, ce ' ‘ t ' ' to a to4 eG ‘ 4 4 ) 7 y wy ‘ j , ' “t hath bade palo Maya tditatdtage fa) - 3 serve seal annhe ye | aired state jaysildielas haired eh diag’ iypidhan he feby ae | ' 3 Aajde ote bet fae fie | . ale | : sihaii 4 . | ‘ehete haijerttatig: bite ots | | : jarbstanitations tanger’ 4 aly phyartten atte ase ae ‘ sviatan ed . 1 Ags ay Pana. 7 pi sy silete ms | i ips fee i) pire ae ; ; af OLE Nils ‘ ‘ : oy ae Y . hoe ; oh (ae : aah mite i aie whe beg ale! | ; . . Pact ; utils i he . - : vet [ Ht 1 ' it \¢ Ve ies tat oa . . apt ten Malt Mtl Sis , | . rere eres iayan . “ ' a | | | ANAT ee 4s. | Natage' , . Satrarte, ree | Bay aiteS{9oree | a ed tleere cies ' i 1 ' : wij t | . Poth petela tale ' Fat 4 a nf nod & FV foagy £59 te" ! . ty ae pi a ede tghdts ' ‘ . ‘ | $ | | | | We a 4 sna gg . ee | say lees mony ‘’ oy 1 ou ¥ oat ‘ : ra . nite Vari : 2 ah sts i pe 7 i “ Teydyaets i ioeat oto4 I ty i 4 Peer erte hs ait Ha . 4s y | «hasay, ahh 9) a care etd Np | . ee ee “is - ive iene ag 3 ; ja itiotiy ay : its ral eH rh wi dui ia foiy fen 5,4 a 7 ae: feat ica Be fagnjulsa H ag ia 4 Lune Spies ire) Vv. 4 \ MADRONO A WEST AMERICAN JOURNAL OF BOTANY VOLUME VI 1941 - - 1942 ansonian NStitys: OSS Meet Be 2 "% O ab Ls RS - National Mus 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 Herszert L. Mason, University of California, Berkeley, Chairman. LeRoy Asrams, Stanford University, California. Epcar ANDERSON, Missouri Botanical Garden, St. Louis. Lyman Benson, University of Arizona, Tucson. Herpert F. Corpetann, Sacramento Junior College, Sacramento, California. Ivan M. Jounston, Arnold Arboretum, Jamaica Plain, Massachusetts. Mivprep E. Marnias, University of California, Berkeley. Bassett Macuire, Utah State Agricultural College, Logan. Marion Ownsey, State College of Washington, Pullman. Secretary, Editorial Board EruHe.t Crum, Department of Botany, University of California, Berkeley. Business Manager WittiAm Hiesey, Carnegie Institution of Washington, Stanford University, California. (09% ‘d 908) ADTIVWVY SIONVUT ‘ONOUAV] “9 DUINTO A 0} dddICsTQUOA ST CONTENTS Some Details of the Reproductive Structures of Sarcodes. Bernice HE. Doyel and L. Marguerite Goss A Study of Isoetes in San Diego County, California | Lowis C. Erickson A New Frasera from Oregon .. Morton EH. Peck and Hlmer I. Applegate New Plants from ‘Oregon... .:. ... i... 00.524 .. Morton EH. Peck Field Characters Distinguishing Pinus ponderosa and Pinus Jeffreyi. Kenneth E. Bradshaw A New Species of Astragalus from Arizona .............. C. L. Porter MiberbulvaGGini SO Weetsenns oa. ..6 4c 8 cuaw Waa dal LeRoy E. Detling A New Species of Paronychia from Mexico ............ .... Harl L. Core Great Basin Plants—III. Caryophyllaceae......._.. .. Bassett Maguire A New Limnanthes from Oregon ........................ LeRoy Abrams RUCVAC WiSHee eee, a tht Gan os tte athe 4 Gab Racha aoknd 28, 58, 90, 142, 206, IN@EES BP aAndeyNG Wii.) 2 kite ae eos sas bo ale Ue ee eb eed 30, 63, 94, 144, 173, Marlarionyimeyucca Whipplel 22.0. 20.5 cus 8S eee Sheek tae Lee Haines Certain North and South American Distributions in | Scirpus. Alan A. Beetle The Problem of Life Zones on Mount Shasta, California. William Bridge Cooke A New Species of Lotus from the Mount Hamilton Range, California. Helen Kk. Sharsmith Proceedings of the California Botanical Society ....... 64, 96, 175, 208, Alpine Flora of San Francisco Mountain, Arizona .... Hlbert L. Little, Jr. Mary Fisk Spencer ..... Frederick Grover New Species of Vascular Plants from the Northw est Coast. George Neville Jones Noteworthy, Plants from Idaho... ....... 62.684 64.5 8.65 Arthur Cronquist MiaskarCedaranm California . oo... 5. ec eee eh te ee Herbert L. Mason Further Studies on Monotropoideae................ Herbert F. Copeland Podocarpus ¢gracilior in Cultivation .. ............4...%. John T. Buchholz The Taxonomic Status of Microsteris Greene ......... Herbert L. Mason The Holacanthoid Plants of North America . _. . Cornelius H. Muller Combinations Proposed in “The Higher Plants of Oregon.” i Morton EH. Peck An Undescribed Species of Stipa from California. G. L. Stebbins, Jr., and R. M. Love A Synopsis of the American Species of Cicuta. Mildred EH. Mathias and Lincoln Constance Great Basin Plants—VI. Notes on Gentiana.......... Bassett Maguire mMrmicauimonlaska and YUKON of. eda ad Palen cee is Bassett Maguire A Comparison of the Embryogeny of Picea and Abies John T. Buchholz Anatomy and Ecology of Ammophila arenaria Link |... . Edith A. Purer An Undescribed Species of Ceanothus from California. Howard E. McMinn Development of the Female Gametophyte in Erythronium helenae and Erythronium tuolumnense .. : Marion S. Cave Some Chemical Properties of Eucalyptus in Relation to Their Evolutionary SLU SMR nite Aare Ath ape MWR Ns puhaid tothcne ng 6 James B. McNair Grassland and Related Vegetation in Northern Mexico Forrest Shreve (CIE OZ SRT Ma ED et POA a eR I’, P. Cronemiller Notes on Polemoniaceae . ._.................. Herbert L. Mason Eburophyton Heller: A Valid Genus of the Orchidaceae. Louis O. Williams : The Significance of Certain Plant Names — .. Carl Sumner Knopf ‘ Notes on the Flora of the Charleston Mountains, Clark County, Nevada. WV Pe SUAS AUST ee oh Se hake ako! Seale sells. Ira W. Clokey : The Type Locality of Polystichum Lemmoni -Underw ood ~=6@ Harold St. John ‘ i 265 239 33 239 Kar Western Noveltiessin, Salixv. <tr e Carleton R. Ball 227 The Genetic Approach to Problems of Rare and Endemic Species. G. Ledyard Stebbins, Jr. 241 he Ly pev of, Clematis hirsutissiman’.... 0.0 _ Ralph O. Erickson 259 Rrancis sWwamaley :Occ 2 ee a. oan ee eae etre eee eee Edna Louise Johnson 260 Indexto! Wolume” Vale 290 oj eee Aa ert Man ee emer tee ere 267 OEP ata cyte hha dl cvs alc ee UNE eh Oe de CN Res CMe HE cle 272 lv SOME DETAILS OF THE REPRODUCTIVE STRUCTURES OF SARCODES Bernice E. Dover anp L. MArGuerire Goss The genus Sarcodes includes but one species, S. sanguinea Torr., the snow flower, a fleshy, saprophytic plant common in the Sierra Nevada in California. This familiar plant has been described in detail by Oliver (6). Each plant consists of a tuft of roots and a stout erect stem which bears spirally arranged scales. The abundant flowers are borne in the axils of the upper scales; the lower part of the stem may be regarded as a peduncle, and the upper as the rachis of a raceme. The stem, scales, pedicels and sepals are densely cov- ered with multicellular glands. All parts above ground are bril- liant crimson. The flowers are ordinarily pentamerous. The sepals are separate; the petals are united for about half of their length and are entirely glabrous. The corolla and calyx are about equal in length; the tips of the petals curve outward while those of the sepals are slightly incurved. The stamens are ten in number, hypogynous, considerably shorter than the petals and slightly shorter than the pistil. The anther is about half the length of the entire stamen. The filament is ribbon-like, entirely glabrous, and attached to the outside of the anther a little below the middle. Dehiscence takes place by a pair of pores on the outer surface of the anther near the summit. The pistil is glabrous. The oblate- spheroidal ovary is marked by ten grooves in the planes of the filaments; at the base it bears a nectary with ten blunt lobes lo- cated between the grooves and projecting between the bases of the filaments. The internal cavity of the ovary is divided by five radiating septa located in the planes of the sepals and bearing massive placentae; in the lower part of the ovary the inner ends of the placentae are fused together so that the placentation is axile, while in the upper part they fail to meet and the placenta- tion is parietal. The placentae bear a very large number of ovules. The cylindrical style, about as long as the ovary, is tra- versed by an open channel into which ridges, being upward con- tinuations of the septa, project. The capitate stigma is five- lobed; the lobes are placed alternately with ridges which project into the style-channel: that is, they are the ends of the carpels. We are able to amend or extend Oliver’s description by obser- vations of the vascular system of the receptacle, the structure and development of the stamen and the embryogeny. The observa- tions are based on material supplied by Dr. Herbert F. Copeland, to whom we are most sincerely grateful. The material was col- lected at Jonesville, Butte County, at an altitude of 5000 feet, and fixed in Bouin’s fluid. In making slides the usual microtechnical methods were employed and safranin and light green were used exclusively as the stains. Manprono, Vol. 6, pp. 1-32. January 17, 1941. JAN « VAN 23 194) 2 MADRONO [ Vol. 6 Vascutar ANATOMY The vascular tissue of the stalk—the peduncle and the rachis of the inflorescence—is to be interpreted as a cylinder, in which, however, the xylem is broken up into separate strands (Pl. 1, fig. 1). There is no sheath of fibers outside the phloem.t| The departure of each leaf trace leaves one gap in the cylinder; the leaf trace is apparently to be interpreted as a single bundle but early in its course it frequently divides into two and before it enters the leaf breaks up intomany. The vascular supply to each flower originates as two bundles, springing from the sides of the leaf-gaps just above the departure of the leaf trace; the two bundles unite in the cortex to form a cylinder, which enters the pedicel. In traversing the pedicel it breaks up into a ring of some five or six bundles. As the bundles enter the receptacle (Pl. 1, fig. 2), a series of six whorls is given off in acropetal sequence: (1) a whorl of five sepal bundles; (2) a whorl of five bundles alternating with the sepal bundles, each of which splits tangentially into (a) several outer bundles that enter the corolla and (b) one inner bundle supplying an antipetalous stamen; (3) a whorl of five antisepal- ous stamen bundles; (4) a whorl of many carpel lateral bundles, arising on the inner sides of the bases of all the stamen bundles, and ascending the ovary wall; (5) a whorl of carpel dorsals ascending the ovary wall in the planes of the petal bundles; (6) a whorl of placental bundles in the planes of the sepal bundles. All the bundles in the ovary wall fade out just before reaching the style. The placental bundles enter the ovary in the planes of the septa and continue into and up the style. They fade out in the stigma. THe STAMEN In the youngest stamens we have seen, the relative sizes of the parts are about as at maturity; that is, the filament is rather 1“My statement [5] that there is a sheath of fibers in Sarcodes was based on inadequate observation”—(orally communicated by H. F. Copeland). EXPLANATION OF THE Ficures. PLATE 1. PuaTE 1. SARCODES SANGUINEA Torr. Fic. 1. Cross section of peduncle, X 4. Fic. 2. Half of a model of the vascular system in the receptacle, seen from the inside, X 16: ca, sepal bundles; co, petal bundles; st, stamen bundles; cd, carpel dorsals; cl, carpel laterals; plac, placental bundles. Fic. 3. Juvenile stamen, X16. Fic. 4. Diagram of a longitudinal section of a stamen, showing the course of the vascular bundle, x16. Fic. 5. Cross section through anther, showing pollen chambers at the plane represented by the line a in figure 4, x 40. Fic. 6. Mature stamen, showing open pores, X16. Fics. 7, 8, 9. Cross sec- tions of mature anther, the planes of which are represented respectively by the lines a, b, c in figure 6, x40. Fic. 10. Cross section of anther wall, being an enlargement of the area marked by the dotted square in figure 9, x 320. 1941 | DOYEL AND GOSS: SARCODES 3 pas plac a AP) eLe. o SOG PLaTE 1. SARCODES SANGUINEA ToRR. zy | MADRONO [Vol.6 longer than the anther and enters on the outer side below the middle (Pl. 1, fig. 4). The stamen bundle bends so as to run not to the summit of the anther but to the lower end. The anther bears two rudimentary descending horns just above the insertion of the filament on the outside and below the pores (PI. 1, fig. 3). In the young anther the usual four microsporangia with wall and tapetum are present (Pl. 1, fig. 5). The two pollen sacs on the inner side of the stamen extend somewhat nearer to the summit than those on the outside. The cells of the epidermis or exothe- cium are tanniniferous except for a patch on the outer side of the upper end of each anther lobe extending across the ends of the two microsporangia of the lobe where the pores are to form. In the youngest material we have examined the pollen grains had already formed. Most of them are four-grooved. As the anther matures the wall between the two microspor- angia of each lobe breaks down (PI. 1, figs. 5, 7, 8); the connec- tive (the wall in the plane of symmetry) also breaks down through a small part of its extent at the base of the anther; for the most part it persists. Where the pores are to form, and below them in the horns, a body of cells within the epidermis becomes differentiated by thick ribbed walls (PI. 1, fig. 10). Dehiscence appears to be accomplished by a contraction of the exothecium which tears the body of thick walled cells in two by a lengthwise slit; the slit becomes a pore rimmed by thick-walled cells (Pl. 1, figs. 6,9). EMBRYOGENY The ovules have an integument of about four layers of cells. In the young ovule a few cells of the outermost layer show indica- tions of tannin being present. Later, nearly all the cells of this layer are tanniniferous. The embryo sac develops in the manner usual in Sympetalae. The archesporial cell forms a megaspore tetrad. The spore at the chalazal end of the tetrad enlarges (PI. 2, fig. 11); its nucleus undergoes three successive divisions which result in four nuclei at the chalazal end and four nuclei at the micropylar end. One nucleus from each end moves toward the center of the embryo sac; there these two nuclei unite to form the endosperm mother nucleus. Cell membranes form around the remaining six nuclei. The three resulting cells at the chalazal end are the antipodal cells. The largest of the three at the micro- pylar end is the egg and the other two are the synergids (PI. 2, fig. 13). Meanwhile, the nucellus and the other three spores have disappeared. Following fertilization and before the embryo divides, the endosperm goes through a series of divisions. Each nuclear division is followed by cell division and the first two divi- sions produce a row of four cells (PI. 2, figs. 14, 15). The sec- ond and third cells of this row (counting from the micropylar end) 1941] DOYEL AND GOSS: SARCODES 5 C; [o J SOSTISCOS. 1 renetooae We" AGS By eS: wee oOr eS tf e oA NAY 6 Hes PiatEe 2. SARcODES SANGUINEA Torr. Longitudinal sections of ovules and young seeds in successive stages of development. Fics. 11-18, x 160. Fic. 19, 6 MADRONO [Vol. 6 divide repeatedly and produce many cells while the first and fourth cells develop into haustoria (Pl. 2, figs. 16-19). At the same time the antipodal cells and the synergids disappear. The zygote first divides into two cells after the endosperm is past the four-celled stage (Pl. 2, fig. 17); of the two daughter cells of the zygote, the one toward the micropyle elongates and forms a suspensor. ‘The suspensor carries its sister cell up to about the level of the wall between the first and second cells of the four celled endosperm. The original suspensor cell undergoes no divi- sions and grows no farther; its sister cell, by a series of divisions, adds to the suspensor, so that the summit of the embryo is carried in among the cells derived from the second cell of the four-celled endosperm. In this region a globular mass of cells, the definitive embryo, is developed (PI. 2, fig. 19). By this time the integu- ment has for the most part been absorbed, except for the outer layer of cells which have become thick-walled; the seed is essen- tially mature. DIscussION Copeland (1, 2, 3, 4, 5) has made several contributions to the knowledge of the details of the reproductive structures of Mono- tropoideae. Our studies show that the details of the reproductive structures of Sarcodes are essentially in agreement with what he has found. The outer whorl of stamen bundles and the carpel- dorsals are in the planes of the petal bundles. The placental bundles are in the planes of the sepals. The embryo sac is nor- mally developed and of normal type. In certain minor details of the development of the embryo, in which, according to Oliver’s figures, Sarcodes is different from other Monotropoideae, we were unable to corroborate his findings; apparently Sarcodes agrees perfectly with other members of the group. The integument of most Monotropoideae is of just two layers of cells and the endosperm is of but few cells. The four-layered integument and the mature endosperm of many cells in Sarcodes are distinctive characters of the genus. The course of the bundle in the anther, running as it does to the base and not to the summit, shows that the end of the anther which is toward the base of the flower is the distal end and the end toward the summit of the flower is the proximal end. The stamen develops, in fact, with the distal end of the filament turned in and down, so that the outer, apparently dorsal, surface of the anther is in reality the ventral surface. This structure of the stamen seems to be typical of the order Ericales. The horns, found in many Ericales on the ventral surface of the anthers, are absent in most Monotropoideae, though conspicuously present in Pterospora. We think it significant that rudimentary horns, previously unreported, are present in Sarcodes. The four-grooved pollen grains of Sarcodes are like those of 1941] ERICKSON: ISOETES 7 Pterospora and Pleuricospora. The pollen grains of Allotropa are three-grooved, while those of Monotropsis have two grooves. Copeland (5) expressed the opinion that Sarcodes together with Pterospora and Allotropa make up the most primitive tribe of Monotropoideae, being the link between the Ericaceae proper and other Monotropoideae. The characters of Sarcodes as we have come to know them lead us to believe that this is the true conception of the line of evolution. Sacramento Junior College Sacramento, California, January, 1940 LITERATURE CITED 1. CoreLtann, H. F. The structure of the flower of Newberrya. Madrofo 2: 137-142. 1934. On the genus Pityopus. Madrono 3: 154-168. 1935. . The reproductive structures of Pleuricospora. Madronfio 4: 1-16. 1937. The structure of Allotropa. Madrofio 4: 137-153. 1938. The structure of Monotropsis and the classification of the Monotropoideae. Madrono 5: 105-119. 1939. 6. Otiver, F. W. On Sarcodes sanguinea Torr. Ann. Bot. 4: 303-326. 1890. oe gol A STUDY OF ISOETES IN SAN DIEGO COUNTY, CALIFORNIA Louis C. Erickson Three species of Isoetes occur in San Diego County, Cali- fornia: I. Nuttall A. Br., I. Orcuttii Eaton, and I. Howellii Engelm. To these species, eleven names or combinations of names have been applied at various times and the validity of at least one of them, J. Orcuttii, has been questioned by Norma E. Pfeiffer, the most recent student of the genus. (Monograph of the Isoetaceae. Ann. Mo. Bot. Gard. 9: 79-232. 1922.) In the present study ecological and morphological aspects of the genus are empha- sized. The writer is indebted to Dr. A. W. Haupt and to Dr. Carl Epling, both of the University of California, Los Angeles, for sug- gestions. The living material studied came from the Kearney Mesa which is about fifteen miles north of the city of San Diego and five to ten miles inland, and from a pool about eight miles farther north. This area is a table-land drained by a system of small streams which have running water only after rains. These streams retain occasional pools along their courses, eventually drying up completely during the long rainless period lasting from May until November or December. Between streams are low mounds covered with a chaparral vegetation, alternating with shallow depressions which retain water throughout the rainy sea- son and in which many small hydrophytes flourish. A more de- tailed account of this region may be found in a recent article by 8 MADRONO [Vol. 6 Edith A. Purer (Ecological study of vernal pools, San Diego County. Ecology 20: 217-229. 1939). Numerous specimens from fourteen localities within the area described were collected at different times of the year. Pre- served material, collected in the same region some years previ- ously by Dr. A. W. Haupt and Dr. O. A. Plunkett, was also exam- ined. In addition, types and representative specimens of the fol- lowing were studied: Isoetes Suksdorfit Baker, I. Nuttalla A. Br., I. Orcutt Eaton, I. Howellii Engelm., I. nuda Engelm., I. melanopoda var. californica Eaton; also representative specimens of I. Under- woodit Henderson, the type of which has been destroyed by fire. The following descriptions have been based on the material col- lected near San Diego. KEY TO SPECIES Corm 3-lobed; velum complete Peripheral strands in leaves 3; megaspores frosted and pre- dominantly stuberculate 2. 0p...) ee 1. I. Nuttallu Peripheral strands in leaves lacking; megaspores usually glossy and predominantly smooth .................... 2. LT. Oreuttu Corm 2-lobed; velum one-third complete ..................... 3. I. Howelli 1. TIsortes Nurtarin A, Br..ex, Engelm.,.Am, Nat. 8): 9215) 1874; Pfeiffer, Ann. Mo. Bot. Gard. 9: 1380. 1922. I. opaca Nutt. ex. Engelm., St. Louis Acad. Sci. 4: 388. 1882. I. Suksdorfu Baker, Handbook of the Fern Allies 1382. 1887. Calamaria Nuttallua Kuntze, Rev. Gen. Pl. 2: 828. 1891. C. Suksdorfi Kuntze, l. c. Corm 3-lobed; leaves 5—75, mean 21, length 2.5—20 cm., mean 8 cm., spreading, usually with a characteristic twist, stomata nu- merous, peripheral strands 3, subterranean part of mature leaves occasionally with some brown pigment, velum complete, mem- branaceous margin up to 5 ecm. in length, tapering gradually; megaspores gray (or sometimes white, gray, and dark brown to black within one sporangium), dark brown to black when wet, 260-560 uy, mean 880 y, the markings variable, from distinctly tuberculate to etuberculate on a frosted or occasionally glazed surface ; microspores 25—81 y, mean 27.5 yu, tuberculate. Isoetes Nuttallii occupies seepage areas along small streams like those found on Kearney Mesa. In contrast with the other two species, I. Nuttall does not grow where water stands, although the plants may be partially submerged after rains. It was noted in several instances that plants growing in the wetter places were being destroyed by worms and bacteria. The soil in which the plants grow varies from a coarse sand containing a small amount of clay to a clay without sand. Many of the plants are shaded by the chaparral shrubs bordering the streams. Variation may be observed in (a) number and length of leaves, (b) color, size and marking of megaspores and (c) depth of the 1941] ERICKSON: ISOETES 9 corms in the soil. The number of leaves per plant varies from 5 to 75; their length from 2.5 to 20 centimeters. The plants which have the largest number of leaves are not always the plants with the longest leaves. Most megaspores are gray, but within some sporangia three colors may be found: white, gray, and dark brown to black. When plotted according to size, the megaspores of some individuals produce a bimodal curve. Others have mega- spores which, when plotted, produce unimodal curves with small dispersion. Between these extremes are all intermediate condi- tions, some producing skewed unimodal curves, and some produc- ing less pronounced bimodal curves. The marking on mega- spores is variable. Ordinarily they are densely covered with tubercles against. a frosted surface. In most sporangia, however, some megaspores usually occur on which the tubercles are wholly or partly suppressed; it is the usual condition for the commissural faces of megaspores to be more prominently marked than the free surfaces. The corms may be covered to a depth of 3 to 4 centi- meters or, on the other hand, may be so shallow that the sporangia are visible; all intermediate conditions may be found. The existence of long-leaved and short-leaved plants, three colors of megaspores in a single sporangia and megaspores which produce bimodal curves when plotted for size may be regarded as evidence of gene mutations. 2. Isontes Orcutrtmn Eaton, Fern Bull. 8: 18. 1900; Pfeiffer, Ann. Mo. Bot. Gard. 9: 182. 1922. I. Nuttall var. Orcutti Clute, Fern Allies 253. 1905. Corm 3-lobed; leaves 8-25, mean 8, length 3-6.5 cm., mean 4 cm., spreading, stomata numerous, peripheral strands none, pig- ment absent from subterranean portions of mature leaves, velum complete, membranaceous margin 1 cm. or less long, narrow; megaspores gray, dark brown when wet, 220-400 y, mean 320 u, smooth and glossy, or rarely frosted, sometimes remotely tubercu- late; microspores 23-30 yp, mean 26.5 uy, tuberculate. Isoetes Orcuttit occupies vernal pools only and is submerged during most of the growing season. After the water evaporates from these pools the plants mature in desiccating soil. In all instances I. Orcuttu was found growing in clay soil which is soft when wet and extremely hard when dry. In contrast to Isoetes Nuttallu, I. Orcuttit presents very little variation. The leaves exhibit a considerable range in number (3-25) and in this species the larger leaf numbers are associated with the longer leaves. In leaf length the variation is slight (3- 6.5 cm.) and does not show the bimodal tendency found in J. Nuttallu. The megaspores are uniformly gray with a character- istically glossy surface. Some of the spores have small remote tubercles, and rarely, some of them may be frosted instead of glossy. The megaspores of J. Orcuttit are much more uniform in 10 MADRONO [Vol. 6 size thar those of either of the other two species. The corms may be covered to a depth of one centimeter at most. Isoetes Orcuttii occupies a different habitat from that of I. Nut- tall: and differs also in the following morphological characters: the leaves are fewer and smaller and peripheral bundles are ab- sent; megaspores are predominantly glossy and smooth and aver- age approximately 60 y less in diameter than those of J. Nuttalli. It is evident that although J. Nuttallii and I. Orcuttii are similar morphologically they differ sufficiently to be considered distinct species. 3. Isorres Howetiu Engelm., Trans. St. Louis Acad. Sci. 4: 385, 7 18823) Preitter’ Ann. ion Bot. Gard. 921139 19225 nuda Engelm., /.c. I. Underwoodii Henderson, Bot. Gaz. 23: 124. 1897. I. melanopoda var. californica Eaton in Gilbert, Working List of N. Am. Pterid. 27. 1901. Corm 2-lobed; leaves 4-56, mean 22, length 5-28 cm., mean 17 cm., spreading, stomata numerous, peripheral strands 4-12, subterranean part of mature leaves usually with abundant dark brown pigment, velum one-third complete, membranaceous mar- gin extending as much as 8 cm. above the soil level and narrowing gradually; megaspores white, tan when wet, 230-600 up, mean 430 u, usually distinctly marked with a combination of tubercles and distinct and anastomosing crests; microspores 27—39 uy, mean 34 yu, tuberculate and occasionally spinulose. Isoetes Howellii is much more widely distributed than either I. Orcuttu or I. Nuttall, for while each of these species is strictly limited to a particular habitat as described previously, I. Howellii may be found in association with either or may frequently occur alone. It occurs in shallow, quickly drying pools, in pools which retain shallow water and muddy soil, in deeper pools, or in stream beds which are often shaded by chaparral plants. The popula- tion of each pool or stream is very limited in area. It is very im- probable that the individuals which occupy one vernal pool or stream interbreed with those of another pool or stream. The sporangia are borne too far below ground level for the spores to be blown by the wind and there are no indications that water runs from one pool to another. | The extent to which these differences in environment influence the plants is not known. Nevertheless, there are striking differ- ences between populations of I. Howellu. These differences are most evident in the number and length of leaves and the size of megaspores. The first two populations which are summarized in Table 1 exist in two pools which are separated by about fifty feet of higher ground. The first of these is a shallow depression in which the water stands for a short time after rains and, with the exception of Isoetes and Pilularia, is quite free from plants during the presence of the standing water. In the other pool the 1941] ERICKSON: ISOETES 11 TABLE 1. Differences between populations of Isoetes Howellii Habitat Leaf length No. leaves Megaspore size 1 partially submerged; 8-18, 14-46, 280-410, not crowded mean 14 cm. mean 26.5 mean 350 u 2 submerged; crowded 10-25, 5-20, 320-480, mean 18 cm. mean I1 mean 400 uw 3 partially submerged; 5-12, 10-31, 300-450, not crowded mean 8.5 cm. mean 17.5 mean 380 wu 4 submerged; crowded 7-16, 4-14, 350-600, mean 11.5 cm. mean 8.5 mean 450 wu 5 shaded 9-18, 7-26, 300-580, mean 13.5 cm. mean 17 mean 430 uw water covers the plants during most of the growing season, since by seepage of water it is kept filled for a longer period. Here the Isoetes plants are crowded and are in association with small spe- cies of sedges. The differences between these populations are not wholly unexpected. In the widely spaced plants of the first population which grow in shallow water and mud uninfluenced by other plants the leaves are more numerous and shorter than in plants of the second population in which the individuals are crowded and submerged. The size of the megaspores also varies, being smaller by an average of 50 y in the plants of the drier more open pool. Another partially submerged population, Table 1 number 8, which grows several miles from the two just described is made up of individuals about two-thirds the size of the partially submerged plants of the first population. Similarly, another submerged crowded population, number 4, is made up of individuals about two-thirds the size of the submerged plants of the second popula- tion, in these the megaspores are decidedly larger by an average of 50 u. The plants thus far considered have been confined to pools. The last population to be considered, number 5, occupies a stream bed and is shaded to a large extent by overhanging chaparral shrubs. These plants differ noticeably from the others by their darker green, more slender leaves. In other characters they are intermediate. The range in variation in the size of megaspores of individual | plants may be very great. Measurements of 400 megaspores from one specimen, indicated a range of from 280—550 u a disper- sion exceeding those given by Miss Pfeiffer for both Isoetes How- ellii and var. minima Pfeiffer. In morphological characters there is complete intergradation between the species and the variety. University of California, Los Angeles, California, July 24, 1939. 12 MADRONO [Vol. 6 A NEW FRASERA FROM OREGON Morton E. Peck ann Emer I. APPLEGATE Frasera umpquaensis sp. nov. Planta biennalis glaberrima; caulis robustus simplex 6—9 dm. altus; folia omnia verticosa foliis verticis 3—4, vel superioribus oppositis, elliptico-oblongis vel late lanceolato-oblongis 1—2 dm. longis ad petiolum brevissimum indis- tinctum contractis; inflorescentia interrupta densa 1-3 dm. longa ramis infimis in axillis foliorum superiorum, bracteis parvis pedi- cellis plerumque brevioribus floribus; lobis calycis linearibus vel lanceolato-linearibus paulum inaequalibus 9-12 mm. longis, quoque ad basin intus cristam setarum brevium ferente; corolla viridescente alba paulo breviore calyce profunde 4-partita cristas setarum longarum inter lobos ferente, lobis ad apicem minute 3—4- dentatis, foviis solitariis magnis profundis suborbiculatis in cir- citu membrana profunde fimbriata marginatis; filamentis anguste linearibus ; capsulis compressis. Glabrous biennial; stem solitary from a short caudex, stout, simple, 6—9 dm. high; leaves all in whorls of 8 or 4 or the upper- most opposite, elliptic-oblong or broadly lance-oblong, 1-2 dm. long, narrowed to very short, ill-defined petioles; inflorescence 1-3 dm. long, interrupted, dense, the lowest branches in the axils of the upper leaves, the bracts above much reduced, the pedicels a little longer to much shorter than the flowers; calyx-segments linear to lance-linear, somewhat unequal, 9-12 mm. long, each with a tuft of short setae at base within; corolla greenish white or pale yellowish, a little shorter than the calyx, deeply 4-parted, with a tuft of long hairs just below each sinus, the narrowly ovate-oblong divisions 3- or 4-toothed at tip, the nectariferous pit near the base large and deep, suborbicular, extending nearly across the segment and bordered all round by a narrow membrane bearing a fringe of long setae, the area below the pit bearing similar setae; filaments filiform, about half the length of the corolla-divisions ; capsules compressed. Type. Anderson Camp, Umpqua-Rogue Trail on the summit of the divide, northwest corner of Jackson County, Oregon, at about 6000 feet elevation, July 11, 1929, Applegate 5930 (Dud- ley Herb., Stanford Univ.). Additional collection: slopes of Abbott Butte, Rogue River National Forest, July 2, 1936, Thomp- son 13067. This large robust plant has the general aspect of Frasera spe- ciosa Dougl. and F. fastigiata (Pursh) Heller but is more closely related to the latter, having quite similar foveae on the corolla, but differing in the setae and in the characters of the calyx. The species is probably confined to the Cascade Mountains along the divide between the upper Rogue and Umpqua rivers. Willamette University, Salem, Oregon, March 25, 1940. 1941] PECK: NEW PLANTS FROM OREGON 13 NEW PLANTS FROM OREGON Morton E. Peck The plants here described have been known to the writer for many years, but he has been hesitant about naming them, hoping that they might be assigned to species already published. This having been found impossible, they are now described as new. Sophora Leachiana sp. nov. Caulis erectus ad basin simplex 3-5 dm. altus minute canescenti-tomentosus; foliis compluribus supra confertis 1-2 dm. longis, foliolis 19-33 late oblongis utroque rotundatis 1.5—2.5 cm. longis tenuibus villoso-tomentosis, subter pallidioribus supra parce adpresso-pubescentibus; racemo ter- minali 7-15 cm. longo floribus 10-25 in pedicellibus 2—5 mm. longis; calyce late tubulari-campanulato supra valde gibboso 7—9 mm, longo lobis brevibus late triangularibus; petalis flavis 9-12 mm. longis; fructu maturo inviso, immaturo valde sursum curvato terete, inter semina constricta, ut videtur, stipite 3-4 mm. longo; seminibus paucis. Stem erect, simple below, 3—5 dm. high, finely grayish-tomen- tose; leaves several, somewhat crowded above, 1—2 dm. long, the leaflets 19-33, broadly oblong, rounded at both ends, 1.5—2.5 cm. long, thin, villous tomentose and paler beneath, thinly appressed- pubescent above; racemes solitary or few, terminal, 7-15 cm. long, the flowers 10—25, on pedicels 5 mm. long or less; calyx broadly tubular-campanulate, strongly gibbous above, 7-9 mm. long, the teeth short and broadly triangular; petals yellow, 9-12 mm. long; mature fruit not seen, immature, strongly curved up- ward on a stipe 83-4 mm. long, few-seeded, apparently constricted between the seeds, gray-tomentose. Type. Rand Ranger station near Galice, Josephine County, Oregon, June 18, 19388, Mrs. Lilla Leach 4343 (type in private herbarium of Mrs. Leach, Portland, Oregon). Sterile material of this interesting plant was collected in 1921 by Douglas C. Ingram (1221) of the United States Forest Service “on the trail to Pea-vine Mt.”, probably within a mile or two of the type locality. Mr. Ingram’s material was doubtfully referred to Amorpha by government taxonomists. Since then Mrs. Leach has collected flowering and young fruiting material near the same locality on three occasions, thus making possible a correct diag- nosis. The plant blooms freely but apparently fruits very spar- ingly. Mrs. Leach has collected very extensively in southwestern Oregon, and would probably have come across the plant else- where were it not of extremely local distribution. I take pleasure in adding this to the list of Oregon plants that bear her name. Sidalcea maxima sp. nov. Caules erecti dense caespitosi 8—12 dm. alti robusti ad basin saepe 1 cm. crassi, glauci glaberrimi 14 MADRONO [Vol. 6 usque ad inflorescentiam; foliis supra glabris subter sparse et minute puberulis, foliis caulinis compluribus inferioribus et radi- calibus 6-10 cm. latis prope ad medium 7-9-fissis, segmentis plerumque leve 3-lobatis, mediis et superioribus 3—5-partitis seg- mentis lanceolatis vel linearibus 3-10 cm. longis; racemis usque ad 2.5 dm. longis infra glabris supra minute puberulis; floribus sparsis; pedicellis brevissimis minute puberulis, bracteis lineari- bus integris subtentis; calyce 8-13 mm. alto sparse et minutissime stellato, lobis angustis triangulari-ovatis; petalis clare roseo-pur- pureis latis apice prope truncato 2—3 cm. longis; carpellis (imma- turis) dorso levibus. Stems in large dense clusters, 8-12 dm. high, robust, often 1 em. thick at base, freely branched from near the base, glaucous and completely glabrous to the inflorescence; leaves glabrous above and nearly so beneath, with only a few minute stellate hairs, the cauline leaves rather numerous, the basal and lower 6—10 cm. wide, cleft nearly to the middle into 7-9 mostly shallowly 3-lobed divisions, the middle and upper cauline parted into 3—5 mostly entire lanceolate or linear divisions 5-10 cm. long; racemes up to 2.5 dm. long, sparsely flowered even in anthesis, the rachis glabrous below, minutely puberulent above; lower flowers of the raceme on slender glabrous stalks 1-8 mm. long, the very short and finely puberulent pedicel subtended by an entire linear bract apparently near the summit of the pedicel; calyx 3-11 mm. high, thinly and very minutely stellate, the lobes narrowly triangular- ovate; petals bright rose-purple, broad, nearly truncate at apex, 2—3 cm. long; carpels (immature) apparently smooth on the back. Type. On moist bank along Dairy Creek, twenty miles north- west of Lakeview, Lake County, Oregon, July 3, 1927, Peck 15435 (Herb. Willamette Univ.). This tall robust plant is characterized by its almost complete lack of pubescence and its large showy flowers. SipaLcEa spicaTa (Regel) Greene var. tonsa var. nov. Folia subter plerumque sine capillis longis; calyx sine capillis longis patentibus, lobis interdum bracteis adpresso-ciliatis; inflores- centia plerumque patentior pedicellis longioribus. Leaves rarely with any long hairs beneath; calyx without any long spreading hairs, but the calyx-lobes as well as the bracts often appressed-ciliate; inflorescence usually less dense, often with longer pedicels. Type. Meadow, Big Summit Prairie, Ochoco National For- est, Oregon, June 30, 1932, Peck 17224 (Herb. Willamette Univ.). Typical Sidalcea spicata is common in the southern counties of Oregon from Curry County to Lake County, reaching its most characteristic development west of the Cascade Mountains. To the north and east of this area it passes gradually into the variety here described. Though numerous intergrades occur along the 1941] BRADSHAW: PINUS 15 indefinite boundary of the ranges, the differences elsewhere are so strongly marked and consistent that it seems advisable to dis- tinguish the two by name. The variety is plentiful throughout most of Oregon east of the Cascades from northern Klamath and Lake counties northward and eastward. Willamette University, Salem, Oregon, March 25, 1940. FIELD CHARACTERS DISTINGUISHING PINUS, PONDEROSA AND PINUS JEFFREYI KENNETH E. BrapsuHaw! Western conifers offer relatively few problems in taxonomic differentiation to the field man, but one frequent source of confu- sion and controversy lies in the similarity between the common western yellow pine (ponderosa pine), Pinus ponderosa Dougl. ex Laws. and its close relative Jeffrey pine, Pinus Jeffreyi Grev. and Balf. ex A. Murr. These species may be found occupying sepa- rate ecological niches (Jeffrey pine has a higher elevational range and occurs on drier sites than western yellow pine and will replace it on serpentine formations at the lower elevations) or they may be found growing intermixed. While certain typical stands or individual trees may be quite readily identified, others defy identi- fication by the use of a simple key and generalized descriptions of the species in question. The following pages contain a list of comparative external features which should greatly facilitate the separation of the two species in the field. Remarks on segrega- tions based on the internal structural and chemical qualities of the wood and foliage, which are essentially tasks for the laboratory, have not been included. Grateful acknowledgement is made of certain technical assistance given the writer by Mr. Lloyd Austin of the California Forest and Range Experiment Station. When using these comparisons, it must be remembered that it is seldom adequate to attempt to identify a tree by using one char- acter to the exclusion of the others. Numerous local strains with distinct morphological and physiological differences result in ex- treme variation. Certain trees have characters of both species, due probably to cases of inter-breeding. At times but one fea- ture, such as characteristic cones beneath an isolated tree, or dis- tinctly brownish inner bark scale surfaces, may be used as a primary distinction which would point to a Jeffrey pine. Certain trees, however, which otherwise resemble western yellow pine have cones similar to those of the Jeffrey pine or the brownish inner bark scale surfaces characteristic of that species. There- 1 Junior Range Examiner, Soil Conservation Service, United States Depart- ment of Agriculture, Berkeley, California (Formerly Assistant to Technician, California Forest and Range Experiment Station, Berkeley). 16 MADRONO [Vol. 6 Younc TREES Pinus ponderosa Fourace: texture relatively fine, yel- low-green, glossy; striae of stomata fine, scarcely distinguishable. Leaver: flaky bark high on slender leader, length of roughened new growth short; dull gray; resin-filled pustules none. Bups: orange or brick red, conic-ovate, acute; scales closely appressed, sur- face with resinous exudations, gener- ally in form of numerous tiny drop- lets. Twics: surface of season’s growth shinging green, previous years’ growth brownish. Bark: soft, somewhat resinous; ridges wide, not much inter-connected; fur- rows Shallow, flaky with bark scales; varying from blackish to yellow- brown, tawny or dull orange; inner surfaces of bark scales powdery dull to brilliant sulphur-yellow. Pinus Jeffreyi Foriace: needles’ thicker, longer, coarser, blue-green; powdery white because of the prominent striae of stomata which are well defined and readily counted. Leaver: flaky bark not extending so high on the somewhat stouter leader, leaving conspicuous long, smooth, sil- very gray area; resin-filled pustules numerous. Buns: darker with purple or chocolate brown tone; somewhat stouter, more elongate, less acute; scales without resinous exudations, tips less closely appressed. Twics: surface of season’s growth green, glaucous or pruinose; previous years’ growth brownish or greenish but retaining the bloom in perceptible density. Bark: hard, non-resinous; ridges nar- row, irregularly connected giving a braided appearance; furrows deep, distinct, not flaky, laminations of bark very distinct on edges; generally dark gray with slight purplish or reddish- brown cast; inner surfaces of bark scales creamy pinkish- or chocolate- brown. fore a dominance of one group of characters over the other must be the basis for establishing an identification, unless the tree in question represents a typical case of direct hybridization. The writer has found that certain characters are best used in the identification of young trees, while others are more serviceable in segregating the older trees. With this consideration in mind, there have been set up practical groups of data for each of these two broad age classes. Trees of intermediate age may be identi- fied by the use of either one of the groups or parts or all of both. Variations in the presence and quality of significant factors with age, size, and condition of the individual trees must be taken into account when using the descriptions contained in this paper. In evaluating the characters of special structures the following modifying conditions should be noted: Fouiace. Foliar differences have been found useful only with the smaller pole-size and seedling trees; they are best used when the two species are growing intermixed so that relative compari- sons can be made. Twies. Color of the twigs or previous years’ growth has been found to be exceptionally constant as a distinguishing feature; it 1941] BRADSHAW: PINUS 17 Op TREES Pinus ponderosa Fourace: often perceptibly yellowish; relatively sparse, in ball-like tufts at ends of branchlets, needles persisting approximately 3 years. Bark: generally tawny, yellow-brown or dull orange; inner surface of scales, especially near ground level powdery dull to brilliant sulphur-yellow, fre- quently a distinctive and reliable char- acter; interior of scale brown or tan; scales somewhat soft, a small piece usually dislodged when lightly struck with fingernail; small dark resin pits present throughout. Opor: slight, resinous. Limps: stout, often grotesquely gnarled and bent in very old trees; compara- tively short, straight and stout in younger mature specimens, sometimes slightly upturned at ends. ConrEs: somewhat ovate; about 3-6 inches long, the spreading scales ap- pearing slender and widely spaced; prickles short, the points over most of the cone protruding outward from the umbos, often hooklike, evident to the touch. Pinus Jeffreyt Foi1acE: somewhat blue-green, darker ; more dense, needles slightly longer and coarser, persisting 5 to 8 years. Bark: externally, color similar to that of P. ponderosa, or dark reddish brown, or wine color; plates similar to those of P. ponderosa or in more typical specimens somewhat narrower with deeper separating grooves; inner surface of scale light creamy pinkish- or chocolate-brown; interior of scale deep reddish; scales more. glossy, harder, merely dented when lightly struck with fingernail; resin pits lack- ing. Opor: rather strong, pleasant, sweet, described as resembling that of pine- apple, mellow apple, vanilla, or violet. Limps: less stout and angled, often slender, elongate, more distinctly up- turned at ends in typical specimens; retained on bole somewhat longer, re- sulting in a longer, more symmetrical crown. Cones: elliptical or long-oval, resem- bling an old-fashioned bee-hive; about 6 to 10 inches long, scales relatively more numerous, stout, closely com- pacted, projecting almost horizontally from the cone axis, thus appearing heavier and denser; prickles long, mostly deflexed, the points straight or even slightly turned in, seldom pro- truding outward except sometimes on the upper 4 to 5 whorls of scales, scarcely perceptible to the touch. is the only reliable means for determining the species of very young seedlings. Bark. Ridge and furrow characters are rather distinctive but hard to see on very small trees just beginning to form bark, and not so apparent on trees over sixteen inches in diameter at approximately breast height. Opvor. The use of odor in identification is best employed on older trees, but may be found helpful to supplement the appear- ance of the bark in young trees which are beyond the stage where the leader and foliage color can be relied upon. Certain indi- viduals of Pinus ponderosa and Pinus Jeffreyi have disturbingly similar odors, and since hybrids become chemically as well as physically intermixed, this feature should not be relied upon solely. Variation in olfactory sensitivity among observers also decreases the value of this method of differentiation. 18 MADRONO [Vol. 6 Cones. Unopened cones of both species are quite as useful for identification as the opened cones. The immature cones of Pinus ponderosa are generally green in color, while the cones of Pinus Jeffreyi are generally purple, but there are so many varia- tions and reversals of this rule that color should not be considered an identifying feature in all localities. Cone lengths are also variable, departing from the approximations presented here to such an extent that, except for specimens showing extremes of size, they should be considered much less reliable than the other features mentioned. Soil Conservation Service, United States Department of Agriculture Berkeley, California, December, 1939. A NEW SPECIES OF ASTRAGALUS FROM ARIZONA! C. L. Porter Astragalus Beathii sp. nov. Radix perennis; caules plurimi, 4—6 dm. longi, striati seu sulcati, glabri; folia 10-15 cm. longa, foliolis 11-21, plerumque oppositis, nunc ellipticis, obtusis, nunc ovato-obcordatis, basi in petiolulum perbrevem attenuatis; racemi 10 ad 20 flori, floribus densis, purpurascentibus; calyx oblongus, pilis albis, dentibus brevibus lanceolato-subulatis; vexillum ova- tum, attenuatum, obtusum, fere 22 mm. longum; carina obtusa; legumen cartilagineum, glabrum, fere 38-4 cm. longum, oblongi- cylindraceum, semibiloculare, polyspermum, sutura superiore ob- tusa, inferiore introflexa; legumen, sectione transversa, rotundum videatur; semina reniformia, fere 3 mm. longa. Plants perennial, many stemmed from the summit of a strong taproot, the stems erect, glabrous, striate to sulecate; leaves pin- nately 11- to 21-foliolate, 10-15 cm. long, strigose when young, becoming glabrate when mature; leaflets varying from elliptical and obtuse in upper leaves to ovate-obcordate in basal leaves, those of the basal leaves often much smaller, subopposite on the rachis, and narrowed below into a very short petiolule; racemes 10- to 20-flowered, the flowers dense, purple; calyx oblong, white- strigose, the teeth short lance-subulate, about one-third the length of the tube; banner ovate, attenuate at base, obtuse at apex, about 22 mm. long, moderately arched; keel obtuse; legume coriaceous when mature, fleshy when young, sessile or subsessile, glabrous, about 8—4 cm. long, 7 mm. wide and thick, oblong-cylindrical, rounded in cross section, the upper suture obtuse, not prominent, the lower suture intruded and forming a thick septum about 2 mm. high within; seeds numerous, reniform, about 3 mm. long. 1 Contribution no. 181 from the Department of Botany and the Rocky Mountain Herbarium of the University of Wyoming, Laramie. 1941 ] PORTER: ASTRAGALUS 19 Piate 3. AstracaLus BeatHi Porrer. Fic. 1, median section of mature pod x5; fig. 2, seed x 5; fig. 3, mature pod in ventral view x1; fig. 4, basal leaf X1; fig. 5, leaflet of cauline leaf <1; fig. 6, flower x 5. 20 MADRONO [ Vol. 6 Type. Two miles south of Cameron, Coconino County, Ari- zona, June 10, 1939, L. N. Goodding, Sel. 34-39 (Rocky Mountain Herbarium; isotype, Missouri Botanical Garden). This species may be referred to section Preusii of Jones (Rev. Astrag. 1923) and to the genus Jonesiella of Rydberg (N. Am. FI. 24: 401. 1929). The elongated, straight, cylindrical, sessile or subsessile pods and the dark purple flowers, as well as the erect many-stemmed growth from a strong taproot will readily dis- tinguish it from any of its near relatives. It appears to be very limited in distribution, being found only in the type locality as far as is known, but there it is quite abundant and conspicuous. The writer takes pleasure in naming this plant for Professor O. A. Beath who first discovered it while on a field trip in connec- tion with his work on seleniferous plants. Since he was unable to obtain mature fruit at the time he first saw it, he informed Mr. L. N. Goodding of the locality and it was through Mr. Goodding that the type material was later obtained. University of Wyoming, Laramie, January 24, 1940. ALBERT RADDIN SWEETSER (1861-1940) In the death of Dr. Albert Raddin Sweetser the Pacific Coast has lost one of its best-loved teaching botanists. Dr. Sweetser was born at Mendon, Massachusetts, July 15, 1861, the son of a Methodist minister. His early education was obtained in Massa- chusetts, in the public schools and at Wilbraham Academy. Entering Wesleyan University, Middletown, Connecticut, in 1880 he received from that institution the degree of Bachelor of Arts in 1884 and of Master of Arts in 1887. Then followed a year at Massachusetts Institute of Technology, where he took special work in chemistry. His first teaching was done in the public schools, Cape Cod, and later at Bucksport, Maine, where he taught in a Methodist seminary. While at Bucksport, in 1888, he was married to Carrie K. Phinney, whom he had met on Cape Cod. His next teaching position was in a Methodist school at Tilton, New Hampshire. In 1893 he entered the Harvard Graduate School of Botany, and remained there for four years. During his last two years at Harvard he was an assistant in botany and at the same time was teaching in Radcliffe College. In 1897 he accepted an invitation to join the faculty of Pacific University at Forest Grove, Oregon. He went to the University of Oregon in 1902 as Professor of Biology, and in 1909 became head of the Department of Botany, a position he occupied until his retirement in 1981. In that year the faculty of the University of Oregon conferred upon him the honorary degree of Doctor of Science. PLatre 4. Atpert RappdDIN SWEETSER. 19441] CORE: PARONYCHIA 21 He died at Eugene, September 12, 1940, at the age of seventy-nine years. Dr. Sweetser was co-author with Mary E. Kent of a small popu- lar “Key and Flora” of Oregon, and the author of a number of shorter scientific articles. During the later years of his life he became greatly interested in the botanical history of the Pacific Northwest, and has left a great deal of collected material on this subject, none of which, unfortunately, has been published. His greatest contribution, however, has been as a teacher, and it was in this field that he himself felt that his most important work lay. His deep understanding of the problems of young people, his kindly personal interest, and his keen sense of humor have endeared him to the many students who have come in con- tact with him during his years of instructing. It was his purpose to build an appreciation of nature in as many people as possible, and to this end he wrote a considerable number of popular botani- eal articles for newspapers and journals and accepted frequent invitations to lecture before various clubs. Another of his major interests was the conservation of the native flora and much of his work was directed to that end. He was a member of Sigma Xi, the American Association for the Advancement of Science, the Medical Society of America, the Society of American Bacteriologists, the Botanical Society of America, the American Association of University Professors, the National Geographic Society, and an associate member of the American Museum of Natural History.—LeRoy E. Det.irne, Uni- versity of Oregon, Eugene. A NEW SPECIES OF PARONYCHIA FROM MEXICO* Earyu L. Core While checking over several sheets of Paronychia from various herbaria, the writer came across a specimen from the collection of Dr. Edward Palmer which apparently belongs to an undescribed species. For this plant the following name is proposed: Paronychia albomarginata sp. nov. Herba perennis, e caudice crasso lignescente; caule ramosissimo a basi incipiente, 4-7 cm. longo, puberulente cum internodis numerosis brevis; foliis lineari- bus vel linearibus-oblongis, puberulentissimis, 3-4 mm. longis, acutis vel mucronatis a stipulis celatis; stipulis argenteis folia aequantibus vel excedentibus; bracteis brevioribus quam flores vix eis paribus; cymis parvis; calycis segmentis 2 mm. longis, puberulentibus, cum marginibus albis conspicuis, in cuspide breve et erecta terminantibus; staminibus sepalorum longitudine dimidiis. 1 Contribution No. 12 from the Herbarium of West Virginia University. 22 MADRONO [Vol. 6 Perennial from a thick woody tap root; stem much branched from the base, 4-7 cm. long, puberulent, with numerous short internodes; leaves linear or linear-oblong, densely puberulent, 3—4 mm. long, acute or cuspidate, hidden among the stipules; stipules silvery, ascending, equaling or exceeding the leaves; bracts shorter than or barely equaling the flowers; cymes small, few- flowered; sepals 2 mm. long, puberulent, with conspicuous white margins, tipped by a short erect cusp; stamens half the length of the sepals. Type. Saltillo, State of Coahuila, Mexico, April 9, 1905, Ed- ward J. Palmer 518 (Britton Herbarium, New York Botanical Garden; co-type in Herbarium of Missouri Botanical Garden). This plant is obviously related to Paronychia Wilkinsonu S. Wats., from which species it differs principally in the length of the sepal awns, which in P. Wilkinsonu are half the length of the sepals or longer and widely spreading, whereas in the present spe- cies they are very short and erect. Other differences are in the leaves, which in P. Wilkinsoni are 4-6 mm. long, in P. albomar- ginata only 3—4 mm. long; in the stipules, which in P. Wilkinsonii are widely spreading, about equaling the leaves, and in the pres- ent species erect, generally exceeding the leaves; and in the size | of the flowers, about 3 mm. long in P. Wilkinsoni, and only about 2 mm. long in P. albomarginata. West Virginia University, Morgantown, West Virginia, March 7, 1940. GREAT BASIN PLANTS—III. CARYOPHYLLACEAE Bassett MAGUIRE The North American pinks have received no serious study? since the revision by Robinson in 1897. Progress in the knowl- edge of our western forms has undergone little advancement since. In attempting to identify new material and to order that of the herbarium more than ordinary difficulty has been encountered, due to this lack of progress and to the multiplicity of forms occur- ring in recent collections. In his treatment of the family, Robin- son (4) constantly referred to the complexity and confusion exist- ing initstaxonomy. Hultén (2, p. 166) has remarked concerning a segregate of Cerastium Beeringianum, ““Owing to the great vari- ability and lack of distinctive characters the taxonomy of the above-mentioned plants is very confusing.” This statement might well apply to the entire family and, in our concern, par- ticularly to the family in our region. 1The first part of the revision of the Western Hemisphere members of Spergularia by Dr. R. P. Rossbach appeared in the March, 1940, issue of Rhodora. 1941] MAGUIRE: CARYOPHYLLACEAE 23 In anticipation of further more extended studies, the follow- ing preliminary report may now be made. Crerastium BreErinaianum Cham. & Schlecht. var. GRANDIFLORUM (Fenzl.) Hultén. From a considerable and diversified population which apparently must come under the above specific name, the variant with petals exceeding 9 mm. (following Hultén) is prob- ably best referred to var. grandiflorum. This variety is apparently a new record from our region. These plants possibly represent also C. pulchellum Rydb. and C. Earlet Rydb. ARENARIA FENDLERI Gray subsp. genuina nom. nov. A. Fendleri Gray, Mem. Am. Acad. n. ser. 4:18. 1849. ARENARIA FENDLERI Gray subsp. brevifolia subsp. nov. Caudici- bus multicipitalibus, caulibus numerosis, plerumque _ basibus foliorum vestito, 4-20 cm. altis; foliis 1-3 (6) cm. longis; in- florescentibus 1-3 (5) ecm. longis, glandulosis; sepalis anguste lanceolatis, acutis; petalis elliptico-oblongis, obtusis vel retusis, 7-9 mm. longis. | Stems numerous, 4-20 cm. high, from a multicipital caudex borne on a thick root, the stem bases conspicuously clothed with grey-brown remains of previous leaves; leaves 1-3 (6) cm. long; inflorescence open or somewhat congested, 1—3 (5) cm. long, con- spicuously glandular-pubescent; sepals narrowly lanceolate, acute, hardly acuminate, the inner more broadly scarious-margined, 5—7 mm. long; petals elliptic-oblong, obtuse or merely retuse, 7-9 mm. long, claws 1-2 mm. long; capsule shorter than the sepals. Type. Meadow, 11,300 feet, Burro Pass, La Sal Mountains, Grand County, Utah, July 18, 1933, Bassett Maguire et al. 17972. Cotypes. La Sal Mountains, Bassett Maguire et al. 1777, 179783, 17974; Abajo Mountains, San Juan County, Utah, Maguire & Redd 1175. (Type and cotypes are deposited at Utah State College.) Plants of meadows or open slopes and ridges from 10,000 to 13,000 feet known only from the La Sal and Abajo mountains of Grand and San Juan counties, Utah, and one collection (Maguire 12254) from Kaibab National Forest, Coconino County, Arizona. This population is distinguished from the Rocky Mountain race by its shorter leaves, longer petals, somewhat broader and less attenuate sepals, and shorter inflorescence. At the highest elevations it passes into the variant, described below, with much reduced leaves and stems, distinctly the alpine ecotype of the plant of subalpine, Canadian, and Hudsonian habitats. ARENARIA FENDLERI Gray subsp. BREviFoLia var. brevicaulis var. nov. Caulibus plerumque 4-8 cm. longis, foliis 1-2 cm. longis. Stems mostly 4-8 cm. long; leaves mostly 1-2 cm. long; sepals occasionally only 4 mm. long. Type. Summit of Pilot Mountain, 12,400 feet, La Sal Moun- 24 MADRONO [Vol. 6 tains, Grand County, Utah, July 11, 193838, Bassett Maguire et al. 17976. Cotypes. La Sal Mountains, Maguire et al. 17977, 17978, 17979, 17980. (Type and cotypes are deposited at Utah State College.) This low compact plant is evidently the western correlative of var. Portert Rydb. (of which the isotype, M. E. Jones 716, is de- posited at this institution) of Arenaria Fendlert subsp. genuina. It is probable that var. brevicaulis represents, at least in part, Are- naria compacta of the manuals of “Utah.” It would seem improb- able that our plant is conspecific with A. compacta Cov. of Cali- fornia, although Coville (1, p. 67) compares it with A. Fendleri, since var, brevifolia is definitely a phase of A. Fendleri, whereas this latter species does not occur to the west of the Great Basin. ARENARIA CAPILLARIS Poir. subsp. formosa (Fischer) comb. nov. A. formosa Fischer, in De Candolle, Prodromus 1: 402. 1824. This entity, apparently not before recorded from Nevada, rep- resents the typical American form of A. capillaris. The polymor- phic population of subsp. formosa has developed a number of vari- ants, the definition of which is not yet clear. This subspecies is represented by the following collections: San Jacinto, Elko County, Nevada, June 138, 1989, Maguire 16811, 16812; Diamond A Ranch, vicinity of Jarbridge, Elko County, Nevada, May 29, 1939, C. York. ARENARIA ABERRANS M. E. Jones. A striking Arenaria suggest- ing A. capillaris but with immense urnulate capsules resembling those of Silene was collected under pine woodland near the south gate of Grand Canyon National Park (Coconino County, Arizona, June 27, 1935, Maguire 12236). These plants are undoubtedly an offshoot of A. capillaris but have undergone such extreme diver- gence as to necessitate recognition as a distinct species. It was not realized that this collection represented the second one made of the recently described A. aberrans M. E. Jones (Contrib. West. Bot. 37. 1980) ascribed to “Box Elder Co., Utah.” The type (W. P. Cottam 4159), however, actually came from Mount Dellan- bough, Mohave County, noeeherr Arizona, about eighty miles west of our station. ARENARIA EastwoopiaE Rydb. var. aDENopHORA Kearney & Peebles. This interesting desert ecotype, recently described (1939) from the vicinity of Tuba, Coconino County, Arizona, ex- tends northward, as might have been expected, into southeastern Utah. Collections of this variant from San Juan County, Utah, are represented by the following: June 29—July 1, 1933, Maguire 2900, 5901, 5902, 5908. Silene Petersonii sp. nov. Rhizomate gracile, cum gemmis conspicuis nodis; caulibus solitariis vel laxe multicipitalibus, 5-15 - 1941 ] MAGUIRE: CARYOPHYLLACEAE 25 cm. altis, simplicibus, aspere retrorse glanduloso-pubescentibus, plus dense nodis; foliis oblanceolatis, 2-3 cm. longis, 4-6 mm. latis, 1-nerviis, aspere glanduloso-puberulis vel scabris; floribus 38-7, raro 1; calycibus campanulato-cylindricalis vel inflatis, 1.5—2 em. longis, membranis, conspicue 10-nerviis (nervis purpureis) glanduloso-pubescentibus, lobis oblongo-ovatis, obtusis, 4-6 mm. longis, glanduloso-ciliolatis, corollis conspicuis exsertis, roseis ad roseo-purpureis, petalis 2-3 cm. longis, limbis cuneatis, tenue den- tatis vel 4-lobatis, lineari-oblanceolatis; capsulis ovato-oblongis, seminibus 2—2.5 mm. latis, tuberculis marginalibus bullatis. Tap root deep set, giving rise to several slender, light brown rhizomes, 1-2 mm. in diameter, producing conspicuous nodal buds, the rhizomes horizontally extensive, branching, giving rise to single or several loosely multicipital branches; stems simple, 5-15 em. high, harshly and retrorsely glandular-pubescent; leaves oblanceolate, 2-3 cm. long, 4-6 mm. broad, broadly or narrowly sessile by means of connate, sheathing hyaline bases, 1-nerved, harshly glandular-puberulent, the upper leaves. somewhat re- duced; flowers 3-7, or seldom only 1, nodding during anthesis, pedicels 1-2.5 cm. long, seldom only 5 mm. long, calyx campanu- late-cylindrical, conspicuously inflated at maturity, 1.5 (1)—2 cm. long, membranous, sometimes purple-tinged, conspicuously 10- nerved (nerves purple), glandular-puberulent, the hairs fre- quently pigmented, lobes oblong-ovate, obtuse, 4 (3)—6 mm. long, glandular-ciliolate; corolla conspicuously exserted, pink to rose-purple, petals 2 (1.5)-—38 em. long, the blade broadly cuneate, irregularly toothed, shallowly lobed, or 4-cleft into linear-oblance- olate lobes, the lateral smaller, appendages inconspicuous, claws broad, gradually narrowed to the base; capsule ovate-oblong, somewhat shorter than the calyx, opening by 6 smooth valves, (rarely 4 or 5), stipe 1-2 mm. long, styles 3; seed 2—2.5 mm. broad, laterally compressed, brown, with a conspicuous broad marginal crest of bullate tubercles. Type. Common on steep, loose, bare, calcareous slopes at 10,900 feet, Skyline Drive, one mile above Baldy Ranger Station, Manti National Forest, Sanpete County, Utah, August 8, 1940, Bassett Maguire 20000 (Utah State College). Cotypes. Loose calcareous sand-clay rim, frequent at 10,500 feet, one-half mile east of museum, Cedar Breaks National Monument, Iron County, Utah, July 12, 1940, Bassett Maguire 19465; frequent in bare gravelly clay on rapidly eroding slopes, Red Canyon, Garfield County, Utah, July 16, 1940, Bassett Maguire 19550. Other collections. Utan. Iron County: Cedar Breaks, August 12, 1988, Hitchcock, Rethke § van Raadshooven 4582, August 7, 19384, Maguire 17977, August 13, 1939, Maguire 17566, June 23, 1940, Maguire 19019, July 12, 1940, Maguire 19451, August 18, 1940, R. T. Clausen & H. Trapido 5109. Garfield County: Red Can- yon, August 9, 1934, Maguire 17987, June 25, 1940, Maguire 19060, 26 MADRONO [Vol. 6 July 16, 1940, Maguire 19533, July 16, 1940, Maguire 19556; Pink Cliffs at 9000 feet, Table Cliff Plateau, Powell National Forest, June 26, 1940, Maguire 19121. Sanpete County: Sky Line Drive, August 8, 1940, Maguire 19989, August 8, 1940, Maguire 20001 (topotypes). It is fitting that this attractive new Silene be named in honor of Dr. E. G. Peterson, President of Utah State Agricultural College, who made possible the establishment of this herbarium, and who since, by constant sympathy and personal encouragement has been most helpful in the progress of its work. Lycunis Drummonpi (Hook.) Wats. In Utah and the Great Basin the tall-stemmed Lychnis has been recognized as represent- ing the three following species: L. Drummondii, with “included corolla” and “sessile capsule’; L. nuda, with “exserted, 4-lobed petals,” “inflated obovate calyx,” and “‘stipitate capsule’; and L. striata, with “exserted 2-lobed petals,” “oblong calyx” and “‘ses- sile ovary.” Specimens recently collected in Utah show these characters in the following combinations, but with all degrees of intermediacy obtaining. Plants of the latter characters show no ecological or geographical segregation, hence are at most only varietally dis- tinct: (a) corolla more or less included, petals bifid or lobed, ovary subsessile or short-stipitate, calyx ovate-oblong; (b) co- rolla more or less exserted, petals (2-) 4-cleft or lobed, ovary sub- sessile or short-stipitate, calyx ovate-oblong. Robinson (4) reduced Lychnis nuda Wats. to Silene pectinata var. subnuda, citing the Watson Humboldt Mountains collection and, without number, two collections of M. E. Jones: “Near Em- pire City and at Franktown, Nev.” A collection from Empire City, Nevada (June 19, 1882, M. E. Jones 3795, Utah State Col- lege no. 5746), although immature, is without question Lychnis and probably represents part of the series cited by Robinson. It would seem then that Robinson was probably in error in the trans- fer. Tidestrom (5, p. 200) accepted the name of Watson as representing a valid species. Nelson (3) reduced the name L. striata to synonomy under L. Drummondii. The writer has not seen the type of Lychnis nuda and realizes that the “inflated calyx” does not obtain in L. Drummondu. How- ever, since the Utah material, which surely represents but a single major species, transcends the characterization of these three for- merly proposed entities, it seems best to treat the plants with exserted and mostly 4-lobed or 4-cleft petals as a variety: Lycunis Drummonpu (Hook.) Wats. var. nuda comb. nov. L. nuda Wats. in King, Geol. Expl. 40th Par. 5:37. 1871. L. striata Rydb. Bull. Torr. Club 31: 408. 1904. Utah State Agricultural College, Logan, Utah, April, 1940. 1941 ] ABRAMS: LIMNANTHES 27 LITERATURE CITED 1. Covirtz, F. V. Descriptions of new plants from southern California, Nevada, Utah and Arizona. Proc. Biol. Soc. Wash. 7: 65-80. 1892. Hutten, Eric. Flora of the Aleutian Islands. Stockholm. 1937. Netson, AvEN. New Manual of Rocky Mountain Botany. 1909. Rozrinson, B. L. [Caryophyllaceae] Synoptical Flora of North America 1: 208-255. 1897. ' 5. Tipestrom, Ivar. Flora of Utah and Nevada. Contrib. U. S. Nat. Herb. 25: 1-665. 1925. p= oS) A NEW LIMNANTHES FROM OREGON LeRoy Asrams Limnanthes Howelliana sp. nov. Caules ex basi 1 vel plures, 12-25 ecm. alti, succulenti, glabri. Folia glabra, infera 5-8 cm. longa, infra divisionibus 2- ad 38-lobatis, supra integris, lineari- oblongis vel linearibus; sepala lanceolata, acuta, 7-8 mm. longa, glabra; petala alba, basi flava, obovata, emarginata, 12-16 mm. longa, apice subtruncato et late emarginato, infra 9-nervia, infra medium sparsim longo-villosa; carpella dense obtuso-tuberculata. Stems one to several from the base, 15-25 cm. high, rather succulent, glabrous. Leaves glabrous, the lower 5—8 cm. long, lower divisions 2- to 3-lobed, the upper entire, linear-oblong or linear; sepals lanceolate, acute, 7-8 mm. long, glabrous; petals white with yellow base, obovate, emarginate, 12-16 mm. long, 6—7 mm. wide at the subtruncate broadly emarginate apex, 9-nerved below, sparsely long-villous below the middle; nutlets thickly beset all over with broad mammillate tubercles. Type. Roadsides and fields near Wilbur, Douglas County, Oregon, April 8, 1894, J. W. Thompson 10161 (Dudley Herbarium 230576). Other collections of this species in Dudley Herbarium are: ditch north of Oakland, J. W. Thompson 4390; Yoncalla, J. W. Thompson 186. All of these stations are in Douglas County on the Umpqua River watershed. Thomas Howell gave a good description of this species in his “Flora of Northwest America” under the name Limnanthes Doug- las R. Br. and I therefore take pleasure in naming it in his honor. It differs from L. Douglasii, a species of the Coast Ranges of cen- tral and northern California, in several characters. The nutlets of that species are smooth or crowned at the apex with a few tubercles, the leaflets or divisions several-toothed or -lobed, whereas in L. Howelliana they are entire or the lower 3-lobed, as in L. alba Hartw. Limnanthes rosea Hartw. has the nutlets thickly beset all over with protuberances, but they are thin, scale-like and whitish. Four other species of Limnanthes have been described from southern Oregon: L. Bellingeriana M. E. Peck, L. floccosa Howell. L. gracilis Howell, and L. pumila Howell. Dudley Herbarium, Stanford University, California, February 5, 1940. 28 MADRONO [Vol. 6 REVIEWS The Cacti of Arizona. By Lyman Benson, J. J. THornseEr, and A. A. Nicuou, with line drawings and three color plates by Lvu- CRETIA BREAZEALE Hamitton. University of Arizona Bulletin vol- ume 11,no. 1, pp. 1-134, with 52 plates, 6 in full color, and colored frontispiece. (Biological Science Bulletin no. 4 [or 5?]) Univer-. sity of Arizona, Tucson, Arizona, January, 1940. $1.00. This semipopular bulletin on the cacti of Arizona was pub- lished to fill the place left vacant by the exhaustion of the supply of Biological Science Bulletin no. 1, by Stockwell and Breazeale, issued in 1933. The discrepancy between the bulletin number shown on the cover and that appearing on the title page is puz- zling, for the cover bears the legend “Biological Science Bulletin No. 5” while on the title page it is called “Biological Science Bul- letin No. 4.”" One also wonders about the date, January 1, 1940, although it is not impossible that the bulk of the stock of bulletins was not distributed until some eleven months later. However, these minor details detract not one whit from the value of this handbook of the Arizona cacti. It is well planned, carefully written, and beautifully illustrated with plates in full color, half-tone, and line drawings. The profuse use of illustra- tions fulfills the authors’ hope that “. . . the more common cacti may be recognized without resort to the keys.” The keys are formal, dichotomously constructed and workable. The number of technical terms used in keys and descriptions has been held to just about the minimum, and those used, tersely defined in a short glossary or in the introductory paragraphs. | Distribution maps show the range, within Arizona, of sixty of the more important species recognized as occurring within the state. It is regrettable that the base maps, upon which the distri- bution is shown in green, were not drawn with bolder lines and with fewer intricate topographical details, for one must compare them with larger scale maps to determine with any assurance the position of many of the limits of distribution. The maps are valu- able in that in spite of the indistinct details they graphically present the general distribution of the species so treated. The taxonomic treatment of the genera is conservative, Car- negiea, Lophocereus, Lemaireocereus, Peniocereus, and Wilcoxia all being lumped in the older genus Cereus. In like manner, Echino- cactus becomes the repository for Ferocactus, Echinomastus, Sclero- cactus, Utahia, and Toumeya, while Coryphantha, Escobaria, and Phellosperma are relegated to synonymy under Mamillaria. This is in keeping with the generic treatment of these groups by several Europeans who have dealt with the cacti during the past decade, and may reflect a reaction to the criticism occasionally voiced concerning the treatment of Britton and Rose to the effect that these two gentlemen recognized too many genera and too few species. The authors’ concept of species seems to be more liberal than their definition of genera, for they recognize ten species of 1941] REVIEWS 29 Echinocereus, the validity of some of which has been questioned by cactus fanciers who usually allow quite narrow specific delimi- tations. Again, both Opuntia ursina and O. erinacea are given full specific value. (I’d do the same myself.) In other instances they have refrained from attempting to evaluate some of the more troublesome complexes in Opuntia and Mamillaria, frankly stating that several more years of critical study must be given these puz- zling groups. In spite of what one’s personal opinion may be concerning the treatment of the genera, the taxonomic treatment in this book is a practical one and doubtless will prove admirably “usable” to many westerners and travelers interested in the south- western flora. The first hand information supplied by these men who have studied the cacti of Arizona at all seasons will be most valuable to botanists to whom opportunities for field work in the Southwest come infrequently. The brief section dealing with the culture and care of cacti is sufficient for handling most of the native species. A. A. Nichol, who wrote this section of the book, refrained from giving detailed specifications, but included the necessary general hints covering the soil, water and light requirements of most of the Arizona cacti. The book is well printed on glazed paper, with paper cover, and at the modest price of one dollar should find wide acceptance among botanists and cactus growers. The illustrations make it an excellent book for the non-professional who is interested in making a “speaking acquaintance’ with the dwellers of the arid Southwest.—Ira L. Wicerns, Dudley Herbarium, Stanford Uni- versity. Flora of Indiana. By Cuarites C. Deam. Department of Con- servation, Division of Forestry. Pp. 1-1236, maps 1—2248, 4 un- numbered maps and frontispiece. Indianapolis, Indiana. 1940. $3.50. (Send order to State Forester, State Library, Indianapo- lis, Indiana.) Dr. Deam has set a new high standard for regional floras. This volume will be a constant source of reference not only for students of the Indiana flora but for all those engaged in floristic studies. Every effort has been made to present a complete and accurate picture of the flora of Indiana. Dr. Deam has incor- porated the results of field studies carried on through years of travel. Every species listed in the flora is based upon actual and accessible specimens and in many cases the identifications have been checked by specialists. Certain parts of the text have been contributed: the genera Carez, Juncus and Luzula by Frederick J. Hermann; Chenopodiaceae by Theodor Just; Crataegus by Ernest J. Palmer. Distribution maps are included for each species. These maps show not only the location but also the time of flower- ing and the herbaria in which the specimens are located. The author has included a discussion of the floral areas of Indiana; a list of names of collecting places; a reference list of 30 MADRONO [Vol. 6 Indiana collectors; and a bibliography of the Indiana flora. Eco- logical notes are given for many of the species and since these are based on the author’s own observations they are of particular value. The flora will undoubtedly be considered a model of excellence and thoroughness. Every student and writer of floras will do well to consult it not only as a model of form but because of the invalu- able information which it contains. Such a volume should be a constant companion in the field; it is to be regretted that it was impossible to publish all this material in a form more suitable for field use.—Mutprep E. Maruias, Department of Botany, Univer- sity of California, Berkeley, California. NOTES AND NEWS RIBES PETIOLARE DOUGL. IN CaLiForNIA. Occurring generally along streams and on moist sites from British Columbia south through the intermountain region to southern Oregon and Ari- zona, Ribes petiolare was first reported from California by George A. Root in 1987 (California Ranger 8°: 4, 1937). The species was found by Root and Hollis Day along Shovel Creek in Shasta National Forest, Siskiyou County, about nine miles south of the California-Oregon boundary line. In the same report Root men- tions finding a few bushes of R. petiolare one and one-half miles north of the boundary line on Kelley Creek near Lakeview, Lake County, Oregon, at the northern end of the Warner Mountains; he did not see it farther south in Modoc National Forest, Cali- fornia. In reviewing Root’s report, Clarence R. Quick (Madrofio 4: 286-290. 1988) mentions observing R. petiolare in abundance on Crane Creek a few miles north of Kelley Creek in Oregon; he also was unsuccessful in finding the species south of the line in the Warner Mountains. In July, 1939, the writer, while examining specimens in the herbarium of the United States Forest Service at Alturas, Cali- fornia, noted a collection of Ribes petiolare from ‘“‘Lost Lake-Silver Creek” in the southern Warner Mountains, Modoc National For- est, made August 18, 1918, by Associate Forester L.S. Smith. It bore the numbers 103838 and 31510, the latter the accession number of the Herbarium of the United States Forest Service, Washing- ton, D. C., to which Mr. Smith sent a portion of his collection. Dr. Frederick V. Coville verified the determination. The follow- ing herbaria have been consulted for records or specimens, but with negative results: University of California, Berkeley, Cali- fornia Academy of. Sciences, Stanford University, United States Forest Service and Division of Plant Exploration and Introduc- tion, Washington, D.C. Apparently Mr. Smith’s collection con- stitutes the earliest known record of the species in California and establishes its occurrence approximately seventy miles south of the Oregon line. 2 1941] NOTES AND NEWS 31 Additional specimens were obtained by G. A. Zentmyer and the writer from the same locality, July 30, 1939, and a portion of this collection has been deposited in the Herbarium of the Uni- versity of California, Berkeley. The species was observed in abundance along Silver Creek and several of its tributaries about one-half mile southwest of Lost Lake (particularly in Sec. 15, T.88 N., R.16E., Mt. Diablo Meridian). It was not found by the writer north of this locality in the Warner Mountains. The occurrence of Ribes petiolare near Lost Lake is considered of significance from the standpoint of blister rust control. The species is one of the most susceptible of western currants and dense concentrations of the bushes grow there in close associa- tion with an abundance of whitebark pine (Pinus albicaulis Engelm.), which according to present information is considerably more susceptible than other western white pines. The rust, how- ever, was not found on either pines or currants at this place.—J. L. Mievxe, Division of Forest Pathology, Bureau of Plant Indus- try, United States Department of Agriculture, San Francisco, California. In a recent issue of “Science” appeared an announcement of the death on August 20, 1940, of Dr. Joseph Burtt-Davy, lecturer in tropical botany at the Imperial Forestry Institute, Oxford. From 1893 to 1908 Dr. Burtt-Davy was connected with the Uni- versity of California, first as a student assistant, later as instruc- tor in botany and assistant botanist in the Agricultural Experi- ment Station. He left California to accept an appointment as agrostologist and botanist in the Department of Agriculture, Pre- toria, South Africa. Soon after the close of the World War he returned to England and was for some time engaged in the prepa- ration of “A manual of the flowering plants and ferns of the Transvaal with Swaziland, South Africa,” parts one and two of which were published in 1926 and 19382 respectively. During his sojourn in California Dr. Burtt-Davy, in connection with his studies on agronomy, visited many parts of the state and made collection of grasses and other forage plants; these collections, amounting to about two thousand numbers, are deposited in the University of California Herbarium. From 1896 to 1900 he pub- lished a number of articles in ‘““Gardeners’ Chronicle,” ‘‘The Pacific Rural Press,” and as bulletins of the University of California Agricultural Experiment Station and the United State Department of Agriculture; he also contributed more than fifty short articles to “Erythea.” While in Berkeley he married Alice Bolton. Dr. Burtt-Davy is well remembered by the older faculty members of the University of California. Important monographic studies published during the current year and not previously noted in MaproNo are: “‘the genus Ellisia,” by Lincoln Constance (Rhodora 42: 38-39. 1940); “‘the genus Dichelostemma,” by Robert F. Hoover (Am. Midland Nat. 24: 463- 32 MADRONO [Vol. 6 476. 1940); “studies in Penstemon VII, the subsections Gairdner- iani, Duesti, and Arenarii of the Graciles... ,” by David D. Keck (Am. Midland Nat. 23: 594-616. 1940); “‘a revision of the North American species of the genus Thermopsis,’ by M. M. Larisey (Ann. Mo. Bot. Gard. 27: 245-258. 1940); “a monograph of the genus Symphoricarpos,” by George Neville Jones (Journ. Arnold Arbor. 21: 201-252. 1940); “‘studies in the genus Hedysarum in North America,” by R. C. Rollins (Rhodora 42: 217-2389. pl. 597. 1940) ; “Spergularia in North and South America,” by R. P. Ross- bach (Rhodora 42: 57-838, 158-198, 203-213. pls. 589-596. 1940 ).—ETHEL Crum. Approximately five-hundred fifty species and varieties of native Californian and cultivated trees are included in a recent publication “Trees of Santa Barbara” by Maunsell Van Renssaeler of the Santa Barbara Botanic Garden, California. Descriptions are brief and non-technical and exact locations of more than eight-hundred specimen trees are given. For convenient refer- ence genera are arranged alphabetically, and both common and scientific names are listed in the index. The book of 141 pages is very attractive in format and is illustrated by a large number of excellent photographs and line drawings. Among those repre- sented by full page photographs are the following native Cali- fornian species: Ceanothus arboreus, Fremontia mexicana, Lyono- thamnus asplenifolius, Pinus Torreyana, Platanus racemosa. (Price: paper, $.75, cloth, $1.50.) —EtTuet Crum. Under the title “Alien plants growing without cultivation in California,’ Dr. W. W. Robbins lists approximately 526 species with comment as to the dates of their introduction into California, their habitats and distribution and their importance as weeds or forage. The pamphlet consists of 128 pages of which thirteen are devoted to introductory discussion, ninety-eight to comment on the species, seven to literature cited and fourteen to a compre- hensive index of common and scientific names. The publication appeared July, 1940, as Bulletin 637 of the Agricultural Experi- ment Station, University of California, Berkeley. At the Thirty-Fifth Annual Meeting of the Botanical Society of America, December 28, 1940 to January 1, 1941, Philadelphia, papers by the following botanists of western states were sched- uled: E. L. Little, Southwestern Forest and Range Station, Ari- zona; James Bonner, J. van Overbeck, California Institute of Technology; D. L. Arnon, Ernest Ball, Donald E. Bliss, R. W. Chaney, R. E. Clausen, Lincoln Constance, Frank Cuneo, Carl Epling, Katherine Esau, A. 8S. Foster, D. R. Hoagland, Flora Murray Scott, F. M. Turrell, University of California; G. H. Ross- bach, Stanford University; Joseph Ewan, University of Colorado. MADRONO A West American Journal of Botany A quarterly journal devoted to important and stimulating articles dealing with plant morphology, physiology, taxonomy, and botanical history. These volumes should be a part of every botanist’s li- brary and should be made accessible to students of all universities and colleges. Volume I, 1916-1929. . . $5.00 Volume II, 1930-1934 .. 5.00 Volume III, 1935-1936 . 5.00 Volume IV, 1937-1938 . 5.00 Volume V, 1939-1940 . . 5.00 Single numbers....... 0.75 The subscription price of MADRONO is $2.50 per year. We solicit your pat- ronage, Address all orders to: William Hiesey Bus. Mgr. Carnegie Institution of Washington Stanford University, California VOLUME VI NUMBER 2 MADRONO A WEST AMERICAN JOURNAL OF BOTANY Contents VariaATIon In Yucca Wurpptel, Lee Haines ........... 0... ccc cee 33 Certain NortH anv Soutsm AMERICAN DiIstTRIBUTIONS IN Scrmmpus, Alan A. Re Ma NC EAE On Sie Ue At EP OL SU 45 Tue Prosiem or Lire Zones on Mount Suasta, Carirornia, William Bridge ita be yh ca aed CUS NL ea ISSR Se GE SERS OOO SpA aT 49 A New Species or Lorus rrom tue Mount Hamitton RANGE, CALIFORNIA, EE STL COE STIPULIO Oe adh Suerte) oe LUNA US VAM a NAM oN OT 56 Reviews: Marion Ownbey, 4 Monograph of the Genus Calochortus (John L. Morrison) ; W. C. Muenscher, The Flora of Whatcom County, State of Washington (L. Constance) ; Jens Clausen, David D. Keck and William M. Hiesey, Experimental Studies on the Nature of Species (Herbert L. TESST NV ACSC I te OEE a ARs me 58 POC TRS Aner INIA EO LE en a Le ee a aM 63 PROCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY ..................-00005. 64 Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania April, 1941 MADRONO A WEST AMERICAN JOURNAL OF BOTANY Board of Editors Hersert L. Mason, University of California, Berkeley, Chairman. LeRoy Asrams, Stanford University, California. Epcar Anpverson, Missouri Botanical Garden, St. Louis. Lyman Benson, University of Arizona, Tucson. Hersert F. Copetanp, Sacramento Junior College, Sacramento, California. Ivan M. Jounston, Arnold Arboretum, Jamaica Plain, Massachusetts. Miuxprep E. Marnias, University of California, Berkeley. Bassett Macume, Utah State Agricultural College, Logan. Marion Ownsey, State College of Washington, Pullman. Secretary, Editorial Board—Etue.t Crum Department of Botany, University of California, Berkeley Business Manager—Wui1aAm HIsesey 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 proofs are returned. Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania, for the CALIFORNIA BOTANICAL SOCIETY, INC. President: Ernest B. Babcock, University of California, Berkeley. First Vice-President: Roxana S. Ferris, Stanford University, California. Second Vice-President: Palmer Stockwell, Institute of Forest Genetics, Placerville, California. Treasurer: Wiiliam Hiesey, Carnegie Institution of Washington, Stanford University, California. Secretary: Lincoln Constance, Department of Botany, University of California, Berkeley. Annual membership dues of the California Botanical Society are $2.50, $2.00 of which is for a year’s subseription to Madrofio. Dues should be remitted to the Treasurer. General correspondence and applications for membership should be addressed to the Secretary. 1941] HAINES: YUCCA WHIPPLEI 33 VARIATION IN YUCCA WHIPPLEI Lee HatInes Yucca Whipplei, a species which is widespread and abundant in southern California, is a member of the family Agavaceae Hutchinson. It ranges from Laguna Seca Chapala in northern Lower California (29° 80’ N. Lat.) to Chew’s Ridge in Monterey County, California (36° 30’ N. Lat.). The species varies greatly in appearance and it is apparent from casual observation that the variants are geographical in some degree. However, the nature of the genus Yucca is such that this variation can be studied ade- quately only in the field. Few herbarium specimens exist and these afford at best only the most general information as to the nature of the plant. The investigation has accordingly been carried on solely in the field. This was particularly necessary since it was found from preliminary observations that the size and conformation of the flower and fruit have essentially the same degree of variability throughout the species and that any conclu- sions must perforce be drawn from a comparative quantitative study of the vegetative characteristics and habit of the inflo- rescence. The author acknowledges his indebtedness to the board of graduate research of the University of California, Los Angeles, for funds with which the extended field work was carried on and his gratitude to Dr. Carl Epling, under whose guidance this work was done, to Dr. H. G. MacMillan for help in the preparation of the figures and to many colleagues for assistance in the field. DISTRIBUTION Before any observations or measurements could be made, it was necessary to discover where, in the vast area within its limits, Yucca Whipplet grows. The distribution area was divided into geographic sections and reconnoitered in an automobile by tra- versing as many of the highways, secondary roads, fire roads, and truck-trails as possible. By traveling approximately four thou- sand miles, the entire area was surveyed and the approximate dis- tribution limits of the species were established (fig. 1). During this reconnaissance, stops were made at favorable localities and observations and measurements were recorded. Such a locality was treated as a field station, designated by a number, and located on a topographic map. In Lower California the distribution has not been determined in detail. The species was observed at the Laguna Seca Chapala, in the mountains east of Rosario, in the chaparral-covered coastal hills near Sacaton between Ensenada and San Vicente, and north of Ensenada. In southern California it is found in the mountains from Campo northward through the Laguna and Balkan moun- Manrono, Vol. 6, pp. 33-64. April 15, 1941. APR 21 194] 34 an MADRONO [Vol. 6 tains to the San Jacinto and Santa Ana mountains. It is con- tinuous in a narrow strip from the San Jacinto Mountains to the San Bernardino, San Gabriel, Santa Susana, and Santa Monica mountains to Mount Pifios and the Tehachapi mountains. Here the area divides, one branch extending northward in the Piute and Fic. 1. Distribution of Yucca Whipplei in California and Baja California. ' G, subsp. Parishii; P, subsp. percursa; I, subsp. intermedia; C, subsp. caespitosa; T, subsp. typica. Greenhorn mountains and to the western margin of Sequoia Na- tional Park, the other passing northward through the Santa Ynez, San Rafael, and Santa Lucia mountains to Chew’s Ridge, north of Tassajara Hot Springs in Monterey County. The species is mari- time from the Santa Monica Mountains northward.’ Southward from Walker Pass to the eastern end of the San Bernardino Moun- tains, it occurs on the margin of the Mojave Desert. The distribution of Yucca Whipplei is not continuous through- out its range. There are major gaps between the Santa Ana and the Santa Monica mountains, between the Santa Monica and the Santa Ynez mountains, between the San Rafael and the Santa Lucia mountains, and between the Walker Pass region and the region along the western border of Sequoia National Park. Within these lesser areas the species is either diffuse and general or may be localized. Throughout its entire range Yucca Whipplei usually occurs in very porous, shallow soil or on rocky outcrops at elevations from sea level up to 6000 or 8000 feet. When grow- ing on ridges or walls it is found on slopes which have an exposure. toward the south. 1941] HAINES: YUCCA WHIPPLEI 35 GrowTH-ForMs It soon became apparent from field observation that at least four types of growth-forms exist in this species and that these forms are correlated with the geographic distribution. These growth-forms may be briefly characterized as follows: (1) a soli- Fic. 2. Yucca Whipplei. <A, a colony, perhaps a single clone, of subsp. percursa, San Rafael Mountains. B, a single plant from such a colony showing a short newly developed rhizome. C, an individual of subsp. intermedia, Santa Monica Mountains. The branch on the left has died back after flowering, that on the right has developed from an axillary bud on the dead branch. D, a small specimen of subsp. caespitosa from Walker Pass showing the method of branch- ing from the original stem. Such proliferation may continue before flowering takes place until a clump of a hundred or more rosettes is formed. tary form, with a simple unbranched, bulbous stem which dies as soon as the plant has flowered; (2) a caespitose form in which the stem branches on the surface of the ground to form a clump of plants composed of from four to one hundred or more distinct rosettes, several of which may send up flower stalks in the same season; in this case, numerous rosettes are formed before any flowering takes place (fig. 2, D); (3) a caespitose form, occasion- ally solitary, in which the stem branches by means of axillary buds which form short lateral branches; in this case new rosettes are usually formed only after flowering and the dying back of a 36 MADRONO [Vol. 6 branch; one or several rosettes may thus be formed, but only one flower stalk is usually produced in any given season (fig. 2, C) ; and (4) a rhizomatous form, the stem branching by means of underground rhizomes to form dense colonies of asexually pro- duced individuals (fig. 2, A, B). The solitary form (1) occurs in San Diego, Riverside, Orange, San Bernardino, and Los Angeles counties in two regions which are geographically isolated. They are indicated in Figure 1 by the letters “T” and “G.” Actually each of these regions is occupied by a distinct form which may be segregated on the basis of size. The caespitose form (2) with many rosettes, of which several flower in each season, occurs on the western margin of the Mojave Desert from the San Bernardino Mountains east and north to Tehachapi Pass and from there into the Piute and Greenhorn mountains. The distribution of this form is indicated in Figure 1 by the letter “C.” The caespitose form with few rosettes (3), only one flowering in each season, occurs throughout the Santa Monica and Santa Susana mountains in the area marked “I.” The rhizomatous form (4) occurs in the San Rafael, Santa Ynez and Santa Lucia mountains; its distribution is indicated in Figure 1 by “P”,. The only point at which any of these races are known to merge and exhibit intermediates lies in the Cajon Pass area where hybridization is apparently frequent between the solitary mon- tane form and the caespitose desert form. There are also two isolated regions in the southern Sierra in which Yucca Whipplei grows: along the western border of Sequoia National Park and on the Middle Fork of the Tule River in the Sierra Nevada foothills. The data regarding these two localities are incomplete and the plants growing there (as well as those of Lower California) have been eliminated from present consideration. It seems probable, that although the proportions are similar to those of the solitary form, they are distinct and form another subspecies because of the nature of branching. VARIATION OF INFLORESCENCE AND LEAVES After discovering that the members of this species possessed four forms of growth habit, the question arose as to whether the morphology of the inflorescence and proportions of the whole plant might not be correlated with the growth habit and geographic distribution. The answer to this question is based upon a statis- tical treatment of the measurements taken at each field station. The following characters were recorded for each of 1114 plants studied in the field (certain other characters were also recorded, but these were found not significant for the present purposes) : (1) the height of the flower stalk, (2) the length of the panicle, (3) diameter of the panicle at its widest point, (4) the diameter of the flower stalk as measured one meter above the surface of the ground, and (5) the length of the leaves. The measurement of characters (1) and (2) were made to the nearest six inches, 1941 | HAINES: YUCCA WHIPPLEI 37 while character (3) was measured to the nearest inch. Ata point one meter above the ground level, the diameter of the flower stalk (4) was measured with outside calipers to the nearest eighth of an inch. The length of the leaf (5) was measured to the nearest inch (fig. 3, C). 3 it —— B Cc E F Fic. 3. Ideographs which summarize the measurements made upon twenty plants of Yucca Whipplei from ten stations. Each pair of stations is from the same geographic region and was so chosen as to illustrate the extremes of variation measured in that region. A, subsp. caespitosa; B, subsp. percursa; D, subsp. Parishii; E, subsp. intermedia; F, subsp. typica. The dimensions mea- sured are shown in C. The stations follow: A, left, station 43, eight miles north- west of Mojave, Kern County; A, right, station 47, 1 mile above Valyermo; B, left, station 16, Santa Lucia Memorial Camp, Monterey County; B, right, sta- tion 51, 5 miles north of Los Olivos, Santa Barbara County; D, left, Arrowhead Springs, San Bernardino County; D, right, station 28, Tujunga Wash, Los Angeles County; E, left, station 26, Malibu Lake, Los Angeles County; E, right, station 15, West Los Angeles; F, left, station 35, 2 miles south of Lake Hen- as San Diego County; F, right, station 21, 10 miles east of Hemet, Riverside ounty. 38 MADRONO [Vol. 6 TABLE 1 Arithmetic Means for Measurements of Morphological Characters of Yucca Whipplet from Each Field Station, Showing the Total Range of Variation in the Species Height : F Field Number of eee pci ee eerie Length Station of Flower me oe e ower | of Leaves Number | Plants || Stalk | Pomicle)) Banicle yy) Stal me) iarimenee : in Feet in Inches Inches in Feet 4 16 8.47 3.53 13.80 1.23 6 14 13.90 6.46 21.70 3.21 fi 12 12.40 5.58 34.20 3.16 8 12 13.00 6.33 25.10 3.02 9 17 7.35 2.70 11.10 1.83 10 19 9.65 4.02 11.70 2.59 12 12 9.07 4.17 13.30 1.96 13 12 8.71 3.92 14.90 1.87 14 12 12.67 6.20 12.67 3.02 15 48 10.45 3.65 17.35 1.42 16 24 11.45 5.95 21.60 2.70 17 24 10.30 4.94 16.60 1.79 18 36 7.65 3.54 12.10 1.88 19 16 9.80 4.15 19.00 3.42 20 12 8.70 3.30 14.50 2.33 21 12 10.00 4.30 14.75 2.26 22 25 9.20 3.83 14.80 3.23 23 12 9.65 4.00 16.20 3.21 24 12 8.76 4.16 17.40 3.16 25 48 7.15 2.98 13.40 1.00 26 52 8.60 Si, 14.10 1.10 27 12 17.40 9.33 33.90 4.71 28 12 16.36 8.96 32.40 4.20 29 i%e) 9.93 3.87 17.30 1.42 30 36 9.13 3.88 14.80 1.31 31 36 9.70 5.03 16.03 2.44 32 13 10.20 3.96 16.05 1.34 33 12 7.68 2.72 9.35 1.39 34 12 8.90 3.22 12.60 1.43 35 24 6.90 2.87 11.10 1.46 36 12 6.92 3.08 12.67 1.58 37 12 9.00 4.21 16.40 2.81 38 12 6.50 2.67 10.20 1.28 39 12 6.45 2.46 11.20 1.23 40 24 8.70 3.50 15.50 1.22 41 26 6.60 2.54 10.00 1.00 42 24 9.05 3.62 14.20 1.44 43 36 6.54 2.56 9.75 1.26 4A 12 6.50 2.85 9.60 1.00 45 12 8.75 3.62 15.30 1.25 46 12 9.17 4.20 17.85 1.36 47 25 8.42 3.58 16.95 1.23 48 25 5.90 2.78 9.16 .86 49 36 7.90 3.30 10.80 1.57 50 13 8.48 3.38 12.10 1.56 51 25 9.18 4,42 15.90 2.09 52 25 8.88 4.44 16.16 1.89 53 50 7.91 3.61 13.50 1.62 54 Le 9.60 4.26 15.50 1.88 76 48 15.10 7.86 23.90 3.64 1941 | HAINES: YUCCA WHIPPLEI 39 In Table 1 are listed the field stations whose locations are indicated by number in Figure 1 together with the number of plants measured at each station and the average value of each of TABLE 2 Arithmetic Mean, Standard Deviation and Standard Error of the Mean for the Characters Measured and Segregated by Geographic Regions and Growth-Forms Yucca Height of Length Diameter piers Length Whipple Flower of of Mower of Subspecies Stalk Panicle Panicle Stalk Leaves a caespitosa, C 7.58 + 102 | 3.27+.587 | 12.06+.247 | 131+ .028 | 23.13 + 340 typica, T 8.32 + .103 | 3.144.060 | 13.30+.240 | 2.14+.050 | 23.93 + .330 Parishii, G 13.16 + 369 | 4.40+.253 | 23.46+.942 | 3.35+.109 | 28.98 + .637 intermedia, I 9.27+ 144 | 3.62 + .662 | 15.42+ .275 | 1.22+.162 | 26.68 + 305 percursa, P 9.33 + .170 | 4.62 + .813 | 16.90 + .292 | 1.99 + 392 | 18.87 + 349 these five characters. The data given in Table 1 show the mean measurements for each station but not the range; that is, the differences between two plants which might be found growing side by side. The ideographs shown in Figure 3 will give an approxi- mate idea of this range, that is, the amount of variation at a given station. The proportions of the individuals, as determined by the measurements indicated, are reconstructed in the ideographs, a method suggested by Edgar Anderson (Ann. Mo. Bot. Gard. 15: 241-832. 1938). Each of the ideographs in Figure 3 represents an individual plant from one of the five geographic regions indi- eated in Figure 1. The left pair of ideographs in each figure represents the largest and smallest plants found at one station, the right pair of ideographs in each figure, the largest and smallest plants found at a second, but widely separated, station within the same geographic region. ‘These have been selected in such a way as to illustrate the extremes of variation found in each of the five geographical regions. The arithmetic mean, the standard deviation wus standard error of the mean were pompuccd for each of the five characters of all the plants measured in each geographical region. These data are given in Table 2. From the arithmetic means ideo- graphs were constructed which represent the mean proportions of the inflorescence and leaves of the yuccas of each of the five geographic regions. These are shown in Figure 4, where “G” represents the solitary form growing in the San Bernardino and San Gabriel mountains; “P” represents the rhizomatous form growing in the San Rafael, Santa Ynez and Santa Lucia moun- tains; “I’’ represents the caespitose form growing in the Santa Monica and Santa Susana mountains; “‘T” represents the solitary form growing in San Diego, Riverside and Orange counties; and 40 MADRONO [ Vol. 6 “C” represents the caespitose form growing on the western margin of the Mojave Desert. These ideographs indicate that there is a recognizable differ- ence in the mean morphology of the inflorescence and propor- tions of the vegetative parts which is correlated with the four forms of growth habit of the stem. This fact is consonant with and gives a quantitative expression of facts which had already been recognized intu.tively from more or less casual observation. The differences between the two solitary forms “G” and “T”’ are so marked that the author proposes to separate them purely on the basis of the size and proportion of the inflorescence, par- ticularly since the two forms are geographically segregated. In order to determine whether the differences presented in Table 2 and Figure 4, are statistically significant, the standard att Fic. 4. Ideographs representing the mean proportions of five regional subspecies of Yucca Whipplei: G, subsp. Parishii; P, subsp. percursa; I, subsp. intermedia; T, subsp. typica; C, subsp. caespitosa. G error of difference was computed for the one hundred possible combinations of comparable characters represented in the ideo- graphs. The resultant differences are recorded in Table 3, and expressed as critical ratios. When the critical ratio has a value greater than three, it is assumed that the differences between the means are statistically significant and that the characters measured may safely be considered as having arisen from sepa- 1941 | HAINES: YUCCA WHIPPLEI AL rate populations in which the true values for these characters are not alike. TABLE 3 Critical Ratios Derived from the Standard Error of Difference between One Hundred Pairs of Means, as Shown typica Parishii intermedia percursa caespitosa So hes, 9.83 6.20 5.05 5.10 ‘S =-1.Pan. 4.83 0.72* 1.81* 0.22* = d.Pan. 10.45 5.80 9.55 3.58 & 4.F.S. 10.10 5.A5 0.38* 2.92 L.Lvs. 7.05 6.13 10.40 1.69* h.F.S. 10.10 11.90 14.60 S 1.Pan. 1.10* 0.25* Vere: ‘& d.Pan. 8.20 6.66 11.70 P dE.S. 10.90 3.34 182.00 L.Lvs. 3.26 10.11 9.05 & hF.S. 0.29* 11.05 3 = 1.Pan. 0.74* 0.39* S d.Pan. 3.70 9.10 one der’ S, 1.82* 0.55* (am livs: 16.80 11.00 = RSE 8.83 = 1.Pan. 0.77* 5 d.Pan. 12.60 Bae dE Ss: 1.73* ela vs. 8.75 * Critical ratios which do not indicate a significant difference between the two means compared. h.F.S.—height of flower stalk. 1.Pan.—length of panicle. d.Pan.—diameter of panicle. d.F.S.—diameter of flower stalk. l.Lvs.—length of leaves. While these ratios are statistically significant, their tax- onomic significance must be interpreted largely upon the basis of experience. These critical ratios suggest that the order of significance of the morphological characters are: the diameter of the panicle, the height of the flower stalk, the length of the leaves, the diameter of the flower stalk, and the length of the panicle. While some of the characters compared in Table 3 are not significantly different, the summation of habit which they express is. It is therefore clear that each geographic race or growth- form may be distinguished solely by mean size and proportions of its inflorescence and leaves. CONCLUSION It has been shown above that Yucca Whipplei is represented throughout its range by four growth-forms and that significant 42 MADRONO [Vol. 6 differences of inflorescence and leaves are correlated with these growth-forms. Furthermore, it has been shown that one growth- form, the solitary type, is characteristic of two forms which differ significantly in habit of inflorescence and foliage and are geo- graphically segregated. It appears, therefore, that Yucca Whipplet is represented by not less than five geographic races. That others may exist seems possible, in so far as the author is able to judge from the limited number of observations which were made on the forms of Lower California and the Sierra Nevada. The question arises, then, as to a suitable taxonomic treatment of these forms. It seems preferable, until there is evidence that more than geographic barriers segregate them, to treat the geographic races, discussed above, as subspecies. TAXONOMY Yucca WuippLer Torrey, Botany of the Mexican Boundary 22, 1859, type collected by Schott at San Pasqual, San Diego County, California (Torrey Herb., N. Y. Bot. Gard.). Y. aloifolia Torrey in Report Whipple Exped., Botany 147, 1856 (not L.). Type collected in Cajon Pass by the Whipple Expedition; since two subspecies with intermediate hybrids occur here it is’ uncertain to which form this reference applies. Hesperoyucca Whipple Baker in Kew Bull. p. 8, 1892. Nearly acaulescent, stem simple or branched; leaves in a basal rosette, 0.125—0.75 inches wide, 1—4 feet long, linear-lance- olate, tipped with a slender spine, fibrous, evergreen, flat to more or less triquetrous, finely striate, their margins serrulate; inflo- rescence 5—20 feet high; panicle oblong, 1.5—-11.5 feet long, 0.5—3 feet in diameter; peduncle 0.5—6 inches thick; perianth segments distinct, nearly equal; filaments clavate, attached to the base of the perianth; capsule short-cylindric, 1.5—2 inches long. KEY TO THE SUBSPECIES Stem branched, only the flowering branch dying back after flowering, both dead and living rosettes usually present. Stem branching by means of rhizomes, forming dense colonies of asexually produced individuals ......... a. subsp. percursa Stem branching on surface of ground, forming a clump of from 4 to 100 or more rosettes; several inflores- ences may arise from a clump of rosettes in a single SEASON aio uhie eye cata ee sce ie pet Ue ae = eae AO a ee b. subsp. caespitosa Stem branching by axillary buds to form short lateral branches; only one inflorescence arises from a group of. rosettes: in <a :Single Season...) ...2.. 0 9. o as c. subsp. intermedia Stem unbranched, solitary, the whole plant dying after flowering. Flower stalk 5-13 feet in height, 1—4.5 inches in diame- ter; panicle 1.5—-7 feet in length, .5—2 feet in diame- ter; San Diego, Riverside and Orange counties and northern: Lower ‘California, 4.2.29), 4... ee ee d. subsp. typica Flower stalk 7-20 feet in height, 1.5-6 inches in diame- ter; panicle 3-11.5 feet in length, 1-3 feet in diame- ter; San Gabriel and San Bernardino mountains ... e. subsp. Parishit 1941] HAINES: YUCCA WHIPPLEI 43 a. Yucca WHIPPLEI subsp. percursa subsp. nov. Type col- lected on Cachuma Mountain in the San Rafael Mountains of Santa Barbara County, California, A. L. Haines (deposited in the Herbarium, University of California, Los Angeles). Caudex longitudine 30-60 cm., in basi rhizomata elongata ferens et ut videtur clonum diffusum faciens; rhizomatibus maturis diametro circiter 2-3 cm., longitudine ad 2 m. et ultra; scapo altitudine ad 5 m. maximam partem circiter 3 m.; scapi in altitudine 1 m. supra humum diametro ad 8 cm., maximam partem 5 cm.; paniculis longitudine ad 3 cm., maximam partem circiter 1.5 m. et diametro in latissima parte ad 70 cm., maximam partem circiter 40 cm.; foliorum longitudine ad 1 m., maximam partem circiter 45 cm. Stem 1 to 2 feet long, producing rhizomes at its lower end; mature rhizomes 1 inch in diameter, 2—6 or more feet in length producing new individuals from the terminal buds to form a dense stand which may be considered as a clone; height of flower stalk 5-16 feet, average 9.3 feet; diameter of flower stalk at one meter above ground .75—3.5 inches, average 1.9 inches; length of panicle, 2—9.5 feet, average 4.6 feet; diameter of panicle at widest point, 6—28 inches, average, 17 inches; length of leaves, 10—35 inches, average 19 inches. This subspecies is a component of both the coastal sage and chaparral formations, occurring mostly between 200 and 2000 feet; it is found in the San Rafael, Santa Ynez and Santa Lucia mountains of Santa Barbara and Monterey counties. b. Yucca WuippLer subsp. caespitosa comb. nov. Y. Whipplei var. caespitosa Jones, Contrib. West. Bot. 15: 59. 1929, apparently based upon plants observed on the desert side of Cajon Pass. Stem branching above the ground to form a crowded, caespi- tose clump of four to a hundred or more rosettes, several branches in a single clump often flowering at one time; height of flower stalk 4-13 feet, average 7.5 feet; diameter of flower stalk one meter above the ground, .5—2.75 inches, average 1.3 inches; length of panicle, 1.5—6 feet, average 3.25 feet; diameter of panicle at widest point 4-24 inches, average 12 inches; length of leaves 12—40 inches, average 23 inches. This subspecies is an associate of the desert woodland, occur- ring particularly with Juniperus californica, usually between 2000 and 4000 feet. It may extend down into the upper limits of the desert shrub formation. It ranges north and west from Arrastre Creek in San Bernardino County along the western border of the Mojave Desert to the region of Walker Pass. ec. Yucca WuipPLe! subsp. intermedia subsp. nov. Type col- lected at Malibu Lake, Santa Monica Mountains, A. L. Haines (deposited in the Herbarium, University of California, Los Angeles). 4A MADRONO © [Vol. 6 Caudex a gemmis axillaribus pauciramosus in maturitate solum; scapo florifero saepius solitario altitudine ad 5 m. et ultra, maximam partem circiter 3 m. et diametro in altitudine 1 m. supra humum ad 5 cm., maximam partem circiter 3 cm.; paniculae longitudine ad 2.5 m., maximam partem circiter 1 m. et diametro in latissima parte ad 60 cm., maximam partem circiter 38 cm.; foliorum longitudine ad 1 m. et ultra, maximam partem circiter 60 cm. Stem branching only by axillary buds after it is well matured; base of stem and roots persistent after flowering to form the subterranean portion of the new plants; only one flower stalk produced in a cluster of rosettes at one time; height of flower stalk 5-15.5 feet, average 9.25 feet; diameter of flower stalk, one meter above the surface of the ground, .5—2 inches, average 1.2 inches; length of panicle .5—7.5 feet, average 3.5 feet; diam- eter of panicle 6—28 inches, average 15 inches; length of leaves 13—40 inches, average 26 inches. The type of branching of this subspecies seems to be inter- mediate between that of subsp. typica and subsp. caespitosa. It is found throughout the Santa Monica and Santa Susana moun- tains of Los Angeles and Ventura counties and occurs from sea level up to 2000 feet. It is an associate of the coastal sage and chaparral formations, particularly the chamisal of the latter. d. Yucca Wurppte1 subsp. typica nom. nov. Y. Whipplei Torr. Bot. Mex. Bound. 22. 1859. Stem simple, solitary, bulbous, dying when fruit has been formed; height of flower stalk 5-13 feet, average 8 feet; diameter of flower stalk, at height of one meter above the ground, 1—4.5 inches, average 2 inches; length of panicle 1.5—7 feet, average 3 feet; diameter of panicle at widest point 6-27 inches, average 13 inches; length of leaves 13—45 inches, average 24 inches. This subspecies has a stout inflorescence with a thick, short flower stalk bearing a heavy panicle of compact flowers. It occurs from 1000 to 4000 feet and ranges through the Laguna, Balkan, San Jacinto, and Santa Ana mountains. It is associated chiefly with the chaparral formation, particularly the chamisal, but occasionally extends into the coastal sage and desert shrub formations. e. Yucca Wuippxer subsp. Parishii comb. nov. Y. Whipplei var. Parishii Jones, Contrib. West. Bot. 15: 59. 1929; type col- lected above Cajon Pass near San Bernardino (?). Y. graminifolia Wood, Proc. Acad. Philad. 20: 167. 1868, based upon plants observed in “‘the mountains twelve miles east of Los Angeles,” perhaps near Monrovia. Stem simple, solitary, dying when fruit has been formed; height of flower stalk 7—20.5 feet, average 13 feet; diameter of 1941] BEETLE: SCIRPUS 45 flower stalk, at height of one meter above the ground, 1.5—6 inches, average 3.3 inches; length of panicle 3—11.5 feet, average 6.5 feet; diameter of panicle 1—3 feet, average 2 feet; length of leaves 17—46 inches, average 29 inches. The largest individuals of Yucca Whipplet occur in this sub- species. It is distinguishable from the other solitary form of the species by the size of its inflorescence. In all dimensions measured the inflorescence of subsp. Parishit is nearly twice as large as the inflorescence of subsp. typica. ‘The subspecies is found on the western slopes of the San Bernardino and San Gabriel mountains where it occurs from 1000 to 8000 feet ele- vation. At lower elevations it occurs with the chaparral and coastal sage formations, at higher elevations it ranges well into the montane forest. Department of Botany, University of California, Los Angeles, September 26, 1940. CERTAIN NORTH AND SOUTH AMERICAN DISTRIBUTIONS IN SCIRPUS ALAN A. BEETLE Recent studies on collections of the cyperaceous genus Scirpus L. obtained in South America by the University of California Botanical Expedition to the Andes has revealed a surprisingly high degree of relationship between the species of this genus in the two hemispheres. Although one species is endemic to the California flora a large number of the species is wide ranging making it necessary to study both near and distant floras to dis- cover their true affinities. The genus has many American species. Nothing approaching a general statement may as yet be given for their distribution. Some range widely, whereas others are very narrowly endemic or occupy discontinuous ranges. The California flora contains sixteen species of Scirpus as fol- lows: S. microcarpus Presl, S. Congdoni Britton, S. criniger Gray, S. paludosus A. Nels., S. robustus Pursh, S. fluviatilis (Torr.) Gray, S. acutus Muhl., S. validus L., 8. californicus (C. A. Mey.) Steud., S. Olneyi Gray, S. americanus Pers., S. nevadensis Wats., S. Clementis Jones, S. carinatus Gray, S. setaceus L. and S. cernuus Vahl. The species known from Argentina were recently well organized by Barros (1). Without following Barros too closely the following entities may be recognized: S. deserticola Phil., S. giganteus Kunth, S. paludosus, S. asper Presl, S. cubensis Poepp. & Kunth, S. cali- fornicus, S. validus, S. Olneyi, S. americanus, S. nevadensis, S. ataca- mensis (Phil.) Boeckl., S. rigidus Boeckl., S. macrolepis Boeckl., S. cernuus and S. inundatis Poir. Scirpus Clementis, S. criniger and S. Congdoni are endemic in the North American flora while S. ataca- mensis, S. rigidus, S. macrolepis, S. giganteus and S. deserticola are 46 MADRONO [Vol. 6 endemic in South America. Nearly fifty per cent of the species comprising these two lists occur both in California and in Argen- tina. An analysis of the distribution of those species which occur: both north and south of the equator should afford some informa- tion concerning the interchange of species in general between the hemispheres. In Scirpus, the section Baeothryon consists of perennials with spikelets uniformly single, terminal, style 3-fid, and achene smooth, brown. It comprises eight species and one variety, namely S. hudsonianus (Michx.) Fern., S. planifolius Muhl., S. Clin- toni Gray, S. rigidus, S. pumilus Vahl, S. Clementis, S. cespitosus L.,. S. cespitosus var. callosus and S. atacamensis. Three members of this group have a circumboreal distribution, and the fact that this. is an old group is attested by the obvious relationship of two Andean endemics to these northern types. Both Scirpus cespitosus var. callosus and S. hudsonianus are found in North America, Europe and Asia. Scirpus pumilus occurs in a few isolated stations in North America, in Europe and has been reported from Tibet (5). The end points of migration down the Rocky Mountains, through the montane regions of Central America and into the Andes, are represented apparently by S. rigidus of Peru and Bolivia and S. atacamensis of Chile and Argentina. That Scirpus planifolius and S. Clintonii are most closely related, within the section Baeothryon, to S. hudsonianus is shown by their mutual possession of cauline leaves and scabrous triangular culms, and is also strongly indicated by the roughly equiformal areas of their North American distribution. On the same morphological grounds the South American S. rigidus is more closely related to these three and less closely allied to its geographical partner S. atacamensis. ‘The last species has smooth, terete culms and cauline leaves reduced to mucronate sheaths, conditions which demon- strate its affinity with S. Clementis and S. pumilus. Acceptance of the latter assumption makes it necessary to postulate one of two possibilities as to their geographic origin: (a) a double migration down the Andes. from North America; (b) a continuous range from North to South America for some prototype which has diverged along parallel lines of evolution subsequent to the geo- graphic isolation of some of its parts. In either case the history of the plants undoubtedly dates back to the Pliocene when, coinci- dent with the earliest invasion of South America by holarctic fauna (6, 9), one or more of the species may be assumed to have had its widest distribution. Fernald (4) has pointed out that the present fragmentary distribution of Scirpus pumilus is probably a direct result of Pleistocene glaciation. Various species in other sections of the genus show an inter- ruption of their ranges in the tropics. Scirpus nevadensis is found to be an uncommon plant in the western United States and Canada occupying strongly saline soils over a wide area. It is rare in 1941] BEETLE: SCIRPUS AT Argentina where it also appears in strongly saline habitats over a wide area. The species is, however, unreported between northern Argentina and Mono County, California. It may have been a coastal, salt marsh type or perhaps was formerly tolerant of fresh or only slightly saline water. Analogies exist today in S. palu- dosus, S. americanus, and S. acutus which grow on the sea margins, on the borders of inland fresh-running streams and lakes and in the saline soils of the Great Basin. The uplift of the Rocky Mountains and the Sierra Nevada initiated the changes which resulted in the disappearance of some of the inland lakes and in- creased the salinity of others. Gradually, in accompaniment with this change, S. nevadensis may have developed an adaptive toler- ance toward aridity. The southern members of the species may have passed through a similar history but the members be- tween were faced with a different problem; either they disap- peared or gave rise to such closely related species as S. Olneyi aun! S. americanus. Scirpus paludosus, S. validus and S. Olneyi are similar to S. nevadensis in the interruption of their ranges by a broad belt of the tropics including at least Central America and South America north of the Amazon but they differ in being much more abun- dant in North America than in South America. There are several possible explanations to account for the existence of this broad hiatus in distribution which is certainly not a reflection of incom- plete collections: (a) the species may be spreading from the northern to the southern hemisphere and have not yet had time to occupy all of their potential area; (b) these species may, per- haps, have been more abundant in the past but are becoming de- pleted in the southern hemisphere due to the impact of changing conditions; (c) greater competition in the southern hemisphere flora may possibly have hindered the spread of these species since they first appeared there; (d) since they are not especially toler- ant of tropical conditions a migration through that area may have been accomplished by a few of the more tolerant biotypes. Scirpus paludosus and S. validus are evidently plastic species to which the first alternative may be applicable but S. Olneyi appears to be a more rigid type and the other alternatives may have to be called into account. Scirpus cubensis which is common in Chile is reported also from Panama by Standley (10). Scirpus inundatus, known from as far south in Chile as the Valdivian Lakes region is also reported from Costa Rica by Clarke (3, p. 460). Scirpus cernuus, a ubiquitous species with a world wide distribution in temperate regions avoids the tropics. Scirpus californicus and S. americanus also have a disrupted range but differ from the preceding species in being widespread and fairly common in both hemispheres. They come nearest to bridging the tropical barrier. Scirpus californicus is known from the San Francisco Bay region south to Guatemala, and from Co- 48 MADRONO [Vol. 6 lombia, Ecuador and Brazil to middle latitudes in South America. To this is added S. californicus var. tereticulmis (Steud.) comb. nov. (S. tereticulmis Steud. Syn. Pl. Cyp. 85. 1855) extending the total range from central South America to the Straits of Magellan. A more uniform climate would undoubtedly amplify this broad latitudinal range. There is geological evidence (2) that uniform, moist climates existed at several different periods of Tertiary time. Scirpus americanus, finally, is known from Nova Scotia and Alberta south to Mexico and from Peru and Brazil to Patagonia. Morphologically similar plants grow indiscriminately in fresh and strongly saline waters. The extreme variability in this species is suggested by the number of taxonomic variations which have been recognized in it. Such a high degree of polymorphy would seem to mark it as a comparatively young species which is reaching the peak of its distribution at the present time. There has now been presented a graded series of geographical distributions from the endemics of section Baeothryon on the one extreme to the ubiquitous Scirpus americanus on the other. The most widely ranging species in the series are those with the great- est variation. This fact suggests that at the time when its number of biotypes and range of tolerance is greatest it is possible for a species to cross well-marked climatic barriers. Probably S. Cle- mentis and S. atacamensis together represent an old species which, with advancing age, depauperization of biotypes and wide sepa- ration, became modified in descent to slightly different end points. Since they occupy areas isolated from other members of their sec- tion this assumption seems more reasonable than to suppose that they never had a sufficient wealth of biotypes to create an over- lapping of their now discrete ranges; that is, chance dispersal or fortuitous origin depend on close relatives living in nearly ad- jacent territories. The species whose distributional pattern is intermediate in character between those of S. Clementis and S. americanus may be classed logically as intermediate in age if they no longer occupy their maximum range, or as young species if they have yet to reach a maximum range. The methods of dispersal of Scirpus are not yet completely analyzed. Most species are stoloniferous perennials which mi- grate easily either up or down stream along muddy river banks. Birds eat the fruits and may carry them for short distances, that is, from one drainage area to another, in their digestive tracts, or. in mud adhering to their feet. Undoubtedly important also is the buoyant nature of the fruits, although this is an extremely variable factor. Achenes of S. americanus are considered to be non-buoyant but are abundant in lake drift (7) while those of S. maritimus L. may float for a month (8, p. 239), not sinking until the pericarp decays. Finally the element of chance dispersal undoubtedly plays a considerable role over a long period of time. The preva- lent uniformity of the aquatic habitat must be largely responsible for the high degree of relationship over wide areas. 1941] COOKE: LIFE ZONES ON MOUNT SHASTA 49 A conclusion similar to that reached by Matthew (6) in con- nection with a study of the evolution of land vertebrates may be drawn. It would seem to be unnecessary to postulate any pro- found change in the existing distribution of land masses to account for the present distribution of Scirpus between the two hemi- spheres. To assume change in climate is probably unnecessary since species are apparently being interchanged between North and South America at the present time, but a more uniform climate would facilitate this process. Such changes as did occur in the past certainly have had their effect on the history of the species concerned, but, apparently, any theory involving cata- clysmic phenomena is not essential to account for the migrations of North and South American species of this genus. Department of Botany, University of California, Berkeley, March, 1940. LITERATURE CITED 1. Barros, M. Ciperaceas Argentineas. Anales Museo Arg. Ciencias Nat. 38: 133-172. 1935. . CuHamperiin, T. C. <A group of hypotheses bearing on climatic changes. Journ. Geol. 5: 653-683. 1897. . Crarke, C. B. The Cyperaceae of Costa Rica. Contrib. U. S. Nat. Herb. 10: 443-471. 1908. . Fernatp, M. L. Persistence of plants in unglaciated areas of boreal America. Mem. Acad. Arts & Sci. 15: 266-268. 1925. 5. Hooxer, J.D. The Flora of British India. 6: 654. 1894. 6. MarrHew, W. D. Climate and evolution. Annals N. Y. Acad. Sci. 24: 171-318. 1915. 7. McArer, W. L. Notes on drift, vegetable balls, and aquatic insects as a food product of inland waters. Ecology 6: 288-302. 1925. 8. Riptey, H. N. The dispersal of plants throughout the world. 1930. 9. Scorr, W. B. A history of land mammals in the western hemisphere. Rev. ed. 1937. 10. Sranpiey, P. C. Flora of the Panama Canal Zone. Contrib. U. S. Nat. Herb. 27: 90. 1928. =~ wo bd THE PROBLEM OF LIFE ZONES ON MOUNT SHASTA, CALIFORNIA WiLt1AmM BrincEe Cooke In 1898 a party led by Dr. C. Hart Merriam made a biological survey of Mount Shasta. In the report (6) published in 1899 Dr. Merriam defined the several life zones he found on the moun- tain on the basis of his earlier discussion of North American life and crop zones (5). The writer, after four years of study, has come to the conclusion that two of the three zones must be revised or clarified (or, as to one of them, completely eliminated) in order that the casual visitor and future worker may be less confused as to the boundaries of the upper zones. The present paper is an 50 MADRONO [Vol. 6 attempt to point out some of the problems encountered and the conclusions resulting from a study of these problems. The Upper Sonoran zone (in Merriam’s terminology) occurs within sight of the mountain in Shasta Valley to the north, in Sacramento Valley to the south, and in the valleys of the Great Basin areas to the east, for example, the areas found in the vicinity of the Modoc Lava Beds National Monument (1). On Mount Shasta this zone does not occur above 4000 feet, the contour line which has been chosen as the lower limit of the area covered by this study. The Transition zone is clearly differentiated on Mount Shasta. Where the base of the mountain extends down to 4000 feet this zone can be divided into two minor belts. The lower of these may be called the Chaparral Belt. The term refers here to a Transition zone chaparral, a broader interpretation than that of Cooper (2). Wherever the Transition zone forest has been dis- turbed on the mountain a dense growth of chaparral has taken its place and to a certain extent has invaded the remaining for- ested portions. Before the Shasta National Forest was estab- lished, great areas of timber were logged. Fire contributed to the further destruction of whatever forest might have been left so that the former forest composition must be reconstructed by study of the accounts of early travelers and from isolated trees and small groves throughout the area. The zone is dominated by Arcto- staphylos patula and Ceanothus velutinus. Additional shrubs in- clude: Ceanothus cordulatus, C. divaricatus, C. prostratus, Castanopsis sempervirens, C. chrysophylla, Purshia tridentata, Pachystima myrsi- nites and others listed in Table 1. Above the chaparral belt there is a narrow belt of forest, typical of the normal Sierran Transition zone except that on the northeast quarter of the mountain there occurs a large stand of Pinus contorta var. Murrayana which almost completely replaces the normal forest association. The normal forest, except where it is invaded by the chaparral, includes the following coniferous species: Pinus ponderosa, P. Lambertiana, P. monticola, P. attenuata, P. contorta var. Murrayana, Abies concolor, A. magnifica var. shas- tensis, Libocedrus decurrens and Pseudotsuga taxifolia. Various spe- cies of deciduous trees and shrubs which form lower stories in this forest are listed in Table 1. Above the Transition zone forest is the Canadian zone as inter- preted by Merriam. Except on the northeast side it is almost wholly composed of Abies magnifica var. shastensis which covers an altitudinal belt of from 500 to 1500 feet around the mountain. At the head of Panther Creek, and between the east side of Gray Butte and the west side of Mud Creek Canyon, and at one or two other small isolated stations, T'suga Mertensiana enters this forest. Locally there may be colonies of from a few hemlocks in the fir forest to a few firs in the hemlock forest. In the vicinity of upper 1941] COOKE: LIFE ZONES ON MOUNT SHASTA TABLE 1 51 Species of woody plants occurring in the Transition Chaparral (Ch), Transition Forest (T), Canadian (Ca) and Hudsonian (H) zones on Mount Shasta, compared with the Zone Indicators of Hall and Grinnell (3) for Upper Sonoran (US), Transition (T), Canadian (Ca) and Hudsonian (H) zones. Abies Abies magnifica var. shastensis concolor . Acer glabrum .. Minis) virigis Var. Sinvata ... 0... 6c ees ce ee ee es mAmelancmer alnifolia <2 .....5...0.0.600eecen enews Arctostaphylos nevadensis Arctostaphylos patula Castanopsis chrysophylla Castanopsis sempervirens Species a) len te) Sep <ol-te? eo 10658) ce: 06) Hen Neg) a Ne: [uy ey ce! (Ol 'o) 0) 10, ©, Jetse, io) re 6: 8 oie! <e? tet <eF Jw: 16; 10: Verce, ce! Se, 19 V9: jer (sce ‘6, Sr Rg Uae We epriel acta! Gey Jol Je% (stot ‘et ta, ce: le Weve: Iwi ouue erie) Jo “vi (sy Ve, sus) “ep eh (ey @) fe: 0) 6) 6," ‘a: 30%, e: 1,8; 26) fe) 26. @eanothus cordulatus ..............0.2.c20.0e0e. Ceanothus divaricatus er er de; 6" seq qantas! ral erwe: fe), iu Jet “en Sey ayesal fe! sw: ce: [o{)6: > net la}, -« Weanotnus PTOStTatus .....6. 6. cst ee eee ewe Ceanothus velutinus Cercocarpus ledifolius Chamaebatiaria millefolium Chrysothamnus Bloomeri var. angustatus Chrysothamnus nauseosus var. occidentalis Cornus californica Cornus Nuttallii Juniperus communis var. montana ............... Kalmia polifolia Libocedrus decurrens Lutkea pectinata eo ee we eiecewisugs. Berle? é2ue: ‘i 16 ‘ellie <e) cess: 16 40. fo 16. Se, Jer iial ie: te: <e" @* “en ley ‘es! BachySrima MyEFSIMItes . 22... edie eee ee ces et eee Phyllodoce empetriformis Pinus Pinus Pinus Pinus Pinus Pinus Pinus Populus tremuloides albicaulis attenuata contorta var. Murrayana Lambertiana monticola ponderosa e) fe) Ketel othe jeuuel sects ‘ey sence! ie lejiset om iwiMei "a: fe? <0! fe) ee: ‘ee, ee) ie) (6: (0 ai Met Well esled te isiten \eigie’ Je. 6: ie. isu ce: 0) 00-8: (8: 61 ie: tv: 8: (0% (@! (et “8” ‘6 et Je: e 78; joi elie. IW) e:isa' Ye Se) <a7 "10: (e. w ‘6 “a: “e060 Ie) 0) leslie’ ve jee. ‘6. i618 ponderosa var. Jeffreyl ....... 00.5, 000086- opus trichoCGarpa .. 1... . 2. eee ea idee epee eet Prunus demisa Prunus emarginata Prunus subcordata et Met e bere: «ee Ow eke, 0 ee el lee 10 © © (6 We 6 6) /0 (0 io, 6 0 eae iotr6) trey teh ce! ve) ist len 6: (ele) .@ ee) ‘e's, ‘6:"8" 6! eee! 6! 16: Ve seudotsu@a taxifOliay ... 6 kd ae Ge nee ecw umsmia tridentata, 020.4466. 04 3 hese st cece cae Ribes Ribes divaricatum Ribes Ribes Ribes cereum .. nevadense Roezlii .. viscosissimum Salix Scouleriana Sambucus caerulea Sambucus racemosa Tsuga Mertensiana Vaccinium caespitosum Vaccinium occidentale Bo 2 wifey ,e! ice) 0) Var se) (oy fo) 98, ee! (8. ‘ole. 16 el ce /0, ja: (fe: et 6: (6. e St geet de efne: cal Ser cet ye. sr \eo (vie) Lente, (on )@:\ie) 0:6. gi te! te. 0) <6: jee" “e! See" <8 mi iw Welke! fos (6 le! ‘wi (ei celle. \e: (ei 1e: 8) va. @. te ve'ue, ©) e106. 16 (6) er 16 sieite se) (od w: le; “euie) fe) se! ie) 10, .0,4le! 16) ‘0:10! 'p! 6: Se, ie: *e: a0 se) oy oe 8. 6 4) 02 Jee; 18! 81 0) isce"ce 8 16) (en'e Cen ol te! (mt ee. fe) ce: 10. ©) e' he wi ie SL cv. ot (6 .8iri8- sel, (eysel diel Ce) 2907.8; sai CoMciel ‘eile. 8! sa neh 8: -e) \0},\0, ‘e818: @, Position on Shasta ChTCa TCaH TCa CaH ChT TCaH ChTCa ChT ChTCaH Ch Ch ChT ChTCa CaH Ca ChTCaH Ch Zone Indicator Ab Ca Ca Ca T T Ca US a T 52 MADRONO [Vol. 6 TABLE 2 Species of herbs occurring in the Canadian (Ca), Hudsonian (H), and Sub- alpine (A) zones on Mount Shasta compared with zonal distribution of previous records (Arctic-Alpine, AA), and with the Hall and Grinnell Indicator lists. Position Reported Indicator Species on Shasta position of Zone Achillea lanulosa var. alpicola ............. H AA Carex BREWER. 208 Ai. cence tne rea CaHA AA AA Carexaphaeocephala «2... has) 7. cee ee ee A Dicentrawmnifloray <2 7. wise ee eae CaH AA Drapavbrewerli. a4 vcs voce a: eee ee wee HA AA AA Erigeron compositus var. trifidus .......... HA AA H Ful Searmana een 20h. ohne ae seeen cake HA AA AA TUNCUSRI Anya acre ks Se ee CaHA AA ewasiaepyemaca, eo. cn: ee ca eee er, eee H AA Wewisiastriphy ais 27 sere ee tees H AA Ca Oxy PiasGig ya. hn cane se AE ence CaHA AA AA Penstemon Menziesii var. Davidsonii ....... CaHA HAA AA Phacelia.dtigidare: Gecuk ..accakl eee ee oe HA AA Ode rImM lel Oe a ee wnks ia ete eee oe Lohan A Polemonium shastense’ 4.2% 4.0 occ ed os os HA AA FPolyconum> minimums 255.2) sckeee eee H Roly sonum, Parnyl <n. ie. siere ee H Polygonum sShastense ¢ .......0 602. eon ses CaHA HA H Potentilla flabelliiolia’: 2*32.o.- 8 ee ee H H Ca Potentilla;Sibwal dil?.4.%.. gan wqeeweree ans oon lal H AA Saxitracva lolmicet <:..0 65.0: ce ee eee ee HA AA AA Silene sWatsSOnit) 2.325 5s ee cee cote ote HA AA Squaw Valley Creek hemlock makes up nearly seventy-five per cent of the forest cover. In the middle and lower portions of the zone a few individuals of Pinus monticola may be scattered among the firs. However, along the west bank of Mud Creek Canyon this species is represented by scattered individuals from the upper limit of erect firs to the middle of the Transition zone forest. One of the interesting features of this zone is the forest on the northeast quarter where the Canadian zone may not exist at all, or where it may be considered as replacing the Transition zone forest. Here species of the Transition zone, including Pinus ponderosa (possibly the variety Jeffreyi) and Abies concolor mingle in a mixed forest with (a) species from the Hudsonian zone, in- cluding Pinus albicaulis and Juniperus communis var. montana; (b) species from the Canadian zone, including Abies magnifica var. shastensis and Pinus monticola; and (c) the anomalous species (at least on Mount Shasta) Pinus contorta var. Murrayana, Castanopsis sempervirens and Ribes cereum. At the borders of the Canadian zone are broad areas transi- tional to the Hudsonian zone above and the Transition zone below. The belt on the lower side includes Abies concolor, A. magnifica var. shastensis and Pinus ponderosa var. Jeffreyi. Inaswamp at Wagon Camp Vaccinium occidentale is dominant. The belt on the upper 1941] COOKE: LIFE ZONES ON MOUNT SHASTA 53 side includes semierect or dwarfed specimens of Pinus albicaulis, Abies magnifica var. shastensis and Tsuga Mertensiana. These transi- tional belts could be labeled with the combined names of their commoner species: ‘Mixed Fir-Jeffrey Pine belt’? and “Pine-Fir- Hemlock belt.’’ In addition to the coniferous trees which have been mentioned as components of the Canadian zone, a number of broadleaf trees and shrubs occur in the area. In the dry for- ests are Ribes cereum, R. viscosissimum, Cercocarpus ledifolius and Populus tremuloides; while in the meadows and spring areas occur Phyllodoce empetriformis, Kalmia polifolia, Lutkea pectinata and Alnus viridis var. sinuata in addition to others listed in Table 1. Arms of the previously mentioned chaparral extend into this zone. It is above the Canadian zone that difficulties are encountered. These difficulties lie in the interpretation of timberline, in the distinction between the Canadian and Hudsonian zones and in the presence or absence of an Arctic-Alpine zone. Merriam considered an Arctic-Alpine zone present on the mountain. He thought that timberline coincided with the high- est limits of Pinus albicaulis, even though at its highest limits all individuals are reduced to mats of from a few inches to a foot or more in height. With this interpretation, timberline is the upper limit of the Hudsonian zone, which is almost exclusively made up of, and is characterized by, Pinus albicaulis. Timberline, then, according to Merriam, divides the Arctic-Alpine from the Hud- sonian zone. Thus timberline extends up most ridges on the mountain to heights of from 9000 to 9500 feet, with occasional arms up to 10,000 feet. The basins formed by Post-Pleistocene glacial cirques, and by moraine materials at about 8000 to 8500 feet would have a much lower timberline, perhaps between 8000 and 8500 feet, depending on the more or less strict application of the term to isolated stands of pines on the faces of minor ridges (with barren gulleys between and below them) and to the pres- ence or absence of trees of any height on the flats below these ridges. The Arctic-Alpine zone, then, would include the areas on ridges on which a fair stand of dwarf pine occurs and which lie in the shade most of the day and thus are considerably cooler in climate. These cooler ridge slopes are on the south and east sides of the ridges and hold snow banks longer, at a lower eleva- tion, than other more exposed areas. In one place, at least, on the north side of Red Butte, there is a small timberline area with a growth of plants above it. This small community was included by Merriam in the Arctic-Alpine zone because of its more severe climate as indicated by an association of plants which includes: Polemonium shastense, Silene Watsonii, Oxyria digyna, Sazifraga Tolmiei and others. However, it was admitted that on top of the Butte, several hundred feet above this colony, there is a fairly large stand of Hudsonian zone timber. A number of similar ‘“Arctic-Alpine” associations occur below timberline as conceived 5A | MADRONO [Vol. 6 by Merriam, some of which grow in climatically restricted areas such as this, while others occur in apparently normal Hudsonian zone Climate. There are other conceptions of what constitutes timberline. We are referring here, of course, to alpine timberline rather than to sea-level poleward timberline, or to those desert or grassland associations on the borders of which tree species drop out in transi- tion belts. United States forest rangers of the Shasta National Forest consider timberline as the upper altitudinal limit of upright tree growth. This is an economic interpretation of the word. In this sense the line or belt between the Canadian and Hudsonian zones is timberline. According to Hopkins (4, 7): “Climatic timberline may be defined as the poleward sea-level or alpine alti- tude limit of upright growth of tree species, under otherwise favorable conditions, as distinguished from dwarf or prostrate forms of the same species, as controlled by the climatic elements of temperature, wind, snow, ice, etc.; and also as distinguished from limits of tree growth due to the absence of suitable soil, moisture, or the presence of local glaciers.”’ In the sense of Hop- kins, then, Merriam’s Shasta Hudsonian zone would occur above climatic timberline (which coincides with the economic timberline of the forest ranger). Proceeding to the problem of the Merriam Arctic-Alpine zone on Mount Shasta, it should be noted that the term “Arctic-Alpine”’ is misleading. Conditions of growth in the arctic circle are dif- ferent from those in an alpine habitat where the air pressure is lower, the proportion of daylight nearly the same as that normal to the latitude under consideration and temperature change dur- ing the growing season more frequent. Since these factors affect food production and other physiological processes of the vegeta- tion the Arctic zone should be differentiated from the Alpine. On Mount Shasta, then, the vegetation may be considered alpine rather than arctic and according to either Merriam or Hopkins the zone would occur above the limits of tree growth. This zone, therefore, if recognized on Mount Shasta, is characterized purely by herbaceous plants. The Alpine zone may be interpreted in terms of associations or an association of plants, in which sense Merriam occasionally used the term in discussing the flora of the mountain. On Mount Shasta a number of species reputedly occur (6) above tree-line. These include: Polemonium shastense, Oxyria digyna, Phacelia frig- ida, Silene Watsonu, Draba Breweri, Carex Breweri, Juncus Parryi, Dicentra uniflora, Saxifraga Tolmiei and others. However, all these species have been discovered below the upper limits of tree growth in various habitats such as draws, flats, gulleys and ridges which occur in Merriam’s Hudsonian zone; all these species have been observed in association with Pinus albicaulis. The only species yet known which occur above timberline as delimited by Merriam and 1941 ] COOKE: LIFE ZONES ON MOUNT SHASTA 55 not also below it are Carex phaeocephala and Poa Pringlet both of which were collected in 1939 along the Summit Trail in an area which Merriam would have called truly alpine since it lies at between 9000 and 10,000 feet, in the cirque in question fully 500 to 1000 feet above the highest limits of his interpretation of tim- berline. Merriam had no temperature records upon which to base his Shasta Alpine zone. He therefore based it upon the association of plants found in an area which should have certain theoretical temperatures. But is ‘this association truly alpine when all but two of its constituents (these two unreported until recently) are found in the Hudsonian zone? Hopkins defines the bioclimatic zones in three major zones and a number of minor zones. The three major zones include: I. “North and south, polar or arctic and alpine, humid or arid major regions of the Frigid Zone.” II. “North and South, tem- perate or intermediate latitudes or altitudes, major arid and humid regions of the Temperate Zone.” III. “Tropical and equa- torial low and high land, humid and arid major regions of the tropical zone.” Four groups of minor zones, of which group 4 is warmest, are described under Major zone I. Upper minor zone 4 is defined as “Arctic and alpine above and below perpetual ice and snow, lower transition or snow line zone.” Under Major zone II, Hopkins describes seven groups of minor zones of which group “1” pertains to the present problem. Under “Upper minor zone 1” the following definition is listed: “High latitudes and alti- tudes, subarctic and alpine regions and areas above climatic tim- berline or its equivalent, ‘timberline zone.’”’ “‘Lower minor zone 1” is “‘timberline.” Mount Shasta has no permanent snow line. The five glaciers which are still active are confined to areas above 10,000 feet and apparently have no immediate effect on the surrounding vegeta- tion. In their vicinity the soil is too poor and the drainage too complete for an association of plants to survive even though no glacier occurred. The permanent snow fields on the mountain are small and scattered and have little effect on the minimal vege- tation in their vicinity largely because they are above the areas where plant life, except lichens and algae, survives even sparsely. From these considerations, an Alpine zone on Mount Shasta is not to be expected. Certainly if permanent snow lines do not exist on the mountain, by definition, Major zone I is absent. The next zone is Major zone II, Upper minor 1. This is subalpine, and from areas of greater distribution of plants, might even be considered upper Hudsonian. However, the finer threads cannot be split until more definite statistics are available for the climate of the mountain, especially in its upper regions, throughout the year. Foster, Warren County, Ohio, December, 1939. 56 MADRONO [Vol. 6 LITERATURE CITED 1. AppLeGATE, EK. I. Plants of the Lava Beds National Monument, California. Am. Mid. Nat. 19: 334-368. 1938. 2. Cooper, W.S. The Broad-Sclerophyll Vegetation of California. Carnegie Inst. of Wash. Publ. 319. 1922. 3. Hatt, H. M. and J. Grinnett. Life Zone Indicators in California. Proc. Calif. Acad. Sci., Ser. 4, 9 (2): 37-67. 1919. 4. Hopxins, A. D. Bioclimatics, a Science of Life and Climate Relations. U.S. Dept. Agr., Misc. Publ. 280. 1938. 5. Merriam, C. H. Life Zones and Crop Zones of the United States. U. S. Dept. Agr., Bur. Biol. Surv. Bull. 10. 1898. 6. Merriam, C. H. Results of a Biological Survey of Mount Shasta, Northern California. U.S. Dept. Agr., Div. Biol. Surv., North American Fauna. No. 16. 1899. 7. Raper, O. Review of Bioclimatics. Ecology 20: 413. 1939. A NEW SPECIES OF LOTUS FROM THE MOUNT HAMILTON RANGE, CALIFORNIA HELEN K. SHARSMITH Lotus rubriflorus sp. nov. Herba annua albosericea, pilis longis; caules 1—5 aliquando decumbentes, 3-10 cm. longi; folia 10-16 mm. longa exstipulata, 4-foliolata imparipinnata, rachibus complanatis, foliolis lanceolatis acutis, 2-11 mm. longis; flores solitarii axillares subsessiles, 6—7 mm. longi; calyx dense pilosus lobis linearibus acuminatis, 4-5 mm. longis, corollam aequantibus; petala rubra in aetate caerulescentia; legumina pilosa, oblonga dehiscentia, 8—9 mm. longa. Slender annual; stems 1-5 (mostly 2-4) from base, 3-10 (mostly 4-8) em. long, usually somewhat decumbent; herbage pilose throughout with long, white, silky, somewhat appressed or spreading hairs; leaves 10-16 mm. long, exstipulate, imparipin- nate; rachis flattened, about 1 mm. broad; leaflets 4, two appear- ing terminal, the remaining two on one side of the rachis, 2—11 mm. long, subsessile, lanceolate, acute, terminal leaflets often slightly faleate; flowers 6—7 mm. long, solitary, axillary, sub- sessile, usually bracteate; calyx 6—7 mm. long, densely pilose; tube 1.5-2 mm. long; lobes linear, acuminate, 4-5 mm. long, equalling corolla; petals glabrous, clear bright pinkish red (aster purple’), fading bluish, claws white, banner 5 mm. long; wings -shorter than or about equalling keel, posterior dorsal lobes large and very narrowly margined with white; keel 5 mm. long, apex somewhat pointed, body very narrowly margined with white dor- sally and posteriorly ; ovary densely pilose, dorsal margin straight, ventral margin curved; style pilose below, glabrous above, curved near base; legumes pilose, 8-9 mm. long, 2.5 mm. wide, oblong, stramineous; seeds 2—4, 2 mm. long, 2 mm. wide, irregularly lens 1 Ridgway, R. Color standards and color nomenclature, plate 12. 1912. 1941] SHARSMITH: NEW SPECIES OF LOTUS 57 Fic. 1. Lotus rubriflorus H. K. Sharsmith: 7, flower; 2, leaf; 3, seed, lateral view; 4, seed, anterior view showing hilum; 4, ovary; 6, banner; 7, keel; 8, wing; 9, interior of calyx. All x5. shaped, angled, notched at hilum, olive green to light brown, usually mottled with deeper brown, the surface often with a bronze sheen. Type. Rolling hills at north end of Adobe Valley, Stanislaus County, Mount Hamilton Range, California, altitude 1600 feet, April 22, 1986, A. M. Carter & H. K. Sharsmith 3544, in flower (Herb. Univ. Calif. No. 605765; isotypes, Gray Herb., U.S. Nat. Herb., Mo. Bot. Gard., Stanford, Kew). Topotype. June 5, 1938, A. M. Carter §& J. L. Morrison 1408, in fruit (Herb. Univ. Calif.; duplicate topotypes deposited with isotypes). 58 MADRONO [ Vol. 6 Lotus rubriflorus has been found only at the type locality, but within the one known colony the plants are abundant. Collec- tions made two years apart suggest that the species is not merely transient at this locality. It inhabits sparse grassland in an open stand of Quercus Douglasii, growing in association with the com- mon vernal annuals of the region, such as Lotus subpinnatus, L. humistratus, Orthocarpus attenuatus, Microseris linearifolia, M. syl- vatica, Pentachaeta exilis, Plagiobothrys canescens. On the basis of the pinnate, short, exstipulate leaves, the small, solitary flowers, and the straight, beakless, dehiscent legumes, Lotus rubriflorus belongs to the subgenus Acmispon (Raf.) Ottley (Univ. Calif. Publ. Bot. 10: 197. 1923). Within this sub- genus, L. rubriflorus is most closely associated with L. denticulatus Greene, L. subpinnatus Lag., and L. humistratus Greene, these spe- cies all having dilated leaf rachises, few, inequilateral leaflets, sub- sessile flowers, and flattened, angled seeds. Although growing in association with Lotus humistratus and L. subpinnatus, L. rubriflorus is a very distinct, easily recognized spe- cies. It most closely resembles L. humistratus in pubescence, length of calyx lobes, and shape and size of legumes, but is more pilose, and the calyx lobes are longer. It differs strikingly from L. humistratus, as well as from L. denticulatus and L. subpinnatus, in the lanceolate leaflets. Furthermore, the flower color of Lotus rubriflorus is exceptional; in North American representatives of the genus the corolla is characteristically whitish or yellow, although often veined, streaked or suffused with red, pink, or purple, or may fade red. In L. rubriflorus the corolla is a uniform red. The following key brings out these distinguishing features and separates L. rubriflorus from those species to which it is most nearly related: Leaflets obovate, often obtuse; flowers cream color to yellow, ; tinged with or fading red or purple 2.4... 4¢20 4. 2240 ee L. denticulatus L. humistratus L. subpinnatus Leaflets lanceolate, acute; flowers bright pinkish red, fading |e} LUG TS) 0 per Mr ta ae mare ene Urn W a ected: MPN OI ee the ya aM Rich Ao tyueee L. rubrifiorus State College of Washington, Pullman, Washington, February, 1939. REVIEWS A Monograph of the Genus Calochortus. By Marion Ownsey. Annals of the Missouri Botanical Garden. Volume XXVIII. Pp. 371-560. St. Louis, December 10, 1940. The genus Calochortus, ranging from Nebraska to the Pacific and from Guatemala to British Columbia, is treated by Dr. Own- bey in three sections, Hucalochortus, Mariposa and Cyclobothra. 1941] REVIEWS 59 The sections are divided into twelve subsections under which are arranged the fifty-seven species and thirteen varieties. Dr. Ownbey’s approach to the problems in Calochortus from the viewpoint of herbarium study, extensive field work, garden cultures and cytological investigation gives his work a definitive- ness possessed by too few monographs. While some botanists may feel that he has admitted to specific rank several entities that might better have received subspecific status, on the whole his decisions seem to be based on a firm foundation of evidence both from comparative morphology and from geographic distribution. The publication of seven new species and ten new varieties should arouse keen interest in the further study of local areas. Of great significance in this connection is the speciation that has taken place in the Siskiyou Mountains of southern Oregon and northern California where four local, very distinct, rarely col- lected species are found. This area should continue to be a fertile collecting ground. Because of the cytogenetic approach and the fact that Dr. Ownbey is now working in the west where garden cultures and field studies may be carried on we await with great interest any additional developments in the phylogenetic analysis of this genus.—Jouwn L. Morrison, Department of Botany, University of California, Berkeley. The Flora of Whatcom County, State of Washington. By W. C. MueEnscHeER. Pp. 1-134, with 10 figures and 1 table. William A. Church Company, Ithaca, New York, 1941. Published by the author. A county flora is so rarely prepared for a portion of any of the western states that the appearance of a less excellent volume than the present one evokes especial comment. The twenty-three hundred square miles comprising Whatcom County occupy the northernmost county of the Pacific Coast and extend in altitude from near sea-level on Puget Sound to 10,750 feet at the summit of Mount Baker, one of the principal peaks of the Cascade Range. With the coastal hills in the Transition life zone, and timberline on Mount Baker at only 6000 feet, all of the boreal zones are thus represented in a relatively small altitudinal range as well as in a small area. With these facts in mind, one will be less astonished at the inclusion of 1042 species and varie- ties. The author contributed an earlier catalogue of the same _area to ““Muhlenbergia” in 1914, and has noted additions to this original list, chiefly in “‘Torreya’”’ and Maprono, during the subse- quent years. A substantial portion of the list is owing to the writer's predilection for aquatic and introduced plants, both of which are too likely to be overlooked by collectors. Although the plant records of the present book are based primarily upon Dr. Muenscher’s own collections, the first set of which is preserved 60 MADRONO [Vol. 6 in the herbarium of Cornell University, he has consulted the col- lections of others as well. The diversity of topics treated in the introduction reflects the author’s wide botanical interests. An interesting feature is the presentation of lists of those species characteristic of such dis- tinctive habitats as alpine meadows, salt marshes, gravelly prairies. and coastal cliffs. A brief account of the zonation of vegetation is handsomely illustrated by carefully selected photographs. A conveniently classified list is offered of plants deemed suitable for cultivation. : In the annotated catalogue, which constitutes two-thirds of the book, the genera and species are alphabetically arranged under the systematically ordered families. Neither keys nor descrip- tions are provided. The abundance and characteristic habitats of each species are described and authenticating specimens are cited for each. It thus becomes possible for the perennially skeptical specialist to check the identity of every item attributed to the group with which he is especially concerned. The taxonomic treatment is, on the whole, conservative, and pretty thoroughly up to date. The current bitter controversy between the advocates of “‘subspecies’” and the proponents of “varieties” is met by calling all subspecific entities, except forms, “varieties,” and retaining the original authors of the trinomials, regardless of whether they designated the subspecific or varietal category. This has the curious effect of attributing “varieties” to such authors as Hall and Clements, Piper, and Pennell, which will probably bring down the wrath of both factions. There is also some inconsistency in the capitalization of specific names, which will doubtless provide ammunition for the arsenal of those advocating uniform decapitalization. This flora is, happily, much more than an unusually complete and attractively prepared catalogue of the plants of one western county. Because of the wide range of habitats and altitudes con- tained in this one political division, it is essentially a catalogue of the flora of western Washington. It is to be hoped that the com- pletion of this very satisfactory study will encourage the prepara- tion of other much needed local floras throughout the west.—- L. Constance, Department of Botany, University of California, Berkeley. Experimental Studies on the Nature of Species. I. Effect of Varied Environments on Western North American Plants. By Jens: Cuiausen, Davin D. Keck anp WitiiaM M. Hirsey. Carnegie Insti- tution of Washington publication 520. Pp. vii+ 452. 1940. Paper, $3.50. Bound, $4.50. This stimulating volume represents the application of methods: in experimental taxonomy to an understanding of the involved intrarelationship of several complicated groups of species. The application of the method to specific problems serves in this case: 1941] REVIEWS 61 as a vehicle to outline the method of research. The groups of plants selected are Potentilla glandulosa and its allies, P. gracilis and its allies, P. Drummondiu and P. Breweri, Horkelia fusca and Hi. californica, the genus Zauschneria, certain groups of Penstemon, the Achillea millefolium complex, the Artemisia vulgaris complex and a large number of well selected minor problems in groups ranging from the Gramineae through the Compositae. The volume is prefaced with a short historical sketch of the research project; a brief tribute is paid to the late Dr. Harvey Monroe Hall under whose able direction the work was begun. It is a point of major interest that the authorship should include a cytogeneticist, a tax- onomist and a plant physiologist. In establishing the experimental method the problem was approached from many points of view and several methods were tried and discarded before the final procedure was adopted. The plant materials selected were those that could be used as clone transplants. A clone is defined by the authors as “‘all asexual propagules derived from one individual.” These were subjected to varied environments: cytogenetic studies were made: a revised taxonomic analysis was presented based upon the results of these experiments. The terminology follows closely that of Turesson. For purposes of review the treatment of Potentilla glandulosa and its allies (section Drymocallis) affords an illustration of the mode of attack. This group had received varied and haphazard treatment at the hands of the old school taxonomists. It was selected because of its widespread vertical and horizontal distri- bution, its great variability and its ready adaptability to garden manipulation. Specifically, California ecotypes from the follow- ing stations were used: Coast Range; Sierra Nevada, foothill, mid-altitude meadow, subalpine and alpine. These differed mor- phologically and showed some variability in seasonal rhythm. They possessed the uniform chromosome numbern=7. There are no genetic barriers between them. Clone transplants of each eco- type were established in California at three stations: a coastal station at Stanford University, a middle station at Mather, and a timberline station near Saddlebag Lake. The latter stations are in the Sierra Nevada. Observations were made to determine constant characters, seasonal differences, effects of light and moisture differences and effects of altitude. The reversibility of environmental modifica- tions was established for some ecotypes by taking clone members of changed individuals back to the original environment. In general it was found that each ecotype retained a certain individuality in all three stations. This was regarded as being due to the sum total of hereditary characters. Each responded to the different environments in a very characteristic way; these changes (modifications) were superimposed over the hereditary characters. While morphological change was slight the annual cycle or response to the rhythm of seasonal events was profoundly 62 MADRONO | [Vol. 6 changed. In some cases, however, this change was not sufficient to insure survival. Cytogenetic studies had as their objective “to interpret the taxonomic status of the various forms by testing the degrees of genetic relationship, to explore their evolutionary background by analysis of gene or chromosome differences and to determine their utility for various ecogenetic investigations.” In determining whether they were dealing with ecospecies or ecotypes the follow- ing criteria were adopted: Crosses between ecotypes would pro- duce a second generation in which the parental genes would be recombined in all proportions without resulting in constitutional weakness of the offspring. In crosses between ecospecies the genic recombinations would produce many constitutionally weak offspring which would barely survive under favorable conditions. Thus there would be constitutional weakness plus environmental unfitness in the offspring of the cross between ecospecies. It has been established in such species of the section Drymocallis as have been studied that the “diversity distinguishing ecotypes and eco- species has been produced by a simple genic differentiation.” Genic incompatability, they report, is slight. It was found that hybrids between the ecotypes were easily produced and fully fertile and the conclusion was drawn that they were true ecotypés of a large and variable ecospecies. The authors conclude that “Potentilla glandulosa presents a picture of evolutionary differenti- ation into ecologic units, the ecotypes, which are not separated from each other by breeding barriers. The ecotypes fit their spe- cific climates and differ by many characters, both physical and morphological. Each of these characters appears to be deter- mined by a series of genes, and is often genetically linked with others. The many differences in genes with minute effect furnish the basis for the multiplicity of intergrading forms observed in nature, and the linkage relations suggest a reason for the limited number of taxonomic units that can be recognized in spite of recombinations and lack of internal barriers.” As a result of many such experimental studies certain general conclusions are drawn relative to the problems of regional differ- entiation. These are discussed in the form of eleven laws gov- erning the nature of modification, its genetic limitations and environmental control and the bearing of modification and genetic differentiation on the development of regional races. Regional races are interpreted as ecotypes or ecospecies which have de- veloped in harmony with their environment through genic differ- entiation of one sort or another and are under the control of natural selection. Regional differentiation implies discrimination between unlike environments. These units, the ecotype and the ecospecies, are regarded as the natural units of ecological and evolutionary behavior. These conclusions are in close agreement with Turesson’s statements regarding the genotypical response of the plant species to its habitat. 1941] NOTES AND NEWS 63 The volume is replete with tables and graphs giving compara- tive measurements of the plant and its parts as it developed in the different environments. Photographic illustrations of plants and of specimen vouchers provide convincing demonstration of the results obtained. This work at once becomes a classic in the field of experimen- tal taxonomy. Regardless of agreement or disagreement as to precise limitation in the definition of species the problem of the pattern of relationship within closely allied groups is clearly set forth by such methods as these. Obviously it will be impossible to subject all plant groups to similar experimentation. It is hoped, however, that enough such work can be done to complete the picture of the nature of relationships in plants so that a pat- tern can be established into which such plants as are not adaptable to this mode of treatment can be placed.—HeErBeErt L. Mason. NOTES AND NEWS The following monographic studies and other important taxo- nomic articles have appeared recently: “preliminary list of des- mids of the Pacific Northwest with descriptions of some new forms,’ by Gordon D. Alcorn (Occas. Papers, Dept. Biol. Coll. Puget Sound, no. 10, 1-200, pls. 1-8. 1940); “‘the North Ameri- can sub-divisions of Ranunculus,’ by Lyman Benson (Amer. Journ. Bot. 27: 799-807, 17 figs. 1940); “a revision of Laurentia and allied genera in North America,” by Rogers McVaugh (Bull. Torr. Bot. Club 67: 778-798. 1940); “new combinations and new names in the Umbelliferae,’ by Mildred Mathias and Lincoln Constance (Bull. Torr. Bot. Club 68: 121-124. 1941); “‘a revi- sion of Choisya,”’ by Cornelius G. Muller (Amer. Midland Nat. 24: 729-742, 7 figs. 1940); “the American species of Hippocra- teaceae,” by A. C. Smith (Brittonia 3: 341-355, 12 figs. 1940); “a revision of the genus Chaenactis,’ by Palmer Stockwell (Con- trib. Dudley Herb. 3: 89-168, pls. 22-45. 1940). Recent articles of general interest to botanists are: “late Terti- ary floras of the Great Basin and border area,’ by Daniel I. Axel- rod (Bull. Torr. Bot. Club 67: 477-487. 1940); “some features of the structure of Toxicodendron diversilobum,’ Herbert F. Copeland and Bernice E. Doyel (Amer. Journ. Bot. 27: 932-939. 1940) ; “history of botanical exploration in Alaska and Yukon ter- ritories from the time of their discovery to 1940,” by Eric Hultén (Botaniska Notizer 1940: 289-846. 1940); “Arizona localities of interest to botanists,” compiled by T. H. Kearney (mimeographed, received 1941) ; ‘additional nomina generica conservanda: Pteri- dophyta and Phanerogamae,” by T. A. Sprague (Kew Bull. Misc. Inf. no. 3: 81-1384. 1940); ‘Gramineae of Nevada,” by Jason R. Swallen (Contributions toward a flora of Nevada, no. 1, mimeo- graphed, Washington, D. C.).—ErTner Crum. 64 MADRONO [Vol. 6 PROCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY October 24, 1940. Meeting, 2093 Life Sciences Building, University of California, Berkeley, at 7:45 P.M. The President, Dr. Ira L. Wiggins, occupied the chair. Dr. Wilfred W. Robbins, Professor of Botany and Botanist in the Experiment Station, Uni- versity of California, Davis, spoke on, “Alien Plants Growing without Cultivation in California.” November 14, 1940. Meeting, 2093 Life Sciences Building, University of California, Berkeley, at 7:45 P.M. The President, Dr. Ira L. Wiggins, occupied the chair. A nominating committee was appointed by the President, to submit nominations for officers of the Society for 1941, and asked to report at the December meeting. Mr. J. W. Stacey, of San Francisco, spoke on, “Distri- bution and Evolution of the Genus Carex in Western North America.”’ December 12, 1940. Meeting, 2093 Life Sciences Building, University of California, Berkeley, at 7:45 P.M. The President, Dr. Ira L. Wiggins, occupied the chair. Mr. William M. Hiesey, Carnegie Institution of Washington, Stanford University, Cali- fornia, spoke on, “Relations between Plants and Climates as Revealed by Transplant Experiments.” January 16, 1941. Meeting, 2093 Life Sciences Building, University of California, Berkeley, at 7:45 P.M. The President, Dr. Ira L. Wiggins, occupied the chair. Miss Ethel K. Crum, Chairman of the Nominating Committee, submitted the names of the following candidates: President, Professor E. B. Babcock; First Vice-President, Mrs. Roxana S. Ferris; Second Vice-Presi- dent, Dr. Palmer Stockwell; Secretary, Dr. Lincoln Constance ; Treasurer, Mr. William M. Hiesey. A motion, made and seconded to accept the report of the nominating committee, was passed unanimously. There were no nominations from the floor. Dr. G. Ledyard Stebbins, Jr., Associate Professor of Genetics, Uni- versity of California, Berkeley, spoke on, “The Genetic Approach to Problems of Rare and Endemic Species.” February 20, 1941. Meeting, 2093 Life Sciences Building, University of California, Berkeley, at 7:45 P. M. The President, Professor E. B. Babcock, occupied the chair. Dr. Ralph W. Chaney, Chairman of the Department of Paleontology, University of California, Berkeley, gave an illustrated lecture on, “Tropical America, a Botancial Garden of the Past.”—L. Constance, Secretary. MADRONO A West American Journal of Botany A quarterly journal devoted to important and stimulating articles dealing with plant morphology, physiology, taxonomy, and botanical history. These volumes should be a part of every botanist’s li- brary and should be made accessible to students of all universities and colleges. Volume I, 1916-1929. . . $5.00 Volume II, 1930-1934 .. 5.00 Volume III, 1935-1936 . 5.00 Volume IV, 1937-1938 . 5.00 Volume V, 1939-1940 . . 5.00 Single numbers....... 0.75 The subscription price of MADRONO is $2.50 per year. We solicit your pat- ronage, Address all orders to: William Hiesey Bus. Mgr. Carnegie Institution of Washington Stanford University, California Bh aes i) VOLUME VI NUMBER 3 Ii MADRONO | A WEST AMERICAN JOURNAL OF | BOTANY | ALPINE Fora or SAN Francisco Mountain, Arizona, Elbert L. Little, Jr. 65 Mane Hisk SPENCER, Mrederick Grover .. 00.0... 6 dee ee ee 82 New Species oF VascuLaR PLANTs FROM THE NortrHwest Coast, George OPV ORESIL UY OR Ci eis CN GOR UM EO Rae Tra i Ma a te 84 | NotTewortHy Piants From [paHo, Arthur Cronquist .................... 87 ALASKA CEDAR IN CALIForRNIA, Herbert L. Mason .................0.00 0055 90 Reviews: R. M. Tryon, N. C. Fassett, D. W. Dunlop and M. E. Diemer, The Ferns and Fern Allies of Wisconsin (Herbert L. Mason); Harold E. 3 Contents | | and Virginia Long Bailey, Forests and Trees of the Western National ii Parks (Ethel Crum); Morton E. Peck, 4 Manual of the Higher Plants . ofOregon (Lincoln Constance) 2...) 2.0050 026022 deen 90 Notts ann News: Field Work of the Bureau of Plant Industry in Nevada His: ¢ (W. Andrew Archer); Three Alien Plants New to Oregon (Lincoln | LTS SGT 5) ) A NESTS RMI NAZIS AC AER aE OE IG 94 PROCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY ...................... 96 | Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania Bi July, 1941 MADRONO A WEST AMERICAN JOURNAL OF BOTANY Board of Editors Hersert L. Mason, University of California, Berkeley, Chairman. LeRoy Asrams, Stanford University, California. me Epcar Anperson, Missouri Botanical Garden, St. Louis. Lyman Benson, University of Arizona, Tucson.» Hersert F. Copetann, Sacramento Junior College, Sacramento, California. Ivan M. Jonnston, Arnold Arboretum, Jamaica Plain, Massachusetts. Mirprep E. Marutas, University of California, Berkeley. Bassett Macume, Utah State Agricultural College, Logan. Marion Ownsey, State College of Washington, Pullman. Secretary, Editorial Board—EtTHeEt Crum Department of Botany, University of California, Berkeley Business Manager—Wiu1iiam HIrsey 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 proofs are returned. Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania, for the CALIFORNIA BOTANICAL SOCIETY, INC. President: Ernest B. Babcock, University of California, Berkeley. First Vice-President: Roxana S. Ferris, Stanford University, California. Second Vice-President: Palmer Stockwell, Institute of Forest Genetics, Placerville, California. Treasurer: Wiiliam Hiesey, Carnegie Institution of Washington, Stanford University, California. Secretary: Lincoln Constance, Department of Botany, University of California, Berkeley. Annual membership dues of the California Botanical Society are $2.50, $2.00 of which is for a year’s subscription to Madrofo. Dues should be remitted to the Treasurer. General correspondence and applications for membership should be addressed to the Secretary. 1941 | LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 65 ALPINE FLORA OF SAN FRANCISCO MOUNTAIN, ARIZONA Exvsert L. Lirrie, Je. San Francisco Mountain, known locally as the San Francisco Peaks, is the highest point in Arizona and has the state’s only definitely alpine area. The disjunct alpine flora of this isolated voleanic cone is, therefore, of special interest. Forty-nine species of alpine vascular plants, including four not recorded previously for the state and several others found nowhere else in Arizona, are represented in a spare-time collection made here by the author in 1938. Although many botanists have visited San Francisco Moun- tain during the past half century, there has been no previous at- tempt to assemble a complete collection of the alpine flora or to prepare a description of the vegetation above the timberline. The author was stationed nearby at Fort Valley Experimental Forest, a branch of the Southwestern Forest and Range Experi- ment Station about nine miles northwest of Flagstaff. From this convenient base he made nine collecting trips to summits of the highest peaks during the growing season from June to September 1938, besides one trip in 1937 and two in 1939. GEOLOGY AND GEOGRAPHY This eroded volcano (text fig. 1) is situated about ten miles north of Flagstaff, Coconino County, northern Arizona (mostly in T. 22 and 23 N., R. 7 E., Gila and Salt River Meridian, center- ing near latitude 35° 20’ N., longitude 111° 40’ W.). From a high timbered plateau within Coconino National Forest about 7000 feet above sea level, it rises abruptly more than a mile to a maximum elevation of 12,655 feet and forms a prominent land- mark visible many miles from all directions. A summary of the geology and geography of San Francisco Mountain, taken mainly from Robinson’s (18) monograph, will serve as a background for the discussion of the alpine plants. The area was at no great height above the sea when volcanic activity in the form of lava flows began in the Pliocene epoch. In early Pleistocene, San Francisco Mountain with four smaller voleanoes was formed from about 38 cubic miles of lava and rose about 8800 feet above the plain. Afterwards the region was raised to its present height above sea level, and nearly four hundred small basaltic cinder cones were formed. Since the voleano became extinct San Francisco Mountain has lost by erosion about three thousand feet or more than one-third 1 Associate forest ecologist, Southwestern Forest and Range Experiment Station, which is maintained at Tucson, Arizona, by the Forest Service, United States Department of Agriculture. Maprono, Vol. 6, pp. 65-96. July 11, 1941. 66 MADRONO [Vol. 6 of its height but only about eight per cent of its volume. No trace of the former crater is left. A large interior valley, called the Inner Basin, was formed on the northeast side and was occupied by a glacier nearly two miles long probably in the late Wisconsin stage. This station was thought to be the southernmost for Pleistocene glaciation in the United States until evidence of glaci- ation was found a few years ago in the White Mountains of southern New Mexico; Pleistocene glaciation occurred also farther south than San Francisco Mountain on San Gorgonio Peak, southern California. As a result of erosion San Francisco Mountain (text fig. 1; pl. 5, fig. 1) now has the shape of a hollow, truncated cone about twelve miles or less in diameter at the base, Fic. 1. San Francisco Mountain, as viewed from Fort Valley Experimental Forest on the southwest, is a symmetrical, eroded volcanic cone. Its three high- est peaks are Humphreys Peak (left), Agassiz Peak (center) and Fremont Peak (right). two miles in diameter at the top, and one mile high. Its slope is about 20 degrees at the top and gradually becomes less towards the base. The irregular rim forms a knife-edge divide between the outer slope and the Inner Basin. Along this crest line, which is continuous except on the northeast, are six peaks above 11,000 feet in elevation. The three highest, projecting above the timber line, are: Humphreys Peak, elevation 12,655 feet, on the north- west; Agassiz Peak, elevation 12,384 feet, on the southwest; and Fremont Peak, elevation 11,984 feet, on the south. Previous BoTAaNicaLt INVESTIGATIONS No references to alpine plants of San Francisco Mountain were found in reports of the United States Government explor- ing and surveying expeditions, though this mountain lay on the 1941 | LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 67 route of several. In the United States National Herbarium are specimens of alpine plants collected on San Francisco Mountain by the following early collectors in the years indicated: E. Palmer, 1869; H. H. Rusby, 1883; M. E. Jones, 1884; Mr. and Mrs. J. G. Lemmon, 1884; E. L. Greene, 1889; F. H. Knowlton, 1889; D. T. MacDougal, 1891, 1898; J. W. Toumey, 1892; E. O. Wooton, 1892; C. A. Purpus, 1900; J. B. Leiberg, 1901; G. A. Pearson, 1908-1909. The Forest Service herbarium at Fort Valley Ex- perimental Forest has specimens of about 25 alpine species col- lected by G. A. Pearson and others. Hoffman (9) in 1877 noted that the timberline was irregular, approaching to within one thousand feet of the summit on the eastern side and five hundred feet on the western. Rusby (19) wrote in 1889 that at the summit grow only lichens and that the alpine flora, of which he named 6 species, begins one or two hundred feet below. This statement, perhaps the source of a similar one by Harshberger (8, p. 583), is not strictly correct. In a list of Arizona plants collected by E. A. Mearns, Britton (2) in 1889 mentioned 23 species from San Francisco Mountain, in- cluding about 6 occurring above timberline. The first meteoro- logical studies in the alpine zone of San Francisco Mountain were made by D. T. MacDougal in August 1898, (3, p. 43-45). There have been four detailed studies of the vegetation of San Francisco Mountain and vicinity from different viewpoints. First of these and the only one to include the alpine zone was Merriam’s (13) biological survey in 1889. A timber survey of the forests of this area was made by Leiberg, Rixon, and Dodwell (11) in 1901 and 1902. Pearson’s (16) detailed study of the forest types of San Francisco Mountain included measurement of environmental factors from 1917 to 1919 at five climatic stations in different zones from the base to the timberline. Hanson (7) in 1922 and 1928 conducted a similar series of climatic stations in his research on the vegetation of the four lower zones at the base of this mountain. The more recent summaries of Arizona vegetation by Nichol (15) and others contributed no additional information on alpine plants. San Francisco Mountain is of special historical interest to biologists because it was here in 1889 that Merriam (13) made one of the first thorough investigations of the plant and animal life of a western locality and worked out his life zones or climatic zones. From the alpine zone, above 11,500 feet, on the “‘bleak and storm-beaten summit” of San Francisco Mountain, Merriam listed 20 species of vascular plants. He recognized also the sub- alpine or timberline zone, a narrow belt at an elevation of 10,500 to 11,500 feet, where the trees became stunted and prostrate, and included a list of 16 species attaining their maximum development here. Later, when he proposed his explanation of temperature control of the geographic distribution of animals and plants, Merriam (14) omitted the subalpine zone, which does not appear 68 MADRONO [Vol. 6 to be distinct ecologically, and did not separate it from the Hud- sonian or spruce zone. Merriam’s two lists totaling 34 species comprise the only detailed published study of alpine and timberline plants of San Francisco Mountain. This collection, which was made by F. H. Knowlton and deposited in the United States National Herbarium, contains specimens of a few other alpine species not mentioned by Merriam. Corallorhiza maculata (listed as C. multiflora) was recorded from timberline zone apparently through error, as the altitude mentioned on the label (Knowlton 106) is 10,000 feet. The author has collected specimens of all the others in Merriam’s two lists except Sagina saginoides (S. Linnaei). Discussion Alpine areas above timberline on San Francisco Mountain occupy about two square miles, nearly all located on the slopes of Humphreys and Agassiz peaks (pl. 5, fig. 1). As here limited the alpine zone is that above the highest wind timber, “krumm- holz,’ or dwarf tree species (pl. 5, fig. 2). On this mountain such alpine scrub is represented by Engelmann spruce (Picea Engelmanni Parry) and bristlecone pine (Pinus aristata Engelm.). Though roads go about half-way up the mountain on four sides, the rocky alpine summits must be reached on foot. Timberline is somewhat irregular and extends nearly one thousand feet higher up the southern and western exposures than the northern and eastern. In ravines on north-facing slopes finger-like areas of alpine vegetation reach down to lower levels. Typical alpine vegetation may occur in protected areas while trees grow on exposed ridges above. Besides the obvious effects of direction of slope and topography, drifting of winter snows is partly responsible for the irregularity of the timberline. After winter storms white clouds of snow are blown off Humphreys Peak into the Inner Basin. The best development of alpine meadows is at an elevation slightly below 11,500 feet at the extreme upper end of the Inner Basin where the deeper snow drifts do not disappear until midsummer. At one place here an alpine meadow merges laterally into a subalpine meadow. ENVIRONMENTAL FacTors. Climatic and edaphic factors affect- ing the alpine vegetation here probably are similar to those of other exposed western peaks. Coville and MacDougal (3) noted that air temperature at the summit of Agassiz Peak had less daily EXPLANATION OF THE FIGURES Puate 5. ALpPIne ZONE oF SAN Francisco Mountain, Arizona. Fig. 1. View from Fremont Peak northwest across the Inner Basin, showing treeless alpine summits of Agassiz Peak (left) and Humphreys Peak (right). Fig. 2. Wind-swept, dwarf Engelmann spruce (Picea Engelmanni) at timberline ridge on west side of Agassiz Peak. Alpine rock field or lichen association in back- ground. Pearson’s timberline climatic station was nearby. Fig. 3. Alpine meadow association, in which the dominant species, Geum turbinatum, forms mats of vegetation on a rocky slope. 1941 | LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 69 ALPINE ZONE OF SAN Francisco Mountain, ARIZONA PuatTeE 5. 70 MADRONO [ Vol. 6 variation than that at lower altitudes but found a greater differ- ence between temperature of soil and air at the top than at lower levels. In the absence of other instrumental records in the alpine zone, it may be of interest to summarize comparable data from Pearson’s (16) timberline station (pl. 5, fig. 2) from 1917 to 1919 to indicate the climatic conditions under which the alpine plants are growing. During these three years the frostless period here, from June to September, varied between 101 and 113 days. Mean air temperature for the period from June to September was 47.9 degrees. Because of temperature inversion, or cold air drainage, night temperatures at timberline were little if any colder than at lower altitudes, but the days were decidedly colder. Annual precipitation at Pearson’s timberline station was slightly higher than at lower zones, totaling 28.03 inches in 1917, 37.74 inches in 1918, and 52.05 inches in 1919. The P/E ratio, or precipitation divided by evaporation, for the interval from June to September was .812. Wind movement on the exposed ridge at timberline was relatively high. During the winter months from November or December to March or April San Francisco Mountain has a white cap of snow, which buries and protects the dormant alpine vegetation. In April and May the snow melts in most places above timberline, exposing the dark-colored rock. However, on shaded slopes and areas of deep drifts, large snow banks remain until early July. In 1938 the last small patch of snow melted in the latter part of August. Frequent thunderstorms of cold rain occur in the summer rainy season from July to September. The first light winter snow in the alpine area usually falls early in October. Above timberline soils are poorly developed and the slopes are largely covered with angular blocks or boulders of andesite and pyroxene dacite. Wind, frost action, and erosion all retard the formation and accumulation of soils on the steep slopes and make the establishment of vegetation difficult. Soil pockets occur in crevices, and in places spreading mats of vegetation hold the shallow soils and help to form more. PuantT associations. The alpine vegetation of San Francisco Mountain is poorly developed and the alpine flora impoverished, in comparison with alpine vegetation and flora of larger, higher mountains. Only two plant associations are distinguished here, the alpine rock field or lichen association and the alpine meadow or Geum turbinatum association. In the xerosere the former asso- ciation gradually merges into and is succeeded by the latter, which is the climax. As no streams, lakes, or permanent snow drifts are present, there is no hydrosere. In contrast, Cox (4) described on James Peak, elevation 13,260 feet, in Colorado, a much richer alpine vegetation. The alpine meadow, or Geum turbinatum association of San Francisco Mountain corresponds with the dry meadow associations on 1941 ] LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 71 James Peak, where Carex rupestris and Geum turbinatum are dom- inant. On James Peak, however, the dry meadow is succeeded by a climax alpine meadow association in which Elyna Bellardi, a species not found on San Francisco Mountain, is dominant. The various communities in hydroseres from wet stony areas, stream and lake margins, and snow drifts through scrub, moor, and wet meadow types to the climax alpine meadow of James Peak all are absent from San Francisco Mountain. ALPINE ROCK FIELD. The alpine rock field or lichen association occupies most of the mountain summit of San Francisco Moun- tain, including exposed peaks and ridges (pl. 5, fig. 2). How- ever, some unstable rock slides are almost without plants. This association is characterized by lichens, principally crustose and foliose, on the rock outcrops with vascular plants scattered among the boulders wherever there is sufficient soil. Even on the highest, exposed peaks a few seed plants can be found. Probably the most abundantly represented species or dominant species is the crustose lichen Rhizocarpon geographicum (L.) Lam. & DC. (de- termination verified by Dr. Carroll W. Dodge). Two species of foliose lichens, of which specimens were collected here previously for the Forest Service by F. W. Haasis and J. O. Veatch and determined by G. K. Merrill, are Lecanora rubina ( Vill.) Ach. and Parmelia conspersa (Ehrh.) Ach. Mosses are less common than lichens and are found in crevices and to a lesser extent on boul- ders. Hypnum revolutum (Mitt.) Lindb. is characteristic of the shaded soil pockets. The moss species most commonly repre- sented on dry outcrops is Grimmia Doniana Sm., which, according to Edwin B. Bartram who determined the two mosses, may be the first record for the state. The only alpine pteridophyte, Cystopteris fragilis, is commonly distributed in crevices. The number of species of flowering plants represented in the alpine rock field is high and includes nearly all occurring above timberline. Of these the mat-forming pioneer species of the alpine meadow, Geum turbinatum, is most conspicuously repre- sented. Five characteristic species of Caryophyllaceae here are Arenaria Fendleri, A. sajanensis, A. verna, Cerastium Beeringianum, and Silene acaulis. Three species of grasses commonly repre- sented are Festuca ovina var. brachyphylla, Poa rupicola, and Trise- tum spicatum. Carea bella is the sedge species most commonly represented, and Luzula spicata the typical rush. Other charac- teristic species in the rock field include Polemonium confertum, Solidago ciliosa, Thlaspi Fendleri, and Pseudocymopterus montanus. Typical pioneer species on rock slides are Geum turbinatum, Primula Parryi, Senecio franciscanus, and Arenaria sazxosa. In shaded crevices and cliffs Heuchera versicolor f. pumila and Oxyria digyna are commonly represented. ALPINE MEADOW. The alpine meadow or Geum _ turbinatum climax association is developed on the more favorable sites, such 72 MADRONO [ Vol. 6 as at the upper end of the Inner Basin and ravines not far above timberline (pl. 5, fig. 3). However, the area occupied is only a small fraction of the alpine zone. Plants of the dominant species, Geum turbinatum, start as pioneers on rocky slopes and especially at the bases of rock slides. They spread vegetatively by creeping stems to form compact mats in which soil is built up and other plants become established to form an almost solid mantle of vege- tation. If any species of vascular plants may be said to be abund- antly represented above timberline, it is this. Other mat-form- ing species here include Potentilla Sibbaldi and Silene acaulis. Most of the species listed under rock fields are commonly represented also in meadows. The following species of grasses and sedges are typical: Festuca ovina var. brachyphylla, Trisetum spicatum, Carea bella, C. albo-nigra, and C. ebenea. Other species of meadows include Luzula spicata, Arenaria sajanensis, Cerastium Beeringianum, and Solidago ciliosa. Scattered or rare in alpine meadows are individuals of the following species not observed elsewhere above timberline: Phleum alpinum, Poa reflexa, Gentiana barbellata, G. monantha, Pedicularis Parryi, Veronica Wormskjoldi, and Erigeron simple.. Mosses of several species are present in alpine meadows but are not common. Liverworts of a single species, Lophozia por- phyroleuca (Nees) Schiffn. (determination by Dr. A. W. Evans), were found above timberline. Not reported previously from Arizona, this species is represented sparingly in alpine meadows and more commonly in the subalpine forest. This climax association is slowly spreading through mat for- mation by the dominant species, and all degrees of intermediate stages between rock field and meadow can be observed. The two associations sometimes are mixed together, for example near the bases of steep slopes where large boulders which have rolled down from above have a vegetation of lichens though surrounded by a climax meadow. Above timberline growth begins in June, or possibly earlier on southern and western exposures. By the middle of June plants of the following species are in flower: Thlaspi Fendleri, Geum turbinatum, Pseudocymopterus montanus, and Polemonium con- fertum. Most of the alpine species blossom in July and August. Among the last species to begin flowering, in August and Sep- tember, are Gentiana barbellata, Erigeron simplex, Senecio francis- canus, and Solidago ciliosa. Plants of Geum turbinatum continue to flower throughout the summer. By the first of October even the last late flowers have been killed by frosts and the vegetation has become dormant for the winter. GroGrAPHic DisTRIBUTION oF Species. Four species of vascular plants found by the author on San Francisco Mountain appar- ently are here first recorded for the state with corresponding extensions of range. All have a northern distribution and reach their southern limits on this and similar mountain summits. The 1941] LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 73 alpines are Juncus Drummondii, Erigeron compositus, and Erigeron simplex. More unusual is Botrychium lanceolatum, a subalpine pteridophyte reaching the timberline. This circumboreal species has been recorded previously from only three western states, Washington, Wyoming, and Colorado. As the author (12) has noted, the station on San Francisco Mountain is the southernmost reported in North America and nearly four hundred miles south- west of the nearest known localities in central Colorado. A few other alpine species here have been found nowhere else in Arizona, but some occur also on Arizona’s second highest mountain, Thomas or “Baldy” Peak, elevation 11,470 feet, in the White Mountains one hundred and fifty miles southeast. Merriam (13) observed that many of the species above tim- berline on San Francisco Mountain are arctic-alpine circumpolar species widely distributed in arctic regions of North America and Eurasia, such as Alaska, Hudson Bay, Greenland, Spitzbergen, and Siberia, and extend far south on summits of such high moun- tain ranges, as the Sierra Nevada, Rocky Mountains, Alps, and Himalayas. However, after an interval of fifty years, there are fewer species in common with these more distant points, owing to increased knowledge and probably also to a trend toward a narrower concept of the species. As a result of such changes and of corrections in nomenclature, only one-half of the 34 species listed by Merriam (138) as occurring in the alpine and timberline zones now bear the same names. The occurrence of disjunct arctic-alpine plants on San Fran- cisco Mountain was explained by Merriam according to the present interpretation, that of migration southward during the glacial period and the ascension of mountain peaks as the tem- perature became warmer. He concluded that, as these species could not have reached here afterwards, the volcano could not be of more recent origin than the glacial period. Rydberg (20) classified the alpine flora of the Rocky Moun- tains into ten groups according to geographical distribution, while Holm (10) in a detailed analysis of the alpine flora of Colorado distinguished four elements as to origin. The 49 alpine species of San Francisco Mountain fall into five more or less natural groups, according to their ranges as given in floras and manuals and in the phytogeographical treatises by Rydberg (20) and Holm (10). These groups are listed below with the abbreviations used in the list of species, the number of species in each, and the per- centage of species in each. Number of Pp : er cent species Arctic-alpine circumpolar species (CP) .... 15 30.6 North American arctic-alpine species (NA) 5 10.2 Rocky Mountain alpine species (RM) ..... 24 49.0 Southwestern species (SW) ............. 3 all Emdemic species (EN) ......52. 6.400880 5 2 4.1 re) MADRONO [Vol. 6 Thus, the relationships of the alpine flora of San Francisco Mountain are quite definitely northern with the floras of arctic and alpine regions. The 15 arctic-alpine circumpolar species, almost one-third of the total, are widely distributed in arctic regions in North America and Eurasia and on alpine mountain summits at lower latitudes. Two of these, Arenaria sajanensis and Stellaria umbellata, are recorded from Siberia but not from Europe, while the rest are listed from both Europe and Asia. A few have even wider distribution. Cystopteris fragilis is almost cosmopoli- tan and ranges to tropical America. Phleum alpinum and Trisetum spicatum are found also in alpine South America. The 5 North American arctic-alpine species have a distribution from Greenland to Alaska, or slightly narrower, and southward on high western mountains but are not listed from Eurasia. The 24 Rocky Mountain alpine species occur in high mountains gen- erally from Montana south to Colorado and Utah and on the highest peaks of New Mexico and Arizona. Some of these are found also in somewhat lower zones. Three southwestern species are confined to New Mexico and Arizona or slightly beyond and apparently have migrated up to the alpine area from lower zones, where they are more typical. Only two alpine species, Senecio franciscanus Greene (5, p. 19- 20), and an undescribed species of Pedicularis, are endemic to San Francisco Mountain, and neither furnishes a clue as to routes of migration. In his monograph Greenman (6, p. 29-30) listed only 6 collections of Senecio franciscanus, all from this mountain. Though Senecio is well represented in alpine regions with several species confined to Colorado, the species apparently most closely related to S. franciscanus are distributed nearby at lower zones in the same area. The undescribed species of Pedicularis is an endemic closely related to the widely ranging species, P. Parryi A. Gray. Several other alpine plants here have been proposed as new species, but these have been reduced to synonymy, with the exception of one or two retained as varieties of other species. However, a large number of endemic species is not to be expected on a geologically young volcanic cone. Most of the alpine species of San Francisco Mountain are found also below timberline, especially in subalpine meadows. The following commonly represented species, as well as a few others, are confined to the alpine zone here: Arenaria sajanensis, Silene acaulis, Polemonium confertum, and Senecio franciscanus. A few, such as Arenaria Fendleri and Thlaspi Fendleri, are equally characteristic of the western yellow pine zone at the base of the mountain. By comparing the alpine flora of San Francisco Mountain with similar plant lists from other southwestern peaks, it should be possible to trace the probable routes of the arctic-alpine plants in migrating to this newly formed volcano. Bailey’s (1, p. 51-53) list of 60 characteristic species of the arctic-alpine zone in New 1941] LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 75 Mexico indicates a much richer alpine flora in the Sangre de Cristo Range at the southern end of the Rocky Mountains more than three hundred miles eastward. The nearest alpine peaks of the main Rocky Mountains are in Colorado about two hundred and fifty miles northeast. Thirty-eight of the 50 alpine species on San Francisco Mountain are included in Cox’s (4) list of 196 alpine species on James Peak, Colorado. The high plateau of Arizona, on which San Francisco Mountain is located, is almost continuous with alpine summits in southern Utah two hundred miles north- ward and connects on the southeast with the White Mountains one hundred and fifty miles distant and with mountains in western New Mexico. RauNkKI4ER’s Lire-rorms. The 48 alpine species of seed plants on San Francisco Mountain were grouped also according to Raun- kiaer’s (17) life-forms. Phanerophytes are absent above timber- line, and only four life-forms are represented. Most of the spe- cies (40) are perennial herbs classed as hemicryptophytes. The remaining 8 are listed below according to life-form. The 5 spe- cies of chamaephytes are: Geum turbinatum, the abundantly repre- sented, mat-forming species; Silene acaulis, a typical cushion species; Arenaria sajanensis, Saxifraga caespitosa subsp. exaratoides var. Lemmonii, and Potentilla Sibbaldi. The single cryptophyte or geophyte species is Senecio franciscanus, which is especially suc- cessful on rock slides because of its creeping rhizomes. There are only 2 species of therophytes or annual herbs, Androsace septen- trionalis var. subumbellata and Gentiana monantha. As others have noted, the distinction between chamaephytes and hemicryptophytes is not sharp, and a few of the species classed as the latter might as well have been called chamaephytes. At any rate, the prevailing life-form here is that of low perennial herbs with rosettes and stems which, often retaining bases of old leaves, elongate slightly each year. The biological spectrum of the alpine zone of San Francisco Mountain and Raunkiaer’s revised normal spectrum are compared below, with percentages of species under each life-form. The slightly higher percentage of chamaephytes above that of the normal spectrum indicates a cold climate intermediate between the temperate hemicryptophytic climate and the chamaephytic climate of arctic regions. Alpine zone of Raunkiaer’s revised : San Francisco normal spectrum P Mountain er cent Per cent Phanerophytes .......... 46 0 @lamaephytes .......... 9 10.4 Hemicryptophytes ....... 26 83.3 Cuyptophytes ........... 6 2.1 MMineroplytes 2........... 13 4.2 16 MADRONO [Vol. 6 List oF ALPINE SPECIES This study is limited to the vascular plants above timberline on San Francisco Mountain. All species normally found at some distance above the irregular line of the last stunted individuals of tree species were included, regardless of their occurrence in other zones. As it was sometimes dificult to determine whether a spe- cies should be classed as alpine, a partial list of subalpine species reaching the timber line or slightly beyond is included as evidence that these species were not overlooked. For example, the shrubby species Juniperus communis var. sibirica and Ribes montigenum, which occur in protected rocky situations slightly above the high- est spruces and pines, are not typically alpine. In naming the specimens the author wishes to acknowledge the valuable assistance of several specialists. The author made deter- minations at the herbarium of the Field Station of the Bureau of Plant Industry at Sacaton, Arizona, and at the United States National Herbarium. Specimens of certain groups were further checked by the following specialists: Pteridophyta by Dr. Wm. R. Maxon, Gramineae by Dr. J. R. Swallen and Prof. J. J. Thornber, Carex by J. W. Stacey, Juncus by Dr. F. J. Hermann, Ranunculus by Dr. Lyman Benson, and Compositae by Dr. S. F. Blake. De- terminations of all the remaining specimens have kindly been checked by Dr. T. H. Kearney. Nomenclature is that of the forth- coming Flora of Arizona, by T. H. Kearney and R. H. Peebles. The names thus are in accord with the International Rules of Botanical Nomenclature and are conservative. Duplicate sets of the author’s specimens have been deposited in the United States National Herbarium, herbarium of the Uni- versity of Arizona, herbarium of the Sacaton (Arizona) Field Sta- tion, and herbaria of the Forest Service at the following localities: Washington, D. C.; Albuquerque, New Mexico; Southwestern Forest and Range Experiment Station at Tucson, Arizona, and Fort Valley Experimental Forest near Flagstaff, Arizona. The 49 alpine species of vascular plants of San Francisco Mountain, including a single pteridophyte and 48 species of flowering plants, are grouped into 35 genera and 17 families. Specimens of two additional species were undeterminable. Larg- est families are Caryophyllaceae (a family well represented at northern points and high elevations) with 8 species, Compositae with 7, and Gramineae with 6 species. Principal genera are Arenaria and Sazifraga with 4 species each and Carez and Potentilla with 8 each. Rydberg (20, p. 92-93) lists from San Francisco Mountain an additional species, Lloydia serotina (L.) Sweet, which is omitted here because no herbarium specimen from Arizona could be located. The species are listed below with brief notes on habitat and abundance. Geographic distribution or range is abbreviated as explained above. 1941 ] LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 77 PoLYPODIACEAE CysTopTERis FRAGILIS (L.) Bernh. Crevices, alpine rock fields; common. CP. GRAMINEAE AGropyron Scrisneri Vasey. Loose rocks and gravel, alpine rock fields; scattered. The weak stems spread out in all direc- tions over the rocks. RM. Festuca ovina L. var. BRACHYPHYLLA (Schult.) Piper. Alpine meadows and rock fields; common. CP. PuHieumM aLpinum L. Alpine zone, confined to climax alpine meadows; rare. CP. Poa REFLEXA Vasey & Scribn. Alpine zone, in meadows; rare. RM. Poa ruPpicota Nash. Alpine meadows and rock fields; com- mon. RM. TrRisETUM spicatum (L.) Richt. Alpine rock fields and mead- ows;common. CP. CYPERACEAE Carex ALBo-NIGgRA Mackenzie. Alpine rock fields and mead- ows; common. RM. Carex BELLA L. H. Bailey. Alpine rock fields and meadows; the most commonly represented species of Carex. Flowers from June to September. RM. CaREX EBENEA Rydb. Alpine rock fields and meadows; com- mon. RM. JUNCACEAE Juncus Drummonpn E. Meyer. This species, previously un- recorded for Arizona, was to be expected here, as it ranges from Alaska south in mountains to New Mexico and California. Rare in one alpine meadow at the upper end of the Inner Basin and commoner in nearby subalpine meadows. RM. Luzuxa spicata (L.) DC. Alpine meadows and rock fields; common. CP. POLYGONACEAE Oxyria pieyna (L.) Hill. Shaded rock crevices of alpine zone; uncommon. CP. CARYOPHYLLACEAE ARENARIA FeNpLERI A. Gray. Typical of lower elevations, a depauperate alpine form common in alpine rock fields. RM. ARENARIA SAJANENSIS Willd. Alpine rock fields and meadows; common. CP. ARENARIA SAxoSA A. Gray. Rock slides in alpine zone but more characteristic of lower zones; common. SW. ARENARIA VERNA L. Alpine rock fields and meadows; com- mon. Nearly all the plants are densely glandular pubescent, but rarely glabrous plants are found. CP. Crerastium Brrerineianum Schlecht. & Cham. Alpine rock fields; very common. NA. 18 MADRONO [ Vol. 6 SaGINna saGINorpEs (L.) Britton. Alpine zone (Knowlton 134, U. S. National Herbarium); rare. Not collected by the author. CP. SILENE acautis L. Alpine meadows and rock fields ; commonly forming cushions or mats. CP. STELLARIA UMBELLATA Turez. Alpine rock crevices and mead- ows; uncommon. CP. RANUNCULACEAE RANUNCULUS INAMOENUS Greene var. ALPEOPHILUS (A. Nels.) L. Benson and var. suBAFFINIS (A. Gray) L. Benson. Var. subaffinis is endemic to San Francisco Mountain. Alpine meadows and in mats of other plants; rare and scattered. RM. Ranuncutus sp. Alpine meadows and in mats of other plants ; rare and scattered. An apetalous plant. Perhaps a reduced form of R. pedatifidus J. E. Smith, which sometimes lacks petals. CRUCIFERAE DrasBa aurEA Vahl var. aureIrorMis (Rydb.) Schulz. Alpine rock fields; common. NA. Drapa crassiFotiA Graham. Alpine meadows; rare. NA. Tuuaspt Fenpter1 A. Gray. Alpine rock fields and meadows; common. SW. CRASSULACEAE SEDUM RHODANTHUM A. Gray. Alpine meadows and rock fields; common. RM. SAXIFRAGACEAE HEUCHERA VERSICOLOR Greene f. pumiLta Rosend. et al. Alpine rock ledges and crevices and also in alpine meadows; common. SW. SAXIFRAGA CAEsPITOsSA L. var. LEmMmMonnt’ Engler & Irmscher. Forming mats on alpine ledges and rock fields and occasionally in alpine meadows; not common. CP. SAXIFRAGA DEBILIS Engelm. Shallow soils in shaded crevices, alpine rock fields; uncommon. RM. SAXIFRAGA FLAGELLARIS Willd. Mainly in crevices of alpine rock fields; not common. CP. SAXIFRAGA RHOMBOIDEA Greene var. FRANCISCANA (Small) Kear- ney & Peebles. Alpine rock fields and meadows; uncommon and scattered in mats of other plants. Var. franciscana perhaps is not worthy of recognition, as it differs only in smaller size and more compact inflorescences. These differences apparently are associ- ated with the alpine habitat. RM. RosacEAkE Geum TuRBINATUM Rydb. Sieversia turbinata (Rydb.) Greene. Dominant in alpine meadows and a pioneer on rocks; by far the most abundantly represented species of flowering plants in the alpine area. RM. 1941] LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 79 PoTENTILLA conciNNA Richardson. Alpine rock fields; un- common. RM. PoTeNTILLA DivERsIFoLIA Lehm. Alpine rock fields and mead- ows;notcommon. RM. PoTENTILLA SipBaLpI Haller f. (Sibbaldia procumbens L.) In mats in alpine meadows and slightly below timberline; common. Cr. UMBELLIFERAE PsEUDOCYMOoPTERUS MoNTANUS (A. Gray) Coult. & Rose. In alpine rock fields and also in alpine meadows; common. RM. PRIMULACEAE ANDROSACE SEPTENTRIONALIS L. var. SUBUMBELLATA A. Nels. Alpine rock fields; common. CP. Primuta Parryr A. Gray. Alpine rock fields, including crev- ices and rock slides, and in meadows; common but scattered. RM. GENTIANACEAE GENTIANA BARBELLATA Engelm. Very rare and scattered in alpine meadows but more typical of lower zones. Plants above timberline are inconspicuous until they flower in September. RM. GENTIANA MOoNANTHA A. Nels. In mats of other plants in alpine meadows; rare. RM. POLEMONIACEAE PoLEMONIUM coNFERTUM A. Gray. Alpine rock fields and also in meadows; common. RM. SCROPHULARIACEAE Pepicuuaris sp. Rare in climax alpine meadows and common below timberline. Previously referred to Pedicularis Parryi A. Gray. However, Dr. Francis W. Pennell, whom I accompanied on a field trip here to collect additional material, is describing the San Francisco Mountain plant as a new, endemic species. EN. Veronica WormskJotpir Roem. & Schult. Commoner below timberline, rare in alpine meadows. NA. CoMPOSITAE ANTENNARIA APRicA Greene. Alpine rock fields; uncommon. RM. ANTENNARIA UMBRINELLA Rydb. Alpine rock fields and mead- ows; uncommon. RM. ERIGERON compositTus Pursh. Reported previously from Green- land to Alaska and south to Colorado, Utah, Nevada, and Cali- fornia. Very rare in an alpine meadow and in rock cliffs at timberline. NA. ERIGERON sIMPLEX Greene. Previously known from Montana south to Colorado, New Mexico, and California. Though this species typically is radiate, some of the plants here are discoid or 80 MADRONO | [Vol. 6 nearly so. Scattered and uncommon in mats of other plants in alpine meadows. RM. SENECIO FRANCISCANUS Greene. Endemic to the alpine zone of San Francisco Mountain, where it is commonly represented on rock slides. Discovered in 1884, but the type was collected by Greene in 1889. EN. , So.ipaGo ciLiosA Greene. Alpine rock fields and meadows; common. RM. Taraxacum sp. Possibly a new record for the state, but the scarce material was undeterminable. In mats of other plants in alpine rock fields; very rare. ParTiaL List or TIMBERLINE SPECIES Specimens of the 24 additional species in this partial list of timberline species were collected by the author in the wind timber or subalpine meadows at timberline or slightly above. Botrychium lanceolatum (S. G. Gmel.) Angstr. Woodsia oregana D. C. Eaton Juniperus communis L. var. sibirica (Burgsd.) Rydb. Bromus ciliatus L. Poa Fendleriana (Steud.) Vasey Poa pratensis L. Sitanion Hystrix (Nutt.) J. G. Smith Carex Haydeniana Olney Carex siccata Dewe Luzula parviflora (Ehrh.) Desv. Anemone globosa Nutt. Aquilegia chrysantha A. Gray Ribes montigenum McClatchie Potentilla Hippiana Lehm. var. diffusa Lehm. Potentilla subviscosa Greene Rubus strigosus Michx. var. arizonicus (Greene) Kearney & Peebles Geranium Richardsonii Fisch. & Trautv. Viola canadensis L. Epilobium angustifolium L. Epilobium saximontanum Haussk. Moneses uniflora (L.) A. Gray Mertensia franciscana Heller Penstemon Whippleanus A. Gray Agoseris purpurea (A. Gray) Greene ? SUMMARY San Francisco Mountain, elevation 12,655 feet, is an isolated voleanic cone located about ten miles north of Flagstaff, Coconino County, northern Arizona. The highest point in the state, it has a disjunct alpine flora which is Rocky Mountain alpine and arctic- alpine circumpolar in relationships. Two plant associations are distinguished in the alpine zone here, the alpine rock field or 1941] LITTLE: SAN FRANCISCO MOUNTAIN, ARIZONA 81 lichen association on most of the rocky summit and the alpine meadow or Geum turbinatum climax association on limited, more favorable sites. Based upon collections by the author in 19388, 49 species of alpine vascular species and 24 additional timberline species are listed with brief notes on habitat and abundance. Four of these are previously unrecorded for the state: Botrychium lanceolatum (subalpine at timberline), Juncus Drummondu, Erigeron compositus, and Erigeron simplex. 12. 13. 14. 15. 16. Li. 18. 19. 20. Tucson, Arizona, May, 1940. LITERATURE CITED Baitey, Vernon. Life zones and crop zones of New Mexico. U. S. Dept. Agr. Biol. Surv., N. Amer. Fauna 35. 100 p., illus. 1913. Britton, N. L. A list of plants collected at Fort Verde and vicinity and in the Mogollon and San Francisco mountains, Arizona, 1884-1888, by Dr. E. A. Mearns, U.S. A. N. Y. Acad. Sci. Trans. 8: 61-76. 1889. Covitte, Freperick V., and Daniet T. MacDovucat. Desert Botanical Laboratory of the Carnegie Institution. Carn. Inst. Wash. Publ. 6. 58 p., illus. 1903. Cox, Cruare Francis. Alpine plant succession on James Peak, Colorado. Ecol. Monographs 3: 299-372, illus. 1933. GREENE, Epwarp L. New or noteworthy species VI. Pittonia 2: 13-24. 1889. GreeNMAN, J. M. Monograph of the North and Central American species of the genus Senecio—Part II. Mo. Bot. Gard. Ann. 4: 15-35. 1917. Hanson, Hersert C. A study of the vegetation of northeastern Arizona. Neb. Univ., Studies 24: 85-178, illus. 1924. HarsHpercer, JOHN W. Phytogeographic survey of North America. 790 p., illus. 1911. Horrman, W. J. The distribution of vegetation in portions of Nevada and Arizona. Amer. Nat. 11: 336-343. 1877. Hoim, TuHeropore. The vegetation of the alpine region of the Rocky Moun- tains in Colorado. Nat. Acad. Sci. Mem. 19 (3): 1-45, illus. 1927. . Lerperc, Jonn B., TuHeopore F. Rixon, and Arruur Dopwetu. Forest conditions in the San Francisco Mountains Forest Reserve, Arizona. U. S. Geol. Surv. Prof. Paper no. 22, Series H, Forestry, 7. 95 p., illus. 1904. Lirrie, Exsert L., Jr. Botrychium lanceolatum in Arizona. Amer. Fern Journ. 29: 36-37. 1939. Merriam, C. Harr. Results of a biological survey of the San Francisco Mountain region and desert of the Little Colorado in Arizona. U. S. Dept. Agr. Div. Ornith. Mammal., N. Amer. Fauna 3. 136 p., illus. 1890. . Life zones and crop zones of the United States. U. S. Dept. Agr. Div. Biol. Surv. Bull. 10. 79 p., illus. 1898. Nicnuot, A. A. The natural vegetation of Arizona. Univ. Ariz. Agr. Exp. Sta. Tech. Bull. 68: 179-222. 1937. Pearson, G. A. Forest types in the Southwest as determined by climate and soil. U.S. Dept. Agr. Tech. Bull. 247. 143 p., illus. 1931. Raunxtiarr, C. The life forms of plants and statistical plant geography. 632 p., illus. 1934. Rosinson, Henry H. The San Franciscan voleanic field, Arizona. U. S. Geol. Surv. Prof. Paper 76. 213 p., illus. 1913. Russy, H. H. General floral features of the San Francisco and Mogollon mountains of Arizona and New Mexico, and their adjacent regions. N. Y. Acad. Sci. Trans. 8: 76-81. 1889. Rypgerc, P. A. Phytogeographical notes on the Rocky Mountain region. II. Origin of the alpine flora. Torr. Bot. Club Bull. 41: 89-103. 1914. 82 MADRONO [Vol. 6 MARY FISK SPENCER Mary Fisk Spencer died in San Diego, California, December 27, 1940. She was within twelve days of completing her one hundredth year. For fifty of those years, in spite of weak eye- sight and handicapped by poor health, she had been an inde- fatigable botanizer; from 1878 to 1914 in Central and Southern Europe, and from 1915 to 1928 in southern California, chiefly in the arid sections of San Diego County. Her collections are found in the leading herbaria of Europe and America and in the private herbaria of many specialists, especially in Europe. Her death marks the passing of an outstanding amateur botanist. Mary Evelyn Fisk was born at Brecksville, Ohio, January 8, 1841, the daughter of John Shipley Fisk and Anne Clapp Clark, both of whom were from New England. MHer father, a master machinist, changed his residence frequently, being in charge suc- cessively of iron works in Knoxville, Tennessee, Franklin Falls, Ohio, and Cleveland, Ohio. As a result of change of residence and financial difficulties Miss Fisk’s education was badly inter- rupted. When she was fifteen she enrolled in the Preparatory Department of Oberlin College, which was then almost the only coeducational institution in the middle west, if not in the entire country. <A year of teaching followed, after which she enrolled in the Literary Department of the College for the years 1858— 1859 and 1860-1861. A bank failure put an end to her college work. Miss Fisk was musically gifted and while at Oberlin must have received considerable musical instruction, since, then as now, Oberlin was giving great attention to music. During the early eighteen-sixties she taught private music classes in and about Cleveland, but about 1865 was able to enter the old Boston (Massachusetts) Music School for study under B. F. Baker; she graduated there, then taught music privately in Boston. How- ever, she wanted further training and by 1874 was studying at the Royal Conservatory of Music in Munich, Germany. After two years of strenuous work she suffered a nervous breakdown, and was thereafter compelled to lead an outdoor life. During this period of European study she met James MacGof- fin Spencer, who had gone to Munich in 1874 to study chemistry. They were married on July 28, 1878, and until September 1914, when they returned to the United States to live, resided in Munich. Mr. Spencer had been for six years Professor of Mathematics in the National Deaf and Dumb College in Washington, D.C. After a severe illness he was obliged to lead a life of leisure, which he diversified by travel and study. Thus it was necessary for both Mr. and Mrs. Spencer to have a rather quiet, outdoor life; and their means, though modest, permitted extensive travel. During the years that Miss Fisk was in Oberlin she became a devoted admirer of Dr. James Dascomb (M. D. Dartmouth, 1833), 5. 6 92 tbout 1 ‘ Cc ken a aph t Photogr x SPENCER SI Fr ARY = 1941 | GROVER: MARY FISK SPENCER 83 who was “Professor of Chemistry, Botany and Physiology” at Oberlin from 18384 to 1878, and Librarian from 1855 to 1873. Through his influence she developed a love of botany, which after her marriage became a life passion. In company with her hus- band she visited most of the countries of Central and Southern Europe, and wherever she went she made extensive collections of the native flora. These collections she exchanged with leading botanists and herbaria, and thus assembled a herbarium of more than 12,000 sheets of vascular plants, chiefly from Central and Southern Europe, with a few from Norway, the British Isles, North Africa and Asia Minor. This herbarium was later presented to Oberlin College. It constitutes the chief part of the European section of the College Herbarium which now contains about 100,000 sheets of vascular plants, and 50,000 of nonvascular plants, housed in steel, moth- proof cases in a fireproof addition to the present botany building. In November 1900 in a letter to Professor Albert A. Wright of Oberlin, who had been Professor of Geology, Botany and Zoology after Dr. Dascomb’s retirement, Mrs. Spencer wrote “I have been collecting for over twenty years, and have a large proportion of the plants of Central Europe which I intend, after I shall have made the collection as complete as I can, to present to Oberlin as an expression of affectionate regard for our dear Prof. Das- comb of blessed memory.” Although her sight and health were even then failing she continued to build up her herbarium, the first installment of which was given to the College in 1908. She stipulated that the collection be maintained as a separate unit, and be called “The Spencer collection of European Plants, in me- moriam Professor James Dascomb.” She herself was in the United States from 1908 to 1910 to visit friends and to supervise the installation of the herbarium. She returned to Europe in the fall of 1910, where she continued to reside until just after the out- break of the first world war. Three years of this time were spent in Corsica, where she made extensive collections that were later given to Oberlin. After January, 1915, Mrs. Spencer resided in San Diego, Cali- fornia, until her death there on December 27, 1940. Mr. Spencer died May 18, 1920, in Burlington, Vermont. In the arid and semiarid areas of San Diego County, Mrs. Spencer found a new and intensely interesting flora. Until her eyesight and health failed completely in 1927 she spent weeks at a time at all seasons of the year collecting in the desert regions as far south as the Mexican border. Many of the plants which she herself was unable to identify were determined at Gray Her- barium, a few by Dr. Greenman at the Missouri Botanical Garden and some of her last collections by Miss Fidella G. Woodcock, Curator of Plants in the Natural History Museum, San Diego. Soon after arriving in San Diego, she become acquainted with Mr. 84 ; MADRONO [ Vol. 6 C. H. Orcutt, from whom she obtained much assistance. A com- plete set of her California collections, with some from Arizona were also given to Oberlin College, the last installment being received in February, 1928. There were approximately 3000 numbers, many species being represented by several specimens collected in different seasons, at different altitudes, and from dif- ferent ecological areas. FReperiIcK Grover, Oberlin College, Ohio. NEW SPECIES OF VASCULAR PLANTS FROM THE NORTHWEST COAST GrorcGe NEVILLE JONES During the preparation of a manual of botany of the North- west Coast, covering the vascular flora of the region west of the Cascade Mountains from British Columbia to Oregon, two addi- tions to the number of described species must be recorded and a new nomenclatural transfer is necessary at this time. In citing specimens, the following abbreviations are used: Gray Herbarium of Harvard University (G); Missouri Botanical Garden (Mo) ; University of Illinois (UI). In treating the willows of the area west of the Cascade Moun- tains it has been necessary to re-examine that small group of species closely related to Salix Geyeriana Anderss. The conclu- sion has been reached that the willow of this group occurring west of the Cascade Mountains is specifically distinct from the eastern Washington and Rocky Mountain S. Geyeriana, and the following nomenclatural combination is herewith proposed: Salix meleina (J. K. Henry) comb. nov. S. Geyeriana Anderss. var. meleina J. K. Henry, Fl. So. Brit. Columbia 98. 1915. 4s Geyeriana meleina Ball, in Abrams, Ill. Fl. Pacific States 1: 506. 1923. Salix Geyeriana is a smaller shrub with densely glaucous branchlets, the pubescence of the leaves is whitish-sericeous, the capsules somewhat longer, and the style obsolete. It is not known to occur west of the Cascade Mountains. Delphinium splendens sp.nov. Herba perennis; radicis fibris elongatis vel subfusiformibus, lignosis; caulibus strictis 1-2 m. altis, simplicibus, glaucis, fistulosis; foliis palmatim 5-—7-lobatis, | 10-30 cm. latis; racemis elongatis, pedicellis glabris, quam flori- bus longioribus, adscendentibus; floribus 15-20 mm. longis; sepalis caerulescentibus, intus glabris, extus puberulis; calcare 8-10 mm. longo, patente vel ad apicem subcurvato; petalis 4, albicantibus, spatulatis, obtusis, 8-10 mm. longis, lamina un- gueque hirtulis; staminibus ca. 25; antheris ovalibus 1.5 mm. longis; folliculis 3, erectis, rectis, glabris vel subglabris, tenuiter 1941 | JONES: NORTHWESTERN PLANTS 85 reticulato-venulosis, 7-12 mm. longis, ad apicem nempe truncatis, in stylum abrupte abeuntibus; pedicellis fructigeris erectis vel adscendentibus; seminibus 3 mm. longis, costis in alas hyalinas productis. Perennial; stem glaucous, leafy, simple, glabrous throughout, hollow, 1-2 m. tall, arising from a fascicle of elongate or some- what fusiform woody roots; leaves palmately 5—7-lobed, 10-80 em. broad, cleft about half way to the middle into oblanceolate, coarsely dentate, cuneate-based acute lobes, dark green, sparsely pilosulous above especially along the veins, pale green and grayish pilosulous-puberulent beneath; petioles glabrous, 10-15 cm. long, longer than the blades ; racemes narrow, many-flowered, 25—50 cm. long; pedicels glabrous, slender, not longer than the flowers, the lower ones 8-15 mm. long at flowering time, ascending; flowers 15-20 mm. long; sepals dull blue, green-veined, glabrous within, puberulent outside; spur 8-10 mm. long, puberulent outside, acutish or obtuse, straight, horizontally spreading, or very slightly curved toward the tip; petals 4, whitish, spatulate, obtuse, 8—10 mm. long, the blade and claw hirsutulous; stamens about 25; anthers oval, 1.5 mm. long, glabrous; filaments 3-5 mm. long, translucent, dilated below, glabrous to sparsely pilosulous; folli- cles 3, erect, straight, glabrous or nearly so, somewhat reticulate- veiny, 7-12 mm. long, the style 3-4 mm. long, the follicle some- what truncate at the apex, abruptly terminating in the style, the fruiting pedicels erect or ascending; seeds 3 mm. long, the angles produced into hyaline wings. The synonymy is as follows: Delphinium scopulorum glaucum sensu Piper, Contr. U. S. Nat. Herb. 11: 280. 1906, Henry FI. So. Brit. Col. 188. 1915; D. glaucum sensu Piper & Beattie, Fl. N.W. Coast 160. 1915, G.N. Jones, Univ. Washington Publ. Biol. 5: 153. 1986, op. cit. 7: 82. 1938, not Wats. Bot. Calif. 2: 427. 1880. Auaska: Chilkat Valley, Walker 1070 (Mo). WasutneTon: [Mount Rainier] Upper Valley of the Nisqually, July 14, 1896, O. D. Allen 248 (type UI, isotype Mo); Swauk River, Sharples 78 (UI); Yakima County, Brandegee 615 (UI); Olympic Mountains, Elmer 2577 (Mo), Piper in 1890 (Mo), G. N. Jones 10735 (UI). This northwestern larkspur has been passing as Delphinium glaucum Wats., a species described from the Sierra Nevada of California in 1876. It grows in subalpine meadows and along streams in the Cascade and Olympic mountains, from Alaska to Oregon. From the Californian D. glaucum, this newly described plant differs principally in its smaller flowers, shorter spurs, shorter pedicels, the smaller whitish petals, glabrous anthers, shorter filaments, shorter follicles with a truncate apex, and the leaves being sparsely pilosulous above and less deeply lobed. It is possible, in fact, to identify sterile specimens by the leaf- characters alone. 86 MADRONO [Vol. 6 Sedum nesioticum sp. nov. Perenne, rosulatum, rhizomate horizontali lignoso; caulibus erectis, e basi excurvantibus, 10—30 em. altis, vulgo sub anthesi aphyllis; foliis plerumque basilibus, numerosis, aggregatis nec non imbricatis, rosulatis, lineari-lanceo- latis, teretibus vel subteretibus, succulentis, bene epapillatis, laete viridibus, nunquam scariosis, 1.5-3 cm. longis, integris; cymis effusis, 4-10 cm. latis, 3—7 dichotomis, floribus pulchre luteis, secundis; petalis discretis, lanceolatis, ca. 1 cm. longis; sepalis aequalibus, lanceolatis, acutulis, 5 mm. longis; staminibus quam petalis subbrevioribus; antheris 1 mm. longis; folliculis 6-7 mm. longis, apice subulato suberecto; seminibus obovoideis, levibus, 1 mm. longis. Perennial, tufted, with a woody horizontal rhizome; stems erect, curved upward at the base, 10-30 em. tall, usually leafless at flowering time; leaves chiefly basal, numerous, crowded but not imbricated, tufted, linear-lanceolate, terete or nearly so, succulent, smooth, not at all papillate, bright green, not becom- ing scarious, 1.5—8 cm. long, entire; cyme loose, 4-10 em. broad, 3—7-forked, the branches becoming divergent or even somewhat recurved, the short-pedicelled bright yellow flowers secund upon the branches; petals distinct, lanceolate, acute or acuminate, about 1 em. long; sepals equal, lanceolate, acutish, smooth, 5 mm. long; stamens slightly shorter than the petals; anthers 1 mm. long; follicles 6-7 mm. long, the subulate tips suberect; seeds obovoid, smooth, striate, 1 mm. long. BRITISH Corea rocky bluffs, Victoria, July 21, 1918, W. R. Carter; C. F. Newcombe 23 (G). Wasuinaton: islets, Gulf of Georgia, L. Ff. Henderson 1686 (type, G); Friday Harbor, S. M. § E. B. Zeller 808 (G); cliffs, Waldron Island, H. C. Cowles 475 (G, Mo). This newly described Puget Sound plant is not accounted for in Fréderstrém’s recent monograph (Acta Horti Gothoburgensis 5-10, App.:1930—-1936) or in the revision of the North American species by Britton & Rose (N. Am. FI. 22: 7-74. 1905). It is evidently related to Sedum stenopetalum Pursh, for which it has commonly been passing, but it is a larger plant than that species. It can be distinguished in the herbarium by the facts that the leaves and sepals are perfectly smooth, not at all papillate, and that the inflorescence is larger and with the branches more diver- gent. The petals are longer, acute or acuminate, but not mu- cronate. The carpels are somewhat larger. So far as is known at the present time S. nesioticum is confined to the islands of the northern part of Puget Sound and adjacent British Columbia, hence the specific name. Sedum stenopetalum Pursh is a perfectly distinct species growing in the mountains from Alberta to New Mexico, and extending as far westward as eastern Washington. University of Illinois, Urbana, September 26, 1940. 1941] CRONQUIST: IDAHO PLANTS 87 NOTEWORTHY PLANTS FROM IDAHO ARTHUR CRONQUIST During the summer of 1939 the writer had opportunity to make extensive collections in Clark and Fremont counties, Idaho. Several specimens apparently represent plants previously not known to occur in the state and others represent range extensions within the state. All numbers cited are the author’s own collec- tions and, unless otherwise noted, duplicates of each number have been deposited in the herbaria of the Utah State Agricultural College, Logan, (UtS), the Missouri Botanical Garden (MB), and the University of Idaho, Southern Branch, Pocatello. AGRopyron SauNnpERsit (Vasey) Hitche. is recorded by Hitch- cock (4) from only two localities, Veta Pass, Colorado, and Salt Lake City, Utah. It was collected in Fremont County three times during the season: along the Idaho-Wyoming border near Squirrel Creek east of Ashton, 1686 (UtS, MB); along railroad tracks at St. Anthony, 1773; subalpine north slopes of Mount Jefferson, southwest of Henry’s Lake, 1874. At the last mentioned station. the species was growing intimately with Agropyron Scribneri Vasey, but without apparent intergradation. | AGROPYRON ScRIBNERI Vasey is recorded by Hitchcock (4) from Montana, Wyoming, Utah, and Nevada, among other states, but not from Idaho. It was collected on subalpine slopes of Mount Jefferson, Fremont County (1873). Carex Hepsurni Boott is not recorded by Mackenzie (6) from Idaho although he mentions having seen specimens from all bor- dering states except Utah. The species was collected at 9000 feet on rocky, subalpine slopes east of Mount Jefferson, Fremont County (1891). CaREx occiDENTALIs L. H. Bailey is recorded by Mackenzie (6) from Utah, Nevada, and southward. The species was collected under an open stand of limber pine, at 9500 feet, on a high ridge above West Camas Creek, north of Kilgore, Clark County (1443) ; it was associated with Poa epilis Scribn. and with several other species of Carez. CARDAMINE uUNIJUGA Rydb. is recorded by Rydberg (8) from Wyoming and Montana. It was collected in standing water an inch or so deep at the edge of Big Springs, Fremont County (1599). SAXFRAGA OPPOSITIFOLIA L. is known in western United States from comparatively few stations. Fernald (1, p. 7) presents a range map showing collections apparently from the Wind River Mountains of Wyoming, Yellowstone Park and adjacent Montana, and again from what appear to be the Bitterroot Mountains, along the Idaho-Montana border. The specimen was found growing 88 MADRONO [Vol. 6 in dense extensive mats at 10,000 feet on north exposures just below the continental divide a mile southeast of the summit of Mount Jefferson, Fremont County. It was almost entirely past flowering at the time of collection (August 1) and hence is repre- sented by only a single scanty sheet (1906, UtS). EupuHorsia Esura L. is a recent introduction from Europe. Hanson and Rudd (8) record it from two localities in Idaho, one from the panhandle at the north end of the state, the other appar- ently from the vicinity of Kilgore, Clark County. It was collected along the roadside near Medicine Lodge Creek, north of Small, Clark County, about forty miles west of the Kilgore station (1931). PensTEMON Wuirtepi Piper subsp. Tristis Pennell & Keck is recorded by Keck (5) from the Salmon River twenty-three miles above Salmon City, and from several localities in Custer County, as far east as Mackay. This species was collected on an isolated sandstone outcrop in a limestone region north of Spencer, Clark County (1339); it was associated with Hackelia patens (Nutt.) Johnst., Phacelia leucophylla Torr. and Castilleja fasciculata A. Nels. It was not seen elsewhere in the area. BRICKELLIA OBLONGIFOLIA Nutt. subsp. typica comb. nov. B. oblongifolia Nutt. var. typica Robins. Mem. Gray Herb. 1: 104. 191%. BRICKELLIA OBLONGIFOLIA Nutt. subsp. linifolia (Robins.) comb. nov. 8B. oblongifolia Nutt. var. linifolia Robins., Mem. Gray Herb. 1 TOAD LOT Ge. This subspecies is recorded by Robinson (7, p. 104) from Colo- rado, Utah, and southward, the Idaho area supposedly being occu- pied by subsp. typica. A collection from the south side of basalt rocks in St. Anthony City Park, Fremont County (1474) is appar- ently subsp. linzfolia. A previous collection from the same station (687) is transitional to subsp. typica. It appears that subsp. lini- folia is the more southerly representative of the species and that the collections cited above represent an extreme northward exten- sion of its range. The habitat from which the specimens were taken simulates very closely what might be expected in hotter and drier more southern localities. In view of this difference in range var. linifolia Robins. may now be given subspecific status. CHRYSOTHAMNUS NAUSEOosUS (Pall.) Britt. subsp. GLAReEosus (Jones) Hall & Clem. is recorded by Hall and Clements (2) from southern Utah and northern Arizona. <A collection from the United States Sheep Station, near Dubois, Clark County (1995), agrees with this form in the glabrous achenes and puberulent involucres, but is taller than reported by Hall and Clements (6 or 8 dm. instead of 3 dm.). A previous collection (854) from the same station was made in 19387. The plant is not generally dis- 1941] CRONQUIST: IDAHO PLANTS 89 tributed through the area, and was found only on a lava reef near the station headquarters. CHRYSOTHAMNUS NAUsEosUs (Pall.) Britt. subsp. LEIosPpERMUS (Gray) Hall & Clem. is recorded by Hall and Clements (2) from southern Utah, southern Nevada, and the borders of California. A form having slightly wider leaves than are accorded to this sub- species, but otherwise agreeing fairly closely with the description given by Hall and Clements, was collected on the southern expo- sure of an extremely dry limestone cliff below the forks of Irvin Creek, northeast of Argora (1965). It will be noted that both of the Chrysothamni referred to above have glabrous achenes, an un- common character in C. nauseosus, and that both were collected in extremely dry rocky situations, more or less simulating the normal habitat of the plants in more southern areas. It is suggested that further exploration may reveal these supposedly southern forms to be more widespread than has been previously believed, occur- ring in hot dry areas often passed up by collectors as being unworthy of inspection. CurysoTHAMNUS Parryr (Gray) Greene subsp. Howarpu (Parry) Hall & Clem. is reported by Hall and Clements (2) from Wyoming and adjacent states, but not from Idaho. This sub- species is apparently represented by a collection from Red Con- glomerate Peaks, east of Argora, Clark County (1940). The plants, growing on bare rocks at 9500 feet, are much reduced and superficially resemble certain species of Haplopappus more than Chrysothamnus. The upper leaves are distinctly elongate and surpass the heads as in subsp. Howardii but the leaves are more nearly glabrous as in subsp. attenuatus (Jones) Hall & Clem. Other characters are more nearly those of subsp. Howardi, and the collection is with some hesitancy so referred. Intermountain Herbarium, Utah State Agricultural College, Logan, March 7, 1940. LITERATURE CITED 1. Fernatp, M. L. Recent discoveries in the Newfoundland flora. Rhodora 35: 1-16. 1933. 2. Hart, H. M. and F. E. Cirements. The phylogenetic method in taxonomy: the North American species of Artemisia, Chrysothamnus, and Atri- plex. Carnegie Inst. of Wash., Washington, D. C. 1923. 3. Hanson, H. C. and V. E. Rupp. Leafy spurge, life history and _ habits. North Dakota Agric. Exper. Sta. Bull. 266: 1-24. 1933. 4. Hircucocx, A. S. Manual of the grasses of the United States. United States Dept. of Agric., Misc. Publ. no. 200. 1935. 5. Kreck, D. D. Studies in Penstemon VI. The section Aurator. Bull. Torr. Bot. Club 65: 233-255. 1938. 6. Mackenzie, K. K. Cyperaceae. North American Flora 18: 1-478. 1931- 1935. 7. Rosinson, B. L. Monograph of the genus Brickellia. Mem. Gray Her- barium 1: 1-151. 1917. 8. Rypgerc, P. A. Flora of the Rocky Mountains and adjacent plains. New York. 1917. 90 MADRONO [Vol. 6 THE ALASKA CEDAR IN CALIFORNIA Hersert L. Mason Many rumors have been current relative to the occurrence of both the Alaska cedar, Chamaecyparis nootkatensis (Lamb.) Spach, and the western larch, Larix occidentalis Nutt., in California. These rumors have been particularly tantalizing since the descrip- tive evidence seemed so convincing. However, they were not vouched for by specimens. Recently, a collection from north- western California clearly demonstrates the occurrence of Chamae- cyparis nootkatensis in our flora. Mr. Oliver V. Matthews of Salem, Oregon, collected the species in 1989 on the northeast slope of Mount Emily in northwestern Siskiyou County and Dr. Doris Gillespie Niles of Humboldt State College, reports having seen it on Little Grayback farther to the west. Both localities are a little over two miles south of the Oregon line. Specimens of the Mount Emily material are deposited in the herbarium at Rancho Santa Ana Botanic Garden, Anaheim, California. Mr. Matthews reports that at Mount Emily the plants are associated with Abies shastensis, Picea Breweriana, Pinus monticola, Libocedrus decurrens ifolia. i and Larus brevifolia Department of Botany, University of California, Berkeley, February 15, 1941. REVIEWS The Ferns and Fern Allies of Wisconsin. By R. M. Tryon, N. C. Fassett, D. W. Duntop, M. E. Diemer. Frontispiece. Pp. v+158 with 214 figures and 76 maps. Published by the Department of Botany, University of Wisconsin, Madison. May, 1940. Price $1.00. The work treats three families of ferns and five families of fern allies as they occur in Wisconsin. The introduction contains directions for the use of the keys and closes with a well directed plug for the American Fern Society. A brief statement of the meteorology, geology, and physiography of Wisconsin precedes the taxonomic treatment. The keys are replete with references to text figures illustrating the terminology, a practice which should facilitate their use. The illustrations are largely halftones, most of which are from excellent photographs; a few have suffered somewhat at the hands of the printer in that their blackness often obscures the detail. Drawings are well done and excellently re- produced. Distribution maps, which are liberally used, give a clear picture of the range of each species. In some cases the limestone areas are mapped showing the correlation between the distribution of the species and the occurrence of limestone. The value of such maps cannot be overestimated. They serve a very useful purpose in interpreting the plant geography of the region as well as in directing botanists to areas where further field work is needed. The chores of the four-fold authorship are divided between a fern specialist, a director of the project, an illustrator 1941] MASON: ALASKA CEDAR 91 PuaTe 7. CHAMAECYPARIS NOOTKATENSIS. A group of small trees (fore- ground) on Mount Emily, Siskiyou County, California. Photograph by Oliver Matthews. 92 MADRONO [Vol. 6 and a photographer. Over all, an angel spread his wings—Martin J. Gillen, “a loyal alumnus, an enthusiastic conservationist.” We wish to encourage more such works and more such angels.—Her- BERT L. Mason. Forests and Trees of the Western National Parks. By Haroup E. Battey and Vireinia Lona Baitey. Pp. x +129 with 71 text figures. Conservation Bulletin no. 6, National Park Service. United States Government Printing Office, Washington, D. C. 1941. Paper, $.25. This valuable contribution to the literature of the national parks begins with a short introductory discussion of forest conser- vation, life zones, structure and functions of a tree and plant asso- ciations. The three general forestal regions (Pacific Slope, Rocky Mountain, Southwestern) in which the western national parks are located are briefly characterized. The forest associa- tions and topographic features of the western parks, including’ those of Alaska and Hawaii, are then treated in some detail. Field keys to the genera and species of trees, non-technical de- scriptions and general information amount to approximately one- half of the content. A well-selected bibliography and an index to the common and scientific names complete the volume which is printed in clear type and beautifully illustrated with half-tones from photographs. The authors are to be congratulated on this attractive and readable booklet which will be very much appreci- ated by botanists and foresters as well as by the vacationing public —EtTHEL Crum. A Manual of the Higher Plants of Oregon. By Morton E. Peck. Pp. 1-866, with 1 plate, 1 map and 90 figures. Binford & Mort, Portland, Oregon, 1941. Price $5.00. — The appearance of a usable manual covering the flora of any state which has not previously been so endowed is an event of the utmost interest to all students of plants. Especially is this true when the book is as thoroughly excellent and generally attractive as this one! There has, heretofore, never been any manual or combination of manuals which would permit satisfactory deter- mination of plants from all parts of Oregon. Furthermore, no one previously has attempted to gather into one volume the vari- ous entities described from the state since the appearance of Howell’s “Flora of Northwest America,’ about forty years ago. A very rough tabulation indicates that approximately 250 of these recently described entities, including two genera, collected into the present book, have never before been available in a manual or flora. If the author had performed no other service than to glean these from the scattered literature, characterize them fully and key them out, he would merit unstinted gratitude. Even those persons who have had considerable experience with the vegetation of the area in question will, I am sure, be amazed at the size of its flora, as revealed in the present volume. 1941] REVIEWS 93 There are included 118 families, 701 genera and 3203 species— more than three-fourths the number of species recorded from Cali- fornia in Jepson’s “Manual,” in 1925. The area of Oregon is only three-fifths that of California. This suggests a degree of diversity for the flora of Oregon at least equal to that of the Cali- fornia flora, which has long been famous for its high degree of diversity coupled with remarkable endemism. An introductory survey of the physiographic and floristic fea- tures subdivides Oregon into the following nine major “‘plant areas’: northern coast, northern coast mountain, Willamette Valley, Rogue-Umpqua, southern coast, southern coast mountain and Siskiyou, Cascade, eastern Oregon (with five subordinate sec- tions) and Blue Mountain. These represent the remarkable blending of at least six floristic elements: the northwestern conif- erous forest, the Cascade, the Rocky Mountain, the Great Basin, the Californian and the Klamath-Siskiyou, a fact which helps to explain the large number of species native to the state. Each area is briefly summarized as to topography, geology, climate and plant assemblages. Important concentrations of endemic species are shown to occur particularly in the Siskiyou and southern coast mountains, Wallowa and Steens mountains, and in the gorges of the Columbia and Snake rivers. This excellent summary is made particularly valuable by the author’s thorough familiarity over a long period of years with nearly every corner of the state. The book is attractively bound in green cloth and is so firmly constructed that it should hold up well even under intensive field usage. Each family, genus and species is carefully keyed out, and for each species there is provided an English name, a full description, and a brief expression of its local and general range. The author bravely discards the traditional “Oregon Myrtle,” often claimed locally to be “native only to Oregon and the Holy Land,” for the more accurate “California Laurel.’ All sub- specific entities are given the status of “variety,” necessitating most of the new combinations proposed. The International Rules _are strictly adhered to in the matter of capitalization of all specific epithets which are derived from personal names, or are generic or vernacular names. There is a notably large glossary of tech- nical terms accompanied by ninety helpful illustrations of plant parts; thirty-six pages of index to families, genera and species, including both technical and English names, and their synonyms, conclude the volume. Mr. J. William Thompson has contributed the entire manu- script for the Pteridophyta and various specialists have prepared keys, or have aided with determinations or advice. The bulk of the taxonomic treatment, however, represents the author’s pains- taking effort to weigh everything that has been written about Ore- gon plants against his own sound taxonomic judgment and his unrivalled knowledge of the area of which he writes. The manual is the culmination of many years of devotion to the flora 94 MADRONO [ Vol. 6 of the state, and is based largely upon his own extensive collec- tions, now permanently housed at Willamette University, Salem, Oregon. From the fine quality of the book, one would not suspect the handicaps under which it was conceived and carried out. Much of the bibliographical work had to be concentrated into infrequent visits to the larger botanical centers, and herbarium study had necessarily to be limited chiefly to the author’s personal collec- tions. Dr. Peck’s achievement is a credit to his own ability and perseverance, and should greatly encourage in the Pacific North- west the study of the science to which he has given so many years of his life. Systematic botany in Oregon will henceforth be sharply divided into the period before and that after the appear- ance of this book. The author has provided a point of reference and of departure for all future work in the state, an achievement whose value can scarcely be overestimated. This manual will stand as a perpetual challenge to those in the smaller institutions and elsewhere, who imagine themselves unable to carry on pro- ductive work because of the lack of facilities or funds, or because of their isolation from the centers of scientific activity. Such obstacles have failed to prevent Dr. Peck from making a bril- liantly successful contribution to systematic botany.—L. Con- STANCE. NOTES AND NEWS Fietp Work oF THE Bureau or Piant INpDusTRY IN NEvapa. For years it has seemed desirable to gather up the rapidly vanishing plant lore of the Indians in Nevada. Stimulated by the interest of officials of the Office of Indian Affairs in Nevada and by the University of Nevada, a special project was initiated by the Bureau of Plant Industry which is interested in the plants them- selves. From this have ramified various excellent projects. Since a wide knowledge of the state flora was necessary for the basic organization of the other data, the collection and classifica- tion of the flora became almost the primary concern. The work has continued for four years, beginning in 1937, and during that time several subsidiary projects have been undertaken. During the first year there were six groups of collectors, most of them hired merely as collectors, who were sent out in pairs to different sections. There were six other collectors who travelled alone or with an assistant. Among these, under the general direc- tion of W. A. Archer of the Bureau of Plant Industry were: Roy A. Allen, Tim Louise Breene, George E. Franklin, Jr., Frank S. Goodner, Newell F. Hancock, William G. Henning, James R. Hen- richs, Philip A. Lehenbauer, Ira LaRivers, Lauritz Lund, William E. McKenzie, Mrs. Edith V. A. Murphey, Loretta R. Miller, Ben- jamin O. Moore, Norman E. Nichols, Benjamin A. Rives, Harry Sampson and Perey Train. After the first year the collections were made by Dr. Archer and Mr. Train. 1941] NOTES AND NEWS 95 The total herbarium collection from the project is 13,798 items, many in duplicate so that much has been left for exchange after the first set was deposited at the University of Nevada and the second in the herbarium of the United States National Arbore- tum at Washington, D. C. The material is also represented at the Gray Herbarium, New York Botanical Garden, St. Louis Botanical Garden, University of California, Stanford University, Catholic University of America, and the Field Museum of Natural History. The labels which appear on these plants give no indica- tion that the work was done under the supervision of the United States Department of Agriculture so that the material will need to be identified merely by the year and the name of the collectors, which have been indicated above. Practically all of the identifications have been done by S. F. Blake, C. O. Erlanson, O. M. Freeman, F. R. Fosberg, F. J. Her- mann, Robert F. Martin, C. H. Muller and J. R. Swallen all of whom are members of the staff of the Division of Plant Explora- tion and Introduction. Certain special groups have been identi- fied by experts outside the Division: Cruciferae by C. Leo Hitch- cock, Saliz by Carleton R. Ball, Gilia by Edgar T. Wherry, Umbel- liferae (in part) by Mildred Mathias, and Scrophulariaceae (in part) by Francis W. Pennell. More recently, as a result of this work, the Division of Plant Exploration and Introduction has started issuing a mimeographed series known as “Contributions toward a Flora of Nevada.” This work is being done by the botanists of the Division in cooperation with the Work Projects Administration of Nevada. It is planned to continue these contributions until all of the plant families of Nevada have been covered. Up to the present there have been about thirty contributions completed, most of them being smaller families, but work is in progress on some of the larger families.— W. ANDREW ARCHER. THREE ALIEN PLants NEw To OreGon. The following notes are based on collections made in the vicinity of Portland, Multnomah County, Oregon. The three plants appear not to have been re- ported hitherto from North America. The specimens cited are deposited in the Gray Herbarium, Harvard University (G), the University of California Herbarium (UC), or the herbarium of the State College of Washington (WS). Cnipium Monnieri (L.) Cuss. Presumably on ballast, Albina, October 25, 1900, Suksdorf 2926 (UC, WS). Adventive from Asia, and apparently not persisting here. MESssERSCHMIDIA siBIRIcA L. (Determined by Ivan M. John- ston.) On ballast, Peninsula Docks, bank of Willamette River at St. John, July 10, 1936, Constance 1801 (G, WS). The species was obtained from a similar situation by Suksdorf, many years ago, and is apparently persisting but not spreading. 96 MADRONO [Vol. 6 LapsaNa APOGONOIDES Maxim. (Determined by G. L. Stebbins, Jr.) Forming ground cover with Lysimachia Nummularia under Fraxinus and Salix at edge of Carex bog, four miles north of Bur- lington ferry, Sauvies Island, May 28, 1940, Constance & Beetle 2677 (UC).—L. Constance. PROCEEDINGS OF THE CALIFORNIA BOTANICAL SOCIETY March 15, 1941. The annual dinner of the California Botani- cal Society was held at the Berkeley Women’s City Club, Berkeley, on Saturday evening, at 6:30 o’clock. Seventy-three members and their guests were present for the banquet. Professor E. B. Babcock, President, acted as toastmaster, and introduced Dr. D. H. Campbell, Dr. G. J. Peirce, Professor H. E. McMinn, Dr. Ira L. Wiggins, and Mrs. S. B. Parish, who sat at the speakers’ table. Professor H. E. McMinn, Professor of Botany, Mills Col- lege, Oakland, was the speaker of the evening. The subject of his address was “‘Botanical Developments in the East Bay Re- gional Parks.” He traced the history of the acquisition and devel- opment of the parklands owned by the East Bay Metropolitan Area, emphasizing particularly their botanical and recreational possibilities. The thoroughly enjoyable evening was in charge of the program committee: Dr. G. Ledyard Stebbins, Jr., chairman; Dr. Adriance S. Foster, and Miss Beryl Schreiber, with valuable assistance from Mrs. E. B. Babcock, Mrs. H. L. Mason and Mrs. Lincoln Constance. April 17, 1941. Third annual ‘Plants and Specimens” meet- ing, 2093 Life Sciences Building, University of California, Berke- ley, at 7:45 P. M., in charge of Dr. G. Ledyard Stebbins, Jr., and Miss Beryl O. Schreiber. The President, Professor E. B. Bab- cock, occupied the chair. Short talks about the specimens or exhibits displayed were given as follows: A new tetraploid Layia, Dr. David D. Keck; Proteaceae, Miss Alice Eastwood; Vegetation type-mapping by aerial photography, Mr. A. E. Wieslander; The phylogeny of the closed-cone pines, Dr. H. L. Mason; The closed- cone pines of Ano Nuevo Point, Mr. Leo Koch; A new hexaploid Solanum, Dr. E. F. Paddock; Artificial production and selection of polyploid grasses for range re-population, Dr. Stebbins. A large number of attractive exhibits, ranging from algae, fungi and lichens to seeds, succulents and unusual cultivated plants were contributed by many exhibitors, including the following: Professor Babcock, Dr. and Mrs. Alan A. Beetle, Mr. and Mrs. Anson F. Blake, Dr. Marion S. Cave, Mr. Yale Dawson, Mr. George B. Furniss, Dr. John W. Gilmore, Dr. Frank Gould, Dr. A. W. Herre, Dr. Robert F. Hoover, Mr. Frank A. Leach, Mr. and Mrs. Forrest McAbee, Miss Elizabeth Morse, Miss Catherine Roberts, Mr. Jack Whitehead and Dr. Ira Wiggins.—L. Constance. MADRONO A West American Journal of Botany A quarterly journal devoted to important and stimulating articles dealing with plant morphology, physiology, taxonomy, and botanical history. These volumes should be a part of every botanist’s li- brary and should be made accessible to students of all universities and colleges. Volume I, 1916-1929. . . $5.00 Volume II, 1930-1934 .. 5.00 Volume III, 1935-1936 . 5.00 Volume IV, 1937-1938 . 5.00 Volume V, 1939-1940 . . 5.00 Single numbers....... 0.75 The subscription price of MADRONO is $2.50 per year. We solicit your pat- ronage, Address all orders to: William Hiesey Bus. Mer. Carnegie Institution of Washington Stanford University, California : a sg GNtAN INSTT; LUME VL“. “ee (hia ONS NUMBER 4 f Hf ; - é i * Ay t> 4 1, V4 We ¢ WN ff iat) Be Ne ff ~~ Gsfrarar ary =~ - ~~ é NAR i] sf ~~ 2 a ey ened Lee pues aD get gas, sarees TS MADRONO A WEST AMERICAN JOURNAL OF BOTANY Contents URTHER STUDIES ON MonotropoweEakE, Herbert F. Copeland ............. 97 DOCARPUS GRACILIOR IN CULTIVATION, John T. Buchholz .................. 119 Tue Taxonomic Status or Microstreris GREENE, Herbert L. Mason ..... ™. 122 its HoracantHow Puants or NortuH America, Cornelius H. Muller ..... 128 Pret CR? Oe RAM nese te etn, 2 be eta tatiana Baaame yas oe tak pee, "133 R. Mm. JSG EIS SOO SOA eR Noe SAE CAR ML Ht LOSME eS Anup in CAT) ois Lees 137 "B Rueelaid): ; Fortunato L. Flemera sa de la Flora del Cuzeo (J. F. Macbride) ; Ivar Tidestrom and Sister Teresita Kittell, 4. Flora of Arizona and New Mexico (Herbert L. Mason) ..... LIRA Nga aA 142 " Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania October, 1941 MADRONO A WEST AMERICAN JOURNAL OF BOTANY Board of Editors Hersert L. Mason, University of California, Berkeley, Chairman. LeRoy Asrams, Stanford University, California. Epcar ANpDERSON, Missouri Botanical Garden, St. Louis. Lyman Benson, University of Arizona, Tucson. Hersert F. Copetanp, Sacramento Junior College, Sacramento, California. Ivan M. Jounston, Arnold Arboretum, Jamaica Plain, Massachusetts. Miuxprep E. Maruias, University of California, Berkeley. Bassett Macume, Utah State Agricultural College, Logan. Marion Ownsey, State College of Washington, Pullman. Secretary, Editorial Board—EtTuHeEt Crum Department of Botany, University of California, Berkeley Business Manager—Wiuti1aAm Hirsey 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 an 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 proofs are returned. Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania, for the CALIFORNIA BOTANICAL SOCIETY, INC. President: Ernest B. Babcock, University of California, Berkeley. First Vice-President: Roxana S. Ferris, Stanford University, California. Second Vice-President: Palmer Stockwell, Institute of Forest Genetics, Placerville, California. Treasurer: Wiiliam Hiesey, Carnegie Institution of Washington, Stanford University, California. Secretary: Lincoln Constance, Department of Botany, University of California, Berkeley. Annual membership dues of the California Botanical 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. 1941] COPELAND: MONOTROPOIDEAE 97 FURTHER STUDIES ON MONOTROPOIDEAE Hersert F. CopeLtanp This paper continues a series (5, 6, 7, 8, 9), of which the account of Sarcodes by Doyel and Goss (12) is to be considered a unit. It records observations on Pterospora, Hypopitys, and Monotropa, and adds to the previous account of Hemitomes (New- berrya). : It has been a pleasure to acknowledge from time to time the unstinted cooperation of various institutions and individual cor- respondents. The following have facilitated the preparation of the present contribution: the Herbarium of the University of California; the Dudley Herbarium of Stanford University; the Herbarium of the California Academy of Sciences; Dr. W. L. Jepson; Mr. Willman Spawn; Dr. W. H. Camp; Dr. P. L. Zimmer- man; and the Oregon Biological Supply Company. PTEROSPORA ANDROMEDEA Nuttall Pterospora andromedea was described by Nuttall (25) as col- lected “In Upper Canada, near the Falls of Niagara. Mr. C. Whitlow.” There has been essentially no nomenclatorial confu- sion as to this plant; no segregation of species or varieties has been proposed; Small (28) cites a single obscure synonym. The plant is common in the mountain ranges of western North Amer- ica; rare eastward to the region of the type locality. The material studied was collected from time to time at Jones- ville, Butte County, California, at an altitude of about five thou- sand feet. There the plant shares the habitat of Sarcodes and Pleuricospora, in forests of fir (Abies concolor). As compared with its congeners, Pterospora emerges from the ground and flowers noticeably later in the season; it is usually in full anthesis late in July. The shoots come up from globular masses of roots. It is noticeable that whereas they emerge in the neighborhood of dead shoots of the previous year, they are not in immediate contact with such dead shoots, nor at a distance to be measured in centi- meters, but a meter or more away. Jepson (19) believes that the plant is monocarpic, and it is probable that he is correct. There is, however, the possibility that the scattered shoots come up from long roots which have emerged from the masses. This is sug- gested by the behavior of the generality of monotropoid plants, in which the genetic individuals are polycarpic, forming shoots year after year as adventitious buds on the same root system. A positive determination of this matter could be attained only by determining the course of individual roots in a mass of humus, and I have not undertaken it. ManroXo, Vol. 6, pp. 97-144. October 15, 1941. a ‘eo a aus 98 MADRONO [Vol.6 The standard accounts of the gross structure of the shoot, as by Jepson and Small, leave essentially nothing to add. The tall and slender stems, densely glandular, are of a light purplish red color. The greater part of each shoot is a rather lax bracted raceme, usually but not always exhibiting orthodox phyllotaxy. The recurved glandular pedicels bear no bractlets. The five glandular sepals are separate. The five petals form a glabrous sympetalous urceolate corolla, yellowish in color, becoming char- taceous in age. Each of the ten stamens bears two horns on the back of the anther. The globular ovary is belted at the base by a nectary from which ten evenly spaced lobes project between the bases of the filaments. Internally, the ovary is five-chambered below, one-chambered above, filled by massive placentae bearing numerous ovules. The stigma is obscurely five-lobed, the lobes opposite the petals, that is, at the ends of the carpels. MacDougal has described the anatomy of the vegetative parts. The stem includes a cylinder of bundles so closely packed as almost to be continuous; around this there is a well developed continuous sheath of fibers. The vascular supply of the flower (pl. 8, fig. 1) enters the re- ceptacle as acylinder. From this there depart radially, first, five sepal bundles, and then, alternating with them, five petal bundles. There are no gaps above these bundles. Five stamen bundles arise as branches from the upper sides of the petal bundles. Above the departure of the petal bundles the stele breaks up into a ring of ten bundles. Of these, the five which are opposite the sepals bend outward and fork periclinally; the outer branches constitute the supply to the five remaining stamens; the inner ascend the ovary wall in the planes of the septa, and are to be interpreted as fused pairs of lateral bundles of adjacent carpels. The five remaining bundles enter the ovary in the planes of the locules, being the planes of the petals and carpels; each of them is a fused pair of ventral bundles of a single carpel. They branch out into the placentae and disappear. In several features the vascular system just described is peculiar. No other monotro- poid plant is known to be without carpel dorsals (though they are not well differentiated from carpel laterals in Sarcodes) ; in none do the placental bundles lie in the planes of the petals. Here it is as though the proper placental bundles had swung outward into the ovary wall, and the carpel dorsals inward, into the placentae. In most plants of the group, the style is supplied by the carpel dorsals, which ascend in the thin bands of tissue between the ridges projecting into the style channel. Sarcodes is exceptional in that the placental bundles supply the style, ascending within the ridges. In Pterospora, lacking carpel dorsals and having the placentals in a peculiar position, the style is without vascular tissue. As is known, the anther (pl. 8, figs. 2-7) projects horizontally toward the style from the summit of the filament, and bears two 1941] COPELAND: MONOTROPOIDEAE 99 horns, inserted respectively on the two sides of the insertion of the filament and projecting toward its base. A vascular bundle runs from the summit of the filament to the inward end of the anther; by this it is known that the inward end is distal, the end bearing the horns proximal, the upper side dorsal, and the lower side ventral. There are four parallel horizontal pollen sacs; the dorsal ones are larger than the ventral. A cross section shows the epidermal cells extended into conical points on the dorsal and ventral, but not the lateral, surfaces. Between the epidermis and the tapetum there are some two or three layers of wall cells; near the horns a few of these cells develop reticulate lignified thicken- ings, as in the endothecium of a typical flowering plant. A similar structure has been reported in Sarcodes, and will be re- ported below in Monotropa; it is less extensive in Pterospora than in these other genera, being apparently merely a relic structure. The cells of the tapetum become binucleate. The pollen grains are four-grooved. Dehiscence of the anther begins by two vertical slits, each of which crosses the proximal ends of the two pollen sacs of one lobe of the anther. Formation of these slits amounts to the same thing as the dehiscence of the anthers of Sarcodes or Monotropsis, being, as I take it, the typical dehiscence of the anthers of Eri- cales. Subsequently two secondary slits develop from the middles and at right angles to the primary ones, that is, in the plane between the two pollen sacs of each lobe of the anther. By a general shrivelling of the walls of the anther, accompanied by growth of the connective, these secondary slits gape widely; it is through them that most of the pollen escapes. Drude’s (138) figure of the fully dehisced anther is accurate, though it gives a false impression that the connective becomes torn in a horizontal plane from proximal to distal. The internal surface of the ovary wall is covered by two layers of well developed fiber-like cells. A complete series of stages of the development of the seed has not been seen, and there is nothing to add to my earlier notes (4). The embryogeny appears to be altogether typical of the group. The integument is of two layers of cells. The wing on the seed is of two layers of cells. It commences to form, by proliferation of the epidermis of the integument at the chalazal end, before the embryo sac is fully developed. Hyporitys Monortropa Crantz Only two species of monotropoid plants were known to Lin- naeus (23). One, known in English as the pine-sap, occurs on all continents of the north temperate zone; the other, the Indian pipe, occurs in North America and eastern Asia, but not in western Asia and Europe. The oldest Latin designation of the pine-sap seems to be Orobanche quae hypopithis dici potest, of Bauhin (1671). It is of course no Orobanche; Tournefort (1706) 100 MADRONO [Vol. 6 gave it as name the adjective Orobanchoides ; Dillenius (1719) took up the substantive designation recorded by Bauhin, as Hypopitys. The Indian pipe, first recorded by Plukenet (1671) under Oro- banche, was named Monotropa by Gronovius (about 1740). Lin- naeus included both species in one genus, for which he used the name Monotropa, the species becoming respectively M. Hypopithys . and M. uniflora. The breach of priority as to the generic name was immediately protested by Hill (16): “Linnaeus takes away its received name hypopitys and calls it monotropa.” Because he was the first after 1753 to use Hypopitys as the name of a genus, Hill is cited as authority for it; this in spite of the facts, that he did not originate it, and that he did not use binomials and cannot be cited as authority for any of the species. Binomials under Hypopitys' were first made by Crantz (10), the pine-sap becom- ing H. Monotropa and the Indian pipe H. uniflora. Inasmuch as the first post-Linnaean authors gave to Hypopitys exactly the extent which Linnaeus had given to Monotropa, it might be held that the two names are exact synonyms, and that Hypopitys is not available as the name of any genus. On the other hand, the pre-Linnaean history shows that the two names are based on different types, and that if the pine-sap is placed in a different genus from the Indian pipe its name is Hypopitys. Such in effect was the conclusion of the pre-type-system authors Nuttall (25) and Bentham and Hooker (1); such was the conclu- sion of Small (27), who made the combination Hypopithys Hypo- pithys. Repeating binomials being excluded by our rules, we must accept the first specific epithet published after 1753; this yields, as has been shown, the combination Hypopitys Monotropa. We might be glad to reject this combination as a matter of taste; it has gone almost completely unaccepted since its original pub- lication; but the rules are designed to spare us the responsibility for a choice. I follow Kamienski (21) and Domin (11; this work has been a most valuable guide to the history and literature) in recogniz- ing only one species of Hypopitys. There are considerable varia- tions, and Small (28) has recognized five species in North America alone; but these variations seem so inconstant as to make the recognition even of varieties a critical matter. The available material preserved in liquid has included sev- eral roots, stems, and flowers collected by Dr. W. H. Camp, in Oregon, Ohio, and Tennessee, and two shoots with nearly ripe fruit collected by Mr. Willman Spawn in Rock Creek Park, Wash- ington, D. C., in July of 19388. 1'Variations in the spelling will be noted. I have not consulted the pre- Linnaean publications. Linnaeus wrote Hypopithys, Hill and Crantz Hypo- pitys. These are mere variant spellings; but we are forbidden by rule to meddle with them. In using the word as a specific epithet, we must follow Linnaeus; in using it as a generic name, we must follow Hill. 1941] COPELAND: MONOTROPOIDEAE 101 Nothing is here added to knowledge of the gross structure. The shoots originate endogenously in roots, a mass of which con- stitutes the permanent organ of the plant. Domin, after a long discussion of the literature, concludes that the underground struc- tures are not true roots, but an axis not differentiated as root or stem, to be called the Prokaulom (anglicizable as procaulon). I have not examined these structures, but the conclusion is inescap- able, from Kamienski’s (20, 21) description and from what is known of the other monotropoid plants, that Domin is mistaken. As Christoph (3) has shown, the occasional more or less com- plete suppression of some of the characters of roots—the cap and endogenous branching—depends on the presence of mycorrhiza. The same effacement of character appears in the roots of other plants when they are beset with mycorrhiza. The shoot is usually yellowish and more or less pubescent. The upper part of it constitutes a bracted raceme. When it emerges from the ground, the raceme is bent to one side, and the buds or flowers are crowded and more or less pendant; later the axis and pedicels become erect and the flowers or fruits become separated. At its maximum the shoot is rarely twenty-five centi- meters tall. The lateral flowers (pl. 8, fig. 8) stand in the axils of bracts whose margins vary from entire to lacerate. There are no bract- lets; four sepals, a lower pair placed laterally and an upper pair placed dorsally and ventrally with regard to the flower; four separate petals with saccate bases, alternating with the sepals; eight stamens; a nectary with eight horn-like lobes arranged in pairs which clasp the bases of the petalad stamens; a pistil, the ovary four-chambered below, one-chambered above, the stigma obscurely lobed, the lobes opposite the petals. Variations in the proportions of ovary and style, as well as in color, pubescence, and the indentation of the margins of the bracts, have been util- ized for subdivision of the species. Older accounts definitely described the terminal flower as pentamerous. I have not found sucha flower. I believe that in the few shoots preserved in liquid which I have examined the ter- minal flower has been suppressed. The highest flower has been tetramerous, the pedicel embraced by two bracts instead of one (evidently as a result of shortening of the last internode, the one above the insertion of the flower), the lower pair of sepals some- what removed from the flower. Domin cites many authors who have disagreed as to whether the outer envelope of the flower is really a calyx, and its segments sepals: Eichler is the chief authority in the affirmative, Baillon in the negative. He quotes observations of Irmisch (17) and Wydler (30) and gives his own, to the effect (a) that the upper pair of leaves of this envelope are, one or both of them, often suppressed; (b) that the lower pair are often somewhat with- drawn from the base of the flower; (c) that these leaves, espe- 102 MADRONO [Vol.6 cially the lower pair, often have buds in their axils. He con- cludes that they are not true sepals, but elevated bractlets in proc- ess of conversion into sepals. It seems to me unreasonable to recognize a calyx in course of coming into existence in any group as advanced as Ericales: rather, any irregularities are to be inter- preted as matters of degeneration. Since Allotropa is the only positively asepalous genus of the monotropoid group; since only Monotropsis and sometimes Allotropa have definite bractlets; we may interpret the facts assembled by Domin in some such fashion as this: the structures of Hypopitys now under consideration are positively sepals; they are affected by a tendency to degenera- tion; it is possible that a tendency to produce bractlets, almost completely extinct in this genus, retains enough strength to affect the course of the degeneration. Kamienski has accounted in full for the anatomy of the vege- tative parts; I have here only to describe the vascular supply to the flower. One bundle from the circle in the stem turns out- ward. It becomes flattened tangentially and presently splits into three branches, of which the middle one supplies the bract while the two on the sides swing together and unite as a cylinder ascend- ing the pedicel. All this is quite the same as in the genera previ- ously studied. The cylinder of vascular tissue ascending the pedicel becomes compressed in the dorso-ventral plane, so as to approximate a four-sided prism (pl. 9, fig. 10). From each of its faces there departs a sepal trace, leaving a small gap or none. The traces to the lateral sepals depart at a much lower level than those to the dorsal and ventral sepals; this is in harmony with Domin’s observations as to the relative positions of the sepals. The petal bundles emerge as broad bands from the angles of the prism; each promptly forks into three, of which the one in the middle is the smallest and descends under the sac of the petal, while the larger lateral ones ascend past the sac. The petalad stamen bundles are not fused with the petal bundles, but are closely associated with them; each originates as a pair of bundles at the sides of a petal bundle, the pair drawing together and unit- ing above the petal bundle. The sepalad stamen bundles emerge at the edges of the siphonostele as it breaks up. The carpel dor- sals, well developed in the ovary walls and style, are only with difficulty traced to their origin; the feebly developed provascular strands that lead into them seem to originate typically as paired strands beside, above, and resembling the petalad stamen bundles. Foster (15) has recently quoted Gregoire to the effect that floral EXPLANATION OF THE Ficures. Puate 8. Piate 8. PTEeROSPORA ANDROMEDEA. Fic. 1. Model of the vascular system in the receptacle X 50. Ca, sepal bundles; Co, petal bundles; St, stamen bundles; Cl, carpel laterals; Pl, placental bundles. Fics. 2, 3. Juvenile stamens, x 10. Fic. 4. Longitudinal section of juvenile stamen, x50. Fic. 5. Cross section of juvenile stamen at the plane marked zw in fig. 4, X50. Fic. 6. Area marked x in fig. 5, X400. Fic. 7. Cross section of dehisced stamen, X50. Hypoprirys MONOTROPA. Fic. 8. Old flower in which the fruit is nearly ripe, x 5. 1941] COPELAND: MONOTROPOIDEAE 103 P{ Piate 8. PTEROSPORA ANDROMEDEA; Hyporirys MONOTROPA 104 MADRONO [Vol. 6 leaves are distinguished from vegetative leaves by acropetal devel- opment of the vascular supply. The carpel dorsals of Hypopitys (and likewise, as will be seen, of Monotropa and Hemitomes) con- stitute an exception to this principle. The breaking up of the siphonostele finally yields one bundle to each lateral placenta, and — two each to the dorsal and ventral placentae. This seems to be an outcome of the bilateral character of the whole vascular sys- tem. It is as though the ventral bundles of adjacent carpels were fused at the sides of the flower, but not at the front and back. It is regretted that no anatomical study has been made of young stamens. The anther opens by two vertical slits at the outer, presumably proximal, end; these slits meet above, separat- ing a small outer valve from a large inner one; and soon the valves liberate the pollen by swinging widely apart. Young stages and old ones (pl. 9, figs. 11, 12) respectively agree exactly with corresponding stages of Pityopus as illustrated by Eastwood (14). It is particularly regretted that the position of the pollen sacs was not ascertained; though it may be presumed that there are four in each anther, lying horizontally, each slit of the anther crossing the ends of two of them. The pollen grains are two- grooved. The fruit is a capsule, its inner surface covered by a single layer of elongate cells not distinguished by staining reactions. The development of the seed has been described in detail by Koch (22). I have seen only one stage, conforming well to Koch’s account, and so beautifully clear that I could not refrain from drawing it (pl. 9, fig. 9). MonoTROPA UNIFLORA L. Of the name of this genus and species enough has been said above. The genus is apparently monotypic; no variation even of varietal rank is recognized as occurring within the United States. Monotropa coccinea Zuccarini, of Mexico and Central America, and M. australis Andres, of Colombia, were treated as varieties by Domin. The available material preserved in liquid has included six collections, as follows: (1) Three shoots purchased some years ago from the New York Biological Supply Company, as a museum specimen, without collection data. (2) Three shoots presented by the Oregon Biological Supply Company; collected by R. E. Griffin, Bullrun, Oregon, July 11, 1989. The fluid preserves the white color of the plants, and makes specimens excellent for mu- seum use rather than for sectioning. (3) A number of shoots presented by the Herbarium of the University of California, with- out collection data. (4) One shoot, with roots, furnished by Mr. Willman Spawn; collected in Rock Creek Park, Washington, D. C., July, 19388. (5) One shoot, with roots, furnished by Dr. P. W. Zimmerman; collected in the arboretum of the Boyce Thompson Institute for Plant Research, New York, summer, 1939. 1941] COPELAND: MONOTROPOIDEAE 105 Placentals Ge WN Pirate 9. Hyporrrys Monotropa. Fic. 9. Longitudinal section of nearly ripe seed X 400. The endosperm, at a certain early stage, is four-celled; the Roman numerals I-IV indicate the derivatives of these cells. Fic. 10. Model of the vascular system in the receptacle X 12.5: Ca, sepal bundles; Cod, petal dorsals; Col, petal laterals; St, stamen bundles; Cd, carpel dorsals. Fes. 11, 12. Dehisced anthers X10. Monorropa unirtora. Fic. 13. Flower X 2.5. Fic. 14. Flower with perianth removed <5. Fic. 15. Pistil x5. Fic. 16. Stigma X5. Figs. 17-20. Anthers X 10. 106 MADRONO [ Vol. 6 (6) Roots, shoots, flowers and fruits furnished by Dr. W. H. Camp; collected in central New York State at various times. As is well known, the erect shoots are in life white, like paraf- fin (the austral races mentioned above are distinguished by red color) ; dried, or placed in most preservatives, they turn black. The stem is clad with spiral scales; study of a single example showed the spiral to be orthodox. The solitary flower (pl. 9, fig. 13) is terminal on the recurved summit of the stem. Domin has quoted many conflicting authorities as to whether or not sepals are present. I find that a varying number of scales may project past the base of the flower. Of these, sometimes none and some- times one is inserted so immediately below the petals that it can be regarded as a sepal. It is not particularly different from the leaves, and I have not found more than one. The five separate petals overlap at the margins so as to form a campanulate corolla about fifteen millimeters long. Each petal is saccate at the base and truncate at the apex, sometimes with an apiculation. There are ten stamens, manifestly in two whorls, the lower opposite the petals. The densely pubescent filaments are curved inward and bear the subglobular anthers pressed against the lower side of the stigma (pl. 9, fig. 14). The ovary is belted at the base by a ten-lobed nectary; the lobes are cylindrical; they are obscurely but perceptibly paired, clasping the bases of the petalad stamens. The ovary is ovoid, marked by five deep longitudinal grooves op- posite the petals, that is, in the median planes of the carpels; and by five shallow grooves between them where the carpels meet (pl. 9, fig. 15). A shallow circular depression at the summit of the ovary is filled by the base of the short, stout, obconical style. The stigma is five-lobed, the lobes opposite the petals and sur- rounding an unusually large crater-like depression which leads into the style passage. The depression is lined by five masses of tissue which stand above the grooves, not the ridges, in the style passage. The surfaces of the five masses of tissue are more or less wrinkled, and the grooves in the style passage are obscurely continued upward upon them to some distance (pl. 9, fig. 16). The stem resembles in its anatomy that of Pterospora rather than those of Monotropsis and Hypopitys. It shows in cross sec- tion a ring of separate strands of xylem and phloem (in most specimens seriously shattered by shrinkage, apparently during fixation, a difficulty commonly encountered also in Sarcodes and Pterospora). Around the cylinder of bundles there is a continu- ous sheath of pericyclic cells with thinly lignified walls, being imperfectly developed fibers. At the summit of the stem the sheath disappears. Exactly ten bundles enter the receptacle (pl. 10, fig. 25). Each of the five lying in the planes of the petals breaks up, typically, into five. Of these, the middle one is the petal bundle; it forks further into three, a small petal dorsal which follows the contour of the sac, and two larger petal laterals which ascend past it. The behavior 1941] COPELAND: MONOTROPOIDEAE 107 ( St “\\ A pr Pa Pirate 10. Mownorropa uniIFtoraA. Fic. 21. Longitudinal section of anther X25. Fic. 22. Cross section of anther at the plane indicated by w in fig. 21 X25, Fic. 23. Area marked X in fig. 22 x 400. Fic. 24. Pollen grain Xx. 900. Fic. 25. Model of half of the vascular system in the receptacle X 12.5: Cod, petal dorsals; Col, petal laterals; St, stamen bundles; Cd, carpel dorsals; Pl, placental bundles. Hemiromes concrstum. Fic. 26. Cross section of primary flower with three secondary flowers in the axils of the sepals x 10. 108 | MADRONO [Vol. 6 of these petal bundles is quite the same as in Monotropsis, Hypo- pitys, and Pityopus. The two bundles adjacent to the petal bundle unite above it to form a petalad stamen bundle, and the two marginal branches unite to form a carpel dorsal which is traced with difficulty here at its origin. As to the five bundles which enter the receptacle between the planes of the petals, each of these forks into three: the middle one supplies a stamen of the upper whorl; the lateral ones enter the placentae, being ventral bundles of adjacent carpels. The figure shows minor deviations from the assumed typical structure as just described: one of the petal lateral bundles does not originate from a proper petal bundle; one of the placental bundles is suppressed. The carpel dorsals enter the style and ascend it for some distance, but they are there poorly developed. If the anther (pl. 9, figs. 17-20; pl. 10, figs. 21-23) is correctly understood, the outer end is proximal, the inner distal, the upper side dorsal and the lower ventral. At anthesis it includes a single chamber. Apparently (these points were regrettably not estab- lished) there are in the juvenile anther four horizontal pollen sacs, of which the ventral pair are much the larger. Dehiscence is through two curving slits at the proximal end; in dried material, and doubtless in life, these slits flare open as gaping pores. Along the margins of the slits the anther walls, elsewhere of one layer of collapsed cells, are of two layers, the walls of the inner layer bearing reticulate lignified thickenings as in a normal endo- thecium (pl. 10, fig. 23). Jepson states that the slits eventually meet, so as to convert the anther wall into two valves. This is not true in such fruiting material as I have seen; on the contrary, the pores retain their individuality until the plant dies and decays. The pollen grains (pl. 10, fig. 24) are three-grooved. The tube nucleus stains poorly and becomes distorted; the generative nucleus remains globular and deeply staining, and is surrounded by a clear space (not shown in the figure), the generative cell. The inner surface of the ovary wall is covered by a single layer of somewhat elongate cells not distinguished by staining reactions. Of the stages in the development of seed, we know only the structure of the mature ovule. It was described and figured long since by Campbell (2) quite as in Hypopitys.? 2 Since the above was written Dr. Zimmerman has had the kindness to send a beautiful collection of fruiting material made in the Arboretum of the Boyce Thompson Institute in late summer of 1940. The seeds are much as in Allo- tropa; they are elongate, having a tail at each end; the: embryo, borne on a suspensor, is usually of two cells; the endosperm has a haustorium at each end. It is now known that haustoria are produced on the endosperm in Sarcodes, Allotropa, Monotropa, and probably (to reinterpret a former observation) in Monotropsis; and that they are not produced in Hypopitys and Pleuricospora. I suppose that the absence of haustoria in these two genera is a result of par- allel evolution and that the classification given at the end of this paper, in which Hypopitys falls near Monotropa and Monotropsis and far from Pleuriscospora, may stand. 1941] COPELAND: MONOTROPOIDEAE 109 HEMITOMES CONGESTUM Gray Under the name of Newberrya, I have given a partial descrip- tion of this rather uncommon plant of the Pacific Coast of North America, and have quoted from Jepson; but Jepson’s description and mine require extension and amendment. The original generic name Hemitomes was rejected by Torrey (29) as inappropriate; and the rejection was maintained by Small (28) on account of the priority of Hemitomus L’Her. Torrey’s ob- jection is of no standing in modern nomenclature; and Small’s is disposed of by the rule (26) ““When the difference between two generic names lies in the termination, these names must be re- garded as distinct, even though differing by one letter only.” Small recognized five species. Jepson has reduced three of these, whose type localities are in California, to synonymy with the type species. With this I fully agree, and I add the one which Jepson omitted as outside his area. JI am glad to remark that Professor John Davidson has tended to support this action, in a paper read at Seattle in June, 1986; and that Dr. W. H. Camp has done so in private correspondence. The synonymy, then, is as follows: Hemitomes congestum A. Gray, Pacif. Rail. Rep. 6: 80. 1857. Newberrya congesta Torr. in Gray, Bot. Calif. 1: 464. 1876. WN. spicata A. Gray, Proc. Am. Acad. 15: 44. 1879. Hemitomes pumilum Greene, Erythea 2: 121. 1894. Newberrya subterranea Eastw., Proc. Calif. Acad. Sci., ser. 3, 1: 80. 1897. Hemitomes spicatum Heller, Cat. No. Amer. Pl. 5. 1898. H. subterraneum Heller, op. cit. Newberrya longiloba Small, No. Amer. Fl. 29: 18. 1914. N. pumila Small, op. cit. The Herbarium of the University of California has a photo- graph of the type of Newberrya spicata, and a photograph and a duplicate of Suksdorf 2168, the type of N. longiloba. I have been particularly helped by Dr. Jepson, who loaned a specimen col- lected by W. G. Wright, the type of H. pumilum. Two collections preserved in liquid have been available: (1) One shoot without roots, furnished by Dr. L. R. Abrams, who collected it in the California State Redwood Park, Santa Cruz County, June 14, 1934. (2) Several shoots and roots collected by Dr. W. H. Camp at Sol Due Hot Springs, in the Olympic Peninsula, Washington, August 5, 1982. Shoots, arising endogenously from roots, vary in height, the flowers being borne approximately at ground level; they expand in ascending, when well developed exceeding two centimeters in diameter at the base of the inflorescence. The inflorescence is essentially a bracted spike, often so brief and compact as to be accounted a head; in Abrams’ specimen the phyllotaxy, both of the leaves below the inflorescence and of the flowers, is orthodox, the apparent divergence being 3/8 or 5/18. Depauperate shoots may bear a single flower; on the other hand, vigorous shoots may bear axillary branches with one or more flowers, and may develop 110 MADRONO [Vol.6 secondary flowers in the axils of the sepals of the primary flowers. This behavior was noted on dissection of the type of H. pumilum; of Abrams’ specimen; and of a specimen by C. A. Reed in the Herbarium of the California Academy of Sciences. Jepson’s words, “Inflorescence . . . composed of short 2- to 5-flowered spikelets” imply that it is normal; but I find it only in a minority of the specimens; I do not find it in Camp’s preserved material. The flowers (pl. 11, fig. 27) are practically always tetramerous; the four sepals are oriented as in Hypopitys and Pleuricospora, dor- sally, ventrally, and laterally, the dorsal and ventral sepals being not infrequently suppressed. The sympetalous corolla is vari- able in length, from less than one centimeter to nearly two centi- meters long. The four lobes alternate with the sepals, and are separate to a depth of more or less than one-third the total length of the corolla. The dried corolla is very fragile, and it is hard to be certain as to how deep the sinuses extend. Newberrya longi- loba was distinguished by particularly deep sinuses, but I do not find these in our duplicate of the type. Indentation of the corolla lobes is variable; the retuse apex mentioned in my former account was merely the character of an individual. Stamens are normally eight, anthers dehiscing by two lengthwise slits on the outside; lobes of the nectary moderately prominent, evenly spaced; car- pels eight, alternating with the perianth parts and stamens, so that the parietal placentae, and the lobes of the stigma, standing above them, are opposite the stamens and perianth parts. Secondary flowers are usually in the axils of the lateral sepals, but may appear also in the axils of dorsal and ventral sepals (pl. 10, fig. 26). In the cluster figured, there is a second pair of scales, like sepals, above the lateral secondary flowers, as if the scales subtending these were bractlets: Hemitomes would have afforded Domin even better evidence than did Hypopitys, that the outer floral envelope is no true calyx. But what is the scale in whose axil is the dorsal secondary flower? And what is the scale on the ventral side of the primary flower, if not a sepal? I re- gard the scales subtending the lateral secondary flowers as sepals; and the scales above these flowers as a secondary pair of sepals, anomalously developed in connection with the anomaly of buds in the axils of the primary pair. The secondary flowers ordinarily have parts in smaller numbers than the primary ones, as two sepals, seven or six stamens, seven, six, or five carpels. They are later in development than the primary flowers. The vascular system in the stem, as reconstructed from sec- tions of Abrams’ specimen (pl. 11, fig. 80), is a cylinder inter- rupted by very large leaf gaps; as there are no other gaps than these, and as the vascular tissue runs in broad oblique bands rather than in slender vertical bundles, it may be regarded as a siphonostele. The leaf traces, emerging at the bases of the gaps, fork into three as they enter the scales. Flower traces are not at all united with the traces to the subtending scales; the supply to 1941] COPELAND: MONOTROPOIDEAE 111 Puate 11. Hemiromes concestum. Fic. 27. Flower x5. Fic. 28. Cross section of juvenile stamen X50. Fic. 29. Pollen grain x 900. Fic. 30. Model of the vascular system in the stem x5. Fic. 31. Model of the vascular system of the cluster of flowers shown in fig. 26 X 15: Br, bundles to the bract; Ca, sepal bundles; Fl:, supplies to secondary flowers; Caz, secondary sepal bundles; Co, petal bundles; St, stamen bundles; Cd, carpel dorsals; Pl, placental bundles. 112 MADRONO [Vol. 6 each flower consists of two bundles springing from the sides of the gap above one of the upper leaves. The figure of the vascular system of a primary flower with its attached secondary flowers (pl. 11, fig. 81) was constructed from a series of sections of Abrams’ specimen, one of which yielded the diagram (pl. 10, fig. 26). The two bundles of the flower trace approach one another, become flattened, and unite as a siphono- stele. Before this union is complete, four sepal traces emerge as single bundles above which there are large gaps. It seems that a secondary flower is typically supplied, like a primary flower, by two bundles springing from the sides of the gap left by the supply to the subtending leaf, which is in this case a sepal. Actually, we find that the secondary flower placed on the left in the figures is supplied as just described; the one on the right is partially so supplied, and partially from the sepal trace; the third, the dorsal secondary flower, is supplied entirely from the trace to the sub- tending sepal. The single bundles to the secondary sepals of the primary flower arise next to two of the secondary flower bundles. Each petal is supplied by two bundles which leave a single gap in the stele. Beyond these the stele emits three whorls each of eight bundles: stamen bundles, one opposite each perianth seg- ment, all apparently in one whorl, not at all. associated with the petal bundles; carpel dorsals, alternating with the stamen bundles, very poorly developed at the base, so that formerly I failed to discover their origin; and placental bundles, opposite the stamen bundles. Juvenile stamens were found in the secondary flowers. The anther is borne vertically on the summit of the filament and in- cludes four parallel vertical pollen sacs, of which the two on the dorsal side are much the larger. (I was formerly mistaken in making the ventral sacs the larger.) Dehiscence is through ver- tical slits on the dorsal sides of the dorsal sacs; the wall between the two pollen sacs of each lobe breaks down, and the whole outer wall swings around to the ventral side of the anther as a broad valve. The cells of the inner surface of the ovary wall are not dif- ferentiated except perhaps by smaller size. No new observations have been made upon the development of seeds; all that is known is that the ovules are essentially like those of Hypopitys and other members of the group. Discussion In concluding the previous paper of this series, I put forward a tentative scheme of classification of the plants construed as con- stituting the subfamily Monotropoideae of family Ericaceae: Tribe Pterosporeae: Pterospora, Sarcodes, Allotropa. Tribe Monotropeae: Monotropsis, Hypopitys, Pityopus, Mono- tropa, Monotropastrum, Wirtgenia? Tribe Pleuricosporeae: Pleuricospora, Newberrya, Cheilotheca? 1941] COPELAND: MONOTROPOIDEAE 113 Observations have now been extended to all of the above- named genera which are native in North America. Not a few of my previous statements have been found to require correction. A summary of the observations constitutes a formidably exten- sive table (Table 1). The data there assembled enable one to construct a tentative phylogenetic tree (text fig. 1) and to recon- sider the proposed scheme of classification. Hemitomes ~ * we Pityopus SS Monotropa Pleuricospora P / Monotropsis | / Allotropa Sarcodes 7 Peerospora rs 7 a ~—S: ue / — Bp eS ie ! Arbutoideae Fic. 1. Apparent phylogeny of the Monotropoideae. Pterospora, Sarcodes, and Allotropa continue to appear more primitive than the other genera. The primitive features are tall and erect habit; bright coloration; axile placentation in the lower part of the ovary; evenly spaced lobes of the nectary; corolla sympetalous, urceolate (in Pterospora and Sarcodes) ; bractlets present (in some specimens of Allotropa only); anthers horned (in Pterospora only) ; anthers not permanently erect. The only single characters which separate these genera as a group from the others are the habit and the coloration; but the com- bination of characters, lobes of the nectary not paired and anthers not erect, is also distinctive. In all this, there is nothing to raise doubt as to the status of the tribe Pterosporeae as a natural group; often a primitive group can be distinguished from several derived groups taken together only by a combination of negative charac- ters. Doubt appears when, on the other hand, we compare these genera with the supposed ancestral group Arbutoideae. The primitive characters of the Pterosporeae are in large part merely the characters of Arbutoideae; only saprophytism and the habit ~ MADRONO 114 SUITES PUB JPA2T PuNnois yyeoys yno ‘sjeqod ‘sjedas snoje}odurAs quid ye ZSullamoy -YYIM Jopul[éAd | pozemusloyipun éA110q a}ytsoddo [eqorsed Ssno1oUl-P -MOT[PA afed | ‘yoo190 ‘[[eurs sauojiwa Fy [P42] punois yyeoys yno syedas sno[e}edt10yo ye SULIOMOY -Y}YIM JopurpAd | poyerusleyipun AL19q opytsoddo [eyorsed Sno.JoUl-p PIM ‘yooi19 ‘Tews | v1odsoai4una]d yyeoys ST[90 07e3u0TO sunod uoyM poylusty yy poylusi[-uou o[nsdvo sjeyod dA0qe [eyoI1ed | snoyezedt.10yo poulpop ‘10ys so[punq jo sult jo 1aAv] GUO [e@PlT[No0] o}1soddo ‘MOTOq JIIXe Sno.19UI-G OPIUM Ioyyel S1opus[s}|'* ndowjouopy yyeoys no sno[e}edr10yo pode ‘.10Ys -Y}IM Jopulpéo | poyerpuotayipun é AL19q dIndsqo [eqorszed SnNO1OUI-f OPIUM Joyyel Stapusys|* sndohzid ST[90 93e.3uU0 TO sunod usyM yyeoys ou poylust[-uou ainsdeo syTeyod dA0Q” [eJoI1ed | snoTeyodi40yo MOTO pourpep ‘104s ‘sgjpunq jo sult) jo 1aAe[ ou0 auouiere| a}1soddo ‘MO]Oq I[IXe Sno1oUl-f 0} OPIUM Loyyed Slapusy[s shjdodiA S]T[90 97e8uU0Ta sunod uoyM Yyyeoys ou poylust[-uou sno[eyoduds poulpop ‘y210ys ‘satpunq jo dura; jo 1aAe[ ou0 A113q d1ndsqo [eqorred Ssno.ouwl-G é oyyM Jaye ‘tapus[s|* sisdo1zouo sno[edase yyeoys S][99 OAT]-19qy glnsdeo syTeyod dA0qge [eyorted | snoreyedts0yod | sodi4js pot YY Jepurpséo jo tov] VUO avoumere| d}1soddo ‘MOTO I[IXe SNOAIUI-G YM oy pods ‘Te 9401401] P doUsoSIyop jo oul, 1eou a[Ays Jo aseq yyeoys Jno UOI}eIPUIAIIp |ynoqe surusdo steyod sA0qe [eyorsed | snoreyoduiAs [[2} A[oze19pow -YJIM Jopurpso yuRos a[usdeo ayisoddo ‘MO[Aq J[IXB SNOJ9UI-G pot *yOa.19 *yn04s SAPOILDY yyeoys syeyod POYluUsly] YIM S][99 BYI[-1Iqy. o[usdeo oyisoddo = jaaoqve [ejor11ed | snoreyodudAs pou sa[punq jo sul. JO S1dAVT OM} [BplorpnooT, ‘dAndsqo ‘MOTOq JIXe SNO19UI-G ystdand aed pata Tle} pLodso1ajd ALBAO JO BULSI}S W19}8 UI [91S aoe fins 1guUy | yA | jo Surqor] UOI}e}UVNe[ IIMO[ | B Ke) (0) | yqeH | snuer) AVAGIOdOULONO|, NVOIWAWY HLUON dO SUMLOVUVHD ‘[ ATAVY, 115 MONOTROPOIDEAE COPELAND 1941] v éSIQOT Jo spuo [eurxoid SaqoT Jo spuo [eurxoid Saqol jo spuo [eurxoid Saqo] Jo spuo [eurxoid Soqol JO Sopis Jo as{MyysueT (Z) ‘SIqO] JO spua [eurxoid (T) é LIS UOT SoeS STVIJUIA ‘[eyUoZzI.LoYy [enboqns sovs [eizUuIA pue [esiop * po}19AUl oinjeu ‘sunoA usy MA [B}UOZILOY jenboqns Soes [B1JUIA puv [esiop ‘91N}CUL Udt{ pe}oaut ‘Sunod Udy M JooI19 IISUO] I1IV sovs [es..0op ‘ pd} AIAUI IISUO| 918 soes [es1op : [eJUOZL.LOY shijzdodhk sisdo1qouo pdo1407)F sapoowwy DLOdsoLayd oo sulei3 usjod UO S9AOOIS jo toquinN S1oyzue jo soUddSTIYIq soevs ualjod pure sisqquy sjedaeo qyusoelpe JO STe1qUuoA pesnjun 10 sa[punq 10430 s[es.op posnj ¢ spedas sTesiop jo 9o1J JaputpAo Jedaeo jo souvyd utr [edaeo ‘s[LOyM OMY poue}}ey sjedaivo yusoe pe Jo sTe1jUsA posn} so[punq 10y}0 | Sajpunq posnj s[es.op SI yore :stedas s[es.iop JO oJ Ayjetqazed pedaeo jo soue|d ur Jedaeo ‘s[LOUM OM} p ynoqe sjedieo yusoe (pe jo do1J s[ei}ueA pasn) IJUUT ‘soypuNnq s[es.op SI Yove S$ syedas s[esiop [ejod 0} posny Jedaeo jo soue|d ut Jedaeo [4oyA 19}n0 JopurpAéo sjedieo sor_punq yusoelpe Jo s[es1op [Tem ALVAO OF sTeijusA posnj Jedaeo I9UUT ‘saTpuNng ST yore ‘stedas d}yUyep ou [eyed 03 posnj so[punq g sTeyusoe|d jo soured ur ‘sno.oumMu [4oyM Ia}no | ynoQge jo suls S[V19}2[ Jedaieo sures Jo Jedaed 0} STe1jUIA posnj Iauul ‘soTpunqg SI yore ‘steyod S[e194e| [eyed 0} pasng ouou jo soueyd utr Jedaeo [LOW 193n0 Japutpso dpoeydooo1 so[pun so[pun so[pun Sse SANS (a wea ciane ade cae eae GHANILNOD ‘[ FIAVY, snusxy [Vol. 6 MADRONO Igpeoiq SUITIL}S YIM | So_puNq 19430 soes [BS1op SUI}eULI][e JO vo1J Soes [eS.1op Jo ‘[edT}AIA s[es.op SUITILYS JO syTes.op ‘ [LOYM 9UO JapurpAo 6 SopIs [esiop ATvUo1edde [edieo souv|d ut ‘g spedivo g Ajyuoredde UadxO1g sauLojiua FY sjedievo qusoe (pe jo do1J Saqol [enboqns STe1jU9A posny IOUUT ‘soTpuNnqg JO SapIs Jo soes [v1]UIA pue s[Tes.op SI ore ‘stedas syTes.op [eyod 03 posnj v ISIMY} SUI [esiop ‘[eoIz19A Jedavo jo sour|d ul Jedaveo [404M 19ynNo JapurtpAo p1odsorrina) d sjedaeo yusoe (pe Bm ero (0) | JO ST@1JUDA solpunq 19430 éSIQOT JO SoVs [V1JUDA s[Tes.op posnjun ‘stedas s[es.op JO 901} so[pung g spuo [euixoid ‘[e}UOZI.LOY Jedavo jo soueyd ul [edaeo ‘S[LOUM OMY ud} Jo dura | ° vdowouopy suoule}s YIM | sotpunq [Iystd é19du0| SUI}PeUII}e 0} popou éS9qQoT Jo soes [P1qUdA s[Tes.iop suowey}s JO s[Tes.op -u0d IOUUI be spuo [eurxoid ‘[eqUOZI.10Y Jedavo sour|d ut ‘g jedivo g ‘s[1oyM OMY JapurpAéo sndofijid suleis usljod gporydoavo1 uo Bie Reeuyae Jo Semone LS ae sTeyusoR[ g Sib as tBont as ee snuexy yo raquinn doUsDSTYIq pue sisyqquy s[A1S | [Tes A1eAQ UIdUI2}S ur 312 S 116 GdIGNTIONOD ‘T ATAV, 1941] COPELAND: MONOTROPOIDEAE 117 which is presumably a result of it are distinctive. Furthermore, the Pterosporeae are notably heterogeneous in anatomy of stem and receptacle, and in corolla, anther, fruit, and seed. I would allow the tribe to stand as a tentative group, and would expect studies of the Arbutoideae to afford evidence as to whether or not it is a natural group, having one origin among Arbutoideae. The Monotropeae, though differing among themselves in vari- ous features, yet exhibit marks of unity; the saccate bases of the petals and the paired lobes of the nectary are associated with a peculiar and definite structure of the vascular supply to the petal. This is safely a natural group. In the corolla, Monotropsis is the most primitive of the genera; but in the ovary, Hypopitys and Monotropa are more primitive, and the feebly developed sheath of fibers is a primitive character in the stem of Monotropa. We may take it that in most respects Monotropsis represents a direct ances- tor of Hypopitys, and Pityopus an only slightly modified descen- dant; Monotropa stands apart from all three, differing not only in the well known solitary flower and in the anatomy of the stem, but also in the number of grooves on the pollen grain. We may safely maintain Hypopitys and Monotropa as separate genera. The tribe Pleuricosporeae has ceased to be tenable as consti- tuted. The common characters of Pleuricospora and Hemitomes, in habit, floral diagram, placentation, and ovules, are features all of which are apparently readily reached by parallel change and some of which are widely distributed. They are outweighed by the differences in pubescence, vascular anatomy of the receptacle, and structure of the anther. Pleuricospora is strictly glabrous. The petal bundles, not forking into two large bundles and one small one, are those of Pterosporeae rather than of Monotropeae. The anther is unique; the only thing elsewhere in the group that suggests it is that of Pterospora. The four-grooved pollen grains again suggest Ptero- sporeae rather than Monotropeae. But we cannot connect this genus with any one genus of Pterosporeae. Hemitomes has the floral diagram of Hypopitys, and, in more extreme form, the irregularity of inflorescence observed in that genus. We may account for the unique petal trace of two bundles by supposing the small middle bundle of the petal traces of the Monotropeae to have been suppressed, this suppression being associated with loss of saccate bases to the petals and pairing of the lobes of the nectary. The slits along the dorsal sides of the dorsal sacs of the anthers seem to amount to a modification of the pores of normal Ericales, which are still present in Monotropsis and Monotropa. The two-grooved pollen grains are those of most Monotropeae. It would be possible to dispose of these genera by assigning one to Pterosporeae, the other to Monotropeae, distinguishing these tribes by glabrous stamens and pistils in the former, pubes- cent stamens and pistils in most of the latter. Such a scheme, 118 MADRONO [Vol. 6 however, would increase the heterogeneity of the Pterosporeae and break the unity of the Monotropeae. I think it best to place each of these genera in a separate tribe, making altogether four tribes to be distinguished as follows (the oriental genus Cheilo- theca remains unplaced; it may perhaps constitute a fifth tribe) : aXe ee a 10. Stamens and pistils glabrous; lobes of the nectary not paired; no gaps above the petal bundles, to which the outer whorl of stamen bundles are more or less fused; grooves on the pollen grain 4 (3 in Allotropa) ; young stems not nodding. 1. Stems erect above ground; red pigment present; anther bent inward; placentation axile below: Tribe Prrro- SPOREAE: Pterospora, Sarcodes, Allotropa. 2. Inflorescence at ground level; red pigment absent; anthers erect; placentation parietal: Tribe Pxirurico- SPOREAE: Pleuricospora. . Stamens and pistil often pubescent; red pigment usually absent (present in varieties of Monotropa, a trace of it in Hemitomes) ; gaps present above the petal bundles; grooves on the pollen grain 2 (3 in Monotropa). 1. Bases of petals saccate; lobes of the nectary in pairs; petals with small dorsal bundles and large lateral bundles; young stems nodding (unknown in Pityopus) : Tribe MonotropeaE: Monotropsis, Hypopitys, Pityopus, Monotropa, Monotropastrum, Wirtgenia? 2. Base of petals not saccate; lobes of the nectary evenly distributed; dorsal bundles of the petals suppressed; inflorescence at ground level: Tribe Hemrromear: Hemi- tomes. | Sacramento Junior College, Sacramento, California, August, 1940. LiTerRATuRE CiTED Bentuam, G., and J. D. Hooxer. Genera plantarum. 3 vols. London, 1862-1883. Campsett, D. H. Monotropa uniflora as a subject for demonstrating the embryo sac. Bot. Gaz. 14: 83. 1889. CuristropH, H. Untersuchungen iiber die mykotrophen Verhialtnisse der “Ericales” und die Keimung der Pyrolaceae. Bot. Centralbl. Beih. 38°: 115-157. 1921. Corrtann, H. F. The development of seeds in certain Ericales. Amer. Jour. Bot. 20: 513-517. 1933. . The structure of the flower of Newberrya. Madrofo 2: 137-142. 1934. On the genus Pityopus. Madrofio 3: 154-168. 1935. . The reproductive structures of Pleuricospora. Madrofo 4: 1-16. 1937. The structure of Allotropa. Madrono 4: 137-153. 1938. The structure of Monotropsis and the classification of the Monotropoideae. Madrono 5: 105-119. 1939. Crantz, H. I. N. Institutiones Rei Herbariae.... 2 vols. Vienna. 1766. 1941] BUCHHOLZ: PODOCARPUS GRACILIOR 119 11. Domin, K. Vergleichende Studien iiber den Fichtenspargel. Sitzungsber. kgl. bohm. Gess. Wiss., II KI., I Stiick: 1-111. 1915. 12. Doyen, B. E., and L. M. Goss. Some details of the structure of Sarcodes. Madrono 6: 1-7. 1941. 13. Drupe, O. Pirolaceae. in A. Engler and K. Prantl. Die natiirlichen Pflanzenfamilien 4°: 3-11. 1889. 14. Eastwoop, Auice. Some new species of Californian plants. Bull. Torrey Bot. Club 29: 75-82. 1902. 15. Fostrr, A. S. Problems of structure, growth and evolution in the shoot apex of seed plants. Bot. Rev. 5: 454-470. 1939. 16. Hitt, Joun. The British herbal. ... London. 1756. 17. Irmiscu, T. Einige Bemerkungen iiber die einheimischen Pyrola-Arten. Bot. Zeit. 14: 585-591, 601-602. 1856. 18. Jepson, W. L. Manual of the flowering plants of California. Berkeley. 1925. 19. ———————.. A flora of California, vol. 3, part 1. Berkeley. 1939. 20. Kamiensk1, F. Die Vegetationsorgane der Monotropa hypopitys L. Bot. Zeit. 39: 457-461. 1881. 21. —_—_————_. Les organes végétatifs du Monotopa Hypopitys L. Mém. Soc. Nat. Sci. Nat. Cherbourg 24: 5-40. 1882. 22. Kocu, L. Das Entwicklung des Samens bei Monotropa Hypopitys L. Jahrb. wiss. Bot. 13: 202-252. 1882. 23. Linnaeus, C. Species Plantarum. 1753. 24. Mac Doveat, D. T. Symbiotic saprophytism. Annals Bot. 13: 1-47. 1899. 25. Nurratt, T. The genera of North American plants. ... Philadelphia. 1818. 26. Renpite, A. B., and others, editors. International rules of botanical nomenclature ... dritte Ausgabe. Jena. 1935. 27. Sma, J. K., and A. M. Var. Report of the botanical exploration of south- western Virginia during the season of 1892. Mem. Torrey Bot. Club 4: 93-201. 1893-1894. 28. Smatt, J. K. [Monotropaceae] in North American Flora 29': 11-18. 1914. 29. Torrey, J. On Ammobroma, a new genus of plants. ... Ann. Lyc. Nat. Hist. New York 8: 51-56. 1864. 30. Wypier, H. Kleinere Beitrage zur Kenntnis einheimischer Gewachse. Flora N. R. 18: 613-617. 1860. PODOCARPUS GRACILIOR IN CULTIVATION Joun T. BucHyHouz > The “African fern-pine,” in recent years popular as a decora- tive tub plant and also planted in the open as an ornamental coni- fer, may become a tree of considerable size. Since mature speci- mens growing in California have produced pollen cones during the past winter, it is now possible to identify the species as Podocarpus gracilior Pilger. The plant has been regarded as a conifer of South African origin and has usually passed under the nursery trade name of “Podocarpus elongata,” which is unquestionably an error. The latter is the legitimate name of a plant of South Africa where there are two narrow leaved species that have been confused and have at one time or another passed under this botanical name. The California exotic has narrow leaves that are somewhat similar to those of Podocarpus elongatus L’Herit. (the earliest described 120 MADRONO [Vol. 6 species in this genus), but the pollen cones and seeds differ con- siderably. It resembles P. falcatus (Thunb.) R. Br. (P. elongata Carr.) more closely than P. elongatus L’Herit., but has green in- stead of brown twigs, leaves that become longer and wider, and pollen cones that are more than twice as long; also the tips of individual microsporophylls (connectives) of the pollen cones are more pointed. In both P. gracilior and P. falcatus the pollen cones are axillary, borne singly or in fascicles of two or three. The seeds of Podocarpus gracilior were brought over from East Africa in 1911 by Mrs. Stewart Edward White. Franceschi, of Santa Barbara, germinated some, if not all, of these. The state- ment is usually current that these seeds were introduced from South Africa (an error which has appeared in print), but the material itself does not agree with the descriptions of either of the narrow-leaved South African podocarps. It fits very closely the description of Podocarpus gracilior. This entire question has been clarified recently by a letter received from Colonel Stewart Edward White in which he states that the tree from which these seeds were collected was located in Kenya, British East Africa. This region of Africa is included in the range given for Podocarpus gracilior Pilger. Specimens of this species at the United States National Herbarium were collected in Kenya by Edgar A. Mearns of the Smithsonian African Expedition which was led by Theo- dore Roosevelt in 1909-1910. The male flowering material col- lected in California in January and February, 1941, in three © widely separated localities agrees in all essential details with the Mearns specimens. Podocarpus gracilior is dioecious, as are nearly all species of Podocarpus. Except for the limited number of original seedlings, the trees in California have been propagated from cuttings. The stock tree used for propagation since about 1922 at the Coolidge Rare Plant Garden Nursery in Pasadena is a male tree. During the past winter Mr. J. J. Mulvihill has kindly sent me a number of reproductive twigs. Thousands of plants have been grown as cuttings from this tree over a period of years and furnished to the nursery trade under the names “African Fern-Pine” and ‘Podo- carpus elongata.” They do very well as tub plants and when these long-suffering decoratives become too old they may be trans- planted to parks and gardens. The writer has seen this conifer used as a street tree in Los Angeles and Pasadena. Apparently they do not become reproductive until they are mature specimens of large size. The largest known specimen, about fifty feet high, is growing at Alameda Plaza in Santa Barbara where two other mature specimens may be seen, none of which had been observed in reproductive condition. However, during July, 1941, Mr. Van Rensselaer of Santa Barbara Botanic Garden found one of these (located in the east section of Alameda Plaza) bearing seeds. Many of the seeds were abortive when less than half grown; some 1941] BUCHHOLZ: PODOCARPUS GRACILIOR 121 Fic. 1. Twigs of Podocarpus gracilior Pilger bearing nearly full grown pollen cones, January, 1941. had enlarged to full size, but all of those which were examined by the writer, were without embryos, with the endosperms shriveled in the manner usual for unpollinated parthenocarpic seeds. With this one exception, all reproductive specimens of P. gracilior thus far observed have proved to be male, although pre- sumably the seeds would produce equal numbers of male and fe- male seedlings. The location of many of the original seedlings is not known. Two specimens growing on the estate of Colonel Stewart Edward White near Burlingame have not been observed in reproductive condition at any time. The three large trees at Alameda Plaza are probably seedlings, and likewise any other old specimens growing in Santa Barbara. It is likely that the tree belonging to Mrs. E. N. Hazard, mentioned by Van Rensselaer (Trees of Santa Barbara 1940, p. 84) is also one of the original seedlings. In 1936 the writer found five or six large specimens on the Dwight Murphy estate at Montecito, Santa Barbara County and these may be seedlings. They have not been observed in reproductive condition. The late Miss Kate Sessions of San Diego informed the writer in 1936 that she had obtained three of the original seedlings from Franceschi. The location of two of them was given. One is growing in the gardens of Julius Wangenhein, 148 West Juniper Street, San Diego. Another is the large tree at the Rosecroft Gardens in San Diego. The latter plant was not reproductive in 122 MADRONO [Vol. 6 1936, but was found with pollen cones during the past winter. Mr. A. D. Robertson furnished the writer with male reproductive specimens from this plant in January and February, 1941. A male tree growing in the Botanical Garden of the University of California, Berkeley, was observed to be in reproductive condi- tion in January, 1941, by Mr. Donald G. Nelson of that institution. The origin of this plant is not known to the writer. Aside from the dozen plants enumerated here, there are probably a number. of other specimens in cultivation on private estates that represent original seedlings, which were distributed in the days before it was discovered that these plants are easily propagated. University of Illinois, Urbana, July 21, 1941. THE TAXONOMIC STATUS OF MICROSTERIS GREENE Hersert L. Mason Perhaps no member of the Polemoniaceae has been so greatly misunderstood as the very polymorphic aggregate species, Phlox gracilis (Dougl.) Greene. It has been variously treated as a mem- ber of the following genera: Gilia, Collomia, Phlox, Navarretia, Polemonium, and is the type species of the genus, Muicrosteris Greene; it has been divided and subdivided into species, sub- species, varieties, subvarieties and forms within these genera ac- cording to the particular whim of the author treating it. The plant ranges from the Pacific Coast to the Rocky Mountains and from temperate Alaska south to Mexico, and recurs in the South- ern Hemisphere in Bolivia, Chile, and Argentina. Essentially an early spring annual, it occurs from the coastal bluffs to timberline. The intent of the present paper is to deal only with the generic position of the aggregate species and not to be concerned with the status and disposition of the smaller taxonomic units. Therefore, the entire group of variants will be treated, for the present at least, as one large, polymorphic species. The species was first collected by Douglas on the banks of the Spokane River [Washington] and given the manuscript name, Collomia gracilis; it was first described by Hooker (6) in 1829 under the name Gilia gracilis with Collomia gracilis Douglas cited as a synonym. In 1887 Greene (4) referred the species to the genus Phlox with the statement: “This interesting plant came to the knowledge of botanists some years in advance of Phlox Drum- mondii Hook. and its allies. It was at first a thing of dubious aspect, not at home either in Gilia or Collomia. But since the dis- covery of the Texan group of annual species of Phlox with peculiar habit, it must have been the mere force of custom which has kept men from seeing that it is an absolutely perfect congener of Phlox Drummondi.” In 1891 (7, p. 433) O. Kuntze, recognizing the page priority of Navarretia over Gilia, made a purely nomencla- 1941] MASON: MICROSTERIS 123 torial shift in the combination Navarretia gracilis (Dougl.) Kuntze. In 1898 Greene (5) erected the genus Microsteris recognizing as species seven segregates of Phlox gracilis. In so doing Greene stated: ““At present I am disposed to adopt it as a principle that species with mucilaginous seeds are nowhere, in this family, to be placed as congeneric with such as have seeds devoid of the gum- miferous coating. This implies the removal of my Phlox gracilis from the genus Phlox.” In his description of Microsteris he states “Calyx, corolla, stamens and capsule wholly as in Phlox.” Thus Greene’s Microsteris hangs by the single character “mucilaginous seeds.” Inthe same year O. Kuntze (8, p. 203) referred Collomia gracilis Dougl. to Polemonium by the simple statement: “P. More- nonsis OK (Collomia gracilis Dgl. non Polemonium gracile).”’ His reasons are forever hidden in parenthetical synonymy. Wecan dismiss without further comment the references to Navarretia and Polemonium. The reference by Douglas to Collomia is understand- able. It was based upon superficial resemblance; furthermore, at that time the genus Collomia had not been clearly circumscribed in the light of the family as a whole. Our problem resolves itself into determining whether Phlox gracilis shall be retained in Gilia as interpreted by Hooker, be retained in Phlox as interpreted by Greene in 1887 or be placed in Microsteris following Greene’s later interpretation. Of subsequent authors most have preferred to follow Asa Gray’s adaptation of Hooker’s treatment in a broad concept of the genus Gilia while only a few have used either Phlox or Microsteris when referring to this species. Brand (2) in his monograph of the Polemoniaceae with its highly elaborated sys- tem of “pigeon holes” chose to place Phlox gracilis in the genus Gilia, subgenus Benthamiophila, section Phlogastrum and _ pro- ceeded to divide the species into fourteen entities in various sub- specific categories. With respect to the generic position of the species I quote from Brand, “‘Species sic intermedia inter genera Phlox et Gilia, ut vix discernere possis, cui generi eam attribuas; a Collomia tamen, quacum plurimi autores junxerunt, calyce, ut cl. Greene docuit, valde diversa.” Although he cited Microsteris as a synonym it is clear from the above quotation that he did not re- gard Microsteris as offering any problem. He was concerned with differentiating Gilia from Phlox. Here again we find but a single character utilized to place the species in Gilia, namely, the fact that the seeds develop mucilage when wetted. Other characters which it possesses that align it with Phloz are treated by Brand as exceptions in Giulia. The most recent treatment that bears on this problem is that of Wherry (9) from whom we quote, “Microsteris. A few diminu- tive western annuals constitute this genus, which has been by vari- ous authors referred to Collomia, Gilia and Phlox. It shows little relationship with the first two genera, and in view of the differ- ence in seeds can scarcely be congeneric with the last, although it may well be a derivative.” Wherry, it will be seen, dismisses 124 MADRONO 7 [Vol. 6 Gilia and Collomia from consideration but parries between Phlox and Microsteris. He finally eliminates Phlox on the basis of ““COROLLA-LIMB small; seeds becoming sticky when moistened,” but he does at least suggest the responsibility of Phlox for the offspring. In an effort to validate the genus Microsteris an- other very insignificant character is added to the one previously utilized, namely the small size of the corolla limb. It is perhaps a reasonable mode of escape when a group of plants does not fit comfortably in any of the related genera to erect a genus for it. However, this procedure should not be adopted until all of the evidence is carefully weighed to determine the precise nature of the differences that seem to make it neces- sary. As pointed out above Microsteris was erected by Greene who listed for it a single character difference from Phlow. Wherry’s additional character of a small corolla limb adds scarcely anything of generic significance. The following tabular arrangement presents the facts pertaining to the development or non-development of mucilage or spiracles in the seed coats of most of the more widely accepted genera or Polemoniaceae. Bonplandia: all species develop mucilage. Cantua: a few species develop spiracles, the rest do not. Cobaea: some species produce spiracles, and other species mucilage. Gilia: very diverse, some species produce mucilage, others do not. The section Ipomopsis, recognized as a distinct genus by Wherry, is about equally divided in this respect. Hugelia: some species produce mucilage, others do not. Langloisia: all species produce mucilage. Leptodactylon: in species examined none produce mucilage. Linanthus: most species produce mucilage, some do not. Loeselia: some produce mucilage, others do not. Navarretia: some produce mucilage, others do not. Phlox: as interpreted by Greene and by Wherry, does not pro- duce mucilage, but if Microsteris is included, will be on the same basis as the other large genera. Polemonium: some species produce mucilage, others do not. The remaining few genera are each very small, and I have not as yet investigated them. But from the above data it would ap- pear that the development of mucilage by the seed coat cannot be relied upon as of primary generic significance. All we can say of Phlox is that in the majority of species the seeds are immutable when wetted. This leaves as a character for the segregation of Microsteris only the small corolla limb. The magnitude of dif- ference here, however, is no greater than the variational limits of corolla size in several other genera of Polemoniaceae, such as Col- lomia, Navarretia and Linanthus. This evidence, it seems, is just cause for denying generic status to Microsteris. When we consider the characters that serve to keep the Phlozr gracilis aggregate out of Gilia we turn from the flower and seed to 1941] MASON: MICROSTERIS 125 Puate 12. Comparison oF PHLOx GRACILIS AND PHLox DRUMMONDII VAR. TENUIS. Fic. 1. Phlox gracilis. Fic. 2. Phlox Drummondii var. tenuis. Fic. 3. Phlox gracilis, flower. Fic. 4. Phlox gracilis, opened corolla. Fic. 5. Phlox gracilis, capsule showing disarticulation of valves. Fic. 6. Collomia, capsule showing valves with margins reflexed. Fic. 7. Campanulate type of capsule found in many species of Linanthus and Gilia. 126 MADRONO [Vol. 6 other parts of the plant. Of the authors who have referred the group to Gilia we find some who regard most of the small genera (Linanthus, Hugelia, Gymnosteris, Loeselia, Collomia, Leptodactylon) as belonging to this genus; others who recognize the small genera mentioned above but who have followed precedent in the disposi- tion of Phlox gracilis. When we exclude from Gilia these small genera there still remains a polymorphic but closely related group of species. The leaves of this remaining group are normally alternate (occasionally through shortening of the internodes they may appear subopposite), and frequently pinnately toothed, lobed or dissected; the corolla lobes are normally entire; the stamens are usually, but not always, equally inserted and equal in length; the capsule valves do not disarticulate on dehiscence but remain united at the base, and although the valves may spread cam- panulately or sometimes reflex on the midvein the capsule falls as a whole (pl. 12, fig. 7); the locules of the ovary are usually more than one-seeded, but occasionally are one-seeded; the seeds are usually small and angular. Phlox gracilis does not conform with Gilia as the following summary of its characters demonstrates: the leaves are pre- dominately opposite (pl. 12, fig. 1), at least below, and are always linear, or oblong and entire; the corolla is salverform, the limb rotate, the lobes frequently emarginate (pl. 12, fig. 3); the sta- mens are unequally inserted and unequal in length (pl. 12, fig. 4); the capsule valves are rigid and disarticulate completely on dehiscence; the locules are one-seeded, the seeds large (pl. 12, fig. 5). Greene was quite correct when he said in his diagnosis of Microsteris, ““Calyx, corolla, stamens and capsule wholly as in Phlox.” And of course Wherry accepts for this group a close relationship to Phlox. It seems that the presence of such typical Phlox characters as the rigid, disarticulating capsule valves and the solitary large seeds in the locules, together with several minor characters which are usual in Phlox and occasional or abnormal in Gilia, throw the weight of the argument to Phloz, not to Gilia. Another line of evidence supporting a relationship with Phlox rather than with Gilia is found in cytological studies; the basic chromosome number in Gilia appears to be n=9 while the basic chromosome number in Phloz is n=7. In Phlow gracilis 2n=14, the count being made from root tip cells. However, in a group with such wide climatic tolerance and such great morphological diversity we may anticipate some polyploidy. Botanists familiar with the genus Phlox only in western North America may be pardoned for hesitating to place P. gracilis, a plant so different from P. Douglasii and P. adsurgens, in the same genus. It is, as Greene points out, only when we take into con- sideration the range of variation of the entire genus that we can hope for a true picture of relationship. In this case the Phlox Drummondii complex of Texas offers a key to the relationship. A collection of Phlox Drummondii var. tenuis Gray from Texas 1941] : MASON: MICROSTERIS 127 (Lindheimer 468) is an excellent example of a connecting type between Phlox gracilis and other members of the genus. A com- parison of figures 1 and 2 (pl. 12) will at once show the great similarity in aspect between the two. Figure 1 represents a plant of Phlox gracilis collected at Tuolumne Meadows, Yosemite National Park, California (Mason 4869). It was especially se- lected for this comparison but is representative of a large segment of the “Microsteris” variants. The evidence of a general similar- ity of aspect substantiated by indisputable Phlox characters up- holds Greene’s first opinion of the generic position of this group. The fact that this western group of plants is related to an eastern group by way of a southern bond is not inconsistent with the growing body of information now being accumulated relative to the history of vegetation in the southwest. Among other genera with related species showing a similar distribution pattern are Juglans, Cercis, Forestiera and Frazinus. This group of trees and shrubs are all represented in fossil floras of Middle Tertiary time and today occur in savanna like floras where Phlox gracilis is a common associate. It would seem that these relationships go back at least as far as the Miocene, if not the Oligocene, in the Sierra Madrean flora of Axelrod (1). Perhaps this region has been the center of origin and differentiation of the entire Pole- moniaceae. Certainly not all Phlox species have had their origin in Keewatin Land as postulated by Wherry (10). If this were true it would be reasonable to expect a higher development of the genus in the old world than is now evident, since migration routes through Beringia would have been available. The occurrence of Phlox in this northern region during the Pleistocene, north and west of the Keewatin center of glaciation is attested by fossil fruits reported and figured by Chaney and Mason (3 p. 17, figs. 34,36). These specimens are strikingly similar to P. sibirica L., a species occurring in the Alaska region today, and ranging west- ward into Siberia. Department of Botany, University of California, Berkeley, August 27, 1941. LIrERATURE CITED — AxeEtrop, D. A Miocene flora from the western border of the Mohave Desert. Carneg. Inst. Wash. Publ. 516. 1939. 2. Brann, A. [Polemoniaceae] in Engler, Pflanzenreich 4°°: 1-203. 1907. 3. CuHaney, R. W., and H. L. Mason. A Pleistocene Flora from Fairbanks, Alaska. Amer. Mus. Nov. no. 887: 1-17. 1936. 4. GREENE, E. L. New or noteworthy species. Pittonia 1: 139-143. 1887. 5. ————————.. Some western Polemoniaceae. Pittonia 3: 299-305. 1888. 6. Hooxer, W. T. Bot. Mag. 56: t. 2924. 1829. 7. Kuntze, O. Rev. Gen. PI. 1. 432-434. 1891. 8. ———_—_—_—_—_.. Rev. Gen. Pl. 3”: 202-203. 1898. 9. Wuerry, Epcar T. A provisional key to the Polemoniaceae. Bartonia 20: 14-17. 1940. 10. ————_—__., Geographic relations in the genus Phlox. Op. cit. 20: 11-14. 1940. 128 MADRONO [Vol. 6 THE HOLACANTHOID PLANTS OF NORTH AMERICA Cornetius H. Mutyer One of the striking botanical phenomena of the southwestern United States and adjacent Mexico is the occurrence of several genera of different families with vegetative characters so similar as to make them confusing. Perhaps the most outstanding such instance is that of the usually leafless green spiny shrubs which resemble Holacantha Emoryi A. Gray, a member of the Simarub- aceae. The similar forms are Koeberlinia spinosa Zucc. in the Koeberliniaceae, Canotia holacantha Torr. in the Celastraceae, and Thamnosma montana Torr. & Frem. in the Rutaceae. These four species are characterized by leaves reduced to scales and early caducous, and spinescent stems persistently green to carry on the photosynthetic process. The lack of leaves and similarity of the spines make these plants rather difficult to dis- tinguish without flowers or fruit. Yet, a series of vegetative characters may be recognized by which the species are readily distinguished. There are other spinescent plants which might be confused with the group here treated. The others, however, are charac- terized by shorter and more slender spines and usually more prominent leaves. For instance, Adolphia in the Rhamnaceae and Forsellesia in the Celastraceae contain species highly similar in habit to the Holacantha-like plants, but in these the spines do not exceed an average of 2.5 millimeters in thickness, and the leaves are prominent throughout the greater part of the growing season. Furthermore, Adolphia is characterized by opposite leaves and branches, while those of the plants here treated are all alternate. Two variants of Koeberlinia spinosa have been distinguished under varietal names. Both of these are characterized by very slender spines which might make them difficult to distinguish from Adolphia except for their alternate habit of branching. Leafless individuals of the leguminous shrubs Cercidium and Cassia armata might become confusing when they lack flowers and fruit. The highly artificial group comprised of the Holacantha-like plants is characterized by the alternate-branching, spinescent stems, the spines usually quite coarse, branches green and photo- synthetically functional for several years, leaves much reduced and early caducous. Johnston (Journ. Arn. Arb. 21: 356-368. 1940) has pointed out the significance of this group of unrelated plants of similar habit as indicative of a relationship between the North American deserts and those of South America where the same habit is common. He showed the habit to be much more general in the South American deserts than in North America. The highly endemic character of all our species except Koeber- linia spinosa would indicate that the plants are relicts of a time when the habit was more common in North American deserts. 1941] MULLER: HOLACANTHOID PLANTS 129 Key Basep oN VEGETATIVE CHARACTERS Leaf scars and branching alternate, leaves inconspicuous and early caducous. Plants glabrous, stem either densely glandular or with minute longitudinal lines of white waxy flakes. Stems densely glandular or warty with translucent glands; buds and spine tips tan; branches with a pair of extra- axillary buds at the bases (one on each side).......... Thamnosma Stems not glandular, with minute longitudinal lines of white waxy flakes; buds dark brown or black, spine tips brown; branches with no extra-axillary buds at the bases (though the buds along the length of the branch may descend to aapoint wear the WaASe) fis dof nar eees eo ps Rane i in aes Canotia Plants with young stems pubescent, neither glandular nor with waxy exudations. Younger stems yellow-green, minutely puberulent with short spreading hairs or these reduced to pustules; branches and spines with a pair of buds at the bases, these extra- axillary. (one on each side) ..............0.. 0c eens Koeberlinia Younger stems gray green with densely matted appressed silky hairs; branches and spines with a single axillary bud and no lateral or extra-axillary buds at the bases .. Holacantha Leaf scars and branching opposite or, if alternate, the leaves conspicuous and persistent ... (genera not treated in this article). THAMNosMA MoNTANUM Torr. & Frem. in Frem., Rep. Exped. Rocky Mts. 813. 18465. Central Arizona north to southwestern Utah, southern Ne- vada, and southern California; most common in western Arizona. A shrub usually 3 to 6 or 7 decimeters tall. (fig. 1). CanoTia HOLACANTHA Torr., U. S. Rep. Survey Railroad Miss. Pac. 4: 68. 1856. Southeastern and central Arizona to northwestern Arizona and doubtfully in the Providence Mountains of southeastern Cali- -fornia. A shrub 8 to 5 or even 6 meters tall. (fig. 2.) KoEBERLINIA SPINOSA Zucc., Abh. Akad. Muench. 1: 859. 18382. Southern Arizona and New Mexico, western Texas, Baja Cali- fornia, northern Sonora and Chihuahua, south through Coahuila and Nuevo Leon to Puebla and Oaxaca. A shrub 0.5 to 2 meters or even a small tree to 5 meters tall. (fig. 1.) KoEBERLINIA SPINOSA Var. TENUISPINA Kearney & Peebles, Journ. Wash. Acad. Sci. 29: 486. 1939. Yuma County, Arizona, and Sonora, Mexico. Differs from the species in its elongate slender spines, blue-green color, and usually longer sepals, petals, and filaments. Typical K. spinosa apparently occurs nowhere west of Tucson, Arizona. This variety is not distinguished from the species on the distribution map. A second variety (K. spinosa var. verniflora Bogusch, Torreya 31: 74. 1931) now scarcely seems worthy of distinction. Al- though this form differs strikingly from the species by its slender spines (and generally reduced size of the organs) and its early flowering (in March and April), its differences are not sufficiently profound nor constant to warrant formal recognition. Nor is [ Vol. 6 ~ MADRONO 130 WOANVLNOW VWSONWVHL X VSONIdS VINITHAGIOy @ es ILYVYM3LG VHLNVOOTOL] O \AMOW VHLNVOOTOL) xX VHLNVOOIOH VILONVS @ Ga el 1941] MULLER: HOLACANTHOID PLANTS 131 there any geographical segregation from the species. The mor- phological basis of this form has been observed by the author in various parts of western Texas and adjacent Chihuahua and Coahuila. In a letter to the author under the date October 7, 1940, Bogusch discusses the variety as follows: “ . observa- tions made in the field upon specimens of Koeberlinia spinosa Zuce. convince me that the variety verniflora described by me is prob- ably a normal reaction of the plant to wound stimulus or repre- sents a form of juvenile growth. From material seen both in the type locality of the Rio Grande Valley of Texas and in the region west of Uvalde, I have seen that new growth which follows ex- tensive injury to the plant often results in being more attenu- ated, both in branches and the spines. The peculiarities asso- ciated with the time of flowering—not necessarily a taxonomic character—may be best explained that in the Rio Grande Valley the vegetation in many respects matures earlier and comes into flower sooner than elsewhere in the state.” Horacantua Emoryr A. Gray, Mem. Am. Acad. ser. 2, 5 (PI. Nov. Thurb.): 310. t.8. 1855. Central and southwestern Arizona to southern California. A shrub 1 meter tall to a small tree reaching 3.5 meters (fig. 2). An undescribed species of Holacantha was recently discovered in northen Mexico. Holacantha Stewartii sp. nov. Frutex procumbens vel ad- scendens 1.5-3(6) dm. altus duplo latior; spinae plusminusve appresso- vel patenti-pubescentes glabratae, papillis minutis exceptis; fructus acutus margine ventrali obtuso-costatus. Low shrub, 1.5—3 dm., rarely 6 dm. tall, usually 2—4 times as broad, procumbent or somewhat ascending, soon leafless, coarsely spiny; stems and spines divaricate; spines 2.5-—6(12) cm. long, (1.5)2.5-3 mm. thick, tips subulate, brown, 3-4 mm. long, branches terete, the immature ones becoming sulcate in drying, glabrate or somewhat spreading-pubescent or appressed-seri- ceous, hairs short, bases persisting as minute papillae; buds in- conspicuous in axils of spines and sparsely scattered along their length, surrounded by small tufts of coarse appressed hairs; leaves quickly deciduous, oblong, acute at both ends, sessile, 5—8 mm. long, 2—2.5 mm. broad, red at vernation (as are the young spines for a time), densely white- or rose-hirsute, becoming green and sparsely hirsute ; flowers dioecious; staminate calyx of 6 ovate, acute, pubescent sepals about 1 mm. long; corolla of 6 fleshy, dorsally pubescent, deeply concave petals with narrow, thin mar- gins, 4 mm. long, 1.75 mm. broad (not flattened), stamens about 12, filaments 1.5—-2 mm. long, broadened basally, strongly hirsute, apex subulate, glabrous; pistillate calyx similar to the staminate; pistillate corolla not seen; staminodes similar to the functional filaments; carpels distinct, apically connivent, stigmas sessile, fused; fruit persistent 1-2 years, 6-carpellate (or carpels fewer 132 MADRONO [Vol. 6 by abortion), carpels distinct, divaricate, lenticular-ovate, acute, ventral margins obtusely ridged, superior, 8-9 mm. long, 5-6 mm. broad, 3—4 mm. thick, glabrous, red or green, surface nearly smooth, lacquered. (fig. 2.) Range: Mexico; western Coahuila and northern Zacatecas. Holacantha Stewartiti is named in honor of Mr. Robert M. Stewart of Santa Elena, Coahuila, whose superior hospitality and whose company on several side trips contributed markedly to the pleasure and success of the several weeks’ work in the vicinity. This species is the second described in this rare and striking genus, the first being Holacantha Emoryi which is confined to Ari- zona and California. From that species H. Stewarti differs in its low sprawling habit, the sparse pubescence of its stems (com- pared with the densely short-tomentose stems of H. Emoryz), the persistence of papillae-like hair-bases, and its usually markedly acute and ventrally ridged carpels. The great discrepancy in the ranges of these two endemics further attests their distinctness. Specimens examined. Coanuiua: Sierra de las Cruces, gulch in limestone hills 0.5 mile north of Santa Elena, August 13, 1940, I. M. Johnston & C. H. Muller 215 (United States National Arbo- retum, USNA, type, sheet no. 96733; Arnold Arboretum, AA) ; southeast base of Sierra de las Cruces, 3 miles northeast of San José, September 5, 1940, 7. M. Johnston & C. H. Muller 1003 (AA, USNA); north base of Sierra de las Cruces, at San Rafael, Sep- tember 8, 1940, I. M. Johnston & C. H. Muller 1084 (AA, USNA); northwest base of Sierra de las Cruces, at San Vicente, September 8, 1940, I. M. Johnston & C. H. Muller 1065 (AA, USNA); 8 to 5 miles south of Laguna de Jaco, September 10, 1940, J. M. Johnston & C. H. Muller 1104 (AA, USNA); north end of Bolson de los Lipanes between E] Almagre and Cerros de Leja, September 12, 1940,I.M. Johnston & C. H. Muller 1239 (AA, USNA). Zacatecas: banks of arroyos in foothills, Hacienda de Cedros, 1908, F. E. Lloyd 191 (United States National Herbarium ). With the exception of the type collection and Lloyd’s from Zacatecas all the plants collected or observed grew in deep, heavy silt flats, usually associated with Koeberlinia spinosa Zuce. Al- though the most luxuriant growth and fruiting occurs in rocky arroyo sites, the species is obviously more at home in the former habitat, as is evidenced by its more frequent occurrence there. The plant is often the only one (or one of a few) on otherwise bare silt. Its procumbent habit serves to bind the soil and forms hillocks down the sides of which the branches sprawl. In two of the seven collections studied fasciated stems were noted. These are flattened and falcate with two ranks of simple normal spines issuing from their edges. It is odd that two such cases of identical abnormality were encountered in so rare a plant, about fifty individuals being observed in the course of a wide and painstaking search. Bureau of Plant Industry, Washington, D. C., February, 1941. 1941] PECK: NEW COMBINATIONS 133 COMBINATIONS PROPOSED IN “THE HIGHER PLANTS : OF OREGON” Morton EK. Peck There has been some question as to the validity of certain new combinations proposed in the ‘“Manual of the Higher Plants of Oregon,’ recently published by the writer. The citations re- quired if one is to adhere strictly to the International Rules (Ch. 3, Sec. 6, Art. 44), are, therefore, published herewith. Numbers in parentheses refer to pages of the “Manual” on which the new combination was made. ToFIELDIA OCCIDENTALIS Wats. var. intermedia (Rydb.) comb. nov. I’. intermedia Rydb., Bull. Torr. Club 27: 528. 1900. (p. 189) BropiaEA Dovuerasu var. Howellii (Wats.) comb. nov. B. Howellii Wats., Proc. Am. Acad. Sci. 14: 801. 1879. (By error as B. grandiflora var. Howellii, p. 199) HaAseEnaRIA ELEGANS var. multiflora (Rydb.) comb. nov. Piperia multiflora Rydb., Bull. Torr. Club 28: 688. 1901. (p. 219) ERIOGONUM OVALIFOLIUM var. ochroleucum (Small) comb. nov. E. ochroleucum Small, Mem. N. Y. Bot. Gard. 1: 123. 1900. (p. 256) ATRIPLEX PATULA var. obtusa (Cham.) comb. nov. A. angusti- folia var. obtusa Cham., Linnaea 6: 569. 1881. (p. 267) Lewista CotyLepon var. Howellii (Wats.) comb. nov. Calan- drinia Howellii Wats., Proc. Am. Acad. Sci. 23: 262. 1888. (p. 281) ARENARIA ACULEATA var. uintahensis (Nels.) comb. nov. A. uintahensis Nels., Bull. Torr. Club 28: 7. 1899. (p. 286) SILENE orEGANA Wats. var. filisecta comb. nov. S. filisecta Peck, Proc. Biol. Soc. Wash. 47: 186. 19384. (p. 296) ANEMONE GLososa var. lithophila (Rydb.) comb. nov. A. lith- ophila Rydb., Bull. Torr. Club 29: 152. 1902. (p. 809) 'AQUILEGIA FoRMosa var. flavescens (Wats.) comb. nov. A. fla- vescens Wats., Bot. King Geol. Expl. 40th. Par. 5: 10. 1871. (p. 312) Dicentra CucuLiaria var. occidentalis (Rydb.) comb. nov. Bicuculla occidentalis Rydb., Bull. Torr. Club 29: 160. 1902. (p. 323) Draza rEPTANS var. Stellifera (Schulz.) comb. nov. D. caro- liniana f. stellifera Schulz., in Engler, Pflanzenr. 4°°: 333. 1927. (p. 326) Descurainta RicHaRDSONI var. viscosa (Rydb.) comb. nov. Sophia viscosa Rydb., Bull. Torr. Club 29: 238. 1902. (p. 840) Descurainia pinnata var. filipes (Gray) comb. nov. Sisym- brium incisum var. filipes Gray, Pl. Fendl. 8. 1849. (p. 340) DescuRAINIA PINNATA var. halictorum (Cockerell) comb. nov. Sophia halictorum Cockerell, Bull. Torr. Club 25: 460. 1898. (p. 340) 134 MADRONO [Vol.6 DEScURAINIA PINNATA var. paradisa (Nels. & Kenn.) comb. nov. Sophia paradisa Nels. & Kenn., Proc. Biol. Soc. Wash. 19: 155. 1906. (p. 340) Descurarnia PInNATA var. Nelsonii (Rydb.) comb. nov. Sophia Nelsoniti Rydb., Bull. Torr. Club 34: 436. 1907. (p. 341) PaRRYA CHEIRANTHOIDES var. lanuginosa (Wats.) comb. nov. P. Menziesii var. lanuginosa Wats. in Gray Syn. Fl. 1: 152. 1895. (p. 352) Streptanthus hastatus (Wats.) comb. nov. Caulanthus_ has- tatus Wats., Bot. King Geol. Expl. 40th. Par. 5:28. pl. 3. 1871. (p. 3538) Sedum glanduliferum (Hend.) comb. nov. Cotyledon glanduli- ferum Hend., Rhodora 32: 26. 19380. (p. 861) Sedum oregonense (Wats.) comb. nov. Cotyledon oregonensis Wats:, ProcAm. Acad, Sci, 18:2 373.5 18625 (paso) PorENTILLA GRAcILIs Dougl. var. dichroa (Rydb.) comb. nov. P. dichroa Rydb., N. Am. Fl. 22: 819. 1908. (p. 893) HorkE.ia Fusca var. capitata (Lindl.) comb. nov. H. capitata Lindl., Bot. Reg. 23, sub. pl. 1997. 1837. (p. 398) HorkELIia Fusca var. pseudocapitata (Rydb.) comb. nov. dH. pseudocapitata Rydb. in Howell, Fl. N. W. Am. 180. 1898. (p. 398) Horkeia Fusca var. parviflora (Nutt.) comb. nov. H. parwi- flora Nutt. ex. Hook. & Arn. Bot. Beechey Voy. suppl. 338. 18388. (p. 399) Horke.ia Fusca var. filicoides (Crum) comb. nov. Potentilla Douglasii var. filicoides Crum. Leafl. West. Bot. 1: 100. 1984. (p. 399) HorkELIA CoNGESTA var. nemorosa (Keck) comb. nov. H. con- gesta subsp. nemorosa Keck, Lloydia 1: 108. 1988. (p. 399) Geum gracilipes (Piper) comb. nov. Potentilla gracilipes Piper, Bull. Torr. Club 27: 392. 1900. (By error as Geum filipes, p. 402) Rosa pisocarPa var. ultramontana (Wats.) comb. nov. R. cali- fornica var. ultramontana Wats. in Brew. & Wats. Bot. Calif. 1: 87. 1876. (By error as R. pisocarpa var. transmontana, p. 404) CERCOCARPUS LEDIFOLIUs var. hypoleucus (Rydb.) comb. nov. C. hypoleucus Rydb., N. Am. Fl, 22: 424. 19138. (p. 407) Astragalus Laurentii (Rydb.) comb. nov. Homalobus Lau- rentii Rydb., Bull. Torr. Club 51:15. 1924. (p. 443) Astragalus subglaber (Rydb.) comb. nov. Homalobus sub- glaber Rydb., Bull. Torr. Club 51:17. 1924. (p. 444) ASTRAGALUS REVENTUs var. Hoodianus (Howell) comb. nov. A. Hoodianus Howell, Erythea, 1: 111. 1898. (p. 444) Astragalus lagopinus (Rydb.) comb. nov. Xylophacos lago- pinus Rydb., Bull. Torr. Club 52: 3872. 1925. (p. 445) AstracaLus Hooxerianus var. siskiyouensis (Rydb.) comb. nov. Phaca siskiyouensis Rydb., N. Am. Fl. 24: 340. 1929. (p. 446) 1941] PECK: NEW COMBINATIONS 135 AstraGaLus Forwoopu var. wallowensis (Rydb.) comb. nov. Atelophragma wallowense Rydb., Bull. Torr. Club 55: 122. 1928. (p. 447) 3 ASTRAGALUS coNJuNcTUs var. Sheldoni (Rydb.) comb. nov. Tium Sheldoni Rydb., N. Am. Fl. 24: 3938. 1929. (p. 448) ASTRAGALUS LENTIGINoOsuUS var. cornutus (Rydb.) comb. nov. Cystium cornutum Rydb., N. Am. FI. 24: 412. 1929. (p. 449) ASTRAGALUS LENTIGINOSUS var. platyphyllidium (Rydb.) comb. nov. Cystium platyphyllidium Rydb., N. Am. Fl. 24: 410. 1929. (p. 449) VIOLA PRAEMORSA var. major (Hook.) comb. nov. V. Nuttalli var. major Hook., Fl. Bor. Am. 1:79. 18380. (p. 486) VIOLA PRAEMoRSA var. linguaefolia (Nutt.) comb. nov. VJ. linguaefolia Nutt., Torr. & Gray, Fl. 1: 141. 1888. (p. 486) VIOLA PURPUREA var. atriplicifolia (Greene) comb. nov. V/V. atriplicifolia Greene, Pitt. 3: 38. 1896. (p. 486) EPILOBIUM GLANDULOSUM var. Cinerascens (Piper) comb. nov. E. cinerascens Piper, Proc. Biol. Soc. Wash. 31: 75. 1918. (p. 494) EPILOBIUM PANICULATUM var. Hammondi (Howell) comb. nov. E. Hammondi Howell, Fl. N. W. Am. 224. 1903. (p.497) MENZIESIA FERRUGINEA var. glabella (Gray) comb. nov. M. gla- bella Gray, Syn. Fl. N. Am. 2: 39. 1878. (p. 542) PuHLox speciosa var. occidentalis (Dur.) comb. nov. P. divari- cata var. occidentalis Dur., Proc. Acad. Nat. Sci. Phila. 3: 97. 1855. (p. 570) Putiox speciosa var. lanceolata (E. Nels.) comb. nov. P. lan- ceolata E. Nels., Rev. W. Am. Phlox 29. 1899. (p. 570) PHLOX LONGIFOLIA var. compacta (Brand) comb. nov. P. Stans- buryt var. compacta Brand in Engler, Pflanzenr. 4°°°: 67. 1907. (p. 571) PHtox tonairo.ia var. longipes (Jones) comb. nov. P. lineari- folia var. longipes Jones, Contrib. West. Bot. 12: 538. 1908. (p. 571) PHLox Lonairouia var. humilis (Dougl.) comb. nov. P. humilis Dougl. ex Hook. Fl. Bor. Am. 2: 72. 1838. (p. 571) PHLox LoNGIFOLIA var. Calva (Wherry) comb. nov. P. longi- folia subsp. calva Wherry, Proc. Acad. Nat. Sci. Phila. 90: 136. 1938. (p. 571) Puiox Doverasnu var. rigida (Benth.) comb. nov. P. rigida Benth. in DC., Prodr. 9: 806. 1845. (p. 571) Puiox Dovetasu var. Hendersonii (E. Nels.) comb. nov. P. condensata var. Hendersonii E. Nels., Rev. W. N. Am. Phlox 14. 1899. (p. 571) PuHtox pirFrusa var. longistylis (Wherry) comb. nov. P. dif- fusa subsp. longistylis Wherry, Proc. Acad. Nat. Sci. Phila. 90: 139. 1938. (p. 572) PHLox pirFusa var. subcarinata (Wherry) comb. nov. P. dif- 136 MADRONO [Vol. 6 fusa subsp. subcarinata Wherry, Journ. Wash. Acad. Sci. 29: 517. 1989. (p. 572) Puiox pirrusa var. scleranthifolia (Rydb.) comb. nov. P. scleranthifolia Rydb., Mem., N. Y. Bot. Gard. 1: 313. 1900. (p.: 572) Putiox Hoop var. canescens (Torr. & Gray) comb. nov. P. canescens Torr. & Gray, Pac. R. Rept. 27: 122. 1855. (p. 572) APLOPAPPUS CARTHAMOIDES var. rigidus (Rydb.) comb. nov. Pyrrocoma rigida Rydb., Bull. Torr. Club 27: 624. 1900. (p. 712) ApLopaPpPpus RAcEMosus var. halophilus (Greene) comb. nov. Pyrrocoma halophila Greene, Leafl. Bot. Obs. & Crit. 2: 16. 1909. (By error as A. racemosus var. halophiloides, p. 712) APLOPAPPUS RACEMOsUs var. duriusculus (Greene) comb. nov. Pyrrocoma duriuscula Greene, Leafl. Bot. Obs. & Crit. 2:16. 1909. — Go 712) APLOPAPPUS RACEMosuUs var. glomeratus (Nutt.) comb. nov. Homapappus glomeratus Nutt., Trans. Am. Phila. Soc. n. ser. 7: 331. 1840. (p. 712) APLOPAPPUS RACEMosus var. brachycephalus (Nels.) comb. nov. Pyrrocoma brachycephala Nels., Bot. Gaz. 37: 265. 1904. (p.— 713) APLOPAPPUS RACEMOSUS var. congestus (Greene) comb. nov. Pyrrocoma congesta Greene, Pitt. 3:23. 1896. (p. 713) APLOPAPPUS LANCEOLATUs var. tenuicaulis (D. C. Eat.) comb. nov. A. tenuicaulis D. C. Eat., Bot. King Geol. Expl. 40th. Par. 5: 160. 1871, (p) 718) APLOPAPPUS HIRTUS var. sonchifolius (Greene) comb. nov. Pyrrocoma sonchifolia Greene, Leafl. Bot. Obs. & Crit. 2:18. 1909. (p.7 18) 7 APLOPAPPUs HiRTUs var. lanulosus (Greene) comb. nov. Pyrro- coma lanulosa Greene, Leafl. Bot. Obs. & Crit. 2: 16. 1909. (p. 713) APLOPAPPUS UNIFLORUs var. Howellii (Gray) comb. nov. A. Howelliti Gray, Syn. Fl. 1°, suppl.: 446. 1886. (p. 713) ANTENNARIA LUZULOIDEs var. oblanceolata (Rydb.) comb. nov. A. oblanceolata Rydb., Mem. N. Y. Bot. Gard. 1: 409. 1900. (p. 741) ARTEMISIA CAMPESTRIS var. Spithamaea (Pursh) comb. nov. 4. spithamaea Pursh, Fl. Am. Sept. 522. 1814. (p. 768) ARTEMISIA CAMPESTRIS var. pycnocephala (Less.) comb. nov. Oligosporus pycnocephalus Less., Linnaea 6: 524. 1881. (p. 768) ARTEMISIA CAMPESTRIS var. pacifica (Nutt.) comb. nov. A. pacifica Nutt., Trans. Am. Phila. Soc. ser. 2,7: 401. 1841. (pp. 768) SENECIO TRIANGULARIs var. trigonophyllus (Greene) comb. nov. S. trigonophyllus Greene, Pitt. 3: 106. 1896. (p. 780) CREPIS RUNCINATA var. imbricata (Babe. & Steb.) comb. nov. C. runcinata subsp. imbricata Babc. & Steb., Carneg. Inst. Wash. Publ. 504: 102. 1938. (p. 804) 1941] STEBBINS AND LOVE: STIPA 137 CREPIS MODOCENSIS var. Subacaulis (Kellogg) comb. nov. C. occidentalis var. subacaulis Kellogg, Proc. Calif. Acad. Sci. 5: 50. 1873. (p. 805) CREPIS ATRIBARBA Var. Originalis (Babe. & Steb.) comb. nov. C. atribarba subsp. originalis Babe. & Steb., Carneg. Inst. Wash. Publ. 504: 162. 1938. (p. 806) CREPIS OCCIDENTALIs var. pumila (Rydb.) comb. nov. C. pumila Rydb., Mem. N. Y. Bot. Gard. 1: 462. 1900. (p. 806) Crepis Baxeri var. Cusickii (Eastw.) comb. nov. C. Cusicki Eastw., Bull. Torr. Club 80: 502. 1903. (p. 806) Willamette University, Salem, Oregon, August 9, 1941. AN UNDESCRIBED SPECIES OF STIPA FROM CALIFORNIA G. L. STepsins, Jr., AND R. M. Love Due to the growing interest in the native forage plants of Cali- fornia, particular attention is now being given by a number of workers to Stipa pulchra Hitchc., one of the most common and val- uable of the native perennial forage grasses in the valley and foothill regions of this state. Several people, including the pres- ent writers, have noticed that this species as recognized in the current manuals actually consists of two distinct types. One, with deep green foliage, relatively broad leaves, stiffer panicle branches, large glumes, thick, fusiform lemmas, and stout, stiff awns, is predominant in the outer Coast Ranges and the wooded parts of the Sierra Nevada foothills. This is typical S. pulchra, of which the type came from Healdsburg, Sonoma County. The other form, with somewhat glaucous foliage, narrower leaves, flexuous, often nodding panicle branches, smaller, narrower glumes, slender lemmas, and slender, often flexuous awns, occurs chiefly in the treeless parts of the inner Coast Ranges, the San Joaquin Valley (in scattered areas undisturbed by cultivation), the valleys of southern California, and the edges of the deserts. The two types have been given different common names, typical S. pulchra being known as purple needle grass, and the slender, interior type as nodding needle grass. The writers have observed these two needle grasses carefully during two seasons of collect- ing in the field, have grown and compiled extensive morphological data on several cultures of each, and have examined their chromo- somes at both mitosis and meiosis. From these observations enough evidence has accumulated to warrant the recognition of the interior type as a distinct species. It may be described as follows. Stipa cernua sp. nov. Folia glauca angusta; panicula ampla, ramis tenuis, flexuosis, cernuis; glumae inaequales, 12-19 mm. longae, 1-1.4 mm. latae, pallidae vel roseo-purpureae, semper 3-nervatae; lemma angusta, 5—10.5 mm. longa, cum fructu 0.6—1 138 MADRONO [Vol. 6 LEGEND @ @ S. PULCHRA O S. CERNUA @® O © S. PULCHRA & S. CERNUA TOGETHER O 3, O [) ® o® © SAN FRANCISCO 'N CALIFORNIA @ Fic. 1. Distribution of Stipa pulchra and S. cernua. mm. crassa, ad basim et supra nervis pubescens, parte superiore glabra; arista 6-11 cm. longa, flexuosa, scabra vel ad basim pubescens. | Mostly in large clumps, the basal leaves numerous, narrow, usually glaucous; culms several, mostly 60 to 90 cm. tall, middle culm leaves 1.2—2.4 mm. broad. Panicle ample, the basal portion often within the lowermost leaf sheath; panicle branches slender and flexuous or cernous; glumes scarious, pale or reddish purple, long acuminate, the lower 12-19 mm. long, the upper somewhat shorter and broader, 1—-1.6 mm. broad, both strictly 3-nerved; lemma 5—10.5 mm. long, 2.2—2.9 mm. broad when unrolled, with caryopsis 0.6—-1 mm. thick at maturity, 5- or sometimes 7-nerved, the callus acute, pubescence dense on the callus and on the lower 1941] STEBBINS AND LOVE: STIPA 139 Piatt 13. INFLORESCENCE AND Fiorets 1n Stipa. Fic. 1. Stipa pulchra Hitche., from Berkeley (Stebbins 2670), panicle branch X .33. Fie. 2. 8S. cernua, from Santa Barbara County (Stebbins 2875), panicle branch x 33. Fic. 3. S. pulchra, from Berkeley, glumes and lemmaX3.3. Fic. 4. S. cernua, type collection, glumes and lemma x 3.3. 140 MADRONO [Vol. 6 TABLE 1 Comparison of the Principal Characteristics of Stipa pulchra and Stipa cernua Width of next to uppermost culmecleat, (tear ee Colorsot leaves tac03 3 eee Character of panicle branches Length lower glume ........ Width upper glume ........ Number of nerves on upper SMMC oes kere Length of lemma’.......... Width of lemma (unrolled) . Number of nerves on lemma Thickness of mature cary- opsis including lemma .... Pubescence of lemma ....... S. pulchra 2.4-6 mm. deep green spreading, or slightly cernuous 15-26 mm. 1.4-2.2 mm. 3-5 75-13 mm. 3.3-4.8 mm. 5-9 1.0-1.4 mm. throughout, or at base and = on nerves to mid- dle or summit S. cernua 1.2-2.4 mm. glaucous slender, flexuous, or cernuous 12-19 mm. 1.0-1.6 mm. 3 5.0-10.5 mm. 2.2-2.9 mm. 5-7 0.6-1.0 mm. at base and on nerves’ one-third to four-fifths of way to summit Lenethoor sawn) 2 oe 60-90 mm. 60-110 mm. Ratio of awn to lemma ..... 5.9-9.8 8.7-14.5 Pubescence of awn _ below nrst-bendée.. Ws. 5 ae eae pubescent scabrous or slightly pubescent | one-fifth to one-fourth of the body of the lemma, the upper part of the lemma scabrous and with the nerves pubescent one-third to four-fifths of the distance to the summit, lemma usually pale straw- or buff-colored, occasionally purplish, brownish, or black- ish; awn 6-11 ecm. long, 8.7—14.5 times the length of the lemma, twice bent at maturity, slender and flexuous beyond the second bend, scabrous or short-pubescent below the first bend. California, from Tehama County south to San Diego County east to the Sierra Nevada foothills, reaching the coast from Mon terey southward, and extending up to about 1360 meters (450¢ feet) altitude in southern California. 7 Type. West side of Cedar Mountain Ridge, southeast of Liver- more, Alameda County, altitude 400 meters (1300 feet), G. L. Stebbins, Jr. 2732, (Herb. Univ. Calif. 641631). The follow- ing specimens in the University of California Herbarium are also typical: west of Orland, Glenn County, Heller 11434; Nacimiento River, Monterey County, Davy 7688; Santa Barbara, Elmer 3874; Mount Pinos, Ventura County, Hall 6426; Mint Can- yon, east of Saugus, Los Angeles County, Munz 6794; near Rialto, San Bernardino County, S. B. §& W. F. Parish 2038 ; San Bernardino Valley, Parish 6204, 11257; near Winchester, Riverside County, Hall 2921; Box Springs Mountain, Riverside County, Hall 2977; San Diego, Brandegee 833. As mentioned previously, Stipa cernua is most closely related to S. pulchra. The differences between the two species are sum- 1941] STEBBINS AND LOVE: STIPA 141 marized in Table 1, and partly illustrated in plate 13. The follow- ing may be used as convenient key characters: ) Leaves green, the middle culm leaves 2.4-6 mm. broad; lemmas fusiform, with caryopsis 1-1.4 mm. thick at maturity; awn stout and stiff, mostly 7-9 times as long as the lemma ....... S. pulchra Leaves somewhat glaucous, the middle culm leaves 1.2-2.4 mm. broad; lemmas slender, with caryopsis 0.6-1 mm. thick at maturity; awn slender, flexuous beyond the second bend, mostly 9-12 times the length of the lemma ......................... S. cernua The distribution of the two species, as determined by the col- lections and observations of the writers as well as by the speci- mens in the University of California Herbarium, is shown in figure 1. It will be seen that they occur together throughout a rather large area, and can often be found growing side by side. In these places they are usually quite distinct, but forms inter- mediate between them do occur. The writers studied particularly these intermediate plants as found in two localities near San Benito, San Benito County, and found them to be completely sterile or nearly so. Although the surrounding plants of S. pulchra and S. cernua were producing good seed in abundance, not a single fertile grain was found on any of the intermediates. The latter showed considerable hybrid vigor, often forming clumps much larger than those of the parent species. In the University of Cali- fornia Herbarium there are three specimens of such sterile inter- mediates: north base of Mount Hamilton, Santa Clara County, Sharsmith 664A; Pacific Grove, Monterey County, Elmer 3507; Las Flores Canyon, Santa Monica Mountains, Los Angeles County, Epling in 1930. Only the latter specimen had anthers with pollen for examination; in it 5 per cent of the pollen grains were large and well filled with cytoplasm, as contrasted with 90— 96 per cent in typical S. pulchra and S. cernua. The chromosome numbers of the two species are reported else- where as 2n = 64, n= 82 for S. pulchra, and 2n=70, n=8385 for S. cernua (Stebbins and Love, Am. Jour. Bot. 28: 371-382. 1941). Three of the sterile intermediate plants from San Benito were dug up and transplanted to pots in Berkeley, and their somatic chro- mosome number was determined as 2n = 67, indicating that they are actually first generation hybrids between S. cernua and S. pulchra. Their meiosis will be studied during the coming season. Except for S. pulchra, S. cernua has no close relative among the North American species of Stipa. It resembles S. comata in its glaucous leaves and long awns, but that species has large, thick lemmas as in S. pulchra, and lacks the collar at the apex of the lemma which is found in both S. pulchra and S. cernua. It is pos- sible that S. cernua is related to some of the numerous South American species of Stipa, but the present writers have not seen adequate material of any of them to judge their relationships. College of Agriculture, University of California, Berkeley, January, 1941 142 MADRONO [Vol. 6 REVIEWS The Evolution of Land Plants | Embryophyta|. By Doveuas Hoventon CampBeLyt. Pp. 1-731 with 351 text figures. Stan- ford University Press. 1940. This large volume has two main elements. It is a résumé of the author’s half century of morphological, chiefly embryological, study; and it is a condensed summary of the work of other writers on the relationships of the higher plants. Of these, the former is the more valuable, which is natural since Dr. Campbell has him- self been the foremost contributor to our understanding of the broader lines of the evolution of the land plants. It is as to these broad lines, where the work of the past fifty years has brought confidence in some things and doubt as to others, that this book registers well the present state of science and can serve as a mile post. The larger part of the book is taken up by the finer classifica- tion, to orders, families, in some places to genera. Here the author depends more upon the views of others, and the presenta- tion is distinctly less authoritative. For example, among the ferns, the Eusporangiatae are well presented, but the treatment of the higher Leptosporangiatae is comparatively weak. In accord with custom, the evidence of paleophytology is treated with respect. This science has of course made progress in various respects. But the reviewer would still recall a remark of Dr. Joseph Hooker. “Amongst the many collections of fossil plants that I have examined, there is hardly a specimen, belonging to any epoch, sufficiently perfect to warrant the assumption that the species to which it belonged can be recognized.” Yet, specific characters may petrify better than those of classes. Consider spermatozoids and the embryo-sac. A discussion of the nature and value of evidence would be a valuable introduction to a book of this kind. Next to its completeness, the most marked characteristic of the book is its lack of dogmatism. Correlated with caution is a tendency to entertain the idea of multiple origin of apparently natural groups, and to admit question, even where affinity seems best established. Two examples: the probable central position of some such plant as Anthoceros in the ancestry of Embryophyta is perfectly presented, and the group “‘Anthocerotes” is made a class, coordinate with Hepaticae and Musci. The latter are treated as probable derived groups, but the derivation seems to be pictured as exceedingly ancient, from primeval Anthocerotes, of which “Of course, the sporophyte . . . must have been much simpler than in any living forms—perhaps comparable to that of such liverworts as Riccia or Sphaerocarpus.” To the reviewer, the stoma, common to Anthoceros and many mosses, provides positive proof that their common ancestor, if not exactly Anthoceros itself, had at any rate a sporophyte independent enough in its nutrition to have evolved this structure. 1941] . REVIEWS 143 As to the angiosperms, polyphylesis is explicitly advocated. The characteristic structure of this group is not seed, nor flower, nor pollen tube, nor trachea; it is the embryo-sac. If the hypoth- esis of multiple origins means that this structure has been evolved several times independently, it is hard to accept. Even more than the stoma, it requires good evidence of repeated evolution before it is questioned as proof of real affinity. There are 351 numbered figures, most of which are composed of a considerable number of drawings, largely original and well reproduced. They add materially to the value of the book. The text is a remarkable mine of detailed information. How much there is of this ntay be shown by the index, which occupies 37 pages of fine print, two columns to the page, and is still incom- plete; thus, under “stoma” there is no reference to the text, and under “‘embryo-sac”’ there is only one.—E. B. Copreitanp, Depart- ment of Botany, University of California, Berkeley. Sinopsis de la Flora del Cuzco. Fortunato L. Herrera. Tomo I. Parte Sistematica. Pag. 1-528. Publicado bajo los auspicios del supremo gobierno. Lima, Peru, 4 de Julio, 1941. This check list of the plants of the Department of Cuzco by the distinguished Peruvian botanist is by far the most complete of several similar works by the same author, the first of which ap- peared in 1919. It lists 2166 species (with a few varieties) 588 of which are cryptogams, about 250 of these being ferns. and fern allies. Even so, the author suggests that probably only about one-half of the species growing within the area have been re- corded. The predominant families are Compositae, Gramineae, and Leguminosae. The names are accompanied, at least for the phaenerogams, by source of publication and citation of specimens. The latter are given in detail, usually including altitude, information which will be invaluable in any study of the flora; habitats, however, are rarely indicated. Often the range of the plant outside of Cuzco, if known, is mentioned; there are some economic notes. An appendix contains descriptions of new species based on the author’s collections. There is also a list of native names and their scientific equivalents, and an index to the genera. In a work of this nature, based of necessity on the literature available—of which there is a good bibliography—there are of course always omissions; on the other hand there are a few addi- tions to the flora of Cuzco. In supplements, which it is to be hoped will be issued from time to time, it would be well to give the source of determination and to indicate where the collections may be consulted in order that identifications may be checked when desired. The work would be more consistently useful, too, if publication citations were always given (which is obviously the intent but they are not infrequently omitted). Most of the typo- graphical errors will easily be corrected; only one mistake in the 144 MADRONO ; [Vol. 6 presentation of the material has been noted, namely the including of the composite Orthopappus, on page 321, in the Melasto- mataceae. The author in preparing the work and the Peruvian Govern- ment in publishing it have made a meritorious contribution to the scientific study of the rich and useful flora of Cuzco; may there be many more similar endeavors based increasingly on the activities of Peruvian students.—J. F. Macsripre, Field Museum of Natural History. A Flora of Arizona and New Mexico. By Ivar Tipestrom and Sister TeresiTa KiTTELL. Pp. xxvi+ 897 with frontispiece. The Sn University of America Press, Washington, D. C., 1941. 5.00. The flora of Arizona and New Mexico, listing 898 genera and 3975 species, is arranged according to the systems of DeCandolle and Bentham and Hooker with some slight emendations, chief of which is the arrangement of the orders and families in a descend- ing numerical sequence as to the number of cotyledons. Hence the Coniferae with many cotyledons come first and the ferns and fern allies with none appear at the close of the work. The keys are brief and to the point and brief descriptions aid materially in amplifying the keys. There is a general citation of habitat and range accompanying each entity. The work is ambitious and as such is worthy but one cannot read it without a feeling of regret. Much of the advance in botany of the past fifteen years is ignored. Many monographs which have appeared during this time are not alluded to either as to the species accepted or in the synonymy. The frontispiece is a map of Arizona and New Mexico showing the major rivers and the two thousand foot contour intervals. There are fifteen circles indicating localities but no evident refer- ence to these in the text. On the other hand the table of contents refers to the map as showing the “belts of vegetation.” The reason for this confusion is not clear to the reviewer. The volume is lithoprinted and would have been materially improved by either a little more space between the species or by underlining the species names. As it stands the pages appear crowded and the typography does not invite the reader’s atten- tion. In many instances the craftsmanship of both typist and printer is definitely at fault—Hersert L. Mason. NOTES AND NEWS On May 24, 1941, the University of California conferred the honorary degree of doctor of laws on Dr. Willis Linn Jepson, Pro- fessor of Botany Emeritus of that institution. The honor is in recognition of Dr. Jepson’s contribution to our knowledge of the California flora and his long and successful promotion of forest conservation in the state. MADRONO A West American Journal of Botany A quarterly journal devoted to important and stimulating articles dealing with plant morphology, physiology, taxonomy, and botanical history. These volumes should be a part of every botanist’s li- brary and should be made accessible to students of all universities and colleges. Volume I, 1916-1929. . . $5.00 Volume II, 1930-1934 .. 5.00 Volume III, 1935-1936 . 5.00 Volume IV, 1937-1938 . 5.00 Volume V, 1939-1940 . . 5.00 Single numbers....... 0.75 The subscription price of MADRONO is $2.50 per year. We solicit your pat- ronage, Address all orders to: . William Hiesey Bus. Mgr. Carnegie Institution of Washington Stanford University, California VOLUME VI NUMBER 5 MADRONO A WEST AMERICAN JOURNAL OF BOTANY gi eT NAEP. mn, nes ‘AQONIAN INS7/FE2S wo iv aN Contents _ A Synopsis or THE AMERICAN Spectres oF Cicuta, Mildred E. Mathias and UIC ONGLADICON hi ween tere nt ete emu O NEU a ye Oa Na Wow hai 145 Great Basin Prants—VI. Nores on GENTIANA, Bassett Maguire ........ 151 Arnica In ALASKA AND YUKON, Bassett Maguire .................0000005 153 _ A Comparison or THE Empryoceny or Picea anp Asixs, J. T. Buchholz .... 156 _ ANATOMY AnD EcoLocy oF AMMOPHILA ARENARIA LINK, Edith A. Purer ... 167 An UNDEscrizep SPeciIes or CEANOTHUS FROM CaLirorniA, Howard EH. Mc- ee coli PE a TRS ECT ES Ee Sea Ca a gaan a etn A ae aE: Gn 171 - Norss anp News: Range Extensions in Species of Western North America, Boykinia Jamesii var. heucheriformis and Saxifraga eriophora (Bassett Maguire), Eriodictyon capitatum (John Tucker); News ............. 173 _ PRocEEDINGs oF THE CALIFORNIA BOTANICAL SOCIETY ..................-. 175 Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania January, 1942 MADRONO A WEST AMERICAN JOURNAL OF BOTANY Board of Editors Hersert L. Mason, University of California, Berkeley, Chairman. LeRoy Asrams, Stanford University, California. Epcar ANnpERsSON, Missouri Botanical Garden, St. Louis. Lyman Benson, University of Arizona, Tucson. Hersert F. Copetanp, Sacramento Junior College, Sacramento, California. Ivan M. Jounston, Arnold Arboretum, Jamaica Plain, Massachusetts. Mirprep E. Maruias, University of California, Berkeley. Bassett Macurre, Utah State Agricultural College, Logan. Marion Ownsey, State College of Washington, Pullman. Secretary, Editorial Board—EtuHet Crum Department of Botany, University of California, Berkeley Business Manager—Wiut1i1am Hirsey 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 proofs are returned. Published at North Queen Street and McGovern Avenue, Lancaster, Pennsylvania, for the CALIFORNIA BOTANICAL SOCIETY, INC. President: Ernest B. Babcock, University of California, Berkeley. First Vice-President: Roxana S. Ferris, Stanford University, California. Second Vice-President: Palmer Stockwell, Institute of Forest Genetics, Placerville, California. Treasurer: Wiiliam Hiesey, Carnegie Institution of Washington, Stanford University, California. Secretary: Lincoln Constance, Department of Botany, University of California, Berkeley. Annual membership dues of the California Botanical 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. 1942] MATHIAS AND CONSTANCE: CICUTA 145 A SYNOPSIS OF THE AMERICAN SPECIES OF CICUTA Mixtprepo EF. Maruias anp LIncoLtn CONSTANCE The water hemlocks are of especial interest because of their poisonous properties, which make them a menace both to livestock and human beings. In a recent paper, Bomhard (1) remarks that, “The extremely toxic character of the underground parts ranks this genus as one of the most virulently poisonous groups of flowering plants native to the North Temperate Zone.” Hence, a great deal of space has been devoted to them in agricultural bulletins, some of which have directed considerable attention to the specific differences allegedly discernible in the subterranean portions of the plants. It is the purpose of the present paper to * evaluate the characters which have been used to delimit species in the genus, and to propose a somewhat revised taxonomic treat- ment. Specific descriptions for Cicuta will appear in a forthcom- ing number of “North American Flora.” In their pioneer revision of North American Umbelliferae in 1888, Coulter and Rose (2) referred the known members of the genus in America to three species, the European Cicuta virosa L. (with two varieties), C. Bolanderi Gray, a plant endemic to the California salt marshes, and C. bulbifera L. Cicuta bulbifera, easily separable from the other species by virtue of the usually asexual reproduction through the agency of bulblets in the axils of the upper foliage leaves, will not enter into the following discussion. This rather simple picture of the genus was upset the following year by Greene (4), who excluded C. virosa from America, and described three additional species from the western United States, C. occidentalis, C. purpurata, and C. vagans. At the same time, he enunciated his faith in the value of vegetative characters in dis- tinguishing the species of water hemlocks in the following words: “In the few families of plants which are, like the Umbelliferae, preéminently natural, the anthological and carpological charac- ters, whether of genera or of species, are apt to be very slight. But here Nature comes usually to the rescue of the despairing car- pological systematist, and gives him good characters for his genera, or for his species, in the vegetative organs. Only by re- garding these latter can a man set good limits to species in such a genus as Cicuta.” Again, in November of the same year, he re-emphasized (5) and elaborated this view, as follows: “The fact is well recognized, or should be, by descriptive botanists, that in herbaceous plants of all kinds, characters of the roots or other subterranean organs are of the very best for specific distinctions. Those of pubes- cence, foliage, and to some extent, of the flower also, are less con- stant within specific limits than are the peculiarities of the root, when the root happens to have peculiarities, which is however by Manproxo, Vol. 6, pp. 145-176. January 17, 1942. 146 MADRONO [ Vol. 6 no means the rule in nature. Most commonly the roots, rhizomes, tubers and other such organs will be much the same throughout. the whole group—a long series of species, or even an entire genus. In an order so extremely natural as that of the Umbelliferae, in which the fruits are so similar that plants of the same carpology are sometimes placed in different genera in deference to merely vegetative differences, it would seem altogether unphilosophical to require of the fruit that it furnish specific characters; or, to assume that unless the supposed members if a genus can be dis- tinguished carpologically the species is but one.” Coulter and Rose, in their later treatment of the Umbelliferae (3), accepted two of the three species Greene had described, but judged C. purpurata to be conspecific with C. Douglasi (DC.) C. & R. They added to the list of species C. Curtissii, conspecific with C. mexicana which they had already described earlier in the same year. Although they accepted Greene’s species, they were not entirely convinced of the validity of the characters he had origin- ally employed in distinguishing them: “We reproduce the above key in the hope that it may be further tested in the field, for with the material at our command we have not been able to follow it fully. While we recognize in Professor Greene’s typical mate- rial the differences suggested, we do not find them constant. The fleshy thickening of the rootstocks and their direction, as well as the thickness and elongation of the roots, seem to vary with the nature of the substratum, as might be expected.” The synoptical key offered by Greene, modified by the inclusion of the species he later described follows. * Root axis very short, nearly or quite erect, not enlarged, its partitions crowded. + Roots all alike, slender fibrous. C. virosa. ++ Main roots coarse, elongated, fleshy fibrous. C. Bolanderi, C. occidentalis, C. purpurata, C. frondosa, C. arguta, C. Sonne. +++ Main roots oval or oblong, fleshy tuberiform. C. maculata, C. bulbifera, C. subfalcata. ** Rhizomatous species; the root axis greatly enlarged, hori- zontal, only partly or not at all subterranean, emitting fibrous roots from beneath. C. vagans, C. californica, C. grandifolia, C. dakotica. Incapable of placement in this scheme are the three following species, also described by Greene: C. valida, “stature of the plant and its underground parts not known’; C. fimbriata, described only from the foliage; C. ampla, “known to me only in the fruiting summit of a single plant.” EXPLANATION OF THE Figures. PLatTe 14. ° Pirate 14. Frurrs or Cicura. Longitudinal view (x 6.5) and cross section (x 9.5). Fras. 1, 2, C. Bolanderi; 3, 4, C. virosa; 5, 6, C. mexicana; 7, 8, C. Doug- lasii; 9, 10, C. Victorinii; 11, 12, C. mackenzieana; 13, 14, C. maculata. MATHIAS AND CONSTANCE: CICUTA 147 1942] Puatr 14. Fruits or Cicuta. 148 MADRONO [Vol.6 Illustrating their scepticism of the very foundation of Greene’s classification, Coulter and Rose proceeded to base their own key on their observation that the “species seem to be best grouped primarily by their oblong or orbicular fruits, further separation being made upon differences in the fruit ribs and in the foliage.” Greene, however, seems not to have been at all discouraged and described C. grandifolia in 1907, and C. frondosa, C. subfalcata, C. dakotica, C. arguta, C. valida, C. Sonnei, C. fimbriata and C. ampla in 1912. Cicuta cinicola A. Nels. was added from Idaho, also in 1912, and recently C. mackenzieana Raup has been described from northern Canada and C. Victorini Fernald from the mouth of the St. Lawrence River. Authors of manuals and floras have not been prone to accept Greene’s second deluge of segregates, but have admitted C. occidentalis or C. vagans or both to their treat- ments of the appropriate areas. In preparing a revision of this genus for our account of the Umbelliferae in the “North American Flora,’ we have had occa- sion to re-examine the material on which Greene based his con- clusions. We are in agreement with him that Cicuta virosa is to be excluded from North America, although its relationship to C. mackenzieana appears to be rather close. We question the whole thesis that the underground parts of the water hemlocks afford specific characters, at least of the nature noted by Greene, and believe that they are so susceptible of modification in relation to soil composition and fertility, and especially to soil aération, as to be essentially useless in this connection. Furthermore, we are unable to attach sufficient importance to the degree of leaf-divi- sion to warrant the retention of C. californica as distinct from C. Douglasii, and find that Coulter and Rose’s distinction between “oblong” and “‘orbicular” fruits sometimes breaks down on the same plant. The diagnostic value of leaf-venation as a means of distinguishing Cicuta from other umbelliferous genera, and also for separating individual species in the former genus, has been stressed recently by Bomhard (1). She distinguishes C. cali- fornica from the other species by the fact that the secondary veins of the leaflets are directed at the marginal teeth, rather than at the sinuses between them. Our studies, however, have not enabled us to find any marked difference in this entity, which would warrant its continued retention as a species. On the other hand, characters of the fruit, especially its com- pression, the proportion between the dorsal and lateral ribs, their relation to the intervals, and the nature of the oil tubes and the seed do seem to enable us to distinguish natural species in the genus. Our treatment, admits the following species for North America: C. Bolanderi, C. meaicana, C. Douglasi, C. mackenzieana, C. maculata, C. Victorinit and C. bulbifera. All of the species pro- posed by Greene will thus be relegated to the synonymy which they appear richly to deserve. 1942] MATHIAS AND CONSTANCE: CICUTA ; 149 Taxonomic TREATMENT Grcuta L. Sp. Pl. 12 255. 1753. Creutaria Lam. FI]. France 3: 445. 1778. Keraskomion Raf. New. Fl. Amer. 4: 21. 18386. Type species: Cicuta virosa L. Axils of the leaves not bulbiferous. Fruit constricted at the commissure; lateral ribs about equalling the dorsals in surface display. Oil tubes large (pl. 14, figs. 2, 6); seed oily, evidently channeled under the tubes. Ribs narrower than the oil tubes: plants of Pacific GoOase Gall MALESHES! fies 8 Sana e ees os 1. OC. Bolanderi. Ribs broader than the oil tubes; plants of coastal eastern and southeastern United States and ECASLEIM, WLCXICO!. . 1g.) eo Sear he eS 2. C. mexicana. Oil tubes small (pl. 14, figs. 8, 12); seed less oily, un- channeled or only slightly channeled under the tubes. Fruit oval to orbicular, at least as long as broad, 2-4 mm. long, 2-3 mm. broad; rays 12-20, 2-6 cm. long; pedicels 3-8 mm. long .............. 3. C. Douglasit. Fruit elliptical, conspicuously broader than long, 1.5-2.2 mm. long, 2-3 mm. broad; rays 7-14, 7-8 cm. long; pedicels 7-12 mm. long ........ 4. C. mackenzieana. Fruit not constricted at the commissure; lateral ribs much broader than the dorsal in surface display. Leaflets coarsely serrate to incised; fruit oval to orbicular, rounded at apex and base; lateral ribs prominent, about equalling the intervals ........ 5. C. maculata. Leaflets finely serrate; fruit ovate, narrowed toward apex, cordate at base; lateral ribs obscure, much narrower than the intervals ................... 6. C. Victorinii. Axils of the leaves bulbiferous ......................0. 7. C. bulbifera. 1. Crcuta BoLtanperi Wats. Proc. Amer. Acad. 11: 189. 1876. Type locality. Suisun, Solano County, California, in salt marshes, Bolander. Distribution. Salt marshes of central and southern Cali- fornia. Representatives. Davy 4106, 6668, 6789, 6877; Heller 7541. 2. CicuTA MEXICANA Coult. & Rose, Proc. Wash. Acad. 1: 145. (January) 1900. C. maculata L. sensu Hemsl. Biol. Centr.-Amer. Bot. 1:566. 1879-81, not C. maculata L. 17538. C. Curtissii Coult. & Rose, Contr. U. S. Nat. Herb. 7: 97. (December) 1900. C. maculata var. Curtissii Fern. Rhodora 41: 439. 19389. Type locality. Coatzacoaleos, Isthmus of Tehuantepec, Vera Cruz, Chas. L. Smith 1161, Distribution. New Jersey to Florida, south and west to Tamaulipas, Nuevo Leon and Vera Cruz. Representatives. Curtiss 6845; Heller 1165; Palmer 445; Pringle 10,804. 38. Cicuta Doverasi (DC.) Coult. & Rose, Contr. U. S. Nat. Herb. 7: 95. 1900. Sium Douglasti DC. Prodr. 4: 125. 18830. Cicuta maculata L. sensu Hook. & Arn. Bot. Beechey Voy. 142. esaiy> not L. 1753. C. californica Gray, Proc. Amer. Acad. 7: 150 MADRONO [Vol. 6 344, 1867. C. crassifolia Nutt. Rept. Wilkes Exped. 17: 3816. 1874. C. virosa var. californica Coult. & Rose, Rev. N. Amer. Umbel. 1380. 1888. C. occidentalis Greene, Pittonia 2:7. 1889. C. occidentalis f. frondosa Greene, op. cit. p. 7. C.purpurata Greene, op. cit. p.8. C.vagans Greene, op. cit.p.9. C.grandifolia Greene, Leafl. Bot. Obs. 2: 124. 1909. C. Douglasii var. occidentalis Jones, Bull. Univ. Mont. Biol. ser. 15, 42. 1910. C. Sonnei Greene, Leafl. Bot. Obs. 2: 239. 1912. C. subfalcata Greene, op. cit. p. 237. C.frondosa Greene, op. cit. p. 236. C. valida Greene, op. cit. p. 238. C. fimbriata Greene, op. cit. p. 240. C. cinicola A. Nels. Bot. Gaz. 54: 141, fig. 1. 1912. C. occidentalis f. californica Wolff ex Engl. Pflanzenr. 4°78: 90: 82. 1927. C. occidentalis f. oregonensi-idahoensis Wolff, op. cit. p. 82. C. occidentalis f. arizo- nensis Wolff, op. cit. p. 82. C. occidentalis f. wyomingensis Wolff, Op City. (82; Type locality. “In America boreali-occid.,’’ Douglas. Distribution. Alberta and Montana to western Alaska, south to California, New Mexico, Arizona and Chihuahua. Representatives. Baker 655; Brown 501; Cusick 2556, 2779; Heller 7174; Nelson & Macbride 1815; Townsend & Barber 57. 4. CICUTA MACKENZIEANA Raup, Journ. Arn. Arb. 17: 279, pl. 197. 1986. Type locality. Sandy margin of a lagoon near the south shore of Lake Athabaska about 1.5 miles west of Ennuyeuse Creek, Canada, August 25, 1935, Raup 6976. Distribution. Hudson Bay to the Mackenzie Basin. Representatives. Macoun 79,261; Raup 6764, 6964. 5. Crcura macutaTa L. Sp. Pl. 1: 256. 1758. C. maculata Lam. Encycl. 2: 2. 1786. C. virosa var. maculata Coult. & Rose, Rev. N. Amer. Umbel. 1380. 1888. C. dakotica Greene, Leafl. Bot. Obs. 2: 237. 1912. C. arguta Greene, op. cit. p. 2388. G: ampla Greene, op. cit. p. 241. C. dakotica var. pseudovirosa Lun- nell, Amer. Midl. Nat. 4: 486. 1916. C. dakotica var. pseudo- maculata Lunell, op. cit. p. 486. Type locality. Virginia, Kalm. Distribution. Prince Edward Island and Quebec to North Carolina and Tennessee, west to North Dakota and Texas. Representatives. Fernald & St. John 1141; M. L. Grant 3238; Heller 1002; Lindheimer 615. 6. Cicuta Victrorinu Fern. Rhodora 41: 441, pl. 561, figs. 1-2. 1939. Type locality. Tidal flats of the St. Lawrence River, Quebec: “greves intercotidales, Cap Rouge pres du Pont de Quebec, 9 aout 1922,” Victorin 15,479. Distribution. Known only from the estuary of the St. Lawrence River, Quebec. Representatives. Fernald & Long 24,249; Victorin 15,480. 1942] ' MAGUIRE: GREAT BASIN PLANTS 151 7, CIcUTA BULBIFERA L. Sp. Pl. 1: 255. 1753. Cuicutaria bulbi- fera Lam. Encycl. 2:3. 1786. Keraskomion bulbiferum Raf. New. Pl4:-21. 1886. Type locality. Virginia, Canada, Clayton. Distribution. Newfoundland and Quebec to Delaware and Pennsylvania, west to British Columbia and Oregon. Representatives. Cusick 2966; Heller §& Heller 551; Sandberg, MacDougal 6 Heller 789. Department of Botany, University of California, Berkeley, May, 1941. LirerAtTure C1rep 1. Bomuarp, Mira1am L. Journ. Wash. Acad. Sci. 26: 102-107. 1936. 2. Couuter, Joun M. & J. N. Rose. Revision of North American Umbelliferae. 129-130. 1888. 3. —. Contr. U.S. Nat. Herb. 7: 93-100. 1900. 4. GREENE, E. L. Pittonia 1: 271-272. 1889. 5, —————_—_—.. Pittonia 2: 1-1l. 1889. GREAT BASIN PLANTS—VI. NOTES ON GENTIANA Bassett MaAGuIRE Continuing the series of minor papers discussing plants of the Great Basin, these notes are concerned with the delineation of a newly recognized geographical population of Gentiana calycosa, and the confirmation of a range extension of G. barbellata. All specimens herein cited are on deposit at the Intermountain Herbarium. The United States Forest Herbarium, Washington, D. C., is designated by the symbol USFH. GENTIANA BARBELLATA Engelm. The following collection con- firms the queried inclusion of the species within the Utah range by Tidestrom. Urau. Sanpete County: frequent, stony, steep, west-facing slopes, summit of Horse Shoe Mountain, South Peak, 12,000 feet, Manti National Forest, August 11, 1940, Maguire 20059. GENTIANA CALycosa Griseb. subsp. typica nom. nov., G. calycosa Griseb. in Hook. Fl. Bor. Amer. 2: 58. 1838. Dasystephana ob- tus.loba Rydb., Bull. Torr. Bot. Club 40: 464. 1918. D. monticola Rydb. l.c. GENTIANA caLycosa Griseb. subsp. asepala subsp. nov. Herbae perennes parvulae; caulibus decumbentibus, 5-10(12) cm. longis; foliis ovatis vel ovato-ellipticalibus 1.0-1.5(1.7) cm. longis; flori- bus solitariis; corollis 3-4 em. longis; calicibus membranaceis, graviter incisis, lobis obsoletis vel inconspicuis subulatisque, rare excedentibus 1-1.5 mm. Low perennial herbs; stems decumbent, 5—10(12) cm. long; leaves ovate to ovate elliptical 1.0-1.5(1.7) cm. long; flowers solitary, corolla 8-4 em. long; calyx membranaceous, character- 152 MADRONO [Vol.6 istically deeply incised on two sides, lobes obsolete, or incon- spicuous and subulate, rarely more than 1—1.5 mm. long. Type. Meadows about seepage areas, southeast slopes, saddle west of Mount Agassiz, 11,500 feet elevation, Uinta Moun- tains, Duchesne County, Utah, August 15, 1933, B. Maguire, Ruth Maguire, & A. G. Richards, 4225. Specimens examined. IpauHo. Custer County: meadow, Steward Canyon, 8000 feet, Lemhi National Forest, July 17, 1981, S. L. Jacobs 90 (USFH, 65577). Idaho County: Floyd Meadow, 6000 feet, Idaho National Forest, August 8, 1930, C. Gray CG57 (USFH, 67318). Lemhi County: Allen Lake Meadow, 8000 feet, Salmon National Forest, September, 1930, A. H. Wheeler 70 (USFH, 84421). Valley County: Nameless Meadow, 6500 feet, Fayette National Forest, September 1, 1930, L. N. Wellman 18 (USFH, 67628) mixed with G. affinis. Blaine County: alpine slopes, base Devil’s Bedstead, Sawtooth Range, 8000 feet, July 28, 1936, J. W. Thompson 13549. Custer County: damp, springy soil, Toxaway Lake, 10 miles west-southwest of Obsidian, Sawtooth Mountains, 8500 feet, August 8-11, 1937, C. L. Hitchcock & J. S. Martin 5749; Mount Hyndman, August 11, 1939, Ray J. Davis 1704. Nevapa. Elko County: meadow, Ruby Ranger Station, 6000 feet, Humboldt National Forest, June 28, 1930, L. E. Mc- Kenzie 28 (USFH, 64353). Uran. Summit County: meadows, Lily Lake, 10,500 feet, 3 miles west of Bald Mountain, Uinta Mountains, August 14, 1983, Maguire et al. 4224; bogs, west shore Henry’s Fork Lake, 10,850 feet, Uinta Mountains, August 4, 1936, Maguire et al. 14381. Uintah County: wet place under spruce, south base of Liedy Peak, 10,000 feet, Uinta Mountains, August 21, 1939, Maguire 17678. Specimens from Oregon intermediate to subsp. typica are: Baker County: Antony Lakes Region, Blue Mountains, 7100 feet, July 28, 1936, J. W. Thompson 13430. Wallowa County: meadows, Mirror Lake, Eagle Cap Peak, Wallowa Mountains, 7500 feet, September 24, 1938, C. W. Shar- smith 3971. The subspecies of Gentiana calycosa may be distinguished as follows: Subspecies asepala Stems conspicuously decumbent, 5-10 (12) cm. long. Leaves 1.0-1.5(1.7) cm. long. Corollas 3—4 cm. long. Calyx membranaceous, deeply incised, lobes obsolete, or inconspicuous and mostly subulate, 1.0-1.5(2.0) cm. long. A Great Basin-Intermountain race, apparently confined to Utah, Nevada — and Idaho. Intermediate plants are known from eastern Oregon. Subspecies typica Stems erect, or slightly decumbent at the base, (8)10—-20 em. long. Leaves (1.5)2.0-3.0 cm. long Corollas 3.5-4.5(5.0) em. long. Calyx membranaceous mainly in the sinuses, not at all or rarely incised, lobes foliaceous, ovate, elliptic, or lanceolate, 5-10(15) mm. long. The population of the Rocky Moun- tains and the Sierra Nevada, ex- tending into Alberta and British Columbia. 1942] MAGUIRE: ARNICA IN ALASKA AND YUKON 153 The smaller, frequently almost procumbent subspecies is pri- marily and sharply set off from the typical population by the critical calyx distinction. It further occupies a clear-cut southern geographical range. Interestingly there is considerable similar- ity in its calyx characters to those of Gentiana Parryi of the south- ern Rocky Mountains, suggesting that this latter species might be likewise a southern, but more complete segregate of G. calycosa subsp. typica. Intermountain Herbarium, Utah State Agricultural College, Logan. May 11, 1941 ARNICA IN ALASKA AND YUKON Bassetrt MAGUIRE For some time the manuscript of a monograph of the genus Arnica has been completed, and is awaiting publication. During the course of the work the writer had been asked by Dr. Eric Hul- tén to contribute the account of Arnica for the “Flora of Alaska and Yukon,” now appearing in parts. Because of the immediacy of the needs of Dr. Hultén, and because appearance of the “mono- graph” does not, in the near future, seem probable, the following new entities, new names, and new combinations, for the most part, are herewith extracted from that study. Only the principal synonomy is given for the new names and combinations. Se- quences and numbering of entities is that employed in the treat- ment prepared for Hultén’s “Flora.” la. Arnica aupina (L.) Olin subsp. angustifolia (Vahl) comb. nov. A. angustifolia Vahl, Fl. Dan. 3. 1816; A. alpina (L.) Olin var. angustifolia Fernald, Rhodora 36:96. 1984. Ib. Arnica atpina (L.) Olin subsp. attenuata (Greene) comb. nov. A. attenuata Greene, Pittonia 4: 170. 1900. Ie. Arnica aLtpina (L.) Olin subsp. tomentosa (Macoun) comb. nov. A. tomentosa Macoun, Ottawa Nat. 13: 168. 1899; A. tomentosa Greene, Pittonia 4: 168. 1900; A. pulchella Fernald, Rhodora 27: 18. 1915. 2a. ARNICA LOUISEANA Farr subsp. frigida (Meyer) comb. nov. A. frigida Meyer ex Ilijin, Trav. Musc. Bot. Acad. Sc. U.S.S.R. 19: 112. 1926; A. nutans Rydb. N. Am. Fl. 34: 828. 1927; A. Sancti- Laurenti Rydb., N. Am. Fl. 84: 828. 1927. a. Var. genuina nom. nov. d. frigida Meyer ex Ilijin l.c. b. Var. Mendenhallii (Rydb.) comb. nov. A. Mendenhalli Rydb. N. Am. Fl. 34: 329. 1927. ce. Var. brevifolia (Rydb.) comb. nov. A. brevifolia Rydb. Nevam. Fl. 84: 329. 1927. d. Var. illiamnae (Rydb.) comb. nov. A. Illiamnae Rydb. Noam. Fl 34-331. 1927. 154 MADRONO [Vol. 6 e. Var. pilosa var. nov. Caulibus 5-10 (15) em. altis, foliis 2.0-4.0 (5.0) cm. longis, denticulatis; herbaceo exigue piloso; pedunculis dense pilosis; pericliniis densus subflavis lanato-pi- losis; capitulis nutantibus, aut plus frequenter erectis; acheniis glabratis, aut plus frequenter sparse super hirsutis. Type. Igloo Creek, McKinley National Park, July 11, 1932, | Joseph Dixon 29. Deposited in the Herbarium of the University of California; isotype at the United States National Herbarium. This population, seemingly confined to the Mount McKinley area, is strongly marked. Plants with erect heads closely re- semble specimens of A. alpina subsp. tomentosa, to which indeed they had mostly been referred. The basally or entirely glabrous achenes, and the yellowish periclinial pubescence seem, however, definitely to relate the McKinley plants to d. louiseana. 4a, ARNIcA corpiFoLIA Hook. subsp. genuina nom. nov. A. cor- difolia Hook., Fl. Bor. Am. 1:33. 18384. a. Var. pumila (Rydb.) comb. nov. A. pumila Rydb. Mem. N. Y. Bot. Gard. 1: 433. 1900. 5a. Arnica Cuamissonis Less. subsp. genuina nom. nov. A. Chamissonis Less., Linnaea 6: 238. 1881. a. Var. Typica Regel, Suppl. Ind. Sachal 151. 1864. b. Var. interior var. nov. Foliis inferioribus plus petiolatis, 2.5-3.5 cm. latis; capitulis 15-18 mm. latis; corollis disci 7-8 (10) mm. longis. Type. Palliser, July 30, 1906, S. Brown 770. Type deposited in the Gray Herbarium; isotypes at United States National Her- barium, New York Botanical Garden, Academy of Natural Sci- ences, Philadelphia, Missouri Botanical Garden. The variety interior is the common inland form, and by far the greatest population, extending southward into Canada. It differs from, although completely intergrading with, the maritime var. typica Regel in having mostly petiolate