi- dental are covered by forest in which several species of pine are dominant. The narrow canyons on the Pacific slopes of the Sierra Madre are occupied by a forest of evergreen oaks and other evergreen or deciduous broad-leaved trees. In their physi- ognomy the types of vegetation resemble those of southeastern Arizona and southwestern New Mexico. The flora of northern Chihuahua is very similar to that of the adjacent parts of New Mexico and Texas, but in each of the types of vegetation there is a strong infusion of endemic or southern species not found in the United States. Desert Laboratory, Carnegie Institution of Washington, Tucson, Arizona, July, 1938. LITERATURE CITED 1. Brann, D. D. Notes to accompany a vegetation map of northwestern Mexico. Univ. New Mexico Bull. 280. 1936. 2. ————_———.. The natural landscape of northwestern Chihuahua. Univ. New Mexico Bull. 316. 1937. 3. Hovey, E. C. The Western Sierra Madre of Chihuahua. Bull. Amer. Geog. Soc. 37: 531-543. 1905. 4. Lumuoitz, C. Unknown Mexico. London, Fisher Unwin. 1905. NOTES ON THE GENUS CEANOTHUS IN CALIFORNIA H. E. McMinn In several genera of California shrubs such as Rhamnus, Ceanothus, Baccharis and Arctostaphylos, there are species which vary in their habit of growth from tall and erect to low, spread- ing, decumbent, or prostrate. The low-growing forms of some of these species are very desirable as ornamentals, especially as ground-covers. In some instances there is at present little evi- dence for the separation taxonomically of the prostrate, decum- bent, and sprawling forms from the erect forms. In others, however, where observations on transplants have been made it seems clear that the low growth-forms which usually inhabit coastal bluffs and adjacent wind-swept mountains are genetically distinct from the erect plants which grow farther inland or away from the effects of the winds from the ocean. One of these forms having retained its low-growing habit in cultivation for over three years is here described as a new variety. CEANOTHUS PAPILLOsUS var. Roweanus var. nov. Frutex com- pactus, sempervirens, 3-9 dm. altus (vel inter alios frutices), usque ad 1.5 m. in latitudine 1.2—-2 m.; ramis arcuatis, ramulis aliquanto horizontalibus; foliis anguste oblongis vel linearibus, 1.2—-4.5 cm. longis, 2-8 mm. latis, pinnate nervatis, supra atro- viridibus, glabris, dense glanduloso-papillatis, subtus dense lanato-papillatis, marginibus revolutis et glanduloso-papillatis, sessilibus vel brevipetiolatis ; flores fructusque ut ei apud speciem. 14 MADRONO [Vol. 5 Compact evergreen shrub, 3 to 9 dm. high (or up to 1.5 m. in competition with other shrubs) and with a spread of 1.2 to 2 m.; branches arched and in somewhat horizontal sprays; leaves nar- rowly oblong or linear, 1.2 to 4.5 cm. long, 2 to 8 mm. wide, pin- nately veined, upper surface dark green, glabrous, densely glandular-papillate, lower surface densely woolly-hirsute, the margins revolute and glandular-papillate, sessile or short-petio- late; flowers and fruit as in the species. Type. Summit of Mount Tranquillon, northwestern Santa Barbara County, California, altitude 2250 feet, March 27, 1938, M. Van Rensselaer 450 (University of California Herbarium no. 600030). Capsules were collected at the same locality by E. D. Rowe July 7, 1938. This variety, to my knowledge, occurs only in a small area in the vicinity of the type locality. Its spreading habit, compact branching, dark green papillate leaves, and dark blue flowers borne in great profusion make it a most desirable ornamental. Specimens which have been in cultivation for over three years at La Purisima Mission Gardens and at the Blaksley Botanic Garden have retained the low spreading habit whereas trans- planted specimens and plants grown from seeds of Ceanothus papillosus Torr. & Gray taken from the Santa Cruz Mountains have maintained the erect habit of growth. I take pleasure in naming this variety in honor of Mr. E. D. Rowe, who first called it to the attention of botanists in 1935. Mr. Rowe, now in charge of the planting at La Purisima Mission, has been in landscape gardening work in Santa Barbara for the past thirty years and has done much to foster an interest in a wide use of native shrubs as ornamentals. For the past ten years I have been interested in studying the distributional range of another group of Ceanothi which have been variously referred to Ceanothus cuneatus Nutt., C. rigidus Nutt., or C. cuneatus var. ramulosus Greene. These shrubs occur in dry rocky or more usually in sandy situations in the Coast Ranges from Santa Barbara County (La Purisima Hills and Burton Mesa, near Lompoc) northward to southern Monterey County and in Santa Cruz and Marin counties. Around Mon- terey Bay they are replaced by C. rigidus Nutt. Since these plants are intermediate between C. cuneatus and C. rigidus and are not associated geographically with either species it seems best for the time being to consider them as specifically distinct. An emended description follows. Ceanothus ramulosus (Greene) McMinn comb. nov. C. cu- neatus var. ramulosus Greene, Fl. Fran., 86. 1891. Coast Ceano- thus. Shrub, 6 to 12 dm. high; branches spreading or arching, or sometimes procumbent; leaves opposite and evergreen, usually not crowded on the branchlets; the blades variable in shape but 1939] EPLING: HYPTIS 15 typically obovate to oblanceolate, sometimes nearly round, 6 to 20 mm. long, 3 to 9 mm. wide, light to dark green above, paler beneath, usually toothed near the truncate or rounded apex, often entire, nearly sessile; flowers lavender, blue, or nearly white, in small peduncled umbels; fruit globose, about 5 mm. in diameter, varying from prominently 3-horned to nearly hornless. February to April. On the bluffs above Point Sal, Santa Barbara County, this species is procumbent to nearly prostrate. On Burton Mesa and on La Purisima Hills, Santa Barbara County, the plants have dark compact foliage and in this character they resemble plants of C. rigidus more closely than those of C. cuneatus. The leaves are often cuneate-oblong and the fruits are often nearly hornless. Plants from seeds collected in Marin County and grown in our trial garden with C. rigidus and C. cuneatus show the distinct long- arching type of branches and pale blue or lavender flowers characteristic of the parent plant. Mills College, California, November 18, 1938. TWO MEXICAN SPECIES OF HYPTIS Cart Eriinec Amongst a recent shipment of Mexican Labiatae collected by Mr. George C. Hinton in the states of Guerrero and Mexico are two undescribed species of Hyptis. Both are referable to the section Buddleioides subsection Umbellatae. As judged from the herbarium specimens both are lovely. Hyptis perpulcher sp. nov. per specim. in Mexico prov. Te- mascaltepec ad Pungarancho in colle a Hinton (no. 8574) lectum constituta est; typum in herb. Univ. Calif. (Los Angeles), iso- typum in herb. Kew. vidi. Herba suffruticosa fragrantissima perpulcher ut videtur cauli- bus ad 4 m. altis solum superne inter flores visis purpureis glaucis ramulis superne hirtellis; foliorum laminis cordatis 12-15 cm. diametro (? et ultra) petiolis 5 cm. longis elatis, in apice acumi- natis, marginibus irregulariter crenato-serratis, paginis ambobus glabris solum subtus ad venas obscure hirtellis; floribus numerosis parvis in umbellis purpureis pedunculis .5—1 cm. longis elatis bracteis parvis linearibus subtentis in paniculis speciosis gracilli- mis dispositis, pedicellis maturis filiformibus 7-8 mm. longis superne incrassatis elatis; calycibus florentibus anguste turbinatis 3 mm. longis extus dense purpureo-pubescentibus, dentibus del- toideis circiter .5 mm. longis, in maturitate tubo 7.5 mm. longo subeylindrato dentibus vix mutatis conniventibus; corollarum caerulearum tubo 2.5 mm. longo. Mexico: Mexico; Temascaltepec; ad Pungarancho in colle. 16 MADRONO [Vol. 5 Hyptis iodantha sp. nov. per specim. in Mexico prov. Guer- rero in regione Mina prope Zapo in colle a Hinton (no. 9844) lectum constituta est; typum in herb. Univ. Calif. (Los Angeles), isotypum in herb. Kew. vidi. Frutex pulcher altitudine ad 3 m. ramulis villosis et inter flores hirtellis; foliorum laminis ovato-lanceolatis 12-15 cm. longis 5—7 cm. latis supra medium longe acuminatis in basi cor- datis, petiolis 1-2 cm. longis elatis, marginibus irregulariter cre- nato-serratis pagina superiore hirtella inferiore incano-tomentosa molle; floribus numerosis in umbellis purpureis pedunculis 1-2 mm. longis elatis bracteis minutis subtentis in paniculis divari- cato-ramosis 20—25 cm. altis dispositis pedicellis maturis filiformi- bus 5 mm. longis superne incrassatis elatis; calycibus florentibus turbinatis 2.5 mm. longis extus dense purpureo-pubescentibus, dentibus deltoideis vix .5 mm. altis in maturitate conniventibus in maturitate tubo 4.5 mm. longo subcylindrato; corollarum vio- lacearum tubo 2.5 mm. longo. Mexico: Guerrero; Mina; Zapo in colle. University of California at Los Angeles, April 20, 1938. THE VIOLETS OF COLORADO Auice Martiau Sports The genus Viola is a characteristic although a minor element of the flora of Colorado. Viola Nuttalliu, a yellow-flowered per- ennial with lanceolate leaves, and V. Rafinesquii, a small blue- flowered annual, both of which flower in April and May, are fre- quent on the eastern plains and the lower foothills. In shady canyons, at lower altitudes, the long-stemmed white-flowered PV. rugulosa and V. canadensis are important. Montane species are nowhere common. Since the “New Manual of Rocky Mountain Botany” (1909) by Coulter and Nelson is now thirty years old, and “The Flora of Colorado” (1906) and “Flora of the Rocky Mountains and Ad- jacent Plains” (1917) by Rydberg are both out of print it seems desirable to prepare a re-description of the species of violets of Colorado. The descriptions have been drawn from herbarium specimens and from the writer’s field observations during two seasons. The University of Colorado specimens of early collec- tion, which were identified by Ezra Brainerd, formed a valuable beginning for the present study. In the “Flora of Colorado,’ Rydberg lists fourteen species and three varieties of violets for the state. Brainerd in Ryd- berg’s “Flora of the Rocky Mountains and Adjacent Plains” credits Colorado with twenty-one species and one variety. In Brainerd’s later work, “Violets of North America” (Bull. Ver- 1939 | SPOTTS: VIOLETS OF COLORADO 17 mont Agri. Exp. Sta. no. 224. 1921) fourteen species are listed for Colorado. The present paper describes eighteen species and mentions three subspecies and one variety. The nomenclature here followed is that of Brainerd’s treatment in the “Flora of the Rocky Mountains and Adjacent Plains” with the following ex- ceptions: Viola montanensis Rydb. is not considered distinct from V.adunca J. E. Smith (thus agreeing with Brainerd’s later treat- ment); V. scopulorum (Gray) Greene is not considered a distinct species but a variety of V. canadensis L. Grateful acknowledgement is made to Dr. Aven Nelson of the University of Wyoming and to Professor Ernest C. Smith of Colorado State College of Agriculture for loan of material, also to Professor C. L. Porter of the University of Wyoming for his examination and report upon a number of specimens. The writer is indebted to Dr. Francis Ramaley of the University of Colorado for helpful suggestions during the work and for his examination of herbarium specimens at the Missouri Botanical Gardens, Uni- versity of Illinois, and Ohio State University. The writer is also indebted to the curators of the above mentioned herbaria. Mr. Joseph Ewan of the University of Colorado has given valuable advice concerning the manuscript. | In the field, the different species of violets may be recognized chiefly by flower color and habit of growth, but in dried speci- mens the color is often faded and growth habit is not always easy of determination. The key given in this study utilizes as a pri- mary segregating character the presence or absence of evident stems above ground and as a secondary division leaf outline and flower color. The presence or absence of stolons is difficult to use in a key since many specimens are collected before the sto- lons have developed. Violet plants may be glabrous, subglabrous or pubescent, yet pubescence is such a variable character that it is not advisable to rely too much upon it in the segregation of species. The length and outline of the spur is often useful in the identification of a species. The shapes of style and stigma vary in different species but are often distorted by drying. Cleistog- amy is frequent among violets, the late summer or early autumn cleistogamous flowers bearing an abundance of seeds. The char- acters of the cleistogamous flowers may be of taxonomic value but cannot always be made use of since herbarium specimens are often collected at a time when the plants are producing only petaliferous flowers. Fruit and seed characters are often valu- able and are used in the specific descriptions in this paper. Most of the violets occurring in Colorado are widely dis- tributed, about half of the species ranging chiefly to the east and the other half chiefly to the west as far as the Pacific Coast states. One variety and three subspecies are confined to the Rocky Mountain region: Viola canadensis var. scopulorum Gray, V. adunca 18 MADRONO [Vol. 5 subsp. radicosa Baker, V. adunca subsp. Ashtonae Baker, V. palus- tris subsp. brevipes Baker. Viola linguaefolia occurs both in the Rocky Mountain area and in the Pacific Coast states and V. biflora is found in the Rocky Mountains and in Alaska. Most of the spe- cies are rare or local in the state. Only the following are at all common or even frequent: V. Nuttallu, V. Rafinesqui, V. rugulosa, V. canadensis, V. nephrophylla, V. adunca, V. bellidifolia. More species of violets occur in the submontane and montane life zones than in the plains area or in the higher zones; many more grow in moist places than in dry situations. Some of the species are confined to one or two life zones, while others occur in at least three zones. The altitudes indicated in the following tabulation are approximate as they vary in different parts of the state: Plants of plains and lower foothills, from 4000 to 6000 feet. Fields, hillsides or forest openings . .. V. pedatifida, V. pratincola, V. Rafinesquii. | Moist woods or streamsides . . . V. canadensis, V. nephro- phylla, V. retusa, V. vallicola. Plants submontane, from 6000 to 8000 feet. Forest openings or hillsides . . . V. Nuttalli, V. pedatifida, V. pratincola, V. venosa. Moist woods or streamsides ... V. adunca, V. canadensis, V. nephrophylla, V. linguaefolia, V. retusa, V. rugulosa, V. Selkirku, V. vallicola. Plants montane, from 8000 to 10,000 feet. Moist shaded situations ... V. adunca, V. bellidifolia, V. canadensis, V. linguaefolia, V. renifolia, V. rugulosa, V. Selkirkiz. é Bogs, streambanks or wet meadows... V. biflora, V. palustris. Rocky hillsides . . . V. Sheltoni. Plants subalpine and alpine, above 10,000 feet. Stream banks or moist ground .. . V. bellidifolia, V. biflora, V. renifolia. Kry To Cotorapo SPECIES OF VIOLA Plants acaulescent (V. palustris stoloniferous) ; flowers borne singly on a scape; petals violet or white, never yellow. Leaves 2-3 ternately parted, minutely pubescent .... 1. V.pedatifida Leaves entire, glabrous or hirtellous. Flowers white, with purple veins. Leaves cordate; plants of bogs and wet meadows, plants stoloniferous+.. 23.0.0) ote 4a. V. palustris subsp. brevipes Leaves reniform; woodland plants .............. 2. V.renifolia Flowers violet, or lilac. Rootstocks slender; length of petals usually under 10 mm. 1939] Leaves hirtellous on upper surface; spur long, enlarged and rounded at base Leaves glabrous throughout; spur short and thick, plants stoloniferous Rootstocks thick; length of petals usually more than 10 mm. Spurred petal glabrous, rounded at apex; leaves subacute or abruptly pointed...... Spurred petal bearded, emarginate or retuse; leaves obtuse to acuminate. Mature leaves obtuse; spurred petal thickly bearded; sepals obtuse Mature leaves often abruptly acuminate; spurred petal slightly hairy; sepals violet, violet, white, or yellow. Annuals; stipules large, leaf-like, palmately pinnatifid or laciniate; petals pale bluish-violet............. Perennials; stipules small, entire or spinulose-serrate. Leaves divided; flowers pale yellow Leaves entire, denticulate, or shallowly lobed. Flowers yellow, often tinged on the back with purple or brown. First peduncles from axils of upper leaves; leaves cordate-reniform First peduncles arising from near base of plant; leaves various. Plant erect; basal leaves 4-9 cm. long, oblong- lanceolate to ovate Plant ascending or widely spreading to pros- trate; basal leaves 2—7 cm. long, of vari- ous shapes. Plant usually dwarf; peduncles seldom 8 cm.; upper leaf-blades less than 2 cm., 3-lobed, hastate .................... Plant larger; all leaves entire, denticulate, or dentate, never hastate. Leaves linear-lanceolate, subacute, the base tapering into the petiole ........ Leaves truncate to subcordate, obtuse. . Flowers violet, or nearly white with purple veins. Flowers violet; stipules spinulose, serrate, or toothed. Plant dwarf, glabrous; stem short and ob- SCURG We ee ere ee ee Sn ee cg Ne 2 Plant generally more than 5 em. tall, puberu- lent throughout or glabrous; stems nu- LLC OUUS erry ees ee a te. eh, Re eo ca Flowers white, purple veined, with yellow base, purple on back; stipules entire, mostly scarious. Leaves hirsutulous beneath, the root leaves from broadly cordate to reniform; plants SPOTTS: VIOLETS OF COLORADO 10. 11. 12. 13. 14, 15. 16. Le 18. 1. Viota PEDATIFIDA Don, Gard. Dict. 1: 320. phinifolia Nutt. in Torr. & Gray, Fl. N. Amer. 1: Prairie Violet. 19 V. Selkirkii V. palustris V. pratincola V.nephrophylla V.retusa V. Rafinesqui V. Sheltonit V. biflora V.linguaefolia V.venosa V. Nuttall V.vallicola V. bellidifolia V.adunca V.rugulosa V. canadensis 1831. V. del- 136. 188388. 20 MADRONO [Vol. 5 Rootstock ascending; plant without stolons, acaulescent, 7-20 em. high, minutely pubescent or almost glabrous; leaf blades 1-7 em. long, 2—8 cm. wide, 2—3 parted, divisions ternately cleft and incised into linear, obtuse, entire or lobed segments, usually trun- cate or cuneate at the base, margins and mid-rib hirsutulous; peti- oles 3-13 cm. long; stipules ovate-lanceolate, acuminate; petals violet, obovate, 10-20 mm. long, half as wide, lateral petals bearded; spur short; sepals linear or lanceolate, acute or obtuse, 6-10 mm. long, 1-3 mm. wide, margins hirsutulous; peduncles of petaliferous flowers taller than the leaves, of apetaliferous flowers shorter, but erect; cleistogamous capsule light gray when ripe, glabrous, 10-15 mm. long; seeds light brown, 2 mm. long. From Ohio northwest through Wisconsin and Minnesota to Alberta, southwest through the prairie and Rocky Mountain states to New Mexico and Arizona. Colorado: moist grassland forest openings, 5,000 to 10,000 feet; infrequent. April—June. 2. VIoLA RENIFOLIA Gray, Proc. Amer. Acad. 8: 288. 1870. V. Brainerd Greene, Pittonia 5: 89. 1902. Kidney-leaved Violet. Rootstock ascending, commonly slender but becoming thick in old plants; acaulescent, 5-10 cm. tall, pubescent to subgla- brous; stolons absent; leaf-blades rather thin, reniform-cordate, 3—6 cm. long and about as wide, the later leaves sometimes blunt- pointed, margins distinctly crenate-serrate; petioles longer than the blades, often twice as long; stipules linear; petals white, the lower frequently with purplish veins, obovate-spatulate, 6—10 mm. long, either beardless or the laterals with small tufts of hairs; spur short; sepals lanceolate, pale margined, half as long as the petals; peduncles from shorter than to slightly longer than the petioles; capsules ellipsoidal, purplish, 10-15 mm. long; seeds about 1.5 mm, long. Newfoundland to Pennsylvania and west through Michigan and Minnesota to Saskatchewan and northward, also in the Black Hills of South Dakota and in the Rocky Mountains to Colorado. Colorado: wet ground in Engelmann spruce forests, 9,000 to 11,000 feet; rare. May—July. 3. Viota Sevxirki Pursh (ex Goldie) in Edinb. Phil. Journ. 6: 324. 1822. Great Spurred Violet. Rootstock slender, creeping or horizontal; plant without sto- lons, acaulescent, 4-10 cm. high, nearly glabrous; leaf-blades thin, broadly cordate-ovate, short acute or obtuse, 1-4 cm. long, as broad or nearly as broad as long, crenate, upper surface hir- tellous; petioles 1.5—7.0 cm. long; stipules ovate, often toothed at apex; petals pale violet, broadly spatulate or obovate, 5-10 mm. long, half as wide, beardless; spur nearly as long as petals, the end enlarged and round; sepals lanceolate or ovate-lanceolate, acute or somewhat acuminate, 4-6 mm. long, 2-3 mm. wide; 1939 ] SPOTTS: VIOLETS OF COLORADO 21 peduncle 3—7 cm. long; peduncles of cleistogamous flowers erect or ascending; capsule subglobose, 4-8 mm. long; seeds 1.5 mm. long, pale buff. New Brunswick, Nova Scotia, and northward; south through New England to Massachusetts and Pennsylvania; west through Ontario and Minnesota to the mountains of British Columbia and south in the Rocky Mountains to New Mexico. Colorado: moist woods and shady ravines from 7,000 to 10,000 feet; rare. April-— June. 4. Viova patustris L., Sp. Pl. 934. 1753. Marsh Violet. Rootstock slender, scaly, creeping or horizontal; plant sto- loniferous, acaulescent, 5-15 cm. tall, glabrous throughout; scapes generally surpassing the leaves, bracteolate at the middle or be- low; leaf-blades thin, ovate-cordate to orbicular or reniform, 2—4 em. long, generally wider than long, the margin crenulate; peti- oles often twice as long as blades; stipules lanceolate, sometimes with a few subulate teeth at apex; petals pale lilac to nearly white with darker veins, obovate or broadly spatulate, 6-10 mm. long, the lateral petals bearded; spur short, thick and rounded; sepals ovate to ovate lanceolate, narrowly white-margined, 5-6 mm. long, obtuse; seeds dark brown, 1.5 mm. long. Labrador south through the mountains of New England, west to Alaska, British Columbia, Washington, Oregon, and northern California, also in South Dakota and the Rocky Mountains to Colorado; Asia and Europe. Colorado: wet or moist soil in shady situations from 7,000 to 10,000 feet; infrequent. May- August. 4a. VIOLA PALUSTRIS subsp. BREvIPES M. S. Baker, Madrofio 3: 235. 19386. ; This subspecies differs from the species in its smaller size and the pure white fragrant flowers with lateral petals entirely beard- less. Estes Park and Moraine Park, Colorado. 5. VIoLA PRATINCOLA Greene, Pittonia 4: 64. 1899. V. Sand- bergu Greene, Pittonia 5: 119. 1908. Meadow or Hooded Blue Violet. Rootstock ascending; plant without stolons, acaulescent, 7-30 em. high, usually glabrous; leaf-blades deeply cordate-ovate tapering gradually to subacute apex or sometimes abruptly pointed, 2-3 cm. wide at petaliferous flowering, 5-8 cm. wide at maturity, as long as wide, crenulate; petioles 2 or 3 times as long as the blade; stipules linear-lanceolate; petals violet, light at base, spatulate, 10-15 mm. long, 4-6 mm. wide, lateral petals somewhat bearded; spurred petal glabrous, rounded at apex; spurs short; sepals ovate-lanceolate, acute or obtuse; peduncles as long as or much longer than the leaves, bracteolate at or above the middle; cleistogamous flowers on short prostrate peduncles; capsules ellipsoidal, 6-10 mm. long; seeds 1.5—2.0 mm. long. 22 MADRONO [Vol. 5 Minnesota and North Dakota to Wyoming and Colorado. Colorado: prairies and hills; open situations, 4,000 to 8,000 feet; infrequent. April-May. 6. VIoLA NEPHROPHYLLA Greene, Pittonia 3: 144. 1896. V. cognata Greene, Pittonia 3: 145. 1896. Rootstock generally ascending, moderately thick; plant with- out stolons, acaulescent 5—15 cm. tall, nearly or quite glabrous; leaf-blades firm, broadly cordate-ovate to reniform, obtuse, 1-3 cm. long, 1-4 em. broad, crenate-serrate, late-season leaves larger; petioles slender, two or three times the length of the blades; stipules ovate-lanceolate; petals violet, lighter at base with purple veining, obovate, large, 10-15 mm. long, half as wide, pubescent to villous, the spurred petal villous, emarginate; spur rounded, about one-fourth the length of petals; sepals ovate to oblong, often rounded, 4-8 mm. long, 2-83 mm. wide; peduncles slender, 4-12 cm. long; cleistogamous flowers ovoid on slender short peduncles; capsules short-ellipsoidal, green, glabrous, 8—10 mm. long; seeds 2.0—2.5 mm. long, olive-brown. South Dakota and west to British Columbia; south in moun- tain areas to New Mexico, Arizona, Utah, Nevada, and California. Colorado: low meadows, moist aspen groves, and sloping ground among bushes, 5,000 to 10,000 feet; moderately frequent. May- July. This is the V. nephrophylla of Brainerd’s account in Rydberg’s “Flora of the Rocky Mountains and Adjacent Plains’ (1917). I have seen no specimens which I could refer to V. nephrophylla as originally described by Greene from western Colorado. Plants of this species are conspicuous because of their firm leaf-blades, long slender petioles and peduncles, and large flowers. 7. VioLta RETUSA Greene, Pittonia 4: 6. 1899. Western Blue Violet. Rootstock ascending, stout; plant without stolons, acaules- cent, 4-15 cm. high, glabrous throughout; leaf-blades at spring flowering broadly ovate or cordate-deltoid, acute or obtuse, 2—4 cm. long, 2-6 cm. wide, finely serrate; aestival leaves wider, reni- form, often abruptly acuminate, the base cordate to truncate-de- current, crenate-serrate; petioles 1-3 times as long as blades; stipules ovate, acuminate, sometimes toothed at apex; petals violet, the upper obovate, 10-15 mm. long, 5-6 mm. wide, lateral petals spatulate, somewhat bearded, spurred petal slightly hairy, retuse; spurs short and rounded; sepals lanceolate, white-mar- gined, acute, 5-6 mm. long, 1-2 mm. wide; peduncle somewhat exceeding the leaves; cleistogamous flowers ovoid, on erect peduncles; capsules ellipsoidal, green, 10-12 mm. long; seeds brown, 2 mm. long. Central Kansas, northwest and west to the mountain front and 1939 ] SPOTTS: VIOLETS OF COLORADO 23 foothill districts of Colorado. Colorado: shady thickets and along streams from 4,000 to about 7,000 feet; moderately fre- quent. May—June. 8. Viota RaFinesquu Greene, Pittonia 4: 9. 1899. V. bicolor Pursh, Fl. Am. Sept. 1: 175. 1814, not V. bicolor Gilib. ‘1781. V. tenella Raf. in Medical Repository, N. Y. ser. 2, 5: 354. 1808 (nomen); Am. Monthly Mag. 191. 1819, not V. tenella Poir. 1810. Wild Pansy. Annual, rootstock ascending, plant caulescent, 1-2 dm. high, lower part of stem somewhat pubescent with recurved hairs, be- coming nearly glabrous with age; stems slender, often branching from base; lower leaf-blades suborbicular, 3-10 mm. long, upper oblong-lanceolate to ovate, 10-20 mm. long, 3—5 mm. wide, entire or crenate-dentate, attenuate at the base, margins minutely pubes- cent; petioles of lower leaves about twice as long as blades, of upper leaves one-third as long as blades; stipules foliaceous, 5—14 mm. long, palmately pinnatifid or laciniate, terminal segment elongate; petals cream-colored, blue or purplish spotted, obovate, 4-9 mm. long, 2—4 mm. wide; spur very short; sepals subulate, about half as long as petals, margins hirsutulous; peduncle 1.5- 3.0 cm. long; capsule glabrous, 6 mm. long or less; seeds light brown, 1.5 mm. long. New York to Michigan, south to Georgia, west through the Mississippi basin to Colorado and Texas. Colorado: moist slop- ing ground of plains and lower foothills, 4,000 to 7,000 feet; locally frequent. April—May. 9. Viota SHEtToni Torr., Pacif. Rail. Rep. 4: 67, t.:2. 1857. V. biternata Greene, Pl. Baker. 3:12. 1901. Fan Violet. Rootstock ascending, deep-seated; plant subacaulescent to caulescent, 5-15 cm. or taller, hirsutulous or nearly glabrous; leaf-blades 1-4 cm. long, and fully as wide, palmately 3-divided, the divisions palmately or pedately 3-parted and cleft; petioles 38-10 cm. long or longer; stipules lanceolate to ovate, scarious; petals yellow, strongly veined, tinged with brown beneath, ob- ovate, 6-12 mm. long, half as wide as long, lateral petals slightly bearded; spur hardly one-third the length of the petals; sepals linear-lanceolate, acuminate, 4-8 mm. long; peduncles 5-15 cm. long; capsule globose, 6-8 mm. long, brown, glabrous; seeds buff, 2 mm. long. Western Colorado to Washington, Oregon, and California. Colorado: rocky hillsides of the western slope of the Rocky Mountains from 7,000 to 9,000 feet; infrequent to rare. April-— June. 10. Viota Birtora L., Sp. Pl. 936. 1753. Alpine Yellow Violet. Rootstock ascending; plant caulescent, 5-20 cm. high; stems 24 MADRONO [Vol. 5 slender, generally 2-leaved and 2-flowered; leaf-blades round- reniform, narrowly cordate at base, rounded at apex, 1-2 cm. long, twice as wide as long, somewhat hirtellous on upper surface and margins, crenulate; stipules ovate; petioles 3-8 cm. long; petals yellow streaked with dark purple-brown, spatulate, 6—8 mm. long; half as wide as long; spur short-conical; sepals linear- oblong, 4 mm. long; peduncles surpassing the leaves; capsule about 6 mm. long, ellipsoidal; seeds about 1 mm. long. Northern Europe and Asia to Alaska; Colorado. Colorado: wet moss and dense herbaceous vegetation in fine alluvial soil along mountain streams from 8,000 to 11,000 feet; rare. April— July. 11. Viota tineuarFo.ia Nutt., in Torr. & Gray Fl. N. Am. 1: 141. 1888. V. flavovirens Pollard, Bull. Torr. Bot. Club 24: 405. 1897. V. erectifolia Nels., Bot. Gaz. 29: 143. 1900. V. gompho- petala Greene, Pl. Baker. 3:11. 1901. Rootstock ascending, slender; plant caulescent or subcaules- cent, erect, 1-3 dm. high, hirtellous to nearly glabrous; leaf- blades narrowly oblong to ovate-lanceolate, mostly obtuse, 5—9 cm. long, 1.5—3.0 cm. wide, entire or repand-denticulate, margins ciliate, veins generally hirtellous; petioles about twice the length of leaf-blades; stipules lanceolate, acuminate; petals yellow, veined with purple, often appearing reddish-brown in dried speci- mens, 10-15 mm. long, 5-8 mm. wide, upper petals obovate, lateral petals spatulate, usually somewhat bearded; spur very short; sepals linear-lanceolate, acuminate, 5-8 mm. long, margins usually ciliate; peduncle generally as long as or surpassing the leaves; late flowers cleistogamous from upper axils; capsule glo- bose, usually glabrous; seeds 2-3 mm. long. Montana to Colorado, west to Washington and northern Cali- fornia. Colorado: mountain valleys or coarse soil near lakes, 8,000 to 10,000 feet; infrequent. June—August. 12. Viota veNosa (S. Wats.) Rydb., Mem. N. Y. Bot. Gard. 1: 262. 1900. V. Nuttallit var. venosa Wats., Bot. King Exped. 35. 1871. V. atriplicifolia Greene, Pittonia 3: 38. 1896. V. Thoriu Nels., Bot. Gaz. 80: 198. 1900. Dwarf Yellow Violet. Rootstock vertical or nearly so; plant caulescent, 5-15 cm. tall, glabrate to pubescent; stems many, ascending, short; lower leaf-blades ovate, obtuse, 1-2 (seldom 3) cm. long and two-thirds as wide, coarsely round-toothed or lobed and tapering into a long margined petiole, the upper leaves often three-lobed hastate ; stipules lanceolate, entire; petals yellow tinged with purple dor- sally (often quite purple in drying), obovate, 6-10 mm. long and half as wide, usually beardless; spur short; sepals hirtellous to pubescent, lanceolate, white margined, one-third to one-half the length of the petals; peduncles 2—5 cm. long, bracteolate half- way up; capsule nearly globose, 4-5 mm. long; seeds about 1 mm. long. 1939 | SPOTTS: VIOLETS OF COLORADO 25 Montana and Wyoming, west through Idaho and Utah to Washington, Oregon and California. Colorado: northwestern part of the state on dry hillsides and in openings of dry conifer- ous forest, 6,000—8,000 feet. April—May. 138. Viora Nurratytyu Pursh, Fl. Am. Sept. 1: 174. 1814. Nuttall Violet. Rootstock ascending, semi-fleshy, occasionally branched or somewhat fascicled; plant caulescent, 5-25 cm. high, pubescent or glabrate to nearly glabrous; stems numerous, spreading rather than erect, generally short unless in a moist situation; leaf-blades oblong-lanceolate or linear-lanceolate, acute or subacute, 2—7 cm. long, 6-20 mm. wide, some puberulent, entire or repand-denticu- late, tapering into margined petioles 3-12 cm. long; stipules lanceolate, entire; petals yellow and at times tinged with reddish- purple on outside, obovate, 8-12 mm. long, half as wide, slightly bearded; spur short; stigma bearded; sepals lanceolate or linear, acuminate, 5-10 mm. long; peduncle 3-8 cm. long; capsule sub- globose, 7 mm. long, brown; seeds brown, 2.5—3.0 mm. long. Manitoba and North Dakota to Missouri and Kansas, south in the Rocky Mountain states to New Mexico and Arizona. Colo- rado: prairies, plains, and foothills; open situations from 4,000 to 8,000 feet; common. April—June. 14, Vioxva vautiicota Nels., Bull. Torr. Bot. Club 26: 128. 1899. V. physalodes Greene, Pl. Baker. 8:12. 1901. Rootstock short, with fascicled fleshy roots, ascending; plant caulescent, low, 4-12 cm. tall, glabrous or pubescent; stems some- what spreading; leaf-blades ovate to lanceolate, obtuse, base rounded, basal leaves often subcordate, 2—5 cm. long, 1.0—2.5 cm. broad, generally puberulent with ciliate margins, entire or ob- securely denticulate; petioles as long as the blades or longer; stipules lanceolate; petals bright yellow streaked with purple, obovate or broadly spatulate, 8-12 mm. long, half as wide, beard- less or slightly bearded; spur short; sepals linear-lanceolate, acuminate, 5-7 mm. long; peduncle shorter than leaves or sur- passing them; capsule broadly ovoid, 5—7 mm. long; seeds 1.5—2.0 mm. long. North Dakota, Saskatchewan, British Columbia, and Wash- ington, south in the Rocky Mountain area to Colorado. Colo- rado: moist valleys, 5,000 to 8,000 feet; infrequent. May—June. 15. VioLa BELLIDIFOLIA Greene, Pittonia 4: 292. 1901. VJ. demissa Greene, Pl. Baker. 3:10. 1901. Subalpine Blue Violet. Rootstock erect or ascending; plant glabrous, dwarf and tufted, 2-6 cm. tall, subacaulescent, the stems short and obscure; leaf-blades round-ovate, 5-16 mm. long, nearly as broad, obtuse at apex, obscurely cordate at base, entire or crenulate; petioles 1-4 cm. long; stipules linear-lanceolate, with a few bristly teeth; 26 MADRONO [Vol. 5 petals violet-purple above, lighter below, obovate, 5-10 mm. long, 3-6 mm. wide; lateral petals slightly bearded; spur half as long as petals, often curved; sepals oblong-lanceolate, 3-5 mm. long, 0.5-1.0 mm. wide; peduncles numerous, 1.5—7.0 cm. long, gen- erally surpassing the leaves, bracteolate above the middle; cap- sule subglobose, 5 mm. long; seeds 1.5 mm. long. British Columbia, Montana and Idaho to Colorado; also in northern California. Colorado: mountain areas from 8,000 to 12,000 feet and higher, often in dense mixed herbaceous vegeta- tion; rather frequent. June—August. 16. Viota apunca J. E. Smith, in Rees, Cycl. 37: pl. 638. 1817. V. subvestita Greene, Erythea 5: 89. 1897. V. montanensis Rydb., Mem. N. Y. Bot. Gard. 1: 263. 1900. V. odontophora Rydb., Mem. N. Y. Bot. Gard. 1: 164. 1900. V. inamoena Greene, Pl. Baker. 3:11. 1901. V.retroscabra' Greene, Pittonia 4: 290. 1901. Hooked Violet. Rootstock ascending; tap root long; plant caulescent, 4—25 em. high, glabrous to glabrate or puberulent or scaberulous; stems numerous; leaf-blades round-ovate or subcordate-ovate, some often ovate-trigonous, 1-4 cm. long, often as broad or nearly as broad as long, crenulate, truncate or cuneate at base, upper leaf-blades narrower; petioles 1-6 cm. long; stipules linear-subulate or linear-lanceolate, nearly entire, toothed or spinulose-serrate; petals violet-purple, obovate or broadly spatu- late, 7-12 mm. long, 4-6 mm. wide, lateral petals bearded; spur 4—7 mm. long, straight, curved, or hooked; sepals linear-lance- olate, 5-7 mm. long, 0.5-1.0 mm. wide; peduncles 2—8 cm. long, often surpassing the leaves, bracteolate above the middle; cap- sule ellipsoidal, 6-7 mm. long; seeds 1.5—2.0 mm. long. New Brunswick, eastern Quebec, and Vermont to Alaska, British Columbia, and California, south in the Rocky Mountain area to Colorado, Utah, and New Mexico. Colorado: hills and valleys; shaded situations, 5,000 to 10,000 feet; frequent. May-— July. 16a. Viota apuNca subsp. AsutonaE M.S. Baker, Madrofio 8: 233. 1936. This subspecies differs from the species in the gla- brous herbage, the somewhat smaller and paler corollas, the ob- long-ovate to oblong-lanceolate (not linear-lanceolate) sepals, the short spur and the emarginate capsule. Cub Lake Trail, Estes Park, Colorado. 16b. Viota apuNcA subsp. rapicosa M. S. Baker, Madrofio 3: 234. 1936. This subspecies differs from the species in having dense short mostly retrorse pubescence throughout, the leaves often acuminate with the base tending to be truncate, and the capsule notched. Kawuneeche Valley, twelve miles north of Grand Lake, Colorado. 17. Viota rucuLosa Greene, Pittonia 5: 26. 1902. V. Ryd- bergit Greene, Pittonia 5: 27. 1902. Rydberg Violet. 1939 ] SPOTTS: VIOLETS OF COLORADO ; 27 Rootstock ascending; plant caulescent, 1.5—4.0 dm. high, pubescent, stoloniferous; leaf-blades thickish, cordate-reniform, abruptly short-pointed, 1-7 cm. long, 2-10 cm. wide, serrate, hirsutulous especially beneath, upper leaves smaller and more often ovate-acuminate; petioles of root-leaves 3-20 cm. long, upper leaves with short petioles, 1-12 mm. long; stipules entire or nearly so, lanceolate, obliquely acuminate, not wholly scari- ous; petals violet or nearly white, purple veined, broadly spatu- late or obovate, 6-12 mm. long, half as wide, lateral petals bearded; spur short; sepals subulate pointed, 3-7 mm. long, 1 mm. or less wide, minutely pubescent, margins white; peduncle 1.5—5.0 cm. long; cleistogamous flowers often appearing later; capsule ovoid to subglobose, minutely pubescent or nearly gla- brous, 6-10 mm. long, greenish-brown; seeds brown, 1.5—2.0 mm. long. | Manitoba, Minnesota and Iowa, west to British Columbia, also in the Rocky Mountain area through Idaho, Montana, Wyo- ming, and Colorado to Arizona. Colorado: woods and shady streamsides from 5,000 to 9,000 feet; locally common. May, July or October. This species is distinguished from the similar V’. canadensis by the presence of stolons and the broader leaf-blades. 18. Viota caNaDENsIs L., Sp. Pl. 986. 1753. V. neomeaicana Greene, Pittonia 5:28. 1902. Canada Violet. Rootstock ascending; plant caulescent, 1.5—-3.5 dm. high, nearly glabrous; leaf-blades thin, cordate-ovate, acuminate or acute, 1.5—6.0 cm. long, 1.5—5.0 cm. wide, serrate, teeth incurved, veins of under surface sometimes pubescent, smaller upper leaves often muriculate to hirtellous; petioles usually longer than blades, those of lower leaves often twice as long; stipules entire, lanceo- late, obliquely acuminate, not wholly scarious; petals violet or nearly white, purple veined, broadly spatulate or obovate, 6—14 mm. long, 3-8 mm. wide, lateral petals bearded; spur short; sepals subulate pointed, 4—7 mm. long, 0.5—1.0 mm. wide, minutely pubescent, margins white; peduncles 1-8 cm. long; capsules ovoid or subglobose, mostly glabrous, 4-6 mm. long, brown; seeds brown, about 2 mm. long. New Brunswick to South Carolina and Alabama, west and northwest through Ohio, Wisconsin, and South Dakota to north- ern Washington and Saskatchewan, south in the Rocky Mountain states through Colorado and Utah to New Mexico and Arizona. Colorado: moist woods and mountain slopes from 5,000 to 10,000 feet; frequent to common. April—June. 18a. VioLa CANADENSIS L. var. scopuLtorUM Gray, Bot. Gaz. 11: 291. 1886. V.scopulorum (Gray) Greene, Pittonia 5:27. 1902. A tufted and somewhat depressed form with small leaves and elongated peduncles, known only from Colorado. University of Colorado, Boulder, January, 1938. 28 MADRONO [Vol. 5 THE GENERA OF THE TRIBE HYDROPHYLLEAE OF THE HYDROPHYLLACEAE LincoLN CONSTANCE A systematic study of Nemophila, which was undertaken with the objective of offering yet another revision of that genus, re- vealed not only difficulties in delimiting the species but also that lines traditionally accepted to separate the genera of the tribe Hydrophylleae did not, in certain cases, conform to natural boundaries. Hence, the study was broadened to include all those plants which have customarily been referred to the tribe, but con- cerned itself especially with Ellisia and Nemophila. The facts appear to necessitate a general realignment of these genera be- fore monographic treatments of the several groups of species can be profitably presented. Asa Gray (7) was the first writer to organize the three genera, Hydrophyllum L., Ellisia L. and Nemophila Nutt., into the tribe Hydrophyleae. He characterized the group principally by the possession of a unilocular ovary, largely filled at certain stages of development by two large fleshy parietal placentae, upon which the ovules are produced, and also by the presence of a single, more or less bifid style. This definition of the tribe has remained substantially unchanged to the present time, and is so accepted by Bentham and Hooker (4), Peter (15) and Brand (5). Some species have been removed by certain authors from each of the three genera under the segregates, Eucrypta Nutt., Decemium Raf. and Pholistoma Lilja, but conservative botanists have, in the main, retained not only the tribal grouping but also the three classical genera. The substitution of Macrocalyx Trew and Nyctelea Scop. for Ellisia and of Viticella Mitch. for Nemophila was based upon purely nomenclatorial considerations and can be disregarded in this discussion inasmuch as Ellisia has been con- served and Nemophila is on the list of nomina generica conservanda proposita. Nemophila has been in common use for more than a century and its retention also is to be advocated in the interest of nomenclatorial stability. Hydrophyllum, distinguished by its perennial or biennial habit, usually basal but always alternate leaves, many-flowered, semi- scorpioid and usually congested inflorescence and exserted sta- mens, has been little confused with either of the other two genera, which are annual with chiefly cauline leaves and usually at least the lowest pair opposite, while the flowers are solitary or few in loose, raceme-like cymes and the stamens are included. Brand (5) has taken up Decemium (treated by Gray as a section of Hydrophyllum) and has constituted it a monotypic genus based upon H. appendiculatum Michx., which differs from members of section Euhydrophyllum Gray chiefly in possessing appendaged calyx-sinuses, an accrescent calyx, a biennial habit and only 1939] CONSTANCE: HYDROPHYLLEAE 29 slightly exserted stamens. Because of the sporadic occurrence of these calycine conditions without close conformance to natural generic lines elsewhere in the tribe, and because the other two features appear to be scarcely of generic importance, it would seem advisable to retain this species within Hydrophyllum. Various sets of characters have been drawn into service to distinguish Ellisia from Nemophila, but the difference most com- monly emphasized—and the only one employed by Brand (5) in his generic key—is the presence in Nemophila of appendaged calyx-sinuses, as opposed to the absence of any such auricles in Ellisia. This lack of appendages on the calyx, indeed, is the only feature, save tribal similarities, possessed in common by all the species usually combined into Ellisia. Bentham (3), who first contrasted Ellisia with Nemophila, laying chief emphasis upon the presence or absence of auricles, clearly questioned the value of this distinction in the following words: ee . in some instances the sinuses (as in some Campanulaceae) are furnished with reflexed appendages, resembling the erect divisions of the calyx in form, but smaller in size. ... these divisions do not indicate any organic modifica- tions in the composition of the calyx, but are merely owing to the prolongation of the united lateral nerves of two adjoining sepals. . . . The character derived from this circumstance must consequently be inconstant, and have little or no relation to general habit . . . and, if that be really the only distinction between Nemophila and Ellisia, it proves the expediency of uniting these two genera ..- Nemophila and Ellisia, when taken together, are a natural group, but are separated by a purely artificial character.” However, Bentham kept the genera separate in this and in a later (4) treatment. The two genera were actually merged by Baillon (2) who, without comment but doubtless influenced by Bentham’s remarks, placed all the species of Nemophila under Ellisia, but did not make the necessary nomenclatorial recombina- tions. Chandler (6), in his excellent revision of Nemophila, noted that, “The question of the relation of Nemophila to Ellisia is vital, but must be left for future consideration.” The auricles of certain species of Nemophila, particularly N. pulchella, are frequently obsolete, so that Brand (5) cited under both genera specimens referable to this one species. Many specimens of Ellisia Nyctelea, type species of its genus, show occa- sional sepaloid teeth which are scarcely distinguishable from auricles. Additional distinctions which have been used to separate the two genera are these: (1) the non-enlargement of the fruiting- calyx in Nemophila, its marked accrescence in Ellisia; (2) the presence of a deciduous cap of colorless cells (the “caruncle,” “cealyptra” or “cucullus’’) at the chalazal end of the mature seed in Nemophila, its complete absence in Ellisia; (3) the presence of corolla-scales in Nemophila, a pair at the base of each filament, as opposed to their obsolescence or absence in Fllisia; and (4) the fact that the corolla usually exceeds the calyx in Nemophila but 30 MADRONO [Vol. 5 is shorter than the calyx in Ellisia. A detailed consideration of this formidable array of differences, however, leads to the dis- covery of important contradictions in all of them. The calyx of Nemophila breviflora, N. Kirtleyi and N. phace- lioides, especially, is more strongly accrescent than that of Ellisia chrysanthemifolia or E. micrantha, whereas the calyx of all species of both genera enlarges to some degree in fruit. Nemophila aurita and N. racemosa entirely lack the distinguishing cucullus on the seed, and that on the seed of N. breviflora, N. microcalyx and N. phacelioides is reduced and persistent. The interstaminal scales in species of Ellisia are often more conspicuous than the squamae in certain collections of Nemophila, many of whose species have forms with the scales reduced to pubescent lines or quite obsolete. The corolla of N. breviflora is definitely shorter than the calyx, that of N. parviflora, N. pedunculata and varieties of N. pulchella often scarcely exceeds it, and that of Ellisia mem- branacea and E. chrysanthemifolia is distinctly longer than the calyx. None of these characters, then, is without striking excep- tions which would seem to diminish the height of the traditional generic barriers. Whereas Nemophila has not been sectionally divided by any- one, its members usually being keyed out on artificial characters, Ellisia has been recognized as a compound group consisting of section Euellisia Gray and section Eucrypta (Nutt.) Gray. The chief criteria employed by Nuttall (14) in founding Eucrypta as a genus were, first, its unique habit of bearing ovules upon both outer and inner faces of the placentae, and, secondly, the produc- tion of two very distinct types of seed in the inner and outer chambers thus formed. The subsequently discovered EF. mi- crantha breaks down the second character because its seeds are homomorphic, but Brandegee’s detection of the occasional pres- ence of ovules on the side of the placentae away from the true locule has led to the association of this species with those which were known to Nuttall and included by him in Eucrypta. The fact that Bentham described Ellisia (Eucrypta) chrysanthemifolia as a congener of EF. (Euellisia) Nyctelea without noting the pres- ence of the “hidden” ovules and continued to maintain this posi- tion for the species even after this peculiarity had been pointed out, undoubtedly exerted a marked effect upon treatments by contemporary and later botanists. The fruit and seed characters of Eucrypta are, in the writer's opinion, of more diagnostic weight than the presence or absence of appendaged calyx-sinuses in determining natural generic lines, as is admirably shown by the fluctuation of the auricle character within Nemophila and Hydrophyllum. Green (8) has adequately summed up the reasons for restoring Eucrypta to generic rank, in the following statement: “These plants are not at agreement with Hilisia in habit. But if they were, capsules of such remarkable structure, and with seeds of two sorts so 1939 | CONSTANCE: HYDROPHYLLEAE 31 strikingly dissimilar, neither sort answering to those of Ellisia or of any other Hydrophyllaceous genus, must, it seems to the writer, establish strongly enough a genus which was long ago well defined by an eminent authority. The name (meaning ‘well hidden’) is very admirably appropriate; for the pair of flat- tened seeds (rarely by the abortion of one ovule, solitary) which lie between the wall of the valve and its placenta, are so closely sealed as to have escaped the detection of that great botanist, the late Mr. Bentham, into whose hands one or both of the species fell at an earlier date than that of Mr. Nuttall’s treatment of them, and who therefore described the plant as if it had been a real Ellisia.” In addition, the species which possess this peculiar condition of ovule formation also agree closely in their usually delicate habit, in the viscid and aromatic condition of their dissected, fern-like foliage, in the hispid pubescence, in their relatively complicated cymose inflorescence and in their narrow and rather shallowly divided calyx and corolla. Their distribution, chiefly in the arid portion of the southwestern United States, is comparable and suggests a common Mexican origin for the group. In the light of the foregoing arguments, the writer proposes to follow Nuttall (14), Greene (8), Heller (10, 11), Abrams (1) and Rydberg (16) in recognizing Eucrypta as a valid genus. Because fruit and seed characters have been given so much weight in the genus Phacelia, and especially in the neighboring family Boraginaceae, the writer anticipated that something of value might be revealed by a comparison of the seeds of those spe- cies comprising Ellisia section Euellisia with those of Nemophila. The seeds of Ellista Nyctelea, E. membranacea, Nemophila aurita and N. racemosa were found to be nearly globose and regularly reticulate or alveolate, and entirely devoid of any cucullus. Those of the remaining species of Nemophila, on the other hand, were mostly ovoid, smooth or tuberculate and variously pitted, but never either reticulate or alveolate, and always with a cu- cullus. The capsules of Ellisia membranacea, Nemophila aurita and N. racemosa are armed with stout prickles and bristles, a condition not duplicated elsewhere in the tribe Hydrophylleae. Furthermore, these three species agree in the possession of a peculiar scandent habit, the prickly armature of the stems and a closely comparable geographic range, which again points to a southern origin. These facts appear to confirm the suggestion that this group is a natural one. The close resemblance of the three species was intimated by Greene (9), who transferred Ellisia membranacea to Nemophila with the remark that this species is, “Thoroughly congeneric with N. racemosa; only empirically placed under ‘Ellisia, notwithstanding the absence of calyx- bractlets.” Similarly, Jepson (12), while restoring this species to Ellisia, referred to the same plant as, “In vegetative habit strikingly similar to Nemophila aurita.” The generic name Pholistoma was proposed by Lilja (13) with Nemophila aurita Lindl. as its type species. Although he based the genus chiefly upon minor characters, the name was effectively 32 MADRONO [Vol. 5 published and a recognizable type species designated, so that it is available for further use. The writer proposes to take up Pholistoma for Ellisia membranacea, Nemophila aurita and N. race- mosa (appropriate recombinations are to be included in a further paper on Pholistoma) because if they were thrown, collectively, into either Nemophila or Ellisia, because of their resemblance to certain species of each genus, the generic characters would be so weakened that Nemophila and Ellisia would have to be re- garded as a single, unwieldy group, to be known henceforth by the Linnean name of Ellisia. Their segregation, it is believed, will achieve a more natural arrangement within the tribe, and free both Ellisia and Nemophila of discordant elements. With the removal of section Eucrypta and the transference of one species to Pholistoma, Ellisia is restricted to the original species, £. Nyctelea, and so becomes a monotypic genus as it was known to Linnaeus. Its globose reticulate seeds are closely similar to those of Pholistoma, but it lacks the scandent habit, possesses a pubescent rather than a prickly stem, an unarmed capsule, and a very small, tubular-campanulate corolla. The predominantly temperate eastern American distribution of Ellisia Nyctelea, although outlying stations occur in the plains area of western Montana and of northeastern New Mexico, contrasts strikingly with the southwestern occurrence of Pholistoma, and indicates that the separation has been one of long duration. The fact that the corollas are narrowly campanulate and usually shorter than the calyx suggests an affinity, pointed out by Ryd- berg (16), with Nemophila breviflora, which, however, has quite different pitted and cucullate seeds, and with Eucrypta, which differs in its viscid herbage, more numerous flowers, weakly enlarging calyx, falsely pentalocular capsule, smooth or corru- gated seeds and its widely different ecological and geographical range. The maintainence of Ellisia as a monotypic genus, then appears to be a satisfactory alternative to the dubious policy of including the very unlike species of Nemophila, Pholistoma, Ellisia and possibly even Eucrypta within the same genus. If all of these were merged, it would be exceedingly difficult to justify the exclusion of Hydrophyllum from the amorphous group which thus would be created. Pollen morphology and chromosome number, it was hoped, might afford additional evidence as to phylogenetic relationships within the tribe. Preliminary and unpublished results seem to indicate, however, that a general similarity in microspore struc- ture and in chromosomal complement prevails throughout the group. The foregoing discussion can now be summarized in the form of a key to the genera as they are accepted by the writer, and which will be treated individually in revisionary papers which are either completed or in the course of preparation. 1939 | CONSTANCE: HYDROPHYLLEAE 33 Key TO THE GENERA Perennial or biennial; leaves chiefly basal and all alter- nate; flowers numerous in scorpioid and often con- gested cymes; stamens exserted ................-.. 1. Hydrophyllum lL. Annual; leaves chiefly cauline, at least the lowest pair usually opposite; flowers solitary or several in loose simple or panicled cymes; stamens included. Herbage variously pubescent, bristly, prickly or gla- brate, but neither viscid nor scented; ovules borne only on the axial face of the placentae. Seeds lacking a cucullus, nearly globose and regu- larly reticulate or alveolate. Succulent, scandent herbs with prickly stems; co- rolla exceeding calyx; capsules armed with stout DricklesvOPr -OLISUIES 4.2% asda 2 tener hee sas ows 2. Pholistoma Lilj. Flaccid herbs with hispid or glabrate stems; co- rolla exceeded by calyx or barely equalling it; capsules pubescent but unarmed .............. 3. Ellisia L. Seeds cucullate, mostly ovoid and smooth or tubercu- late, often pitted and scrobiculate, but neither reticulate nor alveolate ....................... 4. Nemophila Nutt. Herbage viscid and scented; ovules borne on both axial LS and abaxial faces of the placentae ............... 5. Hucrypta Nutt. Department of Botany, University of California, Berkeley, August 28, 1938. LITERATURE CITED Asrams, LERoy. Flora of Los Angeles and vicinity. 321. 1904. Baitton, H. E. Histoire des plantes 10: 397. 1891. BENTHAM, G. Review of the order of Hydrophylleae. Trans. Linn. Soc. London 17: 265-274. 1834. BentTHAM, G., and Hooker, J. D. Genera plantarum 2: 825-827. 1876. Branp, A. Hydrophyllaceae. In Engler, Das Pflanzenreich IV. 251. 29- 57. 1913. Cuanpiter, H. P. A revision of the genus Nemophila. Bot. Gaz. 34: 195. 1902. Gray, A. A conspectus of the North American Hydrophyllaceae. Proc. Amer. Acad. Sci. 10: 312. 1875. . Greene, E. L. Studies in the botany of California and parts adjacent II. Bull. Calif. Acad. Sci. 1: 199-200. 1885. . Manual of the botany of the region of San Francisco Bay. 252. 1894. . Hetrer, A. A. Catalogue of North American Plants north of Mexico. 7. 1898. Botanical exploration in California. Muhlenbergia 2: 163, 233. 1906. Jepson, W.L. A flora of western middle California. 435. 1901. [Lirsa, N.] Tvenne nya vextsligten beskrifne of N. Lilja. Lindblom, Botaniska Notiser 4: 39-40. 1839. . Nouttatr, T. Descriptions of plants collected by William Gambel, MD, in the Rocky Mountains and Upper California. Journ. Philadelphia Acad. Sci., ser. 2, 1: 158-159. 1847. . Peter, A. Hydrophyllaceae. In Engler and Prantl, Das Pflanzenfamilien IV. 3b. 5471. 1895. . Rypserc, P. A. Catalogue of the flora of Montana and of Yellowstone National Park. Mem. N. Y. Bot. Gard. 1: 323. 1900. 34 MADRONO [Vol. 5 A NOTE ON THE OCCURRENCE OF SALVIA IN THE NEW WORLD Cart EruineG The subgenus Calosphace of Salvia includes nearly five hun- dred species, and, due to the localized nature of many of the species, each venture from the beaten path within the tropics brings new species to light. This large group is homogeneous but nevertheless separable into a series of species constellations ninety-two in number. These vary in size from monotypic groups to groups of forty spe- cies or more. Each gives the impression of an immediate origin from a single source through means other than hy- —a bridization, as though through CSE a i a ae es: a spe- y cies. In general the species --= are TT which Here them ie of 1 PEO the same growth form with ‘ . essentially similar habits of inflorescence. They have flow- ers of essentially the same conformation, which differ nevertheless in size and pro- portion. Such constellations differ among themselves by differences in habit, in inflores- cence and in the details of flower structure. Each sug- gests a species group of close relationship and not improb- a SS ably form coenospecies in the Fic. 1. Chart showing the principal sense of Turesson. I have eae areas of Labiatae in the tcated them as sections. The distribution of Salvia subgenus Calosphace in the New World is characteristic of most of the genera of Labiatae which are confined to the New World. The distribution of those genera which are also Eurasian is of a different nature and will be dealt with in a later paper. While it is true that the genus ranges continuously from the Great Lakes to the plains of Argentina it is concentrated in three prin- cipal areas of wide extent and three small subsidary areas which are characterized by endemism. The first mentioned are the highlands of Mexico, particularly the central portion, the north- ern Andes and the Brazilian highlands. The subsidiary areas which are characterized in some degree by endemism are the 1939 EPLING: SALVIA 35 [Vee 7 ISS Oe = ee IB Rela : nae rth = Za Boy eee ie fd Bice aaee abate Zz a x - / q ai , - oe $i = Seas Sper poston / ‘ 50° 120° 90° ‘| (60° g¢ eS Ree o | faa es | ev ~ Y | \ J i (=) a an v —~ = ee ee ee ae ee te ea Fe htt Fic. 3. ae ae showing the distribution of Salvia, subgenus Calosphace, sections Micranthae, Tubiflorae, Purpureae and Rudes 1939 | EPLING: SALVIA 37 =i LP PEE CH an FERC Mid SEGREEE ERLE deal ETE TT TT dededcbai CD Fic. 4. Chart showing distribution of Salvia, subgenus Calosphace, sections Chariantha and Angulatae. highlands of British Guiana, the mountains of Haiti and the mountains of Cuba (including the Isle of Pines). The areas of distribution of the sections are usually restricted and are generally one of three types. The first is illustrated by the sections Chariantha, Flocculosae and Tomentellae. In this case spe- cies of the Mexican highlands find close counterparts in species of the Andes, particularly of Ecuador and Peru. The second is illustrated by the sections Albolanatae, Curtiflorae and Rudes. In this case species of the Mexican highlands find close counterparts in the highlands of Brasil, but not in the Andes. The third case is largely a Caribbean distribution in which the section occurs more or less continuously in the islands of the Caribbean and the isthmus or the northern part of South America, usually at lower elevations. This is illustrated by such sections as Micranthae, Tubiflorae and Purpureae. Only the large section Angulatae is general throughout most of these areas. Not only Salvia subgenus Calosphace but most of the genera of Labiatae which are confined to the New World (in contrast, for example, to those which are circumpolar) conform to this pat- tern; the significance of this fact is unknown. University of California at Los Angeles, April 20, 1938. 38 MADRONO [Vol. 5 NOTES ON THE GENUS ARCTOSTAPHYLOS? A. E. WIrsLANDER AND BeEryt O. SCHREIBER In the mapping of the vegetation types of California and western Nevada, the species of Arctostaphylos, because of their widespread abundance throughout most of the region and be- cause of their great variation, have presented a difficult problem. It has been necessary, in supervising this mapping project dur- ing the past twelve years, to devote much time to the study of this genus in the field. In this study we have had the coopera- tion of the late Dr. Harry S. Yates, botanist of the project, and the active assistance of every field worker on the survey. The senior author has also had invaluable advice and assistance from Dr. W. L. Jepson, of the University of California, not only per- taining to Arctostaphylos but to many other botanical problems concerning the vegetation survey. In differentiating the species of Arctostaphylos the presence or absence of a burl at the base of the stem and the character of the nascent inflorescences have proved most valuable. These char- acters have not been extensively noted or used by previous work- ers in this genus. The presence of a burl is related to the capacity of a species to sprout following fire or cutting. Dr. W. L. Jepson in 1916 pointed out the sprouting character of certain species in contrast to the nonsprouting character of others and suggested the value of this criterion for differentiating species (MaproNo 1: 3-11. 1916). In 1922 he further developed the use of this character as a basis for species differentiation (Maprono 1: 76-77. 1922). The senior author, as a student of Dr. Jepson, had become ac- quainted with this behavior of Arctostaphylos in the Mount Tamal- pais region. When he was assigned to map the vegetation of the state in 1926 and was requested at the same time to determine the sprouting capacity of each species of the chaparral because of its practical bearing on the construction and maintenance of firebreaks, “sprouting” or “nonsprouting”’ became the first point of observation in the field identification of all trees and shrubs encountered. A record of this character was entered on the field 1 Contribution No. 1 from the Vegetation Type Map Herbarium of the Forest Survey Division of the California Forest and Range Experiment Sta- tion. The senior author is in charge of the Forest Survey; the junior author is supervising botanist, Project No. 365—03-3-35, conducted under the auspices of the Works Progress Administration. [ While this article was in press, Dr. W. L. Jepson, to whom a copy of the manuscript was submitted for criticism on November 17, 1938, published two papers, “Embryonic Panicles in Arctostaphylos” (Erythea 8: 97. December 22, 1938) and “Three New Californian Arctostaphyli” (l.c. 8: 99). The use of my name as co-author of A. pilosula Jepson & Wieslander, A. rudis Jepson & Wieslander, and A. silvicola Jepson & Wieslander was unauthorized and with- out my knowledge or consent. A. E. WiesLanpber. | 1939 ] WIESLANDER AND SCHREIBER: ARCTOSTAPHYLOS 39 Fic. 1. Arctostaphylos glandulosa. Development of the burl in a typical sprouting species. label of each herbarium specimen collected. The sprouting char- acter is, in most instances, readily observable but may be ob- secured because of lack of evidence of fire. For this reason field workers were often confused. Either sprout regeneration was lacking because no fire or cutting had taken place since the seed- ling stage or else the layering habit of many of the species was mistaken for sprouting. In an effort to clear up this difficulty we were rewarded by the observation that the burl is not a reaction of the species to mutilation by fire or cutting, as many have understood it to be, but a normal structure which appears early in the life of the seedling as a few nodules, develops steadily with increasing age, and ultimately attains a diameter of a foot or more, even where fires have never occurred. The exterior sur- face of a large, well developed burl is simply an aggregation of latent buds, which sprout as a result of injury by fire or other agencies. Fire may affect the size and shape of the burl, but is not responsible for its existence. The nature and development of the burl in a typical sprouting species is shown in text figure 1. Since, therefore, the presence of the burl can be observed even in very young plants, its value as a taxonomic criterion is greatly enhanced. Nascent inflorescences as shown in text figure 2 develop in most species of Arctostaphylos during the early summer and re- main in this dormant condition until just prior to blooming. In the warm coastal belt of California these structures emerge from the dormant condition soon after the fall rains which initiate growth, but at higher and cooler elevations they remain dormant 40 MADRONO [Vol. 5 until spring. The value of this character in segregating species was first called to our attention in 1931 by Mr. H. A. Jensen, who is associated with us on the type mapping project. On the Hoaglin Quadrangle in Trinity County Mr. Jensen found this character useful in differentiating A. manzanita Parry, A. canescens Eastw., and A. viscida Parry, none of which forms a burl. Al- though previous use has not been recorded in the literature, Dr. Jepson has informed the senior author that he also had noted the value of the nascent inflorescence character of many of the man- zanitas in the Napa region. The usefulness of this character is enhanced by the fact that it is observable over a very long period each year. During subsequent mapping work it has proved an excellent field character for differentiating many of the species. In the following descriptions, special reference is made to the presence or absence of burls and to the nature of the nascent inflorescences. The type collections are filed in the Vegetation Type Map Herbarium, California Forest and Range Experiment Station, located in the University of California Herbarium at Berkeley. Isotypes have been distributed to the United States Forest Ser- vice Herbarium, the United States National Herbarium at Wash- ington, D. C., and the University of California Herbarium. In citing specimens the following symbols represent the various her- baria consulted: VTM, Vegetation Type Map Herbarium; UC, University of California Herbarium; D, Dudley Herbarium, Stan- ford University; CAS, California Academy of Sciences, San Francisco. ARCTOSTAPHYLOS SILVICOLA sp. emend. A. silvicola Jepson and Wieslander, Erythea 8: 101. 1988. Silvery gray shrub, 20-25 dm. high; no burl formation at base; bark smooth, deep dark red; branchlets densely gray- canescent with soft hairs; leaves elliptic to oblong-elliptic, obtuse or apiculate at the apex, rounded-cuneate at the base, densely gray-canescent, or occasionally glabrate, 15-40 mm. long, on petioles 3-8 mm. long; nascent inflorescence (text fig. 1, b) short, 9-13 mm. long, sessile or on peduncles 2—4 mm. long, campanu- late, pendulous or slightly spreading, 1-3 in a cluster, bracts leafy, lanceolate, 4-12 mm. long, the tips slightly recurved, densely canescent, gray green, tinged pink; flowers white, 5-6 mm. long, on glabrous or hairy pedicels; ovary glabrous or rarely hairy; berry globose, slightly depressed at the apex, 5-10 mm., orange brown, glabrous, rarely slightly hairy; nutlets separate. Distribution. Ridges above Mount Hermon and Bonnie Doon, Santa Cruz County, on marine sand deposits, associated with Pinus ponderosa, Arctostaphylos tomentosa, Adenostoma fasciculatum, Ceanothus Ferrisae, Ericameria ericoides, and Lupinus albifrons. Specimens to verify range. Santa Cruz County: Mount Her- mon, A. E. Wieslander 277 (VTM 2467) ; near Bonnie Doon, H. S. 1939 | WIESLANDER AND SCHREIBER: ARCTOSTAPHYLOS Al Yates 5020 (VTM 8019); Big Trees, C. W. Hanks 169 (VTM 8730); Mount Hermon, H. FE. McMinn 1699 (D); Ben Lomond, A.D. E. Elmer 4606 (D, CAS) ; near Tuxido, April 17, 1904, W. R. Dudley (D, CAS); Scott’s Valley, L. R. Abrams 10903 (D); near Eccles, June 27, 1897, W. R. Dudley (D, CAS); above Bonnie Doon, March 29, 1899, W. R. Dudley (D); Glenwood, Feb. 7, 1914, Horace Davis (CAS) ; Mount Hermon Ridge, Beryl Schreiber 2567 (VTM 238756); Felton (Mount Hermon Ridge) C. M. Bel- shaw 1482 (VTM 138639); Mount Hermon, A. EF. Wieslander 701 (VTM 17279). Arctostaphylos silvicola may be distinguished from A. canescens Eastw. by its smaller, elliptic leaves. The nascent inflorescence of A. silvicola is usually pendulous, although occasionally slightly spreading, and the tips of the bracts tend to recurve, whereas in A. canescens they are generally erect or spreading on arcuate peduncles. The ovary and fruits of A. silvicola are glabrous, rarely hairy, while in A. canescens the ovary is densely long white- hairy and the fruits are usually pubescent or sometimes slightly glandular, or glabrate with a glaucous bloom. Arctostaphylos silvicola forms large silvery gray masses in the chaparral and is the dominant member of the vegetation growing under Pinus ponderosa. It also occurs in nearby pure stands as a fire type replacement of Pinus ponderosa in these localities. ARCTOSTAPHYLOS RUDIS sp. emend. A. rudis Jepson and Wies- lander, Erythea 8: 100. 1938. Low shrub, to 15 dm. high, forming burls at the base and fre- quently on the lower branches; bark reddish brown, rough and shreddy; branchlets finely grayzpubescent; leaves elliptic to ob- long, occasionally oval, usually rounded at the base, or occasion- ally truncate, apiculate or obtuse at the apex, glabrous or lightly pubescent especially on the young leaves, bright green, 15-30 mm. long, on pubescent petioles 83-6 mm. long; nascent inflores- cence (text fig. 2,e) short, oblong, 5-6 mm. long, 1—3 at the tips of the branchlets, pendulous, bracts mostly scalelike, ovate to awl-shaped, acute or acuminate, when acuminate the tips re- curved, 2-3 mm. long, reddish or pale green, finely pubescent; flowers 5-6 mm. long, on glabrous pedicels, pinkish; ovary gla- brous; fruit globose, 9-11 mm. wide, glabrous, light brown or reddish brown when mature; nutlets separate. Distribution. Dune sands near Oceano, San Luis Obispo County, south in the vicinity of Lompoc, Santa Barbara County, 150-600 feet elevation, associated with such plants as Arctosta- phylos pechoensis var. viridissima, Ceanothus dentatus var. impressus, Rhamnus californica, Adenostoma fasciculatum, Salvia mellifera, and Ericameria ericoides. Specimens to verify range. Sawn Luis Osispo County: Oceano, N. French 663 (VTM 13592); near Los Berros, A. E. Wieslander 42 MADRONO [Vol. 5 136 (VTM 2528). Santa Barsara County: Corralillos Canyon, A. E. Wieslander 577 (VTM 12520); Surf, G. E. Sindel 179 (VTM 8390); Burton Mesa, H. S. Yates 6534 (VTM 19229); Lompoc Mesa, A. E. Wieslander 235 (VTM 2286); Burton Mesa, H. L. Mason 11770 (UC); Lompoc, March 8, 19387, E. Denys Rowe (UC); Lompoe Moor, Ralph Hoffmann 376 (CAS) ; 4 miles south- east of Lompoc, Rowana S. Ferris 7605 (D); Lompoc Mesa, Beryl Schreiber 2562 (VTM 28749); Lompoc Mesa, A. E. Wieslander 644 (VTM 15478); Point Sal, A. E. Wieslander 8138 (VIM 23744). Arctostaphylos rudis is distinct from all other coastal man- zanitas which form burls. The shaggy fibrous bark separates it from the A. glandulosa Eastw. group and the small, elliptic, gla- brous or glabrate leaves distinguish it from the A. tomentosa Lindl. group. The nascent inflorescences including the pedun- cles are shorter and more slender than in either the A. tomentosa or the A. glandulosa complex, and the fruit is glabrous, polished, considerably larger, and more profuse. Arctostaphylos morroensis sp. nov. Frutex 15-20 dm. altus; caudex etumescens; cortex asper laciniosus porphyreus vel cinereo-fuscus; ramuli pubescentes pilis adpressis, densis brevibus, interdum pilis longis rigidis admixtis ; folia 15-30 mm. longa oblonga vel raro elliptica, plerumque truncata, aliquando basi subcordata vel raro rotunda, apice plerumque apiculato, aliquando obtuso, flavovirentia vel cinereo- viridia, subtus dense tomentosa supra sparse pilosa vel glabra; petioli 2-4 mm. longi; inflorescentia nascens 7-9 mm. longa soli- taria numerosave in axillibus foliorum superiorum aggregata; bracteae foliaceae, lanceolatae minute pubescentes, aliquando pilis rigidis admixtis; pedicelli glabri; corolla alba vel puniceo- tincta 5-7 mm. longa; ovarium dense pubescens; drupa matura diametro 10 mm. globosa vel paullo depressa, aurantiaco-ferru- ginea, sparse pubescens; pyrenae liberae. Shrub 15-20 dm. high; no burl formation at the base; bark rough and shreddy, red brown or gray brown; branchlets ap- pressed-pubescent, and with dense, short, or long bristly hairs, or the latter lacking; leaves oblong to rarely elliptic, 15-30 mm. long, usually truncate but frequently subcordate or rarely rounded at the base, apex usually apiculate but occasionally obtuse, on petioles 2-4 mm. long, yellowish green or gray green, densely tomentose on the under surface, sparingly pubescent to glabrous above; nascent inflorescence (text fig. 2, a) short, campanulate, 10-14 mm. long, 1 to several, clustered in the upper leaves, mostly pendulous, bracts leafy, lanceolate, 7-9 mm. long, and finely pubescent, coarsely bristly or bristles lacking; flowers white to pink tinged, 5-7 mm. long, on glabrous pedicels; ovary densely pubescent; fruit globose, or slightly flattened, orange 1939 ] WIESLANDER AND SCHREIBER: ARCTOSTAPHYLOS 43 brown when ripe, 10 mm. in diameter, lightly pubescent; nutlets separate. Types. Nascent inflorescence: south of Morro Bay (Hazard Canyon), San Luis Obispo County, 200 feet elevation, October 7, 1938, Beryl Schreiber 2554 (VTM 23740). Flowering specimen: 0.4 mile north of Hazard Canyon (south of Morro Bay), 200 feet elevation, January 31, 1936, Ben Bolt 561 (VTM 138800). Fruiting specimen: south of Morro Bay (Hazard Canyon), 250 feet elevation, May 12, 1986, A. E. Wieslander 6359 (VTM 15468). Distribution. Sandy hills south of Morro Bay, 100-400 feet elevation, associated with Adenostoma fasciculatum, Salvia mellifera, Lotus scoparius, and Ericameria ericoides. Specimens to verify range. San Luis Osispo County: Morro Bay, A. E. Wieslander 640 (VTM 15469); Hazard Canyon, A. E. Wieslander 647 (VTM 17235); Los Osos Ranch, N. French 709 (VTM 14085); Osos Creek, B. Bolt 541 (VTM 138783); Valencia Peak, B. Bolt 644 (VTM 14631); Morro sands, Alice Eastwood 14945 (CAS) ; Haynes Ranch, Morro Bay, July, 1912, Mary Page Ingalls (CAS); road to Pecho, Alice Eastwood 15107 (CAS, D); Morro Bay on road to Pecho, February, 1929, Gertrude Sin- sheimer (UC, D). . Arctostaphylos morroensis may be distinguished from Arctosta- phylos tomentosa by the absence of a burl at the base of the shrubs. In all of the A. tomentosa series burls were formed even on the young seedlings, these developing into great massive growths on the mature plants. The nascent inflorescences of A. morroensis are campanulate, while those of A. tomentosa are ob- long and more rigid or spikelike. Characteristically the leafy bracts in A. morroensis are coarsely bristly ciliate but these bristles are occasionally lacking. The bracts of A. tomentosa are usually merely pubescent. The pedicels in A. morroensis are glabrous, whereas in A. tomentosa they are pubescent. Arctostaphylos morroensis is the common shrub in the chapar- ral at the south end of Morro Bay, frequently forming compact masses over extensive areas. It has been reported as far south as Valencia Peak, 2.5 miles south of the type locality (B. Bolt 644 (VTM)). The specific name is derived from the locality where the species is found. Arctostaphylos otayensis sp. nov. Frutex erectus, 10-25 dm. altus; caudex etumescens; cortex laevigatus, atroruber; ramuli breves glandulosi-pilosi; folia 13- 35 mm. longa elliptica vel oblonga, apice acuto apiculatove basi rotunda vel raro truncata utrinque minute pubescentia sub micro- scopio glandulosa raro glabrata cinereo-viridia vel flavovirentia ; inflorescentia nascens breve oblonga 10-20 mm. longa, 3-8, in apicibus ramulorum terminalis, foliis supremis proxissimis sub- tendita, erecta vel alabastris crescentibus paullo arcuato-adcen- dens; bracteae 3-15 mm. longae foliaceae, glanduloso-pilosae ; 44 MADRONO [Vol. 5 pedicelli glanduloso-pilosi; drupa 5-8 mm. diametro globosa pal- lidissimo-ferruginea, nitida glabra, sparse pilosa vel sub micro- scopio glandulosa; pyrenae liberae per occasionem in nucem osseam coalescentes. Erect shrub, 10-25 dm. high; no burl formation at base; bark smooth, dark red; branchlets short glandular-hairy; leaves elliptic to oblong, acute or apiculate at the apex, rounded or rarely truncate at the base, 13-35 mm. long, on petioles 4-7 mm. long, finely pubescent on both surfaces and microscopically glandular, rarely glabrate, gray green or yellowish green; nascent inflorescence (text fig. 2, d) short oblong, 10-20 mm. long, 8-8 in a cluster at the tips of the branches, closely sub- tended by the leaves, erect or slightly curved upwards as the buds develop, bracts leafy, lanceolate, 3-15 mm. long, glandular- hairy, green, tinged red, or yellow green; flowers white, 5-7 mm. long, on glandular—hairy pedicels; ovary glandular—hairy or sparsely glandular-hairy; berry globose, 5-8 mm. wide, very pale brown, shining, glabrous, sparsely hairy, or microscopically glandular; nutlets separate, or coalesced into a solid nut. a i r ty F Ke aD , Fic. 2. Nascent inflorescences in Arctostaphylos: a, A. morroensis; b, A. silvicola; c, A. pilosula; d, A. otayensis; e, A. rudis. Types. Nascent inflorescence: Otay Mountain, San Diego County, 3200 feet elevation, November 8, 1937, H. A. Jensen 437 (VTM 223879). Flowering specimen: Otay Mountain, 3000 feet elevation, January 23, 1934, A. E. Wieslander 350 (VTM 3476). Fruiting specimen: San Ysidro Mountains (on Otay Mountain), 1800 feet elevation, June 11, 1935, A. E. Wieslander 552 (VTM 12510). Distribution. In chaparral on Otay Mountain, San Diego County, 1800-8600 feet elevation, associated with Adenostoma fasciculatum, Arctostaphylos glandulosa, Rhus laurina, Ceanothus cras- sifolius, Chamaebatia foliolosa var. australis, and at the elevation of 1800 feet in the Tecate cypress association associated with Arcto- staphylos bicolor, A. glandulosa, Cercocarpus betuloides, Comaro- staphylis diversifolia, and Chamaebatia foliolosa var. australis. Specimens to verify range. San Dirco County: Otay Moun- tain, A. E. Wieslander 851 (VTM 38277); N. French 279 (VTM 4.936). Piate 4, Buri FORMATION AND EFFECTS OF FIRE ON ArcTrosTapHyLos. Fig. 1. Arctostaphylos rudis showing shreddy bark and burl formation at base. Fig. 2. Arctostaphylos glandulosa (foreground) sprouting from burls after fire; Arctostaphylos otayensis (background) killed by fire. 1939 ] WIESLANDER AND SCHREIBER: ARCTOSTAPHYLOS 45 Arctostaphylos otayensis, although resembling A. glandulosa in many respects, differs from that species in that no burls are formed and the plants are, therefore, killed outright by fire (pl. 4,fig.2). The fruit of A. otayensis is characteristically small and very pale brown in color, shining, microscopically glandular, and usually glabrous but occasionally sparsely hairy. In A. glandu- losa the fruits are red brown in color when ripe and usually very viscid, or glands lacking. The specific name is derived from the type locality, Otay Mountain. ARCTOSTAPHYLOS PILOSULA sp. emend. A. pilosula Jepson and Wieslander, Erythea 8: 101. 1938. Erect shrub, 10-30 dm. high; no burl formation at the base; bark dark red brown, smooth; branchlets pubescent, and with long or short bristly hairs, or these sparse, or occasionally lack- ing; leaves oblong, elliptic to ovate, 15-45 mm. long, on petioles 5-9 mm. long, truncate or rounded at the base, rarely subcordate, occasionally small teeth at the base, apex apiculate, or merely acute, glaucous green or yellow green, (bright green in one form from Sycamore Springs, branches then densely long hairy), glabrous, or sparsely pubescent on both surfaces, young leaves on new twigs densely silvery-tomentose; nascent inflorescence (text fig. 2, c) short, campanulate, 10-14 mm. long, 1-2- rarely 3- clustered at the tips of the branches, mostly pendulous, bracts leafy, 5-14 mm. long, lanceolate, finely pubescent to glabrous, and usually with coarse bristles but these occasionally sparse or lacking, bright green, tinged reddish, or glaucous green; flowers white, tinged pink, 4-7 mm. long, on glabrous or slightly hairy pedicels; ovary usually glabrous (except in one form from near Los Berros) ; berry depressed globose, 8-10 mm. wide, glabrous, orange brown or red brown, usually with blue-black vertical stripes when mature; nutlets separate. Distribution. Chaparral areas of San Luis Obispo County, 200 to 8600 feet elevation, associated with shrubs of Adenostoma fasciculatum, Salvia mellifera, Eriogonum fasciculatum, Lotus sco- parius, and in the La Panza range associated with Pinus Coulteri. Specimens to verify range. San Luis Oxsispo County: Dove, D. I. Axelrod 555 (VTM 17461); Edna, A. E. Wieslander 646 (VIM 17284); Price Canyon, H. C. Lee 427 (VTM 14053); Indian Knob, H. C. Lee 392 (VTM 14019); Sycamore Springs, A. E. Wieslander 580 (VTM 12522); Los Berros Creek, H. C. Lee 425 (VTM 14051); Summit, H.C. Lee 242 (VTM 15247); Porter Ranch, C. M. Belshaw 1690 (VTM 14817); Suey Creek School, H.C. Lee 618 (VTM 15682); Park Ranch, H. C. Lee 578 (VTM 15593); Tar Springs Ranch, H. C. Lee 572 (VTM 15587); New District School, A. D. Gifford 820 (VTM 19640) ; American Can- yon, A. E. Wieslander 261 (VTM 2287); La Panza range, A. E. Wieslander 5388 (VTM 84384) ; La Panza, W. A. Peterson 652 (VTM 19249); La Panza, D. I. Azelrod 588 (VTM 17467); Edna, Beryl AG MADRONO [Vol. 5 Schreiber 2561 (VTM 23747); Pozo, A. D. Gifford 800 (VTM 19625). Arctostaphylos pilosula is one of the dominant species of man- zanita in the chaparral of San Luis Obispo County. There is great variation in color of foliage, the shrubs in different areas varying from glaucous green through gray-green to the bright green form from Sycamore Springs. The amount of pubescence also varies on the shrubs in a given area. The variants are re- tained in the same species on other characters. One peculiarity which field men first observed and used for identifying this species was the bluish-black vertical stripes on the mature fruit. This character has been noted on all fresh fruits, and, in the dried specimens on those fruits which have not become shriveled. ARCTOSTAPHYLOS Parryana Lemmon var. pinetorum (Rollins) comb. nov. A. pinetorum Rollins, Rhodora 39: 462. 1987. Branchlets, petioles, and inflorescence densely glandular- pubescent; nutlets usually separate. Distribution. Mountains of southern California, 5000 to 8000 feet elevation, associated with such species as Pinus Jeffreyt, Pinus Lambertiana, Abies concolor, Quercus chrysolepis, Ceanothus cordulatus, and Castanopsis sempervirens; Uinta Mountains, Utah; Uncompahgre Plateau, Colorado. Specimens to verify range. Los ANagELEs County: Table Mountain, Joseph Ewan 9852 (CAS). San Bernarpino County: Butler Peak, A. E. Wieslander 568 (VTM 12512); Silver Creek, D. I. Axelrod 395 (VIM 9593); Lytle Creek, R. St: John wc? (VTM 1658); Summit Camp Baldy, J. S. Horton (VTM 122638) ; Big Bear, A. Lewis 124 (VTM 1659); Mill Creek, J. S. Horton (VTM 12264); Fawnskin Valley, A. E. Wieslander 569 (VTM 12516); Santa Ana Cafion, S. B. Parish 19291 (UC, D); Bluff Lake, S. B. Parish 3708 (UC); Telegraph Peak, IJ. M. Johnston 1540 (UC, D); Cucamonga Peak, I. M. Johnston 1580 (UC, D); Jenks Lake, P. A. Munz 12745 (UC); Dry Lake, J. Grinnell 2 (UC); Bear Valley, S. B. Parish, June 1886 (UC, D); Mt. San Antonio, LeRoy Abrams 2703 (D). Riversipe County: Tahquitz Valley, H. A. Jensen 443 (VIM 22384). In addition to the above California localities are the following: 31 miles southwest of Whitewater, Uncompahgre Plateau, Mesa County, Colorado, Rollins 1928 (UC); Uinta Mountains, Utah (Rollins l.c. 463). The southern California plant is identical with that described by Rollins from the Uncompahgre Plateau, Colorado. As the Californian plants of this species so closely resemble A. Parryana of southern California in habit and their ranges intermingle it has been found necessary to reduce A. pinetorum to a variety of A. Parryana. The distinguishing character which differentiates the two is the glandulosity of the branchlets, petioles and in- florescence of var. pinetorum, whereas in A. Parryana they are Puate 5. Errecrs OF FIRE ON TWO SPECIES OF ARCTOsTAPHYLOS. Fig. 1. Arctostaphylos patula sprouting from burl after fire. Fig. 2. Arctostaphylos Parryana var. pinetorum killed after fire. 1939 | NOTES AND NEWS AT merely canescent. The nutlets in A. Parryana are most usually coalesced into a solid stone, while in var. pinetorum they are fre- quently separate. A. Parryana usually occurs between 4000 and 6000 feet elevation while var. pinetorum is generally found at higher altitudes although their ranges frequently overlap. Arctostaphylos Parryana var. pinetorum of southern Cali- fornia has long been mistaken for A. patula Greene of the Sierra Nevada. However, A. patula forms a definite burl, whereas 4. Parryana var. pinetorum does not. This has been carefully checked by the field crews of the Type Map Survey and in no instance was a burl found. Layering does occur and might be mistaken for sprouts from a burl. After fire A. Parryana var. pinetorum is killed (pl. 5, fig. 2), while in A. patula regenera- tion takes place by sprouting from the burl (pl. 5, fig. 1). Lack of burl formation in A. Parryana var. pinetorum was observed also by Rollins in the Colorado plants. we C44arsaqnd) aeuisny's ill CsnosgeB) aeurysny'¢ we \ Csnojpewouey Saplouiyssyue's sr Q SOplourysslpue ‘SG o-9 SETTER EN SY 7 4 ' ! 1 60 MADRONO [Vol 5 small spreading capitate glands frequently occur; blades of lowermost deciduous leaves ovate, 1-1.5 cm. long, crenate-ser- rate, borne on petioles .5—2 cm. long, those of the upper gradually diminished, lower leaves subcrenate, the upper oblong-elliptic and entire, subsessile, the median 1.5-3.5 cm. long, 6-12 mm. broad, all pubescent with upwardly curved hairs and usually glandular; flowers axillary, borne mostly above the middle of the plant, pedicels 8 mm. long; lower lip of calyx 4-5 mm. long at flowering, 6 mm. long at maturity, squama then 3.5—4 mm. tall, concave; corolla whitish, its galea and tube 14-21 mm. long, lower lip appressed to laterals; lower stamens seated below middle of tube; nutlets black, verrucose, obscurely banded. In the field this species is readily distinguished by its whitish corolla except in what are apparently rare cases of albinism in other species. However, this is the only single difference I have found which may be advanced to distinguish this species from S. antirrhinoides. In habit and pubescence it is almost identical with that species and in the conformation of its corolla ap- proaches it closely. The nutlets of both are similar, and the range of variation in each is such as to preclude any certain differentiation on these grounds. Measurements of the length and breadth of median leaves (generally the lowermost floral leaves) give almost identical averages, as may be seen from the figure. Only in the conformation of the corolla and particularly the position of the lower stamens with reference to the base of the corolla tube do the averages show a significant hiatus. Even here the extremes of S. antirrhinoides include those of S. califor- nica. ‘The differences show most clearly in the maximum size corollas; these and the averages may be compared in the figure. In respect to corolla conformation S. californica is also very similar to S. Bolanderi. The pubescence as viewed with a lens is usually not glandular although some specimens show a marked sprinkling of slender small capitate glands. As viewed under a compound microscope these glands are usually present in some degree. The area occupied by S. californica is much more restricted than that of S. antirrhinoides. It ranges along the Sierran foot- hills from Tuolumne County northward to Siskiyou County thence southward in the north coast ranges to Alameda County, being found from almost sea-level in the coastal counties to 6500 feet in the Sierra Nevada. It occurs almost wholly within the southern limits of Pseudotsuga tawifolia, both in the Sierran and Coastal forests, growing in seepage spots and on banks above streams or along the streams themselves. The species flowers during June and July. The area occupied may be learned by reference to the map. Following is its distribution by counties: Catirornia. Tuolumne; Calaveras; Amador; El] Dorado; Placer; Nevada; Yuba; Butte; 1939] EPLING: SCUTELLARIAE 61 Plumas; Siskiyou; Humboldt; Tehama; Mendocino; Lake; Sonoma; Napa; Marin; Solano; Alameda. 7. SCUTELLARIA ANTIRRHINOIDES Benth. in Lindl. Bot. Reg. 18: pl. 1493. 1882, based upon a specimen collected by Scouler along the banks of the Columbia River near Ft. Vancouver; the type is in the herbarium of the Royal Botanic Gardens at Kew. Scutellaria viarum Heller in Muhlenbergia 1: 82. 1904, based upon a specimen collected by Heller (no. 5786) along the banks of the Russian River near Windsor, Sonoma County, California ; the actual type is unknown to me. Scutellaria sanhedrensis Heller in Muhlenbergia 1: 31. 1904, based upon a specimen collected by Heller (no. 5894) at Summit Lake on Mount Sanhedrin in Lake County, California; the actual type is unknown to me. Scutellaria nevadensis Eastwood in Bull. Torr. Bot. Club 30: 492. 1908, based upon a specimen collected in Nevada in Elko County in Little Lakes Canyon (Western Stampede) by Beve- ridge (no. 546); the type is in the Herbarium of the California Academy of Sciences, isotypes are in the herbaria of the Univer- sity of California and the New York Botanical Garden and in the Rocky Mountain Herbarium. A perennial herb usually 15-25 cm., rarely 30 cm. tall, creep- ing by rhizomes; stems usually branched at the base, finely pube- scent with upwardly curved hairs, sometimes sprinkled as well with capitate glands; blades of lower deciduous leaves ovate, 1—2 em. long, crenate-serrate, borne on petioles .5-1.5 cm. long, blades of upper leaves gradually diminished, ovate-elliptical, entire, the median 6-15 mm. broad, 15-35 mm. long, pubescent on both surfaces with upwardly curved hairs and sometimes glandular; flowers axillary, mostly disposed above the middle of the plant, borne on pedicels 3-4 mm. long; lower lip of calyx 3—4.5 mm. long at flowering, upper lip 5-6 mm. tall, at maturity, concave; corolla violet-blue, tube and galea 12-22 mm. long, lower lip with two lighter blotches on either side of the middle, closely appressed to the laterals, thus closing the orifice; lower stamens usually seated near the middle of the tube; nutlets black, verrucose, somewhat banded. As pointed out above, Scutellaria antirrhinoides is ee ce able from S. californica predominantly by the position of the sta- mens within the corolla. It is distinguishable from S. angustifolia on similar grounds and also by the conformation of the corolla lips. It differs to a less extent by minutiae of pubescence and habit. The differences of the latter are not so great nor so readily perceived however, and in the absence of flowers these species might readily be confused. But even in size and struc- ture the flowers of both species exhibit a degree of variation sufficient to cause confusion of some specimens and in one region plants commonly occur which are not certainly referable to 62 MADRONO [Vol. 5 either. While S. antirrhinoides approaches S. Austinae and S. angustifolia very closely it is clearly distinct from the immediate allies of those species, S. Brittoni and S. siphocampyloides. The differences in foliage habit may be observed by reference to the chart. The flowers of these species, however, do not have the variability of S. angustifolia and hence the relative position of the lower stamens is always diagnostic. While the habit and pube- scence of S. antirrhinoides are fairly stable the size of the corolla is variable throughout its range, although the proportions are generally fairly constant. There seems to be some correlation between the corolla size and certain geographic areas. While the galea and tube are commonly about 16 millimeters long, those from Nevada average about 18 millimeters and the lowland forms from Humboldt and Del Norte counties average about 20 milli- meters. In conformation the corollas of S. antirrhinoides are inter- mediate with S. angustifolia and its immediate allies on the one hand and S. californica and S. Bolanderi on the other. The extremes lean strongly in both directions. Scutellaria antirrhinoides, the range of which is similar to that of S. nana but more extensive, has the widest distribution of any of the species treated herein, save S. galericulata. Except for the differences in flower size as indicated above, it is fairly stable throughout this range. In coastal Oregon and in California it occurs chiefly below 2000 feet, but in eastern Oregon, Nevada and Idaho it ranges to 5000 feet. The species flowers chiefly in June and July. The area where the greatest variation occurs is in northwestern California and adjacent southwestern Oregon, corresponding in part to Jepson’s Tehaman area (Man. FI. PI. Calif. 12. 1925). The forms in which glandular hairs are abundant and readily perceptible with a hand lens seem to be confined largely to this area and usually occur with eglandular forms. Here also are to be found two anomalous forms, one in which the pubescence approximates that of S. siphocampyloides as it occurs in the coastal ranges of California, the other in which a plant with all the characteristics of S. antirrhinoides is combined with a flower scarcely separable from the smaller-flowered forms of S. Austinae in size, in conformation of the corolla and in the relative position of the lower filaments. A form similar to this intermediate and itself difficult of reference to either species, occurs in the vicinity of Ogden, Utah. Yet S. Austinae and S. antirrhinoides frequently flower side by side with apparently no hybridization. Following is the distribution by counties: Oregon. Washing- ton; Benton; Clackamas; Marion; Wasco; Lane; Curry; Joseph- ine; Jackson; Klamath; Union; Baker; Grant; Harney; Malhuer. IpaHo. Adams; Ada; Owyhee; Blaine; Butte. Nevapa. Hum- boldt; Elko; Pershing; Lander; Eureka. Utan. Uintah Moun- 1939] EPLING: SCUTELLARIAE 63 tains (!). Catirrornia. Del Norte; Humboldt; Mendocino; Lake; Sonoma; Siskiyou; Trinity; Shasta; Modoc; Plumas; Butte; Nevada; Placer; Amador. Following are the localities of perceptibly glandular forms. Orecon. Josephine County: Grants Pass. Douglas County: Comstock. Klamath County: Rock Creek Lodge, west side Upper Klamath Lake. Catirornia. Siskiyou County: Goose- nest foothills, Humbug Creek, Mount Shasta. Trinity County: Coffee Creek at Union Creek, Weaverville. Humboldt County: Trinity River at Willow Creek, Kneeland Prairie, Buck Moun- tain, Thrall, Shasta Valley. Following are the localities of an anomalous form with pubescence similar to that of S. stphocampyloides. Orrcon. Lane County: Spencer Butte. Josephine County: Rogue River near Grants Pass. Jackson County: High Cascade Mountains. Following are the localities of an anomalous form inter- mediate with S. Austinae. OreGon. Harney County: Burns. Grant County: Hamilton. Cattrornia. Siskiyou County: Scott River Valley, Yreka, Cherry Creek, Klamathon, Humbug Creek. Shasta County: near Grisez’s Mill. Uran. Weber County: Ogden Canyon. Morgan County: Peterson Canyon. 8. ScuTELLARIA ANGUSTIFOLIA Pursh, Fl. Am. Sept. 412. 1814, based upon a specimen collected by Lewis and Clark along the Clearwater River near Kamiah, Idaho; the type is in the Acad- emy of Natural Sciences at Philadelphia. Scutellaria veronicaefolia Rydb. in Bull. Torr. Bot. Club 36: 681. 1909, based upon a specimen collected in Nez Perces County, Idaho along Peter Creek by Sandberg, MacDougal and Heller (no. 115); the type is in the herbarium of the New York Botanical Garden. A perennial herb usually 15-30 cm. tall, creeping by slender rhizomes; stems usually branched at the base, generally ap- pressed-hirtellous with ascending hairs, these less often replaced with spreading capitate glands; blades of the lowermost leaves ovate, subcrenate, 1-2 cm. long, borne on petioles .5—3 cm. long, those of the upper gradually diminished, entire, subsessile, oblong-ovate or oblong, appressed-hirtellous with ascending hairs unless capitate-glandular, sometimes, at least the upper surfaces nearly glabrous; flowers axillary, borne on pedicels which are 5-6 mm. long at maturity; lower lip of calyx 4.5-5.5 mm. long at flowering, 5.5—7 mm. long at maturity, the squama then 4-5 mm. tall, concave; corolla deep violet-blue, its galea and tube 20-82 mm. long, arcuate below the middle, lower lip not ap- pressed to the laterals unless at their bases; lower stamens seated somewhat above the middle of the tube; nutlets black, verrucose, sometimes lightly banded. Three species immediately allied to Scutellaria angustifolia: S. Brittonii, S. siphocampyloides and S. Austinae, have usually been re- 64 MADRONO [Vol.5 ferred to that species as varieties. In conformation and size of the flower they are essentially identical. The flowers of S. angustifolia and S. Brittonit are apparently wider in the throat. The averages of measurements show no significant differences. However, they do present recognizable and quite constant dif- ferences in habit. Since this is true and since they occupy areas largely separate and have already been named, I have preferred to maintain them as species. The evidence of their distribution and morphology suggests that they would prove largely inter sterile. Indeed, while the modes of S. antirrhinoides and S. angustifolia are much further apart than the modes of these four species, several forms exist which (as mentioned above) are inter- mediate and dificult to refer to either the first or to the second. Once learned, however, the minutiae of pubescence of these four species as well as the habit, serve to distinguish them. They apparently are not confluent in intermediate areas. The modes of variation of the nutlet patterns are apparently distinct but close, although an insufficient number of mature nutlets is avail- able for a conclusive study. It is not improbable that adequate study of the species in nature would reveal differences more marked than is possible to observe in dried specimens. The range of variation in habit of the four species taken together is considerably greater than in any one of the other species dis- cussed. The differences in habit may be expressed numerically by measurements of the median leaves, correlated with their shapes. Such averages are expressed in Chart 1. Scutellaria angustifolia shows a marked variation in pubescence in which the usual eglandular appressed hairs are replaced by spreading capitate glands. These variants apparently occur in company with the other type and are restricted to an area in the drainage of the Snake River where Idaho, Washington and Ore- gon meet. The species flowers chiefly in May, June and July. Following is the distribution of the species (in the United States by counties) ; in altitude its range lies largely below 2500 feet. British Cotumsia. Ivano. Boundary; Bonner; Kootenai; Latah; Nez Perce; Idaho; Adams; Payette; Boise; Canyon. WasuinctTon. Island; Okanogan; Stevens; Pend Oreille; Spo- kane; Chelan; Klickitat; Whitman; Walla Walla; Asotin. Org- con. Wasco; Polk; Sherman; Umatilla; Union; Wallowa; Baker; Grant; Wheeler; Crook; Deschutes; Lane. The glandu- lar forms occur as follows. IpaHo. Nez Perce County: Peter Creek, Lewiston. Idaho County: Kamiah; Snake River (Willow Creek). Orrecon. Wallowa County: Troy, Deep Creek (Snake River). Wasuineton. Whitman County: Clarkston, Truax. 9. ScuTeLtLaria Britronyu Porter in Bull. Torr. Bot. Club 21: 177. 1894, based upon a specimen collected in Colorado in Clear Creek Cafion by Coulter; the type is in the herbarium of the Academy of Natural Sciences at Philadelphia. 1939] EPLING: SCUTELLARIAE 65 e S.angustifolia (Wn. e S.Austinae Calif) ® S.Brittonii (Rocky Mts) + S. siphocampyloides ® S.? Austinae = S angustifolia (glandular) @ S.Austinae (glabrous) alt > Fic. 2. Map showing distribution of Scutellaria angustifolia and its allies. Scutellaria virgulata Nels. in Bull. Torr. Bot. Club 25: 283. 1898, based upon a specimen collected in Wyoming in the Laramie Hills by Nelson (no. 3218); the type is in the Rocky Mountain Herbarium. | Scutellaria Brittonii var. virgulata Rydb. in Fl. Colo. 296. 1906, based upon the same. A perennial herb generally about 15 cm., rarely 30 cm. tall, spreading by rather thickish rhizomes; stems usually branched at the base, variously pubescent, nearly glabrous, puberulent with retrorse-appressed hairs or with upcurved curled hairs or with either one or the other of these types associated with spreading capitate glands; blades of the lowermost leaves oval, 8-15 mm. long entire, borne on petioles 8-5 mm. long, those of the upper 66 MADRONO [Vol. 5 sessile, ovate-elliptical, entire, gradually diminished upwards, the median 17-35 mm. long, 6-12 mm. broad, entire, frequently somewhat revolute, both surfaces now nearly glabrous, now pubescent and more or less glandular; flowers axillary, borne mostly above the middle of the plant on pedicels 3-4 mm. long; lower lip of calyx 4.5—6.5 mm. long at flowering, 7-8 mm. long at maturity, the squama then 4—4.5 mm. tall, concave; corolla deep violet-blue, its tube and galea 23-32 mm. long, arcuate below the middle, lower lip glabrous, not appressed to the laterals unless at their bases; lower stamens seated somewhat above the middle of the tube; nutlets black, verrucose, banded. From the accompanying diagram it will be seen that the pro- portions of the average median leaves of Scutellaria Brittonii are essentially those of S. angustifolia, although the average dimen- sions are materially different. As a general rule the veins of the leaves are more prominent and more often seven in number as indicated. The texture of the leaves is also different. Although the flowers of S. Brittoni and S. angustifolia are essentially iden- tical in size and proportion, the latter is much more like S. antir- rhinoides in pubescence and habit, than it is like S. Brittoni. The species flowers in May and June. | The species exhibits two pubescence varieties. The first of these is characterized by a clearly defined retrorse pubescence. The second (Scutellaria virgulata) has pubescence which while partly retrorse as seen under a compound lens, nevertheless curls upward to some extent. Both forms may be sprinkled with spreading capitate glands. The two forms are apparently not otherwise distinguishable, and commonly occcur together throughout most of the range of the species. The distribution by counties is as follows. Wyomine. Albany; Laramie. Cotorapo. Larimer; Boulder; Gilpin; Jef- ferson; Denver; Douglas; Clear Creek; Park; Teller; El] Paso; Fremont; Saguache; Huerfano; Garfield; Las Animas. New Mexico. Colfax; Mora; Eddy. 10. ScuTELLARIA SIPHOCAMPYLOIDES Vatke in Bot. Zeit. 30: 717. 1872, based upon a specimen collected in California by Bridges; the type is in the herbarium of the Botanical Institute at Berlin. Scutellaria angustifolia var. canescens Gray in Brewer and Wat- son, Bot. Calif. 1: 603. 1880, based upon three specimens. I consider that collected in the canyon of Pacheco Pass by Brewer (no. 1285) to be the standard. The others were collected by Bolander at the Geysers (no. 3947) and Mount Bullion (no. 4946). A perennial herb generally 20-40 cm. tall, spreading by slender rhizomes; stems usually simple, rarely branching at the base, glandular with spreading capitate glands and pubescent with either ascending or retrorse hairs which are but little curled; 1939] EPLING: SCUTELLARIAE 67 blades of lowermost leaves ovate or oval, subcrenate, 1—2 cm. long, borne on petioles .5—3 cm. long, soon deciduous, those other than the basal prevailingly oblong-elliptical, generally 2-3 cm. long, .5—1 cm. broad, subsessile, obtuse, entire or rarely subser- rate, both surfaces softly pubescent with curled hairs and capi- tate glands; flowers axillary, generally borne above the middle of the plant on pedicels 3-5 mm. long; lower lip of calyx 4.5—5 mm. long at flowering, 5.5—-7 mm. long at maturity, the squama then 4-4.5 mm. tall, concave; corolla deep violet-blue, its tube and galea 24-30 mm. long, arcuate below the middle, lower sta- mens seated somewhat above the middle, lower lip not appressed to the laterals unless at their bases; nutlets black, verrucose, obscurely banded. While indistinguishable from Scutellaria angustifolia in respect to flower structure and proportions, S. siphocampyloides frequently resembles S. californica in habit. This is due to the generally oblong median leaf which is characteristic of both, as well as to a tendency for the lower leaves to be toothed. The species com- prises two well defined and isolated geographic races. The first, which occurs in the foothills of the Sierra Nevada, is distin- guished by the ascendent pubescence, characteristic of most of the group to which this species belongs. The hairs are longer, however, often twice as long as those of S. angustifolia or S. ‘Austinae, and are more spreading. The capitate glands are sel- dom wanting and are well developed. As a result, the pubes- cence is much more noticeable, particularly under a lens, than in any other member of this group. The second form, found in the Coast Ranges south of San Francisco Bay, has essentially the same pubescence but it is clearly retrorse, comparable to that of S. Brittoni but of a different texture. As previously pointed out, an anomalous form of S. antirrhinoides with much the same pubes- cence is known in southwestern Oregon. A form from Tulare County has reduced pubescence scarcely separable from that of S. angustifolia. This species occurs within the oak woodland chiefly below 2000 feet, in rocky soil and in gravelly dry stream bottoms. It flowers chiefly during May, June and July. It is distributed as follows. Catirornia. Plumas County: Prattville, Butt Creek. Tehama County: Mineral. Butte County: Butte Meadows, Chico Meadows. Sutter County: Marysville Buttes. Placer County: Stillwater. Amador County: Ione. Calaveras County: Mercer’s Cave, Reservoir, Angels Camp, Milton, Wallace, North Fork Calaveras River near San Andreas, Harman Peak. Tuolumne County: Sonora, Harden Ranch, Big Oak Flat road, Taylor Hill, Spring Gulch near Bear Creek. Mariposa County: Mount Bul- lion, Greeley Hill above Coulterville. Fresno County: Pine Ridge, Trimmer’s Springs, Big Sandy Creek, Tollhouse, Badger. Tulare County: Homer’s Nose (Sequoia Forest), Middle Tule 68 MADRONO ~ [Vol. 5 River, Paradise Ridge, Kaweah River. Alameda County: Mocho Creek, Cedar Mountain, Oakland Hills, Hayward. Santa Clara County: Coyote Creek, Pacheco Pass, Gilroy, Isabel Creek (Mount Hamilton). San Benito County: Hernandez. 11. Scuretitaria AustTinaE Eastw. in Bull. Torr. Bot. Club 30: 493. 1908, based upon a specimen collected in California in Butte County along Big Chico Creek by Mrs. Bruce (no. 1835) ; the type is in the Herbarium of the California Academy of Sciences; isotypes are in the herbarium of the University of California. Scutellaria linearifolia Eastw. loc. cit., based upon a specimen collected at “San Diego,” California by W. J. Fisher (no. 586) ; the type is in the herbarium of the California Academy of Sciences. A perennial herb generally 10-30 cm. tall, spreading by slen- der rhizomes; stems usually branched at the base either entirely glabrous or puberulent with small ascending curved hairs; blades of lowermost leaves oval or oblong, on petioles about equal in length, soon deciduous, those of all but the basal, oblong-ellipti- cal or linear, obtuse, entire, either wholly glabrous or puberulent with small ascending curved hairs, prevailingly 1.5—3 cm. long, 3-6 mm. wide, subsessile; flowers axillary, disposed above the middle of the plant, borne on pedicels 3-5 mm. long; lower lip EXPLANATION OF CHART 1 Cuart 1. Drawn to scale are figures in which it is sought to indicate the range in habital variation in seven closely related species. As has been pointed out in the text the vegetative parts of one species may readily be confused with those of another in the absence of flowers, or conversely, in the absence of foliage, the flowers of some are difficult to differentiate. The rectangles show for each species indicated the minimum, average and maximum sizes of median leaves. ‘The dotted lines indicate the standard deviation from the mean ex- pressed by the solid lines. Measurements were made upon a varying number of dried specimens. The data involved are as follows: Table 1. Lowermost floral leaf (approximately median) in millimeters g SI S| Bs S| _| a5] 5! 2 | eS es] 2] 2 | 2s N=|3e| ee] 2 | eelx8| ss|] BS ee eta ss LX Si sie wee 3 AE = | a o |a |e < o |e S.- californica :2.2.9... 82 31 | 185) 4 1) Te 7 18 | 4 S. Bolanderi ......... 51 | 42 | 25.5 | 6 10 | 26 13/5) 2 31550 79 S. antirrhinoides ..... 106 | 28 | 17 3.8 10 | 14 6 2.1 2.5 S. Austinae .......-:. 94 | 37 | 22 5.5 12 7.5 Sak) 2 S:) Brittonil 7isc t..o8. 74 | 37 | 24.5 | 5 16 | 15 8 2 5 S. siphocampyloides .. 40 | 44 | 28 6.2 |.15 | 10 5 1s a res S. angustifolia ....... 86 | 32 9) 21 A212 Ld 64) 15 | 3.5 The averages of the last four named are contrasted in figure A. The aver- ages for the three geographical forms of S. Austinae are shown above that name. The average shapes of the leaves of the four last named species are indicated by the outline drawings. 69 SCUTELLARIAE EPLING 1939] "D1.L0]]97 NIG UL UOTPVIIVA Jeo, URIPIUI JO UOT}e}UOSoIda1 DI}eUIMIVASeVIGQ: *[ LAVHD VINOZILSNONY S3GIONINYILNY SS nies S30IOIADNYVSOHdIS VOINYOSJINVS ls ae Sen Kiony MINOLIIE Po SPSabS= SS SSE 55 I¥30NV108 oa 70 MADRONO [Vol. 5 of calyx 4.5 mm. long at flowering, 5-6.5 mm. long at maturity, the squama then 3.5—4.5 mm. tall, concave; corolla deep violet- blue, its tube and galea 23-29 mm. long, arcuate below the mid- dle, lower lip not appressed to the laterals unless at their bases; lower stamens seated somewhat above the middle of the tube; nutlets black, verrucose, obscurely banded. This species is most like Scutellaria angustifolia, differing chiefly in habit. The pubescence, when viewed in a large number of specimens, is seen to be essentially that of S. antirrhinoides but not very different from S. angustifolia. It is not at all glandular or rarely so. The habit of the whole plant may be expressed in the averages of the median leaves as shown in the chart. While essentially of the same shape as those of S. stphocampyloides, they are materially narrower. The species exists in three more or less segregated areas each of which is characterized by a form peculiar to it. The plants of the North Coast Ranges are quite glabrous and the proportions of the median leaves are of a defi- nite order. Somewhat puberulent forms are infrequent. In the northern and central Sierran foothills occurs a second form, the typical (nomenclatorially). It is puberulent with a pubescence very like the eglandular forms of S. antirrhinoides and its median leaves average less in length and in width. In the southern Sierras and especially in the mountains of southern California is a third form (S. linearifolia) with similar pubescence, but with even narrower and shorter leaves; the whole plant is smaller. EXPLANATION OF CHART 2 Cuart 2. Drawn to scale are figures representing the maximum and mini- mum sized corollas of nine species. The conformation of the corollas as shown here is diagrammatic and is not significant. As shown elsewhere, those in the left-hand row are very similar in conformation and are characterized by a closed throat. Those in the right-hand column are likewise similar and are characterized by an open throat. In the case of S. Bolanderi it will be seen that the minimum corolla of subsp. typica is larger than the average of subsp. austromontana. The quadrangles accompanying each are diagrammatic averages of the corolla sizes, indicating length and breadth together with the points of attach- ment of the stamens. In the case of S. Bolanderi figure A represents subsp. austromontana, figure B subsp. typica. In the case of S. Austinae figures C, B and A represent, respectively, the north coast form, the Sierran form, the southern form. It will be observed that while the corolla of the first named form is comparable to those of the three species immediately allied, the average is smaller. This may be due in part to the greater number of southern forms available for measurement. ‘The variation in the width of the throat of these four species may be observed, as well as the variation in place of stamen attach- ment. In the case of S. tuberosa figure A represents subspecies similis, figure C subspecies australis and figure B the intermediate forms. The differences in stamen arrangement between the subspecies is readily perceived. The data were obtained from boiled corollas; they were partly substantiated by field observation. The number of corollas involved in making the diagrams is as follows: S. Austinae, southern form (A) 20, Sierran form (B) 5, north coast form (C) 7, the average of all (D) 32; S. siphocampyloides 15; S. angustifolia 20; S. antirrhinoides 46; S. nana 15; S. tuberosa subsp. similis (A) 18, inter- mediate forms (B) 15, subsp. australis (C) 14; S. Brittoniit 13; S. californica 41; S. Bolanderi subsp. austromontana (A) 7, subsp. typica (B) 7. 1939] BOLANDERI CALIFORNICA ANTIRRHINOIDES EPLING: SCUTELLARIAE AUSTINAE BRITTONII SIPHOCAMPYLOIDES ANGUSTIFOLIA TUBEROSA 71 Cuart 2. Diagrammatic representation of size variation in corollas of Scutellaria. "2 MADRONO [Vol.5 The averages of these forms may be seen by reference to Chart 1. In flower structure and size the species is not materially different from S. angustifolia. The species occurs on dry, usually stony banks or in gravel, frequently along the dry margins of streams and is found chiefly in association with the drier aspects of the yellow pine forest. Its. westward limit corresponds to the western limit of yellow pine in Humboldt County. Although the type was collected along Chico Creek, presumably under oaks, it seldom ranges into the oak woodland. In the Sierra Nevada and North Coast Ranges it seldom occurs above 2500 feet. In southern California it occurs chiefly from 3500-5000 feet. It flowers chiefly during June and July. Following is the distribution by counties: Cattrornia. Hum- boldt; Trinity; Shasta; Tehama; Lake; Napa; Butte; Plumas; Nevada; Yuba; Tulare; Kern; Ventura; Los Angeles; San Ber- nardino; Riverside. University of California at Los Angeles, July 1, 1938. CONE VARIATION IN DIGGER PINE W. PatmMerR STOCKWELL Pinus Sabiniana Dougl., the digger pine, is quite constant in its general appearance, having a forked or loosely branched crown, sparse gray-green foliage and often a leaning posture. The cones are dark, the seeds are large and the seed wings are short and thick. However, the degree of variation in cone size and morphology exhibited by this pine is approached by few others. In central and northern California the cones of digger pine often resemble those of Coulter pine in size and general confor- mation, with hooked spurs as long as two inches from some of the basal scales. Toward the southern end of its range, however, and near the coast, colonies of digger pine are known that pro- duce cones of an entirely different appearance. These cones are short, broad based, massive, woody, carved in appearance, and the scales are tipped with short, heavy down-turned spines. The general aspect of the cone is similar to that of Torrey pine; so striking is this resemblance, in fact, that the botanist may suspect that these two species have been associated in the past, although there is no overlapping of their ranges at the present time. Variation of cone size is as great as variation of cone form in the digger pine. In May, 1938, the writer, accompanied by H. L. Mason, visited a colony of large-coned trees near Bartlett 1939 | STOCKWELL: DIGGER PINE 13 Fic. 1. Size variation in digger pine cones. The large cone is typical of the strain found growing near Bartlett Springs; the small cone represents that which occurs on Figueroa Mountain. Springs, Lake County, California. Many old open cones gathered from the ground were between ten and twelve inches long and one of these measured thirteen inches from the lowest scale to the apex and was seven and three-fourths inches in diameter. This cone, thoroughly dry and empty of seeds, with the peduncle and some of the basal scales missing, weighed two pounds, ten ounces. At the other extreme of size range are old, open cones recently collected at a location reported by Carl Epling on Figueroa Mountain, Santa Barbara County, California. In gen- eral these are short and broad, one of them measuring three and three-fourths inches in length, three and one-half inches in di- ameter at the base, and weighing but four ounces. These ex- amples are not of rare individual cones or of isolated trees, but of local races or genetic strains represented by many trees, a fact which emphasizes their importance to an understanding of species distribution and delineation in the western pines. California Forest and Range Experiment Station, Berkeley, September, 1938. 1h ; MADRONO [Vol. 5 ALICE EASTWOOD’S EIGHTIETH BIRTHDAY ANNIVERSARY Miss Alice Eastwood, for forty-seven years Curator of Botany of the California Academy of Sciences, celebrated on January 19, 1939, her eightieth birthday. To honor her on this occasion more than four hundred botanists and garden enthusiasts met at a luncheon held in the Terrace Ballroom of the Fairmont Hotel, San Francisco. Following the luncheon Dr. F. M. McFarland, president of the Academy, presided at a brief congratulatory ceremony. A resolution by the board of trustees in appreciation of Miss Eastwood’s devoted service to botany as curator in the institution was read by Mr. Norman B. Livermore, president of the board. Miss Eastwood received hundreds of congratulatory letters and telegrams; excerpts from a few of these were read by Dr. Robert C. Miller, Director of the Museum and Steinhart Aquarium of the Academy. Among the valuable gifts received by Miss Eastwood was a collection of the Dennison wild flower paintings presented by Mrs. J. T. Morris and Miss F. P. Morri- son. She received also a book plate, the gift of the California Botanical Club, an organization which she founded in 1891. The plate, designed by Mrs. Alice B. Chittenden, shows redwoods and manzanita in the foreground, and in the distance Mount Ta- malpais, one of Miss Eastwood’s favorite collecting fields. Mr. Allen B. Chickering expressed the regard and esteem of Miss Eastwood’s many devoted friends and admirers and, in apprecia- tion of the work she has accomplished in the crowded quarters of the Academy, offered to contribute the sum of two hundred and fifty dollars as the beginning of a fund for a building to be known as the “Alice Eastwood Herbarium.” Miss Eastwood made ac- knowledgments in a characteristic short speech quoting the verse, “My Birthday,” by Dixie Lee which ends:—“I count my age by friends—not years!” Flowering manzanita (Arctostaphylos manzanita) from the William F. Booth place in Sonoma County was used for table decoration. For many years Miss Eastwood has been especially interested in the native Californian species of this genus and its use on this occasion was especially appropriate. Among organizations represented at the luncheon were: American Fuchsia Society, California Academy of Sciences, Cali- fornia Horticultural Society, California Spring Blossom and Wild Flower Association, San Francisco Garden Club. An attractive biographical folder, sponsored by these organizations, was dis- tributed as a souvenir. The accompaning portrait of Miss Eastwood was taken in January, 1939.—EtTHEL Crum, Department of Botany, University of California, Berkeley, February, 1939. Pruatre 7. Atice Easrwoop. 1939 | HOERL: ARTHROBOTRYUM 75 A NEW SPECIES OF ARTHROBOTRYUM Rutu A. Hoeru A collection of Arthrobotryum by Mr. Harold E. Parks, from Smith River Valley, Del Norte County, California, represents an undescribed species. Specimens of the fungus were sent in April, 1933, to Dr. Lee Bonar of the University of California, Berkeley, who suggested and directed this study. The writer is indebted to Mr. Parks for subsequent collections and field observations, which have shown that the fungus is continuously present in this area. Arthrobotryum spongiosum sp. nov. Mycelium copiosum, compactum massis spongiosis sed densis; hyphae fuscae, profuse ramosae, compactae cellis fere globosis, 10-30 y diametro; synne- mata sparsa, nonnumquam absentia, ferme 1 mm. alta, fusca, composita ex pluribus conidiophoris filiformibus dense compac- tis; capitula globosa, conidia dilute brunnea, cylindrata-ovoidea, 3-septata, leviter stricta ad septas, 32—44 x 10-16 u. Mycelium abundant, composed of spongy but dense masses; hyphae dark, profusely branching, composed of cells almost spherical, 10-30 microns in diameter; synnemata sparse, some- times absent, averaging 1 mm. high, dark brown, composed of Fic. 1. Arthrobotryum spongiosum Hoerl. On bark of Libocedrus decur- rens Torr., X1. 716 MADRONO [Vol. 5 -many closely compacted parallel conidiophores; heads globose, conidia pale brown, cylindric-ovoid, 3-septate, slightly constricted at septae, 32—44 x 10-16 microns. Habitat: on twigs and branches of Libocedrus decurrens Torr., Chamaecyparis Lawsoniana (Murr.) Parl., and Pseudotsuga taxifolia (Poir.) Rehder. Type: on Libocedrus decurrens Torr., above Eighteen Mile Creek, Smith River, Del Norte County, California, April 19, 1933, Harold E. Parks 4422 (U.C. Herb. 596853). Fic. 2. Arthrobotryum spongiosum Hoerl. a, synnema, Xx 50; b, detail of branching hypha, x 430; c, detail of conidiophores and immature conidia, x 430; d, mature conidium, X 430. In most of the described species of Arthrobotryum the super- ficial vegetative mycelium is sparse and the synnemata are numer- ous and gregarious. In this species the opposite situation exists. The outstanding characteristic of the fungus is the thick mat of superficial growth. The mycelium forms compact, black, ir- regular, often more or less confluent masses which are one-half to three-fourths of an inch in depth, and which extend over the branches, giving a general blackened appearance (text fig. 1). Fruiting bodies are scattered and not numerous, and in some col- lections are entirely lacking. Although the synnemata rise from the surface of the mat, they are not easily seen with the naked eye because of their small size and the spongy uneven character of the mycelium. The superficial mycelial mats tend to fall away during the dry season. Conclusive evidence has not been found to indicate that hyphae penetrate into living tissue of the host, and no injury to the tree is apparent. The genus Arthrobotryum has a wide distribution. Species have been described from Italy, Holland, Germany, England, 1939 | COPELAND: JOHN GILL LEMMON 17 India, Ceylon, Borneo, Cuba, Porto Rico, and New Jersey. Arthrobotryum spongiosum approaches A. atrum Berk. & Broome, a European species, in spore size, but other characters widely SE DOTS ONS aT) Dat University of California, Berkeley, March, 1938. A PORTRAIT OF JOHN GILL LEMMON The accompanying photograph of John Gill Lemmon was given to me by Mrs. Stacy Slipner of 3241 Broadway, Sacra- mento, California. Mrs. Slipner, who was born in Sierra County, California, in 1859, was a pupil of Mr. Lemmon’s in the seventies. Her reminiscences regarding the Lemmon family are summarized in the following statements: Frank Lemmon was farming in the Sierra Val- ley during the Civil War. After the war, his brother John, another brother (Judge William Lemmon), and a sister (Mrs. Olson) joined him in California. John had served in the Union Army; Mrs. Olson, also, had served the Union cause by working in hos- pitals. John had been a prisoner of the Confeder- ates [in Andersonville] and had been subjected to severe physical and men- tal strain. He became the school teacher in Sierra Valley. He was a good teacher, popular with chil- dren and parents. Mrs. Lemmon was also a bota- nist; she was a Califor- nian, but not from the mountains. Mrs. Slipner’s reminiscences convey an impression of a gentle- ness which might not be expected in the grimly bearded veteran. The photograph, beneath which is printed the legend “J. G. Lemmon, 4th Mich. Cav.”, bears on the reverse the imprint of a San Francisco photographer. It was evidently made not long after the war, but there is no definite information as to its date. —Hersert F. Copreianp, Sacramento Junior College, Sacramento, California, January, 1939. VA GZ. Lemmon, — : th Atih, Car." 78 MADRONO [Vol.5 REVIEW Phytogeographical Problems of Eastern Canada. By Frere Marie-VicTor1In. Contributions du Laboratoire de Botanique de Université de Montreal, Number 30. American Midland Natu- ralist. Volume XIX, Number 3. Pp. 498-558. The University Press, Notre Dame, Indiana. 1988. This study has arisen from the keen observations of a veteran field botanist, who has spent many years exploring the flora of one of the more interesting regions of North America. As its title implies, the contribution aims chiefly to state certain impor- tant ecological and phytogeographical problems, solutions of which are either not offered or are suggested only as tentative hypotheses. The first problem is that of the occurrence in nature of variations which appear similar to the geneticist’s mutations, and of the existence in such genera as Crataegus and Oenothera of many closely interrelated microspecies. The author concludes that many of these microspecies must have originated since the advent of civilization, but does not discuss the cytogenetic basis for their existence. He considers them important as “traces of discontinuous evolution.”” The principal problem discussed by the author, however, is that of the allogenous elements in the flora of eastern Quebec; those species, either endemic or isolated outliers of characteristically Arctic or Cordilleran species, which persist as relics on the higher mountains, the river gravels, and the exposed shores of the Gaspé Peninsula, the Mingan Islands, and the Island of Anticosti. The author gives careful lists and descriptions of many of these species and their habitats, accom- panied by a fine series of illustrations and maps, which speak eloquently for the thoroughness with which he has studied this flora during a period of more than twenty-five years. As a result of his extensive studies, Victorin has concluded that neither of the two principal hypotheses which have been put forward to account for the presence of these allogenous elements is wholly adequate. He gives careful consideration to the hy- pothesis of Fernald, that they are relics which have persisted in their present localities throughout the Wisconsin glaciation. This hypothesis seems clearly applicable to many of the Gaspé spe- cies, but in Mingania and Anticosti Island, the presence of a lobate ice front must be postulated, with the relic species occur- ring in an ice-free area between the lobes. This view is sup- ported by the localization of the allogenous elements in areas of the south shore of Anticosti that correspond in a certain degree to the portions of the Mingan Islands also occupied by such spe- cies. No geological evidence in favor of this supporting postu- late is presented. Another difficulty with Fernald’s hypothesis is that many of the river gravels of the south side of the Gaspé Peninsula, which is unquestionably glaciated, contain as many allogenous species endemic to these lowlands as do the unglaci- 1939] NOTES AND NEWS 79 ated highlands of the Gaspé. Finally, there is “a number of bi- centric species rather general or frequent on the lower St. Law- rence area whose Cordilleran or Asiatic affinity or identity must be accounted for.” To account for these discrepancies the hy- pothesis of post-glacial migration, recently revived by Wynne- Edwards, is brought up by Victorin. He does not, however, feel that this hypothesis can account for all of the allogenous species, and mentions the interesting fact that many of them, when culti- vated in Montreal, show great vegetative vigor and “‘aggressive- ness.” The thoughtful reader cannot finish this most interesting discussion without agreeing that “the situation seems much too complex for one good simple, schematic and dogmatic explana- tion.”’ One can only hope that the author will continue these studies and will eventually reach a solution satisfactory for the explanation of at least the main features of this fascinating prob- lem.—G. Lepyarp STEBBINS, JR. NOTES AND NEWS New Recorps or ALIEN Piants IN San DiEGo County, CAtLi- FORNIA. Most of the species listed below are here reported from southern California for the first time. I am indebted to Dr. I. M. Johnston, Dr. P. A. Munz, and Mr. J. R. Swallen for some of the determinations. Stations given are in San Diego County, and specimens cited are in the herbarium of the San Diego Natural History Museum. Crencurus EcHINaTus L. La Mesa, October 20, 1937, Gander 4637. Not previously reported from California. PENNIsSETUM RuppEeLu Steud. de atl ee ee ee oe Bg a ee - = ~ eens ee austere a coe Volume I, 1916-1929. . . $5.00 Volume II, 1930-1934 .. 5.00 Volume III, 1935-1936 . 5.00 Volume IV, 1937-1938 . 5.00 Single numbers The subscription price of MADRONO is $2.50 per year. We solicit your pat- ronage, Address all orders to: Dr. David D. Keck, Bus. Mgr. Carnegie Institution of Washington Stanford University, California VOLUME V NUMBER 7 MADRONO @ A WEST AMERICAN JOURNAL OF BOTANY - at | pf, i 4\\ , 4 AN f flo : \ \\ g ? Nee L SG) wep Contents THE PHYLOGENY oF THE ANGIOSPERMS, Herbert F'. Copeland .............. 209 Srupies in WesTeRN Viovets—III, Milo S. Baker ................00.0.0... 218 Dr. SETCHELL AND ALASKA Wittows, Carleton R. Ball ................... 231 A PLEISTOCENE ReEcorpD oF PseupoTsucA MACROCARPA, Herbert L. Mason ... 233 A New Species or Lecipea From Brazit, Albert W.C.T. Herre .......... 235 ADDITIONS TO OUR KNOWLEDGE OF THE FLora oF Mount BAKER, WASHINGTON, Ve MM TECTUSOTVE FMEA Wnt ML HDA meme AU LACT Weal Be PMR ein Tuts, 236 Reviews: Fassett, 4 Manual of Aquatic Plants (Herbert L. Mason); — Ammons, 4 Manual of the Liverworts of West Virginia (Annetta Carter); Jaeger, Desert Wild Flowers (David D. Keck) ............ 237 Nores ann News: Southern Occurrences of Allium crenulatum and Meco- nella oregana (George B. and Ruth P. Rossbach); News ............. 240 Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania July, 1940 MADRONO A WEST AMERICAN JOURNAL OF BOTANY Board of Editors Dr. H. L. Mason, University of California, Berkeley, Chairman. Dr. L. R. Anrams, Stanford University, California. Dr. Lincotn Constance, University of California, Berkeley. Dr. H. F. Coretanp, Sacramento Junior College, Sacramento, California. Dr. A. W. Haupt, University of California at Los Angeles. Secretary, Editorial Board—Eruet Crum Department of Botany, University of California, Berkeley Business Manager—Dr. Davm D. Kecx North Queen Street and McGovern Avenue, Lancaster, Pennsylvania or Carnegie Institution of Washington Stanford University, California Entered as second-class matter October 1, 1935, at the post office at Lancaster, Pa., under the act of March 3, 1879. Established 1916. Published quarterly. Subscription Price $2.50 per year. Volume I, Numbers 1 to 17, complete, $5.00. Volume II, Numbers 1 to 17, complete, $5.00. Volume III, Numbers 1 to 8, complete, $5.00. Single num- bers $0.75. Papers up to 15 or 20 pages are acceptable. Longer contributions may be accepted if the excess costs of printing and illustration are borne by the contributor. Range extensions and similar notes will be published in con- densed form with a suitable title under the general heading “Notes and News.” Articles may be submitted to any member of the editorial board. Manuscripts may be included in the forthcoming issue provided that the contributor pay the cost of the pages added to the issue to accommodate his article. Reprints of any article are furnished at a cost of 4 pages, 50 copies $3.70; 100 copies $4.10; additional 100’s .75¢; 8 pages, 50 copies $5.40; 100 copies $6.00, additional 100’s $1.20. Covers, 50 for $2.50; additional covers at $1.50 per hundred. Reprints should be ordered when page proofs are returned. Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania, for the CALIFORNIA BOTANICAL SOCIETY, INC. President: Dr. Ira L. Wiggins, Stanford University, California. First Vice-President: Dr. G. Ledyard Stebbins, Jr., University of California, Berke- ley. Second Vice-President: Dr. Carl Epling, University of California, Los Angeles. Treasurer: Dr. David D. Keck, Carnegie Institution of Washing- ton, Stanford University, California. Secretary: Dr. Lincoln Constance, 4004 Life Sciences Building, University of California, Berkeley. Annual membership dues of the California Botanincal Society are $2.50, $2.00 of which is for a year’s subscription to Madrofio. Dues should be remitted to the Treasurer. General correspondence and applications for membership should be addressed to the Secretary. 1940] COPELAND: PHYLOGENY OF THE ANGIOSPERMS 209 THE PHYLOGENY OF THE ANGIOSPERMS Hersert F. Corpetann The gist of this paper was presented at a symposium of the Western Society of Naturalists, honoring Dr. Douglas H. Camp- bell, at Monterey, California, December 21, 1939. The title is that of one of Dr. Campbell’s papers (6). I owe it to Dr. Camp- bell to make it clear that some of the opinions stated are not his. The concept of phylogeny is in modern biology intimately bound up with that of natural classification; and the natural classification of the flowering plants is, and has been for some three hundred years, one of the major problems of science. The history of work on this problem may be represented by a phylo- genetic tree, which, by a figure of speech, may be called a phy- logeny of phylogenies (text fig. 1, a) ws ASTE! Es < y ec Wee RENDLE if vy 1925 __ [epics Z nusikurs < - - _ < ne a LES w bs = = a ff : a UMBELILALES jose = pan! eee a) ee a 4 BESSEY “ « ENGLER & 2 US PRANTL e © < $ zs SMPINADALES | CENTIANALES, a BENTHAM - PovenoWiALt 3 & HOOKER i x . i a4 CELASTIRALES 0 oo . vk PASSIFLDRALES | TERS Bap Cenphnes SRICOOe | Seen epee caENCE [EN m 4 9 oO (% ADANSON “ MYRAALES CABrOPHTLLALES ” ou “LINNAEUS ("NATURAL ORDERS") cuumnce nt Loy, Z eanefres § 2 “TOURNEFORT. GALYCINAE Ge aoshues S R 1700 —SOURNEFOR WANA 2 ee oe NUDIFLORAE. < . YL “ RIVINUS MORISON b ec, a CUNG Lae a ALES AUHIN 600 -—- --—-J==-=-—-—--- Fic. 1. Phylogenetic diagrams: a, “phylogeny of phylogenies”; b, phylog- eny of the angiosperms according to Bessey, Bot. Gaz. 24: 177. 1897. The earlier names in this history are now merely of historical interest. It may be worth while, as a matter of historical accu- racy and justice, to justify the position of Linnaeus in the main line of development of the natural system. It is generally known that he appended to his Genera Plantarum (21) a list of fifty- eight natural orders, not described, but identified by lists of in- cluded genera. It is noteworthy that in this list he recognized Maprono, Vol. 5, pp. 209-240. July 10, 1940. jy 1 OO 210 MADRONO [Vol.5 the distinction, discovered by Ray, between monocots and dicots; though he does not use the names of these groups. In effect, he did what we do in any modern manual: he prefixed an acknowl- edgedly artificial key to an attemptedly natural arrangement of the genera. It is not he, but the smaller men who followed him, who may be said to have delayed the development of the natural system by an uncritical acceptance of his artificial system. It is further to be noted that Adanson (1), to whom a place in the history of natural classification is generally conceded, set up a series of families exactly as many as the natural orders of Lin- naeus, many of them being the same groups. Botanists of the early nineteenth century. referred to the “natural orders of Jussieu” as if to imply that Jussieu had founded natural orders, or even the natural system. More justly, we would attribute to Jussieu the popularization of the idea of natural orders. He made the recognition of natural orders a practical convenience by the establishment of a system of named higher groups to include them. The skeleton of his system is as follows (19): I. Acotyledones, i. e., seedless plants: Class I. II. Monocotyledones Stamina hypogyna: Class II. Stamina perigyna: Class III. Stamina epigyna: Class IV. III. Dicotyledones Apetalae Stamina epigyna: Class V. Stamina perigyna: Class VI. Stamina hypogyna: Class VII. Monopetalae Stamina hypogyna: Class VIII. Stamina perigyna: Class IX. Stamina epigyna, antheris connatis: Class X. Stamina epigyna, antheris liberis: Class XI. Polypetalae Stamina epigyna: Class XII. Stamina hypogyna: Class XIII. Stamina perigyna: Class XIV. Diclines irregulares: Class XV; orders Euphorbiae, Cucurbitaceae, Urticeae, Amentaceae, and Coniferae. It was not arbitrarily, but according to precedent, that Jus- sieu treated the category of classes as of rather low rank, and that he designated the classes by number rather than by name. The varying sequence of the hypogynous, perigynous, and epigy- nous subdivisions was evidently intended to provide transitions linking together the series of main groups respectively of mono- cots, apetalae, monopetalae, and polypetalae. These groups as 1940] COPELAND: PHYLOGENY OF THE ANGIOSPERMS 211 main groups of angiosperms, and their subdivision primarily by hypogyny, perigyny, or epigyny, have to a considerable extent remained in use through the hundred and fifty years to the pres- ent time. The sequence in which these groups are arranged has, however, been changed repeatedly. During most of the nineteenth century, the ideas of the de Can- dolles were dominant. The original Candollean arrangement need not be presented... The fundamental difference between gymnosperms and proper dicots having been pointed out by Robert Brown, the following modification of the Candollean sys- tem was put forward by Asa Gray (15): Series I. Phaenogamous or flowering plants. Class I. Dicotyledons. Subclass I. Angiosperms. Div. 1. Polypetalous. Div. 2. Gamopetalous (Monopetalous). Div. 3. Apetalous. Subclass II. Gymnosperms. Class II. Monocotyledons. Series II. Cryptogamous or flowerless plants. The sequence just quoted was followed by Bentham and Hooker (3) in what has turned out to be the ultimate elaboration of the Candollean system. In the mean time the classification of angiosperms had under- gone in Germany an independent development. This culminated in the system of Engler and Prantl (10), summarized as follows: Seedless plants. Gymnosperms. Monocots. Apetalae. Choripetalae (i. e., Polypetalae). Sympetalae (i. e., Monopetalae or Gamopetalae). This system was explicitly an attempt to represent phylogeny. Engler was highly conscious of the prevalence of parallel evolu- tion. He supposed that various groups of Apetalae and Chori- petalae, as well as the whole group of monocots, had originated independently of one another from a hypothetical extinct group of gymnosperms; and that various groups of sympetalae had had an independent origin among the Choripetalae. The Apetalae, Choripetalae, and Sympetalae, then, are to be regarded not as natural groups but as evolutionary levels. The system was pre- sented to the world in an extensive work, useful in the recogni- tion and placing of all the genera, written in a modern language, profusely illustrated, and supported by the prestige of the Ger- -man science of the time. It was generally accepted as the true system; most herbaria and most manuals follow it. 212 MADRONO [Vol. 5 The status of the Englerian system as the accepted system has for half a century made it a prime object of attack. The principle of the representation of phylogeny is universally ac- cepted; but whereas Engler used this principle to justify the resurrection of features of the system of Jussieu, those who have refused to follow him have tended to use it to justify features of the Candollean system. Three systems which exhibit this ten- dency are now to be discussed. 1. Bessey (4) put forward a system represented by the phylo- genetic tree here reproduced (text fig. 1, b). He assumes that the whole group of angiosperms, and its two main subdivisions, monocots and dicots, are natural groups; that among dicots the Choripetalae, and particularly the order Ranales, are primitive and a natural group; but that Apetalae and Sympetalae are not natural groups, each including more than one line of descent from Choripetalae. He drops all three as taxonomic groups: he ar- ranges the orders of dicots in two series, distinguished by hy- pogyny on the one hand as contrasted with perigyny or epigyny on the other. In effect, he makes a primary division by the char- acter which Jussieu had used in making a secondary division. It is an obvious criticism, that perigyny and epigyny cannot be assumed to be the marks of a single evolutionary line: surely, these characters have appeared repeatedly, just as have a/jetaly and sympetaly. The Besseyan system has never commanded wide acceptance, but it must be regarded as a living system, hav- ing been followed in recent works by Clements and Clements (8), Pool (23), and Swingle (25). 2. Hallier (16), like Bessey, regarded Angiospermae, Mono- cotyledoneae, and Dicotyledoneae as natural groups, and Ranales as primitive; but his system does not resemble that of Bessey in detail. He supposed Sterculiaceae to be an important secondary center of variation derived from Ranales. He was followed by Lotsy (22) in a work which was never completed. He has had few other followers; he wasted his energies to an unseemly extent in railing against “Engler und seine geistigen Kinder und Enkel” (17). 3. Hutchinson (18), like Bessey and Hallier, postulates the naturalness of Angiospermae, Monocotyledoneae, and Dicotyle- doneae, and the primitiveness of Ranales. The distinguishing feature of his system is the emphasis placed upon growth form, that is, on the woody as contrasted with the herbaceous charac- ter. Woodiness is assumed to be primitive, and the herbaceous dicots are for the most part arranged in a single derived line of descent. This arrangement is open to essentially the same criticism as that of Bessey, in that it assumes a certain evolu- tionary change to have taken place only once, when it may well have taken place many times. I do not know that this system has been followed in any manuals or herbaria; but it is a living 1940] COPELAND: PHYLOGENY OF THE ANGIOSPERMS 213 URTICACEAE ae Dipterocarpacess Quiinacess Guttifere EUCOMMIACEAE Bixacerr Cochlospermaceae Caryocanacese ALYCANTH Manc FAGACEAE barat cage MERE gpaviacese ROSACEAE Oehrkces Wm Flacourtiaceae Theaceze CASUARINACEAE SN ss Eucryphiacess x Actinidiacege BETULACEAE ‘ ‘ ESCALLONIACEAE : Se Ue! PLATANACEAE . \ Dilleniaceze BRUNELLIACEAE (Myristicaceze) ¢?) Magnoliacece GROSSULARIACEAE MY ROTHAMNACEAE \ won HY ORANGE ACEAE \ MAGNOLIALES Ranales (Some arboreal complex) —k. Campanulatae =~ Ebenales Cucurbitales Begoniale Geraniales fapindales eal Geonnasy* ( Umbelliflorae Gricoccae) a ns Rosales Frimulales) Nudiflorae (sens. lat.) i —plllelebiae) Felygonales ere l ae (inel. Burmana) ees) Lerticillatae) j Fic. 2. Phylogenetic diagrams: a, phylogeny of Urticales and related groups according to Tippo; b, phylogeny of Theineae according to Vestal; ¢, the orders of angiosperms (as limited by various authorities) arranged accord- ing to Mez. system in the sense that it demands consideration whenever the relationships of any family are under discussion. The history just sketched is that of a persistent effort to divide the dicots into a small number of easily distinguishable natural groups. The effort has been a failure; one or more of the subdivisions established by each system have turned out to be products of parallel evolution. The characters used have been matters of gross morphology. The. leaders of botany, the framers of systems, have not been unaware of the necessity of 214 MADRONO [Vol.5 considering all characters. They have used gross characters be- cause these were known, being easily recognizable in dried speci- mens and recorded for the whole range of higher plants. When the Englerian system appeared, no phase of the anatomical, chemical, or physiological branches of botany afforded a body of data adequate for taxonomic application. More recently, sev- eral such bodies of data have accumulated to the extent that their significance can be estimated; four of these are now to be dis- cussed. 1. The leader in the study of the vascular anatomy of the re- ceptacle has been Eames (9), and the center of study has been Cornell. It has been found that in Magnoliaceae and Ranuncu- laceae the vascular supply of the floral parts is essentially that of vegetative leaves. Fusion or suppression of floral leaves is ac- companied—or preceded or followed—by fusion or suppression in the vascular system: the cauline nature of the receptacle, and the foliar nature of the other parts of the flower, may become unrecognizable. It is evident from this body of data that the Ranales are primitive and that there has been much parallel evolution. 2. The main outline of the embryogeny of angiosperms was worked out by Hofmeister. Early comparative studies gave the impression that the group is quite uniform. When, therefore, Treub (28) discovered chalazogamy and other abnormalities in Casuarina, he concluded that this genus should be placed in a class by itself, distinct from both monocots and dicots. Again, when Campbell (5) discovered in Peperomia an embryo sac which does not at all conform to the normal type, he concluded that Pipera- ceae is an ancient group, of distinct origin from the angiosperms with normal embryo sacs. In short, the period when the em- bryogeny of angiosperms was supposed to be quite uniform was followed by one in which it was supposed to be so varied as to indicate multiple origin. Some forty more years have passed; Chicago and Vienna have been centers of embryological study, and Schnarf (24) has prepared a useful compendium of the re- sults up to 1980. It is found that the type of embryo sae which includes eight nuclei derived from one which is produced by re- duction division is indeed the normal type. It occurs in the over- whelming majority of the families; it extends to groups as far apart as Ranales, Campanulatae, and Glumiflorae. From the normal type there are parallel deviations which result, for example, in the same derived type in families as far apart as Liliaceae and Plumbaginaceae. In the order Myrtiflorae, most of the families exhibit the normal type; Onagraceae has a peculiar type of its own; Penaeaceae and the genus Gunnera have developed the same peculiarities as Peperomia. Characters of the embryo sac, then, decidedly make it probable that the angio- sperms are a natural group. The normal type embryo sac is primitive, and deviations from it indicate derivation; but it sur- 1940] COPELAND: PHYLOGENY OF THE ANGIOSPERMS 215 vives in groups otherwise highly advanced. In the several other embryological characters, various features are marks of natural groups: thus a developing endosperm which consists of four cells in a row marks the order Ericales. , 3. The comparative anatomy of woods has been studied not- ably at Harvard. There has been developed a doctrine of the evolution of vessels (11, 12, 18); of wood rays (20); and of wood parenchyma. The anatomy of wood is held not in itself to indicate natural groups, but only degree of evolution; it is used to test hypotheses that this group is derived from that, and yields as answer either “it is possible” or “it is impossible.” The effect has been to demonstrate the parallel evolution of many lines of woody plants from a common source in or near Magnoliaceae. Herbaceous plants are interpreted as having originated re- peatedly in primitively woody groups. Some of the detailed re- sults may be presented. Tippo (27) finds it probable that Faga- ceae and Betulaceae, Urticales, and Casuarina are closely related (text fig. 2, a) all being descended through Hamamelidaceae from Magnoliaceae. Juglandaceae and Aristolochiaceae, which fall near these families in the Englerian system, are found to have nothing to do with them. Vestal (29) has worked out the phylogeny (text fig. 2, b) of the group called Theineae in the Englerian system. He finds it a primitive group, connected di- rectly with Magnoliaceae. The group which Hutchinson calls Bixales is not closely allied with it. This group Theineae or Theales or Guttiferales is of particular interest to me because I believe that both Ericales and Ebenales are derived from it— Ericales from Saurauiaceae, Ebenales from Theaceae. 4. Tests for immunity—‘‘serum diagnosis” or “immune reac- tions’’—were first used for the identification for certain diseases and the bacteria which cause them. An animal, being rendered immune by vaccination to a certain organism, reacts in various ways, by agglutination, precipitation, or anaphylaxis, when ex- posed to protein of the species to which it is immune. It shows the same reactions in lower degree to proteins of related species. These immune reactions are characteristic of proteins in general, not only of those of bacteria. The study of reactions to plant proteins has been carried out chiefly by Mez, of Konigsberg. The work has been reviewed in English by Chester (7). In 1926 it had been carried so far as to yield a complete phylogenetic tree. According to Chester, the pictorial representation pub-— lished at that time is protected by copyright and cannot be repro- duced; it is available in the work of Gortner (14). I have con- structed from the original tree of families a tree of orders (text fig. 2, c). Some of the features of this tree are surprising: our authorities on wood anatomy would not allow the derivation of Amentiferae from Centrospermae, nor the inclusion of Juglan- daceae in Amentiferae, nor the derivation of Bixales from Theales; and for my own part I am not content with the position 216 MADRONO [Vol.5 of Ericales. Theoretically the method is sound; I would con- clude that it has not been adequately calibrated, that the degree to which conclusions can be drawn from the direct results has not been established. With an engine of the power of this one at our disposal, we are not wise in failing to use it when hy- potheses as to relationship are to be tested. It is to be remarked that the use of the method is arduous and exacting, and quite outside of the range of technique in which botanical taxonomists are trained. Apologies may here be offered, in that several other bodies of pertinent data—ecology, chemical characters aside from immune reactions, details of the structure of pollen grains, cytology and genetics—are not discussed. I am not aware that a science of _ systematic physiology has so much as been conceived; but physio- logical data are susceptible of systematic treatment. In the long run, these thing will have to be taken into account. Meanwhile, the four bodies of data discussed are conspicu- ously in agreement with each other and with the views of Bessey, Hallier, and Hutchinson, in making the Angiospermae a natural group and the Ranales primitive. These points should, I think, be accepted as positively established. If the Ranales are primi- tive, the angiosperms are not descended from the specialized group of Gnetineae; nor from the cycadeoids (Wieland, 30), in which the carpels are reduced to stalks bearing solitary ovules; nor from the Caytoniales (Thomas, 26), in which the ovules are enclosed by the incurving ends of blades. No known plant, liv- ing or fossil, has the sort of carpel we require of the ancestors of the angiosperms, except only the genus Cycas; and in features other than the carpels, Cycas is not a good match for the hypo- thetical progenitor of the group. We are forced to postulate as such some extinct group of Cycadineae. This is the conclusion reached long ago by Arber and Parkin (2). From the Ranales the other angiosperms are derived, either directly or through secondary centers of variation, one of which appears to be Theales. They fall into many lines of descent. One such line, derived directly from Ranales, is the whole group of monocots; these are bound together not only by monocotyle- dony but by the whole range of their characters. Others are in general yet to be worked out. It is not probable that any of them will be found at the same time so extensive and so well marked as the monocots. Some or many of them may be defin- able by definite characters, as is the group of monocots. On the other hand, in view of the prevalence of parallel evolution, we should accustom ourselves to the probable necessity of accepting named taxonomic groups like the natural orders of Linnaeus, de- finable only by the list of groups included. Already it seems probable that the line of apetalous trees culminating in Casuarina should be accepted as constituting the order Amentiferae: but it is not easy to frame a list of characters by which it will include 1940] COPELAND: PHYLOGENY OF THE ANGIOSPERMS 217 Hamamelidaceae and Platanaceae while excluding Salicaceae and Juglandaceae. When a group is assigned to its true place, it becomes inex- tricable: every character studied increases the certainty of the assignment. Many current hypotheses as to location show signs of becoming stronger; many families are letting it be known that their true place has been discovered. Not merely within the life- time of contemporary taxonomists, but within a decade or two, there should be few families left to be placed by guess, as being too isolated by evolution, or too poorly known, for definite loca- tion. Demonstration of the true phylogeny of all or of nearly all angiosperms will result automatically in revision of the taxo- nomic system; but it will not determine the names, nor the pre- cise limits, nor in all cases the sequence, of the groups which are to be accepted. One is tempted to discuss features which may make one taxonomic arrangement preferable to another: but they are matters of taste or of mere accident. We may expect pres- ently the establishment of a more sound and more stable taxo- nomic system than we have ever had, being a solution, as to the main outlines, of the long-standing problem of the classification of flowering plants. Sacramento Junior College, Sacramento, California, February 3, 1940. LIvERATURE CITED Apanson, M. Familles des plantes. 2 vols. Paris, 1763. Arper, E. A. N., and J. Parxin. On the origin of the angiosperms. Journ. Linn. Soc. Bot. 38: 29-80. 1907. Bentuam, G., and J. D. Hooker. Genera plantarum. 3.vols. London, 1862-1883. Z Bessey, C. EK. Phylogeny and taxonomy of the angiosperms. Bot. Gaz. 24: 145-178. 1897. 5. CampspetL, D. H. Die Entwicklung des Embryosackes von Peperomia pel- lucida Kunth. Ber. d. deutschen Bot. Ges. 17: 452-456. 1899. 6. ————————_.. The phylogeny of the angiosperms. Bull. Torrey Bot. Club 55: 479-497. 1929. 7. Cuester, K. S. A critique of plant serology. Quart. Rev. Biol. 17: 19- 46; 165-190; 294-321. 1937. 8. CLements, F. E. and E. S. Flower families and ancestors. New York, 1928. 9. Eames, A. J. The role of flower anatomy in the determination of angio- sperm phylogeny. Proc. Congr. Pl. Sci. Ithaca 1: 423-427. 1929. 10. Enoier, A., and K. Prantrit. Die Natiirlichen Pflanzenfamilien. 20 vols. Leipzig, 1887-1909. 11. Frost, F. H. Specialization in secondary xylem of dicotyledons I. Origin of vessel. Bot. Gaz. 89: 67-94. 1930. Sis A 12, ———__—_____ ——_——————-_ J. Evolution of end wall of vessel segment. Bot. Gaz. 90: 198-212. 1930. 13... ————_._ —————_——__ III. Specialization of lateral wall of vessel segment. Bot. Gaz. 91: 88-96. 1931. 14. Gortner, R. A. Outlines of biochemistry. ... New York, 1929. 15. Gray, A. Gray’s botanical text book. vol I. Structural botany... . sixth edition. New York, 1907. 218 MADRONO [Vol. 5 16. Hatimr, H. Vorliufiger Entwurf des natiirlichen (phylogenetischen) Systems der Bliithenpflanzen. Bull. Herb. Boissier ser. 2, vol. 3: 306— 317. 1903. 17, ————————.. Uber die Lennoeen.... Bot. Centralbl. Beih. 402: 1-19. 1923. 18. Hurcuinson, J. The families of flowering plants. I. Dicotyledons. Lon- don, 1926. 19. pe Jussieu, A. L. Genera plantarum secundum ordines naturales disposita. Paris, 1789. 20. Kriss, D. A. Salient lines of structural specialization in the wood rays of dicotyledons. Bot. Gaz. 96: 547-557. 1935. 21. Linnarvus, C. Genera plantarum. ... sixth edition. Stockholm, 1764. 22. Lorsy, J. P. Vortriége iiber botanische Stammesgeschichte. vol. 31. Jena, 1911. 23. Poort, R. J. Flowers and flowering plants. ... New York, 1929. 24. ScunarF, K. Vergleichende Embryologie der Angiospermen. Berlin, 1931. 25. Swineite, D. B. A textbook of systematic botany. New York, 1934. 26. Tuomas, H. H. The Caytoniales, a new group of angiospermous plants from the Jurassic rocks of Yorkshire. Phil. Trans. Roy. Soc. B 213: 299-364. 1925. 27. Trero, O. Comparative anatomy of the Moraceae and their presumed allies. Bot. Gaz. 100: 1-99. 1938. 28. Treus, M. Sur les Casuarinees et leur place dans la systeme naturel. Ann. Jard. Bot. Buitenzorg 10: 145-231. 1891. 29. Vesrat, P. A. The significance of comparative anatomy in establishing the relationship of the Hypericaceae to the Guttiferae and their allies. Philippine Journ. Sci. 64: 199-256. 1938. 30. WieLtann, G. R. American fossil cycads. Carnegie Inst. Publ. 34. 1906. STUDIES IN WESTERN VIOLETS—III Mito S. Baker The investigation of certain problems concerning the violets of the Pacific Northwest called for field study; to this end during the month of June, 1988 and 1939, trips were taken through western Oregon, Washington and British Columbia to visit the localities in which the problems centered. Viota oceLiata Torr. & Gray. Through correspondence with Mrs. Cora Ollivant I had learned that Viola ocellata had been col- lected in the vicinity of Looking Glass Post Office, a few miles southwest of Roseburg, Oregon, on the ranch of Thomas Ollivant. This is a matter of interest to students of Viola since this species was not known with certainty to grow in Oregon, its most northern known limits being northern Humboldt County and central Shasta County in California. In Gray’s “Synoptical Flora,” published in 1897, it was stated that this violet had been collected in the Cow Creek Mountains of Oregon, but diligent inquiry on my part had until now failed to confirm this report. On a hillside at a distance of less than a mile from the ranch house there was a colony of this violet. Although not covering a wide area here, the colony was flourishing and the individuals appeared very similar to typical California plants. Both transplants and herba- 1940] BAKER: WESTERN VIOLETS 219 rium specimens (June 19, 1988, Baker 9033) were secured. The Ollivants assured me that they had collected this species also on Sugar Pine Mountain about five miles west by air line from their ranch. They had also noted its occurrence on another mountain near there. Thus it appears that Viola ocellata is well established in that region. This occurrence suggests that sometime in the remote past this violet had a continuous distribution southward through Oregon and northern California to its present range in Shasta and Humboldt counties but that, for some reason, pre- sumably environmental, it has now disappeared from a strip several hundred miles wide in southern Oregon and northern California. VioLta LANCEoLaTA L. Several years ago, Dr. George Neville Jones collected a very diminutive white marsh violet near Park- land, Washington (Jones 8803). A note was published by him in Madrofio (4: 85. 19388) referring it to V. lanceolata L., a species known hitherto only in the eastern and central United States. My first thought was that he must be mistaken as to its identity. However, after examining Jones’ material, I was convinced of the correctness of the identification. A search of the Parkland region proved unavailing. Dr. Jones, however, had collected another specimen of larger stature near Sea View, Washington (Jones 2115). The next day I went to the mouth of the Columbia River in search of Viola lanceolata along the marshes back of the town of Sea View. Here in a pasture about one mile north of town I found thousands of plants in good flower. The owner of the pasture came along at noontime and found a strange man digging up his land! However, after explanations he was sympathetic and bade me take all of this worthless weed that I wanted. It appears that Viola lanceolata is a pest in the cranberry bogs of this region and yet only one botanist was aware of its existence on the Pacific Coast. The species occurs in at least three spots near Long Beach and Sea View, Pacific County, Washington (June 25, 1938, Baker 9055), and I have no doubt that it can be re-located in the region about Tacoma and Parkland if searched for at the proper season. Indeed, Dr. Jones wrote me that he had collected it at several stations east and south of Tacoma. A trustworthy explanation of this strange quirk in the distri- bution of Viola lanceolata is not yet forthcoming. To the east- ward the species is unknown except in the region of the Great Lakes and farther east and south. Since it is a low altitude marsh plant it could not have migrated across the Rocky Mountains; yet the western plant is typical in all respects except in the small size of its seeds. In the absence of any verified explanation I shall venture a guess as to the origin of these plants, although, frankly, I have little evidence to support this speculation. It is known that Viola lanceolata grows in Venezuela. It is also known that 220 MADRONO [Vol. 5 we have along our coast line a number of coastal species of South America. At the present writing, I have in mind Fragaria chilensis L., Aster chilensis Nees, Madia sativa Molina, Lupinus densiflorus Benth., all of which are fairly common along the Sonoma County, California, coastline. One explanation of the presence of these Chilean species along our own coast is that the seeds were brought here by migratory water fowl. It is common knowledge that thousands of individuals of water fowl including many different species, annually fly northward along our coast, and it is reason- able to suppose that small seeds, particularly of marsh plants, could be easily carried in the mud adhering to the feet of these migratory birds. If the mud or clay were sufficiently adhesive, it seems to me entirely credible that such seeds could be carried even from South America and deposited in the marshes as far north as the Columbia River and Tacoma. Viola superba sp. nov. Radices in plantis maturis, adventi- tiae; herba glabra; caules erecti adscendentes, 25 mm. longi; caulium et rhizomatis internodia longitudine variabiles, 1-9.5 em. longa; foliorum lamina conspicue venata rotunda ovato-cordata, 3.5-4.8 cm. lata, 3-5.2 cm. longa, regulariter serrato-crenata, serrae utrimque 12 ad 14, clare apiculata, sulco in foliis maturis circa 13 mm. alto; foliorum maturorum petioli crassi, .5—22 cm. longi; pedunculi crassi, 5-16.5 cm. longi; bracteolae ovatae lan- ceolataeve adpressae, 2-3 mm. longae, supra medium pedunculi insertae; sepalum inferiorum ovatum, alteri elliptici obtusi scari- osomarginati, 5-6 mm. longi; corolla purpurea ei V. Langsdor ffi subaequalis; flori sicci 2.5 cm. diametro; petala superiora laminis suborbicularibus, lateralia leviter barbata, inferiora nuda, apice paullo retuso, 1.4 cm. lata, 2 em. longa, calcare incluso; calcar conicum eburneum, 2 mm. diametro, 2 mm. longum; stylus cras- sus; foramen circa 5 mm. diametro; capsulae oblongo-ellipticae, 12 mm. longae; semina atro-fulva, 1.5 mm. lata, 2.8 mm. longa, pondere 2.3 mg., caruncula .57 longa seminum haud excedente. Roots adventitious in mature plants as in Viola simulata and V. Langsdorffii but fewer, larger and more woody; whole plant en- tirely glabrous; stems erect or ascending, 2—25 cm. long; inter- nodes variable apparently according to season, 1—9.5 cm. long; root-stocks or buried stems of former years with the same variable length of internodes; leaves conspicuously veined, rounded, ovate- cordate, distinctly apiculate, sulcus deep, averaging 13 mm. in mature leaves, evenly serrate-crenate, 12 to 14 serrations on each side of leaf, width and length approximately equal, 3.5—4.8 cm. wide, 8—-5.2 cm. long; petioles of mature leaves stout, 5-22 cm. long; stipules ovate to lanceolate, entire, becoming scarious, 7-10 mm. long; peduncles stout, 5-16.5 cm. long; bractlets slightly above the middle of peduncle, ovate to lanceolate, appressed, 2—3 mm. long; lower sepal ovate, others elliptical, obtuse, scarious 1940] BAKER: WESTERN VIOLETS 221 Hemrancne ve ak Capnar« wv (arerente Pepe FYieln sasarte Baker Piboctechin ehh ws Fruiting apeotmen: mrgian af peat beg, <8 oy EEE eiles north of Srockinga, doawpiiins Courrky. ee oe Oragen, dune 29, 1835, B. 6. 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