¥ Pe ae Sian pelsercnee Ss crt eae y aes oa - eae ra? ova An Pano pusaetnteryutiar Auaiare meet : = toate ae : r ~ pagent putiatnvtn afew ee Pao ie tay aint ines Ta aesaniaeaes ‘ ‘ Ret ; ane Me na er pense. ke bates “e CMerrenedanyt eee ° ar je Po nr Sa Feet oe pen ween COM GS Worn ew no De GOP ee cage Pet Mee gO re realete owe anaes eres Pe St up ANS: SED SON: Ae Sty Tega San Rae 3 ¢ : ie zi =~ = (3 ole at ay - \ ee > ro ay <* 2. . ‘> ile \ Y Wat? ol PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES THIRD SERIES A Go OP ANT oy WV OE ot 1897-1900 SAN FRANCISCO PUBLISHED BY THE ACADEMY JUN 1419 v4) ly Wy ey yy) CONTENTS OF VOLUME I. Piates I-LIII. GREECE RSI OU LINE SECU N a aha, Baia Gaidl qievare aS ars «io «sso arnaph oboe ONCE ES peers FAM eye etevatene eed eee iar etter ete te a No. 1. A Morphological Study of Naias and Zannichellia. By Douglas Houghton Campbell. (Plates I-V) (Published June 5, 1897.) rd No. 2. Studies in the Herbarium and the Field. I. By Alice East- WOOet Gates? WLW EL) ami he es ha ia's. cle syac.o stoi sw aw enwinnd (Published November 27, 1897.) No. 3. Studies in the Herbarium and the Field. II. By Alice East- MUOOG Ey (erates: VNR): en ee i on) (Published September 24, 1898.) No. 4. Phycological Memoirs. By De Alton Saunders. (Plates XII- PODER DY co ra ate tayeh oocletanctay My aca) troy naka w wvgheiahs teenie ave oe's. ore aMetbdatnrae (Published October 31, 1898.) No. 5. Some Observations on the Development of the Karyokinetic Spindle in the Pollen-Mother-Cells of Cobzea scandens Cav. By Anstruther A. Lawson. (Plates XXXIII- GD ILD Baad ag SA Sar Sone ape FS 8 WORN AS NR SDS (Published November 17, 1898.) No. 6. The Origin of the Karyokinetic Spindle in Passiflora ccerulea Linn. By Clara L. Williams. (Plates XXXVII-XL).... ‘ (Published April 15, 1899.) No. 7. The Nature of the Association of Alga and Fungus in Lichens. By George James Peirce. (Plate XLI)....... (Published June 5, 1899.) No. 8, Californian Hypogzeous Fungi. By H. W. Harkness. Plates PAA UNE CN) YA ta cea cate c ctrate tae heise Nae rhinos wie ~etalle RAs ei Selah (Published July 8, 1899.) No.g. Studies on the Flower and Embryo of Sparganium. By Douglas Houghton Campbell. (Plates XLVI-XLVIII). (Published July 21, 1899.) No. to. A Morphological Study of the Flower and Embryo of the Wild Oat, Avena fatua L. By William Austin Cannon. Wt, eRe eine ire ahs ie ta, aitienon i gana a’e' aces adure ave (Published April 21, 1900.) 71 89 147 169 207 241 293 365 March 18, 1904. A MORPHOLOGICAL STUDY OF NAIAS AND ZANNICHELLIA. BY DOUGLAS HOUGHTON CAMPBELL, Professor of Botany, Leland Stanford Junior University. CONTENTS. PAGE. PLATEs I-V. ; A.—NAIAS. Di hie (GROWING-POINT OF THE, STEM), Wo.00.2 ole'.4 clic esse alee cide bce ales 8 EUR Ue LITLE MIO HIAVIES CERN ety par tis cts) 2 Oc MOMN UW LIE e Siv icles Sieve vale helabeten te eislolon ey 10 ALO Een EA SoS OO OS unter need rn) en )Sn ya Reem ae eet RAL Ta At VS I eS II VRE MERE eH EO ONUIOR SIU ce isa Nevin gH ny ats invite: Sete oa AU Esl aca tale a oat sete cen > 12 MERE IVUATE RS ETIEO WIEN ays cts evolsdaccostetaloh a stelers c/oteye's:o, Sletete ators mvchete ete 12 i), Development af the Polen © 06s occ iewy eae ood es 14 (0) The Germination of the Pollen... «io vin sacs es ea 17 Se EB A LNA TSE OR IGOMTER «5 /sic se elvgtie! aleie eo cisie cl sisiehttaeboer sae 17 Die Mile MOTs PMNS RSI sR hl a's. yyw fap rsye al Sr w\ jalan alla ata,s. o:cha 'n aie, tt oer allah ote 23 NTE aie SOR TATTLE ZACTREGUINNS ee, craredel OY. Ue < s=: yy siel cattet avelapalare duvet oterdiw ia Sites eave, ale See stave stoners 24 yy ME MER ED MANTDRU MONS gee ea pas aie, |g dictate, wrliseie a aiarane aie aide aigive acne RENE a ate Nat 25 VA Ra ney Leas Rs COMMS TIED AD Rast cs". 2.9 valde at aiave de vale ae suv 6 Cale matmielot Revels 30 IDX Tap es LRET Ee SiG Mae elon fey teeta Gia. g ates cralteenetelo taste situs ieeiiaba’s aialincilais latemee nem ages 3I NACE Fe a O Ohare ce hee Ten tesustet a stale sea, tative aharelea bial a a\aiie: e elapassisiates ciate, es a1 NET EE MEIN OSPPORINN sex a cucls sooo) uceisteat tie aie ince iret. ejaiie\ a oie: ajo Sale eyeretatelerenate 34 ; B.—ZANNICHELLIA. IU GENERATAVIORPHOROG Veta sae Ute laeeiee isos Wachee sa celdectaen tents 37 Deets OREM = AER Me ubvay steve tlic, rclu ial eranesi/eve/sl erevace.c 6 ess. ace, a veh) cles teahiehare 38 DEK SV EHEG RROCMS earner errcrese, 8 taane: sre a sleds all ston steyeyeyerei aya oo is vahave avec auetarene ere 4o Nera pera VPA EDT OUTED Rey tavern cue ass teres releuibedvanaretele: ole tals ie,'s s: 01 sptelepelsh shetsbes ote 4o INVADED: aE) RVWAID Fe WED COMO RUS ch Meters cpu erere seater ay tiara! alee, 9. as ualalerarctele oushaele 42 Veeco ECOG MER Viera shoei ene nar Owl DURE rk ao vic 3, a oat a atatele abner alaet® 48 NIRS DE tC OMV ER DONO Alpinnt selena rv asic dibaerats! gla elay auch eveualie/e ekavehey see! araslere 50 NAR INEN EE STENT Weisner e tic ls Cu atgla Wacds watniialc nates dated e Gisaststey sieletecals 50 LEXRSE DED RIN OO EAU enn sae eaten NN Lisa atau Al cle diye us ails ieju) kvtojar o: grace male ‘eleva ieiniaval 50 GE EON DOSER: sever Acree ines el bace in RM a faiesd olla ayers viekcrelcaitelaleole sie eceim 51 PRAT meROIPET Ta ea VR TOSS a ere es or Shel eal te Logie, Glave setback a (Spal vis’ ate ayace 6 st'eiernitals wan era 52 SUMMARY AND CONCLUSIONS Nea UNIAN Sane ReO MERSIN etn ae Thora vara reo Tencieistc siiat raters ateieie ie. «(a stelsieiaieieasouasiye 52 EP ANN CELE ICID ANS ayer te a laeas ice secteu ae Mlondl des iehcleush ateidiaye’lelcieiaiie:'e Sieh sichalete apaverefelels 53 SUPP MENTE AR WON OME. poset coset ernein (Neha Giijecals\ baila ala w/ep = c.cl anlar g ciel euonsterevete 59 FEE LOG RAPER Gas oreee fais fettern ao rclcea niet shovel Snlaayaictelieceve aleve ale: o/sieseh wrosay ratanores oral 60 EPP IUAIN ALON OM EAMES Meta is vaiaaigierciavey js ale tieicho, sizes. g sieie' «vcore ioral srevapalapaileie 62 [1] June 4, 1897. 2 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. IF ONE examines the enormous mass of literature bearing upon the great group of Angiosperms that has accumulated and that is being rapidly augmented, one must be impressed with the very great preponderance of work of an exclusively descriptive character. Whether one examines descriptive floras or monographs of special groups, the number where the descriptions extend beyond a mere enumeration of ex- ternal characters is astonishingly small. While this is usually quite sufficient for the mere identification of a plant and for determining its relation to nearly allied forms, it is quite inadequate for settling questions of relationship between more remote groups, and especially those of obscure affinities. The constant addition of new forms to the already immense list of known species but adds to the confusion which exists, and out of which it seems hopeless to expect ever to bring order. What is needed is not so much additions to this great mass of undigested material, as some beginning of a more thorough study of the materials athand. How greatly would the value of many an important monograph on a genus or family be enhanced did it but include a connected account of the whole life-history of one or two representative forms! Were this done, we should soon have a collection of data upon which to make a beginning, at least, of a classification of the Angiosperms, which would be something more than mere guess work. Such minute histological study of all the phases of development is of especial importance in those simpler groups of both Monocotyledons and Dicotyledons, where the floral parts are very simple, and where doubt arises whether we have to do with structures primitively simple or the result of degradation. The present chaotic condition of taxonomy is beyond question largely due to the superfi- © cial methods which are adopted in classification. An in- heritance from a former scientific era, these primitive methods are held to with a persistency which does not augur especially well for the future. The great advances made of late years in microscopical technique, both in regard to preserving and staining methods, and especially the use of the microtome, have been as yet Bot.—VoL. L.] CAMPBELL—NAIAS AND ZANNICHELLIA. 3 but little utilized by the systematic botanists. While these cannot be expected to be the principal tools of the sys- tematist, they should prove invaluable aids in his work. The improvement in technical methods also makes neces- sary a revision of much of the older histological work, which is often either inaccurate in detail or incomplete. Many of these older contributions, of course, are of the greatest importance, but all the more is it necessary to cor- rect any inaccuracies which they may contain. Naturally, with the improvement in technique there has been an in- creasing number of accurate histological studies of the flow- ering plants, especially investigations bearing upon the development of the reproductive parts. It is these which must serve as the starting point for the accumulation of data for the final classification of the higher plants. It is true that the structure of the flower and fruit of the Angiosperms is of the greatest importance in their classification; but these alone are not sufficient for settling positively questions of affinity, except between nearly related groups. An accurate knowledge of the development and histology of the repro- ductive parts and embryo is also very important in this connection. Not until very much more is known than at present about the life-history of representatives of all the principal types of flowering plants, shall we be in a position to begin to build up a system of classification which we can hope to be even approximately accurate. Among the interesting problems awaiting solution is the character of certain simple Monocotyledons. Indeed, the whole question of the relation of this group to the other flowering plants is one about which there is much disagree- ment. With the hope of perhaps being able to throw some light on this question, and at the same time to call attention to these problems, and perhaps interest other botanists in the same or similar work, the writer decided to begin a series of investigations upon the structure and development of some of the simpler Monocotyledons. The present paper is the first of a series which he hopes to publish from time to time, as the materials become available, While no very 4 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. startling results have been brought out as yet, nevertheless, a number of interesting and more or less important struc- tural details have been brought to light, which it is hoped will serve for the basis of more extended researches in the future. How poorly the affinities of the simpler Monocotyledons are understood is at once evident from the extraordinary divergence of opinion among botanists in their arrangement of genera and families. Thus, /Vazas is sometimes included with Potamogeton and Zannichellia in the family of Naia- dacez(Morong, 1893); while on the other hand, most botan- ists at present consider this family to contain only the single genus JVazas, while Zannichellia is placed in the Pota- mogetonacez (Ascherson, 1889); and it has been recently suggested that it should stand as a type of a separate family, Zannichelliaceee (Schumann, 1892). These instances will serve to show how urgent is the need of a thorough investi- gation of these doubtful forms, and as the two genera, /Vazas and Zannichellia, are structurally among the very simplest of the Monocotyledons, they were selected as the first forms for investigation. Both genera, especially /Vazas, have been examined carefully as to their gross morphology, but the histological details concerning them are very scanty. The most important work done of late years is that of Magnus and Schumann. The former confined his work mainly to Navas, although in his last paper (1894) he also gives some details for Zannichelia; Schumann (1892) has given a fairly full account of the general development of Zanni- chellia; but owing to his depending entirely upon a study of the external structure and not using sections at all, he has made some serious mistakes in his interpretations of the homologies of the floral structures. Most of the work of the earlier botanists was unfortunately inaccessible to the writer, but in Magnus’ first paper (1870) there is a very full and clear résumé of the results of these observations, to which the reader is referred for further details. The writer has confined himself mainly to a study of the Bot.—Vo.. I.] CAMPBELL—NAIAS AND ZANNICHELLIA. 5 development and histology of the reproductive parts of the plants, as the general structure and histology of the vegeta- tive parts has already been, on the whole, accurately de- scribed; nevertheless, it has been found necessary to correct a few points, the most important being the structure of the vascular bundle of /Vazas. The absence of tracheary tissue, which has been supposed to be characteristic of this genus', it is found is only apparent, at least in JV. flexz/is, the only species studied. Here tracheids are always found in the young bundles, but are subsequently destroyed by the great elongation of the parts, so that in the fully developed bundle no trace of them is to be found. This will serve as one more instance of the importance of studying the develop- ment of the parts as well as their structure when fully formed. Most of the results here given were obtained from the study of serial microtome-sections. The material was in most cases fixed with one per cent. aqueous solution of chromic acid, and after thoroughly washing was transferred gradually to alcohol, where it remained until wanted. No other reagent employed gave as good results as chromic acid, although material so fixed, unless very thoroughly washed, is apt to offer resistance to nuclear stains. The _ material was stained zz toto with Czokor’s alum-cochineal, and after dehydrating was imbedded in paraffin. The earlier preparations were passed through turpentine before imbed- ding, but later xylol was substituted with excellent results, the formula being that given by Zimmermann (1893, p. 33), except that a much shorter time than he gives was found to be sufficient. The series of sections was after- wards stained on the slide with a solution of Bismarck brown in 70 per cent. alcohol. In the study of the embryo-sac of Varas, the anilin-safranin method recommended by Schaffner (1896, p. 123) was used to some extent and with good results. lWhile Magnus denied the presence of tracheary tissue, he refers to the work of an Italian, Pollini, who claims to have seen it. 6 CALIFORNIA ACADEMY OF SCIENCES, [PRroc. 3D SER. A.—Naias. The most important contribution to our knowledge of the genus /Vazas we owe to the admirable monograph of the genus by Magnus (1870). In the introduction he has given a very clear and satisfactory résumé of the work done by earlier observers, from Vaillant’s account in 1719, up to the time of the publication of his own memoir. He has also written the account of the Naiadacee in Engler and Prantl’s ‘« Die natiirlichen Pflanzenfamilien’’ (1889), and still more recently a paper in the ‘‘ Berichte der deutschen botanischen Gesellschaft’’ (1894). He clearly made out the relation of the floral parts and the anatomy and histology of the mature plant and fruit, as well as the general facts of germination; but he gives no account of the histology of the growing parts of the flowers. He also gives no account of the em- bryo, beyond a reference to the gross anatomy of the embryo in the ripe seed. Schumann (1892) has also investigated the structure of iVatas, but, like his predecessors, pays no attention to the minute structure of the floral parts. His results agree in the main with those of Magnus, but he does not agree with the latter in considering the envelope surrounding the ovule as a peculiar structure, but, in common with other botanists, regards it as a carpel, a view which Magnus vigorously opposes. My own observations were based entirely upon /Vazas flex- ils Willd., the only available species, and it will be neces- sary to examine critically other species before we can generalize as to results. It is true that all those so far examined agree closely in most of their details; but as in LV. flewilis there is a considerable degree of variation in cer- tain structures, especially in the embryo-sac, it is highly important that these should be compared with the correspond- ing ones in the other species. The material used was collected in the Detroit river, but the species is of wide distribution in the United States and occurs in certain parts of California. Bot.—Vo.u. I] CAMPBELLI—NAIAS AND ZANNICHELLIA., 7 The genus /Vazas is of world-wide distribution, but owing to the great simplicity of the flowers and the variation in the habit of the same species, the limits of the species are very uncertain. There are probably about ten species known (Watson, 1880, p. 191; Morong, 1893, p. 60), of which four occur in the United States, two of these being fcund ie Galnorma, (Watson, I. c.); i. e., WV. flexilis and JV. major (lV. marina L.). They are all entirely submersed aquatics and remarkable for the extraordinary simplicity of the flowers, which consist respectively of a single carpel or stamen; and these are remarkably simple in structure and curiously similar both in their origin and early stages of development. It is generally con- sidered that this extreme simplicity of structure is the result of the aquatic environment, but this is by no means certain, as the flowers cannot readily be considered as derived from any more perfect type, and the genus stands very much isolated. Magnus (1870) has given a very complete and accurate account of the relation of the stem and leaves, and the posi- tion of the flowers. The slender stem has the leaves apparently in whorls of three, but a careful examination shows that in reality one of these three leaves is the basal leaf of a branch springing from the axil of one of the leaves. Except for this basal leaf on each branch, the leaves are in pairs, the lower of each pair always developing a branch from its axil, while the upper one is invariably sterile. A further examination shows that in place of the absent leaf at the base of the branch there is a flower, which may be either male or female in JV. flexz/is, but in JV. major, which is dicecious, is always of the same kind on one plant. These flowers, as Magnus (1. c.) showed conclusively and as we shall see later, are morphologically the equivalents of axes, and the ovule and anther strictly terminal organs. Owing to this formation of a branch in the axil of one of each pair of leaves, the whole plant is bushy in form. Perhaps the most remarkable thing in regard to the flowers is their striking simi- larity, both in origin and structure, the male and female (2) June 1, 1897. 8 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. in the earlier stages being almost indistinguishable. The stamen is invested with envelopes which seem to corre- spond to the outer integuments of the ovule and possibly the carpel. A further discussion of these points, however, must be left for the present. I. Tue GRowInG-POINT oF THE STEM. The relation of the leaves, stems, and flowers has been carefully worked out by Magnus and confirmed by Schu- mann. The writer has nothing to add to their conclusions; but in order to follow out the histological details, a brief review of the facts may be useful. If the terminal bud is carefully dissected out, it will be found that the shoot terminates in a conical tip, which pro- jects considerably above the lateral organs (fig. 1). The leaves arise in pairs, mostly on opposite sides of the stem (fig. 5). The lower one is larger, and its base extends around the stem in the form of a sheath; almost as soon as it is visible, there is formed, just above it, a protuberance (fig. 5, &), which a careful examination shows to be con- siderably extended laterally. Very early this outgrowth shows a cleft in the middle, by which it is divided into two nearly equal parts; of these, one becomes the begin- ning of the branch found in the axil of the older leaf, and the other the rudiment of a flower. At first it is impossible to distinguish between them, but soon they undergo further changes which render their recognition easy. If a section is made, cutting through the centre of the apex, it presents the appearance shown in fig. 5. The pri- mary tissue-systems are clearly recognizable and show the arrangement typical of most angiospermous plants. There is, as usual, a well defined epidermis, all of whose cells are for some time capable of division, but the divisions are always radial, and this epidermis continues without break over the apex, and over the surface of the young appendicular or- gans. Below the epidermis is the periblem, also composed for the most part of a single layer of cells in the younger Bot.—VoL. I] CAMPBELL—NAIAS AND ZANNICHELLIA. 9 parts, but of course later undergoing periclinal divisions. The plerome is well marked, and in longitudinal section shows usually two rows of cells. A single large cell (fig. 5, *), which is sometimes conspicuous, may perhaps be the single initial cell for the plerome, but its position is such as to make it also possible that it belongs to the periblem, and that there are several initial cells for the plerome. In regard to the form of the stem-apex, VV. flexilis re- sembles more nearly JV. graminea, to judge from Schu- mann’s figure (fig. 1), than it does V. major (Magnus, 1870, Pl. IV, figs. 6-11), where the apex is shorter and thicker than in any of the other species. The apex of the stem is seldom perfectly straight, and the first indication of the formation of the lateral appendages is the appearance of two slightly projecting ridges, one immediately above the other and placed upon the convex side of the apical cone (figs. 3, 5, 4, 7). The lower one of these is the leaf and the upper the structure (primordium) which subsequently gives rise to a lateral branch and a flower. A little later there is found on the opposite side of the apical cone and a little higher up than the other leaf, another projecting ridge, which does not, however, show any structures above it. | This is the second leaf of the pair (fig. 5, 7). The tissues of the appendages show the same arrangement as those of the stem-apex and are continuous with them. The tissues of the mature stem are very simple, but their origin is readily traceable to the primary tissues of the apex. As in all aquatics, large air-channels are developed, arising here between the two outer layers of the cortical cells. In the specimen of JV. flexclis figured (fig. 6), there were six of these intercellular spaces (7), and the general structure is much the same as in JV. minor (Magnus, 1870, Pl. VII, fig. 4). lV. major (Magnus, 1870, Pl. VII, fig.2) has a more massive stem, a larger number of air-spaces, and a thicker cortex. There is in all cases a single axial, vas- cular bundle, with a well defined endodermis, showing, when mature, the characteristic radial foldings of the walls, and derived as usual from the innermost layer of the periblem. 10 CALIFORNIA ACADEMY OF SCIENCES. [PRocC. 3D SER. The intercellular spaces are separated by a single layer of cells, and the general appearance of the section is much like that of such aquatic Pteridophytes as Marsilia or Azolla. Like these, the bundle is typically concentric. Magnus(1889, p- 215) states, and his statement has not been contradicted, that there is no tracheary tissue present. This is incorrect, at least for JV. flexzlis. In longitudinal sections of the younger parts of the stem, spiral or annular tracheids are always found, forming a single line in the axis of the bundle. The thickenings on the walls are formed at an early period, and with the rapid growth of the stem in the lower internodes, these primary tracheids are pulled apart and destroyed, leaving the central canal, which Magnus fig- ures. Itmay be noted that Caspary(1858,p.515) noted traces of these trachez in this same species, at the base of the funiculus, but apparently overlooked them inthe other parts of the plant. It is evident from his account, however, that his investigations were not very extensive, and it is by no means unlikely that further investigation, using microtome- sections of the younger parts of the stem, will reveal the presence of tracheids with normal thickenings, in the other species as well as in JV. flexzl’s. In common with other submersed aquatics, /Vazas has the epidermal cells contain- ing chlorophyll, and not noticeably different in form or contents from the outer cells of the cortex. i Par LEAves: The structure and arrangement of the leaves is very sim- ilar in all species of /Vazas yet investigated. The base of the leaf is expanded and forms a sheath surrounding the stem more or less completely. Inthe case of the sterile leaves, i. e., those which develop no axillary products, the base of the leaf envelops the stem completely and one edge overlaps the other, much as is found in the case of the coty- ledon (fig. 80). The marginal cells of the leaf are extended into the brown teeth, which gives the leaves a distinctly serrate outline. Cross-sections of the leaf show a structure Bot.—Vot. I.] CAMPBELI—NAIAS AND ZANNICHELLIA. 1 similar to, but simpler than that of the stem. In WV. flexilis, except in the middle, the leaf is composed of but two layers of cells. The center is traversed by a vascular bundle, much like that of the stem, but without a definite endoder- mis. Inthe younger stages, a single line of tracheids can be seen penetrating into the base of the leaf, but no trace of these is visible in sections of the fully developed leaf. The bundle is surrounded by a layer of large cells, which abut immediately upon the small intercellular space on either side of the bundle. Vazas major (Magnus, 1870, p. 49) differs markedly from all the other species in having a wel} marked epidermis consisting of small cells. This is correlated with the thicker cortex of the stem!. Between the base of each leaf and the stem are two scale-like structures, the ‘* squamule intravaginales,’”’ structures found very generally throughout the Naiadacez and Potamogetonacee. ER re oo as The primary root of the Vazas flexilis is remarkable for the absence of a root-cap, a fact which has apparently been hitherto overlooked. - Unfortunately my material of the mature plants did not include the roots, and I am unable to say whether this condition is also found in the later roots, nor have I been able to find any reference to the subject elsewhere. As to the position of the roots in the fully developed plant, the writer cannot speak from his own observations, and the account given by Magnus (1870, p. 16) is not entirely clear. To judge from his fig- ures, however, it would appear that the roots are pro- duced in regular succession, one to each of the lower nodes of the stem, arising between the two leaves of the node on the side toward the leaf at the base of the branch. lFor further details of the structure of the leaves, the reader is referred to Magnus’ monograph. I2 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. IV. THe FLowers. The flowers of /Vazas represent the simplest form which an angiospermous flower can assume, consisting simply of a single carpel with but one ovule, or of a single stamen, which in most species has but one pollen cavity. In origin and in their earlier phases of development, ovule and stamen present very marked similarity, and this is heightened by the development about the stamen of structures correspond- ing to the integument of the ovule, and of an envelope which imitates very closely the carpellary covering of the ovule; this even extending to the formation of similar teeth at the apex. The resemblance of both to the sporangia of certain Pteridophytes, especially Azo//a, where the macrosporan- gium has a similar investment, is also noteworthy. As we have seen, the rudiment of the flower is formed by the equal division of the primordium, which is formed above the lower of each pair of leaves. At first these are not distinguishable, but very soon one of them becomes somewhat larger than the other (fig. 11, f), and this, which is the young flower, soon develops about it a ring-shaped wall—the carpel or its equivalent—and thus is readily dis- tinguishable from the other protuberance, which is also more pointed and becomes the apex of the lateral shoot. It is thus plain that the origin of the ovule or anther, as the case may be, 1s strictly terminal; i. e., that the flower and the lateral branch are due to the dichotomy of a common primordium and therefore are of equal morphological value. The male and female flowers are scarcely distinguishable at first, as the envelope about each is entirely similar; but as a rule the young ovule is somewhat more slender than the anther. The question of the homologies of the envelopes of the flowers will be better understood after considering the development of the latter. 1. THe Mare Fiower.—Shortly after the separation of the primordium into the rudiments of the lateral branch and flower, the latter grows much more rapidly, and its real nature becomes evident. It begins to broaden at the base Bor.—Vou. L] CAMPBELL—NAIAS AND ZANNICHELLIA. 13 (fig. 8), and this enlargement quickly assumes the form of a ridge running round it and growing up about the apex in the form of a cup-shaped envelope. A longitudinal section of the young male flower at this stage shows a structure very similar, as might be expected, to that of a vegetative shoot. t= is Se & Se. he, a — = = ia al SS ioe 5 MPBELL.| Phare | f t 17 “cae CTT> OP a eg: ea TW Se aur ais YS NB bawhnlon Ay 3 i PLATE II. All figures refer to Natas flexilis. Fig. 27. Mature male flower, showing the long pedicel; X 25. Fig. 28. A male flower which has discharged the pollen, showing the lateral rupture of the floral envelope; X 25. Fig. 29. Longitudinal section of a nearly mature flower, showing a com- plete division of the anther into two loculi; X 4o. Figs. 30, 31. Germinating pollen-spores; X 400. Figs. 32, 33. Pollen-tubes, showing the granular protoplasm at the ex- tremity. In fig. 33, the two generative nuclei, 2, can be seen. Figs. 34-39. Successive stages in the development of the ovule, seen in longitudinal section; < about 4oo, (except fig. 37, which is magnified about 200). The carpel is not shown except in figs. 34 and'37. The archesporial cells are shaded; /, the tapetum, 77'., zz?., inner and outer integuments. Fig. 40. Young female flower, showing the young spiny lobes, x, and stigmatic lobes, v,; one of the latter is still very small; X about 200. Fig. 41. Section of young ovule, with the archesporium divided by longi- tudinal as well as transverse walls. Figs. 42, 43. Young embryo-sacs, with two and four nuclei; X 450. Fig. 44. A mature female flower; X 25. Fig. 45. One of the stigmatic lobes; X 200 Fig. 46. A spiny lobe; X 200. Fig 47. Longitudinal section of the ovary of a mature flower, showing the stylar canal and papille, 4; x about 60. Fig. 48. Young papillze from the upper part of the ovary; X 400. Fig. 49. Older papillz from the same place. Fig. 50. Upper part of mature embryo-sac, showing one of the synergide, s; the egg, 0; and the two polar nuclei in close contact, £; X 400. Fig. 51. The two synergide from a similar one. Fig. 52. Egg apparatus from a slightly younger embryo-sac than that shown in fig. 50. Fig. 53. (a) Upper part of the embryo-sac at the time of fertilization; pf. the pollen-tube, with two generative nuclei; 0, the egg. (6) The large endosperm nucleus, probably formed by the fusion of the two polar nuclei. LITHERITTUNE REV EF [ CAMPBELL Pure IL ins LAA sa \p q pay “e PLATE III. All figures refer to Matas flexilis. Fig. 54. Two sections of an abnormal embryo-sac, in which apparently the upper archesporial cells were not absorbed as is usually the case, and no definite egg apparatus was formed. Fig. 55. Lower part of the embryo-sac shown in fig. 54, with six antipodal cells (four only are shown in the section). Fig. 56. Section of an ovule which apparently had not been fertilized; the embryo-sac is replaced by a mass of elongated cells; X 200. Fig. 57. (a@and 6) Two sections through a normal embryo-sac, showing fertilization; 7, pollen-tube; 0, egg; s, synergidz; 4, endosperm nucleus. (c) Lower part of the same embryo-sac, showing two of the antipodals; x 450 (about). Fig. 58. An abnormal embryo-sac, in which only one cell of the egg apparatus could be certainly distinguished. The end of the pollen-tube was very much enlarged. (6) Antipodal region of the same embryo-sac. Fig. 59. Endosperm nuclei from an embryo-sac containing an advanced embryo. (a) The single large nucleus above the antipodal cells, which under- goes no division; (6) The smaller secondary nuclei from the upper part of the embryo-sac; X 400. Fig. 60. Two sections of the upper part of an embryo-sac, showing the fertilized egg cell, 0, and one of the synergidz, s, which is still intact. Fig. 61. (2) Upper part of embryo-sac, with two-celled embryo; X 250. (6) Endosperm nucleus of the same. Figs. 62-66. Successive stages of the developing embryo, in longitudinal section; X 250. ss, primary suspensor cell; 4, a secondary endosperm nucleus. Fig. 67. Cross-section of an older embryo. Fig. 68. Longitudinal section of an embryo, showing the primary segments I, 2, 3. Fig. 69. Longitudinal section of an embryo, showing the enlargement at the base of the cotyledon. Fig. 70. Two nearly median, longitudinal sections of an older embryo; X 200; s#, stem-apex; cof, cotyledon; in a, the boundary of the plerome is indicated by a heavy line. Fig. 71. Median, longitudinal section of an older embryo. Fig. 72. Similar section of a much older one; X 70. Fig. 73. Longitudinal section of an embryo from a nearly ripe seed; X 4o. Fig. 74. Two sections through the apical region of an embryo of the same age as the one shown in fig. 73. Figs. 75, 76. Median, longitudinal sections through the root of advanced embryos; ~/, plerome; X 200. Fig. 77. Root of a full-grown embryo, showing the formation of inter- cellular spaces. Figs. 78, 79. Two transverse sections of the root just back of the apex; v, vessel; ev, endodermis; 7, intercellular spaces; X 400 pr; LETHARITTUN ine ee - r “2 = 7 Tr Tit a [ CAMPBELE. | 2 RATE 1h, — SPTICE INES AT Tefal Sate OS 2 Po CF oy ’ BA Hh ML Rey Wee OAD at PLATE IV. All the figures refer to Zannichellia palustris L. Fig. 81. Nearly median longitudinal section through the stem-apex; X 7o. The actual apex, 7, was not included in the section, but its position is indi- cated by the dotted line; s#, the sheath below the node; /!, Z?, the two ‘‘spathe leaves;’’ 7, lacunz in the leaves; 9, female flowers; ¢, male flowers; sg, squamule intravaginales; 2’, apex of a lateral branch. Fig. 82. Transverse section of the terminal buds. The leaves are num- bered; otherwise the lettering as in fig. 81. Fig. 83. Longitudinal section of stem-apex; X 200. The apex, +, was divided into two parts, only one of which shows in the section. Fig. 84. Transverse section of the staminal primordium; X 400. Fig. 85. Longitudinal section of the stem-apex after its division into the carpellary branch, 2, and the secondary stem-apex, v,; X 200. Figs. 86-88. Development of the stamen seen in longitudinal section; x 200. Fig. 89. Transverse section of an older anther. Fig. 90. Longitudinal section of an older stamen, showing the peculiar terminal appendage; X 60. Fig. 91. Two nearly mature pollen-spores. In 6, the primary nucleus is dividing. Two free nuclei, 4, are shown, derived from disintegrated arche- sporial cells; X 4oo. Fig. 92. Longitudinal section of young female inflorescence, showing the dichotomy of the plerome; X 400. Fig. 93. A similar section through the margin of an older inflorescence, showing the ovular rudiment, 0, and the subtending carpellary leaf, c. Figs. 94-96. Early stages in the development of the female flower in longitudinal section; X 400. 0, ovular rudiment; cay, carpel. Fig. 97. Section of the young ovule, showing the primary archesporial cell; X 400. Fig. 98. Cross-section of a young flower; X 400. car, Carpellary leaf; 7, the primary tapetal cell. Fig. 99. Longitudinal section of an older flower; X I50. Figs. 100, 101. Longitudinal section of ovules, showing the later divisions in the archesporium; /, tapetum. ’ Fig. 102. An older ovule, the nucleus of the embryo-sac already divided; X 400. Fig. 103. Longitudinal section through the inflorescence, showing its con- nection with the main axis; sh, the involuere. ee ee | CaMpuens | Pure Ww Asean. | oR As f m4 Bs, \ r> oh L} : ~) ‘aw “Xr geese SED : x x a0 is : ad rise iia Core ee FRertNne RE SE , ¥ r " STUDIES (IN > THE HERBARIUM AND THE BIR LD:—=Nortr. BY ALICE EASTWOOD, Curator of the Herbarium. CONTENTS. Piates VI-VII. ik REPORT ON A SMALL COLLECTION OF PLANTS FROM THE WHITE SEV ibis ieae fe [ea ud Ics, 320 OSS AUER ee Pa 71 Di ON SPURLESS HORMS OF AQUILEGIA 2205.0 c ese se ca ew aniemies 76 il.) beeen UNDESCRIBED CALIFORNIAN PLANTS......... 00.0.2 200508 78 TV. Cae MANZANITAS OF Mir: TAMALPAIS....... 0.0... 0.0050 000- 028 81 Ser Svan vole Colm RUATSS, 4 Ube bole bo bed COO ED Soo eIeeIORen Sere noie cio” 86 I.—REPORT ON A SMALL COLLECTION OF PLANTS FROM THE WHITE SANDS OF NEW MEXICO." Some months ago, Professor T. D. A. Cockerell of the College of Agriculture and Mechanic Arts, New Mexico, sent me a small package of plants for determination. They were collected on the ‘‘ White Sands’’ in August, 1896. They hint of a very interesting flora and indicate by a close alliance with well known species and by marked differences from the same, a peculiar environment and probably some 1 The traveler coming down Tularosa Creek, in the Sacramento Mts. of New Mexico, sees before him in the distance what appears to be the sea, with heavy breakers rolling towards the shore. As he descends to the valley, he gradu- ally realizes that the apparent ocean is motionless, and is above, not below, the level of the plain. Coming at length within a few miles of it, he sees before him what are to all appearances great banks of snow; and were it not for the intense heat, the illusion might be complete. Actually arriving at the banks, he finds nothing but pure white sand, piled up perhaps to a height of fifty feet above the plain, from which it rises abruptly, and continuing in un- dulating hillocks as far as the view extends. It is this remarkable formation that is known as the White Sands. Prof. C. H. T. Townsend has described it in similar words in an unpublished paper on the Distribution of Life in the Southwest and Mexico; and I believe almost any one would receive the same impressions on visiting the locality for the first time. L7r] November 23, 1897. 72 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. degree of isolation. The interesting description of the country, which Professor Cockerell has written, throws considerable light on the probable history of these plants. Cnothera tubicula filifolia, var. nov. ‘ PLATE VI, Fic. 1. Stems slender, suffruticose; older parts with grayish brown, shreddy epi- dermis; younger parts and leaves slightly glandular, hoary-puberulent, with spreading short white hairs; leaves sessile, becoming almost terete with the involute margins, 5-15 mm. long, 1-14 mm. or less wide, crowded at the ends of the branches and insmall rosulate clusters in the axils; calyx-tube 3-34 cm. long, funnel-form lobes spotted with dark purple, 1 cm. long, 4 mm. broad, with pointed tip 1 mm. long; corolla 34 cm. in diameter, yellow, tinged with white when withered, petals rhomboidal, slightly acuminate, upper margin wavy; stamens with anthers 1mm. broad, 7 mm. long, equalling the filaments; The exact extent of the White Sands I do not know, but they are probably about forty miles long and a good many miles across. They lie to the east of the San Andreas rangeof mountains. They are composed of pure gypsum, undoubtedly deposited from a salt water lake, which must have been shut off from the sea and by degrees have dried up. Prof. A. Goss, of the N. M. Agricultural Experiment Station, has pointed out to me, that as the lake dried up, the gypsum would be precipitated early, being comparatively insol- uble. Eventually the other salts would be deposited on the top of it. The more soluble salts have long ago been washed away, and we have remaining the beds of gypsum, which now rise considerably above the surrounding plain, the latter doubtless having been lowered by gradual denudation. While the plain itself contains a great quantity of gypsum, the banks are perfectly dis- tinct and well defined—as well defined as a miner’s dump. Somewhere in the sands, I am informed, there is a spring, and water is nowhere far from the surface. It might be thought that no vegetation would grow on pure gypsum sand, but there is a scattered growth, consisting of various shrubby and herbaceous plants. I even founda small poplar, which looked to me like Populus tremu- loides, though not typical; the poplar of the surrounding country, at least of the Rio Grande Valley, across the mountains, is P. fremontit, never P. tremuloides, at so low a level. Oneof the commonest shrubby plants on the sands is a Rhus, while a Bigelovia grows to aconsiderable size. When, last autumn, I visited this locality with Prof. C. H. T. Townsend, I was able to collect a few plants and insects; and my companion went somewhat further than I did, with the result of collecting two species of bees in quantity, and some other insects, which I had missed. Unfortunately, I was anything but well at the time, and we could not delay more than a short while, so that what was obtained was a mere fragmentary sample of the actual fauna and flora of the sands. T. D. A. COCKERELL. MESILLA, N. M., Feb. 28, 1897. : Bot.—VOL. 1.] EASTWOOD—STUDIES FROM THE HERBARIUM. 73 pistil extending 2 or 3 mm. beyond the stamens, stigma round and thick, more than 2 mm. in diameter; ovary 4-toothed at summit; capsule fusiform, contracted under the spreading, 4-toothed, reddish summit, 4-ribbed, splitting into four valves to within 1 mm. of the base and connivent at the top, persistent and becoming woody; seeds dark brown, irregular in outline, becoming mucilaginous when moistened; one row in each cell. This differs from typical @. tubicula chiefly in the very narrow leaves and a more condensed habit of growth, the slender branches becoming almost fasciculate. The pecu- liarities are probably due to poverty in the environment, which by slow starvation would bring about a reduction in the size and form of its organs. Gnothera albicaulis gypsophila, var. nov. PEATE Vii) Fic. 2. Stems woody, corymbosely branched, canescent throughout, with dense, closely appressed pubescence; leaves narrowly oblong to lanceolate, acute, 8-20 mm. long, 2-5 mm. wide, cuneate at base, margin sinuate-dentate, with 3 or 4 short teeth on each side, midrib conspicuous, petioles 2 mm. or less in length; flowers axillary, 3 cm. in diameter; calyx with tube 3 cm. long, slender, divisions 15 mm. long, united in one or two sets, free at the tips and bases; petals white, rhomboidal, as broad as long, narrowed at base toa broad claw, slightly wavy on the upper margin; stamens shorter than the petals, about equalling the style; capsule 3 cm. long, almost perpendicular to the stem, variously curved or even inclined to coil, splitting almost to the base into four narrow valves; seeds 1-2 mm. long, ovate-lanceolate, acute, obtusely angled, usually mottled with purple, minutely tuberculate. The description and figure were drawn from specimens without roots. The older parts of the stems present a ragged appearance due to the shreddy epidermis and the narrow spreading valves of the empty capsules. The desert environment is suggested by stiff, dry stems, close branches and leaves, and rapid development of the fruit. The flowers show but little tendency to become pink as they fade. nothera albicaulis is a variable species, easily recog- nized amid all its forms. Its most constant and distinctive characters are the white, shreddy epidermis, the capsules sessile by a hard, broad base, spreading from the axis so strongly as to make the stem somewhat zig-zig, valves sep- arating almost to the base and widely spreading. 74 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. This variety is nearest to § ¢rzchocalyx Engelmann,’ which has been named Anogra pallida Engelmanni Small.? The description of this variety is meager, but the erect and nu- merous branches of my specimen do not suggest ‘‘ parce ramosa,’’ nor are the seeds obtuse, as stated on page 334, Am. Journ. Sci. (II), Vol. XXXIV, under @. albicaulzs. The leaves approach those of var. v brevizfolza in size. The plate was made by tracing after a photograph. It has, therefore, greater accuracy than would be possible with an ordinary drawing. The flower, however, was drawn from a dried specimen soaked and spread so as to show its shape. Bigelovia graveolens appendiculata, var. nov. PLATE VI, FIG. 3. Stems light green, slightly tomentose except at the densely tomentose leat axils; leaves involute, becoming terete, mucronate; heads in rather few- flowered, corymbiform cymes on pedicels, often divaricately spreading; bracts of the involucre carinate, ciliate, and scarious on the margins, obtuse or acute at the woolly apex; corolla with from one to four yellow, linear appendages of various lengths on the tube, and a few cobwebby hairs on the border. This differs from all other forms of this variable species in the peculiar appendages of the corolla-tube. They are not transformed pappus bristles, being too far up on the tube, rather appearing like an outside corolla, as if the corolla were trying to become double. In other respects it approaches var. hololeuca Gray. Thelesperma gracile Gray. Bidens gracilis, Torrey, Ann. Lyc. N. Y., Vol. II, p. 215. Habttat.—On the Canadian River. The fragmentary specimen from the ‘‘ White Sands’’ shows but little variation from the type; the heads are smaller than those of any of the species represented in the Herbarium of the California Academy of Sciences. 1 Am. Journ. Sci. (II), Vol. XXXIV, p. 335. 2Torr. Bull., Vol. XXIII, p. 176. Bot.—VoL. I.] ZASTWOOD—STUDIES FROM THE HERBARIUM. 75 Muhlenbergia pungens 7hurber.! This is, according to C. J. Croft,’ a valuable forage plant, and is known in Arizonaas Black Grama and Grama China. It was determined by Prof. Scribner, who reports it as *¢ without unusual characters or special peculiarities.”’ There is also a branch of a shrubby Labiate, whose leaves in form and flavor suggest an Audzbertia; but as it is without flowers or fruit, it cannot be identified. 1Proc. Phil. Acad. Sci., 1863, p. 78. 2Proc, Cal. Acad. Sci. (I), Vol. III, p. 205. 76 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. II.—ON SPURLESS FORMS OF AQUILEGIA. Aquilegia differs from allied genera chiefly in having five spurred petals and a column of staminodia surrounding the pistils. In some species a tendency to degenerate from the markedly specialized type exists, and the varieties are characterized by spurless petals. These cause the flowers to superficially resemble those of /sopyrum, Clematis, or Anemone, more than Aguzlegza. Double-flowered forms of Aguzlegia are quite common in some species under cultivation, the stamens either becom- ing distinctly petaloid with spurs, or, together with the pet- als, becoming sepaloid, flat, and destitute of spurs. Agquilegia vulgaris, which is widely spread through Eu- rope and has, probably, been more extensively cultivated than any other species, has several varieties with ecalcarate petals. Two are given in De Candolle’s Prodromus, Vol. I, p. 50: A. stellata, with double flowers, petals flat, spur- less, and colored; A. degener, with double flowers, petals and sepals flat, spurless, and green. These are both figured in Clusius’ ‘‘ Historia Rarorium Plantarum,’’ according to De Candolle. In the ‘‘ Index Kewensis ’’ another variety, A. ecalcarata, is given, which the name indicates to be spur- less; but it is regarded by horticulturists as identical with A. stellata. The attention of the writer has been called to these pecu- liarities by the occurrence of two species in Colorado, which have ecalcarate varieties. The first of these belongs to A. cerulea. Aquilegia cerulea Daileye, var. nov. PLATE VII, Fic. 1. This is in all respects similar to the type except that the spurs of the petals are entirely wanting, sepals and petals flat, blue. The plate indicates the peculiarities of the variety. The plant figured was drawn from specimens sent me by Miss Bot.—VoL. I.] ZASTWOOD—STUDIES FROM THE HERBARIUM. 47 Anna L. Dailey. They were collected at Evergreen, Col- orado, where Miss Dailey noted them during several years. It was her opinion that the plants had become more numer- ous since she first noticed them. Mr. George E. Osterhout of New Windsor, Colorado, reports this form as frequent in Estes Park, Colorado. Some years ago, the writer found several plants with the spurless petals growing among the typical form in Platte Cafion, Colorado. inthe **Gardener s (Chromicle,”’ Vol. XVI, 1881, p: 16; the following note occurs concerning A. cerulea: ‘‘«Mr. E. G. Loder sends us a spurless flower of A guzlegza cerulea quite similar to the European stellate Columbines. The long, white, spurred petals are, in this case, absent and replaced by an additional whorl of ovate, acute, blue sepals. A few years since, Mr. Loder tells us he ‘ collected seeds of A. cerulea at an elevation of 10,300 feet, not far from South Park, Colorado.’ The seeds were sown in North- amptonshire in 1879. A few plants flowered last summer; but this year they have flowered most abundantly and the individual flowers are quite as fine as in their native habitat. Among the seedlings, one plant bears white flowers and others spurless flowers.’”’ Aquilegia micrantha Mancosana, var. nov.. This is the plant that was first described in ‘‘Zoe,’’ Vol. II, 1891, p. 226, as A. ecalcarata. Afterwards it was rede- scribed and figured from better specimens in Proc. Cal. Acad. Sci., Ser. 2, Vol. IV, p. 261. It is without doubt merely a degenerate variety of A. m/crantha, which is ex- tremely variable in both flowers and foliage, as I have in- dicated in Proc. Cal. Acad. Sci., Ser. 2, Vol. VI, p. 280. It seems desirable to give this a more distinctive varietal name, so I take this opportunity and name it after the region where it occurs. To Dr. B. L. Robinson, I am much indebted for the looking up of some references to which I had not access. 78 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. III.—THREE UNDESCRIBED CALIFORNIAN PLANTS. Iris Purdyi, sp. nov. PEATE Vil, ehiGse- Rootstocks slender, scarcely thicker than the fleshy roots; leaves dark green or somewhat glaucous, glabrous, erect or laxly Sonentine , surpassing the scapes, 6-7 mm. wide, 2-4 dm. long, with long, acuminate apex and mar- gins membranous and shortly ciliate, HOEK 15-20 cm. long, slightly flattened; bracts generally overlapping, inflated, glaucous, striate, tinged with rose-color, acuminate; spathes usually 2-flowered, similar to the bracts, but more inflated and more rosy, especially on the margins; pedicels 1 cm. long, about equal- ing the tube of the perianth; perianth with throat slightly dilated above the junction of the style; outer segments oblong, 7 cm. long, 2 cm. wide, rich cream-color, beautifully marked with fine lines of yellow on the claw, and with dotted veins of purple on the spreading blades; inner segments cream- color, somewhat shorter than the outer, widely spreading, linear-oblong, with margins strongly sinuate; stamens with filaments 5 mm. long, 2 mm. broad, narrowing abruptly at the insertion of the anthers; anthers 15-18 mm. long, 3 mm. wide, the polliniferous margins less than 1 mm. wide, edged on each side with purple; style slender, 12-15 mm. long, stigmas about 4 cm. long, including the crests, which are 1} cm. long, laciniate on the outer edge, tinted with pale rose-color to the ridge connecting the lobes of the stigma; stigma scale truncate, slightly undulate; capsules oblong, tapering equally at both extremities, valves about 12 mm. wide and 3 cm. long. This elegant /7zs is the species common in the Redwood region of Mendocino County, around Ukiah. It has here- tofore been included under 7. Dowglascana, which it re- sembles in its narrow, red-based, laxly spreading leaves, its cream-colored flowers, and its habitat. /. Dougdlaszana has always been considered an extremely variable species and includes a great number of forms, some of which may prove, as this had done, when carefully studied and com- pared, to be distinct species. . Lris Purdyi differs from other Californian species of rds in the peculiar bract-clothed, flowering stems. From /. Dougtlasiana it differs in having larger flowers, leaves lighter green, less distinctly nerved, somewhat stiffer, and some- times glaucous. The stigma scale is truncate instead of triangular-acuminate. The stamens are much broader, the capsule shorter, broader, and more uniform at each ex- tremity. The flowers are fewer in the spathes, less exserted, _— Bot.—Vot. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. "79 and on shorter pedicels that elongate but little in age. It forms clumps similar to 7. Douglasiana, but not so widely spreading. It is most fitting to name this species in honor of Carl Purdy of Ukiah, since he first detected its identity as a species entirely distinct from /. Douwglasiana and called at- tention to its peculiar characteristics. Montia rosulata, sp. nov. PEATE WiLL biG. 3° Annual, stems and leaves forming a compact, thick rosette, glaucescent, succulent; radical leaves terete or spatulate, 1-2 cm. long, 2-5 mm. wide; cauline leaves ovate-lanceolate, amplexicaul, 1-2 cm. long, 4-6 mm. wide; inflorescence almost concealed by the leaves, umbellate at first, becoming racemose, each cluster ona short, thick peduncle, subtended by a single lanceolate bract; flowers small, 3-4 mm. in diameter; petals white, unguicu- late, oblong-obcordate; stamens with purple anthers on short filaments; cap- sule inclosed in the calyx, opening by three valves that become involute and acuminate; seeds three, black, glossy, almost 2 mm. long and 14 mm. wide, minutely papillose, especially near the edge; strophiole white, conspicuous without the lens. : The seeds ripen rapidly and are shot to a distance of two or three feet by the elasticity of the valves. The empty valves form a 3-pointed star within the calyx. The flowers are ephemeral and the buds do not open in the house. This species belongs to the group of which J/. perfoliata Howell is the most common, and is intermediate between that species and JV. saxosa Brandegee. It approaches the latter most closely, from which it differs in longer and nar- rower leaves, smaller flowers, fruits, and seeds, and less succulence and density of foliage. The only locality at present known is on Mt. Tamalpais, where it was collected by the writer twice this year, on April 11th and 2oth. On the trail to the Lone Tree, not far from Rock Spring, there is an outcrop of volcanic rock whitened by long exposure to the elements, and here only these little rosettes of flowers and leaves are to be found, scattered around but not abundant, and in a limited area of a few rods. 80 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER. ‘ Newberrya subterranea, sp. nov. PLATE VII, Fic. 4. Stem brown, covered with broad, ovate or orbicular scales, which broaden as the stem becomes bulbous towards the inflorescence; scales surrounding the flowers becoming oblong-lanceolate, acuminate; flowers sessile on a disk- like receptacle 15 mm. in diameter; sepals of two brown, linear-lanceolate scales equalling the corolla, and two white membranous scales variable in length; corolla 1 cm. long, the four or five divisions as long as the tube, 2 mm. broad, oblong, obtuse, or emarginate, ribbed, densely hairy within, sparingly so ex- ternally, with ciliate margins; filaments varying in length, densely clothed with long hairs under the anther, sparsely hairy below, equalling or slightly surpassing the ovary. Anthers 2-celled, the half of each cell only polliniferous, ovate-elliptical; style thick, elongating with age, very hairy under the de- pressed, capitate stigma. The snow-white flowers contrast beautifully with the dark brown scales and stem. In the plant from which the de- scription is drawn, the middle flower is on the plan of five, while all the others are on the plan of four. This seems nearest to //emztomes pumilum Greene, Erythea, Vol. I, Pei Pet. It was communicated by Mr. R. A. Plaskett of Mansfield, Monterey County. He reports that he found it in bloom three inches below the surface of the ground,’ growing in Willow Creek Cajon, under Quercus densiflora. ‘This is the third plant that he has found under such unusual con- ditions, the other two having been lost before they could be sent. They all grew within a radius of forty feet and were from three to eight inches below the surface of the ground. The genus /Vewderrya has heretofore been found only in Oregon and Northern California. The discovery of this interesting species extends its distribution from Mendocino County to the southern extremity of Monterey County. Willow Creek is not far from the southern limit of the red- woods and abounds in the vegetation peculiar to the Redwood Region; so that the conditions are quite similar to those under which the northern species flourish, and it is, there- fore, really not surprising to find this northern genus here. 1Leaf-mould 1 to 1% inches; loam, or black soil, 3 inches; subsoil of coarse, loose, and mouldy nature, light-colored and very gravelly; on north hill-side, under heavy timber (Tanbark Oak and Redwood), where the sun never shines.—R. A. PLASKETT. Bot.—VoL. I.] ZEASTWOOD—STUDIES FROM THE HERBARIUM. 81 IV.—THE MANZANITAS OF MT. TAMALPAIS. Across the Golden Gate from San Francisco lie the picturesque hills of Marin County. These reach their highest point on Mt. Tamalpais, which rises to an altitude of 2604 feet. The rocky slopes of this interesting moun- tain are densely clothed with a shrubby vegetation, except in some spots where a bluish gray volcanic rock becomes exposed. The chief shrubs are Pickeringia montana, Erio- dictyon glutinosum, Castanopsis chrysophylla, Adenostoma fasciculata, several species of Quercus and Ceanothus, Um- bellularia Californica, Vaccinium ovatum, and the different kinds of Arctostaphylos or Manzanita. Of these manzanitas, there are at least four forms that appear quite different from each other; but the amount of variation is so great among some of them, and with one exception they are so different from all forms definitely described, that their identification is a puzzle that has been engaging the attention of the writer for more than a year on frequent trips to the mountain, at all seasons and over its many trails. If the manzanitas in other sections of Cali- fornia present as many variations as these on Mt. Tamal- pais, it will be many years before the genus Arctostaphylos is understood, and then, probably, the results will be secured only through systematic cultivation of the various forms in Botanic Gardens. A beginning is already made in the Botanic Garden of the University of California, where several species are now growing. The inadequacy of printed descriptions and the impos- sibility of deciding, without the types for comparison, to which species some of the Mt. Tamalpais forms are most closely allied, makes it seem necessary to describe and name each form, that seems undescribed, as a distinct species, leaving to the future monographer of the genus the task of assigning definite limits, if that be possible in so polymor- phous a genus, which continually suggests hybridization or a very active and unlimited tendency to vary. 82 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Arctostaphylos nummularia Gray. This is distinct and can not be mistaken for any other species. It is in bloom at some place on the mountain throughout the year. It rarely grows amid dense brush, seeming to prefer somewhat exposed situations, and usually attains a height of about two feet, seldom more than four. It is quite common and always conspicuous, with young stems erect and slender, invested with a halo of white hairs, flowers small and very abundant, and leaves round and shining. The fruit, which is flattened longitudinally and without the mealy pulp of other manzanitas, differing also in the deciduous pericarp, which at maturity leaves the nut- lets naked, sets this apart from all other species and renders the name of manzanita, or little apple, inappropriate when applied to this species. Arctostaphylos glandulosa, sp. nov. Shrubby, never arborescent, 2-8 feet high, young branches straight and erect, older ones divaricate; young shoots glandular-hirsute, with long and short spreading hairs; leaves erect on short petioles, oval to ovate-lanceolate, mucronate, thin, bright green, or sometimes thicker and inclined to be glau- eous, sometimes spinulose-dentate even on flowering stems, an inch or two long, usually glandular-pubescent and glutinous; older stems smooth, dark brownish red, with thicker leaves, inclined to be orbicular, often glaucous, and without glandular hairs; racemes usually few in the panicles, loosely or densely flowered; bracts glandular, persistent, lanceolate-acuminate, green and foliaceous below, usually longer than the pedicels, shorter, often rose-colored and petaloideous above; sepals orbicular, glandular-ciliate, white, membra- nous, stellately spreading after anthesis, often revolute; corolla large, glabrous externally, hairy within, white or pinkish; stamens with filaments densely bearded but not glandular; style shorter than or equalling the corolla; ovary flattened horizontaliy, densely covered with white hairs more or less tipped with rose-colored or white glands; fruit spherical, horizontally flattened, transverse diameter about I cm., vertical diameter 5-7 mm. The immature but fully grown fruit is often tinged with red on one side; when ripe it becomes reddish brown, with the mealy pulp quite abundant. Sometimes it has a purplish bloom, and is generally slightly hairy and glandular. The pyrenz are variously coalescent and frequently united into one. This is the manzanita earliest in bloom on the mountain, the most beautiful, the most fragrant, most variable, and most widely spread. On the upper slopes and towards the Bot.—Vot. I.] ZEASTWOOD—STUDIES FROM THE HERBARIUM. 83 ocean, the most glandular specimens are to be found; while on the Mill Valley Trail, the hills near Larkspur, and those above the reservoir of the San Rafael Water Company (localities somewhat protected from the ocean winds), it becomes almost glabrous except on the pedicels, the ovary, and the young growth, where some glandular hairs are al- ways evident. When fresh shoots spring up where a fire has formerly swept over the mountain, they appear so dif- ferent from the most common form as to suggest, at least, a good variety. The young leaves are often a bright red, appearing like a brilliant flower from a distance. Usually they are green, and often tinged with red, especially on the margins. This species has been included under A. tomentosa, but its peculiar characteristics have been ignored in all descrip- tions of that species. In some of its features it resembles A. Pringlet, which was also formerly included under A. tomentosa. It resembles A. tomentosa in the shape and ar- rangement of the leaves and the erect tendency of the branches. The long, loose, woolly hairs so noticeable on young stems of A. ¢omentosa are in this species replaced by shorter and usually glandular hairs. The ovary and fruit are glandular and this, according to the various de- scriptions, is never the case with A. tomentosa. The glandu- losity is so pronounced in typical specimens of this species that the leaves feel gummy when touched, and stick to each other if placed together. When drying, they are very fra- grant. While this species somewhat resembles A. Priuglec in the character of its pubescence, it differs from it in the shape and texture of the leaves, the persistent bracts, and the flattened instead of conical ovary. Arctostaphylos montana, sp. nov. Shrubby, widely spreading, young branchlets usually erect; lower stems glabrous, dark reddish brown; upper stems canescent, with a dense covering of short curly hairs; leaves venulose, minutely punctate, oval, 1-2 cm. long, with narrow cartilaginous margins and prominent mucro, canescent when 84 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. young, becoming glabrous and glossy when old; petioles stout, 2 mm. long, pubescent, with minute blackish glands intermingled; panicles short, on rather stout peduncles; pedicels white, slender, glabrous, 5 mm. long; bracts short, 2-3 mm. long, canescent at base and minutely glandular, the apex brown, triangular-acuminate, deciduous; sepals green at base, thin and white above, involute, surrounding the ovary after anthesis; corolla small, 5 mm. long, with roundish, reflexed, ciliate lobes, externally glabrous, hairy within; filaments dilated at base, glabrous; ovary glabrous or slightly pubescent, conical; ripe berries glossy, reddish brown, variable in size and shape, 5-10 mm. in diameter, spherical or flattened either longitudinally or transversely, the pyrene either separable or coalescent into one, and extremely variable in shape; pulp thin. This appears to be nearest to A. Hookerz, from which it is chiefly distinguished by thicker, less veiny leaves, shorter and stouter petioles, shorter bracts and longer pedicels, and much thinner pulp. This manzanita loves the exposed ridges, where the bluish gray volcanic rock crops out, and is usually found associated with Quercus dumosa var. bul- lata. The description was drawn from specimens collected near the cypress trees on the trail between the Eldridge Grade and Larsen’s. It is abundant there, forming low, spreading bushes not more than three feet in height. It is also found around the reservoirs of the San Rafael Water Company and along the road from Fairfax to Larsen’s. It is the last species in bloom, not coming into flower before April. The fruit is ripe in September or October and per- sistent for some time after ripening. The new growth is made after the flowering period, the canescent leaves and stems being very conspicuous contrasted with the dark green, glossy, older foliage. Arctostaphylos canescens, sp. nov. Shrubby, forming erect or low, spreading bushes, densely clothed through- out with short white curly hairs, with minute blackish glands intermingled on the stem, petioles, pedicels, bracts, and lower surfaces of the leaves; leaves oval, ovate, or obovate, shortly acuminate, mucronate, about 3 cm. long, 1} cm. wide, on stout petioles 5 mm. long; racemes few or solitary, almost condensed into corymbs, with persistent foliaceous bracts 5-12 mm. long, surpassing the stout pedicels, red-nerved and margined or rosy through- out; sepals oval, white or rose-color, ciliate, densely hairy within, usually sparsely hairy externally, revolute, spreading after anthesis; corolla 7 mm. Bot.—Vo.. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 85 long, with short, orbicular lobes, glabrous externally, hairy within, white or tinged with pink; stamens with filaments somewhat bearded; ovary densely covered with white hairs, but not glandular, slightly rounded at top; style a little longer than the stamens; fruit similar to that of A. glandulosa. This manzanita, while very different in appearance from either A. ol/andulosa or A. montana, has some characteris- tics of both. The condensed raceme and the pubescence are similar to those of A. montana, the pubescence being that of the young growth of A. montana; while the size and shape of the flowers, the foliaceous bracts and stout pedicels resemble those of A. glandulosa. The fruit, also, is large and with abundant pulp. _ It fruits sparingly. Its season of blooming is earlier than that of A. montana, but generally not so early as that of the first blooming bushes of A. elandulosa. The bushes from which the description is drawn grow on the trail from the Eldridge Grade to Larsen’s, and are to be found between the last bunch of cypresses and the Rock Spring. They are close to the trail and conspicuous in contrast with the bright green, very glandular specimens of A. glandulosa in the immediate vicinity. It is abundant on a trail recently cut from West Point to Rock Spring. Pro- fessor W. R. Dudley of Stanford University has sent me specimens of the same species from above Felton and from Loma Prieta in the Santa Cruz Mountains. NA 5 CALIFORNIA EXPLANATION OF PLATE VI. Fig.1. Cnothera tubicula filifolia, var. nov. a. Longitudinal section of the calyx, showing pistil stamens. ete: 6. Node of the stem magnified to show leaves and pubes- cence. ) c. Ripe capsule, actual size. Fig. 2. Cnothera albicaulis gypsophila, var. nov. Fig. 3. Bigelovia graveolens appendiculata, var. nov. oF a. Flower magnified, showing corolla appendages, etc. — 6. Capitula magnified, showing involucre, flowers, etc. | Eastwoon.|Prare VI EXPLANATION OF PLATE VII. Fig. 1. a welt teil OE * 25. Spergularia macrotheca HEYNH. (Tissa.). ......../| * = 26. Platystemon Caltfornicus BENTH... . * = 27. Lepidium bipinnatifidum DESV. (L. Menztesii of Bot. Cal.) 28. Lepidium nitidum NUTT.... . 29. Dithyrea Californica var. maritima DAVIDSON (Biscutella)) 30.1 Oligomeris glaucescens CAMBESS. (O. subulata WEBB.) ..| * 31. Lupinus micranthus DOUGL.. .-..-+ + .+ 252s 0s | * 32. Lupinus albifrons BENTH. (L.Chamissonis)........ |» 2 33. Trifolium microdon H. & A. var. pilosumvar.nov... ... Sue LS SADIE PL DIELS OLDIE, eae wh oy Site ovieh iby se! 1s) ies tise. Le! «=) =)'| 34. Trifolium gracilentumT.&G.......... 35. Lrifoltum stenophyllum NUTT... 2.2. 2 3 2 ss ss 20 despa Pamnert WATS... 6 5 5 sles ue we ee } Byam) Lrsfoleum dichoromsne W SA. so ws ae ees is ss ein LURES ORO East 1a Cyeaaect oe tty Che oO C CL Oe Oma mca Et ne 39. Medicago denticulata WILLD.. . = 40. Astragalus didymocarpus H. & A... ...+..222es Al) Astragalus Traskie Sp. NOVs« = 5 3 6 6 ss ws sss 42. Hosackia venustasp.NOv.........+.-. relic, abroes Asn Erogeum cicutarium THER... se es +e 8% sw os * - 44. Malva pusilla SMITH (M. borealis)... 2... 1.20. * ig 45. Cinothera viridescens HOOK...........2e00. * “s 46. Mesembryanthemum nodifiorum 1... . 46a. Mesembryanthemum crystallinum\,..... . * - 47. Opuntia Engelmanni SALM. vat. littoralis ENGELM?...| * = 47a. Opuntia prolifera ENGELM. . . . 22s es oe ee AS te Daucus PustiusMICHS, . 2 2 sss 1). = se e's * x AQnAprastram angustijoirum NUTT. sles ek se es Re Son wommicsia Meneses IN COB oi isle in ee ee ete es SI. Peucedanum tnsularé sp.nov... ... 2.5.2.5 ee ee area (relia IVEUSRI GRAVES ter aticl eros) ie tol ole! 6 ys. he xdvdluay ao. 2 53-1 Convoluulus macrostegius GREENE..... ...... * ‘a 54. Hehotropium Curassavicuml,.. .. 1s. 1 oe sine ae Ee CCLOCAT YE LthCArIS WICi ee let a ee be) 3 lela 0) ca tea Malts ‘2ZnID ByURg |-eurrese9 equeg * | ‘aquea[Q uBg “puLlMeEy 120 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. ee Be ee ve ie SPECIES. 2 g z E : 5 a | 56.1 Cryptanthe maritima GREENE... + +--+ ses ees 57. Cryptanthe Torreyana GREENE. .++ +++ +--+: * * 58. Amsinckia St. Nicolaisp.NOV...- + +++ +++++-: 59. Amsinckia maritima Sp. NOV... +--+ +++ ee renee ? 60. Lycium verrucosum SO MIOVs | ole o) oh ayes! eh oan eae ee ae 61. Lycium Californicum NUTT... .. ++ + eee ee ees il be * 62. Orthocarpus purpurascens BENTH... +++ ++ ++ - +s cael ese * 63. Plantago insularissp.NOV.... . +++ + eee ee eee 64. Malacothrix indecora GREENE... * 65. Malacothrix implicata sp. NOV... ... +. eee eee ‘3 - * 66. Miucroseris linearifolia GRAY. .1 + ee 22 eet eee ae a PE * 67. Sonchus asper WIEL. 6. 2 tw be a ee . ee dN as * 68. Souchwus tenerrimus Wyss 0 3) sree 6 6 oe ws wie altel ** * 69. Sonchus oleraceus Ty. +--+ see se eee eee ees a “ 3 70. Centaurea MelitenstsVy.?. .. ++ + ee ee eee * a. | LAPLOWESID i one tee Me Yd Lo Wels) (ol els sa las Vila pela, teas eats ge tt ery eaters a Te? « [ow |e |e Ale 73.2 Amblyopappus pusillus H.& A... 2-2 ee eee ees * * * | * * 74. Baeria gracilis GRAY... +--+ ++ +++ ee eee eae * * * * 75.3 Hemtzonia Streetsti GRAY. ..+.++- See et * * 76.1 Leptosyne gigantea KELLOGG. ... ..+ + e+e ee * * * * * 77. Franseria Chamitssonis ESS. var. viscida var.nov... . - * Fransertia Chaprssomts WESS:. se 8 et ne ew | * 78. Franseria bipinnatifida NUTT. var. dubia var.nov. .. . « H” LP Beprnnaifeda. aie okwr ao) ei) SF 0: = yah Sie ls t= eee ig * Fo ee ¥ 79. Baccharis consanguinea DC. ?(B. pilularis)........ * * * * 80. Bigelovia veneta or Grindelia?.....++-+ + es *Closely related species found on some of the other islands but different from the form on San Nicolas. 1 Guadalupe Island, Baja California. 2 Cedros Island, Baja California and S. America. 3 Sau Benito Island and Anacapa Island, Calif. 4 This might more correctly be considered a form of 7. Macrai with the heads on long peduncles. 5 Species reported but not collected. SUMMARY. 82. Species recorded. 64. In common with the mainland, of which about 30 are Californian. 53. In common with Santa Catalina Island. 48. Incommon with Santa Cruz Island. 31. In common with Santa Rosa Island. 31. In common with San Miguel Island. 6. In common with Guadalupe Island; 1 with Cedros Island, 1 with San Benito Island and 1 with Anacapa Island ; 7 species and 3 or 4 varieties peculiar to San Nicolas. Bot.—Vot. I1.] EASTWOOD—STUDIES FROM THE HERBARIUM. 121 II.—NEW SPECIES OF CNICUS FROM SOUTHERN COLORADO AND UTAH. 1. Cnicus bipinnatus, sp. nov. Cnicus Drummondii var. bipinnatus Eastwoop, Zoe, Vol. IV; p. 8. Glaucous and glabrous except for some slight arachnoid tomentum on the stems, petioles, and involucral bracts: stems stout, erect, leafy, 6 dm. or more high, branching from the base and also above; leaves with numerous linear-lanceolate divisions which are 2-6 cm. long, 5 mm. wide and irregularly parted near the base, generally on one side only, into similar lobes varying in length and sometimes as long as the main division, margin laciniate-dentate, spiny, lateral spines 2 mm. long, terminal 4-5 mm. ; radical leaves petiolate, 1%-3 dm. long, 6-10 cm. wide; cauline, sessile, 10-15 cm, long, 5-6 cm. wide: heads corymbose at the ends of the leafy branches, almost sessile, narrow, cylindrical, 4% cm. long, 1 cm. or more wide; involucre of appressed, imbricated bracts, successively shorter, in seven ranks, the lower ones pointed with a weak prickle 3-5 mm. long, the upper attenuate to a scarious tip, minutely puberulent; flowers purple; corolla with tube about half as long as the throat and divisions, throat about one-third as long as the linear, clavate-tipped divisions ; stamens surpassing the corolla ; style straight with the node 24 mm. from the tip; akenes glabrous, shining, flattened, obovate- oblong, 6 mm. long, 3 mm. wide, surmounted with a yellow ring. This is nearer to Cnicus Rothrockid than to Cuicus Drum- mond but differs from the former in its foliage, narrower heads, short, weak spines, and general appearance. It formed a clump two feet or more in height and almost two feet in diameter. Collected in Colorado in Johnston Cajfion, near where it joins the Mancos River, in a locality but rarely visited by white men. Closely related to this is the plant which the writer! described under the name C. Rothrockit var. diffusus. J take this opportunity of giving this, too, specific rank. 2. Cnicus diffusus, sp. nov. Similar to the above in habit and surface: leaves narrower, less deeply divided, with more regular and triangular lobes; spines at the tips of the lobes 1 cm. long, stiff, yellow, those along the margins 1-3 mm. long: heads somewhat broader than the last, with outer involucral bracts tipped with stiff spines from 1-2 cm. long, deflexed-spreading in fruit, inner bracts attenuate ’ 1 Proc. Cal. Acad. Sci., Ser. 2, Vol. VI, p. 303. 3 Sept. 13, 1898, 122 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. to along spiny point; flowers purple; corolla with tube four-fifths of the throat and divisions ; throat about one-third of the tube, divisions linear, abruptly acuminate, 18 mm. long; stamens surpassing the corolla; node of the stigma 2 mm. from the apex ; coma 2% cm. long. Willow Creek, San Juan Co., Utah, Aug. 13, 1895. 3. Cnicus Hesperius, sp. nov. Stem stout, simple, erect, leafy from the base, ribbed, almost glabrous, 3-4 dm. in height, 2 cm. in diameter : leaves 10-15 cm. long, 114-2 cm. wide, upper surface glabrous, lower tomentose, ascending, linear-lanceolate, with numerous rounded lobes, spiny-margined with two to three large, yellow, subulate spines, 5 mm. long, and several shorter ones: heads sessile, crowded, in an oblong, terminal, erect, leafy, glomerule; involucre 2 cm. long, 2% cm. wide, with bracts 3 mm. broad at the yellow, glabrous, ovate base, tapering to a long brown-purple spine 1 cm. long, arachnoid with silky wool except at the glabrous yellow apex: flowers light purple ; corolla tube almost equalling the throat and divisions ; throat fusiform, contracted under the divisions which are linear with thickened apex and about half as long as the throat ; anthers surpassing the petals by 3 mm., sparingly arachnoid except at the pointed tips ; style with inconspicuous node concealed by the anthers and stigma, exserted 2 mm. ; coma I cm. long. Mt. Hesperus on the Bear Creek Divide above timber line, La Plata Mountains, southwestern Colorado, Aug., 1892. Collected by the writer. This Cnzcus is nearest to C. erzocephalus Gray, under which it was placed by the writer in ‘‘ Zoe,’’ Vol. IV, p. 8. A recent inspection of the specimens of Czzcus in the Her- barium of the Academy has convinced me that this deserves to have specific rank and that it is not a hybrid with Czzcus Parryt, as | had formerly conjectured. The involucral bracts are less densely tomentose than in C. eriocephalus, the glomerule erect, the flowers light pur- ple or pink, the entire plant less arachnoid, and the stamens have not only the filaments wooly but also the anthers. I have named it in honor of the mountain on which it is found, the highest in the La Plata Range. There were few individuals growing along the trail leading to the sum- mit of the ridge. A Bot.—Vot. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 123 IlII.—THE COLORADO ALPINE SPECIES OF SYNTHYRIS. One alpine species of Syzthyrzs is now recognized from the mountains of Colorado. This has been aptly named S'. alpina. 1. Synthyris alpina Gray. PLATE IX, Fics. 1a-1d. This is found on most of the high peaks of Colorado, “growing above timber line, in loose, rocky soil. Its low stature and short spike of dark purple flowers superficially distinguish it from the other species. There are always four calyx divisions, variable in shape and size even in flowers from the same spike, and conspicuously fringed with long white hairs. The corolla consists of two parts, united at the base; the upper broadly obovate, entire, and slightly concave; the lower two or three cleft, with laciniate or entire divisions varying in length and breadth. The sta- mens and style are moderately exserted and the flowers are erect. In the little-explored mountains of southwestern Colorado two different species are found above the tree limit. One of these is so distinct that in a group so closely related as that to which Syzthyrzs belongs, it might be taken as the type of a new genus; the other approaches S. alpzna. The placing of these two species in Synthyris necessi- tates a change in the generic characterization of the calyx. Instead of ‘‘calyx with four divisions’’ it must become ‘‘ calyx with two, three, or four divisions.’’ This difference in the number of calyx divisions arises probably from a union of parts not infrequent in other genera. 2. Synthyris Ritteriana, sp. nov. PLATE IX, Fics. 2a-2e. Sparingly pubescent with short, scattered hairs ; scape stout, nearly 3 dm. high, closely covered from the middle of the stem to the spike with foliaceous bracts : leaves radical, 8-10 cm. long, 3% cm. wide, oblong-elliptical, obtuse, 124 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. cuneate and decurrent at base, crenate ; petioles stout, 8-12 cm. long; bracts of the scape alternate, 114-214 cm. long, 10-18 mm. wide, ovate-acuminate, sessile by a subcordate base, crenate-dentate near the apex; floral bracts rhomboidal, acuminate, tapering at base to a short petiole which surpasses the pedicels : flowers white, erect on short pedicels in a spike 5 cm. long; calyx of three divisions varying in shape and size in flowers from the same spike, 4mm. long, orbicular to obovate, entire, toothed or cleft, obtuse or acute, fringed with white hairs less dense and shorter than those on the pre- ceding species ; corolla of two divisions, surpassing the calyx by 2 mm.; upper part broadly obovate, acute, indistinctly spurred at base, ciliate; lower variously cleft with two to three laciniate, ciliate divisions ; stamens with orbicular-ovate anthers 1 mm. broad and filaments inserted at the base of the corolla, surpassing it by 2 mm.; fruit unknown. Collected by the writer in Cumberland Basin, La Plata Mountains, Aug., 1892. It grew in the alpine meadow where moisture was abundant. It is named in honor of Mr. and Mrs. B. W. Ritter of Durango, Colo., to whose kindness I owe the opportunity of visiting these mountains. The plate shows the differences in the floral organs of the three species, all drawn to the same scale. 3. Synthyris reflexa, sp. nov. PLATE IX, Fics. 3a-3d. Glaucous and somewhat viscid; scape erect, 1%-2 dm. high, clothed below the spike with broad, foliaceous bracts: leaves radical, 7 cm. long, elliptical-oblong, obtuse, truncate or cuneate at base, finely crenate, thin in texture ; petiole 3-5 cm. long: bracts of the scape alternate, crowded, 2% cm. long, 2 cm. wide, sessile by a broad subcordate or truncate base, broadly ovate, acute, entire or serrulate near the apex, diminishing upwards; floral bracts ovate to linear, fringed with long white hairs: flowers greenish white, reflexed on pedicels 1 mm. long ; calyx of two divisions 5 mm. in diameter, orbicular, fringed like the bracts; corolla of two parts united at base, the upper broad, somewhat hood-shaped, 5 mm. broad, 8 mm. long, sparingly fringed ; the lower two-cleft 5% mm. long, 4mm. wide, divisions laciniate, 2 mm. long; stamens two, anthers cordate, with cells not con- fluent, filaments inserted at the base of the corolla, surpassing it by 4 mm.; style shorter than the filaments, stigma capitate ; fruit unknown. Collected by the author in Kendall Basin, in the San Juan Mountains, near Silverton, Aug., 1890. It was quite rare. This is the only Syxthyrzs known to have reflexed flowers and two divisions to the calyx. Bot.— VOL. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 125 It is very doubtful if the genus Syzthyrzs will stand in the future as it is now set forth in the Synoptical Flora; nor is it more certain that Professor Greene’s transferrence of all the species to Wadfenza will be final.!’ In a group of genera so closely related and so polymorphous as those included under subtribe Veronicee in Bentham and Hooker’s Genera Plantarum, botanists will always differ in regard to generic limits. A difference in the arrangement of the leaves and habit of growth primarily separated Wulfenta from Veronica; the form of the corolla, the number of calyx divisions, the dehiscence of the anthers and the shape of the seeds sep- arated Synthyris from Wulfenia. Professor Greene has shown the worthlessness of considering the number of calyx divisions as a generic character, and his position is reinforced by the two new species described in this paper and by two described under Wulfenia by Aven Nelson.? Professor Greene, however, does not give a diagnosis of Wulfenia nor tell why it, also, should not be included under Veronzca. If habit of growth is taken as a generic distinction, then Synthyris naturally falls into two genera, one containing S. rotundifolia and S.reniform7s; the other, the remain- ing species and perhaps the original Wudfenca. If other characters are to be taken, many genera would result and the synonymy become much involved. It seems simplest to follow Bentham and Hooker and Gray and leave these uncertain problems to the future; so the new species of the Colorado mountains have been described as Synthyris. 1 Erythea, Vol. II, p. 80. 2 Torr. Bull., Vol. XXV, pp. 281, 282. 126 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. IV.—FURTHER OBSERVATIONS ON THE MANZANITAS OF MT. TAMALPAIS. Since the publication of the article on the ‘‘ Manzanitas of Mt. Tamalpais,’’? I have been able to explore parts of the mountain and the neighboring hills hitherto unknown to me. Scarcely a week has passed that I have not spent Sunday on the hills of Marin County. As a result of these frequent visits and the new territory explored, new facts concerning the distribution, time of blooming, and characteristics of the different species of Arcto- staphylos have been observed. 1. Arctostaphylos nummularia Gray.— This I have found only on Mt. Tamalpais proper. It does not seem to grow on the hills around Fairfax, where the Big Carson and the Little Carson Creeks take their rise, nor on the ridge between Bear Valley and Bolinas. It is abundant on the Boot-jack and Throckmorton Trails, also on the trail leading directly from Mill Valley up the mountain and on the trail from the Potrero to the head of Cataract Gulch. While well formed fruit can be found at almost any time of the year, ripe fruit is seldom seen and never persists as does the fruit of all the other species. 2. Arctostaphylos canescens Hastwood.— This, too, seems to be confined to Mt. Tamalpais and apparently grows only on the southern slope. It loves bleak, gravelly hillsides, where it often holds exclusive possession. On the Throckmorton Trail a few bushes are to be seen, a few more on the Bill Williams Trail; but on the West Point Trail it is abundant, adorning the slopes during the winter with its blooming bushes. On November 7th, the first flowers were seen. The plants on the Throckmorton Trail were beginning to bloom. The delicate pink blossoms have their beauty much enhanced by the gray-green foliage. From this date until See this Vol., p. 81. Bot.— VOL. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 127 February 2oth, it was seen in bloom every Sunday. Well formed fruit was observed on the bushes on the Throck- morton Trail before those on the higher West Point Trail had ceased blooming. Some plants only a few inches high, easily pulled up by the roots, and with but one or two branches, were laden with blossoms. The number of these small plants indicates a species full of vigor, seeking new territory for habitation. The roots of the larger plants are spreading rather than deep and are easily pulled up. Dur- ing May the berries become ripe, and before the other spe- cies show ripe fruit this has usually lost all its berries. 3. Arctostaphylos glandulosa Aastwood.—On Decem- ber roth the first plant of this species was found in bloom. It may have been blooming also elsewhere on the mountain; but two weeks earlier flowers were not seen on the plants near the foot of the Boot-jack Trail, where I have always found them first in bloom. On May rst, fully grown fruit was observed on bushes on the Boot-jack Trail. This species is the most abundant, found everywhere on the slopes and ridges of Mt. Tamalpais and the Fairfax hills. I am still uncertain whether two species ought to be recog- nized where I have described but one or whether, with all its forms, it ought not to be considered a variety of A. tomentosa. | 4. Arctostaphylos montana Zuastwood.—This has a wide range in Marin County. It is usually found only on the uplands and is especially partial to the bluish gray vol- canic areas. Wherever Cupressus Goventana and Quercus dumosa var. bullata are to be found this also will be seen. Besides Mt. Tamalpais, it is found on the Fairfax hills, where it covers large sections of country, being the most abundant manzanita there. It was first seen in flower February twenty-second, on the trail to the source of the Big Carson. Only one bush was in bloom, situated in a warm and sheltered spot. On May ist it was still blooming 128 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. on Mt. Tamalpais, but evidently nearing the end of its season. Having had an opportunity of examining fresh specimens of A. Hookerz in bloom, sent me from Monterey by Miss Marion Rouse, I am confirmed in my belief in the identity of this species as distinct from A. Hookerz. The flowers of the latter are much smaller, the leaves not so thick, and the fruit, as noted before, has much thinner pulp. Bot.—Vot. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 129 V.—TWO SPECIES OF ERIODICTYON HERETOFORE INCLUDED UNDER ERIODICTYON TOMENTOSUM. The genus E4riodictyon was founded on £. crassifolium, collected near San Diego, Calif., and described by Ben- tham in the Botany of the Sulphur, page 37. Previous to this, another species had been described and figured by Hooker and Arnott as Wigandia Californica.’ In the Bot- any of the Sulphur, Bentham transferred this to Hrzodictyon, giving it the appropriate name &. glutinosum. It again suffered a change of name when Torrey?’ restored its earli- est specific name, calling it &. Californicum. Bentham described still another species in the Botany of the Sulphur as &. tomentosum, differing from EZ. crassz- folium in having pedunculate dense cymes, broader leaves, and very numerous, smaller flowers. This Dr. Torrey took as the type under which &. crass¢folium should be included, and suppressed the former name, giving, as his reason, the numerous intermediates observed by Dr. Parry and him- self around San Diego and elsewhere. Gray accepted Torrey’s view with the remark: ‘‘ 4. cras- sifolium Benth. was doubtless rightly united with this by Dr. Torrey, and this name should have been preferred, but the other is good and of the same date.’” Greene described a plant from Monterey County, Calif., collected by Mr. Brandegee in 1885, as true &. tomentosum, and restored the name £&. crasszfolium to the plant of Southern California.* The attention of the writer was attracted to this puzzle by the rediscovery of Mr. Brandegee’s plant near Jolon, in the San Antonio Valley, Monterey County, Calif. Truly it seemed that Professor Greene had solved the mystery; but to be certain, specimens of this from Monterey County and of the large flowered £. crassifolium from San Diego were sent to Kew for comparison with the types. The following 1 Bot. Beech., Pl. LX XXVIII, p. 364. 2 Bot. Mex. Bound. Sur., p. 148. 3 Proc. Am. Acad., Vol. X, p. 331. 4 Bull. Cal. Acad. Sci., Vol. I, p. 201. 130 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. reply was received: ‘* Hrzodictyon crassifolium and £. tomentosum are conspecific and your plant named tomen- tosum is apparently an undescribed species which we also possess from J. G. Lemmon without locality, and from G. R. Vasey, Monterey, 438, coll. July, 1880.”’ Since Lemmon, Vasey, and Brandegee have all passed through Jolon when on their way to the Santa Lucia Mountains in search of Adzes bracteata, doubtless the region of the writer’s collection of this species coincides with theirs and will be considered as the type locality of the new species. 1. Eriodictyon niveum, sp. nov. PLATE X, Fics. 3a-3d. Densely white-tomentose or flavescent; stems three or four feet high, growing in clumps, very leafy below the inflorescence: leaves thick, elliptical- ovate or obovate, 4-6 cm. long, 1-3 cm. wide, entire or crenate, except on the cuneate base, apex acute or obtuse, lower surface reticulate-rugose, upper with veins scarcely discernible; petiole broad, 5-10 mm. long: panicle terminating a long, naked peduncle, compactly or widely branched, the stout branches varying much in length on different plants; cymes densely flowered, with the lower bracts obovate or spatulate, tapering to broad petioles, the upper oblanceolate to linear; flowers small, almost sessile; calyx equalling the corolla tube, divisions linear-subulate, densely clothed with white silky hairs; corolla white or tinged with lavender, 4 mm. long, urceolate, glandu- lar-hirsute externally, glabrous within, tube furrowed longitudinally, slightly contracted under the five small spreading lobes; stamens with the free portion short, inserted below the throat, anthers oval, 1 mm. long; styles shorter than the sepals; capsules orbicular, obtusely 5-angled, tomentose, especially on the angles; seeds four or sometimes five, brown, minutely favose, variable in shape, often keeled, more than 1 mm. long. Collected by the writer near Jolon, Monterey County, Calif., in flower June 1, 1893, and in fruit Sept. 22, 1894. The following specimens, besides, are in the Herbarium of the Academy :— Lobb, San Antonio Valley (date not given); T. S. Bran- degee, Monterey County, 1885; Dr. Palmer, ‘‘ Perhaps from Monterey County,’’ 1876. There is also a specimen without flowers or fruit collected by T. S. Brandegee at Zapato, Fresno County, Calif. The foliage and pubescence are exactly of this species. Bot.—Vot. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 131 2. Eriodictyon Traskie, sp. nov. PLATE X, FIGs. 2a@-2¢. Densely white-tomentose, except the dark-colored, glandular-hirsute calyx: leaves elliptical, 5 cm. long, 15 mm. wide, acute at apex, the base narrowed to a petiole 5mm. long, margins dentate except near the base, veins distinct on the lower surface, barely evident on the upper: panicle slightly surpassing the leaves, with branches spreading or curving upwards, rather slender, glandular and tomentose, bracts from elliptical with dentate margins to lan- ceolate or linear with margins entire; cymes densely flowered: flowers small, on pedicels 1 mm. or more in length; calyx divisions five, narrowly linear, not uniform in length, 4-5 mm. long; corolla purple, the tube equalling the calyx, 5 mm. long, contracted at base and throat, furrowed longitudinally, divisions of the limb irregularly orbicular, not uniform in size, glandular- hirsute externally as well as the upper part of the tube; stamens inserted half way down the tube, almost sessile; style branches glabrous, 1% mm. long; ovary ovoid, glandular-hirsute; fruit unknown. This was discovered May, 1897, on one volcanic upland on Santa Catalina Island, Calif., at an elevation of about 1500 feet, by Mrs. Blanche Trask, the indefatigable local botanist in whose honor it is named. Probably, this is the plant collected by Lyon on Santa Catalina, referred by Dr. Gray to &. tomentosum.’ It approaches &. nzveum but is undoubtedly a distinct species. A plant slightly differing from the above was collected in the Santa Inez Mountains, Calif., by T. S. Brandegee, in 1888. | The peduncles are stout, with thicker and more spread- ing branches, leaves larger and coarser, pedicels much longer, 4 mm. long, while the filaments are short but dis- tinctly evident, the corolla has the same shape, the lobes of the border being not quite so broad. As this, too, is in flower only and very young, comparisons of the fruit and seed cannot be made. The plate shows the floral organs of &. crasstfolium from San Diego, part of the collection sent to Kew, and also those of &. niveum and E. Traskia, all drawn to the same scale, five times the actual size. 1 Suppl. Synop. F1., p. 420. 132 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. VI.—NEW SPECIES OF PACIFIC COAST PLANTS. 1. Campanula angustiflora, sp. nov. PLATE XJ, Fics. 2a-2Cc. Annual, scabrous-hispid; stems slender, angled, 1-3 dm. high, branching diffusely from the base to the top with upwardly spreading branches: leaves sessile, ovate to orbicular, acuminate or acute, deeply dentate, 5-15 mm. long, 3-10 mm. wide: flowers axillary on stout, upwardly spreading pedun- cles, twice to four times the length of the flower; divisions of the calyx linear- subulate, almost equalling the corolla, connivent in fruit; corolla tubular, with five triangular lobes; stamens included, the anthers linear, longer than the thin, broadly triangular-subulate filaments; ovary obovoid, slightly con- stricted at the apex, ribbed; style short, thick, with three revolute stigma lobes; fruit strongly ribbed, irregularly humped with the three valves above the middle; seeds numerous, minute, shining, light brown, with a small darker spot at one end, 3-sided or keeled. This has been included under C. exzgua Rattan by Mrs. Brandegee, who first discovered it on Mt. Tamalpais, July 5, 1886, collecting it again in the same locality June, 1890, and July, 1893. She also found it on Mt. St. Helena, May to July, 1889.1. It was rediscovered on Mt. Tamalpais by Mr. J. W. Congdon, and by the writer near the water tank at the head of the East Fork of Sequoia Cajion on the rail- road track. Besides the points of difference shown by the figures of the two species, there are differences in habit of growth and general appearance. C. exzgua is lower, more slender, less branched, and with the branches divaricately spreading; the leaves are smaller and narrower, and almost hug the stem. The figures are drawn from a specimen collected by Volney Rattan on Mt. Diablo, Calif. It is probably part of the type. In the Herbarium of the Academy there are specimens from Mt. Hamilton, Calif., collected by W. W. Price, similar to those of the Mt. Diablo Campanula. From Priest Valley in Monterey County, Calif., very young specimens of an annual Campanula were collected by the writer, May 12, 1893, resembling C’. angustiflora in 1 Zoe, Vol. I, p. 83. Bot.—Vot. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 133 habit of growth, shape of corolla, and character of stamens and style. The leaves are much narrower, and the calyx divisions vary in length in the same flower and surpass the corolla. This I leave under C’. angustzflora, as these small annual Campanule may be more common than is now supposed and the two species may vary a good deal among themselves. Their season is short, they grow in out-of-the- way places, and they are inconspicuous; so the chances of their discovery and collection are small. ' 2. Romneya trichocalyx, sp. nov. PLATE XI, FIGS. 4a-dc. Perennial, glaucous and glabrous except for the scattered, spreading sete on the peduncles, rhachis, petioles, and lower margins of the leaves; stems many, suffruticose, laxly spreading from the base as they grow older, leafy, branched: leaves rather thin, with conspicuous venation, ovate-orbicular in outline, variable in size and divisions, pinnately 3-5 parted, the lower divi- sions entire, toothed or lobed, the terminal larger, cuneate, 3-5 cleft; leaves on the peduncle closely surrounding the bud, much smaller, the divisions linear, narrower, more numerous, and more setose-ciliate; petioles flat, 5 mm. to 2 cm. long: calyx of three imbricated sepals covered with upwardly appressed, scabrous setze, except near the margins and on the underlapping parts; corolla white, texture crape-like, 8-15 cm. in diameter, variable in the shape, size, and number of the petals; stamens numerous, with linear-oblong anthers and slender filaments, the lower half brownish purple, the upper yellow; styles 1o-11, viscid, incurved; ovary ovoid, densely setose; capsule oblong-ovate, the walls breaking irregularly from the stout, straight ribs of the framework; seeds not seen. This has long been included under Romneya Coulter?, from which it is most markedly distinguished by the setose calyx. There is no doubt of the plant with smooth calyx being true 7. Coulter7, since the description and figure of the type confirm it.’ Miss Kate E. Cole of Oakland first drew my attention to the fact that there were two kinds of Aomneya in cultiva- tion, describing the marked differences between them; but it was not until last fall when I myself saw the two kinds growing side by side in Golden Gate Park, San Francisco, that I began to look up the matter. 1 Lond. Journ. Bot., Vol. IV, p. 75, Tab. III. 134 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. So far as I have been able to discover, there are but two plants of 7. trichocalyx in the Park. Both grow back of the statue of Halleck where two paths meet. A. Coulter grows there too; but the best specimen is to be found near the Haight St. entrance. It is also very abundant in other parts of the Park, being more desirable as a cul- tivated plant owing to its greater size and compact habit. Taking the plants in the Park for comparison, supple- mented by the specimens in the Herbarium of the Cali- fornia Academy of Sciences, the following are the points of differences :— Habit.—R. Coulteri forms large, erect, close clumps, with many strong branches from along the stems. . ¢trichocalyx does not form close clumps because the stems as they grow tall have a tendency to lean over as if too weak to stand; they are more leafy, less branched, and neither so tall nor so stout. Leaves.—The leaves of R. Coulteri are thicker in texture, with fewer and larger divisions, becoming simple on the peduncle but never growing close under the flower, thus leaving the upper part of the peduncle naked. The differences in the leaves, however, are not always to be depended on as the leaves of 2. trichocalyx are so variable, often closely approaching those of R. Coulteri. The upper leaves of R. trichocalyx however always become more dissected on the peduncle and grow close under the flower. Inflorescence.—The peduncles of 2. Coulteri are stouter and more spread- ing than are those of &. trichocalyx. Calyx.—R. Coulteri has a smooth calyx; that of R. trichocalyx is setose. Corolla.—In the specimens in the park, #. Coulteri has larger flowers, with the texture less crape-like than in P. trichocalya. We have specimens of the latter in the Herbarium with corollas fully as large as any of A. Coulteri. Fruit.—When the walls of the capsule break away leaving the skeleton of the pod, the ribs of the pod of R. Coulteri are more slender, becoming atten- uated towards the apex and convolute; those of 2. ¢richocalyx are stouter, uniform, and do not twist around. The figures show the differences in the buds and the leaves of the peduncle. They were drawn from fresh specimens from Golden Gate Park, smaller than ordi- narily because they were the last of the season. Both spe- cies bore fruit, but as it did not ripen, I was unable to com- pare the seeds. The pods from which the comparison was made came from herbarium specimens. Bot.—VoL. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 135 The following specimens of each species are in the Her- barium of the California Academy of Sciences :— Romneya Coultert, Anaheim, July, 1885, M. K. Curran, (with fruit and buds). Baja California, W. G. Wright, (in flower). Golden Gate Park, San Francisco, (cultivated). Santiago Creek, near Orange, Orange Co., Calif., Miss Agnes Bowman, June, 1898, (with flowers and buds). Romneya trichocalyx, Aliso, Baja Calif., T. S. Brandegee, May 30, 1893, (in flower and dry fruit). Sausal, Baja Calif., T. S. Brandegee, June 4, 1893, (in flower). Cafion de Gato, Baja Calif., T. S. Brandegee, June 5, 1893, (stem only). Near Temecula, Riverside Co., Calif., No. 393, S. B. Parish, Oct., 1882, (in flower). Matilija Cafion, Ventura Co., Calif., F. W. Hubby, May 18, 1895, (in flower). Santa Maria R., Santa Barbara Co., Calif., Mrs. Ida E. Blochman, (a bud only). Golden Gate Park, San Francisco, (cultivated). 3. Sedum Congdoni, sp. nov. PLATE XI, Fics. 5a-5d. Stem 1-6 cm. high, simple or branched from near the base, with slender, erect, tortuous branches: leaves alternate, 2-4 mm. long, 1-2 mm. wide: very fleshy, ovate, obtuse, sessile, the place of insertion above the base; flowers yellow tinged with red, sessile in sparingly branched, few-flowered cymes terminating the branches; calyx with five short, broadly triangular divisions, acute and red-tipped; petals five, ovate-lanceolate, less than 2 mm. long, red at the apex; stamens ten, with thread-like filaments shorter than the petals; anthers kidney-shaped; ovaries five, tuberculate near the apex, 1-ovuled; styles curved outwards; fruit unknown. This might be mistaken for Sedum pumilum, since both are small and have one-seeded follicles. The latter has much larger yellow flowers, linear-lanceolate petals, erect styles, and glabrous ovaries. The fringe of hairs at the suture of the follicle is much longer and finer in S'. Bumzlum than in S. Congdoni. The former is farinose when young, becoming glabrous with age. The figures are designed to show the differences in regard to the petals and pistils of the two species. 136 CALIFORNIA ACADEMY OF SCIENCES. [Pxoc. 3D SER. This was discovered by Mr. J. W. Congdon, at Grant’s Springs, Mariposa County, Calif., and collected April 9, 1898. It adds another to his namesakes in Mariposa County, appropriately associating his name with the flora which he has done so much to make known. 4. Cercocarpus Traskie, sp. nov. PLATE XI, Fics. 7a-7e. Tree, 10 to 25 feet high; trunk 2 to 10 inches in diameter, 6 to 8 feet to the lowest branches; bark rough, grayish brown externally, reddish on the inside; upper branches covered with a thin, downy tomentum: leaves orbic- ular to oval, 2-6 cm. long, 1-5 cm wide, with obtuse or acute apex, subcor- date, truncate, or cuneate base; margin revolute from deeply dentate to entire; upper surface dark green, glossy, glabrous, except the downy young leaves; lower surface densely white-tomentose, veins large and conspicuous on both sides; petiole stout, about 5 mm.long. Inflorescence androgynous, the polygamous flowers numerous in axillary umbels; calyx white-tomentose, with tube 1 cm. long and border 5-toothed, open campanulate, 5-8 mm. in diameter, glabrous within; stamens numerous, anthers tomentose, with two linear-oblong cells united only at the insertion of the slender filament; perfect flowers with stigmas curved like shepherd’s crooks, style exserted; akenes t cm. long, linear-oblong, covered with upwardly appressed silky hairs, tipped by the circinate, persistent style, about 5 cm. long, clothed with long, fine, silky hairs spreading horizontally. This, the most beautiful of the Pacific Coast Cercocarpz, was discovered by Mrs. Blanche Trask at the southern part of the island in a volcanic region known as ‘‘ Salte Verde.”’ It is a wild place, too rough for men on horses, with no trails but those made by the goats. Even in winter the heat is great. She writes as follows concerning the place and the trees: ‘‘There are about forty or fifty trees in an arroya so small that there is but room to squeeze through, a southern exposure where Ruin and Earthquake have passed and in whose footprints but few plants have dared to rise.”” The sea dashes at the base of this arroya, the walls of which rise to a height of from 100 to 500 feet. The trees are all isolated, not at all forming thickets. That any one should have found a new tree on an island that so many botanists have visited is surprising; but it is due to the great enthusiasm, the wonderful power of Bot.—VOL. I.] EASTWOOD—STUDIES FROM THE HERBARIUM. 137 exploration, and the intense love for Santa Catalina Island and its flowers which Mrs. Trask possesses. It is with pleasure that I give her name to this tree. Photographs of several of these trees taken recently by Mrs. Trask show them to have widely spreading branches and graceful habit, and to be well worthy of cultivation. The branches are abundantly adorned with rosettes of white tomentose flowers at all the leaf axils, the contrast of which against the dark, glossy green of the upper leaf sur- face is striking and beautiful. The same contrast occurs between the upper and lower surfaces of the leaves, the beauty of which is enhanced by the strong, even venation and revolute margins of the leaves. This tree is unlike the other Pacific coast species and perhaps approaches C. fothergilloides H. B. K., the Mexi- can species, more nearly than any other. It seems to be a type of Cercocarpus isolated and distinct. 5. Calochortus Purdyi, sp. nov. PLATE XI, Fics. 8a-8/. Glabrous and glaucous; stem 2-3 dm. high, rather stout, erect, branching, two to many-flowered, not bulbiferous at base: radical leaf solitary, sheathing the stem, linear-lanceolate, acuminate, 2 dm. long, 1 cm. 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 the bracts, erect in flower, recurved in fruit: flowers broadly open-campanulate; 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, somewhat arched by the nar- row, transverse, semicircular, conspicuous gland, the shallow pit of which is covered by a densely hairy narrow scale; anthers lanceolate, abruptly acumi- nate, cream color or purplish, shorter than the filaments, which broaden to the base; capsule 3 cm. long, 2 cm. wide, broadly elliptical, with the three thin wing-like valves transversely veined. This belongs to the § Aucalychortus according to Wat- son’s arrangement in the Botany of California. In habit it resembles C’. a/bus, but in general is more like a giant C. 138 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Maweanus. Its nearest relative is, however, C. Tolmiez, which belongs to the same region and from which it can readily be distinguished by the absence of the scale cover- ing the gland of the latter. Most of the C. Tolmzez in herbaria is probably this species. It was compared with the original specimen of C. Tolmzez at Kew by J. G. Baker who says: ‘* The Willamette plant differs a good deal from the original C. Tolmzet. C. Tolmzez has pale lilac petals bearded all over the face, no spot, no scale, no obtuse anthers.’ There is no true C. 7olmzez in the Herbarium of the Academy. We have specimens collected by Thos. Howell, from Grant’s Pass, Oregon, from Prairies, West- ern Oregon, April 1881, and from Hillsboro, Oregon, May 1881, all marked C. Tolmzez, but each with a scale covering the gland and with flowers creamy rather than blue. It would seem as if the anthers were variable, since all of C. Purdyt that I have examined have acuminate anthers; but evidently the specimens sent to Kew had obtuse anthers. C. Purdyz grows in the Willamette Valley, in the foot- hills on dry gravelly soil. It is never found in shaded woods. It is named in honor of Carl Purdy who knows Calo- chortt more intimately than anyone, and whose work on the genus in the garden, the field, and the study has accom- plished so much towards determining the true specific limits in this difficult and variable genus.! 1 Since the above description was written and the drawings made, a beautiful figure of this species has appeared in ‘‘ The Gardeners’ Chronicle,’ Vol. I, 1898, p. 305, fig. 147. The flower is much larger than that of any specimen seen by me. It is not unusual, however, that plants have larger flowers under the more favorable surroundings that cultivation produces. Itisa well known fact that Pacific coast species vary consider- ably in the size and vigor of individuals according to the amount of moisture and fertility of the soil. a, Win ran Piety) ; h ¥ had i ih a re EARN UNS Thales Me Alte | Anh MANE Nie Set wi mh fy rn AN iH Viale 140 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. 6. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE VIII. Peucedanum tnsulare, sp. nov. a. and 6. Parts of the same leaf, actual size. c. External view of carpels, magnified as shown in figure. d. Cross-section of a carpel, magnified as shown in figure. Hosackia venusta, sp. nov. a, Calyx. 6, Standard. c, Keel. d, Wing. Figures mag- nified 5 times. Hosackia argophylla GRAY. a, Calyx. 6, Standard. c, Keel. d, Wing. Figures mag- nified 5 times. Abronia alba, sp. nov. a, Flower. 6, Immature fruit. Figures magnified 5 times. Lycium verrucosnm, sp. nov. a. Tip of branch, actual size. 6. Flower magnified 5 times. c. Interior of corolla showing the position of the stamens. d. Interior of the calyx showing the pistil. Astragalus Traski@, sp. nov. a. Fruiting branch, actual size. 6. Flower, twice the actual size. c. Cross-section of the pod. d. Mature pod. Amsinckia St. Nicolai, sp. nov. a, Calyx. 6, Corolla spread open. c, Outline of corolla. d and e, two views of the akene. Figures about 5 times the actual size. Amsinckia maritima, sp. nov. a, Calyx. 6, Corolla spread open. c, Outline of corolla. Enlarged as in fig. 7. [Easrwoon | Prare VIL S TRASKIA .Z457W0Q0. yor CavAcanSct.32Ser Bor VarI. [Eas woon} PLare Vill 7, AMSINCKIA ST.Nit 8 AMSINCKIA MARITIMA. AA&77 3. HOSACKIA ARGOPHYLIA. GHAY. 4 ABRONIA ALBA. /457WOG07. 5. ASTRAGALUG TRASKIS.Z4A57WOQ7. Easrwoon, DEL | / } ia 142 CALIFORNIA ACADEMY OF SCIENCES. _ [PRov. 30 4 EXPLANATION OF PLATE IX. All parts of the flowers are 5 times actual size. i Fig. 1. Synthyris alpina GRAY. | a. Flower. b.,. Calyx, c. Lower part of corolla. d. Upper part of corolla. Fig. 2. Synthyris Ritteriana, sp. nov. a. Flower. - 6. Calyx. candd. Lower part of two different corollas. e. Upper part of corolla. Fig. 3. Synthyris reflexa, sp. nov. a. Flower. 6. One of the equal divisions of the calyx. c. Lower part of corolla. d. Upper part of corolla. od Pretareate: ten Bon vill cists rete ITA. a te Weysabal NTHYRIS ALFINA LALWAAL. 3, OYNTHYRIS fasrwoon, JEL, REFLEXA. LA57WOGD. [Eastwoon | Phare IX. 2 SYNTHYRIS RITTERIANA ZLAS7WOCZ. GG LUE BRITTON GREY, 5S Wa fi h i Bite at ike . CALIFORNIA EXPLANATION OF PLATE X. All figures are 5 times actual size. Fig.1. Eriodictyon crassifolium BENTHAM. a. Part of corolla showing the stamens. 6. Corolla. c. Part of calyx showing the pistil. Fig. 2. Eriodictyon Traskia, sp. nov. a. Interior of corolla showing the stamens. 6. Corolla. c. Calyx spread open showing the pistil. Fig. 3. Lviodictyon niveum, sp. nov. a. Interior of corolla showing the stamens. 6. Corolla of a young flower. c. Corolla with lobes fully expanded. d. Calyx spread open showing the pistil. [Eastwoon | Prare X LERIODICTYON CRASSIFOLIUM AEW7AAM. 2 ERIOUICTYON TRASKIAZ. ZAS7WOLZ. Eastwoon, DEL. ‘'S ERIODICTYON NIVEUM 4457W2QZ. LTE BRITTON & REN, SF; 146 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XI. Campanula exigua RATTAN. a, Flower fully expanded. 4, Fruit. c, Pistil and stamen. Figures 5 times actual size. Campanula angustiflora, sp. nov. a, Flower expanded. 4, Corolla spread open. c, Fruit. d, Pistil and stamen. Figures 5 times actual size. Romneya Coultert HARVEY. a, Bud. 6, Immature pod. Figures actual size. Rommneya trichocalyx, sp. nov. a, Bud. 6, Immature pod. c, Leaf. Figures actual size. Sedum Congdon, sp. nov. a, Plant actual size. 6, Flower. c, Follicle. d, Petal and stamen. The flower and its parts enlarged ro times. Sedum pumilum BENTHAM. a, Petaland stamen. 4, Follicle. Enlarged 1o times. Cercocarpus Traskia, sp. nov. a. Tip of a twig, actual size. 6. One of the larger leaves, showing the lower surface. c. Anther enlarged. d. Fruit. e. Flower. Calochortus Purdyi, sp. nov. Stamen enlarged. Ripe pod. a. Plant, actual size. 6. Petal. c. Scale on petal enlarged. d. Sepal. é. Se LOUSROTON GREYS FASTWOGD. CALUCHORTUS PuRDY? Z4s7waaz. 7 CERCOCARFUS TRASKIS | UMILUM AENTHAM. 5 SS : = =| = 19 =z o ie at aig 3. ROMNEYA CouLTERI 242 4, ROMNEYA: TRICHOCALY? CAMPANULA ANGUSTIFLORA A457WOaD. L CAMPANULA .EXIGUA AATTAW. PHYCOLOGICAL MEMOIRS. BY DE ALTON SAUNDERS. CONTENTS. Pirates XII-XXXII. PAGE Ie SOMEVPACIBIG, COAST EICTOGARPACE Al, oo.) ..6.e6 a,c a2 synveyar cle aheins see erase 147 II. SPHACELARIACEA AND ENCCELIACE OF THE PACIFIC COAST..... 157 EXPAN AION OBI EASES 2 ate ms bytes cieete cf Glos athelerebevelelld aslacaaccte ce 166 I.—SOME PACIFIC COAST ECTOCARPACE. Family ECTOCARPACE C. Ag. Ectocarpacee C. AG., Syst. Alg., XXX, 1824. Emend. Thur. Le Jol. List. Alg. Mar. Cherb., 1863. Plant body arising from a mass of creeping filaments or a disk-like mass of cells mostly monosiphonous, more or less branched ; reproductive bodies of two kinds—plurilocular sporangia formed of numerous small, densely aggre- gated, linear, lanceolate, or ovoid cells; unilocular sporangia cuboidal or globose. SYNOPSIS OF GENERA. The basal part of the plant consisting of branching filaments. The plurilocular sporangia intercalary with the cells of the vegetative EMI CERSGUESGE AU Neo ahah fe ett Meta tare bale) aieha/aic aeigs 2b se 5, dt inle 9-4 bis Pylatella. The plurilocular sporangia not intercalary. Basal filaments mostly superficial .......... PR eon ena ae . Ectocarpus. Basal filaments ramifying through the tissue of the infested plant Streblonema. The basal part of the plant consisting of a cellular, disk-like mass. RVR AAV SOS: <4 a, el Spee ops a pina nc cere be eisle ein e's a) dials) ogee oleate we Phycocelis. 1. Phycocelis Strem/. Phycocelis StraMF., Not., III, 1888, 383. Plant small epiphytic, the basal part consisting of one or two layers, more or less circular in outline, increasing by peripheral growth. Erect filaments present; unilocular sporangia and paraphyses unknown; plurilocular sporangia sessile or stout; zoosporal cells mostly erect in a single row. [ 147] Oct. 29, 1898. 148 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 2. Phycocelis reptans (Crouwan) K7ellm. PLATE XII, Fics. 8-12. Phycocelis reptans (CROUAN) KyELLM. Handbk. I, 1890, 81. Ectocarpus reptans CRovuaN, FI. Finist., 161, 1867. Plant forming rounded patches 1-5 mm. in diameter, the disk-like base of one or two layers of cells 12-15 broad. Erect filaments 2-4 mm. high, unbranched, ending in an abrupt, hyaline hair; cells 8-12 broad, as long to twice as long as the diameter; chromatophores small, oval or band form; plurilocular sporangia lanceolate or linear-oblong, sometimes curved, 25-754 (commonly 40-55) x 12-30, terminal on the erect filaments or rarely sessile on the basal ones. On the leaves and air-bladders of /Vereocystis lutkeanus. Pacific Grove, California. 3. Phycocelis fecunda S/rem/. PLATE XII, Fics. 1-7. Phycocelis feecunda StrazMF., Not., 1888, 383. Spots orbicular, % mm. or so broad. Creeping filaments united to form a disk-like, unistratose, basal part; the erect filaments simple, of 2-6 cells which become transformed into sessile or stipitate, uniseriate, cylindrical, plurilocular sporangia, 27-40 x 6-10; a cluster of long hairs in the center of each spot. On Macrocystis pyrifera, Desmarestia ligulata, and Ptery- gophora Californica. Pacific Grove, California. 4. Streblonema Derb. & Sol. Streblonema DERB. & SOL., Cast. Cat. Pl. Mars. Suppl., 1851, 100. Plant body filamentous, consisting of two parts, the numerous branching filaments ramifying through the tissue of a host plant; erect filaments branched or sessile, usually ending in or bearing long hairs; unilocular spo- rangia large, subglobose; plurilocular sporangia simple or sometimes branched, one to many seriate. 5. Streblonema fasciculatum ( Thur. ) Le Jol. PLATE XIII. Streblonema fasciculatum Tuur. in Le Jou. Alg. mar. Cherb., No. 100, Liste P- 73- Vegetative filaments irregularly much branched, branches alternate, irreg- ular, 6-9” broad, cells 1-3 times as long as broad, slightly constricted at the joints; unilocular sporangia globose or ovoid, sessile or rarely stalked, 25-352 X 21-304, borne on the main filament or the branches; plurilocular sporangia lanceolate or linear, obtuse at the apex, sessile or stalked, borne on the branches near the surface of the infested plant, 40-1007 x 8-12y, often uniserlate. Bor.—Vot. I.] SAUNDERS—PHYCOLOGICAL MEMOIRS. 149 In Nemalion andersontt Farl. San Pedro, California, Aug., 1896. The plant is composed of an irregularly branching mass of filaments running through the soft central filaments of the Vemalion. Some of the branches extend out to the surface and are simple; others are once or twice dichoto- mously branched near the surface. The plurilocular spo- rangia are very irregular, born singly or sometimes aggre- gated on the outer sides of the branches, not at all branched. Long and colorless hairs are born on the branches with the plurilocular sporangia. 6. Ectocarpus Lyngd. Ectocarpus Lyncs., Hydrophyt. Dan., 1819, 130. Plant filiform, branching, attached to the substratum by branching, creep- ing filaments. Erect filaments monosiphonous, branched, occasionally corti- cated by outgrowths of superficial cells ; unilocular and plurilocular sporangia terminal or lateral, never intercalary. 7. Ectocarpus acuminatus, sp. nov. PLATE XIV, FIGs. I-5. Plant minute; creeping filaments forming a compact network; erect fila- ments simple, of the same size from base to apex, a mm. or so high; cells not at all constricted, 12-14 broad, below as long as the diameter, above 2-3-5 times as long ; chromatophores elliptical, much more abundant in the central part of the filament ; plurilocular sporangia sessile on the base of the erect filaments or arising on a long or short stalk from the creeping filaments, lanceolate, very long-acuminate, sometimes tipped with a short hair, often more or less curved, 90-300p (rarely 400) x 20-30/. Forms minute, light brown tufts in the conceptacles of Cystosetra osmundacea. Pacific Grove, California, July, 1896; San Pedro, California, Aug., 1896. The creeping filaments ramify through between the para- physes and attach themselves to the inner wall of the con- ceptacle; the erect filaments extend out of the mouth of the conceptacle a mm. or so; the plurilocular sporangia point toward the opening or extend just out of it. 8. Ectocarpus ellipticus, sp. nov. PLATE XIV, FIGs. 6-9. Plant minute, erect, tufted, attached to the inner wall of the cryptostomata or conceptacles by a colorless mass of creeping, branching filaments. Erect I50 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. filaments unbranched or with a few unicellular branches, 9-12 broad ; cells at base once to twice as long as the diameter, above three times as long; chromatophores oval, numerous; plurilocular sporangia linear or linear-lan- ceolate, 75-Ioop x 18-25, borne laterally at or near the base of the erect filaments or occasionally arising directly from the creeping filaments, sessile, or the lower on short stalks; zoosporal cells 5-8 broad, one to few seriate ; unilocular sporangia sessile, ovate or elliptical, 30-40y x 12-184, borne on the same tufts with the plurilocular sporangia. Forming minute, dark brown tufts 1 mm. high in the cryp- tostomata and conceptacles of Fucus evanescens. Pacific Grove, California. g. Ectocarpus chitonicolus, sp. nov. PLATE XV, Fics. 1-4. Plant 1-2 mm. high; creeping filaments numerous, branched, 15 broad; cells once to twiceas long as the diameter; erect filaments mostly unbranched, 14u broad at base, somewhat narrowed above; cells below %-2 times as long as the diameter, above 2-3 times as long, not at all constricted at the joints; chromatophores numerous, small and oval; plurilocular sporangia lanceolate or narrowly ovate, obtuse at the apex, 90-175” x 20-354, borne laterally in the erect filaments or occasionally on the creeping ones; the lower sporangia on a long or short stalk, the upper sessile. Forms minute tufts on the back of a chiton. Pacific Grove, California. A variable species which approaches &. cylindricus in some of its forms but is separated from it by the more pointed, plurilocular sporangia and the uniformly smaller size of the vegetative parts. 10. Ectocarpus cylindricus, sp. nov. PLATE XVI. Creeping filaments irregularly branched, 12-20y broad; cells 2-3 times as long as the diameter; erect filaments simple or giving off a few short branches below, of the same size throughout, 18-30 broad; cells below and above 2-3 times as long as the diameter, in the central part as long as the diameter, not at all constricted; chromatophores numerous, disk shaped ; plurilocular sporangia lateral, on a one- to several-celled stalk, erect, cylindri- cal or obovate, very obtuse above, 80-2007 x 35-454, opposite, most abundant below; unilocular sporangia on separate filaments, ovate or elliptical, 60-1204 x 30-40, usually on a one-celled stalk, or the upper occasionally sessile, often opposite. The plant forms a compact, velvety mass 2-4 mm. high, of indefinite extent on the stems and rhizoids of Agregia Bot.—VoL. I.] SAUNDERS—PHYCOLOGICAL MEMOIRS. I51 menzzesit, on the fruiting tips of Cystosetra osmundacee, on rocks, on Codium. adherans and C. mucronatum Califor- nicum. Pacific Grove, California, 1895. The form on Cystosecra averages slightly larger and many of the plurilocular sporangia arise directly from the creep- ing filaments; no unilocular sporangia were found on this form. In the forms that grow on a hard substratum, the creeping filaments are short and form a compact network on the surface; in those that grow on a soft substratum (Codium, etc.) the creeping filaments are much elongated and ramify between the loose threads of the host plant. 11. Ectocarpus hemisphericus, sp. nov. PEATE XVII. Plant densely tufted, arising from a compact network of creeping filaments. Erect filaments unbranched at base, primary branches dichotomous, diver- gent, long, not at all narrowed upward, ending bluntly above, secondary branches numerous, short, clustered, many of them ending in a short, blunt hair; cells at base 15-20 broad, 2-3 times as long as the diameter, above 22-3014 broad, 1-2 times as long as the diameter, slightly constricted at the joints ; chromatophores small, numerous, linear or disk-shaped ; plurilocular sporangia lanceolate or ovate, obtuse, short, 30-90% x 14-20p (rarely 304 long), borne laterally on a short stalk in the upper part of the filament; uniloc- ular sporangia short, cylindrical, 30-35 x 20-25, on a one-celled stalk, on the same filament with the plurilocular sporangia. The plant is very abundant, forming dense, hemispherical tufts 2-4 mm. high on Pelvetia fastigiata and Taonza /enne- backere. San Pedro, California, Aug., 1896. But one specimen was found of the form on 7. denne- backere. 12. Ectocarpus hemisphericus minor, form. nov. PLATE XVIII, Fics. 1-3. Vegetative filaments simple or but slightly branched, 16-214 broad. The plant forms small, dark brown tufts 1-2 mm. high on the fruiting tips and upper branches of Fucus harvey- anus. The whole plant is smaller and does not form the dense rounded tufts as in the type. San Pedro, California, Aug., 1896. I52 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 13. Ectocarpus paradoxus pacificus, var. nov. PLATE XVIII, Fics. 4-7. Erect filaments arising from a compact mass of creeping ones, 25-40" broad, divergently branched, branches long, bearing above several short, pointed branchlets ; cells about as long as broad, near the extremities one-half as long, slightly constricted; plurilocular sporangia cylindrical, ovate or lanceo- late, obtuse or abruptly pointed at the apex, borne laterally on the main stem and branches on a long or short stalk, 70-150 x 25-50; unilocular sporangia on short stalk or intercalary, globose, about 3o0y in diameter. Forming small tufts 2-5 mm. high on Fucus evanescens, most abundant on the fruiting tips; differs from the type in its smaller size, in the branching which is never opposite, and in the plurilocular sporangia which are longer and more pointed. 14. Ectocarpus mucronatus, sp. nov. PEATE LX Plant light olive-green, attached at base by a few long, colorless, creeping filaments. Erect filaments 1-5-6 cm. high, loosely intertwined, terminated above bya short, colorless hair, unbranched at base; main filament 30-4op broad; cells one-third to as long as the diameter, below three times as long, not at all constricted, branches spreading, numerous, scattered, mostly short and pointed ; chromatophores disk-form, numerous in each cell; plurilocular sporangia sessile, numerous, 50-155/ X 20-304, ovate or lanceolate, short- acuminate, borne on both the main filament and the branches. Attached to Zonaria tournefortit which was washed ashore. San Pedro, California. Also on Petrospongium berkleyi collected on rocks near San Pedro by A. J. McClatchie (1273). 15. Ectocarpus corticulatus, sp. nov. PLATE XX. Plant erect, 3 mm. to 3 cm. and more high, arising from a small, compact network of creeping filaments. Main filament go-120y broad, irregularly much branched below, above somewhat dichotomously branched ; branches 50-7ou wide, spreading, irregular, some long and bearing many short branches, others short and few celled, all ending abruptly; the main stem and the branches densely corticated ; cells one-half to as long as broad, but slightly constricted ; chromatophores few, large, band-form, each containing several pyrenoids; plurilocular sporangia very variable in size and shape, lanceolate or narrowly ovate, short stalked or sessile, mostly 30-40y x 12-18 Bot.—Vot. L.] SAUNDERS—PHYCOLOGICAL MEMOIRS. 153 (occasionally 50-704 x 20-301), most abundant on the upper side of the ultimate branches but found also along the main stem and primary branches, below apparently arising from the corticating filaments. Forming small, loose masses, 2 mm. to 5 or 6 cm. high, on Desmarestia ligulata, Monterey; on Zostera leaves with Z. granulosus. San Pedro, California. 16. Ectocarpus mitchelle Harv. PLATE XXI, FIGs. I AND 2. Ectocarpus mitchelle Harv., Ner. Bor.-Amer., I, 1851, 143- Ectocarpus indicus Sonv., Zoll. Verz. Ind. Arch. ges. pflanz., 3, 1854. Plant arising from long, creeping filaments 10-15 wide; cells 5-10 times as long as the diameter. Erect filaments unbranched below, bearing above many alternate branches which have numerous short, spreading branchlets ; branches and branchlets pointed or ending in a short hair; main filament 25-402 wide, 1-3 times as long as the diameter ; chromatophores numerous, small, disk-form ; plurilocular sporangia cylindrical, obtuse, 50-1oop x 18-35/, sessile, mostly secund on the upper side of the branches. Forming light olive-green tufts 1-2 cm. high on rocks at the low tide level. San Pedro, California. A careful study of the plant showed it to be identical with the figures and description of Z. indicus given by Askenasy in Ein. Austr. Meersalg., 8, 1894. The same plant was received from Prof. McClatchie named &. wmitchelle. From a careful comparison of the plates and descriptions of E. indicus and E. mitchelle, the two species seem to be identical. Mr. F. S. Collins has received the same plant from San Diego, where it has been called &. vzrescens Thuret. He has ‘‘no doubt that Harvey’s and Thuret’s species are the same.’’ 17. Ectocarpus siliculosus parvus, var. nov. PLATE XXII. Plants tufted, erect, 1-2 cm. high; primary branches numerous, alternate, gradually contracted above into long hyaline hairs, bearing many short, pointed branches. Cells with the main filament 21-30 broad, 1-3 times as long, slightly constricted at the joints ; plurilocular sporangia narrowly lance- olate or conical, 120-290 x 20-27/, pointed and often tipped with a long hair, borne on a long or short stalk, abundant on the main filament and the 154 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. primary branches; unilocular sporangia ovate or elliptical, usually sessile, 35-55" X 20-274, sometimes on the same filament with the plurilocular sporangia. Forming a yellow fleece on sand-colored rocks. San Pedro, California, 1895. Collected by W. A. Setchell (T2713): The Pacific plant differs from the type and from the form Azemadlis in the smaller size of the vegetative parts and the plurilocular sporangia; from the form arctus (Kuetz.) Kuck. in the much narrower vegetative filaments and larger plurilocular sporangia. 18. Ectocarpus confervoides (/toth) Le Fol. Ectocarpus confervoides (RoTH) LE Jou., List. Alg. Mar. Cherb., 75, 1880. Plants 2 cm. to 3 or 4 dem. long, attached to the substratum by horizontal creeping filaments, often entangled at the base; branches alternate, gradu- ally tapering, often corticated, cells of the larger branches 35-50 broad, as long to one-half as long as the diameter; plurilocular sporangia narrowly lanceolate and subulate to ovate and acute, sessile to short or long stalked ; unilocular sporangia ovate, globose, or elliptical. Of this variable, cosmopolitan species two forms have been collected on the Pacific coast. 19. Ectocarpus confervoides pygmaeus (Aresch.) Kjellm. PLATE XV, FIGs. 5-9. Ectocarpus confervoides pygmaeus (ARESCH.) KJELLM., Handbk. I, 18go, 76. Erect filaments arising from a compact substratum of. short creeping fila- ments, unbranched or bearing above a few divergent, long branches, in fig- ures 12-254 in diameter, not at all narrowed above; cells at base 2-3 times as long as the diameter, above once to one-half as long; chromatophores large, irregular, band-form, few in each cell; plurilocular sporangia lateral and terminal, sessile or short stalked, lanceolate or conical, rather obtuse at the apex, 60-Iooy x 20-304; unilocular sporangia lateral, abundant, ovate or globose, 25-40y broad, often two to many seriate. Forming an olive-brown, velvety covering on Desmarestza ligulata and Dictyoneuron Californicum. Pacific Grove, California. Bot.—Vot. 1.] SAUNDERS—PHYCOLOGICAL MEMOIRS. 155 20. Ectocarpus confervoides variabilis, form. nov. PLATE XXIII. Plant 2-8 mm. high, sparingly branched, branches alternate, distant, long drawn out at the tip, cells 10-30% broad, 1-3 times as long as the diameter, not at all constricted ; chromatophores large, band-form, few in each cell; plurilocular sporangia abundant, lanceolate, fusiform or narrowly ovate, usu- ally on a one- to few-celled stalk, borne laterally on the main stem or branches, ‘occasionally arising by a short stalk from the creeping filaments; 75-90 x 16-35; unilocular sporangia globose or ovate, 15-35 wide, single or 2-15 seriate, borne on a one- to three-celled stalk, mostly on the same filament with the plurilocular sporangia but much less common. Forming light olive-brown tufts on Lamznaria andersoniz, Iridealaminarioides, Macrocystis pyrifera, Egregia men- ztestt, Vemalion multifidum, Alaria esculenta, and Lessonia littoralis. A very common species and one that varies considerably in the size of the vegetative filament and the plurilocular sporangia. 21. Ectocarpus penicillatus C. Ag. PLATE XXI, FIGS. 3 AND 4. Ectocarpus penicillatus C. AG., Syst. Alg., 1824, 162. Plant densely tufted, dark olive-green, 1-4 cm. high; main stem and few of the primary branches corticated; cells of the primary branches 20-35 broad, a little shorter than the diameter; secondary branches somewhat secund, short and very numerous; plurilocular sporangia linear or linear- lanceolate, sessile pedicillate, acute 85-1654 x 12-204, borne mostly on the secondary branches. A dark, olive-green plant forming dense tufts on Desma- restia ligulata. Pacific Grove, California. 22. Ectocarpus tomentosus (//uds.) Lyngéd. PLATE XXIV, FIGS. I AND 2. Ectocarpus tomentosus (Hups.) Lvncs., Hydrophyt. Dan., 1819, 132. Plants forming tufts 1-4 cm. long, of densely interwoven, irregularly, much branched filaments forming a rope-like spungy mass. Filaments 6-8 wide ; cells 2-4 times as long as the diameter; branches long, divergent, the ulti- mate ones standing at right angles, curved at the tip; chromatophores large, irregular, band-form, few; plurilocular sporangia linear-oblong, 4o-11ou x 12-18, given out at right angles to the filament, sessile or occasionally short- pedicillate, often curved; ‘‘' sporangia almost elliptical, on short pedicels.” eee ee ee ———— 1 Hauck in Rab. Crypt. Fl. Vol., IT, 330. ° 156 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Common on Fucus evanescens and Fucus harveyanus. Monterey, California. One of the commonest Pacific coast species; where it occurs in large quantities it appears to be very destructive to the ‘‘host’’ plant. The tufts average much smaller than those of the Atlantic coast specimens. 23. Ectocarpus granulosus (Angi. Bot.) Ag. PLATE XXIV, Fics. 3-5. Ectocarpus granulosus (ENGL. Bot.), AG., Sp. Alg. II, 1828. 45. Filaments 1-8 cm. high; main branches mostly opposite, many of them corticated, cells 40-75 broad, one-half to as long as the diameter, secondary branches opposite, short, given off at wide angles, curved at the tips; ulti- mate branches secund, short and acute; plurilocular sporangia abundant, sessile on the secondary and ultimate branches, broadly ovate, obliquely truncate at the base, 60-100 x 30 to 60%; unilocular sporangia wanting. On rocks, Costaria mertensi’, and Nereocystis liitkeanus, Pacific Grove, California; and on Zostera marina, San Pedro, California. The Californian plant agrees with Dr. Farlow’s variety tenuzs in the size and branching of the vegetative filaments, but has the plurilocular sporangia of the type. 24. Pylaiella Bory. Pylaiella Bory, Dict. Class. IV, 1825, 393. Plant filiform, monosiphonous, more or less branched, attached to the sub- stratum by more or less branched creeping filaments. Unilocular sporangia globose, seriate, arising from the transformation of a part of the vegetative cells of any of the branches, opening laterally ; plurilocular sporangia oblong or cylindrical, intercalary, opening laterally, or occasionally terminal and opening at the apex, single or many seriate. 25. Pylaiella littoralis densa, form. nov. PLATE XXV, FIGS. I AND 2, 5 AND 6. Vegetative filaments below densely intertwined into rope-like masses, peni- cillate above, branched. Branches numerous, long, opposite or alternate, bearing many short branchlets; main filament and branches 18-25, cells 1-3 times as long as the diameter, not at all constricted ; unilocular sporangia globose, 2-15 in a series, most abundant in the branches and branchlets, 21-304 in diameter; plurilocular sporangia scarce, mostly in the main fila- ment, 20-50% x 20-304, 2-Io or more in a chain. Bot.—Vou. 1] SAUNDERS—PHYCOLOGICAL MEMOIRS. ES The plant forms dense, ropey, dark olive-brown masses 3-8 cm. long on Fucus sp. Ilwaco, Washingion, near the mouth of the Columbia River. On the southern Californian coast material was collected of what seems to be another well marked variety of P. /itto- valis. As the material was sterile I forbear making any further additions to the long list of varieties and forms of this species. The description of the plant is given below:— 26. Pylaiella littoralis, var. PLATE XXV, FIGS. 3 AND 4. Plant composed of loosely intertwined, light olive-green filaments of indefi- nite length, bearing a few long attentuate branches 20-7oy broad, %-2 times as long as the diameter, not at all narrowed upwards. Main filament and the branches bearing many one- to few-celled branchlets which stand at right angles to the filament. Forming a light yellow flocculent mass of a few cm. long on Amphiroa. San Pedro, California, Aug., 1896. Il-—SPHACELARIACEZ AND ENCCELIACEA! OF THE PACIFIC COAST. Family SPHACELARIACEE (Decne.) Kuetz. Sphacelariacee (DECNE.) KueETz., Linn., XVII, 1843, 93. Plant body arising from a larger or smaller mass of cellular tissue, poly- siphonous, more or less branched, increasing by the division of a large ter- minal cell; reproductive organs (unilocular and a plurilocular sporangia) arising on a longer or shorter stalk directly from the basal tissue or borne laterally on the erect filaments. The family includes at present about eleven genera, only one of which has been collected on the California coast. 27. Sphacelaria Lyng. Sphacelaria LyncsB., Hydrophyt. Dan., 103, 1819. Plant olive-brown, tufted, filamentous, branching; basal plate fastened to rocks or in the tissue of other alge as a substratum; both the axis and I 58 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER: branches terminated by a large apical cell which by transverse, longitudinal, and oblique division produces a plant body, the external surface of which is composed of rectangular cells arranged in regular, transverse bands. Uni- locular and plurilocular sporangia mostly extra-axillary on the branches, on a one- to few-celled stalk; propagula known in most species, on separate plants. 28. Sphacelaria tribuloides J/enegh. Pate XXVI. Sphacelaria tribuloides MENEGH., Lett. al Corinaldi 2, N. 1; Alg. Ital., 336, 1842. Plant densely tufted, olive-brown; primary branches somewhat dichot- omous, appressed, bearing above numerous alternate or opposite branches. Main filament composed of 4-8 siphons, 50-70y wide, the external cells 1-% as long as the diameter of the filament; propagula obcordate on the upper branches ; plurilocular sporangia elliptical or obovate, 75-150 x 50-7ou, ona one- to four-celled stalk. The plant forms dense tufts 1-3 or 4 cm. high, on rocks at the low tide line. Collected by Prof. A. J. McClatchie. Sphacelaria didichotoma, sp. nov. PLATE XXVII. Plant forming small, cushion-like masses 2-4 mm. high. Erect filaments arising from a small compact substratum, bearing many long, spreading, alternate branches; main filament 25-35 broad; external cells % to as long as the diameter of the filament; axis of the filament of four or five siphons; propagula abundant, large, twice dichotomus, often standing at right angles to the main filament; stalk 2-300% long, main branches too-200p. long. Forming small, compact tufts on J/elobesca and Ahn- feldtia. Carmelo Bay, California. Family ENCCELIACE® (Kuetz.) Kjellm. Enceliaceé (KUETZ.) KJELLM., Phyc. Gener., 336, 1843. Reproductive organs developed from superficial cells or from one of the divisions of a superficial cell; plant tissue of parenchymatous structure ; growth intercalary, growing point near the base. Bot.—VoL. I.] SAUNDERS—PHYCOLOGICAL MEMOIRS. 159 SYNOPSIS OF GENERA. Plant body undifferentiated—of similar structure throughout. Plant thin, a flat, leaf-like membrane................... Homeostroma. Piant body differentiated—composed of at least two kinds of cells. Sporangia sunken beneath the surface, plant hollow, cylindrical, or club BARU APOCRINE Tanen sek cor Nis or eras ity citing oi oe Cas een ne ney a era Coilodesme Sporangia on the surface. Paraphyses none, plant a leaf-like membrane. Central layer of tissue composed of large, irregularly rounded (LST: RA ree Chase ney NURS eens ae nee gn eee ne ATE Phyltitis. Central layer of tissue a loose web of slender branching: fila- BRINE Netcast ln a read Waa nh ee ee A aR Endarachne. Central layer of tissue composed of a smaller, compact layer of cells reproducing by unilocular sporangia...... Flalorhipis. Paraphyses unicellular. Riant- poay hollow, cylindrical... 0.5.2 veut. oo Scytosiphon. Plant body hollow, irregulary rounded or oval..... Colpomentia. Paraphyses multicellular; plant body rounded or oval, plants mostly SY TE ADRES CL oe 812 2 ovahin oR ENS colin secd'e Wis Cate ie erat Soranthera. 30. Homeostroma 7. Ag. Homeostroma J. Ac., Anal. Alg. Cont., III, 1896, 3. Punctaria GRev., Syn. in Alg. Brit., XLII, 1830, (in parte.) Plant body membranaceous, of several (2-7) similar layers of cuboidal cells. Unilocular and plurilocular sporangia oblong or oval, formed from the outer layer of cells; paraphyses none, furnished with numerous clusters of hairs. 31. Homeostroma latifolium 7. Ag. PLATE XXX, FIGs. 4 AND 5. Homeostroma latifolium irAci ke Punctaria latifolia, Born. et Thur., Etud., phycol. p. 13. Plant body arising from a minute disk, lanceolate-obovate, 1-5 dcm. long, tapering below to a short stalk 2-10 mm. long, olive-green, plaited on the edges; tissue consisting of 4 or 5 rows of cells; unilocular and plurilocular sporangia scattered, on the same plant. Attached to leaves of Zostera mar7na washed ashore at Monterey, California. 32. Coilodesme Stram/f. Cotlodesme StrazmF., Eing. Meersalg. Isl., 173, 1886. Plant body hollow, cylindrical or oval, on a short, solid stalk which arises from a minute disk-like base composed of two (or three) layers of tissue, an inner layer of large, mostly colorless, cylindrical cells and an outer layer of smaller colored cells. Unilocular sporangia solitary, scattered, developed from the outer layer of cells but surrounded by the continued growth of the outer layer of tissue; plurilocular sporangia, hairs, and paraphyses wanting. 160 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 33. Coilodesme californica (/rupr.) Ajellm. PLATE XXIX, Fics. 1-3. Coilodesme californica (RupPR.) KJELLM. in Engl. and Prantl. Natiirl. Pflanz- enfam., Lief. 86, 202, 1893. Adenocystis californica Rupr., Tange. Ochot, 29. Plant olive-green, hollow, inflated, thin membranaceous or papery, cylindri- cal or ovoid, 2-6 dem. long, %-1 dem. wide, abruptly contracted below into a short, solid stalk 1-3 mm. long. Sporangia abundant, scattered over the whole surface of the plant, ovate, 25-304 x 12-15p. Abundant on the upper branches of Cystosecra osmund- acea. Pacific Grove and San Pedro, California. The plants appear in June as minute, smooth, air-tight sacks filled with a gelatinous liquid which is formed from the degeneration of the central tissue. They gradually increase in size, soon becoming wrinkled and folded and finally torn, especially at the ends. The walls are thin and delicate and are composed of two layers of poorly differen- tiated tissue. The single outer row of cells is rounded and though not well developed it is easily distinguished from the hypodermal layer. The hypodermal layer is composed of two or three layers of irregularly rounded or quadrangular cells; beneath this is the central layer composed of three or four layers of large cylindrical cells surrounded by many smaller ones. The hypodermal layer is supplied with one or two large oval chromatophores, and occasionally one is found in the central layer of tissue. 34. Halorhipis', gen. nov. PLATE XXVIII. Plant body solid, leaf-like, arising from a disk-like base, composed of two layers of tissue, the outer layer consisting of 2-3 rows of cuboidal cells, the inner layer of several rows of large, cylindrical cells; reproducing by unilocu- lar sporangia collected in sori which are distributed over the whole surface of the plant; hairs numerous, forming the center of the sori; plurilocular sporangia and paraphyses wanting. 1 GAs—=sea, pixts—fan. Bot.—Vot.1.] SAUNDERS—PHYCOLOGICAL MEMOIRS. 161 35. Halorhipis winstonii (Azds.) PLATE XXVIII. Punctaria winstonit ANDS., Some New and Old Algze, Zoe IV, 1896, 358. Plant membranaceous, olive-brown, lanceolate, obovate, or spatulate, 8 cm.—2 dem. long, 2-5 cm. broad, usually frayed and torn at the end, gradu- ally narrowed below into a short stalk (2-8 mm. long). Unilocular sporangia elliptical, obovate, or pyriform, 30-45 x 20-304, sori numerous, forming irregular, linear patches of various sizes. On Eegregia menztesiz and on rocks at low tide level. Carmelo Bay, California. In 1895 a study was made of material received from Dr. Anderson, and the distinctness of this plant from Punctaria was at once noticed. During the summer of 1896 several trips were made to Carmelo Bay, fresh material in all stages of growth was obtained from the same locality where the type specimens were collected by Mr. Winston and sent to Dr. Anderson. 36. Phyllitis Awetz. Phyllitis Kuetz., Phyc. Gener., 342, 1843. Plant body solid, membranaceous, lanceolate or linear, tapering toward the base to a short round stem, of two layers of tissue, the cortical layer of small cuboidal cells, the inner layer of oblong colorless cells. Unilocular sporangia linear, covering the entire surface of the plant; plurilocular spo- rangia and paraphyses wanting. 37. Phylilitis fascia (Muell.) Kuetz. PLATE XXX, Fics. 1-3. Phyllitis fascia (MUELL.) KUETz., I. c. Fucus fascia OEDER, Fl. Dan. Tav., 768, 1761. Plant single or clustered, arising from a disk-like base, 2-30 cm. high, 1-5 cm. broad, linear, lanceolate, or elliptical, narrowed at base into a short stem, or sessile. Two forms of this variable species have been collected on the Californian coast: A.—Plants gregarious, linear or oblanceolate, 3-15 cm. long, 3 mm. to I cm. wide, gradually narrowed to a short stalk. On rocks and Phyllospadix sp. Monterey and Pacific Grove, California. 162 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. £.—Plants single or somewhat clustered, broadly lanceolate, plaited on the margin, 5-9 cm. long, 1-2 cm. broad, basal stalk very short or none. San Pedro and Pacific Grove, California. In external appearance this form is almost undistinguish- able from Hudarachne. 38. Endarachne 7. Ag. Endarachne J. Ac., Anal. Alg. Cont., III, 1896, 26. Plant plain, simple, without a rib, composed of three layers of tissue; the axillary tissue of slender, intertwined, articulate filaments, on each side of this a single row of slightly coherent hypodermal cells, outside of which there is a double layer of small cells, slightly elongated in a longitud- inal direction; reproducing by plurilocular sporangia scattered over the whole surface of the plant. 39. Endarachne binghamie 7. Ag. PLATE XXX, Fics. 6 AND 7. Endarachne binghamie J. AG., |. c., 27- Plant clustered, smooth, olive-brown, 5 cm. to a dem. or so high, 1-2 cm. broad, obtuse above, tapering below to a very short stalk. A single dried specimen of this plant, collected at San Pedro, California, was received from Prof. McClatchie about the time of the publication of Agardh’s description; figure 7 was drawn before the publication was received. In external appearance the plant is almost undistinguishable from the broader form of Phy//itis fascia; it is, perhaps, a little thinner and more abruptly contracted at the base. Agardh places it between Scytostphon and Phyliitis. It has been suggested that perhaps all that has been called Phy/- itis on this coast is really Eudarachne. A study of many specimens from Monterey and San Pedro-seems to show that Phylitzs is quite abundant and Hxdarachne is found, so far at least, only on the Southern coast. 40. Scytosiphon Ag. Scytosiphon Ac., Sp. Alg., I, 1823, 160. Plant body filiform when young, tubular when mature, composed of two layers of tissue, the outer of small quadrangular cells, the inner layer of thick- walled, vertically elongated, colorless cells. Plurilocular sporangia developed from the cortical layer of cells, covering the whole surface of the plant; para- physes single celled, oblong-ovate, sometimes wanting. Bot.—Vot. I.] SAUNDERS—PHYCOLOGICAL MEMOIRS. 163 41. Scytosiphon lomentarius (Zyngt.) F. Ag. PLATE XXXI, Fics. 8-10. Scytosiphon lomentarius (LYNGB.), J. AG., |. c., 126. Chorda lomentaria Lyncs., Hydrophyt. Dan., 74, 1819. Plant body unbranched, tubular, arising from short (1 cm. long) filiform stalks, 1-4 dm. long, 1-10 mm. thick; constrictions regular and frequent, occurring at long intervals or entirely wanting. On rocks, Pacific Grove, Monterey, Santa Cruz, San Pedro and San Diego, California (Averill’s set). One form found mostly below the low tide line is broad and short and the constrictions are regular and frequent ; another form occurring on overhanging rocks at or above the high tide line approaches the variety comp/anatus of Rosenvinge. It is longer and more slender than the pre- ceding form and the constrictions are seldom present. Between these two there is such an imperceptible gradation ’ that it is difficult to draw a line of separation. 42. Scytosiphon bullosus, sp. nov. PrATE XXXII, Fics. 1-7: Plant erect, membranaceous, hollow, dark olive-green, cylindrical to broadly ovate, 1-5 cm. high, 1-2 cm. broad, simple or lobed above, nar- rowed at base into a broad disk-like attachment; surface at first smooth, wrinkled, and often torn with age. Plurilocular sporangia, paraphyses, and hairs undistinguishable from those of S. /omentarius. On rocks, exposed at low tide with Leathesza and Ulva. Pacific Grove, California, Aug., 1896. A very variable species but apparently quite distinct from S. lomentarius. 43. Colpomenia Derd. & Sol. Colpomenia Ders. & SOL., Phys. Alg., p. 11, 1856. Plant body globose or oval, hollow, the walls entire or irregularly torn, composed of two layers of tissue ; the inner layer consisting of a few large, rounded, colorless cells, the outer of small quadrangular colored cells. Pluri- locylar sporangia at first forming sori around the hairs, soon spreading over the whole surface of the plant, interspersed with unilocular, clavate, paraphyses. , [2] Oct. 31, 1898. 164 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 44. Colpomenia sinuosa (Roth) Derb. & Soi, PLATE XXXII, Fics. 7 AND 8. Colpomenia sinuosa (ROTH) DERB. & SOL., 1. c. Ulva sinuosa Rotu, Cat. Bot., III, 327, Tab. 12, 1806. Asperococcus sinuosus Bory, of Anderson’s list, etc. Plant body sessile, thin, membranaceous, 4 cm. to a dem. high, sunken and occasionally irregularly torn in the older forms, tissue .255 mm. in thickness, the outer row of cells consisting of one or two rows of cuboidal, colored cells, inner layer of two rows of large roundish cells; plurilocular sporangia 20 x 7p, often containing the zoospores cells in two rows; paraphyses a little shorter and broader, 17 x 1op. The plant is usually found attached to other seaweeds and seems to prefer quiet coves. Carmelo Bay, Monterey Bay, and San Pedro, California. 45. Colpomenia sinuosa expansa, form. nov. PLATE XXXII, Fics. 4-6. Plants aggregated, forming an indefinite expanded mass on rocks; plant tissue .425 mm. in thickness, the inner layer of tissue being composed of 5-7 rows of cells. Santa Catalina Island near Avalon Bay, California. 46. Colpomenia tuberculata, sp. nov. PLATE XXXII, Fics. 1-3. Plant coriaceous, olive-brown, sessile, hollow, hemispherical, 5 cm. to a dem. or more in diameter; surface deeply convoluted, wrinkled, and folded, the whole surface covered in the mature plant with blunt tubercles 1 mm.-1o mm. high, 1-5 mm. broad. Plurilocular sporangia 22-254 x 3-44, composed of 6-8 zoosporal cells in a single row; paraphyses 22 x 5, remaining after the zoospores have escaped. Near San Pedro, California. The plant forms large rounded brain-like masses attached to rocks by the whole under surface; the outer layer of tissue .65—.80 mm. in thickness, composed of 3-5 layers of cuboidal cells, the inner 5-8 rows of large irregular cells. The plant tissue is of firmer texture and much thicker than in C. sinuosa. The structure of the tissue is very similar to that of Hydroclathrus cancellatus. Bot.—Vo.. I.] SAUNDERS—PHVCOLOGICAL MEMOIRS. 165 47. Soranthera Post. & Rupr. Soranthera Post. & Rupr., Illustr. Alg., 19, 1840. Plant body hollow, inflated, composed of two layers of tissue, with a corti- cal layer of small, colorless cells, the inner of large, nearly colorless cells. Unilocular sporangia forming sori which are distributed over the entire sur- face of the plant; paraphyses unicellular, hairs abundant in the center of each sori; plurilocular sporangia unknown. 48. Soranthera ulvoidea Post. & Rugr. PLATE XXIX, FIGs. 4 AND 5. Soranthera ulvoidea Post. & Rupr., l. c. Plants gregarious, sessile, membranaceous, olive-green, globose or oval, 2-8 or 1ocm. high. Sori very abundant, evenly and closely distributed over the whole surface of the plant. Unilocular sporangia clavate, 70-1o00y long, surrounded by numerous linear paraphyses which are nearly twice as long. In sheltered coves, usually on /thodomela larix, occa- sionally on rocks and other alge. Monterey, California. Figs. 1-7. Figs. 8-12. Figs. 1-4. Figs. 5-9. Figs. 1-3. Figs. 4-7. EXPLANATION OF PLATES. ae PAGE PLATE XII. Phycocelis fecunda STRO@MF. 148 Figs. 1-3 on Macrocystis pyrifera. 1 nat. size; 2 and 3 x 450. Figs. 4-7 on Desmarestia ligulata, 6 nat. size; 4, § and 7 x 450. Phycocelis reptans (CROUAN) KJELLM. - TAS ; Figs. 8 and 9 nat. size; 10 x 80; 11 and 12 x 450. PLATE XIilL Streblonema fasciculatum (THUR.) LE JOL. 148 ; Figs. x 450. PLATE XIV. Ectocarpus acuminatus, sp. nov. 149 Figs. 1 and 2 x 80; 3-5 x 450. Ectocarpus ellipticus, sp. nov. Fig. 6 nat. size; 7 x 80; 8 and 9 x 450. 149 PLATE XV. Ectocarpus chitonicolus, sp. nov. 150 Fig. 1 x 60; 2-4 x 450. Ectocarpus confervoides pygma@us (ARESCH.) KJELLM. 154 Fig. 5 nat. size; 6 x 60; 7-9 x 450. PLATE XVI- Ectocarpus cylindricus, sp. nov. 150 Figs. 1 and 6 x 80; the others by 450. PLATE XVII. Ectocarpus hemisphericus, sp. nov. I5I Fig. 1 nat. size; 2x 65, 3-8 x 450. PLATE XVIII. Ectocarpus hemisphericus minor, form. nov. 5a Fig. 1 x 65; 2 and 3 x 450. Ectocarpus paradoxus pacificus, var. NOV. — 152 Fig. 4 nat. size; 5 x 80; 6 and 7 x 450. BoT.—VOL. I.] Figs. Figs. gs. 1, 2, 5, 6 Pylatella littoralis densa, form. nov. PEATE XIX, Ectocarpus mucronatus, sp. nov. Fig. 1 x 80; 2-4 x 450. PLATE 2X. Ectocarpus corticulatus, sp. nov. Pig. 1x6; 2x 80; 3=5 x 450. PLATE XXI. 1,2. L&ctocarpus mitchelle Harv. Fig.1 x 65; 2 x 450. 3, 4. Lctocarpus penicillatus C. Ac. Fig. 3 x 90; 4 Xx 450. PEALE XE Ectocarpus siliculosus parvus, var. nov. Fig. I x 65; 2-8 x 450. PEATE XXIIL- Ectocarpus confervoides variabilis, form. nov. Figs. 1 and 4 x 80; 2, 3 and 5 x 450; 6 one-half nat. size; 7 x 60; 8-10 x 450. PEATE AX IV. . 1, 2. Lctocarpus tomentosus (Hups.) LyNGcs. Fig. 1 nat. size; 2 x 450. . 3-5. LEctocarpus granulosus (ENGL. Bort.) AG. Fig. 3x 6; 4.x 90; 5 x 450. PLATE XXV. Fig. 1 x 80; 2x 6; 5 and 6 x 450. gs. 3,4. Pylatella littoralis, var. PLATE: XXVI. Sphacelaria tribuloides MENEGH. Fig. 1 x 60; 2X 4; 3-9 X 450. SAUNDERS—PHYCOLOGICAL MEMOIRS. 167 PAGE 152 152 153 155 153 155 155 156 155 A 0 en An é 168 Figs. Figs. Figs. Figs. Figs. Figs. Figs. Figs. Figs. Figs. I-3. 4-5- I-7. 8-10. ae aS rt y Mie tea ee CALIFORNIA ACADEMY OF SCIENCES. (Proc. 3D | ; d bet PLATE XXVII. ae PAGE Sphacelaria didichotoma, sp. nov. 158. Figs. 1 and 2 x 50; 3-6 x 450. PLATE XXVIII. Halorhipis winstonii (ANDS.), gen. nov. 160 © Fig. 1x 1%; 2and 5 x 65; 3, 4 and 6x 450. PLATE XXIX. . Coilodesme californica (RUPR.) KJELLM. 160 Fig. 1 nat. size; 2 cross-section, 3 longitudinal section x 450. Soranthera ulvoidea Post. G& Rupr. 165 Fig. 4, a, 6, and ¢ nat. size; 5 x 250. PLATE XXX. Phyllitis fascia (MUELL.) KUETZ. 161 Figs. 1 and 2 nat. size; 3 x 450- Hlomeostroma latifolium |. AG. 159 Fig. 4 nat. size; 5 x 450. Endarachne binghamie J. AG. 162 Fig. 6 nat. size; 7 x 450. PLATE XXxXI. Scytosiphon bullosus, sp. nov. 163 Figs. 1-6 one-half nat. size; 7 x 450- Scytosiphon lomentarius (LYNGB.) J. AG- 163 Figs. 8 and 9 three-fourths nat. size; 10 x 450 PLATE XXXII. Colpomenia tuberculata, sp. nov. 164 Colpomentia sinuosa eapansa, form. nov. 164 Fig. 4 nat. size; 5 x 65; 6 x 450. Colpomenia sinuosa (ROTH) DERB. @ SOL. 164 Fig. 7, a, 6, c nat. size ; 8 x 450. ke [SAUNDERS | PLATE XII Proc. CanAcanSer.i3" cer Bor Vou fee SS A Sea WE Cen LITH BRITTON &AEY, SF: IBD, a LLM. ae | a L4 4 ra ay] LY./ f= HEY om Fine AY 2 5 /LAYUA =k =P TAN BE dha Ee a FHYCOCELI [SAUNDERS | Pare XID ELE, GEL, Ne LITH.BRITTON & REY, 5 F. STREBLUNEMA FAS XCULATUM [THUR] LE Sab, yi [SAUNDERS | PLATE XIV Proc CanAcanScr325er Bor Vol. ’ LIEBRITTON &REY, 57. ELH. MEL. SP NOV. 5P NOV. (au ra ECTOCARFU Fies.1 - 45. ie ACUMINATUS, ECTOCARPUS ELLIPTICUS, as.b - 4. I ““ : e Bis re t At Par [SAUNDERS | PLATE XV. LOU BRATON GREY, 5F: s fy 2 aS es Ly = a, x Opal ee ~ t és ms 4 Be et oO Ss Ee c > aap Ae r= ess GR tp eed S| Sarat cn sa ait my fd aes thy t 2 alee = eee 4 2 on =a] Fe SoM Nor seen alle BISe pee has plleebewees eee O° oko BD OPN N G5 ONO I IK 2000 = x ES os ee LUE DRIITON & REY, SF, | LED. r ye oF FORM. NOV. R, MINO PACIFICUS, VAR. NOV. 5 ECTOCARPOUS HEMISPHERIC NG eee C wu i} x FPARADUX ECTOCARPUS Fries. 4 - 7 wa Proc Cavacan Scr 32Ser. Bor Vail. LITE BHUTTO & REY, SF, ELE. Het, ECTtocARPUS MUCRONATUS, SP NOV. ‘6 Proc CarAcan Scr. 3° Ser. Bor. Vovl. | uate XX. oJ |[SAUNDER LITHERITITON &REY, 5 F ELA DEL ORTICULATUS, = i CTOCARPUS E Ae se F -© i » : neahy , I one a ar Proc. CanAcanScr.3° Sen Bor Vail [SAUNDERS] Prat XXL = \ i) ‘| \i i \ eS C LY EY” Ht ge a a ps I ee ih ELE TEL. LITH BRITTON &REY, SF Fies.1- 2. ECTOCARPOUS MITCHELLA AAR Fies.3-4. ElCTocARPUS PENICILLATUS Z_AB. [SAUNDERS | PLATE XXII. SEAT T Pea at as we ELH Ue. Pita? LEG, * ZITH BRITTON & REY, SF; EGTOCARPUS SILICULOSUS FARVUS, VAR. NOV. DAY a a J. * sn ~~ 5 Th, hea os ASS .* oP. . par w ate ON Fg Re . * bP AT Sh va ’ ‘ . . - “ « Sy Ls . ’ a ee ? ’ aie f : a ‘ wa ® ni - : ¢ Bi a, Rese hat Ys 2 ns a Ar Oe rn & z Proc [arAcan Scr 3*Ser Bor Vou) [SAUNDERS | PLATE XXIII. L 2 Zn CoE eh S58 he i ge SEES LER dy & qe ars ia : artes, § HR ORSON B ZEB Si LE Sa Ga, : my FO ES ey RS <>.0 SS 6S CEEEEAON a aa! CAA ey B%)GSV9 SEA i oe A 330 S8eSe AeA 03 BSeese2 Hey ey a OP ae Prot es ge a c Pam ra & D ay a io as o O EES cE — —— sé I Re Se > Ghia ETS, SE ELLIE see Lote aS BEERS we Sa. CN Sy ees I as Ge & Sy ee ee i SL CJ ee Bue torkcatir 3 Ser Bor Vo. [SAUNDERS | FLaTe XXXII Ba) LITH. BRITTON & REY, 5 ¥: OTT LET, JA TUBERCUOLATA, SP NOV. OLPOMED i yc dja Ait Fies.4- 6. CoLPoMBNIA stntmnsa EXPANSA , FORM. NOV. 5./- 8. COLPOMENIA sinuosa /Ag7/ Dens & Sar. 1G ae SOME OBSERVATIONS ON THE DEVELOP- MENT OF THE KARYOKINETIC SPINDLE IN THE POLLEN-MOTHER-CELLS OF COBZA SCANDENS CAV. BY ANSTRUTHER A. LAWSON. PLATES XXXIII-XXXVI. From the recent investigations of Strasburger (1897), Farmer (1895, 4), Swingle (1897), Harper (1897), Mottier (1898), and others, we learn that centrosomes are present and take an active part in the formation of the karyokinetic spindle in certain of the lower plants. The most complete series of stages yet published showing the behavior of centrosomes in plants is found in the papers of Swingle (1897) and Harper (1897). Swingle investigated the apical cells of Styfocaulon, Harper the developing asco- spores in the ascus of Hrys¢phe. In both of these cases, when the spindle is about to be developed, there is a body present which is surrounded by a system of kinoplasmic radiations. This body divides into two and the daughter- centrosomes thus formed migrate to opposite sides of the nucleus and form the spindle in much the same manner as in animals. Strasburger (1897, @) has described and fig- ured well defined centrosomes in the oégonium of Fucus, and Mottier (1898) has also found them in the tetraspo- rangia of Dictyota. In all of these cases it will be observed that the centro- some is described and figured as taking an active and very - essential part in the development of the spindle. We have thus a very striking parallel between the formation of the spindle in animals and in the lower plants; but when we attempt to extend this parallel to the higher plants we meet Sa Sr ek eo 1Contributions from the Botanical Laboratories of the University of California, No. 3. Presented for the degree of Master of Science. Prepared under the direction of Mr. W. J. V. Osterhout. [169] November 15, 1898. 170 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. with difficulties. In spite of the fact that Guignard (1891) and other writers have described centrosomes in the higher plants, recent investigations have rendered their existence very improbable. These investigations agree in showing that the spindle in its earlier stages possesses several poles, and later, by the fusion of these, becomes bipolar. In such a process of spindle-formation it is difficult to understand how a centrosome can have any part, nor has any observer, Guignard excepted, claimed to discover centrosomes in connection with it. Guignard (1898), however, in a recent article, while admitting that the spindle goes through a multipolar stage in the course of its development, maintains that this does not prove the non-existence of the centrosome. In the pollen-mother-cells of Vymphea and Muphar he describes centrosomes situated at the apices of the cones in the multipolar figures, and also at the poles of the mature spindle. But unfortunately he does not describe or figure a series of stages that would illustrate the behavior of these bodies. Moreover, Strasburger (1897, 6) has already dis- cussed this idea and concludes that it is highly improbable that a centrosome plays any part in this process. Although multipolar spindles were previously described by Belajeff (1894), Farmer (1893), and Strasburger (1896), their significance was not fully understood until Osterhout’s (1897) observations on Aguzsetum were made known. In this paper, which furnishes us with the most complete series of stages of spindle-formation yet worked out, the process is described as follows :— The first thing to be observed in the formation of the spindle in Aguzsetum is a felted zone of kinoplasmic fibres surrounding the nucleus. These fibres grow out from the nuclear wall and take on a radial arrangement. By the coming together of their free ends these threads form a series of cones. The nuclear wall now breaks down and the fibres composing the cones grow in and become attached to the linin and chromosomes. The apices of the cones now approach each other and arrange themselves in Bot.—VOL. I.] LAWSON—COBAAA SCANDENS. r7I two groups and then fuse to form a bipolar spindle. No bodies or granules that could be identified as centrosomes were observed in any stage of the process. It seems very improbable that the occurrence of multi- polar figures is an abnormal phenomenon. They have been found to occur in Leléum by Farmer (1893), in Larzx by Belajeff (1894), and Strasburger (1896). They have been found in Lilium, Fritillaria, Helleborus, Podophyllum, and Pinus by Mottier (1897, a and 6), in Hemerocallis by Juel (1897), in Chara by Debski (1897), and in Zamza by Webber (1897.) The writer has observed them in Hes- perla, Hedera, Disporum, Smilacina, Gladiolus, Jrts, Cobea, and other genera. From this we can only conclude that multipolar spindles are of very general occurrence, and this throws considerable doubt upon the existence of cen- trosomes in the higher plants. ; It must be observed, however, that in all cases where multipolar figures have been described, very little has been observed upon the very earliest stages in the development of the kinoplasmic fibres. It is with the hope of throwing some light on these stages that the following observations upon the pollen-mother-cells of Cobe@a scandens have been made. About two years ago Mr. Osterhout suggested that the writer undertake some investigations upon spindle-forma- tion in anthers of different orders of flowering plants. In undertaking such a task it is very desirable to obtain material with large anthers and large pollen-mother-cells as well as large nuclei. The anthers of Cobea are very good in all of these respects. The anthers were gathered and immediately fixed in the field. The following fixing fluids were used: Alcohol, 95 per cent. Chromic acid, 1 per cent. Flemming’s mixture, chromic-osmic-acetic, strong solution. Wilson’s sublimate-acetic. Boveri’s picro-acetic. Corrosive sublimate, saturated solution in 95 per cent. alcohol. 172 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. The best results were obtained by using Flemming’s strong solution diluted with one volume of water. The material was washed in running water from six to eight hours. It was then carried through different grades of alcohol by means of an apparatus consisting of a tumbler with a cover and a glass funnel. The funnel was of such a size as to be supported in the mouth of the tumbler. A piece of parchment paper was folded and placed in the fun- nel in the same manner as for filtering. Io per cent. solution of alcohol was placed in the tumbler and the anthers were placed in water in the funnel. The mouth of the tumbler was covered and the material was thus allowed to remain for an hour or two. The alcohol in the tumbler was changed at intervals to 25 per cent., 50 per cent. and 95 per cent. solutions. By this means the effect of the rapid change from a weak to a strong solu- tion of alcohol was obviated. The anthers were then thoroughly dehydrated in absolute alcohol. They were then placed in a mixture of bergamot oil and alcohol and then in pure bergamot oil. From the bergamot oil they were transferred to a mixture of bergamot oil and paraffin, and from this to pure paraffin, where they remained at a temperature of 55°C. for twenty-four hours. Microtome sections of 3% in thickness were used. Many stains were tried; especially iron-hematoxylin and Bordeau red, ruthenium red and thionin, etc., but the best results were obtained from Flemming’s triple stain, safranin, gentian violet, and orange G. In the resting condition of the pollen-mother-cell the nucleus is quite large. It contains one or two large nucleoli which stain very readily with safranin and some- times appear to be vacuolated. The chromatin, which is in the skein stage, stains blue with the gentian violet. ‘As soon as the chromatin breaks up and forms the. chromo- somes it stains red with safranin. The chromosomes appear as small oval bodies which are invariably situated in contact with the nuclear wall. The largest number that was observed in polar view was twelve. On the chromosomes Bot.—VOL. I.] LAWSON—COBAA SCANDENS. 173 the writer has made very few observations. They will therefore not be discussed farther. The linin now appears in the form of a lumpy or granular thread; it stains blue and is invariably connected with the chromosomes. The cytoplasm now appears in the form of a clear retic- ulum, as shown in fig. 1. The meshes of this network, which can be traced from the nuclear wall to the cell-wall, appear to be smaller and radially elongated in the imme- diate neighborhood of the nucleus; but as one follows them outwards they increase in size and are comparatively large towards the cell-wall. Scattered irregularly through the cytoplasm are numerous small spherical bodies. These bodies have the appearance of oil-globules in the living cell, but after the cell has been killed in Flemming’s fixing fluid they appear quite black. What the chemical nature or function of these bodies is, the writer is unable to state. The cytoplasm does ‘not maintain its clear, uniform appearance for a very long period, but soon undergoes a remarkable differentiation. This differentiation is shown in figs. 2-5. It is brought about by the gradual accumula- tion of a granular substance which forms a complete zone about the nucleus. The minute structure of this substance is difficult to make out, but it appears to consist of granules which vary in size. While some are very minute, others are comparatively large. They appear to be arranged in such a manner as to give the impression of a foam-structure. This granular.zone is so constant in Cobea and in several other genera observed by the writer, that for con- venience it will be called perikaryoplasm. While it is accumulating, the black bodies, which were previously scattered irregularly through the cytoplasm, take up a definite position. ‘They arrange themselves along the outer margin of the perikaryoplasm, and in section present the appearance of a complete ring, as shown in fig. 5. It is at this stage that the contrast between the outer cytoplasmic reticulum and the perikaryoplasm is greatest. The threads of the outer reticulum now appear to have no 174 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. connection with the nucleus. They can no longer be traced through the perikaryoplasm. Where the outer reticulum abuts on the perikaryoplasm the meshes of the former are so compressed as to give the impression of a loose membrane. These two constituents of the cytoplasm not only differ from one another as regards their structure but their staining properties stand in great contrast. While the outer reticu- lum stains a light or gray-blue with the gentian violet, the perikaryoplasm stains a decided orange with the orange G. The entire cytoplasm thus presents the appearance of two distinct and sharply differentiated zones, differing from one another as regards structure and staining properties. These zones are seen in fig. 5. In case it should be thought that this is due to artifact, it might be well to state here that a large number of living cells were examined, and this striking differentiation of the cytoplasm and the ring of spherical bodies, together with the threads of the outer reticulum, could be readily seen. This, in the writer’s opinion, is sufficient evidence to prove that the phenomenon is a normal one. There seems little doubt that it is the first step towards the formation of the spindle. In fig. 6 we see the first indication of the breaking down of the nuclear wall. It evidently does not break down all at once. It commences to do so at one or more points where it becomes lost in a network which is now developed out of the linin and perikaryoplasm. Figs. 6-8 show stages in the breaking down of the nuclear membrane. It will be seen from these figures that the perikaryoplasm, where it is in the immediate vicinity of the breaks in the nuclear wall, is now undergoing a change. The linin has lost its lumpy appearance and now appears as fine, delicate threads (fig. 6) which are in direct communication with the perikaryoplasm. The latter is losing its granular nature in these places and is taking on the form of a net- work. In fig. 8 we see this network taking on a very definite form. It no longer stains orange, like the Bot.—VOoL. I.] LAWSON—COBAZA SCANDENS, 175 perikaryoplasm from which it develops, but stains deeper and deeper violet like the linin. At the points where the threads intersect granules are seen. In fig. 8 the nuclear wall is no longer continuous but is in the form of fragments. To what extent the linin enters into the formation of the network, the writer is at present unable to say, but it cer- tainly appears to begin its formation. The amount of linin present when the nuclear wall breaks down is not sufficient to form the large central network that we see in fig. 9. This network increases in size and apparently grows at the expense of the perikaryoplasm. In fig. 9 the nuclear wall has entirely disappeared and the area once occupied by the nucleus is now filled by a perfect network which supports the chromosomes within its meshes. These latter bodies appear to be more frequently present in those places where the network is growing outward in the form of projections. The contents of the cell is now beautifully differentiated into three parts: the outer cytoplasmic reticulum which stains light blue, the perikaryoplasm which stains orange, and the central network which stains violet. Between these latter two there is a transition place where the threads of the central network become granular and stain less deeply violet, until finally they become orange and pass over into the granular substance of the perikaryoplasm, where they can no longer be followed. In fig. 9 we see that the central network has atethree places formed definite projections. How many of these are generally formed it is difficult to make out from sec- tions, but the number is nearly always more than two. In fig. 10 we see that the meshes of the network have become elongated in the directions in which the projections extended. In fig. 12 we see that the meshes have a drawn- out appearance, and we have the outline of a multipolar spindle. Figure 11 is probably a later stage than fig. 12, inasmuch as the meshes are drawn out to such an extent as to present the appearance of distinct fibres, and we now have the first spindle-fibres formed. In this figure it will 176 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. also be observed that the nucleolus still persists, which shows that it has not been used to form the central network. This is the latest stage in which the nucleolus was seen, but what eventually becomes of it was not observed. In the next figure we have a characteristic multipolar spindle. Here the fibres are straight and form cones which terminate in sharp points. We see extending from the apices of the cones fibres which have free ends. These fibres eventually become the mantle-fibres which are so characteristic of the mature spindle. The multipolar figure becomes bipolar by the fusion of the cones in much the same manner as it does in Hguzsetum, as shown in figs. 14 and 15, but during the process the mantle-fibres have developed to a considerable length. From the time of the breaking down of the nuclear wall the chromosomes have been in connection with the network from which are developed the spindle-fibres; so that this process differs materially from those cases where the spindle-fibres have been described as growing in and becoming attached to the chromosomes. As shown in figs. 15 and 16, the mature spindle termi- nates in sharp points; there are three kinds of fibres present: The contractile fibres, which are in connection with the chromosomes, are compound in nature, being made up of a number of fine fibres. The continuous fibres extend from pole to pole uninterrupted. The mantle-fibres are remark- able for their great length and early development; they consist of very delicate straight threads; those from one pole cross those from the opposite pole in the manner shown in figs. 15 and 16. It should be observed that although the mantle-fibres are of great length they never extend beyond the perikaryoplasm. After the mantle-fibres have reached their maximum development the contractile fibres commence to draw the chromosomes to the poles. This takes place in the usual way. It will be noticed, however, as shown in fig. 17, that during this process the mantle-fibres appear more divergent from their respective poles. When the chromosomes have BotT.—VOL. I.] LAWSON—COBA:A SCANDENS. O77 reached the poles the mantle-fibres immediately lose their straight appearance and hang loosely alongside of the remaining continuous fibres, as shown in fig. 18. They soon after disappear. Although the continuous fibres now curve out towards the cell-wall as shown in fig. 19, there is no cell-plate formed until after the second division. When the daughter-nuclei have surrounded themselves with a membrane the continuous fibres still persist, being attached to the daughter-nuclei as shown in fig. 19. In this figure we also see that the perikaryoplasm appears to be more plentiful at the ends of the cell where the nuclei lie. It surrounds the nuclei except where the continuous fibres are in connection with the nuclear membrane. The peri- karyoplasm gradually surrounds the nuclei and apparently cuts off the continuous fibres. As shown in fig. 20, they can no longer be traced to the nuclear wall, and the perikaryoplasm has formed a complete zone about each nucleus. The development of the spindle of the second division is identical with that of the first division, but on account of its small size, the stages in the process are much more difficult to work out. In fig. 21 we see the last traces of the continuous fibres, and each of the nuclei is surrounded by a granular zone. It will also be noticed that the black bodies have formed rings at the outer margin of the zones of perikaryoplasm, just as they did in the case of the first division. These zones are as well defined and differentiated from the rest of the cytoplasm in structure and staining properties as in the first division. In fig. 22 we see the two spindles of the second division lying at right angles to one another. In one the spindle is represented only by the cross-sections of its fibres. The other shows the entire spindle with the characteristic mantle- fibres, and the chromosomes on their way to the poles. This figure also illustrates the fact that the perikaryoplasm accommodates itself to the shape of the spindle. In one 178 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. view it appears round and in the other oblong in outline, corresponding to the position and shape of the spindle. In the next figure we see that the spindles are more nearly in the same plane. The mantle-fibres of the two spindles have reached over and have apparently united with one another. In fig. 24 we see three daughter-nuclei in the same plane. By means of the continuous fibres and mantle-fibres they are connected with one another. These fibres now occupy almost the entire cell-cavity. Cell- plates are now formed in the usual way. Swellings appear on the connecting fibres; these increase in size, and finally result in forming cell-walls which separate the daughter- cells from one another. During this stage there appears to be very little of the perikaryoplasm left and what is present is scattered irregularly through the cell. Whether the method of spindle-formation observed in Cobea is of general or exceptional occurrence must be left for future investigation to decide. From Belajeff’s (1894) description of Larzx it may be supposed that a similar pro- cess takes place here, but the origin of the network and the manner in which the spindle-fibres subsequently arise from it were not sufficiently investigated. The develop- ment of spindle-fibres out of a network, as shown in figs. 9-13, recalls Wilson’s (1895) description of the formation of astral rays and spindle-fibres in Toxopneustes. This method of spindle-formation differs decidedly in its earlier stages from that observed by Osterhout (1897) in E-quisetum, although both agree in the formation of a num- ber of poles which subsequently fuse to form the bipolar spindle. We have in Cobea a zone of granular substance, the perikaryoplasm, whose function it is to take part in the formation of a network from which the spindle-fibres are developed; no such zone, however, is present in Aguzse- tum. Similar zones have been figured by Belajeff (1894) (fig. 6) in Larzx and Mottier (1897, a) in Lilium, but they have not been described as taking any part in the formation of kinoplasmic fibres. Bot.—VOL. I.] LAWSON—COBAA SCANDENS, I 79 In all other cases where multipolar spindles have been observed the earlier stages have not been sufficiently inves- tigated to warrant any statement as to whether they follow the Aguzsetum type or that here described for Cobea. From the series of stages here figured the writer can only conclude that centrosomes or directive spheres can take no part in the formation of the spindle, and they con- firm the idea that has already been expressed that in the vascular plants the method of spindle-formation is entirely different from that which prevails in the lower plants and animals. In conclusion I wish to acknowledge my indebtedness to Mr. Osterhout for many valuable suggestions in the prep- aration of this paper. SUMMARY. The observations made upon the formation of the spindle in Cob@a may be briefly stated as follows: A granular substance gradually accumulates and forms a complete zone around the nucleus. This zone is designated perikaryoplasm. Upon the breaking down of the nuclear wall the linin of the nucleus and the perikaryoplasm form a network which occupies the central portion of the cell. This network grows out into several projections which become the cones in the multipolar figures. The spindle-fibres are formed by the elongation of the meshes of the network in the direction of the projections. The cones elongate and become sharply pointed. They fuse in two groups and form the bipolar spindle in the same manner as that observed by Osterhout in Hguzsetum. The mature spindle is characterized by the great length and crossing of the mantle-fibres. The spindle-formation of the second division is identical with that of the first division. No bodies that could be identified as centrosomes were found in any stage of the process. BOTANICAL LABORATORY, UNIVERSITY OF CALIFORNIA, April, 1898. 180 1894. 1897. 1893. 1895a. 18956. 1891. 1808. 1897. 1897. 1897a. 18976. 1898. 1897. 1896. 18974. 18976. 1897. 1895. 1897. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. BIBLIOGRAPHY. BELAJEFF, W. Zur Kenntniss der Karyokinese bei den Pflanzen. Flora, Bd. LXXIX, 1894, p. 430. Depeski, B. Beobachtungen iiber Kerntheilung bei Chara fragilis. Jahrb. f. wiss Bot., Bd. XXX., 1897, p. 227. Farmer, J. B. On nuclear division in the pollen-mother-cell of Lilium Martagon. Ann. of Bot., Vol. VII, 1893, p. 392. Ueber Kerntheilung in Lilium-Antheren besonders in Bezug auf die Centrosomenfrage. Flora, Bd. LXXX, 1895, p. 56. On spore-formation and nuclear division in the Hep- atice. Ann. of Bot., Vol. 1X, 1895, p. 469. GUIGNARD, L. Nouvelles études sur la fécondation. Amn. des Sct. Nat. Bot., Serie 7, Tome XIV, 1891, p. 163. Centrosomes in Plants. Bot. Gaz., Vol. XXV, 1898, p. 158. Harper, R. A. Kerntheilung und freie Zellbildung im Ascus. Jahrb. f. wiss. Bot., Bd. XXX, 1897, p. 249, JueLt, H. O. Die Kerntheilungen in den Pollenmutterzellen von Hemerocallis fulva und die bei denselben auftretenden Unregel- missigkeiten. Jahrb. /. wiss. Bot., Bd. XXX, 1897, p. 205. Mortrier, D. M. Ueber das Verhalten der Kerne bei der Entwickelung des Embryosacks und die Vorgange bei der Befruchtung. /ahrdb. f. wiss. Bot., Bd. XXXI, 1897, p. 125. Beitrage zur Kenntniss der Kerntheilung in den Pollen- mutterzellen einiger Dicotylen und Monocotylen. /ahrd. /. wiss. Bot., Bd. XXX, 1897, p. 169. . Das Centrosome bei Dictyota. Perichte der deutschen Bot. Gesell., Bd. XVI, 1898, Heft. 5. OstERHOuT, W. J. V. Ueber Entstehung der karyokinetischen Spindle bei Equisetum. /ahrd. f. wiss. Bot., Bd. XXX., 1897, p. 159. STRASBURGER, E. Karyokinetische Probleme. /ahrd. /. wiss. Bot., Bd. XXVIII, 1896, p. 151. Kerntheilung und Befruchtung bei Fucus. /ahré. /. wiss. Bot., Bd. XXX, 1897, p. 351. Ueber Cytoplasmastructuren, Kern- und Zelltheilung. /ahré. f. wiss. Bot., Bd. XXX, 1897, p. 375. Wesser, H. J. Peculiar structures occurring in the pollen-tube of Zamia. Sot. Gaz., Vol. XXIII, 1897, p. 453. WItson, E. B. Archoplasm, Centrosome, and Chromatin in the Sea- Urchin Egg. /ourn. of Morph., Vol. X1, 1895, p. 443. SWINGLE, W. T. Zur Kenntniss der Kern- und Zelltheilung bei den Sphacelariaceen. /ahrb. /. wiss. Bot., Bd. XXX, 1897, p. 297. 182 Fig. 1. Fig. 2. Fig. 3. Fig. 4. Fig. 5. CALIFORNIA ACADEMY OF SCIENCES. ([PrRoc. 3D SER. EXPLANATION OF PLATE XXXIII. Figures drawn with Abbe’s camera lucida. Zeiss, Apochromatic Immersion Obj. 12 mm. 1.30 Ap. Compensating Ocular No. 6. A young pollen-mother-cell. The cytoplasm is a clear network which stains a light or gray-blue. The meshes of the network appear to be smaller and radially elongated towards the nucleus. There are numerous spherical bodies scattered through the cytoplasm. They are stained black, probably by the fixing fluid. The nucleus is quite large and shows a large nucleolus and small oval chromosomes which lie immediately upon the nuclear wall. The linin, which stains deep blue, is of a lumpy or granular nature. This figure shows the commencement of a remarkable differentia- tion which takes place in the cytoplasm which immediately surrounds the nucleus. A zone of granular substance, the perikaryoplasm, forms around the nucleus, which differs decid- edly both in structure and staining qualities from the sur- rounding network. While the outer cytoplasmic reticulum stains a light or gray-blue, the granular substance stains orange. This figure shows a later stage with an increase in the quantity of perikaryoplasm about the nucleus, otherwise it does not differ essentially from fig. 2. Shows a still greater increase in the quantity of perikaryoplasm The about the nucleus. The black bodies are now beginning to form a ring at the outer margin of the granular zone. perikaryoplasm has now reached its maximum and stands out in the greatest contrast from the outer cytoplasm. The meshes of the outer reticulum which abut on the perikaryoplasm are small and apparently compressed, giving the impression of a loose membrane which separates the two zones from one another. It is impossible to trace any of the outer meshes into the granular zone. They appear to have no connection with the nuclear wall. The black bodies have now taken up their position at the outer margin of the perikaryoplasm and form a complete ring about it. Fig. 6. It will be noticed that in the preceding figures the linin is more or less of the nature of a granular thread. It now appears to have lost its granular structure and appears in the form of very fine threads. We now have the first indication of the breaking down of the nuclear wall, and at that part where the nuclear wall is broken, the perikaryoplasm is commencing to take on the form of a network and stains violet. a= BD) LITH-BRITTON & SA Lawson. Dez, COBEA SCANDENS Lav 184 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXXIV. Fig. 7. The nuclear wall has nearly all disappeared and the linin has now formed a network. The perikaryoplasm is being gradu- ally transformed into a network of threads which stain deep violet, in contrast to the orange of the surrounding peri- karyoplasm. Fig. 8. This stage is about the same as fig. 7, only the network appears to be of a more definite nature. Fig. 9. The nuclear wall has entirely disappeared and the area once occu- pied by the nucleus is now filled with a clear network of threads; where the threads cross one another granules are present. The entire contents of the cell is now beautifully differentiated into three parts: the outer cytoplasmic reticu- lum which stains light blue; perikaryoplasm which stains orange, and the central network which stains deep violet. The figure alSo shows that the central network has in three places grown out into projections. Fig. 10. This shows the projections to have grown considerably and the meshes of the central network are more elongated, as if pulled out. Fig. 11. In this figure the projections have grown out to such an extent and the meshes have become so elongated that they form dis- tinct fibres. The more definite the fibres, the more deeply vio- let they stain. The figure also shows the vacuolated nucleolus. Fig. 12. This stage is probably earlier than fig. 11, as there are no fibres developed as yet. It shows the central network grown out in three distinct projections, and we have the outline of a multi- polar spindle. 0 Se Gee > , a se A eae ee one . a MA: ieee i is Mat , redder i & REY ree It Tos ix is | isk i 186 Fig. Fig. Fig. Fig. Ay. a7. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXXV. Here we see a characteristic multipolar spindle, the fibres straight and definite, and the cones terminating in sharp points. We also see fibres with free ends projecting from the apices of the cones. These eventually become the mantle-fibres in the mature spindle. It will also be noticed that the chromosomes have taken up a more central position. This figure shows that certain of the poles have approached each other and we have the indication of a bipolar spindle. The mantle-fibres have also grown to a considerable length. This figure shows the mature bipolar spindle. The chromosomes lie in the equatorial plate. The mantle-fibres have developed to an extraordinary length; those from one pole cross those from the opposite pole. The mantle-fibres, although very long, do not extend beyond the perikaryoplasm. The contractile fibres, which are connected with the chromosomes, are plainly compound in nature. There are also continuous fibres to be seen which extend from pole to pole uninterrupted. This stage is a little later than that shown in fig. 15. The chro- mosomes have separated and have commenced to move towards the poles. The poles of the spindle terminate in sharp points. The mantle-fibres have reached their maximum development. By the contraction of the contractile fibres the chromosomes have been drawn toward the poles. The mantle-fibres now appear to be more divergent from their respective poles. The chromosomes have now reached the poles. The mantle- fibres have lost their straight appearance and hang loosely alongside of the remaining continuous fibres. 1a ab § e : ry “¢ ah: “@ ‘oe e = 6 ¢& ° & 3 @ a¢ € %@ 188 Fig. 19. Fig. 20. | ae Fig. 22. Fig. 23. Fig. 24. CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. EXPLANATION OF PLATE XXXVI. Here we see each of the daughter-nuclei surrounded by a membrane, with the chromatin stained blue, and in the form of a necklace. There is also seen a nucleolus in each daughter- nucleus. The continuous fibres still connect the nuclei with one another. The granular substance appears to surround the nuclei except where the continuous fibres are connected with the nuclear membrane. The perikaryoplasm now entirely surrounds each daughter-nucleus. The connecting fibres have curved out toward the cell-wall but can no longer be traced to the walls of the nuclei. The old spindle-fibres have disappeared and each of the daughter- nuclei is surrounded by a definite zone of perikaryoplasm. The black bodies form a ring at the outer margin of each zone. This figure shows the two spindles of the second division lying at right angles to one another. The one is only represented by the cross-sections of its fibres, the other shows the entire spindle with the chromosomes on their way to the poles and with the characteristic mantle-fibres. This figure also shows that the perikaryoplasm has accommodated itself to the shape of the spindles. In one view the zone is round and in the other oblong, corresponding to the size and shape of the spindle. Here the spindles of the second division are more nearly in the same plane than in the preceding figure. The chromosomes are at the poles. The mantle-fibres of the two spindles have reached over and united, thus connecting the nuclei with one another. Here we see three of the daughter-nuclei in the same plane, each surrounded by a nuclear wall. The chromatin is in the spi- reme stage. All the daughter-nuclei are connected by fibres which almost fill the entire cell-cavity. These fibres are curved out toward the cell-wall, and we see swellings on some of them. What little of the perikaryoplasm is left is scattered irregu- larly through the cell. >. A 2 ee, ee. | THE ORIGIN OF THE KARYOKINETIC SPINDLE IN PASSIFLORA CCERULEA LINN. BY CLARA L. WILLIAMS, M. S. PLatEs XXXVII-XL. Recent work on karyokinetic division has made it’ appear very probable that no centrosome is present in the higher plants. In view of this the origin of the spindle becomes a subject of especial interest. The study of karyokinesis in the pollen-mother-cells of Passéf#lora cerulea was taken up in the hope of shedding some light on this question. The origin of the spindle is the sole question considered in this investigation other matters receiving only casual attention. Among the fixing fluids experimented with in the prepar- ation of material were Wilson’s sublimate-acetic, Boveri’s picro-acetic, Flemming’s strong mixture, Flemming’s strong mixture diluted with an equal part of water, and 2 per cent. iridium chloride. When anthers fixed in Wilson’s, Bo- veri’s, and dilute Flemming’s solutions were crushed and examined in their respective fluids they showed such shrink- age that these fluids were not used in making preparations. Good results were obtained with iridium chloride, but no better than with Flemming’s strong mixture (undiluted), which was used for fixing the greater part of the prepara- tions studied. The anthers were left in the fixing fluid for twenty-four hours, and afterwards washed in running water for six hours. 1 Contributions from the Botanical Laboratories of the University of California, No. 4. Presented for the degree of Master of Science. Prepared under the direction of Mr. W.J. V. Osterhout. ‘ [189 | April 13, 1899. 190 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. They were then placed in the inner part of a dehydrator’ and barely covered with water. The outer part of the dehydrator was filled with 95 per cent. alcohol. After remaining in the dehydrator for twenty-four hours, the material was removed to 95 per cent. alcohol and left for six hours. It was then run up successively through a mixture of equal parts of 95 per cent. of alcohol and absolute alcohol, absolute alcohol, a mixture of equal parts of absolute alcohol and bergamot oil, bergamot oil, a mixture of equal parts of bergamot oil and paraffin (43°), and a mixture of paraffin (43°) and paraffin (52°). The anthers remained six hours in each. When in bergamot oil they were placed on the paraffin oven. .Paraffin (52°) was used for imbedding and the microtome sections were cut from 3 to 5 microns in thick- ness. The sections were fixed to the slide either with albu- men (the sections being first spread out on a layer of water over the albumen, after which the slides were placed upon the paraffin oven to dry) or with 80 per cent. alcohol according to Eisen’s method (Eisen 1897). The latter method is preferable as the sections may be stained almost immediately. After mounting, the sections were stained with the Flemming triple stain. They were first placed in safranin, where they remained twenty-four hours. On removal they were decolorized with 95 per cent. alco- hol until the stain was removed from everything except the nucleolus. They were then washed with water and placed in gentian violet, where they were allowed to remain from five to fifteen minutes. There was then poured over the slide in rapid succession, a concentrated solution of orange G, 95 per cent. alcohol, and absolute alcohol. Clove oil was then poured upon the slide and the progress of decolorization watched under the microscope. 1 The dehydrator was made in accordance with the suggestion of Prof. W. A. Setchell, as follows: A funnel of appropriate size is deprived of its neck and supported in the mouth of a tumbler or beaker. A piece of parchment paper folded in the same manner as for filtering is placed in the funnel. A cover fits over the top of the tumbler to pre- vent evaporation. Alcohol is placed in the tumbler; the material is placed in the funnel and covered with water. The rapidity of dehydration depends upon the quantity of water, the strength of the alcohol, and the thickness of the parchment paper, and can be kept under perfect control. Bot.—VOL. I.] WILLIAMS—PASSIFLORA CC2RULEA. I9!I When the sections were sufficiently decolorized the clove oil was removed by placing the slide in xylene for a few moments. The sections were finally mounted in Can- ada balsam. In the young pollen-mother-cell, when the chromatin is still in the spireme stage (fig. 1), the cytoplasm is composed of two distinct elements. One is fibrous and forms a net- work throughout the cell from the nuclear wall to the cell- wall. The other is granular and is uniformly distributed within and upon the meshes of the network. The threads of the reticulum are knotted and sinuous and take the violet stain. The granular part of the cytoplasm usually stains yellow or yellowish brown. About the time when the chromatin thread breaks up a change takes place in the cytoplasm. The meshes of the reticulum immediately sur- rounding the nucleus are drawn out parallel to the nuclear wall and form a sort of weft about it (fig. 2). This con- dition of the cytoplasm is of short duration. Soon the meshes of the cytoplasmic reticulum assume a new arrange- ment. They become drawn out at right angles to the nuclear wall and appear as if radiating from the nucleus (fig. 3). A little later, irregular, deeply staining strands appear in the cytoplasm (fig. 4). For the most part, they are radially arranged and extend toward the cell-wall; some of them reach it. In one or two instances similar stages show groups of fibers outside the nuclear wall. These groups are conical with their bases directed toward the nucleus; they resemble the cones of Hguzsetum (Osterhout 1897, fig. 4). Their subsequent fate could not be deter- mined; no trace of them was seen in later stages. The radial arrangement of the cytoplasmic reticulum spoken of above disappears after a short time, and the fibers of the cytoplasm again form an irregular network as in figs. 1 and 2, The meshes show no tendency to be drawn out in any one direction, although a few deeply staining strands radiate from the nucleus (figs. 6, 7 and 9). Changes in the nuclear wall resulting in its transformation into a meshwork now begin. The first evidences of such change 192 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. are the appearance of granules in the nuclear wall and an irregularity in its outline (fig. 6). Various stages in the transformation of the nuclear wall have been observed (figs. 7, 8, 9 and 10). As the figures show, this change con- tinues until finally no definite nuclear membrane remains; in its stead, there is a zone of deeply staining meshes, the fibers of which contain many granules (fig. 11). This meshwork connects the linin reticulum with the cytoplasmic reticulum, forming one continuous network from cell-wall to cell-wall. In the younger cells there are but few linin threads and these are delicate and stain but slightly. Later, however, they become coarse, exhibit thickenings and increase so that a linin reticulum entirely fills the nuclear space (figs. 8, 9, 10 and 11). The granular constituent of the cytoplasm, which was at first uniformly distributed, soon becomes more or less massed in irregular patches midway between the nuclear wall and the cell-wall (fig. 4), leaving areas comparatively free from granular matter near the cell-wall and near the nuclear wall. This massing of the granular matter con- tinues until a complete dense zone, the granular zone, is formed at the time when linin, nuclear wall, and cytoplas- mic reticulum form an unbroken network from cell-wall to cell-wall (figs. 11 and 12). The granular zone is constant from this time on until the anaphase, and is occasionally seen even after the daughter-nuclei are formed. Sucha zone has been figured by several observers. Juel (1897) shows it as a definite zone in multipolar and bipolar stages; Belajeff (1894) and Osterhout (1897) in the bipolar stage; and Mottier (1897, @ and 6) traces of it in prophase, in multipolar and bipolar stages. What the formation of the granular zone has to do with the origin of the spindle is uncertain. It appears to be simply an accumulation of the granular part of the cytoplasm in a definite area. Soon after the formation of the continuous reticulum fill- ing the cell, the fibers in the vicinity of the old nuclear wall cease to stain more deeply than the other parts of Bot.—Vot. I.] WILLIAMS—PASSITFLORA C@RULEA. 193 the reticulum. The entire network stains uniformly (fig. 12), although the fibers inside of the granular zone differ in appearance from those outside of it. They are delicate and clear cut, while those outside are much thicker, more granular and less fibrous in structure. This difference between the parts of the reticulum inside and outside of the granular zone is more marked in the later stages. The first indication of the formation of the spindle poles is a decided change in the portion of the reticulum sur- rounded by the granular zone. At various points this part of the reticulum is drawn out into cones, which become the poles of the multipolar spindle (fig. 13). Each cone is composed of a reticulum whose meshes become stretched in the direction of the long axis of the cone and finally become fibers, many of which end free. A little later all traces of a reticulum inside of the granular zone disappear. The whole space limited by the zone is filled with free fibers which cross each other in all directions. Many of them converge at their extremities to form the poles of the multipolar spindle (fig. 14). At first the poles point in all directions, but soon they show a tendency to assemble in two groups (fig. 15), which are probably destined to form the poles of the bipolar spindle. The chromosomes assume a more central position than they have had heretofore and are more crowded together. The poles of these two groups gradually fuse (figs. 16 and 17) until finally a sharp-pointed, bipolar spindle results (fig. 18). In the equatorial plate stage, the spindle is usually long, very narrow, sharply pointed, and without mantle-fibers. Sometimes, however, there are long mantle-fibers which extend from the poles and pass obliquely to the cell-wall, the fibers from each pole crossing those from the opposite pole in their course. Often the poles extend to the cell-wall. In the meta- phase, also, the poles may extend to the cell-wall; man- tle-fibers are more often seen in this stage than in the equatorial plate stage. At this time, too, the spindle is usually broader than in earlier stages and its outline is more nearly elliptical. Often, in this stage, the groups 194 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. of fibers which extend from pole to pole are not straight as if stretched, but are more or less wavy (fig. 19). This may be due to the drawing together of the poles. That the poles do approach each other appears probable from the fact that, although in the equatorial plate stage, the poles extend almost or quite to the cell-wall; yet the daughter-nuclei are always some distance from it. The difference in the character of the threads of the inner and outer portions of the cytoplasmic reticulum, which has been mentioned above, increases as spindle form- ation progresses. The threads of the inner portion become less granular and more sharply defined, while those of the outer portion behave in exactly opposite fashion, becoming more and more granular, and finally coming to resemble accumulations of granules rather than threads. Sometimes in the bipolar stage only a few meshes remain and these lie close to the cell-wall (fig. 19). At other times in the same stage such a reticulum still extends throughout the cell (fig. 18). Frequently, when the chromosomes have reached the poles, and also after the daughter-nuclei are formed, branching fibers extend in all directions from the poles almost to the cell-wall. At this time the granular zone may still be discernable, although it may be more or less obscured by the presence of granular matter both inside and outside of the zone (fig. 21). Before long the spindle fibers are separated from the daughter-nuclei by granular matter and each daughter-nucleus is surrounded by a dense granular mass of cytoplasm (fig. 22). Many recent observations on spindle- Sovmaious in higher plants show that the developing spindle passes through a multipolar stage which becomes bipolar by the fusion of the poles, or by.the withdrawal of some of them. Multipolar stages are figured and described by Farmer (1893 and 1895), Belajeff (1894), Strasburger (1895), Osterhout (1897), Mottier (1897 @ and 6), Juel (1897), Webber (1897) and Debski (1897). But the origin of the multi- polar spindle has been worked out in only two or three Bot.—VOL. I.] WILLIAMS—PASSIFLORA CCRULEA. 195 cases. Osterhout (1897) describes it for the spore-mother- cell of EHguzsetum as follows: ‘The process begins by the meshes of the reticulum immediately surrounding the nu- cleus becoming drawn out parallel to the nuclear wall and forming a sort of skein about it. This condition is transitory; soon the cytoplasm immediately surrounding the nucleus does not consist of a network, but of radially arranged fibers which gradually grow longer and push out into the surrounding cytoplasm. At first these cross each other in all directions but soon become parallel. Later, they bend toward each other and form numerous cones which begin to fuse, the nuclear membrane, mean- while, disappearing in places. The threads of the cones penetrate into the nucleus and come into connection with the linin threads. The nuclear wall entirely disappears and the multipolar spindle is formed. Mottier (1897@) describes a similar origin of the multipolar spindle. The following is a summary of Belajeff’s account of its origin in the pollen- mother-cell of ZLarzx: The cytoplasm immediately sur- rounding the nucleus takes on an arrangement which, at first glance, produces the impression of a skein-like mass of threads wound around the nucleus, forming a distinct zone. It really consists of a network, the meshes of which are drawn out parallel to the nuclear wall. (In this, Stras- burger’s work on ZLarzx agrees with Belajeff’s). The nucleus soon becomes filled with a linin network. The nuclear wall disappears entirely and the inner linin network and the outer cytoplasmic one unite to form a continuous reticulum. Fibers pass from the ‘‘ Centralkérper’’ (as Belajeff calls the central network) to the cell-wall, running radially or tangentially through the cell. The fibers, by their contraction, pull the ‘‘ Centralkérper’’ out into a three- or four-angled body. Several fibers unite in a pole which is situated toward the periphery of the cell and is connected by fibers with the cell-wall, as well as with other poles. The chromosomes lie in the middle of the ‘‘ Central- k6rper,’’ surrounded and held in place by a network whose fibers are gradually drawn out to form the spindle-fibers. 196 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. These are fastened to the chromosome in two groups, one at each end. All the fibers of the ‘‘ Centralkérper’’ finally become directed toward one or another of the poles, whose > number is finally reduced to two, probably by fusion. It will be seen that the process in Hquzsetum has but little in common with that in Passzfora, and that the process in Larix, on the other hand, agrees with that in Passzflora in some very essential points. In both Larzx and Passiflora — the cytoplasm and linin take part in the formation of a central network which is pulled out to form cones, which later become composed of free fibers. The cones then fuse to form the bipolar spindle. Various details, how- ever, are different in the two. The peculiar arrange- ment of the cytoplasm showing a zone around the nucleus (fig. 2) in which the meshes are drawn out parallel to the nuclear wall recalls Belajeff’s description, but, as in Equisetum, is of short duration and it is difficult to say what part, if any, it plays in spindle-formation. The same arrangement has been described by Strasburger (1895) for Larix. He finds it to have about the same connection with the formation of the spindle as does Belajeff. Other observers have described a similar arrangement, but do not explain it. The radial arrangement of the cytoplasmic network (fig. 3) corresponds very closely with Belajeff’s description of the state of the cytoplasm of Lavzx before the meshes begin to be drawn out parallel to the nuclear wall. In Passzflora, however, the radial arrangement occurs after this stage. The radial arrangement described by Guignard (1891), Farmer (1893 and 1895), Osterhout (1897), and Mottier (1897a) seems to be due to the presence of free fibers, not to a network. Belajeff speaks of fibers which extend out from the cen- tral network to the cell-wall; these run radially or tangen- tially. The appearance of fig. 4 certainly seems to indicate that there is a similar condition here. Some of the fibers can be traced to the cell-wall, but there is no evidence that they pull the central network out into cones as they Bot.—VOL. I.] WILLIAMS—PASSIFLORA C@RULEA. 197 do in Larix; indeed, their disappearance before the forma- tion of the multipolar spindle would seem to prove that _they have no such function. The unbroken network stage in Passzflora (figs. 11 and 12) is much more striking than in Larzx, and the cones are formed directly by the drawing out of this network without a preliminary converging of radial fibers such as Belajeff describes. The conversion of a reticulum into spindle-fibers has also been described by Wilson (1895). In his work on the egg of the sea-urchin, he shows that the spindle-fibers are differ- entiated out of the linin reticulum. To point out briefly the differences in the mode of form- ation of the multipolar spindle in EAguzsetum, Larix, and Passiflora: in Equtsetum, the cones are formed solely by the converging of radial fibers and only come into contact with the linin network after they have reached their full size; in Larix, they are formed by the converging of radial fibers, and subsequently increase in size by the pulling out of the network at the places where they are attached to it; in Passiflora, they result from the simple drawing out of the network. SUMMARY. The cytoplasm of the cell in the spireme stage is com- posed of two separate elements. First, a reticulum stretch- ing from nuclear wall to cell-wall, and, second, a granular substance uniformly distributed in and upon the meshes of the former. When spindle-formation begins the granular substance collects some distance from the cell-wall and forms a dense zone—the granular zone (figs. I1, 12, 13). The spindle is formed as follows: The nuclear cavity gradually becomes filled with a linin network. The nuclear wall is transformed into a meshwork which connects the linin network with the cytoplasmic reticulum, making a sin- gle continuous network which fills the entire cell. That portion of the network which lies in the center of the cell, 198 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. surrounded by the granular zone (fig. 12), forms the spin- dle. It projects out at various points to form cones in which the meshes are gradually stretched and finally form free fibers (fig. 13). The whole reticulum inside of the gran- ular zone finally becomes transformed into such cones. These at first point in all directions, but finally form two groups (fig. 15). In each of these groups the poles fuse to form a single one, thus producing the bipolar spindle. The spindle is formed by the rearrangement of pre-exist- ing structures, namely, of linin, nuclear wall, and cyto- plasmic reticulum. No centrosome is present, nor could any special spindle-forming substance be recognized. BOTANICAL LABORATORY, UNIVERSITY OF CALIFORNIA, BERKELEY, CALIFORNIA, May 1, 1898. Bor.—VOL. I.] WILLIAMS—PASSIFLORA C@RULEA. 199 1894. 1897. 1897. 1893. 1895. 1891. 1897. 18974a. 18976. 1897. 1895. 1897. 1895. BIBLIOGRAPHY. BELAJEFF, WL. Zur Kenntniss der Karyokinese bei den Pflanzen. Flora, Bd. LXXIX, S. 430-442. Desskt, B. Beobachtungen iiber Kerntheilung bei Chara fragilis. Pring sh. Jahrb., Bd. XXX, S. 227. Eisen, G. Notes on fixation, stains, the alcohol method, etc. Zeitschr. wiss. Mikr., Bd. XIV, S. 195-202. FARMER, J. B. On nuclear division in the pollen-mother-cells of Lilium Martagon. Aznals Botany, Vol. VIII, p. 392. Ueber Kerntheilung in Lilium-Antheren, besonders in Bezug auf die Centrosomenfrage. Vora, Bd. LXXX, S. 56. GUIGNARD, L. Nouvelles études sur la fécondation. Ann. des Sci. Nat. Bot., 7th Ser., Tome XIV, p. 163. JueLt, H.O. Die Kerntheilung in den Pollenmutterzellen von Hem- erocallis fulva und die bei denselben auftretenden Unregelmassig- keiten. Pringsh. Jahrb., Bd. XXX, S. 205. Mortier, D. M. Beitrage zur Kenntniss der Kerntheilung in den Pollenmutterzellen einiger Dikotylen und Monokotylen. Pringsh. Jahrb., Bd. XXX, S. 169. Ueber das Verhalten der Kerne bei der Entwickelung des Embryosacks. Pringsh. Jahrb., Bd. XXXI, S. 125. OstTERHOuT, W. J. V. Ueber Entstehung der karyokinetischen Spindel bei Equisetum. Pringsh. Jahrb., Bd. XXX, S. 159. STRASBURGER, E. Karyokinetische Probleme. /Pringsh. /Jahr6., Bd. XXVIII, S. 151. Wesser, H. J. Peculiar Structures occurring in the Pollen Tube of Zamia. Bot. Gazette, Vol. XXIII, p. 453. Witson, E. B. Archoplasm, Centrosome, and Chromatin in the Sea-Urchin Egg. Journ. Morph., Vol. XI, p. 443. 200 Fig. 1. Fig. Fig. Fig. Fig. Fig. CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. EXPLANATION OF PLATE XXXVII. The Abbe camera was used in drawing. Objective, Zeiss, oil immersion 1/12; ocular, compensating No. 6. Represents a cell with the chromatin in the spireme stage. The cytoplasm is composed of two distinct elements: One is fibrous and forms a reticulum throughout the cell from nuclear wall to cell-wall. The other is granular and is uniformly distributed within and upon the meshes of the reticulum. The meshes of the reticulum are irregular in form and are smaller immediately surrounding the nucleus than elsewhere. Its threads are sinuous and irregularly knotted. The nucleolus is large and contains a small vacuole. The cytoplasm exhibits a peculiar structure. The fibers of the re- ticulum immediately about the nucleus are drawn out into long, narrow meshes parallel to the nuclear wall, forming a sort of weft about it. These appear, at first glance, as long, continuous fibers wound about the nucleus. The meshes of the cytoplasmic reticulum are drawn out at right angles to the nuclear wall and appear as if radiating from the nucleus. The chromosomes are very near the nuclear wall. The nucleolus is still about as large as that in fig. 1 and contains several vacuoles. Linin granules and threads partially fill the nuclear cavity. The granular element of the cytoplasm is beginning to accumulate in dense masses. Deeply staining, irregular strands are to be seen in the cytoplasm. They are radially arranged and some of them extend clear to the cell-wall. At first glance they appear as free fibers, but close examination shows them to be only thickened, more granular strands of the cytoplasmic reticulum. In other respects the figure resembles fig. 3. Three or four conical groups of fibers extend from the nuclear wall into the cytoplasm. Otherwise the figure is very much like fig. 4. The deeply staining, radially arranged, cytoplasmic strands seen in figs. 4 and 5 are less marked. There is a good development of linin. The chromosomes are very near the nuclear wall, which is irregular in outline and is beginning to be transformed into a meshwork. —— Proc CanAcap,Sera= Ser. Bor Voi 202 Fig. Fig. Fig. Fig. Fig. Fig. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XXXVIII. 7. Differs but slightly from fig. 6. The transformation of the nuclear wall has progressed farther and the wall is interrupted at several points. 8. Is similar to figures 6 and 7. The alteration in the nuclear wall, however, is still more marked; the linin is more abundant, its threads are coarser, contain more granules, and stain more deeply. 9g. The cell resembles that represented in fig. 8. The nuclear wall has apparently been transformed into a meshwork which forms one continuous network with the cytoplasmic reticulum on one side and with the linin reticulum on the other. The quantity of linin has increased. 10. Is much like fig. 9, although the transformation of the nuclear wall has not progressed so far. The appearance of this and the two following figures might give rise to the suspicion that the sec- tions of the nucleus are not median and that the appearance of the network filling the nuclear cavity is due to focussing on the inner surface of the nuclear wall. Careful focussing, however, shows conclusively that this is not the case. 11. The nuclear wall, as such, is no longer to be distinguished. In some places it forms a fibrous belt two or three meshes in width. These fibers stain more deeply than any others; they are coarser and contain more granules; they help to form the continuous network which fills the cell. In one place every trace of the nuclear wall is lost; nothing remains to mark the transition from linin reticulum to cytoplasmic reticulum. The granular part of the cytoplasm now forms a complete zone between the fibrous belt representing the nuclear wall and the cell-wall. 12. Every trace of the nuclear wall has disappeared. There is nothing to even indicate its position, and an uninterrupted network stretches from cell-wall to cell-wall. The granular zone referred to in fig. 11 is well marked. “ olin < i ir ee *~ es Doge t a ys 10. PASSIFLORA CIERULEA Z7VNV; LITRASITION & REY, SF “Hie . i th} iy eo) 204 Fig. 13. Fig. 14. Fig. 15. CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. EXPLANATION OF PLATE XXXIX. Shows the part of the reticulum inside of the granular zone begin- ning to be drawn out into cones. In many places, the reticulum is still continuous from the chromosomes to the cell-wall. Represents an early multipolar stage. There is no longer a reticu- lum inside of the granular zone. It has been drawn out at various points to form cones in which the meshes have been gradually stretched until now they form free fibers. These cones and free fibers extend in all directions. The poles of the multipolar spindle lying within the granular zone show a tendency to gather into two groups which are probably destined to form the poles of the bipolar spindle. + Figs. 16 and 17. Represent very late multipolar stages with the poles fusing. Fig. 18. ¥ The granular zone is clearly evident in each. The poles have fused completely, forming a sharply pointed spindle. SITHSRTION SREY SF : 4 PASSIFLORA- CIERULEA oe it it ne at ee oh At Une OY 4 - ‘ ? . ia Ay ms ; j AIT eh ae 206 - 20. es2hs . 22. CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. EXPLANATION OF PLATE XL. Shows a spindle in the anaphase. The granular zone is well marked. From the spindle irregular fibers, probably mantle- fibers, pass out and penetrate the granular zone. Represents a spindle in the anaphase. Branching fibers extend from the chromosome-groups toward the cell-wall. The granu- lar zone is somewhat obscured by a more general distribution of granular matter between the spindle and the cell-wall. Shows the daughter-nuclei formed. The spindle-fibers still con- nect them. Branching fibers radiate in all directions from them. The spindle-fibers are separated from the daughter-nuclei, each of which is surrounded by a dense mass of granular cytoplasm. Pare XL * bi i a ORA CIERULEA LuvN.. ASSIFL : SER. Bor Vonl. Acad Sc1.3 / THE NATURE OF THE ASSOCIATION OF ALGA AND FUNGUS IN LICHENS. BY GEORGE JAMES PEIRCE, Assistant Professor of Plant- Physiology, Leland Stanford Junior University. CONTENTS. PAGE. PLATE XLI. Bae ENT RODE LION tail wate) ha ds gains 8 Bin Jie ey Sidhe voila olen) ie wax may ats eis 207 ee Lae GERMINARION OR SPORES.5,.5 25 «ca acenaite cere tee eee 211 Ill. THE RELATIONS OF GONIDIA AND HyYPH2 AS SHOWN BY GUTOR SEs fe ts secre cielo ah or aihe o eraees o besitos GR eee 216 IV. THe RELATIONS OF GONIDIA AND HypHZ AS SHOWN BY MIGROTOMESSECDIONS co s.c.cc is oo ae nia nes: ee See ee 221 V. THE SIGNIFICANCE OF THE WATER-CONTENT OF RAMALINA EU TL GU TAA TAC aie vare eh taller etc yete ont ats dite see Renee eae 229 VI. THE INFLUENCE OF MECHANICAL STRAINS ON GROWTH..... 231 VII. NucLeus oR PYRENOID AS THE CENTRAL BoDy oF CysTo- CORCUS MGEULS ice sac valuta dina Seine s aslo je ce eee 234 MCP = SSUINUMTARY vice cs crate od A cities ad Sod oheew «loon otiere bre le ae amet ae Ree 236 BBLIOGRAPINY co a) stetcle esta aie nit s)irfeiste mis) 2'2\~)> (<0 sipls a a re ie ee ea ae 239 EXPLANATION OR) PEATE bo hella ance s's'¢ nrota)s# 5) t oc cgiety eens 240 I. INTRODUCTION. Tue last few years have seen a revival of interest, grati- fyingly widespread, in the study of lichens. Except during the few years immediately following the writings of DeBary, Schwendener and Reinke, which wrought a revolution in our conception of lichens, they have been studied mainly by systematists. These found in macroscopic characters all that was required to determine an old or to describe a new species. Every one must have felt, even before Reinke recently recalled the question so forcibly to the minds of botanists, that by no means the last word had been said about these interesting organisms, and that the study of | 207 ] May 13, 1899. 208 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. other than superficial characters would repay the investi- gator. To the physiologist they especially appeal because they are not simple organisms, but an association of two simple organisms. The nature of the association is variously regarded, from pure parasitism of one member upon the other, to associa- tion mutually beneficial, if not absolutely necessary, to the two. Schwendener has always regarded the association as pure parasitism of the fungus upon the alga, and his school advocates the same view. The opposite view is held by Reinke (1894) and his sympathizers, of whom Schneider (1896a) is at the moment perhaps the most conspicuous in this country. Reinke believes that the association is what he names Consortism, and that the ‘‘ Consortium’’ is as ‘*autonomous”’ as any plant. A question to which there are such opposite answers by such eminent investigators cannot be regarded as answered at all. This paper is a contribution to the discussion, and I shall be glad if the account of my work and its results leads at all to the solution of the problem. I must remark at the outset that much of my work may not be new; how much I cannot say, for though I have tried, I have not been able to have access to all the literature on this subject. Further- more, no one should venture, from the study of only a few forms, to draw conclusions regarding a great number, although the careful examination of a few forms will just- ify opinions regarding the others, if only these opinions are held tentatively, subject to modification by further researches. Beginning with Ramalina reticulata Krplhbr., a lichen so large that it forms a conspicuous feature of the landscape in some parts of California, I have examined several species and genera of lichens occurring hereabouts, and not trusting merely to microtome sections of ‘‘ fixed” material, I have had recourse to experiments and cultures on and of living material. The difficulty of minutely studying lichens alive is notoriously great, not least because of their extremely slow growth. Yet in their rates of Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 209 growth there must be great diversity. I am inclined to believe that Aeamalina reticulata grows more rapidly than most. My reasons are based on the universal occurrence of large specimens all about here, and also upon the behavior of this lichen in cultures. As emphasized by Benecke (1898) in a recent paper on the culture of Alga, and earlier by Moller (1887), the difficulty of excluding bac- teria from cultures made from vegetating, and even from spore, material, is very great. The presence of bacteria in a culture will at least delay and obscure, if it does not vitiate, the results; and yet to exclude the bacteria is impossible with our present crude culture-technique. The almost invariably gelatinous nature of the exterior of the lichen body renders the temoval of the bacteria from the surface impossible. During the latter part of a long dry season, the surface of such a lichen as Ramalina reticulata must be so dry, at least by day, that it is not likely to hold the bacteria or the fungus-spores that fall upon it, though they may easily remain caught in some angle or slight irregularity. During and for some time after a rain, ina _fog or on a dewy night, on the other hand, the gelatin- ous surface will be moist, at least sufficiently so in spots to cause whatever bacteria and fungus-spores fall upon it to stick. Moller (1887) recommends washing as thoroughly as possible in clean running water, the aim being to remove by the stream, as well as by floating, whatever may have fallen upon the surface. By this means he partially cleansed the apothecia from which he intended to collect spores for his cultures. Defective as such cleansing is for the apothecia, which are not especially gelatinous on the surface, it is still more so for the much more gelatinous surface of the thallus. The running water does remove some of the foreign objects, but some of the most gelatinous of these (for example, bacteria in the vegetative condition) cling to the gelatinous surface of the thallus in spite of the most thor- ough attempts to dislodge them. For this reason, even if sterilized water be used for the washing, a discourag- ingly large proportion of the thallus fragments used for 210 CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER. culture are accompanied by bacteria which multiply only too rapidly. In spite of thorough sterilization of the slides, cover-glasses, vessels «and instruments used in connection with the cultures, and the utmost pains to pre- vent the introduction into the cultures of organisms from outside, I have failed in most cases to make pure cultures, because the lichen fragments themselves carried bacteria, and substances nutritious for them, into the culture-media. In many cases, however, the number of bacteria was con- stantly small. For most of my cultures I used the ‘‘hanging-drop ”’ method of the bacteriologist, employing for the purpose slides with two concavities, ground sufficiently deep to avoid contact of the drop with the bottom of the concavity. The cover-glasses were sealed upon the slide with vaseline. If the concavities are reasonably large and deep (e. g. 13 mm. diameter, 1 mm. depth) a hanging-drop surrounded by an adequate volume of air, and large enough to supply food-material for several weeks, may be used. It must be admitted, however, that hanging-drop cultures have only a limited usefulness, for the food-supply presently becomes exhausted, and there may accumulate in the drop undue amounts of the dissolved excreta of the organisms present. Various methods have been employed by others in pre- paring material for culture. As culture-media for the spores and lichen fragments, I have experimented with boiled tap-water, with the moisture carried by the spores expelled upon the cover-glass, with a drop of the ordinary nutrient beef-gelatine of the bacteriologist, with similar nutrient agar-agar, and with the distilled water used for general purposes in the laboratory. The distilled water was poisonous. I did not consider it necessary to deter- mine the reason for this, though one would suspect it of being acid and possibly of containing traces of copper or other injurious salts. The distilled water prepared in quantity in the chemical laboratory, and supplied by it to the other laboratories of this University, is sufficiently pure BoT.—VOL. I.] PEIRCE—NATURE OF LICHENS. 211 for ordinary purposes, but it is ill-suited to culture experi- ments on living organisms, as repeated experience shows. The effect of our distilled water on motile bacteria is marked and immediate. Bacteria actively motile in the media in which they are cultivated in the laboratory will continue to be motile at the same rate if placed in a drop of sterilized tap-water. Transferred either from the cul- ture directly or from sterilized tap-water into which they have been inoculated, to a drop of distilled water, they promptly come to rest and will not recover their motility. The unsatisfactoriness of our distilled water for lichen cultures may, therefore, be attributed to some poison in it and not to the lack of oxygen, for boiled tap-water will be at least as free from oxygen as ordinary distilled water. The nutrient agar-agar and gelatine above mentioned were made with tap-water (not distilled water) and had previously been proved to be well adapted to bacteria cul- ture. In the lichen cultures also they were perfectly whole- some, as attested by the only too rapid growth of whatever bacteria were carried into the drops by the spores. Il. Tue GERMINATION OF SPORES. The generally practiced method of obtaining lichen- spores for cover-glass cultures is the following:—the thor- oughly cleansed and sterilized cover-glasses, either dry or with a drop of some sterilized nutrient medium, are so placed that some of the spores discharged from an apothe- cium will strike upon them. The lichen material was pre- pared thus: fresh specimens of /. reticulata, brought into the laboratory from the trees on which they were growing, were brushed and shaken to remove loose dust, etc., from the surface, cut into small pieces, each piece having only one apothecium, and placed on moistened filter-paper in a Petri dish and covered. Near each apothecium, but not touching the lichen, was placed a cover-glass prepared as just described, and supported in such an inclined position that bacteria, etc., which might be floating in the air in the 212 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Petri dish could not settle on the surface intended for the culture. The sterilized cover-glasses, although carefully kept ‘‘butter-side down,”’ might readily become infected by air-currents set up by the necessary movement of the cover-glass and by lowering the cover of the Petri dish. Apart from such accidental infection, the possibility of which was carefully reduced to the minimum, the only bacteria which could get into the cultures were those car- ried in with the spores. The considerable force with which the spores are discharged is indicated, as others have repeatedly shown, by the distance to which they go. This force is developed by the swelling, when wet, of the gelat- inous walls of the paraphyses and other cells forming the contents and the walls of the apothecia. In less than five hours, enough spores will usually be ejaculated upon the cover-glass to justify sealing it over a concave slide. Spores coilected thus upon cover-glasses spread thin with agar-agar or gelatine germinated in seven Gays. At the same time I collected spores upon cover-glasses introduced dry into the Petri dish. Owing to the cooling of the labo- ratory (from morning, when I set cover-glasses to collect spores, to afternoon, when I sealed the cover-glasses on the concave slides) there accumulated on the cover-glasses not only the moisture carried by the spores, but also that con- densed from the damp air in the dish. Thus a supply of water at least adequate for germination was insured, what- ever may be said of the nutrient value of such water. After the same lapse of time (seven days) spores had germinated on these cover-glasses. In all the cultures the number of spores germinated was very small in proportion to the number sown. If I had sufficient confidence in the suitableness of agar-agar and gelatine for lichen cultures, and in the nutritive value of the condensed and spore-carried water, I might draw a definite and significant conclusion regarding the value of the spores of A. reticulata as reproductive bodies. I do not feel jus- tified in doing this, though I will call attention to two things. First, there must be a very considerable number of spores Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 213 discharged, both in nature and in the laboratory, before they are ripe and whenever the apothecia are wet. Second, that by bearing this in mind and by using trustworthy cul- ture media for such cover-glass preparations, subjecting them to otherwise normal conditions of warmth, light and aération, the percentage of ripe spores capable of germina- tion could be determined. From these data the relative values of the spore-method and the purely vegetative method of dispersal could be ascertained. This is, how- ever, hardly necessary, for, as I have shown in a previous paper (1898), the great majority of the specimens of 7. reticulata to be found about here are evidently fragments, while only very few have grown where they are from the spore. This lichen may, earlier in its history, have pro- duced even more spores in each apothecium and more apothecia than now, but this would seem hardly necessary, considering the ease with which fruiting specimens and an abundance of spores may be obtained at all seasons. It seems more probable that in this lichen, as is claimed also for others, superiority of the purely vegetative method of dissemination (by means of fragments torn away by the wind) is resulting in the decrease of the germinating power of the spores. The superiority of the vegetative method of reproduction is two-fold: it insures the presence in the new place of both members of the lichen association—the hyphe and the gonidia, as is the case in the soredia of other lichens — and it results in much wider dispersal than is possible with spores or soredia. The vegetative method of dissemination of this lichen, so evidently a very valuable one and perhaps the most perfect among lichens, may well react on the pro- duction of spores; if not already as to their number, then certainly as to their quality. Although the spores germinated in the same length of time and in about the same numbers (proportionally) in the agar- agar, gelatine and water cultures, the subsequent develop- ment was unequal. In three days after the first germina- tions were observed, the germ-tubes had grown somewhat 214 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. but had not branched in the agar-agar and very few if any more spores had germinated, whereas in the water culture the tubes were longer, some of them had branched, as fig. 3 shows, and the number of spores germinated had increased considerably. In eight days more, that is,in eighteen days after the spores were collected on the cover-glasses, growth had ceased in all the cultures. The differences between the water cultures on the one hand, and the agar and gela- tine cultures on the other, were now still more evident. The germinated and ungerminated spores in the water cul- tures still looked healthy, and the number of bacteria in the cultures was comparatively small. In the other cultures, however, there were many spores evidently dead, the remainder looked unhealthy, the germinated ones least so, and the number of bacteria flourishing on and in the agar- agar and the gelatine was sufficiently large to justify the inference that they were an important if not the sole cause of the unsatisfactory condition of the spores. The contents of the spores which appeared to be alive at all were far less highly refractive than at first; perhaps, but not wholly, because of the consumption of the oil which was stored in them in the form of drops of very consider- able size (figs. 1 a and 46, 2,3 and 4). The death of the spores was marked by certain changes in the cell contents: the protoplasm lost its granular character and gradually became indistinguishable from the wall and the cell-sap, the oil drops spread, became confluent with one another, later appeared to fill the whole cavity of the cell, and finally disappeared altogether, leaving only the empty cell-wall.. Minutely to follow the disorganization-phenomena (‘‘ Dis- organizationserscheinungen ’’) is very difficult, owing to the small size of these two-celled spores. From what I have noticed, I can only suspect, not con- clude, that the spores were killed and disorganized by the products of the discouragingly active bacteria. That the same appearances may be produced in another way is. shown by a water culture made as above described, but with the hollow below the cover-glass filled with sterilized BotT.—VOL. I.] PEIRCE—NATURE OF LICHENS. 215 tap-water. In this preparation the spores all died within three days, the protoplasm disappeared, the oil drops became confluent and filled the cell-cavities. Excess of water, inadequate diffusion of oxygen and carbon-dioxide, checked respiration, the diffusion into the water of some poison contained in the vaseline used for sealing — all or any of these may have been the cause of death and of the breaking down of the cell-contents. Bacteria were practi- cally absent. If it were possible to clean an apothecium so thoroughly that no bacteria would be carried by the spores ejected upon the cover-glass, it seems probable that agar-agar would prove a good medium upon which to cultivate lichens from the spores. This is conceivable but not readily attainable. If practicable, agar-agar cover-glass cultures would be very useful, for they could be minutely watched with the high powers of the microscope. In the cultures just described, made during the latter part of the rainy season, in February, only one cell in each spore put out a germ-tube. Water cultures begun about the middle of March, when it was warmer out of doors and the rains were practically over, were more successful in that both cells of some spores germinated, though in all of these the tubes from the two cells were unequal in length, and only one tube from a spore branched. Having obtained germinated spores by means of these cultures, I attempted to reconstruct a Aamalina by putting these where they could easily reach gonidia from a Rama- lina thallus. The method employed was this: Into a tap- water hanging-drop culture of A. retzculata fragments and isolated gonidia, obtained in the way described on page 218, I introduced by sterilized platinum loop a number of germ- inated spores, stirring gently so as to mix the spores with the gonidia. Germinated spores were thus brought close to isolated and apparently healthy gonidia. One would naturally expect the germ-tubes to be chemotropically attracted by the gonidia and to bend towards them. As - illustrated by fig. 4, the hyphe put forth by the spores did 216 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. not bend at all. The hypha figured was about half as long when the germinated spores were sown in the gonidia culture. It continued to grow straight on, did not branch, and never united with the adjacent or with any other gonidium. The same is true of all the others, and although I kept the preparations until all the gonidia were evidently dead, though the hyphe still looked healthy, no union of hyphz and gonidia took place. Undoubtedly the bacteria, which overran these cultures within a week after they were made, are responsible for the failure of the gonidia to attract the hyphe, of the hyphe chemotropically to respond, and of the two to form a new lichen. That the failure of the attempts just described is not due to an unhealthy condition of the gonidia produced by the isolation is proved on page 220. DeBary (1887) reports from Stahl’s (1877) investiga- tions on the formation of the lichen-thallus, that ‘* smaller branchlets are formed at the points of contact’’ of the hyphe with an algal cell, which closely embrace it and enclose it in fresh ramifications. Whether actual contact is necessary, whether the gonidia stimulate the germ-tubes to bend or to branch, whether the union of gonidium and hypha is always by branches of the germ-tube and not by the tube itself —these are questions which renewed investi- gations only can answer. When, by greater skill and better methods than I have possessed, one is able to put together isolated healthy gonidia and healthy germinated spores, in hanging-drop cultures free from bacteria, the formation and growth of the lichen can be followed, step by step, from the chemotropic behavior of the germ-tubes to the complete investment of the gonidia by the fungus. Ill. Tuer RELATIONS oF GONIDIA AND HypPH AS SHOWN BY CULTURES. Although I have been unable to reconstruct a lichen, still I believe that by other means I have obtained some BoT.— VOL. I.] PEIRCE—NATURE OF LICHENS. 217 information regarding the questions which I hoped to answer in that way. To isolate the gonidia of 7. reticulata without injuring them is a matter of considerable difficulty owing to their position in the compact thallus. The methods described by others do not lend themselves to the species upon which I have worked. Baranetzky (1869) apparently made no attempt to isolate the gonidia from the hyphe directly, con- tenting himself with making thin hand-sections of living lichens (Collema, etc.), and placing them in moist cham- bers. The result was that the gonidia multiplied and grew more rapidly than the hyphe, coming out upon the upper surface of the sections and accumulating there practically free from hyphe. Such a method as this, though satisfac- tory with filamentous gonidia in lichens of comparatively loose texture, is ill suited to lichens of close texture, containing spherical gonidial cells (Cystococcus, etc.) The gonidia do not become free under these conditions. The reason for this may lie in the absence of intracellular haustoria in Baranetzky’s lichens and in the close invest- ment and the penetration of the gonidial cells by the hyphe of the lichens which I have studied. The gonidia of the latter seem to be too completely in the power of the hyphe to be liberated in this way. Teasing out thin sections in water by needles is a slow, tiresome and unprofitable task, for even in this way it is almost impossible to break up the small groups of gonidia closely invested and penetrated by hyphe. These little masses placed in cultures behave like lichens, growing almost imperceptibly; the gonidia do not grow and multiply rapidly and so disrupt the little groups and escape to the surface. When air-dry, 7. reticulata is tough and leathery, difficult to section, readily bent, but not at all friable. Sections of the air-dry thallus cannot be pulled to pieces dry by needles, and, when placed in a drop of water on the slide, expand without at all loosening the hold of the hyphez upon the gonidia. Teasing out sections cut dry and then placed in 218 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. water is as unsatisfactory as though the sections were cut from a moist thallus. The toughness of the thallus is due to the longitudinal direction and the compact arrangement of the rather thick- walled hyphe near the surface. The surface is soft and gelatinous when moist, hard when air-dry. The gelatinous matter of the outer portions of the thallus both holds water very strongly and takes it up very quickly. Air-dry at the end of the long dry season, the lichen still contains 13.9 per cent. of water or nearly one-seventh of its weight, as shown by the following figures: Weight air-dry ..... 4.72 gr. Weight constant at.. 4.06 gr. when dried several hours at 90° C. Weight 10st cuca 0.66 gr. Percentage of water .13.9 The dried lichen must be weighed quickly, for it rapidly absorbs moisture from the air and returns to its original weight. When absolutely dry it is as brittle as it ordinarily is tough, and it may be ground in a mortar to a fine powder. Microscopic examination’ of this powder in water shows it to consist of minute fragments of hyphz interspersed with isolated gonidia apparently uninjured by the grinding. It occurred to me to attempt drying the lichen at a temperature low enough to avoid injuring the gonidia. Warming in an oven for five hours at 36° C. (96° Fahr.), a temperature often attained, and sometimes maintained for several hours, on a summer day, proved to be perfectly harmless to the gonidia, but it did not dry the lichen to the point of brittle- ness. To attain brittleness without further heating, small pieces were kept for four days in a desiccator containing glacial phosphoric acid. Even after this treatment the fragments were not perfectly dry and brittle, though grind- ing rapidly in a mortar yielded powder of which the finest part contained isolated and apparently uninjured gonidia. Some of this was inoculated by needle into hanging drops of sterilized tap-water. Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 219 The individual behavior of the isolated gonidia, fairly scattered throughout the hanging drop, could be easily followed. Many of them were entirely free from any trace of hyphz, while some had short pieces of hyphae still attached to them. Of the latter, some gonidia, as shown by fig. 12, had haustoria in them, while others appeared to have the hyphz only adhering or attached to the outside (fig. 7). A difference in size and in subsequent behavior between those gonidia absolutely free from the fungus, and those upon which and in which fungus fragments still remained, is to be noted. Those gonidia which, though enclosed within the lichen, have not been invested by fungus hyphae, still less penetrated by haustoria, are larger, deeper green, and in every way healthier looking than the others. How is this to be accounted for ? The enclosed but not invested algal cells seem to me to be the only ones in the lichen thallus which are at all advantageously situated. They do have a more con- stant supply of aqueous solutions of food materials than their fellows outside, they are shaded from excessive light, screened from excessive heat, protected from excessive dryness, shielded perhaps from ‘sudden changes of any kind, possibly manured by the excreta of the fungus. In these ways they may be better off than their unhoused fellows, and sometimes at least they are larger. They are not subject to the drain upon their vital energies which those gonidia invested and penetrated by hyphe must withstand if they are to survive. The fungus can obtain from these gonidia with which it is not in permanent and intimate contact only such substances as diffuse from them into the water of the lichen-body; whereas it can withdraw through a haustorium within the cell, and even through the cell-wall with which a hypha is closely in contact, the foods elaborated by the gonidia for their own use. Gonidia not invested by hyphe are to be found in every preparation. They are conspicuous from their larger size, deeper color and generally healthier appearance. It may have been these which suggested the notion that green 220 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. cells are better off as gonidia than as independent alge. The next difference to be noted between the invested and uninvested gonidia is in their subsequent behavior in culture. Within a very few days (at the utmost six) the gonidia which contain haustoria, or which have hyphal. fragments still attached to them, have either divided by internal cell- division at least once, or their cell-contents have contracted away from the wall. Figures 6, 7, 8, 9 show various stages in the division of invested gonidia, in fig. 6 into two, in fig. 7 into four, in fig. 8 another divided into four, in fig. 9 at a the same cell twenty-four hours after, when the wall of the mother-cell had become dissolved, setting free eight daughter-cells. In fig. 9 at the *, and in fig. 10, we have gonidial cells in which the haustoria, broken away from the hyphe which formed and bore them, are plainly visible, the contents of the gonidial cells contracting away from, and in in this way escaping, the haustoria. Later the contracted mass surrounds itself by a new wall and behaves like the uninvested gonidia. These last do not divide in the culture for a very long time. The only difference between the invested and unin- vested gonidia in the culture being in their relation to an organism incapable of manufacturing its own food, it can be safely concluded that the cell-contents of the former divide, or contract away from the wall, in order to eliminate the foreign organism, and to produce individuals which will be free from it. The conditions prevailing in a hang- ing-drop culture cannot be entirely favorable to the green cells; if they were, all the gonidial cells would sooner or later divide. Haustoria within, or hyphal branches closely enwrapping them, must irritate the invested gonidia. The response to this irritation is an effort to get rid of it by contraction or by division. The uninvested gonidia, not subjected to this irritation, do not contract or divide. The gonidia of /. retzculata are in rounded masses fairly compact and held together by hyphe (figs. 5 and 11, and Peirce [1898], p. 412). Many of these masses are composed of small cells, and some of the cells are likely Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 221 to be dividing internally (figs. 6 and 9). From such divi- sions there result new gonidia, but it is not necessary to infer, as some do, that the gonidia are thus only amiably doing their part toward the growth and the permanence of the lichen. On the contrary, as Hedlund (1895) describes for Catillaria denigrata (Fr.) and C. prasina (Fr.), and as I have just shown in the above, we have in both cases merely the attempt on the part of the gonidia to divide in such a way as to exclude the haustoria from as many of the daughter-cells as possible. That the gonidia cannot be benefited, but are evidently injured, by having haustoria within them, I shall show subsequently (page 226) ; and if the haustoria are injurious, is it not sensible to suppose that the gonidia will try to get rid of them ? If they are harm- less or beneficial why should there be frequent divisions which inevitably result in the formation of daughter-cells free from haustoria? Ina culture, the conditions in which seem to be more favorable to the uninjured gonidia than to the much broken and greatly injured hyphe, the gonidia succeed in eliminating the haustoria and in freeing them- selves from enclosing hyphe. Imbedded in the body of the lichen, surrounded on all sides by hyphe, the advantage in the struggle is to the other side; but the struggle and the manner of it are the same. The gonidia in the lichen seek by division to escape the hyphe. Some do—these flourish and grow large; most do not—these divide again as soon as possible and so remain small. IV. Tue ReLaTiIons oF GONIDIA AND HypPH AS sHOWN By MICROTOME SECTIONS. So far as I know, very little use has been made of modern cytological methods in the study of lichens. This is to be regretted, for without recourse to these, the intimacy of the relations of gonidia and hyphe must remain compara- tively obscure. It has already been demonstrated that our culture-methods are still so defective, and the growth of the fungus-component of lichens is so distressingly slow, 222 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. even under the most favorable conditions, that our knowl- edge of lichens, if it is based on hand sections and cultures only, must be incomplete. Hand sections are unsatisfactory at the best because they are so thick that important details are hidden by excess of material. Yet investigations of lichens which are based only on examinations of dead herbarium material, or even of freshly fixed, imbedded and sectioned specimens, are subject to the same danger of one-sidedness which sometimes threatens the work of those who know plants and animals only as embalmed tissues. It is deplorable that we are still utterly unable to obtain, and to keep healthily alive, thin sections of living material. As Fischer (1895) and others have repeatedly emphasized, a technique which is based on the employment of poisonous acids and salts is one which can be implicitly trusted only when their effects on the living protoplasm are exactly known. ‘The danger of being misled by the study of fixed material sectioned by microtome after imbedding is, how- ever, minimized by examination of living material, both sec- tioned and in culture, as a control. When the question before the student can be answered in part at least by deter- mining the relative position of cell-walls, nuclei, etc., the matter is still simpler. Such is in great measure the case with the questions which are under discussion in this paper. From microtome sections of lichens it is possible to obtain clearer views on the structural relations of hyphe and gonidia than can be had otherwise, and from these data, supplemented by culture experiments, conclusions regard- ing their physiological relations may safely be drawn. As reported elsewhere (Peirce, 1898a), the material was prepared as follows: freshly collected specimens of /#. reticulata, Spherophorus globiferus (Li.) D.C., and an Usnea, after being thoroughly wet, were kept in a moist chamber near a window for twenty-four hours. This allowed the gonidia to resume their photosynthetic activ- ities in the light, and the hyphe to become reasonably plump, if they had become dried out of doors. BoT.—VOL. I.] PEIRCE—NATURE OF LICHENS. 223 For fixing I used: Chromic acid, 3 per cent., at room temperature. Chromic acid, 1 per cent., just below boiling temperature. Flemming’s Chrom-osmic-acetic mixture, at room temperature. Corrosive sublimate, saturated solution in 35 per cent. alcohol, at room tem- perature. Corrosive sublimate, saturated solution in 35 per cent. alcohol, just below boiling. The last proved the most successful. Penetration at room temperature is comparatively slow, owing to the large amount of air enclosed in even wet lichens, whereas the air is quickly driven out by the hot liquid. At room tem- perature, the fragments treated with the fixing agent float, and must be exhausted under the pump. Those treated with the hot fixing-solution sink in it almost immediately. There is, therefore, a saving of time, as well as greater certainty of rapid penetration, if the fixing agent is applied hot. Dehydration should not be carried on too rapidly, for the gelatinous walls, especially of the outer hyphe, hold the water. After complete dehydration, the material was transferred from absolute alcohol to equal parts absolute alcohol and xylol, then to pure xylol, and to this were slowly added small fragments of paraffin melting at 55°. At the same time the temperature was slowly raised to about 40°. Paraffin was then added to the point of saturation and the xylol allowed to evaporate. When the xylol was driven off, the specimens were transferred to melted hard paraffin (melting-point about 59°), kept in this for two hours to remove the last trace of xylol and to insure per- fect penetration, then blocked, and sectioned at leisure. The gelatinous hyphe, soft and easily cut when wet, are now hard and rather brittle. For very thin sections, and these yield the most trustworthy results, the knife must be even sharper than for most tissues. The best sections were I » thick, made with a Zimmermann-Minot microtome, and fastened to the slide, after floating and straightening on warm water, by albumen fixative. Some were then stained with Czoker’s Alum-Cochineal and Bismark Brown. Some (2) May 29, 1899. 224 CALIFORNIA ACADEMY OF SCIENCES. [PROC. 3D SER. material was stained zz toto with the Cochineal, but this was less satisfactory than staining on the slide. Other sections were stained with Iodine-green-Fuchsin, washing out with Iodine in 80 per cent. alcohol, and still others with Anilin- - safranin-Gentian-violet-Orange G., according to the direc- tions given by Zimmermann (1893). Both of the double stains and the triple stain were very satisfactory in their different ways. The sections were examined with a Zeiss apochromatic 2 mm. oil-immersion objective and compensa- ting eye-pieces 6 and 12. All of the drawings were made with Abbé camera. Various gonidial cells of A. reticulata containing haus- toria are shown in section in figs. 11, 13, 15. Figures 12 and 14 were drawn from isolated and not from sectioned gonidia, but, for the sake of comparison, they may be referred to here. Figures 11, 12 and 13 show gonidial cells into which the haustoria can have penetrated only comparatively recently, as the protoplasm and chromat- ophores are apparently perfectly normal and uninjured. Figures 14 and 15, on the contrary, are of gonidial cells completely emptied, in which the haustoria still remain, normally plump and with granular protoplasm. Figures 13 and 15 show the haustoria in profile, figs. 12 and 14 end on. In fig. 13, the haustorium, and the hyphal cell of which it is a branch, are shown to be continuous as to both wall and cavity. Figure 15 shows the haustorium within the cell, but the hyphal cell of which it is a branch was below the plane of the section. This gonidial cell was invested by a larger number of hyphe than any other figured, and than the great majority of those examined. In the group of gonidial and hyphal cells shown in fig. 11, two gonidial cells show haustoria within them. It is nothing new to find haustoria in the gonidia of lichens. They have been repeatedly seen and sometimes figured, perhaps most strikingly by Schneider (1896a). It is generally said, however, that the haustoria, though pene- trating the walls of the gonidial cells, do not actually penetrate the protoplasm. Thus Hedlund (1895)—‘‘ Von Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 225 der Seite einer Hyphe wachst ein kruzer Ast heraus, der die diinne Membran vollstandig durchbohrt und innerhalb derselben mehr oder weniger kugelférmig anschwillt. Bei dem Eindringen der Hyphe zieht sich die Hautschicht des Protoplasmas zuriick und bildet eine seichte Einbuchtung, in welcher das angeschwollene Haustorium seinen Platz hat —’’ and Schneider ’—‘‘ The tip of the haustorium may pass through the algal cell-wall, forming a somewhat expanded filament between the wall and cell-plasm. In its highest development the haustorium, often entering the algal cell, develops a much branched network which encloses but does not penetrate the cell-plasm.”’ Neither in my sections, nor in the living gonidia, in which I found haustoria, could I detect the alleged ‘‘ Einbuch- tung’’ of the protoplasm, and certainly the haustoria do penetrate the protoplasm of the gonidial cells of A. reticu- lata, as the figures convincingly show. It may be that, at the time of penetration, the protoplasm may contract away from the intruding haustorium, but this condition is not permanent, the haustorium penetrates the cell, the essential and living part of the cell, as certainly as it penetrates the lifeless cell-wall. Can one imagine that the presence of another body, whether living or lifeless, within the living protoplasm of a cell is not accompanied by profound disturbance of the living protoplasm, is not distinctly irritating to it, is not positively injurious to it? The contrast between the evidently healthy hyphe and haustoria on the one hand, and on the other the haustoria-containing gonidial cells, many of which in sections and in free preparations had contents dead and shrunken or completely gone, at least suggests very strongly, though it does not prove, that the haustoria completely devour the contents of the gonidia. It does not necessarily follow that the penetration of chlorophyll-containing host-cells by haustorial cells is fatal to the host-cells. This is the case, for example, in the host- 1j.c., Pl. Land p. 445. 226 CALIFORNIA ACADEMY OF SCIENCES, [Proc. 3D SER. plants of Cuscuta (Peirce 1893), but no one would venture to intimate that this phanerogamic parasite, containing traces of chlorophyll and therefore less incapable of elaborating non-nitrogenous food than any fungus, does not distinctly injure every cell into which its haustorial cells penetrate. In the lichen, each green cell penetrated by a haustorium is obliged to manufacture non-nitrogenous food in quantity sufficient to supply both the haustorium and itself, and not only for the haustorium but also for the cell of which it is a branch and for the cells adjacent to it. How long cana green cell do this unusual amount of work? Certainly — not for so long as it could do less work. The life of the individual cell is therefore shortened. Overwork, exhaus- tion, death, and finally the complete absorption of its con- tents, follow the penetration of a haustorium. Is the presence of the haustorium so beneficial or so pleasant as to compensate the gonidial cell for its increased work and and decreased span of life ? Turning once more to Cuscuta and its host for compari- son, we find that both host and parasite are multicellular, that owing to the division of labor among the cells one cell can help another. For instance, if the photosynthetic or other food-elaborating activity of a green cell is reduced by the intrusion of a haustorial cell, the former will be sup- plied by its neighbors, in accordance with the laws of diffusion and osmosis, with those substances which it lacks. This is an inevitable physical as well as a regular physio- logical phenomenon, the result of the multicellular condi- tion of the host. Thus the life of the penetrated cell, fed by its neighbors, may be prolonged; the struggle between host and parasite is more nearly equal, each haustorial cell draws upon more than one cell of the host to supply the more than one cell of the parasite. This last is also true of the fungus in the lichen, for there are many gonidial cells. But the gonidial cells are distinct from one another; the fungus is a multicellular plant, with the advantages of one; the gonidia— actually in Ramalina and in many other lichens, vrtuad/y in all—are unicellular plants with their Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 227 corresponding disadvantages. The single gonidial cell must do everything itself or perish. The individual gonidial cells penetrated by haustoria are no better off as individuals than algal cells of the same species outside the lichen: they are obviously worse off. Beside the gonidial cells penetrated by haustoria, there are a much larger number, to be seen both in sections and in free preparations, which are invested more or less closely either by the hyphe themselves or by short branches there- from. These last Schneider (1896a@) calls ‘‘ extra-cellular haustoria.’’ The true haustoria, of which we have just been speaking, he terms ‘‘intra-cellular haustoria.’’ Both are branches of hyphez, are therefore morphologically the same; both are organs of nutrition, are therefore physiolog- ically the same; but it seems hardly desirable to give to organs which do not penetrate, but only clasp, the same name which DeBary (1884) and so many after him have used with the original meaning only—‘‘in’s Innere der Zellen dringende Seitenzweige.”’ Whether we have to do with naked or enclosed masses of living protoplasm, their absorption of food is by the same means—by osmosis. The osmotic absorption of aqueous solutions of food substances into cells enclosed by thin cellulose walls is nearly if not quite as rapid as with naked cells. Through a thick cellulose membrane or through two cellulose membranes which, though closely applied to one another, are thicker than only one, osmosis is slower than through a thin cellulose membrane. Through a gelatinized membrane osmosis is also slower than through a cellulose membrane of the same thickness (Pfeffer 1897). From these considerations we see the means by which both the haustoria penetrating, and the hyphal branches invest- ing, the gonidial cells obtain from them the nutrient solutions which the fungus must obtain from them or die. If the hyphe become gelatinized as to their walls, their absorp- tion of food will be diminished equally with their loss of water. The production of haustoria will, therefore, facili- tate the absorption of adequate amounts of food. In many 228 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. lichens, however, haustoria are never formed; the hyphal branches investing the gonidia osmotically absorb enough food to supply the whole body of the fungus. The principles underlying the absorption, and the results of the absorption, of food by the hyphe from the gonidial cells are identical whether haustoria penetrate or hyphal branches only clasp the gonidial cells. Microtome sections confirm the observations regarding the relative sizes, the number and kinds of division, the general appearance of the gonidial cells invested by and free from hyphe, as reported in the foregoing chapter. Both the fixed and the living specimens show that fungus and alga are associated intimately enough, whether the fungus merely encloses or actively penetrates, for the organism absolutely unable to elaborate non-nitrogenous food to be able to absorb all that it needs from the one that can elaborate it. In both cases the absorption is by osmosis, and it must be almost, if not quite, equally effective in both cases, for the lichens which do not send hyphal branches as haustoria into the gonidia appear to thrive as well as those which do. There must in both cases be irritation of the gonidial cells by the hyphe. The irritation produced by haustoria must, however, be greater, and this conclusion is substan- tiated by the larger number of dead gonidial cells contain- ing haustoria than of those with hyphe merely attached. The observation of haustoria within gonidial cells, though not a step necessary to the conclusion that the fungus is parasitic upon the alga, is confirmatory evidence, for it proves that the fungus has the alga in its power— that it obtains all its food from it, that it irritates and exhausts it in proportion to the intimacy of its relation with the alga. In the other two forms studied —Spherophorus globiferus and an Usnea—microtome sections show that the relation between hyphe and gonidia is intimate, the hyphe closely investing the gonidia. Figure 19 is from a section through an old part of a Usnea, the species of which I did not determine, and shows a hypha, old and thick-walled, still Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 229 closely applied to a gonidial cell which has begun to divide. Figure 20 is from a similar section of Sperophorus and demonstrates the intimacy of contact between the hyphez and a gonidial cell also dividing. I did not succeed in finding haustoria in these two lichens, but from the close- ness of the contact which, as these two figures show, exists between hyphe and gonidia, and from the foregoing dis- cussion of the physiology of this relation, one is certainly justified in concluding that the fungus and the alga stand to each other, in these two lichens as in Ramalina, in the relation of parasite and host. The alge in these three lichens are the same, C'ystococcus humicola, Nag.’ It would be interesting to compare with these forms lichens which have other species of algz as gonidia, but for various reasons I prefer to leave this to others or to later work of my own. V. THE SIGNIFICANCE OF THE WATER-CONTENT OF FRamalina reticulata. The amount of water held, and the force with which it is held, even in very dry air, is significant. The lichen cannot die from drought, and the alga is defended against extreme dryness by the gelatinous enveloping fungus, which holds water enough at least to keep the alga alive, and probably enough to enable it to elaborate some food each day. It is hard to conceive that the alga, supplied with any moisture at all, should be entirely inactive when daily illuminated by the sunlight; and owing to the affinity of the gelatinous outer hyphe for water, the dews and the frequent fogs at night must furnish as frequent additions to the water-content of the lichen. It seems extremely probable, therefore, that the long dry season of California, checking the growth and many other activities of most plants hereabouts, is not suf- ficiently dry to stop the photosynthetic and other activities 1 Professor William A. Setchell, of the University of California, kindly informs me that this alga is now commonly called Chlorococcum humicola (Rab.) Nag. 230 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. of the gonidia of this lichen, much less to kill them; and hence the hyphe, well supplied with food, may also, at least frequently, be active in growth and in other ways. For twelve months in the year these lichens can, and undoubtedly do grow, though the most rapid growth and the most growth must be during the warmer later part of the rainy season. If this is the case with the lichen as a whole, what is the condition of those free algal cells entirely away from and uninvested by fungus hyphe? Throughout a normal dry season on the west side of the Bay of San Francisco, the fogs at night and the dews are sufficiently frequent and heavy to wet fences, tree-trunks, branches, etc., etc. Though this moisture evaporates quickly after the sun appears, I doubt very much that Cyystococcus-like alge are so completely dried by day that they do not revive at night. Indeed, spherical unicellular alge, Cystococcus, Protococcus, etc., although less abundant here on tree-trunks, fences, etc., than in many other parts of the world, can be found alive at all seasons. On almost every fence old enough, one may find the algz alone, or with a few hyphe among them, or with more but still not enough to cover and shield them from dryness, and so on up to mature and fruiting lichens of various species. In this region, at least, the survival of alge like those in lichens is not dependent upon their being protected by fungus hyphe against extreme dryness. It is argued that because lichens are found flourishing where alge alone could not possibly survive — for instance, on dry and sun-baked rocks —that this is evidence that the association of alga and fungus is mutually beneficial. Con- cerning this, 7. reticulata offers a case in point. The alge forming the gonidia of this pendant lichen could not possibly remain alive in the air even if they could float init. Yet, is it any advantage to the gonidial cells of this lichen that they are living, growing, multiplying, where the force of gravitation and the insufficiency of food would forbid alge free from fungus associates to survive? The answer to Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 231 this question will be clearer if we use a homely analogy. A cow would never climb to the top of a twenty-story build- ing, but once elevated to this position in opposition to her ordinary habits and to the force of gravitation, would she be any more advantageously placed than her more common- place relatives in barn and pasture? Though algal cells cannot suspend themselves in mid-air, or live indefinitely exposed to excessive heat and dryness on sun-baked rocks, it does not necessarily follow that they are any better off when associated with fungus hyphe and contributing to the formation of a lichen, than are their more commonplace relatives which are not in the air and have fallen on more advantageous places than exposed rocks. VI. Tse INFLUENCE oF MECHANICAL STRAINS ON GROWTH. As reported in a previous paper on 7. reticulata (Peirce, 1898), the mere wetting of a thallus or thallus fragment, and the subsequent drying, bring about no material change in the length, breadth or thickness. During the rainy season, however, when the lichen remains wet for consider- able lengths of time, growth will take place while the lichen is expanded and soft, thus permanently increasing the dimensions. The lichen will not contract, when it dries again after the rainy reason has passed, to the dimensions it possessed when the rains began; but how much of this increase in size is due to growth solely, and how much to stretching and the consequent change in position of the hyphe, cannot now be stated. It is obvious that this pen- dant lichen, being more than twice as heavy (Peirce, 1898), as well as much softer, when wet than when dry, will be subjected in its different parts to a stretching force vary- ing in the different parts with the weight of the parts nearer the tips. The parts nearest the branch from which it hangs will be subjected to the greatest stretching force or weight, the tip to the least. By stretching force I mean 232 CALIFORNIA ACADEMY OF SCIENCES. [PRoC. 3D SER. in this connection not the forcible expansions which follow the wetting and consequent swellings of a dry lichen, but the actual stretching strain which the distal parts exert by their own weight upon the basal upper parts, a strain which increases with the wetness and with the actual increase in substance. Once thoroughly wet, the lichen will not absorb still more water, but it may grow. We must distinguish, then, between growth which, in this connection at least, is a purely physiological process, and stretching, which is a purely mechanical one. The growth involves an increase in the size of the cells composing the lichen, is made possi- ble by accumulated foods elaborated by the gonidia and hyphe, and is very quickly followed if not accompanied by an actual increase in the weight as well as in the volume of substance either itself living or at least elaborated solely by the living substance. The pull exerted upon itself by the wet and heavy lichen is parallel with the long axis of the thallus; it is stretched lengthwise, while growth takes place in breadth and thick- ness as well as in length. Hegler (1893) has shown for higher plants that a pull exercises two distinct influences; the mechanical one of stretching, the physiological one of irritating —a pull entirely insufficient for the former being in many cases quite sufficient for the latter. The irritation produced by pulling retards growth in the direction of the pull, stimulating it in other directions both as to the individ- ual cells and also as to the tissues and organs as a whole. A pull so light that it does not stretch a stem or branch will lower the rate of elongation, raise the rate and degree of thickening of the walls, especially of those cells which con- tribute most to the mechanical strength of the plant, and in proportion to the actual increase in length there is more than the usual increase in diameter, by thickening the walls of already existing cells as well as by inducing the forma- tion of new cells. Without having made any measurements—a matter requiring infinite patience on account of the low rate of Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 233 growth and because of the many sources of error to be eliminated —and merely from a study of the structure of this lichen (Peirce, 1898), I think we may justly infer in the light of the foregoing discussion, that the increase in length of 7. reticulata is due in greater measure to that stretching of the thallus, by its own weight, which results in bringing the hyphe from their diverse directions into a course nearly parallel with the long axis of the thallus (i. e., parallel with the pull) than to actual growth. The physiological process of growth in length, made less neces- sary if not retarded by the mechanical pull to which this lichen is constantly subject —a pull which is greater when the lichen is softest and weakest —is accompanied by growth in breadth and in thickness and by an increase, by thickening walls, in the mechanical strength of the hyphe. These processes are at least not hindered and, judging from Hegler’s work on higher plants, are likely to be stimulated by the longitudinal pull. Asa result of the growth in thick- ness and breadth, the normal proportions of the thallus will be maintained and, furthermore, more hyphe will be formed which, running at various angles to the long axis, may by the pull be drawn into courses parallel with it. But Hegler has shown one other thing interesting in this connection, namely, that changes in the pull also affect the rate of growth, and that if the pull remains constant for a time the plant will recover its normal rate of growth in length, growing in diameter and increasing in mechanical strength in proportion to the force to which it is subjected. The decreasing weight of our lichen on drying, the gradual diminution of the pulling force, will stimulate it to some growth in length; but because the lichen stiffens as it dries, it cannot grow much at this time, and hence one can con- ceive of little more growth taking place than will suffice to fix the hyphz in the direction into which they have been pulled. Since the pull does not remain constant in nature, winds and varying amounts of moisture forbidding this, the lichen cannot become accustomed to any one amount of 234 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. force, cannot resume its normal rate of growth if, indeed, it may be said to have any one normal rate, though it must and does have normal rates. VII. NucLEeus OR PYRENOID AS THE CENTRAL Bopy OF CystTococcus CELLS. There remains one more point to be discussed in this paper. In two papers which I have been unable to see but which are briefly reviewed in Just’s Jahresbericht (Bd. XXII, 1, p. 148), Dangeard (1894) claims that the body generally regarded as the nucleus of gonidial cells is not nucleus at all but a pyrenoid, the real nucleus being at the side, small, inconspicuous, and formerly supposed to be a vacuole. Differential staining agents are alleged to prove Dangeard’s assertion. As the species of gonidia investigated by Dangeard are not mentioned in the review, I cannot criticise his results and can only advance evi- dence to show that the central body in the cells which serve as the gonidia of the lichens reported upon in this paper, is certainly a nucleus, as the well known nuclear stains and the phenomena of division and cell-division plainly indicate. In a spherical cell, too, one would cer- tainly expect to find the nucleus near if not at the center of the cell under ordinary conditions. In no case have I found a nucleus-like body far from the center of the gonidial cells of the lichens which I have studied. One would expect the nucleus to have the size in proportion to the diameter of the cell which this central body possesses, and not to be small, eccentric and inconspicuous. The form of the nucleus would be likely, for mechanical reasons, to approach that of the cell, though of course this is by no means always the case even in spherical cells. Pyrenoids occur always in chromatophores, as specially differentiated parts of them, not as independent organs. In fig. 16 we have a typical gonidial cell in a thin micro- tome section of A. reticulata, stained on the slide with Iodine- green-Fuchsin. The central body, enclosed in BoT.—VOL. I.] PEIRCE—NATURE OF LICHENS. 235 cytoplasm in which the large rounded chromatophores are imbedded, is dense, granular and provided with a central homogeneous body of its own which, like nucleoli in general stained with this agent, has a deep red color. Figure 17 shows a gonidial cell in the same section in pro- cess of internal cell-division. For the sake of clearness, protoplasm, chromatophores and vacuoles are omitted from the drawing. Each daughter-cell, though not yet provided with its own cellulose wall, contains a dense, granular central mass, from which the nucleolus-like body is absent. This is what one would expect if the central mass is nucleus, for the nucleoli ordinarily disappear before division, reappearing in the daughter-nuclei only after the lapse of some time.’ Figure 18 is still another gonidial cell in the same section, which has divided still further, the eight daughter-cells, most of which are below the plane of the drawing, being already surrounded by their own cellu- lose walls and each containing its own central body. From these figures, from the behavior of the nucleolus- like body both toward the stain and also in division, from the different staining of the central body, the cytoplasm and the chromatophores, and from its position, as far removed _ as possible from all the chromatophores, I cannot do other- wise than conclude that it is a nucleus. Figures 19 and 20 show the nuclei of a gonidial cell of Usnea sp. (?), and of Spherophorus globiferus, respec- tively, dividing and divided. These figures were made from thin microtome sections stained with Anilin-safranin- Gentian-violet-Orange G. The colors, though faint, differ- entiate nucleus and cytoplasm in the manner characteristic of this stain. From the evidence it seems to be clear that the gonidia of these lichens have a central nucleus neither remark- able nor different from. the nuclei of other similar alge whether within or outside of a lichen thallus; that this nucleus is provided, in the ‘‘resting condition,’’ with a typical nucleolus which disappears before the division of the nucleus and the cell; that nuclear division precedes 236 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. cell-division. As to additional details of the processes of nuclear and cell-division, the small size of the objects makes study very difficult, and because this has no direct bearing on the main problem discussed here, I leave the matter as it now stands. VIII. Summary. The results of this examination of a few forms, though admittedly of too few and too like forms to justify anything more definite than an opinion regarding lichens in general, may still be of value as a contribution to the discussion of the relation of fungus and alga in lichens and of the so-called autonomy of lichens. These results may be briefly stated as follows :— I. Both cultures and thin microtome sections demonstrate 1—that hyphez and gonidia are in the most intimate contact; 2—that the hyphe develop branches which may merely clasp the gonidial cells or may, as definite haus- toria, penetrate them; 3—that such clasping or penetration stimulates the gonidia to internal cell-divisions in the effort to form individual cells free from hyphal investment; 4—that the haustoria consume the protoplasmic con- tents of the gonidial cells which they have entered, leaving only the empty cell-wall. II. 1—Since the fungus, being devoid of chlorophyll, must obtain already elaborated non-nitrogenous food; 2—-since the only constituent of the lichen capable of elaborating non-nitrogenous food is the gonidia; 3—and since lichens ordinarily grow where they could obtain little or no non-nitrogenous food from the substratum even if they were dependent upon it for other than mechanical support and mineral salts, —it is obvious that the fungus is fed by the alga; the hyphez, by the gonidia. Bot.—Votz. I.] PEIRCE—NATURE OF LICHENS. 237 III. Though alge may not always grow independently where they are found associated with fungi, forming lichens, it is neither logical nor sensible to conclude that their unus- ual position is beneficial to them. IV. Though the percentage of water retained by the more or less gelatinous constituents of the lichen thallus is higher than that which the gonidia could retain as free organisms, the occurrence of healthy alge on the spots subsequently occupied by lichens demonstrates that free alg can thrive, at least for a time, wherever lichens can. V. Though it may be claimed that the individual gonidial cells live longer in the lichen than the free individual cells of the same species of alga, there is no proof of this, and if there were, it is well known that in their resting forms free alge withstand extremes of heat, dryness, etc., as suc- cessfully as do lichens and lichen gonidia. VI. There is no proof that algal cells serving as lichen gonidia are any better off as to food, protection or situation than the average free algal cells of the same species; whereas it is evident that the fungus portion of every lichen is absolutely dependent upon the gonidia for all of its non- nitrogenous food. VII. As to Reinke’s claim that the lichens should be regarded as plants as truly autonomous as trees and shrubs because they have their own peculiar structure, growth, habit, color, etc., as well as because there can be no lichen without both components, an analogy may assist us to clearer views. Bacteria grown in bouillon cannot form the definite colonies of peculiar and characteristic structure, growth, habit, color, etc., which the same species will form on nutrient agar-agar or gelatine. To the formation of the colony, with its characteristic peculiarities, the living bac- teria and the lifeless but nutritious solid substratum are both absolutely necessary. The bacteria may grow quite 238 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. as well in a liquid medium as upon a solid one, but they will not aggregate into the same form. So the fungus component of a lichen, grown by Moller on liquid media and alone will not form a body like a lichen in structure, growth, habit, color, etc., although it will develop perfectly well. Both substratum and associate will affect the fungus, and the lichen ts the product, the resultant, of all these influ- ences, not of one or two. VIII. The varying mechanical pull exerted by itself and by the wind upon A. refzcul/ata influences its growth in the three dimensions of length, breadth and thickness. IX. The central body of the gonidial cells of Ramalina, Usnea, and Spherophorus, which are Cystococcus humicola Nag., is a nucleus, not a pyrenoid. STANFORD UNIVERSITY, CALIFORNIA, December, 1898. Bot.—VOL. I.] PEIRCE—NATURE OF LICHENS. 239 BIBLIOGRAPHY, BARANETZKY, J. Beitrag zur Kenntniss des selbstiindigen Lebens der Flechtengonidien. /ringsheitm’s Jahrbiicher f. wiss. Botantk, Bd. VII. BENECKE, W. Ueber Culturbedingungen einiger Algen. Bot. Zeitung, 1 Abth., Heft 5. DANGEARD, P. A. In Le Botanist, ate Sér., fasc. 1 and 2, and Comptes rendus de l’ Academie des Sciences Naturelles, Paris, T. CXVIII. -. Just’s Jahresbericht, Bd. XXII, I, p. 148. DeBary, A. Die Pilze, p. 19. Morphology and Biology of the Fungi, Mycetozoa, and Bacteria, p. 398, Oxford. FiscHER, A. Neue Beitrage zur Kritik der Fixirungsmethoden. Anatomischer Anzeiger, Bd. X. HEGLER, R. Ueber den Einfluss des mechanischen Zugs auf das Wachsthum der Pflanze. Cohn’s Beitrage zur Biologie der Pflan- zen, Bd. VI. Also report of HEGLER’s work by PFEFFER in Ber. d. K. Sachs. Ges. d. Wissenschaften: Math.-Phys. Classe, Sitzung 7 Dec., 1891. HEDLUND, J. T. Vorlaufige Mittheilung tiber Flechten. Aotantsches Centralblatt, Bd. LXIII, p. to. M6.ter, A. Ueber die Cultur flechtenbildender Ascomyceten ohne Algen. Jnaug. Diss., Minster. Peirce, G. J. Haustoria of Phanerogamic Parasites. Annals of Botany, Vol. VII, p. 308. On the mode of Dissemination and on the Reticulations of Ramalina reticulata. Botanical Gazette, Vol. XXV, No. 6, Pp. 409. Fixing and Imbedding Lichens. Journ. of Applied Micros- copy. Vol. I, No. 6, p. 99. PFEFFER, W. Handbuch der Pflanzenphysiologie, 2 te Auflage, Bd. I, p. 100. REINKE, J. Die Stellung der Flechten im Pflanzensystem. Prings- heim’s Jahrbiicher f. wiss. Botanik, Bd. XXV. (The literature of the subject is here cited). SCHNEIDER, A. Reinke’s Discussions of Lichenology. Bull. Torrey Bot. Club, Vol. XXIII. Text-book of General Lichenology. STrauL, E. Beitriige zur Entwickelungsgeschichte der Flechten. Leipzig. (Not seen, quoted by DeBary). ZIMMERMANN, A., trans. by HumpHREY, J. E. Botanical Microtec- nique, p. 187. 240 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XLI. The figures were drawn with Abbé camera over Leitz No. 7, or Zeiss 2 mm. apochromatic oil-immersion objectives, and Leitz eye-piece 3 or Zeiss compensating eye-pieces 6 and 12. Ramalina reticulata. Fig. 1, @and 6. Ungerminated spores in water: a@, smaller than the average and probably unripe; 4, larger than the average. x 1300. Fig. 2. Germinated spores, showing breaking up of the oil drops and their re-formation in the germ-tubes ; both cells of each spore germi- . nating. x 1300. Fig. 3. Germinated spore; only one cell putting out germ-tube, but this branched. x 1300. Fig. 4. Germinated spore ; germ-tube growing close to and straight past a healthy gonidium. Is actual contact needed to produce stim- uli? x 650. Fig. 5. Group of invested gonidia and one uninvested gonidium separated by grinding ; showing relative sizes of attached and unattached gonidia. x 650. Fig. 6. Group of invested gonidia set free by grinding; showing gonidia dividing internally into two. x 560. Fig. 7. An invested gonidial cell dividing into four. x 560. Fig. 8. A similar one showing four enclosed daughter-cells. x 560. Fig. 9a. The same cell as fig. 8, twenty-four hours later, showing eight daughter-cells liberated by solution of wall of mother-cell. X 560. Fig. 9. A gonidial cell containing a haustorium; the protoplast contracting and thereby escaping from the haustorium. x 560. Fig. 10. Similar to fig. 9*. x 560. Fig. 11. Group of invested gonidia, showing two haustoria within gonidial cells. x 650. Fig. 12. Isolated gonidium containing haustorium ; gonidium isolated and hypha broken off by grinding. x 1750. Fig. 13. Gonidium, in a section, showing haustorium and the hyphal cell of which it isa branch. 12 and 13 comparatively recently pene- trated by haustoria. x goo. Fig. 14. Gonidium, isolated by grinding, entirely emptied by haustorium. X 1750. Fig. 15. Gonidium, in a section, entirely emptied by haustorium and by investing hypha. x goo. Fig. 16. Unusually large and healthy gonidial cell, free from investing hyphe and showing chromatophores enclosed in_ peripheral cytoplasm ; the central nucleus with its nucleolus and enclosing cytoplasm with radiating strands. x 1750. Fig. 17. A gonidial cell dividing by internal cell-division into four daughter- cells, each with its own central nucleus; daughter-cells not yet surrounded by their own cell-walls. x 1750. Fig. 18. Gonidial cell divided by internal cell-division into eight daughter- cells, each with its own central nucleus and its own cell-wall. X 1750. Usnea. Fig. 19. The central body, nucleus, of a gonidial cell dividing, preliminary to the division of this invested gonidial cell. x 1750. Spherophorus globiferus. Fig. 20. Somewhat later stage in the division of the central body, nucleus, of an invested gonidial cell. x 1750. Proc. Can Acan.Sc1.3" Ser. Bor. Vor] [FerRce] Pirate XI LITH.BRITTON & REY, 5.F. CALIFORNIAN HYPOGAZOUS FUNGI. BY H. W. HARKNESS. PLATES XLII-XLV. INTRODUCTION. For several years the author has contemplated the prep- aration of a monograph upon the Hypogzi of California, and whenever it was possible, search was made in order to secure specimens for the purpose. [III] health has, however, compelled the suspension of work for periods of many months duration, and has also prevented excursions for collecting material at seasons which are unsuited to the invalid. I need not remind those who have given attention to this line of investigation that such a collection could only be brought together by long and persistent effort. Unless by accident, the material is obtainable only by removing the surface of the ground, the common garden hoe being the implement best suited for the purpose. As there are but seldom any surface indications to mark the spot where the tubers grow, it will readily be seen that their collection depends largely upon chance. Expe- rience, however, teaches the collector to seek such loca- tions as are best suited to the growth and development of the desired material; but, with all his knowledge, it is only by persistent effort that he may succeed. It has been too frequently the case that the writer has expended hours of arduous labor without securing a single specimen. The district within the limits of which excursions have been made is bounded upon the north by the California State line, on the south by the Tehachapi range, by the sea- coast on the west, and the valley of the Donner lake upon the east—an area exceeding 400 miles from north to south, {241 ] June 26, 1899. 242 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. and some 300 miles from east to west, and within which are to be found the Coast Range mountains, with the dense forests of Sequoias, and the Sierra Nevada mountains ris- ing to an elevation of 8,000 feet. As will readily be seen, excursions for exploration to localities so remote must be, owing to unfavorable weather and other causes, too often barren of results. Many species may be found soon after the first autumnal rains, especially if the rain is followed by a period of sunshine and moderate heat. If these conditions continue during the entire rainy season much material may be collected during the winter. The most productive season, however, is that of early spring, as it seldom fails during these months that there are warm rains followed by sunny days. If, as it some- times happens, there is an abundant precipitation of moist- ure, good material may be found even late into the spring. After the close of the rainy season but little is to be found, unless it be upon the banks of mountain rivulets, or in a few favored spots where there exists sufficient moisture combined with a suitable soil. There being no visible indications to mark the spot where the fungi abide, to assure success in the search the characters of the trees and shrubs in the locality must be observed, as well as the nature of the soil. At times the fungi may be found beneath and amidst dense masses of decaying foliage of the Sequoias. When so found they will be in the immediate vicinity of the base of the tree. So far, however, as the writer’s observations extend, the time during which the fungi are to be found among the Sequoias is of but short duration, as the mature plants are eagerly sought for by the rodents which are found in numbers in these forests. ‘The most active of these foragers, and con- sequently the most destructive, are two species of wood- rats (/Veotoma). Where these are to be found one must be upon the alert if he would secure mature specimens of the fungus, as the active little fellows are tireless in their search for this dainty. Squirrels and other rodents are but little less active and add to the difficulties of the collector. Bot.—VOL. I.] HARKNESS—CALIFORNIAN HYPOG@I. 243 A few tubers are to be found in the high Sierras at an elevation of 7,000 feet or more; these are generally located at or near the base of Pinus contorta, and occasionally beneath other species of Conifer. So far as my observa- tions extend, this section can be relied upon for specimens only in the late spring, as the snow generally falls too early in the autumn to allow of their development. At a lower elevation—3,000 or 4,000 feet—we find the oaks, under which, at the proper season, good material may be obtained. At these elevations the banks of the streams may yield material as late as July. In the foot-hills, at an elevation of from 1,400 to 1,500 feet, we find the best local- ities for the greatest number of species, which are here most prolific. Upon well drained hillsides, in sandy soil, a variety of Ceanothus is found growing in dense clusters and averag- ing about seven feet in height. When protected from fire, we find beneath these shrubs a large accumulation of decay- ing foliage which serves to fertilize and protect the fungi found here. Upon the plains the Eucalyptus tree has been cultivated to a large extent, and here a few varieties of fungi may be found. It may be broadly asserted that but seldom are species to be found in localities where grass or weeds are seen, as the roots of these plants seems to be inimical to the development of the tubers. Neither are they to be found in places where water accumulates or remains for a length of time upon the surface. The earliest date at which we may hope to find tubers (truffles so-called) is about the first of January. At this time the cell-structure of the glebais in a perfect state but is still destitute of asci or spores, which makes the identifi- cation of species impossible. So far as I have seen, the spore does not arrive at maturity until April; much, how- ever, depends upon the weather. In any event, the tuber is of slower growth than many others of this class of fungi. The genus 7uer is widely distributed throughout California but is exceedingly scanty in numbers. 244 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. As to the economic value of the Hypogei of California: so far, we have found none of the edible varieties of the Continent of Europe, although Zuber Californica is nearly identical with an edible species found in Italy. All of the Californian species are, however, edible, and no doubt would be greatly esteemed as a luxury were it not for the fact that they are so rare as to practically prohibit their use as food. The writer has but recently discovered reliable traces of a variety which, if found in quantity, is certain to take its place as a table luxury, although as yet he has not been able to obtain a single specimen. Some years ago the citizens of Marysville discovered large quantities of an earth fungus which was growing in the vicinity of the city and which was freely eaten by those who were so fortunate as to be aware of its value. In that city they were known as the potato mushroom. A trust- worthy gentleman states that they never appeared upon the surface of the ground, but that the search for them was a comparatively easy matter. The first noticeable sign of their presence was a circular space a foot or more in diam- eter, which was free or nearly so from vegetation. A more careful examination showed minute cracks or fissures upon the surface of the ground, and a moment’s work with trowel or hoe sufficed to unearth a number of white, globose fungi, varying in size from an English walnut to that of a small orange. My informant states that they were gathered with the greatest facility. Another gentleman states that they were found by himself and friends in the vicinity of Sacramento, where they were highly esteemed as a deli- cacy. His description of the method pursued in their col- lection and of the surface indications marking their pres- ence was identical with that of the observer above men- tioned. Neither of the gentlemen could say positively in what month the fungus was found, but both agreed that it was late in the winter or early spring. This will undoubtedly prove to be a Zerfezza, a variety of which is found in northern Africa and also in great abundance in Arabia, being sold in the markets of Bagdad BoT.—VOL. I.] HARKNESS—CALIFORNIAN HYPOGZ@1. 245 as truffles. A TZerfezza has recently been discovered in the valley of the Red river of Louisiana; this was recently distributed by Mr. Ellis in his ‘‘ Centuries’? (No. 1728) as Terfezia leonts. To what extent they are utilized as food in that locality we are not informed. Should the fungus again appear in the Sacramento valley in the same abund- ance as upon previous occasions, it will be sought for, as it is not only of value when fresh, but any excess of the product may be dried and would be an addition to any soup, , and of value for many culinary purposes. _ With these preliminary observations, which I trust may be of service in showing the methods pursued in the quest of material and its geographical distribution, I submit this monograph to the verdict of those who may be interested in this department of botany. I should be lacking in courtesy did I fail to give due credit to my friend, Dr. Gustav Eisen, who has prepared the accompanying illustrations. The obligation is increased when I take into consideration the fact that his time was fully occupied in an entirely different line of biological in- vestigation. Hymenogaster V7it. fymenogaster VitT., Monog. Tub., p. 20. Peridium fleshy or thin, running down into an absorbing base. Cavities at first empty, radiating or irregular. Trama composed of elongated cells, but not of byssoid flocci, and therefore not easily separable. Spores various. (Berk. Outlines Brit. Fung., p. 295.) 1. Hymenogaster versicolor, sp. nov. Subglobose, 2 cm. in diam.; color white turning to pink; common in- tegument thick, corrugated, flakey externally, pinkish beneath, closely adhe- rent; gleba firm; cavities sinuous, minute; spores ovate, attached by a slender but somewhat elongated pedicel, 5x8 yp. Type, No. 174, Harkness Coll. Under small oaks, Bishops, Mill Valley, Marin County, Calif., April. 246 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. 2. Hymenogaster Setchellii, sp. nov. Minute, 1.5 cm. in diam.; color white turning to brown, subglobose, smooth; gleba buff, elastic; cells large, sinuous; sterigmata elongated; spores citriform, guttulate, brown, 7x9 p#. Type, No. 165, Harkness Coll. Under Vaccinium, beneath vegetable humus, Mt. Tamal- pais, Marin County, Calif., April. Named in honor of Professor William A. Setchell, of the University of California. 3. Hymenogaster utriculatus, sp. nov. PLATE XLII, Fics. 6a-6/. Large, roundly gibbous, 2 cm. in diam.; color chocolate-brown; smooth, with slightly concave depressions upon the surface; gleba brown; cells gyrose or subrotund; septa fleshy, elastic, fibrose; spores ovoid, upon a cylindrical pedicel 3 u in length, inclosed within a winged utricle, color citron-brown, 6x10 pL. Type, No. 244, Harkness Coll. Among Sequoias and oaks, Mill Valley, Marin County, Calif., July. The spores of the Hymenogaster are frequently inclosed in a semitransparent utricle; such investment is, however, so far as the writer has observed, uniformly saccate or sub- rotund in outline. 4. Hymenogaster ruber, sp. nov. Oblong, 2 cm. in diam., rugose; peridium pale red, thick, fleshy; gleba brown, septa white; cells minute; spores briefly obovate, rough, 6x8 4. Type, No. 248, Harkness Coll. In the forest, Mill Valley, Marin County, Calif., July. 5. Hymenogaster globosus, sp. nov. Minute, globose, I cm. in diam.; color dirty white; gleba fuscous or nearly black; cells irregular; spores ellipsoidal, 6x 12 p. Type, No. 246, Harkness Coll. In damp ground, beside a rivulet, Mill Valley, Marin County, Calif., July. Bot.—Vot. I.] HARKNESS—CALIFORNIAN HYPOGZI. 247 6. Hymenogaster candidus, sp. nov. Oblong-cylindrical, 3 cm. in diam.; color white; gleba ochraceous; cells large; spores elliptical, guttulate, color brown, 10x 5 4. Type, No. 49, Harkness Coll. Under Pseudotsuga Douglassit, Towle, Placer County, Calif., May. Differing from A. A/otzzi in the form of the fungus and in the shape of the spores. 7. Hymenogaster luteus V77t. Hymenogaster luteus Vitt., Monog. Tub., p. 22, Tab. III, fig, 9. Peridium very thin, soft and silky, white, then brownish, bright yellow within; spores even, ovate or elliptic, oblong, yellow. (Berk. Outlines Brit. Fung., p. 295.) No. 12, Harkness Coll. Under decaying wood, Oakland, Calif., December. 8. Hymenogaster calosporus 7w/. Hymenogaster calosporus Tuu., Fungi Hypo., p. 70, Tab. X, fig. 4. Globosus sat irregularis, szpius depressus aut costato-sulcatus (subtus preesertim), ex albido brunneus et sordidus; peridio tenui vix solubili, humido; lacunis inzequalibus-absque directione, vacuis; septis linea media obscuriore notatis nec scissilibus; pulvinulo subnullo; sporis lanceolato-acuminatis, satu- rate brunneo-rubiginosis, levibus. No. 48, Harkness Coll. In the forest, Towle, Placer County, Calif., May. 9. Hymenogaster muticus B. & Pr. Hymenogaster muticus B. & Br., Ann. & Mag. Nat. Hist., 2d Ser., Vol. II, p- 267. Globose, quite white when young, then tinged with brown and cracked, pale yellow-brown within; spores obovate, oblong, very obtuse. (Berk. Out- lines Brit. Fung., p. 295.) No. 64, Harkness Coll. Under Sequoias, Taylor’s Mills and Mt. Tamalpais, Marin County, Calif., March, ; 248 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 10. Hymenogaster lycoperdineus V7zz. Hymenogaster lycoperdineus Vitt., Monog. Tub., p. 22, Tab. II, fig. 5. Globoso-difformis; peridio albo-fuligineo, sericeo-levi; carne molli elastica dilute fuliginea; cellulis majusculis irregularibus, e basi ad centrum seriatim directis; sporis fusiformibus, pallide fuligineis. (Tul. Fungi Hypo., p. 64.) No. 72, Harkness Coll. Under oaks, Camp Taylor, Marin County, Calif, July. 11. Hymenogaster arenarius 7w/. Hymenogaster arenarius Tut., Fungi Hypo., p. 73, Tab. X, fig. 2. Globosus amorphus obovatus, albidus immutabilis; peridio levi vel inzequali tenuissimo glabro, sicco; cellulis irregularibus, exiguis, parietibus, ferrugineis sporis obrutis; septis albidis subsericeo-nitentibus, tandem aquose obscuris;, sporis minutis citriformibus, in superficie inzequalibus, luteo-brunneis, guttu- lam vix concentricam foventibus. No. 79, Harkness Coll. In the forest, Mt. Tamalpais, Marin County, Calif., March. 12. Hymenogaster pallidus B. & Br. Hymenogaster pallidus B. &. Br., Ann & Mag. Nat. Hist., 1st ser., Vol. XVIII, p. 74. Smaller, rounded, depressed, nearly smooth, white, then dirty tan-color, rather soft, within white, then yellow, then pale brown; sterile base obsolete; spores lanceolate, acute, shortly pedicellate, rather rough. (Berk. Outlines Brit. Fung., p. 296.) No. 81, Harkness Coll. Under oaks, Camp Taylor, Marin County, Calif., March. 13. Hymenogaster caudatus, sp. nov. Large, globose, 5 cm. in diam., color fuscous, rough, caudate; appendages springing from the base; gleba brown; veins olivaceous; spores ellipsoidal, 6x12 pL. The caudal appendage is fleshy, from 1-2 cm. in length by 0.2 cm. in diam., and has a bluntly pointed terminus. The appendage is formed by an aggre- gation of many rootlets which are enclosed in a delicate membranaceous in- vestment. Type, No. 240, Harkness Coll. Beneath Sequoias and oaks, Mill Valley, Marin County, Calif., April. a lf Bor.— Vor. ‘IJ HARKNESS—CALIFORNIAN HYPOGA?1I. 249 14. Hymenogaster olivaceus V7it. Hymenogaster olivaceus Vitt., Monog. Tub., p. 24. Globose, but angular; peridium whitish, then tinged with yellow, rufous when bruised; substance white, then of a dull buff, then rufous-olive, varie- gated with the white trama; spores pedicellate, mucronate, generally smooth. (Berk. Outlines Brit. Fung., p. 296.) No. 167, Harkness Coll. Collected during April, under Seguota sempervirens, at Mill Valley, Marin Co., Calif., and under oaks, at Wire Bridge, Placer County, Calif. 15. Hymenogaster monticolus 7. J/ss. Gautiera monticola Hx., Bull. Cal. Acad. Sci., Vol. I, No. 1, 1884, p. 30. Dark brown, irregularly lobed, 10 cm. in breadth, uniformly about 3 cm. in thickness, nearly plane above and below; stipe short and slender; stroma fer- ruginous brown. basidia apparently two-spored; sterigmata filiform; spores pale brown, elliptic or obovate, apiculate, longitudinally or somewhat obliquely striate, 10-12 by 7-8 ». With the odor of decaying onions. Type. No. 113 (3543), Harkness Coll. Under Seguota gigantea, Mariposa Big Tree Grove, Calif., July. 16. Hymenogaster Bulliardi V77t. Hymenogaster Bulliardi Vitt., Monog. Tub., p. 23, Tab. III, fig. 5. Globosus, ex albido aquilus; gleba firma densa, minutissime cellulosa, demum saturate ferruginea; loculis suboppletis; sporis levibus, late ovatis, breviter obtuseque acuminatis, basi rotundato-obtusis, guttulam crassam sezpius foventibus. (Tul. Fungi Hypo., p. 71.) No. 233, Harkness Coll. Mt. Tamalpais, Marin County, Calif., January. 17. Hymenogaster Behrii De Tonz. Hymenogaster Behrti DE Tont., Syll. Fung., Vol. VII, p. 174. Splanchnomyces Behrii Hx., Bull. Cal. Acad. Sci., Vol. I, No. 1, 1884, p. 30. Cinnamon-brown, irregularly lobed, lacunose, 1-4 cm. in diam.; absorbing base inconspicuous; basidia 2-spored; sterigmata short, filiform; spores very unequal in size, yellowish brown, oval or elliptic, apiculate by the remains of the sterigmata, pitted all over with minute irregular depressions. 10-15 x Io #2. | No. 104 (2911), Harkness Coll. 250 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Growing in vegetable humus, Wildwood Glen, Sausalito, Marin County, Calif., December. i 18. Hymenogaster rufus V7tt. Hymenogaster rufus Vitt., Monog. Tub., p. 23, Tab. ITI, fig. 17. _ Subglobosus; peridio albo-rufescenti subsericeo; basi minuta; carne uni- colore rubro-fusca; cellulis majusculis, irregularibus; sporis obovatis subses- silibus rufis. (Tul. Fungi Hypo., p. 64.) No. 163, Harkness Coll. Under shrubby oaks, Mill Valley, Marin County, Calif., April. 19. Hymenogaster tener Berk. Hymenogaster tener Berx., Ann. & Mag. Nat. Hist., rst Ser., Vol. XIII, p- 349. Small, globose, soft, white, silky; substance pale pink, then greyish-umber; sterile base conspicuous, white; spores broadly elliptic, with a papillary apex, minutely warty. (Berk. Outlines Brit. Fung., p. 296.) No. 11, Harkness Coll. Under oaks, Oakland, Alameda County, Calif. Date not given. Hydnangium Wallr. Hydnangium WAu.tr., in Dietr. Fl. des Keenigr. Preuss., VII, 465. Peridium fleshy or membranaceous. Sterile base none. Trama vesicular. Cells at first empty, then filled with spores. Spores echinate. (Berk. Out- lines Brit. Fung., p. 293.) 20. Hydnangium compactum, sp. nov. Globose, 5 cm. in diam.; color white, smooth; gleba dense, pale orange; cells minute, oblong or ellipsoidal; spores globose, rough, not echinate, white, briefly stipitate, guttulate, 6 # in diam. Type, No. 191, Harkness Coll. Under Ceanothus, Auburn, Placer County, Calif., May. Bor.—Vot. I.] HARKNESS—CALIFORNIAN HYPOGAI. 251 21. Hydnangium album, sp. nov. Globose, color dirty white; peridium attenuate, membranaceous; gleba ochraceous; cells minute; spores spherical, white, briefly echinate, 12 in diam. Type, No. 178, Harkness Coll. In the forest, Calistoga, Napa County, Calif., April and May. Resembling H. candidum Tul., with the exception that the spores are spherical and unusually large. 22. Hydnangium luteolum, sp. nov. Oblong or subrotund, color white turning brown, base not visible; gleba yellowish; cells minute; spores globose, white, briefly stipitate, crowded, echinulate, guttulate, 12 ~ in diam. Type, No. too, Harkness Coll. Found in somewhat sandy soil beneath Libocedrus decur- rens in the mountain region about Alta, Placer County, Calif., July. Closely resembling HY. Carotecolor Tul., except in its inconspicuous base and the form of the spore. Octaviania V7zit. Octaviania Vitt., Monog. Tub., p. 15. Peridium continuous or cracked, cottony, running down into the sterile base. Trama byssoid, easily divisible. Fruit-bearing cavities or cells at first empty. Spores rough. (Berk. Outlines Brit. Fung., p. 292.) 23. Octaviania brunneola, sp. nov. Globose, 5 cm. in diam., common integument smooth, absorbent base dis- tinct, terminating in minute fibrillz; cells various, subrotund or tortuous; basidia 4-spored; spores orbicular, brown, echinulate, 6-10 y» in diam. Type, No. 82, Harkness Coll. Mt. Tamalpais, Marin County, Calif., April. 252 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 24. Octaviania rosea Zk. Octaviania rosea Hx., Bull. Call. Acad. Sci., Vol. I, No. 1, 1884, p. 29. Gregarious, peridium fibrillo-rugose, irregularly lobed, 1-3 cm. in diam., with distinct absorbing base, pale rose color, deepening within; basidia 1-2- spored; sterigmata filiform, capitate, as long as the diameter of the spore; spores globose, hyaline, pale; epispore covered with short, obtuse spines, I4-I7 Us. Type, No. 117, Harkness Coll. Under shrubby oaks, at Golden Gate Park, San Fran- cisco, Calif., January. 25. Octaviania mutabilis Proum. Octaviania mutabilis Roum., Revue Mycologique, Ann. VII, 1885, p. 23. Subglobulosa, alba, tactu vinosa, dein nigrescens, basi fibrillis albis instructa; peridio separabili, tomento fugaci obtecto; gleba primum alba, dein, griseo- brunnea; cellulis irregulariter rotundatis, albidis, szepe interruptis, centralibus majoribus; sporis globulosis, 12-15 in diam., echinulatis, brunneis. No. 138, Harkness Coll. Under Arctostaphylos, Auburn, Placer County, Calif.3 March; Calistoga, Napa County, Calif., March. 26. Octaviania socialis, sp. nov. PLATE XLII, Fics. 5a-5d. Epigzous, large, 8 cm. in diam., surface deeply furrowed, furrows extend- ing to its base; base distinct, with numerous branching threads; gleba rose- pink; cells irregular; spores globose, echinulate, having about ten pointed projections on the circumference, 12-14 # in diam. Type, No. 232, Harkness Coll. In groups upon the surface of the ground beneath Zuca- lypius globulus, Belmont, San Mateo County, Calif., January. 27. Octaviania citrina, sp. nov. Globose, white; common integument flakey, imparting an earthy appear- ance; absorbent base spongy; fibrille wanting; gleba orange, cells irregular, basidia prominent, 4-spored; spores stipitate, brown, globose, echinulate, 10-12 # in diam. Type, No. 157, Harkness Coll. Bot.—VOL. I.] HARKNESS—CALIFORNIAN HYPOGA:1. 253 Collected under Arctostaphylos glaucus at the following localities in California during April: Oat Hill Quicksilver Mine, Solano County; Camp Taylor, Marin County; and Calistoga, Napa County. 28. Octaviania occidentalis, sp. nov. PLATE XLII, Fics. 4a-4d. Large, 2.5 cm. in diam.; color white turning brown, semiglobose; common integument flakey; absorbent base firm, terminating in branching fibrillze; gleba white; cells oblong or subrotund, basidia 4-spored; spores briefly stip- itate, white, globose, echinulate, having about twenty blunt projections on the circumference, 14 # in diam. Type, No. 137, Harkness Coll. Wire Bridge, Placer County, Calif., March. At irregular intervals, a flasked-shaped cystidium which springs from the parenchyma is to be found protruding from amid the true basidia for a distance of 18 » and termin- ating in a conical point. These bodies are filled with what appears to be crystals and are destitute of sporophores. 29. Octaviania compacta 7w/. Octaviania compacta TuL., Fungi Hypo., p. 79. Minor, polyrrhiza, nivea; peridio molliusculo insolubili tomentello-gossy- pino; cellulis rotundis oblongisve, minutissimis, mox obliteratis, oppletis; septis vix conspicuis; sporis innumeris minutissimis, sphzricis, scabriusculis, flavidis, tandem ochraceo-auratis. No. 19, Harkness Coll. Under oaks at Wire Bridge, Placer County, Calif., April. Collected by Chas. L. Phillips. Mill Valley, March. Previously collected at Tamalpais, Marin County, Cah. and Blue Cajfion, Placer County, Calif., under leaves, May- July. 30. Octaviania Stephensii 7w/. Octaviania Stephensii Tur., Fungi Hypo., p. 78. Hydnangium Stephensii Berx., Ann. & Mag. Nat. Hist., rst Ser., Vol. XIII, p. 352, also Vol. XVIII, p. 76. Irregular, oblong, externally rufous, plicato-rugose at the base, cribrose, white within, milky, at length, when exposed to the air, rufous; spores globose, at length echinulate. (Berk. Outlines Brit. Fung., p. 292.) No. 148, Harkness Coll. 254 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Collected at the following localities in California during March: under oaks at Laundry Farm, Alameda County; at Mill Valley and Kents, Marin County; amongst Sequoias at Mt. Tamalpais. Previously collected at Tamalpais, Marin County, Calif and Alta, Placer County, Calif., May—July. 31. Octaviania monticola, sp. nov. PLATE XLII, Fics. 3a-3¢. Large, 3 cm. in diam., irregular, globose, rough, flexible, buff; absorbent base prominent, fibres uniting, with earthy particles enclosed; gleba spongy, chestnut-brown; cells minute; basidia 4-spored, ‘spores ovate, echinulate, IOX12 p#. Type, No. 13, Harkness Coll. Mountain regions in sandy soil, Auburn, Placer County, Calif., April. Hysterangium V77t. Hysterangium Vitt., Monog. Tub., p. 13. Peridium entire, indehiscent, well marked, dissolving readily or splitting off naturally, either thin or somewhat thickish, firm, fibrous or membranaceous, provided with a mycelium; gleba perforated with cells which are at first hollow but are at length somewhat filled up, small, rounded or narrowly linear; par- titions of the cells of very unequal thickness, tough, bearing basidia on both sides; basidia slender, generally 2-spored; spores ellipsoidal. or lanceolate, with short sterigmata, very abundant, smooth and usually pale-colored and pellucid. Fungi of gregarious and hypogzeous habits, of regular spherical shape; polyrrhizous, with the abundant white mycelium which is floccose in a peculiar fashion without being filamentous, either completely or partly cov- ered, or at length attached to a nearly simple, rope-like mycelium and then naked, monorrhizous; with a distinct odor when mature. (Translated from the original. ) 32. Hysterangium cinereum, sp. nov. PLATE XLII, Fics. 2a-26. Oblong-rotund, 5 cm. in diam., color brownish white, elastic. smooth; gleba ashy, cut surface showing abundant veins irregular in outline; spores elliptical, white, 6 x 14 #. Type, No. 31, Harkness Coll. Bot.—VOL. I.] HARKNESS—CALIFORNIAN HYPOGAI. 255 In vegetable humus under Arctostaphylos, Auburn, Placer County, Calif., February. 33. Hysterangium Phillipsii, sp. nov. PLATE XLII, Fics. 1a-16. Large, 3-4 cm. in diam., color rose-pink, fibrillous; rootlets 12 cm. or more in length; gleba of a dark olive or greenish (verdatre) tint, profusely veined, coalescing at the base; cells minute, spores ellipsoidal, in groups of three or four, white, 2 x 54. Type, No. 234, Harkness Coll. Under oaks, Wire Bridge, Placer County, Calif., Janu- ary. Collected by Chas. L. Phillips. 34. Hysterangium occidentale sp. nov. Large, 4 cm. in diam., color dirty white; peridium soluble, subrotund, somewhat flattened or discoid; gleba chocolate-colored; cells gyrose, large; spores ellipsoidal, white, 12 x 74. Type, No. 242, Harkness Coll. Amongst Sequoias, Tamalpais, Marin County, Calif., May. 35. Hysterangium nephriticum Ler. Hysterangium nephriticum Berx., Ann. & Mag. Nat. Hist., rst Ser., Vol. XIII, 1844, p: 350; Tut., Fungi Hypo., p. 82. Depressed, ‘springing from a white, flat, branched, membranous mycelium; peridium firm, elastic, distinct, tomentose; substance pale blue or grey, here and there greenish; cavities radiating from the base; spores minute, oblong, pale clay-color. (Berk. Outlines Brit. Fung., p. 294.) No. 143, Harkness Coll. Under oaks, Auburn, Placer County, Calif., March; Mill Valley, Marin County, Calif., April; Calistoga, Napa County, Calif., May. 36. Hysterangium membranaceum V772. Hysterangium membranaceum Vitt., Monog. Tub., p. 14, Tab. IV, fig. 15. Minus, rotundatum, radicatum; peridio tenui membranaceo albido sub- tomentoso; carne viridiuscula; cellulis difformibus et irregularissimis. (Tul. Fungi Hypo., p. 83.) No. 176, Harkness Coll. 256 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER Mill Valley, Marin County, Calif., April; Auburn, Placer County, Calif., May. 37. Hysterangium stoloniferum 77/. Hysterangium stoloniferum Tur., Fungi Hypo., p. 84, Tab. XI, fig. 8. Sphericum leve glabrum candidum, radice unica funiformi instructum; carne e cceruleo fusca, tenaci; septis crassis; sporis acervatim sordide fuscis, No. 158, Harkness Coll. Collected during April, under oaks, at the following local- ities in California: Calistoga, Napa County; Mt. Tamalpais and Camp Taylor, Marin County; Auburn, Placer County. Previously collected at Tamalpais, March—May. 38. Hysterangium Clathroides V77t. Hysterangium Clathroides Virt., Monog. Tub., p. 13, Tab. IV, fig. 2. Globosum; peridio albido, mycelii gratia polyrrhizo, facile solubili; carne olivaceo-virente. (Tul. Fungi Hypo., p. 80.) No. 156, Harkness Coll. Collected in damp ground at the following localities in California during April: Calistoga, Napa County; Bishops and Tamalpais, Marin County; Wire Bridge, Placer County. 39. Hysterangium australe Seg. | Hysterangium australe SpeGc., Fungi Argent. Pug. IV, n. 237. Primo subglobosum dein ob terre pressionem irregulariter compressum, varie gibbose expansum, magnitudine ludens 5-20 diam., basi manifesta nulla, fibrillis radicalibus perfecte destitutum, album, levissimum, glaberrimum, peridio tenui a pulpa non v. difficile secedente, gleba autem pallide fulvo- olivascente, tremelloideo-subceracea, tubulis numerosis, minutissimis undique irregulariter percursa; tubulis gracilibus 150-250 diam., varie elongatis, vacuis, parietibus sporiferis cinnamomeis; sporis elliptico-elongatis, sursum plus minusve attenuato-rotundatis, deorsum acute attenuato-cuneatis basique truncatis, episporio ubique majuscule undulato-subverruculoso, saturate oliva- ceo-fuligineis, protoplasmate grosse granuloso farctis v. 1-guttulatis 15-20= 8-10 »; stipite longiusculo, gracili hyalino, monospermo fultis. (De Toni in Sacc. Sylloge Fung., Vol. VII, 1888, p. 157.) No. 84, Harkness Coll. Bot.—VOL. I.] HARKNESS—CALIFORNIAN HYPOGA/I. 257 Collected under oaks at the following localities in Cali- fornia during April: Tamalpais and Mill Valley, Marin County; Auburn, Placer County; Calistoga, Napa County. 40. Hysterangium fuscum, sp. nov. Minute, dirty white, globose; mycelium flocculent at base; gleba elastic; veins white; hymenium brown; spores elliptical, 6 x 12 p. Type, No. 177, Harkness Coll. Under Arédutus Menzzeszz, Mill Valley, Marin County, Calif., April. Rhizopogon 7Zz/. Rhizopogon Tut., Fungi Hypo., p. 85, Tab. I, fig. 5, Tab. II, fig. 1, et Tab. XI, figs. 4-5. Peridium continuous or cracked, adhering to creeping branched fibers which traverse its surface. Cavities distinct, at first empty. Spores smooth, oblong- elliptic. (Berk. Outlines Brit. Fung., p. 294.) 41. Rhizopogon aurantius, sp. nov. Subglobose, 2 cm. in diam., color dirty white; peridium attenuate; gleba pale orange, the freshly cut surface showing a creamy exudation; cells large; cell-walls thin but firm; spores subglobose, with colorless oil globule, 8-10 u in diam. Type, No. 74, Harkness Coll. Solitary in dense forests of Sequoias, deeply hidden by decaying vegetation, Mt. Tamalpais, Marin County, Calif., August. Leucophleps, gen. nov. Globose or roundly elongate, color white or citron, dense; gleba multi- locular; cells crowded; veins pearly white; spores spherical or ovoid and borne upon elongated sterigmata. 42. Leucophleps magnata, gen. et sp. nov. PLATE XLII, Fics. 7a-7c. Large, 3 cm. in diam., subglobose or elongate, white, smooth, solid; perid- ium wanting, white; the freshly cut surface sometimes showing a blue tint which soon vanishes; multilocular cells irregularly decreasing in magnitude (2) June 27, 1899. 258 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER. towards the surface and disappearing at the sterile base; veins pearly white; spores single, globose, smooth, enclosed in semi-opaque investments or utricles, with oil globules, supported upon somewhat tortuous sterigmata, 13 # in diam. Type, No. 154, Harkness Coll. Under Acer, Calistoga, Napa County, Calif., April. 43. Leucophleps foveolata, sp. nov. Subglobose, color white or faintly citron; peridium attenuate, minutely pitted; gleba white; veins white; cells rotund; spores white, smooth, guttu- late, 7X12 #. Type, No. 209, Harkness Coll. A second specimen (No. 243) was collected in the same locality. In moist earth beside a rivulet, Mill Valley, Marin County, Calif., July. Differing from Z. magnata in external characters and size of spore. 44. Leucophleps candida, sp. nov. White, 2 cm. in diam., irregular, firm, surface completely studded with depressions of very variable circumference; peridium wanting; gleba of pearly whiteness; cells crowded and plentiful; spores globose, 8 # in diam. Type, No. 207, Harkness Coll. Mill Valley, Marin County, Calif., June. 45. Leucophleps odorata, sp. nov. PLATE XLIII, Fics. 9a-96. Large, 3-4 cm. in diam., color orange, irregularly lobed or oblong, no fibrous attachment, slightly crepitating under pressure. Odor nauseating. Type, No. 251, Harkness Coll. Under oaks, Castle Crag, Shasta County, Calif., July. The marked irregularity in form and color, together with the odor, serves to distinguish this from any other species, and although firm there is distinct crepitus to be observed upon pressure. The cells in the recently cut surface of the fresh plant present a glassy appearance. BotT.—VOot. I.] HARKNESS—CALIFORNIAN HYPOGAI. 259 46. Leucophleps citrina, sp. nov. PLATE XLIII, Fics. 8a-8d. Subglobose, 2 cm. in diam., citron color, smooth; gleba firm, wavy; cells minute; spores roundly elliptical, guttulate, white, two to four spores form upon each basidium; basidia attenuate at the point of attachment, increasing in size towards the apex, oil globules interspersed for its entire length; spores 6x8 p. Type, No. 168, Harkness Coll. Found amidst Manzanitas, Mt. Tamalpais, Marin County, Calif., April. The fungus imparts a red tint to alcohol when immersed. Melanogaster Corda. Melanogaster CoRDAin Sturm’s Deutschl. Fl., Abth. III, Heft 11, 1831, p. 1. Peridium adhering to creeping branched fibres which traverse its surface, without any proper or distinct base. Cells at first filled with pulp. Spores smooth, mostly dark. (Berk. Outlines Brit. Fung., p. 293.) 47. Melanogaster Eisenii, sp. nov. Globose, smooth, 1.5 cm. in diam., color brown; gleba fuscous; veins brown; cells rhomboidal; spores globose, 6-8 » in diam. Type, No. 116, Harkness Coll. Cabo St. Lucas, Baja California, January. Collected by Dr. Gustav Eisen. 48. Melanogaster variegatus 7z/. Melanogaster variegatus TuL., Fungi Hypo., p. 92, Tab. II, fig. 4, et Tab. XII, fig. 6. At first ochraceous, then reddish-ferruginous, minutely downy; walls of the cells dirty-white, yellowish, or orange; pulp black; spores minute. (Berk. Outlines Brit. Fung., p. 293.) No. 145, Harkness Coll. Amongst oaks, Wire Bridge, Placer County, Calif., March. Previously collected at Sausalito, Marin County, Calif., February. 260 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. 49. Melanogaster tuberiformis Corda. Melanogaster tuberiformis CorDA., in Sturm’s Deutschl. Fl., Abth, III, Heft 11, 1831, p. 1, Tab. I. Hypogzeus rotundatus, fusco-cupreus, intus ater, radiculis fibrillosis atris tectus; sporis atris obovatis, deorsum attenuatis. (Tul. Fungi Hypo., p. 95.) No. 5, Harkness Coll. Amongst small oaks, Mill Valley, Marin County, Calif., April. 50. Melanogaster durissimus Cooke. Melanogaster durissimus CooKE., Grevillea, Vol. VIII, p. 94. Subglobosus, compressus, difformis aut sulcatus, levis, durissimus, atro- fuscus, demum nigrescens. Peridio crasso, subnitido; carne mire lacunoso, ochraceo albo; lacunis majusculis, creberrimis, atris. Sporis oblongo-ellip- ticis, inzequalibus, brunneis .oo5-.008 x .003-.005 mm. Odore fortissimo. No. 216, Harkness Coll. Collected amongst oaks at the following localities in Cal- ifornia during April: Auburn and Wire Bridge, Placer County; Mt. Tamalpais and Sausalito, Marin County. 51. Melanogaster aureus 77/. Melanogaster aureus TuL., Fungi Hypo., p. 97. Octaviania aurea Vitt., Monog. Tub., p. 20, Tab. III, fig. 14. Oblongus uniformis, basi radicatus; peridio levi subalbido; carne primo dura aurea, venis albidis (cellularum parietibus) variegata, demum molli et nigrescente; cellulis subcavis. No. 68, Harkness Coll. Amongst oaks, Wire Bridge and Auburn, Placer County, Calif., February. 52. Melanogaster sarcomelas 77z/. Melanogaster sarcomelas Tut., Fungi Hypo., p. 96. Octaviania sarcomelas VitT., Monog. Tub., p. 16, Tab. ITI, fig. 3. Minor, difformis; peridio levi nigro; résticulis nullis; carne nigerrima uni- colore; cellulis regularibus, substantia pultacea sporifera repletis; sporis oval- ibus majusculis, levibus. No. 128, Harkness Coll. Bot.—VOL. I.] HARKNESS—CALIFORNIAN HYPOGA@1. 261 Collected at the following localities in California during April: amongst redwoods, Mill Valley, Marin County; under oaks, Wire Bridge, Placer County; in forest, Laundry Farm, Alameda County. Elaphomyces /Vees. Elaphomyces NEES., in Fries Syst. Myc., Vol. III, p. 21. Common integument thick, hard. Asci globose or obovate. Sporidia con- sisting of several concentric utricles. Internal mass of fungus at length dusty. (Berk. Outlines Brit. Fung., p. 378.) 53. Elaphomyces variegatus V7tt. Elaphomyces variegatus Vitt., Monog. Tub., p. 68, Tab. IV, fig. 4. Mycelium yellow (or yellowish grey), inconspicuous; cortex thick, hard, ochraceous-yellow or golden-yellow, rough, with thick pyramidal and obtuse, or narrow, pointed, and fragile warts, or only granulated; peridium reddish brown and variegated; asci 2-4-spored; sporidia opaque, blackish brown. (Cooke’s Handbook, Vol. IJ, p. 749.) No. 39, Harkness Coll. Under Pseudotsuga Douglassiz, Donner Lake, Nevada County, Calif., July. 54. Elaphomyces Morettii V77t. Elaphomyces Morettiit Vitr., Monog. Tub., p. 71, Tab. IV, fig. 17. Crusta fusco-purpurea; cortice duro, fragili, nigro-brunneo obtuseque ver- rucoso; peridio albido-fusco; sporis fusco-nigrentibus. (Tul. Fungi Hypo., p. I12.) No. 38, Harkness Coll. Under oaks, Santa Cruz, Santa Cruz County, Calif., May. Hydnocystis 7v/. Hydnocystis Tuu., Fungi Hypo., p. 116, Tab. IV, Fig. 7; Tab. XIII, Fig. 2; Tab. XIV, Fig. 1. Receptaculum utriforme, globosufn aut varie sinuoso-anfractuosum, penitus clausum, vel rima basilari inaperta, brevi s. longiuscula et gyrosa, pilisque adpressis confertis fungum intrantibus occlusa et velata quasi dehiscens, in- terius latissime uniloculare et vacuum. Integumentum, receptaculi extima 262 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. corticula, indiscretum, minute papillosum, pilosum et coloratum. Hymenium quod fungi cavitatis paries est, albidum, ex fibris seu filamentis constans long- issime linearibus, subdiscretis aut vix cohzrentibus, parallele e receptaculo prodeuntibus, inzequalibus sterilibusque (paraphysibus), nec non et utriculis immistis paucioribus crassis longe cylindricis obtusis, in filum longum deorsum desinentibus, ascis scil. seu thecis octosporis. Spore uniseriate, sphzricz aut ellipticee, leves, pellucidz, dilute coloratz; nucleo oleoso, tandem homogeneo nec partito. 55. Hydnocystis compacta, sp. nov. PLATE XLIII, Fics. 11a-1I1¢. Minute, reddish brown, subrotund, cavernous without exterior opening, minutely and closely papillose; gleba compact, white; asci cylindrical, obtuse, 175 in length, 8-spored; spores globose, hyaline and unequally papillose, 25 # in diam. Type, No. 98, Harkness Coll. Under Lzbocedrus, Alta, Placer County, Calif., May. Genea V7it. Genea Vitt., Monog. Tub., p. 27. Common integument warty, with an aperture at the apex. Hymenium waved and sinuated, but not forming an intricate mass. Asci cylindrical. Sporidia globose [or subglobose.] (Berk. Outlines Brit. Fung., p. 378.) 56. Genea compacta, sp. nov. PLATE XLIII, Fics. toa-toc. Minute, 1 cm. in diam., irregularly stellate, color light brown, verrucose, dis- tinct base with branching filaments; gleba white, irregularly cavernous; asci cylindrical, stipitate, 270 » in length, 8-spored; spores ellipsoidal, verrucose, 25x18. The fungus is exceedingly rare. The filaments composing the paraphyses are septate, extremely delicate, and of unusual length and wavy. Type, No. 86, Harkness Coll. Found in the forest, Mt. Tamalpais, Marin County, Calif., April. The verrucosity consists in the surface of the spore being covered with small, round, knob-like projections, about sixteen being found in the circumference. Bot.—Vot. I.] HARKNESS—CALIFORNIAN HYPOGA®I. 263 57. Genea arenaria, sp. nov. Minute, color light brown, subglobose, lobed, verrucose, cavernous; gleba white, attenuate; asci linear, 8-spored; spores roundly ellipsoidal, 24 x 18 p; paraphyses brief and not plentiful. Type, No. 42, Harkness Coll. In sandy ground. No locality or date. 58. Genea hispidula Berk. Genea hispidula Brrx., in Tul., Fungi Hypo., p. 121, Tab. XII, fig. 2, et Tab. XIII, fig. 3. Small, brown, externally invested everywhere with rather rigid, adpressed, brown flocci; interior cavity very often simple, with the mouth almost hid- den; radical fibres brown, adhering to the base; spores large, ellipsoid; warts thick and crowded. (Cooke’s Handbook, Vol. II, p. 748.) No. r15, Harkness Coll. Beneath the surface of the ground under trees, San Rafael, Marin County, Calif., May. Under oaks, Wire Bridge, Placer County, Calif., April. 59. Genea verrucosa Vit. Genea verrucosa Virt., Monog. Tub., p. 28, Tab. II, Fig. 7. Very irregular and polymorphous, gibbous, sulcate, or also somewhat many-lobed, black, verrucose ostiolate; ostiola sometimes very broad, rad- ical filaments abbreviated; sporidia broadly elliptic, verrucose. (Cooke’s Handbook, Vol. II, p. 748.) No. 70, Harkness Coll. Amongst decaying leaves under trees, Santa Cruz, Santa Cruz County, Calif., May. 60. Genea spherica 7uw/. Genea spherica Tur., Fungi Hypo., p. 120, Tab. IV, Fig. 2; Tab. XII, Fig. 1, et Tab. XIII, Fig. 6. Regularis et quasi perfecte sphzrica, interdum depressa, atra glabra, verru- cosa, comam ferrugineam siccam spissam longissimamque basi gerens, apice pervia, intus varie labyrintheo-cavernosa, rarius vacua et penitus utriformis. No. 89, Harkness Coll. 264 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. Beneath the surface of the ground under oaks, Contra Costa County, Calif., May. Balsamia V7tt. Balsamia Vittr., Monog. Tub., p. 30. Common integument warty. Hymenium complicated with distinct lacunze not leading to the surface. Sporidia cylindrical or oblongo-elliptic, even, pellucid. (Berk. Outlines Brit. Fung., p. 378.) 61. Balsamia magnata, sp. nov. Subglobose, densely verrucose, 2.5 cm. in diam., color orange red; cavity gyrose; gleba white, firm; asci subovate, 8-spored, 50x38; spores cylin- drical, containing from one to three oil globules, 18 x 8. Type, No. 185, Harkness Coll. In forests, Auburn, Placer County, Calif., May. 62. Balsamia nigrens, sp. nov. Medium, semiglobose, irregular, black, verrucose, warts with polygonal base; openings stellar, sometimes extending through the mass; cavities large and somewhat regular; gleba white, firm, crossed by white wavy lines; asci semiglobose or ellipsoidal, briefly stipitate, 8-spored, 48x 32 4%; spores ob- long-elliptic, guttulate, 26x12. Type, No. 180, Harkness Coll. Beneath Ceanothus, Auburn, Placer County, Calif., May. 63. Balsamia alba, sp. nov. Large, color dirty white, subglobose, fissured, deeply verrucose; gleba firm; asci ellipsoidal, 8-spored; spores cylindrical, guttulate, 12 x 18 yp. Type, No. 129, Harkness Coll. Under oaks, Wire Bridge, Placer County, Calif., Feb- ruary. This fungus is remarkable for the density of the gleba and the small number of its asci. Bot.—VOL. I.] HARKNESS—CALIFORNIAN HYPOG £1. 265 64. Balsamia filamentosa, sp. nov. PLATE XLIII, Fics. 13@-13f Large, oblong or irregularly globose, color ferruginous brown; densely verrucose; gleba filamentous; veins irregular; parenchyma pellucid; asci ellipsoidal, markedly stipitate, 42x24; spores cylindrical, having two to three oil globules, 18 x 12 4. Type, No. 236, Harkness Coll. Under Heteromeles arbutifolia, Auburn, Placer County, Calif., February. The oil globules disappear from view when placed in a medium denser than that of water. 65. Balsamia vulgaris V7ztt. Balsamia vulgaris Virt., Monog. Tub., p. 30, Tab. I, Fig. 2. Major, szpissime sinuoso-exarata vel hinc et illinc excavata, minutissime papillosa, papillis interdum subnullis; lacunis latiusculis gyrosis; septis cras- sis in medio pellucidis; sporangiis paraphyses inter omnino nidulantibus brevioribusque vel breviter exsertis; sporis cylindricis angustis, guttulas oleosas tres subzequales includentibus. (Tul. Fungi Hypo., p. 123.) No. 231, Harkness Coll. Among decaying vegetation in shrubby thickets, Auburn, Placer County, Calif., December. 66. Balsamia platyspora Berk. Balsamia platyspora BERK., Ann. & Mag. Nat. Hist., st Ser., Vol. XIII, 1844, p. 358. Small, globose, rufous, minutely warted, substance pallid yellow, minutely cellulose; sporidia at first broadly oblong-elliptic, with a large globose nucleus, at length slightly elongated trinucleate. (Cooke’s Handbook, Vol. II, p. 747.) No. 222, Harkness Coll. Amongst shrubs under vegetable mould, Auburn, Placer County, Calif., December. 67. Balsamia polysperma V7tz. Balsamia polysperma Vitt., Monog. Tub., p. 31. Minor, anguloso-tuberulosa, papillis minutis congestis ferrugineis vestita; carne subalbida cellulosa; substantia intercellulari (septis) alba opaca; sporis numerosissimis. (Tul., Fungi Hypo., p. 125.) No. 2206, Harkness Coll, 266 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. Under shrubs in sandy soil, Auburn, Placer County, Calif., December. Hydnobolites 7w/. Hydnobolites Tux., Fungi Hypo., p. 126. Integument replaced by white, evanescent down. Hymenium complicated with sinuous lacunz, ending at the surface. Asci elliptic. Sporidia globose. (Berk. Outlines Brit. Fung., p. 377-) . 68. Hydnobolites excavatum, sp. nov. I cm. in diam., color brown; peridium smooth; gleba brown, cavernous; asci subrotund, 8-spored, 65x 54; spores globose, 25 u in diam. Type, No. 189, Harkness Coll. Under vegetable humus in sandy ground, Auburn, Placer County, Calif., May. Hydnotrya B& Br. Hydnotrya B. & Br., Ann. & Mag. Nat. Hist., rst Ser., Vol. XVIII, 1846, p- 78. Common integument minutely papillose, not distinct. Hymenium compli- cated with gyrose lacunz, leading to the surface. Asci oblong. Sporidia globose, tuberculate. (Berk. Outlines Brit. Fung., p. 377-) 69. Hydnotrya cerebriformis, sp. nov. PLATE XLIV, Fics. 19a-19/. Large, 3 cm. in diam., color salmon, subrotund, smooth, brain-like mark- ings upon the surface; gleba white or faintly citron, canals gyrose; hymenium covered with clavate villi between and extending beyond the asci; asci cylin- drical, briefly stipitate; spores globose, brown, foveolate, 25 u in diam. Type, No. 37, Harkness Coll. Among fir trees, Donner Lake, Nevada County, Calif., July. The slight pits upon the spore give it a rough appearance. BotT.—VOL. I.] HARKNESS—CALIFORNIAN HYPOG ZI. 267 Pseudohydnotrya Fischer. Pseudohydnotrya FisCHER, Tuberinez in ENGLER & PRANTL’s Die Naturl. Pflanzenf., Teil I, Abth 1, p. 282. Fungus irregularly rounded, perforated by hollow labyrinth-like passages and chambers which open out at several places on the surface of the fungus, and the walls of which are covered by hymenium. Surface of fungus cov- ered with a pseudo-parenchymatic, hairy peridium, which often penetrates deeply into the interior chambers and then continues directly into the hymen- ium. Hymenium consists of paraphyses and of asci arranged like palisades. Paraphyses cylindrical, septate, at apex knob-like. Asci cylindrical or some- what globular, 8-spored, spores ellipsoidal, smooth, without color, uniseriate or seldom biseriate. (Translated from the original. ) 70. Pseudohydnotrya Harknessii /7scher. Pseudohydnotrya Harknessii FIiscHER, Tuberinee in ENGLER & PRANTL’S Die Naturl. Pflanzenf., Teil I, Abth 1, p. 282. Asci 140-160 4 long, 20-28 » broad, generally 8-spored. Spores 25-28 » long, 14-18 broad. Paraphyses 7-14» thick at the swollen end. (Translated from the original.) No. 1, Harkness Coll. Under shrubs among vegetable humus. Mill Valley, Marin County, Calif., April. 71. Pseudohydnotrya carnea, sp. nov. PLATE XLIII, Fics. 16a-160. Minute, 1.5 cm. in diam., common integument, color pale brown, subglo- bose, irregular, slightly tomentose, chambered; gleba white; parenchyma convolute; asci cylindrical, 125 » in length, 8-spored; spores ellipsoidal, white, guttulate, 22x 15 4; paraphyses hidden. Type, No. 181, Harkness Coll. Among shrubs, Auburn, Placer County, Calif.; May; under oaks, Mill Valley, Marin County, Calif., April. This species is much infested by a parasitic Spheria. 72. Pseudohydnotrya nigra, sp. nov. Large, 2 cm. in diam., color dark brown, inclosed, peridium loosely adhe- rent, tomentose; gleba white; parenchyma convolute; asci 120x8 #; spores guttulate. Type, No, 216, Harkness Coll. 268 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. Under shrubs in firm ground to which it adheres by its hairy investment. Auburn, Placer County, Calif., Novem- ber to April. Stephensia 72/. Stephensia Tux., Fungi Hypo., p. 129. Common integument fleshy, cottony. Base distinct. Hymenium intricate. Asci cylindrical. Sporidia globose, even, at length verrucose. (Berk. Out- lines Brit. Fung., p. 377-) 73. Stephensia bombycina 77/. PLATE XLIV, Fics 18a-18c. Stephensia bombycina TuL., Fungi Hypo., p. 130, Tab. XII, Fig. 4. Genea bombycina Vitt., Monog. Tub., p. 29, Tab. III, Fig. 13, et Tab. IV, Fig. 8; Berx., Ann. & Mag. Nat. Hist., rst Ser., Vol. XIII, p. 357. Subglobose, depressed; peridium rather soft, floccose, irregularly intruded into the cavity, destitute of rooting fibres; flesh gyrose-venose; sporidia pel- lucid, spherical. (Cooke’s Handbook, Vol. II, p. 745.) No. 173, Harkness Coll. Found in forests, March. No locality. This fungus varies slightly from the original description in the spores, which are generally biseriate and marked by irregularly foveolate depressions. Pachyphleus 7z/. Pachyphleus Tur., Fungi Hypo., p. 130. Common integument warty, opening by a terminal aperture. Base distinct. Asci clavate. Sporidia spherical. (Berk. Outlines Brit. Fung., p. 377.) 74. Pachyphleus carneus, sp. nov. PLATE XLV, Fics. 33@-336. Subrotund, 1 cm. in diam., studded with slight warty elevations with stel- late markings at their apex and an irregularly outlined base; gleba citron; veins obscure; asci elongate or ovoid and narrowed at terminus, briefly pedi- cellate, 8-spored; spores irregularly seriate, large, globose, verrucose, 14 # in diam. Type, No. 253, Harkness Coll. Bot.—Vot. I.] HARKNESS—CALIFORNIAN HYPOGALI. 269 Beneath Sequoias, Mill Valley, Marin County, Calif., July. The spores are often seriate, at times, however, nearly biseriate. The fungus resembles P. conglomeratus B. & Br., but has a much larger spore. 75. Pachyphleus ligericus 7w/. Pachyphieus ligericus Tu., Fungi Hypo., p. 133, Tab. XIV, Fig. 5. Exiguus verrucosus et nigricans; sporangiis ovato-globosis; sporarum ver- ruculis crassis obtusisque. No. 44, Harkness Coll. Under pine trees in sandy soil, Towles, Placer County, Calif., May. Myrmecocystis, gen. nov. Fungus minute, irregular, lobed or gibbous, verrucose; gleba chambered by an irregularly stellate cavity not communicating with the exterior; asci subglobose or somewhat elongate, 8-spored; spores rough. 76. Myrmecocystis cerebriformis, gen. et sp. nov. PLATE XLV, Fics. 28a-28e. Minute, 1 cm. in diam., color white or pale citron, lobed, verrucose, enclosed; gleba white, marked by an irregularly stellate-formed cavity with- out regular lines, the structure composed of large and uniform cells; asci subglobose or slightly elongated, 8-spored; spores globose, 24 4 in diam. Type, No. 25, Harkness Coll. In sandy places under oaks, Wire Bridge, Placer County, Calif., May. 77. Myrmecocystis candida, sp. nov. PLATE XLV, Fics. 29a-29¢. Minute, 0.5 cm. in diam., color white, irregular, lobed, verrucose; gleba irregularly chambered; asci subglobose, 8-spored; spores globose, rough. Type, No. 18, Harkness Coll. 270 CALIFORNIA ACADEMY OF SCIENCES. {[Proc. 3D SER. In rich sandy soil under oaks, Alameda County, Calif., June. Differing from J/. cerebriformis in magnitude and in the spore. Geopora Lk. Geopora Hx., Bull. Cal. Acad. Sci., Vol. I, No. 3, 1885, p. 168. Subterranean. Integument woolly, continuous with the trama. Hymenium convolute. Asci cylindrical. Sporidia hyaline, oblong, smooth. 78. Geopora Cooperi /Zz. Geopora Cooperi Hx., Bull. Cal. Acad. Sci., Vol. I, No. 3, 1885, p. 168. Irregularly globular, 2-4 cm. in diam., covered with dense brown wool which is continued inwards on the trama; absorbing base none; hymenium white, not closely packed; asci cylindrical, 8-spored, 220 x 26 4; sporida hya- line, oblong, smooth, with a large shining, eccentric nucleus, 28 x 20 p. Type, No. 106 (3880), Harkness Coll. Haywards, Alameda County, Calif., January. Coll. by Dr. J. G. Cooper: 79. Geopora magnata, sp. nov. PLATE XLV, Fics. 34a-34d. Large, 6 cm. in diam., semiglobose, color brown, with brain-like convolu- tions marking its entire surface; septate hairs minute and in great abundance; gleba white and marked by labyrinthine cavities; asci cylindrical, 8-spored; spores white, destitute of oil globules, roundly ellipsoidal, 14 x 18 #%, much shorter than those of G. Coopert. Type, No. 255, Harkness Coll. Amongst Prunus insignis, Golden Gate Park, San Fran- cisco, Calif., January. 80. Geopora brunneola, sp. nov. Irregularly globose, 3 cm. in diam., color brown, corrugated, tomentose, enclosed; gleba white; hymenium fleshy, densely crowded; asci cylindrical, 80 x 12 #, 8-spored; spores obovate, hyaline, 12x 184. The asci are much shorter than in G. Coopferi and are densely crowded. Type, No. 102, Harkness Coll. Bot.—VOL. I.] HARKNESS—CALIFORNIAN HYPOGA@I. eae ia In sandy ground, Golden Gate Park, San Francisco, Calif., April. 81. Geopora mesenterica, sp. nov. PLATE XLIII, Fics. 12a-12¢. Smooth, irregular, 5 cm. in diam., color dirty white; gleba ferruginous brown; parenchyma convolute; asci cylindrical, 102 x 12 4; with elongated pedicel, 8-spored; spores white, ovoid, smooth, 10 x 12 yp. Type, No. 37, Harkness Coll. Under Ceanothus in decaying vegetable humus, Auburn and Wire Bridge, Placer County, Calif., May. Tuber Wichelz. Tuber MIcHELI, Nov. Pl. Gen., p. 221. Asci short, saccate, disposed in sinuous veins. Sporidia elliptic, reticulate, often echinulate. Peridium warty or tubercled, rarely smooth, without any definite base. (Berk. Outlines Brit. Fung., p. 376.) 82. Tuber (Eutuber) citrinum, sp. nov. PLATE XLV, FIGs. 30@-30¢. Subglobose, 2 cm. in diam., irregular, warty, fissured, color citron, turning to pale brown; gleba white; asci globose, enclosed in wavy filaments; spores two to three, seldom four; spores ellipsoidal, reticulate-alveolate, dark brown, 30 x 42 »; about eighteen alveoli on the circumference of the spore. Type, No. 123, Harkness Coll. In forest, Tamalpais, Marin County, Calif., May. Resembling 7. rapeodorum Tul. excepting in form and the structure of the gleba. 83. Tuber (Eutuber) monticolum, sp. nov. Minute, globose, 1.5 cm. in diam., smooth, color dirty white, fissured, with a slightly defined sterile base; gleba white, firm; asci globose, briefly stipi- tate, 2-4-spored; spores ellipsoidal, dark, reticulate-alveolate, 36 x 25 yw. Type, No. 27, Harkness Coll. Among firs in dense woods in Sierra Nevada mountains at Towle, Placer County, Calif., July. 272 CALIFORNIA ACADEMY OF SCIENCES, [PRoc. 3D SER. 84. Tuber (Eutuber) magnatum Pico. Tuber (Eutuber) magnatum Pico, Melethemata, p. 79. Ochraceo-pallens v. dilute virescens, subleve aut minutissime papillosum, difforme, globoso-angulosum et varie lobatum, basi obconica instructum; venis aeriferis tenuissimis, reticulatis; asci 1-3-sporis; sporis fuscis elliptico- rotundatis et alveolato-reticulatis, retis alveolis amplis. (Tul., Fungi Hypo., p. 150.) No. 62, Harkness Coll. In oak forest, San Rafael, Marin County, Calif., March; under oaks, Wire Bridge, Placer County, Calif., March. 85. Tuber (Eutuber) Borchii V7tz. Tuber (Eutuber) Borchit Vitt., Monog. Tub., p. 44, Tab. I, Fig. 3. Globosum, vulgo regulare, puberulum, albidum, maculis candidis notatum posteaque rufescentibus conspurcatum, intus ex albido fuligineo-violaceum et etiam fusco-nigricans, venis albidis rariusque lineis obscurioribus marmor- atum; sporangiis elliptico-rotundatis, szepius 1I-3-sporis; sporis crassis ovatis reticulato-alveolatis spisseque luteo-brunneis. (Tul., Fungi Hypo., p. 145.) No. 54, Harkness Coll. Amongst decaying leaves of oak, Mt. Tamalpais, Marin County, Calif., June. 86. Tuber (Eutuber) australe Spee. Tuber (Eutuber) australe Spec., Ann. de Sociedad Cientifica Argentina, Vol. XXIV, 1887, p. 122. Globosum v. globoso-trigonum, superne integrum, inferne szepius trilobatum, magnitudine valde ludens (4-30 mill. diam.), lavissimum, sordide album; cutis tenuis a carne inseparabilis; caro compactiuscula, alba, dein grisea, venis parcis crassiusculis, albidis, immutabilibus ramoso-anastomosantibus per- cursa; asci in pulpa dense dispersi, globosi v. globoso-elliptici, primo 2-3 spori, dein szepius 1-spermi v. 2-spermi altero abortivo (70-90 yu x 60-80 #2): spore globoso-elliptice, pallide fulgineo-olivacez, areola hyalina, lata, reti- culato-alveolata cinctz (35-40 4 x 28-30” sine areola; 45-50 4 x 35-38 » cum areola). No. 203, Harkness Coll. Amongst oaks in vegetable humus upon a well drained hillside, Auburn, Placer County, Calif., June. Bot.—VoL. I.] HARKNESS—CALIFORNIAN HY POG AI. 273 87. Tuber (Eutuber) gibbosum, sp. nov. Irregularly gibbose, 2 cm. in diam., color cinnamon-brown; gleba ochrace- ous; septa white, tortuous, obscure; asci subrotund, 3-4-spored; spores dark brown, ellipsoidal, echinate, markedly reticulate-alveolate, large, 24 x 36 ps. Type, No. 162, Harkness Coll. Under oaks, Mill Valley, Marin County, Calif., April. 88. Tuber (Eutuber) excavatum )77. Tuber (Eutuber) excavatum Virr., Monog. Tub., p. 49, Tab. I, Fig. 7. Subglobose, about an inch in diameter; peridium discrete, ochraceous, minutely verrucose, firm; flesh horny, cinerous-red, liver-colored, or tawny; veins pallid-ochraceous; the substance falls away in the center, so as to leave a cavity, which has an opening at the base of the tuber; asci numerous, ellip- soid, 2-4-spored; sporidia ellipsoid, yellowish, or pallid-tawny; epispore largely foveolo-plicate. (Cooke’s Handbook, Vol. II, p. 740.) No. 159, Harkness Coll. Beneath oaks in clayey soil, Laundry Farm, Alameda County, Calif., April. The spore is somewhat larger (42 in diam.) than in the dried specimen, and like it coarsely reticulate. 89. Tuber (Spherotuber) puberulum 2. & Pr. Tuber (Spherotuber ) puberulum B. & Br., Ann. & Mag. Nat. Hist., rst Ser.. Vol. XVIII, p. 81. Gregarious, irregularly sublobate, clothed with short, erect down, which gives it to the naked eye a peculiar pearly appearance; the white spots are very visible, even in dried specimens; peridium very thin and delicate, so that the pinky-brown color of the flesh is apparent through it, often cracked; veins white from.a radiating base, in some individuals very few; sporidia nearly spherical, reticulato-echinulate; odor of the radish. (Cooke’s Hand- book, Vol. IT, p. 741.) No. 36, Harkness Coll. Growing amongst decaying pine bark in the forest, Don- ner, summit of the Sierra Nevada mountains, 7,000 feet, July; under Zzbocedrus, Towles, Placer County, Calif. - (3) July 8, 1899. 274 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. go. Tuber (Spherotuber) Californicum, sp. nov. PLATE XLV, Fics. 31a-310. Subglobose, 1.5 cm. in diam., ochraceous, smooth; gleba firm, brown; veins conspicuous, not plentiful; asci subglobose, not stipitate, 3-4-spored; spores globose, large, brown when mature, reticulate-alveolate, 42 » in diam., about ten alveoli on the circumference. Type, No. 150, Harkness Coll. Under oaks beneath vegetable humus upon a hillside, Laundry Farm, Alameda County, Calif., March. This species is especially notable for the magnitude of its spore. g1. Tuber (Oogaster) Caroli Bonnet. Tuber (Oogaster) Caroli BONNET, Rev. Mycol., Ann. VII, 1885, p. 8. Globosum, brunneo-ferrugineum, verrucis plerumque 5-goniis asperatum, basi squamiformi, eximia instructum; gleba firma, sicca, pallide luteola, dein luteola, venis albis, numerosis, latissimis, e fungi basi exorientibus, gyrosis marmorata, lineis obscuris destituta; ascis globosis v. piriformibus, longe late- que stipitatis, 1-4-sporis; sporidiis ellipsoideis, dense et acute aculeatis, mag- nis, 20-22=14-15, luteo-brunneis. (Paoletti in Saccardo’s Sylloge Fung., Vol. VIII, 1889, p. 894.) No. 149, Harkness Coll. In clayey soil beneath oaks, Laundry Farm, Alameda County, March; Howards, Marin County, Calif., May. g2. Tuber (Spherogaster) candidum, sp. nov. PLATE XLV, Fics. 32a-320. Subrotund, 2 cm. in diam., smooth, color whity brown; gleba light brown; veins attenuate, white; asci subglobose; 3-4-spored; spores globose or ovoid, echinate, brown when mature, 24 » in diam. Type, No. 195, Harkness Coll. Under dense clusters of Ceanothus, Auburn, Placer County, Calif., May. Differing from 7. echinatum Sacc. in the form of the spore. Bot.—VOL. I.] HARKENESS—CALIFORNIAN HYVPOGA‘]. 275 93. Tuber (Spherogaster) Eisenii, sp. nov. Irregularly oblong, 3 cm. in diam., common integument smooth; gleba pale or whitish; veins large; asci ovate, stipitate, 1-2-spored, seldom more than one; spores globose, dark brown, echinate, 18 ” in diam. Type, No. 196, Harkness Coll. In sandy places beneath vegetable humus, Auburn, Placer County, Calif., May. Named in honor of Dr. Gustav Eisen of the California Academy of Sciences. 94. Tuber (Spherogaster) olivaceum, sp. nov. Semiglobose, 2 cm. in diam., color ferruginous brown, smooth; gleba oli- vaceous; veins minute; asci ellipsoidal, markedly pedicellate, 2-4-spored; spores globose, echinate, dark brown, 24 u in diam. Type, No. 197, Harkness Coll. Beneath vegetable humus, Auburn, Placer County, Calif., May. Piersonia, gen. nov. Integument scabrous or warty; gleba showing a multiplicity of brownish dots, orbicular or gyrose; asci nesting together; spores 3-4, alveolate. Named in honor of William M. Pierson, a member of the California Academy of Sciences. 95. Piersonia alveolata, gen. et sp. nov. PLATE XLIV, Fics. 20a-20e. Diameter 1 cm., integument scabrous, color white, turning to sulphur; gleba firm, citrine, cut surface showing a large number of orange-colored dots; asci clavate, 60 x 80 4, pedicel elongated (70 ~), 3-4-spored; spores alve- olate, citrine, 24 # in diam. Type, No. 183, Harkness Coll. Beneath Ceanothus, Auburn, Placer County, Calif., May. 96. Piersonia scabrosa, sp. nov. PLATE XLIV, Fics. 21a-2I1¢e. Semiglobose, irregular, 2 cm. in diam., color chestnut-brown, surface rough; gleba buff; asci obtusely saccate, pedicellate, 4-spored; spores globose, white, alveolate, 20 » in diam. . Type, No. 201, Harkness Coll. 276 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. In the forest, Auburn, Placer County, Calif., June. Differing from the preceding in color and in dimensions of the spore. Delastria 7z/. Delastria Tut., Ann. Sci. Nat., Bot., 2d Ser., Tome XIX, p. 379. Ascomata basi obtusa protuberante instructa, cortice tenuissimo, fibrilloso- byssoideo, passim rimoso aut evanido vestita. Gleba carnosa, humida, mollis, venis candidis anastomosantibus variegata et iisdem in glebulas rotundatas, quasi septis spuriis, divisa. Asci oblongo-reniformes, ampli, 2-4-spori. Sporidia spheerica, reticulato-alveolata, nucleo oleoso donata. (Sacc. Sylloge Fung., Vol. VIII, 1889, p. 904.) g7. Delastria rosea 7u/. PLATE XLV, Fics. 27a-276. Delastria rosea Tut., Ann. Sci. Nat., Bot., 2d Ser., Tome XIX, p. 379. Globosa v. obovata, depressa, vulgo gibberoso-mamillosa, interdumque sulcata et rimosa, fragilis, cortice adpresse byssaceo-tomentoso, subsericeo v. velutino, niveo dein fuscato involuta; gleba ex albido mox amcene rosea, dein rufa, venis candidis, immutabilibus, cum cortice continuis, areolata; ascis inordinatim sparsis, ovoideo-oblongis, szepius incurvis seu reniformibus, nec in modum pedicelli deorsum angustatis, 2-3-rarius 4-sporis; sporidiis, sphzeri- cis, reticulato-alveolatis, subaculeatis, maturis luteolis, 30-40 »# in diam. (Pao- letti in Sacc. Sylloge Fung., Vol. VIII, 1889, p. 905.) No. 182, Harkness Coll. Under shrubs amongst vegetable humus, Auburn, Placer County, Calif., May. . This is the only species found of this genus. The characteristic rose-pink tint shown upon the cut surface of the gleba is still to be seen even after months of immer- sion in alcohol. About fourteen alveoli appear on the circumference. Choiromyces J’7/t. Choiromyces Virt., Monog. Tub., p. 50. Common integument, even. Base definite. Asci clavate. Sporidia spher- ical. (Berk. Outlines Brit. Fung., p. 377-) BoT.— VOL. I.] HARKNESS—CALIFORNIAN HVPOGAI. 277 98. Choiromyces gangliformis )77¢. Choiromyces gangliformis Virt., Monog. Tub., p. 51, Tab. II, Fig. 2. Globosus, levis, fuscus; gleba albida, exiccatione vix mutata, grumoso- ° compacta, venis numerosissimis, interruptis, decolorantibus, mirabiliter anas- tomosantibus et areolas hinc illinc albas filisque minimis concoloribusque varie junctas, gangliformes, inter se relinquentibus; sporidiis sphzericis, verru- cis conicis elongatis asperatis. (Paoletti in Sacc. Sylloge Fung., Vol. VIII, 1889, p. gor.) No. 151, Harkness Coll. Under Arctostaphylos, Calistoga, Napa County, Calif., April. Terfezia Zw/. Lerfezia TuL., Fungi Hypo., p. 172, Tab. VI, Fig. 4; Tab. VII, Fig. 5; Tab. XV, Figs. 3-5; et Tab. XXI, Fig. 15. Integumentum crassum v. tenue, carnosum, clausum continuum v. hinc et illinc parce rimosum, leve, nonnunquam fibrillosum, in cuticulze sorte qua tegitur primo albidum deinque plus minus et inzequaliter fucatum. Moles interior carnosa solida, scil. lacunis destituta, initio pallida et quasi similaris, matura uvida, molliuscula, in massulas s. glebulas rotundatas, rarius diversi- formes, carnoso-humidas v. pulposas, fertiles et varie coloratas divisa, paren- chymate sterili interposito laxiori zerifero pallido maculas que irregulares (areolas) s. venarum species in fungo secto fingente. Sporangia late elliptica v. globosa, utriculis genitivis imposita et in glebulis veluti inordinate creber- rima nidulantia, octospora. Sporze spheric initio conglobate et leves, mature libercze echinate (aliquando insuper reticulatze) diluteque colorate; nucleo tandem oleoso. 99. Terfezia spinosa, sp. nov. PLATE XLV, Fics. 24a-240. Terfezia leonis LANGLotrs in Ellis’ Centuries, No. 1782. Globose, white or citron, smooth; gleba variegated; asci subrotund or briefly elongate, 6-8-spored; spores globose, seldom ellipsoidal, 15-20 2 in diam. Type, No. 1o8a, Harkness Coll. Red River Valley, Louisiana. The spores are furnished with large projections (only about twenty being found upon the periphera of each), which are slightly curved, somewhat blunt at the point, and confluent at the base. As will be observed, there is a marked dissimilarity be- tween this species and 7. /eonzs, a specimen of which 278 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. from the herbarium of Vittadini is in possession of the writer. A spore from this specimen is represented in fig. 25, and agrees in outline with that figured by Tul. in Fungi Hypo., Plate XV, fig, 3. 100. Terfezia Zeynebie, sp. nov. PLATE XLV, Fie. 26. Large, globose, 4 cm. in diam., color white, turning brown; gleba citrine when dried; asci subspherical, 8-spored, 36 x 40 » (briefly pedicellate); spores with spine-like projections, 15 to 18 # in diam. Type, No. 212, Harkness Coll. The valley of the Tigris, Arabia. This fungus was sent to me some years ago by the American Consul at Bagdad. Named in honor of Zeyneb of the Arabian Nights. T. Zeynebie is found in clusters in the alluvial soil of the valley and is readily discovered owing to the fact that slight fissures are to be seen in the overlying earth. The fungus is greatly esteemed as an article of food and is sold in large quantities in the markets of Bagdad. The spine-like projections are short and blunt, generally some little distance apart, about 16 projections appearing on the circumference. Chatin in La Truffe, page 78, describes two species from Mesopotamia, 7. Hafizi and 7. Metaxast. ‘The first, 7. Hafizt, is figured by the author (Plate XV, fig. 1) as being rugose and destitute of spines. Paoletti in Saccardo’s Sylloge Fungorum, Vol. XI, p. 445, refers to the same as possessing a reticulate spore, 18x 20 pw. T. Metaxas’, shown in plate XIII, fig. 2, Ibid., has a much larger spore with a very large increase in the number of its spines. Terfeziopsis, gen. nov. Ascomata smooth, globose or pyriform; gleba veinless, firm; asci globose or ellipsoidal, 2-4-spored; spores globose or ovoid, echinate; spines recurved or hooked. This fungus is nearly allied to Zerfezza, but is separated from Terfezia because of the form of its spore. Bot.—VOL. I.] HARKNESS—CALIIFORNIAN HVPOG, 2 hacen 281 eyanen’ 0.) 1)... i sie OCTAVIANIA . =. +--+ +++ + + +. 251 Stephensias. yo. cos sb sk eee 268 ACER sem eth ayucien bs Fe ean Aes De 260 | bonibycina’s: 5).<) ulalas Soe eee hHranneclas .o. va Ale s.00' A po ee CITSIMA: ca) so we San th per ee COMIPACtA s.\.s a ¢ eee > 253 TERFEZIA Sted er ae eee : Pay y7/3/ MIGHTCOlA® 4 easel 254 Hafizi .......-+.-.-. “ees PATER cats. cole! ed oak eae oe 252 leonis Ai aA RC? 277 occidentalis .......... 253 Metaxasi.... .- ...- +) 218 POSEA! ici dss sete Le ier ek ee 252 Spittosa «<5. 3/2 essa) eM 277 SUP COPSELDS adh ay esa to aT els 260 Zeynebize . ... +.) + ois seers BocIgUA. chistes ene 252 Terfevionsis:.)| |. .)2,.76.. 3 - < 32 298 Stephene.) fs avon see eee 253 lignaria ......-.-.-..-- 279 WETS oe na als da Shed ARS ores Pe 7 | (Eutuber) australe. ....... 272 PACH VPHUGUS,;) a).ls fle cee a ae ahe 268 Borchii....-...-. 272 Gartens: J 6. sh fiche ane eee et ES citrinum....... 271 Lig ericHey) WN, fue tes aca seein 269 excavatum . ... . 273 Bicehomtans oy) ec bie oe Ke 275 gibbosum ...... 273 Mivectatay Otte ti te cote ese 275 magnatum ...... 272 ScabtosaA ss J. c ais eee ees monticolum.... . 271 Pseudohydnotrya.......... 267 (Oogaster) Caroli .....-.-- 274 earned. (. 17) oe ul eiarn et er oD (Sphzerogaster) candidum . ee /-' Harknessi 2 fice Sed co (abe Gy Hisenii .... 275 MISTER Rt Pee) ee 267 olivaceum .. 275 (Sphzrotuber) Californicum . 274 puberulum ... 273 RAIZOPOGOM ane Leecniat oe ete 257 chet ge bel > t- Muu ear eAley P eget eel 257 Ustilago cyanea. 2+. +--+ ee es 281 Bot.—Vot. I.] HARKNESS—CALIFORNIAN HYPOG4&-1, 285 BIBLIOGRAPHY. 1844. BERKELEY, M. J. Notices of British Fungi. Annu. G Mag. Nat. Hiist., 1st Ser., Vol. XIII. London. 1860. Outlines of British Fungology. London. 1848. and C. E. Broome. Notices of British Fungi. Aun. & Mag. Nat. Hist., 2d Ser., Vol. II, p. 267. London. 1846. Notices of British Hypogzeous Fungi. Ann. G Mag. Nat. Fiist., 1st Ser., Vol. XVIII. London. 1885. BoNNET, H. Revue Mycologigue, Ann. VII, p. 8. Toulouse. 1892. CHATIN, Ap. La Truffe. Paris. 1871. Cooker, M. C. Handbook of British Fungi. London. 1879-80. Grevillea, Vol. VIII, p. 94. London. 1831. Corba, A. C. J. In Sturm’s Deutschlands Flora, Abth. III, Heft 11, p. 1. Nuernberg. 1842. Icones Fungorum hucusqgue cognitorum, Tome V, pp. 30, 74. Prage. 1888. Der Toni, J. B. Hymenogastraceai in Saccarbo’s Sylloge Fungorum, Vol. VII, p. 174. Patavii. 1896. FISCHER, Ep. Tuberinez in ENGLER and PRANTL’s Die naturlichen Pfhlanzenfamilien, Teil I, Abth. I, p. 282. Leipzig. 1821-29. FRIES, ELt1As. Systema Mycologicum, Vol. II, pp. 295, 297; Vol. 1884. 1885. 1809. 1729. 1895. 1788. 1885. 1887. 1853. 1845. 1843. 1831. III, p. 21. Gryphiswaldiz. Harkness, H. W. New Species of Californian Fungi. Bw//. Cal. Acad. Sci., Vol. I, No. 1, pp. 29, 30. San Francisco. Fungi of the Pacific Coast. Aull. Cal. Acad. Sci., Vol. I, No. 3, p. 168. San Francisco. Link, H. F. Ods. 1m Ord. Fl. nat., diss. I, in Mag. der. (berlin) Gesellsch. naturforsch. Freunde, Tome III, p. 33 (fide Tulasne, Fungi Hypogei.) MIcHELI, P. A. Nova Plantarum Genera, p. 221. Florentiz (fide Tulasne, Fungi Hypogei). PAOLETTI, J. Tuberaceze in Saccarpo’s Sylloge Fungorum, Vol. VIII. Patavii. Pico. Meletemata inauguralia de fungorum generatione. Turin. (fide Chatin, La Truffe). ROUMEGUERE, C. Fungi Gallici exsiccati. Revue Mycologigue, Ann. VII, p. 23. Toulouse. SPEGAZZINI, CARLOS. Fungi Argentini, Pug. IV, n. 237 (fide Sacc. Syll. Fung.) Las Trufas Argentinas. Ann. Soc. Cien. Argentina, Vol. XXIV, p. 122. Buenos Aires. Tuasnge, L. R., AND CHas. Fungi Hypogei. Paris. Nuovo Giornale Bot. Jtal., Ann. I, Vol. II, pt. 1, p. 63. Firenze. (fide Tulasne, Fungi Hypogzi.) Champignons Hypogés de la famille des Lycoperdacees, ob- servés dans les environs de Paris et les départemens de la Vienne et d’Indre-et-Loire. Ann. des Sci. Nat., Botanique, 2d Ser., Tome XIX, p. 379. Paris. VITTADINI, CAROLO. Monographia Tuberacearum. Mediolani. WaLLroTH. In Dretricn’s Flora des Kenigr. Preuss., Vil, p. 465 (fide Tulasne, Fungi Hypogzi.) 286 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. The colored illustrations have been mostly prepared after fresh material, a few after alcoholic preparations. The microscopic drawings have been made from glycerine preparations with the aid of a Zeiss Apochromat, Obj. 2 mm., Aperture 1.40, Compensating Oculars 8 and 12. Projection on working table. EXPLANATION OF PLATE XLII. Fig. 1. Aysterangium Phillipsti, sp. nov. (a) Vertical section with fibrillose rootlets. (b) Section of gleba. Fig. 2. A ysterangium cinereum, sp. nov. : (a) Fully developed fungus. (b) Transverse section. Fig. 3. Octaviania monticola, sp. nov. (a) External view. (b) Transverse section of same. (c) Sec- tion of gleba enlarged. Fig. 4. Octaviania occidentalis, sp. nov. (a) Section of gleba enlarged; Zeiss AA, Oc. 2. (b) Section showing hymenium and pseudobasidia; Zeiss AA, Oc. 6. (c) Pseudobasidia enlarged, showing spores round or ovoid. (d) Free spore. Fig. 5. Octaviania socialis, sp. nov. (a) Perfect plant. (b) Vertical section. (c) Enlarged spores in situ. (d) Basidia, sterigma, and spores. Fig. 6. AMymenogaster utriculatus, sp. nov. (a) Entire plant. (b) Section of gleba showing cells and hyme- nium. (c) Enlarged section of hymenium. (d, e) Two spores with utricle. (f) Isolated spore. Fig. 7. Leucophleps magnata, gen. et sp. nov. (a) Section of hymenium with basidia and spores. (b) Vertical section showing cell-structure. (c) Optical section of spore with spore investment. Proc. Can Acan.5c1.3” Ser. Bor.Voul ei eal GUSTAV EISEN, DEL LITH BRITTON & REY, SE OLA, SP Nov Fie.4. OCTAVIANIA OCCIDENTALIS, 5P NOV. LEUROPHLEPS MAGNATA, GENET SP NOV. P16. HYSTERANGIUM PHILLIPS, SP NOV. FiG.2.HYSTERANGIUM CINEREUM, SP NOV. FiG.3. OCTAVIANIA MONT FiG.5, OCTAVIANIA SOCIALIS, SR Nov. ['iG.6. HyMENOGASTER UTRICULATUS. SPNov. fit 288 Fig. 8. Fig. 11. Fig. 12. Fig. 13. Fig. 14. CALIFORNIA ACADEMY OF SCIENCES. ' [PRoc. 3D SER. EXPLANATION OF PLATE XLIII. Leucophleps citrina, sp. nov. (a) Section of hymenium with basidia and spores. (b) Basidia isolated, 10 to 12 times the length of the spore (bb); (bbb) oil- globules shown within the basidia. Leucophleps odorata, sp. nov. (a) Mature plant. (b) Vertical section showing cell-structure. Genea compacta, sp. nov. (a) Section of gleba with asci and paraphyses. (b) Ascus with spores. (c) Isolated spore. Hydnocystis compacta, sp. nov. (a) Ascus with spores. (b) Isolated spore. (c) Optical sec- tion of spore. Geopora mesenterica, sp. nov. (a) Ascus with spores. (b,c) Isolated spores. Balsamia filamentosa, sp. nov. (a) Fungus fully developed. (b) Longitudinal section. (c, d) Asci with spores. (e,f) Isolated spores with oil-globules. Balsamia, sp. ? Immature fungus showing development of spores. Sporophaga cyanea, gen. et sp. nov. (a) Optical section of spore of Balsamia vulgaris in the early stage of its development. (b) Section of the same showing the membranous investments of the spore. (c) Aalsamia spore ina more advanced stage of development. (d) Ascus with spores of the host-plant; some of the spores already ruptured and the parasite set free within the ascus. (e) Group of parasites within a spore of the host. (f) Two parasites with hyphe within a spore of the host. (g) Isolated parasite showing a double investment of the same, together with its septate hyphz, the hyphze having a branched terminus. Fig. 16. Pseudohydnotrya carnea, sp. nov. Fig. 17. (a) Entire fungus, hairs faintly shown. (b) Transverse section. Spheria Setchellit, sp. nov. (a) Fragment of host-plant showing an ascus with spores and paraphyses; cell-wall of the perithecia enclosing parasitic spores. (b) Perithecia in different stages of development, one still destitute of spores; the asci of the host-plant faintly shown. (c) Isolated spore greatly magnified. Spores colored black for convenience, in life as in 17c. $o / Va N | ! f \ // \ oA { \ ff G \ | - = | h | ‘| \ y \ | ) \% J \52 wt G | ) f 13¢ / | N u | ga DA me \ \ : =: \ sv one M i \ s 13¢ > = i . ‘sf ee ah 7 : ) Fuad , a f) eH as [ | | rp AN _ 7 } 4 api { (iN » y L~ 5 4 bs j /. 1X x ee) : A r pF J 7 104 =. & a 13 as D it tae = = sj : bn: a : oh ; - ? 7 iy \ ) \ iS } \ ay 2 To\ vate | : Q ; | HAN Y 2 \Vt lie G (> BG j {aloe { \ ve ; } ‘x 10% \ , peice FF I : ' <3 10° a Ra GS se ne s Gus PISEN.DE TERT rieoR - iy STP THe Fi F MPACTA. SP. NO 6 COPHLEPS CITRINA, SP Fie. §. LEUCOPHLEE ! lav. | : co} Pic. 12. GEOPORA MESENTERICA, SP } Fig. 13. RALSAMIA FILAM st Re ash? | 3 Is lu { Fys. 16. PSEUDORYDNOTRYA CARNEA, Vv. Fic. 17 SPHARIA GETCHELLI, S= NOV ci ’ iV tt 5 ; +, ‘ee oy 290 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XLIV. Fig. 18. Stephensia bombycina TUL. (a) Fragment of gleba with asci. (b) Isolated spore showing its rugosities. (c) Optical section showing the periphery. Fig. 19. Aydnotrya cerebriformis, sp. nov. (a) Vertical section of fungus. (b) Enlarged vein-tracing. (c) Asci with paraphyses. (d) Isolated spore showing roughened surface. (e) Optical section of spore. (f) Section of the sur- face of spore magnified. Fig. 20. Piersonia alveolata, gen. et sp. nov. (a) Section of gleba showing veins, cell, and asci. (b) Asci (magnified) with spores. (c) Isolated spore. (d) Optical sec- tion of spore. (e) Section of spore showing alveolate surface. Fig. 21. Ptersonia scabrosa, sp. nov. (a) Section of plant. (b) Enlarged section of cut surface. (c) Ascus with spores. (d) Optical section of spore. (e) Sec- tion of spore surface. Fig. 22. Eudogone malleola, sp. nov. (a) Transverse section of fungus, full size of the mature plant. (b) Conceptacle with contents and fragment of pedicel. Fig. 23. Terfeziopsis lignaria, gen. et sp. nov. (a) Ascus with spores (spores round or slightly ovoid). (b) Optical section of spore (about 70 to 75 spines in its circumfer- ence). (c) Hooked spines on surface of spore. ; 25 ' f woe ) ‘ @ 7 te = : 2.0%, << : « « i] ie 7 P =~ +, Suk ie _ ed 7 1 ® - —_ Sao y" 5 i — SU Song g es ee - =- Proc.CatAcan.Scr.3? Ser Bor.Vol, (Harnness] Flare XLIV Gustav Eisen, Deu. LETH. BRITTON & REY. SP Pi6.1B. STEPHENSIA BOMBYUINA 7iz. Fic, 19. HYONOTRYA CEREBRIFORMIS, SP NOV. Fis. 20. PIERSOMA ALVEOLATA. GEN. ET SP NOV, Fis. 21. PIERSDNIA SCABROSA, SP NOV. Fic. 22. ENDOGENE MALLEOLA, SP NOV. FIG. 23. TERPRZIOPSIS LIGNARIA,GEN.ET SP NOV. mS We as | 292 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. - EXPLANATION OF PLATE XLV. Fig. 24. TZerfezia spinosa, sp. nov. (a) Isolated spore. (b) Spine on surface of spore. Fig. 25. Zerfezta leonis Tul. Isolated spore. Fig. 26. Terfezia Zeynebia, sp. nov. Isolated spore. Fig. 27. Delastria rosea Tul. (a) Section of gleba with ascus and spores. (b) Isolated spore. Fig. 28. Myrmecocystis cerebriformis, gen. et sp. nov. (a) Fully developed fungus. (b) Vertical section. (c) Detail of surface of fungus. (d) Ascus with spores. (e) Detail of surface of spore. Fig. 29. Myrmecocystis candida, sp. nov. (a) Section of gleba. (b) Asci with spores. (c) Isolated spore showing a rough surface. (d) Optical section of spore. (e) Detail of sculptured surface of spore. Asci in c and d im- properly outlined, should appear as in 6. Fig. 30. Zuber (Eutuber) citrinum, sp. nov. (a) Section of gleba with ascus and spores. (b, c) Isolated spores. Fig. 31. Zuber (Spherotuber) Californicum, sp. nov. (a) Section of gleba with asci and spores. (b) Isolated spore. Fig. 32. Zuber (Spherogaster, candidum, sp. nov. (a) Ascus with spores. (b) Isolated spore. Fig. 33. Pachyphleus carneus, sp. nov. (a) Ascus with irregularly serrate spores. (b) Isolated spore. Fig 34. Geopora magnata, sp. nov. (a) Fully developed fungus. (b) Section of fungus. (c) En- larged section of gleba. (d) Enlarged section of hymenium. Paor Cart, Aran Grr 32 Ger B if Proc. Cau Acan.Se1.32 Ser BorVoul [Haruness] Phare XIV. Gustav Eisen Dev. LTH BRITTON S REY, SF F Fic. 24 TERFEZIA SPINDSA.SP NOV Fic. 25 TERPEZIA LEONIS Zi. Fi. 26. TeRrezia ZEYNEBIE,SP NOV. Fic.27 DELASTRIA ROSEA 7iz. 1G.28.MYRMECOCYSTIS CERPBRIPORMIS, GEN. BT SP NOV. Fic. 25. MyRMECOCYSTIS CANOMA.SP NOV Fic. 30. TuBER cITRINUM, SP Nov. Fie3l. TuserR CALIFORNICUM, SP NOV. Fic. 32. TUBER CANDIDUM, SF NOV. FIG.33. PACHYPHLIBUS CARNEDS,SP Nov. Fic.34, GEOPORA MAGNATA, SP NOV. STUDIES ON THE FLOWER AND EMBRYO OF SPARGANIUM. BY DOUGLAS HOUGHTON CAMPBELL, Professor of Botany, Leland Stanford Juntor University. CONTENTS. PAGE. PLATES XLVI-XLVIII. Pepe ETE OWES tera casr tehepeeictshosstetee eras a4 fa nad co siale ia Yoronerel oto ap sas tare eres 295 DWE WSURULIMIE TIODEIO TN NE Ra ocd ice) ek Oa REARS 297 Re CrES ae POD CT Uk ea A are le eee es 298 By, GE Port RR OPAC. 20) sin Aiuto A Bind « ct ratelerd oye wins baa einem es aleaaee ae 299 BIE ¢) 4G CUE TA MELONS ovis 2125 bin a alm ca fll biollalas, 4 Wasa soll ea OD oe atnal teeny 302 IV. THE SECONDARY GROWTH OF THE ANTIPODAL CELLS........ 303 Nea ELE EUMBRY Ojrsc tore carats sits aise tle ns Sates ou nate aie grclareeinial tear aeaae 307 Wilk) CABNORMWALUEIMBRVO-SAGS saleniayeid se nie hie.ctee ae chile wads cabte teller 318 ge IRC AGLI AGRON 3 dh aia Sis ieee nl dip aS io.d's wis! owl er wfeeug sera alaeintnel eee 319 WEE. Tak Aroistatss: OF) SPARGANIUM ... 56.0: 2isc0g cc clet cle omelets 321 BIBEIOGRAPHIVE yaiirelstere atee ofsiey ciate wis tje aaa cnteel her talon e ra eet ate 322 EAPUANATION Gi, PEATES E 2/15 0A. si. tae te we taare em neem 324 THE genus Sfarganium Tourn. includes about a dozen species distributed over the northern hemisphere, but also represented in New Zealand and Australia. The genus is a peculiar one, and there has been much diversity of opin- ion as to its affinities with other Monocotyledons. It has usually been associated with 7yfha in the family Typhacee, but the present view (Engler and Prantl, 1889) is that it should be considered the type of a separate family, Spar- ganiacee, with the Pandanacee or Screw-pines as its nearest allies. The genus has not been extensively studied, but our knowledge of the development of the flower and embryo of the European S. ramosum is fairly complete (Hegel- maier, 1874; Dietz, 1887). The other species have, so far as I know, been treated from the standpoints of the descrip- tive botanist only, and no account of the embryo-sac has [ 293] July 15, 1899. 294 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. yet been published. Some years since I collected a small amount of material of the flowers of the common eastern species, &. eurycarpum Englm., and an examination of this material revealed several puzzling peculiarities of the embryo-sac, which were not clearly understood at the time. Figures of a few of these were published two years ago (Campbell, 1897), but lack of proper material, as well as other work, prevented any further study at the time. During the summer of 1896 a trip to Japan enabled me to procure a few specimens of the Japanese S. longifolium Turcz., and during the past season a good supply of S. szm- plex Huds. was collected in August at Tallac on Lake Tahoe. This species proved very satisfactory for study, and was made the principal subject of my investigations. In September, through the kindness of Miss Eastwood, a quantity of S. Greenzz Morong. collected at Lake Merced, near San Francisco, was sent me, and served to supplement the work done on SS. s¢mplex. My material of the latter species was much more complete than that of any of the others, and moreover it proved easier to handle. The results obtained from a study of this species have been compared, so far as was possible, with those from the other species mentioned. A preliminary statement in regard to S. semplex has already been published in a recent number of the ‘‘ Botanical Gazette’? (Campbell, 1899); but this refers simply to cer- tain of the most striking points concerning the embryo- sac, and no account of the embryo is included. Unfortunately all of the material collected was too old to show the earlier stages of development of the flower, so that the early history of the pollen and embryo-sac could not be traced. In the youngest flowers examined, the structures of the embryo-sac were already complete, and in most cases the embryo-sac was ready for fertilization. Hence in the present paper the study of the flower begins at the time when it is ready for pollination. In the preparations of the material, various reagents were employed in fixing it: one-half to one per cent. chromic acid, BotT.—VOL. I.] CAMPBELL—SPARGANIUM. 295 saturated alcoholic solution of picric acid, saturated alco- holic corrosive sublimate, and weak Flemming’s solution were employed, but of these the corrosive sublimate, on the whole, proved most satisfactory. The material was imbedded in paraffin and the sections were stained, usually with a double stain of Bismarck-brown and safranine, although in some cases they were stained zz foto with alum- cochineal before they were imbedded. kK tas, ProweEr: All of the species of Sfarganium are monececious, both staminate and pistillate flowers being aggregated in dense heads which are either sessile or borne on short pedicels. The lower heads are always pistillate and the upper ones staminate. The latter are usually much more numerous. In S. s¢emplex and its allies the main axis is unbranched, while in the other species, e. g., S. eurycarpum and 5S. Greeniz, the stem is branched. Both sorts of flowers are exceedingly simple in structure. The individual flowers are borne in the axils of small bracts, and are themselves surrounded by a varying number of small membranous leaves which are usually considered to represent a very simple perianth. The male flower is com- posed of three or more stamens, while the female flower has either a single carpel, as in S. s¢zmplex, or a compound pistil composed of two completely united carpels, as in S. ramosum or S. Greeniz. The development of the flower has been studied to some extent in S. ramosum (Dietz, 1887), and is briefly as fol- lows: Upon the primary axis are borne elevations which, in the lower part of the axis, where they are produced in axils of leaves, may develop into secondary axes, but in the upper portions of the axis, form at once the young inflorescences. In S. s¢mf/ex and its allies, this axis does not branch, and all of the lateral axes develop into heads of flowers. 296 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. In S. semplex the lower three or four of these heads develop the pistillate flowers, while the number of staminate heads is usually larger. The individual flowers, according to Dietz (I. c.), arise as small prominences upon the hemispherical fundament of the inflorescence, and upon these prominences arise sec- ondary outgrowths which develop respectively into stamens or carpels. Dietz does not describe in detail the develop- ment of the stamen, and unfortunately my own material was too old to show this, so that it is not now possible to make a comparison with such Monocotyledons as have been investigated. The scales of the perianth arise early, but Dietz does not describe their development. To judge from his somewhat brief account and very dia- grammatic figures, the development of the carpel and ovule is much like that of the low types like Zannzichellia and Lilea, which Sparganium resembles in these respects more than it does 7ypha, with which it has usually been associ- ated. While further investigations are required before this can be decided, it looks as if in Sfarganzum, also, as in other low Monocotyledons, that the solitary ovule is of axial origin and not a product of the carpel. The pistillate flower in S. semp/lex (fig. 1) consists of a single carpel surrounded by about six delicate membrana- ceous scales which form the perianth. Engler (Engler and Prantl, 1889) figures these in this species as having en- tire margins, but the form studied by me had the end of the narrowly spatulate scale sharply toothed (fig. 2), and the most recent figure (Britton and Brown, 1896) of this species shows the same thing, but with fewer teeth than my specimens. A delicate midrib traverses each scale. The ovary in this species merges gradually into the slender style which terminates in the narrowly sagittate stigma which is covered with minute papille. In S. dongifolium, which in many respects resembles 5S. simplex, the style and stigma are both shorter. In a second group represented by S. Greenit and S. eurycarpum the Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 297 pistil is composed of two completely united carpels, and the resulting fruit is two-seeded. In both of these the style is very short or quite absent and the two stigmas are very long (fig. 4). The base of the ovary is also less tapering than in S'. sémplex and the perianth scales are heavier and nearly or quite entire at the apex. S. simplex, which was the species principally studied by me, is a wide-spread form occurring throughout the north- ern hemisphere. In California it is confined to the higher mountains, the specimens collected by me being found near Lake Tahoe which has an elevation of over 6,000 feet above sea-level. The Staminate Flower. In S. s¢mplex the staminate flowers form dense heads, each flower consisting of about six stamens. Just before the dehiscence of the anthers, the filaments elongate very much, so that the head becomes much looser, and the pollen is readily dislodged by the swaying of the slender filaments, much as in the grasses and sedges. The anther has four loculi as in most Angiosperms and offers no specially note- worthy peculiarities. The other species seem to agree closely in the structure of the stamen and were not crit- ically examined. The pollen-spores of S. semplex are small globular cells (fig. 6) and at maturity show the usual division of the nu- cleus into a larger vegetative one, and a smaller generative nucleus, but the latter was not seen to divide again while the pollen remained within the anther. Probably the final division of the generative nucleus takes place within the pollen-tube. Perhaps owing to its small size it was not pos- sible to demonstrate the presence of a definite generative cell, the nucleus apparently lying free in the cytoplasm of the pollen-spore. As the cytoplasm of the ripe spore is very dense and stains strongly, it was not always easy, even in sections, to clearly distinguish the nuclei. The outer spore- membrane is marked with fine granulations but is not very thick. In this species there was almost always present in 298 CALIFORNIA ACADEMY OF SCIENCES. (Proc. 3D SER. the ripe spore a structure which closely resembles the small vegetative prothallial cell in the germinating microspores of such heterosporous Pteridophytes as /soe¢es and Se/aginella. Chamberlain (1895) speaks of such a prothallial cell as an occasional occurrence in Lz/iawm Philadelphicum, but except for this reference I am not aware of any reference to such a structure in the angiospermous pollen-spore. This prothal- lial cell, if such it is, in Sparganium simplex is lenticular in form (fig. 6), and separated from the body of the spore by a very well defined membrane which stains strongly with Bismarck-brown. While it was not possible to demonstrate positively the presence of a nucleus in this cell, there was usually to be seen a central body which stained more deeply than the rest of the cell with the usual nuclei stains and had much the aspect of a partially disorganized nucleus. While it seems very likely that this really represents a prothallial cell, homologous with the similar one in the microspores of Tsoetes, for instance, its real nature cannot be positively as- sumed until its history has been followed. The stamens of S. Greenz? closely resemble those of S. simplex, but the pollen-grains did not show any trace of the prothallial cell, and the epispore is marked with fine reticu- lations. At one point (fig. 7) these were absent, and the clear area probably marks the point at which the pollen- tube emerges. The Pistillate Flower. The pistil may be composed of a single carpel, as in S. simplex and S. longifolium, or of two completely united ones, as in S. Greenzd and S. eurycarpum. In S. simplex (fig. 3) a longitudinal section of the ovary shows the sin- gle anatropous ovule pendent from the upper part of the ovarian cavity. The narrow canal traversing the style opens into the ovary close to the base of the funiculus of the ovule. The funiculus is rather slender and has a single axial vascular bundle. The nucellus is oval and there are two integuments usually found in the ovules of Monocotyledons. Bot.—Vot. I.] CAMPBELL—SPARGANIUM. 299 The outer integument extends above the inner one and its upper margins are somewhat enlarged, so that the micropyle is quite closed. Il. Tue Emsryo-Sac. The youngest embryo-sacs met with had the egg-appa- ratus and antipodal cells developed, and nothing can be stated as to their early history. As the ovule at the time of fertilization is of the ordinary type, it is not probable that there are any marked deviations from the type in its early development. In S. simplex (figs. 3, 8) the mature embryo-sac is broadly oval in outline and is covered at the apex by about two layers of nucellar cells, the central ones somewhat elon- gated, so that the apex of the nucellus is slightly pointed. At the sides there are about four layers of cells in the nucellus, and these remain permanently and are not de- stroyed by the developing embryo-sac as so often happens. Within the embryo-sac the granular cytoplasm is principally confined to a thin parietal layer, except that surrounding the primary endosperm-nucleus there is a considerable amount of granular cytoplasm. In S. Greenii (fig. 5) the two carpels are completely united. A section of the ovary shows a long central pla- centa formed by the coherent inner faces of the carpels, and from the upper part of the placenta depend the two ovules. These correspond in structure with those of S. simplex but are somewhat longer and more slender, although they are less constant in form than in that species. The embryo-sac is longer and narrower and more pointed at the ends. The part of the nucellus above the apex of the embryo-sac is thicker, there being usually about four layers of cells at this point. Sparganium longifolium is much like S. simplex in the form of the pistil, except for the shorter style and small stigma. The ovule is of about the same size and shape, and the embryo-sac, so far as could be judged from ‘the 300 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. small amount of material available, closely resembles that of S'. simplex, and as in that species there are but two, or at most three, layers of cells at the apex of the nucellus. The common eastern SS. eurycarpum, while resembling more nearly S. Greenjz, differs from it in some details. The ovary is broader, due to an enlargement of the outer cells, and there are numerous enlarged cells present which contain bundles of rhaphides, or needle-shaped crystals, presumably of calcium oxalate. These spicular cells also occur in the placenta, and while not entirely absent from the ovary of S. Gveenzz are much less abundant. There are also found on the upper surface of the placenta papillate cells which are much less marked in S. Greenzz. These are doubtless connected with the conduction of the pollen- tubes. The structure of the ovule itself, and the embryo- sac are much alike in the two species. In all the species examined the cells of the nucellus per- sist, and the subsequent enlargement of the upper part of the integuments and the apex of the nucellus gives rise to the peculiar cap (‘‘Samen-deckel”’ of Hegelmaier) which characterizes the ripe seed of all species of Sparganium. The broadly oval embryo-sac of S. s¢mp/ex is scarcely at all narrowed at either end. In all specimens examined the two polar nuclei had already completely fused, and it is ev- ident that the fusion of them to form the primary endosperm- nucleus occurs some time before the fertilization of the egg-cell. The large endosperm-nucleus (fig. 10) is im- bedded in a considerable mass of granular cytoplasm which is elsewhere confined to the thin parietal layer. The posi- tion of the endosperm-nucleus is not constant, and it was found in nearly all positions in the sac; sometimes it lay close to the egg-apparatus, sometimes it was near the antip- odal cells. The egg-apparatus is of the normal structure, and com- pared to the size of the embryo-sac is small. A good deal of variation in the size of the egg and synergide was noted, and these differences did not appear to be neces- sarily connected with the age of the egg-apparatus. The BOT.—VOr, 15] CAMPBELL—SPARGANIUM. ere egg-cell was sometimes in the same plane as the synergide (fig. 8), and sometimes it was lower down and a good deal elongated (fig. 11). In either case it was nearly hyaline, with only a relatively small amount of granular contents sur- rounding the nucleus which is usualky placed near the free end of the ovum, i. e., the end which projects into the embryo-sac (figs. 8, 11). The synergide are alike, some- what smaller than the egg and filled with densely granular cytoplasm in which lies the small nucleus. All the nuclei of the egg-apparatus have a very distinct nucleolus which is especially conspicuous in material stained with anilin- safranine. The rest of the nucleus, however, does not stain strongly with this reagent, the nuclei preliminary to fertil- ization appear to have very little chromatin. The nucleus of the egg is somewhat larger than the synergidal nuclei but otherwise closely resembles them. Above the synergide there was seen in many specimens what looked like the remains of the tapetal cells (fig. 8). This mass stained strongly with Bismarck-brown but showed little structure and probably represented the disorganized remnants of one or more tapetal cells. As the younger stages were lacking in my preparations the origin of this body can only be conjectured. In none of the specimens of .S. sémplex were the separate polar nuclei observed. The large endosperm-nucleus re- sulting from their fusion is, however, conspicuous. It is much larger than the other nuclei of the embryo-sac, and has a very large nucleolus (figs. 10, 11). The antipodal end of the sac in this species is almost flat, although there may be a slight depression where the antip- odal cells are found. The early history of the latter could not be followed, as the material was all too old; but at the time the egg is mature, these cells are remarkable for their very small size, which is in striking contrast to their later development. Not infrequently where these cells had col- lapsed in the process of embedding, they could not cer- tainly be distinguished, and may very easily be overlooked. There was nothing, however, to indicate that they are ever 302 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. really absent. Where they are uninjured (fig. 12) they may usually be seen to be arranged in the same plane and are very shallow, projecting only very slightly into the cavity of the embryo-sac. The cytoplasm of these cells is finely granular but does not readily stain at this stage. The nuclei are very small and inconspicuous, and it is sometimes difficult to demonstrate them at all. In no case did they stain readily with the usual reagents. In S. Greenzt the appearance of the antipodal cells is somewhat different and this resembles more those of other Monocotyledons (fig. 21). The base of the embryo-sac is here prolonged into a narrow cavity within which lie the antipodal cells. ‘These are not usually in the same plane, but one of them lies above the other two, to which it bears much the same relation that the egg-cell does to the syner- gide. The antipodal cells in this species are much larger than in S. szmplex, and the cell contents denser, and the nuclei are larger and readily demonstrated, as they stain without difficulty. So far as my observations go upon 5S. /ongifolium and S. eurycarpum, they are more like S. Greeni than S. szm- plex in the form of the antipodals, although S. longzfolium is to some extent intermediate in character. III. FERTILIZATION. The small size of the pollen-spore and nuclei is not favorable to a study of the details of fertilization, and although in several instances the pollen-tube was detected within the embryo-sac, nothing was observed which indi- cated that the fertilization was in any way different from that of other Angiosperms. The pollen-tube after reach- ing the micropyle pushes down between the cells of the nucellus, which are not injured by its passage. On entering the embryo-sac, it apparently comes into close contact with one synergid, which is probably destroyed in most cases, although this is not necessarily the case, as in one instance, at least, the end of the pollen-tube was seen within the embryo- sac, although both synergide were still intact. The very Bot.—VOL. I.] CAMPBELLI—SPARGANIUM. 303 small size of the generative nucleus makes it extremely diffi- cult to detect, and only a few incomplete observations were made on its behavior after entering the sac. In one case (fig. 9g) a small, deeply stained body was observed within one of the synergide, and this very probably was one of the generative nuclei on its way to the egg-cell. In another instance (fig. 13) there was visible within the egg a small body looking like a nucleus, but not staining as deeply as might have been expected of the male pronucleus. In the same preparation (fig. 14) there was a small granular mass attached to the end of the pollen-tube, and looking as if it might have been discharged from it into the cavity of the embryo-sac, but there was nothing to indicate the discharge of the second generative nucleus and its fusion with the endosperm-nucleus as described by Nawaschin; however, my observations were too incomplete to warrant any posi- tive statement on this point. One of the synergide can usually be detected for a long time after fertilization has been effected, as has been observed in so many other cases. After fertilization has been effected the egg-cell enlarges very little, and after the membrane is developed about it, it remains unchanged for along time. At this time the uni- cellular embryo (fig. 24) is a more or less pear-shaped cell, sometimes in contact with the upper wall of the embryo-sac merely by a very small part of the pointed lower end, some- times having a much broader base of attachment. In this condition it remains until the sac has increased a good deal in size and the development of the endosperm is well ad- vanced. Owing to this slow development of the embryo the earlier stages between the unicellular condition and that shown in fig. 37 were missed. IV. Tue SEcoNDARY GROWTH OF THE ANTIPODAL CELLS. While the embryo-cell remains unchanged for a long time, the other structures of the embryo-sac undergo marked changes. Almost the first visible result of fertilization is a marked increase in the size of the antipodal cells. In 304 ‘CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. S. simplex where this was especially studied, and of which a preliminary account has already been published (Camp- bell, 1899), this is especially noteworthy. Before fertiliza- tion (fig. 12) the antipodal cells are extremely inconspic- uous, but almost immediately after fertilization has been effected they rapidly increase in bulk, and at the same time show other marked evidences of active growth. The cyto- plasm becomes more coarsely and densely granular, and the nuclei enlarge to several times their original size and at the same time take up stains much more readily than before (fig. 26). As the antipodals increase in size they project strongly into the cavity of the embryo-sac and form a con- spicuous nearly hemispherical body. The three original cells now rapidly divide until finally a very large mass of cells (fig. 30) results, probably larger than in any other Monocotyledon. The number of antipodal cells may finally exceed 150, a number greater than that yet recorded for any other Angiosperm. In position and general appearance the group of antipodal cells most nearly recalls that of many Graminee (Hof- meister, 1861; Fischer, 1880, etc.). An important differ- ence is that in the latter the development of the antipodal complex is completed previous to the fertilization of the egg. The method of its development, however, is very similar in the two cases. The embryo-sac in those grasses which have numerous antipodal cells has at first the ordinary number, three, and these subsequently divide to form the larger number ulti- mately developed. Hofmeister (1861) states that in the Triticea the number may be 6-12 and Kérnicke (1896) found 36 or more, a large number, but very much less than the normal number finally developed in Sparganzum simplex. The first division of the nuclei of the antipodal cells occurs shortly after the first division of the primary endo- sperm-nucleus. Following the first division, the increase in size of the antipodal cells and the divisions proceed rap- idly, while the endosperm at this stage remains slightly Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 305 developed. There seems little question, that as Westermaier (1890) pointed out in the grasses, these enlarged antipodal cells replace physiologically for the time being the endo- sperm, and elaborate food materials for the growth of the developing embryo-sac and embryo. The first divisions in both endosperm and antipodal cells take place before there has been any marked increase in the size of the embryo-sac; but later there is a rapid in- crease in the size of the ovule, which probably coincides with the first divisions in the embryo. Kérnicke (1896) in his studies on the antipodal cells of various Graminez was unable to detect any karyokinetic figures in these and was inclined to think that the divisions of the nuclei might be direct. This view appeared to be hardly likely, as the nuclei in the developing antipodal cell of Sfarganium appeared entirely normal, and the cells were actively growing and dividing, not a condition in which one would look for direct nuclear division. Fortunately a preparation was finally secured in which some of the antip- odal nuclei were actually in process of division (fig. 29), and it was seen that the division is the typical karyokinesis, which it is safe to say is the normal type of nuclear division, at least in the early stages. In the case under consideration the endosperm-nuclei were also dividing. In most cases, possibly always, the first division of the nucleus in the antipodal cells in S. sezmflex is not accom- panied by the formation of a division-wall, and the enlarged antipodal cells show plainly two conspicuous nuclei (fig. 29). Later, however, all the nuclei become separated by walls and each of the very numerous antipodal cells has evidently but a single nucleus (fig. 30). In some older ones, however, two nuclei were sometimes seen, so that it would seem that the last nuclear division, like the first one, may be independent of cell division. There is thus a cer- tain analogy with the behavior of the normal endosperm- cells which would indicate that the antipodal cells are not markedly different in nature from the endosperm-cells, ise may be really homologous with these. 306 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. The densely granular cytoplasm of the antipodal cells does not fill them uniformly, but there are usually present large vacuoles (fig. 28). The first division-walls in the antipodals of S’. s¢mp/lex are vertical, so that the mass has the form of a disc, but later transverse and oblique divisions may also occur, although the majority of the walls are ver- tical and the cells become a good deal elongated (fig. 30). As the seed grows older the antipodal cells begin to show signs of disintegration, but in no cases where sections were made had they entirely disappeared, and traces of them probably persist even in the ripe seed. The earlier stages of the antipodals were studied also in S. Greeni and S. eurycarpum, but material was not avail- able for the later development which is probably not entirely like that in S. semplex. In S. Greenzz the antipodals at the time the egg is fertilized are, as we have seen, much more conspicuous than in SS. semplex. They are also differently arranged, not usually lying in the same plane, but one of them being above the other two (fig. 33). In one instance (fig. 16) there was present above the three antipodal cells a large cell which looked like a fourth antipodal, but as this was the only case seen, there was no clue as to its origin. Otherwise the embryo-sac appeared to be entirely normal. In this species (.S. Greenzz) the polar nuclei remained sep- arate, although often in close contact (fig. 20) until after the fertilization of the egg, and in this respect it differs from \S. s¢mplex and probably from SS. eurycarpum. ‘There is also much greater variation in the size of the embryo-sac in S. Green than in S. semplex. As a rule the egg and synergide are larger also than in S. s¢mp/ex and sometimes the former show a reticulate appearance in the cytoplasm, due to the presence of numerous vacuoles (fig. 18), an ap- pearance which was not seen in the specimens of the latter species. The embryo-sac in S. Greenzz is relatively nar- rower than in .S. semplex, and there are three or four layers of cells at the apex of the nucellus, which is also larger than in that species. As we have already stated, the antipodal cells, before the fertilization of the egg, are noticeably Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 307 larger than in \S. s¢mplex, and two are placed below the third one, and lie in the depression at the base of the embryo- sac. This is much narrowed, and may account for the fact that sometimes these two lower antipodal cells take no part in the formation of the group of antipodals formed after fer- tilization (fig. 36). In such cases they remain for a long time quite unchanged, and ultimately show signs of disor- ganization. The upper of the three original cells, however, enlarges greatly and assumes a vesicular form, projecting strongly into the cavity of the embryo-sac. The nucleus of this cell, as in the older antipodal cells of S. s¢mplex, en- larges very much, soon undergoes division, and this is quickly repeated, so that four cells, arranged quadrant-wise, result. Where this was the case division-walls could be seen between the nuclei. Sometimes, however, the lower antipodal cells also enlarge (fig. 35) and the development proceeds much as in S. s¢mplex, except for the relative po- sitions of the cells. How far the antipodal cells may finally develop in S. Greenzz could not be ascertained, as the next stages were absent from my material. So far as could be judged from the few stages examined, the cells show a tendency to become more inflated than in S. semplex and the granular contents are less abundant, but these peculiar- ities, which seem also to characterize S. eurycarpum, may not be constant, and a study of the later stages in these species will be necessary before the question can be defi- nitely decided. V. Tue Empryo. The development of the embryo does not, at first, keep pace with the development of the antipodal cells and endo- sperm, and for some time it remains without any apparent change in appearance. Although a large number of sec- tions were made, it was impossible to find stages between the unicellular embryo and such advanced stages as that shown in fig. 37. It is therefore impossible at present to compare the first divisions in the embryo of S. semplex with the corresponding stages of that in S. ramoswm, the only 308 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. species hitherto examined. As the later stages agree quite closely in the two species, it is not likely that there are any very marked differences in the early stages. Nevertheless, itis much to be regretted that these early stages could not have been compared, and it is to be hoped that these may be examined soon in our American species. According to Hegelmaier (1874) the embryo cell in S. ramosum divides into three superimposed cells, but it is not clear whether the second division is in the upper or lower of the two primary cells. Before any longitudinal divisions are formed there may be one or two more transverse divi- sions, making thus a single row of four or five primary seg- ments in the embryo. A very similar account is given by Norner (1881) for the young embryo of several grasses, except that he criticizes Hegelmaier’s statement as to the time of the fourth and fifth transverse walls, claiming that these are probably not formed until after the first longitud- inal divisions. He concludes that three is the regular number of the primary segments in most Monocotyledons. The more recent investigations in the embryo of the lower Monocotyledons (Schaffner, 1896; Chamberlain, 1895; Campbell, 1897, 1898, etc.) indicate that the basal cell, where this becomes enlarged, does not divide further, and that all the further transverse segmentation is in the terminal cells, but it is probable that in Sfarganzum the segments are the product of the division of the original basal cell. So far as could be judged from an examination of the somewhat advanced embryo of |S. s¢mplewx (figs. 37, 38, 39); it looked as if the first longitudinal divisions in the terminal segment occurred earlier than Hegelmaier gives for 5S. ramosum, and it is doubtful whether here there are more than three primary segments. A comparison of these stages with Hegelmaier’s figures 5 and 13 indicated that possibly his segment 2 really belongs to the terminal segment. The basal segment never becomes enlarged as it does in so many Monocotyledons, but remains usually very small and generally divides early by longitudinal walls into three or four cells, but sometimes a cross-section shows but two Bot.—VOL. I.] CAMPBELI—SPARGANIUM. 309 cells. The absence of the large vesicular suspensor cell found in WVazas, Alisma and other Monocotyledons is largely a physiological phenomenon, doubtless connected with the nutrition of the developing embryo, and is intimately asso- ciated with the degree of development of the endosperm. Hegelmaier (1874) found that in S. ramosum the terminal segment gave origin to all the structures of the embryo ex- cept a portion of the root-apex. As he worked entirely with embryos which were simply rendered transparent, but not actually sectioned, he was unable to make out success- fully all the details of the arrangement of the tissues in the older embryos. Nevertheless, his results, on the whole, were remarkable accurate, and show a close correspond- ence to my own preparations of S. s¢mplex which were obtained from actual sections. He found that the cotyle- don and stem-apex are both products of the terminal seg- ment, and that the plerome of the root is also derived from this segment, and terminates at the wall separating seg- ments I and II. My own studies of the embryo were necessarily confined almost exclusively to S. semplex, as the material collected of the other species did not contain the proper stages of the young seed. The youngest embryos obtained are shown in figs. 37-40. While these agree in general with Hegel- maier’s figures of similar stages in S'. ramosum, there are cer- tain differences which may be noted. The basal, or primary transverse wall (~) which corresponds probably to his wall (7), is regularly lower down than his figures, indicating that the terminal cell of the young embryo is from the first larger than in S’. ramosum. It is possible that his segment II may be included in the portion above the basal wall, but this is not probable, as evidences of median vertical walls extending from the apex of the embryo to the basal wall can generally be made out, and from a study of the older stages it is evident that the plerome of the root extends quite to the basal wall, as Hegelmaier describes for 5S. ramosum. ‘The hypobasal portion of the embryo narrows rapidly, and this portion shows ordinarily but two transverse (2) July 17, 1899. 310 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. segments, instead of the three or four found in S. ramosum. The small basal segment, which is in contact with the apex of the embryo-sac, varies a good deal in size, this being correlated with the form of the original embryo-cell. If this tapers much, so as to have a small point of attachment, the basal segment remains very small and pointed (fig. 42), but if the embryo-cell is flattened where it is in contact with the wall of the embryo-sac, the basal segment of the embryo is correspondingly broader and shows more divisions. A median section of such an embryo shows the limits of the three primary segments still visible (fig. 38). This ter- minal segment is very much larger than the others, and, as already stated, relatively larger than in S. ramosum. The first divisions in all the segments are median vertical ones, and it is clear from studies of transverse sections that there are normally intersecting median walls which divide each segment into four equal quadrants. ‘The divisions are less constant in the basal segment, especially when it is small, and transverse sections of this often show only two or three cells, one or both of the second median walls being sup- pressed. The next divisions are usually vertical also, but there is evidently no absolute rule as to their arrangement. In fig. 40 is shown a series of cross-sections of a young embryo, and it is clear that the divisions are not always entirely alike in the different quadrants of the same section (see fig. 44, 6). Probably, in most cases, the first divi- sion-wall formed in the quadrant is curved, and extends from the quadrant-wall to the periphery. This is then fol- lowed by a series of periclinals which cut off the epidermis. Sometimes, however, it looks as if the first walls were peri- clinals, thus determining at once the separation of the epi- dermal layer. Hegelmaier shows much the same variation in S. ramosum (e. g. his figures 6 and 8). The variations in the basal segment have already been referred to. It may divide into equal quadrants (fig. 44, a), but more commonly there is a suppression of one or both of the second quadrant walls. Where the basal segment Bot.—VOL. I.] CAMPBELLI—SPARGANIUM. 311 is unusually large, transverse or oblique divisions may also occur (fig. 39). A transverse section of the second segment at this stage shows eight peripheral cells surround- ing four central ones. In the upper segment, the next divisions are transverse, and sometimes this gives the appearance of a complete transverse division across, and it is possible that the seg- ment numbered II by Hegelmaier may in some of his figures really refer to a secondary transverse division in the termi- nal segment. The separation of the epidermis is brought about as in the second segment, and soon after another series of similar divisions separates the central group of plerome cells from the periblem. The plerome extends to the basal wall where it ends abruptly, not being found at all in the two lower segments. In cross-section (fig. 44, c) the young plerome shows four cells, one belonging to each quadrant, but there are soon found other longitudinal walls which increase the number of plerome cells. The older embryo is not perfectly cylindrical in form, but is somewhat broader in the plane of the future cotyle- don. Sections made in this direction have the upper seg- ment nearly circular in outline, but the median wall is often somewhat oblique, so that the segment is not always per- fectly symmetrical. In this view the small basal segment usually shows a vertical division into two nearly equal cells. In the cotyledon, which constitutes the major part of the terminal segment, the epidermis is already clearly differen- tiated, but this is not always so evident in the second seg- ment, where the interlayer of cells usually undergoes another periclinal division, which is never the case, so far as I could determine, in the epidermis of the cotyledon. If the section is made at right angles to the face of the coty- ledon (fig. 42) the embryo appears more pear-shaped in outline, and the basal segment may appear undivided. Cross-sections of similar embryos show that in such cases the basal segment is composed of but two or three cells. In a number of instances the second transverse wall was very oblique (fig. 42) and might even intersect the basal 312 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. wall, and such embryos resemble very much the embryo of such grasses as 77rzticum and Hordeum, figured by Norner (1881). Hegelmaier does not show these oblique walls in his figures of .S. ramosum, and in this respect S. semp/lex is more like the grasses than is S. ramosum. The primary tissues are now pretty well defined. The central plerome-strand extends to the basal wall and is sep- arated from the epidermis by one layer of periblem cells (fig. 46). In the segment below, the superficial cells, as we have seen, divide again by periclinal walls, so that there are often two well defined layers of cells, evidently derived from a periclinal division of the primary epidermis. In the basal segment there are seldom more than four cells, and in case there is another segment between the basal one and the second segment (fig. 51) its divisions are quite irregular. . Hegelmaier’s account of the origin of the different mem- bers of the embryo in S. vamosum corresponds closely to what was seen in |S’. semplex. Much the greater part of the embryo is taken up by the cotyledon which elongates rap- idly, so that the embryo soon becomes several times longer than broad, this growth in length being mainly in the coty- ledon, the stem and root remaining short. The stem-apex originates from the terminal segment, but on one side well toward the base. Its position and origin correspond closely to those in the grass-embryo, to judge from Norner’s account and figures (Norner, 1881). In its origin from the terminal segment it differs from such Monocotyledons as Alisma and /Vaias, where the stem-apex belongs to the sec- ond segment. Zannichellca, however, has a terminal stem- apex, and Solms-Laubach (1878) describes several Mono- cotyledons with terminal stem-apices. In Zz/ea (Campbell, 1898) the stem-apex is probably derived from the terminal segment as in Sparganium. Sparganium is, therefore, in regard to the origin of the stem-apex, intermediate between such extreme forms as Zannichellia and the majority of Monocotyledons investigated, in which the stem-apex arises from the second segment. Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 313 Cross-sections of the older embryo are usually more or less oval in outline, the longer axis coinciding with the face of the cotyledon. In sections made just above the basal wall (fig. 47), the first indication of the stem-apex (st) is evident in a more rapid division in the epidermis where the cells are decidedly narrower. At this time, the section of the plerome shows about nine cells, and the original quadrant divisions are very evident. The stem- apex appears to be the product of one only of the quad- rants, and soon becomes sunk in a depression formed by the excessive growth of the basal cells of the cotyledon immediately adjacent to it (fig. 44). The stem-apex later becomes deeply sunk in the narrow cavity formed by the excessive growth of the base of the cotyledon, and this forms a sheath, such as is so frequently met with among the lower Monocotyledons (figs. 49 and 53). The plerome-strand is continued upward into the cotyle- don and downward into the root, but there is no trace of a cauline bundle. The other tissues of the root arise from the cells of the second segment of the embryo. This also divides at first by quadrant walls, and later a central growth of four cells (fig. 44, 6) is separated from an outer row of cells. The latter again divides into two by periclinal walls. The initial group of cells, which contributes to the growth of the periblem, dermatogen and calyptrogen, is derived from the four central cells which may be said to constitute the initial for all the tissues of the root except the central plerome-cylinder. Before the embryo is mature, the second leaf makes its appearance on the side of the stem opposite the cotyledon. The nearly flat stem-apex develops a protuberance on its outer side, and this quickly assumes the form of a short cylindrical body whose tissues are continuous with those of the stem, except that the plerome-cylinder connects with that of the root. The embryo rapidly increases in size as the seed ripens, and finally occupies the whole axial part of the embryo-sac. The space between it and the wall of the sac has in the 314 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER: meantime become completely filled with the endosperm. The full-grown embryo (fig. 52) is a spindle-shaped body, of which the cotyledon constitutes much the greater portion. The outer edges of the sheath at the base of the cotyledon overlap so as to completely conceal the stem-apex and sec- ond leaf (fig. 54). A longitudinal section through the base of the full-grown embryo (fig. 53) shows that the root is very broad and short. Occupying its axis is the conical plerome which in section shows about six rows of cells; this ends at the basal wall whose limits are still discernible. No single initial cell could be made out at the apex of the plerome, nor could any absolute regularity in the segmentation of the terminal cells be recognized. Connected with the main plerome- strand are smaller branches extending into the young leaves, but, as already stated, none is given off to the stem. Be- tween the plerome of the root and the epidermis there are five or six layers of cells. The origin of the other tissues of the root can be traced back to the central group of cells lying next the apex of the plerome and derived from the central cells of the second segment of the embryo. There are probably four of these initial cells, and two can usually be seen in longitudinal sec- tions. Periclinal segments are cut off from those which add to the root-cap, and lateral segments are also formed which, dividing again, form the initials for the dermatogen and periblem. The primary root of the embryo of Sparganium therefore conforms closely to the type described by De Bary (1884), as found most commonly in Monocotyledons. The root-cap is well developed in the older embryo and has the form of a biconvex lens, as the central part of the root-apex is strongly concave. A noticeable difference between the embryo of Spargan- zum and that of most other aquatic Monocotyledons hith- erto examined, e. g. Alisma, WVazas, Lilea, etc., is the ab- sence of the larger vesicular suspensor cell, so conspicuous in those forms. This cell is evidently of great importance in these plants, and directly concerned with the food supply Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 315 of the young embryo. Its absence in Sfarganium, the Graminez, and such Aroids as have been examined is no doubt to be sought in the better development of the endo- sperm in these forms, and possibly also in the presence of numerous antipodal cells, which assume temporarily the functions of the endosperm. The first division of the primary endosperm-nucleus occurs almost immediately after fertilization. At this time the endosperm-nucleus is, usually at least, near the antipodal end of the sac. The divisions are repeated until there are formed in the usual manner many free nuclei arranged in a single layer in the parietal cytoplasm (fig: 30). 4 "These nuclei are more or less flattened and possess a single large nucleolus. At the apex of the sac they are more nu- merous, and the cytoplasm is more abundant and quite sur- rounds the young embryo. As usual, the first cell-division in the endosperm takes place at the apex and proceeds to- ward the antipodal region. While the endosperm-nuclei have been dividing, and the peripheral layer of protoplasm is still very thin, the antip- odals have increased enormously in bulk and divided rap- idly to form the large mass of cells found at the antipodal end of the sac. We have already spoken of the karyoki- netic division of the antipodal nuclei and it is very doubtful whether even in the binucleate cells sometimes found in the later stages this division is ever direct, as it has been found to be in the enlarged antipodal cells of a number of plants (e. g. in Ranunculacee, Mottier, 1895). As the embryo-sac enlarges it changes its form somewhat in SS. s¢mplex, becom- ing relatively much narrower at the upper end which early becomes completely filled with endosperm (fig. 37). In the micropylar end of the sac the protoplasm early be- comes much denser, the nuclei are larger than at the sides of the sac and not confined to a single layer. The nuclei increase a good deal in size before the cell-walls are formed between them (fig. 57)- As Sparganium simplex shows very clearly the endosperm formation, this was studied in some detail. Before the divi- sion-walls are evident, the nuclei are arranged at nearly equal 316 CALIFORNIA ACADEMY OF SCIENCES, [Proc. 3D SrER. distances. The delicate radiating lines connecting them are clearly evident (fig. 57), and in these the cell-plates are soon visible, and the nuclei are thus enclosed in polygonal areas which become cells by this transformation of the cell- plates into cellulose membranes (fig. 58). The cells at the apex of the sac are quite irregular in form and completely surround the embryo (fig. 37). At the sides of the sac these cells seen from the surface are often regularly hexagonal. The endosperm-cells at the apex have much denser con- tents than those at the sides, which are almost transparent. Figure 56 illustrates the way in which the development of the endosperm proceeds centripetally. This section is a nearly median one and shows how the nuclear divisions are mainly in a plane parallel to the periphery of the sac. The first-formed endosperm-cells are at first open on their inner face, and when the nucleus divides a division-wall is found closing up the outer cell, but leaving the inner one also open. This continues, one layer of new cells after another being added to the endosperm, until finally the whole upper part of the sac is filled up, but the lower portion remains open in the center up to a late enlarged period. Free nuclei can always be seen in this open part of the sac, lying in the layer of protoplasm covering the free walls of the inner endosperm-cells. The endosperm-nuclei become a good deal enlarged before the division-walls are formed, and these enlarged nuclei generally show several nucleoli. The younger stages have only a single nucleolus and the outline of the nucleus is round and oval. As they increase in size, it was found that the nucleolus became elongated and constricted (fig. 58), finally resulting in a division into two, generally unequal, portions. This is probably repeated in the case of those which have more than two nucleoli. Each nucle- olus is surrounded by a clear area, while the rest of the nucleus presents a granular appearance. As the nucleus itself becomes more or less distorted in outline with the division of the nucleolus, and often appears lobed, it was at first supposed that we had to do with a case of nuclear Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 317 fusion, but in reality it is more nearly a case of fragmenta- tion. Hegelmaier (1885) figures a number of similar cases, and they have been often observed, but so far as I know the origin of the secondary nucleoli has not been studied. As the seed ripens, numerous very large crystalloids are formed, at least in S. sémlex, in the endosperm-cells sur- rounding the embryo (figs. 63, 64). These are beauti- fully fixed by the action of alcoholic corrosive-sublimate, and stain readily with anilin-safranine. They vary in shape, the most perfect ones being rhombic in outline. Often they were aggregated in large masses, but usually it was evident that these masses were composed of separate crystals. Sparganium simplex may be recommended as an admi- rable subject for demonstrating these bodies. The material collected of the other species was either too young or had not been treated with the proper reagents, so that I cannot speak as to the occurrence of these crystalloids in the endo- sperm of the other species. They have, however, been noted in S. ramosum Sachs (1887). No hard testa is formed about the seed of Sparganium, although this outer part of the nucellus and the integuments persist until a late stage, and probably permanently. Eng- ler’s statement (1889) that the endosperm is only separated from the pericarp by the integuments needs confirmation. In the oldest stages examined by me, there were several layers of nucellus cells still evident. The development of the hard inner pericarp is readily followed. The cells early have thickening layers deposited upon their walls, and these later become very thick and have deep pits developed in them. This sclerenchyma is rather better developed in S'. Greenzi than in .S’. simplex, and sometimes in the former the thickening is so great as to almost obliterate the lumen of the cell. Hegelmaier (1874) has studied in S. ramosum the devel- opment of the curious structure to which he applies the name ‘‘Samen-deckel.’’ The structure of this in S. szm- plex agrees closely with his account. It is formed from an enlargement of the micropylar part of the integuments, 318 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. which become very much swollen, and form two caps, one above the other. The upper part of the nucellus is also included, but the cap is principally derived from the integ- uments (fig. 59). The margins of the inner integument are prolonged upward into a conical beak which fits closely into the opening in the outer integument. The whole structure is enclosed in the space at the top of the ovary which remains as a conical cavity in the pericarp. The fruit of Sparganium is not very unlike the caryopsis of the grasses, but differs in the persistence of the integu- ments and probably also of a portion of the nucellus. Bya further compression and disorganization of these a typical caryopsis would be formed. VI. ABNORMAL EMBRYO-SACS. In Sparganium, as in most other low Monocotyledons hitherto examined, there are occasional deviations from the normal development in the embryo-sac. Some of these have already been noted by the writer (Campbell, 1897) in the case of S. ewrycarpum; indeed, it was these abnormali- ties which directed attention to Sparganium. In both S. simplex and S. Greenzi similar deviations from the type were noted. The most usual form is a multiplication of the nuclei within the embryo-sac without any noticeable increase in its size. As in such cases (fig. 68) the embryo cannot be detected, and the egg-apparatus has either not been formed, or has disappeared, the most probable explanation is that the sac had not been fertilized, but that the vegetative tissue of the gametophyte, i. e., the antipodal cells and the endo- sperm, are capable of a limited growth. This is quite com- prehensible if Sparganzum is, as it seems to be, a very low type of Angiosperm, and the case might be very well com- pared to the limited growth of the prothallium in the hetero- sporous Pteridophytes when fertilization is not effected. Other abnormal embryo-sacs are shown in figs. 65 and 69. In the first case, taken from an ovule of S. semplex, the embryo-sac was very broad and divided longitudinally Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 319 by a membrane—whether of cellulose or not was not deter- mined. Three antipodal cells having the appearance of the normal ones in a recently fertilized sac could be distin- guished. The structures at the micropylar end of the sac were not clearly distinguishable. The extremely broad form of the sac suggested the possibility of there having been two embryo-sacs formed in the ovule, but if this was the case, they had become entirely confluent. Another puzzling form in S. Greenzz is shown in fig. 69. Here the antipodal cells were apparently normal, but the polar nuclei, which were in close contact, were separated from the cavity of the embryo-sac by an evident membrane, and the upper portion of the sac was similarly shut off. Occupying the extreme upper part of this was a body look- ing like a very large nucleus, but the nucleolus (?) did not appear homogeneous, and it is possible that this apparent nucleus was really a cell, as the nucleolus-like body was quite similar in size and appearance to the nuclei of the surrounding nucellar cells. No other nuclei could be cer- tainly made out in this region, except a deeply staining small body close to the wall of a vesicle which lay below the large apical nucleus. If the latter is really a nucleus it probably means that one nuclear division in the upper part of the young embryo-sac had been suppressed, and no egg- apparatus developed. In another specimen of S. Greenzz the embryo-sac was seen to be completely filled with granular cytoplasm, and a similar condition has also been seen in »S. eurycarpum. VII. RECAPITULATION. 1. The stamen in Sparganium is of the usual type: the ripe pollen-spore shows but one generative nucleus, but in S. semplex there is regularly a structure present which prob- ably represents a vegetative or prothallial cell which was not observed, however, in S. Greenzz. 2. The early development of the embryo-sac follows the normal course; the egg-apparatus is smal] in S. szmplex and the polar nuclei fuse completely before the egg is 320 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. fertilized; in S. Gvreenzz they remain separate until after fertilization. 3. The antipodal cells are three in number and in SS. simplex are very inconspicuous; after fertilization there is a remarkable secondary growth in the antipodal cells, re- sulting finally in a large cell-mass, containing in S. semplex over 150 cells. 4. The process of fertilization, so far as it was studied, offered no anomalies. 5. The development of the embryo in S. szmplex agrees closely with that of S. ramosum, studied by Hegelmaier. There are regularly three primary transverse segments, of which the terminal one gives rise to cotyledon, stem-apex, and part of the root. The suspensor remains undeveloped, and in this respect the embryo is like that of the Gramineez. The slight development of the suspensor is associated with the complete investment of the young embryo by the endo- sperm. 6. The stem-apex is lateral in origin like that of most Monocotyledons, but is not developed from the middle of the three primary segments. 7. The primary tissues of the embryo are very early developed, especially in the terminal embryonal segment. The plerome of the root is derived entirely from the terminal segment; the initials for the other tissues of the root arise from the middle segment. 8. The development of the endosperm follows the usual course, but is rather late in forming the first division-walls. The enlarged antipodal cells doubtless function at first as endosperm. , g. The large endosperm-nuclei have often several nucle- oli which are formed by a fragmentation of the original nucleolus. 10. Large crystalloids are abundantly developed in the older endosperm-cells of S. s¢mplex. 11. The inner pericarp develops into sclerenchyma with thick, deeply pitted walls. The development of the ‘‘Samen-deckel’”’ is the same as described by Hegelmaier for S. ramosum. Bot.—VOL. I.] CAMPBELL—SPARGANIUM. 321 VIII. Tue AFrFriniTIEs OF SPARGANIUM. The genus Sparganium has until recently been associated with Zypha in the family Typhacee, but Engler (1889) has pointed out that the differences between 7yfha and Sfar- ganium are so great as to make such a union unwarranted, and he proposes placing Sfarganium in a separate family, following the Pandanacez. Until more is known about the development of the Pandanacez, however, it will be impos- sible to decide how closely the two families are related. A thorough study of some species of Pandanus would be of great value in solving this question. Hegelmaier (1874) has referred to certain resemblances in the embryo of Sfarganium to that of the Graminee, and my own study of S. semplex confirms this. The extraor- dinary character of the antipodal cells also is strongly sug- gestive of the grasses. The probably terminal origin of the single ovule is also like that of the grasses, and this it also shares with most other low Monocotyledons. Indeed the solitary uni-ovulate carpel is probably the most primitive type among the Monocotyledons, and is not the result of a reduction. In this particular S. semf/lex is probably more primitive than the forms like S. Greenzz and S. eurycarpum, in which the ovary is compound. The type of fruit in Sparganzum is not unlike that of the Graminee. While the majority of the latter have a cary- opsis, there are others in which the fruit is nut-like, corre- sponding to that of Sfarganium (see Hackel, 1890). A comparison of some of these forms with Sparganzum would be interesting. The monecious flowers of Sfarganium also are shared by some grasses, e. g. Zea, so that the possibility of an actual relation between these two puzzling families may very well be considered. Of course the resemblances may be purely fortuitous, but it is more likely that they indicate a real, even if remote, affinity, and any further study of the flower and fruit of the Graminez should consider this. In short, so far as a comparative study has been made, Sparganium shows a closer affinity with the Graminez than with any other family, unless possibly the Pandanacee. REP: 1896. 1896. 1891. 1895. 1897. 1898. 1899. 1895. 1897. 1897. 1884. 1887. 1875. 1889. 1879. 1880. 1887. 1890. 1890. 1874. 1885. 1861. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. BIBLIOGRAPHY. Bessey, C. E. The Point of Divergence of Monocotyledons and Dicotyledons. Sot. Gaz., Vol. XXII, September, 1896. Britron, N. L., AND Brown, A. Illustrated Flora of the Northern U. S. and Canada, Vol. I. New York. CampPBELL, D. H. Contributions to the Life-History of Isoetes. Anz. Bot.., Vol. V. New York. Structure and Development of the Mosses and Ferns. Mac- millan, London. A Morphological Study of Naias and Zannichellia. Proc. Cal. Acad. Sct., 34 Ser., Bot., Vol. I, No. 1. Development of the Flower and Embryo in Lilza subulata H. B. K. Ann. Bot., Vol. XII, March, 1898. The Embryo-sac in Sparganium and Lysichiton. ot. Gaz., Vol. XX VII, March, 1899. CHAMBERLAIN, C. T. Embryo-sac of Aster Nove-Angliz. Bot. Gaz., Vol. XX, May, 1895. See Coulter, 1897. CouLTeER, J. M., CHAMBERLAIN, C. T., AND SCHAFFNER, J. H. Con- tributions to the Life-History of Lilium Philadelphicum. Bot. Gaz., Vol. XXIII, June, 1897. De Bary, A. Comparative Anatomy of the Ferns and Phanerogams. Oxford. Dietz, S. Ueber die Entwickelung der Bliithe und Frucht von Spar- ganium Tourn. und Typha Tourn. A7bliotheca Botanica, Heft. V, Cassel. EICHLER, A. W. Bliithendiagrammen. Leipzig. ENGLER, A. ‘Typhacez, Pandanacez, Sparganiacee. Engler and Prantl’s Wat. Pflanzenf., Theil II, Abth. 1. FAMINTZIN, A. Embryologische Studien. Mem. Acad. Imp. Sct. St. Petersb., Ser. 7, Tome XXVI, 1879, No. 10. Reviewed in Just’s Bot. Jahresb., Jahrg. VII, 1883, Abth. 1. Berlin. FiscHER, A. Zur Kenntniss der Embryosackentwicklung einiger Angiospermen. Jenaische Zeit. fur Naturwiss., Vol. XIV. GoeBEL, K. Outlines of Special Morphology and Classifications. Clarendon Press, Oxford. Gray, A. Manual of Botany of the Northern United States. 6th Ed. HaAckEL, E. The True Grasses. Translated from Die Natir. Pfhan- zenf. by F. Lamson-Scribner and E. A. Southworth. Holt & Co., New York. HEGELMAIER, F. Zur Entwicklungsgeschichte monokotyledoner Keime. Zot. Zeit., Bd. XXXII. Morphologie des Dicotyledonen-Endosperms. ova Acta K. Leop.-Carol, Deutschen Akad. der Naturf., Vol. XLIX. Halle. HorMeEIster, W. Neue Beitriige Zur Kenntniss der Embryobildung der Phanerogamen. II.—Monocotyledonen. Leipzig. ‘Aa = Bot.—VoL. I.] CAMPBELL—SPARGANIUM. 323 1896. KGRNICKE, M. Die Entstehung und Entwickelung der Sexualorgane von, Triticum, etc. Verhandl. des Naturhistor. Vereins der Preuss. Rhetnlande, Westfalens in der Regierungsbez. Osna- briick, Jahrg. LIII. 1888. Moronc, T. Studies in Typhacee. Aull. Torrey Bot. Club, Vol. XV. 1893. Motrier, D. M. Embryo-sac of Senecio aureus. oft. Gaz., Vol. XVIII, July, 1893. 1895. Embryology of Ranunculacee. ot. Gaz., Vol. XX, July, 1895. 1881. NORNER, C. Embryoentwickelung der Gramineen. Flora, Vol. LXIV. 1887. Sacus, J. Physiology of Plants. Clarendon Press, Oxford. 1896. SCHAFFNER, J. H. The embryo-sac of Alisma Plantago. Bot. Gaz., Vol. XXI, March, 1896. 1897. Life-History of Sagittaria variabilis. Bot. Gaz., Vol. XXIII, April, 1897. 1897. See Coulter, 1897. 1878. So_ms-LausacH, H. Graf zu. Ueber Monocotyle Embryonen mit scheitelbiirtigem Vegetationspunkt. oft. Zeit., Bd. XXXVI. 1879. STRASBURGER, E. Die Gymnospermen und Angiospermen. Jena. Neue Untersuchungen iiber den Befruchtungsvorgang bei den Phanerogamen. Jena. 1893. TRuE, R. H. Development of the Caryopsis. oft. Gaz., Vol. XVIII, June, 1893. 1880. Watson, S. Botany of California, Vol. II]. Geol. Surv. Calif. 1890. WESTERMAIER, M. Zur Embryologie de Phanerogamen insbesondere tiber die sogenannten Antipoden. Nova Acta K. Leop.-Carol. Deutschen Akad. der Naturf., Bd. LVII, No. 1. Halle. 1893. ZIMMERMAN, A. _ Botanical Microtechnique, translated by J. E. Humphrey. Holt & Co., New York. 324 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XLVI. All figures except 1 and 4 were studied with Leitz lenses and drawn with a Zeiss camera. All figures slightly reduced. Figs. 4, 5, 7, 15 and 23 refer to Sparganium Greenti Morong, the others to S. stmplea Huds. mks - 10. « Ot, - 12. rae 6M 7 pete: p=10: Two female flowers of .S. simplex enlarged about eight times; the one at the right with the perianth removed. A single scale of the perianth more enlarged. Longitudinal section of the ovary showing the single ovule; ma, embryo-sac. Oc. 1, obj. 3. Pistil of S. Greenii; X about 8. Section of the ovary of S. Greenii. Oc. 1, obj. 3. Sections of ripe pollen-spores of .S. simpleac, showing the large vege- tative, and smaller generative, nucleus; fv, prothallial cell. Oc. I, im. x. Sections of younger pollen-spores of S. Greenii; the nucleus is still undivided, and no prothallial cell is visible. The clear space on the surface probably marks the place of exit of the pollen-tube. Upper part of the embryo-sac of .S. stmplea just after the entrance of the pollen-tube; 0, the egg-cell; sy, one of the synergide; /, pollen-tube. Oc. 1, im. +5. Another section of the same embryo-sac; the small dark body within the synergid is probably one of the generative nuclei from the pollen-tube. The antipodal end of the same embryo-sac, showing the single large endosperm-nucleus and two of the antipodal cells. The egg-apparatus and endosperm-nucleus from a mature embryo- . sac. Oc. I, im. 7. (a) Two antipodal cells from a mature embryo-sac; (4) outline show- ing the position of the three small antipodals. Oc. 1, im. 75. # Section of the egg-apparatus showing the entrance of the pollen- tube, f¢. A small body (the generative nucleus ?) is visible within the egg; both synergide are still intact. Oc. 1, im. 7j. Another section of the same; the pollen-tube has apparently dis- charged a small granular mass into the embryo-sac. Section of the nucellus and mature embryo-sac of S. Greenit. Oc. I, obj. 7. A large cell (an extra antipodal ? ), lies above the three antipodal cells. The antipodal region of fig. 16 more highly magnified. 17 and 18. Two sections of the egg-apparatus of S. Greenii. Oc. I, . 5. . 20. - 2. . 22: s 23. im. zs: Embryo-sac of S. Greenii, showing egg, 0, and two antipodals, ant. Os TxOby..7: Unusually large sac of the same species; the section shows the two synergidz and the still separate polar nuclei. Oc. 1, obj. 7. Two of the antipodal cells from the same sac more highly magnified. Section of egg-apparatus of S. Greenii. Oc. 1, im. 5. Endosperm-nucleus from a similar sac. Proc Can Acan Scr3" Ser. Bor. Vowl. [ CAMPBELL] FLare’ XEV] my CALIFORNIA ACADEMY OF SCIENCES. [PRroc. 3D SER, EXPLANATION OF PLATE XLVII. Figs. 33-36 refer to S. Greenii, the others to S. simplex. . 2h. Biot 20: Aas waa) . 20. : 90: ; at, Be: sade me” ATs Apex of fertilized embryo-sac with one-celled embryo. Oc. 1, im. 75. An embryo of the same age, but with a broader base of attachment. One synergid could be still made out. A recently fertilized embryo-sac; one of the enlarged antipodals, hae shown. The endosperm-nucleus has divided once. Oc. I, Obi: 7. The antipodal region of the same more highly magnified; the three enlarged antipodal cells lie in the same plane; one of the two endosperm-nuclei is shown. Enlarged antipodal cells from an older sac; each of the cells shown has two nuclei; the whole number of nuclei was eight or nine. Oc; E, im. is: Two sections of the antipodal group from a still older sac; some of the nuclei are dividing; twelve antipodal nuclei could be distin- guished. Oc. 1, im. 74. Nearly median section of the antipodal group from a much older sac. Oc. 1, obj. 7. Divisions have formed between all the nuclei; numerous free endosperm-nuclei, e7., are present. Part of the lining layer of protoplasm with a single endosperm- nucleus. Oc. 1, im. 7. Surface view of a portion of the nucleated protoplasmic layer lining the wall of the embryo-sac. Antipodal cells from the unfertilized sac of S.,Greentt. Oc. 1, im. 75. Antipodal group shortly after fertilization; the upper cell has en- larged and divided; the two lower ones show indications of disintegration. Antipodal group of SS. Greenii in which all the cells appear active. A somewhat older group of the same species; growth is mainly confined to the upper cell. Upper end of embryo-sac of .S. staplea showing the young embryo surrounded by the endosperm-cells; the central nuclei, e7, are still free. Oc. 1, obj. 7. The embryo from fig. 37 more enlarged. ae section of a similar embryo. Oc. 1, im. 75; 2%, the basal wall. Series of three cross-sections of an embryo of about the same age; a, the lowest segment. Two sections of an older embryo, cut in the plane of the cotyledon. Oc, 1; obj 7. Figs. 42, 43. Longitudinal sections cut at right angles to the plane of the g. 44. 5 ans nAGs a7 . 48. . 49. . 50. et i cotyledon; the basal wall is decidedly oblique. Oc. 1, obj. 7. Three cross-sections of an embryo of about the same age as 42; the lowest segment is divided into four cells. Oc. 1, obj. 7. Upper part of embryo-sac showing the embryo surrounded by the densely granular endosperm-cells; the granular cell-contents of the embryo are not shown. Oc. 1, obj. 7. The same embryo shown in fig. 45, showing the differentiation of the primary tissues, the plerome ends at the basal wall, 22. A are section of an embryo of about the same age. Oc. I, obj. 7. Section in the plane of the cotyledon of a similar embryo, with very regular arrangement of the tissues. Oc. 1, obj. 7. Median section of an older embryo, showing the stem-groove, s?. Oct, Obl 17- Embryo of about the same age, cut in the plane of the cotyledon. Median section of the base of the same embryo, more highly mag- nified; #/., the plerome. Proc. Cat, Acan.Sc1.3? Ser. Bor. Voul. [Camppen.) Pears XLVI 328 Fig. Fig. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XLVIII. All figures except 69, refer to S. simplex. Raye: g- §3- 69. Two sections of a nearly mature embryo. Oc. 1, obj. 3. Root-end of a somewhat younger embryo. The position of the original transverse, or basal wall (2) is still evident, 2’, the second leaf; st. the stem-apex. Oc. 1, obj. 7. Three cross-sections of a mature embryo. Oc. 3, obj. 3. Sections of the young seed before the endosperm has filled the embryo-sac; e, embryo; 77!', 2m”, integuments. Section through the layer of developing endosperm lining the embryo- ‘sac; many of the nucleiare dividing. Oc. 1, obj. 7. Larger endosperm-nuclei from the base of the sac at the time the walls are first beginning to form; some of the nuclei have more than one nucleolus; the fine fibres connecting them are plainly visible. Oc. 1, im. 7. Transverse section of recently formed endosperm-cells showing the nucleolus fragmenting. Oc. 1, im. 5. Upper part of the young seed showing the enlargement of the micropylar portion of the integuments. Oc. 3, obj. 3. Young sclerenchyma-cells from the pericarp. Oc. 1, obj. 7. Fully developed sclerenchyma-cell showing the deeply pitted walls. OG; 15)-0b};, 7: Surface view of the pits more highly magnified. Endosperm-cells from nearly ripe seed, showing large crystalloids. Ce, 5, Ore: 4. Separate crystalloids more highly magnified. Abnormal embryo-sac of .S. simplex; the sac divided longtudinally with the endosperm-nucleus on one side. Three antipodals could be distinguished. Oc. 1, obj. 7. Micropylar end of the same sac. One of the antipodal cells of the same. Abnomal embryo-sac (perhaps unfertilized ?) ; there has been little increase in size of the sac, but numerous endosperm-nuclei are present, and the antipodals have enlarged and divided. Oc. 1, Obj7: Two sections of an abnormal embryo-sac of S. Gveenii,; three antipodals could be seen, and the polar-nuclei (f.7.), in close contact, were separated from the body of the sac. The upper part was also shut off by a membrane, and no definite egg- apparatus could be distinguished. Oc. 1, im. 7. Pate XLII [ CAMPBELL | E = & ao a a & 2 = a B A MORPHOLOGICAL STUDY OF THE FLOWER AND EMBRYO OF THE WILD OAT, AVENA FATUA L. BY WILLIAM AUSTIN CANNON, Assistant in Botany, Leland Stanford Junior University. CONTENTS. Pirates XLIX-LIII. PAGE I. Tee RRO TION cy. dbs cecccids csc c ceca haneee ss yee wn de dt dalemins te 329 II]. ORIGIN OF THE FLORAL ORGANS. ......--ee sere eee eeeeee ceeeee 331 We PeUrre SEAMEN: 225 ol cict udicta Sia s gay vie ole! tip ala: 5 09 @, sini gon Bn bial wate 9 eas 335 BMP clue) EMER VO-SAC» <1= 5 bt) ecco (oo nn ode alesinic sid ale sceale he cla ts Ge ane 337 V. FERTILIZATION AND THE BUILDING OF THE ENDOSPERM......... 342 RON Pre ROWNISE VEN neds cc k vida sna oe nis Sinieiete ee o Seite alewe ones ad eatsis aiamn mms 343 RUN TERREA TIC F-1 his ect ah aloes oie wel sleidalera amie biel éjae otic ee Fa aieehacemiadels 351 Bree TOGRAPHY fi oe ode he cin a tiie rea wiele eine wie hw ni dne nls wala aa nitnin oietee 354 Bee ANA TION. OF (PLATES cfc o)< «524 vlc oes one 08 6 nan cistern bulge arene 356 I. INTRODUCTION. Tue grasses are said by Hackel to be a very isolated family, sharply defined from the other low Monocotyledons to which they are bound by only obscure relationships. For this reason they have been variously placed by systema- tists, but are now generally and probably best regarded as near the Aroids and the Palms. The life history of such grasses as have been studied justifies this conclusion, and it is the object of this investigation to add something to our knowledge of these forms in order that their mutual relation- ships may be better understood. Enough morphological work has already been done on types allied to the grasses to form a basis for comparisons. It remains therefore only for more extended and more thorough studies to be made on grasses before comparisons can be instituted. There have already been a few such studies published. The most important recent contributions to our knowledge of the life history of grasses have been by Lermer and Holzner (1888), and Koernicke (1896). The original of the former con- tribution was unfortunately not at hand and I was obliged [329] April 14, 1900. 330 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. to depend on citations and fragmentary reviews for my knowledge of it. It is an exhaustive work, describing the complete development of the floral and reproductive organs of the cultivated barley as well as its embryology. Koer- nicke, on the other hand, studied the development of the stamen and embryo-sac of the wheat with especial reference to cytological phenomena. ‘Two studies which were made before the introduction of the modern methods of imbedding and sectioning, viz. by Fischer (1880), and Noérner (1881), were consulted in the preparation of this paper. Fischer’s study deals with the development of the embryo-sac of various grasses, while that of Norner treats of the embry- ology alone. All of the contributions mentioned above refer to grasses belonging to the branch of the family known as the Poacez, the members of which, besides being closely related, have very similar habits and habitat. It is to be greatly desired that future studies of the Gramineze may include representative grasses of various habits and habitat of both branches of the family. In this way an adequate knowl- edge of the inter-relationships of the grasses, as well as their relationships to other Monocotyledons may be obtained, and also a better idea of the origin of plant structures. In addition to the papers already named there is a great mass of work on the comparative morphology of the different parts of the mature flower and of the mature embryo. Be- ginning with Malpighi (Omuza opera, 1687), and Gartner one hundred years later, the series extends with little inter- ruption to the present. For a review of these works the reader is referred to the studies by Bruns (1892), and to the recent one by Kennedy (1899). Because the flower of the wild oat is the type grass flower, (it is described and figured in any good text-book of botany), it was selected for this morphological study. The wild oat is supposed to be the ancestor of the cultivated species, or at least the nearest relative of that ancestor, and is one of our commonest grasses. It was presumably introduced into California at an early date from Southern Europe, and has now a wide range throughout the state. Bot.—VOL. I.] CANNON—AVENA FATUA., 43% II. ORIGIN OF THE FLORAL ORGANS. The spikelet of Avena branches sympodially, the apex of the young spikelet developing into flowers and_ finally pistils. Each flower arises below the apex of the pri- mordium of the spikelet and grows at right angles to it, so that the rachilla is sharply angular when young (fig. 1). The primordium of the spikelet consists of three well defined series of tissues: (1) a dermatogen, (2) a single layer of large hypodermal cells—the periblem—which sur- rounds (3) an axial strand—the plerome cylinder. As the primordium increases in length the cells of the epidermis and of the periblem divide by anticlinal walls and form a continuous cap to the plerome. The plerome arises from a single cell, or perhaps a group of cells, that terminates the axial strand (fig. 3). All of these cells are well filled with protoplasm, are without conspicuous vacuoles, and have relatively large nuclei. The floral organs all arise in much the same way upon the periphery of the primordium, beginning with the lower glume and ending with the carpel. Each begins as a ridge of tissue that extends part way around the primordium of the flower.. The origin of each floral organ will be spoken of in the order of its appearance. The lower glume and the young flower apex appear to arise simultaneously just below the tip of the primordium of the spikelet (fig. 4). At first these two rudiments are very similar, but the flower apex soon becomes broader, and the lower glume somewhat deeper by its growth and by the stretching out of that part of the rachilla that is immediately below it. At this time the glume is a semicircular organ, club-shaped in longitudinal section, composed of a well defined dermatogen enclosing about three layers of cells. As the lower glume increases in length it takes a direction which coincides with the direction of that part of the rachilla from which it springs, but it soon bends toward the tip of the spikelet (figs. 1 and 2). At that stage in the development of a flower when all of the floral organs dre first to be distinguished, a narrow outgrowth appears near 332 CALIFORNIA ACADEMY OF SCIENCES. [PrRoc. 3D SER. the base and onthe inner surface of the lower glume. This outgrowth is at first on a level opposite the lodicule, but as the glume increases in length, it is carried upward some- what (figs. r and 2). It becomes longer and broader and forms the lamina of the mature lower glume, while the original rudiment persists as the awn. The upper palet and the lodiculz arise apparently at the same time and from the same rudiment (fig. 5). A cres- cent-shaped ridge of tissue extends nearly around the pri- mordium. ‘This ridge does not form on the anterior aspect, but from the ends the lodicule develop, and from the anterior face the upper palet. The upper palet grows as the adjacent organs develop. When it can first be plainly recognized it is composed of three layers of cells, and later becomes thicker by periclinal divisions of the cells at its base (figs. 5and 6). The upper palet is the last of the protective organs of the oat-flower to develop. The lodicule appear as rounded shoulders jutting out from the lateral surfaces of the primordium and mark the ends of the rudiment that gives rise to the upper palet (fig. 5). The apex of the young flower grows up and leaves the lodicule in a lateral position below its tip. The lodicule are at first triangular with the broad base resting against the surface of the primordium. They gradually acquire the oblong shape characteristic of the mature organs. The stamens of the wild oat arise as conical outgrowths at equal distances apart around the periphery of the apex of the flower. Figure 1 shows the relation of the anterior stamen to the lower glume; and figure 7, which is taken from a longitudinal section made a little to one side of the center of the flower, shows one of the lateral stamens and its relation to one of the lodicule. The stamens develop early, the filament appearing before the lamina of the lower glume is well differentiated and while the carpel is still small. It increases in length by cell-division in all parts and is always relatively slender. The pollen is generally shed before the stigmas of the flower have protruded from the spreading glumes. Bot.—VoL. L.] CANNON—AVENA FATUA. 333 The carpel is the last of the floral organs to arise. It appears as a single ridge of tissue on the anterior side of the floral axis (fig. 6), and gradually encircles the apex of the young flower (fig. 6a). The young carpel is club- shaped in longitudinal section, and its structure is suffi- ciently indicated by the figures. Soon after the carpel has completely surrounded the tip of the flower axis, two pro- tuberances appear on the sides and form the beginnings of the stigmas; later the rim of the carpel closes and shuts in the ovule. A correct interpretation of the relations of the parts of the grass flower is of great importance, because upon it depends a right understanding of the relationships of this isolated family. These relationships are based mainly on the number and arrangement of the floral organs. The origin and development of the gynceceum and androeceum of the grass flower, as well as the origin and development of the organs subtending these, are now fairly well understood, but the morphological significance of some of these organs is very much in doubt. It is not known, for instance, whether the lodicule represent a perianth or not, an impor- tant basal fact in determining the position of the grasses. There are concerning the lodicule two widely different views which have been held by botanists. In the first, the lodiculz are considered to be the stipules of a leaf which has been suppressed, or the parts of a leaf of which the middle is wanting; and the posterior lodicula when present is held to be, according to this hypothesis, a leaf or palet which alternates with the lodicular palet and which there- fore continues the distichous arrangement of the lower glume and upper palet. Or, in other words, according to this interpretation of the grass flower there are represented four alternating palets of which the two lowest persist, the others being rudiments. This explanation of the relation of the lodicule to the other organs of the grass flower is opposed to the law which is sometimes said to govern leaf arrangement among the grasses, by which the leaves of a branch are inserted on a plane at right angles to that of tthe 334 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. leaves of the main axis. Further, this hypothesis excludes the idea of a perianth in the grasses, and with this a very important clue to the connections between the grasses and other low Monocotyledons is overlooked. The second interpretation recognizes a perianth which is represented by the lodicule. The two anterior members are present in most grasses, and the trimerous perianth is completed by a posterior member in the bamboos and certain other grasses. Neither of the hypotheses set up to explain the significance of the lodicule is entirely satisfactory—the first explanation for the reasons given, and the second, mainly because of the position of the lodicule‘in most grasses. The relation of the lodicule to the upper palet in Avena (and in some other grasses as well) in which there is a common origin for both, makes it difficult to look upon the lodicule as members of a perianth, for this, if present, ought to originate and therefore be placed on a higher plane than the upper palet. This intimate connection of the lodiculz and upper palet is met also in Oryza, Zea, and Solenache (Eichler). The lodicule vary greatly in number in the different grasses and assume various positions, as in the Panicez they are mainly outside the upper palet, and in the Ehr- harta-species similar bracts are associated with the lower glume. From this it seems reasonable to conclude that not all of the lodicule can represent rudimentary perianths. In addition to this variation in position of the lodicule there are present in the rye (Eichler) and certain other grasses four anterior lodiculz placed on two planes; the two lower are connected with the upper palet (‘‘ stipular lodicule’’) and the two upper, originating evidently on a higher plane, are regarded as being two members of a perianth (“ peri- anthal lodicule’’). A comparison of the lodicule of Avena with those of the rye shows them to be homologous to the stipular lodicule of the rye, but in Avena the perianthal lodicule are wanting. In the bamboo a trimerous perianth of three lodicule is present, and in addition to this, two stipular lodicule also. The upper lodicule of the bamboo, by reason of their position and number, have been rightly BotT.—VOL. I.] CANNON—AVENA FATUA., 335 considered to make up a perianth. They are homologous to the perianthal lodicule of the rye and cannot, therefore, hold similar relations to the lodicule of Avena. I conclude then from a comparison of the grass flowers that the lodic- ule in Avena do not represent a perianth, but that the perianth is entirely wanting; they are homologous to the stipular lodicule of other grasses. From this it would appear therefore that not all those organs called lodicule are necessarily or even probably the same morphologically. In some grasses they are survivals or rudiments of a perianth, in others they are survivals or rudiments of extra-floral leaves, and in still others, where two rows occur, the inner row is morphologically perianth, the outer extra-floral bracts, and there is no evidence (Eichler) of a double perianth. ~ Ill. Tue STamen. The development of the stamen of Avena, as far as fol- lowed, corresponds rather closely with that of the stamen of wheat, (Koernicke, 1896). The stamens originate as crown-shaped projections at equal distances apart on the periphery of the floral axis. A cross-section shows the young stamen to be oblong in form (fig. 8). By more rapid growth in the portions where the loculi are to appear, the outline becomes irregular and characteristic of the stamens of Angiosperms (fig. 9). A constriction occurs very early at the base of the stamen, which indicates the beginning of the filament (fig. 7). The subsequent growth of the stamen until the time when the pollen grains are ripe is due almost entirely to the elongation of the anthers—the filament growing only immediately before the pollen is scattered. The primary tissues—plerome, periblem, and dermatogen—are clearly marked in the young stamen. As usual in Angiosperms, the pollen is derived from the peri- blem. In the increase in diameter of the anther, preceding the cutting off of the archesporium, the cells of the periblem in the parts where the loculi are to appear become larger 330 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. than the intervening periblem cells and also than those of the plerome. They become separated from the epidermis by periclinal walls of more or less regular formation (fig. 10); but the hypodermal cells between the large periblem cells described divide by anticlinal walls only. This cell- formation results in the lobed condition characteristic of anthers. A cross-section of an anther in which the loculi are visible shows one conspicuously large cell surrounded by two series of periblem cells and the epidermis. Of these three concentric cell series the middle one forms the endothecium, and with the epidermis persists until the anther is ripe. The inner series, according to Koernicke, divides by periclinal walls to form the tapetum and a middle layer, both of which are resorbed in the later stages of the development of the anther. This point, however, I did not see in theoat. The large cells which occupy the middle of each loculus and form, as longitudinal sections show, a single row, constitute the archesporium. Later these divide by radial walls and thus give rise to the pollen mother-cells. A median longi- tudinal section through a loculus in which the pollen mother- cells are formed (fig. 11) shows two rows of spore mother-cells, each of which reaches to the walls of the loculus. The history of the division of the mother-cells into tetrads I did not follow. Koernicke gives a detailed account of this for the wheat, and we may suppose that the history of spore formation in the two nearly related genera is very similar. The mature pollen spore is enclosed by two integuments, the inner of which is very delicate. The spore is densely filled with coarse granules of starch. Three bodies that stain strongly with the reagent used (Haidenhain’s iron- alum-hzmatoxylin) are present in the spore. The largest is oval and stains less strongly than the two spindle-shaped ones. These three are respectively the vegetative and the two generative nuclei. Some difficulty was encountered in distinguishing the nuclei from the cyto- plasm in the generative cells, and it is probable that the Bot.—Vot. I.] CANNON—AVENA FATUA. 337 nuclei form a larger portion of the cell contents than is indicated in the figure (fig. 12). Koernicke does not men- tion a division of the generative nuclei in the pollen grain of wheat, and this would be an apparent exception to the statement by Strasburger (1884), that the division of the generative nuclei is a constant character of the grasses. The generative nuclei in the pollen grain of Avena occupy various positions with relation to the vegetative nucleus, but are always parallel to each other. The vegeta- tive nucleus is oblong and as large as both the other two nuclei combined. The development of the pollen grain was not followed further. IV. Tue Empryo-Sac. The initial cell of the embryo-sac in the oat is of sub- epidermal origin. It is the middle one of the epidermal cells of the primordium, as is generally the case (Fischer), and it is continuous with the axial plerome. At the time when both of the integuments of the ovary are clearly dis- tinguishable, the archesporial cell is easily made out by reason of its relatively large size. At this stage it occupies in the oat, as in the wheat (Koernicke), over one-third of the nucellus, and is eight to ten times as large as the sur- rounding cells. There is no apparent difference at this time between the archesporial cell and those of the nucellus in density of cell contents. When the archesporial cell is first recognizable the ovule is atropous. Parallel with the development of the sister cells and of the embryo-sac, the cells in the rachilla side of the nucellus divide and grow more rapidly than those in the Opposite side, and push the micropylar end over so that finally the micropyle occupies a position almost opposite from that of the archesporial cell. During the revolution of the ovule (macrosporangium) the ovary increases greatly in size, and, from being somewhat flattened at first, it becomes spherical and at last oblong. The size and posi- tion of the ovule is intimately connected with the develop- ment of the archesporial cell and the embryo-sac. 338 CALIFORNIA ACADEMY OF SCIENCES, [PRoc. 3D SER. When the axis of the ovule forms a right angle with that of the flower the archesporial cell has elongated, keeping pace with the elongating nucellus, and its nucleus has undergone one division (fig. 14). The cell is densely filled with granular protoplasm; no vacuoles are visible. There are in Avena as in Ezchhornia (W. R. Smith, 1898) apparently three ways in which the sister cells may be formed. In the first, the nucleus of the archesporial cell divides after the elongation of the cell, the two resulting nuclei divide, and the four sister nuclei lie free in the pro- toplasm of the cell, no walls separating them (fig. 15). Or (second) after the nuclei have divided, walls may appear between them (fig.17). However, owing to faulty methods of staining, I could not in every instance be sure of the presence or absence of the separating walls, but the phenomena were observed at various times during the course of the study. In the third manner of formation of sister cells, dividing walls followed each nuclear division as in the section represented in figure 16. This variation in the formation of sister cells was not described by Koernicke for Triticum, or by, Fischer for any of the grasses studied by him. Fischer did however describe a condition in the genus JZelica which he says is constant, and is analogous to what has been described above for Avena. In Melica the division of the nucleus of the archesporium is followed in all cases by the formation of a dividing wall. The two nuclei thus formed divide each once, but without cell- walls being formed; so that in JZe/ica there are two cells with two sister nuclei in each cell. It would be of import- ance to know whether this condition is as constant as Fischer states, or whether there really is in MJelzca that variation in the formation of the sister cells which is appar- ently the case in Avena and in Ezchhornia. In whatever way the sister cells are formed four super- imposed nuclei always result, and of these the lowest becomes the spore mother-cell, the others becoming resorbed. The uppermost of the sister cells is the first to disappear (fig. 17), and after it the two next lower ones in Bot.—Vot. I.] CANNON—AVENA FATUA., 339 succession, if cell-walls have been formed between them, or simultaneously if the nuclei are not separated by walls (figs. 18 and 19). The macrospore thus formed occupies the position recently occupied by the sister cells, and extends - to the epidermis. No tapetum is formed. When they are present the walls separating the sister cells. are not so strongly swollen as in 7retzcum (Koernicke) and in A Jlope- curus (Fischer, fig. 32). In this regard the oat agrees with the figures given by Fischer for Seslerza. The wall next to the epidermis in Avena is somewhat swollen as in Sesleria (i) ce. figs 10). The macrospore nucleus divides into two nuclei which lie, one in the micropylar end and the other at a point distant about one-third the length of the embryo-sac from its chalaza end. The chalaza nucleus divides at right angles, and the upper nucleus divides parallel to the long axis of the embryo-sac (fig. 20). One large vacuole sepa- rates the two nuclear groups; and another separates the chalaza nucleus from the end of the embryo-sac. Each nucleus of each group divides again, the upper group at right angles to the plane of the first division. The embryo- sac now contains eight free nuclei (figs. 21 and 21a). During these divisions of the nuclei, the embryo-sac has increased greatly in length and diameter, and has absorbed the adjacent cells of the nucellus. Many large vacuoles have appeared in the protoplasm of the embryo-sac; promi- nent among them are the large ones surrounding the proto- plasmic bridge that connects the two groups of nuclei, and the vacuole in the chalaza end. In the micropylar end of the embryo-sac two (sister) nuclei form the synergide; and one that lies at one side of the synergids, the egg; while the sister nucleus of the egg becomes the upper polar nucleus (figs. 21 and 22). Three of the nuclei in the chalaza end of the sac give rise to the antipodal complex, and one forms the lower polar nucleus. The egg is at first spherical, but as it increases in size it becomes oblong, and finally, when nearly ready for fertili- zation, balloon-shaped (figs. 24 and 25). It occupies a 340 CALIFORNIA ACADEMY OF SCIENCES. [PRoC. 3D SER. lateral position as regards the synergidz, which is the posi- tion of the egg in Ses/erza also (Fischer). The synergidz are flask-shaped and do not appear to contain vacuoles. The egg-apparatus increases greatly in size but shows no other change up to the time of fertilization. The upper polar nucleus moves to meet the lower polar nucleus in the region of the antipodal nuclei (figs. 21 and 22). The two nuclei then move towards the egg and remain in close proximity to it, in the protoplasmic bridge that con- nects the egg-apparatus and the antipodals. The polar nuclei do not fuse until about the time of the fertilization of the egg. The exact time was not determined. The nu- cleoli of the polar nuclei are the largest in the embryo-sac and with the ordinary staining reagents become deeply colored. The peculiar behavior of the polar nuclei of the oat does not agree with that in the grasses examined by Fischer; but Koernicke indicates that some such action of the polar nuclei occurs in 7rztzcum, as he states that the two polar nuclei move towards each other. The sides of the polar nuclei that are in contact with one another are flattened, and each cell retains its delicate cell-wall up to the time of fertilization, when they fuse to form the endo- sperm nucleus. The time of the fusion of the polar nuclei of the oat appears to be the same as that in the wheat. There is no rule as to which side of the polar nuclei will come in contact, so that a line separating them may be parallel to the long axis of the embryo-sac, at right angles to it, or it may be at any angle between the two. The endosperm nucleus is imbedded in a dense strand of proto- plasm which connects the antipodals and the egg, and strands radiate from this to the walls of the embryo-sac. This last is especially true of the younger sacs, the proto- plasmic threads disappearing in the older stages. The protoplasmic bridge persists however until the antipodals disappear. The embryo-sacs of all the grasses that have been studied, except Melica (Fischer), show a multiplication of the anti- podal cells. This phenomenon has also recently been BoT.—VOL. I.] CANNON—AVENA FATUA. 341 observed in the family which is probably the nearest ally of the grasses, viz. the Sparganiacee, and in the Araceze.! The antipodal nuclei of the oat begin to divide as soon as the lower polar nucleus is separated from them. They divide repeatedly and each nucleus becomes enclosed in a globular mass of protoplasm. The increase in number is accompanied by an increase in the size of the cells and also of their nuclei. A good idea of this growth of the antipodal complex may be had by comparing figures 22, 23 and 25. The multiplication of the antipodals and their increase in size takes place at the expense of the nucellus (cf. figs. 23 and 25). The greatest number of antipodals is found at the time of the fertilization of the egg. This number was not exactly determined, but it was at least 36, which corresponds with that given by Koernicke for the wheat. There is uniformly in Avena as in Triticum one nucleus in each cell. Fischer describes three antipodal cells only for Alopfecurus and Sesleria, but says that there are four nuclei in each. JZelica, on the other hand, does not show multiplication of the antipodals in any way. When they begin to divide, the antipodal cells of Avena occupy a position near, but not in, the chalaza end of the embryo-sac, and a line drawn through the long axis of the embryo-sac would bisect them. As the embryo-sac develops, the anti- podals assume a lateral position and retain it until they disappear. The antipodal complex shows, in section, a characteristic crescent form; the nuclei are large and stain deeply with the common staining agents. No vacuoles are visible in the young antipodal cells, but they appear as the cells grow, and are increasingly marked during their dissolution. The disorganization of the antipodals begins simulta- neously with the formation of the endosperm, and usually they can no longer be seen at the time when the micropylar end of the embryo-sac is being filled with endosperm. Fig- ure 27 shows the condition of the antipodals when the endo- sperm is beginning to form. A few large vacuoles in each 1 Campbell, Bot. Gaz., March, 1899, and Proc. Cal. Acad. Sciences, 1899. 342 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. cell can be seen. Almost the last stages in the existence of the antipodals is shown in figure 28. The protoplasm of the antipodal cells is the first portion of the cells to disappear; traces of the antipodal nuclei are frequently met when they are quite disassociated from the protoplasm of the cells. The function of the great antipodal complex, in those forms in which it is present, undoubtedly varies with the time of its formation. In the oat, as in the wheat (Koernicke), the antipodals are formed prior to fertilization and disappear with the multiplication of the endosperm nuclei, being con- sumed by the endosperm in its development. In Sfar- ganium (Campbell) the antipodals multiply enormously after fertilization, and evidently function as endosperm, and consequently serve to nourish the growing embryo. V. FERTILIZATION AND THE BUILDING OF THE ENDOSPERM. The pollen-tube enters the embryo-sac by pressing apart the cells of the micropyle. The two synergide do not usually disappear at this time, although one seems to do so. This point needs further study. One, or perhaps the two synergide, may sometimes be observed partly resorbed while the embryo is still in the proembryo stage, and it is likely here, as in the wheat (Koernicke), that both syner- gid are consumed as nourishment for the young embryo. The phenomena associated with the union of the sexual nuclei, as well as the further history of the pollen-tube, the vegetative nucleus, and second generative nucleus were not followed. ; The endosperm nucleus begins to divide about the time when the egg is fertilized; but whether this precedes or follows the union of the nuclei was not determined. At the first division of the egg a few endosperm nuclei may be seen in the thin protoplasmic lining in different parts of the embryo-sac. These nuclei vary a great deal in size, and in shape also, from spindle form to a flattened sphere. Each endosperm nucleus generally contains several (three to eight) nucleoli. In the early stages in endosperm formation BoT.— VOL. I.] CANNON—AVENA FATUA. 343 the nuclei divide in all parts of the embryo-sac_ wher- ever they are. With their increase in number they become larger in size, cease dividing, are usually spherical, and become enclosed in rather dense masses of protoplasm, which later become separated by cell walls. The lumen of the embryo-sac begins to fill first around the embryo, where the endosperm cells that closely invest it are long, some- what crescent-shaped, and very dense. The nucellus and embryo-sac increase greatly in length after the fertilization of the egg, the embryo-sac absorbing more and more of the nucellus cells, until little more than a single layer separates it from the inner integument. A change also takes place in the integuments. Before fertili- zation and more especially during the youngest stages of the embryo-sac, the two integuments are quite similar, each being composed of two layers of uniform cells. After fertilization the outer integument is compressed and finally destroyed, and the inner one takes on heavier cell-walls and becomes the outer seed-coat. VI. Tue Emepryo. The odspore, which is relatively small and enclosed by a delicate membranous wall, increases considerably in size after fertilization and before any division occurs. The cytoplasm is granular and with few vacuoles. An interest- ing variation in the cytoplasmic structure of the cells of very young embryos was met in several instances. The cytoplasm appeared very coarsely granular and _ net-like. No conspicuous vacuoles were present in such embryos; growth and cell-division as far as it had occured were normal. No embryos beyond the proembryo stage pre- sented this peculiar appearance of the cytoplasm. The first division of the egg occurs near the distal end and is transverse, either at right angles to the long axis of the young embryo (proembryo), or at an angle more or less acute to it. I found one abnormal case in the first division of the egg, in which the cell-wall was a longitudinal one. Figure 29 shows an egg in course of the first division 344 CALIFORNIA ACADEMY OF SCIENCES. [Proc. 3D SER. in which the wall is to be at right angles to the long axis of the proembryo. The section was cut not exactly in the median plane, and the figure shows what appears to be a large vacuole near the point of attachment of the embryo. Examinations of the neighboring section show that this is not a vacuole, the only visible vacuole present being in the neighborhood of the dividing nucleus. When the first wall is formed, however, vacuoles occur in both cells, and as the further history of the embryo is traced, the greatest number of these are to be found at and near its base. The second division of the embryo occurs in the basal cell, so that the cell-walls of the pro-embryo are formed in basipetal succession. Ndérner separates the grass embryos into three groups based on the direction taken by the first two walls. His ‘‘ Type I’’ may be represented by figure 31, in which the first wall is at right angles to the long axis of the embryo, and the second parallel to it. Figure 32 represents ‘‘ Type II’ of Nérner in which the first wall is at right angles to the long axis of the embryo, but the second wall cuts this at an acute angle. Such embryos as that shown in figure 30 fall under ‘‘ Type III’’. In this embryo the first wall is at an acute angle with the long axis of the embryo, and the second strikes the first at a sharp angle. As the figures indicate, the three types given by Norner for all grasses studied by him are present in Avena fatua. Of these types the first is perhaps the most common; when the others are met in old embryos, the cell formation is so confusing that it practically defies solution. The third cell-wall is formed in the distal cell of the pro- embryo, bisecting it parallel to the long axis of the pro- embryo, and is the first cell division belonging to the embryo proper (fig. 30). The fourth division is a transverse wall formed in the basal proembryo cell and cuts off the sus- pensor. No subsequent division was observed in the suspensor cell. At this stage in the development of the embryo there are four segments and five cells. There are two distal cells in the first segment, and between them and the suspensor two superimposed cells, of which the second belonged to the proembryo, and the third was derived from Bot.—Vor. I.] CANNON—AVENA FATUA. 345 the basal proembryo cell, and is therefore not homologous to the cells above. For the sake of clearness in tracing its history in the developing embryo this last is designated as segment three. The growth of the embryo is such as to obliterate these primary divisions, but a comparative study shows that prob- ably the cotyledon and stem-apex are derived from the first segment, the hypocotyl and the initials which give rise to the root-cap, the cortex, and the dermatogen of the root from the second, and the coleorhiza from segment III. This is practically the conclusion of Nérner (1. c). As the boundary between the first and second segments par- ticularly, in the advanced embryo, however, is not at all clearly defined, it was not possible for me to determine whether the stem-apex arose from the first segment, as in Lz/@a (Campbell) and others, or from the second, as in Alisma (Schaffner, 1897). In connection with the origin of the members of the embryo it is interesting to note that the position of the root and of the stem-apex in the embryo are the same with relation to the vertical as in the mature plant (which of course is the case in anatropous ovules in general). The reaction of the embryo to the influence of gravitation can here be traced to the one-celled stage. Experiments might be devised which would show not only the immediate influence of gravity on the embryo, but also which would give some grounds on which to base a plausible hypothesis to account for the anatropous condition of the ovule in this and in other plants where this condition occurs, and also its relation to the embryo. Continuing the history of the development of the embryo, the next wall to be formed after the suspensor is cut off, the fifth wall in the embryo, is a vertical one in the second segment. Up to this time in the growth of the embryo the succession in cell formation is quite regular, but after this it varies considerably. The next wall to be formed is generally a vertical one in segment III, making a seven- celled embryo (fig. 33). This segment does not, however, (2) April 17, 1900. 346 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. always divide before quadrants are formed in the first seg- ment. Quadrants are formed in segment I first, but there is no regularity in the quadrant divisions of the other pri- mary segments. Insome embryos quadrants are formed with regularity in all the primary divisions of the embryo, begin- ning with the distal primary segment; butin others, as in the embryo from which figure 35 was drawn, segment III has divided only once, although quadrants had been formed in the second segment, and in the first, octants. Segments I and II show greater regularity in cell formation than the third segment in the early as well as in the later stages in the growth of the embryo. Octants are formed in the distal segment by anticlinal walls which strike the quadrants at an acute angle, gener- ally below the middle, and not as stated by Norner, at right angles to them. Of the octants thus formed the four upper- most form part of the dermatogen and divide subsequently by anticlinal walls only. The octants are usually formed as figure 35 indicates, while segment II is yet in the quad- rant stage. This is, however, not at all uniform, and peri- clinal walls are sometimes met in the second segment before the octants are completely formed (fig. 36). As just stated, the first periclinal divisions occur in the second segment. These are followed by similar division walls in the end segment. Segment III, in which the divi- sions are very irregular, does not take part in the periclinal divisions above mentioned. ‘The plerome is cut off as in Lilea (Campbell, 1898) in the two end segments at about the same time. This was found also in the grasses by Norner (1. c.). In some cases in Avena it occurs, at least, in the end segment first (fig. 38). An examination of the cells of the young embryo shows an increase in number and size of the vacuoles in the micro- pylar end. This peculiarity, together with the irregularity of cell divisions, serves in a way to mark the descendants of the third segment. The suspensor, which is formed when segment III is cut off, remains small, does not divide, and plays an incon- spicuous part in the history of the embryo. A large vacuole Bot.—Votz. I.] CANNON—AVENA FATUA. 347 appears early in that part of the suspensor which touches the micropylar end of the embryo-sac. The protoplasm in the distal end of this cell (i. e. of the suspensor cell) rounds itself off, a membrane separates it from the large vacuole, and thus the embryo is attached to the micropyle by the proximal ends of the side walls of the old basal cell. The dotted lines in figures 33 and 34 indicate this attachment. The embryo remains connected with the micropyle in this manner until about the time when the primary tissues are differentiated, and then the attachment is broken and it lies free in the embryo-sac. The suspensor cell persists to form the end of the coleorhiza. It was not seen to divide, but it is not unlikely that in old embryos it may undergo a few irregular divisions. The behavior of the suspensor resembles that of SZar- ganium (Campbell, 1899), but not of Lz/@a (Campbell, 1898) and Alisma (Schaffner, 1896), in which the suspensor becomes a large and important absorbent organ. The degeneration of the suspensor of Avena is doubtless associ- ated with the development of the endosperm cells, which closely invest the embryo, as well as with the prominent synergidz which here, as in the wheat (Koernicke), serve to nourish the embryo in its youngest stages. After the primary tissues are cut off, the embryo increases in size, mainly in length, by growth which is almost wholly confined to segments I and III. The embryo is club- shaped, pointed at the micropylar and blunt at the distal end. The periblem and plerome cells undergo few longi- tudinal divisions, dividing mostly by transverse walls. This makes the boundaries of the primitive segments difficult to trace. Cross-sections through the middle of the embryo show a plerome of about eight cells surrounded by a peri- blem two or three cells thick. The two are very similar, being composed of relatively large cells. The dermatogen is made up of regular and comparatively small cells. A cross-section near the micropylar end, through segment III, shows a greater irregularity in the structure of the dermat- ogen and internal tissues, which are not differentiated into plerome and periblem. 348 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. After the primary tissues are differentiated there are three very well-marked periods of growth, in which different sections of the embryo take part. The first is that succeed- ing the differentiation of the primary tissues and extends to the appearance of the stem-apex as a rounded elevation of the epidermis on the posterior surface of the embryo, and includes mainly the descendants of segments I and III. The second period is the elongation of that portion of the embryo which is above the stem-apex, the cotyledon, which takes place soon after the appearance of the plumule and plumule-sheath. The third occurs in the region of the radicle, in the descendants of the second segment of the proembryo, and takes place after the cotyledon has elonga- ted. After these periods of growth and before the matura- tion of the embryo, plumule, cotyledon (scutellum), and root develop and grow simultaneously, retaining the same proportions. These periods of growth correspond very well with the further differentiation of the tissues of the embryo. When the stem-apex is first discernible (fig. 41), the embryo is already relatively large, and all traces of its primitive segments are quite obliterated. The embryo at this period is oval with an obtuse cotyledonary and a sharp tapering radicle end. In cross-section, the embryo has changed from circular to oval. The stem-apex appears on the posterior side of the embryo about one-third the dis- tance from the cotyledonary end, and becomes surrounded by a low ridge of tissue. From this, on the side of the stem- apex opposite, springs the epiblast, and on the cotyledonary side merely an augmentation of the boundary which sepa- rates cotyledon and plumule (fig. 41). An examination of the tissues at this stage of the development of the embryo shows several rows of elongated plerome cells in the cotyle- don enclosed by a periblem of large and rather irregular cells. The epidermis of the cotyledon is beginning to assume the palisade-like structure it later develops as an absorbing organ. The elongation of the plerome cells is continued to the region of the stem-apex, but is not found inthe root. This differentiation of the plerome is connected Bot.—VOL. I.] CANNON—AVENA FATUA. 349 with the sudden shooting up of the cotyledon, and doubt- less these elongated cells function as food carriers from the cotyledon to the developing plumule. "The differentiation of corresponding tissues in the root occurs at a much later period, but the primary root tissues are already well defined. The group of cells at the end of the plerome cylinder, which give rise to the root-cap and to the dermatogen and cortex of the root, can be easily distinguished. The ple- rome initials are not developed until later; they do not, in fact, appear until one or two of the leaves of the plumule are cut off, and the embryo is approaching maturity. The plerome of the root is continuous with that of the cotyledon and that of the stem; the cortex and dermatogen of the root are continuous with the periblem of the embryo. A cross-section of the embryo through the region of the root shows a rather large central plerome-complex, as yet undifferentiated, surrounded by a periblem two or three cells in thickness, the outermost layer of which is begin- ning to take on the characters of epidermis. Surrounding all of these are one or two rows of cells which are the sub- epidermal cells of the embryo, enclosed in the dermatogen of the embryo. A somewhat older embryo shows the third stage of growth, that is, the elongation of that part of the embryo within which is the root. This exhibits at the same time a further differentiation of the tissues of the root. A greater number of cells than before has now been cut off from the initials which give rise to the root-cap (by which this organ is very well marked) and to the extra-plerome root tissues. The plerome initials also have become well defined. The tissues of the radicle correspond to those of the root- tip of the mature plant, and agree closely with those of Hordeum figured by Strasburger (1898). The region below the root-cap is made of large, irregular cells, with little protoplasmic contents. These form the coleorhiza of the mature embryo. After the epiblast has been cut off, there appears on the cotyledonary side of the plumule, and just within the ridge of tissues spoken of above as surrounding the stem-apex 350 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. (p. 348), an outgrowth which partly and later entirely encir- cles it. This is the plumule-sheath. Subsequently a leaf, the first one, appears on the side of the stem-apex opposite the cotyledon, which agrees with the position of the first leaf in Hordeum (Lermer and Holzner, 1888). The plumule and first leaf thus appear precisely as the leaves in the mature plant, alternating on the stem. The homologies of the scutellum, epiblast, and plumule- sheath have been abundantly discussed by many writers (see Introduction), and will only be touched upon in this paper. The scutellum is generally regarded as the cotyle- don. Some authors (1) believe the epiblast to be part of the cotyledon, (2) others that it is simply a projection of the nature of a trichome from the hypocotyl, and (3) still others as a second rudimentary cotyledon. The plumule-sheath has been explained in various ways: (1) it is looked upon as an outgrowth of the hypocotyledonary internode, (2) it is regarded as a third leaf alternating with the epiblast, and (3) finally as a ligule-like growth proceed- ing from the scutellum. My observations on the embryo of Avena lead me to agree with Kennedy (I. c.) in his inter- pretation of the homologies of the epiblast and of the plumule-sheath. That is, the epiblast can probably be con- sidered to be a second cotyledon, although it is not homol- ogous to the true cotyledon, and the plumule-sheath may be regarded to be a ligule-like growth proceeding from the base of the cotyledon, forming an integral part of it, and to be homologous to the ligule of the grass blade. While the plumule and plumule-sheath are forming and about the time of the growth in the region of the radicle (referred to as the third period of growth), the cotyledon begins to increase in length and in breadth to form the scutellum of the mature embryo. An examination of the epidermis on the side of the cotyledon next to the endo- sperm shows that the palisade-like cells, which are charac- teristic of the dermatogen of the scutellum in grasses, are already formed. These cells in Hordewm (Lermer and Holzner, 1888) exude an enzyme which dissolves the starch and proteids of the endosperm and renders them Bot.—Vot. I.] CANNON—AVENA FATUA. 351 capable of being absorbed by the embryo in its germination. My attention has been called to the great similarity in appearance and in function between the epithelial cells of the scutellum of grasses and the cells at the tips of haustoria in certain phanerogamic parasites. The likeness between them and the epidermal cells in the haustorium of the Yel- low Dodder is striking (Peirce, 1893). Both are epidermal cells modified to extract food from foreign tissues, and both exude solvents which can dissolve carbohydrates and pro- teids. A further and more careful comparison would be of importance, and might lead to some definite knowledge as to the reason of the peculiar parasitic habit assumed by the cotyledon of grasses. As the region below the stem-apex increases in length, the epiblast, which was at first connected intimately with the plumule-sheath, becomes more and more removed from it toward the radicle end of the embryo. The epiblast increases somewhat in size although it is never as conspicu- ous an organ in Avena as in many other grass embryos. The epiblast is composed wholly of large parenchymatous cells and has no traces of conducting tissue. VII. Summary. The principal points in this paper may be summarized as follows :— I. (a). The spikelet branches sympodially, the flower and finally the ovary terminating the parent axis. (6). The lodiculz and the upper palet originate in the same rudiment. (c). The awn of the lower glume appears before its lamina, the latter being an outgrowth from it. Thus the origin of the parts of the lower glume resemble that of the blade and ligule of the vegetative leaf. (d). There is but a single carpel.. This originates on the anterior side of the floral apex, which it encircles and finally encloses. 352 Ly; III. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. (e). (a). - (b). (c). (@). (@). (d). _(¢). (d). (¢). According to the hypothesis put forward in this paper, attempting to explain the meaning and homologies of the lodiculz in the flower of Avena, they are to be considered as homologous to the stipular lodiculz of the rye. Perianthal lodicule are not present in the flower of the oat. The earlier growth of the stamen is mainly intercalary in the anther; immediately be- fore the pollen is shed the filament elon- gates and causes the anther to protrude from the flower. The archesporium is composed of a single row of cells originating in the periblem. Each spore mother-cell reaches to the tapetum; none being bounded only by its fellows. In the young anther three concentric rows of cells intervene between the loculus and the epidermis. Of these the tapetum and the middle series are resorbed, and the endo- thecium persists to form with the dermat- ogen the anther wall. In the mature pollen-spore the generative nucleus has already divided. The archesporium of the embryo-sac is of hypodermal origin. The sister cells (potential macrospores) are variously formed. The nucleus of the archesporial cell divides to form four sister nuclei. These may be separated as soon as formed by cell-walls, or (2) they may some- what later be separated by the subsequent formation of walls, or (3) they may never become separated, walls never forming. The lowest sister cell becomes the macrospore. There are no tapetal cells in Avena. The antipodals multiply before fertilization of the egg to the number of 36 or more, and disappear as the endosperm develops. Bot.— VOL. I.] IV.-(a@). (6). (c). (@). (0). (d). (e). (Y)- ad CANNON—AVENA FATUA. 353 The synergide serve to nourish the young embryo. The lumen of the embryo-sac begins to fill first in the region of the embryo. All but one or two layers of cells of the nucellus are resorbed by the developing embryo-sac. The outer integument of the ovule is de- stroyed; the inner persists as the seed-coat. The first two cell-walls (which mark off the proembryo) in the development of the embryo appear in basipetal succession, and show the three ‘‘ Types’”’ of Nérner. The cotyledon and the stem-apex are derived from the distal segment of the proembryo; the root, the root-cap and the periblem initials of the root from the middle seg- ment; and the coleorhiza from the basal segment. The suspensor is separated from the embryo by the fourth wall; it does not subsequently divide. The primary tissues are cut off in the first and second segments only; the divisions of the third segment are very irregular. The organs of the embryo originate in the distichous manner characteristic of the vegetative leaves of grasses. The scutellum appears to be analogous to the haustoria of certain phanerogamic parasites. In conclusion I desire to express my gratitude to the ofh- cers of the botanical department, to Dr. D. H. Campbell in particular, for helpful advice and encouragement to me while pursuing the study of Avena. I ought to add also that the work was nearly but not quite finished prior to Dr. Campbell’s departure for Europe in June. STANFORD UNIVERSITY, August 31, 1899. 354 1892. 1892. 1897. 1898. 1899. 1899a. 1883. 1820. 1897. 1897. 1875. 1880. 1788. 1881. 1887. 1899. 1896. 1888. 1881. 1893. 1898. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. BIBLIOGRAPHY. BENTHAM, GEORGE. 4ritish Flora. ath. Ed. 1 Bruns, Dr. Ericu. Der Grasembryo. flora, 1892. CampBELL, D. H. A Morphological Study of Naias and Zannichellia. Proc. Cal. Acad. Sci., 3d Ser., (Bot.) Vol. I, No. 1. Development of Lilzea. Ann. of Bot., Vol. XII, pp. 1-28. March, 1898. Notes on the structure of the embryo-sac in Sparganium and Lysichiton. ot. Gaz., Vol. XXVII, March, 1899. Studies on the Flower and Embryo of Sparganium. Proc. Cal. Acad. Sci., 34 Ser., (Bot.) Vol. I, No. 9. DE CANDOLLE, A. L’Origine des Plantes Cultivées. 1883. Cassini, H. L’Analyse de l’embryon de Graminez. /ourn. de Physique, 1820. T. XCI. CELAKOVSKY, L. J. Ueber die Homologien des Grasembryos. Bot. Zeit., Bd. LV, pp. 141-174. CHAMBERLAIN, CHARLES J. Contribution to the Life History of Lilium Philadelphicum. 2. The Pollen Grain. Sot. Gaz., Vol. XXIII, August, 1897. EIcHLer, A. W. Bliithendiagramme. Leipzig, 1875. FiscHErR, A. Zur Kenntniss der Embryosacentwicklung einiger Angio- spermen. /enaische Zeitschrift, Bd. XIV, p. 90, 1880. GARTNER, J. De fructibus et seminibus Plantarum, 1788. HAcKEL, E. Untersuchungen tiber die Lodiculz der Graser. Eugler’s Bot. Jahr., Bd. 1. p. 336. . Graminee. Engler and Prantl’s Nat. Phanzenf. Theil II, Abth. 2, 1887. KENNEDY, P. BEVERIDGE. The Structure of the Caryopsis of Grasses with Reference to their Morphology and ee Bul ORS: Dept. of Agric., No. 19, 1899. KOERNICKE, Max. Untersuchungen tiber die Entstehung und Ent- wicklung der Sexualorgane von Triticum. Verhandl. da. naturhts- tor. Vereins d. preuss. Rheinlande, etc., Jahr. LIII, p. 149, 1896. LERMER UND Horzner. Beitriige zur Kenntniss der Gerste. Mun- ich, 1888. NORNER, CARL. Beitrage zur Embryoentwicklung der Gramineen. Flora, 1881. No. 16. PEIRCE, GEORGE J. On the Structure of the Haustoria of Some Phanerogamic Parasites. Anz. of Bot., Sept. 1893. Row.Leg, W. W. The Morphological Significance of the Lodicules of Grasses. Sot. Gaz., Vol. XXV, March, 1898. Bor.—Vot. I.] CANNON—AVENA FATUA. 355 1897. SCHAFFNER, J. H. Contribution to the Life History of Sagittaria variabilis. Bot. Gaz., Vol. XXIII, April, 1897. The Development of the Stamens of Typha latifolia. Bot. Gaz., Vol. XXIV, August, 1897. 1898. SmirH, W. R. A Contribution to the Life History of the Pontederia- cee. Sot. Gaz., Vol. XXV, May, 1808. 1884. STRASBURGER, EpuARD. Neue Untersuchungen iiber den Befrucht- ungsvorgang bei den Phanerogamen. Jena, 1884. 1898. Lehrbuch der Botanik. Third edition. Jena, 1898. 1895. VINES, S. H. Students’ Text Book of Botany. 1880. Watson, S. Botany of California, Vol. II. Geol. Sur. Calif., 1880. 1897. 356 CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE XLIX. (Reduced about one-third). Fig. 1. Longitudinal section of a young spikelet: X 57; 4, lower glume; c, carpel; 9, young ovary; 6, young stamen. Fig. 2. Section of older spikelet to show the beginning of the lamina of the lower glume: X 57; Z, lamina; other lettering as in fig. 1. Fig. 3. Median longitudinal section of the primordium ofa spikelet: X 320; d, dermatogen; pv, periblem; A/, plerome. Fig. 4. A similar section, not quite median, showing an older floral rudiment: X 320; st, floral.apex; /g, lower glume. Fig. 5. Longitudinal section of apex of flower in which the upper palet (wp) and lodiculz (/o) appear: X 320. Fig. 6. Longitudinal section of floral apex: X 320; lettering as above. Fig. 6a. Longitudinal section of young ovule with carpel (c): 320. Saal [Cannon] Plate XLIX. RR NEL ate smapeewae aie ss DXPATES OR Vie Proc-anAcan. Ser.32 Ser. Bor. Vol. A FATUA L. AVEN ‘eae 0 Se Vek tb iat ‘, 358 Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. Fig. CALIFORNIA ACADEMY OF SCIENCES. [PRoc. 3D SER. EXPLANATION OF PLATE L. (Reduced about one-fourth). Longitudinal section, not median, of young flower showing a lodic- ula, and stamen: X 320; lettered as above. Cross-section of a young stamen: X 320. Cross-section of a somewhat older stamen than that represented in the preceding figure: X 320. Cross-section of stamen showing pollen mother-cells (fm): X 320. Longitudinal section of stamen of the age shown in the preceding figure: X 320. Section of pollen grain showing one vegetative and two generative nuclei: X 320. Longitudinal section of ovule: X 320; 0, outer integument; 77, inner integument; @, archesporial cell. An ovule with nucellus (zz); and archesporial nucleus that has divided once (forming two sister nuclei): X 320. Four sister nuclei: XX 320. Proc. CanAcap. Ser.37 Ser. Bor. Von. [Cannon] Prats L. 60 wee Amblyopappus pusillus.... . 115, /Noshosohbyof: 1a (ONCE OC mCHOEC ICE eGs critic PALTINSISTC Kal oe siitea sie) (a, veils tente | tars 110, IvCOpSOIdesin «05 to <5. lee 110; SEACLCOSAl si yc) oltre eo eelienien= maritima..... 110, st.nicolai..... 109, PAVESI OTE Doar Muss tirel slay aes su) 's. een eile] te Anogra pallidaengelmanni. .... Aphanisma blitoides. . .. 96, Apiastrum angustifolium.. .. 106, Aguile giay << so) =) 6 - sie (ES) HHS NAR Ut toto eon ac 76, Gatleyce sts ston OPEN OT sacs) tlhe sede vata) ates) on = CCAICALALA Ts: cs ee cuiad stl ater 3) felts 76, micrantha: ye te eie) «i ier tee MaAnCOSANIA . «2 0 «= «os stellata... «9... = eure vulgaris. .... toh mre PATA COCE sete boule stich fe) toys, Nx Ne! eelsel Arbutusmenziesii!, . 0. {205615 s Arctostaphylos. . . . 81, 125, 252, 255, Canescens 25.55 5 5 «= 84, glandulosa........ 82, 85, PIANCMS ie ious) ees us) ese! fespains VO GIZEGH ir. yar ch tatsye prs, (ora veil Velde 84, MT OMEATM As is oust s) tells els 83, 85, reitpeshas (0leh@ley SG Oe 5) Hace - 82, pringlei.. . sah eianiebee <2 TOMENCOSA |) cee ees she BOs PAE ECMAISIS at oteAvey ey 'ah onte. site 114, 115, CALILOTINIGAN sp eneiiai velit) ta) styled eure) © [365] Asperococcus SINUOSUS ». 1.41 + sess Astracalts arrecttsi.) 1.5.) <)lsie) aee didymocarpus........ 102, MOVIN cari aa Cathrotctct eho htc traskkice ky ca scene 102, Atriplex californicum | 5)... /. 96, decumbens 4 ss) che ote 96, ANIAIDe REA i Msraclen chlo ree eeate tae Avena 331, 334, 335, 337, 338, 341, 346, 350, 351, 352, 353 bagbatar. ys fatua Azolla 6.) xg 10%, ‘ola 16 te) ve 6) ema! ie) amie BACCHARIS CONSANGUINEA . 117, 118, pilularis Baeria gracilis Balsamia alba we; Jou lohl mt Sate. te) sel et ke willie te) where ey te.) Mole |e) Ae tie eid ie NISTCUNS) wee ote oe aie Dlatyspotar.i< een kei polysperma Vit SEIS oy ctcta sb ota leaker ts Bidens gracilis BigelOvia oi.) lam OxiyCes). ite o) aiteio uel lngietaatts PMOTECEL So «oss ole ees eaenioee WATICP ALE. 52 V.-uen oli ioe olen Eleocharis PALUSERISIT. waves us Pron 5 ep Eencoeltacese i. (a feie!cel one one cee Endarachie. chs. cise ci eliceok renee 159, jexbefulitsherbt ain nary my Wo ig) otha oS es Endogone..... etree ails MATAEA LS Tec) chic elena tr keene MIACTOCALPA: so bc. of lal ie) oh sie iota MIAILEOLA. . Aen ast enees microcarpa . PISt/OPINIS 6 o.in atts. ‘havens he aatae te Equisetum 41, 42, 170, 176, 178, 179, 195, 196, 197 ee San tee ©) eG ke eke rene Erigeron canadensis........ EPOGichy On): ticles eu ot ke seen Galifornicwi). = (3) a) clve.ssies ee crassifolium ... .. «|. 129,130, SlutinoswMi es. val eel ol user 81, GIVER) cule vai eeeuye ene 130, [PRoc. 3D SER. 119 276 276 155 169 154 55 171 118 118 119 147 149 149 150 154 154 155 152 150 149 156 151 151 153 153 152 152 155 148 154 154 153 155 153 161 338 261 261 261 95 118 158 162 162 279 280 279 280 279 279 191 89 129 129 131 129 131 Bot.—VOL. I.] tomentosum.. - 129, 130, PPASEADE esse sie cei 3 ene ee Erodium cicutarium......-- 104, HEySiphe ss c0s © she) ere ee! Bess Eucalychortus. Eucalyptus globulus Sarg Gai tie Hey pe en COLDS Aes ots, 4) ieueh seis) ial beens Paliewied.ctc 5 a bel erste) © Meloy rieyel’s Franseria bipinnatifida.. . 116, 117; dubia Se ieveqes « 117, chamissonis... bipinnatisecta.......- VISCIdalc ener tee 117, ieitallae tay £y.'6 oe ay wi aie etvatts Maas logis ICUS Ie owt asta) sie EVATIESCENS is \Vareac s+ > Jascia harveyanus....+.+-+-+-. 151, arenaria.... bombycina. » ++ ++ eee tees COMMMACEAL ears ies st hispidula....... Selroaterg Go ecutmomcec, Opcesoin verrucosa... ... - GeEopoTay yal rley she 2 4! ovens 2) t= Ae brunneola.....-.---:+-:-> COOPER. oy. = 2 = saves dementia magnata ...-.-.----:--; 270; mesenterica... GeEraniacedicn is) shinies) <2) sel oye ss Geranium...... ‘ Giliavmulticaulis’s 3 405 <0. ss nevinii..... Gladiolus. ves) Tevakis Glomus macrocarpus... ++ ++ 22 miuicrocarpus.. - Graminec)..s ice a 2 = 8 © 0 = = = 945 Grindelia.... Eatorhipie =). < sess) s = «css 159, winstonii.. . VEG Eta. oie wera ile sire ley estamos: Vel sacs Heliotropium curassavicum . MELTED OLUS. cv ic ye ye. octal ey mt els ase HMemerocallis).5 esses ose sl ee Hemitomes pumilum ...... Hemizonia streetsii....... 116, LOS PELIOS Fo) rome d/#. lay ois) avin) -e)) w)sigel one AuUStrale |S ve Se span lretel talon tales 979 cinerettm.....--.++--- Clathroidesie.. «ey syn easiiemelns cs rarclerbes iG) aa peo clo, a ben be < membranaceum.......-.-. ae nephriticum.......-.--:-. occidentale. ........--. ~~ phiftipeid. 625 Sor 161 Fyyolepesoiaimiholign, yp semiec do ae 6 171 119 171 IRID4A LAMINARIOIDES......-. 171 NEES Lies ee etled alten te et tee . 78; 80 douglasiana.......-+.-. 78, 120 PuULayL sis oa Sahay atte aks 171 TSOGEES sce) lal) aris etd pate ne (6 ee SO; DBs 265 Isopyrum... 1... +e ahamente fore 56 159 KRYNITZKIA m@rilimia.. 1+ +2 0s 159 LOYYEVANA vs os ee veer vee 350 118 TAMINARIA ANDERSONII ...... 104 Warik cwicyisj.e ee abet 171, 178, 195, 196, 103 Weathresialdcrrs setts) sh dination el so tet tats 119 Leguminose.. ..++-++- rn 40 164 246 248 368 Lepidium bipinnatifidium. . . . 98, 119 THEIVPICSIL :, Sasha pac 8 fos ede 98, 119 WitidUM.- 6 = = beet eat OSE LLo Leptosyne gigantea. . . 96, 115, 116, 120 Lessonia littoralis....++-++:+-> 155 Leucophleps...--.-- ale: ee OO: Per Tibi Fe aque ihe reat ae ie? |e 258 Cifrinta, Jie ep foe ie) enone eves 259 FOVEOLATAY Coke) tne peer ane 258 magnata...-.--+-++e> 257, 258 Odoratay ss )20.75 bs \s este 258 Libocedrus....--+-++s+e-: «+ 262, 273 decurrens.....-- oe aay Pye, Eileeas cesses s 2s 296, 314, 345, 346, 347 VA aCe re ss Geni oo (ps, Fee 55, 95 Weis deh: c/ish eee et el to 112, 120 SARALWIS LS. a'r, cot ahs) 2) ee pone! oe 113 Malwa borealis. ..-. + «+++ 104, 119 PuUsillar se sf. Heueeae 104, 119 Wreareiliasty softs (eck ee ate ea eeel © 10 WEey CATS cee oe ttav al a daetou colts Ah sete 138 Medicago denticulata... ~ -102, 119 SACU nee = eb wha mde eee Ok yee, Melanogaster.. .. 2.222 sess 259 BUTEUS feo seer ee Ses ANE 1st] AQHISSININS 2220s, o, aihete tenet) eters 260 eisenid ei. oc wha teaee ene 259 sarcomelaS. 1. + 2 ese ee 260 ta DertLOLmis:. cy. «> soe PL is 260 variegatus .....-s+-+++es 259 bY, (a ble: Wee area ae - 338, 340, 341 Mrelobesia. 235. sred cot cer heney elmer st ia 158 Mesembryanthemum crystallinum. 105 119 MOGIBOLMM s60% ete erie 105, 119 Microseris linearifolia. ..... 114, 120 Microthecium .....--s.s-+s%-e2s-s 281 SORCLER ee Se Te al Ia eS 281 Montia perfoliata .......+-+-s 79 TOSHIATAL - ley hie pees tetas 79 SAROSA) 5 iis pot ee hes hw Lee opis, ans 79 Muhlenbergia pumgens ......-. 75 Myrmecocystis .....-++-- 269 Candidac iets Ses chap ret siee 269 cerebriformis........ 269, 270 NATIADACE . 2... +: - Seaine Cate 36 CALIFORNIA ACADEMY OF SCIENCES. Naias 4, 5, 6, 7, 10, 12, 26, 28, 29, 30, 35, 36, 37, 38, 39, 41, 42, 43, 46, 48, 49, 50, 51, 52, 54, 56, 57; 58, 309, 312, 314 ancistrocarpas” sis, (4 > us hte 22, flexilis ..5, 6, 7, 9, 10, 11, 16, 17, 23, 24, 25, 27, 39, 41, 52, 53, 54, graminea J. ".ic. 2 sation 9, nha b (or: Me eI Bar ec toe rt ou 7 major 7, 9, 11, 16, 22, 23, 25, 31, 55, TUAFEL «te 8 ete ale el a ferles ene podostemion .. 7% i <5 = slew tenwifOlia).. 45. elas s cee 16, Fittishitehcte a a OaeOe cece C6 Nereocystis liitkeanus. ..... 148, Newberrya ecis, i= \eyisue |e eueaatle subterranea . i250 % © so WH PHL sis fo! se oie sie of anya Nyctapinacers: i” so Js «>: cules meet = Nymphea.. COM Pacts ss) oss sa) somes onelee monticola, 263.346 ais stele 6) ie) wits One SOFLOUCLAS o\ 5. «on srs wey Sek eee BOCLAIIG ise pa nor oot a ea leone stephensil) 9) 25;-, Macs reer Cénothera albicaulis....... 73; brevifolia’: . 0.0 st. ehey ete gypsophila. 2.0. Sak cheiranthifolia .. <= <<). » 1045 SPULALIS ss cn el wy oh wee Opie ee trichocal yx s <) io<)/2 a Pan er Poe EUpICwIA ss), sto) bss eee Pilifoliay 5 6 lawas teeter ete viridescensS.. ..-..- Oligomeris glaucescens. ... .99, Sub ara se Sedan Tre cat neta Soy ONAgTACEE 2) 6) sn is ee a Soe = lone COOPUGATIAs, \erielro ie torts ems te 94, 95, 105, engelmanni........ -.94, Wittoralis, © © 2 ve) ses 105, prolifera .9. 2. 2.44 5's 106, Orthocarpus 2.) 5 6) i oo omian purpurascens... ..-+-++-- 112, OEY Zaid: ehsl ool oy jas ote. anos PACHYPHLG:US..»-+2+ «sees CANES. <5 sw) 6. Ss @ ee) or conglomeratus.. . ligericus . ....+ Ast sebeois Pandanacec (lei ele site. -), «hie ere te [PRoc. 3D SER. 31 47 59 23 22 55 ‘ Bot.—VoL. I.] Bandanuss)..0.5.°. Papaveracez....... Parietaria debilis......... 95, PPARSIROLA yo eee eat ie le len iat sitet 196, COBEULEA ae Wihie) 6, se ehsite Seley a IPECIGCATYOeut S's «felis, ele S68 swe GHUEHSIS foe sie shies ehen eis AMBERTIS Fes eigay oy bo eet = 108, Pelvetia fastigiata...... Petrospongium berkleyi...... Peucedanum insulare Phalaris canariensis. ...... 94, HOC Se rete lielia Jue) ain mee oh ed tee ee Phycocelis iewenser Gs EIS By ae Wir o re. ceca ter reptans.. tnt: SmI TTEISte ae tS aida ken ol She 159, 161, PASC Tae Is oh rho ay errereaiie Pans 161, PHyllospagixs.0s ssn «3 « Pickerinpia montana (sj. ss)... = TEE SONTAL te oye ets sania ete) a BULVEOLA ER a cpio ali etien sir el siento SCADEOSA sso) 5 es so ee PTGS cate ell sito OHLOE EAI a ie he) (eis s) fe ney sere pie) te MUTT TES eve: Yael eile) shots terey tits eee PISEIA es aes as fo) MA onaealo hoo Sh SELALIOLES 5 ey Guie sa ue feyedls! cine) site AAPA TAC cialis ie yoasopin Mel eas take Plantago insularis....... 112, PaAcapowulcaye have el alse 6 Rapes ai Platystemon californicus..... 98; Podophyllum..... OlemiOninCerey. 2 \ as) als) shel sire? aide Polygonum aviculare ........ Polypogon monspeliensis ... . 95, BOpHbus fremont. 426 Western sth a hen ELEMUMIOIGES) 5.5 fo, -, settee) (hie sills Potamogeton.... Bit area: aha 4, Potamogetonacee.. . Protococcus.. . Pseudotsuga douglassii. .... 247, Pterygophora californica ... RURELATAUM ot its) ie el OTL OW AE Oy SO Be OL CUORCRIE. Ee DUPIESLOWIG Ca cc sab ath Se Sh ati aniand Os AV AMO TIE fires) Wels <) ia) ois aeunte MIEEOEALISI CS of a as ete s tks hotweniavis ae OIVERCUS.. 92) < Welles © «6 ele siicl c's RAMALINA oe cee 215, 220, 222, 226, Teticulata . 208, 209, 211, 212, 213, 217, 224, 225, 229, 230, 231, 233, 238 INDEX. 321 RESEdACER Ys jc, ret sien eel Petes Nels: io 98 RRIZOpOSon!'s (i aM i Seueteten aeeteteine 118 BUTARHES sia nesA is 1S) hae e 197 Rhodomelavarixs sis sper. snarane 189 RENTS focicar th faln'ds yer, she hat heyitebte) Wel ve We 93 ROQaIIe yas ok ou wien seen ene ay wes 108 COUNLEED acd ie oe tack nese coun 133, 134, 119 trichocalyx..... . . 3133; 134, 151 Rimmex acetosellay ic.) owns teens 152 119 SANICULA MENZIESII.... - 106, 118 Scrophulariacez.... . ~ Bg ote 91 SEY EOSUP MOT sas) foiiesl ot el ol telvonee 159, 147 Diblosws sees Ewe etree ts 148 TORTS EATANS =e pos cio teliad oienewns 148 cotmplanatus ss <5) (s 2 \6 162 Sedum coredont, 5 0-5 eben 162 Pum. soe hey alee eS 161 Selarine lias Tit gee mieh aan 81 Sequoia gigantea... ...... aLieine 275 SEN PCEVIFENS \.)o a) hit ool ls =i 275 SeSlenich ys (yen su cet eam 339, 340, 275 Silene gallica... 40% Ms stems ve 1d 171 Smalacina |5\<, . Sas ye hsweleenpewane 243 Solanacez.. . . 270 Solenache sane oe Romie cee 56 SONCHUS ASPEE <<) sue emons 114, 40 OIETACEUS = (tte ciety cel ote 114, 112 PEMETGUNNIUS Sey yal eke ee 114, 120 CEMULZOTIUS 0/0. fo) ote at La dete 112 Sorantiterats fo yeupueteuet = Valieale 159, 119 ALVOIGES: a) (ots forte latl= Senet one nome 171 Sparvyanacere).. io) «seen! = ee 107 Sparganium.. 45, 49, 55, 57, 293, 295, 89 300, 305, 308, 312, 314, 315, 317, 118 319, 321, 342, 347 72 eurycarpum. 57, 294, 295, 296, 298, Vie 302, 306, 307, 318, 319, 321 36 greenii.. . 294, 295, 296, 298, 299, 36 302, 306, 307, 318, 319, 321 230 longifolium .. . 294, 296, 298, 299, 267 ramosum . . 293, 295, 307, 308, 309, 280 312, 317, 320 267 simplex . . 294, 295, 296, 297, 298, 267 300, 301, 302, 304, 305, 306, 307, 261 309, 312, 315, 317, 319, 320, 321 148 Spergularia macrotheca.. .. . .97, 161 Sphacelaria. . Mes NM Okt Se 2e 159 didichotoma 20.5. 6 ek ee 161 tribuloides), sesso sen name 156 Sphacelariacec, © <)is,s <) 6) st) eee 157 Spbheeriatics. yma ee pee Shc 156 (Hypocrea) setchellii..... ZOPELI sys aye leettell la hie ae or mOBEhid (4 a gine am ele eee 80 Spheeriacessss, 6s) se od tnt ere 127 Sphzrophorus). 2°. . 25 222, 229, 238 globiferus........ 222, 228, 215 Splanchnomyces behrit. . +... + 5s 234 Sporophacars ee sc) ol ae eh eure Cyanea. 2 meee 273 magnatum......-.; a) 6 ol ieue monticolum ........ 271 (Oogaster) caroli......... 274 (Spheerogaster) candidum. .. 274 GCISCHNTT $5 na Chet cee 275 OLVaCeteH.s sss) shee 275 (Spherotuber) californicum.. 274 puberulum). 2.0% 6 aieu «we EV PUA ie) Gave tek oh oh on oae 293, 296, 321 Typhacere co. sss sce 5 es») x ee 293, 321 WE Aconebice ois caital et % leita te co meenrenee 163 SEMMOSA .. s.0 4.0) Raniie ay a, Oye ee Umbellifetce: .'.sica-« © 8 serene 106 Umbellularia californica... .... 81 Wredinee 1) nua sust ee lanes Saree seleks 282 MOLHCACEOR San ne fe) Sale ah eee 95 jeicjcl- A gps al 222, 228, 235, 238 Ustilaginec .) oc: sss 6s me sees 282 Ustilago cyanea. ws e+e eee ene 281 VACCENTOM s/6) oie" ni-e) ols ee ee 246 Ovatum . 2+ -«ces ow ee: =) Blow WerOniCa ss (ois: ale os) soem een notes 125 WIGANDIA CALIFORNICA ..... 129 NA CELPONATEL oo lo ieaneh wiaew san wl ameter aie «125 VAN i Ger et CCRT TAC Re ae 171 Zannichellia 4, 13, 27, 34, 35, 36, 37, 40 41, 42, 48, 49, 50, 52, 53, 54, 55, 57 58, 59, 296, 312 Palustris'., WE ETS BY 3 avg NA eG mah oe pide ale ediecaee Maks ee enka Fests oar 2.00 “« VII (1876), part 1 (volume complete)... 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Nowe ss Some Observations on the Development of the Karyokinetic Spindle in the Pollen-Mother-Cells of Cobeea scandens Cav. BY - ANSTRUTHER A. LAwsSoNn. WitH Four’ PLATEs. Issued November 17, 1898. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1898, wf ; i REA ety hg F it et Poem ae a Sate a eeey y LeATION ze ber eT 2 ee PUB sere Ve sear Ses aes , sae 19.4 mS) Wee OSS a ; - pik \ ic ke) eee! ahs Aste PS EDI ay yo PROCEBEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. THIRD SERIES. Botany. Vorw fo Noss: Some Observations on the Development of the Karyokinetic Spindle in the Pollen-Mother-Cells of Cobzea scandens Cav. BY ANSTRUTHER A. LAWSON. WirH Four PLatEs. Issued November 17, 1898. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1898. fits ih eth ata: PUBLICATIONS OF THE ACADEMY, Ci ontinued. ~“PROCEEDIN GS (osanGy: FIRST SERIES. .) \ > Vol. I (1854-1857), out of print. “TI (4858-1862), out of print. III (1863-1868), parts'1, 2, 3, 4 and 5, out of print. 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By Wathan banks; 2s sf. cakk Ua ieet's a toe Zoology, Vol. I, No. 8—Thé Anatomy of Chelyosoma Productum: Stimpson, By Prank Watts ‘Bancrofts Sin cdi. chee eee Zoology, Vol. I, No:.9—The Gordiacea of Certain American Col- lections, with particular reference to the North American Fauna.—II. By Thomas H. Montgomery, Jr., Ph. D...... Zoology, Vol. I, No. 1o—Observations on Monogenesis in Metridium. By Harry Beal DORR a7 ihe a aceite feb a antec Be Calthm w eed avoiae eo ade Geology, Vol. I, No. ~The Geology, of Santa Catalina Island. By William Sidney Paneier, Sniith baat cae. dae gel e ge Geology, Vol. I,,.No, 2—The ‘Submerged Valleys of the Coast of Cali- fornia, U.S. A., and of Lower. California, Mexico. By ‘ George DAVIS OMe ie oA se Daa antes oe ROE E wala Oe ee eas Geology, Vol. at No. 3—The Development of Glyphioceras and the Phylogeny of the Glyphioceratide. By James Perrin Smith: Geology, Vol. I, No. 4—The Development of Lytoceras and Phyl- loceras. 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Vol. I,, No. 4—Systems of Simple Groups de- rived from the Orthogonal Gioup: By Leonard E. Dickson.. publications should be addressed to J. OB. GUNN, Corresponding Secretary, California: Academy of Sciences, I 35 Mathematical-Physical, Vol. I, No, 1—On Rational Quadratic Trans- 3 ~ * JO 125 All subscriptions, applications for exchanges, and inquiries concerning a San Francisco, California, SAS ay Aetiye #o 5 The Origin of the Karyokinetic pi coerulea Linn. ¥ - “sy Ciara L. Wiuuiams, M. S. , VE a t es pron é ae | roe ii ayy, BN Si vi 7 - WitH Four at te sf ePtat Issued April 15, 1899. w% Pe de ; 4 25 ¥ é cIS Le ODE, PRC ea, + 0s > AY ye oe ") wae tie rman. A, tk ie g la (ut 4 y Th ties: » Chia * x ein fe : ‘ - ne 4 Py es ge ‘ nl EDI A taget . Doueias H a PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. TuIrRD SERIES. Botany. Vow, If Now 6G The Origin of the Karyokinetic Spindle in Passiflora coerulea Linn. EY Ciara L. Wituiams, M. S. WitTH Four PLares. Issued April 15, 1899. SAN FRANCISCO: * PUBLISHED BY THE ACADEMY. 1899. OF ‘THE ACADEMY. Continued. " PROCEEDINGS: ‘(Ocravo)., THIRD SERIES. - f Zoblony; Vol. I, No. 1—Plasmocytes: the Survival of the Cantrosorties and Archoplasm of the Nucleated Erythrocytes, as Free and Independent Elements in the Blood of Batrachoseps atten- Ay kd “3 » -uatus Esch. . By-Gustav Eisen, Ph. D..-... 5... ee eee $1.00, fie on 8 Zoology, Vok I, No. 2—Diemyctylus torosus ; The Litstiictory and” Came h Habits of the Pacific Coast Newt. By William E. Ritter..... . 50 Ms Zoology, Vol. I, No. 3—Scientific Names of Latin and Greek ‘ ey Derivation. By Walter Miller.j. 00. 00.00... Dae case eee ous bagnh ae i Zoology, Vol. I, No. 4—A Genus of Maritime Dolichopodidee New to . America. - By William Morton Wheeler... ©... ...\.0.-- TA. 25 Z oology, Vol. I, No. 5—A Preliminary Account of the Marine Annelids. of the Pacific Coast, with Descriptions of New Species. By Oe.s) # eer bert: Px Johnsom,. 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By: Edwin, Chapin Starksw. 2.2 ena ai 2 aa AG Geology, Vol. I, No. 1—The Geology of Santa Catalina Island. _ By f Sica : William Sidney Tangier Smith........ PAV NE SW emt ARENA 50 - eee: Vol. I, No. 2—The Submerged) Valleys of the Coast. of Cali- fornia, WaSivAsand: of ‘Lower California, Mexico,’ By a George Wavidsann te sic cies RSs hoa vs ea ogni s £5O, 2s Geology, Vol. wf No. 3—The Developmen of Glyphioceras and, the Pi Phylogeny of the Glyphioceratidze. By James PerrinSmith. .35 5 Geology, Vol..I, No. 4—The Development of. Lytoceras_ and Phyl- he loceras. By James Perrin Smith:........... 0+ ose eee e eee 25 M Geology, Nol. I, No. 5—The Tertiary Sea-Urchins of Middle Califor- oy 4 oma, By fohne Gc Merriants iio pun. 38 oes eet ARR Ag Geology, qn I, No. 6—T he Fauna of the Sooke Beds of Vancouver Sage PALULS ep dopa, © aie doje 6, 4 di Slut O48) eee) oO) 4 od wee we eee oe oes ¢ abel vere le Ce 8 0 6) eo wie a dé 0 6 6 byes afeia o “ es ys ‘Alice re aa Ae PW PERNA wed Wola ay pha aeons NY HOE RN Bb tony, Vol. I, No. (ena oe Memoirs. By De ‘Aid Saunders. 75 ‘ etndeds Cav. By Anstruther A. Lawson. eG EAL oN BEOE Bone ‘ = ut siflora coeruila ‘Linn. nt Clara A ai Williams. Se igs eee RCS aN Na PAR RES * egtlettels wed (eiebe 7 mem 8 le 0 87e. Waihomaticat Dhycieal Vol. I, Nas othe: Daerah Geen wat Mee Be Transformation. Py Leonard E, Dickson... +... +.. Fie baie a Ye ae ace y (eat enenee eM) wie Wleye 6 6 enacers se ven Pa acne Spe. AO) Be GUNN, Laing Secretary, ; : California Academy of Sciences, ae San Hepa California, ue wht a “ie HORA RSP a ne eye Ub eS at Ha ip Shee Ae Ne mare y eat PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES. THIRD SERIES. Borany. Vows I Now: The Nature of the Association of Alga and Fungus in Lichens.. BY GEORGE JAMES PEIRCE, Assistant Professor of Plani-Physiology, Leland Stanford Junior Untuersity. WitTH ONE PLATE, Issued June 5, 1899. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1899. ay a TER tg ust I ; ae Rey Aon Vii ae 4 Aa ” PROCEEDINGS CALIFORNIA ACADEMY OF SCIENCES. THIRD SERIES. Botany. Von. f° Nomezee The Nature of the Association of Alga and Fungus in Lichens. BY GEORGE JAMES PEIRCE, Assistant Professor of Plant-Physiology, Leland Stanford Junior University. WitH ONE PLATE. Issued June 5, 1599. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1899. Cae VY “PROCEEDINGS: [OGtAV) Sie fan Ai Nieih\ garth THIRD, SERIES! 2 ¥ i a Zoology, Vol I; ‘No. 1—Plasmocytes; the Survival of the Cash deanies a athe arich: Archoplasm of the Nucleated Erythrocytes, as Free and’: Independent Elements in the Blood of Batrachoseps atten- — ae ‘uatus Esch.) By Gustav Eisen, Phi Deo. isso ee eel eeeeeae : Zoology, Vol..I, No, 2—Diemyctylus torosus ; The Life-History and . eure Habits of the Pacific Coast Newt. By William E. Ritter...2.. 00 ese Zoology, Vol. I, No. 3—Scientific Names of Latin and Greek : Derivation. By Walter Miler ory wi boy cis eed ace 9's Bebe aOR Zoology, Vol. I,,No. 4—A Genus of Maritime Dolichopodidee New. to . ~*" America. By William Morton Wheeler... 02002 05.5.5 6. . Zoology, Vol. I; No. 5—A Preliminary Account of the Marine Atinelids? ae BOE the Pacific Coast, with Descriptions of New Species. By * Herbert P. Johnson, Ph. D...... 2.25. Si devel ‘Zoology, Vol. I, No. 6—Description of-a Species of Fish (ntstsutcaina thas ” owstoni) from Japan, the Type of a Distinct Family of Lam- “noid Sharks. “By David Starr Jordan... 6. eee ete Zooloey, Vol: I, No. \7—Arachnida from Baja California and. Other » Parts of Meco By Nathan Banks yori g Bake ow inee« Zoology; Vol. I, No::8—The Anatomy of Chelyosoma productim Che sk Stimpson. By Frank Watts Bancroft...... Malad Me aca mapa eats ae Zoology, Vol. I, No: 9—The Gordiacea of Certain American Col- ° Mi lections, with particular reference to ‘the North American Bae Fauna.—Il. By. Thomas H, Montgomery, Jr., Ph. 10 agers . Zoology, Vol. I, No. to—Observations on Monogehesis 1 in Metridium. %, sf By Harry Beal Lorre ys A bch age Uys Pi alfaie eke La Zoology, Vol. I, No. 11-~The Osteological Characters of the Genus f Re Sebastolobus. By Edwin Chapin Ptarks\yaist er ehs : ae Zoology, Vol. I, No. 12—Odonata from Tepic, Mexico, with, Supple- mentary Notes on those of Baja California. . By Rae Pes sat OVER tec iosce tant eset Se pet AO cer ee iy steele aaa eae PERS fee Geology, Vol. I, ‘No. 1~—The Geology of ‘Santa Catalina: Island. ‘By eS ites! i William Sidney Tangier Smiths 2: 351% Phe Be Ste ie eta Belo a eee ae 6 Geology, Vol. I, No. 2—The Submerged Valleys. of the Coast of Cait fornia, “U.S. A., and of Lower California, Mexico. By hte lec ah) CON Mea VISOR EGA SOA Soya ana ea samen woe waiiba se Matoma’s ~ Geology, Vol. I, No. 3—The Development of Glyphioceras aad the. ‘Phylogeny of the Glyphioceratide. By James Perrin Smith. —: Geology, Vol, I, No. 4—The Development of Lytoceras and ~Phyl-. © ~~” "loceras. By James: Perrin: Sate hi east ae ae al a Geolozy: Vol. I, No. 5—The Tertiary Sea-Urchins of. Middle Califor- nia. By Ohi GooMerran ice pat. tae CA eee utente i Geology, Vol. I; No. 6—The Fauna ‘of the Sooke Beds of Vancouver 1s era i island... "By Joha Oh Merrianis,2. Botany, Vol. T, No. 2—Studies in the Herbarium atid tia Field: —No. Ais ater: Ree By Alice Eastwood. NN. seal mak Caley ake Coie RS a Liem Peete e wigs aur EAM aed “Botany, Vol. I, No. 3—Studies in ‘the Herbarium and the Field. —No. 2 Sine A ie DS a EN Alice Eastwood.. Ceaveaa Ras eens GOVAN ORLUL arom ryeb a ae aay . Botany, Vol. I, No. 4—Phycological Memoirs. By De Alton Saunders. - Botany, Vol? I, .No. 5—Some Observations on thé Development of the 9... > Karyokinetic Spindle ‘in the Pollen-Mother-Cells of Cobeea: wi . scandens’ Cav. By Anstruther A: Lawson: . 20.02... - Botany, Vol. I, No. 6—The Origin of the Karyokinetic, Spindle i in Pas- * ; ~” siftora. -ceerula: Lirint* By Clara L. Williams. © ov. 32s eo. as. ie es Botany, Vol.I, «No. 7-—The. Nature of ‘the Association of Alga and Niamie ete Fungus in Lichens: By: George James Peirce, ..........-.. Died ite ie aaa Mathematical- -Physical, Vol. I, No. r—On Rational Quadratic Trans- > Lbs oe formations. By M. Wii Flaskell voc Nita hee ND a aay -Mathematical-Physical, Vol. I,. No..2—The Quadratic. ‘Cremona , Ps Transformation. By Leonard E. Dickson... ©. 5.4.4 !0 60 be Gerong toy cee Mathematical: -Physical, ‘Vol. I, No. g—On Curvilinear Asymptotes. a tN WME W Haskell uss 0S aiy os vals niente Sey eae a Sisthcatan Physical Vol. I, No. 4—Systems of Simple Groups: de- ’ Ni ea eet: cate ved fond the Orthogonal Group. By Leonard E. Dickson:. 2s: ©.) 5) All subscriptions, applications for exchanges, and i inquiries: concerning the « publications should be addressed fon: Bea : é wy ‘OE B. GUNN, Corresponding Secretary, California Academy of Sciences, ro ie KO, Californias, . Ai if we 44 A ‘ PROCKEDINGS — CALIFORNIA ACADEMY OF SCIENCES. THIRD SERIES. | Borany. Vou. I, No. 8. Californian Hypogeeous Fungi. BY H. W- HArRKNESs. Wiru Four PLATES. Issued July 8, 1899. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1899. fe AN hd / PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. TuIrRD SERIES. BoTANY. Vons lis Nou ee Californian Hypogeeous Fungi. BY H. W. HARKNEss. WitH Four PLATES. Issued July 8, 1899. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1899. ; Continued. — : - PROCEEDINGS, (Ocravo): es rd = RB cscsoag h SERIES, | ; ee a, rd ee se ey ee a er eas Cr i a ee ee ee ey Vir eee ee ee rhe more ees whee Co ee ee RS Thomas H. Montgomery, Jr., Ph. D...... 125 Zoology, Vol. I, No. 10—Observations on Monogenesis i in Meeiainen . : By Harry Beal Porte: tJ.0/3 wii eece sak ae ok wea ueeeae ‘266 Zoology, Vol. I; No: 11—The Osteological Characters of the Genus Sebastolobus. By Edwin Chapmc Starks, 34444 4) seks mle 125 oe Talay Vol. 1, No. 12—Odonata from Tepic, Mexico, with Supple- mentary Notes on those of Baja California.. By Philip P. Pe LETS cere eno ae OP hr cei ats anos ea Hts ies Sha Na ie ees asd 125 2 2 Geology, Vol. I, No. 1—The Geology of Santa Catalina Island. By “William Sidhey Lanpier Simin, foe rok oes eke owen ves 50 Renlbey, Vol. I, No, 2—The Submerged Valleys of the Coast of Cali- fornia, US. ais and of Lower California, Mexico. By Pee cmete ae George Davirison yr soee Seti. Soho ce he Oe I oa Lx” peas ; Geology, Vol. I, No, 3—The Developinent of Glyphioceras and the ae Phylogeny of the.Glyphioceratide.... By James Perrin Smith. ~ .35 i, Geology, Vol. I, No. 4—The Development of econ and Phyl- “loceras. doy" fames' ‘Perrin Sinith cs 2.3), ak at elem awe Oe one Hier e..: Geology, Vol. I, No. 5—The Tertiary Sea-Urchins of Middle Calton bas ~- nias By OTT, CP WerrAgns Je ats pee su ‘ o Geology, Vol. I, No. 6—The Fauna of the Sooke Beds of Vancouver B95 ca ord lands ..by: John GC. Merriathy 3. Cos a oe te oa ase es - Botany, Vol, I, No. 1—A Morphological Study ey Naias and Zanni- - Gens ~~" chellia. By Douglas Houghton*Campbell ... 22... e000 002, F.00 Paes: Botany, ach I, No. 2—Studies in the Hemaruin andthe Field. No. 1, as By. Alice EASEWOO is ee ty eeu GMS More Oe cae Cou hat ae 25 Botany, Vol. I, No. 3—Studies in the Herbarium and the Field. SING 28 OS By Alito Bastwdodsers, ipo ee Leal) aga 150" af . Botany, Vol. I, No. 4—Phycological Aenions: By De Alton Saunders. ~ .75 _ Botany, Vol. I, No.5—Some Observations on the Development of the hs are Karyokinetic Spindle in the Pollen-Mother-Cells of cai aay _scandens Cav. By Anstruther A.‘Lawson......2...e0.505 135%) : Botany, Vol. I; No. 6—The Origin of the Karyokinetic Spindle i in Pas- Wy ae ao Peter in ner ~~" siflora coerula Linn. By Clara 2.2Williams 5% hea eb ee pear 4. ere eae a Botany, Vol. I, No. 7—The Nature of the Association of Alga and era Rs che es. * Fungus in. Lichens. By George James. Peirce, ..2.....0.... er once NG Botany, Vol. I, No. 8—Californian Hypogzeous Fungi,” By De Wikeeog vas Harkness, Ae O Rie vey STIs Au Veh R aOR OLS Wee UR TE ROR Lak Peet TEE ey Mathematical- -Physical, Voi. I, No. 1—On Rational Quadratic Trans-° ius eet - formations. By M. W, Basket Aad ee ee cee onta, ies Mathemiatical-Physical, Vol. I, No, 2—The Quadratic: Cremona 2 -- ~. . Transformation. By Leonard E, Dickson......s.cssvs-+e Gos I aie _ ’ Mathematical-Physical, Vol..I, No. 3—On Curvilinear ° Asymptotes. cee By Woo dtaekelby oc AN acter a ee OME ae aes _ Mathematical-Physical, Vol. I, No. 4—Systems of Simple Groups de- pe -. rived from the Orthogonal Group... By Leonard E. Dickson... .25 °° | SMALE subscriptions, applications for exchanges, and i airs Long COnErEOe, Oe, Daas : _ publications should be addressed to ‘ais sy Ome GUNN, Corespoadtée Secretary, GP: Bear. California Academy of Seercest: Reena - # ace se aean ‘San Brantyan, Cantona ie 0 $4 Ft Paks. PROCEEDINGS OF THE CALIFORNIA ACADEMY OF. SCIENCES. TutrRpD SERIES. BoTANy. Vou. I, No. Studies on the Flower and Embryo of Sparganium. BY Doucitas HoucutTon CAMPBELL, Professor of Botany, Leland Stanford Junior Universtty. WitrH THREE PLATES. Issued July 21, 1899. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1899. - ws Tha ay Ls rr ae aa | ey ry oy, PUB my it May ‘ / i e | . . ~y f Dy ¥ BAPE TP yl Ay TTEE. “ a ¥ vid ~ WEA cw AY (CW ABA. | ¥ Be ous ga ova ) 5 tb 4% ET: ? CHELL, . eels 4 * Ne Wits ty PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES. THIRD SERIES. Borany. VoL. dt Nooo: Studies on the Flower and Embryo of Sparganium. BY Doueias Houcuton CAMPBELL, Professor of Botany, Leland Stanford Juntor Untuersity. WITH THREE PLATES. Issued July 21, 1899. SAN FRANCISCO: PUBLISHED BY THE ACADEMY. 1899. g Continued. | eee Oe GeeRBince: iBecava: baat? POR TO AMER SSR ERR ESS WR 8.) Be rtad Ver ata OMOEA UeRy 8 og su eres Zoology Vol. I, No. 1—Plasmocytes; the Survival of the Contipstinies Rival Alc and ‘Archoplasm of the Nucleated Erythrocytes, as Free and ~ AE. Rae eal! 7 Independent Elements ‘in the Blood of Batrachoseps atten-— De aa HC ic Mie ated okt uatus Esch.) By Gustav Eisen, Phe Di...) ceric alee etsy $1.00 ae NYAS Zoology, Vol. I, No. 2—Diemyctylus torosus ; The Life History and % PA Habits of the Pacific Coast Newt. By William E. Ritter..... +50. Sea Zoology, Vol. I, No. 3—Scientific Names of Latin and Greek ew MA Derivation. « By, Walter Millers: (sce. ponspieies Vid pliyeke eee Mess Bens Zoology, Vol. I, No. 4—A Genus of Maritime Dolichopodidee New to. «4 America. By William: Morton Wheeler... 22.005... eres +25 +. Zoology, Vol. I, No. 5—A Preliminary Account of the Marine Annelids =~ hone ~~” of the Pacific Coast, with Descriptions of New Species. Bye yi ty er bere Fy. Fobnsenl, Pa Pes Oo. yi cae ti lolein's viet a 5 o Meee gle, am! « 50. Zoology, Vol. I, No. 6—Description of a Species of Fish (Mitsukurina ‘ owstoni) from Japan, the Type of a Distinct Family of Lam- noid Sharks. By David Starr Jordan... 0... ..2..0e.. esses 2B) Zoology, Vol, I, No. 7—Arachnida from Baja California and Other : Parts of Mexico. By Nathan Banks ?))5 sigan bce hx gle / 1.00 “Zoology, Vol. I, No. 8—The Anatomy of Chelyosoma productum Stimpson. By. Frank Watts Bancroft: 3... 02.000. J. ee ene es 150 Zoology, Vol. I,. No. 9—The Gordiacea of Certain American Col- bs) | ee ee lections, with particular reference to the North American MSR Ne ee thy Gules . Fauna.—lII.. By Thomas H. Montgomery, Jr., Ph. D....., 125 Maatsat aed Zoology, Vol. I, No. 1o—Observations on Monogenesis in Metridium. 5 By Harry BORE: TOPEV GUN ia osictkracem hur aibiis oh Abie p pk se OE 25, ’ Zoology, Vol. I, No,, 11—The Osteological Character$ of the Genus ory Sebastolobus. By Edwin Chapin Starks. 20.602 Santi 25 Leoldey, Vol. I, No. 12—Odonata from Tepic, Mexico, with Sibpis. mentary Notes on those of Baja California. By Philip P. ne Dal Vari ny etal ie SPs atalale pt elchiteled deiner Se aig’ CHa won ale 25 Geology, Vol. I, No. 1—The Geology of Santa Catalina ‘Island. By ; William Sidney Tancier Smith i Usb Jae ety tee om erccae .50 Geology, Vol. I, No. 2—The Submerged Valleys of the Coast of Cali- fornia, U. S: A., and of Lower: California, Mexico. By MRCOFTS TIAVIASON S30 FE Gi ae yd sie cece ding Wide wrmgto ew ida Che piney K 50 Geology, Vol. I, No. 3—The Development of Glyphioceras and the rie eee Phylogeny of the Glyphioceratide. By James-Perrin Smith. .35 | Geology, Vol, I, No. 4—The Development. of Lytoceras and Phyl- ya loceras. By James; Perri. Smith ysec 08 2 ue oo alae pen eisne'e 25 Cle NE arabes Geology, Vol. I, No. 5—The Tertiary Sea-Urchins of Middle Califor- Liha Renae. nia. . By JOH CG: Mera. ose Kee, Shag) alee aus 9h 3 My Geology, Vol. I; No, 6—The Fauna of the Sooke Beds of Vancouver ( * 5 sland... “By For Cy Merriany § in ote LON aie wea inka g bee i Botany, Vol. I, No. 1—A Morphological Study of Naias and Zanni- a HA chellia. By Douglas Houghton. Campbell..........--.4.+-: 1.00 Botany, Nel I, No. 2—Studies in the Herbarium andthe Field. —No. 1, ~ yi . By Alice RashWoOd se defo Ofiw wie uo Ve eee Ee S25 ‘Botany, Vol. I, No. 3—Studies in the Herbarium and the Field.—No: 2. Bry ltec® Hasta hse th cles scons NOs Oa ly pets .50° - Botany, Vol. I, No. 4—Phycological Memoirs. By De Alton Saunders. 75 Botany, Vol. I, No. 5—Some Observations on the Development of the. } Karyokinetic Spindle in the Pollen-Mother-Cells ‘of Cobzea ; scandens Cav. By Anstruther A. Lawson... ........0.. .05-. a4 Paap Botany, Vol. I,.No: 6—The Origin of the Karyokinetic Spindle in Pas-- siflora ecenila’ Linn.’ By, Clara Li¢ Walliams cn Lo: fs ye ah oe 35 Botany, Vol.I, No. 7—The Nature of the Association of Alga and ; Fungus in Lichens.. By George James Peirce, .........+..+ 35 Botany, Vol. I, No, 8—Californian Hypogzeous Fungi. By Has Wales 5% ae Lae Res Harkness, Peel othr ew EA ERIE SR BURT GUY T CRC AIL Haein CFL ity Mat pF yi iia rec sits) ‘Botany, Vol. I, No. 9—Studies on the Flower and ‘Embryo of Spar- Cee ihe te na ee ganium. By Douglas Houghton Campbell, ......./.......-. i ASO Ca Man ety gee ’ Mathematical-Physical, Vol. I, No. 1—On Rational Quadratic Cee inf ~ , formations. . By M. W, Haskell... 0c. .c0eu-ebes seetes i ys ae! Meihatiaia dey sir Vol. I, No. 2—The Quadratic Cremona ' Transformation. be Leonard E, Dickson.....0 2.22.02: ‘35, Mathoriacat i se ver , No. 3—On Curvilinear Asymptotes: ABICON KR ae Cds aan au yedaY coe ben's Capa y _.Mathematical-Physical, Vol. I, No. 4—Systems of Simple Groups pay Miso ie rived from the Orthogonal Group. By Leonard E. Dickson... .25 All subscriptions, applications for exchanges, and i aie ag concerning the | may tae “pablicatiqns should be addressed to Lp cat Wie oh De S ER pa CRE at SBN Sect Rt sat O'B. GUNN, Corresponding Secretary, ; j iFM Cte tare AM 4 California Academy of Seleneesy® "AEA Tis hes Be nS PAL gitar ae San Francisco, California, in, FAT EBE TAS hy NGS | PROCEEDI OF SCI ADEMY AC THI ERIES La “y" .) SB) Rab . ORNIA L i RD oO. N Vou Oat, Wild Ls 5 pte. yo E. ~ and 4, 4g e f the a fatua. ven we ats Sa NON, AN STIN land Stanfi te U M ILLIA ity. 8 in Botany 9 par 0 id J Or Assistant oP ITH -! ued A; Iss : Z, 1900. ISCO: il 2 SA Si a sol a. Nod irre Ler kt | i ee _ PUBLICATI ; LAPS: = _ Cuartes H. Giveert, G 2. ek ee ati vag Ae : miiam E. RITTER, - _. _Dovueras H. BELL, ~ ") . “y ‘ 2 ae A Bet : if 4 tf PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, THIRD SERIES. BoTANny. Von. De Nosito: A Morphological Study of the Flower and Embryo of the Wild Oat, Avena fatua L. BY WixtuiaAm AUSTIN CANNON, Assistant in Botany, Leland Stanford Junior University. WirtH FIve PLATES. Issued April 21, 1900. SAN FRANCISCO: PUBLISHED BY THE ACADEMY 1900. PROCEEDINGS OF THE ACADEMY. (OcTAVO.) Third Serves. BOTANY. Vou. 1. No. 1—A Morphological Study of Naias and Zannichellia. By Doug- las Houghton Campbell ....-..---++seeerertetrsrtt i $1<00- No. 2—Studies in the Herbarium and the Field.—No. 1. _ By Alice TeqstyeOde eek cae Coa ee ae ee eee ae ee SRF 25 No. 3—Studies in the Herbarium and the Field.—No. 2. By Alice —Eastwood.....- sees seee ence certs ea wattta Gy tectene et woke oe .50 No. 4—Phycological Memoirs. By De Alton Saunders.....---.+-+- 75 No. 5—Some Observations on the Development of the Karyokinetic Spindle in the Pollen-Mother-Cells of Cobeea scandens Cav. By Anstruther A. Lawson......-----++ erste rtseetrs ests 35 No. 6—The Origin of the Karyokinetic Spindle in Passiflora ccerula [inn > By Clara Ls Williams. 22> + 22+ +8 aeons 35 No. 7—The Nature of the Association of Alga and Fungus in Lichens. By George James Peirce-..2s-.--nrpetrireee tts 35 No. 8—Californian Hypogzous Fungi. By H. W. Harkness....---- Sat i No. 9—Studies on the Flower and Embryo. of Sparganium. By Douglas Houghton Campbell... .-- Se LAE o Wie LS eR Se ae ole ip 50 No. 1o—A Morphological Study of the Flower and Embryo of the Wild Oat, Avena fatua L. By William Austin Cannon...... 50 All subscriptions, applications for exchanges, and inquiries concerning the publications should be addressed to : J. O'B. GUNN, Corresponding Secretary, California Academy of Sciences, San Francisco, California. . se sk, a eos any: Bi i DATE DUE ie) w > fe] x 4 f°] a a < i) — oo rw ere or s : o14 3 5185 00288 39 ) | Ga TU ah . SA NC =f Ms See ~ ¥ 2’ - eae fare eae ee es ‘ CA) 2 Hi Bata 2 Or 6,4 mS ETNA oes (O59 = vad! ee < ~ #, SOEC rs J #, Ta bue tats rt e peeks Rises ty an NG Mheban o OF ee ee Tete teenth a7 rf. iat 4 Ths tel # dete» Ti » ‘ * eSts4 Se a: ~~ Sette ae one atid a ak paPre at ee NI oe vn Pe lerece sh: bee Pi rie por cecotsese sl ¥ feaSe