1; ee Ve a \\C¥ ni ee Ae ‘oe 4 Soe oy OF THE TORREY BOTANICAL CLUB VOL. 40 FOUNDED BY WILLIAM HENRY LEGGETT, 1870 EDITOR EDWARD LYMAN MORRIS * ASSOCIATE EDITORS JEAN BROADHURST MARSHALL AVERY HOWE ERNEST HERBERT MAULE RICHARDS ALEXANDER WILLIAM EVANS ARLOW BURDETTE STOUT NORMAN TAYLOR D CA é 4 Sl NEW YORK 4 1913 '~ PUBLISHED FOR THE CLUB fe ‘ ‘se THE New ERA PRINTING CoMPANY LAN ae & Deceased, September 14, 1913. CASTER, Pa, QK| Tes0s 1914 y 4A CONTENTS Lioyvp, Francis E. Leaf water and stomatal movement in ic and a method of direct visual observation of stomata in situ. - GorTNER, Ross AIKEN, and Harris, J. ARTHUR. On a possible lation: ship between the structural peculiarities of normal and ils fruits of Passiflora heer and some et aa -chemical pipers their expressed juices - 27 RYDBERG, PER AXEL. Studiés on ‘ie Rucky Mountain flora-XXVII - - 43 SMITH, Mae MorGAN. Tetradesmus, a new four-celled coenobic alga (plate 1) Oe eee EvANs, Agee W. See iioomus. Henn toe ys Development of the peristome in Ceratodon purpur - ButTLer, O. A note on he significance of sugar in alees of ae tuberosum (plate 2) - 110 PENNELL, FRANcIS W. Studies in the Agalinanae, a Sah of she Rhinanthaceae - - 119 LEVINE, MICHAEL. Studies i in ‘thie cytology of ihe Hymenomyoetes, espe- cially the Boleti (plates 4-8 ~ 457 SLossON, MARGARET. New ferns from oneal hawace 1 (plat — - 183 pregerie WINIFRED J. A taxonomic cape of the rheahanesr of the Hawaiian Islands—III (plates 9-12 193 PICKETT, F. L. The development of he embry0- “sac of Asiilens tri- phyllum (plates 13 and 1 RicG, Georce B. Is rnd a iste: in cite distributios of Nereocystis Luetkeana? - - 237 BRAINERD, Ezra. Four hybrids of Viola ‘Dedatilidg (plates 15-17) - - 249 BICKNELL, EUGENE P. Viola obliqua Hill and other violets - 261 Knupson, L. Observations on the inception, season, and soatok of cambium development in the American larch sake: sonra = a Koch] (plates 18 and 19) - 71 GLEASON, HENRY ALLAN. Sede a on ie West fallin erates sith one new species from Mexico - - 305 Hoyt, W. D. Some toxic and antitoxic effects i in Sateen col Gamnaves - 333 KUNKEL, OTTo, The production of a pheas iohees xs the aecidiospores of Caeoma nitens Burrill - See - 361 Strout, A. B. A case of ‘at onsen in i accion (lati a - - 367 HARPER, Rotanp M. A botanical cross-section of northern Mississippi ith notes on the influence of soil on vegetation (plates 21, 22) - 77 PENNELL, Francis W. Studies in the A aoioenens a subtribe of the thaceae - 401 ANDREWS, FRANK ial ‘ua ee ae Maia: Seine aicitioge concerning the reactions of the leaf hairs of Salvinia natans_~ - - 441 Harris, J. ARTHUR. On.-the relationship between the number of ovules formed and the capacity of the ovary for maturing its ovules into seeds 447 iii : lv CONTENTS RYDBERG, PER AXEL. Studies on the Rocky Mountain flora—XXIX - 461 Griccs, RoBert F. Observations on the a a ea of the Sugar Grove flora - FromME, F. D. The culture of eae rusts in hs greenhouse - - - as MACKENZIE, KENNETH K. Notes on Carex—VII - 529 MottiErR, Davip M., and NoTHNAGEL, MILDRED. The iaveloniniak and behavior of the cinchincts 3 in the first or heterotypic mitosis of the pollen mother-cells of Allium cernuum Roth (plates 23 and 24) - - 55 Berry, Epwarp W. Contributions to the Mesozoic flora of the Atlantic coastal plain—IX. Alabama - 567 Picarp, Maurice. A bibliography of ees on meiosis 5 et mitosis in Angiosperms~ - - - - CHAMBERLAIN, EDWARD B. Edward tian Morris (Portrait) - - 599 BICKNELL, EUGENE P. The ferns and flowering plants of Nantucket—XI 605 KUNKEL, Otto. The influence of starch, peptone, and sugars on the toxicity of various nitrates to Monilia sitophila (Mont.) Sace. - - 62 PickeTT, F. L. Resistance of the ga coms of anwar rhizophyllus to desiccation - Britton, ELIZABETH Beerrsee: West iidiea sicomiial (plate ue - 653 RYDBERG, P. A. ria etc notes on the “iain Mountain region—I. Alpine on - - 077 SLOSSON, MARGARET. Pet ferns pane opel Ame 18 Gitate 26) - 687 INDEX TO AMERICAN BOTANICAL LITERATURE 89, 131, 187, 243, 295, 373, 457, 523, 591, , 64, én INDEX TO VOLUME 40 erat - - = - . . - - 697 Dates of Publication No. 1, for January. Pages I- 42. Issued 20 February, 1913- No. 2, for February. 43- 96. 18 March, 1913. No. 3, for March 97-136 7 April, 1913 No. 4, for April 137-192 9 May, 1913 No. 5, for May 193-248 20 May, 1913 No. 6, for June. 249-304 18 June, 1913 No. 7, for July. 305-376 18 July, 1913 No. 8, for August. 377-460 13 August, 1913. No. 9, for September. 461-528 10 September, 1913 No. 10, for October. 529-598 15 October, 1913- No. 11, for November 599-652 24 November, 1913- No. 12, for December. 653-712. 6 January, 1914. Errata Page 45, line 8 from bottom, for americanus, read americanum. Page 46, line 1, for elatus, read elatum Page 70, line 11 from bottom, for Oreoxys, read Oreoxis. Page 496, line 14, for paramoena, read peramoena, Vol. 40 No, 1 BULLETIN OF THE TORREY BOTAN ICAL CLUB or JANUARY 10913 Leaf water and stomatal movement in Gossypium and a method of direct visual observation of stomata in situ Francis E. Litoyp In a previous paper* it is shown that the rates of transpiration in cut shoots of the ocotillo, Fouguieria splendens, recorded by simultaneous volumetric readings and weighings, are not paraliel, but that the loss of water from the plant during the day is in excess of that taken up by the cut end of the shoot from the porometer. This result is in general harmony with the findings of Eberdtt with rooted plants of Helianthus annuus. It was not, however, found to be true in my study of the ocotillo that the loss of water takes place at a constant ratio during the hours of day- light, since the whole relation between the income and outgo may be reversed within a short space of time even during daylight by an apparently slight modification of the environmental condi- tions. This ready susceptibility of the plant lent color to the idea that the differences indicated by volumetric and gravimetric readings are measures of differences in the leaf moisture content, to be more briefly referred to as leaf water in the present paper. Determinations of the amounts of water held within the leaf at various times of the day, these being calculated in percentages of dry weight, showed that such: expected differences do occur, _ but the evidence was incomplete in that the amounts of leaf water were not calculated to a constant, since it must be assumed that sped BULLETIN for December 1912 (39: 567-631. pl. 40-45) was issued Jan. 2.] * The relation of transpiration and stomatal movements to the water-content of the leaves in Fouquieria splendens. Plant World 15: 1-14. Ja 1912. Tt See Burgerstein, A. Die Transpiration der Pflanzen 17. Jena, 1904. 1 2 Litovp: LEAF WATER IN GOSSYPIUM the dry weight of the leaf increased during the day. The conclu- sions drawn rested more especially on the assumption that the leaf water at the close of day is not in material excess of that at the beginning, and upon the fact that during the afternoon there is a sharp rise in the amount of leaf water in spite of a constant, if not a still increasing, dry weight. The hope that during the summer of 1911 the more convincing data based upon constant leaf area could be obtained was not realized, and the evidence therefore still remains incomplete. In September 1910 a similar attack was made upon the cotton plant at Auburn, Alabama, in which analogous conditions with respect to leaf water were found. During the present year the question has been further studied, the material being obtained from a pure strain of Dixie cotton, the seeds of which were supplied me by the Bureau of Plant Industry. Plants were grown at Auburn and, under irrigation, at Tucson, Arizona, in the experimental grounds of the Desert Botanical Laboratory of the Carnegie Institution of Washington. My thanks are due to Dr. D. T. MacDougal for his courteous co- operation. The present paper reports upon a portion of the data obtained bearing upon the variation in the amount of leaf water and upon the behavior of the stomata. METHODS DRY WEIGHT AND LEAF WATER. The first two series of leaf samples, the figures for which are given in TABLE I, consisted of leaf material placed in tared, cork-stoppered bottles, weighed, oven-dried thoroughly and reweighed. The samples were taken at two hour intervals. The actual work was done by Mr. C. S. Ridgway, to whom my thanks are extended. The samples on which TABLE III is based were taken at assumed critical times, and treated in the same manner by myself. The data in the remaining tables were obtained by cutting circles of the lamina with a sharp cork borer, placing them forth- with in tared vials and weighing before and after thorough drying. The area of these circles, of which between fifty and sixty were taken for each sample, was calculated, and the exact number of circles taken determined a second time after drying. All data were Lioyp: LEAF WATER IN GOSSYPIUM 3 calculated to 100 square centimeters of leaf blade. In this way the changes in dry weight and in leaf water for each period were determined. STOMATA. The measurements given in TABLE IX are average micrometer measurements of a large number of stomata for each period, made on material fixed in absolute alcohol.* Those in TABLE X are measurements of living stomata made in the field by the following method, first hit upon by me during my work at the botanical laboratory of the Carnegie Institution of Washington, at Carmel, California, during the summer of IgII. A microscope, provided with a condenser and a 4 mm. objec- tive with long working distance, was provided with a cooling cell. This was extemporized by wiring a Soyka flask beneath the sub- stage apparatus. For working in the field, a heavy camera tripod served as a stand capable of ready adjustment in height and position. The illumination consisted of direct sunlight, or when that was not available, a strong artificial light. I used a small miner’s acetylene lamp, on the whole fairly satisfactory. A small arc light probably would be better, and indeed necessary when the leaves are very thick. By means of such strong illumination, properly centered and controlled with the iris diaphragm, it is possible to measure accurately living stomata of leaves with a thickness of 5 mm. without at all injuring them. The method is adapted to the observation of the stomata on a great variety of leaves, including many which are densely covered with trichomes (e. g., Parthenium argentatum Gray; Chenopodium sp.), which being out of focus do not interfere with the vision under strong illumination. The stomata at the bottom of deep pits, such as are found in the Cactaceae, also can be seen, but so far this has involved the removal of a thick slice of the underlying tissues. The method presents the further advantage, one of great * importance in field work, of permitting the repeated observation of the living stomata, or indeed of a single one, during a sustained period, without injury to the leaf or its removal from the plant. This method has, of course, its limitations in view of the size and *Lloyd, F.E. The physiology of stomata. Carnegie Inst. Wash. Publ. 82. 1908. 4 Lioyp: LEAF WATER IN GOSSYPIUM shape of the microscope,* but these limitations are not as strait as one might suppose. For the direct visual observation of the living stomata in situ, for any reasonable desired period, the method above outlined has a unique value for certain purposes. It will be found, I venture to believe, to be invaluable in supple- menting and checking the results obtained by such methods as that of alcoholic fixation (Lloyd, /. c.), of the hygroscope (of whatever type), of the porometer,t and of the stomatograph.t In using the method one must have a care for his eyes. The adjustment of the illumination must be done with the protection of a color screen, and for this purpose a leaf interposed in front of the objective has served well enough. Under field conditions, especially when one is working for a considerable length of time, dark eyeglasses are very desirable, more particularly if the sur- roundings are highly illuminated. Small circular discs of dark- tinted glass in various shades to be placed above the ocular have been found advantageous, more especially when the leaf is rather thin, say as thin as the leaf of the cotton. In the actual observation of the stomata the leaf is placed directly on the stage without any accessories in the form of glass slides or covers. It is held in position by gentle pressure upon it of the index and middle fingers, which, also, embrace the end of the objective. In the eyepiece a micrometer is placed. The absence of a glass slide, which is in any event unnecessary, brings the stomata nearer the focus of the condenser and so improves the definition. As one is dealing with a brilliant area of light, it is quite necessary to have it central, since oblique illumination causes bright reflections from the surface of the stomatal outer vestibule, which may mislead the observer. LEAF WATER IN PERCENTAGE OF DRY WEIGHT If we consult TABLE 1, which contains the percentages of leaf water calculated to dry weight for two parallel series, treated as nearly alike as possible, it will be seen that when so expressed EAS CEREIRRMIESS ATE a Rnke esas iser egal estes ee mio * An instrument of special design, now in course of construction, will obviate many inconveniences, + Darwin, eres boas agate M. On a new method of estimating the aperture of stomata. Proc. Roy. Soc. B. 84: 136-154. 1911. t Balls, W. L. The “Stomatograph.” Nature 87:180. 10 Au ro1t. Lioyp: LEAF WATER IN GosSYPIUM 5 it varies between the extremes of 318 and 220 per cent. The greatest difference between the two figures for any hour is 26, this greatest discrepancy being found at the 8 and at the 14 hour on September 17. If we assume an error of 13 units and apply a corresponding correction by addition to the smallest and by sub- traction from the largest percentage, a difference of 72 units is still afforded. If the correction is similarly applied to the percentages in either series, the difference may be reduced to 46 units, as between the 8 and 14 hour figures. It will appear from these figures that the error of observation is not great, and is probably within five per cent., a conclusion well sustained by the general correspondence of the figures for the two series. It is, further, probably entirely true that the differences are not due to a material error of measurement, but are actual differences in the leaf water of different parcels of material. Since these were not taken by myself, I have compared the differences with those displayed by my data for the ocotillo, upon which I based my conclusions embodied in my paper above referred to,* and I find that dis- crepancies in similar amounts are to be found in them. The same is true of the data which appear in the present account beyond. They may be due to individual differences, or to dif- ferences in groups of plants. under various external conditions, or in the positions of the leaves taken. The general correspond- ence between the two series (FIG. 1) is, however, sufficiently close to warrant drawing the conclusion that during the night there is a rise in the leaf water content (which probably had been pre- ceded by a greater increase previous to the 16 hour) and a decided decrease during the day till 2 P. M. The sharp rise and equally sharp fall indicated between the 18 and 22 hours is puzzling, since if we refer it to change in dry weight, the additional difficulty is introduced of accounting for marked changes in the same. Aside from this peculiarity of behavior the general rise in water content may be referred to the reduction of dry weight, but this involves the assumption that the dry weight was reduced much more rapidly in the period preceding the 18 hour (say between the 14 and 18 hour) than subsequently. A somewhat similarly sharp * The relation of transpiration and stomatal movements to the water-content of the leaves in Fouquieria splendens. Plant World 15: 1-14. Ja 1912. |. Leat Water 4 N 8 240 |- 8 Cy Q 43. 220 |— as OEF - a2 as. 00 1. a 5 Searle | L | | fi 16 Hour 20 22 24 2 4 6 & 10 12 ld Fig. 1. Gossypium herbaceum. Graphs for leaf water for two series and for their averages; and for average transverse stomatal dimensions. : CAOTT WNIdASSOL) NI WALVM AVAT Lioyp: LEAF WATER IN GOSSYPIUM 7 rise in water content is indicated after the 6 hour, September 17; continuing till 8. The tables following show such behavior in only one instance (series 12), but it must be understood that the data therein contained were based on material collected on bright days, while those now being considered were taken on a morning (of September 17) during the first few hours of which a general haze prevailed. The single exception, offered by the old leaves collected on October 13, I91I (series 12, TABLE viit), differs quantitatively, there being here a difference of 15 units (251 to 266 per cent.) as compared with Ig units (284 to 303 per cent.) in the case before us of September 17. The value of the comparison is slightly compromised by the fact that the data of TABLE vill indicate a decrease of dry weight, in an amount that may, however, fall within the limits of error. If no change in dry weight is assumed, the difference in water content in percentage of dry weight would be somewhat reduced. In view of the further fact that the data for both TABLES v and VIII were obtained after an increase in soil moisture, it seems entirely possible that under certain conditions the first two hours of daylight may be accompanied by an increase in leaf water. This involves as a corollary that the leaf, at the hour of sunrise, may not be completely plethoric of water. In order to evaluate with some degree of exactness the meaning of the graph embodying the data of TABLE I, I have assumed that on September 17 the dry weight of the leaves increased in the amounts of 9 per cent. on the original weight in the first four hours (6-10) and 17 per cent. in the whole period of eight hours, namely 6-14. These amounts are taken from TABLE VII, series 8, and represent fairly closely the average performance for young leaves. The weighings for the 6 hour are taken as a basis for the calculation, and the leaf water content is assumed to remain constant, the dry weight only increasing. By comparing the ratios thus obtained with those actually observed, as displayed in TABLE II, we see that the calculated value of the ratio at the 10 hour is less than the observed, and at the 14 hour greater (graph abc, FIG. 1), the latter indicating a net loss of leaf water. If however, an increase a 30 per Soar on the original dry weight be assumed, an a ly justified in view of the data ca aa os 8 Lioyp: LEAF WATER IN GossyPIUM in other tables, a net loss of water would not be discoverable (graph abd, FIG. 1). It is evident, however, from the directions of the graphs that a negative fluctuation was in progress from the Io to the 12 hour, followed by increase afterwards. It is also evident that a positive fluctuation begins after the 14 hour, if we. suppose the datum for the 16 hour, September 16, to apply ap- proximately to the same hour on September 17. Although the above evidence is quantitatively somewhat vague, this result is probably due to the application of assumed dry weights based upon data obtained in the following year. Had such data been obtained at the time, their employment would lead to a surer inference. This appears from similar manipulations of the determinations contained in TABLE 1 of leaf water and dry weight for unknown leaf areas. In the first three columns are given the known data; in the fifth the average dry weights from TABLE v, which were obtained two days later, these being relative to 100 square centimeters of leaf and assumed to be applicable to August 24. Since the two days (August 24 and 26 at Tucson, Ariz.) were practically identical meteorologically, this assumption is probably entirely justified: By dividing the assumed dry weight into the observed dry weight for each hour the approximate area of the leaf sample taken at that hour is obtained. This affords one term of a ratio (column 4), the other of which is the observed leaf water (column 3) by which the leaf water per 100 square centimeters is derived. This, for each of the three hours of observation, is found in column 7 and is plotted in graph 1, FIG. 2. | ; Just above this is graph ta, the coordinate points for which were determined as follows. (See TABLE Iv.) The same ratios of increase of dry weight were assumed as above. One of the observed dry weight readings in TABLE 111 was then taken as a standard, and the dry weights for the other two hours were deter- mined by means of these ratios. By assuming the leaf water to be constant the inverse ratios, expressing leaf water as percentage of dry weight, were obtained (column 2, TABLEIv). By comparing these with the ratios (column 4) of the data recorded for each hour (columns 2 and 4) differences are obtained by subtraction which determine the position of the graph of observed leaf water Lioyp: L&raF WATER IN GossyPpIUM 130+ Leak Water Dry Weight. 6 Hour 8&8 10 l2 M4 Fig. 2. Gossypium herbaceum. Graphs for leaf water and dry weights of leaves for the series indicated by numbers corresponding to their numbers in the tables. The ordinates in this and Fic. 3 are the same for the upper and iower portions but have been shortened between 7o and 130 grams. Young leaves. 10 Litoyp: LEAF WATER IN GOSSYPIUM relative to the graph of constant water when the latter is assumed to be horizontal, as it would be if plotted relative to constant leaf area (column 6). By making the value of the 6.20 hour figure say 160, we obtain results which differ little from those in column 6, TABLE Iti, in calculating which a variable due to a somewhat inconstant relation between leaf weight and area was admitted. It is apparent that a much larger increase in dry weight for the first four hours of the day than that assumed, together with far different march of events during the whole period, must have obtained in order to.obviate the conclusion that on August 24 there was a net reduction in the amount of leaf water followed by a material recovery. In addition to graphs 1 and ta are to be found in FIG. 2 the graphs for the young leaves in the remaining tables. The general similarity of the lot further impels the conclusion that graphs I and ta are not wide of the truth. I have taken the above somewhat indirect method of drawing the conclusions desired with reference to the observations recorded in TABLE Ill, for the purpose of developing a comparison of the observations for the two days, 24 hours apart, at Tucson. We have already examined those of August 24, presented in TABLE II. Those for August 26 are to be found in TABLE v, in which the weights are relative to 100 sq. cm. of leaf blade. Although there was a marked fluctuation in the leaf water on August 26, the loss of water between the 6.20 and 10.20 hours being as great as that on August 24, it is to be noted that the recovery after the 10.20 hour was more rapid on August 26 (graphs 2, 3), and that the absolute amount of leaf water was considerably greater, viz. about 25 per cent., on this day than on August 24. Since the leaves from which the samples were taken were of uniform texture and at similar stages of development, this difference cannot be attributed to variation in these. The difference is due quite probably to the circumstance that on August 25 the ground in which the plants were growing was irrigated. This, if the proper explanation, carries with it the suggestion that it would be well worth while to determine with what precision the soil water conditions may be registered in the condition of the leaves. It is quite possible that the needs of the plant for a water supply Lioyp: LEAF WATER IN GOSSYPIUM 11 to its roots could be determined by following the changes in leaf water from day to day, and so anticipate a condition of necessity which might curtail growth for a period. Obviously such a pro- cedure would be of practical use only where water is under control. We now turn to consider the tables which present data relative to a constant leaf area, namely 100 square centimeters. For the purpose of comparison in what follows, the data em- bodied in TABLES v—vitI inclusive have been segregated in two sets of graphs, those (in F1G. 2) for young or full-grown but not old and indurated leaves, and those (FIG. 3) for old leaves, rather the worse for wear. They were indurated, torn and more or less discolored, but a consistent effort was made to collect the material from leaves with a healthy appearance. The three series (1 to 3 incl.) for Tucson, Arizona, are included if FIG. 2, since the leaves, while mature, were not at all indurated and, indeed, had perhaps not arrived at full growth, though they were well developed. Of the seven series of samples from young leaves one only, viz. no. 4, appears to indicate an aberrant behavior, especially as to increase in dry weight, but evidently so as to the loss of water also. If we disregard this case, for which no explanation is offered, the remainder show a general accord. Of the six remaining cases, two (2 and 3), those for Arizona, show an increase considerably in excess of those for Alabama. The ratio of increase for Arizona is 100 to 120 for the first four hours and 100 to 136 for the whole period of eight hours, beginning at 6.20 for the first series; the ratios for the second series are 100 to I12 and 100 to 124. In Alabama the greatest gain for a like period was in the ratio of 100 to 121, while the lowest was from 100 to 109. Stated in terms of increase of photosynthates, we have for the Arizona plants an accumulation at the rate of 2.25 grams per hour per square meter for the first four hours, and 2.00 grams for the second four hours, for no. 2, TABLE V; the corresponding figures for no. 3 are 1.50 and 1.50. For the young leaves of Ala- bama plants, excluding series 7, TABLE VI, the range of increase (when there was any increase at all) is from 0.5 to 1.6 grams. Sachs’ determinations for the sunflower were 1.88 and 1.7, and for the cucumber 1.5 gr. Brown and Escombe’s datum for the sunflower is, however, 0.714 gr. Although my own data were Lioyp: LEAF WATER IN GOSSYPIUM ——— ————oo —_—_—— -_———— a Dry Weight. L ] l | 6. Seer 8 i | we SH 10 12 14 Fig. 5. Gossypium herbaceum. Graphs for leaf water and dry weights of leaves for the series indicated by numbers corresponding to their numbers in the tables. leaves. Lioyp: LEAF WATER IN GOSSYPIUM | 13 not obtained with a view primarily to obtaining light on the accumulation of carbohydrates, the general correspondence of the Alabama figures with those quoted may be noted. The greater efficiency of the Arizona plants possibly may be due to the fuller development of the leaves. This would be indicated by the performance of the old leaves on October 1 and 8 (series 6 and g), which showed an increase of 30 and 26 per cent., but more surely, if it were not for the general irregularity of their behavior and the entire failure to get an increase at all in series 12 and only a very slight one in series 5. On the other hand the light conditions in Arizona are much more favorable in view of the frequent cloudiness in Alabama. The days on which the Alabama material was collected were partially cloudy, while that from Arizona was taken on continuously clear days. The old leaves betray a much more erratic behavior and the data lead us to no safe conclusion. In the two cases, TABLE VI, series 5; TABLE VII, series 9, however, in which there was a marked increase in dry weight, there was also a marked re- covery of leaf water after the 10 hour. By contrast, however, the leaves that contained the greatest quantity of water, series 12 of October 13, were the least efficient photosynthetically, a circumstance which throws doubt on the inference that the failure of the leaves of series 5, taken from all parts of the plants, to add to their dry weight after the 10 hour, was due to a marked loss of water during the corresponding period. STOMATAL MOVEMENT A glance at FIG. 2 will show that after the 10 hour has been reached there is a distinct tendency to recover the leaf water lost during the earlier hours. It is important to inquire whether this tendency, which begins to make itself evident during the part of the day when transpiration rates may be assumed to be the highest, is in correlation with the behavior of the stomata. Balls,* in Egypt, found by means of his newly devised stomato- graph that the stomata of the cotton slowly close as the hottest part of the day approaches, completing the closure as the sun passes behind neighboring trees at 13.40 hr., remaining closed all * Balls, W. L. The “Stomatograph.” Nature 87: 180. 10 Au 191t. 14 Lioyvp: LEAF WATER IN GOSSYPIUM the succeeding night. They open slowly after sunrise, but more rapidly after the sun strikes the leaf at about 7, attaining the maximum opening at 9, but thereafter closing steadily in spite of brilliant illumination. So far as I am aware this is the only record of the behavior of the stomata in cotton. The measurements given in TABLE x, made microscopically and at the same time by two observers, Mr. C. S. Ridgway and myself, in the manner earlier described, correspond in a rather striking way with Balls’ conclusions, just indicated. Each for- mula represents the minimum and maximum width of the stomatal pores on the upper and under sides of the same leaf. When a very few stomata displayed extreme dimensions, these are enclosed in parentheses. It may be regarded that these would on account of their small numbers affect but little the general diffusive capacity of the stomata taken as a whole. It is evident that during the night and until 6.20 there was no measurable opening movement. This must not be taken to mean that all the stomata were entirely closed. TABLE IX gives stomatal dimensions from preparations made by the alcohol method, in which it is evident that some opening in a few stomata may be observed at any time during the night. The initial opening on September 30, 1911, occurred about 6.30, from which hour on a progressive opening movement was followed, the stomata of the lower faces opening somewhat in advance of those of the upper, though some exceptions to this appear, as at 7.50 and 8; while Livingston and Estabrook* found, in several kinds of stomata, that the upper close and open more rapidly or close more com- pletely than the lower. The maximum average pore width was attained during the period between 7.45 and 9, though the widest openings were seen at 8.20. Because of the extreme sizes of opening then noted, leaves of similar size and exposure were examined to verify the observation, but they were no more seen during the morning, except in one instance an hour later. The closing phase was at once entered upon, but it is evident from the figures that this movement was less uniform in all the leaves taken together than the opening. This may be readily understood * Livingston, B. E., and Estabrook, A. H. Observations on the degree of stomatal movement in certain plants. Bull. Torr. Club 39: 15-22. 12 F 1912. Lioyp: LEAF WATER IN GOSSYPIUM 15 when it is appreciated that the stomata of cotton are quite sensi- tive. I observed, both in Alabama and in Arizona, that at the same hour a leaf in the shade and one in the sun were physio- logically antipodal in this regard. Thus, at 8.30, August 24, at Tucson, Ariz., the stomata of a leaf in the shade were closed both above and below, while those fully exposed to sunlight on another leaf were 0-10u wide above, and 2-44 ‘below. Similar conditions are displayed by the leaves observed at the 1o hour, TABLE x. At the same hour it was noted that the stomata near the apex of a leaf were closed while those near the base were open, a condition readily understandable if wilting is progressive from the apex of the leaf downward. As between young and old leaves, the latter being still physio- logically active, it is not clear that there was a more pronounced tendency in one kind than in the other either toward opening or closing, nor that the leaves near the base of the plant were more favorably placed than those three feet above, at or near the top. Indeed, that the upper leaves close their stomata less early than those near the base seems indicated, and this is in harmony with Rringsheim’s findings,* who believes that the apical parts of the plant withdraw water from the older, the result of more rapid transpiration from the younger organs. The data of the table which we have been considering must be regarded, however, rather as an indication of the usefulness of the method of microscopical observation in the field after the manner already described. The measurements were taken rather at random, and as the plants were nearing the end of their career, the larger leaves being somewhat passées, while the young were from ‘‘second growth” shoots, at the tops of the plants. Their general indications, however, I believe, may be relied upon, namely that the maximum opening of the stomata is reached at about 8.30 in the morning. It has been the invariable experi- ence of three observers during the season of IgII at Auburn, and of one of them at Tucson, that wilting of the leaves becomes first evident at about this hour or somewhat later each day, when sun- light conditions prevail. The reduction of leaf water leading to *Pringsheim, E. Wasserbewegung und Turgorregulation in welkenden Pflanzen. Jahrb. Wiss. Bot. 43: 89-144. 1906. 16 Lioyp: LEAF WATER IN GOSSYPIUM this condition* has been demonstrated quantitatively above, the minimum amount being found, in these determinations, at the 10 hour or somewhat later. It becomes evident that the opening of the stomata is accompanied by a net loss of water by the leaf, more being given off by transpiration than can be obtained to replace it, as I have shown to occur by comparative volumetric and gravimetric determinations in Fouquieria splendens.t The results of similar experiments with cotton may be said, in an- ticipation of their publication, to harmonize fully with those on Fouquieria. Concerning the relative behavior of young and old leaves it may be further pointed out, that so far as the evidence goes in the present paper there is little against either of the alternate views that this loss of leaf water is better resisted by the young or by the old. The numbers of stomata per unit area in the young leaves are considerably greater. An important inquiry at this point is in regard to the relative extent of the internal evaporating surfaces, t but though we do not find the answer, we may be sure that it is sufficient in young leaves to lead to wilting, and that the stomata do not head eff the net loss during the period of advancing temperature, insolation, and evaporating power of the air. It is quite possible that determinations of the amount of leaf water at smaller intervals of time would lead to the finding that the loss is more sudden, after it once begins, in old leaves than in young ones, and that this may be connected with the relative diffusive capacity of the stomata in consecutive intervals of time. I have arrived at this suggestion by a comparison of the dimen- sions of the slightly open pores of young and old stomata, and by calculating their relative diffusive capacities.§ It develops that in old stomata the edges of the pore are more stiffly reinforced, and that in opening they separate from each other throughout t Bergen, J. Y. Relative transpiration of old and new leaves of the Myrtus § Livingston, B. E., and Estabrook, A. H. Observations on the degree of stomatal movement in certain plants. Bull. Torr. Club 39: 15-22. 12 F 1912. Lioyp: LEAF WATER IN GOSSYPIUM iT nearly their whole length at once, making a long narrow opening. Thus I find that the pore lengthens from 12 to 13.5 u in one Case, and from 9 to 10.54 in another, while the width reaches 1.8 and 2 w respectively, these transverse measurements being applicable throughout the major portion of the length of the pore. Such stomata probably attain a relatively high diffusive capacity more quickly than young ones, in which pores, because of the flexibility of their edges, they attain their full length gradually. In the only series of preparations I have at hand the shortest pore I find is 4.5 long and 1 wide. Its index of relative diffusive capacity is 2.12. Assuming that in further opening its dimensions advance to 8.4 by I u (those of a similar stoma in the same prepara- tion) and then to 9 by 1.5 u, the indices would be, for these dimen- sions, 2.9 and 3.67, whereas an older stoma would have measure- ments such as 14 by 0.8 u, or 12 by 1.5 u, with the indices 3.35 and 4.24, respectively. That is, for the same width of pore, the maximum length is reached much more rapidly and a relatively high diffusive capacity more quickly in older thai in younger stomata. If relations such as these obtain, as the evidence before us appears to indicate, we may well think that they may affect the rate of incipient drying,* either with or without appreci- able wilting. Differences of this kind may help to account for the results obtained by Bergen,t who found that the old leaves of sclerophyls transpire more than young leaves which have just attained their maximum area, though he further ob- served that the same relation holds for cuticular transpiration as well. In this regard, Pringsheim’s work, already referred to, may not be overlooked. It is to my mind doubtful, in view of the results on loss of leaf water presented in this paper, that the differences observed by Bergen} as between sun and ghade leaves depend upon the water carrying capacity of the vascular tissues. Turning to the data of TABLES I and IX, as represented in FIG. I, it becomes apparent that the sharp rise in leaf water be- tween the 6 and 8 hours neither harmonizes with the results given in the remaining tables, nor does the curve of stomatal openings * Livingston, B. E. t Bergen, J. Y. Di a tee cab Alors type. Bot. Gaz. 38: 446-451. D 1904. t Bergen, J. Y. Transpiration of sun leaves and shade leaves of Olea europaea sind other tidid Daseloeehgrebon. Bot. Gaz. 38: 285-296. O 1904. 18 Lioyp: LEAF WATER IN GOSSYPIUM for the hours of observation bring out the relations between these and loss of leaf water in sufficient detail. That there should have been increase in leaf water in the amount of about 7 per cent. during the period (6 to 8) when the stomata were opening from 0.7 to 1.4m, must have been due to meteorological conditions, though it is still not clear why the maximum leaf water had not been attained by the 6 hour, unless it should be shown that such a condition does not always intervene even in the early morning hours before sunrise. The flatness of the curve of stomatal opening after the 8 hour, it is likely, does not properly represent the facts, since from the evidence in TABLE xX there was probably a considerable opening and closing between 8 and 10, with the maximum diffusive capacity reached at about g. If, aside from this, the curve of stomatal opening is true, it would seem to indicate that the loss of leaf water from 300 to 240 per cent. took place without a sufficient amount of wilting to involve closure of the stomata, and that the leaf therefore had not passed beyond the condition of drying incipient to wilting. As no observations were recorded to test the truth of this conclusion and no measurements of the meteoro- logical conditions beyond the note that there was a general haziness until the 10 hour were made, we perhaps may be permitted to draw the lesson, that in such studies it is of utmost importance that the integral effect of such conditions should be known before any adequate interpretation of the water relations of the plant may be had. At present the most efficient method to this end is that of measuring evaporation with the blackened porous cup.* Ina future paper it is proposed to display the results which have been obtained in field studies of cotton in the light of such integration. Reference has been made above to the daily wilting of the leaves, observable at first at about the 9 hour. The stomatal measurements would seem to indicate that after this hour there is a rather steady closure. It therefore would be of importance to determine at what hour the maximum transpiration rate occurs when these conditions prevail. In anticipation of such determina- * Marinos B. E. A radioatmometer for comparing light intensities. Plant Wor : 96-99. Ap 1911; Light intensity and transpiration. Bot. Gaz. 51: aa D tort. Litoyp: LEAF WATER IN GOSSYPIUM 19 tions, now possible by the use of the method I have described above, and with reference also to F. Darwin’s* verification in cooperation with Pertz, by means of the porometer, of his earlier work on the temporary opening of stomata during wilting, I measured by weighing the rates of transpiration of a piece of a cotton plant during wilting. After the cutting off, the cut was greased and the piece was suspended on a balance, the loss of weight being determined at intervals of several minutes, beginning at 7.22 when the stomata were open but had not reached their maximum. The leaves were turgid. The figures obtained (TABLE X1) show that between 8.02 and 8.07, following a steady fall in the rate, there was a temporary rise, and while this was very small when compared with Darwin’s data, it may be a phenomenon comparable with that observed by him. However, that the stomata open after wilting sets in still appears doubtful to me. I have watched microscopically the stomata of the cotton, in addi- tion to those of several other species, during the wilting period, beginning with the partly as well as fully opened condition, and I have never observed any such movement. There is at first no change. As wilting sets in, closure commences and proceeds steadily till complete, and I have observed no subsequent opening. _ Obviously further investigation is required to throw light on these differences. CONCLUSIONS In the foregoing paper a method for the direct observation and measurement of stomata im situ is described. The method is ' adapted to field work by night and by day. _ Leaf water, stated in percentage of dry weight, was found to vary in the cotton plant under usual conditions between 318 and 220 per cent. On the day of observation the minimum leaf water content was reached at the 14 hour or thereabouts. This reduc- tion represents a net loss as shown by the determinations made relative to unit area and, therefore, with quantitative regard to dry weight. The amount of loss of leaf water when thus determined is from 7 to 15 per cent. on the initial amount at sunrise, under the condi- Darwin, F., and Pertz, D. F. M. Ona new method of estimating the aperture of stomata. Proc. Roy. Soc. B, 84: 136-154. I9II. 20 Lioyp: LEAF WATER IN GOoSSYPIUM tions prevailing when the observations were made. Among these conditions it may be mentioned incidentally that the soil was well drained and rich in moisture at the time. Severer circumstances © would no doubt effect a still greater loss. The observed loss, how- ever, may be taken as indicative of a usual phenomenon, the reality of which is made evident in an observed daily wilting of the leaves beginning at about the 9 hour, detectable not alone by change in position (since this may occur as a phototropic response) but by flaccidity. The case may be otherwise stated by saying that under usual day conditions, with sunshine, the roots are unable to supply loss of water from the leaves. Balls’* view that the water supply is the limiting factor of growth, and his observation that no growth takes place under the Egyptian sun appear to be quite applicable to Alabama. With regard to the amount of growth in Alabama, preliminary measurements, prompted by the results obtained from the determinations of leaf water, indicate that even under the presumably more favorable humidity condi- tions obtaining here growth does not take place for the major portion of the day, since during the latter part of the growing season an actual daily shrinkage in stem and leaf length has been observed. I can hardly concur, however, with Balls in his view that because growth does not take place in sunshine this is to be interpreted as unfavorable. Comparative measurements on the same variety of cotton obtained in Arizona betray a no more un- favorable reduction of leaf water than in Alabama, when there is sufficient water in the soil. There is evidence that variations in soil moisture are registered in both the absolute leaf water content and in the rate of recovery after the minimum quantum for the day has been reached, while the loss during the first part of the day appears to be less affected. It would seem that the real test is the growth integer for the season, and it is not evident that, with irrigation, the conditions in the semiarid Arizona desert at all events are unfavorable from this point of view. A hot sunshiny day after all may be good for cotton, but this good may not be apparent in growth at the time. This is indicated by the amount of photosynthates formed (measured with small error due to well * Balls, W. Lawrence. The physiology of the cotton plant. Cairo Sci. Jour. 4: i-9. Ji to910; Cotton investigations in 1909 and 1910. Cairo Sci. Jour. 5: 221- 234- S$ 1911. Litoyp: LEAF WATER IN GOSSYPIUM 21 understood causes) by the increase in dry weight. Two series in Arizona gave increases of 24 and 32 per cent. for 8 hours. In Alabama, with the exception of one series of old leaves in which there was no apparent change in weight, the increases ranged from 6 to 25 per cent., the latter being one instance out of a total of seven series. Whatever may be said of increase in dimensions, therefore, it remains the fact, that in spite of the hot unmodified Arizona sun shining throughout continuously cloudless days in August, more energy in the form of carbohydrates was made available than in the similar periods in Alabama. It is proper, however, to recall that the leaves studied in Arizona were of fuller development, while those in Alabama were either rather young (10-15 cm. transverse measurement) or overmature. It is obvious that it will be of great interest to make careful measurements of growth, as indeed of other functions, for com- parison with those of Balls in Egypt. The stomata are practically closed at night, but nevertheless show a tendency to open during the early morning hours.* The more obvious daily opening begins at about 6.30, in Alabama in September, and the maximum is reached at about 8.30 or 9, after which’ closure progresses until 11 or somewhat later. A con- comitant and appreciable wilting takes place, correlated with the reduction of leaf water. During wilting there appears to be no “temporary opening’”’ of the stomata, although I have observed a measurable but not very marked rise in the rate of transpiration about a half hour after wilting starts in, followed by a sudden reduction of rate. *Lloyd,F.E. The physiology of stomata. Carnegie Inst. Wash. Publ. 82. 1908; arwin, Francis, and Pertz, D. F. M. Ona new method of estimating the aperture of stomata. Proc. Roy. Soc. B. 84: 136-154. I9I1I. Gossypium herbaceum. Litoyp: LEAF WATER IN GossyPIUM TABLE I centage of dry weight. Auburn, Ala., Sept. 16-17, r1910. Leaf water in per- Hour Series I Series IT | Average 16 274 280 279 18 266 284 275 20 203 306 299 22 287 263 275 24 282 275 278 2 205 279 287 : 4 285 291 288 6 279 203 286 8 202 318 303 to 293 305 209. - I2 244 244 244 14 220 246 233 TABLE II Observed leaf water expressed in percentage of dry weight and the percentages obtained by assumin g ao per cent. increase and 17 per cent. and 30 per cent. for the second four hours. in dry weight for the first four hours Observed Calculated Hour Dry wt | Water | Per cent. water Dry wt Water | Per cent. 6 4.15 11.79 284 Io 3-70 II.07 299 4.52* II.79 265 .* 14 5.53 12.87 233 4.86T 11.79 243 5-53 12.87 233 5.40t PE70 lo ATO. TABLE III Gossypium herbaceum. Series 1, Tucson, Ariz., Aug. 24, torr. Amount of leaf water in percentage of dry weight and the same calculated to 100 sq. cm., the average dry weights in TABLE V, being assumed as applicable to the material on which this table is based. Observed weights Hour Dry Water 6.20 10.55 34.36 10.00 10.84 28.02 13.30 12.12 28.78 Weight per 100 sq. cm Area SCRE Dry (assumed) Water ee) 2153 0.49 1.59 1918 0.565 1.465 1933 0.62 1.493 * 4.15 plus 9 per cent. T 4.15 plus 17 per cent. t 4.15 plus 30 per cent. Lioyp: LEAF WATER IN GossYPIUM 23 TABLE IV Leaf ater relative ve pee area SOM Dates from. 7 observed weights in TABLE III, g y Nn TABLE V is assumed. Ratios of water (= C) to Observed ratios water to Water to | Leaf water per Hour dry weight dry weight dry weight, sq. cm. area=C 6.20 326 b dale) 326 I0o 100 1.60 10.00 | 285 | 87.5 250 | 79.5 92 1.47 2330 1 age ee 238 73.0 95 1.52 TABLE V Gossypium herbaceum. Tucson, Ariz., 26, 1911. Weight of water and weight in grams per 100 sq. cm. of at at the eae indicated. Two series Wintad Aug. 25. | 6.20 | 10.20 14.15 Series | Dry wt. | Water | Dry wt Water Dry wt | Water 2 0.46 1.96 0.55 | 1.83 0.63 1.94 3 0.50 1.906 | 0.56 | 1.79 0.62 1.96 Ave 0.48 06 = 0.55 1.81 0.62 1.95 TABLE VI Gossypium herbaceum. Auburn, Ala., Oct. 1, 1911. Weight of water and dry weight per 100 square centimeters of leaf at the hours indicated. ies 4, young newly developed leaves near the top of the plant; series 5, mixed, old leaves from various parts of the plant; series 6, old leaves near the top of the plant. 6 10 14.30 | 18 No. : ° Hove | Water Dry wt Water Dry wt Water | Dry wt. | Water 4 0.534 1.74 0.71 1.50 0.58 I.40 0.64 1.50 5 0.50 1.66 0.54 1.36 0.54 1.24 0.56 1.38 6 0.524 1.60 0.66 1.48 0.68 1.58 0.64 1.69 0.515 1.66 0.64 1.45 0.60 I.4I 0,60 1.52 TABLE VII Gossypium herbaceum. Auburn, Ala., Oct. 8, 1911. Weight of water and dry weight per 100 sq. cm. of leaf at the ge: Sadicatna Series 7, material from alternate sides of the same leaves on ten plants; series é. young ares taken indiscriminately from plants in “‘lot 2, closely planted in rows.’ ea series from fresh secondary growth at the tops of plants; series 9, old leaves taken indiscriminately: marked “old.” 6 II.00 / 14.30 No. j : Dry wt Water Dry wt Water | Dry wt Water 7 0.5) 1.56 0.58 1.45 0.58 I.44 8 0.46 1.62 0.50 1.41 0.54 I.41 j 9 0.54 1.4 0.53 1.28 0.68 1.57 VO... 0.50 1.54 0.54 1.38 0.60 1.47 Gossypium herbaceum. Auburn, Ala., Oct. 13, 1911. Lioyp: LEAF WATER IN GOSSYPIUM TABLE VIII Dry weight and weight of water per 100 sq. cm. of leaf at the close of 0.83 inch precipitation during the © preceding 48 hours. Series 10, material taken by a leaf punch from alternate sides of the same leaves ° on ten plants, not more than three circles being taken at one time from a given leaf. aoe leaves (second an taken near the top of plant; series 11, young leaves of the same kind select rial ise as in (1) above ete old leaves near the top of the plants. indiscriminately from ‘“‘lot 2, 1911 "s series 12, mate 6.30 Ir.00 15.00 Series | Dry wt Water Dry wt Water Dry wt Water 10 0.46 1.58 0.53 1.48 0.56 I.51 II 0.45 at 0.47 1.43 0.49 1.36 12 0.70 1.76 0.68 1.81 0.70 1.69 Ave... 0.55 1.63 0.56 1.57 0.58 1.52 TABLE IX Gossypium herbaceum. Auburn, Ala., Sept. 16-17, sae Average transverse dimensions of stomata of the upper and lower sides of the ments are given in parentheses. xtreme measure- TABLE X herbaceum. Auburn, Ala., cia 30, 1ord. Ax Hour diameter, micra 0.7 (4) 6 é ; 0.6 (3) “~ 0.2 (2) 0.2 (3 22 0 (3 re) : oO(1 sie o(1 ‘ o (3) Meee 2", Gossypium in field plants. Averages in bold face 00 6 = = oo 6.20 o = oo 0-2 0 6.30 o-3" ° (0.9) oO ssn o-2” aay oat (0.6) 0-4 = 6. bik. 3 55 o-4" (2) Hour a ‘ 0 (2) : 0.5 os 0.5 (1 : 1.0 (1) 8 1.5 (5) 1.2 (4) is 1.5 (5) ¥ 1.5 oo 2 1.5 (6) 1.2 (4) Stomatal measurements 10.00 Lioyp: LEAF WATER IN GOSSYPIUM 25 o-4(6) o-3(5) (2) 0-3(4) o-(3) 0-(4)’ 0-4(6)’ o~(4) 0-3(6) 0-4 o-5(8)' 0-6 °"> o-(4) 0-3(6) 7) 2) 0-4 0-4(6) toutes ca copay ee 0-4" (0)-5(9)’ 0-4 0-6(8) 0-6(8) 0-4(6) 0-8(12) o-— 0-4(8)’ 4-12 o-4(6)’ 0-8 (0)4-8 0-4(8) 3-8 0-6(8) 0-8(10)’ o~6(10) 0-8 (0)-6(10) 0-10’ (0)-6(10) ona(8) 0-308) 0 o (4) (4-5) (2.5) o-4(8) 0-3(4) (2) o-6(8)’ o-1(4) ~*~ o(t) o(z) ety’ 6G) 0-4 0-2(4) 0-3(6)" 0-3(4) es) o 0 0’ 0 (0) o-10(12) 0(3)_ 0-6 * o2(4) (2) o-8(rt0) o-1(2) o-(8) " (0)2~4(8) (3) PO) o-r 0-2(4) 0(3)' (0)3 us (0)2-4 o-T waa & ae (0)2-8 0-6(8) (0)4-8(10)’ o~8(12)’ 0-8(12)’ 0-6(10) Shade (4-5) o-6(8) o-8(10) (0)-5(7) a> * o-4(6) (4.5) oe Soames: Sunlight sieges Large leaf; sunlight. Two young leaves of same age, twig, and exposure; apex of plant. Two mature leaves on same twig. Two old leaves, apparently wilted. Two small young leaves near base of plant. Two small leaves near base of plant. 26 Lioyp: LEAF .WATER IN GossyPIUM 0(3) 0-2(4) Two very hei leaves at top of plant, same o 0(2) (0.4) light exp nares o(2) _0(2) (0) Two mature leaves tops of neighboring : 0(2)" o(2-6) plants, with same exposures. TABLE XI Gossypium herbaceum. Sept. 27, 1911. Transpiration during wilting. Top of chief shoot, with 3 mature leaves and 5 young leaves. Cut end vaselined. Weighed doors in bright diffused light. Hour Weight, grams | Loss | Rate per minute 7.22 19.3 | 27 19.06 0.24 | .048 32 18.89 0.17 034 38 18.70 0.19 | .032 44 18.56 0.14 -024 50 18.46 0.10 O17 18.36 0.10 O17 8.02 18.26 0.10 O17 07 18.16 | 0.10 | 020 16 18.07 0.09 -OOT 31 17.89 0.18 -0012 40 17.76 0.13 | oorI4 17.65 O.II oOoIt 9.02 17.51 0.14 0012 17.41 0.10 -OOIT 27 17.23 0.18 oor2 his paper is a partial report of work done as an Adams Fund project at ay Aitoies agricultural experiment station. McGit_ UNIVERSITY. On a possible relationship between the structural peculiarities of normal and teratological fruits of Passiflora gracilis and some physico-chemical properties of their expressed juices Ross AIKEN GORTNER and J. ARTHUR HARRIS We present here an account of first studies on the physical properties of the juice expressed from normal and teratological plant organs. MATERIALS The plants furnishing the fruits used were vigorous and normal in growth but were transplanted from the greenhouse to the field too late to attain the largest size or to produce the maximum number of mature fruits before the oncoming of cold weather. We were, therefore, somewhat limited in amount of material, but altogether 10,929 fruits were dissected in obtaining abnormals and normals for checks. Not many more could have been worked over with the facilities available. The fruits for which adequate samples could be secured fell into the following classes :* (a) Normal fruits. Six external sutures, three placentae, no prolification. A sample of such fruits, collected as described below, served as a check for each of the samples of abnormals. (b) Seven external sutures, three placentae, no prolification; 2 samples, I, 2. (c) Eight external sutures, three placentae, no prolification; I sample, 3. (d) Eight external sutures, four placentae, no prolification; 9 samples, 4-12. (e) Six external sutures, three placentae, slight and generally abortive prolification of three external carpels; 1 sample, 13. * For a general account of prolification in Passiflora see a paper by J. A. Harris, Prolification of the Fruit in Capsicum and Passiflora, Ann. Rep. Missouri Bot. Gard. 17: 133-145. 1906. 27 28 GORTNER AND Harris: Fruirs oF PASSIFLORA GRACILIS (f) Six external sutures, three placentae, slight and generally abortive prolification of four external carpels; 4 samples, 14-17. (g) Eight external sutures, four placentae, slight and generally abortive prolification of three external carpels; 1 sample, 18. (h) Eight external sutures, four placentae, slight and generally abortive prolification of four external carpels; 2 samples, 19, 20. (i) Six external sutures, three placentae, large living prolifica- tion of four external carpels; 2 samples, 21, 22. (7) Eight external sutures, four placentae, large living pro- lification of four external carpels; I sample, 23. METHODS The rarity of the abnormal fruits is a source of great difficulty in the collection of the samples. As the fruits were dissected, each abnormal was placed in a dish provided with a ground glass cover and containing bibulous paper saturated with water in order to prevent, as far.as possible, any drying out of the fruits. A normal to serve as a check was at once opened and placed in a similar receptacle. These two were kept side by side until it was necessary or convenient to combine abnormalities belonging to the same type and their check fruits in a pair of larger moist chambers. As soon as a sample of any type conveniently large for the extraction of juice was secured, the collection of another general sample and check was begun. Thus, while the different samples of abnormals came from various plants and were necessarily held for varying lengths of time, the fruits of each sample and of the check with which it was compared were drawn in equal numbers, and at the same time, from the same individual plants and recewed parallel treatment in every detail. . The juice was secured by means of a large “‘beef-juice’’ press. It was filtered clear through a dry barium filter (S. & S. No. 589), and the depression of the freezing point (A) determined in ‘the well-known Beckmann apparatus. The specific gravity of the sap at 20° C. was found by weighing in a pycnometer holding 5.2405 grams of water at 20° C. The concentration of dissolved sub- stances was determined by evaporating a measured volume (10- 15 c.c.) to dryness in glass weighing bottles, first in a water oven GORTNER AND Harris: FRUITS OF PASSIFLORA GRACILIS 29 at the temperature of boiling water and then completing the desiccation by heating to 105° C. for several hours.* Depression of the freezing point, A, specific gravity, d(20°/20°), and total solids in 100 c.c. of juice, s, are the only direct deter- minations entered in the table. From these, osmotic pressure in atmospheres, P, and the average molecular weight of the sub- stances dissolved in the plant sap, M, have been calculated by the formulae P = 12.060A — 0.0214? M = 1890 erage (A 1OG: = d | *It may not be amiss to consider the accuracy of our numerical data. Inas- much as we were not concerned with the absolute depression of the freezing point, but only with the sign and amount of variation between normal and abnormal juices, we have not taken all of the precautions which would be necessary were the absolute values desired. We have found that the degree of pressing the fruits has no influence on the relative values, providing that the normal is treated in the same manner as the abnormal. For example: a sample was pressed lightly and the freezing point of the expressed juice determined; then the residue was pressed so that a considerable amount of juice was again obtained, and this juice was added to that from the first pressing. The absolute values are much different but the relative value remains approximately the same. | Abnormal | Normal Difference ' Light pressing. .. .. | A =0.677° C. A = 0.603° C. | + 0.074° C. eave eteaiee. A A =0.705° C. | A = 0.628° C. + 0.077° C. A constant relative value is also obtained when large samples are divided into two portions and the juice expressed from the second portion after an interval of four days, showing that holding the fruits for several days does not affect the relative values. The absolute and relative values of a pair of samples treated in this manner follow: | Abnormal | Normal | Difference . Set aoe os i Cc First value.......- A = 0.587° C. | A = 0.618° C, | 0.031° C. Four days later.... | A =0.677° C. A = 0.713° C. — 0.036° C, We have not corrected the depression of the freezing point for the amount of supercooling before freezing, for the same reason. That is, we are concerned only ith a relative value. We have had occasion in several instances to repeat a deter- mination, sometimes as much as twenty-four hours later, on the same sample of ie ae ee roa 3 vary from th igi oe | 7 at juice, and in ty ; J y one < See : ‘ Weaw bss 30 GOoRTNER AND HarrIS: FRUITS OF PASSIFLORA GRACILIS The arguments in this paper are in all cases based upon the comparison of the samples of abnormal fruits with their checks. The differences averaged and discussed in the text are in all cases taken as abnormal Jess control, the sign of the difference being positive when there is a greater depression of freezing point, higher osmotic pressure, or higher average molecular weight, in the abnormal fruits. ANALYSIS OF DATA The large table gives the essential constants for the various lots of fruits. The section to the left contains the data for the samples of abnormal fruits, that to the right those for the checks. Consider first the properties of the juice in fruits differing only in the structure of the wall. Series (b) and (c) may be regarded as transitions between the trimerous and tetramerous fruits. In two cases A is higher and in one lower sien 3 in the control samples. The negative difference is only —0.012° and is but twice or thrice the estimated experimental error. The mean difference in depression of the freezing point is + 0.035°. The differences in pressure in atmospheres, P, have the same sign as the differences in depression. In one the difference is — 0.144, in the other two it takes the more substantial values of + 0.638 and + 1.143; the mean of the three is + 0.546. The average molecular weight is in all three cases lower in the abnormal fruits, — 7.41, —4.46 and — 9.02 being the values. The tetramerous fruits (eight external sutures, four placentae, no prolification), class (d), are represented by nine samples, divided as follows: eee as | ge Steen Site Mean value of 5 differences fs WIND 2 OREN rio. ara 3 + 0.002° C, Mea tite | 2 | 7 2.81 Pie ra eas es 6 3 - 0.024 Combining (0), (c) and (d), we have, with headings as above, + 0.010° C. > ; : F : * : . * * : * ; : : : : wo | 4 10 } 8 4 TABLE I PHYSICO-CHEMICAL CONSTANTS FOR JUICE OF NORMAL AND ABNORMAL PASSIFLORA FRUITS Values for abnormal! fruits Val £ i 4 a. Fs M A ae | M Okc ey Depression of 20° Solids in 04 cc Molecular [Osmotic pressure) Depressionof | = 20° ~—_— Solidsin rooc.c. | Molecular | pressure in 4 freezing point | Specific gravity of ju weight in atmospheres | freezing point | Specific gravity of juice | weight [a+ |atmospheres te 0.503° 1.0161 3-042 115.95 6.061 0.515 1.0169 3-349 124.97 | 6.205 2 0.593° I.01Q1 3.466 112.23 7.144 0.540° 1.0173 3.282 116.69 | 6.506 3 0.633° 1.0195 3.600 109.26 7.626 0.568° I.0180 3.448 116.67 6.4 4 0.405° 1.0122 2.209 104.10 4.881 0.358° 1.0114 2.162 115.35 4.315 5. 0.503° 1.0163 3.065 116.86 6.061 0.428° 1.0129 2.356 105.14 5.157 6 0.525° I 2.630 905-54 6.326 0.513° 1.0171 2.720 101.21 6.181 1 0.435° 1.0146 2.282 99 5.242 0.511° 1.0168 2.747 102.70 6.157 8 0.587° 1.0187 3.381 110.53 7.071 0.618° 1.0202 3.5607 110.79 7.445 9 0.558° 1.0175 3-102 106.52 6.729 0.543° 1.0169 3.233 114.27 6.542 10 0.585° 1.0181 3-154 103.29 7.047 0.581° 1.0183 3.001 101.81 7.000 Ir 0.567° T.0161 3.478 118.13 6.831 0.608° 1.0195 3-907 123.87 7.324 12 539° 1.0169 3-204 114.10 6.494 0.527 1.0167 3.270 119.18 6.349 13 0.673° 2 4.237 121.45 8.107 0.614° 1.0184 3.526 110.38 1-397 14. 0.573° 1.0188 2.857 95.12 6.903 0.550° 1.0186 2.798 97.07 6.627 15 0.705° 4.290 117.18 8.492 0.628° 1.0212 3.652 111.62 7.505 16 0,603° 1.0200 3.540 112.70 7.264 0.613° 1.0184 3-212 100.42 7.384 39:1 . .0.869° 1.01 4. 125.10 6.855 0.554° 1.0183 3-523 | 322.26 6.674 1A 0.708° 1.0189 3.801 106.01 8.528 0.683° 1.0202 4.172 | 117.95 8.227 19 0.558° 1.0195 _ —— 6.723 0.585° 1.0204 _—- —=. | 7048 20 0.643 1.0203 3-729 III.51 7.746 0.597° 1.0185 3-354 | 107.79 | 7-192 ar 0.455° 1.0149 —— 5.483 0.548° 1.0180 --— | 6.603 22 0.577° 1.0180 3.584 Litt £3652 6.952 0.533° 1.0170 3.598 | 130.07 | 6.422 23 0.563° 1.0178 3.281 l. «eae.88 6.783 0.599° 1.0198 3-754 | 120.60 7.216 :SIMUVE] GNV YANLYOD STTIOVAD VUOTAISSVG AO SLINAY T€ 32 GORTNER AND Harris: FRUITS OF PASSIFLORA GRACILIS Or disregarding the presence of prolification and adding to these the other fruits which are abnormal (tetramerous as con- trasted with trimerous) in the organization of their ovary wall (i. e. classes (g), (h), and (7)), we get Br eee os 10 6 | + 0.008° C. J: Gere ir sy oR eee cm 3 12 — 4.21 GE ORS Ps Io 5 + 0.151 Apparently, therefore, those fruits which are tetramerous or which show transitions between the trimerous and tetramerous condition, show a greater depression of the freezing point (and consequently a higher osmotic pressure) of their juice and a lower average molecular weightt than the trimerous ones. But the differences are so very slight, and the difficulties and sources of possible error are so many, that further studies will be required to put this conclusion on a sound basis. Turn now from the classification of the fruits according to the characteristics of the ovary wall to a consideration of the question of prolification. Here classes (6), (c), (d) may be left entirely out of account. First dividing the fruits that have prolifications into the classes, trimerous and tetramerous, with respect to the char- acteristics of the ovary wall, without regard to the size or the structure of the included body, we have the following: For trimerous fruits (classes (e), (f) and (i)), Ae Mapes Nesatica aa Mean value of — 5: rie differences es ee eee Ce cue oly 5 2 + 0.016° C lagi eis nko 4 > + 3.21 Wie scnete Binie gtk es 5 I + 0.417 For tetramerous fruits (classes (g), (h) and ( j), * Depression of freezing point is available in one series in which the eee was determined. omit the difference of the freezing points would indicate, inasmuch as colloids do not croleacdhae cee msuence the amount of total solids and through this the average weight. not cacalated, te the cent Point lo avaliable fr one case for which Af and F were not calculated, for the reason noted abo GORTNER AND Harris: FRvuitrS OF PASSIFLORA GRACILIS 33 fa\, PeMinnen Mase eg Eee 2 2 + 0.0029 C. g." Saree eg eet! I 2 ged f Brie our reat Sy att ) I + 0.141 } On combining trimerous and tetramerous (as above) without regard to the character of the proliferous body we have: + o.011° C, Boo ae 7 4 | Much | 5 4 | 0.25 Poa 7 + 0.325 Apparently the proliferous fruits tend to show a greater de- pression of the freezing point and a higher osmotic pressure in their expressed juices than the normal checks with which they were compared. This is true for trimerous and tetramerous fruits alone and for the two classes taken together. Considering the fruits merely as normal and abnormal, we note that these results are in good agreement with those for fruits abnormal only in respect to the structure of the wall. For the trimerous fruits the results for M are, however, not in accord with those for fruits abnormal with respect to the wall only, M being higher in the abnormals than in the controls. For the tetramerous fruits the average molecular weight is again lower than in the controls. Taking both trimerous and tetramerous fruits (with prolifications) together, we find practically no difference between the average molecular weight of the fruits containing supernumerary carpels and that of their controls. Let us now simply classify the fruits as normal and abnormal and compare the 23 samples abnormal in some character with their controls, which are normal in all regards. We have: Positive difference | Negative difference | Mean value of differences Pie dsagslled scien OMAR RE eee Bey see te 15 8 | + 0.0107° C. Mee 7 14 | . — 2.001 Pe oe 15 6 | 0.2275 These results emphasize the conclusions drawn from the individual classes of fruits. * Available for one case where Mand P were not calculated. + Available for two cases where M and P were not calculated. 34 GORTNER AND HARRIS: FRUITS OF PASSIFLORA GRACILIS DISCUSSION AND CONCLUSIONS From the determination of the depression of the freezing point, the specific gravity, and the total solids in the expressed juice of 23 samples of abnormal fruits of Passiflora gracilis and a like number of controls, we are led to the following conclusions: Our experiments indicate that the juice of abnormal fruits has a higher osmotic pressure (greater depression of the freezing point) than that of normals. This is true whether the abnormality be a meristic variation in the fruit wall—i. e. an increase in the number of external sutures or of the number of placentae over the normal condition—or the production of an entirely new structure in the form of an included whorl or whorls of accessory carpels springing from the floor of the fruit (prolification of the fruit).* The average molecular weight of the substances in solution in the plant sap is, apparently, lower in the abnormal fruits, but this is less consistently true for the various classes of structural aberrations recognized. While the findings are fairly consistent throughout, it must be remembered that the problem is surrounded with many difficulties. We have no desire to be dogmatic concerning these conclusions, realizing that a wider series of material than we could possibly obtain is desirable,t and that many questions remain to be in- vestigated.{ Furthermore, it is clear that the whole problem of the nature of the relationship between the structure of the fruits and the properties of the juice remains to be worked out. We only claim to have demonstrated that the physico-chemical properties of the plant sap deserve consideration as a first step in the analysis of the factors involved in morphological variations of the fruit. STATION FOR EXPERIMENTAL EVOLUTION, CARNEGIE INSTITUTION OF WASHINGTON, * In € forthoctaing meee on the morphology of normal and teratological fruits of Passiflora gracilis one of us will show that the oe cf prolification is to some degree correlated with abnormalities of the ovary wal We hope another year to obtain a strain of plants vias a higher percentage of abnormalities or at least to obtain far larger series of dissections {t In particular it will be of great interest to work out in detail the relationship Dewees the properties of the juice of the ovary wall and that of the contained in the case of fruits showing prolifications. Some beginning has been made carpe on this —— but as yet our data are too few to justify the discussion of this and several oth INDEX TO AMERICAN BOTANICAL LITERATURE (1912) aim of this Index is to include all current botanical literature written - Americans, published in America, or based upon American material ; the word Am ica bei cing used in the broadest se Reviews, and papers that age exclusively to forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and no attempt is made to index the literature of bacterio ec An occasional exception is made in favor of some paper appearing in an American periodical which is devoted wholly to botany. Reprints are not mentioned unless they differ from the original in some important particular. If users of the Index will call the veges ane of the editor to errors or omissions, their kindness will be appreciated. This Index is reprinted monthly on cards, ai furnished in this form to subscribers at the rate of one cent for each card. Selections of cards are not permitted ; each subscriber must take all cards published pepe the term of his subscription, Corre- spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club. , Ames, O. Notes on Philippine orchids with descriptions of new species —V: The genus Bulbophyllum in the Philippine Islands. Philip. Jour. Sci. 7: (Bot.) 125-143. 2S 1912. Seventeen new species descri Anderson, P. J., & Anderson, H. W. The chestnut blight fungus and a related saprophyte. Phytopathology 2: 204-210. O 1912. Discusses the Connellsville fungus, for which the name Endothia virginiana is proposed, and concludes that we have in our territory (1) Endothia radicalis (Schw.) Fr., (2) the true blight, E. parasitica (Murrill), and (3) E. virginiana Andrews, F. M. Some variations in plants. Proc. Indiana Acad. Sci. IQII: 279-281.. I9I2. Berger, A. Opuntia tomentella A. Berger spec. nov. Monats. Kak- teenk. 22: 147, 148. 15 O 1912. Banker, H. J. Type studies in the Hydnaceae—IIl. The genus Sieccherinum. Mycologia 4: 309-318. 23 N 1912. Steccherinum Peckii and S. basi-badium spp. nov. are described. Bessey, C. E. Outlines of plant phyla 1-20. Lincoln. 28 S 1912. Third Edition. Bioletti, F. T., & Cruess, W. V. Enological investigations. Calif. Agr. Exp. Sta. Bull. 230: 23-118. f. 7-8. Au 1912. Includes a list of fungi found in must of grapes. 35 36 INDEX TO AMERICAN BOTANICAL LITERATURE Bitter G. Solana nova vel minus cognita—III. Repert. Sp. Nov. 11 201-240, 200 10912. Includes 32 new species in Solanum. Blumer, J.C. The Euphorbias of Tucson and vicinity. Muhlenbergia 8: 97-102. 31 O 1912 Bédeker, F. Mamillaria Verhaertiana Boédeker spec. nov. Monats. Kakteenk. 22: 152-155. 15 O 1912. [lIllust.] Bovell, J. R. The use of entomogenous fungi on scalg insects in Barbados. West Ind. Bull. 12: 399-402. 18S 1912. Brooks, C., & DeMeritt, M. Apple leaf spot. Phytopathology 2: 181-190. pl. 17. O 1912. A disease caused by Sphaeropsis malorum. Brown, W.H. The relation of Rafflesia manillana to its host. Philip. Jour. Sci. 7: (Bot.) 209-226. pl. 12-21. S$ 1912. Cannon, W. A. Structural relations in xenoparasitism. Am. Nat. 46: 675-681. N 1912. [Illust.] Chamberlain, C. J. A round-the-world botanical excursion. Pop. Sci. Mo. 81: 417-433. f. I-10. N 1912. Clark, E. D. The original significance of starch. Orig. Comm, Eighth Internat. Cong. Appl. Chem. 19: 55-69. 1912. Clements, F. E., Rosendahl, C. O., & Butters, F. K. Minnesota trees and shrubs. Rep. Bot. Surv. Minnesota 9: ix-xxi + 11-314. 15 Au 1912. Clute, W. N. A _ problematical fern (Gymnogramma lanceolata). Fern Bull. 20: 43-45. [O] 1912. [Illust.] [Clute, W. N.] Pteridiographia. Fern Bull. 20; 56-60. [O] 1912. Includes notes on (1) new fern pest, (2) walking fern and lime, (3) apogamy in Pellaea, (4) Lycopodium lucidulum porophilum, (5) affinities of Taenitis, and (6 sporophyll zones. Clute, W. N. Rare forms of fernworts—XXII. Fern Bull. 20: 49-52. [O] 1912. [Ilust.] Conklin, G.H. The Hepaticae of the sixth edition at Gray’s Manual compared with the exchange list. Bryologist 15: 88-91. N 1912. Coker, W.C. Achlya DeBaryana Humphrey and the prolifera group. Mycologia 4: 319-324. pl. 78. 23 N 1912. Coker, W. C. Achilya glomerata sp. nov. pl. 79. 23 N 1912. Cook, O. F. Results of cotton experiments in 1911. U.S. Dept. Agr. Plant Ind. Circ. 96: 3-21. 17 Jl 1912 Mycologia 4: 325, 326. Deam, C. C. Additions to the flora of the lower Wabash valley, by Dr. J. Schneck. Proc. Indiana Acad. Sci. 1911: 365-360. 1912. INDEX TO AMERICAN BOTANICAL LITERATURE 37 Deam, C. C. Plants new or rare in Indiana. Proc. Indiana Acad. Sci. 1911: 371-374. 1912. ' Detmers, F. An ecological study of Buckeye Lake. Proc. Ohio Acad. Sci. 5: 5-138. pl. 1-12 + f. 1-31. My 1912. The annotated list of plants includes 19 species of fungi. Detwiler, S. B. Some benefits of the chestnut blight. Forest Leaves 13: 162-165. O 1912. Dodge, R. Further notes on variation in Botrychium ramosum. Fern Bull. 20: 48, 49. [O] 1912. East, E. M. The mendelian notation as a description of physiological facts. Am. Nat. 46: 633-655. N 1912. Emerson, R. A. The inheritance of certain forms of chlorophyll reduc- tion in corn leaves. Ann. Rep. Nebraska Agr. Exp. Sta. 25: 89-124. 1 F 1912. Fawcett, H. S. Citrus scab, Cladosporium Citri Massee. Monthly Bull. State Comm. Hort. Calif. 1: 833-842. f. 253-260. O 1912. Fawcett, H. S. Gum diseases in citrus trees. Monthly Bull. State Comm. Hort. Calif. 1: 147-156. f. 49-53. Ap 1912. Fisher, M. L. Report of the work in corn Simian tn Proc. Indiana Acad. Sci. 1911: 283-284. 1912. French, G. T. Seed tests made at the station during 1910. Ann, Rep. N. Y. Exp. Sta. 30: 69-80. 1912. Gates, F.C. The relation of snow cover to winter killing in Chamae- daphne calyculata. Torreya 12: 257-262. f. 1-3. 10N 1912. Greene, E. L. Certain asclepiads. Leaflets 2: 229-333. 22 O 1912. Greene, E. L. Certain western roses. Leaflets 2: 254-260. 22 O 1912; 261-266. 6N eho Twenty-two new species descri Greene, E. L. Earlier history ot our dogbanes—I. Leaflets 2: 241- GAO. Qa 4) 2612, Greene, E. L. A handful of vetches. Leaflets 2: 267-270. 22 O 1912. Greene, E.L. Miscellaneous specific types—VI. Leaflets 2: 270-272. IgI2 ficdudes new species in a (2), Claytonia (1), Tridophyllum (2) and Sisy- rinchium (1). Greene, E. L. New species of Cicuta. Leaflets 2: ateaee 22 O 1912. Eight new species are described ‘ Greene, E. L. Novitates EROS SEO EEC, § Repert. Sp. Nov. Ir: 108-111. 20 Jl 1912. Includes descriptions of 7 new species of Cercis. 38 INDEX TO AMERICAN BOTANICAL LITERATURE Greene, E. L. Some new lupines. Leaflets 2: 233-236. 22 O 1912. Six new species described Greene, E. L. Some Caictconitns maples. Leaflets 2: 248-254. 220 1912. Greene, E. L. Three new Rhamni. Leaflets 2: 266, 267. 6 N 1912. Grimms, E. J. New and notable members of the Indiama flora. Proc. Indiana Acad. Sci. 1911: 285-289. 1912. [Grout, A. J.] Photographing mosses. Bryologist 15: 97. pl. 4. N 1912. Bryum caespiticium, Hall, J. G. Monochaetia Desmazierit. Mycologia 4: 330, 331. N 1912. A note on the identity of the fungus causing the large leaf spot of chestnut. Halsted, B. D. Shade-induced uprightness in the seaside spurge. Torreya 12: 262. 10N 1912. Halsted, B. D. ° The elongation of the hypocotyl. New Jersey Agr. Exp. Sta. Bull. 245: 3-32. pl. I-12. 25 My 1912. A preliminary study. Harris, J. A. A first study of the influence of the starvation of the ascendants upon the characteristics of the descendants—II. Am. Nat. 46: 656-674. N 1912. Harshbe ger, J. W. A classification of the departments of botany and an arrangement of material based thereon. Science II. 36: 521-525. 18 O 1912. Haynes, C. C. Helpful literature for students of North American Hepaticae. Bryologist 15: 91-93. N 1912. Heald, F. D., & Lewis, I. M. A blight of the mesquite. Trans. Am. Micros. Soc. 31: 5-9. pl. 1. Ja 1912. Heller, A.A. The North American lupines—VIII. Muhlenbergia 8: 103-107. f. 15-17. 310 1912. Includes Lupinus apertus and L. mollis spp. nov. Hesler, L.R. The New York apple tree canker. Proc. Indiana Acad. Sci. 1911: 325-339. f. 1-7. 1912. Davis, B.M. Was Lamarck’s evening primrose (Oenothera Lamarckiana Seringe) a form of Oenothera grandiflora Solander? Bull. Torrey Club 39: 519-533. pl. 37-39. 18 N 1912. Hill, E. J. The fern flora of Illinois. Fern Bull. 20: 33-43. [O] 1912. Hollick, A. Additions to the paleobotany of the Cretaceous formation on Long Island. No. III. Bull. N. Y. Bot. Gard. 8: 154-170. pl. 162-170. 23 N 19%2. Includes Embothriopsis presagita gen. et sp. nov., Laurophyllum ocoteaeoides, Cassia insularis, Myrtophyllum sapindoides, and Ligustrum subtile spp. nov. INDEX TO AMERICAN BOTANICAL LITERATURE 39 Hopkins, C. G. Plant food in relation to soil fertility. Science II. 36: 616-622. 8 N 1912. Horne, W. T. Fungous root rot. Monthly Bull. State Comm. Hort. Calif. 1: 216-225. f. 83-91. My 1912. Armillaria mellea the cause of root rot. Hotson, J. W. Culture studies of fungi producing bulbils and similar propagative bodies. Proc. Am. Acad. Arts & Sci. 48: 227-306. pl. i-12.: O 1912; Includes new species in Cubomia (1), Papulospora (9) and Melampsora (3). Jeffrey, E. C. The history, comparative anatomy, and evolution of the Araucarioxylon type. Proc. Am. Acad. Arts & Sci. 48: 531- 571. pl. 1-8. N 1912. Johnson, A. G. The unattached aecial forms of plant-rusts in North America. Proc. Indiana Acad. Sci. 1911: 375-411. 1912. Johnson, E. C. The smuts of wheat, oats, barley, and corn. U. S. Dept. Agr. Farm. Bull. 507: 3-32. f. 7-12. 8 O 1912 Kingman, C. C. A list of mosses collected in southern California. Bryologist 15: 93-95. N 1912. Lantis, V. Development of the microsporangia and microspores of Abutilon Theophrasti. Bot. Gaz. 54: 330-335. f. I-12. 15 O 1912. Lew's, I. M. A bacterial canker of plum twigs. Trans. Am. Micros, Soc. 31: 145-149. pl. 14. Jl oe A disease caused by Pseudomonas Pru Lindau, G. Einige neue ‘Aastha Repert. Sp. Nov. 11: 122~124. 20 Ji 1912. Includes Geissomeria a Streblacanthus cordatus, and Chaetochlamys pana- mensis spp. nov. from Pana: Lyon, H.L. Tliau, an ea cane disease. R-p. Exp. Sta. Hawaiian Sugar Planters’ Assoc. Bull. 11: 5-28. pl. 1 +f. I-10. S 1912. Includes Melanconium Iliau the imperfect stage of Gnomonia Iliau sp. nov. and notes on M. Sacchari. McLachlan, A. The branching habits of Egyptian cotton. U. S. Dept. Agr. Plant Ind. Bull. 249: 5-28. pl. 1-3 +f. 1. 20S 1912. Mattei, G. E. Osservazioni biologiche sopra alcune Cactaceae. Mal- pighia 24: 341-345. 1912. Melhus, I. E. Culturing o parasitic fungi on the living host. Phyto- pathology 2: 197-203. pl. 20 + f. 1,2. O 1912. ill, E. D. Nomenclatural and systematic notes on the flora of Manila. Jour. Philip. Sci. 7: (Bot.) 227-251. S 1912. Includes Fimbristylis corniculata, Tabernaemoniana subglobosa, Blumea tenera, Limnophila manilensis, and Staurogyne rivularis spp. nov. Merrill, E. D. Notes on the flora of Manila with special reference to 40 INDEX TO AMERICAN BOTANICAL LITERATURE the introduced element. Philip. Jour. Sci. 7: (Bot.) 145-208. 2S 1912. Miller, F. A. The improvement of medicinal plants. Proc. Indiana Acad. Sci. 1911: 309-320. 1912. [Illust.] Moore, G. T. Microorganisms of the soil. Science II. 36: 609-616. N 1912. Murrill,W.A. The Agaricaceae of the Pacific Coast—III. Mycologia 4: 294-308. pl. 77. 23 N 1912. New species are described in Agaricus (9), Stropharia (2), Drosophila (5), and Gomphidius (1). Murrill, W. A. Illustrations of fungi—XII. Mycologia 4: 289-293. pl. 74. 23 N 1912. The following species of Russula are described and illustrated: R. sericeonitens Kaufiman, R. Mariae Peck, R. emetica Fries, R. sulcatipes Murrill sp. nov., R. obscura Romell, R. uncialis Peck, R. foetens, Pers., and R. rubriochracea Murrill sp. nov. Murrill, W. A. New combinations for tropical agarics. abvectita 4: 331, 332. °- 23 NIGt2. Murrill, W. A. The Polyporaceae of Mexico. Bull. N. Y. Bot. Gard. 8: 137-153. 23 N 1912. Includes new species in Coriolopsis (4), Coriolus (6), Favolus (1), Grifola (1), Hexagona (3), Trametes (2), Tyromyces (1), Ganoderma (2), Daedalea (1), Pyropoly- porus (1), Gleophyllum (1), and Lenzites (1). Murrill, W. A. Species of Hydnaceae appear to be scarce on the Pacific coast, as elsewhere. Mycologia 4: 330. 23 N 1912. Short notes on five species collected by the author Orton, C. R. The prevalence and prevention of stinking smut in Indiana. Proc. Indiana Acad. Sci. 1911: 343-346. 1912. Osner, G. A. Diseases of ginseng caused by sclerotinias. Proc. Indiana Acad. Sci. rorr: 355-364. f. I-6. 1912. Osterhout, W. J. V. Some chemical relations of plant and soil. Sci- ence II. 36: 571-576. 1N 1912. Owens, C. E. A monograph of the common Indiana species of Hy- poxylon. Proc. Indiana Acad. Sci. r911: 291-308. f. 1-16. 1912. Pace, L. Parnassia and some allied genera. Bot. Gaz. 54: 306-329. pl. 14-17. 15 O 1912. Petry, E. J. Nutrients in green shoots of trees. Proc. Indiana Acad. Sci. 1911: 321-324. 1912. Piper, C. V., & McKee, R. Vetches. U.S. Dept. Agr. Farm. Bull. 515: 5-28. f. 1-10. 11 N 1912. ; Prescott, A. The tall spleenworts. Fern Bull. 20: 46, 47. [O] 1912. Prince, S.F. Notes on various ferns. Fern Bull. 20: 52, 53. [O] 1912. Pulle, A. Neue Beitrage zur Flora Surinams—III. Rec. Trav. Bot. Néerlandais 9: 125-168. pl. 2, 3. 1912. INDEX TO AMERICAN BOTANICAL LITERATURE 41 Quehl, L. Bemerkungen iiber einige Arten von Mamillarien aus der Untergattung Coryphantha Engelm., Reihe Avulacothele Lem, Monats. Kakteenk. 22: 115-118. 15 Au 1912. Quehl, L. Mamillaria macrothele Mart. Monats. Kakteenk. 22: 145-147. 15 O 1912. Roberts, E.W. The modern theory of the cell as a complex of organized units. Trans. Am. Micros. Soc. 31: 85-113. pl. 6-13 +f. A-K. Ap 1912. Robinson, C.B. Polycodium. Bull. Torrey Club 39: 549-559. 18 N Rolfe, R. A. Eriopsis Helenae. Curt. Bot. Mag. IV. 8: pl. 8462. N 1912, A plant from Peru. Rosenbaum, J. Infection experiments with Thielavia basicola on ginseng. Phytopathology 2: 191-196. pl. 18, 19. O 1912. Rusby, H.H. Newspecies from Bolivia, collected by R. S. Williams—2. Bull. N. Y. Bot. Gard. 8: 89-135. 23 N 1912. One hundred and fifteen new species are described. Schreiner, O. Organic constituents of soils. Science II. 36: 577-587. 1 N 1912. Schreiner, O., & Skinner, J. J. The effect of guanidin on plants. Bull. Torrey Club 39: 535-548. f. 1-6. 18 N 1912. Seaver, F. J. Ancient and modern views regarding the relation of taxonomy to other phases of botanical work. Torreya 12: 262- 264. I10N 1912. Shear, C. L. The chestnut blight fungus. Phytopathology 2: 211, 213.0 1912. Sheldon, J. L. Additions to the recorded mosses of West Virginia. Bryologist 15: 95-97. N 1912. Smith, E. F. Isolation of pathogenic potato bacteria: A question of priority. Phytopathology 2: 213, 214. O 1912. South, F. W. Further notes on the fungus parasites of scale insects. West Ind. Bull. 12: 403-412. 18S 1912. Starr, A. M. Comparative anatomy of dune plants. Bot. Gaz. 54: 265-305. f. 1-35. 15 O 1912. Stevens, N. E. A palm from the Upper Cretaceous of New Jersey. a eae sei. TV. - oe f. 1-24. N 1912. anchorus sp dase oO. "Iris RENEEN Curt. Bot. Mag. IV. 8: a6 8465. N 1912. Sydow, H. & P. Novae fungorum species—VIII. Ann. Myc. 10: nap Teas 1o Au 1912. Cronartium egenulum from Brazil. 42 INDEX TO AMERICAN BOTANICAL LITERATURE Sydow, H. Fungi exotici exsiccati. Ann. Myc. 10: 351, 352. 10 Au 1912. Includes several American species. Taubenhaus, J. J. Root gall diseases of roses, their cause and methods of control. Gard. Chron. Am. 15: 187, 188. f. 1-3. O 1912. Taubenhaus, J. J. A further study of some Gleosporiums and their relation to a sweet pea disease. Phytopathology 2: 153-160. #l. 16+ f.71. Au 1912, T ylor, N. Recent additions to the local flora garden. Brooklyn Bot. Gard. Record 1: 103, 104. O 1912. Thompson, W. P. Artificial production of aleurone grains. Bot. Gaz. 54: 336-338. f. 7. 15 O 1912. Van Hook, J. M. Indiana fungi—II. Proc. Indiana Acad. Sci. 1911: 347-354. fe 4; S 39R2: Vries, H. de, & Bartlett, H.H. The evening primrose of Dixie Landing, Alabama. Science II. 36: 599-601. 1 N 1912. Weingart, W. Cereus vagans Kath. Brand. Monats. Kakteenk. 22: 135, 136. 15S 1912. Weingart, W. . Pilocereus Houlletii Lem. Monats. Kakteenk. 22: 132-135. 15S 1912. [Illust.] Weingart, W. Der Reif an Cereus trigonus Haw. var. Guatemalensis Eichlam und var. costaricensis Web. Monats. Kakteenk. 22: 129- 131. 15S 1912. Weir, J. R. A Botrytis on conifers in the northwest. Phytopathology 2: 215. O 1912. Wherry, E. T. Silicified wood from the Triassic of Pennsylvania. Proc. Acad. Nat. Sci. Philadelphia 64: 366-372. pl. 3, 4. O 1912. Includes Brachyoxylon pennsylvanica sp. nov. Whetzel, H. H. Baldwin spot or stippin. Proc. N. Y. State Fruit Growers’ Assoc. 11: 28-34. 1912. Whetzel, H.H. The fungous diseases of the peach. Proc. N. Y. State Fruit Growers’ Assoc. 11: 21 1-219, 1912; Wright, C. H. Chamaedorea glaucifolia. urt. Bot. Mag. IV. 8: pl. 8457. O 1912. A plant from South America? - Wright, C. H. Furcraea elegans. Curt. Bot. Mag. IV. 8: pl. 8461. O 1912. : A native of Mexico. Vol. 40 No. 2. BULLETIN OF THE TORREY BOTANICAL CLUB —ee ee FEBRUARY 1913 Studies on the Rocky Mountain flora—XXVIII Per AXEL RYDBERG FABACEAE ‘ Thermopsis ovata (Robinson) Rydb. Thermopsis montana ovata Robinson, Contr. U. S. Nat. Herb. 11: 349. 1906. This differs from T. montana not only in its broader leaflets (the only characters given in the original description) but in its spreading leaves, its large stipules, which in the lower leaves are ovate and very oblique, and in its elongate and lax raceme. It differs from T. xylorrhiza A. Nels. in its lax inflorescence and strictly straight pods. Dr. S. Watson in publishing Lupinus Kingii described the plant as being perennial. This mistake of his led him as well as others astray, for he redescribed the same plant a few years later as an annual under the name L. Sileri. This fact has been called attention to several times and, among other places, in my Flora of Colorado. It is, therefore, surprising that the error should be repeated by Coulter and Nelson in the New Manual of Botany of the Central Rocky Mountains, where the description begins: ‘From a perennial rootstock, dwarf, cespitose,’”’ etc., characters which in no way apply to the type in the Gray Herbarium nor to the duplicates in the herbaria of Columbia University and the United States National Museum. Furthermore, Coulter and [The BuLLETIN for January 1913 (40: 1-42) was issued Feb. 20.] 43 44 RYDBERG: Rocky MouNTAIN FLORA Nelson give as a synonym under the same Lupinus aduncus Greene, which is the same as L. argenteus argophyllus, a plant of different habit. The so-called Lupinus rivularis of the Columbia region and extending into Idaho should be known as L. cytisoides Agardh. Miss Alice Eastwood has seen the type of L. rivularis Dougl., which according to her belongs to an entirely different group from the plant called L. rivularis by Dr. Watson in his revision. The following Lupines are to be added to the flora of the Rocky Mountains: Lupinus nootkatensis Donn has been collected in the Rockies of British Columbia and Alberta, L. plumosus Dougl. in Idaho and Utah, L. minimus Dougl. in Idaho and Alberta, L. lepidus Dougl. in Idaho, L. Cusickii S. Wats. in Idaho and Utah, and L. micensis Jones in Utah. Lupinus lupinus Rydb. sp. nov. Perennial with a woody caudex: stems 3-6 dm. high, densely strigose-canescent, sparingly branched; leaves numerous; stipules subulate, about 1 cm. long; petioles canescent, 5-8 cm. long; leaflets 7-9, oblanceolate, usually flat, 3-6 cm. long, appressed- canescent on both sides, less so above; peduncles about 1 dm. long; raceme 5-10 cm. long; bracts lanceolate, acute, 3-4 mm. long, silvery-pubescent, early deciduous; calyx silvery-pubescent, saccate at the base; upper lip scarcely 3 mm. long, the lower fully 5 mm.; corolla about 1 cm. long, dark blue or purple; banner orbicular, pubescent on the back, usually with a light spot in the center; keel strongly curved, rather broad, ciliate on the margins; pod densely villous, about 3 cm. long, mostly 3-seeded. This is related to L. argentinus, L. aduncus, and L. oreophilus, . but differs from the first in its grayish instead of silvery pubescence of the leaves, which are greener above and not conduplicate, and in its less spurred calyx; from L. aduncus in its broader leaves and the shorter upper lip of its calyx; and from L. oreophilus in its broader leaves and saccate calyx. Along streams and in meadows at an altitude of 2,000-3,000 m. Urau: Western Bear’s Ear, Elk Mountains, Aug. 2, IgII, Rydberg & Garrett 9363 (type, in herb. N. Y. Bot. Gard.); also western slope of La Sal Mountains, July 6, 8595, 8596, and 8600; meadow south of Monticello, July 24, 9167; Head of Dry Wash, RYDBERG: Rocky MOUNTAIN FLORA 45 Abajo Mountains, August 11, 9605; Hammond Canyon, Elk Mountains, August 10, 9583. Lotus TENUIS Waldst. & Kit.; Willd. Enum. Hort. Berol. 797. 1809 Lotus tenuifolius (L.) Reich. Fl. Germ. 506. 1830. Lotus Macbridei A. Nels. Bot. Gaz. 53: 221. 1912. In looking over a collection received in exchange from the University of Wyoming, I found a specimen labeled Lotus Mac- bridet A. Nels. n. sp. To my surprise I found that this was a true Lotus, i. e. not belonging to any of the segregates of Hosackia but of the European type. As it would have been exceedingly strange if a species of Lotus in the restricted sense should be found native in America, I turned to our collection of Old World species of Lotus and found that it is the same as L. tenuifolius (L.) Reich. Before I had time to call Professor Nelson’s attention to the fact, his description appeared in the Botanical Gazette. Trifolium macrocephalum (Pursh) Piper, T. plumosum Dougl., T. eriocephalum Nutt., T. spinulosum Dougl., and cyathiferum Lindl. have been collected in Idaho; T. Rusbyi Greene and Medicago hispida Gaertner (M. denticulata Willd.) in Montana. AcMIsPpon Raf. New Flora 1: 53. 1836 I think that this genus should be restored. The Microlotus section sometimes referred to Hosackia, sometimes to Lotus, is out of place in either genus, and Acmispon is the oldest available generic name. Acmispon americanus (Nutt.) Rydb. Lotus sericeus Pursh, Fl. Am. Sept. 489. 1814. Not L. sericeus DC. 18%. Trigonella americana Nutt. Gen. 2: 120. 1818. Hosackia Purshiana Benth. Bot. Reg. under pl. 1257. 1829. Acmispon sericeum Raf. New Fl. 1: 53. 1836. Lotus americanus Bisch. Del. Sem. Hort. Heidelb. 1839. Trigonella sericea Eat. & Wright, N. Am. Bot. Ed. 8,459. 1840. 46 RYDBERG: Rocky MOUNTAIN FLORA Acmispon elatus (Nutt.) Rydb. Hosackia elata Nutt.; T. & G. Fl. 1: 327. 1838. The former of the two species is common on the plains from Minnesota to Arkansas, Sonora, and Idaho; the latter is found in Washington, Oregon, and Idaho. A few more species are found in California. Psoralea stenostachys Rydb. sp. nov. Perennial with a horizontal rootstock; stem adsurgent or erect, branched, sparingly strigose and glandular-dotted, 3-5 dm. high; leaves digitately 3-foliolate; leaflets oblanceolate, 2-4 cm. long, from rounded to acute at the base, mucronate at the apex, sparingly strigose and conspicuously glandular-punctate; peduncles _ 5-15 cm. long; racemes elongate, many-flowered and lax: calyx densely white-strigose; tube 1.5 mm. long; teeth 0.5 mm. long, lanceolate or lance-ovate, acute; corolla white, 4 mm. long; pod densely white-hairy. This species is related to P. lanceolata Pursh and P. Purshit Vail, but differs from both in the elongate racemes and the acute calyx-lobes; from the former it differs also in the hairy pod, and from the latter in the narrower leaflets. It grows on sandy soil at an altitude of about 1,300—1,500 m. Urtan: Government Well, Toole County, June 7, 1900, M. E. Jones 6221 (type, in herb. N. Y. Bot. Gard.); Utah, July 2, 1888, M. E. Jones 1833. Psoralea stenophylla Rydb. sp. nov. Perennial with a horizontal rootstock; stem simple, about 5 dm. high, slender, sparingly strigose and glandular-punctate; leaves digitately 3-foliolate or the lower 5-foliolate; leaflets narrowly linear, 2.5-5 cm. long, about 2 mm. wide, glandular-punctate and sparingly strigose; stipules linear, 5-8 mm. long; petioles about 3 cm. long; peduncles 8-10 cm. long; racemes elongate, 5 cm. long or longer, lax; pedicels usually longer than the calyx; calyx sparingly strigose, conspicuously punctate; lobes triangular, acute, 0.5 mm. long; corolla about 4 mm. long; fruit not seen. This has the narrow leaflets of Psoralea micrantha, but the raceme is elongate and the sepals are acute as in the preceding species, from which it differs in the very narrow leaflets. If it has the densely hairy pod of that species and P. Purshii, it cannot be told from the material, but the young ovaries do not indicate RYDBERG: Rocky MOUNTAIN FLORA 47 such a character. It grows on sandy river banks at an altitude of about 1,600 m. Utau: Proposed dam site, near Wilson Mesa, Grand County, July 1, 1911, Rydberg & Garrett 8367 (type, in herb. N. Y. Bot. Gard.). Psoralea juncea Eastw. was described as being leafless, the leaves being reduced to scales. This is true as far as the stem- leaves are concerned. The basal leaves, which soon wither away, are digitately 3-5-foliolate with lanceolate wee 2-3 cm. long, grayish, strigose and strongly veiny. Psoralea obtusiloba Torrey has been collected in Colorado by Tweedy. Parosela polydenia (Torr.) Heller, P. Fremontii (Torr.) Vail, P. Johnsoni (S. Wats.) Vail, and P. amoena (S. Wats.) Vail have been collected in southern Utah. Phaca ampullaria (S. Wats.) Rydb. Astragalus ampullarius S. Wats. Am. Nat. 7: 300. 1873. Phaca Wardii (A. Gray) Rydb. Astragalus Wardit A. Gray, Proc. Am. Acad. 12: 55. 1877. Phaca subcinerea (A. Gray) Rydb. . Astragalus subcinereus A. Gray, Proc. Am. Acad. 13: 366. 1878. Phaca Cusickii (A. Gray) Rydb. Astragalus Cusickii A. Gray, Proc. Am. Acad. 13: 370. 1878. Phaca sabulonum (A. Gray) Rydb. Astragalus sabulonum A. Gray, Proc. Am. Acad. 13: 368. 1878. Phaca Preussii (A. Gray) Rydb. Astragalus Preussit A. Gray, Proc. Am. Acad. 6: 222. 1864. Phaca serpens (M. E. Jones) Rydb. Astragalus serpens M. E. Jones, Proc. Cal. Acad. II. 5: 641. 1895. Phaca Silerana (M. E. Jones) Rydb. Astragalus Sileranus M. E. Jones, Zoe 2: 242. 1891. 48 RYDBERG: Rocky MOUNTAIN FLORA Phaca jejuna (S. Wats.) Rydb. Astragalus jejunus S. Wats. Bot. King Exped. 73. 1871. Phaca leptalea (A. Gray) Rydb. Phaca pauciflora Nutt.; T. & G. Fl. N. Am. 1: 348, 1838. Not P. pauciflora Pers. 1806. Astragalus leptaleus A. Gray, Proc. Am. Acad. 6: 220. 1864. : Phaca artemisiarum (M. E. Jones) Rydb. Astragalus Beckwithii purpureus M. E. Jones, Zoe 3: 288. 1893. Not A. purpureus Lam. 1783. Astragalus artemisiarum M. E. Jones, Zoe 4: 369. 1894. Phaca pubentissima (T. & G.) Rydb. Astragalus multicaulis Nutt.; T. & G. Fl. N. Am. 1: 335. 1838. Not A. multicaulis Ledeb. 1831. Astragalus pubentissimus T. & G. Fl. N. Am. 1: 693. 1840. Mr. Sheldon placed this between Astragalus crescenticarpus and A. cibarius, two species of Xylophacos; its pod is that of a Phaca. Phaca sesquiflora (S. Wats.) Rydb. Astragalus sesquiflorus S. Wats. Proc. Am. Acad. 10: 346. 1875. Mr. Sheldon associated this erroneously with Astragalus vexilliflexus and other species of Homalobus. It is a true Phaca. Xylophacos cuspidocarpus (Sheld.) Rydb. Astragalus cuspidocarpus Sheld. Minn. Bot. Stud. t: 147. 1894. Xylophacos cibarius (Sheld.) Rydb. Astragalus cibarius Sheld. Minn. Bot. Stud. 1: 149. 1894. Astragalus arietinus M. E. Jones, Proc. Calif. Acad. II. §: 653. 1895. Xylophacos puniceus (Osterh.) Rydb. Astragalus puniceus Osterh. Muhlenbergia 1: 140. 1906. . Xylophacos Zionis (M. E. Jones) Rydb. Astragalus Zionis M. E. Jones, Proc. Calif. Acad. IT. 5: 652. 1895. RYDBERG: Rocky MOUNTAIN FLORA 49 Xylophacos argophyllus (Nutt.) Rydb. Astragalus argophyllus Nutt.; T. & G. Fl. N. Am. 1: 331. 1838. Xylophacos cymboides (M. E. Jones) Rydb. Astragalus cymboides M. E. Jones, Proc. Calif. Acad. II. 5: 650. 1895. Xylophacos musinensis (M. E. Jones) Rydb. Astragalus musinensis M. E. Jones, Proc. Calif. Acad. II. 5: 671. 1895. Xylophacos consectus (Sheld.) Rydb. Astragalus consectus Sheld. Minn. Bot. Stud. 1: 143. 1894. Xylophacos Watsonianus (Kuntze) Rydb. Astragalus eriocarpus S. Wats. Bot. King Exped. 71. 1871. Not A. eriocarpus DC. 1802. Tragacantha Watsoniana Kuntze, Rev. Gen. Pl. 2: 942. 1891. Astragalus Watsonianus Sheld. Minn. Bot. Stud. 1: 144. 1894. Xylophacos utahensis (Torr.) Rydb. Phaca mollissima utahensis Torr. Stansb. Exped. 385. 1852. Astragalus utahensis T. & G. Pac. R. Rep. 2: 120. 1855. Xylophacos inflexus (Dougl.) Rydb. Astragalus inflexus Dougl. in G. Don, Gen. Syst. 2: 256. 1832. Tium eremiticum (Sheld.) Rydb. Astragalus eremiticus Sheld. Minn. Bot. Stud. 1: 161. 1894. Tium atropubescens (Coult. & Fish.) Rydb. Astragalus atropubescens Coult. & Fish. Bot. Gaz. 18: 300. 1893. Astragalus Kelseyi Rydb. Mem. N. Y. Bot. Gard. 1: 241. 1900. Tium arrectum (A. Gray.) Rydb. Astragalus arrectus A. Gray, Proc. Am. Acad. 8: 289. 1873. Astragalus Leibergii M. E. Jones, Proc. Calif. Acad. II. 5: 663. 1895. Astragalus palousiensis Piper, Bot. Gaz. 22: 489. 1896. 50 RYDBERG: Rocky MOUNTAIN FLORA Hamosa calycosa (Torr.) Rydb. Astragalus calycosus Torr. inS. Wats. Bot. King Exped. 66. 1871. Ctenophyllum Grayi (Parry) Rydb. Astragalus Grayi Parry; Wats. Am. Nat. 8: 212. 1874. Cystium platytropis (A. Gray) Rydb. Astragalus platytropis A. Gray, Proc. Am. Acad. 6: 526. 1865. Cystium Coulteri (Benth.) Rydb. Astragalus Coulteri Benth. Pl. Hartw. 307. 1848. Cystium ineptum (A. Gray) Rydb. Astragalus ineptus A. Gray, Proc. Am. Acad. 6: 525. 1865. Cystium lentiginosum (Dougl.) Rydb. _ Astragalus lentiginosus Dougl.; Hook. Fl. Bor.-Am. 1: 40. Par. Cystium araneosum (Sheld.) Rydb. Astragalus araneosus Sheld. Minn. Bot. Stud. 1: 170. 1894. Cystium boiseanum (A. Nels.) Rydb. Astragalus boiseanus A. Nels. Bot. Gaz. 53: 223. 4952, Atelophragma lineare Rydb. sp. nov. Homalobus aboriginum Rydb. Bull. N. Y. Bot. Gard. 2: 176, in part. Igol. Perennial with a woody taproot and short cespitose caudex; stem grayish strigdse, often tinged with purple, 2-4 dm. high; stipules ovate or lanceolate, acute, 2-4 mm. long; leaves 5-6 cm. long; leaflets 9-15, linear, 1-2 cm. long, I-2 mm. wide, grayish strigose; peduncles 5-10 cm. long; raceme 2-3 cm. long, in fruit 6 cm. long; calyx densely black-hairy; tube 3 mm. long; teeth subulate, 2 mm. long; corolla about 8 mm. long, ochroleucous or tinged with purple; keel tipped with dark purple; legume gla- brous, stipitate; stipe 4-5 mm. long; body 25-28 mm. long, con- vexly curved on both sutures, but much more strongly so on the upper; the partial partition very narrow. This is related to A. glabriusculum (A. Gray) Rydb. and A. aboriginum (Richardson) Rydb., but differs from the former in the RYDBERG: Rocky MouNTAIN FLORA 51 grayish pubescence of the leaves, which are strigose instead of villous, and from both in the form of the pod. In both the lower suture of the pod is straight or slightly concavely curved. YuKON TERRITORY: Foot of Lake Lebarge, 1899, J. B. Tarleton 345 (type, in herb. N. Y. Bot. Gard.); Dry Gulch, 1899, Gorman IOr4. ALBERTA: Rocky Mountains, 1857-1859, Bourgeau. Atelophragma Forwoodii (S. Wats.) Rydb. Astragalus Forwoodii S. Wats. Proc. Am. Acad. 25: 129. 1890. Sheldon places this species in the Homalobus, but it is closely related to Atelophragma aboriginum and A. glabriusculum. Atelophragma glabriusculum (Hook.) Rydb. Phaca glabriuscula Hook. Fl. Bor.-Am. 1: 144. 1831. Atelophragma ibapense (M. E. Jones) Rydb. Astragalus ibapense M. E. Jones, Zoe 3: 290. 1893. Atelophragma Arthuri (M. E. Jones) Rydb. Astragalus Arthuri M. E. Jones, Cont. West. Bot. 8: 20. 1898. Onix Mulfordae (M. E. Jones) Rydb. Astragalus Mulfordae M. E. Jones, Cont. West. Bot.8: 18. 1898. This species is the only representative in America of a group of plants segregated from Astragalus by Medicus. The other repre- sentatives are Asiatic. Onix is related to Cystium in having a membranous inflated 2-celled pod, but the pod is triangular in cross-section, the upper suture being acute and the lower more or less sulcate. Microphacos parviflorus (Pursh) Rydb. Dalea parviflora Pursh, Fl. Am. Sept. 474. 1814. Astragalus gracilis Nutt. Gen. 2: 100. 1818. Phaca parviflora Nutt.; T. & G. Fl. N. Am. 1: 348. 1838. Diholcos scobinatulus (Sheld.) Rydb. Astragalus Haydenianus major M. E. Jones, Zoe 2: 241. 1891. Astragalus Haydenianus nevadensis M. E. Jones, Zoe 2: 241. 189g. Astragalus scobinatulus Sheld. Minn. Bot. Stud. 1: 24. 1894. 52 RYDBERG: Rocky MOUNTAIN FLORA Phacopsis scaphoides (M. E. Jones) Rydb. Astragalus arrectus scaphoides M. E. Jones, Proc. Calif. Acad. IT. 5: 664. 1895. Cnemidophacos confertiflorus (A. Gray) Rydb. Astragalus confertiflorus A. Gray, Proc. Am. Acad. 13: 368. 1878. Cnemidophacos argillosus (M. E. Jones) Rydb. Astragalus argillosus M. E. Jones, Zoe 2: 241. 1891. Cnemidophacos reventoides (M. E. Jones) Rydb. Astragalus reventoides Jones, Proc. Calif. Acad. II. 5: 661. 1895. Cnemidophacos reventus (A. Gray) Rydb. Astragalus reventus A. Gray, Proc. Am. Acad. 15: 46. 1880. Kentrophyta tegetaria (S. Wats.) Rydb. Astragalus tegetarius S. Wats. Bot. King Exped. 76. 1871. Homalobus lingulatus (Sheld.) Rydb. Astragalus lingulatus Sheld. Minn. Bot. Stud. 1: 118. 1894. Homalobus exilifolius (A. Nels.) Rydb. Astragalus exilifolius A. Nels. Bull. Torrey Club 26: 10. 1899. Homalobus simplicifolius (Nutt.) Rydb. Phaca simplicifolia Nutt.; T. & G. Fl. N. Am. 1: 350. 1838. Astragalus simplicifolius A. Gray, Proc. Am. Acad. 6: 231. 1864. Homalobus lancearius (A. Gray) Rydb. Astragalus lancearius A. Gray, Proc. Am. Acad. 13: 370. 1878. Homalobus miser (Dougl.) Rydb. Astragalus miser Dougl.; Hook. Fl. Bor.-Am. 1: 153. 1831. Homalobus Dodgeanus (M. E. Jones) Rydb. Astragalus Dodgeanus M. E. Jones, Zoe 3: 289. 1893. Mr. Sheldon placed this next to Astragalus glabriusculus (Hook.) Gray, but its pod has not a trace of a partition and the plant is a true Homalobus, not an Atelophragma. RYDBERG: Rocky MOUNTAIN FLORA 53 Homalobus debilis (Nutt.) Rydb. Phaca debilis Nutt.; T. & G. Fl. N. Am. 1: 345. 1838. Astragalus debilis A. Gray, Proc. Acad. Sci. Phila. 1863: 60. 1864. Homalobus strigosus (Coult. & Fish.) Rydb. Astragalus strigosus Coult. & Fish. Bot. Gaz. 18: 299. 1893. Astragalus griseopubescens Sheld. Minn. Bot. Stud. 1: 24. 1894. Homalobus episcopus (S. Wats.) Rydb. Astragalus episcopus S. Wats. Proc. Am. Acad. 10: 346. 1875. Homalobus collinus (Dougl.) Rydb. Phaca collina Dougl.; Hook. Fl. Bor.-Am. 1: 141. 1831. Astragalus collina Dougl.; G. Don, Gen. Syst. 2: 256. 1832. Aragallus Bigelovii (A. Gray) Rydb. Oxytropis Lambertii Torr. Pac. R. Rep. 4: 80. 1857. Not O. _Lambertiit Pursh. 1814. Oxytropis Lambertii Bigelovii A. Gray, Proc. Am. Acad. 20: 7. 1885. Aragallus plattensis (Nutt.) Rydb. Oxytropis plattensis Nutt.; T. & G. Fl. N. Am. 1: 340. 1838. Lathyrus graminifolius White and L. Torreyi A. Gray have been collected in southern Utah; L. Nuttallii S. Wats. and L. obovatus White in Idaho. EUPHORBIACEAE © * Chamaesyce Parryi (Engelm.) Rydb. Euphorbia Parryi Engelm. Am. Nat. 9: 350. 1875. This has been collected in southern Utah. Chamaesyce exstipulata (Engelm.) Rydb. Euphorbia exstipulata Engelm. Bot. Mex. Bound. Surv. 189. 1859. Euphorbia Aliceae A. Nels. Bot. Gaz. 42: 50. 1906 This has been collected as far north as Wyoming. 54 RypBERG: Rocky MOUNTAIN FLORA ACERACEAE NEGuUNDO (Ray) Ludwig-Boehmer, Def. Pl. 508. 1760 Professor Nieuwland in the American Midland Naturalist* discussed the North American species of box-elder. He used the name Rulac, believing in a pre-Linnaean priority for genera. As both the Vienna Rules and the American Code have adopted 1753 as the starting point for botanical nomenclature, few will follow him in the names adopted. If our box-elders are regarded as generically distinct from the maples, we must use the name Negundo. Professor Nieuwland recognizes six species. I think there should be recognized eight species in North America. The Texan form, Rulac californica texana Pax, is well distinct from Negundo californicum, Professor Nieuwland having overlooked the difference in the fruit, which in the Texan species agrees more with our eastern box-elder and was included in it by Dr. Britton. The following key was prepared by me over two years ago and two new species were named in manuscript. One of these has been described by Professor Nieuwland under the name Rulac Nuttallii; a description of the other is given below. I publish here the key, as several of the characters have not been pointed out by Professor Nieuwland. Branches of the season glabrous or with a few scattered ap- ae haiis; anthers acute, tapering into a tip 4-4 m. long, formed ad the produced connective (in the ae species unknow Fruiting pedicels pe the lower 5~8 cm. long, very slender: fruit glabrous, contracted below into a short stipe. 1. N. orizabense. Fruiting pedicels sparingly pilose: the lower 2-3 cm. long. va and shen finely pubescent; sal latter some- stricted below into a narrow stipe-like base; leaflets broad, toothed, rarely lobed. Ovary and fruit glabrous; om latter slightly or usually not at all constricted be 1 renee logaae 4 lobed, with hair-tufts in the sorte of the v Branches of the season densely velutinous with aa spreading hairs; anthers obtuse, merely mucronate. Leaflets coarsely dentate or lobed; re evident but short. Fruit distinctly constricted at th tipe, densely and clisnieas sie Eehir leaflets * Vol. 2: 129-140. 10911. ore 2 aN. Negundo. 3- N. Nuttallii. RYDBERG: Rocky MOUNTAIN FLORA 65 broadly oval, short-acuminate, usually merely den- tate; the lateral ones often oblique at the base. 4. N. texanum Fruit not at all or slightly constricted at the base; leaflets lanceolate, ovate or obovate, or the ter- minal one rhombic, long-acuminate, usually more or less lobed Fruit glabrous or with a few scattered hairs, ilar to those of the pedicels; mucro the anthers minute or obsolete; leaflets glabrate above in age. Racemes seldom more than 1 dm. long; wings scarcely at all decurrent on the body of the fruit. 5. N. interius. Racemes in fruit 1.5-2 dm. long; wings de- current on the ge of the fruit almost to t m of the s 6. N. Kingii. Fruit ake See mucro of the anthers more distinct, nearly 144 mm. long; leaflets densely pubescent on both sides. 7. N. californicum, fiat 1 + a ays aoe e and evenly Pe LUWaAIOUS LIC Dast, style shades: 8. N. mexicanum. . Negundo orizabense Rydb. sp. nov. A tree with glabrous, brownish twigs; leaves 3-foliolate; pedicels slender, glabrous, 5-10 cm. long; leaflets thin, glabrous or with a few scattered hairs on the ribs below, acuminate at both ends, serrate above the middle, with broadly ovate teeth directed forward and mucronate; the terminal leaflet rhombic-oval, 5-10 cm. long, with petiolules 1-2 cm. long; the lateral ones lanceolate, oval or oblanceolate, short-petioluled; racemes in fruit 2 dm. long or more, the pedicels very long and slender, the lower 5-8 cm. long; samaras ascending, glabrous; body oblong, about I cm. long and 4 mm. wide, acute but not constricted at the base, with one strong and several weak longitudinal veins; wing about 2 cm. long and nearly I cm. wide, somewhat incurved above, not decurrent on the body Mexico: Orizaba, 1853 and 1855, Fred. Miiller (type, in herb. Columbia University). 2. Negundo Negundo (L.) Karsten, Deuts. Fl. 596. 1880-3* DISTRIBUTION: From Ontario and Vermont to Georgia, Mis- souri and Illinois. 3. Negundo Nuttallii (Nieuwl.) Rydb. Acer fraxinifolium Nuttall, Gen. N. Am. 1: 253. 1818. Not sect fraxinifolium Raf. 1808. zk Nieuwland, American Midland Nat 3 Igit. for ry y 56 RypBerG: Rocky MouNTAIN FLORA Rulac Nuitalli Nieuwl. Am. Midl. Nat. 2: 137. IgIt. DISTRIBUTION: From Michigan and Ohio (?) to Kansas, Colorado and Montana. 4. Negundo texanum (Pax) Rydb. Acer Negundo texanum Pax; Bot. Jahrb. 7: 212. 1886. Acer californicum texanum Pax; Bot. Jahrb. rr: 75. 1889. Rulac texana Small, Fl. SE. U. S..743. 1903. DISTRIBUTION: Texas and Oklahoma. 5. Negundo interius (Britton) Rydb. Rulac texana Small, Fl. SE. U.S. 743, in part. 1903. Not Acer texanum Pax. 1886. Acer interior Britton, N. Am. Trees 655. 1908. DISTRIBUTION: From Saskatchewan and Manitoba to Ne- braska, New Mexico, Arizona and Montana. Nieuwland gives Negundo Fraxinus Bourgeau* as a synonym under this. At the place referred to Bourgeau enumerates a number of genera col- lected on May 6. Evidently a comma is omitted between Negundo and Fraxinus. 6. Negundo Kingii (Britton) Rydb. Acer Kingii Britton, N. Am. Trees 656. 1908. Rulac Kingii Nieuwl. Am. Midl. Nat. 2: 139. I9QIt. DISTRIBUTION: Utah and Arizona. 7. NEGUNDO CALIFoRNICUM T. &. G. Fl. N. Am. 1: 250. 1838 Acer californicum Dietr. Syn. 2: 1283. 1840. Rulac californica Nieuw]. Am. Midl. Nat. 2: 139. IQIt. DISTRIBUTION: California and according to Nieuwland extend- ing into northern Mexico. 8. NEGUNDO MEXICANUM DC. Prod. r: 546. 1824 Acer mexicanum Pax; Bot. Jahrb. 7: 212. 1886. Not Acer mexi- canum A. Gray. 1861. Rulac mexicana Nieuwl. Am. Midl. Nat. 2: 140. Igil. DISTRIBUTION: Southern Mexico to Guatemala. * Journ. Linn. Soc. 4:9. 1850. RyDBERG: Rocky MounTAIN FLORA 57 RHAMNACEAE Rhamnus betulaefolia Greene is to be added to the flora; it was collected in southeastern Utah in the summer of 1911 by Professor Garrett and myself. MALVACEAE Dr. Greene* in segregating Eremalche from Malvastrum made this statement: ‘‘and that there exists so much as one real Mal- vastrum north of the Mexican border, I hold to be doubtful.” A little investigation in the history of the genus would show that this statement is untenable. It is evident that Dr. Gray did not base his conception of the genus Malvastrum on the section Malvastrum of Malva of De Candolle, for this section contains the typical species of Malva also. The first subsection of this section of De Candolle’s is Chry- santhae, and some species of this subsection must be regarded as the type of Malva section Malvastrum DC. Of this subsection Dr. Gray remarked: ‘‘If the yellow flowered species with a somewhat different habit and usually a manifest persistent involucre, which forms a second section (the Chrysanthae DC., etc.), are correctly referred to this genus, it will comprise a large number of species _ from tropical and South America, which need an elaborate revision. I enumerate below merely the North American species which are known to me.”’ Furthermore, Dr. Gray did not include in his genus a single species of Malva given by De Candolle. This shows that Dr. Gray based his genus on the North American species and in publishing the genus he gave the name as ‘‘ MAL- VASTRUM Nov. Gen.,’’ without citing De Candolle’s section, al- though he had referred to it a few pages before in a footnote under Callirrhoe. As the type of the genus Malvastrum, therefore, we must desigate the first given binomial under Malvastrum, which is M. coccineum. Of the other species included in the original publi- cation M. Fremontii Torr., M. Wrightii A. Gray, M. grossulariae- folium (Hook.) A. Gray, M. angustum A. Gray, M. Munroanum (Dougl.) Gray, and M. spicatum (L.) Gray are plants of the United States. I agree with Dr. Greene that M. rotundifolium A. Gray and M. exile A. Gray should not be included in Malvas- * Leaflets 1: 207. 1906. 58 RypDBERG: Rocky MOUNTAIN FLORA trum; but I believe that that genus should be merged in Sphaeral- cea. Malvastrum coccineum, the type of the genus, has the habit of the typical species of Sphaeralcea. The fruit is also the same except that the empty non-reticulate portion of the carpel is much reduced. M. grossulariaefolium and M. Munroanum with little more developed upper portions have been tossed back and forth between the genera Malvastrum and Sphaeralcea. Six species should be transferred from Malvastrum to Sphaeralcea under the following names. Sphaeralcea grossulariaefolia (H. & A.) Rydb. (?) Malva Creeana Graham, Bot. Mag. fl. 30698. 1838. Sida grossulariaefolia Hook. & Arn. Bot. Beech. Voy. 326. 1841. Malvastrum grossulariaefolium A. Gray, Mem. Am. Acad. 4: 21. 1849. Sphaeralcea pedata Torr. Mem. Am. Acad. 4: 23. 1849. Malvastrum coccineum grossulariaefolium Torr. Stansb. Exped. 384. 1852. ae Sphaeralcea dissecta (Nutt.) Rydb. Sida dissecta Nutt.; T. & G. Fl. N. Am. 1: 235. 1838. Malvastrum coccineum dissectum A. Gray, Pl. Wright. 1: 17, in part. 1852. Sphaeralcea coccinea (Nutt.) Rydb. Malva coccinea Nutt. Fras. Cat. 1813. Cristaria coccinea Pursh, Fl. Am. Sept. 454. Sida coccinea DC. Prod. 1: 465. 1824. Malvasirum coccineum A. Gray, Mem. Am. 1814. Acad. 4: 21. 1849. Sphaeralcea elata (E. G. Baker) Rydb. - Malvastrum coccineum elatum E, G. Baker, 189Ql. Malvastrum elatum A. Nels. Bot. Gaz. 34: 25. Jour. Bot. 29: 171. 1902. Sphaeralcea digitata (Greene) Rydb. Malvastrum coccineum dissectum A. Gray, part. 1852. Sphaeralcea pedata angustiloba A. Gray, 1887. Pl. Wright. 1: 17, in Proc. Am. Acad. 22: 202. RYDBERG: Rocky MouNTAIN FLORA 59 Malvastrum digitatum Greene, Leaflets 1: 154. 1905. Malvastrum dissectum Cockerell, Bull. Torrey Club 27: 87, mainly. 1900. Malvastrum Cockerellu A. Nels. Bot. Gaz. 34: 24. 1902. Malvastrum dissectum Cockerellii A. Nels.; Coult. & Nels. New Man. Bot. Cent. Rocky Mts. 318. 1909. Sphaeralcea leptophylla (A. Gray) Rydb. Malvastrum leptophyllum A. Gray, Pl. Wright. 1: 17. 1852. Sphaeralcea arizonica Heller, sp. nov. Perennial with a woody caudex branching from the base; leaf- blades reniform to cordate, 3-5 cm. long, densely stellate on both sides, obscurely lobed and crenate; inflorescence paniculate, dense, with short branches; calyx densely stellate throughout; its lobes ovate, acute, about 3 mm. long; petals pink, about 1 cm. long; carpels about 4 mm. long and 1.5 mm. wide, mucronate or short- cuspidate, oblong, only about the aa fourth reticulate. Differing from S. ambigua in the short calyx-lobes and the narrow and dense inflorescence and from S. marginata in the dense stellate pubescence, which extends even to the calyx. Arizona: Flagstaff, June 16,1898, MacDougal 120 (type, in herb. N. Y. Bot. Gard.) ; 30 miles east of Flagstaff, July 18, 1893. Wooton; Fort Verde, May 4, 1888, Mearns 225; same locality 1887, 150; Hol- brook, June 18, 1901, L. F. Ward; Ash Fork, June Io, 1883, Rusby 538. Urau: St. George, Apr. 14, 1880, M. E. Jones 1660; proposed dam site, near Wilson Mesa, Grand Co., July 1, 1911, Rydberg & Garrett 8386; S. Utah, 1877, Palmer; 1874, Parry 25. Sphaeralcea subrhomboidea Rydb. sp. nov. Perennial with a woody caudex, branched at the base; stems stellate, 2-4 dm. high; leaf-blades rhombic in outline, 2-5 cm. long, stellate but not densely so, grayish-green, cuneate at the base, 5-ribbed, 3-cleft about half way down, the divisions 2-4- lobed; inflorescence a dense virgate panicle; calyx densely stellate, 4-5 mm. long; lobes broadly ovate, obtusish; corolla scarlet, 8-9 mm. long; fruit depressed-globose; carpels nearly round, obtuse, the lower half reticulate on the faces; seed solitary, without filiform attachment. 60 RYDBERG: Rocky MOouNTAIN FLORA Nearest related to S. grossulariaefolia but the leaf-blades are rhombic in outline and cleft only half way down, and the terminal lobe is decidedly acute. On account of the leaf-form it may be mistaken for S. Munroana, but the flowers are smaller, the leaves more deeply divided, the fruit is smaller, the carpels less reniform, and the seed without filiform attachment. Utan: Wahsatch County, near Midway, July 6, 1905, Carlton & Garrett 6691 (type, in herb.“N. Y. Bot. Gard.); Fish Lake, around Twin Creeks, Aug. 8, 1905, Rydberg & Carlton 7627. There is a group of plants in Sphaeralcea, however, which differs from the rest not only in habit but also in the character of the fruit. The carpels are not, as in the typical Sphaeralcea, divided into a lower portion, reticulate on the faces and enclosing the seeds, and an upper smooth and empty portion; the whole carpel is in this group smooth and hirsute. Dr. Greene* took out this group and made a new genus under the name of Jiliamna. I think that this was unnecessary, for the plants are evidently cogeneric with the West Indian Phymosia, usually also merged in Sphaeralcea. If the two genera should be merged, the name for the genus would be Phymosia, for it is the older of the two. The species to be renamed under Phymosia are the following: Phymosia acerifolia (Nutt.) Rydb. Splaieicen acertfolia Nutt.; T. & G. Fl. N. Am. 1: 228. 1838. Illamna acerifolia Greene, Leaflets 1: 206. 1906. Phymosia rivularis (Dougl.) Rydb. Malva rivularis Doug!l.; Hook. Fl. Bor.-Am. 1: 107. 1831. Sphaeralcea rivularis Torr. in Gray, Mem. Am. Acad. 4: 23. 1849. Illiamna rivularis Greene, Leaflets 1: 206. 1906. Phymosia grandiflora Rydb. Sphaeralcea grandiflora Rydb. Bull. Torrey Club 31: 565. 1904. Illiamna ane Greene, Leaflets 1: 206. 1906. ae Phymosia Crandallii Rydb. Sphaeralcea Crandallu Rydb. Bull. Torrey Club 31: 564 1904. t Hs zs * Leaflets 1: 205-207. 1906. RypBERG: Rocky MOUNTAIN FLORA 61 Phymosia longisepala (Torr.) Rydb. Sphaeralcea longisepala Torr. Bot. Wilkes Exped. 255. 1874. LOASACEAE Nuttallia* humilis (A. Gray) Rydb. Mentzelia multiflora humilis A. Gray, Pl. Wright 1: 74. 1852. Touterea humilis Rydb. Bull. Torrey Club 30: 277. 1903. Nuttallia integra (M. E. Jones) Rydb. Mentzelia multiflora integra M. E. Jones, Proc. Calif. Acad. IT. 5: 689. 1895. - Touterea integra Rydb. Fl. Colo. 235. 1906. Nuttallia Rusbyi (Wooton) Rydb. Mentzelia Rusbyi Wooton, Bull. Torrey Club 25: 261. 1898. Touterea Rusbyi Rydb. Bull. Torrey Club 30: 276. 1903. Nuttallia lobata Rydb. sp. nov. Perennial with a thick root; stems strict, glabrous or nearly so, white and shining, 3-4 dm. high; leaves 5-8 cm. long, 5-8 mm. wide, narrowly 6blanceolate, sinuately toothed or lobed with short triangular lobes; sepals lanceolate, acuminate, 8-10 mm. long; flowers diurnal, subtended by narrowly linear bracts; petals golden yellow, spatulate, obtuse, 12-18 mm. long; petaloid staminodia similar and almost as large; filaments numerous, the outer dilated ; capsule 15 mm. long, 8-9 mm. thick, acute, almost turbinate at the base; seeds suborbicular, broadly winged. This species is related to N. multiflora (Nutt.) Greene and N. pterosperma (Eastwood) Greene. It differs from the former in the narrow merely toothed or lobed not pinnatifid leaves; from the latter in the acute teeth or lobes of the leaves and the capsule, which is acute not rounded at the base, and from both in the glabrous stem. Uran: Near St. George, 1877, Palmer 172 (type, in herb. Columbia Univ.); 1874, Parry 76; 1902, Goodding 770. Nuttallia acuminata Rydb. sp. nov. Stout biennial; stem 3-10 dm. high, straw-color, white in age, rather dull, densely villous with barbed hairs; lower leaves Sab ie aM RE Gil IGRI sO RIE, ECL SR ie ae ene eae WE SET REST *“ Nuttalle’”’ Rafin. Am. Mo. Mag. (1818): 175. 1818; ‘* Nuttallia”’, Greene. Leaflets x: 209. 10 Ap. 1906. 62 RypBERG: Rocky MOUNTAIN FLORA oblanceolate, 1-2 dm. long, sinuately dentate, densely scabrous with triangular teeth; upper stem-leaves lanceolate, long-acu- minate, pinnatifid with lanceolate or rarely triangular lobes, the lower ones of which are usually large and salient, the base of the leaves, therefore, being very broad and truncate; flowers diurnal; their bracts narrowly linear, entire or with a few narrow lobes; sepals 2-3 cm. long, lance-subulate, long-acuminate, light yellow, about 5 cm. long; outer filaments slightly dilated, the rest fili- form, three fourths as long as the petals; petaloid staminodia none; _ capsule 4 cm. long, 1 cm. thick; seeds obovate, winged. This species has been confused with N. laevicaulis (Hook.) Greene, but differs in the pubescent, duller stem (in N. laevicaulis this is glabrous or with a few scattered stiff hairs, very white and shining), broader petals, more deeply divided upper stem-leaves, which are characterized by their acumination and broad almost subhastate bases. N. acuminata extends farther eastward and northward than N. laevicaulis and is lacking in California. _IpaHOo: Spokane River, Kootenai County, 1892, Sandberg, MacDougal & Heller 651 (type, in herb. N. Y. Bot. Gard.) ; Palouse County and Lake Coeur d’Alene, Aiton 6015. Montana: Emigrant Gulch, 1897, Rydberg & Bessey 4540; Sedan, 1902, W. W. Jones; Garrison, 1895, Rydberg 2737, and C. L. Shear 5248; Helena, 1892, Kelsey. - Wyominc: Between Sheridan and Buffalo, 1900, Tweedy 3617; Gardiner River, 1899, Aven Nelson & Elias Nelson 6000. Uta: City Creek, 1883, Leonard 116 and 227; Beck’s Hot Spring, 1905, Garrett 1595; Antelope Island and Stansbury Island, Stansbury. WasHINGTON: Loon Lake, 1897, Winston; Spokane, 1902, Kraeger 529. ONAGRACEAE Boisduvalia salicina (Nutt.) Rydb. Oenothera densiflora 8 T. & G. Fl. N. Am. 1: 50s. 1840. Oenothera salicina Nutt. in T. & G. loc. cit., aS a synonym. This is quite different in habit from the typical B. densiflora (LindI.) 5. Wats., having the foliage-leaves narrow, linear or linear- lanceolate. It has a much more northern and eastern range, extending into British Columbia and Idaho. RYDBERG: Rocky MOUNTAIN FLORA 63 Epilobium latiusculum Rydb. sp. nov. Epilobium Drummond latiusculum Rydb. Mem. N. Y. Bot. Gard. e270. 1900. To the characters given in the original description may be added that the leaves are distinctly petioled, not sessile as in E. Drummondit. Epilobium platyphyllum Rydb. sp. nov. Epilobium glaberrimum latifolium Barbey, Bot. Calif.1: 220. 1876. Not E. latifolium L. 1753. Epilobium paniculatum, as usually understood, contains several forms or species, connecting on one hand with E. minutum, on the other with E. jucundum. In order to facilitate the further study of the groups, I give the following key of the Rocky Mountain forms. Tube of the hyper funnelform, 1-3 mm. (rarely 4 mm.) pias sunita only slightly exceeding the calyx, 2-3 mm. long; capsule glabrous; tube of hypanthium I-1.5 mm. long. 1. E. Tracyi. Petals pink or purple, 3.5-7 mm. long, about twice as long as the calyx Capsule and iiiceia glabrous or sparingly puberulent. Leaves and bracts very thick, horny at the apex, the latter very short; capsule glabrous; pedicels short. 2. E. subulatum.. Leaves and bracts not very thick, not. horny at the apex; capsule usually puberulent, at least when young; pedicels slender 3. E. paniculatum. Capsule and pedicels cial peaeeneieks ——€, very short. 4, E. adenocladum, Tube of the hypanthium 4-8 mm. long, cylindric or nearly so, abruptly widening into the calyx. Tube of the hypanthium about 4 mm. long; petals 6-7 mm. long. 5. E. laevicaule. Tube of the hypanthium 7-8 mm. long; petals 10-12 mm. long 6. E. Hammondii. Epilobium Tracyi Rydb. sp. nov. Annual; stem 3-8 dm. high, perfectly glabrous, straw-colored; leaves 2-4 cm. long, linear, entire, glabrous; tube of the hypan- thium 1-1.5 mm. long, funnelform; calyx-lobes about 2 mm. long, very acute; petals white, 2-3 mm. long; capsule more or less 64 RypBERG: Rocky MOUNTAIN FLORA clavate, about 1.5 cm. long, perfectly glabrous; seeds obovoid, 1.5 mm. long. This species is related to E. paniculatum but differs in the small white flowers and the perfectly glabrous pod. Ura: Ogden, July 31, 1887, Tracy & Evans 547 (type, in herb. N. Y. Bot. Gard.); Salt Lake City, May 1869, Watson 390. OrEGON: Washington County, July 4, 1894, F. E. Lloyd. WASHINGTON: Spokane, July 11, 1902, Kraeger 152. Ipano: Little Potlatch River, Latah County, June 17, 1892, Sandberg, MacDougal & Heller 477. Montana: Moraine near Polson, August 18, 1901, Umbach. British CoLumBiA: Howser Lake, Selkirk Mts., June 17, 1905, Charles H. Shaw 714. NevabAa: Huntington Valley, August 1868, Watson 390. Epilobium subulatum (Haussk.) Rydb. Epilobium paniculatum subulata Haussk. Monog. Epil. 247. 1884. Epilobium laevicaule Rydb. sp. nov. Annual; stem glabrous, 6-10 dm. high, glabrous and shining; the bark of the lower portion flaky; leaves linear or linear-lanceo- late, 3-6 cm. long; the upper mostly involute, usually entire; tube of the hypanthium about 4 mm. long, rather abruptly widening into the calyx; calyx-lobes 3-4 mm. long; petals rose-colored, 6-7 mm. long; pods clavate, about 3 cm. long, glabrous or almost s0; seeds obovoid, dark; coma dingy. Montana: Manhattan, 1895, Rydberg 2728 (type, in herb. N. Y. Bot. Gard.); Shear 3114; Big Fork, Aug. 3, 1909, Butler 7016. WASHINGTON: Pullman, Aug. 5, 1893, Piper 1631; Spokane, Sept. 1902, Kraeger 536 and 573. IpaHOo: Palouse County, 1892, G. B. Aiton 69; Seven Devils Mountains, Aug. 5, 1899, M. E. Jones 6317. Epilobium Sandbergii Rydb. sp. nov. Perennial by means of turions; stem obtusely angled, 6-10 dm. high, finely puberulent throughout; leaves sessile, ovate, acute, dentate, 3-7 cm. long, pubescent on beneath, except the veins; inflorescence linear-lanceolate, about 5 mm. long; pod 4-6 cm. long, glandular-pilose; seeds 1.5 mm. long, almost peakless; coma tawny. RyDBERG: Rocky MOUNTAIN FLORA 65 - It resembles somewhat E. Palmeri, but the flowers are nearly twice as large. Ipauo: Moist places, valley of Mud Lake, Kootenai County, July 25, 1892, Sandberg, MacDougal & Heller 737 (type, in herb. N. Y. Bot. Gard.). Montana: Bozeman, July 22, 1895, Rydberg 2720. Gayophytum Helleri Rydb. sp. nov. Annual; stem branched with nearly erect, strict branches, 1-3 dm. high, more or less pubescent with spreading hairs; leaves linear, 0.5-2 cm. long, softly hirsutulous; pedicels very short, even in fruit scarcely more than 1 mm. long; sepals and petals scarcely 1 mrh. long; capsules linear, erect, 8-10 mm. long, almost sessile, hirsutulous, not torulose; seeds about 1 mm. long, strigu- lose This resembles G. racemosum in habit and the pod, G. caesiun in pubescence and G. lasiospermum in the seeds. IpaHo: Forest, Nez Perces County, July 16, 1896, Heller 3433 (type, in herb. N. Y. Bot. Gard.). Anogra leptophylla (Nutt.) Rydb. = Oenothera pallida leptophylla (Nutt.) T. & G. Fl. N. Am. 1: 495. 1840. ; Oenothera leptophylla Nutt.; T. & G. FI. loc. cit., as a synonym. Oenothera longissima Rydb. sp. nov. A tall biennial; stem strict, 5-10 dm. high, densely canescent with short crinkled hairs as well as sparingly hirsute; leaves linear or narrowly linear-lanceolate, 1-1.5 dm. long, densely canescent, entire, acute at both ends, the lower short-petioled; spike rather lax: bracts linear-lanceolate, 2-5 cm. long; hypanthium tube 10- 12 cm. long, densely canescent, only slightly widening upwards; sepals linear-lanceolate, about 4 cm. long; free tips about 4 mm. long; petals golden yellow, 4 cm. long; stamens and pistil of about the same length; capsule about 4 cm. long, densely canescent, slightly tapering upwards. This is related to O. macrosceles A. Gray and O. Jamesii T. & G., but differs from the former in being canescent instead of glabrous and in the smaller and narrower bracts, and from the latter in the longer, narrower and entire-margined leaves, and in being more canescent and less hirsute.. It grows on sandy river banks at an altitude of about 1,600 m. 66 RyDBERG: Rocky MOUNTAIN FLORA Uran: Armstrong and White Canyons near the Natural Bridges, Aug. 4-6, 1911, Rybderg & Garrett 9410 (type, in herb. N. Y. Bot. Gard.). Oenothera ornata (A. Nelson) Rydb. Onagra ornata A. Nels. Bot. Gaz. 52: 268. I9gII. Oenothera hirsutissima (A. Gray) Rydb. Oenothera biennis hirsutissima A. Gray, Mem. Am. Acad. 4. 43. 1849. This usually has been regarded as the same as O. Hookeri T. & G. The type of the latter came from California, that of the former from New Mexico. In the plant common in California and the Great Basin, the free tips of the sepals are about 4 mm. long, the pubescence of the leaves is short and that of the calyx not very copious. In the type of O. biennis hirsutissima and other specimens from New Mexico and Colorado, the free tips of the sepals are only 2-2.5 mm. long, the pubescence of the leaves and calyx long and loose, and that of the latter very copious. Oenothera subulifera (Rydb.) Onagra strigosa subulata Rydb. Mem. N. Y. Bot. Gard. 1 : 279. 1900. Not O. subulata R. & P. 1802 Onagra Oakesiana Rydb. Fl. Colo. 244. 1906. Not Oenothera Oakesiana A. Gray. 1867. Chylisma tenuissima (M. E. Jones) Rydb. Oenothera tenuissima M. E. Jones, Proc. Calif. Acad. II. 5: 683. * 1895. Sphaerostigma macrophyllum (Small) Rydb. Oenothera alyssoides villosa S. Wats. Proc. Am. Acad. 8: 591. 1873. Not O. villosa Thunb. 1794-1800. Sphaerostigma alyssoides macrophyllum Small, Bull. Torrey Club 23: 192. 1896. AMMIACEAE Osmorrhiza intermedia Rydb. Washingtonia intermedia Rydb. Mem. N. Y. Bot. Gard. 1: 289. 1900. RYDBERG: Rocky MOUNTAIN FLORA 67 Glycosma maxima Rydb. sp. nov. Perennial; stem 1 m. high or more, puberulent or glabrous, pilose at the nodes; lower leaves twice compound, first pinnate and the lower primary divisions ternate; the upper leaves ternate or twice ternate; leaflets oblong-lanceolate, 5-10 cm. long, minutely puberulent; branches of the umbels 9-12, in fruit more or less spreading; pedicels in fruit 1-1.5 cm. long; fruit fully 2 cm. long, obtuse at the base, contracted above into a beak 2 mm. long; stylopodium conical, 0.5 mm. long, about as long as the styles. This is related to G. occidentalis Nutt., but the fruit is much larger (in G. occidentalis only 12-16 mm., rarely 18 mm. long), and the rays of the umbels are in fruit usually widely spreading, while in G. occidentalis they are nearly erect. The spreading rays sug- gest G. ambigua and G. Bolanderi, but in both these species the stylopodium is flatter. Uran: Mount Nebo, Aug. 15, 1905, Rydberg & Carlton 7585 (type, in herb. N. Y. Bot. Gard.); Rocky Canyon, Provo, Aug. 16, 1887, Tracy 684. Montana: Midvale, July 24, 1903, Umbach 508. Atenta H. & A. Bot. Beech. Voy. 349. 1840 This I think is a good genus, distinct from Carum. Although the fruit is almost the same, the habit is quite different. The habit of Atenia is the same as that of Eulophus. In fact it is hard to distinguish the two genera without mature fruit, both having the fascicled tuberous roots, the narrow leaf-segments, the same inflorescence and flowers. The only essential differences are the deeply concave seed-face with a central ridge and the sev- eral oil tubes in Eulophus and the plane face and solitary oil tubes in Ataenia. The following species are found in the Rocky Mountains: Atenta GAIRDNERI H. & A. Bot. Beech. Voy. 349. 1840 Edosmia Gairdneri Nutt.; T. & G. Fl. N. Am. 1: 612. 1840. Carum Gairdneri A. Gray, Proc. Am. Acad. 7: 344- 1867. Atenia montana (Blank.) Rydb. Carum montanum Blank. Mont. Agr. Coll. Sci. Bot. 1: 9I. 1905. . 68 RypBERG: Rocky MOouNTAIN FLORA Atenia Garrettii (A. Nels.) Rydb. Carum Garrettii A. Nels. in Rose, Cont. U. S. Nat. Herb. 12: 443. 1909. Oreoxis MacDougali (C. & R.) Rydb. Aletes MacDougali C. & R. Cont. U.S. Nat. Herb. 7: 107. 1900. This was doubtfully referred to Aletes by Coulter and Rose. The fruits in the type collection were very young and did not show their true nature. Anyhow, they showed distinct wings, a character inconsistent with the genus Aletes. Professor Garrett and myself collected good fruits in southeastern Utah in the summer of 1911; and these show that the plant is rather an Oreoxis than an Aletes, wings being present and these thick and corky. The two genera are, however, more closely related than has been recognized, having the same cespitose habit, the promi- nent calyx, teeth, etc. Daucophyllum (Nutt.) Rydb. gen. nov. Musenium § Daucophyllum Nutt.; T. & G. Fl. N. Am. 1: 642. 1840, Low cespitose perennials, acaulescent or nearly so, with a branched caudex. Leaves numerous, basal, or 1 or 2 cauline, pinnate or bipinnate with filiform or narrowly linear divisions. Flowers cream-colored to yellow, in dense umbels. Bracts want- ing; bractlets few, narrow, linear. Calyx teeth prominent. Stylopodium wanting. Fruit ovoid or oblong, granular on the intervals. Ribs equal, rather strong, but not at all winged. Oil tubes 2 or 3 in the intervals, 4-6 on the commissural side. Seed terete or somewhat depressed; face plane. The type, Musenium tenuifolium Nutt., was separated as a section in Torrey and Gray’s Flora. The relationship is rather with Harbouria and Aletes than with Musineon Raf. The first- mentioned relationship was recognized by Coulter and Rose (see their Revision, p. 111). It differs from Harbouria in not having thick corky ribs and in having several oil tubes in the intervals. It is still more closely related to Aletes, having the same habit, although narrower leaf-segments, the main differences being, however, the solitary oil tubes in Aletes and 2 or 3 in each interval in Dauco- phyllum, and the concave seed face in the former and the plane — So Ras es RYDBERG: Rocky MouNTAIN FLORA 69 one in the latter. The second species given below was included questionably in Aletes by Coulter and Rose; but in the number of oil tubes and the plane seed face it agrees better with Musenium tenuifolium Nutt. than with the typical species of Aletes. Leaves bipinnate; segments filiform; bractlets not exceeding the pedicels; seed subterete. 1. D. tenuifolium, Leaves pinnate; segments narrowly linear; bractlets longer than the pedicels; seeds somewhat depressed. 2. D. lineare. 1. Daucophyllum tenuifolium (Nutt.) Rydb. Musenium tenutfolium Nutt.; T. & G. Fl. N. Am. 1: 642. 1840. 2. Daucophyllum lineare Rydb. nom. nov. Aletes tenuifolia C. & R. Cont. U. S. Nat. Herb. 7: 108. 1g00. Coriophyllus (M. E. Jones) Rydb. gen. nov. Cymopterus §Coriophyllus M. E. Jones, Cont. West. Bot. 12: 20. 1908. Perennial herbs with more or less fleshy root, somewhat branched rootstock covered with fibrous sheaths, and_ leafy stems with internodes shorter than the leaf-sheaths. Flowers yellow to purple. Bracts none; bractlets present, but narrow. Leaves pinnately dissected, subcoriaceous, rigid, not fleshy, with ovate or lanceolate, cuspidate or spinulose-tipped lobes. Calyx teeth evident. Stylopodium wanting. Fruit orbicular to oval in outline, usually emarginate at both ends, compressed laterally if at all. Ribs with broad wings. Oil tubes 1-5 in the intervals, 2-8 on the commissural side. Seeds little if at alt flattened dor- sally; face deeply grooved. I agree with Mr. Marcus E. Jones that the genus Aulospermum, as constituted by Coulter and Rose, is a rather unnatural one, made up of two groups of quite different habit; but instead of reducing both groups to sections of Cymopterus as Mr. Jones did, I rather regard them as two distinct genera, and adopt for the second group the sectional name first proposed by Mr. Jones. (See the discussion in Cont. West. Bot. 12: 19-20 and 27.) He, however, had the group under two different sectional names. The section is called Coriophyllus on page 20 and Scopulicola on page 27. The following species are found in the Rockies and are dis- US. thus: ‘ 70 RypBerc: Rocky MounTAIN FLORA Wings thickened at the insertion. Leaves ternately bipinnatifid; oil tubes solitary in each i . C. Jonesii. 4 nterval. : Leaves pinnate, with lobed or divided leaflets; oil tubes several in each interval. 2. C. Roses, Wings not thickened at the insertion. Flowers purplish; oil tubes 8 on the commissural side. 3. C. purpureus. Flowers greenish-yellow; oil tubes 4 on the commissural side. 4. C. Bethelt. 1. Coriophyllus Jonesii (C. & R.) Rydb. Cymopterus Jonesit C. & R. Rev. N. Am. Umb. 80. 1888. Aulospermum Jonesii C. & R. Cont. U. S. Nat. Herb. 7: 178. 1900. 2. Coriophyllus Rosei (M. E. Jones) Rydb. Aulospermum Rosei M.E. Jones; C. & R. Cont. U.S. Nat. Herb. 7: 179. 1900. 3. Coriophyllus purpureus (S. Wats.) Cymopterus purpureus S. Wats. Am. Nat. 7: 300. 1872. Aulospermum purpureum C. & R. Cont. U. S. Nat. Herb. 7: 178. 1900. 4. Coriophyllus Betheli (Osterhout) Rydb. Aulospermum Betheli Osterhout, Muhlenbergia 6: 46. I9I0. ‘ PSEUDOCYMOPTERUs C. & R. : This genus is one of the most unnatural in Coulter & Rose’s 4 Monograph. Jones* called attention to this fact, although he included the genus, as well as Oreoxys, Rhysopterus, Aulospermum, — and Pteryxia in Cymopterus, and does not go to the bottom of the facts. The genus as constituted by Coulter and Rose contains at least three distinct groups of plants of little relationship to each : other. The first group contains Pseudocymopterus montanus and its close relatives; the second of P. anisatus and P. aletifolius, and perhaps P. Hendersonii, which I do not know; and the third of P. bipinnatus and probably Cymopterus nivalis S. Wats., of which ee fruit is unknown. P. montanus is the type of the genus, wh latter therefore must be restricted to it and its relatives. Jones * Cont. West. Bot. 12: 24-20. 1908. os RYDBERG: Rocky MounrTAIN FLORA 71 includes P. anisatus and P. bipinnatus in his section Oreoxis, but the genus Oreoxis has all ribs corky and the lateral ones scarcely more prominent than the dorsal ones, the fruit is not flattened dorsally, the styles and sepals are erect. In Pseudocymopterus anisatus the lateral wings are very prominent, the dorsal ribs narrowly winged or some of them merely acute, the styles are recurved, the sepals spreading and one or two of them larger than the rest, and the fruit is decidedly flattened dorsally. The plant is more related to Aletes than to Oreoxis, and P. aletifolius connects it with that genus. It can not be placed in Aletes, however, for in that genus the fruit is not compressed and the ribs not winged. It would be much better to include P. anisatus and P. aletifolius in Pteryxia, as they have the foliage and nearly the same fruit as in that genus, but the strictly acaulescent plant, the narrow and thick wings of the fruit and the very prominent and unequal calyx-teeth would make it rather abnormal even in that genus. Although it does not differ so much in the technical characters of the fruit from the typical Pseudocymopterus, the habit is quite different, so also the texture of the leaves, and in Pseudocymopterus the sepals are minute. It is better to regard P. anisatus as a type of a new genus, Pseudopteryxia Rydb. gen. nov. Densely cespitose, strong-scented, acaulescent perennials with multicipital caudices covered with numerous sheaths of old leaves. Leaves pinnatifid or bipinnatifid with thick, firm, pungent divi- sions. Flowers yellow; involucres wanting; bractlets linear-subu- late, pungent. Calyx-teeth very prominent, spreading, unequal, one or two much longer than the rest. Stylopodium wanting. Fruit oblong, glabrous. Ribs thick, the dorsal and intermediate ones sharp or some of them with narrow wings; the lateral ones with broader wings, distinct from those of the other carpel. Carpels flattened dorsally. Oil tubes 1-3 in the intervals, 2-4 on the commissural side. Seed face plane. Pseudopteryxia anisata (A. Gray) Rydb. 3 Cymopterus (?) anisatus A. Gray, Proc. Acad. Phila (18622 ( 1863. Psciuloeymapbecs anisatus C. & R. Rev. N. Am. Umb. 75. “1888, ; %2 RypBERG: Rocky MOUNTAIN FLORA Pseudopteryxia longiloba Rydb. sp. nov. Densely cespitose perennial with a thick root and short caudex, covered by numerous old leaf-sheaths and petioles; leaves twice pinnatifid, with linear-subulate, pungent divisions; peduncles 2-3 dm. high, stout; bractlets linear-subulate, spreading, often I cm. long; flowers yellow; fruit about 6 mm. long; lateral wings thick, narrow, some of the wings of the dorsal ribs often fully as broad; calyx-teeth less prominent than in P. antsata. This is closely related to P. anisata, differing in the larger fruit (in P. anisata about 4 mm. long), and longer leaf-segments. On account of the long leaf-segments, specimens collected in flower by Carlton and myself were mistaken for Cynomarathrum Nuttall (A. Gray) C. & R.; but good fruit was received in the summer of IQII. Uran: Abajo Mountains, Aug. 17, 1911, Rydberg & Garrett 9761 (fruit; type, in herb. N. Y. Bot. Gard.); also 9760 (fruit); La Sal Mountains, July 7 and 17, 8724 and gors (young fruit); Mountains north of Bullion Creek, near Marysvale, July 23, Rydberg & Carlton 7085 and 7096 (flowers); Mount Ellen, July 24 and 25, 1894, M. E. Jones 5677 (fruit, but poor). Pseudopteryxia aletifolia Rydb. Pseudocymopterus aletifolius Rydb. Bull. Torrey Club 31: 574- 1904. Neither can Pseudocymopterus bipinnatus be retained in the genus; in fact, itis still more out of place. Not only is the habit strikingly different from that of P. montanus, but the fruit is not, as Coulter and Rose described it, ‘‘moderately flattened dorsally,” for the fruit when well developed is moderately flattened laterally, which places it in the other division of the family. Furthermore, the seed face is concave, the bractlets broad and scarious, and a stylopodium, although strongly flattened, is present. Were it not for these characters of the fruit the plant could be placed in the same genus as P. anisatus. As it is, its relationship is with Daucophyllum and Aletes. 1 would place it in Daucophyllum were it not for the winged ribs, the concave seed face and the reflexed style. The fruit is nearer that of Aletes, but the oil tubes are several, the ribs winged, styles reflexed and stylopodium present. Ifa person were using the key given by Coulter and Rose RYDBERG: Rocky MOUNTAIN FLORA 73 in their Monograph and were trying to determine the plant, the key would lead to Aulospermum or Phellopterus, to either of which genera it is not even closely related. Mr. Jones included it in Oreoxis, to which I admit it is related, but the ribs are not corky, the stylopodium present, the styles reflexed, the flowers white, not yellow, and the bractlets scarious. Pseudoreoxis Rydb. gen. nov. Low cespitose acaulescent perennials, with branched caudex. Leaves bipinnate; the segments more or less cleft with small lanceolate divisions. Flowers white in small umbels; bracts want- ing; bractlets ovate or lanceolate, cuspidate or abruptly acumi- nate, scarious, white with a green midrib. Calyx-teeth evident but small. Stylopodium present but low and flat. Styles reflexed. Fruit somewhat flattened laterally, oblong. Ribs all with narrow wings, the lateral ones scarcely wider. Oil tubes 3 or 4 in the intervals, 6-8 on the commissure. Seed face slightly concave. Pseudoreoxis bipinnatus (S. Wats.) Rydb. Cymopterus bipinnatus S. Wats. Proc. Am. Acad. 20: 368. 1885- Pseudocymopterus bipinnatus C. & R. Rev. N. Am. Umbel. 75. 1888. Pseudoreoxis nivalis (S. Wats.) Rydb. Cymopterus nivalis S. Wats. Bot. King. Exped. 123. 1871. I do not hesitate to refer this species to the same genus as P. bipinnatus, although the fruit is unknown, for the habit, and flowers are so closely resembling those of P. bipinnatus. Cynomarathrum latilobum Rydb. sp. nov. Acaulescent perennial with densely cespitose caudex covered by old. broad leaf-sheaths; leaves about 1 cm. long, pinnate, glabrous; leaflets entire or 2- or 3-cleft into broadly lanceolate, reticulate, thick, pointed segments 5-15 mm. long; peduncles I-1.5 dm. long, stout; rays 1-2 cm. long; bractlets linear or lance-linear, 5-6 mm. long; flowers apparently straw-colored or ochroleucous; fruit about 9 mm. long, 6 mm. wide; lateral wings about as broad as the body; dorsal ribs filiform or some of them narrowly winged; oil tubes 2-4 in the intervals, 4-6 on the com- missure, rather obscure. _ The fruit of this species is intermediate between that of C. Nuttallii and C. Parryi, but the plant differs from both, as well as from all the known species, in the broad segments of the leaves. 74 RYDBERG: Rocky MOUNTAIN FLORA The segments resemble those of some species of Cogswellia of the C. triternata group, but the leaves are pinnate, not ternate, the plant has the densely cespitose, sheath-covered caudex characteristic of Cynomarathrum, and the fruit is of that genus, having some of the dorsal ribs winged, and the calyx-teeth are prominent. It grows on sides of canyons at an altitude of 1,600 m. UtaH: Proposed dam site, near Wilson Mesa, Grand County, Utah, July 1, 1911, Rydberg & Garrett 8371 (fruit; type, in herb. N. Y. Bot. Gard.); also 8414 (withered flowers). Cogswellia simplex (Nutt.) Rydb. Peucedanum triternatum platycarpum Torr. Stansb. Rep. 389. 1852. Peucedanum simplex Nutt.; S. Wats. Bot. King. Exped. 129. 1871. Lomatium platycarpum C. & R. Cont. U. S. Nat. Herb. 7: 226. 1900. Cogswellia platycarpa (Torr.) M. E. Jones, Cont. West. Bot. 12: 32. — 8. It was unfortunate that an amendment to the Rochester Code ever was passed at Madison, by which a varietal name could supersede a specific name, and I am glad that the amendment mentioned has been recalled and that we can return to the specific name well known by a long usage. Cogswellia leptophylla (Hook.) Rydb. sp. nov. Peucedanum triternatum leptophyllum Hook. Lond. Journ. Bot. 6: 235. 1847. This species is related to C. simplex, C. triternata, and C. robustior. In general habit, it resembles most the second, but the leaflets are narrower, the fruit is shorter and relatively broader and puberulent. C. simplex has less compound leaves, broader leaflets, larger and glabrous fruit; C. robustior has much broader and more spreading leaflets, longer fruit with very narrow wing. Montana: Helena, June-July, 1891, Kelsey; also May, 1890; University campus and hillsides, Missoula, 1901, MacDougal 130; Old Sentinel, June 12, 1901, MacDougal; Deer Lodge, June, 1888, Traphagen; Mt. Ascension, Helena, 1909, Butler 4057. IpaHo: Hills near Boise, June 7, 1892, Isabel Mulford; Weiser, April 18, 1900, M. E. Jones 6336. New York Botanical GARDEN. Tetradesmus, a new four-celled coenobic alga GILBERT MorGAN SMITH (WITH PLATE 1) METHODS AND MATERIAL The alga described below was cbtained during a study of various algae in pure cultures. The isolation of the different algae was by means of a medium consisting of 0.2% Knop’s (9) solution and 2.0% agar in Petri dish cultures. By this method cultures were obtained which were the descendants of a single cell. Great care was used in the isolation, and only those cultures that were entirely free from other algae and bacteria were used in this study. After the isolation, cultures were made in sterile Knop’s solution in 200 c.c. Erlenmeyer flasks. A complete description of the methods used will be given in a forthcoming paper. The details of the cellular structure were studied in material which had been fixed in Flemming’s weak osmic-acetic-chromic acid mixture diluted with an equal volume of water. For the washing and dehydration of the material a modification of Oster- hout’s (12) method was used. After the material had been al- lowed to stand in the killing solution for 24 hours, as much as possible of the solution was removed from the vial by means of a pipette, and then the vial was filled with distilled water. A celloidin film was prepared by pouring a celloidin solution on a clean surface of mercury and allowing it to harden so that it could be lifted up and placed over the mouth of the vial. The membrane was then allowed to harden still further while on the mouth of the vial. The material was then washed by simply placing it in a dish containing running water. For the purpose of dehydration the vial was transferred to 70 per cent. alcohol for 12 hours and then to two successive portions of 95 per cent. alcohol, at intervals of 12 hours. By means of this method the loss of material through decantation was obviated. The material was imbedded in paraffin and cut on a microtome into sections of from 3 to 5 # in thickness. ' 75 76 SMITH: TETRADESMUS, A NEW COENOBIC ALGA The triple stain of Flemming gave the best differentiation, and all drawings are from preparations made in this manner. I wish to express my thanks here to Professor Charles E. Allen for kind criticism and keen interest shown during the progress of this work and preparation of the manuscript. DIAGNOSIS The alga in question normally occurs in the form of four-celled colonies. The characteristic feature of the arrangement of the cells is that when viewed from the side they are seen to be in two tiers, while in Scenedesmus, apparently the most nearly related form, the cells are all in a single plane. The shape of the cells varies somewhat with their age, the mature ones being more ovoid. The following are the descriptions of the genus and species: Tetradesmus gen. nov. Colonies free, of 4 cells, rarely 1 or 2; cells in two planes, 2 cells in each plane, joined along the longer axes, ovoid with pointed ends; chlorophyl present throughout the cell, pyrenoid single. Repro- duction by autocolonies inside of old cell wall, liberated by rupture of mother cell wall. Coenobia segregata, e cellulis quaternis (rarius 1-2). Cellulae binae latere longiore in seriem duplicem conjunctae, ovoideae utroque polo acutae, homogeneae chlorophyllosae, et pyrenoide singulo praeditae. Propagatio fit autocoénobiis intra cellulam matricalem quae membranae ruptura prodeunt. Tetradesmus wisconsinensis sp. nov. Cells ovoid with sharply pointed ends, 4-5.8 by 12-14.5 mu. Hasitat: Floating in sluggish streams and lakes; Madison, Wisconsin. Cellulae ovoideae utroque polo acutae, 4-5.8 w X 12-14.5 mp. Hasirat: In rivulis lente fluentibus et lacubus libere natantes. Madison, Wisconsin. The characters described above are sufficient, in my opinion, to warrant the assumption that we have a new genus. The objection may be made that this is merely a cultural form of enedesmus acutus Meyen; for according to Chodat (3), Chodat SMITH: TETRADESMUS, A NEW COENOBIC ALGA 77 and Malinesco (4) (5), and Grintzesco (6), Scenedesmus acutus shows great variation under different cultural conditions. The regular arrangement of the cells in a linear series may disappear and the cells become isolated or arranged in the branching chain- like colonies which Naegeli (11) has described under the name of Dactylococcus. Although not at all conclusive, the fact is worthy of mention here that these investigators found no variation of Scenedesmus acutus which looked at all like Tetradesmus. ' The alga in question has been under constant observation for over nine months and cultivated in different media, such as Knop’s solution in different concentrations, Knop’s solution with glucose or cane sugar, Beyerinck’s (2) solution, and Knop’s solution with 0.2-2% sodium chlorid. It has been cultivated under different conditions of illumination and temperature, and under none of these conditions has a colony been found with the cellular arrangement characteristic of Scenedesmus acutus. Con- versely the evidence is just as strong, for I have had several different strains of Scenedesmus acutus under observation for the same length of time and under the same differing conditions, and in these cultures forms resembling Tetradesmus have never been found. If Tetradesmus were a cultural form of Scenedesmus acutus, the change from one form to the other should have been observed in this extensive series of cultures. In the size, shape, and structure of the individual cells the two species also show constant differences. The cells of Tetradesmus wisconsinensis vary in size from 4—5.8 u in width when the alga is grown in 0.1% Knop’s solution; those of Scenedesmus acutus under the same conditions are 6.2—4.5u in width. When the alga is grown in a 1.0% Knop’s solution, the average diameter of the Scenedesmus cell increases to 6-7.8 uw, that of the Tetradesmus cell remaining practically unchanged. With the change to the stronger solution there is no change in the length of the individual cells. This change from the acicular to the ovoid-round shape of the cell in Scenedesmus acutus, when the concentration of the medium is increased, has been already pointed out by Senn (14). On the other hand, Tetradesmus shows no appreciable change in the shape of the cells when the concentration of the nutritive solu- tion is increased. Beyerinck (1) and Senn (14) have shown that 78 SMITH: TETRADESMUS, A NEW COENOBIC ALGA in weaker nutritive solutions the chromatophore of Scenedesmus acutus does not fill the entire cell but that there is a hyaline region at one side. My experiments with Scenedesmus acutus confirm this point; however, the chromatophore in Tetradesmus always completely fills the cell. The pyrenoid of Scenedesmus is con- stantly somewhat larger than that of Tetradesmus, being about 2.5 uw in diameter in the former and 1.5 w in the latter. The arrange- ment of the starch grains differs somewhat, the characteristic segmentation of the grains around the pyrenoid being easily seen in Scenedesmus and only with great difficulty in Tetradesmus. In the older cultures of the former species the cell is usually filled with characteristic angular pieces of ‘‘stroma’”’ starch that have been derived from the pyrenoid; in Tetradesmus kept under similar conditions the ‘“‘stroma”’ starch granules are not very abundant. A good idea of the cellular structure of Scenedesmus acutus may be obtained from FIG. 3. Among other coenobic algae the forms which appear to re- semble Tetradesmus most closely are Selenastrum Reinsch (17) and Lauterborniella Schmidle (13). Selenastrum, as described by West (17), differs from Tetradesmus in that there are more than four cells in some of the colonies, and that the individual cells do not possess a pyrenoid. Dr. Schmidle has kindly compared my material with his type specimens of Lauterborniella and informs me that neither in size, shape, nor arrangement of the cells does Tetradesmus resemble Lauterborniella. MorPHOLOGY The individual cells of the Tetradesmus colony are always more or less ovoid, but in every case they are sharp-pointed at the ends. The cell wall is composed of two parts, as Chodat and Malinesco (4) have shown for Scenedesmus. The inner part is of cellulose and reacts to zinc chloriodid; the outer is composed of a gelatinous material which serves to bind the cells of the colony together. The existence of this outer gelatinous layer can be demonstrated by the use of some such stain as Bismarck brown. There is no central vacuole, but the entire cell except for the nucleus and the pyrenoid is filled with a cytoplasm which is granular or rarely slightly alveolar in structure. The chlorophyl is distributed SMITH: TETRADESMUS, A NEW COENOBIC ALGA 1g throughout the cytoplasm and is not in the form of a definite chromatophore. Every cell possesses a single, eccentrically placed pyrenoid. In preparations stained with the safranin-gentian violet combination, the pyrenoid takes on a brilliant red coloration and the surrounding layer of starch is colored blue. The pyrenoid appears to be an entirely homogeneous body and not composed of lighter and darker regions, as Lutman (10) has shown to be the case in Closterium. On the other hand, the minuteness of the pyrenoid may possibly account for the failure to observe these details in its structure. A method devised by E. M. Gilbert, but as yet unpublished, in which a differentiation of the structure of the pyrenoid of Sphaeroplea was obtained by the use of safranin and Lichtgriin, was also tried, but no differentiation in the pyrenoid of Tetradesmus was observed. Surrounding the pyrenoid is a hyaline region which extends from the surface of the pyrenoid to the starch grains. This possibly may be a region that is com- posed of intermediate compounds in the formation of the starch grains, but this region failed to take any of the different stains used, although Timberlake (15) found that at times a part of it (in Hydrodictyon) stained with orange G. The starch granules surrounding the pyrenoid form a layer which is circular in outline and not angular as in Hydrodictyon (Timberlake) and Closterium (Lutman). Fic. 6 shows that there is some variation in the width of the starch layer and also that the outline of the starch ring may approach the ovoid but never the angular. The drawings also bring out the fact that the pyrenoid is not always in the center of the starch ring but may be towards one side. The starch layer is not a single unbroken ring but is composed of curved plates. Fic. 8 shows this segmentation of the starch layer. The deter- mination of this point is an exceedingly difficult matter, although it may be noted again that the individual starch plates in Scenedes- mus are easily observed. In the case of some of the older cells starch granules may be also found in the cytoplasm some distance from the pyrenoid (FIG. 7). These granules have been given the name of “stroma” starch by Klebs (8), but Timberlake (15) has shown that they are of pyrenoidal origin. The existence of the ‘‘stroma”’ starch may be taken as evidence of a somewhat patho- logical condition, since it is never found (at least in 7 etradesmus) 80 SmiTH: TETRADESMUS, A NEW COENOBIC ALGA in any except the old, yellow-colored cultures. The ‘‘stroma’”’ starch is also produced in much larger quantities in the Scene- desmus cell than it is in the Tetradesmus cell. The nucleus is usually at the inner side of the cell (FIG. 1), but it as well as the pyrenoid may lie in the long axis of the cell (FIG. 2). Frequently the nucleus and the pyrenoid are symmetrically placed in the long axis of the cell. There is always a single nucleole within the nucleus. The chromatin seems to have a very fine granular structure, and the only apparent difference between the cytoplasm and the non-nucleolar part of the nucleus is the greater density of the latter when the material is stained with Heiden- hain’s iron-alum-hematoxylin. REPRODUCTION Reproduction always takes place by the formation of auto- colonies within the mother cell wall. The formation of zoospores or motile gametes was never observed. In this respect the form resembles Scenedesmus, whose sexual reproduction is unknown. The division of the nucleus was not observed with certainty, on account of the minuteness of the nucleus, but stages in the cleavage of the cell were found in sufficient abundance to warrant the assumption that the following is the correct description of the process. The nucleus divides so that the daughter nuclei lie in a line parallel to the long axis of the cell, as shown by a cell containing two daughter nuclei (FIG. 9). This stage was found in great abundance, and the two daughter nuclei were always found at the inner side of the cell. Following this division of the nucleus comes a cleavage of the cytoplasm, so that each half of the protoplasmic mass contains a nucleus (FIG. 104 and 10B). This division takes place by the formation of a cleavage furrow from the outside toward the center. The process seems to be similar to that observed by Harper (7) in the spore formation of certain fungi and by Timberlake (16) in the swarm spore formation of Hydrodictyon. The daughter cells resulting from this division are two naked protoplasts which lie close to one another and are not separated by a wall. The pyrenoid does not divide during this process; consequently, at the end of the first cleavage one daughter SMITH: TETRADESMUS, A NEW COENOBIC ALGA 81 protoplast contains a pyrenoid and one does not. This fact can be determined with the greatest accuracy since the staining re- action of the pyrenoid makes it the most conspicuous structure in the cell. Hundreds of cells at this stage of development have been observed, and in every case the pyrenoid was easily distinguished in one of the daughter protoplasts while the other was empty. At first this cleavage without a division of the pyrenoid was thought to be an abnormality, but the abundance of cells in this condition and the failure to find a single case in which a pyrenoid appeared in each of the daughter protoplasts show that this is the normal condition. The first plane of cleavage is usually approximately at right angles to the long axis of the cell (FIG. 10A), but soon after the cleavage the daughter protoplasts arrange themselves inside of the mother cell wall so that the original cleavage plane appears to be diagonal (FIG. 10B). The stimulus which leads to this twisting of one of the protoplasts upon the other is not known. A second division of the daughter nuclei now occurs. Fic. 11 shows this fully completed, the direction of the division, to judge by the position of the resting nuclei, always being parallel to the primary cleavage plane. There is some variation in the time at which this second division takes place; in some cases it occurs when the primary cleavage plane is only a few degrees from the transverse position, but in others the primary plane has come to lie at an angle of 45 degrees with its original position before the second nuclear division occurs. Fic. 11 shows that after. the second nuclear division a single pyrenoid is still in evidence in one of the protoplasts while there is none in the other. The amount of starch surrounding the pyrenoid is approximately the same as that in the undivided cell, but this is not brought out in the figures, since material which is well stained for the nuclear structures does not show the starch about the pyrenoid. Fic. 10B, from material which was stained especially to bring out the pyrenoid, shows that after the first cell cleavage there is still considerable starch about the pyrenoid. Following the second nuclear division there is a cleavage in each of the protoplasts. These cleavages seem to be usually simul- taneous, but FIG. 12 shows a case in which the cleavage has been 82 SMITH : TETRADESMUS, A NEW COENOBIC ALGA completed in one of the daughter protoplasts and not in the other. The relation of the cleavage planes to one another shows that there are certain mechanical factors governing their position. It is well known [Wilson (20)] that in the cleavage of the animal egg changes often occur in the position of the cells with regard to one | another. Surface tension seems to be one of the factors governing this change.. In moss rhizoids, de Wildeman (18) found that the cross walls are always S-shaped in section, so that although the cross wall as a whole is placed diagonally, yet its junction with the outer walls of the rhizoid is at right angles. This curving of the cleavage planes where they meet the mother cell wall occurs in substantially the same manner in Tetradesmus (FIG. 13, 14). One view of the young four-celled colony, shortly after the second cleavage, when seen from the side, is shown by FIG. 13, but when the young colony is seen in a plane at right angles to that shown in this figure, we observe the very different appearance shown in FIG. 13A. FIG. 13 shows that even after the four proto- plasts of the young colony are formed, one and only one contains a pyrenoid. At this stage, or very shortly after, the old pyrenoid disappears (FIG. 14). This disappearance probably takes place quite rapidly, since no intermediate stages were found between those shown in FIG. 13 and 14. In no case was the pyrenoid recognizable at stages later than those of FIG. 13 and 14. The changes that follow in the young colony might be well said to constitute a period of maturation. The maturation consists in an elongation of each of the daughter protoplasts until it extends the entire length of the mother cell. The protoplasts do not elon- gate so as to lie in one plane, but the two in one (which we may call the lower) half of the cell slip up over the two in the upper half. This slipping occurs along the diagonal plane of the first cleavage- A stage in this slipping is seen in FIG. 15, although the drawing fails to bring out the fact that a portion of each of the two upper cells is hidden by part of the lower pair of cells. When this elongation is completed, the daughter cells are arranged in two planes, so that only two cells may be seen in any lateral view (FIG. 16). The growth of the four daughter cells seems not to be an equal elonga- tion of all parts, but rather an elongation of the end of each of the ; daughter cells farthest from the corresponding apex of the old SMITH: TETRADESMUS, A NEW COENOBIC ALGA 83 mother cell. Asa result the nucleus of each daughter cell is not found in the center of the cell but nearer the end that was originally in contact with the apex of the mother cell wall. Fic. 15 shows this fact, but it must be borne in mind that a portion of the upper pair of cells is concealed by the lower ones. At the beginning of this elongation there is no pyrenoid present in any of the daughter protoplasts (FIG. 15). The further steps in the maturation of the daughter cells consist in the formation of pyrenoids and cell walls. The daughter cells are without pyrenoids until they have reached their full length, and then a pyrenoid suddenly appears in each (FIG. 16). Such an origin of pyrenoids de novo is contrary to the current conception, according to which a pyrenoid always arises by the division of a preéxistent pyrenoid but is in harmony, as will be later pointed out, with Timberlake’s observations in Hydrodictyon. The formation of the new pyrenoid possibly may be a result of the metabolic activities of the nucleus, since the pyrenoid at its first appearance is often very close to the nucleus (FIG. 17). In other cases, possibly representing later stages, the pyrenoid is at a greater distance from the nucleus (FIG. 16). At about the same time as the appearance of the pyrenoid a cell wall is formed around each of the four protoplasts while they are still within the old cell wall. The liberation of the daughter colony is through a longitudinal break in the mother cell wall. End views, such as those shown in FIG. 19 and 20, are especially favorable for studying the relation of the daughter coenobe to the mother cell wall. The opening through which the young colony escapes is a fissure that extends the whole length of the mother cell wall (Fic. 18). The cause of this rupture is probably the growth of the young colony within the old wall. It is not an irregular break but a definite longitu- dinal split, which is always at the side turned away from the other cells of the mother colony. After liberation the young coenobe is composed of cells which are of nearly full length but which are much more acicular than the mature cell (FIG. 17). The further history of the coenobe consists in the growth of the individual cells until they have assumed the shape characteristic of the adult stage. 84 SMITH : TETRADESMUS, A NEW COENOBIC ALGA DISCUSSION Tetradesmus belongs to the family Coelastraceae, recently estab- lished by Wille (19). Tetradesmus shows a close relationship to Scenedesmus. The method of reproduction is essentially the same as that which occurs in Scenedesmus, the chief difference being that at the time of liberation from the mother cell wall the cells forming the coenobe in Scenedesmus become arranged in the form of a plate, while in Tetradesmus they remain rolled up. It seems probable that one form has been derived from the other, but there is little evidence as to the direction that the phylo- genetic development has taken. There is also the possibility that these two forms have been derived from a common ancestor. Tetradesmus may have been derived from Scenedesmus through a loss of the tendency for the cells of the coenobe to unroll and spread out into a linear series after their liberation from the mother cell wall; or some colony of Tetradesmus may have developed this character of unrolling the cells in order to obtain an arrangement of the individual cells better adapted for photosynthesis. Cytological study shows that these forms (Tetradesmus and Scenedesmus) are more closely related to Hydrodictyon and Pedias- trum than has been usually supposed. In the last-named forms reproduction takes place by the formation of a more or less definite number of zcéspores, which then become arranged in the form of the mature colony while they are still inside the old mother cell wall or inside a gelatinous vesicle. In Tetradesmus and Scenedes- mus there is a successive division into a definite number of uni- nucleate masses of protoplasm. These correspond to the zodspores of Hydrodictyon and Pediastrum in every way except for the fact that they have no power of movement. We may easily think of these daughter cells in Tetradesmus and Scenedesmus as zodspores which have lost the power of movement. The loss of power of motility of the zodspores inside the mother cell wall is probably a step in advance of the condition in Hydrodictyon. There is also the possibility that this loss of movement of the zoospores is connected with the smaller number of spores produced and the greater ease with which they get into their permanent positions. pepeinetion in Tetradesmus is completed by a movement of the zc6spores”’ into the position which they will occupy in the SMITH : TETRADESMUS, A NEW COENOBIC ALGA 85 mature colony, and then by the liberation of the young colony from the mother wall. Another similarity between the young daughter cells in Tetra- desmus and the swarm spores in Hydrodictyon is in the behavior of the pyrenoid. In Hydrodictyon, Timberlake (16) noted that after the swarm spores are formed by cytoplasmic cleavage, new pyrenoids appear as small red-staining bodies closely associated with the nuclei. This is exactly what takes place in Tetradesmus and gives us a further basis for considering the young daughter cells of Tetradesmus the morphological equivalents of the swarm spores of Hydrodictyon. As in Hydrodictyon also, the daughter cells at first have no cell wall, but a cellulose wall is formed after the daughter cells have settled down into the position which | they will have in the adult colony. SUMMARY 1. Tetradesmus is a distinct genus, and not a cultural form of Scenedesmus. 2. Reproduction takes place by the formation of non-motile cells (spores) by successive nuclear and cell divisions. These spores then become arranged in the form of the adult colony while they are still inside the old cell wall. The young colonies are liberated by the rupture of the mother cell wall. 3. The pyrenoids in the daughter cells a arise de novo and not by the division of a preéxisting pyrenoid.. UNIVERSITY OF WISCONSIN. BIBLIOGRAPHY . Beyerinck, M. W. Culturversuche mit Zoochlorellen, Lichen- gonidien und anderen niederen Algen. Bot. Zeit. 48: 725. 1890. . Beyerinck, M. W. Notiz iiber Pleurococcus vulgaris. Centralbl. Bakt., Zweite Abt. 4: 785. 1898. Chodat, R. Etude critique et expérimentale sur le polymorphisme Mémoire publié a l'occasion du jubilé de |’Univer- La tv > des algues. sité de Genéve. 1909. Chodat, R., & Malinesco, O. Sur le polymorphisme de Scenedes- mus acutus Meyen. Bull. Herb. Boiss. 1: 184. 1893. Chodat, R., & Malinesco, O. Sur le polymorphisme du Raphidium Braunii et du Scenedesmus caudatus Corda. Bull. Herb. Boiss. I: 640. 1893. > i a “¢ ~I e2) \o ~ to ~J taal Co Ny -) SMITH: TETRADESMUS, A NEW COENOBIC ALGA Grintzesco, J. Recherches expérimentales sur la morphologie et la physiologie de Scenedesmus acutus Meyen. Bull. Herb. Boiss. Ly 34.297: 1902: . Harper, R. A. Cell division in sporangia and asci. Ann. Bot. 13: 467. 1899. . Klebs, G. Fortpflanzungszellen bei Hydrodictyon utriculatum Roth. Bot. Zeit. 49: 789. 1891. . [Knop]. Reference in Kiister, Kultur der Mikroorganismen. Leipzig, 1907. . Lutman, B. F. The cell structure of Closterium Ehrenbergii and C. moniliferum. Bot. Gaz. 49: 241. 1910. . Naegeli, C. Gattungen einzelliger Algen. 1848. . Osterhout, W. J. V. Contributions to cytological technique. V. Embedding microscopic algae. Univ. Calif. Publ. Bot. 2: 85. 1904. Schmidle, W. Beitrage zur Kenntniss der Planktonalgen. Ber. Deut. Bot. Gesell. 18: 144. 1900. . Senn, G. Ueber einige koloniebildende einzellige Algen. Bot. Zeit. 57: 39. 1899. . Timberlake, H.G. Starch-formation in Hydrodictyon utriculatum. Ann. Bot. 15: 619. 1901. . Timberlake, H.G. Development and structure of the swarm-spores of Hydrodictyon. Trans. Wisconsin Acad. Sci. Arts and Lett. 13: 486. 1902. . West, G. S. A treatise on the British freshwater algae. Cam- bridge, 1904. . Wildeman, E. de: Etudes sur ]’attache des cloisons cellulaires. Mém. cour. et mém. des sav. étrang., Acad. Roy. Sci. Lett. et Beaux-arts de Belg. 53: 1. 1893. - Wille, N. Conjugatae und Chlorophyceae. Engler und Prantl, Die Natiirlichen Pflanzenfamilien. Nachtrage zum I Teil, 2 Abt., 236 und 237 Lief. 1911. . Wilson, E. B. The cell in development and inheritance. Ed. 2- N. Y.. 1906. Explanation of plate 1 All figures, with the exception of Fic. 3, are of Tetradesmus wisconsinensis- They were drawn with the aid of the Abbe camera lucida, the drawing being at the level of the base of the microscope, and with the Leitz objective 1/16 and ocular 4- The magnification is about 2,200 diameters. Fic. 1. Longitudinal section of colon Fic. 2. Surface view of colony, showing arrangement of cells. Fic. 3. Colony of Scenedesmus acutus Meyen grown under the same cultural conditions as the colonies of Tetradesmus shown in FIG. I and 2. SMITH: TETRADESMUS, A NEW COENOBIC ALGA 87 Fic. 4,5. Transverse sections of colonies. - 6. Various forms taken by the pyrenoid and the surrounding starch layer Fic. 7. Cell showing ‘‘stroma” starch granules. F - Cell showing the pyrenoid with segmented starch ring. Fic. 9. A cell after the first nuclear division. Fic. 10A, 10B. Completion of first cell division. Fic. 11. After the completion of the second nuclear division. - 12-14. Stages in the second cleavage of the cytoplas IG. I5, 5A. Beginning of elongation of the four daughter ‘tlle Young colony inside of mother cell wall (lateral view). IG. I7. Young colony, immediately after liberation from mother cell. Empty mother cell walls after liberation of young colonies. End view of daughter colony, showing method of its liberation from mother cell walk Fic. Transverse section of coenobe, showing young colony within the wall of one of the mother cells. INDEX TO AMERICAN BOTANICAL LITERATURE (1912) aim of this Index is to include all current botanical literature written by Americans, published in America, or based upon American material ; the word Amer- ica being used in the broadest sense Reviews, and papers that vilicts exclusively to forestry, agriculture, neon manufactured products of vegetable origin, or laboratory methods are not i n no attempt is made to index the literature of bacteriology. An occasional cient is made in favor of some paper — in an American periodical which is devoted wholly to botany. Reprints are not mentioned unless they differ from the original in some important particular. If users of the Index will call the attention of the editor to errors or omissions, their indueas will be appreciated. This Index is reprinted monthly on cards, and furnished in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitted ; each a must take all cards published during the term of his subscription. Corre- ndence relating to the card issue should be ane to the Treasurer of the Torrey aoa! Club Allard, H. A. The mosaic disease of tobacco. Science II. 36: 875, 876. 20 D 1912. Anderson, P. J., & Anderson, H. W. Endothia virginiana. Phyto- pathology 2: 261, 262. D 1912. Atkinson, G. F. Gautieria in the eastern United States. Bot. Gaz. 54: 538, 539. 16 D 109012. Atkinson, G. F. The perfect stage of the Ascochyta on the hairy vetch. Bot. Gaz. 54: 537, 538. 16 D 1912. ; Bailey, W. W. The glory of the morning. Am. Bot. 18: 101-103. N 1912. Barrett, J. T. The development of Blastocladia strangulata, n. sp.. Bot. Gaz. 54: 353-371. pl. 18-20. 13 N 1912. Bartholomew, E. T. Apple rust controllable by spraying. Phyto- pathology 2: 253-257. D 1912. Bean, W. J. Pinus flexilis. Curt. Bot. Mag. IV. 8: pl. 8467. D 1912. A plant from western North Ameri Berry, E. W. Notes on the scabies history of rt walnuts and hickories. Plant World 15: 225-240. f. 1-4. O1 Berry, E. W. Notes on the present status of nk Plant World 15: 169-175. Au ae 90 INDEX TO AMERICAN BOTANICAL LITERATURE Bitter, G. Solana nova vel minus cognita. IV. Repert. Sp. Nov. 1: 241-260. 25 N 1912. Includes four new species from South America Blumer, J. C. Notes on the phytogeography of the Arizona desert. Plant World 15: 183-189. Au 1912. Bouyoucos, G. Transpiration of wheat seedlings as affected by dif- ferent densities of a complete nutrient solution in water, sand, and soil cultures. Beih. Bot. Centralb. 29: 1-20. 15 N 1912. Bower, F. O. The quest of phyletic lines. Plant World 15: 97-109. My 1912. Brooks, A. J. Artificial cross-fertilization of mango. West Ind. Bull. 12: 567-569. 18S 1912. Brown, H. P. Growth studies in forest trees. I. Pinus rigida Mill. Bot. Gaz. 54: 386-403. pl. 24, 25. 13 N 1912. Brown, W.H. The relation of evaporation to water content of the soil at the time of wilting. Plant World 15: 121-134. Je 1912. Chamberlain, C. J. Two species of Bowenia. Bot. Gaz. 54: 419-423- Fe gr me Be es Chamberlain, E. B. A Sedum new to North America. Rhodora 14: 227, 228. 9 D 1912. Choate, H. A. The origin and development of the binomial system of nomenclature. Plant World 15: 257-263. N 1912. Christensen, C. On the ferns of the Seychelles and the Aldabra group. Trans. Linn. Soc. Bot. 7: 409-425. pl. 45. N 1912. Clausen, R. E. A new fungus concerned in wither tip of varieties of Citrus medica. Phytopathology 2: 217-234. pl. 21, 22 +f. } D 1912. Clinton, G. P. Chestnut blight fungus and its allies. Phytopathology 2: 265-269. D 1912. in ase aT ack ere Te Clinton, G. P. The relationships of the chestnut blight fungus. E Science IT. 36: 907-914. 27 D 1912. [Clute, W. N.] Characteristics of our forest trees. Am. Bot. 18: 103-109, N 1912. Clute, W. N. The summer flora of the Chicago Plain. Am. Bot. 18: 97-100. N i912. [Illust.] Coker, W. C. The plant life of es S.C. 1-129. pl. Columbia. 1912. Conard, H.S. The Kellerman plant press. Plant World 15: 1357139 Je 1912. Cooper, W.S. The ecological succession of mosses, As illustrated upo? | 4 Q 4 Isle Royale, Lake Superior. Plant World 15: 197-213. f: 1-6. IQI2. ‘ INDEX TO AMERICAN BOTANICAL LITERATURE 91 Craighead, F. C. Insects contributing to the control of the chestnut blight disease. Science II. 36: 825. 13 D 1912. Dachnowski, A. The nature of absorption and tolerance of plants in bogs. . Bot. Gaz. 54: 503-514. 16 D 1912. Deane, W. A further note on Euphorbia oe sacs de in fruit. Rho- dora 14: 193-196. O 1912. Dewey, L. H. Ramie. U.S. Dept. Agr. Plant Ind. Circ. 103: 3-9. i R2 3 Bor. Dowell, P. A suggestion on the field study of ferns. Am. Fern Jour. 2: 123.18 -D 1912. Evans, A. W. New West Indian Lejeuneae—Ii. Bull. Torrey Club 39: 603-611. 31 D 1912 Includes pease Johnsonii, Leptocolea appressa, and Reciolejeunea Maxonii spp. no Fedde, F. Nite Arten aus der Verwandschaft der Corydalis aurea Willd. von Nord-Amerika. VI. Repert Sp. Nov. 11: 196, 197. 20 O 1912; VII. Repert. Sp. Nov. 11: 289-291. 25 N 1912. _ Includes descriptions of five new species. Fernald, M. L., & Wiegand, K. M. *A northeastern variety of Chelone glabra. Rhodora 14: 225, 226. 9 D 1912. Fritsch, K. Gesneriaceen-Studien. Bot. Zeits. 62: 406, 407. N 1912. Includes Besleria salicifolia sp. nov. from Colombia. Fuller, G. D. Evaporation and the oO of vegetation. Gaz. 54: 424-426. f. 1. 13 N1 Fyles, F. First record of Ste feat Lalebaus + tit Cana: Nat. 26: 116. 18 D 1912. Gager, C. S. Ingrowing sprouts of Solanum tuberosum. 515-524. pl. 36 +f. 1-6. 16 D 1912. Gerard, W. R. Trees that yield butter. 1912. From the Scientific American. Giddings, N. J., & Neal, D. C. Control of apple rust by spraying. paren cd 258-260. pl. 23, 24. D 1912. A preliminary repo: Goodspeed, T. H. Qhantitative studies of inheritance in Nicotiana hybrids. Univ. Calif. Pub. Bot. §: 87-168. pl. 20-34. 6 D 1912. Greene, E. L. Five dwarf lupines. Muhlenbergia 8: 117-119. 7 D Bot. Ottawa Bot. Gaz. 54: Am. Bot. 18: 109-112. N 1g12. Greenman, J. M. Diagnoses of new species and notes on other sperma- tophytes, chiefly from Mexico and Central America. Field Mus. Nat. Hist. Bot. 2: 329-350. 20 D 1912. 92 INDEX TO AMERICAN BOTANICAL LITERATURE Greenman, J.M. New species of Cuban Senecioneae. Field Mus. Nat. Hist. Bot. 2: 323-328. 20 D 1912. Ten new species are described. Gregory, C. T. Spore germination and infection with Plasmopara viticola. Phytopathology 2: 235-249. f. 1-7.- D 1912. Grossenbacher, J. G. Crown-rot of fruit trees: field studies. N. Y. Agr. Exp. Sta. Tech. Bull. 23: 3-59. pl. 1-23. S 1912. Harper, R.M. The Altamaha Grit region in December. Plant World 15: 241-248. O 1912. Harper, R. M. The Hempstead Plains of Long Island. Torreya 12: 277-287. f. 1-7. 13 D 1912. Harris, J. A. A simple test of the goodness of fit of Mendelian ratios. Am. Nat. 46: 741-745. D 1912. Hassler, E. Aristolochiaceae. [In Ex herbario Hassleriano: Novitates paraguarienses. XV.] Repert. Sp. Nov. 11: 176-178. 20 O 1912. Includes Aristolochia glaberrima sp. nov. Hassler, E. Compositae. [In Ex herbario Hassleriano: Novitates paraguarienses. XV.] Repert. Sp. Nov. 11: 165-175. 20 O 1912. Includes new species in Stevia (§), Eupatorium (7), and several new varieties. » Hedrick, U. P., & Wellington, R. An experiment in breeding apples. ‘ N. Y. Agr. Exp. Sta. Bull. 350: 141-186. pl. 1-17. Je 1912. Heller, A. A. The California white fir. Muhlenbergia 8: 121-131. pl. 13-15 +f. 24-26. 27 D 1912. Abies Lowiana Murray. Heller, A. A. The North American lupines—IX. Muhlenbergia 8: 109-116. f. 19-22. 7 D 1912. Includes four new species. Heller, A. A. A small flowered Mimulus. Muhlenbergia 8: 132. 27 D 1912 Mimulus micranthus sp. nov. Hill, E. J. The sand plum in Indiana. Rhodora 14: 196-198. O rise runus Watsoni (Sarg.) Waugh. a ahine ck, A. S. A new species of Andropogon. Bot. Gaz. 54: 424. 13 N 1912. Andropogon Urbanianus Hitchcock. Hofmann, J. V. Aerial isolation and inoculation with Pythium deBaryanum. Phytopathology 2: 273. D 1912. Hopkins, L. S. Further notes on the fern flora of Ohio. Am. Fern Jour. 2: 115-119. 18 D 1912. Howe, R. H. A monograph of the North American Usneaceae. Ann. Rep. Missouri Bot. Gard. 23: 133-146. pl. 7.. 16 D 1912. INDEX TO AMERICAN BOTANICAL LITERATURE 93 Hunnewell, F. W. Galium trifidum at Wellesley, Massachusetts. Rhodora 14: 205, 206. O 1912. Jennings, O. E. A note on the northwestern distribution of the sugar maple. Ottawa Nat. 26: 117, 118. 18 D 1912 Jones, W. R. The digestion of starch in germinating peas. Plant World 15: 176-182. f. 1-7. Au 1912. Kennedy, P. B. Alpine plants—XIII. Muhlenbergia 8: 119, 120. J. 27." 97 D' i912: Kingman, C. C. An American station for Illecebrum verticillatum. Rhodora 14: 207, 208. O 1912. Lamb, W. H. The phylogeny of grasses. Plant World 15: 264-269. N 1912. [IIlust.] Lamb, W. H. A tricarpellary walnut. Torreya 12: 290, 291. f. I. 13 D 1912. Lipman, C. B. The distribution and activities of bacteria in soils of the arid region. Univ. Calif. Pub. Agr. 1: 1-20. 15 O 1912. © Lipman, C. B., & Sharp, L. T. Toxic effects of “alkali salts’’ in soils on bacteria. III. Nitrogen fixation. Centralb. Bakt. Zweite, Abt. 35: 647-655. 15 N 1912. Lipman, J. G., Blair, A. W., Owen, I. L., & McLean, H. C. Factors relating to the availability of nitrogenous plant foods. New Jersey Agr. Exp. Sta. Bull. 251: 3-55. pl. 1-7 +f. 1. 1 Jl 1912. Livingston, B. E. A rotating table for standardizing porous cup atmometers. Plant World 15: 157-162. Jl 1912. [Illust.] Livingston, B. E. A schematic representation of the water relations of plant, a pedagogical suggestion. Plant World 15: 214-218. S 1912. Long, B. Galinm labradoricum in Pennsylvania. Rhodora 14: 199, 200. O 1912. Loomis, M. L. A seedless barberry found at Sherborn, Massachusetts. Rhodora 14: 207. O ea) Berberis vulgaris var. asper Luisier, A. Esbogo de ckaantods brazileira. Broteria 10: 141-172. D 1912 Mastastans: J. M. The relation of plant eeptephans to its environ- ment. Jour. Acad. Nat. Sci. Philadelphia. II. 15: 251-271. 21 Mr 1912. Massey, A. B. Notes on the flora about Clemson College. Charleston Mus. 8: 59-62. N 1912 McAvoy, B. The reduction division in yada, pl. I, 2. 8 N 1912. Bull. Ohio Nat. 13: 1-18. . 94 INDEX TO AMERICAN BOTANICAL LITERATURE Merrill, E. D. New or noteworthy Philippine plants, IX. Jour. Philip. Sci. 7: (Bot.) 259-357. N 1912. Includes two new genera and one hundred new species. Meyer, R. Lchinocactus myriostigma S.-D. var. nuda R. Mey. var. nov. Monats. Kakteenk. 22:.136, 137. 15 S 1912. Meyer, R. Einiges iiber Echinocactus texensis Hopff. Monats. Kak- teenk: 22: 167-169. 15 N 1912. Nelson, A. Contributions from the Rocky Mountain Herbarium. XII. New plants from Idaho. Bot. Gaz. 54: 404-418. 13 N 1912. Includes new species in Melica (1), Calochortus (1), Zygadenus (1), Salix (1), Eriogonum (1), Stellaria (1), Trifolium (1), Crataegus (1), Lupinus (1), Astragalus (2), Lathyrus (1), Viola (1), Cordylanthus (1), Pentstemon (1), and Artemisia (1). Osterhout, W. J. V. Plants which require sodium. Bot. Gaz. 54: 532-536. f. I, 2: 16 D: 1912. Overholts, L. O. Concerning Ohio Polyporaceae. Ohio Nat. 13: 22, 23. 8N 1912. Owen, M.L. Tillaea in Nantucket, Rhodora 14:-201-204. O 1912. Pelourde, F. Observations sur le Psaronius brasiliensis. Ann. Sci. Nat. Bot. IX. 16: 337-352. f. Ig. N- Agi2. Quehl, L. Mamillaria radicantissma Quehl spec. nov. Monats. Kak- teenk. 22: 164-167. 15 N 1912. [Illust.] Ransier, H. E. Outings for Onondaga moonwort and slender cliffbrake. Am. Fern. Jour. 2: 119-121. 18 D I9I2. Rau, E. A. Notes on the flora of Northampton County, Pennsylvania. Torreya 12: 287-289. 13 D 1912. Reed, H. S. Does Phytophthora infestans cause tomato blight? Phytopathology 2: 250-252. D 1912. ; Roberts, J. W. A new fungus on the apple. Phytopathology 2: 263, 264. Digit. Phomopsis Mali sp. nov. Robinson, W. J. A taxonomic study of the Pteridophyta of the Hawaiian Islands—II. Bull. Torrey Club 39: 567-601. pl. 40-44. Rugg, H. G. Whittier’s herbarium. Am. Fern Jour. 2: 121, 122. Safford, W. E. Notes of a naturalist afloat—V. Am. Fern Jour. 2: 97-107. 18D 1912. Sargent, C.S. A Connecticut station for Ilex mollis. Rhodora 14: 205. O 1912. Schaffner, J. H. An undescribed Equisetum from Kansas. Ohio Nat. 13: 19-22. 8 N 10912. Equisetum kansanum Schaffner Sp. nov. INDEX TO AMERICAN BOTANICAL LITERATURE 95 Schlechter, R. Die Orchidaceen-Gattungen Altensteinia H. B. et Kth., Aa. Rchb. f. u. Myrosmodes, Rchb. f. Repert. Sp. Nov. rr: 147-150. 200 1912. Schlechter, R. Orchidaceae novae et criticae. Decas XXXV. Re- pert. Sp. Nov. 11: 41-47. 1 Jl 1912. Includes seven new species from South America. Schreiner, O., & Skinner, J. J. Nitrogenous soil constituents and their bearing on soil fertility. U.S. Dept. Agr. Bureau Soils Bull. 87: 3-84. pl. I-11 +f. r.. 17 D 1912. ; Setchell, W. A. Studies in Nicotiana, I. Univ. Calif. Pub. Bot. 5: I-86. pl. 1-28. 5 D 1912. Shafer, J. A. Botanical explorations in Santa Clara and Oriente. Jour. N. Y. Bot. Gard. 13: 169-172. N 1912. Sharp, L. W. The orchid embryo sac. Bot. Gaz. 54: 372-385. pl. 2I- 23. 13 N 1912. Sherff, E. E. Range extensions of Rhamnus Frangula and Sporobolus asperifolius. Rhodora 14: 227. 9 D 1912. Shreve, F. Cold air drainage. Plant World 15: 110-115. My 1912. Skinner, J. J. Effect of solanine on the potato plant. Plant World 15: 253-256, ft.) ae dea. Smith, E. Etiology of crown galls on sugar beet. Phytopathology 2: 270-272. 1 1G: ; Spargo, M. The heat produced by leaves. Plant World 15: 277-293. D 1912. . Standley, P. C. 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Rev. 5: 593-597. pl. 6-9. N 1912. Wolf, F. A. A field method for sdieeiebaatieas certain orange stock. Alabama Agr. Exp. Sta. Cire. 17: 87-92. pl. I-3 +f. 1-6. Jl 1912. Wolf, F. A. Notes on the anatomy of Opuntia Lindheimeri Engelm. Plant World 15: 294-299. f. 1-ro. D IQI2. Woodward, N.P. An occurrence of Nicotiana rustica in Massachusetts. Rhodora 14: 206, 207. O 1912. Yamanouchi, S. The life history of Cutleria. Bot. Gaz. 54: 441-502. pl. 26-35 + f. 1-15. 16D 1912. Torrey. Crus: * Butt. P VOLUME 40, LATE I r Vol. 40 No. 3 BULLETIN OF THE TORREY BOTANICAL CLUB er ee - MARCH 1913 Development of the peristome in Ceratodon purpureus ALEXANDER W. EvaNs AND HENRY D. Hooker, Jr. INTRODUCTORY In his recent work on the Bryophyta, Cavers (11, p. 182) divides the Bryales, or true mosses, into the four following groups, based on distinctions emphasized by Fleischer: Tetraphidales, Polytrichales, Buxbaumiales, and Eu-Bryales. In this division the peristome is the organ from which the most important differ- ential characters are derived. In the Tetraphidales and Poly- trichales the teeth of the peristome are composed of entire cells; in the Buxbaumiales and Eu-Bryales they are composed of thickened cell walls. Leaving the Encalyptaceae out of con- sideration, the Eu-Bryales may be further divided, following the example of Philibert (’84, p. 67), into the Diplolepideae and the Haplolepideae.. The Diplolepideae with few exceptions have two peristomes, while the Haplolepideae have single peristomes, if such structures are present at all. In the Diplolepideae the two peristomes are derived from two concentric layers of cells in the opercular portion of the capsule. The development of the peristome has been described in several species of this group but is especially well known in Funaria hygrometrica (Goebel, ’87, p. 188; Campbell, ’05, p. 211) and Mnium hornum (Strasburger, 02, p. 491), which may be considered typical representatives. In both these species the outer per- istomial layer is composed of thirty-two longitudinal rows of cells and the inner of sixteen rows. These rows form sixteen groups, [The BuLtetin for February 1913 (40: 43-96) was issued March 18.] 97 98 Evans AND HooKER: PERISTOME IN CERATODON PURPUREUS each of which in cross section shows two cells of the outer layer lying opposite one cell of the inner layer. One of the sixteen teeth of the outer peristome arises in each group and is formed by the deposition of: longitudinal bands of thickening upon the periclinal walls between the peristomial layers. These bands taper to fine points from a broader base. The band deposited by the two outer rows is of fairly uniform thickness throughout and extends as a continuous layer across the thin radial and transverse walls separating the cells. The band deposited by the single inner row corresponds in position and in width with the one just described and is of about the same thickness. Thickening is deposited also upon the transverse walls separating the cells of this row. Upon the disappearance of the portions of the walls which have remained thin the teeth show their mature condition. Each tooth bears upon its inner surface a series of transverse ridges corresponding to the thickened transverse walls, while the smooth outer surface shows the vestiges of the transverse and radial walls, the latter appearing as a zigzag median line running the entire length of the tooth. The delicate inner peristome is formed by deposits of thickening upon the inner walls of the inner peristomial layer. Toward the base the thickening is continuous, but in the upper part it is in the form of isolated longitudinal bands, definite in position and in number. At maturity the continuous portion forms the basilar membrane of the peristome, while the isolated bands form the segments and the cilia. In the Haplolepideae the peristome is likewise derived from two concentric layers of cells, but in this case each tooth is formed by one row of outer cells and two rows of inner cells and therefore shows the zigzag longitudinal line on the inner surface instead of on the outer. In the opinion of Philibert (’88, p. 68), the single peristome of this group is homologous with the inner peristome of the* Diplolepideae. This being the case it is evident that the large-celled inner peristomial layer in the Diplolepideae would be homologous with the large-celled outer layer in the Haplolepideae. The subject of the present study, Ceratodon purpureus, is a characteristic member of the Haplolepideae, and many of the features which the peristome shows, in development as well as in structure, are undoubtedly common to other species of the group. . EVANS AND HOOKER: PERISTOME IN CERATODON PURPUREUS 99 The material, which was collected in the vicinity of New Haven, Connecticut, was studied by means of microtome sections. The mature peristome of Ceratodon purpureus has been repeatedly described in taxonomic treatises, but a brief account of its peculiarities will make the description of the development more intelligible. It consists of sixteen teeth which arise from a continuous basilar membrane. Each tooth is divided almost to the base into two slender tapering branches, which show much the same structure throughout their entire length (FIG. 17, 18). The outer portion of each branch forms a subcylindrical ridge, distinctly contracted at its union with the inner portion, which is broader and more flattened toward the base. Four or five corresponding transverse ridges extend across the branches of each tooth, being close together at the base but farther apart toward the apex. They correspond with the transverse walls of the outer peristomial layer. On the inner surface also vestiges of transverse walls can be detected, but these are unaccompanied by ridges (FIG. 19, 24). Just above the basal undivided portion of each tooth one or more of the transverse ridges extend across from branch to branch, thus leaving median perforations (FIG. 21). The basal portion is especially thick and bears several crowded transverse ridges, which in longitudinal section give the tooth a serrated appearance (FIG. 24). The entire surface of the teeth is covered over with minute spicules. The basilar membrane of the peristome is in the form of a hollow cylinder, sixteen cells in circumference and three or four cells high, representing a part of the outer peristomial layer. The thickened walls in this portion of the peristome are smooth throughout. The early development of the sporophyte in Ceratodon purpu- reus was studied by Kienitz-Gerloff (’78, p. 42), who detected in very young stages a two-sided apical cell cutting off two rows of segments, just as in other members of the Eu-Bryales. Each of these segments, as he further showed, soon became divided into quadrants by an anticlinal wall and then underwent divisions ace to the so-called ‘‘Grundquadrat” or ‘fundamental square” method described below. The embryonic development of Ceratodon is discussed also by Goebel (’87, p. 178), but the most complete account of the process is that published by Kuntzen (12), who, however, pays but little attention to the peristome. 100 Evans AND HOOKER: PERISTOME IN CERATODON PURPUREUS The first developmental study of the peristome in Ceratodon purpureus was made by Lantzius-Beninga (’50, p. 574), who com- pared it with the peristome of Trichostomum tortile, now known as Ditrichum tortile. In his figures of young transverse sections (50, pl. 66, f. 40, 41) he indicated the two layers of cells which give rise to the peristome but represented each layer as being composed of sixteen cells, those of the inner layer alternating with those of the outer. This was proved to be incorrect by Kienitz-Gerloff (’78, p. 44), who stated that, while the outer layer showed sixteen cells in section, the inner showed from twenty to twenty-four cells and that normally two cells of the outer layer bounded three cells of the inner. He showed further that the peristome developed in the amphithecial portion of the sporophyte and that the peristomial cells represented the fourth and fifth layers from the outside except in the region of the operculum, where they represented the third and fourth layers. The studies of Lantzius-Beninga and Kienitz-Gerloff have to do mainly with the region where the peristome is formed and give but few details about the actual development of the teeth, and the same thing is true of most of the work which has been done upon the peristomes of other Haplolepideae. In the present paper the account of the peristome will be divided into two parts. In the first the origin and development of the two peristomial layers will be described; in the second the deposition of the thickenings which constitute the bulk of the teeth will be considered. ae DEVELOPMENT OF THE PERISTOMIAL LAYERS As earlier writers have pointed out, the divisions which take place in the segments cut off from the apical cell of the young sporophyte proceed with great regularity, whether the segment is destined to form a part of the stalk or of one of the regions of the capsule. This regularity is characteristic not only of an indi- vidual species of the Bryales but of the group taken as a whole. ~ It has been recently emphasized by Kuntzen (712) in the particular | case of Ceratodon purpureus and becomes strikingly evident through the study of the operculum and peristome. The.oper- cular segments, like those which go to form the spore-case, are in _ tn Aus ae £ % st OC Se Oe OT ak SR eee ole ens Me a Evans AND Hooker: PERISTOME IN CERATODON PURPUREUS 101 the form of half-cylinders, which become divided into quadrants by anticlinal walls. Cross sections through young sporophytes just below the apical cell show this condition clearly (Fic. 1, 2). It is in the quadrants that the division takes place according to the “fundamental square’”’ method. If transverse walls are left out of consideration each quadrant divides by an anticlinal wall, extending as a curved surface from one of the side walls to the outside wall, into two cells (FIG. 3), a smaller triangular cell (as seen in section) and a larger four-sided cell. The latter soon divides by a periclinal wall into an outer and an inner cell (FIG. 4). According to Kuntzen (12, p. 19) the segments in the stalk sometimes divide in the way just described and sometimes by means of a periclinal wall followed by an anticlinal wall in the outer of the two cells thus formed. In the spore-case he finds the second method only but admits that perhaps the first method is sometimes followed. In either case, after the rearrangement of the walls, the cross section shows four inner cells, the ‘‘funda- mental square,’’ surrounded by eight outer cells. The inner cells constitute the endothecium and the outer cells the am- phithecium. A similar arrangement of the cells can be demon- strated in the young sporophytes of most of the higher bryophytes. In the next stage of development the amphithecial cells divide by periclinal walls, thus giving rise to two layers of eight cells each (FIG. 5). The inner cells, shaded in the figure, will develop into the inner peristomial layer. The corresponding layer in the spore-case gives rise to the outer spore-sac. It is upon the periclinal walls separating the two amphithecial layers that the teeth of the future peristome will be deposited. Marked differ- ences occur in the endothecium and amphithecium with respect to the relative rate of cell division. In the region of the annulus the inner peristomial layer completes its divisions before the endothecium has passed beyond the four-celled stage. In the region of the peristomial teeth the division may be simultaneous or the endothecium may divide first. In the portion of the operculum above the peristome the inner amphithecial layer never passes beyond the eight-celled stage. In the region of the teeth, after the stage shown in FIG. 5, the cells of the outer amphithecial layer divide by anticlinal walls, 102. Evans AND HooKER: PERISTOME IN CERATODON PURPUREUS %' Sse <> Fic. 1-10. Cross sections through the opercular region of young capsules, showing successive stages of development; peristomial layers stippled, X300. 1 and | 2, division of segments into quadrants; 3 and 4, establishment of amphithecium and endothecium; 5, establishment of inner peristomial layer; 6, division of cells in outer amphithecial layer by anticlinal walls; 7, establishment of outer peristomial 8 and 9, division of cells in inner peristomial layer by anticlinal walls; 10, continuation of divisions in outer amphithecial layer and in endothecium. thus raising the number of cells in this layer to sixteen (FIG. 6). Each of the sixteen cells then divides by a periclinal wall so that two layers of sixteen cells each are formed (FIG. 7). The inner layer represents the outer peristomial layer and undergoes no further divisions. It will be noted that the eight pairs of cells in this layer lie opposite the eight cells of the inner peristomial layer. In the spore-case the layer homologous with the outer peristomial layer gives rise to most of the photosynthetic tissue 4 EVANS AND HOOKER: PERISTOME IN CERATODON PURPUREUS 103 including the large intercellular space. The condition shown in FIG. 7 corresponds closely with figures by Kienitz-Gerloff (’78, pl. 1, f. 256) and Kuntzen (’12, f. 10), representing sections through the spore-case. The eight cells of the inner peristomial layer now proceed to divide in a very peculiar way. In each of these cells an anticlinal wall appears (FIG. 8), cutting off either to the right or left a small cell from one fourth to one third as wide as the parent cell. This is followed by a second anticlinal wall cutting off a similar small cell on the other side of the parent cell (FIG. 9). In this way the inner layer becomes divided into - twenty-four cells, each group of three corresponding with one of the pairs of cells in the outer layer. No further divisions are undergone by these twenty-four cells. The arrangement of cells just described seems to be even more constant than Kienitz- Gerloff implies, since he states that the number of cells in the inner layer may vary from twenty to twenty-four. The endothecium and the peripheral layer of the amphithecium play no direct part in the development of the peristome, and yet an account of the cell divisions that take place in them will not be wholly out of place. The divisions in the peripheral layer are not altogether definite. After the sixteen-celled stage, still shown in FIG. 8, the cells divide by anticlinal and periclinal walls without following a rigid system, adjacent cells often dividing in different ways (FIG. 10). In all cases, however, the final condition is° essentially the same, three concentric layers of cells being pro- duced. The outermost layer is composed of (approximately) 128 cells, the middle layer of 64 cells, and the innermost of 32 cells. In the region of the annulus, however, only two layers of cells are formed, thus allowing for the extraordinary development of the annular cells (FIG. 13, 25, 26). The peripheral layers of cells form the operculum, the external walls of the outermost layer becoming strongly thickened. The divisions in the endothecium of the opercular region are essentially like those in the spore-case, as described and figured by Kuntzen (’12). After the four-celled stage, shown in FIG. 8, division takes place according to the “fundamental square” method, eight outer cells and four inner cells being thus formed (FIG. 9). The inner cells repeat this method of division (FIG. 10), 104 Evans AND HooKER: PERISTOME IN CERATODON PURPUREUS while the cells of the outer layer divide by anticlinal walls in a more or less indefinite manner. When the capsule approaches maturity the endothecial tissue within the peristome consists of a relatively small number of large and thin-walled cells, this tissue of course being the continuation of the columella and the sporog- enous tissue in the spore-case. While the segments have been undergoing the anticlinal and periclinal divisions just described, divisions by transverse walls have also occurred in the various layers. Apparently they occur in a rather indefinite way and vary in different parts of the capsule (see Kuntzen, ’12, p. 22, etc.). In the peristomial layers trans- verse walls are especially numerous toward the base and tend, in this region at least, to be more numerous in the outer layer than in the inner (FIG. 13). Toward the apex they are more numerous in the inner layer (FIG. 12). DEPOSITION OF THE PERISTOMIAL THICKENINGS The forty rows of cells taking part in the formation of the peristome may be naturally divided into eight groups of five rows each. In each group three rows of the inner peristomial layer correspond with two rows of the outer layer. A cross section of such a group before the deposition of thickening is shown in FIG. 11. A group gives rise to two teeth of the peristome and the Same processes are repeated in each group. The peristomial cells, before the thickenings are laid down, contain dense cytoplasm and small nuclei with several nucleoli but are usually without vacuoles (FIG. 11-13). In the upper part of the peristome, where the teeth are divided into branches, eight regions of thicken- ing can be distinguished in each group, the four of the outer layer corresponding with the four of the inner layer. Two regions belong to each cell of the outer layer and to the median cell of the inner layer, while one region belongs to each lateral cell of the inner layer. The thickening first makes its appearance in the inner layer (FIG. 14) but soon becomes evident in the outer layer as well (FIG. 15, 16). If two corresponding regions are regarded as forming a single strand, each group will then form four strands, _ representing the four branches of the two peristomial teeth. The deposition of thickening begins in the basal portion of EVANS AND Hooker: PERISTOME IN CERATODON PURPUREUS 105 Fic. 11-16. Cross and radial sections through the amphithecial tissues of older capsules, X300. 11 12, showing the peristomial layers in the region of the teeth, just before the deposition of the thickenings; 13, showing the same stage in the annular region; 14-16, showing early stages in the development of the thicken- ings the peristome but follows essentially the same method throughout the branches of the teeth. The portions of the strands deposited by the inner peristomial cells are broader than those deposited by the outer cells. They are therefore more conspicuous at first, but become less so later on. This is due to the fact that the inner portions of the strands develop to a slighter extent and in thickness only, while the outer portions give rise to the thick 106 Evans AND Hooker: PERISTOME IN CERATODON PURPUREUS subcylindrical ridges already noted in the mature teeth (FIG. 17). The differences between the inner and outer portions become less and less marked toward the apices of the branches (FIG. 18). Upon the transverse walls in the outer peristomial layer thickenings are also deposited. These are continuous with the thickenings on the inner wall and give rise to the transverse ridges of the mature teeth (FIG. 19). In a young stage, seen in tangential section (FIG. 20), the transverse thickenings apparently broaden out the outer longitudinal ridges, and the same appearance is even more ‘ex 9 A Ha ae Fic. 17-19.- Cross and radial a ase in the region of the teeth, showing the peristomial thickenings in their final stage of development, 300; 20, tangential view of young peristome, seen from the outside, X300; 21, basal portion of a fully developed tooth, seen from the outside, X300. striking in an older tooth (FIG. 21). Toward the base of the branches the transverse ridges grow wider and wider until finally some of them span the distance between the branches and coalesce, thus forming a continuous ridge across the entire tooth. Below this region the ridges are all continuous. If Philibert’s ideas regarding the homologies of the peristomial layers are accepted, it becomes evident that the transverse ridges just described corre- spond with those present on the inner surface of the outer peristome in the Diplolepideae. In Ceratodon purpureus, however, the EVANS AND HOOKER: PERISTOME IN CERATODON PURPUREUS 107 ridges become adherent to the inner walls of the peristomial layer instead of to the outer walls, as is the case in the Diplolepideae. In the lower part of a tooth the longitudinal ridges on the outside also show an increase in thickness until they extend more than half way across the cavities of the cells in which they are deposited (FIG. 22, on left). In the basal undivided portion of a tooth these ridges finally coalesce and form a single broad ridge (F1G. 22, Fic. 22~26. Cross and radial sections in the region of the basal portion of the peristome, X300. 22 and 23, showing the portions of two teeth; 24 and 25, showing the same, together with the basilar membrane; 26, showing the basilar membrane only. on right, 23), beyond which the transverse ridges project for only a short distance (FIG. 24, 25). The thinner deposits of thickening formed by the inner per- istomial cells gradually increase in width toward the base until they finally meet and coalesce at the radial walls between the two rows of cells (FIG. 17, 22, 23). The vestiges of the radial walls © form a zigzag longitudinal line on the inner surface of the tooth, and not on the outside as in the Diplolepideae. This line is of course much shorter than in Funaria hygrometrica and Mnium hornum, in which the teeth are not divided into branches. No 108 Evans AND Hooker: PERISTOME IN CERATODON PURPUREUS transverse ridges are developed in the inner peristomial layer, but the vestiges of the walls remain visible as fine lines. When the strands of thickening have reached their full size, the spicules of cell wall substance are deposited in immense number over the entire surface (FIG. 17-19, 21-25). Those formed by the outer cells are considerably longer than those formed by the inner cells, but all are exceedingly minute. The cells which form the basilar membrane of the peristome acquire thick walls also, but the deposits of thickening differ from those which form the teeth. This is especially true of the outer peristomial layer where the thickening extends almost evenly over the inner walls, the radial walls, and the transverse walls but leaves large pits on the outer walls (Fic. 25, 26). In the inner per- istomial layer the thickening is less pronounced and forms a con- tinuous layer over the outer wall. Throughout this portion of the peristome the surface of the thickening remains smooth, no spicules being developed. The basilar membrane lies just within the large cells of the annulus (FIG. 25). Soon after the spicules have been deposited upon the teeth the peristomial cells dry up, the thin parts of the walls shrivel away and disappear, and the teeth become free. In the basilar membrane the pits in the outer wall now appear as perforations. At the same time the operculum becomes separated, and the endothecial tissues disintegrate. The finer structure of the mature peristome and the hygroscopic movements which it executes are fully described by Steinbrinck (’97). SUMMARY The original amphithecium, showing eight cells in cross section, divides by periclinal walls into an inner and an outer layer. The inner peristomial layer develops from the inner amphi- thecial layer, undergoing division by anticlinal walls until it is composed of twenty-four longitudinal rows of cells. The outer peristomial layer develops from sixteen longitudinal rows of cells cut off by periclinal walls from the outer amphithecial layer, after its eight rows have been divided by anticlinal walls; the outer peristomial layer undergoes no further divisions by anticlinal walls. EvANS AND HOOKER: PERISTOME IN CERATODON PURPUREUS 109 Ridges of thickening, representing the future teeth, are laid down upon the periclinal walls between the two peristomial layers. The cells of the peristomial layers form eight groups, each composed of two rows of cells of the outer layer and three rows of the inner layer. Each group gives rise to two teeth. In the upper part of each group eight deposits of thickening are laid down in four strands, representing the four branches of the two teeth; in the lower part only two strands are formed, representing the basal undivided portions of the teeth. In the outer peristomial layer thickenings are deposited also upon the transverse walls, representing the transverse ridges of the teeth. In the undivided basal portion of each tooth a fine median longitudinal line on the inner surface represents the vestiges of the radial walls between two rows of peristomial cells. In the basilar membrane the thickening of the walls in the outer peristomial layer is uniform except in case of the outer walls. YALE UNIVERSITY. LITERATURE CITED Campbell, D. H. (’05). The structure and development of mosses and ferns. Second edition. New York. Cavers, F. (11). The inter-relationships of the Bryophyta. New Phytologist Reprint, No. 4. Goebel, K. (’87). Outlines of classification and special morphology of plants. English translation. Oxford. Kienitz-Gerloff, F. (’78). Untersuchungen iiber die Entwickelungs- geschichte der Laubmoos-Kapsel und die Embryo-Entwickelung einiger Polypodiaceen. Bot. Zeit. 36: 33-64. pl. I-3. Kuntzen, H. ('12). Entwickelungsgeschichte des Sporogons von Ceratodon purpureus. Inaug.-Diss. Berlin. Lantzius-Beninga, S. (’50). Beitrage zur Kenntniss des innern Baues der ausgewachsenen Mooskapsel, insbesondere des Peristomes. Nova Acta Kais. Leop.-Carol. Acad. 22: 560-604. pl. 56-66. Philibert, H. (’84). De l’importance du péristome pour les affinités naturelles des mousses. 2e article. Rev. Bryol. 11: 65-72. Philibert, H. (’88). Etudes sur le péristome. 7€ article. Rev. Bryol. "18: 6-12; 24-28; 37-44; 50-60; 65-69. Steinbrinck, C. (’97). Der hygroscopische Mechanismus des Laub- moosperistoms. Flora 84 Erganzungsbd.: 131-158. Strasburger, E. ('o2). Das botanische Practicum. 4te Auflage. Jena. A note on the significance of sugar in the tubers of Solanum tuberosum O. BUTLER (WITH PLATE 2) Sugar accumulates in the tubers of the potato during the rest period when they are subjected to low temperatures, and, at least this is the general opinion, at the time of germination. In other words sugar develops at a time when the various vitalistic phe- nomena are inactive and again when the life processes are pro- ceeding actively. I During the rest period two factors influence the accumulation of sugar in the tubers of the potato: temperature and oxygen supply. Both these factors may act independently of one another or additively. I shall, however, for convenience, consider them separately. TEMPERATURE The effect of temperature on the accumulation of sugar in potato tubers was studied years ago by Miiller-Thurgau* and, as the results he obtained have not been disproved, I will simply briefly review them, adding some data of my own in confirmation. The belief is very tenaciously rooted in the popular mind that a potato becomes sweet only when it is frozen. A frozen potato never becomes sweet, moreover a potato does not freeze at 0° C., whence probably the origin of this notion. Potatoes, in fact, may be stored at 0° C. for a period of time without being injured, Miiller-Thurgau having kept them at this temperature in some of his experiments for as long as one hundred days.. Taking 0° C., then, as the lowest temperature at which potatoes may be safely stored, Miiller-Thurgau found that when the rates of sugar accumulation at 0° C., 3° C., and 6° C. were compared * Miiller-Thurgau, H. Ueber Zuckeranhdufung in Pflanzentheilen in Folge niederer Temperatur. Landw. Jahrb. 11: 757-828. Vie, 110 BUTLER: SUGAR IN TUBERS OF SOLANUM TUBEROSUM 111 the following relation was approximately obtained, x being the amount of sugar formed at 0° pats” Cc, and at 6° C. He also observed that individual potatoes of the same variety showed marked variations in the rapidity with which sugar accumulated in their tissues, but that if at 0° C. sugar accumulated in relatively small amount in the tubers of a given variety, then no accumulation would occur at 6° C., and conversely, if sugar was formed rapidly at 0° C., then a temperature of 8° C. would be usually necessary to effect the same result. At 10° C. sugar never accumulated. The fact brought to light by Miiller-Thurgau that different varieties of potatoes exhibit differences in the rapidity of sugar accumulation is readily confirmed, as the following table shows. TABLE I SUGAR ACCUMULATION IN DIFFERENT VARIETIES OF POTATOES STORED AT 1°-5° C, FOR 28 DAYS Percentage reducing sugar in potatoes* At beginning of Variety of potato ” experimdnt After 28 days Daily increase ae ee way a 0.59 0.97 +0.013 Piers WOME Se a site 0.88 0.85 —0.001 Rural New Yorker.......... 0.17 I.01 +0.03 Sips aceceia wiwias tee 6 De 0.15 r25 +0.039 BOOTY Ose tecnico oe ee 0.42 0.86 +0.015 Red Alconates vr hs vs 0.10 I.42 +0.047 RBREY Ree eS San ies 0.43 1.10 +0.023 BROW ICEOE oo toe Soars 0.53 1.22 +0.024 PACE OIC 8p Cel nr ees 0.43 1.04 +0.021 Miiller-Thurgau found, taking his results for 3° C., as these more nearly approach the average temperature at which the potatoes were stored in the above experiment, that the daily increase during a period of 30 days was 0.021 per cent, results which are of the same order as those given in the above table. According to Miiller-Thurgau sugar does not accumulate in resting potatoes when they are stored at a temperature of 8° C. * All the quantitative determinations given in this paper were kindly made for me by Mr. F. B. Morrison, Madison, Wis. : 112 Butler: SUGAR IN TUBERS OF SOLANUM TUBEROSUM It would also appear as if potatoes stored at 8°-10° C. not only do not accumulate sugar but do not reconvert it into starch as occurs at higher temperatures (this.is Miiller-Thurgau’s view of the matter), or dispose of it through respiratory activity. The amount of sugar present in the potatoes when they are placed in storage at this temperature suffers little change. This fact is clearly shown in the following table: TABLE II CHANGES IN SUGAR CONTENT OF eines ee: VARIETIES OF POTATOES AFTER STORAGE —11° C. FOR 34 DA Percentage reducing sugar in potatoes Variety of potato pitino At end of 34 days Increase prea a Snare ee re me PCOS 0.59 0.43 —0.16 eivie anew sao 0.88 0.65 —0.23 iti peor Nosker Seong te 0.17 0.24 +0.07 Barly Obie 6.65 oo en ek 0.15 0.16 +0.01 Hatiy Hpee i556 7 oe ea eee 0.42 0.31 —O.1I Red Alcohol 2352 62, cee 0.10 0.17 +0.07 igloo ae ay aie elas 0.43 0.38 —0.05 BlGe Victor se 0.53 0.45 —0.08 MCEOCINIE F506. ke ess 0.43 0.39 —0.04 When potatoes containing sugar are stored at 20° C. the sugar gradually disappears due to increased respiration and recon- version to starch (Miiller-Thurgau). However, if the potatoes remain sufficiently long at 20° C. for germination to be initiated, an increase in the sugar content is liable to take place. The aberrant results in the following table may be accounted for on - these grounds. TABLE III Loss OF SUGAR IN POTATOES STORED AT 20° C. FOR 2I DAYS Percentage reducing sugar in potatoes Variety of potato . ee = At end of 21 days Decrease REUIGNEC oir oe Sees ' 0.59 0.22 ae American Wonder........... 0.88 0.05 —0.83 Rural New Yorker.......... 0.17 trace —0.17 Barty Olio ogee ic as beecs 0.15 0.08 —0.07 Karly Koseo oo o55 oO. 0.13 —0.29 Red Aloghel. .. e.sSie es cys 0.10 0.15 +0.05 Rusty Rose oo sue eos 0.43 0.37 —0.06 Blue Lesee§ Meee wea teak 0.53 0.40 —0.13 MOC OrINCK 56 os 5 oe eae 0.43 0.65 +o.22 BUTLER: SUGAR IN TUBERS OF SOLANUM TUBEROSUM 113 When potatoes containing sugar are placed at 20° C. respiration immediately increases to a very marked extent, while in the case of similar potatoes containing none or only traces of sugar the increase is very slight. The respiration of the potatoes con- taining sugar, while very rapid at first, reaches a maximum in one and a half or two days, and then gradually decreases until it becomes equal to that of potatoes containing little or no sugar. TEMPERATURE 20°C CARBON DIOXIDE RESPIRED IN PARTS PER MILLION U T T T T T T Se Bee Ie ee ee ee DURATION OF EXPERIMENT—IN DAYS Fic. 1. (After Miiller-Thurgau.) Respiration of potatoes containing different amounts of sugar at 20° C. Legend: : Respiration of 50 potatoes containing an average of 1.7814 per cent reducing sugar. : — — — — Respiration of 50 potatoes containing an average of 1.464 per cent reducing sugar ------ Respiration of 50 potatoes containing none or only traces of reducing sugar. The results obtained by Miiller-Thurgau are very striking and are well illustrated in the above graph taken from his work. (FIG. 1.) It is perfectly obvious that the respiratory activity of potatoes foie ez % 3297 wo @ei5+ We 0°G. xe 107 ag Oo ero te z n : Sg 0 : . Be . ¢ Ss fh. Be Oe See 2 ee ee ee ee s~ DURATION OF EXPERIMENT—IN DAYS Fic. 2. (After Miiller-Thurgau.) Influence of change of temperature on res- piratory activity of potatoes initially rich in reducing sugar. is more influenced by the presence of sugar than by temperature. Sugar is essentially a limiting factor. This is well illustrated in the second graph (FIG. 2), in which the respiratory activity of 114 Butter: SUGAR IN TUBERS OF SOLANUM TUBEROSUM potatoes stored at 0° C., then at 10° C., then at 20° C., and again at 10° C., is compared. It will be seen at once that when potatoes were placed at 10° C. respiration immediately increased, then fell gradually until the potatoes were placed at 20° C., when it rose again to the same maximum as before but decreased more rapidly and, after being placed at 10° C. again for a short while, became almost equal to what it had been when the potatoes were stored at 0° C. It should be noted that the experiments just recorded were begun after the potatoes had been kept for 28 days at 0° C., i. e., the potatoes used were rich in sugar. Miiller-Thurgau believed that when potatoes containing sugar were placed at 20° C. the sugar that disappeared was partly used up in respiration, and partly reconverted to starch, as the disappearance of the sugar could not be all accounted for by the amount of carbon dioxide exhaled. In performing his experiments Miiller-Thurgau sliced his potatoes longitudinally, using one half for immediate analysis and the other for analysis at the end of the experiments. Under these conditions, even when the proper corrections for loss of weight due to evaporation are made, some of the sugar will undoubtedly go to the formation of the suber produced along the cut surface, and not unlikely, the increase in the percentage of starch obtained was due to the hydration of the cellulose that had been recently or was being laid down. The question, however, is deserving of further study. OXYGEN We have seen that during the rest period sugar may be made to accumulate in potatoes by storing the tubers at temperatures varying between 0° C. and 6° C., but that at 8°-10° C. the amount of sugar (when any was present) remained stationary. The relation of the oxygen supply to the accumulation of sugar in potatoes therefore may be studied conveniently at 8°-10° C. In the experiment which I have performed on this subject I used the Sir Walter Raleigh variety of potato and the tubers were placed in tubulated bell jars of the usual pattern: Bell jar no. 1 contained 1,934 grams of potatoes, and the tubules were stopped with cotton, thus allowing a free diffusion of air. BUTLER: SUGAR IN TUBERS OF SOLANUM TUBEROSUM 115 Bell jar no. 2 contained 1,936 grams of potatoes and was connected with a water siphon by means of which approximately 8 per cent of the volume of the contained air was renewed daily. Bell jar no. 3 contained 1,897 grams of potatoes and was connected with a water siphon by means of which approximately 0.8 per cent of the volume of the contained air was renewed daily. Within a few days the air in all the bell jars became saturated with water vapor. The experiment was discontinued after 90 days, when the potatoes in the different bell jars were analyzed and found to contain siigar as follows: Bell jar no. I, 0.09 per cent sugar; bell jar no. 2, 0.22 per cent sugar; bell jar no. 3, 0.66 per cent sugar. The effect of reducing the oxygen supply on the accumulation of sugar in potatoes is, therefore, quite marked, though not nearly so notable as that of low temperature. II The view is very generally held that sugar accumulates in potatoes at the time of germination and that this accumulation is essential thereto. According to de Vries* sugar begins to appear in potatoes just prior to germination but before the buds show any signs of growth. Resting potatoes, he says, contain for the most part no sugar, its appearance indicating the initial stages of germination. The sugar first appears in the neighborhood of the eyes, more precisely in the parenchymatous tissue surrounding the bundles leading to them. At first present in small amounts, it soon increases in quantity, developing in all parts of the tubers. During their early period of growth the shoots contain starch but no sugar, though the tubers themselves are full of the latter, the larger amount being in the medulla, the smaller in the cortex. When the shoots are 8 mm. long sugar is found in them only in isolated spots, but somewhat later it is generally distributed, and this condition remains unchanged until they come through the “soil. The accumulation of sugar in potatoes has not, however, as regards germination, any particular physiological significance. As in the case of resting potatoes the amount of sugar found in * Vries, H. de. Keimungsgeschichte der Kartoffelknollen. Landw. Jahrb. 7: 236. 1878. : 116 BuTLER: SUGAR IN TUBERS OF SOLANUM TUBEROSUM germinating tubers is greatly influenced by the milieu in which germination takes place. Potatoes germinating at a relatively low temperature contain more sugar than potatoes germinating at relatively high temperatures, and potatoes germinating in soil contain more sugar than those germinating in a cellar. Again, the distribution of sugar in potatoes germinating in a cellar, for instance, is irregular. All these facts clearly indicate that the metabolism of germination is not concerned primarily in the distribution or degree of accumulation of sugar.* I will give a few examples in illustration. Conditions favorable for germination are not necessarily favorable for the accumulation of sugar. For instance, on the 26th of April, 1912, I examined a number of Early Ohio potatoes, and they contained as a rule no sugar (PLATE 2, FIG. 6) though the sprouts were replete with it. A sample of these potatoes was placed in the ice compartment of an ice chest for 20 days, after which the potatoes composing it contained sugar as indicated in FIG. 2,5. At the same time similar potatoes from the cellar con- tained sugar as indicated in FIG. 1 and 4. It is worthy of note that the potatoes stored in the ice chamber (Fic. 2, 5) resembled very closely the potatoes shown in FIG. 7, as regards distribution of the accumulated sugar. The distribution of sugar illustrated by FIG. 7 was rather common in stored potatoes during March, 1911, which stored potatoes had been at no time subjected to a temperature below 6° C. It is therefore clear that the low tem- perature had not induced irregularities in metabolism. A glance at the plate will also force upon one the conclusion that the degree of germination has no effect upon the distribution of sugar. Neither is there any connection between presence or absence of sugar in the cortex with corresponding changes in the medulla. The medulla may contain sugar even in large amounts and the cortex give no reaction for it (FIG. 3, 7); again the cortex may contain sugar and the medulla be free from it (FIG. 1), though such cases appear to be rarer than the former; again the cortex * The distribution of sugar in potatoes can be followed by boiling thin slices of the tubers in Fehling’s solution for a few minutes, when traces of sugar will be indi- cated by a yellow coloration and an abundance by a red coloration, various shades of orange indicating intermediate amounts. BUTLER: SUGAR IN TUBERS OF SOLANUM TUBEROSUM 117 may contain sugar in large amounts near the suber and be almost free from it near the line of the medulla, which in turn may be more or less replete with it. Conditions illustrated in FIG. 2, 5, 9 are also not uncommon. The distribution of sugar in germinating potatoes, considered with reference to the budding eyes, is also worthy of note. If one cuts longitudinal slices through potatoes in which the apical buds alone have sprouted he will find almost invariably that there is less sugar in the tubers immediately below the shoots than elsewhere. The conditions illustrated in FIG. 2, 5, 7, in which the medulla colors orange in Fehling’s solution except for small areas beneath the insertion of the shoots and around the buds, may be also observed. Qualitative tests having indicated that in germinating potatoes sugar occurred in less amounts near the apices than toward the bases and having also given indications that the bases were usually richer than the middle portions, I thought it would be well to have these results confirmed by quantitative analyses. The results obtained were as follows. TABLE IV DISTRIBUTION OF SUGAR IN GERMINATING POTATOES Percentage reducing sugar in potatoes Remarks Variety Apical zone Middle zone Basal zone Gas a sg 8) se eee nee ane 0.12 .28 59 Single tuber. Composite Hatly ONG 25 63 shies None 15 42 sample of ro tubers. BarhrOMn. 62304 saa ce Trace 35 88 Single tuber. Flashy OM 666 oa ccs ees Trace .46 50 Single tuber Eany Gale oro. os ners None 35 85 Sin le tuber SALTS ORG 255 6 ee oe 0.17 -59 7 Single tuber. TABLE Iv confirms entirely the conclusions deducible from qualitative tests. Potatoes contain little or no sugar near their apices, sometimes nearly as much in their middle as in their basal portions, and sometimes much more towards the base than in the middle. It may be observed that Miiller-Thurgau* found that resting potatoes were richer in sugar at the base than at the apex, and we have seen that this is generally true also of germinating potatoes. * Loc. cit. p. 763. 118 ButTLerR: SUGAR IN TUBERS OF SOLANUM TUBEROSUM The buds in potatoes tend to be more numerous around the apices, and one would expect, therefore, that during the rest period as well as at germination metabolic activity would be greater in the apical portions. One of the consequences of this increased activity in the apices of the resting potatoes would be a lesser accumulation of sugar and the difference in the percentage present, as one proceeded towards the base, would be the more noticeable, the less ready the translocation of the sugar from one part of the tubers to another: from the seats of lesser activity to the seat of greater activity. The data given in TABLE Iv as well as the qualitative results illustrated in PLATE 2 show rather definitely, it seems to me, that there is little if any translocation from remote to budding parts even in germinating potatoes. Were the trans- location of sugar a normal and necessary function in the metab- olism of germination, its distribution would be more regular and constant; one would find a definite relation existing between cause and effect. The distribution of sugar (when present) in germi- nating and resting potatoes is not essentially different,* and it would seem, therefore, that its appearance in quantity just before or at germination should be ascribed, at least in part, to metabolic changes induced by another agent, or other agencies. DurxHaM, NEw HAMPSHIRE. Description of plate 2 Fic. 1. Longitudinal section through Early Ohio from cellar, 16th May 1912. Fic. 2. Longitudinal section through Early Ohio stored in ice chest 20 days, 16th May 1912, Compare with Fic. 1 Fic. 3. Longitudinal section through Rural New Yorker from cellar, March Igit. x ea 4. Longitudinal section through Early Ohio from cellar, 16th May 1912. . 5. Longitudinal section oe Early Ohio stored in ice chest 20 days, 16th ioe Igr2. oe are with FIG Lon pie tudinal section of ae Ohio from cellar, 26th April 1912. Fic. 7. Lon Salta section of Early Ohio from cellar, 8th March rorr. Fic. 8. Leann section through Triumph from cellar, March rort. Fic. 9. Longitudinal section through Triumph from cellar, March 1o1I. “* It is well to point out that germinating potatoes as well as resting potatoes may contain no sugar (cf. FIG. 6). Studies in the Agalinanae,.a subtribe of the Rhinanthaceae * FRANCIS W, PENNELL I. NOMENCLATURE OF THE NEARCTIC GENERA As here defined the Agalinanae constitute a subtribe of the Buchnereae and include a group of closely allied genera, dis- tinguished from Buchnera and its nearer allies only by the normally developed two-celled anthers. These studies concern but a section of this group, all American save for one doubtful record from Madagascar, the genera listed by Von Wettstein in Die Natiirlichen Pflanzenfamilien as Esterhazya Mikan, Macranthera Torr., Seymeria Pursh, Silzia Benth., and Gerardia Linn. In this paper it is desired to place on a firm basis the nomen- clature of the genera occurring in North America north of the Mexican Boundary. This has not proved as easy as anticipated, owing to a misunderstanding of a number of older genera, chief among which is that of Gerardia itself. This paper divides itself naturally into two portions, a history of the genus Gerardia (Plumier) Linn., explaining the reason for its rejection as a genus of the Rhinanthaceae, and a history of the Rhinanthaceous genera proposed from time to time under which our species of this group must now be placed. This paper con- cludes with a summary of the nomenclature it is proposed to follow in these studies. In 1703 the French traveller and botanist, Charles Plumier, a member of the religious order of the Minimi, published a work, “Nova Plantarum Americanarum Genera,” containing descrip- tions of new genera observed during three voyages to America from 1689 to 1697. One of these is the new genus Gerardia. In this work, a volume of 52 pages of Latin text and 40 plates, I find little mention of the portions of America visited, but in the preface to the same author’s “‘ Description des Plantes de L’Amérique”’ more information is given. He tells us that he went *Contribution from the Botanical Laboratory of the University of Pennsyl- vania. 119 120 PENNELL: STUDIES IN THE AGALINANAE first as aid to M. Surian, charged with a commission from the King of France to the Isles Antilles, “‘to make search of all that Nature there produces most rare and most curious.” During the nearly ten years spent in the West Indies, he described, he says, nearly six hundred different plants. Gerardia is both described and figured. The description reads:—‘‘Gerardia est plantae genus flore A monopetalo, perso- nato, cujus labium superius surrectum est, subrotundum & -emarginatum, inferius vero in tres partes divisum, media bifida. Ex calyce autem C surgit pistillum posticae floris parti B, ad instar clavi infixum, quod deinde abit in fructum D oblongum, gibbum, septo medio E, in duo loculamenta divisum, seminibusque foetum orbicularibus F.’’ Then follows the note,—‘‘Gerardiae unicam speciem vidi. Gerardia humilis, Bugulae foliis, Asphodeli radice.”’ The genus so founded by Plumier remained unaltered, and very little known, till the first edition of the Species Plantarum in 1753. Here Linnaeus took it up, and to Plumier’s species which he named Gerardia tuberosa, added four others, purpurea, flava, pedicularia and glutinosa. It is evident, being the species adopted by Linnaeus from the original author of the genus, incidentally also being his first species listed, Gerardia tuberosa L. must be considered the type of the genus Gerardia (Plumier) Linnaeus.* In addition to the striking feature indicated by the specific name, tuberosa is characterized by Linnaeus as “‘Gerardia foltis subovatis tomentosis repandis, longitudine caulis,’ points quite at variance from those of his other species with which the name Gerardia has come later to be exclusively associated. Yet Lin- naeus’ list of species, including tuberosa, with a few additions from time to time, was copied successively from author to author—by Buc’hoz (1778), Lamarck (1786), J. F. Gmelin (1791), etc., to Willdenow (1800), and Persoon (1807). A few authors of this period realized the incongruity of such treatment, and that logically the name Gerardia should apply to tuberosa alone. * Though worked out independently by the writer, this same conclusion was a few months earlier by Dr. N. L. Britton. I am indebted to Dr. J. H. Barnhart for reviewing and confirming my determination of the type of Gerardia. PENNELL: STUDIES IN, THE AGALINANAE 121 First among these was evidently Walter. In his Flora Caro- liniana (1788) so far as our North American species are concerned, he relegates Gerardia to synonymy, and, not wishing to coin a new name, places purpurea, flava and pedicularia in one of his numerous genera called Anonymos. In 1810 in treating Gerardia in Rees’ Cyclopedia Sir James E. Smith, after listing first Linnaeus’ tuberosa, remarks upon its identity being yet doubtful and suggests as a desideratum an examination of its fruit. Then he makes this statement. ‘‘What- ever might be the result of such examination this plant must be the true though it were the only Gerardia, and the rest in that case must have a new generic appellation and character.” This is the first definite assignment of a type species for the genus. As to the further history of Gerardia tuberosa L., after Sir J. E. Smith I find no writer retaining this plant in the Rhinanthaceae. In 1825 Sprengel interpreted it as a synonym of Ruellia rupestris Swartz, an Acanthaceous plant, whence in 1847 it was carried into the new genus Stenandrium of Nees, becoming a synonym of Stenandrium rupestre (Swartz) Nees. If this identification be correct Stenandrium Nees should become Gerardia (Plumier) L. Though antedating the erection of Stenandriwm into a genus, such a change was actually made by Rafinesque in his Flora Telluriana in 1838, where Gerardia tuberosa L., G. rupestris (Swartz) Raf., and G. scabrosa (Swartz) Raf. are cited. In 1835 Bentham in his ‘‘Synopsis of the Gerardieae”’ discusses the past history quite fully, definitely relegates G. tuberosa L. to the Acanthaceae, and endorses the earlier selection, practically made by Sprengel, of G. purpurea L. as the type. This view has been mostly followed till the present day. If Gerardia is properly an Acanthaceous genus what name is to be applied to our familiar North American species commonly so called? Of the Linnaean species of this genus to be retained in the Rhinanthaceae three were North American and one Chinese, the latter however not proving a near ally of the others. Species continued to be added from both hemispheres till as late as 1846 when in DeCandolle’s Prodromus Bentham finally separated the 122 PENNELL: STUDIES IN THE AGALINANAE series into a number of definitely restricted New World and Old World genera. The Old World genera differ from the New in more points than were realized at that time, so may be definitely dismissed from the present discussion. After Linnaeus’ time the first new names proposed were in 1788 Walter’s two genera both named Anonymos. These were well characterized; the one might be typified by Gerardia purpurea L., the other by a new species Anonymos cassioides Walt. Of course the name—or confession of the lack of a name—Anonymos, has no value in nomenclature. In 1791 J. F. Gmelin, reviewing Walter’s work, returned his first Anonymos to Gerardia, but maintained his second Anonymos as a new genus Afzelia. Anonymos cassioides Walt. became Afzelia cassioides (Walt.) J. F. Gmel. This genus also was by later authors returned to Gerardia, and when in 1814 Pursh became convinced of its generic distinctness, finding the name Afzelia meantime applied to a genus of the Caesalpiniaceae, he renamed the genus Seymeria. Afzelia J. F. Gmel. was restored by Kuntze in 1891. The genus to which this name is applied is a definite, natural group of Mexico and the southern coastal plain region of the United States. The next generic description in this group is in 1794 that of Virgularia described by Ruiz and Pavon from Peru, and based upon their V. lanceolata, the specific description of which did not appear till 1798. Though loath with limited material to enter upon any discussion of the South American species of this group, it is necessary to attempt to decide whether this genus can be distinguished from those later proposed to include our North American species. In the differential characters pointed out by Ruiz and Pavon I find, I confess, little that is convincing. The description of a bifid stigma is surely remarkable, but as no later observer has recorded such a structure in this or in any other South American species of this group, there is doubtless here some error. Also the characters depended upon by Martius (1829) have been shown by Bentham (1835) to be untrustworthy. Since 1835 the name Virgularia has been considered a synonym of Gerardia L. Yet an inspection of the species of Virgularia will, I think, show us PENNELL: STUDIES IN THE AGALINANAE 123 sufficient points of contrast. The material before me is all Bolivian,* but may be safely assigned to this genus. . In Virgularia the plant is shrubby, and for our purpose is best distinguished by its tubular, fleshy corolla, mostly red (or some allied shade), after flowering somewhat persistent, shriveling and only tardily falling. These characters appear again, at least in greater part, in the Brazilian Esterhazya Mikan, and in our North American Macranthera Torr. (to be mentioned later), and seem to indicate a sharp distinction between these three genera and the remainder. Virgularia is to be held as a natural, well- marked genus of western South America. The remainder of this paper will deal strictly with the species of the United States, no other generic name having been proposed from Tropical or South America which can affect the nomen- clature of our species. Yet it must be remembered that a or the great center of this group lies in South America, and no complete _ understanding of the inter-relationship of the whole can be gained till the species there are studied. Such study is deferred. The next name that could concern us is in a work descriptive of fruits, Chytra Gaertn. fil., 1805. As only the fruit is shown, and that does not seem conclusive—and as no native country whatever is given—this plant may be left permanently as un- identifiable. Such a solution is suggested by Gaertner’s specific name anomala. Yet it must be recognized that his figure shows a decided resemblance to a ‘‘Gerardia” capsule. ; In 1819 Rafinesque descrided a yellow-flowered, coarse, lanceolate-leaved plant from Western Kentucky, under a new genus, Dasistoma. His description is clear and good, and leaves no doubt that his plant was the one described one year previously (1818) as Seymeria macrophylla Nutt. Rafinesque speaks of a short corolla-tube, rotate, 5-lobed limb, 4 nearly equa stamens, short filaments, glabrous anthers, short, cylindrical style, thick, obtuse stigma,—excellent diagnostic characters for macrophylla, but all impossible for Dasistoma (spelled Dasystoma) as taken up in 1846 by Bentham, and followed in later works. Dasistoma was based upon D. aurea Raf. (1819), antedated by Seymeria macrophylla Nutt. (1818), so the combination becomes * Bang 188, 730, 2530, 2854; Buchtien 129, 789; Rusby 1077, 1078, 1 080, 1081. 124 PENNELL: STUDIES IN THE AGALINANAE D. macrophylla (Nutt.) Raf. (1837). Seymeria macrophylla Nutt. was in 1846 made the basis of a section Brachygyne Benth., which in 1903 was raised to a genus Brachygyne (Benth.) Small, identical with the older Dasistoma Raf. The genus is monotypic. In 1834 Nuttall described a new genus Conradia based upon a large and showy plant, specimens of which he had seen in the herbarium of the Philadelphia Academy of Natural Sciences. His name Conradia was antedated by Conradia Raf. (1825), and by Conradia Mart. (1829), so in 1835, in the account of the plants of Drummond’s collection, the name was changed to Macranthera. Though this series of articles in the Companion to the Botanical Magazine was mostly the work of the editor, Sir W. J. Hooker, Bentham is to be credited for this genus. In the original descrip- tion Le Conte’s and Bentham’s names were both cited after the genus, but in a later article during the same year Bentham tells us Macranthera was a manuscript name used by Dr. Torrey in communicating the plant to Dr. Lindley. Doubtless because the collector of the plant in Torrey’s herbarium (however described two years later as a second species) Le Conte was mentioned, so with Bentham’s explanation Torrey’s name may be con- nected with the plant in question. Conradia Nutt. was based upon C. fuschioides Nutt., the spelling of which was corrected to fuchsioides in the combination Macranthera fuchsioides (Nutt.) Benth. However, as this plant had been previously described by William Bartram in his Travels (1791) as Gerardia flammea, this species must become Macranthera flammea (Bartram) Pennell comb. nov. In 1835 Bentham reviewed this tribe, adding no new generic names, but systematizing and coérdinating the whole, and with another revision in 1846 giving the outline which has been mostly adopted since. f In 1837 Rafinesque undertook the special elaboration of this group in his New Flora of America, adding several new generic names, not giving us a very satisfactory or codrdinated treatment, yet showing nevertheless a surprising insight into the group. As I have already had to refer to several of his names, as he proposed a number of genera which must be adopted, and as since his time no new genera have been proposed which he had not already defined, it will be well to go carefully over his treatment. PENNELL: STUDIES IN THE AGALINANAE 125 In the first place, as already recounted, he definitely carried out Sir J. E. Smith’s suggestion, and transferred Gerardia as a valid genus to the Acanthaceae. . Our Rhinanthaceous species, Gerardia of authors and its near allies, he placed in about eight genera, two of which, Macran- thera ‘“‘Torr.”” Benth. and Seymeria Pursh, were adopted from other authors, and one, Dasistoma, was, as shown above, an earlier genus of his own. His first genus was Aureolaria, created to include the large, perennial, broad-leaved species, and based upon the current interpretation of Gerardia flava L. As his Dasistoma has been wrongly applied to this group, this first name becomes the correct one for these plants. As Gerardia flava L., both by description and the specimen in the Linnaean herbarium, is synonymous with Rhinanthus virginicus L., the type-species of the present genus—our pubescent eastern plant—must be known by the name here given it, Aureolaria villosa Raf. His second genus, doubtfully considered distinct from the last, is nearly as uncertain to the present reviewer. Panctenis was based upon Gerardia pedicularia L. This species and a few close allies agree with Awreolaria in broad leaves, yellow flowers, and awned anthers, but their points of disagreement are equally striking. The annual habit of Panctenis, its corolla pubescent without, its wingless seeds, constitute several points of difference. I prefer to treat it as a subgenus of Aureolaria. The combination Aureolaria pedicularia (L.) was made as a variant by Rafinesque. His third genus, Agalinis, in point of numbers is the most important of all. Based upon Agalinis palustris Raf., this genus was designed to include all the slender, narrow-leaved, purple- flowered plants, which Bentham later (1846) has treated as his section Eugerardia, and recent writers have come to consider true Gerardia. This isa large genus of both North and South America, falling into a number of well-marked subgenera. The type of the genus, Agalinis palustris Raf., is the prevalent plant of moist ground, near marshes, from New England to Carolina, Rafinesque correctly interpreting Gerardia purpurea L. as intended to include all the purple species. However G. purpurea L. is to be typified by the only Linnaean citation with a figure, Plukenet’s ‘‘ Digitalis 126 PENNELL: STUDIES IN THE AGALINANAE virginiana rubra, foliis et facie antirrhini vulgaris,’’ which is unquestionably the current interpretation of the species. A galinis palustris Raf. becomes Agalinis purpurea (L.) Pennell comb. nov. In Tomanthera he placed, as T. lanceolata, a plant in his her-. barium collected in New Jersey by Dr. Cleaver, and cited the species as occurring in Pennsylvania and Carolina. His plant appears to have been undoubtedly Gerardia auriculata Michx., described in 1803 from the prairies of Illinois, his specimen being quite small for the species. Possibly the leaves were abnormally entire, or perhaps they were lobed at base and this feature over- looked. His notice of the rarity of the plant is interesting, agreeing with Dr. Darlington and more recent observers of its sporadic occurrence in the East. With T. lanceolata he correctly but doubtfully associated Gerardia auriculata Michx., though incorrectly as a distinct species. The genus Tomanthera, including two species as now understood, is accordingly based upon T. auriculata (Michx.) Raf. In 1835 Bentham had based his section Otophylla upon Gerardia auriculata Michx., in 1846 raising this to a genus of the same name. Otophylla Benth. (1846) therefore becomes a synonym of Tomanthera Raf. (1837). His Dasistoma of 1819 was here continued, and, as above shown, Seymeria macrophylla Nutt. was identified with it, though as a distinct species. The name Dasistoma was here spelled Dasistema, and D. aurea of 1819 was changed to D. auriculata. Seymeria he adopted unaltered from Pursh. A genus Ovostima was described based upon one species O. petiolata from Florida or Alabama. From the description of the plant, the large smooth corolla, bicuspidate anthers, etc., I believe the plant to have been an Aureolaria, though the descrip- tion of the flower as white is surprising. In Awreolaria the corolla is fleshy and blackens in drying, not thin and apparently white or very pale ochroleucous as described for Ovostima. The plant is left as of doubtful identity. Macranthera was adopted from Bentham, and for the two species that had then been published, M. fuchsioides (N utt.) Benth. and M. Lecontei Torr., he proposed two additional generic names, Toxopus and Tomilix. As M. Lecontei Torr. appears not to have been published till 1837 there is sufficient evidence that PENNELL: STUDIES IN THE AGALINANAE 127 the date on the title page, 1836, is not the date of this volume of Rafinesque’s work. Macranthera fuchsioides (Nutt.) was yet again described by Rafinesque as Russelia flammea based upon Bartram’s incidental description of the plant in his Travels as Gerardia flammea, and the suggestion made that it is possibly to be considered as a new genus Flamaria. An earlier genus of Rafinesque’s, Pagesia (1817), reproduced in the New Flora, has been attributed by various authors to Dasystoma or Gerardia, but certainly does not belong here. Can Pagesia leucantha Raf. be identified with Mecardonia acuminata (Walt.) Small? Finally the genus Dasanthera Raf., under which in the New Flora he placed Gerardia cunetfolia Pursh and G. fruticosa Pursh, has no close affinity with this group, nor even harmony within itself. One species, G. cunetfolia, had been already identified by Bentham (1835) as Gratiola acuminata Walt. (cited as Ell.), so by synonomy would be Mecardonia acuminata (Walt.) Small; the other, G. fruticosa, by the same author in 1835 was considered a Pentstemon, in 1846 as his P. Lewisii, a name which Greene (1892) has changed to P. fruticosus (Pursh). Since 1836 there have been but two new generic names proposed for Nearctic species of this group, Otophylla Benth. (1846), and Brachygyne (Benth.) Small (1903), both of which are antedated by names of Rafinesque’s. The present writer has none to add to the sufficient number already published. ; A summary of the Nearctic genera of this group, with the type species for each, would be: AFzELIA J. F. Gmel.; Linn. Syst. Nat. ed. 13.927. 1791. —Anonymos casstoides Walt. —Seymeria Pursh, Fl. Amer. Sept. 736. 1814. —Anonymos cassioides Walt. DasisToMa Raf. Journ. de Phys. 89:99. 1819. ——Dasistoma aurea Raf. (= Seymerta mac- rophylla Nutt.). —Brachygyne (Benth.) Small, Fl. S. E. U. S. 1073. 1903. —Seymeria macrophylla Nutt. MacranTHERA “Torr.’’; Benth. in Hook. Comp. Bot. Mag. I: 174. 1835. 128 PENNELL: STUDIES IN THE AGALINANAE —Conradia fuchsioides Nutt. (= Gerardia flammea Bartram). —Conradia Nutt. Jour. Acad. Nat. Sci. Phila. 7: 88. 1834 [not Conradia Raf. Neog. 3. 1825; nor Conradia Mart. Nov. Gen. et Sp. 3:38. 1829]. —Conradia fuchsioides Nutt. —Flamaria Raf. New Flor. Amer. 2: 71. 1837. —Gerardia flammea Bartram. —Toxopus Raf. New Flor. Amer. 2:71. 1837 —Macranthera Lecontet Torr. —Tomilix Raf. New Flor. Amer. 2: 72. 37. —Conradia fuchsioides Nutt. AUREOLARIA Raf. New Flor. Amer. 2:58. 1837. —Aureolaria villosa Raf. —FPanctenis Raf. New Flor. Amer. 2: 60. 1837. —Gerardia pedicularia L. (?)—Ovostima Raf. New Flor. Amer. 2:70. 1837. —Ovostima petiolata Raf. AGALINIS Raf. New Flor. Amer. 2: 61. 1837. —Agalinis palustris Raf. (= Gerardia pur- purea L.). TOMANTHERA Raf. New Flor. Amer. 2:65. 1837. —Tomanthera lanceolata Raf. (= Gerardia auriculata Michx.). —Otophylla Benth. in DC. Prodr. 10: 512. 1846. —Gerardia auriculata Michx. UNIVERSITY OF PENNSYLVANIA. LITERATURE CITED | Bartram, William. Travels through North and South Carolina, Georgia, east and west Florida, etc., 412. Philadelphia, 1791. Bentham, George. Synopsis of the Gerardieae, a tribe of the Scrophu- lariaceae. Hook. Comp. Bot. Mag. 1: 198-212. . Bentham, George. Scrophulariaceae, in DeCandolle, Prodromus 0: SII—S§21.-. 1847. - Buc’hoz, P. J. Gerardia. in Histoire Universelle du Regne Végétale 9: 97-99. 1778. Darlington, William. Flora Cestrica, 365. West Chester, Pa. 1837- Gaertner, C. F. De Fructibus et Seminibus Plantarum 3: 184 — pl. 214, fig. 4. Stuttgart, 1805. PENNELL: STUDIES IN THE AGALINANAE 129 Gmelin, J. F. Linné, Systema Naturae, ed. 13. 2: 927-928. Leipzig, 1791. Greene, E. L. Pittonia 2: 239. Berkeley, Calif. 1892. Hooker, W. J. Notice concerning the late Mr. Drummond’s col- lections, made chiefly in the southern and western parts of the United States. Hook. Comp. Bot. Mag. 1:174. 1835. Kuntze, Otto. Revisio Generum Plantarum 1: 457. 1891. Lamarck, J. B. A. P. M. Encyclopédie méthodique, Botanique 2: 688. Paris, 1786. Linnaeus, Carolus. Species Plantarum 610. Stockholm, 1753. Martius, C. F. P. de. Nova Genera et Species Plantarum, quas in itinere per Brasiliam annis 1817-1820 jussu et auspiciis Maximiliani Josephi I Bavariae regis augustissimi suscepto 3: 5-11. Munich, 182 Michaux, André. Flora Boreali-Americana 2: 20. Paris, 1803. Nees von Esenbeck, C. G. Acanthaceae, in DeCandolle, Prodromus II: 283. 1847. Nuttall, Thomas. The genera of North American plants, and a catalogue of the species, to the year 1817 2: 49. Philadelphia, 1818. Nuttall, Thomas. A description of some of the rarer little known plants indigenous to the United States, from the dried specimens in the herbarium of the Academy of Natural Sciences in Philadelphia. Jour. Acad. Nat. Sci. Phila. 7: 88. 1834. Persoon, C. H. Synopsis Plantarum 2: 154. Paris, 1807. Plukenet, Leonard. Almagesti Botanici Mantissa 65. pl. 388, fig. 1. 1700. Plumier, Charles. Nova Plantarum Americanarum Genera 31. pl. I2. Paris, 1703 Pursh, Frederick. Flora Americae Septentrionalis 736. London, 1814. Rafinesque-Schmaltz, C. S. Florula Ludoviciana 48. New York, 1817. Rafinesque-Schmaltz, C.S. Prodrome des noveaux genres de plantes observés en 1817 et 1818 dans l’intérieur des Etats-Unis d’ Amérique. Jour. de Phys. Chim. et Hist. Nat. 89:99. 1819. Rafinesque-Schmaltz, C.S. New flora and botany of North America 2: 58-72. Philadelphia, 1837.* Rafinesque-Schmaltz, C. S. Flora Telluriana 4: 67. Philadelphia, 1838.* * For dates of publication of separate parts of these see Barnhart, Torreya 7: 177-182. 1907. 130 PENNELL: STUDIES IN THE AGALINANAE Ruiz, L. H., & Pavon, J. Florae Peruvianae et Chilensis Prodromus g2. pl. 19. Madrid, 1794. Ruiz, L. H., & Pavon, J. Systema vegetabilium Florae Peruvianae et Chilensis 161. Barcelona, 1798. Small, J. K. Flora of the southeastern United States 1073, 1338. New York, 1903. Smith, J. E. Gerardia, in Rees’ Cyclopedia. 1810.* Sprengel, Kurt. Linnaeus, Systema Vegetabilium, ed. 16. 2: 806-808, 825. Gottingen, 1825. Torrey, John. An account of several new genera and species of North American plants. Ann. Lyc. Nat. Hist. New York 4: 80. pl. 4. 1837.T Walter, Thomas. Flora Caroliniana 169-170. London, 1788. Wettstein, R. v. Scrophulariaceae, in Engler & Prantl, Die Natiir- lichen Pflanzenfamilien 4: 92-93. Leipzig, 1891. Willdenow, C. L. Linnaeus, Species Plantarum, ed. 4. 3: 221. Berlin, 1800. * I am indebted to Dr. J. H. Barnhart for the date of this volume. Tt No date with paper, cited as 1837 in Index Kewensis. INDEX TO AMERICAN BOTANICAL LITERATURE (1911-1913) aim of this Index is to include all current botanical literature written by Americans, apamy in America, or based upon American material ; the word Amer- ica being used in the broadest sense. Reviews, and papers that relate exclusively to forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and made in favor of some paper appearing in an American periodical which is devoted wholly to botany. Reprints are not mentioned unless they differ from the original in some important particular. If u of the Index will call the attention of the editor +o errors or omissions, their ‘aids will be appreciated. This Index is reprinted monthly on cards, and furnished in this form to subscribers at the rate of one cent for each card. Selections of cards are not permitted ; each subscriber must take all cards published during the term of his subscription, Corre- spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club, Arthur, J. C. A new weed exterminator. Science II. 37: 19. 3 Ja 1913. Banker, H. J. Type studies in the Hydnaceae—III. The genus Sarcodon. Mycologia 5: 12-17. Includes Sarcodon radicatus, S. Murrillii, S. fumosus, and S. roseolus, spp. nov. Ball, C. R. The kaoliangs: a new group of grain sorghums. Bs Dept. Agr. Plant Ind. Bull. 253: 5-64. pl. 1,2 +f. I-15. U Ja 1913. Bartlett, H. H. The purpling chromogen of a Hawaiian Dioscorea. U.S. Dept. Agr. Plant Ind. Bull. 264: 5-19. pl. 1 +f.1- 8 Ja 1913. Bates, J. M. Ophioglossum vulgatum in Nebraska. Fern Bull. 20: 67. [Ja 1913] Berger, A. Opuntia Ficus barbarica Berger nom. nov. teenk. 22: 181. 15 D 1912. Bessey, C. E. Some of the next steps in botanical science. 37: 1-13. 3 Ja 1913. Bitter, G. Solana nova vel minus cognita. V. Repert. Sp. Nov. 11: 349-394. 15 D1912; VI. Repert. Sp. Nov. 11: 431-473- 31 D 1912. Includes 17 new species and Bitter, and 6 new species in the section Polybo Brain, Monats. Kak- Science II. several new varieties in the section Tuberarium of tryon. _K. A list of fungi of Cedar Point. Ohio Nat. 13: 25-36: D 1912. : 131 182 INDEX TO AMERICAN BOTANICAL LITERATURE Brannon, M. A. Factors influencing the flora of Devils Lake, North Dakota. Internat. Rev. 4: 291-299. I9QII. Britton, E. G. Ditrichum rhynchostegium Kindb. Bryologist 16: 8. Ja 1913. Campbell, D.H. Some impressions of the flora of Guiana and Trinidad. Pop. Sci. Mo. 82: 19-32. f. 1-3. Ja 1913. Chambers, C. O. The relation of algae to dissolved oxygen and carbon-dioxide, with special reference to carbohydrates. Ann. Rep. Missouri Bot. Gard. 23: 171-207. 1912. Chodat, R. A grain of wheat. Pop. Sci. Mo. 82: 33-45. Ja 1913. Clute, W. N. Polypodium or Xiphiopteris? Fern Bull. 20: 65-67. [Ja 1913.] [IIllust.] Clute, W. N. Priority and fern names. Fern Bull. 20: 68-73. [Ja 1913.] [Clute, W. N.] Pteridographia. Fern Bull. 20: 74-80. [Ja 1913.] Includes notes on: Ferns in bottles; New stations for Florida ferns; Additional fern pests; Number of articles on ferns; A new Equisetum; New southwestern ferns; Death of James Goldie; Two new fern sports; Crested Christmas fern; Bermuda Clute, W. N. Rare forms of fernworts—XXIII. Who can name this fern? Fern Bull. 20: 73, 74. [Ja 1913.] [Illust.] Collins, G. N. Heredity of a maize variation. U. S. Dept. Agr. Plant Ind. Bull. 272: 5-23. pl. 1 +f.12. 21 Ja 1913. Conard, H. S. Revegetation of a denuded area. Bot. Gaz. 55: 80-84. f. 1, 2, 15 Ja 1913. Cook, O. F. Heredity and cotton breeding. U.S. Dept. Agr. Plant Ind. Bull. 256: 5-113. pl. 7-6 +f. 1-19. 13 Ja 1913. Cook, O. F. Morphology of cotton branches. U.S. Dept. Agr. Plant Ind. Circ. 109: 11-16. 4 Ja 1913. Cooper, W. S. The climax forest of Isle Royale, Lake Superior, and its development. I. Bot. Gaz. 55: 1-44. f. 1-14 + map. 15 Ja 1913. Cooper, W. S. A list of mosses collected upon Isle Royale, Lake Superior. Bryologist 16: 3-8. Ja 1913. Dachnowski, A. Peat deposits of Ohio: their origin, formation, and uses. Ohio Geol. Surv. Bull. IV. 16: 1-424. pl. 1-8 +f. I-20. Ap 1912. Darbishire, O. V. The lichens of the Swedish Antarctic Expedition. Wiss. Ergeb. Schwed. Siidpolar-Exp. 4%: 1-73. pl. 1-3 +f. 1: Stockholm. 1912. Davis, B. M. Mutations in Oenothera biennis L.? Am. Nat. 47: 116-121. F 1913. INDEX TO AMERICAN BOTANICAL LITERATURE 133 Durand, E. J. The genus Keithia. Mycologia 5: 6-11. pl. 8i. Ja IQI minis Keithia thujina sp. nov Farlow, W. G. The change toi the old to the new botany in the United States. Science II. 37: 79-86. 17 Ja 1913. Fernow, B. E., and others. Forest conditions of Nova Scotia. i-xi + I-93. Ottawa. 1912. [Illust. + maps. Published by the Commission of Conservation, Canada. Gainey, P. L. The effect of toluol and CS; upon the micro-flora and fauna of the soil. Ann. Rep. Missouri Bot. Gard. 23: 147-169. 1912. , Gates, R. R. Oecnothera and climate. Science II. 37: 155, 156. 24 Ja 1913. Gross, H. Polygonaceae nonnullae novae. Bot. Jahrb. 49: 340-348. I ee “aed ates uruguense and Muehlenbeckia Nummularia, spp. nov- from South Am Hallier, H. Uber friihre Landbriicken, Pflanzen und Vélkerwander- ungen zwischen Australasien und Amerika. Mededeel. Rijks Herb. Leiden 13: 1-32. f. 1, 2. 16 D 1912. Halsted, B. D., and others. Report of the botanical department. Ann. Rep. New Jersey Agr. Exp. Sta. 32: 311-399. pl. eee Ja 1912. Harris, J. A. Biometric data on the inflorescence and fruit of Crinum longifolium. Ann. Rep. Missouri Bot. Gard. 23: 75-99. 1912. Hasse, H. E. Additions to the lichenflora of southern California. 8. Bryologist 16: 1, 2. Ja 1913. Includes Dermatocarpon Zahlbruckneri sp. nov. Hicken, C. M. Nomenclatura botanica. Ilex paraguayensis 6 para- guariensis? An. Soc. Cie. Argentina 73: 360-362. Je 1912. Holden, R. Ray tracheids in the Coniferales. Bot. Gaz. 55: 56-65. Plt, 2.° -¥§ Ja F913. House, H. D. Woody plants of western North Carolina, 1-34. Darmstadt. 1913. Howe, C. D. Distribution and reproduction of the forest in relation to underlying rocks and soils. In Fernow, B. E., Forest conditions of Nova Scotia, 43-93. Ottawa. 1912. [Illust. + maps.] Howe, R. H., Jr. A monograph of the North American Usneaceae. Ann. Rep. Missouri Bot. Gard. 23: 133-146. pl. 7. 1912. Jeffrey, E. C. The history, comparative anatomy, and evolution of the Araucarioxylon type. Proc. Am. Acad. Arts & Sci. 48: 531-571. pl. 1-8 N 1912. 134 INDEX TO AMERICAN BOTANICAL LITERATURE Jewett, H. S. Plagiothecium geophilum (Aust.) Grout. Bryologist 46:8, 9.. -Ja ¥6¥s. Kearney, T. H. The wilting coefficient for plants in alkali soils. U.S. Dept. Agr. Plant Ind. Circ. 109: 17-25. 4 Ja 1913. Kunkel, L. O. A study of the problem of water absorption. Ann. Rep. Missouri Bot. Gard. 23: 26-40. 1912. Includes experiments with cultures of Monilia sitophila (Mont.) Sacc. Learn, C. D. Studies on Pleurotus ostreatus Jacqu. and Pleurotus ulmarius Bull. Ann. Myc. 10: 542-556. pl. 16-18. ai D 10tg: Lillie, R. S. The rdle of membranes in cell-processes. Pop. Sci. Mo. 82: 132-152. f. 1-3. F 1913. Lipman, C.B. Antagonism between anions as affecting ammonification in soils. Centralb. Bakt. Zweit. Abt. 36: 382-394. f. 1-3. 11 Ja 1913. Livingston, B. E. Adaptation in the living and non-living. Am. Nat. 47: 72-82. F 10913. Lloyd, C.G. The polyporoid types of Léveillé at Leiden. Letter No, 36. Mededeel. Rijks Herb. Leiden 9: 1-5. 15 N 1912. Lloyd, C. G. The polyporoid types of Junghuhn preserved at Leiden. Letter No. 37. Mededeel. Rijks Herb. Leiden 10: I-5. 15 WN 1912. ' Lloyd, F. E., & Ridgway, C. S. The behavior of the nectar gland in the cacti, with a note on the development of the trichomes and areolar cork. Plant World 15: 145-156. f. 1-13. Jl 1912. Lodge, C.A., & Smith, R.G. Influence of soil decoctions from sterilized and unsterilized soils upon bacterial growth. Ann. Rep. Massa- chusetts Agr. Exp. Sta. 24: 126-134. Ja 1912. Mackenzie, K. K. A new Carex from Alberta. Proc. Biol. Soc. Washington 25: 51, 52. 19 Mr. 1912. Carex atrosquama Mackenzie. Mackenzie, K. K. Western allies of Carex pennsylvanica. Torreya 13: 14-16. 8 Ja 1913. Includes Carex heliophila sp. nov. Manns, T. F. Two recent important cabbage diseases of Ohio. Ohio Agr. Exp. Sta. Bull. 228: 255-297. f. 1-26. IQII. Maxon, W. R. The tree ferns of North America. Smithsonian Rep. IQII: 463-491. pl. I-15. 1912. Metcalf, M. M. Adaptation through natural selection and ortho- genesis. Am. Nat. 47: 65-71. F 1913. Meyer, R. LEchinocactus macrodiscus Mart. Monats. Kakteenk. 22: 179-181. 15 D 1912. : INDEX TO AMERICAN BOTANICAL LITERATURE 135 Murrill, W. A. The Agaricaceae of tropical North America—VI. Mycologia 5: 18-36. Ja 1913. Includes new species in Gymnopilus (14), Crepidotus (9), and Pholiota (5). Several new combinations are made. Murrill, W. A. Illustrations of fungi— XIII.. Mycologia 5: I-5. pl. 80. Ja 1913. Includes Gyroporus peeian es ) Quél., Ceriomyces auriporus (Peck) Murrill, Rostkovites granulatus (L.) P. an so arpa (Peck) Murrill, C. subglabripes (Peck) Murrill, and C. bicolor bee) rome Norton, J. B. S. Maryland peas ae other harmful plants, Ann. Rep. Maryland Agr. Exp. Sta. 25: 1-71. pl. I-13. 1912. Norton, J. B. Methods used in breeding asparagus for rust resistance. U. S. Dept. Agr. Plant Ind. Bull. 263: 5-60. pl. 1-18. f. 1-4. 13 Ja 1913. Norton, J. B. S., & White, T. H. Rose mildew. Ann. Rep. Maryland Agr. Exp. Sta. 25: 73-80. f. 1-6. 1912. Ohiweiler, W. W. The relation between the density of cell saps and the freezing points of leaves. Ann. Rep. Missouri Bot. Gard. 23: 101-131. pl. 6. 1912. : Osterhout, W. J. V. The effect of anesthetics upon permeability. Science II. 37: 111, 112. 17 Ja 1913. Peltier, G. L. A consideration of the physiology aid life history of a parasitic Botrytis on pepper and lettuce. Ann. Rep. Missouri Bot. Gard. 23: 41-74. pl. 1-5... 1912. Perkins, J. Beitrage zur Flora von Bolivia. Bot. Jahrb. 49: 177-233. 14 Jal Includes descriptions of 20 new species contributed by various authors. (Con- tinued from p. 176. 27 Au 1912.) Quehl, L. Mamillaria Kunzeana Bédeker et Quehl spec. nov. Mo- nats. Kakteenk. 22: 177, 178. 15 D 1912. [lIllust.] Rehm, H. Ascomycetes exs. fasc. 51. Ann. Myc. to: 535-541. 31 D Includes Valsa saccharina Rehm sp. nov. from Canada. ee distributed as Urnula Craterium Fr. and Venturia Cassandrae Peck were obtained fr Safford, W. E. Annona diversifolia, a custard-apple of om foORs Jour. Washington Acad. Sci. 2: 118-125. f. 1-4. 4 Mr 1912. Safford, W. E. Pseudannona, a new genus of Annonaceae from the Mascarene Islands: together with notes on Artabotrys uncinatus and its synonymy. Jour. Washington Acad. Sci. 3: 16-19. 4 Ja 1913. Sammis, E. M. A vacation among the mosses. Torreya 13: I-13. f. 1-5. 8 Ja 1913. 136 INDEX TO AMERICAN BOTANICAL LITERATURE Schaffner, J. H. New and rare plants added to the Ohio list in 1912. Ohio Nat. 13: 36. D 1912. Schreiner, O., & Lathrop, E.C. The chemistry of steam-heated soils. U.S. Dept. Agr. Bur. Soils. Bull. 89: 5-37. 6N 1912. Seckt, H. Céntribucién al conocimiento de la vegetacién del noroeste de la Reptiblica Argentina. An. Soc. Cie. Argentina 74: 185-225. S 1912. Shaw, J.K. Heredity, correlation and variation in garden peas. Ann, Rep. Massachusetts Agr. Exp. Sta. 24: 82-101. Ja 1912. Shaw, J. K. The effect of fertilizers on variation in corn and beans. Am, Nat. 47: 57-64. f. 1, 2. Ja 1913. Stockberger, W. W. A literary note on the law of germinal continuity. Am. Nat. 47: 123-128. F 1913. Smith, C. O. Further proof of the cause and infectiousness of crown gall. California Agr. Exp. Sta. Bull. 235: 531-557, f. 1-28. D 1912. Smith, R. E., and others. Walnut culture in California. California Agr. Exp. Sta. Bull. 231: 119-398. f. I-96. Au 1912. Includes a study of oe walnut organism, Pseudomonas Juglandis Pierce, and other diseases of the walnu Snow, L. M. Pores and retrogressive changes in the plant associations of the Delaware coast. Bot. Gaz. 55: 45-55. f. 1-6. 15 Ja 1913. Stone, G. E. Report of the botanist. Ann. Rep. Massachusetts Agr. Exp. Sta. 24: — [3-107.] Ja 1912. [Illust.] Includes: Diseases more or less common during the year; Rust on Vinca; Bronzing of maple leaves; A notable elm tree; Frost cracks; Some observations on the growth of elm trees; The effects of positive and negative electrical charges on seeds and seedlings. Reprinted with separate pagination, Stone, G. E., & Chapman, G. H. Electrical resistance of trees. Ann. Rep. Massachusetts Agr. Exp. Sta. 24: 144-176. f. 1-3. Ja 1912. Stone, R. E. The life history of Ascochyta on some leguminous plants. Ann. Myc. 10: 564-592. pl. 19, 20. 31D 1912. Tidestrom, I. A new Salicornia. Proc. Biol. Soc. Washington 26: 13, 14. 18 Ja 1913. Salicornia utahensis Tidestrom. Transeau, E. N. The life history of Gloeotaenium. Bot. Gaz. 53: 66-73. pl. 3. 15 Ja 1913. Visher, S. S. The biology of south-central South Dakota. South Dakota Geol. & Biol. Surv. Bull. 5: 61-130. pl. 33-44. Je 1912. Weingart, [W.] Cereus laevigatus S.-D. var. guatemalensis Eichlam- Monats. Kakteenk. 22: 182-185. 15 D 1912. pans : BULL. TORREY CLUB VOLUME 40, PLATE 2 Vol. 40 96°08 BULLETIN OF THE TORREY BOTANICAL CLUB ee ee . APRIL 1913 Studies in the cytology of the Hymenomycetes, especially the Boleti MIcHAEL LEVINE (WITH PLATES 4-8) The discovery of sexual cell fusions in the rusts has given a great impetus to the investigation of the origin of the binucleated cells in the Basidiomycetes. It made clear that cell fusions without immediate nuclear fusions are possible and may result in an undoubted sporophyte with binucleated cells. An added importance is also given to the study of the nuclear divisions in the basidium as the stage at which chromosome reduction takes place. Since the appearance of Brefeld’s familiar observations on the origin of the carpophore of Coprinus stercorarius, it may be regarded as established that no specialized sex organs are necessary for the initiation of the development which leads to the formation of basidia. Brefeld (1876-77) grew the mycelium of this fungus from spores on dung decoction and described and figured two methods of carpophore formation without finding any structures visibly differentiated as sexual organs. He found that the carpophore might have its origin in a single mycelial thread or from a sclerotium, depending on the condition of the culture medium. Brefeld’s results seemed for a time to settle the question as to the existence of any form of sexuality in the Basidiomycetes. The question as to the significance of the hyphal fusions in the my- celium, however, still remained open. Two types of fusions between hyphae in the Basidiomycetes have been long known, [The Butietin for March 1913 (40: 97-136. pl. 2) was issued April 7.] 137 138 LEVINE: CYTOLOGY oF HYMENOMYCETES first the so-called clamp connections and second, the ordinary hyphal anastomoses. Such fusions are, however, well known in other groups of fungi with normal sexual reproduction and the possibility that they may have sexual significance in the Basidio- - mycetes seems remote. Hoffman (1856) was the first to figure the clamp connections between hyphal cells. He clearly described the slender tube joining two adjacent cells just outside their common cross wall and gave the name ‘‘Schnallenzellen”’ or clamp connections to these delicate structures. Brefeld emphasized the fact that clamp connections are not permanently open, for shortly after the tube buds out, a wall appears cutting it off from the cell from which it arose. Eventually, he claims, a second wall is formed separating the clamps entirely from each of the cells which it joins. The cell which puts out the clamp connection is always the one on the apical side of the transverse wall. Harper (1902) does not agree with Brefeld on the point that the fusion tube is cut off from both cells which it joins and interprets the clamps as a means of interchange of food stuffs. Lyman (1907) finds clamp connections arising very early. In one species, Corticium roseo-pallens, clamp connections are found in the germ tubes, while in Corticium subgiganteum no clamps at all are produced. Lyman studied artificial cultures of 75 species of Thelephoraceae, Hydnaceae, and Polyporaceae and finds accessory reproductive bodies, such as oidia in Daedalea unicolor, Lenzites betulina, Polyporus fumosus, Polystictus conchifer, Polystictus versicolor, and in Corticium alutaceum; and chlamy- dospores in Lentodium squamulosum, Poria incrustans, Radulum tomentosum, two species of Hydnum and one of Phlebia, Corticiwm vagum, C. effuscatum, and Michenera artocreas. Bulbils composed of a mass of hyphae were found in cultures of Corticium alutaceum. It would be extremely interesting, in view of the early origin of binucleated cells in the mycelium of many species, to know the number of nuclei in each of these types of asexual spores as af indication of whether they are to be reckoned with the gameto- phyte or sporophyte generation. It has been the common view that clamp connections and hyphal anastomoses have to do with food transportation etc. LEVINE: CyTOLOGY OF HYMENOMYCETES 139 R. Hartig (1885) however believed that such cell unions resemble a sexual act and he compared these phenomena with the fusions in the Conjugatae. He traced the development of Merulius lacrymans from spore to spore and describes hyphal anastomoses and clamp connections as common in its mycelium and crustlike carpophore. Meyer (1896-1902) describes the formation of hyphal anastomoses in Hypomyces rosellus. He states that when one hyphal cell comes in contact with another a fusion may take place. The walls at the point of contact disappear and the protoplasm of the two. unite. Soon afterward a new wall is formed which only incompletely separates the two cells. Meyer believes that the fusions of hyphae by anastomoses, involving the fusion of plasms not closely related, may have the same effect as cytoplasmic fusion in a normal fertilization. In view of the well established fusions found in the rusts, Voss’ claim (1903) that clamp connections and hyphal anastomoses are also present in this group has considerable interest. His observations, so far however, have not been confirmed. Ordinary pit connections for the transfer of food materials are undoubtedly present in the cross walls of the hyphae of all the higher fungi. Strasburger, 1884, maintains for Agaricus campes- | tris, that the cells in the stipe show so-called protoplasmic con- tinuity. Miss Wakefield’s (1909) observations on the conditions: governing the production of the carpophore in Schizophyllum commune and Stereum purpureum are quite in harmony with the view that the origin of the carpophore in the Hymenomycetes is in no way associated with a sexual act. She finds that the production of carpophores depends upon the rate of transpiration. ° Thus carpophores are produced when transpiration is slow while none are formed when evaporation is too rapid. The determination of the number of nuclei in the hyphal cells and the more exact study of their behavior has led to quite new conceptions as to the presence of sex in the Basidiomycetes. De Bary (1866) first observed the nucleus of the basidium in Corticium amorphum. Strasburger (1884) using alcohol and alum haematoxylin was able to demonstrate, beyond a doubt, that there are nuclei in the hyphal cells of Psalliota campestris and in 140 LEVINE: CYTOLOGY oF HYMENOMYCETES the basidium of Russula rubra, and Rosenvinge (1887) extended this conclusion to thirty-five species of Basidiomycetes. Rosen (1892) working on material from Lepiota mucida first observed nuclear fusions in the basidium. He concluded, however, that the secondary nucleus of the basidium resulted from the fusion of six or eight nuclei coming from the hyphae of the lamellae. Wager (1893-’4-’9) gave us the basis for most of our present day conceptions of the nuclear phenomena in the basidium. He followed Rosen in the conception that more than two nuclei may fuse in the formation of the secondary nucleus of the basidium (see table, p. 164). He even suggests the probability of a number of nuclear fusions occurring before the primary nuclei pass into the basidium. Wager observed that in the fusion of the nuclei their chromatic reticula become intermingled and that the nucle- oles finally fuse. He describes the nuclei of the basidium as having the typical structures found in higher plants, such as, chromatin, linin, differentially stained nucleoles, nuclear mem- branes, etc.; and was able to make out karyokinetic division figures. How the spindle is formed Wager could not make out, but believes that it comes in some fashion from an archoplasmic body. The chromosomes are six to eight in number. Dangeard (1895) was the first to establish the fact that only two nuclei fuse to form the secondary nucleus in the basidium of Tremella mesenterica, Dacryomyces deliquescens, Calocera viscosa, Craterellus sinuosus, Bovista plumbea, Nyctalis parasitica, Hydnum repandum, and Polyporus versicolor. He was the first also to affirm the sexual nature of this nuclear fusion in the basidium as well as in the ascus, the teleutospore, and the spore of the smuts. On these facts he bases his well-known doctrine that the Usti- lagineae, Uredineae, Basidiomycetes, and Ascomycetes have 4 sexuality which is essentially equivalent to that of the higher animals and plants. Dangeard claims that the young ascus, the young basidium, the teleutospore, and the smut spore are odgones and develop in one of the following ways. In the first case, the egg germinates by producing a promycelial outgrowth which produces sporidia endogenously as in the ascus, or exogenously as on the promycelium of the teleutospore and smut spore. In the second case the egg becomes segmented into a number of - LEVINE: CyTOLOGY OF HYMENOMYCETES 141 parts, by transverse or vertical walls, as in the Protobasidio- mycetes. In the last case the egg neither divides nor becomes. elongated but produces sporidia on sterigmata. In Tremella Genistae, Dacryomyces chrysocomus, D. deliquescens, and Polyporus annosus, Istvanffi (1895) claims that there is no nuclear fusion in the basidium. He further observed that there are two generations of spores formed on the basidia of Dacryomyces chrysocomus and D. deliquescens. On this point he has been con- firmed by both Juel and Maire, while Dangeard holds that in these forms the secondary nucleus divides only once and only one generation of spores is formed. Istvanffi also describes the formation of two generations of spores from the basidia of Hyd- nangium carneum. He also found uninucleated oidia and chlamy- dospores in Nyctalis asterophora, Psathyra spadiceo-grisea, Stro- pharia melasperma, Galera tenera, and Collybia tuberosa, which would suggest the existence of a fairly long gametophytic stage, which is capable of reproducing itself asexually in these forms. In Merulius fugax he finds that the mycelium is not septate. Juel (1898) points out that Brefeld’s distinction between Protobasidiomycetes and Autobasidiomycetes is supported by cytological evidence since the long axis of the primary spindle is either regularly parallel or perpendicular to the long axis of the basidium in the two groups. My own observations as described below do not show any such constancy in the position of the spindle in the basidium of the Boleti. Juel finds from six to eight chromo- somes, which vary in size, in the basidia of Auricularia mesenterica, and Exidia truncata. Maire (1902), while he emphasizes the existence of an alternation of generations in the Basidiomycetes, makes no contribution to the solution of the question as to the point of origin of the sporophyte. He claims that the cells of the stipe and pileus become multi- nucleated (table, p. 164) by amitotic division. He found, however, that the young mycelium of Coprinus radiatus has uninucleated, cells and produces uninucleated oidia. He shows clearly that the nuclear fusion in the basidium is not a true fecundation and proposes to call it a ‘“‘mixie,” pointing out that the morphological equivalent of sexual cell fusion must come at the origin of the binucleated condition. 142 LEVINE: CyTtToLOGy oF HYMENOMYCETES The basidium is a spore mother cell and not an egg and the nuclear fusion in it is associated with the phenomena of chromo- some reduction. Maire describes and figures a synapsis stage in the prophases of the first division and gives good figures of centrosomes, polar asters, etc., in a number of forms. In Sclero- derma vulgare he found a deeply staining granule on the wall of the resting nucleus, which he believes to be a centrosome. As to the number of chromosomes, Maire is plainly in error. His figures showing two chromosomes are due to poor fixations and the true chromosomes are undoubtedly shown in his protochromo- somes. He first observed the appearance of cytoplasmic threads connecting the nuclei and the sterigmata at the four-nucleated stage of the basidia. He believes that the nuclei move into the sterigmata by the contraction of the threads and thus finally reach the spore. Maire observed this process of nuclear migration in several agarics, a boletus, and a puff ball. Maire also discovered the interesting and as yet unexplained abnormalities of the basidia of Hygrophorus conicus and H. ceraceus. Their subhymenial cells and young basidia are uninu- cleated. The nucleus in the basidium, which is bisterigmatic, divides, forming two nuclei which migrate into the spores. Oc- casionally one of the nuclei in the basidium divides before entering the spore, in this case the two nuclei migrate to one spore while the undivided nucleus goes to the other spore where it divides. In Clavaria rugosa and Cantharellus cinereus he observed as many as three divisions of the secondary nucleus. In the latter species a variable number of spores are formed, from one to five, irrespec- tive of the number of nuclei. The observations of Wager, Juel, and Maire, as to the ap- pearance of the spindles, astral rays, centrosomes, etc., were con- firmed in the main by Ruhland (1901). He describes the nuclear phenomena in a number of Basidiomycetes (see table, p. 164). He denies that four spores are ever formed on the basidium of Hyd- nangium carneum, although the phenomena leading up to the four-nucleated stage are quite regular. He claims that either one or two spores may be formed. When two spores are formed two nuclei pass into each. In case only one spore is formed all four nuclei migrate into it. LEVINE: CyTOLOGY oF HYMENOMYCETES 143 Strands connecting nuclei and sterigmata have been observed also by Petri (1902) and Van Bambeke (1903). Petri holds that these fibers are extensions of the nuclear membranes. He counts five to six chromosomes in the first division; Van Bambeke agrees with Maire as to the number of chromosomes in the Basidio- mycetes. Harper (1902) shows the presence of six to eight chromosomes on the spindle of Hypochnus subtilis. In this species he found all the cells of the thin filmy subhymenial layer binucleated. ; Miss Nichols (1904) finds binucleated cells in the rhizomorphs of Hypholoma perplexum, Crepidotus, Corticium lilacino-fuscum, Dictyophora duplicata, Poria, Pholiota praecox, Lepiota naucina and Lycoperdon pyriforme, and holds that there is in Hypholoma perplexum an uninterrupted series of binucleated cells from the rhizomorphs through the stipe, pileus, trama, and subhymenium to the basidium. The germ tubes from spores of this fungus and Coprinus ephemerus, grown in pure culture, are commonly multi- nucleated; although binucleated cells were found in the mycelia of Coprinus ephemerus twenty-four hours old. Fries (1911!) figures and describes synapsis, longitudinal splitting of the spirem, spindles with centrosomes and astral rays in the basidium of Nidularia pisiformis. He holds that on the primary spindle six to eight chromosomes can be seen. In the second division the chromosomes in the equatorial plate are four in number while the number seen at the poles is but two. He observed also the strands connecting the nuclei and sterigmata and describes granules which he associates with centrosomes, on the walls of the basidium at the points where the sterigmata are to appear. Fries (19112) also confirms in toto Maire’s observations on the nuclear phenomena in Hygrophorus conicus and holds that only one nucleus is found in the subhymenial cell and young basidium. Kniep (1911) grew the mycelium of Armillaria mellea from spores in a gelatin medium, and reports that its mycelial cells are uninucleated throughout. He figures basidia formed directly from this uninucleated mycelium without the development of acarpophore. Whether the bodies Kniep figures as basidia are really these organs seems doubtful. Recent investigators have devoted much attention to the 144 LEVINE: CyTOLOGY OF HYMENOMYCETES nature and function of the cystidium, and Buller and Knoll have ascribed to them hitherto unsuspected functions. Buller (1910) as a result of elaborate studies concluded that the cystidia of the Coprini act as props and emphasizes the importance of the inter- lamellar spaces for the dispersal of the spores. In Inocybe astero- phora Buller describes the cystidia as excreting a ‘mucilaginous substance. Patouillard (1887), Massee (1887, 1894, 1904, 1906), Istvanfhi (1896), Topin (1901), Maire (1910), and Demelius (1911, 1912) hold that the cystidia are organs of excretion or are in some way related to the functions of excretion. Maire (1910) holds with Topin (1901) that the function of the cystidium varies with the stage of its development. In its young stage the cystid- ium is a storage organ of reserve food for the hymenium. Ulti- mately the cystidium becomes an excretory organ. Massee (1887) holds that the cystidia of the gill-bearing fungi are the terminal cells of laticiferous vessels. Their contents escape through a nipple-like filiform attentuation at the apex of the cystidium. Miss Demelius (1911, 1912) is of the opinion that the cystidia serve to protect the fungus from the invasion of insects. She emphasizes their variability in form, as in Collybia radicata, which has spherical, spindle-shaped and finger-like cystidia upon which excretion products may or may not appear. ' Knoll (1912) endeavors to prove that the cystidia are one- celled hydathodes comparable to the active water-excreting cells in the epidermis of the phanerogams. He maintains that trichom- hydathodes, as he calls them, are also found on all parts of the carpophore as well as in the hymenium. They are definitely shaped cells having a slender foot and a much expanded middle region while the upper part forms a sort of neck. The excretion forms a spherical drop at the apex. The fluid excreted contains a jelly which remains after the water has evaporated. Besides water, Knoll believes that the. cystidia excrete by-products of metabolism and so accounts for the appearance of crystals of calcium oxalate on their tops. He also holds that the hydathodes may have a protective function in the case of fungi with exposed hymenial surfaces. For Knoll, the cystidia of the Coprini are aberrant types whose true function remains obscure. LEVINE: CyTOLOGY OF HYMENOMYCETES 145 MATERIALS AND METHODS Spores of Pholiota praecox and numerous Boleti were collected by placing the pilei on circular sheets of filter paper and covering the whole with a bell jar. The spore prints were kept in sterilized Petri dishes. Miss Ferguson’s (1902) method of collecting spores was also used. The Petri dishes with the spore prints were kept in an ordinary refrigerator until needed. A great number of cul- ture media were tried; among those which gave the best results were string beans, and fresh horse manure prepared in a manner similar to that described by Duggar (1901). Cherry agar was prepared by boiling 250 gms. of fresh cherries in a liter of water. The cherries on becoming soft were strained through a cheese cloth and the skins, pits, and stalks were discarded. The juice of the cherries was again boiled with 15-20 gms. of agar agar and the decoction was then filtered and sterilized in an autoclave. Malt- beef agar was made by adding 25 gms. of Loeflund’s malt and 25 gms. of Liebig’s beef extract toa liter of water. This was then mixed with 15-20 gms. of dissolved agar agar and boiled and neutralized with n/1o00 NaOH. The solution was then filtered and sterilized. The spores germinated best in the malt-beef medium. In the case of Pholiota praecox 65 per cent of the spores germinated in this medium. Cherry and malt-beef agars were used for the propagation of the mycelia of several Polypores. The cultures were kept in dark boxes at temperatures varying from 23°—30° C. The early stages of spore germination from cultures in Van Tieghem’s cells were killed and fixed to slides by a modification of Harper’s (1899) and Lutman’s (1910) methods. The cover glass on which the culture is grown is removed from the ring and a few drops of the fixing solution are added. The cover glass with the culture is then inverted over slides covered with a film of albumen fixative and gently tapped till part of the drop falls on it. The spores in the drop become fixed to the slide by the coagulation of the albumen. The slide is then set aside till the liquid partly evaporates. The preparation is hardened by pouring graded alcohol over it, and it is then ready for staining. More fixing solution may be added to the culture drop and the process repeated till all the spores have been removed. A minimum of disturbance of the delicate germ tubes is achieved by this method. 146 LEVINE: CYTOLOGY oF HYMENOMYCETES Older mycelia of Pholiota praecox were obtained by sowing the spores in Petri dishes partly filled with bean, cherry, or malt agar. These were kept in the dark. After three days sufficient growth results to make the fungus visible to the naked eye. The cultures were then transferred to test tubes, from which subsequent cultures were made. Mycelia of Polyporus adustus, P. betulinus, P. destructor, P. versicolor, Collybia velutipes, and Coniophora cerebella were obtained in pure culture from the Association Internationale des Botanistes. These were propagated by transferring small pieces (2-5 mm. sq.) of the mycelium to Petri dishes with malt-beef, cherry, and bean agars. To secure perfect penetration of the fixative in the case of mycelia, the method described by Miss Nichols (1904) was used. Most of my material was collected in the vicinity of New York City. A number of Boleti were also sent to me from Woods Hole. In all, the carpophores of twenty-four species of Boleti and three species of Polypores were studied. Of these Boletus granu- latus, B. castaneus, B. albellus, B. versipellis, B. vermiculosus, B. glabellus, B. chrysenteron, B. indecisus, and B. pallidus were the more favorable for cytological work on the mature carpophore. I have not succeeded in germinating the spores of any of the Boleti. Flemming’s weaker solutions gave the best results, although Merkel’s, Juel’s, and Bouin’s gave fair fixations. Flemming’s triple stain was used. Heidenhain’s iron haematoxylin also gave favorable results. To Prof. R. A. Harper, on whose advice this work was under- taken, I wish to extend many thanks for his kind suggestions and criticisms. SPORE GERMINATION AND MYCELIA As noted above, the question as to the origin of the binucleated cells in the higher Basidiomycetes remains still unsettled. I fin that the spores of Pholiota praecox germinate readily and I have studied the germ tubes and young mycelium with reference to this question. Spores from spore prints, obtained in the manner described above, were sown in Van Tieghem’s cells with bean, malt-beef, and dung decoctions. LEVINE: CyTOLOGY OF HYMENOMYCETES 147 Within six hours, 20 to 30 per cent of the spores will germinate, and in general, there seem to be only slight differences between cultures kept in the dark and those exposed to the light. At the end of twenty-four hours, however, 60 to 70 per cent of the spores had germinated in malt-beef decoctions while only 40 to 50 per cent had germinated in dung decoction. The spores of Pholiota praecox in germinating do not swell or burst but push out a dense globular bud at the apical end, opposite the point of attachment. Germinating spores, at intervals ranging from six to twelve hours, were transferred from the Van Tieghem cells to slides by the modified stippling method, described above. They ‘were then stained with Flemming’s triple stain. The globular bud that first appears from the germinating spore is very dense and at first contains no nuclei. It grows rapidly into an ordinary germ tube and a nucleus appears in it, which is soon followed by another. I have not seen nuclear division figures at this stage, but soon two, four, and more nuclei can be found in the germ tube lying near the spore. In cultures fifteen hours old (PL. 4, FIG. 1) the germ tubes have branched and a large number of nuclei are present. The main germ tube is an outgrowth of the initial globular bud which is more or less permanent, and is still visible in older cultures. The cytoplasm shows a reticulated structure with larger and more or less numerous vacuoles. The nuclei are irregularly distributed through the cytoplasm and show no definitely paired arrangement. Few if any cross walls have been formed at this stage and the hyphal cells are beyond question multinucleated. The nuclei are very small, yet each one shows a distinct nuclear membrane, and a red staining nucleole, while the chromatin is granular and © stains a faint blue. In the further growth of the hyphae up to the forty-eight hour stage, new branches are formed from the bulbous initial bud near the spore, as well as from the main germ tube. Septa are still scarce in these stages; none are shown in FIG. 2, PL. 4. The diameter of the nuclei is nearly equal to that of the hypha. The lateral branches generally show several nuclei which are similar in all respects to those in the main germ tube. The cytoplasm is denser near the apical portion of the branches and fewer vacuoles appear in this region. The cells up to the eee 7 eight hour stage are all multinucleated. 148 LEVINE: CYTOLOGY OF HYMENOMYCETES Malt-beef agar cultures sixty-eight hours old were imbedded in paraffin and sectioned. The cultures were made directly by sowing spores in Petri dishes or the spores were allowed to germi- nate from twelve to twenty-four hours in the Van Tieghem cells and were then transferred to Petri dishes. Both methods gave satisfactory results. The mycelium reaches a diameter of a centimeter on the surface of the agar within three days. Entire masses of mycelium of this size, together with the agar, were cut out and immersed in fixing solutions so as to disturb the hyphae as little as possible. The hardening with alcohols must be very carefully done. If dehydration is too rapid, the agar shrivels. Considerable difficulty is encountered in staining such sections since the agar as well as the fungus takes the stain. In using the triple stain very short exposures and dilution of the orange G are necessary. Sections of material three days old show both binu- cleated and uninucleated cells making up the mycelium. Long multinucleated cells are also found, which are the germ tubes of spores that germinate late. Cultures of the same age do not all of course show the same stage of development. The cytoplasm (PL: 4, FIG. 3, 4) is less dense in these hyphae and stains better. The vacuoles are large and extend across the entire width of the cells. The structure of the nuclei in the uninucleated cells appears very clearly (PL. 4, FIG. 3). The chromatin is composed of delicate strands distributed irregularly through the nuclear cavity. The nuclei of the binucleated cells as shown in FIG. 4, PL. 4, are smaller but show the same structure as those in the uninucleated cells. In cultures three days old, clamp connections and hyphal anastomoses were first noticed. Lyman (1907) reports for Corticiwm roseo- pallens that clamp connections may be found on germ tubes imme- diately after spore germination. Cultures, from spores sown in Petri dishes in malt-beef agar seven days old, make a layer of myce- lium covering the entire surface of the agar. The mycelium from such cultures shows considerable numbers of multinucleated cells. Their cytoplasm is dense, the vacuoles are small and they resemble - the multinucleated cells in the younger cultures described. Binu- cleated cells similar to those observed in cultures three days old appear more frequently and uninucleated cells are also observed — ; occasionally. The latter are long and their cytoplasm shows large ‘a LEVINE: CYTOLOGY OF HYMENOMYCETES 149 vacuoles. Clamp connections may be seen at one or both ends of the cells and are very common. At this stage small concavo-con- vex bodies on both sides of the cross walls of the hyphae appear. These structures stain red with safranin and are dense, homogene- ous bodies. They have been interpreted by Strasburger (1884), Harper (1902), and others as indicating the presence of protoplas- mic connections between the adjacent cells. Hyphal anasto- moses, such as are described by R. Hartig (1885) and Meyer (1896, 1902), are also present. For comparison, I have also studied the mycelium of Collybia velutipes, which grows rapidly in malt-beef and cherry agar. Miss Cool (1912) and Biffen (1899) have both studied its mycelium. I find that the cells of the mycelium are distinctly binucleated. Narrower hyphae densely filled with cytoplasm were also found in sections of this mycelium. The terminal portion of these fila- ments becomes divided into uninucleated cells which give rise to cylindrical uninucleated oidia similar to those described by Biffen for the same species and Miss Nichols (1904) for Coprinus ephem- erus. Clamp connections are regularly present and in thick sections hyphal anastomoses can be also found. The results thus obtained show in the germ tube and early stages of mycelial growth that multinucleated cells predominate in Pholiota praecox, while in cultures two to three days old, multi- nucleated, binucleated and uninucleated cells are all present. The multinucleated cells may belong to younger hyphae as:noted. In older cultures binucleated cells mainly prevail although multi- nucleated and uninucleated cells may be found. The origin of the binucleated condition is not clear, but it is very apparent that it appears early in the vegetative hyphae and persists through- out the subsequent development of the mycelium and the carpo- _ phore. The cells of the mature mycelium of Collybia velutipes are all binucleated and here the binucleated condition may be regarded as fixed for the entire subsequent development, except as in the case of the stipe or the pileus, where the nuclei may divide, forming multinucleated cells. Cultures of four species of Polypores obtained from the Association Internationale des Botanistes were also studied. The € 150 LEVINE: CYTOLOGY OF HYMENOMYCETES species were Polyporus adustus, P. betulinus, P. destructor, and P. versicolor. The first two and the last were also studied by Miss Cool (1912). The material was propagated by transferring small pieces (2-5 mm. sq.) of the mycelia from each culture to a variety of agar media. Soft malt-beef and cherry agars proved the most favorable. Parts of such mycelia were fixed in Flem- ming’s, Juel’s, Bouin’s, and Hermann’s solutions but the most favorable results were obtained with Flemming’s weaker solutions. Clamp connections are of course abundantly present in the cells of these mycelia. I did not specially study the development of the clamps, but it is plain that the mature clamp is not cut off from the two cells which it joins, as Brefeld (1877) held for Coprinus ster- corarius. Only one cross wall is formed separating the clamp from the cell from which it arose. In well stained preparations hemispherical pads similar to those described above are visible on both sides of this cross wall in the clamp. This perhaps indi- cates only a partial closing up of the opening originally present. Similar pads may be seen also on the septa between many of the hyphal cells (pL. 4, FIG. 8). Hyphal anastomoses in these my- celia are also common. The hyphae of these Polypores (PL. 5; FIG. 5, 6) are made up of a series of regularly binucleated cells. In the mycelium of Polyporus versicolor (pL. 4, FIG. 8) un- doubted uninucleated cells are also present but they are not com- mon; the mycelia in cultures of P. destructor and P. betulinus (PL. 4, FIG. 7) may show non-nucleated cells which are joined to adja- cent binucleated cells by hyphal anastomoses and clamps. I have also studied the cells of Coniophora cerebella, also ob- tained from the Association Internationale des Botanistes. They are regularly binucleated; clamps and hyphal anastomoses are also present. It is plain that the binucleated condition is fixed in all these forms long before they proceed to the formation of a carpophore. My mycelial cultures all showed on their surfaces the familiar appearance of droplets of water, varying in size from small glob- __ ules 0.5 mm. in diameter, to large ones 10 mm. in diameter. The small drops are countless while the large ones are relatively few- Regarding Knoll’s (1912) contention that many of the Basidio- mycetes are provided with sp cial hair-like organs, trichome-hyda- LEVINE: CyTOLOGY oF HYMENOMYCETES 151 thodes, for the excretion of water, I may note that my microscopic sections of mycelia in the region where the water appears showed no such specialized organs for its excretion. I found no differ- entiated structure that could be possibly associated with such a function and I am of the opinion that all the mycelial cells can excrete water. CARPOPHORE AND BASIDIA OF THE BOLETI The carpophores of the Boleti because- of their soft fleshy consistency and rapid growth are more favorable for cytological study than the pilei of the Polypores. An abundance of material of the common Boletus granulatus collected in the fall of 1911 proved favorable for fixing and staining. A large number of other forms were also studied for comparison. Pieces of the stipe, ; pileus, ring,* and pores were fixed with Flemming’s weaker solutions and Merkel’s mixture. Flemming’s triple stain was used almost exclusively although a number of preparations were stained with Heidenhain’s iron haematoxylin. Longitudinal sections of all parts of the stipe, at, eae d and below the ring show it is composed of an undifferentiated mass of interwoven hyphae. The hyphal cells toward the center of an old stipe are more loosely interwoven as compared with those near the periphery and form what Ruhland has appropriately called a plectenchyma. The cells vary in diameter and present an appearance in cross section similar to that figured by Harper (1902) for Coprinus ephemerus, although the difference in diameter is not quite so great. The cross walls show the so-called proto- plasmic connections as figured by Strasburger (1884). Clamp connections on hyphal anastomoses are entirely lacking. Many cells in the plectenchyma are binucleated; the majority, however, are multinucleated. The distribution of these cells is not regular although there is a tendency for the cells in the center of the stipe to become multinucleated early. According to Maire this multi- nucleated condition is the result of an amitotic division of the two original nuclei in the cell, though no one has conclusively proved * The presence or absence of a ring is the generally accepted means of separat- g Boletus luteus and B. granulatus. In my m material, both annulate and exannu- Sins forms, otherwise indistinguishable, were collected at the same time and place, 152 LEVINE: CYTOLOGY OF HYMENOMYCETES the presence of such divisions. The cytoplasm in the cells of the stipe contains very large vacuoles. The nuclei are comparatively (PL. 4, FIG. 9) large and have typically red-stained nucleoles and blue chromatin. In very old stipes the comet-shaped nuclei figured by Ruhland also appear. The cell walls of the cells in the ring are mucilaginous and these cells may branch. They are regularly binucleated (PL. 4, FIG. I0). The upper surface of the semiglobular pileus of B. commie is covered by a viscid layer of slime. The flesh is thick; the pores are relatively short. Small cushion-like glandular bodies are conspicuous about their mouths. The flesh of the pileus is made up of a meshwork of inter- twining hyphae with numerous air spaces. The cortical cells of the cap are similar to those in the interior, but their walls are gelatinous. In very old material the mucilaginous layer is very thick and surrounds almost all of the cells down to the pores. The cells of the flesh are short and are almost invariably binu- cleated (PL. 5, FIG. 11). I have found however a few cases of cells with from three to four nuclei. In old material of B. granulatus the trama, the subhymenium, and the hymenium are very distinctly differentiated. The trama is composed of bundles of long parallel hyphae. The straightest filaments are found in the center, while toward the periphery they become more interwoven. The terminal branches of the hyphae of the trama may be traced directly into the subhymenium as is well shown in B. vermiculosus, B. glabellus, and B. pallidus. The cell walls are thick and gelatinous, and take a blue color with the triple stain. Gelatinous material may also fill the intercellular spaces. Two nuclei are found in every cell. Frequently in the nuclei of the ring (PL. 4, FIG. 10), flesh, and trama a darkly staining granule is found on the nuclear membrane, to which the chromatin of the nuclei seems to be attached. The position of this body varies; it is sometimes found lying opposite the nucleole but frequently it is near it. This body resembles the central body described by Harper (1905), for the mycelial nuclei of the mildews- The subhymenial cells are short and are binucleated. Their cell a walls do not become gelatinous, and their cross walls as also those — of the trama and flesh show the characteristic hemispherical pads a oy LEVINE: CYTOLOGY OF HYMENOMYCETES 153 described above. The basidium mother cells in B. granulatus are formed as in all other cases by the division of the subhymenial cells. I have not been able to find nuclear or cell division figures at this stage. The mother cell divides and the outer cell of the two may become either a cystidium or a basidium. Occasionally the last nuclear division is not followed by a cell division in the basidium mother cell of B. granulatus. A four-nucleated cell results (PL. 6, FIG. 45), which elongates and resembles a basidium. It is however decidedly different in appearance from the four nucleated basidium found in later stages. It is slender, more deeply seated in the hymenium and the nuclei are arranged serially. The sub- sequent behavior of such cells was not determined. Paraphyses have been described as elements of the hymenium since the latter part of the 18th century. According to Léveillé (1837) the term was first used by Montagne to signify a sterile basidium, but in more recent years the function of ‘‘space maker”’ has been attributed to them by de Bary (1887), Brefeld (1877), Fayod (1889) and Buller (1910). It seems to me that in all cases the paraphyses are really immature basidia; as is held also by Miss Demelius, who believes that some of them, at least, are, as she describes them, ‘‘derzeit nicht fertile Basidien.’”’ In the Boleti a number of basidia in similarly advanced stages are fre- quently found developing in close contact with each other. The cystidia of the Boleti appear either singly or in clusters. Such clustered cystidia are not uncommon, for Patouillard (1887), Demelius (1912), and Knoll (1912) have also found them. In B. granulatus such clusters are covered by a gelatinous excretion and are visible, as noted above, as minute dark specks about the mouths of the pores and over the surface of the hymenium. The individual cystidium is club-shaped; the entire cluster (PL. 7, FIG. 12) forms a cushion-shaped mass. Cystidia in all stages of de- velopment may be found in a single cluster. Fic. 12 “a,” PL. 7, shows very young cystidia which are approximately the size of the mature basidia. Fic. 12 “‘b,” 12 ‘‘c,” PL. 7, represent later stages of their development. The young cystidium (PL. 5, FIG. 13) shows a dense granular cytoplasm, which is somewhat fibrillar in the upper part of the cell; vacuoles are also present. The young cystidium is regularly binucleated; the nuclei are in all essentials similar to those found in the basidium. : 154 LEVINE: CyTOLOGY OF HYMENOMYCETES The subhymenial cell from which the cystidium arises is pro- portionately larger and may be frequently traced back into a long filament of binucleated cells in the trama (PL. 5, FIG. 15). The cell wall is slightly thickened (PL. 6, FIG. 14) and is entirely covered by the mucilaginous material referred toabove. The evidence is clear that the cystidia, in B. granulatus at least, are glandular structures. The two nuclei are small in proportion to the volume of the cell but the nucleo-cytoplasmic equilibrium is not disturbed since the increase in the size of the cell is apparently due to an increase in the metaplasmic substances which form the source of the material that the cell excretes. The mucilaginous mass stains a faint orange with the triple stain. The outer surface of the gelatinous mass becomes dense and dark in color and ultimately forms a thin pellicle over the entire gland (PL. 7, FIG. 12). The formation of mucilage does not take place through any specialized pore as Massee (1887, 1904) holds, nor is it associated with any localized region of the cystidium as Maire (1910) and Knoll (1912) contend, but apparently takes place over the entire surface of the cystidial cell. Later the cystidium begins to disintegrate. Its cytoplasm shows large vacuoles. The nuclei also begin to show signs of disintegration. The nuclear membrane becomes faint, the nucleoles and chromatin lose their affinity for stains and soon disappear. Subsequent stages show the cell wall shriveled and the nuclei and cytoplasm entirely gone. Single flask-shaped cystidia (PL. 8, FIG. 60) are found in all the species of the Boleti I have studied except B. granulatus. Their nuclei and cytoplasm are essentially similar to those of B. granulatus. Tubular cystidia are found in B. vermiculosus, B. albellus, and B. scaber. They are long cylindrical binucleated cells filled with a granular cytoplasm similar to that described for B. granulatus. The nuclei of the basidia of the Boleti I have studied are quite favorable for cytological study. The chromatin strands stain blue and the nucleoles a bright ruby red, with the triple stain. The nucleoles have about the same size relative to that of the whole nucleus as they have in the higher plants. The cytoplasm of the young basidium (PL. 8, FIG. 46) is finely vacuolar and stains a faint orange. With the growth of the basidium the nuclei (PL. 6 FIG. 16, 17) also increase in size. At the time of fusion their di- re. = a f: 3) =. = Z LEVINE: CyTOLOGY OF HYMENOMYCETES 155 ameter is several times that of the nuclei in the subhymenium. The fusion stages are abundant and easily studied. Thechromatin in the nuclei before fusion loses its reticulated structure and a number of strands appear (PL. 5, FIG. 70). The nuclei in fusing become appressed upon each other and the nuclear membranes: disappear in the region of contact. We have thusa two-lobed stage (PL. 6, FIG. 18) which lasts for some time, showing that the surface tension does not operate to round out instantly the united masses: of the fusing pair, as might be the case if they were merely vacuoles. The nuclear membranes, however, form a continuous boundary (PL. 8, FIG. 47, 48) for the combined nuclear cavities and the con- striction in the plane of fusion gradually disappears. The method of union of the chromatin masses is not easy to make out. The strands soon become mingled so as to be indistinguishable as to their origin. I am inclined to believe, howéver, that the union is nothing more than the approximation of the chromatin strands bringing them into close contact with each other side by side (PL. 8, FIG. 46). At any rate the chromatin masses from the primary nuclei become so intermingled as to leave no visible evidence of their two-fold origin at this stage. The nucleoles approach, come in contact with each other, and eventually fuse (PL. 8, FIG. 49), forming a proportionately larger mass. The basidium continues to grow and larger vacuoles appear in its cytoplasm. The secondary nucleus lies in the center of the basidium and goes through a resting period. At this stage I have also observed a dense oval body which stains red, lying in various positions in the basidia of Boletus granulatus, B. albellus, B. versipellis, B. indecisus, and Strobilomyces strobilaceus. In B. albellus, this body is sometimes found in the upper part of the basidium. Radiating from it are long strands of kinoplasm. In this species (PL. 7, FIG. 61) and in B. granulatus I have also found a similar structure lying between the basidium wall and the nu- cleus. In B. versipellis the body lies near the nucleus and resembles more or less the archoplasmic masses figured by Wager (1894) for Mycena galericulata. I have not been able to connect this body with the processes of nuclear division and I have no proof that, as Wager holds, it is the origin of the centrosomes and karyokinetic figures. 156 LEVINE: CyTOLOGY OF HYMENOMYCETES The first division usually takes place in the upper part of the basidium, although the prophase stages begin while the nucleus is still near its center. The chromatin strands combine to form a long slender thread which forms a more or less irregular coil fill- ing the nuclear cavity. The chromatin filament may become quite thick and thus resemble a post-synaptic spirem (PL. 6, FIG. 19, 20; PL. 7, FIG. 71; PL. 8, FIG. 50). I also found a parallelism of the spirem strands which suggests splitting (PL. 6, FIG. 21). My figures of these stages show a close resemblance to those of Fries (1911') for Nidularia pisiformis. The spirem now becomes segmented as shown in FIG. 22, PL. 6, and each — soon perigee os shorter and denser. The number of these chr tly varies and it is very difficult to count them. During yee division a large vacuole appears at the base of the basidium. The nu- cleus moves toward the apex of the basidium and the spindle figure appears. The nuclear membrane disintegrates and the chromo- somes are found lying in a faintly blue-stained fibrillar spindle whose axis is transverse to the longitudinal axis of the basidium (PL. 6, FIG. 23). The nucleole is found in the cytoplasm (PL. 8, FIG. 52, 53). In some of my preparations the nuclear membrane and nucleole seem to have disappeared before or soon after the spindle is formed, while in others both are present at the equa- torial plate stage (PL. 8, FIG. 51). The direction of the long axis of the spindle in the division of the primary nucleus in the Boleti ; does not always conform to the generally accepted view as em- _ phasized by Juel (1897) and others. I found that while the transverse position is common there are many cases in which the 5 primary spindle may be very decidedly inclined, at least seventy : degrees to the transverse axis of the basidium (PL. 7, FIG. 62). 1°@ have observed such cases in Boletus granulatus, B. chrysenteron, B. badius, B. alutarius, and B. albellus. The poles of the spindle end in small red-stained centrosomes from which long streaming rays extend to the center of the basidium (pL. 8, FIG. 51, 53) The appearance of the polar asters agrees well with that shown by Maire (1902) for B. regius. The cones of astral rays as seen in section resemble diminutive comet tails. The degree to which — the rays are developed depends upon the position of the pole in the basidium. If the pole lies on the wall few or no rays are visible. Vo tA) ie seve t) W = F z LEVINE: CyTOLOGY oF HYMENOMYCETES 157 Compare the astral rays in FIG. 72, 73, PL. 6 and FIG. 51, 53, PL. 8, with those shown in FIG. 24, 62, PL. 7 and FIG. 58, PL. 8. Well- developed polar asters are shown in my preparations of Boletus castaneus, B. glavellus, B. vermiculosus, B. versipellis, B. chrys- enteron, B. punctipes, B. griseus, B. subtomentosus, and B. cyanes- cens. The number of chromosomes is difficult to determine but in favorable sections I have been able to count from six to eight. They lie as shown by Wager (1894), Ruhland (1901), Juel (1897), Harper (1902) and others in the center of the spindle although no characteristically dense equatorial plate is formed (PL. 6, FIG. 72, 73)- The chromosomes are drawn to the poles and at the same time the spindle elongates until its ends (PL. 6, FIG. 25; PL. 8, FIG. 54, 55) touch the basidium wall. Astral rays may be still seen radiating in all directions from the point of contact. The chromosomes become densely aggregated at the poles. I have been able, however, to see distinctly, in perfect fixations, as many as five chromosomes just before they reached the poles (PL. 6, FIG. 25a). Undoubtedly the appearance of two chromosomes in the equatorial plate and diaster stages, as reported by Maire (1902) and Van Bambeke (1903), indicates fusion of the chromosomes due to imperfect fixations. In the reconstruction of the daughter nuclei, a nuclear mem- brane is formed about the chromosomes. The latter begin to stream out (PL. 6, FIG. 26) forming a reticulated structure, and small nucleoles appear. The resulting nuclei resemble in all respects the mother nuclei. In FIG. 56, PL. 8, the two daughter nuclei are shown attached to the basidium wall. Faint astral rays are still visible coming from the point of contact. The old nucleole may be seen lying in the cytoplasm. This persistence of the kinoplasmic rays is conspicuous, but they disappear before the second spindle is formed. The two daughter nuclei prepare immediately for the second division. The prophases are hard to study. I have observed in Boletus versipellis, that the nucleoles come to lie on the side of the nucleus toward the base of the basidium (PL. 8, FIG. 57) and pass out into the cytoplasm, after the disintegration of the nuclear membrane. The spindles show centrosomes with 158 LEVINE: CYTOLOGY OF HYMENOMYCETES long astral rays. I find that in B. versipellis also (PL. 8, FIG. 58) the two secondary spindles may be at an angle of about 10° to the longitudinal axis of the basidium. In these division figures, considerable variation is found in the angle formed by a transverse axis of the basidium and the long axis of the spindle. The nuclear membranes still persist at the equatorial plate stage (PL. 8, FIG. 59). In a polar view of the equatorial plate at least four distinct chromosomes can be seen. In the diaster stage however they are often seen as one or two masses (PL. 6, FIG. 28, 29). I have been able, however, to demonstrate that at the poles also there are more than two chromosomes, as is shown in FIG. _ 27, PL. 6. The secondary spindles may become elongated until the chromosome masses reach the wall of the basidium as in the first division, suggesting that the kinoplasmic rays are attached to the basidium wall and are pulling the chromosomes and centers to it. The young daughter nuclei grow rapidly in size (PL. 6, FIG.: 30, 31) and remain for some time attached to the basidium wall. They then begin to move downward toward the base of the basid- ium and it at once becomes evident that they are connected by faintly stained strands with small granules which lie on the upper wall of the basidium at the point from which they started (PL. 5; FIG. 32). The origin of these granules cannot be easily determined. From their size, color reactions, and position in the basidium it ap- pears that they probably may be the centrosomes, which became fixed to the wall of the basidium in the process of division. The position of the centrosomes on the upper part of the basidium wall indicates the position of the future sterigmata. According to Petri (1902) the strands are the stretched nuclear membranes. I have been able to follow the development of these structures. In the early stages of their movement the nuclei resemble the beaked nuclei (PL. 6, FIG. 33) found in the Ascomycetes during the process of spore formation. As the main body of the nucleus progresses farther from the cell wall the fibrils become longer and thicker and resemble in all respects the strands figured by Maire (1902) and Fries (1911). I believe the fibrillar strand _ ; possibly may be analogous to astral rays; though as Petri suggests — : they may be due to the pulling out of the nuclear membrane. — LEVINE: CyTOLOGY OF HYMENOMYCETES 159 At this stage I have also found another type of fibrils in the cytoplasm. These latter run irregularly but in the main length- wise of the basidium. It may be that they are indications of cytoplasmic streaming. _ As the sterigmata bud out (PL. 6, FIG. 34, 35, 74) the centrosomes and strands are carried upward; the centrosomes remaining at the apex of the sterigma. In mature sterigmata (PL. 7, FIG. 36, 78) the granule lies at the apex and the nucleus is still attached to it by the fibrillar strands. In this stage of development the four nuclei are found at or below the middle of the basidium (PL. 5, FIG. 37) but do not show any indication of fusing as Wager holds for Stropharia stercoraria. A small globular mass of cytoplasm, the spore initial, now appears at the end of each sterigma (PL. 7, FIG. 38). The growth is rapid; later stages are shown in FIG. 39, 40, PL. 7. On the upper, inner surface of the spore wall the centrosome is still visible (PL. 5, FIG. 41) and from it the fibrillar strand passes down through the sterigma to the nucleus in the basidium. It seems that the centrosome marks the apex of growth for the spore as well as for the sterigma. In the spores of B. castaneus (PL. 7, FIG. 75) a number of fibrils radiate downward from the centrosome through the cytoplasm but the strand which runs to the nucleus is thicker than the others. These fibrillar strands connecting the nuclei with the sterigmata and spores are present in practically all the Boleti I have studied. In the basidia of Merulius tremellosus I have also observed them extending from the apex of the spores to the nuclei below in the basidium. In Polyporus brumalis and P. lucidus I could trace them only from the sterigmata to the nuclei. Simultaneously with the development of the spores the nuclei begin to move towards the sterigmata. This migration is probably the result of the contraction of the kinoplasmic fibrils as claimed by Maire (1902). In FIG. 63, PL. 7, of Boletus albellus the nucleus is shown part way through thesterigma. Inthe same spore higher up another spherical, red staining body appears. Its lower por- tion is attenuated forming a long, strand-like fibril which extends in the direction of the nucleus. What the nature of this body is have not been able to determine. The nucleus (PL. 8, FIG. 64) divides soon after entering the spore 160 LEVINE: CYTOLOGY OF HYMENOMYCETES (PL. 5, FIG. 42). I have seen division figures in the spores of Boletus castaneus, B. albellus, B. punctipes, B. cyanescens, B. indecisus, B. glabellus, B. granulatus, B. chrysenteron, B. spectabilis, B. bicolor, B- griseus, B. subtomentosus, B. badius, and Strobtlomyces strobilaceus. The equatorial plate and subsequent stages come out very clearly and well-developed polar asters are present. Long astral rays can be seen radiating from the centers and extending to the ends of the spore (PL. 7, FIG. 79). The spindles are very narrow, but show clearly in the cavity of the nucleus, the nuclear membrane being stillpresent. The chromosomes are sharply differentiated from the spindle fibers (PL. 5, FIG. 76) but are so small and so massed together that their numbers cannot be definitely counted. The position of the spindles is either transverse or parallel to the long axis of the spore. The central spindle stretches in the anaphases and the chromosomes come to lie on the walls on opposite sides of the spore (PL. 8, FIG. 43, 65). The two daughter nuclei show all the essential features of the nuclei in the basidium (PL. 8, FIG. 66). In the spore of B. albellus, a second division occurs. The spore (PL. 8, FIG. 67) becomes very long and the two spindles which appear are similar to those previously described. The spindles may lie parallel or at right angles to each other, the latter case is shown in FIG. 68, PL. 8, where one of a pair of spindles is repre- sented. Fic. 67, 68, PL. 8, show clearly that the number of chromosomes is greater than two, as held by Fries (1911) for the first division in the spore. In the late anaphases, the chromosomes are found at the poles and appear to be fused into one or two masses. The nuclei are reconstructed and four small daughter nuclei result. The division here is not apparently conjugate. The spindles at least are not paired side by side, though the divisions are simul- taneous. In many spores, I have found three nuclei (PL. 8, FIG. 69). Two were small, while the third was larger and had prob- ably not yet divided. It seems probable that the condition found in these spores indicates the initial stage in germination. . The karyokinetic figures in the spores of these Boleti are very distinct and suggest that further study of young mycelia will make possible the settlement of the question as to the first appeat- ance of conjugate division and regularly binucleated cells. ‘ LEVINE: CYTOLOGY OF HYMENOMYCETES 161 Centrosomes and astral rays are strongly and typically developed. Although it is difficult to determine the exact number of chromo- somes certainly more than two are found at all stages. In the first division of the basidium where the spindles are large as many as six to eight can be counted. Aberrant types of basidia have been found in Boletus chrysen- teron, B. punctipes, and B. griseus. The abnormality consists in the appearance of mature sterigma-like projections while the nuclei are still in the process of division. FIG. 77, PL. 5, shows a basidium with one of its four sterigmata well developed. The division figures are perfectly normal, with the exception that they are almost perpendicular to the transverse axis of the basidium. THEORETICAL DISCUSSION I cannot agree with Knoll that the cystidia in the Basidio- mycetes are hydathodes. The cystidia of the Boleti I have studied are evidently modified basidia whose function is in some sense glandular. The quantity of material excreted is very large and in no way resembles the mucilaginous substance found about the trama cells. Just how this substance is excreted by the cystidia is not clear but in all probability it is formed just beneath the cuticle as described by Tschirch (1889) for the gland cells in the higher plants. As I have pointed out, the excretion products in the form of a gelatinous substance may be usually found covering the entire sur- face of the cell. In the case of Boletus granulatus, where several cystidia are found together, a cushion-like gelatinous mass is formed. These masses are the ‘‘granules’’ (PL. 7, FIG. 12) at the mouths of the pores. Knoll admits that a mucilaginous substance accumulates on the upper part of the cystidia of Psathyrella graci- lis, Galera tenera, G. tenuissima, Peniophora globulosa, Paneolus helvolus, and Coprinus lagopus and figures the cystidia of Collybia esculenta, Psathyrella consimilis, and Inocybe trechispora as entirely covered with it. Knoll has done nothing to disprove the contention of Lepeschkin (1906) who showed that the discharge of water from hyphal cells in general depends only upon the condition of the plasma membrane. Biffen (1899) finds that in the case of Collybia velutipes, watery drops may be exuded from 162 LEVINE: CyTOLOGY OF HYMENOMYCETES any part of the carpophore, particularly the pileus, and notes three different kinds of cells covered by a watery exudation. The well-known occurrence of water drops on mycelial cultures is another fact strongly against the conception of special water- excreting organs in the fungi. As I have pointed out, cytological study of the mycelial cells shows no specially differentiated organs for the excretion of water. The mechanical function ascribed to the cystidia by Buller for the Coprini is entirely out of the question in the Boleti. The pores need no such aid to keep them open. I have noticed in many species numbers of spores embedded in the mucilaginous covering of the cystidia. It seems that in this case the cystidia rather interfere with the dispersal of the spores than assist in it. The question as to the method of origin of the binucleated cells and the stage at which they appear in the development of the carpophore is of interest not only with reference to its bearing on the problems of the morphology and phylogeny of the Basidio- mycetes but also and even more in its bearing on the whole question of the nature of sexual reproduction. If a sporophyte with cells containing 2” chromosomes can arise by the simple omission of a cell division in a binucleated cell and if this process can occur at various points in a mycelium either simultaneously or over quite a period of development the fact is of prime significance in the interpretation of gametic unions of the more typical sort. As noted above, the absence of differentiated sex organs at the initi- ation of the carpophore must be taken as an established fact, but the possibility perhaps still remains that the binucleated cells have their origin by the migration of nuclei through clamp connections or hyphal anastomoses. Meyer (1896, 1902) and R. Hartig (1885) have argued on general grounds that such cytoplasmic fusions may have some sexual significance. The observations of Lutman (1910) and Rawitscher (1912) that in the smuts the nuclei do migrate through such connecting tubes are certainly suggestive. _ Voss’ (1902) claim that clamp connections are present in the rusts, if confirmed, must be certainly regarded as good evidence ; against their function as conjugation tubes, since sexual fusions of another type are present in the rusts. Voss’ observa LEVINE: CYTOLOGY oF HYMENOMYCETES 163 however, are not generally accepted, and his figures are incon- clusive. My own studies on the nature of the clamps and hyphal fusions are not conclusive, on this point. As described above, I have observed clamp connections and hyphal anastomoses in cultures three days old, but nothing that would clearly indicate nuclear migrations, though in some cases I have found empty cells adjacent to binucleated cells. It is well established that the binucleated cells do not arise as carpophore initials. They are present long before the carpophores appear and the stimuli leading to the production of the-latter seem to be vegetative and environmental. In order to bring out clearly the stages at which binucleated sporophytic cells are found I have tabulated all the available data as to the number of nuclei in the cells of the mycelia, rhizomorphs, carpophores, etc., of the Basidiomycetes so far studied. Twenty- seven species have been described as having regularly binucleated cells in their mycelia and rhizomorphs. The data show rather clearly that the binucleated condition does not originate at any definite point but may arise anywhere in the mycelium. The germ tubes (PL. 4, FIG. I, 2) are generally multinucleated. The young mycelium has multinucleated cells (PL. 4, FIG. 3, 4) although bi- nucleated and uninucleated cells are also found. Miss Nichols does not believe that there are conjugate divisions in the germ tube but positive observations as to the occurrence of conjugate divisions are sadly lacking. As noted above, I have found no case of conjugate division in the material I studied. As shown in the table, oidia and chlamydospores are reported by Istvanffi (1895), Biffen (1899), Maire (1902), and Nichols (1904), for many forms. I have studied such asexual spores on the mycelia of Collybia velutipes. The evidence seems to be that these spores are usually uninucleated and it would seem natural that the mycelia from which they arise should consist of uninucleated cells. Such spores are certainly not so common in the Basidiomycetes as they are in the Ascomycetes. They probably should be regarded as the asexual reproductive bodies of a gametophyte stage. As yet no binucleated spores from binucleated hyphal cells like the uredospores of the rusts have been found in the rumen NUMBER OF NUCLEI IN THE CELLS OF THE BASIDIOMYCETES* : Rhi Sub- | Young Observer Species Spore | Germ | Myce- iz0-| Stipe | Pileus | Trama| hyme-| ba- | CatPo- tube lium morph nidin’ | sidiam | PHore AURICULARIACEAE Tetvaniie. (2.3. Auricularia Auricula-Judae RRO Pip ava tee VS NEE AR A POs Rae es alle grnass a OURO ig Ue OU laa toc tes Mons Pee RR STR TPNIE NS be Ee Ga a ae ving SU a DSM neemnie ce eee: [ite de Gite avec alate ae 2 MR Said 2 2 Maire ? mesenterica Dicks. Pee ks : As ee 2 or agg degen ha 1S A Fae A MS A 2 2 TREMELLACEAE Juel.... Ps: LEE SIN CASERET eel Cte ee eiiy bos eucnd ann’ BORD ny tos, eh ae caster eco rane cokes Lies area eaand ; wR eee OP Baars a. ys. [Seb acina oes Ree ie ans he ed ante ie Nees ; URAC Sane SEARO OUT sR Si cebe DNMEA Ug (eater an ek ee aA Re Maire.... an Ange se ies) tee ae A MOC Tay gem eMC i, PUT SAIS me SUBR el Wy i oat aati BON nds Pac ty (cy) Bn He WARM O ee ey "eeomstts Gente el Oe Ae tke eae ea DN epee Fig ee ee ee etree sae al oe ow oa ata and Bile Wsase in Maire..........|Vuilleminia pai ok ates '(Nees) Re Naire cee eat : AGED TRS Lael SE ered lB “Ay Hea PARRA, Vol ae ap 2 2 DACRYOMYCETACEAE PR Sty cK DIGLEF UL COP MER cue oss oie SIS POF ee ka nee CT 2 es ee RPS oles MPa Winns Dineen kee es ok 2 2 2 Dangeard....... Ue Fpbscasae. iy See ae bua nde y di Rrege L AS kee as RES ath (4 ee ia Coe EAL MR 2 2 2 2c. Dangeard....... Dacryomyces deliquescens : DENA ER « ; { “ i AN SEE SLT AS SO reas ete neon me NE age on Aegean ALE eae tc AY RPE RAO og aged SRR a 8 Gr Mi eM RM (er 0 eM NEL ani WREb eS ci5s cs ra yt eee SUA Cee Peeing ey Si idicl, cts + Narada su ce. ssc dbase e Bais tities Soersihh ovate Be eee sé ir ie Maire....... te ipa 5 Se 5 oe Beets ca cna at't rE NU | EEN Rant PC A LF 2 2 METEOR oie ies es « Guepinia helvelloides Fr.......... Pe pe ete ene ec a ee ta. Ge SRA TR kG Sass Linceleens vate & nana Pate Cob are 2 2 Ge ere ys ruja (Jacg,): Pats... Se eas he anes, oD ES ape ahs (PS Sas Fave tal CM a eee DEO 2 2 _ EXOBASIDIACEAE Maire. ... .|Exobasidium Andromedae PAPA ey ti tery trie yi ROM Cae ik Iban chee | Cou bet wy llc ates 2 Sng SRS ves as fe eee oe s Sambuct Pers... ceca Bac HS SC Wy Seog 3 cae es to i os 4) A eg ee DP 2 me oe ae ! sacri agg Piva hes hein vitae aces tee RENEE oo 3 SPR tele BE ei Ci epee SNe ey 2 HM ia Weg Pte Pot SHLADAWONAWAPH AO ADOTOLAD :ANIADT NUMBER OF NUCLEI IN THE CELLS OF THE BASIDIOMYCETES,—Continued , ; Sub- | Young . ¢ Germ | Myce- | Rhizo-| gj : Tp - ‘i . | Carpo- Observer Species Spore tthe Rent mci Stipe | Pileus | Trama i io ptiore ee eee ra liaise c.: HODROVG-COV CUCU GS (oo: 2) ae ema taao Cea te ies | een eee Be Pans eS fee Ve alee ad prolate ss Paceocete ate gn MCHO 4) 5. ls ee orticium iain fusca oa: tet eral gic ea Me Ree eee eey CES | cry PSM cc est soe ones -vicldcw eae 6a Mratel oa. MEOUNE TORE go are ge sans aah wea. 2 Ag SD er Ie EI Sos CORSE (et ie Od 2 pay Rl 2 Ree es, Craterellus cornucopioide ar aight e tes ig ete Operas fils. xis site's wicca cs [atatetere B Lome erase Ns eee a 2 2 2 RW ee I i ie i, Ot Petes eters! disk Pant Pe eg we wig cA eee 2 to ees cee yy ASE MISU OE eS ec Ses ee Vin eet al pee lene oe 2 2) (SR MABE oie es. Cyphett cia Sau Bre terres shy ahatay Wide hte aes ae iat bis sles Peiwlod wee 2 Beha. 2 2° FSO R Riare ee. rN ern ere ara arate eRe ies eee ood wie o Lee Soe 2 © pial | eS aa 2 Bh) ie Mare ess. Dictyolus baophias wi iS Se ART ates Bae Scag pet) 3. Lp gee URS TSI ED, et CO ey ESE Sean (We 2 2 eh ot OUCEE BUEBN. oo Fi eo ee exc d.. ST e208 AMR AY Vath Sees | crater eon Lease at a i BR 2) i aaa MOE, ss cess pile quercina ae PRO OS ge ley { eee TS. .. Saco [Wines Oslial anita Kaa gS Feta at 2 Si neeieite Bane. os... Thelephora eo. Es Se iti CN: 0S er Cre |e eae, mena eee ee 2 2 2 1 near Pe MeGR MO rete cer td adn aap Toor, cs laislacod abet ea tis cevdbelew ad 2 2 2 : CLAVARIACEAE MRGUE os Clavaria — 1 Ey EAS A oo a ca | a SNe ER (Mest ashes | geen | Cob come) | eMC Cees 2 2 bbe eee eee CAL] ROE Te ne nei Bis Sones Beever acln cecilia! xl gai etiova, rath [ietalaher Oe opiMah pocaNe) Geer the eos 6-00 2 2 Rosenvinge..... MRRDRER OOD Ys fy tt) cua a. Dee ee tila «a's: Weis ane cds hae sHiece a kiss [a 44 ST CE 1-4 MOLE 6. ok Sue atsba DME cs enue ie laste as Bee eo nah eee ae ates aug ea Pe atond ANG Hn okey pee eee 2 2 Re ciae Dangeard....... YANG FepandNe TAD PEE i BA eS oes one hy oo ee eR PoE ev Fe 2 Peet (baer ee DOORS rst es e PU RG Ue eUk SOT eee ees GLis ss cols aoe COS UE eee 2 Reedy Sew POLYPORACEA PIRES wc ccs en wh gaa ampla (Lév.) . NERS ee es tas a eee Gs a:a o | ei SRG ana at eR RO a | LAN Hates alt 1 A eG ne TOUTS AVMs GAs ater ay Ass She ees rie Nai Cae bates \cisc Jc \ierabeneadia: evierew bataieta en 2 2 ts lash AY See SHLADAWONHWAP AO ADOTOLA)) -ANIADT Y9OT NUMBER OF NUCLEI IN THE CELLS OF THE BASIDIOMYCETES.—Conlinued ‘ Sub- | Young | ., Observer Species Spore | Germ | Myce- | Rhizo-| stipe | Pileus | Trama hyme- ba- | ‘ arpo- - tube lium | morph eli ol eideann (Paes ‘ POLYPORACEAE 2c MOVING? oe S58. RGLOLUS BIADOLIUS fics cre ee AE ee yc eas cue tae, PR oi RRR RT ALS ee ekey rein ian i ea eo 2 2 { y- Levine. ........ IS RIO SARISES See rate oh NEO Sgr Sua Sua th ee aes i ca Salo helstecd oa) 2 2 2 ed Rewine. 6.4. 6... is POMNAMSS Scan PaO ces Caine ene: OP: LES Sane a ee ees sh a aradt | Ce 2 2 ae 4 OTe Gy epg eae ea ee St VORIMS IS TOMI irs are ges am ius 2 eee ES Maa Ore Gain rae atl ay wearer 2 2 ea. 1s 0 Sa a a a SiS OPUS UOTOMELE Io es Sate ye Oak gs fh ante ey Peete ois ae aclu nels cobe oae es 2 2 Sis : PAVING: o4605.. 35 OF Se ORCS DRLEG rs iota ae att cna. Alen tai rele eee are ee braid ss Mavis a tase alee 2 2 { Sees Wervane ee a5 Fistulina hepatica PUR CON Orth nega er Fo oS At DR RE Rennie Boke Karas Gi eee Oa mete Dee ey MS NO er Behe Sate EOS (Pe ea Race eis aie tek Be ee PCs tee etal g oe tig 2-4 2 2 oan Uae So BRAN Ono 6. is Lenzites sy Mia re ee a TTS CREPE Ok wis coh wiceiee [SMe Pet va tae 2 2 2 7 Bb act se ie dstvaniG. 5... Meru uli PEOGM Se eae ee ee hee O ks: iS Du feed ethane (peowenrer a el, Cetage Me abate iC) as ca US ea ED 0 OT eae evar a “lac sahdrnied CVD OP abr ecco Tetris s,s 7 POSS AS Re FG Baa a et Aura ea eee ee Es a ap ee PROS se. os Pra PRUNE Gl inns eit aren eee ne ere GN Raum Lg anys I RS i Bi 8 oak es Pe ae 2 BONS at io “i! Does Oe ie pare M caption sees v. : J my ere eG Le ox eels sides hc Ca prise soo fee Wikis whoa wou 2 2 SE ae Pol: yporus << raga tes “eS ROS giay Be inet! Weoley Dae Bcd Samana ge atl CERES CEE Se GL ah ne ae Levine......... ake a ratta inact: « soem NN ie VU atc ee citer. > ASAE PEER od dinate EN 2c kati RUM ae Vic ARCaH Nee Ras Petvarittres. <3.35 ne BUNGENG ea cs ore Lewy cates { y . Hs (20 Ca aan ih reat Sa ey DRONSY, ep One tee messes (eee Sino Ciera sina ota Seay noS | WAU Se Rial VL meg vs 2 ana nd Reg eee 9 (aa Levine...... v LORE Ear 21g SOR RUA AER AS a un oheae apeee sangre Ma Ure Une (es SN A PRO RINE ec) Si JAI) SOE Nay CMR (atthe an Ante tty ange * Boleti and Polypores in which I found ea spores and binucleated cells in the ee aig and hymenium are Boletus uae, Ba alutarius, B. badius, B. bicolor, B. castaneus, B. chrysenteron, B. cyanescens, B. felleus, B. griseus, B. indecisus, B. luridus, B. Rt: “ naaan B. scaber, B. spectabilis, _ sublomentosus, Strobilomyces strobilaceus, acne Masi P. lucidus, and Merulius 99T SHLAIAWONAWAHF, AO ADOTOLAD) :ANIADT NUMBER OF NUCLEI IN THE CELLS OF THE BASIDIOMYCETES.—Contlinued Sub- Trama | hyme- nium Observer Species POLYPORACEAE Dangeard....... Polyporus versicolor L.......... ais ac o's ns Te ones at ia oe gn tal 1A.) 1s) | OEE Ghia aa agra moe PAPAS fae Ste a AGARICACEAE Wager. . ‘ Amanita muscaria L.........-. (0 ee eae niherina D.C....... PEOIOD es oes Armillaria mellea Pree ile a tie’ SESS SSC) Sane has aN Sen err ne ee Realele eres ¢ Camarophyllus led Wulf... Ripe tse wies Cantharellus ci ve ASS pda ator nae OMe ac. os fe Cimereus Pers... 5... PPORTON Chesed vig ves § bs infundibuliformis es AIS $5) fabacformis oc... ss PONTOt cack oo Clitocybe or Bolt MAOIRE so Al eas cs ura a (Wulf.) Studer Pee seas bassline secre eee eels Istvanfi........ Coll Uybia tuberosa Be ee rs eS oo et ocx cee See ps oY peg ree Ladi velulipes Bevine cee... “ a 1, Gea pean Coprinus ephemerus Wichols..0. 6.5... . if meee. 2... a ra Maire........ ae he: Nichola.....6%... Cre Istvanfi........ Galera tenera Schnee oe PRONG acess as Gomphidins glutinosus Sch ee Ce ee eer oe ae eet et ie tak Yee ee) ee er ee ee ee ar Oe SO ee ee ee ee eee em ee ee else eee el wee eels eee eo Sao Germ | Myce- pore | tube lium Sis Bose peels e cas 2 I Ne wae iaibihie gees pa Dts 2) Soe ea) ger 1 Soares |S aera I 2 BES UE Say | PS ee eT AD site tite eek 5/40 PAREN el aie! 65. Rat eabitare wlan borg iel ss jE Oe PETES SG Cees BRG: 35, SRS 2G ear You” Sats Oe ier (ara en Tere ficceteeleie I oe! a Petree aes oe a FT Oe Jeveees 2 dae, Oa arate eo Ee lisieie ess I Pee ie ie ee ee we 7 SOG [ke Oy ae eS Rhizo- | stipe * orale ip Pileus bes WGC Fe Aaa Ca El ates Pe HE 2 2 atetenete 2 2 aire A, eee Ey, 2 ihe ge oea ne) aayea nalts 2 wees lo 2) 2 fa 9 a WR nortan ipa ge ie ere ee 2 re ee ee oC ae ener ote @ ee we eee Wee Cee Youn oo Carpo- sidium | Phore a esa tuald 2-3 ee Same fee es tM. Bice Ss. Hees MA Ft YA Bei Nils arti cree ee eigen Py BS 2 2 BONS iat D2) paca 2 2-0 year Say Karta 2 eh veka he paras Pe ere Me a A tats ere pia lelimae 2 * al BARE eR Dies ARP ieee naa SALADAWONAWAP AO ADOTOLAD :ANIADT NUMBER OF NUCLEI IN THE CELLS OF THE BASIDIOMYCETES.—Continued 89T ‘ Sub- | Young Observer Species Spore | Germ | Myce- | Rhizo-| Stipe | Pileus | Trama hyme- | __ ba- Carpo- tube lium | morph nium. | sidium | Phor AGARICACEAE « Maire .......|Hygrocybe miniata. BOO Sas ae cient eos Mr ce SLs a alee oapmccH tee dog 2s 2 2 ee Eee ae Hygrop us agathosmus | bays ee PRT Lae aL nS Ree Pipe Ren Obie Feary Pbk gery 2 2 rae eee [ONES ERS Se ee ACCUSES en ae GEN hs eS BP eee Liv sles Cahees 2 SST eae eS 2-00 I a 1 ae ae Be eee Ws COmICHS LS CEE TE OO AOR) BR aerate ean id Pag kk ames I a See OS eee Mt He etraal ane mae hie yk eg ets FS Se aoe eae ee yr , 2 2-4 I a Peay paca ae ee eee Me WCE eis oe eet tw eee Seite s oes clea d Meek ewe 2 2-0O 2 Ria Miia ge Maire. . “ye ifpholoms ese at Mele ea reese. Lc) oS CE AER Daniele: Wescan ese, Renmei 2 2 te, Saran, ae DIA. ices fascicuiare Tiida sc this S08 es Ss ten ta hess Weices Siete ete eee 2 2 eae 10. Pichole.:. 66.6. perplexum Phe o,6.05 060046 fe hc. 2 at ee 2 2 2 2 2 aL Poe ae a a ee 33 SUD elerise tere hs ney liek esa tee eee OHI Lg TRESS (IDO ree, rs tare 2 2 2 Hest Meee 3.42)... Lacthrins déliciosus Te. oc. i ok oe ee as oot SS SENT Rees Nees) Wann See 2 { phat Ne Pe renee bb t eager WA as Goes mee yr OBS on es Oat PNG eet Pik eye: a Vasa a ee a re es ‘hice ee ee Ve) De! aww ie Mies se a eee mn A ad tir ce At oe Bc ay Shichi get CAL ele Mie el Gite whee ea bw 2 Ee eee ayes uhland........ Lepiota cepacstipes REE Pe Beat eee ee) eM > Pe eres Marea cc eee irae) Cane rea oe ath arg Bae od oe eras MIUMERG. as Sy. acino-granulosa Henn Bernabe say es israel he Ma UED WAG y\. Pa MPM, tae wget y aieaevad Sie dos less 2 2 2 EI ees eee de SOM cao a eM Sy Meroe 7 oh PS Vaart iS ene eee 2 5 te ann a ere eee wae ee ee SON Ra ri ae on a hal yaa dale ao eels Lee Uh ck cree 4cy. .|Mar MS RCOTODOUNEE PE es 8 ea ae epee Ramee cre ty ias dios Ue LT MEER VUNG aE MEL ply hk ? he eB ees scene flerialt SUM ae Se mtcne oder th RN less le Peewee ewe a geepo ds Kes 2 BORG eee s ee ae ee me RS Se OBES eR il 2 Pee Bea Te Plea Tae bio pelg ty te reed CoRR SP TG EAE a yong Ves ace 9 8 ohh ps cgaralic ficcia BiBNR’s Bie ee hots 2 ee hte ee ES GEE ye alae s aed es eb 'ee 2 BS ee a, Ni ee hee tase SARS TA COS Saka oa A a eg Ciera FO 2 PEN Oe lain oe PAGE OSes RP CMM eras aa 2 agit tea og ANIADT SALADAWONAWAPT JO ADOTOLAD NUMBER OF NUCLEI IN THE CELLS OF THE BASIDIOMYCETES.—Continued A Sub- | Young Observer Species Spore | Germ | Myce- |} Rhizo-| Stipe | Pileus | Trama| hyme-| ba-_ | Carpo- tube lium morph fe sidium | Phore AGARICACEAE BORE wie 7 1 OLIN PROUCOR oie Gin sis a § Wie wields eee oe Se wy oa) Be Hered ee ek creeds as ch hacwlnen Wate ate Arar ney Nichols : i Eris t GRM ONE a Gccur ene IRI eae; a Se Co ete orcs oeeaes Oe a pe > PREM seh eR aie te La hace IME, RN tae Fo ee (Pleat TOTS SS ae ae PUGS COPUUNUS. 5 yk ee eS Bee SI ate A NE TS os Pansy veep ees ote oye Ss ee) a) { eld MOORS iy vir cics eos Psalliota onan Be aS ney EP eee aid tare oie Sh ale plod mae a Spouse cece SAW woes ¥ 04 SAN ar: OU SS mpes ris Li; Boy ana yar uueretaa a aia ge ca atars Br eee yoke Wee es BOO Vega Ls 2-00 2 Be tse pe: Boren. ; Pate SS Siig eR et cea gn Meas Eee 2 yale le RSE PO OE (ak An a Seen ar RNY SP a 6-8 [vious Strasburger ey : eo ‘Strasburger. . as toy en is Sch. Beers aiue co macan ie aa catn cecum oa AS catia Papeete (ae 0.0, Peete Jat seems tice pn he nie Calle Heep ORG es sees og OL IU A ua Oh a eR Ra Rea Mae Me sired aie a2, Sale a eciny geri p ee a hor wd seo Ae ete hase a Psathyrell crenata aici Les aya PEE ar RCE IBD et Cece Bi Na oa > sel Geet eile I ss Sai ee gee ae 2 Diary ort cut peSemenils pe em a gee es ir et Sn. aes eas 2 2 2 2 Pi ah ae ea ee SASS RARE oh ok ea taney ey eee ners Spr rd cid ch Ca A ROC ee ar re ey Eee PF Nee 2 POR Re Ree see SNS MOUS NN MS SR Ig Si nee Niobe pe I i oa ihe) Sot NE RES Mk roc Th SNES TRE gla Mei a Cae Orc 2 as ME VED. eps Get REO eee ence aaa sey | ee No eS reach oteuda So Ro Po ae Dae eae Oa Siropharia melas perma Sacer Cea Se meee RE emcees mrs al ba a 4 OS ae Ean Rapa spel a tie RS er eae ae mas a eae gc Sec ann Me oan POMPEO DUR ir sh se ete a 8 sg Gen iva ENC a RR ane MOP is dst ee alae ie) oe) 2 > iikaie citings MEME 5. sons S WOME ie cod. c. See hes CEES Ba ete ge igs Cia kote eas tye NS hee te acl chee nye Bid daca PHALLACEAE POROISS..,.... — oe eer sa nat oes Babee bE DRS cls 7 aces AG asp CR ee Se ken ae lb Meee. ee. |\Phallus impudicus L........... Bee Gok a = | SOE, Oe Ua Pe he Eat ee, Cee a" ae, peeere HYMENOGASTRACEAE Van Bambeke. . Hydnangium carneum Wallr................ A ae car ee Be ep Oe) ve pene Pe eres eae 2 Be APS sidan’ Istva : A Re pic ah aera OS 2 5 eet a ae CR AR SM Vea ert NE in poke pots Wide lake teen's teed o Pale ecw he 2 re ee esr Miyh PO esi ee te ses 4 ge ais gee Sh ee a ena P aie i a Wee a Nan cel os ees oo es 2 Tae Rate Ruhland ah bi 2% Rela Sie ae ein greopies POR mea fo be ick | icra Vaal ined ties cos s 2 rae (Saag als ANIADY SHLAQSAWONAWAP, JO ADOTOLA) 691 NUMBER OF NUCLEI IN THE CELLS OF Tue Basipiémycettés—Continued . S Young Observer Species Spore | Germ | Myce- | Rhizo- Stipe | Pileus | Trama | hyme- ba- | Carpo- tube lium | morph nium | sidium | Phore LYCOPERDACEAE Dangeard....... Bovista —. Pere oer ene Lee os SS Sek MG Pails) Ol tere ren ASUS tH Evan reas, panei eruee igo aris eS ae rsa TURD IONUG TERR es ca aie cca ico Pace Sees vice t os Ve SEEPS eS ones cls 2 pe a a PARED ios se coperdon caclatum Bull... . 6 co oo ce cee OES 5 Opie Teas Mopar ees, iS eine ee Gira ay 2-00 2 teed Sher cere MANE ste... “f exci puliforme Scop. <6 co 66. Fcc 8k os Bee POV a Re SR Lee aS Sie ne 2 2 2 DAOHO es ssc fe ptt = 522 BB 0S OSS ala Beets is boty os Rata ee oe he ke 2 ea cy a Prichola.. i. .... Bees: OPM Sok ae areas os" eid egien, © BE emie N elas, Og Paw PgR ced gc o ee Sa UY Stn SAR Gi yon SON LA Se eee IDULARIACEAE WIA ee ves OS MNPSHINS Co re ea oe egret eh ar aieG he eRe rut os Pibey os boi yo ee hee bauble ea. 2 2 2 je ere ae Nidularia ans Bt Bee gues gird y wen atits PSA 2 oC Se Rd GMa ibe Iclcas 4 Panama Waa neat 2 2 Bee so DiS OPIS TAB re eae ees es ee eo PARTS HRS des io Pee eins Bess eee bake Coes wees 2 2 SCLERODERMATACEAE Maire..... + -| Scleroderma WR oo 6g Cane os Se a ei CU SEES Sea DRESS: Saintes! Oke See 2 ae Seana o. = oidia cy. = cystidia c. = conidia M.B. = mycelial basidia ch. = chlamydospores co = multinucleated OLE SALAOAWONAWAH AO ADOTOLA) :ANIART LEVINE: CyTOLOGY OF HYMENOMYCETES 171 The question as to the possible morphological equivalence of the carpophore of the Basidiomycetes and the ascocarp of the Ascomycetes is a difficult one. The development of the ascocarp is in many cases at least initiated by functional or possibly non- functional sex organs. It is possible that in the apogamous Ascomycetes which have been reported as lacking an ascogone, the ascocarp may arise in the same fashion as does the carpophore in the Basidiomycetes. It is to be noted however that binu- cleated mycelia are not so far known to occur in these apogamous Ascomycetes. I am of the opinion that in the Boleti at least none of the cells of the hymenium may be properly regarded as paraphyses. They are all binucleated when young and are all potentially either basidia or cystidia. That certain of these cells serve merely as ‘“‘space makers”’ is an interpretation which appears to me to be without much proof. Certainly there are no paraphyses here which can be compared to those in the Ascomycetes. In the latter the paraphyses are typically gametophytic while in the Basidio- mycetes all the elements of the hymenium are just as certainly sporophytic. The constancy in the occurrence of a nuclear fusion followed by a double nuclear division in the basidium is now generally conceded. Uninucleated basidia such as those of Hygrophorus conicus reported by Maire (1902) and Fries (19117) are apparently rare exceptions. The young basidia of twenty-four species of Boleti and three species of Polypores which I have examined are all binucleated. The two nuclei fuse in all cases and the resulting nucleus divides twice, forming four nuclei. These four nuclei migrate to the spores and there divide again. It is an obvious conclusion that these two nuclear divisions in the basidium involve the reduction of the chromosome number but the small size of the nuclei makes it impossible to reach new or independent conclusions as to the nature of the reduction process. Fries (1911!) and Kniep (1911) hold that the first and second divisions in the basidium are respectively heterotypic and homoe- otypic. But it cannot be said that their figures give any very positive evidence on the large questions here involved. My prep- arations show clearly (PL. 6, FIG. 27) that the number of chromo- 172 LEVINE: CyToLOGyY oF HYMENOMYCETES somes in the second division is certainly greater than two. The number which can be distinguished is variable but more than two are always found. This is true even in the case of the very small karyokinetic figures of the second division in the spore as shown in FIG. 68, PL.8. That we havea reduction division in the basidium comparable to that found in the spore mother cells of the higher plants is hardly to be questioned on general grounds; but the diminutive size of the chromosomes makes the study of the process extremely difficult. It is certainly clear, however, that Maire’s (1902) and Van Bambeke’s (1903) two chromosomes in | both divisions, and Fries’ (1911) two chromosomes in the second division and in the division in the spores are the results of poor fixation causing the fusion of the chromosomes. My observations on the Boleti confirm the views of Wager (1903, 1904), Juel (1897), and Harper (1902), and it cannot be doubted that Maire’s protochromosomes are the true chromo- somes. | Maire (1902) and more recently Fries (1911) have given extremely interesting data as to the migration of the four nuclei from the basidium into the spore. The nuclei according to their observations are drawn into the sterigmata by fibrillar cyto- plasmic strands which extend from the sterigmata to the nuclei. My preparations show clearly also the kinoplasmic strands extend- ing from the nuclei to the points of origin of the sterigmata (PL. 7; FIG. 36; PL. 5, FIG. 37). Petri held that these strands originate by stretching of the nuclear membrane. The daughter nuclei in the telophases become attached to the basidium and when they move ~ downward in their well-known migration, the nuclear membrane is drawn out into a long fibrillar strand. The strand may be com> — pared toa much attenuated beak on the nucleus. Another interpre 2 tation of these strands is possible. As noted in the telophases ° the second division, the daughter nuclei come to lie on the wall of the basidium and apparently their positions mark the point of . origin for the future sterigmata. When the nuclei migrate dow? ward in the basidium, the centrosomes are left behind on t basidium wall. The sterigma, budding out at just this point carries the centrosomes upward (PL. 5, FIG. 37), at its apex. The strands connecting the centrosomes and nuclei are thus carti LEVINE: CYTOLOGY OF HYMENOMYCETES 173 upward with the growth of the sterigma. This growth of the sterigma naturally involves flowage of material into it and that this movement should express itself in the formation of fibrillar strands between the nucleus and the centrosome at the apex of the sterigma is certainly to be expected and indicates the relation of the nucleus to the metabolic activities of the basidium. The strands on this view would be comparable, in general, to the kinoplasmic astral rays. My preparations show further that when the spore buds out on the apex of the sterigma the centrosome is carried up at the apex of the growing spore. The fibrillar strands are extended also into the spore and appear running through its long axis - and maintaining a continuous connection with the nuclei which now lie at the middle or towards the base of the basidium (PL. 7, FIG. 40, 75; PL. 5, FIG. 41). It appears to be generally conceded that these strands are associated in some way with the passage of the nuclei into the spore. My observations in the Boleti certainly give good ground for the contention that they at least direct the course of the nuclei into the spores. The fact that, as I find, these strands extend through the sterigma and into and through the spore body, suggests also that they may be of significance for the apparently difficult process of getting the nucleus through the narrow neck of the sterigma. I find nothing in the Boleti to favor Fries’ (1911') claim that the migrating nuclei undergo a characteristic transformation in passing into the spore. It is, perhaps, not entirely proven that these fibrillar strands are actively contractile kinoplasmic elements which pull the nuclei into the spores but the appearances in the Boleti certainly suggest such a conclusion. Such a conception is, of course, not inconsistent with the view that such strands may also constitute a system of lines of flowage of material into the young spore. SUMMARY 1. Spores of Pholiota praecox germinated in malt-beef extract at room temperature produce multinucleated germ tubes. The mycelia in cultures forty-eight hours old are still composed of long multinucleated cells. In cultures three days old both uninucleated and binucleated cells are found. 174 LEVINE: CyTOLOGY oF HYMENOMYCETES 2. The mycelia of Collybia velutipes, Polyporus adustus, P. betulinus, P. destructor, P. versicolor, and Coniophora cerebella propagated from old cultures are made up of long series of binu- cleated cells. Clamp connections, hyphal anastomoses, and the so-called protoplasmic connections are numerous in all the mycelia. 3. The cells of the mature stipe of Boletus granulatus are all multinucleated, while the cells of the ring are binucleated. The cells of the flesh and trama of B. granulatus are binucleated. The. cells of the subhymenium are binucleated in all the species of Boletus studied. 4. The cystidia of the Boleti occur either singly or in small clusters forming gelatinous granules. In B. granulatus these cushion-shaped ‘‘granules’”’ are abundant at the mouths of the pores and scattered over the hymenium. The individual cysti- dium is binucleated. It is club-shaped and is deeply seated in the hymenium. The cystidia of the Boleti appear to be in some sense glandular in their functions. 5. The nuclear phenomena in the basidium are eieal in » all the species of Boletus examined. Fusion of the two primary — | nuclei of the basidium was observed in Boletus granulatus»B. ver- sipellis, B. glabellus, B. vermiculosus, B. castaneus, B. albellus, and B. chrysenteron. 6. The long axes of the spindles in both divisions are commonly _ transverse to the long axis of the basidium. Variations, however; appear in which the spindles are almost perpendicular to the transverse axis of the basidium. Centrosomes and well-developed astral rays are regularly present. 7. The chromosome number in the first division is from six to eight in Boletus granulatus, B. castaneus, B. albellus, B. vermicu- — losus, B. versipellis, and B. chrysenteron. In the second division the exact number cannot be determined. It is, however, always more than two. 8. At the end of the second division the centrosomes become attached to the walls of the basidium and the four daughter E nuclei are reconstructed in close connection with them. As the : nuclei move downward in the basidium they maintain their — connection with the centrosomes by means of fibrillar strands which are, perhaps, analogous to astral rays. The fibril strands apparently pull the nuclei into the spores. LEVINE: CYTOLOGY oF HYMENOMYCETES 175 g. The centrosomes mark the points of origin for the sterig- mata. They are carried up with the growth of the sterigmata and into the spores. They also apparently determine the apex of growth of the spores. 10. The spores in all the forms studied are uninucleated at first. The primary spore nucleus divides at once. The kary- okinetic figures are small but very sharply differentiated with well- developed centrosomes and polar asters. In the spores of B. albellus a second division occurs. 11. In Boletus chrysenteron, B. punctipes, and B. griseus, basidia with mature sterigmata are found before the completion of the second division. Normal basidia are also present. 12. An alternation of generations comparable to that in the Uredineae is also present in the Basidiomycetes. The sporophyte begins at some indefinite point in the mycelium and extends through the development of the carpophore. COLUMBIA UNIVERSITY, NEw York. LITERATURE CITED 1887. Bary, A. de. Comparative morphology and biology of the Fungi, Mycetozoa and Bacteria. 1899. Biffen, R. H. On the biology of Collybia velutipes. Jour. Linn. Soc. 34: 147-162. 1877. Brefeld, O. Botanische Untersuchungen iiber Schimmelpilze. Heft ITI. 1910. Buller, A. H. R. The function and fate of the cystidia of Coprinus atramentarius, together with some general remarks on Coprinus fruit-bodies. Ann. Bot. 24: 613-628. 1912. Cool, C. Beitrage zur Kenntnis der Sporenkeimung und Rein- kultur der héheren Pilze. Mededeelingen phyto pathologisch Laboratorium ‘‘ Willie Commelin Scholten.’’ 6-36. sterdam. 1894-1895. Dangeard, P. A. Mémoire sur la reproduction sexuelle des Basidiomycétes. Le Botaniste 4: 119-181. Igt1-12. Demilius, P. Beitrag zur Kenntnis der Cystiden. I-II-III Teil. Verh. K. K. Zool.-Bot. Gesells. Wien 61: 278, 322, 378. 1912. Demilius, P. Beitrag zur Kenntnis der Cystiden. IV-V Teil. Verh. der K. K. Zool.-Bot. Gesells. Wien 62: 97-113. 1901. Duggar, B. M. Physiological studies with reference to the germination of certain fungus-spores. Bot. Gaz. 31: 38-66. 176 1889. Igtt. Igi2. IgI2. LEVINE: CYTOLOGY OF HYMENOMYCETES Fayod, V. Prodrome d’une histoire naturelle des Agaricinés. Ann. Sci. Nat. Bot. VII. 9: 181-411. Ferguson, M. C. A preliminary study of the germination of the spores of Agaricus campestris and other Basidiomycetous fungi. U.S. Dept. Agr. Plant Ind. Bull. 16. (x) Fries, R. E. Uber die cytologischen Verhdltnisse bei der Sporenbildung von Nidularia. Zeits. Bot. 3: 145-165. (2) Fries, R. E. Zur Kenntnis der Cytologie von Hygrophorus conicus. Svensk Bot. Tidskrift 5: 241-251. Harper, R. A. Nuclear phenomena in certain stages in the development of smuts. Trans. Wisc. Acad. Sci. Arts & Lett. 12: 475-498. Harper, R. A. Binucleated cells in certain Hymenomycetes. Bot. Gaz. 33: 1-23. Harper, R.A. Sexual reproduction and the organization of the nucleus in certain mildews. Carnegie Inst. Wash. Publ. No. 37. Hartig, R. Der achte Hausschwamm (Merulius asco Fr. ). Berlin. Rev. in Bot. Centralbl. 23: 123-129. Hoffman, H. Die Pollinarien und Spermatien von A 7: Istvanffi, G. von. Uber die Rolle der Zellkerne bei der Ent- wickelung der Pilze. Ber. Deuts. Bot. Gesells. 13: 452- Istvanfi, G. von. Untersuchungen iiber die physiologische Anatomie der Pilze mit besonderer Beriicksichtigung des Leitungssystemes bei den Hydnei, Telephorei und Tomental Jahrb. Wiss. Bot. 29: 391-445. Juel, H. O. Die Kerntheilungen in den Basidien und die Phylogenie der Basidiomyceten. Jahrb. Wiss. Bot. 32: 36I- 386. Kniep, H. Uber das Auftreten von Basidien im einkerniget Mycel von Armillaria mellea F!. Dan. Zeits. Bot. 3? 5297 553- (1) Knoll, F. Untersuchungen iiber den Bau und Funktion der Cystiden und verwandter Organe. Jahrb. Wiss. Bot. 59 453-50I. (2) Knoll, F. Uber die Abscheidung von Flissigkeit an und in den Fruchtkérpern verschiedener Hymenomyceten. Bets Deuts. Bot. eggs 30: 36-44. Lepeschkin, W. W. Zur Kenntnis des Mechanismus der 5 aktiven Wameetumichsiauas der Pflanzen. Beih. Bot. Cen: tralbl. 19': 409-452. eek 1910. 1901. LEVINE: CyToLoGy oF HyMENOMYCETES 177 Lutman, B. F. Some contributions to the life history and cytology of the smuts. Trans. Wisc. Acad. Sci. Arts & Lett. 16: I19QI—1225. Léveillé, J. H. Recherches sur Il’'hymenium des champignons. Ann. Sci. Nat. Bot. II. 8: 321-345. Lyman, G. R. Culture studies on polymorphism of Hymeno- mycetes. Proc. Bost. Soc. Nat. Hist. 33: 125-209. Maire, R. Recherches cytologiques et taxonomiques sur les Basidiomycétes. Théses présentées a la Faculté des Sciences de Paris. Maire, R. Les bases de la classification dans le genre Rus-. sula. Bull. Soc. Myc. Fr. 26: 49-125. Massee, G. On the differentiation of tissues in fungi. Jour. Roy. Micros. Soc. 7: 205-208. Massee, G. A monograph of the Geoglosseae. Ann. Bot. 11: 225-306. Massee, G. A monograph of the genus Inocybe Karsten. Ann. Bot. 18: 459-502. Massee, G. A text book ot fungi, 350. London Meyer, A. Das Vorkommen von Pratcaberbindungen bei den Pilze. Ber. Deuts. Bot. Gesells. 14: 280-281. Meyer, A. Die Plasmaverbindungen und die Fusionen der Pilze der Florideenreihe. Bot. Zeit. 60%: 139-1 78. Nichols, S. P. The nature and origin of the binucleated cells in some Basidiomycetes. Trans. Wisc. Acad. Sci. Arts & Lett. 15: 35-70. Patouillard, N. Les Hyménomycétes d’Europe. Part Perrot, A. Kernfragen und Sexualitat bei Sa pate Stuttgart. Petri, L. La formazione delle spore nell’Hydnangium carneum Wallr. Nuov. Giorn. Bot. Ital. II. 9: 499-514; Rev. in Bot. Centralbl. 92: 86, 87. 1903. Rawitscher, F. Beitrage zur Kenntnis der Ustilagineen. Zeits. Bot. 4: 673-703. Rosen, F. Studien iiber die Kerne und die Membranbildung bei Myxomyceten und Pilze. Cohn, Beitr. Biol. Pf. 6: 237- 264. Rosenvinge, L. K. Suir les noyaux des Hyménomycétes. Ann. Sci. Nat. Bot. VII. 3: 75-91. Ruhland, N. Zur Kenntniss der | ee Karyogamie bei den Basidiomyceten. Bot. Zeit. 59": 187- 178 LEVINE: CyTOLOGY OF HYMENOMYCETES 1884. Strasburger, E. Das botanische Praktikum. Jena. 1889. Tschirch, A. Angewandte Pflanzenanatomie, 193. 1901. Topin, J. Notes sur les cristaux et concrétions des Hyménomy- cétes et sur le réle physiologique des cystides. Thése Pharm, Paris. St.-Germain-en-Laye. 1903. Van Bambeke, C. Sur |’évolution nucléaire et la sporulation chez Hydnangium carneum Wallr. Mém. Acad. Roy. Sci. Belg. 54: 1-44. 1903. Voss, W. Uber Schnallen und Fusionen bei den Uredineen. Ber. Deuts. Bot. Gesells. 21: 366-371. 1893. Wager, H. On nuclear division in the Hymenomycetes. Ann. Bot. 7: 489-515. 1894. Wager, H. On the presence of the centrospheres in fungi. Ann. Bot. 8: 321-334 1899. Wager, H. The sexuality of the fungi. Ann. Bot. 13: 575- — 597: 1909. Wakefield, E. M. Uber die Bedingungen der Fruchtkérper- bildung sowie das Auftreten fertiler und steriler Stamme bei Hymenomyceten. Naturw. Zeitschr. Forst u. Landwirt. 7: 521-551. All the figures were drawn with the aid of a camera lucida and with the Leitz 1/16 objective. Figs. 11, 14, 16-45 were made with ocular 4; distance from camera mirror to drawing board 260 mm. Figs. 1, 3, 4, 5, 6, 9, 10, 46-79 were made with ocular 4; distance from camera mirror to drawing board 170 mm. Figs. 13, 15 were made with ocular 4; distance from camera mit- ror to drawing board 90 mm. Fig. 7 was made with ocular 3; distance from camera mirror to drawing board 90 mm. Figs. 2s — 8, 12 were made with ocular 1; distance from camera to drawing board 90 mm. Explanation of plates 4-8 PLATE 4 Pholiota — Germination tube at the end of fifteen and a half hours. Fic. 2. Same as Fig. 1 at end of forty-eight hour Fic. 3. Uninucleated hyphal cells in mycelium a days old. Fic. 4. Binucleated hyphal cells in mycelium three days old. Ses presents LEVINE: CyTOLOGY OF HYMENOMYCETES 179 Fic. 8. — hyphal cells from old mycelium showing clamp connec- S, anastomoses, hemispherical discs. Boletus ata ne Fic. 9. Cells from the stipe. Fic. 10. Cells from the ring. PLATE 5 Polyporus destructor. Fic. 5. Binucleated cells from old mycelium. Fic. 6. Two adjacent binucleated cells with clamp connections, Boletus granulatus., Fic. 11. inucleated cells from the flesh of the pileus. Fic. 13. Young binucleated cystidiu Fic. 15. Mature and cin young pasa showing connection with hyphal cells in the trama. _ Line over mature cystidium indicates surface of Rare excreti Fic. 32. Cross section of ‘sii showing four daughter nuclei telophase age ge. Fic. 37. Four nuclei a little below the middle of the basidium attached to the tips of the sterigmata by fibrillar strands. Fic. 41. Basidium showing the nuclei in the sterigmata attached to the tips of the spores by fibrillar strands Fic. 42. Nuclear division in the spore in equatorial plate stage. Fic. 44. Same as FIG. 42, diaster stage. Boletus Ep SOnENS, FIG. 70. ary nuclei in basidium Fic. 76. siaaent division in spore ebaiae spindles with centrosomes and astral rays, equatorial plate stage. Boletus chrysenteron. Fic. 77. An abnormal basidium showing one of the sterigmata fully formed while the two daughter nuclei are still dividing. PLATE 6 Boletus granulatus. Fic. 14. Mature and young cystidia. Fic. 16, 17. Young basidia with primary nuclei preparing to fuse. Fic. 18. Basidium showing nuclear fusion FIG. 19, 20. Basidium with nucleus in spirem stage. Lo: Se Fic. 23. First nuclear division showing eight chromosomes, metaphase. Fic. 25. The chromosomes moving toward the poles, ana Fic. 25a. The chromosomes at the poles before fusing, diaster. Fic. 26. Cross section of basidium showing two young danaktec nuclei, telophase. Fic. 27. Diaster, second division, a number of nuclei at each pole. Fic. 28, 29. Cross section of basidium, same as FIG. 27; chromosomes fused into two masses. Fic. 30, 31. Early telophase of the four daughter nuclei. 180 LEVINE: CyTOLOGY OF HYMENOMYCETES Fic. 33. Four nuclei beginning to move toward base of basidium, nucleus to right is attached to centrosome on basidium wall by kinoplasmic fibers. Fic. 34, 35. Early stages in the development of the sterigmata, fibrillar strands connect the nuclei and the tips of the sterigmata Fic. 45. Young four-nucleated hymenial cell in which nuclear division was not followed by cell division. E Boletus castaneus. Fic. 72. Cross section of basidium, first division, the chromosomes massed together Ne ett dene Fic. 73. Same as FIG. 72, six to eight chromosomes. Fic. 74. Nuclei moving to base of basidium. PLATE 7 Boletus granulatus. Fic. 12. A cluster of cystidia forming a gelatinous granule, from the mouth of a S ; Fic. 24. pan division in the basidium, anaphase, showing a number of osomes, the poles lying on the basidium wall. Fic. 36. ena with four nuclei attached to young sterigmata. Fic. 38. Basidium with very young spore buds. Fic.:.30., A se older sot fibrillar strands distinctly visible running the spore body connecting centrosomes an ei. Fic. | 40. older stage; fibrillar strands; nuclei moving toward the spores. ‘ Boletus albellus. Fic. 61. — nucleus with a conspicuous red-staining body lying between and the basidium wall. Fic. 62. ia division, anaphase, with well-marked centrosomes and astral ays rays. Fic. 63. Nucleus in a sterigma. tus castaneus Fic. 71. A spirem stage. Fic. 75. Basidium with the nuclei in the sterigmata, attached by fibers to the entrosomes at the apices of the spore. Boletus cr ahs Fic. 78. Nuclei énacies to the tips of sterigmata by kinoplasmic fibers. Pe. 79. Nuclear division in the spore; spindles with centrosomes and astral rays. Chromosomes in equatorial plate stage. PLATE 8 Boletus granulatus. Fic. 43. Nuclear division in spores, anaphase. Fic. 46. Basidia with primary and secondary nuclei. Fic. 47, 48, 49. Nuclear fusions in the basidium. Fic. 50. Spirem s tight LEVINE: CyTOLOGY OF HYMENOMYCETES 181 Fic. 57. A prophase stage of the second division. Fic. 58, 59. Second division in the basidium, cnene er centrosomes and astral rays, chromosomes in equatorial pla Fic. 60. Cystidium with two sharply differentiated eee to the left is a sidium. a Boletus albellus. Fic. 64. pore with single nucleus. Fic. 65. First division in agp diaster stage. Fic. 66. Binucleated s Fic. 67. Second avon, spindles with centrosomes and astral rays, chromo- somes in equatorial plate stage. Fic. 68. Similar eae only one spindle is figured; several chromosomes are s ; Fic. 69. A spore with three nuclei. New ferns from tropical America—ll MARGARET SLOSSON (WITH PLATE 3) The two species of Dryopteris here described for the first time belong to the group of D. pubescens. Both are from Jamaica, and each bears a most curious and misleading resemblance to the other. For the privilege of describing the first I am indebted to the kindness of Mr. William R. Maxon. This species is based on a single sheet, representing a rootstock and three leaves, two de- tached, originally from the Jenman Herbarium, labelled Nephro- dium luridum Jenman, in Jenman’s hand. It may be described as follows: Dryopteris lurida (Jenman) Underwood & Maxon sp. nov. Nephrodium luridum Jenman MS. Rhizome creeping, furnished with blackish rigid lanceolate or lance-linear acuminate scales up to 6 mm. long, with occasional unicellular gland-like processes and jointed cilia on their margins; similar scales on bases of the stipes; fronds clustered, pubescent, glandular throughout with capitate, often long-stalked and jointed, sometimes forked glands; stipes slender, up to 31.5 cm. long, dark brown at base, upwards brownish or stramineous or greenish, grooved on face; laminae up to 25 cm. long, up to 16.5 cm. broad, green, tinged with olive, ovate-deltoid, tripinnate, abruptly nar- rowed above at about the third or fourth pair of pinnae, their apices acute or acuminate, serrate, giving rise gradually to the pinnae and pinnules; pinnae alternate or opposite, oblique, stalked, mostly asymmetrical, those above the basal mostly ovate-lanceolate to oblong-lanceolate, the second or third pair often subequilateral, those above somewhat cut away beneath at base, the basal pair broadly deltoid or ovate-deltoid, up to 9 cm. broad at base, its inner inferior 2-5 pinnules on either side much longer than the corresponding superior ones and sometimes subbipinnate at base; other pinnules parallel with or overlapping the costa on the inner side, somewhat cut away beneath at base, acute, the larger stalked and obliquely pinnatifid into serrate or entire lobes, (183 184 SLOSSON: NEW FERNS FROM TRCPICAL AMERICA the smaller subdimidiate, serrate and decurrent; texture thin, membrano-herbaceous; pubescence white or whitish, setaceous, multicellular, the facial grooves of the stipes and the backs of the costae thickly coated with short fine soft hairs which spread part way up the midribs of the pinnules, larger coarser stiffer hairs scat- tered over the stipes, costae, backs of the veins on the under surface of the lamina, and between the veins on its upper surface; veins clearly visible, pinnate; sori apical on the veinlets, midway between the midvein and the margin of the lamina, sterile veinlets mostly extending almost to the margin; indusia glandular, not setose; sporangia glab arsely papillose. [PLATE 3, FIGURE I.] = Type in the Underwood Herbarium at the New York Botanical Garden, collected in Jamaica ‘‘1874-79.” The following specimens also are in the Underwood Herbarium: Jamaica: Mt. Diabolo, altitude 609 meters, April 2, 1903, Underwood 1825; vicinity of Hollymount, Mt. Diabolo, May 9; 1903, Maxon 1957; vicinity of Hollymount, altitude about 750 meters, May 25-27, 1904, Maxon 2311, 2260. Dryopteris leucochaete Slosson sp. nov. Rhizome creeping, slightly chaffy; fronds clustered, pubescent — and glandular; stipes up to 35.5 cm. long, brown and slightly — chaffy at base, above greenish to brownish, grooved on face; . scales soft, pale brown, lanceolate, acuminate, up to 3 mm. long, — with slightly ciliate or subglandular margins; laminae up to 20 — cm. long and 25 cm. broad, mostly subpentagonal, commonly — quadripinnate, abruptly narrowed above, apices mostly long- — acuminate and serrate; pinnae oblique, stalked, asymmetrical, © basal pair ovate-deltoid to deltoid; pinnules oblique, stalked to _ decurrent, several of the inferior elongate in the basal pinnae and shortened in the upper; segments oblique, stalked to decurrent, unequally ovate to subtrapezioid, cut away at base, pinnate OF obliquely pinnatifid to serrate; glands capitate, often jointed sometimes forked; pubescence white, setaceous, multicellular, fine and short hairs abundant on the groove of the stipe, backs of costae, and partly on the midribs of the pinnules, larger, coarsef scattered hairs on the general surface of the stipes, costae, backs of the veins, and between the veins on the upper surface of the lamina; veins pinnate; sori apical on the veinlets, mid between the midvein and margin of the lamina; indusia cons ously setose and glandular; sporangia glabrous; spores pa [PLATE 3, FIGURE 2.] a SLosson: NEW FERNS FROM TROPICAL AMERICA 185 Type in the Underwood Herbarium at the New York Botanical Garden, collected on shady forest land in Peckham Woods, Clarendon, Jamaica, altitude 762 meters, May 21, 1912, William Harris 11023. This species differs from D. lurida chiefly in its smaller, soft, pale, almost tow-colored scales, its larger, more finely divided lamina, and its setose indusia. Additional specimens are: Ja- maica; vicinity of Troy, altitude 600-660 meters, June 28, 1904, Maxon 2860; in rocky woodland, near Troy, altitude 701 meters, June 28, 1904, Harris 8770. Explanation of plate 3 Fic. 1. Dryopteris lurida; Jamaica; parts of the type specimen, reduced. Fic. 2. Dryopteris leucochaete; Jamaica; leaf, reduced, Maxon 2860. INDEX TO AMERICAN BOTANICAL LITERATURE (1913) The aim of this Index is to include all current botanical literature written by Americans, published in Ame Aue or based upon American material ; the word Amer- ica being used in the broadest s Reviews, and papers that ee exclusively to forestry, agriculture, horticulture, manufactured products of a origin, or laboratory methods are not included, and no attempt is made to index the literature oe bacteriology. An occasional exception is wholly to botany. Reprints are not mentioned unless they differ from the original in some important particular. If users of the Index will call the attention of the editor to errors or omissions, their kindness will be appreciated. This Index is crate monthly on cards, oe furnished in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitted ; each subscriber must take all nie published rants the term of his subscription, Corre- spondence relating to the card issue. should be addressed to the Treasurer of the Torrey Botanical Club. Adkinson, J. Some features of the anatomy of the Vitaceae. Ann. Bot. 27: 133-139. pl. 15. Ja 1913- Bailey, L. W. Diatoms of New Brunswick. Bull. Nat. Hist. Soc. New Brunswick 6: 387-418. 1913-. Bailey, W. W. A favorite wildflower. Am. Bot. 19: 16-18. F 1913. Epigaea repens. Banker, H. J. Type studies in the Hydnaceae—IV. The genus Phellodon. Mycologia 5: ears 10 Mr 1913. Includes Phellodon carnosus sp. n arnhart, J. H. Catalogue of the Cox — of Darwiniana. Jour. N. Y. Bot. Gard. 14: 2-29. Ja 19 Bendrat, T. A. The flora of Mohawk Hill, Py Y., north of the water- shed. Torreya 13: 45-63. 8 Mr 1913. [Illust.] Berger, A: -Agavé: Haysalde: Gane Ieee eee 1 Pe OPP F 1913. A plant from Mexico Bitter, G. Solana nova vel minus cogane. VII. Repert. Sp. Nov. 11: 481-491. 31 Ja 1913- Nine new ies are described Blake, S. F. A second local record for Rbgnchospors macrostachya. Rhodora 15:19. 7 F 1913- 188 INDEX TO AMERICAN BOTANICAL LITERATURE Britton, N. L. The Charles Finney Cox collection of Darwiniana. Jour. N. Y. Bot. Gard. 14: 1, 2. Ja 1913. Burlingame, L. L. The morphology of Araucaria brasiliensis. I. The staminate cone and male gametophyte. Bot. Gaz. 55: 97- 114. $14, 54 f. P-1T. 15 F 1913. Chamberlain, C. J. Macrozamia Moorei, a connecting link between living and fossil cycads. Bot. Gaz. 55: 141-154. f. I-12. 15 F 1913. Clark, J. J. Heliotropium anchusaefolium. Curt. Bot. Mag. IV. 9: pl. 8480. F 1913. A South American plant. Claassen, E. List of plants collected in Cuyahoga county and new to this county or to Ohio. Ohio Nat. 13:64. Ja 1913. Clements, F. E. The Alpine Laboratory. Science II. 37: 327, 328 28 F 1913. Clute, W.N. The cultivation of the Jris. Am. Bot. 19:6-12. F 1913. [Illust.] Clute, W. N. The mountain spleenwort and its relatives. Am. Bot. 19: 13-15. F 1913. [Illust.] Clute, W. N. The peli nut. Am. Bot. 19: 23, 24. F 1913. [Illust.] Canarium luzonicum or C. commune [Clute, W. N.] The pelican plant. Am. Bot. 19: 19-22. F 1913: [Illust.] Aristolochia gigas. Cook, O. F. A wild host-plant of the boll-weevil in Arizona. Science II. 37: 259-261. 14 F 1913. : Cooper, W. S. The climax forest of Isle Royale, Lake Superior, and — itsdevelopment. II. Bot. Gaz. 55:115-140.f. 15-30. 15 F 1913 Darling, C. A. Spring flowers. i-viii + 1-106. New York. 1 F — 1913. Eames, A. J. The morphology of Agathis australis. Ann. Bot. 27° 1-38. pl. 1-4 +f. 1-92. Ja 1913. East, E. M. A chronicle of the tribe of corn. Pop. Sci. Mo. 82: 225-236. f. 1-12. Mr 1913. j Fedde, F. Neue Arten aus der Verwandschaft der Corydalis aure@ Willd. von Nord-America. VIII. Repert. Sp. Nov. 11: 497-499 — 31 Ja 1913. % Two new species described, Corydalis isopyroides and C. pseudomicrantha. Fernald, M. L. Nuttall’s white sassafras. Rhodora 15: 147% 7 F 1913. Fullmer, E. L. Additions made to the Cedar Point flora cists the summer of 1912. Ohio Nat. 13: 78. 20 F 1913. INDEX TO AMERICAN BOTANICAL LITERATURE 189 Garcia, F., & Rigney, J. W. Grape crown-gall investigations. New Mexico Agr. Exp. Sta. Bull. 85: 3-28. f. 1-3. Ja 1913. Gates, R. R. A contribution to a knowledge of the mutating Oeno- theras. Trans. Linn. Soc. Bot. II. 8: 1-67. pl. 1-6. Ja 1913. Goodspeed, T. H. Quantitative studies of inheritance in Nicotiana hybrids. II. Univ. Calif. Pub. Bot. 5: 169-188. 9 Ja 1913. Gormley, R. The violets of Ohio. Ohio Nat. 13: 56-61. Ja 1913. Gortner, R. A., & Harris, J.A. Ona possible relationship between the structural peculiarities of normal and teratological fruits of Passi- flora gracilis and some physico-chemical properties of their expressed juices. Bull. Torrey Club 40: 27-34. 20 F 1913. Greene, E.L. The two Howells, botanists. Am. Mid. Nat. 3: 30-32. Ja 1913. Harris, J. A. Prolification of the fruit in okra, Hibiscus esculentus. Torreya 13: 33-35. F 1913. Heller, A. A. New western plants. Muhlenbergia 8: 137-142. pl. 16, 17 +f. 28. 31 Ja 1913. Includes Cressa vallicola, C. minima, and Caulanthus senilis, spp. nov. Hesse, E. Echinocactus Graessneri K. Sch. Monats. Kakteenk. 23: 2-6. 15 Ja 1913. [Illust.] Hull, E. D. Ebony spleenwort and shining club moss in northwest Indiana. Am. Bot. 19: 30. F 1913. Hull, E. D. Extended range of Viola pedata L. Rhodora 15: 18, 19. 7 F 1913. Humphrey, L. E. The Ohio dogbanes. Ohio Nat. 13: 79, 80. 20 F 1913. Kittredge, E. M. Some trees and shrubs of Rockland County. Tor- reya 13: 25-33. F 1913. Kunze, R. E. Echinocactus Wislizent Engelm. var. phoeniceus Kunze var. nov. Monats. Kakteenk. 23: 8,9. 15 Ja 1913. Lloyd, F. E. The induction of nonastringency in persimmons at supranormal pressures of carbon dioxide. Science II. 37: 228-232. 7 F 1913. Lloyd, F. E. Leaf water and stomatal movement in Gossypium and a method of direct visual observation of stomata in situ. Bull. Torrey Club 40: 1-26. f. I-3. 20 F 1913. Lunell, J. Adicea. Am. Mid. Nat. 3: 6-12. Ja 1913. Includes Adicea fontana, A. opaca, A. Niewlandii, and A. Deamit spp. nov. Lunell, J. Erigeron in North Dakota. Am. Mid. Nat. 2: 253-258- Jl 1912;—II. Am. Mid. Nat. 3: 1-6. Ja 1913. Includes descriptions of nine new species. 190 INDEX TO AMERICAN BOTANICAL LITERATURE Macoun, J. M. Additions to the flora of Vancouver Island. Ottawa Nat. 26: 143-148. F 1913. Mark, C. G. Notes on Ohio mosses. Ohio Nat. 13: 62-64. f. 2. Ja 1913. Murrill, W. A. The piiuniias of eastern North America. Mycologia 5: 72-86. pl. 85, 86. 10 Mr 1913. Nash, G. V. Winter flowering. Jour. N. Y. Bot. Gard. 14: 43, 44. pl. r11.. F 1913. Nash, G; ¥. Winter protection of plants. Jour. N. Y. Bot. Gard. 14: 30-37. pl. 108-110. Ja 1913. Nichols, G. E. Notes on Connecticut mosses,—IV. Rhodora 15: 3-13. 9 F 1913. Nieuwland, J. A. LEvactoma. Am. Mid. Nat. 3: 57-59. Mr 1913. Nieuwland, J. A. Some midland dogbanes. Am. Mid. Nat. 3: 53-57: Mr 1913. Includes A pocynum Carolini, A. tomentellum, and A. cinereum, spp. nov Parish, S. B. Coreopsis gigantea (Kellogg) Hall. Muhlenbergia 8: 133, 134. f. 27. 31 Ja 1913. Pennell, F. W. Further notes on the flora of the Conowingo or ser- pentine barrens of southeastern Pennsylvania. Proc. Acad. Nat. Sci. Philadelphia 54: 520-534. 30 Ja 1913; 535-539. 13 F 1913. Peck, C.H. New species of fungi. Mycologia 5: 67-71. 10 Mr 1913. Includes Amanita Peckiana Kaufim., Collybia subdecumbens, C. truncata, Ento- a mirabile, Inocybe minima, Leptonia gracilipes, L. validipes, and Puccinia striatospora, spp. nov. Petry, L. C. A protocorm of Ophioglossum. Bot. Gaz. 55: 155-166. f. I-13. 16 F-16913. Piper, C. V. Delphinium simplex and its immediate allies. Contr. — U. S. Nat. Herb. 1§: 201-203. 12 F 1913. a Piper, C. V. The identity of Heuchera cylindrica. Contr. U. S. Nat Herb. 16: 205, 206. 12 F 1913. Includes Heuchera chlorantha sp. nov. Piper, C. V. New or noteworthy eee of Pacific coast plants. Contr. U. S. Nat. Herb. 16: 207-210. 2F 1913 3 Seven new species are described in Alsine i fbionian (1), Oreobroma (1)s , Aster (3), and Arabis (1). A Piper, C. V. On the identity of Poa crocata Michx. Tomeys 13: 3 3 F 1913. Piper, C. V. Supplementary notes on’ American species of Festuca. : Contr. U. S. Nat. Herb. 16: 197-1 12 F 1913. ‘ Includes Festuca sororia sp. nov. ain, D., & Hutchinson, J. Notes on some species of Acalypha oe Bull. Misc. Inf. 1913: 1-28. f. 1-4. Ja 1913. ‘ INDEX TO AMERICAN. BOTANICAL LITERATURE 191 Prescott, A. The common polypody. Am. Bot. 19: 25, 26. F 1913. [Illust.] Quehl, L. Einiges iiber Echinocactus Wislizeni Engelm., Ects. Lecontei Engelm. und Ects. arizonicus. Kunze. Monats. Kakteenk. 23: Q-II. 15 Ja 1913. Robertson, T. B. Further explanatory remarks concerning the normal rate of growth of an individual and its Mone sic significance. Biol. Centralb. 33: 29-34. 20 Ja 1913. Rose, J. N., & Standley, P.C. The American species of Meibomia of the section Nephromeria. Contr. U. S. Nat. Herb. 16: 211-216. pl. §1. 12 F 1913 Includes Meibomia Painteri, M. metallica, and M. angustata, spp. nov. Safford, W.E. Raimondia, a new genus of Annonaceae from Colombia. Contr. U. S. Nat. Herb. 16: 217-219. pl. 52, 53. 12 F 1913. Saunders, C. F. In the home of the fan palm. Am. Bot. 19: I-5. F 1913. [IIllust.] Schaffner, J. H. The characteristic plants of a typical prairie. Ohio Nat. 13: 65-69. 20 F 1913. Includes an enumeration of species of plants to be found. Schaffner, J. H. The classification of plants, VIII. Ohio Nat. 13: 70-78. 20 F 1913. Includes a synopsis of the plant phyla and a classification of the fungi. Scott,W. Aform of Linaria vulgaris. Ottawa Nat.26:129. Ja 1913. Shafer, J. A. Further botanical explorations in Pinar del Rio, Cuba. Jour. N. Y. Bot. Gard. 14: 44-49. F 1913. : Sinnott, E. W. The morphology of the reproductive structures in the Podocarpineae. Ann. Bot. 27: 39-82. pl. 5-9 +f. 1-9. Ja 1913. Small, J. K. A yellow flax from Jamaica. West Indies. Torreya 13:63. Mr 1913. Cathartolinum jamaicense Small, sp. nov. Steele, E.S. Four new species of goldenrod from the eastern United States. Contr. U. S. Nat. Herb. 16: 221-224. F 1913. Sumstine, D. R. Studies in North American Hyphomycetes—Il. Mycologia 5: 45-61. pl. 82-84. 10 Mr 1913. Includes Oosporoidea and Toruloidea, gen. nov., and Oidium Murrilliae, Toru- loidea effusa, F. Unangstii, and Acrosporium Gossypit, spp. Nov. Visher, S. S. Additions to the flora of the Black Hills of South Da- kota—II. Muhlenbergia 8: 135-137- 31 Ja 1913. Wagner, E. Mitteilungen iiber Samlingsaufzucht. Monats. Kak- teenk. 23: 6-8. 15 Ja 1913. — 192 INDEX TO AMERICAN BOTANICAL LITERATURE Weingart, W. Cereus serratus Weing. spec. nov. Monats. Kakteenk. 22: 185-188. 15 D 1912. Wellington, R. Inheritance of the russet skin in the pear. Science II. 37: 156. 24 Ja 1913. Wherry, E. T. Silicified wood from the Triassic of Pennsylvania. Proc. Acad. Nat. Sci. Philadelphia 64: 366-379. pl. 3, 4. 25S 1912. Williams, C. G. The farm grasses of Ohio. Ann. Rep. Ohio Agr. Exp. Sta. 30: 151-174. 1911. — [IIlust.] Wooton, E. O., & Standley, P. C. Descriptions of new plants pre- liminary to a report upon the flora of New Mexico. Contr. U.S. Nat. Herb. 16: 109-196 + vii-xi. p!. 48-50. 12 F 1913. Yamanouchi, S. Hydrodictyon africanum, a new species. Bot. Gaz. 55: 74-79. f. 1-6. 15 Ja 1913. BuLL. TORREY CLUB VOLUME 40, PLATE 3 SLOSSON: NEW FERNS FROM TROPICAL AMERICA PLATE 4 VOLUME 40, TorRREY CLUB . BuLi Buty. ToRREY CLUB VOLUME 40, PLATE 5 coe LEVINE: CYTOLOGY OF HYMENOMYCETES BuLL. TORREY CLuB VOLUME 40, PLATE 6 LEVINE: CYTOLOGY OF HYMENOMYCETES BuLi. Torrey CLuB VOLUME 40, PLATE 7 LEVINE: CYTOLOGY OF HYMENOMYCETES VOLUME 40, PLATE 8 BULL. ToRREY CLuB LEVINE: CYTOLOGY OF HYMENOMYCETES Vol. 40. er A K No. 5 BULLETIN ‘TORREY BOTAN ICAL CLUB A taxonomic study of the Pteridophyta of the Hawaiian Islands—Ill WINIFRED J. ROBINSON (WITH PLATES 9-12) 19. POLYPODIUM L. Sp. Pl. 1082. 1753 A cosmopolitan genus, including ferns of every habitat. Root- stock creeping; leaves articulate; blades various, mostly simple or pinnate; veins free or occasionally anastomosing; sori orbicular, dorsal on the veins, non-indusiate. Type species: Polypodium vulgare L. Leaves simple to deeply pinnatifid, not bearing reddish, resinous glands beneath. Leaves simple Margin entire or very slightly undulate. Rootstock creeping, 5-8 mm. in diameter, the scales light brown; leaves 8-20 cm. X 7.5- 10 mm. Leaves stalked, — with numerous r ish brown ts 2-3 mm. long; margin entire; sori inframedial. P. Hooker. Leaves sessile, pubescent with scattered eis hairs less than 1 mm. long; margin lightly undulate; sori supramedial. eine erect or creeping, less than 5 mm. in diameter, the scales dark brown; leaves P. Knudsenii. 5-7 cm. X 2-5 mm. , Rootstock. erect, rather stout; leaves sub- sessile, linear-lanceolate, entire; sori oval, numerous., Rootstock creeping, wiry; leaves stalked, linear, eighty undulate; sori round, few, distant. P. pumilum. 'P. pseudogrammitis. [The Buttetin for April 1913 (40: 137-192. pl. 3-8) was issued May 9.) 194 RopiINsON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS . Margin crenate to serrate, at least in the lower part; rootstock erect; leaves 10-15 cm. X 2-4 mm. Margin crenate or dentate from base to apex; sori oval, alternate, ees amedial on serrations. P. Haaliliolanum, Margin sinuately pinnatifid below, subentire in in i apical third; sori a inframedial. P. Saffordii. Leaves pinnatifid. asa gn Aonig reduced to a wing along the midrib; er surface and margin marked by scattered, zener glandular hairs Leaves stisptic-lencediate- to elliptic-caudate, m.; sori nearly covering leaf tissue from costa to margin P. sarmentosum. ; -aperee linear to narrowly POS OM 15-25 a 1.52 cm.; sori ae dial. P. Adenophorus. ahlonec ¢ oblong, Sais varying fais Hneok to lanceolate with narrow sinuses between them, to oblong and obtuse with broad sinuses; pellucid lines between the P. pellucidum. vein Leaves soe cee to deeply tripinnatifid, bearing numerous, reddish, resinous glands beneath Leaves bipinnatifid Leaves linear, 5-15 cm. X 5-7.5 mm., pinnae divided into 2 or 3 segments. P. hymenoph: Leaves not linear; pinnae divided into 6 to 15 seg- © : ments. Leafstalk 0.5 mm. in diameter; blades oblong- ovate, 3.5-5 cm. X 1-2.cm.; pinnae compact. P. abietinum. k 1-2 mm. indiameter; blades more saangge Io cm. long. - Leaves subcoriaceous, blades ovate-lanceo- late; pinnae oblong, 20-30 cm. X 6-8 cm.; segments linear; sori narrower than the 3 segments. P. Hillebrandit. Leaves chartaceous, blades elliptic-lanceo- late to elliptic-caudate, 10-35 cm. X 2-5 cm., pinnae lanceolate; segments ob- lanceolate to spatulate; sori as broad as the segments. P. tamariscinum- Leaves tripinnatifid; pinnae 15-25 cm. X 2-3 cm.; seg- oe ments spatulate; sori as broad as the segments. P. Gaudichaudit- Potypopium Hooker! Brack. Fil. U. S. Expl. Exp. 4. 18; Polypodium setigerum Hook. & Arn. Bot. Beech. 103. 1832+ _ Blume. Grammitis setigera J. Sm. Hist. Fil. 181. 1875. TYPE LOCALITY: Oahu. DistTRIBUTION: Polynesia, Hawaiian Islands. ROBINSON: PTERIDOPHYTA OF THE HAWAUAN IsLANDs 195 ILLustRATION: Hook. & Arn. Bot. Beech. pl. 21. f.a. 1832. SPECIMENS EXAMINED: Hawaii, Mann & Brigham 482 N; Robinson 201 V; Maui, Robinson 336 V; 338 V;352 V; Oahu, Heller 2245 C, N; Lichtenthaler N; Hawaiian Islands, Baldwin 8r C, N; Baldwin B; Hillebrand B; ex Herb. Underwood C; Hillebrand N. This is closely allied to P. setosum (Blume) Presl, according to Copeland, Polypodiaceae of the Philippine Islands, Bur. Gov. Lab. Publ. 21: 119. 1905. PoLypopium KnupsEnm Hieron. Hedwigia 44?: 79. 1905 Polypodium samoense var. glabra Hilleb. Fl. Haw. Is. 554. 1888. Not P. glabrum Roxb. Polypodium samoense Christ in Engler, Bot. Jahrb. 23: 358. 1896, in part. Not Baker. TYPE LOcALITY: Kauai, Hawaiian Islands. DISTRIBUTION: On trees, Kauai, Hawaiian Islands. SPECIMENS EXAMINED: Kauai, Baldwin 17 B; Heller 2708 B; C, K, N; Knudsen 7 B; Robinson 405 V; 440 V; 453 V; 826 V; 836 V Hieronymus (Hedwigia 442: 79. 1905) distinguishes P. Knud- sentt from P. samoense by the broader, yellow rather than fer- ruginous, scales of the rhizome; the fewer, smaller hairs upon the leafstalk; the thicker texture and more pointed apex of the leaf; the less evident veins; the larger size of the sori and their greater proximity to the margin of the leaf. Polypodium pumilum sp. nov. Rootstock short, erect, less than I mm. in diameter, covered with dark brown, linear-lanceolate scales 1-2 mm. long and 0.3-0.5 mm. broad at the base, with 6-10 rows of cells in the broadest part; roots numerous; leaves 3-5, simple, nearly sessile, cespitose, linear-oblanceolate, obtuse or bluntly acute at apex, narrowed gradually at the base, 4-7 cm. long, 3-5 mm. wide, the margin subentire, the midrib green, prominent; veins forked once about 2 mm. from the midrib; sori oblong, 1.5-2 mm. long by I-I.5 mm. broad; sporangia flattened laterally, the annulus of 10-12 cells; spores yellow, tetrahedral, 0.040 mm. in diameter, very minutely tuberculate. [Fic. 1.] ) Type collected on Oahu, 1000 m. elevation, 1890, by D. D. Baldwin, distributed by D. C. Eaton as P. sessilifolium (herb. N. Y. Bot. Gard.). : { 196 RoBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS Known only from the type locality. The present species differs from P. Knudsenii Hieron. in its smaller size; in the size of the scales, which in the latter are 3-4 _ Fic.1. Polypodium pumilum, natural size. mm. long and 0.5-0.8 mm. wide, with 10-20 rows of cells in broadest part; and in the smooth margin of the leaf in contra with the hairy margin of P. Knudsenii. POLYPODIUM PSEUDOGRAMMITIS Gaud. Voy. Freyc. Bot. 34, , | 1828 Grammitis tenella Kaulf. Enum. 84. 1824. Not Forster. Polypodium Kaulfussii Presl, Tent. Pterid. 178. 1836. TYPE LOCALITY: Oahu. DISTRIBUTION: On trees, common; Hawaiian Islands. ItLustRaTIons: Hook. & Arn. Bot. Beech. pl. 21. 18 Kunze, Anal. Pterid. pl. 9. {25 -4897. SPECIMENS EXAMINED: Hawaii, Hillebrand B, C; Robinson V; 260 V; 275 V; Oahu, Heller 2215 C; Macrae B; Robinson 2. 9 V; 45 V; 119 V; Kauai, Robinson 446 V; Hawaiian Is Baldwin 32 B; 83 B, C; Gaudichaud B: Wilkes Expedition B, | ROBINSON: PTERIDOPHYTA OF THE HawattANn IsLANDs 197 PoLypopIuM HAALILIOLANUM Brack. Fil. U. S. Expl. Exp, 5. 1854 TYPE LocaLity: Hawaiian Islands. DISTRIBUTION: On tree trunks above 600 m. elevation, Hawai- ian Islands; rare. ILLustTRATION: Brack. Fil. U.S. Expl. Exp. pl. 7. f.4. 1854. SPECIMENS EXAMINED: Hawaii, Robinson 601 V; Oahu, Hille- brand B; Mann & Brigham 178 N; Wilkes Expedition N; Kauai, Lichtenthaler N; Hawaiian Islands, Baldwin 86 B; C; Hillebrand C. Hieronymus (Hedwigia 442: 86. 1905) reduces Polypodium Haaliliolanum to Polypodium subpinnatifidum Blume, but the type of the latter at Berlin has subpinnatifid lobes and acute sinuses, while the leaves of the Hawaiian plant are sinuato-crenate throughout. The sporangia of Polypodium Haaliliolanum are smooth, while those of Polypodium subpinnatifidum are setose. PoLypopium Sarrorpir Maxon, Am. Fern Jour. 2: 19. 1912 Polypodium minimum Brack. Fil. U. S. Expl. Exp. 5. pl. 1. f. 3. 1854. Not Aublet. Polypodium serrulatum var. lata Luerssen, in Wawra, Flora 58: 422. 1875. Polypodium serrulatum Hilleb. Fl. Haw. Is. 553. 1888. Not Mett. TYPE LOCALITY: Oahu. DIsTRIBUTION: Common on trees above 600 m. elevation, Hawaiian Islands. ILLustRations: Brack. Fil. U. S. Expl. Exp. pl. 1. f. 3. 1854; Maxon, Am. Fern Jour. 2: 19. 1912. SPECIMENS EXAMINED: Hawaii, Lichtenthaler N; Robinson 600 V; Oahu, Heller 2905 (type) N, (cotype) C; Mann & Brigham 550 N; Robinson 521 V; Wilkes Expedition N; Kauai, Hillebrand 123 B; Hawaiian Islands, Baldwin 85 B, C; Bishop 81 B; Johnson B; ex Herb. Mt. Holyoke College B; Wilkes Expedition C. PoLYPopIUM SARMENTOSUM Brack. Fil. U.S. Expl. Exp. 8. 1854 Tyrer LocaLity: Hawaiian Islands. DISTRIBUTION: Common on rocks and decasred wood, Hawai- ian Islands. 198 RoBINsoN: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS ILLUSTRATION: Brack. Fil. U.S. Expl. Exp. pl. 2. f. 3. 1854. SPECIMENS EXAMINED: Hawaii, Robinson 203 V; Maui, Batley — C; Robinson 356 V; Oahu, Forbes BM; Heller 2353 C, N; Lichten- 3 thaler N; Mann & Brigham 200 N; Robinson 1 V; 5 V; 46 V; _ 67 V; 117 V; Safford 889 N; Kauai, Robinson 856 V; 418 V; — Hawaiian Islands, Baldwin 82 C, N; Wilkes Expedition C; ex — Herb. John Donnell Smith N. Potypopium ApDENoPHORUS Hook. & Arn. Bot. Beech. 104. 1832 | Adenophorus pinnatifidus Gaud. Voy. Freyc. Bot. 365. 1829. Not Gilib. : TYPE LOCALITY: Hawaiian idence DISTRIBUTION: On tree trunks in the forests above 600 m. elevation. ILLUSTRATION: Hook. & Arn. Bot. Beech. pl. 22. 1832. y SPECIMENS EXAMINED: Oahu, Lichtenthaler N; Mann & Brig- — ham 283 N; Kauai, Knudsen B; Robinson 851 V; Hawaiian — Islands, Baldwin 88 B, C; Wilkes Expedition C, N; ex Herb. q John Donnell Smith N. a : POLYPODIUM PELLUCIDUM Kaulf. Enum. tor. 1824 Polypodium myriocarpum Hook. Ic. Pl. pl. 84. 1837. Polypodium hawatiense Underw. in Heller, Minn. Bot. Stud. s 784. 1897. : preven: Helleri Underw. in Heller, Minn. Bot. Stud. 1: 785-4 1897. Polypodium vulgare Christ, Farnkr. Erde 83. 1909. TYPE LOCALITY: Oahu. 3 DisTRIBUTION: On ground and on tree trunks, at 300-2,000 — m. elevation; Hawaiian Islands. ILLUSTRATIONS: Hook. Ic. Pl. pl. 84. 1837; pl. 944, 94. 1854. : . SPECIMENS EXAMINED: Hawaii, Dale C; Robinson 144 V; 25 267 V; Wilkes Expedition C; Maui, Bishop 83 B; 84 B; Finsch 1 60 B; Robinson 316 V; 324 V; 333 V; Molokai, Hillebrand Oahu, Chamisso B; Forbes 1017 BM; Heller 2075 C; Robinson Kauai, Forbes 268 BM; Heller 2602 C; 2634 C; Hillebrand 54 Knudsen 55 B; 56 B; Robinson 403 V; 482 V; Hawaiian Is ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLAnps 199 Baldwin 89a B; 89c B; 89 C; Bailey C; Douglas 69 B, C; Gaudi- chaud B; Lydgate B; Wilkes Expedition B, C. Polypodium pellucidum is extremely polymorphous, as is shown by the several nominal species that have been segregated at dif- ferent times. Kaulfuss’s type in the Berlin herbarium is very coriaceous and has the intermediate veinlike striations mentioned by this author. The pinnae are rather acute, 4-5 cm. long, with crenate-undulate margins. Divergences from this are seen in: (1) P. Helleri Underw. (type Heller 2602) with longer thinner pinnae; a single incomplete leaf has greatly elongated pinnae; (2) P. hawatiense Underw. with broader (1.5 cm.), shorter (3-4.5 cm.), rounded pinnae, Hillebrand’s var. opacum from Molokai; (3) P. myriocarpum (Hook. Ic. Pl. 1: pl. 84) apparently a mon- strous form with great variability in the pinnae of the same leaf; (4) a form still more common in collections, which has close or spaced pinnae ranging from rounded to pointed on the same leaf. Near the extinct craters, about a mile from the Volcano House, Hawaii, the leaves are folded ventrally upon the midrib. A field study of the range of variations of the species would form an interesting local problem. POLYPODIUM HYMENOPHYLLOIDES Kaulf. Enum. Fil. 118. 1824 Amphoradenium minutum Desv. Prod. 336. 1827. Adenophorus hymenophylloides Hook. & Grev. Ic. Fil. pl. 176. 1829. TYPE LocaLity: Hawaiian Islands. DIsTRIBUTION: Rare, on trees at elevations of 1,000-1,600 m., Hawaiian Islands. ILLustrations: Gaud. Voy. Freyc. Bot. i. 8. Ic. Fil. pl. 176. 1829. SPECIMENS EXAMINED: Hawaii, Robinson 276 V; forest above Cape Lua Pele, Wilkes Expedition N; Wilkes Expedition 19 B, Cc; Maui, Robinson 347 V; Wilkes Expedition B, C, N; Oahu, Beechey C; Diell C; Robinson 87 V; 97 V; Kauai, Heller 2213 C, N; Knudsen 154 B; Robinson 462 V; Hawaiian Islands, Baldwin go B, C, N; Gaudichaud B, C; Hillebrand B; Mann & Brigham 276 N. Hook. & Grev. 200 RoBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS POLYPODIUM ABIETINUM D.C. Eaton, in Mann, Proc. Am. Acad. 7: 219. 1867 Polypodium tamariscinum var. abietinum Hilleb. Fl. Haw. Is. 557. 1888 TyPE LocALity: Hawaiian Islands. DISTRIBUTION: On trees, rare; Hawaiian Islands. SPECIMENS EXAMINED: Hawaii, Wilkes Expedition N; Oahu, Hillebrand B; Remy C; Robinson 32 V; 48 V; Wilkes Expedition N; — Kauai, Heller 2732 C; N; Hillebrand B; Lichtenthaler N; Hawaiian : Islands, Baldwin 94 B; C; Hillebrand B. } Po_yropium HILLEBRANDII Hook. Sp. Fil. 4: 228. 1862 TYPE LOCALITY: Hawaiian Islands. : DisTRIBUTION: At 400-700 m. elevation, rare; Hawaiian Islands. ILLUSTRATION: Hook. Sp. Fil. 4: pl. 279A. 1862. SPECIMENS EXAMINED: Hawaii, Robinson 287 V; Oahu, Forbes. i BM; Robinson 23 V; 123 V; Hawaiian Islands, Baldwin 92 C, Ni Baldwin 93 C; Hillebrand B, C; Safford 891 N; ex Herb. John Donnell Smith N; Wilkes Expedition C. a POLYPODIUM TAMARISCINUM Kaulf. Enum. Fil. 117. 1824 a Adenophorus bipinnatus Gaud. Ann. Sci. Nat. 3: 508. 1824. TYPE LOCALITY: Oahu. DISTRIBUTION: Hawaiian Islands. ILLUSTRATION: Hook. & Grev. pl. 175. 1829. SPECIMENS EXAMINED: Hawaii, Robinson 208 V; 248 V; Oah Copeland N; Heller 2214 N; Lichtenthaler N; Mann & Bright 198 N; Robinson 83 V; 91 V; 92 V; 118 V; 122 V; Safford 892 N Kauai, Robinson 463 V; 495 V; 832 V; Hawaiian Islands, Baldwit gt N; ex Herb. John Donnell Smith N. j POLYPODIUM TRIPINNATIFIDUM (Gaud.) Presl, Tent. Pterid. 1 1836 Ss Adenophorus tripinnatifida Gaud. Ann. Sci. Nat. 3: 508- 1 Voy. Freyc. Bot. pl. 8. 1826. 2 Amphoradenium Gaudichaudii Desv. Prod. 336. 1827. ° TYPE LOCALITY: Hawaiian Islands. DistriBuTION: Above 600 m. elevation, Hawaiian Islands. ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDs 201 ILLUSTRATION: Gaud. Voy. Freyc. Bot. pl. 8. SPECIMENS EXAMINED: Hawaii, Wilkes Expedition N; Maui, Lichtenthaler N; Molokai, Baldwin 93a B; 93b B; Kauai, Remy 16 C; Hawaiian Islands, Hillebrand 122 B; Wilkes Expedition 20 B, C. 20. PHYMATODES Presl, Tent. Pterid. 195. 1836 Tropical plants, mostly epiphytic; rootstock creeping; leaves articulate, distant, simple; veins obscure or prominent, anastomos- ing, with free included veinlets; sorus borne upon anastomosing veins or upon the clavate apex of an included vein, non-indusiate. Type species: Polypodium zosteraeforme Wall. Leaves linear-lanceolate; veins obscure; sori round, nearly covering wn. leaf from costa to margin, light bro Leaves palmately lobed; veins prominent; sori small, dark. P. elongata. P. Spectrum. PHYMATODES ELONGATA Presl, Tent. Pterid. 196. 1836 Polypodium lineare Thunb. FI. Jap. 335. 1784. Not Burm. Pleopeltis nuda Hook. Fil. Exot. 63. 1823. Pleopeltis linearis Kaulf, Enum. 246. 1824. Pleopeltis Thunbergiana Kaulf. Wes. d. Farrenkr. 113. 1827. Polypodium leiopteris Kunze, Klotzsch, Linnaea 23: 319. 1850. Drynaria nuda Fée, Gen. 1: 270. 1850-52. Drynaria elongata Brack. Fil. U. S. Expl. Exp. 42. 1854. Niphobolus linearis Keyserl. Polyp. Cyath. Herb. Bung. 39. 1873. TYPE LocALity: Kosido, Japan. DIsTRIBUTION: On trees and rocks; pone countries. SPECIMENS: EXAMINED: Hawaii, Lichtenthaler N; Maui, Wilkes Expedition N; O. Finsch to C; Kauai, Heller 2533 C; 3533 N; Oahu, Wilkes Expedition C; Diell C; Heller 2005 C, N; 2076 C, N; 2031 C, N; Mann & Brigham 177 N; Robinson 84 V; Safford 893 N; 1887 N; Hawaiian Islands, Baldwin 82 C, N; R. H. Hitchcock C; ex Herb. John Donnell Smith N. PHyMATODES SPECTRUM (Kaulf.) Presl, Tent. Pterid. 197. 1836 Polypodium Spectrum Kaulf. Enum. 94. 1824. Polypodium Thouinianum Gaud. Voy. Freyc. Bot. 348. 1828. 202 ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS Drynaria spectrum J. Sm. Jour. Bot. Hook. 4: 61. 1842. Pleopeltis spectrum Moore, Ind. Fil. 348. 1862. Colysis Spectra J. Sm. Ferns Brit. & For. 98. 1866. TYPE LOCALITY: Oahu. DIsTRIBUTION: On rocks and tree trunks in dense shade of woods, Hawaiian Islands, Sumatra. ILLUSTRATIONS: Gaud. Voy. Freyc. Bot. pl. 5. f. i Bee Nat. Pi.2*;-317. 7. 1644, B«. 1800. SPECIMENS EXAMINED: Hawaii, Robinson 200 V; Maui, Wilkes Expedition C; Oahu, Heller 2118 C; Robinson V; Kauai, Heller 2438 C; Hawaiian Islands, Baldwin 84 C; Miss Sessions C; ex Herb. Kew. C. The leaves of Phymatodes Spectrum vary so much in size and cutting of the lobes as to suggest different species for those found growing upon a single rootstock. 21. PHLEBODIUM (R. Br.) J. Sm. Jour. Bot. 4: 58. 1842 Tropical or subtropical plants, usually epiphytic. Rootstock creeping, I-2 cm. in diameter, paleaceous; leaves articulate; blades pinnatifid or pinnate; primary veins pinnately arranged, the secondary veins anastomosing; sori usually terminal on two included veinlets, non-indusiate. Type species: Polypodium aureum L. PHLEBODIUM AUREUM (L.) J. Sm. Jour. Bot. Hook. 4: 58. 1842 TYPE LOCALITY: West Indies. DisTRIBUTION: On ground and on trees, West Indies, Florida, Mexico, Brazil, Hawaiian Islands. | ILLUSTRATIONS: Plumier, Traité Foug. pl. 76. 1705. PLATE 9- SPECIMENS EXAMINED: Wahiawa Mts., Kauai, Forbes 308 BM. Forbes 308 is the first collection of this species that has been noted for the Hawaiian Islands, and it may be that it is an escape — from cultivation, though the remoteness of the locality where 1t _ was collected makes its origin from a cultivated plant improbable. blue.”’ 22. POLYSTICHUM Roth, Rém. Mag. 2!: 106. 1799 Mostly coarse and rigid terrestrial plants of cosmopolitam distribution; rootstock short, erect; leaves cespitose, not arti : rie eee ee i eee fae. eee Mr. Forbes notes: ‘All the fronds were remarkably glaucous . — ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANps 203 late; blades pinnate to quadripinnate; veins free: sori roundish, dorsal upon the veins; indusium centrally peltate, orbicular. Type species: Polypodium Lonchitis L. Leaves tripinnate, ovate-lanceolate, smooth or s slightly tomentose. P. carvifolium. Leaves bipinnate; stipe and rachis densely covered with linear- , lanceolate scales. Pinnae 1.5-3 cm. apart, sharply aristate. P. haleakalense. Pinnae crowded, margins entire or nearly so. P. Hillebrandii. POLYSTICHUM CARVIFOLIUM (Kunze) C. Chr. Ind. Fil. 70. 1905 Polystichum contifolium J. Sm. Jour. Bot. Hook. 3: 413. 1841. Not Pres ‘ Aspidium curvifolium Kunze, Bot. Zeit. 6: 283. 1848. Aspidium aristatum Sw. var. trots: Wall.; Hook. Sp. Fil. 4: 27. 5868; Dryopteris coniifolia Underw. in Heller, Minn. Bot. Stud. 1: 778. 1897. TYPE LOCALITY: Philippine Islands. DIsTRIBUTION: High plateaus, Soc Jape, Polynesia, Ha- waiian Islands, Australia. SPECIMENS EXAMINED: Heller 2817 C; Baldwin 62 C. The habit, coriaceous texture, and aristate pinnae of this plant place it in Polystichum rather than in Dryopteris. The indusia are sometimes reniform-orbicular, sometimes peltate, on the same leaf. It is very different from the Japanese Polystichum aristatum (Forster) Presl, in rootstock, texture, and the division of the leaf. POLYSTICHUM HALEAKALENSE Brack. Fil. U. S. Expl. Exp. 204. 1854 Aspidium haleakalense Mann, Proc. Am. Acad. 7: 216. 1867. Aspidium aculeatum var. Braunn Hilleb. Fl. Haw. Is. 568. 1888. Not Swartz. Tyrer LocaLity: Region of Sophora, Mauna Kea, Hawaii. DISTRIBUTION: In forests at altitudes of 1,800-2,700 m., Hawaiian Islands. ILLUSTRATION: Brack. Fil. U. 5. Expl. Exp. pl. 28. 1854. - SPECIMENS EXAMINED: Hawaii, Wilkes Expedition N; Maui, Lichtenthaler N; Finsch 45 B; Lydgate B; Lydgate & Baldwin B; Mann & Brigham 481 N; Mann B; ex Herb. Mt. Holyoke College C; Oahu, Macrae B; Hawaiian Islands, Baldwin 61 C, N; ex Herb. John Donnell Smith N. | 204 RoBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS PoLystTicHuM HILLEBRANDII Carruth. in Seem. Fl. Vit. 358. 1873 ; Aspidium Hillebrandi Hilleb. Fl. Haw. Is. 568. 1888. TYPE LOCALITY: Hawaiian Islands. DISTRIBUTION: In forests at altitudes of 1,500-1,800 m., Hawaiian Islands. SPECIMENS EXAMINED: Maui, Bailey C; Baldwin B, C, V! Lydgate B; Hawaiian Islands, Baldwin 60 B, C. 23. CYRTOMIUM Presl, Tent. Pterid. 86. 1836 Rootstock short; leaves clustered, pinnate, somewhat coriace- ous; veins anastomosing; sori globose, dorsal upon free included ‘ veinlets or upon connecting veins; indusium peltate. Type species: Cyrtomium caryotideum (Wall.) Presl. Leaves less than 15 cm. long, with a single row of areolae on either side of the midvein. C. Boydiae. Leaves more than 15 cm. long, with several areolae between midrib 3 and margin. C. caryotideum. Cyrtomium Boydiae (D. C. Eaton) comb. nov. Aspidium Boydiae D. C. Eaton, Bull. Torrey Club 6: 361. 1879. — Aspidium cyatheoides var. depauperatum Hilleb. FI. Haw. Is. 572. — 1888. 4 Dryopteris cyatheoides C. Chr. Ind. Fil. 66. 1906. 7 TYPE LOCALITY: Oahu. e, DISTRIBUTION: On rocky ledges, along streams, Hawaiian 3 Islands. * ILLUSTRATION: PLATE Io. z SPECIMENS EXAMINED: Hawaii, Hillebrand B; Maui, Baldwin . C, V; Oahu, Forbes BM; Rock V. . ; : CyRTOMIUM CARYOTIDEUM (Wall.) Presl, Tent. Pterid. 86. 1836 4 Aspidium caryotideum Wall. Cat. no. 376. 1828. Dryopteris caryotidea Underw. in Heller, Minn. Bot. Stud. 1: 77% 1897. 4 TYPE LocaLity: Nepal. - DiIsTRIBUTION: In forests, India, pes Hawaiian Islands. ILLusTRATIONS: Hook. & Grey. Ic. Fil. pl. 69. 1826; Presl, Tent. Pterid. pl. 2. f. 12. 1836; Brack. Fil. U. S. Expl. Exp- # 16. 1854; Hook. Garden Ferns pl. 13. 1862. ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS 205. SPECIMENS EXAMINED: Hawaii, Lichtenthaler N; Maui, Mann & Brigham 487 N; Wilkes Expedition N; Kauai, Heller 2544 C, N; Hillebrand B; Lydgate B; Lanai, Hillebrand B; Oahu, Forbes 431 BM; Hawaiian Islands, Baldwin 59 B, C; Remy B; Miss Sessions C. Christensen, following Diels (E. & P. Nat. Pfl. 14: 194. 1899), makes Cyrtomium caryotideum a synonym of Polystichum falcatum (L.f.) Diels. However, as Polystichum has free veins and this group has anastomosing veins, they are separated on that character rather than combined on the similarity of their indusia. The type of Polystichum falcatum is from Japan. Thunberg, Fl. Jap. 336. $l. 306. 1784, describes and figures it, but his figure represents a plant with cordate-falcate leaves not auricled, with scales at the base of the leafstalk. The Hawaiian plant resembles Wallich’s specimen at Kew, a tracing from which is in the herbarium of the New York Botanical Garden. 24. TECTARIA Cav. Anal, Hist. Nat. 1: 115. 1799 Rootstock creeping or decumbent; leaves clustered, not articulate, various in form, chartaceous; veins reticulate with free included veinlets; leafstalk purplish brown, smooth above, scaly at the base; sori circular; indusium orbicular, attached at the center. Type species: Polypodium trifoliatum 1 Ss Tectaria cicutaria (L.) comb. nov. Polypodium cicutarium L. Syst. Nat. ed. 10. 2: 1326. 1759. Aspidium cicutarium Sw. Jour. Bot. Schrad. 18002: 36. 1801. Aspidium apifolium Schkuhr, Krypt. Gew. 1: 198. 1809. Nephrodium apifolium Hook. & Arn. Bot. Beech. 105. 1832, Sagenia apiifolia (Schkuhr) J. Sm. Jour. Bot. Hook. 4: 184. 1841. TYPE LOCALITY: Jamaica, B. W. I. (?). DisTRIBUTION: In the tropics, on damp rocks, or in depths of shady forests. ILLUSTRATIONS! Pluk. Almag. 156. pl. 2096. f. 2. 1692; Schkuhr, Krypt. Gew. pl. 56b. 1809; Hook. & Grev. Ic. Fil. pl. 202. 1831. SPECIMENS EXAMINED: Hawaii, Wilkes Expedition N; Maui, Bailey C; Baldwin N; Robinson 314 V; 328 V; 339 V; Oahu, Beechey (ex Herb. Mettenins)’ C; Mann & Brigham 189 N; Robin- 206 ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS son 13 V; 18 V; 135 V; Safford 914 N; 915 N; 916 N; Wilkes Expe- dition N; Kauai, Heller 2842 C, N; Hillebrand 60 B; Knudsen 68 B; Hawaiian Islands, Baldwin 68 B, C, N; Hillebrand (ex herb. Berlin) C; Lindley C; Miss Sessions C; ex Herb. Underw. C; ex Herb. John Donnell Smith N; Wilkes Expedition C, N. 25. NEOTTOPTERIS J. Sm. Jour. Bot. Hook. 3: 409. 1841 A genus of epiphytic ferns, with fleshy, orchid-like roots, found in the forests of tropical Asia, Polynesia, Australia, and Africa. Rootstock erect, short; leaves large, simple, cespitose, sessile or nearly so, spreading, glabrous, entire, arranged spirally with short internodes; veins slender, parallel, close (less than I mm. apart), oblique to the midrib, connected by a transverse intramarginal vein at their apices; sori linear, extending from near the midrib 14 to 2 the length of the veins in the distal half or two thirds of the leaf. Type species: Neottopteris Nidus (L.) J. Sm. NEoTTopTERis Nipus (L.) J. Sm.; Hook. & Bauer, Gen. Ferns pl, 113.Be 1842 4 Asplenium Nidus L. Sp. Pl.-1079. 1753. Thamnopteris Nidus Presl, Epim. 68. 1849. TYPE LOCALITY: Java. DIsTRIBUTION: On trees and on humus soil in forests, usually below 700 m., tropical Asia, Australia, Madagascar, and Polynesia. ILLUSTRATIONS: Hook. & Bauer, Gen. Ferns pl. 113B. 1842; Breyne, Exot. Pl. Cent. pl. 99. 1678. SPECIMENS EXAMINED: Hawaii, Wilkes Expedition N; Maui, Bailey C; Oahu, Didrichsen 3652 C; Gaudichaud B; Hillebrand B; Meyen B; Robinson 17 V; 103 V; Kauai, Heller 205 C; Hawaiian Islands, Baldwin 30 C; Mann & ie 137 N; Safford 31 N; Miss Sessions C. 26. ASPLENIUM L. Sp. Pl. 1078. 1753 A genus of world-wide distribution, especially well represented in the temperate zones, though including many tropical forms. Rootstock erect or creeping; leaves non-articulate, usually cespitose; blades simple to quadripinnate, coriaceous to mem-_ ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDs 207 branaceous; veins free in Hawaiian species; sori linear ‘upon the veins, usually oblique to the midrib or leading vein of the pinna, occasionally parallel with it; indusia attached to veins, opening towards the midrib; sori occasionally athyrioid or diplazioid. Leafblades simply pinnate. ‘innae cau sy to rhomboidal or oblong, subentire. Rootstock creeping; leaves distant. A. unilaterale. haere short, erect; leaves cespitose. Leafstalk dull green, flexuose Pinnae rhomboidal to flabellate, occasion- ally distant, gradually reduced below. A. rhomboideum. Pinnae lente approximate, abruptly reduced bel A. lunulaium. Leafstalk dark Rise or black, ri Bl chartaceous; sori on veins pevatiel to the lower margin, opening upwards. A. monanthes. Blade subcoriaceous; sori on secondary veins, indusium opening toward pri- mary vein. Blade 2.5—10 cm. long; pinnae ovate to suborbicular. A. Trichomanes. Blade 15 cm. long; pinnae oblong. A. pavonicum. Pinnae lanceolate, margin serrate, incised, or lobed. alks green Pinnae auricled at base. A. pseudofalcatum. Pinnae not auricled at base. eafblade coriaceous; basal scales of leaf- stalk ovat Pinnae obtuse at apex; broadly ovate; sorus conterminous with vein; veins obscure. A. Kaulfussii. Pinnae caudate; sorus shorter than vein; veins distinct. A. kauaiense. Leafblade chartaceous; basal scales lance- olate or linear. Leafblade 30-35 cm. long, oblong- ovate; basal scales of leafstalk lan- ceolate. A. enatum. Leafblade 20-25 cm. long, oblong-lan- ceolate; basal leafstalk lin A. Hillebrandii. Leafstalks brown or 5 stee-blue. Pinnae serrate or in < e caudate. imary veins parallel to the midrib. A. contiguum, Primary veins divergent from the mid- rib. A. caudatum. 208 RospINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS Pinnae acuminate. Pinnae cut into rhombic lobes Leafstalk and midrib paleaceous. Leafstalk and midrib smooth. Leafblades more than once pinnate. Bl bhi Pimervectaie to bipinnate, upper basal pinnule auriculate. Binds aad to eT ARI basal pinnules various. Blades bipinnate to tripinnatifid. es chartaceous. Blades under 35 cm. lon Blades oblong, ultimate divisions flabel- late. Blades 3.75-7 cm. X12-18 mm. Blades 20-30 cm. X18—-45 mm. Blades lanceolate, ultimate divisions ovate-lanceolate. Blades over 35 c r ed eee bas patent; pin- kes obliquely ee ee Pinnae lanceolate, approximate; pin- s rhomboidal, basal umaeAlS oc- casa auriculate. Blades coriaceou lades ES ultimate divisions obovate t Pinnae obliquely rhomboidal, approxi- t mate. Pinnae ovate oblong, patent. Blades lanceolate. Ultimate divisions wai or trun- cate, pinnae approxi Ultimate divisions ieee patent. Blades tripinnate to quadripinnate. Blades broadly oblong. Blades deltoid or deltoid lanceolate. lades broadly deltoid, 10-20 cm. long. Blades deltoid lanceolate, 30 cm. or more long. Blades membranaceous; ultimate divi- sions spinulose. Blades thick, chartaceous. Rachis winged; ultimate divisions spatulate, less than 2 mm. broad. . Rachis not winged; osihiane divi- sions 4 mm, or more broad. A, nitidulum. A. horridum. A. glabratum., A. lobulatum. A. varians. A. Macraei. A. Lydgatet. A. Goldmannii. A. acuminatum. A. rhipidoneuron. A. insiticium. > . cuneatum. A. parallelum. A. patens. A. Adiantum-nigrum. A. vexans. : A. schizophylium. A. sphenotomum. | ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDs 209 ASPLENIUM UNILATERALE Lam. Encyc. 2: 305. 1786 Asplenium resectum J. E. Sm. Pl. Ic. Ined. 3. pl. 72. 1790-91. Asplenium amoenum Presl, Tent. Pterid. 107. 1836. Asplenium emargino-dentatum Zenker; Kunze, Linnaea 24: 263. 1851. TYPE LOCALITY: Mauritius. DIsTRIBUTION: In wet, shady localities, Mauritius, Japan, China, Polynesia, Hawaiian Islands. SPECIMENS EXAMINED: Hawaii, Robinson 622 V; Wilkes Expe- dition N; Maui, Bailey C; Lichtenthaler N; Heller 2844 C, N; Robinson 359 V; 366 V; 380 V; Oahu, Arnott N; Beechey C; Mann & Brigham 168 N; Robinson 72 V; 120 V; 122 V; 184 V; Safford 918 N; Kaala Mts., Wilkes Expedition N; Kauai, Forbes 439 BM; Hawaiian Is., Baldwin 37 C, N; Miss Sessions C; Wilkes Expedition €, N; ex Herb. John Donnell Smith N. ASPLENIUM RHOMBOIDEUM Brack. Fil. U.S. Expl. Exp. 156. 1854 Asplenium stoloniferum Presl, Rel. Haenk.1:44. 1825. Not Bory. Asplenium fragile Presl, Hilleb. Fl. Haw. Is. 589. 1888. TYPE LOCALITY: Hawaiian Islands. DIsTRIBUTION: At elevations of 400-700 m., Hawaii and Maui. ILLUSTRATIONS: Brack. Fil. U. S. Expl. Exp. pl. 27. f. 2. 1854; Presl, Rel. Haenk. 1: pl. 6.f.4. 1825. SPECIMENS EXAMINED: Hawaii, Mauna Kea, Wilkes Expedition N; Forest near Cape Lua Pele, Wilkes Expedition N; Maui, Hille- brand N; Hawaiian Is., Baldwin B, C, N; ex Herb. John Donnell Smith N. A. rhomboideum is closely related to the South American A. fragile and to A. viride of N. Europe and Canada. It differs from both in the openness of the fronds and flabellate form of the pinnae as well as in size. Kunze (Klotzsch, Linnaea 13: 140. 1839) in describing A. fragile collected by Ehrenberg in Peru gives the length of the largest fronds as 2 in. He describes the habit as the same as that of A. viride and makes A. castaneum Schlect. (Linnaea §: 611. 1830) and A. projectum Kunze (Linnaea 9: 68. 1834) synonyms. Diels, in E. & P. Nat. Pfl. 14: 235. 1899, gives A. fragile as a doubtful species, while he gives A. castaneum and A. 210 RoBpiInson: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS projectum true specific rank. Christensen (Ind. Fil.) gives the same treatment. ASPLENIUM LUNULATUM Sw. Jour. Bot. Schrad. 1800: 202. 1801 Asplenium erectum Bory, Willd. Sp. Pl. 5: 328. 1810. TYPE LOCALITY: South Africa. DISTRIBUTION: Tropical countries. SPECIMENS EXAMINED: Hawaii, Hillebrand B; Lichtenthaler Nj Maui, Bailey C; Robinson 720 V; 721 V; Kauai, Forbes 377 BM; _ Heller 2845 C; Knudsen 147 B; Hawaiian Is., Baldwin 35 N; Biddlesome N. ASPLENIUM MONANTHES L. Mant. 130. 1767 E Asplenium monanthemum Murr. L. Syst. Nat. ed. 13. 785. 1774+ = Asplenium Menziesii Hook. & Grev. Ic. Fil. pl. r00. 1829. be Asplenium leptophyllum Fée, Mém. Foug. 7. 50. pl. 14. f. 2. 1857+ TyPE LocaLity: Cape of Good Hope. : DISTRIBUTION: At elevations of 900-1,800 m., South America, — Northern Africa, the Azores, Madeira Islands, Hawaiian Islands. InLustRATIONS: J. E. Sm. Pl. Ic. Ined. pl. 73. 1790-91; Brack. Fil. U. S. Expl. Exp. pl. 20. f. 2. 1854; Fée, Mém. Foug. 7. pl. 14.f.2. 1857; Hook. & Grev. Ic. Fil. pl. 100. 1829. s SPECIMENS EXAMINED: Maui, Bailey C; Bishop 77 B; Hille- — brand B; Lichtenthaler N; Kauai, Knudsen 132 B; Oahu, Diell Ci Hawaiian Is., Baldwin 32 B, C; 1838-42; Wilkes Expedition N. — In the Berlin Herbarium there are two types of A. monanthesy | MS ROT en Sf 3 one lax, spreading, with lobed pinnae, proliferous above, the other 2 upright, taller, proliferous from lowest axils of pinnae, yet there is not enough difference to separate them as species. cM ASPLENIUM TRICHOMANES L, Sp. Pl. 1080. 1753 Asplenium densum Brack. Fil. U.S. Expl. Exp. 151. 1854. TYPE LOCALITY: Europe. DisTRIBUTION: On rocks in temperate regions; on mountait sides j in the tropics. : InLustrations: Brack. Fil. U. S. Expl. Exp. pl. 20. f. 3, i 1854. ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLaANps 211 SPECIMENS EXAMINED: Maui, Bailey C; Hillebrand B; Finsch 27 B; Oahu, Mt. Kaa, Macrae B; Hawaiian Is., Baldwin Bo Wilkes Expedition B, C; Lindley C. The difference between A. densum Brack. and the European A. Trichomanes L. consists in the fewer sori (never more than one row in the Hawaiian plant) and the more prominent wings upon the rachis, which suggests the question whether slight differences between plants widely elope geographically should be regarded as specific. ASPLENIUM PAVONICUM Brack. Fil. U. S. Expl. Exp. 150. 1854 Asplenium normale Hilleb. Fl. Haw. Is. 588. 1888. Not Don. TYPE LOCALITY: Saw Mill, Hawaii. DISTRIBUTION: In moist, shady forests, Hawaiian Islands; rare. ILLUSTRATION: Brack. U. S. Expl. Exp. pl. 20. f. 1, ra, 1b. SPECIMENS EXAMINED: Hawaii, Wilkes 50465 N (type); Kil- auea, Lichtenthaler N; Maui, Bailey C; Lichtenthaler N; Kauai, Forbes 160 BM; Heller 2771 C, N; Knudsen 10 B; Robinson 823 V; Oahu, Heller 2218 C; Safford 976 N; 977 N; Hawaiian Is., Baldwin 33 C; Lydgate B; Macrae B; Van Ingen C; ex Herb. John Donnell Smith N; Wilkes Expedition B. A. pavonicum is very nearly related to A. normale Don from Ceylon, but differs in the form of the pinnae and in size. . ASPLENIUM PSEUDO-FALCATUM Hilleb. Fl. Haw. Is. 597. 1888 _ Type Loca.ity: Hawaiian Islands. DistriBuTION: Hawaiian Islands. SPECIMENS EXAMINED: Hawaii, Hillebrand B; Robinson, 628 V; 620 V; 279 V; Wilkes Expedition N; Maui, Lichtenthaler Gs Robinson 716 V; 710 V; ex Herb. John Donnell Smith N; Oahu, Hillebrand B; Remy B; Robinson 144 V; 173 V; seoheenage Is., ex Herb. Mt. Holyoke College C. From the forms of A. lobulatum the pinnae of which are little divided, A. pseudo-falcatum is distinguished by its chartaceous texture in contrast to the coriaceous texture of A. lobulatum. — 212 RoBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS ASPLENIUM KAUuLFusstiI Schlect. Adumbr. 29. 1825 Asplenium protensum Kaulf. Enum. 167. 1824. Not Schlect. Asplenium obtusatum Underw. in Heller, Minn. Bot. Stud. 1: 775. 1897. Not Forst. TYPE LOCALITY: Oahu. DIsTRIBUTION: Higher forest regions, Oahu, Hawaii. SPECIMENS EXAMINED: Hawaii, Remy B; Baldwin B; Oahu, Anderson B; Forbes BM; Heller 2361 C; Day B; Macrae B; Kauai, Robinson &43 V; Hawaiian Is., Lydgate B, C; ex Herb. John Don- nell Smith 5909 N; Wilkes Expedition N. The sori are usually of the Asplenium type but in some cases diplazioid, while in others, the sori borne on adjacent veins face each other. Asplenium kauaiense (Hilleb.) sp. nov. Asplenium Mannii var. kauaiense Hilleb. Fl. Haw. Is. 595. 1888. Asplenium obliquum Underw. in Heller, Minn. Bot. Stud. 1: 775. 1897. Not Forst. TYPE LOCALITY: Kauai. DistTRIiBUTION: Hawaiian Islands. SPECIMENS EXAMINED: Kauai, Knudsen 103 (type) B; 104 B; 105 B; Heller 2486 C, N; Hawaiian Is., Wilkes Expedition N. ASPLENIUM ENATUM Brack. Fil. U. S. Expl. Exp. 153. 1854 Asplenium lucidum Underw. in Heller, Minn. Bot. Stud. 1: 774: 1897. Not Forst. TYPE LOCALITY: Kaala Mts., Oahu. DIsTRIBUTION: Hawaiian Islands. ILLUSTRATION: Brack. Fil. U.S. Expl. Exp. pl. 2z. fer. 1854- SPECIMENS EXAMINED: Oahu, Wilkes Expedition N; Maui, Bailey C; Bishop B; Lydgate B, C; Mann & Brigham 485 Ni Hillebrand B; Kauai, Forbes 53. BM; Heller 2692 C, N; Kauai, Van Ingen C; Robinson 840 V; Oahu, Day 79 B; Hillebrand B; Lichtenthaler N; Safford 871 N;. Wilkes Expedition C, (type) N; Hawaiian Is., Baldwin 38 C, N; Miss Sessions C; ex Herb. John : Donnell Smith 38 N; 39 N; ex Herb. Mt. Holyoke CollegeC. ASPLENIUM HILLesranpit C. Chr. Ind. Fil. 115. 1905 ‘ Asplenium Mannii Hilleb. Fl. Haw. Is. 594. 1888. Not Hook. — TYPE LOCALITY: Waianae Range, Oahu. ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDs 213 - DISTRIBUTION: Hawaiian Islands. SPECIMENS EXAMINED: Oahu, Hillebrand B, C; Safford 917 N; Hawaiian Is., Miss Sessions C. ASPLENIUM CONTIGUUM Kaulf. Enum. 172. 1824 TYPE LocALITY: Oahu, Hawaiian Islands. DISTRIBUTION: Hawaiian Islands. SPECIMENS EXAMINED: Hawaii, Mann & Brigham 160 N ; Maui, Hillebrand B; Robinson 325 V; 382 V; 712 V; Wilkes Expedition N; Lichtenthaler N; Kauai, Knudsen 140 B; Oahu, Bartsch 29 N; Forbes BM; Heller 2115 C, N; Mann & Brigham 156 N; Safford 902 N; 903 N; 911 N; Remy B; Wilkes Expedition N; Molokai, Hillebrand B; Hawaiian Is., Baldwin 4o B, C, N; 42 C; Lindley C. ASPLENIUM CAUDATUM Forst. Prod. 80. 1786 Asplenium spathulinum Hilleb. Fl. Haw. Is. 604. 1888. TYPE Locality: Hawaiian Islands. DISTRIBUTION: Tropical countries. ILLUSTRATION: Brack. Fil. U. S. Expl. Exp. pl. 22. 1854. SPECIMENS EXAMINED: Maui, Hillebrand 13 B; Robinson 632 - V; Kauai, Forbes 636 BM; Knudsen 130 B; 137 B; 141 B; 142 B; 148 B; 149 B; Hawaiian Is., Baldwin B; Gaudichaud B. . Hillebrand, Fl. Haw. Is. 604. 1888, separates Knudsen 141 and 148 as A. spathulinum on the basis that the pinnae in these specimens are more deeply incised than in the others. Robinson 632 V shows both deéply cut and simply serrate leaves on the same plant. Many of the segments are somewhat diplazioid, because sori arise on both sides of the initial vein and each extends along one of the forks. ASPLENIUM NITIDULUM Hilleb. Fl. Haw. Is. 601. 1888 Type LocaLity: Kauapali, W. Maui, Hawaiian Islands. DistrispuTION: Hawaiian Islands. SPECIMENS EXAMINED: Maui, Hillebrand B (type); Bailey C; Lichtenthaler N; Oahu, Heller 2055 C; Hawaiian Is., Baldwin N. This is separated from A. contiguum on its firmer character and the acuminate apices of its pinnae in contrast to the lax and often recurving pinnae of the latter. 214 RoBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS Heller 2055, though soriferous and apparently typical as to size and form of pinnae, has laciniate leaf blades on each plant that present the same ragged appearance that Hillebrand describes for A. contiguum var. laciniatum. Some of the. specimens are diplazioid. ASPLENIUM HORRIDUM Kaulf. Enum. 173. 1824 TYPE LOCALITY: Oahu. ; _ DISTRIBUTION: In valleys above 500 m. elevation; South Pacific Islands and Hawaiian Islands. ILLUSTRATION: Hook. Sp. Fil. 3: pl. 193. SPECIMENS EXAMINED: Hawaii, Remy B; Robinson 204 V; Maui, Bishop 55 B; Oahu, Anderson B; Forbes BM; Hillebrand B; Lichtenthaler N; Mann & Brigham 58 N; Hillebrand B; Safford 908 N; Wilkes Expedition N; Kauai, Heller 2853 C, N; Knudsen 116 B; Hawaiian Is., Baldwin 45 B, C, N; Gaudichaud B; Wilkes Expedition C, N; Miss Sessions C. — Asplenium glabratum sp. nov. Rootstock short, scaly; leaves cespitose; leafstalk 25-50 cm. long, dull green to steel-blue, naked; blade 35-70 cm. X I1I-20 cm., oblong, lanceolate, coriaceous, dull green; pinnae 5-10 cm. xX 11-12 mm., oblong-lanceolate, cut into obliquely obovate, sometimes recurving lobes; veins flabellate; sori on the anterior vein in each lobe parallel to the midrib and also on other veins. Asplenium horridum var. B. Hilleb: Fl. Haw. Is. 604. 1888. Asplenium horridum Kaulf. var. Underw. in Heller, Minn. Bot. Studs 1: 774s 2807: TYPE LOCALITY: Woods of Kahuku and Kahana, Oahu. DISTRIBUTION: Oahu and Kauai, Hawaiian Islands. SPECIMENS EXAMINED: Kauai, Heller 2588 C. (type), N; Hawaii, Robinson 2095 V; Oahu, Hillebrand B. This species is distinguished from A. horridum Kaulf. by the smoothness and dull steel-blue color of its leafstalk and midrib, and by the greater number of sori which extend along the flabellate veins of the pinnae. F ASPLENIUM LOBULATUM Mett. in Kuhn, Linnaea 36: 100. 1869 — TYPE LOCALITY: Oahu, Hawaiian Islands. : DistRisuTION: Hawaiian Islands. ROBINSON: PTERIDOPHYTA OF THE HawatlAN IsLANps 215 SPECIMENS EXAMINED: Hawaii, Mann & Brigham 167 N; Robinson 612 V; 707 V; Wilkes Expedition N; Maui, Bailey C; Robinson 348 V; 371 V; Oahu, Heller 27175 C, N: Mann & Brigham 517 N; Oahu, Robinson 138 V; 146 V; Kauai, Van Ingen C; Miss Sessions C, The following specimens are diplazioid in certain pinnules: Bailey C; Van Ingen C; Meyen B; Miss Sessions C; Wilkes Expedi- tion N ASPLENIUM VARIANS Hook. & Grev. Ic. Fil: pl. 172. 1830 TYPE LocaLity: Nepal. : DISTRIBUTION: Japan, China, Ceylon, Africa, Hawaiian Is- lands. : ILLUSTRATION: Hook. & Grev. Ic. Fil. pl. 172. 1829. SPECIMENS EXAMINED: Maui, Lichtenthaler N; Baldwin 53 B, C, N; ex Herb. John Donnell Smith N. Hooker and Greville state that this is not mentioned in Wal- lich’s list of the plants in the herbarium of the East India Company, though he collected it at Nepal in 1818 and presented the type specimen to Dr. Hooker at Kew (Ms. Herb. Hook.). ASPLENIUM MacrakE! Hook. & Grev. Ic. Fil. pl. 217. 1831 Asplenium strictum Brack. Fil. U. S. Expl. Exp. 168. 1854. Asplenium erectum Macraei Hilleb. FI. Haw. Is. 590. 1888. Asplenium rhizophyllum Heller, Minn. Bot. Stud. 1: 775. 1897. Not Kunze. Asplenium lunulatum var. C. Chr. Ind. Fil. 119. 1906. TYPE LOCALITY: Oahu. DisTRIBUTION: On ground in open woods, Hawaiian Islands. ILLUSTRATION: Hook. & Grev. Ic. Fil. pl. 217. 1831. SPECIMENS EXAMINED: Hawaii, Hillebrand B; Safford 99 N; Oahu, Heller 2117 C, N; Hillebrand B; Mann & Brigham 173 N; Macrae B; Robinson 136 V; 139 V; 164 V; Safford 897 N; Wilkes Expedition N; Kauai, Forbes 390 BM; 2764 C, N; Knudsen 110 B; 133 B; 145 B; 146 B; 148 B; Lichtenthaler N; Wilkes Expedition N; Hawaiian Islands, Baldwin 36 C; Miss Sessions C; ex Herb. John Donnell Smith N; Wilkes Expedition C, N. Hillebrand’s Asplenium erectum e myriophyllum doubtless be- 216 RoBINson: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS longs here. Heller 2764, Knudsen 145, Baldwin 36, and Gaud- ichaud (B) conform to the e type, which has slender, spaced pinnules, while Heller 2117 and Heller 2764 have both open and the compact forms of leaf upon the same plant. Occasionally a sorus is diplazioid. Hillebrand describes A. erectum as having a smooth rachis, while A. Macraei has a winged rachis. The original description of A. lunulatum (Sw. Jour. Bot. Schrad. 18007: 52. 1801) gives the pinnae as rhomboid, ovate, crenulate; the African specimens are simply pinnate, the pinnae being auricled. ASPLENIUM LypGATEI Hilleb. Fl. Haw. Is. 596. 1888 Type LocALity: Niu, Oahu. DISTRIBUTION: Known from type locality only. The plant is represented at Berlin by two specimens collected — by Lydgate in 1871. Asplenium Goldmannii Underw. in herb., nom. nov. Tarachia polyphylla Presl, Epim. 83. 1849. Asplenium polyphyllum Hilleb. Fl. Haw. Is. 607. 1888. Not Bert. TYPE LOCALITY: Oahu, Hawaiian Islands. DISTRIBUTION: Hawaiian Islands. ILLUSTRATION: Mett. Aspl. pl. 5. f. 23. ae SPECIMENS EXAMINED: Oahu, Baldwin 51a B, C; Forbes BM; ‘ Wilkes Expedition N; Lydgate B; Hawaiian Islands (collector — unknown) C. ASPLENIUM ACUMINATUM Hook. & Arn. Bot. Beech. 106. 1832 TYPE LOCALITY: Hawaiian Islands. DISTRIBUTION: Hawaiian Islands. SPECIMENS EXAMINED: Maui, Bailey C; Baldwin 51b Bi 51c B; Hillebrand C; Robinson 376 V; Oahu, Forbes BM; Hille- brand B; Robinson 149 V; 151 V; 186 V; Wilkes Expedition Ni Lichtenthaler N; Kauai, Forbes 85 BM; 140 BM; Robinson 428 Vi : 479 V; 488 V; 497 V; 702 V; Hawaiian Islands, Baldwin 51 Cy a N, C; Mann & Brigham 494 N; Wilkes Expedition N; ex He John Donnell Smith N. ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDS 217 Asplenium rhipidoneuron nom. nov. Asplenium furcatum Hilleb. Fl. Haw. Is. 604. 1888. Not Thunb. TYPE LOCALITY: Haleakala, Maui. DIsTRIBUTION: Hawaiian Islands. ILLUSTRATION: Hook. & Grev. Ic. Fil. pl. 189. 1831. SPECIMENS EXAMINED: Hawaii, Baldwin 50 B, C, N; Wilkes Expedition N; Maui, Bailey C; Baldwin 44 (type) B, C, N; Robin- son 353 V; 302 V; 7960 V; Wilkes Expedition N; Oahu, Mann & Brigham 261 N; Wilkes Expedition N ; Kauai, Forbes 58 BM; Heller 2872 C, N; Van Ingen C; Hawaiian Islands, Baldwin 49 C; Wilkes Expedition C. A. rhipidoneuron may be separated into two forms. Pinnae under 4 cm. long, rhombic. Pinnae 6-15 cm. long, acuminate. This will scarcely separate them as two species, but it is interesting as it suggests the possible crossing of A. lobulatum and A. cuneatum. ASPLENIUM INsITICcCIUM Brack. Fil. U. S. Expl. Exp. 161. 1854 Asplenium cristatum Brack. Fil. U. S. Expl. Exp. 163. 1854. Not Lam. TYPE LocaLity: Hawaiian Is., in forests. DistTRIBUTION: Hawaiian Is., Philippine Is., New Caledonia. ILLUSTRATION: Brack. Fil. U. S. Expl. Exp. pl. 22. f.2. 1854. SPECIMENS EXAMINED: Hawaii, Wilkes Expedition N; Mann & Brigham N; Maui, Bailey C; Hillebrand B; Lichienthaler N; Waiopua, Lichtenthaler N; Oahu, Baldwin B; Forbes BM; Heller 2310 C, N: Hillebrand B; Nuuanu, Robinson 142 V; Hillebrand B; Hawaiian Is., Baldwin 41 C; 43 C, N; Wilkes Expedition N; ex Herb. John Donnell Smith N. A. insiticium Brack. is a more delicate fern than A. lobulatum. The following specimens are diplazioid: Asplenium insiticium grandipinnatum Hillebrand B; Asplenium insiticium pseudoniti- dum Hillebrand B. 218 ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS ASPLENIUM CUNEATUM Lam. Encyc. 2: 309. 1786 TYPE LOCALITY: Jamaica, B. W. I. DISTRIBUTION: West Indies, Philippine, Fiji, Samoan, and Hawaiian Islands. ILLUSTRATIONS: Sloane, Hist. Jam. 1: 46.f.2. 1707. Brack. Fil. U. S. Expl. Exp. pl. 27. 1854. SPECIMENS EXAMINED: Maui, Robinson 344 V; 702 V; Kauai, Heller 2865 C, N; ex Herb. John Donnell Smith N. This differs from A. insiticium in texture and color and is smaller than the average specimen of the latter, though their dimensions overlap. ASPLENIUM PARALLELUM Baker, Hook. & Baker, Syn. Fil. 486. 1874 Asplenium bipinnatum Hilleb. Fl. Haw. Is. 595. 1888. TYPE LocaLity: Niu, Oahu. DISTRIBUTION: Known from type locality only. SPECIMENS EXAMINED: Hillebrand B. This plant is represented by one sheet labelled Asblntum flaccidum which is placed with Asplenium Lydgatei, at Berlin. It differs from A. Lydgatei in being more sturdy in habit and firmer — in texture. ASPLENIUM PATENS Kaulf. Enum. 175. 1824. Not Hook. & Arn. Not Gaud. TYPE LOCALITY: Oahu. DISTRIBUTION: Hawaiian Islands. ILLUSTRATION: Hook. Sp. Fil. 3. pl. 203. 1860. SPECIMENS EXAMINED: Oahu, Hillebrand B; Maui, Hillebrand B, K; Lydgate B. This rare species resembles A. cuneatum in habit in spite of its larger size and tripinnate divisions. ASPLENIUM ADIANTUM-NIGRUM L. Sp. Pl. 1081. 1753 A. patens Gaud. Voy. Freyc. Bot. 320. 1828. Not Kaulf. TYPE LOCALITY: Southern Europe. DISTRIBUTION: Tropical countries. a SPECIMENS EXAMINED: Hawaii, Lichtenthaler N; Mann : ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDS 219 Brigham 259 N; District of Waimea, Wilkes Expedition N; Mauna Kea, Wilkes Expedition N; Mauna Loa, Wilkes Expedition N; Maui, Hillebrand B; Haleakala, Hillebrand B; Finsch 8 B; Wilkes Expedition N; Bailey C; Kauai, Knudsen B; Wilkes Expe- dition N; Hawaiian Islands, Baldwin 48 B, C, N, V; Gaudichaud B; Wilkes Expedition B, C; Douglas B; Hillebrand B; ex Herb. John Donnell Smith N. Gaudichaud’s specimen in the Berlin Herbarium is labelled ‘Forma acutum Bory, var. Onopteris L.,”’ but the plant is not distinguishable by dentation or by other characteristics from other specimens of A. Adiantum-nigrum. ASPLENIUM VEXANS Heller, Minn. Bot. Stud. 1: 776. 1897 TYPE LOCALITY: Slopes of Konahuanui, above Manoa, Oahu. DistrRIBUTION: Hawaiian Islands. SPECIMENS EXAMINED: Oahu, Heller 2058 (type) C; Robinson V; Baldwin V. ASPLENIUM SCHIZOPHYLLUM C. Ch. Ind. Fil. 131. 1905. Not Sw. A. dissectum Brack. Fil. U. S. Expl. Exp. 170. 1854. Not Sw. Type LocaLity: Hawaii. DisTRIBUTION: On ground and on trees, Hawaiian Islands. ILLUSTRATION: Brack. Fil. U. S. Expl. Exp. pl. 24. 1854. SPECIMENS EXAMINED: Hawaii, Baldwin 51d B, C; Hillebrand B, C; Lydgate B; Maui, Hillebrand B; Oahu, Wilkes Expedition N; Kauai, Forbes 361 BM; Heller 2765 C, N; Knudsen 113, 115 B; Johnson B; Hawaiian Islands, Baldwin 49 N; Wilkes Expedi- tion N. ASPLENIUM SPHENOTOMUM Hilleb. Fl. Haw. Is. 599. 1888 TYPE LocaLity: Hawaiian Islands. Distripution: In forests at 1,300-1,600 m. elevation, Hawai- ian Islands. SPECIMENS EXAMINED: Hawaii, Hillebrand B; Maui, Baldwin 52 B; Kauai, Forbes 339 BM; 420 BM; Knudsen 108 B; 117 B; Hawaiian Islands, Baldwin 52 N. 220 RoBINsSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS SPECIES INQUIRENDAE ASPLENIUM PELLUCIDUM Lam. Encyc. 2: 305. 1786 TYPE LOCALITY unknown. DISTRIBUTION: Malaysia-Polynesia. SPECIMENS EXAMINED: Ex Herb. Kunth; Meyen B. One specimen with auricled pinnae 52-jugate, and correspond- ing in other respects to A. pellucidum Lam. is in the Berlin Her- barium and was originally in Kunth’s Herbarium given by Dr. Meyen in 1833. The specimen is no doubt correctly named but correctness of the locality seems doubtful. ASPLENIUM MEIOTOMUM Hilleb. Fl. Haw. Is. 596. 1888 TYPE LOCALITY: Wailupe, Oahu. a DISTRIBUTION: Known from type locality only. SPECIMENS EXAMINED: Hillebrand B. This is represented by a single sterile specimen at Berlin, but Hillebrand’s description proves that this is not his entire collection — of the plant. It may be a teratological form. ASPLENIUM KNUDSENII Hilleb. Fl. Haw. Is. 601. 1888 No specimen so labelled or corresponding to Hillebrand’s : description was found in the Berlin Herbarium. : 27, .ATHYRIUM Roth, Mag. Bot. Roem. & Ust. 2!: 105. 1799 Rootstock short, suberect; leafstalks non-articulate, basal ‘ scales usually thin-walled; blades thin, herbaceous, 1—3-pinnate; veins free; vascular bundles of leafstalk in two strands at base, uniting above; sori linear or curving across the vein, or roundish . at the end of a vein; indusia corresponding with the sorus in shape — or rarely absent. 4 Leafblades bipinnatifid. Sori on distal portion of veins but not protruding beyond margin of lo Sori protruding beyond the margin of the lobe, in some cases apparently stipitate. Leafblades tripinnate to quadripinnate. ri med ssn bos-iogy bine — ae hes one sear eaee linear. ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDs 221 ATHYRIUM DEPARIOIDES (Brack.) Christ, p. p. Farnkr. Erde 223. 1897 [Excl. description and figures] Asplenium deparioides Brack. Fil. U. S. Expl. Exp. 172. 1854 Athyrium proliferum C. Chr. p. p. Ind. Fil. 145. 1905. TYPE LOCALITY: Kaala Mts., Oahu. DIsTRIBUTION: Hawaiian Islands. SPECIMENS EXAMINED: Oahu, Baldwin B; Baldwin 77 B; Forbes BM; Hillebrand B; Lydgate B; Kauai, Forbes 32 BM; Forbes 215 BM; Forbes 238 BM; Heller 2303 C; Heller 2690 C; Baldwin 47b C; Hawaiian Is., Baldwin 47 C. ATHYRIUM PROLIFERUM (Kaulf.) C. Chr. p. p. 145. 1905 Dicksonia prolifera Kaulf. Enum. 225. 1824. Deparia Macraei Hook. & Grev. Ic. Fil. pl. 154. 1829. Cibotium proliferum Presl, Tent. Pterid. 69. 1836. Asplenium deparioides var. y, Hilleb. Fl. Haw. Is. 615. 1888. Athyrium deparioides Christ, p. p. Farnk. Erde 223. 1897. Deparia triangularis Underw. in Heller, Minn. Bot. Stud. 1: 778. 1897. TYPE LOCALITY: Oahu. DisTRIBUTION: In wet woods, 300-600 m. elevation, Hawaiian Islands. ILLUSTRATIONS: Hook. & Grev. Ic. Fil. pl. 154. 1829; Christ; Farnk. Erde 223. 1897. SPECIMENS EXAMINED: Oahu, Baldwin B; Beechey C; Bracken- ridge N; Forbes BM; Lichtenthaler N; Kauai, Heller 2740 C, Heller 2057 C. Arayrium PorreTiaANuM (Gaud.) Presl, Tent. Pterid. 98. 1836 Asplenium Poiretianum Gaud. Voy. Freyc. Bot. 321. pl. 13. 1832. Allantodia scandicinum Kaulf. Enum. 179. 1836. Asplenium scandicinum Underw. in Heller, Minn. Bot. Stud. 1: 775. 1897. Not Presl. TYPE LOCALITY: Oahu. DIstRIBUTION: In wet forests, at altitudes of 200-300 m., Hawaiian Islands. ItLustraTIon: Gaud. Voy. Freyc. Bot. pl. 13. 1832. 222 RoBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS SPECIMENS EXAMINED: Hawaii, Robinson 616 V; 625 V;. Maui, Bailey C; Finsch B; Hillebrand B; Robinson 715 V; Oahu, Chamisso B; Diell C; Forbes BM; Hillebrand 27 B; Macrae B; Meyen B; Kauai, Heller 2623 C; Hillebrand 118a B; Hawaiian Is., Baldwin 53 B, C; Gaudichaud (type) B; Miss Sessions C; Wilkes Expedition G. ATHYRIUM BALpwInit (Hilleb.) C. Chr. Ind. Fil. 140. 1905 Asplenium Baldwinit Hilleb. Fl. Haw. Is. 618. 1888. TYPE LOCALITY: Kauai. DISTRIBUTION: High altitudes, Kauai. SPECIMENS EXAMINED: Kauai, Bailey C; Baldwin 54 (type) B, C, K; Forbes 125 BM; 240 BM; Robinson 4o2 V; 426 V; 458 V; 461 V; 464 V. The sori are subapical in this species and a fibrillose scale appears at the base of almost every pinna, while the sori of A. Poiretianum are typically medial or proximal upon the veins. 28. DIPLAZIUM Sw. Jour. Bot. Schrad. 18007: 61. 1801 Rootstock short, creeping or suberect; leafblades simple to — quadripinnatifid; basal scales of leafstalk clathrate, with or with- — out glands; vascular bundles two at the base of leafstalk uniting above; veins free, once to several times forked; sori linear, 11- dusiate, often double, the indusium opening as in Asplenium. Type species: Diplazium plantaginifolium (L.) Sw. The Hawaiian species of Diplazium form a very natural grouP, more firm in texture and coarser in habit than the species of Athyrium, hence it seems better on account of the limitations of this paper to follow Diels (Eng. & Prantl, Nat. Pflanzenfam. 1°: 224. 1902) than to place these plants in the genus Athyrium, following Milde (Bot. Zeit. 24: 373. 1866) and Copeland (Philipp- Jour. Sci. 3: (Bot.) 285. 1908) Leaves pinnate; rootstock erect or creeping; leafstalks more or less paleaceous at base, with non-glandular scales Rootstock erect; leafstalks stout, dull green, dark pecan’ in drying; basal scales few, light brown; leafblade ovate- lanceolate, 15-30 cm. long, membranous; middle pinnae linear-lanceolate, 12.5-15 cm. 7-22 mm.; basal pinnae ae reduced, the lowest auricled. D. marginale. | Rootstock creeping; leafstalks slender, stramineous; basal. 2 scales dark brown. ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDs 223 Basal scales of eatstales Perit: numerous; leaf- —t¢ oem 1] 1 b lanceolate, blade 45- falcate; besal Slaine sei D. Fenzlianum. Basal scales of leafstalk few, Gucci leafblade 15-30 m. long; middle pinnae oblong-lanceolate, truncate at the base and parallel to the rachis above, oblique below; basal pinnae not reduced. Leaves bipinnate to tripinnate; rootstock creeping; leafstalks paleaceous with oe scales at base when young, D. molokaiense. glabrate with Meir: oa or : cwaie aciona light green, deeply tri- pinnatifid; pinnae oblong-lanceolate; lower pinnae 5~7.5 cm. distant; pinnules lanceolate. Leafblade ovate-oblong, dark green, trip pinnae 12.5-15 cm. distant; pinnules oblong-lanceolate. D. Arnoitii. innate; lower D. sandwichianum. DIPLAZIUM MARGINALE (Hilleb.) C. Chr. Ind. Fil. 235. 1905 Asplenium marginale Hilleb. Fl. Haw. Is. 613. 1888. TYPE LOCALITY: Molokai. DISTRIBUTION: Lower forests, Hawaiian Islands. SPECIMENS EXAMINED: Maui, Lydgate B; Molokai, Hillebrand (type) B; Oahu, Baldwin 47a B; 476 B; Kauai, Knudsen 151 B. 1905 1875. DIPLAZIUM FENZLIANUM (Luers.) C. Chr. Ind. Fil. 232. Asplenium Fenzlianum Luerssen in Wawra, Flora 58:.434. TYPE LOCALITY: Hawaiian Islands. DISTRIBUTION: In isolated localities, Hawaiian Islands. SPECIMENS EXAMINED: Maui, Bailey C; Oahu, Baldwin B; Hillebrand B; Lydgate B; Molokai, Hillebrand B; Hawaiian Is., Hillebrand B. | Diplazium molokaiense nom. nov. Asplenium arboreum Hilleb. Fl. Haw. Is. 609. 1888. Not Willd. Type Locatity: Molokai, Hawaiian Islands. DISTRIBUTION: In wet gulches, Hawaiian Islands. SPECIMENS EXAMINED: Molokai, Baldwin 46 B; Hillebrand (type) B; Oahu, Baldwin B; Kauai, Forbes 438 BM; Knudsen 102 B 1854 Diecazium Arnotti Brack. Fil. U. S. Expl. Exp. 144. 1832. Asplenium diplazioides Hook. & Arn. Bot. Beech. 107. Not Bory. Asplenium Arnottii Baker in Hook. & Baker, Syn. Fil. 240. 1867. 224 RoBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS Athyrium Arnottii Milde, Bot. Zeit. 28: 371. 1870. TYPE LOCALITY: Hawaiian Islands. DISTRIBUTION: Lower forests, Hawaiian Islands. SPECIMENS EXAMINED: Maui, Bailey C; Molokai, Hillebrand B; Oahu, Beechey C; Beratz B; Forbes BM; Heller 2900 C; Hille- brand B; Kauai, Johnson B; Hawaiian Is., Baldwin 55 C; 56 B, C; 57 C; Mrs. Gulick C; ex Herb. Kewensis C. Though the rootstock is prostrate, this fern has almost the appearance of an arborescent form, the leaves reaching a height of from five to six feet. The Hawaiians prize the tips of the young fronds as a salad. DIPLAZIUM SANDWICHIANUM (Presl) Diels, in E. & P. Nat. Pflanzenfam. 14: 228. 1899 Athyrium sandwichianum Presl, Tent. Pterid. 98. 1836. Asplenium sandwichianum Hook. Sp. Fil. 3: 225. 1860. Asplenium brevisorum Baker, in Hook. & Baker, Syn. Fil. Ed. 2. 228. 1874 TYPE LOCALITY: Oahu, Hawaiian Islands. DISTRIBUTION: Hawaiian Islands, Peru. SPECIMENS EXAMINED: Hawaii, Beratz B; Robinson 620 V; Maui, Hillebrand B; Robinson 386 V; Molokai, Hillebrand B; Oahu, Diell C; Forbes BM; Heller 2073 C; Macrae C; Meyen B (cotype); Robinson 148 V; 156 V; 160 V; Hawaiian Is., Baldwin B; ex Herb. Mt. Holyoke College C; Wilkes Expedition C. SPECIES INQUIRENDA DIPLAZIUM SANDWICHENSE Presl, Epim. 85. 1850-52 From Presl’s description this seems to be a form very closely related to Diplazium molokaiense, differing from it in the deeper cutting of the lobes and the greater length of the auricle. Meyen’s type is doubtless still in Presl’s Herbarium in the Bohemian University at Prague. There seems to have been n0 further collection of the plant. 29. SADLERIA Kaulf. Enum. 162. 1824 Sadleria, an endemic Hawaiian genus, is frequently found as 4 pioneer on disintegrating lava rock. It differs conspicuously 1 — ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDS 225 its greater size, rigidity, and the number of its scales, from Blech- num, with which Brackenridge and Gaudichaud united it on the basis of the similarity of its sori. Caudex erect, often arborescent, more or less loti wich scales, or “‘pulu’’; leaves bipinnatifid to bipinnate, 60-180 cm. long in arborescent species, 25-50 cm. in herbaceous forms, usually coriaceous; sori linear upon the intercostal arches on either side of, and parallel to, the midrib of the pinnule, covered by a coriaceous indusium; sporangia short-stalked. Type species: Sadleria cyatheoides Kaulf. Caudex arborescent, 60-300 cm. in height. Leafstalk scaly at base, naked or slightly furfuraceous above; i usually extending nearly the length of the segment. base with light brown scales, 3-4 c chaffy above; blades chartaceous to subcoriaceous, furfuraceous when young, nearly glabrate when ma- ture, oblong-lanceolate, 150-200 cm. long, bipinnate; pinnae linear, 30-60 cm. X 4-7.5cm.; pinnules 3 mm apart. S. Souleytiana. Leafstalk 30-60 cm. long, not sulcate, paleaceous at the base with scales 4-5 ¢ mm., naked ong blades came, glabrate, ovate-oblong, 60-90 long, eg ; pinnae linear-lanceolate, 15-20 cm. 8 a | ; segments of pinnae 2 mm. apart. Leafstalk scaly oes. 20-45 cm. long, sulcate; basal revolute, upper scales ‘ian ss, chaffy; g, acuminate, 45-60 cm. long, bi- S. cyatheoides. scales ribbed, and blades coriaceous, oblong pinnatifid; pinnae oblong, falcate, 10-16 cm. X 15 mm., chaffy along the midrib; sori usually not as long as the segment, often less than half as long. Caudex not arborescent, 2-15 cm. in height; paleaceous as are midribs and costae a I 5- 25 cm. long; scales dark ieee 10-15 cm. long; , oblong-lanceolate, 25-50 cm. long, pinnae 5-Io cm. X 12-25 mm.,; pinnules crowded; margins crenu- late. S. polystichoides. Leafstalk 12-15 cm. long; scales reddish brown, 6-10 cm. long; blades subcoriaceous, ovate-lanceolate, 28- 30cm. long; cm., reduced below, not crowded; S. Hillebranditi. : leafstalk densely : S. unisora. 1857 pinnae 2-4 cm. X 0.75-I margins entire. SADLERIA SOULEYTIANA (Gaud.) Moore, Ind. Fil. 26 Blechnum Souleytianum Gaud. Voy. Bonite Bot. pl. 2. f. 7, 8; pl 134. 1846-49. TYPE Loca.ity: Hawaiian fetalte: 226 ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS DISTRIBUTION: Common in deep wet woods at high elevations Hawaiian Islands, ILLUSTRATIONS: Gaud. Voy. Bonite Bot. pl. 2. f.7,8. pl. 134. 1846-49. SPECIMENS EXAMINED: Maui, Bailey C; Kauai, Heller 2807 C; N; Oahu, Forbes BM; Hillebrand B; Lanai, Hillebrand B; Hawaiian Islands, Baldwin 24 C; N. SADLERIA CYATHEOIDES Kaulf. Enum. 162. 1824 Blechnum Fontanesianum Gaud. Voy. Freyc. Bot. 397. pl. 15. Sadleria pallida Hook. & Arn. Bot. Beech. 75. 1832. Blechnum pallidum Brack. Fil. U.S. Expl. Exp. 133, in part, 1854- TYPE LOCALITY: Hawaiian Islands. DIsTRIBUTION: Lower elevations, Hawaiian Islands and Su- matra. ILLUSTRATION: Gaud. Voy. Freyc. Bot. 24, 15. SPECIMENS EXAMINED: Hawaii, Wilkes Expedition 12 C, Nj ‘ 13 C, N; Maui, Bailey C; Oahu, Chamisso B; Freycinet K; Gaudt- : chaud B; Hillebrand B; Lichtenthaler N; Macrae C; Mann & © Brigham 188 N; Safford 866 N; 867 N; 868 N; ex Herb. Copeland N; Kauai, Forbes 40 BM; Hawaiian Islands, Baldwin 23 B, Nj Miss Sessions C; ex Herb. John Donnell Smith N. : Chamisso’s type in the Berlin herbarium consists of two : sheets: (1) Leafstalk 15 cm. long with rigid scales at base, blade + 18-jugate, 23 cm. wide; (2) fragment without apex, I 16-jugates slightly wider. : i 4 4 Sadleria Hillebrandii nom. nov. of Sadleria pallida Hilleb. Fl. Haw. Is. 582. 1888. Not Hook. & ~ Arn. Bot. Beech. 75. 1832. Blechnum pallidum Brack. Fil. U.S. Expl. Exp. 133, in part. 1854 — Type LocaLity: Kauai, Hillebrand 80 (Herb. Kgl. Bot. Gart- — Berlin). DisTRIBUTION: In dry, exposed places, Hawaiian Islands. ILLUSTRATION: PLATE II. SPECIMENS EXAMINED: Hawaii, Hillebrand B; Maui, Batley C Oahu, Hillebrand 83 B; Robinson 14 V; 19 V; Kauai, Hillebrand 80 (type) B; Heller 2866 C, N. Sadleria pallida Hook. & Arn. proves to be a synonym s ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN IsLANDS 227 “Sadleria cyatheoides Kaulf. The plants confused under this name’ by Brackenridge, Hillebrand, and most later writers form a distinct species, here named for Hillebrand, since he has most fully de- scribed it. This species is one of the first to gain a foothold upon the lava in the floors of the old craters and is very resistant to dry winds on exposed bluffs. SADLERIA POLYSTICHOIDES (Brack.) Heller, Minn. Bot. Stud. 1: 788. 1897 Blechnum squarrosum Gaud. Voy. Bonite Bot. pl. 2. f.1,6. 1846- 49. Without description. Blechnum polystichoides Brack. Fil. U. S. Expl. Exp. 134. 1854. Sadleria squarrosa Hilleb. Fl. Haw. Is. 582. 1888. TYPE LocaALity: Hawaiian Islands. DIsTRIBUTION: In forests on lower slopes of mountains, Hawaiian Islands. ILLUSTRATIONS: Gaud. Voy. Bonite Bot. pl. 2. f. 1,6. 1846- 9. SPECIMENS EXAMINED: Hawaii, Lichtenthaler N; Wilkes Expe- dition N; Oahu, Forbes BM; Hillebrand B; Lydgate B; Heller 2392 C, N; Molokai, Hillebrand B; Kauai, Forbes 216 BM; Hawatian Islands, Gaudichaud B; Baldwin 25 B, C, N. Sadleria unisora (Baker) comb. nov. Polypodium unisorum Baker; Hook. & Bak. Syn. Fil. 307. 1867. Gymnogramme sadlerioides Underw. in igre Minn. Bot. Stud. 1: 781. 18097. Sadleria squarrosa var. depourivait Hilleb. Fl. Haw. Is. 583. Type LocaLity: Hawaiian Islands. DISTRIBUTION: On Sain & walls of high plateaus, Hawaiian Islands. ILLUSTRATIONS: Underw. in Heller, Minn. Bot. Stud. 1: pl. 43. 1897. PLATE 12. SPECIMENS EXAMINED: Kauai, Hillebrand B; Forbes 445 BM; Heller 2863 C; N. . ; This is undoubtedly a valid species, rather than an ecological “variety” of S. polystichoides, as it has been termed by Hillebrand (Hilleb. Fl. Haw. Is. 583. 1888), for true S. polystichoides grows in 1888. 228 ROBINSON: PTERIDOPHYTA OF THE HAWAIIAN ISLANDS the neighborhood of Kilauea, under conditions apparently as xerophytic as those under which S. wnisora is found. 30. DOODIA R. Br. Prod. Fl. N. Holl. 151. 1810 Rootstock short, oblique; leaves clustered, pinnatifid to pin- nate, coriaceous; leafstalk scaly; veins connected by one or more arches parallel to the midrib; sori oblong or slightly curved, in one or more rows parallel to the midrib; indusium membranaceous, opening toward the midrib. Type species: Doodia aspera R. Br. Doop1a KuNnTHIANA Gaud. Voy. Freyc. Bot. 401. 1829 Doodia media Hilleb. Fl. Haw. Is. 584. 1888. Not R. Br. Doodia media var. C. Chr. Ind. Fil. 242. 1905. TYPE LocALity: Hawaiian Islands. a DISTRIBUTION: Common on banks of streams or in wet woods at 500-900 m. altitude; Hawaiian Islands. ILLUSTRATION: Gaud. Voy. Freyc. Bot. pl. 14. SPECIMENS EXAMINED: Hawaii, Robinson 230 V; 241 V; Maui, : Robinson 305 V; 318 V; 350 V; Oahu, Baldwin N; Macrae Bie ‘Robinson 74 V; Safford 895 N; Wilkes Expedition N; Kauai, a , Forbes BM; 252 BM; Heller 2601 C, N; Van Ingen C; Robinson — | 803 V; 809 V; Hawaiian Islands, Baldwin 26 B, C, N; Baldwin 27 B, C; Gaudichaud B, C; Miss Sessions C; ex Herb. J. Donnell — | : : | Smith N. This species has been confused by Hillebrand and others with : D. media R. Br. (type from Australia), the lower pinnae of which become reduced to small triangular auricles. It is safer to main- ae tain the species as distinct under Gaudichaud’s name. An allied 4 | species with widely spaced pinnae, collected in the Fiji Islands by Capt. Wilkes, is mentioned by Brackenridge (Fil. U. 5- Expl. 2 Exp. 138. 1854) as a “var. B.”’ a : Mr. C. N. Forbes’s D. Kunthiana var. depauperata (252 BM), a collected ‘‘on rocky ledges along the main Wahiawa,” is so MUC™ smaller than the other plants of D. Kunthiana as to have gi entirely different appearance. However, as to the form of the leaf, pinnate below, pinnatifid in the upper half, and the form of t linear-lanceolate scales, they are similar. These small plants # fine fruiting condition. a Neer The development of the embryo-sac of Arisaema triphyllum F. L. Picketr (WITH PLATES 13 AND 14) The development of the embryo-sac of Arisaema triphyllum was first studied by Strasburger (1) in 1879, and later by Mottier (2), who published his observations in 1892. Campbell (3, 4) in his studies on the Araceae has referred to the findings of Mottier and used them as a basis for comparison. Gow (5), in his intro- duction to the study of the embryogeny of Arisaema triphyllum, reviewed the development of the embryo-sac, confirmed the findings of the second study mentioned above, and added observa- tions of the division of a primary archesporial cell to form embryo- sac initials. The writer has been able, after the examination of several hundred ovules, to verify most of the stages in the embryo- sac development as set forth in the above-mentioned articles; but has found some notable points of difference touching the formation of megaspores from the primary archesporial cell. On the other hand the writer has found that in some respects the development of Arisaema triphyllum agrees very closely with that of Symplocarpus foetidus as shown by Rosendahl (6) in 1909. The material for the present study was collected in proper season through four years, 1909-12. The various stages were secured from as many habitat conditions as possible. All fixing was done in the field or immediately after the return to the laboratory. Parts of young spikes and separate pistillate flowers of older spikes were fixed, some in strong chrome-acetic acid mixture, and others in chrome-osmic-acetic acid mixture, for 24-30 hours, then washed in running water for 24 hours, and finally gradually dehydrated. The blackening from osmic acid was in some cases removed by treatment of the whole specimens with hydrogen peroxide; but this resulted in considerable shrinkage, so that the specimens were of very little value. The sections were stained with safranin-gentian violet-orange G, with occasional special staining of cell walls with Bismarck brown. 230 PICKETT: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM OVULE AND MEGASPORE MOTHER CELL The ovules occur in groups of two to six arising from the basal placenta in each ovary. As it first appears, each ovule is a rounded conical mass of cells covered by a clearly marked epi- dermal layer, but otherwise undifferentiated (FIG. 4). Soon after the ovule has reached the stage shown in FIG. 4, or about the time of the appearance of the inner integument, the primary arche- sporial cell, or megaspore mother cell or cells, may be found at the apex of the nucellus beneath the epidermis. These cells are hypo- dermal in origin, and are clearly differentiated from surrounding cells by their greater size, denser cytoplasm, and larger nuclei. The time of the differentiation of the sporogenous cells is only approximately the same as that of the development of the first integument as will be seen by a comparison of FIG. I, 2, 3, 6, 8. Fic. 1 and 2 show two and three clearly differentiated megaspore mother cells, and FIG. 6 one such cell with complete periclinal divisions of the epidermal cells, while the integument in each of these ovules shows less growth than in FIG. 3 where no such cell can be with certainty distinguished. From one to four megaspore mother cells have been observed in a single nucellus (FIG. 1, 7, 8, 9" 11). These are usually in the hypodermal layer at first, although in a few instances some were beneath that (FIG. 2, 11, 19). The cells in FIG. If and 19 have been removed from the surface by periclinal division of epidermal cells. In every specimen examined the megaspore mother cells were contiguous, but in no case was there found direct evidence that they had been formed by a division of a primary archesporial cell as suggested by Mottier (2, p. 259): and by Gow (5, p. 40). The position of the cells in FIG. 19 and of the two cells at the left in FIG. 11 indicates that such an origin is possible; but the failure to find other than very rare cases of nuclear activity in cells which have not arisen by the division of epidermal cells discredits the probability of such an origin. The only form of nuclear activity found in clearly differentiated sporo- genous cells was the synaptic or later stages in the tetrad division. The fact that in different ovules showing the same general struc- ture, more than one megaspore mother cell may be found, suggests the origin of such multiple sporogenous cells by simultaneous differentiation from hypodermal cells (see FIG. 1, 2, 8). That PICKETT: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM 231 the division of a primary archesporial cell to form two or more megaspore mother cells or embryo-sac initials is not universal, or even regular, is proven by the presence of many ovules showing the development of but one tetrad of potential megaspores or but one embryo-sac, without evidence of other crushed or damaged initial cells (FIG. 17, 18, 22, 23, 24). Fig. 7, 9, 13 show cross sections of the nucellus with the megaspore mother cells variously arranged. THE TETRAD OF MEGASPORES As the inner integument grows and incloses the nucellus, and the outer integument appears, the megaspore mother cells increase in size, and the epidermal cells of the nucellus undergo periclinal division. This periclinal division, shown at its beginning in FIG. 2 and 5, takes place more rapidly at the apex, sometimes forming a layer of four cells above the megaspore initials (FIG. 16, 18, 22). The division of lateral cells is also irregular and results in a final covering one to three layers of cells in thickness (FIG. 10, II, 24). During these changes in the size and form of the ovule, the mega- spore mother cells increase rapidly in size, becoming somewhat broader and much longer than the surrounding cells. Nuclear changes are also evident, and by the time the outer integument is well started most of the nuclei of the megaspore mother cells show either synapsis or closely related conditions preparatory to the first tetrad division (FIG. 7, 9, 12, 14, 16). The first division of the megaspore mother cell nucleus is quickly followed by the second, the result being a tetrad of potential megaspores (FIG. 18). The first division is usually followed by the formation of a cell wall as in FIG. 22 and 24, but this is not always the case, as 1s shown in FIG. 23. These divisions are usually transverse as shown in the cell at the left in F1G. 16, in the middle cell of Fic. 15a, and in FIG. 24. The resulting megaspores usually lie in a row parallel to the long axis of the nucellus (FIG. 18). That the first division may be longitudinal is shown by preparations similar to that in FIG. 10, and that the later divisions may vary from a trans- verse direction is shown in the upper cell in FIG. 22. From such divisions we might expect the formation of two parallel rows of two megaspores each, or a row of two parallel to the long axis of the nucellus with another transverse pair, as described for Symplo- 232 PICKETT: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM carpus foetidus by Rosendahl (6, p. 3). Where more than one megaspore mother cell is in a nucellus, division of all the nuclei proceeds regularly and may or may not be simultaneous. The most nearly simultaneous cases found are shown in FIG. 9, 19, 19d. Finally, limited space probably causes a crowding in such cases, which results in the checking of activity in some cells and their destruction by the growth of others, as suggested by the cells at the left in FIG. 15, 15a. A few preparations show a cell in a position to indicate its origin as a tapetal cell (FIG. 17). Thesame possibility is suggested by Gow (5, p. 39-40), in FIG. 6, and is also mentioned by Campbell (3, p. 7) for Dieffenbachia Seguine. Such a tapetal structure is shown to be well developed normally in Symplocarpus | foetidus by Rosendahl (6, p. 2, pl. 1. f. 4 and pl. 3. f. 31). The finding of but one such cell showing disorganization as the mega- spores develop seems ‘to the writer sufficient evidence to justify the conclusion that the formation of a functional tapetum in Arisaema triphyllum is unusual or even questionable. As has been stated above, during the enlargement and divisions of the megaspore mother cells, the outer cells of the nucellus undergo periclinal divisions until the sporogenous cells appear quite deeply seated. During the same time the nucellar cells become well filled with starch (FIG. 15 and 17). Asa result of the later gametophyte growth, the lateral portion of the nucellus with its contained food material is consumed, so that the mature embryo- sac is in contact with the inner integument, except at the apex, where a cap of nucellar cells persists (FIG. 20 and 21). The temporary storage of food where it is so readily available for use by the growing embryo-sac may account for the rapid develop- ment of the latter as given below. THE EMBRYO-SAC One of the megaspores develops into the embryo-sac at the expense of the others and of the nucellar tissue. Three nuclear divisions occur in rapid succession as in Symplocarpus foetidus (6, p. 3). The short time required for these divisions is indicated by the fact that out of many preparations of material gathered at one time, and showing about this stage, but very few show PICKETT: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM 233 anything between the first nuclear division of the megaspore (FIG. 20) and the mature embryo-sac (FIG. 20). The daughter nuclei of the first division remain centrally located in the cell until after the second division (F1G. 26). Later a normal egg cell and two synergidae are formed close in the apex of the embryo-sac, and in some cases three antipodals are present (FIG. 20). The antipodal cells are poorly developed however, only occasionally showing typical angiosperm structure, and never showing the remarkable development described by Campbell (3, p. 11) and Rosendahl (6, p. 5) for other members of the Araceae. As far as the writer has been able to determine, the polar nuclei show no such fusion as described by Gow (5, p. 41), but remain separate as in Anthurium (3, p. 15). In every case where a polar nucleus was observed after fertilization of the egg, the lower one was found separate, greatly enlarged, and often disintegrating near the antipodals. Further details concerning the fate of the polar nuclei and of the antipodal structure will be more fully dealt with in a subsequent paper. Where more than one tetrad of potential megaspores are formed in one nucellus, the usual course of events is for some one cell from one tetrad to germinate while the other spores are crushed or broken down. In rare cases more than one embryo-sac have been found developing in the same nucellus. In one ovule two well-developed embryo-sacs were found side by side and of approxi- mately the same size, but unfortunately one lay directly beneath the other in the section. The only preparation showing two such structures in the same section is shown in FIG, 21. These present a marked difference in size as well as deficiencies in antipodal structure. As stated above, such imperfect development of anti- podals is not rare in otherwise normal embryo-sacs. SUMMARY The present study has verified most of the findings of other investigators, but has given results at variance with their reports in regard to the following points. (a) The origin of the several megaspore mother cells from a single primary archesporial cell is doubtful. The first division in the formation of the tetrad has been prob- ably mistaken by earlier investigators for a division of a primary archesporial cell into embryo-sac initials. 234 Pickett: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM (6) The formation of a tetrad of potential megaspores from some of which the embryo-sac or sacs develop, instead of from an archesporial cell, is shown. (c) More than one embryo-sac may be developed in Arisaema triphyllum, as Campbell (4, p. 671) has shown for other members of the Araceae. (d) The fusion of the polar nuclei is doubtful. (e) The antipodal cells rarely develop fully as in typical angiosperms. The writer wishes to express his gratitude to Prof. D. M. Mottier of Indiana University, at whose suggestion the present study was undertaken, for his kindly critical and encouraging words. INDIANA UNIVERSITY, BLOOMINGTON, INDIANA. LITERATURE CITED - * Strasburger. Die Angiospermen und die Gymnospermen. 1879. - Mottier. On the development of the embryo-sac of Arisaema triphyllum. Bot. Gaz. 17: 258-260. Au 1892. N 3. Campbell. Studies on the Araceae. Ann. Bot. 14: 1-25. Mr 1900, 4. Campbell. Studies on the Araceae. Ann. Bot. 17: 665-687. S 1903 Gow. The embryogeny of Arisaema triphyllum. Bot. Gaz. 45: 38-44. Ja 1908. Rosendahl. Embryo-sac development and embryology of Wheto carpus foetidus. Minn. Bot. Stud. 4: 1-10. Je 1909. on a Description of plates 13 and 14 All drawings were made with the aid of a camera lucida. Fic. 1. Longitudinal section of a young ovule with two megaspore mother cells in normal position shortly after the beginning of differentiation. M225: Fic. 2. Longitudinal section of young ovule with three SL fer dif- ferentiated. The two lower cells may be considered as lateral hypoder X225. Fic. 3. Longitudinal section of ovule slightly larger and older than those shown in FIG. I and 2, but without any clearly differentiated sporogenous cells. 225. Fic. 4. Longitudinal section of very young ae before differentiation of ee cells and beginning of integuments. 33 FIG. 5. onevedinat — rat = showing lateral megaspore mother cell, ible t g beginning of p nal division of epidermal a cells. X225. re Me “3 £7 ST Po oe oT ee aa PICKETT: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM 235 G. 6. Longitudinal section of ovule showing one megaspore mother cell and completed periclinal division of epidermal cells. 225. Fr Cross section of nucellus showing layer of 2 to 3 cells surrounding three megaspore mother cells. 335. Fic. 8. Longitudinal section of the whole ovule and part of ovary walls. 225. Fic. 9. Cross section of nucellus similar to ric. 7, but showing different ar- rangement of megaspore mother cells. This shows very nearly simultaneous preparation for nuclear division. an 335. Fic. 10. Cross section of nucellus showing the spindle of a longitudinal first division of a megaspore mother cell. 335. Fic. 11. Longitudinal section of nucellus with four megaspore mother cells. The position of the two at the left suggests a possible origin from a common arche- sporial cell. 225; FIG. 1 Longitudinal section of ovule to show the development of the integu- ments at the time of preparation for first tetrad division. 125. Fic. 13. Cross section of nucellus to show irregularity of periclinal division of outer cells. 335. Fic. 1 Longitudinal section of ovule to show the development of integuments at the time of the first tetrad division. 125. (See FIG. I Fic. 15. Longitudinal section of nucellus showing some sporogenous cells Sraieee by the growth of others. X250. Fic. 15a. Sporogenous cells of FIG. 15. 30. iG. 16. Higher magnification of the nucellus shown in FIG. 14. 250. IG. 17. Longitudinal section of nucellus showing a possible tapetal cell, ¢. I itudinal secti f llus.showing a row of four potential mega- 1G. 1 aaa section of nucellus showing two megaspore mother cells, one below the other. 50. Fic. 19a. Sporogenous cells of FIG. 19. 830. The nuclei are in synapsis. and the wall formed after the first division. This shows the sporogenous cells buri unusually dee 50 IG. 23. ‘ubekaues 22 Fic. 24. A later stage than in ip se x 250. Fic. 25. Section of a the first nuclear division. 830. Fic. 26. Section showing fous nuclear stage in the development of the embryo- 1 c W wall £ e FG REM ee Nan Bes ot 250. Is salinity a factor in the distribution of Nereocystis Luetkeana? * GrorGE B. RicG Under an appointment from the United States Bureau of Soils the writer has investigated the kelps of the Puget Sound region as a source of potash fertilizer. This work was done during the summers of 1911 and 1912. During 1912 attention was given in particular to a study of the effect of fresh water on the growth of the bladder kelp (Nereocystis Luetkeana) as seen in the bed in Freshwater Bay near the mouth of the Elwha River. The Elwha is a snow-fed stream originating on the southwestern slope of Mount Olympus. It flows north and discharges into the Strait of Juan de Fuca some six miles west of the city of Port Angeles, Washington. It is the largest stream flowing into the Strait of Juan de Fuca or Puget Sound from the Olympic Mountains. The monthly maximum and minimum discharge of this river is reported in the Report of the United States Geological Survey,f in cubic feet per second, for the period of October, 1897, to December, 1898. For the portion of the year 1897 covered by this report the maxi- mum discharge occurred in November and was 7,075 second feet, while the minimum occurred in October and was 171 second feet. For 1898 the maximum (3,282 second feet) occurred in June and the minimum (330 second feet) occurred in October. The mean for the period reported is 1,444 second feet. These observations were taken at McDonald, Washington. This is above the outlet of Lake Sutherland and also the outlet of Little River, so that the actual discharge of the river was somewhat greater than the above - figures show. Mr. G. W. Northrup, superintendent of the opera- tion of the power plant of the Olympic Power Company on the Elwha, states that the minimum discharge of the river for the year 1912 has been between 400 and 500 second feet, and that the maximum reaches an enormous amount each year during flood water periods, that is, in May and November. + Published by permission of the Secretary of Agriculture. | * Twentieth Annual Report of the United States Geological Survey, Part IV, Pp. 531 237 Sore va eee, Yater B _ » Sante . CPs. tl xy | Fic. 1. Map of Freshwater Bay, Washington. Drawn from U. S. Coast and Geodetic Survey [Chart no. 6300; kelp bed (Nereo- cystis) shown by ////////; soundings given in pete dotted area indicates shoal water. SILSADONAAN AO NOILASITMISIG AHL GNV ALINITVS “OOTY S&S RiGG: SALINITY AND THE DISTRIBUTION OF NEREOCYsTIS 239 Freshwater Bay does not form a harbor. The distance in a straight line from Angeles Point, which marks the eastern side of the bay, to Observatory Point at its western side is four miles, while the maximum distance to shore in the bay, measured at a right angle from the above line, is only one mile. The strong tidal currents flowing in and out through the Strait of Juan de Fuca sweep freely through this bay. There is a good beach along prac- tically the whole bay. The field observations on the kelp beds in this bay were made on September 11 and 12. On September 11, the surf was so heavy that it was deemed impracticable either to enter the mouth of the river in the 50-foot launch in which the trip was made, or to land with a skiff in the more exposed portion of the bay. The launch was anchored in the more protected portion of the bay behind Observatory Point. A landing was readily made with the skiff from this point. On the afternoon of September 12, the water in the bay was much quieter and we entered the mouth of the river in the launch. There is no kelp at all opposite the mouth of the Elwha River. The river enters the bay by two mouths and the first kelp plants found were about half a mile west of the west mouth. At this end of the bed the kelp plants are scattering and of medium size. A little farther west the bed becomes quite dense, and continues so to a point near Observatory Point. The bed is not closer to the b:ach than one fourth of a mile at any point. The bed is more than 500 feet wide in places and is over two miles long. There is also a good deal of kelp around Observatory Point. The bladder kelp (Nereocystis Luetkeana) is the only kelp found growing in this bed. Macrocystis pyrifera was found floating in the bay, but not attached. Hydrometer readings to determine the density of the water were made at the most eastern point at which kelp was found in the bay, at the west end of the bed, and at séveral intermediate points. Readings for comparison were made at several points in the open Strait also. A reading was made in the mouth of the river and one at a point about 500 feet directly out from the mouth. All readings were taken at a temperature of 15-5° C., that being the temperature to which the zero point of the instru- ment used was adjusted. The water of the river proved to be 240 RiGG: SALINITY AND THE DISTRIBUTION OF NEREOCYSTIS exactly this temperature, while the samples from the kelp bed and the open Strait varied from 12.5° C. to 13.5° C. There was found to be no difference in the hydrometer readings taken in the open Strait and those taken at any of the points where kelp grew inthe bay. The reading taken in the mouth of the river was zero. The reading taken about 500 feet straight out from the mouth of the river showed about one third as great a salinity as the water of the Strait and of the bay. The facts are, then, that a strong tidal current sweeps freely through this kelp bed, and the water at all points in it shows the normal salinity of the water of the Strait. So far as this bed is concerned kelp does not grow in water that has less than the normal salinity. It does not seem quite possible, however, to say posi- tively that the lack of normal salinity is the only inhibiting factor preventing the growth of kelp near the mouth of the river, al- though it seems to the writer that such is probably the case. Outside of the question of the salinity of the water, there are at least three factors that limit the distribution of Nereocystis in the Puget Sound region. These are rocks for anchorage, a strong tidal current, and proper depth of the water. The strong tidal current is present at the mouth of the river, as it is in the other portions of the bay, and the water is of proper depth for kelp, a little distance off shore, as it is in the other portions of the bay, where kelp does thrive abundantly. It may be possible that the silt brought down by the river has so covered the rocks near the mouth of the river as to render the attachment of kelp impossible. Two sets of samples were collected—one set from the extreme western end of the bed (over three miles from the mouth of the river) and one set from the extreme eastern end (the portion of the bed nearest the mouth of the river). The analysis of these samples is shown in the following table. The above figures should not be construed as indicating posi- tively any characteristic difference in the potash content of the kelp from the two ends of the bed. It will be noted that sample number 3 has a much higher potash content than any other sample reported in the table. Neglecting this one, the samples from the west end of the bed average about one per cent higher in potash than the samples from the east end. This, however, should be a ah R1IGG: SALINITY AND THE DISTRIBUTION OF Nereocystis 241 considered in the light of the fact that two samples of fronds collected by the writer in the Brown’s Island bed near Friday Harbor, Washington, in 1911, and analyzed by Dr. J. W. Turren- tine of the United States Bureau of Soils differed in their potash content by 9.7 per cent.* COMPOSITION OF KELP SAMPLES (Nereocystis Luetkeana) FROM FRESHWATER Bay, PUGET SounpD. Collected by George B. Rigg. Analyses by E. G. Parker and J. R. Lindemuth, U. S. Bureau of Soils. Nitrogen determinations by J. C. Trescott, U. S. Bureau of Chemistry. is Potash, | Iodine,| Soluble Organic | Ash, No. Locality a of | per Nitrogen, | per salts, matter, per plan cent per cent. : ent. | percent. | percent. | cent 1 | From west Fronds | 18.04| 2.26 | 0.23 44.17 51.13 | 4.80 2 end of bay | Fronds | 17.61 22% 24 -70 51.11 | 4.72 3 Stipes 31.62 1.21 o 63-75 33-15 -} 3-46 4 | Fromeast | Fronds | 16.92/ 2.57 0.20 43-37 51.16 | 5-47 5 end of bay Fronds | 17.05 2.71 0.24 | 41.53 51.67 30 6 ronds | 17.32 2.53 | 0.28 | 45.40 50.12 | 4.48 7 Fronds O.90°t <2.54 0.19 42.88 52.19 | 4.908 8 Stipes 16.50 BS 4 0.30 59.24 37-40 | 3-36 9 Stipes 16.72 1.46 0.20 57.06 39.02 3-92 It is quite possible that in some states of tide and current the water may be less saline at the east end of the bed than at the west end, and that this may account for the slightly lower potash content in the samples from the east end. There seems to have been little previous work done on the influence of the varying salinity of water on the growth of the Laminariaceae. Setchellf says that “Exact figures are wanting”’ and that ‘Recourse may be had only to general experience and statements can be couched only in general terms.’’ He says that a few Laminariaceae ‘‘ascend tidal rivers to a slight extent.’ The writer has not found any instance of this in the Puget Sound region, but his investigation particularly of the smaller leaf-like species has not been by any means complete. : Setchell notes the fact that Fucaceae and Ulvaceae are found in tidal rivers. This is true in the Puget Sound region. It is interesting in this connection to note that a Pelvetia plant has even been found growing in a salt marsh in this region. The specimen * Sixty second Congress, Second Session. Senate Document No, 190, p. 220. tT Ibid., p. 135. 242 Ricc: SALINITY AND THE DISTRIBUTION OF NEREOCYSTIS referred to was found by Dr. T. C. Frye in a marsh near Toke Point, Washington. While the writer has not made any detailed investigations in regard to the growth of Laminariaceae near the mouth of any other river than the Elwha, his general observations as well as his conversations and correspondence with other observers lead him to believe that kelps are to be looked for in this region only where the water has practically the normal salinity of sea water. UNIVERSITY OF WASHINGTON, SEATTLE. INDEX TO AMERICAN BOTANICAL LITERATURE (1910-1913) aim of this Index is to include all current botanical literature written by Americans, published in pale or based upon American material ; the word Amer- ica being used in the broadest se Reviews, and papers that sans exclusively to forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and no attempt is made to index the literature of bacteriology. An occasional exception is made in favor of some paper appearing in an American period wholly to botany. Reprints are not mentioned unless they differ from the original in some important particular. If users of the Index will call the attention of the editor ‘to errors or omissions, their kindness will bé appreciated. This Index is reprinted gorgls on cards, and furnished in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitted ; each subscriber must take all pee published during the term of his subscription, Corre- spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club, Ames, O. Orchidaceae novae et criticae insularum Philippinarum. Leaflets Philip. Bot. 5: 1549-1588. 12 D 1912. Forty-two new species are described. Andrews, A. L. Notes on North American Sphagnum. gist 16: 20-24. Mr 1913. Andrews, A. L. Philological aspects of the “Plants of Wineland the Good.”’ Rhodora 15: 28-35. 4 Mr 1913. Bachmann, F. M. The migration of Bacillus amylovorus in the host tissues. Phytopathology 3: 3-14. pl. 2,3 +f. 1, 2. F 1913. Barss, H. P. Cherry gummosis. Biennial Crop, Pest and Horti- cultural Rep. Oregon Agr. Exp. Sta. 1911-1912: 199-217. f. 10-19. 10 Ja 1913. Caused by Pseudomonas Cerasus. Bean, W. J. Prunus pennsylvanica. IV. Bryolo- Curt. Bot. Mag. IV. 9: pl. 8486. Mr 1913. Blewitt, A. E. Notes on Euphorbia Cyparissias L. Rhodora 15: 43. 4 Mri Bédeker, he ‘Manilare Wrightii Engelm. Monats. Kakteenk. 23: 20, 21. 15 F 1913. [Illust.] " Bérgesen, F. Some Chlorophyceae from the Danish West Indies. Bot. Tidssk. 32: 241-273. f. I-17. 25 Je 1912. 243 244 INDEX TO AMERICAN BOTANICAL LITERATURE Britton, N. L. Botanical exploration in Bermuda. Jour. N. Y. Bot. Gard. 13: 189-194. pl. 103-107. D 1912. Cannon, W. A. Some relations between root characters, ground water, and species distribution. Science II. 37: 420-423. 14 Mr 1913. Cockerell, T. D. A. Natural selection. Pop. Sci. Mo. 82: 388-396. Ap 1913. Cook, O. F., & Doyle, C. B. Three new genera of stilt palms (Jriartea- ceae) from Colombia, with a synoptical review of the family. Contr. U.S. Nat. Herb. 16: 225-238. pl. 54-65 +f. 41. 21 F 1913. Acrostigma, Catostigma, and Catoblasius gen. nov. Dixon, H. N. A remarkable form of Dicranella heteromalla Schimp. Bryologist 16: 29, 30. Mr 1913. Reprinted in part from the Journal of Botany for October, 1912. Evans, A. W. Revised list of New England Hepaticae. Rhodora 15: 21-28. 4 Mr 1913. Fernald, M. L. Alnus crispa (Ait.) Pursh, var. mollis (Fernald), n comb. Rhodora 15: 44. 4 Mr 1913. Fernald, M. L., & Wiegand, K. M. The variations of Luzula cam- pestris in North America. Rhodora 15: 38-43. 4 Mr 1913. Freeman, E. M. Harry Marshall Ward. Phytopathology 3: 1, 2. Freeman, G. F. Southwestern beans and teparies. Arizona Agr. Exp. Sta. Bull. 68: 573-619. pl. r-10. 30 Au 1912. Gandara, G. Morfologia de las raices de las plantas. Mem. Soc. Cie. “Antonio Alzate” 30: 7-10. f. I-12. O 1910. Gregory, C. T. A rot of grapes caused by Cryptosporella viticola. Phytopathology 3: 20-23. f. z, 2. F 1913. Grout, A. J. Collecting mosses in Florida. Bryologist 16: 27-29- Mr 1913. Giissow, H. T. Report of the Dominion botanist. Rep. Exp. Farms (Canada) 1912: 191-215. pl. 6,7 +f. 1-4. 1912. Giissow, H. T. Powdery scab of potatoes. Spongospora subterranea (Wallr.) Johns. Phytopathology 3: 18, 19. pl. 4+f.2. F 1913- Harper, R. M. Botanical evidence of the age of certain ox-bow lakes. Science II. 36: 760, 761. 29 N 1912. Hastings, E. G., Evans, A. C., & Hart, E. B. The bacteriology of cheddar cheese. Centralb. Bakt. Zweite Abt. 36: 443-468. f. 7, 2 15 F 1913. Hedgcock, G. G. Notes on some western Uredineae which attack forest trees. II. Phytopathology 3: 15-17. F 1913. Hedrick, U.P. The domestication of American grapes. Pop. Sci. nex : 82: 338-352. Ap 1913. [Illust.] D INDEX TO AMERICAN BOTANICAL LITERATURE 245. Hill, E. J. The annulus of Tortella caespitosa (Schwaegr.) Limpr. Bryologist 16: 17, 18. Mr 1913. Holdt, F. von. Forstwissenschaftliches aus Nordamerica. Mitteil. Deuts. Dendr. Gesells. 1912: 114-123. 1912. [Illust.] Hubbard, F. T. Further notes on the geeiains of Essex County, Massachusetts. Rhodora 15: 36-38. 4 Mri Humphrey, C. J. Winter injury to the white a ecm 3? 62, 63. F 1913. Jackson, H. S. Apple tree anthracnose. Biennial Crop, Pest and Horticultural Rep. Oregon Agr. Exp. Sta. 1911-1912: 178-197. f. 1-9. 10 Ja 1913. Caused by a cei: Neofrabraea malicorticis (Cordley) Jackson, proposed as the type of a new genus. Jewett, H.S. A moss ‘“‘ washing machine.” Bryologist 16: 25-27. f. I. Mr 1913. Johnson, A. G. Canada-thistle and its eradication. Agr. Exp. Sta. Circ. 32: 1-12. f. 1-3. Ja 1912. Kennedy, P. B. Studies in Trifolium—VII. Muhlenbergia 9: 1-29. pl. 1-5. 7 Mr 1913. Includes Trifolium insularum, T. Helleri, T. aioe esa hsp T. Traskae, and T. mercedense spp. nov. Loomis, M. L. Some extensions of local ranges. 4 Mr 1913. Lunell, J. Erigeron in North Dakota. Am. Mid. Nat. 2: 253-258. 1 Jl 1912. Lunell, J. A new Gutierrezia from Oregon. 195. 18 Mr 1912. : Gutierrezia ionensis sp. NOV. Lunell, J. A new Laciniaria from F lorida. 164. 15 Ja 1912. Laciniaria Deamiae sp. nov Lunell, J. New plants from Minnesota—II. 162. 15 Ja 1912 Lunell, J. New pits from North Dakota—VI. Am. Mid. Nat. 2: 142-149. Nuo11r;—VII. Am. Mid. Nat. 2: 153-159 15 Ja 1912; —VIII. Am. Mid. Nat. 2: 185-188. 18 Mr 1912;—X. Am. Mid. Nat. 3: 12,13. 6 Ja 1913. Lunell, J. Some new Laciniariae. Purdue Univ. Rhodora 15: 44. Am. Mid. Nat. 2: 194, Am. Mid. Nat. 2: 163, Am. Mid. Nat. 2: 159- Am. Mid. Nat. 2: 169-177- 18 Mr 1912. Includes Laciniaria Deamii sp. nov Lunell, J. Synonymy alterations. MacClement, W. T. Some conditions Ottawa Nat. 26: 153-160. Mr 1913. Am. Mid. Nat. 2: 264. Jl 1912. of progress in the plant world. 246 INDEX TO AMERICAN BOTANICAL LITERATURE Macoun, J. M. Additions to the flora of Vancouver Island. Ottawa Nat. 26: 160-168. Mr 1913. Continued from p. 149. Maxon, W. R. A new genus of davallioid ferns. Jour. Washington Acad. Sci. 3: 143, are = Mr 1913. Sphenomeris Maxon, gen. Melhus, I.E. Septoria ee in relation to pea blight. Phytopathology 3: 51-58. pl. 6. F 1913. mpi E. D. Notes on Philippine Euphorbiaceae. Philip. Jour. Sci. : shee o, D 1912. : ha ew genus ee and new species in Antidesma (1), Bridelia (1), ROLE 8 (1), Cyclostemon (3), Excoecaria (1), Macaranga (4), Mallotus (7), Ostodes (1), Phyllanthus #6 Sapium (1), Sauropus (1), and Trigonostemon (2). Merrill, E. D. On the identity of Evodia triphylla. Philip. Jour. Sci. 7: (Bot.) 373-378. D 1912. Merrill, E. D. The Pineda monument and the probable site of the first botanic garden in the Philippines. Philip. Jour. Sci. 7: (Bot.) 363-371. pl. 22. D 1912. Meyer, R. Echinocactus texensis Hopff. Monats. Kakteenk. 23+: 28; 29. 15 F 1913. Meyer, R. Uber Echinocactus Pfeifferi Zucc. Monats. Kakteenk. 23: 1g, 20. 15 F 1913. Morse, M. Inorganic colloids and protoplasm. Science II. 37: 423- 425. 14 Mr 1913. Murrill, W. A. Collecting fungi in the Adirondacks. Jour. N. Y. Bot. Gard. 13: 174-178. 23 N 1912. Nieuwland, J. A. Box elders, real and so-called. _ Mid. Nat. 2: 129-142. N 1IgII. Includes Crula gen. nov. Nieuwland, J. A. New plants from various places. Am. Mid. Nat. 2: 178-185. 18 Mr 1912. ; Includes A pocynum glaucum and Persicaria amurensis spp. nov. Nieuwland, J. A. Notes on our local plants. Am. Mid. Nat. 2: 164, 165. 15 Ja 1912;—II. Am. Mid. Nat. 3: 14-22. 6 Ja 1913;— III, Am. Mid. Nat. 3: 41-47. Mr 1913. Nieuwland, J.A. Our amphibious Persicarias. II. Am. Mid. Nat. 2: 201-247. My 1912. Includes Persicaria tanaophylla and P. carictorum Spp. nov. Nieuwland, J. A. A question of nomenclature. Am. Mid. Nat. 2: 258-262. 1 Jl 1912. Nieuwland, J. A. Silene conica in Michigan. Am. Mid. Nat. 2: 264- I Jl 1912. INDEX TO AMERICAN BOTANICAL LITERATURE 247 Nieuwland, J. A. Some local albino plants. Am. Mid. Nat. 2: 265, 266. Jl-1912. Ochoterena, I. Memoria sobre las plantas deserticas mexicanas. Mem. Soc. Cie. ‘Antonio Alzate”’ 30: 171-182. pl. 4-6. Mr 1911. Pennell, F. W. Further notes on the flora of Conowingo or serpentine barrens of southeastern Pennsylvania. Proc. Acad. Nat. Sci. Philadelphia 64: 520-539. 13 F 1913. Porsild, M. P. Vascular plants of west Greenland between 71° and 73° N. Lat. Meddelelser om Grénland. 50: 351-389. f. I-14. 1912. Prain, D., & Hutchinson, J. Notes on some species of Acalypha. Kew Bull. Misc. Inf. 1913: 1-28. f. 1-4. Ja 1913. Quehl, L. aa die Preisverzeichnisse der Kakteen. Monats. Kak- teenk. 23: 18, 19. 15 F 1913. Reed, C. A. Wild flowers of New York. 1-46. Mohonk Lake, N. Y. 1912. [Illust.] Robinson, C.B. Roxburgh’s Hortus Bengalensis. Philip. Jour. Sct. 7: (Bot.) 411-419. D 1912. Rouaix, P. El arbusto llamado ‘“‘hoja-sen” en los estados fronterizos (Flourensia cernua D.C.). Mem. Soc. Cie. ‘Antonio Alzate Yad: 3201-303. Je 1910. Rydberg, P. A. Studies on the Rocky Mountain flora—XXVIII. Bull. Torrey Club 40: 43-74. 18 Mr 1913. Includes Aas eoKs Coriophyllus, Pseudopteryxia and Pseudoreoxis gen. nov., and new species in Lupinus (1), Psoralea (2), Atelophragma (1), Negundo (1), Sphae- valcea (2), Nuttallia gee were (5), Gayophytum Sage Oenothera (1), Glycosma (1), . Pseudopteryxia (1), Cynomarathrum (1), and Cogswell ta Schmid, G. Die Bliite des Cereus pees: S.-D. Monats. Kak- teenk. 23: 29. 15 F 1913. [Illust.] Serner, O. Kreuzungsresultate bei phyllokakteen. Monats. teenk. 23: 17. 15 F 1913. Shear, C. L. Endothia Gasicilt (Schw.). Phytopathology 3: 61. Kak- F 191 Smith, ow Some successful inoculations with the peach crown gall organism. and certain observations upon retarded gall formation. Phytopathology 3: 59, 60. F 1913. Smith, G. M. Tetradesmus, a new four-celled coenobic alga. Bull Torrey Club 40: 75-87. pl. 1. 18 Mr 1913. Tetradesmus wisconsinensis gen. et sp. nov. South, F. W. Some root diseases of permanent crops in the West Indies. West Ind. Bull. 12: 479-498. 18 5S 1912. Spaulding, P. Notes on Cronartium Comptoniae. Phytopathology 3: 62. F 1913. . 248 INDEX TO AMERICAN BOTANICAL LITERATURE Speare, A. T. Fungi parasitic upon insects injurious to sugar cane. Rep. Exp. Sta. Hawaiian Sugar Planters’ Assoc. Bull. 12: 5-62. pl. 1-6 + f.. 15-2.) D 1912. Includes Entomophthora pseudococci and Aspergillus parasiticus spp. nov. Stapf, O. Ruellia Harveyana. Curt. Bot. Mag. IV. 9: pl. 8485. Mr 1913. A plant from Mexico. Stout, A. B. A biological and statistical analysis of the vegetation of a typical wild hay meadow. Trans. Wisconsin Acad. Sci. 17: 408 469. pl. r8-23. O 1912. Vaupel, F. Vier von Ule in Nordbrasilien und Peru gesammelte Kakteen. Notizbl. Kénigl. Bot. Gard. Berlin 5: 283-286. 25 Ja 1913. : Webster, H.S. Grape culture in Pennsylvania. Perinsylvania Dept. Agr. Bull. 217: 7-66. f. I-57. 1912. Includes notes on the fungus diseases of grapes. Weed, C. M. Portraits of American trees, native and naturalized, with a guide to their recognition at any season of the year and notes on their characteristics, distribution and culture. House and Garden Q: 269-275. Je 1906. Weingart, W. Cereus Dismiss S.-D. Monats. Kakteenk. 23: 30. 15 F 1913. Wilcox, E.M. Smuts on Nebraska cereals. Nebraska Agr. Exp. Sta. Bull. 131: 3-16. f. 1-13. 3 Au 1912. Williams, R. S. The genus Husnotiella Cardot. Bryologist 16: 25: Mr 1913. Wolf, F. A. A new Gnomonia on hickory leaves. Ann. Myc. 10: 488- 491. pl. 15. 31 O 1912. Gnomonia Caryae sp. nov. Wollenweber, H. W. Pilzparasitire Welkekrankheiten der Kultur- pflanzen. Ber. Deuts. Bot. Gesells. 31: 17-34. 27 F 1913. Wollenweber, H. W. Studies on the Fusarium problem, Phyto- pathology 3: 24-50. pl.5 +f.12. F 1913. Includes descriptions of Fusarium redolens, F. conglutinans, and F. Sclerotium: spp. nov. Bui. ToRREY j f sy CLuB VOLUME 40, PLATE 9 PHLEBODIUM AUREUM L. BuLL. TORREY CLUB VOLUME 40, PLATE 10 ae ee ep ees ee CYRTOMIUM BOYDIAE (D. C. EATON) W. J. ROBINSON BuLL. TORREY CLUB VOLUME 40, PLATE II bE en re SADLERIA HILLEBRANDII W. J- ROBINSON BULL. ? IE L. TorRrY CLUB VOLUME 40, PLATE 12 PLANTS OF THE HAWAI! EQTED ON THE 1SLANt UAH, ON KAHOLUAMANOS, ASOVE wamera Gr A A Mejia, Semremnen 24-90, U6 SADLERIA UNISORA (BAKER) W. J. ROBINSON VOLUME 40, PLATE 13 Buy. Torrey CLus wéby ane, pees PICKETT: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM VOLUME 40, PLATE 14 BuL_. TorrReEY CLuB AAANATINNIEDcavanabeg Oh li\ann ihn Co SN FRA UIS x Ks + a ee Sn UN a ie oe a ere, 78 he SD) ) See ee Sl neo ss if v the See < } = ca Ny ee t zs a ee os se Ee it ee C5 Tago fou : ibe "ans of op 5. GS an OA vere GUNERDCEAIG HOTT | WN Lait TTY ‘ 1 ’ 4 A ye Sg “, I ‘ ! er Bh PICKETT: EMBRYO-SAC OF ARISAEMA TRIPHYLLUM Vols 40 See No. 6 BULLETIN OF THE TORREY BOTANICAL CLUB ee JUNE, 1913 Four hybrids of Viola pedatifida Ezra BRAINERD (WITH PLATES 15-17) The hybrids between Viola pedatifida and allied species are in several respects the most interesting among the 75 or 80 that have appeared in the genus as represented in North America. At least two of the four are remarkably hardy, almost immune from attacks of fungus, and comparatively fertile; they are there- fore well suited for experimental cultures. The marked contrast in leaf outline displayed in the parents of the several crosses affords a fine opportunity for studying in detail the many diverse forms of leaf that emerge in the offspring of the hybrid. The other opposed parental characters, relating to pubescence, color of capsule, color of seed, length of peduncle, and villosity of spur petal, also lead to results well worthy of careful study. One of these hybrid plants I have had in hand for eight seasons and have raised from it over 450 offspring, extending through four generations. It was discovered at Yorkville, Ill., May 1905, by Miss Mary O. Pollard, a former pupil, and may be briefly described as follows: 1. Viola papilionacea X pedatifida hyb. nov. Rootstock stout, at length extensively branching horizontally; leaves broadly deltoid-ovate in outline, cleft into 7-11 linear or oblong lobes, the middle lobe much the widest, glabrous, though the margins are often scabro-ciliolate; petals violet, the odd one more or less villous; cleistogamous flowers on erect or ascending peduncles, intermediate in length to those of the parent species; capsules 8-12 mm. long, infertile, averaging in 12 capsules 8 249 250 BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA seeds; seeds brown, 2 mm. long; offspring notably diversiform. (PLATE 15, Aa.) Several sowings have been made of the seeds of this hybrid; but the ten plants of the first brood (236), whose offspring have been raised for two succeeding generations, are the only ones that will be here discussed. The forms of leaf found in the ten plants and in their respective broods of offspring are presented in the first half of TABLE I. The second half presents the corresponding facts regarding sixteen offspring of brood 236, plant 3, whose leaf had a hybrid form, quite the same as that of the mother plant from Illinois. TABLE I CULTURES OF VIOLA PAPILIONACEA XPEDATIFIDA FOR FOUR GENERATIONS V. papili h lan f ee ni She SERS Aart Pie As Sixteen Fs; offspring of 236-3 Their wu. L leaf cleft: ¥ di pe oie Ra Sait 1a ne 0 a Their offspring—Fy Ten Fs offspring of hybrid Brood | Leaf Number and | | Their offspring—F3 468 form Brad | form | Total B inprrses Ra ARSE ease Pe teh fist ar eaeae ai tg a I Nccndbpiy and form| ! A |Aa| a Brood. |... Sees ps } | A Aa| a No. 1} A 60 12 | *22 | : - No. 2) Aa 606 Bt Sil bees No. 2 A 595° 30 30 INo. 3| Aa | 607 | 4| 6| 3 13 No. 3, Aa ee ae 3} s2 [No 4) 4 Speen um eee. | 59 8|20| 8 No. 5 a 609 512785 No. 4) Aa boy gan eae ots tee 10 |No. 6] A 610 | 20 20 No. 6| @ | 508 | 16 | 16 INo. 7} a | Grr | [xx] a No. 7 A 599 16 16 |No. 8] A 612 14 4 No. 8 Aa 600 | 1|10| 3 14 |No. 9} @ | 613 16 16 No. 9 Aa O08 fo 2g INO. 11; Aa | 6x5 | 3} 6) 3) 32 No. 10) Aa | 602 | 6 3. | 45 |No. 12} Aa | 616 | 3 7| 3| 13 No. 11} A 603 | 13 13 |No. 13 GETZ | 32 | 12 No. 12) Aa 604 | 3] 5| 2 to |No. 14| A 618 |12 | 12 0. 15 a r2|° 12 Foo ai 199181 33 | 389 WN 6) 48 | Geo 2] 7] 4) | 105 _|No. 17) @ } 621 | oud Roe rio tA eta t | 86 | 216 Si ee From Aa’s.../ 15 |31|18| 64 The cultures show that in this hybrid the Mendelian law controls in a general way the inheritance of leaf form, though there is no dominance, the hybrid leaf being intermediate between the leaves of the two parents. The plants that resemble V. pedatifida in having parted leaves always produce offspring with parted leaves; those that resemble V. papilionacea in having uncut leaves always produce offspring with uncut leaves; while those BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA 251 that have the compromise leaf reproduce plants with three forms of leaf incision, as did the original hybrid. Also in the relative number of these forms there is an approximation to the Mendelian ratio 1 :2:1. In the above 26 plants (broods 236 and 468), whose forms were verified by their offspring, there are 9 A’s, 11 Aa’s, 6 a’s, the theoretical ratio being: 63 A’s, 13 Aa’s, 63 a’s. In the 169 offspring of Aa plants, given above for the third and fourth generations, there are 44 A’s, 84 Aa’s, 41 a’s, the theoretical ratio being: 42} A’s, 844 Aa’s, 42} a’s. Here we find, as usual, that the larger the number of individuals the closer the normal ratio is realized. But besides general conformity there are also departures from the strict Mendelian law. For one thing the hybrid or inter- mediate leaf varies in different individuals, inclining now more to the form of the one parent species and now more to the form of the other. Also the reversionary forms, designated as A and a, are rarely complete reversions. The A plants, though stable in producing like parted leaves in succeeding generations, do not have leaves as deeply parted as in V. pedatifida; and the a plants, though plainly uncut and stable, usually have teeth noticeably longer than in normal V. papilionacea, sometimes even pectinate. (PLATE 15, FIG. a.) Another cause often conspires to increase these differences in leaf pattern: the presence of minor hybrid characters that inde- pendently adjust their special conflicts of hybridity. For example, the leaf of V. pedatifida is usually truncate or even cuneate at the base, that of V. papilionacea usually cordate. A hybrid offspring may inherit the broad truncate base of the former with the uncut margin of the latter. Sometimes in the hybrid leaf the lobes are entire, and obtuse at the tip; sometimes, as in the normal leaf of V. pedatifida, the lobes are again cleft or toothed on the outer margin, and acute at the tip. In these various ways there han arisen in the numerous progeny of the hybrid under discussion a considerable diversity of foliage, such as would present insoluble difficulties to a taxonomic student, who did not know that these diverse forms all came from one individual, by close-fertilized reproduction, in the short period of three or four years. The extreme differences are such as would 252 BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA warrant the making of several distinct species, according to the hasty methods of ordinary practice. The hybrid V. papilionacea X pedatifida seems not to be rare in the Middle West. I cite a few interesting examples: M. A. Castleton 25, Vinita, Okla., April 18, 1891; distributed as “ V. palmata.”’* From the United States National Herbarium in 1911 was distributed with printed ticket: ‘‘ Viola Bernardi Greene, Freeport, Ill., Charles F. Johnson, May 15, 1899; determined by Dr. E. L. Greene and Philip Dowell.’’ The plant is quite the same as the hybrid under discussion from Yorkville, Ill., and seems to represent Dr. Greene’s present conception of his species. V. indivisa Greene is also a derivative form of this hybrid; the ‘type’ from Prairie Junction, Minn., E. L. Greene coll., July 7, 1898 (Pittonia 5: 124. pl. 13. 1903). Also along railway, Naper- ville, Ill., Z. M. Umbach, May 18, 1897. (Cf. Leaflets 1: 182. 1906.) 2. Viola pedatifida X sagittata hyb. nov. Plant becoming cespitose, the rootstock dividing into several erect branches; leaves that develop after petaliferous flowering finely pubescent especially beneath and on the upper portion of the petiole, the blades subcordate-ovate in outline (the width about 3 the length), cleft into 6-8 oblong-linear lateral lobes and a broad slightly toothed terminal lobe, the leaves of late summer relatively broader; petals violet, the three lower villous; apetalous flowers and fruit on erect peduncles as long as the petioles; auricles of sepals long and divergent; capsules green, 6-10 mm. long, often quite infertile; seeds intermediate to those of the two parent species in size and color; offspring much unlike each other in - foliage, but blades always incised or coarsely toothed toward the base. (PLATE 16, FIG. Aa.) This hybrid first attracted my attention in a parcel of violet specimens collected in central Illinois by Mr. V. H. Chase, and sent me in November 1907 for determination. It was found in undisturbed prairie soil along the right of way of the Rock Island and Peoria Railroad, just north of the south boundary of Stark County. At the same place and time were collected V. pedatifida and pubescent V. sagitiata, the three plants bearing the conse- cutive numbers 1356-7-8. The anomalous plant impressed me as * Regarded as V. viarum by Mr. Pollard, and apparently the basis for accrediting this species to Ind. Terr. in Britton’s Manual, p. 636. 1905. BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA 253 distinct from V. pedatifida X sororia, discussed below, and as a cross between the two species with which it grew. Mr. Chase, to whom I appealed for living plants, found that the station had been recently burned over; but the following May he discovered another colony along the railway a half mile farther south ( V. H. Chase 1619). The stocky specimen sent was easily divided, and six or eight vigorous plants were obtained during the season of 1908. Mr. Chase reported that the pubescent V. sagittata ‘ was very abundant, thousands of plants cover the ground with a blue carpet, mostly where the land was a little low and damp. _ V. pedatifida seemed to prefer rather drier ground. The hybrid was invariably with V. pedatifida, on fairly dry soil; and V. sagittata was never more than a few rods away.” During the season of 1909 I grew nineteen offspring of Chase 1619, and they gave abundant evidence as to the taxonomic status of the mother plant. Leaves of nine of these offspring are figured in PLATE 16 and indicate something of the marked diversity of form resulting from the combination, in the leaf of the original hybrid, of at least four pairs of opposed characters,* that blend or segregate, independently and variously, in the several offspring. 3. Viola pedatifida < sororia hyb. nov. Becoming cespitose with multicipital caudex; leaves that expand at petaliferous flowering 9-13-cleft, the lateral lobes broadly linear, usually with one or two coarse teeth on the outer edge toward the apex, the middle lobe much broader and incised on either side, the upper face somewhat hirtellous, the lower surface and the petioles villous; the leaves of summer larger and less deeply cleft; apetalous flowers on rather short, erect or ascend- bearing 5-20 brown seeds 2 mm. long; offspring markedly dis- similar. Not rare on prairies of the Middle West. (PLATE 17.) I am greatly indebted to the kindness and skill of Mr. Chase for the abundant and excellent material used in the study of this hybrid. Collectors in this region-know that the native flora of the open prairies is now largely restricted to untilled strips of land * These are: V. pedatifida V. sagittata T. Outline - broadly flabelliform lanceolate 2. Form of base truncate or cuneate cordate or subcordate coarsely toothed at base 3. Incision 2-3-ternatel finely pubescent y dissected 4. Pubescence margins and veins hirtellous 254 BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA along the borders of railways. On May 16, 1909, Mr. Chase, whose bicycle was adjusted to run on rails, traversed in 5 hours the 24 miles between his home at Wady Petra and the town of Galva. In order to make the return by train, he says, ‘I could not stop to hunt along the way; but whenever I saw a cut-leaved violet that was not V. pedatifida I stopped for it.”” The twelve numbers of living plants collected on this trip reached me safely, and all have flourished in the Vermont garden. The status of hybrid plants in the wild is well shown by a detailed study of these specimens and of their offspring; for all but two sterile plants have been reproduced by seed. The main points regarding them, that have a bearing on the present problem of hybridism, are presented in the following tabular synopsis (TABLE II): TABLE II VIOLET PLANTS COLLECTED BY V. H. CHASE IN CENTRAL ee May Sig 1999) bial Pig ie pend h noe oe a pares characters Soe "Seed color A paoer V. pedatifida D { 1951 re ie | 7 pedatif = 1956 | (parted) |(glabrous) ioeem | ales V. sororia Willd... I2Q1 | B fD 66 educn) (villous) | hte (brown) V. papilionacea Ph. 1958 a D 66 | 1950 Aa ' sterile o V. he psc 1952 . b | Dd 7 sicsanaes 1949 ae ie BARE Bie 93 1947 Aa b A . tee 1957 Aa Bb | sterile 0 V. pedatifida 1955 A ras Ce 4 Da 10} X sororia 1953 Aa c pias Br 1954 Aa Ce Dd 8 3 5008 AB. | gine Dd 18 Nos. 1951 and 1956 are V. pedatifida; and I have added to the list, though not collected May 16, 1909, V. sororia, the other parent of the hybrid under discussion; specimens of this had been previously sent me by Mr. Chase from four stations in his vicinity. No. 1958 is the prairie form of V. papilionacea, named V. pratincola by Dr. Greene (Pittonia 4: 64. Ji 1899). These three are common and widespread species of the Prairie States from Canada to Texas; the first and third reach westward to the mountains of Colorado. Four of the numbers are the hybrid V. papilionacea X peda- BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA = 255 tifida, already discussed in this paper. Of these no. 1950 flowers freely in May with showy flowers, that often appear also in July and August; but apetalous flowers are rare, and neither sort has been found to produce seed. The three. others are also nearly sterile, bearing only 5-10 seeds to a capsule; but none of the three turns out to be like the Pollard plant from Yorkville in being a first cross (or F,), the form to be selected as the starting point for experiments on the laws of inheritance in hybrid offspring. No. 1952 has an uncut leaf as in V. papilionacea and a green colored capsule as in V. pedatifida, both recessive or reversionary characters never found in a first cross. No. 1949, on the other hand, has the hybrid leaf and the hybrid capsule color but the buff seeds of V. pedatifida and is therefore another subhybrid. No. 1947 consists of five plants, the seeds of which differ in color, and the leaves of which, though somewhat incised, display at least three unlike patterns; the five plants therefore must be considered the offspring of an earlier hybrid. The remaining plants of the 1909 collection are equally variant formas but of V. pedatifida X sororia. No. 1957 is of dwarf habit and has the compromise or hybrid leaf; but though vigorous and multiplied by division into eleven plants, it has failed to yield a single seed. No. 1955 (six plants by division) has a leaf more deeply cut that the others, and this style of leaf reappears in all its offspring. In this we see a reversion, far from complete but stable, to the leaf of V. pedatifida. No. 1953 (again six plants by division) has the pure green capsules of V. pedatifida, as have also its offspring, and so is another subhybrid. But the two remaining numbers, 1954 and 1948, seem to be the desired first product of hybridism, all the four pairs of opposed characters in the double parentage appearing in a compromise form in both numbers. A flowering specimen of 1954 was distributed in my “ Violets of eastern North America, 1910,” no. 121; and in no. 122 are shown two sister offspring, one with the uncut leaves of V. sororia, the other with the parted leaves of V. pedatifida.* No. * In one of our large herbaria, where the work of mounting is done by novices, these two offspring were considered too unlike to appear on the secs sheet. : the plant with uncut leaves was mounted over ticket 122; while the sister plant with parted leaves was placed on the sheet with no. 121, which indeed it more closely resembled. That two plants so dissimilar should come from one self-fertilized parent has seemed incredible even to certain “ . 256 BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA 1948, being unusually fertile for a hybrid, was chosen for the basis of a somewhat detailed study of the reproductive behavior of a tetrahybrid. During the season of 1910 twenty-one plants were grown from the seeds of Chase 1948. In August nearly all bore cleistogamous flowers, which matured several capsules of seeds. These were sown about December 1 in shallow boxes and placed in a cold frame with no protection from the winter weather but a covering of burlap. In the spring of 1911 all but seven of these sowings gave broods of F3 offspring, containing each 6-18 plants. These have been carefully observed for two seasons and the characters of each plant noted as respects leaf incision, pubescence, color of capsule, and color of seed, four qualities in which the parent species were opposed. In each of these four qualities the plant resembled either V. pedatifida, V. sororia, or their hybrid; and in most instances the data were at hand, and clear enough, to determine at once this resemblance by inspection. In the case of the sixteen F, plants of brood 781, here made use of, the characters were verified by the behavior of their offspring, the reversionary forms always proving stable, the hybrid forms always unstable. The details of this experiment are given in TABLE II, in which the symbols Aa, Bb, Cc, Dd denote the blend or hybrid character. The statements made above regarding the marked diversity of leaf pattern in the offspring of V. papilionacea X pedatifida and the departures from strict Mendelian law are equally true of the analogous hybrid V. pedatifida X sororia. The imperfect rever- sions in leaf form are shownin PLATE 17,¥IG. A.B, A.Bb, and A.b, compared with the leaf of V. pedatifida figured above them. But in this hybrid the same phenomena are observable also in the varying colors of capsule and of seed. In all three pairs of char- acters the stable reversions marked A, C, and D are not complete reversions. The darkest capsule or seed found in the Fs brood is much lighter than the capsule or seed of V. sororia. It is further to be observed that though the Mendelian law leads us to expect on the average one of each of these reversions in every four offspring, we have here only one of each in the sixteen offspring. At the same time the hybrid forms are in excess of the BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA ALY | normal average (one half of the whole number); instead of 8 of each we have 10 Aa’s, 12 Cc’s, and 10 Dd’s. And it should be remembered that this statement is not based solely on the appear- ance of the F, plant but on the fact that the reversionary charac- ters, A, C, D, were found to be stable in reproduction; while the hybrid characters were found to be unstable. For example, with the exception of no. 4, which had only two offspring, each of the ten Aa plants in brood 781 gave 2-7 plants with uncut leaves. TABLE III VIOLA PEDATIFIDA XSORORIA, CHASE 1948, AND ITS OFFSPRING HybridF, | 4a | 36 | a@ | De Forms of sixteen F,, offspring Fs offspring of brood 781 Brood Foliage ‘Capsule | Seed | Brood Size Exhybrids SSE ee es ere re | | | | | | a.B.C.d FST nO. Fes) a | Ba Ge a4 d 853 13 plants 8{5 ica 2 a.B.C.d FBE NOV. 43 | a | B Ce | d 855 ro plants i. eBid 781 no. 4 | AG Bb Cece) Dad" 4856 2 plants 0 | | | (1 A.Bicd WEY ho. Gila ee Cc. Da 858 16 plants | 3,1A.b.C.d | | |” [a Abed 78I no. 7 Aa | b Ce Dd 859 7 plants | o : BAC. 781no. 8 Aa Cc d | 860 6 plants | 4 13 “B.C d 1 A.B.C.D 344 A-B.c.D : 1 a.B.c. 781 no. Io Aa b Ce) Dd 862 14 plants a.b.c | | 7S NO. Go) As B | &e D 861 16 plants | c | Dd 863 rr plants | | | I 2{ 781 no. 13 a Bb Coo e 865 2 plants o <> | a.B.c. 78rno.14 | a | B c Dad | 866 | 9 plants | 6 i? ge 781 no.15 | Aa’ | Bb | C..| d | 867 | 15 plants [1 © 4.B.C.d 78Ino.16 | Aa Bb Cee 868 10 plants ° oe r a.B, 781 no.17 | Aa Bb | Ce ie a 869 18 plants | 2 { = eoheg 81 no. b | Ce | Dd | 87 8plants (1 a.b.c.d ER hire me | (1 AbCd 781 no.20 Aa b | Cc | Dd | 872 | 14 plants | 311 A.bc.d ; 1 a.b.c.d | ae ae eer _ And not only in the second but also in the third generation of this hybrid the number of plants having the positive character seems to fall short of the Mendelian requirements. This appears if 258 BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA we note the ratio in which the several hybrid characters in brood 781 segregate in the F; offspring: The 10 Aa plants had 113 offspring: 23 A’s, 54 Aa’s, 36 a’s 20:48 :32 The 7 Bb plants had 74 offspring: 14 B's, 35 Bb’s, 25 b’s Ss Ss 10:47:34 The 12 Ce plants had 130 offspring: 23 C’s, 67 Cc’s, 40 c’s ratio in ‘% 18:51:31 The 10 Dd plants had 93 offspring: rg D’s, 44 Dad's, 36 d's 14:47:39 Total 410 73 normal ratio: 25:50:25 Combining these results, we find in 410 instances of the reproduc- tion of a hybrid character, that instead of 1024 reversions to the positive character there are only 73; instead of 2 5 per cent, only 18. However, the determination of the characters in these 171 F; offspring rests only upon their appearance, having not been as yet verified by observing the behavior of the F, offspring. Of this generation 45-50 broods are hoped for by another season from seed already sown. But we seem to be already justified in the suspicion that in a species-hybrid as complex as the one under experiment, where the opposed characters of the parents appear in a blend or intermediate form, this form may acquire a certain degree of fixity, whereby the reversions to the positive type in a pair of opposed characters are less complete and less frequent than in normal Mendelian segregation and the reversions to the negative type more frequent. If we might assume that the gametes holding the positive character were more or less impure, while the gametes holding the negative character were pure, the situation would be fairly well accounted for. In the great diversity of forms displayed in the 171 F; offspring the most interesting group are the 41 exhybrids, in which all of the four characters under study are reversionary and constant. Among these we find three plants in which the four characters of V. pedatifida, A.b.c.d, reappear, one plant in each of the broods 858, 863, and 872. What may be called a form of V. pedatifida with uncut leaves, a.b.c.d, is found once in each of the broods 863, 871, 872. A pubescent V. pedatifida, A.B.c.d, occurs in brood 858. A purple-capsuled V. pedatifida, A.b. C.d, occurs in both 858 and 872. Similarly, we have a cut-leaved V. sororia, A.B.C.D, in brood 861; a green-capsuled V. sororia, a.B.c.D, once in brood 861 and three times in brood 866; and a buff-seeded V. sororia, a.B.C.d, in broods 853, 855, 860, 867, and 869, eleven plants in all. In short, all but five of the sixteen possible combi- BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA 259 nations in fours, of these eight pure elementary characters, are to be found in these 41 plants. As showing how a hybrid tends in successive generations to eliminate its hybrid characters, ever becoming simpler and finally pure, we note that starting with the F, plants, necessarily hybrid in all the opposed characters of the two parent species, we have in the next generation only two such hybrids, and in the 171 plants of the 2d generation, none; indeed, the law of probability calls here for only one in every 256 offspring. _ My first recognition of V. pedatifida X sororia was in a package of living plants sent May 22, 1907, by Dr. H. V. Ogden of Mil- waukee, collected at Upper Nemahbin Lake, Wis., growing with both parents, and considered by him as ‘‘ doubtless a hybrid.”’ The same thing, however, had been sent me some three years earlier by Dr. Greene as a specimen of his V. Bernardi, collected by himself at Dixon, IIl., June 18, 1898. In Leaflets 1: 184. Ja 1906, the plant is transferred to V. perpensa, then first de- scribed. I have recently examined the three other specimens there cited and regard them all as forms of V. pedatifida X sororia.* Also V. fallacissima Greene, Leaflets 1: 185. Ja 1906, is another form of the same hybrid from western Missouri— Bush 141, Lee’s Summit, Mo., July 8, 1899. Other specimens are: E. J. Palmer 3345 and 3393, Webb City, Mo., April 19 and May 5, 1911; Mary O. Pollard 6, Yorkville, Ill., May 16, 1909; L. M. Um- bach, prairies, Clarendon Hills, Ill., June 21, 1899—distributed in 1911 from United States National Herbarium as “‘ V. palmata.”’ 4. Viola nephrophylla X pedatifida hyb. nov. V. Wilmattae Pollard, Proc. Biol. Soc. Wash. 15:178. Au 1902T Foliage much as in V. papilionacea X pedatifida, nearly glabrous, palmatifid with several narrow lateral lobes; corolla ‘deep violet, 2 cm. broad ”’; petals markedly villous and sepals with slightly scarious margins, as in V. nephrophylla. It has not been practicable to secure living plants for cultures. But the alleged parent species were both growing in the cafion, * I therefore wish to correct my statement in Bull. Torrey Club 37: 584. 1910 that V. perpensa Greene is the western form of V. palmata—a too hasty inference from the fact that a V. palmata specimen in my herbarium from Aurora, Ill., was named V. Bernardi by Mr. C. L. Pollard. : pe is 404,924 United States National Herbarium, Mrs. Wilmatte P. { The ty Cockerell, Sapello Cafion (c. 8000 ft. alt.), Beulah, New Mex., May 5, 1901. 260 BRAINERD: FOUR HYBRIDS OF VIOLA PEDATIFIDA and the status of the plant is strikingly analogous to that of numbers 1 and 3 above described. The supposed hybrid has been recently again collected, by Mr. Paul C. Standley.* The leaves in his specimens are cut about halfway to the midrib; but on the same sheet are two detached leaves parted as in V. pedati- jida, indicating that this species grew with the anomalous plant. MIDPLEBURY, VERMONT Explanation of plates PLATE 1 . Figures 3 natural size Aa. Viola papilionacea X pedatifida Brainerd, transplanted, May 1905, from York- ville, Ill., Mary O. Pollard coll.; ex horto Middlebury, Vt., May 14, 1910. 9. 109 Brainerd’s violets of eastern No. Am., Ig10. A. Fs offspring of Aa, having stable leaf pattern resembling that of V. pedatifida; ex horto (brood 612) Middlebury, Vt., June 2, 1912. a. F:2 offspring of Aa, having stable leaf pattern resembling that of V. papilionacea; ex horto (brood 236 plant 6) Middlebury, Vt., June 3, Iga9. PLATE 2 Figures 4 natural size A. Leaf of Viola pedatifida Don, south of Wady Petra, Stark Co., Ill., V. H. Chase 1356, May 26, 1907. a. Leaf of pubescent V. sagittata Ait., south of Wady Petra, Ill., V. H. Chase 1524, July 28, 1907. Aa. Viola pedatifida X sagittata Brainerd, transplanted, May 3, 1908, from along railway 1} miles south of Wady Petra, Ill., V. H. Chase I610; ex horto Middle- bury, Vt., Sept. 7, 1900. I~13. Characteristic leaves of nine F» offspring of above hybrid; ex horto (brood 630) October 1909. Figures } natural size A leaf of Viola pedatifida Don, transplanted, May 1909, from near Galva, Ill. V. H. Chase 1951; ex horto (brood 851) Oct. 3, rorr. A leat of V. sororia Willd., transplanted, May 1909, from Yorkville, Il., Mary O. Pollard coll.; ex horto (brood 707) Aug. 31, 1910. Hybrid leaf of V. pedatifida Xsororia Brainerd, transplanted, May 1909, from along railway, Stark Co., Ill., V. H. Chase .B.—a Aa.B, F2, plant 9 of brood 781, stable as to pubescent leaf only a.B, F2, plant 14 of brood 781, stable as to pubescent uncut leaf. A.Bb,F3, plant 8 of brood 868, stable as to palmatifid leaf only. Aa.Bb,¥ 2, plant 17 of brood 781, hybrid in bott pul df @.Bb,F3, plant 8 of brood 867, stable as to uncut leaf only. A.b, Fs plant q of brood 995, stable as to glabrous palmatifid leaf. Aa.b, F2, plant 7 of brood 781, stable as to glabrous leaf only. 4.b, Fz, plant 0 of brood 998, stable as to glabrous uncut leaf. * United States National Herbarium, Standley 6072, near the Sierra Grande 2100-2925 m. alt.), Union Co., New Mex., June 18, IOI. Viola obliqua Hill and other violets EUGENE P. BICKNELL Viola obliqua—the name is become anathema! Venerable indeed, yet ffom of old misunderstood even by those who have sought to do it honor, rejected, reinstated, and at last altogether cast out, it may be deeméd a matter for apology that it should be now once again brought forward. Nevertheless I ask a further hearing in its behalf—the fraternity of violarians must be my judge. The name seems first to have emerged into the modern light in the Illustrated Flora. If recollection ke not at fault, I myself had some part in this. And the view then shared with the author of that work, Hill's illustration before us, that this discredited name was perfectly available for exact use, has not suffered any change. I have not turned to Hill’s much ridiculed plate from that day until this writing, nor do I suppose that Doctor Britton has, yet, viewing it together now we are at agreement as before. More redoubtably than any other writer, more picturesquely, Doctor Greene has used his slings and arrows against this name.* Yet, as his page presently allows us to see, with friendly purpose! His onslaught—assuredly not to be withstood—finally by a hairs- breadth evades a fatal issue. With fine dexterity the all but destroyed thing has been rescued and, on the instant, sent forth with now well-established rights—for how shall it ever again be assailed with better success? Yet somewhere wasa miscalculation. Later writers, and there have appeared not a few, have.approved the name as extinct, perhaps not stopping to apprehend this reinstate- ment or the dryness of Doctor Greene’s closing avowal ‘‘ the most common of all East American violets . . . I am confident it can never be proven that it is not Viola obliqua Hill.” Doctor Greene has made his hypothetical objector say of Hill’s plate that ‘‘ it does not half represent any violet that ever grew in any country. It is glaringly false in representing flowers erect on * Pittonia 3: 142-143. 1896. > 261 262 BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS peduncles perfectly straight to the very summit.” Thus is the overtechnical critic at first gaily allowed his fling. But this imagined sceptic need have made less allowance for the “ un- botanical draughtsman ” than it is implied he should have done. Had Doctor Greene’s: keen eye ever scanned the woodland floor among the hills along the Hudson, or on Long Island, it must have seen in apparition before it this same pictured violet of Hill’s side by side with its living counterpart having flowers postured in the self same upwardly oblique way and even strictly erect! Here then was the living vindication of Viola obliqua Hill and Sir John had waited nearly a century and a half to be put in credit. The surprising thing is that his figure ever should have come under any doubt. It utters authenticity. Its entire composition proclaims that it could have been in no part extemporized. Be- yond peradventure we see in it the copy of an actual plant worked over by a conscientious but not a facile draughtsman. No inattentive sketch, no artist’s fiction, would have been cumbered with the needless and inartistic detail shown in this cut nor, like it, reveal the painstaking effort of a careful but none too practised hand. Nor, as to the flowers should it have been forgotten that Hill put in print, and he had the living plant, that they were “ oblique,”’ and Aiton that they were “ erect.’”’ I do not know whether this violet remained in cultivation in England up to the time when Aiton wrote or whether he had ever seen it in growth. But if his description was drawn up from herbarium specimens having the petals partly discolored from drying it offers an explanation of his use of the word straminea in giving the color of the flowers. By fault of this word, nothing else can explain it, the history of the blue-flowered Viola obliqua has come confusingly in touch with violets so remotely related to it as the white- or creamy-flowered Viola blanda and the yellow-flowered Viola rotundifolia, As for our plant which so perfectly upholds Hill’s illustration I doubt not that ‘t may be found bearing its upwardly looking flowers over a far wider range than where I myself have seen it growing, for it is none other than the common violet we have been taught to call Viola afinis LeConte. The Illustrated Flora was therefore right in restoring the name Viola obliqua, although BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS 263 not drawing the specific lines so closely as we may now do; Doctor Greene was right in supporting the name with his endorsement; Mr. Pollard in Britton’s Manual was exactly right in allowing Hill’s name to displace LeConte’s. Notwithstanding all this the name Viola obliqua will be searched for in vain in the violet writings of the day. The plant figured by Hill was not that extreme form of the species of slighter figure and narrower leaf that we seem to have set up as typical of Viola affinis. But it was of the same flexuous habit, the same grouping of foliage and flower, the same deep cordation and acuteness of the expanded leaf; the blades had the same pronouncedly crenate-serrate marginal pattern, and some of them were quite sufficiently narrow and attenuate to satisfy the most exacting affinis standard. It is no objection that other leaves were broadly ovate. Forms of the species, very usual forms, do produce just such broadly ovate leaves and in like way associated on the same plant with the more characteristic nar- rower ones. Such plants, and others much more strongly grown than the medium plant portrayed by Hill, are not possibly to be kept distinct by a name from the most reduced and delicate forms of the series, for the extreme phases are everywhere inextricably blended together through every avenue of intergradation. It must not be inferred that there is a particular form of Viola ‘obliqua in which the flowers at some stage of their growth always become upturned. This trait of the flower is no more than a ~ tendency in the general species which, in some plants, or colonies of plants, may be perfectly realized, while in others it is not seen at all - or proceeds no further than an opening out of the crooked tip of the peduncle causing the flower to stand away from the scapeina horizontal or an upwardly oblique position just asit may in many another violet. Noris the erect position of the flower at all extra- ordinary among violets. It is occasionally seen in other species although in no other known to me does it come to a well-established trait, and in no other than this, except rarely, have I seen the flowers strictly ‘‘ erect or peduncles perfectly straight to the very summit.” Just as our medium plant passes down into its smaller and more delicate forms so also, and with as gradual transformation, does it grow up into an every way larger violet having thicker 964 BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS leaves, obtuse and of less pronounced crenation, and more striking flowers of deeper hue and bluer tone of color. Of late years much has been made of this larger violet, and forms which cluster about it, under the designation Viola papilionacea Pursh. Let us digress upon this rather ostentatious newcomer among our named violets. For myself I have never quite succeeded in finding out what was the touchstone of ‘‘ Viola papilionacea.” Nor does there appear to be perfect accord among its sponsors as to its exact credentials. Mr. Pollard in first taking up the name* introduced us to a wholly glabrous plant, describing accurately the fine violet to which we have just adverted. But the specimens he put out} were at some discord with his description, showing us a violet having a characteristic pubescence on the petioles. Inrespect of this pubescence the specimens coincide with Doctor Greene’s understanding of ‘‘ papilionacea’’t and with the admirable drawing of Mr. Holm which supplements his description. Mr. Stone, both by description and illustration, reports the plant as bearing pubescence on the petioles.§ Mr. House as well.||_ Doc- tor Brainerd, on the other hand, although taking a broader treat- ment, seems more in accord with Mr. Pollard in his description of a wholly glabrous plant** as also is Doctor Dowell}}—something like an even division between the smooths and the roughs. All this is not making too much of a little pubescence, for the glabrous and the pubescent plants differ by far more than this one character. And the strain of discrepancy running through the discussions of * papilionacea ”’ is reason enough why this conjectural species has not been received by all of us with any such compelling sense of recognition as, for instance, all felt towards “ Viola cucullata’”’ - the instant that Doctor Greene gave us the cue. The glabrous ‘‘ papilionacea”’ is found on grassy banks or at the borders of meadows along descending places from woodlands ar a Re ee = iat aentrg 26: 136. 1 Tt North Am. Violaceae. Determined rat distributed by Prof. Edward L. Greene ee Mr. Charles Louis Pollard, No. } Pittonia 4: 140-141. 1900. § Proc. Acad. Nat. Sci. Phila. 55: 670-671. || Bull. Torrey Club 32: 258. 1905. ** Rhodora 6: 15. 1904; Bull. Torrey Clu’ 3 :500. Ioto. tf Bull. Torrey Club 37: 164. 1910. 1904. BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS 265 where Viola obliqua grows. In these near meadows or by brooks that traverse the same woodlands ‘ Viola cucullata”’ raises its long-stemmed flowers of twofold blue. In many ways the smooth “ papilionacea’’ is strikingly intermediate between these two plants. Doctor Brainerd has opened our eyes to violet hybrids. Have we not found one here? I recall all these violets as they grew in profusion about my former home on the Hudson and seem to see, I did not see it then, our woodland and our meadow violet meeting in such places as I have described and crossing over into each other—how else than through free hybridi- zation? And the perplexing intermediate examples that I had then tried to sort into species seem, as the specimens are turned to now, to lend strong confirmation to such an hypothesis. If, then, we have indeed here come upon the truth, our ‘‘ papilionacea”’ is seen to be not all that we have been asked to believe. Yet by this very reduction it takes a clearer outline as a plant marked by a characteristic hairiness on the convex side of the petiole, often localized just below the blade, but often, also, thinly diffused on the lower surface of the lamina, precisely as Mr. Holm’s drawing so faithfully portrays. The upper face of the blade is by no means always glabrous, but what pubescence may later appear there is not often very obvious. By this at first strictly dorsal pubescence it is a marked plant, for it should be noted of the glabrous ‘‘ papilionacea ’—whether the hybrid, if so it proves to be, or the enhanced Viola obliqua—that the slight pubescence it may sometimes show has its site on the upper surface of the blade just where we find evidences of it in the nearly glabrous Viola cucullata. It is a marked plant also, in its group, by deeply cordate and crenate-dentate leaves which in age so open out their cordation as to become subtruncate at the base, and it is an especially noteworthy violet by reason of a ready tendency to semi-domestication. It will be well here to turn to yet another one of our violets, the pubescent Viola sororia Willd. In woodlands wherein this species and Viola obliqua are in free growth together they may be found, it is no uncommon thing, blending the one into the other in perfect confluence. Among these plants of mixed strain we recognize, now Viola obliqua, changed only by the beginnings of 266 BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS pubescence, now its too intimate associate, still to be called Viola sororia but taking a more gracilent habit and narrower form of leaf. More intermediate in the series are plants of stronger growth and in these we seem to see our pubescent “ papilionacea,’’ never, perhaps, exactly as we know it in its semi-domesticated state but so much the same that the origin of our plant would seem to be disclosed as though in an open book. We have been here freely following appearances and may have been easily mislead in thus seeming to have traced our plant back to the mode of its beginning. - Nothing like demonstration has assured us. But, if it were indeed a proved thing, even then our semi-domesticated “ papilionacea,"’ far along in its generations, would be manifestly of a higher category than those chance hybrids we seem to come upon in the making. Our plant thrives in places whence both of its suggested parents have disappeared, or perhaps have never been. It is become independent of the parental aid, is self per- petuating. It is, by continuous descent from season to season, no longer a hybrid, but rather a species whose hybrid origin goes back —how far we may not know. Such a plant is surely to be ac- corded its own distinctive name. I have not found that such a name has ever been given unless the pleasing one Viola laetecaerulea of Doctor Greene may happily prove to be available. Most certainly we cannot continue to call this plant Viola papilionacea Pursh. We turn to Pursh and read under this name, it is so clear we cannot be mistaken, the quite sufficient description of no other violet than our common one of boggy meadows and wet places that we have been calling Viola cucullata Aiton. Of this violet there is an open meadow form, it has doubtless been remarked by all of us who have given any field attention to violet matters, that has definable points of difference from more usual phases of the plant and this, may we not say unmistakably, is the form more particularly held in view by Pursh. In my collections formerly made about Van Cortlandt Park I find specimens of this plant put aside as far back as 1895 under a herbarium name given with reference to the notably triangular cordate and acute leaves well-developed as early as the time of flowering. The leaves are not strongly cucullate nor strictly gla- brous nor is their marginal pattern at all pronounced (‘ triangu- BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS 267 laricordatis acutis crenatis subcucullatis glabriusculis’’—Pursh's de- scription could scarcely be more exact) ; the peduncles, at flowering, little if at all surpass the leaves in height (pedunculis longitudine foliorum); the flowers, compared with those of the Viola obliqua series, are explicitly more papilionaceous, allowing for the fanciful application of this adjective to the: flower of a violet, (‘‘ petalis obovatis: 3. inferioribus infra medium barbatis conniventibus, 2. superioribus reflexis’’).* Point by point Pursh’s description meets the distinctive characters of this plant, proving slightly inexact only in respect of the variable bearding of the petals, for the odd one, although sometimes slightly bearded, is prevailingly glabrous. The bearding dusted with pollen readily becomes Pursh’s ‘“ yellow down.’ ‘‘ Flowers blue, elegantly striated,”’ points with unmistakable indication to the flowers of our meadow violet distinguished above all others by the delicacy and sharp beauty of their dark penciling. ‘‘ In wet places’ would scarcely be particularly affirmed of any other blue-flowered heart-leaved violet, although Pursh does say of his other one “In grassy wet places,’ wherein, however, is more of a distinction than might appear to one not well knowing our violets in the places where they grow. Upon the face of the evidence this other violet of Pursh’s, his ‘‘ Viola cucullata Ait.,’”’ was our Viola obliqua. Its glabrous leaves, cucullate only at the base, had more promi- nently indentured margins than the crenate leaves of his Viola papilionacea, he expressed the difference by calling them “ ser- rate ’’; the scapes were shorter, an essential distinction; the petals were obliquely bent, therefore the upper pair less characteristically reflexed and the others more open. The two plants are thus placed by Pursh in unmistakable apposition. The lateral petals are described as being merely ‘‘ bearded,” not, as in his papili- onacea, ‘‘ infra medium barbatis,” an acute distinction, not to bear a too literal rendering, but quite true in the sense that the bearding in the one species is more restricted and relatively mcre basal on the petal than in the other. Mr. Stone is, I think, the only recent writer who has called attention to the more forward extension of * It should be said that this description applies more particularly to the flower in its earlier and its later stages—I know of no violet the flower of which at the stage of fullest development is not more or less widely expanded. 268 BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS the bearding in Viola obliqua—his Viola affinis.* ‘‘ Flowers blue, white at their base,”’ points also to Viola obliqua for, while the petals of all these violets are white at their base, the flowers of this one are lighter towards the throat where those of our “ cucullata’”’ are characteristically of deeper hue, often in sharp contrast with the pale blue surrounding parts. It is not to be believed that Pursh did not know both of these common violets, and I have been at pains to determine and to lay stress upon each of his described species the better to emphasize the identity of his Viola papilionacea, for it would appear that by right of priority this is the true name of our meadow violet that we have been miscalling Viola cucullata Ait. If a decade and more ago we knew as much about our violets as we know today, however scant our present knowledge may be, Aiton’s Viola cucullata would scarcely have been construed in terms of our meadow violet that, as we have just seen, Pursh called Viola papilionacea. Doctor Greene in pointing out to us this distinct but long hidden species did not adopt for it a doubtfully applicable name without using a deliberate mark of interrogation.7 However unmistakable the description of any later author may be, there is not one word by Aiton himself that can be deemed distinctive of this plant. Quite otherwise. His Viola cucullata had, for instance, subterete scapes shorter than the leaves, which were attenuate at the apex, and the petals, the upper pair not being reflexed, were white at their base. His entire description differs in no essential from the description of Hill’s Viola obliqua, nor does it fail at any point to apply to that plant. That species must have been the very one he had before him but, believing Hill’s plant to be characterised by erect flowers, partly stramineous in color like some of the European species, he very naturally considered his own plant to be distinct by reason of blue inverted flowers on scapes reflexed at the apex, which characters, although common to all violets, he is particular to report. Can it be doubted that the name Viola cucullata Aiton is but a synonym of Viola obliqua Hill and that our meadow violet that we have allowed to bear Aiton’s name should now inherit from Pursh the name Viola papilionacea? * Loc. cit., p. 671. } Fittonia 3: 143. 18096. BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS 269 Yet another violet should here receive a word. Little recog- nition has been accorded to Viola domestica. I ask myself, Can it be alone from force of first impressions that this violet remains to me one of the most individualized and set apart species of its group? No other one is altogether glabrous. I have given the closest scrutiny to very many growing plants and failed to detect on even one so much asa single hair. Rare examples viewed by lens do show some obscure appressed spiculae near the margin of the leaf on its upper face, but such plants are so unusual as to suggest some admixture in the strain. By this rather remarkable absence of pubescence this violet is at marked variance from the one that has been combined with it under ‘ papilionacea,”’ in which some dorsal hairiness on the leaf is so constant a character. There is other evidence that these two violets are of c_llateral rather than lineal relationship. In more than one direction they disclose an obviously different course of growth. In the pubescent one, I know not what other name to call it by than Viola laetecae- rulea Greene, the scapes at flowering are erect, bearing the flowers high, even above the leaves, later becoming flexuous or sometimes declined; in Viola domestica they are always shorter than the leaves and tend to rise obliquely, sometimes bearing the flowers out around the sides of the tuft. The light green leaves of Viola laetecaerulea at flowering time are normally deeply cordate, later becoming dilated and taking a more or less subtruncate base; those of Viola domestica are from the first openly cordate or sub- truncate and show much less change of form with age. They are of a strikingly bright deep green and when young somewhat succulent and shining. The flowers are unlike those of Viola laetecrerulea, or any other violet known to me, having longer more twisted often narrowly rhomboid petals of deeper hue intensifying into a dark true purple. The upper pair when in ultimate position are not only reflexed but deflexed backward by a downward twist from the base. Viola laetecaerulea is one of our earliest flowering blue violets, Viola domestica the latest of all. While the former is partly domesticated the latter must be, I think, considered as wholly so. _ It is found by fence rows, in old orchards, yards and abandoned grounds, growing in rich but never in wet soils and in shade or partial stade as if it had come originally from 270 BICKNELL: VIOLA OBLIQUA HILL AND OTHER VIOLETS the woods. But I have never found it in a really wild state, that is to say, never far away from the habitations of man. It is of more gregarious habits than any of our wild species, often colonizing so thickly over wide spaces as to crowd out most other plants and, until taking its strong later growth, is low and somewhat spreading of leaf and peduncle, keeping close to the ground; not least to be noted is its relatively narrow range of variation, which is un- usual among the violets of its immediate relationship. It has been classed with ‘‘ papilionacea’’, although its nearest ally is doubtless Viola obliqua; but it is too signally different from that species to be forced upon it. Because hybridization may have at some period entered into the history of Viola laetecaerulea, of domestic tendencies, there might be reason for inquiring whether Viola domestica had not a like origin, but, if Viola obliqua is or was one parent, where is the other? Should the indications reported in this paper not be mistakenly understood the facts before us would be these: That the name Viola obliqua Hill belongs to the common and widely variable violet that we have been calling Viola affinis LeConte. That no such species exists as the supposed one we have been calling “Viola papilionacea,” this being a mixture, partly an accentuated phase of Viola obliqua, partly a hybrid of that species with our meadow violet, partly also a cross of Viola obliqua with Viola sororia and partly a well appointed violet, perhaps descended from such a cross, which may be called, pending proof, Viola laeteczerulea Greene. That the Viola cucullata of Aiton is a synonym of Viola obliqua Hill, and that our meadow violet that we have known as cucullata was first described by Pursh, receiving the inalienable name Viola papilionacea. That Viola domestica is, by attributes of form and habit, invested with a signal indi- viduality and that notwithstanding its domesticated nature the source of its origin does not yet appear. NEw YORK ; : ; ve, ‘ Observations on the inception, season, and duration of cambium development in the American larch {Larix laricina (Du Roi) Koch,] * L. KNupson (WITH PLATES 18 AND 19) INTRODUCTORY During the past twenty-five years very little attention has been devoted to a minute study of the diameter increase in trees. Comparatively little is known concerning the season of wood formation; and with respect to the region of the tree in which diameter increase first begins, the evidence is contradictory. With the object of determining the part of the tree in which cambial activity begins, as well as to determine the season of growth, investigations were begun during the season of 1909 and continued in 1911. The results obtained from the study of mate- rial collected these two seasons form the basis of this paper. The subject was suggested by Prof. W. W. Rowlee, and to him, as well as to Prof. B. M. Duggar, the writer is indebted for helpful suggestions. HISTORICAL The work of von Nordlinger, Th. Hartig, Robt. Hartig, Mer, and others has thrown some light upon the extent and duration of cambial activity. They have found, in general, that under forest conditions growth first begins in the youngest twigs and then proceeds downward into the older regions. Less work has been done on the cambial activity in isolated trees. Concerning the region of first cambial activity, Th. Hartigt concluded that it occurred in the youngest twigs and then gradu- ally extended downward. In a 30-year old Pinus sylvestris, and also in oak, cambial activity began almost simultaneously over the entire trunk, while in larch and maple of the same age cambial * Laboratory of Plant Physiology, Cornell University, Contribution No. 8. + Hartig, T. Anatomie und Physiologie der Holzpflanzen, 368. 1878. 271 272 KNUDSON: CAMBIUM DEVELOPMEMT IN AMERICAN LARCH activity began from two to four weeks later in the lower part of the trunk than in the twigs. Asa result of extensive investigations, Robt. Hartig* advanced the idea that cambial awakening was dependent upon temperature and that therefore the thickness of the bark, temperature of the soil moisture, and insolation, were important factors. He found in an isolated 10-year old Pinus sylvestris, that cambial activity had begun two weeks earlier than in isolated 35- and 65-year old trees, and four weeks earlier than in a 1o0-year old tree grown under forest conditions. The comparisons were all made at a height of 6 meters. He also found that under natural forest conditions the growth of the annular ring of Scotch pine, Norway spruce, and European larch at a height of 27.5 meters, was on June 9 respectively 66 per cent, 56 per cent and 75 per cent completed. Going toward the base the growth decreased, and at a height of 1.5 meters the percentage of the annular ring com- pleted was 35, 21 and 18 per cent respectively. In isolated trees of Scotch pine and Norway spruce the growth on July 9 was ap- proximately the same in all parts of the trunk. Under forest conditions growth was found to begin in the twigs and proceed downward, the cessation of growth following the same order. According to Mer,} the cambial activity in oak, beech, bass- wood, fir, and other trees of twenty-five years of age and under begins in the youngest twigs. In older trees cambial activity is described as simultaneous at the bases of the branches and trunk. He states also that in a single cross-section cambial activity may be evident on one side and not on another. Hastings} found that in broad-leaved trees increase in diam- eter did not begin until the buds had opened. He found that growth first begins in the 1-year old twigs, and later it occurs in 2- and 3-year old twigs. When wood is forming in 5- or 6-year old growth there is simultaneous development over the entire tree. In pine it begins first in the 2- and 3-year old twigs. In the hemlock the growth was first observed in the 6-year old twigs, * Hartig, R. Das Holz der deutschen Nadelwaldbaiume, 35-38. 1885. {t Mer, E. Sur les causes de variation de la densité des bois. Bull. Soc. Bot. France 39: 95-105. 18092. t Hastings, G. When increase in thickness begins in our trees. Science II- 12: 585. 1900. KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 273 while in Taxodium distichum the same conditions prevail as in the broad leaves. Buckhout* made during a period of four years caliper measure- ments of European larch at intervals of five days during the growing season. The measurements were made at breast height and the age of trees experimented upon is given as 45 years. He found that the formation of leaves was coincident with the begin- ning of diameter increase. The beginning of this increase was close to April 25, during the four years. He found a gradual increase from this date until about July 1, when further growth in diameter practically ceased. The data secured from this method of measurement are not, as he himself realized, entirely conclusive, on account of the errors which may result from the swelling and shrinking of wood and bark with the varying moisture content. From the horticultural side gross investigations have been made by Keffer,t Goff,t Cranefield,§ and others on the duration of growth in fruit trees. Their work is concerned with the develop- ment of shoots and on the duration of wood increase as determined by the readiness with which the bark could be peeled. The work of these men will be considered in a subsequent paper. METHODS OF INVESTIGATION For the investigation during 1909, four larch trees of approxi- mately thirteen years of age were used. These trees are hereafter designated for convenience as trees A, B, C, and D. The trees originally grew in a swamp in Oswego County, New York, but were transplanted in 1902 to the nursery on the Cornell University Campus, on land which slopes gently to the west and is of a well- drained, heavy clay soil type. The trees were planted four feet apart and were shaded on the east and west sides but not on the Sosy ie ee PS” Lae ee oo el ae Sar a ee lens * Buckhout, W. A. The formation of the annual ring of wood in European larch and the pine. Forestry Quarterly 5: 259-267. 1907. + Keffer, C. A. The early growth and training of apple trees. Tenn. Agr. Exp. Sta. Bull. 14: 1-16. » EB rie The resumption of root growth in spring. Wisconsin Agr. Exp. Sta. Ann. Rep. 15: 220-228. 1898. § Cranefield, F. Duration of growth period in trees. Ann. Rep. 17: 300-308. 1900. Wisconsin Agr. Exp. Sta. 974 KNUDSON: CAMBIUM DEVELOPMENT OF AMERICAN LARCH north or south. Trees B, C, and D were very uniform as regards size and form. Tree A, although of the same age, was slightly smaller. In’ order to determine the region of growth inception it was necessary to take material from the apex of the tree to the base. The larch has, at intervals, whorls of branches, the number of which agree approximately with the age of the tree. The trees used had each ten such whorls and material was removed from below each whorl, at different times throughout the growing season. Cuttings were made only from the south side of each tree. The first cuttings were made a few inches below each whorl of branches and the subsequent cuttings were made a few inches below the preceding and a little to one side. In obtaining the material for study two incisions, 2 cm. apart, were made through the bark and into the wood to a depth of 1 cm. and the piece then removed with a small knife. The injured area was then filled with grafting wax. The material collected from trees A and B was fixed in a solution consisting of 33 parts glycerine, 35 parts alcohol, 30 parts distilled water, and 2 parts glacial acetic acid. The material kept in this solution was in excellent condition for sectioning, though, of course, no good fixing of the protoplasmic structure was obtained. That collected from trees C and D was fixed in Gilson’s solution and kept, by mistake, in 95 per cent alcohol. When attempts were made to section it, several months later, considerable difficulty was experienced, because of brittleness. Attempts to soften the material, by allowing it to remain in equal parts of glycerine and alcohol, and also in glycerine alone, proved futile. The greater part of the material was sectioned without imbedding, but some of it was necessarily imbedded in celloidin. The sections were cut from 20 to 40 yw in thickness and stained with safranin and Delafield’s haematoxylin of the formula so commonly used for wood staining. The methods employed during the season of 1911 are described subsequently. INVESTIGATIONS OF 1909_ The first cuttings were made on April 19 and at this time the buds located on the 4-, 5-, and 6-year old wood had opened, KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 275 the leaves being 1/16 of an inch in length. On the younger wood, the buds were less advanced. This was more marked in the 1- and 2-year old wood. This slower development toward the terminal shoot and apex of the branch held true also for the catkins. The same condition was noted also in several larches which in Sep- tember produced a new growth of leaves, the result of a drouth, followed by favorable conditions. This earlier development of leaves on the older wood is significant in the light of the subsequent facts concerning the inception of cambial activity. Cambium in resting condition.—According to Sanio and other investigators the cambium proper consists of a tissue but one cell in thickness, which cells by division produce a row of xylem mother and a row of phloem mother cells. Each of these rows divides and gives rise respectively to two rows of potential xylem cells and two rows of potential phloem cells. Except by careful cytological study the row of true cambium cells cannot be distinguished from the neighboring cells. The term cambium has been, therefore, generally applied to that tissue which lies between the visibly differentiating phloem and xylem. The cambium tissue is com- posed of a number of rows of cells, which cells are characterized by their thin walls, dense protoplasmic content, and, viewed in cross section, rectangular shape. In trees in the resting condition it would be reasonably assumed that the true cambium comprises the first row of cells just without the xylem. The cells bordering this row on the outside would then be considered as phloem. As a matter of fact, however, sections made from cuttings obtained from the trunk of larch on November 13 exhibit just outside of the xylem a distinct tissue 34 u in diameter, consisting of five or six rows of cells in thickness. The cells of the outer five rows are not visibly distinguished from cells of the inner row, but are distin- guished from the adjacent phloem cells by their size and proto- plasmic content. See FIG. 1 and 3. Because of the similarity of all of these cells I have considered the six rows as comprising the cambium tissue, the term cambium being employed in its generally applied sense. Inception of cambial activity and development of phloem—The material collected on April 19 showed that cambial activity had begun. The layer of six cambium cells had increased in diameter. 276 KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH The outer cells of this tissue were losing their rectangular shape and assuming more nearly that of a square, as viewed in a cross section. See FIG. I, 2, and 3. In studying the slides made from material collected on April 19, it was found that in the 5th, 6th, and 7th cuttings of tree A the cambium had developed to a greater extent than in the other cuttings. Not only had the cambium increased in diameter, but seemingly new cells had been formed. This increased development near the middle was maintained until May 25. The average increase in number of phloem cells by May 25 was only 1.8, but the diameter increase of the cambium and phloem was nearly 100 per cent. Up to this time no xylem what- _soever had been developed. It appears therefore that the earliest growth consists in an enlargement of the cambium tissue with the gradual transformation of the peripheral cells into phloem tissue. The old phloem cells adjacent at this time are becoming compressed due to the pressure brought about by the transformation of the cambial cells. Compare FIG. 1, 2, and 2. In TABLE I are given the figures obtained by the measurement of the diameter of phloem and cambium tissues in trees A and C. The figures for the diameter of the cambium tissue during the resting period (cutting made November 13) are given for com- parison. The figures included under the dates April 19 to May 25 inclusive refer to tree A. From June 3 to July 6 the figures refer to tree C. As indicated previously, the six-celled layer adjacent to the xylem is considered the cambium. Although transformation of the peripheral cells had occurred, it is difficult to state which cells are cambium and which cells are phloem. Consequently the six rows, despite the transformation, I have considered as cambium. Any cells in excess of the six rows, which lie within the old com- pressed phloem cells, I have considered as new phloem, After May 25, when xylem and phloem were both developing rapidly, the cambium tissue was still considered as a tissue of six rows of cells. It was difficult to select always the six most uniform rows, but in general the error was slight and at most of little consequence. From an examination of the table it may be seen that up to May 25 the middle regions show the greatest growth. From April 19 to May 25 the increase in phloem was gradual, but from May 25 TABLE I NUMBER OF ROWS OF NEW PHLOEM CELLS AND DIAMETER OF PHLOEM AND CAMBIUM TISSUE IN TREES A AND C Nov. 13 gue: 19 » | eo Mag 5 E, Mas: rr a3 ae 25 & ‘ies 3 ioe Gs 15 June 30 eS Se ee Po ‘ Sarasa CRS aa A nae eater No. "ORE No. — No. ape No. a 78% a No. og | No. a we pe No. | ~~ Below 1st whorl ....| 0 | Sec @. Sh iO ae 16 ax 6 58 | | 51 | 7\t22 | 4 7 146 Below 2d whorl ....! 0 | 34 |o 54 ry | 58 I SP ole. PR eo! sy | 9 | I70 Below 3d whorl . oO; 34 {0 | 58 |x | 68 |o } 58 Jo | 5: l4 | 75 ro | 184 | 12 224 4th whorl ©} 34 |2 | 58 | P12 6F -<\o Sho) OR ee eS. ro) 489 >) & |) 284: | ts 206 Below 5th whorl OR ae Ere GR Sle q1 2 Wh) S| ae (et BS ae Sed lp | eye | tg 278 Below 6th whorl Ose Wat or r | 68 2 WAS a lay Sits 2| 221 OQ. | axr 14 273 Below 7th whorl. Ot) 34-12 1 OB. tt 1 6s I 58 ir { 68 Bhi OE [12|207 | 9 | tor 13 262 Below 8th whorl Oo} 34 |o } 58 | o } 48 I SI} 2 1) 6b 32.1 58.) 24) 90° |r4>] a1 I4 278 Below oth whorl O0| 34 |0 | 44 jo Ly Suse fe Be ae eo corn gs me Ba |\— 1%. |} 30 — roth whorl OP SR PO AA bol ey cy 44 (0 | 51 | SI 12 | 204 iro 204 — - — — — a tee eee ee Average... ©} 34 |0.6) 54.7 (0.6) 57. 57-5 5 | o7 Sete | 61.4 18 63.3 ne | 190.7 (9.6 189 | 13.6 | 268 L1@ HOAV'T NVOINANY NI INANdOTIAAG WAIEWVD : NOSGANY 978 KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH to June 3 the growth was markedly increased, while from June 3 to June 15 seemingly little growth occurred. After June 3 it appears that relatively few new phloem cells were formed, but growth consisted more of an enlargement of the cells already formed. Some of. the figures for June 15 are less than the corres- ponding figures for June 3. This may be perhaps explained by the fact that the cuttings were made somewhat lower, or better perhaps by the fact that in obtaining these cuttings they were not taken on a line directly below the preceding, but to one side. Mer (loc. cit.) draws special attention to the well-known obser- vation that wood growth is not always uniform on all sides of the tree. rH PreMed aad cee etait 4 Sagessesas Cutting. DiacraMi. Diameter of cambium and new phloem at various different dates. A, Nov. 13; B, April 19; C, May 25; D, June 3. KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 279 By July 6 the phloem was nearly complete with respect to cell numbers; for, the average annual number of rows of phloem cells produced during a period of three years was found to be seventeen. At this time the cells were all very regular in form and no visible differentiation had occurred. In DIAGRAM I are represented the diameter measurements of the phloem at several different dates. In trees B and D the growth is similar to that in trees A and C, though more vigorous growth resulted in tree B than in tree A. In this tree also the development of cambium consists first in a transformation of the peripheral cells. In this case, however, the number of rows of phloem cells produced by May 25 was 5.6 with nearly a 200 per cent increase in diameter. The detailed figures are given in TABLE II. TABLE II NUMBER OF ROWS OF NEW PHLOEM CELLS AND DIAMETER OF PHLOEM AND CAMBIUM TISSUES IN TREES B anp D Nov. 13 May 5 | May 11 May 25 | June 8 June 18 Region of cutting coe ee ; 2 ce Dia. | Dia. No. _ No. oa No. ee No, ot No. i No.) pr Below 1st whorl..... o| 34 |— — | 75 |2 ' 68 | 8 | 170 11 | 228 Below 2d whorl ..... ON SRS Sag S58. gop tae rr aes (ae 98 Below 3d whorl ..... 34°13 68 4 | 82 |4 6|— | — |11 | 286 Below 4th whorl..... o | 34 |4 | ro2 |4 | 102 |6 92 |12 | 221 |14| 262 Below 5th whorl..... 0 | 34 | § 85 | 6 | tis eee | 262 Below 6th whorl..... 0 | 34 |4 68 }8 | 107 |7 92 |I0 | 190 13 | 238 Below 7th whorl... .. | OSF ars) ts ee ' So A oe mca poppe 279 Below 8th whorl. .... o| 34 14 | 85 oe — |6 | 92} 8 | 170 |—| Below oth whorl... .. Ot Gare aa ot es 6 705) 8: | 293.412 | 245 Below toth whorl....| 0 34°13 | Wer. 6 awe mags Bie ee as 238 | 34 |3.1| 72 | 4.8 89 Is.3| 90 9.5 186 12 | 250 Average. .....-. | 9 Development of xylem.—While in tree A there was a gradual increase in phloem from April 19 to May 25, yet no xylem cells had been formed. During the week of May 25 to June 3, coincident with the marked increase of phloem, a very marked increase of xylem occurred, over one third of the xylem being completed - during these seven days. TABLE III gives the figures obtained for the number of rows of xylem cells formed, and the diameter of the xylem tissue in the various cuttings at the different dates. There is included also the diameter of the xylem tissue formed in tree B during the years 1909 and 1910. In the first column the 280 KNuDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH figures refer to tree A. The figures from June 3 to July 6 refer to tree C. TABLE III NUMBER OF ROWS OF NEW XYLEM CELLS FORMED AND DIAMETER OF NEW XYLEM TISSUE IN TREES A AND aA May-25 June 3 June 15 June 30 July 6 | 1909 | 1910 Region of cutting Bee be . eee | | p; 2 Rl worn ewes Dia. No. No. aa) No. | ay | No. | — | No. | — | pe 8 Below 1st whorl ..| o 15 | 262 | 25 544| 43. |1,054! 646} 986 Below 2d whorl . ° 3 833! 55 1,564, 1,598|1,377 Below 3d whorl 0 = [20° - }°578"1.33 48 |1,462 | 1,853|1,870 ow 4th whorl co) 29 | 697 | 43 |1,122| 50 |1,462 1,785|1,870 Below 5th whorl o 28 | 748| 45 (|1,292| 55 1,530! 1,360/1,955 Below 6th whorl re) 23 646 | 46 (1,326) 58 ,666 I,275|1,224 Below 7th whorl te) 24 | 648 | 36 |1,122] 57 3 734|/1,.428 Below 8th whorl..| 0 22 | 714/36 11,156)57 (1,632,65 1,972|2,363/1,59 Below oth whorl..| o 1 607+) a8 | 50) | — | 62 /|1,870/1,666]1,088 Below roth whorl. oO 20 | 663 | 27 | 833/58 (1,564 58 (1,802 | | | | | — beac Average... i... | O°) 22-5 (636 }-35.2 999.6. 53-3 1,493 61.6 |1,881 1,586 1,488 y | 1 eee From the table it cannot be determined exactly in which part of the tree xylem growth first begins. The greatest growth by June 3 was below the fifth whorl of branches while the least growth was just below the first whorl of branches. In general, by June 3 more xylem cells were formed in the middle of the tree than at either the top or base. This would tend to indicate that the first growth of xylem occurred in the middle of the tree below the fifth whorl of branches. TABLE IV NUMBER OF ROWS OF NEW XYLEM CELL S AND DIAMETER OF NEW XYLEM TISSUE IN TREES B anp D May rr May 25 June 8 | June 18 Region of cutting ar age nee 4 TRSNEP Nos Me | | No: aire ee gone Below 1st whorl. 205605222 o ty) (3) 22 442 43 | 1,003 Below 20 whort, . 4.05 5.553 0 ce) 0 40 647 53 | 1,326 Below 3d whorl, 02: (3) ts) o — — 55 1,394 Below: 4th whorl. 0. 0 4 ia: 35 918 50 1,258 Below: sth what. 3. 5 8. o —_—- |— — 45 | 1,202 Below 6th whorl. ............ ° 5 55 32 850 46 | 1,122 Below 7th whorl........-.... oO 2 20 — sie 55 1,258 Below 8th whorl............. oO 0 to) 34 850 -- — Below oth whorl............. 0 ° ° 30 782 50 | 1,292 Below. roth whorl... oO r9) o 49 1,292 Average 45 Uc a ee ° 1.2 14 32 nas |! ag 4 -3aa8 TABLE V TOTAL NUMBER OF ROWS AND DIAMETER INCREASE OF PHLOEM, CAMBIUM, AND XYLEM CELLS IN TREES A AND C | Nov, 13 | Apr. 19 Apr. 26 May 5 May 11 May 25 June 3 June 15 | June 30 July 6 Region of cutting an a tr 4 ae Seay ape Pm ae No. _ No. oe No. - No. ee No. nay No. are No. cin Po tees ios No. bs Below 1st whorl. 6 bo aes 34 | 5 aa eel At ai) Seg rea elo 28| 384 | 38 — — | 2d whorl ., Go) e416 Ste Sah Grol ar 6 | 58 oy ee —/| — | 48 1,003 | — — Below 3d whorl ....| 6 34 |6 Sr 16 TN © ae Gf Sr oura 1s 75 —) — 49 I,t0T | 66 1,686 Below 4th whorl 6 34 |8 68 6 62 |8 51 8 | 68 101° 975 45| 884 | 58 1,305.) 71 I,758 Below 5th whorl ee Te OE BCE aE lg. | 9E Oren) 68 9| 85 |45| 952 | 60 | 1,468) 75 1,741 Below 6th whorl Go a4 17 15. \8 68 |8 | 75 8 | 72 O°} 75 41 | 867 | 62 1,536 | 78 1,045 Below 7th whorl....| 6 | 34 |7 58 19 65-217, | g8 4 |. 68 Ht OL Pris ah eal Wa te I,312'| 76 1,842 : Below 8th whorl....| 6 | 34 |6 st Oe 7A te a 4 | 61 bp ae a 3 42 15038:| 156 1,332 | 77 1,910 | 86 | 2,251 Below oth as ne We Or ae LG 44 |6 Br BE vette 6 | 51 —| — 45 1,057 | — = 2,125 Below toth whorl. 6 34 16 44 |6 Sto: 6s sr tes HL 9 6 | 51 38| 867 | 43 1,037} — — | 76} 2,040 Average. .... tery 3 6 34 l6 4l 53-7 | 6.6 56 7.1| 55.8 | 7.1! 60.8 ! 7.5] 63.3 |40| 820 | 51.2 | 1,184 | 73.8 | 1,813 | 82 | 2,138 NosdaANy I8Z HOUV] NVOMANY NI LININdOTIAAG WaAIlEKv> 282 KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH While in trees A and C the region of the first growth of xylem could not be definitely located, more fortunate results were ob- tained with tree B. No xylem was formed in tree B until May 25 and then neither at the top nor at the base of the trunk, but in the middle region. By May 25 then the xylem in that part of the tree between the fourth and fifth whorls of branches was four cells in thickness,—while between the sixth and seventh whorls it was five cells and between the seventh and eighth it was two cells in diameter. The most marked development again occurred im- mediately after May 25. The detailed figures are given in TABLE Iv. Season and duration of diameter increase-—In order to bring out more clearly the period of greatest diameter increase, the combined values for increase of phloem, cambium, and xylem, in trees A and C are given in TABLE vy. - From the table it is at once evident that very little growth took place previous to May 25, and the growth which occurred was confined, as before indicated, entirely to the cambium and phloem. From May 25 to June 3a very marked increase occurred, and during that time in the middle regions of the trunk nearly one half of the total diameter increase was completed. In TABLE VI is given the daily average increase in rows of cells and also the daily diameter increase in trees A and C. TABLE VI Period May 25-June 3 June 3-June rs / June 15-June fe June 30-July 6* No. rows of cells......... | 4.2 Z; 1.4 I. Dia. in microns.......... 94.5 | 30. | 4o. 48. By July 6 the ring of xylem was almost complete, judging from the extent of the summer wood which was then in the process of formation, though not yet completely developed. Unfortunately no further series of cuttings were made after July 6 to determine the succession with regard to growth cessation. In TABLE VII is given the combined diameters of the phloem, cambium, and xylem tissue in trees B and D, as well as the total number of cells. * For cutting nos. 9-10 only. KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 283 TABLE VII TOTAL NUMBER OF NEW ROWS AND DIAMETER OF PHLOEM, CAMBIUM IN TREES B AnD D Nov. 13 | Mays «| Mayxixz May 25 June 8 June 18 Region of cutting Fa tags eee ER d 3 ; se No. ae No.| sir | No. res | No. i No. ce Below 1st whorl....| 6 | 34 |— 75| 8 | 68 |36 | 612/60 |1,231 Below 2d r .|6| 34 | 6| 34] 9| 58/12 | 89 |57 | 840) 70 [1,544 Below 3d whorl 6 | 34 | 9 | 68.120 | 88 (0-3) 9S 1,680 Below 4th whorl 6 | 34 |10} 102/10] 102 |16 {143 {53 {1,071} 70 {1,520 Below 5th whorl aad dee | ea eo | Ss NTA hs ire a ee re 1,486 Below 6th whorl 6 | 34 |10| 68/15} 108|18 (147 | 48 |1,040/65 |1,506 Below 7th whorl 6 | 34 |—| — bs ae Birt Be S| eg ieee i af foams BR 8g Below 8th whorl OG 5421085 pea toe ae 92 | 48 so Below oth whorl....| 6 | 34 8 61 —| — /|12 76 |44 | 955/68 (1,537 Below 1oth whorl...| 6 | 34 | 9) 75 —!|— |— |— |— | — |62 {1,530 Average . hee eee OL Ba 9! 72 11! 901 12.6 'TO1.5 ATS | 917! 69.4'1,508 The season of diameter growth in larch is relatively short. Practically all of the growth occurred during the month of June. The small increase produced from the time of bud opening, April 19 to May 25, consisted entirely of cambium and phloem develop- ment. The diameter increases of trees A and C and B and D during the season 1909 are graphically represented in DIAGRAM II. INVESTIGATION OF IQII Since the studies made in 1909 indicated that growth in diameter first occurred in the middle region of the trunk and not at the base or apex, it seemed advisable to continue these studies with respect to the development in the lateral branches. The investigation was made with the object of answering the three following questions: (1) Does inception of diameter increase begin first in the topmost branches, in the middle, or in the basal, branches? (2) In what part of the individual branches does diameter increase begin? (3) Does xylem development in the lateral branches precede that in the trunk? For this investigation a larch tree of approximately Io years of age was used. It was growing isolated, about 100 feet from the trees mentioned above, growing in the nursery row. For con- venience this tree is hereafter designated as tree E. On May 22, from each of the whorls of branches from apex to base, one branch CAMBIUM DEVELOPMENT IN AMERICAN LARCH 284 KNUDSON : =e TESTER BREE a = aa “pe Berle A te Hit eee Hames Petal i B, diame . » mae | a ac aim ee APRIL ID = 2 y A, diameter increase in trees A and . * vane re om ae oa Syaepns a aD inti 4 HEHE DIAGRAM II. trees C and D. KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 285 was removed. The removal, as before, was made from the south side of the tree. The branches were numbered according to their position on the tree, the topmost branch being No. 1, and the basal branch No. 7. From the individual branches there was removed from each season’s growth a cutting; from the topmost branch two such cuttings were removed and from the basal branch TABLE VIII NUMBER OF ROWS OF XYLEM CELLS AND DIAMETER INCREASE OF XYLEM TISSUE IN THE CUTTING FROM LATERAL BRANCHES OF TREE E Region of cutting on lateral No. of rows of new | Diameter increase No. of branch branch xylem cells of xylem Below rst whorl Below 2d whorl Below Ist whorl Below 2d whorl Below 3d whorl Below Ist whorl Below 2d whorl Below 3d_ whorl Below ss whorl WH WwW Ww bo @Sorwrnmoonroons Fc moon Hms9 07 OOO 080 espe ae ~4 ne Lal ww ; w .~ WR Lal Below 7th whor Below 1st whorl ’ Below 2d whor Below 3d whor Below 4th whor Below 5th whor Below 6th whor Below 7th whor _ Below 8th whorl Ll cal oOo OC SOMO O Re ieee | SIT ITTTIAADADAGCUUUUNUNUNEERREAWWWHNHNRNHH ooo] Oo Oo — o 9 4 ey co 3S 4 rp ° bo Pah JOSCOHOHOONONHOCONDONOHHRONHNHNOONNOOOOSO seven cuttings. These cuttings were fixed, sectioned, and pre- pared as before. In the following table the figures given refer only to the number of cells in the new xylem layer and to the diameter of that tissue. The diameter increase over the entire tree is represented in DIAGRAM III. Ms 286 KNupDsON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH In order to show more clearly the results obtained and to aid in the interpretation the following two tables have been compiled from the preceding. In the TABLE Ix are presented for each of the branches the average number of rows of xylem cells and the average diameter increase of the xylem tissue for the entire lateral branch. The averages are obtained from the figures for each individual cutting. DIAGRAM 111. Diameter increase of xylem in various parts of tree E on May 2 TABLE IX AVERAGE DIAMETER INCREASE OF XYLEM IN LATERAL BRANCHES Branch Average no. of rows of new Average diameter increase xylem cells of xylem Busca pacaea he dae eee 0 LE is Se Bee eee ts) : ; jh eee rey Ratalienl Moire OU cy I 1s B AY sala) waste die nie oy cae menue te 1.25 ee ae ef wipe skh wae Ragin ee ee emg 0.83 I4.D yu PPE Se a Fee Ok, CUE me are See eke ¥ Zs ¥ VEE ss cities eee oa bees 0.25 : i - From the above table it is evident that the least growth occurred in the top and basal branches and the greatest growth KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 287 in the middle branches. This increase corresponds with the re- sults secured previously on the diameter increase in the trunk, in which it was found that the growth first began in the middle region. In regard to the second question as to the region of the branch in which growth first occurs, TABLE X is suggestive. The figures of the following table are the averages of the seven apical cuttings, the seven basal cuttings and the average diameter of xylem in the middle regions of the seven branches. TABLE X AVERAGE DIAMETER INCREASE OF XYLEM AT APEX, MIDDLE, AND BASE OF ALL LATERAL ‘CHES Region of Diameter increase cutting of xylem PGE oo ceria pis os oon pe ee RN RE eee a ER area jet as 20.8u Mitidle 5 ae a ris Me EO Oh per oes 7.5 pe Basal 2 205.2b i es Fae a ee, i ae ea 2.5 From previous results it was expected that the growth would first occur in the middle of the branch, but the evidence indicates that the development of xylem in the lateral branches begins at the apex and then continues towards the base. The opening of the buds, however, does not follow the same order. On April 26, 1911, the lateral branches of a tree in the nursery row were ex- amined and it was found that buds on the one and two years old wood had not yet opened, on the third year wood the leaves were just protruding while on the older wood the buds were fully opened. In regard to the third question, “‘ Does diameter increase in the twigs and branches precede that in the trunk?” the folldwing observations were made. From the same tree from which the - cuttings from the lateral branches were removed and on the same day,—namely, May 22, cuttings were removed from the apex to the base of the trunk. Seven such cuttings were removed from the south side of the tree, one cutting just below each whorl of branches. From a study of slides prepared from the material the following figures have been obtained. (See TABLE x1.) There are also included in TABLE XI figures derived from cuttings made on June 6. A comparison of the figures for May 22 with the figures for growth in the lateral branches (see DIAGRAM ut) reveals the fact 288 KNupSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH TABLE XI NUMBER OF ROWS OF XYLEM CELLS AND DIAMETER INCREASE OF XYLEM IN TREE E Diameter increase of No. June 6 Region of cutting | No. May 22 Sylera May s2 Diameter increase of xylem June 6 Below rst whorl... re) — | Below 2d whorl ... 3 5 | ie a Below hor 5 on ai 7 139.5 u Below 4th whorl 4 £85. oe —— Below 5th whorl 4 88.8 u 9 | 256° 7 Below 6th whorl... 7 185 yp 17 } 444 p Below 7th whorl...| 9 220.4 ub 20 | 48r yu AVeTage oii. 6558 5.8 139 FB g 286 fi 3g that except for the apical part of the trunk the growth there has been much greater than in the branches. The average diameter increase of xylem in all the branches was by May 22 only 10.7 up, and the average number of cells in the new xylem layer only 0.62. The greatest diameter increase in the branches was only 37 yp, while the average for the trunk was 5.8 cells with a diameter of 139 u. If reference is made to TABLE vit it will be seen that no growth occurred in branch No. 2 while in the trunk at this region the increase was 55.5 u. The average diameter of xylem in branch No. 7 was only 4-5 p, while in the region of the trunk from which the branch was secured the diameter increase amounted to 229 p. It is very evident, therefore, that in a larch of this age under isolated conditions growth does not begin first in the twigs and then extend to the trunk, but rather it begins first in the trunk. The figures in TABLE XI give no strong indication of the region in which growth first began. The diameter increase was greatest just below the 4th, 6th, and 7th whorls of branches. No increase was manifest near the apex. : TABLE XII Region of cutting No. of rows of xylem Dia. increase in trunk cells of xylem’ Below: 3st whorl. os syn Cutting lost Below 5th whorl............. 4 Itt pe Below 7th whorl. . 2... 202.6... 9 2224 Below 9th whorl... .......... 5 129 u In order further to check the work of 1909 a few cuttings were made from another tree in the nursery row on May 22. This tree KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 289 was now approximately 15 years of age. Cuttings were removed from the south side of the trunk below the Ist, 5th, 7th, and 9th whorls of branches. The results from the measurement of the xylem are found in TABLE XII. While no conclusions can be drawn respecting the inception of growth the indications again. point to the middle region. The figures are of interest mainly because of the fact that growth of xylem has occurred before May 25, which was the date of xylem development in 1909. FACTORS INFLUENCING DATE OF XYLEM FORMATION Temperature, moisture, and insolation are important factors in all growth phenomena. In 1909 and 1911 the opening of buds of larch began April 19. In 1909 the xylem formation began on May 25, while in 1911 the xylem formation began about May 20.* The temperature, moisture, and sunshine figures for the intervals between the bud opening and wood formation are found in the following table. The mean temperature for the day is considered - as the number of heat units for that day, and the total heat unitsT is considered as the sum of these mean temperatures. Period | Total heat units | Hours of sunshine Precipitation April 19—May 25, 1909... 1,810 | 234.5 | 2.89 April 19—May 20, IgII... 1,769 257- yA Oe The number of heat units then during the interval before wood formation in 1911 was almost as great as that during the longer period of 1909, while the amount of sunshine was greater during the 1911 period. The amount of precipitation at this time of the year was not important as sufficient water was available both - seasons. In 1911, however, the mean temperature for the three days preceding wood formation ranged from 76° F.-78° F. and for the three days preceding this period the range of the mean temper- atures was 62° F.69° F. During 1909 the nine days preceding wood formation ranged in mean temperature from 50° F.-54° F. except one day which had a mean temperature of 58. The three * The exact date of xylem formation was not determined. The observation on May 22 showed a diameter increase of 6 cells. + No daily mean temperature was below 34° F. 2990 KnupsON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH days preceding wood formation in 1909 had 39.4 hours of sun- shine; the same period in 1911 had 26.2 hours of sunshine. It is very probable therefore that during 1911 the relatively higher temperature for the six days preceding wood formation was important in hastening the inception of xylem formation. SUMMARY RESULTs OF 1909. (1) Cambium and phloem development.— The cambium in the trunk during the resting condition consists of six rows of cells 34 » diameter. On April 19 the first material collected exhibited an increase in diameter of the cambium tissue. The cambium cells were all enlarged while the outer cells were in the process of transformation, changing from a rectangular to square shape as viewed in cross section. In the middle regions of the trunk on April 19 an increase in phloem cells was evident. The increase of cambium and phloem was gradual over the entire trunk in tree A up to May 25, the greatest increase being main- tained in the middle region. Similar conditions were found in tree B, though here the increase of new phloem cells was more marked. The greatest growth of phloem occurred immediately after May 25 and was coincident with the greatest development of xylem. ; (2) Xylem development.—-In tree A no xylem was formed before May 25. In tree B a few xylem cells were formed by May 25 in the middle regions of the trunk. Growth of xylem was almost simultaneous, however, in all parts of the trunk. The greatest growth occurred immediately after May 25 and in tree C the xylem was nearing completion by July 6. RESULTS OF I9gII.—(1) Growth in diameter in the lateral branches begins first in the middle branches and is followed by that in the basal and apical branches. (2) In the individual branches growth begins first at the apex and then descends towards the base. (3) Diameter increase of the trunk precedes that in the branches and twigs. ; (4) Temperature and insolation conditions in 1911 induced wood formation five days earlier than in 1909. (5) No direct evidence was secured in tott concerning the KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 291 region in the trunk of first cambial activity. Indications pointed again to the middle and basal regions. DIscussION OF RESULTS In a number of anatomical text books it is stated that the xylem development precedes that of the phloem. This idea is conveyed rather ambiguously by Stevens.* In the American larch the development of phloem certainly precedes that of xylem and its most rapid development is coincident with that of the xylem. Brown’sf figures indicate that in Pinus rigida a similar condition prevails. The results obtained by the writer do not agree with those of Th. Hartigt with 30-year old European larch, wherein diameter increase near the base of the trunk was two to four weeks later ' than that in the twigs and branches. The factors which may operate to cause this difference are considered subsequently. My results agree with those of Brown§ who finds that in Pinus rigida the first diameter increase of xylem begins a few meters below the apex. The work of Brown was done during the same period as that of the writer and on trees in a plot adjacent to those used in this investigation. Respecting the date of diameter increase Buckhout states that in European larch it is coincident with leaf formation. It is very probable that the diameter increase at this time is due mainly to a swelling of the tissues. In my investigations the development of xylem began a month later than the beginning of leaf formation. From observations made during the past two years with a consider- able number of trees and from the results of other investigators it seems probable that in general growth in diameter does not begin until the leaves have been fully developed and have been suffici- ently active in food making to supply the requirements of rapid cell formation. The reserve foods stored up in the fall are prob- ably largely utilized in leaf and also in blossom formation, when the latter precede the formation of leaves. * Stevens, W. C. Plant anatomy, 2nd Ed., p. 170. 1910 + Brown, H. P. Growth studies in forest trees, I. Pinus rigide Mill. Bot. Gaz, § Brown, H. P., loc. cit. 992 KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH What factors operate to cause growth inception in a particular part of a tree? Robt. Hartig* believed that temperature was the most important factor, consequently insolation, temperature of the air and of the soil moisture, and thickness of the bark are the essential factors which determine the region of first diameter increase. No doubt these factors are important, as considerable evidence indicates that in old trees diameter increase is delayed at the base of trunk where insolation is poor and the bark is thick. In young trees, however, these are not the only factors. In the larch trees of 13 years of age the diameter increase did not begin first in those regions with the thinnest bark and best insolation. The thickness of the bark at the apex, middle and base of the tree A was respectively 596 u, 1,937 wu, and 3,278 uw. So also in the isolated tree E the inception of diameter increase did not occur in the parts of the tree best insolated and with the thinnest bark, but rather in the thicker barked and more poorly insolated parts of the tree, namely the middle and basal regions of the trunk. In the individual branches, however, growth in diameter began in the - regions of thinnest bark and of best insolation. | Whittenf has shown that the color of the bark may be impor- tant in the time of growth inception of buds. The color of the bark may be a factor in determining the region of diameter increase in young trees of larch. The color of the bark of the apical part of the trunk in spring is yellowish to greenish, becoming darker towards the base. The darker color, because of its capacity for heat absorption, may counteract the insulating effect of the thick bark, consequently the diameter increase begins in the basal and middle regions. The growth begins first in the middle regions because the bark here is of the same color as that of the basal regions and is only about half as thick. The fact that in the branches and twigs of larch the opening of buds on the apical regions is retarded is suggestive of the influence of the color of the bark. The bark of the apical regions being of lighter color less heat is absorbed, its temperature therefore is lower and the devel- opment of the buds is slower. This is in agreement with the * Hartig, R., loc } Whitten, J. C. pe protection of the peach. Missouri Agr. Exp. Sta. Bull. 38: 140-164. 1897. KNUDSON: CAMBIUM DEVELOPMENT IN AMERICAN LARCH 293 results obtained by Whitten* on the date of blossoming of green- and purple-twigged peach trees. The temperature of the soil moisture and of the air, and the thickness and color of the bark are not the only factors operative in inducing diameter increase. Certainly as regards all of the above factors the branches are all more favored than the trunk of the tree, yet in the isolated tree E the diameter increase began first in the trunk. Food supply may be a factor in stimulating the cambium of the trunk to diameter increase, before there is develop- ment in the branches. It is not possible to state the influence of food supply. Controlled experimental work with this as well as with the other factors is necessary to give an idea of the influence of each in stimulating the cambium to activity. In conclusion it should be stated that considerably more work is necessary in order to establish the region of the tree in which cambium activity first begins. No doubt different conditions will be found for trees of the same species of different ages, and for trees of different species and genera. Investigations of this character are important because only by such studies can the factors be determined which stimulate cambium activity. It is essential also to determine the period and extent of phloem and xylem formation in trees and the duration of cambium activity. Much more work remains to be done along this line. Such work is of especial importance in fruit culture and in a subsequent paper the results of such an investigation will be presented. CORNELL UNIVERSITY. Explanation of figures Fic. tr. Cutting no. 6 made from trunk on November 13, showing cambium tissue a to a, consisting of six rows of cells; X475. Fic. 2. Shows extent of cambium in basal cutting, No. 10, made on May 5. Compare with Fig. 2; 475 Fic. 3. Cutting no. : taken from trunk of tree A on May 5. Note the increased diameter and cell differentiation; 475. f Fic. 4. Cutting no. 5 made on May 25 from tree A. Xylem formation not yet begun; X33. Fic. 5. Cutting no. 4 from tree C made on June 3; X33- Fic. 6. Cutting no. 7 from tree C made on June 30; Fic. 7. Cutting no. 8 from tree C made on July 8. wood is beginning; Cambium a to a. XK33- Differentiation of summer X33: aS ee * Whitten, J. C., loc. cit. 3 Ramee apace id? ee ; Bes ie es : mere INDEX TO AMERICAN BOTANICAL LITERATURE (1910-1913) The aim of this Index is to include all current botanical literature written by Americans, published in America, or based upon American material ; the word Amer- ica being used in the broadest sense. Reviews, and papers that relate exclusively to forestry, egriculture, es cena manufactured products of vegetable origin, or laboratory methods are not i ed, and no attempt is made to index the literature of bacteriology. An ‘soli: cence is made in favor of some paper appearing in an American periodical which is devoted wholly to botany. Reprints are not men aa unless they differ from the original in some important particular. If users of the Index will call the attention of the editor *o errors or omissions, their kindness will be appreciated. This Index is reprinted monthly on cards, and ‘famished i in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitted ; each Bondoc must sae all cards published during the term of his subscription, Corre- pondence relating to the card issue should be addressed to the Treasurer of the Torrey hasan Club, Abrams, L. The gymnosperms growing on the grounds of Leland Stanford Jr. University. Dudley Memorial Volume (Stanford Univ. Publ.) 81-110. 1913. [Illust.] Albert, F. El verdadero pino Oregon. Pseudotsuga taxtfolia. Bol. Bosques, Pesca i Caza 1: 444-461. f. 1-14. Ja 1913. Alsberg, C. L., & Black, O. F. Contributions to the study of maize deterioration. U.S. Dept. Agr. Plant Ind. Bull. 270: 5-48. pl. 1. 11 Mr oe a Biochem and Ererscaccsic investigations of Penicillium puberulum and Penicillium acai Bartlett, H. H. PEO studies on Oenothera,—II. The delimi- tation of Oenothera biennis L. Rhodora 15: 48-53. pl. 102, 103. 12 Ap 1913. Bean, W. J. Pyrus ioensis. Curt. Bot. Mag. IV. 9: pl. 8488. Ap 1913. A plant from central United States. Beardslee, H.C. An acre of Lysurus. In Lloyd, C. G., Mycological notes 38: 515, 516. N 1912. ore Beardslee, H.C. A much named agaric. In Lloyd, C. G., Mycological notes 38: 524. f. 519. N 1912. Berger, A. Cereus Bridgesii S.-D. Monats. Kakteenk. 23: 44, 45. 15 Mr 1913. : 295 296 INDEX TO AMERICAN BOTANICAL LITERATURE Blakeslee, A. F., & Jarvis, C.D. Treesin winter. Their study, plant- ing, care and identification. 1-446. New York. 1913. [Illust.] Bédeker, F. Uber einige Coryphanthen und deren Bliiten. Monats. Kakteenk. 23: 45-47. 15 Mr 1913. Britton, E. G. Wild plants needing protection. 6. “Wild Azalea’”’ (Azalea nudiflora L.). Jour. N. Y. Bot. Gard. 14: 79-81. pl. 114. Ap 1913. Britton, N. L., & Rose, J. N. ae in Cactaceae—I. Contr. U. S. Nat. Herb. 16: 239-242. pl. 66-73. 10 Ap 1913. Includes new species in epee (2), Epiphyllum (1), Hylocereus (1), Nyc- tocereus (1), Opuntia (1), and Witiia (1). Brown, H.B. Studies in the development of Xylaria. Ann. Myc. 11: 1-13. pl. 7, 2. 15 Mr 1913. Brown, W.H. The relation of the substratum to the growth of Elodea. Philipp. Jour. Sci. 8: (Bot.) 1-20. F 1913. Brown, W. H., & Graff, P. W. Factors influencing fungus succession on dung cultures. Philipp. Jour. Sci. 8: (Bot.) 21-29. F 1913. Bunzel, H. H. A biochemical study of the curly-top of sugar beets. U.S. Dept. Agr. Plant Ind. Bull. 277: 5-28. 13 Mr 1913. Campbell, D. H. The morphology and systematic position of Caly- cularia radiculosa (Steph.). Dudley Memorial Volume (Stanford (Univ. Publ.), 43-61. f. 1-72. 1913. Campbell, D. H. William Russel Dudley. ‘Dudley Memorial Volume (Stanford Univ. Publ.) 11-15. 1913. Cannon, W. A. Some relations between salt plants and salt-spots. Dudley Memorial Volume (Stanford Univ. Publ.) 123-129. 1913. Claassen, E. Cualoplaca pyracea (Ach.) Th. Fr., a crustaceous lichen on the sandstone sidewalks of east Cleveland, Ohio. Ohio Nat. 13: 99, 100. 25 Mr 1913. Cockerell, T. D. A. Fossil flowers and fruits. ty 75,76. Ap 1913. Includes descriptions of Sambucus Ellisiae and Phalaris (?) g Torreya 13: nov fe Cocks, R. S. Leguminosae of Louisiana. Bull. 1: v-vi+1-26. S 10910. [Illust.] Collins,G. N. A variety of maize with silks maturing before the tassels. U.S. Dept. Agr. Plant Ind. Circ. 107: 3-11. f. 1-3. 7 F 1913. Collins, G. N., & Kempton, J. H. Inheritance of waxy endosperm in hybrids with sweet corn. U.S. Dept. Agr. Plant Ind. Cire. 120: 21-27. f.1r. § Ap 1913. : Cook, O. F. The abortion of fruiting branches in cotton. U: S. Dept. Agr. Plant Ind. Circ. 118: 11-16. 22 Mr 1913. Louisiana Nat. Hist. Surv. INDEX TO AMERICAN BOTANICAL LITERATURE 297 Cook, O. F. Durango cotton in the Imperial Valley. U. S. Dept. Agr. Plant Ind. Circ. 111: 11-22. f. 1-5. 1 F 1913. Cook, O. F. Leaf-cut, or tomosis, a disorder of cotton seedlings. U.S. Dept. Agr. Plant Ind. Circ. 120: 29-34. f. 1. 5 Ap 1913. Cook, O. F. A new ornamental palmetto in southern Texas. U. S. Dept. Agr. Plant Ind. Circ. 113: 11-14. 15 F 1913. Inodes exul Cook. Cook, O. F. Wild wheat in Palestine. U. S. Dept. Agr. Plant Ind. Bull. 274: 5-56. pl. 1-15. f. I-11. 3 Ap 1913. Daines, L. L. Comparative development of the cystocarps of Anti- thamnion and Prionitis. Univ. Calif. Pub. Bot. 4: 283-302. pl. 32- egg. SE Air por: Dewey, L.H. A purple-leaved mutation in hemp. U.S. Dept. Agr. Plant. Ind. Circ. 113: 23, 24. 15 F 1913. Dillman, A. C. Grasses for canal banks in western South Dakota. U. S. Dept. Agr. Plant. Ind. Circ. 115: 23-31. f. z, 2. 1 Mr 1913. Dudley, W.R. The vitality of the Sequoia gigantea. Dudley Memorial Volume (Stanford Univ. Publ.) 33-42. 1913. Dutton, D. L. Additions to the lichen flora of Vermont. Bull. Vermont Bot. Club 8: 16,17. Ap 1913. East, E. M. Inheritance of flower size in crosses between species of Nicotiana. Bot. Gaz. 55: 177-188. pl. 6-10. 15 Mr 1913. Eckerson, S. A physiological and chemical study of after-ripening. Bot. Gaz. 55: 286-299. 15 Ap 1913. Eggleston, W. W. A trip across Vermont by Luigi Castigilioni, an Italian nobleman, in August, 1785. Bull. Vermont Bot. Club 8: 21,22. Ap 1913; Estee, L. M. Fungus galls on Cystoseira and Halidrys. Univ. Calif. Pub. Bot. 4: 305-316. pl. 35. 31 Mr 1913. Includes Guignardia irritans Setchell & Estee, sp. nov. Fairchild, D. Some Asiatic Actinidias.. U. S$. Dept. Agr. Plant Ind. Circ, 110: 7-12. pl. r, 2+f. 1, 2. 18 Ja 1913. Fairchild, D., & Simmonds, E. The grafted papaya as an annual fruit tree. U.S. Dept. Agr. Plant Ind. Circ. 149: 3-13. f. 1-4. 29 Mr 1913. Fawcett, W. New plants from Jamaica.—II. Ap 191 Includes Coccoloba Priorii, C. nigra, and C. neglecta spp. nov. Fernald, M. L. Some noteworthy varieties of Bidens. 74-78. 23 Ap 1913. Fernald, M. L. Some North avnextbatt relatives of Polygonum mariti- mum. Rhodora 15: 68-73. 23 Ap 1913. Jour. Bot. 51: 123-125. Rhodora 15: 298 INDEX TO AMERICAN BOTANICAL LITERATURE Fernald, M. L., & Wiegand, K. M. A northern variety of Erigeron ramosus. Rhodora 15: 59-61. 12 Ap 1913. Erigeron ramosus septentrionalis var. nov. Gates, R. R. A new QOenothera. Rhodora 15: 45-48. pl. 100, I0T. Mr 1913 Oenothera angustissima sp. nov. Gates, R.R. Tetraploid mutants and chromosome mechanisms. Biol. Centralb. 33: 92-99. 20 F 1913; 33: 113-150. f. 1-7. 20 Mr 1913. Goodrich, L. L. H. Flora of Onendaga County as collected by the members of the Syracuse Botanical Club. 1-210. Syracuse, 1912. [Ilust.] Goodspeed, T. H. On the partial sterility of Nicotiana hybrids made with N. sylvestris asa parent. Univ. Calif. Publ. Bot. 5: 189-198. 21 Mr 1913. ; Harger, E. B. Some plants of the Southbury Triassic area. Rhodora 15: 65-68. 23 Ap 1913. Hawkins, L.A. The effect of certain chlorides singly and combined in pairs on the activity of malt diastase. Bot. Gaz. 55: 265-285. 15 Ap 1913. Hawkins, L. A. Experiments in the control of grape anthracnose. U.S. Dept. Agr. Plant Ind. Circ. 105: 3-8. pl. 1, 2. 10 F 1913 Hemenway, A. F. Studies on the phloem of the dicotyledons—II. The evolution of the sieve tube. Bot. Gaz. 55: 236-243. pl. II. +f. 1-3. 15 Mr 1913. Higgins, B. B. The perfect stage of Cylindosporium on Prunus avium. Science II. 37: 637, 638. 25 Ap 1913. The perfect stage is named Coccomyces hiemalis sp. nov. Hoffman, C. Paraffin blocks for growing seedlings in liquid culture solutions. Bot. Gaz. 55: 244-248. f. 1-3. 15 Mr 1913. Holden, R. Cretaceous Pityoxyla from Cliffwood, New Jersey. Proc. Am. Acad. Arts & Sci. 48: 609-624. pl. 1-4. Mr 1913. Includes Pinus protoscleriopitys, Pityoxylon foliosum, and P. anomalum, spp. nov. Howe, I. A. New botanical finds for St. Johnsbury. Bull. Vermont Bot. Club. 8: 15, 16. Ap 1913. Howe, R. H. A further note on the Linnean Herbarium. Torreya 13:77, 78. 6 Ap 1913. Hubbard, F.T. A Panicum unreported in New England. caveat u 15: 64. 12 Ap 1913. Kellerman, K. F. The excretion of cytase by Penicillium pinophilum. U.S. Dept. Agr. Plant Ind. Circ. 118: 29-31. f. z, 2. 22 Mr 1913. Kellerman, K. F. Soil bacteriology as a factor in crop production. U.S. Dept. Agr. Plant Ind. Circ. 113: 3-10. f. r. 15 F 1913. INDEX TO AMERICAN BOTANICAL LITERATURE 299 Kellerman, K. F. Testing cultures of nodule- forming bacteria. U.S. Dept. Agr. Plant Ind. Circ. 120: 3-5. f. 1. 5 Ap 1913. Kempton, J. H. Floral abnormalities in maize. U. S. Dept. Agr. Plant Ind. Bull. 278: 5-18. pl. 7, 2+f. 1, 2. 2 Ap 1913 Kirk, G. L. Some West Haven plants. Bull. Vermont Bot: Club 8: 47; 18. Ap 19613: Knowlton, F. H. The fossil forests of Arizona. Am. Forestry 19: 207-218. Ap 1933. Kunze, R.E. LEchinocactus Wislizeni Engel. and Echinocactus Lecontei Engel. Torreya 13: 73-75. 6 Ap 1913. Lawrence, W.H. Bluestem of the black raspberry. Washington Agr. Exp. Sta. Bull. 108: 3-30. f. 1-49. O 1912. Caused by Acrostolagmus caulophagus sp. nov. Lawrence, W. H. Plant diseases induced by Sclerotinia perplexa nov. sp. Washington Agr. Exp. Sta. Bull. 107: 3-22. f. I-9. O 1912. Lloyd, C. G. Mycological Notes 38: 510-524. f. 510-519. N 1912. [Illust.] Lyon, T. L., & Bizzel, J. A. The influence of alfalfa and of timothy on the production of nitrates in soils. Centralb. Bakt. Zweite Abt. 37: 161-167. 29 Mr 1913. Maublanc, A. Sur une maladie des feuilles du papayer ‘Carica Papaya.” A Lavoura 16: 208-212. 1913. [Illust.] sed by Sphaerella Caricae (Speg.) Maublanc, the ascigerous stage of Cer- cospora Caricae Speg. The same paper is published also in Portuguese under the title ‘‘Sobre uma molestia do mamoeiro (Caryca Papayal, L.),’’ 204-208. May, J.B. A teratological specimen of Cypripedium acaule. Rhodora 15: 73,74. 23 Ap 1913. McAtee, W.L. A list of plants collected on St. Vincent Island, Florida. Proc. Biol. Soc. Washington 26: 39-52. 22 Mr 1913. McBeth, I. G., & Scales, F. M. The destruction of cellulose by bacteria and filamentous fungi. U.S. Dept. Agr. Plant Ind. Bull. 266: 9-52. pl. 1-4. 21 F 1913. : McKee, R., & Ricker, P. L. Nonperennial Medicagos: the agronomic value and botanical relationship of the species. U.S. Dept. Agr. Plant Ind. Bull. 267: 5-38. pl. 1-13. 21 F 1913. : ; McMurphy, J. The Synchytria in the vicinity of Stanford University. Dudley Memorial Volume (Stanford Univ. Publ.) 111-114. pl. 1, 2. 1913. Includes Synchytrium Amsinckiae sp. nov. _ Meade, R. M. Methods of securing self-pollination in cotton. U. S. Dept. Agr. Plant Ind. Circ. 121: 29, 30. f. 1. 12 Ap 1913. 300 INDEX TO AMERICAN BOTANICAL LITERATURE Meade, R. M. Supernumerary carpels in cotton bolls. U.S. Dept. Agr. Plant Ind. Circ. 111: 25-28. f. 1, 2. 1 F 1913. Merrill, E. D. A flora of Manila. Bur. Sci. Manila Pub. 5: 1-490. 31 D 1912. Merrill, E. D. Studies on Philippine Rubiaceae, I. Philipp. Jour. Sci. 7: (Bot.) 31-62. pl. rz. F 1913. Thirty-five new species are described. Meyer, R. Echinopsis calochlora K. Sch. Monats. Kakteenk. 23: 33, 34. 415 Mr 1913. Montgomery, E. G. Experiments in wheat breeding: experimental error in the nursery and variation in nitrogen and yield. U. S. Dept. Agr. Plant Ind. Bull. 269: 5-61. pl. i-4+f. 1-22. 24 Ap 1913. Nash, G.V. Thecedar of Lebanon. Jour. N. Y. Bot. Gard. 14: 86-89. pl. 175. Ap 1913. Newcombe, F.C. The scope and method of state natural history surveys. Science II. 37: 615-622. 25 Ap 1913. Nichols, G. E. A simple revolving table for standardizing porous cup atmometers. Bot. Gaz. 55: 249-251. f. 74. 15 Mr 1913. Northup, Z. The influence of certain acid-destroying cee sa lactic bacteria. Mich. Agr. Exp. Sta. Bull. 15: 3-35. Je Oakley, R. A., & Garver, S. Two types of ie SE in alfalfa. U.S. Dept. Agr. Plant Ind. Circ. 115: 3-13. f. 1-8. 1 Mr 1913. Orton, W. A. Powdery dry-rot of the potato. U. S, Dept. Agr. Plant Ind. Cire. 110: 13-15. 18 Ja 1913. Otis, C. H. Michigan trees. A handbook of the native and most important introduced species. Michigan Univ. Bull. 14: i-xxxii + 1-246. Mr 1913. [Illust.] Includes a short introduction by G. P. Burns. Parish, S.B. The California Paroselas. 15 Ap 1913 Includes Parosela neglecta sp. nov. Bot. Gaz. 55: 300-313. f. 1-5: Peirce, G. J. Studies of irritability in plants.—III. Volume (Stanford Univ. Publ.) 62-80. f. 14, Pierce, N. B. A new walnut. Juglans quercifolia sp. nov. Pool, V. W., & McKay, M. B. The control of the sugar-beet leaf-spot. U. S. Dept. Agr. Plant Ind. Cire. 121: 13-17. 12 Ap 1913. Quehl, L. Mamillaria echinoidea Quehl spec. nov. Monats. Kakteenk. 23: 42, 43. 15 Mr 1913. {Illust.] Rangel, E. Grave molestia do coqueiro (Cocos sta i Ld Lavoura 16: 199-201. 1913. {Ilust.] Dudley Memorial 1913. Science II. 37: 613, 614. 18 Ap 1913- INDEX TO AMERICAN BOTANICAL LITERATURE 301 Rigg, G. B. The effect of some Puget Sound bog waters on the root hairs of Tradescantia. Bot. Gaz. 55: 314-326. 15 Ap 1913. Roberts, J.W. The ‘“‘rough bark” disease of the yellow Newton apple. U.S. Dept. Agr. Plant Ind. Bull. 280: 5-16. pl. 1-3+f. 1, 2. 12 Ap 1913. : A disease caused by Phomopsis Mali. Rood, A. N. Juncus monostichus in Ohio. Rhodora 15: 62. 12 Ap 1913. Saccardo, P. A. Notae mycologicae. Ann. Myc. 11: 14-21. 15 Mr Includes Aecidium zonatum, A. Theveliae, ea uberata, Macrophoma mexicana, and Didymopsis phyllogena, spp. nov Mexico. Sargent, F. L. Plants and their uses. i introduction to botany. i-x+1-610. f. 1-384. New York, 1913. Seeley, H. M. Some features of the dandelion. Bull. Vermont Bot. Club 8: 13, 14. Ap 1913. An abstract. Setchell, W. A. Mushrooms and _ toadstools. Univ, Calif. Agr. Exp. Sta. Circ. 84: 1-4. Ja 1913. Shantz, H. L. The effects of artificial shading on plant growth in Louisiana. U.S. Dept. Agr. Plant Ind. Bull. 279: 5-31. pl. 1-6 +f. I-Iz. 16 Ap 1913. Sharp, K. D. Summer ina bog. 1-149. Cincinnati, 1913. Shear, C. L., & Wood, A. K. Studies of fungous parasites belonging to the genus Glomerella. U.S. Dept. Agr. Plant Ind. Bull. 252: 5-110. pl. 1-18 +f. I-4. 25 Ja 1913. Slosson, M. The stag-horn ferns. Jour. N. Y. Bot. Gard. 14: 63-67. pl. 112, 113. Mr 1913. Small, J. K. The genus Malpighia in Jamaica. Torreya 13:77- Ap Malpighia Harrisiit Small sp. nov. described. Jour. N. Y. Small, J. KK. Report on exploration in tropical Florida. Bot. Gard. 14: 81-86. 6 Ap I913. Smith, M. V. (Mrs. C. H.) An old-time herbarium. Bull. Vermont Bot. Club 8: 19-21. Ap 1913. An abstract. : Speare, A. T., & Colley, R. H. The artificial use of the brown-tail fungus in Massachusetts, with practical suggestions for private experiment, and a brief note on a fungous disease of the gypsy caterpillar, 5-31. pl. r-8+f. 1, 2. Boston, 1912. Sprague, T. A. Hypericum Kalmianum. Curt. Bot. Mag. IV. 9: pl. 8491. Ap 1913. 302 INDEX TO AMERICAN BOTANICAL LITERATURE Starr, A.M. Poisoning by Ginkgo. Bot. Gaz. 55: 251. 15 Mr 1913. Steele, E.S. An investigation of ‘‘Laciniaria scariosa.” Torreya 13: 78,79. 6 Ap 1913. Stone, G. E. Alist of plants growing without cultivation in Franklin, Hampshire and Hampden Counties, Massachusetts. i-vii-+1-72. Amherst, 1913. Stone, G. E. Magnolia tripetala in Springfield, Massachusetts. Rho- dora 15: 63. 12 Ap 1913. Sturgis, W. C. The Myxomycetes of Colorado.—II. Colorado Col. Publ. Sci. 12: 435-454. pl. 2. Ap 1913. Includes Fuligo megaspora, Didymium anomalum, and Enerthenema syncarpon, spp- nov. Sudworth, G. B. Forest atlas. Geographic distribution of North American trees. Part 1—Pines. Maps 1-36. 1913. U.S. Dept. Agr. Forest Service Publication. Taylor, N. A plant new to the state of New York and the local flora range. eesti 13:78. 6 Ap 1953. Adoxa Moschatellina Underwood, J. G. Notes on flora of Stratton, Franklin, Berkshire, Newport, Island Pond, and Hartland. Bull. Vermont Bot. Club 8: 8-12. Ap 1913. Underwood, Pe G. Phlox diverienia in Vermont. Rhodora 15: 79. 23 AplI Underwood, “ G. Plants new to Nistiadll. collected in 1912. Bull. % Vermont Bot. Club 8: 22, 23. Ap 1913. Victorin, —. Notes on the occurrence of interesting forms of Cyperaceae in Quebec. Ottawa Nat. 27:15, 16. 21 Ap 1913. Wagner, E. Allerlei aus dem Kakteenkasten. 43, 44. 15 Mr 1913. Wakeman, N. The Monardas. A phytochemical study. Bull. Univ. Wisconsin Sci. Ser. 4: 81-128. Au 1911. _Also reprinted with separate pagination. Monats. Kakteenk. 23: Wellington, R. Studies of natural and artificial parthenogenesis in the genus Nicotiana. Am. Nat. 47: 279-306. My 1913. Wheeler, L. A. Additions to the flora of West River Valley in 1912. Bull. Vermont Bot. Club 8: 14, 15. Ap 1913. An abstract. White, O. E. The bearing of teratological development in Nicotiana on theories of heredity. Am. Nat. 47: 206-228. f. 1, 2. Ap 1913- White, T.H. Tomato variations induced by culture. Maryland Agric. Exp. Sta. Bull. 173: 121-133. f. 1-5. Ja 1913. INDEX TO AMERICAN BOTANICAL LITERATURE 303 Wight, W. F. North American species of the genus Amygdalus. Dudley Memorial Volume (Stanford Univ. Publ.) 130-137. 1913. Includes Amygdalus Harvardii sp. nov. Wilcox, E. M., Link, G. K. K., & Pool, V. W. A dry rot of the Irish potato tuber. Nebraska Agr. Exp. Sta. Research Bull. 1: 5-88. pl. 1-28+f. 1-15. 1 Mr 1913. Williams, A. Carnivorous plants of Ohio. Mr 1913. Ohio Nat. 13: 97-99. 25 TORRE VIOLA PEDATIFIDA XSAGITTATA AND LEAVES OF ITS PARENTS AND OF NINE OFFSPRING Buy. Torrey Cius VOLUME 40, PLATE t7 V. sororia V.pedatifida SPRING LEAVES OF VIOLA PEDATIFIDA XSORORIA, OF ITS PARENTS, AND OF NINE OFF But. Torrey CLus VOLUME 40, PLATE 18 KNUDSON: CAMBIUM IN THE AMERICAN LARCH VOLUME 40, PLATE 19 But. TorREY CLUB Msn tree A peste without conspicuous lateral veins beneath. Vernonia angustata. 2. Leaves broadest near or above the middle, veiny. Vernonia gnaphaliifolia. The status of the first three species in the key above is still open to question and may require future adjustment. Gardner’s type locality for V. araripensis was in Brazil, and it is scarcely probable that the same species occurs in the Antilles, especially as far as Cuba or Santo Domingo. Authentic specimens of Lessing’s V. stenophylla have not been seen, and the plants referred to his name differ in some slight features from the original - description. Both species are accepted solely on the authority of certain European students of the genus. GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 309 Vernonia corallophila sp. nov. Stem erect, herbaceous, 4 dm. high, virgate, finely striate, strigose-pubescent; leaves sessile, narrowly linear, rigid, revolute, one-nerved, I-3 cm. long, strigose-hispid and punctate above, densely strigose-pubescent beneath; inflorescence cylindrical or sparingly branched; heads 11-flowered, in the axils of the upper leaves or rarely 2 or 3 together on a short ascending lateral cyme; bracteal leaves resembling the cauline, but reduced in size and the upper barely exceeding the heads; involucre turbinate, 5-6 mm. high, its scales all straight, erect, very loosely imbricated, strigose-pubescent, the outer subulate, the inner narrowly oblong, long-acuminate; achenes pubescent, 2 mm. long; pappus nearly white, the inner series 4-5 mm. long. Type, Britton 1939, from a coral limestone beach at the United - States Naval Station, Guantanamo Bay, Oriente, Cuba, March 17-30, 1909, deposited in the Herbarium of the New York Botanical Garden. The species is readily distinguished by its slender virgate inflorescence and small leaves from V. stenophylla with its long leaves and divaricate inflorescence. Vernonia angustata sp. nov. Vernonia sublanata angustata Gleason, Bull. N. Y. Bot. Gard. 4: 177. 1906. Herbaceous, height not stated; stems slender, puberulent or thinly pubescent, sparingly branched above; leaves thin, flat, spreading, narrowly oblong with parallel sides, 3-5 cm. long, 4-8 mm. wide, obtuse or rounded, mucronate, entire, acute at the reduced in size, and the upper only 1 cm. long; heads sessile, distinctly secund, about 18-flowered; involucre broadly campanu- late, 6-7 mm. high, its scales loosely and irregularly imbricated, strigose-pubescent, acuminate or sharply acute, the outer lanceo- - late, the inner narrowly oblong; achenes pubescent, 2 mm. long;. pappus white, the inner series 6 mm. long. 310 GLEASON: STUDIES ON WEST INDIAN VERNONIEAE VERNONIA GNAPHALIIFOLIA Rich. in Sagra, Hist. Fis. Pol. Nat. Cuba 11: 34. 1850 Vernonia sublanata Gleason, Bull. N. Y. Bot. Gard. 4: 177. 1906. Dr. Urban has established the identity of these two species by the examination of authentic material, and they are accordingly united here. The species appears to be common and widely distributed in Cuba from the province of Havana eastward. SPECIES-GROUP DIVARICATAE The history of this group of Vernonieae begins with the publication of V. divaricata by Swartz in 1806. This was followed in 1831 by Lessing’s V. acuminata, and no further addition was made to the group until 1906, when Gleason described V. albicoma and V. expansa. During the last half-century V. divaricata and V. acuminata have been usually considered identical. The whole group, so far as known, is confined to Jamaica, and the ample collection in the Herbarium of the New York Botanical Garden includes seven distinct species. At the present time the chief difficulty lies in determining to which of these seven species the old names divaricata and acuminata belong. Certain characters given by the authors in their original descriptions serve to exclude one or another of the species, and by this process of exclusion it is possible to arrive finally at a reasonable conclusion. It may be safely affirmed that the two species described below are the only ones examined to which these names can consistently be given, without doing violence to some important feature of the original description. VERNONIA DIVARICATA Sw. FI. Ind. Occ. 3: 1319. 1806 A straggling shrub, 1-2 m. high; leaves spreading, thin, bright green, elliptical or elliptical-lanceolate, 4-7 cm. long, 1.5-2.5 cm. wide, acute or subacuminate, entire, narrowed to an acute base, minutely puberulent above, thinly pubescent and finely punctate with minute pellucid glands beneath, sessile or with petioles 1-5 mm. long; inflorescence of numerous, lax, loosely flowered, di- varicate cymes, bearing each 5-15 secund heads, and frequently prolonged into a leafy shoot; bracteal leaves oblanceolate to narrowly oblong, barely exceeding the 11—13-flowered heads; in- volucre campanulate, about 4 mm. high; scales thinly strigose- GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 311 pubescent, the inner oblong, gradually narrowed to a rounded tip, the outer triangular-ovate, obtuse or acute. The following specimens are referred here: Jamaica: Harris 8205, altitude 500 m.; Britton & Hollick 1996, altitude 700 m.; Britton 431, 679; Cockerell. - VERNONIA ALBICOMA Gleason, Bull. N. Y. Bot. Gard. 4: 185. 1906 m woody, slender; leaves thin, bright green, elliptical or elliptical-lanceolate, 5-6 em. long, 1.5-2.3 cm. wide, acuminate, subentire, tapering at the base, glabrate above, very finely pubes- cent or glabrous beneath; petioles 5~7 mm. long; upper leaves somewhat reduced; inflorescence of 1-3 curved spreading terminal cymes bearing each 5-10 sessile secund 18-flowered heads; bracteal leaves lance-oblong, the middle ones about equaling the involucre, which is broadly campanulate or hemispheric, 5 mm. high; scales erect, loosely and irregularly imbricated, nearly glabrous, the outer lanceolate, acuminate, the inner oblong, acute; pappus white. Campbell 6152, from the foot of Long Mountain, Jamaica, alti- tude 115 m. VERNONIA ACUMINATA Less. Linnaea 6: 663. 1831 Shrubby, height not stated; stem finely cinereous-puberulent; leaves thin, bright green, broadly ovate-lanceolate, 6-9 cm. long and half as wide, long-acuminate, entire, acute at the base, very sparsely and minutely puberulent on both sides, on petioles about mm. long; inflorescence of numerous widely spreading freely branched cymes, the ultimate branches bearing 3-7 heads; rameal leaves lanceolate, 2-3 cm. long; bracts oblong-lanceolate to linear, those at the ends of the cymes only 6 mm. long; heads 18-flowered ; involucre campanulate, about 5 mm. high, its scales all obtuse or the outer acute, pubescent, especially the outer series, the exposed portion of the inner series oblong. Wight 20, from wooded hillsides, Port Antonio, Jamaica. VERNONIA EXPANSA Gleason, Bull. N. Y. Bot. Gard. 4: 186. 1906 A straggling shrub 2.5-3 m. high, branching above; leaves numerous, rather crowded, thin, divaricate, ovate, 3-5 cm. long, I.7-2.7 cm. wide, acute or subacuminate, entire, obtuse or broadly rounded at base, sparsely and finely pubescent, or nearly glabrous ~ above; petioles 3 mm. long; upper and rameal leaves smaller, broadly elliptical, abtuse; inflorescence of several short cymes bearing each 3-6 heads; bracteal leaves equaling or much shorter 312 GLEASON: STUDIES ON WEST INDIAN VERNONIEAE than the involucre; heads crowded, sessile, 11-flowered; involucre broadly campanulate, 4 mm. high; scales all appressed, closely imbricated, glabrous or nearly so, rounded at the tip, the outer broadly ovate, the exposed portion of the inner ovate. Jamaica: Harris 8796, near Troy, altitude 2,000 ft., Britton & Hollick 1956, woods, Bluefields Mountain, altitude 750 m. Vernonia pluvialis sp. nov. Apparently erect and branched only near the top, height 4 m. or more; stem finely striate, glabrate, or thinly pubescent, especially above; leaves spreading, or usually ascending, firm and rigid, oblong-ovate to subrhomboid, broadest conspicuously above the middle, the principal ones 3-5 cm. long by 1.1—1.9 cm. wide, acute, entire and subrevolute, gradually narrowed to an obtuse base, minutely puberulent on both sides and glandular-punctate below; veins not prominent, sessile or on petioles 1-2 mm. long; inflorescence of several short few-headed cymes in the axils of the upper leaves, forming a compact leafy panicle; bracteal leaves barely exceeding the heads, ovate to ovate-lanceolate; heads crowded in clusters of 2-7, or sometimes single, sessile, not at all secund, 8- (rarely 5-) flowered; involucre narrowly campanulate or subcylindric, 5-7 mm. high, its scales puberulent, well imbri- cated in several ranks, the outer ovate-triangular, acute or apicu- late, the inner becoming oblong and slightly narrowed to an obtuse or rounded tip; achenes densely hirsute, not mature in the specimens examined; inner pappus pale brown, 4-5 mm. long on the immature achenes. Type collected by Forrest Shreve on Blue Mountain Peak, Jamaica, May 14, 1906, and deposited in the Herbarium of the New York Botanical Garden. Other sheets in the same herbarium are Nichols 20, from moist woods, Morce’s Gap, altitude 5,000 feet, Nichols 120, from the summit of Blue Mountain Peak, and E. G. Britton 3856, from Sir John Peak. Vernonia proclivis sp. nov. ‘Shrubby, about 2 m. high; stem sparingly branched, finely striate, glabrate or thinly puberulent; cauline leave sconspicuously — reflexed, bright green above, pale green beneath, thin, elliptic- oblong, widest at or near the middle, 6-8 cm. long by 2-3 cm. wide, ' distinctly acuminate, entire, gradually narrowed or subacuminate at base, glabrous and finely glandular-punctate with transparent globules above and minutely black-punctate beneath; veins promi- GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 313 nent beneath; petioles 3-4 mm. long; rameal and bracteal leaves similar, but much smaller; inflorescence cylindrical or pyramidal, composed of numerous short, divaricate, irregularly branched cymes, forming a terminal panicle, and bearing each 4-10 sessile, crowded, 8-flowered heads near their tips; involucre narrowly campanulate, 4-5 mm. high, its scales erect, regularly imbricated in several ranks, thinly puberulent or glabrate, the outer ovate, with acute or apiculate tips, the inner oblong, slightly narrowed to a rounded tip; flowers purple to pale lavender; immature achenes densely pubescent; outer pappus pale brown, 0.4 mm. long, the inner brown, 4 mm. lon Type, Britton 102, from Morce’s Gap, near Cinchona, Jamaica, September 2-10, 1906, in the Herbarium of the New York Botan- ical Garden. Three other sheets in the same herbarium agree with it perfectly and are referred to the same species: Marble 188, from Cinchona, E. G. Britton 3856, from the Blue Mountains, and Britton 4055, from the Parish of St. Thomas. The latter was collected in March; the achenes are all discharged and the dry and brown involucre is widely open. Vernonia reducta sp. nov. Shrubby, freely branched above, height not stated; stem striate, glabrate below, becoming puberulent in the inflorescence; leaves spreading, firm, bright green above, paler beneath, narrowly elliptical-oblong or somewhat oblanceolate, broadest at or near the middle, the principal ones 4-4.5 cm. long by 1.2-1.6 cm. wide, sharply acute or subacuminate, entire or finely denticulate, gradually narrowed at the base, minutely puberulent on both sides and finely glandular-punctate beneath; petioles 2-5 mm. long; cymes numerous, terminating the stem and the upper branches, and forming a large loose pyramidal panicle; rameal and bracteal leaves resembling the cauline, but becoming narrower and smaller toward the ends of the branches, and finally barely exceeding the heads; heads crowded in clusters of 2-7, sessile or nearly so, not at all secund, 5-flowered; involucre narrowly campanulate or subcylindric, 5-6 mm. high, its scales puberulent on the back and regularly imbricated in several ranks, outer scales triangular, acute, and somewhat arachnoid-ciliate, the inner be- coming oblong-linear, puberulent or glabrate, narrowed to an obtuse apex; achenes about 2 mm. long, hirsute; pappus rufescent, the outer series 0.5 mm., the inner 4-5 mm. long. Type, Britton 203, from Sir John Peak, Jamaica, September 314 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 2-10, 1906, in the Herbarium of the New York Botanical Garden. Two other sheets in the same herbarium are Britton 151, from New Haven Gap, and Shreve, from Sir John Peak. The seven species of the group may be distinguished by the following key: A. Heads more or less secund, 11—18-flowered; involucre cam- panulate to hemispheric, distinctly spreading when press-dried; its scales loosely and irregularly imbri- cated in fe I. Principal leaves about 3 times as long as broad. a. Heads 11—13-flowered; pappus brown. Vernonia divaricata. b. Heads 18-flowered; pappus white. Vernonia albicoma. 2. Principal leaves twice as long as broad, or less a. seesnisigs broadly ovate-lanceolate, distinctly acu- nate; exposed portion of inner involucral ic oblong; heads 18-flowered; cymes freely branched. Vernonia acuminata. b, Leaves broadly ovate, acute or subacuminate; ex- posed portion of inner involucral scales ovate; heads r1-flowered; cymes sparingly branched. Vernonia expansa. B. Heads not at all secund, 5-8-flow wered; involucre narrowly Dp dried; its begs rather uniformly imbricated in several ran 1. Leaves sprea rs or ascending, nearly or quite sessile, distinctly oblong-obovate or subrhomboidal, 30-50 m. long, acute; heads 8- (rarely 5-) flowered. Vernonia pluvialis, 2. Leaves proportionately narrower or larger, “nauiess near the middle, more or less acuminate a. Leaves elliptical-oblong, about 25 X65 mm., con- spicuously reflexed; heads 8-flowered. Vernonia proclivis. b. Leaves narrowly elliptical-oblong, about 14 X45 mm., spreading; heads 5-flowered. Vernonia reducta. SPECIES-GROUP FRUTICOSAE When a sheet of Santo Domingan material came to hand, identified by Dr. Urban as authentic V. fruticosa Sw., it was at once seen that the species was entirely distinct from V. rigida and from the whole group to which V. rigida belongs. It was also seen that its nearest congeners were to be found among some un- described species from eastern Cuba, with which it is accordingly grouped. As in several other species-groups, the chief similarity between the species included is in the general habit. The most GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 315 striking common feature here is the broad, rugose or buHate leaf with reticulate venation, closely invested beneath by a dense tomentum. In the Jamaican species-group Permolles, with simi- larly bullate or rugose leaves, the pubescence is densely sericeous- hirsute rather than closely tomentose, and in V. yunquensis the character of the involucral scales separates the species at once. The seven species may be distinguished as follows: I, Santo Domingan species; leaves oblong-ovate, truncate or su eens at base, obtuse, repand; bracteal leaves not ced; bgey: very flexuous. Vernonia fruticosa, TT; Pitts speci A. Inner in ae scales obtuse or rounded; leaves ovate-lanceolat B. Inner involucral scales sharply acute or acuminate. eaves oval to slice obtuse or narrowed at the base, rugose and finely papillose- pubescent above. a. Leaves more than half as broad as long, very lunt or rounded at the apex; straggling plant with white flowers. Vernonia calophylia. b. Leaves less than half as broad as long, nar- rowed to an obtuse or subacute apex; plant erect, with light blue flowers. Vernonia vicina. 2. Leaves of a lanceolate or ovate type, broadest near t neate or subcordate base, strongly bullate and rugose above. a. Leaves acute or barely obtuse, 1 coceopsesite beneath; inflorescence loo n- gated; inner involucral ec broadest just below the middle. Vernonia neglecta. b. Leaves obtuse or rounded; inflorescence com- pact; cymes abbreviated; inner invo- lucral scales broadest near the base. * Leaves ovate-oblong, crenate or repand, brown-tomentose beneath; involucre densely pubescent; inner scales merely Vernonia desiliens. i acute. Vernonia calida. ** Leaves broadly ovate or oval, entire or somew pubescent or glabrate; inner scales sharply acuminate. Vernonia semitalis. VERNONIA FRUTICOSA (L.) Sw. FI. Ind. Occ. 3: 1323. 1806. Stem very slender, freely and loosely branched, glabrous or puberulént in the inflorescence; leaves firm, ovate-oblong, 1-2 cm 316 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE long by half as wide, obtuse or rounded at the tip, irregularly crenulate or subentire, rounded or subcordate at base, dark green and thinly but softly pubescent above, densely gray-tomentose beneath; petioles 1-2 mm. long; upper and terminal branches floriferous and strongly flexuous; bracteal leaves resembling the cauline in shape and size; heads single, sessile, 21-flowered ; involucre broadly turbinate or campanulate, 5 mm. high, its scales all erect or somewhat spreading, very loosely imbricated, puberu- lent or glabrous, the outer subulate, the inner narrowly lanceolate and long-acuminate with a subulate tip; achenes hirsute; pappus nearly white, 4 mm. long. For the identification of this obscure species of Swartz we are indebted to Dr. Urban. The description above is based on a single sheet, Fuertes 655, from the Province of Barahona, Santo Domingo, at 350 m. altitude. Vernonia desiliens sp. nov. Herbaceous, sparingly branched, 3-5 dm. tall; stem rather prominently striate, glabrous below, becoming puberulent in and near the inflorescence; leaves firm and coriaceous, ovate-lanceolate to narrowly elliptical, the largest on the type sheet 8—8.5 cm. long and 2.3-2.9 cm. wide, obtuse, entire or obscurely and shallowly crenulate, obtuse or rounded at the base, with petioles 1-3 mm. long, rugose, glabrous and shining above; veins elevated beneath and prominently reticulated, the principal lateral ones strongly ascending and almost parallel to the leaf-margin; the interstices of the reticulations closely gray-tomentose; cymes apparently one or two, terminal and from the upper axils, spreading, 1-2 dm. long, very flexuous; bracteal leaves resembling the cauline but narrower in shape and gradually reduced in size, oblong or narrowly ellip- tical, 3-5 cm. long, 5-12 mm. wide, spreading, those at the ex- tremity of the cyme oblong-linear, 2 cm. long; heads sessile, single in the axils, 1-2 cm. apart, 21-flowered; corollas purplish; in- volucre turbinate, thinly pubescent, 9-10 mm. high; the lower outer scales short, minute, triangular, acute and apiculate, and closely imbricated; the inner scales much longer, linear-oblong, obtuse; achenes strigose-pubescent; pappus light brown, the inner series 6 mm. long, minutely barbellate, the outer series 0.5 mm. long, somewhat paler. Type, Shafer 3232, growing among rocks near water, Arroyo del Medio, Oriente, Cuba, at 450-550 m. altitude, January 20, 1910, deposited in the Herbarium of the New York Botanical Garden- GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 317 Vernonia calophylla sp. nov. A straggling shrub, 1 m. high or less; stem slender, striate, freely and loosely branched. cinereous-puberulent below, becoming tomentose in the branches; leaves firm and rigid, ovate to sub- rotund, the principal ones 20-25 mm. long by 13-18 mm. wide, rounded or obtuse at the apex, entire, somewhat revolute, rounded or subcordate at base, dark green, rugose, and papillose-pubescent above, closely invested beneath with silver-gray tomentum; veins conspicuous, the lateral ones ascending and the veinlets promi- nently reticulated; petioles 1-2 mm. long; heads in the axils of the upper leaves, forming cymes 10-16 cm. long, sessile, 18-21- flowered; bracteal leaves resembling the cauline, but smaller, I—1.5 cm. long and more densely pubescent above; corollas white; involucre campanulate, 5 mm. high; scales somewhat arachnoid- puberulent, especially near their tips, the outer ones narrowly triangular, subulate, the inner linear-oblong, acute; achenes pubescent, 2 mm. long; pappus nearly white, the outer series 0.8 mm., the inner 4-mm. long. Type, Shafer 8102, from Camp La Gloria, south of Sierra Moa, Oriente, Cuba, December 24-30, 1910, deposited in the Herbarium of the New York Botanical Garden. Vernonia vicina sp. nov. An upright shrub, 3-6 dm. tall, sparingly branched; stem finely sleet glabrate below, puberulent above, and in the branches closely cinereous-tomentulose; leaves firm and rigid, spreading, dark green above, elliptic to elliptic-oblong, the principal ones 3-4.5 cm. long by 1-1.5 cm. wide, broadest at or near the middle, narrowed to an obtuse or subacute tip and to an acute _ base, entire, somewhat revolute, rugose above, papillose-pubescent when young, becoming glabrate with age, closely invested beneath with thin yellowish brown or cinereous tomentum, especially between the veins; lateral veins ascending, but curved and ap- proximate near the margin; veinlets prominently reticulated ; petioles curved, 1-2 mm. long, tomentose; upper leaves like the lower, but smaller, eventually only 1 cm. long by 4 mm. wide, bearing heads in the upper 4-5 axils; heads about 26-flowered; flowers light blue, involucre campanulate, 5-6 mm. high, its scales closely imbricated, glabrous or thinly puberulent, outer scales subulate, somewhat spreading, inner scales lance-oblong, sharply acute; achenes pubescent, 2 mm. long; outer eee white, 0.8 mm. long; inner series almost white, 4 mm. long. Type, Shafer 8202, from Camp La Gloria, south of Sierra Moa, 318 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE Oriente, Cuba, December 24-30, 1910, deposited in the Herbarium of the New York Botanical Garden. Vernonia neglecta nom. nov. V..Wrightit Griseb. Cat. Pl. Cuba 144. 1866. Not V. Wrightii Sch.-Bip. 1863. Vernonia gnaphaliifolia Gleason, Bull. N. Y. Bot. Gard. 4: 178. 1906. Not V. gnaphaliifolia Rich. 1850. Apparently erect and suffruticose, sparingly branched, 6-15 dm. high; stem and branches slender, obscurely striate, finely gray-tomentulose; leaves firm, flat, spreading, narrowly ovate- oblong to ovate-lanceolate, the largest on the type sheet 6—7 cm. long by 2 cm. wide, acute, or subobtuse and minutely apiculate, entire, rounded, truncate, or even subcordate at base, dark green and bullate above, papillose-puberulent when young, becoming glabrous and shining at maturity, softly gray-tomentose or almost villous beneath, especially along the elevated, prominently reticu- somewhat scarious on the margin; achenes pubescent, 1.5 mm. long; outer pappus white, 0.9 mm. long, the inner series very pale _ brown or almost white, 4 mm. long. Type, Wright 1309, on banks of cliffs near Monte Verde, eastern Cuba, deposited in the Gray Herbarium of Harvard University. The specimen bears two labels, one dated “Dec. 27,” the other “Jan.—Jul. 1859.” On the left-hand side of the same sheet is another specimen of the same species, also numbered 1309, and collected in eastern Cuba, Sept. 1859-Jan. 1860. Another sheet of the same species in the Gray Herbarium is Wright 2788: Vernonia calida sp. nov. : Shrubby, freely branched, 5 dm. tall; stem obscurely striate, thinly puberulent or finely cinereous-tomentulose in the inflores- cence; leaves spreading, thick, rigid, ovate-oblong, the principal GLEASON; STUDIES ON WEsT INDIAN VERNONIEAE 319 ones 3.5-5 cm. long and 1.5-2 cm. wide, obtuse or rounded at the tip, crenate or repand, broadly rounded or subcordate at base, strongly bullate, dark green, minutely papillose-pubescent when young, soon becoming glabrate or scabrellate and shining above, densely and softly brown-tomentose beneath; petioles 1-3 mm. long; lateral veins prominent, strongly curved and soon confluent; veinlets prominently reticulated; cymes few, simple or sparingly branched, 6-12 cm. long, the rachis densely tomentose; bracteal leaves resembling the cauline in shape, but smaller, crenulate or entire, the upper I-1.5 cm. long; heads rather crowded, single or frequently two at each node, about 21-flowered, separated by inter- nodes I-2 cm. long; corollas pink; involucre broadly campanulate, mm. high; scales densely pubescent, closely imbricated, or somewhat spreading at the tip, the outer subulate, the inner narrowly triangular-lanceolate and acute; achenes pubescent, 1.5 mm. long; pappus yellowish brown, the outer series 0.7 mm., the inner 5 mm. in length. Type, Shafer 8408, in dry soil, Sabanilla to Yamuri Arriba, Oriente, Cuba, January 30, February 1, 1911, deposited in the Herbarium of the New York Botanical Garden. Vernonia semitalis sp. nov. Shrubby, 6-9 dm. tall, freely branched above; stem striate, leafy, thinly brown-tomentose, especially on the younger branches; eaves numerous and crowded, thick, rigid, somewhat revolute, divaricately spreading, ovate or ovate-triangular, broadest near the base, 1.5-2 cm. long by I-1.3 cm. wide, obtuse or rounded at the apex, entire, truncate or subcordate at base; upper surface shining, glabrous or scabrellate, strongly bullate; lower surface closely invested with a thin gray-green tomentum; veins elevated beneath, the lateral ones ascending and confluent near the margin; veinlets prominently reticulated; upper leaves resembling the lower ones and scarcely reduced in size, bearing heads in their axils and forming several crowded cymes 10-15 cm. long; heads about 21-flowered, secund, the lower separated by internodes I cm. long, the upper approximate; corollas white; involucre cam- panulate, about 5-6 mm. high; outer scales triangular-subulate and pubescent, the inner narrowly triangular, sharply acuminate, glabrous or nearly so; achenes pubescent, 1.5 mm. long; pappus _ nearly white, the outer series 0.7 mm., the inner 4 mm. long. Type, Shafer 4176, from pine land, altitude 400 m., along the trail from Rio Yamaniguey to Camp Toa, Oriente, Cuba, February 22-26, 1910, deposited in the Herbarium of the New York Botan- ical Garden. 320 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE SPECIES-GROUP SAGRAEANAE This group is distinguished at once from other West Indian species by the large glabrous achenes. The leaves also are usually thick and firm or coriaceous, entire or with spinulose teeth. Most of the species have been in the past poorly represented in American herbaria, and some of them have been seldom collected since their original discovery. In the revision, Vernonia rigida Sw. and Vernonia fruticosa (L.) Sw. were regarded as identical and referred to this group, to which the name Rigidae was applied. Since that time, specimens of V. fruticosa have again been collected, and the species is seen to belong to a different group. The Jamaican V. rigida, also, is described with pubescent achenes, a character which removes it at once from this group. In 1836 De Candolle described V. Sagraeana from Cuba, the first known species of the group. This was followed in 1850 by V. Valenzuelana of Richard. In 1863 Schultz examined the recent Cuban collections of Wright, and added three species, leptoclada, inaequiserrata and Wrightii, and a fourth, Sprengeliana, based on a plant collected by Bertero in Santo Domingo. Grisebach added a variety, inaequiserrata angustifolia, also collected by Wright in Cuba. That left the group with seven species and one variety: and, so far as known to the writer, no authentic collection of any of these was made or at least recognized for forty years. Further difficulty was added by the confusion of numbers of some of Wright’s collections, so that at least two different species have masqueraded in herbaria under wrong names. One case of this confusion was recognized in 1906 by Gleason, who remedied it by the description of V. viminalis. Since 1906, the collectors of the New York Botanical Garden, in their diligent explorations of Cuba, have recollected four of these old, imperfectly known species, and have added three entirely new forms, which are here described. The group as a whole is one of the most easily recognized of all the West Indian species. It is characterized especially by a high involucre and by large, glabrous, obscurely ribbed achenes, with a prominent basal callus, and the large, firm or rigid leaves. GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 321 I, Species of Santo Domingo; leaves oblong, very tomen- subcordate base; inner involucral scales rounded and apiculate at the tip. - Vernonia Sprengeliana, II. Species of Cub A. Leaves ca coriaceous, more or less revolute and hining above, oblong or Bids tes ces a aaie dentate or entir Heads wered; sateen cylindric. Vernonia purpurata. . Heads ak 18 flowers or more; involucre cam- panulate a. Leaves narrowly oblong, at least three times as fie as broad, ipa or with numerous minute spinulose teeth. Vernonia Valenzuelana. b. Leaves Coca oblong, a twice as long as Ass entire, or with a few LS barely spreading, the inner acute, : e outer mucronate. Vernonia leptoclada, ** Outer scales conspicuously squarrose or reflexed, all sharply acuminate or subulate Vernonia Wrightit. B. Leaves thin or firm, bat coriaceous, flat, serrate, ptisadedicatas, or entire I. Leaves puberulent or glabrous beneath. a. Outer scales of the involucre short, acute or mucronulate, appressed. Vernonia Sagraeana, b. Outer scales o e involucre elongated, subulate, pape Vernonia aronifolia. ae Leaves, Laie beneath. involu et chia sharply acumi- e, densely e. Vernonia viminalis. b. ae opoee As gem rounded to a mucronate apex, or acute. * er scales obtuse or subacute; muddle scales without scarious margin; leaves entire or obscurely spinulosedentiulate Vernonia fallax. ** Inner scales shsaiates the middle with a scarious margin. ° Leaves narrowly linear-oblong, entire. Vernonia aceratotdes. °° Leaves narrowly elliptic-oblong, sharply serrate. Vernonia inaequiserrata. Vernonia Sprengeliana Sch.-Bip. The type specimen cited by Schultz is Bertero 507 and the recent description by Gleason is : 322 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE based ona single sheet of Wright, Parry, & Brummel 273. An excellent collection of this rare species, Fuertes 1388, has been recently distributed and agrees perfectly with the original descrip-. tion and with that of Gleason (Revision, 184). Vernonia purpurata sp. nov. Shrubby, 2-2.5 m. tall; stem stout, coarsely striate, thinly tomentose below, becoming densely so in the inflorescence; leaves crowded, heavy, rigid, coriaceous, divaricate, elliptic-oblong, ob- tuse or subacute, entire or irregularly repand, obtuse or rounded at base, strongly rugose above, but glabrous and shining except for some thin pubescence along the midvein, minutely puberulent along the veins beneath; veins elevated on the lower surface, the lateral veins prominent, ascending, the veinlets small and closely reticulated; petiole 2-4 mm. long, tomentose; inflorescence small, irregular, composed of several short (2-6 cm.) leafy cymes, bearing each 4-10 heads; rameal leaves resembling the cauline, but two thirds as long; bracteal leaves narrowly oblong or oblong- linear, 10-15 mm. long, not present below many of the heads; heads sessile, secund along the cymes or aggregated at their tips, 8-flowered; corollas white; involucre narrowly cylindric, 6 mm. high; scales closely imbricated, appressed, sharply acute, the lower ovate-triangular, pubescent, the middle ones with a ovate-triangular exposed portion, ciliate, glabrous on the back, the inner entire, puberulent on the back, purple-brown at their exposed tips; achenes glabrous, immature in the type specimen; pappus pale yellow-brown, the outer series 1.3 mm., the inner 7 mm. long. Type, Taylor 544, from Jiquarito Mountain, Sierra Maestro, eastern Cuba, altitude 1,020 m., September 18, 1906, deposited in the Herbarium of the New York Botanical Garden. The lower leaves are lacking from the type plant. The crowded upper leaves are remarkably uniform in size, 4-5 cm. long by I.5-I.9 mm. wide. On another branch is the base of a leaf which measures 3 cm. wide, indicating that the lower leaves are considerably larger than the upper. While the species certainly belongs to this group, it is distinguished ae all the others known by its few-flowered heads. Vernonia Valenzuelana Rich. A shrub 1.2 m. high, on dry ferruginous soil, southeast of Paso Estancia, Oriente, Cuba, Shafer 1705. GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 323 Vernonia leptoclada Sch.-Bip. A specimen agreeing perfectly with the original description has been recently collected by Shafer, 8145, from Camp La Gloria, south of Sierra Moa, Oriente, Cuba. It is described as a straggling shrub 3.5 m. tall, with white flowers. Vernonia Wrightit Sch.-Bip. Ex descr. A shrub, 1 m. high, freely branched above; stems pubescent; leaves heavy and coria- ceous, spreading or ascending, ovate-oblong to oblong, 3.5 cm. long by 1.2 cm. wide, sharply acute or mucronate, revolute, entire or with a few remote spinose teeth, rounded or even subcordate at the base, very scabrous above but not pubescent, minutely puberulent under the lens beneath; petioles 1-2 mm. long, pubescent; cymes long and spreading, with numerous heads; bracteal leaves like the cauline, but gradually reduced to 1 cm. long; heads sessile, secund, about 21-flowered; involucre about 7-8 mm. high, outer scales ovate, acuminate into a squarrose or recurved tip, the inner erect or somewhat spreading, sharply acute or subulate; pappus pale brown. Shafer 7738, from a dry serpentine hill near El Yunque, Oriente, and Shafer 3072, from pine lands at 500-650 m. altitude near Woodfred, Oriente, are referred here. They agree in most essential points with Schultz’ description of V. Wrightw (Journ. Bot. 1: 234. 1863), but lack the glabrous leaves narrowed at both ends and the sordid-purple pappus. Schultz’ species is based on Wright 1309. As is well known, the Wright numbers are much confused, and frequently shelter more than one species. The only available specimen of this number belongs to an entirely different species. Vernonia Sagraeana DC. A shrub 1.5 m. tall, growing on banks at an altitude of 325 m., near El Cuero, Oriente, Britton & Cowell 12704. Vernonia aronifolia sp. nov. - 324 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE mm. long, standing at right angles to the margin, obtuse or sub- acute at base, minutely puberulent beneath, especially along the veins, glabrous above; veins prominent, light green, the lateral ones arcuate-ascending, branched and reticulated, especially toward the margin, one of the smaller marginal veins prolonged into each denticulation; cymes terminal and lateral, spreading, unbranched, 15-20 cm. long, very leafy, bearing 6—10 heads; bracteal leaves resembling the cauline in shape and texture, or varying to broadly oblong-elliptic in shape; the lower approximat- ing the cauline in size, the upper gradually reduced to half the size, but always greatly exceeding the flowers; heads sessile, single in the axils, about 34—47-flowered; corollas white; involucre hemispherical or broadly campanulate, 8-9 mm. high, in dried specimens about 15 mm. broad; scales thinly puberulent, im- bricated only at the base, straight or erect in bud, becoming spreading or flexed in fruit, the outer lance-linear, subulate, the inner linear-oblong, acuminate to a subulate tip; achenes smooth; outer pappus white, 0.8 mm. long, inner series pale brown or nearly white, 8 mm. long, minutely barbellate. Type, Shafer 13514, collected from high rocks in limestone hills, vicinity of Sumidero, Province of Pinar del Rio, Cuba, August 2, 4, 1912, deposited in the Herbarium of the New York Botanical Garden. The collection is represented by two sheets, one includ- ing apparently the top of the plant, and the other two detached lateral branches. Vernonia fallax sp. nov. Shrubby, I m. high; stem erect, sparingly branched, finely striate, closely gray-tomentose, especially above; leaves firm, reticulated; petioles 1-2 mm. long; inflorescence pyramidal, ter- minal, of about 4-10 short, spreading or recurved cymes bearing each 3-7 heads; heads secund, sessile, about 21-flowered; involucre broadly campanulate or subhemispherical, 6-7 mm. high, its scales appressed, closely imbricated, pubescent, especially near the tip, outer scales ovate-triangular, acute and cuspidate, the inner subacute or rounded at the tip; outer pappus white, 0.7 mm. long, the inner pappus very pale brown, 6 mm. long; achenes glabrous. GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 325 Type, Britton & Wilson 5478, from a hillside, altitude 500 m., in the Trinidad Mountains, Province of Santa Clara, Cuba, March 12, 1910, deposited in the Herbarium of the New York Botanical Garden. The bracteal leaves have all fallen off the type specimen, except a few fragments, and the leaves are crowded on short lateral branches. The inflorescence is thus left at the end of a naked peduncle 2-3 dm. long, giving the specimen an aspect entirely unlike other species of the group. It is scarcely to be expected that the same peculiarity will be maintained in other collections of the species. Vernonia aceratoides sp. nov. Vernonia inaequiserrata angustifolia Griseb. Cat. Pl. Cuba 144. 1866. Slender and probably herbaceous; stem finely striate, closely gray-tomentulose; leaves firm, spreading or ascending, narrowly oblong-lanceolate or lance-linear, the principal ones 7~8 cm. long and I-1.2 cm. wide, acute and mucronulate at the tip, entire or somewhat repand; obtuse or rounded at the base, minutely scabrellate above and puberulent along the midvein, finely brown- tomentulose beneath; veins prominent beneath and conspicu- ously reticulated; petioles 2-3 mm. long; inflorescence terminal, of about 3 short divaricately spreading cymes, bearing each six or seven secund heads; bracteal leaves oblong, the upper ones not exceeding the heads, and all proportionately broader than the cauline; involucre narrowly campanulate, 5-6 mm. high; scales closely and regularly imbricated, appressed, the outer ovate- triangular, cuspidate, the inner with an ovate exposed portion, rounded and apiculate at the tip. Grisebach’s variety was based on a specimen of Wright 2784; the preceding more detailed description is based on a sheet of the same number in the Herbarium of the Missouri Botanical Garden. SPECIES-GROUP LONGIFOLIAE The herbarium of Dr. Otto Kuntze contained a good specimen of a Vernonia from St. Thomas, collected by Kuntze himself in 1874, and labeled Vernonia Thomae Benth. It can not be dis- tinguished, however, in any essential character from Vernonia albicaulis Pers., and the two species may henceforth be considered identical. This disposition of V. Thomae was suggested before by Gleason (Revision, 191), although at that time the two were 326 GLEASON: STUDIES ON WEST INDIAN VERNONIEAE kept separate since good material for examination was lacking. The island of St. Thomas is accordingly added to the known distribution of V. albicaulis. It has been collected in St. Thomas by others also, and is represented in the herbarium of the Field Museum by two sheets, Millspaugh 522 and Eggers 34. Vernonia longifolia Pers. To the distribution of this species may be added St. Martin, Boldingh 2641, and Montserrat, Shafer 172, 589, 659, O61. Lepidaploa, Scorpioideae reductae Vernonia arctata Gleason. The species was originally described (Bull. Torrey Club 33: 185. 1906) from New Providence Island, of the Bahama group, but is now known to occur also throughout Andros Island, Small & Carter 8506, 8613, 8759, 8890, Brace 5170, 6754, 6926, 7138. Field data show that the flowers vary from purplish white to bright rose-purple and that the plant reaches a height of 2 meters. Vernonia bahamensis Griseb. Reported by Gleason (Bull. Torrey Club 33: 187. 1906) from Fortune Island and Inagua, it is now represented also by specimens from Crooked Island, Brace 4851; Acklin’s Island, Brace 4330; Salt Cay, Millspaugh & Millspaugh 9249; Long Cay, Brace 4152, 4020, 4115; Mariguana, Wilson 7461; Castle Island, Wilson 7783; Cotton Cay, Mills- paugh & Millspaugh 9362; North Caicos, Wilson 7721, Millspaugh & Miullspaugh 9175; East Caicos, Millspaugh & Millspaugh 9082; and South Caicos, Wilson 7688. The last specimen cited has leaves subacuminate or merely acute at the base, 5 cm. long by 2.5 cm. wide, and in general closely approximates V. albicaulis Pers. The Scorpioideae reductae have been considered (Gleason, Revi- sion, 165, 166) as related by origin to the species-group Longifoliae, to which V. albicaulis belongs, and the theory is strengthened by the strong superficial resemblance just mentioned. It is interest- ing to note that V. bahamensis occupies the southeastern portion of the Bahama archipelago, nearest the area of the Longifoliae, and that the particular specimen comes from South Caicos, which is almost the extreme southeastern island of the group. Vernonia complicata Griseb. In the type collection, Wright 2790, the leaves are all entire, subrotund, and about 5 mm. aie GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 327 An excellent specimen, Britton 2225, recently received at the New York Botanical Garden from Guantanamo Bay, in extreme eastern Cuba, has leaves of the same character on the old shoots, while on the young branches they are flat or undulate and 10-15 mm. long. The Guantanamo plant is described as a shrub 1 m. tall with purple flowers. Lepidaploa, Scorpioideae aggregatae Vernonia Thomae Benth., included here by Gleason (Revision, 191), is now regarded as identical with Vernonia albicaulis Pers. Urban (Symb. Antill. 7: 421. 1912) has recently added a species, so that the number in the group remains four. They may be distinguished as follows: A. Achenes pubescent; outer pappus conspicuous, it I ht 1 tk the white bristles than of the inner series; leaves 2-3 cm. long. Vernonia buxifolia (Cass.) Less, B. Achenes glabrous and glandular; outer pappus minute, its scales not sharpl istin- guished in width from those of the inner series Vernonia Tuerckheimii Urban. 2. Leaves minutely puberulent or glabrous h; pappus yellowish or tawny; heads 8-flowered. Vernonia montana Gleason. C. Achenes densely hirsute; outer pappus conspic- uous, its scales chaffy and fimbriate; leaves 4-5 cm. long, closely gray-tomentose, aud with prominent veins beneath. The first three species are all very similar in habit and structure, and are all natives of Hispaniola. There is no doubt that they are closely related. The character of the pappus and the larger leaves indicate that V. buxifolia is the primitive form. The dis- tribution (Cuba) and the general habit of V. yunquensis, especially of the leaves, as expressed in pubescence and venation, separate it sharply from the first three species, and imply that it may logically constitute another species-group. Vernonia segregata sp. nov. ___ A straggling or vinelike shrub, reaching a height of 2.5 ™.; stem obscurely striate, closely pubescent; the branches olive- Vernonia yunquensis Gleason. 328 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE brown and finely tomentulose; leaves numerous, crowded, firm, dark green, oblong to elliptic-obovate, broadest at or above the middle, the principal ones 2.5-4 cm. long and I-1.5 cm. wide, obtuse or subacute, entire, obtuse or rounded at the base, scabrel- late and resinous-punctate above, glabrous beneath and densely punctate with resinous globules and impressed black glands; midvein puberulent, the lateral veins inconspicuous; petioles I-2 mm. long; inflorescence terminal, irregular in shape, consisting of several short, simple or sparingly branched cymes 2-4 cm. long, naked below, and bearing 2-8 crowded heads in a terminal sub- capitate cluster, or at the base of the branches; bracteal leaves I—3 subtending each cluster of heads, resembling the cauline in shape, 5-15 mm. long; heads about 8-flowered; corollas white; involucre campanulate, 3-4 mm. high, its scales loosely and ir- regularly imbricated, appressed at the base, but spreading at the tip, stiff and firm in texture, the outer narrowly triangular-lanceo- late, long-acuminate, the inner narrowly oblong-linear, tapering gradually to the acuminate puberulent apex; achenes thinly pubescent, 2 mm. long; pappus nearly white, the outer series I mm., the inner 4 mm. long. Type, Shafer 4050, from rocky river banks in the vicinity of Camp San Benito, Oriente, Cuba, altitude 900 m., February 24, 1910, deposited in the Herbarium of the New York Botanical Garden. Other sheets in the same herbarium, all collected in the mountains of Oriente, are Shafer 8051, stated to be vinelike and 8 feet high; Shafer 8216, 1.5-2 feet high; and Shafer 4446, described by the collector as an herb three feet high with purple flowers. Notwithstanding field differences in the color of flowers or texture of stem, all four numbers clearly belong to the same species. The relationship of V. segregata is puzzling. The subcapitate clusters clearly represent a modification of a scorpioid type, and most closely resemble the inflorescence of the Scorpioideae aggre- gatae. The involucre is quite different, however, from that of typical members of the group. For the present, it has been con- sidered advisable not to assign the species to any group. SPECIES-GROUP HAVANENSES In recent work on Vernonia, V. havanensis and V. Ottonis have been considered identical (Revision, 192). The large series of specimens now available for study permits the ready separation — of two species, with characters so typical that to each can be GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE 329 assigned the proper specific name without difficulty. In addition, a new species has been collected by Shafer and is described below. With the exception of V. pallescens, whose position in this group 1s somewhat uncertain, the group is distinguished by similari- ties in habit. The leaves are of a comparatively broad type, widest near or usually above the middle, and with the serration most prominent on the distal half. The involucre scales are regularly pubescent in two areas on the back, one on each side of the mid-nerve. The four species may be distinguished as follows: A. —— strictly scorpioid; the cymes many-headed ongated; heads all sessile. Vernonia pallescens. B. Aeeiatiat freely branched and subpaniculate, some of the heads pedicellate; leaves with a tendency to be broadest above the middle; scales glandular on the back, e. I. Heads with 18 flowers or more; involucres 5-8 mm. high, or some of the scales 10 mm. long, distinctly purple-tinged; leaves essentially glabrous on both sides; pappus white, or with a faint brownish yellow Vernonia havanensis. 4 Fairey Ore ene, involucres 3-4 mm. high, ob- scurely or not at all tinged with purple; leaves scabrous above; pappus pale brown a. Heads 11~13-flowered; inner ia obtuse or subacute; inflorescence divaricate. Vernonia Oitonis. b. Heads 5-flowered, aggregated in subcapitate clusters at the ends of the branches, forming a pyramid or subhemispheric inflorescence; inner scales acute. Vernonia Orientis. Vernonia pallescens Gleason. The species certainly differs phylogenetically from the rest of the group, as shown by its inflorescence and its geographical distribution. It is included in the group merely for lack of a better place to put it. VERNONIA HAVANENSIS DC. Prodr. 5: 37- 1836. Vernonia stictophylla Wright, Sauv. Anal. Acad. Ci. Habana 6: 176. 1869. The specimens at hand fall into two groups, the first with leaves long-attenuate at base, almost sessile, and thin in texture; the second with leaves cuneate into a distinct petiole and firm in texture. No other characters for their separation have been 330 GLEASON: STUDIES ON WEST INDIAN VERNONIEAE found. The first group includes the type collection for Wright’s species. Both series have been collected, so far as data are given, in the province of Pinar del Rio. VERNONIA OTTONIS Sch.-Bip. Linnaea 20: 508. 1847. Vernonia hieracioides Griseb. Mem. Am. Acad. 8: 511. 1860. Vernonia cubensis Griseb. Cat. Pl. Cuba 144. 1866. Except one collection from Santa Clara (Leon 1315) and one from the Isle of Pines (Curtiss), all the sheets examined are from Pinar del Rio. They show considerable variation in the pubes- cence, serration, and texture of the leaves, but can not be further separated. The specimens include cotypes of both Grisebach’s species, and agree perfectly with Schultz’ description. Vernonia orientis sp. nov. Shrubby, as much as 6 m. in height, apparently not extensively branched; stem coarsely striate, glabrate below, becoming cinere- ous-puberulent in the inflorescence; leaves rigid, dark green, spreading, oblanceolate, the principal ones 9-11 cm. long by 2.5-3.-5 cm. wide, abruptly short-acuminate or sharply acute, remotely dentate with sharp salient teeth, chiefly above the middle, attenuate from below the middle to a cuneate base, very scabrous above, minutely puberulent and scabrellate beneath; veins elevat below, only the midvein and its lateral branches prominent; petioles 5-10 mm. long; inflorescence terminal, broadly pyramidal or subhemispheric; cymes freely branching, ultimately bearing 2-6 heads aggregated or subcapitate near the tips; bracts subulate, 3-5 mm. long; heads 5-flowered; involucre 3-4 mm. high, cam- panulate; scales ovate to ovate-oblong, sharply acute or sub- " acuminate, essentially glabrous but glandular on the back; achenes sparingly pecs outer pappus minute, the inner pale yellow- ish brown, 4 mm. Type, Shafer 3509, from Sierra Nipe, near Woodfred, Oriente, Cuba, altitude 450-550 m., January 10, 1910, deposited in the Herbarium of the New York Botanical Garden. The two collections, both from Oriente, are the only —s of the group from this part of the island. It is distinguished from the other members of the group at a glance by its inflorescence, and also by the involucral scales and the number of flowers. GLEASON: STUDIES ON WEstT INDIAN VERNONIEAE 331 Lepidaploa, Paniculatae dichotomae VERNONIA MENTHAEFOLIA (Pépp.) Less. Linnaea 4: 268. 1829. Eupatorium menthaefolium Pépp. in Spreng. Syst. 3: 412. 1826. Vernonia Grisebachii Sch.-Bip. Jour. Bot. 1: 231. 1863. | The original description of this species by Péppig is too brief to be of any value at the present time. But his specimens were preserved, and examined later by both Schultz and Lessing. Les- sing gives a detailed description, based on these types, stating that the heads are many-flowered and 3 lines high. Schultz’ descrip- tion, referring without doubt to the same specimens, or to dupli- - cates of them, indicates that the heads are 11-flowered and the involucre hardly 1 line high. He then described V. Grisebachii, as cited above, to include the forms with large heads, based on Wright 1305. Examination of an ample series of specimens at the present time reveals but one species, agreeing with Lessing’s and Schultz’ descriptions, but never with the small heads ascribed by the latter to V. menthaefolia. In the series examined are two of Wright's collections, 282 and 2792, and Shafer 8811, which was found by Dr. Britton to agree with the specimen of Wright 1305 in the Kew herbarium. Throughout the series the heads have II to 18 flowers, and the involucres are 4-5 mm. high. The leaves show considerable variation, from narrowly oblong-lanceo- late, acuminate at both ends, to ovate, rounded at the base and acute at the apex. These characters are not sufficiently definite or constant to permit the recognition of two species. V. menthaefolia is the most abundant species of the genus in Cuba, judged from the frequency of its collection, and occurs throughout the island. Among recent accession to the Herbarium of the New York Botanical Garden is an Eremosis from the state of Durango, which differs distinctly from all the fifteen described species of the genus. Eremosis ovata sp. nov. _ Shrubby; height and habit not stated; stem obscurely striate, closely cinereous-pubescent, becoming tomentulose in the inflores- cence; leaves thick, firm, ovate to ovate-elliptic, 7-10 cm. long, 4-5 cm. wide, obtuse or subacute, entire, obtuse at base, dull green, minutely and softly tomentulose above, densely cinereous- 332 GLEASON: STUDIES ON WEsT INDIAN VERNONIEAE tomentose beneath; veins elevated below, the lateral ones promi- nent and ascending, the veinlets inconspicuous; petiole 8-13 mm long; inflorescence broadly pyramidal or hemispheric, about 2 dm. wide; rameal leaves elliptic, about 2-6 cm. long, otherwise like the cauline; heads 4-flowered, in clusters of 3-8, on pedicels 2-5 mm. long; involucre narrowly campanulate, straw-colored or pale brown, 5-6 mm. high, outer scales short, broadly ovate, obtuse to subacute and apiculate, irregularly arachnoid or tomen- tulose, inner scales deciduous, oblong or ovate-oblong, acute, glabrous, or with minute patches of thin tomentum near the tip; achenes pale brown, 3 mm. long, prominently ribbed, thinly hirsute with oo hairs; pappus white, 8 mm. long, the outer series much shorter Type, Palmer 139, from San Ramon, Durango, Mexico, de- posited in the Herbarium of the New York Botanical Garden. In general habit and shape of leaf, Evemosis ovata most closely resembles Eremosis Steetzii (Sch.-Bip.) Gleason, but is distin- guished at once from this one-flowered species by its four-flowered heads. Its nearest relatives are probably to be found among the three-flowered species, such as Eremosis Palmeri (Rose) Gleason, from which it differs in the broad ovate leaves and dense tomentum. The presence regularly of four flowers in each head is a peculiar feature, hitherto unknown in the genus. It is paralleled in a way, however, by the occurrence of two flowers instead of one in the heads of certain specimens of Eremosis tarchonanthifolia (DC.) Gleason. UNIVERSITY OF MICHIGAN, ANN ARBOR. Some toxic and antitoxic effects in cultures of Spirogyra * W. D. Hoyt In view of difficulties commonly experienced in maintaining algal cultures in the laboratory, it seemed desirable to make an experimental study of some of these difficulties, as was suggested to the writer by Professor Georg Klebs, and the present publication deals with a portion of such a study. The experimentation was carried out in the Botanisches Institut at Heidelberg in 1909 and 1910, the facilities of these laboratories being made available through the kindness of Professor Klebs. The experiments to be considered below bear upon the preparation of a nutrient solution suitable for algal growth under laboratory conditions, and under- take a partial analysis of the relations of the aqueous aaa eas to the success or failure of algal cultures in glass. The alga used for this study was Spirogyra longata (Vauch.) Kg.,f brought from Algiers in the spring of 1906 and kept since then (until the winter of t909-10) in a north window of the Botanisches Institut at Heidelberg, with only small additions of tap water from time to time. At the time of beginning these studies the culture was healthy and showed vigorous growth, and the material seemed excellently suited for experimentation of the sort in hand; all the filaments had developed under closely similar conditions and were obviously adjusted to the conditions existing in the laboratory. In the experimentation it was thus possible to alter the culture medium without greatly changing the other environmental factors—e. g., light, temperature—under which the plants had developed. The last consideration is an important * Botanical Contribution from The Johns Hopkins pas No. 29. t Determined from Einfachste Lebensformen des Tier- und Pflanzenreiches, Eyferth, B., 3 Aufl. 1900. 333 334 Hoyt: CULTURES OF SPIROGYRA one; results obtained with material brought from various other conditions to the laboratory cannot be interpreted as caused by changes in the nature of the culture solution alone, a point well emphasized by some of the considerations to be brought out below. The method here followed was to transfer a few (about 30 to 50) filaments from the stock culture to about ten cubic centimeters of the medium to be tested, in a covered glass dish. These trans- fers were made late in the afternoon, and the cultures were sub- jected to microscopic examination, without removal from the culture dishes, on the next morning and on succeeding days until final results were obtained. The experiments lasted for periods of from one to eighty-six days. The condition of each culture was recorded in the following terms. Excellent means that, at most, only a few cells showed injury; good, that considerably more than half of the filaments seemed uninjured; poor, that more than half were injured; dead, that practically every cell was dead. The tables used in this paper have been constructed from the daily records by stating in similar terms the general condition of the cultures throughout the experiments. Precautions were of course taken to have the dishes and instru- ments as clean as possible. Any dish once used for a poisonous solution was discarded at the end of that experiment, and dishes were sometimes interchanged and furnished with fresh supplies of the media to be tested. The different lots of special distilled water did not enter into experiments until they had been found to be harmless to the plant here used. In most cases about the same amounts of media were used in the different experiments; the quantity. of solution as related to the number of cells in any culture is to be considered as an important factor in all experiments bearing upon the relation of the nature of the medium to the behavior of organisms existing therein. I. NUTRIENT SOLUTIONS The nutrient media used were all prepared according to the formulae given by Kiister (10), being those of Sachs, Knop,; Molisch, and Crone.* These were afterward diluted to the — * These solutions contained salts in the following proportions: Hoyt: CULTURES OF SPIROGYRA 335 required concentration with either ordinary or nontoxic distilled water. The reactions to litmus paper of the solutions thus pre- pared were as follows: Sachs’s, very slightly acid; Knop’s, acid; Molisch’s, slightly acid; Crone’s, neutral. The salts used were, in every case, those of Merck. The results obtained from the various cultures in these nutrient solutions are given in TABLE I, where each culture is denoted by a letter indicating the condition of the culture during the period of the experiment, the number of letters thus denoting, in each case, the number of times that the experiment in question was per- formed. The durations of the cultures are also given. The same notation is used in TABLES II, II, and v. Using nontoxic distilled water, prepared in a manner to be described later, the best growth was shown with all the media in concentrations containing from 0.05 to 0.1 per cent of total salts. In these concentrations of the solutions of both Molisch and Crone, Spirogyra was kept growing vigorously, in apparently perfect condition, for a period of about two months, at the end of which time the experimentation was discontinued. Crone’s solution seemed slightly more favorable to the organism than did that of Molisch; Sachs’ solution produced a fair growth; while Knop’s medium was distinctly unfavorable, probably because of the marked acidity, free acid being injurious to many algae (10). No doubt still other media might prove as favorable as those of Molisch and Crone. When ordinary distilled water was used in preparing the solu- tions a good growth was never obtained, but the results in both Molisch’s and Crone’s solutions seemed slightly better in con- centrations of from 0.5 to 1 per cent than in weaker solutions. This will be discussed later. The addition of one per cent of agar to the nutrient solutions, the Spirogyra filaments being laid on the surface of the coagulated — pinecone ay Sacus Knope MOLiscH CRONE I g. KNOs, 1 g. Ca(NOs)2, 1 g. (NH«)2HPOs, 1 g. KNOs, 0.5 g. NaCl, 0.25 g. KNOs, 0.5 g- KH2POs, 0.5 g. MgSO«, 0.5 g. CaSOx, 0.25 g. MgSOz, 0.5 g- MgSO, 0.5 g- CaSOx, 9.5 g. MgSOu, 0.25 g. KH2POx, 0.5 g. CaSOu, 0.25 g- Cas(POs)2, 0.5 g-CaHPO,s, 0.125 g. KCL, Trace FeSO.. 0.25 g. Fes(POs)s. Trace FeClo. Trace FeCl: 336 Hoyt: CULTURES OF SPIROGYRA medium in a covered Petri dish, produced slight improvement in all the 0.1 per cent solutions from ordinary distilled water. A good growth was obtained on similar agar plates prepared from 0.5 per cent solutions (in ordinary distilled water) according to the formula of Sachs, of Molisch, and of Crone.* Since the solutions here employed were all prepared from water and salts, it will be expedient to present the following considera- tions under the two headings: (II) Water and (III) Salts. II. WATER Many authors have found that tap water and distilled water are toxic to organisms, an excellent review of the literature of this subject being given by Livingston (12). Most workers have attributed the toxic effects of ordinary distilled water to small amounts of various metals—especially copper—taken by the water from the still and from the supply pipes. Lyon (19) and Bullot (6), however, found that water distilled in glass was markedly toxic, thus indicating that part of the poisonous action may be due to the presence of volatile substances. Lyon attributed this effect to small quantities of ammonia in the water. Similarly Livingston (12) showed that redistillation from glass to glass, while improving the quality of his ordinary distilled water, did not render it harmless. He obtained evidence indicating that part of the toxic bodies present in ordinary distilled water thus redistilled are volatile and reappear in the distillate, while part are nonvolatile and remain in the undistilled residue. The same writer prepared nontoxic water by shaking ordinary distilled water with highly absorbent solids, especially with purified lamp black, and then filtering out the solids. He concluded that the solids had absorbed, or at least removed from solution, both the volatile and the nonvolatile toxic substances. In the present investigation both tap water and ordinary dis- tilled water were found to be markedly toxic to Spirogyra, and were subjected to various treatments in an effort to obtain some evidence concerning the nature of the poisonous substances in- volved. The tap water was drawn from the faucet of the laboratory * Knop’s solution was so acid that 1 per cent agar would not harden ina 0.5 per cent concentration of the medium. Hoyt: CULTURES OF SPIROGYRA 337 supply, directly into the culture dishes. The distilled water used was from the usual supply of the laboratory, obtained from a local druggist. It had been distilled in a copper still and kept in glass. All such water used in these experiments was from the same original supply and may thus be considered as uniform. The results obtained in the study of the physiological properties of various waters are given in TABLES II and III. Tap water was usually fatal to the plant within a few days, although its toxicity varied slightly at different times (II, 1*). This water, when distilled with glass boiler and glass condenser, Was as toxic as the untreated water (II, 2). The distillate was then sampled at different times during the run, by placing culture dishes at the outlet of the condenser and allowing them to receive the water directly; the alga was introduced into these water samples after they had cooled. Samples collected at the beginning and end of the run were as toxic as the untreated water (II, 3, 3d). About the middle of the run the distillate collected was found to. be decidedly less injurious than the untreated tap water (II, 3a). Concentration of the tap water to one tenth its original volume by boiling (II, 4) and also by allowing evaporation to proceed at a temperature much below boiling, till only about a sixth of the original volume remained (IJ, 5, 6), produced marked improve- ment over the original untreated water, but concentration by evaporation to one third of the original volume did not improve its quality (II, 7). It will be noted, however, that the untreated water of this particular test was more toxic than in the two former experiments. In general, concentration seems to have improved the quality of the water, probably by driving off some of the volatile toxic substances, though it is of course not to be forgotten in this connection that the nonvolatile constituents were much more concentrated in the treated water than in the original. The effect of high temperature without concentration was also tested. Heating tap water in the autoclave for 15 minutes, at a temperature of 144° C., destroyed all the toxic material, or else rendered it nontoxic; in water thus treated Spirogyra filaments ee ee a * Roman numerals followed by Arabic, in parentheses, refer to table and experi- ment, respectively. 338 Hoyt: CULTURES OF SPIROGYRA water drawn from the tap at the same time, nearly all the fila- ments succumbed within a few hours (II, 8). Heating tap water to a temperature of 100° C. in a steam sterilizer for 45 minutes produced no improvement; the filaments of Spirogyra died almost as quickly in this as in untreated water (II, 9). Treatment of the tap water of the Heidelberg supply by the method of shaking with finely divided carbon, as employed with such marked effect by Livingston in the preparation of culture media from his ordinary distilled water, produced no improve- ment. In this treatment Merck’s ‘‘animal charcoal’? was em- ployed as absorbing solid, without preliminary treatment of any sort. While it seems probable that there may be essential differ- ences between the various forms of finely divided carbon now upon the market, as to their efficiency in the water treatments here con- sidered, no comparisons have been made in this regard. The form here used was chosen merely because of its convenience. Although, as has been pointed out, simple distillation of the Heidelberg tap water from glass into glass failed to correct the toxicity of the water, it was found that this same process of dis- tillation rendered the water nontoxic to the Spirogyra here em- ployed (II, 10) when animal charcoal was present in the retort during the distillation. In this treatment the boiler was a flask of Jena glass having a capacity of one liter; the condenser was of glass, provided with a bend to prevent the entrance of spray from the boiling liquid and thoroughly washed by use; and the receiver was a flask of Jena glass. Nocork or rubber or material other than glass was used on the condenser. The usual charge was about 800 cc. of tap water with from five to ten grams of Merck’s animal charcoal. Boiling was not allowed to become very violent and the ‘process was stopped when the charge had been reduced to about one eighth of its original volume. In water thus prepared Spiro- gyra filaments remained in apparently healthy condition until the appearance of injury from lack of nutrient salts. This method of distillation from animal charcoal showed some variation in the quality of the water produced (II, 10), but the differences exhibited were not as great as those obtained with different portions of water from the stock culture (III, 36). The water distilled from charcoal was frequently tested throughout Hoyt: CULTURES OF SPIROGYRA 339 the experiments,—by determining its effect, when used alone, upon Spirogyra filaments. Out of more than ten runs the distillate from but a single one was found to be decidedly toxic to the alga. The cause of this single failure was not investigated. The new method was uniformly employed for the preparation of nontoxic water throughout the course of these studies. Several experiments were carried out to throw light upon the question of the manner of operation by which distillation from animal charcoal may produce such a marked effect upon the physio- logical properties of the Heidelberg water. Not only the distillate but also the undistilled and concentrated residue (after removal of the charcoal by filtration through paper) showed marked improvement as compared with the untreated tap water (II, 11, 12). Furthermore, a high degree of toxicity was produced in the practically nontoxic water, either of the filtered residue or of distillate, by the addition of a portion of the used charcoal that had been retained in the filter (II, 11, 12a, 12c). From the above observations it appears clear that the animal charcoal in the boiler of the still must have acted to remove toxic substances from the water, although, as has been stated, such action could not be detected at ordinary temperatures. It also appears that charcoal which has been the agent in this removal of injurious material becomes itself able to reproduce toxic properties in nontoxic water to which it may be added. It may be therefore considered as at least highly probable that the action of the finely divided carbon of these distillation experiments was that of an absorber rather than that of an oxidizing or other catalytic agent. Of course it is open to question, whether the toxicity reproduced in harmless water by used animal charcoal may be caused by the same substances as was the original toxicity of the tap water before treatment. The state of our knowledge of these matters precludes their further analysis at this time. One experiment was made to show the relative resistance to the toxic influence in tap water of two portions of Spirogyra which were originally from the same source but which had been subjected for a time to different conditions. One sample was taken from the stock jar and another from a vigorous culture which was origi- nally from the stock culture and then had grown for the last 68 340 Hoyt: CULTURES OF SPIROGYRA days in 0.1 per cent Crone’s solution. The two samples contained approximately equal numbers of filaments. They were well rinsed in nontoxic water and then placed together in untreated tap water in a covered glass dish. Within a few hours the filaments from the stock jar showed decided injury, and, at the end of six days, when the experiment was discontinued, about one third of them were dead (II, 13). The filaments transferred from Crone’s solu- tion showed no signs of injury during the period (II, 14). This experiment emphasizes the facts, well recognized, but not always considered by workers with cultures, that the internal conditions of organisms are fully as important in determining their behavior as are the conditions of the surroundings, and that the past history of organisms (the sort of surroundings under which they have lived) greatly influence their internal nature. Comparable results cannot be expected, in such work as this, unless the living material used has been kept under uniform conditions for a considerable time previous to experimentation. Turning to the results obtained with ordinary distilled water (TABLE Itt), this was found to be extremely toxic, killing all the filaments of Spirogyra, except in a single instance,* in from one to nine days (III, 1). Redistilling this water in glass produced a decided improvement in some cases, but none in others (III, 2-6). Similarly, the samples of water obtained as above described, at the beginning, middle and end of the distillation run, varied greatly and inconsistently among themselves. The undistilled portion left in the boiler after redistillation had been stopped was always just as toxic as untreated water (III, 2a, 3c, 4e, 5e). Making from the distillate or from the undistilled residue (III, 4, 5) a 0.5 per cent concentration of Crone’s solution produced no improvement, so that this addition of nutrient salts failed to counteract the toxicity of these two waters. The addition of animal charcoal to the boiler during redistillation in glass usually produced no improvement over the water redistilled without the solid (III, 7, 8)- Furthermore, the undistilled residue from such redistillation (after — removal of the carbon) showed no marked improvement. Boiling with or without carbon (III, 2-10) is thus seen to have produced * In this case an unusually small amount of distilled water was used, in pro- portion to the number of filaments in the culture. Hoyt: CULTURES OF SPIROGYRA _ 341 no correction of the toxic conditions of ordinary distilled water. Heating in-an autoclave for 15 minutes at 144° C. made ordinary distilled water decidedly less injurious, but did not render it entirely nontoxic (III, 11). Dilution with an equal volume of physiologically pure water (III, 12) or with water from a vigorous culture of Nitella (III, 13) did not make the ordinary distilled water less toxic, but dilution with an equal volume of a colloidal platinum solution (III, 14) rendered this water less injurious.* The addition to ordinary distilled water, at the time of adding the alga, of filter paper (III, 15), cotton (III, 16), kaolin (III, 17), quartz sand (III, 18), or small amounts of chalk (III, 19, 20), produced no improvement; but the addition of abundant chalk (III, 21), lime (III, 25), broken agar sticks (III, 27), or dry sphagnum moss (III, 31) usually rendered the water nontoxic. Spirogyra filaments died within a few hours in ordinary distilled water to which a small amount of agar solution was added (III, 28), they lived nearly two months on the surface of a 1 per cent. solution of agar made with this same water (III, 29), but died within three days on the surface of a 2 per cent agar solution (III, 30). In erdinary distilled water shaken with chalk (III, 22), wood charcoal (III, 23), or garden soil (III, 24) and decanted after the subsidence of the solid, little or no improvement was observed at first, but decided improvement was shown after a few days. Addition to the water of sediment from a vigorous culture of Nitella (III, 32) produced better growth than that occurring in untreated water, while sediment from the stock culture of Spiro- gyra (III, 33) produced no improvement. It was found that the alga filaments themselves may exert a profound influence upon the toxicity of the solution in which they are immersed. A transfer of filaments was made, in the usual manner, from the stock culture to a dish of ordinary distilled water (III, 34). These filaments were dead within eighteen hours. They were then removed from the culture and replaced by a new transfer of filaments from the stock culture. These also died within eighteen hours, and were replaced as before. The third * This colloidal platinum solution was kindly prepared by Dr. W. Fraenkel in the laboratory of Prof. G. Bredig. It contained 0.0096 g. of platinum per 100 c.c, } 342 Hoyt: CULTURES OF SPIROGYRA transfer was killed in about two days, and the material which then replaced it lived fifteen days, but was in apparently poor condition at the end of that time and was replaced. The material of this fifth transfer preserved an apparently healthy condition for forty- four days, at which time the experiment was discontinued. The toxic water had been réndered nontoxic simply by treatment with alga filaments. A clearer example of the effects of plants in re- moving or counteracting the toxic substances of a solution would be difficult to imagine. Similar results were obtained by Dandeno (7) upon growing corn seedlings in toxic solutions of HCl and H.SO,. Toxic water may be greatly improved for Spirogyra by previous treatment with another alga. A mass of Ulothrix (with a few Spirogyra filaments) was allowed to remain twenty-eight days in ordinary distilled water (III, 35), while a control of a similar dish with another sample of the same water, but without alga, stood beside the first. At the end of this period the Ulothrix was re- moved and Spirogyra from the stock culture was transferred to each of the two dishes. In the water which had been previously treated with Ulothrix the Spirogyra lived, albeit in rather poor condition, for twenty-two days, while in the untreated water of the control all filaments were killed within eighteen hours. In connection with the above observations on the effectiveness of alga filaments in correcting the toxicity of ordinary distilled water, it should be noted that, in all these experiments, the amount of Spirogyra added, in proportion to the amount of medium employed, exerted a considerable influence upon the results. A large amount of the alga was able to live in a small amount of a decidedly toxic medium. In all such cases the filaments on the outside of the mass were killed while those within remained in good condition. Similar results have been obtained by Nageli (20), Deherain and Demoussy (8), Bullot (6), Dandeno (7), and Bokorny (3). In the same connection, water from the stock culture of Spirogyra, tested by the method used in these experi- ments, allowed a good growth (III, 36), but seemed less favorable than the water from a vigorous culture of Nitella (III, 37). Although some of the experiments which have been considered were not repeated as many times as might be desirable, the different Hoyt: CULTURES OF SPIROGYRA 343 experiments appear to corroborate each other, and seem to throw some light on the general nature of the toxic substances present in these waters. It will be noticed that'the effect'of distillation and of high temperature was often different in the case of tap water and in that of ordinary distilled water. Some of these differences are summarized in TABLE Iv, where a plus sign indicates improve- ment, a minus sign indicates no improvement. When both signs occur, different tests were in disagreement. The facts of TABLE Iv indicate that the toxic materials present in the tap water and in the ordinary distilled water here studied were, in part at least, different substances; whatever may have been the nature of these substances those in the ordinary distilled water were mainly very resistant to heat. As to the general characteristics of the toxic substances in the ordinary distilled water, there appears no reason for doubting that they were probably, for the most part, metallic in their nature, derived in some way from the distillation apparatus. This is the conclusion reached by many authors who have studied the toxicity of ordinary distilled waters since the time of Nageli, and such supposition introduces no difficulty in the interpretation of the experiments that are reported here; it might be expected that metallic poisons, whether dissolved as salts or ions or existing merely in suspension, would not be very sensitive to correction by heat. The effect of redistillation might be to transfer a portion of the poisons from retort to receiver, thus leaving the distillate more or less toxic, and at the same time concentrating another portion (nonvolatile with steam and not sensitive to the tempera- tures employed) in the retort. It has been shown by Livingston et al. (11), Livingston (12), Wheeler and Breazeale (37), Schreiner and Reed (27, 28, 29), Shorey (30, 31), and Lathrop (26), that certain organic substances Present in various soil extracts are toxic to wheat seedlings. Since, as was pointed out by Livingston (12), tap water may be considered as a natural soil extract, the possibility is suggested that it may be toxic on account of poisons derived from the soil. At the same time, it is of course probable that the toxicity of the Heidelberg tap water here tested may have been due, to a greater or less extent, to substances having their origin in the supply pipes. 344 Hoyt: CULTURES OF SPIROGYRA It may be supposed that somewhat complex poisons emanating from the soil would be largely volatile (either as such or as their decomposition products, e. g., ammonia), and would thus be removed from the retort in the process of distillation. If such volatile material were still toxic and were retained in the receiver, the distillate would of course exhibit toxic properties. But the logical possibilities are here very numerous and a more detailed a priori consideration of this problem would be out of place without much more thorough experimentation than is now available. The toxicity of the waters here dealt with was probably due, in every case, to more than one substance; different portions of tap water varied in their injurious effects, different portions of ordinary distilled water were differently affected by redistillation in glass, and other lines of evidence might be deduced from the present studies to indicate that both waters contained more than one kind of injurious material. It appears highly probable that the beneficial effect on the ordinary distilled water of chalk, lime, agar, sphagnum moss, soil, and finely divided carbon, was due to their adsorptive action, a conclusion similar to that reached in the work of Nageli (20), Breazeale (5), Livingston et al. (11), and Livingston (12). In some cases all the poisonous substances seem to have been removed from solution, for the treated water was entirely nontoxic. The non- toxic water obtained by the distillation of tap water from animal charcoal contained more dissolved electrolytic material than did the toxic water prepared by simple distillation, as was shown by conductivity measurements kindly made upon these waters by Dr. W. Fraenkel, but the total salt content of the charcoal-treated water was calculated to be less than 0.01 per cent, so that the removal of toxicity seems not to be related to an increase in salt content. Furthermore, similar decrease in the toxicity of the water was brought about by substances which were chemically very unlike the charcoal used in these distillations. The small amount of chalk that failed to produce improvement when added to ordinary distilled water was much in excess of the amount dissolved, so that the marked beneficial effect of a larger amount of chalk cannot be considered as due to the addition of calcium to the water. The improvement produced by colloidal platinum and Hoyt: CULTURES OF SPIROGYRA 345 by agar jelly appears also to have been due to adsorptive action, though a discussion of this matter will not be given place here. It is interesting to remark that addition of sand, cotton, and filter paper failed to produce any correction of the ordinary distilled water of these experiments, although these substances have been found by Nageli (20), Dandeno (7), True and Oglevee (36) and Jensen (9), to be active in improving other toxic solutions. The difference thus noted may be of course due either to a difference in the toxic substances dealt with or to differences in the solids used. Breazeale (5) obtained similar negative results upon adding quartz flour, filter paper, paraffin, or sand to solutions of H:SO, which were toxic to corn seedlings. He suggested that the failure of these substances to reduce the toxicity of the solutions was due to the high concentration of HzSO, necessary to produce toxic action, and the consequent relatively slight adsorption of the acid by the solids. This explanation can not, however, apply to the results described above, since in these experiments the amounts of the toxic substances were extremely small. In considering the behavior of Spirogyra in different nutrient media (see p. 335), it has been observed that the optimum con- centration of the solution appeared to be a function of the kind of water used. With solutions that contained salts in the propor- tions adopted by Molisch and Crone and were prepared with non- toxic water, the optimum concentration for the growth of Spirogrya was found to be from 0.05 to 0.1 per cent, while similar solutions prepared with ordinary distilled water exhibited an optimum con- centration ten times as great. It thus appears that the toxicity of the ordinary distilled water was more or less overcome by the larger amount of nutrient salts present and that, conversely, the retarding effect of relatively high salt concentration was counter- balanced by the presence of the toxic material of the water. The influence of dissolved salts in overcoming the toxicity of solutions has been noted by other authors,* but the whole question here raised is not at present in a state to warrant an attempt at detailed discussion. It is obviously related to the general problem of physiological antagonism. Some points on salt antagonism will be brought forward in the succeeding section. es ad * Breazeale (4), Livingston ef al. (11), Livingston (12), Sumner (35), Schreiner and Reed (28), and Schreiner and Skinner (32, 33)- 346 Hoyt: CULTURES OF SPIROGYRA The beneficial effect of the action of alga filaments, in rendering originally toxic water less toxic to other filaments of the same or another form (see p. 341-2), has been noted already. While the evidence at hand suggests the probability that this effect may be due to an absorptive (or adsorptive) action on the part of the filaments,—Nageli (20), Deherain and Demoussy (8),—and that it may be therefore directly comparable with similar effects pro- duced by other solids with very large surface exposure, it must be nevertheless remembered that such an effect may be also partially or wholly due to chemical alterations of the originally toxic substances, as if brought about by enzymatic or other material emanating from the plant. No evidence is available for a decision in this matter. The effects produced upon Spirogyra in toxic water, by sediment from the stock culture of this plant (III, 33) or from a Nitella culture (III, 32), together with the compara- tive growth of the alga in water from the two cultures just men- tioned (III, 36, 37), may suggest a possible excretion from Spiro- gyra of material more harmful to itself than to Nitella. It appears that the question of toxic and antitoxic excretions may enter into problems of algal growth just as it has come to play so important a part in the physiology of the bacteria and fungi and, recently, in that of the higher plants. III. Sats It has been frequently shown for both animals and plants that the toxicity of many solutions of single salts may be counteracted - by the presence of other salts in the same solution, without regard to the physiological effect of the latter salts when employed singly. Loew (15, 16, 17, 18) would restrict this conception of the antagonistic action of two ions to the single case of calcium and magnesium. Benecke (1) showed that the toxicity of other salts than those of magnesium is counteracted by calcium, but ascribed the antagonistic action to calcium only. Osterhout (21, 22, 23, 24, 25) has, however, observed antagonism between fourteen different pairs of ions, including potassium, sodium, ammonium, calcium, magnesium, strontium, and barium. Loeb (13) reported that the toxic effect of a pure NaCl solution, on the eggs of Fundu- lus, was partly counteracted by each of a number of salts, including Hoyt: CULTURES OF SPIROGYRA 347 the bases: calcium, barium, magnesium, strontium, cobalt, aluminium, lead, zinc, and chromium. The experimentation about to be reported, upon the relation to Spirogyra of the salt content of the medium, is merely qualitative, the aim being to gain some knowledge of a number of possible conditions in algal-cultures rather than to investigate any of the chemical questions here encountered. No attention has been | given to the chemical characteristics of the salts employed, the solutions having been prepared on the percentage basis, and the terminology of percentage will be retained in the discussions. The experiments are partly repetitions of those of Benecke (r). Aside from the effect of dissolved salts to counteract the toxicity of salt solutions, these experiments also include a number of tests bearing upon the influence of undissolved materials upon this same sort of toxicity. Throughout these experiments no water was used which had not been shown to be nontoxic; it had all been distilled from animal charcoal in glass, condensed in glass, and collected in glass. Precautions were always taken to avoid possible contamination of the dishes used, especially in the case of the cultures without calcium. All salts used, except the CaCOs, were those of Merck. The experiments are listed, with their results, in TABLE V. For the study of the influence of calcium in the medium a cal- cium-free solution was prepared from equal portions by weight of KNO;, K,HPO,, KCI, and MgSO;. . This solution was tested in two concentrations, 0.4 and 0.1 per cent (of total salt content by weight), and both proved highly injurious to the Spirogyra here employed (V, 1). The addition of calcium-free animal charcoal produced no improvement in either case (V, 2), but common wood charcoal, pulverized, which was shown by test to contain some calcium, improved the weaker solution (V, 3). In the weaker solution a small amount of CaCl produced about as marked improvement as did wood charcoal (V, 4), a trace of CaCO; allowed still better growth (V, 5), while an abundance of the last- named, only slightly soluble salt kept the alga in excellent condi- tion during the entire period of the experiment—35 days (V, 5). These same experiments, performed with solutions prepared from ordinary distilled water, gave the same results, except that the 348 Hoyt: CULTURES OF SPIROGYRA solutions thus made were all more toxic than those from nontoxic water. A solution similar to the one without calcium, but with mag- nesium also absent, was prepared with equal parts by weight of KNO;, KzgHPO,, and KCl. This was more toxic than the calcium- free solution with magnesium present; both 0.4 and 0.1 per cent concentrations were fatal to the plant within two days (V, 6). Animal charcoal produced no improvement in the stronger solu- tion (V, 7). The addition of calcium carbonate in considerable excess of its solubility enabled the plant to grow for several days (V, 8). Asolution containing 0.1 per cent of the three potassium salts and 0.1 per cent of CaCl, was fatal within three days (V, 9), while the addition of 0.1 per cent of MgSO, to the 0.3 per cent solution of potassium salts prolonged the life of the organism for six days, but did not completely counteract the toxicity (V, 10). The last is a repetition of the first experiment of this section, where the organism lived 28 days (V, 1). A simple solution of magnesium sulphate was fatal to the plant in concentrations ranging from 0.4 to 0.01 per cent (V, II). Animal charcoal produced improvement only in the weakest of these concentrations (V, 12). Wood charcoal (with some calcium present) slightly improved both the 0.1 and the 0.4 per cent. solutions (V, 13). While the simple solution of MgSO, killed the organism in two or three days, a 0.4 or 0.1 per cent solution of MgSO, plus 0.1 per cent of CaCl, (V, 14), as well as 0.1 per cent solution of MgSO, plus 0.1 per cent of KCI (V, 15), allowed the alga to live fourteen days. In these experiments the addition of potassium was just as efficient in counteracting the toxicity of magnesium as was the addition of calcium. To determine whether the effect of CaCO; in counteracting the toxicity of MgSO, is to be considered as due solely to the addition of calcium to the solution or partially to other causes, the following experiments were carried out. A saturated solution of CaCO; was prepared and to this was added 0.4 per cent of MgSQ,. At the same time, to a 0.4 per cent solution of MgSO, was added an excess of finely divided CaCO . In the former solution, the alga was markedly injured within nineteen hours, and was totally dead at the end of fourteen days (V, 16). In the solution contain- Hoyt: CULTURES OF SPIROGYRA 349 ing solid CaCO 3, however, the alga remained in good condition for seven days and had many filaments apparently uninjured at the end of fifteen days, when the experiment was discontinued (V, 17). These and other lines of evidence obtained during the progress of this work seem to indicate that the beneficial effect of the powdered CaCO; was due in part to other causes than the mere addition of calcium to the solution, probably to its adsorptive action. This suggestion is further supported by the results of Livingston et al. (rr) Snyder and Cook (34), Breazeale (5), and Schreiner and Reed (28). The colloidal solution of platinum already referred to rendered a 0.01 per cent solution of MgSO, decidedly less toxic (V, 18), but the addition of 0.008 per cent PtCl, to a 0.01 per cent MgSO, solution produced no observable improvement. Although, as was noted above, the addition of 0.1 per cent of KCl rendered a 0.1 per cent solution of MgSO, less toxic, a 0.1 per cent solution of KCI alone was decidedly more toxic than this mixture (V, 20), and a 0.4 per cent solution of KCI was fatal within two days (V, 19). This result is in agreement with the work of Osterhout (22). The colloidal solution of platinum produced improvement in a 0.1 per cent solution of KCI (V, 21). Addition of 0.1 per cent of CaCl: to a 0.1 per cent solution of KCI rendered the latter slightly less toxic (V, 22), although a 0.2 per cent solu- tion of CaCl was fatal within a few hours or days (V, 23). Two experiments were made bearing upon the relative endur- ance, in solutions of the above salts, of samples of Spirogyra originally from the same source, but subjected for a time to dif- ferent surroundings. One portion of an alga culture which had been growing vigorously for sixty-six days in 0.1 per cent Crone’s solution was rinsed in nontoxic water and placed in a solution containing 0.1 per cent of KCI and 0.1 per cent of CaCh (V, 24) and another portion of the same culture was likewise rinsed and placed inao.2 per cent solution of CaCk (V, 25). On the following day were added to both of these cultures samples of Spirogyra from the stock jar. In both cases the solution was injurious to both samples of the alga, but decidedly less so to the filaments from Crone’s solution than to those from the stock culture (V, 22, 23). This result again emphasizes the well-known fact, already men- 3850 Hoyt: CULTURES OF SPIROGYRA tioned, that the internal conditions or the physiological state of organisms used in experimentation of this sort must always be taken into account. In this portion of the work, no evidence was obtained for a specific action of the calcium salts in antagonizing other salts. The action of CaCl, in antagonizing MgSO; and KCl was no more pronounced than was that of KCI in counteracting the toxicity of MgSO, and of CaCh, and that of MgSO, in counteracting the toxicity of KCl and of CaCl. These results seem strongly to support the conception of Loeb (14) and Osterhout (22, 25) of the importance of physiologically balanced solutions. MgSO, CaCl, and KCl, when used separately, and a mixture of three potassium salts were all extremely toxic in the present investiga- tion, but when the salts of any two of these bases were brought together (as in combinations of MgSO, and CaCk, MgSO, and KCL, KCl and CaCl) the resulting solution was much less toxic. A mixture of the three potassium salts here employed was as toxic as KCl alone and only when salts of all three metals were present in favorable proportions was good growth obtained. Results similar to these were reported by Bokorny (2). As it has been shown by several workers that Spirogyra requires calcium, these facts cannot be interpreted by supposing that this alga differs from higher plants in being able to survive in the absence of this element. The improvement produced in the weaker toxic salt solutions by animal charcoal may be considered as probably due to the adsorptive action of the finely divided carbon. The beneficial effect of the addition of colloidal platinum to toxic solutions may have been likewise due to its adsorption of the toxic salts. The author is gratefully indebted to Prof. B. E. Livingston for helpful suggestions in the preparation of this paper. SUMMARY 1. Cultures of Spirogyra longata (Vauch.) Kg. were kept under laboratory conditions in a vigorous state during the entire period of the experiments,—more than 60 days. Crone’s solution was the best of the nutrient solutions tested, and that of Molisch was almost as satisfactory, but the solution of Sachs seemed slightly Hoyt: CULTURES OF SPIROGYRA 351 less favorable, while that of Knop was distinctly unfavorable, probably because of its marked acidity. 2. Both the tap water and the ordinary distilled water of the Heidelberg Institut were markedly toxic to this Spirogyra, even after distillation (or redistillation) from glass to glass. 3. The toxicity of the tap water was partially removed by con- centrating the water to a fraction of its original volume, and was entirely removed by heating to 144° C. or by distillation in glass from animal charcoal; but the water was not greatly improved by simple distillation or by heating to 100°C. The toxicity of ordinary distilled water was partly or wholly corrected by the presence in the culture of chalk, lime, solid agar, dry sphagnum moss, colloidal platinum, or of other adsorbents; it was partially corrected by redistillation or by heating to 144° C. The presence in the cultures of filter paper, cotton, sand, kaolin, or CaCl, was without effect, and redistillation in glass from animal charcoal did not produce nontoxic water. 4. The results obtained seem to indicate that the toxic ma- terials present in the tap water were mostly volatile (perhaps Organic substances derived from the soil), with a relatively small amount of nonvolatile material probably emanating from the supply pipes. The toxic substances in the ordinary distilled water were mostly nonvolatile (probably derived from the sup- Ply pipes and from the still), with a small amount of volatile matter carried from retort to receiver with the distillate. 5. When nontoxic water was used, the best growth was ob- tained in nutrient solutions containing from 0.05 to 0.1 per cent of total salts; but when ordinary distilled water was used, the optimum concentration of similar nutrient solutions was ten times as great. 6. With KNOs, KsHPO,, KCl, MgS0,, and CaCh (singly or in various combinations), a mixture of the three potassium salts was as toxic as KCI alone; KCl, MgSQu, and CaCh were all extremely toxic when used singly; but mixtures of any two of these three were less toxic than the solution of a singleone. Good growth was obtained only when salts of all three metals were present in favor- able proportions. The toxicity. of MgSO, appeared to be as completely counteracted by KCl as by CaCk. 352 Hoyt: CULTURES OF SPIROGYRA 7. Weak solutions of certain toxic salts were improved by the addition of calcium-free animal charcoal or colloidal platinum. It seems indicated that the effect of powdered CaCOs, in counter- acting toxicity due to nutrient salts, was due in part to the adsorp- tive action of the solid. THE JoHNS HOPKINS UNIVERSITY. Hoyt: CULTURES OF SPIROGYRA 363 TABLE I CULTURES OF SPIROGYRA IN NUTRIENT MEDIA PREPARED WITH DISTILLED WATER Nature of culture medium No. of | Author and concen-| Kind of dis- | Agar added! Duration of cul- Condition of culture | tration, per cent of | tilled water | to medium, tures, days plants * ? total salts in used in me- | fer cent by medium dium weight I Sachs, 0.05 — 3 2 0.05 nontoxic —_ 4 E 3 0.10 ordinary 10, 20 Pd, Pd 4 0.10 oO. 1.0 34 P 5 0.10 nontoxic 40, 18, 44 E, Eg, Gp 6 0.50 ordinary — 3,14 D,D 7 0.50 oO. 1.0 60 E 8 0.50 nontoxic — 50 G 9 1.00 ordi — 2 D Io I.00 nontoxic — 44 P II Knop, 0.05 ordinary oe 3:3 D,D 12 0.05 nontoxic — 48 Gp 13 0.10 di — 3 D 14 0.10 do. 1.0 49 Pd 15 0.10 pence a 18, 31, 44,11 | G,t Pd, Pd, Pd 16 0.50 i a 18, 49, I G,t is D 17 0.50 salen “= 23;.2 Pd, D 18 1.00 ordinary — Pd 19 1.00 | nontoxic —< I D 20 | Molisch, 0.05 ordinary — 3 D 2r 0.05 nontoxic — 48 E 22 0.10 | ~ ordinary ee 7 Pd 23 0.10 oO. 1.0 49 - 24 0.10 nontoxic 62, 62, 40,40 | E,G,G,G 25 0.50 ordinary 555 48, 22, 14 G, Pk. Pd 26 0.50 do. 1.0 66 E 27 0.50 nontoxic sae 50 Eg 28 T.00 ordinary —_— 46 P 29 T.00 nontoxic eae: 51 P 30 Crone, 0.05 ordinary = 3 D 31 0.05 nontoxic — 48 E 32 0.10 ordinary 3 D 33 0.10 ; 1.0 8 Pd 34 0.10 | nontoxic 23, 40, 40,51 | E, E, E.G 35 0.50 ordinary aa 48, 13, 14 Eg,t Pd, Pd 36 0.50 do. 1.0 E 37 0.50 nontoxic — 50 2 38 I.00 ordi ca 82 G 39 1.00 nontoxic 51 Gp * The ene nee ob the cont is indicated as follows: E, excellent, G, 7, good; P, poor; D, dea cimacoengr ier by each of ae letters—as, Eg indicates a condition between excellen , ete, the experiment wae i . oO several notations giv en rele ete single experiment. The numbers in eceding column (duration in days) refer to the experiments in the same order as do the letters; thus, I nico condition and the first number denoting duration both refer to the same experimen etc. +The amount of medium used was unusually small, in comparison with the number of filaments in the culture. 354 Hoyt: CULTURES OF SPIROGYRA TABLE II CULTURES OF SPIROGYRA IN TAP WATER, WITH TREATMENTS AND ADDITIONS , Duration No. of ns of Condition culture Treatments of, and\additions to, tap water cultures, of plants* days 2 Ue $4,477 IP, DED 2 paige gibt on to glass 13,44, 2: (P-Pd,D , first portio at, P,D 3a rote 2, middle portion 3, 12 Gp, Gp 30 |As 2, final po 12,3 Pop oiled to A ae ori E812 Eg, P 5 vaporated Hi orig vol G 5a ntreated (control to 5 I Ps Evaporated to 1/6 orig. vol I Gp 6a |Untreated (control I Pd if Evaporated to 1/3 orig. vol I D 7a |Untreated ac to 7) r D 8 {15 mins. at 1 37 E 8a Ppa: ‘onteol to 8) 37 Pd 9 |45m 46 Pd 10 Distillate ees animal charcoal, glass to glass 1 to 44 |E, E, E, Eg, Eg, Eg, Eg, G, P Ir |Boil ith animal charcoal to 1/8 orig. vol., filtered| 26 Ila naar from 26 11b |Do., charcoal rinks Ir added to culture. 26 Gp 12 |As 11, filtered 16 G 12a |As ft, not filtered 16 Gp 12b |As 11a 16 G 12zc |As11b 16 Gp 13 |Untreated plants from stock jar} 6 G 14 |As 13, but plants from Crone’s sol.f 6 E * The condition of th 1 as foll E, excellent; G, good; P, poor; D, dead. Acombinati of letvecn dent condition between that denoted by each of these letters, as: Eg indicates a condition between excellent and good, etc. Where the experiment was repeated, each of the several notations given refers to a single experiment. The numbers in the preceding column (duration in days) refer to the experiments in the same order as do the letters; thus, the first letter denoting con- dition and the first number denoting duration both refer to the same experiment, etc. Tt Nos. 13 and 14 were simultaneous, in the same dish Hoyt: CULTURES OF SPIROGYRA TABLE III 355 CULTURES: OF SPIROGYRA IN ORDINARY DISTILLED WATER, WITH TREATMENTS AND No. of Duration micas Treatments Additions canis. ger sit. days I |Untreated — 54,9, 6,1 (Gp, D, D, D 2 Distillate, gina to glass — ee G 2a concentrated, residue from — I D 2 Distillate, glass to ~_ to 1/8 orig. vo st faa _— I D 3a |As3, middle portion — 4 Eg 36 3, final portion — 4 G 3¢ |Boiled, ee to 1/8 orig. vol., residue fro: — I D 4 poner glass Pig _ to 1/9 vol., first port —_ 9 G 4a re . middie portion = os P 4b |As 4, final portio: — 9 4 4¢ |As 4, all portions mixed _ 9 P 4d |Distillate, glass to glass Salts to form 0.5 pe. pate ssol.} 9 D 4e Boiled, concentrated, residue from a 9 D 4f |As 4e Salts to form 0.5 p.c. Crone’ssol.| I D 5 gligand — ws ore to 1/16 i 12 Eg 54 Ag: : Sade pe Fos a T2 Pd 56 5, —_ ion — 12 D 5¢ |As 5, all portions mixed — 12 P 5d Distillate glass to glass Salts to form 0.5 p.c. Crone’ssol.| 12 Gp 5e Boiled, concentrated. residue from 55 — 12 Fg 5f |As 5e Salts to form 0.5 p.c.Crone’ssol.| 1 6 |Distillate, glass to glass, first por- ti — 4 Pd 7 ee. es animal charcoal, lass t — 46 E 8 As 7, to 8 ote Mi first portion — 2,12 D,D 8a As 8, mi — — 3, 12 G, Ye d 8b |As 8 detent pea 12,1 P,D 8c Boiled wth aiiio seit charcoal, fil- tered, concentrated residue from 8b — 3) 12 ° a eg 9 (Boiled 3 me D Io sue nnd Sith ss ooh charcoal, not : D Iz. its pons at 144° C. a 39 G 12 adie: xic vere to do wna vl a,1,% D, D, D 13 do. Nite alba a double prams I D 356 Hoyt: CULTURES OF SPIROGYRA TABLE III, continued. Duration a -abediok Treatments Additions cohen, - aaa days | ] 14 do. Colloidal Pt sol. | : to double vol.) 26 Gp 15 do. Filter er 10, ¢ Pd, D 16 do. Cotto ;2 D, D 17 do. Kaolin ak D,D 18 do. Quartz sand i, z DD, 19 do. Pigoes trace a D 20 do. do., exce: 5 D 2I do. do., ahanaaice 23, 56 E, Eg 22 |Shaken with chalk, decanted after| 22a \As 22, decanted after 10 days — 9, 10 G, Pd bY seein with wood charcoal, de- nted — 53 r 23a Abe 23, decanted after 4 days — 30 G 24 |Shaken with aga decanted after sever i day: 44 Gp 25 #=|Untrea : Lime, abundance| 2 E 26 do CaCle, O 0.4 p.¢ D 27 do Agar sticks 20 E 28 do. Agar, not hard enin: I D 29 do Agar, I ~.c., alga ons 54 G 30 do Agar, 2 p.¢., as 29 ra) 31 do hagnum mo 23, 23. E, E, Pd 32 do Nitella sediment | 48 G 33 do og 5 D 34 do. I D 34a |Water from 34, alga removed Alga bro by resh material pe sek jar} 1 D 34b |Water from 34a, alga removed do. 2 D 34c |Water from 34), alga removed do. 15 9 34d |Water from 34c, alga removed do. 44 E 35 |Treated with Ulothrix,} removed after 28 days do, 22 te 35a (Untreated (control to 35) _ _ I D 36 Mele from stock Shack cul- ps 52, 47,9» 4 E, Eg, G, Gp 37. |Water from Nitella culture — 53,9 E,E * The condition of the plants is indicated as follows: E, excellent; G, good; P, poor; D, dead. A friana oe: of ieee a a oe be in i that denoted by each of these letters—as, Eg nt and good, etc- Where the experiment was eee ted, each of the several notations given refers to a single experiment. The numbers in the preceding column (duration in days) refer to the experiments in the same order as do the letters; thus, the first letter denoting condition and the first number denoting duration both refer to the same experiment, + Ulothrix killed except in center of mass. Hoyt: CULTURES OF SPIROGYRA 357 TABLE IV EFFECT OF VARIOUS OF TAP AND DISTILLED WATER UPON THEIR TOXICITY Tap Ordina Treatment Water Distilled Water Distilled in glass (distillate)........ so cent Distilled in glass (undistilled residue).............-+++ + on Distilled with carbon (distillate). ... mip =a Distilled with carbon (undistilled residue)...........-- + = Pleated t6:r44 "Coe ee ac ee ee o +* * Still toxic, but much improved. 358 Hoyt: CULTURES OF SPIROGYRA CULTURES OF SPIROGYRA IN SALT SOLYTIONS PREPARED FROM NONTOXIC WATER, WITH ND WITHOUT ADDITIONS Salts in solution, per cent es No. of Additians Duration of} Condition of culture | ¢ NO; [KeHPO,| MgSO,| KCI Oe TS ee ET: | Ost 0.1 0.1 0.1 — 13, 28,8 | Gp, Gp, Pd Ia@ | 0.025 | 0.025 | 0.025 | 0.025 — 26, 29 Gp, P 2 o.1 0.1 0.1 0.1 Animal charcoalf 2a | 0.025 | 0.025 | 0.025 | 0.02 0. 26 Gp Si 0.1 0.1 0.1 Wood charcoal I5 Pd 3a | 0.025 | 0.025 | 0.025 | 0.025 2 35 G 4 do., do. do. do. jCaCle 28 G . do. do. do. do. |CaCOs, t 35 Eg 5a ‘ do. do. do. |CaCO; abundance 35 E 6 0.133 | 0.133 — | 0.13 2 D 6a | 0.03 0.033 — | 0.033 — 2 D 7 0.133 | 0.133 | — | 0.133 |Animal charcoalf 3 D 8 5 do, —_— do. |CaCOsz 6 Be 9 0.1 0.1 — |o.1 CaCle 0.1 p.c. 5 D 10 do. do. | 0.1 do. —_ 6 Pd II ae — 10.4 — a 3,2 D,D Ila _ — 0.1 —_ —_— a2 DD. D 11b —_ — | 0.05 —_— — 2 D Tie _ — | 0.01 —_— 6,2 Pd, D 12 a — \o4 — jAnimal charcoalt 2 D I2a —_ — “| O2% —s do. 2 D 12b — — | 0.05 _— do. 2 D I2¢ — — | 0.01 oo do. 9,5 PoP 13 — — |04 — |Wood charcoal{ re do: 13a _ — |o.1 — i Pd 14 — — |04 — |CaCle 0.1 fa 15 P 14a ee — | 0.1 — : baa Pd 15 — do. | 0.1|| 14 P 16 — — |o4 — |CaCOs ings pater 14 Pd 17 _ do. — |CaCOs, in I Gp 18 — — | 0.01 —_— voyensie cl. Pt 90 p.c.| 19 Gp 18a —_ do. — |Pt Cl 8 p.c. 2 D 19 — — |0.4 2 D 20 — _— — |0.1 a 6 Pd 21 — a — do. |Colloid. sol. Pt 12 - : 22 — — do. |CaCl: 0.1 p.c. 7,6 Gp,° Pd 23 — —_ — — jCaCl: 0.2 p.¢ a2 D,AD 24 — — o.r |CaCle o.t p.¢ 4 G° 25 — — —_ — |CaCle 0.2 p.c. 7 G4 * Th d ] E, excellent; G t ; P, poo D, dead. A combination of letters denotes a condition between. oat ae oe indi each of these letters—as, Eg cates a condition between excellent and good, etc. Wh experiment was , each of t veral ions gi refers to a single experiment. The numbers in the preceding (duration in days) refer to the Sagundeprninge in the same order as do the letters, ur 2 the — letter denoting condition and the first number denoting duration both refer to the same experiment, etc. + Culture 23 days old showed n um with acetic acid and ium oxalate TA culture 14 days old denied no Rag re Containing some calcium. || Showed no calcium. ° Spirogyra from culture in Beacon’ s solution, 66 days old, was used in 24 and 25- The first test of the stock material (22) and of material from Crone’s solution (24) bie mage carried out in the same he second of the Spirogyra (23) and of material from Crone’s stock kale (25) were nerranronand carried out in the same . ie) 19. 20, Hoyt: CULTURES OF SPIROGYRA 359 LITERATURE CITED Benecke, W. Ueber die Giftwirkung verschiedener Salze auf Spirogyra, und ihre Entgiftigung durch Calciumsalze. Ber. Deuts. Bot. Ges. 25: 322-337. 1907. Bokorny, Th. Ueber den Einfluss des Calciums und Magnesiums auf die Ausbildung der Zellorgane. Bot. Centralbl. 62: 1-4. 1895. Bokorny, Th. Nochmals iiber die Wirkung stark verdiinnter Lésungen auf lebende Zellen. Arch. Ges. Physiol. 110: 174- 226. 1905. . Breazeale, J. F. Effect of the concentration of the nutrient solu- tion upon wheat cultures. Science II. 22: 146-149. 1905. . Breazeale, J. F. Effect of certain solids upon the growth of seed- lings in water cultures. Bot. Gaz. 41: 54-63. 1906. Bullot, G. On the toxicity of distilled water for the freshwater Gammarus. Univ. Calif. Publ. Physiol. 1: 199-217. 1904. Dandeno, J. B. - The relation of mass action and physical affinity to toxicity. Amer. Jour. Sci. IV. 17: 437-458. 1904. Deherain, P. P. & Demoussy, E. Sur la germination dans l’eau distillée. Compt. Rend. 132: 523-527. 1901. Jensen, G. H. Toxic limits and stimulation effects of some salts and poisons on wheat. Bot. Gaz. 43: 11-44. 1907- sc latnr Ernst. Kultur der Mikroorganismen. Leipzig. 1907. Livingston, B. E., Britton, J. C. & Reid, F. R. Studies on the properties of an unproductive soil. U. S. Dept. Agric., Bur. Soils, Bull. 28. 1905. Livingston, B.E. Further studies on the properties of unproductive soils. U.S. Dept. Agric., Bur. Soils, Bull. 36. 1907. Loeb, J. Studies in general physiology. Chicago. 1905. Loeb, J. The dynamics of living matter. New York. 1906. Loew, O. The physiological réle of mineral nutrients in plants. U. S. Dept. Agric., Bur. Plant Ind., Bull. 45. 1903. Loew, O. Note on balanced solutions. Bot. Gaz. 46: 302, 303. Eoew, O. Bull. Col. Agric. Tokio 7: 7- 1906 (quoted verbatim by Benecke, Ber. Deuts. Bot. Ges. 25: 324- 1907). Loew, O. & May, D. W. The relation of lime and magnesia to plant growth. U.S. Dept. Agric., Bur. Plant Ind., Bull. I. 1901. Lyon, E. P. A biological examination of distilled water. Biol. Bull. 6: 198-202. 1904. Nageli, C. von. Ueber oligodynamische Erecheinungen in lebenden Zellen. Neue Denkschr. Schweiz. Gesell. Naturforsch. (Nouv. 360 Hoyt: CULTURES OF SPIROGYRA Mém. Soc. Helvétique des Sci. Nat.) 33: Abth. I., second con- tribution, pages I-52. 1893 21. Osterhout, W. J. V. Extreme toxicity of sodium chloride and its prevention by other salts. Jour. Biol. Chem. 1: 363-369. 1906. 22. Osterhout, W. J. V. On the importance of physiologically balanced solutions for plants. Bot. Gaz. 42: 127-134. 1906; 44: 259- 272. 1907. 23. Osterhout, W. J. V. The antagonistic action of magnesium and potassium. Bot. Gaz. 45: 117-124. 1908. 24. Osterhout, W. J. V. Die Schutzwirkung des Natriums fiir Pflanzen. Jahrb. Wiss. Bot. 46: 121-136. 1908. 25. Osterhout, W. J. V. The nature of balanced solutions. Bot. Gaz. 47: 148, 149. 1909. 26. Schreiner, O. & Lathrop, E. C. Examination of soils for organic constituents, especially dihydroxystearic acid. U. S. Dept. Agric., Bur. Soils, Bull. 80. 1911. 27. Schreiner, O. & Reed, H. S. Some factors influencing soil fertility. U.S. Dept. Agric., Bur. Soils, Bull. 40. 1907. 28. Schreiner, O. & Reed, H.S. Ceitain organic constituents of soils in relation to soil fertility. U. S. Dept. Agric., Bur. Soils, Bull. 47. 1907. 29. Schreiner, O. & Reed, H.S. The toxic action of ae organic plant constituents. Bot. Gaz. 45: 73-102. 30. Schreiner, O. & Shorey, E. C. The isolation of sera organic substances from soils. U.S. Dept. Agric., Bur. Soils, Bull. 53. 1909. 31. Schreiner, O. & Shorey, E. C. Chemical nature of soil organic matter. U.S. Dept. Agric., Bur. Soils, Bull. 74. 1910. 32. Schreiner, O. & Skinner, J. J. Some effects of a harmful organic soil constituent. U.S. Dept. Agric., Bur. Soils, Bull. 70. 1910. 33- Schreiner, O. & Skinner, J. J. Organic compounds and fertilizer action. U.S. Dept. Agric., Bur. Soils, Bull. 77. 1911. 34. Snyder, A. H. & Cook, C. L. The maintenance of fertility. Ohio Agric. Exp. Sta., Bull. 167. 1905. 35. Sumner, F. B. Further studies of the physical and chemical. rela- tions between fishes and their surrounding medium. . Jour. Physiol. 19: 61-96. 1907. 36. True, R. H. & Oglevee, C. S. The effect of the presence of in- sain ae on the toxic action of poisons. Bot. Gaz. Q: I-21. 37- wise: H. I. . Breazeale, J. F. On the causes of unproduc- tivity in a Rhode Island soil. Ann. Rep. R. I. Agric. Exp- Sta. 1905: 286-323. 1906. The production of a promycelium by the aecidiospores of Caeoma nitens Burrill Otto KUNKEL INTRODUCTION Except in the genus Endophyllum, so far as known, the aecidio- spores of the rusts give rise to sporophytic mycelium. Its sporo- phytic nature is shown by the fact that each cell contains two nuclei that have been produced by successive conjugate divisions of the two nuclei in the aecidiospore. The aecidiospores of some of the species of Endophyllum, however, produce a promycelium. Its cells are uninucleate and belong, therefore, to the gameto- phytic generation. This fact has been well established through the work of Tulasne (10), Sappin-Trouffy (8), Maire (5) and others. Hoffmann has shown that nuclear fusion occurs in the aecidiospores and that this fusion nucleus divides to form the nuclei of the sporidia. This interesting form has received con- siderable attention in recent years, because its life history offers suggestions regarding evolution in the rusts. While engaged in a study of the effects of media on the germina- tion of various spores I was surprised to find that the aecidiospores of Caeoma nitens taken from leaves of Rubus frondosus Bigel. on germination produce a promycelium in much the same way as do the aecidiospores of Endophyllum Sempervivi Lév. My first suspicion that this might be due to the influence of the medium upon which the spores had been placed proved groundless, for further tests showed that they germinate in exactly the same way when placed in distilled water or in tap water. The production of a promycelium by these aecidiospores seems to have escaped the observation of the numerous students of this common and widely distributed rust. Galloway (6) figured them germinating in water and shows that the germ tube becomes more or less septate. He did not, however, observe the production of sporidia 361 362 KUNKEL: PROMYCELIUM OF CAEOMA NITENS and his drawings hardly suggest the promycelial nature of the germ tube. Clinton (1) has also studied the germination of these spores and gives more than twenty figures, showing them in various stages of development, but he failed to find any trace of sporidia. He mentions having observed the spores in water cultures during two or three days and gives a detailed account of their germination. He also observed the germination of the spores on the moist surface of young blackberry leaves. Olive (7) and Kurssanow (4) have investigated Caeoma nitens from the cytological standpoint. They have shown that sexual fusions occur in the base of the caeoma and that the aecidiospores typically contain two nuclei. OBSERVATIONS The aecidiospores used in this study of their germination were obtained from well infected leaves of Rubus frondosus. The leaves were collected in three different parts of Van Cortlandt Park, New York City. All of the spores germinated in the same fashion. The cultures were made in Petri dishes containing distilled water, tap water, and agar media of various kinds. The spores were simply dusted over the surface of the medium and generally gave a high per cent of germination. The cultures were kept at room temperature (about 23° C.). The promycelium ordinarily consists of five cells (FIG. I, @), four of which bear sporidia. Stained preparations show that the stalk cell is without a nucleus and that the other four cells contain one nucleus each (Fic. 1, b), thus demonstrating the promycelial nature of the germ tube. Frequently the promycelium consists of only four cells and in that case each cell is capable of producing a sporidium (FIG. I, c). There are also instances where the promy- celium consists of less than four cells, but in such cases two or more sporidia are generally produced on the same cell. The promycelium sometimes consists of more than five cells (FIG. 1, d). Five or six sporidia are sometimes produced by the same promy- celium, but thisisa very rarecase. In FIG.1 , eis showna basidium with sporidia in different stages of development; it also shows one sporidium that has fallen off from its sterigma and has begun to germinate. * KUNKEL: PROMYCELIUM OF CAEOMA NITENS 363 The sterigma arises as a pointed outgrowth from a promycelial cell. It tapers evenly and is generally slightly curved rather than straight. Its diameter at the base varies between three and five microns, while at the tip its diameter measures from one and one half to two and one half microns. It normally reaches a length of from fifteen to twenty microns, but in some cases it becomes much longer. The sporidium, as is the rule in the rusts, is a typically kidney- shaped spore, slightly narrower at its basal than at its distal end. Maturing in a short time after the aecidiospore begins to germinate, it soon falls off from its sterigma and, on a moist surface, germinates Fig. 1. mycelium; d, aecidiospore producing an abnormal promycelium; ¢, germ aecidiospore with promycelium bearing sporidia in different stages of development. in a few hours. Sometimes it produces a secondary sporidium, but more often it may develop a corkscrew-shaped germ tube that frequently reaches a length eight or ten times that of the sporidium. Occasionally this germ tube shows branching. The diameter of this germ tube is always much less than the diameter of the nor- mal promycelium. * 364 KUNKEL: PROMYCELIUM OF CAEOMA NITENS Although there is considerable variation in the length and shape of the promycelium when grown on various media, under fairly similar conditions it is rather constant both in size and in shape. Generally it reaches a length of three or four times the diameter of the spore, but sometimes it becomes much longer than this. The average diameter of the spore, based on a measurement of twenty-five spores taken at random, is twenty-one microns. The average length of the mature promycelium, based on the same number of measurements, is seventy-eight microns. DIscussION Tranzschel (9) and Clinton (2) both claim to have established by infection experiments that Puccinia Peckiana Howe is the teleutostage of Caeoma nitens. Tranzschel removed three healthy blackberry plants from the park of the St. Petersburg’ Forestry Institute. One of these plants he grew under a bell jar in the laboratory of the Institute; the other two plants were placed in a garden. The three plants were inoculated with spores of Caeoma nitens. Later in the season all of the three plants became infected with Puccinia Peckiana. Tranzschel had no control plants in his experiment and he makes no mention of having observed the plants in the park of the St. Petersburg Forestry Institute at the time his plants showed infection with the Puccinia. Clinton transplanted blackberry plants from the forest to his greenhouse. He inoculated two of these with the aecidiospores of Caeoma nitens taken from infected blackberry plants. In a little less than two months both of these plants showed infection with Puccinia Peckiana, while several other blackberry plants that had been used as checks showed no infection. It is worth noting that three raspberry plants which he inoculated with spores of Caeoma nitens taken from infected raspberry plants, failed to become infected. All plants used had been free from both the Caeoma and the Puccinia during the previous summer. It is hardly to be expected that the change from sporophyte to gametophyte should occur in two different places in the life history of the same rust, but further experiments are needed in order to settle this point. The production of a promycelium by the aecidiospores of KUNKEL: PROMYCELIUM OF CAEOMA NITENS 365 Caeoma nitens suggests that it is a short-cycled rust in no way connected with Puccinia Peckiana and in its life history quite comparable to Endophyllum Sempervivi. The perennial mycelium lives over winter in the tissues of the host. Cell fusions at the base of the chains of aecidiospores produce sporophytic cells from which the aecidiospores arise by conjugate division of the two cells that fuse. This gives binucleated aecidiospores and intercalary cells. The fact that the aecidiospores germinate in a promycelium suggests that here, as in Endophyllum, nuclear fusion takes place _ in the aecidiospore and we may expect that the sporidia produced by the promycelium reinfect the host and produce a mycelium with uninucleated cells, thus completing the life cycle. If, on the other hand, Puccinia Peckiana is a stage in the life history of this fungus, the problem is not so simple and no similar case is known among the rusts. As was pointed out in the introduction, the fact that sexual fusions occur previous to the production of aecidiospores is well established. The above described germination of these spores makes clear another stage in the life history of this fungus, and raises a number of interesting questions regarding its position among the rusts. I shall test during the coming summer the evidence regarding the connection supposed to exist between Caeoma nitens and Puccinia Peckiana, and shall also undertake a cytological study of both of these forms. The wide distribution and economic importance of Caeoma nitens, its abundant production of spores and the ease with which they may be germinated and studied makes it a favorable object for class use. It is, therefore, highly important that all stages in its life history should be thoroughly understood. SUMMARY 1. The aecidiospores of Cacoma nitens on germination regu- larly produce a promycelium. 2. This promycelium normally consists of five cells. The stalk cell contains no nucleus, but the other four cells contain one nucleus each. _ ; 3. Each uninucleated cell bears a sporidium on a sterigma. 4. These sporidia germinate immediately by producing either a secondary sporidium or a germ tube. 366 KUNKEL: PROMYCELIUM OF CAEOMA NITENS 5.. The production of a promycelium by these aecidiospores suggests that Caeoma nitens is a short-cycled rust and casts doubt on the connection supposed to exist between this fungus and Puccinia Peckiana. 6. Caeoma nitens is the only rust of the caeoma type, having aecidiospores that are known to produce a promycelium. The other rusts having aecidiospores that are known to function as teleutospores, belong to the one genus Endophyllum. I am greatly indebted to Dr. R. A. Harper and Dr. W. G. Marquette for advice and criticism while engaged in this work. COLUMBIA UNIVERSITY, NEw York CIty. BIBLIOGRAPHY 1. Clinton, G. P. Orange rust of raspberry and blackberry. III. Agr. Exp. Sta. Bull. 29: 273-300. 1893. 2. Clinton, G. P. Relationship of Caeoma nitens and Puccinia Peckiana. Bot. Gaz. 20: 116. 1895. 3. Hoffmann, A. W. H. Zur Entwicklungsgeschichte von Endo- phyllum Sempervivi. Centralbl. Bak. 2 Abt. 32: 137-158. 1912. 4. Kurssanow, L. Zur Sexualitat der Rostpilze. Zeits. Bot.2: 81-93- TQIO. 5. Maire, R. L’évolution nucléaire chez les Endophyllum. Jour. de Bot. 14: 369-382. 1900. 6. Newcombe, F. C. & Galloway, B. T. Perennial mycelium mt the fungus of the blackberry rust. Jour. Mycol. 6: 106-107. 1890. 7. Olive, E. W. Sexual cell fusion and vegetative nuclear divisions in the rusts. Ann. Bot. 22: 331-360. 1908. 8. Sappin-Trouffy, P. Recherches histologiques sur la famille des Urédinées. Le Botaniste 5: 59-244. 1896. 9. Tranzschel,W. Culturversuche mit Caeoma interstitiale Schlechtd. (= C. nitens Schw.). Hedwigia 32: 257-259. 1893. Tulasne, L. R. Second mémoire sur les Urédinées et les Ustilaginées. Ann. Sci. Nat. IV. 2: 77. 1854. m4 9 A case of bud-variation in Pelargonium A. B. STOUT (WITH PLATE 20) The precise nature of bud-variation is not satisfactorily known. No adequate classification of the various kinds of bud-variation has been made. Recent investigations regarding the nature of plant chimeras indicate that some of the phenomena generally considered as bud-variation are associated with chimeras and are to be explained by the nature of the chimeras. Observations and experiments are now being made at the New York Botanical ~ Garden on various types of bud-variation. In the studies on Pelargonium one case has arisen which seems of special interest in its bearing on the nature of bud-sports from plants that are chimeras. Baur (1909 a and 6; 1911) has recently shown that the variega- tion in the case of the ‘‘albomarginatae” varieties of Pelargonium zonale is due to the presence of white and green cells which are sharply distinct and which occupy a characteristic position in relation to each other. It has been long known that the paler tissue of these variegated plants owes its characteristics to a lack of chlorophyl in its cells. Baur shows that the plastids are present in the white cells but are colorless. Baur further claims that this arrangement of green and white cells in the leaf can be explained by the arrangement of the corre- sponding tissues in the growing point and actually shows that in the plants whose leaves have layers of white cells on the exterior there is in the apex of the stem a cap of white cells over the greener cells beneath. In such plants the relative position of the white and the green cells is maintained throughout the development of the leaves. : By further study Baur (1909 a; 191 1) found that in other cases these two kinds of cells may be variously arranged with reference to each other. In some plants various stems and leaves show a 367 368 STOUT: BUD-VARIATION IN PELARGONIUM sectoral arrangement with a more or less bilateral distribution of the two kinds of cells. In certain types of Pelargonium the green cells are outside as one or two layers covering the white cells. In some individuals streaks of one of the tissues are mingled with the | other. Baur is able to interpret the conditions in these various forms by using the conception developed by Winkler (1907) in his re- markable discoveries regarding the so-called chimera-nature of graft hybrids. Baur introduced the term periclinal chimera for the condition where the peripheral cell-layers are different from the enclosed tissues and sectoral chimera for the cases where there is more or less of a bilateral or radial distribution. The term hyperchimera, first suggested by Strasburger (1909), is used for the cases where there is a more or less intimate mixture of the different kinds of cells. On these various chimeras of Pelargonium, wholly green or wholly white shoots may arise. This is due to the fact that the two kinds of cells which are maintained by the cell divisions in the meristematic regions become segregated in the growing points, the process not being essentially different from that by which peripheral, sectoral, or hyperchimeras arise. The various types of these Pelargonium chimeras are familiar to horticulturists and have been propagated rather widely by cuttings, thus preserving quite uniformly the different forms. The periclinal chimeras having white peripheral cell layers are com- monly cultivated forms on account of the striking effect of the white-margined leaves. One of these varieties is known by the trade name of Madame Salleroit. During the summer of 1912 a plant of this type which was grown in an outdoor bed at the propagating houses of the New York Botanical Garden produced a branch in which the relative position of the two kinds of cells is reversed. When the cutting was made during the early part of the summer it possessed only leaves with the white margin. In October of that year, when first brought to the attention of the writer, the plant, ap- peared as shown in the photograph here reproduced (see PLATE 20). Two branches, one the main and the other a lateral branch, bore leaves with the white cell layers placed externally to the green as Stout: BUD-VARIATION IN PELARGONIUM 369 | was the case in the plant from which the cutting was made. The leaves were quite uniform in shape and in coloration and are quite typical for the variety known as Madame Salleroi. Dr. L. H. Bailey kindly made the varietal determination from leaves taken from these branches and further states in a letter to the writer that this variety has not been thus far placed specifically. As shown in PLATE 20, the enclosed green cells fail to develop uniformly toward the margin of the leaves, thereby leaving an irregular marginal zone of pure white cells. In the central portion of these leaves the enclosed green tissues show through the white. On the third branch, which is a lateral one, the leaves are quite different. They are larger and the surface is green to the extreme margin. These leaves are not, however, of a uniform green, for through the central portion of each there is an irregular palmate- shaped area of lighter green which is due to white cell-layers enclosed between the upper and the lower green layers. In other words the place relationship of the white and the green cells is here reversed from what it is in the main part of the plant. In this branch the plant has literally turned itself inside out. Micro- scopical examination of free-hand sections confirmed the super- ficial observations as to the color relations. In the black and white plate accompanying this article the general pattern in the leaves is well shown by the different shades. Since this plant has been under observation, about twenty leaves have matured on this branch. As shown in PLATE 20 the amount and distribution of the white tissue varies in the different leaves. In some leaves there are small flecks of white scattered through the green. A few of the leaves when about half developed show traces of a dark zonal band which is a feature of various showy- leaved Pelargoniums. When these leaves are mature, however, this zonal band is faint. Baur notes a case of bud-variation identical with this one. He states (1909 a, p. 333), that the plant which he designated as Pel. 9 had white-bordered leaves but produced in 1908 a br anch having wholly green leaves but which were plainly of a yellowish green in the center. The anatomical studies of these leaves (1909 @, p. 345) showed that the white cells were enclosed by the green cells, 3870 Stout: Bup-VARIATION IN PELARGONIUM In the light of Baur’s anatomical studies this sort of bud-varia- tion is readily understood as due to a mechanical readjustment of the two kinds of cells already in the growing points. This par- ticular type involves more extensive rearrangement than the cases where pure green or pure white branches are produced by the development of a bud containing only one kind of cells to the exclusion of the other. For the development of a branch re- versing the position of the two kinds of cells as described above there must be a breaking out of the enclosed green cells in the growing point and a growth of both green and white cells in such a manner that the green cells surround the white cells. It may be that in this case the green cells break forth at two separate points not far distant and that in further growth they meet, enclosing the white cells. On the main portion of the plant here shown, the mature leaves possess over their whole surfaces two peripheral cell-layers that are white. To maintain this relationship the cell divisions which give rise to these layers must occur only in planes which are at right angles to the surface of the leaf, The outer layers do not contribute to the vascular tissues and the inner green tissue does not form epidermis, a fact clearly shown by Baur. In the sporting branch, however, the green cells get to the surface and form the epidermis as well as some of the mesophyl and vascular tissue, while the white cells cease to form epidermis and now contribute only to the inner tissues. The cells preserve the green and white character of their chromatophores but take on different structures or different functions according to position and environment. © In his interesting report of results of anatomical and hereditary studies of variegated varieties of Pelargonium, Baur was not especially concerned with the evidence of interaction between the two kinds of cells, the white and the green, where both exist in the same leaf. His photographs, however, show the same sort of difference which have appeared so strikingly in the case here under consideration. The marked differences between the two kinds of leaves pro- duced on this plant (PLATE 20) make it clear that the outer layers largely determine the size of the leaves and the depth of the lobing- When the green is outside the leaf is larger, more deeply lobed and Strout: BupD-VARIATION IN PELARGONIUM 371 like those which are borne on the branches that are composed purely of green tissue. When the white is outside, the leaf is much smaller and more like those leaves which are composed only of white tissue. Several plants of Pelargonium Madame Salleroi of the same clone as the plant producing this bud-variation have been under observation in the propagating houses. Several of these have produced leaves composed wholly of white cells. These have been of practically the same shape and size as the variegated leaves on the same plant. The pure white cells are, of course, dependent upon the green cells for carbohydrate food. In the case of a chimera relationship with the green cells enclosed, there may be mechanical and chemical stimuli to the overlying white cells that result in a slightly larger leaf. This effect is, however, not marked. All the potentialities of a large and deeply lobed leaf are present in the green cells of a typical leaf of the Madame Salleroi variety. When these green cells get to the exterior those potentialities find expression, but as long as the peripheral layers of white are uni- formly maintained, there is no visible evidence that these poten- tialities exist. Their suppression may be due chiefly to mechanical limitations imposed by the peripheral layers of white cells which decrease the number of cell divisions. In the various plant chimeras there is an association of more or less independent and different kinds of cells. In the chimeras resulting from grafting, the two kinds of cells may be decidedly different, producing, when separate, two distinct types of leaves, but when associated together, forming leaves of still different Patterns. The various chimeras produced by Winkler (1907 and 1909) and the chimera Crataegomespilus Asnieresti (see illustration by Baur 1911, pl. VIIZ) illustrate this phenomenon. In addition to such mechanical and physical interactions, Winkler (1910) has Presented some evidence that there may be a vegetative fusion of cells in graft tissues producing what he would consider as the only true graft-hybrid, and he further holds that hybrid modifications may also result from the migration of such substances as atropin or nicotine between stock and scion. These facts indicate that the general phenomena of plant chimeras have a very direct bearing on theories of morphogenesis 372 Strout: BUD-VARIATION IN PELARGONIUM and that cellular interaction is a potent factor in influencing cell differentiation and in determining the physical characteristics of such organs as leaves. The bud-sports which arise from the various Pelargonium chimeras are, we may say, rather simple cases of variation due to a mechanical rearrangement of the kinds of cells already present. Their appearance is a confirmation of the rule that like produces like in its application to cell lineage rather than evidence of spon- taneous somatic mutation or variation. In this case the real variation occurs when the white cells appear as the progeny of green cells. The frequency of this spontaneous variation in Pelargonium (and in other cases as well) and the real nature and the causes of the process.are problems for future solution. New YorK BOTANICAL GARDEN. LITERATURE CITED Baer, E. (19092). Das Wesen und die Erblichkeitsverhaltnisse der ‘‘varitates albomarginatae hort.” von eee zonale. Zeits. f. ind. Abst. u. Vererbungslehre. 1: 330- (1909b). Propfbastarde, “ae: oe Pe und Hyperchi- maren. Ber. Deuts. Bot. Ges. 27: 603-605. pach Einfiihrung in die experimentelle Vererbungslehre. Strasburger, E. (1909). Meine Sgelaeea zur Frage der Propf-_ bastarde. Ber. Deuts. Bot. Ges. 27: 511-528 Winkler, H. (1907). Uber Propfbastarde und pilanalich Chimaeren. Ber. Deuts. Bot. Ges. 25: 568-576. (1909). Weitere Mitteilungen iiber Propfbastarde Zeits. Bot. I: 315-345. (1910). Ueber das Wesen der Bluse Ber. Deuts. Bot. Ges. 28: 116-118. INDEX TO AMERICAN BOTANICAL LITERATURE (191-1913) The aim of this Index {s to include all current botanical literature written by Americans, published in America, or based upon American material ; the word Amer- ica being used in the broadest sen views, and papers that eee exclusively to. forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and no attempt is made to index the literature of on: An occasional exception is made in favor of some paper appearing in an American periodical which is devoted wholly to botany. Reprints are not mentioned rinfeas they differ from the original in some important particular. If users of the Index will call the attention of the editor to errors or omissions, their kindness will be appreciated. This Index is reprinted monthly on cards, and furnished in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitted ; each subscriber must take all cards published during the term of his subscription, Corre- spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club, Ames, A. A new wood-destroying fungus. Bot. Gaz. 55: 397-399- f. 1-6. 15 My aN Poria atrosporia sp. n e Amundsen, E. O. Black rot of the navel orage. (Alternaria Citri Pierce and Ellis.) Monthly Bull. State Comm. Hort. Calif. 2: 527- 537- f. 324-330. My 1913: Ashby, S. F. Banana diseases in Jamaica. Jamaica Dept. Agr. Bull. II. 2: 95-128. pl. 21-28. Ja 1913. Ashby, S. F. Diseases of cocoes and other crops. Jamaica Dept. 913- Atkinson, G. F. Is the biennial habit of Oenothera races constant in their native localities. Science II. 37: 716, 717-9 My 1913. Bartlett, H. H. Systematic studies on Oenothera,—lll. New species from Ithaca, New York. Rhodora 15: 81-85. 19 My 1913. Includes Oenothera nutans Atkinson & Bartlett and O. pycnocarpa Atkinson & Bartlett, spp. nov. Blake, S.F. Forms of Ophiogl. Rhodora 15: 86-88. f. 1. “to My 168 <* Blake, S. F. Two aro cas of Panicum calliphyllum Ashe. 15:99, 100, 19 My Blewitt, A. E. Scirpus figs in Connecticut. Rhodora 15: 98, 99 19 My 1913. lontum in eastern North America: Rhodora 373 374 INDEX TO AMERICAN BOTANICAL LITERATURE Blumer, J. C. Ein Vegetationsbild aus Arizona im Sommer. Bot. Jahrb. Beibl. 50: 1-10. 15 Ap 1913. Bower, F. O. Sir Joseph Dalton Hooker. Bot. Gaz. 55: 384-391. 15 My 10913. Drost, A. W. The Surinam Panama disease of the Gros Michel banana. Jamaica Dept. Agr. Bull. II. 2: 128-149. pl. 20-30. Ja 1913. Translated by S. F. Ashby from Surinam Dept. Agr. Bull. 26. Collins, G. N., & Kempton, J. H. Effects of cross-pollination on the size of seed in maize. U.S. Dept. Agr. Plant Ind. Circ. 124: 9-15. 3 My 1913.” Cook, O. F. Relationships of the false date-palm of the Florida Keys, with a synoptical key to the families of American palms. Contr. U.S. Nat. Herb. 16: 243-254. pl. 74-77. 14 My 1913. Cook, M. T., & Martin, G. W. The Jonathan spot rot. Phyto- pathology 3: 119, 120. Ap 1913. Coville, F. V. Directions for blueberry culture. U. S. Dept. Agr. Plant Ind. Circ. 122: 3-11. 19 Ap 1913. Edgerton, C. W. The stem rot or Hawaiian “iliau’’ disease of sugar cane. Phytopathology 3: 93-98. pl. 8. Ap 1913. omonia Iliau. Emerson, R. A., & East, E. M. The inheritance of quantitative characters in maize. Nebraska Agr. Exp. Sta. Research Bull. 2: §-120. f. 1-21. 1 Ap 1913. Farwell, O. A. Duboisia Hopwoodii—A histological study. Merck’s Report 20:—[1-10]. f. 7-7. My 1o11. Felippone, F. Contribution a la flore bryologique de 1l’Uruguay. Fasc. 2: 1-38. Montévidéo 1912. Includes Barbula uruguayensis Broth. sp. nov. Fernald, M.L. A northeastern variety of Carex paeouter Rhodora 15: 92,93. 19 My 1913. Fink, B. The nature and classification of lichens—II. The lichen and its algal host. Mycologia 5: 97-166. My 1913. Fitzpatrick, H. M. A comparative study of the development of the fruit body in Phallogaster, Hysterangium, and Gautieria. Ann. Myc. 11: 119-149. pl. 4-7+f. 1-7. 30 Ap 1913. Gardner, N. L. New Fucaceae. Univ. Calif. Publ. Bot. 4: 317-374- bl. 36-53. 25 Ap 1913. des ee Setchell & Gardner, gen. nov., Pelvetiopsis Gardner, alid nelu gen. nov., Hali ca Gardner, Blossevillea Brandegeei Setchell & Gardner, and Cystoseira rane saa spp. nov. Goodspeed, T. H. Notes on the germination of tobacco seed. Univ. Calif. Publ. Bot. 5: 199-222. 15 My 1913. INDEX TO AMERICAN BOTANICAL LITERATURE 375 Gould, H. P., & Fletcher, W. F. Apples and peaches in the Ozark region. U.S. Dept. Agr. Plant Ind. Bull. 275: 5-95. pi. 1-6+f. 1-6. 6 My 1913. Graves, A.H. Notes on diseases of trees in the southern Appalachians —I. Phytopathology 3: 129-139. f. 1-10. Ap 1913. Harper, E. T. The probable identity of Stropharia epimyces (Peck) Atk. with Pilosace algeriensis Fries. Mycologia 5: 167-169. My Hassler, E. Oenotheraceae.—J. In Ex herbario Hassleriano; Novitates paraguarienses. XVI.]_ Repert. Sp. Nov. 12: 39, 40. 20 Mr 1913. Includes Jussieua bullata sp. nov. . Hedgcock, G. G. Notes on some diseases of trees in our national forests. III. Phytopathology 3: 111-114. Ap 1913. Herre, A.W. C.T. The lichens of Mt. Rose, Nevada. Bot. Gaz. 55: 392-396. 15 My 1913. Includes Acrospora thermophila sp. nov. Holden, R. Some fossil plants from eastern Canada. Ann. Bot. 27: 243-255. pl. 22, 23. Ap 1913. Howe, R. H. An additional note on Nantucket lichens. Rhodora 15: 93, 94. 19 My 1913. Hull, E. D. Botrychium obliquum and var. dissectum in Berrien Co., Michigan. Torreya 13:112. 6 My 1913. Jehle, R.A. The brown rot canker of the peach. Phytopathology 3: 105-110. pl. ro. Ap 1913. Klugh, A.B. Notes on the algae of Georgian Bay. Rhodora 15: 88-92. 19 My 1913. Includes Rivularia laurentiana sp. nov. Knight, L. I., & Crocker, W. Toxicity of smoke. Bot. Gaz. 55: 337- 371. f. 1-4. 15 My 1913. : Knowlton, F. H. Results of a paleobotanical study of the coalbearing rocks of the Raton Mesa region of Colorado and New Mexico. Am. Jour. Sci. IV. 35: 526-530. My 1913. Knuth, R. Drei neue Arten von Oxalis aus Siid-Amerika. Repert Sp. Nov. 12: 36, 37. 20 Mr 1913. Includes Oxalis lancifolia, O. integra, and O. fuegensis, spp. NOV. Knuth, R. Geranium Purpusti, spec. nov. aus Mexiko. Repert. Sp. Nov. 12: 40, 41. 20 Mr 1913. ; Levine, M. Studies in the cytology of the Hymenomycetes, especially the Boleti. Bull. Torrey Club 40: 137-181. pl. 4-8. 9 My 1913. Melhus, I. E. Silver scurf, a disease of the potato. U.S. Dept. Agr. Plant Ind. Circ. 127: 15-24. f. 1-4. 17 My 1913. 376 INDEX TO AMERICAN BOTANICAL LITERATURE Meyer, R. Echinopsis albispinosa K. Sch. Monats. Kakteenk. 23: 61-63. 15 Ap 1913. Murrill, W. A. Illustrations of fungi—XIV. Mycologia 5: 93-96. pl. 87. My 1913. Includes illustrations of six spécies of Venenarius Nichols, G.E. The vegetation of Connecticut. I. Phytogeographical aspects. Torreya 13: 89-112. f. 1-6. 6 My 1913. Norton, J. B. S. Jonathan fruit spot.. Phytopathology 3: 99, 100. Ap 1913. O’Gara, P. J. _ Studies on the water core: st apple. Phytopathology 3: 121-128. f. r, 2. Ap 1913. Osterhout, G. E. Two new species of Carduus from Colorado. Muh- lenbergia 9: 54-56. 19 My 1913. Carduus modestus and C. acuatus Osterhout. Overton, J. B. Artificial parthenogenesis in Fucus. Science II. 37: 841-844. 30 My 1913. Pickett, F. L. The bacelemei of the embryo-sac of Arisaema tri- phyllum. Bull. Torrey Club 40: 229-235. pl. 13, 14.. 20 My 1913. Piper, C. V. The wild prototype of the cowpea. U.S. Dept. Agr. Plant Ind. Circ. 124: 29-32. 3 My 1913. . Quehl, L. Mamillaria Thornberi Orc. Monats. Kakteenk. 23: 51, 52: 15 Ap 1913. Rigg, G. B. Is salinity a factor in ae distribution of Nereocystis Leutkeana? Bull. Torrey Club 40: 237-242. f. r. My.1913- Rehm, H. Ascomycetes exs. Fasc. 52. Ann. Myc. 11: 166-171. 30 A Plants distributed as Tympanis Fraxini, Phaleoderris Heliopsidis, Pezizella on- tariensis, Botryosphaeria Hamameéelidis, Pseudotthia Symphoricarpi and Uncinula were collected in North America eae H. Ascomycetes novi. Aas. Myc. 11: 149-155. 30 Ap 1913: cludes Naevia canadica, Ombrophila limosa, Pezicula Vecuates Diatrypa potella, and Mycosphaerella lageniformis, spp. nov. from Rorer, J. B. The use of the green muscardine in ‘he control of some sugar cane pests. Phytopathology 3: 88-92. pl. 7. Ap 1913- Schlechter, R.S, Burmannia Wercklei Schltr. nov. spec. aus Costa Rica. Repert. Sp. Nov. 12:35. 20 Mr 1913. Shear, C. L. Some observations on phytopathological problems in Europe and America. Phytopathology 3: 77-87. Ap 1913- Slosson, M. New ferns from tropical America—IlI. Ball Torrey Club 40: 183-185. pl. 3. 9 My 10913. Includes Dryopteris lurida (Jenman) Underwood & Maxon and D. leucochaete Slosson, spp. nov BULL. TORREY CLUB VOLUME 40, PLATE 20 STOUT: BUD-VARIATION IN PELARGONIUM Vol. 40 | No. 8 BULLETIN :. OF THE TORREY BOTANICAL CLUB A botanical cross-section of northern Mississippi, with notes on the influence of soil on vegetation RoLtanp M. HARPER (WITH PLATES 21, 22) INTRODUCTION The coastal plain of the southeastern United States seems to be more diversified geographically in Mississippi than in any other state, with the possible exception of Florida; and the correlations between geology and vegetation are more obvious there than in any other part of the coastal plain, unless it is in western Alabama. These interesting correlations were graphically described by Dr. E. W. Hilgard in his epoch-making ‘‘Geology and Agjiculture of Mississippi” in 1860, in the fifth volume of the Tenth Census reports in 1884, and in his text-book on Soils* in 1906. On pages 490-492 of the book last named there is a special phytogeographical sketch of the northern end of the state, between latitudes 34° and 35°, accompanied by an outline map showing the soil provinces or geographical divisions, and a table giving in very condensed form the chemical and physical characters of the soil and a list of a few of the more characteristic trees in each region. In the summer of 1911 I had occasion to cross the northern half of Mississippi twice, first a little south of the portion last mapped by Dr. Hilgard, and then almost through the center of it. In so doing I crossed all but two or three of the divisions on his map (which form belts approximately parallel to the Mississippi * Reviewed in Torreya 7: 170-175. [The Buttetin for July (40: 305-376: oh mn was issued 18 Jl 1913.] 377 378 HARPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI River, instead of to the coast as in most other parts of the coastal plain, and therefore running north and south), and I took ad- vantage of the opportunity to make a more complete analysis of the vegetation of each division than had been attempted in that part of the country before. The observations made on that trip, besides embodying some previously unrecorded facts, have led to some conclusions which seem sufficiently new to be offered to the botanical public. . Previous literature. Besides the works of Dr. Hilgard already noted, and a few papers on particular regions, which will be men- tioned farther on, the following have an important bearing on the phytogeographical problems of northern Mississippi. Campbell & Ruffner. A physical survey extending from Atlanta, Ga., across Alabama and Mississippi to the Mississippi River, along the _ line of the Georgia Pacific Railway, embracing the geology, topog- raphy, minerals, soils, climate, forests, and agricultural and manu- facturing resources of the country. 8vo. 147 pp. and 2 folded maps. New York, 1883. (Pages 92-96 and 102-107 relate to Mississippi.) a B. Hurt. Mississippi: its climate, soil, productions, and agri- cultural capabilities. U.S. Dept. Agric. Misc. Spec. Rep. no. 3- 89 pp. 1883?. (Contains an annotated list of trees, among other things.) Sargent & Mohr. (Forests of) Mississippi. U. S. Tenth Census 9: 530-536, and colored map. 1884. W. J. McGee. The Lafayette formation (in Mississippi). U.S. Geol. Surv. Ann. Rep. 121: 451-461. 1892. L. C. Johnson. (Underground waters of) Mississippi. U. S. Geol. Surv. Water Supply & Irrigation Paper 114: 171-178. f. 23 (geological map). 1905. A. F. Crider. Geology and mineral resources of Mississippi. U. S: Geol. Surv. Bull. 283. 99 pp. 1906. Crider & Johnson. Summary of the underground-water resources of Mississippi. U. S. Geol. Surv. Water Supply & Irrigation Paper 159. 86 pp. 6 plates (including colored geological map), 11 text- figs. 1906. A. F. Crider. A provisional [sic] geologic and topographic map of Mississippi. 20 X 28 in., colored. First published in 1907, and issued in connection with several different bulletins of the Mississippi State Geological Survey. HARPER: BOTANICAL CROSS-SECTION OF Mississippr 379 C. E. Dunston (of the [U, S.] Forest Service). Preliminary examina- tion of the forest conditions of Mississippi. Mississippi State Geol. Surv. Bull.7. 76 pp. (Not dated, but apparently published two or three years ago). Some of these publications are more useful to botanists than their titles would seem to indicate. There are very few references to the area under consideration in purely botanical literature, doubtless chiefly because nearly all the plants growing there, as far as known, are widely distributed, as I have already pointed out in the case of northwestern Alabama and the Delaware peninsula,* and can be studied more conveniently elsewhere. Itinerary. On June 6, 1911, I crossed the eastern boundary of Mississippi at McCrary, in Lowndes County, and continued to Columbus, Artesia and West Point by the Mobile & Ohio R.R. In the next few days I traveled almost due westward from West Point, in about latitude 33° 30’, on the “Southern Railway in Mississippi” (formerly Georgia Pacific Ry.), stopping at Carroll- ton, Greenwood, Itta Bena, Stoneville and Greenville, and making short excursions on foot from each of these places. On the 1oth I crossed the Mississippi River at Greenville by going ten or twelve miles upstream to Luna Landing, Arkansas (the nearest railroad point on the other side), which gave me a view of the | banks of the river for that distance. On the 16th I re-entered Mississippi near Mineral Wells, in DeSoto County, traveled south- eastward on the Frisco System (formerly Kansas City, Memphis and Birmingham Ry.) to Tupelo, then southward on the Mobile & Ohio to West Point and Artesia, and back to Alabama the same Way as before. Over a year later, namely, on August 31, la I came into the state near Corinth, in Alcorn County, traveling _ Southward on the Mobile & Ohio R.R. to Tupelo, West Point, etc. The notes made on this last trip have been combined with those of 1911 in making up the list of plants for one of the regions, as will be explained presently. Method of treatment. The various soil belts described by Dr. Hilgard are not as easily recognized now as they were when he knew them best, for the increase of population and cultivated fields generally tends to obliterate geographical distinctions. And * Torreya 7: 44-45. 1907; 9: 217. 1909- 380 HarPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI as I crossed the boundaries of most of the regions at the rate of 25 to 40 miles an hour, and without ever having seen them before, I was unable to discern some of the distinctions that have been made by those who have explored the territory on foot and made careful studies of the stratigraphy. In the following pages the results of my superficial study of the more conspicuous vegetation of these belts, as far as I could recognize them, are set forth. Notwithstanding its superficiality, this has one advantage over previous studies of Mississippi vegetation in being quantitative. Under each belt or region named the geology and topography are briefly described, and the percentages of lime, potash, and phosphoric acid in the soil, taken from Hilgard’s report on cotton production in the fifth volume of the Tenth Census, are indicated in nearly every case, for purposes of comparison. Additional literature is cited for some of the regions. All those crossed east of Carrollton and Holly Springs extend southeastward into Ala- bama, where they have been recently described, with quantitative analyses of the forests of each, in my geographical report on the economic botany of that state (Geol. Surv. Alabama, Monograph 8. 228 pp. June, 1913). The plants identified in each belt are divided first into trees, shrubs and herbs, and then arranged in order of abundance or frequency, with a number indicating how many times each was seen, except in the case of belts so narrow that the frequency numbers would be too small to have much significance. Species seen not more than once in a distance of fifty miles are usually omitted. The names of evergreens are printed in heavy type and those of vines in italics. THE REGIONS IN DETAIL From the Alabama line to about three miles west of Columbus, a distance of about 12 miles, the country is underlaid by the Eutaw formation, one of the divisions of the Cretaceous, but the surface is mostly a sandy loam of much more recent age, presum- ably the Lafayette.* As the railroad in this short distance crosses the Buttahatchee and Tombigbee Rivers and traverses a few * A description of this part of Mississippi by Dr. Hilgard can be found in the 5th volume of the Tenth Census, pages 296-298. HARPER: BOTANICAL CROSS-SECTION OF Mississippr 381 miles of the bottoms of the latter, the vegetation visible from the train is mostly that of river-bottoms. And as the soil is so fertile that most of it is under cultivation by this time, there is not much natural vegetation left. Most of the shrubs and herbs seen were introduced species, and it is hardly worth while to enumerate them. The commonest trees seem to be Pinus Taeda, Salix nigra, Liquidambar, Taxodium distichum, Quercus Phellos and Ulmus alata, in the order named. Cretaceous prairie region. From McIntyre (three miles west of Columbus) to Artesia and West Point, thence westward about to the line between Clay and Webster Counties, the country is characterized by a newer Cretaceous formation, the Selma Chalk or Rotten Limestone, with little or no Lafayette loam over it. On the return trip I entered the same belt near the northeastern corner of Pontotoc County, and traversed it lengthwise from Tupelo to Artesia. This is the ‘northeastern prairie region’’ of Hilgard’s Mississippi reports, a direct continuation of the central prairie region or black belt of Alabama, which has been well described by Smith,* Mohr,t} and several less familiar writers. This prairie region is gently undulating or “‘rolling,’”’ with very few springs or small streams. Its soil, mostly a gray calcareous clay, was once considered the most fertile in the state, with the possible exception of that in the ‘‘delta’’ (described farther on), and consequently most of it has been long given over to agricul- ture. Analyses of this soil published by Dr. Hilgard show 0.99- 1.37% of lime (CaO), 0.33-0.86% of potash (K,O), and 0.03-0.27% of “ phosphoric acid” (P.0;). With the possible exception of corn, cotton always has been the principal crop (about 20% of the total * Tenth Census 6: 55-58, 68, 128-140 (pages numbered correctly at bottom). 1884; Geol. of Coastal Plain of Ala. 281-285, 350-352, 533-535> 538-539, 576-577> 585, 605-608. 1894. , ee + Plant Life of Alabama 97-106. 1901. (This however includes two adjoining (op. cit., 10-12), and Crider (U.S. G. S. Bull. 283: 17-19), and in the government soil in both states. 382 HARPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI area having been devoted to that crop as long ago as 1880), but in recent years, especially since the approach of the boll-weevil, a great deal of alfalfa has been raised. A large amount of land also is now devoted to pasturage, as in many other parts of the world where agriculture has been long established. In such a fertile region forests, especially primeval forests, are of course scarce. Some parts of it indeed are said to have been treeless when first discovered, whence the name “‘prairie’’; but it would be hardly possible to determine the location and extent of the original prairie spots with any degree of satisfaction now. However, this is one of the few parts of the South where one can see fields and pastures on the sky-line in many places instead of all woods. The remaining forests are principally of two kinds: oak groves on broad low knolls of poorer soil (Lafayette?), and bottom- land forests near some of the creeks and rivers. The latter doubt- less owe their preservation to the fact that the earlier settlers found such land too difficult to drain, and often too insalubrious to live in; but the growing population continually requires more land, and the bottom-land trees are gradually disappearing before the axe of the farmer. Pines are not seen at all in this part of Mississippi, except an occasional solitary specimen of Pinus Taeda or P. echinata, presumably introduced. Although very different geologically, and not very similar in climate, there are some striking resemblances between the present appearance of this region and the prairie region of Illinois. Both have comparatively level topography and fertile grayish to blackish clay soil, and streams which fluctuate considerably and are muddy most of the time. Both have very little woodland at the present time (they were much less alike in this respect before the country was settled, though), and almost no evergreens. Luxuriant corn-fields make up a large part of the summer land- scape (in Illinois the crop is nearly all corn, but in Mississippi it is about half cotton), and finally in both regions the population is about as dense as extensive agriculture alone will support, and consequently it is practically at a standstill outside of the manu- facturing cities. Just as in the Cretaceous region of New Jersey and Delaware,* the herbaceous flora recognizable from a train consists mostly of * See Bull. Torrey Club 37: 425. 1910; Torreya 12: 22%. 1921. HARPER: BOTANICAL CROSS-SECTION OF Mississippi 383 weeds; but even the weeds are rather characteristic of this region, both in Mississippi and in Alabama. The following list of plants is derived from 128 miles of travel through the prairie region in 1911, and from 69 miles over part of the same route (viz., from Tupelo to Artesia and McIntyre) in 1912. The country between Corinth and Tupelo, traversed on the 1912 trip, although mostly underlaid by the same formation, is not taken into consideration _ in the statistics because it has much more of the superficial sandy loam (Lafayette?), a difference which is strongly reflected in the vegetation. The figures for the two years are kept separate, because one set of observations was made in early summer and one in late summer, which makes a considerable difference in the aspect of the herbaceous vegetation. The first figure in each case is for June, 1911, and the second for August, 1912. Species seen less than three times in traveling these 197 miles are omitted. TREES HERBS 24+7 Quercus stellata 31 +18 Sorghum Longer 21+5 Salix nigra oe +10 os deltoides quidambar Styraciflua ; i: Ss ee Phellos 24+o0 Anthemis Cotula ba Ww 13+5 Quercus falcata? 8+15 Helenium tenuifolium 8+3 uercus marylandica 20+0 Da cus” * s : 7+3 Platanus occidentalis o+19 oe . ricer a 6+4 Hicoria ovata? 19 +0 tag 7+0 Ulmus alata? eh — lone oen: 2 ti I+9 etoc sp. Hele ae nigra 6+4 Ambrosia wee 3+3 Quercus pagodaefolia nie aan serotin 6+0 ee triacanthos (intro- nia Apocyniim Re cata m o+7 Chama ta fasciculata : se dots — (introduced) ets Stoners pevecke vaal 5+0 Hicoria alba? 6+0 Ratibida pinnata 4+2 Quercus lyrata? 4+2 Acne iactatatuns *. x, 6 sp- 3+0 Fraxinus sp. o+ greene bie = cata Curtis sanguinalis 2+1 Cercis Canadensis o+5 Syn I+2 Juniperus virginiana o+5 Bidens sp thiu oO u ee pe Boltonia asteroides 19+4 Prunus angustifolia 4+o Asclepias tu 13+1 Sambucus canadensis 4+0 4+5 Cephalanthus occidentalis 2+2 Typha latifolia 7+1 Brunnichia cirrhos o+4 Solidago sp- 6+ coma radicans o+4 Croton capitatus 3+2 Rhus glabra o+4 Carduus sp- 2+ assafras variifolium 3+0 Andropo: sco 3+0 Smilax glauca 3+0 Cyperus pseudovegetus 3+o Rubuss o+3 Conoclinium coelestin 2+ ria macrosperma o+3 § tiphium | terebinthinaceum Ga ap Gpoconaies indicus 384 HarPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI Assuming the above figures to represent correctly the relative abundance of the species, less than 2% of the woody plants (i. e., individuals, not species) are evergreen. The significance of this fact will be discussed farther on. The oaks in this region, as in the ‘“‘delta’”’ region to be de- scribed below, are rather puzzling, especially to one who has never seen them before and has no opportunity to stop and examine them closely in the light of descriptions or specimens. Dr. Hilgard* has described some curious forms of oaks in this very region, which ought to be investigated by a competent taxono- mist. The comparatively large number of herbs, and the occur- rence of a few genuine prairie species, such as Ambrosia bidentata, Silphium laciniatum, S. terebinthinaceum, S. perfoliatum, Mesadenia tuberosa, and Polytaenia Nuttallu (the last three seen only once, and therefore not listed), are reminders of the prairie conditions that once existed in this region. Pontotoc Ridge. The next younger formation in Mississippi is the Ripley, the uppermost division of the Cretaceous. Its strata are more sandy and therefore less easily eroded and dis- solved than the Rotten Limestone, and they give rise to a belt of hills, rather rugged for the coastal plain, known in Mississippi as the Pontotoc Ridge. According to Dr. Hilgard’s analyses its soil contains 0.17-0.28% of lime, 0.15-0.37% of potash, and 0.08-0.11% of phosphoric acid; or approximately half as much of these important materials as in the prairie belt.t The Ripley formation and its corresponding topography are wanting in the latitude of Columbus and Greenville, but on the return trip I traversed it for about 15 miles, between New Albany and Sherman. Although the soil of the Pontotoc Ridge is perceptibly less fertile than that of the neighboring prairies, it is mostly under cultivation now, and one does not get a very accurate idea of its native vegetation by merely crossing it once on a fast train. In the following list the numbers are omitted, because they would be too small to have much significance. * Soils 491, 404, 498-502. } For descriptions of this part of Mississippi see Hilgard, Geol. & Agric. Miss. 83-92. 1860; U. S. Tenth Census §: 221-223, 292-294. 1884; and the U. S. soil survey of Pontotoc County by Bennett and Winston, 1907. HARPER: BOTANICAL CROSS-SECTION OF Mississipp1 385 TREES SHRUBS Salix n Sambucus canadensis Liauidambar Slater Robinia Pseudacacia Cornus runus angustifolia Pinus e china ta Sassafras variifolium Platanus occidentalis Rhus glabra Quercus alba Puieaa f a ERBS Cercis canadensis Asclepias tuber Liriodendron Tulipifera Tripsacum dactytoiies Populus del Cicuta C Quercus marylandica Sorghum Halepense Quercus stellata (and several less frequent weeds) A noteworthy difference between this list and the one next preceding is the presence of Pinus echinata* and Liriodendron, species which do not grow in the richest soils. I am not sure that any of the shrubs and herbs listed could have been found in pre- historic times in the places where I saw them. Post-oak flatwoods. The oldest Eocene formation in Missis- sippi underlies the “‘ post-oak flatwoods,” which lie immediately west of the Pontotoc Ridge in Union County and of the prairie region in Clay County, and extend southeastward into Alabama. This belt is nowhere more than 15 miles wide, and where I crossed it on the westward trip its boundaries seemed so ill-defined, that I could not very well separate my notes on it from those on ad- jacent regions. On the way back, however, I had little trouble in locating its western edge near Hickory Flat, and its eastern edge at the western base of the Pontotoc Ridge, near New Albany. The Lafayette formation seems to be absent in the flatwoods, and the Eocene strata have weathered into a very stiff clay, con- taining, according to Dr. Hilgard’s analyses, 0.08-0.18% of lime, 0.25-0.75% of potash, 0-0.05% of phosphoric acid, and 0.17- 0.85% of magnesia. Both its physical and its chemical properties make this soil ill adapted to agriculture, and the region is very thinly settled. This part of Mississippi has been described by Hilgard,j Hurt,t and E. J. Hill,§ and in the government soil surveys of Pontotoc, Clay and Oktibbeha Counties. 20129 Pie, Sareenld wag of the pine forests of Mississippi (opposite page 53° of the oth volume of the Tenth Census) does not indicate the occurrence of any pine at all in this region t Geol. & Agric. Miss. 273-288; Tenth Census 5: 224-227, 294-296. $ Op. cit., 12-14. § Torreya 6: 231-232. 1906. In this interesting paper, based on observations made in Oktibbeha County in 1858, the post-oak flatwoods west of Starkville are 386 HARPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI Hickory Flat and New Albany are only 14 miles apart, and between those points I had less than twenty minutes in which to gather the following list of plants. TREES HERBS Pinus wesc tie Sorghum Halepense Quercus stella Helenium tenuifolium a eae Typha latifolia nigr E igeron ramosu Faais grandifolia Tripsacum dactyloides ‘(Quercus Phellos Rudbeckia hir Quercus marylandica Scirpus teases ‘Quercus falcata SHRUBS Sambucus canadensis Rhus glabra Arundin aria Cephalanthus occidentalis This list is too short to draw many important conclusions from, but the fact that a pine stands at the head is significant. The second tree, Quercus stellata, is the one which gives the region its name. Forests of very similar aspect can be seen in the “barrens” of extreme northern Alabama and in the Paleozoic flatwoods of the Coosa valley in Georgia and Alabama. Eocene red hills. The more hilly portions of the Eocene (i. e., excluding the flatwoods) are traversed by the Southern Railway from the eastern part of Webster (formerly Sumner) County to Carrollton, and by the ‘‘Frisco”’ from the Mississippi River to Hickory Flat. Geologists subdivide the Eocene into several formations, but with the exceptions named above and below, I was not able to correlate these with any marked differences in topography or vegetation. The Southern Railway keeps pretty close to a small river nearly all the way across Webster County, and.that probably makes more difference in the appearance of the country between that county and those immediately west of it than the difference in age of the underlying rocks does. West of a line drawn from about Holly Springs to Carrollton, though, the Eocene is covered by a superficial formation which affects the vegetation enough to warrant a geographical separation. called pine-barrens, evidently on account of the contrast with the pineless prairie region bordering them on the east. (The “ Aletris obovata” mentioned, judging from the EAN given, and a specimen which I have since seen in the author’s her- barium, seems to have been wrongly identified. The locality is several hundred miles pees: any known station for that species, and in a very different kind of country.) HARPER: BOTANICAL CROSS-SECTION OF MissiIssipp1 387 The typical Eocene country of northern Mississippi is moder- ately hilly, with reddish to yellowish clayey soil, some of it derived from the weathering of the Eocene strata, and some belonging to the much more recent Lafayette. According to Dr. Hilgard’s analyses these soils contain 0.05—0.09% of lime, 0.09-0.24% of potash, and 0.02-0.09% of phosphoric acid; which is a smaller percentage of the first two than in any other region here described. Like most other parts of northern Mississippi, it is now mostly under cultivation, and primeval forests are scarce. The same belt extends northward into Tennessee and eastward to South Carolina, if not farther.* The following plants were noted more than once in Webster and Montgomery Counties and the eastern half of Carroll on June 7th, or between Holly Springs and Hickory Flat on the 16th; a total distance of 77 miles. Sea HERBS 32 Pinus echina 15 Plantago aristata ay Liguidambar Shyractihes tr Rubdeckia 21 ix n 8 Anthemis Cotula 9 Taxodi ar diiatle hum 47 Physostegia s 7 Liriodendvon Tulipifera 4 Erigeron ramosus 7 Quercus Phellos 5 Daucus pusillu 7 Quercus falcata 4 — caroliniana 6 Pinus Taeda 4 urus cernuus 6 Quercus marylandica 4 Cicuta Curtissii 5 Quercus al 3 ghum se 5 Cornus florida 3 Helenium tenuifolium 4 Quercus nigra 3 Cyperus pseudovegetus 3 Carpinus caroliniana 3 Ascle tube 3 Fagus grandifolia 3 Boltonia asteroides 3 Platanus occidentalis 3 Juncus aristulatus 2 Quercus stellat 2 Tripsacum dactyloides 2 Ulmus 2 Eryngium yuccaefoliu 2 Quercus Michau 2 Koellia flexuosa 2 Populus deltoides 2 Rhexia lanceolat 2 Cercis canaden 2 Typha latifolia — 2 Ca pean dentate 2 Ambrosia artemisiaefolia 2 Juncus effusu’ —_—_ 2 Panicum scoparium 18 Sambucus canadensis 2 Craccea virginia 13 Prunus angustifoli 5 Brunnichia cirrhosa 5 Rhus glabr 3 Alnus rugosa 3 Rhus copallina 2 Rubus sp 2 Cephalanthus Sa 2 Robinia Pseudaca th Census 5: 231I- * For descriptions of this part of Mississippi see Hilgard, Ten For notes on the 235, 301-302; and the U.'S. soil survey of Montgomery County. * South Carolina end of the same belt see Bull. Torrey Club 37: 411. 1910; 38: 225. EOE: 388 HarPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI The pines are probably relatively more abundant now than they were originally, on account of their tendency to spread in old fields, so that the apparent proportion of evergreens among the woody plants (19 per cent) may be too large. But it is very evident that the soil of this region is better adapted to evergreens than is that of most of the other regions discussed in this paper. Ericaceae seem to be entirely absent, and native monocotyledons are not conspicuous. (There is no telling how many of the shrubs and herbs listed are really indigenous here, but probably not more than half.) Of the species enumerated, Taxodium and Brun- nichia are almost confined to the coastal plain, and Quercus Michauxti and Populus deltoides mainly so as far as the south- eastern states are concerned; but the other species are pretty widely distributed. Yellow loam region. Between Memphis and Holly Springs the topography is much the same as in the region just described, but the surface is covered with a few to several feet of loess, a buff- colored very fine-grained somewhat calcareous silt,* which makes the vegetation considerably different. This is the ‘yellow loam region”’ or ‘‘brown loam table-lands” of Hilgard, which has no counterpart anywhere to the eastward. Dr. Hilgard’s analyses show in this soil 0.24—0.25% of lime, 0.30-0.55% of potash, 0.07% of phosphoric acid, and 0.31-0.48% of magnesia. Almost every acre of it, except on the immediate banks of streams, has been cultivated at some time or other, and much of it is now badly gullied. No primeval forests were seen, and no shrubs or herbs other than weeds, but probably nearly all the trees listed below grew in the same region in prehistoric times, even if not in exactly the same places where they are now. The following list covers 44 miles, about 12 of which were in Tennessee, but so close to the Mississippi line as not to cause any appreciable error. TREES 3 Liquidambar Styraciflua 12 Salix nigra 2 Platanus occidentalis : ee falca 2 Gleditschia triacanthos saree 2 —- virginiana + tte stellata 2 Quercus alba * For dexpiptions of this soil see the works of Hilgard and McGee mentioned near the beginning of this paper, also Hilgard, Am. Jour. Sci. gt: 319-321. 1866; T. O. Mabry, Jour. Geol. 6: 273-302. 1897. HARPER: BOTANICAL CROSS-SECTION OF Mississipp1 389 TREES (continued) HERBS I Quercus nigra 5 Plantago aristata I Quercus Michauxii 3 Rubdeckia hirta I Taxodium P goserats: 2 ~~ I Quercus v 2 Asclepias tuber I Pe Tulipifera 1 Tripsacum dactyloides r Daucus A sae s SHRUBS 1 Euphorbia sp. I2 Prunus angustifolia rr Sambucus canadensis ni Rhus aes nigra is partly evergreen (probably less so here than nearer the coast, though), but with this possible exception no evergreens were seen. Ericaceae and Cyperaceae are likewise rare or absent. Bluff region. Carrollton is near the brow of a line of bluffs which border the flood-plain of the Mississippi River all the way from Memphis to Vicksburg, and form one of the most prominent topographic features in Mississippi. From Carrollton station (North Carrollton P. O.) the railroad descends rapidly for six or eight miles to the foot of the bluffs, following the valley of a creek. I walked the railroad from North Carrollton to Mal- maison, which is near the edge of the flood-plain, and from there northward up into the bluff hills about two miles and back. Geologists generally map the surface formation of these bluffs (‘cane hills,’ they are called farther south) as loess, but what I saw of it in Carroll County looked almost exactly like the “second bottom” loam along many rivers in the coastal plain of Alabama, and quite different from the more typical loess which I saw a few days later in Arkansas, and between Memphis and Holly Springs. The steeper slopes of the bluffs are strewn in many places with subangular cherty pebbles. The plants seen along and near the railroad and creek between Carrollton and Malmaison are mostly species characteristic of bottom-lands, as follows. TREES Taxodium distichum _— Michauxii hooey Styraciflua " Acer Negundo Ss : Oo lyrata Diktannec occidentalis Jug nigra Cercis canadensis sro dron Tulipifera Carpinus caroliniana aarp aad Betula nigra Ulmus Morus rubra 390 HARPER: BOTANICAL CROSS-SECTION OF MISsSISSIPPI RUBS HERBS Sambucus canadens Cicuta Curtissii Cephal ae ocuidentals Saururus cernuus Brunnichia c rrhosa Juncus ndinari sisete cs ommelina hirtelia Hydrangea quercifolia Carex oe is? pst ordata Homalocenchrus oryzoides Rhus g Onoclea sensibilis ree catia arborescens Carex lupulina? Rosa carolina Spirodela polyrrhiza Itea virginica Panicum scoparium Teucrium s Diodia teres* Euphorbia corollata etc. Up on the hills north of Malmaison the vegetation is much like that of ordinary dry woods at moderate altitudes throughout the South, as the following list shows. TREES HERBS Pinus echinata Cracca virginiana ercus marylandica Pteridium aquilinum or orida Meibomia laevigata Quercus falcata Mei ia Michauxii Quercus stellata Koellia flexuosa Que ‘us Psoralea pedunculata Hicoria alba Ante Z nnaria plantaginifolia Quercus coccinea pears acrostichoides Mitchella repens HRUBS Dioscorea villosa Ceanothus americanus Hydrangea quercifolia Judging from the vegetation, the soil of these hills must be considerably less fertile than that of the same line of bluffs farther south, as described by Dr. Hilgard (who apparently never visited Carrollton and vicinity). Pinus echinata, the commonest tree, does not seem to have been reported from this part of Mississippi at all before. It seems never to grow on loess (according to Call, Geol. Surv. Arkansas 1889?: 184. 1891), or in any other very rich soil. This and several other species in the list can stand frequent forest fires, but Hydrangea and the three evergreen herbs and two vines grow mostly in ravines, where they are pretty well protected from fire. The *‘ Delta’ (PLATE 21). Next is the “‘ Yazoo delta”’ or Missis- sippi bottom. Many of its features can be matched fairly well in * On sand-bars along the creek between Carrollton and North Carrollton. This species grows in quite a variety of habitats, mostly unnatural, but they all have at least one character in common: exemption from fire HARPER: BOTANICAL CROSS-SECTION OF Misstssippr 391 the flood-plains of smaller rivers, but they are here exhibited on a far larger scale than anywhere farther east. On the Mississippi side the ‘‘delta’”’ extends from a few miles north of the northern boundary of the state down to Vicksburg, nearly three degrees of latitude, or 200 miles in a straight line, and has its maximum width of 60 miles about midway, or just in the latitude where I crossed it. Generally speaking, it is a vast plain, sloping gently southward, at a rate of about a foot to the mile, traversed by numerous crooked, sluggish, muddy rivers and bayous, whose banks are usually a little higher than the interstream areas. The larger rivers of the delta, such as the Yazoo and Sunflower, are thirty or forty feet deeper at high water than at low water, have a perceptible current, and are navigable for steamboats most of the year, while the smaller bayous are stagnant much of the time, and fluctuate comparatively little. At least 90% of the area is or has been subject to occasional inundation from the spring floods of the Mississippi River (‘‘Father of Waters”), but the building of levees along the banks of the great river, in the last half century or so, has considerably restricted the overflows. The soil is mainly a fine gray silt, coarsest on the banks of the larger streams, where the current in times of flood is swiftest. Analyses published by E. A. Smith and E. W. Hilgard show that it contains 0.26-1.35% of lime, 0.30-1.10% of potash, and 0.11— 0.30% of phosphoric acid. This is one of the most fertile soils known, according to Dr. Hilgard (Soils, 116, 345), and very little commercial fertilizer has been used on it as yet. Most of the area seems to be under cultivation now (cotton being the principal crop), but there is still considerable primeval forest with splendid. hardwood timber.* wer There is much valuable information about this region in Humphreys and Abbot’s voluminous government report on the- physics and hydraulics of the Mississippi River, 1861 (reprinted with additions in 1876). Other references are: E. A. Smith,. Proc. A. A. A. S. 20: 251-262. 1872; C. G. Forshey, Proc. A. A. A. S. 21: 78-111. 1873; Hilgard, Tenth Census §: 85-88, 241-247, 319-321. 1884; Campbell & Ruffner, op. cit. 93-96, 105-107; © OY ae te the middle of the delta, but from all accounts the northern half Seems to be most extensively cultivated, and forests consequently more prevalent. in the southern half. 392 HARPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI Hurt, op. cit. 19-24. The latest paper on the area under con- sideration is Bulletin 244 of the Office of Experiment Stations, United States Dept. Agriculture, a ‘‘Report on the Belzoni* drainage district in Washington County, Mississippi,’’ by H. A. Kipp, 1912. This comprises 55 pages, a map and several diagrams; and from the list of bench marks occupying the last six pages one ‘can get a crude idea of the trees of the region and their relative abundance. Running north and south near the middle of the ‘‘delta’”’ is a narrow low ridge, described by Smith and Hilgard as the ‘‘dog- wood ridge,’’ and said to be above the reach of all floods. Where I crossed it, about Itta Bena, this ridge seems to be only a few inches high, and it would be hardly noticed by any one not making a special search for it. Even the inhabitants of that neighbor- hood who were interviewed on the subject seemed never to have heard of it. (It is doubtless more noticeable farther north.) I walked along or near the ridge from Sheppardtown to Itta Bena, eight miles, but the vegetation along there did not seem different enough from that of the rest of the region to be listed separately. The following plants were seen more than once on June 7th to 9th, inclusive, from the train between Malmaison and Green- ville, and on side trips on foot from Itta Bena, Elizabeth and Stoneville, a distance of about 75 miles in all. TREES 2 Morus rubra 31 Taxodium distichum 2 Carpinus caroliniana 26 pags erg Styraciflua 2 Ilex opaca 25s 2 Nyssa sylvatica 18 Chedicetie triacanthos 2 Hicoria ovata 17 Populus deltoides 2 Quercus texana? 12 Ul lata? II Quercus pagodaefolia SHRUBS . fone Phellos 25 Pode apices cirrhosa niflor: 18 Sabal é ni sotaas easton 6 Sambucus ‘canadens 5 Quercus Michauxi: 5 Arundin panies 4 Platanus cocoate 3 Cashelent ras occidentalis 4 Que cus ni 3 Tecoma radicans 4 Fraxinus s = i 2 Rhus radicans 4 Celt 2 Bignonia crucigera 4 Acer vabeaie tridens? 2 Berchemia scandens 3 Planera aquatica 2 Adelia acuminata 3 Quercus lyrata us americana? * Also (perhaps more commonly) spelled Belzona. ft Some of this may be Ulmus crassifolia, which I was not acquainted with at that time. HARPER: BOTANICAL CROSS-SECTION OF Misstssippr 393 HERBS 14 Anthemis oe 2 Carex Crus-corvi 2 Ambrosia Se oe us 4 Sorghum Wiiewe 2 Saururus c 2 Monarda credo 3 Dracopis amplexicaulis There are probably few places in temperate regions where one can see more species of trees in traveling a similar distance through an essentially homogeneous region. Six of those listed are oaks, and if I had been more familiar with the Mississippi valley repre- sentatives of that genus I might have identified still more. The only evergreen tree in the list is I/ex opaca,* and that is confined to the highest and driest spots, and constitutes considerably less than I per cent of the arboreal vegetation. As in other hardwood regions,} the woody plants greatly outnumber the conspicuous native herbs. The abundance of woody vines seems to be char- acteristic of alluvial habitats and some other rich soils, in various parts of the world.t The herbs listed are nearly all weeds. Taxodium distichum, the commonest tree, is not found on the banks of the navigable streams of the delta, for it apparently cannot stand more than ten or twelve feet of average seasonal fluctuation of water.§ It abounds along the smaller bayous or “lakes,” and in the interstream swamps described by Smith, Hilgard and others. At the time of my visit the branches of most of the cypress trees in this region and among the bluffs near Carrollton had many branches dead at the tips. The cause of this condition was not ascertained, but drainage operations may have had something to do with it. Populus heterophylla, which is said by Sargent|| to be especially abundant in this region, I saw only once, and that in the outskirts of the city of Greenwood, where it might not have been in- digenous. Besides the species to be noted presently as conspicuous by their absence all the way across the state, the following were not seen at all in the ‘‘delta”’: tree in New England and in * This seems to i the only angiosp iS several interior s t See Bull. Sain ed He Ae Igo. t See Torreya ro: § See Science II. mee prio 28 No 1912. || Silva N. A. 9: 164. 1896. " 394 Harper: BOTANICAL CROSS-SECTION OF MISSISSIPPI Terrestrial oe Quercus falcata Pinus (all species) Quercus marylandica aca us virginiana Liriodendron ag rugos Sassafras Nyssa biflora rhe nes stellata Banks of the Mississippi (PLATE 22). Of the banks of the “Father of Waters”’ I can say very little, having seen them only for a few miles, between Greenville, Miss., and Luna Landing, Ark., on June roth. Nuttall passed by there in January, 1820, and pub- lished some observations on the river-bank vegetation in his ‘Journal of travels into the Arkansa territory”’ the following year. Lyell traversed part of the same route in March, 1846, and his observations can be found in his ‘‘Second visit to the United States,’ 2: 163-164. 1849. On the inner sides of bends Salix nigra (?) is seen everywhere on sand-bars between high and low water marks, and Populus deltoides on more silty soil a little higher up. These trees can probably stand as much seasonal fluctuation as any in the world. They seem to prefer soils poor in nitrogen and rich in inorganic plant foods, and they must be regarded as the pioneers for that particular type of soil. On the outer sides of bends, where the river is eroding its banks, Liguidambar, Arundinaria, and various other species characteristic of river-bottoms are visible, together with occasional specimens of Taxodium in the swamps farther back. CONCLUSION Notable absentees. Of the plants which are common in other parts of the coastal plain and rare or absent in northern Mississippi the following occur to me. Nyssa biflora was seen only twice, once in Clay County and once in Lowndes; and Juniperus was seen only once on the IgII trip, that in Lowndes County. (It is rather.common in the black belt of Alabama.) Ferns are rare, as are nearly all pine-barren plants. The following were not seen at all. Pinus palustri Myrica cerifera Taxodium tenbekcattens Magnolia (all species) Tillandsia* Ericaceae Orchidaceae * In crossing the ‘‘delta’’ I must have been pretty close to the range of Tillandsia usneoides, for Nuttall (Travels, 228), in floating down the Mississippi on Jan. 22, HARPER: BOTANICAL CROSS-SECTION OF Mississippr 395 Relation of flora to precipitation and soil texture. The absence of Takodium imbricarium, Magnolia glauca, and other bog plants from northern Mississippi is correlated with the seasonal distribu- tion of rainfall, among other things. In the greater part of the coastal plain, summer is the rainy season; but in the “ Mississippi embayment” portion, which is farthest inland, the winters are wetter than the summers, just as in the interior hardwood region, of which this might be regarded as forming a part. At Water Valley, Yalobusha County, which is pretty close to the center of the northern half of Mississippi, meteorological records for a period of twenty years show that only 30.3 per cent of the normal annual precipitation comes in the four warmest months, June to September, and 41.9 per cent in the six warmest months, May to October.* (March is usually the wettest month and October the driest.) This type of seasonal distribution of rainfall makes all streams, and the ground-water too, high in spring and low in fall; while in the pine-barren portions of the coastal plain the greater evap- orating power of the sun in summer is largely counterbalanced by the increased rainfall at that season, and consequently the water- level is much more uniform there, and conditions are favorable for the development of peat and of bog plants. Ponds and swamps are scarce in northern Mississippi, except in the ‘‘delta,’’ where they are caused by the topography, in spite of the climatic conditions just described. Another factor perhaps still more important in determining 1820, first met it at Cypress Bend, about 20 miles below the mouth of the Arkansas River; and one of the bends near aay is named “Spanish Moss Bend.”” The same plant was reported from along the Cumberland River in Stewart County, Tennessee, by Dr. J. M. Safford, state ign of Tennessee, in Garden & Forest 3: Si. * For data of seasonal distribution of rainfall in some other parts of the coastal Vv. plain see Bull. Torrey Club 37: 415~416 (footnote). Ann. Rep. 3: 21 5. Iori. The interior hardwood region has been br usually a little warmer than April; and furthermore it is usually drier than April in the regions that have dry summers, and wetter than April in the regions that have wet summers, so that the figures for May to October give greater contrasts than do those for April to September. 396 HARPER: BOTANICAL CROSS-SECTION OF MISSISSIPPI the character of the flora of northern Mississippi is the prevalence of clayey soils, contrasting strongly with the sand of the pine- barrens nearer the coast. Sand is here chiefly confined to the beds of creeks and rivers, as it is in most of the interior hardwood region. The character of the soil may not be wholly independent of the seasonal distribution of rainfall, but it would be too much of a digression to discuss the matter here. Suffice it to say that it happens that in the Eastern United States most regions with wet winters and dry summers have fertile clayey soils, and where the reverse is true sandy soils predominate. Some relations of vegetation to soil chemistry. Dr. Hilgard, in his earliest and latest books (1860 and 1906), and in various other works, has always stressed the importance of chemical composition of soil, especially the percentage of lime, in determining the character of the vegetation. In his ‘‘Soils,’’ page 490, as already noted, he gives the lime percentages for each soil belt of northern Mississippi, and describes the corresponding vegetation briefly. Similar correlations have been made in Europe by a number of investigators, and the fertility of calcareous soils has been long proverbial. Limestone is one of the commonest and most easily recognized minerals, so that such correlations are easily made; but in the light of the observations made on this trip, and other recent investigations, it is highly probable that some of the other mineral ingredients of soils which do not manifest themselves so conspicuously may be equally important to vegetation. In the case of lime there always has been a difference of opinion as to just how much of it a soil should contain to be called cal- careous. According to Dr. Hilgard, in Europe a soil is not usually called calcareous unless it effervesces with acid, which requires about 5% of lime (calculated as CaO)*; while in this country many soils containing only 1% of lime differ as much in their vegetation from those which have less as they do from some derived from nearly pure limestone. He concludes that calcareous soils are distinguished better by their vegetation than by any arbi- trary chemical stahdard; but here another difficulty is encoun- tered. Just what is calciphile vegetation? ; Various European investigators, Schimper} for example, have * In this connection see Plant World 15: 300-301 T See pages 94-106 of his Plant roam de English aes 1903. HARPER: BOTANICAL CROSS-SECTION OF MissiIssippr1 397 published lists of calciphile plants for particular regions, and in temperate eastern North America certain plants, mostly trees, have become by common consent, as it were, accepted as indicators of calcareous soils. Dr. Hilgard lists quite a number of these in each of his books, and even goes farther and distinguishes dif- ferent forms.of the same species characteristic of different kinds of soil. But none of these writers seem to mention any characters which their calciphile plants have in common, so that if a person totally ignorant of mineralogy should travel around the world he could identify the limestone regions only by knowing the individual species which have been listed as calciphile; and the flora changes almost completely every thousand miles or so in temperate regions, As a matter of fact, the supposed lime-loving trees listed by Hilgard and others in this country do have some characters in common. They are all deciduous except the cedar (and I have recently shown that that is not necessarily calciphile*), and many of them have durable dark-colored heart-wood, thin leaves, and large seeds. But trees with similar characters, and indeed most of the same species, can be also found in many places where the soil is poor in lime or at least not commonly regarded as cal- careous, and associated with other species which have not been hitherto regarded as calciphile. Coville in his work in Arkansas a quarter of a century ago found that the difference in the vegeta- tion of sandstone and limestone areas in close proximity was more quantitative than qualitative; i. e., the species were nearly the same, but their relative abundance differed considerably in the two areas.t : Comparatively few observations on calciphile vegetation in tropical and cold-temperate regions seem to have been made. In Schimper’s Plant Geography less than a page (380) is devoted to the effects of lime on vegetation in the tropics, and he believes its influence to be less there than in temperate regions. In ex- treme southern Florida, including the Keys and most of the main- land south of Miami, the rock is nearly all limestone,{ and there * Torreya 12: 145-154. I9gI2. t See Arkansas Geol. Surv. Rep. 1888!: 246-247- 18 . tI have no analyses of the mainland rock, but that a the Upper Keys is sai to be over 90% calcium carbonate. 398 Harper: BOTANICAL CROSS-SECTION OF MISSISSIPPI is very little soil on top of it. But almost none of the trees regarded as calciphile in the states farther north are found there, and the question arises, are the numerous species of trees growing on limestone in South Florida (nearly all of them tropical) true calciphiles? They are certainly very different in aspect from the supposed lime-loving trees of Mississippi, being nearly all ever- green. Furthermore, some of the commonest species, the pine especially, flourish equally well in sandy soils a little farther north, which are very poor from an agricultural standpoint. Let us see now if there is not some soil ingredient other than lime which is of fundamental importance to vegetation. One of the most convenient and at the same time perhaps the most sig- nificant characters of arboreal vegetation that can be expressed quantitatively is the percentage of evergreens, and this has been already indicated roughly for most of the soil belts described above. The percentages of certain soil ingredients also have been given, and it will be noticed that in the regions under consideration the evergreens can be correlated with potash just about as well as with lime, evergreens being scarcest in the soils richest in potash. This relation is still more apparent when we compare northern Mississippi, where nearly all the soils are pretty well supplied with potash, with Florida, where soil conditions are very different. Florida has a larger proportion of evergreens than any other state in the Union, and at the same time its soils are poorest in potash, though fairly well supplied with lime.* It is altogether likely that the limestones of South Florida above mentioned are deficient in potash, though I have too few data on this point as yet. Statistics collected by Hilgard in his book on Soils show that the ‘percentage of potash in tropical soils is usually less than in those of ‘temperate regions; and it is barely possible that the prevalence of evergreens in the tropics may be due partly to this fact, and not to climate alone, as has been hitherto supposed. In temperate * On page 11 of Bulletin 85 of the U. S. Bureau of Soils, published late in r9t2, there is a table of the average chemical composition of the soils of each state, based on over a thousand analyses. The average of 88 samples from Florida is 0.03% of potash and 0.31% of lime. The corresponding figures for four typical hardwood states are as follows: Kentucky, 92 samples, 0.35% potash, 0.18% lime; Ohio, 57 samples, 0.25% potash, 0.29% lime; Tennessee, 144 samples, 0.28% potash, 0.20% lime; West Virginia, 14 samples, 0.53% potash, 0.16% lime. HARPER: BOTANICAL CROSS-SECTION OF MiIssIssIpPI 399 climates evergreens abound in and near peat bogs and are scarce in clayey soils, and it is well known that potash is scarce in peat and abundant in clay. Just why potash should be antagonistic to evergreens is an ecological problem that need not.be discussed here; for at present I am merely pointing out the geographical correlation. The reasons why this correlation has not been made before are probably first because potassic rocks are not conspicuous and identifiable at sight like limestone, and second because the percentage of potash in the soil varies between much narrower limits than does that of lime, and most soils in inhabited regions have enough potash for the average plant. But it has been proved by many experiments that in an artificial soil containing no potash at all nothing will grow; and it is therefore reasonable to assume that between this artificial condition produced in the laboratory and natural soils containing an average amount of potash there must be intermediate stages where the vegetation is very different from what it is on the average soil. As potash seems to be more abundant in leaves than in any other part of a plant it is natural that a deficiency of this substance should manifest itself first in the leaves, and that plants which do not have access to much potash should have smaller leaves than those of rich soils, and keep them longer. GEOLOGICAL SURVEY OF ALABAMA, UNIVERSITY Explanation of plates 21, 22 Fic. 1. Level dry woods about two miles north of Sheppardtown, Leflore County, approximately on the “ dogwood ridge.” The most conspicuous trees are Nyssa sylvatica (in foreground), Quercus nigra, and Ilex opaca. June 8, 1911. Fic. 2 at Quito, Leflore County, looking west from wagon bridge. Trees mostly Taxodium distichum and Nyssa uniflora. The dead branches in the water indicate the absence of current. June 8, 1911. Fic. 3. Eroding bank of Mississippi River on outer side of bend about two miles above Greenville, Washington County. Shows Taxodium distichum, Populus tern) bank of river a little farther upstream, June 10, rgort. Nearer view of same (eas showing numerous vines, Platanus occidentalis, etc. Studies in the Agalinanae, a subtribe of the Rhinanthaceae* FRANCIS W. PENNELL II. SPECIES OF THE ATLANTIC COASTAL PLAIN Following a survey of the nomenclatural history of the group of Rhinanthaceous plants, which we would term for convenience the Agalinanae, there should properly come some comparative morphological study of the several genera. It has seemed advis- able, however, to defer such for the present, in order first to present the results of a season’s field-study of the group in the Coastal Plain of the South Atlantic and East Gulf states. The present paper includes a revision of the species known to occur in the Atlantic Coastal Plain from New Jersey to eastern Louisiana. The flora of the northward extension of this area— Long Island and southeastern Massachusetts—has not been included, as this region is so narrow and broken, and the Coastal Plain flora so attenuated, as scarcely to make such inclusion desirable. From New Jersey southward and westward to central Alabama the Fall Line has been followed as the natural inland boundary. But toward the west instead of turning northward to include the lower Mississippi Valley, an arbitrary line has been drawn on the 33d parallel to the Mississippi River, the western limit of this study. It is believed that while sufficiently diverse the flora of the region included has so much in common, and so much in contrast to the districts northward and westward as to warrant a special consideration. Four genera of Agalinanae occur within this area. Macranthera is monotypic and wholly restricted to t Afzelia with two species east of the Mississippi River 1s nearly sO restricted, Aureolaria, primarily a genus of the Appalachian ‘district, has several species adapted to this region, while aesins , the largest genus, here reaches its greatest diversity and abundance. * Contribution from the Botanical oe ad of the University of Of these his region, Pennsylvania. 402 PENNELL: STUDIES IN THE AGALINANAE In the late summer and early autumn of 1912, I was able to spend over two months in field-study of this group, and the following itinerary indicates the more important points in the Coastal Plain where collections were made: Abita Springs, St. Tammany Parish, Louisiana............. August hie Catalpa, West Feliciana Parish, Louisiana................. - o-2 Biloxi, Marrison Co, Mississippi). .s oe 6 be hs os tS 29-20 ‘Theodore, Mobile Ca, Alabama, 23.0024 525 8: August 30-Sept. 4 mene, Momie CO Alabama fo eos a op epee Sept. Bay Minette, Baldwin Co., sirainaniaie Sieae Pa Seat Sta te a sree ib bone gi 8 Hiow, Santa Rosa: Co. Ploridaes. 2S oe ee ee * 9 Milligan, Santa Rosa Co., iets Wrasse: y wor tars ecnt@ena eave I ae ae) P4orala, Covington Co-, Alabama : 2.0. 20 ee x II-15 Gripiey,- Washington Co,; Mlorida) 20, 272i ee es e 16-18 Apalachicola, Franklin Co., wloda Silky GA cA VERS MER eee 43 19-21 allahassee, Leon Co., F Ee aS bases ae Se 8c oe Se ie he eae ve 22-24 Ss ERS WARN (Ore PIOTIda oo oes pase si aee eae oe ss 25-26 Monticello, Jetferson Co., Florida... 665 sy a7 Helena Thomas Co. Geeteth 5 eo Ae Ska ri 28-30 ta, Lowndes Oo., Gaderitg 4 eae es oes Cie hes Oct. I Gide tip Coc Georgii io. loco ieee) irs a2 2-4 Eknigiag, Comee Co., Georgia. os. os ig oe ee es 4 yetoss; Ware. Coz, Georgia cncoce focus ik eee ee =~ 5-6 Jacksonville, Duval Co., Florida fd eet Da Rate wie: hag Sree a 47-8 Brunswick, Glynn Co. Georgia. 2.4... <0 os bocnc san zy 10 Charleston, Charleston Co., ae WAROURE UE SUS os eG kee - II-13 nks Corner, Berkeley Co., South Carolina.............. sh I4 Wilmington, New Hanover Co., North Carolina........... ifs I5—16 Rocky Mount, Nash Co., North Rs ie rac etiniatbie we weataie > 17 Weldon, Halifax Co., North Carolina..................... “s 18 Altogether about 300 numbers of Agalinanae were collected, and a field-study of each species made. These collections and notes form the basis of this report. In connection with this material I have also reviewed the specimens from the Coastal Plain in the herbaria of the following institutions: United States National Museum, Missouri Botanical Garden, Field Columbian Museum, Biltmore Herbarium, New York Botanical Garden, Academy of Natural Sciences of Phila- delphia, University of Pennsylvania, Charleston Museum, Tulane University, Florida Agricultural College, also in the private herbaria of Dr. E. L. Greene, C. C. Deam and H. H. Bartlett. To the custodians and owners of all of these I am greatly in- debted. PENNELL: STUDIES IN THE AGALINANAE 403 Also I am indebted to Dr. N. E. Brown of Kew Gardens, England, for consulting Bentham’s types in their collections, portions of three of which were kindly sent me; to Dr. A. B. Rendle of the British Museum for examining Walter's types; and to Dr. C. H. Ostenfeld of Copenhagen, Denmark, for comparing the type of Gerardia tenuifolia Vahl. Of the thirty-four species and subspecies in the following pages, all but one have been seen and studied in the field. This exception, Agalinis oligophylla, is a Louisiana species not in flower in August when I was there. All new species here proposed have been seen growing, and compared with allies in their native environment. It has been a matter of no small satisfaction that herbarium material since reviewed has fallen so readily into the species deemed valid in the field. A word may be said concerning field variation in this group. The one species of Macranthera and our two of Afzelia, as well as all our species of Agalinis seem relatively uniform and constant. Breaks between species in the last are often slight but field-study shows them to be true—I have observed little in this genus to suggest hybridism or pronounced subspecific variation. On the other hand, in Aureolaria, especially in the subgenus Panctenis, conditions seem much more complicated, and such a species as Aureolaria pectinata can be viewed only as composed of a number of strains. In the following treatment the endeavor is made to include quite fully diagnostic characters in the key, while, to save space, only in the case of new species are specific descriptions given. Full synonomy is included so far as Coastal Plain species are’ concerned. Type localities are quoted from the original descrip- tions, and where types have been examined the fact is noted. No types have been found for Rafinesque’s species. The type of Gerardia tenuifolia leptophylla Benth.* has not been as yet identi- The flowering and fruiting seasons are from specimens seen, — but are necessarily incomplete. North and south the flowering season is about the same for a given species, possibly a little later south. All definiteness in season seems to be lost in southern * Hook. Comp. Bot. Mag. 1: 174- 1835. “ Jacksonville, Louisiana.” G. tenui- folia filiformis Benth. in DC. Prodr. 10: 518. 1846, is the same plant. 404 PENNELL: STUDIES IN THE AGALINANAE Florida. Distribution notes are made as definite as practicable.* All localities from which specimens have been seen are listed, and in the case of specimens of my own collecting numbers are given. Of course many specimens have been seen with data too vague for such classification. In the fuller revision of this group upon which the writer is working it is intended more fully to index specimens seen. In prosecuting this study I have been most deeply indebted to the following three gentlemen: to Mr. Roberts Le Boutillier, of Wayne, Pa., whose generosity enabled me to accomplish the field-work required, to Dr. Roland M. Harper whose knowledge of the flora of the Southeast, most generously imparted, has made my search far more successful than it otherwise could have been, and to Dr. John M. Macfarlane, under whose direction and con- stant support this study has been made. Key to the coastal plain genera Corolla See orange, its base thickened, fleshy, Sel ercoam shriveling and blackening before falling. Filaments equal, meatgnce Ay pubescent with beaded hairs. aia sacs paid parallel, opening their entire length. Capsule ovoid, inate, densely and closely pubescent. Seeds win ged. 1. Macranthera. Coil: Suis inflated throat or spreading lobes, yellow or purple, not fleshy nor semi-persistent. Filaments not long-exserted, pubescence not beaded. Corolla nearly rotate, tube short, lobes longer than tube, yellow. Filaments nearly equal, about the length of the corolla-tube; anther-sacs closely parallel, opening by short apical slits. Capsule ovoid, acute. Seeds wingless or winged. 2. Afzelia. Corolla with inflated throat, tubular-campanulate, lobes much shorter than tube. Filaments didynamous, included; anther-sacs opening their entire length Corolla yellow. Anther-sacs parallel, awned at base. Capsule acute to acuminate. Seeds wingless or winged. 3. Aureolaria. * In summarizing local distribution I have largely adopted the floristic areas out= outlined in the following publications: W. Stone—Plants of Southern New jersey=—— ep. New Jersey State Mus. [1910]: 1912; F. Shreve et al.—Plant Life of Maryland—Maryland Weather Service 3: r910; R. M pag of the coastal plain—Bull. Torrey Club 37: 405-428, 1910; R. M. Har tamaha Gri Region of Georgia—Ann. New York Acad. Sci. 17: 1. a R. M. Harper— Preliminary Report on Peat—Florida Geol. Sury., Ann. Rep. 3: 199-375. I9II; C. Mohr—Plant te of irairan Teer: trib. U. S. Nat. Herb. 6. 1901; R. M- Harper—Geograp p Geol. Surv. Alabama Monograph 8. 1913- PENNELL: STUDIES IN THE AGALINANAE 405 Corolla pink or purple. Anther-sacs more or less di- vergent, obtuse to mucronate-awned at base. Capsule rounded at apex. Seeds wingless. 4. Agalinis. 1. MACRANTHERA “Torr.”’; Benth. in Hook. Comp. Bot. Mag. 1: 174. 1835-6 One species: I. MACRANTHERA FLAMMEA (Bartram) Pennell, Bull. Torrey. Club 40: 124. 1913 Gerardia flammea Bartram, Trav. 412. 1791. “Stony gravelly heights’”’ along Tensaw River* near ‘‘ Taensa,’’ Alabama. No type specimen known to exist. Identified by Mohr, Plant Life of Alabama, 15. 1901. Conradia fuschioides Nutt. in Jour. Acad. Nat. Sci. Phila. 7: 88. 1834. No locality given. Type, without data, seen in Herb. Acad. Nat. Sci. Phila. Macranthera fuchsioides (Nutt.) Benth. in Hook. Comp. Bot. Mag. 1: 174. 1835-6. Macranthera Lecontei Torr. in Ann. Lyc. Nat. Hist. New York 4: 80. pl. 4. 1837. ‘‘In dry pine woods on the Alatamaha, in Liberty County, Georgia.”” Type; without data, seen in Herb. Columbia University. Russelia flammea (Bartram) Raf. New Fl. Am. 2: 71. 1837. Flamaria coccinea Raf. New Fl. Am. 2: 71. 1837. Additional name for Russelia flammea (Bartram) Raf. Toxopus gymnanthes Raf. New Fl. Am. 2: 72. 1837. tional name for Macranthera Leconte Torr. Toxopus calycinus Raf. New Fl. Am. 2: 72. 1837. Additional name for Macranthera fuchsioides (Nutt.) Benth. Tomilix bracteata Raf. New Fl. Am. 2: 72. 1837. Additional name for Macranthera fuchsioides (Nutt.) Benth. Macranthera fuchsioides Lecontei (Torr.) Chapm. Fl. So. ug. S. 297. 1860. Conradia Lecontei (Torr.) Kuntze, Rev. Gen. 1: 459. 1891. Though somewhat variable, when in flower the calyx-lobes are mostly entire, linear, relatively short as compared with corolla-tube [= M. Lecontei], later the corolla shrivels and Addi- * Eastern arm of Mobile River. 406 PENNELL: STUDIES IN THE AGALINANAE contracts, but persists, while the calyx-lobes continue to grow, becoming quite lobed and leaf-like [= M. fuchsioides]. Flowers, August to October. Fruit, September to October. DisTRIBUTION: Borders of wet sandy thickets, in lower coastal pine belt and coast district, southern Georgia and northern Florida to southern Mississippi. Frequent in pine hill region of northwestern Florida, and in southern Alabama. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: Georgia: Probably Liberty Co., LeConte; Thomasville. Florida: Quincy; Beverly (4681); Argyle; Crestview; Milligan (4595); Milton (4564); Bluff Springs. Alabama: McRae (4641); Bay Minette (4553); Mobile; Whistler (4534); Theodore (4406, 4459, 4462). Mississippi: Ocean Springs; Biloxi. 2. AFZELIA J. F. Gmel. curante L. Syst. Nat. ed. 13. 927. 1791 Stem closely pubescent, viscid; leaves pinnatisected, ati lanceolate or broader; calyx-lobes lanceolate; corolla dee pdpied pubescent without, its lobes ovate, 3-3.5 mm. ce nthers lanose with yellow hairs on dorsal side of connective; ek ovate, oa! mm. long, densely shott tomentose with brown hairs; ds winged. Plant low, 2-6 dm. tall, widely branched. 1. A. pectinata. Stem ates pubescent, scarcely glandular; leaves pinnatifid, : segments filiform; calyx-lobes linear; corolla pale yellow, gla- brous without, its lobes lanceolate, 1.5-2 mm. aaa anthers glabrous; capsule inversely pyriform, 4-4.5 mm. long, glabrous; seeds wingless. Plant tall, 5-10 dm., virgately branched. 2. A. cassioides. I. AFZELIA PECTINATA (Pursh) Kuntze, Rev. Gen. 1: 457. 1891 Seymeria pectinata Pursh, Fl. Am. Sept. 2: 737. 1814. “In South Carolina. Catesby—yv. s. in Herb. Sherard.”’ Seymeria Jacksoni Ell. Sketch 2: 123. 1824. ‘Sent to me from Louisville, Ga., by Mr. Jackson.” Type seen in the Elliott Herbarium at the Charleston Museum. Seymeria heterophylla Raf. New Fl. Am.2: 68. 1837. ‘‘Alabama and Georgia, my specimen from Leconte.” Flowers, August to September. Fruit, mid-September to October. DistTRIBUTION: Dry sandy pineland in the coastal plain from PENNELL: STUDIES IN THE AGALINANAE 407 South Carolina to Mississippi, south in the Florida peninsula to Miami. Frequent in southern Georgia and southern Alabama and in Florida. Nearly restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: South Carolina: Aiken. Georgia: Augusta; Thomson; Louisville; Brunswick (4845); Way- cross (4780); Thomasville (4732) ; Leslie (4760) ; Columbus. Florida: Jacksonville; South Jacksonville; San Pablo (4802); St. Augustine; Tocoi; Eustis; Palm Springs; Clarcona; Georgiana; Miami; Marco; Tampa; St. Marks (4705, 4715); Quincy; River Junction; Liberty Co.; Fort Gadsden (4686); Apalachi- cola (4674); Chipley (4645); Ponce de Leon (4656); Milligan (4585); Milton (4568); Santa Rosa Island. Alabama: Auburn; Abbeville; Ozark; Florala (4630); Greenville; Wilcox Co.; Mobile Co. Mississippi: Waynesboro. 2. AFZELIA CASSIOIDEs (Walt.) J. F. Gmel. curante L. Syst. Nat. ed. 13. 927. 1791 Anonymos cassioides Walt. Fl. Carol. 171. 1788. No type locality given, presumably from Berkeley Co., South Carolina. Gerardia cassioides (Walt.) Pers. Syn. 2: 154. 1807. Seymeria tenuifolia Pursh, Fl. Am. Sept. 2: 737- 1814. New name for Gerardia cassioides (Walt.) Pers. Flowers, September to: mid-October. Fruit, October. DistriButTion: Moist to dry pineland, mostly sandy, in the coastal plain from North Carolina to Florida and Louisiana. Frequent in the Wilmington pine barrens, occurs also near Fayette- ville, North Carolina, and occasional or frequent southward. Most abundant in flat pine woods of southern Georgia, northern Florida, and near the Gulf coast to Louisiana. In the Florida peninsula reaching Bradentown on the west coast. Mostly in the coastal plain, casually inland in northern Georgia, Alabama, and south- _ astern Tennessee. PLANTS AND SPECIMENS EXAMINED: : North Carolina: Wilmington (4900, 4919); Fayetteville. South Carolina: Columbia; Santee Canal; St. Johns; Cooper 408 PENNELL: STUDIES IN THE AGALINANAE River; Monks Corner (4878); Otranto (4872); Summerville; Charleston (4866); Beaufort. Georgia: Thalmann (4809) ; Coffee Co.; Naylor (4743); Moultrie; Thomasville (4725); Leslie. Florida: Jacksonville; St. Augustine; Bradentown; Tampa; Lake City; Monticello (4719); St. Marks (4713); Quincy; River Junction; Fort Gadsden (4691); Apalachicola (4678); Chipley (4649); Ponce de Leon (4653); Milligan (4588). Alabama: Auburn; Abbeville; McRae (4639); Bay Minette (4552); Mobile Co. Mississippi: Meridian; Fontainebleau; Ocean Springs; Biloxi; Nicholson. Louisiana: Covington (4217); Hammond. 3. AUREOLARIA Raf. New Fl. Am. 2: 58. 1837 Corolla glabrous without; calyx-lobes entire; seeds winged. Plants perennial, not glandular. Aureolaria (sensu strictu). Capsule glabrous. Flowers evidently ee St glabrous, more or less glaucous. ath. Stem quite glaucous. Bracts ofa lanceolate type, some at least dentate like the leaves. Capsule 15-20 mm. lon Stem pres glaucous. Bracts of Leal . A. virginica. la aces about 12-15 mm. long. 2. A. reticulata. th. ee of aspatulate type,entire. 3. A. dispersa. bescent. ant pu throughout. Flowers very short-pediceled. 4. A. villosa. te; seeds wingless. Plants annual, Panctenis (Raf.) subgenus nov.* glandular. Leaves and stem minutely pubescent or mi- nutely glandular. Fedioes mostly over bescent. Stem-leaves mostly 3 em. long. * Gerardia sect. Pedicularioides Benth. in Hook. Comp. Bot. Mag. £: 205+ 183 rt if sectional names have full standing, has priority over Panctenis Raf. New Fi. aa 0. 28375 PENNELL: STUDIES IN THE AGALINANAE 409 Pedicels mostly shorter than or equaling the bracts. 5. A. pedicularia. Stem-leaves mostly less than 2 cm. long. Pedicels longer than the bracts. 6. A. pedicularia caesariensis. Leaves and stem minutely glandular- pubescent. Leav PROS si Ay ose of the stem mos ge 2-3 cm. long. 7. A. pedicularia carolinensis. Leaves and stem densely dgahtendbaaian edicels mostly less than 10-12 mm. long, shorter than the calyx. Leaves pinnately lobed, lobes sharply pectinate. mispheric. Capsul Stem-leaves all spreading. Flowering pedicel 5-10 mm. long. Flowers about 5 mm. long. Plant widely branched, uppermost leaves smaller, but not ex- cessively reduced. 8. A. pectinata. eam teay en, at least the upper, appresse es Flowers about 40-45 m. g. Plant virgately branched, uppermost leaves much reduced 9. A. pectinata floridana. 1. Aureolaria virginica (L.) Pennell, comb. nov. Rhinanthus virginicus L. Spec. Plant. 603. 1753. “Habitat in Virginia.” Specimen in Gronovius’ herbarium identified by Pursh. As specimen in Linnaean Herbarium bears hand- writing of Linnaeus the younger, and was probably a later addition, I presume Gronovius’ plant to be the type. For dis- cussion see Kuntze, Rev. Gen. 1: 460. 1891. G rardia flava L. Spec. Plant. 610. 1753. “Habitat in Virginia, Canada.” Specimen in Linnaean Herbarium identified by Bentham in Hook. Comp. Bot. Mag. 1: 198. 1835-6. Anonymos flava (L.) Walt. Fl. Carol. 170. 1788. As to synonymy, not description, the latter probably applying to Aureolaria villosa Raf. Gerardia lauca Eddy in Med. Repos. N. Y. IInd. Hexade, 5: 126. 1807. Plandome, Long Island. C. W. Eddy. Gerardia quercifolia Pursh. Fl. Am. Sept. 2: 423. 1814. “On the banks of rivers in rich shady elie os caer sam to Carolina.” _ : 410 PENNELL: STUDIES IN THE AGALINANAE Aureolaria glauca (Eddy) Raf. New Fl. Am. 2: 60. 1837 Dasystoma quercifolia (Pursh.) Benth. in DC. Prodr. 10: 520. 1846. Dasystoma flava (L.) Wood, Class Book. 529. 1861. As to synonymy, not description, the latter applying to Aurevlaria villosa Raf. Gerardia virginica (L.) Britton, Prelim. Catal. N. J. Pl. 40. 1888. Dasystoma virginica (L.) Britton in Mem. Torrey Club 5: 295, 1894 Flowers, in New Jersey, September. DIsTRIBUTION: Frequent on rich slopes, often rocky, in wood- land inland, very rare in the Coastal Plain. On wooded slopes at a few points in southern New Jersey. SPECIMENS EXAMINED: New Jersey: New Egypt; Nesco; Fairton. 2. AUREOLARIA RETICULATA Raf. New Fl. Am. 2: 59. 1837 Aureolaria reticulata Raf. ‘Florida and Alabama.’ No type _ known to exist. Dasystoma bignoniiflora Small in Bull. N. Y. Bot. Gard. 1: 285. 1899. ‘Collected by Dr. Burrows, at Tampa Bay, Florida, in 1834.’’ Type seen in Herb. Columbia University. Flowers, late-August to mid-October. Fruit, September to October. DISTRIBUTION: Sandy ravines and moist woodland, both in calcareous and siliceous districts, in the coastal plain from North Carolina to central and western Florida. Apparently most frequent in calcareous region of northern and western Florida. Restricted to the coastal plain; possibly better considered a sub- species of A. virginica replacing that species in the southern coastal plain. PLANTS AND SPECIMENS EXAMINED: North Carolina: Fayetteville. South Carolina: Santee Canal; St. Johns; Monks Corner (4875) > Eding Island. Georgia: Thomasville (4723); Leslie (4765). Florida: Tampa Bay; Monticello (4720) ; Tallahassee (4690, 4698); , River Junction; Marianna; Milton (4565, sae PENNELL: STUDIES IN THE AGALINANAE 411 3. Aureolaria dispersa (Small) Pennell, comb. nov. Dasystoma quercifolia intermedia Benth. in DC. Prodr. 10: 520. 1846. No type locality given, nor type specimen known to exist. Description would indicate this species. Dasystoma dispersa Small, Bull. Torrey Club 28: 452. Igot. “Louisiana: Feliciana, Carpenter; type in the herbarium of Columbia University.’”’ Type seen in Herb. Columbia Uni- versity. Gerardia dispersa (Small) K. Schum. in Just’s Bot. Jahresber. 29: 580. 1903. Flowers, August to September. Fruit not seen. DIsTRIBUTION: Sandy thickets and oakland, frequent in coastal pine belt near the Gulf, southern Alabama to eastern Louisiana, reaching Wilkinson Co., Mississippi. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: : Alabama: Baldwin Co.; Crichton (4521); Hollander’s Island (4504). Mississippi: Ocean Springs; Biloxi (4384); Long Beach; Wilkinson Co. Louisiana: Abita Springs (4117, 4245); Covington; Feliciana. 4. AUREOLARIA VILLosA (Muhl.) Raf. New Fl. Am. 2: 59. 1837 Aureolaria villosa (Muhl.) Raf. No type locality given, nor type specimen known to exist. Gerardia villosa Muhl. Catal. 58, 1813, and Gerardia heterophylla Muhl. 1. c. are treated as nomina subnuda. Dasystoma pubescens Benth. in DC. Prodr. 10: 520. 1846. “In Americae sept. civitatibus orientalibus frequens.” This species has been commonly identified as Gerardia flava, but appears not to be G. flava L. Flowers, late-May to mid-August. Fruit, late-July to October. DistriBuTION: Widely distributed in the eastern Unit States, more common above the fall line. Frequent or occasional in sandy soil in the coastal plain from New Jersey to Louisiana. Not seen from the Florida peninsula. In New Jersey frequent in middle and Cape May districts, less so in the pine barrens. 412 PENNELL: STUDIES IN THE AGALINANAE PLANTS AND SPECIMENS EXAMINED: New Jersey: Spotswood; Farmingdale; Hornerstown; Medford; Locust Grove; Westville; Clarksboro; Swedesboro; Atco (3540); Hammonton; Cape May Court House (3986) ; Bennett; Cold Spring; Cape May. Maryland: Chestertown. Virginia: Naucks (2471); Cape Henry; Franklin. North Carolina: Wilmington. South Carolina: Santee Canal; St. Johns; Bluffton; Aiken; Graniteville. Georgia: Macon. Florida: Jacksonville; Chattahoochee River. Alabama: Ozark Evergreen. Louisiana: Pearl River. 5. AUREOLARIA PEDICULARIA (L.) Raf. New Fl. Am. 2: 61. 1837 Gerardia pedicularia L.Sp. Pl.611. 1753. ‘Habitat in Virginia, Canada.” Anonymos pedicularia (L.) Walt. Fl. Carol. 170. 1788. As to synonymy, probably not description. Panctenis pedicularia (L.) Raf. New Fl. Am. 2: 61. 1837. Dasystoma pedicularia (L.) Benth. in DC. Prodr. to: 521. 1846. Variable, the following subspecies local, and more or less sharply defined. Flowers, mid-August to mid- September. Fruit, October. persisting through the winter. DIsTRIBUTION: Widely distributed and frequent in the northern states above the fall line. Rare in the coastal plain where mostly replaced by the following subspecies. Material seen nearly all from New Jersey, where it is rare or absent in middle district, somewhat more frequent in the pine barrens and evidently approaching the subspecies caesariensis. (Transitional specimens indicated by asterisk.) PLANTS AND SPECIMENS EXAMINED: New Jersey: Middlesex Co.; Brindletown*; Atsion*; Hammon- ton*. Pennsylvania: Tinicum (3589). PENNELL: STUDIES IN THE AGALINANAE 413 6. Aureolaria pedicularia caesariensis Pennell, subsp. nov. Annual. Stem ro dm. tall, widely branching, minutely and closely pubescent, very sparingly glandular below. Leaves sessile, ovate-lanceolate, less than 2 cm. long, pinnatifid, segments irregularly crenate-dentate, equaling or longer than the width of the central portion of the lamina, minutely and closely pubescent. Pedicels slender, glandular, 15 mm. long, longer than the calyx, longer than, mostly twice exceeding, the bracts. Calyx-tube glandular with short-stalked glands, equaled by the dentate calyx-lobes. Corolla 35 mm. long, yellow. Capsule 10-12 mm. long, elliptic-ovoid, minutely glandular, surpassing the calyx-lobes. Type, Atco, Camden Co., New Jersey, Sept. 7, 1911, F. W. Pennell 3545 in Herb. University of Pennsylvania. Flowers, mid-August to mid-September. Fruit, October, per- sisting through the winter. DistriBuTION: Dry sandy pine and oak woods, frequent in the pine barrens of New Jersey, where it mostly replaces the species. PLANTS AND SPECIMENS EXAMINED: New Jersey: East Plains; Bamber; Woodmansie; Taunton; Atco (3545, 3627); Middletown, Cape May Co. 7. Aureolaria pedicularia carolinensis Pennell, subsp. nov. , Annual. Stem 10 dm. tall, widely branching, minutely glandular-pubescent. Leaves sessile, ovate-lanceolate, 2-3 cm. long, pinnatifid-lobed, segments somewhat crenate, shorter than the width of the central portion of the lamina, minutely glandular- pubescent. Pedicels slender, glandular, 10-20 mm. long, longer than the calyx. Calyx-tube glandular with short-stalked glands, mostly exceeded by the incised-dentate calyx-lobes. Corolla 35 mm. long, yellow. Capsule 12 mm. long, elliptic-ovoid, minutely glandular, equaling or barely surpassing the calyx-lobes. Type, savannahs near Mill Pond, Wilmington, North Carolina, June 23, 1909, J. M. Macfarlane in Herb. University of Pennsyl- Vania. Flowers, June to September. Fruit, October. Distripution: Dry sandy pine and oak woods, pine barrens of southeastern North Carolina, where it probably replaces the species. PLANTS AND SPECIMEN EXAMINED: North Carolina: Wilmington (4925) (J. M. Macfarlane in 1909; in 1911); ‘Southeastern North Carclina,” W. W. Ashe. 414 PENNELL: STUDIES IN THE AGALINANAE 8. Aureolaria pectinata (Nutt.) Pennell, comb. nov. Gerardia pedicularia pectinata Nutt. Gen. Plant. N. Am. 2: 48. 187-65: In the sandy pine forests of Carolina and ‘Georgia.’”’ No type in Herb. Acad. Nat. Sci. Philadelphia. Gerardia pectinata (Nutt.) Benth. in Hook. Comp. Bot. Mag. 1: 206. 1835-6. Panctenis pectinata (Nutt.) Raf. New Fl. Am. 2: 61. 1837. Dasystoma pectinata (Nutt.) Benth. in DC. Prodr. 10: 52. 1846. Very variable, as here understood, including a number of strains. . Flowers, July to September. Fruit, September to October. DISTRIBUTION: Dry sandy pineland, especially hilly, in the coastal plain, South Carolina, through southern Georgia, pine hills of northwest Florida, to Alabama and Mississippi, locally frequent extending above the fall line. In some of its forms occur- ring throughout the Gulf and Lower Mississippi Valley states. PLANTS AND SPECIMENS EXAMINED: South Carolina: Eutawville; Santee Canal; Monks Corner; Cooper River; Beaufort. Georgia: Augusta; Thomson; Americus. Florida: De Funiak Springs. Alabama: Florala (4625); Flomaton; Prattville; Tensaw; Deer Park; Mobile; Spring Hill (4532). Mississippi: Meridian; Jackson. 9. Aureolaria pectinata floridana Pennell, subsp. nov. Annual. Stem about 10 dm, tall, much branched, glandular- villose. Branches virgate, ascending. Leaves sessile, ovate, those of the stem less than 2 cm. long, pinnatifid, segments pec- tinately toothed, glandular-villose; at least the upper leaves ascending or appressed to the stem or branches, the uppermost much reduced. Pedicels stout, 5 mm. long or less, shorter than the calyx, glandular-villose. Calyx-tube glandular-villose, much exceeded by the lanceolate, pectinate teeth. Corolla 40-45 (-50) mm. long, yellow, no purple markings within tube (in type specimens). Capsule 12-14 mm. long, broadly ovoid, glandular-pubescent, shorter than the calyx. Type, Fort Gadsden, Franklin Co., Florida, Sept. 20, 1912, F. W. Pennell 4683, in Herb. University of Pennsylvania. PENNELL: STUDIES IN THE AGALINANAE 415 This may be Gerardia pedicularia pectinata Nutt., as the men- tion of short pedicels and very large flowers would suggest, but as floridana does not occur in South Carolina, though probably in the flat pine woods of southeastern Georgia, I retain the name for the prevalent species of the district cited. I have seen no old collections of floridana from Georgia. Flowers, May to mid-October. Fruit, June to October: DIsTRIBUTION: Dry sandy pineland. Flat pine woods of Florida and southern Georgia, south to Polk Co., Florida; re- placing the species. PLANTS AND SPECIMENS EXAMINED: Georgia: Thomasville (4724). Florida: Jacksonville; Pablo; St. Augustine; Eustis; Orange City; Lake Brantley; Polk Co.; Tampa; Marion Co.; Tallahassee; Fort Gadsden (4683); Apalachicola. 4. AGALINIS Raf. New FI. Am. 3: G1, 3847 Perennial, from a running rootstock, glabrous throughout. Pedi- cels erect, 5-20 mm. long. Corolla slightly fleshy, pink, with darker spots, but no yellow lines within throat, pubescent with- out, pubescent within at base of upper lobes, 35-40 mm. long. Capsule globose, 5-6 mm. long. Seeds dark brown. [Lini- foliae.] 1. A. linifolia. Annuals, fibrous rooted, in most at least the upper surface of the leaf scabrous. Pedicels ascending or spreading. Corolla membranous, purple or pink, mostly with darker spots and two yellow lines within throat. Capsule poe or ovoid. Corolla with lobes all spreading, pubescent within at base of upper lobes, more or less pubescent without. Seedsdark brown. Plantsdarkeningindrying. Calyx- Leaves uniform, linear to cageimantane: Inflorescence of normal racemes; cs pedicels less than 1 with Anther-sacs o Plant fleshy, ee below, with elongated racemes a cels reaching ro mm. io 3 2. A. maritima. 416 PENNELL: STUDIES IN THE AGALINANAE Leaves and calyx-lobes acute to acumi- nate. Anther-sacs mucronate to t ba minutely awn not fleshy, more branched. Pedicels less m. long. Cisolie purple or yellow lines and darker Plants dull-green or present. Plants uniformly than 5 pink-purple, 2 spots purplish, much darkening in d rying. Stem smooth or minutely scabrel- 1 ilary us. fascicles slightly or not developed, if present shorter than eaves. Corolla 20-35 mm. long, rose-purple. Corolla on mm. long, on vide nt pedicels 2-5 mm. long. Stem egeringty scabrellous. to te. triangular-subu- Corolla 20-25 mm. long. vi es narrowly ear to almost sea » Calyx- lobes triangular- subulate to subu- late. Stem essen- ry. Stem-leaves nar- rowly 1.5-3 ng, mostly longer e inter- nodes, mostly curling in drying. Branches mostly simple, ascend- ing, virgate. fascicles relatively well 3. A. purpurea. PENNELL: STUDIES IN THE AGALINANAE 417 developed. Pedi- cels 2-5 mm. long. 4. A. virgata. Stem-leaves I-2.5 veloped. g in drying. 5. A. pinetorum. almost fili- form, I- cm. long, curling drying. 6. A. delicatula. Corolla 15-18 mm. long, pink-purple. Flowers nearly sessile, on pedicels less than 2 mm. long. 7. A. Harperi. Stem scabrous. Axillary fascicles aaa evclned mostly ‘ ualing lea Corolla RE OEE no sed lines 8. A. fasciculata. early so. Axillary fascicles abundantly developed. 9. A. georgiana. 418 PENNELL: STUDIES IN THE AGALINANAE growth of stem apex, frequently appear- ing terminal. Pedicels over 5 mm. lon Stem pay oan Anther-sacs_ evidently ed mm. long. Corolla 25-30 mm. long. 10, A. pulchella. Stem glabrous, or essentially so. Anther- acute to minutely mucronate- awned at base, pu nt, glabrous over much of dorsal surface. Seed- coat with dark-brown thas areas between these elongated, scar cely ee and scarcely or not reticu- lated. Stem-leaves alt to filiform, semi-fleshy, long. Axillary fascicles snceaien developed. Pedicels 10-35 mm. long. Corolla 25-30 mm. long. 11. A. filifolia. Stem-leaves opposite. Axillary fas- : cicles scarcely developed or none. Stem-leaves narrowly linear to filifo: ridges. Plants densely and repeatedly branched, usually PENNELL: STUDIES IN THE AGALINANAE 419 Pedicels less than 10 mm. long, mostly shorter than he bracts 5-3 Leaves filiform-seta- ceous. Flowers con- spicuously “ter- minal.” 13. A. setacea. Pedicels, at least in fruit, 25-50 mm. long, slender, four to five times exceeding the bracts. Co- roll branched, the lower ranches widely ascend- ing. Leaves narrowly filiform. 14. A. laxa. Stem-leaves filiform-subulate, 1 cm. long or less, much shorter than internodes. -coat with broader areas. Pedicels 5-15 mm. long. Calyx-lobes acute, 0.5 mm. long. rolla 15-20 mm. long. Stem terete, grooved, not angular 15. A. oligophylia. Leaves dimorphic, basal oval- peas eaeiatiiae: cauline, minute, scale-like, appressed. Pedice less than 5 mm. long, many tee appearing to terminate minute axillary branches. Calyx- lobes minute, subulate. Corolla 15-20 mm. long, pink-purple. Stem mostly 4-angled, often pubes- cent at base. 16. A. aphylla. Seeds yellows brown. Plants light green, not darken- g tying. _ pink. Flowers on pedi oe longer than bracts. Calyx-tube reticu- late-venulose, lobes Sica callose. [Erectae.] Stem-leaves 2-2.5 cm. long, filiform-linear, acute. olla 15 mm. long, lobes more or less emar- ginate, 2 yellow lines and purple spots evident. 17. A. decemloba. Stem-leaves 1-1.5 cm. long. Corolla 15-20 mm. long, lobes more or less emarginate, 2 yellow lines and purple spots well developed. Pedicels slender. Leaves nearly filiform, acutish or acute. Plant relatively lax. 18. A. tenella. 420 PENNELL: STUDIES IN THE AGALINANAE Corolla — mm. long, lobes “Sg to tru 2 yellow lines and pur Sonate ee Seieloned or EN ee stouter. Leaves linear, widening ane obtusish to obtuse. Plant strict, stiff. 19. A. erecta. Corolla bilabiate, two upper lobes ascending over stamens and style, glabrous within at base of upper lobes. Seeds dark brown. [Tenuifoliae.] Corolla pubescent without, 2 upper lobes two-thirds length of lower 3, minutely ciliate, concave-arched. Pedicels, if exceeding the bracts, less than twice their length. Corolla 10-18 mm. long, purple. Leaves linear, ee of the stem aes broadly so 20. A. tenuifolia- orolla g without, 2 upper lobes less tans one-half length of lower 3, conspicuously ciliate, flattened. Pedicels at least three times the length of the racts. Leaves filiform, those of the stem 1.5—2 cm. long. Pedicels 15-30 mm. long. Corolla rose-pink, t5-18 mm. long. Plant reaching 8 dm oe stich branched. Leaves minute, scale-like, those of the stem less . long. Pedicels 5-10 mm. long. meget ee aes Io0-13 mm. long. " Plant less than 5 dm. tall, sparingly very laxly branched. 22. A. filicaulis. , 21. A. divaricate- AGALINIS LINIFOLIA (Nutt.) Britton in Britt. & Br., Ill. Fl. ed. 2. 3: 209. 1913 Gerardia linifolia Nutt. Gen. Pl. N. Am. 2:47. 1818. ‘‘HAB. From Wilmington, North Carolina to Florida.’ Type, labeled “Carolina,” seen in Herb. Acad. Nat. Sci. Philadelphia. Agalinis perennis Raf. New Fl. Am. 2: 63. 1837.. “My speci- men is from Florida.” Flowers, mid-August to October. Fruit, September to DISTRIBUTION: Wet pineland, and especially margins of ponds, in the coastal plain. Ellendale, Delaware; from Wilming- ton, North Carolina to southern Florida and near the Gulf coast to Louisiana. Most abundant in flat pine woods of southern Georgia and Florida, less frequent in the west Florida pine hills and in the Altamaha grit region of Georgia, occasional north of the Savannah River or west of Florida. Restricted to the coastal plain. PENNELL: STUDIES IN THE AGALINANAE 421 PLANTS AND SPECIMENS EXAMINED: Delaware: Ellendale. North Carolina: Wilmington; Brunswick Co. South Carolina: Hartsville; Santee Canal; Eutawville. Georgia: Everett; Brunswick (4823); Waycross (4790); Naylor (4745); Thomasville (4729); Sumter Co.; Columbus. Florida: Tisomia (4813); Baldwin; Jacksonville (4794); St. Nicholas; San Pablo (4807); Cutler; Homestead; Camp Long- view; Fort Myers; St. Petersburg; St. Marks (4714); Quincy; Fort Gadsden (4690); Apalachicola; Chipley (4648, 4666); Ponce de Leon (4654); Paxton (4600). Alabama: Elmore. Mississippi: Biloxi. Louisiana: Hammond. 2. AGALINIS MARITIMA Raf. in Med. Repos. New York. IInd. hex. 5: 361. 1808 (as Gerardia); New Fl. Am. 2: 62. 1837 Gerardia maritima Raf. ‘‘Found in the islands of Egg-Harbour, in New Jersey.’’ No type known to exist. Gerardia purpurea crassifolia Pursh Fl. Am. Sept. 2: 422. 1814. “In salt marshes, near New York.”’ Gerardia maritima grandiflora Benth. in Hook. Comp. Bot. Mag. I: 208. 1835-6. ‘Texas, Drummond, Ist Coll.” Gerardia spiciflora Engelm. Bost. Jour. Nat. Hist. 5: 227. 1845. New name for Gerardia maritima grandiflora Benth. Gerardia maritima major Chapm. Fl. So. U. S. 300. 1860. “Brackish marshes, Apalachicola, Florida.” There are several collections of Chapman’s of this species, but none, indicated as type, has been seen. Flowers, April (on the Gulf coast) to mid-September. Fruit, August to September. : DISTRIBUTION: Salt marshes, along the Atlantic and Gulf coasts from New Jersey to Louisiana, locally frequent. Extending to Maine northward, in the south to Texas, Yucatan, Cuba and the Bahamas. Along the Gulf coast and Atlantic coast in Florida a much branched plant, reaching 6 dm. tall, leaves and calyx- lobes more liable to be acutish, pedicels mostly exceeding the bracts, corolla reaching 20 mm. long, and anther-sacs lanose- 422 PENNELL: STUDIES IN THE AGALINANAE pubescent, northward becoming progressively smaller and simpler, till a plant of but 0.5—1.0 dm. tall in Maine, leaves and calyx- lobes constantly obtuse, pedicels shorter than the bracts, corolla. scarcely 15 mm. long, and anther-sacs nearly glabrous. PLANTS AND SPECIMENS EXAMINED: New Jersey: Keasbey; Long Beach; Forked River; Barnegat Pier; Beach Haven; Atlantic City; Ventnor; Mays Landing; Ocean City; Palermo; Sea Isle; Peermont; Cape May Court House (2604); Holly Beach; Five-mile Beach; Cold Spring (2157); Cape May. Delaware: , Bernhardt. Maryland: Ocean City. Virginia: Parksley; Walnut Point. North Carolina: Ocracoke Island; Beaufort. Florida: Titusville; Eau Gallie; Marco; Sanibel; Pine Island; Tampa; Long Key, Pinellas Co.; Hernando Co.; St. Marks (4702); Apalachicola. Alabama: Mobile Co. Mississippi: Ship Island. Louisiana: Breton Island. 3. AGALINIS PURPUREA (L.) Pennell, Bull. Torrey Club 40: 126. 1913 Gerardia purpurea L. Sp. Pl. 610. 1753. ‘Habitat in Virginia, Canada.” | Anonymos purpurea (L.) Walt. Fl. Carol. 170. 1788. Gerardia purpurea grandiflora Benth. in Hook. Comp. Bot. Mag- I: 208. 1835-6. ‘‘ New Jersey.” Agalinis palustris Raf. New Fl. Am. 2: 62. 1837. ‘‘Near marshes. . . . From New England to Carolina.”’ Agalinis longifolia Raf. New Fl. Am. 2: 62. 1836. ‘Near streams New Jersey to Virginia.” (?) Agalinis corymbosa Raf. New Fl. Am. 2:63. 1837. ‘‘Caro- lina and Florida.” Flowers, mid-July to mid-October. Fruit, late-September to October. DisTRIBUTION: Moist sandy soil, nearly throughout the coastal PENNELL: STUDIES IN THE AGALINANAE 423 plain. Abundant from New Jersey to South Carolina, in New Jersey in the middle district and coastal strip, absent from the New Jersey, probably also from the Wilmington, N. C., pine barrens; southward abundant near the coast in Georgia, extending to the Florida Keys, of occasional occurrence in the Altamaha grit region of Georgia, more common inland. Westward occa- sional, especially near the Gulf coast, and probably frequent north of the pine belts. A characteristic plant of coastal regions, edges of salt marshes. Widely distributed in the eastern United States, most abundant in the coastal plain. PLANTS AND SPECIMENS EXAMINED: New Jersey: Keasbey; South River; Spotswood; Keyport; Farm- ingdale; New Egypt; Burlington; Delair; Griffith’s Swamp; Kirkwood; South Westville; Clarksboro; Mickleton; Swedes- boro; Deal; Belmar; Chadwick; Toms River; Seaside Park; Forked River; Waretown; Barnegat Pier; Cox’s; Manahawken; Absecon; Atlantic City; Ventnor; Ocean City; Ocean City Junction; Avalon (4004); Wildwood; Middletown; Cape May Court House (2602, 2603); Bennett; Cape May. Pennsylvania: Tinicum (3598). Delaware: Porter; Ellendale; Rehoboth. Maryland: Kent Island; Abbey’s Island; Laurel; Lanham; Buena Vista (2640); College Park; Hyattsville; Bladensburg. District of Columbia: Anacostia; Terra Cotta (2678, 2679); Holmead Swamp; District Line (2639); Washington. Virginia: Alexander Island (2671); Arlington (2672); Alexandria; Four-mile Run; Seven Pines (4946); Smith’s Island; Norfolk; Virginia Beach; Munden. North Carolina: Weldon (4948); Rocky Mount (4932); Wilming- ton (4914, 4927); Southport. South Carolina: Ebenezer; Santee Canal; Monks Corner (4876); Otranto (4869); Charleston; Yemassee (4850, 4854); Bluffton; Aiken. Georgia: Thomson; Thalmann (4811); Waycross (4784); Fitz- gerald; Naylor (4746, 4753); Thomasville (4735A); Cordele (4769); De Soto (4758); Leslie (4767). Florida: Jacksonville (4799); San Pablo (4806); Miami; Black 424 PENNELL: STUDIES IN THE AGALINANAE Point; Homestead; Camp Longview; Long Prairie; Bull Key; Cape Florida; Fort Myers; Lakeland; St. Petersburg; Lake City; St. Marks (4703); Apalachicola. Alabama: Mobile. Mississippi: Ora; Long Beach; Pass Christian (4357). 4. AGALINIS VIRGATA Raf. New Fl. Am. 2: 62. 1837 Agalinis virgata Raf. ‘‘Glades of Pine woods in South New Jersey near Mullica Hill, &c.’’ No type known to exist. Gerardia racemulosa Pennell Torreya 11: 15. 1911. ‘“Type.— Parkdale, Camden Co., N. J., F. W. Pennell 2692 Coll. Sept. °27, 1910, in Herb. Acad. Nat. Sci. of Phila.” : Flowers, September to mid-October. Fruit, mid-September to October. = DIsTRIBUTION: In the coastal plain, New Jersey, and North and South Carolina. In New Jersey restricted to the pine barrens where frequent; frequent or common in the Wilmington pine barrens; frequent or otcasional in pineland in South Carolina. Replaces A. purpurea in typical pine barrens. Restricted to | the coastal plain. PLANTS AND SPECIMENS EXAMINED: New Jersey: Hornerstown; Forked River; Pasadena; Atsion; Hammonton; Egg Harbor City; Parkdale (2692, 2604, 3584: 3626, 3808); Winslow Junction (in 1908). _ North Carolina: Newbern; Wilmington (4902, 4921). South Carolina: Monks Corner (4877); Eutawville; Charleston. - 5. Agalinis pinetorum Pennell, sp. nov.. Annual. Plant 6-8 dm. tall, much branched, branches ascend- ing, long and slender. Stem slightly angular, glabrous or nearly so. Leaves opposite, stiffly spreading, narrowly linear, thickened, very scabrous on the upper surface, those of the stem 2-2.5 cm. long. Axillary fascicles scarcely or not developed. Racemes of 8-14 mostly opposite flowers. Pedicels very short, in flower 2 mm. long or less, hardly longer in fruit. Calyx-lobes triangular- subulate, I-1.5 mm. long. Corolla 20-25 mm. long, pubescen without, pubescent within at base of upper lobes, rose-purple, 2 — yellow lines and purple spots within throat below; lobes all spread- PENNELL: STUDIES IN THE AGALINANAE 425 ing, rounded to emarginate, ciliate. Filaments lanose; anther- sacs oblong, lanose, minutely mucronate-awned at base, 2 mm. long. Style filiform 6-10 mm. long. Capsule globose, 5 mm. long. Seeds not seen. Type, St. Marks, Wakulla Co., Florida, Sept. 26, 1912, F. W. Pennell 4708, in Herb. University of Pennsylvania. Flowers, late-September to October. DisTRIBUTION: Moist soil, in pineland, southern Georgia and northern Florida. Probably common in Altamaha grit region, frequent through flat pine woods of northern Florida, west to the Apalachicola River. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: Georgia: Waycross (4781, 4791); Douglas (4775); Naylor (4750); + Thomasville (4734, 4738); Cordele (4770, 4771, 4773). ; Florida: Jacksonville (4795); St. Marks (4708); Fort Gadsden (4688); Apalachicola. 6. Agalinis delicatula Pennell, sp. nov. Annual. Plant 6-8 dm. tall, branched, branches long and very slender. Stem angular, glabrous. Leaves opposite, spreading, almost filiform, scabrous on the upper surface, those of the stem I-2 cm. long. Axillary fascicles scarcely developed. Racemes of 8-12 mostly opposite flowers. Pedicels very short, in flower 2 mm. long or less. Calyx-lobes subulate, 1.5-2 mm. long. Corolla 20 mm. long, sparingly pubescent without, pubescent within at base of upper lobes, rose-purple, 2 yellow lines but (in this collec- tion) no purple spots within throat below, lobes all spreading, rounded to emarginate, minutely ciliate. Filaments lanose; anther-sacs oblong, lanose, mucronate-awned at base, 1.5 mm. -long. Style filiform. Fruit not seen. Type, Ponce de Leon, Holmes Co., Florida, Sept. 17, 1912, F, W. Pennell 4661 in Herb. University of Pennsylvania. Nearest to A. pinetorum of which this may possibly prove a form, but the plants seen appeared strikingly distinct, and occurred in a different region, west of the known range of that species. Flowers, mid-September to October. DIstRIBUTION: Most sandy pineland, western Florida. Possibly frequent in the west Florida pine hills. Restricted to _ the coastal plain. Type only seen. 426 PENNELL: STUDIES IN THE AGALINANAE 7. AGALINIS HarPERI Pennell in Small, Flora of Miami, 167. 1913 Agalinis Harperi Pennell. ‘‘Type.—St. Marks, Wakulla County, Florida. F. W. Pennell, 4707 coll. September 25, 1912”’ in Herb. University of Pennsylvania. Annual. Plant 4-8 dm. tall, relatively sparingly branched. Stem slightly angular, glabrous or nearly so. Leaves opposite, rather stiffly spreading, narrowly linear, scabrous on the upper surface, those of the stem 2-3.5 cm. long. Axillary fascicles scarcely or not developed. Racemes of 8-20 mostly opposite flowers. Pedicels very short, in flower less than 2 mm. long, hardly longer in fruit. Calyx-lobes triangular-lanceolate to tri- -angular-subulate, 1 mm. long or less. Corolla 15-18 mm. long, pubescent without, pubescent at base of upper lobes within, pink- purple, 2 yellow lines within throat below, small purple spots mostly along these; lobes all spreading, rounded to truncate, ciliate. Filaments lanose; anther-sacs oblong, lanose, acute at base, 1.5 mm.long. Capsule 4-5 mm. long. Seeds reticulated with dark brown ridges, areas between these relatively dark, with finer cross lines. Flowers, mid-September to mid-October. Fruit, October. DISTRIBUTION: Moist sandy pineland, borders of salt marsh, etc. Flat pine woods’ of southern Georgia, south through the Florida peninsula. Apparently most frequent in the Everglades. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: Georgia: Thalmann (4810); Thomasville (4726). Florida: Camp Longview; St. Marks (4701, 4707, 4711). 8. AGALINIS FASCICULATA (EIl.) Raf. New Fl. Am. 2: 63. 1837 Gerardia fasciculata Ell. Sketch 2: 115. 1824. ‘‘Grows prin- cipally in lands subject to occasional inundation from the ocean—on Eding’s Island near Beaufort very common.” Type seen in the Elliott Herbarium at the Charleston Museum. Gerardia purpurea fasciculata (Ell.) Chapm. Fl. So. U. S. 300. ' 1860. Flowers, August to October. Fruit, September to October. DISTRIBUTION: Moist to dry sandy, loam, or clay soil; depres- sions among sand-dunes of beach, edges of salt marsh, or loam soil PENNELL: STUDIES IN THE AGALINANAE 427 in limestone districts, etc.; especially in old fields, almost the only Agalinis to be met with in cultivated soil. In the coastal plain, Norfolk Co., Virginia (where perhaps introduced); South Carolina to Louisiana, mostly near the coast. Common near the coast in South Carolina and Georgia, and apparently through most of the Florida peninsula. Apparently absent or rare in Altamaha grit region, west Florida pine hills, or inland from the coast in Ala- bama. In clay soil in Georgia north of the Altamaha grit. Near the coast westward to Louisiana. Abundant in alluvial districts near the lower Mississippi River, and in the hill country of West Feliciana Parish. Extending westward to Texas, and north through the lower Mississippi Valley. mparnad introduced on ballast at Philadelphia. PLANTS AND SPECIMENS EXAMINED: Virginia: Northwest. South Carolina: Santee Canal; St. Johns; Otranto (4868); Sulli- van’s Island (4860); Charleston (4863); Edisto Island; Aiken; Yemassee (4849); Beaufort; Hilton Head; Bluffton; Goat Island. Georgia: Brunswick (4818); Cumberland « Island; Waycross (4792); Naylor (4747, 4751); Valdosta (4740); Thomasville (4735); Cordele (4755, 4772); Leslie (4761, 4766). Florida: Jacksonville (4793); St. Augustine; Orange City; Eustis; Minneola; Clarcona; Killarney; Merritt’s Island; Fort Lauder- dale; Miami, Alapattah; Cocoanut Grove; Homestead; Biscayne Bay; Big Pine Key; Sanibel; Polk Co.; Tampa; St. Petersburg; Fort King; Gainesville; Lake City; Monticello (4718); St. Marks (4706, 4717); Tallahassee (4695, 4697); River Junction (4669); Apalachicola (4675, 4680). Alabama: Mobile; Cedar Point. Mississippi: Ocean Springs; Biloxi (4370); Pass Christian (4356); Natchez. ‘ Louisiana: Mandeville; Covington; Amite City; Hammond; Slaughter (4267); Baines (4276); Catalpa (4303, 4304, 4330). \ 9. Agalinis georgiana (C. L. Boynton) Pennell, comb. nov. Gerardia georgiana C. L. Boynton in Biltm. Bot. Stud. 1: 148. 1902. ‘‘In the pine barrens near Cordele, Dooly County, 428 PENNELL: STUDIES IN THE AGALINANAE Georgia, in September, 1901. . . . in moist sandy soil in pine barrens. . . . The type specimens are deposited in the Bilt- more Herbarium.” Type, collected Sept. 18, 1901, seen in the Biltmore Herbarium. Flowers, mid- to late-September. Fruit, late-September to October. DISTRIBUTION: Dry sandy or clay soil, in pineland. Southern Georgia, southern Alabama, and northern Florida. Apparently occasional in flat pine woods of southern Georgia and northern Florida; near Cordele, Georgia; frequent in pine hills of western Florida and southeastern Alabama. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: Georgia: Valdosta (4739); Thomasville (4728); Dooly Co. Florida: Fort Gadsden (4693); Chipley (4665); Ponce de Leon (4662); Milligan (4586). Alabama: McRae (4609); Florala (4629, 4632). 10. Agalinis pulchella Pennell, sp. nov. Annual. Plant 6-10 dm. tall, widely branching. Stem angular, scabrous. Leaves spreading, linear, scabrous on the upper surface, those of the stem opposite, 2-3 cm. long. Axillary fascicles conspicuously developed, but mostly share? than the leaves. Racemes of 4-6 mostly alternate flowers, uppermost one, by arresting of stem-apex, often suggesting a tennitial flower. Pedicels slender, in flower 15-30 mm., in fruit 25-40 mm. long, much longer than the bracts. Calve-lobes minute, less than .5 mm. long, subulate. Corolla 25-30 mm. long, minutely pubescent without, pubescent within at base of upper lobes, rose-purple, 2 yellow lines and relatively large purple spots within throat below; lobes all spreading (upper much reflexed), notched at apex, ciliate. Filaments lanose; anther-sacs ovate, evidently short-awned, densely wooly over entire surface, 3 mm. long. Style slender, 10-20 mm. long. Capsule globose, 5-6 mm. long. Seeds reticu- lated, with dark-brown ridges, areas between these more or less broadly hexagonal, pale, not finely reticulated. Type, Ponce de Leon, Holmes Co., Florida, Sept. 17, 1912, F. W. Pennell 4658, in Herb. University of Pennsylvania. Flowers, September. Fruit, October. PENNELL: STUDIES IN THE AGALINANAE 429 DistriBuTION: Dry open, sandy pineland, southern Georgia, and northern Florida, westward to Louisiana. Frequent in the Altamaha grit region of Georgia, the flat pine woods southward to the Gulf coast in northern Florida, through the pine hills of western Florida and southeastern Alabama, and in Mobile Co., Alabama. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: Georgia: Waycross (4779); Douglas (4776); Moultrie; Thomas- ville (4731); Cordele. Florida: Gadsden Co.; Fort Gadsden (4692); Chipley (4650, 4663); Ponce de Leon (4658); Milligan (4587). Alabama: McRae (4642); Theodore (4427, 4452, 4454, 4455: 4493, 4515). | 11. AGALINIS FILIFOLIA (Nutt.) Raf. New Fl. Am. 2:65. 1837 Gerardia filifolia Nutt. Gen. Pl. N. Am. 2: 48. 1818. ‘““HAB. In West Florida. Dr. Baldwyn.” No type in Herb. Acad. Nat. Sci. Philadelphia. Flowers, September to early-October. Fruit, October. DIsTRIBUTION: Rather dry, sandy pineland, southern Georgia and Florida. Frequent or common in flat pine woods of southern Georgia and northern Florida, south through the Florida peninsula. to Miami, west to Apalachicola, and along the coast to Santa. Rosa _ Island, possibly reaching extreme southern Alabama. Plant from Santa Rosa Island, remarkably fleshy, possibly in brackish situation. PLANTS AND SPECIMENS EXAMINED: Georgia: Sunbury; Brunswick (4821, 4828); Waycross (4785); Naylor (4752); Valdosta (4741). Florida: Jacksonville (4800); South Jacksonville; St. Nicholas; San Pablo (4803); Mayport; Pablo Beach; St. Augustine; Clarcona; Tillman; Miami; Manatee; Fort Gadsden (4694) ; Apalachicola (4671, 4673); St. Vincent; Santa Rosa Island. Alabama: , Gates (probably really from Florida.) 12. Agalinis Holmiana (Greene) Pennell, comb. nov. Gerardia Holmiana Greene, Pittonia 4: 52. 1899. “Plentiful in open pine and oak groves along Michigan Avenue south of 430 PENNELL: STUDIES IN THE AGALINANAE the Soldiers’ Home grounds near Brookland, D. C., collected by Mr. Holm and the writer, 20 Oct., 1898.’’ No specimen of this date seen, but one in Herb. N. Y. Bot. Gard., of Dr. Greene’s collecting, from Brookland, D. C., dated Oct. 16, 1898, may stand as the type. ; Flowers, late-August to mid-October. Fruit, mid-September to October. DIsTRIBUTION: Dry sandy pineland. Long Island to Florida and Alabama; irregularly distributed. Common in the pine barrens of New Jersey, sparingly in the middle district of the same state; common on thé Potomac formation between Baltimore and Washington; common in Wilmington pine barrens of southeastern North Carolina, and probably so near the coast to Charleston, South Carolina; inland probably in the fall line sand hills through South Carolina and Georgia, apparently into Alabama; at Tampa, Florida. Rather narrower-leaved and more setaceous in true pine barrens. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: New Jersey: South River; Spotswood; Freehold; Tomlin; Frank- linville; Pasadena; Woodmansie; Jackson; Atco (3544, 3628); Malaga; Egg Harbor City; Absecon. Maryland: Glen Burnie; Riverdale; Lanham; Cherry Grove (2644); Sligo Mill Road (2657); Oxon Hill. District of Columbia: Takoma (2655); Lamond (2656); Terra Cotta (2680); Brookland (2662, 2669); Washington. North Carolina: Newbern; Wilmington (4904, 4023, 4929); Fayetteville. South Carolina: Columbia; Charleston (4864); Aiken. Georgia: Augusta; Burke Co.; Butler. Florida: Tampa. Alabama: , Metsner(?). 13. AGALINIS SETACEA (Walt.) Raf. New Fl. Am. 2: 64. 1837 Anonymos setacea Walt. Fl. Carol. 170. 1788. No type locality given, presumably should be from South Carolina, but appar- ently from farther west. Type in the British Museum iden- tified by Dr. A. B. Rendle as agreeing with my number 4757- PENNELL: STUDIES IN THE AGALINANAE 431 Gerardia setacea (Walt.) J. F. Gmel. curante L. Syst. Nat. ed. O27 - 1991. Gerardia Plukenetit Ell. Sketch 2: 114. 1824. ‘‘Grows in wet spungy soils, very common between the Oakmulgee and Chata- houchie Rivers.’’ Type seen in the Elliott Herbarium at the Charleston Museum. Statement of habitat probably due to confusion with A galinis pinetorum Pennell. Agalinis Plukenetii (Ell.) Raf. New Fl. Am. 2: 63. 1837. Agalimis setacea (Walt.) Raf. As to synonymy, not description, _ the latter probably applying to A. erecta (Walt.) Pennell. Gerardia filifolia Gatesii Benth. in DC. Prodr. 10: 518. 1846. “In Alabama (Gates!).’”’ Type in the Kew Herbarium, identified, from fragment sent me, as this species. Flowers, mid-September to October. Fruit, not seen, probably late-October to November. DistrRiBpuTION: Dry open sandy pineland. In the coastal plain from western Georgia and northern Florida to eastern Mississippi. Frequent from Sumter County, Georgia, westward through the pine hills of western Florida and southern Alabama to southeastern Mississippi, abundant toward the coast. One record from the Florida Keys,—Pine Key, Blodgett—likely due to mixing of labels. Occasional above the fall line in northern Ala- bama and northern Georgia. PLANTS AND SPECIMENS EXAMINED: Georgia: Cobb (4757); Cuthbert. Florida: Apalachicola (4672); Milligan (4583, 4584); Milton (4509, 4570). , See Alabama: Auburn; Wright’s Mill; Tuskegee; Clayton; Florala (4623); Bay Minette (456z); Mobile; Spring Hill (4524); Crichton (4523); Theodore (4420, 4457, 4401, 4517). Mississippi: Meridian; Biloxi (4382). 14. Agalinis laxa Pennell, sp. nov. Annual. Plant 6-10 dm. tall, widely and very laxly branched. Stem nearly terete below, slightly angled above, glabrous. Leaves ‘spreading, opposite nearly throughout, narrowly linear to nearly filiform, nearly glabrous, those of the stem 2-3 cm. long. Axillary fascicles scarcely or not developed. Racemes of 3-8 mostly 432 PENNELL: STUDIES IN THE AGALINANAE opposite flowers. Pedicels in flower 15-30 mm., in fruit 25-50 mm. long, 4-5 times exceeding the bracts. Calyx-lobes minute, less than 0.5 mm. long, acute. Corolla 15-18 mm. long, minutely pubescent without, pubescent with pink hairs within at base of upper lobes, pink-purple, 2 yellow lines, and small purple spots especially along these, within throat; lobes all spreading, rounded, ciliate with pink hairs. Filaments sparingly lanose, upper shorter pair nearly glabrous; anther-sacs ovate, mucronate at base, pale, lanose, 1.5 mm. long. Style 4-5 mm. long. Capsule globose- ovoid, 4-5 mm. long. Seeds nearly black, small for the genus, with relatively heavy longitudinal and connecting ridges. Type, Brunswick, Glynn Co., Georgia, Oct. 10, 1912, F. W. Pennell 4824, in Herb. University of Pennsylvania. Flowers, late-September to October. Fruit, late-October. DISTRIBUTION: Dry sandy pineland, river sand hills and old dunes, near the coast, South Carolina to Florida. Frequent from Brunswick, Georgia to Jacksonville, Florida; no specimens seen from farther south on Atlantic Coast. One specimen from Her- nando County, Florida. Probably frequent on river sand hills in lower Georgia and northeastern Florida. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: South Carolina: Monks Corner (4880). Georgia: Brunswick (4824); Bonnyman (4778); Waycross (4783). Florida: Jacksonville; San Pablo (4805); Pablo Beach (4801); Hernando Co. 15. Agalinis oligophylla Pennell, nom. nov. Gerardia Plukenetit microphylla A. Gray, Syn. Fl. N. Amer. II. I: 293. 1878. ‘Louisiana, Drummond, Hale. Keys of Florida, Blodgett, &c.’’ Louisiana material to be counted as typical. I have not seen the type. For Florida citation see under A galinis setacea (Walt.) Raf. Gerardia microphylla (A. Gray) Small, Fl. S. E. U. S. 1077. 1338. 1903; not Agalinis microphylla Raf. New Fl. Am. 2: 65. 1837. DisTRIBUTION: Probably moist pineland, southern Louisiana, east of the Mississippi River at Jackson, East Feliciana Parish, more frequent westward. One old specimen seen labeled, probably incorrectly, as from Alabama. Restricted to the coastal plain. PENNELL: STUDIES IN THE AGALINANAE 433. SPECIMENS EXAMINED: (2?) Alabama: , J. Torrey. Louisiana: Jackson. 16. AGALINIS APHYLLA (Nutt.) Raf. New Fl. Am. 2: 65. 1837 Gerardia aphylla Nutt. Gen. Plant. N. Am. 2: 47. 1818. “HAB. From North Carolina to Florida, where it was first detected by Dr. Baldwyn.” Type seen in Herb. Acad. Nat. Sci. Philadelphia; accompanied by fruiting plant of Agalinis erecta (Walt.) Pennell. Gerardia aphylla grandiflora Benth.* in Hook. Comp. Bot Mag. 1: 174. 1835-6. “‘Jacksonville.”’ Drummond. - Agalinis microphylla Raf. New Fl. Am. 2: 65. 1637. “In Florida, collected by Leconte (Collins herb.).”’ Flowers, mid-September to early November. Fruit, October to November. DIsTRIBUTION: Moist sandy pineland near the coast, North Carolina to Florida and Louisiana. . Occasional from the Wilming- ton pine barrens southward, through North and South Carolina; most abundant in the flat pine woods of southern Georgia and northern Florida, frequent in the Altamaha grit region of Georgia; the west Florida pine hills; less frequent westward to Louisiana. Apparently does not occur in the Florida peninsula. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: North Carolina: Wilmington. South Carolina: Santee Canal. Georgia: Thalmann (4808); Brunswick (4819); Waycross (4789); Coffee Co.; Naylor (4748). Florida: Tecate (4814); Jacksonville (4798); South Jacksonville; St. Marks (4712); Fort Gadsden (4682); Apalachicola (4676); Chipley (4647, 4664) ; Point Washington; Ponce de Leon (4655); illigan. Alabama: Mobile; Spring Hill. ) # Dr, NE. Browa has kindly consulted the Bentham correspondence at Kew, which makes it evident that this author is responsible for the treatment of Gerardia in Hooker’s account of the plants of Drummond's collections. Also it seems evident. that the paper containing this was not published until early in 1836. 434 PENNELL: STUDIES IN THE AGALINANAE Mississippi: Ocean Springs; Biloxi; Pass Christian; Nicholson. Louisiana: Abita Springs. 17. Agalinis decemloba (Greene) Pennell, comb. nov. Gerardia decemloba Greene, Pittonia 4: 51. 1899. “Plant not uncommon about Brookland, D. C., inhabiting grassy knolls and hillsides bordering on pine woods.’’ A specimen in Herb. N. Y. Bot. Gard. collected by Dr. E. L. Greene at Brookland, D. C., in Oct. 1898, may stand as the type. Flowers, late-August to mid-September. Fruit, late- Scotenbee to October. DIsTRIBUTION: Dry soil, light sand or clay, in the coastal plain in Kent Co., Delaware, frequent on Potomac formation near Washington, D. C.,-and probably occasional south North Carolina. Occasional inland in the Piedmont region from Penn- sylvania to North Carolina. Apparently has a fragmentary distribution, but not well understood. PLANTS AND SPECIMENS EXAMINED: Delaware: Felton. Maryland: Buena Vista (264r); Lanham; Forest Glen; Silver Springs. District of Columbia: Takoma Park (2654); Brookland ae 2661, 2677, 4950). 18. Agalinis tenella Pennell, sp. nov. Annual. Plant 5-8 dm. tall, laxly branched, branches slender. Stem angled, glabrous throughout. Leaves spreading, linear- filiform to nearly filiform, acutish to acute, opposite nearly through- out, those of the stem I-1.5 (~2) cm. long. Axillary fascicles none. Racemes of 8-12 mostly opposite flowers. Pedicels slender, in flower 8-20 mm. long, in fruit reaching 25 mm. long. Calyx-tube reticulate-venulose; lobes minute, apiculate. Corolla 15-20 mm. long, minutely pubescent without, pubescent with pink hairs within at base of upper lobes, rose-pink, 2 yellow lines and small diffused purple spots within throat; lobes all spreading, more or less emarginate, ciliate. Pilamenes lanose especia toward apex; anther-sacs lanceolate, mucronate at base, 2 mm. long, villose with pink hairs 1.5 mm. long. Style slender, 5-8 mm. long. Capsule globose-ovoid, much flattened at base, 4 mm. a Seeds narrow, yellowish-brown. PENNELL: STUDIES IN THE AGALINANAE 435 Type, Thomasville, Thomas Co., Georgia, Sept. 28, 1912, F. W. Pennell 4727, in Herb. University of Pennsylvania. Flowers, mid-September to mid-October. Fruit, October. DiIsTRIBUTION: Dry sandy pineland, in the coastal plain from South Carolina to Florida and Alabama. Occasional in lower South Carolina; most abundant in Altamaha grit region of Georgia; less frequent in upper edge of flat pine woods of southern Georgia entering north central Florida at Gadsden County, and in middle Georgia entering east central Alabama at Lee County. In the Altamaha grit region common, mostly replacing A. erecta Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: South Carolina: Otranto (4871); Eutawville; Yemassee (4853). Georgia: Waycross (4782, 4786); Naylor (4744); Douglas (4777); Moultrie; Thomasville (4727); Cordele (4768, 4774); Cobb _ (4750); Leslie (4764). | Florida: Chattahoochie. Alabama: Auburn. 19. AGALINIS ERECTA (Walt.) Pennell, in Small, Fl. Florida Keys, 133- 1913 Anonymos erecta Walt. Fl. Carol. 170. 1788. No type locality given, presumably from Berkeley County, South Carolina. There is no type in the Walter collection in the British Museum. Of the species occurring in Berkeley County, those which best answer the description are Agalinis Holmiana (Greene) Pen- nell, Agalinis laxa Pennell, and the following. Of these the first two are long-pediceled, and very lax, the bracts in the first are scarcely conspicuously shorter than the peduncles, moreover both are relatively infrequent. The third species, the following, in its strict erect habit, its pedicels not con- spicuously long, but with bracts conspicuously shorter, seems seems best to fit Walter’s description; moreover, it appears to be much the most abundant species of the district. Gerardia erecta (Walt.) J. F. Gmel. cwrante Linn. Syst. Nat. ed. 13, 928. 1791. 4 5 436 PENNELL: STUDIES IN THE AGALINANAE Gerardia setacea parvifolia Benth.* in Hook. Comp. Bot. Mag. I: 174. 1835-6. ‘Jacksonville.’ Drummond. Agalinis obtusifolia Raf. New Fl. Am. 2: 64. 1837. ‘‘West Tennessee, Alabama and Florida.’’ Description in greater part or entirely of this species, though the Tennessee specimen could hardly belong here. Gerardia parvifolia (Benth.) Chapm. Fl. So. U. S. 300. 1860. Agalinis parvifolia (Benth.) Small in Britt. & Br. Ill. Fl. ed. 2. @<-212.-° 1913. Flowers, early-September to mid-October. Fruit, October. DistRIBUTION: Moist to dry sandy pinelands, in the coastal plain from North Carolina to Florida and Louisiana. Occasional or frequent in eastern North Carolina; common in the Wilmington _ pine barrens, and southward near the coast to Charleston, South Carolina; less frequent in the Altamaha grit region and inland in Georgia; common through the flat pine woods of Florida, south through the peninsula and on the Florida Keys; common westward through the pine hills of west Florida, decreasing inland in south- eastern Alabama; and common near the Gulf coast westward to eastern Louisiana. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: North Carolina: Rocky Mount (4933); Wilmington (4970, 4915, 4920); Brunswick Co. South Carolina: Monks Corner (4879); Otranto (4870). Georgia: Sunbury; Coffee Co.; Moultrie; Thomasville (4733); Leslie. Florida: Tisonia (4815); Jacksonville ou San Pablo (4804); Green Cove Springs; St. Augustine; Big Pine Key; Marco; Fort Myers; Polk Co.; Tampa; Lake City; St. Marks (4710); Fort Gadsden (4685); Apalachicola; Chipley (4646, 4667); Ponce de Leon (4659); Paxton (4640); De Funiak Springs; Milligan (4596). Alabama: McRae (4614); Florala (4634); Bay Minette (4548, 4562); Mobile; Spring Hill (4526); Theodore (4428, 4453); Hollander’s Island (4503). Mississippi: Waynesboro; Ocean Springs; Biloxi (4399); ‘Manuel; Gulfport; Long Beach; Pass Christian (4363); Nicholson. * See footnote on page 433. PENNELL: STUDIES IN THE AGALINANAE 437 Louisiana: Pearl River; Bayou Lacombe; Abita Springs (4226, 4227, 4231). 20. AGALINIS TENUIFOLIA (Vahl.) Raf. New Fl. Am. 2: 64. 1837 Gerardia tenuifolia Vahl, Symb. Bot. 3: 79. 1794. ‘‘Habitat in America septentrionali.”” Type in Herb. Universitetets botan- iske Museum, Copenhagen, Denmark, collected by Von Rohren, and said to be probably from Philadelphia, is iden- tified by Dr. C. H. Ostenfeld as agreeing with material (my number 2681) sent from eastern Pennsylvania. Flowers, mid-August to mid-October. DIstRIBUTION: Moist to dry sand or loam, deciduous or mixed woodland, widely distributed and common through the eastern United States above the fall line, in the coastal plain locally frequent, especially in limestone districts. In New Jersey occa- sional in the middle and Cape May districts, occasional south- ward near the fall line; in Sumter County, Georgia; frequent in red loam soil in central northern Florida; in limestone in south- eastern Alabama and western Florida; and in alluvial soil in southern Alabama and Louisiana. A larger plant southward. PLANTS AND SPECIMENS EXAMINED: New Jersey: New Egypt; Camden; Clarksboro; Swedesboro;~* Bennett; Cold Spring. Delaware: Van Dyke. Maryland: Ardwick (2645); Oxon Hill. , District of Columbia: Brookland (2658). Georgia: De Soto (4759); Leslie. Florida: Monticello (4721); Tallahassee (4699); Chattahoochie; River Junction (4670); Aspalaga; Paxton (4601). Alabama: Chapel Hill, Covington Co. (4619); Florala (4597, 4606); Cocoa; Mobile; Crichton (4522). Mississippi: Meridian; Jackson. Louisiana: Mandeville(?); Catalpa. 21. Agalinis divaricata (Chapm.) Pennell, comb. nov. Gerardia divaricata Chapm. FI. So. U. S. 299. Mar. 26, 1860. ‘Low sandy pine barrens.’ No type indicated, but abundant 438 PENNELL: STUDIES IN THE AGALINANAE material of this species collected and distributed by the describer. Gerardia Mettauert Wood, Class Book 530, Dec. 1, 1860. ‘Wet sandy places, Middle Fla. (Dr. Mettauer).’’ Type seen in Herb. Columbia University. Gerardia Mettaueri clausa Wood, Class Book 530, Dec. 1, 1860. “With the others,” i. e. the species and G. Mettaueri nuda Chapm. Flowers, September to peseick: Fruit not seen. DIsTRIBUTION: Dry sandy pineland, western Florida and adjacent southeastern Alabama. Abundant through the west Florida pine hills, eastward through the middle Florida flat woods to Apalachee Bay. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: Florida: St. Marks (4716); ‘near Tallahassee; Gadsden Co.; Carrabelle; Fort Gadsden (4684, 4687); Apalachicola (4679); Chipley (4644, 4668); Ponce de Leon (4657); Portland; Milligan (4593); Milton (4572). Alabama: Florala (4622, 4624, 4627, 4633). 22. Agalinis filicaulis (Benth.) Pennell, comb. nov. _Gerardia aphyilla filicaulis Benth.* in Hook. Comp. Bot. Mag. I: 174. 1835. “Jacksonville.” Drummond. Gerardia filicaulis (Benth.) Chapm. FI. So. U. S. 299. Mar. 26, 1860. Gerardia Mettuaeri nuda Wood, Class Book 530, Dec. 1, 1860. Under Gerardia nuda Wood (1870), ‘‘ Middle FI. (Dr. Mettauer, 1855).” Gerardia nuda Wood, Am. Bot. and Flor. 231. 1870. Flowers, mid-September to early November. F ruit, October to November. DIsTRIBUTION: Moist grassy sandy pineland, in the coastal plain from southern Georgia and northern Florida to eastern Louisiana. Frequent in the flat pine woods of southern Georgia and northern Florida; occasional or rare in the Altamaha grit region; frequent in the pine hills of western Florida and adie" * See footnote on page 433. PENNELL: STUDIES IN THE AGALINANAE 439 southeastern Alabama; westward near the coast to southern Mississippi and probably eastern Louisiana. Restricted to the coastal plain. PLANTS AND SPECIMENS EXAMINED: Georgia: Waycross (4788); Naylor (4749); Tyty. Florida: Jacksonville (4796); St. Marks (4709); Fort Gadsden (4689); Apalachicola (4677); Ponce de Leon (4660); Paxton (4643); De Funiak Springs; Milligan (4594). Alabama: McRae (4608); Florala (4626). Mississippi: Ocean Springs; Biloxi; Pass Christian. Louisiana: ‘‘ New Orleans.” 7. Drummond. UNIVERSITY OF PENNSYLVANIA. Some observations concerning the reactions of the leaf hairs of Salvinia natans FRANK MARION ANDREWS AND Max Mapes ELLIS If one observes plants of Salvinia natans which are in an active condition, it will be observed that the hairs on the leaves show drops of liquid which they secrete. It was noticed from time to time that some of the leaves of the Salvinia which was kept growing in a large tub in the green-house bore small drops of dark colored liquid. Frequently a dead dipterous insect was found on the surface of a leaf somewhat enveloped in a white fiingus which was also attached to the leaf hairs. These occurrences coupled with the superficial resemblance of the leaves of Salvinia to those of Drosera suggested the possibility that this fern might also be able to absorb food from decaying matter on the leaves. It was first desirable to see if organic matter would go into solution on the leaves of this plant. Experiment 1. Two large battery jars of Salvinia were iso- lated from the tub and small soft-bodied dipterous and thysanuran insects, such as were found in and about the tub, were crushed and placed upon many of the leaves. By the end of a week all of these insects were covered with a white fungus, apparently the same as that previously noted on the Salvinia in the tub. During the second week almost all of this fungus disappeared and most of the inserts were reduced to drops of dark colored liquid. Onlya few of the insects dried up. This experiment makes it entirely probable that some of the dark colored drops found on the leaves of the Salvinia in the tub contained organic remains. This reduc- tion of the organic matter to a solution may have been the result of any or all of these three actions, (1) simple decomposition, (2) action by the Salvinia or (3) action by the fungus. Three experi- ments were made to ascertain whether or not the Salvinia con- tributes to this decomposition of the organic matter. Experiment 2. Several large crystallizing dishes were filled with fresh Salvinia. On the surface of about every other leaf 441 442 ANDREWS AND ELLIS: REACTIONS OF HAIRS OF SALVINIA was placed a small piece, approximately one millimeter cube, of the white of a hard boiled egg. The weight of the cubes was not great enough to break down the leaf hairs on which they rested. Similar cubes of the boiled white of egg were placed on clean glass in the dishes just above the surface of the water, as controls. During the first twenty-four hours all of the white of egg, both control and experiment, became rather transparent. At the end of forty-eight hours several of the leaves were removed and ex- amined under a microscope. The cubes of egg had retained their shape perfectly, that is, there had been no rounding off of the edges as has been previously noted in the digestion of the white of egg by Drosera. A certain amount of the white of egg had been removed, however. Each of the leaf hairs on which the cube was resting had penetrated it and reached nearly through the block of boiled egg. The extreme tips of the prongs of each hair were thus firmly imbedded in the white of egg, but the remainder of the leaf hair scarcely touched the cube, being smaller than the chamber it occupied. This little chamber resembled a hole made in ice with a warm metal rod, being slightly larger than the leaf hair at every point except the tip. In addition to this reaction by the leaf hairs supporting the cube of white of egg, the row of hairs immediately around the cube which were not under it but which just touched it had also reacted. These hairs had bent in on all sides, penetrating the cube of white of egg in the same manner as those hairs on which the cube was resting, except that they entered the cube from the side; that is, there seemed to have been a positive chemotaxis on the part of the leaf hairs with reference ‘to the white of egg. This.experiment was continued for several days, with many plants and always with the result as just stated. Special precaution was necessary to keep the cubes of boiled egg from drying out too quickly. However, the white of egg did not injure the leaves in any way as far as could be determined during the six days it rested on them. Experiment 3. Other cultures of Salvinia were isolated and small drops of uncooked white of egg placed on most of the leaves. Control drops of water of the same size were also placed on many leaves. The drops of both white of egg and water rested as tiny spheres on top of the leaf hairs for the first. twenty-four hours. q ANDREWS AND ELLIS: REACTIONS OF HAIRS OF SALVINIA +443 During the second twenty-four hours the drops of white of egg in many instances broke, forming irregular patches which sank down to the surface of the leaf, completely surrounding the hairs which had supported them. At the same time the neighboring leaf hairs bent over into the white of egg in much the same manner as noted regarding the cubes of cooked white of egg. The control drops of water did not change meanwhile, remaining as spheres on top of the leaf hairs. Several things might have caused the drops of white of egg to act in this manner, but in the light of the solvent action of the leaf hairs on the cubes of cooked white of egg it seems probable that the leaf hairs themselves were responsible for the change which caused these drops of raw white of egg. to change. These cultures were continued for eight days. During this time several of the drops of white of egg became infected with the white fungus already mentioned. The result was the same as that with the crushed insects. The white of egg be- came steadily more liquid until it was reduced to a fluid almost as mobile as water. By close observation it could be determined that in those cases where the tiny spheres of white of egg did not break and run down on to the surface of the leaf the amount of white of egg had decreased. Experiment 4. On the leaves of fresh cultures of Salvinia drops of uncooked yellow of egg were placed. Controls were established by drops of the yellow of the egg being placed on clean glass in the dishes with the Salvinia, just above the surface of the water. The color of the egg on the leaves became noticeably paler than the control during the first twenty-four hours. At the end of forty-eight hours it had spread over a small area of the leaf surface in the same manner as the uncooked white of egg had done in the previous experiment. The color of this egg on the leaves was several shades lighter than the control and of a quite different consistency, being more like cream than the control. The same chemotaxis was displayed by the hairs surrounding the area covered by the egg. Experiment 5. It was the object of this experiment to ascer- tain whether the plant was profiting by the presence of the decay- ing or soluble organic matter on the surface of its leaves. Four different jars of Salvinia were prepared. 444 ANDREWS AND ELLIs: REACTIONS OF HAIRS OF SALVINIA In jar Number 1 the Salvinia was floated on distilled water. In jar Number 2 on a nutrient solution containing the following substances: ere WE ee ae eee eis Oh oy eee 1000.0 cc. WO cust Niet ie 2k en Pd ke PR es I.0 gm. RCOPCTE SUMO MDLG 20's Sidiice. Wee cds Se aCe 0.5 gm. PAA PCH BIN DUALE i <6. ook lc cs eae oe ce oa 0.5 gm. PICS FINOM PALE oak ais ee bk et aia 0.5 gm. Jar Number 3 contained a nutrient solution the same as that used in Number 2, except that the potassium nitrate was omitted. On the leaves of the Salvinia in this jar were placed cubes of cooked white of egg, drops of uncooked white of egg and crushed insects. Jar Number 4 also contained a nutrient solution the same as Number 3, that is, without the potassium nitrate, but the leaves in this jar were free from foreign substances. | The Salvinia in jar Number 1 began to change color on the second day. The otherwise bright green color of the leaves be- came dull and by the fifth day there were distinct yellow spots on the leaves. In a week all of the leaves in this jar were of a uniform brownish-yellow color. In Salvinia care must be taken to keep the light conditions for good growth as nearly constant as possible. It frequently happens that Salvinia plants that have been growing in somewhat weak light will, when brought into very bright light, lose their green color and die. This, however, may be prevented by gradually bringing them into strong light which they may then stand without the least i injury. The plants in jar Number 2 remained quite normal throughout the entire experiment, which lasted fifteen days. The leaves in jar Number 3 began to lose their color in six days and those in Number 4 four days. Both those with and without foreign matter on them finally lost all their color. By the end of the first week, however, there was a decided difference between the leaves in the two jars. Almost all of the leaves in both jars were turning yellow but those in jar Number 3 retained their color, especially, in the area immediately around that covered with organic matter. The ninth day all the leaves in both jars were yellow, excepting a few in jar Number 3. These few were still a little green in the area around the organic matter. i ANDREWS AND ELLIS: REACTIONS OF HAIRS OF SALVINIA 445 They continued to fade and by the fifteenth day all were entirely yellow. This experiment shows that the plant can obtain a certain quantity of its food from the decomposing organic matter offered to it in the way above described. The amount that can be taken up by the leaf hairs, however, is small and is insufficient to supply the demands of the plant. The Salvinia with organic matter on its leaves seemed to have an advantage but this advantage in nature is only occasional or as chance offers. “Experiment 6. Pieces of cinder and iron filings were carefully laid on the leaves and allowed to remain for several days. No reaction of any sort occurred among the hairs of the leaves. SUMMARY One of the most interesting results was the chemotactic re- action of the leaf hairs. This was very distinct in every case, excepting experiment 6. ) The experiments proved that the leaf hairs are capable of exerting a distinct solvent action on the organic matter placed on them. Experiments 2, 3 and 4 show this. This solvent action does not serve to remove any objectionable organic matter but the plant profits by the food derived by this action. The latter is well shown from the observations made, especially those of experiment 5. No experiment showed the leaves suffering from the presence of small amounts of organic matter on their leaves and on the contrary the leaves thus treated were the last to lose the green color as in experiment 5. The Salvinia may get only a small quantity of food in this way, as is indicated by the rapid decline of the plants even in experiment 5 when placed in a solu- tion free of potassium nitrate. The positive chemotaxis of the leaf hairs, the solvent action on the cubes of cooked white of egg and the discoloring of the yellow of the raw egg coupled with the fact that in the normal habitat this fern frequently has the bodies of small soft insects which are finally dissolved on its leaves, shows at least that some food is taken in by these leaves when it is offered. : INDIANA UNIVERSITY, BLOOMINGTON w * On the relationship between the number of ovules formed and the capacity of the ovary for maturing its ovules into seeds J. ARTHUR HARRIS I. INTRODUCTORY REMARKS In a series of papers published in part only and which need not be cited here, I have attempted by the use of the modern higher statistics to analyze the internal factors influencing seed production. | This work has consisted chiefly in determining the correlations between the degree of development of various somatic organs and the fertility of the fruit. Certain peculiarities of the fruit itself have been also considered in their relationship to capacity for seed production or to the characteristics of the seed formed. A question of considerable interest upon which very little has been published is that of the relationship between the number of ovules formed by a fruit and its capacity for maturing these ovules into seeds. To the data of this problem the present paper is a contribution. In it only questions of fact will be considered, for it is quite premature to essay any interpretation of observed relationships in more general terms. The problem in hand is to determine whether ovaries with a number of ovules above the average are more (or less) capable of developing their ovules into seeds than those below the average. A priori, ovaries with more than the mean number of ovules must be expected to produce on the average absolutely more seeds than those with less than the mean number. A Posteriori, this condition is found almost without exception. But the question which interests the physiologist is whether ovaries of any class with respect to number of ovules are more capable of seed produc- tion as shown by their developing a higher proportion of their ovules into seeds. This problem is not only of importance from the standpoint of 447 448 HARRIS: NUMBER OF OVULES FORMED the physiology of seed production, but of considerable interest from the fact that in Staphylea there is a selective mortality with respect to number of ovules, ovaries with a smaller number of ovules being less capable of developing into mature fruits than those with a larger number.* The determination of the existence of such a relationship, especially the measurement of its intensity, presents considerable difficulty. It is desirable that degree of interdependence shall be expressed in terms of correlation, but there are dangers of spurious coefficients. Again, the work so far done has demon- strated very low correlations between somatic characters, or those of the fruit, and seed production. One would, therefore, anticipate very slight relationships between the number of ovules formed per fruit and its capacity for seed development. In such cases very large and numerous series of data are desirable. II. METHOD OF ANALYSIS I believe the following method of reasoning is valid. In a series (a population, to use the technical term) of poly- spermous fruits only a portion of the ovules develop into mature seeds. Let o be the number of ovules formed and s the number of seeds matured per fruit; both are variable; s is always some pro- portion of o. If there be no relationship between the absolute number of ovules formed and the capacity of the fruit for maturing its seeds, the most probable number of seeds for any pod is 9, where ‘ p a s/o, i the bars indicating the mean value of the two variables. Now let s = s— po, or the deviation of any individual s from its probable value, on the assumption that the chances of an indi- vidual ovule developing into a seed are independent of the number of ovules in the ovary in which it is produced. The correlation coefficient between o and 2, r,z, should furnish the information we need. A convenient formula for Ps determination of this relation- * Harris, J. Arthur, Biometrika 7: 452-504. 1910; Science II. 32: 519-528. I9Q10; et pos Mo. 78: 521-538. I9I1 3 Harris: NUMBER OF OVULES FORMED 449 ship was most kindly worked for me while engaged on this problem at University College, London, by Professor Karl Pearson.* III. Discussion OF DATA This formula has been so far applied to the problem of fecundity in plants only in the case of the fruits of Cercis, Robinia and Sanguinaria and of the inflorescence of Staphylea, Celastrus and Crinum.t Only one or two published series were available for each of these species. Because of the delicate relationships which are ordinarily found between fertility and other characterst it is essential for trustworthy results that the relationship be worked out on as large a series of material as possible. It will be also advantageous if one can include a large number of sub-series differing from each other in the conditions to which they have been subjected but each homogeneous in itself. The only such series of data is that used for the working out of the relationship between bilateral asymmetry and fertility and fecundity in Phaseolus vulgaris.§ For each of these 53 series, embodying altogether over 170,000 pods, I have carried through the arithmetical routine necessary for the determination of 7.2, the correlation between the number of ovules formed per pod and the deviation of the number of seeds from their probable ‘value on the assumption of there being no relationship between the number of ovules formed and the capacity of the pod for maturing its seeds. - TABLE I gives the results. The key letters indicating the individual series permit easy reference to other published informa- tion concerning them. The second column shows the number of pods upon which the determinations are based. The third gives the coefficient of correlation between the number of ovules and Harris, J. Arthur, The Correlation between a Variable and the Deviation of a Dependent Variable from its Probable Value. Biometrika 6: 438-443. 1909. } For the results for the fruits of Cercis, and Robinia and the = GE of Staphylea see the original paper on methods. For Sanguinaria see Biometrika 7: 321-324. 1910. For Celastrus see Missouri Bot. d. Ann. Rep. 20: Ce 1909, and for Crinum see Missouri Bot. Gard. Ann. Rep. 23: 85-89. 1912. t For a review of the pertinent literature for plants see Biometrika 8: 52-65. Igtt. J. Arthur, in Roux’s Archiv. f. Entwicklungsmech. Organism. 35: 500- 522. 1912. ; 450 Harris: NUMBER OF OVULES FORMED TABLE I Sales N Fxg %oz and Probable Error 92/27 oz Sra tas ete ayer 1804 -374 —.040 =.016 — 2.52 1 1S DEG eaeh a nen OA Mey Ara 8043 “357 —.014+.008 — 1.80 BGG aes 806 107 189 +.023 — 8.27 Ta S sore goa sats 346 2177 —.234+.034 — 6.82 BW ees 467 128 —.290 +.029 —10.15 na re pee ener ee 696 -542 —.0I12 +.026 — .4 MS es oe cis Stare 6310 -247 —.102 +.006 —17.25 CE ois ot 5251 -461 —,I0I =.009 —10.98 GCs 3502 sSET —.045 =.0II — 3.94 CAST ee es ee 2656 -489 —.068 =.013 — 5.20 GG Soest 1438 -362 —.069 =.018 — 3.88 Le 4 0 eee ae eee 1227 -426 —.020 +.019 — 1.04 GER 2 ee 807 -386 —.022 =.024 — .95 LOSE for rama Oy es eae 5141 -444 +.012 +.009 -+- 1.23 jg Beep nas Sien are bv insieae 14029 -511 —.015 +.006 — 2.68 MBC Ue set 2355 -504 +.121 +.014 + 8.82 Ae ee ce ki 11230 -426 —.057 =.00 — 9.02 HBC es Se. 2614 501 +.10I +.013 + 7.69 FEED aS aye witless 5581 -418 040 +.009 — 4.47 BI Aes ure ae aos 1733 -509 +.086 +.016 + 5.32 RD eee ees 5449 500 +.026 + .009 + 2.87 TEDDC ys Baie niece 2308 -464 +.097 =.014 + 6.07 } SD enn Rage mendes sac 1473 521 +.055 =.018 + 3.17 ED} 9 Bp h en Gareserae entree 1827 -496 —.046 =.01 — 2.91 DDG ciate II59 -463 +.066 +.020 + 3.35 Ls D Pear oear ees Ue 2018 444 —.018 =.015 — 1.17 DDDG 7 Haatieeses 1542 -429 +.047 +.017 + 2.77 DI es Saviaeae: 5955 -548 +.029 +.009 + 3.29 WHC. ese eee 2077 -409 +.045 +.015 + 3.05 LG) yo oS pecesrliemaey ariae oR 5019 -446 052.010 — 5.48 DHRC rhea 2404 -438 +.055 +.014 + 4.06 WSOos she Hae 2569 B3t —.064 =.013 — 4.79 Base ae 6605 505 +.071 +.026 + 2.90 USSG 3 ee eek 1888 332 —.065 +.01 — 4.21 RISE oe oe ee 3406 -470 —.0AI +.012 — 3.60 USEC ee 1570 -322 —.091 +.017 — 5.38 WSHE 2 ca a 1743 -366 —.035 +.016 — 2.18 USHAC 3 ssa s 1936 -318 —.106+.015 — 6.95. PUD bea s ele ee 802 -376 —.018 =.024 — 74 ISON ssn ose 1810 «245 —.154+.016 — 9.91 USD es eo ae eee 851 -370 —.027 +.023 — I.17 USDC Ecko es 1619 +315 —.1I16 +.017 — 7.05 PSC ae aes 2876 -344 —.056 +.013 — 4.50 PSS eS Wi ee 7809 -367 —.049 =.008 — 6.42 BOSE oes: ore 2457 -4II +.035 +.014 + 2.57 MSH eee re ores 4541 -458 004 +.010 pas pels | Go grep ors cs anne 2117 -268 —.155+.014 —10.84 PSHE 26s eos. 3837 -398 —.028 +.011 — 2.59 PSBHC. oo ee 3180 Oe he 089 +.012 > 7-46 SD aa ay 1449 -433 +.002 +.018 + .12 BG ee oats wien 1506 -267 —.134+.017 — 7.83 PSII oo tk 1556 -421 —.016 +.017 — .92 FSDDC #2 o Fes 2646 328 —.I10I +.013 er ets HARRIS: NUMBER OF OVULES FORMED 451 number of seeds per pod, 7., while the fourth contains the correla- tions between the number of ovules per pod and the deviation of the number of seeds from their probable value. All of the values for 7,, are positive, and of a substantial order of magnitude. Both positive and negative coefficients for ros occur, and the values are low throughout. Of the 53 determina- tions, 38 are negative and 15 positive in sign. Were there no biological relationship between o and s an equal number of positive and negative values would be expected. Thus there is a deviation from equality of 11.5 * 2.46,* which is probably significant. Thus there is apparently a distinct negative relationship be- tween 0 and z. This conclusion is supported by restricting the constants upon which it is based to those which are more probably statistically significant with regard to their probable errors. Thus I find that for the 41 constants 2.5 and more times their probable error 28 are negative and 13 are positive. Of the 26 constants which are over 4 times their probable error, 21 are negative and 5 are positive. Of the 13 which are over 7 times their probable error, II are negative and 2 are positive. Taking means of the ratio of the correlation coefficients to their probable errors, I find a mean ratio of 5.10 for the negative values and of 3.87 for the positive. Thus the negative constants are as a whole more trustworthy than are the positive, although some of the positive values must be certainly regarded as trustworthy statistically. Consider next the relative magnitude of the positive and nega- tive coefficients. The 38 negative constants give a mean value of — .0732 while the 15 positive correlations give a mean value of + .0529. Thus the negative coefficients are numerically larger than the positive. I now split the materials up into the individual varieties. The results are conveniently summarized in TABLE Il. Positive coefficients occur only in Navy, Ne Plus Ultra and White Flageolet. In the two latter, only 3 of the 22 constants are positive, and neither of the three can be safely regarded as significant with regard to its probable error. In Navy, quite different conditions seem to prevail. In both strains 6 of the 9 * Calculated from .6745 V 53 X.5 X-5- 452 Harris: NUMBER OF OVULES FORMED constants are positive. Numerically, the positive constants aver- age higher than the negative, and in comparison with their probable errors they seem to be more trustworthy, the values of the ratio of 7,z to-its probable error averaging higher in the positive than -------~—0 | PLUS ° 1 i] I ZERO BAR ; “maT li ° ZERO BAR L | maT : “ TT T 1 ad see ga nee oa DtaGraM I. Intensities of correlation. Oneinterval on marginal scale = .052- in the negative cases. Just the reverse of this condition is found everywhere else. All of the positive values which are over thrice their probable error fall in the Navy strains. 1 Mea we fs a 2606 Ea Ba ONE ache eee a, pe leg ca ieg etic eae Ey Cea aa ts PP ee ee ts ae Pe RE TR Some SRG ak PT cee be IAGRAM 2. Si of corr ee gre agua y the rati their probate eel One interval on marginal scale 454 HARRIS: NUMBER OF OVULES FORMED These relationships are forcibly shown in DIAGRAMS I and II. In both of these the intensity of the correlation is shown by the & TABLE II Variety | oo of | Mean 7,,, Mean a Gol Wake ee. | BAUR pian rg mie Oe was Ss Sees — — — WEBNS tlk oi fsa ee 5 Sas | 3 —.o81 4.20 PAC WAR or oes Re ee Ss PRG ar ot ee es ae ne ws eel — — — TT es at cbn ie aaa Fo bee le 2 —.262 —8.49 wiS s -Shilogien accel see ae ee canis EPRI Social eeth te Wes oe Sean oa 8 —.055 —5.46 NAVY Theo ore cen ue eee eee | BMG eel ee ise 6 +.074 +5.48 MINUS ore es ean cen teens 3 —.037 —2.70 INN UBD csi bar ahs a ee agape Ee Plus. ies oy tas Ca 6 4.050 +3.28 einen eee Sad eae ES 3 —.039 —3.19 Ne Poses ‘Ulisa ft ie or eae sa ae ight Calas See re ee I +.017 +2.78 CE ay oe ee ee oe ee Io —.072 — 4.60 noe Fagediet ; Peewee ie ce nis der ec haub fal psi Dechert 2 +.014 “F135 vhs dive taba Wie hy ch epterarsenmtane) SUA Rees 9 —.070 — 5.40 length of the vertical lines. The broken ones extending above the zero bar measure positive values; the solid ones extending below the zero bar indicate negative coefficients. In DIAGRAM 1 the lengths of the lines are in terms of the correlation coefficients, Yoz; iN DIAGRAM 2 they are in terms of the ratio of the correlation to its probable error. In both, the large central figure represents the distribution of values for the whole material. This is analyzed into the Navy series, shown in the lower corner, and into all other series, shown in the upper right hand corner of the figures. The difference in the contribution of these two elements to the general series is very striking. IV. RECAPITULATION AND DISCUSSION In dwarf varieties of garden beans, Phaseolus vulgaris, there is but a slight relationship between the number of ovules per ovary and its capacity for maturing these ovules into seeds. So lax is this correlation that in working with only moderately large samples both positive and negative values of the iencrmr may be found in the same strain of material. Harris: NUMBER OF OVULES FORMED 455 Such a relationship does, however, exist. So far as the ma- terials available may be considered as representative of the species it is generally negative, i. e., as the number of ovules formed in- creases the capacity for maturing these ovules into seeds decreases. This conclusion is supported by the facts that the negative correla- tions are significantly more numerous than the positive, they average larger numerically, and they have a higher degree of trustworthiness with regard to their probable errors. In some varieties, however, the correlations seem to be gen- erally positive. This is true for the common Navy. All other varieties so far as studied—White Flageolet, Ne Plus Ultra, Burpee’s Stringless, Golden Wax and Black Wax—show exclu- sively or preponderantly negative correlations. Concerning the explanation of this relationship no suggestion can be made. Such an attempt would be quite premature until ample quantitative data on the nature (sign) and intensity of the relationship in a considerable series of varieties are available. Anyone venturing to suggest explanations must also fully realize that the problem is an exceedingly complex one, involving many difficulties which need not be enlarged upon here. But as matters of biological fact the results seem definitely established, and represent one further step in the analysis of the problem of fertility and fecundity in plants. CoLtp Sprinc Harsor, N. Y. INDEX TO AMERICAN BOTANICAL LITERATURE (1913) The aim of this Index is to include all current botanical literature written by Americans, published in America, or based upon American material ; the word Amer- ica being used in the broadest sense. Reviews, and papers that relate exclusively to forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and no attempt is made to index the literature of bacteriology. An occasional exception is made in favor of some paper appearing in an American periodical which is devoted wholly to botany. Reprints are not mentioned unless they differ from the original in some important particular. If users of the Index will call the attention of the editor to errors or omissions, their kindness will be appreciated. This Index is reprinted monthly on cards, and furnished in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitted ; each subscriber must take all cards published during the term of his subscription, Corre- spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club, Andrews, A. L. Sphagnaceae. N. Am. Fl. 15: 3-31. 14 Je 1913. Andrews, A. L. Sphagnales. N. Am. Fl. 15: 1. 14 Je 1913. Anthon, S. I. The bitterroot. Am. Bot. 19: 45-48. My 1913. Brown, H. B. Form and structure of certain plant hybrids in com- parison with the form and structure of their parents. Mississippi Agr. Exp. Sta. Technical Bull. 3: 3-54. f. I-33: Ja 1913. Brown, P. E. A study of bacteria at different depths in some typical Towa soils. Centralb. Bakt. Zweite Abt. 37: 497-52!- 22 My 1913. Blakeslee, A. F. A possible means of identifying the sex of (+) and (—) races in the mucors. Science II. 37: 880, 881. 6 Je 1913. [Illust.] Britton, E.G. Andreaeales. N. Am. Fl. 15: 33. 14 Je 1913. Britton, E.G. Archidiaceae. N. Am. Fl. 15: 45; 46. 14 Je 1913. Britton, E.G. Bruchiaceae. N Am. Fl. 15: 47-54. 14 Je 1913- : Britton, E.G. Bryoxiphiaceae. N. Am. Fl. 15: 69, 70. 14 Je 1913. Britton, E.G. Ditrichaceae.. N. Am. Fl. 15: 55-67-14 Je 1913. Britton, E.G. Seligeriaceae. N. Am. Fl. 15: 71-75- 14 Je 1913. Britton, E. G., & Emerson, J.T. Andreaeaceae. N. Am. FI. 15: a 39. 14 Je 1913. 457 458 INDEX TO AMERICAN BOTANICAL LITERATURE Britton, E. G., & Williams, R. S. Bryales. N. Am. FI. 15: 41-43. _ 14 Je 1913. Britton, N. L., & Brown, A. An illustrated flora of the northeastern United States, Canada and the British possessions 1: i-xxix + I- 680; 2: 1-735; 3: 1-637. New York. 1913. [Ed. 2.] Britton, N. L., & icc J. N. The genus Epiphyllum and its allies. Contr. U. S. Nat. Herb. 16: 255-262. pl. 78-84. 6 Je 1913. Includes Epiphyllum URED E. Nelsonii, E. pumilum, spp. nov.; Eccre- mocactus Bradei gen. et sp. , and Strophocactus gen. nov. Clute, W. N. A curious nila habitat. Am. Bot. 19: 49,50. My 1913. [Illust.] Clute, W. N. The origin of the Plum Island flora. Am. Bot. 19: 41- 44. My 1913. [Illust.] Clute, W. N. An ornamental garden plant. Am. Bot. 19: 61, 62. My 1913. [Illust.] [Clute, W. N.] The slender cliff-brake. Am. Bot. 19: 56, 57. My 1913. _ [Illust.] Clute, W. N. The spider flower. Am. Bot. 19: 53-55. My 1913. [Ilust.] [Clute, W. N.] Spring flowers of prairie woods. Am. Bot. 19: 58, 59- My 1913. [Illust.] Copeland, E. B. Notes on some Javan ferns. Philip. Jour. Sci. 8: (Bot.) 139-145. pl. 2-4. My 191 Includes Cyathea proegithat Athyrium pulcherrimum, A. subscabrum, and Polypodium javanicum, spp. ni Copeland, E. B. On Phollitis in Malaya and the supposed genera Diplora and Triphlebia. Philip. Jour. Sci. 8: (Bot.) 147-155. Pl. 5-7. My 1913. Dexter, J. S. Mosquitoes pollinating orchids. Science II. 37: 867- 6 Je 1913. Drennan, G. T. Narcissus biflorus. Am. Bot. 19: 51, 52. My 1913- Emerson, R. A. The possible origin of mutations in somatic cells. Am. Nat. 47: 375-377-. Je 1913. Hansen, J. Mosses of the vicinity of St. John’s University, College- ville, Stearns County, Minnesota. Bryologist 16: 42-45. f. I- My 1913. Hasse, H. E. The lichen flora of southern California. Contr. U. S. Nat. Herb. 17: 1-132 + vii—xii. 9 Je 1913. Heald, F. D. The dissemination of fungi causing disease. Trans. Am. Micr. Soc. 32: 5-29. Ja 1913. Heald, F. D., & Gardner, M. W. Preliminary note on the relative prevalence of pycnospores and ascospores of the chestnut-blight : fungus during the winter. Science II. 37: 916, 917- 13 Je i de INDEX TO AMERICAN BOTANICAL LITERATURE 459 Howe, R. H. Lichens of Mount Katahdin, Maine. Bryologist 16: 33-36. My 1913. Hedrick, U. P. A striking correlation in the peach. Science II. 37: 917, 918. 13 Je 1913. Hubbard, F. T. On Eragrostis cilianensis (All.) Vignolo Lutati. Philip. Jour. Sci. 8: (Bot.) 159-161. My 1913. Hull, E. D. Notes on the fern flora of Michigan. Am. Bot. 19: 48. My 1913. Jordan, W. H. Studies in plant nutrition. II. N. Y. Agr. Exp. Sta. Bull. 360: 53-77. f. 1-8. F 1913. Kaiser, G. B. Slime mould growing on a moss. Bryologist 16: 45. My 1913. Keith, S. C. Factors influencing the survival of bacteria at tempera- tures in the vicinity of the freezing point of water. Science II. 37: 877-879. 6 Je 1913. Merrill, G.K. Florida lichens. Bryologist 16: 39-41. f.z. My 1913. Millspaugh, C. F. The living flora of West Virginia. West Virginia Geol. Surv. 5 (A): 1-389, 454-486. 1913. [IIlust.] McDermott, F. A. A tetracarpellary walnut. Torreya 13: 137-139. f. I. 9 Je 1913. Maza, M. G. de la. Determinacion de plantas Cubanas. Fanero- gamas. 1-58-+ 1-4. Habana. 1913. Merrill, G. K. Lichens from Java. Torreya 13: 133-137. 9 Je 1913. Nelson, A., & Macbride, J. F. Western plant studies. I. Bot. Gaz. 55: 372-383. 15 My 1913 Includes 12 new species in pte Te (1), Clematis (1), Delphinium (1), Horkelia (1), Astragalus (1), Nemophila (1), Phacelia (1), Oreocarya (1), Castilleja (2), Pentstemon (1), and Erigeron (1). Nieuwland, J. A. Abnormal fruits of Juglans regia. Am. Bot. 19: 59, 60. My 1913. Northrup, Z. The influence of certain acid-destroying yeasts upon lactic bacteria. Centralb. Bakt. Zweite Abt. 37: 459-490. 22 My 1913. Prescott, A. Goldie’s shield fern. Am. Bot. 19: 63s 64. . My 1913. Nephrodium Goldieanum. Quehl, L. Mamillaria dolichocentra Lem. und ihre Verwandten. Monats. Kakteenk. 23: 69, 70. 15 My 1913. Purpus, J. A. Ariocarpus trigonus K. Schum. Monats. Kakteenk. 23: 65-69. 15 My 1913. [Illust.] Robinson, W. J. A taxonomic study of the Pteridophyta of the Hawaiian Islands. III. Bull. Torrey Club 40: 193-228. pl. 9-12 + f.z. 20 My 1913. Includes — of Polypodium pumilum, a kauaiense, and A, glabratum, spp. n 460 INDEX TO AMERICAN BOTANICAL LITERATURE Small, J. K. Flora of Miami. i-xii + 1-206. New York. 1913. ‘Descriptions of the seed-plants growing naturally on the Everglade keys and in the adjacent Everglades, southern peninsular Florida.”’ Small, J. K. Florida trees. A handbook of native and naturalized trees of Florida. i-ix + 1-107. New York. 1913. Stetson, S. The flora of Copake Falls, New York. Torreya 13: 12I- 133. f. I-4. 9 Je 1913. Swingle, W. T. Type of species in botanical taxonomy. Science II. 37: 864-867. 6 Je 1913. Tidestrom, J. Novitates florae utahensis. Proc. Biol. Soc. Washing- ton 26: 121, 122. 21 My 1913. Delphinium pinetorum, Eriogonum Kearneyi, Oreocarya Shanizii, and Mertensia Sampsonii, spp. nov Trelease, W. Agave in the West Indies. Mem. Nat. Acad. Sci. 11: 1-298. pl: I-1I6. 1913. Includes forty new species. Weingart, W. Cereus Boeckmannii Otto. Monats. Kakteenk. 23: 49, 50. 15 Ap 1913; 70-72. 15 My 1913. Weingart, W. Cereus lepidanthus Eichlam. Monats. Kakteenk. 23: 52. 15 Ap1913._ [Illust.] Weir, J. R. Destructive effects of Trametes Pini and Echinodontium tinctorum. Phytopathology 3: 142. Ap 1913. Weir, J. R. Some observations on Polyporus Berkeleyi. Phyto- pathology 3: 101-104. pl. 9. Ap 1913. White, D. The fossil flora of West Virginia. West Virginia Geol. Surv. 5 (A): 390-453, 488-491. 1913. White, D. A new fossil plant from the state of Bahia, Brazil. Am- Jour. Sci. IV. 35: sg Si f. 1-3. Je 1913. Alethopteris Branneri sp. n Williams, R. S. pap macrocarpum Card., in Florida and Funaria rubiginosa, sp. nov. Bryologist 16: 36-39. pl. 4. My 1913. Wolf, F. A. Abnormal roots of figs. Phytopathology 3: 115-118. pl. 11. Ap 1913. Woodward, R. W. Cyperus Grayii in Rhode Island. Rhodora 15: 100. 19 My 1913. Woodward, R. W. Extended range of some Connecticut plants. Rhodora 15: 94-96. 19 My Io1 Wylie, R.B. A long-stalked Elodea flower. Iowa State Univ. Bull. 6: 43-50. pl. 1, 2. 26 Ap 1913. BULL. TORREY CLUB ‘OLUME 40, PLATE FIGURE I FIGURE 2 HARPER: NORTHERN MISSISSIPPI BuLL_. ToRREY CLUB VOLUME 40, PLATE 22 FIGURE 3 FIGURE 4 HARPER: NORTHERN MISSISSIPPI \ Vol. 4C No. 9 BULLETIN OF THE TORREY BOTANICAL CLUB SEPTEMBER, 1913 Studies on the Rocky Mountain flora— XXIX \ ; Per AXEL RYDBERG *% MONOTROPACEAE . Hypopitys latisquama Rydb. sp. nov. Plant pink, 1-3 dm. high, more or less short-pubescent above; scales of the stem broadly ovate, obtuse, I-1.5 cm. long; flowers usually 10-15; sepals spatulate or cuneate, 8-10 mm. long, abruptly acuminate, ciliate; petals cunate or obovate, 11-12 mm. long, rounded‘and sinuate at the apex, pubescent and ciliate, fila- ments and style copiously hairy; stigma retrorsely bearded. This is closely related to H. lanulosa (Michx.) Nutt., but differs in the large and btoad scales on the stem and the larger flowers. MOonrtTANA: Bre Mountains, July 28, 1896. Flodman 708 (type, in herb. N. Y. 3ot. Gard.). . Wyominc: 1873, Parry 196. WasHINGTON: Olympic Mountains, Elmer 2464. \ PRIMULACEAE Primula specuicola Rydb. sp. nov. Perennial with a short »otstock; leaves 5-13 cm. long, thin, slightly farinose when young, in age glabrate, with winged petioles; blades spatulate or elliptic, obtuse\at the apex, sinuate-dentate; scape I-I.5 cm. long; umbels 10-20-flowered; bracts linear-subu- late, thin, 5-1o mm. lorg, slightly gibbous at the base; pedicels 5-10 mm. long in flower or 1-4 cm. long in fruit; calyx densely farinose; tube deeply campanulate, 3-5 mm. long; lobes linear- oblong, 2.5-3.5 mm. long, obtusish; corolla-tube yellowish, 8-10 [The ButueTin for August (40: 377-460. pl. 21, 22) was issued 13 Au 1913.] 461 462 RYDBERG: STUDIES ON THE ROCKY MOUNTAIN FLORA mm. long, 1.5 mm. in diameter; lobes cuneate, merely emarginate with .a broad sinus, dark violet, about 3 mm. long; stamens in- serted in the middle of the corolla-tube; capsule about 6 mm. long. This species is related to P. farinosa L. and P. incana M. E. Jones, but differs from both in its very thin leaves, more exserted corolla-tube and slender bracts. In P. incana M. E. Jones (P. americana Rydb.), the only other species of the group in the Rocky Mountains, the bracts’are thick, almost fleshy, obtusish, lanceolate, and often nearly equaling the pedicels. In the bracts and inflor- escence, it resembles more P. farinosa L. of Europe and north- eastern America. P. Ellisiae of the Sandea Mountains of New Mexico, though belonging to this group and of the same habit, has much larger flowers, the lobes of the corolla being 8-10 mm? long. P. specuicola grows only in loose soil, under overhanging cliffs in the alcove-like heads of the canyons, characteristic of the limestone bluffs of San Juan River. Urtau: Along San Juan River, near Bluffs, Aug. 25-29, I9II, Rydberg 9882 (type, in herb. N. Y. Bot. Gard.); same locality, Feb., 1912, Edna Scorup, and in 1895, Alice Eastwood. Androsace albertina Rydb. sp. nov. Cespitose perennial, but scarcely pulvinate; leaves narrowly oblanceolate, about 1 cm. long, sparingly ciliate, not carinate; scape 5-10 cm. long, slender, sparingly hairy; bracts linear- lanceolate, 3-4 mm. long; pedicels 3-5 mm.; calyx-lobes elliptic, obtuse; corolla-lobes 2-3 mm. long. This is most like the European A. Chamaejasme Host, but the leaves and bracts are narrower. It differs from A. carinata Torr. in the narrower leaves, not carinate' beneath, less pulvi- nate habit, longer peduncles, longer pedicels, and smaller flowers. ALBERTA: Lake Agnes, National Park, Banff, Aug. 1897, Mr. and Mrs. C. Van Brunt 77 (type, in herb. N. Y. Bot. Gard.); Jumping Pound Creek, June 14, 1897) Macoun 23478; Rocky Mountains 1858, Bourgeau. Montana: Yellow Mountain, jane 24, 1897, R. S. Williams. Androsace simplex Rydb. sp. nov. Annual; leaves oblanceolate, 3-6 mm. long, acute, entire, minutely puberulent; scape usually solitary, erect, very slender, 2-7 cm. high; bracts oval or lance-oval, 2-4 mm. long; pedicels RYDBERG: STUDIES ON THE RocKy MOUNTAIN FLORA 463 5-15 mm. long, suberect or strongly ascending; calyx-tube obpyramidal, about 2 mm. long; lobes lanceolate, about 1.5 mm. long, acute; corolla small, shorter than the calyx. This is related to A. occidentalis, but the plant is more delicate, the scapes solitary, bearing a 1-4-flowered umbel with ‘strongly ascending or nearly erect pedicels, the bracts smaller and dis- tinctly acute. Montana: Missoula, May, 1897, Elrod & assistants 33 (type, in herb. N. Y. Bot. Gard.). Utau: Near Salt Lake City, May 1882, M. E. Jones. British CoLtumsia: Lytton, April 17, 1889, Macoun. Dr. Greene separates an American species Androsace capil- laris Greene from the Asiatic A. filiformis Retz, and claims that the former is a perennial. All American specimens that I have seen are, however, annuals, and I can see no reason for such a separation. Dodecatheon Jaffreyi Moore has been collected near Sawtooth, Idaho, by Evermann. GENTIANACEAE Anthopogon ventricosum (Griseb.) Rydb. Gentiana ventricosa Griseb. in Hook. Fl. Bor.-Am. 2: 65. 1838. Anthopogon Macounii (Holm) Rydb. Gentiana Macounii Holm, Ottawa Nat. 15: 110, 179. 1901. . Anthopogon tonsum (Lunell) Rydb. sp. Nov. Gentiana detonsa tonsa Lunell, Bull. Leeds Herb.. 2:7. 1908. This is closely related to A. Macounii (Holm) Rydb., but differs in the glabrous filaments, a character not pointed out by Dr. Lunell. Amarella tortuosa (M. E. Jones) Rydb. Gentiana tortuosa M. E. Jones, Proc. Calif. Acad. II. 5: 707. 1895. Amarella ventorum Rydb. sp. nov. Gentiana arctophila densiflora Torr. Fremont’s Rep. 94. 1845. Not G. arctophila densiflora Griseb. ~ Low annual or biennial, branched near the base; stems 5—10 cm. long, branched, internodes shorter than the leaves; basal 464 RYDBERG: STUDIES ON THE Rocky MOUNTAIN FLORA leaves oblanceolate; stem-leaves linear or linear-lanceolate, about 2 cm., acute; flowers 1-3 in the axils; pedicels 2-8 mm. long; calyx-tube about 2 mm. long; lobes linear-lanceolate, 3-5 mm. long, acute, scabrous on the margins; corolla about 5 mm. long; lobes ovate, obtuse or acute; crown none. This little Amarella lacks the setaceous fimbriate crown at the base of the corolla-lobes and therefore should be classified with the arctic or subarctic A. propinqua (Richards.) Greene, and A. arctophila (Griseb.) Greene, but the corolla-lobes are acute or obtuse, instead of cuspidate. Wyominc: Wind River Moutainns, Aug. 4, 1843, Fremont. Dasystephana oregana (Engelm.) Rydb. Gentiana oregana Engelm.; A. Gray, Syn. Fl. 2!: 122. 1878. Dasystephana glauca (Pall.) Rydb. Gentiana glauca Pall. Fl. Ross. 2: 104. 1784. Dasystephana calycosa (Griseb.) Rydb. Gentiana calycosa Griseb. Gen. et Sp. Gent. 292. 1839. Dasystephana monticola Rydb. sp. nov. Gentiana calycosa stricta Griseb. Gen. et Sp. Gent. 292. 1839. Gentiana calycosa monticola Rydb. Bull. Torrey Club 24: 252. 1897. Dasystephana obtusiloba Rydb. sp. nov. Cespitose perennial; stems erect or ascending, about 1 dm. high; internodes short, equaling or a little longer than the leaves; leaves very broadly ovate, 3-5-ribbed, usually acute at the apex and subcordate at the base; calyx-tube broadly turbinate, 5-6 mm. long; lobes broadly oval, rounded at the apex, often over- lapping, about 8 mm. long; corolla dark blue, about 3.5 cm. long; lobes rounded at the apex; lobes of the plaits about half as long as the corolla lobes. This is related to D. calycosa, but differs in the lower habit and rounded corolla-lobes. MontTANna: Mary Baker Lake and Sperry Glacier, Aug. 21, 1901, Vreeland 1162 (type, in herb. N. Y. Bot. Gard.); Lake MacDonald, Aug. 22, 1901, Umbach 371; Mount MacDonald, July 25, 1900, Elrod & assistants; Silloway Peak, July 17-19, 1901, MacDougal 692; Blackfoot Indian Reservation, Aug. and Sept. 1909, Gilman Thompson. RYDBERG: STUDIES ON THE RocKy MOUNTAIN FLORA 465 Swertia Fritillaria Rydb. Glabrous, light green, perennial; stem 1.5-3 dm. high; basal leaves and lower stem-leaves alternate, 6-10 cm. long, thin, long- petioled; blades obovate, spatulate, rounded at the apex, abruptly contracted into winged petioles of about the same length; middle and upper stem-leaves all alternate or a single pair of opposite ones, oblanceolate or oblong; inflorescence rather lax, elongate; pedicels 1-2 cm. long; sepals lanceolate, about 6 mm. long; corolla-lobes lanceolate, mostly acute, greenish white along the midrib and azure along the margins, dotted all over with dark blue spots in the manner of many species of Fritillaria; filaments more or less dilated, some of them very broad; glands inconspicu- ous with rather long blue fringes. Uran: Wet places in caynons: Big Cottonwood Canyon, Au- gust 4, 1905, Garrett 1566 (type, in herb. N. Y. Bot. Gard.). APOCYNACEAE . Amsonia Eastwoodiana Rydb. sp. nov. Perennial, with a short woody base; stem 3-5 dm. high, gla- brous; stem-leaves lanceolate, usually narrowly so, 3-5 cm. long, glabrous, acute at each end; leaves of the numerous strongly ascending branches linear; calyx-lobes subulate, 2 mm. long or longer; corolla 16-20 mm. long; tube narrowly trumpet-shaped; lobes nearly 4 mm. long; pod 5-8 cm. long, about 8 mm. thick, constricted and often breaking off between the seeds, 3—5-seeded; seeds oblong, about 1 cm. long and 6 mm. thick. This is most closely related to A. brevifolia, having the same flower and fruit, but the plant is in habit more like A. Fremontii, for which it has been mistaken. The latter has still longer calyx-lobes which are narrower, and its pod is not restricted between the seeds. In canyons of desert regions. _ Uran: Moab, July, 1911, Rydberg & Garrett 8468 (fruit, type, in herb. N. Y. Bot. Gard.); Willow Creek Canyon, August, 1895, Alice Eastwood 73 (fruit). ARIZONA: Ten miles east of Holbrook, June 22, 1901, L. F. Ward (flowers); Lee’s Ferry, 1890, M. E. Jones. Amsonia texana (A. Gray) Heller of the Flora of Colorado and Coulter & Nelson’s Manual is A. latifolia Jones. i ae ! SY. On 4 "en, i Pg, meee — 4 oR | A Pin se chek mM Elep « neon. OF ius Dice OHIO m | rere! t, : | se “ice y Cawnence Fic. 2. Range of Oxydendrum arboreum in Ohio. the northeastern quarter of the state extending south to the present area along the lines of Merriam’s map. This may be illustrated by the distribution of Lycopodium in Ohio (FIG. 1). Almost as conspicuous is a second group which extends north from the Ohio River and occupies a triangular area with its apex at Sugar Grove. The sorrel-tree, Oxydendrum arboreum, is a oe example (FIG. 2). 492 GriIGGS: THE SUGAR GROVE FLORA Each of these when the whole range of the plants concerned is taken into consideration is found to be a composite of several types of distribution. These together with others belonging to types not so conspicuously homogeneous within the state may be classified as follows: A. ALLEGHENIAN PLANTS ON THE SOUTHWESTERN EDGES OF THEIR RANGES. Type range, BETULA LUTEA (FIG. 3). by Ca of yt <= J 4 BY ta (ae é a wee r ° a pa . a & — Oo Xo) ps, yo A s eg ye % I bY Fic. 3. Range of Betula lutea, This list includes beside the Alleghenian plants which go no further west than the Lake Superior region, some Canadian plants which stretch across the continent. Though these are more northerly and fewer of them reach Ohio, the ranges of those that we do have are so similar to the Allegheny type that they are inseparable. Capnoides sempervirens (FIG. 4) and Cornus canadensis, which terminates about twenty miles north of our area, are typical examples. Although this is a very homogeneous group of plants, conforming very closely to the typical range, many of them are also found in outlying stations far removed from the main range, as for example Blephariglottis lacera and Tsuga canadensis. There is also a tendency which may become more evident when more cdllections are available, for some of them to extend into south- western Ohio and southern {Indiana, e. g. the chestnut. This list includes 39 species as follows: Griccs: THE SUGAR GROVE FLORA 493 Fic. 4. Range of Capnoides sempervirens. Achroanthes unifolia Aronia nigra Aster macrophyllus Betula lutea Blephariglottis lacera Capnoides sempervirens Chimaphila maculata Chrysosplenium americanum Circaea alpina Cypripedium acaule Cypripedium reginae Epigaea repens Fraxinus nigra* Gaultheria procumbens Gentiana crinita Isotria verticillata Juncoides s altuensts Lycopodium clavatum Lycopodium complanatum Lycopodium lucidulum Lycopodium lucidulum poro- philum Lycopodium obscurum Lysias orbiculata Lysimachia quadrifolia Melampyrum lineare Panicularia elongata Panicularia pallida Parnassia caroliniana Polygonum arifoliumt Pyrola elliptica Pyrola rotundifolia : * The Ohio and Indiana (fide Coulter) distribution would indicate that this belongs in group F, but I follow Hough’s map and place it here. It is unknown south of Columbus. Tt Too widely extended in Indiana and Georgia to be typical. 494 GRIGGS: THE SUGAR GROVE FLORA Rubus odoratus Rynchospora glomerata Sambucus pubens Saxifraga virginiensis Trollius laxus Tsuga canadensis Unifolium canadense Viola rostrata. B. APPALACHIAN AND New ENGLAND SPECIES ON THE WESTERN EDGES OF THEIR RANGES (reaching Maine but not extending west of Lake Erie). Type range, SERICOCARPUS ASTEROIDES (FIG. 5). q é Fic. 5. Range of Sericocarpus asteroides. a K aw 72 ae * - tlw o) be —" % oi 0 Fic. 6. Range of Asplenium monitanum. The lines separating this group of plants from the preceding and following categories are somewhat arbitrary. There are many plants of evident boreal affinities which are now confined to the Appalachians. Their ranges form a continuous series between the typical Alleghenian, extending from Newfoundland to Lake Superior, down to those like Abies Fraseri and T suga carolipiana which are confined to a small area in the highest part of the southern mountains. Those which reach Ohio, however, seem to fall rather naturally into the two categories here listed. Those of the first group number 14 and include: Aster divaricatus Carex costellata Castanea dentata Dasystoma laevigata Eatonia nitida Mieracium paniculatum Hieracium venosum Kalmia latifolia Panicularia acutiflora Pinus rigida © Quercus Prinus Rhododendron maximum Sericocarpus asteroides Viola rotundifolia. Griccs: THE SUGAR GROVE FLORA 495 C. APPALACHIAN PLANTS (from southern New York or Con- necticut to Ohio and south through the mountains). Type range, ASPLENIUM MONTANUM (FIG. 6). These ranges are in some cases difficult to distinguish from those of the Carolinian plants because their northern boundaries nearly coincide and because of the tendency to spread through southern Ohio into Indiana toward the Ozarks. In such cases the general affinities of the plant have been the criterion for decision. Thus Aruncus Aruncus is placed here because the same or a closely related species is found on the Pacific coast to Alaska thereby clearly indicating its boreal affinities although its distribution in the eastern United States is apparently clearly Carolinian. We have 12 plants belonging to this category as follows: Aruncus Aruncus Phlox stolonifera Asplenium montanum Phacelia dubia Asplenium pinnatifidum Pinus virgimiana Azalea lutea Silene rotundifolia Cardamine rotundifolia Stachys cordata Oxydendrum arboreum Viola hirsutula. 0 CAROLINIAN PLANTS ON THE NORTHERN EDGES OF THEIR RANGES. Type range, PASSIFLORA LUTEA (FIG. 7). These are typically plants of southern or even subtropical Pe affinity whose northern limits are largely determined by lati- [3¥t--_'/~ wz| —- tude. As might be expected, there is no such uniformity in ~— the northern ranges of these t ley he plants as in those of the first group; Ilex opaca, Quercus mary- eee = landica, and Liguidamber Styraci- Fic. 7. Range of Passiflora lutea. jflua, though members of this group, just reach the southern extremity of Ohio and do not ~ come within 75 miles of Sugar Grove. The typical*members of this group extend straight across the country at about the latitude of Philadelphia, but there is a strong tendency in many Carolinian plants like Andropogon virginicus (FIG. 8) to extend up the coastal 496 Griccs: THE SuGAR GROVE FLORA plain through’ New Jersey to Long Island or even to Massachusetts. These are starred (*) on the list. The coastwise distribution t¢ of such plants finds its most striking exemplification in the occurrence Fic. 8. Range of Andropogon virginicus. of such a plant as Schizaea pusilla in Newfoundland.t The + Carolinianf'plants which terminate at Sugar Grove number 32 and include: *Andropogon virginicus Hydrangea arborescens Aralia spinosa Iris cristata *Aristida dichotoma Koellia incana§ *Asclepias variegata *Lechea racemulosa *Ascyrum multicaule Lobelia leptostachys§ *Betula nigra ; Lobelia puberula Blephariglottis paramoena Napaea dioica§ Carduus virginicus Panicum polyanthes *Cassia nictitans Panicum stipitatum§ Chrysopsis Mariana Passiflora lutea Cunila origanoides Porteranthus stipulatus§ Dentaria heterophylla Quercus minor Diospyros virginiana *Solidago erecta Eupatorium aromaticum Stylosanthes biflora Eupatorium coelestinum Trichostema dichotomum§ *Eupatorium rotundifolium Trifolium reflexum§ * Extending up the coastal plain into Long Island or New England. is interesting phenomenon has at <€ its bearing discussed by Hollick, Plant Distribution as a Factor in the Interpretation of Geological Phe- nomena with especial reference to Long Island and vicini rans. N. Y. Acad. t See Fernald, 1. c. § Not typical Carolinian plants. Of those marked thus, Koellia incana, Panicum Griccs: THE SUGAR GROVE FLORA 497 E. MISssISsIPPIAN PLANTS ON THE EASTERN EDGES OF THEIR RANGES, Type range, ISOPpYRUM BITERNATUM (FIG. 9). ae oe —— me Fic. 9. Range of Isopyrum biternatum. These are mostly plants characteristic of the great forest which once covered the Mississippi Valley and number 15 including: Aesculus octandra* Afzelia macrophylla Asclepias Sullivantii Bidens aristosa Brauneria purpurea* Dodecatheon Meadia* Fraxinus quadrangulata Hypericum Drummondii Isopyrum biternatum Psoralea Onobrychis Quamasia hyacinthina* Smilax ecirrhata* Sullivantia Sullivantii Valeriana pauciflora* Veratrum Woodit F. PLANTS ON THE SOUTHERN EDGES OF THEIR RANGES. Type range, SCUTELLARIA GALERICULATA (FIG. I0). This appears to be a miscellaneous aggregation without much similarity in range except that they are northern but not moun- tain plants. Probably further study and comparison would dis- cover common characteristics as conspicuous as in other groups. Some of them like Anemone canadensis are bounded by the Basin stipitatum, and Trichostema dichotomum have boundaries running from northeast to southwest instead of east and west while Napaea dioica reverses the case and is reported northwestward as far as Minnesota. Porteranthus stipulatus and Trifolium reflexum are transitional between this and the next group in that they do not cross the mountains but stop in western New York. Lobelia leptostachys also is not known much beyond the mountains and is likewise transitional to the next group. * Also known locally further east but the main body of the range stops in Central Ohio. 498 GricGs: THE SUGAR GROVE FLORA Others like Salix amygdaloides have a wide of the Great Lakes. distribution westward but taper eastward in a triangular area in western New York, thus conforming to Harsh- with its vertex egies SEER 7 » ) ae \ 4 f ~ Tee ot Os 4 bape ee \ =, By a) fs ye ro e e< = em (eee is ae ~TS az Dt TR Range of Scutellaria galericulata. Fic. tro. berger’s* map of-the Ohio-Tennessee area. 9 plants have been classed here as follows: Salix amygdaloides Anemone canadensis Cornus stolonifera Saxifraga pennsylvanica Dasyphora fruticosa Scutellaria galericulata Solidago juncea. Pedicularis lanceolata Populus tremuloides In addition to those given above there is one anomalous case which fits into no natural geographical range which I can discover. Viburnum dentatum comes down to our area from the northeast and meets Viburnum molle which comes up from the southwest. The characters which separate these species moreover do not hold in this region. It is evident therefore either that we do not under- stand these species and have only one of them even though there is a distinct variation from one part of the state to the other, or that they are not good species. The presence at one place of so many species on the edges of * Harshberger, J. W - Phytogeographic Survey of North America. Veg. der Erde 13: facing 790. IQIt. - Griccs: THE SUGAR GROVE FLORA 499 their ranges affords a favorable opportunity to study also their behavior. ‘Are they rare or abundant? Does their reproductive apparatus function normally? Do those plants on their northern _ edges behave differently from those on their southern? The eastern from the western? Is it possible to assign any reasons for the location of their termini here rather than fifty or a hundred miles beyond? These questions will be considered in a following paper. OHIO STATE UNIVERSITY, ” CoLumBus, OHIO. The culture of cereal rusts in the greenhouse F. D. FROMME The desirability of maintaining cultures of parasitic fungi on the living host in the greenhouse for purposes of study and physio- logical experimentation is obvious, particularly so with those obli- gate forms that cannot be cultivated on artificial media. Methods of culture of the powdery mildews of the grasses and other members of the Erysipl , and of a species of Cystopus, etc., Peronosporaceae, have been placed on an exact basis by the work of Reed (28, 29) and Melhus (23). No exact data of this nature, however, are available for the rusts. Although a vast number of infection experiments on this group have been made in recent years, these have dealt only incidentally with conditions governing spore germination, infection, and spore formation, and in many cases our knowledge on these points is based on field experiments con- ducted under conditions not subject to rigid control. There are a great many scattered details in the literature as to conditions affecting the development of the rusts. I shall summarize o those that bear more especially on the problems of growing rusts in the greenhouse. Smith (33) found that dew is of more importance in determining the prevalence of asparagus rust than rainfall. When little dew is formed infection cannot occur and sporulation may be checked, or altered, with a substitution of the teleuto for the uredo stage. This may occur in midsummer on the vigorously growing host as the result of excessive atmospheric dryness. He states, moreover, that abundance of soil moisture, instead of favoring rust develop- ment, acts as a check by giving the host greater vitality. This is in agreement with earlier observations by Stone and Smith (34, 35). They found asparagus beds on light, dry soil heavily rusted while those on heavy, moist soil were comparatively free from rust. Sirrine’s (32) observations also support the conclusion that dew is the most important agent in the spread of asparagus rust. 501 502 FROMME: THE CULTURE OF CEREAL RUSTS Morgenthaler (25) finds that mechanical injury of the leaves of the host favors teleuto as compared with uredo formation in the case of Uromyces Veratri. Tischler (36) found that shoots of Euphorbia Cyparissias infected with Uromyces Pisi became free from the rust mycelium when grown in a warm (25-27° C.) greenhouse. Many factors doubtless influence the germination of rust spores. They frequently germinate in a ‘few hours after being placed in water, as noted by de Bary. Often, however, though collected fresh, they fail to germinate for no apparent reason. This “capricious”’ germination has been noted by a large number of careful observers, and Eriksson, because of this uncertainty, does not consider the aecidiospores of the rusts of the cereals im- portant factors in their dissemination. Schaffnit (31) explains this ‘‘capricious’”’ germination on the ground that unless the spores are mature internally before detachment from their stalks they are incapable of germination. Complete maturity is at- tained only at a sufficiently high temperature (20°-25°) in an atmosphere calm enough to prevent their premature detachment. These conditions are not always realized in nature, hence the lack of uniformity in the results of germination tests. Freeman (13) and Klebahn (19) both have found that spores which germinate poorly in water may produce an abundant infection on the host, and therefore argue that germination tests are not conclusive unless conducted on the host. Klebahn states that aecidiospores of Peridermium Strobi germinated slightly or not at all in water, very vigorously on the Ribes host and less vigorously, but abundantly, on a decoction of Ribes. Sappin- Trouffy (30) likewise noted a marked difference in the germination of aecidiospores of Coleosporium Senecionis in water and in a decoction of Senecio vulgaris. Schaffnit (31) on the other hand, obtained no increase in host decoctions over the germination in water nor any effect attributable to a mechanical excitation by the substratum. Marshall Ward (37), was unable to find any effect of raw or cooked extracts of various bromes on the germina- tion of the uredospores of Puccinia dispersa. The effect of various chemicals on rust spore germination has been investigated by Wiithrich (40) and Carleton (6). Wiithrich FROMME: THE CULTURE OF CEREAL RUSTS 503 (see TABLE 1) has determined *the inhibiting action of different concentrations of various acids and salts on the germination of aecidio- and uredospores of P. graminis. Carleton finds that com- pounds containing Hg, Cu, Fe, Pb, Cr, and strong acids inhibit the germination of uredospores of Puccinia rubigo-vera, P. graminis and P. coronata and that those which contain O, Na, K, Mg, S, C, and NH; in large proportions are favorable to germination. The effect of temperature on spore germination has also received attention. Eriksson (9) found that aecidiospores of Aecidium Berberidis, A. Rhamni, and Peridermium Strobi and uredospores of Puccinia glumarum, P. graminis and P. coronata often germinated more readily at a few degrees below zero C. and on melting ice than at higher temperatures. Uredospores of P. dispersa according to Marshall Ward (37) germinate after freezing in ice for ten minutes. He attributes any increase in vigor obtained in this way to the drying action of the freezing and not to the low temperature. They germinated also at 27° but failed to do so at 30° and were killed at 65° to 70°. The optimum is near 20°. They germinate readily, if the spores are properly ripened and fresh and the temperature does not rise above 25°, in light, darkness, or red light, but less readily in blue light. Gibson (14) reports tests with uredospores of Puccinia Chrysanthemi as follows: Fifty per cent germination at 6°-6.5°, free germination between 7° and 21°, all at 21°-25°, one eighth at 24°-25°, and none at 30°. If kept dry at 35° for eighteen hours and then removed to 17° they germinate freely. Johnson (18) has determined the minimum, optimum, and maximum tempera- tures for germination of uredospores of the cereal rusts. These are: for Puccinia graminis on wheat, oats, and barley, 2° to 31°; for P. rubigo-vera on rye, 2°-30°; and for P. coronata on oats, 7°-8° to 30°. The optimum was determined by the general vigor of the germination tube and for all forms studied lies between 12° and 17°. This is somewhat lower than for P. dispersa as deter- mined by Marshall Ward. Johnson suggests that these low cardinal temperatures may explain the difficulty of obtaining infections in very warm greenhouses and on hot summer days and may account for the observation that rust epidemics are favored by subnormal temperatures at critical infection periods. 504°. FROMME: THE CULTURE OF CEREAL RUSTS The length of time during which uredospores of P. graminis retain their vitality was found by de Bary (3) to vary between one and two months, while the aecidiospores of the same form lost : their color and capacity for germination in one month. Marshall Ward (38) obtained germination with uredospores of P. dispersa after having kept them in a dry state for 61 days. Miss Gibson (14) reports a germination of 25 per cent. with uredospores of P. Chrysanthemi after storage of 71 days but none a week later. One aecidiospore of a sowing of Phragmidium Rosae-alpinae germinated after storage of 82 days. Barclay (2) found uredo- spores of some forms capable of germination during periods of from two to eight months (see TABLE 1) in the Himalayas. Bolley (5) obtained a 5 per cent germination with uredospores of P. graminis after exposure to air and sunlight during the month of August. Many observers have shown that various rusts are able to winter over in the uredo stage in some regions, but these data do not involve the determination of the actual time during which the spores are viable. Gibson (14) has shown that a large number of uredospores will germinate on the leaves of the wrong host and that their germ tubes will enter the stomata without, however, producing an infection. In these cases the end of the germ tube dries up in the substomatal chamber without further development. Because inoculation does not always result in infection, Marshall Ward (37) would distinguish sharply between these terms which are often used interchangeably. The passage of the germ tube into the host should be spoken of as inoculation and the subsequent development in the tissue of the host as infection. Pole Evans (rr) has investigated the entrance of the germ tubes of uredospores of P. graminis, P. glumarum, P. simplex, and P. coronifera into the stomata of their respective hosts and the establishment of the mycelium in the tissue of the host. The uredospores germinate within twenty-four hours and the infection is well established by the third day. When the germ tube reaches a stoma it forms a swelling or appressorium directly over it. A branch from the appressorium next enters the stomatal slit and forms a large vesicle in the substomatal chamber into which the contents of the appres- FROMME: THE CULTURE OF CEREAL RUSTS 505 sorium and germ tube are poured. One or more infecting hyphae are now sent off from the substomatal vesicle and these imme- diately establish connections with the surrounding host cells by means of haustoria. The early stages of development and the position and shape of the substomatal vesicle, and the number of infecting hyphae arising from it, are morphological distinctions between the different species. In the rusts the period between inoculation and sporulation is known as the incubation period. This apparently varies some- what with different species but the normal range for the uredo as reported by a number of authors lies between eight and twelve days. Marshall Ward has noted that the normal incubation period is shortened during clear sunny weather and Iwanoff (16) found that shading delayed aecidium-formation in P. graminis. What stimulus, or stimuli, determine the entrance of the germ tube into the stomata of the leaf has not been established. It is perhaps most generally held that the host exerts a chemical influ- ence on the germ tube. If this is true it is apparently not a specific influence, since germ tubes have been shown to enter the stomata of quite the wrong host. Massee (22) endeavored to demonstrate ‘a positive chemotropism but was unable to eliminate the effects of hydrotropism from the experiment. Marshall Ward (37) has noted an apparent heliotropic curvature in the germ tubes of P. dispersa. Balls (1) placed uredospores on a rubber film provided with small holes. Laboratory air was on one side of the film while the air on the other side was saturated with moisture at 23°. The germ tubes entered the holes and grew through into the region of higher pressure of water vapor. He believes that growth towards greater moisture will explain the entrange of the germ tube into the host. A recent article by Melhus (24) on the culture of parasitic . fungi deals with the culture of P. Helianthi, P. coronata, P. graminis and P. Sorght. The-methods employed by him consist in spraying the plants to be inoculated with a spore suspension, covering with a hood and placing in a refrigerator or humidity-box for twenty-four hours, at 14° for P. Helianthi, 16° for P. coronata, and 18° for P. Sorghi. He finds that P. coronata will not maintain itself - even though supplied with plenty of host material. It is more 506 FROMME: THE CULTURE OF CEREAL RUSTS difficult to hold in culture than the sunflower rust, a fact which is attributed to the slower growth of the oat plants. Reinoculation is necessary about every three or four weeks. P. Sorghi is more easily cultured than the cereal rusts and Melhus has propagated it both winter and summer. The amount of infection increases as the corn plants grow larger and the fungus spreads from one culture to another by natural agencies. I have tabulated the principal recorded observations on the influence of various conditions on spore germination and develop- - ment in the rusts in TABLE 1. These scattering and somewhat fragmentary records illustrate the incompleteness of our knowledge of rust physiology. CULTURE METHODS Preliminary experiments were made to determine how long single infections would maintain themselves under greenhouse conditions and whether they would self-propagate to any extent. The rusts used were Puccinia dispersa Erikss. on rye and Puccinia coronifera Kleb. on oats. The seedlings were grown in 5-inch pots and infections were secured by atomizing them with a uredo- spore suspension followed by covering with a bell jar for twenty- four hours. The incubation periods for both forms averaged about. twelve days in this preliminary work. The infections secured maintained themselves for about two weeks after the first ripening of the pustules. After this time the number of pustules visibly decreased through withering and dying off of infected leaves and eventually the cultures became entirely free from infection. Close association of non-infected with infected plants did not produce new infections. No marked difference in susceptibility between old and young plants was apparent but the younger were. found more desirable for inoculation because their greater compactness facilitated uniform covering with the spray. My further experiments were planned to meet various require- ments. First a method to maintain cultures of as nearly as pos- sible constant virulence, with the fewest necessary transfers, over extended periods of time, was tested out. Such a method is suited, for example, to test the possibility of maintaining the rust for long periods in the uredo stage and for the study of the effects of such conditions of growth on its virulence, incubation period, TABLE I on Ate? S. at. BD ee 1 re Spore Temperature Chemicals* Length of incu- ron ee om gts Moisture Host | Host Storage | Variation in =n Min. | Opt. | Max. Inhibiting |°**T8°¢ | ighe | | % Puccinia graminis........ PE SOO ROE occis 4 es 6 ars ee ek mga: Be) OE aR Enea er are eee I-2 months | Bi, putea eee 8 days 2—- hours in | cad | Puccinia graminis........ PORE Pike oa eee i ey “a I. OW be Cawubesek ys SWORE lo epee ated aan cakes 0.65 NasCOs | 0.1 2H2O4 0.0 CeHsOr _Puccinia graminis........ Il |Wiithrich 20-21° We WY re 8 acti rey hs ks he oe fs 6h Se ok l oe c ca ca bwen ee ts hours|o.r HCl | Puccinia graminis........ © IORI Nc ee cas cbeaalenters tee ei Sect pe beck he eG Agi. 1) Baek Soe mene een z 0.01 ZnSO, lengthens 0.0r ZnCl incubation o0.or CuSO, ME Peete | ice We wted ecu sae ooo blame ee ue os ewe ae Wen ee Puccinia graminis........ Be cE ONS ores eta cee gk wtlv ace as ry Evans weather favors elopment Puccinia graminis........ II |Eriksson | Low Eee nine ig RC Ge ae cles PCN eee a ioe Oy es oe Re Se cious” Puc inia coronata........ It [Eriksson Low “emperatures SOU er sec a VIG. Sere eee ea by k « Fabs} oo. SAU Se Ne rat peewee nde cious" Puccinia glumarum....... Il |\Eriksson bee sos stig sohin enh CO, wha cemmge = SSS Sane ie Gee Pamir eerie adres ra seat alee een enen accelerate cious’’|} 0.01 FeSO« 0.001 ZnSO, * All data as"to chemicals are Wiithrick's. SIsnd TVAaXHO JO AMOLTND AH] :aANWOUy LOS TABLE I—Continued Species of rust yay Author se he sv | : oat ine res ai Vatia aie nee oe Malstare Favor- Eanible sate cel oat seernee o light Min. | Opt. | Max | ing | Puccinia graminis........ Ec Wt ich cites | oes ev uke eee ae cs eG os RS heals Be hee ahs wo eae ae DOR Pd ee eam eG 0.0001 HgCle nia oo Bae as II |Johnson PSA Score og BR: 6 ik BREE Wi) Se SURRY Bape) Sei Game Me esr OMNES, We pee iene sme te WRAL acide PR Pace re Tae ewes II |Johnson Pee Ree SOC Bg ene ste ss co eb ee Oh Rae od oe oe ee ed oe te ou Dees Puccinia volian bss II |Johnson ye ge eRe ye SE RR Cie eS i RR RRR Ria US a emi GUN Me ACE MoM iy, Puccinia cco tbtalea ee is TE FC ArIObon Ciao seco ia eb eee Na 2 ev (eon tae, eae Ry Di waco Aen samy Men aects Se haar lal BM tant Oa Mai K Cu Puccinia graminis........ 5S: PION Poa care or. eee Mg PEE CL was CEES UN Eien Pohe ace DEL «cL eee Ss Pb Puccinia coronata........ Tae CARICOM Tp woes. emusued eee ee ee Ca Se have CaP Ente) see Man tes ede teed OVE el cen rran coon NHs | Alka- loids SESS II |Melhus 16° nee i RE Css inf eee ts she Sone a Pelee s Saad Vib ls ve eb odiccs el Soka daye Puccini Helionthi. II |Melhus ne Ble ag Cesc cua, 0 Re AMES FERONR Signi Muri Renae Se io Mees Ua Sea ne sae days Pi, Pactra peat. LLM De i ee Ma es 58s bg Bo cA cH le das a oo 8 days Pree at a ee Aer Pee entee Coe nee OMROMEIS COE ich octet bes echoes Pe dicv ier sew okalesccbesise eles bes ua. Puccinia rubigo-vera...... Be ONO M Mec w iin ate. Oper mt Ruane ei i plat ep TS EES MR ea By re RI) ee tn ag Puccinia dispersa........ IE. |Marshall |ro-12°| 20° |26-27°).....5...]...... CO:z |No in-|...... 61 days in dark 7-10 days Ward fluence nd in red or : blue light Pwemueemespersa........| Il (Freeman |......)......[...... ONCER DOOTL yeaa his os. |. oss is ae ca Worst y Pe cee Cel a ee RN Pe ae as in water euccemta AsPoragi.,.....| 1, 11 |Smith |....../...... es a [RM Ti hae es Lae eles foc eed te ocean Pickens fee ke oe dor ts kaa sential for]. infection 80¢ AWWOUJ SISNad Tvaaao AO AAALTIAD AH] TABLE I—Continued Gunicies wl-cuie nig are ae Temperature ___ Chemicals f ae a ieee of i ese ~~ Moisture ,, | Host | Host Storage Variation in 4 Favor- | Inhibit- | extracts light Min. Opt. Max. ing ing Phragmidium Rosae-alpinae) 1 |Gibson |...... EES, OR RES a: ea Mana | Germ OS GB YS Oh cerca: Pane Se ae tubes enter wrong host Puccinia Chrysanth EG ee aie oe aap ed te ad ee ce ne re aD ia Me ee! nee aa Mir: ae Puccinia Chrysanthemi II |Jacky We OWE Bi i i eh al oc tence hee ee dec due doetce hese. — 25 Phragmidium obtusum.. . II |Dietel 4- std ydrption OREEE eae AMPA. care WE eigisces BEM Ci lego sacs < « howe ore Liebe an Too bean ice and s _ Peridermium Strobi...... We aS CP aes oe oee cn Weatener iy orci ie... Fair | A- iB CCRS “iuscet ces take hes bee rss A, water bund : ant Peridermium Soraueri..... ce his sn cee yeah: eye a ed MO aye) He se at, ee pa ss Coleosporium Senecionis RI OG freee s kc eae vot) gat qt, ee ed Be PRES ERS a ah = ne ite DER Fete Ae al pag isan sar Trouffy none in bund- wat ant Urompess Pist 0065.5... ee rere ere mi EY gt ic, «ds cos hes ks lochs obec Ceshovn ee oe, moist air at 25-27 Ui IIE cose i se cs Il arclay )..2%.'. Se TSE a Ob RI COIN i RP, rR Re Rislee | Meee: FAAS Paces yumy pm ae As sche aie Wil Uredo Gomphrenatis . . I wham ca Rs) ots lee PD oe a one SSS le gs ee a eae ie eS PING. FAL rors pete aces Sse Prenanthis...... smb ich ti tear pac MRO OR as | CS RE a a ace Oa 7 mo CE MY CPeOU RIC Daan Ca aa a es Puccinia Caricis- Ao Ao gal Peer Eee bn © oe ae ere Vite oan AIT Wikies Sta wrh oe | scve'w ache ee oe Coie celle BPs RY (OR cgi Pt“ CMR SNR a Jromyces Vossiae........ I OMEN CU ema oceht Whe wernt. User teh... VIS ocks cell oawien L ip Sa, eats SNE Se Raat BBN | Reta ty alge Puccinia oa Saf ee ET ie eben wer Rig ls oo. cleaves sal oaaeed See, FP ai Ar ses eee cases. ess Ss Jromyces Pist.......... ON so lo Os ee ig at ri Pe, ol os cele ced catcdcues We TS AG, OO: Coe Re ee. oa be eo Melampsora Lini....... POY eriee eee les WD ESR a eT aN Ls cas oa ce ov clacee ocx 2 Witte TE te ae ei hes coc ee kd nia flosculosorum ven GO BOGEN Be IS AR NRE IC TEN RD © Oe a Se eee Une BMG! 0G. eee en cee ss Dee te SLSQUY TWHEAD JO AAALTND AH] :ANWOUT 510 FROMME: THE CULTURE OF CEREAL RUSTS etc. It was found that it was necessary to transfer the infection to new cultures once a month. Five ripe, oval pustules were selected for each inoculation. The spores from these were removed with a scalpel and immersed in 25 c.c. of water in the bottle of the atomizer. This was then shaken vigorously to secure uniform distribution of the spores before application and the culture was subsequently covered with a bell jar for twenty-four hours. ““Spring’’ rye and “ Kherson”’ oats were used exclusively. About twenty-five seeds were sown to a 5-inch pot. Six transfers at intervals of a month were made. At first seed for the next culture were sown two weeks prior to the date set for transfer, but a further simplification of method was secured by sowing a month prior to inoculation. Thus sqgwing and transfer could be made at the same time. Plants were a month old when inoculated and two months old when abandoned. Well-infected cultures on both oats and rye were maintained for six months in this way. No attempts were made to measure exactly the degree of infection secured but the pustules were seemingly as numerous at the end of the period as at the beginning. Germination tests of spores in drop cultures made at various times gave 50-75 per cent. germination in six to twelve hours. The _ incubation periods during the six months were quite uniform. Twelve days was the longest incubation period recorded and ten days the shortest. Cultures made in this way show relatively few sori. The method provides for maintenance of a rust culture but it does not provide an abundant supply of infected plants at all times. During the incubation period the pustules have disappeared from the old cultures and have not matured on the new ones. To have abundantly infected plants continuously available the following method was used. Transfers were made once a week instead of once a month. Since two weeks were required for complete ripen- ing of the pustules it was necessary to run alternate series of host cultures. One series was ready for transfer one week and the other series the week following. Seeds for subsequent cultures of each series were sown at the time of inoculation and the seedlings were thus two weeks old when the rust was transferred to them. Cultures of P. coronifera have been maintained for eight months FROMME: THE CULTURE OF CEREAL RUSTS 511 by this method and the fungus has gone through thirty-seven generations of the uredo stage with no decrease in. virulence. It was soon found that even more simple methods of inocula- tion are equally if not more efficacious than the application of spore suspensions with an atomizer. The plants to be inoculated are first thoroughly atomized with water. A well-infected culture pot bearing fully ripe pustules is then held in a horizontal position immédiately above them and given a vigorous shaking. The spores that fall from above are caught in the small drops of water provided by the spray. If the culture used is heavily infected the falling spores may be seen as yellow clouds. The inoculated plants are then covered for twenty-four hours. Four pots of seedlings can be inoculated simultaneously from the same culture and a uniform degree of infection secured on all four by placing them close together and holding the culture somewhat higher than for inoculating a single pot. It was found that spraying prior to inoculation could be dispensed with but the pustules secured were somewhat less numerous than when the spray was applied. To secure the best results the culture used for transfer must be heavily infected and the transfer made shortly after the ripening of the pustules. If transfer is delayed more than a week after sporula- tion begins, inoculation with a spore suspension must be resorted to. : Very numerous and uniformly distributed pustules were se- cured and maintained by this dry-spore method of inoculation. The approximate number of pustules per plant in a culture was determined in the following way. * For facility in counting the surface area of the pot it was divided into smaller areas with strips of cardboard. The number of plants and infected parts on each of the smaller areas was then easily ascertained. Ten plants were then taken at random from the culture, removed to the stage of a binocular, and the number of pustules on the infected parts determined. TABLE 1 shows the amount of infection on a typical culture. In this culture all of the first leaves were infected, 62.5 per cent of the second leaves, and 29.1 per cent of the sheaths. The average number of pus- tules on the first leaves of the ten selected plants was 574.4, on the second leaves, 22.9, and on the sheaths, 1.4. The small number of 512 FROMME: THE CULTURE OF CEREAL RUSTS pustules on the second leaves was due to the fact that only their tips were exposed at the time of inoculation. The number of pustules on the upper and lower surfaces of a leaf is sometimes equal and two pustules are often situated directly opposite each other. The average number of pustules per plant maintained in the mass culture experiments was about 200-300. The lowest average recorded for a culture was 161.6, the largest 598.7. The largest number of pustules counted on a single plant was 996 (plant 3, TABLE II). TABLE II. ANALYSIS OF INFECTION ON CULTURE 7C Treatment: Innoculated by dry spore method, covered with bell jar. Number of plants, 72. Date planted, January 1. Date innoculated, January 13. Date of sporulation, January 24. Number of parts Part of plants No. of parts infected Per cent. infected 72 First leaves ie 100 72 Second leaves 45 62.5 72 Leaf sheaths 2I 29.1 — NUMBER OF PUSTULES ON TEN PLANTS SELECTED AT RANDOM No. of ; First leaves Second leaves plant SIME MEE eee oe : Leaf Total, entire Lower surface | Upper surface Sane haa rend — 207 246 27 30 I 601 2 215 341 12 2I 5 594 3 506 441 18 31 o 996 4 444 521 7 II ms 986 5 234 336 4 5 0 579 6 225 249 0 o ts) 474 7 376 403 10 13 3 805 8 99 140 oO oO 2 241 9 53 TI5 I 3 ° 172 Bae) 189 314 I4 22 is) 539 Total 2638 3106 93 136 14 5987 Ave...... 263.8 310.6 9-3 13.6 1.4 598.7 Ave. 574.4 22.9 1.4 598.7 NORMAL DEVELOPMENT OF THE UREDOSORI OF P, CORONIFERA The period of incubation for the rust in the open greenhouse varied between eight and eleven days from October to December. Twelve days was the constant incubation period during December but this decreased to nine days in the latter part of January and FROMME: THE CULTURE OF CEREAL RUSTS 513 remained at nine days throughout February and March. The sequence of stages during a nine days incubation period may be divided into two periods. First, a period of vegetative develop- ment during which no evidence of infection*is seen. Second, a fruiting period during which the stages in the formation of the pustules are apparent. The first period occupies five days after inoculation and the first visible evidences of pustule formation become apparent on the sixth day. The leaves on this day have a faint mottled appearance which is due to the presence of small areas that are lighter in color than the surrounding leaf tissue. These areas are visible only by transmitted light. On the seventh day the light areas become more conspicuous and their boundaries more sharply defined. The areas become slightly swollen on the eighth day and a light orange color is apparent. During the next twenty-four hours the development of the pustules is rapid. They continue to swell until the epidermis of the leaf is ruptured by a longitudinal slit and the orange mass of spores is extruded. The mass of spores hangs together for a time but breaks apart on drying and falls from the leaf as separate spores or in small groups when the leaf is disturbed. OBSERVATIONS ON CONDITIONS AFFECTING SPORE GERMINATION, INFECTION, AND RATE OF DEVELOPMENT Effect of moisture It has been demonstrated a number of times during this culture work that a humid atmosphere provided by covering with a bell jar is necessary to secure infection. Abundant moisture may be supplied at the time of inoculation and still the plants will not become infected, in the greenhouse, unless they are covered soon afterwards. The drops of water apparently dry up before germi- nation and infection result. The per cent of saturation in the greenhouse in which my experiments were made as obtained from the hygrometer records, averages about 75~80 per cent with a temperature range between 55 per cent. and 85 per cent. Al- though the conidia of Erysiphe graminis infect the cereals spon- taneously under these conditions, the uredospores of P. coronifera will not do so. To test the possibility of providing conditions of humidity 514 FROMME: THE CULTURE OF CEREAL RUSTS under which the rust might become self-propagating by close association of cultures, a sash frame culture box, 3 ft. square, was made of five window sash. The humidity maintained here when cultures were growing in it was quite constant and averaged 93 per cent. with occasional fluctuations of 2-3 per cent. Even insucha humid atmosphere new infections occurred only sparingly, al- though cultures were sprayed and heavily inoculated. To obtain good infections it was necessary to cover the cultures with bell jars as in the open greenhouse. _ A direct comparison of the effects of covering and not covering in the culture box was obtained by a statistical study. Two pots of seedlings, each seven days old, were inoculated simul- taneously from the same culture, after which one was covered for twenty-four hours and the other was left uncovered in the culture box. During the remaining eight days of the incubation period they were exposed to equal conditions of humidity. The differ- ence in the degree of infection obtained on the two cultures was covered culture if that obtained on the covered is regarded as the normal. The difference in the degree of humidity to which the cultures were exposed for twenty-four hours after inoculation could not have been more than 7 per cent, as the average in the culture box was 93 per cent and the atmosphere under the bell jar was _ presumably saturated. It is rather striking that this difference of 7 per cent should have produced a difference in degree of infec- tion of 94 per cent. The spores that produced the 6 per cent normal infection on the non-covered culture probably germinated more rapidly or were more favorably located with reference to moisture than the bulk of the spores. Effect of temperature To test the effect of different temperatures on the degree and rate of development of P. corontfera, two cultures of the same age were inoculated simultaneously from the same stock culture. Immediately afterwards one was placed in the greenhouse ‘‘stove”’ * FROMME: THE CULTURE OF CEREAL RUSTS 815 where the temperature ranges between 20° and 30°.* The temperature here was quite constant for each twenty-four hours throughout the experiment. It did not fall below 25° from 10 A.M. to 5 P.M. nor rise above 20° from midnight to 6 A.M. The other culture was placed in the greenhouse where the tem- perature fluctuation was between 14.5° and 21°. Here the tem- perature remained quite constant at 16° for the greater part of the twenty-four hours. It reached 21° for a short period at noon and 14.5° at midnight. Both cultures were covered after inoculation. The first visible signs of infection became apparent on the culture in the ‘‘stove’’ on the fourth day after inoculation and fully ripe pustules were produced on the seventh day. Evidences of infection did not become visible on the culture in the greenhouse until the seventh day after inoculation, which was the date of sporulation for the culture in the ‘‘stove,” and ripe pustules were not formed until the twelfth day. The high temperature of the ‘‘stove,”’ which was an average increase of about 7-8°, was apparently responsible for a marked increase in the rate of development of the fungus and a decrease of five days in the incubation period. No differences in the degree of infection secured on the two cultures were apparent. The experiment was repeated immediately, but without a control, and the incubation period for this culture in the “stove” was but six days. This is the shortest incubation period I have observed. Further cultures gave results as follows. No.’7 in the ‘‘stove,’’ incubation period seven days. The temperature for each day was 20° from 6 P.M. to 6 A.M. and 25-30° from 8 A.M. to 2 P.M. Three controls in the greenhouse, incubation period nine days. Temperature 10-15.5° from 6 P.M. to 6 A.M. and 15.5-21° from 9 A.M. to 5 P.M. No. 8 in the ‘‘stove,’’ incubation period nine days. There were many minor fluctuations in this period. The maximum was 30°, the minimum 8°. The general range was distinctly lower than in the preceding test. Two controls in the greenhouse, incubation period twelve days. Temperature 10- 15.5° from 6 P.M. to 6 A.M. and 15.5-21° from 10 A.M. to 3 P.M. * The temperature records were obtained with Richard Fréres’ self-recording thermometers. 516 FROMME: THE CULTURE OF CEREAL RUSTS Effect of light To determine the effect of light exclusion on spore germination and rate of development, four culture pots of the same age, seven days, were inoculated simultaneously. Immediately afterwards one of the four was transferred to a physiological dark room which joins the greenhouse. A continuous circulation of air between the two rooms is maintained by an electric fan. Thus the average degree of humidity of the dark room, which is about 80 per cent., does not fall below that of the greenhouse although the range of fluctuation, which is from 60 per cent. to 95 per cent., is somewhat greater. The other three cultures were placed in the culture box as controls. The culture was exposed in the dark room for three days and at the end of this time was returned to the culture box. The plants at this time were quite as green and fresh as those of the control cultures and could not be distinguished from them. The incubation period for the three controls was eight days, while that of the culture left three days in the dark room was eleven days. At the time of sporulation some of the leaves of this culture showed signs of yellowing at their tips. No pustules were pro- duced on these discolored areas but on the normal green parts they were as numerous as on the controls. The difference of three days in the incubation periods is exactly equal to the period of light exclusion and indicates a complete arrest of the development of the fungus in the dark room. The effect of light exclusion during the latter part of the incubation period was also tested. Four cultures were inoculated and placed in the culture box. Four days later one of them was transferred to the dark room, where it was left four days, and then returned to the culture box. No signs of infection were visible on it at this time, while the unripe pustules on the controls were plainly visible. The pustules on the controls ripened on the ninth day, while three additional days, twelve days in all, were necessary for a similar development of the culture that had been in the dark room. By excluding light four days in the latter part of the normal incubation period, the maturation of the rust had been delayed three days. This shows that even after the fungus has become well established in the host its development is strongly retarded in complete darkness. FROMME: THE CULTURE OF CEREAL RUSTS 517 The average difference in degree of humidity, of approximately 13 per cent, between the culture box and the dark room could not have been an important modifying factor in these results, as I have found that the incubation period is not modified by relative degrees of humidity after the first twenty-four hours. Thus parallel sets of cultures, when grown in the culture box and in the greenhouse under similar conditions of light and temperature but with the same difference in humidity that is found between the culture box and the dark room, had the same incubation period. Likewise when a culture was kept under a bell jar during the entire incubation period, with occasional removal for change of air, there was no difference between its incubation period and that of the control in the greenhouse that was covered for the first twenty- four hours only. The question naturally arises how this retardation of the growth of the fungus is brought about. It may be the direct effect of total absence of light on the fungus itself. Then, again, it is possible that the fungus simply suffers from lack of food, since the host is incapable of assimilation in the darkness. It seems hardly pos- sible, however, that such a complete inhibition in the growth of the fungus should have resulted in the brief time involved unless it is dependent on the transition products in photosynthesis. This latter possibility is by no means inconceivable and should this explanation prove the correct one it could be made the basis for an explanation of the obligate parasitism of the rusts and their inability to develop on any form of artificial medium. VIABLE PERIOD OF UREDOSPORES Two series of tests were made to determine the period through which the uredospores of P. coronifera would retain their vitality. Ripe spores were removed from pustules on the cultures and placed in small gelatine capsules which were then stored in the laboratory at room temperature. The first set of spores was stored on March 13. A drop culture made on this date gave a high per cent of germination in twelve hours. Drop cultures of the stored spores were made on March 109, 23, and 28 and April 12 and 30. From 9 to 16 per cent. of the spores sown germinated in each of these tests. No further trials were made until June 1 and at this time 518 FROMME: THE CULTURE OF CEREAL RUSTS the spores were colorless and failed to germinate. The last germination obtained was after 48 days of storage and the spores had lost their capacity for germination at some time between 48 and 80 days. Another lot of spores were stored on November 26. These germinated in gradually decreasing per cents on January 8, 18, and February 2, 11, 18. The length of time required for the develop- ment of a germination tube increased from a period of 6-12 hours in November to a period of 36-48 hours in February. In the last drop culture made, on February 18, only 29 spores of 1,013 tested germinated. These few spores, about 0.2 per cent, had germinated after 84 days storage. . CONTROL OF MILDEW The powdery mildew, Erysiphe graminis, became so abundant on the oats cultures that measures for its control became necessary. The mildew spreads rapidly in the greenhouse and outgrows the rust to such an extent that rust culture work may be seriously interfered with. Attempts at exclusion of the mildew by isola- tion of the cultures and inoculation with uredospores selected from apparently non-mildewed leaves proved unavailing. The control and total exclusion of the mildew was achieved by treatment with sulphur dust. This was applied with a powder gun, twenty-four hours after rust inoculation. The plants were also atomized with a weak solution of sulphuric acid (1/1000) prior to the application of sulphur and were covered afterwards for twenty-four hours. With this treatment no mildew adie while control cultures became heavily infected. This work was undertaken at the suggestion of Prof. R. A. Harper, to whom I am indebted for suggestions and criticisms. SUMMARY 1. Two of the cereal rusts, Puccinia dispersa Erikss., on rye, and P. coronifera Kleb., on oats, have been cultured in the uredo stage, on the living hosts in the greenhouse, for a consecutive period of six months, from December 1912 to June 1913, by the transfer of infection once a month. P. coronifera was also cultured for a period of eight months, from September 1912 to May 1913, FROMME: THE CULTURE OF CEREAL RUSTS 519 with transfer of infection once a week. During this period the rust went through 37 generations of the uredo stage. No decrease in the degree of infection secured resulted from such continuous culture. 2. The average degree of infection maintained in mass cultures was approximately 200 pustules per, plant. The largest number of pustules counted on an individual plant was 996. 3. P. coronifera does not self-propagate to any extent even when abundant host material is supplied and a constant humidity of 93 per cent is maintained. 4. High humidity is the essential factor in securing successful inoculation with uredospores of P. coronifera. No infections resulted when cultures were exposed in an atmosphere of 75 to 80 per cent of humidity, and at 93 per cent only 6 per cent of the normal degree of infection was obtained. Normal infections were secured only when cultures were covered with a bell jar for twenty- four hours subsequent to the application of spores. 5. The rate of development of P. coronifera increased with tem- perature increase. A decrease in the normal incubation period of five days, or 41 per cent, was produced in the ‘‘stove” where the temperature ranged from 20° to 30° while the range at which the normal cultures were grown was 14.5° to 21°. 6. Total light exclusion either early or late in the incubation period checks the development of P. coronifera and results in an almost complete cessation of growth. 7. Uredospores of P. coronifera when stored at room tempera~ ture gradually lose their capacity for germination. A 0.2 per cent germination was obtained after storage of eighty-four days. COLUMBIA UNIVERSITY, NEW YORK. LITERATURE 1. Balls, W. L. Infection of plants by rust fungi. New Phytol. 4 18, 19. 1905. 2. Barclay, A. On the life-history of Puccinia coronata, var. hima- lensis. Trans. Linn. Soc. II. 3: 227-242. pl. 56. 1891. 3- Bary, A. de. Untersuchungen iiber Uredineen. Monats. Akad. Berlin 15-50. pl. 1. 1866. 4. Bary, A. de Neue Uiterescueees iiber Uredineen. Monats. Akad. Berlin 205-216. pl. 1. 1867. 520 FROMME: THE CULTURE OF CEREAL RUSTS 5. Bolley, H. L. Einige Bemerkungen iiber die symbiotische Myko- plasmatheorie bei dem Getreiderost. Centralbl. Bakt. 4: 887-896. 1898. 6. Carleton, M. A. Studies in the biology of the Uredineae. Bot. Gaz. 18: 447-456. pl. 37-39. 1893. 7. Dietel, P. Beitrage zur Morphologie und Biologie der Uredineen. Bot. Centralbl. 32: 246-249. pl. 3. 1887. . 8. Eriksson, J., & Henning, E. Die Hauptresultate einer neuen Un- tersuchung itiber die Getreideroste. Zeitschr. Pflanzenkrank. 4:66-73. 1894. 9. Hosp j. Uber die Férderung der Pilzsporenkeimung durch Kalte. Centralbl. Bakt. 12: 557-565. 1895. 1o. Eriksson, J. Die Getreideroste. Stockholm 1896. - 11. Evans, B. P. The cereal rusts. Ann. Bot. 21: 441-466. pl. 43. 1907. 12. Evans, B. P. South African cereal rusts. Jour. Agr. Sci. 4: 95- 104. IQII. 13. Freeman, E.M. Experiments on the brown rust of bromes (Pucci- . nia dispersa). Ann. Bot. 16: 487-494. 1902. 14. Gibson, C. M. Notes on infection experiments with various Ure- dineae. New Phytol. 3: 184-191. pl. 5-6. 1904. 15. Hitchcock, A. S., & Carleton, M. A. Kansas State Agr. Coll. Exp. Sta. Bull. 38. 1893. 16. Iwanoff, B. Untersuchungen iiber den Einfluss des Standortes auf den Entwickelungsgang und den Peridienbau der Uredineen. Centralbl. Bakt. 18?: 265-288, 470-480, 655-672. 1907. 17. Jacky, E. Der Chrysanthemum-Rost. Zeitschr. Pflanzenkrank. 10: 132-142. 1900. 18. Johnson, E. C. Cardinal temperatures for the germination of uredospores of cereal rusts. Abstract. Phytopathology 2: 47-48. I912. 19. Klebahn, H. Die wirtwechselnden Rostpilze. Berlin 1904 20. Maire, R. Recherches cytologiques et taxonomiques sur les Basidiomycétes. Bull. Soc. Mycol. 18: Append. 1-209. Pl. 1-8. 1902. 21. Maire, R. La biologie des Urédinales. Prog. Rei Bot. 4: 109- 163. = 108%, 22. Massee, G. On the origin of parasitism in fungi. Phil. Trans. 197: 7-23. 1904. 23. Melhus, I. E. Spore germination and infection in certain species of Oomycetes. Univ. Wisconsin Res, Bull. 15: 25-84. pl. I-10. 1gIt. 30. 6. w 37- 38. 39. 40. FROMME: THE CULTURE OF CEREAL RUSTS 521 - Melhus, I. E. Culturing of parasitic fungi on the living host. Phytopathology 2: 197-203. pl. 20. 1912. Morgenthaler, O. Uber die Bedingungen der Teleutosporenbildung bei den Uredineen. Centralbl. Bakt. 27?: 73-92. 1910. Plowright, C. B. British Uredineae and Ustilagineae. London 1889. . Pritchard, F. J. A preliminary report on the yearly origin and dissemination of Puccinia graminis. Bot. Gaz. 52: 169-192. . Reed, G. M. Infection experiments with Erysiphe graminis DC. Trans. Wisconsin Acad. Sci. 15: 135-162. 1906. Reed, G. M. Infection experiments with Erysiphe Cichoracearum DC. Univ. Wisconsin Bull. 250. Sc. ser. 3: 341-416. 1907. Sappin-Trouffy. Recherches histologiques sur la famille des Uré- dinées. Le Botaniste 5: 59-244. 1896. Schaffnit, E. Biologische Beobachtungen iiber die Keimfahigkeit und Keimung der Uredo- und Aecidiensporen der Getreideroste. Ann. Mycol. 7: 509-523. 1909 Sirrine, F. A. Spraying for asparagus rust. N. Y. Agr. Exp. Sta. Bull. 188: 234-276. 1900. Smith, R. E. The water relation of Puccinia Asparagi. . Bot. Gaz. 38: 19-43. 1904. Stone, G. E., & Smith, R.E. The asparagus rust in Massachu- setts. Matsichinewtis Agr. Coll. Bull. 61: 1-20. pl. 1,2. 1899. Stone, G. E., & Smith, R. E. Relationship existing between the asparagus rust and the physical properties of the soil. Massa- chusetts Agr. Coll. Exp. Sta. Ann. Rep. 12: 61-73. 1900 Tischler,G. Untersuchungen iiber die Beeinflussung der Euphorbia Cyparissias durch Uromyces Pisi. Flora 104: 1-64. I91I. Ward, H. M. On the relations between host and parasite in the bromes and their brown rust. Ann. Bot. 16: 233-315. 1902. Ward, H. M. Further observations on the brown rust of the bromes. Ann. Mycol. 1: 132-151. 1903. Ward, H. M. The histology of Uredo dispersa and the “ myco- plasm” hypothesis. Phil. Trans. 196: 29-46. pl. 4-6. 1903. Wiithrich, E. Uber die Einwirkung von Metallsalzen und Sauren auf die Keimfahigkeit der Sporen einiger der verbreitetsten parasitischen Pilze unserer Kulturpflanzen. Zeitschr. Pflan- zenkrank. 2: 81-94. 1892. INDEX TO AMERICAN BOTANICAL LITERATURE (1910-1913) aim of this Index is to include all current botanical literature written by Americans, published in oo or based upon American material ; the word Amer- ica being used in the broadest s Reviews, and papers that ick exclusively to forestry, eps! horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and no attempt is made to index the literature of bacteriology. An occasional exception is made in favor of some paper appearing in an American periodical which is devoted some important particular. If users of the Index will call the attention of the editor *o errors or omissions, their kindness will be appreciated. This Index is reprinted monthly on cards, and furnished in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitted ; each sana must take all cards published during the term of his subscription, Corre- ndence relating to the card issue should be addressed to the Treasurer of the Torrey Hee cal Club, Aaronsohn, A. The discovery of wild wheat and its possibilities for the United States. Chicago City Club Bull. 6: 167-175. 9 Je 1913. Arthur, J.C. Uredinales on Carex in North America. Mycologia 5: 240-244. Jl 1913. Bailey, W. W. November waifs. Am. Bot. 17: 98-100. N 1911. Banker, H. J. Type studies in the Hydnaceae—V. ores genus Hydnellum. Mycologia §: 194-205. Jl 1913- Includes descriptions of five new species. Barrett, O. W. The Philippine coconut industry. Philip. Bur. Agr. Bull. 25: 1-67. pl. 1-19 +f. I, 2. 1913. Includes considerable information of a botanical nature. Bean, R.C. Some Maine plants. Rhodora 15: 134, 135. 1 Jl 1913. Berger, A. Agave Warelliana. Curt. Bot. Mag. IV.9: pl. 8501. Je A Mexican can plant. ‘Bessey, C. E. Root punctured by root. Am. Bot. 17: 103. WN Ig1I. [Illust.' : Bessey, C. E. Some statistics as to the flowering plants. Science II. 38: 234, 235. 13 Au 1913. Bicknell, E. P. Viola obliqua Hill and other violets. Bull. Torrey Club 40: 261-270. 18 Je 1913. Bolley, H. L. The complexity of the microorganic population of the soil. Science II. 38: 48-50. 11 Jl 1913. 523 524 INDEX TO AMERICAN BOTANICAL LITERATURE Brainerd, E. Four hybrids of Viola pedatifida. Bull. Torrey Club 40: 249-260. pl. 15-17. 18 Je 191 iola papilionacea X pedatifida, V. pedatifida X sagittata, V. pedatifida X sororia, and V. nephrophylla X pedatifida, hyb. nov. Brainerd, E. Is Viola arenaria DC. indigenous to North America? Rhodora 15: 106-111. pl. 104. 11 Je 1913. Brainerd, E. Notes on new or rare violets of northeastern America. Rhodora 15: 112-115. 11 Je 1913. Britton, E. G. Wild plants needing protection. 8. ‘American or mountain laurel” (Kalmia latifolia L.). Jour. N. Y. Bot. Gard. 34: 121-123. pl. 117... Je 1913. Britton, N. L. Addison Brown. Jour. N. Y. Bot. Gard. 14: 119-121. Je 1913. [Illust.] Brown, S. Lophiola aurea Ker. Bartonia §: 1-5. Je 1913. tus Chamberlain, C. J. The oriental cycads in the field. Science II. 38 164-167. 1 Au 1913. Clark, E. D., & Smith, C. S. Toxicological studies on the mushrooms © Clitocybe illudens and Inocybe infida. Mycologia 5: 224-232. pl. gr. fl xon4. Clute, W. N. The smooth or meadow phlox. Am. Bot. 17: 97, 98. N 1911 ; Cockerell, T. D. A. The seedling of Phyllocarpus. Bot. Gaz. 55: 460. f. I. 16 Je 1913. Connell, A. B. Ecological studies on a northern Ontario sand plain. Forest. Quart. 13: 149-159. Je 1913. Crump, W.B. The coefficient of humidity: a new method of expressing the soil moisture. New Phytologist 12: 125-147. f. 1. 29 My 1913: Davis, B. M. A catalogue of the marine flora of Woods Hole and vicinity. U.S. Dept. Com. & Labor Fisheries Bull. 31: 795-833. 1913. Davis, B. M. General Cuaactectatics of the algal vegetation of Buzz- ards Bay and Vineyard Sound in the vicinity of Woods Hole. U.S. Dept. Com. & Labor Fisheries Bull. 31: 443-544. Charts 228-274. 1913. Dunn, S.T. The genus Marah. Kew Bull. Misc. Inf. 1913: 145-153- f. 1-5. My 191 Includes Marah micranthus and M. major from Lower California. Fairman, C. E. Notes on new species of fungi from various localities. a 5: 245-248. Jl 1913. udes descriptions of ten new species in Septoria (1), Sphaeropsis (2), Hender- sonia me is os (t), Pyrenochaeta (x), Coniothyrium (1), Diplodia (1) and Cry ptodiscu INDEX TO AMERICAN BOTANICAL LITERATURE 525 Fernald, M. L. An albino Kalmia angustifolia. Rhodora 15: 151, 152. Au 1913. Fernald, M. L., & Wiegand, K. M. Calamagrostis Pickeringit Gray var. debilis (Kearney), n. comb. Rhodora1s5: 135,136. 1 Jl 1913. Fernald, M. L., & Wiegand, K. M. Two new Carices from Newfound- land. Rhodora 15: 133, 134. 1 Jl 1913. Carex gracillima var. macerrima and C. lenticularis var. eucycla, var. nov. Field, E.C. Fungous diseases liable to be disseminated in shipments of sugar cane. U.S. Dept, Agr. Plant Ind. Circ. 126: 3-13. f. 1-7. 10 My 1913. Forbes, C. N. Anenumeration of Niihau plants. Occasional Papers, Bishop Museum 53: 17-26. f. I-4.. 1913. Includes Euphorbia Stokesit sp. nov. Forbes, C. N. Notes on the flora of Kahoolawe and Molokini. Occa- sional Papers, Bishop Museum 5%: 3-16. f. I-7. 1913 Fraser, W. P. Further cultures of heteroecious rusts. . Mycologia 5: 233-239. Jl 1913. ’ Fuller, G. D. Reproduction by layering in the black spruce. . Bot. © Gaz. 55: 452-457. f. 1-6.. 16 Je 1913. : Garman, H. The woody pints of Kentucky. — eee Exp. Sta. Bull. 169: 3-62. f. z-20. 1 Ja 1913. Gerste, A. Notes sur la médecine et la botanique des anciens Mexi- — cains. 1-191. Rome. 1910.. [Ed. 2.] Graves, A. H. A case of abnormal development of a short growth in Pinus excelsa.: Torreya 13: 156-158. f. 1. 8 Jl 1913. Harper, R.M. Economic botany of Alabama. Part I. Geographical seh Geol. Surv. ete Monograph 8: 1-222. f. ‘I-63. Je 19 Hayes, . XK, East, E. M., & Beinhart, E. G. Tobacco breeding in Connecticut. Connecticut Agr. Exp. Sta. Bull. 176: 5-68. pl. 1-12. My 1913 Heller, A. As Acmispon in California. aerate 9: 60-65. 30 Je 1913 Includes ‘aici gracilis, A. sparsiflorus, A. aestivalis, and A. glabratus, spp.~ no Holden, W.P. The fixation of nitrogen in Colorado soils. Colorado. Agr. Exp. Sta. Bull. 186: 3-47... My 1913. Holway, E. W. D. North American Uredineae 1: 81-95. us 37-44. It Je 1913. Includes Puccinia poromera, P. Pseudocymopteri and P. Cynomarathri, spp. nov. Humphrey, L. E. The.genus Fraxinus in Ohio. Ohio Nat. 13: 185- 188. Je 1913. 526 INDEX TO AMERICAN BOTANICAL LITERATURE Jones, D.H. A morphological and cultural study of some azotobacter. Centralb. Bakt. Zweite Abt. 38: 14-25. pl. 1-4. 21 Je 1913 Kellerman, K. F., & Leonard, L. T. The prevalence of Bacillus radicicola in soil. Science II. 38: 95-98. 18 Jl 1913. Knowlton, C. H., and others. Reports on the flora of the Boston district—XVII. Rhodora 15: 122-132. 1 Jl 1913. Knudson, L. Observations on inception, season, and duration of cambium development in the American larch. [Larix laricina (Du Roi) Koch.] Bull. Torrey Club 40: 271-293. pl. 18, 19. 18 Je 1913. Kunkel, O. The production of a promycelium by the aecidiospores of Caeoma nitens Burrill. Bull. Torrey Club 40: 361-366. f. I. 18 Jl 1913. Lamb, W. H. A key to common Nebraska shrubs. Univ. Nebraska Forest Club. Ann. §: [1-7]. 1913. Reprinted without pagination. Land, W. J. G. Vegetative reproduction in an Ephedra. Bot. Gaz. 55: 439-445. f. 1-5. 16 Je 1913. Lemoine, [M.] Mélobésiées. Revision des Mélobésiées antarctiques. In Charcot, J., Deuxiéme expédition —— francoise (1908- 1910). 1-67. pl. 1, 2. 1913. Lipman, C. B., & Wilson, F. H. Toxic inorganic salts and acids as affecting plant growth. (Preliminary communication.) Bot. Gaz. 55: 409-420. 16 Je 1913. Long, B. Southerly range extensions in Antennaria. Rhodora 15: 117-122; t Jl 1913. Maxon, W. R. Saffordia, a new genus of ferns from Peru. Smith- sonian Misc. Coll. 614: 1-5. pl. r, 2 +f. 1. 26 My 1913. Includes Saffordia induta sp. nov Maza, M.G.dela. Determinaci6n de plantas cubanas (fanerégamas). [1], 2, [3-9]. Habana. 1912. Melchers, L. E. The mosaic disease of the tomato and related plants. Ohio Nat. 13: 149-173. pl. 7, 8 + f. 1. Je 1913. Morse, W. J. Powdery scab of potatoes in the United States. Science I]. 38: 61, 62. 11 Jl 1913. Murrill,W.A. The Agaricaceae of the Pacific coast—IV. New species of Clitocybe and Melanoleuca. Mycologia 5: cates Ji 1913. Includes 21 new species in Clitocybe and 25 in Melanole Norton, A. H. Plants from islands and coast of aan, Rhodora 15: 117-136. 1 Jl 1913. Norton, A. H. Some noteworthy plants from the islands and coast of Maine. Rhodora 15: 137-143. Au 1913. INDEX TO AMERICAN BOTANICAL LITERATURE 527 Orton, W. A. Potato-tuber diseases. U. S. Dept. Agr. Farmers’ Bull. 544: 3-16. f. 1-16. 25 Je 1913. Osterhout, G. E. Concerning some species of Agoseris in Colorado. Muhlenbergia 9: 65, 66. 30 Je 1913. Osterhout, W. J. V. Protoplasmic contractions resembling plasmo- lysis which are caused by pure distilled water. Bot. Gaz. 55: 446- 451. f. 1-6. 16 Je 1913. Parish, S. B. Additions to the known flora of southern California. Muhlenbergia 9: 57-59. f. 2. 30 Je 1913. Reddick, D. The diseases of the violet. Trans. Massachusetts Hort. Soc. 1913: 85-102. pl. 1, 2. 1913. Reed, H. S., & Cooley, J.S. The transpiration of apple leaves infected with Gymnosporangium. Bot. Gaz. 55: 421-430. f. I. 16 Je 1913. Rigg, G. B. The distribution of Macrocystis pyrifera along the Ameri- can shore of the Strait of Juan de Fuca. Torreya 13: 158, 159. 8 Jl 1913. Rodman, R. S. Hieracium florentinum at Wellesley Hills, Massachu- setts. Rhodora 15: 116. 11 Je 1913. Rogers, S. S. The culture of tomatoes in California, with special reference to their diseases. Univ. Calif. Agr. Exp. Sta. Bull. 239: 591-617. f. 1-13. Je 1913. Roland-Gosselin, R. Are the species of Rhipsalis discovered in Africa indigenous? Torreya 13: 151-156. 8 Jl 1913. From Bull. Soc. Bot. France 59: 97-102. 1912. A translation by E. G. Britton. [Rolfe, R. A.] New orchids: decade 40. Kew Bull. Misc. Inf. 1913: 141-145. My 1913. Includes Sielis barbata, Cycnoches Ci —. Sage, J.H. Arenaria curotiniata i in Rhode Island. Rhodora 15: 115. II Je 1913. Schindler, A. K. Einige Bemerkungen iiber Lespedeza Michx. und ihre nachsten Verwandten. Bot. Jahrb. 49: 570-658. 17 Je 1913. Seaver, F. J. Some tropical cup-fungi. Mycologia 5: 185-193. i. 88-90. Jl 1913. Selby, A. D. Disease susceptibility of apple varieties in Ohio. Ohio Agr. Exp. Sta. Circ. 113: 53-56. 15 Ap 1913- Shear, C. L., & Stevens, N. E. Cultural characters of the chestnut- blight and its near relatives. U. S. Dept. Agr. Plant Ind. Circ. 131: 3-18. 5 Jl 1913. Small, J. K. Flora of the southeastern United States. i-xii+1- -1394. New York. 1913: [Ed. 2.] tdi: f America 528 INDEX TO AMERICAN BOTANICAL LITERATURE Smith, J. D. Undescribed plants from Guatemala and other Central American Republics——XXXVI. Bot. Gaz. 55: 431-438. 16 Je 1913. Includes descriptions of 12 new species in Rheedia (1), Caryocar (1), Maytenus (1), Meliosma (1), Phyllocarpus (1), Serif he Gilibertia (2), Basanacantha (1), Perymenium (1), Arctostaphylos (x), and Cor z. Smith, H. H. Thomas Howell. "he it 55: 458-460. 16 Je 1913. [Ilust.] Spaulding, P. The present status of the white-pine blister rust. U. S. Dept. Agr. Plant. Ind. Circ. 129: 9-20. f. r-6. 7 Je 1913. . Spegazzini, C. Contribucién al estudio de las Laboulbeniomicetas argentinas. An. Mus. Nac. Hist. Nat. Buenos Aires 23: 167-244. f. 1-97... 19%3. Describes three new genera and twenty-five new species. Spegazzini, C. Mycetes argentinenses. VI. An. Mus. Nac. Hist. Nat. Buenos Aires 23: 1-146. f. 76-99. 1912. Describes a large number of new species. Sprague, T. A. Hypericum aureum. Curt. Bot. Mag. IV. 9: pl. 8498. Je 1913. A plant from southeastern United States. Standley, P.C. Five new plants from New Mexico. Proc. Biol. Soc. Washington 26: 115-120. 21 My 1913 ‘ Nuttallia Springeri, easeamgar 7 australis, pense arenaria, A. hirtella, and Chrysothamnus elatior, spp. n Stapf, O. Amelanchier Miscsgk Curt. Bot. Mag. IV. 9: pl. 8499. Je 1913. A North American plant. Stewart, A. Expedition of the California Academy of Sciences to the Galapagos Islands, 1905-1906. VII. Notes on the lichens of the Galapagos Islands. Proc. California Acad. Sci. IV. 1: 431-446. 17 D 1912. Sturgis, W. C. On Stemonitis nigrescens and related forms. Bot. Gaz. 55: 400, 401. 15 My 1913. Swingle,W.T. The present status of date culture in the southwestern states. U.S. Dept. Agr. Plant Ind. Circ. 129: 3-7: . 7 Je 1913: Williams, A. Caryophyllaceae of Ohio. Ohio Nat. 13: 176-184. Je 1913. Wolden, B.O. Asters. Am. Bot. 17: 100-102. N 19It. Wolden, B. O. Taste of poison ivy. Am. Bot. 17: 102. N 1911. Woodward, R. W. Juncus dichotomus in Rhode Island. Rhodora 15: 15:. Au rgt3. Vol. 40 No. 10 BULLETIN OF THE TORREY BOTANICAL CLUB ee ree OCTOBER, 1913 Notes on Carex—VII KENNETH K. MACKENZIE CAREX UMBELLATA AND ITS ALLIES A study of a number of recent collections, chiefly from the western part of the United States, has brought about the con- clusion that in addition to Carex umbellata Schk. and Carex deflexa Hornem. and their allies, there are a considerable number of ad- ditional species belonging to the group Montanae, characterized | by the development of pistillate spikes on subradical peduncles or on very short culms. A natural division of these species can be made from certain characters taken from the perigynia. While all of the Montanae have the perigynia strongly 2-ribbed or 2-keeled, certain species of the group now under consideration found in California also have the outer face finely many-nerved. In all other members of the group the perigynia are nerveless or at most have on one face a few partly developed nerves on the lower half of the body. Next to be separated are two closely related species of the southeastern United States (Carex nigro-marginata and Carex floridana). These are to be distinguished by the fact that while the spikes are on very short culms and may appear basal they are not on basal peduncles. Each culm really bears one to several pistillate spikes sessile at the base of the staminate spike; and elongated radical peduncles bearing only pistillate spikes are not ‘normally developed, as in the remaining species After making this division probably the most satisfactory’ and [The ButteTIN for September (40: 461-528) was issued 10 S 1913-] 529 530 MACKENZIE: NOTES ON CAREX natural method of telling C. déflexa and its allies from C. umbellata and its allies is that there are always pistillate spikes at the base of the staminate in the former, and the lowermost of these has a leaflet-like bract exceeding the inflorescence. This bract is rarely auriculate, and if colored at all at base is purplish-brown tinged. In the other group all the pistillate spikes are frequently subradical, but when there is a pistillate spike at the base of the staminate its bract is squamiform and shorter than the inflorescence, or else, if rarely longer, it is auriculate and strongly reddish-tinged at base. Carex deflexa has a close ally in C. Rossii, the two species having been often confused. However, in C. deflexa itself the rootstock is very slender, horizontally creeping and short-stoloniferous; the perigynium is very small (about 2.5 mm. long) and strongly exceeds the scales; and the beak is short and but inconspicuously bidentate. In C. Rossi, as stated by Dr. Holm (Am. Journ. Sci. IV. 16: 37. 1903), the rootstock is ‘much more robust, relatively shorter and ascending, not horizontal and not stoloniferous in the stricter sense of the word.” This last species, too, has larger, longer-beaked and conspicuously bidentate perigynia, with relatively longer scales. Carex brevipes W. Boott, a little-known species of the far western mountains, seems to be valid. It has the habit of growth of C. Rossti, but the short-beaked, narrow perigynia are about as small as those of genuine C. deflexa. A very interesting collection made by Mr. W. W. Eggleston in the northern part of New Mexico in IQII, contains two new species belonging to the group of species now under discussion. One of these is related to Carex deflexa and its allies. It differs, however, in having few-flowered pistillate spikes even on fully developed plants. The beak of the perigynium is obliquely cut at apex and becomes slightly bidentate in age, and the margins are scarcely serrulate. Another plant collected b? him, now named Carex geophila, differs from the real Carex umbellata in having a globose perigynium body fully 1.75 mm. or more wide; the beak is not strongly 2-edged. The eastern species grouped with Carex umbellata have the perigynium body triangular-globose and but I-1.5 mm. wide, while the beak is strongly 2-edged Another species most closely related to Carex geophila occurs MACKENZIE: NOTES ON CAREX 531 along the Pacific coast from San Francisco to Vancouver Island. It furnishes the basis for the reports of the occurrence of Carex umbellata on the Pacific coast, and specimens of it are probably also responsible for the large perigynia at times attributed to that species. This plant, which I am calling Carex brevicaulis, has pistillate spikes maturing 1-4 strongly pubescent perigynia and the culms are little fibrillose at base. Carex geophila on the other ° hand has pistillate spikes maturing some 5-15 puberulent peri- gynia and the culms are extremely fibrillose at base. Several distinct species have been referred to Carex umbellata Schk. The genuine plant is distinguished by its deep green narrow erect leaves, the old reaching an extreme width of 2.5 mm., and the width of the young ones at fruiting time averaging about 1.5— 1.75 mm. The cuspidate pistillate scales are normally lanceolate or narrowly ovate-lanceolate, and usually do not entirely conceal © the lower part of the perigynia. The long-beaked perigynia are 3 mm. long or more and short-pubescent, and the beak is markedly bidentate. In parts of the country, especially in the north and west, the place of Carex umbellata is often taken by a closely related species, the perigynia of which are smaller (3 mm. long or less) and have a short (0.5 mm. long) shallowly bidentate beak. The scales are ovate and vary from acutish to cuspidate, and in the basal spikes tend strongly to conceal the lower part of the perigynia. This plant is the var. brevirostris Boott; the C. abdita of Mr. Bicknell. My own experience is that this species occurs in limestone districts and that Carex umbellata is a species not found in such districts. A closely related species ranging from Missouri to Texas has also been separated. Thoroughly distinct is the coastal plant described by Professor Fernald as var. tonsa (Proc. Am. Acad. 37: 507. 1902), and by Mr. Bicknell raised to specific rank (Bull. Torrey Club 35: 492. 1908). While the long glabrous perigynia make it well marked, the wide stiff spreading leaves referred to by Mr. Bicknell are even more characteristic, and in the field distinguish the plant at a distance from Carex umbellata when growing with it. Carex tonsa is an abundant ea in parts of the pine barren country of New Jersey. 532 MACKENZIE: NOTES ON CAREX The main distinctions between those members of the Montanae which are here discussed and which differ from all the other members of the group in having the pistillate spikes or some of them on short culms or on basal peduncles and so hidden among the leaves, — be vianges eget as follows: 5 ae i ly many trongl sees Kalen putblihstioned, eae to cuspidate; perigyni body globose; staminate spikes many-flowered; ae pistillate spikes on elongated, very slender peduncles. 1. C. globosa. Scales reddish-brown tinged, cuspidate or long-awned; enn ium body oval; staminate spikes few-flowered; basal pistillate spikes on short erect peduncles. 2. C. Brainerdii. II. Perigynia strongly 2-keeled, otherwise nerveless (except t times obscurely coarsely nerved at bas @). A. secre and staminate spikes closely contiguous (the ms often very short). Pera normally triangular; scales strongly dark- Pig aay stolons short, ascending; culms strongly fibrillose Achenes meals nie ate abe sept 3 at eet one a tely dark little fibrillose at base. B. Lower pistillate spikes widely separate (on subradical peduncles). 3. C. nigro-marginata. 4. C. floridana. Bract of sowent non-basal pistillate spike leaflet-like, exceeding culm; if at all colored, purplish-brown siiad at base. Rootstock slender; culms Saga loosely ces- pitose (perigynia 2.5 mm. long, short-beaked, shallowly bidentate; staminate spike 2. o mm. long, inconspicuous); northeastern species. . C. deflexa. Roots nas stout; culms de nsely satis species. vines acre spikes normally several- to y-flowered; perigynium beak biden- : ae the margins ciliate-serrulate. Perigynia 2.5-3 mm. long, the beak 0.25— 0.75 mm. long, pete bidentate. 6. C. brevipes. sietensi? Sie 5 mm. long, the beak longer, biden q. -C. Rosstt. Upper sacs spikes 1-3-flowered; _peri- gynium beak obliquely cut, in age biden- tulate; the margins little if at all ciliate- serrulate (New Mexico). 8. C. pityophila. Bract : lowest non-basal A — if pres- , squamifor er than culm; or, Mi lone. srt aise paket reddish-brown tinged a MACKENZIE: NOTES ON CAREX 533 jeighog body globose, 1. ‘ a mm. or more wide; not strongly 2-ed tees spikes with si 5 geo peri- gynia; culms extremely fibrillose at base Mexico). 9. C. geophila. Pistillate spikes with 1-4 strongly pubesc perigynia; culms little fibrillose at oat (Pacific slope). 10. C. brevicaulis, oe body Baa iain nt I-I.5 mm. ; beak hits 2-edge Ma 2.5-4 . long, gir ie leaf- blades slender, light green, ascending, 5 Short-stoloniferous, the new shoots phyl- lopodic, the sheaths eis | filamentose (perigynium beak very short; achen roughened). 11. C. microrhyncha. Densely cespitose, the new shoots aphyl- lopodic, erect-ascending, the sheaths filamentose. Perigynia 2.5-3 mm. long, the beak less than es length of body; h, shining, minutely oe piaoretse oid. 12. C. abdita. Perigynia 3.25-4.25 mm. long, the nutely roughened, oblong-obovoid. 13. C. umbellata. Perigynia 3.5-4.5 mm. long, glabrous except long beak; leaf blades stiff, deep green, spreading, 2.5—4 mm. wide. 14, C. tonsa. I. CAREX GLOBOSA Boott, Proc. Linn. Soc. 1: 259. 1845 Carex umbellata var. globosa (Boott) Kiikenth. in Engler, Pflan- zenreich 47°: 453. 1909- Clumps medium-sized, from ‘elongated slender rootstocks, stoloniferous, the culms 15-35 cm. high, phyllopodic, slender, exceeding leaves, roughened on angles above, strongly fibrillose and more or less reddened at base. Leaves with well-developed blades five to eight to a fertile culm, the lowest clustered, the upper more or less widely separate, the blades flat with revolute margins, I.5-2.5 mm. wide, varying in length from 3 cm. to 3 dm. (on sterile culms), attenuate-pointed, strongly roughened; terminal spike staminate, erect, short-peduncled, 1-2 cm. long, 2.5-4 mm. wide, many-flowered, the scales oblong-obovate, obtuse or acute, purplish brown with lighter center and hyaline margins; pistillate spikes two or three (with additional basal ones on long capillary > 534 MACKENZIE: NOTES ON CAREX peduncles), approximate, erect, sessile or a little peduncled, 5-10 mm. long, 4-8 mm. wide, short-oblong or suborbicular, containing 4-10 ascending, rather closely arranged perigynia in few ranks; bracts leaflet-like, shorter than to a little exceeding inflorescence, little sheathing, occasionally with a purplish tinge at base; scales ovate, obtuse to cuspidate, purplish with lighter center and hy- aline margins, somewhat wider and longer than perigynia; peri- gynia 5 mm. long, the body globose, 2.25 mm. wide, puberulent or short-pubescent, 3-angled, noticeably nerved, abruptly nar- rowed to a prominent stipitate base and abruptly beaked at apex, the beak 0.75-1.25 mm. long, strongly bidentate; achenes ob- tusely triangular, closely fitting within perigynia, short-obovoid, 2 mm. long; stigmas three. All specimens seen referable to this species are from the coastal counties of California and the islands off the coast. The range so shown is from Santa Cruz Island and Santa Barbara north as far as Sonoma County. The specimens from the Yosemite Valley referred to this species seem more properly referable to the following species. SPECIMENS EXAMINED: CaLiFoRNIA: Oakland, Bolander (C., L. S.*); ‘California,”’ Bolander, 1865 (L. S.); Oakland, Bolander 20 (H.); Oakland, Bolander 2295, May, 1866 (H.); Island of Santa Cruz, Brandegee, Apr. 1888 (N. Y.); Monterey County, Davy 7366 (P.); Sonoma County, Congdon 84, May 29, 1892 (P.); Mt. Tamalpais, Marin County, Heller 5716, June 18, 1902 (L. S., N. Y.); San Diego, Brandegee, 1889 (N. Y.); Sonoma County, Congdon, May 30, 1895 (L. S.); Santa Lucia Mts., Monterey County, Davy 7724, May, June, 1901 (H.); Santa Barbara, Brewer 303 (H.; L. S.); Brown 370, June 1897 (N. Y.); Redwoods, Marin Co., Bolander V+W, 1866 (C.). 2. Carex Brainerdii sp. nov. In large clumps from slender, elongated rootstocks, the culms from very short to 15 cm. high, phyllopodic, reddened and slightly * a to the abbreviations of the names of herbaria used in the present paper: arvard University; L. S., Leland Stanford Junior University; N. Y., New — York es Garden; P., S. B. Parish; B., Ezra Brainerd; K. M., Kenneth K. Mackenzie; D. C., Geological Survey of Canada; N., U.S. eas C., Colum- bia University; Piper, C. V. Piper. MACKENZIE: NOTES ON CAREX 535 fibrillose at base, much exceeded by leaves, slender, very rough on the sharp angles; sterile shoots strongly aphyllopodic. Leaves with well-developed blades four to eight to a fertile culm, the blades flat with slightly revolute margins, 1.5—3 mm. wide, those of the fertile culms short, of the sterile up to 25 cm. long, much roughened on both sides and averaging wider than those of the fertile; terminal spike slender, staminate, 5-8 mm. long, about 0.5 mm. wide, sessile or short-peduncled, few-flowered, the scales lanceolate or ovate-lanceolate, acuminate with 1-3-nerved green center and broad hyaline margins richly chestnut-tinged; pistillate spikes 4-6, 1-4-flowered, the upper 2 or 3 approximate, sessile or short-peduncled, the others basal, widely separated and strongly peduncled (the erect peduncles not much elongated), the zigzag rachis often terminating in a sterile flower; bracts of upper spikes well-developed, green, hyaline-margined and chestnut-tinged, all or only the lowest exceeding inflorescence, the lower more or less strongly sheathing, the upper sheathless or nearly so; scales ovate or ovate-lanceolate, cuspidate or long-awned with several-nerved green center, and with wide, strongly hyaline reddish-brown tinged margins, usually slightly longer but narrower than perigynia; perigynia softly short-pubescent, green, usually reddish-brown tinged, 4.5 mm. long, the body oval, 2.5 mm. long, 1.75 mm. wide, strongly 2-ribbed, finely many-nerved on outer face, nearly or- bicular in cross-section, strongly stipitate (1 mm.), abruptly con- tracted into the serrulate, hyaline-tipped, bidentate beak 1 mm. long; achenes triangular with strongly convex sides, closely en- veloped by perigynia, 2.25 mm. long, nearly 1.75 mm. wide, truncate at apex, round-tapering at base; style slender, not en- larged at base, readily detached; stigmas three. The type specimen (herb. Brainerd) of this distinct plant was collected by Dr. Ezra Brainerd (z21) in El Dorado County, California, on July 19, 1897, on a mountain north of Slippery Ford in the Sierra Nevada Mountains. A duplicate of the type is in the Gray Herbarium. Bolander 6196, collected in 1866 in the Yosemite Valley (H.; L. S.), a specimen collected by Mrs. Austin, in 1877, in Plumer County (H.), and H. E. Brown 370, collected near Sisson, Siskiyou County, California (N. Y.) are probably to be referred to this species. None of these specimens, however, is in very good condition. 3. CAREX NIGRO-MARGINATA Schwein. Ann. Lyc. N. Y. 1: 68. 1824 Carex lucorum var. nigro-marginata Chapm. Fl. S. E. U. S. 539. ~ 536 MACKENZIE: NOTES ON CAREX Densely cespitose, the stolon short, ascending; culms 2—10 cm. high, much exceeded by the leaves, triangular, rough on the angles, very strongly fibrillose at base, several together with a common cluster of leaves at the base. Leaves numerous and conspicuous, the blades from very short to 35 cm. long, 2-4 mm. wide, flat, very rough, the mid-nerve prominent; staminate spike sessile, 5-8 mm. long, 2-3 mm. wide, the ovate scales obtusish, brownish with greenish midrib and narrow hyaline margin; pistillate spikes two or three, sessile, erect, contiguous or the lower slightly separate, the upper at the base of the sessile staminate spike, orbicular or ovoid-orbicular, 4-7 mm. long, 3-5 mm. wide, closely flowered, with about 6-15 ascending perigynia; bract of lower spike well developed, green, attenuate, 1.5 mm. wide at base, 5-25 mm. long, and from shorter than to exceeding the inflorescence, the upper bracts similar but shorter; scales ovate, acutish to short-cuspidate, from slightly shorter to slightly longer than the perigynia, with a broad strip of green in the center and con- spicuous brownish black margin, or at times in immature speci- mens the brownish black margin narrow and inconspicuous; perigynia 3.5 mm. long, the body oval, 1.75 mm. long, tapering into the stipitate base 0.75 mm. long, the beak 1 mm. long, the body compressed-orbicular and somewhat obscurely triangular in cross-section, I mm. wide, narrowly ridged along the sides, otherwise nerveless, it and the beak minutely puberulent, the beak bidentate; achenes short-triangular, 1.5 mm. long, closely fitted to the perigynia; stigmas three. Differs from Carex floridana in (1) its normally triangular achenes, (2) strongly dark-margined scales, and (3) often stiff culms, which are (4) strongly fibrillose at base, and (5) in its short ascending stolons instead of long creeping ones, thus making the plant more densely cespitose. Forms of Carex varia Muhl. in which the leaves much exceed the culms may key into this species or the following one. Their narrow slender leaf-blades, and the lack of dark margins to their scales distinguish such specimens from the present species, and the absence of the long stolons and the lenticular achenes of Carex floridana serve to distinguish that species. SPECIMENS EXAMINED: NEw York: Babylon, Leggett, May 21, 1869 (C). NEw Jersey: Landisville, Atlantic Co., Gross, 1872+1883 (N. Y.) and May 28, 1898 (K. M.); Tuckerton, Mackenzie, MACKENZIE: NOTES ON CAREX 537 May, 1911 (K. M.); South Lakewood, Mackenzie 4536, May 15, 1910 (K. M.); Milford, Porter, May 27, 1867+1868 (Ne ¥33 Lakewood, Torrey Club, May 28, 1898 (N. Y.); Holland Station, Garber, June, 1868 (C); “‘New Jersey,”’ Smith (B). PENNSYLVANIA: Monroe, Bucks County, Ruth, May, 1885 (N. Y.); French Creek, Brinton, May 22, 1892 (C). DELAWARE: Centreville, Commons, May, 1864 (N. Y.). District OF CoLuMBIA: Steele, April 29, 1899 (K. M.); M. A. Curtis, 1845 (C). NortH CARrotina: M. A. Curtis (N. Y.); Chapel Hill, Ashe 3006 (N. Y.), April, 1897 (B); Salem, Schweinitz (C) ; ‘‘ North Carolina,” Chapman (C); Hunter (C); Tryon, Brainerd, April 14, 1909 (B). SoUTH CAROLINA: Greenville, Mackenzie 2991, April 2, 1908 (K. M.); Clemson, Oconee County, House 3150, March 20, 1907 (Nevo: ALABAMA: Auburn, Earle, April 10, 1901 (N. Y.); ‘‘ Alabama,” Peters, 1867 (N. Y.). Louisiana: Leavenworth, 1845 (C). TENNESSEE: Broad River, Rugel (D. C.). ARKANSAS: Leavenworth, 1845 (C). Missouri: Campbell, Bush 6597, April 19, 1912 (K. M.). 4. CAREX FLORIDANA Schwein. Ann. Lyc. N. Y. 1: 66. 1824 Carex lucorum var. floridana Chapm. FI. S. E. U.S. 539. 1860. Carex nigro-marginata var. subdigyna Bockl. Linnaea 41: 220. 1877. ; Carex nigro-marginata var. floridana (Schwein.) Kiikenth. in Engler, Pflanzenreich 4”: 444. 1909. : Culms very slender, or capillary, erect or spreading, from very short to 2 dm. high, roughened on the angles, exceeded by the leaves, coming up in small clumps and long-stoloniferous, slightly filamentose at base. Leaves largely basal, those on the culms abortive or very short, the basal leaves 2-3 dm. long, 1-2 mm. wide, flat or somewhat involute (especially near the base), more or less glaucous and roughened; staminate spike one, terminal (but exceeded by the contiguous pistillate spikes), very short and in- conspicuous, few-flowered, sessile, 3-5 mm. long, 0.75 mm. wide, the closely appressed scales lanceolate, obtuse, with green midrib and white hyaline margins; pistillate spikes two, sessile, very 538 MACKENZIE: NOTES ON CAREX close together, short-oblong, 4-8 mm. long, 2.5—-4. mm. wide, 4-8- flowered, the perigynia appressed-ascending; bracts lanceolate or ovate-lanceolate, white-hyaline with green midrib, acuminate or cuspidate, shorter than the subtended spike; scales oblong or ovate-oblong, acuminate to obtusish, white-hyaline with green midrib, rather wider but shorter than the perigynia; perigynia puberulent, spindle-shaped, 3.25 mm. long, the body oval, com- pressed-orbicular in cross-section, 1.5 mm. long, 0.75 mm. broad, tapering to the stipitate base 1 mm. long, and rather abruptly into the slender beak 0.75 mm. long, the orifice entire or nearly so, the body nerveless except for the prominent decurrent edges of the beak; achenes normally lenticular, closely fitting the perigynia, the face oblong-elliptic, 1.75-2 mm. long, 1 mm. wide; stigmas two. SPECIMENS EXAMINED: GeorGiA: Rocky Comfort Creek, Louisville, Jefferson Co., Harper 2105, April 9, 1904 (N. Y.). FLoripA: Jacksonville, Curtiss 4128, April 3, 1893 (C), 4639, March 24, 1894 (C), 6127, March 31, 1898 (K. M.); Hibernia, Canby, March 1869 (C); “Florida,” Chapman (N. Y. and C); “Florida,” Keeler (C); Banks of Little River, Chapman (C); West Florida, Chapman, 1836 (C); Apalachicola, Biltmore 1783 (N. Y.); “Florida,” Chapman, from. Dr. Lemann, 1847 (N. Y. ex herb. Boott). Louisiana: Jackson, Ingalls, February (C); Hale 707 (C). Mississippi: Ocean Springs, Earle, Feb. 1889, very young (N. Y.). Texas: Big Sandy, Bush 2877, April 7; 1902 (NEY) 5. CAREX DEFLEXA Hornem. Plantelaere, ed. By 0S OSB: 1621 Carex varia var. minor Boott, in Hook. Fl. Bor.-Am. 2: 223 (in part). 1840. Carex pilulifera var. deflexa Drej. Rev. Crit. Car. Bor. 54. 1841. “ Carex Novae-Angliae Schw.” Boott, Ill. Car. 2: 96 (in part). 1860. Carex pilulifera forma, Béckl. Linnaea 41: 216. 1877. Carex deflexa var. Deanei Bailey, Mem. Torrey Club 1: 42. 1889. Clumps small or medium-sized, stoloniferous, the rootstocks slender, horizontally creeping, branching; culms 3-10 cm. high, very slender, exceeded by the leaves, smooth, the fertile mostly phyllopodic. Fertile culms with several leaves with well-developed MACKENZIE: NOTES ON CAREX 539 blades inserted towards the base, the blades ascending, 1 mm. wide, usually less than 6 cm. long, roughened on the margins and towards the apex; leaves of sterile culms more numerous and with longer and somewhat wider blades; staminate spike solitary, erect, sessile, 2-4 mm. long, 0.5-I mm. wide, inconspicuous and often exceeded by the closely contiguous pistillate spikes, the scales ovate-lanceolate, closely appressed, acute, straw-colored with hyaline margins and tinged with reddish brown; pistillate spikes one or two, sessile or short-peduncled, approximate, suborbicular or short-oblong, 2-4 mm. long, 3 mm. wide, containing about 2-8 ascending perigynia, normally with an additional, widely separated, basal spike; lower bract 5-10 mm. long, not sheathing and hardly colored at base; the upper much shorter; scales ovate, acute or short-acuminate, wider but shorter than the perigynia, tinged with reddish brown, the midrib green and the margins hyaline; peri- gynia puberulent, obovoid, obtusely triangular, nerveless or nearly so, the body 1.5 mm. long, I mm. wide, tapering to a stipitate base 1 mm. long, and abruptly contracted into a short (0.5 mm. long) beak with emarginate or shallowly bidentate orifice; achenes triangular (rather obtusely), short-oval, 1.5 mm. long; stigmas three The arstien forms of Carex varia Muhl. often resemble forms of Carex deflexa in which the basal spikes are not developed. They are, however, strongly cespitose and have squamiform bracts, which are normally hyaline-margined at base and do not usually exceed the culms. SPECIMENS EXAMINED: Arctic AMERICA: Norway House, Richardson (N.Y. and C.). QUEBEC: Montmorency Falls, Macoun 67592, June 28, 1905 (N. Y.); Calumet, Macoun 7405, June 9, 1891 (D. C.). ONTARIO: Sudbury Junction, Macoun 30979, May 24, 1884 (D.C) NEw Brunswick: Kent County, Brittain 30974, 1888 (D. C.); Prince Edward Island, Macoun 10702, June 26, 1888 (D. C.). Nova Scotia: Truro, Macoun 10704, June 12, 1883 (D. C.). British CoLtumBIA: Latitude 54°, Macoun 30970, June 10, 1875 (D. C.); McCloud’s Lake, Macoun 30972, June 24, 1875. Maine: Mt. Desert Island, Faxon, June 23, 1891 (N. Y.); Orono, Fernald, May 15, 1902 (N. Y.; K. M.); and May 29, 1890 (C.); Ft. Fairfield, Fernald 149, July 6, 1893 (C.); Mt. Desert 540 : MACKENZIE: NOTES ON CAREX Island, Rand, June 21 and 23, 1891 (C.); Piscataquis County, Fernald 269, July 5, 1895 (C.). New Hampsuire: Franconia, Faxon, June 9, 1893 (N. Y.); Lisbon, Graves, June 10 and 13, 1893 (C.); Crawford Notch, Faxon, June 7, 1878 (C.); Wing Road, Pringle, June 4, 1878 (B). VERMONT: Ripton, Brainerd, May 25, 1878 (B); June 7, 1879 (B); June 10, 1892 (B; N. Y.); Ripton, Brainerd & Eggleston, June 9, 1898 (K. M.); Lyndon, July 2, 1875 (N. Y.); Groton, Pringle, June 13, 1879 (B). New York: Lake Placid, Peck, June 12, 1897 (N. Y.); White Face Mountain, Parry, Aug. 1851 (C.). Massacuusetts: Hawley, Forbes, May 30, 1905 (K. M.). MicuiGan: Keweenaw County, Farwell 745, July 1, 1890 (Cb 6. CAREX BREVIPEs W. Boott, in S. Wats. Bot. Calif. 2: 246. 1880 Carex globosa var. brevipes W. Boott, in S. Wats. Bot. Calif. 2: 485. _ 1880. Carex deflexa var. Boottii Bailey, Mem. Torrey Club 1: 43. 1889. Carex pilulifera var. Novae-Angliae F. Kurtz, Bot. Jahrb. 19: 419. 1894. Carex Rossii var. brevipes (W. Boott) Kiikenth. in Engler, Pflan- zenreich 4%: 452. 1909. In dense clumps from stout, matted, ascending rootstocks, not stoloniferous, the culms from very short to 18 cm. high, phyllopodic, reddish-purple tinged and more or less strongly hidden at their base, slender, roughened on the angles above; sterile culms aphyllopodic. Leaves with well-developed blades 4-8 to a fertile culm, the blades flat with slightly revolute margins, I.5-2.5 mm. wide, up to 15 cm. long, roughened towards apex; terminal spikes staminate, slender, short-peduncled or sessile, 4-12 mm. long, 2.5 mm. wide, several-many-flowered, the scales ovate, acute or cuspidate, about 3-nerved, reddish- or light purplish-brown with lighter center and narrow hyaline margins; pistillate spikes 3-5, usually 10-20-flowered, the upper one or two approximate, from sessile to strongly peduncled, the others widely separated, basal, long-peduncled, the perigynia in several ranks, ascending; bract of lower non-basal spike leaflet-like, exceeding inflorescence, green, slightly purplish-auricled at base; scales ovate, acute to cuspidate, I-3-nerved, green with narrow MACKENZIE: NOTES ON CAREX 541 hyaline margins and more or less strongly purplish-tinged, about the width of but shorter than mature perigynia, exposing the upper part; perigynia small, 2.5 to nearly 3 mm. long, 1.25 mm. wide, green, puberulent, the body little longer than wide, obscurely triangular in cross-section, 2-ribbed, otherwise nerveless, from short to rather strongly stipitate, abruptly contracted into the slender, minutely serrulate, slightly colored or hyaline-tipped shallowly bidentate beak 0.25-0.75 mm. long; achenes triangular with strongly convex sides closely enveloped by perigynia, nearly 2 mm. long, truncate at apex, round-tapering at base; style short and slender; stigmas three. This species, lly collected in the Sierra Nevada Mountains of California, has no immediate relationship with Carex globosa Boott, to which it has been referred. As pointed out by Prof. Bailey, its real relationship is with Carex deflexca Hornem. and its allies. It resembles real Carex deflexa in its small perigynia, 2.5-3 mm. long with shallowly bidentate beak. It, however, is densely cespitose with stout rootstocks, while Carex deflexa is stoloniferous and has slender rootstocks. The staminate spikes in the plant of the Sierra Nevadas are more developed, the perigynia seem less strongly stipitate, the spikes are usually more flowered, and the plant is much stiffer. Carex Rossi, with larger deeply bidentate perigynia, seems constantly. different. SPECIMENS EXAMINED: CaLiForNIiA: ‘“‘Lake Tahoe to Bear Valley,” Kelloce, and “Rocky Mts. California,” Kellogg (type sheets in Gray Herbarium) ; Sierra Nevada, Braman & Kellogg, June 9, 1870 (N. Y.); Sierra Nevada Mts., El Dorado County, Brainerd 116, July 19, 1897 (H; Brainerd). “WASHINGTON: Wenatchee Mts., Kittitas County, Elmer 453; June 28, 1897 (Piper). 7. Carex Rossi Boott, in Hook. Fl. Bor.-Am. 2: 222. 1840 Carex Novae-Angliae var. Rossii Bailey, Bot. Gaz. 10: 207. 1885. Carex deflexa var. Rossii and var. media Bailey, Mem. Torrey Club 1: 43. 1889. Carex Novae-Angliae var. deflexa Bailey, Proc. Am. Acad. 22: . 124. 1886. Carex deflexa var. Farwellii Brit., Brit. & Br. Ill. Fl. 1: 334. 1896. - 542 MACKENZIE: NOTES ON CAREX Carex Farwellit (Brit.) Mackenzie, Bull. Torrey Club 37: 244. I9I0. Clumps medium-sized, densely or loosely cespitose, hardly stoloniferous, the rootstocks ascending; culms 5-20 cm. high, slender, usually exceeding the leaves, smooth or slightly roughened under the inflorescence, aphyllopodic or phyllopodic; basal spikes usually numerous. Fertile culms with several leaves with well- developed blades inserted towards the base, the blades ascend- ing, I-2 mm. wide, usually less than 6 cm. long, roughened on the margins and towards the apex; leaves of sterile culms more numerous and with longer and somewhat wider blades; stami- nate spike sessile or nearly so, erect, 3-8 mm. long, 1 mm. wide, exceeding the contiguous pistillate spike, the scales ovate-oblong, closely appressed, acutish, with green midrib and hyaline margins and strongly tinged with reddish brown; pistillate spikes one or two, sessile or short-peduncled (and with some additional widely separated basal ones), approximate or somewhat separate, sub- orbicular or short-oblong, 3-5 mm. long, 3-4 mm. wide, containing 3-10 ascending perigynia; lower bract 0.5-5 cm. long, not sheathing and hardly colored at base; the upper much shorter; scales ovate, acute to acuminate, or cuspidate, wider but shorter than the mature perigynia, strongly tinged with reddish- or purplish-brown, the midrib green and the margins hyaline; perigynia short-pubescent, 3-4-5 mm. long, obovoid, obtusely triangular, 2-ribbed but other- wise nerveless or nearly so, the body 1.5-2.5 mm. long, 1 mm. wide, tapering to a stipitate base 0.5—-1.5 mm. long, and abruptly contracted into a bidentate beak 0.75-1.25 mm. long; achenes tri- angular (rather obtusely), short-oval, 1.5 mm. long; stigmas three. This characteristic species of the western mountains has been given a great deal of study by me in an endeavor to ascertain definitely whether it represented an aggregate of more than one species or not. As a result I have come to the conclusion that but one variable species is represented and that the notable variations shown in individual plants are to be explained by en- vironmental conditions. The species varies from densely cespitose in alpine or subalpine situations to loosely cespitose in more pro- tected localities; these more loosely cespitose plants have the leaf-blades of the sterile culms much more developed and in all respects show a stronger vegetative growth; the scales of the basal spikes are inclined to be strongly cuspidate’ and the perigynia range in length from 3.75 to 4.5 mm. with a beak as long as the y- Such plants answer to Carex Farwellii and are quite dif- MACKENZIE: NOTES ON CAREX 543 ferent in appearance from the densely cespitose plants with much less developed vegetative growth, acute scales, and _ perigynia ranging from 3 to 4 mm. in length, with beak shorter than the body, which represent the other extreme of the series. However, all kinds of intermediate combinations appear, and make separation impossible except arbitrarily. Other plants with narrow involute leaves collected in very dry situations also look quite different, but their peculiar aspect seems wholly due to their dried-up condition, as no structural differences have been found. The species as here treated ranges from Vancouver Island and the Canadian Rockies south through Washington and Oregon to the higher Sierras of California and eastward in the higher moun- tains of Nevada and Utah. It seems common in Colorado, Wyoming, Montana, and Idaho. It has been found in northern Michigan. It is to be expected in northern New Mexico and possibly Arizona, and its northern range in western Canada is yet to be ascertained. SPECIMENS EXAMINED: CANADA: Rocky Mt. Park, Macoun 64056, July 8, 1904 (N. Y.); between Kettle and Columbia Rivers, Macoun 63318, June 6, 1902, and 63319, July 9, 1902 (N. Y.), and 63320, July 19, 1902 (N. Y.); Nanaimo, Vancouver Island, Macoun 76742, July 13, 1908 (N. Y.); Medicine Hat, Assiniboia, Macoun 7402, June 4, 1894 (C.); Mountain Post, Assiniboia, Macoun 10780, June 11, 1895 (N. Y., H); Revelstoke, Shaw 834, July 6, 1905 (N. Y.); Revelstoke, Macoun 57A, May 19, 1890 (C.); Lake Louise, Macoun 64055, July 20, 1904; Mt. Arrowsmith, Rosendahl 2023, June 28, 1907 (N. Y., H); Qu’Appelle Valley, Macoun 49, June 22, 1879 (H); Yale, Macoun, June 17, 1876 (H); Grand Valley, Macoun 261, June 16, 1880 (H); Trail to Asalkan Glacier, British Columbia, Brainerd, Aug. 11, 1897 (B); Vancouver, Macoun, July 27, 1887 (B); Calgary, Macoun 25460, June 7, 1897 (D. C.); Mt. Arrow- smith, Vancouver Island, Macoun 30969, July 17, 1887 (D. C.); Rogers Pass, Selkirk Mts., Macoun 30968, July 29, 1890 (D. C.); Elbow River, Rocky Mts., Macoun 25461, July 15, 1897 4 bog tees Esquimault, Vancouver Island, Macoun 376, June 9, 1893, and 544 MACKENZIE: NOTES ON CAREX _ Mt. Benson, Macoun 377, July 10, 1893 (D. C.); Nanaimo and Horne Lake, Vancouver Island, Macoun 20286, June 4, 1887, and 10706, July 25, 1887 (D. C.); Kananaskis, Rocky Mts., Macoun 32016, June 15, 1885 (D. C.); Spy Hill, Saskatchewan, Macoun 72789, July 1, 1906 (D. C.); Clearwater River, Macoun 32007, July 8, 1888 (D. C.); Lytton, British Columbia, Macoun 32008, April 17, 1889 (D. C.); New: Westminster Junction, British Colum- bia, Macoun 80878, April 19, 1889 (D. C.). WASHINGTON: Bingen, Suksdorf, May 12, 1909 (N. Y.); Mt. Paddo, Suksdorf, Aug. 9, 1909 (N. Y.); Olympic Mts., Elmer 2718, June 1900 (N. Y., Piper); Mt. Rainier, Piper 2537, Aug. 15, 1895 (Piper, N. Y.); West Klickitat County, Suksdorf 276, April 30, 1885 (C.); “Washington,” Henderson, 1892 (C.); Mt. Adams, Howell, Aug. 15, 1882 (H); Cascade Mts., Allen 168, June 4, 1895 (N. Y., H, Piper); Klickitat River, Suksdorf 48, June 2, 1883 (H); Cascade Mts., Vasey, 1889 (Piper); Whitman County, Piper 3094, July 20, 1899 (Piper): Blue Mts., Horner 480, July 29, 1897 (Piper); Mt. Rainier, Piper 2552, Aug. 1895 (Piper); Mt. Adams, Henderson 2094, Aug. 10, 1892 (Piper); Olympia, Henderson, 1892 (Piper); Hangman Creek, Spokane County, Sandberg & Leiberg 30, May 17, 1893 (Piper); Wenatchee, Brandegee 1145,1883 (D.C.). EGON: Union County, Cusick 1322, 1886 (Piper) ; “Oregon,” Cusick (Piper) ; Wallowa Mts., Sheldon 8535, July 12, 1897 (K. M.); Lake County, Eggleston 6850, June 5, r911 (N); Mt. Adams, Chickering, Aug. 1882 (D. C.) CALIFORNIA: Sierra Nevada, Kellogg (N. Y.); Cisco, Sierra Nevada, Kellogg, June 10, 1870 (H); Hat Creek, Shasta County, Eggleston 7382, 7434, 7435, July 31, 1911 (N) also 7485, Aug. 2, 1g1r (N); Summit, Placer County, Heller 9853, July 16, 1909 (K. M.); Pyramid Peak, El Dorado County, Hall & Chandler 4749, Aug. 1-2, 1903 (H). | Utan: Big Cottonwood Canyon, Salt Lake Co., Garrett 1658, Aug. 21, 1905 (N. Y.); Marysvale, Jones 5343, May 31, 1899 (N. Y., K. M.); Alta, Jones 1204, Aug. 7, 1879 (C.); Hornwood Canyon, Watson 1260, July 1869 (C.). IpaHo: Kootenai County, Sandberg, MacDougal & Heller 234, May 23, 1892, and 841, Aug. 5, 1892 (N. Y.); Nez Perces County, A.A. & E. G. Heller 3388, July 9, 1896 (C., Piper); Kootenai MACKENZIE: NOTES ON CAREX 545 County, Sandberg, July 1887 (N. Y.); Sweetwater, Heller, July 1896 (K Montana: Bozeman, Flodman 289, July 7, 1896 (N. Y.); Little Belt Mts., Flodman 288, Aug. 1896 (N. Y.); Helena, Kelsey, July 12, 1892 (N. Y.); Little Belt Mts., Rydberg 3377, 3802, Aug. 1896 (N. Y.); Spanish Basin, Madison Range, Flodman 287, July 18, 1896 (N. Y.). CoLtorapo: Vasey (N. Y.); Chamber’s Lake, Baker, July 13, 1896 (N. Y.); La Plata Mts., Baker, Earle & Tracy 685, July 14, 1898 (N. Y.); Chamber’s Lake, Crandall, July 25, 1894 (N. Y.); Chamber's Lake, Colo. Agri. College 2549, July 28, 1889 (N. Y.); Beaver Creek, Colo. Agri. College 2558, July 19, 1898 (N. Y.); Pagosa Peak, Baker 237, Aug. 1899 (N. Y.); Cameron Pass, Baker, July 14, 1896 (N. Y., C.); Middle Park, Beardslee, Aug. 1892 (N. Y.); Colorado Springs, Jones 59, May 14, 1878 (N. Y., C., B); Silver Plume, Rydberg 2416, and Shear 669, Aug. 21, 1895 N. Y.); Twin Lakes, Wolfe 1058, 1873 (C.); Mt. Helen, Mac- kenzie 307, Aug. 1901 (K. M Wyominc: Big Horn Mts., Sheridan County, Tweedy 2246, July 1899 (N. Y.); Black Rock Creek, Teton Forest Reserve, Tweedy gor, Aug. 1897 (N. Y.); Teton Pass, Merrill & Wilcox 1249, July 13, 1901 (N. Y.); Madison Canyon, A. & E. Nelson 6761, Aug. 29, 1899 (N. Y.); Battle Lake, A. Nelson 3046, Aug. 16, 1897 (N. Y.); Yellowstone Park, Williams, 1888 (N. Y.); Yellow- stone Park, A. & E. Nelson 6361, Aug. 8, 1899 (N. Y., K. M.); La Plata Mines, A. & E. Nelson 5148, Aug. 25, 1898 (K. M.); Ten Sleep Lakes, Nelson 2972, Aug. 19, 1897 (K. M.). MIcuiGANn: Clifton, Keweenaw Co., Farwell 244, June, 1890 (Col., H). * 8. Carex sity oohtia | sp. nov. In large dense clumps from slender tough ascending forking rootstocks, not stoloniferous, the culms from very short to 15 cm. high, aphyllopodic, reddish brown and more or less fibrillose at base, usually shorter than leaves, slender, very rough on the sharp angles; sterile culms aphyllopodic. Leaves with well-devel- oped blades 5-10 to a fertile culm, the blades involute but flat above with slightly revolute margins, slender, at flowering time 0.75-1.5 mm. wide, 2-25 cm. long, much roughened; terminal spike staminate, slender, 4-8 mm. long, 1.5 mm. wide, more or less 546 MACKENZIE: NOTES ON CAREX strongly peduncled, few-several-flowered, the scales ovate or obovate, obtuse to acute, I-3-nerved, purplish brown with lighter midvein and conspicuous hyaline margins; pistillate spikes 2-5, usually 2—5-flowered, the upper one or two approximate or little separate, sessile or peduncled, the others widely separate, basal and strongly peduncled, the perigynia erect-ascending, the rachis zigzag; bract of upper spike green, scarcely sheathing, slightly purplish-tinged at base, normally exceeding inflorescence; scales ovate, acute to short-cuspidate, with several-nerved green center and hyaline margins and more or less strongly purplish-brown tinged, nearly as long and nearly as wide as, but not enveloping or concealing perigynia; perigynia sparingly puberulent, green, 3-5-4.5 mm. long, the body short- to long-oval, 2.25-3 mm. long, 1.75 mm. wide, 2-ribbed and otherwise nerveless or nearly so, triangular-suborbicular in cross-section, strongly stipitate (0.75-1 mm.), abruptly contracted into the scarcely ciliate-serrulate, hyaline-tipped, obliquely cut, in age shallowly bidentate beak, 0.75-1 mm. long; achenes triangular with strongly convex sides, closely enveloped by perigynia, 2-2.75 mm. long, nearly 1.75 mm. wide, truncate and slightly apiculate at apex, rounded at base; style slender, not enlarged at base, readily detached; stigmas three. The type specimen was collected by Mr. W. W. Eggleston (6605) southeast of Tierra Amarilla, Rio Arriba County, New Mexico, in the pifion belt at an altitude of 2,320 meters in the spring of 1911 (sheet 660821, United States National Herbarium). His numbers 6536, 6540, 6542 and 6610, collected in the same locality, also represent this species. Fendler’s 889 collected in New Mexico in 1847 (H) also belongs here. Carex geophila differs in the characters given in the key and in addition the. present species has narrower leaves, more slender staminate spikes, few-flowered pistillate spikes, strongly reddened culm bases and scarcely ciliate-serrulate perigynium beak. 9. Carex geophila sp. nov. In large very dense clumps from tough, rather slender, much branched rootstocks, not stoloniferous, the culms from very , slender, very rough on the sharp angles; sterile culms strongly aphyllopodic. Leaves with well-developed blades 5-10 to a fertile culm, the blades flat with slightly revolute margins, 1.5-2.5 mm. wide, 2-15 cm. . MACKENZIE: NOTES ON CAREX 547 long, much roughened; terminal spike staminate, slender, 5-9 mm. long, 2.5 mm. wide, more or less strongly peduncled, several— many-flowered, the scales ovate, acute or short-acuminate, many- striate, purplish brown with lighter center and conspicuous white-hyaline margins; pistillate spikes 2-5, usually 5—15-flowered, the upper one or two usually approximate, sessile or short-pe- duncled (sometimes absent), the others widely separated, basal and strongly peduncled, the perigynia in several ranks, ascending; bract of upper spike (where spike is present) well developed, green, somewhat sheathing, slightly brownish-red tinged, shorter than inflorescence; scales ovate, acute to short-cuspidate, those of the upper spikes reddish brown with 3-nerved green center and white- hyaline margins, those of lower spikes slightly if at all reddish- brown tinged, all from slightly shorter to slightly longer and wider than but not enveloping or nearly concealing perigynia; perigynia puberulent, green, 3.25—4 mm. long, the body suborbicular, 2.25-2.5, mm. long, 1.75 mm. wide, 2-ribbed, otherwise nervelezs or more or less strongly nerved at base on one face, nearly orbicular in cross-section, strongly stipitate (0.5-0.75 mm.), abruptly con- tracted into the serrulate, slightly hyaline or purplish-tipped bidentate beak, 0.5-0.75 mm. long; achenes triangular with strongly convex sides, closely enveloped by perigynia, about 2.25 mm. long, nearly 1.75 mm. wide, truncate and slightly apicu- ate at apex, round-tapering at base; style slender, not enlarged at base, readily detached; stigmas three. The type specimen, collected by Mr. W. W. Eggleston (6584) at Tierra Amarilla, Rio Arriba County, New Mexico, in the spring of 1911, isin the United States National Herbarium (sheet 660800). His numbers 6614, 6474, 6466, 6550, 6556, 6458, and 6593 from the same locality also belong here, as does also his 6655 collected near Chama, Rio Arriba County. The species is also represented by a’specimen in the Gray Herbarium, collected by Dr. Greene April 22, 1880 (deep shady canyon of Mineral Creek in the Mo- gollon Mountains). 10. Carex brevicaulis sp. nov. In dense clumps, stoloniferous, the culms phyllopodic, 5-10 cm. high, slender, exceeded by the leaves, sharply triangular, very rough on the angles, reddish brown and more or less fibrillose at base. Leaves with well-developed blades 6-10 to a fertile culm, clustered near base, the blades flat, 1.5-3.5 mm. wide, usually 2.5-7.5 cm. long, roughened above and towards apex; leaf-blades of sterile culms 5-12 cm. long; terminal spike stam- 548 MACKENZIE: NOTES ON CAREX inate, few-flowered, short-peduncled, 6-9 mm. long, 1.5-2 mm. wide, the scales narrowly ovate, acute to short-cuspidate, reddish brown with light-colored midrib and center and white-hyaline, non-ciliate margins; lateral spikes 2, 3 or 4, pistillate, 4-6 mm. long and nearly as wide, the uppermost sessile near base of stam- inate, the second (if present) sessile and somewhat remote, the others far remote, basal, slender-peduncled, the maturing peri- gynia I-4, erect-ascending, the upper flowers not developing; bract of upper spikes leaflet-like, shorter than or rarely slightly exceeding culm, widened at base into reddish brown auricles with hyaline margins; scales ovate, acute to short-cuspidate, reddish brown with light-colored midrib and center and white- hyaline, non-ciliate margins, narrower and shorter than the mature perigynia; perigynia about 4 mm. long, loosely short- pubescent, more or less yellowish- Siren tinged, stipitate, the body globose, 2.25 mm. wide, 2-ribbed, abruptly contracted into the slender serrulate, rather shallowly bidentate beak 1 mm. long; achenes triangular-globose, the sides strongly convex, closely enveloped by perigynia, 2 mm. wide and slightly longer, abruptly rounded at base and apex, slightly stipitate; style slender, enlarged at base, deciduous, short; stigmas three. SPECIMENS EXAMINED: BritTisH COLUMBIA: Victoria, Macoun 76706, June 12, 1908 (N. Y.); Vancouver, Macoun, May 30, 1873 (H); Macoun 32014, May 8, 1875 (D. C.). WasHINGTON: Whidbey Island, Gardner 343, May 29, 1897 (Piper). OREGON: Yaquina Bay, Howell 2994, May 1886, type (N. Y-); Wilkes Exped. 1834—1842 (C.). CALIFORNIA: San Francisco, Kellogg, May 1880 (N = 11. Carex microrhyncha sp. nov. In large stools, spreading by short stolons; culms from very short to 15 cm. high, mostly much exceeded by the leaves, slender, triangular, rough on the angles, reddish-brown tinged and strongly fibrillose at base, phyllopodic; sterile culms phyllopodic, termi- nating the short stolons, their sheaths little if at all filamentose. Leaves numerous, erect or ascending, light green, from very short to 20 or 30 cm. in length, at flowering time 1.5—-2.5 mm. wide, in age up to 3 mm. wide, flat with somewhat revolute margins, very rough above; terminal spike staminate, sessile or short- peduncled, 5-10 mm. long, I.5-2.5 mm. wide, the scales obovate, rd MACKENZIE: NOTES ON CAREX 549 obtuse or acute, reddish brown with lighter midvein and hyaline margins; pistillate spike usually present at base of staminate, sessile or short-peduncled, globose-oblong, 3.5 mm. wide, 4-7 mm. long, its bract squamiform, reddish-brown tinged and exceeded by culm, the basal spikes 2 or 3, subglobose, 4-6 mm. long, 3.5- 4.5 mm. wide; scales broadly ovate acute or short-cuspidate, about length of but wider than perigynia and largely concealing them, strongly several-nerved, greenish or hyaline; perigynia 2.25-3.25 mm. long, the body short-oval, triangular-orbicular in cross-section, 1.25 mm. wide, short-pubescent, 2-ribbed, otherwise nerveless, tapering or contracted into a short stipitate base 0.5 mm. long, abruptly contracted into the short (0.5 mm. long) beak, less than half length of body, the beak 2-edged, hyaline-tipped, at most obscurely bidentate; achenes triangular, oblong-obovoid, filling perigynia, minutely stipitate, dull or silvery blackish, the superficial cells conspicuous, the sides convex and angles blunt and prominent; style slender; stigmas three. This species bears such a strong superficial resemblance to Carex abdita Bicknell that it was not until I came to examine the achenes that I found out that the plants were distinct. The achenes in fact much more resemble those of true Carex umbellata Schk., as described by Mr. Bicknell (Bull. Torrey Club 35: 491), and as the perigynia are those of Carex abdita, I began to doubt the excellent achene characters brought out by him. However, still further study brought out the differences shown in the key in the manner of growth and in the sterile shoots as com- pared both with C. abdita and C. umbellata. The perigynia, too, are much more concealed by the scales than in either of these species. SPECIMENS EXAMINED: / Missouri: Dodson, Jackson County, Mackenzie, May 10, 1896, and May 14, 1899, type (K. M.); St. Louis, Riehl, 1838 (C); St. Louis County, Eggert, April-May, 1887 (H). INDIAN TERRITORY: Limestone Gap, Butler (H). Texas: Blanco River (Dew. Herb., H); “‘Texas,’”’ Leavenworth (C); Dallas, Reverchon, March 1877 (C). 12. Carex Appita Bicknell, Bull. Torrey Club 35: 492. 1908 Carex umbellata var. brevirostris Boott, Ill. Car. 2:99. pl. 294. 1860. ‘Very densely cespitose, culms from very short to 15 cm, high, 550 MACKENZIE: NOTES ON CAREX mostly much exceeded by the leaves, slender, triangular, rough on the angles, reddish-brown tinged and strongly fibrillose at base, phyllopodic; sterile culms aphyllopodic, erect, the sheaths filamentose. Leaves numerous, the blades erect or ascending, light green, from very short to 20 or 30 cm. in length, at flowering time about 1.5 to 2.5 mm. wide, in age up to 3 mm. wide, flat with somewhat revolute margins, very rough above; terminal spike staminate, sessile or short-peduncled, 5-10 mm. long, 1.5—2 mm. wide, the scales obovate, obtuse or acute, reddish brown with lighter midvein and hyaline margins; pistillate spike usually present at base of staminate, sessile or short-peduncled, globose- oblong, 3.5 mm. wide, 4-7 mm. long, its bract squamiform, reddish- brown tinged and exceeded by culm, the basal spikes 2 or 3, short, oblong, 5-9 mm. long, frequently staminate at apex; scales ovate, abruptly acute or acuminate, longer and wider than peri- gynia but not concealing them, strongly several-nerved, greenish or hyaline, the upper at least reddish-brown tinged; perigynia 2.25-3.25 mm. long, the body subglobose, triangular-orbicular in cross-section, 1.25 mm. wide, short-pubescent, 2-ribbed, otherwise nerveless, tapering or contracted into a short-stipitate base, 0.5 mm. long, abruptly contracted into the short (0.5—1 mm. long) beak, less than half length of body, the beak 2-edged, hyaline- tipped, at most obscurely bidentate; achenes triangular, orbicular- obovoid, filling perigynia, brownish, sessile, shining, irregularly pitted, the sides convex and angles sharp and narrow; style slender; stigmas three. Differs from Carex umbellata in the small, short-beaked, ob- scurely bidentate perigynia, and in the achenes. A northern species extending south to Delaware and Indiana. SPECIMENS EXAMINED: CANADA: Quesnelle, Macoun, May 29, 1874 (H); Norway House, Richardson 323 (H); Rocky Mts., Richardson (H); Carlton House (H); Norway House, Richardson (H); Victoria, Vancouver Island, Macoun 16673, May 7, 1875 (D. C.); Fraser River Valley, Macoun 32015, May 18, 1875 (D. C.); ‘Rocky Mts.,” Drummond (D. C.); Hastings County, Ontario, Macoun 32019, June 15, 1865 (© .), MAINE: Vassalboro, Fernald & Chamberlain, May yy, 1902 (K. M.); Bangor, Knight, May 14, 1905 (K. M.). VERMONT: Snake Mt., Brainerd, June 11, 1897 (B). New HampsHIRrE: Mt. Willard, Faxon, June 7 (N. Y.). MACKENZIE: NOTES ON CAREX 551 MassacHUsETTs: Deerfield, Cooley (C); Boston, W. Boott (63 CONNECTICUT: Bridgeport, Eames, May 28, 1908 (N. Y.). RHODE ISLAND: Thurber, May 1846 (N. Y.). New York: Richmond Hill, Long Island, Bicknell, May 11, 1904 (N. Y.); Jamaica, Bicknell, May 19, 1905, type (N. Y.); Sparrow Bush, along Delaware River, Britton, May 30, 1903 (N. Y.); Yonkers, E. C. Howe, May 1876 (N. Y.); Whitesboro, Oneida County, Haberer 5176, June 19, 1883 (N. Y.); New York, LeRoy (C.). NEw Jersey: Hoboken, Torrey, May 1824 (C.); Andover Junction, Mackenzie 4869, May 1911 (K. M.); Columbia, Macken- zie, May 1913 (K. M.); Cranberry Lake, Mackenzie 2608, June 9, 1907 (K. M.); south of Port Jervis, Mackenzie 4584, May 30, 1910 (K. M.). DELAWARE: Townsend, Commons, May 17, 1883 (N. Y.). INDIANA: Ripley County, Deam 10578, May 19, 1912 (K. M.); Clarke County, Deam 10494, May 8, 1912 (K. M.); Wells County, Deam, April 30, 1899 (K. M.). 13. CAREX UMBELLATA Schkuhr, Willd. Sp. Pl. 4: 290. 1805 Carex umbellata var. vicina Dewey, Am. Jour. Sci. 10: pl. D. f. 13 1626; 32°. 317. 2620; . Very densely cespitose, culms from very short to 15 or 20 cm. high, much exceeded by the leaves, triangular, rough on the angles, reddish-brown tinged and strongly fibrillose at base, phyllopodic; sterile culms aphyllopodic, erect, the sheaths filamentose. Leaves numerous and conspicuous, the blades from very short to 20 cm. long, at flowering time 1.5-2 mm. wide, in age wider, very rough, flat with somewhat revolute margins; terminal spike staminate, short-peduncled, 8-12 mm. long, i.5-2.5 mm. wide, the scales obovate, obtuse or acute, reddish-brown with lighter midvein and hyaline margins; pistillate spike usually present at base of staminate, sessile or short-peduncled, globose-oblong, 3.5 mm. wide, 4-7 mm. long, its bract squamiform and exceeded by culm, the basal spikes oblong, 4-10 mm. long, 3.5—4.5 mm. wide; scales lance-ovate, short-cuspidate to acuminate, from slightly shorter to slightly longer and rather wider than the perigynia, but not concealing them, those on the shorter culms hyaline with green several-nerved center, those on the longer culms similar, but the margin tinged with reddish brown; perigynia 3.25-4.25 mm. long, 552 MACKENZIE: NOTES ON CAREX the body short-oval, triangular-orbicular in cross-section, about 1.75 mm. long and 1.4 mm. wide, short-pubescent, 2-ribbed, otherwise nerveless, abruptly short-(0.5 mm. long) stipitate, abruptly con- tracted into a beak about length of body, the beak 2-edged, hyaline- tipped, bidentate; achenes triangular, oblong-obovoid, filling peri- gynia, dull or silvery blackish, minutely roughened, the superficial cells conspicuous, sessile, the sides convex, and angles sharp and narrow; style slender; stigmas three. Carex pennsylvanica Lam. and Carex heliophila Mackenzie at times develop pistillate spikes on subradical peduncles. The long stolons characteristic of these species afford the easiest means of distinguishing such specimens from Carex umbellata and its allies. It is possible that these specimens represent, to some extent at least, hybrids with the Carex wmbellata group as they are treated by Kiikenthal, but my own inclination is to regard them as above. SPECIMENS EXAMINED: CANADA: Bic, Rimouski County, Guide Forbes, June 23, 1905 (K. M.); Point Pleasant, Nova Scotia, Macoun 16674, June 18, 1883 (D. C.); Edmonton, Ontario, White 32017, May 24, 1893 (D.€.); VERMONT: Middlebury, Brainerd, May 24, 1878, and June 9, 1891 (B); Winooski, Brainerd, June 5, 1897 (B); Chipman Hill, Brainerd, May 30, 1897 (B). MICHIGAN: Port Huron, Dodge, June 3, 1894 (K. M.); Lake Harbor, Umbach, May 28, 1898 (K. M.); Orion, Oakland County, Wheeler, May 30, 1895 (C); Grand Lodge, Wheeler 47, May 5, . 1890 (C). MAINE: Orono, Merrill, June 5, 1898 (N. Y.); Orono, Fernald, June 30, 1890. MASSACHUSETTS: South Ashburnham, Forbes, May 30, 1904; (K. M.); Manchester, Chamberlain (N. Y.); Cambridge, Mrs. Britton, May 12, 1889 (C); “‘Mass.,”” Dewey (D. C.). New York: Yonkers, E. C. Howe, May 1880 (N. Y.); ‘‘New York,” Crawe (C); ‘New York,” Gray, 1846 (C); Highlands of New York, Torrey (C). NEw JERSEY: High Point, Sussex County, Mackenzie 4564 and 4571, May 29, 1910 (K. M.); Tuckerton, Mackenzie, May 1911 M.). MACKENZIE: NOTES ON CAREX 553 14. CAREX TONSA (Fernald) Bickn. Bull. Torrey Club 35: 492. 1908 Carex umbellata var. tonsa Fernald, Proc. Am. Acad. 37:507. 1902. Densely cespitose, freely short-stoloniferous; culms 2-15 cm. high, much exceeded by the leaves, sharply triangular, strongly roughened on the angles, strongly reddish-brown tinged and fibrillose at base, phyllopodic; sterile culms aphyllopodic, the sheaths little filamentose. Leaves numerous and conspicuous, the blades spreading, deep green, 2.5-4 mm. widewithrevolute margins, 5-25 cm. long, rough towards the long attenuate apex; staminate spike 6-12 mm. long, 2-3 mm. wide, the obovate scales acute, reddish brown with greenish or straw-colored center and white- hyaline margin; pistillate spike occasionally present at base of staminate, sessile or nearly so, erect-ascending; basal pistillate spikes 2-3, on long slender peduncles, short-oblong, 6-1omm. long, 4.5-6 mm. wide, containing 3-20 closely packed appressed- ascending perigynia in several ranks; bract at base of uppermost spike setaceous, not sheathing, from much shorter than to slightly exceeding spike; scales conspicuous, ovate, shori-cuspidate to acute, wider and from slightly shorter to slightly longer than perigynia, whitish or straw-colored hyaline, with green midrib, the upper often with purplish brown margins; perigynia 3.5-4.5 mm. long, the body broadly oval, 1.75 mm. long, 1.25 mm. wide, tapering into the stipitate base 0.75 mm. long and the beak 1.75- 2.5 mm. long, the body compressed-orbicular and obscurely triangular in cross-section, 2-ribbed, otherwise nerveless or nearly so, very sparsely short-pubescent above and on the strongly 2- edged bidentate beak; achenes short-obovoid, triangular, filling perigynia, brownish, shining, pitted, the superficial cells incon- spicuous, substipitate, 1.5 mm. long, closely fitting perigynia, the sides convex and angles sharp and narrow; style slender; stigmas thr SPECIMENS EXAMINED: CanapDa: Lake Ellen, Ontario, Macoun 32020, July 1, 1884 (D. C.); “British N. W. America,” Richardson (D. C.); Chalk River, Ottawa Valley, Macoun 16675, May 30, 1884 (D. C.); Cape a L’Aigle, Quebec, Macoun 67504, July 27, 1905 (D. C.); Truro, Nova Scotia, Macoun 32018, June 14, 1883 (D. C.). Marne: Orono, Fernald, May 15, 1902 (N. Y., K. M.); also May 29, 1890 and June 2, 1890 (C). Micnican: White Hall, Wheeler, June 27, 1900 (D. C.). 554 MACKENZIE: NOTES ON CAREX MASSACHUSETTS: South Dennis, C. N. Brainerd, May 1878 (B); Nantasket, E. Brainerd, June 11, 1896 (B). RHODE IsLAND: Cumberland Mills, Collins, May 29, 1892 (C). New York: Wading River, E. S. Miller (N. Y.); Valley Stream, Bicknell, May 23, 1908 (N. Y.); Woodmere, Bicknell, May 10, 1908 (N. Y.). New Jersey: South Lakewood, Mackenzie 4544, May 15, 1910 (K. M.); Tuckerton, Mackenzie, May 1911 (K. M.); Lake- wood, Torrey Club, May 28, 1898 (N. Y.); ‘‘New Jersey,’’ Parker (N. Y., D. C.); “New Jersey,’ Knieskern (N. Y.); Forked River, ‘Torrey Club, May 29, 1896 (C); Tom’s River, Britton, May 23, 1885 (C);Stelton, Mackenzie 3024, May 3, 1908 (K. M.); Yardville, Mackenzie, May 1913 (K. M.). DistTRIcT OF COLUMBIA: Tacoma Heights, Williams, March 26, 1898 (K. M.); Holm, May 1899 (D. C.). NEw YorK City The development and behavior of the chromosomes in the first or heterotypic mitosis of the pollen mother-cells of Allium cernuum Roth Davip M. MotTtTiER AND MILDRED NOTHNAGEL (WITH PLATES 23 AND 24) It may seem to the reader of cytological literature that whoever offers a contribution upon Allium might well preface his remarks with an apology for so doing. However, certain favorable forms of both plants and animals will doubtless ever remain objects of investigation. On looking about for favorable material for class use, the senior author came upon a species of wild onion, common in certain localities in Indiana, namely, Allium cernuum Roth, a species which is regarded as more favorable than the much used Allium Cepa. A study of mitotic phenomena has been made in both vege- tative and microspore mother-cells, and the results obtained and conclusions reached differ so much in certain respects from those of Bonnevie (’11), as set forth in a recent contribution on Allium Cepa, that we have decided to present the results of our observa- tions on the pollen mother-cells at this time, reserving an account of the process of nuclear behavior in vegetative cells for a future publication. In describing the resting stage of the nucleus in the pollen mother-cell of Allium Cepa, Bonnevie (’11, 197) asserts that a number of threads radiate from a chromatin knot or lump (Chro- matinknoten), being continued into the meshes of the nuclear net. “Yom Chromatinknoten sieht man eine Anzahl Fadchen, die in den Maschen des Kernnetzes ihre Fortsetzung finden, radiar ausstrahlen.” With the disappearance of the anastomoses between these radiating threads, the latter gradually became more distinct throughout their entire length, at the same time appearing always zigzag or spirally twisted. Precisely the same structure is reported for the resting nucleus of somatic cells. Following this 555 556 MOor;rTriER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM structure there takes place (I. c. 197) a pairwise conjugation of the chromosomes, which finds its culmination in synapsis. This conjugation is accomplished by the lateral fusion in pairs of the threads radiating from the chromatin knots (I. c. fig. 20). Soon after this (I. c. 198) such nuclei become more irregular, for, in addition to the chromatin knots or lumps, there appear other dense accumulations of chromatin, so that, as a result, the original radial arrangement of the threads is no longer recognizable (I. c. fig. 21). In the light of their own preparations as compared with Bonnevie’s figs. 18-21, the writers are convinced that Bonnevie is describing the appearance of very poorly fixed and poorly stained nuclei. That Bonnevie has failed to distinguishabetween good and bad fixation is clear to the writers from the following (Il. c. 198): “Ja, das Zusammenlaufen = Chromatinsubstanz kann soweit gehen, dass alles Chromatin des I iner einzigen, optisch schwer analysierbaren Masse zusammengeballt erscheint.”’ This is true, but such phenomena do not represent normal steps in the mitotic process; they are largely artifacts. It is true, as has been pointed out some years ago by one of us (Mottier, ’07, fig. 15), that chromatin granules may sometimes form accumula- tions either by themselves or grouped about the nucleolus, but such phenomena are to be regarded more on the order of chance occurrences than as representing significant and regularly appearing stages of the nucleus. It is conceivable that such accumulations of chromatin granules may be run together or fused by the reagents, and it is highly probable that the Chromatinknoten were formed in this manner. We do not find these masses of chromatin in our preparations. Atany rate the Chromatinknoten of Bonnevie’s figs. 18-21 are not to be regarded as of any consequence in the normal process of mitosis, as will be seen from what follows. THE RESTING NUCLEUS AND SYNAPSIS The nucleus of the resting stage in the pollen mother-cells - of Allium cernuum Roth presents the well-known net or reticulum of linin upon which are distributed with more or less regularity the chromatin particles or granules. From one to several nucleoli of varying sizes are present. Fic. 1 illustrates the structure of MOTTIER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM 557 the nucleus in a pollen mother-cell soon after the last somatic division. The structure of the whole cell is the same as that of any somatic cell from any meristematic region. The growth period of both cell and nucleus now begins, and the very marked increase in the size of the nucleus as compared with that of the cell is very conspicuous in this as well as in other species of Allium (Fic. 2, 7, 8, 9). In Fic. 2 the nucleus is almost if not quite as large as it ever becomes. The nuclear reticulum is uniform, and the nucleoli may or may not be evenly spaced in the cavity of the nucleus. They do not lie in the same plane, and in making the drawing the focus was necessarily changed. Fic. 2 and all others represent rather thick sections of cells. Sometimes the chromatin granules form larger and smaller aggregates, which may be grouped about the nucleoli or removed from the latter, but we do not find large fused masses of chromatin such as Bonnevie has figured and described as ‘‘Chromatinknoten.”’ A glance at FIG. 2 shows further that there may be a tendency to form a thread, that is, ‘there will be seen stretches of linin in which the granules are arranged in lineal series. As pointed out by one of us (Mottier, 07) for Lilium Martagon, there is a tendency in Allium cernuum to form a delicate thread or spirem just before or as the nucleus passes into the synaptic contraction. Fic. 2 is about ready to begin the contraction of its net into the compact mass. FIG. 3 is a faithful attempt to illustrate the nuclear structure passing into synapsis, and FIG. 4 is a similar stage but includes the whole nucleus. These two figures were found in the same section of the loculus, in which were to be seen variously different stages of the early contraction. The writers wish to state most emphatically that there is no evidence of the fusion of two spirems during the contracting process. The spirem is formed directly from the nuclear network in the only way possible for a net to make a continuous thread, namely, by the breaking or dissolving of threads of certain meshes and the fusion of others. In the fusion of meshes several threads are seen to unite just as frequently and as certainly as one may find the fusion of only two threads. The appearance of the lateral union of two threads of certain meshes in several parts of the nucleus previous to synapsis is the strongest evidence, in the opinion of the writers, that those observers can bring forward 558 MorrieER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM who hold to the doctrine that two spirems fuse side by side during, or prior to, synapsis and who deny that the somatic chromosomes are arranged end to end ina lineal series to make the continuous hollow spirem. These authors ignore the fact that three or more threads fuse in the formation of the spirem from the net as well as only two, and as will be pointed out in a subsequent paragraph, they omit from consideration and from their series of figures the most difficult and perhaps the most important steps in the forma- tion of the bivalents from the hollow spirem. FROM SYNAPSIS TO THE BIVALENTS The stage of FIG. 4 passes directly into the closely contracted mass of FIG.5. While the chromatin is still in this state, the thread gradually shortens and thickens into a heavy cord. Even before an appreciable loosening up of the synaptic ball it is readily seen that a thick spirem, or cord, is forming from the slender thread, and, as soon as the contracted mass loosens (Fic. 6), the correct- ness of this interpretation is beyond doubt. We have in our preparation transitional stages between FIG. 5 and 6, but it was not deemed necessary to include these in the series. The thick cord thus developed now becomes distributed throughout the nuclear cavity. In Allium it is relatively thick, apparently rather uniform in structure, though sometimes lumpy, and in many cases numerous delicate threads extend from the spirem to the nuclear membrane or between adjacent or parallel portions of the cord (Fic. 7, 8). A nucleolus is usually present. At this stage a longitudinal split may be sometimes seen, but this phe- nomenon is rather the exception than the rule (Fic. 8). This fission always closes up and the two halves become so closely applied or fused that the double nature of the thread, if really present, is completely concealed before any indication of cross segmentation is discernible. Following the stage of the loose hollow spirem, the same undergoes a rearrangement before transverse segmentation, which results in a twisting, looping, and an entangling of its parts. This phenomenon found in the lilies and in other plants is known as the second contraction. In the lilies there is a central knotted or entangled portion of the spirem from which extend somewhat MOTTIER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM 559 radially loops and straight stretches of the cord, the latter with free ends. In Allium cernuum we do not find this typical appear- ance observed in Lilium. There is usually a tendency for the spirem to mass or become more closely entangled near the center of the nuclear cavity with a looping in the freer parts as shown in FIG. Io and 11, which represent the less complicated condition, but, as a rule, the entanglement is so complicated that it is not possible to follow definitely more than a few loops or turns of the eatire cord. If, for example, the spirem of FIG. 10 or II were bunched together more closely near the center with a greater twisting of the loops, we should have the more complicated state referred to above. The complexity of this step is increased by the fact that the spirem usually becomes more lumpy, or thicker places alternate with others more attenuated, just prior to, or as this rearrangement is ushered in (FIG. 9). Sometimes when the rearranged condition is not too confused, the spirem seems to be undergoing cross segmentation (FIG. 10), but whether this is the rule we are unable to say. It is certain, however, that in all, or in nearly all cases, the transverse segmentation of the spirem is accomplished during the entangled condition, or the stage of the second contraction. As segmentation is taking place there is always a violent twisting about each other of the two members of the bivalents, for as soon as the bivalents can be recognized as such, they invariably present the appearance of FIG. 12, save that they are more closely bunched together. Ordinarily they are heaped up in a morecompact mass. For the illustration we have selected a nucleus in which a less entangled massing of the bivalents is present (FIG. 12). That each bivalent, or the majority of them, represents a loop of the spirem, the two sides of which have twisted about each other, and not the two halves of the longitudinally split spirem, is in our opinion beyond question. The longitudinal split of the thread seen in the spirem disappears from sight, and, if it be present during the stages described, the two halves are so closely applied that no trace of the fission can be seen. Almost without exception the two members of each bivalent are twisted about each other, some tightly, others loosely (F1G.12,13). Inall cases the bivalents, as soon as formed, are massed and entangled into a confused heap. 560 Mor?rrieR AND NOTHNAGEL: CHROMOSOMES OF ALLIUM Later they separate and become irregularly distributed within the nuclear cavity (Fic. 14). At the same time they show a tendency to untwist, and as this is brought about the fact that many represent loops of the spirem is strikingly manifested. Bonnevie figures the looped and twisted condition of the bivalents, but the manner in which they originate from the spirem is not satisfactorily shown. In fact Bonnevie does not seem to have taken cognizance of the stages which we have described as the rearrangement of the spirem, or the second contraction, and this author has not, in our opinion, shown how her FIG. 33 is derived from FIG. 30. In our opinion her figures not only disprove the very thing she attempts to demonstrate, but lend support to the view set forth in the foregoing paragraphs, namely, that the two members of each bivalent represent pieces of the spirem that were previously arranged end to end and not the longitudinal halves of parts of the spirem. At the time of cross segmentation of the spirem the chromo- somes have attained their largest size.* During the formation of the spindle and later they seem to undergo a condensation by which their size is much reduced. Their number is seven or eight. While seven only were counted in some cases, the writers are inclined to regard eight as the correct haploid number. FROM SPINDLE TO DAUGHTER NUCLEI As is so well known, the spindle develops first as a multipolar complex which gradually becomes bipolar. Within the complex of spindle fibers the chromosomes are usually crowded so that the transition from the tightly twisted state of the two members to the large open ring-shaped structure that appears rather constantly in the equatorial plate of the mature spindle, is not readily followed (Fic. 15-17). The ring-shape of the bivalents in the spindle is doubtless the most striking phenomenon in the whole mitotic process of Allium cernuum. This form of chromosome is brought about by the fact that the chromosomes are attached to the spindle fibers at a point about midway between the ends. As the two members of each bivalent thus attached are drawn apart by the * The reader should bear in mind that Fic. 9, 1o, 11, 12, 14, 18 and 19 are more highly magnified than Fic. 13, 15, 16, 17, etc. MOotTTIER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM 561 spindle fibers, each is bent at the place of attachment and a ring results. In FIG. 16 some of the rings are seen from the edge while others have their flat sides turned toward the observer, In many of these rings it is still very evident that they were loops of the spirem as shown above. However, the open or closed ring is not the only form seen in the spindle stage. Sometimes the bivalents appear as two straight or crooked rods attached at or near the ends to the fibers, or X- and Y-shapéd forms may appear. One of the most conspicuous phenomena in the whole mitotic process of this species, and one which is very significant when viewed in the light of the kinetic processes involved in mitosis, is the shape of the chromosomes just before the appearance of the multipolar spindle and at the stage of the mature spindle (Fig. 13, 16, 17); In FIG. 13 the members of each bivalent, or at least the large majority of them, are tightly twisted about each other, while in FIG. 16 and 17 they appear just as uniformly as open rings. The question arises: what kinetic forces are responsible for the twisting in FIG. 13, and what for the condition of FIG. 17? A discussion of this interesting question would extend far beyond the limits of this paper, and we shall merely venture the opinion that neither magnetic nor osmotic activities seem applicable to the phenomena under consideration. During metakinesis, that is, just at the instant when the segments are separated, each shows iis longitudinal fission. Each half-ring, which is made into a U by the pull of the spindle fibers, is now a double U. Frequently just before metakinesis this longitudinal fission can be seen when the free ends of the bivalent are turned directly toward the observer (Fic. 18). In FIG. 19, an anaphase, the double U-like nature of the daughter segments is clearly shown. It may be remarked in passing that the large size and elongated form of the chromosomes of the mature spindle, and the fact that the halves of the U’s and v’s tend to separate from each other during the anaphase, make counting perplexing and uncertain, in spite of the large size and small number. The shape and position of the various chromosomes as they pass to the poles seem to speak strongly in favor of a pull being exerted by the spindle fibers, which are in reality fine colloidal 562 Mot?rrieER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM threads and not expressions of osmotic currents. In fact it seems extremely difficult to bring any of these phenomena under explanations based upon osmotic activity. On arriving at the poles, the chromosomes become closely crowded together in a manner well known for nearly all plants. In the organization of the daughter nuclei, the chromatin does not pass into the finely divided state by the processes of reticu- lation, alveolization or fragmentation as is characteristic of many gymnosperms and dicots. The various segments do elongate, however, to three or more times their original dimensions, be- coming somewhat lumpy or irregular in outline, and finally form a sort of interrupted spirem which is seen as a series of longer or shorter loops or turns passing from the pole to the anti-pole side of the nucleus (Fic. 22). This figure represents a daughter nucleus seen somewhat obliquely from the polar side. The course of the spirem is usually more irregular than in this figure, there being many more short and abrupt genuflections or kinks. We have spoken of this spirem as discontinuous, for the reason that what are regarded as free ends can be found. These free ends are sometimes joined by very delicate threads like the anastomosing threads extending between parallel parts of the spirem in all cells whether purely vegetative or sporogenous. If the apparently free ends were connected by thicker threads, the spirem could then be spoken of as continuous. Whether or not this spirem is continuous or interrupted is of no theoretical importance. The side of each loop, or turn, is wavy or zigzag, due, of course, to the lack of space in the nuclear cavity for the placing of the greatly elongated segments. Whether continuous or interrupted, the whole forms a sort of crown or wreath open both at the pole and anti-pole sides. We assume that the adjacent or parallel sides of the loops are homologous with the sides of the U's or V's that pass to the poles during the previous anaphase. We do not find that the loops or ends of this spirem unite at the polar side to form ‘“‘Chromatinknoten”’ either in these nuclei or in somatic cells that are normally preserved. A comparison of a daughter nucleus (FIG. 22) with a granddaughter nucleus (Fic. 23) shows that the arrangement of the chromatin is similar and that it is due to similar causes which may be and probably are purely mechanical. MOTTIER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM 563 SUMMARY The resting nucleus prior to synapsis consists of a reticulum of linin and chromatin granules and of one or more nucleoli. The ‘“‘Chromatinknoten”’ of Bonnevie are not present. Before synapsis there is, as in Lilium, a tendency to form a delicate continuous thread or spirem. There is no union of two spirems in synapsis. Synapsis is a real contraction of the nuclear net and not a growing away of the nuclear membrane from the nuclear network as claimed by Lawson. The spirem is a direct transformation from the nuclear net. The hollow spirem is a thick chromatin cord in which a longi- tudinal split is only occasionally seen and only in parts of the same. This split whenever present always closes up completely before the cross segmentation. The rearrangement of the spirem takes place which is referable to the second contraction described for the lilies and other plants. This results in an entanglement of loops and parallel parts of the spirem which twist upon each other. During this rearrangement the transverse segmentation of the spirem occurs. Each bivalent chromosome is formed by an approximation, usually side by side, of different lengths of the spirem, which may have appeared as loops or otherwise. Each bivalent is, therefore, to be regarded as two somatic chromosomes that were previously arranged end to end in the spirem. The approximation of two somatic chromosomes, side by side, or otherwise, or their adherence end to end to form bivalents, is not known as synapsis in botanical literature, nor is it properly called a conjugation. The prevalent form of bivalent upon the mature spindle is the large ring, although other forms exist. The daughter segments split longitudinally during shisha: This fission may be looked upon as a preparation for the second, or homotypic, mitosis. In the construction of the daughter nuclei, the chromatin does not pass into a finely divided state. The chromatin segments elongate greatly, becoming wavy or zigzag, and form an inter- rupted spirem by the union of a number of the free ends. This spirem is disposed in the form of a wreath or crown open at both 564 Morrier AND NOTHNAGEL: CHROMOSOMES OF ALLIUM the polar and antipolar sides. The ends of the chromatin segments do not fuse into ‘‘Chromatinknoten”’ in the daughter nucleus. INDIANA UNIVERSITY, BLOOMINGTON LITERATURE CITED Bonnevie, K. (’11). Chromosomenstudien, III. Chromatinreifung in Allium Cepa (o"). Archiv Zellforschung 6: 190-253. I9QII. Lawson, A. A. (’11). The phase of the nucleus known as synapsis. Trans. Royal Soc. Edinburgh 47: 591-604. IgII. Mottier, D. M. (?03). The behavior of the chromosomes in the spore mother-cells of higher plants and the homology of the pollen and embryo-sac mother-cells. Bot. Gaz. 35: 250-282. 1903. (07). The development of the heterotypic chromosomes in pollen mother-cells. Ann. Bot. 21: 309-347. 1907. (09). Prophases of heterotypic mitosis in the embryo-sac mother-cell of Lilium. Ann. Bot. 23: 343-352. 1909. . Explanation of plates 23 and 24 All figures were drawn trom enn with the aid of the Abbé camera lucida and with Leitz 1/12 immersion and ocular IV or age Zeiss apochromatic immersion mm., apert. 140, and seiautn g ocular 12. gnification of figures 3, 4; 9» 10, II, I2, 14, 18 and 19 about 2000; all other Plt about 1600 IG. I. Young pollen mother-cell soon after the last somatic divieion Fic. 2. Pollen mother-cell near the close of the period of — ay: ie nucleus is almost as large as it ever becom hin section of a ea passing into synapsis; the net-work is form- inga ‘hiresa or spirem. Fic. 4.. A similar stage, showing nearly the whole nucleus. These two figures were near each other in the same “leas FIG. 5. yl is complet Fic. 6: The cen mass ae up. The very long, slender thread has shortened into a eee Fic. 7 e thick, pies spirem. Fic. 8. The same stage as Fic. 7. A longitudinal split is seen in two or three Fic. 9. The spirem as it frequently appears before the rearrangement into the second contraction. At this stage the spirem may appear lumpy or with thicker and thinner portio FIG. 10, II. ial steps of the rearrangement. The chromatin cord is, as 4 rule, much more entangled than in these figures. Indications of transverse segmen- tation are seen in FIG. Io. Fic. Segmentation is about oouulessd, The two members of each bivalent are ROS ‘eben each ot Fic. 13. Similar to es preceding; the twisting is more pronounced in all chromosomes. Fic. 14. Chromosomes beginning to untwist preparatory to the formation of the spindle. MOTTIER AND NOTHNAGEL: CHROMOSOMES OF ALLIUM 565 Fic. 15. Multipolar stage of spindle. Fic. 16, 17. The fully developed spindle. Nearly all of the chromosomes are rings, Fic. 18. Spindle showing three chromosomes with the metaphase espa At the right a ring-shaped chromosome seen flatwise. In the center the two what curved members of the bivalent are stretched out tangentially upon the spin tlle e. The four free ends seen in these two chromosomes are indications of the longitudinal split. The chromosome at the left is a ring seen from the Fic. 19. An anaphase; the longitudinal fission of pies dicaghter chromosome is very evident. 1G. 20. A polar view of an anaphase. The halves of the longitudinally split daughter segments tend to separate. Fic. 21. A typical anaphase in longitudinal section. Fic. 22. A polar view of a daughter nucleus showing arrangement of the spirem into a system of loops. Fic. 23. One cell of a tetrad, or granddaughter cell. The disposition of the chromatin is the same as in the daughter cells at the corresponding stage. Contributions to the Mesozoic flora of the Atlantic coastal plain— . Alabama* Epwarp W. BEerry The Tuscaloosa formation as developed in western Alabama has been known for over fifty years to contain remains of fossil plants, and that it contained a large and varied Cretaceous flora has been known since Dr. Eugene A. Smith published a brief list of species in 1894. The principal items in the history of the study of this formation and its flora may be briefly enumerated as follows: The Tuscaloosa formation was named by Smith and Johnson in 1887 (U. S. Geol. Surv. Bull. 43: 95) from the city and river (now usually known as the Warrior or Black Warrior River) of that name in Alabama. Earlier observers had noticed the pres- ence of sands and clays below the recognized Cretaceous and above the Carboniferous, Professor L. Harper, the state geologist of Mississippi, mentioning them in print as early as 1856 (Proc. Acad. Nat. Sci. Phila. 8: 126-128) and suggesting that their age is perhaps Permian or possibly Triassic. The same year Prof, Alex. Winchell mentioned the Tuscaloosa mottled clays, calling attention to the contained vegetable remains “appearing like the stems and leaves of dicotyledonous plants.” He doubted their Triassic age and in his table of formations they appear in the Lower Cretaceous (Proc. Am. Asso. Adv. Sci. 10: 92. 1856). Meek and Hayden in discussing (Proc. Acad. Nat. Sci. Phila. 9: 117-133. 1857) the Alabama Mesozoic mentioned wood and leaves and correlated the lower part with the lowest Cretaceous of New Jersey and Nebraska. Their lithologic characterization clearly indicates that they are discussing the Tuscaloosa, and they say that although the weight of the evidence favors the correlating * Published by permission of the Director of the United States Geological Survey. The present paper is a brief abstract of the systematic chapter of a Mono- graph of the Upper Cretaceous floras of the eastern Gulf Coastal Plain, submitted for publication by the U. S. Geological Survey, this study being a part of the Coastal Plain Investigations directed by T. Wayland Vaughan. 567 568 BERRY: MESOZOIC FLORA OF ATLANTIC COASTAL PLAIN of these beds with the Neocomian of the Old World positive evi- dence is lacking that a part may not be older than Cretaceous. Subsequently, Professor Hilgard (Geol. and Agr. Miss. 61. 1860) described the beds in Mississippi beneath his Tombigbee sands as the Eutaw group and referred them to the Cretaceous. The following year Meek and Hayden restated their views and defi- nitely correlated the beds in Alabama with the Dakota of the Western Interior (Proc. Acad. Nat. Sci. Phila. 13: 419-421. 1861). Again in 1876 Meek (U. S. Geol. Surv. Terr. 9: 38-42) re- affirms his belief that the basal Cretaceous of Alabama is of the same age as the plastic clays of New Jersey and the Dakota sandstone of the Upper Missouri section. All of these geologists failed to discriminate the Tuscaloosa from the overlying sands and laminated clays of what is now known as the Eutaw formation. The first reasonably complete account of the Tuscaloosa formation is given by Smith and Johnson in the publication previously alluded to. From the attitude, lithologic ch ter, and stratigraphic position of the beds they cor- related the Tuscaloosa with the Potomac of the Middle Atlantic slope, which had just been named and briefly described by McGee (Rep. Health Officer Dist. of Columbia for the year ending June 30, 1885: 19-21, 23-35), a natural correlation since the Potomac as understood in the earlier days of its study included beds which according to the opinions of different students were referred to various levels ranging from the Triassic to the Cretaceous and which subsequent study has shown to constitute a series of well- marked formations, the oldest of Neocomian age and the youngest of Cenomanian age. From the year 1883 down to the present Dr. Eugene A. Smith, the distinguished state geologist of Alabama, has added to our knowledge of these deposits, being assisted in the earlier years by L. C. Johnson and D. W. Langdon, Jr. The discovery of all of the noteworthy localities for fossil plants is due to their efforts. In 1884 some leaf impressions collected by Langdon in Bibb County were submitted to Leo Lesquereux, among which he recognized a species of Podozamites which he thought might indicate a pre-Cretaceous age. Lesquereux afterward determined BERRY: MESOZOIC FLORA OF ATLANTIC COASTAL PLAIN 56 a small collection of leaves from the Tuscaloosa beds at Tuscaloosa but this list seems never to have been published. In 1886 Smith and Langdon discovered several localities for fossil plants in the vicinity of Tuscaloosa (Cottondale, Snows Place, Tuscaloosa) and the next year the United States Geological Survey sent Professor Fontaine into the field. The latter made large collections of mostly fragmentary material from these outcrops as well as from one or two other outcrops near the town of Tuscaloosa. In 1892 Professor Lester F. Ward visited Alabama and in company with Dr. Smith made extensive collections from Glen Allen and Shirleys Mill. These collections received a preliminary study by Professor Ward, who furnished a list of 35 species which was published by Smith in 1894 in his Report on the Geology of the Coastal Plain of Alabama. This list enumerated the following forms: Andromeda ae Newb. = Andromeda grandifolia B Andromeda ORE ae Hollick (No- vae-Caesareae) omeda Parlatoriit Hee Wellingtoniana aig = Aralia edium Newb. = Cin- namomum Newberryi Berry Cladophlebis parva Font. = Cladophlebis alabamensis Ber Cycadinocar pus circularis Newb. Czekanowskia capillaris nage * Dewalquea groenlandica Diospyros primaeva Hee Eucalyptus attenuata eae * Ficus lanceolato-acuminata Newb.* oe Woolsonit Newb. riodendropsis Bu sate : ewb. ae nies agnolia alterna 28g Magnolia siiicahets Newb. Magnolia glaucoides Newb. = Magnolia Boulayana Lesq. Magnolia longifolia Newb. = Magnolia Newberryi Berry Magnolia speciosa Heer Myrsine borealis Hee Populus apiculata Saee: = Cordia apicu- ata Berry Proteoides daphnogenoides Heer = Ficus daphnogenoides Berry Pterospermites modestus Lesq.* Tricalycites papyraceus Newb Widdringtonites Reichii (Ett. : Hee Sequoia gracillima (Lesq.) Newb. = as Pea ponties Reichii ung ) Heer lla V sky Sequ Sequoia Reichenbachi eae ) Heer In preparation for my work I spent the field season of 1909 in Alabama, revisiting all of the known plant localities and making extensive collections. In company with Dr. L. W. Stephenson the Warrior and Tombigbee river sections were studied by means * Not recognized by me. 570 Berry: MEsoOzoIc FLORA OF ATLANTIC COASTAL PLAIN of a launch trip from Tuscaloosa down to the Eocene contact at Moscow; the Coosa and Alabama rivers were traversed from Wetumpka to Montgomery; the Chattahoochee River from Columbus to Gainesville; the upper Tombigbee River in Missis- sippi and various localities in Tishomingo, Prentiss, and Itawamba counties, Mississippi, were explored. I have also had the benefit of the collections and notes made by Dr. L. W. Stephenson in his extensive field and office studies on the stratigraphy and paleo- zoology of the Cretaceous of the Eastern Gulf area, as well as the extensive collections previously made for the United States Geological Survey by Smith, Fontaine, and Ward. All of the types and duplicate material are in the collections of the United States National Museum. Recognizable fossil plants have been found at the following localities in Alabama and a single locality in northeastern Miss- issippi: near Iuka, Mississippi; Glen Allen, Shirleys Mill, Tusca- loosa, Cottondale, Snow Place, Sanders Ferry Bluff, Whites Bluff, and several other localities in Alabama where only one or two species have been found. The identifiable species other than those new to science are enumerated in the following notes. LocALITY NEAR IuKA, MISSISSIPPI This is the most northerly known plant-bearing outcrop of the Tuscaloosa formation. It is situated in a cut on the Southern Railway 13 miles east of Iuka in Tishomingo County, and while near the base of the formation in this county it is younger than the plant-bearing Tuscaloosa localities in Alabama. The following species associated with water worn pellets of amber occur at this outcrop: Andromeda Wardiana Lesq., Andro- vettia carolinensis Berry, Sequoia Reichenbachi (Gein.) Heer. LOCALITY NEAR GLEN ALLEN, ALABAMA This outcrop is in a cut of the St. Louis and San Francisco R. R. about one-quarter of a mile east of Glen Allen near the northern boundary of Fayette County. The following species occur here: Andromeda grandifolia Be Bauhinia marylandica Berry erry Andromeda Novae-Caesareae Hollick — Cinnamomum Newberryi Berry Andromeda Parlatorit Hee Cissites formosus Heer BERRY: MESOZOIC FLORA OF ATLANTIC COASTAL PLAIN 571 Cornophyllum vetustum Newb. ycadinocarpus circularis Newb. Diospyros primaeva Hee Diospyros shaver Stes fas Ficus pays gt (Heer) Berry Ficus Krausiana Lesq. Liriodendropsis simplex Newb. Lycopodium cretaceum Be Magnolia Lacoeana Lesq. Magnolia Newberryi Berry Magnolia speciosa Heer Marattia cretacea Velenovsky (?) in lis Heer Myrsine Gaudini (Lesq. Pter phagenbiics: Car Oeneners Berry Salix Lesquereuxii B Tricalycites AeA wh. Widdringtonites aad eset ) Heer Zizyphus lamarensis Berry LocaLity AT SHIRLEYS MILL, ALABAMA This outcrop is about twenty-five miles south of Glen Allen in southern Fayette County at a point where the old Fayette-Tusca- loosa coach road descends to the Davis Creek bottom. The following species occur at this locality: ndica Berry Brachyphyllum macrocarpum formosum Ber Iry arpolithus floribundus Newb. Celastrophyllum Brittonianum Hollick e Ce se salieri m Newberryanum Hollick m Newberryi Berry Cuchi: aligerum (Lesq.) Berry olutea obovata Berry Crotonophyllum panduraeformis Berry Dammara borealis Heer pices ed acutus Heer ewalquea Sm i Berry Dickaoni pvaletdie Heer — ee. Lesq. reo (Heer) Berry Inga cretacea Lesq. Laurophyllum Juglans arctica Heer Kalmia Brittoniana Hollick nervillosum Hollick Laurus plutonia Hee Leguminosites sa citonis Lesq. Liriodendropsis Segnan ene Newb. j rd ia hakcoues Lesq. Palaeocassia laurinea as Panax cretacea Heer Persoonia Lesquereuxti Knowlton Phaseolites ade Lesq. Protoda a speciosa Hollick & Jeffrey pista: carolinensis Berr Sequoia heterophylla Velenovsky Tricalycites papyraceus Newb. Widdringtonites Reichii (Ettings.) Heer Widdringtonites subtilis Heer 572 BERRY: MESOZOIC FLORA OF ATLANTIC COASTAL PLAIN LOCALITIES NEAR TUSCALOOSA The following list embraces species occurring at several out- crops in and near the town of Tuscaloosa in Tuscaloosa County: Andromeda grandifolia Berry Ficus Woolsoni Newb. Andromeda Parlatorii Heer Magnolia speciosa Heer Diospyros primaeva Heer Salix flexuosa Newb Ficus daphnogenoides (Heer) Berry Salix Lesquereuxii Berry LOCALITY NEAR COTTONDALE This locality is along the public road about 10 miles east of Tuscaloosa and two miles southeast of the town of Cottondale in Tuscaloosa County. The following species have been identified from this outcrop: Andromeda Parlatorii Heer Liriodendron Meekii Heer Baukinte cretacea Newb. Magnolia Capellinii Heer Bauhini Sregaaee Pee Magnolia longipes Newb Celasitrophyllum cre Magnolia speciosa Hee Celastrophyllum eae ee ek Malapoenna cretacea ail ) Knowlton Celastrophyllum undulatum Newb. Myrica emarginata Heer Cinna u yt Myrsine Gaudini (Lesq.) Berry Citrophyllum aligerum (Lesq.) Berry Persea valida Hollick Cocculus cinnamomeus Velenovsky (?) Phaseolites formus Lesq. Diospyros primaeva Heer Pinus raritanensis Berry Ficus cece (Heer) Berry Platanus latior (Lesq.) Knowlton Ficus inaequalis Les Populus hyperborea Heer Ficus Krausiana pitt Protophyllocladus subintegrifolius (Lesq-) Ficus Woolsoni Newb. Berry Geinitzia formosa Heer Pterospermites carolinensis Berry Tle i Lesq. Salix Lesquereuxii Berry Juglans arctica Heer Sassafras acutilobum Lesq. Laurus plutonia Heer Sequoia Reichenbachi (Gein.) Heer LocaLITy ON SNOW PLANTATION Two plant-bearing outcrops occur on the Snow Plantation about nine miles southwest of Tuscaloosa. These are known in the literature as ‘‘Upper Ravine” and ‘‘Big Gully, Snow Place”’ and are in enormous gullies eroded into the upland from the west bank of the Warrior River. The following species occur here: Abietites foliosus (Font.) Berry paps borealis Heer Andromeda grandifolia Berry Dicksonia groenlandica Heer rdeseie goed -Caesareae Hollick - Stephensoni Berry arlatorit Heer Eucalyptus Geiniizi Heer papain carolinense Berry Eucalyptus latifolia Hollick Celastrophyllum crenatum Heer Ficus crassipes Heer # BERRY: MESOZOIC FLORA OF ATLANTIC COASTAL PLAIN 573 Ficus daphnogenoides (Heer) Berry Salix Lesquereuxti Berry Ficus Krausiana Heer Sequoia ambigua Heer Laurophyllum angustifolium Newb. (?) Sequoia fastigiata (Sternb.) Heer i inata r yrica emarginata Hee Sequoia Reichenbachi (Gein.) Heer yrsine borealis Heer Tricalycites papyraceus Newb. Podozamites marginatus Heer Widdringtonites Reichii (Ettings.) Heer Salix flexuosa Newb. Widdringtonites subtilis Heer SANDERS FERRY BLUFF This locality is on the west bank of the Warrior River about eleven miles southwest of Tuscaloosa in the county of that name. The following plants occur at this outcrop: Acerates amboyensis Berry Salix flexuosa Newb. Ficus crassipes Heer Salix Lesquereuxit Berry Ficus Krausiana Heer Waites BLUFF OUTCROP This locality is on the right bank of ‘the Warrior River in northeastern Green County, three hundred and nine miles above Mobile and near the top of the Tuscaloosa formation. The , following species have been identified from this outcrop: Brachyphyllum macrocarpum formosum Sequoia heterophylla Velenovsky Berry Sequoia Reichenbachi (Gein.) Heer Dewalquea Smithi Berry Widdringtonites Reichii (Ettings.) Heer In addition to the well-known Cretaceous species in the fore- going lists the Tuscaloosa formation has yielded upwards of fifty new species which are described in the following genera: Aralia, Calycites, Capparites (2), Carpolithus, Cassia, Celastrophyllum (5), Cladophlebis, Cocculus (2), Conocarpites, Eorhamnidium, Equtse- tum, Eugenia, Ficus (3), Grewiopsis (2), Hymenaea, Junger- mannites, Leguminosites (3), Lycopodites, Malapoenna, Menisperm- ites (2), Myrica, Oreodaphne, Persoonia, Phyllites (2), Piperites, Platanus (2), Populites, Proteoides, Sapindus, Sapotacites (3), and Sphaerites. The flora as a whole comprises over 150 species, of which over 40 per cent. of the genera are not represented in the existing flora. None of the species survive into the lower Eocene. Eighty-seven genera segregated into 48 families in 31 orders are represented, the most abundant orders being the Ranales with 15 species, the Coniferales with 14 and the Urticales with 8. The largest single genus is Celastrophyllum with 12 species. The — 574 Berry: MEsozoiIc FLORA OF ATLANTIC COASTAL PLAIN Dicotyledonae of the Tuscaloosa formation number 123 species, ‘distributed in 34 families in 21 orders. The Choripetalae number 107, the Gamopetalae but 16 forms. The flora as a whole is a lowland coastal flora, many of the species being strand types. It indicates a land surface of rather uniform topography, an abundant and well-distributed rainfall, equable temperatures of warm temperate or subtropical type, with slight seasonal changes. Meager floras are found also in the younger Cretaceous strata of the Eutaw and Ripley formations, but these are not included in the present contribution. JoHNS Hopkins UNIVERSITY, BALTIMORE A bibliography of works on meiosis and somatic mitosis in the Angiosperms MAURICE PICARD The accompanying bibliography was prepared, in the first instance, for the compiler’s personal use. He publishes it, be- lieving that there is need of such a means of reference to the works on meiosis and somatic mitosis in the plants already studied, and that, by such publication, interest may be aroused in the groups hitherto neglected. As far as the writer knows, there is no such bibliography in existence at the present time, nor has the plan of citing literature on mitosis with reference to the systematic position of the plants studied ever been adopted. Inasmuch as the object of the bibliography is to provide a working basis for further research no attempt has been made to make the citations on the individual plants exhaustive. It is believed, however, that from the citations given one can obtain references to all the literature. Works published before 1880 have not been cited; the citations extend to May, 1913. Works on the morphological development of the male and female gametophytes have been mentioned only when they contain matter of cytological interest. The writer has used his own discretion with respect to articles of questionable relevance, and also in deciding whether or not incidental references to somatic mitosis should be cited. The bibliography was compiled chiefly at the libraries of Cornell University, Columbia University, and the New York Botanical Garden; and my thanks are due to their librarians for courtesies shown me. I am also obliged to Professor G. F. At- kinson, Professor R. A. Harper, and Dr. A. B. Stout for access to some otherwise unobtainable articles, and to Professor G. F. Atkinson for examining the manuscript sheets. The writer is aware that such a bibliography as here presented must be inadequate in many ways, and he hopes that those who can will acquaint him with omitted references. The nomenclature employed is that of N. L. Britton’s ‘‘ Manual 576 PICARD: BIBLIOGRAPHY of the Flora of the Northern States and Canada,”’ 2d ed., 1907, when the plants cited are contained in this volume. In the case of forms not within the range of this work, the nomenclature follows the rules laid down by the International Botanical Congress at Vienna. Where the two systems differ, the designation of the Vienna code is added in parentheses. BIBLIOGRAPHY ANGIOS PERM AE—MonocoTyLEDONES NAIADALES. NAIADACEAE Potamogeton foliosus Wiegand, K.’99, Bot. Gaz. 28: 328-359. Naias marina Guignard, L. ’98, Arch. Anat. Micr. 2: 455- 509. Guignard, L. ’99, Compt. Rend. Acad. Sci. Miiller, H. A. C. ’12, Arch. Zellforsch. 8: I-51. Zostera marina Rosenberg, O. ’o1, Bihang Ké6ngl. Sv. Vet.- Akad. Handl. 27°: 1-24. Rosenberg, O. ’o1, Meddel. Hég. Bot. Inst. Stock. ALISMACEAE Sagittaria latifolia Schaffner, J. H. ’07, Ohio Nat. 7: 41-48. GRA MINALES—GRAMINEAE Triticum vulgare Kérnicke, M. ’97, Untersuchungen tiber die Entstehung und Entwicklung der Sexual- organe von Triticum mit besonderer Be- riicksichtigung der Kernteilung. Bonn Diss. T. vulgare, Hordeum Nakao, M. ’11, Jour. Coll. Agr. Tohoku Imp. distichon, Secale cere- Univ. Sapporo, 4°. ale, T. vulgareXS. cereale T. vulgare, Aegilops Bally, W. ’12, Ber. Deuts. Bot. Ges. 39: ovata 163-172. Oryza sativa Kuwada, J. ’10, Bot. Mag. Tokyo, 24: 267- Saccharum oficinarum Franck, W. J. ’11, Somatische Kern- en Cel- deeling en Microsporogenese bij het Sui- kerriet. Diss. Delft. Amsterdam. (Bot. Centralbl. 120: 644.) Zea Mais - Kuwada, J. ’11, Bot. Mag. Tokyo, 25: 163- 181; 405-415. PicarD: BIBLIOGRAPHY 577 CYPERACEAE Carex acuta Juel, H. O.’oo0, Jahrb. Wiss. Bot. 35: 626-659. Carex aquatilis Stout, A. B. ’12, Arch. Zellforsch. 9: 114-140. ARALES—ARACEAE Arisaema triphyllum Atkinson, G. F. ’99, Bot. Gaz. 28: 1-26. .Peltandraundulata Duggar, B. M. ’oo, Bot. Gaz. 29: 81-098. (virginica), Symplo- carpus foetidus Arum maculatum Rosenberg, O. ’05, Bot. Not. 1905: 1-24. Richardia africana Overton, J. B. ’09, Ann. Bot. 23: 19-61. X YRIDALES—CoMMELINACEAE Tradescantia virginiana Strasburger, E. ’80, Zellbildung und Zell- teilung. Jena. Heuser, E. ’84, Bot. Centralbl. 17. Guignard, L. ’84, Ann. Sci. Nat. Bot. VI. 17: 5-59. Guignard, L. ’85, Ann. Sci. Nat. Bot. VI. 20: 310-376. Guignard, L. ’91, Ann. Sci. Nat. Bot. VII. 14: 163-296. Strasburger, E. ’oo, Uber Reduktionsteilung, etc. Hist. Beitr. 6: Jena. Mottier, D. M. ’03, Bot. Gaz. 35: 250-292. Strasburger, E. ’o04. Sitzungsber. Kén. Preuss. Akad. Wiss. 18: 587-614. Miyake, K. ’o5, Jahrb. Wiss. Bot. 42: 83-120. Farmer, J. B., & Shove, D. ’05, Quart. Jour. Micr. Sc. 48: 559-569. Mottier, D. M. ’07, Ann. Bot. 21: 309-347. LILIALES—MELANTHACEAE Abama ossifraga Berghs, J. ’05, La Cellule 22: 139-160. (Narthecium ossifra- . gum Grégoire, V. ’05, La Cellule 22: 221-276. LILiac Hemerocallis fulva Tangl, E. ’82, Denkschr. Kais. Akad. Wiss. Wien, 45: 67-86. Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. Juel, H. O.’97, Jahrb. Wiss. Bot. 30: 205-226. Strasburger, E. ’00, Uber Reduktionsteilung, etc. Hist. Beitr.6. Jena. 578 i Allium Erythronium Fritillaria PicarD: BIBLIOGRAPHY Strasburger, E. ’80, Zellbildung und Zell- teilung. Jena. Guignard, L. 84, Ann. Sci. Nat. Bot. VI. 17: 37 59- Guignard, L. ’85, Ann. Sci. Nat. Bot. VI. 20: 310-372. . Strasburger, E. ’88, Uber Kern- und Zell- teilung, etc. Hist. Beitr. 1. Jena. Schaffner, J. H. ’94, Bot. Gaz. 19: 444-459. Ishikawa, C. ’97, Jour. Coll. Sci. Imp. Univ. Tokyo, 10: 193-224. Schaffner, J. H. ’08, Bot. Gaz. 26: 225-238. Némec, B. ’99, Jahrb. Wiss. Bot. 33: 313-336. Strasburger, E. ’oo, Uber eae McComb, A. ’oo, Bull. "Cece Club 27: 451-459. Merriman, M. L. ’04, Bot. Gaz. 37: 178-207. Berghs, J. ’04, La Cellule 21: 171-189; 383-397. Miyake, K. ’o5, Jahrb. Wiss. Bot. 42: 83-120. Grégoire V. ’05, La Cellule 22: 221-376. Grégoire, V. ’06, La Cellule 23: 309-358. Grégoire, V. ’07, La Cellule 24: 369-420. von Derschau, M. ’o7, Beih. Bot. Centralbl. 22: 167-190. Lundegardh, H. ’10, Sv. Bot. Tidskr. 4 174-196. Lundegardh, H. ’12, Jahrb. Wiss. Bot. 51: 236-280. McComb, A., ’oo, Bull. Torrey Club 27: 451- Schaffner, J. H. ’or, Bot. Gaz. 31: 369-387: Schaffner, J. H. ’07, Ohio Nat. 7: 41-48. Strasburger, E. ’82, Arch. Mikr. Anat. 21° 476-590. Went, F. ’87, Ber. Deuts. Bot. Ges. 5: 247-258. Strasburger, E. ’88, Uber Kern- und Zell- teilung. Jena. Belajeff, Ww. ’92, Sitzungsber. Warsch. Natur- Ges. 25: Ap. u. Mai. bf PICARD: BIBLIOGRAPHY 579 Hyacinthus orientalis Scilla non-scripta Scilla sibirica Paris quadrifolia Bellevalia romana Nothoscordum fra- grans Funkia Belajeff, W. ’94, Flora 79: 430-442. Strasburger, E. ’95, Jahrb. Wiss. Bot. 28: 151-204. Strasburger, E., & Mottier, D. M. ’97, Ber. Deuts. Bot. Ges. 15: 327-332. Mottier, D. M. ’97, Jahrb. Wiss. Bot. 30: 169-204. Belajeff, W. ’97, Ber. Deuts. Bot. Ges. 15: 345-349. Belajeff, W. ’98, Ber. Deuts. Bot. Ges. 16: 27-34. Sijpkens, B. ’04, Rec. Trav. Bot. Néerl. 1: 160-218. von Derschau, M. ’07, Beih. Bot. Centralbl. 22: 167-190. Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-587. Went, F. ’97, Ber. Deuts. Bot. Ges. 5: 247-258. Strasburger, E. ’88, Uber Kern- und Zell. teilung. Jena. Hyde, E. ’o09, Ohio Nat. 9: 539-544. Overton, J. B. ’93, Vierteljahr. naturf. Ges. Ziirich 38. Schniewind-Thies, J. ’01, Die Reduktion der Chromosomenzahl und die ihr folgenden Kernteilungen in den Embryosackmut- terzellen der Angiospermen. Jena. Ernst, A. ’02, Flora 91: 1-46. Berghs, J. ’05, La Cellule 22: 201-214. Grégoire, V. ’06, La Cellule 23: 309-358. Guignard, L. ’85, Ann. Sci. Nat. Bot. VI. 20: 310-372. Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. iu Strasburger, E. ’00, Uber Reduktionsteilung, etc. Hist. Beitr. 6: Jena. Strasburger, E. ’05, Jahrb. Wiss. Bot. 42: I-71. e, K. ’05, Jahrb. Wiss. Bot. 42: 83-120. Miyak Grégoire, V. ’05, La Cellule 22: 221-376. 580 Lilium PICARD: BIBLIOGRAPHY Sykes, M. G. ’08, Arch. Zellforsch. 1: 390- 398; 525-527- Strasburger, E. ’80, Zellbildung und Zellteil- ung. Jena. Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. Flemming, W. ’82, Arch. Mikr. Anat. 20: 1-86. Guignard, L. ’84, Ann. Sci. Nat. Bot. VI. 17: 5—59- Guignard, L. 85, Ann. Sci. Nat. Bot. VI. 20: Sele * Siresbaaer, 3 . °88, Uber Kern- und Zell- teilung, etc. Jena. Guignard, L.’91, Ann. Sci. Nat. Bot. VII. 14: 163-2096. Zimmermann, A. ’93, Beitr. Morph. Physiol. Pflanzenzelle 2. Farmer, J. B. ’93, Ann. Bot. 8: 392-396. Belajeff, W. ’94, Flora 79: 430-442. Strasburger, E. ’95, Jahrb. Wiss. Bot. 28: 151-204. © Sargant, E. ’95, Jour. Roy. Micr. Soc. 283- 287. Farmer, J. B., & Moore, J. E. S. ’95, Anat. Anz. 11: 71-80. Farmer, J. B. ’95, Flora 80: 56-67. Dixon, H. H.’ 95, Proc. Roy. Irish Acad. III. 3: 707-720. Sargant, E. ’96, Ann. Bot. 10: 445-477: Sargant, E. ’97, Ann. Bot. 11: 187-224. Mottier, D. M.’97, Jahrb. Wiss. Bot. 30: 169- 204. Schaffner, J. H. ’97, Bot. Gaz. 23: 430-452- Strasburger, E., & Mottier, D. M. ’97, Ber- Deuts. Bot. Ges. 15: 327-332. Belajeff, W. ’97, Ber. Deuts. Bot. Ges. 15: 345-349. Mottier, * M. ’98, Jahrb. Wiss. Bot. 31: 125-15 stasburge, E. ’98, Jahrb. Wiss. Bot. 31: 511-59 PICARD: BIBLIOGRAPHY 581 Galtonia candicans Agapanthus, Tricyrtis Anthericum, Asphode- lus Yucca Grégoire, V. ’99, La Cellule 16: 233-298. Strasburger, E. ’oo, Uber Reduktionsteilung, etc. Hist. Beitr. 6. Jena. Dixon, H. H. ’o1r, Notes fr. Bot. School of Trinity Coll. Dublin. Mottier, D. M. ’03, Bot. Gaz. 35: 250-282. Berghs, J. ’04, La Cellule 21: 171-189. Allen, C. E. ’04, Bot. Gaz. 37: 464-470. Allen, C. E. ’05, Ann. Bot. 19: 189-258. Allen, C. E.’o5, Jahrb. Wiss. Bot. 42: 72-82. Miyake, K. ’o5, Jahrb. Wiss. Bot. 42: 83-120. Farmer, J. B., & Moore, J. E. S. ’05, Quart. — Jour. Micr. Sc. 48: 489-557. Schaffner, J. H. ’06, Bot. Gaz. 41: 183-191. Schaffner, J. H. ’07, Ohio Nat. 7: 41-48. Grégoire, V. ’07, La Cellule 24: 369-420. Mottier, D. M. ’07, Ann. Bot. 21: 309-347. Strasburger, E. ’08, Jahrb. Wiss. Bot. 45: 478-570. Mottier, D. M. ’og, Ann. Bot. 23: 343-352. Zimmermann, A. ’93, Beitr. Morph. Physiol. Pflanzenzelle 2. . Schniewind-Thies, J.’o01, Die Reduktion, etc. Jena. Strasburger, E. ’04, Sitzungsber, Kon. Preuss. Akad. Wiss. 18: 587-614. Strasburger, E. ’05, Jahrb. Wiss. Bot. 42: I-71. Miyake, K. ’05, Jahrb. Wiss. Bot. 42: 83-120. Digby, L. ’o09, Ann. Bot. 23: 491-502. Digby, L. ’r0, Ann. Bot. 24: 727-757. Farmer, J. B., & Digby, L. ’10, Rep. Brit. Assoc. Adv. Sci. Sheffield. ee L. ’84, Ann. Sci. Nat. Bot. VI. 17: Sick E. ’80, Zellbildung und Zell- teilung. Jena Kérnicke, M. on Sitzungsber. Niederrhein. Ges. Natur- und Heilkunde Bonn. 4, III. Miiller, C. ’10, Jahrb. Wiss. Bot. 47: 99-117. Bonnet, J. ’12, Arch. Zellforsch. 7: 231-241. 582 PICARD: BIBLIOGRAPHY CONVALLARIACEAE Asparagus officinalis Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. Salomonia biflora, Cardiff, I. D. ’06, Bull. Torrey Club 33: (Polygonatum biflo- 271-306. rum Convallaria Strasburger, E. ’84, Neue Untersuchungen tiber den Befructungsvorgang bei den Phanerogamen, etc. Jena. Wiegand, K. ’99, Bot. Gaz. 28: 328-359. Wiegand, K. ’oo, Bot. Gaz. 30: 25-47. Lerten J. ’o1, Die Reduktion, ete. “ena Berghs, Pe ibe La Cellule 22: 41-54. Vagnera (Smilacina) Lawson, A. A.’11, Trans. Roy. Soc. Edin. 47: 591-604. Trillium Atkinson, G. F. ’99, Bot. Gaz. 28: 1-26. Ernst, A. ’02, Flora g1: 1-46. Grégoire, V., & Wygaerts, A. ’04, La Cellule 21: 5-76. AMARYLLIDACEAE Alstroemeria Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. Guignard, L. ’84, Ann. Sci. Nat. Bot. VI. 17: 5-59. Galanthus nivalis Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. Guignard, L. ’91, Ann. Sci. Nat. Bot. VII. 14: 163-296. Leucojum Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. Went, F. ’87, Ber. Deuts. Bot. Ges. 5: 247- Scrastrese E. ’88, Uber Kern- und Zell- teilung, etc. Hist. Beitr. 1. Jena. Guignard, L. ’91, Ann. Sci. Nat. Bot. VII. 14: 163-296. Strasburger, E. ’95, Jahrb. Wiss. Bot. 28: 151-204. Strasburger, E. ’00, Uber Reduktionsteilung, ete. Hist. Beitr. 6. Jena. MORNE eta bh rie ACS PICARD: BIBLIOGRAPHY 583 Narcissus Went, F. ’87, Ber. Deuts. Bot. Ges. 5: 247- 258. IRIDACEAE Tris Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. Strasburger, E. ’95, Jahrb. Wiss. Bot. 28: 151-204 Strasburger, E. ’oo, Uber Reduktionsteilung, etc. Hist. Beitr. 6. Jena. Kérnicke, M. ’o1, Sitzungsber. Niederrhein. Ges. Natur- und Heilkunde Bonn. 4, III. Strasburger, E. ’05, Jahrb. Wiss. Bot. 42: I-71. Miyake, K. ’05, Jahrb. Wiss. Bot. 42: 83-120. Gladiolus hybrid Metcalf, H. ’o1, Proc. Nebraska Acad. Sci. 7: : 109. ORCHIDALES—ORCHIDACEAE Listera Guignard, L. ’84, Ann. Sci. Nat. Bot. VI. 17: 5-59- Guignard, L. ’85, Ann. Sci. Nat. Bot. VI. 20: 310-372. Guignard, L. ’91, Ann. Sci. Nat. Bot. VII. 14: 163-296. Rosenberg, O. ’05, Bot. Not. 1905: 1-24. Orchis Guignard, L. ’82, Ann. Sci. Nat. Bot. VI. 14: 26-45. Strasburger, E. ’95, Jahrb. Wiss. Bot. 28: 151-204. Limodorum abortivum Guignard, L. ’97, Ann. Sci. Nat. Bot. VIIL. 6: 177-220. SCITA MINALES—MUSACEAE Musa - Tischler, G. ’10, Arch. Zellforsch. 5: 622-670. CANNACEAE Canna indica Wiegand, K. ’oo, Bot. Gaz. 30: 25-47. Kérnicke, M. ’o1, Sitzungsber. Niederrhein. Ges. Natur- und Heilkunde Bonn. 4, ITI. ANGIOSPERMAE—DICOTYLEDONES CASUARINALES—CASUARINACEAE Casuarina Juel, H. O. ’03, Flora 92: 284-293. URTICALES—MorackEaE (URTICACEAE) Morus indica Tahara, M. ’o9, Bot. Mag. Tokyo, 23: 343-353. 584 PICARD: BIBLIOGRAPHY Tahara, M. ’10, Bot. Mag. Tokyo, 24: 281-298. Cannabis sativa Strasburger, E. ’10, Jahrb. Wiss. Bot. 48: 428-520. URTICACEAE a dioica, Elato- ee E. ’10, Jahrb. Wiss. Bot. 47: 245-2 SA setingues sein aa ca Viscum album Guignard, L. ’85, Ann. Sci. Nat. Bot. VI. 20: 310-372. POLYGONALES—POLYGONACEAE Rumex Patientia, Strasburger, E. ’80, Zellbildung und Zellteil- Rheum undulatum ung. Jena. Fagopyrum esculentum Stevens, N. E. ’12, Bot. Gaz. 53: 277-308. CHE NOPODIALES—NYCTAGINACEAE Mirabilis hybrid Tischler, G. ’08, Arch. Zellforsch. 1: 33-151. PORTULACACEAE paksicass virginica Cardiff, I. D. ’06, Bull. Torrey Club 33: 271- 306. CHENOPODIACEAE Beta Strasburger, E. ’80, Zellbildung und Zellteil- ung. Jena. CARYOPHYLLACEAE Melandrium rubrum Strasburger, E. ’10, Jahrb. Wiss. Bot. 48: 427-520. RA NALES—NYMPHAEACEAE Nymphaea alba, Nu- Guignard, L. ’97, Ann. Sci. Nat. Bot. VIII. phar luteum (Nym- 6: 177-220. pbhaea lutea) Lubimenko, W., & Maige, A. ’07, Rév. Gén. Bot. 19: 404-425; 433-458; 474-505- Nymphaea alba Strasburger, E. ’oo, Uber Reduktionsteilung. etc. Hist. Beitr. 6. Jena. RANUNCULACEAE Clematis recta ‘Guignard, L. ’85, Ann. Sci. Nat. Bot. VI. 20: 310-372. Trollius europaeus Lundegardh, H. ’o9, Sv. Bot. Tidskr. 3: 78- 124. Helleborus foetidus _ Strasburger, E. ’88, Uber Kern- und Zellteil- ung, etc. Hist. Beitr. 1. Jena Mottier, D. M. ’97, Jahrb. Wiss. Bot. 30: 169-204. sires toe PICARD: BIBLIOGRAPHY 585 Mottier, D. M. ’98, Jahrb. Wiss. Bot. 31: 125-158. Overton, J. B. ’05, Jahrb. Wiss. Bot. 42: 121- 153. Berghs, J. ’05, La Cellule 22: 141-160. Thalictrum purpuras- Overton, J. B. ’09, Ann. Bot. 23: 19-61. cens Aconitum Napellus Overton, J. B. ’05, Jahrb. Wiss. Bot. 42: 121-153. Paeonia spectabilis Overton, J. B. ’93, Vierteljahr. Naturf. Ges. Zirich 38. MAGNOLIACEAE Magnolia Guignard, L. ’97, Ann. Sci. Nat. Bot. VIII. 6: 177-220. Magnolia, Lirioden- Andrews, F. M. ’o2, Beih. Bot. Centralbl. 11: dron 134-142. BERBERIDACEAE Podophyllum peltatum Mbottier, D. M. ’97, Jahrb. Wiss. Bot. 30: 169- 204. Strasburger, E. ’00, Uber Reduktionsteilung, etc. Hist. Beitr. 6. Jena. Kérnicke, M. ’o1, Sitzungsber. Niederrhein. Ges. Natur- und Heilkunde 4, III. Mottier, D. M. ’03, Bot. Gaz. 35: 250-282. Mottier, D. M. ’05, Bot. Gaz. 40: 171-177. Overton, J. B. ’05, Jahrb. Wiss. Bot. 42: 121- 153- ; Mottier, D. M. ’07, Ann. Bot. 21: 309-347. CALYCANTHACEAE © Calycanthus floridus | Overton, J. B. ’05, Jahrb. Wiss. Bot. 42: 121-153. Overton, J. B. ’09, Ann. Bot. 23: 19-61. ERATOPHYLLACEAE deol submer- Strasburger, E. ’02, Jahrb. Wiss. Bot. 37: 477-526. PA Pa VERALES—PAPAVERACEAE Corydalis cava Strasburger, E. ’82, Arch. Mikr. Anat. 21: 476-589. CRUCIFERAE Hesperis matronalis Strasburger, E. 80, Zellbildung und Zellteil- ung. Jena. 586 PICARD: BIBLIOGRAPHY Bursa (Capsella), Si- Laibach, I. ’07, Beih. Bot. Centralbl. 22: symbrium, Brassica, I9I-210. Stenophragma, Alys- sum, Iberis, Lunaria Bursa (Capsella),[Zos- Rosenberg, O. ’08, Flora 93: 251-2 59. tera and Calendula briefly, also] SARRACENIALES—DRosERACEAE Drosera Rosenberg, O. ’99, Meddel. Hég. Bot. Inst. Stock. 24: 1-126; ’or, 4": 1-21. Rosenberg, O. ’03, Ber. Deuts. Bot. Ges. 21: I1O-II9. Rosenberg, O. ’04, Meddel. Hég. Bot. Inst. Stock. 6": 1-13 Rosenberg, O. ’04, Ber. Deuts. Bot. Ges. 53- Berghs, J. ’05, La Cellule 22: 139-160. Rosenberg, O. ’06, Kjellman Bot. Stud. 237- 243. Rosenberg, O. ’o09, Kéngl. Sv. Vet.-Akad. Handl. 43:.3-63. Rosenberg, O. ’o9, Sv. Bot. Tidskr. 3: 163. ROSALES—PapILionacEaE (LEGUMINOSAE) Laburnum Strasburger, E. ’07, Jahrb. Wiss. Bot. 44: 482-555. Lathyrus odoratus Gregory, R. P. ’o05, Proc. Cam. Phil. Soc. 13: 148-157. . Phaseolus _ Zimmermann, A. ’96, Morph. und Physiol. pfl. Zellkernes. Jena. Wager, H. ’o4, Ann. Bot. 18: 29-55. Mano, T. Martins ’os, La Cellule 22: 55-7 78. Pisum Strasburger, E. ’80, Zellbildung und Zellteil- ung. Jena. Cannon, W. A. ’o3, Bull. Torrey Club 30: 519-543. Strasburger, E. ’o07, Jahrb. Wiss. Bot. 44: 482-555. Strasburger, E. ’11, Flora 102: 1-23. Vicia Faba Zimmermann, A. ’96, Morph. und Physiol. pfl. Zellkernes. Jena. Strasburger, E. ’00, Uber Reduktionsteilung, etc. Hist. Beitr. 6. Jena. a PICARD: BIBLIOGRAPHY 587 McComb, A. ’oo, Bull. Torrey Club 27: 451- 459. Gardner, B. ’o1, Contr. Bot. Lab. Univ. Pa. 2: 150-182. Karpoff, W. ’04, Unters. aus Mosk. Landwirt. Inst. 1 Karpoff, W. ’04, Trav. Inst. Agron. Moscou. ea H. ’10, Sv. Bot. Tidskr. 4: 174- PE, H. ’10, Jahrb. Wiss. Bot. 48: 285-378. Fraser, H., & Snell, J. ’11, Ann. Bot. 25: 845-855. Lundegardh, H. ’12, Jahrb. Wiss. Bot. 51: 226-280. GROSSULARIACEAE (SAXIFRAGACEAE) Ribes hybrids Tischler, G. ’06, Jahrb. Wiss. Bot. 42: 545- ; 578. ROSACEAE Rosa, Rubus, Alchem- Strasburger, E. ’04, Jahrb. Wiss. Bot. 41: alla 88-164. Rosa Rosenberg, O. ’09, Sv. Bot» Tisdkr. 3: 150- 162. Fragaria elatior Strasburger, E. ee pairs Best., etc. Hist. Beitr. 7. a. Potentilla hybrid Tischler, G. ’08, set Zellforsch. 1: 33-151. GERA NIALES—TROPAEOLACEAE Tropaecolum majus Strasburger, E. ’80, Zellbildung und Zell- teilung. Jena. RUTACEAE Dictamnus albus Strasburger, E. ’82, Arch. Mikr. Anat. 21: 89. Bde tine E. ’88, Uber Kern- und Zell- teilung, etc. Hist. Beitr. 1. Jena. Strasburger, E. ’07, Jahrb. Wiss. Bot. 44: 482-555. Osawa, ’12, Jour. Coll. Agr. Tokyo, 4: 83-116. EUPHORBIACEAE Strasburger, E. ’10, Jahrb. Wiss. Bot. 48: 428-520. Citrus Imp. Univ. Mercurialis annua 588 PICARD: BIBLIOGRAPHY Malte, M. O. ’10, Embryologiska och cyto- logiska undersékningar 6fver Mercurialis annua. Diss. Lund. SAPINDALES—ACERACEAE Acer Cardiff, I. D. ’06, Bull. Torrey Club 33: 271- 06. Darling, C. A. ’o09, Bull. Torrey Club 36: 177- 199. Darling, C. A. ’12, Bull. Torrey Club 39: 407- 410. MALVALES—MALVACEAE Gossypium hybrid Cannon, W. A. ’03, Bull. Torrey Club 30: 133-172. Balls, W. L. ’10, Ann. Bot. 24: 653-665. Lavatera Byxbee, E. S. ’oo0, Proc. Calif. Acad. Sci. III. 2: 63-82. PARIETALES—PAssIFLORACEAE Passiflora caerulea Williams, C. L. ’99, Proc. Calif. Acad. Sci. III. 1: 189-206. THY MELEALES—THYMELEACEAE Daphne, Gnidia, Wick- Strasburger, E. ’o9, Zeitpunkt Best., etc. stroemtia indica Hist. Beitr.7. Jena. MYRTALES—ONAGRACEAE Onagra (Oenothera) Gates, R. R. ’07, Bot. Gaz. 43: 81-115. and Oenothera Geerts, J. M. ’07, Ber. Deuts. Bot. Ges. 25: I9I-195. Lutz, A. M. ’07, Science, II. 26: 151- 152. Gates, R. R. 08, Bot. Gaz. 46: 1-34. Geerts, J. M. ’08, Ber. Deuts. Bot. Ges. 26: 608-614. Geerts, J. M. ’09, Réceuil Trav. Bot. Néer- landais 5: 93-209. Lutz, A. M. ’og, Science, II. 29: 263-267- Gates, R. R. ’09, Bot. Gaz. 48: 179-199. Gates, R. R. ’o0g, Arch. Zellforsch. 3: 525-552- Davis, B. M. ’o09, Ann. Bot. 23: 551-5713 "10, 24: 631-651. Gates, R. R. ’10, Proc. Int. Zool. Cong., Cambr. Mass. Davis, B. M. ’113, Ann. Bot. 25: 941-974- Ate Koen PICARD: BIBLIOGRAPHY 589 Gates, R. R. ’11, Bot. Gaz. 51: 321-344. Gates, R. R. ’12, Ann. Bot. 26: 993-1010. PRIM ULALES—PRIMULACEAE Primula Gregory, R. P. ’09, Proc. Cam. Phil. Soc. 15: Digby, L. ar Ann. Bot. 26: 357-388. GENTIANALES—OLEACEAE Syringa hybrid Juel, H. O. ’00, Jahrb. Wiss. Bot. 35: 626- 657- Tischler, G. ’08, Arch. Zellforsch. 1: 33-151. GENTIANACEAE Gentiana procera Denniston, R. H. ’13, Science, II. 37: 383- 384. ASCLEPIADACEAE Asclepias syriaca Stevens, W. C. ’98, Kansas Univ. Quart. 7: 77-85. Strasburger, E. ’o1, Ber. Deuts. Bot. Ges. Ig: 450-461 Gager, C. S. i Ann. Bot. 16: 123-148. POLEMONIALES—POLEMONIACEAE Cobaea scandens Lawson, A. A. ’98, Proc. Calif. Acad. Sci. III. 1: 169-188. ; SOLANACEAE Solanum tuberosum Mano, T. Martins ’o05, La Cellule 22: 55-75. SCROPHULARIACEAE - Pedicularis sylvestris Guignard, L. ’84, Ann. Sci. Nat. Bot. VI. 17: BIGNONIACEAE Bignonia venusta Duggar, B. M. ’99, Bull. Torrey Club 26: 89-105. RUBIALES—RUvBIACEAE Houstonia coerulea Stevens, N. E. ’12, Bot. Gas: 53: 277-308. CAPRIFOLIACEAE Sambucus nigra Went, F. ’87, Ber. Deuts. Bot. Ges. 5: 247-258. Strasburger, E. 88, Uber Kern- und Zell- teilung, etc. Hist. Beitr. 1. Jena. ADOXACEAE Adoxa Moschatellina© Lagerberg, T. ’06, Kjellman Bot. Stud. 80-88. CAMPANULA ee Kirkwood, J. E. ’07, Bull. Torrey Club 34: 221-242. 590 PicARD: BIBLIOGRAPHY Cucurbita Pepo Zacharias, E. ’95, Flora 81: 217-266. Zimmermann, A. ’96, Morph. und Physiol. Pfl. Zellkernes. Jena. Bryonia hybrid Tischler, G. ’06, Ber. Deuts. Bot. Ges. 24: 83-96. Tischler, G. ’08, Arch. Zellforsch. 1: 35-151. CAMPANULACEAE Campanula grandis Overton, J. B. ’o05, Jahrb. Wiss. Bot. 42: “2421-152. ' Overton, J. B. ’09, Ann. Bot. 23: 19-61. Micrampelis Kirkwood, J. E. ’07, Bull. Torrey Club 34: 221-242. CICHORIACEAE and COMPOSITAE Silphium Merrell, W. D. ’00, Bot. Gaz. 29: 99-133. Tanacetum vulgare Rosenberg, O. ’05, Bot. Not. 1905: 1-24. Antennaria Juel, H. O. ’00, Kéngl. Sv. Vet.-Akad. Handl. Crepis 33°: 1-57; ’05, 30%: 1-21. Rosenberg, O. ’09, Sv. Bot. Tidskr. 3: 64-77. Beer, R. ’12, Ann. Bot. 26: 705-726. Mieracium Juel, H. O. ’o5, Kéngl. Sv. Vet.-Akad. Handl. 391: I-21. Rosenberg, O. ’07, Sv. Bot. Tidskr. 1: 398- 410. Rosenberg, O. ’07, Bot. Tidskr. 28: 144-170. Taraxacum Juel, H. O. ’04, Ark. Bot. 24: 1-9. Juel, H. O. ’o5, Kéngl. Sv. Vet.-Akad. Handl. 39‘: I-21. - Rosenberg, O. ’o9, Sv. Bot. Tidskr. 3: 150- 162. Achillea Millefolium, Lundegardh, H. ’o9, Sv. Bot. Tidskr. 3: 78- Calendula officinalis, 124. Anthemis, Matricaria Chamomilla Calendula officinalis, Beer, R. ’12, Ann. Bot. 26: 705-726. Anthemis, Matricaria Chamomilla Chondrilla juncea Rosenberg, O. ’12, Sv. Bot. Tidskr. 6: 915- gI9. Tragopogon pratensis, Beer, R. ’12, Ann. Bot. 26: 705-726. Doronicum plantagi- ' neum CORNELL UNIVERSITY, ITHACA. INDEX TO AMERICAN BOTANICAL LITERATURE (1910-1913) The aim of this index is to include all current botanical literature written by Americans, published in America, or based upon American material ; the word Amer- ica being used in the broadest sense. Reviews, and papers that relate exclusively to ‘hoe, agriculture, horticulture, manufactured products of vegetable origin, or snboratory methods are not included, and R a ' some important particular. If users of the Index will call the attention of the editor to errors or omissions, their kindness will be appreciated. This Index is reprinted monthly on cards, - furnished in this form to subscribers at the rate of one cent for each card. Selections of cards are not permitted ; each subscriber must take all cards published Fass pe term of hie subscription, Corre- spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club, Albert, F. La Tuia jigante. (Thuya plicata.) Bol. Bosques Pesca i Caza 1: 586-591. Mr 1913. [Illust.] Alway, F. J. Studies on the relation of the non-available water of the soil to the hygroscopic coefficient. Nebraska. Agr. Exp. Sta. Research Bull. 3: 5-122. f. 1-37- 25 Je 1913. Ames, A. A consideration of structure in relation to genera of the Polyporaceae. Ann. Myc. 11: 211-253. pl. 10-13. Je 1913. Ames, O. Orchidaceae: Illustrations and studies of the family Orchidaceae. The genus Habenaria in North America. Fascicle i-xiv + 1-288. pl. 60-79. “Boston, 1910. Ami, H. M. Preliminary lists of the organic remains occurring in the various geological formations comprised in the map of the Ottawa District, including formations in the provinces of Quebec and Ontario, along the Ottawa river. Ann. Rep. Canada Geol. Surv. II. r2: 51G-77G. D 1901. Andres, H. Zwei neue Pirolaceae aus der Subsection Erxlebenia (Opiz) H. Andres nebst einigen Bemerkungen zur Systematik der hei- mischen Arten. Verhandl. Bot. Ver. Brandenburg 1912: 218-227. Sind, Be] AOTs. Pirola paradoxa sp. nov. from Washington. Andrews, F. M., & Ellis, M. M. Some observations concerning the reactions of the leaf hairs of Salvinia natans. Bull. Torrey Club 40: 441-445. Au 1913. 591 592 INDEX TO AMERICAN BOTANICAL LITERATURE Arthur, J. C., & Kern, F. D. The rediscovery of Peridermium pyri- forme Peck. Science II. 38: 311, 312. 29 Au 1913. Babcock, E.B. A new variety of Juglans californica Watson. Science II. 38: 89, 90. 18 JI 1913. Bachmann, F. M. The origin and development of the apothecium in Collema pulposum (Bernh.) Ach. Archiv Zellforschung 10: 369- 430. pl. 30-36. 8 Jl 1913. Becker, M.A. The preservation of our wild fowers. Jour. N. Y. Bot. Gard. 11: 169-175. Jl 1910. Berry, E. W. A fossil flower from the Eocene. Proc. U. S. Nat. Mus. 45: 261-263. pl. 21. 13 Je 1913. Black, C. A. The morphology of Riccia Frostii Aust. Ann. Bot. 27: 511-532. pl. 37, 38. -Jl 1913. Chrysler, M. A. The origin of the erect cells in the phloem of the Abietineae. Bot. Gaz. 56: 36-50. f. 1-12. 16 Jl 1913. Cockerell, T. D. A. A wine-red sunflower. Science II. 38: 312, 313- 29 Au 1913. Collins, G. N. Mendelian factors. Science II. 38: 88, 89. 18 J! 1913. Collins, G. N., & Kempton, J. Inheritance of waxy endosperm in hybrids of Chinese maize. IV*. Conférence Internat. Génétique, Paris 191%: 347-357. 1913. Cook, M. T., & Taubenhaus, J. J. The relation of parasitic fungi to the contents of the cells of the host plants. (II. The toxicity of vegetable acids and the oxidizing enzyme). Delaware College Agr. Exp. Sta. Bull. 97: 3-53. f. 1-43. 1 J! 1912. Copeland, E. B. Some ferns of northeastern Migtsaas Leaflets Philip. Bot. 5: 1679-1684. 28 Je 1913. Includes Angiopteris Elmeriana, Cyathea Warihon, C. dimor photricha, C. cinerea, Dryopteris urdanetensis, and Athyrium propinquum, spp. nov ; Copp, G. G. The protection of native plants. Jour. N. Y. Bot. Gard. 7: 26-29. F 1906. Cosens, A. A contribution to the morphology and biology of insect galls. Univ. Toronto-Stud. Biol. 13: 297-387. pl. 1-13. 1912. Crandall, C. S.! Mosquitoes pollinating orchids. Science II. 38: 51: 11 Jl 1913. Darbishire, O. V. Lichens collected during the 2d Norwegian Polar Expedition in 1898-1902. 1-68. pl. 1, 2, Kristiania 1909. Rep- Sec. Norw. Arct. Exped. ‘‘Fram” 1898-1902. No. 21. Davis, B. M. Genetical studies on Oenothera—IV. The behavior of INDEX TO AMERICAN BOTANICAL LITERATURE 593 hybrids between Oenothera biennis and O. grandiflora in the second and third generations. Am. Nat. 47: 449-476. f. I-15. Au 1913; 547-57 ka Fe £05 FF 1901S, Davis, B. M. The problem of the origin of Oenothera Lamarckiana De Vries. New Phytologist 12: 233-241. f. 7. 26 Jl 1913. Eastham, J. W. Useful wild plants of Canada. Ottawa Nat. 27: 40-43: 12 Jl 1913. Ellis, M. M. Seed production in Yucca glauca. Bot. Gaz. 56: 72-78. 16 Jl 1913. Elmer, A. D. E. A few new Polygalaceae. Leaflets Philip. Bot. 5: 1671-1678. 21 Je 1913. Includes Securidaca atro-violacea multiramosum, and X. subglobosum. Elmer, A.D.E. New Anonaceae. Leaflets Philip. Bot. 5: 1705-1750. 19 Jl eS New speci ibed in Unona (4), Uvaria (4), Xylopia (1), Saccopetalum (1), Polyalthia (8), Phaeanthus (1), Oxymitra (2), Orophea (3), Mitrephora (4), Meiogyne (2), Goniothalamus (3), Deprananthus (1), and Artabotrys (2). Elmer, A. D. E. Palawan Acanthaceae. Leaflets Philip. Bot. 5: 1685-1704. 3 Jl 1913. Includes one new species each in Strobilanthus, Eranthemum, Dicliptera, Ruellia, Hallieracantha, and Lepidagathis; two new species in Hypoestes and three in Gymno- stachyum, Evans, A. W. Notes-on North American Hepaticae. IV. Bryologist 16: 49-55. f. 1-7. Jl 1913. Includes Cololejeunea setiloba sp. nov. Fairchild, D. The discovery of the chestnut bark disease in China. Science IT. 38: 297-299. 29 Au 1913. Fawcett, H. S. Citrus scab (Cladosporium Citri Massee). Univ. Florida Agr. Exp. Sta. Bull. 109: 51-60. f. 24-31. My 1912. Fawcett, H. S. Stem-end rot of Citrus fruits. (Phomopsis sp.). Univ. Florida Agr. Exp. Sta. Bull. 107: 3-21. f. 1-9. O 1911. Ferdinandsen, C. Fungi terrestres from northeast Greenland (N. of 76° N. Lat.) collected by the ‘‘Danmark-expedition’” 1906-08. Meddelelser om Grénland 43: 137-145. pl. 9. 1910. Includes Calvatia arctica Ferdinandsen & Winge sp. nov. Floyd, B. F., & Stevens, H. E. Melanose and stem-end rot. Univ. Florida Agr. Exp. Sta. Bull. 111: 3-16. f. 1-9. D 1912. Forbes, S. A., & Richardson, R.E. Studies on the biology of the upper Illinois River. Bull. Illinois State Lab. Nat. Hist. 9: 481-574. pl. 65-85. Je 1913. vy Py ieee es | | ; [ | ee floriferum xX £ £ 594 INDEX TO AMERICAN BOTANICAL LITERATURE Gallée, O. Lichens from northeast Greenland (N. of 76° N. Lat.) collected by the ‘‘ Danmark-expedition’’ 1906-08. Meddelelser om Gronland 43: 183-191. I9QI0. Gallte, O. Saxifragaceae 2. The biological leaf-anatomy of arctic species of Saxifraga. Meddelelser om Grénland 36: 239-294. f. I-29. 1910. Gleason, H. A. The relation of forest distribution and prairie fires in the middle West. Torreya 13: 173-181. Au 1913. Gow, J. E. Observations on the morphology of the aroids. Bot. Gaz. 56: 127-142. f. I-47. 14 Au 1913. Hamet, R. Sedum Carnegiei, a new species of the family Crassuiaceae from the herbarium of the Carnegie Museum. Ann. Carnegie Mus. 8: 418-420. Mr 1913. Harper, R. M. A botanical cross-section of northern Mississippi, with notes on the influence of soil on vegetation. Bull. Torrey Club 40: 377-399. pl. 21, 22. Au 1913. Harris, J. A. Note on the alpine dwarfing of Polygonum Bistorta. Torreya 13: 182-184. f. 7. Au 1913. Harris, J. A. On the-relationship between the number of ovules formed and the capacity of the ovary for maturing its ovules into seeds. Bull. Torrey Club 40: 447-455. f. 1, 2. Au 1913. Hassler, E. Apocynaceae. [In Ex herbario Hassleriano: Novitates paraguarienses XVIII.] Repert. Sp. Nov.12: 257-264. 20J11913- Includes Aspidiosperma Rojasii and A. Quirandy Hassler, spp. nov. and several new varieties of the latter species. Hassler, E. Malvaceae. [In Ex herbario Hassleriano: Novitates para- guarienses XVIII.] Repert. Sp. Nov. 12: 264-269. 20 Jl 1913- Includes Malvastrum guaraniticum, Sida rupicola, and S. margaritensis spp- OV: Hassler, E. Oecnotheraceae—II. [In Ex herbario Hassleriano: Novi- tates paraguarienses XVIII.] Repert. Sp. Nov. 12: 269-278. 2° Ji 1913. Includes 54 new varieties, forms, or subspecies. Hayden, A. An ecological study of a prairie province in central Iowa. Proc. Iowa Acad. Sci. 18: 55, 56. 1911. An abstract. Heald, F. D., & Studhalter, B.A: Preliminary note on birds as carriers of the chestnut blight fungus. Science II. 38: 278- 280. 22 Au 1913. Hesselbo, A. Mosses from northeast Greenland (N. of 76° N- Lat.) INDEX TO AMERICAN BOTANICAL LITERATURE 595 collected by the ‘‘ Danmark-expedition”’ 1906-08. Meddelelser om Grénland 43: 171-180. pl. 11, 12. 1910. Includes Bryum Myliusii sp. nov. Hibbard, R. P. The antitoxic action of chloral hidieie’ upon copper sulphate for Pisum sativum (Preliminary contribution). Centralb. Bakt. Zweite Abt. 38: 302-308. 30 Jl 1913. Hitchcock, A. S. Mexican grasses in the United States National Her- barium. Contr. U.S. Nat. Herb.17: 181-389+vii-xiv. 15 Jl 1913. Includes 23 new species in Andropogon (2), Aristida (2), Campulosus (4), Des- champsia (1), Lasiacis (1), Melica (1), Muhlenbergia (1), Paspalum (3), Poa (4), Senites (1), Sorghastrum (1), Sporobolus (1), Syntherisma (1), Trisetum ( 1), Trinochloa (1), and Tristachya (1). Holden, R. Contributions to the anatomy of Mesozoic seatices - No. I. Jurassic coniferous woods from Yorkshire. inn. Bot. 27: 533- 545. pl. 39, 40. Jl 1913. Howard, W. L. An experimental study of the rest period in plants. Univ. Missouri Agr. Exp. Sta. Research Bull. 1: 5-105. Ap 1910, Hutchinson, J. Parthenium argentatum A. Gray. Hooker's Icones Plantarum. IV. 10: pl. 2998. p. 1-3. Jl 1913. Hutchinson, J. Podachaenium eminens. Curt. Bot. Mag. IV. 9: $l. 8502. Jl 1913. A plant from Central America. Jones, L.R. A plea for closer interrelations in our work. Science II. 38: 1-6. 4 Jl 1913. Knupp, N. D.. The flowers of Myriophyllum spicatum L. Proc. lowa Acad. Sci. 18: 61-73. pl. 1-4. I9I1I. Lacy, M.G. A discussion of the results obtained by crossing Zea Mais L. (Mais Djagoeng) (—Reanu luxurians Dur.—teosinte) and Eu- chlaena mexicana Schrad. Am. Nat. 47: 511, 512. Au 1913. Lind, J. Systematic list of fungi (Micromycetes) from northeast Green- land (N. of 76° N. Lat.) collected by the ‘‘ Danmark-expedition” 1906-1908. Meddelelser om Grénland 43: 149-162. pl. 10. I9gI0. Includes Ascospora veges Hendersonia gigantea, Coniothyrium Lesquerellae, and Pyrenophora filicina, spp. n Lindly, J. M. Flowers of ae County. Proc. Iowa Acad. Sci. 18: 19-24. I9QII. Livingston, B. E. Climatic areas of the United States as related to plant growth. Proc. Am. Philos. Soc. 52: 257-275. pl. 9-11. Ap 1913. 596 INDEX TO AMERICAN BOTANICAL LITERATURE Loesener, T. Mexikanische und zentralamerikanische Novitaten. IV. Repert. Sp. Nov. 12: 217-244. 25 Je 1913. Includes new species as follows: pitts Daagoeame H. Gross, Coccoloba oaxacensis H. Gross, Pisonia linearibracteat erl, Heliocarpus Caeciliae Loes., utilon Seisdigason Ulbrich, Sphaeralcea eahat Ulbrich, Hauwya Donnellsmithit cle: H. longicornuta Loes., Xylopleurum deserticolum Loes., and Cordia Langlassei Loes. Macallum, A. B. Surface tension and vital phenomena. Univ. -Toronto Stud. Physiol. 8: 3-82. pl. 7. 1912. BSc in English in revised form from an article “‘ Oberflachenspannung und Lebense nungen,”’ in vol. 11, Ergebnisse der Physiologie. McBeth, 3 S Cellulose as a source of energy for nitrogen fixation. U.S. Dept. Agr. Plant. Ind. Circ. 131: 25-34. 5 Jl 1913. Macbride, T. H. Notes on Iowa saprophytes. Proc. Iowa Acad. Sci. 18: 57-60. I9DI. Marie-Victorin, . Découverte du Lycopode petit-cypres dans les Laurentides. Naturaliste Canadien 39: 166-170. My 1913. Martin, J. N. The physiology of the pollen of Trifolium pratense. Bot. Gaz. 56: 112-126. f. r 14 Au 1913. Matthew, G. F. A new flora in the older Palaeozoic rocks of southern New Brunswick, Canada. Trans. Roy. Soc. Canada III. 6: 83-99- Be te 2. tka. Melhus, I. E. The powdery scab of potato (Sbongaspora Solani) in Maine. Science II. 38: 133. 25 Jl 1913. Merrill, G.K. New and interesting lichens from the state of Washing- ton. Bryologist 16: 56-59. Jl 1913. Includes Bialora myriocarpella sp. nov. Meyer, R. Einiges iiber Echinocactus longihamatus Gal. und seine Varietaten. Monats. Kakteenk. 23: 91-93. 15 Je 1913- Meyer, R. Uber Echinocactus haematacanthus Monv. Monats. Kakteenk. 23: 94-96. 15 Je 1913. Mickleborough, J. A report on the chestnut tree blight, the fungus, Diaporthe parasitica Murrill. 1-16. f. 1-4. Harrisburg. My 1909- [Illust.] Pennsylvania Department of Forestry. Miller, F. A. Breeding medicinal plants. Am. Jour. Pharm. 85: 291- 301. f. 1-4. Jl 1913. Nathorst, A. G. Contributions to the carboniferous flora of north- eastern Greenland. Occurrence of the plant-fossils. Meddelelser om Grénland 43: 339-346. pl. 15, 16 +f. 1-4. 22 Mr 1911. INDEX TO AMERICAN BOTANICAL LITERATURE 597 Nichols, G. E. Summer evaporation intensity as a determining factor in the distribution of vegetation in Connecticut. Bot. Gaz. 56: 143-152. 14 Au 1913. Orton, W. A. The development of disease-resistant varieties of plants. IV°. Conférence Internat. Génétique, Paris 1911: 247-265. f. I-9. 1913. Ostenfeld, C. H. Marine plankton from the East-Greenland Sea (W. of 6° W. Long. and N. of 73° 30’ N. Lat.), collected by the ‘‘ Dan- mark-expedition” 1906-1908—I. List of diatoms and flagellates. Meddelelser om Grénland 43: 259-285. f. I-11. 1910. Ostenfeld, C. H., & Lundager, A. List of vascular plants from north- east Greenland (N. of 76° N. Lat.), collected by the “ Danmark- expedition”’ 1906-1908. Meddelelser om Grénland 43: 1-32. fl. 1-6 +f. I-3. 1910. Ostrup, E. Diatoms from northeast Greenland (N. of 76° N. Lat.), collected by the “ Danmark-expedition”’ 1906-08. Meddelelser om Grénland 43: 195, 196-256. pl. 13, 14. 1910. Pammel, E. C., & Clark, C. Studies in variation of red clover. Proc. Iowa Acad. Sci. 18: 47-54. 1911. [Illust.] Pammel, L. H. Some fungus diseases of trees. Proc. lowa Acad. Sci. 18: 25-33. 1911. [Illust.] Pammel, L. H., & King, C. M. Pollination of red clover. Proc. lowa Acad. Sci. 18: 35-45. 1911. [Illust.] Pennell, F. W. Studies in the Agalinanae, a subtribe of the Rhinan- thaceae. Bull. Torrey Club 40: 401-439. Au 1913. Includes descriptions of 5 new species. Prain, D., & Hutchinson, J. Notes on some species of Acalypha. Kew Bull. Misc. Inf. 1913: 1-28. f. 1-46. Mr 1913. Quehl, L. Allerlei aus dem Kakteenkasten. Monats. Kakteenk. 23: 93, 94- 15 Je 1913. Reimer, F. C., & Detjen, L. R. Self-fertility of the scuppernong and other muscardine grapes. North Carolina Agr. Exp. Sta. Bull. 209: 5-23. f. I-13. S 1910. Ricker, P. L. Directions for collecting plants. U. S. Dept. Agr. Plant Ind. Circ. 126: 27-35. f. 1-5. 10 My 1913. Rolfs, P. H., Fawcett, H. S., & Floyd, B. F. Diseases of citrus fruits. Univ. Florida Agr. Exp. Sta. Bull. 108: 27-47. f. 10-23. N 1911. Rolfe, R. A. Stanhopea convoluta. Curt. Bot. Mag. IV. 9: pl. 8507. Au 1913. A plant from Colombia, South America. 598 INDEX TO AMERICAN BOTANICAL LITERATURE Saxton, W. T. The classification of conifers. New Phytologist 12: 242-262. f. I. 26 Jl 1913. Selby, A.D. A brief hand-book of the diseases of cultivated plants in Ohio. Bull. Ohio Agr. Exp. Sta. 214: 307-456 + i-vii. f. 1-106: Mr 1910. Shannon, C. W. The trees and shrubs of Oklahoma. Oklahoma Geol. Surv. Cire. 4: 1-41. Mr 1913. [Illust.] Shear, C. L., & Stevens, N.E. The chestnut-blight parasite ( Endothia parasitica) from China. Science II. 38: 295-297. 29 Au I913. Simon, C. E., & Wood, M.A. On the inhibitory action of certain anilin dyes upon bacterial development. Proc. Soc. Exp. Biol. & Medicine 10: 176-178. 21 My 10913. Smith, J. D. Undescribed plants from Guatemala and other Central American republics. Bot. Gaz. 56: 51-62. 16 Jl 1913. Twenty new species are described. t Smith, J. D., & Rose, J. N. A monograph of the Hauyeae and Gongylocarpeae, tribes of the Onagraceae. Contr. U. S. Nat. Herb. 16: 287-298. f. 45-54. 23 Au 1913. Includes ‘Burragea and X ylonagra gen. nov. and Hauya Rusbyi sp. nov. Stevens, F. L., & Hall, J.G. Diseases of economic plants. °i-x 35 513. f. I-219. New York, 1910. Stout, A. B. Tomato-nightshade chimeras. Jour. N. Y. Bot. Gard. 14: .145-150. pl. t2z. Au 1913. Surface, F. M. |The result of selecting fluctuating variations, data from Illinois corn breeding experiments. IV°. Conférence Inter- nat. Génétique, Paris rort: 222-255. 1913. Swingle, W. T. Chaetospermum, a new genus of hard-shelled citrous fruits. Jour. Washington Acad. Sci. 3: 100-102. f. 7. 19 F 1913- Swingle, W.T. Le genre Balsamocitrus et un nouveau genre voisin, Aeglopsis. Bull. Soc. Bot. France 58: (Mém.) 225-245. pl. se +f. , B. Mr 1912. ae Balsamocitrus gabonensis sp. nov. and Aeglopsis Chevalieri gen. et sp. NOV- ‘Swingle, W. T. Observations sur les quelques espéces. indo-chinoises des genres Atalantia et Glycosmis. -Notulae Syst. 2: 158-163. f. J: 25 Mr rogrz2. Swingle, W. T. Variation in first generation hybrids (imperfect dominance): its possible explanation through zygotaxis. IV. Con- férence Interna. Génétique, Paris 1911: 581-594. f. I-10. 1913: Buti. ToRREY CLUB VOLUME 40, PLATE 23 MortieR AND NOTHNAGEL: CHROMOSOMES OF ALLIUM CERNUUM VOLUME 40, PLATE 24 BuLi. TORREY CLUB MorttieR AND NOTHNAGEL: CHROMOSOMES OF ALLIUM CERNUUM Vol. 4C No. 11 BULLETIN OF THE TORREY BOTANICAL CLUB NOVEMBER, 1913 Edward Lyman Morris EpwWarp B. CHAMBERLAIN (WITH PORTRAIT) The sudden death of Edward L. Morris on September 14, last, is a loss not easy to estimate. He was by preference a student of systematic botany, but long experience as a successful teacher, and active work as a museum curator, gave him a breadth of scientific training and an appreciation of popular and scientific points of view that are not often combined in those whose position makes them the interpreters of science to the public. Those who knew Mr. Morris feel that their personal loss over- shadows everything else. He had to an unusual degree the genial charm of manner that makes and retains friends, and a loyalty that takes no account of time or effort spent in helpful service. His daily life was so full of patience, cheerfulness, and sympathy, that few realized how great, at times, was his own burden. Even when his own life was full of trouble, he was never without the characteristic cordial greeting for everyone he met. With high ideals and the strong convictions that accompany them, he was frank in the expression of opinion, but at the same time modest in statement and considerate of the views of those from whom he differed. The affectionate respect of his associates and the codperation they gave him ee striking testimonials to his own personality. In his own work Mr. Morris set for himself a severe standard, and demanded like faithfulness from others; yet he worked with [The BuLLETIN for October (40: 520-598. pl. 23, 24) was issued 15 O 1913.] 599 600 CHAMBERLAIN: EDWARD LyMAN Morris an enthusiasm that enlivened dry detail and routine. Whatever he did showed painstaking method, sincerity of purpose, and devotion. As a colleague has written of him, ‘‘He put his heart as well as his conscience into his work.’’ He was a patient in- vestigator and a close observer, but willing to defer any conclusion until he had a first-hand knowledge of the facts. This desire for truthfulness made him chary of publication; there was so often some minor point that required more study for a complete under- standing of the case. The facts of Mr. Morris’s life are, briefly, as follows:—He was born at Monson, Mass., October 23, 1870, the son of Edward Franklin Morris and Louise Janette Clapp, and his youth was spent in the vicinity of his birthplace. At as early an age as eleven, he began the systematic study and collection of the plants of the township, continuing his collecting throughout his pre- paratory school course.. Entering Amherst College in the autumn of 1888 from Monson Academy, he was given special credit for the botanical work already done, and an opportunity to continue it in conneetion with the college museum. During his third and fourth college years he had especial privileges for advanced work in botany and zodlogy, and was in charge of the college museum. Obtaining the bachelor’s degree from Amherst in 1891, he spent one year at the Museum of the Worcester Natural History Society, one year in graduate study at Harvard, and two years as instructor at Amherst, which conferred upon him the degree of M.A. in 1895. Of his work at Amherst Professor Tyler says, in a letter re- cently received: ‘It always seemed as if he was working purely for the enjoyment of it. He was a very hard worker, and made his students work. The best men did so because they caught his spirit, the others because they had to. He made things very clear, and always knew what to tell and how much to leave to the student to find out. He never made his teaching a mere memorizing of dry details.”’ In 1895 Mr. Morris removed to Washington, D. C., and for twelve years was connected with the school system of that city, for the last seven years being head of the department of biology- His progressive teaching here developed a course in biology . = CHAMBERLAIN: EDWARD LYMAN Morris 601 that reached the life and activities of young people. He always taught as if in the laboratory, arousing the interest of his stu- dents, stimulating them to seek first-hand knowledge, teaching them that biology was a matter of everyday life, and training them to think clearly and hard. During the year 1907, Mr. Morris resigned his position in Washington to become curator of natural science in the Museum of the Brooklyn Institute of Arts and Sciences, a position which he held at the time of his death. For one year, also, after the resignation of Dr. Lucas, he was acting curator-in-chief. This position gave him the chance to develop the ideas of the educative value of museum collections that he had long held. Believing that exhibition specimens should always arouse the desire for further information, he not only saw that the books furnishing such knowledge were close at hand, but insisted that every visitor should have, if he desired, the opportunity of personal conversation with the curator. In addition to professional duties, Mr. Morris found the time to take an active part in general scientific work, his own motto for such things being, ‘‘Make both tie for first place.” In Washington he was a member of the botanical, biological, and entomological societies, and of the Cosmos Club, but most closely identified with the Washington Biologists’ Field Club, of which he was a founder and leading spirit. No one who knew him at ‘“‘The Island” could fail to see how very close to his heart the success of the Club was, or ‘forget how enthusiastically he entered into the plans for its development. Even after leaving Washington he kept in close touch with all that went on at the Club, and never failed to revisit it when circumstances permitted. He had made a large collection of plants from the Club property, which he hoped to make the basis for a detailed catalogue. For years Mr. Morris’s especial pleasure had been the syste- matic study of the Plantaginaceae, of which he had accumulated a large amount of material, many species being represented by alcoholic as well as ordinary herbarium specimens. It was a source of keen regret to him that increasing duties encroached upon the leisure hours that he preferred to spend upon his col- lections. The characteristic desire for accuracy delayed the publication of many conclusions that had already been attained, 602 CHAMBERLAIN: Epwarp LyMAN Morris conclusions that it is hardly possible to reconstruct from the notes available. Besides what has already been mentioned, Mr. Morris was for a short time associate editor of School Science and an associate examiner on the College Entrance Examination Board. At the time of his death he was editor of the Torrey Botanical Club, of which he had been a member since 1901. In 1898 he collected for the United States National Herbarium on the Florida Keys, and in 1900 was an assistant upon the staff of the 3 United States Fish Commission in West Virginia. For four years he was a special plant expert of the Department of Agriculture, doing field work in Oregon, along the Great Lakes, and in Iowa. In 1908 Mr. Morris was secretary of the Nomenclature Com- mission of Section G of the American Association for the Advance- ment of Science, and in 1911 was elected a fellow of the Association. Mr. Morris was twice married, his first wife being Florence Syvret, of Charlton, Mass., who died in 1903. In 1907 he married Mary E. Bedell, of Washington, D. C., who, with a son, survives him. | NEw York City BIBLIOGRAPHY The following list contains the titles of all of Mr. Morris’s publications that it has been possible to find in the limited time available. Corrections or additions will be greatly appreciated. . Plant study. Plant World 2: 77. F 1899. 2. A revision of the species of Plantago commonly referred to Plantago patagonica Jacquin. Bull. Torrey Club 27: 105-119. 24 Mr 1900. gyrea, P. dura, P.inflexa, P. brunnea, P. fastigiata, P. tetrantha, spp. nov-; P. aristata Nuttallii subsp. nov.; P. lanatifolia (Coult. & Fish.) Small, P. erecta, P. scariosa, nom. no 4. Baiedeeas hederaceum in America. Proc. Biol. Soc. Washington 13: 157, 158. 13 Je 1900. 4. Some plants of West Virginia. Proc. Biol. Soc. Washington 13: 171-182. 31 O 1900. Polypodium vulgare oreophilum Maxon and Vernonia gigantea pubescens, sub- spp. nov. 5- North American Plantaginaceae—Il. Bull. Torrey Club 28: 112- 122. pl. 12. 2 Mr 1901. 7 P. verticillata, P. picta, P. oblonga, P. ignota, P. speciosa, P. obversa, spp. nOV- on) fea - ~ oO lol Ny CHAMBERLAIN: EDWARD LYMAN Morris 603 . A correction of Vernonia gigantea pubescens. Proc. Biol. Soc. Vernonia maxima pubescens nom. nov. Plantago septata E. L. Morrissp. nov. In Britton, N. L., & Rydberg, An enumeration of the flowering plants collected by R. S. Williams and J. B. Tarlton. Contributions to the botany of the Yukon Territory 4. Bull. N. Y. Bot. Garden 2: 182. 27 My 1901. Botanizing in and around a lake. Plant World 4: 109, 110. Je ICol. . Plants for the aquarium. School Science 1: 95. 1901-2. - “Occasional ’’ leaves of Trillium. Plant World 5: 92, 93. pl. 13. My 1902. . Abnormal Trilliums. Plant World 6: 87-89. Ap 1903. [Illust.] . The bush morning- glory. Plant World 7: 109-113. pl. 5, 6. My 1904. Ipomoea lepiophyila Torr . Plantago (Plasiavinelia) coelorhiza Morris & Macloskie, n.s. In Macloskie, G. Flora of Patagonia. Reports of the Princeton University Expeditions to Patagonia 8: 734. pl. 25 C. S 1905. North American Plantaginaceae—III. Bull. Torrey Club 36: 515-530. 1 O 1909. P. xerodea and P. pusilla Engelmannii, nom. nov. . The nomenclatural authority for Gonionemus Murbachii. Proc. Biol. Soc. Washington 22: 179-182. 30 O 1909. . Herbarium suggestions. Torreya 11: 145-149. f. I-3. 19 Jl 1911. . The germination of cat-tail seeds. Torreya 11: 181-184. f. I, 2 12 $ 1911. . [Review of] Stewart’s Botanical survey of the apie i Islands. Torreya 11: 222, 221. 18 O 1911. - Museum cata’ogues. Proc. Am. Assoc. Museums 5: 35-38. IQII. . [Review of] Dinsmore’s Plants of Palestine. Torreya 12: 34-36. 15 F 1912. - An apparently new record for Rubus Chamaemorus Linnaeus. orreya 12: 88. 17 Ap I9I2. . The possibilities of botanical exhibits. Proc. Am. Assoc. Mu- S ums 6: 105-108. ICI2. - The museum point of view in botany. Proc. Am. Assoc. Museums 7: 83-85. 1913. The ferns and flowering plants of Nantucket—xXI EUGENE P. BICKNELL CELASTRACEAE *CELASTRUS SCANDENS L. Rare; it is found sparingly on Coskaty entwined with wild rose and red raspberry near the harbor shore, and in Shawkemo, where it is massed thickly along a low bank back of the beach. Flower buds June 4, 1909; first flowers June 2, I9II. ACERACEAE ACER RUBRUM L. In swamps and low grounds. It is commonly of no greater stature than a shrub, but in sheltered thickets becomes a well- developed tree, and in Beechwood has attained a height of not less than thirty to thirty-five feet, the trunks thirty to thirty-five inches in basal girth. The leaves of different trees are widely vari- able, sometimes appearing much like those of Acer carolinianum, again taking an elongated form with narrowly cleft attenuate and sharply cut lobes. *ACER CAROLINIANUM Walter. Frequent in boggy thickets, sometimes side by side with those forms of Acer rubrum with which it is most sharply in contrast. It is a tree of marked individuality when appointed in its true features but these are not always well expressed and its divergence from the red maple cannot be said to have passed into a fixed separation. In its most characteristic forms the small thickish leaves, rather clustered at the ends of the branchlets, are of rounded outline and broadly notched into three short lobes, the blades, only 4-6 cm. long and wide, having the upper surface of a dark shining green, the lower surface peeroruuely whitened and more or less pubescent. 605 606 BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET *ACER SACCHARINUM L. A. dasycarpum Ehrh. | About two miles from the town scattered among an open growth of pines off the Wauwinet road are to be found a few small silver maples not over three or four feet in height. They were first observed in 1909. *ACER PLATANOIDES. The Norway maple has been little used on Nantucket, but is self-seeded in planted grounds and has occasionally grown up into small trees in neglected places. *ACER PSEUDO-PLATANUS L. 2 Many fine sycamore maples shade the streets of the town and produce a numerous progeny.of seedlings some of which persist and grow into small treesin out of the way places. In full flower along the streets June 6, 1909. The three introduced maples here mentioned have only the slenderest claim to be included in the island’s wild flora and are reported mainly for purposes of. record as widely cultivated trees which are tending to become naturalized. BALSAMACEAE IMPATIENS BIFLORA Walt. Common in low grounds often bordering thickets or growths of rankly growing taller plants. A form occurs having very pale: or whitish spotted flowers. First flowers July 1; 1912; blooms through September. VITACEAE Vitis Lasrusca L. Very common generally and in many places conspicuous from its luxuriant growth. It thrives in low thickets draping the shrubbery and strays into open places, trailing among the grass and herbaceous plants or even sprawling in bare sandy fields. Flower buds June 12, 1909; first flowers June 17, 1908; June 18, 1910; well-formed green fruit June 27, 1912. The fruit may be of the largest size and deep purple or amber purple in color, or much smaller, more numerous and crowded BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 607 in the clusters, and greenish or greenish purple even when fully ripe. This pale-fruited form is locally abundant on Marthas Vineyard, where it is often wholly green at maturity and is known to the islanders as the white wild grape. I did not myself see it growing there, but, on Sept. 28, 1911, was shown several large sacks filled with the perfectly ripe green fruit which, in gathering, had been kept separate from the usual purple kind. VITIS AESTIVALIS Michx. Found only on the eastern side of the island, where it is frequent or locally common in wet or dry thickets sometimes actually intertwined with Vitis Labrusca. It occursin Shawkemo, Pocomo, Coskaty, and Squam, and south to Tom Never’s Pond. Comes into flower rather later than V. Labrusca. Flower buds very small June 7, 1911, and June 13, 1908; first flowers June 26, 1910; still in bloom July 11, 1912. It appears to fruit only sparingly on Nantucket, although bearing abundantly on Marthas Vineyard. Fruit small and green Aug. 13, 1906; becoming purplish Sept. II, 1907. On Marthas Vineyard much of the fruit was still unripe Oct. 5, 1912. A very old vine near Abram’s Point measured twenty-one inches around close to the base and seventeen inches a foot above. PARTHENOCISSUS QUINQUEFOLIA (L.) Planch. In thickets, either in low grounds or on the dry plains, some- times trailing over banks clothed with crisp lichens and bearberry or even thriving in exposed white sand. Flower buds _ barely visible June 1, 1909, and June 14, I9II; no open flowers up to July 12, 1912. The leaflets vary from glabrous to thickly pubescent with silvery hairs on the lower surface and to some extent on the upper surface also; this pubescence may extend thinly along the petioles but seems always to be absent from the branchlets and - tendrils. It is a character that has been adduced as distinctive of Parthenocissus hirsutus (Donn) Small, but as to the pubescent - Nantucket plant there seems little reason to doubt that it is merely a condition of the common Virginia creeper. The leaves of young plants are often very pubescent, and in older plants the lower leaves may be pubescent and the later ones quite glabrous. 608 BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET MALVACEAE MALVA ROTUNDIFOLIA L. An abundant weed, flowering freely from May through Sep- tember and doubtless until frost. *MALVA VERTICILLATA L. The herbarium of the Nantucket Maria Mitchell Association contains a specimen of this mallow, collected by Mrs. Nellie F. Flynn, bearing the record ‘‘ Waste place, Sept. 22, 1902.”’ *MALVA MOSCHATA L., Several white-flowered plants in full bloom July 9, 1912, ina vegetable garden at Surfside; Surfside, Aug. 1909, Mrs. Mary A. Albertson; lane off Madequet road, 1905, Mrs. Eleanor W. Morgan, fide F. G. Floyd. *ALTHAEA ROSEA Cav. Freely spontaneous by street sides and in neglected places about the town and appearing occasionally in waste lots in the suburbs. The seedling plants begin to spring up at the end of May. Just in flower in several waste spots July 12, 1912. Histscus MoscHeurtos L. When in bloom the rose-mallow is conspicuous at a number of the shore ponds on the northern and eastern sides of the island, but it seems to be quite wanting about the ponds on the south shore. At most of its localities it is not abundant, although grow- ing in profusion at a few places. It is found at Capaum Pond, Reed Pond, Monomoy, Shimmo, Squam Pond, and on Coskaty, where it was in full bloom Aug. 16, 1906. Flowers observed as late as Sept. II, 1907. Mr. Floyd’s notes refer to a form having white flowers with a crimson eye found by a small pond in Monomoy by Miss Mary Foster Coffin. It is not improbable that this may have been Hibiscus oculiroseus Britton, which is not rare on Long Island. HY PERICACEAE ASCYRUM HYPERICOIDES L. Long known from Nantucket, the northeastern limit of its range, but not at all a scarce plant there, as has been supposed. BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 609 It is, however, confined to the eastern side of the island, where it is locally common from Wauwinet to Quidnet, extending west to Beechwood and south to beyond Sachacha Pond. In full flower Aug. 7, 1906; continues to bloom until late in September. It often spreads out into patches of considerable size, which become noticeable from their light green color as early as the middle of June: HYPERICUM ADPRESSUM Bart. Common about several ponds in Polpis and Saul’s Hills; west of Sachacha Pond; Waqutuquaib Pond; Miacomet Pond; a single early flower July 11, 1912; in full flower Aug. 7, 1906; Sept. I, 1904; some flowers remaining Sept. 18, 1907. The young plants have become several inches high by the middle of June. Early in the season this St. John’s-wort may be seen in small ponds either wholly submerged or showing emersed leafy tips. Later, when the waters have fallen, such plants often develop with unusual vigor, becoming fully two feet high with the leaves pro- portionately enlarged and the submerged portion of the stem greatly thickened with spongy tissue (var. spongiosum Robinson). Where colonies of the plant extend back from a flooded shore a complete gradation may be traced from this spongiose aquatic condition to the more usual terrestrial state. The latter comes earliest into bloom, the most dwarfed examples of the driest Situations flowering first and often precociously. The spongiose tissue is doubtless homologous with the aeren- chyma produced on the floating stems of Decodon verticillatus. A slight but evident spongiose enlargement of the lower part of the stem is sometimes seen in Hypericum canadense and in Hyperi- cum boreale when these low ground plants grow in very wet places. HYPERICUM PERFORATUM L. One of the bright-flowered weeds of fields and waysides, and Scattered widely over the plains and commons. First flowers June 27, 1910; June 29, 1912. Hypericum punctratum Lam. Not common but found sparingly at a number of widely Separated stations, mainly on the eastern side of the island; not 610 BiIcKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET observed west of Maxcy’s and Hummock Ponds. Plants of full size June 15, 1911; small flower buds June 11, 1912; in full flower and with some mature pods Aug. 16,1906. So far as observed, the Nantucket plant has always sessile broadly clasping leaves. *HYPERICUM BOREALE (Britton) Bicknell. This is the commonest Hypericum of the island, abounding in low grounds, damp or wet sandy places, and pond shores. It is sometimes aquatic, inhabiting deep water with the habit of a Callitriche, the elongated leafy stems either wholly submerged or their tips emersed. In wet sand it may become strongly stolonif- erous, putting forth prostrate basal offshoots which reach a length of several inches and root at intervals, sending up small flowering stems and terminating in a cluster of stems from the rooted tip. The young plants become recognizable early in June. Just in flower Aug. 13, 1906, remaining in bloom through September. HYPERICUM MUTILUM L. ; Common in low grounds, often with its characters unusually well emphasized, the broadly clasping leaves becoming as large as 3 cm. long by 2 cm. wide. The earliest leaves are observable at the end of May and the young plants take definite form early in June. In full flower Aug. 13, 1906; flowering through September. *HyPERICUM MAjus (A. Gray) Britton. Infrequent, growing in damp places. West and southwest of the town; Trot’s Swamp; Miacomet Pond; Quaise. Just in flower Aug. II, 1906; in full flower Sept. 8, 1904; Sept. 12, 1907. HYPERICUM CANADENSE L., Common in low grounds and wet sandy places. Leaves often almost filiform linear. Plants very small May 30, 1909; a single early flower June 20, 1908, and July 3, 1912; in full flower and with mature capsules Aug. 13, 1906; continues in flower through September. *Hypericum dissimulatum sp. nov. Erect, often from an oblique or horizontal rooting base, commonly 1.5-3 dm. high, exceptionally up to 5.5 dm., not often BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 611 branched below the middle; leaves narrowly oblong, obtuse, sessile or subclasping, 3-5-nerved, 1-3 cm. long, 2-6 mm. wide; branches slender, openly ascending, bearing dichotomous many- flowered bracteolate cymes, the bracts subulate; sepals oblong to lanceolate, obtuse or acutish, equaling or shorter than the cap- sules; capsules greenish to reddish purple, small, 2-4 mm. long, ellipsoid to conic-ovoid. Maine to Maryland and North Carolina. Type from Nan- tucket, damp roadside west of the town, Sept. 20, 1899, in flower and fruit, in herb. N. Y. Botanical Garden. Also collected on Nantucket Sept. 8, 1904, Miacomet Pond, and Sept. 9, 1904, near the town. This plant has been known to me for many years, having been collected first in York County, Maine, then on Nantucket, on Marthas Vineyard, where it is more common than I have found it elsewhere, and on Long Island. It is found in -damp sandy places, usually growing with H. canadense, H. majus, H. mutilum, and H. boreale, one or all, and is not less distinct in appearance from each of them than are they among themselves. It differs from H. canadense in broader often subclasping leaves, _ more diffuse inflorescence, and smaller often ellipsoid capsules. Narrower leaves, more spreading and compound inflorescence, and smaller capsules distinguish it readily from H. majus, while it stands apart from H. mutilum by stricter, less branched habit, narrower less clasping leaves and longer, or more ellipsoid, purple capsules. Certain specimens approach H. canadense in the form and color of the pods, other examples seem nearer to H. mutilum, and it may well be questioned whether it be not a hybrid of these . two species or, indeed, partly of H. canadense and H. boreale as some specimens might seem to suggest. But all of our small St. John’s-worts of this group are nearly related and, considering the extended coastwise range of H. dissimulatum, as good reasons appear for viewing it as one of a chain of close species as for surmising that it may be a cross. In addition to material from Maine, Nantucket, Marthas Vineyard, and Long Island, collected by myself, the following specimens may be cited: In herb. N. Y. Botanical Garden: — RHODE IsLaAnD: Kingston, Aug. 21, 1906, E. S. Reynolds. 612 BIcKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET PENNSYLVANIA: Smithville, Lancaster County, J. K. Small & Io Fe Carter. MARYLAND: Hyattsville, Aug. 13, 1904, H. D. House. NorTH CAROLINA: Mica, June, 1898, C. W. Hyams. In herb. Columbia University: New York: Springfield, L. I., 1896, Elizabeth G. Knight; New Dorp, Staten Island, Aug. 31, 1890, N. L. Britton. SAROTHRA GENTIANOIDES L. Abundant in dry sandy places, stems appearing June 15, 1911; in full flower in September. The plant may be actually minute, its simple stem bearing only a single flower, or densely branched to form a firm convex mass I-1.5 dm. in diameter. TRIADENUM VIRGINICUM (L.) Raf. Very common in wet swamps and about the borders of muddy ponds. Earliest leaves May 31, 1908; no flowers remaining in September. ELATINACEAE ELATINE AMERICANA (Pursh) Arn. Common in some of the sandy ponds, growing in shallow water near the shore.. Observed especially in Maxcy’s Pond, Miriam Coffin Pond, and Miacomet Pond. At Maxcy’s Pond on Sept 12, 1907, it grew as profusely on the damp sand where the water had receded as beneath the surface along the shore. At one spot in heavy mud ten yards or more from the water’s edge it had formed compacted moss-like mats, some of them six inches across, a mode of growth remarkably unlike that of the submerged plant. Correlated with this difference in habit ran a variation in characters which was brought out strikingly by comparison of the living plants. In water and on damp sand the individual plants were separate in growth, uniformly simple-stemmed, and whitish or pale green in color. In the mud form the matted stems were often divergently much branched and the general color a lively green tinged with reddish or purple, these tints deepening on the cap- sules into bright crimson; instead of greenish white the petals were rose color and were sometimes as large as 1.5 mm. in breadth. The capsules, some being four-valved, were larger than those of BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 613 the submerged plant and of a distinctly different form, depressed- subglobose and wider than long instead of broadly obovoid and longer than wide; actual measurements were 2 mm. wide by I mm. long in the terrestrial plant and only I-1.5 mm. wide by I mm. long or more in the normal aquatic form. : CISTACEAE CROCANTHEMUM CANADENSE (L.) Britton. Helianthemum canadense Michx. The typical plant is not common and is rather local in its distribution, giving place to the following, which is everywhere abundant. It is however frequent in the oak barrens towards Siasconset and is found sparingly in Quaise, on the plains towards the south shore, on Great Neck and elsewhere. No flower buds visible June 3, 1909; first flowers June 11, 1909; in full flower June 19, 1910; a few flowers remaining June 30, 1912. Reduced petaliferous flowers are often produced in September. *Crccanthemum dumosum sp. nov. Similar to Crocanthemum canadense but lower and of more branched and spreading habit, commonly diffuse and semi- prostrate or ascending, the pubescence somewhat more densely and softly canescent, intermixed with scattered non-stellate longer hairs and some minute glandular hairs of a reddish color; leaves smaller and shorter than those of C. canadense and of a more bluish green color, mostly oval and elliptic and obtuse, often very small and crowded on the short divergent branchlets; flowers slightly paler than in C. canadense; mature calyx often larger, the sepals very broad and mostly acuminate, usually bearing reddish papillae on the outer surface and reddened glandu- lar or viscid hairs in the pubescence; primary inflorescence an ascending succession of single petaliferous flowers succeeded by rather numerous flowers intermediate in size and character be- tween these and the later apetalous ones. Well marked and abundant all over Nantucket, combining with such common and characteristic island plants as Amelanchier nantucketense, Ilex fastigiata, and Linum intercursum to stamp the flora with a signally distinctive character. It is found also on Marthas Vineyard and on the Hempstead Plains of Long Island. 614 BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET Blooms rather earlier than C. canadense. First flowers May 31, 1909, and quite generally in bloom June 1; June 3, 1911; still some flowers June 26, 1910. At one station a number of clus- tered plants bore flowers so pale in color as to appear almost white. | Type from Nantucket, Sept. 21, 1899, in herb. N. Y. Botanical Garden. The typical form of the plant has an unlikeness to typical Crocanthemum canadense greater than appears between some other closely allied species within the genus, and this diversity of aspect becomes especially striking when, as is sometimes the case, the two are found growing near together. Typical C. canadense is a taller erect plant with lighter-colored stems and longer and more slender and simple ascending branches, narrowly oblong or oblanceolate leaves tapering to the base and the acute apex, brighter green on the upper surface and less densely pubes- cent. Ordinarily it holds very true to these characters, showing little tendency to marked variation. In several instances where the two plants growing near together allowed a close comparison of the open flowers, those of C. dumosum were seen to be notably the larger, the acuminate sepals reaching a length of 8-10 mm. and reddened with glandular hairs and papillae, while those of C. canadense, narrower and mostly obtuse, were but 5—7 mm. long and only obscurely if at all glandulose. These differences are not, however, always so well marked. Nevertheless C. dumosum is evidently a strongly established derivative of C. canadense, even if it be not yet wholly disconnected from that species. It has been a recurring source of confusion to not a few Nantucket collectors and it seems altogether expedient to dispose of it as 4 stumbling block by giving it identity by a name. *CROCANTHEMUM MAJus (L.) Britton. Helianthemum majus B.S.P. Rather common on the plains towards the south shore; else- where very local although widely scattered, but wanting over 4 great part of the north and east sides of the island. No visible flower buds June 22, 1910, July 2, 1912; first flowers July 10, 1912- small petaliferous flowers sometimes appear in September. BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET .615 *Crocanthemum propinquum Bicknell. Helianthemum propinquum Bicknell. Rather local, but not uncommon in dry open places or along sandy roadways through pine barrens. Common on Marthas Vineyard. In full bloom June 26, 1910; not many flowers left June 29, 1912; a few belated flowers July 11, 1912. This plant, not at all uncommon from Nantucket to western Long Island and doubtless further south, appears to remain almost unknown to botanists and seems not to have been reported by any collector since it was first described in Britton’s Manual over eleven years ago. In the seventh edition of Gray’s Manual it has been quite misunderstood, being mistaken for the plant described in this paper as Crocanthemum dumosum and referred to as being probably only a stunted form of C. canadense. I know the plant now much better than when I ventured to give it a name and have found no reason to doubt that it is an unequivocal species, that is to say, one that is organically discrete from those allied species which most nearly approach it, however. close the degree of their relationship. Narrow indeed is the interval between this plant and those other convergent species whose distribution it partly shares. But I have not found in this any proof of consanguinity but rather an example of the exceeding closeness in which specific lines may run in perfect security from coalescence or entanglement. The plant is to be viewed critically especially in its relation to C. majus. Its. clustered primary flowers at once give this indication and mark its distinctness from C. canadense. Singularly enough, however, in the later stages of its growth it more nearly resembles the latter, agreeing in color of foliage and slender ascending branches surpassing the primary inflorescence. This character of the mature plant sketches it out clearly from C. majus, of strict habit and short close branches, but in its unbranched early- flowering stage, then also of paler foliage, it is almost a reduced counterpart of the larger plant. To review its differences from Crocanthemum majus, it is a much smaller and more slender and flexuous plant, at length more openly and slenderly branched, less densely canescent from the first and finally much greener, the leaves narrower and more 616 BIcKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET obtuse, often spatulate-linear, and usually on more obvious petioles; the primary inflorescence is of more delicate and open structure, the flower buds elliptic in form rather than ovoid, the calyx becoming notably larger, 8-10 mm. long, and often strongly reddish-tinged, thus equaling in length the largest calices of C. canadense as well as corresponding in color, although with narrower sepals and wanting the characteristic pilose hairs; the narrow outer sepals are shorter than in C. majus, and the even _smaller petals are of rather a brighter yellow; the primary capsules are smaller, thinner-walled, and less broadly ovoid, longer than wide instead of wider than long, and are without the umbonate tip; it is, in fact, much more like the capsule of C. canadense, although smaller and narrower; the papillose seeds are also much like those of C. canadense. There is nothing in all this that denotes the plant to be neces- sarily of mixed strain, nor do my observations lead me to believe that it is a hybrid. It does indeed possess in combination the early flowering time of Crocanthemum canadense and the smaller pale yellow flowers of C. majus, together with the slender branch- ing of the one and the clustered petaliferous flowers of the other, yet its capsule has not its counterpart in that of either, nor is it intermediate with them, being smaller and less broadly ovoid. The plant stands apart from these companion species also in its small size and more delicate structure, in the prevailing form of the leaves and in its non-cespitose habit. Its slender stems, although sometimes loosely clustered, commonly arise at distinct, even remote intervals along tortuous elongated rootstocks, forming open groups or larger patches, sometimes several feet in diameter. It is rarely found associated with more than one of its close allies, often, indeed, occupying territory where not either one of & others is found at all. The relationship of Crocanthemum propinquum to the little- known C. georgianum of the southern states is evidently close, although, according to Dr. Small, the latter possesses the very distinct character, as compared with the northern group of species, of bearing pe petaliferous and apetalous flowers in the same clusters. SON SENSES Soa anon ae BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 617 HUDSONIA ERICOIDES L. Few plants of Nantucket spread over the island more widely or in greater abundance than this little heathlike species and not one is more conspicuous in the landscape when in full bloom. Nor is there any other that, at flowering time, puts its scene in color with quicker transformation, for there come seasons when it bursts into bloom on all sides in the hours of a single hot morning. Earliest flowers May 30, 1909, quite generally in bloom June 2; first flowers June 4, 1911, in the early morning, everywhere in flower by noon; abundantly in bloom June 7, 1908, inflorescence becoming brown by the 13th and but few flowers remaining on the 18th; in the season of 1910 it had passed flowering in exposed places June 20, although still blooming freely in the shade of pine groves. After full bloom it remains for one or two weeks the season’s most conspicuous flower, spreading its sheets of gold along the roadways and over acres of plain and hillside, a radiant sight. A few days later the flowers are withered and the wide tracts that had glowed with their color become brown and rusty as if seared by fire. In open sandy places where this plant has formed the compact circular cushions that are one of its modes of growth, the flowers usually open first close to the ground on the side towards the morning sun, blending together in patches of expanding brightness as they continue to unfold. Gradually as the sun rises overhead the glow of color creeps back along the borders of the tuft, some- times uniting around its circumference in a golden ring. Soon afterwards the entire tuft has become an unbroken mass of bloom. Often in midsummer these cushion-like tufts even in the hottest and most exposed sandy spots remain fresh and green in bright contrast to their parched surroundings, calling to mind so remote a comparison as the stones along a woodland brook covered with green moss. In open pine scrub south of the town on June 5, I9I1, several patches of this plant, all near together, bore flowers of palest sulphur-yellow, in striking contrast to the normal bright yellow flowers everywhere about them. 618 BIcKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET HUDSONIA TOMENTOSA Nutt. Very abundant, blanketing the dunes and reaches of white sand back of the beaches and occurring on sandy exposures all over the island. It is sometimes found in association with the preceding but seems not to mix readily with any other plant. Close to blooming June 3, 1911; in full flower June 7, 1908, June 15, 1910, and some flowers remaining June 27; last flowers July 2, 1912, on the exposed ocean front at Siasconset, where many plants flower later than in more protected parts of the island. Ordinarily it begins to flower a little earlier than Hudsonia ericoides. LECHEA MINOR L. Abundant on the eastern side of the island from Wauwinet to Saul’s Hills and Siasconset, extending west. to Shawkemo and through the South Pasture to Surfside; not seen on the western side of the island. The season’s shoots a few inches high June 23, 1910. LECHEA VILLOSA EIl. Much less common than the preceding but like it restricted mainly or entirely to the eastern side of the island, having a scattered distribution from Wauwinet to Siasconset and . the South Pasture and from Pocomo to Shawkemo and Saul’s Hills. LECHEA MARITIMA Leggett. One of the island’s most common plants, appearing every- where in dry sandy soil, even to the tops of Saul’s Hills. It makes its best growth in pure sand, where it becomes widely branched and densely canescent. In less simple soils amid the low vegetation of the moorland or in partial shade it is more thinly canescent and shorter-branched, having a narrower panicle and closer inflorescence. Such forms take on a likeness to Lechea juniperina that seems almost to shadow the origin of that more northern species. Sometimes on rising ground in open growths of pines or other trees it may become very slender and greener, with more scattered leaves and branches, more slenderly branched and open panicle of longer-pedicelled flowers and rather larger fruiting calyx—var. interior Robinson. In full flower Sept. 3, 1904, Sept. 11, 1899; small new shoots BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 619 June 15, 1910. In the autumn, sometimes as early as September, the basal shoots may be found beneath the surface of the sand so densely invested with white pubescence as to appear as if coated with hoar frost. *LECHEA LEGGETTII Britton & Hollick. L. moniliformis Bicknell. Not rare on the eastern side of the island from Reuwindt to Polpis, Gibbs’ swamp and Tom Never’s swamp; one station near Madequecham Pond on the south shore. It is found in low grounds spreading to dry sandy levels near wet places; in one instance it grew on the border of a sphagnum bog, and in another in wet soil along a brackish marsh. Plants 6 inches high June 24, 1910; flower buds well advanced Aug. 7, 1906; some mature pods Aug. 31, 1904. This, in its extreme phase, is the plant described by me some years ago as Lechea moniliformis. The type specimens, as well as others like them from Long Island, mark a pronounced departure from typical L. Leggettii. Other specimens from Nantucket and Long Island are less distinctive and I am in doubt whether it is well to rate the plant as other than a variety of the common species. Nevertheless, it has points of distinction which need no second glance to impress any one who may be familiar with the common inland form of the species, for L. moniliformis would appear to be a plant of the coastal plain, and there is as yet no evidence that it does not belong exclusively among our coastal plain species. Moreover it shows this difference in habits from the more inland plant of dry open places and hilly ground, that it is of low grounds often of wet and brackish soils. A better knowledge may show that its distinctive name should be restored, but for the present let it be merged with L. Leggettit. I take to be typical of the latter the plant that I used to find among the hills and rocky outcroppings along the Hudson near New York and which, found also in New Jersey and on Staten Island, largely made up the material studied by Leggett and by Britton & Hollick. - compared with this = _ distinguishing characters of L. e the slender a gated flowering branchlets and the markedly secund and moniliform inflorescence, for in the 620 BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET i typical plant the ultimate inflorescence takes a short corymbulose _ rather than a slenderly racemulose plan. These flexuous branch- lets are often borne on short spreading branches crowded on the upper part of the stem, producing a broadly ovoid or obovoid, often dense and very leafy panicle instead of a more oblong and open one. The leaves, similar to those of L. Leggetti proper, are rather longer and more tapering acute and narrowed into more evident petioles 1-2 mm. long. The mature calyx and pod is commonly larger and more elliptic than in the typical plant and is usually further distinguished by its decidedly purplish color; also the capsule is rather more exserted and often more distinctly short-stipitate, and the general pubescence is sparser and of rather longer and looser hairs. Note.—Lechea racemulosa Lam. was attributed to Nantucket by Mr. Leggett and is reported by Mrs. Owen as having been found there by Mr. Dame. There would seem to be little reason to doubt that these records were based on mistaken determinations. VIOLACEAE VIOLA PEDATA L., The commonest blue-flowered violet of Nantucket, broadcast on the plains and commons and among open growths of scrub pines. The flowers are often small for the species and of deep color, varying to pale lilac and sometimes pure white. The spring flowers of Nantucket are late in coming, and this violet, which on Long Island colors acres of the Hempstead Plains from April, in early seasons, until the middle of May or; in later seasons, till the end of the month, is commonly in full bloom on Nantucket from late in May until after the middle of June. In the forward season of 1908 no flowers were to be found after June 15, but the following year children on their way t0 school were seen carrying large bunches on June 6, and it was blooming in profusion as late as June 12. Flowers are occasionally produced in midsummer and, more frequently, in September. On Sept. I, 1904, among scrub pines where, earlier in the yea! fire had passed, destroying the herbage, many of these violets had sprung up afresh and were in full bloom. The leaves of all differed curiously from their normal form, being narrowly to BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 621 broadly cuneate and flabellately cleft into irregular lobes of varying length and breadth. Plants with similar leaves collected June 12, 1908, were rooted deep in heavy yellow sand and, like those of the burned-over tract, had doubtless suffered some disturbance of their normal course of growth. *VIOLA OBLIQUA Hill. Viola affinis LeConte. See Bull. Torrey Club 40: 261-270. 1913. On a shaded bank at Watts Run, an abundant growth, and sparingly in a not distant thi€ket in Squam; also in the shade of a willow by a bog hole west of Trot’s Swamp. In full flower as late as June 9, 1909. Becoming 3.5 dm. high, or more; leaves thin, from narrowly to broadly cordate-ovate, attenuate to acuminate, acute, in. age widely dilated at base and broader than long, the largest 9 cm. wide, the upper surface with some minute appressed hairs; sepals ovate to ovate-lanceolate, obtuse, flowers often becoming upturned; peduncles of apetalous flowers of very variable length even on the same plant, declined, ascending or sometimes strictly erect and over 1.5 dm. high; capsules mostly blotched with purple, sometimes pale, the expanded valves 7-10 mm. long; seeds pale. *VIOLA PAPILIONACEA Pursh. Viola cucullata of authors, not Aiton. See Bull. Torrey Club 40: 261-270. 1913. Found only in a boggy meadow about a mile west of the town, growing sparingly with Viola lanceolata; in full flower June 1, 1909. Plants rather small, somewhat tufted from multicipital root- stocks; scapes mostly not longer than the leaves; leaf blades cor- . date-ovate to triangular-cordate, crenulate-serrate, thinly pubes- cent on the upper surface with appressed silvery spiculae; sepals narrowly lanceolate, sometimes elongate, ciliolate; flowers pale blue, or deeper blue, much darker towards the throat. VIOLA LAETECAERULEA Greene. V. papilionacea of authors, in part, not Pursh. See Bull. Torrey Club 40: 261-270. I913. Found only in the town, where it is frequent by streetsides and in shaded yards, often forming close beds, and appearing as 622 BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET if introduced. -In flower June 3, 1909, June 7, 1911. Petioles more or less pubescent dorsally, sometimes densely villous, but more often glabrous, except towards the base of the blade; blades mostly with some pubescence beneath at the base or along the veins. When growing in damp shaded yards the leaves are thinner and brighter green, resembling those of Viola obliqua; plants more deeply set in looser and drier soils have duller leaves of thicker texture, the blades broadly reniform with wide sinus and rounded to the short-pointed apex, the margins more closely erenate-serrate; capsules green. VIOLA FIMBRIATULA Sm. Excepting Viola pedata no other blue-flowered violet is common on Nantucket. Therefore it might be thought that the purity of the fimbriatula line would be wholly uncontaminated, and that variation in the species might be seen in its intrinsic phases free from any influence of hybridization. Nevertheless, the variation shown under this insular seclusion is not less remarkable than is commonly the case elsewhere, where associated species may be supposed to have had their influence. The more common form on Nantucket has ovate-oblong subcordate leaves little if at all incised and often as long as the petioles. A coarser form has longer petioles and larger blades, which become 5 cm. or more wide across the subtruncate base. In bare spots on clayey soil are found very small forms with ovate to ovate-lanceolate subentire leaves narrowed into short petioles and crowded in a close rosette against the ground. In shade among the Miacomet pines there is a form having considerable pubescence but otherwise showing something of the aspect of Viola sagittata, many of the narrow and long-petioled leaves being rather deeply cordate and saliently dentate at the base with upcurved acute teeth and, notwithstanding their pubescence, appearing bright green and shining on the upper surface; the flowers are deep purple with the rather narrow petals often crenulate and obscurely pointed. A series of Nantucket specimens was submitted to Doctor Brainerd who says of them, referring especially to the sagittata- like plants: “Your plants are not strictly hybrids but intermediate BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET 623 forms which have probably resulted from hybridization in the indefinite past. We have no pure Viola sagittaia in Vermont, but most of your odd forms turn up from time to time in the Champlain valley.” *VIOLA FIMBRIATULA Sm. X OBLIQUA Hill. _ Asingle cluster growing with Viola obliqua on a shaded bank at Watts Run, June 9, 1909. Petioles and peduncles pubescent with short spreading hairs, the leaf blades similarly clothed on the veins beneath, sparsely appressed-pubescent on the upper surface, ciliate; later leaves ovate-oblong and openly cordate, or somewhat attenuate-triangu- lar from a subtruncate base, coarsely and rather closely sinuate- dentate towards the base, the largest 6 cm. wide by 9 cm. long on petioles 10-15 cm. long; apetalous flowers few, mostly weakly developed, the buds lanceolate and acute, their petioles ascending- horizontal or declined. This plant grew in shade beside a mass of Viola obliqua. No Viola fimbriatula was found with it, but the specimens are obvi- ously intermediate between these two species, and on Nantucket no others are possibly to be assigned as parents. *VIOLA ODORATA L. An old garden plant of the town, here and there strayed along streetsides and established in neglected yards. One particular tuft has grown for many years in a crevice between the paved sidewalk and a brick wall on North Water Street. It was first noticed in 1899, and has been found at the same spot on every subsequent visit to the town, evidencing both the tenacity of the plant and the undisturbed repose of the town streets. VIOLA LANCEOLATA L. Very common in bogs and low grounds and in wet sandy soil about the borders of ponds. In full flower May 30, 1908, June 3, 1909, June 3, 1911, and still commonly in flower June 15; some flowers remaining June 25, 1910. Plants growing with Viola pallens and appearing more or less intermediate with it, and others approaching Viola primulifolia are not improbably hybrids. 624 BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET *VIOLA PRIMULIFOLIA L. Less common than the ieatic and often found in drier soils. In full flower May 31, 1908, June I, 1909; last flowers June 17, 1910, June I5, I9II. VIOLA PALLENS (Banks) Brainerd. Common in open sphagnum bogs and meadows and in damp thickets. No flowers left May 31, 1908, June 7, 1909; still bloom- ing June 3, 1910, a few last flowers June 8, IgII. A form of distinct appearance was found in several wet sphag- num bogs, especially in one near Shawaukemmo Spring. It is strictly glabrous throughout, the scapes and petioles delicately streaked with pink, the leaf blades unusually thick and veiny, becoming as large as 5 cm. in breadth, and varying in shape from long-ovate and deeply cordate to broadly cordate-reniform; petioles sometimes 9 cm. long; longer peduncles 1.5 dm.; capsules green; seeds 1-1.25 mm. long, dark gray to nearly black when mature. Doctor Brainerd, who has examined specimens, regards it as a form of Viola pallens. Note-—Viola blanda Willd. which proves to be common on Marthas Vineyard is to be looked for on Nantucket. NEw York City The influence of starch, peptone, and sugars on the toxicity of various nitrates to Monilia sitophila (Mont.) Sacc, OTTO KUNKEL INTRODUCTION Winogradsky and Omeliansky (7) found that the addition of .05 per cent of glucose or asparagin to nitrite-media hindered the development of the nitrate-bacteria. Peptone at a somewhat greater concentration was also detrimental. These substances are widely used in the preparation of culture media, but have been given little attention by investigators of the problems of toxicity. Fluri (2) has reported that the salts of aluminum render the protoplasm of Spirogyra and other water plants permeable and are, therefore, injurious to these plants. He found, however, that if glucose, glycerin, or isodulcitol are mixed with the alumi- num salt it loses its power of rendering the protoplasm permeable. Thus the toxicity of aluminum salts to these plants depends upon whether or not glucose, glycerin, or isodulcitol are present in the medium. Quite recently Schreiner and Skinner (4) have reported that the toxicity of the organic poison cumarin is counteracted by phosphates, thus indicating a relation between the organic and the inorganic part of the medium. In view of these results it has seemed worth while to investigate the influence of organic substances on the toxicity of inorganic salts. It has been my object to determine the toxicity of different salts in the presence of certain inorganic substances that are much used in the preparation of culture media. MATERIAL AND METHODS I have used the fungus Monilia sitophila in all of my experi- ments and have found it well suited to my purpose. Some of the things that recommend it are as follows: (1) It is a rapid grower. On a favorable medium at room temperature it will produce spores 626 KUNKEL: INFLUENCE OF STARCH, PEPTONE, AND SUGARS ON in fifteen hours. (2) It is able to use as a partial source of its food supply a rather large number of organic compounds. Went (6) has found that it grows well on media containing any one of the following substances: maltose, trehalose, raffinose, saccharose, cellulose, starch, fats, and proteids. This makes it especially suitable for experiments in which the organic part of the medium is to be varied. All of my cultures were grown in Petri dishes that were pre- viously immersed for at least ten hours in cleaning solution made according to Duggar’s method (1). In the preparation of media and for rinsing glassware redistilled water was used. In the tables that follow, the zero sign indicates that none of the spores on the medium in question had germinated, the minus sign indicates that germination and microscopic growth had taken place, while the plus sign indicates that growth was visible to the naked eye. All cultures were incubated at room temperature (about 22° C.)- In order to determine whether or not the toxicity of various salts to Moniha sitophila is influenced by sugars, starch, or pep- tone, each of these organic substances was used separately in testing the toxicity of the inorganic salts. I have designated the highest concentration of a salt that would. permit germination of the spores in a given medium, as the limit concentration for that medium. A number of preliminary experiments were made to determine the approximate value of the limit concentration for each salt in each medium used. The results obtained in a final set of experiments are shown in the tables given below. EXPERIMENTAL Monilia will produce a considerable growth of mycelium and will ripen spores on a medium made by adding 5 grams of corn- starch to 95 cubic centimeters of redistilled water. The results obtained in a series of experiments in which different inorganic — salts were added -to this medium are shown in TABLE I.. The concentrations varied from 1.33 molar for potassium nitrate to .000004 molar for zinc nitrate. The table shows at a glance the sorte toxicity of the nitrates used, in the order in which they are arranged in the table TOXICITY OF VARIOUS NITRATES TO MONILIA SITOPHILA 627 TABLE I THE TOXICITY OF INORGANIC SALTS TO MONILIA SITOPHILA GROWN ON STARCH MEDIA m Growth after 11 days 1.33 molar solution of potassium nitrate + 5%starch........ _ 1.06 “a “ ‘ ‘ ae + * SUPT Ao ogee aan a 80 “ ‘“ ‘e ‘6 af + hy Sie teary EN a az 53 ‘ “ “ “ ‘6 + *“ Sy Snare ag aa 4% 26 “ “ ‘s “ se + * ee pans ae eke + 13 ‘a “ ‘6 ‘e min + * Pes Rais ee + 667 ‘ ss * calcium <5) + “* PAO SRE Ee Rs a 533 “ “ ‘s ss + “ ice gett a of 400 “ “ “ “ a + “* TR re ase a7 267 “ “ “ “ a + * Seal sik lene ea i 133 + ‘ ‘es “e ss + i 2s Heme Seren Me + .067 “ “ ames “ Be ree a gt ya eae * 034 “ ‘“ ‘a “a as + * pen PANY cate ate = 667 “a “ “ sodium “s + * Raia 4 wauigee te) 533 “ “ “ “8 + * sasha ca pene oO .400 “ “ “a “ sd + * etek SO ae acs = -267 “ “ “ “es ate + ‘ Pies Sea) Sel is 133 “as ‘e ‘“ “a i + e pean erik las ar .067 se “ “ “ s + * ieee Py em a 034 ‘“ se “a “ ts + * Barer gis Saiaa ye ne 187 “ “ “ barium se + * ee cae Stes co) 156 “ “ “ “ “ + ‘“ Pees Sas caattnce = «125 “ “ “ “s es + “ Tie Gone achat a 004 “a “ “ “ as + * ee ee Par aa 062 rv ‘e “a ‘s + * geal ei eho ete + .031 “a “ec ‘a “s ad + ew Foe gad a aeracea ae .007 “a “ ‘s “ $e + * Lu ge: ualgen pa oe .500 “ ‘“ “urea as + “ Eee De phates (0) .250 “ “ “ “ te + * feo pee ae ae fe) 125 “ ‘“ “ ‘e “ + " pe aah gil koe apa f?) .063 ‘ ‘ ‘“ “s “ + * Bete Mike esas ce) .050 és “ “ “ “ + * LA Dae anara nt Ueane n .025 “a “ ‘s “s “ + a ch See tee + hig ae cee See weer ce + 006 “ ‘ es “a st + * neghiits gages rant ae -050 " * “ammonium “ ic ie eens : .038 “ “ 4 + * Lp RR enc ral Gi? ws .025 ‘“ “ ‘“ “ “ + * hi, any ate — .013 ‘ ‘ “ ‘“ a“ + Me pees + .006 ‘ a “ “s “ + * one aioe ogee + -0075 “ “ “ aluminum “ + * eRe See oO = 3 ‘“ “ “ 6s “ + * Lae er eee aie °o 0050 ‘ “ ‘ “s “ + ne Sa ee nae oe ee 18) .0038 ‘a “ “6 ‘“ “ + * Siem cog a ar ehaceee 0 = = 628 KUNKEL: INFLUENCE OF STARCH, PEPTONE, AND SUGARS ON -0006 molar solution of aluminum nitrate + 5% starch........ + -00133 rs * ferric a +." Se ha hears (a) -00067 ips =e sae fe + * ea ee ee os _ -00053 8 ie " 3 = + * Pe Seen Stary ta) -00040 © rf a aes a + * ete va Seay te _- 00027 « = heres a = + * Oe ian Oe Ne + 00013 oy fs fe sss mn + * Sea vo es + 00007 Ai re 5 ae = + * il A cepa ee + 00133 “a: ee * silver sin + ‘* Bae A laces Saat g fe) 00067 ie £3 sy =F ei + * ed ely Se Ts, oe oO 90053.“ Sr + * HES ry RR oe 0 00040 He fe Peas: - + * seis ietie at tits fo) 00027 ste Hi ~ = om + * Perea ne way oO 00013 x ‘fi enc: i + * ida Ranga = 00007 t <é Seinees + * ee Aiea ace eee + 000338 “ ey mate a bt a a ose Sate a Se ha oO 000254 “ ss : = ya + * eas eoearee pune oO ooo169 = * s ee Sere, ¥ + “ ee ss ees 0 000082“ ee NOTE tre i + * Bo neers acs oO 00043“ si ee ss SSE an ee ere fe) 000034 “ wes Rue ie + * steep aE oO oooo25 “* be Rueitons S + * ein bene diag case — oooor7 =“ 2 ies HY + * EE Saae eerie ae ooo008 —* re Pee 6 + “ Pe Neen 4 Bl 000004 “ a pees & + * Pe epee ce eee = 5000 = ia “ammonium tartrate + “ ep A pine fo) 2500 = ee : ~ Zs + * Sar eater a oO 1250 ae Stee, *t + *“ ebro eee to) 0625 = - eS te + ‘* ghe cease ieee fe) 0500 = i pores s + * eee eer fe) -0250 = Me ence 4a fe paras septs ee Shr = 0125 = : haces a + * fr lege a dacaes Be 0063 3 % oe ees @ apt Geena eer oe aon = redistilled water + Ser a we = from potassium to zinc. Monilia made sufficient growth to be easily visible to the naked eye in the starch medium containing potassium nitrate at a concentration of 0.8 molar or less, while at a concentration of 1.33 molar the spores germinated and pro- duced microscopic growth. In a starch medium containing cal- cium nitrate at a concentration of 0.667 molar, the spores germi- nated but produced such a small amount of mycelium that it could be observed only with the aid of the microscope., It is interesting to note that after three days no germination had taken place in this medium. This shows that in such concentrations growth is very slow. In all media containing calcium nitrate at a concentration of 1.33 molar or less, there was abundant growth. TOXICITY OF VARIOUS NITRATES TO MONILIA SITOPHILA 629 When sodium nitrate at a concentration of 0.533 molar was combined with the starch medium no germination occurred. When the concentration was 0.400 molar a small amount of growth was obtained. In still lower concentrations enough mycelium was produced to be easily visible to the naked eye. It was found that growth was much retarded in media containing the higher concentrations of sodium nitrate. This shows that near the limit concentration the rate of growth of Monilia, de- creases as the amount of sodium nitrate is increased. Barium nitrate in the starch medium at a concentration of 0.187 molar inhibits the germination of the spores. At a con- centration between 0.156 molar and 0.094 molar, the spores germinate but produce such a small amount of mycelium that it is not visible except under the microscope. At a concentration of 0.062 molar or less, good growth was obtained. When the concentration of urea nitrate in the starch medium Was 0.125 molar, the spores were unable to germinate. When the concentration was reduced to 0.063 molar, the spores germinated but produced such a small amount of mycelium that it was not visible to the naked eye. Good growth was obtained when the concentration of urea nitrate was 0.05 molar or less. In starch media containing ammonium nitrate at a concen- tration of 0.05 molar none of the spores germinated, but when the concentration was 0.038 molar, they germinated and produced microscopic growth. When the concentration was reduced to 0.025 molar or less, abundant growth occurred. Aluminum nitrate in starch media at a concentration of 0.0025 molar inhibits the germination of the spores. At a concentration of 0.0013 molar the spores germinate and produce microscopic mycelia, while at concentrations of 0.0006 molar or less, good growth is obtained. Ferric nitrate in starch media is quite toxic to the spores of Monilia. In concentrations of 0.00027 molar or less, good gr owth was obtained, but in more concentrated media little or no growth occurred. Silver nitrate is even more toxic than ferric nitrate; when its concentration was 0.00027 molar, the spores germinated, but Produced only microscopic mycelia. In more concentrated media no germination took place. 630 KUNKEL: INFLUENCE OF STARCH, PEPTONE, AND SUGARS ON Zinc nitrate in starch media is far more toxic than any of the other nitrates used; at a concentration of 0.000034 molar, it inhibited the germination of spores. In a concentration of 0.000025 molar, the spores germinated and produced microscopic mycelia, but when the zinc nitrate was used at a concentration of 0.000017 molar or less, abundant growth was obtained. Ammonium tartrate is the only organic salt that was tried in these experiments. In starch media it is slightly more toxic than ammonium nitrate. Ina concentration of 0.025 molar, the spores germinated but produced only microscopic mycelia, while in a like medium containing the same concentration of ammonium nitrate abundant growth was obtained. As shown by TABLE I zinc nitrate is the most toxic substance used in starch media. If its limit concentration be taken as one then the limit concentrations of the other nitrates in the same medium may be expressed, by comparing equimolecular con- centrations, approximately by the following numbers: silver nitrate, 5; ferric nitrate, 26; aluminum nitrate, 52; ammonium nitrate, 1,520; urea nitrate, 1,600; calcium nitrate, 16,560; and potassium nitrate, 53,200. To show at a glance the relative toxic values of the various substances used, they are given in the order of their toxicity in TABLE II. That toxicity does not seem to be related io the valence of the kation is also shown by this table. TABLE II THE RELATIVE TOXICITY OF VARIOUS SALTS IN STARCH MEDIA Valence of kation Toxic concentration Monovalent.......... 133 molar potassium nitrate oh. Re Eee ree 464 : calcium - Monovalent.......... 400 = sodium ch age pee ies 156 “ barium Monovalent.......... 040 : urea $3 Monovalent.......... 038 " monium * onovalent.;.. 0... 025 . ammonium tartrate Trivadlents oon As 0013 in aluminum nitrate eat eR ASS 00067“ ferric z Monovalent.......... 00013 ** silver m8 Divelentcc0 ok: 000025“ zine ef The table shows, in the case of each salt, the greatest com centration at which growth was obtained when the salt was used ? TOXICITY OF VARIOUS NITRATES TO MONILIA SITOPHILA 631 in starch media. A comparison of equimolecular concentrations shows that of all the nitrates used, potassium nitrate is the least toxic and zinc nitrate is the most toxic to Monilia when it is grown in a starch medium. Beginning with the least toxic, the order of toxicity of the nitrates in starch media is as follows: potassium nitrate, calcium nitrate, sodium nitrate, barium nitrate, urea nitrate, ammonium nitrate, aluminum nitrate, ferric nitrate, silver nitrate, and zinc nitrate. Having thus determined the degree of concentration at which the different nitrates are toxic to Monilia sitophila in starch media, experiments were made in which the organic part of the medium was varied for the purpose of determining whether or not the toxicity of these salts can be modified by the presence of one or another of the organic sub- stances commonly used in making media. The organic substances tried are peptone, glucose, fructose, and galactose. The results obtained in these experiments are shown in TABLES III to VIII. As shown by TABLE III, barium nitrate in peptone media at a concentration of 0.133 molar inhibits the germination of the spores of Monilia. In a starch medium containing barium nitrate at a concentration of 0.156 molar, the spores germinate and produce a small amount of mycelium. This shows that barium nitrate is more toxic in peptone media than in starch media. No concentration of barium nitrate shown in TABLE III was of sufficient strength to inhibit germination and growth in the presence of glucose. There was, however, a very small amount of mycelium in the media containing the barium nitrate at a concentration of 0.167 molar. This indicates that the toxic dose in glucose media is near the concentration 0.167 molar. The toxicity of barium nitrate in starch media and in glucose media is approximately the same. Its toxicity in peptone media is much greater than in glucose media or in starch media. In fructose media its toxicity is approximately the same as in starch and. glucose media but is much less than in peptone media. At a concentration of 0.1 molar, barium nitrate in the presence of galactose inhibits the germination of spores. Its toxicity in galactose is approximately the same as in peptone, but is much greater than in starch, glucose, or fructose. At a concentration of 0.033 molar, aluminum nitrate in peptone 632 KUNKEL: INFLUENCE OF STARCH, PEPTONE, AND SUGARS ON TABLE III THE TOXICITY OF BARIUM NITRATE IN PEPTONE, GLUCOSE, FRUCTOSE, AND GALACTOSE MEDIA edium Growth after 11 days. .167 molar solution of barium nitrate + 5% peptone......... Z .133 ‘“ “ “ 6s re + * ae so fare) ata Oo 067 “ «6 i ee + ‘* lS eae eas ey cope 033 ry 6 “ ‘ ies + * Peds wh nye eas ee cae + 17“ ‘ “ ‘ a“ + * ee Rain Oe + 007 “ ‘6 ‘6 #6 L — Re er pact Aenea + 167“ “ Siete eT eae RROOSE: SMEs ss = sag te “ as hs a OA es ene arias ate = 002 ie Wee 2 Se aa pte ira Corea ras ~ 067 >“ eB Gaye as eine at Shh oe * 033 oe bi bad ae ae + 5 Seay tres RS Boe asi o17 “ ‘“e “6 +s be + * = Fare eye TS = 007 aE SE ee Lee See + 005 ae eae ls, Hein ASRS a 004 “ “ 66 6 iy + * sista A OG tangs cid 167. * 4 ie ee = fee trmetose sas, 4s s = 133°“ “ “ “ ys + * spies hes Zo “Se eae Bane ic ane, a ets Maa roe - day ee ee oe ane Tee - 033." re So He ny od eg emg = O17 ‘ ‘6 se ‘6 FE + 1s Stee ede ails ake ae 007“ “ ik ee Reet tegen ee Ef 005 “ ‘ ‘ e + * Prog oe Wap + 004“ a be eae Oa ISS ahenuope s e,: 167 ‘“s “ ‘s “ id + * Feat Mapa Gear et oO 133. * ee ee ae” Fe eS ees 0 a8 és Sane ty tee a: er ia eta tel fa) 067 “ ‘6 +e + * RG Tere rarer Fk = 033. a oe a clog. Kas eo = O17 ‘ 46 s as +f + * RIG epee SDT Ae + 007“ eae ig PTE My ae eRe te iL coos“ ‘ “ a“ oe + * il nine ge tee ee + .004 =“ o i Hy Sk eyo te RePeae ae SRE + distilled water + ‘‘ peptone......... ees . ee 50 ODMR Saline + we Slee oo EEC SOEE bor aan cise i " es palaetoee cis gs es +: media inhibits the germination of the spores; in a concentration of 0.017 molar, the spores germinate and produce microscopic mycelia. Aluminum nitrate in starch media is more than ten times as toxic as in peptone media. Its toxicity in glucose media is approximately the same as in starch media. It is much less TOXICITY OF VARIOUS NITRATES TO MONILIA SITOPHILA 633 TABLE IV THE TOXICITY OF ALUMINUM NITRATE IN PEPTONE, GLUCOSE, FRUCTOSE, AND GALACTOSE MEDIA ium Growth after 11 days .167 molar solution of aluminum nitrate + 5% peptone....... 2240 .133 “ ‘“e “ “ be +s Peres lg

on f=.) ~J TOXICITY OF VARIOUS NITRATES TO MONILIA SITOPHILA 639 ammonification by Bacillus subtilis, Bot. Gaz. 48: 105-125. 1909. . Schreiner, O., & Skinner, J. J. The toxic action of organic com- pounds as modified by fertilizer salts. Bot. Gaz. 54: 31-48. 1912. . Ssadikow, W. S. Ueber den Einfluss des Strychnins auf Bakterien. Centralbl. Bakt., 1 Abt. 60: 417-425. I9QII. . Went, F. A. F.C. Monilia sitophila (Mont.) Sacc., ein technischer Pilze Javas. Centralbl. Bakt., 2 Abt. 7: 544-550; 591-598. 1901. . Winogradsky, S., & Omeliansky, V. Ueber den Einfluss der organ- ischen Substanzen auf die Arbeit der nitrifizierenden Mikrobien. Centralbl. Bakt., 2 Abt. 5: 329-343. 1899. Resistance of the prothallia of Camptosorus rhizophyllus to desiccation * F. L. PIcKetr Camptosorus rhizophyllus (L.) Link is found growing with mosses and lichens on the shaded surface of dry limestone ledges and on detached limestone slabs in open ravines and torrent beds. Only rarely have groups been found in well-shaded or continually moist places in this region (southern Indiana). Growing in places thus exposed without constant water supply, the plants are subjected to brief periods of abundant moisture (during and immediately after precipitation) which alternate with longer periods of drought. That plants with a delicate prothallial stage in their life history could secure and retain residence under such conditions has been a cause for surprise. The drought-resisting power of some greenhouse cultures of this fern grown in the spring of 1912 suggested a possible adapta- tion to its well-known xerophytic habitat. In an attempt to ‘determine lo what extent this ability to withstand drought might be a factor in adaptation, fronds with mature spores were collected in October, 1912, and cultures were made as usual on sterilized soil in clay saucers. These cultures were subjected to a variety of conditions to be later enumerated. An attempt was made to obtain information on the following points: the uniformity of spore germination and prothallial ‘development, the ability of prothallia to resist or survive natural ‘drought conditions, and the ability to survive conditions leading to complete desiccation. Fronds were collected on October 26, 1912, and kept between sheets of filter paper in a book in the laboratory. The sporangia were lightly crushed to free the spores and then sown on thoroughly sterilized soil, November 22, 1912. The cultures were kept in the greenhouse and were protected by bell-jars supported on * The writer has been unable to find any literature bearing upon this subject, for the prothallia of this or any other homosporous fern. 641 642 Pickett: RESISTANCE OF PROTHALLIA TO DESICCATION small blocks of wood to provide adequate ventilation. Some cultures were exposed to direct sunlight while others were exposed to strong diffused light only. The temperature conditions were the same, 20-25° C. The soil was kept moist, not wet, but was allowed to show a dry surface for a period of 12 to 24 hours once each week. The first green was noticed December 17, at which time the prothallia when examined with a microscope were found to be composed of two to ten cells each. These cultures showed good growth, seeming to suffer no injury from the dry periods. Those in the sunlight showed rather a more rapid development than those in diffused light. On February 11, 1913, the culture in sunlight contained prothallia in good condition but varying in size from merely germinated spores up to plants 2-3 mm. broad with but very few antheridia or archegonia. The culture in diffused light did not show similar prothallia until March 21, 1913. The spores of Camptosorus rhizophyllus germinate very irregu- larly. Twelve weeks after the spores were sown, a small bit of soil—not over 3 mm. square—removed from a culture where the plants seemed most thrifty, showed all stages in development from spores with the perinium just ruptured to prothallia bearing mature antheridia and archegonia. Many spores retain their vitality up to May in the dry atmosphere of the laboratory, and fronds collected in March furnished viable spores for cultures. The long dormant period of spores on the moist soil of cultures suggests that they might remain so on the soil of their habitat through the winter season. | Four methods of reducing water content were used. First, a glycerin desiccator was used, consisting of two glass vials through which, by means of an aspirator, a current of air was drawn after passing through two U-tubes containing glycerin and crumpled filter paper. Heaviest c.p. glycerin was used. The aspirator was arranged so that the air of the vials was changed about one hundred times each twenty-four hours. Second, cul- tures were left under bell-jars, exposed to the warm air and full sunlight of the greenhouse. Third, cultures were left under bell- jars, exposed to the dry air and diffused light of the greenhouse lobby or vestibule at a slightly lower temperature, 16-20° C This place represents as nearly as possible the natural growth PICKETT: RESISTANCE OF PROTHALLIA TO DESICCATION 643 conditions. In the second and third cases full ventilation was secured by allowing the jars to rest on blocks of wood 2 cm. high. Fourth, large portions of a culture were placed in a desiccator over c.p. sulphuric acid and the whole apparatus was kept in a cool, moderately lighted location. In all cases, when soil and pro- thallia were removed from a dry chamber they were placed in contact with moist soil under a bell-jar subject to full diffused light at a temperature of 16-20° C., for recovery. Examination for dead prothallia was made after three or four days under such conditions. Extreme care was taken at all times when removing portions of cultures to or from the dry chambers, to leave the prothallia as far as possible undisturbed and uninjured. Results—glycerin desiccator.—A large portion of the soil of a culture was removed to the vial of the desiccator and allowed to remain in the dry air undisturbed, except as small portions were removed for recovery and growth. Specimens placed in the desiccator on March 22 seemed to revive completely up to April 22, at which time a few small dead prothallia were found in a portion removed. Another portion removed April 29 showed about 50 per cent of the prothallia dead. The last of the soil and prothallia was removed May 5. All the smaller plants and all but a very few of the larger plants were dead at that time. In this set of experiments a very few fully matured prothallia survived continuous exposure to dry air for forty-four days. Very few were damaged by such exposure for a period of thirty days. That the recovery was complete in case of survival is proven by the continued growth of the prothallia and their later production of sporophytes. That vigorous desiccation follows immediately after the plants were placed in the vials is shown by the fact that the soil had given up all free moisture in twenty-four hours after being placed therein. As a check, at the beginning of this experiment a clump of thrifty mature prothallia of Onoclea Struthiopteris was placed in one of the vials. After forty-eight hours’ exposure to the dry air not one plant recovered. A similar set of experiments was arranged with the whole apparatus exposed to the direct sunlight. Most of the prothallia so exposed were dead at the end of twenty-eight days, and all were dead after thirty-five days of exposure. 644 PiIcKETT: RESISTANCE OF PROTHALLIA TO DESICCATION Results—sulphuric acid desiccator.—A portion of soil bearing prothallia was removed to a porcelain dish in a desiccator con- taining c.p. sulphuric acid. The lid was sealed down with vaseline and the apparatus placed in diffused light at a temperature of 16-20° C. After eighteen hours the prothallia showed a marked yellowish color. A portion removed after four days showed a recovery of but two or three per cent. The plants removed at this time recovered very slowly, requiring a week to resume a normal appearance. Results—normal dry air.—The soil mass of a culture was divided into two approximately equal parts and carefully removed to sterilized clay saucers. One portion was placed under a bell-jar_ exposed to the full sunlight in the greenhouse. The other was kept under a bell-jar exposed to full diffused light and at a temper- ature of 16-20° C., an average of four degrees lower than that for the first portion. After three weeks, but a few of the plants exposed to direct sunlight recovered, and all were dead after five weeks of such exposure. Of the second lot not exposed to direct sunlight a portion removed after thirty-four days showed almost complete recovery. After fifty-five days, about 25 per cent of the prothallia recovered. After sixty-three days, a very few of the largest plants recovered and continued to grow. The conditions under which the second group of plants was kept approximate very closely the summer conditions in the regular habitat of Camptosorus rhizophyllus. The results of that set of experiments certainly suggest an explanation of this plant’s abundant growth under such conditions. It should be noted here, as above, that full recovery of the plants has been demonstrated by their continued growth and later production of sporophytes. If mature prothallia can withstand continuous drought for two months, they would certainly survive the difficulties of the average season after late March, at which time spores may germinate outside. The occasional rains through the summer would make possible recovery, fertilization, and the coguape of sporophytes. In connection with this set of experiments another should be noticed. A culture was prepared as for all the above on No- vember 25, 1912. The drainage was such that a part of the soil PICKETT: RESISTANCE OF PROTHALLIA TO DESICCATION 645 surface was dry at all times, except when occasionally flooded. The remaining portion was moist as in the other cultures. All gradations of moisture were in evidence in and between the two. regions. Small prothallia were seen on December 18 on the damp soil. On March 1, 1913, most of the prothallia of this region showed mature sex organs. The drier portions at the latter date showed germinating spores, dwarf male prothallia, and later stages in development up to mature plants. After March 1, this culture was watered irregularly and was allowed to. become quite dry each time before more water was applied. After April 15 the culture was screened from the direct sun. Water was occasionally applied so as to flood the whole surface. On May 5 there were many living prothallia and several young sporophytes. On June 9 there were several large, living pro- thallia and the young sporophytes were uninjured, although during the previous month the culture had been four times dry, once for seventy-two hours. SUMMARY As has been stated above, the experiment of subjecting pro- thallia to normal dry air without direct sunlight,—continuous conditions approximating the average of varying conditions found in nature,—has shown that the production of mature prothallia under such conditions is possible. The other experiments of subjecting prothallia to more thorough desiccation in the glycerin desiccator and over sulphuric acid show the possibility of sur- viving the extreme conditions found in nature. There can remain but little doubt that the drought-resisting character of the prothallia is a very effective factor in the adaptation of Camptosorus rhizophyllus to its habitat. INDIANA UNIVERSITY, BLOOMINGTON INDEX TO AMERICAN BOTANICAL LITERATURE (1913) of this Index is to include all current botanical literature written by Americans, published in America, or based upon American material ; the word Amer- ica being used in the broadest s Reviews, and papers that ore exclusively to forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and no attempt is made to index the literature of bacteriology. An occasional exception is made in favor of some paper appearing in an American periodical which is devoted wholly to botany. Reprints are i a unless they differ from the original in some important particular. If users of the Index will call the attention of the editor to errors or omissions, their kindness will be appreciated. This Index is ae monthly on cards, a furnished in 1 this form to subscribers at the rate of one t for each card, Selections of cards are not permitted ; each subscriber must wee ‘a cards published during the term of his subscription, Corre- spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club, Anthon, S. I. The surprise lily. Am. Bot. 19: 81-83. Au 10913. [ llust.]} Calochortus macrocarpus. Bailey, W. W. Transient botanizing. Am. Bot. 19: 98, 99. Au 1913. Bartholomew, E. T. Black heart of potatoes. Phytopathology 3: 180-182. pl. 19. 14 Au 1913. Bitter, G. Solana peruv., aequat., boliv. Bot. Jahrb. Beibl. 50: 5°-67. 19 Au 1913. Includes 7 new species in Solanum. Blodg-tt, F. M. Hop mildew. Cornell Univ. Agr. Exp. Sta. Bull. 328: 281-310. f. 93-1 1. Mr 1913. A disease caused _by Sphaerotheca Humuli. Borgesen, F. The marine algae of the Danish West Indies. Part I. Chlorophyceae. 1-160. f. 1-126 + chart. Copenhagen. 1913. Includes Pringsheima Udoteae, Cladophora uncinata, C. corallicola, and Avrain- villea Geppii, spp. nov. Brand, A. Polemoniaceae peruvianae et bolivienses. Bot. Jahrb. Beibl. 50: 50-52. 19 Au 1913. Includes Huthia longiflora sp. nov. Brown, P. E. Media for the quantitative determination of bacteria in soils. Centralb. Bakt, Zweite Abt. 38: 497-506. 9 Au 1913. 647 648 INDEX TO AMERICAN BOTANICAL LITERATURE Burger, O. F. A bacterial rot of cucumbers. Phytopathology 3: 169, 170. 14 Au 1913. Clute,W. N. The day lilies. Am. Bot. 19: 86-90. Au 1913. [Illust.] Cogniaux, A. Cucurbitaceae andinae. Bot. Jahrb. Beibl. 50: 75-76. 19 Au 1913. Includes A podanthera eriocalyx sp. nov. Cogniaux, A. Melastomataceae peruvianae. II. Bot. Jahrb. Beibl. 50: 31-33. 19 Au 1973. Includes Tibouchina fulvipilis and Axinaea Weberbaueri, spp. nov. Collins, F. S. Three plants with extension of range. Rhodpra 15: 169, 170. 1S 1913 Panicum Bicknellii, Potentilla tridentata, and Juncus bufonius var. halophilus. Collins, G. N. Mosaic coherence of characters in seeds of maize. U.S. Dept. Agr. Plant Ind. Circ. 132: 19-21. 19 Je 1913. Cook, M. T., & Schwarze, C. A. A Botrytis disease of dahlias. Phyto- pathology 3: 171-173. pl. 17. 14 Au 1913. Dammer, U. Solanaceae americanae. II. Bot. Jahrb. Beibl. 50: 52-58. 19 Au 1913. Includes 6 new species in Dunalia. Fawcett, H. S. Two fungi as casual agents in gummosis of lene trees in California. Phytopathology 3: 194, 195. Je 1913. Also published in Monthly Bull. State Comm. Hort. Calif. 2: 601-617. f. 340-351. Au 1913. Botrytis vulgaris and Pythiacystis citrophthora. Fernald, M. L. A peculiar variety of the canoe birch. Rhodora 15: 168, 169. 1S 1913. Ferry, R. Etude sur les PORES Premier Supplément Rev. Myc- IQII: 1-96. pl. -8. 14 Je 1911. Fletcher, E.F. Further wool-waste plants at Westford, Massachusetts. Rhodora 15: 172. 1S 1913. Fromme, F. D. The culture of cereal rusts in the greenhouse. Bull. Torrey Club 40: 501-521. 10S 1913. Geckler, A. Echinocactus Droegeanus Hildm. Monats. Kakteenk. 23: 122. 15 Au 1913. Gilg, E. Gentianaceae andinae. Bot. Jahrb. Beibl. 50: 48-50. 19 Au 1913. Includes five new species in Gentiana and one in Macrocarpaea. Gilg, E. Malesherbiaceae andinae. II. Bot. Jahrb. Beibl, 50: 11 12. 19 Au 1913. Includes Malesherbia Weberbaueri and M. scarlatiflora, spp. nov, INDEX TO AMERICAN BOTANICAL LITERATURE 649 Griggs, R. F. Observations on the geographical composition of the Sugar Grove flora. Bull. Torrey Club 40: 487-499. f. I-10. 10S 1913. Giissow, H. T. The barberry and its relation to black rust of grain, Phytopathology 3: 178, 179. 14 Au 1913. Harris, J. A. An illustration of the influence of substratum hetero- geneity upon experimental results. Science II. 38: 345, 346. § S.1013. Harter, L. L., & Field, E. C. A dry rot of sweet potatoes caused by Diaporthe Batatatis. U.S. Dept. Agr. Plant Ind. Bull. 281: 7-38. pl. 1-4+f. 1-4. 1 My 1913. Heribert-Nilsson, N. Oecnothera-problemet. Svensk Bot. Tidsk. 7: 1-16. 25 Mr 1913. _ [Illust.] Herter, W. José Arechavaleta. Monats. Kakteenk. 23: 125-127. 15 Au 1913. - Hill, G. R. Respiration of fruits and growing plant tissues in certain gases, with reference to ventilation and fruit storage. Cornell Univ. Agr. Exp. Sta. Bull. 330: 377-408. Ap 1913. Hull, E. D. Advance of Potamogeton crispus L. Rhodora 1§: 171, 172. 3 @ 1013. Johnston, J. R. The nature of fungous diseases of plants. Porto Rico Sugar Producers’ Assoc. Cire. 2: 3-25. f. 1-9. My 1913. Jostmann, A. Pilocereus lanatus (H. B. K.) Web. var. Haaget (Po- selg.) Schelle. Monats. Kakteenk. 23: 125. 15 Au 1913. [Illust.] Kellerman, K. F. The use of congo red in culture media. U.S. Dept. Agr. Plant Ind. Circ. 130: 15-17. 21 Je 1913. Keyser, A. Variation studies in brome grass. (A preliminary report). Colorado Agr. Exp. Sta. Bull. 190: 3-20. pl. I-19. Je 1913. Kranzlin, F. Amaryllidaceae peruvianae, bolivienses, brasilienses. Bot. Jahrb. Beibl. 50: 2-5. 19 Au 1913. Includes Collania Herzogiana, Bomarea Ulei, B. Loreti, and Eucharis Ulei, spp. nov. : Kranzlin, F. Buddleiae americanae nonullis gerontogaeis adjectis. Bot. Jahrb. Beibl. 50: 33-47. 19 Au I913. Includes 22 new species in Buddleia. Kranzlin, F. Calceolariae peruv., aequat., argent. Bot. Jahrb. Beibl. 50: 67-75. 19 Au 1913. Includes 9 new species in Calceolaria. Loesner, T. Celastraceae andinae. II. Bot. Jahrb. Beibl. 50: 8-10. 19 Au 1913. Includes Maytenus apurimacensis and M. andicola, spp. nov. 650 INDEX TO AMERICAN BOTANICAL LITERATURE Meinecke, E. P. Notes on Cronartium coleosporioides Arthur and Cronartium filamentosum. Phytopathology 3: 167, 168. 14 Au 1913. Morse, W. J. Some borrowed ideas in laboratory equipment. Phyto- pathology 3: 175-177. pl. 18. 14 Au 1913. Muschler, R. Caryophyllacea aequatoriana. Bot. Jahrb. Beibl. 50: 5,6: 19 Au 1913. Drymaria adiantoides sp. nov. Muschler, R. Compositae peruv. et boliv. II. Bot. Jahrb. Beibl. 50: 76-108. 19 Au 1913. Includes 29 new species in Mikania (1), Liabum (9), Gynoxis (6), Chuquiraga (3), Onoseris (4), Barnadesia (5), Mutisia (3), and Jungia (3) Muschler, R. Crucifera peruviana. Bot. Jahrb. Beibl. £0: 7, 8. 19 Au 1913. Cremolobus stenophyllus sp. nov. Orton W. A. International phytopathology and quarantine legis- lation. Phytopathology 3: 144-151. 14 Au 1913. Pilger, R. Biologie und Systematik von Plantago § Novorbis. Bot. Jahrb. 50: 171-287. f. 1-30. 19 Au 1913. Includes Plantago taraxacoides, P. hypoleuca, P. pseudomyosuros, P. chubutensis, — P. ecuadorensis, P. Berroi, P. achalensis, P. Buchtienti, P. macropus, P. been? P. Arechavaletai, P. _ventanensis, P. denudata, P. valida, spp. nov., and many 2 varieties from Amer. Pilger, R. ae peruviana. Bot. Jahrb. Beibl. 50: 1, 2. 19 Au 1913. Trichoneura Weberbaueri sp. nov. Pilger, R. Rosacea peruviana. Bot. Jahrb. Beibl. 50:8. 19 Au 1913- Prunus huantensis sp. nov Prescott, A. The moonwort. Am. Bot. 19:95-97. Au 1913. [Illust.] Putnam, B.L. Our native Aguilegia. Am. Bot. 19:97. Au 1913- Rydberg, P. A. Studies on the Rocky Mountain flora—X XIX. Bull. Torrey Club ae 461-485. 10S 1913. Includes 26 new species in Hypopitys (1), Primula (1), Androsace (2), Amarella (1), Hydrophyllum (1), purehtes (1), Dasystephana (2), Amsonia (1), Cress@ (1), Cuscuta (1), Gilia sok gunn (1), Polemonium (2), Phacelia (1), Oreocarya (2)s Pentstemon us (1), and Castilleja (1). Saunders, s ri nis soaps. Am. Bot. 19: 99, roo. Au 1913- Schellenberg, G. Berberidacea peruviana. Bot. Jahrb. Beibl. 50: 6; 7. 19 Am-198s. Berberis peruviana sp. nov. INDEX TO AMERICAN BOTANICAL LITERATURE 651 Schellenberg, G. Frankeniacea peruviana. Bot. Jahrb. Beibl. 50: 10) 31.19-A-1913 Frankenia peruviana sp. nov. : Shear, C.L. The type of Sphaeria radicalis Schw. Phytopathology 3: 191, 192. 14 Au 1913. Stewart, V. B. The fire blight disease in nursery stock. Cornell Univ. Agr. Exp. Sta. Bull. 329: 317-371. f. 172-126. Ap 1913. Sturgis, W. C. Herpotrichia and Neopeckia on conifers. Phyto- pathology 3: 152-158. pl. 12, 13. Je 1913. -Swingle, W. T. Feroniella, genre nouveau de la tribu des Citreae, fondé sur le F. oblata, espéce nouvelle de |’Indo-Chine. Bull. Soc. Bot. France §9: 774-783. pl. 18. 18 F 1912. Sydow, H. & P. Novae fungorum species—X. Ann. Myc. 11: 254-271. f. 1-8. Je 1913. Includes Sphaerulina salicina from haa sige Dakota, Phyllachora atro-maculans and Dothidella Picramniae from Costa Taubenhaus, J. J. The black rots of the sweet ibis Phys: pathology 3: 159-166. pl. 14-16. Au 113. Includes Sclerotium bataticola sp. nov. Thomson, R. B. On the comparative anatomy and affinities of the Araucarineae. Philos. Trans. Royal Soc. London B 204: I-50. pl. 1-7.. 7 My 1913. Trelease, W. Furcraea peruviana. Bot. Jahrb. Beibl. 50: 5. 19 Au 1913. Furcraea occidentalis sp. nov. Tullsen, H. Pentstemon grandiflorus. Am. Bot. 19: 84, 85. Au 1913. [Illust.] Urban, I. Plantae novae andinae imprimis Weberbaurinae. VI. Bot. Jahrb. Beibl. 50: 1-108. 19 Au 1913. Consists of 19 separate papers here indexed under their respective authors: Bitter, Brand, Cogniaux, Dammer, Gilg, Kranzlin, Loesener, Muschler, Pilger, Schellenberg, Trelease, and Vaupel. Vaupel, F. Cactaceae andinae. Bot. Jahrb. Beibl. 50: 12-31. 19 Au 1913. Includes Cephalocereus melanostele, Cereus acanthurus, C. apiciflorus, C. petalus, C. decumbens, C. micranthus, C. i napenowe , C. squarrosus, C. Weber Echi: us aurantiacus, E. = S serach, E. Weberbaueri, sit nocact dactylifera, and O. ignescens, spp. nov. Vaupel, F. Sieben neue Cactaceae. Monats. Kakteenk. 23: 105-107. 15 Jl 1913. : Vaupel, F. Verzeichnis der seit der Herausgabe des I. Nachtrages zu K. 652 INDEX TO AMERICAN BOTANICAL LITERATURE Schumann’s ‘‘Gesamtbeschreibung der Kakteen’’ (1903) neu beschriebenen und umbenannten Gattungen und Arten aus der Familie der Cactaceae. Monats. Kakteenk. 23: 11-14. 15 Ja 1913; 23-27. 15 F 1913; 37-41. 15 Mr 1913; 56-60. 15 Ap 1913; 72-78. 15 My 1913; 81-88. 15 Je 1913. Weingart, W. Cereus Hirschtianus K. Schum. Monats. Kakteenk. 23: 108-111. 15 Jl 1913. Weir, J. R. Auricularia mesenterica (Dicks.) Pers. Phytopathology 3: 192. Au 1913. Wellington, R. Mendelian inheritance of epidermal characters in the fruit of Cucumis sativus. Science II. 38: 61. 11 Jl 1913. Wernham, H. F. New Rubiaceae from tropical America—II. Jour. Bot. 51: 218-221. Jl 1913 : Includes eleven new species in Pteridocalyx (1), Tournefortiopsis (1), Gonzalea (4), Machaonia (3), Malanea (1), and Cephaelis (1). Wester, P. J. Annonaceous possibilities for the plant breeder. Philip. Agr. Rev. 6: 312-321. pl. 2-7. Jl 1913. Westling, R. Uber die griinen Spezies der Gattung Penicillium. Arkiv Bot. rz’: 1-156. f. 1-81. 1911. Discusses several American species. Williams, R.S. Dicranaceae. N. Am. Fl. 15: 77-158. 8 Au 1913. Includes new species in Dicranella (2), Dicranum (1), and Campylopus (4). Williams, R.S. Leucobryaceae. N.Am. Fl. 15: 159-166. 8 Au 1913- Includes Octoblepharum erectifolium Mitten sp. nov. Wilson, G.W. Fusarium or Verticillium on okra in North Carolina? Phytopathology 3: 183-185. Je 1913. Wolf, F. A. Melanose. Phytopathology 3: 190, 191. 14 Au 1913- Yamanouchi, S. The life history of Zanardinia. Bot. Gaz. ee 1-35. pl. 1-4+f. 1-24. 16 Jl 1913. York, H. H. The origin and development of the embryo sac and embryo of Dendrophthora opuntioides and D. gracile{is]. 1. Bot. Gaz. 56: 89-111. pl. 5, 6 +f. 1, 7, 8, 12a, 20, 21. 14 Au 1913; II. Bot. Gaz. 56: 200-216. pl. 7. 17S 1913. “Vol. 4C No, 12 BULLETIN OF THE TORREY BOTANICAL CLUB DECEMBER, 1913 West Indian mosses—| ELIZABETH GERTRUDE BRITTON (WITH PLATE 25) A. West INDIAN MOSSES KNOWN TO LINNAEUS In Linnaeus’ Species Plantarum* 8 generat and 103 species of mosses are recognized, of which only 2 are known to be tropical American in their distribution, ranging from southern Florida to South America. The first of these tropical species is Bryum albi- dum L. (p. 1118) known to Dilleniust as Bryum nanum, lariginis foliis albis, and now known as Octoblepharum albidum (L.) Hedw., with the type locality on the island of New Providence in the Bahamas. The other species, Rhizogonium spiniforme (L.) Bruch was the first species of Hypnum named by Linnaeus and it also was based on a Dillenian description and plate.§ He called it ‘the Herring’ s-Bone Hypnum” and his specimens were sent to him from Mt. Diabolo, Jamaica, by Sir Hans Sloane. Its range through the tropics is even wider than that of Octoblepharum, including the Islands of the Pacific; both species are known to occur not only throughout the American tropics but also in Asia and Africa. Tr Sihapesits ecees pani eos 2 5. Polyirichum. ..i...006.4. 3 2. PRESCUM i eee 3 6: MUM, So oy ei eet 18 Bo ROMUNGHS <0 ics ce we ow 4 Pa og 1, aA Parone ie ae ert 30 S Mis ak eae ees 3 BN ss Raat 40 103 t Historia Muscorum 364. pl. 46. f. 2. eet te Historia Muscorum 332. pl. 43. f. 6 {The ButLeTIn for November (40: ares Spabeal was issued 24 N 1913.] 653 654 BrITTON: WEsT INDIAN MOSSES 1. OCTOBLEPHARUM ALBIDUM (L.) Hedw. Descr. 3: 15. 1791 Bryum albidum L. Sp. Pl. 1118. 1753. Type LocaALity: New Providence, Bahamas. DISTRIBUTION: Florida and the Bahamas, ercedtiowt: the West Indies; in Cuba, Jamaica, Haiti and St. Domingo, Porto Rico, Dominica, St. Thomas, St. Kitts, Montserrat, Guadeloupe, Martinique, St. Lucia, Grenada to Trinidad; also in South America and tropical regions of Africa and Asia. ILLUSTRATIONS: Dill. Hist. Musc. pl. 46. f. 21; Hedw. Descr. 3: pl. 6A; Card. Rech. Anat. Leuc. pl. 12. f. 61. ExsiccaTaE: Sull. Musci Cub. Wright. 55. 1861; Husnot, PI. Ant. Fr. 127. 1868; Austin, Musci App. Suppl. 478. 1874: Ren. & Card. Musci Am. Sept. Exsicc. 213; Holz. Musci Acroc. Bor. Am. 57; Small, Mosses S. U.S. 52. 2. RHIZOGONIUM SPINIFORME (L.) Bruch, Flora 29: 134. 1846 Hypnum spinaeforme L. Sp. Pl. 1122. 1753. Mmium spiniforme C. Mill. Syn 1: 175. 1849. Type Loca.Lity: Mt. Diabolo, Jamaica, Hans Sloane. DIstTRIBUTION: In wet woods, in tropical regions of all portions of the world. Southern United States: Georgia, Alabama, Louisiana, Florida; Jamaica, Cuba, Haiti, Porto Rico, Guadeloupe to S. America; Mexico, Guatemala, Costa Rica, and Panama. . ILLUSTRATIONS: Sloane, Hist. Jam. pl. 25. f. 4.1707; Dill. Hist. Musc. 332. pl. 43. f. 8. 1741. ExsICCATAE: Sull. Musci Cub. Wright. 58. 1861; Husnot, PI. Ant. Fr. 752. 1868; Austin, Musci App. Suppl. 576. 1874; Ren. & Card. Musci Am. Sept. Exs. 64; Holz. Musc. Acroc. Bor. Am. 174; Pringle, Musci Mex. 10482. Another of the Linnaean species, Hypnum cuspidatum L., has been found in the high mountains of Jamaica. Pogonatum ur- nigerum L.. Sp. Pl. 1109. 1753, was also credited to Jamaica, fol- lowing Dillenius, who quotes Hans Sloane’s History of Jamaica and mistook his f. 5, pl. 25, for this European species. F. 4 of the same plate is unmistakable for Rhizogonium spiniforme. B. West INDIAN MOSSES KNOWN TO OLOF SWARTZ In his Prodromus, Swartz* retained 5 of the generic names * Olof Swartz, Nova Genera & Species Plantarum seu Prodromus, etc. 138-14?- 1788. BRITTON: WEsT FNDIAN MOSSES 655 used by Linnaeus* and enumerated 41 species, which as at present recognized belong to 37 different genera, three of these having their type localities in Hispaniola (Haiti and Santo Domingo), all the rest in Jamaica. In studying the collections made by Mr. Wm. Harris in Jamaica and ourown later collections, a special effort has been made to obtain an accurate knowledge of these Swartz types and Dr. A. Le Roy Andrews, of Cornell University, very kindly consented, when he visited Stockholm in the summer of 1912, to examine these types for me and compare them with specimens from our own collections in Jamaica, sent as duplicates to the Naturhistoriska Riksmuseum. Dr. Andrews was able to see and compare the original specimens with ours in all but two cases: Bryum parasiticum Sw. |= Syr- rhopodon parasiticus (Sw.) Besch.] and Hypnum congestum Sw. [= Pleuropus congestus (Sw.) Broth.], which species we have not yet been able to recognize, the former being from Hispaniola and the type lacking in Swartz’ herbarium, the latter from Jamaica and Haiti. We suspect from the illustration given by Hedwig that the latter is probably referable to Palamocladium Bonplandt (Hook.) Broth., which Brotherus later refers to Pleuropus, though he states that he has not seen specimens of Pleuropus congestus. In 1806 Swartz discarded his Linnaean limitationst and adopted some of the generic changes proposed by Hedwig (1792), to whom he sent specimens of most of his West Indian mosses, from which almost all of Hedwig’s plates were drawn. This eliminated Fentinalis and Mnium from the West Indies and added seven generat and three species to the list given in the Prodromus; he further amplified his list by giving more in detail the stations and habitats. These are translated and quoted in the following list of species in the sequence enumerated by Swartz, with their modern Names, synonyms, and distribution as at present known to us from the West Indies: * Fontinalis, Polytrichum, Mnium, Bryum, and Hypnum. + Fl. Ind. Occ. 3: 1759-1841. 1806. t Encalypta, Trichostomum, Tortula, Dicranum, Pterogonium, Neckera, and kea, ligt 656 BrITTON: West INDIAN MOSSES 1. Neckera jamaicense (Gmel.) E. G. Britton, comb. nov. Fontinalis crispa Sw. Prod. 138. 1788. Not Hypnum crispum L. op. Fl. F124. (91753. Hypnum jamaicense Gmel. Syst. Nat. 2: 1341. 1791. Neckera undulata Hedw. Descr. 3: 51. 1792. Neckera undulata Hedw.; Sw. Fl. Ind. Occ. 3: 1780. 1806. Neckeropsis undulata Kindb. Eu. & N. A. Bryin. 1:20. 1897. HABITAT AND TYPE LOCALITY: ‘‘On trunks of trees in dense low woods, Jamaica.” DIsTRIBUTION: Not uncommon on trees from Florida and the Greater and Lesser Antilles to Trinidad; Mexico, Guatemala, and “Panama; also in South America. _ Ittustrations: Dill. Hist. Musc. 294. pl. 32. f. 8; Hedw. Descr. pl. 21 (from Swartz’ type). ExsIccaTarE: Austin, Musci App. Suppl. 529; Sull. Musci Cub. Wright. 75; Husnot, Pl. Ant. Fr. 155; Grout, N. A. Musci Pleur. 230. 2. NECKERA DISTICHA (Sw.) Hedw. Descr. 3:53. 1792 Fontinalis disticha Sw. Prod. 138. 1788. Neckera disticha Hedw. Descr. 3: 53. 1792. Neckera disticha Sw. Fl. Ind. Occ. 3: 1784. 1806. Neckeropsis disticha Kindb. Eu. & N. A. Bryin. 1:20. 1897. HABITAT AND TYPE LOCALITY: On trunks of trees, Jamaica and Hispaniola. DISTRIBUTION: Less common, on trees, Florida and the Greater and Lesser Antilles to South America; in Central America from Mexico to Panama; also in Africa. ILLustRatTion: Hedw. Descr. pl. 22 (from Swartz’ type). ExsiccaTaE: Austin, Musci App. Suppl. 530. 3. PTEROBRYUM FILIcINUM (Sw.) Mitt. Jour. Linn. Soc. 12: 425+ 1869 Fontinalis fiicina Sw. Prod. 138. 1788. Neckera filicina Hedw. Descr, 3: 45. 1792. Pilotrichum filicinum P. Beauv. Prod. 83. 1805. Neckera filicina Sw. Fl. Ind. Occ. 3: 1788. 1806. Pireella filicina Cardot, Rev. Bryol. 40: 18. 1913. BRITTON: WEST INDIAN MOSSES 657 HABITAT AND TYPE LOCALITY: “‘ Near Coldspring, high moun- tains of southern Jamaica, on trunks of trees.” DISTRIBUTION: Jamaica and Cuba. ILLUSTRATION: Hedw. Descr. pl. 18 (from Swartz’ type). ExsIccaTAE: Sull. Musci Cub. Wright. 76. This species has immersed capsules and seems to belong where Mitten has placed it. 4. PILOTRICHUM HYPNOIDES (Sw.) P. Beauv. Prod. 83. 1805 Fontinalis hypnoides Sw. Prod. 138. 1788. Neckera hypnoidea Hedw. Descr. 3: 43. 1792. Neckera hypnoides Sw. FI. Ind. Occ. 3: 1790. 1806. HABITAT AND TYPE LOCALITY: ‘‘ Jamaica, on trunks of trees in high mountains.”’ _ DISTRIBUTION: Jamaica to Trinidad. ILLUSTRATION: Hedw. Descr. pl. 17 (from Swartz’ type). 5. CRYPHAEA FILIFORMIS (Sw.) Brid. Bryol. Univ. 2: 251. 1827 Fontinalis fiiformis Sw. Prod. 138. 1788. Neckera filiformis Hedw. Descr. 3: 41. 1792. Neckera filiformis Sw. Fl. Ind. Occ. 3: 1786. 1806. HABITAT AND TYPE LOCALITY: ‘‘ Hispaniola, in arid regions .on branches of Haematoxylon campechianum.” DIsTRIBUTION: Jamaica, Cuba, and Santo Domingo; also in South America, Central America and Mexico (Guatemala and Yucatan). ILLUSTRATION: Hedw. Descr. pl. 16 (from Swartz’ type). ExsIccaTaE: Sull. Musci Cub. Wright. 67. 6. PoGONATUM TORTILE (Sw.) Brid. Bryol. Univ. 2: 108. 1827 Polytrichum convolutum Sw. Prod. 139. 1788. Not L. 1753. Polytrichum convolutum Hedw. Sp. Musc. 94. 1801. Pogonatum convolutum Beauv. Prod. 85. 1805. Polytrichum tortile Sw. Fl. Ind. Occ. 3: 1839. 1806. Polytrichum domingense Brid. Mant. 201. 1819. Polytrichum cubense Sull. Proc. Am. Acad. 5: 281. 1861. Polytrichum glaucinum Besch. Ann. Sci. Nat. VI. 3: 210. 1876. Polytrichum Husnotianum Besch. |. c. Polytrichum crispulum Besch. 1. c. 211. 658 BRITTON: WEST INDIAN MOSSES Polytrichum laxifolium Besch. 1. c. 211. Polytrichum Pleeanum Besch. |. c. 212. Polytrichum Sintenisit C. Mill. Hedwigia 37: 222. 1898. Polytrichum (Catharinella) obscuro-viridis C. Mill. Hedwigia 37: 223. 1898. HABITAT AND TYPE LOCALITY: “On clay banks, high moun- tains of southern Jamaica.” DisTRIBUTION: Cuba, Jamaica, Haiti, Porto Rico, Guadeloupe, Martinique, Dominica, Grenada, and Barbados. ILLUSTRATION: Hedw. Sp. Musc. Pl. 20. ExsIccaTAE: Sull. Musci Cub. Wright. 57; Husnot, Pl. Ant. Fr. 153. This species varies greatly according to habitat, whether dry or wet, sunny or shady. It usually grows on roadside banks of. the hard red clay, on the dry or southern sides of the West Indian islands and under such conditions, does not attain the lax, long leaves that are produced in shady moist valleys. Micro- scopic sections of the leaves show the lamellae to be somewhat variable but all of one generally uniform character, and though the serrations of the margins are more or less variable, the teeth being at times appressed and at others spreading, we find no constant differences between them. The presence of teeth on the back of the costa is just as true of P. tortile Sw. as of P. glauci- num Besch. 7. BREUTELIA TOMENTOSA (Sw.) Sch. “In Hb.’ Paris, Index Bryol. 1: ed. 2.173. 1904 Mnium tomentosum Sw. Prod. 139. 1788. Bryum tomentosum Sw. Fl. Ind. Occ. 3: 1837. 1806. Bartramia macrocarpa Hampe, Linnaea 32: 141. 1863. Bartramia macrotheca Hampe, Ann. Sci. Nat. V. 3: 373- 1866. Breutelia macrotheca Jaeg. Adumb. 1: 556. 1873-74. HABITAT AND TYPE LOCALITY: ‘“‘ On the edge of woods high mountains of Jamaica.” DisTRIBUTION: Jamaica and Guadeloupe to South America; also Mexico and Costa Rica. ILLusTRATIONS: Hooker, Musci Exot. pl. rg. 1818. From original specimen of Swartz. E. & P. Nat. Pfl. 1°: 656. f. 498: 1904. BRITTON: West INDIAN MOSSES 659 8. PHILONOTIS SPHAERICARPA (Sw.) Brid. Bryol. Univ. 2: 25. 1827 Mnium sphaericarpon Sw. Prod. 139. 1788. Mnium sphaericarpum Hedw. Descr. 3: 93. 1792. Bryum sphaericarpon Sw. Fl. Ind. Occ. 3: 1835. 1806. Bartramia sphaericarpa Mitt. Jour. Linn. Soc. 12: 261. 1869. HABITAT AND TYPE LOCALITY: “In shady mossy places, summits of mountains of southern Jamaica.’ DISTRIBUTION: Florida, Jamaica, Porto Rico, St. Thomas, St. Kitts, St. Vincent, Martinique, and Guadeloupe; Honduras to South America. ILLUSTRATION: Hedw. Descr. 3: pl. 38A. 9g. DITRICHUM RUFESCENS (Hampe) Broth. in E. & P. Nat. Pfl. 1°: 300. I9QOI Mnium strictum Sw. Prodr. 139. 1788. Not Ditrichum strictum Hampe. 1867. Trichostomum strictum Sw. F1. Ind. Occ. 3: 1761. 1806. Trichostomum pallidum strictum Schwaegr. Suppl. 21: 77. 1823. Leptotrichum rufescens Hampe, Linnaea 31: 521. 1862. Cynontodium strictum Mitt. Jour. Linn. Soc. 12: 42. 1869. Cynontodium rufescens Mitt. Jour. Linn. Soc. 12: 44. 1869. Leptotrichum mexicanum Sch. ; Besch. Mém. Soc. Sci. Nat. Cher- bourg 16: 174. 4872. Leptotrichum capillifolium Sch.; Jaeg. Adumb. 1: 388. 1871-72. Leptotrichum pseudo-rufescens C. Miill. Bull. Herb. Boiss. 5: 554. 1897. HABITAT AND TYPE LOCALITY: “Jamaica, on shady slopes in sandy wet soil among other mosses, cold places.” DISTRIBUTION: Jamaica, 1,500-2,100 meters. Also, Mexico to Colombia. ILLUSTRATION: Schwaegr. Suppl. fl. 123. 10. TORTULA AGRARIA (Sw.) Sw. Fl. Ind. Occ. 3: 1763. 1806 Bryum agrarium Sw. Prod. 139. 1788. Bryum acuminatum Sw. Prod. 139. 1788. Barbula agraria Hedw. Descr. 3:17. 1792. Barbula Raui Aust. Bull. Torrey Club 6:43. 1875. 660 BRITTON: WEST INDIAN MOSSES HABITAT AND TYPE LOCALITY: “Jamaica and Hispaniola, in sugar fields and on calcareous rocks.”’ DistRIBUTION: Florida and Texas. Common in the Bahamas on limestone rocks, whence it was known to Dillenius. Jamaica, Cuba, Porto Rico, Guadeloupe, Antigua, Montserrat to Trinidad and South America; also in Mexico. ILLUSTRATION: Hedw. Descr. pl. 6B, from original specimens collected by Swartz in Jamaica and Santo Domingo. De _BRyum ACUMINATUM Sw. Prod. 139. 1788. (See 10) 12. SYRRHOPODON LYCOPODIOIDES (Sw.) C. Miill. Syn. 1: 538. 1849 Bryum lycopodioides Sw. Prod. 139. 1788. Dicranum? lycopodioides Sw. F1. Ind. Occ. 3: 1766. 1806. HaBITAT AND TYPE LOCALITY: ‘‘ Jamaica; in moist shady woods, on high mountains.” DIsTRIBUTION: Jamaica, Santo Domingo, Haiti, Porto Rico, Guadeloupe, and Martinique to Trinidad. ExsiccaTAE: Husnot, Pl. Ant. Fr. 151. 13. SYRRHOPODON PARASITICUS Besch. Ann. Sci. Nat. VIII. 1: 298- 1895 Bryum parasiticum Sw. Prod. 139. 1788. Encalypia parasitica Sw. Fl. Ind. Occ. 3: 1759. 1806. Calymperes parasitica Hook. & Grev. Edinb. Jour. Sci. 1: 131: 1824. The type cannot be found at Stockholm in Swartz’ herbarium. A fragment of the type specimen exists at Kew, and Mitten had only two leaves of it. He states that it is very close to Calymperes Richardi but the illustration given by Schwaegrichen of the calyp- tra and the description given by Swartz, ‘‘ Calypira longa subulata, non laxa, pallida, ore aequali, latere demum fissili”’ disprove this, and it is evident, either that Schwaegrichen was mistaken in figuring a calyptra which resembles that of a Macromitrium or it is a species of that genus, which is very common in Jamaica. Mitten referred a specimen collected by R. Spruce in South America (no. 2) “ this species but that proves to be a true Calymperes. BRITTON: WEsT INDIAN MOSSES 661 The duplicate type from which Schwaegrichen’s plate was drawn has been loaned to us from Geneva and corresponds with all of this plate except the calyptra, which is lacking; but the hyaline basal cells are not clearly indicated. It is evidently a species of Syrrhopodon with entire leaf margins bordered by elon- gated cells and does not agree with any known to us thus far from the West Indies. HABITAT AND TYPE LOCALITY: Hispaniola. ‘‘On branches of Haematoxylon and Mimosa Unguis-cati.” DIsTRIBUTION: Known only from the original collection. ILLUSTRATION: Schwaegr. Suppl. 1: 60. pl. 177. 1811. 14. HoLomirrium cALycinum (Sw.) Mitt. Jour. Linn. Soc. 12: 60. 1869 Bryum calcycinum Sw. Prod. 139. 1788. Weisia calycina Hedw. Sp. Musc. 70. 1801. Cecalyphum? calicinum Beauv. Prod. Aetheog. 50. 1805. Dicranum calycinum Sw. FI. Ind. Occ. 3: 1768. 1806. HABITAT AND TYPE LOCALITY: ‘‘On roots of trees in high moun- tains. Jamaica.’ DIsTRIBUTION: Known only from oe ILLUSTRATION: Hedw. Sp. Muse. #l. 14. f. I-5- 15. FIssIpENS PALMATUS (Sw.) Hedw. Descr. 3:69. 1792 Hypnum palmatum Sw. Prod. 140. 1788. Dicranum palmatum Sw. F1. Ind. Occ. 3: 1774-: 1806. Skitophyllum palmatum De la Pyl. Jour. de Bot. II. 4: 146. 1814. HABITAT AND TYPE LOCALITY: ‘‘In shady clayey places at roots of palms. Jamaica. Collected also on high trunk of Areca oleracea, in a cavity filled with rotten leaves.” DIsTRIBUTION: Jamaica, Cuba, and St. Thomas. ILLustrations: Hedw. Descr. pl. 30A. 1792 (from Swartz’ type); De la Pyl. Jour. de Bot. pl. 35. f. 6. 1814. ExsICcATAE: Sull. Musci Cub. Wright. 7. 16. FIssIDENS POLYPODIOIDES (Sw.) Hedw. Descr. =; 63. 1792 — Hypnum polypodioides Sw. Prod.'140. 1788. 3 Dicranum polypodioides Sw. Fl. Ind. Occ. 3: 1772- 1806. Skitophyllum polypodioides De la Py]. Jour. de Bot. II. 3: 153. 1814. 662 BRITTON: WEsT INDIAN MOSSES HABITAT AND TYPE LOCALITY: ‘‘On the ground in shady mossy slopes in high mountains, Jamaica.” DistRIBUTION: Georgia, Alabama, Florida, and Louisiana; Jamaica, Cuba, Haiti, Porto Rico; Dominica, Guadeloupe, and Martinique, to South America; also, Mexico, Guatemala, and Panama. ILLUSTRATIONS: Sull. Icon. Musc. pl. 27; De la Pyl. 1. c. pl. 38. fF: To. ExsiccATAE: Drummond, Musci Am. ed. 2. 38; Sull. & Lesq. Musci Bor. Am. ed. 2. 87: Sull. Musci Cub. Wright. zo; Small, Mosses So. U.S. 9; Husnot, Pl. Ant. Fr. 133. 17. FISSIDENS ASPLENIOIDES (Sw.) Hedw. Descr. 3: 65. 1792 Hypnum asplenioides Sw. Prod. 140. 1788. Dicranum asplenioides Sw. Fl. Ind. Occ. 3: 1770. 1806. Skitophyllum asplenioides De la Pyl. Jour. de Bot. II. 4: 156. 1814. Fissidens Barbae-montis C. Miill.; Ren. & Card. Bull. Soc. Roy. Bot. Belg. 311: 152. 1892. Fissidens costaricensis Besch. Bull. Herb. Boiss. 2: 390. 1894. HABITAT AND TYPE LOCALITY: “On mossy rocks in high moun- tains of Jamaica.” DISTRIBUTION: Jamaica and St. Kitts; also Mexico and Costa Rica. ILLustRatIONS: Hedw. Descr. pl. 28 (from type); De la Pyl. Jour. de Bot. II. 4: pl. 38. f. 8, 9. EXSICCATAE: Pringle, Musci Mex. 10,503. 18. PHYLLOGONIUM FULGENS (Sw.) Brid. Bryol. Univ. 2: 671. 1827 Hypnum fulgens Sw. Prod. 140. 1788. Pterigynandrum fulgens Hedw. Descr. 4: 101. 1797. Pterogonium fulgens Sw. Fl. Ind. Occ. 3: 1776. 1806. ? Phyllogonium viride Brid. Bryol. Univ. 2: 673. 1827. Phyllogonium aureum Mitt. Journ. Linn. Soc. 12: 424. 1869. Phyllogonium globitheca C. Mill. Bull. Herb. Boiss. 5: 563. 1897: HABITAT AND TYPE LocaLity: ‘Dependent from branches of trees in high mountains of Jamaica.” BRITTON: WEst INDIAN MOSSES 663 DISTRIBUTION: Jamaica, Cuba, Haiti, Porto Rico, St. Kitts, Antigua, Montserrat, Guadeloupe, Martinique, St. Vincent, and Grenada to Trinidad. Also in South America. ILLUSTRATION: Hedw. Descr. pl. 39 (from type). ExsIccaTAE: Sull. Musci Cub. Wright. 737; Husnot, Pl. Ant. Fr. 754. There is some doubt as to what the type specimen of Phyl- logonium viride of Bridel is. The type locality is Brazil and it is just possible that the name may antedate either P. immersum Mitt. or P. Serra C. Miill. Both these species were distributed by E. Ule in his Bryotheca Brasiliensis no. 81 from Serra Geral, Prov- ince of Santa Catharina, Brazil. All the West Indian specimens, so-called, are referable to P. fulgens. 19. LEPIDOPILUM DIAPHANUM (Sw.) Mitt. Jour. Linn. Soc. 12: 382. 1869 Hypnum diaphanum Sw. Prod. 140. 1788. Hypnum diaphanum Hedw. Sp. Musc. 243. 1801. Hypnum? diaphanum Sw. Fl. Ind. Occ. 3: 1828. 1806. Pterygophyllum diaphanum Brid. Bryol. Univ. 2: 345. 1827. Hookeria diaphana W.-Arn. Disp. Musc. 56. 1825. HABITAT AND TYPE LOCALITY: ‘In depressions, mountains of Jamaica. Mixed with Marchantia and Jungermannia.” DiIstRIBUTION: Jamaica, Martinique. ILLustRATIONS: Hedw. Sp. Musc. pl. 61. f. 1-6 (from type). 20. CyYCLODICTYON ALBICANS (Sw.) Broth. in E. & P. PA. 1°: 935- 1907 Hypnum albicans Sw. Prod. 140. 1788. Hypnum albens Gmel. Syst. Nat. 2: 1343. 1791. Leskea albicans Hedw. Sp. Musc. 218. 1801. Leskea albicans Sw. Fl. Ind. Occ. 3: 1811. 1806. Hypnum pallidum Brid. Musc. Rec. 2?: 127. 1806. Pterygophyllum albicans Brid. Bryol. Univ. 2: 349- 1827. HABITAT AND TYPE LOCALITY: ‘On old and rotten trunks of trees, temperate regions of Jamaica.” DiIstTRIBUTION: Jamaica, Guadeloupe, St. Vincent, and Mexico. ILLustRaTIONS: Hedw. Sp. Musc. 218. pl. 54. f. 13-16 (from type). ExsiccataE: Pringle, Musci Mex. 10,664. 664 BRITTON: WEST INDIAN MOSSES 21. HoMALIA GLABELLA’ (Sw.) Mitt. Jour. Linn. Soc. 12: 458. 1869 Hypnum glabellum Sw. Prod. 140. 1788. Leskea glabella Hedw. Sp. Musc. 235. 1801. Neckera glabella Sw. Fl. Ind. Occ. 3: 1782. 1806. HABITAT AND TYPE LOCALITY: ‘‘On trunks of trees in mountains of Jamaica.” DisTRIBUTION: Jamaica, Porto Rico, Guadeloupe, Mexico, Costa Rica, to Venezuela. ILLUSTRATION: Hedw. Sp. Musc. I. 59 (from type specimens). 22. METEORIOPSIS PATULA (Sw.) Broth. in E. & P. Pfl. 1°: 825. 1906 Hypnum patulum Sw. Prod. 140. 1788. Hypnum patulum Hedw. Sp. Musc. 279. 1801. Hypnum? patulum Sw. Fl. Ind. Occ. 3: 1832. 1806. Leskea remotifolia C. Miill. Linnaea 19: 216. 1847. Meteorium stellatum Lorentz, Moosst. 165. . 1864. Meteorium flaccidum Mitt. Jour. Linn. Soc, 12: 443. 1869. Meteorium tenue Sch. Besch. Mém. Soc. Sc. Nat. Cherbourg 16: 227%. 4872. Meteorium diversifolium Besch. 1. c. Meteorium torticuspis C. Miill. Bull. Herb. Boiss. 5: 204. 1897: HABITAT AND TYPE LOCALITY: ‘‘On roots and branches of trees near the summits, mountains of Jamaica.” DistTRIBUTION: Florida, in hammocks near Cutler, J. K. Smell; Jamaica, Haiti, Porto Rico, Guadeloupe, Martinique, Dominica, Montserrat, St. Vincent, Grenada, and Trinidad to South America; also Mexico, Guatemala, Honduras, Costa Rica, an Panama. ILLUSTRATION: Hedw. Sp. Musc. fl. 73. ExsIccaTAE: Sull. Musci Cub. Wright. 80; Husiot Pl. Ant. Fr. 168; Pringle, Musci Mex. 15,136. This is a common and variable species in the tropics and accordingly has received a variety of names. There seems to be no reason for maintaining two sections and such a host of names — in this genus, for according to Mitten and R. S. Williams the following also are syrionyms of this species: M. aureo-nitens Hampe (not Hook.), M. barbipendulum C. Miill.; M. cirrifolium Schw-, BRITTON: WEsT INDIAN MOSSES 665. M. chiriquense Ltz., M. Eurhynchium C. Miill., M. Filicis C. Miill., and M. subambiguum (Hampe) Paris. 23. MITTENOTHAMNIUM REPTANS (Sw.) Card. Rev. Bryol. 40: 21. 1913 Hypnum reptans Sw. Prod. 140. 1788. Hypnum reptans Hedw. Sp. Musc. 265. 1801. Hyvypnum reptans Sw. Fl. Ind. Occ. 3: 1819. 1806. Microthamnium reptans Mitt. Jour. Linn. Soc. 12: 506. 1869. Hypnum pseudo-repians C. Miill. Bot. Zeit. 14: 439. 1856. Microthamnium Turckheimti C. Mill. Bull. Herb. Boiss. 5: 215. 1897. Microthamnium minusculum C. Mill. Bull. Herb. Boiss. 5: 565. 1897. Stereohypnum reptans Fleisch. Hedwigia 47: 275. 1908. HABITAT AND TYPE LOCALITY: “‘On earth and trunks of trees, interior of Jamaica.”’ . DIsTRIBUTION: Cuba, Jamaica, Guadeloupe, Martinique, Mexico, Guatemala, Nicaragua, Costa Rica, Panama, and South America. ILLUSTRATION: Hedw. Sp. Musc. Pl. 68. 24. POROTRICHUM FASCICULATUM (Sw.) Mitt. Jour. Linn, Soc. 12: | 468. 1869 Hypnum fasciculatum Sw. Prod. 140. 1788. Hypnum fasciculatum Hedw. Sp. Musc. 245. 1801. Hypnum? fasciculatum Sw. Fl. Ind. Occ. 3: 1827. 1806. Thamnium fasciculatum C. Mill. Hedwigia 37: 260. 1808. HABITAT AND TYPE LOCALITY: ‘‘On roots of trees; high moun- tains of Jamaica.” DIsTRIBUTION: Jamaica, Porto Rico, and Trinidad to South America. | ILLustRATIONS: Hedw. Sp. Muse. pl. 62. f. 8-ro (from Swartz’ specimens). 25. HYPOPTERYGIUM TAMARISCI (Sw.) Brid. Bryol. Univ. 2: 715. I 827 | Hypnum Tamarisci ‘Sw. Prod. 141. 1788. | Leskea Tamariscina Hedw. Sp. Musc. 212. 1801. 666 BRITTON: WEsT INDIAN MOSSES Hypnum Tamarisci Sw. Fl. Ind. Occ. 3: 1825. 1806. Hypopterygium brastliense Sull. U.S. Expl. Exp. 26. 1859. | ?Hypopterygium pseudo-tamarisci C. Miill. Linnaea 38: 645. 1874. HABITAT AND TYPE LOCALITY: ‘On trunks of trees, creeping among mosses in the cold regions of Jamaica.” DISTRIBUTION: Jamaica, Cuba, Haiti, and Porto Rico; also in Mexico, Guatemala, Costa Rica, and South America. ILLUSTRATIONS: Hedw. Sp. Musc. pl. 62. f. 8-zo; Sull. U. S. Expl. Exped. pl. 26. ExsiccaTAE: Sull. Musci Cub. Wright. 230; Pringle, Musci Mex. 70,497. 26. PILOTRICHELLA FLEXILIS (Sw.) Jaeg. Adumb. 2: 162. 1875-76 Hypnum flexile Sw. Prod. 141. 1788. Leskea flexilis Hedw. Sp. Musc. 234. 1801. Hypnum? flexile Sw. Fl. Ind. Occ. 3: 1830. 1806. Meteorium flexile Mitt. Jour. Linn. Soc. 12: 438. 1869. Neckera cochlearifolia C. Mill. Syn. 2: 130. 1851. Neckera turgescens C. Miill. Syn. 2: 131. 1851. Pilotrichella eroso-mucronata C. Miill. Bull. Herb. Boiss. 5: 563. 1897. Pilotrichella recurvo-mucronata C. Miill. Bull. Herb. Boiss. §: 563- - 97. . . HABITAT AND TYPE LOCALITY: “Summits of mountains in Southern Jamaica.” DIsTRIBUTION: Jamaica, Cuba, Haiti, Porto Rico, Guadeloupe; Mexico, Guatemala, Nicaragua, Costa Rica, Panama; Colombia, Ecuador, Bolivia, and Brazil. ILLUSTRATION: Hedw. Sp. Musc. pl. 58. EXsICcATAE: Pringle, Musci Mex. 10,420, 10,468, Grout, N. A. Musci Pleur. 389. 27. PAPILLARIA NIGRESCENS (Sw.) Jaeg. Adumb.1:169. 1875-76 Hypnum nigrescens Sw. Prod. 141. 1788. Hypnum nigrescens Hedw. Sp. Musc. 250. 1801. Pierogonium nigrescens Sw. Fl. Ind. Occ. 3: 1778. 1806. Neckera nigrescens Schwaegr. Suppl. 32. 1828. ' BRITTON: WeEstT INDIAN MOSSES 667 Meteorium nigrescens Mitt. Jour. Linn. Soc. 12: 441. 1869. Papillaria nigrescens Donnellii Aust. Musci App. Suppl. 14. 1898. HABITAT AND TYPE LOCALITY: ‘On branches of trees, high mountains of Jamaica. Collected on Anacardium occidentale.” DISTRIBUTION: Louisiana, Florida, and the Bahamas; Jamaica, Cuha, Haiti, Porto Rico, Barbados, and Trinidad to South America; also in Lower California, Mexice, Guatemala, Costa Rica, and Panama. Also in South America. ILLUSTRATIONS: Hedw. Sp. Musc. pl. 65. 1801 ; Schwaegr. Suppl. pl. 244. 1828; Bryologist 7: 14. 1904. ExsiccaTAE: Austin, Musci App. Suppl. 533, Sull. Musci Cub. Wright. 83. The var. Donnellii is simply a xerophytic condition in which the leaves fall off and the terminal branches become brittle, thus propagating the species; in fact, the fruit is seldom found. Austin and J. D. Smith collected it at Caloosa, Florida, in 1876-78 and Mr. Severin Rapp has reported it from Sanford. In all our Jamaica collections I have found it but once, on a calabash tree. 28. PRIONODON DENsUS (Sw.) C. Miill. Bot. Zeit. 2: 130. 1844 Hypnum densum Sw. Prod. 141. 1788. Hypnum densum Hedw. Sp. Musc. 282. 1801. Hypnum? densum Sw. FI. Ind. Occ. 3: 1829. 1806. Neckera crassa Hornsch. FI. Brasil. 1: 56. 1840. Pilotrichum densum C. Miill. Syn. 2: 160. 1859. HABITAT AND TYPE LocaLity: ‘‘In Blue Mountains, southern Jamaica, on roots of trees.”’ DistTRIBUTION: Jamaica, Cuba, Haiti, Mexico, Costa Rica, and Panama, 1,500-2,000 ft.; also in South America. ILLUSTRATION: Hedw. Sp. Muse. $l. 74 (from Swartz’ type): Bot. Zeit. 2: pl. 1. FExsiccaTakg: Pringle, Musci Mex. 10.483. 29. Pritorricnum composituM (Sw.) P. Beauv. Prod. 82. 1805 Hypnum compositum Sw. Prod. 141. 1788. Neckera composita Hedw. Sp. Musc. 203. 1801. Neckera composita Sw. Fl. Ind. Occ. 3: 1792. 1806. 668 BRITTON: WEsT INDIAN MOSSES HABITAT AND TYPE LOCALITY: “On trunks of trees in woods, interior of Jamaica.” DISTRIBUTION: Jamaica and Grenada (‘‘Costa Rica’’?). ILLUSTRATIONS: Hedw. Sp. Muse. pl. 46. f. 8-13 (from Swartz’ type). 30. LEPIDOPILUM POLYTRICHOIDES (Sw.) Brid. Bryol. Univ. 2; 269. 1827 Hypnum polytrichoides Sw. Prod. 141. 1788. Hypnum polvirichoides Hedw. Sp. Musc. 244. 1801. Orthotrichum polytrichoides Brid. Musc. Recent. 2?: 31. 1801. Neckera polytrichoides Sw. Fl. Ind. Occ. 3: 1794. 1806. Lepidopilum polvtrichoides var. costaricense Ren. & Card. Bull. Soc. Roy. Bot. Belg. 32!: 192. 1893. Hookeria Carionis C. Mill. Bull. Herb. Boiss. 5: 205. 1897. HABITAT AND TYPE LOCALITY: ‘‘On branches of trees and shrubs, also on rocks, mountains of Jamaica and Hispaniola.” DISTRIBUTION: Jamaica, Cuba, Haiti, Porto Rico, Guadeloupe, Martinique, Montserrat, and St. Vincent to South America: also, Mexico, Guatemala, Costa Rica, and Panama. ItLustraTions: Hedw. Sp. Muse. pl. 61 (from Swartz’ type); Schwaegr. Suppl. 3: pl. 232. ExsIccaTAE: Husnot, Pl. Ant. Fr. 156. 31. HELICODONTIUM CAPILLARE (Sw.) Jaeg. Adumb. 2: 225- 1876-77 Hypnum capillare Sw. Prod. 141. 1788. Leskea capillaris Hedw. Descr. 4: 25. 1793. Leskea capillaris Sw. Fl. Ind. Occ. 3: 1813. 1806. HABITAT AND TYPE LOCALITY: ‘On trunks of trees, interior of Jamaica.” DISTRIBUTION: Jamaica, Cuba, Haiti, Porto Rico; also in Mexico and South America. ILLUSTRATION: Hedw. Descr. 4: pl. ro. ExsIccaTAE: Sull. Musci Cub. Wright. 70; Panels Musci Mex. 750. | BRITTON: WEST INDIAN MOSSES 669 32. RHACOPILUM TOMENTOSUM (Sw.) Brid. Bryol. Univ. 2: 719. 1827 Hypnum tomentosum Sw. Prod. 141. 1788. Hypnum tomentosum Hedw. Descr. 4: 48. 1793. Hypnum tomentosum Sw. F1. Ind. Occ. 3: 1823. 1806. Rhacopilum tomentosum var. gracile Besch. Mém. Soc. Sci. Nat. Cherbourg. 16: 257. 1872. HABITAT AND TYPE LOCALITY: ‘ On roots of trees near rivers, temperate regions of Hispaniola.”’ DISTRIBUTION: Louisiana, Bermuda, Cuba, Jamaica, Haiti, Santo Domingo, Guadeloupe, to Trinidad and South America; Mexico, Costa Rica, Guatemala, Nicaragua, and Panama; also in Asia and Africa. ILLUSTRATIONS: Hedw. Descr. pl. 19; Bryologist 10: fl. 5. ExsIccaTAE: Sull. Musci Cub. Wright. 74; Pringle, Musci Mex. 10,501. 33. CALLICOSTELLA DEPRESSA (Sw.) Jaeg. Adumb. 2: 352. 1875-76 Hypnum depressum Sw. Prod. 141. 1788. Leskea depressa Hedw. Sp. Musc. 215. 1801. Leskea depressa Sw. Fl. Ind. Occ. 3: 1804. 1806. HABITAT AND TYPE LOCALITY: ‘‘On bark of trees, mountains of Jamaica.” DIsTRIBUTION: Jamaica, Cuba, Porto Rico, Haiti, and Guade- loupe. ILLustraAtiIons: Hedw. Sp. Muse. $l. 53. f. 1-7 (from Swartz’ type). 34. Clastobryum trichophyllum (Sw.) E. G. Britton, comb. nov. : PLATE 25 Hypnum trichophyllum Sw. Prod. 141. 1788. Hypnum trichophyllum Hedw. Sp. Musc. 274. 1801. Neckera trichophylla Sw. Fl. Ind. Occ. 3: 1798. 1806. Lepyrodon trichophyllus Mitt. Jour. Linn. Soc. 12: 422. 1869. Leucodon trichophyllus Jaeg. Adumb. 2: 122. 1877. Lepyrodon trichophyllus robustior Besch. Ann. Sci. Nat. VI. 3: 224. 1876. 670 BRITTON: WEsT INDIAN MOSSES Palamocladium trichophyllum C. Miill. Flora 82: 465. 1896. Palamocladium trichophyllum subtile C. Miill. Hedwigia 37: 240, 1808. Orthothecium trichophyllum Fleisch. F1..Buit. 3: 667. 1906. Plants light yellowish green, glossy; stems rooting and creeping, with simple erect branches, often 2 cm. high and prolonged into slender flagellate branchlets bearing foen septate gemmae in clusters in the axils of the upper leaves; branch-leaves crowded, spreading, glossy, strongly plicate when dry, lanceolate-acuminate, 3-5 mm. long, ecostate, margins plane, serrate; cells linear, walls porose, slightly thickened, alar cells shorter and broader, curved, forming a small, serrate auricle. Autoicous, perichaetial leaves shorter, paler, more suddenly subulate, more sharply serrate. Seta erect, straight or flexuose, red, 15-25 mm. long; calyptra cucullate; capsule erect, ovoid-cylindric, sometimes contracted below the mouth when dry, 2-3 mm. long, lid rostrate; annulus none; walls with irregular square or hexagonal cells 27-54 mu long X 27 wide; neck short, stomatose; peristome double; teeth incurved, brown, narrow, not perforate, papillose, with slightly t trabeculate lamellae; endostome paler, also papillose with a short minutely papillose, unequal in size, 5u-16, maturing in winter. Forming bright glossy mats in shade on trunks and roots of tree-ferns and palms on high mountains, rarely on rocks. Fruit rare! HABITAT AND TYPE LOCALITY: “On bark and trunks of old trees, Jamaica.” DISTRIBUTION: Jamaica, Cuba, Porto Rico, Haiti, Santo Domingo, St. Kitts, Dominica, Martinigue. Guadeloupe, St- Vincent, Montserrat, and Trinidad to Venezuela. ILLustraTIons: Hedw. Sp. Muse. pl. 71; E. & P. Nat. Pf. xo: 7942); Sook ExsiccaTarE: Husnot, Pl. Ant. Fr. 183, as Meteorium sericeum Sch On account of the rarity of its fruit this species has been placed in a variety of genera none of which seem to me to be correct. Its double peristome and different habit remove it from Lepyrodon - and its tropical distribution from Orthothecium, the species of which are alpine or arctic and subarctic. Its relationship however seems to me to be more with the Entodontaceae, where Fleischer BRITTON: WEsT INDIAN MOSSES 671 has placed it; the presence of septate gemmae, and the ecostate leaves and more or less imperfect endostome, show its relationship to Clastobryum indicum Dozy & Molk. as figured by Brotherus (E. & P. Nat. Pfl. 13: 874. f. 640. 1907) but the leaf cells are porose and the walls are thickened as shown on the same page in f. 639 of C. planulum Mitt. Clastobryum americanum Cardot, originally described from Mexico, also occurs on the slopes and summit of Sir John Peak above Cinchona, in the Blue Mountains of Jamaica, and Mr. R. S. Williams has collected it in Bolivia at 8,000 ft. near Cargadera in 1902. 35- THUIDIUM MICROPHYLLUM (Sw.) Jaeg. Adumb. 2: 251. 1876-77 Hypnum microphyllum Sw. Prod. 142. 1788. Hypnum microphyllum Hedw. Sp. Musc. 269. 1801. Hypnum microphyllum Sw. Fl. Ind. Occ. 3: 1821. 1806. Hypnum calyptraium Sull. Pac. R. R. Rep. 4: 190. 1856. HABITAT AND TYPE LOCALITY: “‘On roots of trees, Jamaica.” DISTRIBUTION: Canada to Florida and the Bahamas, Jamaica, Cuba, and Mexico. ILLUSTRATIONS: Hedw. Sp. Musc. #/. 69 (from Swartz’ type); Sull. 1. c. pl. roo. ExsIccaTAE: Sull. Musci Cub. Wright. 99. 36. SEMATOPHYLLUM CAESPITOSUM (Sw.) Mitt. Jour. Linn. Soc. 12: 479. 1869 Hypnum caespitosum Sw. Prod. 142. 1788. Leskea caespitosa Hedw. Sp. Musc. 233. 1801. Leskea caespitosa Sw. Fl. Ind. Occ. 3: 1807. 1806. Rhaphidostegium caespitosum Jaeg. Adumb. 2: 454. 1875-76. Hypnum loxense* Sull. Proc. Am. Acad. Arts & Sci. 5: 287. 1861. Not Hooker, 1822. HABITAT AND TYPE LOCALITY: “On roots of trees, mountains of Hispaniola.” DistriBuTION: Cuba, Jamaica, Haiti, Porto Rico, Guadeloupe, * The real Sematophyllum loxense (Hook.) Jaeg. has been found in Cuba. 672 BriTTON: WEstT JNDIAN MOSSES and Martinique, to Trinidad and South America; also Mexico and Costa Rica. ILLUSTRATION: Hedw. Sp. Musc. #!/. 40. 37. SEMATOPHYLLUM PUNGENS (Sw.) Mitt. Jour. Linn. Soc. 12° 477. 1869 Hypnum pungens Sw. Prod. 142. 1788. Hypnum pungens Hedw. Sp. Musc. 237. 1801. Leskea pungens Sw. Fl. Ind. Occ. 3: 1806. 1806. Pungentella pungens C. Miill. Hedwigia 37: 260: 18098. HABITAT AND TYPE LOCALITY: “ Roots of trees in moist woods, mountains of Jamaica.” DISTRIBUTION: Jamaica, Cuba, Porto Rico, Virgin Islands, Guadeloupe, Martinique, and Dominica to South America; also Mexico and Guatemala to Panama. ILLUSTRATION: Hedw. Sp. Musc. pl. 60 (from Swartz’ type). EXsICCATAE: Sull. Musci Cub. Wright. 104; Husnot, Pl. Ant. Fr. 786. 38. PLEUROPUS CONGESTUS (Sw.) Broth. E. & P. Nat. Pfl. 1: 1138. 1908 Hypnum congestum Sw. Prod. 142. 1788. Hypnum congestum Hedw. Sp. Musc. 283. 1801. Leskea congesta Sw. Fl. Ind. Occ. 3: 1809. 1806. Homalothecium congestum Jaeg. Adumb. 2: 311. 1877-78. HABITAT AND TYPE LOCALITY: ‘‘On old trunks of trees, interior of Jamaica.” : DISTRIBUTION: Jamaica, Haiti, Montserrat, and Dutch Guiana. ILLUSTRATION: Hedw. Sp. Muse. pl. 74. f. 4-7. 1801. Excepting for the illustration given by Hedwig, little is known of this species in modern times. Mitten and Brotherus had not seen specimens. At the British Museum there is a specimen labelled ‘‘Leskea congesta Sw. Ind. Occ. ex Cl. Swartzio. J. Vahl,” which is evidently a mixture of Palamocladium leskeoides and Clastobryum trichophyllum. Hedwig’s description calls for a plant with entire somewhat secund, falcate leaves and a horizontal capsule, characters which do not agree with either of the species named above. BRITTON: West INDIAN MOSSES 673 The synonymy of Palamocladium is as follows: Palamocladium leskecides (Hook.) E. G. Britton, comb. nov. Hookeria leskeoides Hook. Musc. Exot. pl. 55. 1818. Leskea Bonplandi Hook.; Kunth. Syn. Pl. Aeg. 1: 61. 1822. Hypnum Bonplandi C. M. Syn..2: 463. 1851. | Homalothecium Bonplandi Jaeg. Adumb. 2: 379. 1875-76. Palamocladium Bonplandi Broth. Bot. Jahrb. 24: 281. 1897. Isothecium Bonplandi haitense Ren. & Card. MS. in herb. Pleuropus leskeoides Hook. MS. in Herb. 39. ORTHOSTICHOPSIS TETRAGONA (Sw.) Broth. E. & P. Nat. Pfl. 1°: 805. 1906 Hypnum tetragonum Sw. Prod. 142. 1788. Hypnum? tetragonum Hedw. Sp. Musc. 246. 180!. Hypnum? tetragonum Sw. FI. Ind. Occ. 3: 1833. 1806. Pterigynandrum aureum Brid. Mant. tot. 1819. Pterigynandrum quadrifarium Brid. Bryol. Univ. 2: 194. 1827. Isothecium tetragonum Brid. Bryol. Univ. 2: 377. 1827. Neckera quinquefaria C. Mill. Syn. 2: 124. 1850. Neckera tetragona C. Miill. Syn. 2: 125. 1850. Meteorium tetragonum Mitt. Jour. Linn. Soc. 12: 431. 1869. Pilotrichella tetragona Besch. Mém. Soc. Sci. Nat. Cherbourg 16: 224. 3872. HABITAT AND TYPE LOCALITY: ‘‘On trunks of trees, near summits of mountains in Jamaica.” DISTRIBUTION: Jamaica, Cuba, Santo Domingo; to Trinidad and Guiana; also in Mexico, Guatemala, Honduras, Nicaragua, Costa Rica, and Panama. ILLUSTRATION: Hedw. Sp. Musc. #l. 63. 1801 (from Sweartx’ type). This moss is not uncommon in Jamaica and was known to Hans Sloane* and Dillenius,t who called it ‘the square-branched Hypnum from Jamaica.” Both of these authors figured it rather poorly. * Hist. Jam. 1: 68. pl. 25. f. 3. 1707. tT Hist. Musc. 335. pl. 43. f. 73. X741- 674 BRITTON: WEstT INDIAN MOSSES 40. SCHLOTHEIMIA TORQUATA (Sw.) Brid. Bryol. Univ. 1: 323. 1826 Hypnum torquatum Sw. Prod. 142. 1788. Hypnum torquatum Hedw. Sp. Musc. 246. 1801. Neckera torta Sw. Fl. Ind. Occ. 3: 1800. 1806. Schlotheimia torta Schwaegr. Suppl. 12: 39. 1816. Schlotheimia pellucida C. Mill. Bull. Herb. Boiss. 5: 561. 1897. Schlotheimia undato-rugosa C. Miill. Hedwigia 37: 238. 1898. HABITAT AND TYPE LOCALITY: ‘‘On old mossy trunks of trees in woods, mountains of Jamaica.” DISTRIBUTION: Jamaica and Cuba, 5,000-6,000 ft. alt. ILLUSTRATIONS: Hedw. Sp. Muse. $l. 63. f. 4-7 (from Swartz’ type). ExsIccaTAE: Sull. Musci Cub. Wright. 52. 41. MACROMITRIUM cCIRRHOsUM (Sw.) Brid. Bryol. Univ. 1: 316. 1826 Hypbnum cirrhosum Sw. Prod. 142. 1788. Anoectangium cirrhosum Hedw. Sp. Musc. 42. 1801. Neckera cirrhosa Sw. Fl. Ind. Occ. 3: 1802. 1806. Schlotheimia cirrosa Schwaegr. Suppl. 3}. 1827. HABITAT AND TYPE LOCALITY: ‘‘On trunks of trees, temperate parts of Jamaica.” DISTRIBUTION: Jamaica, Gabe: ‘Haiti, Santo Domingo, Porto Rico, St. Kitts, Guadeloupe, Martinique, Montserrat, and Trinidad to South America; also, Guatemala and Panama. ILLUSTRATIONS: Hedw. Sp. Muse. pl. 5; Schwaegr. Suppl. - 201A. EXsICCATAE: Sull. Musci Cub. Wright. 52; Husnot, Pl. Ant. Fr. 144. 42. ‘THUIDIUM INVOLYENs (Hedw.) Mitt. Jour. Linn. Soc. 12: 575- 3 1869 Leskea involuens Hedw. Descr. 42275-17048. Leskea involvens Swartz, Fl. Ind. Occ. 3: 1815. 1806. HABITAT AND TYPE LOCALITY: ‘With Helicodontium capillare on trunks of trees, interior of Jamaica.” BrittON: West INDIAN MOSSES 675 DISTRIBUTION: Jamaica, Cuba, Haiti, Porto Rico, Guade- loupe, and Barbados to South America; also Mexico (Yucatan). ILLUSTRATION: Hedw. Descr. pl. 11 (from Swartz’ type). ExsiccaTAE: Sull. Musci Cub. Wright. 98. 43. Turckheimia linearis (Sw.) E. G. Britton, comb. nov. Tortula linearis Sw. Fl. Ind. Occ. 3: 1765. 1806. _ Barbula linearis Brid. Mant. Musc. 88. 1819. Trichostomum lineare Broth. E. & P. Nat. Pfl. 1°: 394. 1902. HABITAT AND TYPE LOCALITY: “‘On dry calcareous rocks, His- paniola.”’ DISTRIBUTION: Jamaica, Cuba, and Haiti. Our spécimens from Jamaica and Cuba have a well-developed, slender peristome, which disappears from the old capsules. I believe this species to be congeneric with Turckheimia guate- malensis Broth., which also shows traces of a peristome though the capsules are all old. The section of the leaf in T. linearis is remarkable for having two rows of guide-cells of about Io cells each in the costa, with a stereid band both aboveand below. The costa is rather broader thanin T. guatemalensis and smooth on the dorsal side, showing as a prominent white rib tothe leaf. It is papil- lose on the upper surface and the cells of the blade bear several minute papillae on both surfaces. This peculiarity of the costa removes Turckheimia linearis from Trichostomum; and although there is but a single row of guide-cells in T. guatemalensts, their macroscopic resemblance is so close that they appear to be congeneric. 44. IsopTERYGIUM TENERUM (Sw.) Mitt. Jour. Linn. Soc. 12: 499. 1869 Hypnum tenerum Sw. Fl. Ind. Occ. 3: 1817. 1806. HABITAT AND TYPE LOCALITY: ‘“‘On trunks of trees, mountains of Jamaica.” DIstTRIBUTION: Jamaica, Cuba, Haiti, Guadeloupe, Martinique, St. Lucia, to Trinidad and South America; also Bermuda, and Louisiana to Florida. ExsiccaTaE: Sull. Musci Cub. Wright. 107. 676 Britton: West INDIAN MOSSES According to Dr. Andrews’ notes ‘‘the Swartz specimens, which are deposited in the collections of the Naturhistoriska Riks- museum at Stockholm, are distributed through the herbarium of non-Scandinavian mosses, which are, in general, arranged after Paris’s Index. Packets are generally uniform, one to many on the herbarium sheet. Swartz’ specimens are recognizable by labels in his handwriting included in the packet, by the kind of paper with water-mark to which he pasted them and references of others to the origin of specimens.” We have seen specimens of all but two of these species, and have duplicates of many of them; it is therefore our intention to distribute sets of these and other West Indian mosses, in exchange for other exsiccatae and duplicates from the West Indies, Central America, and South America. NEw York BOTANICAL GARDEN Explanation of plate 25 Clastobryum trichophyllum (Sw.) E. G. Britton The figures were drawn from magnifications three times as great as expressed in the numbers, which represent the magnifications of the figures as they stand in the Or tee lant, natural size. 2. Portion of branch reser rene flagellate branches and gemmae, xX 23%. 3- Outline of stem leaf, XK 1 4,5. Outlines of sare sabes X 1634. 6. Apex of leaf, * 1 7. Basal portion of co showing the auricle, « 108. 8. Median cells, & 26 9. Apex of leaf FSH Ks the = in the walls of the apical cells, X 263. Cross section of leaf, X 1 11. Branch with gemmae, x aE 12. Gemma, X 138. 13. Cross section of stem, & 1 14. Perichaetial bud, leaves of one side removed to show the paraphyses and I 17. Stoma from base of capsule, X 19 18. Portion of perenanne and upper st of capsule, X I95. 19. Spores, X 195. Phytogeographical notes on the Rocky Mountain region |, Alpine region P, A. RYDBERG The alpine region, roughly speaking, is the region between the perpetual snow and the timber line. THE UPPER LIMIT, THE PERPETUAL SNOW LINE A perpetual snow line cannot be spoken of in the southern Rockies. Even the highest peaks do not have a perpetual snow cap like Mt. Shasta or Mt. Hood. This is probably due to the less amount of moisture and precipitation. It is true that many of the peaks have perpetual snow on them, but this snow is mostly in the form of snow-drifts and small glaciers, especially on the northern or northeastern side. The amount of snow depends to a great extent on local conditions, as for instance on an exposure > to the northwestern winds or partial protection from the direct action of the summer sun. The Snowy Range of Colorado has more snow than the much higher Gray’s Peak, Sierra Blanca, or Mount Massive. In the Canadian Rockies and especially in the Selkirk Mountains the conditions are different and more like those of the European Alps. There the highest peaks have a perpetual snowcap and the glaciers extend far down in the valleys. In northern Montana, as for instance in the Sperry Glacier region, are found the only places in the United States where in the Rockies there are glaciers of any great extent, notwithstanding the fact that the Montana mountains are considerably lower than those of Colorado. ; THE LOWER LIMIT, THE TIMBER LINE The timber line is by no means a well-defined boundary line. It is in reality a broad zone in which the woody vegetation gradually thins out from the dense forest to the last krumholz. In nature there is not found any sharp line between two regions, but only a gradual transition zone between them. 678 RYDBERG: PHYTOGEOGRAPHICAL NOTES Different authors have fixed the timber line differently, as for instance: 1. At the forest line; i. e., where the continuous forest stops. 2. At the grove line; i. e., where the trees cease to form com- munities of larger or smaller size. 3. At the tree line; i. e., where the arboreal species cease to form trees. 4. The absolute timber line: i. e., where these species disappear altogether, even as krumholz. To me it seems superfluous to consider more than two of these “lines,” viz. the ‘forest line” and the ‘‘absolute timber line,” which may be called the Lower and the Upper Timber Lines. But to fix the limit between the alpine region and the subalpine region at either of the two would be erroneous in certain respects. The region between these two timber lines is a transition zone between the two, or it may be still better called a zone of strife. A continuous warfare goes on between the forest and the alpine grass- land. A seed from the forest succeeds in germinating between the low alpine plants. A tree grows up. The alpine plants are smothered in the shade. More tree seeds have a chance to germi- nate and a grove is formed and the forest region is carried upwards. On the other hand, snow and wind kill the trees on the edge of the forest or the grove, the shade is gone and the alpine plants soon take possession. But more on this subject below. The width of this transition zone or zone of strife depends on many factors. In one place the forest meets a steep cliff and stops abruptly. In such places there is no transition zone. In other places the lower timber line has been pushed down by a ledge of snow thick enough to last the larger part of the summer and having the power of smothering the trees, but not the herbaceous alpine vegetation. Still above the lower timber line at these places, there might be found isolated trees or groves of trees a thousand feet higher up, especially on higher ground, where the snow has not been so deep. In other places the lower timber line might have been pushed down by wind, not so much by its mechanical force as by its desiccating effects. In treating the alpine region, I would be inclined to place the boundary at the lower timber line, i. e., the forest line, so as to RYDBERG: Puy APHICAL NOTES 679 include all open spaces of the transition zone as these have a flora alpine in character. In treating the subalpine region, how- ever, I would place the boundary at the absolute timber line, so as toinclude the groves, isolated trees, and krumholz as well. The groves, if of any size, contain not only the trees themselves, but also wood plants and underbrush belonging to the subalpine region. They are either encroaching on the region above or are themselves remnants of a former forest. So are also the isolated trees and krumholz, although not associated with other plants of the forest. In a mountain region extending through twenty degrees of latitude, from lat. 35° to lat. 55° (the Rockies north of 55° have not been considered in my work), the altitude of the timber line necessarily varies greatly. In Colorado the lower timber line is found at an altitude of between 3,200 and 3,400 m. and the upper at 3,400 to 3,500 or rarely 3,600 m. In Montana the lower one is at 2,200-2,500 m. and the upper 2,500-2,700 m. In the Cana- dian Rockies they are even lower. FACTORS GOVERNING THE TIMBER LINE The conditions that have been given as causing or modifying the timber line are: 1. A decreased temperature during the growing season. 2. Too short a growing season. Late frost on account of lack of protection from snow. Strong desiccating winds. Deep snow. Form of precipitation. Large mountain masses. Exposure to and protection from direct sunlight. 9. Physiographical barriers. 10, Ecological barriers. 11. Economic timber line. « PI AKL SG . Low TEMPERATURE It is natural that too low a temperature should be one of the important factors causing the disappearance of the forest. The temperature during the winter has, however, very little influence 680 RYDBERG: PHY APHICAL NOTES upon the growing of trees. It is shown that the temperature in temperate regions in the winter often is much lower than in many places in the arctic. It is doubtful if the temperature in the alpine region of Colorado ever becomes as low as on the plains of Mon- tana, where some years ago it was recorded as 65° F. below zero. The only place within the alpine region of the Rockies where a record has been kept during the winter is on the top of Pikes Peak, and here only for a few vears. No such low temperature has been recorded there. It is during the growing season that a low temperature limit for forest growths can be spoken of, for during the winter, when the life functions of the plants lie dormant, a few degrees more or less makes in reality no difference. Képpen claims that no trees can grow at a place where the mean temper- ature during the warmest months of the year does not reach 10° C. Schroeter gives a table taken from the records of the meteorological central station at Ziirich in which are given the mean temperatures in July at fifteen stations at the timber line in Switzerland. The mean temperature ranges from 7.75° C. at Zermatt to 15.4° at Monte Generoso. This shows that the timber line may reach a little higher than the isotherm 10° C. and in other cases not reach it. Schroeter is particular enough to mention that at Monte Generoso the low timber line is not an artificial one made by man. See below. The arctic timber line seems to be more coincident with the isotherm 10° C. for July than the timber line in the mountains. SHORT GROWING SEASON Another cause of the timber line is the shortness of the growing season. This, it may be, is just as important a factor as the pre- ceding. When speaking of the arctic timber line, it is easy to see that this factor acts parallel to the preceding, for near the sea- level places of the same isotherm in the summer have about the same length of summer, but not so in the mountain regions. In the heads of valleys, where big snowdrifts are formed during the winter and melt late in the summer, and along glaciers and per- manent snow the frost is kept longer in the ground and the growing season is naturally shortened. Therefore, in many places in the Rockies, the timber line is a thousand feet or more lower in the valley heads than on the slopes on the sides. The shortness of RYDBERG: PHYTOGEOGRAPHICAL NOTES 681 the season may have also another effect on the timber line, i. e., the seeds would not have time to ripen. This may be of great importance in accounting for the arctic timber line, but it can have very little influence on the alpine timber line, for most of the conifers that reach the timber line have winged seeds, which are easily carried above the line of maturing seed and then can germinate there. LATE FROST Another factor which has been given as having effect on the timber line is late frost in the spring, killing the new sprouts. As the conifers, which are most affected, do not readily produce a second crop of shoots the same season, the forests after a few repeated frosts will soon be killed. In such a way large districts of pine forest were destroyed in Montana a few years ago. STRONG DESICCATING WINDS One of the most important factors is strong wind. This factor has been much underestimated in earlier times, but later writers on the phytogeography of the arctic regions have recognized it more and more. In my belief it is one of the most important factors in the Rockies. The trees at the timber line and especially those few isolated stragglers above the real forest line show marked effects from the wind. The trees are not only low, stunted, gnarled, ragged, with enormously elongated lower branches often spreading on the ground, but conspicuously one-sided, telling at the glance the direction of the prevailing winds. But the me- chanical influence of the wind is not the most important, however. Of greatest importance are its desiccating effects, especially in the winter. This effect of the wind has been recognized even in arctic regions, but it must be taken into consideration still more in the mountains. The timber line is much lower on the north side of the Alps than on the south side. This is due not only to the difference in temperature (for the difference in altitude should not be so great), but still more to the desiccating northern winds. In some places in Montana these winds are northerly, but in southeastern Colorado and southern Utah they are from the south- west, and it is on this side of the mountains that the timber line is the lowest. In the Abajo Mountains of southeastern Utah, for 682 RYDBERG: PHYTOGEOGRAPHICAL NOTES instance, there is no timber line at all on the southern and western sides, for no timber is growing between the semi-arid cedar-pinyon belt and the top of the mountains. The whole southern and western slopes of the mountains proper are covered by a semi-arid grass formation. The highest peaks (altitude about 11,000 feet) just reach the timber line on the eastern and northern sides, only one or two hundred feet belonging to the alpine region. The desic- cating effects of the winds are increased by the thinness of the atmosphere. DEEP SNOW Deep snow is also a factor. As the desiccating wind lowers the altitude on the wind-swept ridges so does the snow in the heads of the valleys. I have already mentioned that the great snow- drifts or glaciers here shorten the growing season. But the snow- drifts have also a direct mechanical influence on the timber line in the way of smothering the tree vegetation. Herbs and low shrubs can withstand being covered by snow much better than a tree, for their growing season does not begin before the snow is practically off the ground, while the tops of the trees may be above the snow and exposed to the summer heat months before the snow cover of their roots and lower branches has melted. The lower portion of the tree is cut off from the air while the upper portion is already in vital activity. It is easy to distinguish trees stunted by the action of the wind from those stunted by the smothering snow. In the former the lower branches are enormously developed compared with the upper, and often creeping along the ground, while in the latter the lower branches are dead and covered by fungi or their mycelia. The usual condition in the Rockies is, that wherever there is 2 large valley head, where the snow has a chance to lodge, this is always devoid of trees, except in places of higher ground, where the snow-drift has not been so deep and has had time to melt earlier in the summer. On such higher places there are often groves or isolated trees. The absence of trees in such a valley head is due less to the shortness of the season, produced by the snow, than to the smothering of the tree vegetation. RYDBERG: Puy APHICAL NOTES 683 FORM OF PRECIPITATION Another important factor influencing the timber line is the form of precipitation. In high altitudes the air is too rare to hold much moisture and the rain falls at the least lowering of the tem- perature. The rain falls therefore either in the form of mists or in light showers, which only wet the surface of the ground. It may be sufficient to keep alive the low rosettes or cushions of the alpine vegetation, but it is not sufficient for the deep-rooted trees. Furthermore, if a little heavier rain should come, the water would rush down the steep slopes of the mountains, not having time to sink down into the ground. The tops of the mountains are there- fore arid, because the air is too rare to hold much moisture and quickly gives it up in light showers. Nowhere in the Rockies proper is the moisture very great. In the foot-hill regions and on the surrounding plains the temperature is too high in the summer to allow any precipitation. These zones are therefore also arid. It is at middle elevations that the precipitation is the greatest. The air here is dense enough to hold more moisture and the tem- perature low enough to allow precipitation. It is also at middle altitudes that we find the forest areas in the Rocky Mountain region. LARGE MOUNTAIN MASSES In the Swiss Alps, observations have been made that in regions of large mountain masses, as for instance in the Monte Rosa region and the Engadine and others, the timber line is higher than on isolated mountains. I have not seen any satisfactory explanation of this fact. It may be due partly to the fact that the central mountains of such massed groups are more or less pro- tected from the desiccating winds. It may be due also to the circumstances that in the winter more snow lodges between the mountains, the melting of it is more retarded, and the water is more arrested in its downward course by the trees and their roots. The air in the summer time would be therefore, from the evapo- ration, more loaded with moisture, which would naturally also benefit the mountain tops. Whatever the real cause may be, it seems as if the observations made in Switzerland hold good in the Rockies. From my own experience, I know that the timber line in the isolated Belt Mountains and Crazy Mountains in Montana 684 RYDBERG: PHYTOGEOGRAPHICAL NOTES is much lower than in the main Rockies, as for instance in the Yellowstone Park. The Belt Mountains would not be high enough to have a timber line if they were in the Rockies. So also in the Wahsatch and La Sal Mountains, the timber line is much lower than in the Rockies of Colorado. Even in the Colorado Rockies themselves, the timber line seems to be higher in places where the mountains are more massed. So for instance is it higher on Mount Massive and other mountains around Leadville than on the more isolated Pikes Peak, Sierra Blanca, or Longs Peak. I understand that on the isolated Mount Shasta, the timber line is much lower down than on the peaks of the Sierra Nevada, but here it may depend upon the proximity of the ocean, the greater moisture, and the consequently larger snowcap on Mount Shasta. EXPOSURE TO SUNLIGHT Exposure to the direct sunlight and protection from it evi- dently also have influence on the altitude of the timber line, though perhaps not so much as one might expect. The insolation on the mountain tops in direct sunlight is very great. Schroeter esti- mates that on the top of Mont Blanc it is 26 per cent stronger than in Paris. The Rockies of Colorado have about the same height as Mont Blanc and are situated from 5° to 8° farther south, and the insolation is fully as great. The amount of light and heat which can be absorbed by the plant is therefore much greater on the mountain tops than on the plains. The radiation is also very great in the higher altitudes so that the temperature in the shadow is much lower. According to Schroeter the timber line lies 100-200 meters higher on the southern side than on the northern, and DeCandolle claims that the limiting line of vege- tation, of plants in general, is at an average of 200-300 meters higher on the equatorial side. These statements cannot be veri- fied in the Rockies. The timber line in Colorado is, perhaps, higher on the northern side, but this is probably due to other conditions. The timber line trees of Colorado are mainly Picea Engelmannii, Abies subalpina, and Pinus aristata. The first two are trees that need a great deal of moisture, and their seedlings require shade. These two trees are therefore more confined to the more shady and wetter northern slopes, where they also RYDBERG: PHYTOGEOGRAPHICAL NOTES 685 extend higher up. Pinus aristata is a tree that stands much more drought and is found more on the southern slopes, but it is a tree of little value as a forest tree, growing scatteringly only. Tome it appears to be a species which has passed its best wadaondd oizalcty and is in process of dying out. PHYSIOGRAPHICAL BARRIERS One of the conditions modifying the altitude of the timber line is to be found in physiographical barriers. Among these may be counted snowdrifts and glaciers, but these have been already mentioned. Besides these, the most important are precipitous cliffs and rock-slides. Very little needs to be said about these barriers. Neither gives the forest trees a chance to grow. Meeting oneof these barriers, the timber may cease to grow thousands of feet below’ the physiological timber line. Wherever a steep cliff arrests the forest, many of the alpine plants will be found growing in the crevices, hundreds or even a thousand feet lower than usual, and there are a few plants characteristic of the rock-slides. These may be best included in the alpine vegetation. ECOLOGICAL TIMBER LINE Sometimes an ecological timber line is mentioned, i. e., where bacteria in the soil and other organisms necessary for the growth of trees cease to exist. Theoretically, I can easily see that such a timber line may exist, but practically I have no information that such a one is found in the Rocky Mountains, distinct from the merely physiological one. No investigation in this line has been made. ECONOMIC TIMBER LINE In Switzerland there exists also an economic timber line. The alpine meadows are there used as summer pastures for sheep and goats. These animals make depredations on the young trees and hinder the spreading of the forest, but in many places the subalpine forest is actually cut down by men to make room for more pastures. In either case the alpine conditions will be brought further down the mountains and the timber line lowered. Such an economic timber line cannot be said to exist in the Rockies. 686 RYDBERG: PHYTOGEOGRAPHICAL NOTES ALPINE VEGETATION After having discussed the causes of the timber line, it is easier to define what an alpine plant is. In short, it is a plant that can endure the climate of the mountains above the timber line. It is a plant that requires less heat during the growing season than the forest trees, or that can survive a shorter growing season, or is less affected by frost, and besides can better withstand desiccating winds, deep snow, reduced precipitation, etc., or a combination of such conditions. Some authors claim that alpine and arctic plants are xerophytes, but they are not necessarily so. While most of the plants of alpine and arctic regions can withstand a great deal of drought, in fact are xerophytic plants, it is not the case with all. Not a few of the arctic-alpine plants require a great deal of moisture, growing only below and around snowbanks, or in springy or boggy ground, as for instance several species of Ranunculus, Saxifraga (in extended sense), Salix, and many grasses and sedges. There are in the arctic-alpine regions even true aquatics, as for instance among the phanerogams, Catabrosa aquatica, Phippsia algida, Sparganium minimum, and S. hyperboreum, and a few species of Potamogeton. NEw YorxkK BOTANICAL GARDEN New ferns from tropical America—lll MARGARET SLOSSON (WITH PLATE 26) Trichomanes rhipidophyllum Slosson, sp. nov. Rhizome creeping, about 5 mm. in diameter, thickly tomentose; stipes brown and tomentose to within about 1 mm. of the lamina, then green, slightly hairy, and winged by the narrowly and abruptly decurrent base of the lamina, in fertile fronds 5-7 mm. long, in sterile 1.5-3 mm. long; laminae shining bright green, delicately papyraceous, the fertile 0.8-1.1 cm. long, 0.7-1.3 cm. broad, suborbicular, almost semicircular, or subtrapezoid, at apex slightly once cleft, at base subtruncate or cuneate, the sterile 0.4-1.3 cm. long, 0.5-1.5 cm. broad, round-reniform to ovate or broadly obovate; margins irregularly undulate or sublobate, marginal hairs few, slender, simple or binate; surfaces or at least the lower one with a few short club-shaped mostly 2-celled hairs on the veins; veins few, distant, in the sterile fronds subflabel- lately forked, midveins of fertile fronds flexuose, with about 3 pairs of branches forked 1-4 times, spurious veins short, almost none; cells of the lamina between the veins variable, mostly narrow, 0.056- 0.116 mm. long, 0.020-0.040 mm. broad, their walls 0.004-0.008 mm. broad; in- volucres 1-2, borne at the base of the central cleft of the lamina, almost wholly = Trichomanes rhipidophyllum. exserted, the lips 1.5 mm. broad, their 1, cellular structure of the mar- - brown border 1-4 cells deep; spores glo- gin of the lip of the involucre, , 0.044—-0.056 mm. in diameter. nlarged; - cells ep between Type in the Underwood Herbarium rather eb aapal dowd dak! at the New York Botanical Garden, collected on a tree in a damp forest near Onaca, Colombia, at an altitude of 760 m., Aug. 24, 1898-99, Herbert H. Smith 2445. A note on the label says ‘not observed elsewhere.” a é 687 ca 688 SLOSSON: NEW FERNS FROM TROPICAL AMERICA not likely to be mistaken for any other. Marked by its bright shining green color, rounded, undulate or not more than sublobate margins, and few flabellately forked veins tapering toward the apex. From Trichomanes sphenoides Kunze, which also has fla- bellately forked veins tapering toward the apex, it may be easily distinguished by the greater distance between the tips of the true veins, varying from .5 to 2 or 2.5 mm., and by the very thick walls of the cells of the laminae. Polystichum machaerophyilum Slosson sp. nov. Rhizome erect, its scales light brown, 2.5-6 mm. long, ovate to oblong, fimbriate or subfimbriate; scales of lower part of the stipes similar, passing into minute scales scattered over both sur- faces of the frond, which are elongate-caudate, entire or slightly toothed, from a short more or less fimbriate-ciliate base; fronds up to 51 cm. long, up to 7.1 cm. broad; stipes 2.5-20.5 cm. long, brownish-stramineous or greenish, channeled on the face, rounded at the back, the upper part channeled also or flattened on the sides; rachis similar to upper part of the stipe; laminae brownish olive, lanceolate or lance-linear, slightly tapering toward base, pinnate, the apex in young fronds often deltoid-acuminate, in mature fronds of two kinds, the first subcaudate, elongate-linear or lanceolate, subentire or undulate or serrate-lobate, often pro- liferous at tip, the second flagelliform and proliferous at tip, naked or with a few small scattered pinnae, both kinds sometimes on the same frond, the second terminating the first; pinnae mostly alternate, 8-27 to a side, usually approximate or distant, the lower short-stalked, the upper sessile or subsessile or adnate, often passing into short variously shaped often obovate segments, principal pinnae broadly cuneate-hastate, above the auricles deltoid to linear, the basal margins entire, the outer margin sub- entire or undulate or with one or more minute occasionally spines- cent teeth or in large fronds irregularly crenately lobed, the lobes sometimes mucronate, basal auricles and apex of pinnae spinescent; texture coriaceous-chartaceous; venation pinnate, obscure, veins very oblique, about 1-4 times forked, the basal auricles with distinct midveins; sori usually about midway between the midvein and the margin of the frond, or nearer the midvein, rarely a few submarginal; indusia more or less erose or with a few cilia; spo- rangia glabrous; spores cristate. Type in the Underwood Herbarium at the New York Botanical Garden, collected at Arroyo del Medio, above the falls, Sierra SLosson: NEW FERNS FROM TROPICAL AMERICA 689 Nipe, Oriente, Cuba, altitude 450-550 meters, December 22, 1909, J. A. Shafer 3262. The label reads: ‘‘Shaded rocks near water.” This plant is closely related to both Polystichum ilicifolium Fée and P. triangulum (L.) Fée, and very likely may be a hybrid between the two. Numerous specimens have been seen, all Cuban, and excepting Wright’s specimens, all from the Province of Oriente. Wright’s specimens bear the indefinite inscription of ‘“‘Cuba’’ and ‘‘Cuba orientale,’ but are dated 1859, 1860, and 1865. During the first two years Wright is known to have collected in the Province of Oriente, but in 1865 he is believed to have collected only in the western part of Cuba.* | Polystichum machaerophyllum in a mature state is easily dis- tinguished from P. triangulum by the peculiar apices of the fronds, varying from long-drawn-out to flagelliform, non-proliferous to proliferous. It is more likely to be confused with P. tlicifolum, but may be known by the proportionately broader and shorter laminae; their darker olive-green color, resembling that of P. triangulum; and the larger and longer pinnae, distinctly biauricu- late at base, with the part above the basal auricles not short and margined with large sharp oblique spinescent teeth, as in P. ilici- folium, but more or less extended and subentire or very slightly toothed or crenately lobed, the lobes entire or minutely mucronate. The indusia are peculiar, varying from only slightly erose to markedly so with a few cilia. The indusia in P. triangulum are entire, and in P. ilicifolium vary from markedly erose to con- spicuously long-ciliate. P. decoratum Maxon, the only other Cuban Polystichum known with fronds flagelliform at apex, may be readily recognized by its pinnae widely excised, not auricled, at base on the lower side. The following specimens of P. machaerophyllum at the New York Botanical Garden and in the U. S. National Herbarium in Washington have been examined: Cusa: Camp La Gloria, south of Sierra Moa, Oriente, Dec. 24-30, 1910, Shafer 8096; bank of river among stones, Camp La Barga, Oriente, altitude 450 meters, February 22-26, 1910, Shafer 4127; on moist rocks, Cooper’s Ranch, base of El Yunque Mountain, Baracoa, March, 1903, Underwood & Earle 1179, 1180; * See L. M. Underwood, A Summary of Charles Wright’s Explorations in Cuba, Bull. Torrey Club 32: 298, 300. 1905. 690 SLosson: NEW FERNS FROM TROPICAL AMERICA vicinity of Baracoa, February 1-7, 1902, Pollard, Palmer & Palmer 237; “in Cuba Orientale,” 1859, 1860, C. Wright 828 in part; without specific locality, 1865, C. Wright 828 in part.* Explanation of plate 26 1-3. Trichomanes rhipidophyllum; 1, rootstocks and leaves, slightly reduced; 2, 3, sterile and fertile frond, enlarged, showing venation; H. H. Smith 2445 unnumbered figures. Polystichum MARE OP REM: 4, indusia, en- pinnae do not show clearly in the photograph, owing to a slight infolding of the dried specimens.) *In the Underwood Herbarium and in the U. S. National Herbarium the remainder of Wright’s no. 828 is P. decoratum. INDEX TO AMERICAN BOTANICAL LITERATURE 1907-1913 The aim of this Index is to include all current botanical literature written by Americans, published in America, or based upon American material ; the word Amer- ica being used in the broadest s Reviews, and papers that peice exclusively to forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, an gees is made to index the literature of bacteriology. An occasional exception is made in favor of some paper appearing in an American periodical which is devoted eee to botany. Reprints are not mentioned unless they differ from the original in some important particular. If users of the Index will call the attention of the editor *o errors or omissions, their kindness will be appreciated. This Index is reprinted monthly on a and furnished in this form to subscribers at the rate of one’ cent for each card, Selections of cards are not permitted ; each subscriber must take all cards published during the term of his subscription, Corre spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club, Bailey, V. Life zones and crop zones of New Mexico. U. S. Dept. Agr. Biol. Surv. N. Am. Fauna 35: 7-100. pl. 1-16 +f. 1-6. S 3 1614, Bartlett, H. H. Inheritance of sex forms in Plantago lanceolata. Rhodora 15: 173-178. 17 O 1913. Berry, E. W. Contributions to the Mesozoic flora of the Atlantic coastal plain—IX. Alabama. Bull. Torrey Club 40: 567-574. 15 O 1913. Blackader, E. H. The shade trees of Ottawa. Ottawa Nat. 27: 31- 36. 17 My 1913; 27: 38-40. 12 Jl 1913. Blake, S. F. A redisposition of the species heretofore referred to Leptosyne. Proc. Am. Acad. Arts & Sci. 49: 335-346. S 1913. Includes Stephanopholis gen. nov., Coreopsis parvifolia sp. nov., and many new names and combinations. Blake, S. F. A revision of Encelia and some related genera. Proc. Am. Acad. Arts & Sci. 49: 346-398. pl. 1. S 1913. Includes Simsia setosa, S. submollicoma, S. eurylepis, S. jamaicensis, and S. triloba, spp. nov. Briggs, L. J.. & Shantz,H. L. The water requirement of plants. I, Investigations in the Great Plains in 1910 and 1911. U.S. Dept. Agr. Plant Ind. Bull. 284: 5-49. pl. 1-11 + f. 1, 2. 16 O 1913; II. A review of the literature. U.S. Dept. Agr. Plant Ind. Bull. 285: .f. 7-6. 60 1913. 5-96. f. | gas 692 INDEX TO AMERICAN BOTANICAL LITERATURE Britton, E.G. Wild plants needing protection. 9. ‘‘Flowering Dog- wood”’ (Cynoxylon floridum). Jour. N. Y. Bot. Gard. 14: 133, 134. pl. r20. Jl 1913. Britton, N. L. Four undescribed West Indian sedges. Torreya 13: 215-217. 25 1913. Stenophyllus homey S. portoricensis, Fimbristylis inaguensis, and Rynchospora bahamensis, spp. n Brooks, C. ee blotch and apple fruit spot. Phytopathology 3: 249, 250. Au 1913, Brown, P. E. Methods for bacteriological examination of soils. - Centralb. Bakt. Zweite Abt. 39: 61-73. 27 S 1913. Brown, W.H. The phenomenon of fatigue in the stigma of Martynia. Philip. Jour. Sci. 8: (Bot.) 197-201. Jl 1913. Burnham, S. H. A supplementary list of plants of Copake Falls, New York. Torreya 13: 217-219. 2S 1913. Christensen, C. Filices Purdomianae. Bot. Gaz. 56: 331-338. 15 O 1913. Includes Athyrium Sargentii, Cheilanthes lanceolata, Dryopteris Purdomii, D- sericea, Matteuccia intermedia, and Polystichum gracilipes spp. nov., all from northern hina. Coons, G. H. A preliminary host index of the fungi of Michigan, . exclusive of the Basidiomycetes, and of the plant diseases of bacterial and physiological origin. Ann. Rep. Michigan Acad. Sci. 14: 232- 276. 1912. East, E.M. ‘Xenia and the endosperm of angiosperms. Bot. Gaz. 56: 217-224... 17 & 1083. Fernald, M. L. Carex tincta a valid species. Rhodora 15: 186, 187- 17 O 1913. Fernald, M.L. The indigenous varieties of Prunella vulgaris in North America. Rhodora 15: 179-186. 17 O 1913. Fink, B. Botanical instruction in colleges. Proc. Ohio Acad. Sci. 6: 72-87. 1913. Fulton, H. R., & Winston, J. R. Some important diseases of field crops in North Carolina. Bull. N. Carolina Dept. Agr. 182: 5-24- Ap 1913. Gager, C. S. Botany. Bull. Univ. Missouri Sci. Ser. 1: 149-173. J! 1913. Garrett, A. O. Some introduced plants of Salt Lake County, Utah. Torreya 13: 237-241. 14 0 1913. . INDEX TO AMERICAN BOTANICAL LITERATURE 693 Gates, F.C. The vegetation of the region in the vicinity of Douglas Lake, Cheboygan County, Michigan, 1911. Ann. Rep. Michigan Acad. Sci. 14: 46-106. pl. 5-21. 1912. Gilbert, E. M. Biologic forms of black knot. Phytopathology 3: 246, 247. Au 1913. Gilbert, W. W. Cotton anthracnose and how to control it. U. S. Dept. Agr. Farm Bull. 555: 1-8. f. 1-8. 7 O 1913. Gile, P. L. Relacién entre los terrenos calcA4reos y la clorosis de la pifia. Puerto Rico Estac. Exp. Agr. Bull. 11: 7-53. pl. 2, 2. 24 FL-29%3. An English edition of this was published 7 N I9II. Goddard, H. N. Can fungi living in agricultural soil assimilate free nitrogen? Bot. Gaz. 56: 249-305. f. 1-18. 15 O 1913. Griffiths, D. Einige neue Opuntioideen. Monats. Kakteenk. 23: 130-140. 15S 1913. [Illust.] Includes seven new species in Opuntia and one in Nopalea. (Translated from the English by F. Vaupel.) Hackel, E. Gramineae novae—X. Repert. Sp. Nov. 12: 385-387. 25 S 1613. Includes Ichnanthus Damazianus sp. nov., from Brazil. Harper, R. M. Five hundred miles through the Appalachian Valley. Torreya 13: 241-245. 14 O 1913. Harter, L. L. Foot rot, a new disease of the sweet potato. Phyto- pathology 3: 243-245. f. r, 2. Au 1913. Plenodomus destruens sp. nov. on stems of Ipomoea Batatas. Hartley, C. Bark rusts of Juniperus virginiana. Phytopathology 3: 249. Au 1913. Hartley,C. Twig canker on black birch. Phytopathology 3: 248-249. Au 1913. Hassler, E. Anacardiaceae. In Ex herbario Hassleriano:. Novitates paraguarienses. XIX. Repert. Sp. Nov. 12: 373,374- 15S 1913. Hassler, E. Compositae—II. In Ex herbario Hassleriano: Novitates paraguaiienses. XIX. Repert. Sp. Nov. 12: 367-371. 1 S 1913. Hassler, E. Leguminosae. VII. In Ex herbario Hassleriano: Novi- tates pataguarienses. XIX. Repert. Sp. Nov. 12: 371-373. 1 $ 1913. : Hassler, E. Novitates argentinae. I. Repert. Sp. Nov. 12: 201, 202. 30 My 1913; II. Repert. Sp. Nov. 12: 365-367. 1 S 1913. 694 INDEX TO AMERICAN BOTANICAL LITERATURE Hayes, H. K. The inheritance of certain quantitative characters in tobacco. Zeits. Induk. Abstammungs- und Vererbungslehre 10: 115-129. f. 1-8. Je 1913. Heald, F. D. The symptoms of chestnut tree blight and a brief description of the blight fungus. Pennsylvania Chestnut Tree Blight Commission Bull. 5: 3-15. pl. 1-16. 15 My 1913. Hedgcock, G. G., & Long, W.H. Notes on cultures of three species of Peridermium. Phytopathology 3: 250, 251. Au 1913. Hedgcock, G. G., & Long, W. H. An undescribed species of Peri- dermium from Colorado. Phytopathology 3: 251, 252. Au 1913. Peridermium Betheli sp. nov. Holm, T. Phryma leptostachva 1..,.a morphological study. Bot. Gaz. 56: 306-318. pl. 8-ro. 15 O 1913. Knowlton, C.H. Festuca octoflorain Vermont. Rhodora1s: 187, 188. 17 O 1913. Knowlton, C. H., and others. Reports on the flora of the Boston district, XV. Rhodora14:107-113. 14Je1912;--XVI. Rhodora 15: 54-59. 12 Ap 1913;—XVII. Rhodora 15: 122-132. 1 1913;— XVIII. Rhodora 15: 144-151. ‘11 Au 1913. Knudson, L. Imbedding and warming stand. Bot. Gaz. 56: 339, 340- J. 4, 2. 15 O 1913. Kuyper, J. Veslag van den plantkundige. Vers. Dept. Landbouw Suriname 1912: 6-20. 1913. Includes a list of fungi found during the year. Lindau, G. Einige neue Acanthaceen aus Zentralamerika. Repert. Sp. Nov. 12: 423-426. 25 S 1913. Lloyd, C. G. Synopsis of the genus Cladoderris. 1-12. f. 520-539. Cincinnati. Jl 1913. Mackenzie, K. K. Notes on Carex—VII. Bull. Torrey Club 40: 529-554. 15 O 1913. Includes Carex Brainerdii, C. pityophila, C. geophila, C. brevicaulis, and C. microrhyncha, spp. nov. McAtee, W. L. Some local names of plants. Torreya 13: 225-236. 14 O 1913. McCurdy, H. M. On certain relations of the flora and vertebrate fauna of Gratiot County, Michigan, with an appended list of mammals and amphibians. Ann. Rep. Michigan Acad. Sci. 14: 217-225. 1912. Merrill, E. D. The botanical exploration of Amboina by the Bureau of Science, Manila. Science II. 38: 499-502. 10 O 1913. INDEX TO AMERICAN BOTANICAL LITERATURE 695 Merrill, E. D. Studies on Philippine Melastomataceae, I. Philip. Jour. Sci. 8: (Bot.) 207-250. Jl 1913. Thirty-nine new species described. Merriman, M.L. Nuclear division in Spirogyra crassa, Bot. Gaz. 56: 319-330. pl. TI, 12. 15 O 1913. Metcalf, H. The chestnut bark disease. Yearbook Dept. Agr. 1912: 361-372. pl. 34-37. 1913. Mez, C. Additamenta monographica 1913. Repert. Sp, Nov. 12: 411-421. 25S 1913. Includes new species in Nidularium (1), Aregelia (1), Tillandsia (3), Vriesea (2), Lindmania (1), Puya (1), Pitcairnia (3), Brocchinia (1), Hohenbergia (1), Aechmea (1). Mottier, D. M., & Nothnagel, M. The development and behavior of the chromosomes in the first or heterotypic mitosis of the pollen mother-cells of Allium cernuum Roth. Bull. Torrey Club 40: 555- 565. pl. 23, 24. 15 O 1913. Nichols, G. E. The vegetation of Connecticut.—II. Virgin forests. Torreya 13: 199-215. f. I-5. 25 1913. Olive, E. W. Intermingling of perennial sporophytic and gameto- phytic generations in Puccinia Podophylli, P. obtegens, and Uromy- ces Glycyrrhizae. Ann. Myc. 11: 297-311. pl. 15. Au 1913. Osterhout, W. J. V. The organization of the cell with respect to permeability. Science II. 38: 408, 409. 19 S 1913. Pammel, L. H., & King, C. M. Four new fungous diseases in Iowa. Iowa Agi. Exp. Sta. Bull. 131: 199-221. f. 1-13. Ap 1912. Picard, M. A bibliography of works on meiosis and somatic mitosis in the angiosperms. Bull. Torrey Club 40: 575-590. 15 O 1913. Praeger, W. E. Plant breeding. Ann. Rep. Michigan Acad. Sci. 14: 22-32. 1912. , Quehl, L. Beschreibung einiger Kakteenbliiten. Monats. Kakteenk. 23:.120.. 35 S 1013. Rolfe, R. A. Catasetum wists cciaa Curt. Bot. Mag. IV. 9: pl. 8514. S 1913. A plant from Peru. Rolfe, R.A. Rosa foliolosa. Curt. Bot. Mag. IV.9: I. 8513. S1913.- A plant from North America. Rolfe, R. A. Stanhopea grandiflora. Curt. Bot. Mag. IV. 9: pl. 8517. O 1913. A plant from Ecuador. Rydberg, P. A. Studies on the Rocky Mountain flora—XXVI. Bull. 696 INDEX TO AMERICAN BOTANICAL LITERATURE Torrey Club 39: 99-111. 18 Ap 1912;—XXVII. Bull. Torrey Club ote 301-328. 23 Jl 1912. Includes Dipterostemon and Hesperochloa gen. nov. and new species in Salix (1), Celtis nb penis (2), Parietaria (1), Eriogonum (3), Rg (1), Chenopodium (1), Atriplex (2), Eurotia (1), Limnia (1), Cerastium (1), Alsine (1), Arenaria (1), Ranunculus (1), Thalictrum (1), Delphinium (5), Cheirinia mr Sophia (1), Arabis (2), Parrya (1), Smelowskia (1), Deschampsia (1), and Anticlea (1). Saccardo, P A. Notae mycologicae. Ann. Myc. 11: 312-325. Au 1913. Includes Macrophoma Brenckleana and Fusicoccum dakotense spp. nov. From North Dakota Sargent, H. E. Luzula campestris, var. frigida in New Hampshire. Rhodora 15: 186. 170 1913. Sargent, C. S. Trees and shrubs; illustrations of new or little-known ligneous plants. 2: 1-56. pl. r00—125. S1907; 57-116. pl. 126-150. My 1908; 117-189. pl 151-175. Je 1911; 190-278. pl. 176-200. Au 1913. Includes descriptions of eighty-seven new species and a large number of varieties. Schenck, H. Acaciae myrmecophilae novae. Repert. Sp. Nov. 12: 360-363. ° 1 S-1913: Includes nine new species. Shull, C. A. Semipeimeability of seed coats. Bot. Gaz. 56: 169-199. FF to. 17 S613: Small, J.K. Shrubs of Florida. i-x + 1-140. New York. 451913. Small, J. K., & Carter, J. J. Flora of Lancaster County: being descriptions of the seed-plants growing naturally in Lancaster County, Pennsylvania. i-xvi + 1-336. New York. 3S 1913- Smith, G. M. The use of celloidin membranes for the demonstration of osmosis. Bot. Gaz. 56: 225-229. f. 1-3. 17S 1913. Solms-Laubach, H. Graf zu. Tietea singularis. Ein neuer fossiler Pteridinenstamm aus Brasilien. Zeits. Bot. 5: 673-700. pl. 6, 7: Au 1913. Sprague, T. A. Nautilocalyx pallidus. Curt. Bot. Mag. IV. 9: #1. 8519. O 1913. A plant from Peru. Stapf, O. Utricularia longifolia. Curt. Bot. Mag. IV. 9: pl. 8510. S 1913. Stoddard, E. M., & Moss, A.E. The chestnut bark disease. Endothia gyrosa var. parasitica (Murr.) Clint. Connecticut Agr. — Sta. Bull. 178: 5-19. f. 1-8. $1913. [Illust.] INDEX TO VOLUME 40 New names and the final members of new combinations are in bold face type. Abama vegeta 577 Abies Fraseri, ay subalpina, 684 Abietites folidet sy | Acer, 588; eatoruienias ae californicum texanum, 56; carolinianu 05; dasy- carpum, 606; raxnifolium, 5 53 esta om 5 56; Ne- nies Rivers 6063 Pseudo- Sai Cob: rubrum, 605; rubrum tridens, 392; sacchiaveninn. 606; tex- num, geanconleinas 54, 605 erates amboyensis, 571, ere Achillea Millefolium, 590 Achroanthes unifolia, 493 onan Bed americanum, 45; elatum, 46; seri Aconitum “Napellas, 585 eno- 1 392 Adenophorus bipannatus, 200; hym h es pin 198; nnatifidus, tripinnatifida, 200 bags ecidium Berberidis, 503; Rhamni, 503 egilops o 76 Aesculus seh ee OE —. 122, 127; 401, 403, 404, 406; ssioides, 122, 406, 407; macrophylla, be of the Rhinan- 401 - 34; corymbo ; 434; dalicitala 0, 437; erecta, 435, 431; fasciculata, 417, 426; fili- 8; filif caulis, 420, 438; olia, 418, 429; sobeeine: 417, 427; Harperi, 417, 426 olmi: 418, 420, 435; laxa, 41 431, 435; linifolia, 415, 420; longifolia : itima, 415, 421 ophy 432, 433; obtusifolia, 436; oligophylla, 403, 419, 432; palustris, 125, 120, 128, 422; i , 436; perennis, 420; pinetorum, 417, 424, 425; Plukenetii, 431; pulchella, 418, 428; purp 126, 416, 422; setacea, 419, 430, 431; tenella, 419, 434; tenuifolia, 420, 437; virgata, 417, 424 Al 573; lanugi- Alt | Agapanthus, 581 fee aricus campestris, Alabama, perep ese ns the Mesozoic flora o f Atleatie tale plain, 567 Alchemifla, mee Aletes, 68, 69, 71; MacDougali, 68; Monat 3 Alismacea e, 576 Allantodia scandicinum, 2 Allium, 555, 558, 578; Seoe 555; cernu- Scere 5 manita muscaria, 167; pantherina, 167 aan 464; arctophila, 464; pinqua, 464; ‘tortuosa, 4633 ventorum, 403 Ambrosia artemisiaefolia, a ao 387; bidentata ota 84; trifida, Amelanchier tucketense, 61 iy Ammiaceae, Ampelopsis cordata, 3 Am om ae nium Gautichaudii, 200; min- Amsinciia spina, 481; micrantha, 481; 1r ‘Anissbia pedis. no Eastwoodiana, 465; Fremontii, 4 texana, 465; 6 gra > alae ia, 569; Novae- =e ae Parlatorii, 569-572; Wardi- Adeopcene scoparius, 383; virginicus, 495, 49 A cen albertina, =oeg Cte 463; carinata, 462; Chama me, 462; fili- rn apr ociehtniie gest simplex, pee rtele ne oe 497, 498 Angiosperma' e, 5 iosperms, a baiieieohy of works on mei a somatic mitosis in the, 575 Anoectangium cirrhosum, 674 Anogra leptophylla, 6 Anonymos, 122; cassioides, 122, 127, 407; 697 698 | INDEX erect 435; flava, 409; pedicularia, a to) 0 Anthemis, age Cotula, 383, 387, 393 Anthericum Anthopogon Mecoun, 463; tonsum, 463; bir aeners 463 ramet Antirr i 3s 484; Kingii, 484 ypandieayin’ 465 toca oo 383 Aracea Aragalls *Bigelovii, 53; plattensis, 53 Aral Aveta. 5733 oe ae 569; spinosa, 496; Wellingtonia Arisaema triphyllum, ee 232, 234, 577 Me eee tripylium, ee development of the embryo-sa Aristida Pispeialiony oe Armillaria mellea, 167 Arum sag ost 577 Aruncus Aruncus, 495 oO ig macrosperma, 383, 390, 392; Asclepiadaceae, 466 pias se pent labriformis, ae; exicana, ovation. 3 WG: enienar 4973 syriaca, ; evagior 383, 385, 387, 389; a, 4 Ascyrum Snlas 608; multicaule, Ae officinalis Asperugo proc ccumbens we tenellus, 481 pert t bs Bi cra se eee Braunii, 203; al olium, 2 he- me sana 204; haleakalense, ; Hillebrandi, 20 Mate sic einai 208, 216; Adiantum-nigrum, 208, 218; am rigs ae ee pool rs um, 210, 216; erectum M ei, : ianum, 223; cidum, 218; fragile 209; furcatum, 217; tum. ities pseudonitidum, 217; kauai- ense, 207, 212;7Kaulfussii, 207, 212; Knudsenii, 220; leptophyllum, 210; lobulatum, 208, 211, 214, pA 212; lunulatum, 207, Bao 215): 210; Lydgatei, 208, 216, 21 ; Macraei, 208, M ‘3 215, 216; Mannii, abe "Man nii kauai- ense, 212; marginale, 223; meiotomum, ; esii, 210; wine: ; 210; monanthes, 207, 210; monta 494, 495; Nidus, 206; nitidulum, “208 lucidum, 220; pinnatifidum, 405 Poiretianum, 221; polyphyllum, 216; projectum, 209, 210; protensum, 212; pseudofalcatum, 207, 211; resectum, 209; rhipidoneuron, 2 17; rhizo- phyllum, 215; “ mb ones 207, 200; sandwichianum, 224; scandicinum, 221; , sclanoche llum, 208, ae spathulinum, 213; sphenotomum, 208, 219; stoloni- ferum, 209; strictum, 215; Tricho- manes, 207, 210, 211; peter ae wie varian S, 208, 215; vexans, 20 ride, 209 Aike kazes utahensia, 466 Aster divaricatus, 494; macrophyllus, 4 Astragalus ampullarius, 47; araneosus, ill 50; argillosus, 52; argophyllus, 49; ieti + arrec ctu seopubescens, 53; Haydenianus ma- jor, Sr Hayde nianus nevadensis, 51; ibapense, 51; ineptus, 50; inflexus, 49; jejunus, a Kelseyi, 49; lan lan antes Ss e Leibergii, 49; lentiginosus, 50; taleca i: ingulatus, 52; iser, 52+ Mulfordae, 51; multi » 48; Si- nensis, 49% 1 ygeger 49; platytropis, 50; Pre 47; pubentissimus, 48; sek a8 reventoides, 52; reventus, 52; sabulonum, 47; scobina sepia serpens, 47; sesquiflorus, 48; Sileranu 47; simplicifolius, 52; strigosus, 53: subcinereus, 47; tegetarius, 52; utahen: sis, 49; vexilliflexus, 48; Wardii, 47+ Watsonianus, Psd Zionis, 48 Atelophragma, cathe ec 50, 513 age 51; Porvoudi « glabrius- culum, 50, 51; ibapense, fe ‘lineare, 50 INDEX ge ni! i. be gpm 67; Garrettii, 68; ; Arnottii, 224; Bald- Be D deparilded 220, 221 m, 220-222; proliferum, 220, Athyrium, 20, 2 Jonesii, 70; purpureum, a R 125, te) Aureolaria, 26, 128, 401, 403, 404, 408; dispersa, 408, 411; glauca, 410; pectinata, 403, 409, 414; pectinata i 14; pedicularia, 125, 408, 400, Auricularia Fuateateaee, ee mesen- terica, 141, 164 grog oe get ampla, 165 ’ Azalea lutea, 495 yg neste 637 Balsam e, 6 Barbula agus 659; linearis, 675; Raui, 6 Bartramia macrocarpa, 658; macrotheca, 658; sphaericarpa, 659 Bauhinia cretacea, 571, 572; marylandica, 579-572 i Bellevalia romana, 579 Sie yg ia scandens, 92 Berry, E. Contributions to the Mesozoic § fines of the oes coastal cae Alabama, Bet: Bete ee. 492, 493; a a 496 wae mae of works eiosis and matic mitosis in os Peenionteery e ferns and flower- 699 161, 166, 175; indecisus, 146, 155, 160, 166; pallidus, 146, 152, 166; punctipes, 6 157, 160, 161, 175; regius, 156, 166; scaber, 154, 166; spectabilis, 160, 166; subtomentosus, 157, 160, 166; vermicu- losus, ae 152, 154,157, 166, 174; versi- pellis, I55, 157, 158, 166, 174, 180 Boltonia ieee 383, 387 bbe cig e, 479 ical cross-section of northern Miss- notes on the influence of 377 B ne, 124, 127 Brachyphyllum Thee oe formosum, 571, 573 BRAINERD, E. Four hybrids of Viola Brauneria purp Breutelia Ln ented a 58; tomentosa, 6 Britron, E.G. West Indian mosses—I, Brunnichia, 388; cirrhosa, 383, 387, 390, 392 Bryonia, 590 Bryum, 653; acuminatum, 659, 660; agrarium, 659; albidum, 653, 654; calcycinum, 661; lycopodioides, 660; nanum, 653; parasiticum, 655, 660; sphaericarpon, 659; tomentosum, 658 Bachan: II9 B ursa, 586 ButTLer, O. A note on the significance of sugar in iy tubers of Solanum tuberosum, II0 Caeoma nitens, 361-366 Caeoma nitens Burrill, The production “ . laa deg by the aecidiospores olets, 26 Bidens, aristosa, 497; sp-, 383 Calendula, 586; officinalis, 590 Bigno cigera, 392; venusta, 589 Callicostella depressa, 669 Blechnum, 225; esian 26; | Callirrhoe. oe, 57 u ange epebtian sono, 227; Callisteris arizonica, 472; attenuata, 471; Souleytianum, 225; arrosum, leucantha, 471 Biepherigie foisilgs as: 493; pera- | Callitriche, Calocera cornea, 164; viscosa, 140, 164 Calycanthus floridus, 585 Calycites, 573 Bois vee salicina, 62 Boleti, a in the cytology of the ymen H mycetes, es Boletus albeilus, 146, I54-T: 56, 159, 160, 166, 174, 180, 181; alutarius, 156, 166; badius, 156, 160, 166; bicolor, 160, 166;|C castaneus, 146, staat =. 160, 166, 174, 179, 180; 146, 156, 157, 160, I6I, 174, ros sere pene 157, 160, 166; flavus, 165; glabel- lus, 146, 152, 157, 160, 166, 174; us, 146, 151-156, 160, I61, 166, 174, i, 180; griseus, 157, 160, ymperes, 660; parasitica, 660; Rich- i, 660 Camarophyilus virgineus, 167 ampanula Camptosorus rhizophyllus, 641, 642, 644, Canna indica, oa Cannabis sativa Cantharellus Aharon 167; cinereus, 142, 167; infundibuliformis, 167; tubae- formis, 167 700 Capnoides semier virehs- bags Capnorea incana, 4 bape peer 479; Watsonia Capparites, 573 Capsella, 586 oeirani Sg orp 495 Carduus, 383; nicus, 4 496 Carex abdita, von 533, 5493 acuta, 5773 534 etexa, 541; No- net 538; pilulifera ophila P gu mbellata, 549-552; umbellata brevirostris, 5 531; umbellata vicina, 551; varia, oe 539; varia omepe 38 Carex um none its allies, ad Carpin coraiiae a, 387, 3 Gatnaliciie 5733 fisribundas, pe 571 m, 67; Gairdneri, 67; Garrettii, 68; m, 6 Case of bud variation in Pelargonium, A, 307 Cassia, 573; nictitans, 4 Castanea dentata, 3 Castile hispida, 485; babip cash 484 Casu Celastracea Celas inet 28 BTSs laminae cTt. carolinense, 572; » 569, 572; decurrens, 571; slicheosrase 571, 5723 Newberryanum, 571; undulatum, 569, 2 astrus, 449; scandens, 605 Celtis, 383, 392 Centratherum chine Cephalanthus Esceatety 383, 386, 387, 90, 392 Ceratodon,99; purpureus, 98-100, 106, 109 Ceratodon purpureus, Development of the peristome in, 97 rma s- INDEX Ceratophyllum submersum, 585 a oa 449; canadensis, 383, 385, 387, Clesckinn 383 Chamaec rista fasciculata, 3 se Scomtectge coy ‘ ta, 3 "Aan: 53 CH “pga ward Lyman go eke a Aas maphila i Rg 493 oecteenen juncea, 590 Chrysanthae, 57 Chrysopsis Mariana , 496 Chrysosplenium americanum, 493 rea tnnataities ma, 66 Chytra, 123 Cibotiu mp rolife Cicu a Cartiball, 383, 38 Be, 387, 390 ibanciaed intermedium, 569; New- Cissites ne ot 570 Cistaceae, 61 Citrophstium aligerum, 571, 572 Citr Cladophiebis, par IE coal aed 306; ee Clastobryum americanum Rae paige planulum, 671; a Bad ting pris 672, 67 Clavaria aig 165; rugosa, 142, 165 ver micu lari 573; alabamensis, 569}; ytoni elon red 584 Clete tia | recta, 584 ae tocybe saath: 167; aurantiaca, x4 Clitopis orcella, 167 Closte: ee (3 Cc peasita argillosus, 52; conferti flo oo 52; reventoides, 533 vaveaiticd. Eceeen scandens, 589 na tc 57: Oona te 572 Coelastra Cogewellia toptopsyils, 74; platycarpa, 74; robustior, 74; simplex, 74; triternata, Coleosporium Senecionis, 502, 509 Collomia aristella, 475, 476; inconspicua, 476; linearis subulata, 475, 4793 tinctoria, 475, 476; tinctoria subulata, ta, 161; radicata, 144; Collybia yet re I, 167; velutipes, 146, 149, ommelinaceae, 577 Coniophora cerebella, 146, 150, 165, 174 INDEX Conocarpites, 573 Conoclinium coelestinum, 3 Conradia, 124, 128; paper aN 124, 128, 405; Lecontei, 405 Contributions to the Mesozoic flora of the Atlantic coastal plain—IX. Ala- bama, 56 Convallari Convallariaceae, 582 Convolvulaceae, 466 ‘Conyza chinensis, 306; odorata, 306; patula, 306 Cocdaut ephemerus, 143, I5I, ee re se 161; radiatus, 141, rarius, 137, ye. tuberosus, et Conta apiculata , 569 Coryd a, 585 cies olanchan! A843 bicolor, 484; ramosa, 484; Wrightii gag or 60; Bethe, x Jonesii, 70; Ss, 70; Rosei Pee dos ilu cin shone Cornus canadensis, 492; abies 385, 387, 90; stolonifera, 498 Corticium alutaceum, 138; amorphum, 139; uscatum, 138; lacteum, 165; lilacino-fuscum, 143, 165; roseo-pallens, 38, 148; subgiganteum, nae vagum, ; racca virginiana, 387, 390 Crataegomespilus Asnieresii, 371 Craterellus cornucopioides, 165; sinuosus, 140, 16 petty teers 143, 167 Crepis, Cressa 3 depressa, 466; erecta, 466 Crinum, 449 Cristaria coccinea, 58 Crocanthemum canadense, 613-615; du- mosum, 613-615; georgianum, 616; Bi aoe. a Ponts opinquum, 615, 616 roton capita intact panduraeformis, 571 Cryphaea filiform 57 ‘Cryptanthe Sitges 481; calycosa, 481; flaccida, 481; osa, 481; ct Cucurbita Pepo, 590 Culture el sears! rusts in the greenhouse, e, Cunila ec caatiee 406 Cuscuta curta, 466; Gronovii curta, 466 Cuscutaceae, 4! Cyanopis, aes pubescens, 306; villosa, 06 gf luowepe pe 306; chinense, 306; pubes- m, 306 cya hire hirsutus, 170 cadin us circularis, 569, 571 idee teen» eihdndine: 663 701 pedis carn 69, ee. anisatus, 71; bipin- us, de Jonesii, 70; nivalis, 70, 73; ouren reus, 70 ynoglo onic officinale, os Serta cs mn tobem, 73; Nuttallii, 72) 1375 SB Rae fs ere 659; strictum, 6 Cyperaceae Cyperus a 383, 387, 393 Cyphella rng — digitalis, 165 Cypripedium a e, 4933 — Coles aihins, 204%, Boydia e€, 204; tideum, 204, 205 Cystium, 51; araneosum, 50; boiseanum, 50; Coulteri, 50; ineptum, 50; lenti- ginosum, 50; platytropis, 50 Cystopus, 501 Czekanowskia capillaris, 569 €, 493 caryo- Dacryomyces Ce aap a ss I41, 164 Dactyloc Hee Daedalea unico, 138 Dalea parviflora, 51 Dammara borealis Gyr, See , 588 IAI; 123; 326, 411; flava, 4 ar0; siege 4 — queria, 410; luercifolin inter- Dascophyilum, 68, 72; lineare, 69; tenui- Daucus us pusilg a 387, 380, 393 apes us, 609 Dentaria heterophyl, 496 De ei, 221; triangularis, 221 | Dermatophyllites acutus, 571 , = some in t the first or heterotypic mitosis of the pollen mother cells of Allium cernuum Roth, The, 555 asthe of no embryo-sac of Ari- a triphyllum, The, 2 pevekopmiont Of a sestitnaie in Cera- todon purpureus, 97 Dewalquea groenlandica, 571, 573 Dickson Nien a: 571, 572; pro- lifera, 2 Dieranum aspleniides, 662; calycinum, 660; palmatum, 569; Smithi, Dictamnus albus, 587 702 Dictyolus bryophilus, 165; oe 165 poe fase aah — 143, 87 Didymoglos: Dieffenbachia ‘Sekine: 232 Diholcos scobinat : Diodia tere (a) Dioscorea villosa, 390 Diospyros amboyensis, 571; primaeva, se HUT. Ags Seder tacts 571; vir- ana, 388 , 496 Diplaziom, soy say ease 223; Fenzli- 223; marginale, eure ense, 223, 224; plantagini- folium, 222; sandwichianum, 223, 224 a eae rufescens, 659; tortile, 100 Dodec n Jaffrey eae Meadia, 497 Dosdia, ahs asper % Kunthiana, 228; Kunthiana Paint 25 228; media, 228 Doronicum cise eg 590 Dracopis amplexicaulis, 393 Drosera, 442, Drynaria _cloneata, 201; nuda, 201; spectr Drvopteris, 203; caryotidea, 204; niifolia, 203; cyatheoides, 204; leuco- chacte, 184; lurida, 183, 185; pubes- 183 Diycpreices Stephensoni, 572 Eatonia nitida, 494 Eddya hispidissima, 481 Edosmia er eri, 67 Edward Lyman Siurite 599 Elatinace: vee 612 latine americana, 612 Elatostema, 58 ELLIs, ANDREWS, F. M., &. m: observations concerning the deg of the leaf hairs of Salvinia Perce foliosa, 478';. penduliflora, 479; salina, 479; scopulina, 479 Encalypta asitica, 660 Endophyllum, hc. 366; Sempervivi, 361, Endothia parasitica, 488 Entodonaceae, 670 mes adium, 63; Drum- m, 63; laevicaule, 63, 6 tiusc lum, 63; minutum, 63; Palmeri, : aniculatum, 63, : ner ore ebay tees Ss Sand emocarya muricata, 481 Eremosis, 306; ovata, 331, 332; Palmeri, FE ita, 465, 578 - ord St INDEX 332; Steetzii, 332; tarchonanthifolia, 2 eae 394 rigeron ramosus, 386, 387 Sota IIQ, 123 Eucalyptus Sie a, 569; Geinitzi, 572; os st lia, 572; nervosa, 569; parvifolia, eos ccity 573 Eulophus, 67 Eu upatoria ci Gaur 306 ie evar aromaticum, 496; coelesti- m, 496; m menthaefolium, 331; rotundi- eae, orollata, 390; Cyparissia S,.502* colioclare 53; Par- ryi, 53; Sp. 3 9 Euphorbia cea hra Euphrasia mollis, 485 ns, A. W., & Hooker, H. D. JR. Developmen the seiistaaae in Ceratodon pares 97 Exidia truncata, I41, Exobasidium pie Rare 164 Fabaceae, 43 Fago pyrum stg tum, 584 Fagus, 394 andifolia, 386, 387 Ferns arid " aivet ring plants of Nan- tucket—XI., The, 605 Ficus, 573; crassipes, 572, 573; daphno- genoides, 569, 571-573; inaequalis, 569, 571, 572; Krausiana, 571-5733 lanceolato-acuminata, 569; Woolsoni, 569, 571, 572 bag oats ‘asplenioides, 662; Barbae- m , 662; costaricensis, 662; pal- s, 661; polypodioides, 661 Fistulina hepatica. lamaria, 127, I 405 Fontinalis, 653, a Ag crispa, "6h: disticha, filiformis, 657; Fouquieria, 16; splendens, Four hybrids of Viola sr Bor 249 hese prety 587 Fr; gee a, 4933 quadrangulata, 497; D. The culture of pans tae in the greenhouse, oF: Funaria hygrometrica, 97, - ia, 579 Galanthus nivalis, 5 ~~ tenera, I4I, oe 167; tenuissima, Gattoutla candicans, 581 INDEX 703 Gaultheria array: ag 493 uxii, Gaura Micha Gayop ibcact acum, 65; Helleri, 65; lasios m, 65; racemosum, 65 Geaster Smbriatas, 170 Geinitzia formosa, 571, 572 Gentiana Arebobiba denistiora: A463; caly- sa, ; calycosa monticola, 464; calycosa stricta, 464; crinita, 49 det , 463; glauca, 464 acounii, a 463; oregana, 464; procera, 580; tortu- osa, 463; ventricosa, 463 Gentianaceae, 46 Gerardia, 119-122, 125, 129, 408; aphylla, As3* aphylla oe 438; aphylla g iculat randiflora, 433 a, 126, 128; cassioides, 407; nettle. Lay; - loba, 434; dispersa, ; divaricata, -* I27; georgiana, 427; glauca, 409; glutinosa, 120; Holmiana, 429; lini- folia, 420; maritima, oie maritima grandiflora, 421; maritima major, 421; Mettaueri, 438; Mettaueri clausa, 438; Mettaueri nuda, 438; microphylla, 432; nuda, 438; parvifolia, 436; pectinata, 414; pedicularia, 120, 121, 125, 128, 412; pedicularia pectinata, 414, 415; Ptukeneti, 431; Plukenetii micro- , 432; purpurea, 120-122, 125, ? assifolia t mulosa, 424 pig. To Fs brosa, med ana shi east 436; setacea, pete aay 403, Gilia ag ‘eileen aa aggregata dgesii, 472; arenaria, ; arenaria rub 25 tella, 75» 476; ei eae gf a ; arizonica, 472; attenuata, 471, 472; Burleyana, 470; caespitosa, 473-475; can 4 7 00 472; iberidifolia, 468-471; ‘thientiuen 73; longi » 471; se hg 469, 470; ultiflora, 471, 473; n 479; palmi- frons, 470; pulche! Ha, se apd) pungens, 473, 474; pungens squarrosa, 474; ri Syed ula, 475; rosea 470; 472; sinister, 476; sinuata, 472, poet spergulifolia, 469-471; spicata, 469, 471; = si oe 468, 470; stra- mine ubnuda, 473, 475; 'te _ tuba pte ‘iigde, 469-471; Tweedy Gladiolus, 583 ig ert H. A. Studies on = West ere Vernonieae, with new s from Mexico, 30 Gleditsia triacanthos, 383, 388, 392 eats were la, 571 tie mbigua, 67; Bolanderi, 67; ay occidentalis, 67 Gnidia er Gomphidius glutinosus, 167 Gortner, R. A., & Harris, J. A. On “ possible relationship between the ructural p ities of normal and tau falaat fruite of Passiflora gracilis and some phys Parca properties of their cxpreneed Fi ay — 588; he tows 65: 0,13; ete 576" yaaa 576 Grammitis setigera, 194; tenella, Gratiola ce oat E27: eR le 484; 127 a: Observations on the ried composition al the Sugar eave flora, 487 Gruvelia, 479; pusilla, 479; setosa, 470 Guepinia helvelloides, Rage od rufa, 164 Gymnogramme sadleri Haematoxylon, 6 sya campechianum, 657 Hamosa ig HARPER, ay M. A botanical cross-sec- tion of northern Mississippi, with notes poet the influence of soil on vegetation, stan J. A. On the relationship be- tween the number of ovul e a ARRIS, J. A., &. On a _ossible “relations between the of normal and teratologica AE of P Passiflora gracilis ventas omg properties s their on Heleniam tes elias ped 486. 387 um canade ense, 613; majus, 614; propinquum, 615 Helianthus Perigiat I Helicodontium capillare, 668, 674 mapearas | Scetitinn 584 Hemerocallis fulva, 577 Hesperis matronalis, 585 Heterocodon rariflorum, 485 704 Hibiscus Moscheutos, 608; oculiroseus, Hicoria alba, 383, 390; ovata, 383, 392 Hieracium, 590; paniculatum, 494; veno- sum, 494 Holomitrium pera oa 661 oe glabella, 664 alobus, py SI, 52} aboriginum, hh 53; Dodge: 673; con- ett ; Carionis, 668; eoides re orbit distichon, 576 Hosackia, 45; elata, = Purshiana, 45 Houstoxts coerulea + lab ge so a Hoy . D. Some toxic and antitoxic piel in cultures of Spirogyra, 333 Hudsonia tomentosa, 618; ericoides, 617, 618 Pienig ate ses ee 579 Hydnan » 141, 142, 169 Rediuwe Tae pee fepuntiing: i 140, 165 Hydrangea, sort arborescens, 390, 496; Ie create drodictyon, ss 80, 83-85 Suan eae, 4 INDEX 671; nigrescens, 666; pallidum, 663; palmatum, 661; patulum, 664; poly- podioides, 661; Dolytrichotdes, 668; pseudo-reptans, pungens, 672} reptans, 665; spnaetorme, 654; ama- risci, 665, 666; 675; tetra- onum, eS 560: tor g 673; serdar quatum, 674; trichophyllum, 669 Hypochnus Sambuci, 164; subtilis, 143, 164 Hypomyces rosellus, Hypopitys lanulosa, bod latisquama, 461 ypopterygium brasiliense, 666; pseudo- tamarisci, 666; Tamarisci, 665 Iberis, 586 Ilex fastigiata, 613; Masoni, 571, opaca, 393, 394, 495 Illiamna, 60; acerifolia, 60; angulata, 60; 572; rivularis, 60 Impatiens niet Index to Amer aes +37; 4 24. 591, 647; 69 Iniuence a starch, po Aa and sugars on the toxicity of various nitrates to Monilia sitophila (Mont.) Sacc., The, apace literature, 3» 205, 373) 457, 523, In an cretacea, 571 bye cybe asterophora, 144; trechispora, Iris, 583; cristata, 496 Is cit a factor in the distribution of Nereocystis Leutkeana? 237 neA. 306; ovata, 301 pees odes Watson, 478; occidentale | Isopterygi tenerum, = Isopyrum biternatum Natl sephe fla te 168 gigi es _Bonplandi "bartinee. 6733 Hygrophorus cheer gare mus, 168; ceraceus, tetragonum, 673 2, 168; conicus, 142, 143, 168, 171; | Isotria vetticiliate, 493 lucorum, 168 tea virginica, 390 Hymenaea, 573 Hypericaceae, 608 Juglans arctica, de 572; nigra, 389 Hypericum adpressum, 609; adpreecim Juncoides saltuen spongiosum, 609; boreale, 609, 611;| Juncu saletdiaven: pes effusus, 387, 390 canadense, 609-611; dissim ; 610, Fenverinicat tes, ; me mmondii, 497; majus, 610, Juniperus virginiana, 383, 394 611 eee 610, 611; perforatum, 600; sick Brittoniana, 571; rai oa 494 gam aouemeniabiad 168; fasci- Kentrophyta wes etaria culare, 168; rplexum, 143, 168;| KNup eek on the cubticcida. 168 etica: : son, and duration a Hypnum, 653, 673; albens, 663; albicans, cambium eS ee in the Ameri 663; as eB 662; Bonplandi, 673; s laricina (DuRoi) Koch}, caespitosum, 671; atum, 671 27 are, 668; cirrhosum, 674; com-| Koellia flexuosa, 387, 390; incana, 496 positum, 667; congestum, 655, 672;| KuNKEL, O. The influence of starch, cuspidatum, 654; crispum, 656; den-| peptone, and gars the toxicity sum, 667; depr ressum, 669; diaphanum, of ous nitrates to Monilia sitophila 663; fasciculatum, 665; flexile, 666;| (Mont.) Sacc., 625; The production fulgens, ri glabellum, 664; jamai-| of a promycelium by the aecidiospores cense, 656; loxense, 671; microphyltum, | of Caeoma ni Burrill, 361 INDEX Laburnum, 586 Lactarius delicious, 168; piperatus, 168 ie e, 48 m amplexicaule, 481 pee erecta, Larix laricina ee “Roi Koch, tg! tions on the inception, season, ile eid of cambium, development in merican larch Latha —— is, 53% Nuttallii, 53; obovatus, 53; odor (ae PGE SF Soa 573; nervil- axsag m, 571 Laurus plu tonia, a 572 § eadharboiate lla, Lavatera, 588 Leaf water and stomatal movement in iz ° jor mh Lechea 618 terior, 8 5 Leggett Peps 620; mari- a i LS: Se monliformis sa, sa, 496 620; villosa, 6 mphalobot des,. vase na, ig t eseide’ 166 Lepidium i. 383 phanum, Lepidopilum dia 663; poly- Sicheles , 668 Lepiota cepaestipes, 168; lilacin no-granu- pe i mucida, 140, 168; naucina, pipiens PEE yak aor 474; eu pungens, 473; 473" Hooker, 474; pungens, pasa ai caespitosum, ap caagg 8 — um, 659; mexi- 659; pseudo-rufescens, 659; ee oe trichophyllus, 669; Lepyrodon peboeae ee) has io Leskea albicans, 663; Bonplandi, 673; capillaris, 668; caespitosa, 671; con- Dee 672; depressa, 669; flexilis, 666; gla , 664; involvens, ou: pungens, 672; re motifolia , 664; Tamariscina, 665 Leucodon ~ eichonee nae prea Leucojum, 582 LEVINE, M. Stu in the cytology of the Prato oc 28 especially the Boleti, Liliaceae, 577 Liliales » 577 Lilium, 559, 580; Martagon, 557 Limodorum abortivum, 583 Linum intercursum ae Liquidambar, 381, 304; Styraciflua, 383, 572; 9 lia, 569, 571, constricta, 571; simplex, 569, 571 705 Listera, 583 ppc ee arvense Le , 481 is Se a af water and stomatal geet in Gossypium and a method of ark visual observation of stomata ins Loasa ine Lobelia eiadion 496, 497; puberula, Lobrlincene, 85 matium platycarpum, pe 45; americanus, 7 45; Macbridei, 45; sericeus, 45; tenuifolius, 45; tenuis » 45 us, 445 argen nteus argo- 44; a 433 ome a; lupinus, 44; micen 44; 44; nootka- ten fi i oreo pila "443 plumosus, 44; rivularis, 44; "Siler ri, 43 Lycoperdon caelatum, 170; excipuliforme, £795 gemmatum, 170; pyriforme, 143, peopndites ry: Lycopodium, 489, 491; sppaeaeage 489, pres complan: , 489, 493; creta- Lysimachia pe a 493 , MACKENzIE, K. K., Notes on Carex— Macranthera, Lid, 123,...124, Li 137; 401, 403-405; flammea, 4055 fuchsioides, 124, 126-128, 40 et fuchsi- oides Lecontei, 405; Lecontei, 126, 128, Macrocystis pyrifera, 239 Macromitrium, 660; cirrhosum, 6 Mex nolia, 304, 585; arian lag 569; auri- 569; Bou 569, 571 Capellinii, 572: scary 395% : prt 569; Hollicki, 571; Lacoeana, 571; longifolia, 569; abn S7t, 5725 Newberryi, 569, 571; obtusata, 571; 7 speciosa, 569, 571, 572 Malapoenna, 573; cretacea, 572; falci- folia, 571 — (4 6 Pagan 58; Creeana, 58; ta, 608 vularis, 60; rotundi- folia, 608; sia ag Malvaceae, 57, 608 alvas Pare Y 54 aber muetang 57; coc- cineum, 57, 58; ¢ m dissectum, 58; coccineum ica coccineu grossu mesg lium, or > eicecnn, 59; digitatum, 59; dissectum, 59; dissectum Cockerl, 593 pas 58; exile, 57; Fre tii, 57; cesulariaefolium, i 58; ark ed an Munroanum, 5 706 58; Aergzemt mpeess 57; spicatum, 57; Wrightii Marasmius a tamales 168 Marattia cretacea, 571 ~~ tricar ia, —_ ardon ace phocooeaey 453 hispida, 45 Meibomia laevigata, 390 ; Michauxii, 390 Melampsora Lini, Melampyrum lineare, 493 M drium rubrum, 584 Melanthaceae, 577 Me s 383 iflora humilis, 61; multi- flora integra, 61; Rusbyi, 61 Sere ait ame annua, 587 Mertensia alpina, eye coriacea, 481; brachycalyx, 481 neeolata, 481; longiflora, 481; mutans ape per rplexa, 481; pulchella, 4 erulius fugax 166; lacrymans, 139, 166; Penciians, I59, 166 4 rsifolium, 664 ureo- nitens, 664; rpc 0 664; oreo’ quense, 665; te a ‘ibs flaccidum, 664; flexile, 666; escens, a: i 0; cettaeieh t 66 subam- sericeum, 670 AS biguum, 665; tenue, ba: chiens, 6733 ; torticuspis, ie M seme cranthes diffusa, 475 Microlotus, 45 Micromeria Douglasii, 481 Microphacos parviflorus, 51 Microsteris gracilis, 47 M — = eB pas 665; rep- 8; 665: T ponents 665 bili, 478, ; foliosa, 478; salina, 479; §' #0 imulus caval nalis, 483, 484; East- woodi Mirabili s, 584 Mississippi, A botanical cross-section of northern h neat on the influence wi of soil es vegetation, 377 so vibe 390 Mittenothamni a strictum, 659; tomentosum, 658 da iodor: , — 482; Nuttallii, 482; pectin sperma rotundifolia, ae: isn 625- Monnioty leone ha Monotropaceae, 4 Morus indica, wae rubra, 383, 380, 392 Morttier, D. M., & ‘AGEL, M. INDEX The development and behavior of the hromosomes in the first or heterotypic mitosis of the pollen mother-cells Allium cernuum Roth, 555 Muciporus corticola, 166 Musa soba Musenium, 68; rr papheter 68, 60 Mycena galericulata, 155, 168 i aew. 573; cerifera, 394; emarginata, STIASTS Myrsine borealis, 569, 571, 573; Gaudini, 571, Myzorrhiza pinetorum, 485 ae oi ag Tr “i ferns and flowering plants of—XI, 60 — ng 496, 497 S, 583 Saale pre ™ oe 666; crassa, composita, 667; 667; icha, 656; ina, 656; ental ig pyre glabella, 664; hypnoidea 657; jamai- cense, 656; nigrescens, 666; poly- trichoides, 668; quinquefaria, 673; tetragona, rsd torta, 674i trichophylla, 69; turge 666; ulata, 65 Neckiteea sacs "saci: 656;‘undulata, 656 Negundo, ee seb 34-56; fraxini- folium » 553 interi 5 56; mexican 55; Nuttallii, 54, 55; cts aenes, 54, 553 ema $5: 56 acladu ede eng og Nescicoesie: 4 206; Nid Nephrodium apiftium, pomp geet 183 a, 237-241 ereocystis auch coin. Is salinity a factor in the distribution of, 237 ropical ew ferns America—II, 183; —III, ace globosa, 170; pisiformis, 143, 156, I Niphebol us linearis, 201 Nitella, 341, 342, 346 Note on ad renaperss of egg in the tu of Solanu ar Notes et Carex: EOvIL. Nothoscordum Ris ao N uphar ingrrearr oe Nuttallia acuminata humilis, 61; integr: ee 61; laevis, ple lobata, 63; arg ees 6x; p erosperma, byi Nyctalia asterophora, 168; I4I, para- Nyssa biflora, 304; Snowiana, 571; syk vatica, 392; uniflora, 3 INDEX Observations on the ee compo- sition of mn the miteokite, season, ration of cambium development Sr Acree pote (Larix laricina pads Koch), 2 Pixies. cage albidum 653, 654 ther: al yssoides villosa, 66; Oeno 588; reer hirsutissi ma, 66; densiflora, 62; hirsutissima, 66; Hookeri, ef! nay a 65; leptophylla, sk longissima, 65; acrosceles, 65; ornata, 66: valida pi i sale Se cufleting: 62; subulife ra 66; tenui et a Soinite. a relationship between the ructural pe ities of normal and tieaboloeilal | er of Passiflora gracilis = reap eters properties ed juices, 2 mber nd the aoauity of ry for maturing its ovules into eae: ia Onagra, 588; Oakesiana, 66; ornata, 66; strigosa subulata, 66 eae 2 » 480; multicaulis, Palmer, yo: pustulosa, 480 cog )reoxis, ny 70, 71, 73; MacDougali, 68 Jrobanchacea Irthocarpus “rte idus, ne Sbarro’ eapet telvencia \rthothecium, 670; tr ichophylum, 670 chairing polytrichoides, 66 iva, 576 idee i termedia, 66 mite: 127, 128 Wostima, 126, 128; petiolata, 126, 128 Ixydendrum arboreum, 491, 495 \xytropis Lambertii, 53; Lambertii Big- elovii, 53; plattensis, 53 eooeeoogsssss Paeonia spectabilis, 585 Pagesia, 127; leucantha, 127 cea laurinea, 571 ‘Palamocladium Bonplandi, 655, 673; leskeoides, 672, 673; trichophyllum, 670; in raga re subtile, 670 Panax cretacea, 5 Panctenis, 125, 728, 403, 408; pectinata, 412 161 Pani uti, 494; elongata, 493; iia th am 390° se es 496 . 287 707 Papillaria nigrescens, 666; nigrescens Donnellii, wed ae aoa ifolia, 579 arnass: wearer 493 ial rae amoena, 47; Fremontii, 47; polydenia, arthenium arg Parthenocissus ante 607; quinque- olia, Paspalum, 383 Passiflora oe 588; gracilis, 27-34; lutea, 495, 4 Paxillus prreeeak: 168 ectoc , 479; miser, 481; penicillata, 481; setosa, 479 rum, 84 Pedicularioides, 408 Pedicularis centranthera: 485; flammea, 485; Jana 485; lanceolata, 498; Oederi, 485; sylvestris, 589 Pelargonium, 367-372; Madam Salleroi, 368-371; eee 36 ogee ium, A case of bud-variation in, Peltandra undulata, 577 Pelve Peniophora bene 161; quercina, $65 PENNEL Studies in the Agalina a a bere of the Rhisekthetiee. 119, Pentstemon, — 7 acuminatus, 482; al- bertin uni- lateralis, 482, 483 mium Soraueri, 509; Strobi, 502, 3. pn Permolles, 315 Persea valida, 572 fia, 5733 Lesquereuxii, 571 cedanum simplex, 74; triternatum Teptophy lum, 74; triternatum platy- Phaca a mpullaria, 4 artemisiarum, tities S32; Cusickii, ft debilis, 5s: 48; le ptalea, ollissim: acelia Pema 4793 . isolok phe crenu- missa, 479; dubia, 495; lata aecabalincis: yrs v scehpamatotlas 4793 708 hispida, 479; humilis, A783 integrifolia, 479; luteopurpurea, 4 orbicularis, 479; Palmeri, 479; pe § 479; illa, amosissim 79 Phacopsis scaphoides, hallus impudicus, 169 Phascum, 65 um, 653 Phaseolites formus, 571, 572 Phaseolus, a vulgaris, 449, 454 Phellopteru INDEX 385, 386-390; palustris, 3094; ath 572; rigida, 291, 494; pes 22% 272; Taeda, 381, 382, 387; virgini- i Piperites, eella icine, 656 Pisum eeswitens arizonicus, 481 Planera aquatica, 392 Plantago aristata, 383, 387, 389 atior, 572; yl Pea Philibertlla cynanchoides, 466; hetero- | Platanus, 573; | a, 466 383, 385-389, Philonotis sagged tee 659 Pleopeltis linearis, 201; n Pte 201; spec- Phippsia or ida, trum, 202; Thunbergiana, Phlebia Pleuropus, 655; congestus, et 672; Phlebo diu leskeoides, 673 Phlox Nr Bo oe albomarginata, 467; | Pluteus cervinus, 169 alyssifolia, 467; alyssoides, 467, 468;| Podophyllum peltatum, 5 omon » 468; montana | Podozamites, 568; margina us, 573 prostrata, 468; bryoides, 467; caespi-| Pogonatum convolutum, 657; tortile, t 467, 468; collina, 467, 468; con- 657; urnigerum, 654 densata, 467; costata, 467; Covillei,| Polemonium californicum, 477; coeru- 467; dasyphylla, 468; densa, 468; eum, 477; columbianum, 477; deli- diapensioides, 467; Douglasii, 468; catum, 476, ; Grayanum, 478; glabrata, 467, 468; Hoodii, 467; Hoodii Haydenii, 477; intermedium, 478; glabrata, 468; Hookeri, 474; Kelseyi, mellitum, 473; xi m, ; mon- 467; multiflora, 7 8; muscoides, trosense, 477; occidentale intermedium, 466, 467; Stansburyi, 468; stolonifera, 478; p ium, 476, ; pil —_ 4953 hector saa 467, 468; viridis, 468 perms lucifera, 168; praecox, 143-140, Rosae- 169, 173 Phragmidium obtusum, 509; 509 ogonium aureum, 662; fulgens, 662, 663; globitheca, 662; immersum, 663; tra, 663; viride, 662, 6 elongata, 201; Spec- ihe cag 201; trum, 201, 202 Phymowa 60; acerifo 60; lia, 60; Crandallii, andifior ora, 60; longisepala, 61; ulaeie. Physostegia sp., Poe erene | gic on the Mountain region—I. Rocky Alpine region, 67 oe M. A bibliography of works on osis —_ somatic mitosis in the angi perms, 575 Picea Engelmannii, 684 Pickett, F. L. The phlei cid of the e embryo-sac of Arisaema triphyllum istance of the fearkell ia of osorus rhizophyllus to desic- gee Pilotrichella eroso-mucronata, 666; flex- ilis, 666; recurvo-mucronata, 666; na ae , 667; compositum, ay m, gee hypnoides, 657 Polyporus acanthoides, rrimum, 477; scopulinum 478; Tevisii, ary; tricolor, 476; viscosum, 477, 478 Polemoniacea eg Polygonum arifo um, 493 Polypodium sence, 194, 200; AGG: rum, 195; H oe pas [os fleri, 198, ieende aa leon, Hookeri, 193, 1943 hymenophylloides, 94, 199; gawd fussii, 196; 193, 195, I 201; rae tos rod pecutatonl lata, 197; sessilifolium, 55 ih ee 194; Spectrum, 201; aici a iscinum 194, 200 dum, 200; unisorum, gare, 193, 198; zosteraeforme, = adust 146, 150, 166, 174; annos 146, 150, 66” 4, 178; 159, 166; destructor, 146, Pinnulaceae, 461 Pinus saan 684, 685; echinata, 382, 150, 166, 174, 179; fumosus, 1383 INDEX lucidus, te 166; versicolor, 140, 146, 150, 167, Rotpaichais: 202, 203; 390; ari ristatum, ‘ah: vances, coniifolium, 203; decoratu 689; fa — m, 2053 haleakalense, Sox: Hille- bra 204; ilicifolium, 689; machaerophitum, 688-690; triangu- lum ge gos p od 203; Polystcts conchifer, By versicolor, 138 Polytaenia Nuttallii, Paleiiebies. 6533 se weet 657; crispulum, 657; cubense, 657; doit inin- gense, ; glaucinum, 657, 658; Reasiteieector 657; laxifolium, 658; b curo-v 56s decgrrncs 658; Sintenis teen: ‘tortile, 657, Populites, 573 pera apiculata, 569; deltoides, 383- 2, 394; heterophylla, 383, 490; Lowarnien: 572; tremuloides, le 143, 167; incrus ; rotrichu ciculatum, 66 Hoyt ieee stipulatus, 496, 407 Po ee 686; foliosus, a Potentilla, 58 Prim . 580; americana, 462; inosa, a ee neana, 462; specuicole, Hoty 462 ater densus, 667 Produc of gv mycelium by seidopore of Caeoma nitens La Bog ok = Ronee ghd 569 Protodamm 571 Protophiyllocladus subintegifotiue, ci I sil ; is Psathyra spiaidico Saeiaae I4I, 169 habe consimilis, 161; crenata, 169; disseminata, 169; gracilis, 161 Pseudocymopterus, 70; aletifotius 70-72; 0-73; it, Pseudopteryxia, EN aletifolia, 723 detente: 71, 72; » 72 Pseudoreoxis, 733 "Siplihaten, 73; nivalis, juncea, eolata, 46; Psoralea 47; icrant lat ; Stenophylla, 46; stenostachys, a Pteridium reabergint Pteridophytes of the Hawaiian ide III, A taxonomic study of the, 193 erigynandru rum aureum, 673; fulgens, aca quadrifarium, 673 erobryum filcinura, 656 Pterogonium fulgens, 662; nigrescens, es carolinensis, 571, 5723 560 modestus 3 e| Resistance of the p 709 ppt orga albicans, 663; diaph- um, 663 Peeryxi a, 70 Puccinia Acetosae, 5009; = at ie 508; pvr filicinae, 509; Chrysanthemi, 509; coronata, we 505- ors oro- 504, 506, 0-519; cispersa, 8, 518; Vicaainenrans 509; 09; igo-vera, ee simplex, 504; Sorgh 505. 506, 5 Pasigenielle pungens, 672 Pyrola elliptica, ey rotundifolia, 493 Quamasia hyacinthina, 497 Quercus alba, 383-388, 390; coccinea, 390; falcata, 383, pasion 390, 304; lyrata, 383, 389, 392; marylandica, ni 385, 388, 390, 394, 395; Michauxii, 387- 389, 302; satu 496; nigra, ais, We, 2; pagodaefolia, 383 iS stellata, 383, 3 300, 394; texana, 392; velutina, 389 Radulum tomentosum, 138 Ranunculus, 686 Ratibida pinnata, 383 rothallia of Campto- sorus rhizophy thes to desiccation, 641 , 669 Rha hooves aaa sum Rhamnaceae, 57 Rhamnus becataetelis: 57; tenax, 571 scape paigcany. enatts caespitosum, 671 Rheum undulatum, 58. Rhexia eae a, 38 7 Rhinanthaceae, Studies in the Agalina- £ the, nae, a subtribe o II9 Rhinanthus virginicus, 125, 409 a rte gn e one 654 Rhodod maxim my 494 77 IGG, Is salinity a factor in one? distribution of Nereocystis Luetkeana Robinia, 449; phere vite 385, 387, 3890 OBINSON, W. J., A taxonomic study 0 of the Pteridophyta pra the Hawaiian Islands—III, ange Rosa, 587; carolina Rubus, 587; Sree eas 361, 362; odo- ratus, 494 sie angen ee 387, 389 Rue apes re californica, 56; californica exana, 54; Kingii, 56; mexicana, 56; Naitanii 54, 56; texana, 56 710 x Sp., 383; hy eigen 584 Rusciie flammea » 405 Russula integra, Spies tts 169; rubra, | 140, 169 Phytogeographical | I. Alpine region, 6773 Rocky Mountain flora—XXVIII, 43; —XXIX, 461 ’ Rynchospora glomerata, 494 be glabra, 3 Saccharum dfacinaearh: 576 Sek. 224; cyatheoides, 225-227; Hillebrandii, 225; pallida, Rt poly- stichoides, 225, 227; Souleytiana, 225; squarrosa, 227; squarrosa aoa erata, 10s: 295,- 327 498; flexuosa, rans 571-573; nigra, 2 Mee. 441-445; natans, 441; Some | rvations concerning ne reactions of ihe teat hairs of, 441 Sambucus canadensis, 383, 385-387. 389, 390, nee septa 589; pubens, 494 Sanguina a, 449 Sapi 3 Sapotacites, 573 Sarothra gentianoides, 612 Sassafras, 394; acutilobum, folium, 383, 385, 389 Saururus cernuus, 387, 390, 393 er a, 686; pennaylvanica, 498; vir- giniensis, 494 Scenedesmus, 476-85; acutus, 76-78, 86 572; varii- Schiz Schlotheimia apa ane Becers 674; torquata, 674; torta, 674; undato- rugosa, 674 ae non-scripta, ligt sibirica, 579 us Eriophoru Sclevonarein waleare po 170 Scopulicola, 6 gatae, 328 y scorphullariaceae, 482 galericulata, 497, 498 schacinn effusa, 164; quercina, 164 Secale cereale, 576 oO 5 th tinh dp) th 4 a 4 a gens, 672 Senecio vulgaris, 502 Sequoia ambigua, 573; fastigiata, 573; gracillima, 569; heterophylla, 569, le saesian yen 569, 579, 572, . | Sida coccinea, af. enastrum, 78 sematophyllum caespitosum, 671; pun- = INDEX | Sericocarpus cc pghntae 494 j meria, I19, 2, 125, 127; hetero- oe 406; Jacksoni, 406; macro- hvylla, 123, 124, 126, 127; pectinata, yee ot issecta, 58; grossu- laractoti, oe pee e a, 495 oes um, ciniatum, 383, 384; sn ie terebinthinaceum, «at 29 Be uheleed 586 Sitilias caroliniana, 387 | Skitophy llum asplenio ides, 662; matum, 661; polypodioides, 661 sper on, M. ew ferns from tropical America—lII, 383; III, 687 Sm the ecithata, nid glauca, 383 Situ, G. radesmus, a new ‘ian celled pare a. 45 Solanum tuberosum, 589; A note on the significance of sugar in the tuber of, pal- Solidago erecta, 496; a. 498; sp., 383 ions concerning the reac- Gos of aad leaf hairs of Salvinia na ie ns, Som aii ogre effects in cul- cues a9 Mechs rghum halepense, as 385-387, 389, a Sparassis crispa, 165 Sparganium hypoboreum, 686; minimum, 86 eau 60; acerifolia, 60; | a, Oey arizonica, 59}. coccines, mig Crandallii, 60; digitata, 5 is- | secta, 58; elata, 58; diflora 0; grossulariaefolia, 58; leptophylla, 59; oe cage 61; aE ae baat - | a, 60; pedat begs 58; ri aac Me Sat erncry Fae 59 | var eto ee Sphaeroplea Sphacrostisma alyseodes macrophyllum, Sphagn £955 Spirodela polyrrhiza, 3 pirogyra, : S8 358, pee longata, 333, 350 e toxic and antitoxic effects enandrium, 121; Coen 121 Stenophragma, 5 Stereohypnum reptans, 665 Stereum purpureum, 139 Stout, A. B. A case of bud-variation in Pelargonium, 367 INDEX ae val sav I5 S 160, 166 Stropharia melas I4I, 160; eee tet a, © ies. pike ae 159, Studies in the Agalinanae, a subtribe of e Hymeno- ycetes, array the Boleti, 137 Studies on the Rocky ae flora— XXVIII, 43; XXIX, Studies on i West aed che spo with one new species from Mexico, 305 Stylosan vid 3 bi flora, 496 arid Sillivantil 497 Swertia Fritillari Fides foetidus, 220, 232; 577 Syntherisma sanguinalis, 383 Syringa, 589 Syrrhopodon, lycopodioides, 660; 661; parasiticus, 660 Tanacetum vulgare, 590 telaaeim polyphylla, 216 acum, 590 Taxodium, 388, 304; distichum, 273, 381, , 389, 392, 393; imbricarium, 394, og Taxonomic study of the Sc ii of the Hawaiian Islands 193 m n tarla ccueintl (76-80, 82, 84; wisconsinen- sis, 76, 77, 8 Tetradesmus, a new four-celled coenobic al ga, Teucrium, 390 Thalesia ‘minut: 485; purpurea, Sedi, 48 4% “emia m purpurascens, 584 Tha nc bopopaigee 665 Tham s Nidus, 20 rhekehen E Gaharentath: 165; palmata, 16 Thermopsis montana, 43; montana ovata, 43; ovata, 43; xylorrhiza, 43 Thuidium involens, 674; microphyllum, huja ee 490 Tiana sia Tiu ctum, Pores atropubescens, 49; acces mitic fg Tomanthera, ios 128; lanceolata, 126, 128 milix, 126, 128; bracteata, 405 Tortula agraria, 659; linearis, 675 by ag humilis, 61; integra, ee > Resbui: Toxopus, 126, 128; calycinus, 405; gym- nanthes Tragopogon pratensis, 590 711 See Genistae, 141, 164; mesenterica, jade enum virginicum, 612 Tricalycites papyraceus, 569, 573, 571 chomanes rhipidophyllum, 687, 690; sphenoides, 6 Trichostema cane a sg 497 Trichos oe beech Tri om chostomum, 675; * ety 675; palli- m watts , 659; strictum, pee ortile, 100 Tricyrtis, 581 t dong oo Siedecalitnps 45; eriocephalum, 45; ephalum, 45; plumosum, 45; eee. 496, 4973 Rusbyi, 45; spinulosum, 45 Trigonella americana, 45; sericea, 45 tS prac i 82 Triphyearia hispida, Tripsacum m dactyloides, yore 386, 387, 380 Triticum vulgare, 576; le cereale, 576 ; us, 494 Tropaeo majus, 587 es S aeatensli, 492, 494; caroliniana, Por-khaienia guatemalensis, 675; linearis 75 Typha latifolia, 383, 386, 387 Ulmus ea a 383, 387, 392; ameri- cana, ;Sp., 38 othrix, are Unifolium canaden: se, 494 Uredo Bupleuri, 509; Gomphrenatis, 5090 Uromyces Pisi, 502, 509; Veratri, 502; 485; Vossiae, 509 U rtica dioica, 584 Vagnera, 582 Valeriana mre 497 Veratrum W' i, 497 Verbena y aaitberonlodies A481; een es 481; can naden: nsis, 481; ciliata, Gooddingii, 481; remota, 481 Verbenaceae, 481 Vernonia, 305, 306, 325, 328; pee 305, 310 321, 325; acuminata, 314; albicaulis, ee albicoma, i 31I, 314; amar 307, 308; stata, 308, sees anus 308; arborescens, ae 308; arctata, 326; aronifolia, 321, 323; bahamensis, 326; buxifolia, as: calida, 315, 318; calo- vila, 375, 3173 chinensis, 306; com- rall 08, plicata, 326; ophila, 308, 300; bensis 0; desiliens, 315, 316; divaricata, 310, 314; Lc SB 310, 318; Grisebachii, sis, 328, 329; hieracioides, 330; tha, _ 307; iserrata, 320, 321; inae- 712 INDEX quiserrata angustifolia, 320, 325; lepto- 259, 260; pedata, 620, 622; pedatifida, clada, 320, 321, 323; lon et hae 330: 249-260; pedatifida XX sagittata, 252, menthaefolia, 331; mon a7; 260; pedatifida < sororia, 253-260; neglecta, 315, 318; orientis ae : perpensa, 259; pratincola, 254; primuli- Ottonis, 305, 328, 329, 330; pallescens, folia, 623, 624; rostrata, 494; rotundi- 329; pluvialis, 312, 314; proclivis, 312, folia, 262, 494; sagittata, 252, 253, 314; purpurata, 321, 322; reducta, 313, 60, 622, 623; poeta 254-260, 265, 314; rigida, 314, 320; Sagra 7320; 266, 270; Wilmattae, 259 321, 323; segregata, 327, 328; ice Viola obliqua Hill aaa other violets, 261 315, 319; Sintenisii, igs ; Viola pedatifida, Four hybrids of, 249 _Sprengeliana, 320, 321; ste een Violaceae, 620 308, 300; stictophylla, 329; sublanata, 305, 310; sublanata angustata, 309; Thomas, 305, 325, 327; Tuerckheimii, 305, mo et onan 320-322; at na, 317; viminalis, 320, 32 Wrightii, 305, 318, 320, 321; saree aed sis, 315; 32 Vernonieae, Studies on the West Indian, (a) Veronica arvensis, 484; ei 484; peregrina, 484; xalapensis, Viburnum dentatum, 498; mae 498 Vicia Faba, 586 Viola affinis, AON 263, 268, sly yooh B di blanda, pea) 621, 623; nephroph yl 259; ne neil ila x 270, eh pten ees Bie 250-25 jestidnaaee x pedatifida, Virgularia, 122, 123; lanceolata, 122 Viscum album, 584 Vitaceae, 606 Vitis aestivalis, rltey igure 606, 607 Vuilleminia comedens, 164 Washingtonia intermedia, 66 Weisia calycina, 661 West Indian mosses—-I; — Wickstroemia indica, 588 Widdringtonites Reichii, subtilis, 571, 573 569, 571, 5733 Xanthium s ; Xylophacos. asi et p pete 493 ee 48; conse » 49; cuspidocarp » 485 andes 4 Pe aflenus, 49; m asisah sis, 49; D utahensis, 49; nace saat ‘“ itn: s, 48 Xyridales, 577 Yucca, 581 Zea Mai 7 Zizyphus Jamarensis 571 Zostera, 586; marina, 576 Buti. 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