Ngee: : Has pas Gr ry pee > = a & +. i ad. 1 = iAP, PL, be MAE ee Bua) phew Vibe i Dts as PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES THIRD SERIES DEN OOo Vows ff 1900-1904 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1904 CONTENTS OF VOLUME II. PLates I-XXXII. PAGE AMMEN dias 9s ue tates onic mw aeis Coin Oe EM Rak Rae Bde R a ST AON RY MA weds i RCOMECNIES Soro sche eee ADS alee Bl iieke ETAL IEN A ieiar avian paraeals sens eieine baat s iii No. 1. Nitophylla of California. Description and distribution. By Chas. Palmer, Nott., 1) (Riatesil— ai ciae scutes ako areca oaeities: I (Published August 3, 1900.) No. 2. The Development of the Karyokinetic Spindle in the Pollen- Mother - Cells of Lavatera. By Edith Sumner Byxbee. (Plates mx PT) SC aia s toe ictete bebe teemerie cesses sieve oa os 63 (Published October 31, 1900.) No. 3. Studies on the Coast Redwood, Sequoia sempervirens Endl. By George James Peirce. (Plate XIV )s.255.020.66...5-<: 83 (Published April 4, 1901.) No. 4. A Revision of the Genus Calochortus. By Carl Purdy. (Plates PR SINR ite oo. sccvecelblatarvolelsiawe vis |e 0:0 ais.tche eR eeeMEmEne Ts ee leet ache i 107 (Published December 14, 1901.) No.5. A Group of Western American Solanums. By S. B. Parish... 159 (Published October 23, 1901.) No. 6. An Account of the Species of Porphyra Found on the Pacific Coast of North America. By Henri T. A. Hus. (Plates bb, Eh, 6.0) | AR ee Pe ERS ab seer ge 173 (Published January 4, 1902.) No. 7. Some new Species of Pacific Coast Ribes. By Alice Eastwood. (Plates DEX EER IV ite ina cc as ac erie aomaem eee apy ema s 241 (Published April 14, 1902.) No. 8. Cell Studies. I. Spindle Formation in Agave. By W. J. V. Osterhout, (Plates SO V-XAVILI) cs ce saemanne cores 255 (Published May 5, 1902.) No. 9. New Species from the Sierra Nevada Mountains of California. By Pailice Bast wore icies'e tvs oirin's ia = pila orinlure’ whaeialen’e avy 285 (Published June 3, 1902.) No. 10. The Root-tubercles of Bur Clover (Medicago denticulata Willd.) and of some other Leguminous Plants. By George Jaiies: Peirces. (Plate SORL A) ca. wove s¥/s) iia cle thoiarnie ueniaie 295 (Published June 21, 1902.) No. 11. Spindle Formation in the Pollen-Mother-Cells of Cassia to- mentosa L. By Henri T. A. Hus. (Plates XXX-XXXII) 329 (Published April 30, 1904.) SAGA Se rere are Serre ieee Aer aee tense Rest py ite eomor at fe bis are lo Mratarelarduedete Pen stats oeele 355 April 30, 1904. LIBRARY NEW YORK Bol ANICAL GARDEN i. SCIENCES ay), uf Ca . Preps I % ry Vor. IL, No. a a! e ae: Mt abst oy t ti pe 4 i fe d hi B; ARLES PauMEeR Nott — RM wie Me oi i Issued Ne es A « f a oy / i moe ry , ae fy Ae ra Gay. ky ee ee pees & cra | ‘EDI Lu PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Tuirp SERIES BoTany | Vata tly Ne ur Nitophylla of California. Description and Distribution g@RA Qk Ry 2 WHEW YORK BOTAKHICAL £ ; ~ i7 gt & bd BY CHARLES PALMER NotT Wiru NINE PLATES Issued August 3, 1900 * SAN FRANCISCO PUBLISHED BY THE ACADEMY 1900 NITOPHYLLA OF CALIFORNIA. DESCRIPTION AND DISTRIBUTION.' BY CHARLES PALMER NOTT. CONTENTS. PvLates I-IX. PAGE Foe PEE ROIIUC TIONS: , 0 /silu< a ctohieinepemivenes asics y nacies Qabyiaiwiele a's ae 2 Bie AERIS IVEY 5 2 a ain ttrniort nisi M vit Laie Mailaalamions ols intdeies ie Mania meal os oi 2 PTE) AGEmeRAL, OESCRIP BION: 2529s oh 255 gee id 6 Dea eens Seed id aed aie 5 SYNOPSIS OF GENERIC CHARACTERS),......00) leo os ck ele bein 5 REMARKS ON SOME GENERIC CHARACTERS AS ILLUSTRATED BY. CALIFORNIAN SPRCIES (2% fats 32 244 solbaiiaae cane ae +s 7 BRE EFOSC AIC FIONA Sco stakes Cte Sohn Heeb ale atlas cs a LRULROE OF, « SUBSIFOLUNGAS 54 1c tin wictewind oe kin slew en tee as 7 ROCUTON OF THIZGlaS 0. 2 vernarawia ORG Sia ee ola 8 ORSRCOLS GND. INNOVATIONS 5)! os san alana ee oa eee ae 9 ME TEE EEE ECON 6 oa. 5's eB ais Sa ook mee ae OE a 9 Te ET NE OTT EEE LY MAORI NS oe en een ME 9 SHALE RL BIGMENTNG Sou ss che sins Varennes pn a Pic sae 9 Form and Branching as Specific Characters........... so) Boy 17) oa Ae Ae EASE aleve mieg 2S ape eres CREE Oe ee Io DUT EIEI ED a EAE att Oi dlse eas abc et i: b-4hd oo Ske Sta Tea eae ee Io VETOES Ti Seth tla aie Ga) Oa amie! s bm Deen Lo eRe ed Se II VETIS I etic any eR hs in 0) f Sine aS ce aa A eee oe II matte amd Paved CRONGCLSS: sor also deat da ae II DP CAUEESOE NE ee at ad «(ty n'aigins)si at date At ee eee eR TE tat ee 12 RTP Divaid Vite areas ok ne Ine Pa ape ieiatte's) acs roar Sea Re 12 UF ORF CCETIO I eh i sin 2) Wi gta dS u's 2 Ea Saige a Ae es ig) ALE CEIIE ENO CYSOGATDS: oo dce eee a 14 POE STEGER A gina ei. Syoid'>: asa a Se ee 14 Geographical Distribution of the Genus ............... 14 TT OIETIEG I 6 op NE Ghat tS chee su « Sete, Grek AR 15 IV. DESCRIPTION AND DISTRIBUTION OF SPECIES.............2 .00008 15 KrY TO THE CALIFORNIAN SPECIES OF NITOPHYLLUM...... 15 DV PIL CP MINOT OMEN aE ie ob da kh recaps the 2 ese ek 32 ON LAP NEI cratic sensed e's Semis eo een fas Pelee eae: 29 Die PIE LE SULAIOIE SAS Son 5 BE ga AS ch ons SMa Gd le ti ee aaa eR 27 Be COMMBCME Nei 0 xi Satins dara\ np Ret Chea obre stan GinwE PRN Cate 39 DN CORPO MII ATA 0 hs \(5 nisi n's wena tic aed’ alin eRe tia nL tt 34 DUERLDELENOUMUIE occa ha nse tas ald es Larne eRe tie ees 16 AMERY CUI EMM ete UE aca | Batatss Su lnnais Gis hoe: Ra dekh SOR 6 « 22 ANA MURROT AIH EME DS rex gate) hs PEP Sade Gan Nie steed hat ics 26 DN EOF DPI hos diy wis ean. too hake id Usted oa aa ane 24 Ds MELLEL gaa te tae tock s AIG (cian aes ook SITE ates 21 TAPES OF DISTRIBUTION OF SPRCIBS! 6). 5 4201.0 eee das 43 LITERATURE AND EXSICCAT# CITED IN THIS PAPER...... 2.2... 000008 45 PRA Rian UA Riese Sere. hic SE mice wd sich ge gd Wl wis wht sbpu olip wikis w 46 1 Contributions from the Botanical Laboratories of the University of California, No. ro. {1] July 24, 1900, 2 CALIFORNIA ACADEMY OF SCIENCES, [PRoc. 3D SER. I. INTRODUCTION. Our knowledge of the genus /Vtophyllum Grev. as rep- resented upon the coast of California and adjacent shores has been hitherto of a fragmentary and limited nature. The number of species found in the region named, the points of their occurrence, and their identity with the forms already known, or their title to recognition as distinct spe- cies, have been for some time largely matters of conjecture. With a view of providing a more connected account of these species, so far as data could be obtained, the follow- ing descriptions and accompanying notes upon distribution are presented. II. History. The first notice of Californian /V:tophylla, so far as the writer is aware, was made by W. H. Harvey (1858, Pt. II, p. 104, Supp., p. 128), who mentions the species of /Vzto- phyllum then known upon the western coast of North America. Two forms, JV. fryeanum and JV. laceratum (LV. violaceum )', were mentioned as occurring at Golden Gate, San Francisco Bay, California. Still later, in a ‘‘ Notice of a Collection of Algz made on the Northwest Coast of North America, chiefly at Vancouver’s Island, by David Lyall, in the years 1859-61.’’ Harvey (1862, p. 170) gives the original description of MHymenena latissima (LV. latissimum), specimens of which were dredged, or found adrift in Esquimault Harbor, Vancouver’s Island, B. C. The next mention of Californian /Vtophylla is found in the ‘‘Bidrag till Florideernes Systematik” of J. G. Agardh (1871, p. 49), where reference is made to Harvey’s species Hymenena latissima Harv. (LV. latissimum ). A noteworthy addition to the number of North American species was made when Professor W. G. Farlow (1875, p- 365) published a ‘‘List of the Marine Algz of the United States, with Notes of New and Imperfectly Known Species.”’ 1 The names in parentheses are those applied to the species as recognized in this paper, where the quoted name differs. Bot.—Vot. II.] MWOTT—CALIFORNIAN NITOPHYLLA. 3 In this paper were enumerated the following six species: NV. (Neuroglossum) anderson J. Ag. (IV. andersonia- num); NV. fissum J. Ag. (WV. ruprechtianum); NV. fryeanum Harv.; WV. laceratum Grev. (LV. violaceum); LV. latissimum J. Ag.; WV. ruprechtianum J. Ag. In the following year Farlow (1876, p. 695) published a second list, which included all the above mentioned species with the addition of JV. areolatum D. C. Eaton (JV. Jatis- simum) and JV. spectabile D. C. Eaton. The most comprehensive account of /Vitophyl/um yet pre- sented was that contained in the Epicrisis Floridearum of J. G. Agardh (1876, pp. 446-472, 698-7or). This account of the genus included all of the species then known to occur on the west coast of North America, with the excep- tion of WV. spectabile D. C. Eaton. To the forms men- tioned by Harvey and Farlow were now added JV. mudte- lobum J. Ag., WV. violaceum J. Ag., and WV. flabelligerum J. Ag. (LV. ruprechizanum). Farlow (1877, pp. 238, 245), in a paper discussing some aloz new to the United States, comments on some of the species mentioned by him in the two papers of 1875-76, and also gives the original description of JV. spectadbzle D. C. Eaton. Between 1877 and 1898, the literature pertaining to Cali- fornian WVitophylla consists of lists of the forms occurring on the coast, with the exception of Hervey’s ‘‘Sea Mosses”’ (1881), a popular work, in which some of the character- istics of color, size and venation of the JVtophylla are described. Several other writers should here be noticed. ‘Dr. C. L. Anderson (1891, p. 224), M. A. Howe (1893, p- 67), Daniel Cleveland, and A. J. McClatchie (1897, p. 358) have published lists of coast forms. No additions were made to the number of west coast forms by any of these writers except the last named, whose paper contains the first mention of the occurrence of JV. uncinatum J. Ag. upon the Californian coast. The latest and most complete statement in regard to west coast Witophylla is found in the volume published by. J. G. Agardh (1898). 4 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. According to Agardh twelve species of JVztophyllum and two species of /Veuroglossum are assigned to the west coast of North America. The complete list is as follows, viz.: LV. farlowianum J. Ag. (WV. ruprechtianum); N. flabelli- gerum J. Ag. (lV. ruprechtianum); NV. fryeanum Uarv. (LV. fryeanum); LV. latissimum J. Ag.; WV. macroglossum J. Ag. (LV. latissimum); NV. multilobum J. Ag.; WV. marginatum J. Ag. (LW. ruprechtianum); N. ruprechtianum J. Ag.; LV. spectabile Eaton.; JV. stenoglossum J. Ag. (LV. violaceum); NV. uncinatum J. Ag.; WV. violaceum J. Ag.; Neuroglossum andersontanum J. Ag. (Wit. andersonianum); MWVeurogtos- sum lobuliferum J. Ag. (Vit. violaceum ?). Agardh, basing his distinction between species upon dif- ferences in color, texture, form of frond, and position of cystocarps, as well as upon the more reliable characters, such as shape and position of sori, and venation of frond, regards all these forms established by him as valid species. It should be observed, however, in regard to these forms, that the descriptions are in many cases, as admitted by the author himself, drawn up from fragmentary or imperfect specimens. Further, the characteristics which are em- ployed to a considerable extent by Agardh as distinctions between species, and even between subsections, are vari- able to a marked degree. Observation of a considerable range of forms by the writer has led to the conclusion that sufficient allowance has not been made for the variations which require that great freedom should be used in defining the boundaries of species occurring on the Californian coast. A distinction should be made between the more variable characters such as color, texture, form of frond, and posi- tion of the cystocarps, and the less variable characters, such as shape and position of sori, and venation of frond. After a careful examination of the descriptions of the species as formulated by Agardh, and further study and comparison of the variations of the plants themselves, the writer is obliged to conclude that the degree of elasticity which seems desirable in considering Californian forms has not been permitted in regulating the limits of a species, or Bot.—VoL. II.] MOTT—CALIFORNIAN NITOPHYLLA. 5 else access has been had to material which has not come under the writer’s observation. A further consideration of these forms will be found in the remarks upon species in a later portion of this paper. The writings of the three authors mentioned above, viz., Harvey, Agardh, and Farlow, constitute the important liter- ature upon Californian J/Vztophylla, and their work alone will be considered in the further discussion of the species. III. GENERAL DESCRIPTION. The fuller discussion of the species of /Vtophyllum of the Californian coast will be advanced by some special treat- ment of the prominent morphological characters and geo- graphical distribution of the genus itself, for the sake of the increased light thrown by such treatment upon like points in connection with the forms to be hereafter discussed. The following synopsis exhibits the principal characters of this genus and will be followed by a discussion of some special features of morphology and distribution, as illus- trated by Californian species. SyNopPsIS OF GENERIC CHARACTERS. Frond either erect or exhibiting a prostrate and an erect portion. Prostrate frond creeping, linear, or irregularly expanded or membranous, occasionally with midrib, nerves, or veins; with or without rhizoids; variously lobed, divided or branched, occasionally proliferating; branches rising at intervals into erect fronds; margin entire, serrate, dentate, crenate, undulate, or lobed; sometimes forming offshoots and innovations. Erect fronds rising from holdfast or prostrate frond, singly or several together; sessile, subsessile, or stalked; flat and linear, or membranous, variously lobed, divided, forked, segmented, and branched; frequently proliferating ; 6 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. - with or without midrib, nerves or veins; with margin entire, serrate, dentate, crenate, undulate, orlobed. Branch- ing subdichotomous, subpinnate, subpalmate, or palmate, with branches or segments entire, linear or expanded, sometimes much prolonged. Stalk linear, flat, with or without distinct midrib; frequently becoming thickened and cylindrical through wearing away of margin of frond and renewed growth of remaining portion; frequently twisted by wave action; often persistent and freely proliferating. Midrib, when present, usually conspicuous, narrow or wide, simple below, sometimes branched above, sometimes evanescent or dividing into flabellate or anastomosing nerves, frequently becoming thickened, stout and persist- ent, freely proliferating. Nerves usually conspicuous, occu- pying body of frond, margin, or apices, usually branching freely, flabellate, free, or anastomosing, sometimes dividing into minute and inconspicuous veins. Veins inconspicuous or microscopic, occupying body of frond, margin or apices, simple or branching, flabellate, free, or anastomosing, com- monly evanescent and indistinguishable from ordinary tis- sue of frond. Sporangia found on both surfaces of the frond, usually in locally thickened portions, in sori of varied shape, con- taining tripartite tetraspores. Sori minute and scattered over the entire surface of the thallus, or large, forming orbicular patches disposed irregularly over the surface; or linear marginal patches; or lines arranged radially along the margin; or borne on marginal or surface proliferations of varying size. Antheridia developed from the superficial cells of the thallus, forming whitish patches scattered over the surface of the frond, the latter frequently becoming rugose. Cystocarps scattered over both surfaces of the frond, or arranged along the margin, or borne on marginal prolifera- tions, usually large, projecting beyond the surface of the thallus, opening by a carpostome. Bor.—Vot. Il.] MO7T7—CALIFORNIAN NITOPHYLLA. | REMARKS oN SoME GENERIC CHARACTERS, AS ILLUS- TRATED BY CALIFORNIAN SPECIES. The Prostrate Frond.—The prostrate creeping frond possessed by many species of Nitophyllum deserves special consideration. Agardh makes some use of this character in separating the genus into subsections, but the importance of this structure to the plant and the extent to which it may be developed have not, so far as it has been possible to learn, been very fully demonstrated. Influence of Substratum.—The character of the sub- stratum upon which the plant is located and the extent to which it is exposed to the dashing, drawing, or swirling force of the waves affect both the amount of growth and the shape of the prostrate frond. The various species of Vtophy//um occur in a variety of situations, from about mid-tide mark outward and down- ward to deep water. They grow in some cases upon the piles of wharves, where the prostrate frond must take advan- tage of cracks in the wood or roughenings of the surface to secure a foothold. Other species are found upon bare rock- surfaces, exposed to the dash of breakers. The most com- mon situation is that of those species which inhabit sheltered rock crevices or pools surrounded by rocks which protect them from the force of the waves. In such spots there will usually be found upon the rocks a rich growth of Bryozoa and Porifera, whose sponge-like substance affords an excel- lent foothold for the plants and is conducive to a free devel- opment of the prostrate frond. Other algz, notably the Corallines, afford by reason of their jointed structure and rough surface, excellent habitats. In general, the charac- ters of the prostrate frond correspond to those of the erect portion, but there are marked exceptions. JV. latissemum, for instance, possesses a membranous, broadly divided, erect frond, while that of JV. andersontanum is very much branched. Two forms in the same genus could hardly seem more widely different, yet the prostrate frond of the two species is very much alike where growing under 8 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. approximately similar conditions of substratum and wave action. Under other conditions the prostrate frond of JV. latissimum may become widely linear and irregularly lobed, lose the toothed margin which forms one of the points of resemblance to JV. andersonianum, and acquire a midrib and nerves, in this case resembling the prostrate frond of JV. ruprechtianum. Again, VV. multilobum, which as a rule selects sharply inclined or vertical rock-surfaces as a habi- tat, forms by means of its prostrate frond orbicular patches composed of the closely overlapping, broadly lobed, and membranous portions of the creeping, prostrate frond. The room afforded on the bare rock apparently favors the radial development here exhibited; while the necessity for secur- ing adequate thickness and firmness to meet the dash of the waves has led to the close overlapping or dovetailing of the various divisions of the prostrate frond. ‘ It has already been remarked that the prostrate part of the plant resembles in many respects the erect portion which rises from it. This statement usually holds good with regard to the shape and branching, but does not apply to the venation. As a rule, a midrib or nerves are lacking in the prostrate frond of those species whose erect fronds are provided with such structures. Formation of Rhizoids.—A number of species also exhibit a response apparently to the stimulus of contact, by send- ing short processes or rhizoids from the surface of the frond to the substratum to which they adhere. These pro- cesses have been observed on the under surface of the prostrate frond of JV. ruprechtianum, N. violaceum, IV. multilobum, NV. harveyanum and JV. corallinarum, and are recorded for several other species. Still more remarkable is the instance observed of the formation of these rhizoids in the case of a plant of JV. violacewm. This specimen had wrapped itself around portions of the thick frond of Prio- nitis lanceolata, to whose surface numerous processes sent forth from the surface of the Vztophyllum in contact with the Prionztis had attached themselves. Bot.—Vou. II.] MOZT—CALIFORNIAN NITOPAYLLA. 9 Offshoots and [nnovations.—A point of further interest in connection with the development of the prostrate frond is found in the formation of offshoots and innovations. In the first case, slender branches may arise from the margin of the older portion of the frond. These grow, secure attach- ment for themselves, and separate from the parent frond, and later give rise in turn to erect fronds. In the second case, by the growth and branching of the prostrate frond, an extended structure is produced, the ramifications of which become separated from each other by the decay and disappearance of the older portions, thus forming innova- tions in a manner similar to the process occurring in the Bryophytes. The Erect Frond.—The erect part of the plant commonly rises singly from the prostrate portion, but occasionally the fronds are clustered together, as in /V. multclobum, where they are grouped in the middle of the orbicular patch formed by the prostrate frond. Szze.—The height and breadth of the erect frond is an extremely variable character. JV. corallinarum, occurring upon Corallina chilensts, does not reach a height of 2 cm.; while JV. sfectabile is reported by D. C. Eaton as reaching 50—60 cm. Shape and Branching.—Great diversity exists in the shape and branching of the erect frond. Some species are broadly membranous, and but slightly lobed or divided. Good examples of this type are JV. spectabile, VV. latissimum and JV. fryeanum. At the other extreme may be placed such a finely dissected and abundantly branched species as lV. andersonianum. Between these two opposing types are found all gradations of frond division and arrangement of branches. It is not uncommon to find in one and the same species forms exhibiting a tendency to become broadly membranous, or very much divided and branched. For instance, in /V. ruprechtianum, there seems to be a ten- dency toward the flabellate or expanded type of frond, though the typical specimens of the species are character- ized by division of the frond into linear, much prolonged IO CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. branches. JV. violaceum, on the other hand, varies in the direction of the finely dissected type of frond. JV. fryeanum is a membranous form which often becomes narrowly lacin- iate, with the segments much prolonged. JV. andersonz- anum, which has been instanced as an example of a finely dissected frond, becomes, when growing in quiet water, very broadly linear, with the amount of branching largely reduced, and the expanded branches-very regularly arranged. Form and Branching as Specific Characters.—Some allu- sion has been made in the foregoing pages to the value of such characters as form and branching of the erect frond for distinctive purposes in describing a species. The ques- tion was raised then as to the advisability of making too narrow limits for a species upon such distinctions as form and branching, without regard to the effect of environ- mental factors. The remarkable variations in these two respects existing within the limits of a single species empha- size this fact, and further call attention to the influence of environmental relations upon form. Stalk.—The stalk, which characterizes many species, varies in length, width and thickness. The plant, by vari- ations in the length of the stalk, may be sessile, subsessile, or long-stalked. In some forms the stalk is narrow and somewhat thickened; in others it is furnished with a thin, expanded margin. It may vary in thickness from a few cell-layers to a thick, almost fleshy tissue. A midrib, when present, usually runs through the median portion of the stalk. Increase in thickness may take place through a growth of the superficial cells, or through an increase in the number of cells composing the central layer. Frequently the thin margin becomes worn away, and this is accompanied by an increase in thickness of the median portion of the frond, so that the stalk becomes cylindrical. Midrib.—The midrib presents considerable variation in form and extent. In some species it may be distinguished only as a slight thickening of the median portion of the stalk or frond, while in others it is more highly differenti- ated, appearing as a ridge of considerable prominence. It Bot.—Vot. Il.] MOTT—CALIFORNIAN NITOPHYLLA. II may be either simple and unbranched, as is commonly the case in WV. multilobum, or it may become considerably divided, as in WV. ruprechtianum. Frequently it does not extend in the frond beyond the upper portion of the stalk or lower segments. In other cases its ramifications reach out into the branches almost to their tip, and there evan- esce, or divide into nerves. Nerves.—These structures are a characteristic feature of the frond in several species, reaching a high degree of development in some forms. In NV. latissimum no midrib is present, but the large membranous frond is supported by a network of intersecting nerves and veins of considerable prominence. The other species are not distinguished by such a full development of these structures. Usually the nerves are limited to the outer margins or apices of the frond, where they become flabellate or anastomose freely with each other. Veins.—More or less conspicuous veins constitute a note- worthy structural element in some species, especially in JV. ruprechtianum and LV. violaceum. In these plants, particu- larly where the frond is at all flabellate, a rich development of the finer venation may be seen, whose ramifications extend in a flabellate fashion throughout the frond, or, in some cases, anastomose with one another, finally becoming free. In WV. fryeanum, N. uncinatum, and LV. corallinarum the midrib and conspicuous veins are entirely wanting and the only trace of venation is seen in the microscopic veins which characterize these species. These minute structures are frequently very delicate and invisible to the naked eye. They extend as a rule throughout the frond, branching freely or anastomosing. In /V. fryeanum, however, the del- icate veins, in nearly every case, become somewhat stouter toward the base of the erect frond, where they form a more or less conspicuous fan-shaped area. A single species, /V. spectabile, is destitute of any sort of veins. Variable and Fixed Characters.—It has been said in a preceding portion of this paper that stress should be laid 12 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. upon the greater value of certain characters as specific distinctions and the less value of others. The more variable characters there mentioned have been sufficiently discussed, and the less variable, more important specific distinctions, based upon the character of the venation and shape and position of the sori, may here be treated. Venation.—The Californian /Vztophylla may be separated into groups, distinguished from one another by the charac- ter of the venation. In that group which includes the greater number of forms, the species possess a midrib, nerves, and conspicuous veins, developed to a’ greater or less extent according to the species. Some forms exhibit only the midrib, which passes over at once into the undiffer- entiated frond as in /V. andersonianum. Others, again, are provided with a midrib which divides into more or less con- spicuous nerves, the latter passing again into the ordinary frond, e. g., WV. harveyanum and LV. multilobum. Still other forms, such as JV. violaceum and JV. ruprechtianum, show a full development of midrib, nerves, and veins, the last named structures usually conspicuous, and either anasto- * mosing or remaining free and flabellate. A second group consists of forms in which no midrib is present, but the sur- face of the frond is marked by a network of reticulate nerves and veins, as in JV. /atissimum. 'The forms of the third group, including JV. fryeanum, NV. uncinatum and LV. corallinarum, are destitute of midrib and nerves, and are provided with scarcely perceptible, usually microscopic veins, which either anastomose with one another, or remain free. A fourth group comprises forms which wholly lack venation of any kind whatsoever, as e. g., /V. spectadbile. Sort.—The sporangia of Vitophyllum are gathered together into sori of varying shape and size. These latter structures may be employed as reliable specific distinctions in discriminating between species. The Californian plants, taken as a whole, show a considerable range of forms as regards the shape of the sorus, and likewise a considerable variety in its position. Within the limits of certain species, e. g., LV. violaceum and lV. ruprechtianum, these variations Bor.—Vor. II.]} MWO7T7T—CALIFORNIAN NITOPHYLLA. 13 are remarkable. Notwithstanding this fact, the variations can be so expressed in terms of size, shape and position that clear distinctions can be drawn between species. In some cases the sporangia occur in narrow lines of varying width arranged in a flabellate fashion on the upper divisions of the frond, extending perhaps from its median portion to the margin—their regular position in /V. harveyanum and a disposition frequently seen in JV. ruprechtianum—perhaps only found as a fringe just within the margin, as often observed in JV. ruprechttanum. Other forms have the sori in linear strips or patches along the margin of the frond, an arrangement best seen in JV. vzolacewm. In another spe- cies, /V. multzlobum, the sori are irregular in shape, rounded or linear-elliptical, with their longer dimensions extending transversely across the frond upon whose upper segments they are borne. The sori occur in other instances as rounded or elliptical patches borne either singly upon distal lobes, as in /V. andersonianum, or scattered irregularly over the surface, as in JV. spectabcle, sometimes displaying a ten- dency to arrange themselves into lines, as is well shown by LV. fryeanum. In yet other plants the sori are minute, rounded structures, densely aggregated between the anas- tomosing nerves and veins of the frond, e. o., JV. latissimum. Proliferation.—The capacity for proliferation possessed by some species is very great. To such an extent does this phenomenon take place, that specimens are often found in which the primary frond has remained compara- tively undivided, and has produced from its margin numer- ous proliferations which exceed in size the primary frond itself. The favorite point for the production of prolifera- tions is along the margin of the frond, especially after this has been weathered by the waves. In such cases the original frond may be reduced to a narrow strip, which will produce proliferations surpassing in size the original portion of the plant. In some species, particularly in JV. ruprechit- anum, the frond becomes reduced to the midrib; this per- sists for a longer or shorter period, and proliferates very freely, so that by this process the plant practically becomes 14 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. perennial. Some species, again, notably JV. vzolaceum and LV. ruprechtianum, bear marginal proliferations in great numbers, upon which appear the sporangia and cystocarps. Antheridia and Cystocarps.—The antheridia and cysto- carps need be mentioned only in a cursory way, inasmuch as they take no prominent part in the identification of the species. The antheridia first make their appearance as whitish patches distributed rather evenly over the surface of the frond. As they mature, the frond becomes wrinkled or rugose to a marked degree. Antheridial plants have now been found in three species, viz., VV. fryeanum, JV. latissimum and JV. spectabile, in considerable abundance. The cystocarps are conspicuous structures, making their appearance irregularly over the surface. In some species the most weathered or reduced plants seem to be favored in the production of cystocarps; and in this case the latter sometimes occur in clusters of proliferations produced from the surface or worn margins of the frond. Parasites.—In only one instance have the plants of /Vito- phyllum been found to harbor parasites. As a rule, the species are quite free from epiphytes and parasites. Gonimophyllum buffhamz Batters has been found by the writer (1897, p. 81) upon JV. ruprechtianum. The plants are a pale pink color, contrasting strongly with the deep lake red of the host plant. The parasite seems to prefer the lower portions of the frond as its habitat, occurring on the /V7fo- phyllum near its base much more frequently than elsewhere. It grows in patches, extending in some cases an inch in breadth, while the individual fronds reach a height of ten millimeters. So far, Gonimophyllum has been found only upon the tetrasporic /V. ruprechtianum. Agardh (1898, p. 88) is of the opinion that Gonzmophyl- Jum is a monstrous form of JV. laceratum. He suggests further that JV. /aceratum occasionally may be hermaphro- ditic, contrary to the normal course of development, and that this monstrous form may be the structure concerned in the phenomenon. Geographical Distribution of the Genus.—The genus NVitophyllum is distributed universally throughout the oceans Bot.—Vou. II.] MOTT—CALIFORNIAN NITOPAYLLA. 15 of the globe. It has been found in the Arctic and Antarc- tic oceans, along the shores of the continents bordering upon the Atlantic and the Pacific, in the Mediterranean Sea and the Indian ocean, and in the waters of Australia. The coast of the British Islands is particularly rich in species, while some of the finest examples of the genus come from Australia. About seventy species are known, of which number ten occur on the west coast of North America. Of the ten species thus credited, eight are limited to California or neighboring shores. A ninth species, /V. uxcinatum, is one of the oldest known forms, and is more widely distributed, being found in the Adriatic Sea, on the coast of Australia, in California, on the shores of Europe, in the upper Atlan- tic and Mediterranean Sea, and in New Zealand waters. The tenth species, /V. harveyanum, is an inhabitant of New Zealand coasts. Flabitat.—The species of /Vztophyllum occur on other algz and on Phyllospadix; on bare rocks exposed to the . dash of the waves or protected from their violence; or upon rock surfaces which have become thickly coated with Cor- allines or various Porifera and Bryozoa; or upon the piles of wharves on the surface of the wood. The plants range from the littoral zone between tide marks to the sublittoral zone and have been dredged at a depth of 12-15 fathoms. IV. DESCRIPTION AND DISTRIBUTION OF SPECIES. KEy TO THE CALIFORNIAN SPECIES OF NITOPHYLLUM. Frond provided with midrib only, or with midrib and nerves, or midrib, nerves, and usually conspicuous veins. Frond with midrib only, margin of frond serrate or dentate, sori in rounded patches on distal lobes of frond. N. andersonianum (32). Frond with midrib and nerves, but without veins. Sori forming flabellate lines on upper segments of frond. N. harveyanum (29). Sori forming transverse patches on upper segments of frond. N. multilobum (27). 16 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. Frond with midrib, nerves and usually conspicuous veins. Color a pale violet through bright violet red to purple violet ; thin and papery in texture; sori forming narrow flabellate lines on upper divisions of frond, or linear patches along margin, or rounded patches borne on the crispate margin or on mar- ginal proliferations, or appearing in both lines and patches. NV. violaceum (39). Color a bright red to blackish red; firm and leathery in texture ; sori forming wide flabellate lines on upper divisions of frond, or rounded patches on marginal or surface proliferations, or appearing in both lines and patches... V. ruprechtianum (34). Frond without midrib, but provided with network of conspicuous nerves and WEISS: iahichs Mest ua veils} Sa yn oo oe oes Sen a ae euree es ae NV. latissimum (16). Frond without midrib, nerves, or conspicuous veins, but provided with microscopic veins. Frond ample, erect, membranous below, branching above; sori form- ing elliptical patches scattered over surface of frond. N. fryeanum (22). Frond erect, branching throughout, apices of ultimate branches re- curved or hookeGweniau. wee: ues cre tate NN. uncinatum (26). Frond minute, creeping, ultimate lobes becoming free; sori forming rounded patches on free lobes; on Corallina chilensis (and other. alant)k stties tans & wie is deals tae a S55 NN. corallinarum (24). Frond destitute of veins, membranous; sori forming elliptical patches scat- tered over Surface Of frond..)s..2 2.22 J2sc.5 s0eec N. spectabile (21). N. latissimum 7. Ag. Epicrisis Floridearum, Contin, Spec. Gen. et Ord. Alg., 1876, p. 464. Hymenena latissima Harvey, W. H., Proc. Linn. Soc., (Botany) Vol. VI, 1862, p. 170. N. latissimum AGARDH, J. G., Bidrag Florid. Sys., 1871, p. 49. FARLow, W. G., Proc. Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for 1875, p. 695, 1876. NV. areolatum Eaton, D. C., Farlow, 1. c., p. 695. N. latissimum AGARDH, J. G., Epicrisis Florid., Contin. Spec. Gen. et Ord. Alg., 1876, pp. 464 and 699. Fartow, W. G., Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 238. FARLow, W. G., ANDERSON, C. L., and Eaton, D. C., Algz Amer.-Bor. Exsiccateze, No. 68, 1878. HERVEY, A. B., Sea Mosses, 1881, p. 175. ANDERSON, C. L., Zoe, Vol. II, 1891, p. 224. Howe, M. A., Erythea, Vol. I, 1893, p. 68. McC LaTcuie, A. J., Proc. So. Cal. Acad! Seti; Volo jiise7, 93,1358. Nori, C. Vga Phyc. Bor.-Amer., Coins, F. S., HoLpen, I., and SETCHELL, W. A., Fasc. VII; No. 335, 1897. TILDEN, J. E., American Algz, Century III, No. 212, 1898. AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol. DI, Pi 3, 1898, p: 83. NN. macroglossum AGARDH, J. G., |. c., p. 84. Bot.—Vot. II.]} MOZTT—CALIFORNIAN NITOPHYLLA. 7 Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, slender, linear, much branched; without rhizoids, midrib, nerves, or veins; margin serrate or toothed. Erect frond subsessile or shortly stalked, flat, membranous; without mid- rib, but with numerous, conspicuous nerves and veins; palmately divided or segmented, frequently proliferating; with margin usually entire, occasionally minutely serrate. Segments linear or wedge-shaped, usually expanded and lobed at apices. Stalk, when present, usually short, occasionally longer, formed by the wearing away of frond on margins at base and subsequent thickening of nerves and remaining portion of frond. Nerves and veins numerous, conspicuous, extending over almost entire surface of frond, evan- escent in outer lobes, branching and anastomosing, frond thus conspicuously areolate. Proliferations usually numerous, especially upon more reduced portions of frond, membranous, segmented or lobed, nerved and veined. Sporangia in minute sori scattered over both surfaces of the frond between the nerves and veins, in the conspicuous areoles. Antheridia forming dull whitish patches over the entire surface, the latter becoming ridged, indented, or rugose. Cystocarps numerous, large, irregularly disposed over both surfaces, pro- . jecting beyond the surface. Remarks on the Species.—The color of JV. latissimum does not change to any extent when dried. In the living state the tint is a deep, sich, lake red, becoming a shade darker in the dried condition. Plants frequently occur which attain a length of 35-40 cm., with the frond divided into numerous segments from 3-5 cm. in width. The spe- cies is one of the handsomest and most luxuriant in habit of any of the forms inhabiting the coast. LV. latissimum is characterized by its broadly expanded fronds, divided into large, rounded lobes and segments, and by the extensive network of prominent nerves and veins covering the surface. This conspicuous network gives to the surface a markedly areolated appearance, which serves to distinguish the plants from all others of the genus as represented on Californian shores. The frond frequently proliferates along the margin, thus giving rise to numerous ovate or lanceolate proliferations, which possess a well marked venation and often produce sporangia and cysto- carps. Proliferations likewise appear freely from reduced portions of the frond. (2) July 26, 1900. 18 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. The creeping, much branched, claw-like, prostrate frond possessed by JV. Jatissémum is an excellent adaptation for the purpose of enabling the plant to retain its foothold upon the rough and seamy piles and rock surfaces which are its habitat. Very little variation has been observed in the prostrate frond; and it seems to be a well defined and spec- ially differentiated structure. Tetrasporic plants are of the most common occurrence. Sporangia are produced very abundantly over the entire surface, between the nerves and veins, thus adding to the distinctive areolate aspect of the plant. The sori are minute and crowded together, giving the impression of large sori completely filling the areoles. Antheridial specimens have been observed in but one locality. They reacha large size, being among the most magnificent examples of the species. In the early stages of development the antheridia form pale, whitish patches. Later, they become more evident, and when well developed, cause the entire frond to appear dull reddish white. The surface of the frond is then decidedly rugose. A comparison of JV. latissimum with a specimen of JV. hillie Grev., distributed by Le Jolis in Algues Marines de Cherbourg, No. 215 (in herbarium W. A. Setchell), shows a striking similarity between the two species in texture, venation, and character of lobes or branches. A series of specimens of JV. Az//i@ in the herbarium of Professor W. G. Farlow shows that this plant varies greatly as to prominence of veins. Careful comparisonof JV. /at/ssimum with strongly veined specimens of JV. Az/iie brings out a strong resem- blance between the two. With the material in hand, how- ever, it is not advisable to do more than point out the possi- bility that the two species may be identical. It is desirable to call attention at this point to the latest views of Agardh (1876, pp. 464 and 699, 1898, pp. 83-85) concerning JV. Jatisstmum. According to Agardh, JV. Jatissémum is a form which was first collected by David Lyall at Vancouver Island, B. C., and described by Harvey (1862, p. 170) under the name Bot.—Vot. II.] MOTT—CALIFORNIAN NITOPAYLLA. 19 flymenena latissima. A similar form was received some- what later by Agardh from Golden Gate, San Francisco Bay, collected by Berggren. Further than this, corre- sponding forms were issued in the Alg. Amer.-Bor. Exsicc. (1878, No. 68), collected by Dr. C. L. Anderson, at Santa Cruz. Again, according to Agardh, there is still another form, viz., JV. macroglossum, which is referred to Californian shores. This plant was collected originally by Berggren also at the Golden Gate, and at first was included by Agardh in LV. latessimum. Specimens collected by the writer (1897, Fasc. VII, No. 335) at the type locality are said by Agardh to be identical with the plant first referred by him to J. latissomum and now included under JV. macroglossum. The writer has now examined in the field and herbarium a wide range of plants comprehended under these two species of Agardh. Specimens, collected at numerous localities and at various times in the year along the coast from San Pedro northward to Puget Sound entrance, provide a fair amount of material as a basis for estimating Agardh’s spe- cies. An especially abundant series of forms was collected by the writer between January and May at Golden Gate, San Francisco Bay, the type locality for Agardh’s /V. mac- roglossum. ‘This collection included numerous specimens of both the species above referred ‘to as established by Agardh. A discriminating examination of Agardh’s descriptions shows, as far as the writer can determine, that JV. Jatzss¢mum differs from /V. macroglossum in being of a paler color, possibly less luxuriant in habit, with by no means such a well developed system of venation. The nerves and veins anastomose less freely, project above the surface of the frond less prominently, or not at all, as is more commonly the case, and soon evanesce into the frond. The areoles are larger and more conspicuous by reason of the broad, flat aspect of the nerves and veins. The sori are fewer in number and are most numerous about the borders of the areoles. 20 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. The range of forms examined by the writer with these points in mind exhibits, it is true, the structural differences pointed out by Agardh. But further, it should be observed that the species here in question, as field studies show, is apparently a form occurring between December and August, reaching its finest development in February and March, and at that time exhibiting the characters of /V. macroglossum. The forms collected from April to August show the charac- ters of lV. /atissimum, with frequent occurrence of transi- tion forms between the two species. The evidence collected by the writer in field and herba- rium points to the conclusion that /V. macroglossum and JV. latissimum are seasonal variations of the same species, that is, JV. latissimum. Habitat.—On piles of docks and wharves, or on sloping rock surfaces where the wave action is not violent; lower littoral to sublittoral zone. Distribution.—At various points along the coast from San Pedro to Vancouver. Localities. —San Pedro! (Mrs. S. P. Monks); Santa Barbara! (Dr. and Mrs. L. M. Dimmick; Mrs. S. P. Cooper); shores of San Luis Obispo County! (Mrs..R. W. Summers); Pacific Grove! (M. A. Howe; Mrs. J, My Weeks; C. P. Nott); Santa Cruz! (Dr. C. L. Anderson; Mrs. J. M. Weeks); San Francisco! (G. W. Lichten- thaler); San Francisco Bay entrance (Golden Gate) at Fort Point! (M. A. Howe; W. "A. Setchell; W.]74% Osterhout; C. P. Nott); at Lime Point! (C. P. Nott); Fort Ross! (W. A. Setchell); Klatsop, Oregon, (L. F. Hen- derson, in Herb.; W. G. Farlow, fide W. A. Setchell) ; Port Orchard, Washington! (J. E. Tilden); Esquimault Bay, Vancouver Island, B. C. (Harvey, in Proc. Linn. Soc., (Botany) Vol. VI, 1862, p. 170; Farlow, Proc. Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365); Puget Sound! (N. L. Gardner). Bot.—Vot. Il.] MWOTT7—CALIFORNIAN NITOPHYLLA. 21 Nitophyllum spectabile D. C. Eaton. In FARLow, Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 245. Nitophyllum spectabile FARLOw, W. G., Report U. S. Fish Comm. for 1875, p. 695, 1876; Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 238. Eaton, D. C., in Farlow, 1. c., p. 245. FARLow, W. G., ANDERSON, C. L., and Eaton, D. C., Algz Amer.-Bor. Exsiccatz., No. 67, 1878. HERVEY, A. B., Sea Mosses, 1881, p. 174. ANDERSON, C. L., Zoe, Vol. II, 1891, p. 224. AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol. DMs Pi3, S98, p..43- Synopsis.—Frond both prostrate and erect. Prostrate frond thin, linear, creeping, destitute of venation; becoming thickened when weathered; branch- ing irregularly; branches rising into erect fronds. . Erect frond sessile or subsessile, flat, membranous; destitute of venation; irregularly oblong, deeply pinnately lobed, occasionally palmately segmented, sometimes proliferating; margin entire, sinuate, or lobed. Segments linear, lanceolate, ovate, or cuneate, frequently deeply lobed at apices. Sporangia in elliptical sori, disposed at nearly regular intervals over both surfaces of the frond. Antheridia in whitish patches over entire surface of frond, giving to latter an areolate aspect. Cystocarps numerous, conspicu- ous, irregularly disposed over both surfaces, projecting beyond the surface. Remarks on the Species.—This plant retains, when dried, the bright, rosy red hue which characterizes.it in the living state. The species is said by Professor Eaton, who estab- lished it, to reach a length of 50-60 cm. It is one of the largest and finest species of the coast. The general aspect of the frond is much like that of /V. fryeanum. It differs from that form, however, in not possessing any kind of venation. k Comparison of JV. spectabile with LV. ruthenicum (P. & R.) Kjell. aroused a suspicion that the two forms might be iden- tical. A more careful examination showed that, in speci- mens of JV. ruthenicum received from Professor Kjellman, the plants were ‘‘obsoletely veined below,’’ as is stated in the description of /V. ruthenicum (1889, p. 25, Pl. I, figs. II-12), and as is shown in the figure. JV. spectabzle, on the contrary, is totally destitute of venation. All the evi- dence at hand demonstrates that /V. sfectabzle apparently is a distinct species. - The writer is indebted to Professor W. A. Setchell for the following note upon JV. spectadzle, through the courtesy of Professor W. G. Farlow, who kindly permitted an exam- ination of his specimens of /Vztophyl/a in connection with 22 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. the preparation of this paper. Professor Setchell says, ‘‘Specimens in herbarium of Professor Farlow (ex. herb. Acad. Petrop.) labelled ‘Aglaophyllum ruthenicum, Exp. Lutk. ad litora Americane borealis-occidentalis, Ross’ are young (about an inch high, with no fruit present), and might be the young plants of JV. spectabzle. In the herba- rium of Farlow are also several specimens from St. Paul Islands, in Behring Sea (legit White), which might be JV. spectabile.”’ The account of this species was advanced to a great extent by examination of a large collection of material from Mrs. J. M. Weeks, of Santa Cruz, Calif., who made an especial effort to secure antheridial, tetrasporic, and cysto- carpic plants. The material thus obtained confirms the conclusion that /V. spectabzle is entitled to rank as a distinct species, as established by Professor D. C. Eaton. Habitat. —On rocks ? or other algz, sublittoral to elittoral zone. Dredged in 12-15 fathoms, Monterey Bay, Calif. Distribution.—Along the coast from Santa Monica north- ward to Santa Cruz. Localities.—Santa Monica! (Miss S. P. Monks); Paci- fic Grove, in Monterey Bay! (C. P. Nott); Santa Cruz! (Dr. C. L. Anderson, fide Eaton in Farlow, Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 245; Mrs.-J. M. Weeks). Nitophyllum fryeanum /arlow. Algze Amer.-Bor. Exsiccatz, No. 69, 1878. Nitophyllum fryeanum Harvey, W. H., Ner. Bor.-Amer., Supp., 1858, p. 128? (See remarks on species). FARLOw, W. G., Proc. Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for 1875, p. 695, 1876. FARLow, W. G., ANDERSON, C. L., and Eaton, D. C., Algze Amer.-Bor. Exsiccatzee, No. 69, 1878. HERvey, A. B., Sea Mosses, 1881, p. 176. ANDERSON, C. L., Zoe, Vol. II, 1891, p. 224. Howe, M. A., Erythea, Vol. I, 1893, p. 68. McCratrcuig, A. J., Proc. So. Cal. Acad. Sci., Vol. I, 1897, p. 358. AGARDH, J. G., Contin. Spec. Gen. et Ords Ale: Volkan Pt 3, 1898.74: Synopsis.—Frond both prostrate and erect. Prostrate frond much reduced, flat, membranous, lobed, without venation or rhizoids. Erect frond sessile or subsessile, flat, membranous, with microscopic veins; »«-dichotomously or palmately branched or segmented; margin entire, Bot.—Vot. II.] MWO77—CALIFORNIAN NITOPHYLLA. 23 serrate or toothed. Segments linear, frequently prolonged, occasionally expanded and lobed at apices. Veins not numerous, extending through the frond, branching and anastomosing. Proliferations minute or wanting, appearing along the margin of the frond. Sporangia in small elliptical sori scattered over the entire frond, tending to become arranged into lines. Antheridia in whitish areolate patches scattered over surface. Cystocarps conspicuous, irregularly disposed over both sur- faces, projecting beyond the surface. Remarks on the Species.—The color of WV. fryeanwm is very attractive, being a bright rosy red in both the living and dried states. Some of the plants reach a height of 15 cm. The size and more especially the shape is subject to considerable variation. The frond may be short and deeply lobed, or long and branching, with the branches consider- ably prolonged and linear. N. fryeanum is noteworthy as being one of the three forms of the coast which possess delicate, microscopic veins. With the exception of this character, V. fryeanum and JV. spectabile have many points in common. The last named species, however, is destitute of any kind of venation. It is a matter of some doubt whether Harvey’s (1858, Supp., p- 128) original description of this plant does not better apply to V. multzlobum. His mention of a lobed and crenulate margin does not seem to hold good for JV. frye- anum. WHarvey’s name, however, was applied to the speci- mens issued in the Alg. Amer.-Bor. Exsiccate (1878, No. 69) and Agardh (1898, p. 74) also retains Harvey’s name, with an additional reference to the specimens above men- tioned. It seems advisable, therefore, to retain this name for the plant here dealt with, which is identical with that published in the Alg. Amer.-Bor. Exsiccate. There were reasons for supposing that VV. fryeanum might be referred to WV. ruthenicum (P. & R.) Kjellman (1889, p. 25, Pl. I, figs. 11-12), with which it agrees to a certain extent in the characters of venation and sori. Care- fully selected specimens were forwarded to Professor Kjell- man, who replied that the two species were not identical. The plants exchanged with Kjellman for purposes of 24 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. comparison, while not wholly inducing the writer to accept Kjellman’s conclusion as to the non-identity of the two spe- cies, yet do not furnish sufficient reason for declaring them identical. Until a more extended comparison can be made of a wide range of forms, the writer prefers to leave the species as established in the Alg. Amer.-Bor. Exsiccate (1878, No. 69). The writer further takes this opportunity to express his obligations to Mrs. J. M. Weeks for material of /V. fryeanum collected by her, which permitted a careful study of anther- idial, tetrasporic and cystocarpic plants. Habitat.—No reliable data are at hand concerning the habitat of this species. Some twenty-five specimens seen by the writer were all washed ashore from deep water. The plant presumably occurs upon rocks, and other alge, in the lower sublittoral, and perhaps elittoral zone. Distribution.—Known to occur with certainty at but two localities on the Californian coast. At Golden Gate, San Francisco Bay, it apparently has not been collected since the first specimen, if indeed it was identical, was secured by A. D. Frye and forwarded to Harvey. Localities.—Pebble Beach, Monterey Peninsula! (Miss Bayles); Santa Cruz! (Dr. C. L. Anderson; Mrs. J. M. Weeks); Golden Gate, San Francisco Bay ? (A. D. Frye, jide Harvey, Ner. Bor.-Amer., Supp., 1858, p. 128). Nitophyllum corallinarum, sp. nov. Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, flat, membranous, with microscopic veins and with rhizoids; lobed and branching, with branches becoming erect at intervals, margin entire. Erect frond subsessile, shortly stalked, flat, membranous, with microscopic veins; ovate-spatulate to elliptical, two to three times longer than broad; sub- dichotomously lobed or divided, margin entire. Segments minute, ovate, oblong or cuneate. Stalk very short, narrowly linear or cylindrical, passing into a midrib, the latter extending throughout frond, usually branching and free. Sporangia large, prominent, in sori of varying shape and size, solitary or clustered on body of frond or its segments. Femarks on the Species.—The form from which the syn- opsis of this species is drawn up was obtained at San Diego by Mrs. E. Snyder, and sent to the writer by Mr. F. S. Bot.—Vou. II.] MO7T—CALIFORNIAN NITOPHYLLA. 25 Collins. It is the only specimen which, to the writer’s knowledge, has thus far been seen on this coast. LV. corallinarum is avery minute plant in comparison with the other Vitophylia of the coast. Corallina chilensts, upon which the WVtophyllum grows, attains a height between 8 and 12 cm., and in the lower portion a width of 5 cm. or more. The branches of the Coralline are arranged pin- nately along the main axis from base to apex of the plant. The general outline is very regularly fan-shaped. The epi- phytic Wetophyllum extends, by means of its prostrate frond, over the entire surface of the Coralline, to which it adheres firmly by means of the rhizoidal processes that are produced abundantly from the surface in contact with the Coralline. From the prostrate frond rise at regular intervals erect branches which are shortly stalked and expand into ovate- spatulate or elliptical fronds, which may reach a height of 7 mm. and a width of 3 mm. The color is a rosy red to dull carmine. So completely is the Coralline enveloped by the Nitophyllum, that the natural color of the former, as well as its jointed structure, is very much obscured. The thin and delicate frond of JV. corallinarum presents the tessellated surface characteristic of the genus. Throughout both the prostrate and erect fronds may be distinguished minute microscopic veins, which branch more or less freely and remain free. The sporangia are prominent and large, considering the general minute size of the plant. They occur in clusters rather than in sori of definite shape, and are borne upon the body of the erect frond or onits segments. Tetraspores of unusually large size are formed in the sporangia. This plant may ultimately be found to be identical with some species of Europe. Such a determination must be left, however, to future observation upon more abundant material. Habitat.—Epiphytic on Corallina chilensis. Distribution.—Concerning the distribution of JV. coradl- narum little can be said. The Coralline upon which it grows is found along the coast from San Diego northward 26 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. to Fort Ross, but no observations are known to the writer on the occurrence of JV. corallinarum at any other place than the type locality. Locality.x—San Diego! (Mrs. E. Snyder in herb. F.S. Collins). Nitophyllum uncinatum 7. Ag. Spec. Gen. et Ord. Alg., Vol. II, Pt. 2, 1852, p. 654. Nitophyllum uncinatum McCvatcuiE, A. J., Proc. So. Cal. Acad. Sci., Vol. I, 1897, p. 358; also in Phyk. Bor.-Amer. CoL.ins, F.S., Holden, I., and SETCHELL, W. A., Fasc. VII, No. 337, 1897. AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol. III, Pt. 3, 1898, p. 65. Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, narrow, linear, thin and membranous; frequently weathered, reduced, and thickened along median portion, with microscopic veins, and with rhizoids; margin entire, or toothed; branching, branches rising into erect fronds at intervals. Erect frond sessile or subsessile, flat and linear, thin and delicate, some- times thickened in median portion, with microscopic veins; branching sub- dichotomously from the base upwards, with margin entire, or occasionally toothed. Branches linear, occasionally expanded, or acuminate, frequently recurved or hooked at apices. Veins microscopic, extending throughout frond, occasionally branching and anastomosing. ‘* Sporangia in solitary disc-like sori, on the upper branches ”’ or “‘in round- ed sori’’ on the outer branches. (See Remarks on Species.) Antheridia as yet unobserved. Cystocarps minute, marginal or submarginal, produced at infrequent intervals, projecting slightly beyond surface." Feemarks on the Species.—V. uncinatum has a bright, rosy red tint when alive, usually changing to a dull purplish or brownish red when dried. The fronds may attain a length of ro-15 cm. The plant is one of the more delicate species of the coast, as may be seen in the thin and mem- branous character of the frond. Throughout the narrow, linear segments extend microscopic veins, which, with the numerous recurved or hooked apices of the branches, may be regarded as the prominent morphological characters.” 1 Account of cystocarps from specimen in Hauck und Richter, Phykotheca Univer- salis. Fasc. VII, No. 306, 1889. 2 Nordhausen (Pringsheim’s Jahrbiicher f. Wiss. Botanik, Band XXXIV, Heft 2, 1899. p- 263) finds that the hooked apices of the branches of N. uacinatum serve as climbing organs. Bot.—VoL. II.] MOTT—CALIFORNIAN NITOPHYLLA., 27 No fruiting specimens of /V. uucinatum, so far as can be ascertained, have been reported from Californian shores. The plant seems to be an exclusively southern form on this coast, having been collected only at San Diego and San Pedro, in southern California. From these two localities numerous and abundant collections have been taken, none of which, however, have revealed fruiting specimens. There is a strong probability that the plant propagates itself largely, if not entirely, by vegetative means. It occurs commonly on Phyllospadix, in quiet water, conditions of substratum which would favor the active development of the prostrate frond. Its local abundance is shown by its occurrence in such quantities as sometimes to clog the nets of fishermen. The description of the sori, as given in the synopsis of the species, is taken from Agardh (1852, p. 654, 1876, p. 465, 1898, p. 65). The account of the cystocarp is based upon an examination of the specimen issued in the Phyko- theca Universalis (see note under synopsis of species). There seems to be little doubt that the species of this coast is identical with the European plant. Hlabitat.—In quiet water, on other algew, and on Phyllospadix. Distribution.—lV. uncinatum is a cosmopolitan species, limited in its local distribution, having been found at but two points on the coast. ’ Localities.—San Diego! (Herb., F. S. Collins); San Pedro! (Mrs. E. A. Lawrence; A. J. McClatchie; Mrs. S. C. Purdy; W. A. Setchell). Nitophyllum multilobum 7. Ag. Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 698. Nitophyllum multilobum AGaArpuH, J. G., Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 698. FARLow, W. G., Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 238; ANDERSON, C. L., Zoe, Vol. II, 1891, p. 224. CoLtins, F. S., HoLpEn, J., and SETCHELL, W. A., Phyc. Bor.-Amer., Fasc. VII, No. 336, 1897. AGARDH, J. G., Con- tin. Spec. Gen. et Ord. Alg., Vol. III, Pt. 3, 1898, p. 45. 28 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, irregularly expanded and membranous, sometimes a rounded expansion, sometimes irregularly branched or lobed; with rhizoids; without midrib, nerves or veins; margin entire, sinuous, or somewhat lobed. Erect frond slightly stalked, flat and linear, with distinct midrib; simple below, subdichotomously or subpalmately segmented, not proliferating; mar- gin sinuous, toothed or incised. Segments decidedly bullose, usually obtusely lobed, with sinuous or toothed margin. Stalk short, linear, flat, occasionally becoming thickened. Midrib well developed, rather wide, frequently branched, soon evanescent. __ Sporangia formed in large, irregularly oblong, frequently lobed or confluent sori, transversely placed upon the segments of the frond. Antheridia as yet unobserved. Cystocarps few, large, conspicuous, scattered over both surfaces. Remarks on the Species.—In the fresh state /V. mu/tz- Jobum is a dark red to dull carmine, becoming a burnt car- mine to blackish red when dried. The plant is a dwarf one, rarely reaching a height of 9 cm. The predominant characters of /V. multilobum are seen in the rather prominent development of the midrib, in the bullose aspect of the frond, and in the peculiar transverse sori. The first named structure is confined to the lower portion of the frond, where it is visible as a definite thick- ening of the median part, though it does not project promi- nently above the surface. At its upper extremity it fre- quently branches, and the resulting portions evanesce very soon into the ordinary tissue of the frond. The bullose frond of WV. multilobum is an important feature in the appearance of the tetrasporic plant. Usually the surface of the segments which form the upper portion of the plant exhibits this trait. Here the surface is alternately raised and depressed, while the margin becomes crinkled and lobed. The sori, together with the bullose aspect, furnish the most certain means of identifying the species. No other plant of the coast possesses such a characteristic feature as these transverse sori, usually produced in great abundance on the segments of the frond. | The characteristic transverse sori serve to distinguish JV. multilobum from lV. harveyanum, with which this plant other- wise has several points in common. In JV. harveyanum, however, the sori form flabellate lines on the segments of Bot.—Vot. II.] MOTT—CALIFORNIAN NITOPAYLLA. 29 the frond. In color, the two species are much the same. The midrib of JV. harveyanum is usually more pronounced than that of JV. multclobum. NV. harveyanum is much slen- derer than JV. mu/tzlobum, and may attain a height three or four times that of the latter. ffabitat.—On bare rock surfaces or on rocks coated with Corallines, from high water mark to the sublittoral zone. Distributcon.—Limited at the present time to the Cali- fornian coast. Has now been reported from Carmel Bay northward to Cape Mendocino. Apparently a northern form. Localities.—Carmel Bay! (C. P. Nott); Pacific Grove! (C. P. Nott); Santa Cruz! (Mrs. J. M. Weeks); Land’s End, San Francisco! (W. A. Setchell; C. P. Nott); Golden Gate, San Francisco Bay (Berggren, fide J. Agardh, Epi- crisis Floridearum, 1876, p. 698; W. A. Setchell); Lime Point, San Francisco Bay! (C. P. Nott); Dillon’s Beach (W. A. Setchell); Fort Ross! (C. P. Nott); Cape Men- docino (C. G. Pringle, in herb., W. G. Farlow, fide W. A. Setchell). Nitophyllum harveyanum 7. Ag. Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 462. Nitophyllum harveyanum J. Ac., Phyk. Bor.-Amer. CoLuins, F. S., Ho pen, I., and SETCHELL, W. A., Fasc. KIV, No. 693, 1900. Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, linear, flat; without rhizoids, destitute of midrib, and not proliferating; branching, branches becoming erect at intervals; margin entire, serrate, or somewhat laciniate. Erect frond stalked, flat, linear; with midrib and flabellate nerves; branch- ing, rarely proliferating, margin entire, or somewhat laciniate. Branches palmate or subpalmate, linear or becoming expanded, occasionally lobed or cleft. Stalk flat, linear, with distinct midrib, becoming thickened and cylin- drical through wearing away of margin and renewed growth of median por- tion. Midrib narrow, conspicuous, branching above, becoming divided into flabellate nerves, the latter conspicuous, branching freely, remaining free and flabellate. Veins minute or wanting. Sporangia in linear sori extending flabellately from the nerves to the mar- gin of the frond. Antheridia as yet unobserved. Cystocarps large, promi- nent, irregularly disposed, projecting beyond the surface. 30 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Remarks on the Species.—iV. harveyanum varies in color in the living state from deep salmon red to dull carmine, becoming purplish to blackish red when dried. The plant may reach a height of 20 cm., but is as a rule 6-10 cm. The prostrate frond in /V. harveyanum does not develop so extensively as in other species. It is destitute of midrib and nerves, and js not specially thickened. JV. harveyanum is a sparingly branched form as regards its erect frond and the whole plant is rather stiff and unyielding, even when freshly taken from the water. The midrib is conspicuous, tapering slightly towards its upper extremity, and rather suddenly becoming divided into flabellate nerves. The margin in the lower portions of the frond and upon the stalk frequently wears away, the remaining median portion then becoming thickened and cylindrical. In the branches the margin occasionally is serrate or laciniate. LV. harveyanum was first collected on this coast at Land’s End, San Francisco, by Professor W. A. Setchell. It grew in company with JV. mudtilobum, to which, at this locality, it bears some resemblance, on account of its size and vena- tion. Professor Setchell, however, upon noting the non- bullose character of the frond and the flabellate arrangement of the sori, so different from the transverse sori of JV. mz/- tilobum, concluded that the plant was a distinct species. The writer, when examining the plant in connection with other material secured by him at Fort Ross, was of the opinion that it must be the plant described by Agardh (1876, p- 699) under the name JV. flabelligerum, although pre- viously the conclusion had been reached that Agardh’s JV. flabelligerum was but a form of LV. ruprechtianum. Sufficient comment has been made already upon the dis- tinctions to be drawn between JV. harveyanum and JV. mul- tilobum. It is desirable, however, to point out here some of the differences existing between JV. harveyanum and cer- tain forms of JV. ruprechtianum. There is enough of simi- larity between certain variations of the latter species and LV. harveyanum to give reason for the suspicion that the two are identical. The examination of a good range of Bot.—Vot. II.] MWO7TT—CALIFORNIAN NITOPHYLLA. 3 specimens of /V. ruprechtianum soon brings to light the variation in that species, however, and helps to establish its non-identity with JV. harveyanum. Certain forms of WV. ruprechtianum exhibit a pronounced dark purplish red tinge, both in the fresh and dried states, and are somewhat stiff and brittle. The segments of the frond are narrower, and more or less prolonged. Such forms almost invariably bear flabellately arranged, linear sori, and on the whole, present the distinctive characters of NV. harveyanum. Between such an extreme variation as this and the typical WV. ruprechtianum, however, there may be found every gradation in color, form of segments, and position and shape of sori, which are discussed more in detail under WV. ruprechtianum. The form in question, however, usually retains enough of the distinctive color and venation of VV. ruprechtianum to enable it to be recognized. The writer is further indebted to Professor Setchell for a comparison made by him between specimens of JV. harvey- anum in the herbarium of Professor Farlow, and plants from this coast. The specimens in Professor Farlow’s herbarium are from New Zealand, and are designated as NV. harveyanum by Agardh. The resemblance in habit between these and plants collected by the writer at Fort Ross is very striking, both in the tetrasporic and cystocarpic plants. Judging from Agardh’s description and from this comparison of specimens, there seems to be good reason for keeping this species under lV. harveyanum. Habitat.—lNV. harveyanum is found most frequently upon very much exposed rock surfaces which are bare or coated with Corallines, at extreme low tide-mark in the littoral and sublittoral zones. Distribution.—Along the coast from Santa Cruz north- ward to Puget Sound. Apparently a northern form. Localities. —Santa Cruz! (Dr. C. L. Anderson); San Francisco! (G. W. Lichtenthaler); Land’s End, San Fran- cisco! (W. A. Setchell; C. P. Nott); Duxbury Reef! (W. A. Setchell); Fort Ross! (C. P. Nott); Puget Sound! (Thomas Stratton). 32 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER: Nitophyllum andersonianum 7. Ag. Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 474.1 Nitophyllum (Neuroglossum) andersonit FArRLow, W. G., Proc. Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for 1875, p. 696, 1876. Neuroglossum andersonianum AGARDH, J. G., Epicrisis Floridearum, Con- tin. Spec. Gen. et Ord. Alg., 1876, p. 474. Nitophyllum andersonit Hervey, A. B., Sea Mosses, 1881, p. 177. Nitophyllum (Neuroglossum) andersonti ANDERSON, C. L., Zoe, Vol. II 1891, p. 224. Nitophyllum andersonit Howe, M. A., Erythea, Vol. I, 1893, p. 68. Mc- CLATCHIE, A. J., Proc. So. Cal. Acad. Sci., Vol. I, 1897, p. 358. Neuroglossum andersonianum AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol. III, Pt. 3, 1898, p. 122. Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, slender, linear, without rhizoids; branching irregularly, occasionally prolifer- ating; without midrib, nerves, or veins. Margin beset at regular intervals with spine-like, sometimes recurved, pinnate teeth. Branches becoming erect at intervals, expanding into erect fronds. Erect frond shortly stalked, linear, flat, simple below, branching above, with midrib; margin serrate, dentate, or beset with numerous spine-like, pinnate teeth. Branches subpinnately arranged, linear, or alternate at base and expanding at their apices, usually much prolonged. Stalk linear, flat, with definite midrib and thin margin, the margin sometimes disappearing and the median portion becoming thickened, almost cylindrical. Midrib of vary- ing width, becoming prominent, in some cases thickened, almost cylindrical, branching and evanescent in upper portions of frond. Sporangia in rounded sori, the latter usually large and conspicuous, at the apices of the upper, sometimes expanded, branches. Antheridia and cysto- carps so far unobserved. Remarks on the Species—The color of JV. andersont- anum varies from bright red to dull carmine when alive, becoming a burnt carmine in the dried specimens. More often the plant has the darker hue mentioned above. The frond may attain a height of 20 cm. The - prostrate frond is commonly slender and much branched, showing much similarity to the corresponding portion of JV. /atissimum. It is destitute of midrib and nerves, and seldom becomes thickened or broadly linear. The erect frond branches freely, while its divisions exhibit considerable variation in width. In some plants they are very slender, linear, and much divided or branched. In 1 This plant was here for the first time described. It had, in 1875, been mentioned by Farlow (cf. citations) uuder the name Nrtophyllum (Neuroglossum andersonti) J. Ag. Ms, Bot.—Vot. II.] MO7TT—CALIFORNIAN NITOPHYLLA. 33 others the segments are quite broad and very regularly pin- nately arranged. The cause of this variation apparently may be found in the environment. When exposed to vio- lent wave action the fronds become extensively branched. In comparatively quiet waters, on the other hand, the ex- panded frond reaches its widest development. The pre- dominant characteristic of /V. andersonianum is the produc- tion, along the margin of both the prostrate and erect frond, | of numerous pinnately arranged, spine-like, minute projec- tions or teeth. The midrib, in its normal state, is the slightly thickened median portion of the frond, due to an increase in size of the cells of the central layer. Unless the frond is stimulated to further growth by injury, the midrib remains in this state, and, on reaching the upper branches of the frond, soon evanesces. Under the process of weathering, however, the margin becomes worn away. This seems to incite the cells of the median portion to renewed growth, with the result that the stalk and definite midrib become thickened and almost cylindrical. This spe- cies does not commonly proliferate. The tetrasporangia are as yet the only reproductive structures observed. The antheridia and cystocarps have not, so far as can be learned, been seen in the species. It seems to the writer desirable to allow this species to remain under /Vtophyllum rather than to assign it to Veuro- glossum. The habit, sori, and inner structure furnish evi- dence for thus placing it. Since it was first reported from this coast, doubt seems to have existed in the minds of writ- ers as to whether it was a /Vitophyllum or a Neuroglossum. The synopses already in existence of the genus /Vewrog/os- sum and of the present species do not at all permit a defi- nite idea to be gained of the internal structure, and widely varying statements exist as to the position of the sori. But the careful study of an extensive range of specimens has served to establish the writer’s belief that for the reasons above mentioned the species can be referred to Vitophyllum until a comparison can be made with type specimens: (3) July 28, 1900, 34 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Habitat.—On other alge and on rocks covered with Porif- era and Bryozoa at low tide-mark in the littoral and sub- littoral regions. . Distribution.—lV. andersonianum has now been observed along the coast from San Pedro northward to Carmel Bay. It has never been recorded elsewhere than from the coast of California. Localities.—San Pedro! (Mrs. E. A. Lawrence); Santa Barbara! (Dr. L. N. Dimmick; Mrs. 5S. P. Cooper); shores of San Luis Obispo County! (Mrs. R. W. Sum- mers); Carmel Bay! (C. P. Nott); Pacific Grove! (Mrs. J. M. Weeks; M. A. Howe); Santa Cruz! (Dr. C. L. Anderson, Mrs. Boardman). Nitophyllum ruprechtianum 7. Ag. Bidrag till Florideernes Systematik. Lunds Universitets Arsskritft., Tome WWADUIS stoly rus goyy Fos te Nitophyllum ruprechtianum FarLow, W. G., Proc. Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for 1875, p. 696, 1876, AGARDH, J. G., Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 470. Nitophyllum flabelligerum AGARDH, J. G., loc. cit., p. 699. Nitophyllum ruprechtianum Hervey, A. B., Sea Mosses, 1881, p. 178. ANDERSON, C. L., Zoe, Vol. II, 1891, p. 223. Hows, M. A., Ery- thea, Vol. I, 1893, p. 68. McCratcuis, A. J., Proc. So. Cal. Acad. Sci., Vol. I, 1897, p. 358. TILDEN, J. E., American Algz, Century III, No. 213, 1898. AGARDH, J. G., Contin. Spec. Gen. et Ord. Alg., Vol. Til, Pt: 32,1898, p94: Nitophyllum marginatum AGARDH, J. G., loc. cit., p. 93. Nitophyllum farlowianum AGARDH, J. G., loc. cit., p. 95. Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, flat, narrowly membranous, or linear; with rhizoids; frequently proliferating, lobed or branching; often provided with midrib and nerves; margin entire, cuneate, or lobed; frequently forming offshoots or innovations. Erect frond stalked, with midrib, nerves, and veins; branching, very often proliferating; margin entire, cuneate, undulate, crispulate, laciniate or lobed, often beset with minute proliferations. Branching subdichotomous or sub- palmate, with branches linear and often much prolonged, occasionally alter- nate below, becoming expanded and cuneate above, frequently divided or lobed. Stalk linear, flat, with definite midrib, very often becoming cylin- drical through wearing away of margin and. thickening of midrib, frequently twisted by wave action, often persistent and freely proliferating. Midrib Bot.—Vot. II.] MOT7T—CALIFORNIAN NITOPAYLLA. 35 conspicuous, branching, sometimes divided into usually conspicuous flabel- late nerves and veins, or unfrequently remaining undivided and evanescent; frequently weathered, thickened, persistent, and proliferating freely from sides and end. Nerves flabellate, free or anastomosing, often conspicuous, or inconspicuous and evanescent, sometimes dividing into minute veins. Proliferations produced very freely, on stalk, on margin of frond or on reduced frond, minute and rounded or large, linear, cuneate, frequently lobed or divided, with midrib and flabellate nerves, often bearing sori and cystocarps. Sporangia in linear sori flabellately disposed about margin of frond, or in linear or irregular submarginal patches, or upon proliferations abundantly produced along margin of frond or upon its surface. Antheridia as yet unobserved. Cystocarps large, infrequent, projecting, irregularly disposed over both surfaces, or gathered together along the margin, or borne upon marginal or surface proliferations. Ltemarks on the Species.—Considerable variation in color may be observed in JV. ruprechttanum. The plants, when young, are often bright red, becoming dull red or carmine with increasing age. The proliferating fronds often exhibit this change in color. When dried, the plant becomes a deep carmine to blackish red. A length of 20-30 cm. is not uncommon, as the plant is vigorous in its growth. Fre- quently a large number of abundantly branched fronds develop from a single stalk. Good specimens often form masses 30 cm. in diameter and 30 cm. or more in height. The prostrate frond in JV. ruprechtianum is developed rather more extensively than in any other species of the coast. It is usually membranous in character, and may either be undifferentiated to any extent or occasionally be provided with midrib and nerves. Owing to the free devel- opment and frequent branching of the prostrate frond, there is formed on the substratum an extensive ramification whose outer ends, by the decay or accidental rupture of the older portions of the frond, become separated, and constitute the starting point for a new frond. . The predominant characters of JV. ruprechtianum are displayed in the robust habit, the abundant proliferations, and the variety in the position of the cystocarps and sori, and the shape of the latter. Proliferation takes place more abundantly in JV. ruprecht- tanum than in any other of the Californian forms. The frond very commonly becomes worn away by the action of 36 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. the waves, and by friction upon other alge and upon rocks. The portions thus reduced persist for a considerable time and give rise to numerous proliferations which exhibit all the characters of the original frond. It seems strongly probable that the remarkable degree of variation in the shape and position of the sori in JV. ruprechtianum has been a fruitful source of error to those who have been called upon to identify the forms belonging to this species. It is difficult, perhaps, without extensive examination of plants on the shore at all seasons, to realize what a diversity of form may be found within the specific limits. Especially is this diversity important in considering the sori, which are usually much employed in establishing specific distinctions. The examination of abundant material of JV. ruprecht- zanum has shown that the plants fall into three groups dis- tinguished by the differences in the method of production of the sori, without regard to other features. In one group the sori are produced on the upper, flabellately expanded segments of the frond, and are arranged in rather wide linear sori, or lines, distinct or occasionally confluent, and ex- tending in a more or less connected fashion from the median portion of the segment to the margin. A second group exhibits these linear sori usually confined, however, to the apices of the segments, while along the margin are pro- duced numerous minute proliferations upon which are borne sori having the form of rounded patches. In a third group, the sori are confined to the marginal proliferations or to the similar proliferations appearing upon both the margin and the surface. While, in general, these variations in the posi- tion and shape of the sori are seen on different plants, yet it is of great importance in employing them as specific char- acters to keep in mind the fact that these three different dispositions of the sori are likewise found on one and the same plant. A similar habit with respect to the production of sori is seen in JV. violaceum, where, however, the linear sori are much narrower and the marginal ones are often widely Bot.—VoL. II.} MOTT—CALIFORNIAN NITOPHYVLLA. a linear and extend in some cases a considerable distance along the margin. When compared with JV. harveyanum, which also bears linear sori arranged likewise in flabellate fashion, it is seen that, while the resemblance in the pro- duction of the sori is strong, /V. harveyanum possesses a darker purplish tint, is not so robust in habit, and is desti- tute of such a well developed system of venation as belongs to LV. ruprechtianum. The agreement between JV. ruprechtcanum and JV. viola- ceum in the production of sori and venation is much more marked, yet the two can be distinguished by the character- istic violet hue and papery texture of JV. v7olaceum, in con- trast to the dull red color and leathery texture of JV. ruprechtianum. The same variety of position as is seen in the case of the sori may also be observed in the cystocarps. These struct- ures may be borne by JV. ruprechtianum, either upon the surface, when they are scattered at irregular intervals over the entire surface or form a border just within the margin, _ or they may be borne singly or several together on surface. or marginal proliferations. The existence of such numerous transition forms between the two extremes of surface and marginal production of the sori, a fact clearly established by a careful examination of material from a long extent of coast, points to the conclu- sion that within the limits of the species known as JV. ruprechtianum it is possible to include a wide range of forms characterized by the features already pointed out, and that sufficient grounds do not exist for the establish- ment of several species among which these forms may be distributed. The evidence afforded by these transition forms should, therefore, be employed in examining the species estab- lished by Agardh (1876, p. 699; 1898, pp. 93-96), viz., LV. flabelligerum, N. marginatum and JV. farlowtanum, which, so far as the writer can determine from the de- scriptions, have been split off from JV. ruprechtianum. The characterization of these three species is made to 38 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. depend upon differences in texture, branching, venation and sori. In the foregoing pages an attempt has been made to show the amount of variation possible in these characters, and to emphasize their relative value for spe- cific purposes. The numerous collections made between widely distant limits along the coast at various points and at different seasons of the year, as well as all the available material at hand in herbaria, have failed to yield forms which could not be referred to JV. ruprechtianum as defined in the foregoing account of the species. A word with respect to the general shape and segmenta- tion of the frond may be added in regard to the variation existing in /V. ruprechtianum. Forms may be found that tend to become elongated, with much prolonged, narrow, and acute segments. Near to these may be placed forms like- wise with the segments prolonged, but flabellately expanded, and with rounded apices. The tendency to expansion is seen to best advantage, finally, in plants that become divided into a few broadly wedge-shaped segments, with these in turn somewhat deeply lobed, with the lobes rounded as in the flabellate type. Flabitat.—lV. ruprechtianum especially occurs on rocks usually covered with Corallines or Bryozoa and Porifera, among which the prostrate frond attains a rich develop- ment, in deep rock pools on gently sloping shores, littoral to sublittoral zones. Distribution.—Along the coast from San Diego, Califor- nia, northward to Port Orchard, Washington. Localities.—San Diego! (D. Cleveland); Point Loma! (Miss Minnie Reed); La Holla! (Miss Minnie Reed); San Pedro! (Mrs. S. P. Monks); Santa Barbara! (Dr. and Mrs. L. M. Dimmick; Mrs. S. P. Cooper); shore of San Luis Obispo County! (Mrs. R. W. Summers); Port Harford! (W. A. Setchell); Avila Beach! (Miss Mabel Miles); San Simeon! (E. Palmer); Carmel Bay! (C. P. Nott); Pacific Grove! (M. A. Howe; Mrs. J. M. Weeks; C. P. Nott); Santa Cruz! (Mrs. Boston; C. L. Ander- son; Mrs. J. M. Weeks); Duxbury Reef! (W. A. Setchell) ; Bot.—Vot. Il.] MOTT—CALIFORNIAN NITOPHYLLA. 39 Dillon’s Beach! (W. A. Setchell); Fort Ross! (W. A. Setchell; C. P. Nott); Port Orchard, Washington! (J. E. Tilden). Nitophyllum violaceum 7. Ag. Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 700. Nitophyllum laceratum Harvey, W. H., Ner. Bor.-Amer., Vol. II, 1858, p. 104. FARLOw, W. G., Proc. Amer. Acad. Arts and Sci., Vol. X, 1875, p. 365; Report U. S. Fish Comm. for 1875, p. 695, 1876. Nitophyllum violaceum AGARDH, J. G., Epicrisis Floridearum, Contin. Spec. Gen. et Ord. Alg., 1876, p. 700. FARLOow, W. G., Proc. Amer. Acad. Arts and Sci., Vol. XII, 1877, p. 238. Hervey, A. B., Sea Mosses, 1881, p. 180. ANDERSON, C. L., Zoe, Vol. II, 1891, p. 224. Howe, M. A., Erythea, Vol. I, 1893, p. 68. McCratcuir, A. J., Proc. So. Cal. Acad. Sci., Vol. I, 1897, p. 358. Nott, C. P., in Phyc. Bor.-Amer., CoLuins, F.S., HOLDEN, I., and SETCHELL, W. A., Fasc. VIII, No. 389, 1897. AGARDH, J. G., Contin., Spec. Gen. et Ord. Alg., Vol. III, Pt. 3, 1898, p. QI. Nitophyllum stenoglossum AGARDH, J. G., loc. cit., p. 92. Neuroglossum lobuliferum ? AGARDH, J. G., loc. cit., p. 121. Nitophyllum violaceum formum crispulum SETCHELL, Phyk. Bor.-Amer. Co.uins, F. S., Houpen, I., and SETCHELL, W. A., Fasc. XIV, No. 694, 1900. Synopsis.—Frond both prostrate and erect. Prostrate frond creeping, broadly linear, or membranous, with rhizoids, branching, without midrib or nerves; margin toothed or laciniate. Erect frond stalked, flat, linear, occasionally with midrib, with flabellate nerves; subdichotomously or subpalmately divided into numerous segments, in some cases finely laciniate, frequently proliferating; margin entire, finely serrate, crispate, or toothed; segments or branches in some cases broadly obcuneate, in other cases narrow, becoming broadly linear, much prolonged and flabellately expanded at apices or remaining linear. Stalk short, soon merging into the frond, usually without midrib, but with flabellate, sometimes anastomosing nerves extending into the branches; or long, somewhat nar- row and thickened, almost cylindrical, and again merging into the flabellately nerved branches. Nerves not very conspicuous, usually extending through- out the frond from base nearly to apex, becoming divided into flabellate, frequently anastomosing veins, the latter soon evanescent. Sporangia in narrow lines flabellately disposed, occasionally confluent, on upper segments of frond, or placed singly or in clusters along the margin, or upon marginal sporophylls, the latter appearing at intervals or in a dense fringe along the margin. Antheridia as yet unobserved. Cystocarps large, projecting, irregularly disposed over both surfaces, or submarginal, or upon marginal proliferations. Feemarks on the Species.—The color of JV. violaceum varies from a pale violet through bright violet red to purple violet, in both the living and dried states. The plants attain 40 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. a height of 20-25 cm. Though not so robust in habit as lV. ruprechtianum, still some specimens of VV. violaceum are vigorous in their growth, branching freely and forming large, handsome plants. The plant is papery or parchment- like in texture, rather brittle when dried, and does not adhere well to paper. lV. violaceum, as before remarked, agrees with JV. ruprechtianum in many morphological details, such as char- acter of branching, general shape and position of sori, and venation; but it may be distinguished from that species by its different color and texture, and the minute differences in size and shape of the sori. An extremely wide range of variation may be seen in the amount of dissection which the frond of JV. violaceum undergoes. On the one hand, there is found a form in which the frond becomes divided from the base into a great number of slender, much prolonged branches, which divide again and again, until finally the apices of the ultimate branches are prolonged in a flabellate fashion sufficiently to show the specific characters of venation and color. On the other hand, there occur forms rather broadly membra- nous at base, that divide into a few broadly obcuneate seg- ments cleft from the outer edge into narrower portions which are prolonged into lobes, again exhibiting the charac- ters of the species. Between these extreme types may be found intermediate forms that in some cases vary toward the finely dissected frond, in others, toward the broadly membranous frond. Stunted’ and weathered plants also occur in considerable numbers in late winter and spring. The frond in these instances is occasionally thickened and rather fleshy, while the sori are gathered into clusters of mar- ginal proliferations or remnants of the former margin. Pro- liferation takes place, though not to the same extent as in LV. ruprechtianum. ‘The proliferations are, as a rule, small, and of varying size. They usually bear sori. In common with JV. ruprechtianum, IV. violaceum chal- lenges attention by reason of the peculiarities of its sori, which exhibit again the same range of variation in regard Bot.—Vot. II.} MOTT—CALIFORNIAN NITOPHVLLA. 41 to their shape and position that was observed in JV. ruprecht- zanum. There are to be found forms with the sori arranged in flabellate, occasionally confluent lines, differ- ing, however, from lV. ruprechtianum in the relative nar- rowness of the lines. Numerous transition forms occur which combine the flabellate linear sori with rounded. or linear patches upon the margin. Finally, frequent instances may be found of plants bearing the sori as rounded or semicircular patches along the margin, or upon marginal proliferations. The same conclusion that was reached in regard to the limits of the species in the case of JV. ruprechtianum may be applied to LV. vzolaceum, for here almost exactly similar conditions exist as to specific characters. The variations are in similar directions. Extended study of the forms from numerous localities has shown that they may be referred along the lines there laid down, which leads to the conclu- ‘sion that /V. violaceuwm is a species with limits sufficiently wide to include all the forms common to the coast. The two species established by Agardh (1898, pp. 92 and 121), viz., /V. stenoglossum and Neuroglossum lobuliferum, seem, therefore, to the writer, as nearly as can be deter- mined from the descriptions, to be forms of JV. vzolaceum. Hlabitat.—On rocks covered with Corallines, Bryozoa and Porifera, in sheltered situations 6r in rock pools on gently sloping shores, and on piles of wharves, in the littoral and sublittoral zones. Distribution.—Along the coast from San Pedro north- ward to Fort Ross. Localities.—San Pedro! (Mrs. Lawrence; Mrs. S. C. Purdy); White’s Point! (A. J. McClatchie); Santa Bar- bara! (Dr b.. N. Dimmick; Mrs. 5. P. Cooper); San Simeon! (E. Palmer); Carmel Bay! (C. P. Nott); Pacific Grovel( Mi AD Howe; ©. P: Nott); Santa ‘Crug! (Dr. C. L. Anderson); Land’s End, San Francisco! (C. P. Nott); Fort Point, San Francisco! (M. A. Howe; W. A. Setchell; C.P. Nott); North Beach, San Francisco! (W. A. Setchell; C. P. Nott); Golden Gate, San Francisco Bay (Berggren, 42 CALIFORNIA ACADEMY OF SCIENCES. [PRoC. 3D SER. fide J. G. Agardh, Epicrisis Floridearum, 1876, p. 700) ; San Francisco! (G. W. Lichtenthaler); Lime Point, San Francisco Bay! (C. -P. Nott); Duxbury Rees] (WwW. 22 Setchell); Fort Ross! (C. P. Nott). By way of summarizing some of the features mentioned in the foregoing account, the following table, showing the distribution of species within Californian limits, is incorpo- rated. It will be seen that ten species occur on the coast. These have now been reported from twenty-eight localities, ranging from San Diego northward to Puget Sound, Wash- ington, and points on Vancouver Island, B. C., embracing twenty degrees of latitude and fourteen hundred miles of coast line. It may be pointed out that there is here a coastal distribution equal to that of the Atlantic shores of Europe and the Mediterranean together. In the light of the evidence collected from a wide range of material in field and herbarium, the ten species recog- nized may be considered valid until more extended com- parison with European specimens proves their identity with previously described species. Bot.—Vot. Il.]} MOT7T—CALIFORNIAN NITOPAYLLA. TABLE OF DISTRIBUTION OF SPECIES. (San Diego, Calif., northward to Esquimault Bay, B. C.) Si eseate Wit Seth eel ae Pen g S se lass > 3 Ka SS eet Se hee Ae cd ed ae | ee eg Si esti Sears 5 = | 8 : = 3 -|* 3 AMO ICZ Or ity 16 aise oe late sece x RO soma series. * [ING 0) Ra Sr * SaMpedrOcdas soece ceeds e * Wihitte’SiPointe5 visas secs Soicae Wonicas masses & Salita Babbatas ence ves see 2S * a San Luis Obispo County.| * * * Port Harfords.. 25.36 2.4). * Avila each tctss sete artes 5 San Simeon: . .: 62... << « WanmeleBayirceien Sees nase % x 2 Rachie Grover. ssceo ase * * * * *e Seite Civesueon suscosen * * # * % * * TEANGISMELN GE: siere aarecs cores # ‘ HOt Ott terscoe esses * Golden Gate............ xP |x ee PNGHiaY BEACH 2 ace iis ceu San PPAMCiseOs yo. 25scne s # % ime; Botte oe sss ieee * 2 Duxbury Reef. .......\...... * # Dillon's Bede. ie...s0,<% “ = Ont ROSSia. cs uae eee * * * Cape Mendocino........ * Klatsop, Oregon........ # Port Orchard, Wash..... * * Puget Sound, Wash..... * Esquimault Bay, B. C.... * “UNZDULIUN * AT *WN9dN1 020 * AT 44 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. The writer is deeply indebted to Professor W. A. Setch- ell, upon whose suggestion this paper was undertaken, not only for collections of material and the use of his private herbarium, but more especially for his discriminating advice and helpful suggestions throughout the entire work, as well as for a comparison of some coast forms with specimens in the herbarium of Professor W. G. Farlow. Further acknowledgment is made to Professor Farlow, who kindly permitted an examination of the /V/ophylla in his collection, and to Mr. F. S. Collins and Dr. C. L. Anderson, both of whom courteously forwarded to the writer all the /Vzto- phyla contained in their herbaria. Acknowledgment should also be made to Mr. Collins for the specimen from which the description of JV. corallinarum was drawn up. To Mrs. J. M. Weeks the writer is especially indebted for collections which permitted a much more comprehensive account of certain species, and also for the privilege of examining specimens in her herbarium. The herbarium of the Uni- versity of California was consulted frequently in the prepa- ration of this paper. Special acknowledgment is due to Professors O. P. Jenkins and G. C. Price for their courtesy in extending to the writer during several seasons the privi- leges of the Hopkin’s Seaside Laboratory at Pacific Grove, California, thus facilitating the examination of a large amount of living material, at different times in the year. UNIVERSITY OF CALIFORNIA, BERKELEY, June, Ig00. Bot.—Vot. Il.] MOTT—CALIFORNIAN NITOPHAYLLA. 45 LITERATURE AND EXSICCATZ CITED IN THIS PAPER. 1852. AGARDH, J. G. Spec. Gen. et Ord. Algarum, Vol. II, Pt. 2. 1871. Bidrag till Florideernes Systematik. Lumnd’s Untiversitets Arsskrift, Tome VIII. 1876. Epicrisis Floridearum. Contin. Spec. Gen. et Ord. Algarum. 1898. Contin. Spec. Gen. et Ord. Algarum, Vol. III, Pt. 3. 1878. Algze Americanz-Borealis Exsiccate. FARLOw, W. G., ANDERSON, C. L. and Eaton, D. C. 1891. ANDERSON, C. L. List of California Marine Algze. Zoe, Vol. II, No. 3. 1877. Eaton, D. C., in Farrow. Proc. Amer. Acad. Arts and Sci., Vol. XII, p. 245. 1875. FARLOw, W.G. List of the Marine Algz of the United States, etc. Proc. Amer. Acad. Arts and Sci., Vol. X. Id. Report U. S. Fish Comm. for 1875. On Some Algze New to the United States. Proc. Amer. Acad. Arts and Sci., Vol. XII, No. 20. 1858. Harvey, W. H. Nereis Boreali-Americana. 1862. Notice of a Collection of Algze made on the Northwest Coast of North America, chiefly at Vancouver’s Island, by David Lyall, in the years 1859-61. Proc. Linn. Soc., (Botany), Vol. VI. 1889. Hauck und Ricuter. Phykotheca Universalis, Fasc. VII. 1881. Hervey, A. B. Sea Mosses. 1893. Hower, M. A. A Month on the Shores of Monterey Bay. rythea, ; Vol. I, No. 3. 1889. KjyELLMAN, F. R. Om Beringshafvets Algflora. XK. Svenska Vetensk.-Akad. Handl., Tome XXIII, No. 8. 1897. McCratcuir, A.J. Proc. So. Cal, Acad. Sct., Vol. I. 1899. NorpDHAUSEN, M. Zur Anatomie und Physiologie einiger ranken- tragender Meeresalgen. Pringsheim’s Jahrbiicher f. wiss. Botantk, Band XXXIV, Heft. 2. 1897a. Notr, C. P. Some Parasitic Floridez of the Californian Coast. Erythea, Vol. V, No. 9. 1876. 1877. 18976. In Phykotheca Borealis-Americane. COLuins, F.S., HOLDEN, I. and SETCHELL, W. A. Fasc. VII. 1897¢. 1. c. Base, VIE. 1897. Phykotheca Borealis-Americane, Cou.uins, F. S., HOLDEN, I. and SETCHELL, W. A. Fasc. VII. 1900. | l.c. Fasc. XIV. 1898. TILDEN, J. E. American Algz. Century IIT. 46 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. EXPLANATION OF PLATE I. Nitophyllum latissimum. Fig. 1. Tetrasporic plant of late spring, with broad, flat veins and large areoles. One-third natural size. Fig. 2. Cystocarpic plant, otherwise as in fig. 1. One-third natural size. Fig. 3. Antheridial plant, mature, showing in the lower third the character- istic wrinkled surface of the antheridial plant. One-third natural size. oe [Nott] PLATE T. ea 1 PRoc.CALACAD. Ser.3" SER. Bor. Vol FHOTO-LITH. BRITTON & REY, 5.F- NITOPHYLLUM LATISSIMUM. 48 CALIFORNIA ACADEMY OF SCIENCES. (Proc. 3p SER. EXPLANATION OF PLATE II. Nitophyllum latissimum. Fig. 4. Typical cystocarpic plant, showing portion of prostrate frond, vena- tion, proliferation, cystocarps, and a portion of the creeping, claw-like prostrate frond. One-third natural size. Nitophyllum spectabile. Fig. 5. Typical tetrasporic plant, showing prostrate frond, young and mature erect fronds, and position and shape of sori. One-third natural size. ] Piate. IL. ak j (Nt ER. Bort. Vou Il uJ Panic CALAGAD. Sc1.47 FHOTO-IITH BRITTON & REY, SF. ILE CTA Fis.5. NITOPHYLLUM SPEI M LATISSIMUM. LLU HY DH U 16.4. NIT E i= be, Ne at at oy = b) 7 ¥ =~ ‘ Py al ‘ ate Res mm : : oe - 50 Fig. . IO. EL. Pa 9 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE III. Nilophyllum fryeanum, Cystocarpic plant, showing disposition of cystocarps and segments of frond. One-third natural size. Tetrasporic plant, showing palmate frond with much prolonged segments, arrangement of sori and delicate veins. One-third natural size. Tetrasporic plant, showing venation and attenuate segments of frond. One-third natural size. Cystocarpic plant, showing pinnately arranged segments of frond. One-third natural size. Nitophyllum corallinarum. Tetrasporic plant, showing frond creeping over branches of Corad- lina chilensis, and sori borne on free, erect fronds, e. g., at x and x’. One-half natural size. Nitophyllum multilobum, Tetrasporic plant, showing habit and characteristic transverse sori. One-half natural size. Nitophyllum uncinatum. Sterile plant, showing character of branching and «characteristic recurved apices of branches. One-half natural size. Proc.CaLAcan. 5c1.32 Ser. Bor. Voult. PHOTO -TATH. BRITTON & REY, 5.F MULTILOBUM. TCPHYLLUM EANUM. Fig. li. Nt Fig 12. NITOPHYLLUM UNCINATUM Fies.6-9. NITOPHYLLUM FRY! F1e.10. NITOPHYLLUM CORALLINARUM, SP NOV. ft wel 52 Fig. Fig. 3. 14. Bais a ee CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE IV. Nitophyllum harveyanum. Slender form, tetrasporic, showing habit. One-half natural size. Robust form, showing linear sori and flabellately expanded habit. One-half natural size. Nitophyllum ruprechtianum. Prostrate frond of the stout membranous type. One-half natural size. Nitophyllum fryeanum. Prostrate frond of the delicate membranous type, showing rhizoids as minute processes on surface. One-half natural size. Nitophyllum andersonianum. Prostrate frond, of the freely branching type, with portions of the bases of erect fronds. One-half natural size. Proc.CALAcAn. Sc1.3? Ser. Bor. Von ll [Nort] Plate IV PHOTO-LITH. BRITTON & REY, 6 F- Fies 13-14. NITOPHYLIUM HARVEYANIM. Fic.18. NITOPHYLLUM FRYEANIM FIG.15. NITOPHYLLUM Beer HTIANUM Fig.17. NITOPHYLLUM ANUERSONIANUM = i < alae nie a es » 4 -. eae a4 Fig. Fig. Fig. Fig. 20. 21. CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER, EXPLANATION OF PLATE V. Nitophyllum andersonianum. Robust form, sterile, showing usual character of branching, with branches flabellately expanded at apices. One-third natural size. Broad membranous form of quiet waters, sterile. One-third natural size. Slender form, sterile, showing fine dissection of frond. One-third natural size. Transition form, tetrasporic, much branched, showing tendency to become membranous. One-third natural size. te. LATE V p I [Nort | Proc.CALAcan. Sci.3? SER Bor. Vou] & REY, ITH. BRITTON NITOPHYLLUM ANDERSONIANUM 7 - at e pu ae ia raat AT mi } \ or hin ie id oS ‘bat Foye BY i aS) ‘a AD ‘ aus * } , ' ee) : r y . x AYO aS hg f A NES . ‘ . c a . ’ ’ - . ‘ dl - . . z a i . ih (itn : A 4 ™ e - * aig Ae j " ie 4S ¥ 1 4 ‘ a 4 . , . 4 n J . i ¥ j ”: + f it J 2 ie > 2 * ry . 56 Fig. Fig. Fig. Fig. 22; 25: 24. 25. CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. EXPLANATION OF PLATE VI. Nitophyllum ruprechtianum., Expanded, membranous frond, sterile. One-fifth natural size. Examples of weathered and proliferating plants. One-fifth natural size. Slender, much prolonged type, minutely proliferating along margin, sori on proliferations. One-fifth natural size. Typical plant, showing stalk, branching, venation, and proliferations, with sori on proliferations. One-fifth natural size. Proc. CALACAD. Sc1.a? SER. Bor. Vou ll. [Nort] PLATE VI PHOTO -LITH. BRITTON & REY, 5.F NITOPHYLLUM RUPRECHTIANUM 58 20: es CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE VII. Nitophyllum ruprechtianum. Typical plant, showing stalk, branching, venation, proliferations, and sori on proliferations. One-third natural size. Portion of frond, showing flabellately disposed sori. One-third natural size. Same, showing sori (linear) confined to apices of branches. One- third natural size. Weathered frond, cystocarpic, with prostrate frond. One-third natural size. Portion of frond, showing linear sori confined to ends of branches and rounded sori on marginal proliferations. One-third natural size. LI q IER. BOT. Vi BS ae PRoc.CAL.ACAD. SCI Me 2) v » “= 60 Fig. Fig. F ig. Fig. Fig. Fig. Fig. Su S2. 33: 34. 35: 36. 37- CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. EXPLANATION OF PLATE VIII. Nitophyllum violaceum. Intermediate form, showing segments of frond, with crispate mar- gin and marginal sori. One-fifth natural size. Typical form, showing: slender, linear branches, marginal prolifer- ations, and sori forming linear patches along margin or rounded patches on the proliferations. One-fifth natural size. Intermediate form, with branches flabellately expanded at ends; sori as in preceding figure. One-fifth natural size. Robust, membranous form, sterile. One-fifth natural size. Reduced and weathered form, the conspicuous sori placed singly or together along the margin, or borne on proliferations. One- fifth natural size. Weathered form, as in preceding figure. One-fifth natural size. Finely dissected form. One-fifth natural size. Proc.CaAL.Acan. Sc1.32 SER. Bor. Vout. [Nort] PLATE VIII PHOTO-LITH. BRITTON & REY, ©.F- NITOPHYLLUM VIOLACEUM 62 Fig. Fig. Fig. Fig. Fig. Fig. 38. 39- 4o. Al. 42. . 43. - 44. 45- ig. 46. CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. EXPLANATION OF PLATE IX. Nitophyllum ruprechtianum. Portion of tetrasporic plant, showing rather wide, linear, flabellately disposed sori. Four-fifths natural size. Same, showing transition from linear, flabellate sori to rounded sori borne on marginal proliferations. Four-fifths natural size. Same, showing sori borne only on marginal proliferations. Four- fifths natural size. Nitophyllum violaceum. Portion of tetrasporic plant, showing narrow, linear, flabellately disposed sori. Four-fifths natural size. Same, showing transition from linear, flabellate sori to rounded, inframarginal sori. Four-fifths natural size. Same, showing rounded inframarginal sori. Four-fifths natural size. Nitophyllum latissimum. Detail of tetrasporic sori, showing venation, and areoles occupied by minute sori. Four-fifths natural size. Nitophyllum andersonianum. Portion of tetrasporic plant, showing sori as rounded or elliptical patches on distal segments of frond. Four-fifths natural size. Nitophyllum multilobum. Portion of plant, showing characteristic transverse sori. Four-fifths natural size. My “ EEN te : An ne eae The: Development of the ‘Karyokinetic "Spindle. in n the Pollen-MotherCells q At a et + Hot Lavatera | Ebr Sumver ‘Byxpee, ee new Pthatta eth ok? _ PUBLICATION. COMMITTEE, +H, Guzexn, Chairman. ‘ Ae G P, Rixroxo. Toes ae 2 A Helo % X my oly hs $4 lav of yy) Nid "EDITORS. or BO PANTCAL ‘PUBLICA ATION. ale q) 4 | Dovenas H. Campsett, Ma th) {5 ha: Wea ; PROCREDINGS As ri . ‘ OF THE CALIFORNIA ACADEMY OF SCIENCES THIRD SERIES BoTANY Vou? Il, No? 2% ae . The Development of the Karyokinetic 4 _ Spindle in the Pollen-Mother-Cells of Lavatera “A Toe tL 2 WEW YORK - RBOTANICAL SA RHEM: BY EpitH SUMNER BYXBEE Witru Four PLATES Issued October 31, 1900 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1900 THE DEVELOPMENT OF THE KARYOKINETIC SPINDLE IN THE POLLEN-MOTHER-CELLS OF LAVATERA|! BY EDITH SUMNER BYXBEE. PLATES X-—XII i. THE way in which the spindle is formed varies widely in the different families of plants that have been studied. In the generative cells of higher plants, the spindle seems to be always multipolar at first, but the formation of the poles may proceed in several different ways. With a view to shedding further light on this question, the pollen-mother- cells of a species of Lavatera were selected for study.’ This plant blooms throughout the year, so that the mate- rial is plentiful at all times, while the arrangement of the flowers in dense racemes and the number of anthers ina flower make it easy to obtain cells in all stages of division. The pollen-mother-cells are large. They may be examined before preserving, so that much timeis saved. The flowers were usually gathered in the morning, as it was found that then the cells were dividing more rapidly. The anthers were examined by crushing them, either with or without the addition of one per cent. glacial acetic acid. All heads in a favorable condition were then immediately dropped into the fixing fluid. It may be mentioned here that all the more striking appearances observed in preserved material were also seenin the fresh. The granular zone, especially, stands out with great distinctness by reason of the strong refrac- tive properties of its granules. 1 Contributions from the Botanical Laboratories of the University of California, No. 13. Presented for the degree of Master of Science. Prepared under the direction of Dr. W. J. V. Osterhout. * Owing to the destruction of the plants, it has been impossible to determine the spe- cies with exactness. Itis probably either Lavatera unguiculata Desf. or L. micans L. [63] October 29, 1900. 64 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. The following fixing fluids were tried: Flemming’s strong mixture; one per cent. chromic acid; two per cent. iron trichloride; Wilson’s corrosive sublimate-acetic; Bove- ri’s picro-acetic; Lindsay’s potassium bichromate-platinum chloride-osmic-acetic; one per cent. palladium chloride and one-half per cent. iridium chloride. Flemming’s strong mixture was used undiluted and also with the addition of one, two and three parts of water. The quantities of chromic and acetic acids in the original mixture were also varied. Dilute solutions were found to shrink the cells much more than the strong ones. Varying the amount of chromic acid did not improve the action of the fixing fluid. An increase in the amount of acetic acid, however, gave the best results obtained. Flemming’s strong mixture with an excess of acetic acid was therefore almost exclu- sively used. Fair results were also obtained with palladium chloride and iridium chloride to which a small amount of glacial acetic acid had been added. After remaining in the fixing fluid for twenty-four hours, the anthers were washed in running water for six hours. They were then placed in a dehydrator’ for twenty-four hours, with 95 per cent. alcohol below and distilled water above. Some alcohol was then removed from the material and mixed with an equal volume of 95 per cent. alcohol. The material was transferred to this stronger mixture for two hours. By repeating this process three or four times the material was brought into 95 per cent. alcohol without shrinkage. It was left in 95 per cent. alcohol for twenty- four hours. It was then placed for six hours in each of the following, successively: absolute alcohol; absolute alco- holand bergamot oil (equal parts); bergamotoil; bergamot oil and paraffin, 47° (equal parts); paraffin 47°; paraffin 47° and paraffin 54° (equal parts); paraffin 54°. Sections 3 to 4 » in thickness were cut with the Minot wheel microtome. Of the stains tried, Flemming’s triple stain (safranin, gentian violet and orange ‘‘G’’) gave the best results. , 1 For a description of the dehydrator see Lawson, 1898, and Williams, 1899. Bot.—VOL. II.] BYXBEE—LAVATERA. 65 The pollen-mother-cells are large, and the diameter of the nucleus is equal to fully one-half that of the cell. The chromatin thread is thick and stains blue with gentian violet. The large nucleolus stains red with safranin. It containsa single vacuole. The linin either forms a complete network filling the whole nucleus, or is present as broken threads attached to the chromatin and nucleolus. In neither case does it stain. The cytoplasm is made up of two constituents, one fibrous, the other granular. The fibrous part forms a network com- posed of delicate threads crossing each other in every direction. These threads stain deep blue with gentian violet. The other element is composed of small granules, varying somewhatin size. These granules are scattered throughout the network, both between and upon the fibers, usually in sufficient quan- tity to give the cytoplasm a cloudy appearance. ‘They have a tendency to collect in small, denser masses at the inter- section of the fibers of the network. This tendency is especially noticeable in the earlier stages. A little later the granular matter is distributed more evenly. It stains a brownish yellow with orange ‘‘G’’. A cell in this stage is shown in fig. 1. The first change that occurs in the cytoplasm is the elon- gation in a direction parallel to the nuclear wall of the row of meshes immediately surrounding the nucleus (fig. 2). This process goes on until three or four rows of meshes outside the nucleus have become pulled out in this way (fig. 3). ‘These meshes become so long and narrow that, often, on casual inspection, there seem to be threads wound round and round the nucleus. On close examination, however, the meshes can always be seen. Within the nucleus the linin network has meanwhile broken up somewhat; the threads lose their smooth, transparent appearance and begin to stain blue. One or more additional vacuoles appears in the nucleolus, which begins to stain purple rather than red. Soon after this, the granular substance, which hitherto has been equally distributed throughout the cytoplasm, begins to collect in a denser mass immediately about the nuclear 66 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. wall, covering the elongated meshes of the cytoplasmic reticulum (fig. 4). It increases rapidly in quantity about the nucleus without decreasing throughout the rest of the cytoplasm. It is always densest close to the nucleus and from there shades gradually out into the cloudy mass of the cytoplasm. As this granular substance begins to accumulate, the meshes of the cytoplasmic reticulum, with the exception of a narrow zone surrounding the nucleus, become radially elongated (figs. 4 and 5). This arrangement, however, does not seem to have any particular significance and soon disappears. Ina very short time a zone of granular matter has collected about the nucleus, occupying from one-half to one-third of the space outside of it. It becomes so dense that it entirely obscures the reticulum within it, except that a few fibers may sometimes be visible close to the nuclear wall (figs. 5 and 6). By this time, the elon- gated meshes of the cytoplasm immediately surrounding the nucleus have been transformed into free fibers, which lie between the granular zone and the nuclear wall. Occa- sionally there is a felt of fibers bounding the outer edge of the granular zone, and, by reason of their deep blue color, standing out conspicuously against the yellowish brown granular matter (figs. 6 and 8). This is by no means constant. By the time the granular zone is completely formed, the radial arrangement of the reticulum outside of it has entirely disappeared (fig. 6). The granular zone is composed of a dense mass of granules, most of which are larger than those that gave the cytoplasm its cloudy appearance in the earli- est stage. In this form it remains unchanged until the end of the anaphase. While the granular zone has been gathering, the linin in the nucleus has increased somewhat in quantity and in staining power. About this time, the cytoplasm loses its regular structure, as shown in figs. 6, 7, etc. Soon the fibers immediately outside of the nuclear wall come into clearer view, as though the granular matter had withdrawn Bot.—Vot. II.] BYXBEE—LAVATERA. 67 from the wall a little, or had been used up at its inner edge (figs. 7 and 8). These fibers, as has already been stated, are probably derived from the elongated meshes of the cytoplasm. They are of much greater diameter than the linin threads, are smooth and stain deep blue. The extent to which these fibers are visible varies greatly in different cells. At some point the nuclear wall disappears and through the gap thus formed the fibers immediately without the nucleus begin to grow into the cavity (fig. 9). At first, these fibers can be distinguished from the linin threads of the nucleus by their greater diameter and smoother appear- ance. As the nuclear wall continues to disappear, how- ever, and the linin to thicken up, the fibers from within and without the nucleus mingle in an interwoven mass in which those of different origin cannot be distinguished (fig. 10). By the time the nuclear wall has entirely disappeared, the nuclear cavity is filled with a mass of interwoven fibers which is usually densest about the circle of chromosomes which marks the situation of the old nuclear wall (fig. 10). In some part of the nuclear space the fibers crowd closer together to form a denser mass and at the same time tend to range themselves so that they lie more or less parallel to each other (figs. 11 and 12). Soon a number of projec- tions appear in this mass as though it were being drawn out at a number of points. Figure 13 shows an earlier and fig. 14 a later stage in this process. This continues until a dis- tinct multipolar spindle is formed (fig. 15). Soon two principal groups of fibrous cones can be distin- guished in the multipolar spindle and in each of these groups one cone becomes the most prominent. Into these two all others are soon absorbed by the continued straight- ening out and converging of the fibers. Figure 16 shows a spindle in which the cones have almost reached their final position and in which there is only a trace of the third cone. The completion of this process brings the chromosomes into position at the equatorial plate just at the time that the perfect bipolar spindle is formed. 68 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. The completed bipolar spindle lies with its poles close against the inner edge of the granular zone, which has as- -sumed an oval shape to accommodate itself to the outline of the spindle. The spindle has sharply pointed ends (fig. 17). It is composed of two sets of fibers. One set runs from pole to pole forming the central spindle. The other runs only from the poles to the chromosomes to which the fibers are attached in bundles. These fibers contract and pull the daughter chromosomes toward the poles. When this process is completed the fibers of the central spindle no longer appear straight but have assumed a wavy appearance. As the daughter chromosomes approach the poles, the mantle fibers appear (fig. 18). About this time the granular zone loses its definite outline. It begins to break up in the plane of the equatorial plate of the spindle and gathers about the poles in two masses in which the daughter chromosomes lie em- bedded. ‘These two masses remain connected by a shell of granular matter which outlines the old spindle. Figure 19 shows the beginning of this process and fig. 20 a later stage. The cytoplasm seems to contain a greater quantity of gran- ular matter than at any previous time. Within these two granular masses are formed the daughter nuclei. These have at first a decided indentation on the side toward the spindle (fig. 20). Later they become spherical (fig. 21). The daughter nuclei are thus from the first surrounded by a granular zone which, by the time they are completely formed, has become relatively as wide and as dense as that about the mother nucleus. The central spindle fibers seem to disintegrate and when the nuclei are ready for the second division remain simply as lines of granules connecting the two granular zones (fig. 21). The second division, as far as could be observed, exactly repeats the process of the first. It was impossible to follow, under the dense granular zone, the elongating of the meshes about the walls of the nuclei, but the concentric lines of fibers were visible in many cases in the narrow space between the zone and the wall. The breaking down of the nuclear wall, the growing in of the fibers, and the formation of the multipolar and bipolar spindles occur as in the first division. Bot.—Vot. II.] BYXBEE—LAVATERA. 69 The planes of division of the daughter nuclei do not seem to be at all constant. Sometimes they divide in the same plane, sometimes in planes at right angles to each other, and there are all possible transitions between thesetwo. Figure 22 shows a cell in which the planes are at right angles to each other and fig. 23 one in which they are nearly parallel. The four daughter nuclei resulting from the second divi- sion become surrounded by granular zones just as did the nuclei resulting from the first division. These zones are usually very broad and dense. Connecting them are the mantle fibers across which the cell-plates are formed later on (fig. 24). The granular zone persists even in the pollen-grain, at least while it is young (fig. 25). It usually occupies at least one-third of the cell space outside of the nucleus. The cytoplasm outside of the zone also contains a great deal of granular matter. The most important fact in the method of spindle forma- tion above described is that the spindle is formed from free fibers and not from a network. That part of the cytoplas- mic reticulum which aids in the formation of the spindle is converted into free fibers at an early stage, long before the nuclear wall breaks down (figs. 5, 6, etc.). The linin net- work of the nucleus breaks up at an even earlier time. The fibers derived from these two sources become inter- woven but never form a true network. The granular zone, too, is more than usually prominent in Lavatera. Its significance will be discussed later. The higher plants hitherto most exhaustively studied, as E-quisetum (Osterhout, 1897), ZLarzx (Belajeff, 1894), Cobea (Lawson, 1898), and Passzflora (Williams, 1899), all show a certain general resemblance to each other and to Lavatera in the method of forming the spindle in the repro- ductive cells. No two of them, however, agree fully in the details. In all of them the first changes in the cyto- plasm are either a radial elongation of the meshes of the reticulum or a parallel drawing out of the first two or three rows adjacent to the nucleus. 7O CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. In Zarzx the radial precedes the parallel elongation, and is of short duration, to judge from Belajeft’s figures. In Equisetum, Passiflora, and Lavatera the parallel elongation is the first change observed in the cytoplasm; in Cobea alone it has not yet been observed. In Larix and Lavatera this condition persists. In the former case, the elongated meshes become part of the central network; in the latter they become transformed into free fibers. In Passzflora and A guzselum this condition is transitory. In Hguzsetum the parallel elongated meshes are drawn out into an indefi- nite mass of radially arranged fibers which grow out into the cytoplasm, become parallel to each other, and finally bend together into groups, so that, before its breaking down, the nuclear wall is surrounded by a number of cones. When the nuclear wall disappears, the fibers of these cones grow into the nuclear cavity, come into contact with the linin threads and the chromosomes, and form a multipolar spindle. In Passiflora the radial elongation of the meshes of the reticulum persists for some time. Some of the threads stain more strongly and present an outline suggesting the cones in Aguisetum, though they are not formed of free fibers. This condition of the cytoplasm is transitory and seems to have nothing to do with the formation of the spin- dle. This is formed directly from the network resulting from the union of the linin reticulum of the nucleus with that portion of the cytoplasmic reticulum immediately out- side of the nucleus. On the breaking down of the nuclear wall these unite to form a continuous network which fills the entire space within the granular zone. The network becomes pulled out at a number of points, and is changed into free fibers which form the multipolar spindle. In Larix, Cobea and Lavatera, no cones are. present before the nuclear wall disappears. It would seem that the development of these cones is correlated with the slight development of the granular zone. It hardly seems that it would be possible to have such cones in forms like Cob@a and Lavatera, where the granular zone is very dense. In Bor.—VOL. II.] BYXBEE—LAVATERA. 70 Larix, where there is less granular matter, long fibers extend out from the central network—which results, as in Passiflora, from the union of cytoplasmic and linin threads— to form by their contraction the cones of the multipolar spindle. In Cobea and Lavatera the spindle formation goes on within the dense granular zone. In Cod@a a network is formed as in Larix and Passiflora. In Lavatera the fibers seem to be always distinct and bunch themselves together into a dense mass. The multipolar spindle is formed by a pulling out of the network (Coéea), or mass of fibers (Lav- atera), as in the other cases. The method of formation of the bipolar from the multi- polar spindle does not seem to vary in the various cases. One of the principal differences, then, in the method of the formation of the spindle in the various plants studied, seems to be the time at which the free fibers are formed from the reticulum of the resting cell. In some cases this occurs very early, as in Hguzsetwm and Lavatera, which, however, differ widely in other respects. In other cases, as Larix, the spindle itself seems to be a network much stretched out. The granular zone, whichis so conspicuous in Coé@a and Lavatera, has been figured in most of the papers on the division of the ‘generative cells of the higher plants. Oster- hout (1897) figures it in the bipolar stage in Hguzsetum and Mottier (1897 a) in Podophyllum and Felleborus. Mottier (1897 6) also speaks of its presence at several stages in the divisions in the embryo-sacs of the Lz/zacee, but does not seem to regard it as constant in or characteristic of these divisions. Juel (1897) figures it as a prominent ring in Hemerocallis, but does not discuss it at any length. In Passifiora, also, it forms a well marked zone. While it is so commonly, perhaps invariably, present in the reproductive cells at the time of their division, it has not been observed in any of the dividing vegetative cells that have been studied. This fact seems to indicate a connection with the two rapid divisions of the reproductive cells for 72 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. which it apparently furnishes food material. The granules, too, strongly suggest the yolk granules present in animal eggs. They appear in small quantities in the pollen-mother- cells at an early period, but their exceedingly rapid increase at the time when the first steps in the actual formation of the spindle takes place strongly impresses one with the idea that the cytoplasm is busy producing nutritive material to serve the cell through the period of its activity. The manner in which the granular zone accumulates sug- gests the gathering of the deutoplasm in animal eggs ( Wil- son, 1896, p. 115 et seq.). In Pass¢fora it appears first in patches scattered through the cytoplasm and later gathers into a somewhat loose. ring at some distance from the nucleus. This is the way in which the yolk collects in the egg of Diemyctylus and other Amphibians, as described by Jordan (Wilson, 1896, p. 116). In Cobea and Lavatera the granular substance appears first close around the nucleus and spreads out from this as in the trout (Henneguy) and cephalopods (Ussow) (See Wil- son, 1896, p. 117). The exceedingly dense zone formed by this substance in Zavatera exactly resembles that figured by Van Bambeke for a fish (.Scorfena) (Wilson, 1896, p. 116, fig. 59, C). In some cases, as Cobea, the granular matter is used up as the divisions are completed. In Lava- tera, however, it constantly increases in quantity up to the formation of the pollen-grains in which it is present, at least in the younger stages. It seems probable that it per- sists in them and serves them for food during the develop- ment of the pollen-tube and the succeeding divisions. Bot.—VOL. II.] BYXBEE—LAVATERA. 73 SUMMARY. 1. The cytoplasm of the young pollen-mother-cell is made up of two constituents—a fibrous network and a granular substance. 2. The spindle is formed in the following manner :— a. The meshes of the network, close to the nuclear wall, pull out in a direction parallel to the wall, forming a felt of fibers about the nucleus. 6. The granular constituent of the cytoplasm col- lects in a wide, dense zone about the nucleus. c. The linin increases in quantity. d. The nuclear wall breaks down and the fibers outside begin to grow into the nuclear cavity. ée. The cytoplasmic and linin fibers form a mass in which the chromosomes lie. jf. The mass of fibers projects out at a number of points, forming the multipolar spindle. g. Two of the cones become more prominent than the others, which they finally absorb, thereby forming the bipolar spindle. 3. The process in the second division exactly repeats that in the first. 4. The granular substance, forming the dense zone, is comparable with the deutoplasm of animal eggs. 5. Finally, the spindle is formed directly from elements —cytoplasmic and linin reticula—present in the cell from the first, and not from any special spindle-forming sub- stance, or by the aid of centrosomes. BOTANICAL LABORATORY, UNIVERSITY OF CALIFORNIA, October, 1899. s¥ae> | Bot.—VOL. II.] BYXBEE—LAVATERA. 75 1894. 1897. 1898. 18974a. 18976. 1897. 1897. 1896. 1899. BIBLIOGRAPHY. BeLAjEFF, W. Zur Kenntniss der Karyokinese bei den Pflanzen. Flora, Bd. LXXIX, p. 430. Juet, H. O. Die Kerntheilung in den Pollenmutterzellen von Heme- rocallis fulva, und die bei denselben auftretenden Unregelmassig- keiten. Pringsh. Jahrb., Bd. XXX, p. 205. Lawson, A. A. Some Observations on the Development of the Karyokinetic Spindle in the Pollen-Mother-Cells of Cobzea scan- dens Cav. Proc. Cal. Acad. Sci., 34 Ser., Bot., Vol. I, p. 169. Mortirer, D. M. Beitrage zur Kenntniss der Kerntheilung in den Pollenmutterzellen einiger Dikotylen und Monokotylen. Pringsh. Jahrb., Bd. XXX, p. 169. Ueber das Verhalten der Kerne bei der Entwickelung des Embryo-sacks, etc. Pringsh. Jahrb., Bd. XXXI, p. 125. OstERHOuT, W. J. V. Ueber Entstehung der karyokinetischen Spin- del bei Equisetum. Pringsh. Jahrb., Bd. XXX, p. 155. STRASBURGER, E. Ueber Cytoplasmastructuren, Kern- und Zelltheil- ung. Pringsh. Jahrb., Bd. XXX, p. 375. Witson, E. B. The Cell. New York. 1896. Wixuiams, C. L. . The Origin of the Karyokinetic Spindle in Passi- flora coerulea Linn. Proc. Cal. Acad, Sci., 34 Ser., Bot., Vol. I, p- 189. 76 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE X. All figures were drawn with the Abbe camera lucida: objective, Zeiss oil immersion 1/12, compensating ocular No. 6. Fig. 1. A young pollen-mother-cell. The chromatin thread is beginning to break up. The cytoplasm is composed of two elements, one a fibrous network, the other a granular substance. Fig. 2. A little laterstage. The first row of meshes adjacent to the nuclear wall has begun to pull out. Fig. 3. A number of rows of meshes adjacent to the nuclear wall have be- come elongated. The chromatin thread has broken up. Fig. 4. The granular substance has begun to collect about the nuclear wall. The outer meshes of the cytoplasm have become radially elon- gated. The nucleolus shows four vacuoles. Fig. 5. The granular zone has increased in width. Fibers can be seen between it and the nuclear wall. The linin is thickening up. Fig. 6. The granular zone is completely formed. The radial arrangement of the meshes outside of it has disappeared. Proc.CALACAD.Sci.a2 SER.Bor. VoLIl. 7 HEL. EDTA S. ITH. BRITTON & REY, 5.F. Ean tw td af maths heey al ee aT nea teenie f on | i i oa be Pa tsa H hi ey | iA snee 1) » wis! a i 78 Fig. Fig. Fig. Fig. Fig. Fig. Io. Ii. 12. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XI. The space between the granular zone and the nuclear wall has widened and in this space numerous fibers are seen. The cyto- plasm has a less regular structure. About the same stage as that shown in fig. 7, but there are many more fibers outside the nucleus. There are also strongly staining fibers in the cytoplasm. The nuclear wall has disappeared at one point and the fibers out- side the nucleus are growing into its cavity. The linin has increased in quantity. The nuclear wall has completely disappeared and the space within the granular zone is filled with a mass of interwoven fibers; those near the center of the space are linin. The fibers begin to straighten out and arrange themselves in groups, in which they are parallel to each other. A more advanced stage in this process. - ee 5 separ See: 9. este anise ‘. ‘ be vt ta .y LIITH.BRITTON & REY, 5.F, at: + 8% on aes ee Co a TT Le , - zp 2 1 a? Cea —— ‘e is) iW eta x, Wes ve 80 Fig. Fig. Fig. Fig. Fig. Fig. > Fig. 13. 14. I5. 16, rps 18. 19. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XII. The fibers begin to converge at a number of points to form cones. The cones project out farther from the mass of fibers. The fibers have straightened out. The cones are in two groups. The cones have nearly fused to form the bipolar spindle. A completed bipolar spindle. The daughter chromosomes have nearly reached the poles. There are a few mantle fibers. The fibers of the spindle are no longer straight. The granular matter is becoming thinner at the equator and collect- ing more ‘densely about the poles of the spindle. There are traces of radiations which seem to be formed of granular matter extending out from the poles. PROD LALACAD, SETA? See Bor VoL IT. ot Ses wah te »?. EMTH 5. BYXBEE, DEL. [ByxpEr] PLATE XIL LiTH.BRITTON & HEY, 6.FL tn ; 82 Fig. Fig. Fig. Fig. Fig. Fig. 20. 21. 22. 23. 24. 25. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XIII. The daughter nuclei are formed. They are at this time crescent shaped as seen in section. There is still a shell of granular mat- ter bounding the remains of the spindle. It is seen in section as two lines. The daughter nuclei have become spherical. The granular matter has increased in quantity. A spindle of the second division. The other spindle, whose posi- tion is indicated by the circular mass of granular matter, lies at right angles to the first. Two spindles of the second division that lie nearly parallel to each other. The granular mass between them is not of equal density throughout. The four daughter nuclei, each surrounded by a dense granular zone. They are connected by mantle fibers. A young pollen-grain. The nucleus is surrounded bya dense gran- ular zone and the cytoplasm outside of this contains much gran- ular matter. Bor. Voull. eal RARAGANS SETA PROG Der asats ah “e - : EQITH &.5YXBEE, DEL. a? ” IITH. BRITTON & REY, 5E [BYXBEE | PLATE XIII. an ~d Pa —t is 7 — co raf lal “ ra « 4 nam A) e c i * » 4 i fr " r “ 7. . -< ‘ . it 8) ihe) Oe 4 Dos» | SO ee Wo Bist? rey ¥ EES cs ey { * i | Turrp Serre et age 5 iy i. ; Va zy - I J _ sempervirens E si ae aa re Gsorce Janns Peres — § ) A ibe aM ee pine 5 at a Associate Professor of Plant Physiology. ; = i : tet ‘Stanford Jr. With Ong Piate Tssued April 4, 1901 x Ba RTE at ‘ PUBLISHED BY THE ACADEMY Ate f ae a Oe ih he ys en 4 Bes ah ct ‘ a vw _“~ en -\ - B79 , * | 5 ‘ é ) x at E. R eA ‘ TO Gah Bete ERTL AA Shares mae oe - PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES THIRD SERIES BoTany Vou. Thi Mor, 4 Studies on the Coast Redwood, : Sequoia sempervirens Endl. 8 ep Wi 2 WY YORK ROTA HIIC A Senne a BY GEORGE JAMES PEIRCE Associate Professor of Plant Physiology, Leland Stanford Jr. University. WITH ONE PLATE Issued April 4, 1901 SAN FRANCISCO PUBLISHED BY THE ACADEMY _ Igol EC , ; 4 Las 7 if baie: vs } 1) itt t oC OP laa, ay ' i i vi \ DS ) i af Vi STUDIES ON THE COAST REDWOOD, SEQUOIA SEMPERVIRENS ENDL. BY GEORGE JAMES PEIRCE. Associate Professor of Plant Physiology in the Leland Stanford Juntor University. CONTENTS. PLATE XIV. PAGE. I. THe VEGETATIVE MODE OF REPRODUCTION IN THE REDWOOD... 83 Il. PECULIARITIES OF SOME VEGETATIVELY PRODUCED YOUNG EST V VO OID Sestae a ee ee ore PO felons ee acdc bin ei loeree kone Eee 88 PAL aVAGETA TION: foteriaeta sober o siaiee. scree girs nts We ee 88 TBP WIRTBEMISWEAN. vice vig heeds Oe) soles! 518)s So sleln do's bOnn eae em 89 Ill. Tue SIGNIFICANCE OF THE WHITE REDWOODS IN CONNECTION WITH OUR CONCEPTIONS OF PARASITISM AND OF HEREDITY..... 100 I. Tue VEGETATIVE MopE oF REPRODUCTION IN THE REDWOOD. Tue coast redwood (Seguoza sempervirens Endl.) is one of the comparatively few coniferous plants which reproduce themselves vegetatively by suckers. The majority of the Coniferz multiply only sexually, by seeds. Seeds offer the means by which plants may be carried in the dormant or resting condition for long distances, thus permitting ex- tended distribution. If the seeds retain their vitality, they offer also the means by which the species as well as the life of the individual may be maintained for long periods. A forest composed of other coniferous trees than the redwood will usually come to an end when the trees are felled. The coniferous forest may be succeeded by a wholly worthless growth of ‘‘scrub’’ oaks and other deciduous leaved plants, as for example on the Ossipee Plains in southeastern New Hampshire, or by hardwood forest of diverse composition, as in other parts of New England, or it may be followed by desolation, bare sand or rock, as in the region around | Truckee and about Lake Tahoe in California and Nevada. Not so with the redwood forest. [83] March 14, Igol, 84 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. In the National Geographical Magazine for May, 1899, Gannett asserts that a region naturally forested with red- wood will not become reforested with the same tree if the standing timber is felled. Hesays(l.c., p. 151) ‘‘Nowhere is there any young growth. The youngest trees, which are found only in the northern portion of the [redwood] belt, are several hundred years of age. When the timber has been cut, there is no sign of reproduction from seed. In many localities sprouts are starting from stumps in the cut areas, but even this form of reproduction is limited. In- deed, everything seems to indicate that for some reason, probably a progressive drying of the climate, the forest environment is not favorable to the growth of redwood, and that with the clearing away of the present forests, the end of the species as a source of lumber will be at hand.’’ Gannett furnishes a photograph of sprouted redwoods in a cut area. It is true, as Gannett says, that the major part of the red- wood forest is north of San Francisco, especially in Hum- boldt County, California; but in the Santa Cruz Mountains, south of San Francisco, there are more than merely ‘‘ scat- tered groves’ of redwood trees. The amount of redwood lumber here cut is evidence that the redwood has attained profitable size and that it still occurs in profitable quantity. Redwood forests must have been abundant in the mountains between the southern arm of the Bay of San Francisco and the ocean, even within comparatively recent years. Much of what was once forested land is now tilled, but by the roadsides and along the fences one sees great blackened stumps which prove the recent presence of redwood forest. In the cafions and on the steeper hillsides, where land can- not under present conditions be profitably cultivated. and need not be used for pasture, some of the old redwoods remain. In these same places young redwoods are coming up, some from the stumps, more from the uninjured and still living underground parts of trees which have been felled, and some from seed. So far as this region is con- cerned, the alarm which Mr. Gannett’s remarks arouse seems not to be well founded. BoT.— VOL. II.] PEITRCE—SEQUOIA SEMPERVIRENS. 85 The redwood is a tree of fairly rapid growth. I can not base my opinion on measurements, but I believe that the sprouts from the stumps and underground parts of old red- woods which have been felled grow faster than plants from the seed. It appears, from what I shall presently report, that in the living underground remnants of old trees there are great quantities of reserve food which are avail- able for the nutrition of sprouts. These sprouts or suckers are not wholly dependent upon the food they themselves elaborate. So long as their connection with the parent tree continues unbroken, and the remnants of the old tree retain their vitality, the young trees can use the food stored in the parent. For this or for some other reason, the young trees which begin as suckers and sprouts attain a considerable height and diameter within a few years. Young trees of this sort, in the cafions and on the mountain sides in various places which I have visited, not far from the Stanford Uni- versity, already have very considerable dimensions. Such trees may be seen above Portola and above Los Gatos. In comparison with the great redwoods of the virgin forest to the north, these young trees are very small. They are still absolutely as well as relatively of no value as timber, but they have been growing only a few years. If they are allowed to continue to grow, if they are reasonably pro- tected against drought by having the watershed above and about them as little disturbed as possible, I can see no reason why valuable redwood timber should not continue to be produced in these cafions at least. Mr. Gannett’s account of the lumbering operations in the northern redwood forest reveals one, and it seems to me an adequate, reason why the redwood forest is succeeded neither by a second growth of seedlings of its own sort nor by a vigorous growth of sprouts from the stumps and under- ground parts. Because the redwood is free from resin and contains much water, the freshly felled trees do not burn readily. For this reason, the lumbermen commonly clear away the rubbish around the trunk of a felled tree simply by setting fire to the brush. In this way the foliage and 86 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. smaller branches are consumed and the main trunk becomes accessible, blackened by the fire, the bark more or less burned through, but the wood uninjured. A fire hot enough to burn up so much green rubbish, though not hot enough to impair the value of the great felled trunks as lumber, is surely hot enough to do great damage to the superficial parts of the stump if not to kill it. Such a fire would probably heat the ground enough and deep enough to injure or kill the underground parts, and it would surely destroy all seeds not deeply buried in the soil. Land cleared in any such way as this usually has to be restocked by plants that wander in, their seeds being blown or brought in by wind, animals, man, etc. It seems to me, therefore, that the habit not of the redwood but of the lumberman is responsible for the failure of the northern redwood forest to renew itself. In view of these facts, is it not unnecessary to imagine any harmful change, if change at all, in the climate of the Pacific Coast since the redwoods have lived here ? Young redwood trees grown from the seed under some- what artificial conditions often send up suckers from the trunk at or slightly below the level of the ground. A con- siderable number of redwoods in the Arboretum of Stan- ford University are doing so. In some instances this may be due to injury to the upper parts of the tree by fungus or animal enemies, but apparently not in all. It seems to me much more likely that the more abundant branching at the bases of these young trees is a reaction to the larger amount of light which falls upon the surface of the soil and upon the lower parts of the trees in the comparatively open Arboretum than in the natural forest. But the Arboretum is not only more brightly lighted than the forest; it is warmer by day and colder by night during some if not all seasons of the year; it receives much less moisture both as rain and as fog than the naturally forest-clad hill and mountain sides; and the soil is not able to retain so much moisture because the ground is naked or nearly so. In the natural forest, unharmed by sheep or man, the ground is covered by a thicker or thinner layer composed of humus, decayed Bot.—VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS, 87 leaves, leaves only recently fallen, and a great variety of small plants—mosses, lichens, etc.—which form a turf. This covering of the forest floor, as every one knows, is the most important natural means of holding back water, restraining it from too rapid flow, and holding it against evaporation. Some or all of these differences between the natural and the artificial habitat of these trees may act upon them as stimuli to which the production of suckers is the visible reaction. The Palo Alto, the only large redwood tree still standing on the floor of the Santa Clara Valley, so far toward the Bay of San Francisco, has been subjected to many disturb- ing influences. Its crown has been seriously injured and its underground parts have been subjected to changed environment. The proximity of the railway embankment and bridge have caused changes in the drainage, both sur- face and subsoil, and other disturbances less evident must also have occurred. Around the base of this old tree, growing thickly and closely about it, is a brush or thicket of suckers. No young trees have grown up around the parent, forming a little grove such as one sees around the stump of a felled or fallen redwood of advanced age. Only these suckers are formed, close around the trunk, and these are not likely to attain any considerable height or size. So far as I know, it is only when conditions are unlike those prevailing in the natural forest, or when an old tree has been felled or injured or at least considerably disturbed above or below ground, that it sends up suckers from the trunk or stump, or that young trees come up from the re- moter underground parts. These last often make circular groves of greater or less size, known as ‘‘redwood temples.”’ Even in the group of giant redwoods at Felton, near Santa Cruz, one sees this arrangement clearly marked. The suckers and sprouts may attain great size in the course of time, as some of these giant redwoods show. In the production of suckers or sprouts from the trunks and underground parts of Seguota sempervirens, we see the vegetative mode of reproduction engaged in bya species of the Coniferz; but this recourse to the vegetative mode of 88 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. reproduction is probably the result of some external stimu- lus or stimuli acting upon the plant which, under unchanged conditions, would continue to reproduce itself by seeds if at all. This method of vegetative reproduction is probably sufficient, even under present climatic and other conditions, to secure the continuance of redwood forests in the regions where they now occur, provided lumbering operations are so conducted that the production of suckers and sprouts is not made impossible by destructive fires. Il. PECULIARITIES OF SOME VEGETATIVELY PRODUCED Younc REDWooDs. A. Fasciation. Fasciation of the young suckers coming up around the trunks of redwoods is not uncommon. The view advocated by Frank (1896), that they are the consequences of an excess of food substances, is strengthened by the time and manner of their appearance. Frank says that fasciations on other plants appear especially when the ordinary branches have been removed or injured in any way. We have seen that the redwoods produce suckers probably only when stimulated to do so by external influences, espe- cially by the removal or at least the injury of the parts above ground. The wound, or other injury, which stimulates the redwood to form suckers, may occur when there is such an abundance of food in immediately usable form, that the production and growth of suckers is so prolific as to insure the fusion of the adjacent parenchymatous parts of the very young branches. In the autumn and early winter, I have had no difficulty in finding fasciated redwood suckers in the Arboretum of Stanford University; they are very notice- able. In the spring and summer months they are by no means so common. I have found no new ones this spring, though there are many young suckers on the redwoods in the Arboretum. Summer, at least the earlier half of the dry season, is the time of food manufacture and storage. In the latter half of the dry season little food can be manu- factured because only little water is obtainable. In autumn Bot.—VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. 89 the food manufactured and stored becomes available and is used after the first rains have made it possible for growth to be renewed. If, as was the case last winter, there is much mild damp weather, growth will be luxuriant, the stored food will be freely used, the conditions for sucker-formation and for fasciation will coincide. In the spring, especially after a mild winter, during which much growth and comparatively little food manufacture have taken place, the stores of food having been considerably reduced, growth will be less luxu- riant and food manufacture will become more necessary and more active. We see, then, some reasons for the formation of the fasciations and for the time of their appearance. B. Albinism. The most remarkable and, I am surprised to find, not an especially rare peculiarity of the suckers or sprouts which come up from the stumps or from the old roots of felled or fallen redwood trees, is that they are sometimes perfectly white. My attention was first attracted to this peculiarity when, in the fall of 1898, a student brought some redwood twigs bearing white leaves into the Botanical Laboratory of this University. On inquiry I learned where these white redwoods were growing, and in the fall of 1899 I went to the spot. These are the only white redwoods which I have seen growing, but I have heard of others much larger and one which must be several years old was brought to the laboratory from the ‘‘ Redwood Retreat,’’ about twelve miles from Gilroy. This last I planted in my garden, but it lived only a short time, whether because it was injured in the transplanting or because it could not bear transplanting late in the spring I do not know. Its behavior before it died I will speak of presently (p- 95). The white redwoods which I have visited are on the sum- mit not far from the stage road between La Honda and Redwood City, and on the line to the left of the road (as one goes toward La Honda), where the forest gives place to open fields. The tallest redwood tree in view marks the spot where the white ones grow. This tall tree is one of a gO CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. number which are several decades old. They, and other smaller young trees of various ages, have come from the stump and roots of a much older tree which must have been very large when it was felled. The old stump has been repeatedly burned under and into, apparently by camp fires, but the heat could not have been enough to do more than local damage, and even this is not great. There is little left of the old stump above the general ground level, but as the hillside falls away abruptly at that point, a good deal of the underground part of the old tree is more or less exposed. Because of the irregularity of the surface of the hillside just there, the old tree sent its roots out more irregularly than is usually the case and the trees which have sprung up from them are not symmetrically placed. There is a thicket, but not a circle or ‘‘ temple,’’ of redwoods. All of these second growth trees are perfectly normal, as far as I could see. One buttress of the old parent tree, instead of sending up a few more or less scattered sprouts which grow up fairly rapidly and, within a season or two take on the characters of young trees in bark, foliage, and manner of branching, produces branches or bunches of scrubby, thickly set, short and slender sprouts or suckers. These are perfectly white as to leaves. The youngest parts of the stems are of the same ghostly color as the leaves. These white suckers may attain a height of thirty (30) centimeters in the course of one season. They began growing early in April this spring (1900), and they go on growing till hard frost comes. In the same length of time, and with a simi- lar origin, a green sprout or sucker would make two or three or more times this growth in length. The white suckers increase in thickness proportionally to their growth in length, that is, slowly, but the surface of the stem be- comes brown and develops cork sooner than the correspond- ing parts of the green suckers. This precocious cork-for- mation is not accompanied by other means of protection or by such vigor that the white suckers survive the hard frosts of winter. Even this last winter, milder than usual, was fatal to the white suckers; they were killed down to or just Bot.—VOL. II.] PETRCE—SEQUOIA SEMPERVIRENS. gi below the surface of the soil. The green suckers, on the other hand, are enough tougher to survive the winter. In this spot, therefore, white suckers with parts above ground which are two years old are not tobe found; but near and just under the ground are well-formed buds which, surviving the winter cold, form the next year’s growth of white suckers. In this difference in ability to resist cold, we have one of the physiological differences between the dependent white suckers and the independent green suckers. Whether this is merely a coincidence or a fundamental difference in vigor which forces the white suckers, unable to form chlorophyll, to draw food from the parent if they are to survive, who can tell? This difference between white and green suckers is not everywhere visible. White redwoods of fair height and age are reported, indefinitely to be sure, from various places in the Santa Cruz Mountains. I have seen white redwoods several years old. These, however, came from places of lighter frosts, if any frost at all touches them during the winter. Turning now to the anatomy of the white and the green suckers, we see certain peculiarities in the white which demand remark. The leaves evidently present the most marked differences. The leaves of the white suckers are similar in size, form, kind and arrangement to those of the green. There are two kinds of leaves on both green and white suckers—the early or young form, large, long, few, scattered along the stem or branch—and the later or mature form, smaller, shorter, more numerous, regularly placed along the branches, giving to these leafy branches the typical flat and thin dorsi-ventral aspect as compared with the more nearly radial arrangement in the young plant. These two forms, found on the suckers, are also found on seedlings. According to Goebel (1898), the young form is to be re- garded as presenting the original leaf-form and leaf-arrange- ment in the Gymnosperms. The rudimentary characters to be discussed later, which are found in the leaves of the white suckers, would lead one to believe that sooner or later 92 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. suckers would be found which retain this primitive charac- ter for considerable periods, if not throughout their exis- tence. If Goebel’s other supposition is true—that the basal branches, showing the young form of leaves, disappear by correlation as the mature form develops—one would infer the greater physiological perfection and effectiveness of the mature as compared with the young form. But in the white suckers the main function of leaves, that of photosyn- thetically manufacturing non-nitrogenous food, is entirely suppressed—the mature form of white leaves being as im- potent as the young form. Two other important functions of leaves—that of securing and controlling transpiration, and that of securing and controlling the aération of the plant-body,—would apparently be as perfectly prepared for and accomplished by the one form of leaves as the other, provided the texture and the surface-area of the two kinds of leaves are equal. In these two respects the mature leaves are superior to the young form. There is, therefore, some reason, other than the conception of an ‘‘inherited tendency,’ for the development of the mature form of leaves in the white suckers. Since the Seguodas are geologically such old plants, it is interesting to have the young form of leaves so clearly marked and so constantly recurring. There is a striking resemblance between some of the green suckers in their young condition and the great fronds of Cycas revoluta, the leaves of the redwood resembling the leaflets of Cycas in form, thickness and arrangement. Can this be a hint as to the origin, perhaps the common origin, of the Coniferz and the Cycadacee ? Comparing the mature form of leaves of green and of white suckers from the same localities, one finds that, despite the superficial likenesses, there are decided struc- tural differences. These are at once evident in cross- sections of the leaves, as shown in figs. r and 2. Figure 1 is a diagram of a cross-section of a small green redwood leaf, the single vascular bundle occupying the centre of the leaf, one resin-tube lying under it, the other two resin-tubes Bot.—Vot. II.] PEIRCE--SEQUOIA SEMPERVIRENS. 93 being located at the ends. The upper surface of the leaf is convex, the lower concave, the upper surface and the edges being greatly strengthened by the thickened and heavily cutinized walls of the epidermal cells and by the underlying single layer of sclerenchyma. Beneath this is the very per- fectly developed palisade parenchyma, extending from edge to edge of the leaf. The remainder of the mesophyll is com- posed of simple, unbranched parenchyma, enclosing many intercellular spaces, and bounded on the under side of the leaf by the single layer of fairly thick-walled epidermal cells. In contrast to this, fig. 2, a similar diagrammatic view of the cross-section of a larger white leaf shows a less convex upper, a less concave lower, surface, and the almost or quite complete absence of sclerenchyma cells except at the edges of the leaf (see fig. 9). The most striking difference in the structure of the two leaves, however, consists in the complete absence of palisade parenchyma from the white redwood leaf. The remainder of the mesophyll is com- posed of somewhat larger parenchyma cells than in the green leaves, and the intercellular spaces are also slightly larger. Examination of these cross-sections under higher magni- fication (figs. 7-10) reveals still more plainly the contrast between the white and the green leaves. Figure 7 is a detail from near the middle of the green leaf represented in fig. 1. Figure 7 shows the thick-walled epidermal, the thicker walled sclerenchyma, cells, and the regular palisade- parenchyma cells. These contain many chloroplastids, slightly larger than the starch-grains indicated in the figure. There are many of these starch-grains imbedded in the cytoplasm. Vacuoles are numerous, and evident. The cytoplasm and nucleus are sharply differentiated. Figure 8 represents the corner of fig. 1 cut off by the dotted line. In fig. 8 are shown the very strong sclerenchyma cells im- mediately underlying the epidermis at the edge of the leaf, and extending almost continuously along the under side to and around the resin-tube. This last is large and bounded by many suitably supported, thin-walled, glandular cells. 94 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. At the edges, as well as elsewhere throughout the palisade and mesophyll tissues of the green leaves, starch-grains occur in considerable numbers in all the cells. The amount of starch present undoubtedly varies at different seasons of the year; but since these green leaves were collected at the same time as the white ones, and near them, comparison of the starch content is justified. Figure 9 represents under higher magnification the part of fig. 2 cut off by the dotted line. As this shows, scleren- chyma cells are found at the edges of the white leaves, but they are not so firm as in the green leaves. The resin-tube is decidedly smaller and bounded by cells evidently less active than the glandular cells of the resin-tubes in the green leaves. There is no starch in any mesophyll cells of the white leaves. The cytoplasm and nucleus are not easily distinguishable from one another in most of the mesophyll cells. The cytoplasm presents a thoroughly disorganized or, as one may more truthfully say, a by no means organ- ized, appearance, neither vacuoles nor plastids being dis- cernible in most cells. The vacuoles occur in few cells only and are unlike those of green mesophyll cells. The plastids are variable. In sections of white redwood leaves from the summit near where the Redwood City—La Honda stage road crosses the first ridge between here and the sea, I have entirely failed to detect even rudiments of plastids or chromatophores. There are granules and gran- ular aggregations in the cells, as fig. 9 shows, but material carefully fixed in Flemming’s weaker mixture of chrom- osmic-acetic acids and stained by two-tenths per cent. acid fuchsine in distilled water* failed to exhibit any structures which I could positively identify as even rudimentary chro- matophores. On the other hand, the material from near Gilroy, treated in exactly the same way, contained chro- matophores which ranged in size from those about half as large as the average chloroplastids in the normal green leaves down to indistinguishable rudiments. * See Zimmermann-Huniphrey, Botanical Microtechnique, pp. 196, 202, etc. Bot.—VOL, II.] PETRCE—SEQUOIA SEMPERVIRENS. 95 In this connection I may state that the white redwood taken from near Gilroy and planted in my garden had at least one leaf which, after the little tree was transplanted and before it died, became pale green over half its surface on either side of the midrib. I have made no attempt as yet either to transplant, or to disconnect from their parents without transplanting, the white redwoods growing on the summit near the La Hondaroad. This I shall do presently. Obviously the white redwoods must turn green if they are to survive after being severed from the parent. Some white redwoods can do this more readily than others, the condi- tion of the chromatophores being one of the factors con- trolling this change. Why there should be these differences in the rudimentary condition of the chromatophores of white redwoods we can understand only after determining the reason for the production of any white leaves at all. Comparing white redwoods with cedars, which, in culti- vation and in nature, not infrequently produce white leaves or green leaves striped or otherwise variegated with white, we find the cell-structure as well as the general conditions for the nutrition of the plants quite unlike. ‘The mesophyll cells in the white parts of green variegated leaves, and of white leaves, contain less protoplasm (Frank, 1896) than do normal green cells, the cytoplasm forming a comparatively thin layer lining the cell-wall, the greater part of the cell- cavity being filled with the more than usually abundant cell-sap. Chromatophores, if visible at all, are colorless, small, and scarcely denser than the cytoplasm, or otherwise distinguishable from it, but they vary in this respect with the degree of whiteness of the leaves. The white redwoods are similar, the cells of the whitest containing no structures recognizable as chromatophores, while those leaves which contain visible chromatophores are not perfectly white. But between the unorganized though abundant protoplasm in the mesophyll cells of the white redwood, and the meagre but very definitely situated protoplasm of the white meso- phyll cells of cedar and similar plants, there is a great dif- ference. This we may perhaps account for thus. 96 CALIFORNIA ACADEMY OF SCIENCES. [PRoC. 3D SER. Cedars and other plants with white or white variegated leaves are independent, manufacturing in the leaves and other parts that are green the non-nitrogenous foods needed. Any reduction in the number of green leaves, or in the number of chlorophyll-containing-cells in the mesophyll, is a reduction in the capacity of the plant to manufacture food; and if all the leaves turn white the plant will die as soon as the food is consumed which was elaborated and stored while its leaves were green. The turning white, or the failure to become green, of the leaves or any part of the leaves of cedars, etc., is a variation neither useful nor permanent; it is really a morbid condition. White redwoods, such as I have seen and here describe, are not independent. They absorb from the still living underground parts of the parent tree the non-nitrogenous foods (starch, sugar, etc.) manufactured in its own green leaves and stored in its own underground parts. These stores of food are very great in old redwoods. When for any reason a sucker starts with none of its leaves green, it is exactly as well off, so long as the store of food in its par- ent lasts, as if its leaves were green and as if it could manu- facture food for itself. The activities and possibilities of the white sucker are not abruptly terminated by the exhaustion of its own very limited store of food. It can and does draw on its parent for much food. In the variegated cedar we have some leaves shirking their function as food-manufac- turing organs, either because they were defective from the time of their origin at the growing-point, or because they have developed this pathological condition subsequently. The white redwoods, on the contrary, are the vegetatively produced offspring of a wholly independent organism which live as parasites. They take on some of the characters of parasites, as is shown by the absence of palisade cells in the leaves, and by the rudimentary condition of the chromato- phores and other protoplasmic contents of the mesophyll cells. They are also less vigorous and grow less rap- idly than wholly independent though similarly produced individuals. BotT.—VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. 97 Other differences in structure between the white and the green leaves of redwood may be mentioned. As shown by figs. 11, 12 and 13, the walls of the epidermal cells of the white are not as thick as those of the green leaves. Figure 11 shows the epidermal cells around and above the some- what depressed stoma on a white leaf. Figure 13 is a similar view of a part of the surface of a green leaf. Both figures are of a stoma from the upper surface. Though in the green and in the white the mouth of the stoma is about equal in size, the adjacent epidermal cells are smaller as well as thin- ner walled in the white redwood. Figure 12 represents the guard-cells of the stoma, only the upper part of which is shown in fig. 11. There are no chromatophores in the guard-cells, but the nuclei are well differentiated. Though the numbers of stomata on the under side of the green and the white leaves are about equal, there are more stomata on the upper surface of the white leaves than of the green. Figures 3 and 4 represent very diagrammatically the shape and size, but exactly the numbers, of the stomata in equal areas of epidermis from the upper side near the midrib of a green and of a white leaf from redwoods in the Santa Cruz Mountains. Figure 5 indicates the number of stomata in an equal length of epidermis similarly situated but from a green leaf from one of the redwoods in the Arboretum of Stanford Uniyersity. Figure 6 is another strip from a white redwood leaf from the mountains. Two facts are demonstrated by these figures: first, that the white leaves always have more stomata on the upper side than do the green ones; second, that in the green leaves the number of stomata on the upper side of the leaf is proportioned to the humidity of the region in which the tree grows. The sec- ond is a fact well known and understood; the first is new and not easy to understand. In all probability it will be found that the white redwoods occur where transpiration is not so great asin many places in the mountains where green redwoods occur. Ina thicket there would evidently be less rapid transpiration than in the open at one side or above the thicket. The white redwood suckers which I have seen are 98 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER, certainly in a situation in which transpiration can not be very rapid while the suckers are actively growing. We come now to consider what causes the suckers of certain redwoods to be white while others are green. It is evident that the white redwoods are not white from etiola- tion, for other suckers similarly situated, and other plants all about, are as green as usual. Nor can lack of iron be the cause, for the same reason. So far as I can see, the only reason for these plants being white is that the leaves form, and attain nearly or quite their full size, at a season when there is insufficient warmth for the formation of chromatophores and chlorophyll pigment, though enough for growth. This is in perfect harmony with Sachs’s obser- vation (1864), since extended by Frank (1895), that seed- lings growing and buds unfolding at low temperatures produce leaves yellowish or white, either wholly or in patches. I had occasion to notice this phenomenon par- ticularly during the past winter in the leaves of Bur Clover (Medicago denticulata Willd). In December and January there were warmth, moisture, and light enough for a lush vegetation composed of the common annuals, but the nights were chilly and the days not warm. There was an unusual amount of variegation in the leaves of the common weeds. That there was light enough for chlorophyll formation is evident from the fact that Bur Clovers growing in the labo- ratory had no white or variegated leaves. As Sachs proved by experiment, the plant must have a certain minimum amount of warmth in order to form chlorophyll. This min- imum, higher than the minimum for growth, of course varies with the species. The white redwoods growing on the crest near the La Honda road are killed down to the ground each year by the frost. This is evidence of considerable cold. In January of this year (1900), when I visited these white redwoods, they had been frosted down but, buried in the leaf-mould covering the branch of the old stump from which the white redwoods spring, were many buds, healthy, with well formed but perfectly white leaves. On that summit no temperature Bot.— VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS, 99 records are kept. I therefore cannot tell the temperatures prevailing when these buds were forming in darkness under- ground. The darkness is of no significance, for all suckers begin in darkness and ordinarily their leaves are then green. Insufficient warmth seems, then, to be the reason why the chromatophores and the chlorophyll pigments do not form in the cells of these growing leaves. Frank (1895) says that it frequently happens that plants with leaves white because of cold at the time of their form- ation often retain these white leaves.even into the summer, subsequent warmth being insufficient to stimulate to chro- matophore and chlorophyll formation. On all such plants, however, the leaves developed later than the white ones, and when the temperature is higher, are green. If this were not the case, the plants would die. In the white redwood we have a different state of affairs. Although the first leaves borne on a shoot of one year’s growth may all have been formed in the cold, late in the previous year, and therefore may not be able to turn green, the leaves later formed, and the internodes forming or at least elongating later, when there is sufficient warmth, one would expect to find green. On the contrary, once started as white redwoods, the suckers continue white as to leaves and young cortex for an indefinite time. The differences in the color, and in the condition of the chromatophores between the white redwood leaves from near the La Honda road, and those from near Gilroy, may be accounted for thus. It is probable that near Gilroy, at least in the spot where the white redwoods grow, the tem- perature is not so low as on the exposed summit crossed by the La Honda road. While the temperature is low enough to prevent chlorophyll formation, it is not low enough com- pletely to suppress the formation of chromatophores, and by no means low enough to interfere with growth. Slight variations in low temperatures at the times when the buds, from which suckers spring, are forming, might permit the formation of green buds, of yellowish buds with rudimentary chromatophores, and of white buds with no chromatophores (2) March 18, rgor. I0O CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. at all. Certain it is that all the white redwoods I have seen or heard of grow where the temperature is low in autumn and winter. Ill. Tue SIGNIFICANCE OF THE WHITE REDWoOODS IN CONNECTION WITH OUR CONCEPTIONS OF PARASITISM AND OF HEREDITY. White redwoods are wholly dependent, absolutely para- sitic, plants which are in their first generation. They are not the offspring of other white redwoods, they are not the descendants of a long line of more and more dependent, more and more degenerate, organisms. Their parasitic characters have been acquired, or developed, during the brief course of their own existence, but they possess some parasitic characters not yet acquired by plants which have been semiparasitic for no one knows how long. Phora- dendron, Viscum, and the other ‘‘green parasites’’ have long lived at the expense of the other plants upon which they grow; but though attached to their hosts, these para- sites manufacture in their own green leaves their own non- nitrogenous foods. As I have shown elsewhere (1893), the ‘‘green parasites”’ which have been studied differ from completely parasitic flowering plants (e. g. Cuscuta, Brugmansia, Rafflesia) in the completeness of the connection between the tissues of host and parasite effected by the haustoria. In the com- plete parasites, xylem and phloém of the parasite are directly connected with the xylem and phloém of the host by means of xylem and phloém tissues which are continuous through- out the haustoria. In the ‘‘green parasites,’’ on the other hand, only the xylems of host and parasite are directly con- nected. This anatomical difference may be considered the reason for the difference in the degree of parasitism in these two sets of plants, or we may conceive that, so long as the parasite remains green, and therefore able to manu- facture its own food, a complete connection with both sets of conducting tissues in its host is unnecessary and unformed. There is at present no means to decide which Bot.—VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. IOI is true. In the first place, we do not know the actual func- tions, and cannot determine the entire significance, of the phloém tissues in higher plants. In the second place, so long as Trelease (1894), or any one else, can say without proof of error, that Leztneria floridana, a tree, contains no sieve-plates and inferably no sieve-tubes, the elements of phloém commonly regarded as most essential, it is impossi- ble to conclude that sieve-tubes are indispensable or that a phloém as well as xylem connection between host and para- site is essential for complete parasitism. Until the chemical physiology, and not the anatomy only, of the relation exist- ing between parasite and host in Viscum, Phoradendron, Cuscuta is worked out, it cannot be known how significant and important are the tissue connections effected by the haustoria. As to the significance of these tissue connections, the conditions presented by the white redwood may furnish some idea. The dependent white redwoods are branches of independent parents and are therefore connected from their beginnings, xylem with xylem, phloém with phloém, and parenchyma with parenchyma, with their parents. By means of these connections the adequate supply of foods, as well as of food-materials and water, by the parent to its offspring, from old redwood host to parasitic sucker, is assured from the first. On the other hand, a parasite attacking the tree from the outside must establish these connections. It may establish them only imperfectly, as in Viscum, or completely, as in Cuscuta. The white redwood, with its perfect connection with the parent, offers the coun- terpart of the condition which accompanies complete para- sitism, and though the leaves persist as such, they are struc- turally no longer perfect leaves, and physiologically only partly so. Because of its perfect connection with the host, the white redwood is able immediately to develop some feat- ures of the characteristic structure of parasites. This is especially interesting because the white redwoods are the vegetatively produced offspring of independent plants, themselves the descendants of generations of inde- pendent plants. The suckers of redwoods inherit the 102 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. tendency, which one would expect to find firmly fixed, to develop into independent plants, green and manufacturing their own foods. But these young suckers, originating when the weather is too cool for chlorophyll and chromatophore formation, though warm enough for a certain amount of growth, beginning with white leaves and growing well enough during the season without manufacturing food for themselves, form no chlorophyll even in the leaves and internodes later developed. The effect of low temperatures when the first leaves were forming was either upon the protoplasm itself, preventing its forming chromatophores and chlorophyll, or upon the chemical processes by which these organs and substances are produced. One naturally as- sumes the former—that the powers of the protoplasm are lessened by low temperatures. In these first leaves, the protoplasm is prevented by the cold from following its in- herited tendency to produce chromatophores and chloro- phyll. In the leaves later formed, the inherited tendency to form chromatophores and chlorophyll is not interfered with by cold, but it does not cause these leaves to become green. They do not need to be green; the plant obtains food enough without turning green and manufacturing its own food. The inherited tendency is not aroused by hun- ger into action. The stimulus needed to set it in operation not being given, the inherited tendency remains dormant as long as the white suckers remain connected with the parent. The parasitic habit forced upon the young white sucker by its inability to manufacture its own food, and the parasitic characters assumed by the young white sucker, are continued as the plant grows. Continued healthy existence in spite of inability to manufacture food induces in new leaves and cortex those characters found in the earlier and older ones. Environment, the influence of certain stimuli, induce a re- action ordinarily characteristic of species of plants which have been parasitic for generations. The white redwood serves as an index of the relative powers of heredity and of environment, or, more definitely, of heredity and of the influence of, and the power of Bot.—VOL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. 103 reaction to, certain stimuli. There being no need to man- ufacture food, the food-manufacturing apparatus is not formed, a parasitic habit being successful so far as the individual is concerned, the inherited habit is not entered upon. That the need to manufacture food would have an effect upon the development of the white suckers is indicated by the behavior of the white sucker which grew near Gilroy and which I planted in my garden where it could obtain little if any organic matter as food. It died soon after transplanting, but ot untzl one leaf had become pale green. The effect of cutting white suckers away from the parent stock and from their supply of manufactured food I shall test presently by experiment on the redwoods near the La Honda road; but this experience is not without significance. At least it strengthens my contention that inherited tendency is less strong than environment, and that in some cases, at least, inherited tendency must be called into action by some specific stimulus or combination of stimuli operating upon the plant from outside itself. In our white redwoods, the descendants of an exceedingly ancient race of trees in which heredity should be proportionally strong, we have a certain amount of evidence that the irritability and the power of response of the organism to external influences are stronger than its heredity. May not this always be the case? May it not be that what we call heredity is really the response to similar stimuli and combinations of stimuli occurring in orderly succession in the course of nature? STANFORD UNIVERSITY, CALIFORNIA, June, 1900. Bot.—VoL. II.] PEIRCE—SEQUOIA SEMPERVIRENS. 105 1895. 1896. 1899. 1898. 1893. 1864. 1894. 1893. BIBLIOGRAPHY. Frank, A. B. Krankheiten der Pflanzen. Bd. I. Breslau. The same, Bd. III. GANNETT, HENRY. The Redwood Forests of the Pacific Coast. Nat. Geograph. Magazine, Vol. X. GOEBEL, Kar_. Organographie der Pflanzen. Bd. I. Jena. PEIRCE, GEORGE J. On the Structure of the Haustoria of Some Phanerogamic Parasites. Annals of Botany, Vol. VII. SACHS, JULIUS von. Ueber den Einfluss der Temperatur auf das Ergriinen der Blatter. Flora, 1864. Reprinted 1892, in Gesamm, Abhandlungen, Bd. I. TRELEASE, WILLIAM. Leitneria floridana. Siath Ann. Report Missouri Bot. Garden. ZIMMERMANN, A.- Botanical Microtechnique. Translated by J. E. Humphrey. New York. 106 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Io. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XIV. All figures were drawn with an Abbé camera lucida. Cross-section of green redwood leaf, showing position of vascular bundle and resin-tubes, the well developed palisade parenchyma and sclerenchyma. X 30. Diagrammatic. Cross-section of a larger white redwood leaf, showing the absence of palisade parenchyma and sclerenchyma. X 30. Dia- grammatic. Strip of epidermis from upper side of green redwood leaf, showing distribution of stomata near midrib. X 60. Similar strip from white redwood leaf, showing distribution of stomata in an equal area. %X 60. Figures 3 and 4 are from mountain redwoods. Similar strip from green redwood leaf, showing number and dis- tribution of stomata. X 60. Similar strip from white redwood leaf, showing the same. X 60. Figure 5 is from redwood growing in Arboretum of Stanford University, fig. 6 from mountain redwood. Part of green leaf shown in fig. 1, from above vascular bundle, showing palisade parenchyma, sclerenchyma, starch-grains, and other contents of palisade cells. X 300. Part of figure cut off by dotted line, showing starch-grains and other contents of mesophyll cells, thick-walled sclerenchyma and epidermis at edge of leaf, strengthening cells around resin-tube, GiGi B00: Corresponding part of fig. 2, a white redwood leaf, showing unor- ganized contents of mesophyll cells, lighter strengthening tis- sues, etc. %X 300. Showing structure of stoma of white redwood and absence of chlorophyll and starch-grains. X 300. Figs. 11 and 12. Two views from the surface of stoma from white redwood leaf; fig. 11 from above, showing auxiliary cells (Vebenzellen), fig. 12 from further down, showing guard cells. Stoma closed. X 350. Fig. 13. Surface view of epidermis and stoma from green redwood leaf, showing larger and more vigorous epidermal cells, and decidedly thicker walls of epidermal cells. X 350. — PRoc.CAL.ACAD. Scr.3G” SER Bor. VoL Il / ey f t @ & / 2S Sr D Seat I Ye / oe * SEER Scer | va a l | ee = ca} : pee at as= — . oo ES SS nes r—) — — —> S = — = => —_> — => DS = a SS aS a Ee = o Pe = ad = Sa — oOo — Dd o = = => = 7 = > ao <= = cd Gi cer, FARTS OF LEAVES OF SEQU [Petce] PLATE XIV PHOTO -LITH. BRITTON & REY, SF. » Loner MPERVIRENS LAZZ ; - ¥ wa SCIENCES fet ae aN é ¢ i ¥ S L HED BY THE ns Ree Sanat I piceiatee Rais ea AS i Ao Nh . 5 ale bs ies NS wh ape La } , i i TION COMMITTEE We ¥ Enel 5 f’ yy ( ~ “ ea ah WN ee Cuartes H. Gitpert, Cha PPA hy ie Leta on mA : f ake Bea 7 me ’ 5 hi i ‘ cat A sich ‘ S 44a ORS OF BOTANICAL PUBLICATIONS Sot MOURN NI: Oe beh Rr SN ae ad SercHELL TOR ‘% BE ora due ace “iy PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES TuIRD SERIES BoTANy Vous Hy Nok'4 ie A Revision of the Genus sg Calochortus BY CARL. vEpy a _ Issued December 14, 1901 SAN FRANCISCO PUBLISHED BY THE ACADEMY . I90l 1 a ‘ \ Pa gat Re W te ay Pal etnas 2 at . es 1A APL TD Mia abe © SPR AN Ic +e mY A REVISION OF THE GENUS CALOCHORTUS. BY ‘CARL, BURDY- PLATES XV-XIX. INTRODUCTION. THE most widely diffused as well as the handsomest of the liliaceous plants of the Pacific Coast are the Calochorti. On the north they reach British America; one species is to be found as far east as Nebraska; several are natives of northern Mexico; and within these limits no considerable section of country is destitute of some species. While the range of the genus is so immense, that of sev- eral of the species is also very extensive. What a diversity of conditions C’. uzttalliz meets in its range from the west- ern side of the Sierra Nevada to Nebraska, and from the Snake River to Arizona. C. uztidus is found from the meadows of eastern Oregon to the shores of Yellowstone Lake, C. albus from San Diego to Tehama, and many others are scattered over hundreds of miles. A species distributed over a region so varied in soil, climate, and alti- tude cannot but be variable; and it hardly need be added that the genus Calochortus is a very difficult one for botan- ists to deal with. I long since became convinced that it is only in the gar- den, where plants from different localities can be grown under identical conditions, that the relationship between apparently different forms can be satisfactorily determined. For some years I have grown a large variety of Calochorti in my grounds, and have had nearly every known species under cultivation, often in many forms. The culture of Calochorti is most interesting, though not unattended with cultural difficulties; but the beautiful flowers amply repay all efforts, and the garden has proved the identity of forms [ 107 J November 27, Igor. 108 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. apparently different; here, also, variations attributed to environment are shown to be constant. In the garden, too, strains, which from a botanist’s standpoint seem scarcely distinguishable, show marked differences in vigor, flower- ing time, or immunity from disease. It is a peculiarity of our liliaceous plants, that asa rule in a given locality there is little variation from a well marked type, as little, indeed, as may be found between flowers growing upon the same plant. Hundreds and thousands of flowers may be picked, all conforming closely to this type. In another locality, the same species will be found marked- ly different. The difference between the forms in the two localities may be slight, consisting merely of a marking or a slightly varying leaf, habit, or gland; yet the variant, once noted, is found to be constant. In Calochortus color forms are frequent, the flowers from one bulb retaining the same tints under any and all conditions. The difference between forms from different localities is rather that which florists designate by the word ‘‘strain’’ than what is usually under- stood to constitute a botanical species or variety. In cultivation it has frequently been found that a very slight variability in strains is accompanied by a marked constitutional difference. In two beds of Calochortus venus- tus, planted in the same soil, and separated only by a thin board, it would puzzle a botanist to state wherein the plants vary. They come from widely separated localities, and the difference is one more easily detected by the eye than con- veyed by words. In one bed, two-thirds of the leaves are already destroyed by mildew (otrytzs), while in the other, not one leaf is injured; and such is the case whenever and wherever the two are planted. Many similar instances occur in other species, but a single one is sufficient to show that the slight variations which the eye detects are not the only ones. Such strains are present in nearly every species of Ca/lo- chortus. The range of a strain may be very local—a few miles square—or it may be found over half the length of a state. In Calochortus venustus one strain runs through all Bot.—VOL. II.] PURDY— CALOCHORTUS. 10g the plants found for hundreds of miles along the Sierra; another strain is found in the same species occurring in the Coast Range and over an equal area. In some of the more variable species there are several strains. In many of the Calochorti the gradations from one species to another are so slight thatit is impossible to separate them. The extreme types on which the species are founded are easily distinguishable, but a perfect chain of variations links them closely together. There is no doubt that C. weedzz, C. plummere, and C’. obtspoénszs are variations of a greater species. While, as before stated, it is the rule that a given locality produces specimens conforming closely to a type, yet this is not always the case. In some localities the variations are bewilderingly numerous. I have seen places where hun- dreds of flowers of C. venustus could have been selected, each differing in color and markings from the rest. Why a species that remains so true to a type in some localities should vary so remarkably in others is a subject that will not be discussed at present. Hybridization will account for it in some instances, while in others it is hardly a tenable hypothesis. I cannot say that I attribute any material share in the origin of the many strains or varieties to hybridization, although among the Calochorti it is not infrequent. Such crosses as C. albus and C. benthamz, C. maweanus and C. pulchellus are frequently met with, but I have never yet seen one that was fertile. Again, varieties of a species, e. g., C’. luteus var. oculatus and C. luteus var. cttrinus, readily cross and produce fertile hybrids. Over a small area in- numerable cross-breeds may be found, but afew miles away the two varieties will be found separate and varying as little as in any locality. Then again, hybridization often will not take place between two apparently very closely related species. I have often seen C. vesta in flower surrounded by large numbers of C. /uteus var. oculatus or var. citrinus, but not a plant could be found that in any way indicated hybridization; while last summer, in the Sierra Nevada, I IIO CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. visited a spot where C. /uteus var. oculatus and C. luteus var. citrinus had hybridized to a remarkable degree. A few yards away,indeed mingling with them, were thousands of C. venustus. Within a quarter of a mile I do not doubt there were fifty thousand plants in flower, yet close search failed to reveal one that in any way suggested a cross of C. venustus with any of the variations of C. Zuteus. In the ‘‘ Botany of California,’’ Vol. II, published in 1880, Watson described twenty-six species of Calochortus. The work was carefully done, though the material at com- mand was meager as compared with that obtainable now; yet nearly every species recognized by Watson stands good to-day. Many new species have been added, but by the exploration of new territory rather than by the subdivision of old species. The work of preparing the present paper has been facili- tated by the courtesy of the California Academy of Sciences and the University of California, in allowing me to inspect their herbarium specimens. Mr. J. W. Congdon of Mari- posa very courteously permitted an examination of his material, and to Mr. G. W. Hansen I am indebted for a set of specimens from Amador County. My personal collec- tion, including both herbarium and living specimens, covers a wide range; still, with these facilities, probably as good as can be obtained anywhere, the material is painfully unsatis- factory in some species, several of which are represented in the best herbariums by a single specimen, if at all. With each year appear many new forms, even from Cali- fornia. Last season brought three new species, and many striking variations of old species were added to the already large assortment. The field is immense and has never been properly worked over. In view of these facts, it seems the wisest course to disturb existing nomenclature as little as pos- sible. As to whether a given degree of difference warrants a specific or a varietal name seems to me to be very largely a matter of personal opinion. While one can hardly agree with the author who designates a color form by a specific name, it will probably be consulting the convenience of Bot.—VoL. II.] PURDYV— CALOCHORTUS. Vag at botanists to allow such a name to stand for the present, especially as a more extensive knowledge of the subject may result in still further changes. The pressing need is for a work containing descriptions of all known species of Calochorti, together with such grouping as will readily con- vey to the student the relationship of the various species. In the ‘‘ Botany of California,’’ the types of the species known to Watson are usually very accurately described. The only criticism to be made is that in many instances he was acquainted with but afew representatives of the species. Nineteen years have added much to our knowledge of the range of the various forms of Calochorti, but it is still far from complete. In the notes on distribution of species, the range as I have accurate knowledge of it is given. A species may, and in many cases doubtlessly does, extend over a far wider range than that with which it is credited. The measurements of any portion of a plant as given in published descriptions of Calochorti are of little value, and are apt to be misleading. Environment makes the greatest difference in the size of the plants. Take, for instance, those of the woods, such as C. albus, C. pulchellus, and the elegans group; the variations are almost limitless. Especially after a forest fire is growth luxuriant. Plants which under adverse conditions have leaves but a few inches in length, and few-flowered, slender stems, will, under more favorable circumstances, produce great leaves a foot or two long, stout stems eight inches to two feet in height, and a dozen or more fine flowers. Especially do the plants of the Mariposa sec- tion respond prolifically under fertile conditions. Ifthe season be dry, the plants are sparsely scattered and but a few inches above ground; but let the season be one of great rainfall, they fairly hide the ground with tall, many flowered stems upholding numerous large blossoms. But while measurements based upon a series of specimens are almost valueless, proportional measurements of the parts of the same flower are often of great importance, the proportions between the parts being usually the same whatever the size of the flowers. I12 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. In the present paper I have departed from the usual method by describing a single type plant and indicating the variations in the notes. Personal experience has proven that the fuller a description is, the more value it has in the determination of specimens; details seemingly of little im- portance at the time when the specimen was described, are often indispensable in the work of later students, making it an absolute necessity for them to refer to the type specimen in order to determine what was really described. This brevity in some of the earlier descriptions makes them entirely valueless for determination. Having no personal knowledge of the Mexican species outside of the herbarium, I have not included them in this revision, being unable to add anything of value. There is a large and almost unknown field in Arizona and western New Mexico which will probably yield several new species. Only the description and when possible the locality (the original locality is quoted in nearly every instance) of the species have been given in the following pages, the synonymy having been omitted because of its often doubtful nature. KEY TO THE SPECIES OF CALOCHORTUS. Section |. ucalochortus. Flowers or fruit nodding; petals incurved or strongly arched; gland trans- versely crested or hairy; capsule nodding, with thin acute or winged cells; leaves long and glossy, not channeled. Group 1. GLOBE TULIPS. Type of Group C. albus. Flowers subglobose, nodding. Woodland plants; California. Flowers white; petals covered with scattered silky hairs within. 1, C. albus. Flowers rose color; petals silky within, partly opening out. Foot- hills of Fresno and Tulare counties, California..2. C. amcenus. Flowers light yellow; petals silky within, gland bordered with stiff hairs which cross each ofher...>.:.-..-....+.. 3. C. pulchellus. Petals very strongly inarched, not silky within, but margin thickly set with short, stiff hairs; gland like last........ 4. C. amabilis. Bot.—VOL. II.] PURDY — CALOCHORTUS. II3 Group 2. STAR TULIPS. Type of Group C. elegans. Flowers campanulate, erect or ascending; capsule nodding (except in No. 13); stem low and flexuous (In 14, 15, 16, stout and erect), not bulbiferous or very seldom so. * Petals covered with hairs, and with a transverse scale covering upper part of the gland. Woodland plants. Flowers yellow. Foothills of the Sierra Nevada...5. C. benthamt. Flowers white or purplish blue, covered with long erect hairs; cap- sule oblong-elliptical; stem branching........ 6. C. maweanus. 6a. C. maweanus var. major. 66. C. maweanus var. roseus. Flowers blue, covered with silky hairs, longer and slenderer than the last; capsule orbicular; inflorescence umbellate. 7. C.ca@ruleus. Flowers greenish white; petals with very narrow scale and covered with long hairs. Oregon and north..8. C. elegans. 8a. C. elegans var. nanus. Flowers yellow, green tinged; petals strongly inarched and pit deeply ‘set: Mt. Jefferson, Oregon)..........2....45 9. C. lobbit. ** Petals with a transverse scale closely appressed over upper portion of gland, nude or nearly so. Woodland plants; in dry soil. Flowers white with a single tuft of a few hairs at each end of scale on petals; plants very low and slender. Sierra Nevada. 10. C. nmudus. Flowers white with scant hairs on lower third; plants taller than the last. Vicinity of San Francisco Bay........ 11. C. umébellatus. *** Petals nude or only lower portion hairy; flowers campanulate; plants growing in open wet meadows. Flowers lilac, hairy on lower third; one or several bulblets on stem DEO ey Tlie SONTAG 6 4.32 Siene wie 5 4 dom owes oc esermere 12. C. unifiorus. Flowers white, not bulbiferous; capsules erect...13. C. shastensis. **** Petals covered with silky hairs; flowers and stems stout and erect. Closely related to Group 1 of Mariposa. Plants growing in open fields or hillsides. Mt. Shasta, California, and north. Flowers blue; petals without scale, covered with long silky hairs. Mt. Shasta, California (?), and Willamette Valley, Oregon. 14. C. tolmiet. Flowers white; petals with scale, otherwise the same as the last. Willamette Valley, Oregon ......62..0 foc sees 15. C. purdyt. Flowers straw color; petals without scale, otherwise like C. tolmieit. Lake Pend d. Oreille, Idaho ...... 16. C. apiculatus. II4 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. SecTIon II. Mariposa. Flowers open-campanulate; gland usually densely hairy; capsule and pedicels erect. Group I. OREGON MARIPOSAS. Type of Group C. xitidus. Capsule as in Section I, but erect; leaf (as in Section I) long and glossy, not channeled. Oregon and northeast. * Petals with an indigo blotch in the center. Flowers large, white to lavender ...............+-. 17. C. mtidus. **Petals not spotted in the middle; flowers lilac, smaller than in the preceding. Flowers strongly arched and barred with yellow..... 18 C. greenet. Flowers less arched; stem bracted midway...... 19. C. pavonaceus. Similar to last; stem not often bracted midway. 20. C.longebarbatus. Flowers white, densely hairy above gland.......... 21. C. howellit. Group 2. Rocky MountAIN MARIPOSA. Type of Group C. gunnisoni. Gland transverse and narrow; leaf usually as in Section I. East of. the Rocky Mountains... 2: 3.vscesneae ss 22. C. gunnisonti. Group 3. WEED’s MarRIPosa. Type of Group C. weedii. Petals covered with slender hairs; capsule narrowly oblong, with thick, obtusely angled cells; radical leaf as in Section I, solitary, long, shining, and not channeled; bulb heavily coated with coarse black fiber. Flowers orange or rarely pink or white.............. 23. C. weedit. Flowers purple...23@. C. weedii var. purpurascens (C. plummere). Petals brownish, short, truncate, not equaling sepals. 230. C. weedii var. vestus. 23c. C. weedii var. obispoénsis. Group 4. GOLDEN BowL Mariposas. Type of Group C. clavatus. Petals yellow, lower half covered with clavate hairs; radical leaves linear and deeply channeled. Stem stifly zigzag; 22.052 Seeppeenetep eriele nes wis oie 24. C. clavatus. Stem not zigzag. Doubtfully placed in this group, but has no cla- vate hairs...25. C. concolor, sp. nov. (C. luteus var. concolor ). BotT.—VOL. II.] PURDY — CALOCHORTUS. II5 Group 5. Type of Group C. kennedyt. Petals nearly naked; gland round, small, and densely hairy with matted hairs; leaves ashy blue, linear, deeply channeled. Desert plants. Petals vermilion or Orange: ..1.2s0.5454 2s heee soos 26. C. kennedy. Flowers clear yellow ; petals densely hairy below; capsule narrowly OUIGHE Sewer < Dak webu hho octane Ree er aaa Br. a GUECUS. Group 6. BuTrerFLy TULIPS. Type of Group C. venustus. Petals slightly hairy below, usually oculated and brilliantly colored; gland prominent, round or lunate; leaves linear, channeled. California. Flowers yellow; petals not oculated; gland lunate; capsule attenuate from a broad base; plant dwarfed............... 28. C. luteus. Flowers yellow or lemon, otherwise same as in var. oculatus. 28a. C. luteus var. citrinus. Flowers white, yellow or lilac; petals oculated, gland lunate. 286. C. luteus var. oculatus. 28c. C. luteus var. robusta. Flowers lilac or white; gland narrow, doubly lunate...29. C. vesta. Flowers white, cream, lilac, purple, red or pink; petals oculated, in some varieties with a red blotch above eye; gland round; capsule linears 20.025). as00 30. C. venustus. 30a. C. venustus var. roseus. 306. C. venustus var. eldorado. 30c. C. venustus var. purpurascens. 30d. C. venustus var. sulphureus. Group 7. Litac MArRIPposAs. Type of Group C. splendens. Petals white, lilac, or purplish, not oculated, more or less hairy; gland small, round, and densely hairy; leaves linear, channeled. Flowers lilac-purple; petals hairy on lower third. 31. C. splendens. Flowers lilac to salmon, short yellow hairs on lower third of petal; plants low and slender....... 31a. C. splendens var. montanus. Flowers large; petals pale lilac with cobwebby hairs on middle CRINGE tetera artes reicteraiel a siaiale store’ 316. C. splendens var. major. Much larger and stronger than type; hairy on lower third of petal. 31c. C. splendens var. rubra, Flowers white; gland ill defined; plants more slender than last. AES A MIERCTD Gna sas 0 p.ne ¢)<10 0 sen aseehboun 32. C. palmer. Flowers white or lilac with dark maroon spot at base of petal; cap- sule oblong. Resembles C. splendens........ 33. C. cataline. Flowers smoky white; stems stout and umbellate. 34. C. invenustus. Flowers similar to last; petals deeply pitted...... 35. C. excavatus. 116 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Flowers purple; sepals obtuse; stem flexuous and weak, almost CRESDING. 5. oi Sues mans acme ix editegy ste ioe beeen 36. C. flexuosus. Flowers white. Resembles C. splendens........... 37. C. dunnit. Group 8. GREEN BANDED MARIPOSA. Type of Group C. macrocarpus. Petals purplish lilac, with a greenish line down the back, obovate- acuminate. Stems stout and rigid; leaves linear and deeply channeled. North- eastern California to eastern Washington and southern Idaho. 38. C. macrocarpus. Group 9. SEGo LILIEs. Type of Group C. nuttallii. Petals white, lilac, yellow, or pink; gland round; stem prominently bulbiferous at base, umbellate. Flowers as described above. Stout desert plants of the Great Basin and Sastwatdly 12.5.5. \\js..swammen aay waleespm ’< 39. C. nuttallit. Flowers smaller, usually white; anthers sagittate. Slender Alpine pants: Sierta Nevada. po. cerssinacwes seaess 40. C. leichtlinit. DESCRIPTION OF SPECIES. Calochortus. Perianth deciduous, of six distinct, more or less concave segments, the three outer (sepals) greenish and more or less sepaloid, the inner (petals) mostly broadly cuneate-obovate, usually with a conspicuous glandular pit near the base, and variously colored. Stamens six, on the base of the seg- ments, included; anthers linear to oblong, basifixed, dehiscent laterally. Ovary sessile, triquetrous and three-celled, many ovuled; stigmas sessile, recurved, persistent; capsule elliptical to lanceolate, membranous, three- angled or three-winged, mostly septicidally dehiscent; seeds numerous, two rows in each cell, somewhat flattened, with a thin membranous white or brownish, often loose, testa. Stems usually flexuous and branching from mem- branous or but fibrous-coated corms; leaves few, linear-lanceolate, radical and cauline, the latter alternate and clasping; all with many nerves and transverse veinlets. Flowers one to twenty, showy, terminal, paniculate or umbellate. The above generic description is in greater part that of Sereno Watson as given in the ‘‘Botany of California.”’ The genus is confined to western America, from Nebraska to the Pacific Ocean, and from northern Mexico to British America. Section I. ucalochortus. Flowers or fruit more or less nodding; petals strongly incurved or arched, with a broad, transversely crested or more or less hairy pit above the base; Bot.—VOL. II.] PURDY— CALOCHORTUS. si Thy sepals naked, rarely spotted; capsule elliptical or broadly oblong, deeply tri- quetrous and septicidal, the thin compressed lobes acute or winged; seeds ascending, close and pitted, the testa mostly brownish. Group 1. GLosBe TULIPs. Flowers subglobose, nodding. Woodland plants; California. ‘ 1. Calochortus albus Dozgl. Calochortus albus DouGLas in itt. Cyclobothra alba BENTHAM, Trans. Hort. Soc., N. S., Vol. I, p. 413, Pl. XIV, fig. 3; Bot. Register, Vol. XX, 1835, Tab. 1661. Stem stout, glaucous, usually branching, a foot or two high; radical leaves a foot or two long, 8-12 lines wide, lanceolate-acuminate; bracts large and foliaceous, I-5 inches long, 4-8 lines wide; sepals shorter than petals, Ovate-acuminate, greenish white; petals pure white, purplish at base, ovate- orbicular, acutish, 12-15 lines long, with scattering long silky hairs above the gland; gland lunate, shallow, with four transverse imbricated scales, fringed with close short yellow or white glandular hairs; anthers oblong- obtuse, mucronate; ovary attenuate above; capsule 1 or 2 inches long, 6-12 lines broad, abruptly short-beaked; seeds brown, pitted. The original specimens are in all probability from Mon- terey, as Douglas visited there, where the species is plentiful. C. albus is found in the Coast Range of California, from San Francisco Bay south to San Diego County, and in the Sierra Nevada Mountains, from Butte County south to San Diego County. There is quite a difference between the plants of the Coast Range and those of the Sierra Nevada. The flowers of the former are darker in color, often tinged with rose, and with petals thicker, more strongly inarched. The petals never open out sufficiently to show the inside of the flower, which after being in bloom a few days is half opened. Calochortus amenus Greene is really a color form of this Sierran form of Calochortus albus; but owing to the fact that the original locality of C’. a/dus is unknown, the writer hesi- tates to erect either the form from the Coast Range or that from the Sierra Nevada into a new species. Several variations of the form of the Coast Range have been described, some as species and some as varieties; but I fail to discover any characters by which they may readily be recognized. 118 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3p SER. 2. Calochortus amenus Greene. Calochortus amenus GREENE, Pittonia, Vol. II, 1890, p. 71. Similar to C. albus, but lower and more slender; petals a rosy pink with the gland rose-purple, scarcely at all arched, and opening in full bloom. Found in the higher foot-hill region of Fresno and Tulare counties, California. ‘«* Mountains east of Visalia, California.’’ 3. Calochortus pulchellus Dozg/. Calochortus pulchellus Douctas in litt. Cyclobothra pulchella BENTHAM, Trans. Hort. Soc., N. S., Vol. I, 1835, p. 415, Pl. XIV, fig. 1; Bot. Register, Vol. XX, 1835, Tab. 1662. Stem stout, glaucous, usually branching, 8 to 16 inches high; radical leaves a foot long, 6 to 12 lines wide, lanceolate-acuminate; bracts large and folia- ceous, 2 to 3 inches long on the same plant, 4 to 6 lines wide; sepals shorter than petals, ovate-acuminate, yellow tinged with brown on the back; petals canary-yellow, ovate with the base cuneate, obtuse at apex, 9-12 lines long, with scattering long silky hairs above the gland, and bordered with short stiff hairs; gland deep, protruding outwardly, bordered with stiff hairs which cross each other; anthers oblong-obtuse; ovary elliptical to elliptical-orbic- ular, abruptly short-beaked. The original specimens of this species were collected by Douglas prior to 1835. The exact locality is not given; but the only place in which the species has since been found is in the Mount Diablo region, a section which was easily accessible at the time of Douglas’ visit to California, and often visited by his Mexican-Californian hosts. Although in a region much visited by botanists since then, no specimens were to be found in any of the herba- riums of this State up to the year 1897, when C’. pulchellus was collected by Miss Alice Eastwood of the California Academy of Sciences. . The very name Calochortus pulchellus had been appropri- ated by another yellow-flowered species of the same group, which is described and named below as C. amabzlis, and which is clearly a distinct species. Following are the original descriptions of the species copied from the Botanical Register :— Bot.—VOL. II.] PURDY— CALOCHORTUS. 1m ge) Cyclobothra pulchella.—‘‘Umbellis 2-3 floris, pedunculis bracteis breviori- bus, floribus globosis, petalis ovatis obtusis serrulato-fimbriatis fovea valdé excavata extus callosd, sepalis ovato-lanceolatis acuminatis vix brevioribus.’’ Calochortus pulchellus.—‘‘Caulis erectus, teres, glaber, subcorymbosus, apice magis ramosus quam in precedente, et humilior. Folia plana, acumi- nata, minus glauca; superioribus brevioribus. Pedunculi bracteis foliaceis breviores, bini ternive. Flores globosi, minores quam in precedente, lutei. Sepala virescentia, viridi-striata, petalis paululum breviora, acutissima. Pet- ala ovata, barbata, fimbriata, basi glabra: fovea nectarifera pilis abscondita.”’ a 4. Calochortus amabilis, sp. nov. Stems stout, usually branching in pairs, 8 to 12 inches high, glaucous; radical leaves 10 inches long, 4 to 6 lines wide, lanceolate-acuminate, tinged with purple; bracts large and foliaceous, 2 to 3 inches long, 4 to 6 lines wide; sepals shorter than petals, ovate, shortly cuneate at base, sharply acuminate or even mucronate at apex, yellow tinged with brown on the back; petals clear yellow, ovate, with a short claw, obtuse at apex, naked but margined with a close row of short stiff hairs, very strongly inarched so that the tips of the petals overlap each other much like a child’s pin-wheel; gland very deep, projecting upwards and outwards like a knob, lined with short stiff hairs which cross each other; anthers oblong-obtuse; ovary elliptical, short-beaked. C. amabilis is found on the hills along the north side of San Francisco Bay, from the redwood belt to the Sacra- mento foot-hills, as far north as Burnt Ranch, Trinity County, California. The species has been distributed in large numbers among the flower-growers of the world as Calochortus pulchellus, which it resembles in habit and size. The latter is more closely allied to C. albus, having the same globular flower and petals silky-haired within. It is also of a much lighter shade of yellow, and never could be confused with C’. ama- bilis by anyone who had seen both. Fa, > Group 2. Star TUuLips. Flowers campanulate, erect or ascending; capsule nodding (except in No. 13); stem low and flexuous (in 14, 15, 16, stout and erect); not bulbiferous or very seldom so. *Petals covered with hairs, and with a transverse scale covering upper part of the gland. Woodland plants. 120 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 5. Calochortus benthami Aaker. Calochortus benthami BAKER, Journ. Linn. Soc. (Bot.), Vol. XIV, 1875, p. 304; also Bot. Mag., Pl. 6475. Stems several-flowered, very flexuous, branching, dividing into pairs 7 inches high; radical leaf linear-lanceolate, shining, 4 lines wide, exceeding stem; bracts ovate-lanceolate, long, acuminate, 2 to 4 inches long, 3 to 5 lines wide; sepals 8 to to lines long, narrowly ovate, mucronate; petals yellow, with a yellow claw, naked, obovate, rounded above, a little longer than sepals, the upper portion of the gland covered by a narrow crescentic scale which is densely bordered above with short yellow hairs, some of which are clavate; anthers lanceolate-acute, capsule nodding, nearly orbicular, 6 to 9 lines long. Found in the lower Yellow-Pine belt of the Sierra Nevada, from Mariposa to Butte counties. ‘¢ California.”’ The description is drawn from rather an extreme speci- men. The plants are often very slender, simple, and but three to four inches in height. The flowers vary but little. In some sections flowers with the claw dark red or nearly black are common (C’. wallacez Hort.). 6. Calochortus maweanus Lezchilin. Calochortus maweanus LEICHTL. ex BAKER, in Journ. Linn. Soc. (Bot.), Vol. XIV, 1875, p- 305; also Bot. Mag., Pl. 5976. Resembles the preceding; stem usually branching, very slender and flexu- ous, 3-Io inches high, with from a few to ten flowers; leaves glaucous, much exceeding stem, 2-6 lines wide; bracts lanceolate, narrow, Io lines or more long; sepals ovate-lanceolate, acute or acuminate; petals a little longer, broadly ovate, acute; gland covered above with a narrow transverse scale, immediately above the scale densely hairy, the entire surface thickly bearded with long, erect, white or bluish hairs; anthers lanceolate-acuminate, 2-3 lines long; capsule oblong-elliptic. This is the type which is found in the Coast Range, from San Francisco Bay, at least as far north as Trinity County, California, and western Oregon. ‘‘California.’’ 6a. C. maweanus var. major, var. nov. This variety is twice as large as the type, from which it differs in its much stronger habit and lighter color. BotT.—VOL. II.] PURDY— CALOCHORTUS. I2I Grows in the Yellow-Pine belt, Butte County, California. 66. C. maweanus var. roseus, var. nov. The flowers of var. roseus are tinged with rose; the bulb is distinctive, having a smooth, mahogany-colored coat. Its habitat is western Oregon. 7. Calochortus ceruleus Watson. Calochortus ceruleus WATSON, Proc. Amer. Acad., Vol. XIV, 1879, p. 263. In general resembling a small specimen of C. maweanus. The plants are very slender; leaves and bracts narrower, pedicels more slender; flowers almost always in an umbel, petals more rhombic in outline, claw more slen- der, scale broader and fringed, the remainder of petal densely covered with long slender silky hairs; anthers oblong-obtuse; capsule orbicular, not beaked, 6 lines long. Specimens of this species show but little variation. ‘‘California (in the Sierra Nevada, Placer to Plumas counties).”’ 8. Calochortus elegans Pursh. Calochortus elegans PursH, Fl. Am. Sept., Vol. I., 1816, p. 240. Scape very slender, 4-8 inches high; leaves lanceolate-acuminate, nar- row, exceeding scape; flowers one to four in umbel, bracts one-half length of pedicels, acuminate from a base 2 lines wide; sepals ovate-acute, greenish white without, lighter within, purplish at base; sepals obovate-obtuse, whitish or tinged slightly with green, with purple spot on claw, covered thickly with rather short soft hairs, which are white on upper and purple on lower por- tion, excepting that the margin and a band around upper portion of petal is naked; scale narrow, ascending, and deeply fringed, covering about one-third the width of claw; anthers long, acuminate; capsule elliptical, rounded at each end. The type specimens were collected by the Lewis and Clark expedition on the headwaters of the Kooskoosky in Idaho (?). The writer was never able to obtain specimens of this species until just as the present paper was going to press, when flowers which are unquestionably the true C. elegans were received from a collector in western Idaho, near Spokane. As the original description of the species is very brief, the fuller description, as given above, was drawn 122 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. from these specimens. There is much difference in general appearance between the species and var. zanus, and in my opinion the variety will eventually follow the other forms which were originally included under C. elegans, and be given specific rank. For the present, however, it is left under the varietal name. No more definite locality than western Idaho can be given for the type. Some Californian specimens have been referred to C. elegans but the writer has never seen any which come under the species and does not believe that either the type or any of its variations are found in California. The Californian plants referred to as C. elegans are C. nudus. 8a. C. elegans var. nanus Woop. (Proc. Phil. Acad., 1868, p. 168.) Dwarf and very slender, leaves very narrow; petals more hairy and ciliate, often acute or even acuminate.—C. lyallii Baker, Journ. Linn. Soc. (Bot.), Vol. XIV, 1875, p. 305. ‘‘High hills, Yreka [California]. Also on Mt. Hood [Oregon].”’ The description is that of Watson in the Botany of California. The variety is known to the writer as it grows on Mt. Hood and on Mt. Adams. Watson evidently had small specimens, as under favorable conditions it is quite as strong as any of the preceding. The color is a delicate cream. we g. Calochortus lobbii, sp. nov. Calochortus elegans var. lobbii BAKER, Journ. Linn. Soc. (Bot.), Vol. XIV, 1875, P- 305- Stem 3 to 5 inches high, not so slender as the preceding; leaf a little exceed- ing stem in length, 3 to 5 lines wide, lanceolate, abruptly acute; sepals ovate- lanceolate, acute, greenish with a dark spot below, 6 to 8 lines long; petals a little longer, white tinged with green, broadly rhombic-ovate, very deeply pitted, the pit showing as a prominent knob on back of petal; scale very narrow, deeply bordered with long, feathered, hairy fringes, and concealed in the recess of pit; above the scale there is a nectar-producing gland covered by a broad band of agglutinated feathered hairs, above this band lower half of petal densely hairy with silky hairs, upper half less densely hairy; fila- ments subulate; anthers oblong-acuminate, ending in a hook-like cusp; cap- sule narrowly beaked. Bot.—VOL. II.] PURDY— CALOCHORTUS. 123 So far found only on Mt. Jefferson, Oregon. “i Srecon.’; The above was identified for the writer by Baker. As the other varieties which Baker mentions in his ‘*Tulipez’’ have been erected into species, and this is more distinct in character than any of them, it seems proper to raise it to specific rank. — ** Petals with a transverse scale closely appressed over upper portion of gland, nude or nearly so. Woodland plants; in dry soit. v 10. Calochortus nudus Watson. Calochortus nudus Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 263. Calochortus elegans var. subclavatus BAKER, Journ. Linn. Soc. (Bot.), Vol. XIV, 1875, p. 305. Low and slender, scape 2 to 4 inches high, with a single leaf 3 to 6 inches long, 3 to 6 lines wide, light green, of even width for most of length, abruptly acute; flowers one or more, in all specimens examined in an umbel if more than one; sepals narrowly oblong-ovate, acute, shorter than petals; petals greenish white or lilac, greenish at base, obovate, somewhat acute, denticu- late above, 5 to 7 lines long, the same in width, entirely nude except for a tuft of two or three short stiff hairs at each extremity of the narrow, closely appressed scale which covers the upper margin of gland; anthers blue, oblong, two-thirds the length of the subulate filaments. Probably C. elegans var. subclavatus of Baker. On north sides of high mountains in the pine forests of the Sierra Nevada, from Tulare to Plumas counties; in loose dry soils. ‘‘California (in the Sierra Nevada, Yosemite Valley to Plumas County).”’ The type as described from Tulare County is white, but there seem to be variations tending to lilac, and in some sections a nude petal. This is the smallest flowered of all the Calochorti. v 11. Calochortus umbellatus Wood. Calochortus umbellatus Woop, Proc. Acad. Nat. Sci. Phila., 1868, p. 168. Cyclobothra elegans var. Torr. ?, Pac. R. R. Rept., 1856, Vol. IV, p. 146. Calochortus collinus LEMMON, Erythez, Vol. III, 1895, p. 49. Stem low and branching, 3-15 inches high, flexuous; radical leaf exceed- ing stem, narrow, 3-4 lines wide; bracts foliaceous, acuminate, 1-4 lines (2) December 3, 1901. 124 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. long, 3-4 lines broad at base; flowers five to ten; sepals greenish white, ovate-lanceolate, acute; petals white, broadly fan-shaped, denticulated, obtuse; scale triangular, ascending, appressed on upper portion of gland, which has many short white hairs just above it, remainder of petal nude; fil- aments slender; anthers 1 line long, oblong-obtuse; capsule short, oblong, 6 lines wide, 8 lines long. A native of California, found on the lower mountains and hills back of Oakland, Berkeley, and Mills Seminary, and on Mt. Tamalpais. ‘‘Oakland, California.’’ The species was long confused with C. wnzflorus, and was described as C’. collinus by J.G. Lemmon. Professor E. L. Greene called the writer’s attention to the earlier description of Wood. C. umbellatus can be confused only with C. uniflorus, from which its lack of bulblets and the situation in which it grows, as well as its color, easily distinguish it. It varies _but little. *** Petals nude or only lower portion hairy; flowers campanulate; plants growing in open wet meadows. 12. Calochortus uniflorus Hook. & Arn. Calochortus uniflorus Hoox. & ARN., Bot. Beech. Voy. Suppl., 1841, p. 398, Tab. XCIV. Stem low, flexuous, but often stout, usually branched, 4-8 inches high, with one to four bulblets below the surface; radical leaves broad, 4-6 lines wide, exceeding stem; bracts linear-lanceolate, long and conspicuous; flowers four to ten, in one to three umbels, on long flexuous pedicels 3-10 inches long; sepals ovate, lanceolate-acuminate, greenish lilac; petals cuneate, somewhat truncate, denticulate, 10-12 lines long, color lilac, often with a purple spot on each side of the scale, naked above, sparingly hairy imme- diately above the gland; gland shallow, not pitted, a narrow triangular scale appressed upward over upper center; filaments slender; anthers obovate- obtuse, lilac, 2 lines long, one-half length of filaments; capsule elliptical. Except in the spots on petal or sepal there are few color variations. From Monterey, California, northward in the Coast Range to Grant’s Pass, Oregon. Found in wet meadows. C. uniflorus was originally described by Hooker and Arn- ott from specimens collected on the Beechy Expedition. The specimens were without doubt collected either in the Bot.—VOL. II. ] PURDY — CALOCHORTUS. 125 vicinity of San Francisco or at Monterey. A good cut accompanies the description. C. lilacinus was first described by Kellogg from specimens collected near Calistoga, Napa County, California. Dr. Kellogg’s original description, together with a water color drawing, are in the Herbarium of the California Academy of Sciences. After carefully comparing all the data contained in the descriptions of C. uniflorus and C. lilacinus with both fresh and herbarium specimens from Monterey, Calistoga, and a number of other localities, the writer is thoroughly convinced that the two species are the same. The description of C. untflorus was published much earlier than that of C. /z/a- cinus; it is drawn from a fair specimen of the species. It is obvious that Dr. Watson confused C. /z/acinus with the quite different C. wmbellatus. The writer has seen C’. wuzflorus in meadows high in the mountains where the plants grew very low and slender and were only one- to three-flowered. In rich ground several bulblets are produced annually, and if left undisturbed large and dense masses are formed, sometimes hundreds of them to the square foot. In soft ground the stems are apt to run along close to the surface a few inches to a foot before coming through, and in these situations plants a foot high with pedicels ten inches long are not uncommon. v 13. Calochortus shastensis, sp. nov. Scape low, slender, 4 to 10 inches high, but unusually erect, with a single shining light green radical leaf 3 to 6 inches long, of almost uniform width (3 to 6 lines), but abruptly acute at apex; bracts lanceolate, 6 lines long; sepals long, ovate, acute and acuminate, greenish without, lighter within, purple spotted near base; petals white or lilac, broadly fan-shaped, somewhat truncated above, denticulate, naked except that some few specimens have a few hairs above the narrow, fringed, ascending scale which divides the gland; anthers linear, obtuse, slightly sagittate; capsule as in preceding but erect. Found in open moist meadows in the vicinity of Sissons, California, at the base of Mt. Shasta, and about springy places on the western flank of the mountain. C. shastensis has long been known and collected as C. nudus, which it closely resembles in flower but from which 126 CALIFORNIA ACADEMY OF SCIENCES, [PRoc. 3D SER. it is clearly distinguished by the erect capsule. Itis a curious fact that a species linking the small Calochorti of the wet lands with the C. zzt¢dus section should be found at the very point where the latter terminates its most southern extension. The true C. xudus, it will be noted, grows only on dry slopes in the Sierra Nevada, from Plumas County, Cali- fornia, southward. GIANT STAR TULIPS. **** Petals covered with silky hairs; flowers and stem stout and erect. Closely related to Group 1 of Mariposa. Plants growing in open fields or hill-sides. Mt. Shasta, California, and north. 14. Calochortus tolmiei Hook. & Arn. Calochortus tolmiet Hook. & ARN., Bot. Beech. Voy. Suppl., 1841, p. 398. Stem stout, erect, usually branched, 9 to 18 inches high; leaves 4 to 6 lines broad, not greatly exceeding the stem, bracts foliaceous; petals 9 to 15 lines Jong, very broadly obovate and scarcely acute, rather deeply pitted, covered and fringed with long purple and white hairs; gland without scale, but the upper circular edge with a dense fringe of reflexed hairs; anthers lanceolate and acuminate, 2 to 3 lines long; capsule broadly elliptical, acutish at each end, ro to 15 lines long. C. tolmiez is frequently confused with C. purdyz. The writer has seen specimens of the true C’. ¢o/mzez from the hills west of the Willamette River in Oregon. Dr. Watson also refers specimens from Mt. Shasta, California, and Mt. Adams, Washington, to this species. The original locality is ‘‘ Banks of the Walamet River.’’ 15. Calochortus purdyi Hastwood. PLATE XV. Calochortus purdyi Eastwoop, Proc. Cal. Acad. Sci., 3d Ser. (Bot.), Vol. I, p. 137, Pl. XI, figs. 8a-8/ Glabrous and glaucous; stem 2 to 3 dm. [18 to 16 inches] high, rather stout, erect, branching, two to many-flowered, not bulbiferous at base; radical leaf solitary, sheathing the stem, linear-lanceolate, acuminate, 2 dm. [8 inches] long, 1 cm. [6 lines] wide, the upper surface bright green, the lower glaucous and ribbed with the filiform nerves; bracts foliaceous, lanceolate-acuminate, amplexicaul, upper ones opposite; pedicels equalling or slightly surpassing Bes. Bot.—VOL. II.] PURDY— CALOCHORTUS. 127 the bracts, erect in flower, recurved in fruit; flowers broadly open-campanu- late; sepals from elliptical to narrowly ovate, abruptly acuminate, tinged with purple on the outer surface, purple-veined on the inner, two-thirds as long as the petals; petals broadly obovate-cuneate, acute or rounded at apex, creamy white or tinged with purple, bearded all over the inner surface with long hairs which are white on the upper half of the petals, purple on the lower, some- what arched by the narrow, transverse, semicircular, conspicuous gland, the shallow pit of which is covered by a densely hairy narrow scale; anthers lan- ceolate, abruptly acuminate, cream color or purplish, shorter than the fila- ments, which broaden to the base; capsule 3 cm. [one-fourth inch] long, 2 cm. [Io lines] wide, broadly elliptical, with the three wing-like valves trans- versely veined. The above is the original description as given by Miss Eastwood. ‘Willamette Valley [Oregon], in the foot-hills.”’ C. purdyi may be distinguished from C’. dol/mzez by its color and the absence of the gland. The two are fre- quently confused and many of the specimens in herbariums labeled C. tolmzez are in reality C. purdyz. Through C. howelliz, C. tolmiez and C.. purdyz are closely related to C. longebarbatus; while the large forms of C. maweanus link them to the e/egans group. 16. Calochortus apiculatus Aaker. Calochortus apiculatus BAKER, Journ. Linn. Soc. (Bot.), Vol. XIV, 1875, P- 3095. Stem stout, erect, a foot to 18 inches high; the single radical leaf 6-12 inches long, 3-9 lines wide; bracts linear, acuminate, 1-3 inches long; sepals lanceolate, greenish white, acute, 6-9 lines long; petals straw colored, broadly obovate, one inch long, distinctly pitted, above with only scattering hairs, pit densely hairy and without scale; anthers 4 lines long, acuminate; filaments as long; capsule 12-15 lines long, narrowly oblong. ‘‘Columbia brittanica ad montes Pend Oreille et Koot- enay.”’ The above is drawn from Baker’s original description and from letters. The writer has no personal knowledge sot the species: Section II. JMMarzposa. Flowers and fruit erect on stout pedicels; flowers open-campanulate; gland usually densely hairy; capsule (except in Group 1) narrow, with thick lobes, 128 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. septicidal; sepals often hairy or subglandular, or spotted within; seeds ascending and somewhat turgid, with white, loose and spongy, minutely tesselated testa. Group 1. OREGON MARIPosAs. Capsule with thin, acute or winged cells (as in Section I, but erect), also resembling Section I in the solitary, flat, shining leaf. Closely linked to the preceding group of Eucalochorti (Giant Star Tulips). Oregon and to the northeast. * Petals with an indigo blotch in the center. 17. Calochortus nitidus Doug/. Calochortus nitidus Douc as, Trans. Hort. Soc., Vol. VII, 1830, p. 277, Tab. UX; fig: a: Stem bulbiferous near the base, slender, stiffly erect, not bracted in the middle, bearing an umbel of 2-4 flowers subtended by 2-4 linear bracts; ped- icels often 3 inches long; sepals ovate-lanceolate, long-acuminate, exceeding petals; petals white to lavender, a conspicuous indigo spot in the middle, 2 inches long, the same in width, broadly cuneate, rounded above, with a small rounded gland densely matted with short hairs, and scattering long hairs around and above; filaments filiform, winged below; anthers linear- oblong, two-thirds as long as filaments; capsule elliptical-orbicular, strongly winged and crested. Described from specimens from Union, Oregon. ‘¢Columbia britannica’’ (Dougl.); ‘‘Oregon’”’ (Spaulding). This species is the largest flowered of the group; it is found in wet fields, from eastern Oregon through Idaho to Montana and northeastern Nevada. Specimens from Yel- lowstone Lake are yellow. The purple blotch on the petal seems to be distinctive. ** Petals not spotted in the middle; flowers lilac, smaller than in the preceding. 18. Calochortus greenei Watson. Calochortus greenei WATSON, Proc. Amer. Acad., Vol. XIV, 1879, p. 264. Stem stout, branching, often a foot high or more, 2-5-flowered; leaf about equalling the stem, an inch broad; bracts narrow, elongated; sepals greenish with more or less of lilac within, and with a yellowish hairy spot above the base; petals broadly fan-shaped and obtuse, 1% to 1% inches long, lilac, somewhat barred with yellow below, strongly pitted and arched, the lower part densely covered with very long yellow hairs, upper part of the blade more thinly hairy, not ciliate; pit densely villous above a broad transverse, laciniate scale; anthers broad, acute or obtuse, % inch long; capsule an inch long, 4 to 6 lines broad, attenuate into a stout beak. Bot.—VOL. II.] PURDY — CALOCHORTUS. 129 “Siskiyou County, California (Greene); Multnomah County, Oregon (Howell).’’ The (lescription and locality are copied from the original of Watton, but the latter habitat is incorrect, as Howell’s specimens are C’. longebarbatus. While at work on this revision the writer had not access to specimens of authentic C’. greenez, the examples from Oregon and Washington referred to this species being in reality C. longebarbatus Watson. Specimens from Modoc County, California, a little east of the type locality, appear, however, to be the true C. grcenez. Professor Greene, the discoverer of the species, is of the opinion, which the writer also shares, that Dr. Watson con- fused two distinct species in his description. C. greenez is described as having the erect capsule characteristic of the section, while Professor Greene reported that some of his specimens had the nodding capsule of the Eucalochorti. From the evidence, it would seem that the densely hairy, strongly arched Calochorius described by Dr. Watson is properly another one of the Giant Star Tulips; but this cannot be definitely determined until there is fuller material to work with. 1g. Calochortus pavonaceus Fernald. Calochortus pavonaceus FERNALD, Bot. Gaz., Vol. XIX, 1894, p. 335. Stem erect, stiff, slender, a foot or so high, with a single linear bract near the middle and two or more below the umbel; radical leaf flat, shining, lan- ceolate, not channeled; flowers 1-4 in an umbel; pedicels 134-3 inches long, exceeding bracts; sepals purplish, ovate-lanceolate, acuminate; petals 1% inches long, cuneate-obovate, denticulate, with a broad claw, color lavender to purple, with a circular band above the small round gland, which is covered with densely matted yellow hairs; a few hairs on the margin and silky hairs sparingly scattered over lower third; filaments slender, winged, exceeding the obovate-obtuse anthers, which are 4-5 lines long; capsule elliptical, acutely angled and strongly beaked. The description is from specimens collected near the type locality, ‘‘Pullman, Whitman Co., Washington.’’ 130 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 20. Calochortus longebarbatus Watson. PLaTE XVI, Fic. 1. Calochortus longebarbatus Watson, Proc. Amer. Acad., Vol. XVII, 1882, p. 381. Near C. pavonaceus. Stem slender, a foot or so high, bulbiferous at base, stiffly erect; inflorescence umbellate if more than one flowered, short linear bracts subtending the umbel or a single flower on a stem; true radical leaf one, lance-linear, acute, 2 to 3 lines wide, nearly as long as or exceeding stem, a linear bract-leaf on lower part of stem just above radical leaf; sepals ovate-lanceolate, acuminate, a little shorter than petals, greenish lilac within; petals lavender, lighter below, with a dark purple circular band above gland, obovate-cuneate, denticulate, margin naked; gland small, roundish, covered with densely matted brown hairs, with some long silky hairs above and beside it; filaments slender, winged below, 2 to 3 times as long as anthers; anthers narrowly ovate, obtuse; mature capsule g to ro lines long, strongly winged and beaked. ‘“‘In low grassy grounds, Falcon Valley, Klickitat County, Washington Territory.’’ The description given is of average specimens from the type locality, and from Hood River, Oregon. From the description it will be seen that C. l/ongebarbatus closely resembles C. pavonaceus except that the bracts are near the base instead of the middle, there are no hairs on the mar- gin of the petals, and the filaments are somewhat longer. The two may run into each other, but to definitely ascer- tain whether this is the case specimens from a much wider range are needed. 21. Calochortus howellii Watson. Calochortus howellit Watson, Proc. Amer. Acad., Vol. XXIII, 1888, p. 266. Stem erect, a foot or more high; radical leaf solitary; cauline leaf narrow and short; sepals ovate and shortly acuminate; petals white, an inch long, denticulate, slightly ciliate near the base, covered within with short crisped hairs, those above the gland denser and dark greenish; gland transversely oblong, densely covered with short yellow hairs; anthers oblong-acute, and apiculate, 3 lines long; capsule elliptical, acute, 9 lines long. ‘¢Found near Waldo, Oregon, in 1884, and at Roseburg in 1887.” BoT.—VaL. II.] PURDY— CALOCHORTUS. 131 The above description is drawn from specimens collected near the type locality. 3 } Group 2. Rocky Mountain Mariposa. Gland tiansverse and narrow, extending across petal from side to side; leaf usually as in Section I. 22. Calochortus gunnisoni Watson. Calochortus gunnisoni Watson, Bot. King’s Rept. Ex. goth Par., p. 348. The usually single radical leaf linear-lanceolate, acute, flat, and not chan- neled; stem bulbiferous at base, a foot or so high, erect; inflorescence usually umbellate, the umbels 1-4-flowered and subtended by two or more rather broadly lanceolate, acuminate bracts; one or several cauline leaves below the umbel; sepals narrowly ovate, acute, with scarious margins, yellowish within, often marked with dark blue; petals creamy white, often banded with dark blue above gland, cuneate, with a broad claw, usually rather truncated above, sometimes slightly rounded, 12-15 lines long; gland narrow, extending trans- versely nearly from side to side of petal, and covered with short, dense, glandular clavate hairs; anthers equalling filaments, ovate, cuspidately acute; capsule cylindric-triangular, Described from an average sized specimen from Boulder, Colorado. The species is found on the eastern side of the Rocky Mountains from Nebraska to New Mexico. ‘‘Rocky Mountains of Colorado. Collected also in Utah by Gunnison.”’ The color of the petals is variable, flowers from different localities showing all the gradations from white, through pink, lilac, purple, and blue, with considerable variety in the color of the bands marking the lower portion of the petal; but a greenish tinge and greenish hairs characterize them all. Some of these color forms have been named. Group 3. WeEEpD’s MarIposa. Petals covered with slender hairs; capsule narrowly oblong, with thick obtusely angled cells; radical leaf as in C. albus (Section 1), solitary, long, shining, and not channeled; bulb heavily coated with coarse black fiber. 132 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 23. Calochortus weedii Wood. PLATE XVII. Calochortus weedit Woon, Proc. Acad. Nat. Sci., Phila., 1868, p. 169. Corm deep seated, heavily coated with coarse dark brown fiber; stem 1-3 feet high, stout and flexuous, usually branched, leafy, one to many flowered, not bulbiferous; radical leaf lanceolate-acuminate (the same as in C. aléus), solitary, broad, shining, flat and not channeled; cauline leaves broad, acumi- nate; sepals often exceeding petals, narrowly ovate-lanceolate, acuminate, yellowish within, often with a hairy spot at base, with scarious margins; pet- als cuneate, denticulate, either rounded or more or less truncate above, 12-15 lines long, orange colored and covered with long silky yellow hairs, each set in a dark brown spot, upper quarter usually naked; gland small, circular to oblong, densely matted with short hairs; filaments filiform, exceed- ing the oblong anthers; capsule narrow, attenuate upward, 1% inches long. Described from specimens from San Marcos, San Diego County. **San Diego.’’ C. weedzz in its various forms is found in the Coast Range of California, from San Luis Obispo County south to Lower California. It is easily distinguished from other Mariposa Tulips by the single large radical leaf (one to two feet long by five to eight lines wide), the very heavy, coarse, stringy, fibrous coating of the bulb, and the long silky hairs springing from the brown dots on the petals. The species is almost as variable as C. venustus, each variation having its own range, where it is found to the exclusion of all others. Beginning at San Diego, we have the typical C. weedzz, a large orange-colored flower, covered with yellow hairs and dotted with brown. In this form the petals are usually full, although occasionally somewhat truncated. Nearly all of the C’. weedz¢ in San Diego County, whether in the interior or on the coast, conform to the type. 23a. C. weedi var. purpurascens Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 265. (C. plummere GREENE, Pittonia, Vol. II, 1890, p. 70). Going north to Los Ange- les and San Bernardino, we find a broad belt of country where the flower of C. weedzz is still full petaled, but the color is lilac or lilac purple. While possessing hardly any \q BotT.—VOot,. II.] PURDY— CALOCHORTUS. 133 analytical points of difference, it may be said that in cultiva- tion this form proves to be a stronger and hardier plant than the type. A curious fact in connection with the bulb is also worthy of mention. The coarse fibrous coating which the variety has in common with the type is often found to cover not one bulb, as is usual, but two large bulbs of about equal size, laid so flatly face to face that outwardly the bulb retains the round form of the species, and it is only by b-eaking the coating that the presence of the second bulb is discovered. This habit of the bulb can hardly be called oifsetting, as it is impossible to call either the parent bulb. In the San Jacinto Mountains this form varies to pale pink and white. It is the C. weedi var. albus of horticulture. A fine plate of this variety may be found in ‘‘The Gar- den,” Veb..2,. 1895. 236. C. weed? var. vestus. Continuing still further north to Santa Barbara, we again find the species, but the petals are much more truncated and curiously fringed with brown hairs, while the color is reddish brown. 23c. C. weedi var. obtspoénsis, var. nov. (C. obisfo- énsts LEMMON, Bot. Gaz., Vol. XI, 1886, p. 180.) This is an extreme form found in San Luis Obispo County. The petals of the flowers are still more truncated than in the preceding, and near the town of San Luis Obispo, which I think is the northern extreme of its range, the flowers assume a most fantastic form, the brownish petals being so much truncated that the sepals far exceed them, and the hairs which are scattered in typical specimens here seem to be condensed upon the small remaining surface of the petals. ‘‘On dry, stony hills near San Luis Obispo, Cal.’’ It is, of course, impossible to enumerate here all the many gradations in this plant, which forms one of the most interesting studies in plant variation. Group 4. GoLpEN Bow. Mariposas. Petals yellow, lower half covered with clavate hairs; radical leaves long- linear and deeply channeled. 134 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. 24. Calochortus clavatus Watson. Calochortus clavatus Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 265. Stem stiff, very stout, strongly zigzag, branching, 1-3 feet high, if few flow- ered umbellate, if many flowered with a terminal umbel and short laterals at the axils of the rather large cauline leaves; the single radical leaf glaucous, linear, deeply channeled, often 1-2 feet long; cauline leaves as broad and prominent; pedicels stout, 2-5 inches long; sepals acuminate, broadly ovate- lanceolate, nearly as long as petals, yellowish within, often greenish without, spotted, with dry scarious margins; petals rather truncate, 18 lines wide, 21 lines long, broadly fan-shaped and strongly arched, with a broad claw, the gland deeply pitted, with yellow hairs; the side of gland and the lower half of petal densely hairy with long yellow hairs, each tipped with a knob-lik2 point, and purplish red at base; color a rich yellow, with claw often reddisi brown and a reddish brown band above the hairy zone; perianth bow}- shaped (‘‘formed like a broad-based cup’’); filaments slender, a little exceed - ing the ovate-oblong, obtuse, purplish-brown anthers; capsule narrow, attenuate above and below, 3 inches long. Described from medium sized plants. Found on dry rocky points, usually in volcanic soils. The species is widely but peculiarly distributed from Newhall (Los Angeles County) to San Luis Obispo; it is also found in the Sierra Nevada near Pleasant Valley (El Dorado County), and on White Rock, an isolated quartz rock- mass in the plains of Mariposa County. Doubtlessly found also on lava formations at many intermediate points. ‘‘California (San Luis Obispo; J. G. Lemmon, 1878).’’ This Calochortus is the stoutest stemmed, tallest, and largest flowered of all the Calochorti. The heavy, strongly zigzag stem, yellow bowl-shaped flowers, and clavate hairs, are strongly marked characters shared in by no other mem- ber of the genus. The knob-like tips of the hairs are trans- lucent, having the appearance of little icicles. But although the prominent characters remain constant, there is con- siderable local variation. ; In Ventura and Los Angeles counties (Piru City to New- hall), for instance, the flowers are a very rich yellow and the plant is rather low and stout. Near the city of San Luis Obispo, the plant has the same habit, but the upper half of each petal is backed with olive which showing through gives a peculiar changeable color effect. In El Bor.—VOL. II.] PURDY — CALOCHORTUS. 135 Dorado County the form is very tall and large, often three feet in height, with flowers five inches across, and of a lighter yellow. The hairs are longer and the knob-like tip is smaller. Y 25. Calochortus concolor, sp. nov. Calochortus luteus var. concolor BAKER, Plate in “ The Garden,’’ Dec. 7, 1895. Bulb large, reddish; radical leaves narrow, glaucous, deeply channeled; one or more cauline leaves below umbel; stem 2 feet in height, one to several flowered, if more than one flowered, umbellate; pedicels stout, 1 to 3 inches long; umbel subtended by linear bracts; all parts of plant very glaucous, lower stem and sepals with a bluish bloom; sepals ovate-acute, with scarious margins, yellowish within, brownish on back; petals a deep rich yellow, tending toward orange, broadly fan-shaped, 15-18 lines long, and as broad as long, slightly rounded above, the lower third densely hairy with long erect yellow hairs; gland small, oblong; anthers yellow, linear, exceeding fila- ments; capsule strongly triquetrous, lance-linear, attenuate above (imperfect in type specimen). Described from a large plant collected by Mr. D. ‘Cleve- land at Laguna, on the edge of the desert, San Diego County. This is one of the desert species found in rocky soil in various places on the desert side of San Diego County and in the Cuyumaca Mountains, also at Mill Creek, near San Bernardino, and in Hermit Valley, Riverside County. The writer does not agree with Mr. Baker in referring this spe- cies to C. Juteus; its affinities, if any, are with C. nuttalliz or C. clavatus. Following C. clavatus it is the largest flow- ered, and the showiest of the yellow Mariposa lilies. a Group 5. Petals nearly naked, gland round, small, and densely hairy with matted hairs; leaves ashy blue, linear, deeply channeled. Desert plants. 26. Calochortus kennedyi Porter. Calochortus kennedyi PorTER, in Coutt. Bot. Gaz., Vol. II, 1877, p. 79- Plate in ‘‘ The Garden,”’ Feb. 11, 1893. Stem very low, rather stout, often only 1-4 in. high, 2-4-flowered; radical leaves linear, 2-10 in. long, channeled, very glaucous, as are the stem and bracts; flowers produced in an umbel, which is subtended by short bracts; pedicels 3-4% inches long; sepals ovate-oblong, about equalling petals, obtuse 136 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. or acute, with white scarious margins, vermilion inside, often spotted brown near base, brownish without; petals cuneate, rather truncate, 10-15 lines long, not quite as broad as sepals, a dazzling vermilion, naked above, a few scat- tering hairs below; gland small, round, densely matted with short hairs and bordered nearly black; filaments one-half as long as the oblong-ovate, obtuse, brownish purple anthers, which are 3-4 lines long; capsule 14~2 in. long, 4-5 lines wide, attenuate above. Found in the arid regions of southeastern California from near the Ojai Valley (Ventura County) and Tehachapi Station, along the eastern flank of the southern continua- tion of the Sierras into San Diego County, eastwardly to the Argus Mountains in southwestern Nevada, and throughout Arizona. ‘¢ Kern County, California.”’ In California the flowers are a vermilion color, in Arizona and Nevada, orange. While the stem may reach eighteen inches in height, it often hardly rises above the ground, and the meager ashy foliage gives little promise of the dazzling flower. It is probable that in eastern Arizona and in Nevada C’. kennedyz will be found merging into C. aureus. 27. Calochortus aureus Watson. Calochortus aureus WATSON, Amer. Nat., Vol. VII, 1873, p. 303. Low, 4-6 inches high, with a single linear carinate radical leaf, 3-4 inches long; scape short, 1-2-flowered, the single pair of bracts linear, 2 inches long; sepals greenish-yellow, with a dark-purple spot near the base, oblong- or ovate-lanceolate; petals broadly cuneate, 15 lines long, bright-yellow, with a small well-defined, circular, densely hairy gland near the base, and a lunate purplish spot above it; young capsule narrowly oblong, not winged. ‘“* On sand-cliffs, Southern Utah.’’ The writer has no acquaintance with the species except as scant herbarium specimens. ‘The description and local- ity are the original of Watson. Group 6. BuTTERFLY TULIPs. Petals slightly hairy below, usually oculated and brilliantly colored; gland prominent, round or lunate; stem bulbiferous at base, erect, slender, branching; radical leaves, usually a pair, channeled, linear, slightly glaucous, not ashy blue as in the desert species; cauline leaves and bracts narrow. Bot.—VOL. II.] PURDY— CALOCHORTUS. 137 C. luteus and C. venustus are the true Mariposa or Butter- fly Tulips. It would be difficult to name any other group of plants in which can be found such a wonderful diversity of color and marking. The colors range from white through lilac to purple, from delicate pink through light reds to deep glowing reds; buffs, yellows and citrons are all present; and the tintings, oculations, and blotchings are bewilder- ingly numerous. Watson bases his division of the group on the differences in the gland: that of C. /uteus being from transversely oblong to narrowly lunate; that of C. venustus from round to longitudinally oblong. While the casual observer would place all together, Watson’s division is a true one. In C. luteus the capsule is acuminate from a triangular base, in C. venustus, linear. 28. Calochortus luteus Dozg/. PLATE XVI, Fic. 2. Calochortus luteus DouGt., Bot. Register, Vol. XIX, 1833, Tab. 1567. Plants usually dwarfish, 8-1o inches high, stem slender, stiffly erect, bul- biferous at base, often branching; bulblets enclosed within sheath of stem; radical leaves linear, channeled, 1-3 lines wide, bright green, slightly glau- cous; cauline leaves or bracts linear, 1-3 inches long, peduncles 1-4 inches long; sepals narrowly ovate-lanceolate, acute, 12 lines long, 2-3 lines wide, yellowish within; petals cuneate, as long as broad, slightly rounding above, yellow or tending toward orange, not oculated, but having penciled lines radiating from gland to center of petal, claw broad; gland rather broad, lunate, densely hairy with ascending matted yellow hairs, a very few scatter- ing hairs above reaching to middle of petal; stamens about equalling style, filaments slender, a little longer than the light yellow, oblong-linear, acute anthers; mature capsule acuminate from a triangular base, 14-2 inches long. The description is drawn from strong specimens collected at Monterey. ‘¢ California.’’ The type described of C. /uteus follows the coast-line from Anderson Valley (Mendocino County) to San Diego County. It grows in adobe soil and is dwarfish. Speci- mens from Mendocino, San Francisco, Monterey, and San Diego counties are exactly alike. 138 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Farther in the interior, from Sonoma County (Sebasto- pol) to the Sacramento Valley, and south along the foot- hills and higher plains of the east side of the San Joaquin Valley, a second form grows, exactly like the type except that it is tall and slender. This is the form commonest in the Sacramento and San Joaquin plains. By slight grada- tions through a multitude of forms, a dark eye develops, and the habit becomes more luxuriant, until we have C. luteus var. citrinus. 28a. C. luteus var. citrinus. This variety is merely a color variation of that which follows, the color varying from yellow to deep lemon, and the central spot being darker and not oculated. This variety is found at its best in the Russian River region from Hopland, Mendocino County, to the town of Sonoma, in Sonoma County. In the latter place, the color is a deep citron and the eye has disappeared. At various other points, for example, eastern Lake County, Linden (San Joaquin County), near Placerville (El Dorado County), and in various localities in the foot-hills of the Sierra this form is approached. 286. C. luteus var. oculatus. This variety is usually much taller, larger flowered, and often many flowered; petals white to lilac or purple, often 2 inches long, the same in width, the center of each showing an oblong dark brown spot oculated with yellow, and many radiating pencilings; sepals also oculated. The type of C. luteus var. oculatus as above described is common in the Coast Range from Shasta County to San Francisco Bay. It is to be expected that it would occur as far south as Monterey County, but the writer has seen no specimens from that region. In the foot-hills of the Sierras, from Butte to Kern coun- ties, there is a form slightly differing from the type. The petals have less yellow in the middle third, the blotch is smaller, the gland rather deltoid than lunate, and the pen- cilings are lighter—all trifling differences which only close study will detect; yet they are quire sufficient to separate the oculatus of the Coast Range from that of the Sierras. C. luteus var. oculatus of the Sierra foot-hills is very con- stant. While in some localities it crosses with var. cztrznus, Bot.—Vot. II.] PURDY — CALOCHORTUS. 139 specimens from near Chico (Butte County), from Placer- ville (El Dorado County), and from Dunlap (Fresno County), are identical in every respect. As before stated, C. luteus var. citrinus and var. oculatus hybridize readily, and in many localities where both are found, there are cross breeds in endless variety, running as is usual in such cases to greater extremes than either parent. In Mendo- cino County, from Ukiah to Hopland, and in El Dorado County, near Camp Creek, these crosses are particularly plentiful and beautiful. There are other strains of C. /uteus, each found in a more or less extensive territory. 28c. C. luteus var. robusta (C. venustus var. robusta (Hort.).) In por- tions of El] Dorado County a form approaching C. ocwlatus is found growing in wet grounds (oftener in wire grass lands), which is there dwarfish and with three to five almost spheroid bulblets on the stems. The colorings are very rich; the forbidding surroundings seem to have developed unusual vitality in this form, for in ordinary soil the bulbs produce unusually tall, stout stems but retain the bulb ferous habit. 29. Calochortus vesta Purdy.' Calochortus vesta PurDy, The Garden, Oct. 12, 1895, with colored plate; Gardeners’ Chronicle, 1895, Part II, page 14. Tall, large flowered, stem leafy, pedicels elongated, 8-12 inches long; petals more narrowly cuneate than in C. luteus var. oculatus, white tinged with lilac (with rare albinos), instead of an oculated spot having a broad red- dish or dark brown band across the middle; gland narrow, doubly lunate, extending across from side to side of petal; bulblets long and slender, I to 4 to the stalk, not enclosed in sheath of stem, but set at an angle. While C. vesta is grouped with C. luteus and C. venustus, it is a strongly marked form which does not hybridize with either. C. vesta is found in adobe soil (sticky black or blue clay) from Sonoma and Napa counties to Humboldt County, California. 1C. vesta and the varieties of C. venustus—var. roseus, vat. purpurascens, and var. eldo- rado—were first mentioned in a catalogue of bulbs issued by the writer. C. vesta was first described in an unsigned article in ‘‘ The Gardeners’ Chronicle” (August, 1896). It was so named in honor of the author’s wife. In the same article, C. venustus var. roseus and var. purpurascens were also described under the names C. voseus aud C. purpurascens, thus giving specific rank to the catalogue varieties. A fine colored plate of C. westa, C. venustus (the type, known as var. voseus), and C. venustus var. purpurascens, was published in ‘‘ The Garden” (Ijondon); and a good plate of C. venustus (type), C. luteus (type), and C. luteus var. citrinus, appeared in the same mag- azine 1883 (?). (3) December 5, Igor. 140 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Calochortus bulbs move from year to year by means of a spongy process growing from their base, at the end of which the new bulb is formed. By this means the bulblets of C. vesta, being set at an angle, travel some distance before at maturity they take a vertical position. By digging into a patch where C. vesta has been undisturbed for some years, the ground will be found literally full of bulbs, the immature ones placed at every angle. In most bulbiferous Mariposas, the bulblets being placed inside the stem sheath, and parallel to the stem, grow straight down and form a close clump. C. luteus var. robusta has a habit somewhat similar to that of C. vesta, though not so marked; and in C. venustus var. purpurascens the habit is exactly the same as in C. vesta. 30. Calochortus venustus Doug/. Calochortus venustus DOUGL. ex. BENTH. in Trans. Hort. Soc., Ser. II, Vol. I, 1835, p. 412, Tab. XV, fig. 3. Stem stiff, erect, usually branching, 4 inches to 3 feet in height, bulbiferous, with a single bulblet at base; one or two linear, radical leaves, 1-3 lines wide, quite glaucous; pedicels 2-5 inches long; sepals oblong-lanceolate, 4 lines wide, 18 lines long, acute, as long as petals; petals broadly obovate- cuneate, slightly rounding above, wider than long, 16 lines long, 20 lines wide, color white to lilac above, carmine below, oculated like C. luteus, with reddish pencilings on each side of eye, and a transversely oblong, rose- colored blotch near apex, claw reddish brown; gland roundish or oblong, densely matted; filaments a little exceeding the narrowly oblong, obtuse anthers; capsule linear, 2-3 inches long. Described from a specimen collected in San Benito County, California. | C’. venustus is even more variable than C. Juteus. While C’. luteus is almost always found in clay, sometimes heavy clay, C’. venustus usually grows in light open ground, often in sandy soil. Bor.—VoOL. II.] PURDY— CALOCHORTUS. I41I 30a. C. venustus var. roseus, The Garden, Oct. 12, 1895. The typical form described above is known as var. roseus, or C. roseus. 306. C. venustus var. eldorado, var. nov. Differs from var. vroseus in having the more cuneate petals always narrower than long. Found in all colors from white through lilac to purple, from pink through many shades to deep claret, with or without rose-colored blotch near the top of petal. The plants are stout, usually tall (1-2 feet), branching, and large flowered. C. pictus (The Garden, Oct. 12, 1895) is merely a white form of the above variety. 30c. C. venustus var. purpurascens, The Garden, Oct. 12, 1895. This is a strong growing variety, like the type but taller; flowers lilac to purple, petals oculated and more leafy, but without the rose-colored blotch. The plants are very bulbiferous, producing 1-4 bulblets a year, which scatter as do those of C. vesta; they are not enclosed in a stem sheath. Found growing in adobe (heavy clay) soils, from San Luis Obispo County to Suisun Bay, California. 30d. C. venustus var. sulphureus, var. nov. As in the preceding, like the type but taller; petals a light, warm yellow, with eye in center, and rose- colored blotch at top. Occurs at Newhall and Alcalde (Kern County), in the lower end of the San Joaquin Valley. C. venustus is as widely scattered as C. /uteus, and has many strains, some local, others widely spread. There are two principal strains. One commences in the Coast Range at Antioch (Contra Costa County) and extends southward into the interior or dryer portion of the Coast Range to Los Angeles County. The type described, as found from Antioch to Paso Robles, always has a rose-colored blotch at the top of the petal. In this strain the petals are broader than long, giving a very full flower. At Paso Robles there is a break, and we find a wonderful variety of color forms, from white to purple, and from pink to deep red, always with a rose a ea a aaae 1 The original C. venustus purpurascens Watson was described from specimens from Kern County, in the southern Sierras, and is without doubt only the purple extreme of the El Dorado strain. The name has for years been used for a purple variety of the Coast Range form, which is distinct in every way. As the latter is perfectly distinct, with features by which it can be readily identified, and the former is a mere color extreme, occurring usually with other colors, the rule of priority could, I think, be shaded a little, and the name be used for the Coast Range type. 142 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. blotch at the top of the petal. From this point south to Los Angeles County the forms seem to change frequently. At Preston and Newsom’s Springs (San Luis Obispo County) there is a pretty white form, in color exactly like C. cataline (white, tinged lilac), destitute of both eye and rose-colored blotch, but with the ovate dark brown spot on the claw of the petal, as in C. cataline. At Santa Barbara the type with the rose blotch reappears, and it is found at intervals southward. In the Ojai Valley and at Newhall there is a white-flow- ered variety with a rose blotch and narrow petals, closely resembling the white-flowered plants at Paso Robles. It is also tall and slender. At Newhall a new color strain comes in, quite novel in C. venustus; this (var. sulphureus) is light yellow, narrow petaled, and has the rose blotch. At Elizabeth Lake, a point about midway between Newhall and Alcalde, the species is again seen in the infinite variety of color found about Paso Robles and in the southern Sierras, but with the additional yellow form. Newhall is the most southern point from which the writer has specimens of C. venustus. Mr. S. B. Parish has written a very interesting descrip- tion of these color forms. The Eldorado strain, so termed by the writer in a cata- logue of bulbs, is found in the Sierra Nevada. It begins to appear a little north of Camp Creek (El Dorado County), high up in the foot-hills in the upper Yellow-Pine belt (growing oftenest in open woods), and extends at a nearly uniform altitude in the Sierras south to the Tehachapi and Tejon mountains. The colors of the flowers are richer than those of the Coast Range forms. They vary from white to lilac and deep purple, from pink to deep red, and sometimes light cream flowers are found. ‘The eye is dark brown, not much oculated, the blotch at the top of the petal varies from rose to gold, and flowers can be found with rose or gold rays across the entire top of the petal; but the rose blotch at the apex, instead of being a constant feature as is Bot.—VOL. II.] PURDY— CALOCHORTUS. 143 usual in the Coast Range forms, is rare. At all points from which specimens of this form have been collected the variety of color is shown to vary greatly in proportion; sometimes nineteen-twentieths of the flowers are white, again red or purple will predominate, and this in localities only a few miles apart; but no single color form is ever found to the exclusion of all others. Group 7. Litac Mariposas. Petals white, lilac, or purplish, not oculated, more or less hairy; gland small, round, and densely hairy; leaves linear, channeled; capsules linear except in C. cataline. 31. Calochortus splendens Doug’. Calochortus splendens DouGL. ex BENTH. in Trans. Hort. Soc., Ser. II, Vol. I, 1835, p. 411, Tab. XV, fig. 1. Stem slender, often bulbiferous at base, usually a foot or two high; sepals often spotted purple, ovate-acuminate, about equalling petals; petals broadly fan-shaped, circular above, 1-5 lines long, 18 lines broad, upper half naked, scattering short hairs around the gland and on lower third, the claw very short; color from lilac to purple, lighter about gland, usually reddish purple on claw; gland small and round with a mound of matted agglutinated hairs which are scarcely distinguishable as hairs, and are frequently absent; anthers purple, obtuse, one-third as long as filaments; capsule linear. Described from strong plants from San Diego, California. In horticulture the type is known as C. splendens var. atroviolacea. It isfound from San Diego to Santa Barbara counties, and on the coast islands. 31a. C.splendens var. montanus, var.nov. Like the type but very slender, 8-12 inches high, smaller flowered, and more densely hairy about the gland with short yellow hairs; color lilac to salmon pink; often bulbiferous. Described from specimens from Raynetta, San Jacinto Mountains. 316. C. splendens var. major, var.nov. This resembles var. montanus but is not bulbiferous, is stouter and much larger flowered; petals pale lilac, lighter below, with very long tangled hairs scattered on middle third. Described from specimens from Monterey County, Cali- fornia. Found also in San Luis Obispo County. 144 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. The figure shown in Douglas’ plate is that of the form here described as the type. Baker, in his Tulipe, gives an accurate description of this same form, calling it Calochortus splendens; but Watson, in the ‘‘Botany of California,’’ has described as the type the form here given as var. major. The marked difference between these two forms is in the hairs. In the type the hairs are short and found on the lower third of the petal; in var. major they are on the middle third of the petal, and are long and cobwebby. In addition to this, the latter variety is of a stronger habit, has larger, lighter colored flowers, and is without bulblets. 31c. C. splendens var. rubra, a form found in eastern Lake and Napa counties, is singularly enough in gland and hairs almost exactly like the type at the other extremity of the range. It is a very large plant, with a deep-seated yellow bulb, tall (1-3 feet), stout stem, and a large reddish lilac, pink or purplish flower, the petals quite hairy with short hairs on lower third, and with a deep reddish purple claw. C. splendens var. montanus has been mistaken for C’. pal- mert, but it is closely related to the type; in fact, aside from the color of the hairs above the gland, and the more slender habit, it hardly differs from the type except in its habitat. The writer has it from ‘‘cienagas’’ (wet springy spots) near Tehachapi, from wet spots near Bear Valley (San Bernar- dino County), and from wet meadows in the San Jacinto Mountains. The other three forms always grow in dry, rocky soil. 32. Calochortus palmeri Watson. Calochortus palmeri Watson, Proc. Amer. Acad., Vol. XIV, 1879, p. 266. Stem very slender, lax and flexuous, a foot or two high, 1-7-flowered, bulbiferous near the base; sepals with narrowly acuminate recurved tips, spotted; petals 6-12 lines long, white (or yellowish below) with a brownish claw, and with scattered hairs around the ill-defined, broad, densely hairy gland; anthers obtuse, 3 lines long, capsule very narrow, an inch long or more. ‘‘California, near the Mojave River.’’ The above description and locality are quoted from the original of Watson. Bot.—Vot. II.] PURDY— CALOCHORTUS. 145 The writer has never seen a specimen which conformed to the description; all of those seen from Southern Cali- fornia collections were either C. cnvenustus or C. splendens var. montanus. 33. Calochortus cataline Watson. Calochortus cataline Watson, Proc. Amer. Acad., Vol. ang 1879, p. 268. Calochortus lyoni Watson, Proc. Amer. Acad., Vol. XXI, 1886, p. 455. A foot or two high, general habit as in C. splendens; stem bulbiferous at base; sepals broadly ovate-lanceolate, acute, a little shorter than petals, greenish, spotted dark purple on claw; petals cuneate, longer than wide, rounded above, narrowing abruptly to a short narrow claw; color from white tinged with lilac, to lilac purple, with a large ovate purplish-maroon mark, the apex covering claw and the rounded end above gland; gland oblong, densely matted with short agglutinated hairs, a few scattering hairs around and above; filaments slendér, about four times as long as the yellowish, oblong-obtuse anthers; capsule oblong, rounding at both ends. Described from specimens taken near Los Angeles, California. ‘‘Santa Catalina Island.’’ [California. | C. cataline is found from Los Angeles and San Bernar- dino along the coast to Santa Barbara, and on all the islands along the coast. It is one of the least variable and best marked of all the Mariposas. The oblong capsule is a distinctive feature, while the peculiar ovate mark on the petals is equally characteristic. The species was first described by Watson from a plant in the capsule, the specimens having been collected on Santa Catalina Island. Later, he described the same spe- cies, under the name of C. lyonz, from flowers collected at Los Angeles and Newhall by Dr. Asa Gray and W.S. Lyon. C. lyoni also grows on Santa Catalina Island, and there is no doubt of its identity with C. cataline. In an able article in ‘‘Erythea,’? Dr. Davidson of Los Angeles shows that the two species are the same. 34. Calochortus invenustus Greene. Calochortus invenustus GREENE, Pittonia, Vol. II, 1890, p. 71. Stem very stiff, stout as compared with C. splendens, strongly bulbiferous at base, the bulblets large, ovate-oblong as in C. nuttallii; radical leaf linear, 146 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. deeply channeled; plant very glaucous, as in most desert species; inflores- cence umbellate or at least the larger divisions of stem so; umbels, or branch divisions, subtended by three or four lanceolate bracts an inch or two long; sepals ovate-lanceolate, acute, considerably shorter than petals, yellowish green within; petals obovate-cuneate, short clawed, rounding above or abruptly acute, smoky white, claw purplish; gland small, oblong, densely hairy with matted hairs, yellowish short tangled hairs above gland; anthers 2-3 lines long, not equalling filament; capsule 3-4 lines broad in the middle, tapering both ways. The description given is drawn from specimens taken at Bear Valley (San Bernardino County), California. ‘‘Higher mountains to the westward of the Mojave Desert.”’ C. invenustus was discovered by the namer, Professor Greene, near Tehachapi. It is also found in the Tejon Mountains, and doubtlessly grows all along the line from there to Bear Valley. Specimens from the former locality are identical with those here described. The characteristics of the species place it between C. nuttallic and C. splendens. The prominent offset, stiff, stout stem, and tendency to flower in umbels, are strongly suggestive of C. nuttalliz, while the flowers resemble those of C. splendens var. montanus. 35. Calochortus excavatus Greene. Calochortus excavatus GREENE, Pittonia, Vol. II, 1890, p. 71. Resembling the last [C. zzvenustus], but the bracts ovate-lanceolate, scar- ious almost to the striate-veined middle portion, their acuminate tips re- curved; petals white shaded with lurid purple above, but dark purple below and about the broad obovate hairy gland, which is deeply impressed, appear- ing like a yellow saccate body on the outside of the petal; stamens as in the last, but anthers dark maroon. ‘‘From Bishop Creek, Inyo County, California, collected by Mr. W. H. Shockley (No. 427).”’ Having little knowledge of this species, the writer has quoted the original description and locality of Professor Greene. From specimens seen, the species would seem to come between C. znvenustus and C. nuttalliz. BotT.—VOL. II.] PURDY— CALOCHORTUS. 147 36. Calochortus flexuosus Watson. Calochortus flexuosus Watson, Amer. Nat., Vol. VII, 1873, p. 303. Stem slender, very flexuous or almost decumbent, a foot or so high, branching; bracts linear or lanceolate, 6-15 lines long; sepals oblong-lanceo- late, greenish with a deep purple spot; petals broadly obovate-cuneate, 12 to 15 lines long, purple, claw deep purple; gland obscure, purplish yellow, scat- tering glandular hairs above gland; capsule broadly oblong. The description given is substantially that of Watson. ‘¢ Southern Utah and Northern Arizona.’’ In 1897, the writer received a number of fresh flowers from St. George, Utah, the type locality. They showed the color to be a deep, rich purple (the flowers are some- times white), the markings varying considerably, showing bands or spots on either petal or sepal; scarcely any two of the flowers were alike in this particular. The very flex- uous habit is distinctive. The stems are often even decum- bent (‘‘creeping’’ a correspondent has it). Found also in southern Nevada. This Calochortus grows on the hills in red granite soil. Mr. S. B. Parish collected a species in the desert region of Southern California, which he identified as C. flexuosus. This, he writes, grows in tufts of grass in saline meadows. It is, however, hardly probable that the same species grows under such diverse conditions. / 37. Calochortus dunnii, sp. nov. Stem not bulbiferous at base, a foot or two high, slender; leaves linear, deeply channeled; sepals ovate-acute, with white scarious margins, a little over one-half the length of the petals, never recurving, light green without, greenish white within, faintly spotted; petals broadly cuneate, as broad as long, rounded above, white, with a reddish brown transverse band above the gland; gland small and round, densely hairy with short matted hairs, short scattering hairs on each side of the gland only; capsule linear as in C. venustus. Described from specimens flowering in the writer’s grounds; they were originally collected near Julian (San Diego County), California, by the veteran naturalist, George W. Dunn, in whose honor the name is given. So 148 CALIFORNIA ACADEMY OF SCIENCES, [PRoc. 3D SER. far, this pretty new species is known only from the type locality. It forms a connecting link midway between C. splendens and C. venustus. Group 8. GREEN BanpED Mariposa. Petals purplish lilac, with a greenish line down the back, obovate-acuminate. 38. Calochortus macrocarpus Dozg/. PLATE XVIII. Calochortus macrocarpus DouGu., Trans. Hort. Soc., Vol. VII, 1830, p. 276, Tab. VIII. Stem bulbiferous at base, stout, erect and rigid, 1-2 feet high, one or more flowered; the single radical leaf, linear, deeply channeled; cauline leaves 3-5, narrow and convolute; sepals about equalling petals, acuminate, purple inside or tinged with purple, darker at base, sometimes spotted and hairy, with a broad scarious margin; petals obovate, narrowly acute or acuminate, or often distinctly cuspidate, a deep purple lilac, lighter at base, with a broad light-colored claw 1%-2% inches long and half as wide, a greenish line down the middle, the lower third above the gland with scant glandular hairs; gland oblong, densely hairy; anthers purple or yellow, lanceolate, obtuse, 4-6 lines long, about equalling filaments; capsule attenuate upward, 1%-2% inches long. In sandy deserts from northeastern California (Modoc and Lassen counties) to eastern Washington and into Idaho. A very clearly marked species varying but little. There is a white form near Pullman, Washington. The Indians of the northwest are very fond of the bulbs. Group 9. Serco LILIEs. Flowers lilac, white, yellow or pink; gland round; stem prominently bul- biferous at base, umbellate. 39. Calochortus nuttallii Zorr. & Gray. Calochortus nuttallii Torr. & Gray., Pacif. R. R. Rept. (Bot.), Vol. II, 1854, p. 124. Stem erect, stiff, a foot or two high, with a large oblong bulblet at base; infloresence simply umbellate, 1-5 flowered; radical leaf linear, deeply chan- neled; cauline leaves 1 to 3, narrow, glaucous, revolute; sepals ovate-lanceo- late, with scarious margins, yellowish within, with or without a dark spot at base, which is sometimes hairy, much shorter than petals; petals broadly BotT.—VOL. II.] PURDY — CALOCHORTUS. 149 obovate-cuneate, with a rather narrow claw, abruptly acute or rounding above, white, yellowish below, with a small round or oblong gland which is covered densely with agglutinated hairs, a few long hairs scattered about the gland; anthers oblong-obtuse, more or less sagittate at base, shorter than filaments; capsule lanceolate, 1-2 inches long. Described from specimens from southern Utah. C. nuttaliit has the widest range of any Calochortus. It is found from the western flank of the Sierras to Nebraska, and from the Snake River to New Mexico. In color it varies greatly and in other characters slightly. The region of its habitat is too little known to enable any one to mark the range of the variations definitely. In Nebraska and eastern Colorado it is yellow; in many places in central Colorado the color varies wonderfully from white and pink to purple; from Salt Lake to Reno, Nevada, and southern Utah, but one form seems to be common, that which is the type here described; at Pine Valley, southern Utah, the flowers are cream colored; and at Dimee, New Mexico, they are orange. 40. Calochortus leichtlinii! Hooker. Calochortus leichtlinit Hooker, Bot. Mag. Tab. 5862. A dwarfed or alpine form, quite different from any of the other forms, often but an inch or two in height; stem more slender; anthers strongly sag- ittate; petals smoky white with a dark spot. Found in the Sierras at an altitude of from 6,000 to 9,000 feet. There is another variety of this dwarfed alpine Calochor- tus also found in the Sierras, but the description of this form will be reserved until there is fuller material to draw from. 1C. letchtlinii is included under C. xuttallit by Watson, in the Botany of California, Wol, U1, p: 177: : Re] A ' ‘ ve pissy a " “1 eli) 7 : \ ay : 7 , mes a 1 me ; ais , Teeth j a d hain La 5% | a 150 CALIFORNIA ACADEMY OF SCIENCES. (PROC. 3D SER. EXPLANATION OF PLATE XV. Calochortus purdyi. a, Plant, actual size. 6, Petal. c, Stamen, enlarged. d, Sepal. e, Scale on petal, enlarged. f, Ripe pod. 1 “ ‘= rf ‘ ae ie e rat 4 halo ‘ mk te how aero Gf ; aS. OU cre a iS TR SN a a >> Proc. CaLArag. 9c1.32 Ser. Bor Voll PHOTO-LITH. BRITTON & REY, SEL PR TU CALOCHOR' fh arr f Mn aa f Mii At At a MAF ins 4 et Giieh be Re i ; f } ty Linh a Tel) Tid Wit ay (OPTS ane cD ore } He i TT AY f; hei. ' hy why yi GL ie ei ena a) De i Matiet i i Pai Wa ah, ay (is ee! Chien dts he ae \ > tel Wry Wie it Ay! f CALIFORNIA ACADEMY OF SCIENCES. (Proc. 3p SER. a a — a eee : — a EXPLANATION OF PLATE XVI. Fig. 1. Calochortus longebarbatus. a, Petal. | 6, Petal, less enlarged. ‘ Fig. 2. Calochortus luteus. eS a4 a,b, Petals. ; ¢,d, Capsule at different stages of development. fr 7 -~ PrRoc.CALACAD. Scr.32 SER Bor VouIt. [PuRDY| PLATE XVI FHOTO-LITH BRITTON & REY, 5.F. 1, CALOCHORTUS LONGIBARBATUS. 2, LALOCHORTUS LUYEUS. | ?, ‘ nay Wf, f re tec abe e f Male YL Wy , a f EXPLANATION OF PLATE XVII. ) Calochortus weedii. a, 6, Capsule at different stages of development. cs Sepal. d,e, Petals. “" & , Ss hile i wi}, Pe ae er W rie ter age vis ft ay ai Be ead Ppaghs . sede ange a Rs A PRoc.CAL.ACAD. Sc1.32 SER. Bor. Von IT. [PuRoY| PLATE XVII FHOTO -LITH. BRITTON & REY, 5.F. ALOCHORTUS WEEUII. ji CALIFORNIA ACADEMY OF SCIENCES. (Proc. 3b SER. / EXPLANATION OF PLATE XVIII. Calochortus macrocarpus. , am on : ‘ s i - SS Sao Peery he jaar T oe ip~! at it eae 8 io Mis Proc. CAL.ArAn, $c1.32 SER Bor. VoLll (SyRoY| PLATE XVIII Wy “ \ NY S < a .\ EN ff FHOTO “ITH. BRITTON & REY, SF. mu | TIM rm IATITTTC CALUEHDORTUS MACROCARFPUS. ui i) } ean phen hay Pi eh 158 CALIFORNIA ACADEMY OF SCIENCES. EXPLANATION OF PLATE XIX. Petals of Calochortus species. Veep Leal oD MND C. amabilis. C. benthamt. C. maweanus. C. amenus. C. albus. C. vesta. C. gunntsoni. C. longebarbatus. C. nuttallit. C. venustus. C. venustus roseus. C. cataline. C. howellit. C. dunntt. C. clavatus. C. howellii, flower. C. dunnit, flower. C. clavatus, flower. 7 fF ‘on aT | ip r wich” e Cae Ae Pay e - PrRoG.CAL ACAD. SC1.37 SER Bor. Vou Il. [Puroy| PLATE XIX PHOTO -LITH BRITTON & REY, S.F oe he ae ty per i ee >? 27 L901 23, i vf re sf IT R mE. LLIA WI PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES TuirRD SERIES BoTANY Vou.) I, Ne. 5 A Group of Western American Solanums ers uEw YOR poTANte : GARDE BY his eens Issued October 23, 1901 SAN FRANCISCO PUBLISHED BY THE ACADEMY I9OI A GROUP OF WESTERN AMERICAN SOLANUMS. BY S. B. PARISH. Tux first notice in botanical literature of any of the group of Solanums which is the subject of the present paper was in 1826, when in the Memoirs of the Academy ot. ot. Petersburg the name Solanum umbelliferum was bestowed by Eschscholtz on a plant from ‘ Nova California.’? As the original description is not easily accessible, a transcrip- tion of it, for which I am indebted to Dr. J. N. Rose, is subjoined in the foot-note.’ The floral characters here assigned are common to a whole series of plants that the student of Californian botany finds it unsatisfactory to unite, and as difficult to separate on stable lines; nor are the vegetative characters defined with sufficient accuracy to enable one to determine which of these plants the author had in hand. aa 14. Solanum umbelliferum /sch. Solanum umbelliferum Escu., Mem. Acad. Petersb., Vol. X, 1826, p. 283. ‘““(Tnerme, foliis integerrimis, calycibus quinquedivisis, staminibus zequali- bus, inflorescentia terminali.) ““Caule suffruticoso, erecto, foliis ovalibus, acutis, integerrimis, pubescenti- bus; umbellis terminalibus. “In fruticetis Novee Californiz. “Caulis orgyalis, suffruticosus, fistulosus, angulatus, pubescens; ramis sub- herbaceis, nutantibus, tomentoso pubescentibus. ‘‘Folia alterna, petiolata, ovalia, acuta, integerrima, utrinque pubescentia, vix pollicaria, caulina interdum late ovata sesquipollicaria. ‘Flores terminales umbellati; umbella plerumque quadriflora, interdum bi- vel-triflora; involucrum parvum urceolare, integrum, pubescens; pedunculi zequales, elongati, pubescentes. Calyx urceolaris quinquefidus pubescens, laciniis acutis. Corolla calyce triplo major, dilute violacea, quinquefida, extus pubescens. Antherze flavee. ‘‘Bacca magna purpurea.” The “‘involucrum parvum’? is, of course, the organ better designated by Dr. Gray in the Synoptical Flora as a ‘‘cupulate node’ at the insertion of the pedicels on the peduncle. [159] October 23, 1901. 160 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. Dunal (1852), in De Candolle’s Prodromus, substantially repeated the above diagnosis for the original species, and added to it two others. The significant points in the descriptions of these species are that the one, S. genis- lotdes, is characterized as ‘‘ramulosissimts pilosis,’’? and the other, S. californicum, as ‘‘ tomentosis candicantibus.’’ ‘The first of these was founded on a specimen collected in Cali- fornia by Douglas, and the latter on another plant from the same region and collector, and one of Pavon’s from ‘‘Nova Hispania.’”’ In 1876, Dr. Gray, in a contribution to the Proceedings of the American Academy, reduced Dunal’s two species to the original Eschscholtzian S. wmbelliferum, making the branched hairs of the pubescence the essential character of that species, and at the same time proposing a new species, S. wanti, for the reception of certain plants which had since come to hand, and in which the pubescence was of ‘‘simple and few-jointed hairs, some of them glandular.’’ No type is specified, but reference is made to specimens collected by Xantus de Vesey, Bigelow, Anderson, and Lemmon. A variety, S. xanti wallacez, was proposed for a plant collected on Santa Catalina Island by Wallace. There are preserved in the Gray Herbarium of Harvard University authentic specimens of all the above species, those representing Eschscholtz’s, and Dunal’s types coming from the Herbarium of Trinity College, Dublin. By an examination of these it is possible to ascertain exactly what were the plants intended by the different authors, and what more recently collected material may be included with them. A specimen collected by Hartweg is accompanied with a note in Dr. Gray’s hand, certifying it to be an original of Eschscholtz’s S. umbelliferum. It has a stem moderately hirsute (but not canescent) with a mixture of unbranched and few-branched hairs in about equal proportion. The leaves are ovate, obtuse at base, about two centimeters long, sparsely hirsute, the hairs short and mostly unbranched. All the hairs are unilocular and not glanduliferous. Bot.—Vot. Il.] PARISH—WESTERN AMERICAN SOLANUMS. 161 An original Douglasian specimen represents S. gents- totdes. 'The stems are slender; the leaves few, somewhat fascicled, minute (5-6 mm.), and ovate; the hairs on the peduncles are mostly branched, on other parts of the plant they are entire, or a few once-branched. As in the first species, all are one-celled and glandless. The specimen has the appearance of coming from a starved plant, as Dr. Gray suggests. Coulter’s No. 590, also from Trinity College Herbarium, is a like form. A specimen of the plant of Douglas, on which S. calt- fornicum was based, has stout stems canescently tomentose, with hairs all of which are branched. The leaves are broadly ovate (2-4 cm.), cuneate at base, and sparsely hirsute with mingled unbranched and few-branched hairs. The struc- ture of all the hairs is as in the other species. A specimen by Fremont, mounted on the same sheet, is a better repre- sentative of this form, and has like characters. The plant collected by Xantus de Vesey, No. 73, at or near Fort Tejon, California, may be taken as the type of the species which bears his name. It has oblong leaves (circ. 3 cm.) mostly acute at summit and cuneate at base, except a few leaves lobed and truncate at base. The whole plant is moderately hirsute with rather short plurilocular hairs. Anderson’s, Lemmon’s, and Bigelow’s plants are on the same sheet. The first is quite like that of Xantus; the other two are mere fragments with similar pubescence and oblong leaves, which are obtuse or subcordate at base, and probably were taken from plants whose lower leaves were cordate. Wallace’s Santa Catalina Island specimen (type of the variety wad/ace?) is likewise fragmentary. It consists of the summit of a branch with an ovate leaf (9 cm.) and a few young leaves, and an umbel of flowers with large corollas (4 cm.); the pubescence of the stem is of long tawny hairs, multilocular and viscid-glandular; the leaf is nearly glabrous above and tawny-villous below. All the hairs appear to be unbranched, but other specimens commonly show some few-branched hairs. 162 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. From the above notes on these type specimens it will be seen that the character which has been relied upon as dis- tinctive of S. wmbelliferum, namely, ‘‘ hairs branched,’’ is only partially applicable, not one of them being without a considerable admixture of unbranched hairs. Indeed, I have not been able to find a single specimen of this species, among the many examined, in which more or less unbranched hairs could not be detected. On the other hand, many plants are found with a few, often a very few, branched hairs intermixed with the prevalent simple hairs. More- over, the character of the pubescence, at least in most specimens of .S. wmbelliferum, is different on the various parts of the same plant. On the young stems, notably towards the tips, it is exclusively, or nearly so, of many- branched hairs, while on the leaves, notably on the older ones, it is largely, sometimes exclusively, of unbranched hairs." A more satisfactory character is found in the structure of the hairs. In S. wmbelliferum these, whether branched or unbranched, are without cell divisions and are not glanduliferous. In the plants which have been referred to S. wantz the hairs consist of elongated cells, some of the cells usually evacuate and collapsed, or atrophied, so that the hairs have a peculiar unevenness; many of them are tipped with black globular glands, causing the plant to be more or less viscid. In glabrate forms the hairs are very short and mostly reduced to a single cell, but they remain glanduliferous. Unfortunately, this character is not entirely constant, and it is possible to find specimens on which there are branched hairs which are also plurilocular and glanduliferous. The leaves of the group exhibit a wide range of variation in shape, passing from orbicular to oval, oblong, elliptical, and even lanceolate; the apices are either acute or rounded, 1JIn a specimen collected by Kellogg and Harford near Bear Harbor, on the northern coast of California, a third form of hair occurs on the oldest stems. These are densely tomentose with the usual mixture of branched and unbranched hairs, from which stand out scattered spinose branched hairs, 2-4 mm. long. Bor.—Vot. II.] PARISH—WESTERN AMERICAN SOLANUMS. 163 the bases attenuate, cuneate, rounded, truncate, or cordate. Often a part of this range of variability may be seen ona single plant, and series of specimens can be arranged readily passing by intergradations from one extreme to the other. A tendency to segmentation is manifested by the occasional occurrence on a specimen of a few leaves with a pair, or even two pairs, of basal lobes. The leaves are either smooth or papillose-roughened beneath the pubes- cence, and are either entire or crenate margined, the latter character appearing to be constant and of some diagnostic value. The floral characters are practically the same throughout the group, except that S. wal/acez usually has a narrower and deeper lobed calyx than the others. The corollas are from two to five centimeters in diameter, and in color vary from light to very dark violet, the centers having green markings; the long, bright yellow anthers are sagittate at base, and are on very short, stout filaments; the style exceeds them one-half to one-third its length, is usually straight, but the included portion is sometimes bent. The fruit is very little known; it may possibly afford some satis- factory characters when better understood. It is a smooth, globular, many-seeded berry, about two centimeters in diameter, or perhaps sometimes smaller, In S. wadlacez it is certainly dark purple when ripe, and this color has usually been assigned to the fruit of all the members of the group; but according to Professor Greene’ the ripe fruit of S. umbelliferum is ‘‘yellow;’’ that of S. xantz I have never seen more than a light or whitish green when apparently ripe. Such being the generally inconstant characters of this group of plants, it is evident that their satisfactory segrega- tion is a matter of no little difficulty. The exercise of that botanical industry which multiplies ‘‘species’’ by the min- ute description of individuals might reap here an abundant harvest. On the other hand, a rigorous insistence on sharp 1 Man. Bay Region, p. 268. 164 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. and absolute delimitations, which should exclude all inter- grading forms, would reduce the whole group to a single heterogeneous species, a disposition, it is safe to say, unac- ceptable to any one who has studied these plants. Avoid- ing these two extremes, I have attempted a classification with reference to certain diverging lines of development which are manifested, without insisting upon definite cleav- ages, which do not exist. Such differences, as every botanist is aware, are more easily perceived in the exam- ination of copious material than characterized. In the present instance the facts as they are in nature are repre- sented by such a treatment; for we have here a series of plants in which variation has outrun the processes of selec- tion, and in which the connecting forms yet remain to unite the diverging lines of evolution. It has also the advantage of preserving the species recognized by Dr. Gray, who with his accustomed discernment fixed upon the best diag- nostic character, that of the pubescence, although it does not possess the definiteness he seems to have supposed. To the group I have prefixed an Arizona plant with a very different corolla, but with further characters shared by the other members of it. KEY TO THE GROUP. Corolla deeply 5-cleft; nodes of the peduncle obsolescent. 1. S. arizonicum. Corolla angulately 5-lobed; nodes cupulate, prominent. Leaves mostly linear-lobed at base. 2. S. tenutlobatum., Leaves seldom lobed. Plants viscidulous; hairs unbranched. Leaves crenate; corollas large. Stems long-hirsute. 3. S. wallacet. Hispidulous or glabrescent. 4. S. wallacei viridis. Leaves with entire margins. Acute or merely obtuse at base. 5. S. xanti. Cordate or subcordate at base. 6. S. xanti intermedium. Attenuate at base, small. 7. S. xantt glabrescens. Plants not viscid; hairs branched. Stems villous. 8. S. umbelliferum. Stems canescently tomentose. 9. S. umbelliferum californicum. Bot.—VoL. II.] PARISH—WESTERN AMERICAN SOLANUMS. 165 * Suffrutescent or suffruticose plants; peduncles lateral, or by the suppres- sion of the growing apex apparently terminal; styles clavate; fruit a many- seeded berry. + Corollas small, 5-cleft; peduncles slightly thickened at the articu- lation of the pedicels. Vv 1. Solanum arizonicum. Barely suffruticose or even herbaceous; stems 3 m. high, not striate or angled, pubescent with unbranched hairs, the upper part canescent, as are the lower surfaces of the leaves; leaves ovate-lanceolate, 2-3 cm. long, prom- inently anastomose veined, the lower half of the margins coarsely toothed; flowers in small corymbs (about 7-flowered); peduncles surpassing the leaves; pedicels short, 2-5 mm.; calyx 3 mm. high, the lobes ovate; corolla light purple, pubescent without, 5-6 mm. wide, 5-cleft nearly to the base into ovate-acuminate lobes; anthers 3 mm. long, on filaments 1 mm. long; style hirsute below; fruit not seen. Hlabitat: ot Springs, Arizona (397 Toumey, June 17, 1892 [N]).! + + Corollas rotate, angulately 5-lobed, violet, with green markings at base; peduncles thickened into a cupulate node at the articulations of the slender pedicels. ++ Pubescence of several-celled, unbranched hairs. V 2. Solanum tenuilobatum. Suffrutescent, stems slender, angled, glabrescent below, hirsutulous above with short, one- to several-celled, non-glanduliferous hairs; leaves linear to narrowly oblong, 2-3 cm. long, the midrib prominent, all but the uppermost with a pair of hastate linear lobes at base; umbels 1-4-flowered; corolla 12-15 mm. wide; fruit not seen. Flabitat: Mexico,—Lower California (probably near Ensenada, C. C. Parry, April, 1882, type [G]); (Carrizo Creek, Brandegee, April 19, 1893 [A]). 1 The letters in brackets denote the herbaria in which specimens are deposited. [A] Herbarium California Academy of Sciences, San Francisco; [G] Gray Herbarium, Har- vard University; [N] National Herbarium, Washington; [P] Herbarium of S. B. Parish, San Bernardino, Calif.; [U] Herbarium University of California. It isa pleasure to record my thanks to Miss Alice Eastwood, Dr. B. I,. Robinson, Dr. J. N. Rose, and Dr. Willis L. Jepson, for the opportunity of examining the collections of which they are the custodians. Iam also under obligations to Mr. H. M. Hall for speci- mens and other favors. 166 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. vA 3. Solanum wallacei. Solanum xanti wallacei Gray, Proc. Am. Acad., Vol. XI, 1876, p. 90; in Brew. & Wars. Bot. Calif., Vol. I, 1876, p. 539; Syn. FI., Vol. II, Pt. 1, 1878, p. 229. GREENE, Bull. Cal. Acad. Sci., Vol. I, 1885, p. 226; id., Vol. II, 1887, p. 408. Lyon, Bot. Gaz., Vol. XI, 1886, pp. 204, 334, 336. BRANDEGEE, Zoe, Vol. I, 1890, p. 143. FRANCESCHI, Zoe, Vol. IV, 1893, p. 137. Davrpson, Pl. Los Angeles Co., p. 21 (1896). TRASK, Erythea, Vol. VII, 1899, p. 140. Solanum xanti WATSON (not Gray), Proc. Am. Acad., Vol. XI, 1876, p. 117. Suffrutescent, ‘‘ often forming round masses;’’ stems about a meter long, densely tawny-villous with long, multilocular, viscidly glanduliferous hairs which are unbranched, or usually a few once-branched; leaves thickish, sometimes pustulose, usually less densely villous than the stems, crenate margined, the lower ample, cordate, the upper ovate, rounded, or subcordate at base; calyx narrowly funnel-form, deeply cleft, or wider and less deeply divided; corolla 2-4 cm. wide; style glabrate, or villous below; ripe fruit dark purple. Flabitat: Islands off the Coast of California and Lower California, and near the seacoast in Central California. California (Brandegee [G]); Central California (429 Palmer [A]); Los Angeles County (Santa Catalina Island, Wallace, type [G]; 76 Lyon, 1885 [G]; McClatchie, Nov., 1893 [Ps Mrs. Trask; Dec., 2804); Ni) Santa Barbara County (Santa Cruz Island, Greene, 1886 [A]; U.S. S. Albatross, Feb., 1889 [A]) (Santa Rosa Island, Brandegee [A]) (Santa Barbara [no collect- or’s name] Jan., 1892 [A]) (Santa Inez Mountains, Brande- gee, 1888 [A]}); San Louis Obispo County (Mrs. Blochman, May, 1893 [A]; 428 Palmer, June, 1876 [A]); Marin County (Miss Eastwood, Oct., 1896 [A]). Mexico (Guadaloupe Island, 62 Palmer, 1875 [G]; Greene, April, 1885 [P]; Franceschi, Jan., 1893 .[A (much reduced form), N, P, U]). ¥ 4. Solanum wallacei viridis. Stout, erect, glabrate, or above hispidulous; hairs mostly reduced to a single cell; leaves ovate, cordate, or rounded at base; calyx cup-shaped, with short lobes. Bot.—Vot. Il.] PARISH—WESTERN AMERICAN SOLANUMS. 167 Habitat: Central California, near the coast. California (586 Coulter [G]); Monterey County (Pacific Valley, Miss Eastwood, May, 1897, type [A, P]) (Santa Lucia Mountains, Willow Creek, R. A. Plaskett, Feb., 1898 [A]); Marin County (Mt. Tamalpais, Miss Eastwood, May 30, 1896 [A}). V 5. Solanum xanti Gray. Solanum xanti Gray, Proc. Am. Acad., Vol. XI, 1876, p. 90; in BREW. & Wats. Bot. Calif., Vol. I, 1876, p. 539; Syn. FI., Vol. II, Pt. 1, 1878, p. 229. Davipson, Pl. Los Angeles Co., p. 21 (1896). McC tatcuiE, FI. Pasadena, p. 641 (1895). | CoviLLE, Death Valley Rept., pp. 167, 251 (1893). Stems slender, 3-10 dm. long, woody, or at high altitudes herbaceous from a lignescent base, the younger angled, moderately villous, with many-celled, unbranched hairs, most of them gland-tipped; leaves ovate, ovate-oblong to oblong-lanceolate, 1-4 cm. long, acute or obtuse at base; corolla 1-2 cm. wide; mature fruit apparently light green in color. ‘ Hairs with some of the cells atrophied, and sometimes a few are once- branched. Habitat: Throughout California, except in the desert region, ascending to 6,500 feet altitude in the mountains, and reaching the borders of Arizona and Lower California. California (428 Palmer, 1876 [N]; 186 Thomas Bridger [N], a canescently tomentose form); central California (429 Palmer, 1876 [N]) Sierra Nevada Mountains (Lemmon, 1875 [N]); Alameda County (Piedmont, F. W. Koch, March, 1895 [U]); Calaveras County (1358 Davy, May, 1898 [P, OIE ‘‘Viscid to the touch, herbage malodorous;’’ leaves thin, neurose) ; Fresno County (Fort Miller, Heermann, July, 1853 [N]) (Toll House, 2,050 feet altitude, 7 Hall & Chandler, June, 1r900 [P]) (Pine Ridge, 5,000 feet altitude, 93 Hall & Chandler, June, 1900 [P]) (Dinkey Creek, 5,300 feet altitude, 355 Hall & Chandler, June, 1900 [P]) (North Fork King’s River, 6,000 feet altitude, 448 Hall & Chandler, July, r900 [P]); 168 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. Inyo County (Willow Creek, 789 Coville & Funston [G, N, P]); Kern County (Fort Tejon, 73 Xantus, 1857-8 [G, N]); Los Angeles County (San Gabriel, 108 Brewer [G], the specimen in [U] under this number is of the var. glabrescens) (Borders of Mojave Desert, Antelope Valley, Pringle, May, 1882 [N]) (Elizabeth Lake, 1888 Parish, June, 1887 [P]) (Saugus, Brandegee [A]) (Antelope Valley, 2343 Davy [U]); Mariposa County (Yosemite Valley, Mrs. Dodd [U]); Mendocino County (Eel River, W. G. Wright, 1894 [A]); Placer County (Truckee River, July, 1886, 49 Sonne [A]; 398 Sonne [P]); Plumas County (Mrs. Austin [G]); San Bernardino County (Bear Valley, altitude 6,500 feet, H. M. Hall, July, 1899, [P]; 3382 Parish, June, 1894 [N]) (Bloomington, Parish, March, 1897 [P]) (Reche Cafion, Parish, 1897 [P| )'s Santa Barbara County (Bartlett Cafion, 131 Rothrock, 1875 [G]); Santa Clara County (Saratoga, 254 Davy, Sept., 1893 [U]}): Sierra County (Webber Lake, Lemmon [G]); Sonoma County (Freestone, Miss Eastwood, March, 1899 [A]); Tulare County (Mineral King, Brandegee [A]). Arizona (Palmer, 1869 [N]) (Central Arizona, 427 Palmer, 1876 [N]) (Fort Apache, 607 Palmer, June, 1890 [G, N]). Mexico,—Lower California (Ensenada, 3711 Jones, April, 1882 [G], a transition to var. glabrescens) (San Pedro Martir, Brandegee, May, 1893 [A]). v 6. Solanum xanti intermedium. Solanum xanti CoviLLe (not Gray), Death Valley Rept., p. 257 (1893). Stems woody, lax, up to 2 m. long, viscid, leaves cordate to oblong, at least obtuse at base, 3-15 cm. long; corollas 2-4 cm. wide. Few-branched hairs are often present, indicating a transition to S. wmbel- liferum; while in size and shape of leaf this form passes into S. wadlacet through its variety vzr7dis. Bot.—Vot. Ul.] PAR/ISH—WESTERN AMERICAN SOLANUMS. 169 Habitat: California, from Sonoma County southward, chiefly in the foot-hills, but ascending the mountains to 8,000 feet altitude in Southern California. * California (Bigelow [N]; Chas. Sayre, 1875 [N]); Kern County (Havilah, 1064 Coville & Funston, June, 1891 [N]); Los Angeles County (Cucamonga, Bigelow, 1853-4 [G]) («* Cahuenga Pass,” 189 Brewer LG iC san thes nando Valley,’’ 189 Brewer [U]) (‘‘ San Fernando Plains,”’ 207 Brewer [U]) (‘‘ Santa Susana Mountains,” 207 Brewer [G]) (Compton, McClatchie, 1897 [P]) (Pasadena, Mrs. Brandegee [A]); Marin County (Redwood Canon, Miss Eastwood, March, 1896 [U]): Monterey County (Santa Lucia Mountains, 440 Gack. Vasey, 1880 [N]; 20 R. A. Plaskett, Feb., 1898 [N]); Napa County (Jepson, May, 1897 [P, U]); Riverside County (Santa Ana River, 141 H. M. Hall, May, 1895 [P]); San Bernardino County (San Bernardino, 441 G.R. Vasey, 1880 [N]); 4388 Parish, May, 1897, type [G, N, P, U]) (San Antonio Mountains,—Lytle Creek, altitude 5,750 feet, H. M. Hall, June, 1899 [N, P]; Swarthout Cajion, altitude 6,500 feet, H. M. Hall, June, 1899 [G, N, P, UJ); Santa Barbara County (Santa Inez Mountains, G. W. Dunn, May, 1891 [A]; Sonoma County (191 Samuels [N], a form with shorter hairs) ; Tulare County (Long Meadow, altitude 8,000-9,000 feet, 206 Palmer, June, 1888 [N]). . 7. Solanum xanti glabrescens. Stems woody, slender, 10-15 dm. long, glabrate, or above hirsutulous with short, mostly one-celled hairs; leaves smaller (2-6 cm.), oblong, elliptical or lanceolate, mostly attenuate or acute at base; corolla 2 cm. wide. Habitat: From southern Oregon, throughout California (excepting the desert region), to northern Arizona and Lower California. Also doubtfully reported from New 170 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. Mexico. This is the most widely distributed form, but apparently confined to lower altitudes. Stems usually lax, and seeking support from other shrubs, but in open ground, _notably near the coast, forming low, compact clumps. - Oregon (Josephine County, Howell, May, 1884 [G]). California (294 Fremont, 1846 [G]; ‘‘R. N. A.’’ 1896 [N]) (Sierra Nevada Mountains, Lemmon, 1875 [N]) Southern California (285 Parry & Lemmon [G}]); Alameda County (Berkeley, April, 1900, H. M. Hall [P]); Butte County (Clear Creek, 191 H. E. Brown, April, 1897 [A, N]) (Little Chico, Mrs. C. C. Brown, April, 1897 [A]); | Calaveras County (Mokelumne Hill, 83 Blaisdell [A}) ; Colusa County (Epperson’s, Mrs. Brandegee [A]); Lake County (Mrs. Brandegee, July, 1884 [A]) (Snow Mountain, Mrs. Brandegee [A]); Los Angeles County (San Gabriel Canon, 108 Brewer [U]) (Compton, McClatchie, 1896 [P]); Modoc County (Goose Valley, M. J. Baker [U]) (Little Hot Springs Valley, Baker & Nutting, July, 1894 [U]); Monterey County (Santa Cruz, 2223 M. E. Jones, June, 1881, in part [A]); Napa County (Zem Zem, Jepson, July, 1892 [U]) (Vaca Mountains, R. H. Platt, March, 1898 [-A]); Placer County (Mrs. M. M. Hardy, 1893 [A]) (Apple- gate, Mrs. H. Smith [A]); Riverside County (San Jacinto River, 3115 Leiberg, March, 1898 [N]); San Bernardino County (San Bernardino, 4384 Parish, May, 1897, type [G, N; P, U]); San Diego County (San Diego, Cleveland, 1874, 1875 [G]; April, 1881 [P]; Dec., 1883 [A]; Mrs. Brandegee [A]; Greene, March, 1885 [A], a very leafy form; Miss Cummings, April, 1896 [G]) (Alpine, Mrs. Brandegee [A]) (Temecula Canon, Greene, 1885 [A]) (Fallbrook, Parish, Nov., 1891 [P]) (Witch Creek, Alderson, May, 1894 [P]) (San Isabel, A. W. Henshaw, April, 1893 [N]; H.M. Hall, May, 1899 [P]) (Oceanside, 4437 Parish, June, 1897 [A, G, N, P, U], a compact, maritime form) (‘‘ Southwestern Bot.—Vot. II.] PARISH—WESTERN AMERICAN SOLANUMS. 171 part of Colorado Desert,’’ Orcutt, April, 1889 [N], but an error in locality is probable) ; Siskiyou County (Yreka, 877 Greene, June, 1876 [G]). Arizona (Fort Mojave, Cooper [G]). New Mexico (‘‘Chiefly in the Valley of the Rio Grande, below Dojia Ana,’’ rorr Mexican Boundary Survey [N], perhaps an error of locality). Mexico (Lower California, Pringle, April 6, 1882 [N)]. A transition to S. tenuzlobatum). ++ ++ Pubescence of one-celled hairs, at least those of the stems mostly many-branched, not gland-tipped. va 8. Solanum umbelliferum Zsch. Solanum umbelliferum Escu., Mem. Acad. Petersb., Vol. X, 1826, p. 283. Duna_ in DC. Prodr., Tome XIII, 1852, p.86. Gray, Proc. Am. Acad., Vol. XI, 1876, p. 90; in BREw. & Wats. Bot. Calif., Vol. I, 1876, p. 539; Syn. Pi, Vol. Ll, Pt. 171878, p..220; Solanum genistoides DUNAL, in DC. Prodr., Tome XIII, 1852, p. 86. Suffrutescent or suffruticose, stems slender and erect, 1 m. or more in length, moderately hirsute, the hairs glandless and without cell divisions, mostly branched, but some simple, and these often predominating on the leaves; leaves thin, ovate to oblong, obtuse or somewhat acute at base. flabitat: Coast counties of central California as far south as Santa Barbara. A doubtful form from Mexico. = California .(Hartweg, type [G]; 587 Coulter [G]; Bloomer [G]; Kellogg [G]; Douglas, type of S. genzs- tordes |G]; 590 Coulter, same form [G]); Alameda County (Berkeley, Greene [A]; McLean [U]) (Sunol, Congdon, May, 1892 [P]; Jepson, March 9, 1900 [P]); Mendocino County (Bear Harbor, 717 Kellogg & Harford, July, 1869 [N]); Monterey County (Fremont, Jan. 31, 1846 [G]); San Francisco County (Bigelow, 1853-4 [G]; Kellogg [G]; Bolander, 1866 [N]); San Mateo County (Bolander, 1892 [G]) (Crystal Springs, Miss Eastwood, April, 1896 [A, U]); Santa Clara County (Stanford University, 104 C. Rutter, Feb. 10, 1892 [N]); SN aM ts ps Clg SS ate Uhl ae emia ier ieee Race Bh i sd a " 172 CALIFORNIA ACADEMY OF SCIENCES, [PRoc. 3D SER. Santa Cruz County (Santa Cruz, 2223 M. E. Jones, in part [A]) (Glenbrook, Santa Cruz Mountains, H. Davis, April, 1899 [A]). Mexico (San Martin Island, 30 Anthony [G, N]. Nearly leafless, leaves small, orbicular, trifoliately lobed; probably distinct). v 9. Solanum umbelliferum californicum. Solanum californicum DuNAL, in DC. Prodr., Tome XIII, 1852, p. 86. Solanum umbelliferum GREENE (not Escu.), Man. Bay Reg., p. 267 (1894). Stems stout, erect, densely and canescently tomentose with many-branched hairs, those of the leaves sometimes in part unbranched; fruit said to be ‘‘vellow”? when mature. Habitat: Coast Mountains from San Francisco to Santa Barbara, and possibly on the borders of Nevada. California (Douglas, type [G]; Fremont, 3d Exped. [G]; 589 Coulter [G]; Brandegee [G]) (Santa Maria Mountains, Mrs. Watts [A]); Contra Costa County (Oakly, March, 1900 [P]); Fresno County (Alcalde, Mrs. Brandegee [A]) (Huron, Miss Eastwood, May, 1893 [A]); Monterey County (Monterey, Fremont, 3d Exped., Jan. 31, 1846 [N]; 633 Brewer [G]; 633 Guirardo [U]; Bran- degee [A]) (Pacific Grove, Tidestrom, Jan., 1893 [U], a stout form with ovate leaves, 3-4 cm. long, same as Guirardo’s plant) (Nacimiento River, Miss Eastwood, May, 1897 [A]) (Los Burros Trail, Miss Eastwood, May, 1897 [A]); | San Benito County (New Idria, Miss Eastwood, May, 1893 [A]):; San Francisco County (Lone Mountain, 14 Kellogg & Harford [A]); . Santa Barbara County (San Rafael Mountains, H. C. Ford, 1887 [G]) (Howard Cajion, Miss Eastwood, May, 1896 [A], albino) (Duford’s Ranch, Miss Eastwood, May, 1896 [A]) (Sespe, May, 1897, F. W. Hubby [P]). Nevada (Carson City, Anderson, 1865 [G], but an error in the locality label is probable). America, wary 4, 1902 A al Sele a as Ges an HE ACADEMY Fs a agra Nay te z ie % LICAT B PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES THIRD SERIES BOTANY Von. Ti Nor, 6 r An Account of the Species of Porphyra Found on the Pacific Coast of North America BY rad be Hews LT, A. .ius NEW YGH« eROTANES AL, Sanne ™ Witru THREE PLATES Issued January 4, 1902 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1902 AN ACCOUNT OF THE SPECIES OF PORPHYRA FOUND ON THE PACIFIC COAST OF NORTH AMERICA. BY HENRI T: AL BUS.” CONTENTS. PLATES XX-XXII. DRE UISTOR VE wa corastoskathas sl satel wore sible coke slateutie stele s cheaper e enter e 173 UU MORPHOL OG Vite vie cioteias arereid (e acavenscis Tate ote de 4 Saleem ae tev as eS 175 DELP REDISTRIBUTION: seis eens aes chose staie (x terest ioteia sects rales oor ate Lanter ee ae amen ee 192 TV. | DESEREPTION OR: OPHGIBS! SLi: o5)s. da lujee see bacaltnte eve eee 6 oe mnare lead 195 KEyiro THE PAGIRIC COAST SPECIES)! (.4icc (case wel oper 195 Nis CBCONOMIG) WISE SIAM: W15 crta ved oaitin we eiskrael Gelatts qetskes wie snbe eeewaews 230 WEE IME TELODS wate has cyita lobule Soipseteus Socks adel niaria atctehstereierstcraravere io Gl ste neaepenan ai 231 EVMBRATURE WAND EX SICCAT Ae @1TED \cticysrseasielan slseiel oalcterecabielel ste) cue ie ene terets 234 EXPLANATIONMORSELATES | 4.1 fo'b cise: s ox Mists cia cite aicts) aisteielelalete a) ose hstan epee 236 I. HustTory. From a systematic point the genus Porpfhyra has been scantily dealt with. Created in 1824, by C. A. Agardh, to contain those species of ‘‘ U//va’’ which possess a red color- ing matter, it has since been fully treated by but two authors—J. G. Agardh (1882) and J. B. de Toni (1897). The former author was the first to distinguish between monostromatic and distromatic species. ‘This idea was car- ried still farther by Kjellman, who in 1883, in his ‘‘Alge of the Arctic Sea,’’ distinguished between the genus Por- phyra and the subgenus Dzploderma, the latter to include all distromatic species. The name Diploderma was changed by de Toni (1897) to Wildemanza. L. Kolderup Rosenvinge (1893) used the name /Por- phyra for both monostromatic and distromatic species, retaining Diploderma as a subgeneric name. ‘This appears *Contributions from the Botanical Laboratories of the University of Cal- ifornia and presented in partial fulfillment of the requirements for the degree of M.S., May, 1899. Prepared under the direction of Professor W. A. Setchell. [173 ] December 14, 1901. 174 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. to be a step in the right direction, since, as Rosenvinge points out, the distromatic species are frequently monostro- matic in portions of the frond. The writer has often found specimens of P. miniata, P. tenuissima and P. abysstcola which were partly monostromatic, partly distromatic in the purely vegetative portions of the fronds. Even though the distromatic character is far more con- stant than was originally supposed, it seems to the writer there exists no sufficient reason to subdivide the genus, since the plants agree so entirely in habit and external characters as to be readily recognized by the collector as belonging to the genus Porphyra. Other contributors to our knowledge of the genus Por- phyra are Foslie (1890) and Strémfelt (1886), both of whom chiefly investigated the waters of Northern Europe. The species of the genus Porphyra in Asiatic waters have been little studied. Suringar (1870) mentions P. vu/- garis as occurring in Japan. Afterwards Kjellman (1897) studied the species of Porphyra of the coast of Japan, and in his paper enumerates six new Japanese species. One of the first to mention Porphyra in America was Ruprecht (1852). In an account of a species of Phyllo- spadix collected by Wosnessenski near the mouth of the stream Slavjanka (Russian River!), he refers to a para- sitic species of Porphyra, occurring on the blades; ‘‘gegen die Blattenden zu, finden sich kleine parasitirende Exem- plaren von Porphyra.’’ From the fact that P. nazadum And. is the only species of Porphyra occurring with any regularity on Phyllospadix, it is more than probable that this is the species referred to. Harvey (1858) in his account of American Algz, men- tions but a single species, P. vulgaris, found on both the east and west coast. He is inclined to unite P. vulgares Ag., P. laciniata Ag., P. purpurea Ag., P. linearis Grev., and P. amethystea Kiitz. under the name P. vulgaris. Later, Farlow (1881) described P. /acinzata Ag. as a cosmopolitan species, and mentions P. J/eucosticta Thur. as probably occurring in New England, but not yet certainly observed. Bot.—Vot. IL] HUS—PORPHYRA. 175 Collins (1882) reported P. /aciniata from the east coast, and later (1884) P. leucosticta and P. minzata. J. Agardh (1882) reports P. coccinea (P. naiadum?) and P. perforata from the Pacific Coast. Since then Dr. Ander- son (1892) has added two new species to the number, P. naiadum and P. nereocystis. Miss Tilden in Century III of ‘‘Algze of North America”’ has distributed four species of Porphyra from the Pacific Coast under the names P. miniata, P. naiadum, P. leuco- sticta, and P. lacinzata. Up to 1898, however, there had been reported from the west coast of North America but four distinct species. In the winter of 1897 Professor Setchell suggested that the writer investigate the peculiar base of P. naradum And., but specimens of Porphyra gathered on collecting trips, and a consideration of those in the herbarium, showed such a variety of morphological and anatomical characters that the desirability of a collection and investigation of the species of Porphyra occurring on this coast became appar- ent; itis the results of these investigations which are set forth in the following paper. Il. Morpuo.oey. The shape of the fronds of the various species of Porphyra is exceedingly variable, but that of most of them can be reduced to the elongated type of frond. The variation is between linear and oblanceolate, and nearly all the species mentioned in this paper exhibit both characters at various periods of their existence. A striking exception to this rule is Porphyra perforata tf. lanceolata, which, as a rule, is constantly linear; yet there are specimens in our herbarium which are decidedly lanceolate. On the other hand, I believe there are but few mature specimens of P. fenwzssima, if any, which ever exhibit a linear form. The nearest approach the plant makes to the linear form is when young; it then-possesses an oblong outline. Closely connected with the shape of the fronds are their length and width. These three characters seem to be 176 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. closely related and are evidently determined by the same conditions, viz., age, zone, and locality. It would be diffi- cult to say which of these three agencies exerts the greatest influence on the plants. All seem to be of equal importance. A very good illustration is yielded by P. perforata. While young specimens of P. perforata (three to four centimeters long) are usually irregularly expanded, with a tendency towards the orbicular, we notice that very soon a change in the shape of the frond takes place. Asa rule, specimens five centimeters or more in length already show the type, lanceolate with undulate margin; but older specimens, and this applies particularly to those found in the lower part of the litoral zone, and in the upper part of the sublitoral zone, possess a great width and are frequently much lobed and laciniate. Especially those plants which grow on the flat surfaces of rocks, for instance on reefs, show a marked 1so- diametric development. But if the plant grows pendant from an overhanging rock, it develops the elongated type of frond. Another condition, and dependent upon locality, is the movement of the water. Plants growing where they are continually exposed to the wash of the waves, back and forth, and from side to side, show far less marked longi- tudinal development than those which are exposed to the movement of the water in but one direction. This is very well illustrated by P. xazadum And. f. major. The author had an opportunity to observe this plant growing on Zostera in the lagoon at Bolinas, Marin County, California. The lagoon, which is long and narrow, was, in the summer of 1899, protected by a high bar, so that at the rise of the tide the water flowed in very regularly for a number of hours, till the turn of the tide, when it flowed out as regularly. The blades of Zostera and the fronds of P. nazadum were bent in the direction taken by the water and the latter showed a marked elongation, so great, indeed, that the writer felt entitled to consider them a special form of the species, since they were fully twice as long as the blades of P. naiadum growing on Phyllospadix, and since there existed some other minor differences as well. The latter form was consequently designated as P. nazadum f. mznor. Bot.—Vot. II.] HUS—PORPHVRA. Ba br An interesting instance of great length attained by exposure to the motion of waves in one direction was found in a specimen of P. perforata f. lanceolata, which grew on a rock buried in the sand of the gently sloping shore of the Presidio, San Francisco, California. This specimen attained a length of 325 centimeters, which to the author’s knowledge is the greatest length ever attained by any specimen of this species of Porfhyra. This extraordinary longitudinal devel- opment (the average length is but thirty to forty centime- ters) was evidently due to the plants being stretched out at full length every time a wave rolled in or went out. P. nereocystis, growing in three to five fathoms of water on the stipes of /Vereocystzs litkeana, often attains a great length, specimens of over three meters in length having been collected at Monterey. A specimen of P. variegata collected at Santa Cruz by Dr. Anderson measured seventy- nine centimeters. On the other hand, we have found a fertile specimen of P. perforata but two centimeters long. But the plant which in its adult stage is the smallest of all Pacific Coast species of Porphyra is P. natadum, which often bears fruit when but one centimeter long. " The width of the fronds also varies considerably. While the writer has measured specimens of P. nereocystis which were fully forty-nine centimeters in diameter, some mature specimens of P. perforata f. lanceolata collected by Dr. W. A. Setchell at Monterey, California, measured in their widest part but twenty-five hundredths of a centimeter. In regard to the part age plays in the determination of the shape of the fronds, it must be said that while the younger plants asa rule possess the elongated type of frond, the older plants generally have a greater width. A microscop- ical examination reveals the fact that the divisions of the cells of the younger fronds are usually parallel to each other and at right angles to the longer axis of the frond. In the older plants, where the development is more isodiametric, we can readily recognize more or less isodiametric groups of cells, which evidently arose from a single cell. Of course, environmental conditions have much to do with this. 178 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. The base in Porphyra varies from cuneate to cordate, or even umbilicate, and is sometimes cucullate, as in P. nereo- cystis. These various forms of base depend just as much as size, etc., upon age, zone, and locality. An umbilicate base is found on the fronds of plants growing on flat sur- faces (P. laciniata f. umbzlicalis); one is liable to meet with a cordate base in the older fronds; while a cuneate base is found on the fronds of plants which grow in exposed places. In the genus Porphyra, we may distinguish between two kinds of attachment, the one cushion-shaped and paren- chymatous, the other discoid and rhizoidal. The latter form of base has been amply discussed and illustrated by Bornet and Thuret (1878), and it will suffice to say here that the cells in the immediate neighborhood of the base produce rhizoid-like projections, in thickness from two- tenths to one-tenth the diameter of the cell, which grow down through the jelly between the cells and the cuticle. That these hyphe actually grow, and that the older the plant grows the more of these projections are produced, is demonstrated by treating the base of a young specimen of Porphyra perforata with Schultze’s macerating fluid. This dissolves the jelly, and the weight of a cover-glass crushes the preparation sufficiently to show the details. From each of the thick-walled cells near the base, a hypha may be seen growing out, which may be longer or shorter, some being even but a few microns in length. Their course is more or less direct. Most of them extend down to the ~substratum. Consequently, though the frond about one centimeter above the base is normal, the part lower down is very much thickened by an ever increasing number of these projections, which finally form a dense network, in which it is impossible to trace the individual hyphe. According to Agardh (1882), these hyphez possess no septa, at least, he has been unable to see them; for he observes: ‘‘ Hec fila radicantia Porphyre mihi semper inarticulata obvener- unt, * * * .’? They are long, slender, tapering threads, averaging one to two microns in thickness. Of course, the longer they become, the less evident their tapering nature BotT.—VOt. II.] HUS—PORPAYRA. 179 is. Infact, the diameter of the larger hyphe appears to be the same for the whole length. While the majority agree in this regard, if we follow them down to the base a differ- ence soon becomes apparent. Some of the hyphe come to an abrupt end, their diameter remaining constant, the con- tents remaining hyaline and parietal, and no septa being present; but others show a greater or lesser increase of thickness at the tip for a greater or lesser length, and a few even branch or at least show indications of branching (Pl. XX, figs. 7-10). In some of these, septa have been dem- onstrated. The swollen ends contain protoplasm. Whether these ends are to be considered as haustoria, and whether the hyphe enter the cells or intercellular spaces of the host- plant, or whether they merely adhere to the substratum, are questions to which the author can give no definite reply. In sections of the base of a specimen of Porphyra perforata which grew on Phyllospadix, it was impossible to deter- mine the course of the hyphe. ‘The same was true for plants growing on wood. Young specimens of Porphyra perforata growing on barnacles were treated with one per cent. nitric acid; but after dissolving the calcium salts, it was impossible, partly owing to the confusing mass of par- asitic algz which flourished in large numbers on and in the shells, but especially to the presence of chitin, to follow the hyphz in their course. The ever increasing number of hyphe adds considerably to the thickness of the frond, the latter within one-half a centimeter of the disc often measuring two hundred microns or more, while the strength must be increased a hundred- fold. The cells which give rise to these hyphe, especially those situated more towards the disc, are but imperfectly seen, even in a specimen not quite four centimeters long, being obscured by the hypha-like projections which sur- round them. It is easy to conceive that being thus partly excluded from the light these cells should undergo some change. They lose their purplish color, have yellow- brown cell-contents, and their walls are considerably thickened. 180 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. At first it seemed exceedingly improbable to the author that the cells to which the function of the attachment of the frond was delegated should finally produce fruit; but a knowledge of the fact that the formation of spores in these cells was observed by Bornet and Thuret (1878) led him to make a more careful study of the basal cells. Up to this time, however, the author has been unable to demon- strate a single cell which both emitted a hyphal thread and. bore fruit, though the oldest obtainable specimens were investigated. It must therefore be concluded that if ever the contents of these cells are transformed into spores, this must be but rarely the case. The areolate, lighter colored portion of the frond, about one centimeter in diameter, directly surrounding the attach- ment found in all specimens of Porphyra as known to the writer, with the exception of P. natadum, probably finds its reason in two causes. The first is the partial loss of color of the cells near the base, caused by decreased activity owing to the large number of rhizoid-like projections which separate them from the surface of the frond. The other cause may be looked for in the large number of rhizoid-like projections with hyaline walls, resulting in an increased thickness and consequently increased density of the lowest part of the frond. Some of the species of Porphyra are slightly stipitate. On this coast only P. /eucosticta shows this to any marked extent. The stipes appear to possess the same structure as the discs. A cushion-shaped base is, as far as the writer is aware, found in but a single species of Porphyra, viz., P. natadum And., a species peculiar to the Pacific Coast. P. natadum has been found growing on eel-grass, either on Phyllo- spadix in exposed places in the sublitoral zone or on Zos- tera, sheltered, in lagoons. On examining the blades of Phy//lospadix during the win- ter months, we find here and there small reddish brown, cushion-shaped growths, which to the superficial gaze appear like colonies of diatoms. Continued observation shows a gradual increase in the number of these wart-like, Bot.—Vo_. II.] HUS—PORPHYRA. 181 more or less flattened structures. Finally, they cover the blades of eel-grass in such large numbers that they grow next to and over each other, and lose their natural hemi- spherical shape, obscuring the normal color of the eel-grass, and giving a rough appearance to the blade. As the season advances, examination with a lens shows a greater or smaller number of short, blunt protuberances .issuing from the wart-like growths. Under the microscope they appear to be composed of a number of cells placed end to end. Further observation demonstrates the fact that these cells, by division in two planes, give rise to a monostromatic frond. From this it is but a step to estab- lish a genetic connection between the hemispherical struct- ures on eel-grass and the fully grown fronds of P. natadum on the same host-plant. Evidently we have the prothalloid form of P. nazadum before us. This was already suspected by Dr. Setchell when he called my attention to the matter. The prothallium, when young, consists of but a single layer of cells, placed side by side on the blade of the eel- grass. For a certain length of time these cells continue to divide in a single plane. After that, division in the second plane begins to take place, gradually giving rise to the wart-like growths referred to above. In section they appear to consist of layers of large, thin-walled, parenchymatous cells (Pl. XXI, fig. 19). The cells of the central layers possess ordinary cell-contents, but only a very small chro- matophore. The two or three outer layers are made up of slightly smaller cells, and possess a large chromatophore. The cells of the layer adjacent to the surface of the blade of the host-plant also contain a large chromatophore. Upon these latter cells evidently devolves the function of attach- ing the prothallium to the eel-grass. Each cell is extended so as to form a short, sharp, unicellular rhizoid. The writer has been unable to determine with any satisfaction whether these rhizoids entered the cells of the host-plant or not. Careful sectioning and staining has failed to reveal anything of the kind. But in material which had been shrunken by reagents, only the rhizoids at the periphery of the cushion-shaped base were attached to the eel-grass, the 182 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. central part of the base having shrunk away, exposing the rhizoids. From this it would appear as if the rhizoids entered, to a slight extent only, the cuticula of the host- plant. The young fronds arise from the cells of the external layer of the prothallium. Evidently any cell may give rise to a frond by a division in one plane, in advance of the surrounding cells which form part of the external layer of the cushion-shaped base. When in this manner a filament of some five or six cells has been formed, the cells of the filament begin to divide in two planes, thus giving rise to a membranous frond, the length of which when fully grown seldom exceeds six centimeters. The number of fronds a prothallium may give rise to appears to be indefinite, every cell of the outer layer of the base seemingly being capable of producing a frond. A frond may be formed during the first stage of the existence of the prothallium, cases having been observed where frond formation evidently took place when the prothallium was but two cells in thickness. The formation of a frond by a cell of the outer layer does not mean cessation of growth for the other cells of the outer layer, since frequently a frond may be found, the base of which lies in a depression of the prothallium several cells deep. An attempt was made to ascertain, if possible, if when the frond has reached a certain size, the cells of the frond, in the neighborhood of the base, produced rhizoid-like pro- jections such as are found in the corresponding cells of other species of Porphyra. But an examination of the bases of a large number of mature fronds of P. nazadum failed to reveal these structures. When Porphyra naiadum occurs on Zostera, it produces the same wart-like growths, but only on the extreme mar- gins, not on any part of the surface of the blade, and they appear to be smaller. Other species of Porphyra probably occur but seldom on eel-grass. The only species found by the writer to occur occasionally on the same host-plant were P. perforata, P. laciniata, and P. abysszcola. Bot.—Vot. II.] HUS—PORPHYVRA. 183 The value of the prothalloid base lies evidently in the power to form a large number of fronds rapidly, which in the production of new fronds in case of accident is clearly of great importance. As far as the author is aware, P. nazadum has never been found growing on any substratum other than eel-grass, nor has a cushion-shaped base such as here described ever been found in any other species of Porpfhyra. The nearest approach to such a description is that of the base of P. coccinea J. Ag., such as is found in Agardh’s ‘Till Algernes Systematik’’ VI (1882). But judging from this account, the base is hollow, being formed by the involution of the edges of the young frond. When the frond grows older the base finally flattens out. It is worthy of note, that in connection with P. coccinea Agardh mentions a Porphyra occurring in large numbers on the Pacific Coast; it grows on seaweeds (!), and judging from the description given might possibly be P. nazadum. The color of the fronds of the different species of Por- phyra is such as to lead one to place the genus among the Floridex. But the color is far from being constant. A hundred different shades may be met with, for even the color of the different fronds belonging to one species varies; so that an attempt to describe a species by the color would be futile. While the color of one species (P. tenuissima ) is, as a rule, a delicate pink, others are a bright red or even crimson, as P. abysstcola. P. laciniata exhibits a decidedly purple color, while P. Zerforata appears mostly yellow-brown. The frond of P. variegata is crimson when sterile, while when fruiting it acquires the beautiful varie- gated appearance indicated by its name. An important fact is that the color of herbarium specimens generally changes. This was most notable in P. perforata. Specimens which when collected had a yellow-brown tint generally became a deep blue-purple. Some of the fronds of P. nereocystis underwent a change in the herbarium, while others retained the original dull brown-red color. Especially did specimens which were rough-dried and afterwards soaked in fresh or salt water for mounting 184 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. purposes seem to change color on drying. A similar change was noted in sheets of Asakusa Nori (see Economic Uses.) which were of a yellow-brown color when bought in San Francisco, but having been kept for several months in a closed paper box turned purple. A sheet of Asakusa Nori which had accidently been left partially exposed to the air and light for about a month, showed after that time a bril- liant violet coloring in the exposed portion, while that part of the sheet which was not exposed retained its original yellow-brown tint. As far as can be judged from the statements of various authors, as well as from our own observations, it appears that the color of certain species varies according to the locality. This is well illustrated by P. /eucosticta Thur. It seems that the European specimens of P. /eucosticta are of a distinctly yellow color when fresh, and when dried a deli- cate purple-pink tint. But the specimens of P. /eucosticta found on the Pacific Coast, if gathered early in the season, are deep-pink, becoming lighter as the season advances. From the above, it will be seen that specimens of Porphyra should, whenever possible, be mounted fresh; that even then the color is of small value from a systematic point of view; and that it is most undesirable, in fact, impracticable, to use the color of a frond as the criterion for the species, though it is often of great value in indicating its position. It remained for J. G. Agardh to call attention to the monostromatic and distromatic nature of the fronds of the different species of Porphyra. These characters have been found to be absolutely constant in all species, with the exception of those belonging to what we may call the ‘‘miniata’’? group, which includes besides P. mznzata, P. amplissima, P. tenuissima, and P. abyssicola. ‘The. first three species are, as a rule, distromatic, though places may be found which exhibit a monostromatic character, especially towards the edges. Fronds of P. abyssicola, which species was first described by Kjellman as monostromatic, have been found by Rosenvinge and by the author to sometimes exhibit a distromatic character, either through the whole frond or in portions of it. Bot.—Vot. II.] HUS —~PORPHYRA. 185 Kjellman applies the name Dzploderma to all distromatic species, but the above mentioned results lead the writer to agree with Rosenvinge in applying the name Porphyra to all members of the genus, while retaining Dzfp/loderma (Wildemania de Toni) as a subgeneric name for the di- stromatic fronds, the more so as in habit and external char- acter the monostromatic and distromatic species agree in all respects. Two sources of confusion in determining the number of layers in the frond exist. The first is the age of the frond, the second, the formation of reproductive cells. Young fronds of distromatic species are frequently monostromatic. This monostromatic character persists in the vegetative por- tion of the frond even after the fruit has been formed (P. abysstcola) (Rosenvinge, 1893). The same results were obtained by the author in his observations on P. abysszcola, but he cannot confirm Rosenvinge’s statement, that in the distromatic forms the inferior portion of the thallus is com- posed of a single layer of cells. ‘‘ Dans les formes distro- matiques, du reste, la partie inférieure du thalle est composée d’une seule assise de cellules’’ (Rosenvinge, 1893, p. 84). On the contrary,if we do not consider those of P. abysszcola, but few, if any, distromatic specimens were found which were monostromatic at the base. As a rule, whenever a frond of a distromatic species was partly monostromatic, the monostromatic portion was found in the region of the tip at the edge. Monostromatic species practically become distromatic as soon as the reproductive bodies begin toform. The second division of either the antheridium or sporocarp-mother-cells takes place ina direction parallel to the surface of the frond, and in this manner gives rise to two layers of cells which are often difficult to distinguish from purely vegetative cells; so that in deciding their nature, the cells of the sur- rounding tissue must be taken into account. The thickness of the fronds of the various species of Porphyra is more or less variable, and though not an abso- lutely specific character, taken in conjunction with other 186 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. characteristics it is of great value, especially in indicating the position of a sterile frond. As a rule, the thickness of the fertile part of the frond is much greater than that of the sterile part. This seems to be due to the swelling of the jelly surrounding the repro- ductive bodies at the time of ripening. Also, wherever we have a dicecious frond, or where the antheridia and sporo- carps are born on separate portions of the plant, the thickness of the antheridial portion is greater than that of the sporo- carpic portion. This would bear out our hypothesis that increased thickness is due to the swelling of the jelly, since there exists a larger number of partitions consisting of jelly between the antherozoids than between the carpospores. P. variegata exhibits a very marked thickness and strati- fication of the jelly-walls surrounding the vegetative cell. The walls of the vegetative cells of all fronds of this species examined were of this nature; consequently the author feels entitled to consider this a diagnostic character, the more so as the only other Porphyra which possesses much thickened cell-walls differs widely from P. varzegata in habit and external characters. The plant here referred to is P. perforata f. segregata. The walls not only of the vegetative cells but also of the reproductive cells are much thickened, especially those produced by the first reproductive division of the antheri- dium. In fact, the upper and lower groups of antherozoids are noticeably separated, which lends the cross-section of the antheridial portion of the frond a most characteristic appearance. The outer jelly-walls of nearly all the fronds examined were infested with bacteria which formed narrow lines per- pendicular to the surface of the frond, reminding one of the canals formed by the ‘‘spermatium’’ at the time of the fer- tilization of the ‘‘procarp’’, as described by Berthold (1882). These ‘‘ canals’’ were found in the jelly surrounding the vegetative and antheridial cells, as well as in that surround- ing the sporocarps. In regard to the shape of the vegetative cells, it may be said that while in the monostromatic fronds the cells are o Bot.—Vot. II.] HUS—PORPHYRA. 187 either cubical or more frequently higher than broad, the vegetative cells of the fronds of the distromatic species vary, as a rule, from cubical to broader than high. Excep- tions to this rule are found in P. nereocystzs, a monostromatic species which sometimes possesses cells which are broader than high, and in P. vartegata. While the vegetative cells of the younger sterile fronds of the latter species are usually square, the vegetative cells of the older fertile fronds are much higher than broad and often have a fusiform appear- ance. Judging from the fact that the vegetative cells are found between the reproductive cells, it is suggested that the shape of the former is due to pressure exerted by the reproductive cells, which before dividing gorge themselves with protoplasm, and when fully ripe swell to an abnormal size, owing to the partial dissolution of the jelly partitions separating the individual spores. During the study of the species of Porphyra of the Paci- fic Coast, the fact gradually made itself felt that the repro- ductive bodies are of the greatest diagnostic value, and that habitat, color, and thickness of frond can only be used to determine species in connection with the number of divi- sions of the antheridia and sporocarps. Since the object of this paper is merely to give a syste- matic account of the species of Porphyra of western North America, and it is not designed to throw light on the sexu- ality or nonsexuality of the genus Porphyra, the author uses the terms sporocarp and antheridium merely to indicate the larger and smaller bodies, which by some are believed to play a part in sexual reproduction, without necessarily ascribing a sexual character to these bodies. The same is true for the asexuality of the monospores. But it must be said that in no case even the slightest indication of sexuality has been observed, though many sections of sporocarps in all stages of development were examined. Neither has the author been able to observe an amceboid movement of the liberated carpospores, nor flagelliform appendages to the bodies contained in the antheridia, nor any movement on the part of these bodies, though observations were made to 188 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. determine this point if possible. The results obtained by the writer therefore agree rather with those obtained by Reinke and Bornet than with those of Berthold and others. While some species of Porphyra are moncecious, others are dicecious. In some cases fronds have been found which evidently approached closely to those of a monc- cious species, but which differed from it in being dicecious and in exhibiting some slight differences of habit, etc. Such fronds have been referred to the original species but were separated from it under a form-name. We may distinguish two forms of moncecious fronds. In the one, antheridia and sporocarps are separated in patches. These patches are usually sharply defined, especially at the edges of the frond, owing to the lighter color of the ripe antheridia and to the more intense color of the ripe sporo- carps. This arrangement is usually met with in the fronds of the monostromatic species. In the other form, the antheridia and sporocarps occur side by side, so that the frond has a uniform color. This occurs in the distromatic species, and more particularly in what the writer has found convenient to designate the ‘‘mznzata group,’’ which in- cludes P. amplissima, P. miniata, P. tenuissima and P. ? abysstcola. Vegetative cells are frequently found mixed with the spo- rocarps, among both monostromatic and distromatic species, and in larger or smaller patches. While the distromatic spe- cies show this constantly, the monostromatic species often fail to show these vegetative cells among the sporocarps. Hardly ever has the author found any vegetative cells mixed in with the antheridia of the monostromatic species. Among the sporocarps there appear frequently bodies which by various authors have been called monospores. They seem to be formed by the arrest of division in one of the segments of the sporocarp.or by one of the vegetative cells lying among the sporocarps. What differentiates them from the vegetative cell proper is a greater thickness of the cell-wall and a larger amount of protoplasm. The chromato- phore may be seen lying in or near the center of the cell. Bor.—Vot. II.] HUS—PORPHYRA. 189 These monospores can easily be distinguished from the dead vegetative cells lying among the sporocarps. The dead cells possess likewise a thick wall but apparently contain yellowish, homogeneous, highly refractive cell-contents, in which no chromatophore can be discerned. Whether monospores are sexual or not, or whether they possess any reproductive power, the author has, notwith- standing a series of careful experiments, been unable to determine. The reproductive bodies are usually first formed at the margins and gradually spread over the whole frond. Bornet even observed spores in the basal cells of P. laciniata. Under the microscope we can trace the various stages of division from the original vegetative cell to the fully ripe sporocarp. This is especially easy in the species where antheridia and sporocarps occur side by side in patches. Observation shows that each vegetative cell gives rise to a single sporocarp. The sporocarp by two more or less simultaneous divisions at right angles to each other and to the surface of the frond finally consists of four segments. In some species division proceeds no farther, and four carpo- spores are the result; but in other species, where the fully ripe sporocarp contains more than four carpospores, the cruciate division is followed by a division parallel to the surface of the frond, giving rise to eight segments, which by further cruciate division perpendicular to the surface of the frond in each of the resulting cells may give rise to thirty-two carpospores.' In the formation of the antheridia, starting from the vege- tative cell equivalent to the mother-cell of a sporocarp, there first takes place a vegetative cruciate division perpendicular to the surface of the frond, which gives rise to four anther- idial cells. Consequently, owing to this additional vegetative 1The author understands under “cruciate’”’ division, two divisions in different direc- tions, at right angles to each other and to the surface of the frond, and which are simul- taneous or nearly so. Under ‘‘transverse’”’ or ‘‘parallel” division the writer understands a division parallel to the surface of the frond. Though these divisions are, of course, but seldom if ever strictly parallel or at right angles, the use of the terms “cruciate” and “parallel”’ is a great convenience, doing away with a lengthy explanation. (2) December 1g, Igor. bee) CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. division of the antheridium-mother-cell each sporocarp cor- responds to four antheridia. The author distinguishes between a vegetative division and a reproductive division by the aid of the partition walls laid down by these divisions. A jelly-wall formed after a vegetative division is, as a rule, much thicker than one formed after a reproductive division. Furthermore, a wall of the latter kind dissolves when the frond is fully ripe, so that the reproductive bodies become arranged more or less irregularly; something which is very clearly shown in the antheridia of P. leucosticta. The first reproductive division of the antheridium is parallel to the surface of the frond, corresponding to the first transverse division of the sporocarp, and is followed by a cruciate division in both segments. In fact, parallel and cruciate divisions alternate until the number of antherozoids peculiar to the species has been formed. The only differ- ence, therefore, between antheridia and sporocarps lies, if we do not consider their origin, in the larger number of divisions which the former undergo. Though the manner of division of antheridia and sporo- carps is fairly constant, yet a large number of variations take place. The most frequent among these is the direc- tion of the last division in either antheridia or sporocarps, which is not necessarily parallel or perpendicular to the surface of the frond, but is often oblique, and is occasion- ally omitted altogether in some of the segments of the spo- rocarp or antheridium. Rarely an additional division takes place in some or all of the segments of the sporocarp or antheridium. Cases have been met with, where the vegetative division of the antheridium-mother-cell is not as evident as usual, and the whole vegetative cell apparently becomes an antheridium, so that four times the usual number of anther- ozoids are formed. Occasionally the first cruciate division of the sporocarp- mother-cell is vegetative instead of reproductive, and only one-fourth the usual number of carpospores are found. Bor.—Vor. II.] HUS—PORPHYRA. IQI With the material at the disposal of the author, he has been able to distinguish between four types of division of the reproductive bodies, the differentiation into types being based upon the number of antherozoids and carpospores produced. The first is the Porphyra perforata type. Here thirty-two carpospores are produced, the sporocarp under- going first a cruciate division, followed by a parallel division in each of the segments, which is again followed by a cru- ciate division of each segment. If we represent a vegeta- tive cell by a cube, and indicate the two horizontal lines respectively as a@ and 6, and the perpendicular as c, we can, by the aid of the formula 32 ze 2 =) readily form a diagram such as is represented in fig. 25, which shows the manner of division. The antherozoids contained in each antheridium number, in the P. ferforata type, 128, and are formed by alternat- ing parallel and cruciate divisions, the first division of the antheridium being parallel to the surface of the frond.’ The manner of division may be represented by the formula 128 (AS Ae , a) (fir. 28). "To, this. type belonewZ: perforata, P. perforata f. lanceolata, P. perforata f. segre- gata and P. nereocystis. The second is the P. /eucosticta type. Here eight carpo- spores arise from a cruciate division of the sporocarp- mother-cell, followed by a parallel division of the four segments. This may be represented by the formula WBE 8 (< Bt =) (fig. 24). The antherozoids contained in each antheridium of the fronds of species belonging to the P. deucosticta type num- ber sixty-four. They are formed by first a parallel division of the antheridium, followed by a cruciate division, after which a second parallel and a second cruciate division take 1It must be remembered that an antheridium is but one-fourth as large as a sporo- carp, the first division of the antheridium-mother-cell, which corresponds to both a vegetative cell and a sporocarp-mother-cell, being vegetative and cruciate. 192 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. place in each segment, so that the arrangement of the anther- wa %b € Dies sha =) (fig. 27). To this type belong P. leucosticata, P. laciniata, and it may be predicted, judging from the carpospores, that P. naiadum also belongs to this section, the antheridia in this species being as yet undetected. Among the distromatic forms the remainder of the types are found. First we have the P. amplissima type, with eight carpospores and sixteen antherozoids, and the form- Bab ie wa %b ¢c ule 8 (< ae 5) (fig. 24) and 16 (4 ) (fig. ozoids is represented by the formula 64 ( Brine 26); then the P. mznzata type, with four carpospores and eight antherozoids, and the formule 4 (=. & c) (fig. 23) and 8 (4 = as , ) (fig. 25). To this belong P. mzn- tata, P. tenuissima and P. abyssicola. Of the two other distromatic species which occur on the Pacific Coast, but one form of fruit has been found, so that they can hardly be brought forward as types. Ill. DuisTRIBUTION. It is almost impossible to obtain a correct idea of the dis- tribution of the older species of Porphyra, as frequently the name P. /aciniata was applied to various species which since have been separated from it. However, the author believes that it may be said with some degree of certainty that P. laciniata occurs on the western shores of Europe, from the Norwegian Polar Sea (71° N. lat.) to the Mediterranean (40° N. lat.) and on the Atlantic coast of North America from Greenland (67° N. lat.) to New Jersey (40° N. lat.). It has never been authoritatively reported from the eastern shores of Asia; for though older authors have mentioned it, yet P. daciniata was not included by Kjellman (1897) among the Japanese species, and he even expresses some doubt as to its occurrence. On the Pacific Coast of North Bor.—Vot. II.] HUS—PORPHYRA. 193 America P. /acindata has been reported from Orca, Alaska, from Yakutat, and from Amaknak Island (between 61° and 54° N. lat.). P. dactniata f. umbilicalis was reported by Professor Setchell (1899) from the Pribilof Islands. Porphyra leucosticta Vhur. does not appear to possess such a wide range as the species just discussed. It is found on the Atlantic coast of England, Germany, and France, and appears to be abundant in the Mediterranean. Collins (1884) and Holden (1897) have detected it on the eastern shores of North America, and while it may have a wide distribution on the Pacific Coast, it has as yet been reported from buta single locality, Monterey Bay, California (36° 45/ Nilat:,).. Up to the present time the members of the ‘‘mznzata’’ group, under which the author includes P. amplissima, P. mintata, P. tenuissima,and P. abyssicola, have been reported by European collectors only from the more northern latitudes (60°-80° N. lat.). On the west coast of North America these species have not been reported from so far north, ranging between 36° 45’ and 60° N. lat. P. amplissima was first found by Kjellman (1883) in the Norwegian Polar Sea, and it has since been reported from both the east and west coast of Greenland. Kyjellman did not detect this species in any of the localities visited by the Vega Expedition. Since then it has been collected at Orca, Alaska (60° 30’ N. lat.), and near Coupeville, Washington (48°. 30—,N. lat. y. P. tenuissima occurs on the shores of Norway, Iceland, and Greenland. It has never been found in the Bering Sea, but it has been collected at Sitka, Alaska (57° N. lat.). P. miniata is met with on the coasts of Norway, and attains its greatest latitude on the northwest coast of Spitz- bergen (79° 49 N. lat.). It is also found on the east coast of Norway and in Baffin Bay, whence it descends to New Foundland. There exists some doubt in the author’s mind as to the occurrence of the typical P. mznzata on the Pacific Coast. Only an extensive collection of specimens gathered on 194 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. numerous expeditions along the coast can lead to a definite conclusion. But the occurrence of a variety designated by us as P. miniata f. cuneiformis is an undoubted fact. It has been collected in the Gulf of Alaska (60° N. lat.), at Coupeville, Washington, and as far south as Monterey Bay, California (36° 45’ N. lat.). P. abyssicola is reported by Kjellman and others from the north coast of Norway and Russia and from Greenland. What the author believes to be P. abyssicola has been col- lected on the Pacific Coast at Whidby Island, Washington (48° 10’ N. lat.). Five species of Porphyra appear to be peculiar to the Pacific Coast. In some cases it is possible: to ascribe a reason for this. P. mereocyst?s, which as far as can be judged selects (Vereocystes liitkeana exclusively as its host- plant, is necessarily limited to the region of distribution of this species of WVereocystis. It has been reported from St. Paul, Kadiak Island (57° 30’ N. lat.), from Coupeville, Washington, and from the Californian shores (33°40' N.lat.). P. naiadum, growing on Zostera and Phyllospadix, seems limited to the Pacific Coast. Though other species of Porphyraoccur on Zostera, both in Europe and on American shores, yet no case is known to the author where P. nacadum was found growing on eel-grass in waters other than those of the Pacific, where it extends from Coupeville, Washing- ton (48° 10’ N. lat.) to San Diego, California (32° 20 N. lat.). P. perforata, so closely allied to P. laciniata, attains nearly the same northern latitude as the latter species, but extends far lower down the Pacific Coast. Of the two varieties of this species the author has been able to find only one, P. perforata f. lanceolata, at San Francisco and at Monterey, while the other occurs from Washington to Mexico (San Roque) (47° 30°-27° 8%’ N. lat.). P. variegata was first found by Kjellman at Bering Island (Vega Expedition). Since then it has been reported by various collectors along the Pacific Coast, from Whidby Island, Washington, to San Pedro, California (48° 10-33° 40’ IN. tat.) Bot.—Vot. I1.] HUS—PORPHYRA. 195 P. occidentalis has been found in but a single locality, Monterey Bay, California (36° 45 N. lat.). DISTRIBUTION OF THE PACIFIC COAST SPECIES OF PORPHYRA. hy| ty] Syl ty] be] be] be | Be] 8] 8) slsialsialsialglsieis Si Xs Ss Sy lees Su Sea WS S/S/2/ 8/8) S18) S$ Sls] s Y | Og 2.| a R 5 Ss 5 a) S > Si lee yee i S sh less S/S SSF) Si 81S) S18 = S/R) 81 § on Nae sal 3 a a s Sy a : . * Behring Sea. 5 ot x | * | x x | * « | Gulf of Alaska. Q Vancouver, Whidby Island, 2 * * * * ¥ * * o Seattle. 2 x |* | * San Francisco. =I erate * # | # |'% || # Monterey Bay. > * Santa Barbara. 5 * er ee ke San Pedro, San Diego. S x « | « | New England. x | x | West Europe. x | « | Mediterranean. «| «|* | * » | Greenland. a) ® |e [x x | Arctic Ocean and Spitzbergen. IV. DerscrRIPTIONS OF SPECIES. Kry TO THE PaciFic Coast SPECIES OF PORPHYRA. zt. Fronds monostromatic : 0... sc ccdiens ncn ee ee ne eeien esos snare cimes 2 fOGS CISEFOMIAUC, ecu coca Heeie sx ews ane e eciainin es soi eee Mabareyae 7 Fronds monostromatic or distromatic........---- bth Athen Mb teebearete II 2. Base cushion-shaped, consisting of parenchymatous cells... P. naiadum Base discoid, consisting of agglutinated, rhizoid-like cells...........- 3 3. Thirty-two spores in each sporocarp ....----+2eeeerersssree rete cees 4 Eight spores in each sporocarp ...-..-+sseeeree eerste cesses steers: 6 A, PPG MCE CIONIS Fis <0 aha Joie wa as os bn es elcid sin atm An mete amis aay 5 Fronds dicecious ........-----202 sees seee eee P. perforata f. lanceolata 5. Fronds brown-purple, 45-150/ HIMLC ates siete ec srrooictecterevtetanetatele P. perforata Fronds brown-purple, 6oy thick, with thick partition walls. .....-..---- P. perforata f. segregata Fronds red-purple, 25-60 thick, with thin partition walls. .P. nereocystis 6. Fronds MOncecious. : 2... 12. cc eee scenes see ee soenmeen P. leucosticta Fronds dicecious, ...... 2... scce cece ceceee cece nseene cneee P. laciniata 196 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 7. TONGS MONOSCIOUS |: jsxs n= \worels! Aes, QOS \N0 CO SLY e ee ih 2 Seok oO oo Oe U Row 8883} Soe oy, : [Hus] PLATE XX. WUT Reeo ORE) Ly a VAY Ps iy gee L345 Les SoS Hl vie NT) Mi Hi He a; m Ky ite cH ) a Pao Wea ss: eg + 4 a 8.87 pl Bg ¢ ‘si ats it Ret eee eed) . ai endless: p oe Te SSR "Jeovobal Sh yy ih = Q¥ : is SS Ss C2 Og, rosea See 19 Ca Nepery 5 CRS Litre) NHN : RY ey QQ r Gaeconsras SAN OTQOALX2RO So ISOS 206 CO (os Ip YS =m pee es eed se Se Gp » & QE psitscai= CREO ORS FHOTO-LITH. BRITTON & REY, SF. ORPHYRA VARIEGATA. F1G819-22 PoRBHYRA NAIADTIM. ues 4 1/0 \p ay aaa * my ian 240 Fig. Fig. Fig. Fig. Fig. 22. 24. 25. , 26. rf 28. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXII. Diagrams illustrating Formation of Reproductive Bodies. Cube, representing cell of Porphyra sp., which has undergone a cruciate division perpendicular to the surface of the frond. For- mula for sporocarps, 4 (Rade ee G ) : ara e The same, after undergoing a division parallel to the surface of the frond in each segment. Formula for sporocarps, 8 ( Le <). Bi) aKa The same after undergoing another cruciate division perpendicular to the surface of the frond in all segments. Formula for sporo- carps, 32 (Pe 7 <); for antheridia, 8 (44, wee <) The same after undergoing another division parallel to the sur- face of the frond in all segments. Formula for sporocarps, 64 (=; be It for antheridia, 16 (2% Bo =). 4 4 4 2 2 4 The same after undergoing another cruciate division perpendicular to the surface of the frond in all segments. Formula for anther- idia, 64 {2% «2 Se 4 4 4 The same after undergoing another division parallel to the sur- face of the frond in all segments. Formula for antheridia, 128 (ae. Ao =) F 4 4 8 [Hus] PLATE XXII CEGGESE S| : QOS : SSSA : QUeasese ot cueeusser al Eee SS ee SS a ES Ie ES, Proc.CALACAD. SC1.a” Ser. Bor. VoLtil ON OF REPRODUCTIVE Booies oF PORPHYRA. ] FORMAT Tarp: SeRis Sones of Pacific DT IRA Se are ee ALICE. EastTw oop NE wrator of the Department ye Sitany es BY Y rH AcaDEMy i's Nien Sy : t =) i rain eC Mi; ) MITTEE | PUBLICATION COM ; hte Ua a 14 ; ~ Rat 7 ergs Maes ite, TR Foi ‘ eee GILBERT, Chairman ye {SCAT we PROCEEDINGS Fi OF THE CALIFORNIA ACADEMY OF SCIENCES z TuirRD SERIES q BoTANny Von. ff, No. 7 # Some New Species of Pacific Coast Ribes BY AuicE EASTWOOD ¥ Curator of the Department of Botany WitH Two PLATES Issued April 14, 1902 SAN FRANCISCO PuBLISHED BY THE ACADEMY 1902 4 ‘. . ARS hy Set Py eT all SOME NEW SPECIES OF PACIFIC COAST RIBES. BY ALICE EASTWOOD. Curator of the Department of Botany. PLATES XXIII AND XXIV. Tue genus Ades is represented on the Pacific Coast by four subgenera, which include an indefinite number of species and forms. A recent attempt on the part of the author to identify the unnamed specimens of this genus, which had for years been accumulating in the Herbarium of the California Academy of Sciences, has brought to light the following species which seem to be undescribed. The first six belong to the suborder Ribesia; of these, numbers one and two fall into the group typified by Azdes sanguineum Pursh, number three into that typified by 7. malvaceum Smith; numbers four, five and six into that typi- fied by A. nevadense Kellogg. The last three belong to the suborder Grossularia. Some botanists might include the first six under /zes sanguineum, the last three under /?. menzzesiz. This aggre- gation may satisfy the amateur to whom generic differences are sufficient, but the real student desiring to learn the truth regarding a genus will find it a source of great confusion, and altogether unsatisfactory. While it is to be kept in mind that nature knows no boundaries, and that orders, genera, and species are divi- sions made by man for his own convenience, yet these methods of classification have a scientific value beyond that of pure utilitarianism, and ought to show as far as possible the life-history of a group of related plants and of the entire plant world, when the knowledge of man makes it possible. [ 241 ] April 14, 1902. 242 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Yr, 1. Ribes brandegei, sp. nov. PLATE XXIII, Fics. 1a AND 10. Shrub with erect, branching stems unarmed; older bark dull brown; young bark glossy, not shreddy. Leaves three-lobed, reniform, 3-4 mm. long, and of about equal width; upper surface sparsely pubescent with fine, silky hairs, mostly on the veins, and with scattered sessile or shortly stipitate glands; lower surface pale green, with appressed, glandular hairs, veins at base vil- lous; margin incised-dentate, glandular-ciliate; petioles generally shorter than the blades, glabrous or clothed with a fine pubescence under the gland-tipped hairs; stipular dilation 3 mm. broad, fringed with uneven, glandular hairs. Inflorescence racemose, erect in flower, the peduncles equalling or longer than the flowering portion, generally surpassing the leaves; pubescence as on the petioles; bracts foliaceous, oblanceolate to obovate, acuminate, incised, glandular-ciliate; flowers three to ten, on slender, erect pedicels which later become as long as the flower. Calyx rose-color, 8 mm. long, pubescent on the outside, glandular at base, puberulent within; divisions as long as the tube, oblanceolate to obovate, cucullate at summit, 2 mm. wide. Petals white, half as long as the calyx divisions, 1.5 mm. wide, orbicular- spatulate, on short claws. Stamens with slender filaments, 1.5 mm. long; anthers oblong, tipped with a blunt mucro. Style two-cleft at apex, with the stigmas broad. Berry glabrous, globular. This species is related to Azbes sanguineum Pursh from which it differs in the pubescence, inflorescence, and shape of floral organs, as can be seen by the figures. , Collected by Mr. T. S. Brandegee, in whose honor the author takes pleasure in naming it, first at Sierra de Laguna, Lower California, January 25, 1890, later in the mountains of the Cape Region, March 26, 1892. 2. Ribes scuphami, sp. nov. PLATE XXIII, Fics. 2a AND 26. Shrub with the upper bark reddish, shreddy, puberulent, unarmed. Leaves orbicular, three- to five-lobed, truncate to reniform at base, 2-5 cm. wide, about as long, unevenly dentate; upper surface pubescent with crisp, spread- ing hairs; lower, canescent with matted hairs; stipular dilation of the petiole broad, glandular, and tomentose, fringed with glandular hairs; petioles about as long as the blades, with pubescence like the stipules. Racemes numerous at the ends of the branches, 9 cm. long, slender, when flowering erect on peduncles which are shorter than the leaves; bracts oblanceolate, red, gland- ular, 8 mm. long, denticulate at apex; pedicels filiform, erect, a little longer than the bracts. Flowers subtended by two small, red bracteoles which are soon deciduous. Calyx rose-color, with tube 5 mm. long, divisions Bot.—VOL. II.] EASTWOOD—PACIFIC COAST RIBES. 243 linear-oblong, 7 mm. long. Petals white turning reddish, oblanceolate, cune- ate, 4 mm. long. Stamens a little shorter than the petals; anthers globular. Ovary sparingly pubescent, and with scattered, stipitate glands. This is nearest to /Pzbes sanguineum Pursh. It differs especially in having the racemes erect in flower, also in the more slender flowers with narrowed divisions. This species is the most beautiful of all belonging to the group of which Lf. sanguineum is the type. It was collected on Smith River, Del Norte County, California, by Major J. R. Scupham, May, 1898. It is a pleasure to name this plant in honor of one who has brought many interesting plants to the herbarium of the California Academy of Sciences from little explored parts of California. ra 3. Ribes indecorum, sp. nov. PLATE XXIII, Fics. 3a AND 36. Shrub with erect stems, having dark brown, shreddy bark on the older growth, the younger parts tomentose and glandular. Leaves three-lobed, 2-4 cm. long, 2-3 cm. wide, finely rugose on the upper surface, clothed with stipitate glands, and a fine, sparse, silky pubescence; lower surface white with a felt-like tomentum, and with a few gland-tipped hairs on the veins; margins irregularly, doubly. crenate; petioles stout, shorter than or equalling the blades, glandular and tomentose, the stipular dilation (as wide on each side as the petiole) fringed on the margin with uneven, gland-tipped hairs. Inflorescence racemose, spreading or pendent, in fruit surpassing the leaves; flowers sessile but erect; peduncles short; bracts foliaceous, almost equalling the flowers, lanceolate, 6 mm. long, 2 mm. wide, with the margins fringed with long, gland-tipped hairs. Peduncles stout, glandular, and tomentose. Flowers at base subtended by two membranous, glandular, and tomentose bracteoles; calyx-tube more than twice as long as the broad, rounded divi- sions; these tomentose and glandular on both sides, almost 2 mm. wide; petals orbicular, reniform, 1 mm. wide, crenulate, on very short and broad claws. Stamens as long as the petals, on stout, short, deltoid filaments; anthers .75 mm. long, longer than the filaments. Style stout, hairy at base, two-cleft at apex, with broad, yellow stigmas; ovary tomentose and somewhat glandular. Collected by the author at Cajon Heights, near San Diego, California, March 14, 1891. There is also a specimen in the Herbarium of the California Academy of Sciences collected by Dr. George Thurber at San Pasqual, San Diego County. It is labeled Arzdes sanguineum, No. 606. 244 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Ribes indecorum is nearest to ARzbes malvaceum but differs most noticeably in the much smaller and sessile flowers. The floral organs, too, are not the same. V 4. Ribes ascendens, sp. nov. PLATE XXIII, Fics. 4a anp 46. Erect shrub unarmed, with gray-brown bark on older stems, younger stems paler and shreddy. Leaves three- to five-lobed, orbicular, reniform, 3-6 cm. wide, 2-5 cm. long, crenate-dentate, almost glabrous on the upper surface, the lower clothed with fine, spreading pubescence; petioles equalling or shorter than the blades, glandular; stipular dilation narrow, fringed with long, gland-tipped hairs. Peduncles generally surpassing the leaves, at first erect, later nodding, glandular-pubescent; flowers crowded at the summit of the peduncle, which is naked for more than half its length; bracts oblanceo- late, rounded at apex, 7 mm. long, 2 mm. wide, with gland-tipped hairs on the surface and margin; pedicels half as long as the bracts, lengthening with age, and recurving upwards, so that the berries are erect. Flowers subtended by two membranous bracteoles which are soon deciduous. Calyx open- campanulate, rose-color, the tube about half as long as the divisions; these ovate, obtuse, 3.5 mm. long, 2.5 mm. wide, slightly pubescent. Petals white, orbicular, narrowed to a short, broad claw, 2 mm. wide. Stamens not equal- ling the petals, filaments linear, anthers oblong. Ovary clothed with gland- tipped hairs. Berry veiny, sparingly glandular, becoming 7 mm. or more in diameter. This species is near 7’. nevadense Kellogg, but the racemes are ascending when in flower. The floral organs also differ in shape. The type was collected by the author at Millwood, (Sequoia Mills) Fresno County, California, in flower, May 4,1895; in fruit, July 18, 1893. There are specimens from General Grant Grove in the same vicinity, and from Coburn’s Mills in Fresno County, collected by T. S. Brandegee; the former, July, 1892, the latter, May 29 (year not given). 4a. Lftibes ascendens jaspere, var. nov. (Plate XXIV, fig. 5). This is similar to the species in general appear- ance, but the sepals and petals are more orbicular, the filaments are shorter, and the anthers broader and more orbicular. It is named in honor of Mrs. William Jasper, who sent the specimen from San Emidio Cajfion, Kern County, California, May, 1895. Bot.—VOL. II.] EASTWOOD—PACIFIC COAST RIBES. 245 v 5. Ribes hittellianum, sp. nov. PLATE XXIV, Fics. 6a AND 60. Erect shrub with spreading branches, 2-3 feet high; bark smooth, un- armed, gray-brown, shreddy on the younger branches. Leaves three- to five-lobed, orbicular, reniform or truncate at base, 3-4 cm. wide, 2-3 cm. long, irregularly dentate and somewhat revolute, rugulose veiny, glabrous on both sides but with some scattered glands on the lower; petioles about as long as the blades, sparingly tomentose and glandular; stipular dilation broad, truncate, membranous, as wide on each side as the petiole. Racemes 1-2 cm. long, at first erect, later nodding, but with the pedicels erect; flowers three to eight, crowded, subtended by deciduous bracteoles. Calyx tubular- campanulate, with the tube 1 mm. long, the divisions rose-color, oblong, obtuse, 4 mm. long, 2 mm. wide. Petals white, narrowly oblong, three- fourths as long as the sepals and one-half as wide. Stamens with subulate filaments, half as long as the sepals; anthers orbicular. Stigmas two, capitate; ovary and immature fruit clothed with stipitate glands. This species belongs in the group of which A. nevadense Kellogg is the type. It differs from that species in the inflorescence and the shape of the floral organs. Collected near the head-waters of Cafion Creek, Trinity County, California, not far from Twin Lakes, July 9, 1901, and named in honor of Mr. Carlos T. Hittell, one of the party on a trip to these little known mountains. of 6. Ribes glaucescens, sp. nov. PLATE XXIV, Fics. 7a AND 70. Unarmed shrub with older bark gray-brown, younger bark bright brown glossy, shreddy. Leaves three-lobed, orbicular-reniform, about 3 cm. long, 3-5 cm. wide, irregularly dentate, glabrous except for some minute glands on the lower surface, glaucescent, paler on the lower surface; petioles about as long as the blade, minutely puberulent, with the stipular dilation on each side narrower than the petiole, and sparingly fringed with glandular hairs. Inflor- escence in fruit spreading or erect, generally shorter than the leaves, rather loosely flowered with from five to ten flowers; peduncles as long as the raceme, striate, puberulent; pedicels slender, becoming 5 mm. long, shorter than the brown, membranous, gland-tipped bracts. Flowers subtended by two small, deciduous, reddish bracteoles. Calyx open-campanulate, with very short tube, and spreading divisions; these rose-color, oblanceolate, 4 mm. long, 1.5 mm. wide, glabrous. Petals white, spatulate two-thirds as long as the sepals, denticulate near the apex. Stamens half as long as the sepals, with suborbicular anthers and broad filaments. Ovary glabrous except for the scattered stipitate glands. 246 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. This species is related to Azbes nevadense Kellogg from which it differs in the glaucous color of its foliage, the racemes erect in fruit, and the shape of the floral organs. It was collected by the author on Mount Shasta, August 13, 1893. No note was taken of the exact locality, but it must have been some place on the trail from Sisson. v 7. Ribes oligacanthum, sp. nov. PLATE XXIV, Fics. 8a AND 80. Erect, branching shrub with the younger stems puberulent, older gray- brown, not shreddy, somewhat tortuous, unarmed, except for the simple or triple thorns at the leaf-axils; these often short and abortive. Leaves broadly ovate or orbicular, three- to five-lobed, 2-3 cm. wide and about as long, with margin laciniate-dentate, base truncate, but cuneate where it joins the petiole, glabrous; petioles slender, as long as or twice as long as the blades, glabrous or puberulent; stipular dilation twice as wide as the petiole, ciliate with white, silky hairs. Peduncles filiform, one- to three-flowered, 1-2 cm. long; pedicels almost as long, together becoming 4 cm. long, slightly puberulent; bracts broadly ovate to orbicular, acuminate, three-angled, clasping, reflexed. Calyx 12 mm. long, the linear, acute segments more than twice as long as the tube, 2 mm. wide, apparently white, veined with parallel veins that branch near the apex, puberulent or glabrous. Petals involute, obovate when spread out, irregularly denticulate at apex, 3 mm. long, veins palmate. Filaments almost twice as long as the petals, dilated at base; anthers oblong- ovate, cordate at base, tipped with a recurved mucro. Style divided two- thirds of its length, slightly exserted. Young fruit puberulent, clothed with a few long spines (about ten) each about 5 mm. long. Related to Arzbes californicum Hook. & Arn. from which it differs in the sparsely spinous fruit, the glabrous leaves, the lax inflorescence, and the shape of the floral organs. Collected by the author on the road between Jolon and King City, in Monterey County, California, near Mans- field’s Ranch, ten miles from King City, May, 1897. / / 8. Ribes sericeum, sp. nov. PLATE XXIV, Fics. 9a-9/. Erect, branching shrub, several feet high; stems clothed with numerous fine, weak, short prickles, which are gland-tipped on the young shoots, also with short, close, silky pubescence; axillary thorns three, orange-color, stout, united, the middle one longest, more than 1 cm. long, broadening at the Bot.—VOL. II.] EASTWOOD—PACIFIC COAST RIBES. 247 base, pubescent and glandular on the lower part, glabrous on the upper. Leaves thin, three- to five-lobed, broadly ovate-orbicular, reniform or trun- cate at base, 2-4 cm. long, not quite so wide, incised-crenate, clothed with fine, white, silky hairs which are appressed or spreading, also with fine gland-tipped hairs; petioles about as long as the blades, more glandular and more spreading-pilose, dilated only at the very base, and without the appear- ance of stipules. Peduncles one- to three-flowered, slender, erect, with pubescence like the petioles; pedicels about half as long; bracts orbicular or three-lobed, foliaceous; bractlets similar but smaller. Flowers 2 cm. long, open-campanulate in the bud. Calyx with the divisions at length reflexed, longer than the tube and the ovary, oblong, purplish red, greenish near the apex, softly silky villous on both sides; tube campanulate, veined, slightly glandular at base. Petals white, 5 mm. long, involute, erose along the almost truncate apex. Stamens with filiform, purple filaments, exserted beyond the sepals in the opening flower, and also beyond the pistil; anthers narrowly linear-oblong, almost 2 mm. long, obtuse. Pistil two-cleft for 2mm. Ovary densely clothed with horizontally spreading fine, silky hairs mixed with some longer, glandular hairs, the glands purple. Fruit purple, clothed with short, weak bristles and scattered hairs. Some of the bristles retain the purple glands on the fruit. Collected in flower by Mr. R. A. Plaskett, at Spruce Creek; also at Gorda, in flower and fruit. Collected by the author at Pacific Valley, with immature fruit. 2. ser7- ceum flowers in December and January and fruits in June. At Point Sur specimens were collected by the author in June, 1893, with very large, pear-shaped fruit, almost 4 cm. long, and specimens with globular fruit were collected at about the same time at Slate’s Hot Springs. All these localities are on the coast of Monterey County, California, at the base of the Santa Lucia Mountains, and the range extends from south of Point Gorda to north of Point Sur. fribes sericeum is related to F. subvestitum Hook & Arn. but it has different leaves, different pubescence, and the floral organs are not the same. 8a. Libes sericeum viridescens, var. nov. The variety is similar to the type, but the flowers are smaller and green- ish, the leaves are more densely clothed with silky white hairs, and are more orbicular-reniform. The peduncles in the specimens examined all have single flowers. This variety was collected by R. A. Plaskett at Gorda, Monterey County, California, January, 1898. 248 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. y g. Ribes hystrix, sp. nov. PLATE XXIV, Fics. toa-1od. Shrub several feet high, with light brown, tortuous branches, minutely pubescent and thickly beset with stout, rigid, horizontal, yellow prickles, some gland-tipped, generally small on the new growth, and increasing in size with age; axillary thorns triple, stout, distinct at base, middle one longest, becoming 15 mm. long, lower part pubescent, upper, glabrous. Leaves thin, three-lobed or some five-lobed, with the basal lobes small, 2-4 cm. wide, orbicular-reniform, incisely dentate, minutely pubescent and dotted with sessile glands on the lower surface, almost glabrous on the upper; petioles about as long as the blade, tomentose and slightly glandular. Peduncles one- to three-flowered, 1-2 cm. long, ascending, slender, sparingly pilose and clothed with gland-tipped hairs; pedicels less than half as long, occasion- ally longer; bracts orbicular or lobed, clasping, acuminate to obtuse. Calyx pubescent and glandular; tube a little longer than the ovary; divisions 1 cm. long, surpassing the rest of the flower when reflexed, 3 mm. wide, lower part purple, near the apex greenish, obtuse. Petals white, broadly obovate when spread out, acute, narrowed to a short claw, involute, 4 mm. long. Stamens with broad filaments dilated at base, as broad and long as the anthers; these about 3 mm. long, sagittate at base, tipped with a blunt mucro. Styles surpassing the sepals in the opening flower, divided about half; stigmas small, capitate; ovary globular, tomentose, and densely clothed with purplish bristles, some near the calyx gland-tipped. Fruit purple, more or less densely clothed with stiff, spreading prickles, 2-5 mm. long. This species is nearest to A. menzzesez Pursh but differs in the glandular pubescence, the shape and texture of the leaves, the size of the flower, and shape of the parts. Collected in flower by Mr. R. A. Plaskett, at Gorda, Santa Lucia Mountains, California, December, 1897. The fruiting specimens were collected by the author at Pacific Valley, in the same vicinity, May, 1897, and June, 1893. Bot.—VOL. II.] EASTWOOD—PACIFIC COAST RIBES. 249 Kry TO THE SPECIES OF PaciFic CoAsT RIBEs. The following key, made by the author for convenience in looking up the described species of Pacific Coast Azbes, is appended, as it may be useful to others under the same circumstances. Chrysobotrya SPACH. GOLDEN CURRANT. Calyx with long tube; flowers yellow, berry smooth, leaves convolute in bud; stems without thorns or prickles. 1. &. tenutflorum LINDL., Bot. Reg., Vol. XV, Tab. 1274. Oregon and California. 2. R, aureum Pursn, Fl. Am. Sept., Vol. I, p. 163. Oregon and Washington. Ribesia BERLANDIER. WILD CURRANT. Calyx-tube cylindrical to rotate; berries smooth; leaves plicate in bud; stems without thorns or prickles; flowers in racemes. Calyx-tube cylindrical. 3. &. bracteosum Doucu., Hook. Fl. Bor. Am., Vol. I, p. 233. Northern California to Alaska. 4. &. cereum Dovuct., Bot. Reg., Vol. XV, Tab. 1263. California to British Columbia. 5. &. viscosissimum PursH, FI. Am. Sept., Vol. I, p. 163. Sierra Nevada of California to British Columbia. 6. R. sanguineum PursH, Fl. Am. Sept., Vol. I, p. 164. Northern California to British Columbia. 7. R. scuphamt, sp. nov. Del Norte County, California. 8. RR. brandeget, sp. nov. Lower California. 9. &. glutinosum BENTH., Trans. Hort. Soc., Ser. II, Vol. I, p. 476. Coast Mountains of California. 10. R. malvaceum SmitH, Rees. Cycl., Vol. XXX. Coast Mountains of California. 11. &. palmert VasEY & Ross, Proc. U. S. Nat. Mus., Vol. XI, p. 529. Lower California. 12. &. indecorum, sp.nov. San Diego County, California. 13. R. nevadense KELLOGG, Proc. Cal. Acad. Sci., Vol. I, 1855, p. 65 = (R. sanguineum variegatum Wats., Bot. King’s Exped. tooth Par.) Sierra Nevada of Central California. 14. &. ascendens, sp. nov. Sierra Nevada of Central California. 15. &. hittellianum, sp. nov. Trinity County, California. 16. RR. glaucescens, sp. nov. Mount Shasta, California. 17. &. viburnifolium Gray, Proc. Am. Acad., Vol. XVII, p. 202. Lower California, and islands off the coast of Santa Barbara, California. 250 18. 19. 20. 21. 22. 29. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Calyx rotate or saucer-shaped. R. prostratum UV HERI, Strip. Nov. 3, Tab. 2. British Columbia and northward. R. hudsonianum Ricuarps., Frank. Journ. 2nd Ed., App., p. 6. British Columbia and northward. . migratorium SuKsporF, Deutsche Bot. Monats., Bd. XVIII, p. 86. Washington. .laxiflorum Pursu, Fl. Am. Sept., Vol. II, p. 731. Wash- ington and northward. . ciliosum Howe .., Fl. N. W. Am., p. 208. Mount Hood, Oregon. . erythrocarpum COvILLE & LEIBERG, Proc. Biol. Soc. Wash., Vol. X, p. 132. Crater Lake, Oregon. yy BR DW Grossularia A. RICHARD. GOOSEBERRY. Stems thorny under the fascicles, generally prickly besides; leaves plicate in bud; flower solitary, in corymbs, or racemes. 24. 25. 26. a, 28. 29. 30. aly ae 33: 34. 35- 36. 37- 38. Berry prickly or glandular. R. menziesii Pursu, Fl. Am. Sept., Vol. II, p. 732 (= 2. ferox SmitH, Rees. Cycl., Vol. XXX. [Index Kewensis]). Coast Mountains of California. . hystrix, sp. nov. Santa Lucia Mountains, California. . californicum H. & A., Bot. Beech. Voy., p. 346. Coast Mountains of California. . occidentale H. & A., Bot. Beech. Voy., p. 346. Coast Moun- tains of California. A doubtful species. . oligacanthum, sp. nov. Santa Lucia Mountains, California. . victoris GREENE, Pittonia, Vol. I, p. 224. Coast Mountains of California. . mariposanum CONGDON, Erythea, Vol. VII, p. 183. Mariposa County, California. . subvestitum H. & A., Bot. Beech. Voy., p. 346. Coast Moun- tains of California. . sericeum, sp. nov. Santa Lucia Mountains, California. . lobbii Gray, Am. Nat., Vol. X, p. 274. Northern California to Vancouver. . marshallii GREENE, Pittonia, Vol. I, p. 31. Northern Cali- fornia to Oregon. _amictum GREENE, Pittonia, Vol. I, p. 69. Sierra Nevada of California. . wilsonianum GREENE, Erythea, Vol. III, p. 70. Mountains of Kern County, California. . cruentum GREENE, Pittonia, Vol. IV, p. 35. Coast Mountains of California to Oregon. . aridum GREENE, Pittonia, Vol. IV, p. 35. Sierra Nevada in Kern County, California. Sie ee ae ee Sed SRR re ULES Bot.—VOoL. II.] EASTWOOD—PACIFIC COAST RIBES. 251 39. R. hesperium McC.iatcuir, Erythea, Vol. II, p. 79. Los Angeles County, California. 40. R. amarum McC Latcuiks, Erythea, Vol. II, p. 79. Los Angeles County, California. 41. &. montigenum McC.atcuik, Erythea, Vol. V, p. 38. 42. FR. lacustre Porr, Encycl. Suppl. II, p. 856 (= &. echinatum Dovuct., Bot. Reg., sub. Tab. 1349 [Index Kewensis] ). California, northward. 43. FR. lacustre var. molle Gray, Bot. Cal., Vol. I, p. 206. Sierra Nevada Mountains, northward. Berry smooth. 44. RR. divaricatum Douct., Trans. Hort. Soc., Vol. VII, p. 515 (= R. villosum Nutt., T. & G. Fl. N. Am., Vol. I, p. 547). California. : 45. R. oxyacanthoides L., Sp. Pl., p. 201 (= FR. saxosum Hook., Fl. Bor. Am., Vol. I, p. 231 [Index Kewensis]). Sierra Nevada of California to British Columbia. 46. FR. gracile Micux., Fl. Am. Bor., Vol. I, p. 111. Oregon to British Columbia. 47. FR. niveum LINDL., Bot. Reg., Vol. XX, Tab. 1692. Washington. 48. R. leptanthum Gray, Mem. Am. Acad., N. Ser., Vol. IV, p. 53. Sierra Nevada of California. 49. R. cognatum GREENE, Pittonia, Vol. III, p. 115. Oregon. 50. R. lasianthum GREENE, Pittonia, Vol. III, p. 22. Sierra Nevada of California. 51. &. velutinum GREENE, Bull. Cal. Acad. Sci., Vol. I, No. 3, p. 83. California to northern Oregon. 52. R. qguercetorum GREENE, Bull. Cal. Acad. Sci., Vol. I, No. 3, p. 83. Coast Mountains of California. This is the same as R. leptanthum brachyanthum GRay, according to Greene. 53. Rk. ambiguum Wats., Proc. Am. Acad., Vol. XVIII, p. 193. Washington. This has been changed to R. watsonianum KOEHNE. 54. R. montanum Howe.., Fl. N. W. Am., p. 210. Siskiyou Mountains, Oregon. This has been changed to #. dinomt- natum HELLER. Robsonia BERLANDIER. Stems thorny; parts of the flower commonly four; calyx with erect lobes. 55. . speciosum PursH, Fl. Am. Sept., Vol. II, p. 731 (= &. stamt- neum SMITH, Rees. Cycl., Vol. XXX = R. fuchsioides BERL., Mem. Soc. Geneva, Vol. III, Pt. 2, p. 58.) Monterey, Cali- fornia, southward. The references not being available, the following species could not be placed :— 56. R. roezli REGEL, Gartenfl., p. 226. Am. Bor. occ. [Index Kewensis]. 57. &. spethianum KOEHNE. 252 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXIII. Fig. 1. Ribes brandegeti, sp. nov. a. Diagrammatic representation of the flower; x 5. 6. Leaf, natural size. Fig. 2. Ribes scuphami, sp. nov. a, Diagrammatic representation of the flower; X 5. 6. Leaf, natural size. Fig. 3. Ribes indecorum, sp. nov. a. Diagrammatic representation of the flower; X 5. 6. Leaf, natural size. Fig. 4. Ribes ascendens, sp. nov. a. Diagrammatic representation of the flower; X 5. 6. Leaf, natural size. Fig. 5. Ribes ascendens jaspere, sp. et var. nov. [Eastwoou| PLATE XXIII. PRoc.CALACAD. Scr.32 SER Rot VoLIL ITTON & REY, 5. -LITH. BRI PHOTO nV. ep «TT oF, NL .RIBES SCUPHAMT, wn iat pd ae aa rm co poy pa JUV. i oP ET VAR} 254 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. EXPLANATION OF PLATE XXIV. Fig. 6. Ribes hittellianum, sp. nov. a. Diagrammatic representation of the flower; X 5. 6. Leaf, natural size. Fig. 7. Ribes glaucescens, sp. nov. a. Diagrammatic representation of the flower; x 5. 6. Leaf, natural size. Fig. 8. Ribes oligacanthum, sp. nov. a. Diagrammatic representation of the flower; x 5; dotted lines indicate the shape of the ovary and calyx-tube. 6. Leaf, natural size. Fig. 9. Ribes sericeum, sp. nov. a. Flower, natural size. 6. Buds, natural size. c. Leaf, natural size. d. Petal, spread out; X 2. é. Petal as it appears in flower; X 2. J. Stamen; X 2: Fig. 10. Ribes hystria, sp. nov. a. Flower and bud, natural size. 6. Leaf, natural size. G6. (Stamens) >< 2: ad.) Petal: 72: Proc .GALAGAn, Scr.3" SER Bor VoL. A.EASTWOOD, DEL. [Eastwoou] PLATE XXIV PHOTO -lITH. BRITTON & REY, 5.F. F16.6. RIBES HITTELLIANLM, SE NOV. FIG.4. RIBES OLIGACANTHUM, SE NOV. Fig.7. RIBES GLAUCESCENS, SE NOV. EI G 4. RIBES SERICEIM, SP NOV. Fig.10. RISES HYSTRIX, SP NOV. arn oe Sh af Mae OY \ f ARTY pote’ Ad es iy os . we _ CALIFORNIA ACADEMY OF SCIENCES - ‘Tarrp Series ey tie ) 2m 2)-§ pindle Formation in » 'e Wal bo es OS Tera our!) > “4 es hs > fe A bus bey \ : a SS j BNR Sig N “ie eons ayy _ Assistant Professor of Botany in the University of California a SUR & e" _. “Wrru. Four Pates. - gee SGN aRS 3a ore ba F : 4 MRE St Bg URL aS SAN FRANCISCO PUBLISHED BY THE ACADEMY para POLE lah ; . asf RG iis ‘ Ae AN tg Le ‘ re IK) iF y Aiea KS we + a PROCEEDINGS > J 3 OF THE S CALIFORNIA ACADEMY OF SCIENCES 4 THIRD SERIES “d BoTANY Vou, i. No: Ss es a. Cell Studies Be I. Spindle Formation in Agave ee BY W. JV cOSte2 rn our Assistant Professor of Botany in the Untversity of California WitH Four PLATES - s LIBRARY NEW YORK. Issued May 5, 1902 BOTANICAL GARDEN SAN FRANCISCO PUBLISHED BY THE ACADEMY 1902 CELL STUDIES.’ I. SPINDLE FORMATION IN AGAVE. BY W. J. V. OSTERHOUT, Assistant Professor of Botany in the University of California. CONTENTS. PLatEs XXV-XXVIII. I. INTRODUCTION AND TECHNIQUE. j} II]. SprnDLE FORMATION IN AGAVE. » First Mitosis: PAGE. 1. PROGENETIC STAGE.........-- sda Ss dbetiRos slp leieins it e.e.0 piney prea 261 DB GRMETIC STAGE) 5 alae olen cle'p ie 6 = gins ote hisiei 9,0 oinybie pin ec0r= {nme Heisler 262 PERSSTERLAR STAGE 5 ol ote ceva o'tleia riot sie mdi aie ern nieve > mY eg er Ae 265 HL) ASCHCULARG STAGE 2.0). 4-2 52.5jplei win cla ce eb wis #2 taba mains aiaielsoma ne a5 266 By) BAPOLAR STAGE ¢ 10 p55 vole Uh ois ret ae. e sink nis podem sepa sleet seas 266 Second Mitosis: F-) IMERODUCTION wc silt 62) = tid eesine es ans aienls plae vidal jinn erie eels bre vite 267 2.) (PROGENETIC | SENG, sns 5.045 60 \ce as pitta ot » aie ele in) weit rele es Bem 268 BULUGENMIIC STAGIs 0.225 daray eter eer tn te We acces Beaman sone 268 Be STE DLAR LAGE cola tu xo os ath p Say) sient = ot ents i ak alent ales a 269 EME AGCTCULAR SHAGE 203i cinjnpsiai + r NEW SPECIES FROM THE SIERRA NEVADA MOUNTAINS OF CALIFORNIA. BY ALICE EASTWOOD, Curator of the Department of Botany. Most of the plants described below were collected on a trip to the south fork of Kings River from Millwood to the Kings-Kern Divide (Harrison’s Pass), and to Kearsarge Pass. The trail lies near the boundary of Fresno and Tulare counties, and is one of the best known of the south- ern Sierra Nevada. It is a country of magnificent forests, beautiful mountain meadows, rocky slopes down which dashing torrents rush, and cafions through which the river flows serenely but swiftly. The upper elevations are characterized by jagged peaks and ridges which are clothed with everlasting snow and enormous granite boulders, and are gemmed with little lakes of great beauty. The collection was made between July 2nd and rath, 1899. To the care and assistance of Messrs. Pierson Dur- brow, S. L. Berry, and Benjamin Brooks, members of the party, the success of the trip is due. The types are in the Herbarium of the California Acad- emy of Sciences. Vv 1. Streptanthus gracilis, sp. nov. Annual or biennial from a slender tap-root, with delicate branches, chiefly from the base, 1-2 cm. high, glabrous, glaucous. Basal leaves orbicular to narrowly elliptical or spatulate, sinuate-dentate to obtusely lobed or lyrate, tapering to long slender petioles as long as or twice as long as the blades, together 1-3 cm. long; cauline leaves linear-oblong to ovate, entire to crenately lobed, 5 mm. to 2 cm. long, auriculate at base, either sessile or on very short petioles. Racemes few-flowered, those from the slender basal branches one- to six-flowered; pedicels erect, 1-5 cm. long, generally shorter than the calyx; bracts wanting except with the lowest flowers, which are in the axils of the [ 285 ] June 2, 1902. 286 _ CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. upper leaves. Calyx rose-purple, urceolate, narrowed under the spreading lobes; the lobes obtuse, membranously margined, and undulate, becoming as long as the tube formed by the united sepals; disk-like receptacle conspicu- ous. Petals rose-pink, the slender claws exserted, blades spatulate to obovate-orbicular, half as long as the claws (3 mm.), and more than twice as broad (2 mm.), sinuate. Stamens in three pairs, with filaments all distinct, one pair with filaments 1 mm. long, the next pair 2 mm., and the third 5 mm.; the anthers of the two shorter pairs 3 mm. long, those of the longest pair not quite 2 mm. Pods (immature) erect, linear, glabrous, tipped with a broad, sessile stigma; the largest 1.5 cm. long, 1 mm. wide. Seeds not known. This delicate little plant was collected by the author below timber line on the trail from East Lake to Harri- son’s Pass (the type locality), and by Miss Catherine E. Wilson, in 1898, on the trail to Bull-frog Lake. It seems to be rare. 2. Polygonum exile, sp. nov. Annual, with slender, wiry stems, erect, four-ribbed, about 3 dm. high, glabrous or minutely puberulent, especially near the top, with few erect, virgate branches from near the base. Leaves erect, linear-lanceolate, or terete from the inrolling of the margins, jointed to the ochrez, cuspidate, the upper surface minutely dotted; lowest leaves 2 cm. long, diminishing upwards, shorter than the internodes; ochrez with three hyaline, long-acumi- nate divisions. Flowers solitary or few at the nodes, erect, sessile, or on short pedicels; divisions of the perianth extending to below the middle; white, pinnately veined with green or rose-colored veins, elliptical, hooded at apex, 2.5 mm. long, investing the ripe seed. Stamens three, with ovate, long-acuminate filaments half as long as the perianth; anthers minute. Akene three-angled, rhombic in outline, acute at each end, 2.5 mm. long, brown, glossy, minutely papillate; styles very short, deflexed. This is similar in general appearance to P. dougtasi Greene, but differs in having but three stamens and in the erect flowers. Collected by the author in Kings River Cafion, July 4, 1899. 3. Eriogonum scapigerum, sp. nov. Caudex branched, the divisions clothed with the dead brown bases of former leaves. Leaves all radical, oblong to orbicular, 5-15 mm. long, densely white-matted-tomentose on the lower surface, becoming somewhat glabrate on the upper, with undulate margins, obtuse apex, and cuneate base, tapering abruptly to the long petiole; petiole 3-5 cm. in length, flat, with a central rib, and broadening at base for about half the length. Scapes many, very slender, glabrous, 5-17 cm. high, terminated by a solitary head BoT.— VOL. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 287 not more than 1 cm. in diameter. Bracts united, ternate, the lobes deltoid- acuminate, dark red, glabrous, except for the long, white-woolly hairs on the margin; bractlets at base of involucres similar but smaller, with the lobes more deeply divided on one side. Involucres turbinate, glabrous, indistinctly ribbed, woolly-ciliate along the entire to undulate margins; pedicels ex- serted, distinctly jointed to the perianth at apex; divisions orbicular to obovate, hooded at apex, glabrous on the outside, hairy within at the base ; the outer ones broader, 2 mm. long. Stamens slightly exserted, anthers two-lobed, suborbicular. Akenes three-sided ; styles spirally coiled. This is near &. nudum Douglas, of which it may be only an alpine variety. It looks quite different from the common form as found in the Coast Mountains. Collected by the author on Harrison’s Pass, above tim- ber line, at an elevation of almost 14,000 feet, July 9, 1899. vie 4. Garrya pallida, sp. nov. Branching shrub, several feet in height; older stems glabrous, dark brown; younger ones cinereous, with densely appressed, silky pubescence. Leaves oval, elliptical, ovate, or obovate, becoming stiff and thick with age, pale green and glaucous, strongly veined, cinereous, with an appressed pubes- cence of fine silky hairs on both surfaces but denser on the lower, and becoming sparser with age; tapering somewhat at each end, the apex ab- ruptly acuminate, with the point recurved, margin entire, thickened, rarely slightly undulate; blade 3-7 cm. long, 2-4 cm. wide; petiole stout, .5-1.5 cm. long. Pistillate spikes pendent, solitary or clustered, 4-6 cm. long; lowest bracts deeply cleft, long-acuminate, upper ones cleft above the middle, abruptly acuminate, silvery, silky canescent from the densely appressed hairs; ovaries ovate, on short, thick pedicels, with pubescence similar to that of the bracts; styles divaricate; berries becoming almost glabrous. The staminate flowers have not been seen. This species is nearest to G. fremont: Torrey, and is found in the southern Sierra Nevada. The type was col- lected in Kings River Cafion, July, 1899. Specimens from San Emidio Cafion, Tejon Pass, and Tehachapi, all in Kern County, also from the region of the Kaweah River, in Tulare County, seem to be the same species. 4 5. Convolvulus berryi, sp. nov. Stems perennial, trailing but scarcely twining, 5 dm. or more long, with few branches from near the root, densely white-woolly throughout, with fine, spreading hairs. Lower leaves on petioles longer than the blades; upper shorter, 1-4 cm. long, broadly deltoid, with spreading sinus or Sagittate with the sinus less open, apex mucronate, the two basal angles 288 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. lobed with mucronate or obtuse lobes, 2-6 cm. wide at base, 2-5 cm. long. Flowers chiefly near the base, on long peduncles surpassing the leaves, often upwardly curving, 5-8 cm. long; bracts similar to the upper leaves, close under the flower, 1 cm. wide, 1.5 cm. long. Sepals ovate-oblong, unequal in breadth but of the same length, hairy with appressed hairs except on the membranous margins of the inner ones, mucronate, 14 mm. long, the broad- est Io mm. wide. Corolla yellowish white, hairy on the angles and at their tips, 4cm. long. Stamens with narrow sagittate anthers, 7 mm. long ; fila- ments hairy below, attached to the corolla for half their length. Style as long as the stamens (2.5 cm.), with ovate-lanceolate stigmas, 3 mm. long, I mm. wide. This beautiful species comes nearest to C. vz/losus Gray, from which it differs in the bracts, peduncles, and stamens, also in the broader leaves and longer peduncles. The pubescence is less velvety. It is also near C. tomentellus Greene, from which it differs in almost the same organs. Collected at Millwood by the author, July, 1893 and 1899, also near Converse Basin. The plant is named in honor of Mr. S. L. Berry. Vv 6. Castilleia brooksii, sp. nov. Perennial, branching from the base, but with a few short branches above, 3 dm. high, glandular, viscid throughout and with a pubescence of uneven, silvery, weak, jointed hairs. Leaves sessile, linear-oblong, entire to three- lobed, about 2 cm. long, less than 5 mm. wide; lobed leaves generally subtending the branches, the divided portion half the length. Branchlets terminated by short, compact spikes, with subsessile flowers; bracts variously and unevenly lobed, with the tips colored. Calyx a little longer than the corolla tube, obliquely gibbous at base, equally cleft before and behind, with divisions shorter than the tube, each two-cleft, with unequal, triangular-sub- ulate, one-nerved, obtuse divisions, 3 mm. long. Corolla 2.5 cm. long; galea longer than the tube, straight at first but later curving outwards, having three blunt teeth at apex, the middle one smallest; lower lip truncate, 3 mm. long, 2 mm. wide, the sharply acute teeth incurved, folds noticeable. Stamens exserted, with filaments smooth and anthers narrowly linear, with unequal cells. Ovary glabrous, obliquely acuminate; stigma clavate, exserted from the top of the galea. This comes under the group to which belong C. parviflora Bongard, and C. mznzata Douglas, with neither of which it agrees. The flowers are yellowish red but probably vari- able in color, as in most species of this genus. Collected by the author on the trail up Bubbs Creek, early in July, 1899, and named in honor of Mr. Benjamin Brooks. Bot.—Vot. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 289 V 7. Castilleia disticha, sp. nov. Perennial, erect, 6 dm. high, branching from the base and also above with generally short, slender, spreading branches; somewhat viscid and with a close, often somewhat scanty, cinereous pubescence mixed with longer, jointed hairs. Leaves linear, acute or obtuse, the lower 4 cm. long, 3 mm. wide, diminishing upwards, sessile by a truncate or subauriculate base, distinctly three-veined, the middle vein most conspicuous; margin entire or undulate-crisped. Inflorescence spicate, elongating in fruit, especially on the main stem, 1-2 dm. long; flowers after anthesis distichous, becoming more or less remote, sessile or almost so, with the capsule appressed to the stem, the calyx and corolla persisting and spreading; bracts foliaceous, the upper ones, only, colored, variously toothed, with the middle tooth longest, equalling or shorter than the corolla. Calyx slightly surpassing the corolla tube, about equally cleft before and behind for half the length; each division tipped with red, two-cleft, with triangular, subulate, unequal lobes, 2-3 mm. long, three-nerved, thin in texture, somewhat gibbous, but not broadest at base. Corolla red, 3 cm. long, with galea as long as the tube, truncate or emarginate at apex; lower lip three-toothed, the middle tooth much smaller than the lateral, separated by a broad sinus, thin, not callous. Stamens exserted, filaments glabrous; anthers narrow, with unequal cells, more than 2mm. long. Stigma exserted from the summit of the galea, capitate. Cap- sule obliquely oblong-ovate, 8-rio mm. long, chartaceous; seeds elliptical to orbicular, light brown, invested with a membranous, foveolate outer coat. This is more closely allied to C. minor Gray and C. sten- antha Gray than to any of the perennial species. Its ses- sile or almost sessile flowers, more brightly colored and differently shaped, together with the different habit of growth, mark it as distinct. The type was collected by the author at Converse Basin, on the trail to the south fork of Kings River, July, 1899. The species is also found at Millwood, where it was col- lected the same year by the author, and by Mr. T.S. Bran- degee, July 19, 1892. / f v - 8. Castilleia nana, sp. nov. Low, 3-6 cm. high, with several stems from a woody caudex, which is thickly clothed with the dead stems of former seasons ; somewhat cinereous and viscid, the pubescence of the inflorescence of longer, jointed, arachnoid hairs. Leaves simple and linear or three- to five-divided, with narrow, linear, acuminate lobes which are shorter or longer than the undivided portion, together 1-2 cm. long. Flowers in heads terminating the stems; bracts broad, similar to the upper leaves, embracing the sessile flowers and more than twice as long; the division terete from the involute margins, once and a half to twice as long as the lower, undivided part, which is 4 mm. wide 290 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. and 5 mm. long. Calyx glandular, subequally cleft before and behind, with the divisions twice as long as the tube, surpassing the corolla, each with two narrow, linear lobes 5-6 mm. long, extending below the throat of the in- conspicuous corolla. Corolla 7 mm. long, the lower lip longer than the galea, the three lobes thin, ovate-orbicular, obtuse, 2 mm. wide, somewhat saccate below; galea with the middle portion of thicker texture than the membranous sides, three-toothed at apex. Stamens with the upper pair of anthers exserted, the lower included, each distinctly two-celled, clothed with a few long hairs at base. Pistil capitate, .5 mm. in diameter, exserted; ovary lanceolate. Fruit and seeds unknown as the plants were too young. This is an alpine species found only above timber line. It is related to the group formerly included under Ortho- carpus, section Castilleioides, but differs from all other species in the peculiar corolla, it being the only species described with the galea shorter than the lower lip. Collected by the author on Harrison’s Pass, above East Lake, July 9, 1899. There are either one or two other low, alpine species in the same region which seem to be related to C. pallida Kunth, but the material is too scanty for satisfactory deter- mination. \ g. Mimulus bioletti, sp. nov. Annual, branching diffusely, chiefly from the base, about 2 dm. high, glandular-villous throughout, except the corolla. Leaves near the root spatulate; cauline leaves rhombic obovate to lanceolate, thin, tapering to a broad petiole, or sessile, sparingly serrulate or entire, 2-4 cm. long, 2-15 mm. broad. Flowers axillary, on slender, upward-spreading peduncles, almost as long as the internodes, generally shorter than the subtending leaves. Calyx tubular, 8 mm. long when in flower, 12 mm. in fruit, often purplish-dotted below the middle, plicately carinate-angled, the ribs rugose, rounded; divi- sions deltoid, with the margins involute, the obtusely pointed apex spreading outwardly. Corolla as long again as the calyx, with ampliate throat and scarcely two-lipped border, crimson, the upper lip with a yellow blotch dotted with crimson in the throat; limb 12-15 mm. across, with divisions rounded, crenulate, or entire. Stamens and style included; the former four in two sets, each united by the anthers, one set longer than the others; anthers ciliate, explanate, one above the other; stigma bilammelar, cuneate in out- line. Capsule included in the rigid calyx-tube, obtusely four-ribbed, opening at the sides from the base up; placenta free, except at the top; seeds numer- ous, minute. This belongs to section Eumimulus Gray, and is most closely related to MZ. palmeri Gray. It differs chiefly in the larger flowers and different calyx. Bor.—Vot. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 291 Collected in Hetch-Hetchy Valley, Tuolumne County, by Mr. F. T. Bioletti, in July, 1900. ¥ 10. Phacelia stimulans, sp. nov. Stems tall, simple from a branched caudex, becoming 5-6 dm. high, erect, sparsely leaved, generally flowering from the middle, viscid-pubescent, and clothed besides with fine, long, stinging hairs. Radical leaves forming a rosulate cluster, simple or with a few lobes at base, ribbed between the hispid veins, elliptical, acuminate, 3 cm. long; petioles very hispid with spreading hairs. Spikes of the panicle simple, the lowest and uppermost geminate, horizontally spreading, somewhat distant, 5-6 cm. long; pedun- cles very glandular, becoming shorter near the top; pedicels capillary, half as long as the calyx. Divisions of the calyx oblong-spatulate, hispid, net- veined, shorter than the corolla, surpassing the capsule. Corolla tubular, the lobes conniving after anthesis and persistent, held to the calyx by the tangling together of the long, persistent stamens and style. Filaments exserted, conspicuously clothed with long white wool. Capsule ovate- acuminate, hispid; seeds ovate, brown, not glossy, pitted. This is allied to P. cércinata Jacq. f., but is entirely unlike any of the described species which were formerly included under that species. On account of the stinging hairs of the stems and leaves it might be confused with P. nemoralis Greene; but this has an altogether different habit, pubescence, and inflorescence. Collected by the author, July, 1899, in Kings River Cajion, not far from the swampy meadow near which campers stop on the way to Bubbs Creek. / Vv 11. Gilia sparsiflora, sp. nov. Annual, a foot or so high, branching above, with slender, spreading stems, minutely glandular-pubescent. Leaves few, terete from the infolding of the margins, about an inch long, tipped with a short bristle. Flowers few, termi- nating the branchlets, two to three in the clusters, sometimes solitary in the upper axils; bracts keeled at base, three-lobed, the middle lobe much larger than the lateral, all subulate-aristate, surpassing the flowers. Calyx mem- branous between the ribs, clothed with dense, white, cottony wool, the un- equal, aristate-subulate divisions as long as the corolla tube. Corolla salver- form, tcm. long, white with some purple dots in the funnel-form throat; the divisions elliptical-obtuse, half as long as the tube. Stamens equally in- serted, with arrow-shaped anthers, obtuse at apex, exserted from the throat of the corolla. Capsule oblong, 1 cm. long; seeds few, oblique at base, three- sided, generally with rounded angles, developing mucilage and spiracles. 292 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Gilia sparsifiora belongs to the same group as G. ver- gata Steud. Collected by the author in Kings River Cajion, in July, 1899; and also along Bubbs Creek trail. v 12. Cryptanthe vitrea, sp. nov. Annual, with several stems from a tap-root, 1-2 dm. high, very hispid throughout with white bristly hairs which are pustulate at base. Leaves linear, 1-2 cm. long, 2 mm. wide, strongly nerved. Flowers sessile, in numerous short spikes from almost the lowest axils, on slender peduncles. Calyx 3 mm. long, the sepals conniving to form a tube around the nut- lets, the tips free, densely clothed with long stiff bristles, 3 mm. long, and also a fine, white, hispid pubescence. Corolla only about 2 mm. long, with a small limb. - Nutlets, only two maturing, ovate, obtuse, almost 2 mm. long and 1 mm. wide at base, very glossy brown mottled, sharply tuberculate, attached to the gynobase for the entire length, the groove closed except at the forks. The species comes nearest to C’. muriculata Greene, but the nutlets are broader, the flowers much smaller, and the entire plant so floriferous that in the dried specimen it is almost impossible to distinguish the peduncles. Collected by the author on Bubbs Creek trail, July 5, 1899. 13. Aster durbrowi, sp. nov. Stems erect, perennial from creeping root-stocks, disposed to grow in tufts, 1-3 dm. high, sparingly leafy, erect, branching only at the inflores- cence, glabrous and green except for some white woolly hairs which are sparse on the lower part, but which make the upper part almost cinereous. Radical leaves on long winged petioles which are dilated and clasping base, lanceolate, the blades about as long as the petiole, together 5-8 cm. long, 5-10 cm. wide, entire or distantly serrate, ciliate on the margin, the hairs becoming longer towards the base of the petiole and decurrent on the stem; cauline leaves similar but sessile by an auriculate or cordate clasp- ing base, the upper ones broadening and becoming shorter and more pointed, the lower ones narrowing towards the insertion. Heads cymose, the branchlets terminated by one to four middle-sized heads, 1-2 cm. across, on short bracteate pedicels; involucre of six rows of imbricated bracts with green, foliaceous tips, not spreading, the inner ones with purplish acuminate tips, the outer linear, mucronate, glabrous except for the ciliate margins, distinctly one-nerved and chartaceous at base; rays pistillate, reddish purple, I cm. long, 1.5 mm. wide, dentate at apex, sparingly ciliate on the lower part; disk flowers purplish; corolla shorter than the pappus, which extends Bot.—Vot. II.] EASTWOOD—NEW PLANTS FROM CALIFORNIA. 293 about te the exserted style and stamens; the tube slightly pubescent. Akenes (immature) hispid with dense, white, upwardly appressed hairs; pappus simple, scabrous. This is near A. yosemztanus Greene (A. ascendens yosem- ztanus Gray), of which it may prove to be a variety. The habit of growth is quite unlike that of the above species, while its larger heads, sparingly leafy stems terminated by few heads, and the auriculate- or cordate-clasping cauline leaves make it appear even more distinct. It is the com- mon aster of the wet meadows at the upper altitudes in this region. Collected in Horse Corral Meadow, July 11, 1899, and named in honor of Mr. Pierson Durbrow. v 14. Madia villosa, sp. nov. Stems simple, erect from an annual root, about 3 dm. high, slender, vil- lous with long, white, soft, spreading hairs, also becoming glandular near the top with black, stipitate glands. Lower leaves generally opposite, upper alternate, linear or the lowest oblanceolate, sessile, entire, or glandular- serrate with distant teeth, revolute, strongly ribbed, villous, with the hairs finely pustulate on the older leaves, obtuse at apex, 4-7 cm. long, 2-5 mm. wide. Heads few, terminating slender peduncles near the top of the stem, the uppermost first in bloom, clothed with few small bractlets; outer bracts of the involucre 8 mm. long, the foliaceous tips equalling that which encloses the akene, linear acuminate, villous and glandular, half as long as the deeply three-lobed rays, these often with a reddish-brown spot at base; inner bracts scarious with short foliaceous tips; disk flowers all sterile, the corollas as long as the abortive akenes, together 8 mm. long, the tube somewhat villous and the lobes clothed at tip with spreading hairs; fertile akenes flattened laterally, semilunate, black and brown mottled, minutely papillate in rows, 1.5 mm. wide, 4 mm. long, glabrous; anthers purple and exserted, giving a purplish color to the disk. This is nearest to Wada corymbosa DC. (Madaria cor- ymbosa Greene). It is also close to M7. hispida Greene, and really seems to be intermediate between the two species. The type was collected at Converse Basin, July 12, 1899. It was also found on Bubbs Creek trail. { af ivy hy wii yi iss Turd SERIE Bae 4 S be bercles of | % -denticul ata ‘Willd. ) ms ¥ fe ~ ot of Som e Other Leguminous Plants. se be James PEIRCE 7 ait > 4 4 hb xen: 1 ( H i ys © Associate Professor of Plant Physiology in ‘the Leland Stanford Junior University =e dA PER on a PLT 2k ye nt AUT Se ; ‘ Epa ee a ee 5 at vores Re rae Dae re ¥ «Issued June 21, 1902 SPSS bad dey ea ea Emr a ea ea D BY THE ACADEMY BSON “W. Ho PH Ae © E Jos a 1S) DITORS Se ae AL ie PF v Douce! PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES TurirRD SERIES BOTANY Vow. Hy No. to The Root-tubercles of Bur Clover (Medicago denticulata Willd. ) and of Some Other Leguminous Plants. BY GEORGE JAMES PEIRCE Associate Professor of Plant Physiology in the Leland Stanford Junior University / WitH ONE PLATE Issued June 21, 1902 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1902 nea Tepe | rey ih i ia, vA THE ROOT-TUBERCLES OF BUR CLOVER (MED- ICAGO DENTICULATA WILLD.) AND OF SOME OTHER LEGUMINOUS PLANTS. BY GEORGE JAMES PEIRCE, Associate Professor of Plant Physiology in the Leland Stanford Junior Untversity. CONTENTS. PLATE XXIX,. 5 PAGE D9 INTRODUCTION: AND. METHOD 2 23.) .dc)-4.0 one oy eke eon seen 295 II. ORIGIN AND MORPHOLOGY OF ROOT-TUBERCLES..........00 000. 298 III. THE Form AND DISTRIBUTION OF ROOT-TUBERCLES............ 312 IV.) [ne SrRUCTURE? OF ROOT-TUBERCEES © wiciccle aisles cleta te “ea iste ne 316 SSUES are! ais fe s(x wie oh) aiaiaiein wie or cde cea sR erally a mieteterevaide ieee eh eretee 324 PVEHIOGRAPEY ¥\5.2)5 6. dass soon aie sli bh at ates eral Aee: atts Kaien ale Getera tele 327 EXPLANADION: OF PLATE. /o/.102'cisaele crclevemeclonimerctecetnes ieaiaiciens 328 I. InrTRODUCTION AND METHOD. SOME time ago, on casually examining some hand-sections of the root-tubercles of Bur Clover (Medicago denticulata Willd.), I was struck by the great differences between the cells containing bacteria or bacteroids and those in which there were none. In these sections, the bacteria-containing cells looked so unhealthy, as compared with the cells free from bacteria, as plainly to suggest that the intimate rela- tions of bacteria and leguminous cells were not mutually advantageous, but that the bacteria were parasitic. There- upon I began a careful microscopic study of the root- tubercles of Bur Clover and other leguminous plants’ in 1 Bur Clover was especially favorable for my work because, at most seasons, I could get living plants from out of doors very near the laboratory, and I could grow such ma- terial as I needed at other times from seed very quickly in the laboratory. Besides this, however, I have studied Lupinus micranthus Doug]. var. bicolor Watson, L. rtvularis Doug]. var. latifolius, Melilotus parvifiora Desf., Medicago sativa \,inn., Hosackia subpinnata Torr.and Gray. The points which I wished to determine are essentially the same in all of these, and hence my descriptions, though specifically of Bur Clover, are applicable to the others. [ 295 ] June 17, 1902. 296 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. order, if possible, to ascertain the real relations of the tubercle organism to the cells in which it is found. This I believe I have done. The bacteria are parasites, not bene- fiting but injuring, if not finally killing, the cells in which they occur. Whether the association of these bacteria with a leguminous plant benefits the plant as a whole is another question, answerable not from microscopic examination but solely by experimental cultures. According to Frank (1890, p. 109), the Legumznose@ are not all similarly affected by the bacteria. Some are greatly benefited, stimulated to in- creased growth and other activities, while in others the bacteria are ordinary parasites, not benefiting the host in any way in return for the food derived from it. Without implying whether the results of infection benefit or injure the plant as a whole, one may speak of the roots or cells of leguminous plants being zzfected by tubercle bacteria. I shall later, however, take occasion to discuss whether the presence of the tubercle organism is really beneficial. The material studied was either fresh, growing out of doors wild or sown in boxes in the laboratory, or alcoholic. The latter was fixed in Flemming’s chrom-osmic-acetic mixture, dilute, and after washing for twelve to twenty-four hours in running water was dehydrated and kept in go per cent. alcohol. These tubercles, which were of different sizes, ages and conditions, according to the season, were imbed- ded in paraffin melting at 54° C., sectioned and mounted in the usual way. The youngest tubercles scarcely turn brown in the fixing fluid, but older ones become brown or almost black. In any case I transferred the slides, after the paraf- fin had been removed by turpentine, to a solution contain- ing one part commercial peroxide of hydrogen in twenty parts 80 per cent. alcohol. In this solution they remained a half hour, or until the sections were no longer in the least brown, and were then run down into water for staining. The method of staining which I found most useful is a combination of Flemming’s well known and now very popu- lar triple stain—anilin safranin, anilin gentian violet, and orange G.—with Ehrlick’s method of staining cover-glass BotT.—VOL. II.] PEIRCE—ROOT-TUBERCLES. 297 preparations of bacteria. The stains were made up according to the directions given in Humphrey’s Zimmer- mann’s Botanical Microtechnique (p. 186), and were used as Hof (1898) directs, except that after the sections had been for not more than two minutes in the gentian violet solution they were rinsed with water and placed for a half hour or longer in Gramm’s iodine solution to differ- entiate the bacilli and the infection threads from the cyto- plasm. Hof says that the sections may be left from two to three minutes in the anilin gentian violet. I often found this quite too long, and had difficulty in washing out enough of the violet without taking out the safranin also. One minute is usually long enough for these tissues. Washing off the Gramm’s iodine with water, the slides were then allowed to remain for one to two minutes in staining bottles containing orange G; they were then washed with absolute alcohol so long as gentian violet came off abundantly or needed to be removed (as shown by microscopic examina- tion), were cleared in clove oil, and mounted in xylol balsam. I decidedly prefer clove oil to xylol for clearing, as it aids in the differentiation for which this staining method is so highly prized. So far as my experience goes, this method of fixing and staining is perfectly certain to demonstrate the infection threads and to differentiate the bacteria in the cytoplasm and in the unstained matrix of the threads. I am, therefore, somewhat at a loss to understand the difficulties reported ‘by some authors in staining tubercles and their contents. Miss Maria Dawson (1899, p. 8) reports, for example: ‘‘For some time I made use of both hand-sections and microtome sections of paraffin material. The latter method I afterwards abandoned, however, since I found the tuber- cle tissues very difficult objects to stain upon the slide, and also ordinarily thin hand-sections serve better for the exam- ination of the filaments within the cells—a point to which I wished to devote special attention.’ With the stains Miss Dawson used on hand-sections, and of which she speaks 298 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. favorably, I, too, failed to get satisfactory results from microtome material, butowing to the success of my own method I did not try hers very long. II. Oricin AND MorPHoLtoGy oF ROOT-TUBERCLES. Although most that I can say about the entrance of the tubercle bacteria into the roots of leguminous plants has already been reported by others, I wish to describe the infection in Bur Clover (Medicago denticulata Willd.) from the beginning, and to discuss some of the stages in the process. I must also frankly admit that I do not know all that has been written on the subject, for the literature is copious and scattered, and I have been able to see only the papers herein referred to. I therefore bespeak lenient criticism of my acquaintance with the literature, remote- ness from the centres of scientific and other work making it very difficult to secure the papers, and even references to the papers, of my subject. How the tubercle bacteria in the soil come into contact with the root-hairs of the leguminous plants which they attack is not known. The majority of authors consider these bacteria (Bacillus radicicola Beyerinck, /hizobium leguminosarum Frank) only slowly motile if motile at all. In artificial cultures they are usually quite motion- less (Migula, 1900, p. 772). Zinsser (1897, p. 447) says they are small and actively motile. Miss Dawson (1900, p. 59) reports that in drop-cultures, a week or more old, the chains become motile, the shorter moving more rapidly than the longer, but none actively, and the motion resembles the pendulum movement of Oscz//atorza filaments. In younger drop-cultures, containing 2.5 per cent. gelatine, I have seen this movement. The movement of the chains formed in artificial cultures may be only a feeble index of a much more active movement of the separate bacilli when these occur in natural conditions. Artificial cultures are unsatis- factory at best, and it may very well be that the tubercle bacilli are actively motile, for a time at least, in damp soil. BoT.—VOL. II.] PEIRCE—ROOT-TUBERCLES. 299 It is well known that many bacteria are actively motile only for a short time after division. When the conditions in the soil—whatever these conditions may be—favor the repro- duction of these bacilli, the young and separate rodlets may be able to move with a fair degree of rapidity. They may possibly be stimulated into motility by the proximity of the root of some leguminous plant, and may be characteristically attracted to it by the substances diffusing from it into the soil-water (see Czapek, 1896). At first glance this seems improbable because of the very small proportion of root- hairs attacked to the total number of root-hairs formed; but on this point the following observations throw some light. In the field of a Leitz objective III and ocular 3 I counted one hundred root-hairs on the sides of a young lateral root. Of these hairs one was infected. There must have been an equal number of root-hairs on the top and bottom of the root as it lay on the slide. The zone of hairs was about five times as long as the distance through which I counted hairs. This would make the total number of hairs on this small root at least one thousand. I searched carefully, but found no other infected hair anywhere on this rootlet. The proportion of infection in this case is as one to a thousand. The rootlet examined was from a young Bur Clover seed- ling growing in sandy soil in the laboratory. There must have been a large number of Bur Clover bacteria in the soil used, for it was taken from a spot where Bur Clover throve last year and again this. Out of doors I fancy that the number of root-hairs would be greater in the same sandy soil than in the laboratory, for I watered my indoor mate- rial, and that outside was watered only by the rains. The number of root-hairs attacked is probably no greater, how- ever. The proportion of one to a thousand is therefore conservative. If, then, these bacteria are motile only slowly, if at all, it is apparently the mere chance of a root- hair’s growing to or very near where the bacteria are which makes infection possible. We may infer either that the 1 This I estimated with considerable precision by means of a mechanical stage. 300 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. bacteria are not motile or move only slowly, or that it is chance which governs the infection of the root-hairs. This latter inference is encouraged by a statement of Miss Daw- son’s (1899, p. 21), that she saw ‘‘ on a very small piece of a lateral root from one of the plants no less than twenty- seven hairs, side by side, with well grown infection tubes within them. This observation may serve to show how successful the attacks of this organism may be, provided suitable conditions can be arrived at.’’ To ascertain whether the number of infections in root- hairs in nature is smaller than it might be under other conditions, I tried the following experiment. Three layers of filter paper, moistened with tap-water, were laid in each of four small saucers and covered by tumblers. These were sterilized in an Arnold steam sterilizer on three suc- cessive days. A half dozen Bur Clover seeds were then placed under each tumbler on the damp filter paper. The seeds had just been removed from the little bur-like pods, dipped in corrosive sublimate solution of one to one thou- sand, and rinsed in distilled water which had been repeat- edly boiled. The filter-papers were watered daily with boiled distilled water. In a few days the seeds germinated. When the roots had grown to an inch or so in length and had developed many root-hairs, they were watered with boiled distilled water in which had been ground healthy growing tubercles. The water therefore held tubercle bacteria in suspension. The next day nearly every hair in the field on one side of a root was found to be enlarged and twisted at the ends and showed the beginning of an infection thread. Given the contact or close proximity of the tubercle bacteria with the root-hairs, infections may take place in great numbers simultaneously, at least when the roots are very young. Whether the roots are always susceptible, or whether older root-hairs or root-hairs on older roots are susceptible, is another question. Any change in the composition of the walls of the root-hairs may affect their solubility or at least their permeability by the bacteria (cutinization?). Bot.—VOL. II.] PEIRCE—ROOT-TUBERCLES. 301 It would appear from another experiment that not all con- tacts and infections of root-hairs with tubercle bacteria lead to the formation of tubercles. Among the seedlings grow- ing in sterilized moist chambers, I infected some with bacteria from a gelatine culture of Bur Clover tubercle bacteria. The next day showed a great increase in the number of bacteria, but the tips of the root-hairs, though bent in many instances, were not coiled in the manner usual in infections, but instead, were cut off into short sausage- shaped, often non-nucleated segments. In this way the bacteria which have entered a root-hair are excluded from the more vital parts of the root, just as gonidial cells in lichens are known to exclude the haustoria of the fungus by so dividing that only one daughter-cell contains any part of the haustorium which has penetrated the mother-cell (Hedlund, 1892; Peirce, 1899). Once given the contact with the bacteria, the root-hairs can become infected; but these infections may be resisted by the leguminous plant by cutting off the infected parts. I am by no means ready to attach especial weight to the result of this experiment for the following reasons: First, I did not repeat the experiment, important though it would be to prove that the root-hairs do cut off the infected por- tions; second, this result followed the infection of the sterile root of a seedling, not by bacteria suspended in water but by stroking the root with a platinum needle which had been dipped into a culture of the bacteria. By this means not only bacteria but also their accumulated pro- ducts in the culture-medium were put upon the root. It might well be that these products, rather than the bacteria themselves, so irritated the root-hairs that they segmented as above described. It would be interesting to follow this matter to a decisive conclusion, but it was not possible at the time to do so, and this point was not directly connected with the main object of this investigation. It may be that by similar means the root-hairs, and thereby the roots, of other than leguminous plants resist and escape infection by the bacteria which so characteristically affect the 302 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. leguminous plants growing in the same soil and under the same conditions. This possibility also deserves the test of experiment. Returning now to the question of the motility of the tu- bercle bacteria, we see that the experiments just described indicate that fewer infections take place in nature than when many bacteria are brought directly into contact with young and sterile roots, but the experiments leave the matter of the motility of the bacteria still undecided. The behavior of the bacteria in artificial culture is inconclusive, and ap- parently we cannot now imitate the conditions which prevail in the soil. Figures 1-3 show root-hairs of Bur Clover plants infected by tubercle bacteria. Figure 2 shows the lower and longer of the two hairs in fig. 1 more highly magnified. Figure 3 is a hair on another plant drawn with the same magnification as fig. 2 (300). In these figures, as in those of Frank, Dawson, and many others, it is noticeable that on the con- cave side of the curved tip of the root-hair there is a small mass of bacteria, this mass being continuous with the line or strand of bacteria extending through the hair. The wall of the hair seems intact and uninjured except where the small mass of bacteria is. At this point there is no ap- parent rupture of the wall. The wall may be actually per- forated, though to see this with the mass and the strand of bacteria in place would be very difficult or impossible even in very thin sections. It is much more probable that the wall is merely softened, the cellulose digested at the point where the bacteria are—a soft place large enough in area to permit the bacteria to enter either by actual locomotion or by the formation and growth of new cells in this direction. The little group of bacteria on the surface and near the tip of a root-hair is very often at the point of greatest curv- ature of the hair. This curvature is due to the bacteria. The bending is the evident response to irritation. The irritation may consist in the softening and partial solution of the cell-wall by enzyms formed by the bacteria—a mechan- ical irritation—or in the stimulation of the cell by the same Bot.—VoL. II.] PEIRCE—ROOT-TUBERCLES. 303 or other substances. In the one case we have a traumatropic bending, in the latter a chemotropic. (See Spalding, 1894.) This bending is entirely different in appearance and dis- tinct in cause from that which carries the root-hairs closely around particles of soil. This last is due to irritation by contact (thigmotropism) and by water (hydrotropism). Since the majority of infected root-hairs show the bend- ing at or near the tip, as shown in figs. 1-3, we may infer that the bacteria enter uninjured hairs which are able by growth curvatures to respond to mechanical or chemical stimuli. If the hairs were broken, the ability to respond, and the responses (curvatures) would be greatly lessened, and instead of a short, close spiral (figs. I-3), or a pro- nounced bend, we should have little or no curvature. The curvature of a broken hair is doubtful, and for mechanical and physiological reasons certainly difficult to understand. The roots of young Bur Clover grown in sandy soil in the laboratory showed very few broken hairs when I dug up the plants to search for infected hairs. The soil was very friable, but even then I expected to find more hairs broken as the result either of taking up the plants or of their growth in the soil. It would appear, then, that these bacteria are able to soften or dissolve cell-wall, and when they come into con- tact with a root-hair, enter it, whether it is broken or not. The very slow movements, which are all that most observers report having seen in these bacteria, their ability to soften or dissolve cellulose, the small number of infected hairs, and the small number of broken hairs, make Fischer’s graphic description (1897, pp- g1-2) of how the infection of leguminous plants takes place at least doubtful though no less graphic and interesting. He says: ‘‘ Die feinen Wur- zelharchen einer jungen, noch knéllchenfreien Legumino- senpflanze schieben und drangen sich iiberall zwischen die Bodentheilchen ein, um hier Wasser und mineralische Salze aufzunehmen, ja sie scheiden sogar besondere Stoffe aus, um die Erdteilchen, mit denen sie dicht verkleben, zu lésen. So wird schon die unverletzte Oberflache der Wurzeln 304 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER. chemotaktisch wirkende Stoffe vielfach absondern. Dazu kommen noch zahlreiche verletzte Wurzelhaare oder andere leichte Wunden der Wurzel, die anlockend auf Kndllchen- bakterien wirken werden, wenn diese in den wasserer- fiillten Raumchen zwischen den Bodentheilchen herum- schwarmen. Wovon hier die Bakterien leben, bedarf noch weiterer Untersuchung, denn sie miissten hier natiirlich mit bescheideneren Kohlenstoff- und Stickstoff quellen vorlieb nehmen als in der Reinkultur mit Asparagin und Zucker. Gerade solche Stoffe, besonders das chemotaktisch sehr wirksame Asparagin ist in den Keimpflanzen der Legumi- nosen stets reichlich enthalten und wird bei jeder Verletzung der Wurzel hervortreten. So konnte ihm wirklich die Rolle des Anlockungsstoffes fur die Kndéllchenbakterien zufallen, die in ein aufgerissenes Wurzelhaar genau so einschwarmen wiirden, wie in eine mit Asparagin gerfiillte Kapillare.’’ Fischer’s conviction, expressed at length and supported as well as possible by example and argument (1. c., pp. 131-2), that bacteria do not enter uninjured plant-cells and hence cannot produce disease by being par- asitic on or in plants, is probably responsible for this state- ment, which seems to me the opposite of correct. If it can be shown that bacteria of any one species penetrate the cell-wall of healthy uninjured plants, producing unusual growths therein, Fischer’s contention that there are no bacterial diseases of plants breaks down. On this question Smith (1901) takes issue with Fischer, and seems to prove that there are bacterial diseases of plants. To help himself over the unavoidable difficulty of the tubercle bacteria entering the roots of plants, Fischer says (l. c., p. 92): ‘*Ja es scheint sogar, als ob die Legu- minosen durch Auflockerung der Zellwande an manchen Wurzelhaaren u. s. w. die Anlockung der bacterien vorbe- reiten. In dicht gedrangten Ziigen dringen sie von der Ober- flache der Wurzel in deren Inneres vor, wobei ihnen wie- derum die Leguminose den Weg zu ebnen scheint dadurch, dass sie die schwer durchdringbaren Zellwande etwas auf- lockert.’’ Such a loosening or softening (Auflockerung) BotT.—VOL. II.] PEIRCE—ROOT-TUBERCLES. 305 of the cell-walls is well enough known in other cases; but it is not the host which softens its own walls in order to facilitate the entrance of a foreign organism, but rather the foreign organism which, by enzyms secreted by itself, softens or dissolves the walls of its host which lie across its path of growth.* Having entered the root-hair by softening or dissolving a small portion of the cell-wall, and moving or growing through this, the tubercle bacteria multiply rapidly, forming a thread-like zoogloea from the infection spot along the hair into the epidermal cell of which the hair is a branch (figs. 2and 3). From the epidermis the infecting zoogleea grows fairly straight into the underlying cortical parenchyma. Figure 4, drawn from one of a series of thin microtom sec- tions stained as previously described, indicates the course of the infecting strand (purple). This course is nearly, though not quite, straight toward the central cylinder of the root, for within a series of five or six sections—a distance of 20-30 ~—the infection thread was traced from the base of the root-hair (7. 2. in fig. 4) to one cell (10) in the layer next to the endodermis of the central cylinder. The cells in this layer are distinguished from the cells of the cortical parenchyma by somewhat larger and denser nuclei. This layer is the one from which the lateral roots arise. The direction of the infection thread—which is solid, and is incorrectly termed infection ‘‘ tube ’’—is too regular not to encourage one to suppose that the course of the growing strand of bacteria is determined by attraction exerted by the host-cells upon the bacteria. This then is chemotropic growth of the strand or, if the bacteria are motile in the cells, chemotactic movement of the bacteria. The course of the thread is toward the conducting tissues of the host. This is similar to the growth of the haustoria of Dodder, Cassytha, Viscum, Phoradendron, and other phanerogamic parasites (Peirce, 1894; Cannon, 1901). The growth does 1 See De Bary (‘‘Morphology aud Biology of the Fungi, Mycetozoa, and Bacteria,” Oxford, 1887) and many others as to this in fungi, and Peirce (Annals of Botany, 1894) as to Dodder. 306 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. not extend into the central cylinder and the conducting tis- sues, so far as I have seen. Instead, in the layer of cells just outside the endodermis of the root, division takes place in the cell into which the infection thread has penetrated and in the cells adjacent to it. The daughter-cells grow, repeated divisions and growth follow, and there arises a conical mass of cells which are somewhat larger, and which contain more protoplasm, than the adjacent cortical paren- chyma cells. This conical mass is the young tubercle. At first all of its cells are merismatic, but later the divisions become more and more limited to the cells near the rounded apex of the blunt cone. Thus a regular cambium is differ- entiated in the tubercle. This cambium, as shown in fig. 7, lies near the tip of the tubercle, and forms a bowl-shaped or shallow thimble-shaped layer. The growing tubercle pushes out the overlying cortical parenchyma and epidermis, forming an increasing swelling on the side of the root. Cortical parenchyma and epider- mis, at least for a time, nearly keep pace with the growth of the tubercle. Thus, although the cortical cells are com- pressed somewhat, the epidermis is not ruptured, and the tubercle does not burst out of the side of the root as a lateral root does. The layer of cells which, on infection, gives rise to the tubercle, forms new cells not only centrifugally but also centripetally, so that by these new and growing cells the tubercle is pushed outward, away from the central cylinder. In this way the cylindrical mass of the root itself is kept fairly uniform. An older tubercle appears to be attached to the root. Only by tracing its development can one see that it originates internally. ‘The course of development was only very imperfectly traced by Schneider (1893); hence his bold and erroneous statement that ‘‘ tubercles seem always to develop exogenously.”’ Frank (1890, p. 70) states that the tubercles are new organs formed and well nourished by the plant. He com- pares them with galls which are formed by plants at the points attacked by parasites (insects, worms, etc.), and Bot.—VOL. II.] PEIRCE—ROOT-TUBERCLES. 307 adds, to strengthen his comparison, ‘‘ die Wurzelknéllchen sind kein Organwelches der Leguminose urspriinglich eigen ware, ebensowenig wie dies bei den anderen Pflanzen der Fall ist, sondern eine erst von dem Rhizobium angeregte, dann aber selbst aufgebaute Bildung.’”’ Further on, he says that he has repeatedly seen lupines, cultivated in sterilized and uninfected soil, which formed swellings on the roots closely resembling young tubercles, but showing neither infection threads nor any traces of the cell-contents char- acteristic of true root-tubercles. He accounts for this not on the ground of slight infections producing only abortive tubercles, but on the hypothesis that the lupines, accus- tomed for thousands of years to symbiotic existence with the tubercle bacteria, have so firmly acquired the habit of forming tubercles that they begin to form them even before and without infection. On these points I wish again to call attention to the fact that the tissues of the tubercle orig- inate from the same layer of cells as gives rise, by similar divisions, to the lateral roots (see figs. 4, 5; 6). When one compares a very young mass of tubercular tissue, still enclosed in the cortex of the root, with a very young lateral root also still enclosed in the cortex of the root, the resemblance between the two structures is strong. Figures s and 6 show this. Figure 5 is a diagram of a section in which a tubercle and a lateral root are growing side by side and from the same layer. In the figure the tubercle is to the left, the lateral root to the right. Figure 6 is a drawing of tubercle and lateral root on a larger scale, the root to the right, the tubercle to the left. In the tubercle some infection threads show. The tubercle has the same form as the root, but shows no differentiation among its cells. hie lateral root already shows a differentiation of dermatogen and there is a foreshadowing of the vegetative point. Central cylinder and periblem are not yet distinguishable. The cells of the tubercle are larger than those of the lateral root, but the nuclei of the tubercle cells are not proportion- ally larger and most of them are actually no larger than those of the lateral root. 308 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. As tubercle and lateral root grow, the resemblances be- tween the two decrease and finally disappear altogether. The tubercle has no cap! and no central strand of con- ducting tissue. The tubercle cells differentiate into definite tissues more slowly than do those of the lateral root, but near the tip of the older tubercle there is a mass of meriste- matic cells similar to the growing-point of a lateral root (see fig. 7). This meristem forms cells forward and backward as does the growing-point. The central mass of the tuber- cle is proportionally much larger than the central cylinder of the lateral root, but it is wholly undifferentiated. The cortex of the tubercle contains vascular bundles, small and separated from each other by considerable spaces of paren- chyma (see fig. 7), and is enclosed by layers of cork-cells. These may and usually do become powdery on the surface and rub off as the tubercle forms, just as the cap cells do from the tip of a root. In point of origin and in their earliest growth, tubercle and lateral root are similar. In subsequent growth they are more and more dissimilar. Morphologically, then, the root- tubercles are lateral roots. Though called into activity by very different causes, the cells of the pericycle give rise by division to masses of cells which, on the one hand, develop into tubercles, on the other develop into lateral roots. In the one case we know the stimulus which causes the tuber- cle to form. It is the infection of the root-cells down to the innermost layer of the periblem by bacteria. Do lateral roots form as the result of external stimuli or are they the effects of causes internal to the plant? The latter is the less likely from the fact that the size, number, and position of lateral roots varies in plants of the same species according to the soil, to the number and kind of other plants living in the same soil, to the distribution of moisture and other matters in the soil, and to a great number of other factors not now recognized. This subject merits investigation. As 1 According to Life (Botanical Gazette, April, 1901) the roots infected by Axabena and certain other organisms in certain species of Cycas also have no caps, yet he unhesitat- ingly describes them as roots. Bor.—VoL. II.] PEIRCE—ROOT-TUBERCLES. 309 plants are studied with a view to ascertaining the effects of each factor in the environment, it will become more and more evident that many of the effects which are now attributed to internal causes, lumped together under the name inheritance or distributed among the various func- tions of the organs of the body, are the reactions of the parts to stimuli exerted upon them from outside. If the formation of lateral roots by the division of the cells of the pericambium should prove to be the result of external stimuli, it will be found that these stimuli operate upon the cells immediately concerned. The diverse development of tubercle and lateral root, the result of the persistence of the different stimuli which called them into existence, obscures the common morphology of the two organs so that it is only natural that Frank should have called the tubercles new organs. If the tubercles were the result of hypertrophy indiscriminately among the cells of the cortex of the root, as may be the case in several of the species which Frank studied and described, this would be no evidence that the tubercles are morphologically roots. But in Bur Clover the case is clear. Doubtless the same can be made out in other leguminous plants when the origin of the tubercles is studied in properly stained microtome sections of young roots. I have not examined Frank’s Lupinus, bean, pea, etc., for it is a matter costing much time and patience to find just the right stages, and I preferred to study my own plants and to leave a review of the origin and structure of the tubercles of the plants stud- ied by Frank to a later time or to others interested. In Bur Clover, at least, I think I have advanced strong evidence against Frank’s contention that the tubercles are new organs. As to his hypothesis, involving inheritance as one of the factors in their origin and development, that lupines grown in sterilized and uninoculated soil may form small enlargements like young tubercles, I think my demon- stration of the common origin of tubercles and lateral roots also has some significance. Assuming that the soil remained sterile, which is not wholly probable, the plant might begin 310 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. to form lateral roots which, for some unknown reason, aborted while still in the cortex of the mother root. Lat- eral roots are known to do this, and if the abortion took place early enough, the root character of the new formation might be lost if it had already developed. It seems to me much more probable, however, that both causes were in operation; that the soil did not remain sterile; that the plants were infected by so few or so feeble tubercle-bacteria that the tubercles stimulated to begin to form aborted because the infection was not strong enough. If the leguminous plant, or its separate cells, and the bacteria are parasitically associated, the plant would resist the entrance and growth of the bacteria, and would be much more likely to succeed in this if the attacking bacteria were few or feeble. Over- coming the bacteria, the stimulus to tubercle formation ceases, the tubercle remains rudimentary. ‘That infection of sterilized soil by the tubercle-bacteria is possible, and even difficult to avoid, is known to all who have worked on the subject. This, then, rather than inheritance, accounts for the rudimentary tubercles which Frank describes. The bacteria in the infection thread, which grows through the root-hair and the cortical parenchyma cells of the root to the pericambium layer, multiply, but they multiply most rapidly in the infected cells farthest from the surface of the root. New threads form, which grow out into and infect the cells of the mass of new cells composing the embryo- tubercle. Thus a majority of the cells in the young tubercle contain bacteria. Though infected cells do divide (see pp. 322-323), they probably divide less often than the uninfected cells. The primary infection is in a nearly straight line from the root- hair inwards. The infection of the daughter-cells com- posing the embryo-tubercle is accomplished by branching infection threads growing in fairly straight lines radiating from the base of the tubercle. In this way the cells near the base of the growing tubercle are most infected, those near the tip least. It may be in consequence of this that the cells at and near the tip of the tubercle retain their Bot.—Vot. II.] PEIRCE—ROOT-TUBERCLES. 311 merismatic quality, and that they form the bowl-shaped layer or layers of meristem which continue the growth of the tubercle (see fig. 7). The infection threads grow out toward the tip of the tubercle (fig. 10), but the meristem continues to form new cells between itself and the cells containing bacteria and infection threads. By the layer or layers of uninfected daughter-cells the meristem protects sts own cells from infection. Perhaps because they escape infection, they retain their ability to divide. If they can be prevented from forming a sufficient number of daughter- cells to enable them to escape infection, what will be the result? To answer this, and some other questions, I tried the following experiment. I imbedded young tubercles on growing roots of potted plants of Bur Clover in plaster of Paris, according to the method devised by Pfeffer (1892) and used by his pupils (Newcombe, 1894, Richter, 1894, etc.). The roots were disturbed as little as possible and were put back carefully in the soil as soon as the plaster had been applied to the tubercles. The plants were grow- ing in coarse sandy gravel, so that it was not difficult or dangerous to lift out one or more roots when the soil was well loosened by being soaked with water. After the lapse of twenty-nine days, I again took up the roots, cutting them off for careful examination. Two tubercles had been firmly held at the base by the plaster, but had broken the casts sufficiently at the tips to grow fresh and pink out of their investments. New tubercles seemed to have been formed since the roots were operated upon. One tubercle, which had stayed in its cast, was taken out and sectioned by hand. It was evidently dying. The bacteria in it seemed active, but were fewer and much smaller than in unconfined tuber- cles. There was no meristem; the cells of the tubercle were in their definitive condition; there were smaller quan- tities of starch in this than in ordinary tubercles. What was true of this tubercle was equally true of others which had failed to break and grow out of their casts. (2) June 18, 1902. 312 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. It would seem, then, from the results just described, that the tubercle meristem is preserved from loss of its meris- matic qualities by escaping infection, and that the presence of bacteria in the cells ultimately costs them their power of division (see pp. 322-323). The meristem near the tip of the tubercle is a survival, as is the vegetative point at the root-tip, of the merismatic cells which constitute the tubercle and the root in their embryonic condition. The result of imbedding an infected tissue, a root-tuber- cle, in plaster, is different from that described by Newcombe (1894), who worked on healthy plants. He says that the external mechanical resistance causes developing cells to attain their definitive condition more slowly, continues the merismatic power and activity of the cambium cells, and prolongs the life of such cells as ordinarily die early. Newcombe worked on healthy plants, and his results exhibit the effect of pressure only on growing cells and organs. In my experiment the pressure which checked the growth of the tubercle-cells may not have mechanically affected the bacteria. Since the bacteria and the cells of the tubercle are competitors, the plaster investments handicapped the latter to such an extent that the ultimate results of bacterial activity appeared earlier than in unconfined tubercles, the bacteria gaining an advantage. To the action of the bac- teria rather than of the plaster cast are due the early loss of merismatic power and the early assumption of their definitive condition by the cells of the tubercle. Ill. Tue Form AND DISTRIBUTION OF ROOT-TUBERCLES. The form and distribution of root-tubercles merit some discussion, and since those of Bur Clover are typical I will continue to describe them. The tubercles grow both in length and in thickness at the ends, not at their bases, and thus become club-shaped. They may and often do branch. The growth takes place solely at the tip, because the only meristem is there. The tubercle tissue is supplied with food through vascular bundles which are neither large Bor.—Vot. II.] PEIRCE—ROOT-TUBERCLES. 313 nor numerous but adequate for a time. A lateral root grows both at the tip and throughout its length, thickening and elongating. In this way its cylindrical form is main- tained. The lateral root contributes food-materials and water in increasing amounts to the plant which forms it. The tubercle receives food from the plant. Perhaps it contributes also to the nutrition of the plant. Experiment so far seems to justify this belief. But if the tubercle were altogether beneficial and increasingly so, one would suppose that it would grow at the base, by secondary thickening, as well as at the tip, by primary growth, in order through increasing conducting tissues to contribute more and more to the nutrition of the plant. The absence of such secondary growth and the ultimate fate of the tubercle—dying and being cut off—suggest that the leguminous plant limits as far as possible the supply of food to the tubercle, and finally stops it. Herein we have another item of evidence against Frank’s hypothesis that the leguminous plant encourages tubercle formation. It does not cut off the tubercle immediately ; the irritation which results in tubercle formation is too great and the osmotic demand for food is too strong to be resisted at once by the plant. Only after a time is tubercle growth checked—perhaps by the remoteness of the tubercle meristem from the source of food-supply—and later still, the tubercle is cut off. The tubercles may be rosy pink at and near the tip, creamy white elsewhere, nearly or quite the same shade as the roots bearing them. Later, the oldest tubercles, those nearest the surface of the soil, may branch, taking on a flat, though thick, fan shape. By no means do all of the tubercles branch. Those very near or almost on the surface of the soil do not, and in the lower half of the infected portion of the root-system I have seen almost no branched tubercles. The difference in age between the branched and unbranched tubercles in the same or the adjacent layers of soil is not sufficient to account for the difference in form. The branched tubercles are the first ones to lose the plump and healthy appearance of active life; they grow thin and 314 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. shriveled. They have grown fast, attained maturity early, and they die young. The reasons for this are probably two: Furst, in the upper layers of soil, which are certainly best aérated, the bacteria in the tubercles obtain the uncom- bined nitrogen which they absorb and fix (Mazé, 1897) more readily and more abundantly than those in tubercles farther down; hence, second, they grow and multiply more rapidly, the tubercle-cells are irritated proportionally. Be- cause the rapid growth and multiplication of the bacteria, and, probably as a consequence, of the tubercle-meristem cells also, are not uniform, branching occurs as a result of some parts of the tubercle growing faster than others. The greater activity of the bacteria and of the host-cells in these branched tubercles is not accompanied by adequate, much less proportional, growth of the base, and of the conducting tissues in the base, of the tubercle. These tubercles sooner cease to receive as much food from the leguminous plant as they need, and hence are the first to die. Since these branched tubercles are the largest and contain most bac- teria, one would suppose they would benefit the plant more than the smaller ones (if any tubercles are beneficial), and that they would be best supplied with afferent and efferent conducting tissues, as indicated by the proportional size of their bases. This is not the case. The vertical distribution of tubercles on the roots has been reported by Frank (1890, pp. 22-3). The greatest number and the largest tubercles occur on the lupine within seven centimeters of the surface of the soil, and there is a rapid decrease in both number and size as the depth increases, till below fifty-three centimeters none was found. The distribution on Bur Clover roots corresponds. The strictly aérobic character of the tubercle bacteria, as shown by artificial cultures, accounts for this, but the relations of these organisms to the uncombined nitrogen of the air as well as to the oxygen should be borne in mind when these bacteria are said to be aérobic. The distribution of air in a soil varies with the nature of the soil—a well drained gravelly soil being well aérated to a greater depth than a Bot.—VOL. II.] PEIRCE—ROOT- TUBERCLES. 315 heavy, compact, clay soil. It also varies with the tillage. I have found tubercles much lower on the roots of Bur Clover growing on a heap of gravel than in an undisturbed and compact clay. Mechanical reasons are insufficient to account for this. The only inference to be drawn is that the bacteria are limited in their natural distribution to those soils and those layers of soil which contain considerable volumes of air, for only there will they find enough oxygen and nitrogen for their needs. It is the general habit of leguminous plants to send their roots fairly deep into the soil. In a natural field, or one returned to a state of nature, where the soil is covered by a mixed vegetation, it is found that different plants send their roots to different depths. In this way the resources of the soil are more perfectly exploited by the plants and destructive competition is avoided. But it is to be noted that of those plants which send their roots deeper, many are members of the Legwminose. Is this merely to escape competition with other seed-bearing plants, or to reach a more abundant and constant supply of water, or to escape the attacks of the bacteria which cause them to form tuber- cles? It is mainly the Leguminose which are successfully attacked by tubercle bacteria, and they, as a rule, send their roots fairly deep into the soil. Furthermore, the number of roots increases with the distance from the sur- face. It would appear not inconsistent with the evidence so far obtained, to suppose that the habit of the Leguminose of sending their roots deep, and of causing them to branch copiously only after they have reached some depth, is one means which these plants have of avoiding the attack of tubercle bacteria. Alfalfa or Lucern (Medicago sativa Linn.) has notori- ously long roots. They are reported to go down to the permanently water-bearing levels of the soil. The best chance for examining these roots would be offered when a well is being dug where Alfalfa is growing. I have not been so fortunate as to have such a chance, but digging around Alfalfa plants growing as weeds in a grass field 316 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. shows that below a depth of twenty centimeters from the surface the number of tubercles decreases rapidly. The roots of this plant are perennial, and the new roots each season are most of them formed so far below the levels in which tubercles ordinarily occur on the roots of leguminous plants that this plant should form a good test object of the vertical distribution of tubercle bacteria in the soil. Com- pared with such annuals as Bur Clover, a member of the same genus, there were far fewer tubercles on Alfalfa roots than on Bur Clover at the time that I dug around Alfalfa (December). There were young roots near the surface as well as further down, but the greater number of young roots must have been formed far below where I reached by digging, for there were not enough young roots above to meet the needs of the plant. Alfalfa and many other perennial Leguminose, may therefore form the majority of their new roots each year so deep in the soil that they can- not become infected. As the tubercles are not perennial, whatever advantage may accrue to the perennial legumi- nous plant by its association with bacteria would be limited in time and quantity to the early life of the individual, when its roots were all in the layers of soil containing active tubercle bacteria. The question is well worth study. IV. THE STRUCTURE OF ROOT-TUBERCLES. The structure of a tubercle is shown somewhat diagram- matically in fig. 7. This is a sketch, at a magnification of thirty-five diameters, of a section of a young and still grow- ing tubercle. The section is parallel with the long axis of the tubercle and at right angles with the root. The cam- bium of the tubercle lies between a and 4, parallel with and in the broken curved line. This meristem is composed of two or three layers of cells. Those toward the periphery of the tubercle as well as those toward the center divide, the cells toward the periphery differentiating rapidly into cells which round off from one another and form the pow- dery, cap-like tissue which wears away but protects the meristem within just as the root-cap protects the BotT.—VOL. II.] PEIRCE—ROOT-TUBERCLES. 317 growing-point of the root. The cells toward the center differentiate somewhat diversely according as they become infected by bacteria or continue free from them. The un- infected cells remain comparatively small, and present the characters of ordinary parenchyma cells, the protoplasm becoming vacuoled and containing numerous starch grains. There may be several vacuoles in these cells or one tra- versed by strands of cytoplasm. The infected cells grow larger and in their definitive con- dition are from half as large again to twice as large as the uninfected cells. This increase in size may be attributed to one of three causes: First, to the stimulation of the protoplasm by the bacteria and the substances produced by them in the cells; second, to the actual irritation (in- flammation) of the protoplasm; and third, to the increased pressure set up in the cell by the rapidly multiplying bacteria. By the plaster of Paris method we can test the relative value of two of these influences; the third must be de- termined by ascertaining microscopically the actual con- dition of the protoplasm of infected cells. On imbedding the young tubercles of growing roots, as above described, the pressure normally or abnormally developed in the cells will be resisted by the plaster, the cells expanding against the plaster will be subjected to compression. Nine days after the tubercles were enclosed in plaster, I opened the casts and sectioned the tubercles by hand. There were many more starch-grains in the uninfected cells than in ordinary tubercles; I saw no infection threads (I did not fix and use the triple stain previously described and hence infection threads might have been present which escaped my notice), the bacteria and bacteroids were smaller, and the general appearance suggested that the leguminous cells were better able to bear confinement than the bacteria were. Increase in size both of the tubercle and of its com- ponent cells being prevented by the plaster investment, the bacteria have no increasing space in which to grow, and continuing to multiply, for a time at least, they remain 318 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. small, in consequence consuming less of the food provided for them by the leguminous plant. Starch can therefore accumulate in the uninfected cells from which the infected cells are osmotically supplied with food. Depositing the non-nitrogenous food in solid form (starch) of course re- duces the turgor pressure in the cells of the tubercle and thereby reduces the resistance to the plaster investment. The component cells of the tubercle enclosed in plaster of Paris remain smaller than in unconfined tubercles, the in- fected and uninfected cells being more nearly equal in size. Enclosing a tubercle in plaster may diminish the stimulus or inflammation produced in the cells by the bacteria, but as the bacteria survive and multiply there can be no great diminution of their chemical effect on the cells in which they occur. The physical result, pressure, is much more affected by the plaster investment. Since the infected cells remain more nearly the same in size as the uninfected ones, the inference is plain that the excessive increase in size of the infected cells is due to increased pressure in them. The infected cells, as shown by figure 8, are thin-walled and contain only one large vacuole. This is not traversed by cytoplasmic strands. The quantity of bacteria may vary in infected cells, and with this there is a corresponding vari- ation in the appearance of the cells. Thus, fig. 8 shows a typical infected cell in which the bacteria have multiplied enormously, while fig. 9, magnified about one-third larger, represents a cell in which there are comparatively few bac- teria, most of which are at the point indicated by the line from &. In the cells containing relatively few bacteria there may be some starch-grains, as indicated by the line ‘from S. In unstained hand-sections, the degree to which the older parenchymatous cells in the central part: of a tubercle are infected is indicated at a glance by the amount of starch in the cells, the cells with the average amount of bacteria containing no starch-grains, the cells with no bac- teria containing many starch-grains, the cells with few bacteria containing starch grains in inverse proportion to BotT.—VoL. II.] PEIRCE—ROOT-TUBERCLES. 319 the number of bacteria. The cells with few or no bacteria receive more non-nitrogenous food than they consume. The excess they deposit in solid form as starch. The cells with many bacteria presumably receive at least as much non-ni- trogenous food; but either they themselves or the bacteria in them consume this so that there is no excess to deposit. From the cells toward the center of the tubercle the new cells formed by the tubercle meristem are infected by means of infection threads running fairly straight toward and into the daughter-cells of the meristem (see fig.10). This figure, magnification two hundred, was drawn from a thin micro- tome section of a young tubercle, and is colored as nearly as possible like the cells of the preparation. The prepara- tion was stained, as previously described, by Flemming’s triple stain, Gramm’s iodine solution being used after the anilin gentian violet in order to differentiate the strands of bacteria. The cell walls are drawn in black, though they were only very faintly stained and of course were not black. The cytoplasm is brownish yellow from orange G., the nuclei a somewhat deeper shade of the same color, the nucleoli red from anilin safranin, the infection threads pur- ple from anilin gentian violet. In a fresh preparation, whatever starch grains are present in the section are stained the usual color by the iodine, but this color is fugitive. The arrow to the left in this figure (fig. 10) shows the direction in which the tubercle meristem lies. It is evident from this figure that the infection threads run very definitely toward the new cells formed by the merismatic layer. Since all the food of the tubercle cells comes from the opposite direc- tion, from the root, there must be some other directive influence than this food to cause the infection threads to — grow so definitely toward the tip of the tubercle. This influence must come from the cambium or from its daugh- ter-cells, and must consist in the substance or the products of these cells rather than in the food supplied to them. If we are to assume any chemotactic influence, it must be exerted by some diffusing substance or substances and 320 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. hence must be by the products and not by the living sub- stance of the meristem and its daughter-cells. The direction of growth of these infection threads cannot be determined by the oxygen or nitrogen (or both) of the air, for if this were the case, we should find strands of bacteria running from the central cells in all directions toward the periphery of the tubercle. This is not the case. The strands run toward the growing-point of the tubercle. In consequence, the daughter-cells successively formed by the repeated divi- sion of the cells of the meristem become infected. Not only do the infection threads run definitely toward the growing-point of the tubercle; they also grow toward the nucleus of each cell which they enter. This statement has been repeatedly made and denied in papers on the subject of root-tubercles. In hand sections, especially if the mate- rial were not carefully fixed and differentially stained, it would be easy to find evidence in support of the affirmation and of the denial. Microtome sections, differentially stained as before described, of carefully fixed growing tubercles of the species of leguminous plants which I have especially studied, show that in most cases the infection threads run definitely toward the nuclei of the tubercle cells. This is evident in fig. 10. Figures 11 and 12 also show this. In fig. 13 are shown two tubercle cells in which the main in- fection thread is not directed toward the nuclei, but the lower of these two cells shows that the infection thread bends toward the nucleus. In the next section of the series (not figured) a branch runs from the main infection thread to the nucleus. In the cell shown in fig. 11 the infection thread is divided, one part running beneath, the other above the nucleus. For the sake of clearness this upper part of the thread was omitted in drawing. The nucleus of the upper cell in fig. 13 was not in the plane of the sec- tion. An adjacent section (not figured) contains this and has a branch of the main infection thread running to it. There must be some reason for this definite growth of the strands of bacteria toward the nuclei of the cells which Bot.—VoL. II.] PEIRCE—ROOT-TUBERCLES. 321 they enter.’ The effect on the nuclei is marked, as will be shown presently. - The changes which take place in infected cells as they develop are indicated in figs. 14 a, 6, ¢, d. These repre- sent successive cells from the meristem backward toward the center of the tubercle, which is shown in fig. 7. A part of the cambium layer at a—é in fig. 7 is shown at #-y in fig. 14 a. This series is stained by Flemming’s triple stain, but Gramm’s iodine was not used. For this reason the bacteria and infection threads are not differentiated. The magnification of figs. 14 4, c, d is 300, that of fig. 14 a is 270, hence 14 6 and 14 @ do not meet exactly. The fig- ures in the series had to be drawn separately by reason of the limited field of the objective which was used to give the necessary magnification. Two cells of the tubercle cambium are shown at +-y. Recently formed daughter-cells lie toward the outside as well as toward the center of the tubercle. The cell z is already beginning to show the effects of infection, vacuoles of considerable size, which later become confluent, form- ing in the cytoplasm, and a distinct vacuole, like a halo, appearing around the nucleolus. As shown by the cells further toward the center of the tubercle, the nucleolus is the first part of the nucleus to be evidently affected by the presence of tubercle bacteria in the cell. It is the first part of the cell to decrease in size and to disappear. If the nucleolus is in fact an accumulation of food in the nucleus, one would expect it to disappear, to be consumed, when- ever there arose a special need of food in the nucleus, or even in the cell as a whole perhaps. Furthermore, the 1W. Magnus (i900) discusses the endotrophic mycorrhiza fungus of WNeottia nidus avis I,. in this connection, stating (pp. 7-9) that the hyphe of this fungus do not grow toward the nucleus of the cell with any regularity, but that in many other parasitic fungi the hyphz do grow toward and around the nucleus, in some cases, however, with no greater regularity than toward and around starch-grains or other solid contents of the cell. He adds (p. 61): ‘‘Dass sich parasitare Pilze mit ihren Haustorien oft an den Zellkern legen und sich in seiner Nahe eigenthiimlich verzweigen, gestattet keinen Riickschluss auf die Bedeutung des Kernes als Nahrungscentrum der Zelle.’?’ To con- clude, because fungus hyphe or strands of bacteria grow toward the nucleus ofa cell which they have entered, that the nucleus is the center of nutrition of the cell is illogical; but it may well be in these cases that the nucleus contains or produces sub- stances which nourish and attract the parasites. 322 CALIFORNIA ACADEMY OF SCIENCES [PRoC. 3D SER. greater solubility of the substance of the nucleolus than of the substance of the nucleus and cytoplasm would make it the first to disappear under the attack of a parasite. In fig. 14 4 two characters of the cells are especially noticeable: the nuclei are becoming disorganized, cavities taking the place of the nuclear substance; and the cyto- plasm is no longer so dense as in the younger cells shown in fig. 14 a, nor does it stain so deeply with the orange G. The degeneration of the nucleus is especially marked in the cells c, 7, A. These cells have lost their nuclei, except the last traces of acromatic substance, before they have themselves grown to their full size. The two upper cells of fig. 14 ¢ have reached the maxi- mum size of infected cells. It is impossible to say whether they have grown to this size after their nuclei were reduced to the condition depicted or while their nuclei were being destroyed. I cannot say whether the smaller cells c, 2,4 of fig. 14 6 would ever have grown to the size of the upper two in fig. 14 4, but the unusual growth of infected cells is due to the bacteria, and it is the bacteria which also destroy the nucleus or cause it to be destroyed. In the upper cells of fig. 14 c the cytoplasm is no longer the clear color pro- duced by orange G. This brownish yellow color is dulled by the purple of the now extremely numerous bacteria. The smudgy appearance of the cytoplasm of the cells in 14 d is due to the mixture of the purple stain of the bacteria and the brownish yellow of the cytoplasm, an effect which is very striking in the preparations. The cells of fig. 14 d also show the greatly reduced nuclei and the large central vacuoles. Fig. 8 represents a cell from near the center of the tubercle shown in fig. 7. The cytoplasm is crowded with bacteria, the central vacuole is very large, the nucleus is reduced to an elongated lumpy mass as seen in section, or to a thin lumpy plate in the entire cell. When infected cells contain any considerable number of bacteria, they cease to be able to divide. Freshly infected daughter-cells of the cambium layer do divide, as fig. 15 shows. One is inclined to say that the disturbing, if not Bot.—VOL. II.] PEIRCE— ROOT-TUBERCLES. 323 already destructive, effect of the bacteria upon the nucleus and other parts of the cells causes them to lose their power of division. As has been previously pointed out (p. 311), be- tween the tubercle meristem and the infected cells there normally lie two or more layers of uninfected daughter- cells. These divide also and grow. By this means the meristem cells are kept from infection. When, however, the growth of these daughter-cells is prevented, and the rate of division of the meristem cells is reduced by such mechanical pressure as a plaster of Paris investment of the whole tubercle, the infection progresses to and into the merismatic cells and they presently lose their merismatic character and take on their definitive form. One result of the division of infected cells is the produc- tion of new cells already infected and therefore not requiring the entrance of infection threads. As shown by Hedlund (1892) and myself (1899), the gonidia of lichens frequently divide in such directions as to exclude the haustoria from some of the daughter-cells. No such result can follow when a cell contains a number of minute parasites distrib- uted in its cytoplasm. The cell may divide under these conditions, but the daughter-cells share the parasites as well as the substance of the mother-cell. The figures above referred to, especially figs. 14 a, 4, ¢, d, and fig. 8, plainly show that the presence of tubercle bacteria is not beneficial to the cells which contain the bacteria. So far as the relations of the bacteria and their host-cells are concerned, no one would hesitate to call the association a clear case of parasitism of bacteria in the cells. Whether it is a benefit to the leguminous plant to form tubercles, to harbor bacteria in these, to have the cells of the tubercles destroyed, and all the food supplied to the tubercle consumed, is another question. The most careful experimentation and the critical examination of the results of experiment have so far led to the generally accepted belief that the association of bacteria with the roots of leguminous plants is beneficial to the leguminous plants. I find it hard to understand how association with bacteria 324 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. which destroy the cells in which they occur can benefit the larger member of the association. The bacteria are known to fix free nitrogen from the air. By this means they form in the cells of their host-plants nitrogenous compounds which the host may use. Apparently the bacteria form more than enough of these valuable nitrogen compounds in the cells of their hosts to compensate for the extra material used in forming and maintaining tubercles the cells of which are ultimately destroyed by the bacteria. This implies a mar- velous balance of profit and loss, the more remarkable since the profit apparently exceeds the loss. One point more needs to be made clear. Miss Dawson (1899, p. 14) says that it is difficult to conceive how such strictly aérobic bacteria as these can flourish in the cells of such compact tissue as composes the tuber- cle. This difficulty is of her own conceiving, for do not the cells of the tubercles respire and are they not neces- sarily supplied with oxygen for respiration? Again, inter- cellular spaces in the infected tissue do occur, as figures 14 5, 14 c, 13, 12, 15, 8, and g plainly show. Even if intercellular spaces did not occur, as asserted by Schneider (1893, pp. 786 and 787), the existence of the living cells of the tubercle tissue would prove the presence of sufficient quantities of oxygen, if not of nitrogen, in the tissue and therefore in the cells. Unless we are to imagine anaérobic respiration for these cells, it is unnecessary to assume it for the bacteria which infest them. Fischer (1897, note 63 to p- 92), shows this clearly, and my sections reveal the pres- ence of intercellular spaces through which a diffusion of gases could take place even if the diosmosis of solutions of the gases concerned were inadequate to supply the demand. SUMMARY. 1. Though the bacteria which form root-tubercles on leguminous plants are usually only slowly motile, if motile at all, in artificial cultures, this proves nothing as to their motility in the soil. Bot.—VOL. II.] PEIRCE— ROOT-TUBERCLES. 325 2. Io. The proportion of root-hairs infected to the total num- ber formed is small, in one case computed to be 1:1000. Given the contact or close proximity of tubercle bac- teria with the root-hairs, infections may take place in great numbers simultaneously, at least when the roots are very young. Infection of the root may be resisted by cutting off the infected ends of the root-hairs. The tubercle bacteria enter and infect a root-hair by softening or dissolving a small portion of the wall and moving or growing through this. There is no evidence that they usually enter through broken root-hairs, and the curvatures of infected root-hairs are evidence against these hairs having been broken at any time. The infection thread grows fairly straight, being chemotropically attracted, through the cortical paren- chyma, from the root-hair to the layer of cells next outside of the central cylinder of the root. The tubercles originate only endogenously and from the same layer as gives rise to lateral roots. We may therefore conclude that the tubercles are mor- phologically lateral roots, though greatly modified by the influence which caused them to be formed. Tubercles form only as the result of stimulation by bacteria. Do lateral roots form as the result of internal causes or of external stimuli? The growth of the tubercle is apical, the daughter-cells of a bowl-shaped terminal meristem constituting the growing part of the tubercle. There is little or no secondary growth in thickness. Because of this, the conducting tissues do not keep pace with the growth of the tubercle. The growth of the tubercle is correspondingly limited. Tubercles are largest and most numerous near the surface of the soil. It is possible that perennial Leguminose form few if any tubercles after their roots have grown deep into the soil. 326 Il. 12. ie 14. THe 16. 17. CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. The presence of bacteria in the cells of a tubercle pre- vents the infected cells from forming starch granules. Uninfected cells do not attain the size usually reached by infected cells. The larger size of in- fected cells is due to increased pressure, probably also the greater irritation, in these cells. The bacteria cause the degeneration and almost com- plete destruction of the nuclei of the cells in which they occur. The infection strands grow definitely, chemotropically, toward the daughter-cells formed by the tubercle meristem, and seem also to grow definitely toward the nuclei of the cells into which they penetrate. Infected cells soon lose their power of division, though not of growth. The presence of bacteria in the cells of the tubercle is injurious to these cells, and the relation of the bac- teria to their host-cells is parasitism. It is difficult to understand how the leguminous plant as a whole can profit by an association which is injurious and finally destructive to the cells in which the bacteria occur. Intercellular spaces occur in the tissues of root- tubercles. Even if they did not, it would not be necessary to assume that the bacteria live anaérobi- cally therein, since the tubercle cells do not live anaérobically. LELAND STANFORD JR. UNIVERSITY, PALO ALTO, CALIFORNIA, July 27, 1901. Bot.—Vot. II.] PEIRCE—ROOT-TUBERCLES. 327 1890. 1893. 1898. 1899. 1900. 1897. 1900. 1896. 1892. 1899. 1894. 1893. 1897. IgoI. 1894. Igol. 1893. I9oI. 1892. 1894. 1894. 1897. 1900, BIBLIOGRAPHY. FRANK, A. B. Ueber die Pilzsymbiose der Leguminosen. Sonder- abdruck aus den Landwirthschaftlichen Jahrbiicher. Berlin. ZIMMERMANN, A. Botanical Microtechnique. New York. Transl. by Humphrey. Hor, A. C. Histologische Studien an Vegetationspunkten. oft. Centralblatt, Bd. LXXVI, pp. 65-69. Dawson, Maria. Nitragin and the Nodules of Leguminous Plants. Phil. Trans. Roy. Soc. MicuLa, W. System der Bakterien. Bd. I, 1897; II, 1900. Jena. ZINSSER, O. Ueber das Verhalten von Bakterien, insbesondere von Knollchenbakterien, in lebenden pflanzlichen Geweben. /ahrdé. /. wiss. Bot., Bd. XXX. Dawson, Marta. Further Observations on the Nature and Functions of the Nodules of Leguminous Plants. P#il. Trans. Roy. Soc. CzaPeK, F. Zur Lehre von den Wurzelausscheidungen. /ahré. /. wiss. Botanik, Bd. XXIX. HEDLuND, T. Kritische Bemerkungen tiber einige Arten der Flech- tengattungen Lecanora, Lecidea, und Micarea. Bihang till K. Suen. Vet.-Akad. WHandl., Bd. XVIII. PEIRCE, G. J. The Nature of the Association of Alga and Fungus in Lichens. Proc. Cal._Acad. Sct., 3d Ser. (Bot.), Vol. I, No. 7. SPALDING, V. M. The Traumatropic Curvature of Roots. Ann. Botany, Vol. VIII. PEIRCE, G. J. Structure of the Haustoria of Some Phanerogamic Parasites. Ann. Botany, Vol. VII. FIscHER, A. Vorlesungen tiber Bakterien. Jena SMITH, E. F. Entgegnung auf Alfred Fischer’s ‘‘ Antwort’ in be- treff der Existence von durch Bakterien verursachten Pflanzenkrank- heiten. 2ter Theil. Centralbl. f. Bakteriol., Parasitenkunde u. infektionskrankh., 11 Abtheilung. (The special literature is here cited.) Peirce, G. J. A Contribution to the Physiology of the Genus Cus- cuta. Ann. Botany, Vol. VIII. Cannon, W. A. The Anatomy of Phoradendron villosum Nutt. Bull. Torr. Bot. Club, Vol. XXVII. SCHNEIDER, A. The Morphology of Root-tubercles of Leguminosz. American Naturalist, Vol. XXVII. Lire, A. C. The Tuber-like Rootlets of Cycas revoluta. Pot. Ga- zette, Vol. 31. PFEFFER, W. Ueber Anwendung des Gipsverbandes fiir pflanzen- physiologische Studien. er. d. K. Sachs. Gess. d. Wiss. Newcomes, F.C. The Influence of Mechanical -Resistance on the Development and Life-period of Cells. Bot. Gazette, Vol. XIX. RICHTER, J. Ueber Reaktionen der Characeen auf dussere Reize. Flora, Bd. LXXVIII. Mazé. Fixation de I’azote libre par la bacille des nodosités Légumi- neuses. Ann. de l’Inst. Pasteur, T. x t. Macnus, W. Studien an den endotrophen Mycorrhiza von Neottia Nidus avis L. /ahrd. f. wiss. Bot., Bd. XXXV. 328 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER. EXPLANATION OF PLATE XXIX. Most of the figures were drawn with the aid of a Leitz drawing prism ocular, the rest with an Abbé camera lucida. Fig. 1. Two root-hairs of Bur Clover infected by tubercle bacteria, showing the characteristic bending at the point of infection; x 50. Fig. 2. The lower of the two root-hairs of fig. 1. The mass of bacteria in the concavity of the coil and the infection thread running from this point through the hair; x 300. ; Fig. 3. Another infected and coiled root-hair with the infection thread growing close to the nucleus of the hair; x 300. Fig 4. A section showing the course of the infection thread from the base of the hair (7. 4.) to the layer of cells which gives rise to the tubercle, the same layer which, under other conditions, would give rise to a lateral root; x 360. Fig. 5. A section showing a young tubercle (left) and a young lateral root (right) developing from the same layer; x 20. Fig. 6. The same section enlarged. In the lateral root (right) differen- tiation of tissues is already taking place, but notin the tubercle (left): x 360. Fig. 7. Longitudinal section of a tubercle, young and still growing. The largest part of the tubercle is composed of bacteria-containing cells. Outside of this mass is the comparatively thin layer in which are the small and scattered vascular bundles. Beyond this is the protective tissue, pow- dery on the outside, which is continuous with the cortex of the root. At a-b is the tubercle meristem, which forms daughter-cells both forward and backward, as does the growing-point of a root; x 35. Fig. 8. Atypical infected tubercle cell, thin-walled, with the cytoplasm surrounding a large single vacuole, and the nucleus reduced to a small lumpy plate. Cytoplasm stained with orange G., but the color obscured and dulled by the purple of the bacteria stained by anilin gentian violet. The nucleus is purplish from the mixture of anilin safranin and anilin gentian violet, a wholly different color from that of a healthy nucleus stained and differentiated by Flemming’s triple stain and Gramm’s iodine solution. Intercellular spaces are shown; x 660. Fig. 9. Tubercle cell containing only small number of bacteria (at 2) and enclosing some starch-grains (.S). The nucleus has degenerated only some- what and the vacuolization of the cytoplasm has not progressed far; x 820. Fig. 10. Section of a tubercle, part of tubercle shown in fig. 7, near the meristem. The direction in which the meristem lies is indicated by the arrow at the side. The section was stained by Flemming’s triple stain and differentiated, after the anilin gentian violet, by Gramm’s iodine. Thus the infection threads are clearly brought out. Note the course of the infection threads, definitely toward the tubercle meristem and generally toward the nuclei of the cells entered; x 200. : Fig. 11. One cell from fig. 10 (nearly in the center of the figure) showing the solid infection strand (zoéglcea) in which the separate bacteria can be distinguished; x rooo. Fig. 12 and 13. Tubercle cells showing the infection threads growing in definite direction, generally toward the nuclei of the cells; x 820. Fig. 14, a-d. A series of cells from the meristem backward in a longitu- dinal section of a tubercle. The section is stained by Flemming’s triple stain, but not differentiated by Gramm’s iodine, hence infection threads do not show. In fig. 14 a, two meristem cells show at 2-y, with daughter- cells both forward and backward. Z=already infected and degenerating cell. In 14 6, degeneration of the cell and especially of the nucleus is shown by cells c, 7,4. Degeneration of the nucleus, obscuring of the color of the cytoplasm by the many bacteria in it, and the formation of the char- acteristic vacuole shows in 14 ¢, especially the two upper cells. Fig. 14d shows the same still more markedly, especially the uppermost cell. In this series the effect of bacterial infection of the cells is clearly exhibited; 14 a, x 270; 14 b-d, x 300. Fig. 15. Division of an already, though recently, infected daughter-cell of the tubercle meristem; x 300. LITH DAUTTON & TUKy.5 ] [PEIRCE ROOT-TUBERCLES OF Bur: CLOVER. 3? Ser. Bor. VouI. An. StI Proc Cans PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES Tuirp SERIES BoTANY . Vou. Il, No. 11 Spindle Formation in the Pollen-Mother-Cells of Cassia tomentosa L BY Henry To AGA Gs WitH THREE PLATES Issued April 30, 1904 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1904 “commitre® on PuBLt we —— >, + Mis, Chairman oo Fuser re oe PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES THIRD SERIES BoTANY Vou ih, No it Spindle Formation in the Pollen-Mother-Cells Of Gla ssia tomentose -L BY Hrewirt (FAs relas WitH THREE PLATES Issued April 30, 1904 SAN FRANCISCO PUBLISHED BY THE ACADEMY 1904 } f Dill al slat SPINDLE FORMATION IN THE POLLEN- MOTHER-CELLS OF CASSIA TOMENTOSA L. BY HENRI T. A. HUS. CONTENTS. PLates XXX-XXXII. PAGE PRESEN UCEION G5 Go oa lelareis Ua wieints ete Grlersleepiaamp nme eave maelal Male ic Eis 329 WOBSEREPTION Cinco sior es 8s keg Rice Wk suave late wy gelatine) uiyy miaveyiahe Pa siprn eye yiia ie eae MO SCUGSIOINN oii cie sao kb dco tartine S siemied armie etnue alesse alana seieterars Chala els rie 341 The Nature of the Multipolar Spindle. ......---.--++++++++++ 341 The Origin of the Cones of the Multipolar Spindle.......-... 343 SU TNR YT ie ot x wha! asses gow psenleldvore Sis oie miele walneal Tole iae oe 346 Pr BWACGRADELVT oo ici: dniere: o dosle eis wiatelh nc emerohodnpelrnetarniose mies ca be esenai hina os 348 EXPLANATION OF PLATES .......0eeececcescese cosscncrcsseccsssceess 350 INTRODUCTION. Cassia tomentosa I.. possesses a number of qualities which particularly recommend this plant to the cytologist. Chief among these are the large number of buds produced and the protracted flowering period. The stages of cell- development which are of greatest interest at the present time are passed through very rapidly, and are also the most difficult to fix; hence the collection of an immense amount of material is necessary to ultimate success. At the same time the protracted flowering period affords an opportunity to determine at leisure which is the best fixing-medium and what shall be the duration of the fixing-period. The following fixing-fluids were tried: [329] April 6, 1904. 330 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. z. Strong Flemming: I per cent. chromic acid 15 pts. 2 percent. osmic acid 4 pts. Glacial acetic acid E pt. 2-6. No. 1 diluted with 1-5 volumes of water. I per cent. chromic acid 15 pts. 2 per cent. osmic acid 4 pts. I per cent. glacial acetic acid 1 pt. 8-12. No. 7 diluted with 1-5 volumes of water. 13: I per cent. chromic acid 15 pts. 2 per cent. osmic acid 4 pts. 14-18. No. 13 diluted with 1-5 volumes of water. IQ. Y% per cent. chromic acid 15 pts. 2 per cent. osmic acid 4 pts. Glacial acetic acid I pt. 20. No. 19 diluted with 1 volume of water. 202 1. © Det Cent. Chromite acta. 22. . 2 per cent. osmic acid. 23 I per cent. osmic acid. 24-26. 1, 5 and 10 per cent. aqueous solution of bichro- mate of potash. 27. Tellyesniczky’s fluid (Arch. mikr. Anat. 52:247, 1898) : Bichromate of potash 3 gm. Glacial acetic acid 5 cc. Distilled water LOOCG; 28-32. No. 27 diluted with 1-5 volumes of water. 33. Miiller’s solution: Bichromate of potash 2-2% pts. Sulphate of soda i pt. Distilled water 100 _ pts. 34. Zenker’s mixture: Corrosive sublimate 5 per cent. Glacial acetic acid 5 per cent. Miiller’s solution 90 per cent. 35. ‘Saturated aqueous solution of bromine 1 pt. 1The material was immersed for two hours in this solution, and then removed to Flemming’s strong mixture, where it remained eight hours. Bot.—VOoL. II.] HUS—CASSIA TOMENTOSA L. 331 Distilled water 9 pts. 50. ¥y% per cent. iodine 1% per cent. potassium iodide Ppt Distilled water Distilled water 9 pts. I cannot here enter upon an extensive discussion of the effects of these fixing-fluids. It is sufficient to say that material fixed for ten to twelve hours in Flemming’s strong mixture showed as a rule no shrinkage; while material fixed in other solutions (especially in the very dilute ones) or left longer than twelve hours in Flemming’s strong mix- ture, showed considerable shrinkage, with the exception perhaps of that fixed with Tellyesniczky’s fluid, which gave fairly good results. In fixing, two rules were observed: one, always to fix in the field, since only in material fixed in this manner were some of the earlier stages found in any number; the other, to keep the material submerged, for only then could the necessary penetration of the fixing-fluid take place. Frequently air-bubbles adhered to the anthers, especially if the latter were not separated, causing them to float. To prevent this a very simple device was used: a disc was cut off the lower end of the cork of the neckless fixing-bottle and attached to a glass rod passing through the cork; this kept the material submerged, while a notch in the edge of the disc allowed the escape of any air-bubbles. After fixing, the material was washed from six to eight hours in running water; a longer stay in water apparently injured the finer structure of the cytoplasm. At this stage the material was sorted and then dehydrated. It appeared here that a prolonged stay in weak alcohol produced shrinkage. After reaching 95 per cent. alcohol, the material remained for periods of about twelve hours each, in absolute alcohol, a mixture of equal parts of abso- lute alcohol and bergamot oil, bergamot oil, a mixture of equal parts of bergamot oil and 43° C paraffin, 43° C paraffin and 54° C paraffin. It was then imbedded in 71° C 332 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. paraffin, the use of which was found necessary to obtain good ribbons. Sections 3-5 mw. thick were cut and fixed on the slide by the water-albumen method. Flemming’s triple stain was used, with some modifica- tions. The length of time for each stain was ascertained separately. It was finally found advisable not to use the safranin solution usually employed, but to substitute for it Babes’ safranin A, consisting of a mixture of equal parts of a saturated alcoholic solution of safranin and a saturated aqueous solution of safranin, because the former had a tendency, when old, to give a muddy appearance to the cytoplasm. In the case of orange G, even weak solutions took out too much blue. By substituting for this a solution of iodine and potassium iodide of the same strength and composition as the fixing-fluid mentioned under number 36, much better results were obtained. In fact the potassium iodide iodine seemed to fix the blue in the fibers, so that absolute alcohol and clove-oil could be relied upon to wash out the superfluous blue. The staining process was as follows: After dissolving the paraffin in xylol and removing all traces of the latter by a double washing in 95 per cent. alcohol, the slides were placed in the safranin for five minutes and then decolorized in absolute alcohol to which 0.1 per cent. hydrochloric acid had been added. After washing thoroughly in water to remove all traces of the acid, the slides were placed for exactly five minutes in a concentrated aqueous solution of gentian violet, and then washed off in water; after which they were immersed for twenty seconds in the solution of iodine and potassium iodide above referred to. This appeared to fix the blue in an admirable way, so that after dehydrating in absolute alcohol for one second, the prepa- rations, in clove-oil, could be watched under the microscope until the cytoplasm was perfectly clear, the fibers a dark blue, and the granular zone brown-violet. The nucleolus was a bright red as well as the chromatin, except in the earlier stages, when the chromatin stained blue. Bot.—VOL. II.] HUS—CASSIA TOMENTOSA L. 333 DESCRIPTION. The nuclei of the pollen-mother-cells of »Cassza tomentosa, in the stage indicated in figure I, show a nucleolus, which is sometimes vacuolate, and a broken up chromatin thread. The former stains red, the latter blue or red. The cyto- plasm is composed of two parts, one apparently fibrous,’ the other granular. The first is composed of thinner or thicker fibers running in every direction, but more especially from the nuclear wall to the cell-wall, so that the meshes which they form are elongated in a radial direction (fig. I). The meshes are smallest in the neighborhood of the nucleus, and gradually increase in size toward the periphery. The other component of the cytoplasm appears in the form of larger or smaller granules situated between and upon the fibers, and showing a tendency to accumulate in larger or smaller masses, especially at the intersection of the fibers. The fibers can be stained blue with gentian violet, while the granules stain brown-violet. The whole presents a very uniform appearance, except here and there where a thicker fiber or a denser accumulation of granules shows more prominently. Scattered irregularly through the cell may be seen small dark round bodies, apparently oil globules, to judge by their appearance in fresh material. It may be of interest to note that while these bodies were very numerous in some Cases, especially in the early stages, in others they were rare and were then found but seldom in the later stages. Gradually the cytoplasm around the nucleus begins to change in appearance: the change takes place in a single layer of meshes immediately adjacent to the nuclear wall; these meshes become smaller, due perhaps to the interpola- tion:of new fibers, and to the appearance immediately around the nucleus of granular matter, situated upon and between the fibers, often completely obscuring them. The meshes now become elongated parallel to the nuclear wall; 1 The description of the structure of the cytoplasm refers to microscopical appearance only, without indicating whether it isa reticulum or a foam-structure. 334 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. they become very narrow, being usually from two to three times as long as broad. The fibers are covered with gran- ules; they begin to stain a deeper blue than the other fibers. This appearance extends only partially around the nucleus, usually about half way (fig. 2.) Changes now begin to take place in the cytoplasm farther away from the nucleus: the cytoplasm loses its uniform appearance; here and there fibers have begun to thicken, or granules have accumulated upon them. These fibers stain a deep violet and show sharply against the lighter background (fig. 3). They are arranged in more or less conical groups with their bases directed toward the nucleus. Very gradually, almost imperceptibly, the granules around the nuclear wall increase in number, and a second layer of elongated meshes is laid down next the one already formed. Inthe meantime the granules situated upon and between the fibers have disappeared, and the fibers have become thick and smooth (fig. 3). We nowsee the nucleus partly surrounded by a number of rectangular meshes elongated in a direction parallel to the nuclear wall. These processes seem to continue for some time, until finally three or four layers of elongated meshes have been laid down around the nucleus. Then the radial fibers of these meshes begin to disappear, leaving a zone composed of fibers running parallel to the nuclear wall. That this zone is not attached closely to the latter can be easily observed in later stages when one or more spaces exist between the nuclear wall and the fibrous felt-like zone, which stands out clearly as a dark blue band. In the meantime granules have continued to accumulate around the nucleus, and the meshes of the cytoplasm adjoin-" ing the felt-like zone have grown smaller. Together they form a distinct zone (the granular zone) which possesses a distinct outer boundary. The cell now presents the appearance indicated in figure 4. The chromatin appears in the form of lumps, which like the nucleolus, stain a brilliant red. Here and there linin, in the form of violet-staining granular threads, may Bot.—VOL. II.] HUS—CASSITA TOMENTOSA L. 335 be seen connected with the chromosomes. The nucleus is more or less completely surrounded by a zone of smooth parallel fibers which stain a dark blue. Next these lies the granular zone, occupying about half the remaining cell-space, and composed of a reticulum of small, more or less isodiametric meshes, between which (especially at the intersection of the threads) a great number of larger and smaller granules may be seen. Surrounding this zone (which stains brown-violet) is a dark blue line apparently composed of fibers upon which granules, large and small, are situated. These fibers are not continuous, and the number of granules upon them is not the same in all places, so the ring which they form is an irregular one. It stains a deep blue, very effectually marking the boundary between the brown-violet granular zone and the outer gray-yellow cyto- plasm, which is composed of fibers forming large irregular meshes. Upon and between these fibers are larger or smaller masses of granules. The granular zone does not present an absolutely uniform appearance; in places fibers upon which granules have accumulated stand out clearer than others. Usually they run parallel to, or at a small angle with the nuclear wall. At first only one or two are seen, but they rapidly increase innumber. Soonthose nearest the nuclear wall seem to establish connection with some of the fibers of the felt-like zone (fig. 5); gradually other, deeply- staining fibers are added, all running at a greater or lesser angle to the nuclear wall, and finally arranging themselves in conical groups with their bases resting on the fibrous zone. Not only one cone like this may be observed, but several (fig. 6). During this time the number of prominent fibers in the granular zone continues to increase. Fre- quently a space can be observed, at least in places, between the nuclear wall and the felt-like zone. At about this stage the nucleolus begins to disappear; in the preceding one, however, the nucleolus was still present, and the chromatin was in the form of curved, rod-shaped chromosomes, or else in the form of rings, formerly con- sidered so typical of the Liliacez. 336 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. ‘The linin of the nucleus has gradually become more prominent, staining a darker blue; but while in the pre- ceding stage it showed as fine granular threads, it now (fig. 6) appears as smooth fibers, running through the nucleus, and usually more or less parallel to each other and to the axis of the largest cone. The fibers stain blue. The cones, which are composed of granular threads, increase in number. Some are larger; some smaller. ‘ Usually one has the ascendency over the others, and this, as arule, is the first cone formed; it appears ordinarily at one end of the ellipsoidal nucleus. As has been said, it is usually parallel to the axis of this largest cone, and con- sequently more or less parallel to the longer axis of the nucleus, that the linin threads are directed. This seems to indicate a polarity of the cell. In figure 6 we find the cones composed of granular threads; these threads are in connection with the fibers of the felt-like zone. In the next stage we find cones, large and small, composed of smooth fibers. These fibers appar- ently originate from the felt-like zone. But do they push out of their own accord? or are they pulled out by the granular fibers? or are the cones a result of a combination of the granular fibers and those of the felt-like zone? Judging from my preparations the last seems to be the most probable. ; As soon as the fibers of the cone have become smooth, the nuclear wall just below the cone breaks down, and the fibers penetrate into the nuclear cavity, where they anasto- mose with the linin threads. The two kinds of fibers can hardly be distinguished, for while it is true that the fibers originating from the cytoplasm are thicker than those formed from the linin, even this difference very soon dis- appears. Even at this early stage the fibers arrange themselves in bundles (fig. 7), an arrangement which is far less promi- nent in the fully developed multipolar stage, but which reappears in the bipolar spindle. The chromosomes retain their original position for a long Bot.—VOL. II.] HUS—CASSTA TOMENTOSA L. 337 time after the nuclear wall has disappeared. Only when the multipolar spindle has been fully formed, and in fact not until a rearrangement of the poles (with a view to the formation of a bipolar spindle) has begun, do they begin their migration toward the spot where the equatorial plate is to be formed. Throughout the formation of the cones and the subsequent changes from multipolar to bipolar spindle, no indication of a centrosome could be observed. As soon as the fibers composing the cones have become smooth, the nuclear wall breaks down and the blue-staining fibers penetrate the nuclear cavity, where they lie in contact with the bright red chromosomes. The multipolar spindle thus formed has a variable number of cones; the largest number observed in a single section was eight (fig. 10) ; but as the cell was always cut into a number of sections, the number of cones must be greater. The granular zone now presents a uniform appearance, and stains from yellow-brown to brown-violet; the granular threads which were formerly so prominent have disappeared ; nothing remains but small meshes which are fairly uniform in size. Between and upon the threads composing the meshes a large number of granules can be observed. The poles protrude into the granular zone. Surrounding the granular zone we find an irregular line still staining a deep blue. The cytoplasm outside this line retains the same structure as in the last stage. In the formation of the multipolar spindle a highly inter- esting phenomenon was observed. It has been stated before that, as a rule, a cone situated at one end of the nucleus had from the first an ascendency over the others; and while it was not observed that another cone (situated directly opposite this one) was always the second to be formed, yet one could not fail to see that after a little while two cones (one at each end of the nucleus) were more prominent than the others, or were even the only cones present, in which case the nuclear wall disappeared beneath these cones while persisting everywhere else. Such a case is shown in figure 8. ° In nearly every instance this promi- 338 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. nence of two cones could be noted. If my observations are correct, we have in Cassa tomentosa a most interesting transition stage between the multipolar polyarchal and the multipolar diarchal spindle. The largest number of poles of the multipolar spindle observed in cross section was eight; the most frequent number noted in a section was from four to six. In what manner the multipolar spindle finally became a bipolar one could not be determined with certainty; the cones approached each other in two groups (fig. 12), finally forming the bipolar spindle. The cell now presents the appearance indicated in figure 13. The chromosomes lie in the equatorial plate. In polar view, twelve chromosomes can be observed. The spindle has sharp-pointed poles. Apparently there are here three kinds of fibers; some having the appearance of strands are attached to the chromosomes; some run from pole to pole; while from each pole mantle-fibers diverge into the granular zone. ‘The fibers stain a deep blue; the granular zone is yellow-brown; but the irregular line around the granular zone is as blue and sharp as it was before. The poles of the spindle project into the granular zone. The wandering of the chromosomes toward the poles now begins in the usual manner; the continuous fibers still run from pole to pole; while the mantle-fibers become more prominent. At this stage we find, sometimes inside the granular zone, sometimes outside it, small round bodies which stain red, sharply contrasting with other similar bodies, lying on the outskirts of the granular zone, which stain a deep violet, and look like the oil-globules previously observed. Finally the chromosomes reach the poles, and the contin- uous fibers as well as the mantle-fibers assume a wavy appearance. The mantle-fibers during the last stage diverge more than formerly... Why the continuous fibers should assume this wavy appearance is not clear. Certainly the 1 Perhaps it would be well to state here that these mantle-fibers were not equally prominent in all preparations. In fact in some cases no trace of them could be found. Bot.—VOoL. II.] HUS—CASSIA TOMENTOSA EL. 339 suggestion of Williams (1899, p. 194; vide Went, 1887, p. 258, figs. 8-10) that it is caused by the drawing together of the poles, appears very acceptable. In the case of Cassza tomentosa the spindle poles either reach to the outer edge of the granular zone, or else pass through it (figs. 13, 14), yet the daughter nuclei are always formed well within it. The most interesting point at this stage is the behavior of the granular zone, which shows a tendency to accumu- late around the daughter nuclei as they are about to be formed. Gradually it surrounds them on all sides except where the connecting fibers remain. As soon as a wall is formed around the daughter nuclei, they become entirely surrounded by granular matter. It was the last stages of this process which were particu- larly easy to follow in my preparations. To say that the granular matter encroached upon that part of the cell which contained the yet remaining continuous fibers would not be quite true; the fibers seemed rather to thicken at various places in the immediate neighborhood of the daughter nuclei, and this apparently at the cost of the immediately adjacent parts of the fibers. This process continued until nothing was left of that part of the continuous fibers but a granular thread, which sooner or later lost its continuity, so that gradually the place where the threads were, was entirely taken up by granules. This process takes place very gradually, beginning at the sides of the daughter nuclei and proceeding toward the axis of the cell. At first the meshes formed by the transformation of the connecting fibers into granular matter are fairly large, but soon they can no longer be differen- tiated from those of the rest of the granular zone. This change, especially in its later stages, is very slow, and may not be consummated until the daughter nuclei are already forming spindles. The daughter nuclei now lie in a granular mass in no way different apparently from that which surrounded the mother nucleus. The shape of the granular zone is not quite circular, but more or less ellipsoidal, slightly con- stricted about the middle. 340 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. The shape of the daughter nuclei is decidedly ellipsoidal, with the longest axis at right angles to the main axis of the previous spindle. The daughter chromosomes are irregular in shape: sometimes they take the form of lumps; at other times they appear as chains of granules. They stain red and are connected by bands of finely granular matter. A body which seems to be a nucleolus appears in each daughter nucleus. Owing to the presence of a dense granular zone around each daughter nucleus, it was impossible to follow in this case the stages just preceding the formation of the multi- polar spindle; yet what could be seen was sufficient to indicate that these stages agreed in the main with the cor- responding stages of the mother nucleus. The differences observed are to be explained by the shape of the spindle, which in all cases is of a pronounced multipolar diarchal type. The first indication of spindle formation that could be observed was the appearance of a mass of clean-cut, closely-woven fibers surrounding each daughter nucleus. These fibers were particularly distinct where the nuclear wall had shrunken somewhat. They stained a deep blue (fig. 17). The multipolar spindles of the daughter nuclei are formed in a manner similar to that of the mother nucleus. Linin fibers become more and more prominent within the daugh- ter nucleus; they run more or less parallel to the longest axis of the nucleus in which they are formed. Cones begin to appear here and there on the nuclear wall, usually at the ends; they increase in number. In some places the nuclear wall breaks down; the cytoplasmic fibers enter the nuclear cavity, and become merged with the threads of nuclear origin. Finally a multipolar spindle is formed, which, as has been previously stated, is of a pronounced multipolar diarchal type. The narrow, compressed appearance of the entire multipolar figure is here very striking, far more so than in the multipolar spindle of the mother nucleus. In no case was as large a number of poles obsetved as in the Bot.—Vot. II.] HUS—CASSIA TOMENTOSA L. 341 corresponding stage of the mother nucleus. The planes in which the spindles of the daughter nuclei lie are sometimes at right angles, sometimes parallel to each other. Finally the daughter chromosomes reach the poles and become surrounded with a membrane. The chromatin is in the form of granules, arranged in some places to form a thread, and interconnected by bands of the same finely granular substance which we saw connecting the chromo- somes of the daughter nuclei. The nucleolus, which dis- appeared at the time of the dissolution of the membrane of the daughter nuclei, now again makes its appearance. The granular zone surrounds each of the four daughter nuclei very much as in the stages just following the first division of the mother nucleus; but instead of filling up the entire space between the nuclei, it merely surrounds them, | leaving a clear space in the middle through which fibers may be seen running from one nucleus to the other. DISCUSSION. In the formation, development, and subsequent behavior of the spindle there are two points of special interest; the nature of the multipolar spindle, and the origin of the cones. Tun NATURE OF THE MULTIPOLAR SPINDLE. In the first as well as in the second divisions of the pollen-mother-cells of Cassza tomentosa there is more or less approach to the spindle formation described for vege- tative cells. Instead of a multipolar polyarchal spindle Anlage (Strasburger, 1900, p. 121) such as we usually meet with in pollen-mother-cells, we have here a more or less multipolar diarchal spindle. At the very beginning we meet with an indication of this. The nucleus is usually ellipsoidal. The first cone (afterwards the largest cone) appears ordinarily at one end of the ellipsoidal nucleus. The second cone formed lies in many instances directly opposite the first one, or in other cases, the cone which 342 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. occupies this position becomes, as a rule, more prominent than those surrounding it. These two cones remain the most prominent ones. More or less parallel to the axis of these cones, and consequently more or less parallel to the longer axis of the nucleus, run the linin threads. The multipolar diarchal nature of the spindle is even more prominent in the second division. Thanks to the numerous investigations of recent years, we are now able to compare a number of modes of spindle formation. Strasburger (1900, p. 118) distinguishes three types of spindle formation; the multipolar polyarchal, the multipolar diarchal, and, opposed to these, the bipolar diarchal. Multipolar polyarchal spindles are characteristic of the first division of pollen-mother-cells (Lzlium, Passtflora, Lavatera, Cobea, Gladiolus) and of spore- and embryosac- mother-cells (Osterhout, 1897; Strasburger, 1900, p. 121). In Cassia tomentosa we have a multipolar diarchal spindle Anlage in the first division, which thus forms a connecting link between the multipolar polyarchal spindle Anlage of reproductive cells and the multipolar diarchal spindle de- scribed for vegetative cells, i. e., the root-tips of Vzcza (Strasburger, 1900, p. 116), of Adium (Nemec, 18992), and of ferns (Hof, 1898, p. 169). In Wymphea alba (Strasburger, 1900, p. 122) the spindle Anlage of the mother nucleus does not differ essentially from that in the daughter nuclei described by Strasburger for /rzs squalens, where it is multipolar diarchal. It seems therefore that there exists no sharply marked difference between spindle formation in reproductive and in vegetative cells, and that the two processes are closely related. . As suggested by Strasburger (1900, p. 122), the various modes of spindle formation would readily lead one to suppose them influenced by polarity: ‘‘ Eine Pollen- und Sporenmutterzelle kénnte auch in ihren Protoplasten multipolar sein und so die polyarche Anlage der Kern- spindel bedingen. Auch liesse sich denken, das die Bot.—VoL. II.] HUS—CASSIA TOMENTOSA L. 343 diarche Anlage der Kernspindeln in Gewebezellen eine Folge ihrer longitudinalen und radialen Polaritat sei.”’ Polarity is in many cases probably determined by causes outside the cell; for though in the first division of the pollen-mother-cells and in most divisions in vegetative cells, the long axis of the spindle coincides with the long axis of the cytoplasmic mass, yet there are numerous cases (both in reproductive and in vegetative cells) where the long axis of the spindle is placed at right angles to the long axis of the cell. Instances of these are the spindles of the daughter nuclei in pollen-mother-cells, where the nuclei have probably a reciprocal influence. Though the axes of the two spindles are generally parallel and at right angles to the longest axis of the cell, yet it sometimes occurs that the two spindles lie in planes which are at right angles to each other. More striking are those divisions in the cambium cells where the axis of the spindle lies at right angles to the long axis of the cell. It seems probable that we must in many cases look out- side the cell for the cause of the polarity, as is especially indicated by the observations of Nemec (/7ora, 18990, p- 219) on cell-division in potato-tubers, where, in a cell (the surrounding cells having died) the spindle formation did not take place in ‘‘ monaxial’’ fashion, but cones were formed on all sides. THE ORIGIN OF THE CONES OF THE MULTIPOLAR SPINDLE. But few cytologists have investigated closely the origin of the cones of the multipolar spindle. The first to devote his special attention to this subject was Belajeff (1894), in whose paper *‘Zur Kenntniss der Karyokinese bei den Pflanzen’’ were published the results yielded by his obser- vations on the division of the pollen-mother-cells of various species of Zarzx. Since then, in addition to Strasburger’s observations, the spore-mother-cells of Hguzsetum (Oster- hout, 1897), and the pollen-mother-cells of Cob@a (Lawson, 344 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. 1898), Passiflora (Williams, 1898), Lelzum (Grégoire, 1899), Lavatera ( Byxbee, 1900), Gladiolus ( Lawson, 1900), and now Cassia, have been studied more particu- larly in this regard. The results of these studies are a number of observations which agree in the main, though in minor details they frequently differ. It may therefore be of interest to compare the various modes of formation and development of the cones of the multipolar spindle in the spore- or pollen-mother-cells of these plants. . About even the earliest stages the observations frequently differ. In Larzx, Cobe@a, and Cassza the first stages ob- served are described as showing a more or less radial arrangement of the meshes of the cytoplasm (in Codea in the immediate neighborhood of the nucleus only). Similar observations have been made on the pollen-mother-cells of Lilium speciosum (Grégoire, 1899), and by myself on those of Lilium Humboldiz. In Eguisetum and Gladiolus this radial arrangement is not specially mentioned; but in regard to Passifora and Lavatera it is distinctly stated that this radial arrangement of the cytoplasmic meshes does not appear until an elongation of the meshes parallel to the nuclear wall has taken place. In all plants mentioned (with the exception of Codec) this parallel elongation of the cyto- plasmic fibers has been observed and followed with great care, especially in ZLavatera. Here the manner of the formation of the meshes is identical with that observed in Cassia, except perhaps, that in Cassza the fibrous zone in its first stages is not so complete. But sooner or later the number of rows of parallel elongated meshes increases to such an extent that the felt-like mass is easily discernible. At this stage it is figured by a number of authors. Stras- burger (1888) figured and mentioned it; but Belajeff (1894) was the first to describe this felt-like zone: ‘‘ Die erste Verdnderung, welche ich bei in Theilung begriffenen Zellen habe beobachten kénnen, war die Bildung einer den Kern umhiillenden, dichten, filzartigen Schicht, welche auf den ersten Blick als eine concentrisch um den Kern gewun- Bot.—VOoL. II.] HUS—CASSIA TOMENTOSA L. 345 denener Fadenknauel erscheint. Eine nahere Untersuch- ung feinster Schnitte zeigt jedoch, dass diese Filzschicht aus der Kernwandung parallel in die Lange gezogenen Schlingen (Maschen) besteht.”’ This description would fit the felt-like zone occurring in the pollen-mother-cells of all the plants just mentioned; but Belajeff’s words, ‘‘ Die obenbeschrieben Umgruppirung der Plasmafaden ist, wie es scheint, die Folge ihrer Contraction und des Zusammenziehens der Schlingen (Maschen) um den Zellkern,’’ do not agree with the views of others in regard to this matter. Both Byxbee and Williams are of the opinion that the meshes become elongated because of a drawing-out. This then would be caused, not by a con- traction of cytoplasmic threads, but by their expansion. What causes this expansion is not yet explained, but an increase in size of the nucleus is not improbable, and how- ever slight, would produce the appearance described. Observations on Cassza tend to confirm this supposition. But whatever the manner of its origin, a weft-like zone seems to be formed around the nucleus in nearly all cases. The manner in which the fibers of cytoplasmic origin form cones seems to differ in the various plants observed. In this connection it is necessary to call attention to the fibers which may be seen in the cytoplasm at the time of the formation of the fibrous zone. In Cassza these fibers appear when the first changes in the cytoplasm immediately adjoining the nucleus begin to take place. At various points the presence of granular fibers, which stain a deep violet and which are sometimes arranged in conical groups, be- comes apparent. They remind one of the ‘irregular, deeply staining strands”’ of Passzflora ( Williams, 1899, p.191) or of the cones of Eguzsetum (Osterhout, 1897, p. 161, fig. 4). In Cassza these fibers apparently establish a connection with the fibers of the fibrous zone, which latter gradually become parallel, or nearly so, to the thicker granular fibers running at a greater or lesser angle to the nuclear wall. The origin of the cones is apparently determined by the thicker granular fibers. No drawing-out of the cones takes place. (2) April 15, 1904. 346 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. In regard to the granular zone, which is so prominent in Cassia, little remains to be said. This zone has been de- scribed and figured for several plants such as Hemerocallis (Juel,1897), Larix, Equisetum, Lilium, Passiflora, Lavatera, Cobea, Gladiolus, Podophylium and Helleborus (the last two by Mottier, 18972) but always in reproductive cells at the time of division, and not in vegetative cells at the same stage. The partial disappearance of the granules during the formation of the fibrous zone seems to indicate that they are an accumulation of material ready for immediate use. The large quantity present in reproductive cells which divide twice in rapid succession points to the same thing. And though the granular zone is present even in the last stages of the second division, though not always so promi- nently (Lawson, 1898, p. 178), and in fact after the pollen- grains have separated (Byxbee 1900, fig. 25), it must be remembered that a large quantity of food material is needed within a comparatively short time of activity, not only by the pollen-mother-cell but also by the pollen-grain. As Byxbee (1900, p. 72) points out, the manner in which the granular zone accumulates, suggests the gathering of deuto- plasm in animal eggs (Wilson, 1896, p. 115 e¢ seqg.). SUMMARY. 1. The cytoplasm of the young pollen-mother-cell is made up of a network of more or less radially arranged fibers, upon and between which larger and smaller granules are found (fig. 1). 2. The spindle is formed as follows: a. The meshes adjacent to the nuclear wall be- come smaller, and elongated parallel to the nuclear wall (fig. 2). b. A granular zone accumulates around the nucleus. In the cytoplasm, especially in the peripheral part, appear deeply-staining rough fibers, frequently arranged in conical groups (fig. 3). Bot.—VoL. II.] HUS—CASSIA TOMENTOSA L. 347 c. The fibers forming elongated meshes around the nucleus become smooth (fig. 3). d. A felt-like zone is formed partially or com- pletely surrounding the nucleus; granular linin threads appear within the nucleus; the granular zone now takes up about one-half the remaining space (fig. 4.) e. The deeply-staining rough fibers of the cyto- plasm, united into cones, establish connection with those of the felt-like zone (fig. 5). f. The linin threads of the nucleus become more prominent and finally smooth. They run parallel to each other and to the axis of the cone which has the ascendency over the others. g. As soon as the threads of a cone become entirely smooth, the nuclear wall breaks down at the base and the linin and kinoplasmic fibers anastomose. The fibers become grouped into bundles (fig. 7). h. A multipolar spindle is formed, two cones of which, situated opposite each other, are more prominent than the rest. Sometimes but two cones are present (fig. 8). The two promi- nent cones finally absorb the others, thus forming a bipolar spindle. 3. The spindle formation for the second division takes place in the manner described for the first, but is even more pronouncedly multipolar diarchal. 4. The spindles of the daughter nuclei sometimes lie in planes which are sometimes at right angles, sometimes par- allel.to each other. 3. The spindle formation in Cassia tomentosa L. forms a connecting link between the multipolar polyarchal spindle Anlage ordinarily met with in dividing pollen- , spore- , and embryosac-mother-cells, and the multipolar diarchal spindle- Anlage described for vegetative cells. In conclusion I beg to acknowledge my great indebted- 348 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. ness to Dr. W. J. V. Osterhout, at whose suggestion the work was undertaken, and under whose direction it was for the greater part carried out. Berkeley, California, and Amsterdam, Holland, October, Igor. Bor.—Vot. II.] HUS—CASSIA TOMENTOSA L. 349 1894. 1900. 1899. 1808. 1897. 1898. 1900. 18974. 18976. 1899a. 18996. 1897. 1896. 1888. 1900. 1884. 1887. 1896. 1899. BIBLIOGRAPHY. BELAJEFF, W. Zur Kenntniss der Karyokinese bei den Pflanzen. Flora, Bd. LXXIX, p. 430. ByxpeE, E. S. The Development of the Karyokinetic Spindle in the Pollen-Mother-Cells of Lavatera. Proc. Calif. Acad. Sct., 3d Ser. (Bot.), Vol. II, No. 2. GrécorrE, V. Les cinéses polliniques chez les Liliacées. La Cellule, T. XVI, 2° fasc., p. 235- Hor, A. C. Histologische Studien an Vegetationspunkten. Sof. Centralblatt, Bd. LX XVI, p. 65. Just, H. O. Die Kerntheilungen in den Pollenmutterzellen von Hemerocallis fulva, und die bei denselben auftretenden Unregel- massigkeiten. Jahrb. f. wiss. Bot., Bd. XXX, p. 205. Lawson, A. A. Some Observations on the Development of the Karyokinetic Spindle in the Pollen-Mother-Cells of Cobzea scan- dens Cav. Proc. Calif. Acad. Sct., 3d Ser. (Bot.), Vol. I, p. 169. Origin of the Cones of the Multipolar Spindle in Gladiolus. Bot. Gazette, Vol. XXX, p- 145. Mortisr, D. M. Beitrage zur Kenntniss der Kerntheilung in den Pollenmutterzellen einiger Dicotylen und Monocotylen. /ahro. f. wiss. Bot., Bd. XXX, p. 169. Ueber das Verhalten der Kerne bei die Entwickelung des Embryosacks und die Vorgange bei der Befruchtung. /ahrd. f. wiss. Bot., Bd. XXXI. Nemec, B. Ueber die karyokinetischen Kerntheilung in der Wur- zelspitze von Allium Cepa. Jahrb. f. wiss. Bot., Bd. XXXITI, p- 313. Ueber Kern und Zelltheilung bei Solanum tuberosum. Flora, Bd. LXXXVI, p. 214. OsterHouT, W. J. V. Ueber Entstehung der karyokinetischen Spindel bei Equisetum. Jahrb. f. wiss, Bot., Bd. XXX, p. 159. Rosen, F. Beitrage zur Kenntniss der Pflanzenzelle. Cohn’s Beitrage zur Biologie der Pflanzen, Bd. VII, p. 249. STRASBURGER, E. Ueber Kern und Zelltheilung. Jena. Ueber Reduktionstheilung, Spindelbildung, Centrosomen und Cilienbildner im Pflanzenreich. Jena. TANGL. Zur Lehre von der Kontinuitat des Protoplasma. Sitzder. d. k. k. Akad. d. Wiss., Bd. XC. Went, F. A. F. C. Beobachtungen iiber Kern-und Zelltheilung. Ber. d. d. Bot. Geselisch., Bd. 11, Heft 7, p. 247. Wison, E. B._ The cell in development and inheritance. New York. Wiis, C. L. The Origin of the Karyokinetic Spindle in Passi- flora coerulea Linn. Proc. Calif. Acad. Sct., 3d Ser. (Bot.), Vol. I, p. 189. 359 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXX. All figures were drawn with the Abbé camera lucida. Objective, Zeiss Hom. Imm. 73; Comp. ocular, No. 6. Fig. Fig. Fig. Fig. Fig. Fig. I. A young pollen-mother-cell. The cytoplasm consists of elon- gated meshes. Change in cytoplasm. The meshes of the layer immediately adjacent to the nucleus become smaller. Beginning of accu- mulation of granular matter around the nucleus. Formation of violet-staining fibers in conical groups in the cyto- plasm. The fibers parallel to the nuclear wall have become smooth. A dense granular zone has accumulated around the nucleus. Granular fibers have appeared in the granular zone. Formation of cones. More or less parallel linin threads are seen in the nucleus. 352 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXXI. Fig. 7. The nuclear wall breaks down. The fibers are grouped in bundles. Fig. 8. Nucleus with two especially prominent cones. Figs. 9-10. Multipolar spindles; these occur but rarely. Fig. 11. Indication of bipolar spindle. Fig. 12. Perfect bipolar spindle. 7 : - ah a, 2 oo eS ‘i ee, ee 7 - 2 - ; 5 354 Fig. Fig. Fig. Fig. Fig. Fig. Ta: 14. 15. 16. 17- 18. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXXII. Daughter chromosomes proceeding toward the poles. Daughter chromosomes at poles. Several small, red-staining bodies present in granular zone and cytoplasm. The daughter nuclei are surrounded by membranes. Felt-like zone around daughter nuclei. Multipolar diarchal spindle of daughter nuclei. Bipolar spindle of daughter nuclei. INDEX TO VOLUME II. New names in heavy-faced type; synonyms in ttalis. AGAVE..... 264, 265, 267, 271, 272, 273 Calochortus macrocarpus. . . . 116, 148 americana........ + + 260, 274 maweanus.. . 109, 113, 120, 121, 227 Aglaophyllum ruthenicum ...-- 22 A ATOL 40) ion om es 113, 120 PUNT ene eros a en ene ened one, ™ 342 TOSEOUG gel cso, suet aioe 113, 121 Aster ascendens yosemitanus. . .- 293 nity Gude) Oe GLeeL eG: ic 107, 114, 126, 128 GueDLOwdle 100 pavonaceus... ..- - 114, 129, 130 DICEUST ohana 1s 6) emen ene emine han 141 CALOCHORTUS . 107, 108, 110, 112, 116, 129 plummere...----- 109, 114, 132 134, 147, 149 pulchellus . . . 109, 111, 112, 118, 119 albus 107, 109, 111, 112, 117, 118, 119, 131 purdyi....-.---- 113, 126, 127 amabilis.......-- 112, 118, 119 purpurascens. + +++ +++++ 139 AMCEUUS toe tues Gos, « s LL2;) 1175218 POSCUS A Tea rem espe) aneselion x - 139, 141 apiculatus.....---+-- 113, 127 Shastensis....-+«- +. 113, 125 SLES ied cee i crarene Los 136 splendens 115, 116, 143, 144, 145, 146, 148 HET Ali ersy ei os ace 109, 113, 120 atroviolacea.....--+-+-+ 143 CELUICUSH is ees 2) one oie LS major... ....- - 115, 143, 144 Gafalaticelicn. ts 26> 115, 142, 143, 145 montanus . 115, 143, 144, 145, 146 ClavatLUSie ci she oie os 114, 134, 135 TUDUA Pheri Alo iste elt 115, 144 COTEAUSTM ni See eco ie sini eas 123, 124 tolmiei. ....... -- 113, 126, 127 CONCOlOL = hes Sle alae 114, 135 umbellatus ..... 113, 123, 124, 125 Ut IT Sires =) foray os 116, 147 uniflorus.....-.-- 113, 124, 125 elegans... . - 111, 113, 121, 122, 127 venustus.. 108, 109, 110, 115, 132, 137 ABIL EF Ae. sees Le eee 122 139, 140, 141, 142, 147, 148 SOETEEYE S On Od Cn ae Tis yl 22) aN eldorado .... . .115, 139, 141 subclavatus . +. +++ eee 123 purpurascens. . 115, 139, 140, 141 ExGavattsis 9 « .6 6 3 «© 115, 146 FOPUSIED ic ee aioe els (6 tee 139 flexilosusi..... « «- - + - « «116, 147 TOSEMUS: chee at oes 115, 139, 141 greenei....+--+--- 114, 128, 129 sulphureus .. . . . 115, 141, 142 gunnisoni....---+-+:-: 114, 131 vesta ....-.-- 109, 115, 139, 140, 141 howell... 26s ee ss 114, 127, 130 WAIIACENL 2. loica) ot iesel =) = He Chit 24) invenustus..... ~- - 115, 145, 146 RCCELULM ai ioe.a) cae oi istrel fen ot 109, 114, 132 kennedyi.....--.--- 115, 135, 136 AlfauiSiavasdics cient el © toicl aie 133 TeiCHEMEI a temel f- 10 be iiss 116, 149 obispoensis....--- 114, 133 Niaciiniswatciciewetes «0h 3) 2 Ie ae 125 purpurascens... ...- - - 114, 132 Lobbites sf eici ee see tens 342 GREHOCARPUS) ou spells sw spare, eusimeeaD @ECHIALOLIA es ss. te else pee sie 298 PASSIFLORA .. 69, 70, 71, 72, 342, 344,.345 346 Phacelia circinata.....-.+.-+-.-. 291 MeETIOLAIB asi ele.s «<..5 ivi, 8) steele Stimnlans. <2: 6 * 615 6 tyete 291 Phoradendron......-- 100, 101, 305 Phyllospadix . . 15, 27, 174, 176; 179, 180 194, 207, 212, 213, 214, 233 SCOMEIn Mba cis at els. 'el'e! Sate 212 Podophyllum .... ...-.--+ +71, 346 Polygonum douglasii ......--- 286 PS Were AU eee RCS I 286 Porphyra . 173, 174, 175, 177, 178, 180, 182 183, 184, 185, 186, 187, 188, 192, 194 195, 197, 200, 210, 212, 213, 214, 216 219, 220, 221, 224, 228, 229, 230, 231 abyssicola .174, 182, 183, 184, 185, 188 192, 193, 194, 195, 196, 219, 223, 224 ainethysteay. = 3 - f=: 6. se! +) 2) 174 amplissima.. . 196, 205, 215, 216, 221, 222, 224 atropurpurea .- +++ +2 ress 199 COCCINEA.. 2 + + + 2 ® laciniata . . 174, 175, 182, 183, 189, 192 193, 194, 195, 196, 197, 198, 199, 204 208, 230 umbilicalis.. .. . . 178, 193, 199 leucosticta . 174, 175, 180, 184, 190, 191 192, 193, 195, 198, 199, 200, 201, 205 207, 220. 228 APE ye ais satus. Us Sci ealh ena HLA miniata . . 174, 175, 184, 188, 192, 193 195, 197, 218, 219, 220, 221, 222, 224 cuneiformis .....- 194, 196, 218 FeOHUESSEE seo ee eee «0h 220 naiadum. . 174, 175, 176, 177; 180, 181 182, 183, 192, 194, 195, 212, 213, 214 223, 231, 232, 233 major...-.--- « -176, 213; 214 PRN OW es) = nereocystis . . . 175, 177, 178, 183, 187 191, 194, 195, 210, 211, 212, 229, 230 occidentalis ......-- 195, 196, 228 perforata . . 175, 176, 177, 178, 179; 182 183, 191, 194, 195, 197, 200, 202, 203 204, 205, 206, 208, 209, 211, 212, 216 230, 231 lanceolata. .175, 177, 191, 194, 195 208, 210 segregata . . . - 186, 191, 195, 207 INDEX. Porphyra purpurea. ......... tenuissima . . . 174, 175, 183, 184, 192, 193, 195, 196, 219, 220, 221, 224 variegata..177, 183, 186, 187, 194, 196, 225, 226, 227, 228, 231 VUIGAYEIS. » «2+ 174, 196, 202, 204, Prion ers pie, exe) alow ant) sey ievesua eats TATICEOMALAY ol apiiet ta [ul wife Wet Mell ota) ta Pyropia californica... +++ +24 wi id wh jot toy 6! (oh cot We wee moa chityaibim on crs actoieola) HG o6o,c ambiguum.. . amictum... Beyer es ee ascendens 50.) 4) 35) en os 244, JASPELLEY sje! =) = os: sts a AUTEM I oie See a Sen sae Rafat binominatum.... bracteoOSunys weiss. oie a le ae brandevei .- 0). i). 6). «242, COLE TH a),c, h lelvelin bo) rome Ninn aon tails (Othbiormehto bits o Aiguopol cto & 246, GULLOSUHIN Ve 3 cots) Shou ie bal sire) eliceste cognatum....+--+-2+-s+-ee-s Copa besre hab booWr ry wy Ie ein ech itch Our fahhiphateentiherll greg: Goch Sic ch ded - echinatumt.. . + + sees 3 erythrocarpum.........-- fuchsioides.. ++ ++ +> Rie etal glaucescens......-. - +245, glutinosum....-..--+ PEACE Nay pyettta sens pelea) rest isnia HeESPEEUIE heii") 5 [ieee ie =) hittellianum ...... .. 245, hudsonianum.. . hysteis. 2). es yess oles 248,, indecortum ......- 243, 244, TACHSERE S22 Ge lio aiiail toy hues) yainers molle.. . lasianthum.. . pba hiteynsbs?l, Mier og OC moon PEO Bok leptanthum..)...).. 25% = brachyanthum .. 2+ + +e ees malvaceum........ 241, 244, mariposanum ......-. -- SnALSHALIs sproier to sikst elie ne) Joyal menziesii. migratorium....-.-+--+-+-:--; montanuwmt.. ++ + montigenum......--+-+-:- “nevadense . . . 241, 244, 245, 246, Hivenin's ¢ |. 6- % OcciIdeNntales ic eek = feline © oligacanthum.......- 246, oxyacanthoides..... palmeri.... 358 Ribes prostratum..... GUWETCELOLHIN |, ye ious tedl ot =. se TORZII ict smells & sanguineum.... VATLELOLUME . . » = « wo ee SAXOSUM1.«« « « scuphami SETICCUA so) Si Haw yo, 246, viridescens....... SPaciiawuIn Js.) as eked shies! he SPeEClOSHM os 9s sis, oh sts Pe SLAMEREUM oa te ce le, a ies 0 ye Te subvestitum ... tenuiflorum.... VOLUN s,s eee eich ee VADUEHMILOI EE cl iks ny eee WACOCTIS( vim es We Cele tah al se THEASUGD Ga Welty 1a. state io eae c sk WISCOSISSIMNUM. (6 6 a. mes watsoniantim...- fic. «8208 wilsonianum..... RIBeCSIA eld car baer ate arneee er ok ODSGIMIA:. clcsiitis vane tee tel aie eis SEQUOIA SEMPERVIRENS .... Solanum arizonicum...... COMPOWMICUE saa iss oo 160, genistoides.. .. tenuilobatum .. .. . 164, “ owe CALIFORNIA ACADEMY OF SCIENCES. 250 251 251 249 249 251 249 250 247 251 251 251 250 249 251 249 250 251 249 251 250 249 251 87 165 172 171 171 214, 223, 224 [PRoc. 3D SER. Solanum umbelliferum 159, 160, 162, 163, 164, 168, 171, 172 californicum.. .. . .164, 172 wallacel). > Goto 163, 164, 166, 168 WATAGIS i .cP is wets asta 164, 166, 168 xanti. . 160, 162, 163, 164, 166, 167, 168 gilabrescens.. . . . 164, 168, 169 intermedium..... - 164, 168 QUBLIQCEL 0s 1s =)» . 160, 161, 166 Streptanthus gracilis... . 285 TEV AN dh! alone was ts Sokiontey Meg te Mone aan 173 atropurpurea.. .....+<«s-e 202 TRESRIATA Hh 2 Fs Beg eee - 196 miniata.. . BY Ae yr epee 218 PUTpULEA Fl Se ss el ee eee Ae 218 Visera, -. ow the eee eee 342 Viscuiitiors con cues bea - 100, 101, 305 Wildemanta......-+ wis) os ODT Sy LSD: amplissima .. «eee 215 TALTHIAT EN a bie es fe) te Puna be 196 pea a AP arvan s lc! ess ew ee 218 perforata we wer sre neva 202 PemgsSthe ds Meh ea cin hat oe 220 AGE PUA: Wks bie: tien) oe ta is es 225 ZOSTERA. . . . 176, 180, 182, 194, 207, 213 7 nird Ss eries. : BOTANY. VOL. ie : No. Hey Morplclosica Study of Naias and Zannichella. sane Hones cy - Jas Houghton Campbell - DA RTE er koa Cows EE es FEB E OO ae -No. -2—Studies in the Herbarium and the Field. No. Ie BY: Alice ob ie “Bastw0od, cosbse csv ecns obebe tees seen pea tese renee nes els, Speen ae No. _3-Siudies in. the Herbarium. and the Field. —No. 2. ‘By. Alice Lo oe | es PastwOods as ots et eecagkee pe cera ate Mart ewan A HOF en ae No. _-4—Phycological ‘Memoirs. By De “Alton Gquiders. oss acd ek? OTS Ce aa “No. 5—Some' Observations on the Development of the Karyokinetic (ee Beret “are - Spindle in the Pollen-Mother-Cells of Cobzea scandens Cav. ee act caer ts ~ By Anstruther A. TeaWBOMl 6. 3s cic. 5s Shas eniec ene ste toeyes a5 ae OS 6—The Origin of the Karyokinetic Spindle in ‘Passiflora coerulea Be aes ake: “Linn, By Clara L, Williams. 00.0.0 s06ss.-ccrussetesseees 36 eee No. eaEeNe Nature of the Association of Alga and Fungus it in Lichens: toe ‘ By George James Peitcess i se etks copes ee eae en Gee ee No. 8—Californian Hypogzeous Fungi. By H. W. Syidcaces: Se 7! ber a iS - No. g—Studies on the Flower and Embryo of ‘Sparganiam. By Soak ar ae ~~. Douglas Houghton Campbell. . .: Mi REA pararie wes we Cant Weer aN gO No. to—A_ Morphological Study of the Flower and Enibrys- of thee v5 oe Brace © Wild Oat, Avena fatua L. By William Austin Cannon...... +50 en er : ene 4 Saeco Be a oe ~~“ VOL. Il. eae pee ; : & No, 1_Nitophylla of California. Description and Distribution. By “2 Charles Palmer Nott.. 00 ..0cc.0s.0 feeee cecepe teen rence 7S: “No: 2—The- Development of the. Karyokinetic Spindle in the Pollen- Bite erat? ¥ %; - Mother-Cells of Lavatera. By Edith Sumner Byxbee . op eee “No. 3—Studies « on the. Coast Redwood, Sequoia pire ek aes Hal ay "By George James Peirce... 22. ..eeee cecens ce teenee etee es BS inaoe No. 4—A Revision of the Genus Calochortus.. By Carl Purdy. i fie a5OUe ee - No. 5—A Group of ‘Western American Solanums. By S. Be Parish: 2.254 sy _ Now 6—An ‘Account of the Species of Porphyra, Found on. the Pacific + Coast of North America, - By Henri T. LR 2 CARRE R RGRS PRA (0 at Ps es No. 7—Some new Species ‘of Pacific Coast Ribes. By Alice Eastwood 25> » No. 8—Cell Studies: I. Spindle’ Formation in ‘Agave. By W. J. A! Be te | es SOsterhQut. chs opace 5 Fee crore Coren Snes teat eee ae BOseS 5 No. g—New Species. eon the Sierra Nevada Mountains of California. aerate By Alice Eastwood. ....-.5ce0 tenes entenerstetreces rent YT ga No. to—The- Root-tubercles of Bur Ciitier (Medicago dendeutaia Willd.) and of some other Leguminous Plants. Sa: See ee George James Dire ee i ca EE De RE Bo irae a ae Ege Wie No. ards Formation in the Pollen-Mother-Cells of Cassia laos So i : dey cie aaed Ll. ree Henri Ty A. Bus: a eras Porbstp “ine ey othe. he hg ATV ae iy ii oi ‘ yh ih ' ——= PN eines HL AT a } | Il | 0288 39 3 5185 0 | |