OF THE TORREY BOTANICAL CLUB eer VOL. 46 FounDED BY WILLIAM HENRY LEGGETT 1870 EDITOR ALEXANDER WILLIAM EVANS ASSOCIATE EDITORS JEAN BROADHURST MICHAEL LEVINE JAMEs ARTHUR HARRIS GEORGE ELWoopD NICHOLS MARSHALL AVERY HOWE ARLOW BuURDETTE STOUT NorMAN TAYLOR NEW YORK 1919 PUBLISHED FOR THE C Tue New Era PRINTING COMPANY LANCASTER, PENNSYLYANIA CONTENTS STEINBERG, R.A. As study of some factors influencing the stimulative action of ie zinc sulphate on the growth of Aspergillus niger— A comparison of tWo Slrains Of fue tite ie (pate 2) i Gs ig Ge ae Se a ¥ Wituiiams, R. S. Notes on some western Lichens.............-..-e-ceceeee 21 Brown, ELIZABETH DOROTHY WUIST. Apogamy i in Camptosorus rhizophyllus ie Pee ee, GOL 6A I EE ee Ch er ay 27 RAMALEY, FRANC ds at diff Ititud Colorado 37 OSTERHOUT Sas. 7 ‘Additions vi the flora: ot Colorada. a Oe: Sz Levine, MicuareL. The sporadic a ulate of non-edible mushrooms in cultures of Agaricus campestris (plates 3-5)... 0270 oe ee. 57 WEATHERWAX, PAUL. Gametogenesis sal fecundation in Zea Mays as the basis of xenia and heredity in the endosperm (plates 6, 7).........-..+0.005 43 ANDREWS, F. M., and Brats, C. C. The effect is soaking in water and of Seration-an tie growth Of Zea Mavs. ie ee pcb ee eee eS or ARTHUR, JOSEPH CHARLES. News Ale OF Uredinede-—KEs oa ee 107 Brown, Forest B. H. The preparation and treatment of woods for micro- es ie ee ee ei es ev bcc Cw eb bede ave 127 STEWART, E.GrRaAcE. Mucilage or slime formation in the cacti (plate 8)........ 157 Evans, ALEXANDER W. A taxonomic study of Dumortiera........../...... 167 PENNELL, FRANCIS W. Notes on plants of the southern aoa States—V.. 183 GRIFFITHS, Davip. New and old species of Opuntia (plates 9, 10)........... 195 Wi111aMs, R.S. The genus Desmatodon in North America (plate 11)....... 207 ASHE, W. W. Notes on trees and shrubs in the vicinity of Washington....... 22% GLEASON, HENRY ALLAN. Taxonomic studies in Vernonia and related gene 235 Recorp, SAMUEL J. Storied or tier-like structure of certain dicotyledonous NI ee ens ie i ee a ak ae et eae 253 WEATHERWAX, Paut. The ancestry of maize—a reply to criticism.......... 275 Berry, Epwarp W. A new Matonidium from Colorado, with remarks on the distribution of the Matoniaceae (plates 12; 13)... 2... 2... eee ee ee 285 RypsBerc, P. A. Phytogeographical notes on the Rocky Mountain region— VIII. Distribution 7 the Montane poe NS angst Serer tne 1g? Sel Ata 295 Cavace, MaArGARET B. The develop truct f the bulb in Cooperia Pirnmmondit (ties Ta-16) ae i a se nae ee ae 337 PENNELL, FRANCIS W. A brief conspectus of the species of Kneiffia, with the characterization ob. a new allied Benus. {o65 ccyc eee iS ae eee 363 Hitrcucock, Romyn. Preliminary note on a differential staining of the cyto- Aah ts CUA ACCRC Ss fi go eg Po ac es es Pie 5 ss i 375 SMITH, CHARLES PIPER. Studies in the genus peg — Pua; ec; 389 ARTHUR, J. C., and Mains, E. B. Grass rusts of unusual structure.......... 4it BICKNELL, EUGENE P. The ferns and flowering plants of ae! 423 Tuurston, H. W., Jr. Sex in the Conjugatae and the relative frequency of the difteress) types of COMpUSALION... . oo rs eee so a ee ees 441 iv DATES OF PUBLICATION LEVINE, MICHAEL. Studies on plant cancers—I. The mechanism of the formation of the leafy crown gall (plates 17, 18)... ........0-.-- eee ceees Woopsurn, WILLIAM L. Preliminary notes on the embryology of Reboulia Pep nehiCe tiiate TO yee err Ck. ee Seg kG vd Sao ee STEVENS, NEIL E. The development of the endosperm in Vaccinium corym- INDEX TO AMERICAN BOTANICAL LITERATURE, 31, 65, IOI, I5I, 189, 227, 279, 329» 381, 417, 453, 481. TDR: TOT VOR ME CAG eng ce oo ee ee es es ee 489 No. 1, for January. Pages 1-36. Issued January 20, 1919. No. 2, for February. 37-72. February 25, 1919. No. 3, for March. 73~-106. March 10, 1919. No. 4, for April. 107~156. May I, 1919. No. 5, for May. 157-104. May 23, 1919. No. 6, for June. : 195~234. July 8, 1919. No. 7, for July. 235~284. July 31, 1919. No. 8, for August. : 285~334. September 27, 1919. No. 9, for September. 337-388. October 17, 1919. No. 10, for October. 3 November 5, 1919. No. 11, for November. 423-460. December 8, I919- No. 12, for December. 461~502. December 31, 1919. Sisee —e line 12, for “ curea”’ read ‘‘ aurea.’ 199, ine 17, for “‘that to”’ read “‘ to that.’’ a 207, line 8, Dlesmatodon] neomexicanus is a synonym of D. plinthobius (as ated on page 212) and not a species of Tortula, Page 261, line 12, for “* Cladastris’’ read ‘' Cladrastis.” Page 305, line 2 from bottom, for ‘‘ ophoides"’ read “ _SiRniass, a Page 306, line 13, for “ racemosa” read “‘ racemosus. Page 308, line 5, omit ‘* Aster laevis.” Page 312, last line, for ‘* Graphalium” read *‘ Gnaphalium.” Page 318, line 15, for ‘‘ Tolmei”’ read ‘‘Tolmiei.” Pages 335 and 336 are aR Page 412, line 15 from ** Phaksopsora” read “' Pha hakopsora.”’ Page 415, line 8 from Scie for * “ _— wg soroides”’ read ‘* Pela eoldes. Page 430, line 16, for ‘‘ memoralis”’ lis.”” Page 431, line 8 from bottom, for “ preany ‘read “* namie: os BULLETIN OF THE TORREY BOTANICAL CLUB JANUARY, 1919 A study of some factors influencing the stimulative action of zinc sulphate on the growth of Aspergillus niger. Il. A com- parison of two strains of the fungus. R. A. STEINBERG (WITH PLATE I AND FOUR TEXT FIGURES) It will be apparent to those concerning themselves with the “stimulative” action of zinc sulphate on the growth of Asper- gillus niger that marked discrepancies exist between the results reported by different investigators. These variations are most apparent in the yields (dry-weight per 50 c.c. of nutrient solution) obtained in the apparent absence of zinc, the maximum yield in the presence of the optimum concentration of zinc, and the mini- mum concentration of zinc necessary for the maximum yield. Discussion of the variations apparent in the results reported will be confined to the following tabulation of the minimum con- centration of zinc for maximum growth, as given by various auth- ors: Wee a a i er hg ee ee 10.61 mg. Zn/L Weibel 9.10 ~ (9) 8 Sones eee ae aS hs a tag ght rahe al he 9 14 oe 8.41 = Weick ce ne ee er ee 0.09 us Poitier (NF 6 ec os es cree ee eee S ea 0.10 “ The differences are evidently too large to be laid to experimental errors. The experiments taken up in this paper are concerned with observations made on two strains of Aspergillus niger over a [The BULLETIN for December (45: 477-519) was issued December 23, 1918.} 1 > STEINBERG: GROWTH OF ASPERGILLUS NIGER period of somewhat more than six months. The evidence, though fragmentary, since many of the experiments had other purposes in view than that for which they are here utilized, is sufficiently full to be of value in indicating some of the factors responsible for the variations noted above.* From cultures used in previous studies (labelled as originally from the “‘ Internationalstelle fiir Pilz-Kulturen, Amsterdam’’), two one-spore strains were isolated in the following manner: A series of platings and transfers to agar (1 per cent. each of peptone, sucrose and agar) indicated that the cultures could be divided microscopically into two groups based on the relative abundance of yellow pigment in the hyphae. Two tubes were selected, one showing no yellow pigment and the other the maxi- mum amount of yellow pigment; these served as the source material for the isolation of two one-spore strains. Single drops of an agar dilution of the spores from these cul- tures were placed on sterile cover-glasses, and the cover-glasses inverted on Van Tieghem slides. These were placed in the thermo- stat at 30-31° C. After twenty-four hours the slides were exam- ined (Leitz: 4 ocular, 8 objective) and only those retained in which but one conidium, and this germinated, could be seen. The bit of agar containing this single germinated spore was at once trans- ferred from the coverglass to an agar slant. In another twenty- four hours these transfers had fruited and were again immediately transferréd. Great care.was taken in making this transfer not to touch the substratum but only the tops of the conidiophores. The cultures thus finally obtained are assumed to be one-spore cultures. They will be referred to as the W and Y strains, re- spectively. The presence of yellow pigment is more conspicuous in the latter. The second transfer was considered advisable as a check on the possible presence of other conidia (ungerminated) overlooked in the microscopic examination made after the first twenty-four hours. Since a mycelium arising from a conidium does not fruit within twenty-four hours, any ungerminated conidium accident- * In view of the results obtained by me, some of the experiments should be re- peated and elaborated, but since this work has been indefinitely interrupted as a result of my being drafted into the army, I am publishing the data as they stand. STEINBERG: GROWTH OF ASPERGILLUS NIGER 3 ally overlooked will not as yet have had time, even if it did de- velop, to have resulted in spore formation at the time of the sec- ond transfer. The two strains thus isolated were kept on agar slants, of the composition given above, in the thermostat, at 30-31° C. Sub- cultures for the inoculation of the flasks (containing Pfeffer solu- tion) were made on bread and were also kept in the thermostat at 30-31° C. The agar stock cultures were carried in duplicate, transfers being at irregular intervals. A duplicate stock culture once having been used either for the preparation of new stock cultures or of a bread culture was placed aside as unfit for further use. The practice was also made of preserving a single dupli- cate stock culture, unused, from each transfer. The methods used in the experiments described below have already been given in detail in a previous publication (6). I may note briefly that all cultures were kept at 30-31° C., and that 150 c.c. pyrex Erlenmeyers were used. The Pfeffer nutrient solu- tion was made up with water redistilled through glass, “Crystal Domino” cane-sugar, and Merck’s ‘‘Reagent” NH«NO; and MgSO,.-7H20; the other compounds being the Baker ‘‘ Analyzed.” In one or two instances, in which Kahlbaum’s ‘Zur Anal.” MgSO,.-7H,0 was used, this fact is noted. Each culture in the pyrex flasks contained 50 c.c. of Pfeffer solution, the dry-weight or yield representing therefore the amount of tissue formed by the organism in such a culture. Ingculations at all times were from bread cultures. The period of growth of the cultures was a each case seven days. The existence of quantitative differences between the eo strains was quite clearly indicated in the very first experiment (Exp. I Ww eg f Nl ei ee es a ee owe + o> 0.237 gram 0.546 gram Ob Mg. ZNfls 2 eae O08 2": Bd d, OR a The yield of the W strain is less than half that of the Y strain in the apparent absence of zinc, whereas it is slightly greater than that of the Y strain in the presence of 0.1 mg. Zn/L. 2 ee a er * The detailed data for each experiment are given in the Appendix. \ 4 STEINBERG: GROWTH OF ASPERGILLUS NIGER The next experiment (Exp. IT) gave the following results. ING ANGE eae Peale Ga Vote es ESS 0.317 gram 0.480 gram WOT es A a ee ae 4 O.565. 0.781 On the assumption that both strains could, even in the presence of the optimum concentration of zinc, produce not more than one gram of tissue, it is evident that a higher concentration of zinc would be necessary in the case of the W than of the Y strain in order to obtain the maximum yield. The determination of the minimum concentrations of zinc required for the maximum production of tissue was made in quite a simple manner. An experiment was first performed in which zinc in widely different concentrations was used and the values desired thus located approximately as being between two of these concentrations. The next experiment was with concentrations be- tween these limiting values and differing amongst themselves by amounts of 0.01 mg. Zn/L. The first experiment which served for the approximate loca- tion of the zinc optima is here summarized (Exp. III). oe 0.0 mg. oT Gi cick ei crue ere oae ae a 0.217 gram 0.302 — O00T 50 ae Ae eee ee ais oT: 0.620 0.05 a ici eke Arent: ON Ma a eney o7ag. = O47; 4° O.1 ee OA eae a ate a Gace kaa Geeta ray dy dees 0.805 “ I.0 Re ee PN car ONs a ce i arss COIS. o.798° .** These values are shown plotted in TExT FIG. 1. The minimum . _ concentration of zinc capable of bringing about maximum growth is, we note, approximately 0.1 mg. Zn/L for the W strain and about 0.05 mg. Zn/L for the Y strain. We observe, in addition, that the maximum yield of the former is slightly the greater. The more accurate location of the values sought was attempted in a series of cultures differing in zinc content by 0.01 mg. Zn/L (Exp. IV). WwW y: 0.0 mg. Zn/L | 0.119 gram 0.0 mg. Zn/L | 0.343 gram 0.07 = | 0.805 °"* ho 08 ze | 0.344" 0.08 i. 0.900 ‘* bo O03 re 0.803 0,09 he | 0.940 “ i 0.04 ct | 806 “* 0.10 - 0.825 “* ' 9.05 be 0.847 * STEINBERG: GROWTH OF ASPERGILLUS NIGER 5 Reference to figures given for Exps. III and IV indicates that the yield concentration curves intersect in both cases at approxi- mately 0.075 mg. Zn/L. With this concentration the yields with both strains under the conditions employed should be identical and equal to about 0.85 gram. é S m Grams Yield 02 06 06 | Milligrams Zinc per Liter _Fic. 1. For explanation see text. The number of concentrations employed was evidently not suf- ficient to establish the precise location of the minimum zinc con- centration producing the maximum yield, more particularly in the case of the Y strain; and, furthermore, a comparison of the - yields obtained in this experiment with those obtained in the pre- viously cited experiments shows that the organisms are not re- acting uniformly to what are intended to be identical cultural conditions. This is brought out more conspicuously if we include in our consideration the results obtained several months later (Exes. VIII and IX). OD TN Pe oo ele See pie ect 0.136 gram 0.205 gram Ce te ee eae .287 26a 0.1 nT ek Giapeen re algirig’ aca Sue uo ga oe es C872" 0.5 = 0543" 0.826 * TO Fu Pama ne oor —*: 6.8590" 5.0 Ore ey aaa 0.963“ CIS 10.0 7 0.906 “* ie iy & ea 15.0 Prat i cleat ele e ele sees a ewe n002 0.740." 20.0 “ge ne O.00r Oa, 25.0 Tec? Ue irecbieta eee ermcar dig oe Reale «te O45 =" O7s4: For these experiments also the yields have been plotted against zinc concentrations (TEXT FIG. 2). 6 STEINBERG: GROWTH OF ASPERGILLUS NIGER On comparing the results obtained in this experiment with those of the preceding, we become aware of a curious phenom- enon—the action of suboptimal zinc concentrations on the growth has become less effective. Whereas in Exp. II the addition of io - 9 ag gf acanane FETE a is be =—=n mn pe We bts ap tt ES u See ~s CeCe n5 a e it rs Oo A +t i +t ° 2 18 20 ae 24 26 6 8 lO 12 14 16 MILLIGRAMS ZINC PER LITER Fic. 2. For explanation see text. 0.01 mg. Zn/L to the cultures resulted in yields of 0.565 gram and 0.781 gram for the W and Y strains respectively, in Exps. VIII and IX the corresponding yields were only 0.287 gram and 0.283 gram. Even with apparently zinc-free cultures the same change had taken place. In the earlier experiment just mentioned the yields in the apparent absence of zinc were 0.317 gram and 0.480 gram for the W and Y strains respectively; while now they are 0.136 gram and 0.205 gram. It would seem that for suboptimal zinc concentrations (or more precisely, concentrations below 0.1 mg. Zn/L), the initial differences between the W and Y strains with respect to the extent of growth have largely disappeared. The validity of this assumption will, however, again be discussed. Finally, after culturing for thirty-one weeks on the peptone- sucrose agar, the tenth generation transfers of the two strains re- sulted in the following yields when inoculated into the flasks con- taining as usual each 50 c.c. of Pfeffer solution (Exp. XVIII). WwW ¥ OO ME MN. cu... ee enreg ay 0.177 gram ° 0.222 gram Tee tebe aie ie Gedy apa OO Coat aba eg O87. 5 0.440 “ 0.02 Beg leley Cee enan gre ig 0:38). > O.501r .-** 0.03 Me cue eee dee Cay s 7 0.563: °° 0.04 ee eS a eo oa om ee 0.343" " 0.642 >=" O06 oe ee oe 9.534." 0.707“ STEINBERG: GROWTH OF ASPERGILLUS NIGER 7 WwW Y 0.06 mg. sl Bp he Ss ey ey accel 0.688 gram 0.793 gram 0.07 BE Sret ia lalg woateraty cies “eels GOOF." O75 57° OCU Fhe Se ae aig Sie ee eek ce 0.758 0.734 0.09 AAUP chaise Swans MECCA ere + O7OLs 7" 0.770°.;"' 0,10 SP eee sat en ccan i g wee Uuite 0.800 “ 0.799.“ The minimum concentrations of zinc necessary for the forma- tion of maximum dry-weight is for the W strain probably greater, in this case, than 0.1 mg. Zn/L, and for the Y strain, approximately 0.06 mg. Zn/L. The yields in the absence of zinc are a trifle higher than in Exps. VIII and IX. In the presence of 0.01 mg. Zn/L the yield of the W strain is identical in Exps. VIII and XVIII; whereas the yield of the Y strain, though 0.283 gram in the preceting: ex- periment, is in the present trial 0.440 gram. Many of the cultures in Exp. XVIII were photographed and \ Y ; ee Oe | i TI a8 any ‘asi Be BE DR Be PREECE tt Pi Po = wv rl ae es ~~ acl br a ae oe Mo = o & S Ee © r 0.0! 0,02 0.03 0.04 0.05 0,06 0,07 0,08 0,09 0.10 Milligrams Line per Liter Fic. 3. For explanation see text. are reproduced in PLrateE 1. In Text Fic. 3 the yields are shown plotted against the zinc concentrations. To facilitate a survey of the changes in the yields obtained under supposedly identical conditions as these experiments pro- gressed, the yields in ‘‘zinc-free’’ media for the entire period are summarized in the following table: Both strains, it will be noted, showed a rapid decrease in re- spect to their capacity for growth in the Pfeffer solution. The two strains, nevertheless, are distinct in that the W always forms a smaller amount of tissue than the Y strain. The fact that a decrease in the capacity for growth has indeed taken place will 8 STEINBERG: GROWTH OF ASPERGILLUS NIGER Weeks si isolation | Number of transfers Ww ¥ ‘ I Original | 0.237 gram 0.546 gram 3 I On aa 3.486." 5 2 0.217 0,302 = 8 3 | O1I9. O.545 9 - 6.105. O308 rae) 3 O:530): C270 G5 19 5 05354" O.4400-°: 20 6 O,1g0- — 22 6 | — 0:205.° 7" 23 7 O:520 20. — 24 7 0.086 “* 25 pole | Osho st 8.41862 26 8 | Oras." 27 8 Gage: o: — 28 9 Oxs7 0.232 29 10 @:08s C4708 30 Io or30 Ors 31 Io ap) ty a ae C272 become more evident, in spite of the confusing experimental vari- ations, on reference to the effect of constant amounts of zinc on- the growth at different periods of time. Summarizing the yields obtained for the entire period with concentrations of 0.01 and 0.10 mg. Zn/L we find: Weeks since pooner of oon mg. nil. | oro mg. Ene isolation ransfers w | y | w Y I Origi | | 0.903 g o.81I g 3 Ai 0.565 g. | 0.781 g — os 5 2 | fe a By say 0.805 ‘* 8 3 _ 0.635": ° 6 0.287 “ —_ 0.851 “ 2 6 _ 0283."° — 0.823" 30 10 0.198 “* 0.268 *‘ _— oa 31 10 0.287 “ 0.440 “ ooeo © | O70." The results obtained in ‘‘zinc-free”” media and with media con- taining 0.01 mg. Zn/L are shown graphically in TEXT FIG. 4. There is, it will be seen, a perceptible decrease in the yields obtained with both strains with time. While experimental variations due to at present unascertained conditions tend to obscure this phenomenon, a survey of the yields for identical zinc concentrations cannot but make evident that the capacity for growth of the organisms in the presence of small amounts of zinc has decreased. Not only the apparently zinc-free but also the zinc cultures exhibit the same result. Whereas strain STEINBERG: GROWTH OF ASPERGILLUS NIGER 9 W when grown in the absence of zinc resulted initially in a yield of 0.237 gram, and the Y culture of 0.546 gram, the final yields were 0.177 gram and 0.222 gram. Similarly for the cul- tures grown in the presence of 0.01 mg. Zn/L, the initial yields were 0.565 gram and 0.781 gram and at the time of interruption of os A OS NCCC Nwebloie aay yslololm aay CT — A Oe CCEE eee SN Cece POCO EEN Pere os ARES on erect S22 SR2222-.—_ me te 2/8 VEER P St 209 ana an oor es oo e718" O8225°,"" gaa yp. 2 a O.872°.5" 0.302 “ 0.620 “ [yeas ies eae 0.798 “ EXPERIMENT IV 8 weeks W 3 No zine 0.07 mg. Zn/L 0.08 mg. Zn/L 0.09 mg. Zn/L o.10 mg. Zn/L 0.143 gram 0.833 gram 0.947 gram 0.935 gram 0.806 gram 0:092 <5. O85o! G.005 -. = 0.885 “* iy £1 Aas O19. G685° " O.593°.5- Horr: ** 080200" O16 3. 0.786 “‘ OB57 6.052" O870. = 0.44. et 1. O@6r = OBA 2! Tort... i tb leis O16 i" | 0.805! 0.900 ."* 0.940 “ o8as ¥3 No zinc 0.02 mg. Zn/L 0.03 mg. Zn/L 0.04 mg. Zn/L | 0.05 mg. Zn/L 0.307 gram 0.818 gra 0.757 gram 0.786 gram ~=—séiO«wWBQ gram. 6377 5°. 0.830." OS3r oc: 1 aha 0.85 i G.200.7<;* O85e 3% C786. 2" 0.828: VO Oey 0,369)...“ 0.856. “* O:8465 5S! 0.757. Bae 3S Saks O3660.0-- 6.856." 0.704 Lon a leaded | O:878 087 0.343 “ 0.844 ae 0.803 “e 0.806 ae | 0.847 “ee EXPERIMENT V 9 weeks, no zine W3 | Y¥3 0.127 gram | 0.327 gram * 0.908 i { .362 a G.078 0.545 4" O.085 3" | 0.206. erg “ L268. * o1os 2 | 6,307. 16 STEINBERG: GROWTH OF ASPERGILLUS NIGER EXPERIMENT VI 10 weeks W3 x3 No zinc 0.09 mg. Zn/L No zinc oo2 mg. Zn/L 0.104 gram 0.822 gram 0.301 gram 0.822 gram 0.005 °° 0;80200" 0,264 °°" O.02T 05" O;Fa8 Oo i700" "" O.315..7- EOGs oo. 0.173 “e 0.752 6é 0.244 “ 0.840 se og se aaa 0.816..." O25" 5 OB7A 0.139 =“ 0.792“ 0.276." 0.902 “ EXPERIMENT VII 19 weeks, no zinc Ws Ys5 0.133 gram 0.218 gram Ota) O.252 —. O.146 0.263 5 Ae Fi eas e250 o* O.12i C218. 1" O14 = 6.2405"! EXPERIMENT VIII 20 weeks, W 6 No zinc o.or mg. Zn/L o.1 mg. Zn/L 0.5 mg, Zn/L 1.0 gm. Zn/L 0.138 gram 0.298 gram 0.835 gram 0.817 gram 0.932 gram 0.137 OS cs 0.528 .-** CO Se Oe Retater tl 0.80 bi 0.174 ® 0485." O.Bse | C82e 2)" O.072 00°F 0.104 if 0.200." O.835 °° 0.840 ‘ 0.910 “ 0.125 O26 s 0.895 6.916"? 0.958. * 0.536. > 0.287 2" | 08 me pec 0843.20 O05." 5.0 mg. Zn/L 10.0 mg. Sn/L 15.0 mg. Zn/L 20.0 mg. Zn/D 25.0 mg. Zn/L 0.914 gram 0.880 gram | 0.905 gram 0.947 gram 0.944 gram 0.932 . 0.869 “* 6.871 * 0.860 “ 0.052. “ SO. 0.903 “* 0.894 “* G.o0r) “* oni 0.967 fc 0.041 \* 0.022." 0.014 0.943 “* 1.024 0.939 “* 0.917 “ 0.883“ 0.973.“ 0.963 . — 6,906.“ 0.9025'7 @.901 «°° oo45> sa a > STEINBERG: GROWTH OF ASPERGILLUS NIGER I7 EXPERIMENT IX 22 weeks, Y 6 No zinc | o.or mg, Zn/L | o.r mg. Zn/L | 0.5 mg, Zn/L 1.0 mg. Zn/L 0.242 gram | 0.270 gram 0.843 gram | 0.808 gram 0.854 gram GOI86: = | Gigaenc O836" ** Poumon mes (e 0.86520" C203 OTR: =" 0,825" ."" Fy Lanta 0;3843°° "7 re 20 ¢ ae | G2500° 0.798 * OS45 O:864°05° eet ipele bape a Spa | EPCS CMa as" Coy. O05 |... | O26 oe" 0.823 “ te G.820.(.5' 0.859 “* 5.0 mg. Zn/L | to.0 mg. Zn/L | 15.0 mg, Zn/L | 20.0 gm. Zn/L 25.0 mg. Zn/L 0.750 gram 0.791 gram 0.700 gram 0.706 gram 0.755 gram O.736 " a7s9 2" ri lS ge ore Faery O98 oc 0.739 =“ 0.774 “ 0.760 ** 0.747 “* 0.739 “ D.7A0). 717 vob fev egies O7O8 0.770 - 5 De NS 0.750" (Pu at as O: FAG 0 = eae ar et anes } Orgy 1 Ae ae @ Sea O746. 0° Re ar see LE, aaa EXPERIMENTS X, XI AND XII No zinc X, 23 weeks : XI, 24 weeks XII, 25 weeks W> | W7 Ww; Wi | Y7 | Vx Lee ' 0.211 gram 0.078 gram | 0.170 gram | 0.101 gram | 0.270 gra | 0.164 gram 0.907 0.082 “ eies eo) eae oS 0.262 * 0.196 “ 0.091 “‘ O,t48-)) 0.066 “ | 00! Rae nC (i ae ae 0.087 <<" C125." 0.098 ‘“ | OBO 4 Cage se Gaia OOOO. * O1§s =" O09 6 ol O60 [0.252 °° { \ pees gene 0.086 “ TRE ck 0.098 “* oc 4 6.216% EXPERIMENTS XIII AND XIV W 8, no zinc XIII, 26 weeks XIV, 27 weeks 0.131 gram 0.136 gram O25": 0.200 “* aM aie O:1390°> "7 OF C555... 0.196. 0.095 “* 0.426..." O5AL 18 STEINBERG: GROWTH OF ASPERGILLUS NIGER EXPERIMENT XV 28 weeks, no zinc Wo Yo 7W3 ik See 0.184 gram 0.247 gram 0.110 gram 0.255 gram Ye oa eee age Ao. aia 0.070 0,267>. Ors 0248" O:t80 i." C20r 0.597855" 0.1904: ** 0.168 °° e269 = 40100 8.220 °° "* Gist) =: G.42G5° >" O.257)-"" Oeag: 0 G47 6.369..2%* EXPERIMENT XVI 29 weeks, no zinc W 10 —8W3 +8W3 Yu10 —Y3 | +8 V3 0.070 gram | 0.145 gram | 0.114 gram | 0.167 gram | 0.167 gram | 0.101 gram COGE Sor! 0.%44 ** OSGeo 2" OO: poe Gaede 0.19 +5 O04 0.094 “ 0.108" O76". o 0.196 “ 0.363 50°" O:083. ERE ee 0.004 "* Crype 7's O.180° 0 "% O,.I04 = O76: > Ota 0.103..." 0.199 «“* OF47 * aid epee 0.085 135°" O154 < O76 2 O74)“ 0.179.“ EXPERIMENT XVII 30 weeks W 10 Y 10 No zinc _ oor mg. Zn/L No zinc o.or mg. Zn/L 0.113 gram 0.195 gram 0.179 gram 0.199 gram O.210 11 O.268."' 0.1 iy G:358 ** oO 137 oe 0.226 ae o 195 ee 0.269 ae O20; G60.) ** O.376. O.202 7 0. 150 “e 0.140 as ry 0.223 4é G130...“ o.168* oiler 0.268. ."' 7W4 7V¥4 No zinc 0.01 mg. Zn/L No zinc 0.01 mg. Zn/L 0.115 gram 0.212 gram 0.159 gram 0.197 gram 0.115 6.8572" O.476 7% 0.276 * t2r 7) Dus OF aad O.78S (2° BAG 0.084 : Orgs. Se ow GetPos 0478” 0.112 0.208 O.286 0.246: 0.109. .** O.166 :°! O75. Peay © o° Gegichs STEINBERG: GROWTH OF ASPERGILLUS NIGER EXPERIMENT XVIII 31 weeks W 10 No zine o.or mg. Zn/L 0.02 mg. Zn/L 0.03 mg. Zn/L 0.04 mg. ZnL 0.203 gram 0.271 gram 0.386 gram 0.473 gram 0.525 gram OX74 0.40812" Een ds eee cr de ra ate Sines 0.140 <<": : O.106 5" oe ss ae oe ee an 0.2875" 0.387 °°" ways a i se RI _ 005 mg. Zn/L| 0.06 mg- Zn/L | 0.07 mg. Zn/L | 0.08 mg. Zn/L | 0.09 mg. Zn/L | 0.10 mg. Zn/L 0.466 gram |} 0.704 gram | 0.698 gram | 0.868 gram | 0.722 gram | 0.821 gram aHoee =" oh oly Satie G.605° 7° 6.647557" 0.79 5 Hore.°-*? a eT ee 0.6885)" 0.607. “ O86 o76re 6800" Y ro * No zinc o.or mg. Zn/L 0.02 mg. ZnL 0.03 mg. Zn/L 0.04 mg. Zn/L 0.186 gram 0.524 gram 0.517 gram 0.578 gram 0.650 gram C2127. O56. ** 0.485 “* was 6.637. °% 0428557 G.252 >" O22e a. 0.222. > >" Cae O.ROL' O.564. 6." 0,642 °’ 0.05 mg. Zn/L| 0.06 mg. Zn/L | 0.07 mg. Zn/L | 0.08 mg. Zn/L | 0.09 mg. Zn/L o.to mg. Zn/L 0.674 gra 0.862 gram | 0.765 gram | 0.753 gram | 0.765 gram | 0.794 gram Ojsg o424° * O7AT OFES. C765.) 0.804 “ 0.707 0.793.°°° 0.7537 944 1 AIO.” 799 .* 20 STEINBERG: GROWTH OF ASPERGILLUS NIGER LITERATURE CITED 1. Javillier, M. La presence et le rdle du zinc chez les plantes. Thése. Paris. 1908. 2, Ono, N. Uber die Wachsthumsbeschleuniging einiger Algen und Pilze durch chemische Reize. . Jour. Coll. Sci. Imp. Univ. Tokyo 13: 142. 1900. 3. Raulin, J. Etudes chimiques sur la végétation. Ann. Sci. Nat. Bot. V. 13: 93. 1869. 4. Richards, H. M. Die Beeinflussung des Wachsthums einiger Pilze durch chemische Reize. Jahrb. Wiss. Bot. 30: 665. 1897. 5. Richter, A. Zur Frage der chemischen Reizmittel. Die Rolle des Zn und Cu bei der Ernahrung von Aspergillus niger. Centralbl. Bakt. II 7: 417. 1Igor. Steinberg, R. A. A study of some factors influencing the stimula- tive action of zinc sulphate on the growth of Aspergillus niger. 1. The effect of the presence of zinc in the cultural flasks. Mem. Torrey Club 17: 287. 1918. 7. Thom, C., & Currie, J. N. The Aspergillus niger group. Jour. Agr. Research 7:1. 1916. de Explanation of plate 1 All photographs from Exp. XVIII First Row. Wstrain. No zinc added to the cultures. SECOND ROW, strain. No zinc added to the cultures. THIRD ROW. W strain. Cultures contain from left to right 0.01, 0.02, 0.03, 0.04, FourTH ROW. W strain. Cultures contain from left to right 0.06, 0.07, 0.08, 0.09, 0.10 m: FIFTH ROW. Y strain. Cultures contain ‘iiss left to right 0.01, 0.02, 0.03, 0.04, 0.05 m SIxTH ROW. Y strain. Cultures pans from left to right 0.06, 0.07, 0.08, 0.00, oO. /L. SEVENTH ROW. Cultures from left to right: W zinc-free, Y zinc-free, W 0.1 mg. Zn/L, Notes on some western Lichens R. S. WiLtrams Having occasion recently to write up the labels for a collec- tion of lichens made in the Yukon region, in the years 1898 and 1899, and also for a small collection I made chiefly in the Blackfeet Indian Reservation of northern Montana, in the summer of 1897, I found a number of unnamed species and also various corrections which it seemed to me might be made. SPECIMENS FROM YUKON The Yukon specimens were often in none too great quantity as they were picked up as occasion permitted without any previous preparation for that line of work. Duplicates of most of the Yukon collection I sent to the late T. A. Williams, of Washington, and to Miss Cummings, of Wellesley, and perhaps two thirds of these were determined by Mr. Williams, but no list published. Some years later another not very complete set was sent to Dr. R. Heber Howe and of these, determinations, partly made by Dr. Hasse, were published in the Bulletin of the Torrey Botanical Club (38: 287-293. 1911). I now find in going over the original collection a number of species not seen by Dr. Howe and also some discrepancies owing, doubtless, partly to mixtures, lack of material or poor specimens. — I shall first mention the Yukon plants not sent to Dr. Howe. They are the following well-known and widely distributed species: LECIDEA DECIPIENS (Ehrh.) Ach. Dawson, on rock, April, 1899 (49). LECIDEA GRANULOSA (Ehrh.) Poetsch. Hills above Lake Lindeman, May, 1898 (50). CLADONIA PYXIDATA (L.) Hoffm. Lake Lindeman, May, 1898 (4a). RHIZOCARPON GEOGRAPHICUM (L.) Lam. & DC. Dawson, April, 1899 (82). Ze WILLIAMS: NOTES ON SOME WESTERN LICHENS RHIZOCARPON PETRAEUM (Wulf.) Massal. Dawson, very common (Z07). PELTIGERA APHTHOSA (L.) Hoffm. Lake Lindeman, May, 1898 (34). NEPHROMOPSIS CILIARIS (Ach.) Hue. Lake Lindeman, April, 1898 (76). CETRARIA ISLANDICA CRISPA Ach. Lake Lindeman, April, 1898 (15a). PARMELIA PHYSODES (L.) Ach. Dawson, common, October, 1898. One species referred to in Howe’s list, however, I do not find in my collection, namely, that under No. 11 in his enumeration, which is called Biatora franciscana Tuck. This was part of my collection number 56; in what is left everything is evidently Lecanora calcarea (L.) Nyl., as determined by Dr. Hasse. The remaining notes on Yukon specimens are taken up in order, according to Dr. Howe’s numbers, and relate to changes, the numbers in parentheses being my collection numbers. 8. LECIDEA TESSELLATA.Floerke. The specimen (52) was sent to Dr. Hasse, and referred to Lecidea, but was ‘‘too fragmen- tary for a satisfactory examination,” in his opinion. 33. LECANORA LENTIGERA (Web.) Ach. On earth of river bluff just below Dawson, April, 1899 (64); evidently this, a fine and apparently rare species. Another specimen, however (87), questionably referred to L. crassa, under the impression that it was not distinct from L. lentigera, is certainly quite different from either It isevidently L. thamnoplaca Tuck., not previously known north of | Montana and but rarely collected. It was first obtained by Bo- lander in Nevada and next by Dr. Coulter in Yellowstone Park. 34. The specimens under this number, called Lecanora thamno- placa Tuck., Dawson, April, 1898 (52), are all in my packet Lecanora fruticulosa (Dicks.) Ach. 35. The two collections here referred to Lecanora pallida have rather too large spores for that species. One (63a) has ellipsoid spores f n, which I should refer to L. tartarea (L.) Ach.; the other WILLIAMS: NOTES ON SOME WESTERN LICHENS 4) (00); with spores ns a, 1 should think Ochrolechia pallescens (L.) 7 siti The spores of L. pallida (Schreb.) Schaer. are given as =r ary otherwise O. pallescens is very similar to L. pallida. 36. Called Lecanora subfusca var. argentea Ach. This is cer- tainly the same as No. 34, or L. fruticulosa, previously credited in North America only to California, I believe. 39. Called Lecanora epibrya Nyl. Dawson, July, 1898 (30). My specimens apparently not distinct in any way from the pre- ceding. 40. In my set none of the specimens under this number (88) are distinct from No. 34. Lecanora castanea (Hepp) Th. Fr., as given in Howe’s list, does not seem to be represented. 53. All poor specimens under this number (97), apparently charred by fire. They were doubtfully referred to Ephebe pubes- cens (L.) Fr. Collected at Dawson, July, 1898 (97). I think the species is rather Lecidea globifera Ach., from which I believe Biatora Russelliit Tuck. is not distinct. 56. This is typical Solorina saccata (L.) Ach., rather than the var. spongiosa, which it is called. Collected at Dawson, August, 1898 (35). The other specimen under the name S. saccata, from Lake Lindeman, May, 1898 (22), seems to be rather ocmtig: hypnorum (Wahl.) Koerb. The spores are not more than - Be and eight in the ascus. Solorina saccata has spores mostly four 6-55: 18-28 © Three other species referred to in the list proved to be inde- terminable in the duplicates sent out. I believe them to be as follows: ; CLADONIA GRACILIS CHORDALIS Floerke. Lake Lindeman, May, 1898 (6). BUELLIA COLLUDENS (Nyl.) Tuck. Dawson, December, 1898 (82), with no spores; Dawson June, in the ascus and™ 16 ; : 1899 (82a), with spores two-celled, about “— p, pale and eight in the ascus. 24 WILLIAMS: NOTES ON SOME WESTERN LICHENS LECIDEA DECIPIENS (Ehrh.) Ach. Dawson, on rock, April, 1899 (49). A widely distributed species. SPECIMENS FROM MONTANA ENDOCARPON TORTUOSUM Herre. Near Big Badger Creek, Blackfeet Indian Reservation, Sep- tember, 1897 (92). This species was described by Herre in 1911 from specimens collected near Reno, Nevada. I find no other record except these Montana specimens. The genus is peculiar in having oblong, muriform, brown spores, mostly two in the as- , ‘ ‘ Qo cus; in this species measuring about = ENDOCARPON PUSILLUM Hedw. Heart Butte, September, 1897 (zoo). A much smaller species than the preceding, closely adnate to rocks and forming black- ish discolorations. Spores similar to, and nearly as large as, the preceding, measuring about = p. LECIDEA AMYLACEA Ach. Henry Mountain, Blackfeet Indian Reservation, August, 1897 (95). Weseem to have in the museum no specimen of this from North America. It is credited to Greenland and to Utah by Tuckerman. LECIDEA ARMENIACA (DC.) Fr. Blackfeet Indian Reservation, August, 1897 (87); fruiting. This species has been rarely collected in North America, and not before in fruit, I believe. In many respects, the plant is not so very unlike Lecanora esculenta of Arabia, which is supposed to be the manna of the ancient Jews. The taste is evidently quite similar. ACAROSPORA RHAGADIOSA (Ach.) Fr. On perpendicular walls of sandstone just below the Great Falls of the Missouri, February, 1889 (go); in fruit. This appears to be the only collection made of this species in North America. It seems a well-defined species, known previously only from Europe. The determination was by Nylander. WILLIAMS: NOTES ON SOME WESTERN LICHENS 25 ACAROSPORA CERVINA (Wahl.) Koerb. Two-medicine Lake, Blackfeet Indian Reservation, August, 1897 (97). This is a small western species, on rock, with apo- thecia more or less immersed and the ascus crowded with minute spores about 4 pu. It has been mostly collected in California, under the name Lecanora fuscata (Schrad.) Th. Fr. This seems to be its most northern record. ACAROSPORA CHLOROPHANA (Wahl.) Massal. Henry Mountain, Blackfeet Indian Reservation, at 7000 ft., August, 1897. One of the most conspicuous of yellow, rock-loving species. The Golden Gate (or Gardiner) entrance to Yellowstone Park takes its name, I believe, from the quantity of this lichen covering the rock-walls near. LECANORA ATRYNEA (Ach.) Nyl. Near Two-medicine Lake, August, 1897 (90). The speci- mens more closely resemble some from the Pyrenees, determined by Nylander, than those from California collected by Herre. The California specimens are the only ones of North America in the museum, but it is mentioned in Miss Cummings’s list of Alaska species as credited to that region by Dr. Almquest. LECANORA THAMNOPLACA Tuck. Columbia Falls, November, 1893, in fine fruit (gr); also col- collected east of the Rocky Mountain Divide in Montana. BLASTENIA FESTIVA (Fr.) Hasse. Near Forty-mile Creek, Blackfeet Indian Reservation, August, 1897 (z06). A small and inconspicuous rock-loving species. These Montana specimens appear to be the only ones collected outside of California in this country, but I believe are correctly referred here although the spores are not polar-bilocular as ordi- narily occurs in this genus. Hasse states that the spores may be simply bilocular. RINODINA CHRYSOMELAENA (Ach.) Tuck. Forty-mile Creek, Blackfeet Indian Reservation, August, 1897 (116). This species has been but rarely collected and not before to the westward of the Mississippi, I believe. New York BotanicaL GARDEN . % : ie < “ 2 é P is » bP) oe . 2 A i a * ; y blige cd Z es is mn if _ ‘ : " ¢ " Fae \ =a . t yh ante 2 ‘3 ? oe 7 neat ; e . 4 pigs Bu i rf z eae Apogamy in Camptosorus rhizophyllus* ELIZABETH DorotHy Wuist BROWN (WITH PLATE 2) The life history of Camptosorus rhizophyllus (L.) Link, in both generations, has been studied in detail by Pickett.t An ecolog- ical study of the prothallia has also been published by the same author,f but no mention is made of observed cases of apogamy either in the field or in cultures. Neither has apogamy been recorded for this fern by any other investigator. The case of apogamy described in this paper occurred in a culture of the fern which had been made, along with cultures of other ferns, to determine if apogamy could be induced by the mod- ification of external conditions. MATERIAL Collections of the fertile fronds of Camptosorus rhizophyllus were made during the month of August from plants growing on the limestone cliffs of a glen in the vicinity of Ithaca, New York. They were allowed to dry by exposing the sealed envelopes, in which they had been placed in the field, to sunlight before an open window for about a week. For future use the envelopes were placed in a pasteboard box in the laboratory. Cultures were made by sowing the spores thickly on 26 c.c. of Knop’s full nutrient solution, to which had been added a drop ofal per cent. solution of ferric chloride, in small glass capsules. The formula of Knop’s solution is as follows: Ma a ea eee 0.25 xan CoINOps Foe Se ph ee Se ge a ey I.00 POs a ea OOO es eee es 0.25 * Be cow ee we oa eA eae pe ot pee e a Re Oh ag * Contribution from the Osborn Botanical Laboratory. The development of the prothallium of Camptosorus rhizophyllus. Bot. Gaz. 57: 228-238. pl. 12,77 +f.1-8. I914. “Some ecological adaptations of certain fern prothallia—Camptosorus rhizo- phyilus are Asplenium platyneuron Oakes. Amer. Jour. Bot. 1: 477~498. pl. 49,50 +f.I-19. 1914. ! 27 28 Brown: APOGAMY IN CAMPTOSORUS RHIZOPHYLLUS After the spores had been sown the cultures were placed be- fore an east window and the culture solutions were not renewed. DEVELOPMENT OF THE PROTHALLIA Germination began in about one week after the spores were sown and the growth and development of the prothallia were rapid at first, then slower as the amount of nutrient solution in the capsules decreased. The size and shape of the prothallia varied greatly. In the more crowded regions only a few devel- oped into more or less heart-shaped prothallia, while the others in these regions were ribbon-shaped, irregular or branched. In the less crowded regions, especially near the margin of the cultures, more developed into heart-shaped prothallia. All the prothallia were smaller than when developed in cultures where the nutrient solution was renewed from time to time. Many of the prothallia showed the great irregularity of margin and the various types of marginal outgrowths described by Pickett. Antheridia in large numbers developed on the ribbon-shaped and branched prothallia, while the heart-shaped prothallia developed meristem which bore first antheridia and later archegonia. The latter evidently did not function, as no normal sporophytes were formed, although large quantities of antherozoids were present. In about ten weeks after germination began, an apogamous sporophyte was found on one of the larger prothallia near the margin of the culture, in one of the less crowded regions. Ina few weeks after the apogamous sporophyte developed, the nutrient solution in the capsules having been exhausted, new nutrient solu- tion was added. The stimulating effect of this new solution could soon be noted by the appearance and growth of the prothallia and by the development of a few normal sporophytes. However, this stimulating influence seemed of short duration, as the culture soon began to decline. A number of the prothallia in the more crowded regions turned brown and appeared dead, while no more sporo- phytes were produced. When the solution was again exhausted new was added and again the stimulating influence could be noted, but it was not as noticeable as it had been before. The prothallia showed increased vigor, although no more sporophytes developed. Many of the prothallia which appeared dead gave rise to prolif- BROWN: APOGAMY IN CAMPTOSORUS RHIZOPHYLLUS 29 erations which developed into prothallia bearing antheridia. Thus by adding new solution from time to time when the old became exhausted the culture has been kept living up to the present time. Some of the prothallia appear normal, having a meristem bearing both antheridia and archegonia, while others have given rise to proliferations which in turn bear antheridia. DESCRIPTION OF THE APOGAMOUS GROWTH The prothallium on which the apogamous outgrowth occurred was somewhat irregular in shape and only one cell in thickness (Fic. 1, P). No sinus was present and at the point where one usually occurs in larger and older prothallia a marginal structure, or lobe, had formed (Fic. 1, L). The apogamous outgrowth began its development as a swell- ing in the region of the prothallium where the meristem begins to form. This swelling continued to grow, forming a cylindrical pro- cess several cells in thickness which became somewhat narrower as it developed. After growing for some time in this manner re- version to a narrow prothallus-like structure, only one cell in thickness with an irregular broadened apex (Fic. 1, AO), took place. At one point a branch, irregular in shape and itself show- ing a slight tendency to branch, developed (Fic. 1, B;). On the opposite side of the prothallus-like oA aka near the ewes an- other branch was formed. This branch-p elongated, one cell in thickness with a bibad slightly heart-shaped apex (Fic. 1, Bz). Rhizoids developed from some of the marginal cells near the broadened apex (Fic. 1,R). On one side of the elongated part of this branch-prothallium, the marginal and ad- jacent cells were smaller and more numerous. From this region an archegonium, normal in appearance, developed (Fic. 1, A:). The venter of the archegonium was not imbedded in the tissue of the prothallus but the whole archegonium extended beyond the margin of the prothallus (Fics. 2, 3). Two archegonia developed on the apogamous growth near its point of origin (Fic. 1, As) and just beyond this point a cluster of tracheids was formed in the tissue of the outgrowth (Fic. 1, T). As well as could be observed, owing to their imbedded position, these tracheids resembled those of a normal sporophyte. 30 Brown: APOGAMY IN CAMPTOSORUS RHIZOPHYLLUS DIscUSSION Experimental data on induced apogamy would seem to indi- cate that bright light and insufficient moisture for fertilization were the controlling factors for the forms studied. In this partic- ular case an insufficient food supply seems of greater importance than these two factors for the following reasons. Apogamy was never observed in other cultures of this fern under the same light conditions where the nutrient solution was renewed frequently. The prothallium upon which the apogamous outgrowth occurred, while developing near the margin of the culture, was crowded by other prothallia and shaded by them. Sufficient moisture for fertilization was available, as the prothallia developed upon a liquid medium. Later, when the nutrient solution was renewed, thereby supplying sufficient nourishment, normal sporophytes developed in the culture, although conditions of light and moisture remained unchanged. The fact that the part of the apogamous outgrowth which was last to develop showed a reversion from a more or less complex structure several cells in thickness bearing tracheids to a simple prothallus-like one only one cell in thickness and branched indicates a lowered vitality of the outgrowth. Goebel considers a reversion to a juvenile form as being the result of unfavorable con- ditions. The unfavorable condition in this instance was doubtless an insufficient food supply. However, the effect of this factor in inducing apogamy will be discussed at a greater length in a future paper on induced apogamy in Phegopteris polypodioides Fée. Since only one case of apogamy was observed in the experi- mental cultures, it would seem that apogamy is of very rare occur- rence in Camptosorus and not easily induced. Explanation of plate 2 Fic. 1. Prothallium showing apogamous outgrowth, X 120: P, prothallium; AO, apogamous outgrowth; Aj, Ag, archegonia; T, tracheids; Bi, Bo, branches; R, thizoids; L, lobe of prothallium. Fics. 2, 3. Surface view and median optical section view of the archegonium borne on the margin of the branch of the apogamous outgrowth, & 275: N, neck; C, canal; V, venter; E, egg; P, prothallium. INDEX TO AMERICAN BOTANICAL LITERATURE 1915-1918 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 pee exclusively to forestry, agriculture, horticulture, manufactured products of vegetable origin, or laboratory methods are not included, and ea mpt is made to index the literature of bacteriology. An occasional exception is n favor of some paper appearing in an American periodical which is devoted ae to botany. Reprints are not mentioned unless they differ from the hl in some important particular. If users of the Index will call the attention of the edito: to errors or omissions, their kindness will be appreciat dex is reprinted monthly on cards, and farnishea i 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- ndence relating to the card issue should be addressed to the Treasurer of the Torrey 1 Club. Spo! Botanical C Angli, J. La ‘‘Araucaria araucona” (Mol.) Koch (= Araucaria im- bricata R. Pav.) y su resina sus relaciones con las demas coniferas. Bol. Acad. Nac. Cien Cérdoba 23: 1-84. f. I-21. 1918. Anthony, R. D. Methods and results in grape breeding. Proc. Soc. Hort. Sci. 1914: 81-86. Mr 1915. Atkinson, G. F. The genus Endogone. Brooklyn Bot. Gard. Mem. r: 1-17. 6 Jl 1918. Atkinson, G. F. The genus Galerula in North America. Philos. Soc. 577: 357-374. 19 Au 1918. Nineteen new species are described. Blakeslee, A. F., & Avery, B. T., Jr. A vegetative reversion in Por- tulaca. Brooklyn Bot. Gard. Mem. 1:18. 6 Jl 1918. _ [Abstract]. Britton, E.G. Musci. In Britton, N. L. The flora of the American Virgin Islands. Brooklyn Bot. Gard. Mem. 1: 103, 104. 6 Jl 1918. Britton, N. L. The flora of the American Virgin Islands. Brooklyn Bot. Gard. Mem. 1: 19-118. f. z. 6 Jl iy Includes chapters contributed by A. W. E W. Riddle, E. G. Britton and M. A. Howe. New species are d described in Opuntia vate ‘ef ermees (1), Lecania (1), and Miciiciste (x). Burns, G. P. Weather conditions and plant development. Brooklyn Bot. Gard. Mem. 1: 119-122. 6 Jl 1918. 31 Proc. Am. a2 INDEX TO AMERICAN BOTANICAL LITERATURE Burt, E. A. The Thelephoraceae of North America—IX. Aleurodis- cus. Ann. Missouri Bot. Gard. 5: 177-203. f. I-14. 205 1918. Seven new species in Alewrodiscus are described. . Connors, C. H. Heredity studies with the carnation. Proc. Soc. Hort. Sci. 1914: 95-100. Mr I9I5. Coons, G. N. Seed tuber treatments for potatoes. Phytopathology. 8: 457-468. f. 1-6. 12S 1918. Cook, M. T. Modern applications to botany. Brooklyn Bot. Gard. Mem. 1: 123-127. 6 Jl 1918. Dixon, H. N. The mosses collected by the Smithsonian African Ex- pedition 1909-10. Smithsonian Misc. Col. 697: 1-28. pl. 1, 2. 8 O 1918. Twelve new species in various genera are described. Dudley, P. H. Fungi the cause of decomposition of timber. Wood- Preserving 5: 26-35. f. r-10. S 1918. East, E. M. Intercrosses between self-sterile plants. Brooklyn Bot. Gard. Mem. 1: 141-153. 6 Jl 1918. Elliot, C. Bacterial oat blight. Phytopathology 8: 489, 490. 12S 1918. Evans, A. W. Hepaticae of St. Croix, St. Jan, St. Thomas and Tor- tola. In Britton, N. L. The Flora of the American Virgin Is- lands. Brooklyn Bot. Gard. Mem. 1: 104-109. 6 Ji 1918. Fairman, C.E. New or noteworthy Ascomycetes and lower fungi from New Mexico. Mycologia 10: 239-264. S 1918. New species are described in Diatrype (1), Didymella (2), A piosporella (1), Lep- tosphaeria (2), Gibberidia (1), Pyrenophora (1), Hendersonia (5), Microdiplodia (3), Phyllachora (1), Hysterium (1), Patellea (1), Phoma (2), Dothiorella (1), Placosphae- ria (1), Contothyrium (1), Ascochyta (1), Ascochytula (1), Stagonospora (1), Cryptostictis (1), Camarosporium (2), Arthrobotryum (1), and Rhabdospora (1). Fink, B. A new genus and species of the Collemaceae. Mycologia 10: 235-238. pl. 73. 25 § 1918. Collemodes Bachmanianum. Goodspeed, T. H., & Clausen, R. E. An apparatus for flower meas- : urement. Univ. Calif. Publ. Bot. 5: 435-437. pl. 54+f. 1. 25S 1918. SOHN Graves, E. W. My experiences with a fern garden. Am. Fern. Jour. 8: 71-76. S 1918. Greenman, J. M., & Pfeiffer, N. E. A new Selaginella from Mexico. Ann. Missouri Bot. Gard. §: 205-210. pl. rz, 12. 20S 1918. Selaginelia Landii sp. nov. INDEX TO AMERICAN BOTANICAL LITERATURE 33 Giissow, H. T. Drouth injury to McIntosh apple. Phytopathology 8: 490, 491. f. r. 12S 1918. Giissow, H. T. Observations on obscure potato troubles. Phyto- pathology 8: 491-495. f. 2-5. 12 S$ 1918. Harper, E. T. Hvypholoma aggregatum and H. delineatum. Myco- logia 10: 231-234. pl. r2. 255 1918. Harper, R. A. Binary fission and surface tension in the development of the colony in Volvox. Brooklyn Bot. Gard. Mem. 1: 154-166. pl. 2+ f. 1-4. 6 Jl 1918. Harper, R. A. Organization, reproduction and inheritance in Pedi- astrum. Proc. Am. Philos. Soc. 837: 375-439. pl. 5, 6+ f. 1-35. § 1918. Harris, J. A. Further studies on the interrelationship of morphologi- cal and physiological characters in seedlings of Phaseolus. Brook- lyn Bot. Gard. Mem. 1: 167-174. 6 Jl 1918. Harris, J. A. The interrelationship of the number of stamens and pistils in the flowers of Ficaria. Biol. Bull. 34: 7-17. 1918. Harshberger, J. W. American heaths and pine heaths. Brooklyn Bot. Gard. Mem. 1: 175-186. f. 1-8. 6 Jl 1918. Hollick, A. Some botanical problems that paleobotany has helped to solve. Brooklyn Bot. Gard. Mem. 1: 187-190. 6 Jl 1918. Hottes, A.C. Nature’s garden across southern Canada. Jour. Inter- nat. Gard. Club 2: 338-351. $1918. [Illust.] Howe, M.A. Algae. In Britton, N. L. The flora of the American Virgin Islands. Brooklyn Bot. Gard. Mem. 1: 116,117. 6 Ji 1918. Hubert, E. E. Fungi as contributory causes of windfall in the north- west. Jour. Forestry 16: 696-714. O 1918. Jeffrey, E. C. Evolution by hybridization. Brooklyn Bot. Gard. Mem. 1: 298-305. pl. 5. 6 Jl 1918. Jennings, O. E. An annotated list of the Pteridophytes of northwest- ern Ontario—II. Am. Fern Jour. 8: 76-88. S 1918. Kryshtofovich, A. N. On the Cretaceous age of the ‘Miocene Flora” of Sakhalin. Am. Jour. Sci. 46: 502-510. S 1918. Kunkel, L.O. A method of obtaining abundant sporulation in cultures of Macrosporium Solani E. & M. Brooklyn Bot. Gard. Mem. 1: 306-312. f. 1-4. 6 Jl 1918. Lankester, C. H. Lycastes in Costa Rica. Orchid Rev. 26: 181. Au 1918. 34 INDEX TO AMERICAN BOTANICAL LITERATURE Loeb, J. The law controlling the quantity of regeneration in the stem of Bryophyllum calycinum. “ses Gen. Physiol. 1: 81-96. f. I-5. S 1918 Machidane: J. M. Synchronism in plant structures. Brooklyn Bot. Gard. Mem. 1: 313-326. 6 Jl 1918. Marsh, C. D., Clawson, A. B., & Marsh, H. Larkspur or “poison weed.” U.S. Dept. Agr. Farm. Bull. 988: 1-15. f. 1-6. Jl 1918. Marsh, C. D., Clawson, A. B., & Marsh, H. Stagger grass ( Chro- sperma muscaetoxicum) as a poisonous plant. U.S. Dept. Agr. Bull. 710: 1-14. f. 1-8. 13S 1918. Maxon, W. R. Further notes on Pellaea. Am. Fern Jour. 8: 89-94. S 1918. Merrill, E. D. New plants from Sorsogon Province, Luzor, Philip. Jour. Sci. rz: (Bot.) 1-35. Ja 1916. Forty new species in various genera are described. Merrill, E. D. The systematic position of the “rain tree,” Pitheco- lobium Saman. Jour. Washington Acad. Sci. 6: 42-48. 19 Ja 1916. Metcalf, H. The problem of the imported plant disease as illustrated by the white pine blister rust. Brooklyn Bot. Gard. Mem. 1: 327~ 333. pl. 6,7. 6 Jl 1918 Moore, G. T. Eras notes—III. A wood-destroying alga, Go- montia lignicola, n. sp. Ann. Missouri Bot. Gard. 5: 211-222. pl. £S*78. : AC A IOTR, Mottet, S. Les Garrya. Rev. Hort. 90: 152. 16S 1918. [Illust.] Murrill, W. A. The rosy-spored agarics of North America. Brooklyn Bot. Gard. Mem. 1: 334-336. 6 Jl 1918. Myers, C. E. Study of the inheritance of size and productiveness in pedigreed strains of tomatoes. Proc. Soc. Hort. Sci. 1914: 26-33. Mr 1915. Nieuwland, J. A. “Fairy circles.” Am. Mid. Nat. 5: 230, 231. -$ 1918. Nieuwland, J. A. Heterothrix (B. L. Robins.) Rydb. a synonym, and other notes. Am. Mid. Nat. 5: 224, 225. S$ 1918 Nieuwland, J. A. Teratological notes. Am. Mid. ‘Nee Sr oaar, S 1918. ; Olive, E. W. The cytological structure of Botryorhiza he eto Brooklyn Bot. Gard. Mem. 1: 337-341. fl. 8 6fir Osterhout, W. J. V. A method of oo EY fol os Physiol. 1: be Bt i... S tds INDEX TO AMERICAN BOTANICAL LITERATURE 35 Osterhout, W. J. V. The nucleus as a center of oxidation. Brooklyn Bot. Gard. Mem. 1: 342-347. 6 Jl 1918. Osterhout, W. J. V., & Haas, A. R. C. On the dynamics of photo- synthesis. Jour. Gen. Physiol. 1: 1-16. f. 1-5. S 1918. Packard, C. Difference in the action of radium on green plants in the presence and absence of light. Jour. Gen. Physiol. 1: 37, 38. 1918. Pearson, G. A. The relation between spring precipitation and height growth of western yellow-pine saplings in Arizona, Jour. Forestry 16: 677-689. f. 1-3. O 1918. Peltier, G. L.. & Neal, D. C. Overwintering of the Citrus-canker organism in the bark tissue of hardy Citrus hybrids. Jour. Agr. Research 14: 523, 524. pl. 58. 9 S 1918. Pierce, R.G. Additional list of state and national quarantines against the white pine blister rust. Phytopathology 8: 484-486. 12S 1918. Piper, C. V., & Beattie, R. K. Floraof the northwest coast. i-iv + 1-418. Lancaster. 1915. f Piper, C. V., & Beattie, R. K. Flora of southeastern Washington and adjacent Idaho. i-xi+ 1-296. Lancaster. 1914. Pring, G. H. Cycads. Jour. Internat. Gard. Club 2: 365-377. S 1918. [Illust.] Rathbun, A. E. The fungous flora of pine seed beds. Phytopathol- ogy 8: 469-483. 12S 1918. Reed, G. M. Physiological specialization of parasitic fungi. Brook- - lyn Bot. Gard. Mem. 1: 348-409. 6 Jl 1918. Reeves, F.S. An investigation in tomato breeding. Proc. Soc. Hort. Sci. 1914: 24-26. Mr 1915. Riddle, L. W. Lichens of St. Thomas. In Britton, N. L. The flora of the American Virgin Islands. Brooklyn Bot. Gard. Mem. 1: 109-15. J. 3: .G JL 1918. Opegrapha acicularis, Lecania euthallina, and Blastenia nigrocincta, spp. nov., are described. Rolfe, R. A. Gongora latisepala. Curt. Bot. Mag. IV. 14: pl. 8766. S 1918. A plant from Colombia. Rolfe, R. A. Govenia tingens. Curt. Bot. Mag. IV. 14: pl. 8768. S 1918. A plant from Peru. ‘ [Rolfe, R. A.] Orchids of Panama. Orchid Rev. 26: 179-181. Au 1918. 2 4 36 INDEX TO AMERICAN BOTANICAL LITERATURE Rowlee, W. W. Relation of marl ponds and peat bogs. Brooklyn Bot. Gard. Mem. 1: 410-414. f. 1-3. 6 Jl 1918. Rumbold, C. Laboratory notes on cultures of Endothia parasitica A. & A. Notes on the color reactions of reproductive and vege- tative hyphae of E.. parasitica when treated with chemicals. Phy- topathology 8: 495-499. f. 6. 125 1918. Shannel, A. D., Scott, L. B., & Pomeroy, C.S. Citrus fruit improve- ment: a study of bud variation in the Marsh grapefruit. U.S. Dept. Agr. Bull. 697: 1-112. pl. I-11 +f. I-14. 27S 1918. Shear, C. L. Pathological problems in the distribution of perishable ~ plant products. Brooklyn Bot. Gard. Mem. 1: 415-422. pl. Q-II. 6 Jl 1918. Shreve, E. B. Investigations on the imbibition of water by gelatine. Science I]. 48: 324-327. 27 5 1918. Shreve, F. The Jamaican filmy ferns. Am. Fern Jour. 8: 65-74. pl. 4.> Sy 1618: Shull, G. H. The duplication of a leaf-lobe factor in the shepherd’s- purse. Brooklyn Bot. Gard. Mem 1: 427-443. f. 1-4. 6 Jl 1918. Shufeldt, R. W. Grasses, sedges and some September flowers. Am. Forestry 24: 551-555. f. 1-8. S 1918. Sinnott, E. W. Isolation and specific change. Brooklyn Bot. Gard. Mem. 1: 444-447. 6 Jl 1918. Skottsberg, C. Botanische Ergebnisse der schwedischen Expedition nach Patagonien und dem Feuerlande 1907—-1909.—V. Die Vege- tationsverhaltnisse langs der Cordillera de los Andes S. von 41° S. Br. Kungl. Svensk. Vetenskapsakad. Handl. 56: 1-366. pl. 1-23 + f. I-24. 1916. ‘ Includes the new genera Pycnophyllopsis and Xerodraba and 31 new species in various genera. Smith, E. F. Frank N. Meyer. Science II. 48: 335, 336. 4 O 1918. Smith, E. F. The relations of crown-gall to other overgrowths in plants. Brooklyn Bot. Gard. Mem. 1: 448-453. 6 Jl 1918. Stewart, F. C. Tubers within tubers of Solanum tuberosum. Brook- lyn Bot. Gard. Mem. 1: 423-426. f. 1-3. 6 Jl 1918. Stone, G. E. Contact stimulation. Brooklyn Bot. Gard. Mem. 1: 454-479. f. I-4. 6 Jl 1918. Taylor, N. A quantitative study of Raunkiaer’s growth-forms as il- lustrated by the 400 commonest species of Long Island, N. Y. Brooklyn Bot. Gard. Mem. 1: 486-491. 6 Jl 1918. BULL. TORREY CLUB VOLUME 46, PLATE I STEINBERG: GROWTH OF ASPERGILLUS NIGER BULL. TORREY CLUB ° es ae \ eine > ANN A PESTER 5 SY) ART Me LS? Baal) eee cL S. 8, a ie ca Se, ae, SY We 7 9. oa > an “I> CA ) \, VOLUME 46, PLATE 2 BROWN: APOGAMY IN CAMPTOSORUS RHIZOPHYLLUS Vol. 46 No. 2 BULLETIN OF THE TORREY BOTANICAL CLUB FEBRUARY, I919 Xerophytic grasslands at different altitudes in Colorado FRANCIS RAMALEY (WITH TWO TEXT FIGURES) The first comprehensive study of xerophytic grassland in Colorado was reported by Shantz (16) for the mesas at Colorado Springs. Later the same author (17, 18) published articles dealing with grasslands of the plains. Important work in the Pikes Peak region has been done by Clements (1, 2) and by Schneider (15). During the past few years the present writer (6, 7, 8, 9, 10, 11, 12) has published a number of papers dealing with dry grassland of the montane zone at Tolland, Colorado. In a contribution devoted chiefly to successions from marsh vegetation Robbins (14) also has given some points on dry grassland at Tolland. A rather brief but very clear and definite account by Vestal (20) of foothill grasslands, chiefly near Boulder, is included in a survey by him of the various associations of the foothill zone. An earlier paper by the same author (19) characterized the grasslands of the plains in the same locality. Fuller (3) has recently made a comparison of the dry grassland at Tolland with black soil prairie in Illinois. Up to the present time there is no printed comparison of xerophytic grasslands at different altitudes in Colorado. Since - these communities occur in large or small stretches all the way from the plains to alpine heights it is evident that very decided ecological and floristic differences must exist. These differences — it is the aim of the present paper to report. Observations have been made on some of the areas for ten seasons. Certain localities have, however, been visited only a few times. During the Pprog- [The BULLETIN for January (46: 1-36. pl. 1, 2) was issued January 20, 1919.1 a7 38 RAMALEY: XEROPHYTIC GRASSLANDS AT ress of the work a large collection has been accumulated which is kept as a special ‘dry grassland herbarium.’”’ Air temperature records have been kept for the growing season at Tolland (8,889 ft.) and these have been compared with reports from the govern- ment stations at Boulder (5,340 ft.) and at Corona (11,660 ft.). Soil temperatures have been carefully studied at Boulder and at Tolland, while occasional records have been made in the higher altitudes. The present work does not in any sense cover the whole Pe ae ak ‘ oan ae oe oe Fic. 1. sii of Colorado showing general oe features. The Conti- nental Divid zig-zag course north and so he foothills at the moun- tain front are indicated by the series of eat age Cae and Tolland are to the northwest of Denver. of Colorado (Fic. 1) but merely the area tributary to the Uni- versity at Boulder and the Mountain Laboratory at Tolland. Field work has been done chiefly in a rectangle about 40 miles east and west and 30 miles north and south with altitudes from 5,000 to 14,000 feet above sea level (Fic. 2). In the following synopsis of xerophytic grasslands, descriptions are omitted because these are available in the writings of Shantz, Vestal, Pound and Clements (5), and the writer. Nichols’s (4) exposition of ecological concepts has been followed in choosing names for the different communities. The nomenclature of species is that of Rydberg (13). ; DIFFERENT ALTITUDES IN COLORADO 39 SYNOPSIS OF XEROPHYTIC GRASSLANDS IN NORTHERN CoLoRAbo! A. Associations of mixed character belonging to the plains region; not clearly dominated by grasses or sedges: 1. Gutierrezia-Artemisia Association. 2. Plains Ruderal Association. 3. Chrysothamnus Association. 4. Artemisia ‘filifolia Consociation (of the Sand Hills Mixed Association). B. Associations dominated by sedges; mountain communities not represented on the plains (Ramaley, 11): 5. Carex stenophylla Grassland Association ; foothills and montane. 6. Carex Rossii Grassland Association; montane and subalpine. 7. Carex siccata Grassland Association; montane and subalpine. 8. Carex elynoides Grassland Association; subalpine and alpine. C. Associations of the Mixed Dry Grassland Type: 9. Inceptive Dry Grassland Association of the Foothill Zone (the Foothills Primitive Grassland of Vestal, 20). 10. Derivative Dry Grassland Association of the Foothill Zone (the Foothills Mixed Grassland of Vestal, 20). 11. Inceptive Dry Grassland Association of the Montane Zone. 12. Derivative Dry Grassland Association of the Montane Zone. D. Associations dominated by one or a few species of grasses: 13. Short Grass Association; plains and mountain front; domi- nated by Bouteloua and Bulbilis. 14. Wheat Grass Association, mountain front chiefly; dominated by Agropyron Smith. . Andropogon Bunch Grass Association; plains and mountain front. 16. Porcupine Grass Association; plains and foothills, occasionally montane; dominated by Stipa 17. Sand Hills Mixed Association; plains; dominated by Cala- movilfa and Andropogon. 1It is often wean to decide just what communities should be included as “grassland.” The writer omits lichen and mat associations, since they hove almost i on Saas these are best known as ‘“‘scrub.” The Prairie Grass Association (of Vestal) is not included, as it is mesophytic. 40 RAMALEY: XEROPHYTIC GRASSLANDS AT 18. Festuca Bunch Grass Association; montane; dominated by tall cespitose species of Festuca. 19. Muhlenbergia-Danthonia Consociation (of the Derivative Dry Grassland Association of the Montane Zone). | Platieville # seer Oe ro, sere, - 2. Map of that part of Colorado readily studied botanically from the University of Colorado at Boulder and from the Mountain Laboratory at Tolland. Contour lines are shown for 5,000 ft., 6,000 ft., 8,500 ft., and 11,000 ft. The moun- tain front is at about 6,000 ft. THE Mixep Dry GRASSLAND TYPE The term ‘‘Mixed Dry Grassland,” as used by the writer, in- cludes all mountain grasslands of mixed character, growing gener- ally in coarse-grained soil. In the preceding synopsis the com- munities numbered from 8 to 12 inclusive belong here and also Nos. 1 and 19. Early stages of the Short Grass Association, No. 13, are also of the Mixed Dry Grassland type. The Porcupine Grass Association, No. 16, when it occurs in montane situations may resemble ordinary montane dry grassland, and the Festuca Bunch Grass Association, No. 18, is, in places, not so very dif- ferent. Indeed, any of the grassland associations of the mountains may belong to this general type. While there is no difficulty in recognizing inceptive and deriv- ative stages of the various dry grasslands the species concerned DIFFERENT ALTITUDES IN COLORADO 41 are much the same. Thus, in the montane area, practically all the species present in the early stages persist throughout; the relative proportions change, however, and more species enter the association as the soil becomes finer grained and as humus accu- mulates. Occurrence of these grasslands is wide spread. In the lower part of the foothill region typical vegetation is a coniferous savanna. In the interspaces between trees there is dry grassland. Many intervales, locally known as ‘‘parks,’’ are almost entirely without trees. In both foothill and montane areas many stream terraces, alluvial fans, and south-facing hill slopes support dry grassland. In the subalpine zone conditions for forest develop- ment are more favorable, and grassland is likely to be confined to burned areas or to wind-swept south exposures. ENVIRONMENTAL INFLUENCES The stations where most of the collections were made, with their altitudes, are as follows: Boulder, 5,340 ft.; mountain front, mesa area. Crescent, 7,457 ft.; lower foothills. Rollinsville, 8,367 ft.; upper foothills. Sulphide, 8,508 ft.; upper foothills. Smartweed Lake, 8,420 ft.; upper foothills. Tolland, 8,889 ft.; montane. Bryan Mountain, 11,000 ft.; subalpine. Boulder, Crescent, Sulphide and Bryan Mountain are in Boul- der County; Rollinsville, Smartweed Lake and Tolland are in Gil- pin County (see Fic. 2). Many other stations through both of these counties have been visited. The soil of all the dry grasslands is disintegrated rock, rather coarse grained, often with pebbles and boulders. It readily permits penetration of rain water but it dries quickly. As shown in a previous paper (10) the water content is low, averaging about 7 per cent, but the wilting coefficient is correspondingly low, about 5; figures being for 3 dm. depth. The soils warm up quickly through direct insolation and show a higher temperature than the air. Floristic differences in the dry grasslands do not depend on soil quality for this is much the same in all, but rather on air temperature (TABLE I), soil temperature (TaBLE II), and pre- 42 RAMALEY: XEROPHYTIC GRASSLANDS AT TABLE I MEAN TEMPERATURES IN DEGREES FAHRENHEIT. THOSE AT TOLLAND AND BRYAN MOUNTAIN ARE ESTIMATED, EXCEPT FOR JULY AND AUGUST AT TOLLAND Boulder 5,340 ft. Tolland 8,889 ft. | Bryan Mt. 11,000 ft. MOY. eta i es 56 } 42 35 BAG: Ree ios nee Nie Uae gie 65 | 51 44 BY iG oe at ee i 70 57 50 ANCNM Eyes oA ae es 70 | Sk 48 Septeniber ny ests eee: 64 } 46 39 TABLE II SOIL TEMPERATURES IN DEGREES F AHRENHEIT OF DRY GRASSLAND AT THREE TYPICAL ALTITUDES; NUMEROUS OBSERVATIONS AT 3 DM. DEPTH Boulder 5,340 ft. Tolland 8,889 ft. | Bryan Mt. 11,000 ft. Mayer a a ie oe 55 FUNC. er eel Ran 64 54 PS sy a oe 72 64 58 AUBUSt Ce Poo ee 70 62 56 pepteiber G8 os Oe, sy 64 58 ; Sa Note.—Since readings of soil temperature were taken in the daytime they are somewhat above the true average. Data for Bryan Mountain are rather fragmen- tary and figures may have to be revised later. Probably the May soil temperature at Bryan Mountain is very low, close to freezing in the early part of the month. cipitation. The mean annual rainfall at Boulder is 18 inches, at Tolland probably 28 inches, and at Bryan Mountain it can safely be estimated at 38 inches. April is the wettest month at all the sta- tions. At the higher altitudes the greater rainfall and cooler tem- perature make the grasslands more mesophytic than those of plains and foothills. SYSTEMATIC LIST OF SPECIES A list of the species characteristic of xerophytic grasslands at various elevations is given below. Only spermatophytes and pteridophytes are recorded; lower plants, aside from parasitic fungi, are almost entirely unrepresented in dry grassland. It is of importance to note that in montane and subalpine situations, due to the more congenial climatic features mentioned above, a num- ber of species, which in the lowlands are confined to moister soil, may be regular components of dry grassland. This distinction is brought out in the list: the letter x signifies that the species is present in dry grassland at that altitude; the letter 0 means that the species is found in moister places rather than in dry grassland at the altitude indicated. The letter s means that the species occurs at the altitude named, but only in special situations (where warm or protected) and is not a true component of dry grassland. DIFFERENT ALTITUDES IN COLORADO 43 Sub- | Mon- | Upper | Lower — alpine | tane | foothill | foothill | front Selaginellaceae | a GSS See ees Padre’ x | x igi alee a Avian bi MOPUME Se. SCUIA Ue esp ees ch - x x - - EL pet oO Nein Vespamerssiy sorority eos = x x x DSCUdOT OPENS os ie ea Ae a ~ * x oO o oy: SCHOPUSONE 5 he eee ees es _ x x x s e EVEOMENS Se oe a ees OC Ses Se - = - ne SMUNIES G5 er a - $ x i * wy sss ae wa FOG Oar oie eee. ~ - x x x (ede Pa eee ee Chee - x x o te) Anbropobon "Halli. ete ie hae iy weak oe — = = ce x tip leaded Bags: ee ayo Nee - = = _ x Aristida tit G Pen re: Prey nw oes eas Aus - - = x < - Avena REE is ices CR eae 9 he eee - x - - - MM OTtOntGhG ok ee RPE Canes x - = -—" = uacpharcaias on frieholepie!. re ae ie oe estou x x - - Bouteloua curtipe ihe ek ae a eee - - - * x aaa 26h eae Ace Popes Peeters: - _- x x x i harsh os Pe 5 as es - — x x x Bromus irisecormis (introduced) See ish oe - = i x mm pelligwus. 05.5850 3 Ue hs es = x x _ = Bulbilis dusstendas TERN PN Mote iuaig eee a Ta ere - - - - x Calamagrostis gate Soca Sore Die tek ere ai * x x ~ al Danthonia interme Ese San rasta dca’ Weel < x - - ois TSE. CURA Gas ot a cus ee x x aed Eas Elymus PIOUS SOS ie Here pe ee = _ = CoS Erioco iin bemenvides:: uh OG ea Diss ae alee ec = “ta "as ee Festuca brachyphylla..........++++: eran a ata. 4 - - fe ines yg 2, ee emer a pee RAO as Ne ees x x = SS RIMMISPOTE: os Peete he eee x x = 9 fe e Pr ae be ide Aes & es ~ - - = x MEE i ce Serer aa. iene ad ence x x - - ‘2 | $G¢sMORIENE. OS. oi pug Rr tees sana FORM eel oe eas wee x + x Cer stium PCCHIR PION 5 So ee ae Been - - Peet. te BAEK POdUM 3 oy Cea et - - _ x 4 oa CRINDESIFE ao a es - ~ x x x ie SCI Oe oe Pe ed eee x x - - Caryophyllaceae Silene EDM 5 hae Sas 6 oR ne Oe et ea: # - = ee EGU ia OS See Na ae ate or oy - x x - - Ranunculaceae Delphinium -Seeal on yea oh a eels - - - eae Clie Vos anes wae ree Cane cn - - - - x Pulsatilla lndiatotane te Bye ear et eee Le o 0 Fe ae Papaveraceae | Argemone intermedia... o.oo ee ee es ie i Picci ae Brassicaceae Chetrintaiaspera. 280 oo, Sa SS Fire eons = eo x x = DOPVIMNE os oie ae et Po iceeines - =_ seh x x ae econ iam PES Oe un ara Bae x % x - COLE oat rae OA bees 0a - x x x - Draba coloradensis SAIS Tite pep Sl Aten de erence - _ on x x PEGI GCHSELOTUIN «ons 6 CE Ree ees — ~ x = x aes montana Sia opi oRoy ae eae euece wore a x x x Pivades JRORRRG OO a a ae - “ x x x T Ria ht COIGFAOEASE ooo eae oe ss x a x x x o r SCENES Cia eee x. x - - - : Crassulaceae Sedum steno pelaline. 64 ost i os % # x x x Saxifragace: Micranthes rhomboidea. . 0... 6. ce ee es x < x o oO Rosaceae Drymocallis sco ERD Bree ig Ny aaa Fast the = x x x x Potentilla COMIN A a ER a ee vies okie [ x x ae a ' MER nL oe ae ot Oe es Set - x x x x i H EPPIAKG O08 SS Be ee Oe ~ x x x £ a Sirigcse i ie ee oe eee - x x x x Arogeilas al albitoru CS sos cae eee - oe x x x She games an x x va gy c Lamberti Ai RR este OL ae c iiuis Soap - x x x x Gaara Gace See Seas eae ae x x bg ate Astragalus jae oct hee Op aes ~ x x = = SUCCHIONINMGE Cok 6 os fe ees ~ ome a x * Heal OES a Re eee eer ge re ie _ x x x x Orophaca tridactylica...........--- cuca. Mines - ge ~ x x Petalostemon oligophyllus.......02. 0000-2055 — ae a = i PUPDUICNS oe ee ie Va ee ee nas = a ae Psorvates Grg0PRgla 65. a ee a Ss jae ee / sy in x Psoralea tenuiflora........ Pe ee ae ve 2 . 46 RAMALEY: XEROPHYTIC GRASSLANDS AT sear ar ee Sub | Mom. | tipper | Lowen alpine | tane | foothill | foothill) front Fabaceae—continued 1 93tt: Drurenondid ns 8 oes Seo eis 2 ead ss ors aE (eer — x x Xylophacos Pabres Wiveg ace mae Sure LeU, tae vas Se eae. ee - - erent re a ar ee vos - } = fey} eae x sd SHOPEMGMM SR 2 eee oe — - x x x Geraniaceae | Geranium Fremon 7) lt OER SNAP eT See ak ee apa Baar ak ies ea 3 x | = x POTTY eS GEST i ie Se ees ws - | = cA BE x Linaceae | Lent LOWEST a a cc. eee —_ * < x x Euphorbiaceae | LUAVMAIUS TOOMSINS oy BV Cee eee. et ads © x x & Malvaceae | Sphaeralced cocesneg) os ors eae ee ~ | — - % x Violaceae | Viele Natalie Oe ee a = ~ - x x Cory phantha pcksecnarddeests ah ee es ee - | ~- x x x POORO SO oo LO eae ia ot ee . x & Opuntia fragilis Gate ae es: Cid eae ip _ x x acantha. <<)... ; POT ESR PO ae te me - 3 x ee | Goward coccinea co ee Peat egeecne ee — |= - bs * te MU ae ee a ee et - | = _ a = Mertolix servrulata 8G ae i ee - - - x < Am eae Pleas OCOHE Sore or ke on ee pane — = az = Cog. swellia macrocar po Ba iy eno. Sa UN - | - - x ~x Meer ei. Geek ee bee ee —- |} = - x x Harbour trachea ey oes a a ee — [oA x L& x OE. ESOGVICOIMIN 8 Ss OE Sis - ae - x % pacar jhe uae ewes pay ee ~ x = Alec OD PRR GS CO Soe Oe a ae eee x jo - — ~ Pseudocymopterus tenutfolius ... 0.0.0.0. 0... - | x x - rimulaceae ANGPOSECE MENUDO ke ee ces x % = ; CHIPWIGHS 5 ec ee sae hs ~ x x = shbumbellna co. sos i ia a a. ~ ntianaceae Dasystephana Bigelovis. 3 os. 6 4 - _ x x at Tessaranthium oe peek are ae at mar ec.” x x oO pia Acerates snus neg eee OE ue eee - = - ~ x OP ne Es Rd ioe al SUM - - - * Adare ps pieneriig Se RE RO PEs Oe Sarena! ~ - 3 tem: Colloerta ikedess ooo ooo ree a - ~ x x ° a Me eae We Pa ee PEA rt - x x = Co PPNOME So ee ay Co eee aware ~ ~ a x x Phlox mulbifiors.- wR SMe ge wie Wind Ueacg e aes ~ eke eo te fey ea RSs ou mS orate “ - -_ -—- _ DIFFERENT ALTITUDES IN COLORADO 47 j Peacmenipapeiaaals Sub- | Mon- | Upper | Lower eng alpine | tane | foothill | foothill | gont Hydrophyllaceae Phacelia gor ag We 222 i ee ee ee - x * x x Sey ON Cee es er ee x o - - = Boragi inaceae M ertensia — SO deg hoc cee aa aed Ng cae ee - ¢ - - - BAER es Cr tes aa eS x - - - ~ = lanceoiat a ea ei i ant yea atte ee are - = x x x Ne TULCPEIOr a i oe is ee Be a - x x - - es rennet PRN. RENCE S ety eee - - x ~ — te a ie Pe neal eaten Sew oes x _ - - - CCGCGE VG OP GOlE oo ia ee phen, tae ee ies - s 5 x x Onosmodium eeibaiaa RS SEAS a eG Glangende ste ae — = = —_ x Verbenaceae V FRC PV ACIBOSD socio 6s ees ake CE - = re ne: x Lamiaceae Fi cdeoiie MASPias ors eae he a Ge ee eS oA las 564 = os x Monorda pectimata.. (oti es 7% Sigs ake epee ss a ag x hul Castilleja cechientans: Cesk 6 Ree os x - - = ap SESSELEAOPE ik Od Ok Hoe ca k 2a ms OF - ~ - oe oe Urihocar bus tiles. oo ies eS ae eer es - x x pa fe PCnislemor OAS oo Nein Oke a ne ts o> - a a x x Pla taginaceae Siies PIS. 2 eee Ores Cree os eo - — x x x Rubiacea CraLfene DETERS eGo, BN Cs ee ee = x x o o Santalaceae Comindta Panda oop ieee oi es - x x & x Campanulaceae COM POwUld DesOdeS ee oe eee ae x x Ps o 0 Ambrosiaceae AMOTOSIG PSUOSTGCRVG © oo i co sn ied bik ee s 5 x cs pened Achillea lamulosa oreo ee a a es = x x x o o Antennaria ana phaloides Pere Gees ape Suni ee ces x oe x = ey Vaees Wert ar ec ot Cahora Mat < x x x x a: ROS Cs Poh pe Ata hosts a - x x x - Bs ee ose - ~ = _ x ie WICIOPIVNG. oa a ss - x x x x Artemisia COMOROS eee ee - x x x x PSO i ee eee in es oS - - x x x 6 Farmoddl 0 oe ee ee eee x E x : PSNR os pink oy ts acee ewe aes ite ~ x # x : ae , or Joop Ce eerie ree ~ x x co x : Palesone fs soe eR Eee + - - * Y Aster phan De a eas a _ er on we x Pore oo ease i ee oe: — s x x Chrysopsis asprelok Piet yey ees eh ea pee Loe x x x x foliosa .... roe ia LT a DT ee dttit Ge Ceaieal * The species of crear wrsegne? one Tye of separation and it is siti that what the writer h nd C. hispida should be combined, also that te ted a Jove and ln ae ingle. 48 RAMALEY: XEROPHYTIC GRASSLANDS AT Sub- | Mon- | Upper | Lower | — A alpine | tane | foothill | foothill | front Carduaceae—continued Chryso ae ee Nearer a regane ait seach ib a ~ - x f AMER oe. Wispida eo one e oe es panels grit gp AS NORE one Ge aN hes a aes Bie ae eae x Pade a Cirsium Bias ok by ein oe ai ae a CEE Sa a ree Dane bo = es | x | # COFGODSES FERCLOVE Cis PS ose te wars - | = ee CEN he PV ERPPOM COMES. C08 os hey Ca ek ns Med oe | - | - ae | ze x 3 SMS acca ese ees corel Se ene tes aia ag ‘ ME ed eae rs ee ae pete aes x | x x : (il AO EMUPES CPx Os ce eee En wale fence x pees eae x Goslerdia areas oe ee ees { o— x x o o Grindelia ryt te ee AS | mf oe x - | = = BSG ss Uae Oe ea aeey peeae / - | - en laas © eae e SOUGIIOSG i Eee es —_ ~ Bemba. EDGE ae OLE is eee ae atte _ / x pee Oe there NN niliertesia: Saran oc. s fic sb es os ies ES - | = - | s x Eiéhamthas Pevolanis 3 oo 5 es Sad eid eae _ x | ie Hymeno peers: es Beg rere Sele i gaits a es = x x Kuhnia glutino Boers er erneeeaite semen yt Me renee sin aes - ~ x Laciniaria paced Ge onl Da esate yioe eu ate - | = - “ x M achaeranthera koe Sia ee ee ae nbd ods - % x sng Hey any eee 9 we ghey x 3 a dries oe kW Ste es en ee satu Mele” x nent Ratibida iucgae Pe ee OPE ED ae SS ee - oe Senecio oblanceotaius 9 25 ea ee) - | = | x | Me 3 ~ WOME SOL ed ee eae ee as es oie ie x gee epee fs POMS 6 ioe ana ee eee rT be eehils come - x OME, 0 tens ate Pig es ee - x x = WEREPIROPNIRE i de Oca Ae 0 = - Sideranthus ef Se Scenes Pes Ru ee alee. - - | x Solidago CONCIRNG Ss es Vs oe ee fo x x = - . deck ca ce ia Cie Oy eas x x - - - MISSONTIER HS 6S Phe cae ee ~ = x 5 MORE OS Ores ee ee - ~ - x x er OFEOPRYNG 22 8 os CEE een bos o a - Tetraneuris Pict VEG Te ge Oe ee tas [ - - x alt ced ea Pees bodes) —-l = LowestnS PYSMAEIE ee es ee ~ | = FOWRSONMIE CX8CODO Se os ee fame a - x x rs BFONOUIORG oe a ee - | ~ & ee ee Cichoriaceae A goseris eet ies CDE oe Oe bern et eee ae x x % pe ame ar wales Ge eee x x o eons juncea rg eect age gr — |o- | - — x FLORISTIC DIFFERENCES OF DRY GRASSLANDS AT DIFFERENT ALTITUDES The foregoing systematic list gives in brief form the chief facts of altitudinal distribution of the various species in the dry grasslands. At the mountain front, i. e., on the mesas and mesa terraces, there are 160 species. In the ok foothills ite muimber DIFFERENT ALTITUDES IN COLORADO 49 is reduced to 139, in the higher foothills to 130, in the montane area to 107, and in the subalpine area to 50. It will be seen that there is a reduction in species roughly corresponding to increase of altitude. But the change from the montane dry grassland (at Tolland) to the subalpine dry grassland (at Bryan Mountain) is very abrupt. It is sometimes difficult to decide whether a given species is to be included as properly belonging to a certain grassland. The attempt has been to exclude stragglers and accidental members, in other words to keep the lists reasonably small. While the systematic list ,tells what particular species are found at the various stations it gives little idea of the true ap- pearance of each grassland area. At the mountain front, as already noted, there is a great variety of plants. The average height of the vegetation is taller than elsewhere; many of the grasses and other plants are 3-6 dm. tall, and some are taller. Certain species are conspicuous which either do not occur at all _ higher up or else are infrequent, as, for example, Agropyron spi- catum, the Andropogons, Bromus brizaeformis; Yucca glauca, the Delphiniums, the Petalostemons, Tium Drummondii, the various Cactaceae, the Gauras, Helianthus petiolaris, Laciniaria punctata, and Rattbida columnifera. At Tolland the number of species is still ie but a certain few are especially abundant and dominate wide areas, as, for example, the following: Mertensia Bakeri, which flowers in May; Aragallus Lambertii, which flowers in early July; Sedum stenope- talum, in mid-July; Campanula petiolata and Orthocarpus luteus, in August. The average height of vegetation in midsummer is 2-3 dm. Grasses and sedges are abundant, especially where the soil has considerable humus. Here the dry grassland becomes what the writer calls the Muhlenbergia-Danthonia Consociation (of the Derivative Dry Grassland Association of the Montane Zone.) ‘The dry grassland at Bryan Mountain (Carex elynoides Associ- ation) has a rather striking appearance because of the tufted masses of the dominant plant. This is, however, of low growth and the association does not look at all like the bunch grass asso- ciation of the plains. Conditions are somewhat mesophytic. on account of low temperature and frequent showers, so that many 50 RAMALEY: XEROPHYTIC GRASSLANDS AT of the plants recorded as belonging to the dry grassland would also be included in a list of mesophytes. One-third of the species are grasses. The systematic list includes a total of 256 species in 131 genera and 35 families. The families best represented with the number of genera and species in each are shown below: Poaceae! iso eer. ae 25 66 UB ALB A ceca sok cS te & COAG oe a3 SA's, ics Pabacese 552. hse 9 16 Polygonaceae.......... ye : Aminiaceae.... 2.00.65. eee fUsinaceae. 0. fo ss eee es Boraginaceae.......... =, ae | Brassicaceae........... 6: 10 Carduacene «2225 iS: 23 55 Only 5 species range the whole distance from mountain front to subalpine zone. Of the 50 species at Bryan Mountain 29 extend down to Tolland and only 5 to the mountain front at Boulder. Of the 160 species at Boulder 40 range up to Tolland and 5 continue to Bryan Mountain. Floristically, then, the dry grasslands at Boulder and at Bryan Mountain are almost totally different, while a very considerable similarity exists between those of Boulder and Tolland and of Tolland and Bryan Mountain. SUMMARY The foregoing paper is based upon a study of grasslands in northern Colorado from the plains to the subalpine zone. A list is presented of all recognized xerophytic grassland communities in northern Colorado. From this list the author selects for com- parison those of the ‘mixed dry grassland type.” Environmental influences, the soil, temperature and rainfall are considered; the very low air and soil temperatures of the subalpine region being noted. To show floristic changes with altitude a systematic list of dry grassland plants is printed and the altitudinal distribution of each species indicated. At the mountain front there are 160 species in the dry grassland, while the numbers at other stations are as follows: lower foothills 139, upper foothills 130, montane area 107, subalpine area 50. Of the last named nearly one-half are strictly high-altitude forms which do not extend down even to the montane zone. It is pointed out that a number of species belonging to moister situations ‘in the lowlands are able to enter dry grasslands of higher altitudes because of lower temperature DIFFERENT ALTITUDES IN COLORADO 51 and greater rainfall there. A brief statement is made of differ- ences in the grasslands not at once apparent from the systematic list. Floristically the dry grasslands of the mountain front and of the subalpine zone are almost totally different, but considerable similarily exists between those of the mountain front and of the montane zone. UNIVERSITY OF COLORADO, OULDER, COLORADO. LITERATURE CITED Clements, Frederic E. Formation and_ succession herbaria. Univ. Nebraska Stud. 4: 329-355. 1904. ————. Research methods in ecology. Lincoln. 1905. Fuller, G. D. A comparison of certain Rocky Mountain grassland with the prairie of Illinois. Trans. Illinois Acad. Sci. 8: 1-10. | . 5 ath Nichols, George E. The interpretation and application of certain terms and concepts in the ecological classification of plant com- munities. Plant World 20: 305-319, 341-353. 1917. . Pound, Roscoe, & Clements, Frederic E. Phytogeography of Nebraska. 2d edition. Lincoln. 1900. . Ramaley, Francis. Botany of northeastern Larimer County, Colorado. Univ. Colorado Stud. 5: 119-131. f. 1-10. 1908. 7- —————.. The amount of bare ground in some mountain grass- lands. Bot. Gaz. 57: 526-528. 1914 8. —————. The relative importance of different species in a moun- tain grassland. Bot. Gaz. 60: 154-157. I9I5. 9. —————. Quadrat studies in a mountain grassland. Bot. Gaz., 62: 70-74. 1916. 10. ——————. Dry grassland of a high mountain park in northern Colorado. Plant World 19: 249-270. f. 1-6. 1916. 11. —————.._ The role of sedges in some Colorado plant communities. Amer. Jour. Bot. 6. 1919. Not yet published. 12. Ramaley, Francis, & Elder, Mary Esther. The grass-flora of Tol- land, Colorado, and vicinity. Univ. Colorado Stud. 9: 121-141. PO fe HE nS 13. Rydberg, P. A. Flora of the Rocky Mountains and adjacent plains. New York. 1917. 14. Robbins, W. W. Succession of vegetation in Boulder Park, Colorado. Bot. Gaz. 65: 493-525. f. I-14. 1918. = on on -_ 9. RAMALEY: XEROPHYTIC GRASSLANDS IN COLORADO . Schneider, Edward C. The succession of plant life on gravel slides in the vicinity of Pike’s Peak. Colorado Coll. Publ. Science Ser. 12: 289-311. f. 1-6. I9g11. . Shantz, Homer LeRoy. A study of the vegetation of the mesa region east of Pike’s Peak. Bot. Gaz. 42: 16-47, 179-207. 1906. ——_———. Natural vegetation as an indicator of the capabilities of land for crop production in the Great Plains area. U. S- Dept. Agr. Bur. Pl. Ind. Bull. 201. - ¥Orr. ————. Plant succession on abandoned roads in eastern Col- orado. Jour. Ecol. 5: 19-42. f. 1-23. 10917. Vestal, Arthur G. Prairie vegetation of a mountain-front area in Colorado. Bot. Gaz. 58: 377-400. f. 1-9. 1914 : Foothills vegetation in the Colorado front range. Bot. Gaz. 64: 353-385. f. 1-8. 1917. Additions to the flora of Colorado GEO. E. OSTERHOUT 1. Nuttallia hastata sp. nov. Biennial, 4-8 dm. high, scabrous throughout as other like species of the genus, branched from the upper half, or even lower, the branches strict and somewhat fastigiate, flowers from the ends of the stem and branches, somewhat cymous; lower leaves narrowly linear, 5-10 cm. long, sinuate and bluntly dentate, tapering to a petiole, upper leaves linear-hastate, attached by a broad base, deeply dentate with sharp teeth, acuminate; flowers white, petals eight to ten, about 2 cm. long, the alternate ones smaller and more pointed, but seldom anther-bearing; capsules variable in length, the longer ones about 3 cm. long, four- to five- angled, usually a pair of bracts at the base; seeds under a good lens thickly and evenly papillose, wing-margined. I have collected Nuttallia hastata at different times in several localities in Colorado, North Park, Middle Park, and Glenwood Springs, and not far from Jelm Post Office, Wyoming, by the road- side, on the east side of the Laramie River. The specimens which I have selected for the type were collected near Walden, North Park, Colorado, August 6, 1918, No. 5796. It does not closely resemble N. nuda (Pursh) Greene or N. stricta (Osterhout) Greene, but is what I suppose has been classified as N. Rusbyi (Wooton) Rydb. among the Colorado species. The flowers, however, are white, not yellow; and the capsule is angled, not cylindrical. Professor Wooton, in the original description of N. Rusbyi, observed, ‘‘stigma three-parted,” but this is characteristic of a number of species, perhaps of the genus. There is a peculiarity in the species of Nuttallia in that the flowers do not hold their color in the dried state. In the ‘Flora of the Rocky Mountains and adjacent plains,’’ Dr. Rydberg says of the white flowered species, ‘‘ petals straw-colored,” and this is their appaerance by the time they usually reach the herbarium, but at the time of opening they are white, as far as I am acquainted with them, pure white. 53 54 OSTERHOUT: NEW PLANTS FROM COLORADO 2. Phacelia formosula sp. nov. Biennial, 1.52 dm. high, stem single and upright, the flowering branches from near the top, or more commonly branched from near the base, the assurgent branches almost equaling the main stem, densely pubescent and somewhat hispid, becoming more glandular and hispid above; leaves lanceolate, or some of them oblanceolate, in outline. 5-7 cm. long pinnate with leaflets 5-10 mm. long, 3-5 mm. wide, entire or toothed, a terminal portion lobed or pinnatifid, not much enlarged, hispid with rather short hairs, the upper more glandular; inflorescence of several branches of two-ranked scorpoid racemes becoming 3-5 cm. long, corolla blue, its Jobes rounded and entire, the filaments and style long- exserted; mature capsule the length of the calyx lobes, or very nearly so, 4 mm. long, four-seeded, the seeds 2.5 mm. long, rounded on the back, foveolate, the hollowed ventral side with a salient ridge lengthwise through the middle, the edges rounded. I am indebted to Mr. J. Francis Macbride of the Gray Her- barium for the information that Mr. C. F. Baker’s No. 758, col- lected at Ouray, Colorado, in 1901, is the original specimen from which Phacelia Bakeri (Brand) F. Macbr. was described. It is possible that P. formosula may be considered as closely related to P. Bakeri, which Dr. Rydberg has made a synonym of P. glandu- losa Nutt., or may be it might be considered the same species. But P. formosula is a rather smaller plant, the leaves are smaller, the hispid pubescense is not so coarse, the calyx lobes are narrower, and the capsule is smaller and shorter pediceled. The difference is also notable in the character of the seeds. Dr. Gray, in. the Synoptical Flora of North America, observed of the group in which P. glandulosa is placed, “seeds oblong or elliptical, flatter and thin- ner”’ than those of the group in which P. crenulata Torr. is placed, and it might be added that their edges are surrounded by a thin flat margin. P. formosula has ‘‘seeds with excavated ventral face divided by a salient ridge,” and rounded edges, like those of P. crenulata or P. corrugata A. Nels., except that they are not corrugated on the edges. The seeds are like those of P. splendens Eastw., only smaller. The only locality from which I have P. formosula is North Park, Colorado; and the type specimens were collected near Walden, along the road descending to the Michigan Creek, August 6, 1918, No. 5794. : OsTERHOUT: NEW PLANTS FROM COLORADO 55 3. Oreocarya monosperma sp. nov. Biennial or short-lived perennial, stem single or several from the crown of the root, 2-3 dm. high, hispid with coarse spreading hairs and a finer pubescence beneath, branching from about the middle into a thyrsoid-paniculate inflorescense; lower leaves oblanceolate, obtuse, with the petiole 3-8 cm. long, about a cm. wide, becoming smaller upward, sessile, very hispid on both sides, the coarse hairs pustulate at base, a fine indument beneath; caylx lobes narrowly linear, in fruit becoming 8-9 mm. long, hispid and pubescent as the stem; corolla white, the tube the length of the calyx lobes, 4 mm. long, the anthers attached at the throat, filaments almost none; nutlets, only one maturing, ovate, mar- gined, the margin upturned, slightly more than 3 mm. long, lightly cross-ridged, and tuberculate between the ridges. Oreocarya monosperma in appearance is much like O. thyrsiflora Greene; the two plants are about the same size, but the thyrsus is not so long in comparison with the stem; the nutlets are darker in color, ridged more, and less tuberculate, one instead of four. Collected at Trinidad, Las Animas County, Colorado, July 20, 1918, No. 5754. 4. Mertensia Clokeyi sp. nov. Stem slender, 2.5—3 dm. or occasionally 4 dm. high, glabrous or thinly appressed pubescent, branching from the upper third into a loose paniculate inflorescense, the peduncles often long and the flowers clustered at their ends; leaves rather remote on the stem, the lower narrowly linear, 3-4 cm. long, 3-4 mm. wide, the upper lanceolate, or narrowly ovate-lanceolate, acuminate, all sessile by a broad base, pubescent on both sides, the upper thinly with appressed, sharp pointed hairs, the lower less appressed; pedicels strigose pubescent, calyx 3.5 mm. long, the lobes 2.5 mm., lanceolate, glabrous, the edges ciliate; corolla 10-11 mm. long the tube half the length; filaments long, about the width of the anthers. Mertensia Clokeyi was collected by Mr. Ira W. Clokey, of Denver, at Lake Eldora, Boulder County, Colorado, in woods, altitude 9,300 ft., No. 3161. It belongs to the M. /Janceolata group. The leaves are pubescent on both sides, the pedicels strigose, and the calyx glabrous, except the ciliate edges of the lobes; in these respects it is like Mertensia media Osterhout, but the leaves are much broader and different in shape. 56 OSTERHOUT: NEW PLANTS FROM COLORADO “5. Agoseris frondifera sp. nov. An acaulescent perennial, the scape occasionally bearing a leaf, and usually two large leafy bracts at the base of the involucre, 1-2 dm. high, glabrous except for the tomentum at the base of the involucre and some slight tomentum on the stem, mainly at the base; leaves narrowly oblanceolate, from 1 dm. to almost 2 dm. long, glabrous but not glaucous, entire or with several narrow lobes 1-2 cm. long; involucral bracts 2 cm. high, in three series, the inner narrow and scarious-margined, the outer broad, ab- ruptly acuminate, longer and covering the inner, with a long tomentose pubescence especially on the margins; rays yellow; achenes striate for 8 mm., the beak 6 mm. long, the pappus white, 10 mm. long. Agoseris frondtfera is related to A. montana Osterhout; the bracts are like that species, and occasionally A. montana bears a leaf on the scape. It isin every waya smaller plant than A. mon- tana, and the leaves are narrower. It was collected by Mr. Ira W. Clokey of Denver, as his label reads, at Camp Pitts, Boulder County, Colorado, in woods, altitude 9,600 ft., August 16, 1918, No. 3114. 6. ONOPORDUM TAURICUM Willd. On July 22, 1918, I stopped for a few minutes at the Green- horn Post Office in Pueblo County, Colorado, and across the road from the post office, growing in abundance along a little ravine was what I took to be some European Cirsium. The heads of purple flowers made an attractive showing. Mr. J. Francis Mac- bride of the Gray Herbarium identified it as O. tauricum Willd. The sporadic appearance of non-edible mushrooms in cultures of Agaricus campestris MICHAEL LEVINE (WITH PLATES 3-5) In studying the culture and development of Agaricus cam- pestris during the winter and spring of 1916-1917, I visited some of the largest establishments of commercial mushroom growers in the East and thus have had an opportunity to study the fleshy fungi, other than the commercial varieties of Agaricus campestris, which appear sporadically in the mushroom beds. A number of such types for this country have been recorded by Peck* and others, The appearance, in the beds of one of the largest mushroom growers in New York City, of great numbers of Panaeolus veneno- sus, a very poisonous mushroom recently described as a new species by Murrill,t furnished material for the special study of its phy- siological and toxicological properties and I have published my results along this line elsewhere.{ As this fungus seems quite dangerous I shall give here some results of observations made on the growth habits and describe another variety or ‘form in which it sometimes occurs. Panaeolus venenosus Murrill (PLATE 3, FIGS. I-8).—This species is of interest to mycologists, since up to the present time it has been found only in two widely separated mushroom houses in the vicinity of New York City, in beds spawned for Agaricus cam- pestris. The problem of its origin or occurrence in the wild state still remains unsolved. The plants studied were found in several different mushroom houses. In greenhouses which had been imperfectly darkened, better developed plants appeared, such as are shown in Fics.2 and 6. The plants grew in small fairy rings, mostly one to two feet in diameter, and in the darker mushroom houses were made conspicuous by the markedly | developed white * N. Y. State Mus. Bull. 157: 67-68, = IOTX; Bull. 150: 43: 1910. +A very dangerous mushroom. Mycologia 8: 186, 187. 1916. The physiological properties of two species of poisonous mushrooms. Torrey Club r7: 176-201, pl. r, 2. 1918. 57 Mem. 58 LEVINE: SPORADIC APPEARANCE OF NON-EDIBLE MUSHROOMS tomentose bases of the stipes, as shown in Fic. 7. My observa- tions as to the diagnostic characteristics of the fungus agree with the description given by Murrill. The spore print, however, although blackish in general, appears to have a delicate purplish hue. The shape of the spores is like that of the other species of Panaeolus. In my spore prints there is a great number of trans- lucent, possibly immature spores. The following characters are generally present. The sporophores are 6-12 cm. tall; they are cespitose or gregarious. The pileus is 2~5 cm. in diameter and is rather fleshy. Its shape differs with the age, being campan- ulate in young specimens, later becoming plane and umbonate. The surface is moist and hygrophanous and is fulvous to isabelline in color (Ridgw.) when young, and dark bay when mature. In mature specimens the surface becomes wrinkled, as shown in FIG. 1. The gills are adnately attached and fuliginous in color with a grayish white edge. The stipe is fleshy, but hollow in the center, and in length is approximately two to three times the diameter of the pileus. The surface is striate and is covered with small hair-like scales. The base of the stipe is, as noted above, con- spicuously covered with a white tomentum. The obviously different looking plants, shown in PLATE 4, FIGS, 12-14, appeared during the month of April in one of the greenhouses in which the floor below the benches was used for mushroom culture. These plants were not abundant but were collected twice during the month. The whole of the material weighed about 40 grams, most of which was used in the experiment to determine the physiological and toxological properties. It was found that the plant was poisonous in the same degree as P- venenosus. The sporophores in general are like those of P. veneno- sus but they differ materially in the length of the stipe. The stipe is 2-4 cm. long and 5-8 mm. thick; that is, one fifth to one third the size of the stipe of P. venenosus. It is striate, however, and is somewhat covered with hair-like scales, as in P. venenosus. It is hollow and tapers slightly toward the base. Its color is fulvous and darker at the base than the stipe of P. venenosus and the tomentum at the base, noted in P. venenosus, is poorly de- veloped and may be lacking. The taste and color are like those of P. venenosus. It may be regarded as a form of that species. LEVINE: SPORADIC APPEARANCE OF NON-EDIBLE MUSHROOMS 59 Panaeolus campanulatus L.,* which was the first member of this genus to be recognized as poisonous, also appeared in the mushroom beds. These plants appeared late, however, after the crop of Agaricus campestris mushroom was exhausted. The size and abundance of the sporophores might make them tempting to the uninitiated. The characteristics of the plants observed were identical with those already described for this species. The color of the pileus may be more accurately described as tilleul buff (Ridgw.) at the center and darker at the margin. Panaeolus retirugis Fr. (PLATE 2, FIGS. 9-11) was also found in © the beds mixed in with the sporophores of Agaricus campestris. It was the next most common to P. venenosus. A large number of these. plants were collected during the month of April and they had been fairly common earlier. Species of Panaeolust in general have been regarded as suspicious, especially P. retirugis. This plant was studied toxicologically by Ford and I have also found it to be poisonous to the same degree as P. venenosus when applied to the gastrocnemius muscle of the frog and the vagus nerves of frogs and turtles. The characters of the plant, as grown in dung- piles and well-manured lawns, are well known and the specimens collected by me were fairly typical. The occurrence of Panaeolus campanulatus and P. retirugis in the mushroom beds may be accounted for by the coprophilous nature of these plants. It is possible that the mycelium of these species of Panaeolus are brought into the mushroom houses with the manure or introduced by flying spores in early autumn. The other possibility lies in the method employed for obtaining commercial spawn. It appears that some of the spawn makers in the eastern United States are following the method employed in England and France for obtaining a commercial spawn.t This consists of making trenches in the sod, where “spontaneous” mycelium or sporophores of Agaricus campestris appear in the oe See ‘Mclivaine ; & Macadam. One thousand American Fungi 386. 1900 See Krieger, C.C.L. Note on the reported poisonous properties of Copriwus comatus. Indiana. Mycologia 3: 200-202. 1911. Also Murrill,W.A. Anew poi- sonous mushroom. Mycologia 1: 211-214. 1909. tSee Duggar, B. M. The cultivation of the mushroom. U. S. Dept. Agric. Farmers’ Bull. 204; 1-25. 1911; The — * mushroom growing and mushroom spawn making. U.S. Dept. Agr. Bur. Plant Ind. Bull. 85: 1-60. pl. 1-7, 1905; Mushroom growing 92. New York. fore 60 LEVINE: SPORADIC APPEARANCE OF NON-EDIBLE MUSHROOMS pasture, and filling them with thoroughly fermented manure. The vigorously growing mycelium spreads into this manure after several weeks. When this occurs, the manure is taken out of the trenches and slowly dried. This constitutes “virgin spawn,”’ from which the commercial spawn is made. By this method it is conceivable that the mycelia of a number of fungi may be found growing in the “virgin spawn” and be propagated in the commercial spawn. This at least seems to be a plausible explana- tion for the introduction of P. venenosus into mushroom houses. This is supported by the fact that the fungus appeared only after the beds were spawned; as mentioned above it appeared in two widely separated mushroom establishments, both of which, however, were using the spawn from the same spawn maker. How P. venenosus escapes observation in the field is still a question that remains unsolved. It is possible that the plant is a species of Psilocybe or Inocybe made aberrant by cultural conditions. Another possibility may be that the mycelium never has conditions in the open favorable to the development of sporophores but produces fruit bodies under cultivation only. Clitocybe dealbata Sow. (PLATE 2, FIGS. 15-17). A species of Clitocybe which I have identified as belonging to the variable species known as Clitocybe dealbata also appeared. These plants were found in several mushroom houses in great abundance, growing in large clumps in the beds of Agaricus campestris. The Clitocybe species were found in these houses from January to May. There are three recognized varieties of Chlitocybe dealbata Sow.., namely, var. minor Cooke, var. deformata Peck, and var. sudorifica Peck. The var. sudorifica was later made a species by Peck. The essential diagnostic characteristics of these plants, as given by: Peck follow. C. dealbata has a white, fleshy pileus with a Wavy margin; the gills are close, thin, adnate, and white in color; the stipe is fibrous, equal, and stuffed or hollow. Var. minor differs from the typical form of the species in its smaller and more regular form, its opaque gills, and the pleasant farin- aceous odor. Var. deformata has a thin white, and very irregular pileus with a wavy or lobed margin; the gills are adnate or LEVINE: SPORADIC APPEARANCE OF NON-EDIBLE MUSHROOMS 61 slightly decurrent. C. sudorifica Peck has an irregularly shaped pileus which often becomes lobed; the gills are adnate and slightly decurrent; the pileus is watery when moist and whitish or grayish white when dry. This species was confused with C. dealbata, but the presence of sudorific properties forms a good basis for distinguishing them. The size of the spores in all these forms is approximately the same. While C. dealbata and its varieties are generally known to inhabit lawns and grassy places, the typical form of the species and the var. deformata have been reported growing in mushroom houses. As far as may be judged from the descriptions and the study of Peck’s original herbarium material the species and varieties are separated only with great difficulty, yet I believe that the specimens I found are more like the typical C. dealbata, although I have not studied the toxicological properties of the juices of these various plants. Ford* and his collaborators found that the juices of C. sudorifica were very toxic to guinea pigs and rabbits. Peck de- scribed earlier the sudorific effect induced by eating this plant. Gillott and Clark and Smitht have recognized and studied the poisonous properties of other species of the genus. In view of the toxic properties of the members of the genus the abundance of this species in a commercial mushroom house is of interest. As in the case of Panaeolus the species may have been introduced into mushroom houses through commercial spawn or flying spores. Tricholoma melaleucum Quél. PLATE 5, FIGS. 21-23, shows a fungus which I found but once in the mushroom houses. The sporophores were discovered in an old mushroom bed. I am not altogether sure of its identity. There are slight discrepancies between my specimens and the description of this species given by Murrill§ (Melanoleuca melaleuca). These differences may cone be accounted for by the abnormal conditions under which d, W. W., & Sherrick, J. L. On the properties of several species of the Peyporaca and a new variety of or . peau dealbata sudorifica Pk. Jour. .and Exp. Ther. 2: 549-558. 1911; Further observations on Fungi, particularly Eidaiste ene gts Pk., Pholiota autumnalis ee and Inocybe decipiens Bres. Jour. har Exp. Ther. 4: 321-332. 1913- + Bick médicale sur l’empoisonnement par les champignons. Lyon. 1900. t Toxcological studies on the mushrooms Clitocybe Pee and tedinte infida. Mycologia 5: 224-232. pl. or. 1913. N. Am. Fl. 107: 7. 1914. 62 LEVINE: SPORADIC APPEARANCE OF NON-EDIBLE MUSHROOMS these plants grew. However, the description and published illustrations* of Tricholoma melaleucum best fit this plant. The sporophores I found are somewhat larger than the type described by Murrill, whose herbarium material I compared with my plants. The pileus is larger than this type and measures 3-9 cm. in diameter, while in height the plant agrees with the description. The pileus is thin, convex to plane and depressed. The margin is lobed and may become divided into a number of segments as shown in Fic. 23. The color of the pileusis drab to light drab (Ridgw.). The gills in these specimens were not very white, as described by Murrill, but drab gray. The spores of this plant agree perfectly with those of T. melaleucum. The stipe is even, although it may be enlarged at the top and at the base. The surface is reticulately veined as shown in Fic. 21 and in this respect is unlike the description. Its color agrees with the description. Up to the present T. melaleucum has been reported as found only in woods, fields and lawns. Its habit is solitary and the plants found in the mushroom beds were solitary, although they tended to be cespitose. Peziza domiciliana Cooke} (PLATES 4, FIGS. 18-20). The Peziza shown in Fics. 18-20 is common in all the mushroom houses studied. It makes its appearance as early as November and can also be found late in the spring. The plants generally appeared before and during the growth of the mushrooms. I am indebted to Dr. Seaver for the identification. While the plants agree in the main with the description given for P. domiciliana, there are some differences. The apothecia grow singly or gregariously but never caespitose. The outer surface of the cup has a white granular appearance and the color of the hymenium is pale ochra- ceous salmon (Ridgw.) when young, to Dresden brown, when old, instead of ochraceous buff or dungy buff as given by Overholtz and Seaver. While the spores are ellipsoidal and hyaline when young, and in this agree with the description already published, they are slightly smaller in size. A considerable number of species of Coprinus us in the * See Barla. Les via etdancics des Alves Matias, pl. 46,f.8-15. 1888. t See Seaver, F. J. Development of the cup Fungi. Mycologia 8: 195-198. pl. 188, 789. 10916, LEVINE: SPORADIC APPEARANCE OF NON-EDIBLE MUSHROOMS 63 compost after the beds were made and several species of Poria were found on the wooden framework in some of the more moist houses, but I shall not report on them at this time. DEPARTMENT OF BOTANY, COLUMBIA UNIVERSITY. Description of plates 3-5. In PLATEs 3 and 5 the natural sizes of the plants are represented; in PLATE 4 the figures are slightly reduced. PLATE 3 Fics. 1-4. Young stages in the development of Panaeolus venenosus Murrill. Fics. 5-8. Mature stages of Panaeolus veneno Fics. 9-11. Stages in the development of een retirugis Fr. PLATE 4 Fic. 12. Longitudinal section of a mature carpophore of Panaeolus venenosus with a short stipe. Fic. I Young carpophore of Panaeolus venenosus with a short s tipe. Fic. 14. The under surface of the pileus of Panaeolus venenosus with a short stipe. Fics. 15-17. Clusters of Clitocybe dealbata Sow., showing the upper and lower surfaces of the pilei Fics. 18-20. iauive stages in the development of Peziza domiciliana Cooke. PLATE 5 Fics. 21-23. Diff t stag fat 4 hat t cl ly resembles Tricholoma melaleucum Quél. INDEX TO AMERICAN BOTANICAL LITERATURE 1911-1918 aim of this Index is to include all current botanical literature written by Americans, PRS in America, or based upon American material ; the word Amer- ica epi d in the broadest sense. and papers that ‘ate exclusively to forestry, agriculture, horticulture, manatoed products of vegetable origin, or laboratory methods are not included, an attempt is made to index the literature of bacteriology. An occasional exception is ie 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. s Index is reprinted monthly on cards, and furnished in this form to subscribers at ads rate of one cent for each card, Selections of cards are not permitted ; each subscriber must wi 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. Agrelius, F. U. G. Botanical notes. Trans. Kansas Acad. Sci. 28: LO7-187.. “IOIS: 3 Agrelius, F. U. G. Botanical notes, 1917. Trans. Kansas Acad. Sci. 28: 118-120. 1918. Andrews, E. F. Adaptation and natural selection. Bot. Gaz. 66: 382. 160 1918. Andrews, F. M. Anthocyanin of Beta vulgaris. Proc. Indiana Acad. Sci. 1917:167. 1918. Andrews, F. M. The effect of centrifugal force on plants. Proc Indiana Acad. Sci. 1917: 175. 1918. Andrews, F. M. Improved forms of Maximow’s automatic pipette. Proc. Indiana Acad. Sci. 1917: 169-173. f. 1-4. 1918. Andrews, F. M. Stoppage of a sewer line by roots of Acer Saccharum. Proc. Indiana Acad. Sci. 1917: 165. I1g18. Andrews, F. M. Studies on pollen. Proc. Indiana Acad. Sci. 1917: Arthur, J. C. Uredinales of Guatemala based on collections by E. W. D. Holway.—II. Aecidiaceae, exclusive of Puccinia and form- genera. Am. Jour. Bot. 5: 420-446. 9 N 1918;—III Puccinia, ex- clusive of species on Carduaceae. Am. Jour. Bot. 5: 462-489. 30 N 1918. ncludes new species in Ravenelia (5) Uropyxis (1) Skierka (1), Pucciniosira (1), Uromyces (3) and Puccinia (11). 65. 66 INDEX TO AMERICAN BOTANICAL LITERATURE Ashe, W. W. Additions to the arborescent flora of North Carolina. Jour. Elisha Mitchell Sci. Soc. 34: 130-140. S 1918. Includes A melanchier austro-montana sp. nov. and many new combinations. Bailey, I. W., & Tupper, W. W. Size variation in tracheary cells: I. A comparison between the secondary xylems of vascular Crypto- gams, Gymnosperms and Angiosperms. Proc. Am. Acad. Arts and Sci. §4: 149-204. f. -6. S$ 1918. Beals, C. C. The effect of aeration on the roots of Zea Mays—l. Proc. Indiana Acad. Sci. 1917: 177-180. f. 1-3. 1918. Bicknell, E. P. Aster laevis. Addisonia 3: 47, 48. pl. 104. S 1918. Blake, M. A. Distribution of peach yellows in nursery stock. Proc. Soc. Hort. Sci. 1910: 46-51. Ap I9gII. Blake, M. A. Factors which determine color in the forcing of roses. Proc. Soc. Hort. Sci. 1910: 19-27. Ap 1911. Britton, N. L. Byron David Halsted. Jour. N. Y. Bot. Gard. 19: 221. S 1918. Britton, N. L. Opuntia Opuntia. Addisonia 3: 49, 50. pl. 105. S 1918. Brown, E. W. Regeneration in Phegopteris polypodioides. Bull. Torrey Club 45: 391-397. f. 1-3. 19 O 1918. Brown, W. H. The fungi cultivated by termites in the vicinity of - Manila and Los Bafios. Philip. Jour. Sci. 13: (Bot.) 223-231. O44. TAGs. Cockerell, T. D. A. Notes on the flora of Boulder County, Colorado. Torreya 18: 177-183. 29 O 1918. Coit, J. E. The brown spot of the navel orange. Proc. Soc. Hort. Sci. 1910: 62-67. Ap IgIt. Coker, D. Revision of the North American species of Encalypta. Bull. Torrey Club 45: 433-449. pl. 13, 14. 15 N 1918. Coker, W. C. A visit to Smith Island. Jour. Elisha Mitchell Sci. Soc. 34: 150-153. pl. 10-16. S$ 1918. Cowgill, H. B. Cross-pollination of sugar cane. Jour. Am. Soc. Agron. 10: 302-306. N 1918 Crandall, C. S. The vitality of pollen. Proc. Soc. Hort. Sci. 1912: 121-130. Mr 1913. Dixon, H. N. Uganda mosses collected by R. Diimmer and others. Smithsonian Misc. Col. 69: 1-10. pl. 7. 21 O 1918. Eight new species in various genera are described. INDEX TO AMERICAN BOTANICAL LITERATURE 67 Dorsey, M. J. The inheritance and permanence of clonal varieties. Proc. Am. Soc. Hort. Sci. 1916: 41-71. Mr 1917. Dorsey, M. J. Sterility in the grape. Proc. Soc. Hort. Sci. 1912: 149-153. Mr 1913. Dudgeon, W. Morphology of Rumex crispus. Bot. Gaz. 66: 393-420. pl. 17-19 + f. I-21. 15 N 1918. . Dufrenoy, J. Pine needles, their significance and history. Bot. Gaz. 66: 439-454. f. 1-29. 15 N 1918. Dymond, J. R. Seeds. Ottawa Nat. 32: 52, 53. S 1918. Ferris, R. S. Taxonomy and distribution of Adenostegia. Bull. Torrey Club 45: 399-423. pl. ro-12. 19 O 1918. Adenostegia Hanseni, A. parviflora, A. littoralis, A. Helleri, and A. palmata, spp. nov., are described. Fisher, D. F. Apple powdery mildew and its control in the arid regions of the Pacific northwest. U. S. Dept. Agr. Bull. 712: 1-28. pl. 73 + 7.7; 2. 260 1958. Fitch, C. L. Field studies of a leafroll disease of potatoes. Proc. Soc. Hort. Sci. 1912: 44-51. Mr 1913. Fitzpatrick, H. M. The cytology of Eocronartium muscicola. Am. Jour. Bot. 5: 397-419. pl. 30-32. 9 N 1918. Free, E. E. A colloidal hypothesis of protoplasmic permeability. Plant World 21: 141-150. Je 1918. Fromme, F. D. An automatic spore trap. Phytopathology 8: 542- 544. f.7. O rgr8. : Gleason, H. A. On the development of two plant associations of ‘northern Michigan. Plant World 21: 151-158. Je 1918. Goodspeed, T. H. Method of replacing paraffin solvent with paraffin. Bot. Gaz. 66: 381, 382. 160 1918. Graff, P. W. Philippine Basidiomycetes—III. Bull. Torrey Club 45: 451-469. pl. 15. 15 N 1918. Harper, R. M. The American pitcher-plants. Jour. Elisha Mitchell Sci. Soc. 34: 110-125. pl. 2-6. S 1918. Harrington, G. T., & Crocker, W. Resistance of seeds to desiccation. Jour. Agr. Research 14: 525-532. 16S 1918. Hartley, C. Stem lesions caused by excessive heat. Jour. Agr. Research 14: 595-604. f. 1. 23 S 1918. Harvey, R. B. Hardening process in plants and developments from frost injury. Jour. Agr. Research 15: 83-112. pl. 7-11, A. + f. 1-3. 14 O 1918. 68 INDEX TO AMERICAN BOTANICAL LITERATURE Hayes, H. K., & East, E. M. Improvement in corn. Connecticut Agr. Exp. Sta. Bull. 168: 1-21. pl. 1-4. Je 1911. Hedgcock, G. G., Bethel, E., & Hunt, N. R. Pifion blister-rust. Jour. Agr. Research 14: 411-424. pl. 54-57 + f.1. 2S 1918. Hedrick, U. P. Natural resistance to disease in fruits. Proc. Soc. Hort. Sci. 1912: 106-114. Mr 1913. Hedrick, U. P., & Wellington, R. The hereditary transmission of char- acters of apples. Proc. Soc. Hort. Sci. 1911: 19-29. F 1912. ‘Heimlich, L. F. The trees of White County, Indiana, with some reference to those of the state. Proc. Indiana Acad. Sci. 1917: 388-471. pl. I-32. 1918. Higgins, J. E. Sex in Carica papaya and its relation to breeding and cult re. Proc. Soc. Hort. Sci. 1910: 75-78. Ap 1911. Hill, A. W. The genus Caltha in the southern hemisphere. Ann. : 421-435. f. 7-10. Jl 1918. “gens ue C. alata, and C. phylloptera, spp. nov., are described. Hodgson, R. W. A sterigmatocystis smut of figs. Phytopathology 8: 545, 546. O 1918. Hoffer, G. N., & Glanasoff, D. Corn-rootrot and wheat scab. Jour. Agr. Research 14: 611, 612. 23S 1918. Preliminary paper Holmes, J. S. Extension of the range of Prunus umbellata into North Carolina. Jour. Elisha Mitchell Sci. Soc. 34: 126-129. pl. 7, 8 S 1918. Hotson, J. W. Sphagnum as a surgical dressing. Jour. Am. Peat Soc. 13: 195-226. f. 7-18. O 1918. Howard, W. L. The rest period in plants. Proc. Soc. Hort. Sci. 1910: 33-46. Ap IgII. Howe, M. A. Calcareous algae from Murray Island, Australia, and Cocos-Keeling Islands. Papers Dep. Mar. Biol. Carnegie Inst. Washington 9: 291-296. pl. 97, ae 2: fo 98. 16 Au 1918. Carnegie Inst. Washington Publ. no. Jackson, H. S. The Uredinales of Sie Sci. 1917: 311-385. 1918. Jackson, H.S. The Uredinales of Indiana—II. Sci. 1917: 133-137. 1918. Jackson, H. S. The Ustilaginales of Indiana. Sci. 1917: 119-132. 1918. Proc. Indiana Acad. Proc. Indiana Acad. Proc. Indiana Acad. INDEX TO AMERICAN BOTANICAL LITERATURE 69 Johnston, J. R. Algunos Hongos entomogenos de Cuba. Mem. Soc. Cubana Hist. Nat. ‘‘ Felipe Poey”’ 3: 61-82. pl. 1,2. 1918. Includes Spicaria aleyrodis and Torrubiella lecanii, spp., nov. Johnston, J. R., & Bruner, S. C. Enfermedades des naranjo y otras plantas citricas. Cuba Est. Exp. Agron. Bull. 38: 1-54. pl. 1-15. Au 1918. Jones, D. H. Bacteria—friends and foes. Ontario Dept. Agr. Bull. 265: 1-99. Au rgr8._ [Illust.] Includes chapters on bacterial diseases of plants and bacteria of the soil and manure pile. Knowlton, C. H., Ripley, W. S., & Weatherby, C. A. Preliminary lists of New England plants,—-X XVI. Rhodora 20; 182-185. 11 N 1918. Kunkel, L. O. Tissue invasion by Plasmodiophora brassicae. Jour. Agr. Research 14: 543-572. pl. 61-80 +f. 12,2. 16S 1918. Long, B. Eragrostis peregrina a frequent plant about Philadelphia. Rhodora 20: 173-180. 11 N 1918. MacCaughey, V. The Hawaiian sumach. Neneleau; Rhus semialata var. sandwicensis Engler. Torreya 18: 183-188. 29 O 1918. Mackenzie, K. K. Charles Keene Dodge. Torreya 18: 188-190. 29 O 1918. Mann, A. Diatoms from Murray Island, Australia. Papers Dep. Mar. Biol. Carnegie Inst. Washington 9: 297. 16 Au 1918. Carnegie Inst. Washington Publ. 213. McClintock, J. A. A disease of coldframe parsley caused by Sclerotinia libertiana. Virginia Crop Exp. Sta: Bull. 18: 379-384. f. 84-86. I Ja 1916. McMurran, S. M. Preventing wood rot in pecan trees. U. S. Dept. Agr. Farm, Bull. 995: 1-8. f. 1-10. Jl 1918. McNaught, J. B. The algae of Kansas reservoirs. Trans. Kansas Acad. Sci. 28: 121-128. 1918. McWharf, J. M. Echinacea: its combination and use. Trans. Kansas Acad. Sci. 28: 179-180. 1918. Markle, M.S. A comparison of the plant succession on Hudson River limestone with that on Niagara limestone, near Richmond, Indiana. Proc. Indiana Acad. Sci. 1917: 109-113. 1918. Markle, M. S. Notes on microscopic technique. Proc. Indiana Acad. Sci. 1917: 115-117. 1918. {Illust.] Nash, G. V. Callicarpa japonica. Addisonia 3: 45. pl. 103. S 1918. 70 INDEX TO AMERICAN BOTANICAL LITERATURE Nash, G. V. Cornus mas. Addisonia 3: 41. pl. ror. S 1918. Nash, G. V. Crassula portulacea. Addisonia 3: 57. pl. 109. S 1918. Nash, G.V. Ilex serrata argutidens. Addisonia 3: 51. pl. 106. S 1918. Nash, G. V. Magnolia kobus. Addisonia 3: 55, 56. pl. 108. S 1918. Nash, G. V. Othonna crassifolia. Addisonia 3: 53. pl. 107. S 1918. Nelson, J. C. Notes on the flora of Lake Labish, Oregon. Torreya 18: 191-195. 29 O 1918. Nichols, G. E. The sphagnum moss and its use in surgical dressings. Jour. N. Y. Bot. Gard. 19: 203-220. pl. 216-218. S 1918. Norton, J. B. S. Some unusual tomato variations. Proc. Soc. Hort. Sci. rg10: 71-75. Ap I19II. Nothnagel, M. Resistance of Mucorzygotes. Proc. Indiana Acad. Sci. 1917: 181-187. 1918. Noyes, H. A. Reaction of culture media. Proc. Indiana Acad. Sci. 1917: 149-162. 1918. Noyes, H. A., Trost, J. F., & Yoder, L. Root variations induced by carbon dioxide gas additions to soil. Bot. Gaz. 66: 364-373. f. I-9- 16 O 1918. Osner, G. A. Additions to the list of plant diseases of economic im- portance in Indiana. Proc. Indiana Acad. Sci. 1917: 145-147. 1918. Ottley, A. M. A contribution to the life history of Impatiens Sultant. Bot. Gaz. 66: 289-317. pl. 14, 15. 16 O 1918. Pammel, L. H. Prickly lettuce. Rhodora 20: 180, 181. O 1918. Pfeiffer, N. E. The sporangia of Thismia americana. Bot. Gaz. 66: 354-363. pl. 16. 160 1918. Pipal, F. J. A suspected case of stock poisoning by wild onion (Allium canadense). Proc. Indiana Acad. Sci. 1917: 139-143. 1918. [Illust.] Price, H. L. Some results of cabbage crosses. Proc. Soc. Hort. Sci. 1g10: 53-60. Ap IgII. Reddick, D., & Stewart, V. B. Varieties of beans susceptible to mo- saic. Phytopathology 8: 530-534. O 1918. Reed, H. S. Absorption of sodium and calcium by wheat seedlings. Bot. Gaz. 66: 374-380. f. 1. 160 1918. Reimer, F.C. Self-sterility of rotundifolia grapes. Proc. Soc. Hort. Sci. r9t0: 27-32. Ap Igit. Reynolds, E. S. Two tomato diseases. Chiytonatticlogy 8: 535-542. j. 1,2 Q 1918. INDEX TO AMERICAN BOTANICAL LITERATURE i Robinson, B. L. A descriptive revision of the Colombian Eupatoriums. Proc. Am. Acad. Art. & Sci. 54: 264-330. 8 O 1918. Robinson, B. L. Diagnoses and notes relating to tropical American Eupatorieae. Proc. Am. Acad. Arts & Sci. 54: 235-263. 8 O 1918. New species are described in Eupatorium (37), Fleischmannia (1) and Kuhnia (1). Robinson, B. L. Keyed recensions of the Eupatoriums of Venezuela and Ecuador. Proc. Am. Acad. Arts & Sci. 54: 331-367. 8 O 1918. Rolfs, P. H., & Fawcett, H.S. Fungus diseases of scale insects and white fly. Florida Agr. Exp. Sta. 119: 71-82. f. 14-31. N 1913. Rusby, H. H. Viburnum prunifolium. Addisonia 3: 59. pl. I70. S 1918. Sampson, A. W. Climate and plant growth in certain vegetative asso- ciations. U.S. Dept. Agr. Bull. 700: 1-72. f. 1-37. 14 O 1918. Sargent, C. S. Notes on North American trees—III. Tilia. Bot. Gaz. 66: 421-438. 15 N 191 Five new species and 7 varieties are jel. Shaw, W. R. Some microtechnical methods and devices. Philip. Jour. Sci. 13: (Bot.) 241-261. f. 1-5. Jl 1918. Schneider, C. Notes on American willows—II. The species related to Salix glauca L. Bot. Gaz. 66: 318-353. 16 O 1918. 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Forestry 24: 612-614. O 1918. Weatherwax, P. ioonrend technique for corn pollination. Proc. Indiana Acad. Sci. 1917: 105-107. f. 1, 2. 1918. Weatherwax, P. Variation and varieties of Zea Mays. Proc. Indiana Acad. Sci. 1917: 99-103. 1918. Wellington, R. Raspberry breeding. Proc. Soc. Hort. Sci. 1912: 155-159. Mr 1913. Wernham, H. F. Tropical American Rubiaceae—X. Jour. Bot. 55: 336-341. 1917;—X. The genus Manettia. Jour. Bot. 56: (Suppl.) 1-8. 918 Includes description of six new species in Psychotria (2) and Palicourea (4). White, O. E. Inheritance studies on castor beans. Brooklyn Bot. - Gard. Mem. 1: 513-521. pl. 23-28. 6 Jl 1918. Wieland, G.R. A study of some American fossil Cycads. Part VIII. Notes on young floral structures. Am. Jour. Sci. 46: 645-650. f. I. N 1or18. Wilson, O. T. A storage fermentation of dasheens. Phytopathology 8: 547-549. f. 1. O 1918. Wolfe, J. J. Alternation and parthenogenesis in Padina. Jour. Elisha Mitchell Sci. Soc. 34: 78-109. pl. r. S 1918. BULL, TORREY CLUB VOLUME 46, PLATE 3 Oo 4 * we ED) z LEVINE: NON-EDIBLE MUSHROOMS 4 46, PLATE VOLUME TORREY CLUB BULL. ) MUSHROOMS NON-EDIBLE LEVINE: BULL. TORREY CLUB VOLUME 46, PLATE 5 LEVINE: NON-EDIBLE MUSHROOMS Vol. 46 No, 3 BULLETIN OF THE TORREY BOTANICAL CLUB MARCH, I919 Gametogenesis and fecundation in Zea Mays as the basis of xenia and heredity in the endosperm PauL WEATHERWAX (WITH PLATES 6 AND 7 AND TWO TEXT FIGURES) In the almost endless number of varieties of maize there is afforded excellent material for experimental work on heredity, as is indicated by the part that the plant has played in the genetic studies of the past twenty years; but, unfortunately, maize offers at the same time limited opportunity for satisfactory cytological work. In addition to its other advantages, maize has a variable endosperm, in the transmission of whose characteristics we get not only a deeper insight into the true nature of the angiosperm endosperm but also some interesting checks on general theories of heredity. But many of the conclusions drawn from these endosperm studies have been based upon the assumption of certain facts concerning gametophyte development and a single description of the fecundation process, of which no figures were given. It was for the purpose of putting the cytological side of the question upon a substantial basis that this study was under- taken. The structure and development of the spikelets have been described in other papers (23, 24), and only the essential details will be repeated. THE MEGASPORE Before the integuments have made any considerable progress in enclosing the nucellus, the archesporium becomes visible as a [The BULLETIN for February (46: 37-72. pl. 3-5) was issued February 25, 1919] 73 74 WEATHERWAX: GAMETOGENESIS IN ZEA Mays large hypodermal cell near the tip of the nucellus (PLATE 6, FIG. 1). It soon divides periclinally, giving rise to a parietal cell, the tapetum, and a megaspore mother-cell; but no wall is formed, and the tapetal cell is immediately consumed (Fic. 2). The parietal layer of tissue, five to eight layers of cells in thickness, which is ultimately present (Fic. 16), is formed from epidermal cells by periclinal divisions (Fic. 2 and TExT-FIG. 1). In another connection (24, p- 492) the statement was made that, inasmuch as no evidence had been seen of the ALK disorganization of any of the fs MAL megaspores, all four probably ae e AY (} J, As, 7] sa S ; ’ p EY LAN Wiper “ functioned as in the lilies. NY But it has since been pointed out to me by Mr. E. G. a as Binucleate stage oF megaspore Anderson, of Cornell Univer- Tice uns cage a es ie ity, tat it eneh be the eae, ment; w, ovary wall; ”, tip of nucellus. the results of certain experi- ments on heredity in the en- dosperm must receive a new interpretation. Consequently, a thorough investigation of the development of the megaspore and embryo sac was thought profitable. The heterotypic division of the mother-cell is, in all essential details, similar to that in other plants (Fics. 3, 4). The chromo- somes are small and the karyokinetic figures indistinct; and the material is not satisfactory for giving any light on controverted questions of cytology in general. The bivalent character of the chromosomes at this stage is, however, beyond question. A wall is laid down between the two cells resulting from this division (Fic. 5). The more deeply-seated of these two cells next divides, giving rise to two megaspores (FIG. 6). The other cell begins to divide, but the process is arrested, usually at about the spindle stage, by the incipient disintegration of the dividing cell and the adjacent megaspore (Fic. 7). Thus, only the chalazal one of the four megaspore potentialities persists (FIG. 8). The absorption of the other three is not completed until the embryo sac is almost mature (Fics. 8-12). | WEATHERWAX: GAMETOGENESIS IN ZEA Mays 75 THE EMBRYO SAC The germination of the megaspore is the same as that in most plants where an ordinary seven-celled embryo sac is to be formed. In the two-nucleate stage (FIG. 9) there appears a large central vacuole, accompanied by a smaller one at the chalazal end of the cell; these vacuoles persist through subsequent stages until the ultimate organization of the embryo sac. The position of the spindles following the four-nucleate stage (FIG. 11) and the arrange- ment of the following eight nuclei (Fic. 12) substantiates the observation made on numerous other plants that one of the polar nuclei is the sister of the egg. When the membranes are first formed (Fic. 12), dividing the gametophyte into cells, the antipodals are much larger than the cells of the egg apparatus; but, by the time the embryo sac is fully organized (PLATE 7, Fic. 13), the egg and the synergids have greatly enlarged, and, with little or no increase in size, the antipodal cells have begun to divide, forming the multicellular antipodal tissue which seems to characterize the embryo sacs of most grasses. Meanwhile, the polar nuclei have come together near, or in contact with, the plasma membrane of the egg. After its organization, the embryo sac continues to increase in size, at the expense of the nucellus, until, at the time of fecundation, it has a volume five to ten times as large as it had at the eight- nucleate stage. This growth is accompanied by rapid changes in other parts of the pistil as well. The nucellus grows rapidly and pushes the integuments out against the ovary wall, which, growing less rapidly, is kept tightly stretched over the turgid ovule. The style, or “silk,” increases from a length of one or two centimeters to its full length, which may be forty centimeters or more in some varieties, and develops its numerous stigma hairs. Guignard (10, p. 44) has given a good description of the embryo sac. It occupies relatively a very small part of the nucellus (Text FIG. 2). The cells of the egg apparatus are very large (Fics. 13, 16). The dense cytoplasm of the pear-shaped synergids shows the longitudinal striations characteristic of these cells in many other plants. The nuclei are located well toward the micro- pylar ends of the cells and react so feebly to stains as often to be indistinguishable. The egg is larger and more rounded than a 76 WEATHERWAX: GAMETOGENESIS IN ZEA MAys synergid, and its cytoplasm is much less dense. The nucleus is almost centrally located and is closely surrounded by most of the cytoplasm of the cell. It is a little larger than that of a synergid and always stands out distinctly; the nucleolus is large, but the chromatin granules are small and scattered. The polar nuclei resemble the egg nucleus but are often a little larger. They are usually surrounded by a dense mass of cytoplasm, which often makes detailed observa- tion difficult. They do not fuse until after the entrance of the pollen tube to the em- bryo sac, even though pollina- tion be artificially delayed until long after the normal time. The antipodal tissue con- tinues to grow until the time of fecundation, often consist- ing of fifty or more cells, some Fic. 2. Longitudinal section of female z floret, for — of orientation, X 75: /, of which may have more than lemma; ?, palea; e, embry m,nucellus; one nucleus. A number of o, outer ei ks #, inner ‘[itepankent= these have been observed in m, micropyle; ¢c, stylar canal; #, course of Gets pollen tube after reaching the ovary; s, the Process of division, and, rudimentary stamen. while accurate counts of the chromosomes have not been made, there is nothing in the appearance of the karyokinetic figures to indicate ,that there are any more than the haploid number as has been reported for many other plants. The theoretical signific- ance of this antipodal tissue in the grasses has never been fully investigated, and it may help some time to explain the female gametophyte of the angiosperms. It is probably the equivalent of the endosperm of gymnosperms, but in maize it is completely absorbed soon after fecundation ~~ never becomes a part of the endosperm of the mature seed. ~ WEATHERWAX: GAMETOGENESIS IN ZEA Mays fas THE POLLEN TUBE As has already been noted (24, p. 487), the generative cell of the pollen grain has divided, forming two very small, crescent- shaped sperms, before the dehiscence of the anther. The pollen grain finally comes to rest on the silk where it is probably held by some sticky exudation that may be instrumental in initiating germination. The pollen tube emerges very soon and makes its way down through one of the stigma hairs to the body of the silk (Fic. 14). Here it pushes its way between the parenchyma cells, absorbing food from them, no doubt, until it reaches one of the vascular bundles, which it follows toward the ovary. At the time of its emergence from the pollen grain, the tube is greatly exposed to desiccation, and this continues until it has made its way into the body of the silk. This, together with the fact that the pollen grains themselves dry up very readily and are seldom viable for more than a day or two, probably accounts for the importance to the corn crop of warm rains at the time of polli- nation. In some kinds of corn the silks may attain a length of fifty centimeters or more, and there are probably few species of plants in which the pollen tube has farther to grow. Consequently, it has been an interesting problem to determine the time elapsing - between pollination and fecundation. A number of experiments were made to determine this, and the best results were obtained with a small variety of sweet corn with silks about twenty-five centimeters in length. Silks that had been properly protected were pollenized at 2:45 P.M. on a warm, foggy day in early Sep- tember, with pollen that had been shed on the forenoon of the same day, and ovules and segments of silks were fixed at various intervals for two or three days after pollination. It was found that within two hours the pollen tubes were well established in the silks. This result has also been verified by several experiments in the laboratory (Fic. 14). Ovules fixed twenty-five hours after pollination showed fecundation and stages just before (Fic. 16) and just after fecundation. These results could, of course, be modified by extensive experiments with silks of different lengths and pollen of different ages and at different temperatures, degrees of humidity, etc., but that is rather the problem of the physiologist. 78 WEATHERWAX: GAMETOGENESIS IN ZEA MAys The aim in presenting here the meager data available is to show the great rapidity with which the pollen tube grows. It is inter- esting to note that Jensen (13, p. 15) finds a somewhat greater time (32 to 40 hours) between pollination and fecundation in wheat, although the style of the latter is much shorter than in maize. It is not to be inferred that there is at any one time a living pollen tube reaching from the stigma hair to the embryo sac; and this statement is probably very generally true of all angiosperms. Observation of the pollen tube in the silk is difficult, but it is probable that there is not more than a centimeter or two of it alive at any one time; and the living portion seems to become shorter and broader as it approaches the ovule. It is in most cases practically impossible to find any trace of the pollen tube in _ the tissue through which it has passed. One of the difficulties in attempting to observe fecundation is in being able to identify the sperms when they reach the embryo sac. The protoplasm of the pollen tube, as it grows down through the silk, is so dense and so coarsely granular that it is almost im- possible to distinguish with certainty the sperms and the tube nucleus. Both can be made out with a fair degree of certainty, however, in the less dense contents of the tube in the ovary cavity as it approaches the embryo sac; and they are, of course, visible in the pollen grain before germination. The tube nucleus is irregular — in size and shape and does not seem to have a definite membrane. There is no visible difference between the two sperms. They are very small, slender crescents, with the ends drawn out to very fine points (Fics. 15, 16). The nucleus, which seems to consist almost wholly of chromatin, constitutes the greater part of the cell and forms the middle of the crescent; no nucleolus has been observed. There is only a small quantity of cytoplasm, and noth- ing of its details can be made out. The size of the sperm is remarkably correlated with the long distance through which it must be carried from the pollen grain to the embryo sac. Practically all varieties of maize are fertile inter se; and this means that any pollen grain, even though it be produced by a plant characterized by short silks, is capable of producing a pollen tube as long as is necessary, as is determined by the length of the silk of the variety pollenized. | WEATHERWAX: GAMETOGENESIS IN ZEA Mays 79 The path of the pollen tube on reaching the ovary has never been satisfactorily explained. True (20, p. 217) quotes Hackel (11) as saying that the outer integument acts as a conducting tissue. Guignard (10, p. 43) says that the tube enters the ovary by way of the stylar canal. My preparations have not given conclusive results, but .they do not tend to substantiate these opinions. The pollen tube seems to grow downward from the base of the style until it reaches the inner epidermis of the wall of the ovary (TExT FIG. 2). Remnants of it can be traced just below the epidermis to a point near the micropyle, where it enters the ovarian cavity. After following a very crooked course here, it finally enters the micropyle, forces its way between the cells of the parietal tissue, and enters the embryo sac (FIG. 16). FECUNDATION The contents of the pollen tube may be emptied between the cells of the egg apparatus or into any one of them; but it is usually emptied into a synergid. The membrane of the other synergid often seems to break at about the same time, emptying its contents into the cavity of the endosperm cell. The tube nucleus is soon lost to view, if it was visible at all; and, because of their extremely diminutive size, it is very difficult to follow the sperms beyond this point. The cytoplasm of the area to be examined is very dense and is filled with numerous globules of various sizes and shapes (FIGS. 13, 16), produced, no doubt, by the mixing of the cell contents brought together by the entrance of the pollen tube. The disorganizing synergids give rise to cytoplasmic differentia- tions that are especially confusing. Material fixed with reagents containing osmic acid presents almost hopeless complications; but, with chromo-acetic acid as a fixing agent, followed by Flem- ming’s triple stain, the sperm nuclei can be identified. In several preparations a small red object, which is like a sperm nucleus in size, shape, and granular appearance, has been seen sticking to the egg nucleus (Fic. 17); and a similar body has, in a number of instances, been seen attached to one (Fic. 18) or both (Fic. 19) of the polar nuclei, which were in the process of fusion. No fusing polar nuclei have been seen without this sperm-like body. On this basis, then, it seems certain that the so-called “‘double fecundation”’ takes place in maize. 80 WEATHERWAX: GAMETOGENESIS IN ZEA MAYS In 1901 Guignard (10) reported having observed this process, but his otherwise excellent paper was not illustrated. While his results have been generally accepted by morphologists, and made the basis of extensive work by geneticists, the desirability of a verification is attested by the number of students who are known to have attempted at different times to repeat his work. If negative results were generally published, we should, no doubt, have a much more voluminous literature on this point. In one preparation the writer observed the two fusions taking place simultaneously (Fics. 17, 18), but in a majority of cases one preceded the other. Whether or not there is a definite order in which the fusions occur has not been determined, because it is almost impossible to determine by appearance alone whether a cell under observation is an egg or a zygote. It is true in all cases, however, that several free endosperm nuclei are formed before the first division of the fecundated egg. At the time of fecundation the egg nucleus is often seen to have moved to one side of the cell (Fic. 16), and the polar nuclei may also migrate within a limited range, often approaching the micropylar end of the embryo sac. Whether or not there is any significance to be attached to these migrations has not been de- termined. These fusions take place while the egg nucleus and the polar nuclei are in a resting condition (Fics. 17-19). The chromatin is gathered into numerous round, globular bodies, some of them almost as large as a sperm; and these granules are loosely connected by strands composed of finer granules of similar appearance. That these chromatin bodies are not the “prochromosomes”’ of some authorities is attested by their number, which is far too great. XENIA As long as maize has been cultivated, it has been noted that, when white and colored varieties of some kinds were grown close together, the ears of the former were likely to bear a few colored seeds. The American Indians are said to have observed this and to have attributed it to the intermingling of the roots underground. Later, civilized man attributed the phenomenon to some effect of cross pollination, but its mechanism long remained a mystery. WEATHERWAX: GAMETOGENESIS IN ZEA MAyYs 81 In 1881, Focke (9, p. 511) coined the word xenia to apply to the immediate effect of foreign pollen upon maternal tissue. Nawaschin’s discovery (16) that in many plants, one of the sperm nuclei enters into the makeup of the endosperm, suggested an explanation of the mechanism of xenia in maize; and Guignard’s discovery of ‘“‘double fecundation”’ in maize, in 1901 (10), left no room for reasonable doubt as to the mechanism of xenia. The immediate effects of cross pollination, that is, such effects as may immediately be observed in the seed or fruit in which the hybrid embryo is borne, are probably not so common as has often been supposed; and by no means are all of them illustrations of xenia, in the true sense of the word. There is a common belief that pumpkins growing near water- melons will, by hybridization, impair the quality of the latter in the first generation; and numerous other instances have been cited of slight changes in the quality of fruits, supposed to be due to the immediate effects of cross pollination. The experimental demonstration of most of these, with races known to be genetically pure for the characteristics considered, has not been accomplished. Another type of the phenomenon is afforded by hybrids between varieties of peas and other plants, which are variable as to the color or physical character of the cotyledons. Here a dominant character may, through hybridization, appear in seeds borne on plants pure for the recessive character. But this is merely the early recognition of a hybrid by means of characteristics that are differentiated in the embryo. Bailey and Gilbert (1) must have had in mind such phenomena as these when they made the errone- ous statement (p. 327) that xenia occurred in peas. In contrast with these phenomena is the immediate effect upon the endosperm, so well known in maize. This has also been demonstrated in teosinte-maize hybrids (21), and has been re- ported in a few other crosses between different varieties of cereals. The primary essential for xenia is variability of endosperm in plants that will hybridize, and for this reason, maize when used for one of the parents, at least, furnishes the best-known illustration. In a recent review of the whole question of influences following fecundation, Waller (22) suggests that the term xenia be reserved for the phenomenon limited to the endosperm of angiosperms and 82 WEATHERWAX: GAMETOGENESIS IN ZEA MAys due to the entrance of a sperm into the constitution of the primary endosperm nucleus. It is a direct result of the introduction of hereditary factors into the endosperm and not to be attributed to enzymatic action or other stimulating influence. To include “those influences which follow fertilization but are remote from it’’ (p. 282) and are ‘‘due to the developing zygote”’ (p. 284), he proposes the term ectogony. Whether or not the new term is applicable, is not a question to be decided in this connection; but the distinction between xenia and other influences less directly connected with fecundation isa timely one. Xenia may be defined, then, as any effect that may bé produced upon the endosperm of an angiosperm by pollination with pollen from a plant having a different kind of endosperm. Of course, it is not to be expected that xenia will occur in all such crosses. Correns (3, pp. 411-414) has outlined a number of cases in which xenia will or will not occur in maize, and East and Hayes (7, p. 103) have condensed all the available data into a law of xenia. For our purpose, it may be said that xenia will occur in any cross in which the male parent possesses the dominant and the female the recessive of an allelomorphic pair of endosperm characters, or when the two parents possess respectively two characters whose interaction is necessary for the production of a visible effect. When dominance is incomplete, or when inhibiting factors are present, complications are introduced which need not be discussed here. THE ENDOSPERM OF ANGIOSPERMS The variability of the maize endosperm, with the accompanying phenomenon of xenia, gives it a prominent place in any explana- tion of the endosperm of angiosperms. This tissue has been ex- plained in many ways, but its true significance still offers an un- solved problem. The correct explanation, if it is ever found, will probably develop from researches on the phylogenetic origin of the angiosperms. In most of the plants in which fecundation has been studied, one sperm has been found to enter into the constitution of the endosperm, and the phenomenon is believed to be of general oc- currence. But there are many exceptions which serve to com- WEATHERWAX: GAMETOGENESIS IN ZEA Mays 83 plicate the problem. A good résumé if these is given by Coulter and Chamberlain (5, pp. 165-186), and more are being added from time to time. Miss Sargant (17, p. 702) stated clearly the problems involved in the interpretation of the triple fusion and reviewed the explana- tions that had been given up to 1900. Although her paper ap- peared very early in the investigation of the question from the modern point of view, it showed such a clear insight into the nature of the problem that the data that have accumulated since that time have afforded little foundation for further constructive work. Some have considered the endosperm merely a belated pro- thallium, the triple fusion of nuclei being non-sexual and of no more significance than the nuclear fusions that often occur in vegetative tissue. Strasburger (18, p. 308) calls the triple fusion a ‘‘vegetative fertilization,” as contrasted with the ‘‘generative fertilization” of the egg. These types of fecundation resemble each other in the transmission of certain hereditary characters, but they differ in the nature of the new individuals produced. Again (17, pp. 704-706), the endosperm may be looked upon as a monstrous embryo, its aberrant nature and limited development being determined by the antipodal one of the polar nuclei, inas- much as its chromosome number is known to be irregular in many plants. According to this view, the endosperm is neither sporo- phyte nor gametophyte, but a new generation, characteristic of the angiosperms alone, and for which Trelease (19) has proposed the name xeniophyte, because it is the generation in which xenia may occur. All agree upon the function of the endosperm as an organ of nourishment for the embryo, and its rapid growth has been attrib- uted to its sexual, or pseudo-sexual, origin. Collins and Kempton (2) and Jones (14) have even shown that hybrid endosperms, produced by crossing varieties of maize that have been selfed for some time, show a perceptible increase in size over those of the parent races. This is attributed to the increased vigor usually resulting from hybridization. Miss Sargant (17, pp. 709-710) has suggested that the introduction of the paternal elements into the primary endosperm nucleus may be a device of the plant to pro- 84 WEATHERWAX: GAMETOGENESIS IN ZEA Mays duce a food of proper quality for the hybrid embryo. In the light of later work on xenia, we should be compelled to infer from this that the peculiar endosperm qualities necessary in such cases were the dominant characters, since we have no evidence that, ex- cept in special cases to be mentioned later, the recessive characters of the male are ever effective in the endosperm; they are not visible, and there is no future generation in which they might be detected. Moreover, in a cross between sweet and starchy varieties, the _ former being the female, there is produced a starchy endosperm, which is harder to digest than the sweet one that would have been produced had the triple fusion not occurred. The part played by the maize endosperm, then, is to complicate the problem and at the same time act as a check on our solutions. Were it not for xenia and the attendant hereditary phenomena, the logical disposal of the endosperm would be to call it gameto- phytic tissue resulting from a triple vegetative fusion to which no great significance could be attached: and this would probably have been done long ago had the peculiarities of the maize endosperm not been known. But the transmission of hereditary characters to this tissue, as illustrated in maize and a few other species, strongly suggests its parallelism with the sporophyte and saves us from an incorrect explanation of the ques ion. HEREDITY OF ENDOSPERM CHARACTERS Without committing ourselves as to the most logical interpre- tation of the endosperm in general, we may, for the purpose of ex- plaining genetic data, adopt the convenient expedient of consid- ering the endosperm of maize a monstrous sporophyte, a sort of sister of the embryo. It derives a vigorous growth stimulus from the triple fusion in which it originates. It passes through a series of tissue differentiations, none of which, however, resemble very closely those characteristic of the embryo. It never reaches sexual maturity, and, consequently, has no descendants. Because of the sporophytic nature of the endosperm, Mendelian principles of heredity have been applied to its study in maize, with results unique in many instances, and all dependent, more or less, upon the cytological facts here set forth, or, at any rate, nor at variance with them. WEATHERWAX: GAMETOGENESIS IN ZEA MAys 85 _ The maize endosperm is either sweet or starchy. The starchy tissue occurs in two forms, one corneous and translucent, and the other soft and white; and different proportions and variations in the arrangement of the soft and corneous portions gives rise to dent, flint, pop, and soft types. The mature seeds of the sweet varieties are always wrinkled; they are to be understood as having pop, dent, flint, and soft potentialities remaining invisible because of limited starch development. The starchy condition is dominant to the non-starchy, but dominance among the variations of the starchy condition is a less definite thing. The corneous endosperm is either white or yellow; the yellow color, which has been found to be due to more than one hereditary factor, is dominant. It may appear in either the starchy or the sweet endosperm. The aleurone is red, purple, or colorless. Two hereditary factors are necessary for the production of the red color, and these, interacting with a third, produce purple. The presence of either color combination is dominant to its absence, but other color genes are also present, at least one of which is an inhibiting factor. Correns found (4) in certain crosses that a white variety pol- lenized with pollen from a purple did not always produce purple aleurone, although the embryos of the same seeds proved to be hybrids. He explained this non-appearance of xenia by assuming that the recessive factors carried by the two maternal nuclei entering into the primary endosperm nucleus were dominant over the one factor introduced by the sperm, although the latter was ordinarily dominant. East and Hayes (7, pp. 58-59), having found a better explana- tion of this aberrant result, attacked Correns’s hypothesis on the ground that quality and not quantity of chromatin is the deter- mining factor. In this argument they failed to distinguish between the idea of double quantity and quality acting twice. In later experiments (12, p. 12) they found that when reciprocal crosses were made between a soft and a corneous variety, the quality of the endosperm produced was always determined by the female parent. At the same time it was shown that a soft white female crossed with a flinty yellow male produced soft yellow seeds. The appearance of xenia in color showed that the peculiar 86 WEATHERWAX: GAMETOGENESIS IN ZEA Mays results of the reciprocal crosses were not due to a failure of the sperm to fuse with the polar nuclei, as Webber had suggested (25, pp. 34-37) in explanation of a similar occurrence. Therefore, they were forced to conclude that two applications of the one factor may dominate one application of the other. Results leading to the same conclusions have since been se- cured with color combinations. If a plant heterozygous for purple aleurone be selfed, four types of endosperm should be produced, depending upon the number of times the factor combi- nation for purple is present. And when such an experiment is made, there are produced, besides the proper number of white seeds, visibly different types of purples, appearing in significant ratios. Emerson (8) has recently given a detailed report on a number of experiments of this kind. It may be said, therefore, that the solution proposed by Cor- rens was a valid one, although it did not apply to the problem that he had in hand. By taking advantage of the triple parentage of the endosperm, this series of experiments provided the first direct evidence of the cumulative effect of repeated applications of the same factor, which is the basis of the multiple factor hypothesis. But these theoretical conclusions have all been based upon the assumption of certain cytological facts not hitherto demon- strated. If, in the formation of the maize embryo sac, all four megaspores functioned, which the writer (24, p. 492) at one time thought probable, the two polar nuclei, coming respectively from the two nuclei resulting from the heterotypic division, would be genetically different in a hybrid plant. Therefore, a sweet-starchy hybrid, crossed with pure sweet, should produce all starchy seeds, since the two polar nuclei together would represent an entity unaffected by segregation and always carrying the dominant factor. But, as a matter of fact, when a cross of this kind is made,ai:!I ratio is produced. The assumption that only one megaspore survived had good precedent in Koernicke’s work (15) on Triticum, recently verified by Jensen (13); but now it is definitely known that the same is true of maize. _ In some crosses of white varieties with purple or red, a mottled aleurone is produced; and in sweet-strachy crosses a few seeds have been found with the endosperm bilaterally divided—one WEATHERWAX: GAMETOGENESIS IN ZEA MAys 87 half starchy and one half sweet. Webber (25, pp. 34-37) found an explanation for such irregularities.in the assumption that the sperm might sometimes fail to fuse with the polar nuclei, and thus be left to divide independently, or might fuse with only one of them, the other dividing independently. In either case there would be two, possibly genetically different, sources of endosperm formation, which might account for the two colors of aleurone. It has since been found that aleurone mosaics are amenable to Mendelian principles, but for the divided endosperms East and Hayes (7, pp. 34-35) consider Correns’s explanation well-founded. This assumption requires that either the sperm or one of the polar nuclei, both of which have been considered gametic in nature, develop without uniting with another gamete, which would be a very unusual behavior. If such an explanation of the occurrence is necessary, it seems more reasonable to suppose that the sperm united with the polar nucleus coming from the micropylar end of the embryo sac, since it is the sister of the egg and necessarily much like a gamete, and that the antipodal one of the polar nuclei, with its possibly irregular chromatin organization, divided inde- pendently, as its near relatives, the true antipodals, regularly do in maize and other grasses. Yet this would take from one polar nucleus much of its gamete-like nature and might seriously inter- fere with its ability to transmit hereditary characters in the ortho- dox fashion necessary for some of the other phases of our theory. In fact, any assumption of this kind will lead to the conclusion that something, elsewhere considered gametophyte, here contri- butes directly to endosperm formation and shows endosperm (xeniophyte) characteristics. : The responsibility for these rare occurrences of bilaterally differentiated seeds may be placed upon still another hypothesis which involves valid cytology and is not in conflict with Mendelism. The division of the primary endosperm nucleus may be heterotypic in a way. In fact, it is well known that the subsequent divisions often show fewer than 3x chromosomes; some chromosomes must, therefore, at some time, pass bodily to the poles of the spindle without themselves dividing, and, if one of these carries the gene for starchiness, reduction necessitates a sort of Mendelian segre- gation. 88 WEATHERWAX: GAMETOGENESIS IN ZEA MAys In the frequency with which these bilaterally differentiated endosperms occur—about one in 10,000—East (6, p. 220) has likened them to ‘‘bud sports.’” This explanation is not materially different from the one suggested above, although the cytologica phenomena accompanying the formation of “bud sports’’ are not well understood at present. In the endosperm, where chro- matin behavior seems much less regular than in ordinary embryonic tissue, the chances for such occurrence seem exceptionally good; but, neither in the division of the primary endosperm nucleus nor in the arrangement of the free nuclei in subsequent stages, is there anything to account for the perfect symmetry and fine line of demarcation characterizing these seeds. SUMMARY Of the four potential megaspores in maize, only one persists and becomes functional. The embryo sac is a modified form of the ordinary seven-celled type. The antipodals undergo division, forming a large mass of tissue before the time of fecundation. The sperms, which are exceedingly small, are matured inside the pollen grain before the dehiscence of the anther. External conditions and the length of the silk determine the time required for the growth of the pollen tube, but fecundation is known to have taken place in some instances within a little more than twenty-four hours after pollination. Guignard’s work on double fecundation in maize is verified. Both sperms are functional, one uniting with the egg to give rise to the embryo, and the other entering into the constitution of the primary endosperm nucleus. This is the cytological basis of xenia and the attendant hereditary phenomena. Immediate effects of pollination made manifest outside the embryo, if such really occur, should not be called xenia. The triple fusion takes place almost simultaneously with fecundation of the egg, but the endosperm develops much more rapidly than the embryo. In connection with genetic studies, the assumption that the endosperm is sporophytic in its genetic behavior seems consistent. The cytological data substantiate practically all the assumptions WEATHERWAX: GAMETOGENESIS IN ZEA MAys 89 of cytological fact upon which recent experiments on endosperm heredity in maize have been based. I take opportunity here to express my obligations to Professor D. M. Mottier for the suggestion of this problem and for valuable assistance and criticism in connection with the study. = NS w on - ~] INDIANA UNIVERSITY LITERATURE CITED . Bailey, L. H., & Gilbert, A. W. Plant-breeding. New York. 1915. . Collins, G. N., & Weniepteii J. H. Effects of cross pollination on the size of seed in maize. U.S. Dept. Agr. Circ. 124. 1913. Correns,C. Untersuchungen iiber die Xenien bei Zea Mays. Ber. Deutsch. Bot. Ges. 17: 410-417. 1899 Bastarde zwischen Maisrassen mit besonderer Beriick- sichtigung der Xenien. Bibl. Bot. 53: 1-161. pl. 1, 2. 1901. . Coulter, J. M., & Chamberlain, C. J. Morphology of angiosperms. New York. 1903. East, E. M. Xenia and the endosperm of angiosperms. Bot. Gaz. 56: 217-224. 1913. . East, E. M., & Hayes, H. K. Inheritance in maize. Connecticut Agr. Exp. Sta. Bull. 167. 1915. . Emerson, R. A. A fifth pair of factors, Aa, for aleurone color in maize, and its relation to the Cc and Rr pairs. Cornell Agr. Exp. Sta. Mem. 16. 1918. . Focke, W. O. Die Pflanzenmischlinge. Berlin. 1881. . Guignard, L. La double fécondation dans le mais. Jour. de Bot. 15: 37-50. 1901. . Hackel, E. True grasses. English translation. New York. 1890. 9 . Hayes, H. K., & East, E. M. Further experiments on inheritance in maize. Connecticut Agr. Exp. Sta. Bull. 188. 1915. . Jensen, G. H. Studies on the morphology of wheat. Washington Agr. Exp. Sta. Bull. 150. 1918. . Jones, D. F. Bearing of heterosis upon double fertilization. Bot. Gaz. 65: 324-333. 1918. . Koernicke,M. Untersuchung iiber die Entstehung und Entwick- lung es Sexualorgane von Triticum mit besonderer Beriick- sichtigung der Kerntheilungen. Verhandd. Naturhist. Ver. Preuss. Rheinl. 53: 149-185. pl. 5 +f. A-C. 1896 . Nawaschin, S. Neue Beobachtungen iiber Befruchtung bei Frit- illaria tenella und Lilium Martagon. Bot. Cent. 77: 62. 1899. 90 WEATHERWAX: GAMETOGENESIS IN ZEA MAys 17. Sargant, E. Recent work on the results of fertilization in angio- sperms. Ann. Bot. 14: 689-712. 1900. 18. Strasburger, E. Einige Bemerkungen zur Frage nach der “doppel- ten Befruchtung” bei den Angiospermen. Bot. Zeit. 58: 293- 316. 1900. 19. Trelease, W. Two new terms, cormophytaster and xeniophyte, axiomatically fundamental in botany. Proc. Am. Phil. Soc. 55: 237-242. 1916. 20. True, R. H. On the development of the caryopsis. Bot. Gaz. 18: 212-226. pl. 24-26. 18093. . de Vilmorin, P. L. A hybrid Froth teosinte and corn. Bull. Soc. Bot. France. 54: 39-42. pl. I. 1907 22. Waller, A, E. Xenia and other influences following fertilization. Ohio Jour. Sci. 17: 273-284. 1917. 23. Weatherwax, P. Morphology of the flowers of Zea Mays. Bull. Torrey Club. 43: 127-144. pl. 5,6 +f. 1-4. 1916. 24. —_———._ The development of the spikelets of Zea Mays. Bull. Torrey Club 44: 483-496. pl. 23 + f. I-33. 1917. 25. Webber, H. J. Xenia, or the immediate effect of pollen in maize. U. S. Dept. Agr. Div. Veg. Phys. and Path. Bull. 22. 1900. Explanation of plates 6 and 7 PLATE 6 All figures X 725 Fic. 1. The archesporial cell. Fic. 2. Megaspore ca and disorganized tapetal cell. Fic. 3. Megaspore mother voll in synapsis Fic. 4. Heterotypic spindle. Fic. 5. Dau pet gum pone from reduction division. Fic. 6. aspores, ss, and micropylar ster cell in process of division. Tw Fic. 7. Sie daughter cell and one megaspo e beginning to disintegrate: s, functional megaspore. Fic, 8. Functional megaspore absorbing the other three. Fics. 9-12. Steps in the germination of the megaspore and organization of the embryo sac: ss, synergids; e, egg nucleus; ~~, polar nuclei; a, antipodal cells. PLATE 7 Fic. 13. Embryo sac soon after organization, X 725: ss, synergids (position shown by dotted lines); e, egg nucleus; , polar nuclei; a, antipodal tissue. Fic. 14. Germinating pollen n grain on stigma hairs, X 200 Fic. 15. Sperm, X 1750. Fic. 16. Pollen tube entering embryo sac, X 385: 0, outer integument; #7, inner integument; tf, pollen tube; w, ovary wall; s, sperms; , tube nucleus (probably); e, egg nucleus; ?, polar nuclei in contact; a, one synergid; other synergid not shown. Fic Sperm nucleus, s, in contact with egg nucleus, X 1500 Fie: 18, 19. Sperm nuclei, ss, in contact with the fusing polar mack. X 1500. The effect of soaking in water and of aération on the growth of Zea Mays F. M. ANDREWS AND C. C. BEALS (WITH FIVE TEXT FIGURES) The first of these experiments was to ascertain the best length of time required to soak the grains of Zea Mays for growth to begin. The grains we used were one year old. Some experiments of this kind have been made,* but since the time varies to some extent in the grains of Zea Mays of different kinds and ages, it was de- sired to determine it for the material at our disposal. The second object was to determine the effects of aération on the seedlings of Zea Mays under different conditions. 1. EXPERIMENTS IN SOAKING GRAINS OF ZEA Mays a. Time required for maximum growth In order to ascertain the proper length of time to soak Zea Mays for maximum growth the grains were kept in tap water for varying lengths of time, and then in wet sawdust long enough to total forty-eight hours. The experiments were kept under normal growing conditions. Tastes I-IV give the average length of growth of five grains. The unsoaked grain used as a control made no external growth in any of the experiments. TABLE I TABLE II Hours Growth Hours Growth soaked mm. in mm. RO pi ha oil oe eee 8.6 Ee Parr 5.2 E97, ies So a a 10.8 Wr ies axes 11.2 SOc eee os 12.6 NOG ai a ee II.5 OT Cees ei ee 10.2 co haere rey 13-4 Pt, patna ean es 4.6 cP a eas eT Pas 11.6 Be ae eee ws 3-0 GA coyeweyeewn ee 4.8 Kors geen a3 II. Uredinia amphigenous on brownish or purplish spots, early or somewhat tardily naked, cinnamon-brown; urediniospores globoid or broadly ellipsoid, 23-32 by 27-35; wall cinnamon- brown, rather thick, 1.5-2.5 u, closely and finely echinulate, the pores prominent, equatorial, 3 or 4, covered with swollen, hyaline cuticle. III. Telia disposed similarly to the uredinia, chestnut-brown; teliospores oblong or clavate, 18-26 by 37-48 u, rounded at both ends or slightly narrowed below, usually not constricted at septum; wall dark chestnut-brown above, lighter below, 1.5-2.5 u thick, thickened 6-8 4 at apex, sometimes also thickened at one side; pedicel golden-brown, short. On Leptoloma cognatum (Schultes) Chase (Panicum cognatum ~ Schultes), Auburn, Lee County, Alabama, August 31, 1890, II, iii, Geo. F. Atkinson 1586 (type); Ellsworth County, Kansas, July 27, 18096, II, iii, C. H. Thompson; Austin, Texas, February 27, 1901, II, W. H. Long, Jr. 82; Austin, Texas, November 10, 1914, II, Lewis & Tharp 4z; Stillwater, Oklahoma, July 27, 1915, II, C. D. Learn 128; Austin, Texas, October 29, 1915, II, II, B. C. Tharp. The type collection had the host given as probably Muhlenbergia diffusa. The material is scanty and without in- florescence. Its identity was ascertained through the painstaking examination made by Mrs. Agnes Chase. On July 16, 1915, she wrote; ‘The specimen marked ‘Muhlenbergia diffusa’ I make Leptoloma cognatum. They have the same texture and ligule, and the little erect bit of firm tissue on either side at the summit of the sheath, where in aged leaves it tears loose from the thin ligule as found in Leptoloma.’’ The teliospores were dis- covered on type material in April, 1911, but it was not until the host was positively identified that the relationship of the rust could be worked out. Puccinia Cockerelliana Bethel, sp. nov. O. Pycnia amphigenous, few, 107-128 » broad. I. Aecia hypophyllous or caulicolous, in small groups 3 mm. or less across, low cupulate; peridium soon disappearing; peridial cells rhombic, 16-23 by 27-35, the outer wall thick, 6-8 x, smooth, the inner wall thinner, 2—3 », coarsely verrucose; aecio- spores angularly globoid, 18-24 by 20-29 »; wall nearly or quite colorless, moderately thick, 1.5-2.5 , finely and evenly verrucose. II. Uredinia epiphyllous, intercostal, oblong-linear, 0.5~I 114 ARTHUR: NEW SPECIES OF UREDINEAE mm. long, cinnamon-brown; urediniospores ellipsoid, 19-26 by 24-32 w; wall colorless or nearly so, 1.5-2 yu thick, finely echinulate, the pores obscure, probably scattered. II. Telia similar to the uredinia, long covered by the epidermis, becoming dehiscent by a longitudinal slit, grayish-black; telio- spores not surrounded by stromal hyphae, cylindric or clavate- cylindric, 13-21 by 50-85 wu, rounded or truncate above, tapering to base, the upper cell about one third length of spore, slightly or not constricted at septum; wall chestnut-brown above, paler below, thin, - about I uw, moderately thickened above, 3-7 u; pedicel short, tinted. On Thalictrum Fendleri Engelm., Gunnison County, Colorado, September 2, 1899, I, E. Bartholomew; Eldora, Colorado, 9,000 feet alt., July 25, 1910, I, E. Bethel (Barth. N. Am. Ured. 6176); Eldora, Colorado, June 24, and July 2, 1911, I, E. Bethel; Trout Lake, 10,000 feet alt., August 2, 1912, I, F. D. Kern 5107; Trimble Springs, nine miles from Durango, Colorado, 7,500 feet alt., August 4, 1912, I, F. D. Kern 5303. On Festuca Thurberi Vasey, Eldora, Colorado, 9,000 feet alt., September 17, 1910, III; same, May 20, June 24, July 4 (type), October 7, 1911, III; same, June 30, 1912, III; same, September 19, 1914, III; same, July 22, 1916, III, all collected by E. Bethel; same, August 25, 1911, II, Bethel & Kern; Fremont Station near Manitou, Colorado, August 25, 1916, II, III, J. M. Bates 6486. As early as 1910 Mr. Bethel wrote in transmitting specimens that these forms of rust on Thalictrum and Festuca, found growing together and apparently genetically connected, were noticeably different from the forms belonging under Puccinia Clematidis. He then transplanted healthy plants of both hosts to his garden in Denver, and the following year began a series of cultures, using both aeciospores and teliospores, which has extended to the present time. Some of the results of these numerous trials have been transmitted to the writer; the publication of a full account of the work, however, is contemplated by Mr. Bethel, who has supplied the name and nearly all the information regarding the species, and it is deemed neither necessary nor courteous to give more than a bare statement in this connection. Mr. Bethel also sent telial culture material several times to the writer, from which only one successful germination of spores was obtained. A culture followed, the result being recorded in a report of cultures for 1915 (Mycologia 8: 132. 1916) under the name Puccinia A gropyrt. ARTHUR: NEW SPECIES OF UREDINEAE 115 Puccinia Cockerelliana is conspicuously different in both gross and microscopic appearance from P. Agropyri, which occurs on the same and other similar hosts. The aecia are more crowded, more robust and more inclined to be bullate; the aeciopsores are larger and have much thicker walls. The telia are on the rough or morphologically upper side of the leaf and situated between the veins, instead of on the smooth side of the leaf or on sheaths and stems asin P. Agropyri. The telia are also larger, and become normally dehiscent and uncovered after a time. The teliospores are very long and slender, and have no enveloping stromal mass, as commonly found in strictly subepidermal forms. The species is named by Mr. Bethel in recognition of the emi- nent services rendered science in many fields by Professor T. D. A. Cockerell, of the University of Colorado, Boulder, Colorado, and especially to recall his help in studying the flora of Colorado, including the rusts and other fungi. Puccinia inclita sp. nov. II. Uredinia amphigenous, tardily naked, brownish-yellow; urediniospores globoid or ellipsoid, 22-26 by 24-32 #; wall pale yellow or brownish-yellow, thin, I-2 u, coarsely and sparsely echinulate with elongated and sharply pointed projections, the pores obscure, doubtless 3 and approximately equatorial. III. Telia similar to the uredinia, chocolate-brown; teliospores broadly ellipsoid or oblong, 26-29 by 35-40 #, rounded at both ends, very slightly or not constricted at septum, often with meso- spores intermixed, 23-27 by 26-29 »; wall chestnut-brown, 2.5- 3.5 « thick in upper cell and somewhat thinner in lower cell, thicker above, 3-6 u and often slightly lighter in color, smooth; pedicel golden-brown or paler, the diameter uniform, about one and one half times length of spore. On Ichnanthus pallens (Sw.) Munro, Mayagiez, Porto Rico, March 2, 1916, II, Whetzel & Olive 396; El Yunque, Porto Rico, April 12, 1916, II, II], Whetzel & Olive 397 (type). On Oplismenus hirtellus (L.) R. & S., Las Marias, Porto Rico, July 10, 1915, Il, F. L. Stevens 8118. The striking appearance of the urediniospores with their prominent echinulation caused the first collection studied, that by Professor Stevens on Oplismenus, to be referred to the South American Uredo Olyrae P. Henn. (see Mycologia 8: 21. 1916), but 116 ARTHUR: NEW SPECIES OF UREDINEAE it wasafterward placed under Uredo paspalicola P. Henn. (see Myco- logiag: 92. 1917). The other two collections, those by Whetzel & Olive on Ichnanthus, were placed under Puccinia substriata Ellis & Barth. (see Mycologia 9: 73. 1917),a species now consid- ered to include Uredo paspalicola. More intimate and extended - comparisons, however, have led to the belief that this form should be separated from the widely distributed and variable P. substriata. The most characteristic features are the large and usually pale urediniospores, with their thin walls, sparsely covered with spine- like points. The spores were at first thought to be thick-walled (see Mycologia 8: 22. 1916),an error due, as in many other cases, to mistaking the dense ectoplasm of the cell as part of the wall. Puccinia Coelopleuri sp. nov. O. Pycnia amphigenous or petiolicolous, crowded in groups I-2 mm. in diameter, noticeable, subepidermal, globoid, 96—112 u in diameter; ostiolar filaments short. II. Uredinia amphigenous; the primary form sometimes peti- olicolous, crowded, circinating about the pycnia in round or oblong groups 2-10 mm. long, oblong, 0.5-4 mm. long, early naked, pulverulent, cinnamon-brown, ruptured epidermis conspicuous, the secondary form scattered over the surface of the leaves unaccom- » panied by pycnia, oval, 0.2-0.5 mm. long, otherwise like the pri- mary form; urediniospores broadly obovoid or ellipsoid, 22-26 by 27-35 u; wall cinnamon-brown, 2~-2.5 u thick, much thicker above, 3-7 u, moderately echinulate, the pores 3, equatorial, covered by the colorless swollen cuticle. III. Telia mostly hypophyllous, scattered, oval, 0.2-0.5 mm. long, early naked, somewhat pulverulent, chocolate-brown, rup- tured epidermis evident; teliospores cylindric, oblong or oblong- clavate, 15-20 by 37-63 », rounded at both ends, or slightly nar- rowed below, not or slightly constricted at septum; wall dark cinnamon-brown, thin, 1-2y, slightly thickened at apex, 3-4n, smooth; pedicel colorless, fragile, short. On Coelopleurum Gmelini (DC.) Ledeb., Juneau, June 20, 1917, No. 355, July 18, 1918, No. 481; Mendenhall, June 24, 1917, No. 366, September 9, 1917, No. 387 (type), August 3, 1918, No. 487; Haines, August 21, 1918, No. soz, all from Alaska, and collected by J. P. Anderson. A conspicuous and, on account of its large ~ spores, a striking species. ARTHUR: NEW SPECIES OF UREDINEAE 117 Puccinia parca sp. nov. II. Uredinia hypophyllous, scattered or irregularly grouped, roundish, 0.3-0.6 mm. across, brownish-yellow, pulverulent; urediniospores narrowly ellipsoid or obovoid, 13-16 by 19-26 y; wall brownish- or light-yellow, very thin, 1 u or less, moderately and inconspicuously echinulate, the pores indistinct, 2-4, equa- torial or superequatorial. . II. Telia similar but slightly larger than the uredinia, dark cinnamon-brown, pulverulent; teliospores ellipsoid or oblong, 15-19 by 29-42; wall cinnamon-brown, rather thin, 1-2 4, thickened into a hemispherical hyaline papilla over the pores, 4-5 u, smooth; pedicel colorless, one half length of spore or less, fragile. On Tiniaria scandens (L.) Small (Polygonum scandens L.), Flatbush, Long Island, New York, October 5, 1889, II, III, J. L. Zabriskie 703; Stelton, New Jersey, September 7, 1892, II, III, Byron D. Halsted (Seym. & Earle, Econ. Fungi 367, type); Laurel Springs, northwestern North Carolina, September 20, 1904, II, Ill, H. H. Hume 278. As long ago as October, 1905, Professor Holway called the attention of the writer to the peculiarities of the rust issued by Seymour & Earle in their Economic Fungi under the name of P. mammillata. Professor Holway pointed out that that species has rough spores and of a different shape from this material. He thought it might be a new species, but later in his North American Uredineae (1: 40) placed it doubtfully under P. septentrionalis Juel. P. septentrionalis is a boreal species on Bistorta viviparum, having its aecia on Thalic- trum alpinum, and in America has been taken in Alaska and New- foundland. The rust in question agrees with P. septentrionalis in the character of its teliospores, as Professor Holway pointed out in detail, but differs from it in having slightly narrower uredin- iospores, with thinner walls, of a lighter and more yellowish color. The species is markedly distinct from P. Polygoni A. & S., the common rust on the same and related hosts, both in the teliospores and urediniospores. The pores of the urediniospores are difficult to make out, but are usually three and approximately equatorial, while in the more common P. Polygoni they are distinct and two in the upper part of the spore. 118 ARTHUR: NEW SPECIES OF UREDINEAE Puccinia gentilis sp. nov. II. Uredinia hypophyllous, scattered, round or elliptic, 0.3—-0.8 mm. across, pulverulent, cinnamon-brown; urediniospores oblate- spheroid, 23-26 » broad by 19-24 u long, or globoid to obovoid, 21-23 by 21-28; wall dark cinnamon-brown, 1.5-2 y thick, moderately and strongly echinulate, the pores 2 or 3, subequatorial or approximately equatorial. Ill. Telia hypophyllous, similar to the uredinia, becoming pulverulent, blackish-brown; teliospores broadly ellipsoid, 27-32 y 35-45 mu, rounded above and below, not constricted at septum; wall chocolate-brown, thick, 3-5 wu, thickened over the germ-pore into a yellowish umbo, 7-10 p thick, moderately verrucose with markings uniting into short irregular lines giving a coarsely verru- cose appearance; pedicel colorless, with thin walls, 1 u or less, twice length of spore On Salvia aleansdne Rose, Oaxaca, Mexico, October 21, 1899, II, iii, EZ. W. D. Holway 3699; Salvia sp., Oaxaca, Mexico, October 18, 1899, II, III, E. W. D. Holway 3666 (type). The species is in many respects similar to Puccinia mitrata Syd., but the uredinio- spores are larger and more prominently echinulate. Puccinia prospera sp. nov. II. Uredinia amphigenous, scattered, round, 0.2-0.8 mm. in diameter, pulverulent, cinnamon-brown; urediniospores oblate- spheroid, 25-30 broad by 20-234 long; wall cinnamon-brown, 1.5 thick, moderately and rather strongly echinulate, the pores 2 or 3, subequatorial. III. Telia hypophyllous, similar to the uredinia, pulverulent, chestnut-brown; teliospores broadly ellipsoid, 27-32 by 35-40un, rounded above and below, not constricted at septum; wall chest- nut-brown, rather thin, 1-2.5 4, thickened over the germ-pores, 5-7 u, obscurely verrucose-rugose; pedicel colorless, fragile, once to once and half length of spore, thin-walled, fragile. On Salvia microphylla H. B. K., Toluca, Mexico, September 17, 1898, Il, E. W. D. Holway 3136; Pachuca, Mexico, October 5, 1899, II, III, E. W. D. Holway 3579 (type). Another species of the P. mitrata group, the teliospores being of the same size as those of that species, but with the thickness of wall and its markings like P. farinacea Long. The urediniospores also have the size of P. farinacea. ARTHUR: NEW SPECIES OF UREDINEAE 119 Puccinia massalis sp. nov. O. Pycnia amphigenous and caulicolous, rather conspicuous, subepidermal, 112-120 yu broad. I. Aecia amphigenous and caulicolous, crowded in irregular groups 2-10 mm. across on the blades, often 6-20 mm. long on the stems, petioles and veins, causing distortion of the host, cylindric, ‘0.5-0.8 mm. in diameter, 0.5-1 mm. high; peridium with erect margin, erose or somewhat lacerate; peridial cells rhombic, 19-27 by 32-45 u, the outer wall 7-10 thick, smooth, the inner wall 5-10 yu, closely tul late ; aeciosp ellipsoid, 16-18 by 22-27 u; wall colorless, 1.5-2 thick, very closely and inconspicuously verrucose. II. Uredinia amphigenous, scattered, round, 0.8-1.2 mm. in diameter, pulverulent, cinnamon-brown; urediniospores irregularly ellipsoid, obovate or oblong, 18-24 by 27-35 4; wall light cinna- mon-brown, I—1.5, thick, moderately echinulate, the pores 2, equatorial. - II. Telia similar to the uredinia, pulvinate, chocolate-brown ; teliospores ellipsoid, 23-32 by 39-48 u, rounded above and below, slightly or not constricted at septum; wall dark chestnut-brown, thick, 3-5 u, thickened and lighter-colored over the germ-pores, 9-10 », smooth, the pore of upper cell apical, of lower cell usually half way to hilum or occasionally at septum; pedicel colorless, once to thrice length of spore. : . On Helianthus ciliaris DC., Las Cruces, New Mexico, October 27, 1892, I, III, E. O. Wooton; Ysleta, Texas, February 24, 1914, Ill, Arthur & Fromme 5704; Mesilla Park, New Mexico, October 7, 1914, III, and May 9, 1915, I, W. A. Archer; Albuquerque, New Mexico, December, 1914, III, W. H. Long 5183; Brazito, New Mexico, June 15, 1915, I, II, I], W.A. Archer (type). This is a much more robust species than Puccinia Heliantha Schw., to which most of the collections here cited have been referred (Mycologia 8:159. 1916). The aecia are much larger, and cause swelling of the adjacent tissues, even giving the appearance in some cases of being a systemic rust. Both the aeciospores and teliospores have far thicker walls. Puccinia invelata Jackson, sp. nov. O. Pycnia epiphyllous, gregarious, few, inconspicuous, sub- epidermal, 115 u broad. I. Aecia hypophyllous, solitary or in groups of two or three; peridium lacerate; peridial cells ellipsoid, 14-18 by 27-35 u, over- 120 ARTHUR: NEW SPECIES OF UREDINEAE lapping, the wall 3 » thick; aeciospores ellipsoid, 16-23 by 24-26 yp; wall 1-1.5 » thick, closely verrucose IJ. Uredinia hypophyllous, scattered, roundish, small, 0.2—-0.4 mm. across, moderately pulverulent, cinnamon-brown ; uredinio- spores globoid, ellipsoid or obovoid, 20-26 by 24-29 ; wall cin- namon-brown, 1.5-2 u thick, prominently echinulate, the pores 2, surrounded by an indistinct smooth area, equatorial. Ill. Telia hypophyllous or somewhat amphigenous, scattered, roundish, small, 0.1-0.4 mm. across, early naked, compact, chestnut-brown; teliospores ellipsoid or obovate, often somewhat irregular, 19-26 by 32-45, obtuse or rounded above, more or less narrowed below, slightly constricted at septum; wall cinna- mon-brown, laminate, thin, 1.5-2.5 » thick, much thicker above, 7-10 uw, with distinct and somewhat paler umbo; pedicel colorless, once to once and a half length of spore, usually breaking away. On Verbesina montanoifolia Robs. & Greenm., Patzcuaro, Mexico, October 16, 1898, ii, III, 3000 (Barth. Fungi Columb. 5055); Morelia, Mexico, October 8, 1899, ii, III, 3592 (Barth. N. Am. Ured. 426); Patzcuaro, Mexico, October 10, 1899, O, I, I, III, 3606-7 (type), all collections by E. W. D. Holway. The species differs from P. abrupta Diet. & Holw., which occurs on the same and other species of Verbesina, by the strongly obovate teliospores with somewhat narrowed base, paler and thinner walls, and more fragile pedicel. The Verbesina rusts have recently been studied by Professor H. S. Jackson, who has separated this form as a new species and supplied the name and diagnosis. Uromyces Shearianus nom. nov. Aecidium Atriplicis Shear, Bull. Torrey Club 29: 453. 1902. Uromyces Atriplicis Arth. Bull. Torrey Club 45: 141. 1918. Not U. Atriplicis McAlpine, 1906. In the preceding number of this series the writer transferred the early specific name of this rust to another genus without noticing that it had already been used in that connection for a wholly unlike rust occurring in Australia. Although regretting the unnecessary synonym I am pleased to have the opportunity to dedicate the species to Dr. C. L. Shear, who first described it, and who has done notable service in many ways in the field of mycology. ARTHUR: NEW SPECIES OF UREDINEAE 121 Uredo biporula sp. nov. II. Uredinia hypophyllous, chiefly scattered or somewhat gregarious, orbicular, 0.2—-0.3 mm. in diameter, soon naked, pul- verulent, dark cinnamon-brown; urediniospores triangular-obo- void, 21-23 by 23-26 w; wall dark cinnamon-brown, I-2 yu thick, moderately and finely echinulate, the pores 2, basal, and close to the hilum. On Salvia fulgens Cav., Amecameca, Mexico, October 31, 1899, E. W. D. Holway 3758. This collection, in which the ured- inia are abundant and conspicuous, shows the unique character among Salvia rusts of two basal pores in the urediniospores. Uredo amicosa sp. nov. II. Uredinia hypophyllous, scattered, bullate, 0.2-0.4 mm. in diameter, opening by a central pore; paraphyses thickly im- bricated, the united bases forming a tissue-like lining to the sides of the sorus, the long free ends cylindric or fusiform-cyclindric, 10-16 by 67-112 , acuminate or acute, the wall colorless, thick- ened to nearly or quite obliterate the lumen; urediniospores epedicillate, angularly oblong, ellipsoid, or obovoid, 23-34 by 40-60 u; wall golden-brown, 2-3 u thick, sometimes twice as thick at apex, sparsely and strongly echinulate, the pores obscure, possibly 3 or 4 and equatorial. On Chrysophyllum Cainito L., mesas near Mayagiiez, Porto Rico, March 29, 1917, H. E. Thomas 264. There are many sapotaceous rusts, but this one on the common star apple of the tropics seems to be different from any heretofore described. The rather conspicuous sori are abundantly scattered over the under side of the leathery leaves. The spores may possibly be catenu- late, as no pedicels can be detected, either attached to the spores or at the bottoms of the sori. Two species of Uredo have been described on undetermined species of Chrysophyllum from Brazil, but both of them have much smaller spores and quite different sori, judging from the descriptions, no specimens having been seen. They are Uredo chrysophyllicola P. Henn. (Hedwigia 41: 106. 1902) and U. Chrysophylli Sydow (Hedwigia 49: 78. 1909). i Uredo ignava sp. nov. II. Uredinia amphigenous, numerous on light brown dis- colored areas, roundish or oblong, 0.2-0.6 mm. across, pulverulent, 122 ARTHUR: NEW SPECIES OF UREDINEAE cinnamon-brown; paraphyses peripheral, numerous, hyphoid, incurved, 10-13 by 29-45 y, the wall pale cinnamon-brown or - sometimes colorless, I-1.5 4 thick, frequently thickened 3-5 w on moderately echinulate, the pores obscure, probably 4 and equa- torial. On Bambos vulgaris Schrad., Santiago de las Vegas, Cuba, January 29, 1916, J. R. Johnston 424 (type); Maricao, Porto Rico, March 15, 1916, Whetzel & Olive 428, 429; Mayagiiez, Porto Rico, March 20, 1916, Whetzel & Olive 427. The rust produces an abundance of pale yellowish sori on both sides of the leaf, but especially beneath, and more or less discolors the tissues. It has been referred to Uredo paspalicola (Uredinales of Porto Rico, Mycologia 9: 92. 1917; Uredinales of Cuba, Mem. Torrey Club 17: 165. 1918), which it much resembles. There are a number of bamboo rusts. The material listed here has been carefully compared with Puccinia corticioides Berk. & Br. (Syd. Ured. 1263), P. Kusanoi Diet. (Syd. Ured. 1239, 1313, 1373), P. longicornis Pat. & Har. (Syd. Ured. 2314), and one other species distributed by Kingo Miyabe as P. Phragmitis, all from Japan, and P. Bambusarum (P. Henn.) Arth. from South America, as well as P. Arundinariae Schw. from North America, all on species of Arundinaria or Bambos (or Bambusa as the generic name is often written). It has also been compared with the de- scriptions of other species on these hosts. Aecidium Clemensae sp. nov. OQ. Pycnia epiphyllous chiefly, few, crowded opposite the encircling aecia, noticeable, reddish-brown, subepiderma I. Aecia hypophyllous, crowded in orbicular groups 3-8 mm. across on much larger slightly discolored spots, cupulate or cylin- dric, 0.2-0.4 mm. in diameter; peridium revolute, coarsely lacerate; peridial cells rhomboidal, 32-47 » long, somewhat overlapping; aeciospores angularly globoid or broadly ellipsoid, 19-27 by 24- 27 u; wall colorless, 1-2 u thick, moderately and closely verrucose. On Cissus incisa (Nutt.) Des Moulins, Fort Sill, Oklahoma, June 19, 1916, 11931 (type), July 27, 1916, 11931a, both collected by Mrs. Joseph Clemens. This aecial rust is undoubtedly heteroe- cious. It has some resemblance to the aecia of Puccinia subnitens ARTHUR: NEW SPECIES OF UREDINEAE 123 Diet., but possesses much larger spores. It is a pleasure to have the opportunity to name this rust in honor of an indefatigable and able collector, who has contributed largely to the wealth of material for study in the rusts as well as in many other groups of plants. The material has been secured from many regions during her world-wide travels, and has been most generously placed at the service of students. Aecidium Bourreriae Holway, sp. nov. OQ. Pycnia amphigenous, few in small groups, noticeable, subepidermal, 128-160 u broad. I. Aecia hypophyllous, loosely grouped, short-cylindric, 0.2-0.3 mm. in diameter; peridium fragile; peridial cells rhombic in side view, 13-15 by 23-24u, slightly overlapping, the outer wall 6-7 uw thick, transversely striate, the inner wall 3.5-4.5 thick, closely and rather prominently verrucose; aeciospores globoid or ellipsoid, 19-23 by 23-26 uw; wall colorless, rather thick, I.5-2.5 u, closely and finely verrucose. On Bourreria havanensis Miers, Nassau, New Providence, Bahama Islands, March 2, 1903, E. W. D. Holway. Professor Holway distributed this collection under the name here used, but left the publication of the description to some one else. Aecidium Chamaecristae sp. nov. Aecidium Cassiae Ellis & Kellerm. Trans. Kans. Acad. Sci. 10: 91, hyponym. 1887. Not A. Cassiae Bres. 1891. O. Pycnia amphigenous, subepidermal, noticeable, in small groups, 70—96 uw in diameter. I. Aecia amphigenous, loosely grouped upon reddish spots 2-8 mm. across, cupulate, 0.1-0.2 mm. in diameter, short; peridium recurved, erose; peridial cells rhombic or rhomboidal, 16-23 by 26-35 u, considerably overlapping, the outer wall thick, 9-10 u, transversely striate, smooth, the inner wa! thinner, 4-6 yu, closely verrucose; aeciospores globoid or ellipsoid, 15-19 by 18-25 u; wall colorless, thin, I-1.5 yw, finely and closely verrucose. On Chamaecrista fasciculata (Michx.) Greene (Cassia fasciculata Michx.), Manhattan, Kansas, 1886, W. A. Kellerman (Ellis & Ev., N. Am. Fungi 1825); Lincoln, Nebraska, May 28, 1902, John L. Sheldon; same, May 29, 1902, George G. Hedgcock. The species has apparently never been described, and the name long since chosen was already in use when given. The original collection 124 ARTHUR: NEW SPECIES OF UREDINEAE issued as No. 1825 in Ellis & Everhart’s North American Fungi may be accepted as the type. The collections give the host as Cassia Chamaecrista, which by some taxonomists is considered to be the same as C. fasciculata. The fungus much resembles Aecidium Torae P. Henn., occurring on Cassia Tora in Ceylon and Africa, but varies in seemingly important ways, particularly in having well-developed pycnia, which are absent in the foreign collections examined. Aecidium modestum sp. nov. O. Pycnia Se een inconspicuous, subepidermal, globoid, 160-220 pw in diameter. I. Aecia eautiedisiies in groups 4-10 mm. long, short cylindric, 0.3-0.4 mm. broad; peridium erect, erose or somewhat lacerate; peridial cells seen with difficulty in side view, in face view very irregular in shape, 19-23 by 35-58 u, the outer wall thin, about I-I.5 u, almost smooth, the inner and side walls 2-3» thick, closely and prominently verrucose; aeciospores ellipsoid or oblong, 23-26 by 27-32 uw; wall colorless, 1-1.5 w thick, finely and closely verrucose. On Zephyranthes sp., near Ixmiquilpan, State of Hidalgo, Mexico, 1905, Rose, Painter & Rose 8952. The species differs from A. Zephyranthis Shear, also from Mexico, in possessing larger pycnia and aeciospores, and in having more delicate peridial cells. Aecidium ingenuum sp. nov. O. Pycnia hypophyllous, discoid, inconspicuous, subcuticular. 80-130 uw broad by 40-50 pw high, without ostiolar filaments. I. Aecia hypophyllous, closely packed in rows, nearly cylindric to tongue-shaped, often confluent; peridium erect, 0.5-o.8 mm high, very delicate and fragile; peridial cells in radial section narrowly oblong or linear, 10-16 by 32-42 4, somewhat over- lapping, the outer wall 1-2 w thick, smooth, the inner wall 3-5 u thick, closely verrucose with slender tubercles; aeciospores globoid or broadly ellipsoid, 13-16 by 16-23 yu; wall colorless, 2-3 u thick, half the thickness being due to the close, rather fine and somewhat deciduous tubercles. On Picea canadensis (Mill.) B. S. P., Fish Creek, Wisconsin, June 30, 1913, J. J. Davis; Solon Springs, Wisconsin, June 17, 1914, J. J. Davis; and Walden, Vermont, June 8, 1917, C. L. Orton, communicated by C. R. Orton (type). This is the first ARTHUR: NEW SPECIES OF UREDINEAE 126 Peridermium on Picea having subcuticular pycnia to be found in North America. In Europe one such form is known, which was shown by cultures made in 1916 by Ed. Fischer (Mitt. Nat. Ges. Bern 1916: 131. 1917) to belong to Pucciniastrum sparsum (Wint.) Ed. Fisch., on Arctostaphylos. The American form doubt- less belongs to some species of Pucciniastrum, but probably not to P. sparsum, as the spores are considerably smaller than in the European form of that species. PURDUE UNIVERSITY, LAFAYETTE, INDIANA The preparation and treatment of woods for microscopic study* Forest B. H. Brown (WITH SIX TEXT FIGURES) CONTENTS PAGE Bs INTRODUCTION (05) fi gc so Che ee ea eae ees 127 SUANIOSIMENT OF MICROTOME (© 600 ye eae eee gp a ke ag) 129 3. PREPARATION OF MATERIAL FOR SECTIONING .......-..-0-0-ccucecvcecece 132 BC CULLING THE BOOCKS (0075 44 oo ee ee ee. 132 By REMOVING (SAE BER er gs cate eS et ts eae ree dee rl pees 133 Ci; BOR TERING (THE TISSUM Ss |) Oia ees oa et ee es pee 133 Disc TOBIN ridin ease aca ei Pues ait Ee Lk ote i bd CS Gece 135 Ee SECTIONING WITH. THE MICROGTOME 23.05 6s 25 ln oe cc Nh 136 HARD GRETIONS ho sta) oar eee AP tear gs 5 ok By 137 4. bikinis OF Anat BP IAACERATION SS ig ho eee er as 137 S--LERINITION OF ANATOMICAL DETAIES) 3) 20550 tissy Bh Fee a. 138 EERE STAINES 5 oat OMe es ei RG Lh We ae ig 138 B. DUE RD ENTIAL GUVRNCTION 0p wie fi ee Pea ee i eG I4t 6. ccknath eis ee ee ee ee I4I : Pi CREE CLO S re ea a a ite, Eis ee ae 141 els MSIE SO CRNBr SEG NR ct een hing aie ete A haere chk Waimea dds cw ig bay ate 142 GUMS ei es rede geete teh cats Ok suc Ve eas ed aces 142 D. ESSENTIAL OILS, RESINS, AND GUM-RESINS..........5..-0¢0 ce ceeus 146 ee PAE Ne ae eo M Ee hice cee ound Os Mey Ms SEE eas wala woe 146 Bris ENING oc. er ae oe NPR Sage Way See tg Me ah lar ges yeu Dy 4 aun pew 6 @laay 146 ee eh, CRUSE RE Sy eee pas eta g Wits ok pile b Woe Ge viele ob 146 HO CHRMICAL (GROWTH -RINGS oe ey Sie tLe ee eos Sele bee ane 147 PROG EC RMMIRATION LEST. eo ig eo oy ee os ke eve 148 er (Sita rem pee ee ee ea os a eg ae pa aye 149 1. INTRODUCTION By reason of extreme variation in the mineral, resin- or gum- like content, and in the texture, hardness, and other properties peculiar to the stem tissues of tropical woody plants, the task of preparing these for microscopic study is ordinarily a difficult one, particularly in the case of an extensive series composed of numerous unrelated species of widely differing ecological types. The pub- lished technique relating to the preparation of wood for slides has been worked out primarily from a study of the comparatively soft * Contribution from the Osborn Botanical Laboratory. 127 128 BROWN: Woops FOR MICROSCOPIC STUDY tissues of the woods of temperate regions, to which anatomical work has, in large part, been confined. Such tissues often cut satisfactorily without treatment, while even oak and hickory, among the hardest of such woods, if taken from fresh (green) material, may be satisfactorily sectioned along any plane, by a microtome, without any preparatory treatment. Tropical wood collections, on the other hand, ordinarily embrace a high proportion of species the tissues of which, particularly when selected from dry material, can only be cut after a more or less prolonged treat- ment, the nature of which varies considerably according to the structural features of the species or type. Moreover, after sections of sufficient clearness have been obtained, to an even greater degree than in temperate woods, there are many anatomical characters, often of the greatest interest, which cannot be satis- factorily observed without the use of reagents, stains, or media of definite refractive properties. To meet the particular needs in this field, there is insufficient information at hand, and, further, little seems to have been added in recent years. Papers marking distinct progress in anatomical work too often neglect to publish essential details in the methods employed. During the years 1916-1918, the writer carried out an extended investigation of the woods of Hawaii. In the course of these studies it was found necessary to devote a very considerable amount of the time to the preparation of slides, and altogether more than three thousand permanent mounts were made, including approximately one hundred macerations. In connection with this work, various accepted methods were tested and a number of new ones were devised. In the present paper, the technique employed is described in some detail, particular attention being called to certain heretofore undescribed methods of treat- ment and to a number of improvements on the methods in in common use. While emphasis is thus placed upon the treat- ment of the highly complex woody tissues of tropical dicotyledons, the suggestions embodied should be of value in the treatment of other woods than tropical, or of other than woody tissues, in that, to a certain extent, the treatment of some of the cellulose tissues of the stem, particularly the collenchyma and phloem layers, has been included. * BROWN: Woops FOR MICROSCOPIC STUDY 129 The writer is indebted to Professors J. W. Toumey, S. J. Record and others, of the Yale School of Forestry, and to Pro- fessors A. W. Evans and G. E. Nichols, of the Department of Botany, for the use of authentic material, for the facilities essen- tial for accurate work, and for many helpful suggestions and criticisms. 2. ADJUSTMENT OF MICROTOME For sectioning, Thomson’s modification of the Jung-Thoma sliding microtome (’10), supplied with Walb blades 170 mm. in length and 35 mm. in width, was used. To secure the best results, it was found essential that careful attention be paid both to the sharpening of the knife and to its adjustment on the carriage of the microtome. Briefly, the knife was first sharpened to a wedge- shaped edge of which the two planes were inclined at an angle 6 (Fic. 1) of 20° to each other. At the same time, perfect axial Fic. 1. Diagram to show edge of sliding microtome knife cutting section 10 ¢ thick. § (representing downward inclination) = 4.° @ (representing acuteness of edge after being ground with back raised by cylindrical clamp) = 20. alignment of the cutting edge was secured in the manner to be described presently. In mounting on the carriage, the blade was first inclined downward so as to form an angle 6 (Fic. 1) of 4° between the lower plane (OC) of the knife wedge and OE, the plane in which a given tissue is to be sectioned (vertical incli- nation). The knife carriage was then revolved in a horizontal 130 Brown: WooDs FOR MICROSCOPIC STUDY plane so that the edge of the blade lay at a definite angle with reference to the direction of movement of the carriage on the slide- way. For general work, a horizontal angle of inclination of about 20° was found satisfactory, but to obtain extremely thin sections, as in imbedded material, an angle of about 5° proved to be best. Since angle 6’ (Fic. 1), which the planes of the average micro- tome knife make with one another, is ordinarily more acute than at the edge honed to 20°, this difference must be allowed for in adjusting the vernier. Thus if 0’ is 13°, the vernier should read- 7.5° to give an inclination of 4°, an excellent angle for most work. A few woods containing gummy or resin-like material cut better at a somewhat greater inclination, as 6°. On the other hand, if the blade tends to ‘‘pull into”’ the tissue, a more acute inclination may be needed. It is doubtful if cutting edges with 6 less than 20° should ever be used in cutting woody tissues. The more acute edge is too easily injured to be used in cutting many of the harder woods, such as Pandanus, even when these have been carefully treated. The 20° edge, on the other hand, has sufficient strength in blades of good quality to appear undamaged under the microscope after use in cutting any of the treated material, and at the same time it is sufficiently acute to meet every requirement. The chief objection to using a more obtuse edge is that the section, especially at steep inclinations of the blade, is too sharply bent at O, Fic. 1, in cutting, so that it tends to curl or even roll up. It is partly for this reason, also, that the downward inclination of the knife (angle 8) should not be greater than necessary. But even if the inclination is correct, the quality of the work which the instrument is capable of doing may be seriously im- paired by an imperfectly aligned cutting edge. This, of course, is a point which does not have to be taken into account at all in using a rotary microtome. For the sliding microtome, it is not sufficient that the edge shall coincide with a single plane, as OD, Fic. 1. Errors of alignment from heel to point may still be present by reason of which an otherwise well sharpened edge would not coincide with a plane passing through O at right angles to OD. Here an error so slight as to escape detection by the eye may be the entire cause for poor work. For example, let the error of BROWN: WOoOoDS FOR MICROSCOPIC STUDY 131 alignment, 7, midway between heel and point be 0.3 mm. with reference to a plane meeting the bisecting plane OD at right angles and in contact with heel and point. Let angle 6 = 20°, angle a = 10°, angle 6 = 4°, and angle a+ 8 = 14°. Distance y = r sin (a + 8), representing the error of alignment of the knife edge with reference to the plane of section, will then be 72 u. In other words, when heel and point of blade are in contact with the plane of section, the edge at mid-distance will be 72 » above this plane; whence it follows that with each full heel-to-point horizontal stroke in cutting, as when revolved horizontally at an acute angle of about 4° with reference to the direction of motion, the edge will twice traverse a vertical distance of 72 through the tissue, destroying the section. With the edge thus imperfectly aligned, the knife can be used to advantage in a sliding microtome only when placed less obliquely to the direction of motion than is essential for the cutting of thin sections, as at 20°-40°, in which position sections are cut with fractional strokes considerably less than the length of a 170 mm. blade. To give the true edge essential to accurate work in cutting woody tissues, the use of a special type of hone is necessary. Such a hone consists of a piece of plate glass as wide as the knife is long, with a true plane surface covered with Diamantine Powder and oil or water, so that the knife remains constantly in contact with the hone throughout its length during the process of grinding. An excellent abrasive may be prepared by grinding two fine Belgian or carborundum hones together, and collecting: the fine powder thus produced on the surface of the glass. This is not so harsh as the No. 1 Diamantine. Having ground the blade on the glass hone until the edge at both sides makes perfect contact with the surface, a brief grinding on a Belgian hone (2 x 8 in.) wet with 30 per cent glycerine will often give an edge sufficiently even and sharp for general work. The knife should be drawn across the hone obliquely, heel and edge forward, alternately upon each side. But, for best results, this grinding should be followed by sharpening upon a flat leather-surface hone, the blade in this case. being pushed obliquely, point and back forward, along the hone. The surface of this hone may be kept in condition by the use of any of the fine abrasives used for this purpose on razor strops. A strop 132 BROWN: Woops FOR MICROSCOPIC STUDY of the hanging type should not be used in that it tends to round the planes of the knife wedge. The wedge angle is best regulated by the use of a cylindrical metal clamp fastened to the back of the knife during the process of sharpening. In addition to the possession of (1) true axial alignment , the edge should (2) be sharp enough throughout its length to cut a hair 7 t Tt tdeediee el ee en Fic. 2. Block showing tropical wood (Te- coma sp.) with tissues properly oriented for sectioning in cross, radial, or tangential planes. heart-wood; S, sap-wood. The rays (M, R) are low and narrow; the vessels (V), num- erous and small. G, growth-ring. at least one full growth-ring (G, Fic. by contact and (3) should appear perfect'y even and without nicks under the low power of the microscope. The number of sections which may be cut without re-sharpening the knife is ordinarily small; with diffi- cult material, frequently as few as two or three perfect sections. 3. PREPARATION OF MATERIAL FOR SECTIONING A. CUTTING THE BLOCKS In preparing material for sectioning, the blocks must be shaped with refer- ence to the plane or planes in which the sections are to be cut. Thus a block in- tended for a cross, radial, and tangential series of sections may have a radial length of 2 cm., a vertical height of 8 mm., and a tangential thickness of 6 mm. Sections cut from the three planes of such a block will ordinarily include 2) and the full height of the BROWN: WooDs FOR MICROSCOPIC STUDY 133 rays. Occasionally a species with higher rays or other special features may require blocks of larger dimensions. When present, both heart-wood (H) and sap-wood (S$) may be included. The block should be carefully trimmed with a knife so that the tissues are perfectly oriented with respect to each of the planes intended for sectioning. The medullary rays in particular should coincide as perfectly as possible with the radial plane. In tropical woods, the rays are often narrow, visible only under a lens, and curved, so that more than ordinary care is necessary in trimming this surface. Before proceeding with the treatment, the blocks should be numbered for sake of record. Some use a system of notches along the edges. Perhaps one of the best methods is to carve Roman numerals on the radial face not intended for sectioning, underscor- ing IX to distinguish it from XI (N, Fic. 2); also, many tropical woods require an arrow to indicate the direction of growth. B. REMOVING THE AIR After the blocks have been cut the desired shape, air should be extracted as far as possible from the cell lumina by alternate boiling and cooling in water. The use of an Eimer and Amend aspirator No. 3250 after boiling greatly hastens the process. This treatment should be continued until the lightest blocks sink and little or no air comes from the tissues when the aspirator is applied. Most blocks sink in a few hours, but species with numerous tyloses may take a longer time. Thus, blocks of Rhus semialata Murr. var. sandwicensis Gray remain floating five days during the boiling and cooling process, or two days when boiling periods are followed by the application of the aspirator. C. SOFTENING THE TISSUES Nearly all xerophytic species of tropical woods require a long treatment in strong hydrofluoric acid, often extending over several weeks, before they can be sectioned. Rain forest and bottom- land woods are frequently soft, but even such woods usually cut with clearer outlines when treated for at least a few days in acid. For treating material with hydrofluoric acid, the blocks, after removal of air, may be placed in a wide-mouthed glass bottle, care- 134 BROWN: WooDs FOR MICROSCOPIC STUDY fully coated inside with hard paraffin, or, still better, in cups pre- pared by cutting empty hydrofluoric acid bottles. Strong hydro- fluoric acid is added to cover the material, and the container then corked or covered with a hard paraffin plate for a period varying from a few days to six weeks or longer, the length of time being determined by removing blocks at intervals of one to several days and testing with a sharp scalpel until they are found to be suf- ficiently soft to cut easily in transverse section. In the writer’s experience, tissues are rarely injured by leaving them a long time in hydrofluoric acid. Thus, in case of a young stem of Dracaena curea Mann, deli- cate tissues, such as the phloem, cambium, undifferentiated parenchyma, and other thin-walled tissues, together with the bun- dles of needle-like raphides of calcium ox- alate, were all left uninjured by long treat- ment in acid. In one instance, a block of Santalum Freycinetianum Gaud., of the size and shape described for Fic. 2, was placed wiki eo in acid on September 6, 1917, and left until eins: March 26, 1918, when it was washed and sectioned. Though in moderately strong acid for over six months, all structures, including the mineral crystals, were in perfect condition. Hydrofluoric acid probably softens the tissues mainly, if not entirely, by the removal of silica (desilicification). But other minerals would probably be acted upon. For example, calcium would form the insoluble calcium fluoride which would remain in the wall. Curiously, crystals of calcium oxalate in crystal paren- chyma or idioblasts usually remain nearly or quite intact long after the wood has been sufficiently softened to cut well. Even in maceration by Schultze’s solution, crystals may be uninjured. Dr. A. J. Hill suggests that, in case of the hydrofluoric acid treat- ment, the crystals may have been protected by the formation of an insoluble film of calcium fluoride, the possibility of which is seen from the following equation: CaC,0, + 2 HF = soy + H2C20, nsol, BROWN: Woops FOR MICROSCOPIC STUDY 135 The presence of such a film was indicated, though not certainly proven, by refraction tests. Another possible agency of protection may consist in a thin and presumably impermeable organic mem- brane closely applied to the surface of the crystal. Such a mem- brane may be detected by zinc chlor-iodide or by the haematoxylin stain. Also, a thicker mucilaginous membrane is usually visible outside the inner thin membrane. The resistant character of these organic coverings is indicated from the fact they may be little if at all acted upon by Schultze’s solution. As soon as the tissues have been sufficiently softened to cut. well, they should be washed in running water for about four days to remove all acid. An arrangement like that shown in Fic. 3 is recommended both for this purpose and for washing the sections at a later stage. The blocks are placed in a short jar the top of which is covered with cheese-cloth through which the pointed end of a glass siphon tube is pushed. The long arm of the siphon should extend to the bottom of the jar. The short arm takes its water from a somewhat taller jar placed beneath a running tap. After washing, the blocks should be covered with glycerine where they may remain until needed for sectioning. The effect of the glycerine is such that any tissues which have become too brittle, in a few hours, become sufficiently flexible to cut well with the microtome. D. IMBEDDING Celloidin method.—Certain woods of peculiar structure, like Pisonia, in which portions are composed of very soft unlignified tissues, and other woods, in case it is desired to cut extremely thin sections, must be imbedded in celloidin before cutting. In imbedding, the general procedure followed was that described by Plowman (’04). Solutions of Schering’s celloidin in a mixture composed of equal volumes of ether and absolute alcohol may be prepared in 2 x 8 cm. shell vials in concentrations of 2, 4, 6, 8, 10, 12, 14, 16, 18, and 20 per cent. The blocks to be imbedded are first treated in hydrofluoric acid and washed as above described. Next, in the usual manner, they are gradually transferred to 100 per cent alcohol. During the latter stages of this dehydration process, they are left two days in each alcohol, and the absolute alcohol is changed at least once. They are now transferred to a 136 Brown: WooDs FOR MICROSCOPIC STUDY mixture of ether and alcohol, in equal volumes, and finally through the graded series of celloidin solutions, being left about twenty- four hours in each concentration. On reaching the 20 per cent solution, a pressure-resisting container should be used. A brass case with screw top, such as a microscope objective box, will answer the purpose, or the corked imbedding vial may be wrapped tightly with strong cord. After adding sufficient solid celloidin to thicken the solution as much as possible, it is placed in a paraffin oven at 50-60° C. for three days. During this time, the contents of the vial are kept under pressure by the confined gases with the object of forcing the celloidin more perfectly into the tissues. - The imbedded blocks are then removed, hardened in chloroform for twelve to twenty-four hours, and placed in a mixture of equal parts of 95 per cent alcohol and glycerine until ready for sectioning. Paraffin method—Only very soft tissues should be imbedded in paraffin. Woody tissues, even after being well softened by acid, are likely to become too hard during the process of imbedding. in paraffin to cut well with the microtome. E. SECTIONING WITH THE MICROTOME Unimbedded material.—After softening the tissues as already described, the material is ready to section. In cutting sections, the blade of the knife should be kept flooded with a 15 per cent solution of glycerine in 95 per cent alcohol. Sections are best removed from the blade by a fine camel’s hair brush and are transferred to water. From blocks prepared as above described, cross-sections may be cut as thin as I 0-15 u; radial sections, 8-20 p; and tangential sections about 7 4. Where, for special purposes, such as the study of the detailed structure of pit membranes, it is necessary that the sections be considerably thinner than this, the material must be imbedded in celloidin as above described. Imbedded material—To section material after imbedding in celloidin, the blade may be moistened either with 95 per cent alcohol or with the alcohol-glycerine mixture. Sections of woody tissues may then be cut to less than 3 in thickness. To obtain very thin sections, it is essential not only that the knife be sharp, but that it be ground to an alignment sufficiently true to admit of the use of the blade at very oblique horizontal inclinations with refer- BROWN: WoOoDs FOR MICROSCOPIC STUDY 187 ence to the direction of motion, such as 4°. Thin sections such as these require great care in handling. In most cases, particularly in the longitudinal planes, only extremely small pieces may be cut, since such sections, being less than the diameter of the cells, are often sections of single cells and may even be too small to be seen without a lens. After washing the sections in water to remove all glycerine, the celloidin may be removed by covering the sections for several hours with the ether-alcohol mixture. The process of clearing may go on very slowly and more than twenty-four hours may be required to remove all celloidin. The sections should now be placed on a slide in dilute albumen fixative (one drop to 2 c.c. of distilled water) and warmed slightly until dry. After standing over night, they are ready for staining. For staining, Koplin jars are best used, but otherwise the procedure is the same as for unimbedded material to be described in connection with differen- ‘ tial stains. F, FREE HAND SECTIONS Since small unstained radial and tangential sections mounted in glycerine ordinarily show the essential anatomical features, the following method, though especially adapted to coniferous woods, is suggested, in connection with macerated preparations, to assist in the rapid identification of material. The radial and tangential surfaces are first cut true with a knife, then a sharp razor is drawn lightly over the surface allowing it to cut thin fragmentary pieces. These are placed in a drop of water on a slide and held over a flame until boiling temperature is reached. Alcohol is now dropped upon the section until all air has disappeared from the cell-lumina. After a brief staining in aqueous potassium iodide solution of iodine, the sections are ready to mount in glycerine. 4. PREPARATION OF MATERIAL BY MACERATION Many tropical woods contain substances in the ray cells and in other elements, by reason of which details of pitting and similar features are obscured. Macerations will be found satisfactory in such instances and the following modification of Schultze’s method is suggested for their preparation :— 138 Brown: WooDs FOR MICROSCOPIC STUDY 1. Cut chips to expose a radial length of 2 cm., an axial length of I cm., and a tangential thickness of 2 mm. 2. Place these in a numbered test tube, cover with water, and keep at boiling temperature in the water bath for about an hour. The hot water is then replaced by fresh cold water and the tube reheated. Alternate cooling and boiling periods should be continued for at least five changes. 3. Cover the material with 50 per cent nitric acid and add a small amount of potassium chlorate (as much as may be taken upon the point of a small knife blade). It is now kept at boiling temperature until the pieces have whitened and commenced to fray, when cold water is poured on, causing the macerated material to settle. The acid solution is then replaced with water and al- lowed to heat with the purpose of removing the acid solution. 4. Transfer the material to a watch-glass where it may be teased apart with a small size artist’s brush. By means of a pipette, change the water several times or until all particles of foreign matter and traces of chemicals have been removed. 5. Replace the water by 95 per cent alcohol for 30 minutes or until all air has been removed from the cell-lumina. 6. Replace the alcohol by a 2 per cent solution of Bismarck brown in 70 per cent alcohol, in which the material should stand for twelve to twenty-four hours. 7- To remove excess stain, wash quickly in alcohol and transfer to glycerine, which should be changed if greatly colored by the stain. 8. Mount in glycerine and cement with Brunswick black or gold size. Preparations such as these are almost essential for the working out of the details of vascular anatomy. If a water bath capable of receiving twenty test tubes is used, but little time need be consumed in making the preparations. The action of the chemi- cals proceeds so slowly that there is little danger of tissues being destroyed. 5. DEFINITION OF ANATOMICAL DETAILS A. BY DIFFERENTIAL STAINING In the writer’s experience, Haidenhain’s iron-haematoxylin has proved the best all-round stain for bringing out the anatomical BROWN: WooDs FOR MICROSCOPIC STUDY 139 structure of woody tissues. This is true both for the relatively thick sections cut from unimbedded blocks and for the very thin sections obtained by the celloidin method. In these latter sec- tions, for example, it is possible with this stain to bring out such structures as the reticulated thickenings of pit membranes as well as similar features not visible by other means. Before staining, sections cut from the unimbedded blocks or imbedded sections on slides are washed in three or more changes of water to remove glycerine. The sections are then covered with a 2.5 per cent aqueous solution of iron-alum (ferric ammonium sulphate) for five to twelve hours. Next, they are washed in run- ning water for five minutes (or two minutes in case of slide material), stained with 0.5 per cent aqueous solution of haema- toxylin (Chamberlain, ’15, p. 41) for twelve to twenty-four hours, washed in water for three minutes, and then again treated with the iron-alum solution. The sections are left in the iron-alum until they become light gray, but, for best results, the material should be watched under the microscope, so that the process may be stopped the moment the clearest definition is reached. Sections are then washed in 1unning water for three hours or longer to iemove all trace of iron-alum. In many cases, this washing also serves to remove precipitates and other clouding matter from the tissues so that the clearness and transparency of the section is greatly improved. It is often well to extend the period of washing to twenty-four hours or longer. For this purpose the siphon apparatus shown in Fic. 3 is useful, since it allows a thor- ough percolation of water through the sections for any length of time without danger of mechanical injury to the tissues or of loss of material. The sections rise flat to the top directly in the out- flowing tap water which, from a public supply system, is usually sufficiently alkaline to give a clear blue color to the haematoxylin, hence being preferable to distilled water for this purpose. For best results, woody tissues stained as above should be counterstained with safranin. This stain is made up by combining equal portions of a 1 per cent solution of alcohol-soluble safranin in 95 per cent alcohol and a 1 per cent aqueous solution of water- soluble safranin. For extremely thin membranes, where a very weak counterstain is desirable, one to three minutes in the safranin 140 BROWN: WoopDs FOR MICROSCOPIC STUDY may be sufficient; but for general work, longer periods up to two hours, giving a heavy but not too opaque stain, are desirable. De- hydration is accomplished by washing quickly in 95 per cent alco- hol, then successively in absolute alcohol and xylol. The sections are mounted in dammar or balsam. Sections which tend to curl badly after cutting, as in case of many woods with thick-walled elements, may be placed, as soon as cut, between two glass slides and allowed to dry before staining. In most cases, the section may then be placed free in the stain, but occasionally it may be necessary to proceed with one or more steps of staining before removing from the slides. For a number of purposes, the above process, with slight modification, may be used successfully in staining the soft cellulose tissues of the bark; but for this purpose Congo red is superior to safranin as a counterstain. Where shrinkage due to dehydration is not too great and the cell contents stain black, as in Malus or Pyrus, preparations showing clearly such details as the reticulate thickenings, pits, and even protoplasmic bridges through the pit-membranes of the hypodermal ‘collenchyma may be obtained by the following process :— I. Sections 6 w in thickness are cut from green material col- lected in winter, to show pits and protoplasmic bridges in section, or 10 uw in thickness to show pits and reticulations in surface view. 2. Place for five hours in 2.5 per cent aqueous solution of iron-alum. _ 3. Wash five minutes. 4. Stain about twelve hours in 0.5 per cent aqueous solution of haematoxylin. 5. Wash five minutes. 6. Differentiate in a 2.5 per cent iron alum solution until sections are gray. 7. Wash three hours. 8. Stain in a saturated aqueous solution of Congo red one to five minutes for pits in section, ten to thirty minutes for pits in surface view, one to two hours for reticulation of thin end-walls. 9. Dehydrate in absolute alcohol one minute. 10. Clear in xylol one minute. 11. Mount in dammar. BROWN: WoOoDS FOR MICROSCOPIC STUDY 141 Also, longitudinal sections of phloem 6 uw in thickness, when treated as above, will show the lattice-like arrangement of sieve areas in the lateral walls of the sieve tubes. A convenient method of numbering and labeling slides is as follows. By means of a camel’s hair brush, cover the entire upper surface of the slide outside the coverslip with dilute dammar (mounting consistency diluted twenty to thirty times with xylol). The brush is best fixed in the cork of a bottle used to contain the solution. In a few seconds the records may be written with Hig- gin’s waterproof ink, and index arrows sketched to point to any special part of the mount, if desired. The surface should then again be brushed with dammar that the slide may be freely handled or washed without injury to the writing. B. BY DIFFERENTIAL REFRACTION The laws of refraction may be employed not only to give clear definition to details in outline, but also to determine physical or chemical qualities of cell-walls or cell-contents. In either case characters may often be determined by refraction more readily and with greater precision than by the reaction of stains or of chemical reagents. Refraction opens'a wide field for research in plant histology and will be treated in some detail in a subsequent paper dealing with refraction of light in relation to plant tissues. 6. MICROCHEMICAL REACTIONS Characters readily observed by microchemical means often have a systematic value, or are of ecological or physiological in- terest, or are closely identified with the peculiar qualities of color, hardness, hygroscopicity, porosity, durability, strength and other properties upon which depends the special value of any given wood in the arts. In the following paragraphs are indicated several important characters frequently found in woody tissues of warm climate trees, together with a number of microchemical tests helpful in connection with their demonstration. A. CELLULOSE Sections are placed twelve hours or longer in a solution made by adding one drop of concentrated aqueous Congo red to 20 c.c. 142 Brown: WooDs FOR MICROSCOPIC STUDY of distilled water. Cellulose tissues turn deep blue as soon as transferred to 10 per cent hydrochloric acid. If mounted in glycerine (acidulated by adding one drop of strong hydrochloric acid to 10 c.c. of glycerine) and well cemented with Brunswick black or gold size, the blue color will remain several months without fading. B. CELLULOSE-LIGNIN The above test for cellulose may be varied by transfercing the section for a brief period to a strong aqueous solution of anilin chloride immediately after the treatment with Congo red, the remainder of the treatment being the same. Lignified tissues appear yellow; cellulose, blue. Cc. GUMS Gummy substances, both those which are soluble in water and those which swell without going into solution (mucilages), are constantly met with in woody tissues, occurring in the lumina of vessels, tracheids, and other cells, or in intercellular canals or cavities, these latter resembling resin cavities. Certain forms of each (gums and mucilages) are believed to play the r6le of reserve material (Haas and Hill, ’13, p. 125; Griiss, ’96). Fibers, composed in large part of mucilaginous inner layers, are also of frequent occurrence. The gummy nature of amorphous bodies observed in permanent mounts is usually indicated by shrinkage checks or other evidences of contraction on dehydration, as in Fic. 5 (see further below). Gums may be further distin- guished from resins by their insolubility in chloroform. Mucilaginous laminae in fibers (‘‘gallertartige Verdickung”’ Sanio, 63, p. 105; Solereder, ’08, p. 1143: ‘ Hemicellulose,” Griiss, ’96; Schellenberg, ’05: ‘‘ cellulose,” Potter, ’o4: ‘‘mu- cilaginous fibers,” Sachs, '75, pp. 35-36; Jeffrey, ’17).—Fre- quently woody fibers, particularly of xerophytic species, are composed of one or more clear, gelatin-like concentric inner layers which shrink greatly on drying (Fic. 4); in young tissue, they turn deep violet with zinc chlor-iodide. Solereder ('08, p. 1143) records twenty-two families in which such fibers occur, and his list is not complete. In a given wood, fibers of this description may occur either sporadically or in regular distribution ; Brown: Woops FOR MICROSCOPIC STUDY 143 they may be few in number or may compose nearly or quite 100 per cent of the prosenchyma. In some cases (Robinia), the purple reacting lamellae of fibers may function as reserve material, and be more or less completely dissolved in the growing season (Schel- lenberg, ’05; Griiss, ’96). In this connection, twigs of Acer rubrum L., Cercis canadensis L., and Robinia Pseudo-Acacia L., gathered in winter condition, sectioned, and treated with zinc chlor-iodide, make favorable material for study. A number of distinctive propention are exhibited by such fibers: a. Hyg ——Mucilagi layers readily take up water, swell without see sabes selabiou, and shrink correspondingly on ; B 0 5 10 R 15 20 25 u T Fic. 4. Abbé camera drawings of a laminated fiber of Xylosma hawatiense Seem. in cross-section. © mut after desiccation. B, after treatment with water. 1 yers have swollen, but the mucilaginous layers (G) more than the ligno-cellulose outer portion (ZL). The pode pits (P) come into alignment after soaking. One of the pits remains permanently in connection. T, tangential direction. R, radi direction. drying. If cross-sections 15-30 » in thickness are mounted on a slide in water, then dehydrated by flowing absolute alcohol be- neath the coverslip, the mucilaginous layers may be observed to shrink greatly, in the majority of cases, drawing away from the 144 ~ Brown: Woops FOR MICROSCOPIC STUDY thin outer ligno-cellulose layer usually present (Fic. 4, A). Fibers in which the mucilaginous layers remain in contact with the outer layers on drying ordinarily show large shrinkage checks extending radially outward from the center, through the muci- laginous portion. Dehydration may be completed by placing the slide upon the water bath for a few seconds. In stained sections mounted in balsam or dammar, mucilaginous membranes are usually shown in the contracted state. In glycerine, water, or other media in which the tissue has been mounted without dehydration, the mucilaginous layers appear in the swollen con- dition. Abbé camera outline sketches of single fibers before and after dehydration, made with high power and extended draw tube to obtain the greatest possible enlargement, accurately show the amount of shrinkage as in Fic. 4, A and B. The mucilaginous core of such fibers often contracts over 25 per cent of both radial and tangential dimensions on drying, and swells rapidly to original size on admission of water. The outer ligno-cellulose layer and the middle lamella (M) which is here indistinguishable from the primary thickening of the fiber, on the other hand, show relatively little change in dimensions. The shrinking and swelling of the mucilaginous thickenings is partly independent of the other parts of the tissue, so that, as in the case of the Hawaiian woods ex- amined by the writer, the wood tissue itself was not observed to swell or shrink in proportion to that of the mucilaginous layers of the fibers. However, blocks of wood in which mucilaginous fibers were abundant, as, for example, Xylosma, were found to shrink as much as I2 per cent tangentially and 7 per cent radially on drying from saturation. 6. Reaction to stain and other reagents.—Stains and chemical reagents react differently according to the age of the tissue; also, after material has been softened in hydrofluoric acid, neither stains nor chemical reagents give characteristic reactions. In old tissue (heartwood), haematoxylin, Bismarck brown, and anilin blue stain the mucilaginous layers with varying in- tensity as a substance of variable composition. Very commonly stains, particularly the safranin, are readily extracted by the al- cohol washes or other treatments in the staining process, so that the mucilaginous layers appear slightly or not at all stained in the BROWN: Woops FOR MICROSCOPIC STUDY 145 finished mount. With phloroglucinol and hydrochloric acid, a more or less pronounced red reaction is usually obtained; with zinc chlor-ioidide, a yellow, brownish, or sometimes purplish color. In young tissue (twigs, sapwood), the mucilaginous membranes stain deeply with haematoxylin, or Congo red; with zinc chlor- iodide they turn deep purple or violet. c. Fracture-—In maceration, mucilaginous fibers are usually extremely brittle. A fragment ordinarily shows conchoidal fracture across the mucilaginous core. Woods in which mucilagi- nous fibers are abundant are likely to be brittle. 50 100 py Fic. 5. Abbé camera drawings of vessel in Tabebuia chrysantha Nichols, (?), show- ing surface of insoluble gum (mucilaginous) plate (M). A, after dehydration; B, after admission of water beneath coverslip. The shrinkage crack (C) closes in B. Mucilage or insoluble gum in cells, vessels, or canals.—The mucilaginous or gummy substances, which occasionally fill cer- tain cells, vessels or intercellular cavities in dicotyledonous woods, respond to water tests in much the same way as the mucilaginous portion of mucilaginous fibers. Here the substance does not ordinarily pull away from the sides of the cavity or canal on dehydration, but cleaves apart in long wide gaps extending, usually, across the center. Such shrinkage cracks may be observed to close up tightly soon after admitting water at the edge of the coverslip, and drawing it through by means of blotting paper at the opposite side (Fic. 5). The water test serves also to distin- guish the gums or mucilages from the resins, which, though they may shrink on dehydration, do not swell in water. 146 BROWN: Woops FOR MICROSCOPIC STUDY D. ESSENTIAL OILS, RESINS, AND GUM-RESINS Substances of this description occur abundantly in woody tissues of tropical dicotyledons, in the'cell lumina or in intercellular canals (Guignard, ’92; Record, ’18), in the form of globules, transverse plates or irregular masses. As before indicated, such substances may readily be distinguished from the gums by the fact that after drying they do not swell in water; also, by their solubility in chloroform. The gum-resins, however, may have some of the properties of each component. To determine the solubility in chloroform, sections (particularly those from seasoned blocks) should remain twenty-four hours in the solvent. The essential oils, which often possess an aromatic odor, resemble the resins except that such bodies readily dissolve when absolute alcohol is drawn beneath the cover glass. E. FATS : The fatty oils resemble, in appearance, the essential oils with which they often occur in the same cell. Both are stained by Sudan III. The fatty oi’s are distinguished from essential oils ‘from the fact that absolute alcohol dissolves few fatty oils, but does dissolve the essential oils; strong potassic hydrate saponifies fatty oils, but not essential oils, although some complex resins which break down into a fatty component may appear to be saponified; a temperature of 130° C. volatilizes essential oils but not the fats. F. TANNIN The presence of tannin in tissues is readily demonstrated by familiar tests. A bluish or greenish black is caused by a neutral- ized solution of ferric chloride, and a reddish yellow by ammonium molybdate. G. MINERAL CRYSTALS Vertical strands of short cells, each with a single crystal of calcium oxalate nearly filling the cavity, are of frequent occurrence in tropical woods, being found in over 26 per cent of the Hawaiian woods studied by writer. Less frequently they occur in ray-cells and other tissues. Their composition is determined from the fact that calcium oxalate dissolves without effervescence in BROWN: WooDSs FOR MICROSCOPIC STUDY 147 hydrochloric acid, but is unaffected by acetic acid. Calcium carbonate, which is of comparatively rare occurrence in crystalline form, dissolves in either acid with effervescence. The fact that each mineral crystal is often closely incased in a resistant, nearly impermeable membrane may make it necessary to free the face of the mineral from its coating before the test can be applied. This may be done by holding a thin section in a pair of forceps, igniting, and allowing to burn or char only suffi- ciently to free the crystal,or a portion of it. Care must be exercised, for the calcium oxalate, with sufficient heat, is changed to calcium ca bonate and finally to calc um oxide. : H. CHEMICAL GROWTH-RINGS Some woods without any visible structural growth-rings, may yet exhibit what appear to be definite seasonal variations in the chemical composition of the wood, so that distinct chemical rings es pad Se & ees A, O) (estes AGI ED &) , Rae a 2 HOt ot root Ort POS Oat RHE S] Ho Cea tird bovay Vee WER Pelt OTOL PES St sy ett tek AA I: we a ab FAGt FOLOL O cs] a ee 14a fa A HON aGug § " Ti do) PS a= Seg" \ 9) ae. ra S o> yr ah e05 of «| ZFS) iO Y] jee x re hh tS Se Y¥ (OIF a) Ne rel ch, ‘SJ ie (| qin \ Vie Ps SE 4 wa co 0 med 100 Chemical growth-ring in Xylosma hawaiense Seem. Arrow indicates dicection of growth. Z, lignified zone. A slight retardation of ewe is indicated by somewhat shortened ray-cells at R, and occasional slightly compressed fibers (F)- may be demonstrated by microchemical means. The term chemi- cal growth-rings is here suggested for such zonal variations in the 148 BROWN: WooDs FOR MICROSCOPIC STUDY cellulose, lignin, mucilage, or other components of the cell-wall, which appear correlated with seasonal growth. The evidence that such zones of chemical variation actually represent true growth-rings is not without structural confirmation. For example, on following carefully a chemical ring (Fic. 6) which appears sharp on its outer face, as in Xylosma hawatiense Seem., such evidences as the presence of short ray cells (R) or slightly flattened fibers (F) would seem to indicate a slight retardation of growth at that place. In case of woods in which chemical rings are present due to seasonal variations in lignification, such rings are readily demon- strated by placing cross-sections 15-20 thick in an alcoholic solution of phloroglucinol for two minutes, then treating with strong hydrochloric acid. Definite chemical rings appear, which are sharply defined on the outer face, blending gradually into deeper shades toward the interior. Similarly, the cellulose test may reveal zonal differences in the amount of cellulose. Another type of growth-ring is defined by seasonal zones of mucilage- reacting fibers, but such zones, though occurring in regular zonal alternation, may not be sharply defined on either face. In Alphitonia excelsa Reiss., such mucilaginous chemical rings are plainly visible without a lens. 7. Liguip PENETRATION TEST It not jadvegnente happens that tyloses, gums, and other bodies which may fill the lumina of vessels and other conducting elements in the region of the heartwood, are displaced or lost during the process of sectioning or staining, so that, from a micro- scopic examination, one may fail to make an accurate estimate of their presence or abundance. The following test is of material aid: Alcohol is dropped upon the transverse surface of a dry block of wood. If tyloses, gums, or other bodies are abundant, the liquid spreads out over the surface; if absent, the liquid quickly disappears in the tissues and soon runs through to the opposite end. Thus, in case of the red oaks, only a few seconds are re- quired for the liquid to appear at the opposite end of blocks six inches or more in axial thickness. Woods, in general, separate rather definitely into two classes, those which are penetrable and those which are not. BROWN: Woops FOR MICROSCOPIC STUDY 149 8. SUMMARY In cutting thin woody sections with a sliding microtome, it is essential, for accurate results, that careful attention be paid to sharpening and adjusting the knife. The blade must. be ground to perfect axial alignment. The wedge-shaped edge should have an angle of approximately 20°; for general work this gives better results than a more acute edge. In softening tissues of tropical woods for sectioning, strong hydrofluoric acid may be used in preference to weak. The length of time required to soften tissues of tropical woods varies from a few days to several weeks, little if any injury being done to the tissues by remaining a. long time in acid. The process of demineralization of the cell-wall may be completed without per- ceptible effect upon the outline of the calcium oxalate crystals contained in the cells, beyond the fact that the refractive properties may be changed. In the maceration of woody tissues, Schultze’s method may be employed with safety if equal volumes of acid and water are used. Fibers with one or more mucilaginous inner layers are frequent in the xylem of tropical trees. Perhaps the most constant proper- ties of the mucilaginous layers consist (1) in their marked swelling in water and (2) their brittleness on drying. With stains and reagents, reactions differ with the age of the tissue. Thus in young tissue (twigs, sapwood) gathered during the period of rest, a purple color is obtained with zinc chlor-iodide; haematoxylin, anilin blue, or Congo red stain deeply. In old tissue (heartwood), the mucilaginous layers react, as a rule, yellow or brownish with zine chlor-iodide, and take stains irregularly. Woods in which mucilaginous fibers are abundant may shrink greatly on drying or swell correspondingly on wetting, though not to the same degree as the mucilaginous membranes. Woods without structural growth-rings may possess chemical rings demonstrable by microchemical means. E UNIVERSITY LITERATURE CITED Chamberlain, C. J. ('15). Methods in plant histology. Ed. 3. Chicago. Guignard, L. ('92). L’appareil sécréteur des Copaifera. Bull. Soc. Bot. France 39: 233-260. f. I-13. 150 BROWN: WooDs FOR MICROSCOPIC STUDY Griiss, J. (’96). Uber Lésung und Bildung der aus Hemicellulose bestehenden Zellwande und ihre Beziehung zur Gummosis. Bibl. Bot. 39: 1-13. pl. 1. Haas, P., & Hill, T. G. ('13). An introduction to the chemistry of plant products. New York. Jeffrey, E. C. (’17)._ The anatomy of woody plants. Chicago. Plowman, A. B. (’04). The celloidin method with hard tissues. Bot. Potter, M. C. (04) On the occurrence of cellulose in the xylem of woody stems. Ann. Bot. 18: 121-140. pl. 8. Record, S. J. (’18). Intercellular canals in dicotyledonous woods. Jour. For. 16: 429-441. f. 1-8. Sachs, J. (’75). Text-book of botany. English translation. Oxford. Sanio, C. (’63). Vergleichende Untersuchungen iiber die’ Elementar- organe des Holzkérpers. Bot. Zeit. 21: 85-91; 93-98; 89-III; 113-118; 121-128. pf. 4. Schellenberg, H. C. (05). Uber Hemicellulosen als Reservestoff bei unseren Waldbaumen. Ber. Deuts. Bot. Ges. 23: 36-45. Solereder, H. ('08). Systematic anatomy of the Dicotyledons. Eng- lish translation. Oxford. Thomson, R. B. (’10). A modification of the Jung-Thoma sliding microtome for cutting wood. Bot. Gaz. 50: 148-149. f. I. INDEX TO AMERICAN BOTANICAL LITERATURE 1917-1919 e aim of this Index is to include all current botanical literature written by Re published in a i or based upon American material ; the word Amer- ica bei ait in the broadest sense. Reviews, and pap hat relate exclusively to forestry, agriculture, horticulture, elniilace products vegetable origin, or laboratory methods are not included, and attempt is made to index the literature of . An occasional exception is fae in favor of some paper earing in an American periodical which is devoted 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 farnished i 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, Anderson, J. P. Annotated list of plants collected in Greeley and Wallace Counties, Kansas. Iowa Nat. 3: 26-44. Ap 1917. Arthur, J.C. Uredinales of Guatemala based on collections by E. W. D Holway—IV. Puccinia on Carduaceae, form-genera and index. Am. Jour. Bot. 5: 522-550. 9 Includes new species in Puccinia (9), Uredo a) and Aecidium (tr). Ballard, W. R. Strawberry notes. Maryland Agr. Exp. Sta. Bull. 211: 51-76. f. 1-3. Ja 1918. Barker, E. E. Heredity studies in the morning-glory (Jpomoea pur- purea [L.] Roth). Cornell Agr. Exp. Sta. Bull. 392: 1-38. pl. 1-3. Ji 1917. Barrus, M. F.° Varietal susceptibility of beans to strains of Colleto- trichum lindemuthianum (Sacc. & Magn.) B. & C. Phytopathology 8: 589-614. pl. 1-5. D 1918. Berry, E. W. A Cretaceous Hymenaea ee Alabama. Am. Jour. Sci. 47: 65-68. Ja 1919. [Illust.] erry, E. W. The history of the linden and ash. Plant World 21: 163-174. Jl 1918. Bicknell, E. P. Ilex verticillata. Addisonia 3: 71, 72. pl. rio. 30D 1918. 151 152 INDEX TO AMERICAN BOTANICAL LITERATURE Bisset, P. Prolification in a double-flowered form of Calendula officinalis. Jour. Heredity O: 324-425. J. 12,.13. .12 D 1918. Brooks, .M. M. Comparative studies on respiration—III. The effect of ether on the respiration and growth of Bacillus subtilis. Jour. Gen. Physiol. 1: 193-201. f. 17-5. N 1918. Brown, B.S. Fruiting of apple trees every other year. Jour. Heredity 9: 304-306. f. 3. 12 D 1918. Brown, E. D. W. Apogamy in Camptosorus rhizophyllus. Bull. Torrey Club 46: 27-30. pl. 2. 20 Ja 1919. Brown, F. W. Orange like fruit from a lemon tree. Jour. Heredity 9: 308-310. f. 4-6. 12 D 1918. Chapman, G. H. Mosaic disease of tobacco. Massachusett Agr. Exp. Sta. Bull. 175:°73-117: pl. 1-5... My 1917. . Cruess, W. V. The fermentation organisms of California grapes. Univ. Calif. Publ. Agr. Sci. 4: 1-66. pl. 1, 2 +f. 1-15. 31 D'1918. Curtis, O. F. Stimulation of root growth in cuttings by treatment with chemical compounds. Cornell Agr. Exp. Sta. Mem. 14: 71-138. f. -8. Au 1918. Dixon, R., & Fitch, F. E. The human side of trees. i-xxi + 1-199. New York. 1918. Dosdall, L. Overwintering of the aeciospores of Cronartium ribicola Fisher. Phytopathology 8: 619. D 1918. Elliott, J. A. Storage rots of sweet potatoes., Arkansas Agr. Exp. Sta. Bull. 144: 1-15. pl. 1-4. My 1918. Elliott, J. A. Wood-rots of peach trees caused by Coriolus prolificans and C. versicolor. Phytopathology 8: 615-617. f. 1, 2. D 1918. Ewing, E. C. A study of certain environmental factors and varietal differences influencing the fruiting of cotton. Mississippi Agr. Exp. Sta. Bull. 8: 1-93. D 1917. Fitzpatrick, T. J. Corylus rostrata in lowa. Iowa Nat. 3: 44, 45. Ap 1917. Fitzpatrick, T. J. The fern flora of northeastern Iowa. Am. Fern Jour. 8: 97-103. 20 Ja 1919. Floyd, F. G. A crested fern used in landscape planting. Am. Fern Jour. 8: 110-114. 20 Ja 1919. Folsom, D. The influence of certain environmental conditions, espe- cially water supply, upon form and structurein Ranunculus. Physiol. Researches 2: 209-276. f. I-24. D 1918. INDEX TO AMERICAN BOTANICAL LITERATURE 153 Fox, A. Nut fall and leaf droop of coconut palms in Cuba. Cuba Rev. 17: 12-15. Dig18._ [Illust.] Free, M. Effect of low temperatures on greenhouse plants. Brooklyn Bot. Gard. Rec. 8: 14-18. Ja 1919. Freeman, E. M_ The story of the black stem rust of grain and the barberry. Minnesota Agr. Ext. Div. Bull. 27: 1-8. f. 1-5. Ap 1918. Gardne-, N. L. New Pacific coast marine algae—III. Univ. Calif. Publ. Bot. 6: 455-486. pl. 38-41. 3 D 1918. Includes new species in Placoma (1), Dermocarpa (2), Xenococcus (3), Pleurocapsa (2), paoculnel (1), Phormidium (1), Symploca (2), Microcoleus (2), Calothrix (2), Dichothrix (2), Rivularia (1), and Brachytrichia (x Gleason, H. A. Echinacea purpurea. Adiuobhs 3t.67,- 66. D1, Trg. 30 D 1918. Goodspeed, T. H., & Hodgson, R. W. Note on the effects of illumi- nating gas and its constituents in causing abscission of flowers in Nicotiana and Citrus. Univ. Calif. Publ. Bot. 5: 439-450. 28 D 1918. Gustafson, F. G. Comparative studies on respiration—II. The effect of anesthetics and other substances on the respiration of Aspergillus niger. Jour. Gen. Physiol. 1: 181-191. f. 1-5. N 1918. Harper, R. M. A sketch of the forest geography of New Jersey. Bull. Geog. Soc. Philadelphia 16: 107-124. O 1918. Harvey, R.B., & True, R. H. Root absorption from solutions at mini- mum concentrations. Am. Jour. Bot. 5: 516-521. f.z, 2. D 1918. Hayes, H.K. Natural crossing in wheat. Jour. Heredity 9: 326-330. Piae tse 12D sore. Heinicke, A. J. Factors influencing the abscission of flowers and partially developed fruits of the apple (Pyrus malus L.). Cornell Agr. Exp. Sta. Bull. 393: 45-113. f. 1-8. Jl 1917. Hodgson, R. W. An interesting bud-sport in the Washington navel orange. Jour. Heredity 9: 301-303. f. 2. 12 D 1918. Kezer, A., & Sackett, W. G. Beans in Colorado and their diseases. Colorado Agr. Exp. Sta. Bull. 234: 1-32. f. 7-6. Mr1918._ [Illust.] Killip, E. P. A year’s collecting in northeastern United States. Am. Fern Jour. 8: 121-126. 20 Ja 1919. Livingston, B. E. Porous clay cones for the auto-irrigation of potted plants. Plant World 21: 202-208. Au 1918. [Illust.] Loeb, J. Amphoteric colloids—II. Volumetric analysis of ion- protein compounds: the significance of the isoelectric point for the purification of amphoteric colloids. Jour. Gen, Physiol. 1: 237-254. jf. 1-8 WN 1918. 154 INDEX TO AMERICAN BOTANICAL LITERATURE Loeb, J. The physiological basis of morphological polarity in regener- ation—I. Jour. Gen. Physiol. 1: 337-362. f. 1-18. 20 Ja 1919. Longyear, B. O. The dandelion in Colorado. Colorado Agr. Exp. Sta. Bull. 236: 1-35. f. 1-16. Ja 1918. MacCaughey, V. The genus Morinda in the Hawaiian flora. Plant World 21: 209-214. Au 1918. MacCaughey, V. The strand flora of the Hawaiian Archipelago—ll. Ecological relations. Bull. Torrey Club 45: 483-502. 23 D 1918. Massey, L. M. The diseases of roses. Trans. Mass. Hort. Soc. 1918: 81-101. pl. 1, 2. 1918. Massey, L. M. Rose diseases. Am. Rose Annual 1917: 92-101. Fi Dy Fir 1ORF: Matz, J. Report of laboratory assistant in plant pathology. Florida ’ Agr. Exp. Sta. Rep. 1917: 87R-94R. f. 9-15. My 1918. Includes notes on pecan dieback and an undescribed Gnomonia on pecan leaves. McClintock, J. A. Sweet potato diseases. Virginia Truck Exp. Sta. Bull. 22: 455-486. f. ro8—r2r. 1 Ja 1917. Melhus, I. E., & Vogel, I. H. Cabbage diseases. Iowa Yearbook Agr. 18: 435-438. f. 1-3. 1918. Merrill, E. D. Oreomyrrhis borneensis Merr. sp. nov., an interesting addition to our knowledge of the Malayan flora. Am. Jour. Bot. §: 514, 515. pl. 30... D 1918. Murrill, W. A. Three young Crusoes. i-xii + 1-218. pl. 1, 2+ jf. 1-83. New York. 1918 Contains some botanical information of the West Indies. Nafziger, T. E. How sorghum crosses are made. Jour. Heredity 9: 4at, 320: 12 DD 7018; Nash, G. V. Salvia farinacea. Addisonia 3: 77, 78. pl. r19. 30 D 1918. Nash, G. V. Spiraea Thunbergii. Addisonia 3: 63, 64. pl. 112. 30 D 1918. Newman, C. C., & Leonian, L. A. Irish potato breeding. South Carolina Agr. Exp. Sta. Bull. 195: 1-28. f. r-19. Je 1918. Oberholser, H. C. The status of the genus Orchilus Cabanis. Proc. Biol. Soc. Washington 31: 203, 204. 30 D 1918. Osborn, H. F. Algonkian bacteria and popular science. Science II. 46: 432-434. 2 .N 1917. Osterhout, W. J. V. Comparative studies on respiration—I. Intro- duction. Jour. Gen. Physiol. 1: 171-179. f. 1. N 1918 INDEX TO AMERICAN BOTANICAL LITERATURE 155 Osterhout, W. J. V. A comparative study of permeability in plants. Jour. Gen. Physiol. 1: 299-304. f. 1, 2. 20 Ja 1919. Osterhout, W. J. V. Endurance of extreme conditions and its relation to the theory of adaptation. Am. Jour. Bot. 5: 507-510. f. zr. D 1918. Osterhout, W. J. V. A simple method of demonstrating the production of aldehyde by chlorophyll and by aniline dyes in the presence of sunlight. Am. Jour. Bot. 5: 511-513. D 1918. Osterhout, W. J. V., & Haas, A.R.C. The temperature coefficient of photosynthesis. Jour. Gen. Physiol. 1: 295-298. 20 Ja 1919. Oswald, W. L., & Boss, A. Minnesota weeds—III. Minnesota Agr. 4: Exp: Stay Ball. 176: 6-42. f.1-25. JV 1018, Pennell, F. W. Symphoricarpos Symphoricarpos. Addisonia 3: 61, 62. pl. I1z. 30 Di1918 Petrak, F. Die nordamerikanischen Arten der Gattung Cirsium Bei. Bot. Centralb. 32: 223-567. f. 7. 1917. Includes many new combinations. ~ Piper, C. V. Some western species of Lathyrus. Proc. Biol. Soc. Washington 31: 189-196. 30 D 1918. Pulling, H. E. Root habit and plant distribution in the far north. Plant World 21: 223-233. f. 7. S 1918. Ramaley, F. Notes on dune vegetation at San Francisco, California. Plant World 21: 191-201. f. 1-4. Au 1918. Ridgway, C.S. Ernst (4, p. 162) D. trichocephala includes larger forms on the whole than D. velutina, the thallus attaining a maximum width of 3 cm., but it includes also forms which are no wider than those of D. velutina. The size seems to be very strongly influenced by external conditions. he branching of the thallus is usually dichotomous but some- times ventral. The sexual receptacles are terminal, the branches bearing them being variable in length and normally limited in growth. Not infrequently, however, a sexual branch innovates at the apex, the innovation growing in the same direction as the branch and broadening out from a stalk-like base. Such an innovation is usually found in connection with a male receptacle or a female receptacle which has failed to be fertilized; but even a sterile branch, the growth of which has been arrested in some way, is sometimes induced to innovate. Apical innovations in Dumor- tiera have commonly been regarded as somewhat abnormal, caused perhaps by unfavorable environmental conditions. In D. calcicola, where they occur more regularly, at least on sexual individuals, they give rise to jointed sympodial systems of very striking appearance and have been emphasized by Campbell (2, p. 334) as a_ specific peculiarity. Unfortunately similar sympodia have been recorded by Schiffner (10, p. 275) in speci- mens of “D. hirsuta” from New Guinea and by Ernst (4, p. 161) in specimens of D. velutina from Java, so that they are by no means confined to D. calcicola. EvANs: TAXONOMIC STUDY OF DUMORTIERA 171 In its histological features the thallus differs strikingly from most of the other genera of the Marchantiaceae, since it lacks, either partially or completely, the usual dorsal epidermis and system of air-chambers. , The simplification in structure which is thus exhibited is regarded as a derived condition, associated in some way with the usual moist and shaded habitat of the plants. By the earlier writers air chambers were supposed to be invariably absent. Taylor, indeed, in his account of Hygropyla irrigua (26, p. 391), described a system of branched and anastomosing lines (rami) on the upper surface of the thallus but did not intimate that there was any connection between these lines and chambers. Many years later Leitgeb (7, p. 308), working mainly on Taylor’s species, confirmed his observations and showed that the lines were ridges, representing the boundary-walls of vestigial air chambers. In the vicinity of the apex he was able to demonstrate a short-lived epidermis with distorted pores and he noted that epidermal fragments or isolated cells sometimes persisted for a while on the boundary walls of older chambers. Close to the receptacles he observed the frequent occurrence of elongated papilliform cells in the spaces enclosed by the boundaries and pointed out the homology between these and the green cell-chains _ filling the air chambers of Marchantia. In the forms which Leitgeb studied these papilliform cells seem to have been restricted to the vicinity of the receptacles. At any rate he made no mention of them in other parts of the thallus, and the figures which he afterwards published (8, pi. 8, f. 8-11) show a smooth superficial layer of cells. A few years later Spruce (16, p. 566) gave a detailed account of “D. hirsuta,” based primarily on South American material and apparently found a very different state of affairs. According to his descrip- tion the superficial cells are “‘ papilloso-prominulae” and give the living plants a velvety appearance. If there were exceptions to this condition in any of his plants he did not allude to them. Goebel (5, p. 224, f. 63) described the surface of “D. hirsuta” in much the same way and figured the papilliform cells very clearly. Unfortunately he did not state the source of his material. He contrasted the species with an unnamed form from Ceylon, in which he found vestigial chambers but no surface papillae, this 172 Evans: TAXONOMIC STUDY OF DUMORTIERA form thus agreeing essentially with D. irrigua, as described by Leitgeb. Soon afterwards Campbell (1, p. 49), in specimens of “D. trichocephala,’ from the Hawaiian Islands, found neither papilliform cells nor vestigial chambers. The descriptions of Leitgeb, Spruce, Goebel and Campbell thus record the following three types of surface in Dumortiera: (1) with both vestigial chambers and papilliform cells; (2) with vestigial chambers but without papilliform cells; (3) with neither vestigial chambers nor papilliform cells. When these types are clearly defined, as they sometimes are, they seem to yield excellent characters for the separation of species, although writers have expressed dissenting views regarding their value. Schiffner, for example, in his description of D. velutina (12, p. 26), emphasized the crowded papilliform cells as one of the features of the species, while Stephani claimed that such cells were present in all the species and that their persistence in D. velutina merely indicated a shaded environment. Coker (3, p. 226) explained the presence or absence of vestigial chambers in much the same way. In the vicinity of Chapel Hill, North Carolina, he found that plants of D. hirsuta growing in shaded, rather dry localities showed such chambers clearly, while plants in more exposed, wet localities were perfectly smooth; and he attributed these differences to environmental factors. Schiffner (14) criticized the views of Stephani and Coker. He maintained that the degree of develop- ment which the air chambers exhibited was not directly caused by the environment but that it depended upon inherited qualities. To support his statements he showed that “‘D. trichocephala”’ (with greatly reduced chambers) and D. velutina (with better-developed chambers and papilliform cells), often grew under exactly the same conditions in the primeval forests of Java and Sumatra and that both maintained their distinctive peculiarities. He showed further that plants of D. velutina from an almost dark well were exactly the same as those from sunny paths. His most convincing arguments, however, were drawn from plants which had been cultivated under identical conditions for about twenty years in the botanical garden at Prague. These plants represented D. velutina and D. irrigua, and their distinctive differences persisted unchanged. : Evans: TAXONOMIC STUDY OF DUMORTIERA 173 Some of Goebel’s recent observations (6, p. 628, 629) help to confirm Schiffner’s deductions. He described a Brazilian Dumor- itera, presumably a form of D. hirsuta, which lacked vestigial air chambers completely and found that none were formed even when the plants were cultivated under the same conditions as D. velutina, which developed its normal chambers and papilliform cells. He called attention to the fact that very young plants of Marchantia lack air chambers and he therefore interpreted the Dumortiera without chambers as a juvenile condition. In case air chambers are never formed there is simply, in his opinion, a persistence of the juvenile state. If air chambers are formed the presence or absence of papilliform cells indicates a greater or less advance beyond the condition without air chambers. . In other words there are certain races or varieties or ‘‘species’’ of Dumor- tiera which are never able, even under the most advantageous of conditions, to advance beyond the state without air chambers, while other “‘species’’ can advance to various stages beyond this state. Goebel mentioned also a plant of “D. hirsuta,’ which gave rise to an adventive branch without chambers. Since “D. hirsuta,” in his conception of the species, normally develops chambers, this branch was interpreted a as a reversion to a juvenile state. According to the writer’s experience, although the three types of thallus are often distinctly marked, they are by no means invariably so. Even an individual thallus may sometimes bear crowded papilliform cells in an older portion and be smooth or nearly so nearer the apex; another may form vestigial air chambers for a while and then continue its growth without them. Such cases are further examples of reversions and indicate that these may be induced without the intervention of adventive outgrowths. The power to revert, which more or less advanced types thus clearly possess, complicates still further the conditions found in. the genus and adds to the difficulty of defining specific limitations. If a form with crowded papillae represents a ‘‘species,’’ a smooth form growing in the same area may represent merely a juvenile condition of the same thing or it may represent a “‘species’’ which can not advance. At the same time it must be admitted that smooth or nearly smooth states are the only ones which 174 Evans: TAXONOMIC STUDY OF DUMORTIERA occur over extensive areas, according to the information at hand. In Europe and Africa, for example, no forms with crowded papilli- form cells have as yet been reported, although some of the plants have vestigial chambers while others have none. In such areas, therefore, the difficulties of distinguishing between forms with crowded papillae and smooth forms are eliminated, although the difficulties of distinguishing between forms with air chambers and those without them still remain. In the writer’s opinion the latter distinction is less significant than the former, and it seems inex- pedient at the present time to attempt to use it in the delimitation of species. The receptacles of Dumortiera, which represent stalked branch- systems, have been repeatedly described (see, for example, Ernst, 4, Pp. 173-178). The stalk of the male receptacle is extremely short but shows two rhizoid-furrows, agreeing in this respect with the much longer stalk of the female receptacle. The disc of the male receptacle is not clearly lobed and the antheridia are not arranged in radiating rows, although they clearly arise in acropetal succession. The disc of the female receptacle develops normally from eight to sixteen groups of archegonia on its lower surface. hen the receptacle is young there are no distinct marginal lobes, but these become evident later on in case fertilization has taken place, the groups of archegonia being situated beneath the lobes. The involucre is thick and fleshy and shows a small apical opening; on its surface it bears scattered bristles. Similar bristles, which have been interpreted as modified rhizoids, occur on the upper surface of the disc, sometimes abundantly and some- times sparingly, sometimes restricted to the marginal portions and sometimes more evenly distributed. Vestigial air chambers are not developed, and the surface-cells, although sometimes more or less convex, do not form papilliform outgrowths, even when the vegetative thallus forms them abundantly. The features used in separating species have been drawn mainly from the female receptacle and relate more especially to the mar- ginal lobes and the number and distribution of the dorsal bristles. | According to Stephani (25) the receptacle of D. trichocephala is strongly convex and very bristly, while that of D. hirsuta is less convex, with the bristles confined to the marginal portions. He EvANs: TAXONOMIC STUDY OF DUMORTIERA 175 describes further certain ridges or rays on the dorsal surface alternating with the lobes; these ridges branch by forking, the branches extending along the margins of the lobes. In D. tricho- cephala the branches are not connected in any way and thus leave sharp sinuses between the lobes; in D. hirsuta the branches are connected by thallus-substance, and the sinuses thus appear lunulate. These distinctions would be very helpful if they were at all constant but, in the writer’s experience, this is not the case. All gradations occur between strongly bristly receptacles and those with marginal bristles only, while the ridges with their branches, although sometimes fairly distinct, are often vague and evanes- cent. Neither is there any correspondence between differences in the thallus and differences in the female receptacle. A smooth or nearly smooth form, for example, may bear receptacles with many bristles scattered over the surface or with only a few bristles restricted to the margin. The characters assigned to the female receptacle of D. velutina seem at first sight to be more trustworthy. According to Schiffner (12, p. 26) the disc is depressed-conical with broad lobes and very shallow sinuses, the upper surface bearing few or no bristles. The figures published by Ernst (4, pl. 18, f: 1) and by Campbell (2, text-f. 3) represent the upper surface as perfectly smooth, but otherwise agree with Schiffner’s description. In material of D. velutina from Java, determined by Campbell, the receptacles agree closely with his figures and show no bristles. This is not the case, unfortunately, with the specimens distributed by Schiff- ner in his Iter Indicum, one of which came from Java (No. 27) and the other from Sumatra (No. 32). In these the immature receptacles are more or less bristly and some of the bristles are scattered over the upper surface. The older receptacles tend to be less bristly, but their marginal sinuses are often deeper than -Schiffner’s description and the published figures indicate. It would thus appear that the characters assigned to the female receptacle by Schiffner were subject to variation on therefore untrustworthy. Although the antheridia and archegonia are usually borne on separate receptacles in Dumortiera, bisexual receptacles have long been known. They were first described by Taylor (26, p. 391) 176 EvANs: TAXONOMIC STUDY OF DUMORTIERA in D. irrigua, where they seem to be somewhat of ararity. Ernst, however, demonstrated their frequency in Javan specimens of ““D. trichocephala”’ (4, p. 207). He found them also in D. velutina but much more rarely and associated this difference with the prevailingly monoicous inflorescence of ‘‘D. trichocephala”’ and the prevailingly dioicous inflorescence of D. velutina. Whether or not bisexual receptacles are frequent in American forms of Dumortiera remains to be determined. The spores of Dumortiera vary in color from pale to dark brown. The tetrahedral form persists until maturity, although the ridges separating the faces are sometimes difficult to distin- guish. The surface-markings are remarkably uniform. Each face bears a series of minute and crowded papillae or short lamellae, which are usually irregularly distributed but which sometimes show a slight tendency to be arranged in short rows. No reti- culum is developed. With respect to size statements in the liter- ature are not in accordance. Stephani (25) gives a diameter of 25 u for D. trichocephala and of 34 u for D. hirsuta; for D. velutina he gives no measurements. Ernst’s figures are considerably higher (4, p. 178); he gives a length of 45-60 uw and a width of 35-50 un, without distinguishing between D. trichocephala and D. velutina. Campbell (2, p. 329) states that the spores of D. trichocephala are about 20 uw long while those of D. velutina are about 29 u. The writer has examined spores of various forms from widely separated stations and finds that the long diameter measures 20-30 p, a considerable range being often present in a single capsule. Ap- parently little help can be obtained from the spores in the separa- tion of species. The preceding discussion brings out the untrustworthy nature and inconstancy of certain differential characters which have been employed in defining the species of Dumortiera. Those drawn from the size and method of branching of the thallus seem especially unreliable. Those drawn from the female receptacle and the spores are scarcely more satisfactory. On the basis of characters drawn from the structural features of the vegetative thallus, the two following species may be distinguished, and these are the only ones which the writer would recognize at the present time: Evans: TAXONOMIC STUDY OF DUMORTIERA 177 Upper surface of thallus smooth or nearly so throughout ae: often showing vestigial air chambers). . D. hirsuta. Upper surface of thallus with crowded papilliform cells, at least in certain portions (always showing vestigial air chambers). 2. D. nepalensis. I. DUMORTIERA HIRSUTA (Sw.) Nees Marchantia hirsuta Sw. Prodr. Fl. Ind. Occid. 145. 1788. Dumortiera hirsuta Nees, Nova Acta Acad. Leop.-Carol. 12: 410. 1824. Marchantia irrigua Wils.; Hooker, Brit. Fl. 2: 106. 1833. Hygropyla irrigua Tayl.; Mackay, Fl. Hibern. 2:54. 1836. Dumortiera irrigua Nees, Naturg. Europ. Leberm. 4: 159. 1838. _?Dumortiera hirsuta angustior G. L. & N. Syn. Hep. 544. 1846. ?Dumortiera hirsuta intermedia G. L. & N. I. c. ?A skepos brevipes Griffith, Not. Pl. Asiat. 2: 340. 1849. Dumortiera hirsuta irrigua Spruce, Trans. Bot. Soc. Edinb. 15: 566. 1885. It seems unnecessary to give a list of the many specimens which the writer has referred to D. hirsuta. They represent a very extensive geographical range, for the most part tropical, and include a few sterile and poorly developed plants, which are not altogether above suspicion. This is especially the case when such plants were found in regions where D. nepalensis also occurs. The specimens examined came from the following states, countries, and islands: Pennsylvania, North Carolina, South Carolina, Kentucky, Tennessee, Georgia, Florida, Alabama, Missouri, and Arkansas; Mexico, Guatemala, Nicaragua and Panama; Bermuda; Cuba, Hispaniola, Jamaica, Porto Rico, Montserrat, Guadeloupe, Martinique and Grenada; Colombia, Peru and Bolivia; Venezuela, Brazil and Paraguay; Ireland, England (East Sussex) and France (Hautes-Pyrénées); Madeira and the Canary Islands; Fernando Po and Kamerun; China, India (in- cluding Nepal and Ceylon), French Indo-China and Japan; the Philippine Islands, Java and the Hawaiian Islands. It has unfortunately been impossible to secure the actual type specimen of Marchantia hirsuta for examination. The specific name “hirsuta” is here associated with the form without surface papillae, because a Swartzian specimen in the British Museum (kindly examined by Mr. Gepp) is smooth and because the 178 Evans: TAXONOMIC STUDY OF DUMORTIERA smooth form predominates in the West Indies. Out of over fifty West Indian specimens examined (including fifteen from Jamaica, the type-locality for M. hirsuta) only two showed papil- lae. Of course this evidence is not absolutely conclusive, and if it should ever be proved that the original specimen of M. hirsuta represented the papillose form, the synonymy of the species would have to be revised. There is no such doubt about Marchantia irrigua. The Irish specimens of Dumortiera are all smooth on the upper surface, with _ rather vague vestiges of air chambers, and (as Schiffner has shown) never produce papillae even under cultivation. The vars. angus- tior and intermedia of the Synopsis need further study. Var. angustior was based on Mexican plants with a strongly setulose female receptacle, while var. intermedia was based on plants from Peru and South Africa with fewer bristles on the receptacle. Neither of these varieties is known to the writer from authentic material. The same thing is true of Griffith’s Askepos brevipes, which was based on Indian specimens. Schiffner reduced Griffith's genus to synonymy under Dumortiera in 1893 (9, p. 35) but has apparently made no definite reference to its single species. Ac- cording to Griffith’s description and figures A. brevipes is clearly a Dumortiera. It is here referred to D. hirsuta, rather than to D. nepalensis, on the basis of one of Griffith’s specimens in the Mitten Herbarium. This specimen came from Dehra, India, and is labelled ‘‘Askepos’”’; the thallus shows a smooth dorsal surface with vestigial air chambers, and the female receptacle bears scattered bristles. 2. DUMORTIERA NEPALENSIS (Tayl.) Nees prokalios nepalensis Tay}. Trans. Linn. Soc. 17: 392. pl. 15, f. 2. 1836. Marchantia trichocephala Hook. Icon. Pl. pl. 158. 1837. Dumortiera nepalensis Nees, Naturg. Europ. Leberm. 4: 169. 1838. Dumortiera trichocephala Nees, 1. c. 4: 499. 1838. Dumortiera hirsuta latior G. L. & N. Syn. Hep. 544. 1846. Dumortiera hirsuta trichopus Spruce, ios Bot. Soc. Edinb. 15: 567. 1885. Evans: TAXONOMIC STUDY OF DUMORTIERA 179 Dumortiera velutina Schiffn. Denkschr. Math.-Naturw. Cl. Kais. Acad. Wiss. Wien 67: 156. 1899. Dumortiera calcicola Campbell, Ann. Bot. 32: 334. pl. 8 +f. 0. 1918. According to the knowledge at hand the range of D. nepalensis includes neither Europe nor Africa; otherwise it corresponds pretty closely with that of D. hirsuta. In certain parts of its range, such as the southern United States and the West Indies, it has been rarely collected; in other parts of its range, such as southeastern Asia, it seems to be more abundant. The following specimens have been examined :— GEORGIA: Forest Falls, Decatur County, 1901, R. M. Harper 1193a (apparently mixed with D. hirsuta). FLoripa: Gainesville and vicinity, 1909, R. M. Harper 13; 1915, N. L. T. Nelson 100; Pineola, 1918, J. K. Small. Mexico: Honey, Puebla, 1908, Barnes & Land 520, 526; Jalapa, Vera Cruz, 1908, Barnes & Land 564. HonpuRaAs: trail near Rio Platano, 1903, P. Wilson 682. Jamaica: near Newhaven Gap, 1906, A. W, Evans 604. Porto Rico: between Adjuntas and -Ponce, 1906, M. A. Howe 1256. CoLomsiA: Bogota, W. Weir. Ecuapor: near Bafios, R. Spruce (distributed as D. hirsuta var. in Hep. Spruceanae). Peru: Monte Guayrapurina, R. Spruce (distributed as D. hirsuta var. trichopus in Hep. Spruceanae). VENEZUELA: near Caracas, 1854, Burchel; valley of the Rio Limon, Aragua, 1913, H. Pittier 6070. Cutna: Hongkong, collector unknown. Inpra: Nepal, 1820, N. Wallich (type of Hygropyla nepalensis) ; Sikkim-Himalaya, 1889, Fathers Decoby & Schaul 321 (determined by Schiffner as D. velutina). FEDERATED MALAY StATEs: Taiping Hills, 1912, D. H. Camp-. bell. Japan: Uzen, 1888, M. Miyoshi 4; Tokio, 1898, K. Miyake 1; Kioto, G. Shimadzu & Co. 12. PHILIPPINE IsLANDS: Mt. Banjao, Tayabas Province, Luzon, 1907, F. W. Foxworthy 2395; Benguet Subprovince, Luzon, 1901, 180 Evans: TAXONOMIC STUDY OF DUMORTIERA E. D. Merrill 7901; 1910, E. Fénix 12814; 1907, A. D. A. Elmer 8614; Bontoc Subprovince, Luzon, 1910, Father Vanoverbergh 875; near the Shibuyan River, Davao District, Mindanao, 1904, E. B. Copeland 985. SuMATRA: foot of Mt. Singalang, 1894, V. Schiffner (distri- buted as D. velutina in Iter Ind. 32). Java: Buitenzorg, F. A. W. Miquel; same locality, 1894, V. Schiffner (distributed as D. velutina in Iter Ind. 27); same local- ity, 1906, D. H. Campbell; Tjibodas, Preanger Province, 1894, V. Schiffner (distributed as D. hirsuta var. latior in Iter Ind. 24). Borneo: Bidi Caves, Bau, Sarawak, 1913, D. H. Campbell (type of D. calcicola). HawallAN IsLAnDs: without definite localities, D. Dousiss 71 (type of Marchantia trichocephala); 1864-70, H. N. Bolander; 1876, J. Bailey 1; Honolulu, Oahu, 1892, D. H. Campbell; Panoa, Oahu, 1895, A. A. Heller 2330; Manoa Valley, Oahu, 1917, D. H. Campbell; Oahu, 1918, H. L. Lyon; West Maui, 1875, D. D. Baldwin. SAMOAN IsLANDs: Utumapa, Upolu, C. & L. Rechinger (dis- tributed as D. velutina in Krypt Exsic. Mus. Palat. Vindob. 1391); without definite locality, 1888, Frances C. Prince. The type specimen of Hygropyla nepalensis in the Taylor Herbarium shows a thallus with numerous surface papillae asso- ciated, as Taylor’s figures indicate, with a bristly female recep- tacle. The type specimen of Marchantia trichocephala in the same herbarium shows a thallus with still more numerous papillae and female receptacles which are still more bristly. Both speci- mens clearly belong to the same specific type. The var. latior of D. hirsuta, which is listed above in the synonymy, was probably an admixture; it was based on specimens from various parts of the world and may have included forms which would now be referred to the true D. hirsuta. The var. trichopus, however, is _undoubtedly a true synonym of D. nepalensis, as the specimens distributed by Spruce clearly indicate. The last two species which are listed among the synonyms have already been discussed to some extent in the preceding pages. D. velutina is distinguished from strongly papillose forms of D. nepalensis by certain features of the female receptacie, but Evans: TAXONOMIC STUDY OF DUMORTIERA 181 these features (according to the evidence at hand) are too incon- stant to be considered of specific value. In D. calcicola, as already pointed out, the characters drawn from the size of the thallus and from the sympodia developed by the sexual plants are of little significance. The characters drawn from the immature female receptacles with unfertilized archegonia and from the male re- ceptacles are likewise of doubtful value. Unfortunately the papilliform cells of the thallus are rather scantily developed in the type material, but this condition is often pupien ey in un- doubted D. nepalensis. For valuable assistance in the preparation of this paper the writer would express his thanks to Professor W. G. Farlow, Professor D. H. Campbell, Dr. Marshall A. Howe and Miss Caroline C. Haynes. Through their generosity it has been pos- sible to study the types of D. irrigua, D. nepalensis, D. tricho- cephala, D. dilatata and D. calcicola, as well as other. interesting and authentic material. SHEFFIELD SCIENTIFIC SCHOOL, ALE UNIVERSITY. LITERATURE CITED 1. Campbell, D. H. The structure and development of the mosses and ferns (Archegoniatae). London and New York. 1895. Studies on some East Indian Hepaticae. Ann. Bot. 32: 319-338. pl. 8,9 +f. I-10. 1918. 3. Coker, W. C. Selected notes. II.—Liverworts. Bot. Gaz. 36: 225-230. f. I-4. 1903. 4. Ernst, A. Untersuchungen iiber Entwicklung, Bau und Ver- teilung der Infloreszenzen von Dumortiera. Ann. Jard. Bot. Buitenzorg II. 7: 153-223. pl. 18-24. 1908. 5. Goebel, K. Wasserpflanzen. Pflanzenbiologische Schilderungen 2: 217-386. pl. 26-31 +f. 63-121. Marburg. 1893. 2. 6. Organographie der Pflanzen. 2d ed. 2. Bryophytes. Jena. I915. 7. Leitgeb, H. Ueber die Marchantiaceengattung Dumortiera. Flora 63: 307-312. eat beactitmmen iiber die Lebermoose. VI. (Schluss)- Heft. Die Marchantieen und allgemeine Bemerkungen iiber Lebermoose. Graz. 1881. 182 ‘oO Evans: TAXONOMIC STUDY OF DUMORTIERA . Schiffmer, V. Marchantiaceae. Engler & Prantl, Nat. Pflanz- enfam. 1°: 16-38. f. 6-2z. 1893. ———. Ueber exotische Hepaticae. Nova Acta Acad. Caes. Leop.-Carol. 60: 219-316. pl. 6-19. 1893. . Ueber einige Hepaticae aus Japan. O6esterr. Bot. Zeitschr. 49: 385-395. 1899 ie Hepaticae der Flora von Buitenzorg 1. Leiden. 1900. - Neue Materialien zur Kenntniss der Bryophyten der atlantischen Inseln. Hedwigia 41: 269-294. 1902. Ueber Dumortiera. Hedwigia 43: 428, 429. 1904. Studien iiber die Rhizoiden der Marchantiales. Ann. Jard. Bot. Buitenzorg II. Suppl. 3: 473-492. f. 1-3. 1909. . Spruce, R. Hepaticae of the Amazon and of the Andes of Peru and Ecuador. Trans. Bot. Soc. Edinburgh 15. 1885. . Stephani, F. Contribuigoes para o Estudo da Flora a Alvica: Hepaticae. Bol. Soc. Broteria 4: 1-15. pl. 1-3. 1886. Hepaticae africanae. Bot. Jahrb. 8: 79-95. pi. 3, f. 1-9. 1886. Westindische Hepaticae. Hedwigia 27: 276-302. i. II-I4. 1888 Hepaticae costaricenses. Bull. Soc. Roy. Bot. Belgique 31: 175-182. 1892. : Hepaticarum species novae V. Hedwigia 33: 1-10. 1894. Hepaticae africanae. Bot. Jahrb. 20: 299-321. 1895. Hepaticae japonicae. Bull. Herb. Boiss. 5: 76-108. 1897. Hepaticae sandvicenses. Bull. Herb. Boiss. 5: 840-849. 1897. Dumortiera R. Bl. ay [In Species Hepaticarum 1: Bae | Bull. Herb. Boiss. 7: 222-225. 1899. . Taylor, T. De Marchanteis. Trans. Linn. Soc. I7: 375-395. pl. T2-25; 1826. Notes on plants of the southern United States—V FRANCIS W. PENNELL KALMIELLA HIRSUTA (Walt.) Small Sandy scrub-land, between Theodore and Hollanders Island, Mobile County, Alabama, September 3, 1912, Pennell 4513. POLYCODIUM FLORIBUNDUM (Nutt.) Greene Open pine-land, Biloxi, Harrison County, Mississippi, August 28, 1912, Pennell 4405. | SABATIA ELLIoTTIm Steud. Moist scrub-land, between Theodore and Hollanders Island, September 3, 1912, Pennell 4512. Dasystephana tenuifolia (Raf.) Pennell, comb. nov. © Diploma tenuifolia Raf. Fl. Tell. 3: 27. 1837. “‘ Florida . . . seenin the herb. of Torrey.”” The type, labeled in Rafinesque’s hand- writing ‘‘G. tenufolia Raf. Monog.,” is in the Columbia Uni- versity Herbarium at the New York Botanical Garden. It bears data of collection, “‘ Florida, Mr. Croom, 1832, flowers white.” In the American Journal of Science for October, 1833 (25: 69), H. B. Croom records ‘‘Gentiana alba (White flowered Gentian)”’ as growing in ‘‘ wet pine woods’”’ in Middle Florida, a region de- fined as ‘‘ that tract of country which lies between the Suwanee River on the east, and the Apalachicola on the west.’’ His “‘ observations were chiefly made . . . about twenty miles west of Tallahassee, about thirty miles from the Gulf of Mexico, in latitude about 30° 30’.”” The plant was seen in bloom January 1-5, 1833, and must certainly be the same species as the plant sent Dr. Torrey. Dasystephana tenuifolia is most nearly allied to D. Porphyrio (Gmel.) Small, long known as Gentiana angustifolia Michx. Rafinesque briefly distinguishes the two by certain features, the fs 183 184 PENNELL: PLANTS OF SOUTHERN UNITED STATES most obvious of which is the white corolla of the southern plant. Southern botanical authors treating of the Carolina flora, describe the blue-flowered species, but Chapman, working on the Gulf coast, knew only the white-blooming low-growing plant. 32. f. 1324-1357. ‘Cincinnati. D_1918. Lorenz, A. Nardus stricta in the White Mountains. Rhodora 21: 2, 23. 30 © toig, MacCaughey, V. The pala or mule’s-foot fern (Marattia Douglasit (Presl.) Baker) in the Hawaiian Archipelago. Torreya 19: 1-8. 21 F 1919. McRostie, G. P. Inheritance of anthracnose resistance as indicated by a cross between a resistant and a susceptible bean. Phyto- pathology 9: 141-148. 8 Mr i919. Maxon, W. R. A new Selaginella from Oklahoma and Texas. Proc. Biol. Soc. Washington 31: 171, 172. 30 D 1918. Selaginella Sheldoni Maxon. Maxon, W. R. Notes on American ferns— XI. Am. Fern Jour. 7: 104-106. 23 F 1918—XII. Am. Fern. Jour. 8: 114-121. fl. 6. 20 Ja 1919;—XIII. Am. Fern Jour. 9: 15. Mr 1919. Merrill, E. D. New or noteworthy Philippine plants— XIV. Philip. Jour. Sci. 13: (Bot.) 263-333. S 1918. Eighty-four new species in various genera are described. Miller, L. E. In the wilds of South America. i-xiv + 1-424. New York. 1918. [Illust.] Miller, W. D. A distinction between two Carices. Rhodora 21: 23, 24. 10F 1919. INDEX TO AMERICAN BOTANICAL LITERATURE 231 Mitchell, J. A. Bear clover ( Chamaebatia foliolosa Benth.). Jour. For- est. 27: 39-43. Ja 1919. Morrison, B. Y. An autumn blooming Iris. Jour. Internat. Gard. Club 2: 599-601. Dig18. _ [Illust.] Morse, A. P. Amaranthus Powellii and Digitalis lanata in New Eng- land. Rhodora 20: 203. 27 Ja 1919. Moxley, G. L. Adiantum capillus-veneris L. forma cristatum f. nov. Am. Fern Jour. 9: 27, 28. Mr 1919. Munson, T. V. Improvement of quality in grapes. Proc. Soc. Hort. Sci. 1905: 19-24. Mr 1908. Murrill, W. A. George Francis Atkinson. Jour. N. Y. Bot. Gard. 19° 314, 415. 22018. Nanz, R. S. The Southern limit of Encalypta laciniata. Bryologist 2273.90 F F919. Nash, G. V. Hardy woody plants of the New York Botte! Garden. Jour. N. Y. Bot. Gard. 19: 222-225. S 1918; 293-296. N 1918; 315-318. DD. 918. Nelson, J. C. The name Toxylon again. Am. Bot. 25: 21-23. F 1919. Nicholson, W. E. A reminiscence of the late Dr. Emil Levier. Bryol- ogist 21: 85, 86. 15 Ja 1919. Orton, W. A. On methods of breeding for disease-resistance. Proc. Soc. Hort. Sci. 1907: 28. Mr 1908. Ann. Osmun, A. V. Common potato diseases and their control. Rep. Massachusetts State Board Agr. 65: 125-133. f. 1-8. 1918. Osterhout, G. E. Additions to the flora of Colorado. Bull. Torrey Club 46: 53-56. 25 F 1919. ribuideccaits hastata, Phacelia iI Meriensia icin Oreocarya monosperma and A goseris frondifera, spp. nov., are described. Osterhout, W. J. V. Phe anni of the process of death. Jour. Biol. Chem. 31: 585-589. f. r. 1917. Osterhout, W. J. V. Edward Strasburger (1844-1912). Proc. Am. Acad. Arts & Sci. 51: 927, 929. D 1916 Osterhout, W. J. V. Some aspects of the temperature coefficients of life processes. Jour. Biol. Chem. 32: 23-27. f. Z- 1917- Peirce, G. J. What kind of botany does the world need now? Science, II. 49: 81-84. 24 Ja 1919. Pennell, F. W. Corrections of names of Colombian plants. N. Y. Bot. Gard. 19: 319. D 1918. Jour. 232 INDEX TO AMERICAN BOTANCIAL LITERATURE Pennell, F. W. Concerning duplicate types. Torreya 19:13, 14. 21 F 1919. Pennell, F. W. Some remarks upon Limosella. Torreya 19: 30-32 F 1919. Piper, C. V., & Coe, H.S. Rhizoctonia in lawns and pastures. Phyto- pathology 9: 89-92. pl. 7, 8. 26 F 19109. Pope, W. T. The banyan and some other closely allied species. Ha- waiian Forest. & Agr. 6: 121-129. Mr 1909. [Illust.] Ramaley, F. Xerophytic grasslands at different altitudes in Colorado. Bull. Torrey Club 46: 37-52. f. 1, 2. 25 F 1919. Reddick, D. Vern Bonham Stewart. 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Calif. 8: 10-12. f. 6-8. Ja 1916. . Schacke, M. A. A chromosome difference between the sexes of Sphaerocarpos texanus. Science Il. 49: 218, 219. 28 F 1919. Schlechter, R. Kritische Aufzahlung der bisher aus Zentral-Amerika bekanntgewordenen Orchidaceen. Beih. Bot. Centralb. 36: 321-520. 1918. Includes the new genus Epilyna and 84 new species in various genera. Shamel, A. D. Bud variation in dahlias. Jour. Heredity 9: 362-364. J. 2i, 12... 6 F 166. INDEX TO AMERICAN BOTANICAL LITERATURE 233 Shaw, J. K., & Norton, J. B. The inheritance of seed coat color in garden beans. Massachusetts Agr. Exp. Sta. Bull. 185: 59-104. Ji 1918. Shufeldt, R. W. February—and plant life still sleeps in northern climes. Am. Forest. 25: 868-875. f. 1-14. F 1919. Smith, E. F. The cause of proliferation in Begonia phyllomaniaca. Proc. Nat, Acad. Sdi.g: 36,.37)..15 F i019. Soth, B. H. The cushion pink. Am. Bot. 25:1. F 1919. [Illust.] Spragg, F. A., & Nicholson, J.W. Rosenrye. Jour. 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Torrey CLusB WILLIAMS: DESMATODON Vol. 46 No. 7 BULLETIN OF THE TORREY BOTANICAL CLUB ‘ JULY, 1919 Taxonomic studies in Vernonia and related genera Henry ALLAN GLEASON In preparing the manuscript for the tribe Vernonieae, to be published in the North American Flora, several new species and varieties were recognized and a few nomenclatorial changes became necessary. One of the latter involves the erection of a new genus. Since the form ot the North American Flora demands relatively concise descriptions and permits no critical discussion, the present paper is issued in advance. A variety, in the opinion of the writer, represents a group which is not worthy of specific rank, yet demands recognition in the intensive study of a species. In general, varieties are not ad- mitted into the North American Flora. Specific descriptions are drawn broadly enough to include all the varieties, while the varietal names are cited among the synonyms. The same treat- ment will be used for the varieties published here. Although this paper and the manuscript for the North Amer- ican Flora have been prepared at the New York Botanical Garden and are based primarily on the collections there, the writer has been greatly assisted by material from many American herbaria and takes this opportunity of expressing his thanks: to Mr. C. C. Deam and Mr. J. Lunell for material from their private herbaria; to those in authority at the University of California, the Field Columbian Museum, the University of Illinois, the Iowa Agri- cultural College, the University of Kentucky, the University of [The BULLETIN for June (46: 195-234. pl. 9-12) was issued July 8, 1919] 235 236 GLEASON: TAXONOMIC STUDIES IN VERNONIA Minnesota, the Ohio State University,and the University of Wis- consin for the loan of material; to the directors or curators of the Brooklyn Botanic Garden, the Gray Herbarium, the National Herbarium, the New England Botanical Club, and the Phila- delphia Academy of Sciences for the facilities of their herbaria and libraries; to Dr. B. L. Robinson, Dr. J. M. Greenman, Mr. W. R. Maxon, and Mr. E. E. Watson for information, assistance, and criticism; and especially to the University of Michigan, where part of the work was done. VERNONIA BORINQUENSIS Urban In describing the species, Urban took as the type a form with hirsute achenes, resinous-dotted leaves, and short straight ap- pressed hairs on the lower leaf surface. This is the commonest form in American herbaria. Urban’s variety Stahlii covers a much rarer form with glabrous achenes and without resinous dots on the leaves. ' Two other forms also occur, which are deserving of varietal names. Vernonia borinquensis resinosa var. nov. Achenes glabrous; leaves conspicuously dotted with glands and resin on the lower surface: otherwise like the typical form of the species. TYPE: ex herb. E. W. D. Holway, collected at Cayey, Porto Rico, January, 1911, and deposited in the herbarium of the New York Botanical Garden. Vernonia borinquensis hirsuta var. nov. _ Achenes hirsute; leaves densely sericeous-hirsute beneath with bent or curved hairs 2-4 mm. long, nearly or completely concealing the resinous dots; otherwise like the typical form of the species. Type: Britton, Stevens, & Hess 2471, collected in a wooded valley, Rio de Maricao, Porto Rico, at an altitude of 500-600 meters, April 2, 1913, and deposited in the herbarium of the New York Botanical Garden. Field notes indicate that it is a vine, two meters long. The species and its three varieties may be distinguished as follows: £ GLEASON: TAXONOMIC STUDIES IN VERNONIA 237 Achenes hirsute; leaves with resinous dots or glandular pits n the lower surface. Pantene of short, straight, appressed hairs seldom — oO. ot mm. in length. V. borinquensis. Pubes leaf-surface of bent or curved hairs 2-4 mm. long. V. borinquensis hirsuta. Achenes glabrous. Glands on the lower leaf-surface present. V. borinquensis resinosa. Glands on the lower leaf-surface none. V. borinquensis Stahlit VERNONIA SERICEA L. C. Rich. Vernonia phyllostachya (Cass.) Gleason. We are indebted to Ekman for the application of the correct name to this well-known species. Specimens in American her- baria show a great variation in the size, proportion, and density of pubescence of the leaves, without offering legitimate opportunity for the separation of varieties. VERNONIA GNAPHALIIFOLIA Rich. Until the last decade this species was poorly represented in American herbaria. The collections of the New York Botanical Garden now include an ample series of specimens, sufficient to give some idea of the range of variation within the species. On ~ casual inspection, the specimens fall into two groups, characterized by wide and narrow leaves. The latter come from the provinces of Santa Clara, Camaguey, and Oriente, that is, from eastern Cuba; while the former are from Santa Clara, Havana, and Matan- zas. The wide-leaved forms have leaves from 9 X 19 mm. to 21 X 51 mm., and the ratio of length to width varies from 2.1 to 3.4. The leaves of the narrow-leaved specimens range in size from 4 X 23 mm. to 14 X 61 mm., and their ratio from 3.2 to 5-7. They make accordingly a continuous series. In all cases the leaves are somewhat revolute and densely sericeous. beneath. All specimens agree in the characters of inflorescence and flower- structure except one, Shafer 2958, from Holguin, Oriente, which lacks the resinous dots on the principal involucral scales. Another specimen differs so much from the specific type that it may be described as a new variety. 238 GLEASON: TAXONOMIC STUDIES IN VERNONIA Vernonia gnaphaliifolia platyphylla. var. nov. Leaf-blades broadly elliptic-ovate, less than twice as long as wide, flat, not revolute at the margin, closely and finely gray- tomentose beneath; otherwise resembling the typical form of the species. Type: Brition, Cowell, and Shafer 12,933, collected at En- senada de Mora, Oriente, March 26-29, 1912, and deposited in the herbarium of the New York Botanical Garden. It is described as a shrub, one meter high, and is the only broad-leaved representa- tive of the species so far known from eastern Cuba. It is also the only specimen examined within the species without the revolute leaf-margin and with distinctly tomentose pubescence. VERNONIA ICOSANTHA DC. Ekman has pointed out the peculiar nomenclatorial confusion attached to this well-known species of the Lesser Antilles, and has chosen to apply to it the name Vernonia arborescens (L.) Sw. In describing Conyza arborescens, Linnaeus had before him not only the plate of Plumier, portraying the Vernonia of the Lesser Antilles, but also an actual specimen of a different species from Jamaica. A comparison of his text with the plate shows that the description could not have been taken from the plate alone, but was based primarily on the specimen. The latter accordingly becomes the type of the species and retains the specific name arborescens, and V.icosantha remains the first valid name for the species of Martinique and Guadeloupe. Vernonia Shaferi sp. nov. Stem shrubby, 1-2 m. high, the young branches closely cinereous-pubescent, becoming glabrate the second year; leaf- blades ovate-lanceolate or elliptic, the largest 4 X 12 cm., the upper much smaller, all acuminate, entire, acute at the base, dark green, minutely papillose-pubescent, and very sparsely casieouceaetend above, paler green but otherwise the same below; heads about eighteen-flowered, crowded in leafy secund cymes at the ends of the branches of the season; bracteal leaves oblong or oblong-ovate, acute, 5-10 mm. long; involucres broadly turbinate o campanulate, 6 mm. high, the scales rather closely imbricate, erect or appressed, narrowly oblong-lanceolate, acuminate, irreg- ularly pubescent and ciliate, and usually resinous toward the GLEASON: TAXONOMIC STUDIES IN VERNONIA 239 tip; achenes hirsute; pappus white, its bristles 6.5~-7 mm. long, the paleae very irregular in length, as much as 1.5-2 mm. long, minutely erect-ciliate; flowers white or pink. Type: Shafer 172, collected in Montserrat, January 23, 1907, and deposited in the herbarium of the New York Botanical Garden. The same herbarium also contains sheets of three other collec- tions made on the same expedition, Shafer 589, 650, and 661. This handsome species is obviously closely related to V. longi- folia Pers., as shown by the shape and pubescence of the leaves, the inflorescence, and the character of the involucral scales. It is distinguished from that species at sight by the white pappus, as well as by the slightly larger heads, the much larger pappus- bristles, which are only 4-5 mm. long in V. longifolia, and the unusually long, barely ciliate paleae. It isa pleasure to name the species in honor of its first collector, the late John A. Shafer, who discovered several other interesting Vernoniae in the West Indies. VERNONIA RACEMOSA Delp. Vernonia racemosa Delp. Mem. Accad. Torino II. 14: 396. 1854. Vernonia araripensis Gleason (in part), Bull. N. Y. Bot. Gard. 4: 181. 1906, not Gardn. Vernonia sericea L..C. Rich. subsp. racemosa Ekman, Ark. Bot. 13:85. 1914. The two sheets in the herbarium of the New York Botanical Garden, which were referred by Ekman to V. racemosa as a sub- species of V. sericea, differ in certain features from V. sericea, with which they are not associated geographically, and agree better in general character with the linear-leaved species of Hispaniola and adjacent Cuba. The species-group Arborescentes, which in- cludes V. sericea, has leaves of a broad type, not revolute, and comparatively few spreading cymes near the ends of the branches, producing a rather short and broad inflorescence. V. racemosa and its allied species have revolute, linear leaves and: narrow, elongate inflorescences, composed of relatively short and few- headed cymes distributed over a considerable length of the axis. While this latter character is largely one of habit, the narrow revo- lute leaves afford a ready and accurate means of distinguishing it and its allies from the Arborescentes. 240 GLEASON: TAXONOMIC STUDIES IN VERNONIA VERNONIA RIGIDA Sw. Collected originally by Swartz in Jamaica in the middle of the eighteenth century, this excellent species has since been practically lost. Few botanists have examined Swartz’s originals and as a result the name V. rigida has been applied by various collectors and students to several entirely different species. The genuine V. rigida was finally rediscovered by Wm. Harris at Upper Claren- don, Jamaica, December 27, 1917, and good specimens are now deposited in the herbaria of the Field Columbian Museum and the New York Botanical Garden. The species is obviously allied with the well-known Jamaican V. acuminata Less. Vernonia Sagraeana angusticeps (Ekman) var. nov. Vernonia angusticeps Ekman, Ark. Bot. 13:14. 1914. The sheet of Wright 284, on which the species of Ekman was based, in the Gray Herbarium agrees with the species in every particular except the small number (twelve) of flowers in the head and the consequently more nearly cylindrical involucre. SPECIES-GROUP BUXIFOLIAE In the mountains of Haiti and Santo Domingo occurs a group of three poorly known species of Vernonia. Only eight speci- mens have been examined in American herbaria by the writer, while Ekman mentions twelve in various European collections. One of these, the last to be described, is V. Tuerckheimii Urban, which seems to a well-marked and easily recognized species. Another is a form with small leaves, averaging only 7 X 12 mm., and a short, cylindrical or ellipsoid involucre 3-4 mm. in diameter, or spreading under pressure to 5 mm. For convenience this will be designated here as species A. The third, here referred to as species B, has leaves averaging 10 X 23 mm., and a turbinate or _ almost salviform involucre, spreading at its mouth to a width of 7-9 mm. even when not pressed. While the involucre of species A offers no noteworthy feature, that of species B is remarkable for its imbrication. It is composed of five vertical but gently spiral rows of scales, with ten to thirteen scales in each row, beautifully imbricated, and with the outer ones gradually reduced at the acuminate base of the head. Ekman had before him tly tah as GLEASON: TAXONOMIC STUDIES IN VERNONIA 241 only species A; at least he cites two collections which cover that species in the New York collection and fails to mention the peculiar involucre of species B, which is utterly unlike that of any other North American species. This species A is designated by him V. buxifolia (Cass.) Less., although he has not seen Cassini’s type, and as a synonym he adds V. domingensis (Spreng.) DC., the type of which he has seen. These two names have been re- garded as synonymous for over eighty years. The writer, in 1906, considered that species B was the true V. buxifolia, and described species A as new under the name V. montana. From this de- scription and the cited specimen Ekman recognized that V- montana was co-specific with the plants which he had examined, and reduced V. montana to synonymy. At the same time he was unable to match Gleason’s description of V. buxifolia (species B) with anything he had seen in European collections and decided that it was probably a new species. When he examined the New York material of these in 1914 he annotated the sheets of species A as genuine V. buxifolia and those of species B as “V. buxifolia forma.” . Cassini described the involucre of Lepidaploa buxifolia as turbinate and regularly imbricated, with an assemblage of short rounded scales covering the summit of the peduncle at the base of the head. In this feature it can agree only with species B. De Candolle, in examining the type of Sprengel’s Proustia domin- gensis, used terms which certainly apply to species B, but which do not emphasize the peculiar involucre. One can scarcely imagine that he would have passed by such a striking feature if the specimen had exhibited it. The writer is therefore convinced of the justice of maintaining V. buxifolia for species B, as he did in his revision in 1906. For species A, he must be guided by the negative evidence of De Candolle and the positive statement of Ekman in regard to V. domingensis, use that name for species A, and relegate his own V. montana to synonymy. Vernonia morelana sp. nov. A shrub 3-5 m. high, branching above; stems striate, closely gray-tomentose, becoming glabrate with age; leaf-blade firm, dull-green, ovate-oblong, 3 X 7.5 cm., on tomentose petioles 8mm. 242 GLEASON: TAXONOMIC STUDIES IN VERNONIA long, entire or with a few low teeth, obtuse or rounded at the base and apex, closely scabrous-pubescent above, finely gray-tomentose beneath; upper and rameal leaves similar but smaller and more densely tomentose, those in the cymes broadly ovate to subrotund, 5-15 mm. long; cymes freely branching, forming a hemispheric cluster 2 dm. wide at the end of the branches; heads 21-flowered; involucre campanulate, 4-4.5 mm. high, its scales regularly im- bricate, all appressed or barely spreading at the tip, outer and middle scales ovate to ovate-oblong, sharply acute or cuspidate, tomentose-ciliate and often puberulent on the back, inner scales lanceolate, sharply acute or subacuminate, 4 mm. long, nearly or quite glabrous; achenes thinly pubescent and densely resinous- glandular; pappus white or very pale tawny, the bristles 6 mm. long, the paleae narrow, 0.6-0.8 mm. long. Type: Pringle 7697, collected by streams, Cuernavaca, More- los, Mexico, altitude 5,000 feet, March 16, 1899, and deposited in the herbarium of the New York Botanical Garden. While this species clearly resembles the other members of the species-group Deppeanae in leaf-habit, inflorescence, and pubes- cence, it is distinct in its glandular achene, a structure not ob- served elsewhere in the group. It resembles V. canescens H.B.K. in its white pappus, but differs in its pubescence and broader - involucral scales. It approaches V. Deppeana Less. in its leaf- pubescence, but differs in its white pappus, sharper scales, larger involucre, and broader paleae. It is even more widely separated from the other species of the group. Vernonia salamana sp. nov. bove, finely pubescent beneath, especially on the prominent campanulate, 4-5 mm. high, its scales loosely but regularly im- bricate, pale green with a darker spot near the tip, ciliate, puberu- GLEASON: TAXONOMIC STUDIES IN VERNONIA 243 lent on the back, obtuse to broadly rounded at the tip, the mid- vein becoming prominent near the apex and usually prolonged into a minute mucro; achene minutely pubescent on the ridges; pappus pale tawny, its bristles 4 mm. long, the paleae narrowly linear, about 0.4 mm. long. Type: Maxon & Hay 3385, collected on dry plains near Salama, Guatemala, January 22, 1905, and deposited in the herbarium of ~ the New York Botanical Garden. Vernonia salamana apparently finds its nearest relative in the well-known V. patens H.B.K., with which it agrees in its achenes, pappus, involucral scales, olivaceous leaves, and the finely pubes- cent lower leaf-surface. The shape of the leaf-blade is unlike that of V. patens and is closer to that of V. Deppeana Less. The pubes- cence on the upper side of the leaf and the prominent venation are distinctive. VERNONIA MOLLIS H.B.K. A plant collected by the Brothers Seler, number 3377, has been identified by Hieronymus as V. mollis H.B.K. and distributed to American herbaria under that name. Ekman has considered the specimen as V. canescens H.B.K., to which it is obviously closely related, but from which it differs in the flat leaves and the tomen- tose lower leaf-surface, in its involucral scales, which are all sub- ulate, and in its general habit. At the same time, it is doubtful if it is the true V. mollis. This is a Colombian species, described as having leaves sericeo-lanuginous beneath. This character is met with in certain Colombian specimens in the New York collec- tions. : Vernonia ctenophora sp. nov. . Stem herbaceous, at least 4 dm. high, finely striate, thinly pubescent and resinous-dotted; leaves sessile or with petioles 2-4 mm. long; leaf-blades thin, ovate-lanceolate, entire, acuminate, obtuse or rounded at the base, as much as 5 cm. long by 2 cm. wide or the upper somewhat smaller, thinly puberulent and con- spicuously glandular-dotted above, closely and finely gray- pubescent and resinous on the surface beneath and sparsely pubescent on the midrib and the obscure lateral veins; inflores- cence of two or three elongated, erect, terminal or subterminal cymes; bracteal leaves resembling the cauline and _ progres- sively " smaller, the upper only 15-20 mm. long; heads 20- 35 mm. apart, sessile, 18-21-flowered; involucre campanulate, 244 GLEASON: TAXONOMIC STUDIES IN VERNONIA 6-7 mm. high, its scales irregularly but rather closely imbricate, the outer and middle with triangular-ovate or oblong appressed bases and long subulate tips, the inner linear-oblong, rather abruptly acuminate and most of them subulate-tipped, and all thinly pubescent with dark-colored hairs and sparsely resinous; achenes thinly pubescent, sharply ribbed, 1.5 mm. long; pappus - white, its bristles 4 mm. long, prominently barbellate, the paleae little wider than the bristles, 0.6 mm. long, sharply ciliate with salient teeth. Type: E. A. Goldman 508, collected at Apazota, Campeche, Mexico, December 30, 1900, and deposited in the United States National Herbarium as sheet 396871. Vernonia ctenophora is a member of the species-group Argyro- pappae, as indicated by the resinous-dotted leaves and the long subulate involucral scales. Within the group it is most closely related to V. hirsutivena Gleason, from which it is distinguished by the conspicuously barbellate pappus bristles, the sharply ciliate and shorter paleae, the thinly pubescent involucre and achenes, the comparatively thin pubescence on the veins, and the numerous resin-dots on the upper surface of the leaf. VERNONIA MISSURICA Raf. Vernonia illinoensis Gleason, Bull. N. Y. Bot. Gard. 4: 211. 1906. Throughout its wide range, which is more extensive and covers more diverse environmental conditions than that of any other species in the United States, V. missurica exhibits a considerable variation in structure. This variation pertains chiefly to the inflorescence and leaf-pubescence and less to the characters of involucre and achene. | Judged from herbarium evidence and field experience, the species is best developed in Indiana, Illinois, and northern Mis- souri, where it is by all odds the most common species of the genus. Here the inflorescence is broad, freely branched, with many heads, and relatively flat, and the leaves are thinly but closely tomentose beneath with cinereous multilocular hairs which cover the surface and veins alike. Farther to the north- east, at the border of its range in Michigan and Ontario, the mullti- locular hairs on the leaf-surface are relatively fewer and usually GLEASON: TAXONOMIC STUDIES IN VERNONIA 245 replaced by short straight conical hairs. The leaves are smaller and proportionately narrower, and frequently with a basal taper. Because of these structures, the species is frequently mistaken for V. altissima Nutt., from which it may be distinguished by the ‘resin-dots on the leaf, involucral scales, and achenes. This extreme form has been described by Daniels under the name V. michiganensis, and the same thing from Ontario appears in several herbaria under another unpublished name. Throughout this whole region, from Michigan to Missouri, the plants exhibit generally rounded to subacute, purple involucral scales, imbricate in relatively few series, and a purple pappus. This is the form described by the writer as V. allinoensis. | West of this region the species is much less common, and the herbarium material has been collected in widely scattered localities as far south as southern Texas. In general, the collections from west of the Mississippi and south of the Missouri Rivers have a loose open inflorescence, sharper and frequently smaller involucral scales, imbricate in relatively many series, and frequently green instead of purple, and a pappus which soon becomes tawny in color when exposed to light. But these differences are not con- stant nor even coincident on the same plant, so that even well- marked varieties can not be accurately distinguished. Yet in four cases out of five the geographical origin can be correctly guessed merely by a glance at the involucre of the plant. Another area in which a form of the species occurs is the coastal region of southern Mississippi and Alabama. Whether similar plants also occur in southern Louisiana and southeastern Texas is not known definitely, but the inference is that they do. This extreme southeastern form differs in certain features from the species, so that it is recognizable at a glance, and it may be de- scribed as a variety. - Vernonia missurica austroriparia var. nov. Inflorescence more or less elongate, very loose, open, irregular, and few-headed; leaf-blades broadest distinctly above the middle, thinly tomentose beneath or merely pubescent; resinous glands on the leaves, scales, and achenes as in the typical form of the species. ; 246 GLEASON: TAXONOMIC STUDIES IN VERNONIA Type: Tracy 8015, collected at Tensaw, Alabama, August 18, 1904, and deposited in the herbarium of the New York Botanical Garden. Other specimens are: Tracy 6970 from Ocean: Springs, Mississippi; Tracy 478¢ 4780 from Coopolis, Mississippi; and various specimens collected “by Mohr at or near Mobile, Alabama. In some specimens the leaf-pubescence is reduced in amount until the leaf resembles that of V. altissima. Such plants have been in- cluded under this variety because of the size of the heads, the char- acter of the involucre and inflorescence, and the presence of resin. _ Vernonia aborigina sp. nov. Stem sc eR striate or ribbed, covered with a brown tomentum becoming thicker above; leaf-blades ovate- lanceolate, conuests denticulate with low ascending callous teeth, acuminate, narrowed below into an obtuse or rounded sessile or subsessile base, 8-15 cm. long, scabrous above with short papillose hairs, densely brown-tomentose beneath; inflorescence rather small and compact, 11-13 cm. wide and bearing relatively few (about 30) heads; bracteal leaves on the cyme-branches lance- oblong, 10-15 mm. long; heads large, in fruit about 12 mm. high, containing in the single head counted 68 flowers; involucre broadly rounded at the base, 7-8 mm. high, expanding to 16-18 mm. wide at maturity; involucral scales closely and regularly imbricated, the outer minute and triangular and all squarrose or recurved at the tip, acute, sparingly ciliate along the brown margin, resinous- glandular and thinly puberulent along the purple central two thirds, elsewhere green and glabrous, tipped with a rather promi- nent carinate midvein which is frequently prolonged into a short mucro; achenes 3.5 mm. long, olive in color, with low ridges and broad flat furrows, conspicuously glandular in the furrows, minutely and sparsely puberulent on the ridges; pappus-bristles reddish-tawny, almost plumose below, merely barbellate above, 7 mm. long, paleae as long as the diameter of the achene, narrowly linear, equaling in width or barely wider than the bristles The type was collected by P. H. Rolfs in Oklahoma, west of Fort Smith, Arkansas, August, 1891, and is deposited in the herbarium of Iowa Agricultural College as sheet number 32272. No other material has been examined. The description of this new species is offered with considerable reluctance, because of the great variabilty of the western species and their known tendency to hybridize. In several features it differs from all other western species. The large heads, the un- GLEASON: TAXONOMIC STUDIES IN VERNONIA 247 usual number of flowers in each head, the olive-colored, glandular achenes, and the reddish, conspicuously barbellate pappus dis- tinguish it from all others. The squarrose scales separate it from all others except V. Baldwini Torr., the very narrow paleae from all except V. interior Small and the species-group Fasciculatae. It is to be hoped that further collecting in a somewhat neglected region may bring to light additional material. - Vernonia fasciculata nebraskensis var. nov. Leaves shorter than in the typical form of the species, narrowly lanceolate, denticulate, acute, pale-green or yellowish green; heads closely crowded. Type: Rydberg 5400, collected in Kearney County, Nebraska, July 14, 1900, and deposited in the herbarium of the New York Botanical Garden. Numerous other sheets occur in all larger herbaria; in fact, almost all of the Nebraska specimens labeled V. fasciculata are to be referred to this variety. The species proper occurs only along the Missouri River, so far as known to - the writer, while the variety extends westward more than half the length of the state. Although the brief varietal description apparently offers but little evidence for the separation of a variety, nevertheless the Nebraska specimens in any large herbaria all look alike, all look different from the rest of the species, and can be separated at a glance even by a person not familiar with the genus, as the writer has been able to demonstrate. VERNONIA ALTISSIMA PUBESCENS (Morris) Daniels Inner involucral scales tipped with a short, straight,’ flat, erect, linear tip, not over 2 mm. long. While Morris’s variety was originally separated primarily by the character of the foliage, this is, as has been pointed out by Blake, due to some unusual pathological or teratological condition. The shape of the scales, on the other hand, is found as described - above not only in Morris’s type but in several other herbarium sheets from the Alleghenian region, from Pennsylvania and Ohio south to Alabama and South Carolina, and in isolated collections of Wilkinson from Mansfield, Ohio. 248 GLEASON: TAXONOMIC STUDIES IN VERNONIA “ Vernonia altissima brevipappa var. nov. Lower surface of the leaf-blades with a few multilocular hairs along the veins; paleae only 0.1-0.3 mm. long; otherwise as in the typical form of the species. Type: collected by J. Schneck at Mt. Carmel, Illinois, August 13, 1891, and deposited in the herbarium of the University of Illinois. / Vernonia altissima laxa var. nov. Heads smaller than in the species, 13~21-flowered; inflorescence broad, irregular, very loose and open; inner involucral scales often apiculate, the middle scales frequently sharply acute; resin-dots none; leaf and pubescence as in the typical form of the species. Type: Harper 1936, collected at Newton, Georgia, August 19, 1903, and deposited in the herbarium of the New York Botanical Garden; duplicates of the type are in the Gray Herbarium and the herbarium of the Field Columbian Museum. Other speci- mens of this southern or coastal plain variety are Tracy 8046 and Eggleston 5142. / Vernonia flaccidifolia angustifolia var. nov. Leaves narrowly lanceolate, denticulate, only 1-2 cm. wide; otherwise as in the typical form of the species TyPE: ex herb. Torrey, collected in Alabama and deposited in the herbarium of Columbia University; another sheet in the New York collections is from Georgia, ex herb. Chapman. 7 Vernonia ovalifolia purpurea var. nov. Leaves with numerous multilocular hairs on the veins of the lower surface, the upper leaves subtending the branches truncate on rounded at the sessile base; pappus purple; otherwise as in the typical form of the species. Tyre: F. S. & E. S. Earle 99, collected at Auburn, Alabama, July 22, 1899, and deposited in the herbarium of the New York Botanical Garden. Vernonia jucunda sp. nov. Stem apparently herbaceous, at least 4 dm. high, erect, sparingly branched, densely villous when young, becoming floccose when older and glabrate at about 3 dm. from. the summit, faintly GLEASON: TAXONOMIC STUDIES IN VERNONIA 249 striate and reddish brown under the pubescence; leaf-blades ovate- oblong, as much as 38 X 80 mm., narrowed toward the sessile base or into a short margined petiole, undulate and irregular at the entire margin or rarely with a few low salient teeth, frequently a little revolute, acute at the apex or short-acuminate into a small subulate tip, dark-green and rugose above with impressed veins and scabrous with papillose hairs or hair-bases, gray or nearly white beneath with a close fine tomentum; lateral veins promi- nent, ascending and straight almost to the leaf-margin; upper and bracteal leaves similar but smaller; heads five-flowered, in a sympo- dial raceme, standing opposite and a little (2-3 mm.) below a bracteal leaf which later bears secondary heads in its axil, primary heads eight to ten; involucre 8 mm. high, 2.5 mm. wide, with its stiff scales imbricate and appressed at the base and squarrose at the tip, lanceolate-oblong to ovate-oblong, broadest below or near the middle and long-acuminate into a subulate, glabrous, terete, callous tip, the outermost green, one-half the length of the purple inner ones, and all papillose-villous with erect hairs on the exposed portion; corolla apparently pale purple, its tube glabrous, not ampliate above, 5 mm. long, its lobes glabrous, 3 mm. long by 0.6 mm. wide, with parallel sides and triangular tip; filaments glabrous, attached at two thirds the height of the tube; anthers 2.8 mm. long, minutely rounded at the triangular tip, their alge bases ee mm. long; style hairy along the upper I.4 m its branches 1.4 mm. long, tapering, hairy on the outer side; ioe 2.5 mm. long, shallowly ten-ribbed, pubescent with shott erect hairs on the ridges; pappus-bristles pale tawny, 6.5 mm. long, barbellate; paleae linear-lanceolate, 1.0-1.1 mm. long by 0.09— 0.18 mm. wide, trough-shaped and pubescent on the inner face. Tyre: Purpus 7060, collected in the Sierra de Tonala of Chi- apas, Mexico, October, 1913, and deposited in the herbarium of the University of California as number 173434. Other sheets of the same collection are in other American herbaria and agree in every particular with the type. Vernonia jucunda is the first species of the section Stenocepha- lum to be discovered in North America. Other members of the section are South American; one species, probably undescribed, occurs in Colombia and a number in Brazil. The section is characterized by few-flowered heads set a short distance below the bracteal leaf, by an involucre constricted at the throat and com- of subulate, more or less squarrose scales, and by leaves which are usually revolute and tomentose beneath. 250 GLEASON: TAXONOMIC STUDIES IN VERNONIA Among the many interesting plants collected by Charles Wright in Cuba, his number 2789, described by Grisebach as Vernonia lepidota, is one of the least known. Wright described it as a suffruticose plant, ascending on bushes to a height of about 3 meters, with purple flowers. The Gray Herbarium contains a large specimen which shows the foliar characters very well, but is too immature for a careful study of the floral structures, achenes, or pappus. Ekman examined three sheets of the same number in European herbaria, including Grisebach’s type, and has published the first good description of its reproductive structures. A few of his observations have been verified at the Gray Herbarium. Ekman points out that its anthers and styles agree with those of the genus Vernonia, but that its pappus is entirely different. The inner pappus is composed of not more than seven flattened bristles, and the outer of scales which are coalescent into a cylin- drical tube with lacerose margin. This feature alone is sufficient to warrant the erection of a genus for it, which may appropriately be named in Ekman’s honor. EKMANIA gen. nov. Inflorescence a corymbiform cluster, freely branched and beset with petiolate bract-like leaves; heads homogamous, few- flowered; involucre of a few series of closely appressed scales; corolla glandular without; style and anthers as in Vernonia; achene glabrous, ten-ribbed; pappus biseriate, the outer of a cylindrical tube with lacerose margin, the inner of five to seven stout flattened bristles; stem and foliage lepidote. Type species: Vernonia lepidota Griseb. Ekmania lepidota (Griseb.) comb. nov. Vernonia lepidota Griseb. Cat. Pl. Cub. 145. 1866. Leaf-blades elliptic-oblong, the larger ones 3.5 X 8 cm., the upper smaller, thinly silvery-lepidote above, densely fulvous- lepidote beneath, entire, obtuse or subacute, prominently veined; the larger bracteal leaves petiolate, 10-13 mm. long and one- nerved, the others more crowded distally and gradually reduced in size to short subterete scales 2 mm. long, closely appressed to the involucre and distinguished from it chiefly by their lepidote pubescence; involucre 3 mm. high, its brown scales trough- shaped or boat-shaped, acute, pubescent or scurfy on the back. GLEASON: TAXONOMIC STUDIES IN VERNONIA 251 Oliganthes Milleri (Johnston) comb. nov. Vernonia Milleri Johnston, Proc. Am. Acad. 11: 698. 1905. The structure of the pappus of this little-known species from the island of Margarita indicates clearly its affinity with Oligan-_ thes rather than with Vernonia. Piptocoma rufescens latifolia var. nov. Leaf-blade narrowly ovate or elliptic, two to three times as long as wide, broadly obtuse or rounded at the apex, abruptly narrowed at the base, its tomentum loose and thin; petiole 1 cm. long; involucre larger and its scales more tomentose than in the typical form of the species. Type: Britton, Britton & Shafer 104, from a coastal thicket on Water Island, St. Thomas, January 31 to February 4, 1913, and deposited in the herbarium of the New York Botanical Garden; a second sheet in the same collection is Britton & Shafer 845, from a coastal thicket on Salt Island, Tortola. The species of Elephantopus occurring in the southeastern United States have long been a difficult problem for taxonomists. E. carolinianus Willd. and E. nudatus Gray are quite distinct and have generally been recognized. Gray, in the Synoptical Flora, in 1886, combined E. tomentosus L. with E. elatus Bertol., and in general Bertolini’s species was not recognized until 1901. Baker then regarded it as distinct and gave some valuable com- parative measurements. While the difference in length of in- volucral scales and pappus is distinct in mature heads, it is not always reliable at earlier stages. Possibly for this reason, Baker’s measurements have been neglected by recent authors, and Small has separated the two species chiefly on characters of leaf and pubescence, which are unfortunately exceedingly variable. One diagnostic character has however been overlooked. In E. tomentosus the hairs on the midvein are generally reflexed, while on the leaf-surface proper many or all of the hairs point backward. In E. elatus the hairs of the midvein are longer and stiffer and point distinctly forward. Separated by this character, many specimens usually referred to E. tomentosus are found to belong to the other species, having also the short, heavily invested scales 252 GLEASON: TAXONOMIC STUDIES IN VERNONIA and short pappus ascribed by Baker to E. elatus, and only the leaf-shape of E. tomentosus. These may be described as a va- riety. Elephantopus elatus intermedius var. nov. Leaf-blade elliptic or oblong, abruptly narrowed to the base, less than three times as long as wide; stem comparatively short and sparingly branched ; pappus sometimes as much as 5 mm. long. Type: Tracy 4741, collected at Coopolis, Mississippi, Sep- tember 8, 1898, and deposited in the herbarium of the New York Botanical Garden. The variety is represented in herbaria by numerous specimens, and is distributed from southern Mississippi east along the coastal plain to Georgia and south to Lee and Dade Counties in southern Florida. | NEw York BoTANIcaAL GARDEN - Storied or tier-like structure of certain dicotyledonous woods* SAMUEL J. RECORD A considerable number of dicotyledonous woods are character- ized by a storied or tier-like arrangement of their secondary ele- ments. Such woods exhibit on longitudinal section, typically on the tangential, fine transverse lines, bands or striations commonly called “ripple marks.’’ These markings may be distinct to the unaided eye, may be at or near the limit of vision, or may require the lens. In some cases they may be more readily seen without the lens than with it, or they may be fairly distinct under the lens but indistinct or apparently absent under the compound micro- scope. In specimens with very dark and infiltrated heartwood the markings may be obscured there but show plainly in the sap- wood; usually, though, they appear to best advantage in the heart- wood. In woods with this type of structure the cambial cells are in tangential as well as radial seriation and part or all of the elements to which they give rise preserve this arrangement. Or- dinarily it is only in stems of considerable thickness that the tiers assume a marked regularity, but the diameter a stem must attain before the storied structure becomes pronounced is subject to wide variation, depending not only on the species but also on the rate of growth. In certain stems of Leguminosae and Zygophyl- laceae the tiers were found to be fairly regular in what was appar- ently the second annual ring, less than one eighth inch from the pith. In general, though, this feature is wanting in the twigs of herbarium material. The storied structure may not extend to all of the elements. In the extreme and also the most common form (Zygophyllaceae, Bignoniaceae, Leguminosae in large part, and others) the rays, vessel segments, tracheids, wood fibers, and wood parenchyma strands are all in horizontal seriation. On the tangential surface the cross sections of the rays appear as short, closely spaced, par- * Contribution from the Yale School of Forestry, No. 4. 253 254 RECORD: STRUCTURE OF CERTAIN WOODS allel lines, often of very regular length and arrangement, while separating the tiers is a fine line of lighter or darker shade than the remainder of the wood. These lines are made by the junctions of the tiers of wood cells and vary in appearance according to the elements involved in a particular area, whether fibers with inter- lacing tips or vessel segments and wood parenchyma cells with abrupt terminations. In some instances, also, the local aggre- gations of pits in the fiber walls where the lumina become con- stricted tend to increase the refraction there and make the lines more distinct. According to Von Héhnel (1, p. 33) this is very pronounced in Bocoa provacensis Aubl. (Inocarpus edulis Forst.); his findings are confirmed by Wiesner (5, pp. 15, 950), who figures the pits. In the cases investigated by the present writer such pit-areas were noted in several woods, as for example, in Machae- rium, Physocalymma(?), and Erythrina. Where the rays are all storied they occupy the median portion of each tier, the height of which is usually considerably greater than the height of the rays. Consequently there is room for considerable variation in the height of the rays without interrupt- ing the regularity of the transverse markings. It is not uncom- mon to find some of the rays grown together at the margins and thus occupying two or three stories, as for example, in Cercis, Crescentia, Ormosia and Ougenia. Where the rays are in perfect seriation, a section between two tiers misses them completely. In most instances, however, such a section shows rayless gaps, the width of which depends upon the regularity of stories. In some storied woods, especially in the Bombacaceae, Mal- vaceae and Sterculiaceae, the rays are of two general sorts, large and small. Here only the low rays are in seriation and they may be so few, comparatively, and so over-shadowed by the large rays that the ‘‘ripple marks”’ are indistinct except in proper light and may be easily overlooked in casual inspection. In some of these woods the markings may be plainer without the lens than with it and, because of the very limited field under observation, may not be distinguishable at all under the compound microscope. In both of these types the elements other than the rays are normally storied. The vessel segments and vascular tracheids correspond in length to the height of each tier, though occasional RECORD: STRUCTURE OF CERTAIN WOODS 205 short segments are found which appear to be subdivisions. The wood parenchyma strands show a uniformity in length with the vessel segments, but in a few cases where parenchyma is scanty the vessel segments and tracheids (when present) seem to be en- tirely responsible for the ‘‘ripple marks.” This was found to be the case in Dalea spinosa Gray, Artemisia tridentata Nutt., and Bigelovia graveolens Gray. The number of parenchyma cells per strand is not constant but in most of the woods investigated was found to be two or four; sometimes only one (an intermediate or substitute fiber). Itis not uncommon to find that the metatracheal strands are composed of two cells and the paratracheal of four. Where the parenchyma is abundant, the component cells of the strands may be uniformly disposed and thus give rise to a secondary seriation which, es- pecially if the cells are large, is readily visible under the lens. In this case the height of the tiers in only one half or one fourth that of the ordinary tiers. In species of Bombax, Ceiba (Eriodendron) and Heliocarpus the number of cells per strand is four; in Char- pentiera, Diphysa and Lonchocarpus, two; in Gossypium and Ptero- cymbium, two in the metatracheal and four in the paratrachea'. The same structure obtains in some of the finer-textured woods, but the small size of the cells usually renders it indistinct or in- visible under the lens. It was found fairly distinct in Ichthyo- methia piscipula (L.) Kuntze (mostly two cells per strand) and less so in Tabebuia Guayacan Hemsl. (four cells per strand). The libriform fibers and fiber-tracheids (collectively, wood fibers) may or may not show distinct seriation in the tangential section. In all ordinary cases the fibers are much longer than the vessel segments or the height of the tiers. In many cases there is a widened middle portion equal to about one third of the total length of the fiber and corresponding to the length of the original cambial cell. The attenuated ends of the fibers of one tier are forced during elongation between the fibers of the tiers immediately above and below and, in consequence, a cross section through the middle portion of a tier will show (under the compound micro- scope) alternate rows of large and small cells, the latter being twice as many as the large ones. A section through the junction of two tiers will show fewer cells and they will be more nearly uniform as 256 RECORD: STRUCTURE OF CERTAIN WOODS in size. In many woods the fibers show a gradual instead of ab- rupt diminution in caliber and the feature just mentioned is absent or indistinct. It may require careful maceration to deter- mine whether or not the fibers are storied, particularly in woods with irregular rays. The fact that the fibers are so interlaced and dove-tailed together prevents woods with storied structure from being weakened or made brittle thereby. In Herminiera elaphroxylon Guill. & Perr., the wood cells (" pali- sade tracheids’’ of Jaensch, 3, p. 269) are parenchymatous and arranged in regular stories. Some of the cells are subdivided but not very regularly. The low uniseriate rays are storied while the large ones (which may contain vessels) are not. The vessel seg- ments are mostly of the same length as the height of the tiers but according to Wiesner (5, p. 20) some of them may be subdivided. The wood fibers, which are distributed in narrow layers, are three times the length of the other elements and have localized pit areas at the constrictions. Solereder (6, p. 276) figures Aeschynomene sp. with the parenchymatous wood cells much pitted at the ends. The present writer did not have opportunity to study woods of this genus but found much the same structure in Erythrina spp., except that all or nearly all of the rays are large and the wood fibers in the scattered bands are apparently not storied. The parenchymatous cells making up the ground mass are large, thin-walled, blunt- ended, and usually subdivided into two cells about the vessels. The markings vary from distinct in some specimens to barely vis- ible without lens in others. Where the elements of the wood are definitely storied it is, of course, to be expected that the same arrangement will likewise appear in the secondary phloem. In the limited number of stems with bark examined by the writer this was found to be the case and in some instances, Dalea spinosa for example, the feature was more distinct on the inner surface of the bark than in the wood- An unusual case was found in Olneya tesota Gray, where the inner surface of the bark exhibits, under the lens, fine but distinct and fairly regular cross-lines (150-160 per inch) without, apparently, any corresponding structure in the wood. Under the compound microscope the vessel segments and wood parenchyma strands are indistinctly storied, but the rays and apparently the fibers are RECORD: STRUCTURE OF CERTAIN WOODS aoe not, while the vertical elements of the secondary phloem are in horizontal seriation. The distinctness of the cross-lines in the inner layers of phloem was found to be largely due to highly re- fractive callus on the sieve plates. The storied structure of wood occurs throughout a consider- able range of families and orders as will appear from the following list, which is arranged according to the Engler and Gilg (1912) system of classification. The first numeral following the family refers to the number of different genera in which the feature has been noted by the writer; that in parentheses, to the total number reported. . UrtIcCALES: Ulmaceae, 1 (1); Moraceae, I (I). ‘ oe Amarantaceae, I (I). : Moringaceae (1). . ROSALES: faes (7); Leguminosae, 4o (51). . GERANIALES: Zygophyllaceae, 3 (4); Rutaceae, 1 (1); Simarubaceae, 3 (4); Meliaceae, 6 (7). 6. SAPINDALES: Hippocastanaceae, 1 (1); Sapindaceae (1). 7. MALVALEs: Tiliaceae, 5 (8); Malvaceae, 4 (4); Bombacaceae, 3 (3); Ster- culiaceae, 7 (8). 8. MyRTIFLORAE: Lythraceae, I? (12). 9. EBENALES: Ebenaceae, I (1). . TUBIFLORAE: Bignoniaceae, 3 (3). Iz. CAMPANULATAE: Compositae, 3 (3). ‘ npBW DN H ae q ° & Ps o Hw ° Consideration of the above list leads to the conclusion that there is no close correlation between the form of wood structure under consideration and the various types of floral organization which serve as the basis for the classification followed. There seems to be a correlation, however, between this storied structure and certain types of differentiation of the wood elements. The vessel segments have abrupt ends and simple perforations, and the elongated or scalariform type of pitting is rarely found and then only in the tracheidal vessels and vascular tracheids. The wood fibers are for the most part provided with simple pits or with pits having only small and indistinct borders. The pits between vessels and parenchyma are usually small and _half- bordered. It is also interesting to note that with very few ex- ceptions (notably Aesculus, Cercis, Cytisus, Diospyros and Tilia) the woods are of tropical or subtropical origin. There is no family, with the possible exception of the Zygo- 258 RECORD: STRUCTURE OF CERTAIN WOODS phyllaceae (only a few woods of which were studied), in which all of the woods are characterized by horizontal ranking of their elements. In many cases it appears to be a generic character but more often it is specific only. A few instances have been noted where the feature is not constant for a given species and may be of only local occurrence in the same specimen. One of the best examples of this is furnished by Swietenia, where ‘“‘ripple marks”’ exhibit wide variation in regularity and occurrence and are about as likely to be absent as present. In general, however, the con- stancy of the feature seems to bear a direct relation to the degree of regularity of the cross-markings. It is most dependable when all of the elements are involved and especially when the rays are fine ahd of uniform height. In this investigation attention has been given to the elements storied, the regularity and visibility of the transverse markings and to the height of the tiers in each wood examined. In most in- stances only the gross and lens characters are given since they are most readily employed in practice. There are so many factors entering into the question of distinctness or visibility that the particular observations of the writer may not always apply. For instance, in Guaiacum the markings, though extremely fine, are distinct under the lens in the sapwood and may be indistinct or even invisible in very resinous heartwood. Usually the visibility is materially increased by moistening the surface, but in some in- stances the opposite effect is produced. According to Von Hohnel (2, p. 2) the distance between the cross-lines varies from one half to one eleventh of a millimeter, and for a given species is constant. In order to determine whether or not there is a definite tier-height for each species the present writer counted the number of tiers per inch of length on the tangential section of every specimen examined. An inch, divided into quarters, was marked off and the count and recounts made under the lens for the entire inch, the quarter-marks serving as an additional check. In the larger specimens, at least two dif- ferent inches were counted. In some instances the markings, be- ing fairly distinct without a lens but indistinct with it, were counted with the naked eye or with the aid of a reading-glass. Where the rays are not in seriation the cross-lines are likely to be RECORD: STRUCTURE OF CERTAIN WOODS 259 indistinct, and the most satisfactory results can be secured by counting the vessel segments. In the Zygophyllaceae the lines, though very uniform and readily visible under the lens, are so very closely spaced (about 250 per inch) that is it practically im- possible to make an accurate count without the use of the com- pound microscope. By using low power and reflected light the smooth surface of a small specimen can be readily examined and measurements made without preparing thin sections. Considerable variation in tier-height was found not only in different specimens of the same species but also in the same speci- men. Even within a given inch of length the number per quarter may show a variation of from two to five as a result of the occa- sional to frequent bifurcation of the tiers, with consequent narrow- ing at the point of branching. The greatest variation in tier height within a species was noted in Diospyros virginiana L., where the counts per inch on eleven different specimens were as follows: 57, 57, 58-60, 60, 61, 62, 63, 66-68, 70-72, 72-74, 81-82; a maximum difference of 25 per inch, or nearly 45 per cent on the basis of the minimum. In only one other case did the variation exceed 20 per cent, and almost always it was less than 10 per cent. In 86 woods of the Leguminosae the number of markings per inch were as follows: under 100, 29 per cent; 100 up to 125, 34 per cent; 125 up to 150, 21 per cent; 150 to 190, I2 per cent. Nearly every wood showing between 125 and 190 “ripple marks”’ per inch belongs to the Leguminosae. The investigated woods of the Zygophyllaceae have considerably more than 200 per inch, mostly about 250, and may, by means of this feature alone, be readily separated from all of the others. This fact has proved of practical value in distinguishing the wood of true lignum-vitae from its various substitutes. The present investigation fails to confirm certain findings of other writers. Von Hdhnel (1, p. 38) states that the rays are storied in Parkinsonia, but there was no evidence of this in the specimens examined by the writer. The grenadillo wood is re- ferred to as “Inga vera W.?” (p. 39) but this is probably incor- rect, inasmuch as various specimens of Inga spp. were found to be without any tendency to storied structure. The name “ grena- dillo’”’ is applied to several woods, one of which is Brya Ebenus DC. Another wood with “ripple marks” was provisionally identified 260 RECORD: STRUCTURE OF CERTAIN WOODS by the same author as Conocarpus erecta (Combretaceae), but au- thentic specimens of this species are wholly without this structure, as are all other representatives of the family that were examined. Wiesner (5, p. 997) states that in Olea europea L. the rays (under the lens) produce a fine wavy cross-striping on the tangential section. Specimens of the wood examined by the writer exhibit no tendency to “ripple marks,” and Miiller (Atlas der Holz- structur, 1888, pp. 88, 91) gives as one of the distinguishing fea- tures of this wood the irregularity in size of the rays. All of the foregoing woods are excluded from the table given below. The only representative of the Lythraceae included in the list is Physocalymma scaberrimum Pohl, variously known as Bra- zilian tulip-wood, rose wood, ‘‘pao de rosa,” “‘cego machada,” “grao de porco”’ and ‘‘sebastiao de arruda.”’ Writers all seem to be in agreement in referring this wood to the species mentioned, but the specimens examined by the writer, which are evidently the same as those described by Wiesner (5, p. 975), appear to belong to the Leguminosae. They certainly do not resemble other avail- able material of the Lythraceae. Another wood of which there is some doubt is Ferolia guya- nensis Aubl. (= Parinarium sp.) of the Rosaceae, mentioned by Von Hohnel (1, p. 43), who calls it ‘‘Ficatin- oder Kénigsholz.”’ According to Stone (Timbers of Commerce, 1904, p. 101) this spe- cies supplies the wood from Guiana commonly known as “‘ washiba”’ or bow-wood. Auth®ntic specimens of washiba and bow-wood, collected by the Forestry Officer of British Guiana are Tabebuia spp. The tier height in the latter is about 0.25 mm., while accord- ing to Von Héhnel the height of the “‘ Markstrahlstockwerke”’ in the wood of Ferolia is about 0.15 mm. If Von Hoéhnel’s measure- ment refers only to the height of the rays the discrepancy dis- appears, since this agrees closely with the writer’s observations. No other wood of the Rosaceae has been reported as having a tier-like structure. The satinwood, Chloroxylon. Swietenia DC., is listed with the Rutaceae (following Engler), rather than with the Meliaceae, be- cause the wood so closely resembles those of the Rutaceae and has so little in common with the others. No other representative of the Rutaceae has been found with storied structure, while various representatives of the Meliaceae exhibit a more or less RECORD: STRUCTURE OF CERTAIN WOODS 261 pronounced tendency in this direction. It is interesting to note, however, that the number of tiers per inch in the latter woods (so far as known to the writer) is between 45 and 55, while in the satin- wood it is about 80, Most of the woods with ‘‘ripple marks’’ belong to the Legu- minosae. Following isa list of woods examined by the writer in which the storied structure was not observed, though a few (marked with an asterisk) have been reported by other writers and are included in the list. Some of the more common synonyms are placed in parentheses. These woods are: Adenanthera, Albizzia*, Bowdichia (?), Browneopsis, Cassia*, Cercidium, Cercidophyllum, Cladastris (Copaiba), Copaifera, Cynometra, Daniellia (?), Dimorph- andra, Enterolobium, Erythrophloeum, Eysenhardtia, Gleditsia, Gymnocladus, Haematoxylon, Inga (see page 259), Intsia, Kingio- dendron (?), Lysiloma, Mimosa, Pahudia, Parkia*, Parkinsonia, Peltogyne, Peltophorum (?), Piptadenia, Pithecolobium, Prioria, Prosopis, Pterodon, Robinia, Sindora (Vouacapoua), Zygia (?). In this investigation all of the woods in the extensive collec- tions of the Yale School of Forestry were gone over. Since many of these are from tropical regions, of which our botanical knowledge is far from complete, it is by no means certain that all of the spe- cific identifications are correct, though special effort was made to eliminate errors or indicate the doubtful cases. A considerable number of woods, mostly belonging to the Leguminosae, have been omitted because their botanical status has not been deter- mined. The following table is published with a view of calling attention to an important feature of wood structure and with the hope of stimulating further investigation in this direction. The writer will ale ra assistance in correcting and extending the data. NOTEs: Unee. oe not eared are Pee npape woods nar ia wad raga = of the rays, th I ““Wood p refers to the secondary seriation produced by the individual cells of the wood paren- strands; not visible without lens. ‘‘Fiber pits storied” refers to the hori- scuial seriation of special pit areas on.the wood fibers. here not noted does not ily mean that the feature is absent, since not all of the woods were examined microscopically. ~‘‘ Number of tiers per inch” applies exclusively to measurements by the writer and is the basis for ‘‘tier-height.'"’ Adash indicatesa tendency. Data from other writers are credited to them and are as nearly complete as their descrip- tions will permit. TABLE OF WOODS WITH TIER-LIKE ee ve SHOWING THE ELEMENTS STORIED, THE GECULARITY: AND VISIBILITY OF THE TRANSVERSE S (‘‘RIPPLE MARKS”’), AND MEASUREMENTS OF TIER-HEIGHT | Elements Regularity | Visibility | Measurements storied of lines a ° a BB |2 ach Scientific name : Common name Country g E 28 § c| E 4 2 z $ Tier-height | Number of S| 2/4 S| » bo 5] | | 3} 3 | tterhelgnt) tiers per g S ed Be £13/ 8] 5| > Fy 8 in mm, inch 3) si8a/ 5) Bele al ei zi 2/2 fat | a) 8 3/4 A m AMARANTACEAE : | CHARPENTIERA OVATA Gaudich. Papala Hawaii x| X x x 0.22—0.23 | ITO-I15 BIGNONIACEAE CRESCENTIA CUCURBITINA Black a Florida x x x 0.22—0.23 | 110-115 UJETE L. : Calab Porto Ric x x x 0.2I-0,22| II5-120 TABEBUIA Dowrt- Sur Rose rine Genk “ape ope Pe x Rix x 100 N Hem Yellow guayacan Pana x x x x 90- 92 oe ead C rriseb. Toro-ratai faint x x x 0.17-0.18 | 140-145 “4 P. IR eae prieto West aa Mexico | X x x te) = PENTAPHYLLA Hemsl. (Ro ‘an x x x 9 : HUMANNIANA Urb, Roble colorado Porto Rico x x x 0.29-0.31| 80- 85 sp. (No. 58) Roble Honduras ms —| x m4 0.26-0.28| 90- 96 ed sp. (No. 97) 2 "i x x x “ nt (No. 378) si Colombia x x x 0.32-0.33|} 76- 78 . p. (No. 351) Apamate Venezuela x 5 x .32 ae a Hackia British Guiana x x > 0.26 9 = sp. Washiba Bow-wood x x x 0.24 102-103 TECOMA ARALIACEA DC, Surinam Groenhart a x = 4 0.3 mesial ae Araguaney Venezuela x x x 0.2I-0.22| II5—120 2 * LSUCOKYL Roble Porto Rico x x x 0.24-0.25 | 100-105 de OBTUSATA ae cr. I pe Liais) Lapacho Argentina x x I 120 GIS SdOOM NIVLYaD JO HANLINALS :dkxoory TECOMA ss DC. (Hohnel, 1, p. 41) x x| X 0.25 NS Juss West Indies x x 0.21 a ri ers gt Lapacho Argentina x xX x 0.22—0.23 | I12—-II5 “= sp. (No, 114 " e x * a 0.24 105-107 . - (market _ gay Bethabara x K x 0.28 90 4 p. (2 specime Brazil x x x 0.25-0.26| 95-100 BOMB eo. BoMBAX CYATHOPHORUM (Casar.) Schu cualbaiaged Daher philigg Imbirussi Brazil x| X x x X| 0.63-0.71 | 35- 40 Bompax tec 1 Didu Andamans x| X x x X| 0.42-0.43 | 58- 60 sp.? (Ni 0. oes 3) Tolu ‘ Colombia —| X x x 0.40 63 ed sp.? (No. 1570) Majagua - x x x 0.42-0.45| 55- 60 CEIBA PENTAND na Gartn. (Erioden- dron pina C. eiba Cuba x| X x x 0.42 70: CUMINGIA cite stag Vidal Gapas-gapas Philippine Islands | X 4 x 0.33 75 COMPOSITAE ARTEMISIA TRIDENTATA Nutt. Sage Brush California Xx x x 0.13 200 + BACCHARIS SAROTHROIDES Gray 5 x x —| XX] | 0.20-0.21 | 120-125 BIGELOVIA GRAVEOLENS Gray Colorado x x X|X<} 0.13 + 200 + EBENACEAE DIOSPYROS VIRGINIANA L. (11 specimens) | Persimmon Eastern United States x Pare x 0.30-0.44| 57- 82 HIPPOCASTANACEAE AESCULUS OCTANDRA Marsh. (A. flava | Buckeye Central United Ait.) States x x x X| 0.36-0.43| 58- 68 LEGUMINOSAE oe PROCERA ong : Benth. (Acacia procera Willd.; Héhnel, 1, p. 38) Tropical Asia x 0.20 A ain E sp. (Solereder, 6, p. 275) | Solah India x x x ANDIRA ANTHELMINTHICA Benth. (Hohael, p. 35) Brazil? x x 0.25 SGOOM NIVLYSO JO AMNLINULS :dxoory €9G TABLE OF WOODS WITH TIER-LIKE STRUCTURE, SHOWING THE ELEMENTS STORIED, THE REGULARITY AND VISIBILITY OF THE TRANSVERSE ued LINES (‘‘RIPPLE MARKS’ *), AND MEASUREMENTS OF TIER-HEIGHT—Contin Lp eg mapianty Visibility Measurements glE It id 4 Scientific name Common name Country , 3 er: 8 3 5 2 3 3 unin af g S 5 Sia Eo a a‘ 4) 3 8 Tier-height gee er #| 2/8") 4) g| fi] 8 ei ei ga) aoe ae ci hil = al ed i] 3 £18] 3| Bla #8] 2] 2] 2/2 < 4 4 fe Pd 5 - a pO fe os LEGUMINOSAE—Continued ANDIRA INERMIS A. Yaba Cuba x] x x x 0.26 93- 96 (Wiesner, 5, P- 947) Partridge Wood Tropical America | X x x APULEIA PRABCOX Mart Ibira pere Argentina x x x 0.24-0.26| 95-105 : : be x x x 0.28-0.29 0 S sp. (No. 2497) Faviero Brazil x x ~ 95 cosy gemeatapa yore Balf. f. (Saupe, 4) ; BAPHIA NITID d. (Wiesner, 5, p. 936) |} Camwood Tropical Africa x x x ; BAUHINIA bcutara Pit tier Panama x pat x 0.22—0.23 | IIO-115 ETICULATA DC. (Héhnel, 1, p. 3 Africa? x x BELAtRsA MUCRONATA Griseb. Yamaquey Cuba x > 4 oe 0.13-0.14| 180-190 sprnosa A, Rich. x x x 0.14 175-180 Brya EBenus D Green Ebony West Indies x x x 0.15-0.16 | 160-165 CARSALPINIA CORIARIA Willd. Di vi-div i Curacao x ~ x|— 0.2I-0.23 | 110-120 : x —i xX x 0.18 136 a be rf India x x x 0.19-0.20 | 126-132 sg EBAN! ‘ o ferro Brazil x x x 0.25-0.26} 95-100 My eat St ei ALi Brazil-wood bs x x x ‘ 90 « MELANOCARPA Griseb. Guayacan Argentina x x —j|X} | 0.18-0.19 | 130-135 4 : s x x x 0.17-0.18 | 140-145 bis Sappan L. Sappan-wood Philippine Islands | X X| X x 0.26 905 SGOOM NIVIYHD JO AMNLOAALS :-daooay CAESALPINIA TINCTORIA Domb. (Héhnel, I, p. West Indies? x x : 0.33 — Fistuta L. (Héhnel, 1, p. 37) India? x x 0.17 Cast — RMUM AUSTRALE A. Cunn. & He lack Bean Australia x Mi | LX 0.31-0.33| 75-80 Abe epee PATINENSE Pittier Amarillo de Guaya- | Panama x 0.23-0.24| I05-IIO quil ‘ x x - ROBUSTUM Mart. Colombia x x x 0.23 ss sp. ? (ten specimens) Arariba Brazil x x x 0.21-0.25 | 100-200 e Guayacan Jobo South America x 0.25 100 -CERCIS CANADENSIS Ls Redbud Central United x States Fel eee set 0.16-0.17 | 150-160 ‘* SIL1guAstTRUM L. (Wiesner, 5, p x 930 Judas Tree Europe

ms in mm, fe Cg} 9 > ail Sie aS 2) 8/83| | gia) 8 3) 8] 2/2 Zl ale | e/a] 5] 3/a) 8]8) LEGUMINOSAE—Continued | PTEROCARPUS cars - Brazil x x x 0.26 05 SAN Red Sanders dia x x x ; 113 PTEROGYNE en “Tul oie specimens) | Ibiraro or Viraro Argentin x x x 0.25-0.28| 90-100 sp. ereipo enezuela x x x 0.26 04 ~SOPHORA CHRYSOPHYLLA See i waii x x x 0.16 155-160 SECUNDIFLORA Lag. Frijolita Southwest United States x x 0.16 160 SWARTZIA GAILLARDI Pittier Panama x x x 0.22—0.23 | IIO-II5 —— enth. : x x x 0.25 100—102 - TOMENTOSA DC, (Stone and Freeman, ‘ . 55) amara British Guiana x ? x TAMARINDUS INDICA Tamarind Hawaii x x x 0.31 80 TIPUANA Hoping Benth. Tipa Argentina x x x 0-0. 120-125 s 0. 897 - ws x x x 0.18—-0.19 | 130-135 WALLACEODENDRON CELEBICUM Koord. Banuyo Philippine Islands | X x x 0.33-0.36 O- 7 LYTHRACEAE (see page 260) ABERRIMUM Pohl? Brazilian Tulip Brazil x | X x 0.16 160 MALVACEAE GOSSYPIUM ARBOREUM L, Cotton Costa Rica x Xx x 0.20 85 896 SGOOM NIVLIYHD AO AMNLINULS :aqxooay GOSSYPIUM DRYNARIOIDES Seem Kokio Hawaii x x x Histscus aspirant oy Haesk. Philippine Islands | X x x 0.33-0.36| 70-°75 MACROPHYLLUS Rox Java x x * chine Blume (M. &J., 7, 1,2. 375) iy x x Hreiscus TILIaceus L. . Malubago Philippine Islands | X|— +S roe Xx 0.28-0.30| 85— 90 IDA DENSIFLORA Hook. & Loro blanco x x x 0.23 I ESPES PULNEA Corr. aL (aL Ba p. 389) India > eee x x 0.25 100 cape Carapa ee, Aub Andiroba South America x X| |x 0.50 50 ATA ttn OM. Sia Be ty p. 2 Java x x Satay sp. (four specimens) Cedro Tropical America | X |X| Xx 0.45-0.50} 50- 55 HICKRASSIA TABULARIS A. Juss. (Chuk- rasia tabularis A. Juss.) Chittagong Wood | India x x! Xx| xX 0.31-0.32| 78- 82 ENTANDROPHRAGMA CANDOLLEI Harms Sapeli Africa x xX| xXx 0.5 50 GUAREA TRICHILOIDES L. (Hodhnel, 1, p. 39) x x KAYA sp. African Mahogany } Africa x X| x\ X x ego 50 SwIETENIA MAHAGONI Jacq. Mahogany, Caoba | Tropical America | X ¥54 coral uaa [Pa 0.46-0.54| 46- 54 MORACEAE ‘ Ficus ALTissimA Blume India x x\" x 0.38-0.40| 62- 66 “© BENJAMINA L. Balete Philippine Islands | X Xx x x 0.39 64 MORINGACEAE gken seme co Gartn. (HGh- Griesholz BS crane x| X| Xx Skosonas ih (M.&J., 7, II, p. 514) | Java x x x ROSACEAE (see page 260) euwrony Smetana Aubl.? (Parinarium | Ficatin- or Kénigs- ; Héhnel, 1, p. 43 holz Guiana x x SGOOM NIVLYHD JO AYNALIANLS :deoory 69Z TABLE OF WOODS WITH TIER-LIKE STRUCTURE, SHOWING THE ELEMENTS STORIED, THE REGULARITY AND VISIBILITY OF THE TRANSVERSE LINES (‘‘RIPPLE MARKS”), AND MEASUREMENTS OF ; TIER-HEIGHT—Contin nue Elements Regularity Visibility Measurements storied | on lines s A \ a Scientific name Common name Country ol alee & a Be Ne Be : s] a oe - -} 3 BY | 5 ?| Tier-height | Number of g| 8/8) 21 a| £| | sta) e/g, 8) mmm | “See 4 v = val els inc f\atalsigizea aie ae ‘| al aie glale|E|S/5/a/$/2 RUTACEAE : || | CHLOROXYLON SWIETENIA DC. Satinwood India x x |x 0.31-0.32-| 78- 82 SAPINDACEAE | SAPINDUS SENEGALENSIS Poir. (Héhnel, 1, 43) Africa? x x) xX x 0.20 SIMARUBACE PICRAENA EXCELSA oe itive, I, p | 41; Wiesner, 5, p Jamaica Quassia | West Indies —| x ? x ‘PicRASMA JAVANICA Bliss (M.&J., 7, II, Java x x x 0.30 “SIMARUBA AMARA Aubl. Pao parahyba Brazil x x x 0.43 58— 60 GLauca DC, Bitterwood Florida x xi—| |x 0.50 5I-— 52 SURIANA MARITIMA L, ? Palo corra Curacao > x x 0.21 0 ST. iSite ogame GUAZUMA ULMIFOL Guasima Curacao x x x 0.30-0.31 | 80- 85 HERITIERA coonrtse ‘Ait Dungon late ‘| Philippine Islands x —| X x 0.31 80 i MINOR Lam. i India x para tea’ x 0.31-0.32| 77- 80 a ie f x x | x 0.27 92 ea x x 0.31-0.33| 74- 80 KLEINHOVIA Hospita L. OLE SGOOM NIVIYSD JO AMNALINALS :adaoosy MeEvocuia inpicA Gray (M.&J., 7, I, p : Java Mire x x PTEROCYMBIUM bewsioedsion R.Br. (M.& In7% LP ‘3 x“ x x 0.40 eS oe Daeiline Merr Philippine Islands x| XX x x 0.45-0.50| 50- 55 PTEROSPERMUM D IVERSIFOLIUM Blume Java x a tee x\— 0.50 feted Jungh. (M.& . TT 4s D. 4 . x xXl— x\— 0.50 PTEROS bine ecudlieccitne Lam. India x|— x 4 X| 0.30-0.31 | 80- 85 Cae 1, p. 42) (0.25,H.) Bayok es Islands |X x x 0.38 66 STERCULIA SOETIDA L. (M.&J., 7, p. 424) x| X ULIFOLIA Ro Dalibuda In ale x| X x * 0.36 70 TARRIETIA AKCYRODENDKON Benth. Crow’s-foot Elm Australia x x x 0.36 70 VANICA Blum umbayao Philippine Islands x x x 0.36-0.41 | 60- 70 us Maange Mia. (M.&J., 7, I, p. 44 Java x x x TARRIETIA SYLVATICA (Vidal) Merr. Dungon Philippine Islands x x x 0.32-0.33|} 83- 85 “ “ iri « “a x xf | fea eo ee 0.26—-0.28 90- 95 TILIACEAE | | BERRIA ae LA Roxb. Java Pd oor x x 0.25 INQUELOCULARIS T.&B. (M.& | JB 492, 496) oe x x| xX COLUMBIA SERRATIFOLIA DC, Analin Philippine Islands _—) X ie le LN 0.31-0.33] 75—- 80 DIPLODISCUS PANICULATUS Turcz. Baloba e = Mis x x 0.31-0.33 | 75— 80 GREWIA ee 4 Juss. Java x x x E A Vahl (M.&J., 7, I, pp. ‘éo. ee x x x 0.23 Grewia ROLFE! Merr. Danglin Ptipin Islands x x x 0.24-0.25 | 100-105 we TILIAEFOLIA Vahl | Ind x x x 0.23 IIo HELIOCARPUS ger eae sie Shs Amor seco Argentina x x x 0.33 os CULATUS Turcz. Pan x| X x x 0.38 65 PENTACE piety "casi. (M.&J., 7; Java x x x 0.40 ScHOUTENIA OVATA Kérth. (M.&J., 7, I, p. 522) 4 px Mi XI 0.22 SGOOM NIVLYSD AO FANLINALS :dkxOouy TZ TABLE OF WOODS WITH TIER-LIKE STRUCTURE, SHOWING THE ELEMENTS STORIED, THE REGULARITY AND VISIBILITY OF THE TRANSVERSE LI inued NES (‘‘RIPPLE MARKS’’), AND MEASUREMENTS OF TIER-HEIGHT—Contlinue Risen | moparity Visibility Measurements pis : C : a E & a Scientific name Common name ountry | 3 a2 3 a ‘4 3 3 ; ig : Number of ae ) T| 2] Sle Tier-height | “tiers per | 2 aa "a 4 s ie o & 1 Tam, inch a of S| > =| .8| = 3 3| £183) 8) Se) 89/2 8) 2) 2 <4(MiF jmlmis| Eales 71 eee TILIA AMERICAN Basswood Eastern United States x x|— x X| 0.42-0.45| 55— 60 ** CORDATA Mill. var. JAPONICA Japan x yd |< X| 0.48 52 es Seen ang Vent. Basswood Southeastern United States x xI— x X|0.40-0.41 | 60- 62 ‘* MANDSHURICA Rupr China x x MEP 0:45: 56 ‘* PUBESCENS Ait. Trecsed, 8, p. 76) | Basswood Eastern United States x x|— x | 0.41-0.43]} 58- 60 ULMACE HOLoPpTELEA wiser ps is (Rox) Planch ___ (Ulmus atta India x x x 0.22—0,23 | IIO-II5 ZYGOPH ae BULNESIA ARBOREA ba ) ipa ra Mane x ma x 0.10 50 RMIENTI Palo santo Arge x x x 0.10 240-250 Guatacum pasamchoye: Gray West Coast Mexico | X x x 0.00 260-270 NALE L. Lignumvitae West Indie x x x 0.10 250-260 Kg : i Florida x x x 0.09 260-270 LARREA CUNEIFOLIA Cav. (Solereder, 6, Argentina? Lannea D oa reel Cav. Javilla . xX\— x 0.09 270-280 Pore addy ROM ETRA Ruiz & Pav. (Gua oom hygrometricum Bail.; Hohnel,_ ‘. p. 40) Guayac Chile x x x 0.09-0.10 GLE SGOOM NIVLYS) AO AMALIAULS *TaOoay RECORD: STRUCTURE OF CERTAIN WOODS 273 SUMMARY Storied or tier-like arrangement of part or all of the elements is characteristic of many dicotyledonous woods representing a wide range of natural orders and families. Thestoried structure is found also in the secondary phloém The structure is exhibited on longitudinal sections, typically the tangential, as fine transverse striations (‘‘ripple marks’’). These are often visible without a lens. ‘‘Ripple marks” are constant enough in stems of considerable thickness to serve as a valuable diagnostic feature. In some woods pit areas on the fibers are in seriation. In some instances, also, the cells of the wood parenchyma strands are arranged in a secondary seriation, visible under the lens. A table is appended giving for each wood the various elements storied, the uniformity and distinctness of the markings, and the height of the tiers. LITERATURE CITED — Héhnel, Franz, Ritter von. Uber stockwerkartig aufgebaute Holz- korper. Ein Beitrag zur Holzanatomie. Sitzungsb. math.-naturw. Cl. Akad. Wissens. Wien 89': 30-47. 84. Ueber den etagenférmigen Aufbau einiger Holzkérper. Ber. -_ . 2. Deutsch. Bot. Ges. 2: 2-5. 188 3. Jaensch, Th. Zur Anatomie einiger Leguminosenhdlzer. Ber. ‘Deutsch. Bot. Ges. 2: 268-292. pl. 5. 4. 4. Saupe, A. Der anatomische Bau des Holzes der Leguminosen und sein systematischer Werth. Flora 70: 259-268, 275-282, 307— 922;°323-435- 1867. 5. Wiesner, Julius. Die Rohstoffe des Pflanzenreiches 2. Leipzig. .: 9904. 6. Solereder, Hans. Systematic anatomy of the Dictoyledons. Eng- edition. Oxford. 1908. re , J. W., & Janssonius, H. H. Mikrographie des Holzes der auf te vorkommenden Baumarten. Leiden. Band I, 1906, Band IT, 1908. . Record, Samuel J. Tier-like arrangement of the elements of certain woods. Science II. 35: 75-77- I912- . Stone, Herbert, & Freeman, W.G. The timbers of British Guiana. London. 1914. + ee \o Ws og teas | iis ie 45 The ancestry of maize—a reply to criticism PAUL WEATHERWAX In a recent article by H. J. Kempton (1), of the: Bureau of Plant Industry, exception is taken to some statements made by me ina paper appearing last September in the Bulletin of the Torrey Botanical Club (45: 309-342), on the evolution of maize. Inasmuch as some real errors are pointed out in this criticism, their correction is in order; but, as to certain other points upon which our data differ, I have nothing to add to my original paper. Our differences of opinion on matters of interpretation and evalu- ation of facts would probably not be diminished by further dis- cussion; and any criticism on ethical grounds of the spirit of Kempton’s paper and his imputations of the writer’s ignorance are left to the fair judgment of the reader. But Kempton’s man- ner of treating details and his intermingling of the irrelevant with the relevant have clouded the issue, and a brief presentation of its present status seems appropriate. It is said (p. 4) that I have misrepresented Collins (2) by con- fusing bracts with prophylla and by substituting ‘‘one-rowed”’ for “single-rowed” in a quotation. Both points are well taken, for no misrepresentation was intended. These errors had been discovered long before the appearance of Kempton’s paper, and steps had already been taken to correct them; this is a welcome opportunity to do so. _ The error in speaking of the husk of the maize ear as pro- phylla (pp. 314, 330) does involve an unintentional misrepre- sentation of Collins’s statement; but this point is beside the ques- tion, for the true prophylla of all kinds of maize, including pod corn, have buds in their axils, which is the significant point. Whether or not these buds develop far enough to be visible externally is a quantitative matter. The difference between “one”’ and ‘‘single”’ in the quotation (p. 321), describing the pistillate spike of teo- sinte, is appreciated, and it is unfortunate that the substitution was made. But the bearing of this point upon the evolution of 275 276 WEATHERWAX: THE ANCESTRY OF MAIZE the ear is unchanged, for the fact remains that Collins (p. 525, footnote) was using the structure of the spike of teosinte in an attempt to explain the appearance of ears with odd numbers of tows in some hybrids between maize and teosinte; and a “‘single- rowed”’ spike, such as that of teosinte, is of no avail in getting around the difficulty, for it has two rows of functional spikelets, not to mention the aborted ones. Kempton’s support (pp. 6-7) of the theory that the ear of maize originated by fasciation is based chiefly upon certain true- breeding races having bifurcated ears and a bifurcated central spike in the tassel. But our only evidence of the regressive nature of these mutants is the fasciation theory itself, and reasoning of this kind does not make much progress. The fact that ears hav- ing rows not in multiples of four could not have been formed by the union of spikes having four rows each is evaded by Kempton’s assumption (p. 7) that a row of pairs of spikelets or the pedicelled spikelets of two such rows were aborted. Abortion of parts isa common occurrence in maize, but our best evidence of this is in finding their rudiments; and, if abortion is thought to be respons- ible for the occurrence of ears with ten, fourteen, or eighteen rows, the burden of proof is with those who reject a more simple and more direct explanation of the situation. Such an explanation is afforded by the theory that the ear and the central spike of the tassel have developed from a branched inflorescence like that of some of the sorghums. The assumption here involved is that the pairs of spikelets are the remnants of reduced branches; and ma- terial illustrative of intermediate steps in this reduction is afforded by branch corn, which was unknown when Collins (p. 526) re- jected this theory. Advocates of the hypothesis that maize arose by hybridization find the necessity for their position in the fact that ordinary maize has so many primitive characteristics and seems to occupy an intermediate position between teosinte, which is more highly specialized, and pod corn, which seems to be more primitive than ordinary maize. A detailed morphological study shows that the differences in specialization here concerned are neither so great nor so significant as was supposed when the hypothesis was first advanced: but fur- ther, discussion of this point is waived for the present. WEATHERWAX: THE ANCESTRY OF MAIZE 20 The primitive characters of the maize plant might cause some difficulty in connection with any theory to the effect that maize descended from teosinte itself or from some plant much like teo- sinte; but probably no one at present gives serious consideration to any such theory. In descending directly from a primitive ancestor, the plant need not have made equal progress in all re- spects, and a combination of highly specialized characters with others relatively simple would be expected. Probably no one ap- preciates more fully than does the systematist the significance of this principle. The weakest spot in the hypothesis of the hybrid origin of Zea lies in the fact that pod corn has not been shown to be essentially different from ordinary corn except in the possession of enlarged bracts and of functional pistils in the tassel. In Collins’s argu- ment, pod corn has no exact identity. Sometimes it has ears, and sometimes it is earless. He seems to have selected from all the characteristics possessed by this ill-defined variety those tend- ing to uphold his hypothesis. Kempton admits as much (p. 7) when he attributes the presence of buds in the axils of the leaves of pod corn to the recent origin of this variety from ordinary corn. The problem for which they offer no solution involves a means of distinguishing the characters of the orthodox podded type—one of the hypothetical ancestors of Zea, according to their view— from other characters received from ordinary maize by this recent mutant. It is true that the hybrid-origin hypothesis itself will distinguish them, but, when we apply it, we find ourselves dealing in circular logic. It is generally admitted that the podded types now in existence are recent mutations rather than original forms; but our criterion for the regressive nature of some of their charac- teristics is in the character of the grasses held to be closely related to maize. The idea of the hybrid origin of maize has had the misfortune to be based upon a hypothesis not substantiated by subsequently discovered facts. It has never been very enthusaistically accepted by botanists in general, because it seems unreasonable and taxes the imagination unnecessarily. It has had its good effect in pro- moting further investigation, but it has run its course and has been - found wanting." 278 WEATHERWAX: THE ANCESTRY OF MAIZE On the other hand, comparative morphology, which has been one of the most reliable and productive agents in establishing lasting theories of evolution, points out clear-cut, logical, reason- able evidence of the direct origin of Zea, codrdinately with Eu- chlaena and Tripsacum, from an ancestor long ago extinct. There has been pointed out no accepted fact whose rational interpreta- tion discloses any inconsistency in the theory. The three genera are different simply because they have lost different organs that were possessed by their progenitor and have specialized others to different degrees. INDIANA UNIVERSITY, BLOOMINGTON, INDIANA LITERATURE CITED 1. Kempton, J. H. The ancestry of maize. Jour. Wash. Acad. Sci. Oi 3-81 Toss. 2. Collins, G. N. The origin of maize. Jour. Wash. Acad. Sci. 2: 520-530. 1912. INDEX TO AMERICAN BOTANICAL LITERATURE 1911-1919 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 p ages that relate exclusively to forestry, agriculture, wetniy a rarreneniar products of vegetable origin, or laboratory methods are not included, an no attempt is made to index the literature of bacteriology. An occasional naan 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 edi to errors or omissions, their kindness will be a iated. This Index is reprinted monthly on cards, and saleuets in this form to subscribers at the rate of one cent for each card, Selections of cards are not permitied ; each subscriber must take 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 iestisa Club. Allen, E. R. Some conditions affecting the growth and activities of Azotobacter chroococcum. Ann. Missouri Bot. Gard. 6:1-44. pl. 1 J. te -¥ 4910. Allen, E. R., & Davisson, B. S. An all-glass nitrogen apparatus. Ann. Missouri Bot. Gard. 6: 45-48. pl. 2. F 1919. Alway, F. J., McDole, G. R., & Trumbull, R. S. Relation of mini- mum moisture content of subsoil of prairies to hygroscopic coeff- cient. Bot. Gaz. 67: 185-207. 18 Mr 1919. Apolinar Maria, Hno. El eucalipto (Eucalyptus globosus Lab.) Bol. Soc. Cien. Nat. La Salle 6:68-74. Je 1917; 85-87. Jl 1917; 107- 112. Au. 1917: 126-128. S 1917; 139-142. O 1917; 150-152. N 1917; 6: 13-20. Ap 1918. Armell, H. W. Die Moose der Vega-Expedition. Ark. Bot. 115°: I-1IT. 33918, Arthur, J. C., & Bisby, G. R. An annotated translation of the part of Schweinitz’s two papers giving the rusts of North America. | Proc. Am. Philos. Soc. 57: 173-292. 1918. Atkinson, G. F. Relationships within the Rhodosporeae. Bot. Gaz. 67: 266, 267. 18 Mr 1919. 279 a 280 INDEX TO AMERICAN BOTANICAL LITERATURE Bailey, V. General features governing life in Glacier National Park, [In Bailey, V. & Bailey, F. Wild Animals of Glacier National Park] 15-24. pl.1+/f.z. Washington. 1918. Bancroft, R. L. Experiments to determine whether one form of © phosphorus in a complete nutrient plant solution is more available for the growth of buckwheat seedlings than are other forms. New Jersey Agr. Exp. Sta. Ann. Rep. 38: 423, 424. 1917. Britton, E. G. Plants in ornament. Bull. Met. Mus. Art 14: 51, 52. 1919. [Illust.] Britton, N. L. Dr. Henry Allen Gleason, appointed first assistant. Jour. N. Y. Bot. Gard. 20: 39, 40. F 1919. Brooks, C., Cooley, J. S., & Fisher, D. F. Apple-scald. Jour. Agr. Research 16: 195-217. f. I-II. 24 F 1919 Brotherus, V. F. Contributions a la flore bryologique de 1’Argentine. Ark. Bot. 156: 1-15. 1918. hirteen new species in various genera are described. Brotherus, V. F.. Moseniella, un nouveau genre des mousses ~~ Brésil. Ark. Bot. 157: 1-3. pl. r. 1918. Buscalioni, L., & Muscatello, G. Studio anatomo-biologico sul Gen. “Saurauia’”’ Willd. con speciale riguardo alle specie americane. Malpighia 28: 49-81. 1917; 331-370. pl. 5-10. 1918. Buscalioni, L., & Muscatello, G. Studio monografico sulle specie americane del gen. ‘“Saurauia’” Willd. Malpighia 28: 1-48 pl. I, 2. 1917; 314-330. 1918. Bush, B. F. The Missouri muhlenbergias. Am. Mid. Nat. 6: 17-20. Mr 1919. Candolle, C. de. Begoniaceae Centrali-Americanae et Ecuadorenses. Smithsonian Misc. Coll. 69%: 1-10. 9 Ap 1919. Seventeen new species are described. Chase, A. Some causes of confusion in plant names. 17: 159-162. F 1919. Chavarria, A. P. Apuntes para el estudio de la flora nacional. Bot. Soc. Cien. Nat. Inst. La Salle 6: 135-138. O 1917. Cockerell, T. D. A. Hybrid perennial sunflowers. Bot. Gaz. 67: 264-266. 18 Mr igig. Collins, J. L. Chimeras in corn hybrids. f. 1-6. 8 Mriog19. [Illust.] Condit, I. J. & Stevens, H. J. ‘‘Die-back”’ of the fig in California. Mon. Bull. State Comm. Hort. Calif. 8: 61-63. f. 34, 35. F 1919. Jour. Forestry Jour. Heredity 10: 3-10. INDEX TO AMERICAN BOTANICAL LITERATURE 281 Cook, M. T. Potato diseases in New Jersey. New Jersey Agr. Exp. Sta. Circ. 105: 1-38. f. 1-19. Ja 1919. Cook, M. T. Report of the department of plant pathology. New Jersey Agr. Exp. Sta. Ann. Rep. 38: 523-563. pl. 1-6. Cook, M. T., & Helyar, J. P. Diseases of grains and forage crops. New Jersey Agr. Exp. Sta. Circ. 102: 1-16. f. 1-4. 7 N 1918. Cordero, M. Prolegomenos para e! estudio analitico del agave. Bol. Dir. Est. Biol. Mexico 2: 235-240. Jl 1917. [Illust.] Deane, W. Amsinckiain New England. Rhodora 21: 38-40. 25 Mr 1919. Detmers, F. Two new varieties of Acer rubrum L. Ohio Jour. Sci. 19: 235-237. pl. 12, 13. F 1919. Dodge, C. W. Tyrosin in the fungi; chemistry and methods of study- ing the tyrosinase reaction. Ann. Missouri Bot. Gard. 6: 71-92. F-¥e F919: Dorety, Sr. H. A. Embryo and seedling of Dioon spinulosum. Bot. Gaz. 67: 251-257. pl. ro-rz. 18 Mr 1919. Duggar, B. M. & Dedge,C. W. The use of the colorimeter in the indi- cator method of H ion determination with biological fluids. Ann. Missouri Bot. Gard. 6: 61-70. f. z. F. 1919. Fairchild, D. Testing new foods. Jour. Heredity 10: 17-28. f. o-13- 8 Mr 1919. Fawcett, W., & Rendle, A. B. Notes on Jamaica plants. Jour. Bot. 57: 65-68. Phyllanthus minor, P. inaequaliflorus and P. Coxianus, spp. nov., are described. Fernald, M. L. Carex flava, var. gaspensis in Vermont. Rhodora 21:40. 25 Mr 1919. Fernald, M.L. Helianthemum Bicknelliiand H. propinquum. Rhodora 217.46, a7~: 25.Mr 1919. Fromme, F. D., & Murray, T. J. Angular-leafspot of tobacco an un- described bacterial disease. Jour. Agr. Research 16: 219-228. pt. 25-27. 24 F 1919: Bacterium angulatum sp. nov. Gager, C. S. Horticulture as a profession. Science II. 49: 293-300. 28 Mr 1919. Garrett, A. O. Spring flora of the Wasatch region. i-xii + 1- 106. Salt Lake City, 1911; i-xii + 1-139. Lancaster. 1912; i-xii + 1-144. Lancaster. 1917. First, second and third editions. 282 INDEX TO AMERICAN BOTANICAL LITERATURE Goodspeed, T. H. Notes on the germination of tobacco seed—III. Note on the relation of light and darkness to germination. Univ. Calif. Publ. Bot. 5: 451-455. 3 Ap 1919. [Grosvenor G.] American berries of hill, dale, and wayside. Nat. eog. Mag. 35: 168-184. pl. 1-8. F 1919. Illustrated by Miss M. Eaton. Hallier, H. Ueber Aublet’s Gattungen unsicherer oder unbekannter Stellung und iiber pflanzengeschichtliche Beziehungen zwischen Amerika und Afrika. Mededeel. Rijks Herb. 35: 3-33. 29 Ja 1918. : Hallier, H. Ueber Patrick Browne’s Gattungen zweifelhafter Stellung. Mededeel. Rijks Herb. 36: 1-6. 8 F 1918. Halsted, B. D. Possible correlations concerning position of seeds in the pod. Bot. Gaz. 67: 243-250. 18 Mr 19109. Halsted, B.D. Report of the department of botany. New Jersey Agr. Exp. Sta. Ann. Rep. 38: 371-408. 1917. Hamlin, A.D. F. Plant formsin decorative art. Bull. Met. Mus. Art 14: 49, 50. 1919. [Illust.] Harper, R. M. A new method of mapping complex geographical features, illustrated by some maps of Georgia. School Sci. & Math. 18: 699-708. N 1918. four maps of Georgia show ‘‘geographical” divisions, and the prevailing soil-type, commonest trees, and leading crops in each of these divisions of the state. Henrard, J. T., & Thellung, A. Lepidium flavum Torrey var. apterum nob. Mededea! Rijks Herb. 34: 1, 2. 29 Ja 1918. [Illust.] Herzog, T. Die von Dr. Th. Herzog auf seiner zweiten Reise durch Bolivien in den Jahren 1910 und 1911 gesammelten Pflanzen. Mededeel. Rijks Herb. 29: 1-94. 29 D 1916; 33: I-19. 28 Ja 1918. Includes the new genera Corollonema, Schwenkio psis and Saccanthus and 83 new species in various genera Hitchcock, A. S. Report of the committee on generic types of the Botanical Society of America. Science II. 49: 333-336. 4 AP I9I9. Hoagland, D. R. Note on the technique of solution culture experi- ments with plants. Science II. 49: 360-362. 11 Ap 1919. Horne, W. T. Oak-fungus disease, oak-root fungus disease, fungus root-rot, toadstool root-rot or mushroom root-rot. Mon. Bull. State Comm. Hort. Calif. 8: 64-68. f. 36-39. F 1919. Hotson, J. W. Sphagnum from bog to bandage. Publ. Puget Sound Biol. Sta. 2: 211-247. pl. 31-48. 5 Mr 1919. INDEX TO AMERICAN BOTANICAL LITERATURE 283 Hurd, A. M. The relation between the osmotic pressure of Nereo- cystis and the salinity of the water. Publ. Puget Sound Biol. Sta. 2: 183-193. 5 Mr 19109. Irwin, M. Comparative studies on respiration—VI. Increased pro- duction of carbon dioxide accompanied by decrease of acidity. Jour. Gen. Physiol. 1: 399-403. f. 1, 2. 20 Mr 1919. Itano, A., & Neill, J. Influence of temperature and hydrogen ion concentration upon the spore cycle of Bacillus subtilis. Jour. Gen. Physiol. 1: 421 —428. f. 1, 2. 20 Mr 1919. Jagger, I. C. Control of vegetable diseases. Cornell Ext. Bull. 19: 2915-2936. f. 91-113. Je 1917. Lyman, G. R. The unification of American botany. Science II. 49: 339-345. 11 Ap 1919. Lynge, B. Ueber einige Regnellschen Parmelien aus Matteo-Grosso, Brasilien. Ark. Bot. 15): 1-4. 1918. Parmelia fatiscens and P. coccinea, spp. nov., are described. McClelland, T. B. Influence of foreign pollen on the development of vanilla fruits. Jour. Agr. Research 16: 245-252. pl. 31-35. 3 Mr 1919. MacDougal, D. T., & Spoehr, H. A. The origination of xerophytism. Plant World 21: 245-249. O 1918. McDougall, W. B. Development of Stropharia epimyces. Bot. Gaz. 67: 258-263. f. I-10. 18 Mr 1919. Martin, W. H. A comparative study of salt requirements in solution cultures for the growth of buckwheat to the flowering stage and to maturity. New Jersey Agr. Exp. Sta. Ann. Rep. 38: 419-422. 1917. Matz, J. Algunas observaciones respecto a la Sarna (scab) del citro en Puerto Rico. Rev. Agr. Puerto Rico 2: 40, 41. F 1919. Matz, J. Enfermedad de la rdiz de la cafia de Azicar. Rev. Agr. Puerto Rico 2: 38, 39. F 1919. Mousley, H. The orchids of Hatley, Stanstead County, Quebec. Ottowa Nat. 32: 144-147. F 1919. Murrill, W. A. Trees and children. Jour. N. Y. Bot. Gard. 20: 61-63. Mr i919. Neller, J. R. The influence of the roots of growing plants upon the activity of soil microérganisms as indicated by the production of carbon dioxide from the soil. New Jersey Agr. Exp. Sta. Ann. Rep. 38: 414, 415. 1917. 284 INDEX TO AMERICAN BOTANICAL LITERATURE Ness, H. Field experiments with crown gall, 1913-1917. Texas Agr. Exp. Sta. Bull. 211: 1-21. f. 1-7. I Olaya, R. M. La ciencia y el mana de los hebreos. Bol. Soc. Cien. Nat. Inst. La Salle 4: 42-46. Ap 1916. Osterhout, W. J. V. A comparison of permeability in plant and animal cells. Jour. Gen. Physiol. 1: 409-413. 20 Mr 1919. Osterhout, W. J. V. Decrease of permeability and antagonistic effects caused by bile salts. Jour. Gen. Physiol. 1: 405-408. f. r, 2. 20 Mr 1919. Potts, A. T. The fig in Texas. Texas Agr. Exp. Sta. Bull. 208: I=qt. f. 1-13... Ja 1or7. Rigg, G. B. Early stage in bog succession. Publ. Puget Sound Biol. Sta. 2: 195-210. pl. 29, 30. 5 Mr 1919. Rosen, H. H. A preliminary note on a bacterial disease of fox tail. Science II. 49: 291. 21 Mr 1919. Rosendahl, H.V. Filices novae. Ark. Bot. 14!8; 1-5. gy I-3. 1916. Adiantum madagascariense, Asplenium Afzelii, Diplazium latisectum and Odon- tosoria, spp. nov., are described. Rydberg, P. A. A genus of plants intermediate between Petalostemon and eee Sodas N. Y. Bot. Gard. 20: 64-66. Mr 1919. Thornbera gen. nov. Sargent, C. S. ea on North American trees. IV. Bot. Gaz. 67: - 208-242. 18 Mrigrg. Populus Palmeri, P. texana, Betula Eastwoodae, spp. nov., and many new varieties are described. Schneider, C. Arbores fruticesque chinenses novi. IV. Bot. Gaz. 64: 137-148. pl. 15. 15 Au 1917. Describes 8 new species in Salix and 1 in Alnus. Shive, J. W. Comparison of salt requirements for young and for mature buckwheat plants grown in sand cultures. New Jersey Agr. Exp. Sta. Ann. Rep. 38: 409-413. f. z. 1917. Shufeldt, R. W. Various parasitic plants; with an owl story. Am. Forestry 25: 937-941. f. 1-9. Mr 1919. Small, J. K. Ferns of Royal Palm Hammock. i-vii + 1-39. i. 1. New York. 1918. [Illust.]_ Small, J. K. Narrative of a cruise to Lake Okeechobee. Am. Mus. Jour. 18: 685-700. D 1918. [TIllust.] Small, J. K. The prickly-pears of Florida. Jour. N. Y. Bot. Gard. 20; 21-39. pl. 224-226. F 1919. Includes Opuntia lata Small, O ammophila Small, O. keyensis Britton, and O. sebrina Small. Vol. 46 No. 8 BULLETIN OF THE TORREY BOTANICAL CLUB AUGUST, I919 A new Matonidium from Colorado, with remarks on the distribution of the Matoniaceae EDWARD W. BERRY (WITH PLATES I2 AND I3 AND TWO TEXT FIGURES) In 1916 Professor Cockerell, of the University of Colorado, published a brief note* based upon plant material collected from the supposed McEIlmo formation of southwestern Colorado. The subsequent discovery that this supposed Lower Cretaceous flora was stratigraphically above a black shale horizon carrying a considerable dicotyledonous flora led to the sending of all of the material to the present writer for a more critical study than Professor Cockerell was able to devote to it. The discussion of the question of the stratigraphic relations and age of these two floras is reserved for a subsequent communi- cation, the present paper being devoted to the interesting questions of habit and distribution derived from a study of the abundant relics of Matonidium preserved in the later of the two floras above mentioned. The Colorado remains, which Professor Cockerell identified as Matonidium Althausii, or Matonidium Goepperti as some paleo- botanists prefer to call the species on the ground that the older and correct name is less familiar, are superficially very much like that well-known and rather variable species. Botanists who * A Lower Cretaceous flora in Colorado. Jour. Wash. Acad. Sci. 6: 109-112. pl. I, 2. 1916, [The BuLLETIN for July (46: 235-284) was issued July 31, 1919.] 285. - 286 BERRY: A NEW MATONIDIUM FROM COLORADO depreciate specific differentiation and who, like Professor Seward, are more interested in the former distribution of generic types, would not hesitate to refer the Colorado form to Matonidium Althausit. Such questions are always difficult, and tastes and temperaments differ with individual workers. One point of view may serve one purpose and the other another. All that need be insisted upon in the present connection is that the framing of loosely defined specific limits as opposed to finely discriminated ones effectually obscures whatever value the fossils may have for the elucidation of geologic chronology or problems of the former time and avenues of migration, and is a method that has gradually become almost obsolete in the allied field of paleozodlogy. It would seem a priori incredible that a single botanical species should range through the Jurassic and Lower Cretaceous and extend over at least two continents in opposite hemispheres; nevertheless it is impossible to differentiate the British Jurassic from the Wealden forms or these from the still younger Russian occurrences, and the specimens referred by Ward to Matonidium Althausii from the Albian Fuson formation of the Black Hills* in this country cannot be distinguished from the earlier European occurrences of that species. Recognizing fully the individual variability in Matonidium Althausit and the variation in the form and size of the pinnules from different parts of a single frond, the Colorado Matonidium nevertheless presents a number of apparently constant peculiari- ties that serve to mark it as a slightly but consistently different type. Among these distinctive features might be mentioned the stouter stipe and rachises, the more numerous pinnae, which appear to have been at least twice as numerous as in Matonidium Althau- sit, judging from the frond bases of the latter as figured by Ettings- hausen, Schenk and Seward. The pinnae are more slender in the Colorado form and the pinnules are shorter and wider with fewer and more nearly circular or isodiametric sori. For example, in the form of Matinidium Althausii described from the Black Hills the falcate slender pinnules are twice as long as in the Colo- rado material, and each bears about twelve pairs of sori. The Portuguese specimens as figured by Heer} show eighteen pairs of * Ann. Rep. U. S. Geol. Surv. 197: 653. pI. 160, f. 5-8. 1899. t Sece. Trab. Geol. Port. 16. pl. 15, f. 1-6. 1881. BERRY: A NEW MATONIDIUM FROM COLORADO 287 sori to each pinnule, Schenk’s admirable figures* show eight or nine pairs, and they are very numerous in Trautschold’s figures of the Russian material.t The most elongate and largest pinnules of the Colorado species bear but five or six sori on the distal half of the lamina and but six or seven on the proximal half, while the most abundant remains are those with short and broad pinnules bearing three sori on their distal halves and four sori on their proximal halves. I regard these features and others to be mentioned as of specific value and propose that the Colorado specimens be discriminated as Matonidium americanum sp. nov. Matonidium Althausiit Cockerell, Jour. Wash. Acad. Sci. 6: 111. f.2. 1916. [Not (Dunker) Ward.] Cycadospadix ? sp. Cockerell, Idem. 110. f. 1. This species also includes formswhich Cockerell suggested might represent Todites, and Weichselia or Cladophlebis. The equiset- aceous stems mentioned as probably representing Eguisetum Burchardti (Dunker) Brongniart are probably fragments of the large stipes of Matonidium. The striations are somewhat irregu- lar and do not alternate at the nodes, there are no traces of nodal sheaths and the apparent jointing appears to be due to shrink- age cracks of a collapsed and consequently furrowed stipe, since a similar state of preservation is seen in the objects that were referred to Cycadospadix, which latter are clearly the apical portions of Matonidium stipes and only most superficially suggestive of Cycadospadix, and strictly comparable with similar remains of Matonidium Alihausii as figured by Schenk,t Ettingshausen§ and Seward. || Matonidium americanum may be characterized as follows: Digitate fronds with the same habit as Matonidium Althausu or the existing Matonia pectinata, with long stout stipes about I cm. in diameter as preserved in a partially flattened condition. Stipe surface somewhat regularly furrowed, but whether this was * Palaeont. 19: 220. pl. 27, f. 5; pl. 28, f. 1, 2; pl. 30, f. 3; pl. 42, f. rz. 1871. + Nouv. Mém. Sci. Nat. Moscou 13: 28. pl. ro, f. 3. 1870. t Palaeont. 19: pl. g2 f. 7. 1871. § Abh. k. k. Geol. Reichs. 13: pl. 5. f. 4, 6. 1854. || Jurassic Fl. 1: pl. rr, f. 1. 288 Berry: A NEW MATONIDIUM FROM COLORADO an original feature or is due to collapse during fossilization is undeterminable. Distad the stipe expands into a flabellate recurved or reflexed ‘‘collar’’ which shortly divides to form the axes of the pinnae, whose attitude in life was transverse or re- flexed with respect to that of the vertical stipe. These reflexed collars’’ are especially common in the rocks and are the objects which Cockerell referred to Cycadospadix. They are well shown in PLATE 12, FIGS. 9-I2. The width of the “collar” before it split up into pinnae varies from specimen to specimen and is not always clear, although it is narrowest medianly. No traces of pinnules have been found in actual connection with these stipes although they are in close association. The divisions of the ‘‘collars,’’ however, do not terminate as they would if the fossils were of the nature of Cyca- dospadix, and the breaks at the ends are obviously mechanical and not natural. The evidence of their reflexed form is also clear, and the size of the segments, their condition of preser- vation and furrowing are in exact agreement with the speci- mens of proximal pinnae axes. The number of pinnae was large but somewhat variable. The ‘‘collar’” shown in Fic. 9. indi- cates four lateral pinnae at each end of the collar and thirty- one additional intercalated pinnae, making thirty-nine in all. Fic. 10 shows clear traces of at least twenty-five pinnae, and Fic. 12, which is the counterpart of the apex of the specimen shown in Fic. 11, shows the bases of thirty-six pinnae. The pinnae were therefore much more numerous than the published figures indicate for Matonidium Althausii and also somewhat more numerous than the normal number in the modern Matonia pecti- naia. The exact length of the pinnae is unknown since all are preserved in a fragmentary condition, but judging by the frag- ments of proximal, medial and distal parts preserved, and the size and degree of tapering of the rachises, they could not well have been shorter than 20-25 cm., or about the same length as in the admirable specimen of Matonidium Alihausii figured by Schenk* from the German Wealden. Their width, however, was much less than in that species, the largest certainly identified (PLATE 12, FIG. 8) being but 11 mm. across, while the more abund- * Palaeont. 19: pl. 28, f. 1. 1871. | BERRY: A NEW MATONIDIUM FROM COLORADO 289 ant remains (PLATE 12, FIG. 2) areabout 8 mm. inwidth. Several sterile fragments which I have referred to this species on the basis of general form and association are somewhat larger(PLATE 12, FIG. 7) being 15 mm. wide. These show a narrower rachis with slightly less obtuse pinnules which are greatly crowded, and may represent fragments of a contemporaneous species of Cladophlebis. Fertile fragments of pinnae present a very different appearance when the opposite faces are viewed. From below the rachis appears very stout and the laminae of the pinnules is entirely hidden by the large sori except for a broad band in the position of the midvein. Viewed from above the rachis is much narrower, the pinnules evidently having been inserted near the upper margin of the rachis as shown in the section (PLATE 13, FIG. 4). The pinnules are short and broad, more or less falcate, obtusely rounded at their tips. All have revolute margins and they are opposite or sub-opposite in position. Their substance is thick and the venation cannot be made out in any of the preserved material. Distad along the rachis the pinnules gradually become shorter and somewhat broader relatively (PLATE 12, FIG. 6), eventually becoming very small and coalescent basally (PLATE 12, FIG. 3, 4), their revolute margins giving them a pointed appearance. Proxi- mad along the rachis the pinnules gradate through short and broad forms to short wide scallops as shown in PLATE 12, FIG. 5. The sori are in two rows separated by a broad space in the region of the midrib. They diminish in size toward the pinnule tips and are prevailingly circular except where mutual crowding causes the lateral margins to be somewhat flattened. They stand out from the surface of the pinnules as prominent umbos and a ‘pronounced umbilicus on the indusium marks the position of the central soral axis. The indusia are peltate and are intact in all of the Colorado material so that the number of sporangia can not be made out. In Matonidium Althausii the sori are said to have been more numerous than in the modern Matonia pectinata, sections of a sorus of the latter species being introduced for com- parison on PLATE 13, FIGS. 1, 2. _In the bulk of the material each pinnule shows three or four upper and four or five lower sori (PLATE 13, FIG. 3). These were the only forms in the collection which Professor Cockerell had labelled Matonidium Althausu. In the 290 Berry: A NEW MATONIDIUM FROM COLORADO largest fruiting material there are six or seven sori on the distal halves of the pinnules and seven or eight on their proximal halves, (PLATE 13, FIG. 6), a less number than in Matonidium Althausu, as already remarked. The sori also differ from those of the latter in being isodiametric instead of transversely elongate and more or less rectangular. The material, which was collected by J. T. Duce, comes from the divide between Cutthroat Gulch and Hovenweep Canyon west of Dolores and is contained in a light somewhat ferruginous sand- stone containing little quartz and a great deal of feldspar, and apparently corresponds to what Cross correlated with the Dakota _ sandstone in the San Juan region of Colorado. The family Matoniaceae is one of exceptional botanical in- terest. Its sole existing genus, Matonia R. Brown (1830), con- tains but two known species which differ remarkably in vegeta- tive habit and are both confined to the uplands of the Malay Peninsula and the island of Borneo. The older and only species known up to 1888 was commonly associated with the family Cyatheaceae, although its intermediate character had usually _ resulted in its being set apart as a special tribe. It has also been associated with the Gleicheniaceae and the Polypodiaceae. Christ (1897) and most subsequent workers have regarded Matonia as the sole existing type of a distinct family, and an admirable account of the genus and of some of its fossil relatives was given by Seward in 1899.* The extinct genus Matonidium, to which the Colorado plant is referred, was a characteristic type of the Jurassic .and especially of the Lower Cretaceous. Although known as early as 1843 its botanical affinity was first recognized by Schenk in 1871 and it has subsequently been found to have been widespread in the European region and more sparingly represented in the western United States. Most of the occurrences have been referred to the single species Matonidium Althausii, although Krassert has described a well characterized species from the Upper Cre- - taceous (Cenomanian) rocks of Moravia, which appears to have also been present in rocks of the same age in Bohemia. * Phil. Trans. Roy. Soc. Lond. 191B: 171-209. pl. 17~20. 1890. { Beitr. Palaon. Oesterreich-Ungarns 10: 119. pl. rz, f. 1; pl. 12, f. 1, 2; pl. 17, f. 10. 1896 ee ; Berry: A NEW MATONIDIUM FROM COLORADO 291 These occurrences are assembled upon the accompanying sketch map (TEXT FiG. 1) and are shown within the enclosed areas in Europe and North America. The area of distribution of the two existing species of Matonia is shown on the map in solid black. A second rather well-known extinct genus belonging to this family is the genus Laccopteris, which was founded by Presl in 1838. Laccopteris was especially characteristic of the late Triassic and hence was considerably older in its inception than Matonidium. Its relationship with Matonia was first clearly defined by Zeiller* as 2 ay ] Rha he Wit eo. | | es Royyd’ | moe 7 ok yy & : fe & a FE a li as ch Fic. 1. Sketch map showing the area of distribution of the two existing species of Matonia (solid black), the fossil occurrences of Matonidium (black disks within the enclosed areas), and the fossil occurrences of Laccopteris where they do not coincide with occurrences of Matonidium (L). in 1885. Reference to the accompanying map (TEXT FIG. 1), where the occurrences of Laccopteris are indicated by L, shows that it greatly extends the Mesozoic range of the family and gives the Matoniaceae representation in the far north (Greenland and Spitzbergen), in the Asiatic region and in Australia. Other and less well known fossil genera which have been referred to the Matoniaceae include Phlebopteris Brongniart, Microdictyon Sa- porta and Knowltonella Berry. * Bull. Soc. Bot. France 32: 22. 1885. 292 Berry: A NEW MATONIDIUM FROM COLORADO VETUAAALPAAAATILG2-723-74-72244-2-224 YUE! MUTA SS SSSSSSSSSSSSSESSSS Meumnnnneennre eS SSS ; M ¥ 4 v4 ¥ vy VY 2 Ne \y ij Fe Y Y Ve i Wr Y NA V7 YF yer NY Nv, Vea Va iE ws S V4 VE ye NY VA eb Ni 7 We Ni Ny 4 ne NZ 7 Ve % if Ny VF, VS “ NY Ne, \ YE we WA WG yi U4 We eS NZ SZ (Bs N24 NZ S WES S2NZ NE i NG? VG PS SBNE VE és SON ANG Pr Z WE yess UBNB Nr phe EES YANG WZ \ les NING NE \ We NZ OZ YES NN IAN prs SAE V7, VA pees SABNEVE \(\ ASS NAA ee AAA Ss NBO YAY UU AY &. y ul! 4} : VA Ui ft; 4, th NN Fic. 2. Restoration of Matonidium americanum Berry, two fifths natural size. Berry: A NEW MATONIDIUM FROM COLORADO 293 The accompanying restoration of Matonidium americanum, shown in TEXT FIG. 2, is based upon a considerable amount of ma- terial which, although fragmentary, supplies data for all parts of the frond. Its abundance in the small collection from this one locality would seem to indicate that the species was gregarious, sending up its stout stipes from a creeping rhizome as does the modern Matonia pectinata. The reduced figure was made from a life size drawing made by plotting the various fragments of the different parts to scale, and the arrangement of the pinnae is based upon their disposition in Matonia pectinata and the evidence derived from the numerous fossil frond bases (‘‘collars’’) preserved. Whether the stipe in the fossil species was as elongated as in the modern Matonia pectinata is not known. At the apex the stipe of Matonidium americanum was less distinctly dichotomous with less scirpoid branches than in Matonia pectinata, one side of the stipe becoming concave and the whole forming a reflexed fan- like base from which the rachises of the numerous pinnae proceded. In life these were not upright as in the restoration but were turned back ninety degrees or more giving the fronds an almost peltate appearance. 294 x Loma CPWAKRY DH Fic. 1. sporangia. 1G. 2. annuli. Fic. 3. sori, X 10. Fic. 4. Fic. 5. xX 10. Fic. 6. below, X 1 Berry: A NEW MATONIDIUM FROM COLORADO Explanation of plates 12 and 13 PLATE 12 MATONIDIUM AMERICANUM Berry Under side of fertile pinna showing stout furrowed rachis. Under side of fertile pinna showing normal size and form. Upper side of apical ghia of pinna. Portion of same, Basal part of pinna a below. Upper part of pinna from above. Maximum size of rare occurrence, from above. Fragment from below showing maximum development of sori. Apex of stipe showing indications of 39 pinnae. Apex of stipe with 25 pinnae Apex of stipe of another specimen. Counterpart of apex of specimen shown in 11 with indications of 36 PLATE 13 Fics. 1,2. MATONIA PECTINATA R. Br. Lamina and sorus in vertical section showing indusium (black) and. Sorus in transverse section showing central pillar and sporangia with Fics. 3-6. MAaTONIDIUM AMERICANUM Berry Two pinnules of normal form from below to show form and position of Same in longitudinal vertical section, x 10. A ative ‘ le int ti showi J t—2 Two pinnules of the large form with maximum number of sori, from Phytogeographical notes on the Rocky Mountain region Vill. Distribution of the Montane plants P. A. RYDBERG The Montane Zone or Pine Belt of the Rocky Mountain region includes approximately the areas between the altitudes of 2,500 and 3,000 m. in southern Colorado, between 1,800 and 2,500 m. in southern Montana, and between 1,200 and 1,800 m. in Alberta, where the Canadian Pacific Railroad crosses the mountains. A general description of the Zone has been given in a previous ar- ticle.* The Subalpine Zone of the Rockies comes in contact with the Hudsonian or Eastern Subarctic Zone along the foothills of the Rockies from the headwaters of the Peace River northward, and here the species of the East and of the Rockies more or less in- termingle. But the relation between the Montane Zone of the Rockies and the Canadian or Eastern Boreal Zone is very different, - for the latter, in its typical development, reaches its western limits in the region of the Lake of the Woods and Lake Winnipeg. The most representative species of the Canadian Zone, Strobus Strobus and Pinus resinosa, reach their northwestern limits in this region and it is only in the transition belt between the Canadian and Hudsonian Zones, which extends along the height of land between the Saskatchewan and Athabasca Rivers, that species of the Canadian Zone range farther west. This transition belt is characterized by mixed woods of Pinus Banksiana, belonging to the Canadian, and of spruces and larches belonging to the Hud- sonian Zone. North of the upper Athabasca River and Beaver River practically the whole country is Hudsonian, while south of the northern branch of the Saskatchewan the prairies and the plains begin. These latter grassland formations belong to the Transition or Sub-boreal Zone. The number of plants common to the Canadian Zone of the East and the Montane Zone of the Rockies is small, outside of * See Bull. Torrey Club 42: 11-25. I915.- 295 296 RYDBERG: PHYTOGEOGRAPHICAL NOTES the common transcontinental species that grow along the water- courses and species which are common to the Hudsonian-Subalpine Zone and the Canadian-Montane Zone. None of theconifers are common to the two regions, and among the trees which occur in common we find only a few species of Salix, Populus, and Betula. If we consider the relationship between the Montane plants of the Rockies and those of the Pacific Coast Mountains, however, we find the conditions quite different. The Cascade Mountains are directly connected with the Rockies in the north and there are several mountain chains interposed between the Cascades and the Rockies in British Columbia, separated from one another only by narrow river-valleys. The Montane Zone of the Cascades, therefore, is practically continuous with that of the Rocky Moun- tains, and the two regions have many plants in common, the leading conifers not excluded. Many of the Pacific species have emigrated into the Rocky Mountains, especially into the northern Rockies, and many Rocky Mountain species into the Cascades. As has already been pointed out in earlier articles of mine, the Rockies may be divided into two principal parts, the Northern Rockies, extending from the Yukon Territory to northern Wyo- ming, and the Southern Rockies, of southern Wyoming, Colorado and northern New Mexico. Each of these main divisions may be subdivided, and the following districts may be distinguished: Northern Rockies 7. Big Horn District 1. Canadian Rockies 8. Black Hills District 2. Main Range, Montana Southern Rockies 3. Selkirk-Bitterroot District 9. Main Range District 4. Belt Mountains District 10. Uintah-Wasatch District 5. Yellowstone District 11. Sevier District 6. Sawtooth District Leal 2. La Sal-Abajo District The main range of the Northern Rockies north of Butte, Montana, is rather homogeneous and the change in the Montane flora seen in proceeding northward is rather gradual, although many of the high northern species found in the Canadian Rockies (District 1) are not found in Montana. South of Butte the main range (District 2) becomes lower and less distinctly Montane until the neighborhood of Yellowstone Park is reached. Here it rises higher and the Alpine Zone is present in the Bear Tooth, Sh Teton, Gros Ventre, and Mae ica ih RYDBERG: PHYTOGEOGRAPHICAL NOTES 297 Wind River Ranges. This region may be regarded as a distinct subdivision (District 5). Numerous species belonging to the Cascade Mountains have invaded the Selkirk Mountains in British Columbia and the Bitter- root Mountains between Idaho and Montana, and the flora here has become more or less Pacific in its character. This region (District 3) includes also more or less the western slopes of the main range in British Columbia and northern Montana. The mountains of Central Idaho, such as the Sawtooth, Sal- mon River, and other ranges (District 6) are characterized in the Montane Zone rather by the lack of many species found in the main range than by the introduction of any considerable new element. In the Submontane Zone, however, the character of the flora apparently is more like that of the mountains of the Great Basin than that of the main range, although the flora of this region is not so well known as might be desired. The isolated mountains of central Montana, such as the Belt Mountains, the Snowy Mountains, the Crazy Mountains, and the Little Rocky Mountains, together with the Cypress Hills in Canada (District 4), being rather low, are characterized by a meagre Montane flora without any additional element. This can also be said of the Big Horn Mountains of Wyoming (District 7), although these are much higher and contain a few strictly endemic species. The Black Hills of South Dakota and Wyoming are also to be counted as a subdivision of the Northern Rockies (District 8), although they contain quite a number of species belonging to the Southern Rockies as well as many belonging to the Canadian and Alleghanian Zones. In the Southern Rockies the main range (District 9) includes all of the mountains in Colorado and northern New Mexico, together with the Sierra Madre, Medicine Bow, and Laramie Ranges in Wyoming. Only a few peaks of the latter reach the Montane Zone. © The Uintah and the northern part of the Wasatch Mountains in Utah (District 10), although geologically belonging to the Northern Rockies, have a flora closely resembling that of the Southern Rockies. In the Wasatch Mountains, however, there 298 RYDBERG: PHYTOGEOGRAPHICAL NOTES are found quite a number of species which have immigrated from the Northern Rockies or from the mountains of the Great Basin. The southern part of the Wasatch Mountains, including the Sevier Range, the Henry Mountains, and other ranges west of the Colorado of the West (District 11), have a flora characteristic of the Basin Mountains. The Montane flora consists to a great extent of species common to the Rockies, the Basin Mountains, and the Sierra Nevada. The La Sal and Abajo Mountains, in Southeastern Utah (District 12) have a mountain flora almost the same as that characteristic of the main range of the Southern. Rockies in Colorado. The plants of the Montane Zone of the Rocky Mountains may be classified in the following categories. These are practically the same as those in the Subalpine Zone, with, however, some modi- fication. I. Transcontinental Species. II. Species common to the Rockies and the ede Zone of the East. , III. Species common to the Rockies and the Pacific Mountains. IV. Endemic species. I. Transcontinental Species The transcontinental species, as well as most of those com- mon to the Rockies and to the East, consist partly of forest species, most of which have migrated around the Saskatchewan Plains, partly of water, meadow, and thicket species which have followed the watercourses across the plains. The former consist to a great extent of species common to the Subalpine and Montane Zones in the Rockies and hence also to the Hudsonian and Cana- dian Zones of the East; the latter consist mostly of species found also in the Submontane and Transition (or Alleghanian) Zones. Many of the water and bog plants, however, are not found on the plains and hence must have followed the woods. A. TRANSCONTINENTAL SPECIES RANGING THROUGHOUT THE ROCKIES Here I have included species which are found in the main ranges of both the Northern and Southern Rockies. Many of RYDBERG: PHYTOGEOGRAPHICAL NOTES 299 these may be lacking in one or more of the districts or subdivisions, especially in Districts 8, 11, and 12, but some also in Districts 4, 6, and 7. I. PLANTS WITH BOREAL-SYLVAN DISTRIBUTION In this discussion the word ‘‘sylvan’’ applied to a plant does not mean that it grows only in the deep woods, but that its distribu- tion has taken place in connection with the Northern Woods, and that it does not grow in the prairie or plains regions. The plant may be a forest species in the true sense or it may grow in open woods, thickets, or among rocks in more open places. The essen- tial point is that its migration east or west has taken place around and north of the plains, and not across them along the water- courses. To this category belong the transcontinental trees and most of the shrubs. The most important of the former is the quaking aspen, Populus tremuloides, also found in the Subalpine Zone. : In this and subsequent lists, species which are marked are rare in the Southern Rockies; those marked ‘‘}”’ attain their best development at higher altitudes and reach the Alpine zone; those marked ‘‘t”’ develop best lower down and reach the plains; those followed by ‘‘(Eur.)”’ are found also in Europe and usually also in northern Asia. The nomenclature is that of the author’s Flora.* the KD a. Forest species Trees Populus tremuloides . Salix Bebbiana Populus balsamifera** Betula papyrifera** Shrubs Juniperus sibirica (Eur.) Lepargyraea canadensis Dasiphora fruticosa (Eur.) Arctostaphylos Uva-ursi (Eur.) Chamaepericlymenum Linnaea americana candense Distegia involucrata * Flora of The Rocky Mountains and Adjacent Plains. New York. 1917. 300 RYDBERG: PHYTOGEOGRAPHICAL NOTES Herbs Phleum alpinum (Eur.) Avena striata Danthonia spicata Poa compressa (Eur.) Allium sibiricum (Asia) Lysiella obtusata Peramium decipiens Cytherea bulbosa (Eur.) Corallorrhiza multiflora Tium alpinum (Eur.) Chamaenerium spicatum (Eur.) Heracleum lanatum Moneses uniflora (Eur.) Pyrola uliginosa Pyrola asarifolia** Pyrola chlorantha (Eur.) Pyrola ellipiica Erxlebenia minor (Eur.) Ramischia secunda (Eur.) Monotropa uniflora Veronica serpyllifolia (Eur.) Galium boreale (Eur.) Galium triflorum Specularia perfoliata Botrychium Lunaria (Eur.) Botrychium virginianum (Eur.) Botrychium silaifolium** Filix bulbifera Filix fragilis (Eur.) tlix montana (Eur.) Polystichum Lonchitis Theiypteris Dryopteris (Eur.) Asplenium septentrionale (Eur.) Asplenium Trichomanes (Eur.) Pteris aquilina (Eur.) Cryptogramma acrostichoides b. Aquatic and bog species The following water and bog plants probably reached the Rockies by the way of the Northern Woods: Shrubs Salix chlorophylla Betula glandulosa Herbs Sparganium minimum (Eur.) Triglochin palustris (Eur.) Scirpus caespitosus (Eur.) Carex diandra (Eur.)t Calamagrostis Langsdorfii (Eur.) Carex gynocrates (Eur.) Deschampsia caespitosa Catabrosa aquatica (Eur.) Panicularia nervata Panicularia borealis Pamicularia septentrionalis Carex disperma (Eur.) Carex canescens (Eur.) Carex brunnescens (Eur.) Carex paupercula (Eut.) Carex aquatilis (Eur.) Eriophorum angustifolium (Eur.) Bistorta vivipara (Eur.)T Scirpus pauciflorus (Eur.) Alsine borealis RYDBERG: PYTHOGEOGRAPHICAL NOTES 301 Sagina saginoides (Eur.) Mimulus moschatus Thalictrum alpinum (Eur.)t Limosella aquatica (Eur.) Batrachium flaccidum (Eur.) Limosella tenutfolia (Eur.) Ranunculus reptans Veronica Wormskjoldit{ Cardamine pennsylvanica Elephantella groenlandica Subularia aquatica (Eur.) Senecio pauciflorus Epilobium alpinum (Eur.)t Lycopodium annotinum (Eur.) Epilobium Hornemannii (Eur.)+ c. Various mesophytes A few plants which are neither aquatics nor forest species have invaded the Rockies from the north. These are species that grow among bushes, among rocks, or on hillsides. Torresia odorata (Eur.) Juncoides parviflorum (Eur.) Calamagrostis purpurascens Juncoides intermedium Poa crocata Juncoides spicatum (Eur.)t Festuca rubra (Eur.) Blitum capitatum (Eur.) Bromus ciliatus Moehringia latifolia (Eur.) Carex praticola Moehringia macrophylla Carex concinna Viola adunca Carex Halleri (Eur.) 2. PLANTS WITH RIPARIAN OR CAMPESTRIAN DISTRIBUTION These plants have crossed the continent over the plains, following mostly the watercourses. All of them are found in the foothills also and most of them attain their best development on the plains and _ prairies. a. Aquatic plants Sparganium angustifoliumt Panicularia grandist Potamogeton natans (Eur.)t Eleocharis palustris (Eur.)f Potamogeton alpinus (Eur.)t Eleocharis acicularis (Eur.){¢ Potamogeton lucens (Eur.)t Eriophorum gracile (Eur.)t Potamogeton foliosust Scirpus validust Triglochin maritima (Eur.) Lemna trisulca (Eur.)t Alisma brevipes Lemna minor (Eur.)t Sagittaria latifoliat Persicaria coccineat Phragmites Phragmites (Eur.)t Batrachium trichophyllum (Eur.)t 302 RYDBERG: Batrachium Drouetu (Eur.)t Ranunculus Purshitt Sisymbrium Nasturtium-aquati- cum (Eur.){ Tillaeastrum aquaticum (Eur.){ Callitriche palustris (Eur.)t Callitriche autumnalis (Eur.)t PHYTOGEOGRAPHICAL NOTES Hippuris vulgaris (Eur.)t Sium cicutaefolium Menyanthes trifoliata (Eur.)t Veronica americanat Utricularia vulgaris (Eur.)t Utricularia minor (Eur.)t b. Bog and wet meadow plants Tree Salix cordata Herbs Phalaris arundinacea (Eur.)t Alopecurus aristulatust Calamagrostis elongatat Calamagrostis canadensis} Beckmannia erucaeformist Carex leptalea (Eur.) Carex aureat Carex Buxbaumii (Eur.)t Carex lanuginosa Carex viridulat Carex rostratat Ranunculus sceleratus (Eur.) Halerpestes Cymbalariat Argentina Anserina (Eur.)t Gnaphalium uliginosum (Eur.) Equisetum arvense (Eur.) c. Meadow plants Agrostis hyemalist Muhlenbergia Richardsonis Poa annua (Eur.)t Poa triflora (Eur.)t Poa pratensis (Eur.)t Festuca octoflorat Hordeum jubatumt Carex stenophyllat Carex interior Juncus bufonius (Eur.)t Polygonum ramosissimumt Capnodes aureumt Arabis ovatat ' Draba nemorosat Viola nephrophyllat Viola septentrionalist Artemisia biennis To this category belong also some of the escaped cultivated plants and common weeds, such as Phleum pratense (Eur.)t Dactylis glomerata (Eur.)t Syntherisma Ischaemum (Eur.) eth nesee Botrys (Eur.)t Bursa Bursa-pastoris (Eur.){t Carum Carut (Eur.)f Plantago major (Eur. 5 RYDBERG: PHYTOGEOGRAPHICAL NOTES 303 d. Hillside plants Panicum Huachucae _ Balderdykia Convolvulus (Eur.) Ibidium strictum Pulsatilla ludoviciana Polygonum Douglasii B. TRANSCONTINENTAL SPECIES RESTRICTED TO THE NORTHERN ROCKIES Nearly all of the transcontinental plants restricted in their distribution to the Northern Rockies are of boreal-sylvan distribu- tion, whether they are forest species or not. In spreading across the continent, they have followed the northern woods around the plains north of the Saskatchewan and then south in the mountains. In some cases the species have not extended very far south— have not even entered the United States; in other words, their distribution in the Rockies is limited to District 1, the Canadian Rockies. Others have traveled farther south and invaded Dis- trict 2, or even Districts 3 and re About one third have spread further south into District 5 and from there to Districts 6, 7 and 8. Two are even found in the Uintah-Wasatch District of the Southern Rockies. I. SPECIES REACHING AT LEAST THE YELLOWSTONE DISTRICT a. Forest plants Sabina horizontalis Androsace septentrionalis (Eur.) Melica Smithit Valeriana septentrionalis Oryzopsis pungens Aster major Carex Peckii Aster Lindleyanus Streptopus amplexifolius Youngia nana (Asia){ Rosa acicularis (Asia) Aspidium viride (Eur.) Osmorrhiza divaricata b. Water and bog plants Eriophorum Scheuchzeri (Eur.)t Comarum palustre (Eur.) Eriophorum Chamissonis (Eur.)} Equisetum palustre (Eur.) Carex livida Equisetum fluviatile- Juncus Richardsonianus Equisetum laevigatum 304 RYDBERG: PHYTOGEOGRAPHICAL NOTES c. Cliff plant Antiphylla oppositifolia (Eur.)+ Of these species Oryzopsis pungens, Carex Peckii, Osmorrhiza divaricata and Juncus Richardsonianus have extended their range into the Black Hills of South Dakota, and Osmorrhiza divaricata and Youngia nana into the Uintah Mountains of Utah. 2. SPECIES REACHING THE MAIN RANGE IN MONTANA OR THE BITTER-ROOTS IN IDAHO, BUT NOT FURTHER SOUTH a. Forest plants Actaea rubra Lycopodium obscurum (Eur.) Ribes Hudsonianum Lycopodium complanatum (Eur.) Dryopteris Filix-mas (Eur.) Lycopodium clavatum (Eur.) Dryopteris dilatata (Eur.) Lycopodium sitchense b. Water and bog plants Shrubs Salix pedicellaris Salix candida Herbs Rynchospora alba (Eur.) Castalia Leibergii Scirpus subterminalis Drosera rotundifolia (Eur.) Scirpus atrocinctus Drosera longifolia (Eur.) Carex vaginata (Eur.) Geum macrophyllum Carex limosa (Eur.) Mertensia paniculata Carex scirpoidea (Eur.) Equisetum sylvaticum (Eur.) Carex lacustris (Eur.) Equisetum scirpoides (Eur.) Carex Crawfordii Lycopodium inundatum (Eur.) Carex flava (Eur.) Pellaea glabella, growing among exposed rocks, has a peculiar distribution. Though it is not found in the Southern Rockies it is found in the Black Hills and the hilly country of western Ne- braska and reappears in eastern Kansas. RYDBERG: PHY APHICAL NOTES 305 3. SPECIES LIMITED TO THE CANADIAN ROCKIES a. Forest species Carex aenea Coptis trifoliata (Eur.) Carex durifolia Thelypteris Phegopteris (Eur.) Cypripedium passerinum Dryopteris fragrans (Eur.) Lysias orbiculata . Woodsia glabella (Eur.) Ophrys convallarioides Lycopodium alpinum (Eur.) Comandra livida b. Water and bog plants Herbs Eriophorum callitrix (Eur.) Carex militaris Eriophorum opacum (Eur.) Carex deflexa Scirpus pumilust Oxycoccus Oxycoccus (Eur.) Carex chordorrhiza Oxycoccus macrocarpus II. Species common to the Rockies and the Canadian Zone The plants common to the Rockies and to the Canadian Zone of the East consist either of eastern species, whose ranges extend west into the Rockies, or of Rocky Mountain species which have invaded the East. The distribution of the former in the Rockies is very similar to that of the transcontinental species just treated. Some of them -have followed the northern woods, others the watercourses across the plains. The distribution area of some reaches the Southern Rockies, while that of others stops in north- ern Wyoming, in northern Montana, or in the Canadian Rockies. Among the Rocky Mountain species which have emigrated east, some have reached the Lake Superior region, while others are found as far east as the Gaspé Peninsula of Quebec. A. EASTERN SPECIES, EXTENDING SOUTH INTO THE SOUTHERN : ROCKIES 1. Plants of boreal-sylvan distribution a. Forest species Oryzopsis asperifolia Vagnera stellata Agrostis oreophila Coeloglossum bracteatum Cinna latifolia (Eur.) Peramium ophoides Carex Parryanat Corallorrhiza Corallorrhiza (Eur.) 306 RYDBERG: PHYTOGEOGRAPHICAL NOTES Claytonta virginica Prunella vulgaris (Eur.) Ranunculus micranthus Clinopodium vulgare (Eur.) Fragaria americana Pedicularis canadensis Rubus pubescens Linnaea americana Viola Selkirkii Erigeron droebachiensis (Eur.) Viola rentfolia Botrychium simplex (Eur.) Viola canadensis Botrychium neglectum (Eur.) Circaea alpina (Eur.) - Cryptogramma Stelleri Aralia nudicaulis Selaginella selaginoides b. Water and bog plants Alsine alpestris (Eur.) Epilobium adenocaulon Alsine crassifolia Petasites sagittata Parnassia parviflora Nabalus racemosa Geum rivale (Eur.) Lactuca spicata Viola palustris (Eur.) Equisetum pratense (Eur.) 2. EASTERN PLANTS WITH RIPARIAN OR CAMPESTRIAN DISTRIBUTION Muhlenbergia racemosa Sullivantia Hapemanii Juncus Vaseyi Vicia trifida Carex lanuginosa Vicia americana Urtica gracilis — : Apocynum androsaemifolium Carex siccata Macrocalyx Nycteleat Thalictrum dasycarpum Plantago eriopodat Lepidium densiflorumt Rudbeckia hirta Vitis riparia, Juncus Torreyi, and Asplenium platyneuron, eastern species, have reached the Southern Rockies in Colorado, but are not found in the Northern. 3. PLANTS OF THE GREAT PLAINS Some of the plants of the Great Plains extend up into the Montane Zone. Although they do not belong to the Canadian or Alleghanian Zones of the eastern United States, they are immi- grants from the East and may be included here. | Calamagrostis micrantha _ Plantago Purshii Polygonum buxiforme Laciniaria punctata Xanthoxalis Bushti Chrysopsis villos Viola pedatifida Solidago glaberrima Anogra latifolia Equisetum variegatum Anogra coronopifolia RYDBERG: PHYTOGEOGRAPHICAL NOTES 307 B. EASTERN SPECIES, EXTENDING INTO THE NORTHERN ROCKIES ONLY I. SPECIES REACHING AT LEAST THE YELLOWSTONE REGION Carex Richardsonu Phaca americana Carex eburnea Hedysarum americanum Heuchera hispida Of these, Heuchera hispida and Phaca americana even reach the Black Hills and western Nebraska. 2. SPECIES EXTENDING ONLY TO NORTHERN MONTANA OR IDAHO Carex tenera — Chiogenes hispidula Carex pedunculata Melampyrum lineare Parnassia palustris (Eur.) Thelypteris Robertiana (Eur.) Mitella nuda Dryopteris cristata (Eur.) 3. SPECIES LIMITED TO THE CANADIAN ROCKIES Trees Picea canadensis Picea mariana Shrub Salix pellita Herbs Carex atratiformis Primula mistassinica Vagnera trifoliata Petasites palmata Coptidium lapponicum (Eur.) _—Pteretis nodulosa Ribes glandulosum Dryopteris intermedia Geum perincisum _ Of these, Picea canadensis and Pteretis nodulosa are found also in the Black Hills. C. Rocky MouNTAIN SPECIES WHICH HAVE EMIGRATED EAST- WARD The following plants have extended their ranges as far east as eastern Minnesota, western Ontario, upper Michigan or Hudson Bay. Shrubs Rosa Bourgeauiana - Amelanchier alnifolia 308. RYDBERG: PHYTOGEOGRAPHICAL NOTES Herbs Viola rugulosa Aster Wilsonit Monarda menthaefolia Solidago pulcherrima Androsace subumbellata Erigeron glabellus Aster laevis Another species, Dryas Drummondii, has even reached the Gaspé Peninsula,’ Quebec. III. Species common to the Rockies and the Pacific Mountains A. SPECIES FOUND IN BOTH THE NORTHERN AND SOUTHERN ROCKIES, AS WELL AS THE CASCADES AND THE SIERRAS a. Forest species Nearly all of the plants belonging in this category have passed from the Rockies to the Pacific Mountains, or vice versa, in the . north where the two mountain systems are connected, and not across the Great Basin. Trees Pseudotsuga mucronata Apinus flexilis . Pinus Murrayana Salix Scouleriana Shrubs Odostemon A quifolium Pachystima Myrsinites Ribes viscosissimum Gaultheria humifusa Rubacer parviflorum Vaccinium scoparium Echinopanax horridum Herbs Oryzopsis Bloomert Ranunculus Douglasi Festuca subulata Ranunculus Bongardii Elymus glaucus Actaea arguta Carex Bolanderi Thalictrum sparsiflorum Carex athrostachya Aquilegia coerulea Veratrum speciosum Osmorrhiza obtusa Vagnera amplexicaulis Pectianthia pentandra Vagnera lilacina Fragaria bracteata Trillium ovatum Geranium viscosissimum Piperia unalaschensis Geranium Richardsonu RYDBERG: PHYTOGEOGRAPHICAL NOTES “309 Linum Lewistt Achillea lanulosa Circaea pacifica* Artemisia frigida (Asia) Glycosma occidentalis . Arnica cordifolia Chimaphila occidentalis Senecio pseudaureus Pyrola picta Hieracium albiflorum Pterospora Andromedea Athyrium alpestre (Eur.) Androsace filiformis (Asia) Athyrium cyclosorum (Eur.) Polemonium. occidentale Cryptogramma densa Collinsia parviflora Polypodium hesperium Pedicularis racemosa Cryptogramma densa, Athyrium alpestre,and Osmorrhiza obtusa reappear eastward on the Gaspé Peninsula, Quebec. When their distribution in Canada becomes better known, they may prove to belong among the transcontinental plants. Thalictrum sparsi- florum extends east to the Hudson Bay, Elymus glaucus to Upper Michigan, and Collinsia parviflora and Achillea lanulosa to western Ontario; Echinopanax horridum is found near the shores of Lake Superior. b. Water and bog plants In this class have also been included many plants of the wetter meadows and copses; in other words, the class consists of plants which probably have spread along the watercourses. This means, in this case, mostly along the Columbia River and its tributaries, for the Frazer River drains mostly the Cascades and other ranges west of the Rockies. Only the headwaters of this stream are in the Rocky Mountains, though north of the region here considered and mainly within the Subarctic Zone. The Colorado of the West runs for hundreds of miles in a deep canyon, does not touch the Sierras, and therefore can play practically no part in the distribution of the Montane plants. ’ Shrubs Salix glaucops Kalmia microphylla Alnus tenuifolia Herbs Muhlenbergia comatat Agrostis grandis Muhlenbergia filiformis — Agrostis asperifoliat 310 RYDBERG: PHYTOGEOGRAPHICAL NOTES Ranunculus Eschscholtzit Campe americanat Agrostis vartabilts Graphephorum Brandeget Danthonia californica Danthonia unispicata Carex Raynoldsit Carex tenuirostris Carex simulata Carex athrostachya Carex Kelloggit Lemna gibba (Eur.) Juncus Mertensianus Iris missouriensts Polygonum Watsonit Bistorta bistortotdest Claytonia lanceolata Parnassia fimbriata Micranthes arguta Epilobium brevistylum Epilobium occidentale Amarella strictifiora Mimulus Langsdorfii Mimulus Lewisiit Pedicularis bracteosa Galium subbiflorum Aster occidentalis Crunocallis Chamissonis Alsine strictiflora Alsine laetat Aster Burketi Erigeron salsuginosus Rudbeckia occidentalis Alsine calycantha Senecio triangularis Nymphaea polysepala Arnica longifolia Thalictrum sparsiflorum (Asia) Agoseris elata Myosurus aristatus : Of these, some have spread also to the East, as Crunocallis Chamissonis and Galium subbiflorum to Minnesota, Linum Lewistt to the Black Hills and Nebraska, Alsine strictiflora to Ontario, and Thalictrum sparsiflorum and Alsine laeta to Hudson Bay. c. Various mesophytes In this category are included plants that grow in open places, as dry meadows, table-land, hillsides and cliffs. Many of these grow also at lower altitudes; in the foothills and even on the plains. Many are also common to the mountain chains of the Great Basin. Trees or shrubs Cercocarpus ledifolius Limnobotrya montigena RYDBERG: PHYTOGEOGRAPHICAL NOTES Eriocoma hymenoidest Poa longtligula Poa Sandbergu Hesperochloa Kingtit Agropyron Smithiit Elymus condensatust Carex Douglasiit Carex Hoodit Carex phaeocephalat Carex obtusata (Eur.) Carex Rossii Juncus longistylis Juncoides comosum Eriogonum stellatum Eriogonum umbellatum Eriogonum ovalifolum Polygonum sawatchense Sarcobatus vermiculatust Eurotia lanatat Lewisia redeviva Oreobroma pygmaea Cerastium strictum (Eur.) Arenaria congestat Arenaria Burkett Anemone globosa Draba lutea (Eur.) Arabis retrofracta Sedum stenosepalum Lithophragma bulbifera Lithophragma parviflora Petrophytum caespitosum Potentilla Bakeri Potentilla diversifolia Drymocallis glandulosa Lupinus tenellus Lupinus argenteust Herbs Viola venosa Viola linguaefolia Epilobium paniculatumt Gayophytum intermedium Gayophytum ramosissimum Gayophytum racemosum Oenothera Hookert Lavauxia flava Leptodactylon pungens Leptodactylon Nutallia Gilia aggregata Collomia linearis Hydrophyllum capitatum Lappula floribunda Cryptantha Torreyana Pentstemon procerus Castilleja linariaefolia Campanula petiolata Macronema suffruticosum Aster campestris Antennarta rosea Antennaria oblanceolata Gymnolomia multiflorat Balsamorrhiza sagiitata Wyethia amplexicaulis Helianthus petiolarist Madia glomerata Chaenactis Douglasi Artemisia incompta Artemisia tridentatat Arnica pedunculata- Arnica fulgens Agoseris lacintata Woodsia oregana Woodsia scopulina Si RYDBERG: PHYTOGEOGRAPHICAL NOTES Of these Eriocoma hymenoides, Juncus longistylis, Arabis re- trofracta, Lithophragma bulbifera, L. parviflora, Petrophytum caes- pitosum, Drymocallis glandulosa, Epilobium paniculatum, Arnica pedunculata, and A. fulgens extend as far east as South Dakota and Nebraska; Gilia aggregata extends to Minnesota, reappearing farther east on the Gaspé Peninsula; Carex Rossii and Draba lutea to upper Michigan; Woodsia scopulina and W. oregana are found on the Gaspé Peninsula. B. RocKY MOUNTAIN SPECIES FOUND IN BOTH SOUTHERN AND NORTHERN ROCKIES, WHICH HAVE SPREAD INTO THE CAS- CADES, BUT ARE NOT FOUND IN SIERRA NEVADA a. Forest species Tree Picea Engelmanni Shrubs Sorbus scopulina Rubus melanolasius Herbs Calamagrostis luxurians Carex Geyert Opbhrys nephrophylla Razoumofskia americana Razoumofskia Douglasit Atragene columbiana ‘Ozomelis stauropetala Conioselinum scopulorum Ligusticum tenuifolium A pocynum ambigens Aster Geyert Erigeron macranthus Erigeron speciosus Erigeron conspicuus b. Water and bog plants Salix exigua Poa leptocoma Poa interior .- Poa Olneye Agrostis humilis Agrostis idahoensis Limnorchis stricta Limnorchis borealis Limnorchis viridiflora Ibidium porrifolium Shrub Herbs Alsine obtusa Ranunculus alismaefolius Ranunculus cardiophyllust Trollius albiflorus Delphinium multiflorum Argentina argentea Dodecatheon parviflorum Castilleja exilis Graphalium sulphurescens RYDBERG: PHYTOGEOGRAPHICAL NOTES 313 Of these Argentina argentea extends east to South Dakota and Ranunculus cardiophyllus to eastern Saskatchewan. c. Various mesophytes Melica spectabilis Melica bella Bromus polyanthus Carex nubicola Carex pachystachya Carex Geyer Carex filifoliat Carex scopulorum Anticlea elegans Juncus confusus Allium Geyert Delphinium Nelsoni Thlaspi Nuttallii Draba nitida Peritoma serrulaitum Leptasea austromontana Potentilla Nuttallia Drymocallis corymbosa Sieversia grisea Sieversia ciliata Thermopsis montana Lupinus caespitosus Astragalus striatus Astragalus goniatus Viola vallicola Phlox caespitosa Gilia pulchella Phacelia sericea Castilleja lancifolia Castilleja lauta Castilleja hispida Orthocarpus luteus Coleosanthus grandtflorus Aster apricus Antennaria flavescens Antennaria concinna Atnennaria corymbosa Antennaria pulcherrima Antennaria anaphaloides Artemisia Michauxiana Artemisia canat Artemisia arbuscula Artemisia tripartita Arnica Parryt Arnica mollis Arnica Rydbergit Senecio serra Selaginella densa Of these, Delphinium Nelsonii, Anticlea elegans, Juncus confusus, and Allium Geyert extend east to South Dakota or Nebraska, Astragalus striatus to Minnesota, and A. goniatus to Hudson Bay. 314 RYDBERG: PHYTOGEOGRAPHICAL NOTES c. PACIFIC SPECIES, WHICH HAVE INVADED THE NORTHERN ROCKIES, BUT NOT THE SOUTHERN I. SPECIES WHICH HAVE REACHED ONLY THE SELKIRK-BITTERROOT DISTRICT a. Forest species The species listed here have probably followed the mountain chains in their migration from the Pacific Mountains by the way of the Cascades and the Selkirks into the Rockies. Trees Strobus monticola Tsuga heterophylla Larix occidentalis Thuja plicata Abies grandis Taxus brevifolia Shrubs Ribes nevadense Azaliastrum albiflorum Rubus nivalis Vaccinium parvifolium Rubus spectabilis Linnaea longiflora Ceanothus prostratus : Herbs Carex laeviculmis Cytherea occidentalis Untfolium dilatatum Corallorrhiza Mertensiana Allium validum Trautvetteria grandis Disporum oreganum Mitellastra caulescens Clintonia uniflora Pectianthia Breweri Piperia multiflora Therophon majus Piperta elegans Chimaphila Menziesii Lysias Menziesii Castilleja pinetorum Eburophyton A ustinae b. Water and bog plants These have probably followed the Columbia River and Frazer River and their tributaries up into the mountains. Shrubs or trees Salix Lemmoni Salix sitchensis RYDBERG: PHYTOGEOGRAPHICAL NOTES Agrostis foliosa Calamagrostis Cusickit Graphephorum muticum Tofieldia occidentalis Cardamine oligosperma Lupinus polyphyllus Trifolium longipes Epilobum oregonense Epilobum glaberrimum Herbs Dodecatheon Jeffreyi Anthopogon simplex Mimulus nasutus Mimulus primuloides Mimulus Breweri Polystichum munitum Isoetes Howellit Isoetes Nutiallit C. Various mesophytes 315 These have probably spread along the foothills from the Cas- cades to the Selkirks and Bitterroots, or even across the Columbia Plains, as most of them are also found in the Submontane or Transition Zone. Stipa Thurberiana Festuca viridula Hordeum murinum (Eur.) Carex concinnoides Carex spectabilis Carex Mertensti Eriogonum pyrolaefolium Delphinium depauperatum Pulsatilla occidentalis Arabis suffrutescens Herbs Phoenocaulis cheiranthoides Sedum Douglasit Heterisia Meriensiana Dasystephana oregana Stenotus stenophyllus Antennaria confinis Balsamorrhiza deltoidea Arnica Menztesit Cheilanthes gracillima Selaginella Wallacet 2. SPECIES WHICH HAVE CROSSED THE MAIN RANGE OF THE ROcKIES IN MONTANA AND ALBERTA . Pinus ponderosa Ribes laxiflorum Rubus pedatus Forest species Trees Apinus albicaulis Shrubs Acer Douglasii 316 Melica subulata Aqutlegia formosa Tiarella untfoliata RYDBERG: PHYTOGEOGRAPHICAL NOTES Herbs Adenocaulon bicolor Pyrola dentata b. Water and bog plants Poa nervosa Naiocrene parvifolia Viola Macloskeyt Trifolium Beckwithit c. Various mesophytes Lupinus laxiflorus Phlox Douglasu Pedicularis contorta Arnica diversifolia Carex Preslit XA erophyllum tenax Anemone Drummondii Smelowskia ovalis Arabis Lyallit Adenocoulon bicolor reappears on Lake Superior and Trifolium Beckwithii in eastern South Dakota. 3. SPECIES WHICH HAVE REACHED THE YELLOWSTONE PARK a. Forest species Calamagrostis Suksdorfi Hypericum Scouleri Ozmorrhiza brevipes Salix A ustinae Panicum thermale Tofieldia intermedia Juncus nevadensis Stipa Elmer Stipa oregonensts Calamagrostis rubescens Carex Jonesit Carex nervina Carex luzulina Kelloggia galioides Aster integrifolius Antennaria racemosa b. Bog plants Shrub Herbs Alsine crispa Ranunculus alismellus Cardamine Breweri c. Mesophytes Potentilla Blaschkeana Potentilla glomerata Fragaria platypetala Dasystephana calycosa Townsendia scapigerat Balsamorrhiza terebinthaceat RYDBERG: PHYTOGEOGRAPHICAL NOTES Olt 4. SPECIES WHOSE DISTRIBUTION EXTENDS EVEN INTO THE UINTAH AND WASATCH MouNTAINS OF UTAH a. Forest species Shrubs Vaccinium occidentale Sambucus coerulea Herbs Aquilegia flavescens Apocynum pumilum Bicuculla unsflora b. Water and bog plants Shrub Ledum glandulosum Herbs Ruppia pectinata Alsinopsis occidentalis Limnia asartfolia Alsine brachypetala Limnia sibirica (Asia) Dodecatheon alpinum Limnia perfoliata Aster oreganus c. Mesophytes * Agrostis Thurbertana Arabis Lemmonit. Arenaria Douglasit Lupinus leucophyllus Thalictrum occidentale Gayophytum diffusum Paeonia Brownit Gayophytum pumilum Thlas pi californicum Lappula diffusa Draba oligosperma D. PLANTS COMMON TO THE NORTHERN ROCKIES AND THE Cas- CADES, BUT NOT FOUND IN THE SOUTHERN ROCKIES OR IN SIERRA NEVADA 1. SPECIES FOUND AS FAR SOUTH AS THE YELLOWSTONE PARK REGION a. Forest shrubs — Spiraea densiflora Menziesia ferruginea Spiraea lucida Menziesia glabella Salix 1dahoensis Salix Geyeriana Agrostis oregonensis Carex Piperi Carex microptera Caltha leptosepala Lupinus Burke Epilobium delicatum Sitanion montaum Elymus nitidus Carex Tolmet Eriogonum Piperi Spraguea multiceps Silene Lyallit Silene oregana Silene multicaulis RYDBERG: PHYTOGEOGRAPHICAL NOTES b. Bog plants Shrubs Alnus sinuata Herbs Angelica Lyallii Dodecatheon conjugens Aster Jessicae Senecio subnudus Botrychium Coulteri c. Various mesophytes Thalictrum columbianum Delphinium Nuttallianum Arabis albertina Lupinus leucopsis Lupinus sericeus Cordylophorum suffruticosum Amarella anisosepala Dasystephana monticola Pentstemon crassifolius Castilleja pallescens Valeriana ceratophylla Aster conspicuus Antennaria flagellaris Antennaria Howellii Artemisia floccosa Arnica gracilis Senecio Howellii Hieracium cynoglossoides Lupinus sericeus and Aster conspicuus extend east to the Black Hills. 2. SPECIES WHOSE RANGE EXTENDS EVEN INTO THE UINTAH- WASATCH REGION Juncus Regelii Silene columbiana Ranunculus limosus Delphinium bicolor Arabis rupestris Arabis microphylla Potentilla dichroa Castilleja Tweedyi Eucephalus elegans Hieracium albertinum Hieracium griseus Gnaphalium proximum RYDBERG: PHYTOGEOGRAPHICAL NOTES 319 3. SPECIES FOUND IN THE MAIN RANGE IN MONTANA AND ALBERTA BUT NOT FARTHER SOUTH Shrubs Salix Fernaldi Sorbus occidentalis Philadelphus Lewisu Vaccinium globulare Ribes petiolare Luetkea pectinata Rosa nutkana Herbs Poa Vaseyochloa Viola orbiculata Xerophyllum Douglasu Valeriana Scouleri Juncoides glabratum Penstemon Lyallit Erythronium grandiflorum Dodecatheon cylindrocarpum Eriogonum polyphyllum Dodecatheon viviparum Eriogonum depressum Castilleja Suksdorfit Arenaria nardifolia (Asia) Castilleja lutea Silene repens (Asia) Antennaria Howellit Atragene grosseserrata Antennaria luzuloides Arabis Nuttallit Erigeron Howellit Arabis furcata Aster modestus Heuchera glabra Aster diabolicus Heuchera grossulariifolia Aster Sayianus Micranthes aestivalis Achillea fusca Potentilla Drummondii Cirsium Macouniu Epilobium luteum Selaginella montanensis 4. CASCADE MOUNTAIN SPECIES WHICH HAVE EMIGRATED INTO THE SELKIRK-BITTERROOT REGION Shrub Salix commutata Herbs Alopecurus pallescens Eriogonum thymotdes Poa Cusicku Claytonia chrysantha Agropyron lanceolatum Alsine washingtoniana Elymus Howellii Arenaria cephaloidea Carex stenochlaena Aquilegia columbiana Allium fibrillum Sedum Leibergu Allium Cusickii* Hemieva ranunculifolia 320 RYDBERG: PHYTOGEOGRAPHICAL NOTES Aruncus acuminatus Trifolium plumosum Angelica Piperi Osmorrhiza Leibergit Osmorrhiza purpurea Ligusticum Canbyt Ligusticum Leibergu Chamaepericlymenum unalas- chense Moneses reticulata Pyrola bracteata Oxycoccus iniermedius Pentstemon pinetorum Veronica Cusickiu Synthyris major Castilleja cervina Razoumofskya Laricis Valeriana sitchensis Castilleja Vreeland Aster Cusicku Balsamorrhiza Careyana Saussurea americana Lactuca multifida Polystichum Andersont Thelypteris Oreopteris (Eur.) E. PLANTS COMMON TO THE SOUTHERN ROCKIES AND THE SIERRA NEVADA These plants have spread across the Great Basin, their seed being carried by birds or wind from mountain to mountain by way of the numerous though low parallel mountain chains within the Basin. Most of them are xerophytic, the rest mesophytic. Species in the following list marked ‘‘tt’’ extend into the Sub- montane Zone; those marked “tt” are not found east of the Wa- satch Mountains. Pinus aristata Elymus. simplest Muhlenbergia gracilistt Poa Fendleriana Carex fissuricolat{. Carex epapillosatt Rumex hymenosepalustt Amaranthus Powellit Quamoclidion multiflorumtt Oreobroma nevadensis Trees Abies concolor Herbs Erocallis triphylla Alsine Jamesii Lepidium nesinuaunes’ Heuchera rubescenstitt Sericotheca glabrescens{ttt Drymocallis pumilatt Trifolium Rusbyitt Phaca Hookerianatitt Hamosa calycosatitt Kentrophyta tegetariatt Hypericum formosum}{ RYDBERG: PHYTOGEOGRAPHICAL NOTES 321 Viola Beckwithiitt : Dugaldia Hoopesiit+ Viola Sheltont Senecio filicifoliustttt Microsteris micranthat} Polystichum scopulinumtt Hydrophyllum alpestrett Selaginella Watsonitt Macronema discoideum Selaginella Underwood IV. Endemic Rocky Mountain species As the endemic element of the montane plants is very large, consisting of about 1040 species, or over 53 per cent of the whole number, it is hardly practicable to list them all. I shall therefore merely give the number of species found in each category and mention specifically only a few in each class whose distribution is particularly characteristic or of special interest. A. ENDEMIC PLANTS COMMON TO THE SOUTHERN AND NORTHERN ROCKIES Many of the endemic plants are of wide distribution, their range extending from Colorado or northern New Mexico to Mon- tana or even further north. To this category belong the following trees: Pinus scopulorum, Populus angustifolia, Betula fontinalis, and six species of Salix (three of these usually mere shrubs). Among the shrubs, Sambucus melanocarpa, Ceanothus velulinus, and Svida instolonea are the most common and most widely distri- buted. The category contains 6 trees, 12 shrubs, and 217 herbs, and if to these are added 7 grasses and 6 other herbs belonging to the plains and occasionally reaching the Montane Zone, the whole number of species is 248. Of these some extend outside of the Rocky Mountains, as for instance Svida instolonea, which reaches to Manitoba and Kansas, Drymocallis fissa the Black Hills, and Scrophularia occidentalis North Dakota and Oklohoma. B. PLANTS ENDEMIC TO THE SOUTHERN ROCKIES ONLY I. PLANTS FOUND BOTH IN THE MAIN RANGE AND IN THE UINTAH- WASATCH REGION To this category belong two trees, Picea pungens and Populus Wislizenii, together with 13 shrubs and 197 herbs, or in all 212 species. Of these some are limited to the very southern slope of gee RYDBERG: PHYTOGEOGRAPHICAL NOTES the Rockies and are in reality immigrants from the Upper Sonoran region, as for instance, Populus Wislizenii, Grossularia leptantha, Blepharoneuron tricholepis, Calamagrostis scopulorum, Festuca arizonica and Fragaria ovalis. As I have stated elsewhere, the interchange of flora between the Southern Rockies and the Northern does not take place so much along the continental divide in central Wyoming as from the Wasatch Mountains, over the Bear River Mountains and the Tetons, to the Northern Rockies. Several southern species are found in the two intermediate ranges mentioned and _ several northern ones in the Wasatch. These southern species are as follows: Shrub Salix Wolfit Herbs Stipa Vaseyi Primula Parryi Rumex densiflorus Orthocarpus purpureo-albus Ranunculus intertextus Penstemon subglaber Delphinium occidentale Penstemon Rydbergii Delphinium reticulatum Aster Canbyi Cardamine cordifolia Senecio perplexus Arabis divaricarpa Senecio rapifolius Potentilla filicaulis Senecio uintahensis Sidalcea neomexicana Cirsium Eatoni Amarella monantha Leontodon scopulorum The following plants of the Southern Rockies extend into the Black Hills or western Nebraska. Poa andina Ceanothus Fendleri Eriogonum pauciflorum Cynomarathrum Nuttallii Arabis Fendleri Dodecatheon radicatum Draba auriformis Sambucus microbotrys Saxifraga simulata Thelesperma gracilis Potentilla propinqua Senecio spartioides Opulaster monogynus Senecio rapifolius RYDBERG: PHYTOGEOGRAPHICAL NOTES 323 2. PLANTS FOUND IN THE MAIN RANGE BUT NOT IN THE UINTAH- WASATCH REGION To this category belong nearly half of the endemic plants of the Southern Rockies. The list comprises 6 shrubs and 252 herbs, but no trees. Some of these are restricted to the southern slope only and may be considered as immigrants from the Upper Sonoran region. Among these are three of the four fernworts endemic to the Southern Rockies: Cheilanthes Fendleri, Notholaena Fendleri, and Selaginella mutica. A fourth fern, Woodsia mexicana, is also found in the Black Hills and in Minnesota, and ranges southward into Mexico. 3. PLANTS RESTRICTED TO THE UINTAH-WASATCH REGION This category comprises 7 shrubs and 71 herbs, but no trees. Many of these plants are also found in the mountains of the Great Basin. Some of them, as Fendlerella utahensis, Chamaebatiaria Millefolium, Arctostaphylos platyphylla, Phaca serpens, and Phaca Sileriana are evidently immigrants from the Upper Sonoran region. There are 47 local endemics found in Wyoming and south- eastern Idaho which occur nowhere else in the Rockies. Of these maybe one third should be counted as belonging to the southern Rockies. If so, the total number of endemics restricted to the southern Rockies would be about 560 species. C. PLANTS ENDEMIC TO THE NORTHERN ROCKIES ONLY 1. PLANTS OF GENERAL DISTRIBUTION WITHIN THE NORTHERN ROCKIES Fully one-third of the endemic species of the Northern Rockies are of general distribution and extend as far south as the Yellow- stone Park Region. Among these are included two trees, Picea _albertiana and Betula utahensis, 4 shrubs and 102 herbs; altogether 108 species. Of these the following extend south into the Uintah- Wasatch region. Juncus Tweedyi Delphinium bicolor Cardamine multifolia Aconitum divaricatum Ranunculus saxicola Draba andina 324 RYDBERG: PHYTOGEOGRAPHICAL NOTES Arabis oreophila Swertia congesta Arabis exilis Synthyris laciniata Micranthes Greenet Orthocarpus Tolmiet Potentilla ovina Aster amplifolius Drymocallis foliosa Machaeranthera viscosa Trifolium scariosum Erigeron tenellum Angelica Roseana Arnica arcana Dodecatheon salinumt The following reach the Black Hills: Alsinopsis dawsonensis Aragallus gracilis Atelophragma glabriuscula Aragallus spicatust Atelophragma Forwood11 Aster meritus Homalobus dispar Cirsium Drummondii Aragallus villosus 2. SPECIES FOUND IN MONTANA AND NORTHERN IDAHO AND NORTHWARD This category contains one tree, Betula subcordata, two shrubs (Vaccinium sp.), and 31 herbs; altogether 34 species. Of these, Aragallus splendens extends eastward to Minnesota, Vaccinium membranaceum to upper Michigan, and V. ovalifolium to the Gaspé Peninsula. 3. LOCAL SPECIES OR SPECIES OF VERY RESTRICTED RANGE The local species of Wyoming and eastern Idaho number 47 (of which perhaps one third should be accredited to the Southern Rockies while 6 are also found in the Uintahs or Wasatchs), those of Montana 19, those of western Idaho 14, those of the Black Hills 5, and those of the Canadian Rockies 22 (among the latter one tree, Betula alaskana, and three shrubs); in all, 107 local species. If all categories of endemic species are considered, the number restricted to the Northern Rockies includes altogether about 230 species, and the number endemic to the — Mountains as a whole 1,040 species. SUMMARY Within the Montane Zone in the Rocky Mountains are found about 1900 species. Of these, approximately 50 per cent* are * All percentage figures in the following paragraphs are computed with reference to the total number of species. RYDBERG: PHYTOGEOGRAPHICAL NOTES 325 Montane plants in the ‘restricted sense, i. e. plants which attain their best development within this zone. Of the rest, many reach their best development in the Subalpine Zone above, and many others in the Submontane Zone below. A few alpine plants are sometimes found as low as the Montane Zone and several species from the Great Plains or from the Sonoran Zone are occasion- ally found as high up. Of the plants found in the Montane Zone, 245, or less than 13 per cent, are transcontinental, i. e. they are found both in the East and on the Pacific Slope, as well as in the Rockies; 176 of these, or 9 per cent of the total flora, are common to the Northern and Southern Rockies, another 1 per cent extend as far south as Wyoming, and 1 per cent are limited to the Canadian Rockies. Besides the transcontinental plants, there are 84 species which are common to the East and the Rockies but have not reached the Pacific Slope. If to these are added a score of western plants which have emigrated eastward as far as the Great Lakes and Hudson Bay, some of them even to the Gaspé Peninsula, Quebec, there are in all about 350 species, or nearly 1834 per cent of the flora, which are common to the East and to the Rockies. The larger portion of these, 250 species or over 13 per cent, are found in both the Northern and Southern Rockies, and 100, or more than 5 per cent, in the Northern only. None of the Montane plants are common to the East and the Southern Rockies only. The number of species common to the Rockies and the Pacific Mountains is much larger: if the transcontinental species are excluded, about 565, or nearly 30 per cent; or, if these are included nearly 43 per cent of all the plants found within the Montane Zone of the Rockies. Nearly 450 of the plants common to the Rockies and the Pacific Mountains (the transcontinental ones included), or nearly 24 per cent of the whole number are found in both the Northern and Southern Rockies; 350, or about 18 per cent, are found in the Northern Rockies but not in the Southern, and not quite 2 per cent occur in the Southern but not in the Northern Rockies. Of the species common to the Rockies and the Pacific Mountains (the transcontinental ones not included), about 300, or nearly 16 per cent, are found both in the Cascades and the Sierra Nevada; about 225, or nearly 12 per cent, are found in the 326 RYDBERG: PHYTOGEOGRAPHICAL NOTES Cascades alone, and less than 2 per cent in the Sierra Nevada alone. If the transcontinental plants are added, the percentage for the plants found in both the Sierras and the Cascades and for those found in the Cascades alone would be increased to about 20 per cent of each. The ratio for those found in the Sierras alone would remain less than 2 per cent. This refers of course to the Montane plants only. In the Submontane region the ratio would be much greater. The endemic element consists of 1,046 Montane plants or over 53 per cent of the whole number. Of these, 245 or nearly 13 per cent are common to the Northern and Southern Rockies, 560 or nearly 29 per cent being restricted to the Southern Rockies, and 230 or 12 percent to the Northern. Of the latter about 10 per cent extend as far south as Northern Wyoming and I per cent are restricted to the Canadian Rockies. Of the species restricted to the Southern Rockies over 11 per cent are common to the Main Range in Wyoming, Colorado, and New Mexico and the Uintah- Wasatch region, while 13% per cent are restricted to the former and nearly 4 per cent to the latter. Of those restricted to the Northern Rockies, 8 per cent are found in Northern Wyoming and a little over I per cent are restricted to the Canadian Rockies. If the flora of the Canadian Rockies were better known this latter number probably would be much larger. We must also remember that only the region south of latitude 55° is here considered. It may also be of interest to see how the number of species found in the Southern and Northern Rockies would compare, if all categories of Montane plants are taken in consideration. There are over 40 per cent common to both, nearly 28 per cent restricted to the Southern Rockies and 32 per cent to the Northern. The species found in the Southern but not in the Northern Rockies consist almost wholly of endemic forms, less than 2 per cent being common to the Southern Rockies (and most of these found only in Utah) and the Sierra Nevada and 3 species only being Eastern nontranscontinentals. Those found in the Northern Rockies and not in the Southern consist of over 3 per cent transcontinental species, less than 2 per cent being common to the Northern Rockies and the Eastern Canadian Zone, nearly 15 per cent common to the former and the Pacific Mountains and 12 per cent endemics: altogether 32 per cent of all Montane species. RYDBERG: PHYTOGEOGRAPHICAL NOTES 327 As stated before, the interchange of species between the two great divisions of the Rockies has taken place from the northern part of the Wasatch Mountains over the Bear River Mountains and the Teton Mountains to the Northern Rockies, or vice versa, rather than along the continental divide in central Wyoming. Among the Montane plants, I have listed 69 northern species, or over 3% per cent of the total flora, which are found in the Wasatchs but nowhere else in the Southern Rockies, and 23 Southern species, or over I per cent, which are found in southern Idaho or in the Teton Mountains, and nowhere else in the Northern Rockies. A good deal could also be said about the distribution of the plants in the Black Hills, a meeting place of plants from the North- ern Rockies, the Southern Rockies, the Canadian and Alleghanian Zones of the East, and of the flora of the Great Plains, and I hope to take up this subject at some future time. NEW YorRK BOTANICAL GARDEN INDEX TO AMERICAN BOTANICAL LITERATURE 1917-1919 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 pa “pe that relate exclusively to forestry, agriculture, horticulture, lactate products of vegetable origin, or laboratory methods are not included, an no attempt is made to index the literature 7 Sek tyes An occasional exception is made in favor of some paper appearing in an American periodical which is devoted at to botany. Reprints are not fe a wl they differ from the original in important particular. If users of the Index will call the attention of the editor ao 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. Bryological notes—V. Scapania nimbosa from Nor- way. Torreya 1g: 49-51. 14 My 1919. Andrews, E.F. The Japanese honeysuckle in the eastern United States. Torreya 19: 37-43. f. 1, 2. 14 My 1919. Arthur, J. C. New names for species of Phanerogams. Torreya 19: 48, 49. 14 My Ig19. Arthur, J.C. New wa ag of Uredineae—XI. Bull. Torrey Club 46: 107-125. 1 My1 New species are peaeiy in Puccinia (8), Uredo (3), and Aecidium (5). Arthur, J. C. Research as a university function. Science II. 49: 387-391. 25 Ap 1919. Beardslee, H. C. A new species of Amaniia. Jour. Elisha Mitchell Soc. 34: 198, 199. pl. 30, 31. Mr 1919. Amanita mutabilis sp nov. Berg, A. A simple method of distinguishing nematode galls of wheat from bunted kernels. Phytopathology 9: 181,182.f.7. 2Ap 1919. Berry, E. W. Geologic history of the locust and its allies. Plant World 21: 284-298. f. r, 2. N 1918. Blakeslee, A. F. & Avery, B.T. Mutations in the jimson weed. Jour. Heredity 10: 111-120. f. 5-15. 25 Ap 1919. 329 390 INDEX TO AMERICAN BOTANICAL LITERATURE Butler, A. E. Notes by a collector in the Colorado Rockies. Nat. Hist. 19: 170-181. Mriogrig._ [Illust.] Bouquet, A. G. B. Pollination of tomatoes. Oregon Agr. Exp. Sta. Bull. 157; 1-29. f. 7-5. Mr 1919. Brown, F. B. H. The preparation and treatment of woods for micro- scopic study. Bull. Torrey Club 46: 127-150. f. 1-6. 1 My 1919. Chamberlain, E. B. Anacamptodon splachnoides var. Tayloriae in Missouri. Bryologist 22: 16. 30Ap 1919. Chamberlain, C. J. The living cycads. i-xiv+1-172.f. 1-91. Chi- cago. 30 Ap 1919. Clokey, I. W. Carex notes. Rhodora 21: 83-85. 9 My 1919. Includes Carex arapahoensis and C. subimpressa, spp. nov. Coit, J. E., & Hodgson, R. W. An investigation of the abnormal shedding of young fruits of the Washington navel orange. Univ. Calif. Pub. Agr. Sci. 3: 283-368. pl. 25-g2+f. 1-9. 4 Ap 1919. Coker, W.C. The Hydnums of North Carolina. Jour. Elisha Mitchell Soc. 34: 163-197. pl. 1-29. Mr 1919. vat aragamaan ferrugipes, H. carolinianum and Phellodon Cokeri Banker, spp. nov., are described. Collins, F.S. The green algae of North America, second supplementary paper. Tufts College Stud. 47: 1-106. pl. 1-3. 1 Je 1918. Collins, F.S. A working key to the genera of North American algae. Tufts College Stud. 48: 1-50. N 1918 Coulter, J. M. The botanical opportunity. Science II. 49: 363-367: 18 Ap rgI9. Cox, P. Dirca palustris L. in New Brunswick. Ottawa Nat. 32: 170. Mr 1919. Davis, J. J. North American Ascochytae. Trans. Wisconsin Acad. Sci. 19: 655-670. I9gI9. Davis, J. J. Notes on parasitic fungi in Wisconsin—IV. Trans. Wisconsin. Acad. Sci. 19: 671-689. 1919;—V. 690-704. 1919; —VI. 705-727. 1919. [Hlvet.] Twenty-three new species in various genera are described. Deatrick, E. P. The effect of manganese compounds on soils and plants. Cornell Agr. Exp. Sta. Mem. 19: 371-402. F 1919. Durfee, T. Lichens of the Mt. Monadnock region, N. H.—No. II. Bryologist 22: 15, 16. 30 Ap Ig19. Durand, E. J. Encalypia laciniata in central New York. ee 22: 13. 20 Ap 1916: INDEX TO AMERICAN BOTANICAL LITERATURE 301 Ensign, M.R. A staining method for vascular tissue. Phytopathology g: 180. 2 Ap 1919. Ensign, M. R. Sweet potato mosaic. Phytopathology g: 180, 181. 2 Ap 1919. Fairchild, D. Present condition and opportunity of the American Genetic Association. Jour. Heredity 10: 65-67. 5 Ap 1919. Farwell,O.A. Bromelica (Thurber): a new genus of grasses. Rhodora 21: 76-78. 9 My 1919. Fernald, M. L. Lithological factors limiting the ranges of Pinus Banksiana and Thuja occidentalis. Rhodora 21: 41-67. 14 Ap 1919. [Illust.] Fitzpatrick, H. M. George Francis Atkinson. Science II. 49: 371, 372. 18 Ap 1919. Fromme, F. D. Cedar rust. Virginia Hort. Soc. Ann. Rep. 23: I-11. 1918. Fromme, F. D., & Wingard, S. A. Bean rust: its control through the use of resistant varieties. Virginia Agr. Exp. Sta. Bull. 220: 1-18. pl. 1-5. N 1918. Giddings, N. J. Infection and immunity in apple rust. West Vir- ginia Agr. Exp. Sta. Tech. Bull. 170: 1-71. pl. 1-11. D 1918, Gould, H. P. Fig growing in the South Atlantic and Gulf states. U.S. Dep. Agr. Farm. Bull. 1031: 1-45. f. I-24. Mr 1919. Gray, J., & Peirce, G. J. The influence of light upon the action of stomata and its relation to transpiration of certain grains. Am. Jour. Bot. 6: 131-155. f. 7-18. 1 My 1919. (Greenman, J. M.] Early collections in the Garden Herbarium. Missouri Bot. Gard. Bull. 7: 29-35. pl. S11. Mr 1919. [Greenman, J. M.} Plants in the aroid house. Missouri Bot. Gard. Bull. 7: 35-38. pl. 7-5 Mr 1919. Harper, R. M. Some vanishing scenic features of the southeastern United States. Nat. Hist. 19: 193-204. F 1919. [Illust.]. Hayes, H. K., & Stakman, E. C. Rust resistance in timothy. Jour. Am. Soc. Agron. 11: 67-70. F 1919. Hoagland, D. R. Note on the technique of solution culture experi- ments with plants. Science II. 49: 360-362. 11 Ap IgI9. Haas, A. R. C. Respiration after death. Bot. Gaz. 67: 347-365- f. 1-3. 18 Ap 1919. Hottes, A.C. Commercial plant propagation. 1-180. f. I-105. York. 1918. New oon INDEX TO AMERICAN BOTANICAL LITERATURE Howe, M. A. Dahlias and their culture. Jour. Hort. Soc. New York 11: 285-301. pl. 45+. 1-3. Reprinted abridged and with substitute illustrations in Gard. Chron. Am. 23: 152-154. My 1gI9. Johnson, A. T. Arbutus Menziesii. The Garden 83: 124. 15 Mr 1919. Johnson, J., & Milton, R.H. Strains of white burley tobacco, resistant to root-rot. U.S. Dept. Agr. Bull. 765: 1-11. f. 7-4. 18 Ap 1919. Jones, L. R. Our journal, Phytopathology. Phytopathology 9: 159- 164. 2 Ap 1919. Lee, H. A. Plant pathology in Japan. Phytopathology 9: 178, 179. 2 Ap 1919. Leighty, C. E., & Hutcheson,T. B. On the blooming and fertilization of wheat flowers. Jour. Am. Soc. Agron. 11: 143-162. f. II, 12. Ap 1919. Lloyd, F.E. The origin and nature of the mucilage in the cacti and in certain other plants. Am. Jour. Bot. 6: 156-166. 1 My 1919. Love, H. H., & Craig, W. T. The synthetic production of wild wheat forms. Jour. Heredity 10: 51-64. f. 1-9+frontispiece. 5 Ap 1919. Lowe, R. L. Collecting in Arkansas. Bryologist 22: 14,15. 30 Ap 1919. Lyman, G. R. The unification of American botany. Science II. 49: 339-345. 11 Ap 19109. Lyon, T. L. Influence of higher plants on bacterial activities in soils. Jour. Am. Soc. Agron. 10: 313-322. D 1918. MacMillan, H. G. Fusarium-blight of potatoes under irrigation. Jour. Agr. Research 16: 279-303. pl. 37-41. 17 Mr 1919. Macoun, J. M. Habitat of Carex Franklinii Boott. Ottawa Nat. 32: 169. Mr 1919. Martin, J. N. Botany for agricultural students. i-x-+1-585. f. 1-488. New York. 1919. McClelland, T. B. Vanilla: a promising new crop for Porto Rico. Porto Rico Agr. Exp. Sta. Bull. 26: 1-32. pl. 1-3+f. 1-4. 17 AP 1919. Moore, G. T. Botanical participation in war work. Science II. 49: 269-274. 21 Mr 1919. Murrill, W. A. The natural history of Staunton, Virginia. i-xiii+I- 266. pl. 1-4. New York. _ Nichols, G. E. Lophiola aurea in Nova Scotia. Rhodora 21: 68. 14 Ap 1919. INDEX TO AMERICAN BOTANICAL LITERATURE ooo Nichols, G.E. Raised bogs in eastern Maine. Geog. Rev: 7: 159-167. Tod, Bear FOt0. Orton, C. R., & Kern, F. D. The potato wart disease. Pennsylvania Agr. Exp. Sta. Bull. 156: 1-16. f. 7-4. Mr 1919. Pennell, F. W. Eysenhardtia. N. Am. Fl. 24: 34-40. 25 Ap 1919. Seven new species are described. Piper, C. V. New Pacific coast plants. Proc. Biol. Soc. Washington 32: 41-44. 11 Ap 1919 Includes Cryptantha suffruticosa, Sidalcea nelsoniana, Stachys caurina, S. con- fertiflora, S. ciliata macrantha, Penstemon deserticola, and Circium oreganum, spp. nov. Popenoe, W. The avacadoin Guatemala. U.S. Dept. Agr. Bull. 743: I-69. pl. 1-23. 17 Ap 1919. : Pomeroy, C. S. Bud variations in sugar cane. Jour. Heredity 1o: 129-135. f. 16, 17. 25 Ap 1919. Reed, H. S., & Holland,R. H. The growth rate of an annual plant Helianthus. Proc. Nat. Acad. Sci. 5: 135-144. f. 1-3. Ap 1919. Roberts, H. F. The founders of the art of breeding. Jour. Heredity 10: 99-106. f. 1-4. 25 Ap 1919; II. Jour. Heredity 10: 147-152. f. 1 + frontispiece. 26 My 1919; III. Jour. Heredity 10: 229-239. f. I-20. 15 Jl 1919. Robinson, B. L. An unusual Daucus Carota. Rhodora 21: 70, 71. 14 Ap 1919. Rock, J. F. A monographic study of the Hawaiian species of the tribe Lobelioideae family Campanulaceae. i-xvit+1-394. pl. 1-217. 1919. Includes new species in Lobelia (4) and Cyanea (2). Rolfe,R: A. Govenia lagenophora. Curt. Bot. Mag. IV. 15: pl. 8794. Mr 1919. A plant from Mexico. Rolfe, R. A. Jsabelia virginalis. Curt. Bot. Mag. IV. 15: pl. 8787. Mr 1919. A plant from Brazil. Rose, R. C. After-ripening and germination of seeds of Tilia, Sam- bucus, and Rubus. Bot. Gaz. 67: 281-308. 18 Ap 1919. Rosendahl, C. O. Variations in the flowers of Erythronium propullans Gray. Torreya 19: 43-47. f. 1-3. 14 My I919. Rowlee, W. W. Synopsis of the genus Ochroma, with descriptions of new species. Jour. Washington Acad. Sci. 9: 157-167. 19 Mr 1919. Six new species are described. Rydberg, P. A. Key to the Rocky Mountain Flora. 1-305. New York. 1919. i-xiiit+1-216. pl. 1-4. 1919. 334 INDEX TO AMERICAN BOTANICAL LITERATURE Rydberg, P. A. (Rosales) Fabaceae. Psoraleae. N. Am. Fl. 24: 1-64. 25 Ap 1919 Includes Hotta, a a Psoralidium, Pediomelum, Psorobatus, Psoroden- dron, Psorothamnus, and new species in Hoita (4), Psoralidium (1), Pedio- melum (7), Amorpha an Baeuadee (2), Psorothamnus (4), and Parosela (8). Schneider, C. Notes on American willows—IV. Species and varieties of section longifoliae. Bot. Gaz. 67: 309-346. 18 Ap 1919. Several new varieties are described. Shear, C. L. First decade of the American Phytopathological Society. Phytopathology 9: 165-170. 2 Ap 1919. Shufeldt, R. W. Mandrakes, wild lupine, and notes on American snapping turtle. Am. Forestry 25: 995-1000. f. 1-9. Ap 1919. Skinner, J. J.. & Reid, F. R. The influence of phosphates on the action of alpha-crotonic acid on plants. Am. Jour. Bot. 6: 167-180. f. I-9.. 1 My 1919. Smith,G. M. A second list of algae found in Wisconsin lakes. Trans. Wisconsin Acad. Sci. 19: 614-654. pl. 10-15. Includes the new genus Plankiosphaeria and new species in Piniksepiers (1), Asterococcus (1), Oocystis (1), Tetraedron (1), Characium (1), and Chlorolotrys (1). Stakman, E. C., Levine M. N., & Leach, J.G. New biologic forms of Puccinia graminis. Jour. Agr. Research 16: 103-105. 20 Ja 1919. Standley, P. C. A neglected Solidago name. Rhodora 21: 69, 70: 14 Ap 1919. Stevens, N. E. Keeping quality of strawberries in relation to their temperature when picked. Phytopathology9: 171-177. 2Ap 1919. Stevenson, J. A. A check list of Porto Rican fungi and a host index. Jour. Dept. Agr. Porto Rico. 2: 125-264. Jl 1918. Stout, A.B. Bud variation. Proc. Nat. Acad. Sci. 5: 130-134. f. 18. Ap I9g19. Tisdale, W. H. Physoderma disease of corn. Jour. Agr. Research 16: 137-154. pl. A, B, to-17 +f. 1. 3F 1919 Tottingham, W. E. A preliminary study of the influence of chlorides on the growth of certain agricultural plants. Jour. Am. Soc. Agron. 11: 1-32. Ja 1919. Transeau, E. N., & Tiffany, H. New Oedogoniaceae. Ohio. Jour. Sci. 19: 240-242. pl. 14. F 19109. O. hystricinum and Bulbochaete Bullardi, spp. nov., are described. Van Fleet, W. New everbearing strawberries. Jour. Heredity 10: 14-16. f. 6,7. 8 Mr 1919. BULL. TORREY CLUB VOLUME 46, PLATE 12 3 4 5 j 8 6 \ \\\ WZ Zz ae Ss > S - = \ MATONIDIUM AMERICANUM BERRY But. TORREY CLUB VOLUME 46, PLATE 13 <2 ROE L2 <2 1,2. MATONIA PECTINATA R.Br. 3-6. MATONIDIUM AMERICANUM Berry Vol. 46 No. 9 BULLETIN OF THE TORREY BOTANICAL CLUB SEPTEMBER, 1919 The development and structure of the bulb in Cooperia Drummondii MARGARET B. CHURCH (WITH PLATES 14-16 AND NINE TEXT FIGURES) A survey of the literature dealing with the Amaryllidaceae and closely related families reveals the lack of detailed study of bulbs, such as modern methods of microtechnique and the com- pound microscope make possible. The usual textbook drawings of bulbs are extremely hazy as regards the nature of the vegetative point and even of the neighboring and slightly older members. The present study is submitted with the hope of supplying a more complete knowledge of the development and structure of a typical bulb, together with diagrammatic representations of the same. HISTORICAL Herbert (13), in his discourse on the Amaryllidaceae, states that the genus Cooperia has ‘‘black shelly seeds.” He received his bulbs of Cooperia Drummondii (12) from travelers and grew them in his greenhouse or outdoors—according to season—as he did his other plants for study. ‘One bulb in six months produced five successive scapes,’’ is his experience. ‘‘As soon as the seed on one scape is ripe another seems ready to rise. . . . Its habit appears to be to flower successively from earliest spring till September with leaves principally in autumn or winter.”” Flower- ing in the greenhouse at Brown University was at its best during March and February. No successive flowering of any one plant occurred extensively. Indeed it was deemed good fortune if {The BuLLETIN for August (46: 285-336. pl. 12, 13), was issued September 27, 1919.] 337 338 CHURCH: THE BULB IN COOPERIA DRUMMONDII several out of a dozen bulbs containing immature flowers blossomed at all. Although no criticism is intended here on either hand, it seems fair to state that Herbert was not justified in assuming that he could determine in European greenhouses or gardens the habit of bulbs native to Texas. He found his Cooperia Drum- monduw with a ‘‘flower [which] expands always in the evening, usually not perfect after the first night.’’ Nocturnal flowering is an anomaly, as Herbert himself states, and it seems probable that he was overzealous here in his observations, since the writer over a period of three years observed no new flowers of Cooperia Drummondti except in the morning and also observed no flowers opening except in the bright morning light. In fact the flowers in the greenhouse rarely shoot up as late as after eight o’clock in the morning. It is claimed that the name “‘rain lily”’ is applied to this amaryllid on its native ground, because it does shoot up into full bloom on a bright morning after a period or night of rain. Irmisch (16-18) described in detail the structure and develop- ment of many bulbs among the Liliaceae and Amaryllidaceae, and a short article on bulbs contemporary with Irmisch’s works was published by Koch (20) in 1849. In this last paper bulbs are classified as perennial, periodic and biennial. We find in years subsequent to 1850 no important work undertaken on the bulb excepting the biological studies of Rimbach (23-28) published around 1897. | A number of articles, important indirectly if not directly, may, however, be recorded here for the future reference of other students. In 1870 Hanstein (10) published an article on embryo development in both monocotyledonous and dicotyledonous plants. The species of plants discussed in this work are of no immediate interest, yet the article deserves study and consideration because its author is a clear-headed pioneer in showing the relation of physiology. to morphology and anatomy. Baillon (1) comes closer to the subject at hand while describing what seem to have been bulbiform abnormalities in the development of the embryos of Hymenocallis, Crinum, etc. The anatomy or structure of the vascular traces and secondary roots, and the relation of leaves and shoot axis in certain monocotyledons, e.g. Allium Cepa and Lilium Martagon, are well comprehended by Falkenberg (7)- CHURCH: THE BULB IN COOPERIA DRUMMONDII 339 The work of Solms-Laubach (30) is known and well established in reputation. Tschirch (32) describes some interesting observa- tions in regard to the cotyledon of Lupinus during its development in the seedling. Baranetsky’s work (2) on the development of vegetative points in monocotyledons is exceedingly valuable, discussing as it does among other points the development of subepidermal cambium, foliar buds, ensheathing leaves, internodes and permanent tissues for nine specific monocotyledons. It enters into details of anat- omy further than the present paper makes any pretense of doing. A paper on the germination of the Amaryllideae by Worsley (36), delivered before the Royal Horticultural Society, is in- accurate and needs no consideration. The structure of a garlic bulb has been described in a general fashion by Highfield (14). The duration of bulb parts, the maturity of seedling plants, and the placing of the bulb slightly below the surface of the ground by means of roots formed from the base of the cotyledon are noted briefly. Howard (15) has experimented with the mature plants of Cooperia Drummondii, while working on the summer rest period of plants. He does not mention any studies with “‘offsets.”’ METHODS AND MATERIAL The mature bulbs for this study of Cooperia Drummondu were secured from the campus of the University of Texas through the courtesy of Dr. H. H. York, of Brown University. A portion of the bulbs planted in the greenhouse at Brown University flowered freely and produced seed, which on maturity was used to secure all young plants. The methods of microtechnique usually followed were found - unsatisfactory, since the bulbs of Cooperia Drummondii appear to be particularly troublesome material for embedding. Their delicate scales contain starch, a mucilaginous slime which may coagulate during killing and fixation, and crystals of calcium oxalate (see Menz, 22). The scales and stem-axis are composed of parenchyma cells and vascular bundles, which resist the section- ing individually. Material was fixed in an aqueous solution of Picro-acetic* acid allowed to act for twenty-four hours. A water * 1 gram of picric acid, I c.c. of acetic acid, and 100 c.c. of water. 340 CuHuRCH: THE BULB IN COOPERIA DRUMMONDII solution of the fixing agent was found preferable to an alcoholic solution, since in fixing material such as Amaryllis the alcohol evidently causes a coagulation of substances present in the scales and possibly in the bulb axis. To hasten the penetration of the material by the fixing agent, the object as soon as dropped into the acid was placed in a chamber from which the air was exhausted until the large size of the bubbles arising from the cut surface of the tissue indicated a fair approach to a vacuum. The air was allowed to replace the partial vacuum very slowly without outside aid. The use of the exhaust chamber was also resorted to with every change of alcohol, alcohol and ether, and ether. The fixation was practically perfect owing to more immediate and entire penetration. It did not, however, secure even moder- ately good fixation of incipient roots which were wholly within the bulb axis. Prior to killing and fixation, the bulb was trimmed down to about six scales, if the purpose was merely the examination of the youngest portions. Thorough fixation was thus more easily secured and the vegetative point was not crushed by the heavier parts. Washing was carried on in water for two days and completed in the low grades of alcohol, such as 20-70 per cent. The general celloidin method was followed from here on until the actual point for embedding was reached. At this point in the work, the object, after the removal of the superfluous celloidin, was hardened in 70 per cent alcohol and then in chloroform for two hours. It was transferred from the chloroform to 85 per cent alcohol. The usual paraffin method of imbedding was now followed (5)- Material prepared in this way sectioned with unusual smoothness and ease. The rotary microtome was used in all paraffin work except where bulbs or pieces of material were too large for the section block. Such large pieces of material were mounted on blocks of wood, trimmed out underneath until small enough to be placed in the jaws of the sliding microtome clamp. In handling individual paraffin sections as large as, or larger than, two inches square, a vessel of warm water was kept at hand. The sections placed carefully on the surface of the warm water uncurled readily, with no danger of cracking. It is necessary to see that the water is not CuurcH: THE BULB IN CooPERIA DRUMMONDII 341 too warm at first. A few changes of the water to warmer or reheating of the water by placing the container in another vessel of warmer water will be found useful. MATURE EMBRYO The structure of the mature embryo as seen in relief exhibits a scarcely protruding pocket, which protects the leaf-primordium and the vegetative point by surrounding their outer surface as they lie against the base of the cotyledon. The sheath-like base of the cotyledon has no vascular system and is very simple in structure. The long spindle-shaped cotyledon is succulent and projects into the mass of endosperm, absorbing food for the de- veloping embryo by means of a layer of thin-walled parencyhma- tous cells running parallel with the longitudinal axis of the embryo. The suspensor, which is composed of either one or two cells, still holds the embryo in position at maturity. The number and size of the leaf-primordia, developed at the time the seed is ripe, are not at all constant but depend on the individual embryo. No cases have been noted, however, where the first leaf-primordium had not made its appearance at this period. The primary root is protected by a well-developed root cap even as early as the maturity of the seed. Wordsell (35) has recently given to botany a comprehensive paper on the monocotyledonous embryo, where extensive dis- cussions and a bibliography relating to the cotyledon may be found. His opinion is exactly that of Celakovsky, namely, that the cotyledon is an equivalent of the capsule of the moss sporo- gonium, just as the hypocotyl is of the seta. Goebel (8) states in regard to the cotyledon of the seed plants: ‘‘I need only say here that the cotyledons, which so frequently differ in form from the foliage leaves, are merely arrested forms of these, the arrest being sometimes permanent, sometimes transient.” Lyon also has con- tributed (21) to discussions of the true morphological position of the cotyledon. At best a discussion becomes one of terminology unless combined with detailed anatomical observations on serial sections of embryos at successive ages of development. The present writer is inclined to agree with Lyon, stating that as the cotyledon of Cooperia Drummondii does not appear to arise as an 342 CHURCH: THE BULB IN COOPERIA DRUMMONDII exogenous lateral outgrowth upon the growing point of a stem as do the later foliar structures, therefore, said cotyledon, occurring in the mature seed plant at the base of the primary stem, is more closely related to the nursing foot of the bryophytes. The succu- lent cotyledon of the young plant of Cooperia Drummond in anatomical structure and size is well adapted to serve as an absorption organ. SPROUTING OF THE SEED The ripe seed of Cooperia Drummondii was described by Herbert (13) as having a brittle, shiny black seed coat and in- clined to be wedge-like in shape. The seed coats are slightly torn at the time of sprouting as the root pushes through the very small micropyle, but they are never split apart into halves. On February 20, 1914, the seeds from a ripe pod were immediately placed upon wet filter paper on the sides of a moist chamber at ordinary room temperature. Three days later—February 23— the seeds had sprouted and the roots on the seedlings averaged 4 mm. in length; on February 25 the root length was Io-12 mm., while at thirty days of age the root length was 65 mm. If, how- ever, the seeds were well dried and allowed to remain dry for several months at the temperature of the laboratory they did not sprout so readily. The future root and shoot of the embryo are pushed out of the seed coats by a lengthening of the cotyledon, as well as by the growth of the root- and shoot-regions. Solms-Laubach (30) notes a similar condition in Heterachtia. The portion of the cotyledon between the micropyle region of the seed coats and the surface of the soil becomes a brilliant green, indicating the formation and presence of chlorophyll here. The cotyledon therefore functions in part as a leaf, aiding the leaf blades in the photosynthetic processes of the plant’s activities. The leaf grows upward to the light above the ground, while the cotyledonary sheath serves as a protecting sheath in the same © fashion as the encircling leaf base of any subsequent leaf serves as a sheath for the next younger leaf (Fic. 4). The region bounded by the root, the cotyledon and the plumule was designated by Richard (29) in 1808 as the ‘“‘tigelle,” corresponding as he said to the Latin term “‘cauliculus.”’ The ‘‘tigelle’’ he defines as CuHuRCH: THE BULB IN COOPERIA DRUMMONDII 343 merely a “‘prolongement”’ of the ‘“‘radicule,’’ explaining that ‘‘la Tigelle (Cauliculus) se confond d’une part avec la Radicule dont elle n’est qu’un prolongement et se termine de I’autre a la base de la cavité cotylédonaire.” Jussieu (19) agrees with Richard and also connects the ‘‘tigelle’’ up with the term ‘‘carnode” of Cassini and with Brongiart’s ‘‘mealy body” found in Lemna, although the describer says the latter even may be a cotyledon. At this region designated as the ‘‘tigelle” the writer noticed in Cooperia Drummondu that the cells divide in planes at variance with the tissues of the shoot above it and the root below it (Fic. to, h). Here they divide in both the horizontal and tangential planes with little lengthening and practically no broadening, while in the shoot and root the cells broaden relatively rapidly and also elongate greatly. The area enclosing the axis of the first leaves and the growing point (the plumule, the ‘‘gemmule”’ of early writers) is potentially cotyledonary. Lyon (21) states that the maximum development in this area takes place at the point or points that are in the most favorable position to function, and that in monocotyledons the maximum development occurs only on one side. And, further, that cotyledons do not arise as exogenous lateral outgrowths upon the growing point of a stem as do all later foliar structures. The gross anatomy of Cooperia Drummondii indicates that there is a point of maximum development and also one of decidedly mini- mum development in the cotyledonary region. Also the anatomy of the developing embryo shows us that the cotyledon does not develop from the vegetative point, nor does the cotyledonary sheath. The function of each of these portions is peculiar to itself, the maximum point of development elongating into a haustorial organ with the function of absorbing food for the germinating embryo, the point of minimum development not elongating but serving as a protective covering or a sheath to the primordia or a pocket in which the gemmule is deposited, accord- ing to the older writers, asa ballisina cup. The gemmule is thus defined by Richard (29): ‘‘La Gemmule (Gemmula) est la petit corps simple ou composé qui nait ou du fond de la cavité coty- lédonaire, qui le referme étroitment.’”’ The early scientists noticed that their ‘‘gemmule’—the vegetative point plus an 344 CHURCH: THE BULB IN COOPERIA DRUMMONDII incipient leaf or so—could be sighted through the ‘‘fente.”’ As Jussieu (19) states: ‘‘. .-. et ce ne fut qu’apres des recherches répétées et minutieuses que je parvins 4 m’assurer que le cotyledon présentait en effect une petite fente vers sa base.’”’ They specu- lated as to whether or not this gemmule were naturally_detachable from the rest of the structure. Brown (3) speculates in his work and merely shoulders enough responsibility to say that the gemmule in bulbiform amaryllids can be seen and that it escapes through this opening, “petite fente.”’ A careful microscopic examination of slides of a five-day-old Cooperia Drummondii seedling reveals cell structure indicating that root contraction has already begun. Such a condition has been noted by Hallstrém-Helsinke (9) in Urginea maritima, by Rimbach (23-28) and by De Vries (34) among the dicotyledons and also in Hyacinthus orientalis. Rimbach appears to be the best-known investigator volunteering a hypothesis of the relation of this contraction to root tissues. It is the intention of the writer to discuss his deductions and those of others later. THE SEEDLING A five-day-old plant of Cooperia Drummondii (Fic. 1) shows no outward indication of bulb formation. However, the region h does limit the territory of the root and of the shoot. The primary root is a tap root, in appearance stout, tapering slightly downward, and colorless. The shoot consists of the first leaf blade or lamina, 1,; the cotyledon, c (from which in the preparation drawn portions of the seed coats and of the endosperm tissue, e, had been removed) ; and the cotyledonary sheath. Acentral longitudinal section of this individual (Fic. 2), under the low power of the microscope, shows in addition the sheath, 0, of the first leaf and the primordia of the second leaf, J, and 62, together with the vegetative point, pm. The stippled strips represent approximately the space occupied by the vascular traces, no branch of which goes to the cotyledonary sheath, cs. Fic. 3 takes to one side the youngest area and repre- sents exactly its cellular structure. In this region, surrounded by the first leaf of the young plant, the cells are actively dividing. The cells at 2, and 2 indicate not only that the second leaf has already become differentiated from the primordial meristem, pm, CHURCH: THE BULB IN COOPERIA DRUMMONDII 345 but also that as a whole the meristematic tissue of this second leaf occupies an area placed diagonally to the longitudinal axis and surrounding the primordial meristem like a collar. The primordial meristem is-often definitely composed of paired cells (Fics. 3 and 6, pm), as Carano (4) has recorded for Yucca. Each new leaf arises from a group of subepidermal cells (Barenetskey, 2), which become meristematic and divide tangentially. Thus a new tissue is formed which, as it pushes out, becomes a future leaf. As the first leaf is thrust out far from the cotyledonary sheath into the air (FIG. 4), the cotyledon elongates and curves downward until it is parallel with itself and the vertical shoot, thus passing through an angle of 180 degrees (FIG. 4). THE TWO-MONTH-OLD PLANT When the food supply has been exhausted or drawn upon to the necessary extent the cotyledon, having no further need to serve as a haustorial organ, becomes shriveled up and dies (Fics. 5, and 14). With this dying off of the cotyledon the dry, hard seed coats with any residue of endosperm drop to the ground or at least cease active relations-with the ~\ cotyledon. \ The second leaf blade usually reaches daylight be- tween the fifty-fourth and the sixty-first day (Fic. 14). As the first leaf sheath is surrounded by the basal sheath of the cotyledon, so this second leaf sheath is surrounded by the basal sheath of the first leaf, in respect to which it is distichously placed. In Tulipa the leaf lamina, as is well known, does not always develop. Henry (11) makes note of this fact in his discourse, ‘‘Beitrage zur Kenntniss der Laub- knospen.”” Even in young bulbs of Tulipa the aborted scale-like leaf alternates with a true leaf bearing a green lamina. In Cooperia Drummondii, however, the lamina always develops and as a result we have in this member of the Amaryllis family no mere sheath- or scale-leaves. The lamina of the leaf of Cooperia Drummondii appears to develop simultane- ously with its respective base. The incipient bulb is now readily recognizable (Fics. 5 and 14). Its outer and only scale is formed bulb; scale = 5 cm. 346 CHURCH: THE BULB IN COOPERIA DRUMMONDII of the cotyledonary sheath. The bulge shown (Fic. 14) is due to a young root. This first adventitious root, 7, originates (Fic. 8) at the node of the second foliar leaf and, growing downward, cuts its way through the first leaf and the cotylar sheath. A slightly older root with a well differentiated root cap and primary tissues (dermatogen, periblem, and pleurome) is shown in Fic. 18. This seedling is from the same planting as that of Fic. 8. The young root arising endogenously in the bulb- or stem-axis will, together with similar companions, replace the tap or primary root. The primary root is still active at this time. The secondary roots, which become functionally the principal roots of Cooperia Drummondti, are adventitious roots, derived from an active cambium tissue below and to one side of the growing point. They force their way through the tissue of the bulb-axis and form at first roots to one side of the early dying primary root and later a complete ring of such roots. The roots of Cooperia Drummondii have root hairs. The vascular bundles which develop in the secondary roots anastomose with the bundles already formed in the stem-axis. The complete working out of the origin and arrangement of the vascular traces in the stem-axis of Cooperia Drummondti would be very difficult, so complicated is the anasto- mosis. Both the primary root and the secondary roots show a wrinkling on their surface which is connected with the drawing down of the bulb into the soil by contraction of said roots. A bulb may be drawn down in this fashion to a depth of over seven inches. The puckered or wrinkled condition of the primary roots (Fics. 8, 15, 17 and 18) is due to contraction of at least some of the tissues. This phenomenon occurs in the adventitious roots also. Its hypothetical mechanics will be discussed later. The outlay of root tissue in FIG. 17 has a narrow slice marked off. This slice has been divided into three sections—1, 2 and 3. If this narrow strip of root is studied with the high power of the microscope we have under observation cellular tissues similar to those of FI. - 19, where I represents the vascular region; 2, the parenchyma; and 3, the cork and other dead tissues. Evidently the root con- traction has entirely damaged only the outer cells, which are now dead tissue. The inner cells have in some way accommddated themselves in part to the strain. The trace is still intact although CHURCH: THE BULB IN COOPERIA DRUMMONDII 347 its individual cells are irregular in diameter, presumably from the effects of some pressure. The parenchyma cells outside the trace are still equipped with cytoplasm and nuclei, although some force has pushed them by each other and flattened their nuclei. ROOT CONTRACTION The first reference in the literature to root contraction, accord- ing to De Vries (34), is that of Tittmann (31) in Flora for 1819. Under ‘‘Wurzelbildung”’ Tittmann describes the contraction of the roots in Daucus Carota as follows: ‘‘Der Stengel wird dicker und kiirzer, zieht sich gleichsam in die Erde hinein, oder wird von derselben angezogen, und man findet dann nach einiger Zeit die langen Saamenblitter dicht auf der Erde ausgebreitet. Unter- sucht man in dieser Periode die Wurzel, so hat sich die Granze zwischen ihr und dem Stengel, der sich nun auch in der Erde befindet, ganz verloren und ausgeglichen.” De Vries :(34) himself worked principally with dicotyledons, experimenting with eighteen species of plants. He studied the effects of exposure to air, to water and to salt solutions on strips, separate pieces of definite tissues, and single cells of roots. He measured changes in length, width, and volume of tissues and cells. He considered roots from plants still bearing succulent cotyledons to those two years of age. He studied young roots and roots two years old. Each experiment is considered in itself and in the light of previous experiments, until at last he concludes that root contraction is not due to intake of water but to changes in turgor and remarks that the thoughtful reader will ask, “durch welche Ursachen die ungleiche Dehnbarkeit der Zellhaute selbst bedingt wird.” Rimbach discusses root contraction as it occurs in Colchicum autumnale (26), Arum maculatum (28), and Allium ursinum (27). He found that, in the case of naturally deep-set bulbs of Allium ursinum,a depth of 10-15 cm. was attained after several years by the contraction of the roots. In explanation as to how contraction in roots comes about he states, that, while the root tip holds firmly to the earth particles, the endodermal cells contract longitudinally as a result of a tendency to lengthen radially and tangentially, which results in the contraction of the root. The bulb yields to 348 CuHuuRCH: THE BULB IN COOPERIA DRUMMONDII this pull, which results from the contraction of the roots, and is gradually placed more deeply in the ground with each new set of roots. Even the upward development of the shoot, where the lowest and oldest scales are each formed at a slightly higher level, does not counteract entirely this forced downward movement of the bulb. Therefore the bulb is continually placed more deeply in the soil. In Arum maculatum Rimbach (28) explains that the active portion of the root consists of the cells directly below the epidermis and the ‘‘hypodermal layers.’’ These cells shorten in their length as much as one half and their radial walls lengthen somewhat. The vascular bundles and their accompanying cells are not active. However, the latter respond sufficiently to the vertical pressure caused by the shortening of the parenchymatous" cells just out- side them to show definite wrinklings along their walls, and the tracheal tubes themselves shorten between the rings. Rimbach’s careful observations and well-chosen biological experiments are not supported by his theoretical conclusions. Because he has accurately noted that endodermal cells lengthen radially, he proceeds to state that they shorten longitudinally _and that therefore the whole root contracts, with the result that the bulb is set more deeply into the ground. All other tissues are passive, yet they must be shortened in the shortening of the whole root. Without sufficient morphological proof Rimbach claims that the outer or cork cells are crushed and killed, and that the vascular cells are thickened. The outer cells are crushed doubtless, but do they die first or are they killed by the crushing? His further con- clusion that the vascular cells thicken may be true, but where is his proof? To return to the active or endodermal cells, how can we accept the statement that because a cell lengthens in one direction therefore it must necessarily shorten in another? Vol- ume ‘being constant and cell wall composition and tension being similar at every point this would be true; but Rimbach did not investigate these points. : Rimbach states that ripples may be seen on the root surface even to the root-ends with the naked eye. Microscopic slides have proved to the writer that the process of root contraction is well under way in a five-day-old seedling of Cooperia Drummondit CHURCH: THE BULB IN COOPERIA DRUMMONDII 349 at points not visible to the naked eye. The ripples in individual cells are not confined to radially placed cell walls, as Rimbach finds them to be in the species which he investigated. On the contrary they may be found on walls running in any direction. . In some stages they are so fine as to be distinguishable only with the oil immersion objective, but always numerous cells with rippled walls may with care be detected in the parenchyma cells of the root with a combination of 10 ocular and 8 objective. The warped and altogether disorganized condition of the epi- dermis and ‘‘hypodermal’”’ layers in the contracted roots of Cooperia Drummondii may be comprehended, if we study the tissues represented by the blackened areas in FIG. 17, section I. It seems safe to accept these facts: (1) roots do shorten; (2) the parenchymatous tissues of the root are the seat of this activity; (3) the cork and the vascular trace are passive; (4) the cork is ultimately crushed; (5) there is a region where one can see wrink- lings and measure shortening, a second region where no wrinklings are visible yet where one can measure shortening, and an un- changed region (Rimbach); (6) in dicotyledons the trace becomes visibly curved inward and outward in a wavy fashion, while in monocotyledons the vascular bundles remain practically straight (De Vries). : What remains to be determined constitutes a problem of botanical research as yet unsolved. We may hope that some worker with an interest in morphology as well as physiology may master this problem by a study of serial sections of young roots and a consideration of the physical relation of turgor and biochemical alterations in the protoplast and cell membrane. All the roots of a plant may not shorten equally. Phaedranassa chloracea, according to Rimbach (25), has a main root which shows no shortening. In other cases none of the roots shorten. Tulipa, according to Déring (6), is a case in point. The bulb here would not be pulled down since its roots spread out almost parallel with the surface of the soil. THE SCALES The scales, that is, the leaf sheaths, which constitute the greater portion of the bulb, are differentiated into certain tissues, represented diagrammatically in Fic. 7. Ascale has an epidermal 350 Cnurcu: THE BULB IN COOPERIA DRUMMONDII layer on each surface. The several rows of cells directly below the outer epidermis contain starch, while the cells below the inner epi- dermis do not. The starch-filled cells measure 59-158 uv in length and 59-69 » in width, with their greatest dimension running parallel to the longitudinal axis of the bulb. The vast majority of such cells average 158x59u. The cells between the cells containing starch and the accompanying cells contain raphides in the first seven or so scales, but rarely in the older scales. The amount of starch also is less in the older scales than in the more actively functioning portions of the bulb. There appear to be, however, sphaero-crystals stored in the older scales. There is noticeably less starch and fewer raphides in the scales immediately rae , : base bt Fic. 2. Fic. 3. Fic Diagram of Cooperia Drummondii as it would appear if the internodes devtaped I, lamina of a leaf; b, basal sheath; 0, leaf subtending flower; f, flower; n the lower part of the figure a central longitudinal cyon is represented, showing pincigake the relation of the basal sheath to the main axis; the relation of the flower and its subtending leaf to the main axis is problematic with the evidence at hand Fic. 3. Diagrammatic semi-transparency of the youngest portion of the bulb of Cooperia Drummondii: pm, the primordial area, below which are the primordia of the youngest or uth leaf; gree: older leaves are marked Int» In—2, and Ins; their respective bases, Bn1, ba—2 and : CHuRCH: THE BULB IN CoOoPERIA DRUMMONDII 351 following the flower stalk which has most recently died, and finally neither starch nor raphides are to be noticed in the outermost and oldest scales. The leaf bases in the youngest portion of the bulb— the only portion within the bulb itself where the leaf base in cross section is scarcely continuous with the lamina—have no starch stored in them. The leaf accompanying the flower is supplied with starch even when it is very young. Young roots that have not yet pierced the bulb-axis contain raphides. The starch grains and raphides, therefore, are most prevalent in the younger scales. Here cells with raphides may be found among the tissues contain- ing starch or scattered between the starch-bearing cells and the accompanying cells, but most commonly they are found in rows of cells. An individual cell in such a row is about twice as long as a starch-bearing cell or often still longer. The sheath or bundle of crystals practically fills the cell containing it. Measurements of the raphide-bearing cells show that their dimensions measure 277-475 wu in length and 79-99 uw in breadth. The remainder of a scale is composed of vascular bundles and their accompanying parenchyma, the cells of which are intermediate in length between the cells filled with starch and those with raphides. The dimen- sions of these rather simple cells range from 178x119» to 297 x 69 b. Miss Menz (21) has made observations similar to the above in connection with Amaryllis, Zephyranthes, Sternbergia, Crinum and Allium, finding in the scales of these genera starch, raphides of calcium oxalate and also a latex-like or slimy substance stored as reserve material. DEFINITION OF A BULB Bulbs, according to Irmisch (16), have been looked upon as roots, underground shoots, and downward growing shoots, while Irmisch himself concludes with the emphatic statement that a bulb is a bulb—an organ peculiar to certain plants and distinctive in itself. A bulb is a shoot in which the internodes have not developed or have developed only to a small extent (TEXT FIGS. I, 4 and 5). The peripheral layers of the bulb consist of the bases of the foliar sheaths. As there are no internodes in the shoot of Cooperia Drummondit its foliar sheaths never can form an exterior covering ooe CHURCH: THE BULB IN COOPERIA DRUMMONDII to internodes but instead surround the next younger sheath, as they themselves are surrounded by an older sheath. If a shoot, developed as is usual in herbaceous plants, corre- sponds to a telescope drawn out for observation of the stars, a bulb finds its correspondential in the same telescope pushed in— i.e. a bulb is a foreshortened or a ‘‘telescoped”’ shoot (TEXT FIG. 2). A bulb of Cooperia Drummondii is probably never raised above the earth’s surface. It is constantly being set deeper into the ground. The oldest tissue composing the base of the truncate bulb axis is continually sloughing off, while the youngest tissue is continually formed at a point farther and farther away from the point where the primary root originated. This setting of the bulb deeply into the ground is due to root contraction, which has been explained as far as is possible with our present knowledge before us. An old bulb of Cooperia Drummondii may ultimately bear a neck 15-18 cm. in length. *‘OFFSETS”’ » arise between the mature scales as protrusions. When condi- tions fostering vegetative pro- The ‘‘offsets”’ or “‘splits pagation occur, the ‘‘splits”’ develop (Fics. 12 and 13; TEXT FIGS. 4 and 5) between the scales or leaf sheaths and break their way through the outer and older bulb scales, in an effort to reach the soil. They develop on alternate sides of the axis and frequent- ly occur between every five Fic. 4. “Offset’’ readjusting itself after eS nae pais iatCe i an unfavorable start; roots diagrammatic, to be considerable variation Le. cpg due to contraction omitted; in regard to this matter of Fic. 5. Sis. arrangement. Fic. te iia propagation displacin; H bay © Regn roots and leaves diagrammatic; enry (11) notes tha Ga gea arvensis, G. stenopetala and CuurRCcH: THE BULB IN COOPERIA DRUMMONDII 353 Tulipa sylvestris show secondary bulb formation from the original bulb developed to extreme abnormality, yet every bulb so formed is always a leafy shoot. That is, such a bulb is a secondary or lateral shoot, or a terminal shoot which will eventually separate itself from the growth of previous years. Vigier (33) relates a few casual but interesting experiments with bulbil formation in cuttings of Lillium candidum. Bulbils did not form on the cut- tings, unless the end of the cutting and thus the terminal bud were injured accidentally or intentionally. Nipping the terminal bud of horticultural plants to produce lateral branching is a common horticultural practice. Regarded as lateral branches these bulbils are unique only in their being vegetative outgrowths which can of themselves reproduce the plant vegetatively, where the usual secondary branch can not do so without the gardener’s aid. The bulbils of L. candidum, formed in the light above ground, were in Vigier’s experiments green with purple spots, while those formed on the part of the cutting under the soil were white. There can be no proof brought up against the statement that there is pri- marily no difference between these bulbils of L. candidum formed below or above the ground and the offsets, splits, or brood-bulbs of Cooperia Drummondii, for instance. There is no difference even between the bulbils formed in Vigier’s experiments: whether the said bulbils have green and purple pigment or are colorless; whether they are formed above or below the surface of the ground; or whether we are considering a bulbil or what is recognized commonly as a lateral shoot. In each case we have a shoot— a structure which is still a shoot none the less, whether it be a main or lateral shoot, or an artifically aborted shoot, or (as is the bulb) a naturally aborted shoot, in which the internodes have elongated little if at all. THE MATURE BULB Irmisch (16, 17) describes the structure of immature and mature. bulbs of Amaryllis formosissima (now Sprekelia for- mosissima). His work presents to us a most careful re- search, recorded by the observer's skilfully executed drawings. While further investigation since 1850-60 causes us to feel certain in regard to points which are here discussed rather lengthily and 354 CHURCH: THE BULB IN COOPERIA DRUMMONDII left undecided even then, and to disagree with some conclusions which Irmisch does draw, yet the ground is covered so minutely and the observations are presented so carefully that the work is worthy of our close consideration. According to Irmisch Amaryl- lis formosissima has in the mature bulb, considered first of all from the outside toward the inside, three or four leaves with Fic. 6. Central longitudinal section of youngest portion of mature bulb; camera-lucida drawing of portion cut off between dotted lines at center of PLATE 16, IG 17: black areas represent the flower and its subtending leaf; stippled areas indicate the portions of the leaves which becomes laminae; cross-hatched areas show the leaf bases; plain areas are primordial tissue; it is presumed that the oldest leaf shown is the sixth leaf—an arbitrary choice. Scale = 0.1 mm. closed bases. Such a leaf he terms “Schale,’’ which literally translated into English means ‘“‘skin” or “hull” and again has the derived meaning, ‘“‘bowl.” This seems a most apt scientific term, for one can readily understand that if the center of a bulb is removed, leaving only a few outer scales there is left merely a few skins, bowl-like in shape. Following the three or four leaves with closed bases, there is one leaf with an open base which sub- tends a flower scape. Such a leaf is termed by Irmisch ‘‘Schuppe,” which means “‘scale”’ (as of a fish) or ‘‘shovel” in English. Apart from the blade the open based leaf accompanying the flower of Cooperia Drummondii is shaped decidedly like a scale and with CuHuRCH: THE BULB IN COOPERIA DRUMMONDII 355 the blade attached becomes the shovel. In the case of Amaryllis formosissima there is in the axis of this open based leaf an axillary shoot or secondary branch whose terminal bud becomes a flower. This flowering scape has two leaves which are not normally devel- 9) Rey) oped. The flowering branch is followed by J three or four leaves with closed bases similar to the first group, but always growing smal- ler and more rudimentary as the primordia PIS: 7 Cross-sec- : tion diagrams after Ir- are approached. An old bulb might have isch: a, of Amaryllis within it four flowers with their accompany- formossissima; b, of Ga- ing ‘‘Schuppen”’ and thirteen to fifteen scales. [@”/#™s ot Leucojum ; 2 : in Galanthus and Leu- Usually all laminae are dead at flowering com Irmisch found times, Irmisch states, and the new leaves alternation between come always from that portion of the bulb “Schuppe” and ened ae ae Al Th ; 1 “Schale’’; in Cooperia Inside of the last wer = occasional p,.immondii and open leaf does not alternate with the closed Amaryllis formossisima leaves, so that diagrammatically Irmisch’s in- en-based leaf te fat: iit 3 ted as.in TEXT ‘subtending the flower oe eee phe ecteiasrem ta 4 - does not alternate with FIG. 7,a. In Galanthus and Leucojum, how- _ the closed leaves. ever, Irmisch does find alternation between Schuppe and Schale (TExtT FIG. 7, 6). The present writer has ob- served for Cooperia Drummondit the same relation of the two types of leaves as Irmisch did for Amaryllis formosissima, which again relates Cooperia closely to Amaryllis. In Cooperia Drummondit, however, the axillary bud develops only one bract besides the flower from its terminal primordium. This is represented dia- grammatically by TEXT FIGs. 2 and 9. The shovel-like leaf accompanying the flower is open at the base in Cooperia Drummondii whether the flower matures or not, but in Narcissus and Leucojum the base of the same type of leaf becomes closed if the flower never develops. In Cooperia Drum- mondii this leaf may belong to the lateral axis which gives rise to the spathe-like bract and the flower. Cooperia Drummondit has no stipules unless the base opposite the lamina was once formed by the fusion of such (TEXT FIG. 8). It seems, however, as if the base were here derived from a slight and all-encircling outgrowth of the primordial meristem of each individual leaf primordium, Sy 3 pos i" o 356 CHURCH: THE BULB IN COOPERIA DRUMMONDII The mature plant of Cooperia Drummondii has a subglobose bulb, increasing in size with age. The outermost scales have become thin, dry, brown membranes, whose venation is readily n-2 re aris a eet “syne | 4 G. 8. Central portion of cai showing immature flower; presuming there This is in accordance with an immature flower, and 0, its subtend- ing leaf with an open base. Scale = 0.15 mm. seen. As one goes inward, tearing away the scales, those with dried edges are suc- ceeded by fresher and thicker scales. Every third scale is separated from the next younger or older scale, according as we work inward or outward, by a scale accompanying a flower peduncle, and this arrangement is repeated in accord with the age of the plant, the parts always grow- ing smaller and more rudimentary. An old bulb may contain the vascular traces and dried stalks of numerous past or un- developed flowers, and immature flowers for three successive flowerings (see TEXT FIG. 9). Irmisch (16) gives a type classification of bulbs supplemented with examples, Latin designations and a set of symbols. The first type is designated as ‘‘Gagea-Arten,” and the symbol expression corresponding is G+Z, where G is equivalent to ‘‘ Gegen- wart’’ and Z equals ‘‘ Zukommen,” next year’s development. Therefore all bulbs similar to Gagea consist of G (present, active portions) and Z (tissues capable of repeating the life story next year). symbols 1V+G-+Z. Another type, e.g. Galanthus nivalis, is represented by the IV is equivalent to parts of the preceding year which are now present as ‘‘ Nahrblatter” or sheaths containing stored material. Yet another type, Amaryllis formosissima, may be represented by 2V + G + Z, where the parts of two previous years still remain. Cooperia Drummondiui might be represented by nV + G + Z where n varies largely with environment as ner as with the age of the plant. The following scheme, illustrating a mature bulb of Cooperia Drummondit, may likewise be considered in connection with TEXT FIGS. 2, 6, 8 and 9: CHURCH: THE BULB IN COOPERIA DRUMMONDII 357 MAIN AXIS OF BULB Three leaves with closed bases (Schalen). Lateral axis: One leaf with open base (Schuppe), position problematic; One bract subtending flower; One flower (terminating lateral axis). Three leaves with closed bases. Etc. to vegetative point. Galanthus and Leucojum also have open-base leaves succeeded by a flower stalk, but their open-base leaves alternate with the closed-base leaves. The open leaves do not alternate with the Fic. 9. Diagrammatic longitudinal section of mature bulb showing the main axis drawn in thin lines: the flowering axis (?) in heavier lines; the section between the dotted lines may be seen enlarged in TEXT FIG. 6. closed-base leaves in either Amaryllis formosissima or Cooperia Drummondii, but with each other,(TEXT FIG. 7, @ and b). The 358 CHURCH: THE BULB IN COOPERIA DRUMMONDII flower peduncle, terminal in respect to the lateral axis, is subtended by a spathe-like bract. CONCLUSIONS 1. The mature seed of Cooperia Drummondii germinates with little difficulty. 2. A five-day-old seedling shows a tap root, hypocotyl, and one young leaf blade. 3. A ten-day-old seedling exteriorly shows no indication of bulb formation. 4. Seedlings by their fortieth to fiftieth day of growth have developed a typical bulb formed from the closed bases of alternate leaves. 5. The bulb is set deep into the ground by a contraction of the ' roots, which begins as early as the fifth day. 6. Amature bulb is composed of a thickened axis (with morpho- logically aborted internodes), from which develop (in a down- ward direction) adventitious roots, piercing the outer edge of the axis in a circle, and (in an upward direction) scales (leaf bases), flower scapes (each with its accompanying open base leaf), and leaves. 7. The flower scape and the leaf accompanying it alternate with the next younger corresponding leaf and scale; they arise between every third and fourth scale. : | 8. Vegetative buds of a type called ‘‘offsets’’ among horti- culturists develop between the scales whenever conditions of environment are unfavorable to flowering. 9. A bulb is a foreshortened vegetative shoot which appears aborted, because the internodes do not develop, while the nodes are thickened by irregular and considerable multiplication of cells. This paper was presented in 1918, in partial fulfillment of the requirements for the degree of Doctor of Philosophy at Brown University. The writer begs to acknowledge the assistance given by Professor Harlan H. York of the Botanical Department. LITERATURE CITED 1. Baillon, M. H. Sur le développement et la germination des bulbi- formes des Amaryllidées. Bull. Soc. Linn. Paris 1: 4-5. 1874- Oo as on caer oO Ke) ° iS) Lael ~ CHURCH: THE BULB IN CooPERIA DRUMMONDII 359 . Baranetzky, M. J. Sur le développement des pointes végétatifs i Hil. des tiges chez les Monocotylédones. Ann. Sci. Nat. Bot. V 3: 311-365. pls. 14-16. 1897 - Brown, R. Prodromus Florae Novae Hollandiae et insulae Van Diemen. London. 1810. . Carano, E. Sulle formazioni secondarie nel catle delle Monccatile. doni. Ann. di Bot. 8: 1-42. pls. 1-4. 1910. . Church, M. B. Celloidin-paraffin methods. Science II. 47: 640. 1918. Doring, E. Das Leben der Tulpe. Sondershausen. 1910. . Falkenberg, P. Vergleichende Untersuchungen iiber den Bau der Vegetationsorgane der Monocotyledonen. Stuttgart. 1876. . Goebel, K. Organography of plants. English translation. Ox- ford. 1900-0 . Hallstrém-Helsinki, K. H. Ueber die Keimung von Urginea maritima Baker. Schweiz. Wochenschr. fiir Chem. und Pharm. 49: 89-91. 6 figs. I911. . Hanstein, J. Entwicklung des Keimes der Monokotyle und Dicotyle. Bot. Abhandl. 1: 1-112. pls. r-1r8. 1870 . Henry, A. Beitrage zur Kenntniss der Laubknospen. Dritte Abtheilung. Nova Acta Acad. Caes. Leop. Carol. 21: 277-292. pls. 16, 17. 1844. . Herbert, W. An appendix (to the Botanical Register), containing a treatise on bulbous roots. London. 1821. Amaryllidaceae; preceded by an attempt to arrange the monocotyledonous orders, and followed by a treatise on cross- bred vegetables and supplement. London. 1837. Highfield, E.G. The structure of a garlic bulb. Naturalist 1912: 331-3362 1012; Howard, W. L. An experimental study of the rest period in plants. The summer rest of bulbs and herbaceous perennials. Second report. Missouri Agri. Exp. Sta. Research Bull. 15: 1-25. figs. 1-8. Ap 1915. Irmisch, T. Zur Morphologie der monokotylischen Knollen- und Zwiebelgewachse. Berlin. 1850. Notes on the development of bulbs and tubers (abridged from the German original). Jour. Roy. Hort. Soc. London 8: QI-124, 207~—221. 38 figs. 1853. Beitrage zur Morphologie der monokotylischen Gewachse. 1. Heft. Amaryllideen. Halle. 1860. Jussieu, A.de. Mémoire sur les embryons monocotylédons. Ann. Sci. Nat. Bot. II. 11: 341-361. pl. 17. 1839. 360 CHURCH: THE BULB IN COOPERIA DRUMMONDII 20. Koch, K. Lilienpflanzen und Zwiebelbildung [in Beitrage zu einer Flora des Orientes]. Linnaea 22: 213-219. pl. 2. 1849. . Lyon, H. L. The phylogeny of the cotyledon. Postelsia Igor: 57-86. 1901. . Menz, J. Beitrige zur vergleichenden Anatomie der Gattung AI- lium nebst einigen Bemerkungen iiber die anatomischen Bezie- hungen zwischen Allioideae und Amaryllidoideae. Sitzungsb. Kais. Akad. Wissens. Wien 119: 475-533. pl. 1-3 +f. I-16: 1910 . Rimbach, A. Ueber die Ursache der Zellhautwellung in der Endo- dermis der Wurzeln. Ber. Deuts. Bot. Ges. 11: 94-113. 1893. Zur Biologie der Pflanzen mit unterirdischen Spross. Ibid. 13: 141-155. pl. 14. 1895. Ueber die Tieflage unterirdisch ausdauernder Pflanzen, Ibid. 14: 164-168. 1896. Ueber die Lebensweise des Arum maculatum. Ibid. 15: 178-182. pl. 5. 1897. Lebensverhéltnisse des Allium ursinum. Ibid. 15: 248- 252. pl. 8. 1897. Biologische Beobachtungen an Colchicum autumnale. Ibid. 15: 298-302. pl. r2. 1897. . Richard, L. C. Demonstrations botanique, ou Analyse du fruit considéré en géneral. Paris. 1808. ' . Solms-Laubach, H. Ueber monocotyle Embryonen mit scheitel- biirtigem Vegetationspunkt. Bot. Zeit. 36: 65-74, 81-93. pl. 4. 1878. . Tittman, D. J. A. Botanisch-karpologische Bemerkungen. Flora 2: 651-666, 667-671. 1819. . Tschirch,A. Physiologische Studien iiber die Samen, uatinaontiese ; der Saugorgane derselben. Ann. Jard. Bot. Buitenzorg 9: 143- 183. pl. 20-25. 1891. . Vigier, A. Formation de bulbilles sur le lis blanc. Rev. Hort. 78: 406. 1906 906. . Vries, H.de Ueber die Kontraction der Wurzeln. Landw. Jahrb. 9: 37-80. 5 figs. 1880. - Wordsell, W. C. The morphology of the monocotyledenous em- bryo and of that of the grass in particular. Ann. Bot. 30: 509- 524. f. I-10. 1916. . Worsley, A. Germination of Amaryllideae. Jour. Roy. Hort. Soc. 28: 420-423. f. 10-112. 1903-1904. CHURCH: THE BULB IN COOPERIA DRUMMONDII 361 Explanation of plates 14-16 The following remarks refer also to the text figures. FIGS. 7, 12 and 13 in the plates and TExT FIGS. I, 4, 5 and 9 are not camera lucida drawings. All other and Lomb camera lucida was use e Zeiss camera lucida was employed. Leitz-Wetzler compound microscope was used as a rule. IG. 10, however, was made with a Zeiss microscope. The jongitudinal sections were cut in a plane at tion is indicated for most of the figures. The following symbols are employed: b, leaf base; c, cotyledon; cs, cotyledonary eheath; eé, sisi amie f, flower stalk; i flower trace; hf, bulb-axis or ‘‘tigelle’”’; /, lam o, open-base leaf; pc, pro- mbium; pcs, procambial strand; pm, Fe ete ae stem; 7, root; sc, closed leaf; < old flower stalk; #, testa. PLATE 14 Fic. 1. Five-day-old seedling; one half of the seed coats and the endosperm has been removed to show the cotyledon; 4h is the hypocotyl, and /, the first leaf; see Fics. 2 and 11; scale =I cm. Fic Central longitudinal section of five-day-old. seedling, showing the sheath, ‘ of the first leaf, i, and the primordia of the second leaf, 2: and be, together with the em point, pm; the stippled areas show the location of vascular traces; scale = Fr differentiated from the primordial meristem, pm, but a so that as a whole the meristematic tissue of this second leaf occupies an area ia diagonally to the reset: axis and surrounding the primordial meristem like a collar; scale = © gets 4. Ten-day-old seedling; as the first leaf is thrust out far from the coty- isionary sheath into the air the cotyledon elongates and curves downward until it is parallel with itself and the vertical shoot, having passed through an angle of 180 degrees; scale = 1Ic Fic. 5 Thirty-seven -day-old seedling; 0: is the base of the first leaf, h, and Is is the second leat; ale = 1 cm. Fic. 6. cae of the thirty-seven-day-old seedling; scale = 0.1 mm.; 4 and ba oe be tranaos agram ees relative proportions of na cells, raphide-bear- + 7 ing ae sak beaten cells and vascular traces; scale = PLATE I5 Fic. 8. Longitudinal section (not quite through ais) of thirty-seven-day-old Aes nee young root cutting way through base of aa first leaf; this root arises e and at the base of the second leaf; Fic. 9. Primordia of the point to sinty- aay ae shown in Fic. 14; not a central section; scale = 0.1m Fic. 10. Longitudinal section mae central) through slightly developed plant to show region, 4, dividing shoot and root in its early formation; h is the first leaf. Primordia of bulb shown in Fic. 15. 362 CuuRCH: THE BULB IN COOPERIA DRUMMONDII Fic. 12. Detached young ‘‘offset’’ with two scales already formed; it arises Sais between the mature scales as a ae on. . 13. Young “‘offset’’ h has pierced the outermost scale of the bulb; root contraction in old bulb not eas, leaves and roots diagrammtaic; scale = PLATE 16 Fic. 14. Fifty-four- to sixty-one-day-old seedling. The cotyledon has become shriveled up and dead. As the first leaf is surrounded by the basal sheath of the cotyledon, so this second leaf sheath is surrounded by the basal sheath of the first leaf, in respect to which it is distichously placed; see Fig. 9; scale = 1 cm Fic. 15. Longitudinal section of thirty-one-day-old bulb; see Fic. 11; scale = 0.1 mm. Fic. 16. Longitudinal section (not central) of thirty-one-day-old bulb; 63 and 13 are the points at which the primordia of the third leaf are forming; otherwise the symbols are as in preceding drawings; scale = 0.1 mm. Fic 17. Longitudinal section (not central) of thirty-seven-day-old bulb; 1/4 is the fourth leaf, not a central section; the puckered or wrinkled condition of the primary root is due to contraction, a normal phenomenon; the outlay of root tissue has a narrow slice marked off into three sections—r, 2, and 3—for details of which see Fic. 19; scale = 0.1 m Fic. 18. Longitudinal section through thirty-seven-day-old seedling with a young root showing a well-differentiated root cap and primary tissues (dermatogen, periblem and pleurome); sca G. Showing under high-power magnification the root tissues laid off between the numbers 1-3 in Fic. 17; for further explanation consult text; scale A brief conspectus of the species of Kneiffia, with the characteriza- tion of a new allied genus FRANCIS W. PENNELL It has recently been shown that the best-known specific name among our sundrops must be transferred from one species to another. While such a change is always peculiarly unfortunate, the desire for a definite nomenclature makes it unavoidable. In tradition the name ‘“‘fruticosa’”’ had been handed down as applying to the glandular-fruited element of the aggregate at first known by that name, whereas study of the Clayton Herbarium specimen upon which Linnaeus based the species showed that the name must be associated with the plant bearing on the capsule glandless incurved hairs. Nomenclature should follow definite rules of procedure, but surely scientific truth may raise the question as to the advisability of continuing the name ‘‘fruticosa” for any species of a group of plants with herbaceous, strictly annual stems. May we not plead the right to reject a proved nomen falsum? Moreover, from this standpoint in freeing our most widely known Kneiffia from the onus of the word “‘fruticosa,” Dr. Blake has hardly improved nomenclature—surely not in the opinion of our genetical friends— by the substitution of the name “‘Aybrida.” One of the incentives to the present study has been the hope of finding for this species some appropriate name. Another incentive has been the desire to place correctly a plant characteristic of the restricted but most unique prairie near New York City, the Hempstead Plains. This plant, which seems amply distinct, appears below as K. velutina. I present the results of this study with hesitation. Species- lines have not always been found clear, and in any genus so near to Oenothera one may expect the same tendency to split into incipient species. However, before this genus likewise is selected 363 364 PENNELL: THE SPECIES OF KNEIFFIA for intensive cultivation and study, it may be well, from the viewpoint of the taxonomist, to present an outline of its com- position. For this study I have examined specimens in the herbaria of the New York Botanical Garden, United States National Museum, Missouri Botanical Garden, Academy of Natural Sciences of Philadelphia, University of Pennsylvania and Charleston Museum. Stem-leaves broadly linear to ovate. Bracts similar leaves, linear, longer than the capsules. Stigmas linear, conspicuous. Wings of capsule pronounced. KNEIFFIA. Mature srtens clavate-linear, not stipitate. Hypanthium 20-25 . long. Sepals with salasie aac ee tips, 2-4 mm. long. Petals 20-25 mm. long. Stem, leaves and capsules hirsute. Hypanthium ro-12 mm. long. Sepals with shorter tips, strigose or somewhat agbred ing-pubescent. Petals 1 tem, leaves. and iio ia or appressed-pubesce Mature capsule-body aa to oblong, more or less stipitate. i. K. pratensis. 2. K. sessilis. of earlier flowers 12-25 mm. long. ee capsule-body decidedly clavate, scent with normally incurved sande hairs, in Nos. 8 and 9 with some interspersed gland-tipped hairs. Mature eacinls about as wide as long, usually much shorter than the stipe. Capsule with minute appressed : hairs. Leaves broadly linear, strigillose to glabrous. Basal leaves narrowly oblanceolate. 3. K. subglobosa. silky. 4. K. arenicola. Mature capsule-body longer than wide, longer than, about equaling or sometimes shorter than the stipe. Basal leaves broadly lanceolate to ovate. Stipe and capsule-body both pubes- cent with glandless hairs, the PENNELL: THE SPECIES OF stipe frequently equaling or sometimes exceeding the cap- sule-body. Stipe of capsule in fruit equaling, or somewhat exceeding the permanently pubescent body. Plant Plant rue spreading Stipe of capsule in fruit iets the body, which tends to become glabrat Main stem-leaves 7-9 cm. long, glabrate. Capsule- body pubescent with mi- ; nute hai aia bie pubescent body. Stipe always shorter than the capsule-body. Leaves nearly linear, finely pu- bescent to glabrate, 5-9 cm. long. Capsule-body clavate. Leaves ee ea softly densely : pubescent, em. long. Capsule-body St ae Mature capsule-body oblong or nearly so, bescent with short straight gland- tipped hairs, or becoming glabrate. Main stem-leaves linear-lanceolate to 1 lanceolate-ovate, scarce aler neath. Stem pubescent to sid glabrous. Petals of earlier flowers 18-25 mm. long. Stem ‘pare aikicen to gla- brous. Leaves lanceolate. Stipe actrees than the capsule- body Stipe longer than the capsule- body. Leaves usually nar- rower. Stem more or less hirsute. Leaves olate-ovate. Inflorescence KNEIFFIA 365 5. K. frutic ae Se ie Gain 6. K. riparia. 4. K. brevistipata. 8. K. semiglandulosa. 9. K. velutina. 10. K. tetragona. toa. K. tetragona longistipata. usually congested. 1ob. K. tetragona hybrida. 366 PENNELL: THE SPECIES OF KNEIFFIA Main stem-leaves lanceolate-ovate to ovate, glaucous th Ste glabrous or rarely obscurely pubes- cent. Petals of earlier flowers 25-30 mm. long. 11. K. glauca. Petals of earlier flowers 5-15 mm. long. I florescence in a usually over one half height of plan Capaule oe with incurved glandless - Petals of earlier flowers I0O-1I5 mm. None: Inflorescence erect from the 12. K. Spachiana. Pee sparsely pubescent with gland- tipped hairs. Petals of earlier flowers 5-10 mm. long. Young inflorescence nodding. 13. K. perennis. Stem-leaves filiform-linear to filiform. Bracts del- , toid-ovate to ovate, shorter than the capsules. Stigmas very short, scarcely appearing as lobes. Wings of capsule ridge-like. PENIOPHYLLUM. KNEIFFIA Spach Knetffia Spach, Hist. Veg. 4: 373. 1835. Type species, Oenothera glauca Michx. I. KNEIFFIA PRATENSIS Small Knetffia pratensis Small, Fl. SE. U.S. 842, 1335. 1903. ‘Type, Jefferson Co., Mo., Eggert, June 11, 1878, in Herb. N. Y. B. ey Type seen. Kneifia Sumstinei Jennings, Ann. Carnegie Mus. 3: 480. Ppl. 19. 1906. “Dry upland field near Kittanning [Pennsylvania], D. R. Sumstine, June, 1905. Type specimens in the Pennsyl- vania Herbarium of the Carnegie Museum. Acc. No. 2905.” Isotype seen in the herbarium of the New York Botanical Garden. Oenothera pratensis Robinson, Rhodora 10: 34. 1908. Prairies and woods, Ohio to Wisconsin, Iowa and Arkansas; introduced near New York City, Pittsburgh and Washington. 2. Kneiffia sessilis Pennell, sp. nov. Stem 3-4 dm. tall, pubescent with ascending hairs. Leaves 6-9 cm. long, acutish, densely strigose-pubescent with ascending hairs, in age somewhat glabrate. Bracts much exceeding the PENNELL: THE SPECIES OF KNEIFFIA 367 capsules. Hypanthium 10-12 mm. long. Sepals 15-16 mm. long, long-attenuate; tips more or less free in the bud. Petals triangular, 15-17 mm. long. Filaments less than one half the length of the petals. Anthers 6-7 mm. long, yellow. Stigmas becoming one half length of style. Capsule linear, at least 9 mm. long, densely pubescent with ascending to appressed hairs; not seen mature. Type, Little] Rf{ock], Ark[ansas], collected in flower June 2, 1885, Dr. H. E. Hasse; in the herbarium of the New York Botan- ical Garden. Also ‘‘La. Hale,’ in the United States National Herbarium, with longer hypanthium, broader leaves and less dense pubescence apparently belongs to this species. 3. KNEIFFIA SUBGLOBOSA Small Kneiffia subglobosa Small, Bull. Torrey aol a* 177-5). 4800; ‘‘North Carolina and Georgia.’ Type, ‘‘on the slopes or summit of Stone Mountain, De Kalb County, Georgia,” col- lected in fruit September 6-12, 1894, J. K. Small, seen in the herbarium of Columbia University at the New York Botanical Garden; isotypes in the herbaria of the United States National Museum and the Missouri Botanical Garden. I have found no basis for the crediting of this =— to North Carolina. Open rocky slopes, over granite, central Georgia to central Alabama. 4. KNEIFFIA ARENICOLA Small Kneiffia arenicola Small, Fl. SE. U. S. 842, 1335. 1903. “Type, Biltmore Herb., no. 5649d, in Herb. N. Y. B. G.” Type, “sand hills, Augusta, Georgia,” collected in fruit July 27, 1900, seen in the herbarium of the New York Botanical Garden; isotype in the United States National Herbarium. Sand-hills and dry pine-barrens, in the Coastal Plain, South Carolina to southern Mississippi. 5. KNEIFFIA FRUTICOSA (L.) Raimann Oenothera fruticosa L. Sp. Pl. 346. 1753. ‘Habitat in Virginia.” Type, Clayton 36, is identified by Dr. S. F. Blake (Rhodora 20: 51. 1918) as the plant here considered. 368 PENNELL: THE SPECIES OF KNEIFFIA Oenothera florida Salisb. Prod. 278. 1798. New name for O. fruticosa L. . Oenothera linearis Michx. Fl. Bor. Amer. 1: 225. 1803. ‘‘Hab. in Carolina superiore.’’ Type not seen or verified. Kneiffia angustifolia Spach, Nouv. Ann. Mus. Par. 4: 367. 1835. “Habitat in Georgia, Carolina, et Virginia.’’ Type not seen or verified. Kneiffia linearis Spach, Hist. Veg. 4: 376. 1835. Kueiffia longipedicellata Small, Bull. Torrey Club 23: 178. 1896. “West Virginia to North Carolina and Florida.”” Type, “Albemarle Co., Virginia,’ collected May 21, 1889, W. C. Rives, seen in the herbarium of Columbia University at the New York Botanical Garden. Kneiffia fruticosa Raimann; Engler & Prantl, Nat. Pflanzenfam. gt: 3145 180s) Oenothera longipedicellata Robinson, Rhodora 10: 34. 1908. Open soil, sandy or barren, mostly in the Coastal Plain although extending inland upon suitable soils (as in the serpentine barrens of southeastern Pennsylvania and eastern Maryland), Long Island to Florida, Missouri and Texas. 5a. Kneiffia fruticosa humifusa (Allen) Pennell, comb. nov. Oenothera fruticosa humifusa Allen, Bull. Torrey Club 1: 3. 1870. Type, “Montauk Point Long Island. [7. F. Allen.] July 1869,” seen in the herbarium of Columbia University at the New York Botanical Garden. Knetffiia linearis Alleni Britton, Mem. Torrey Club 5: 235. 1894. Based on Oenothera fruticosa humifusa Allen. Knerfia Alleni Small, Bull. Torrey Club 23: 177. 1894. (?) Oenothera linearis Eamesii Robinson, Rhodora 10: 34. 1908. ‘Sandy shore of a salt pond, Stratford, Connecticut, E. H. Eames (type hb. Gray).’’ Perhaps rather a form of the species, approaching humifusa. (?) Ocenothera friticosa Hamestt Blake, Rhodora 20: 50. 1918. Sandy soil, eastern Long Island, New York, and perhaps on the Connecticut coast. Said to be of very distinct appearance and, locally near Montauk, to be extremely abundant. Specimens elsewhere along the coast approach the habit of this. PENNELL: THE SPECIES OF KNEIFFIA 369 6. KNEIFFIA RIPARIA (Nutt.) Small Oenothera riparia Nutt. Gen. N. Amer. Pl. 1: 247. 1818. ‘Has. On the banks of Cape Fear river, Wilmington, North Carolina, in situations subject to inundation.”’ Kneiffia riparia Small, Fl. SE. U.S. 842. 1903. Swamps along the Cape Fear River, near Wilmington, North Carolina. Little known, although recently re-collected by Dr. J. M. Macfarlane, C. S. Williamson, etc. Possibly not a distinct . species. 7. Kneiffia brevistipata Pennell, sp. nov. Stem 2-4 dm. tall, pubescent. Leaves 3-6 cm. long, linear- lanceolate, acutish, densely strigose-pubescent with ascending hairs, the young leaves very silky. Bracts slightly exceeding the capsules. Hypanthium 6-9 mm. long. Sepals 7-9 mm. long, acutish; tips not free in the bud. Petals triangular, 12-15 mm. long. Filaments about one half length of petals. Anthers 3-3.5 mm. long, yellow. Stigmas less than one half length of style. — Capsule-body clavate-oblong, 5-6 mm. long; wings raised beyond ridges; pubescent, becoming glabrate, especially distally; on a stipe less than its own length. Type, Poplarville, Mississippi, collected in flower and fruit, July 7, 1891, S. M. Tracy 1681; in the United States National Herbarium. Dry pine ridges, southern Alabama to eastern Louisiana. 8. Kneiffia semiglandulosa Pennell, sp. nov. Stem 3-6 dm. tall, finely pubescent, becoming glandular above, purple-red. Leaves 5-9 cm. long, lanceolate-linear, acutish, finely pubescent with ascending hairs to glabrate.. Bracts much exceeding the capsules. Hypanthium 10-20 mm. long. Sepals 10-13 mm. long, acuminate; tips not or slightly free in the bud. Petals triangular, 15-25 mm. long. Filaments about one half length of petals. Anthers 5-6 mm. long, yellow. Stigmas less than one third length of style. Capsule-body clavate, 8-10 mm. long, wings exceeding the prominent ridges; finely pubescent with spreading gland-tipped, usually also with some incurved glandless, hairs, often glabrous or nearly so; on.a stipe less than its own length Type, Biloxi, Mississippi, collected in flower April 21, 1891, S. M. Tracy 5064, in the United States National Herbarium. Pine-land, West Florida to southern Mississippi; on ballast at Wilmington, North Carolina. 370 PENNELL: THE SPECIES OF KNEIFFIA 9. Kneiffia velutina Pennell, sp. nov. Stem 2-4 dm. tall, pubescent. Leaves 2-4 cm. long, narrowly or broadly lanceolate, acute to acutish, densely soft-pubescent. Inflorescence less than one fourth height of plant. Bracts little exceeding, the upper shorter than the capsules. Hypanthium 9-12 mm. long. Sepals 10 mm. long, somewhat attenuate, so that buds are slightly caudate. Petals triangular, 15-20 mm. long. Filaments about one half length of petals. Anthers pale yellow. Stigmas about one third length of style. Capsule-body oblong, slightly clavate, 7-9 mm. long; the wings almost equaling width of body; pubescent with incurved hairs and, especially proximally, with some finer gland-tipped hairs. Type, dry sandy soil, Garden City, Long Island, New York, collected in flower June 23, 1902, F. A. Mulford; in the herbarium of the New York Botanical Garden. Dry sandy soil, apparently restricted to the Hempstead Plains of western Long Island, from which numerous collections have been seen. 10. Kneiffia tetragona (Roth) Pennell, comb. nov. Oenothera tetragona Roth, Catalecta 2: 39. 1800. A garden plant, for which an American origin is stated. No specimen seen but the full description would apply to the plant here considered. Oenothera fruticosa ambigua Nutt. Gen. N. Amer. Pl. 1: 247. 1818. ‘Has. Common around Philadelphia.” Oenothera incana Nutt. l. c. 247. 1818. ‘“‘Has. In dry woods, Maryland.—Dr. W. C. Barton, v. s. in Herb. Barton.” Oenothera pilosella Raf. Ann. Nat. 15. 1820. ‘Indiana, near Evansville.”’ (?) Oenothera canadensis Goldie, Edinb. Phil. Jour. 6: 325. 1822. “Island of Montreal.’’ The size of flowers would indicate that this plant must be K. tetragona. ‘ Apparently far out of the normal range of the species (see Macoun, Cat. Canad. PI. 1: 172. 1883). Oenothera ambigua Spreng: Syst. 2: 229. 1825. Oenothera serotina Sweet, Brit. Fl. Gard. 2: pl. 184. 1826. A garden plant, of which no specimen seen. Kneifia suffruticosa Spach, Hist. Veg. 4: 374. 1835. ‘‘Croit dans les Etats Unis, depuis la Géorgie jusqu’ au Canada.” PENNELL: THE SPECIES OF KNEIFFIA 371 Kneifia shee pan Spach, /. c. 375. 1835. ‘‘Oenothera serotina Sweet. .. Oenothera fruticosa phyllopus seen Bot. Mag. 64: sub pl. 3345. 1837. ‘‘ Bot. Mag. t. 332.” Oenothera fruticosa incana Hook. I. c. sub pl. 3545. _ 1837. Oenothera fruticosa hirsuta Nutt.; T. & G., Fl. N. Am. 1: 496. 1840. Oenothera hybrida ambigua Blake, Rhodora 20: 52. 1918. Dry soil, barrens, etc., New York to Alabama, Tennessee and southern Michigan; the commonest Kuneiffia of the Piedmont and Alleghanian floras. 10a. Kneiffia tetragona longistipata Pennell, var. nov. Leaves linear-lanceolate. Stipe equaling capsule-body. Type, woods near Clemson, Pickens County, South Carolina, collected in flower May 12, 1907, H. D. House 3340; in the herbarium of the New York Botanical Garden. Piedmont Region, North Carolina to Georgia. 10b. Kneiffia tetragona hybrida (Michx.) Pennell, comb. nov. Oenothera hybrida Michx. Fl. Bor. Amer. 1: 225. 1803. ‘Hab. in Carolina superiore.” Knetffia floribunda Spach, Hist. A el 4: 376. 1835. ‘‘Croit dans le midi des Etats-Unis.” Mountains of North Carolina, there nearly or quite replacing the typical form of the species. As examples may be cited: Biltmore Herb. 669b; Heller 263; and Standley 53606. 11. KNEIFFIA GLAUCA (Michx.) Spach Oenothera glauca Michx. Fl. Bor. Amer. 1: 224. 1803. ‘‘Hab. in sylvis remotis et occidentalibus flumini Mississipi confini- bus, versus regionem IIlinoensium.” Surely the plant here considered, a not known to occur so far west as the type station. Oenothera Fraseri Pursh, Fl. Amer. Sept. 2: 734. 1814. “In South Carolina Fraser . . . v. v. in Hortis.”’ Kneiffia glauca Spach, Hist. Veg. 4: 374. 1835. Kneiffia Fraseri Spach, 1. c. 375. 1835. oie PENNELL: THE SPECIES OF KNEIFFIA Oenothera fruticosa Fraseri Hook. Bot. Mag. 64. sub pl. 3545. 1837. Oenothera fruticosa glauca Lév. Monog. Onothera 107. 1902. Wooded mountain-slopes, southern Virginia to northern Georgia, eastern Tennessee and eastern Kentucky. 12. KNEIFFIA SPACHIANA (T. & G.) Small — Oenothera Spachiana T. & G., Fl. N. Am. 1: 498. 1840. ‘Texas, Drummond.” Kneifia Spachiana Small, Bull. Torrey Club 23: 179. 1896. Oenothera fruticosa race Spachiana Lév. Monog. Onothera 106. 1902. Sandy prairies, Kansas and northwestern Arkansas to Louis- iana and Texas. : 13. Kneiffia perennis (L.) Pennell, comb. nov. Oenothera perennis L. Syst. ed. 10, 998. 1759. Canada. Oenothera pumila L. Sp. Pl. ed. 2. 493. 1762. ‘‘Habitat in America septentrionali.”’ Oenothera chrysantha Michx. Fl. Bor. Amer. 1: 225. 1803. “Hab. a Quebec usque ad sinum Hudsonis.” Oenothera pusilla Michx. 1. c. 225. 1803. ‘‘Hab. in rupibus, ad lacus Mistassins.” Kneiffia chrysantha Spach, Nouv. Ann. Mus. Par. 4: 368. 1835- Kneiffia Michauxit Spach, Ann. Sc. Nat. Bot. II. 4: 167. 1835- Based on Oenothera chrysantha Michx. Knetffia pumila Spach, Hist. Veg. Phan. 4: 377. 1835. Ocnothera pumila chrysantha Gordinier & Howe, FI. Rensselaer Co., N. Y. 14. 1894. ‘“‘Poestenkill, Howe.”’ . Oenothera pumila rectipilis Blake, Rhodora 19: I10. 1917. ‘NEw Brunswick: dryish rocky ground, Petit Rocher, Gloucester Co., 21 Aug. 1913, Blake 5513 (Type in Gray Herb.).””. The unusual state in which the pubescence is spreading may be considered as a form. Dry fields, Nova Scotia and Quebec to Minnesota and North Carolina; northward to Hudson Bay; also on St. Pierre Island. PENNELL: THE SPECIES OF KNEIFFIA 373 PENIOPHYLLUM?* Pennell, gen. nov. Slender glabrous herb, with virgately branched stem. Leaves of two forms, the basal petioled, ovate, less than 2 cm. long, the numerous stem-leaves scattered, filiform-linear to filiform, longer. Flowers in spikes terminal on the stem and branches. Bracts deltoid-ovate to ovate, shorter than the capsules. Sepals partially cohering, reflexing in two pieces. Petals about 4 mm. long, tri- angular. Filaments unequal, glabrous. Anthers oblong, glab- rous. Style glabrous. Stigma broad, capitate, the four lobes scarcely or not distinguishable. Capsule 4-6 mm. long, ellipsoid, sharply 4-angled, not stipitate. Seeds angled, brown, I mm. long, irregularly clustered. Type species, Oenothera linifolia Nutt. 1. Peniophyllum linifolium (Nutt.) Pennell, comb. nov. Oenothera linifolia Nutt. Jour. Acad. Nat. Sci. Phila. 2: 120. 1821. ‘Habitat: On the summits of arid hills and the shelv- ings of rocks, near the banks of the Arkansas [T7.. Nuttall].” Specimen, labeled ‘‘Arkansa, Nuttall,’’ seen in herbarium of Columbia University at the New York Botanical Garden. Kneiffia linifolia Spach, Nouv. Ann. Mus. Par. 4: 368. 1835. Kneifiia linearifolia Spach, Ann. Sc. Nat. Bot. II. 4: 167. 1835. Based upon Oenothera linifolia Nutt. Sandy soil, prairies and open woodland, southern Missouri and Kansas south to western Louisiana and eastern Texas; also on granite in central Georgia and central Alabama. NEw York BOTANICAL GARDEN * From rnvior, thread, and $vAdor, leaf. Preliminary note on a differential staining of the cytoplasm of Characeae Romyn HITCHCOCK The observations here recorded were made principally on two species of Nitella, but they are believed to apply generally for plants of this family. Perhaps the means of differential staining of living cell contents will find useful application in the study of other plants, but thus far my attention has been confined to the charas. Although it is my intention to continue observations in this direction, this should not deter others from work on the same lines. The field opened for investigation by this means is large and of much interest. It has long been known that the peripheral layer of cytoplasm in a developed internodal cell of Chara is relatively dense and viscous, while that within is notably thinner. By the use of neutral red the cell contents of a Chara plant may be differentially stained while the plant is living. The color is readily taken up from a dilute solution. An elongated internodal cell thus stained shows the cyclosis more clearly and beautifully than in the natural condition. Observing such a stained cell there is seen a large central cylinder of a pronounced cardinal red or wine color, bordered on either side with a narrow line of green. Within the colored cylinder, or vacuole, are numerous sus- pended granules, vesicles and spherical elements of various kinds, the nature of which is, for the most part, undetermined. Some of these are deeply colored and can be more advantageously studied than in the natural state. Indeed, they take the stain before the vacuole shows a general coloration. Probably these various structures have never before been so clearly seen and under such favorable conditions for study. The borders appear green because the light passes through the marginal chloroplasts regularly arranged next to the cellulose wall. But through this green sheen may be clearly seen the outer layer : 375 376 Hitcucock: DIFFERENTIAL STAINING of denser cytoplasm, carrying small, uncolored granules and some spherical plasmic bodies in suspension, in active cyclosis, closely following the cell wall. This outer layer, which, for convenience, may be here specifically designated as protoplasm, is quite un- colored. Thus we have, if we imagine a cross section, the cellulose wall lined with the chloroplasts in a thin, reticulated stroma, then the thin layer of uncolored, moving protoplasm, within which is the strongly colored, wide cylinder. The line of demarcation between the colored cylinder and the uncolored protoplasm is as sharply defined as a thin cell wall. Clearly it is the so-called vacuole wall. This line is wavy and constantly changing as it yields to the irregularities in the thick- ness of the protoplasm stream. The stained contents of the vacuole participate, however, in the movement of the protoplasm without, following precisely the same course. Moreover, deep within the vacuole it may be observed that the neutral or indif- ferent line, which marks the separation of the currents moving in opposite directions, indicates also an invisible barrier within the vacuole, which cannot be easily crossed by stray elements from the circulating stream. A sharply defined differentiation of the cell contents has thus been effected. The significance of it is not yet clear. It is not possible at present to state precisely what particular parts or elements in the cell sap have taken the stain, or whether any color is held in solution. By far the greater number of the cor- puscles or spherical elements so evident in the circulating stream of charas, are suspended in the cell sap, not in the protoplasm. The Schleimblaschen of Naegeli, later named Wimperkorperchen and mentioned by Allen as ‘‘ciliated”’ granules of protoplasm, are often deeply stained, but apparently not always. The name is misleading since there is no evidence of the presence of motile cilia on these bodies. We have some admirable hypotheses concerning osmotic action in the plant cell. There are membranes permeable and selec- tively permeable, solutes obedient to the rules established for their guidance, whereby the whole train of physical phenomena becomes clear. It now becomes desirable to know how the coloring matter makes its way into the vacuole of the Nitella cell. The HitTcHcock: DIFFERENTIAL STAINING 377 solution passes directly through the outer membrane and the rotating protoplasm, without coloring or affecting them in any way, so far as can be observed. It also passes through the vacuole wall. Apparently some ingenuity will be required to explain the observed phenomenon by osmotic action. One is inclined to regard it as indicating a general permeability of the tissues to the colored solution, which happens to be one not injurious to life processes. The color is strongly held in the vacuole and in un- colored water cyclosis continues for many days. Perhaps it is true, as some have supposed, that living proto- plasm will not take up coloring matter. We are told that when the nucleus stains the cell is dead. We have seen, however, that protoplasm permits passage of the coloring matter used in these experiments. Should we, then, conclude, that the contents of the vacuole which stain so deeply are therefore dead matter? Such a view is not sustained by the observed active circulation of the colored cell sap along the definite lines established. A PECULIAR PLASMIC STRUCTURE IN THE NITELLA CELL Among the many corpuscular elements of the cytoplasm, of which we have almost no understanding, some are of such remark- able character as to justify particular mention, although they cannot yet be satisfactorily described. They are curious spherical masses of granular matter, of extreme plasticity, greatly varying in size up to 0.1 mm. in diameter. The granules are in a state of constant agitation, as though the mass were seething with life. Within these spheres may be seen from one to twenty or more concave, saucer-shaped disks, 5-15 u in diameter, with contours more or less irregular, in rapid rotation, like so many wheels. They move freely about in the sphere, constantly changing the plane of rotation, sometimes on edge, sometimes lying flat but always twisting about and changing position. The nature of the movement is suggestive of cilia, but thus far none have been detected. These have seemed to me as perhaps connected with the de- velopment of chloroplasts, many of which are found freely circu- lating in the cytoplasm. Although this suggestion rests upon a very weak foundation, I find it difficult to resist the impression from observations up to this time. ore.. Hitcucock: DIFFERENTIAL STAINING These structures were first observed by me on June 18, 1918, in a variety of Nitella opaca Ag., found at Ithaca, New York, although not recorded in the local flora. Since then I have re- peatedly observed them in another species of Nitella, not yet seen in fruit, and also in Chara coronata, var. Schweinitzii. From this it may be inferred the structures are common to this family of plants. The only mention I have found in the literature of anything resembling these structures is in an admirable paper by Goeppert and Cohn,* where they are rather imperfectly described, not quite as I have seen them. Doubtless the description relates to the same structures. The authors associated them with the formation of starch, , IrHaca, NEW YorRK SUPPLEMENTARY NOTE The long, hyaline rhizoids of Nitella afford a more favorable means of demonstrating the selective coloring. The vacuole of a rhizoid cell becomes: deeply colored, while the thin, uncolored outer stream is in active rotation. The remarkable fact, already mentioned, that only living cells become colored, is more clearly manifest in the small and transparent rhizoid cell. In any mass of rhizoids by far the greater number will be dead cells. These remain colorless in the dye, while the living cells begin to color immediately. So long as cyclosis continues in a cell, that cell will take the color. If the cell is dying and the cyclosis is weakening it will not stain so deeply as in active life; but whenever a trace of color is seen cyclosis can be detected. When cyclosis has ceased no coloration what- ever is visible. When a stained cell dies the color soon disappears from the vacuole, doubtless by diffusion in the outer water. It would seem as though we had in this a test for living matter! But what kind of living matter has the vacuole and how does it differ from the denser, outer protoplasm? Or, have we instead, a test for vitality in a cell? Speculation as to the significance of these facts is at present unprofitable. Osmotic action affords no explanation of a concentration of a solute by passing through a * Ueber die Rotation des Zellinhaltes in Nitella flexilis. Bot. Zeit. 7: 665-673> 681-691, 697-705, 713-719. pl. 10. 1849. HircucockK: DIFFERENTIAL STAINING 379 ~~ membrane; yet, clearly, the dye is taken from the dilute, outer solution and concentrated within the walls. With the printer’s proof at hand I would like to add that the vacuole contents may be strongly colored by reason of an acid reaction. Ordinary water, or distilled water, usually is slightly alkaline, slowly changing neutral red to yellow or orange. When Nitella is grown in water slightly acidified with acetic acid, the vacuole becomes much more deeply colored than the culture solution. INDEX TO AMERICAN BOTANICAL LITERATURE 1918-1919 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 stag used in the broadest se Reviews, and papers that neue exclusively to piaeah agriculture, horticulture, csiteenued products of vegetable origin, or laboratory methods are not included, an no attempt is made to index the literature af bacteriology. ee occasional tepid 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 Se in some important particular. If users of the Index will call the attention of the edito 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. Atanasoff, D. A novel method of ascospore discharge. Mycologia II: 125-128. f. 1-3. .7 Je 1919. Bacon, A. E. Along highway and cross-country in Oklahoma. Ver- mont Bot. & Bird Club Bull. 4 & 5: 8-12. Ap 1919. Bailey, I. W. Depressed segments of oak stems. Bot. Gaz. 67: 438-441. f. -4. 19 My 1919. Beach, W. S. The Fusarium wilt of China aster. Michigan Acad. Sci. Ann. Rep. 20: 281-308. pl. 18-22 +f. 28. 22 N. 1918. Bennett, C. W. Soft rot of pepper caused by Bacillus carotovorus. Michigan Acad. Sci. Ann. Rep. 20: 351, 352. pl. 38-4go. 22 N 1918 Bertoni, M. S. Contributiones preliminares al estudio sistematico, biologico y economico de las plantas del Paraguay. An. Cien. Paraguayos 2: 135-142. Ja 1918. Includes Cedrela tubiflora and Chorisia Josephinae, spp. nov. Bertoni, M.S. Essai d’une monographie du genre Ananas. An. Cien. Paraguayos 2: 250-322. Ja 1918. Bertoni, M. S. Graminaceas de las regiones forestales litorales del Alto Parana (Paraquaya, Brasilefia y Argentina). An. Cien. Para- guayos 2: 143-166. Ja 10918. ; 381 382 INDEX TO AMERICAN BOTANICAL LITERATURE Bertoni, M. S. La Stevia Rebaudiana Bertoni. La Estevina y la Re- baudina nuevas substancias edulcorantes. An. Cien. Paraguayos 2: %29-134.. -Ja-1918. Bessey, E.A. Rose M. Taylor. Phytopathology 9: 212, 213. 23 My 1919. Bethel, E. Puccinia subnitens and its aecial hosts—II. Phyto- pathology g: 193-201. 23 My 1919. Bonar, L. The rusts of the Douglas Laké region. Michigan Acad. Sci. Ann. Rep. 20: 277, 278. 22 N 1918. Bowles, C. W. Systematic botany. Am. Bot. 25: 57-63. My 1919. Brandes, E. W. Distribution of Fusarium cubense, E. F. S., the cause of banana wilt. Michigan Acad. Sci. Ann. Rep. 20: 271-275+ 22 N 1918. Brotherton, W. E: Note on inheritance in Phaseolus. Michigan Acad. Sci. Ann. Rep. 20: 152. 22 N 1018. Brown, W. H. The theory of limiting factors. Philip. Jour. Sci. 13: (Bot.) 345-351. f. z. N 1918. Brown, W. H., & Trelease, S. F. Alternate shrinkage and elongation of growing stems of Cestrum nocturnum. Philip. Jour. Sci. 13: (Bot.) 353-360. N 1918. Campbell, D. H. The derivation of the flora of Hawaii. 1-34- Stanford University. 1919. Leland Stanford Jr. Univ. Publ. Univ. Series. Clark, F. R. Bud formation of plant hypocotyls. Michigan Acad. Sci. Ann. Rep. 20: 146. 22 N 1918. Claassen, E. Mosses of several Ohio counties. Ohio Jour. Sci. 19: 362-366. Ap Igig. Clute, W. N. Officinal rch of the United States. Am. Bot. 25: 47-50. My roto. Cobb, F., & Bartlett, H. H. A case of Mendelian segregation in Oeno- thera pratincola. Michigan Acad. Sci. Ann. Rep. 20: 151. 22 N 1918. . Coons, G. H. Michigan plant disease survey for 1917. Michigan _ Acad. Sci. Ann. Rep. 20: 425-450. pl. gi-50. 22 N 1918. Coons, G. H. The soft rot of hyacinth. Michigan Acad. Sci. Ann. Rep. 20: 353, 354. .22 N 1918. INDEX TO AMERICAN BOTANICAL LITERATURE 383 Darlington, H. T. Weed immigration into Michigan. Michigan Acad. Sci. Ann. Rep. 20: 261-267. 22 N 1918. Dodge, C. K. Contributions to the botany of Michigan—II. Univ. Mich. Mus. Zool. Misc. Publ. 5: 1-44. 13 Jl 1918. Evans, A. T. Embryo sac and embryo of Pentstemon secundiflorus. Bot. Gaz. 67: 427-437. pl. 12. 19 My 1919. Evans, A. W. A taxonomic study of Dumortiera. Bull. Torrey Club 46: 167-182. 23 My 1919. Farwell, O. A. Necessary changes in botanical nomenclature. Rho- dora 21: 101-103. 21 My ro19. Farwell, O. A. Notes on the ae flora. Michigan Acad. Sci. Ann. Rep. 20: 161-195. 22 N1 Many new varieties and bbls e are ae Farwell, O. A. The Trillium grandiflorum group. Michigan Acad. Sci. Ann. Rep. 20: 155-159. 22 N 1918. Farwell, O. A. The yellow flowered cypripediums. Michigan Acad. Sci. Ann. Rep. 20: 197, 198. 22 N 1918. Fernald, M.L. Bidens connata Muhl., var. gracilipes n. var. Rhodora 21: 103, 104. 21 My 1919. Fernald, M. L. Rubus idaeus and some of its variations in North America. Rhodora 21: 89-98. 21 My 1 Flynn, N. F. Plants new to Vermont collected at Woodstock. Ver- mont Bot. & Bird Club Bull. 4 &5:7. Ap 1grg. Flynn, N. F. A word concerning xanthiums. Vermont Bot. & Bird Club Bull. 4 & 5:20. Ap 1919. Fraser, W. P. Cultures of heteroecious rusts in 1918. Mycologia #22 129-134. 7 Je roto: Free, M. Effect of low temperatures on greenhouse plants. Brooklyn Bot. Gard. Rec. 8: 14-17. Ja 1919. Galloway, B. T. Giant crowngalls from the Florida everglades. Phytopathology 9: 207, 208. pl. ro. 23 My 1919. Gardner, M. W. The mode of dissemination of fungous and bacterial diseases of, plants. Michigan Acad. Sci. Ann. Rep. 20: 357-423. 22 N 1918. Gates, F.C. A comparison of the trees found in cemeteries of Hancock County, Illinois, with the native trees of the same region. Michigan - Acad. Sci. Ann. Rep. 20: 245, 246. 22 N 1918. 384 INDEX TO AMERICAN BOTANICAL LITERATURE Giddings, N. J., & Berg, A. A comparison of the late blights of to- mato and potato. Phytopathology 9: 209, 210. pl. rz. 23 My 1919. Gleason, H. A. Notes of the introduced flora of the. Douglas Lake region. Michigan Acad. Sci. Ann. Rep. 20:153. 22 N 1918. (Abstract.) . Gleason, H. A. Scirpus validus for demonstrating procambium. Michigan Acad. Sci.. Ann. Rep. 20: 153. 22 N 1918. (Abstract.) Graves, A. H. Some diseases of trees in Greater New York. My- cologia 11: 111-124. pl. ro. My 1919. Griggs, R. F. The beginnings of revegetation in Katmai Valley. Ohio Jour. Sci. 19: 318-342. Ap 1919. [Illust.] Giissow, H. T. The Canadian tuckahoe. Mycologia 11: 104-110. - 7-9. 7 Je 1919. Grifola Tuckahoe sp.. nov. is described. Haas, A. R. C. Effect of anesthetics upon respiration. Bot. Gaz. 67: 377-404. f. I-4. 19 My Ig19. Harvey, L. H. A coniferous sand dune in Cape Breton Island. Bot. Gaz. 67: 417-426. f. 1-8. 19 My 1919. Harvey, R. B. Importance of epidermal coverings. Bot. Gaz. 67: 441-444. f. 1, 2. 19 My 1919. Hibbard, R. P. Salt ratios in soil cultures. Michigan Acad. Sci. Ann. Rep. 20: 147-150. 22 N 1918. Holman, R. M., & Reed, E. Notes on the phyotplankton and other algae of Douglas Lake and vicinity. Michigan Acad. Sci. Ann. Rep. 20: 153, 154. 22 N 1918. Howe, I. A. Some new stations for rare plants in northeastern Ver- mont. Vermont Bot. & Bird Club Bull. 4 &5: 15-19. Ap 1919. Hunt, N. R. The ‘‘iceless refrigerator” as an inoculation chamber. Phytopathology 9: 211, 212. pl. r2. 23 My 1919. Jennings, O. E. Potamogeton Vaseyi in northeastern Ohio. Ohio Jour. Sci. 19: 343. Ap 1919. Johnston, E.L. Thesandlily. Am. Bot. 25: 52-54. My 1919. Kern, F.D. North American rusts on Cyperus and Eleocharis. Myco- logia 11: 134-147. 7 Je 1919. Puccinia abrepta, P. liberta and Uredo incomposita, spp. nov., are described. INDEX TO AMERICAN BOTANICAL LITERATURE 385 Kitchin, P. C. The relation between the structures of some coniferous woods and their penetration by preservatives. Michigan Acad. Sci. Ann. Rep. 20: 203-220. pl. rz. 22 N 1918. Kotila, J. E. Frost injury of potato tubers. Michigan Acad. Sci. Ann. Rep. 20: 451-460. pl. 51, 52. 22 N 1918. Lee, H. A., & Merrill, E.D. The susceptibility of a non-rutaceous host to citrus canker. Science II. 49: 499, 500. 23 My 1919. Lyman, G. R. The advisory board of American plant pathologists. Phytopathology 9: 202-206. 23 My 19gI19. Macbride, J. F. Certain North American Umbelliferae. Contr. Gray Herb. 56: 28-35. 31 D 1918. Macbride, J. F. Further new or otherwise interesting Liliaceae. Contr. Gray Herb. 56: 1-20. 31 D 1918. Includes Dasylirion longistilum sp. nov. and several new names and new com- binations. Macbride, J. F. Reclassified or new Compositae, chiefly North Ameri- can Helenieae. ‘ Contr. Gray Herb. 56: 36-50. 31 D 1918. Includes Aster deserticola sp. nov. : Macbride, J. F. A revision of Mentzelia, section Trachyphytum. Contr. Gray Herb. 56: 24-28. 31 D 1919. Macbride, J.F. A revision of Mirabilis, subgenus Hesperonia. Contr. Gray Herb. 56: 20-24. 31 D 1918. Macbride, J. F. Various American spermatophytes, new or trans- ferred. Contr. Gray Herb. 56: 50-61. 31 D 1918. Guatteria boyacana, Duguetia vallicola, Machaerium Whit setae Cryptantha echinosepala, C. quentinensis, and Plantago Parishii, spp. nov., are des ribed. MacCaughey, V. The mangrove. Am. Bot. 25: 42-46. My 1919. MacDougal, D. T., Richards, H. M., & Spoehr, H. A. Basis of suc- culence in plants. Bot. Gaz. 67: 405-416. 19 May 1919. Makemson, W. K. The leaf mold of tomatoes, caused by Clado- sporium fuluum Cke. Michigan Acad. Sci. Ann. Rep. 20: 31I- 348. pl. 23-37. . 22 N 1918 Maxon, W. R. A new Cheilanthes from Mexico. Proc. Biol. Soc. Washington 32: 111, 112. 21 My 1919. Cheilanthes castanea. McCall, A. G., Norton, J. B. S., & Richards, P. E. Abnormal stem growth of soybeans in sand cultures with Shive’s three-salt solution. Soil Sci. 6: 479-484. pl. 1,2. D 1918. 386 INDEX TO AMERICAN BOTANICAL LITERATURE Millspaugh, C. F., & Sherff, E. E. Revision of the North American species of Xanthium. Field Mus. Nat. Hist. Publ. Bot. 4: 9-49. pl. 7-13. Ap I9gI9g. Includes descriptions of 4 new species. Murrill, W. A. Illustrations of fungi—XXX. Mycologia 11: 101- 103. pl. 6. 7 Je 1919. : Ganoderma Tsugae, G. sessile, Inonotus dryophilus and Tyromyces Spraguei are illustrated in color. Nelson, J. C. The gender of Rumex. Am. Bot. 25: 55, 56. My 1919. Nelson, J. C. Species east and west. Am. Bot. 25: 70, 71. My I9I9Q. Newcombe, F. C. Behavior of plants in unventilated chambers. Michigan Acad. Sci. Ann. Rep. 20: 145, 146. 22 N 1918. Oflive,] E. W. Key to some of the principal families of flowering plants. Brooklyn Bot. Gard. Leaflets 7: [1-4] 21 My 1919. Osterhout, W. J. V. Antagonism between alkaloids and salts in rela- tion to permeability. Jour. Gen. Physiol. 1: 515-519. f. 1-3. 20 My 1919 Pengelly, M. Demonstration of methods for the study of stomatal action. Michigan Acad. Sci. Ann. Rep. 20: 154. 22 N 1918. Pennell, F. W. Notes on plants of the southern United States—V. Bull. Torrey Club 46: 183-187. 23 My igr19. Includes Acerates hirtella sp. nov. Pollock, J. B. Blue-green algae as agents in the deposition of marl in Michigan lakes. Michigan Acad. Sci. Ann. Rep. 20: 247-260. pl. 10, 17. 22 N 1918. Pollock, J. B. The longevity in the soil of the Sclerotinia causing the brown rot of stone oon Michigan Acad. Sci. Ann. Rep. 20: 279, 280, 29 NY Seeley, D. A. The cS of the growing season in Michigan. Michi- gan Acad. Sci. Ann. Rep. 20: 223-232. f. 22-25. 22 N 1918. Shreve, F. The establishment of desert perennials. Jour. Ecology 5: 210-216. 3 Ja 1918. Shufeldt, R. W. Phytophotography or the science of photographing flowers. Am. Forestry 25: 1069-1074. f. 1-8. My 1919. Soth, B. H. The arctic gentian. Am. Bot. 25: 41. My 1919. [Il- lust.] byes’ INDEX TO AMERICAN BOTANICAL LITERATURE 387 Stewart, E. G. Mucilage or slime formation in the cacti. Bull. Torrey Club 46: 157-166. pl. 8. 23 My 1919. Swingle, W. T. Merrillia, a new rutaceous genus of the tribe Citreae from the Malay Peninsula. Philip. Jour. Sci. 13: (Bot.) 335-343. PS 6 0.°°N TO1B: Tanner, F. W. Bacteriology and mycology of foods. i-vi-+1-592. pl. 1-10 +f. 1-85. New York. 1919. Van Fleet, W. New pillar rose. Jour. Heredity 10: 136-138. f. 78, Ig. 25 Ap 1919. Rosa soulieana X R, setigera. Van Pelt, W. Onion diseases found in Ohio. Ohio Agr. Exp. Sta. Month. Bull. 4: 70-76. f. 1-6. Mr 1919. Wakeman, N. A. Pigments of flowering plants. Trans. Wisconsin Acad. Sci. 19: 767-906. 1918. Waterman, W. G. Ecology of the pine forest relic at Interlochen, _ Michigan. Michigan Acad. Sci. Ann. Rep. 20: 241-244. f. 26. 22 N 1918. Watkins, S. L. The western Azalea. Am. Bot. 25: 51, 52. My ot baton Weatherby, C. A. Further notes on Impatiens ete Rhodora 21: 98-100. f. z-6. My 1g19. Weatherby, C. A. An omission in the preliminary list of New England Ranunculaceae. Rhodora 21: 104. 21 My 1919. Weatherby, C. A. Long Pond. Rhodora 21: 73-76. 9 My 1919. Weaver, J. E. The quadrat method in teaching ecology. Plant World 21: 267-283. f. 1-7. N 1918. Westgate, J. M. Report of the Hawaii Agricultural Experiment Station, 1918. 1-55. pl. 1-11. Washington. 10 Ap 1919. Wheeler, L. A. Additions to West River Valley flora. Vermont Bot. & Bird Club Bull. 4 & 5:8. Ap 1919. Whetzel, H. H. Cooperation among plant pathologists. Cornell Countryman 16: [1-3] F 1919. Whetzel, H. H. George Francis Atkinson. Bot. Gaz. 67: 366-368. 18 Ap 1919. [Portrait.] Williams, E. F. George Golding Kennedy. Rhodora 21: 25-35. 25 Mr 1919. [Portrait.] 388 INDEX TO AMERICAN BOTANICAL LITERATURE Wolkoff, M. I. Effect of ammonium sulfate in nutrient solution on the growth of soybeans in sand cultures. New Jersey Agr. Exp. Sta. Ann. Report 38: 416-419. 1917. Woodcock, E. F. Structure of mature seed of Eriogonum microthecum. Michigan Acad. Sci. Ann. Rep. 20: 233-236. pl. 13. 22 N 1918. Woodward, R. W. Two Festuca varieties. Rhodora 21: 72. 14 Ap 1919. Yates, H. S. Some recently collected ee fungi—II. Philip. hea Sci. 13: (Bot.) 361-385. N11 rty-one new species in various genera are a en H.W. Ballota hirsuta, a recent adulterant for Marrubium vulgare. Am. Jour. Pharmacy g1: 147-156. f. r-9. Mr 1919. Zavitz, C. A., & Squirrell, W. J. Hay and pasture crops, grasses, clovers, etc. Ontario Dept. Agr. Bull. 269: 1-55. F 1919. [Illust.] Zeller, S. M., & Dodge,C.W. Arcangeliella, Gymnomyces,and Maco- ieee in North America. Ann. Missouri Bot. Gard. 6: 49-59. f. 1-3. F 1919, ‘tadities weniger caudata, Gymnomyces Gardneri and Macowanites echino- Sporus, spp. n CHURCH: COOPERIA DRUMMONDII BULL. TORREY CLUB ee fe Yi KOSH Ly VOLUME 46, PLATE I5 e?' on Nee CHURCH: COOPERIA DRUMMONDII VOI UME 46, PLATE 16 Bui. TORREY CLUB | Goel - dead tines <0 [ af vascular region - 1 | parenchyma es LC AC a CREE (((daccaq [ee a AWeguag] CHURCH: COOPERIA DRUMMONDII Voi. 46 No. 10 BULLETIN OF THE TORREY BOTANICAL CLUB ee OCTOBER, 1919 Studies in the genus Lupinus—lV. The Pusilli CHARLES PIPER SMITH (WITH TEN TEXT FIGURES) INTRODUCTION In my second and third papers of this series (Bull. Torrey Club 45: 1-22, 167-202. 1918), I treated the Microcarpi, the larger and more neglected division of Watson’s subgenus Platycarpos, and I now present my review of the species and varieties properly to be included in the smal'er and much less neglected group, the Pusilli. ; The following list includes the published names that need to be considered in this paper: pusilus Pursh, Fl. Am. Sept. 2: 468. 1814. brevicaulis Wats. Bot. King’s Report 53. 1871. Kingu Wats. Proc. Am. Acad. 8: 534. 1873. Silert Wats. ibid. 10: 345, 1875. Shockleyt Wats. ibid. 22: 470. 1887. capitatus Greene, Pittonia 1: 171. 1888. odoraius Heller, Muhlenbergia 2:71. 1905. scaposus Rydberg, Bull. Torrey Club 34: 45. 1907. rubens Rydberg, ibid. 34: 45. 1907. dispersus Heller, Muhlenbergia 5: 141. 1909. flavoculatus Heller, ibid. 5: 149. 1909. intermontanus Heller, ibid. 8: 87. 1912 argillaceus Woot. & Standl. Contr. U. S. Nat. Herb. 16: 1913. ab Jaap R aaa In my discussion of L. malacophyllus Greene, I indicated a disposition to question the inclusion of this species in the Micro- [The BuLLetIN for September (46: 337-388. pl. 14-16) was issued October 17, 1919.] 389 390 SmitH: STUDIES IN THE GENUS LUPINUS carpi, but so placed it on the basis of its verticillate inflorescence. The non-ciliate keel and the floral bracts, however, connect it with the Pusilli. To contrast it further with ZL. subvexus trans- montanus C. P. Smith (Bull. Torrey Club 45: f. zo), with which it is confused, I insert here drawings of my own, FIG. 43. I now con- sider. it better to compromise the situation by recognizing three divisions of the subgenus, and accordingly propose the following classification : Flowers not verticillate; keel glabrous. PUSILLI. Flowers, at least the lower, in whorls. Keel glabrous; flowers 10-12 mm. long. MALACOPHYLLI. Keel ciliate above near claw; flowers 11-19 mm. long. MICROCARPI GEOGRAPHICAL DISTRIBUTION The Pusillt are peculiar to western North America. Typical L. pusillus is primarily a species of the Atlantic drainage of the Rockies, from Alberta and North Dakota to western Kansas LO8 Fic. 43.. LuPINUS MALACOPHYLLUS Greene. A. A. Heller 9606 (CPS). and New Mexico. All the other forms of the group, however, are plants of the Great Basin and Mexican Plateau, from central Washington and southwestern Wyoming to Arizona and southern California. DIAGNOSTIC CHARACTERS Authors’ ‘original descriptions’? are not reprinted in this paper. The diagnoses here presented have no pretense of being complete, but are intended to sum up briefly the seemingly more dependable characters. These descriptions, therefore, allow for the evident variations observed by me in the specimens listed. The upper calyx-lip calls for special mention. While this — seems to be relatively constant in size and shape in most of the forms of this group, its variability in L. brevicaulis should be duly SMITH: STUDIES IN THE GENUS LUPINUS 391 recognized. This variability paratlels equal diversity in L. subvexus and L. densiflorus, as recently shown by me (Bull. Torrey Club 45:5. f. 7, 16, 24, 30, 31, 35, and 42). In like manner, varia- tions in length of peduncles and details of branching are not accepted by me as specific in character unless supported by recog- nizable differences in the floral parts or fruits. | This conclusion is followed in my treatment of the variable L. Kingii, where these variations are evidently superficial rather than fundamental. My measurements of the floral parts are from the softened material, mounted in water, hence should agree rather with the fresh flowers than with dried material. An allowance of about 10 per cent should be made when measurements of dried flowers are compared: with my figures. MATERIAL EXAMINED AND CITED | Seven institutions have contributed to this paper by making it possible for me to examine their specimens of this group. The names of these institutions are given below, with the respective abbreviations used in the citations. ‘‘CPS” indicates that the specimen cited is in the private collection of the writer, and ‘‘FS” indicates a specimen owned by the United States Forest Service. B, Brooklyn Botanic Garden; CAC, Colorado Agricultural College; G, Gray Herbarium of Harvard University; NY, New York Botanical Garden; RM, Rocky Mountain Herbarium, University of Wyoming; T, Torrey Herbarium (et New York Botanical Garden) ; UC, University of California; US, United States National Herbarium. Key to the species d varieties of the Pusilli Flowers usually crowded into head-like racemes. ms short, seldom over 1 cm. long; leaves crowded basally; upper calyx-lip not over 2 mm. long, bifid, tiednbed or nearly obsolete. 8. L. brevicaulis. Stems elongated, several cm. long; ieayes scattered; pper calyx-lip 3-6 mm. long, ‘ Pubescence copious, that of the pedicels and calyx long and spreadin ya. L. Kingii. Pubescence scant, that of ie pedicels and calyx ~ short and appressed. 4b. var. argillaceus. 392 SMITH: STUDIES IN THE GENUS LUPINUS Flowers scattered in elongated racemes. : owers distinctly ascending in anthesis. 8. [L. scaposus.] Flowers spreading in anthesis. ; Ra ensely pubescent with hairs about I mm. ong; pods smooth or scaly on the sides. Flowers about 6 mm. long; banner angled at apex; pods ovate, 10-15 mm. long; ovules two. 9. L. Shockleyi. Flowers 10-12 mm. long; banner rounded at apex; pods oblong, about 20 mm. long; ovules two-four. 10b. var. pilosellus. Loosely villous or varying to glabrate Stems almost or quite glabrous, calyx strictly so; pods eee smooth or scaly nm the sides, about 2 m. long, ovules two-six; seeds 2-3 mm. across. toa. L. odoratus. Stems obviously villous, at least the lips of oe e calyx villous; pods usually villous on the sides; ovules mostly two. Racemes obviously exceeding the foli- ge; pods ovate, about 10 mm long, Mid villous on the sides; seeds 2-3 m Branches pHa pies the ial peduncle erect and early- flowering. Ita. L. rubens. ranches’ early-developing, all floriferous and ast sea a 116. var. flavoculatus. Racemes equalled or su by the foliage; pods eeechoping usually somewhat constricted near the middle, shaggy on the sides, about 20 mm. long; seeds about 4 mm. eo Flowers 10-12 mm. long; lower calyx-lip over twice as long as ide. 12a. L. pusillus. Flowers 7-8 mm. long; lower calyx-lip barely twice as long wide. 12), var. intermontanus. 7a. Lupinus Kine Wats. Proc. Am. Acad. 8:534. 1873. [FIG. 44]. See Lupinus Silert Wats. ibid. 10: 345. 1875 (in part). Lupinus capitatus Greene, Pittonia 1: 171. 1888. Stems well-developed, erect with ascending branches or varying to widely spreading, 1-4 dm. tall, branched at the base or more commonly well above the base, spreading-villous; leaves usually SMITH: STUDIES IN THE GENUS LUPINUS 393 scattered, glabrous above or more or less hairy; peduncles variable in length, the short capitate or subcapitate racemes exceeding the foliage or surpassed by same; flowers crowded, I0O-II mm. long, commonly few; pedicels 1-2 mm. long, villous; calyx bracte- olate, villous, upper lip 4-7 mm. long, bifid, the divisions slender, lower lip 5.5-7 mm. long, three-toothed; petals blue or purple, narrow, banner 8-10 mm. long, 3-5 mm. wide, usually acute at apex, wings 6-8 mm. long, 2-3.5 mm. wide, oval or oblong, keel straight, 6-8 mm. long, about 3 mm. wide; pods rhombic-ovate, about 10 mm. long; seeds about 2 X 2.5 mm. Watson soon recognized the affinity of his supposed perennial, L. Kingii, and his annual, L. Sileri, and reduced the latter to synonymy. This viewpoint has been followed by Coulter (Man. Bot. Rocky Mts. 273. 1885), Coulter and Nelson (New Man. Rocky Mt. Bot. 469. 1909), Rydberg (Fl. Colo. 100, 1906; FI. a SS ag a fem 3 1 2 See 5 G. 44. LUPINUS Kincn Wats. 1. Korstion & Baker 131 (FS); 2. C. A. Purpus bie aay, 3. S. Watson 234 (US); 4. C. F. Baker 439 (US); 5. M. E. Jones 5641 (US). Rocky Mts. 469. 1917), and Heller (Muhlenbergia 6:71. 1910). Heller and Rydberg have also referred Greene’s L. capitatus to Watson’s species. Wooton and Standley (Contr. U.S. Nat. Herb. 19: 340. 1915), however, have recognized L. Kingii and L. Sileri as distinct species, each represented in New Mexico, and have segregated L. argillaceus as a new species. My study of the specimens listed below prevents my recognition of more than one species here, though as least one variety probably deserves at- tention. I can not see specific characters in the variations in length of peduncles, or in the extremes of types of branching (widely divaricate to ascending) here; but the more sharply de- fined variation in pubescence seems to be of sufficient definition for use in varietal recognition. Greene attributes to L. capitatus 394 SMITH: STUDIES IN THE GENUS LUPINUS an “entire” lower calyx-lip; but the several Coconino County specimens examined by me have the normal three-toothed lower lip. Utan. Beaver County: Beaver, Aug., 1882, M. E. Jones (G). Garfield County: Panguitch Lake, 7 Sept., 1894, M. E. Jones 60151 (US). Piute County: Grass Valley, L. F. Ward 648 (US); Grass Valley, Aug., 1882, Powell Exped., L. F. Ward (G). San Juan County: mesa east of Monticello, July, 1911, P. A. Rydberg & A. O. Garrett 9214 (US, NY). Sevier County: Burrville, July, 1894, M. E. Jones 5641 (US, RM); Loa Pass, July, 1894, M. E. Jones 5641 (NY, UC); Fish Creek Canyon, July, 1909, A. O. Garrett 2532 (NY). Summit County: Parley’s Park, July, 1869, S. Watson 234 (US). County not given: 1872, Capt. Bishop (US); Juniper Mountains, 1898, C. A. Purpus 6256 (US). Cotorabo. Archuleta County: Piedra, July, 1890, C. F. Baker 439 (G, NY, RM, US); Pagosa Springs, Aug., 1904, E. O- Wooton (US). Costilla County: Garland, July, 1896, C. L. Shear 3682 (NY); San Acacio, Culebra Creek, July, 1912 E. R. Warren (RM). Delta County: without definite locality, June, 1892, J. H. Cowen (US). Gunnison County: Gunnison Water- shed, July, 1901, C. F. Baker 452 (G, NY, RM, UC, US). LaPlata County: Durango, July, 1896, F. Tweedy 452 (US). LaPlata or Montezuma County: Parrott City, Rio Mancos trail July, 1875, Hayden Survey 1197, T. S. Brandegee (UC); trail, between Rio de los Mancos and Rio !a Plata, July, 1875, 7. 5. Brandegee (G); Thompsons Park, LaPlata Mountains, July, 1898, C. F. Baker, F. S. Earle, & S. M. Tracy 441 (G, NY, UC, US). County not given: Colfax, Sept., 1877, T. S. Brandegee (UC); 1896 F. Clements 318 (NY). New Mexico. Bernalillo County: Tijeras, Sandia Moun- tains, May, 1914, C. C. Ellis 430 (US). Dona Ana County: Organ Mountains, May, 1905, EF. O. Wooton (RM). Grant County: GOS Ranch, Aug.-Sept., 1911, J. M. Holzinger (US). McKinley County: Cooledge [Guam], July, 1889, M. Hopkins (US). Rio Arriba County: vicinity of Chama, July, 1911, P. C. Standley 6793 (US); Dulce, Jicarilla Apache Reservation, Aug., 1911, P. C. Standley 8071 (US); El Rito, Aug., 1904, E. O. Wooton (US). Sandoval County: Rito de las Frijoles, Aug., 1902, T-. SMITH: STUDIES IN THE GENUS LUPINUS 395 D. A. Cockerell 19 (US); Rito de las Frijoles, Aug., 1910, W. W. Robbins (RM). San Juan County: Tunitcha Mountains, Aug., ee okee te Standley 7786 (US). Socorro County: Mogollon Mountains, middle fork of the Gila, Aug., 1900, E. O. Wooton (US); Wheelers Ranch, July, 1906, E. O. Wooton (RM, US). ~ Arizona. Apache County: White Mountains, Aug., 1903, D. Griffiths 5273 (US); Alpine, July, 1912, L. N. Goodding 1258 (RM). Cochise County: Bisbee, July, 1912, Z. N. Goodding 1258 (US). Coconino County: Cosnino, July, 1883, H. H. Rusby (US); Flagstaff, Aug., 1884, M..E. Jones 3947 (NY, RM, US); San Francisco Mountains, 2 Sept., 1889, F. H. Knowlton 174 (US); Flagstaff, June, 1892, J. W. Toumey 573 (US); Grand Canyon, July, 1892, J. W. Toumey 575 (G, US); Grand Canyon, July, 1892, E. O. Wooton (US); Flagstaff, Sept., 1895, J. W. Toumey (NY, UC); Flagstaff, July, 1897, C. L. Harris 913 (US); Flagstaff, 1900, C. A. Purpus 8083 (US); Flagstaff, June, 1901, J. B. Leiberg 5522 (US); Flagstaff to Mugillo Mountains, July, 1903, D. Griffiths 4974 (US); Coconino Forest, Aug., 1908, G. A. Pearson rz (US); Grand Canyon, July, 1912, V. Rattan (UC); Grand Canyon, June, 1913, A. E. Hitchcock 39 (US); Flagstaff, Aug., 1915, A. S. Hitchcock (US); Kaibab Forest, 1918, Korstion & Baker 131 (FS). Navajo County: Taylor, June, 1897, W. Hough 72 (US). Pima County: Sabina Canon, Catalina Mountains, Apr., 1892, J. W. Toumey 572 (US 211833 in part). Yavapai County: Ashfork, July, 1903, D. Griffiths 4873 (US). County not determined: Hell’s Canyon, May, 1883, H. H. Rusby 1402 (US); Hell’s Canyon, June, 1883; H. H. Rusby 552, 554 (UC); Nagles Ranch, 13 Sept., 1894, M. E. Jones 6054¢ (US). 7). Lupinus Kingii argillaceus (Woot. & Standl.) ‘comb. nov. [Fic. 45.] Lupinus argillaceus Woot. & Standl. Contr. U. S. Nat. Herb. 10: 137. - 1013. Differs from the typical form of the species in the sabicoebiace being shorter and subappressed, especially that of the calyx and pedicels being appressed and inconspicuous. No other characters seem to separate this from the widely branching individuals of the obviously villous species. 396 SMITH: STUDIES IN THE GENUS LUPINUS New Mexico. San Miguel County: near Pecos, Aug., 1908, P. C. Standley 40974, 4975 (NY, US). McKinley County: Rio Zuni, July, 1892, E. O. Wooton (US 735190 and 562216). SESS ( OI Fic. 45. Lupinus KINGU ARGILLACEUS (Woot. & Standl.) C. P. Smith. 1. P.C. Standley 4975 (US); 2. E. O. Wooton (US 562216); 3. J. Wolf 195 (US). CoLtorabo. Mesa County: Loma, Rio Grande, Sept., 1873, . Wolf 195 (US). mM . LUPINUS BREVICAULIS Wats. Bot. King’s Report 53. 1871. [Fic. 46.] Lupinus dispersus Heller, Muhlenbergia 5: 141, 1909. Lupinus scaposus Rydberg, Bull. Torrey Club 34: 45. 1907. Low, less than 1 dm. tall, densely villous, stems short, seldom over I cm. long; leaves crowded basally, glabrous above: peduncles 3-6 mm. long, the subcapitate racemes about 2 mm. long, com- monly elongating after anthesis; flowers 6-8 mm. long, crowded; pedicels 1-2 mm. long, villous; calyx bracteolate, villous, upper lip 1-2 mm. long, truncate, two-lobed or bifid, lower lip 4-6 mm. long, two- to three-toothed, or rarely entire; petals bright blue or pale-tinted, banner about 6 mm. long, 2-4 mm. wide, apex angled or rounded, wings 4-6 mm. long, 2 mm. wide, keel 4-6 mm. long, straight; pods ovate, about 10 mm. long, ovules two or three, seeds 2 mm. long. From the first I experienced difficulty in attempting to follow others in separating L. brevicaulis and L. dispersus. Careful attention to the original descriptions shows few differences brought out, except as to the calyx-lips. Quoting from Watson: . upper lip very short, truncate, obsoletely toothed, membranous, lower entire or obscurely toothed. Quoting from Heller’s ideaees of his species: SMITH: STUDIES IN THE GENUS LUPINUS 397 - . . upper calyx lobe less than 2 mm. long, 2-parted almost to the base, the divisions with a broad somewhat U-shaped sinus; lower lobe nearly 4 mm. long, prominently 2-toothed, the sinus narrow. .. . I have dissected flowers and made drawings from fourteen of the numerous collections examined and here listed. As shown in Fic. 44, both the lower and upper calyx-lips exhibit considerable Fic. 46. LUPINUS BREVICAULIS Wats. 1. A.A. Heller 8646 (US); 2. M. E. Jones 3814 (US); 3. Mrs. J. Clemens (CPS); 4. M. E. Jones 5273 (US); 5. M. E. Jones 5130y (US); 6. M. E. Jones 5272b (US); 7. D. Griffiths 3055 (US); 8. E. O. Wooton (US 737532); 9. C. F. Baker 437 (US); 10. J. B. Leiberg 2108 (US). variation. The lower lip is entire in only one of these fourteen cases, i.e. Heller’s Reno collection. Five cases show a three- toothed lower lip, four cases the two-toothed condition, and two collections show both two-toothed and three-toothed individual calyces. Rydberg’s L. scaposus is evidently a pathological phase of L. brevicaulis. I have examined the type collection (T, G, RM) and cannot find justification for recognizing this form even as a variety. Three other collections, from as many states, include specimens of this variation. These are: ‘‘Jones 5273, Kanab, Ut.’’ (RM 14246); ‘““‘Lemmon, Camp Lowell, Ari.’”” (G); and ‘“‘Watson 223, Coyote Mts., Nev.’’ (T). We should not overlook Watson’s own record (1871, p. 53) which reads: A form of this was collected with some of the peduncles much elongated and bearing above the leaves loose racemes of reduced florets, apparently perfect in all their parts, but usually sterile (223). I see only the one species in this series of specimens. If the extremes of western Nevada were alone before me, I could readily accept Heller’s segregate as a variety of Watson’s species; but 398 SMITH: STUDIES IN THE GENUS LUPINUS careful study of all the material considered shows no sufficiently constant or fundamental differences available for a satisfactory varietal classification. OrEGON. Lake County: Chewaucan Marsh, 2 June, I9II, W. W. Eggleston 6771 (US); Fifteen Mile Creek, Warner Valley, 16 June, 1911, W. W. Eggleston 6983 (US). Harney County: base of Steins Mountain, 30 May, 1885, 7. Howell (US, G). Malheur County: Barren Valley, 1885, W. Cusick 1250 (US); Dry Creek hills, June, 1901, W. Cusick 2554 (US, G, NY, UC, RM); Barren Valley, June, 1896, J. B. Leiberg 2198 (US, G, UC). Nevapa. Humboldt County: Coyote Mountains, June, 1868, S. Watson 223 (US 41363, T); Humboldt Valley, 1868, S. Watson 222 (T). Washoe County: Reno hills, June, 1900, S. G. Stokes (NY); Truckee Pass, June, 1906, P. B. Kennedy 1333 (US, NY); Truckee Pass, June, 1907, A. A. Heller 8646 (US, B); Reno, 1 May, 1910, A. A. Heller roo0o6 (US, B, NY, RM). Mineral County: Hawthorne, June, 1882, M. E. Jones 3814 (US, NY). Esmeralda County: Candalaria, W. H. Shockley 5r (UC). Nye County: Tonopah, June, 1907, W. H. Shockley 86 (UC); Rhyolite, June, 1907, W. H. Shockley 54 (UC). Lincoln County: Caliente, May, 1902, L. N. Goodding 942 (RM). Utan. Weber County: Ogden, June, 1903, S. G. Stokes (US). Davis County: Antelope Island, June, 1869, S. Watson 222 (US). Salt Lake County: near Garfield, May, 1909, Mrs. J. Clemens (US, NY, RM, CPS). Juab County: Diamond Valley, 28 Apr., 1894, M. E. Jones 5125c (US). Piute County: Marysvale, May, 1894, M. E. Jones 5338i (US). Washington County: La Verken, May, 1894, M. E. Jones 5183b (US); Silver Reef, May, 1894, M. E. Jones 5149g, 5151 (US); Santa Clara Valley, Apr., 1894, M. E. Jones 5139h, 5139y (US); Spring Dale, May, 1894, M. E. Jones 5261h (US). Kane County: Kanab, May, 1894, M. E. Jones 5273 (US, NY, UC, RM). Cotorapo. Garfield County: Glenwood Springs, June, 1899, G. E. Osterhout (T, G, RM). Delta County: Surface Creek, March, 1892, C. A. Purpus 175 (UC); Hotchkiss, June, 1802, J. H. Cowen 147 (NY, G); same, J. H. Cowen 648 (CAC); Delta, June, 1909, L. Tidestrom 2198 (US). Montrose County: Naturita, May, 1914, E. Payson 306 (NY, RM). La Plata County: Dur- SMITH: STUDIES IN THE GENUS LUPINUS 399 ango, June, 1898, C. S. Crandall (RM). Archuleta County: Arboles, June, 1899, C. F. Baker 437 (US, G, NY, UC, RM). New Mexico. McKinley County: Fort Wingate, 1883, W. Mathews (G). Socorro County: Patterson, June, 1892, E. O. Wooton (US 241178, US 737533). Dona Ana County: plains at base of Organ Mountains, 1852, C. Wright 1362 (US 20950-2); Filmore Canyon, Organ Mountains, May, 1905, E. O. Wooton (US). Grant County: Copper Mines, June, 1851, Mex. Bound. Survey 287 (NY); River Mimbres, May, 1851, G. Thurber 335 (T); Silver City, 1891, G. C. Nealley 39 (NY); same, G. C. Nealley 40 (US); Silver City, May, 1906, V. Bailey (US). County not _ given: without definite locality, 1869, E. Palmer (US 20576, T): Mesa Top, 1883, C. D. Walcott 28 (US). CuinuAHuA. Ojo de Vacate, June, 1851, G. Thurber 335 (NY). ARIZONA. Cochise County: Camp Lowell, 1880, J. G. Lem- mon (G). Santa Cruz County: Tumacacori and vicinity, 1903, D, Griffith 3055 (US). Pima County: Canaca to Arivaca, 1903, D. Griffith 3570 (US). Yavapai County: Ash Fork, May, 1903, D. Griffith (US). Coconino County: base of San Francisco Mountains, July, 1884, J. G. Lemmon & wife (US, UC); Hell’s Canyon, May, 1883, H. H. Rusby 554 (US, T); Turkey Tanks, . Flagstaff, June, 1891, D. T. McDougal (US); Pipe Spring, May, 1894, M. E. Jones 5272b (US). Mohave County: Hackberry, May, 1884, M. E. Jones (US 220209); near Kingman, June, 1893, NV. C. Wilson (UC). CALIFORNIA. San Bernardino County: New York Mountains, near Leastalls, June, 1915, S. B. Parish 10259 (UC). Inyo County: Bishop Creek, May, 1886, W. H. Shockley 424 (UC). 9. LuPINUS SHOCKLEYI Wats. Proc. Am. Acad. 22: 470. 1887. [Fic. 47.] More or less acaulescent, 1-2 dm. tall, densely pubescent with hairs about 1 mm. long, spreading on the stems and petioles, sub-appressed on the leaves; peduncles 5-10 cm. long, racemes lax, 3-6 cm. long, equalled or surpassed by the foliage; flowers about 6 mm. long; pedicels slender, often curved, about half as leng as the flowers, spreading-pubescent; calyx hairy, the upper lip 3 mm. long, bifid, lower lip 3-4 mm. long, minutely three-toothed or 400 SMITH: STUDIES IN THE GENUS LUPINUS sometimes bifid; petals “blue or purple’’ or pink, banner 5-6 mm. long, 4 mm. wide, angled at apex, wings 5 X 3 mm., keel straight, 4 mm. long, 2 mm. wide: pods ‘“‘oblong ovate,” ciliate on the margins, scaly on the sides, ovules two, seeds rough, about 3 mm. in diameter, pale in color. ArIzoNA. Mohave County: Kingman, May 1884. J. G, Lemmon & wife (UC). NEvapA. Esmeralda County: Sodaville, W. H. Shockley 367 (T, UC); same locality, June, 1888, W. H. Shockley 43509 (NY)- Mineral County: Hawthorn, June, 1882, M. E. Jones 3812 (NY, RM, US). . CALIFORNIA. Kern County: Tehachapi, June, 1884, K. Brandegee (UC). Riverside County: Palm Springs, Apr., 1905, H. M. Hall 5762 (UC); Point of Rocks, Whitewater, Apr., 1911, TZN WoW? ‘Fic. 47. Lupinus SHOCKLEYI Wats. 1. M.E. Jones 3812 (US); 2. J. G. Cooper (US 344389). E. E. Schellenger (UC). San Bernardino County: Camp Cady, Mojave River, June, 1861, J. G. Cooper (US); Browns Ranch, Mojave Desert, May, 1882, S. B. Parish (UC) ; Morongo, Colorado Desert, March, 1882, S. B. Parish 1270 (NY; from the herbarium of H. E. Hasse); Mojave River, E. Palmer (NY; from the her- barium of P. V. Le Roy); Barstow, K. Brandegee (UC); Daggert, May, 1917, K. Brandegee (UC); Kelso, May, 1917, K. Brandegee (UC); Colorado Desert, Apr., 1905, T. S. Brandegee (UC). San Diego County: Borregos Spring, T. S. Brandegee (UC). The New York sheet from Dr. Hasse’s herbarium (S. B. Parish 1270) includes two specimens of L. Shockleyi and one specimen each of L. odoratus, L. concinnus, and L. microcarpus var. ruber. Mr. Parish informs me that his No. 1270 belongs to a collection of L. odoratus taken in May, 1882, while his visit to the Morongo was fff April of that year. It seems evident, therefore, that this label cannot be accepted as correctly applying to any of the % SMITH: STUDIES IN THE GENUS LUPINUS 401 specimens assembled on the sheet. The boundary line sep- arating Riverside and San Bernardino Counties passes through the Morongo country and it seems to be impossible to determine on which side of the line were taken the Morongo specimens here referred to San Bernardino County. Palmer’s specimen from Le Roy’s herbarium is a striking vari- ation towards L. odoratus and probably should have been given more serious attention here. Ioa. Lupinus oporATus Heller, Muhlenbergia 2: 71. 1905. [FIG. 48.] Nearly or quite acaulescent, glabrous or sparsely villous, 1-2 dm. tall; peduncles commonly erect, equalling or exceeding the leaves, racemes 5-8 cm. lofig; flowers 10 mm. long, well-scat- tered; pedicels 4-6 mm. long, glabrous; calyx glabrous, upper lip about as broad as long, 2 mm. long, entire or notched, lower lip about 5 mm. long, entire or obscurely three-toothed; petals blue or purple, the banner with a yellow center, Io mm. long and 10 mm. wide, wings 8 mm. long, 5-6 mm. wide, keel 7-8 mm. long, the point decidedly sees pods oblong, 17-20 mm. long, smooth or scaly on the sides, villous on the margins, ovules two to six, seeds rough, about 3 mm. across, unmarked (?). Fic. 48. Lupinus oporatus Heller. 1. S. B. Parish 4957 (US); 2. A. E. D. Elmer 3624 (US); 3. A. i Heller 8207 (CPS). y This excellent species is of special interest both because of its number of ovules and its limited supply of hair-covering. The former character suggests relationship to certain species of the annuals of the subgenus Lupinus; but its cotyledons are sessile and its relationship is evidently closer to L. pusillus. There should be no further excuse for confusing it with L. brevicaulis. CALIFORNIA. Los Angeles County: Dry Lake, Antelope Valley, May, 1896, J. B. Davy 2223 (UC); Lancaster, June, 1902, Fy 402 SMITH: STUDIES IN THE GENUS LUPINUS A. E. D. Elmer 3624 (B, G, NY, US); Lancaster, May, 1909, K. Brandegee (UC). Kern County: Antelope Valley, May, 1905, F. Grinnell (US); North Fork Kern River, June, 1888, &. Palmer 112 (US); Green Horn Mountains, June, 1888, E. Palmer 112 (T); near Randsburg, May, 1906, H. M. Hall & H. P. Chandler 7285 (UC); Randsburg, May, 1913, K. Brandegee (UC). San Ber- nardino County: Fort Mojave, J. G. Cooper (US); Cajon Pass, east slope, June, 1861, J. C. Cooper (US); Mojave River, June, 1876, E. Palmer 84 (G); Mojave Desert borders, May, 1882, S. B. & W. F. Parish 96 (T), 1269 (G); Mojave Desert, May, 1899, C. J. Pringle (G, US); Hesperia, June, 1901, S. B. Parish 4057 (NY, US); same locality, May, 1892, S. B. Parish 2354 (T); same locality, June, 1895, S. B. Parish. 3774 (G, UC); Mojave, Apr., 1905, Mrs. C. DeKalb (NY); Barstow, May, 1905, H. M. Hall 6166 (B, UC); five miles west of Barstow, March, 1914, Mable Mint- horn (UC); Kramer, Apr., 1905, A. A. Heller 7673 (B, NY, UC); same locality, May, 1909, K. Brandegee (RM, UC); same locality, 1919, K. Brandegee (UC); Blacks Ranch, near Fremont Peak, May, 1906, Hall & Chandler 6843 (B,G, UC); Deadman’s Point, Apr., 1916, S. B. Parish 10787 (UC). Inyo County: near Olancha, Apr., JT. S. Brandegee (UC); Owens Lake, Apr., 1900, S. W. Austin 22 (NY); Laws, May, 1906, A. A. Heller 8207 (B, G, NY, UC); Big Pine, June, 1906, Hall & Chandler 7285 (RM, UC); NEvapDA. County not given: Great Basin, 1875, J. G. Lemmon (US 20973). Washoe County: without definite locality, May, 1875, J. G. Lemmon (G). Arizona. Mohave County: near Kingman, June, 1884, J.G. Lemmon (G); Hackberry, May, 1884, M. E. Jones 66 (G). 106. Lupinus odoratus pilosellus var. nov. A L. odorati differt caulibus petiolisque conspicue _pilosis: leguminibus oblongis, 2 cm. longis; seminibus 2-4. Stems and petioles conspicuously pilose with short white spreading hairs; pods oblong, 2 cm. long, two- to four-seeded. CALIFORNIA. San Bernardino County: Nipton, June, 1905, K. Brandegee (Type, UC 180117); Mohave River, May, 1876, E. Palmer 84 (UC 82240). SMITH: STUDIES IN THE GENUS LUPINUS 403 Ita. LupINUS RUBENS Rydberg, Bull. Torrey Club 34: 45. 1907. [Fic. 49.] Branched at or near the base, 6-15 cm. tall, villous, axial peduncle usually erect, commonly flowering. before the branches develope, racemes exceeding the foliage; flowers 6-12 mm. long, approximate or well scattered; pedicels 1-2. mm. long, usually glabrous but sometimes more or less villous; calyx-lips setose- villous, the cup commonly free of hairs, upper lip entire or notched, lower lip entire or two- or three-toothed; petals as in L. odoratus, the banner with a yellow center; pods ovate, 8-12 mm. long, - usually two-seeded, more or less villous on the sides; seed charac- ters yet to be determined. This is a variable species intermediate between L. pusillus and L. odoratus. One extreme, when in flower, is hardly separable from the New Mexican variation of L. pusillus, while other forms approach closely to L. odoratus. Critical cases will evidently have to be judged in terms of fruiting material. Fic. 49. Lupinus RUBENS Rydberg. 1. M. E. Jones 5138 (US); 2. M. E. Jones 4413 (US); 3. M. E. Jones 5024i (US); 4. E. D. Ball (CPS); 5. J. G. Cooper (US 344379). Utan. Emery County: Green River, May, 1890, M. E. Jones (US); same locality, May, 1909, Z. D. Ball (CPS). Grand County: Grand River Canyon below Moab, July, ro11, collector _ hot given (NY). Kane County: Kanab, Mrs. A. P. Thompson ( Washington County: St. George, Apr., 1880, M. E. Jones 1657 (NY, RM, T, US); same locality, Apr., 1884, M. E. Jones 5110] (US); Santa Clara Valley, Apr., 1884, M. E. Jones 5138 (NY, UC, US). County not ascertained: southern Utah, 1872, 2. 404 SMITH: STUDIES IN THE GENUS LUPINUS Capt. Bishop (US), 1873, Capt. Bishop (G), 1874, C. C. Parry 4, 42 (7). or. ArIzoNA. Navajo County: six miles northeast of Winslow, May, 1go1, L. F. Ward (US). Coconino County: ten miles north of Tuba, May, tgo1, L. F. Ward (US). Mohave County: Hack- berry, May, 1884, M. E. Jones 4413 (US). County not deter- mined: no locality, 1869, E. Palmer (US 20944, two upper speci- mens); Beaverdam, Apr., 1894, 7. E. Jones 50241 (US) Nevapa. Clark County: Cottonwood Springs, Vegas Valley, 1891, V. Bailey 1876 (US). Lincoln County: Caliente, Meadow Valley Wash, May, 1902, L. N. Goodding 942 (RM). County not given: Dry Lake, Apr., 1905, L. N. Goodding 2234a (G, NY, RM, UC). . CALIFORNIA. Inyo County: Wild Rose Springs, Panamint Mountains, May, 1915, S. B. Parish (UC). San Bernardino County: Cajon Pass, 1860-1861, J. G. ‘Cooper (RM, left-hand specimen); Providence Mountains, near summit, May, 1861, J. G. Cooper (US); New York Mountains, June, 1915, S. B. Parish 10073 (UC); Good Springs, May, 1915, K. Brandegee (UC); Cima Station, May, 1915, K. Brandegee (UC 180152, upper specimen). 11d. Lupinus rubens flavoculatus (Heller) comb. nov. [FiG. 50.] Lupinus flavoculatus Heller, Muhlenbergia 5: 149. 1909. Differs from L. rubens proper in the early development of floriferous, widely spreading branches, in the longer pedicels (1-4 mm.), and in the lower calyx-lip being usually two-toothed. This much resembles various specimens of L. odoratus, but seems to be fairly constant in its differences as to pubescence and fruit. It varies easily into that variable species, and my disposition of some of the specimens cited here is not beyond fair questioning. One of the specimens (UC 149911) is evidently a pathological variation. Nevapba. Nye County: Rhyolite, May, 1907, W. H. Shockley 43 (UC); Rhyolite, May, 1909, A. A. Heller 9669 (B, NY, RM, UGC, CPS). CALIFORNIA. Inyo County: Darwin Mesa, Yucca Valley, May, 1891, F. V. Coville & F. Funston 877 (US); Boundary Can- yon, Grapevine Mountains, June, 1891, F. V. Coville & F. Funston 977 (G, T, US); Lone Pine, May, 1913, K. Brandegee (UC). SMITH: STUDIES IN THE GENUS LUPINUS 405 San Bernardino County: Barnwell, May, 1911, K. Brandegee (UC 149898, 149909-10-11); Cima Station, May, 1915, and Lp Wy = Fic. 50. LUPINUS RUBENS FLAVOCULATUS (Heller) C. P. Smith. A.A. Heller 669 (CPS June, 1915, K. Brandegee (UC). Kern County: South Fork Kern River, 1898, C. A. Purpus 5714 (UC, US). Modoc County: Surprise Valley, May, 1879, J. G. Lemmon (G). 12a. Lupinus pusiILLus Pursh, Fl. Am. Sept. 2: 468. 1814. [F1G. 51.] Stems well-developed, branches ascending to widely-spreading, loosely villous, 1-2 dm. tall; peduncles 1-2 cm. Jong, racemes 3-5 cm. long, mainly equalled or exceeded by the leaves; flowers 10- 12 mm. long, well scattered; pedicels about 2 mm. long, villous; calyx villous, upper lip broader than long, 2 mm. long, the short teeth acute or rounded, lower lip over twice as long as wide, 5-6 mm. long, the apex entire and acuminate, acute, or rounded, or occasionally three-toothed; petals Blatahy purplish, or almost Fic. 51. Lupinus pusit_us Pursh. 1. V. Bailey ro (US); 2. M. E. Moodie 946 (US); 3. A. Nelson 8258 (US); 4. E. Palmer 82 (US 291129). white, banner 10-11 mm. long and 9-10 mm. wide, wings 8-9 mm. long, 5-6 mm. wide, keel 8 mm. long, 3.5 mm. wide, the point somewhat upturned; pods 20-25 mm. long, lance-oblong, acute at the apex, slightly constricted near the middle, two-seeded; seeds about 4 mm. across, rugose, apparently unmarked. 406 SMITH: STUDIES IN THE GENUS LUPINUS The typical form of this species is mainly confined to the Atlantic drainage of the Rocky Mountain region, the exception being its occurrence on the “Snake Plains’’ of southern Idaho. The New Mexican plants referred here may prove to be a separable variety, as the racemes often exceed the foliage and the plants suggest the larger phases of L. rubens. Fruiting material, with matured seeds, is much desired from New Mexican localities and Green River, Utah. The Great Basin form, separated by Heller, deserves recognition as a variety, some intermediates occurring. ALBERTA. Bow Valley County: West Fork Water Coulee, near Rosedale, June, 1915, M@. E. Moodie 946 (G, US). Medicine Hat County: Medicine Hat, May, 1894, J. Macoun 4069 (G). ALBERTA or SASKATCHEWAN. .West of South Saskatche- wan, June, 1879, J. Macoun tog (G); north of Cypress Hills, Aug., 1880, J. Macoun (T); Cypress Hill, July, 1895, J. Macoun 10412 (G). IpAHO. Elmore County: King Hill, July, 1911, Nelson & McBride 1111 (RM). Ouyhee County: Murphy, July, rort, Nelson & McBride 1030 (RM). Bingham County: Blackfoot, July, 1893, E. Palmer 275 (US). Lincoln County: Bluelakes, June, 1893 E. Palmer 82 (US). Montana. Cascade County: Great Falls, July, 1885, R. S. Wiliams 214 (US); Great Falls, Sept., 1886, F. W. Anderson (UC); Falls of the Missouri, July, 1887, F. W. Anderson 1422 (NY). Dawson County: near Glendive, 1883, L. F. Ward (US); Glendive, June, 1908, B. T. Butler 2005, 2006 (NY). Still Water County: Still Water, July, 1889, F. Tweedy 3 (NY). Cus- ter County: Miles City, June, 1893, F. E. Lloyd (NY). Yellow- stone County: Custer, May, 1890, J. W. Blankinship (UC). Park County: Upper Yellowstone, July, 1899, J. W. Blankinship (G, RM). Locality not given: 1890, P. A. Rydberg 397 (NY); June, 1890, J. W. Blankinship 156 (US). YELLOWSTONE Park. Yellowstone, V. Havard (US); eleven miles above Glendive, July, 1883, L. F. Ward (US); mouth of Gardiner River, F. Tweedy (US). Wyominc. Albany County: Laramie River, Laramie Peak, 1864, R. B. Hetz (US). Converse County: Lost Springs, July, 1915, W. L. Hees 113 (RM). Crook County: Pine Ridge, July, SMITH: STUDIES IN THE GENUS LUPINUS 407 1909, M. Cary 365 (US). Diobrara County: Van Tassell, Doug- las, June, 1899, H. M. Barrow 35 (RM). Johnson County: Buffalo, 23 July and 26 July, 1898, 7. A. Williams (US). Laramie County: Fairbanks, July, 1894, A. Nelson 490 (G, RM, T, UC, US). Natrona County: Alcova, July, 1901, L. N. Goodding 157 (RM). Platte County: Wheatland, June, 1901, A. Nelson 6258 (B, G, NY, RM, UC, US). Washakie County: Worland, June, 1909, B. C. Buffum (RM). County not given: 1893, B. W. Evermann (US); four miles below U. L. Ranch, July, 1896, F. H. Knowlton 159 (US); Sweetwater River, July, 1898, E. Nelson 4989 (CAC, RM); Cheyenne River, E. J. Wallace (RM). : CoLtorapo. Denver County: Denver, May, 1873, J. M. Coulter (US); Denver, June, 1873, J. Wolf 20 (US); Denver, _ Platte River, June, 1878, M. E. Jones 199 (NY); Denver, July, - 1885, G. W. Letterman (US). Fremont County: Canon City, 1872-1873, 7. S. Brandegee (UC). Larimer County: LaPorte, June, 1895, J. H. Cowen 149 (G, NY); Ft. Collins, W. F. Marshall 734 (CAC, RM, US); Fort Collins, June, 1898, C. S. Crandall (RM); Fort Collins, June 1915, J. H. Cowen 113 (US), same, June, 1893 (G). Lincoln County: Hugo, July, 1905, C. D. Marsh (US). Ouray County: Ridgeway, Aug., 1894, F. Tweedy 240 (US); Uray [Ouray], May, 1900, J. E. Payne (CAC). Prowers County: Lamar, May, 1892, C. S. Crandall (CAC). Pueblo County: Pueblo, June, 1891, E. Penard (T): Pueblo, June, 1897, collector not given (CAC); Pueblo mesas, May, 1900, P.A. Rydberg & F. V. Vreeland.5941 (NY, RM). Weld County: Crow Creek, June, 1896, F. H. Knowlton 97 (NY, US); New Windsor, June, 1895 and 1901, G. E. Osterhout (NY);same locality, June, 1905, G. E. Osterhout (G, RM). County not given: June, 1970, E. L. Greene 67 (G); June, 1890, C. H. Hall (B). NortH Daxota. Barnes County: Valley City, Cheyenne Valley, July, 1903, M. A. Barber 176 (G). Billings County: Marmarth, July, 1909, V. Bailey ro (US). Morton County: Parkin, June, 1916, V. Bailey (US). County not given: Bad Lands, July, 1891, H. L. Bolley 132 (RM). SoutH Dakota. Fall River County: without definite locality, Aug., 1911, S. S. Visher 2703 (RM); Fall River Falls, June, 1892, 408 SMITH: STUDIES IN THE GENUS LUPINUS P. A. Rydberg 599 (US). Harling County: Steppe, July, 1910, S. S. Visher 231 (RM). Meade County: Smithville, June, 1894, V. Bailey to (US). Perkins County: Bixby, July, 1912, S. S. Visher 615 (RM). Washington County: Sheep Mountain and sand hills south, July, 1911, S. S. Visher 2406 (NY). County not given: near White Earth River, Missouri, J. N. Nicolet 161 (G); north edge of sand bluff, S. A. Skinner 177 (RM). NEBRASKA. Brown County: Longpine, June, 1897, J. M. Bates (RM). Cherry County: Valentine, June, 1891, J. ,M. Bates (G, T). Frontier County: near Curtis, June, 1891, P. A. Rydberg 46 (NY). Keith County: without definite locality, June, 1890, G. D. Swezey (T). Dawes County: Crawford, June, 1891, J. M. Bates (B). Kansas. Finney County: Garden City, May, 1891, H. W. Menke (T). Graham County: without definite locality, June, 1897, A. S. Hitchcock 81a (G, NY, RM, US). Logan County: - without definite locality, May, 1895, A. S. Hitchcock 81 (G, NY, RM, US). County not given: western Kansas, 1885, E. N. Plank 45. te New Mexico. San Juan County: Aztec, May, 1899, C. F. Baker 435 (G, NY, RM,* US). Dona Ana County: near Organ Mountains, Apr., 1852, Mexican Boundary Survey 287 (NY, two sheets); opposite Frontera, March, 1852, C. Wright 1362 (US 20950-1). Without locality: Mex. Boundary Survey (US 20941, upper and left-hand specimens); 1851, C. Wright 1302c (G); May, 1901, F. K. Vreeland 821 (NY). ARIZONA. Pimta County: Tucson, March, 1852, c C. Parry (T). Without locality: 1869, E. Palmer (US 20944, two upper specimens). 12b. Lupinus pusillus intermontanus (Heller) comb. nov. [Fic. 52 Lupinus intermontanus Heller, Muhlenbergia 8:87. 1912. * The ‘‘RM”’ sheet of this collection has Baker’s number given as ‘‘ 433,” is labelled L. aduncus Greene, whence Heller has annotated this sheet as follows: ““Apparently immature L. rubens Rydb. Baker has mixed labels. L. aduncus is a perennial. I have the real thing here now, also under 433.’’ It is evidently a part of Baker’s collection 435, which seems to have been originally determined as L. arizonicus. I may be wrong in assigning it to L. pusillus a of to L. rubens. Matured plants with pods and seeds are needed for 7 SMITH: STUDIES IN THE GENUS LUPINUS 409 Plants usually more congested than in the species; flowers appreciably smaller, 7-9 mm. long; pedicels and calyx-cup glabrous or sub-villous, lips villous, lower lip less than twice as long as broad, 3-4 mm. long, entire or three-toothed at apex; banner 7-8 mm. long, 5-6 mm. wide, wings about 3 X 6 mm., keel 6 mm. long, straight or the acumen slightly upturned. CALIFORNIA. Modoc County: Surprise Valley, May, 1879, J..G. Lemmon (G). Inyo County: Deep Spring Valley, 1898, C. A. Purpus 5807 (UC, US). NevapA. Clark County: Jean, May, 1915, K. Brandegee (UC). Esmeralda County: Candelaria, W. H. Shockley 270 (UC). Humboldt County: Unionville, June, 1868, S. Watson 227 in part (US). Ormsby County: Carson City 1864, C. L. Anderson 84 Fic. 52. Lupinus PUSILLUS INTERMONTANUS (Heller) C. P. Smith. 1. A. A. Heller 0500 (CPS); 2. Merrill & Wilcox 725 (US). (G); Empire City, June, 1882, M. E. Jones 3813 (NY, US). Washoe County: Wadsworth, May, 1909, A. A. Heller 9599 (B, G, RM, UC, US, CPS); same locality, June, 1913, K. Brandegee (UC). OrEGON. Harney County: Alvord Desert, June, 1896, J. B. Leiberg 2432 (US); same locality, July, 1898, W. C. Cusick 2020 (G, RM, UC, US). Lake County: Drews Valley, July, 1893, Mrs. R. M. Austin (UC). WASHINGTON. Franklin County: Pasco, June, 1892, L. F. Henderson 2362 (G); same locality, May, 1899, C. V. Piper 2982 (G. NY). Walla Walla County: Wallula, May, 1883, T. S. Bran- degee 706 (G) ; same locality, May, 1896, T. Howell 1920 (NY); same locality, May, 1903, J. S. Cotton 1028 (US); Walla Walla, Capt. Wilkes 955 (US 21026, right-hand specimen only*); same locality, May, 1903, J. S. Cotton 1028 (G); plains of the Columbia River, * The left-hand specimen is L. subvexus fluviatilis, in fruit. 410 SMITH: STUDIES IN THE GENUS LUPINUS 1826 ex Herb. Benth. (G). Yakima County: Yakima River near Morgan’s Ferry, June, 1884, W. N. Suksdorf 269 (G); west of Mabton, June, 1903, J. S. Cotton 1115 (US). IpAHO. Owyhee County: Murphy, July, 1911, Nelson & McBride 1030 (RM). Twin Falls County: Twin & Shoshone Falls, July, 1911, Nelson & McBride 1333 (RM). Wyominc. Big Horn County: Greybull, May, 1910, M. Cary 521 (US). ? Sweet Water County: thirty-five miles north of Point of Rocks, June, 1901, E. D. Merrill & E. N. Wilcox 611 (G, US); near Washington Ranch, June, 1901, Merrill & Wilcox 725 (G, RM, US). UtaH. Weber County: Ogden, 1872, Hayden Survey (US). Beaver County: Frisco, June, 1880, M. E. Jones (US). Kane County: Kanab, May, 1894, M. E. Jones 5286f (NY, UC, US); twenty miles south of Pahria, May, 1894, M. E. Jones 5292c (US). : ARIZONA. Coconino County: Moqui, May, 1869, E. Palmer (G). Cotorapo. Huerfano County: Walsenburg, July, 1896, C. L. Shear 4753, 4800 (NY). It is interesting to note that this small-flowered variety is almost entirely confined to the Pacific and Great Basin drainage. Grass rusts of unusual structure* J. C. ARTHUR AND E. B. MAINS (WITH TWO TEXT FIGURES) The leaves of species of Olyra belonging to the tribe Paniceae and of species of bamboos are frequently much alike in form and physical characters, although systematists place the genera nearly at the extremes of the grass series. The resemblances are often so great that only an experienced agrostologist can feel sure of their identity when not accompanied by inflorescence. It is not surprising, therefore, that the rusts on these genera, as well as their hosts, should have been confused. Thus Uredo Olyrae P. Henn., described in 1904, to which true Olyra rusts have been referred, has recently been found to be on some species of the bamboo genus Arundinaria, and Uredo ignava Arth. on the genus Bambos has only recently been separated from material formerly placed with paniceous rusts. Not only is there a curious resemblance between the leaves of these two groups of hosts, but also an equally interesting re- semblance between some of the rusts that inhabit them. This was first noticed in the uredinia having delicate incurved para- physes and spores that are more or less thin-walled and colorless. Later it was found that the presence or absence of these weakly differentiated paraphyses is not by itself an important diagnostic character, as the forms called Uredo paspalicola without paraphyses and U. Stevensiana with paraphyses have been shown to be only extremes of one species having all degrees of intergradation. In the study of this sort of rusts it became more and more necessary to use a razor and make sections of the sori in order to be surer of their structure. In this manner the paniceous species, Puccinia tubulosa, P. inclita, and P. imposita, were segregated and gradually circumscribed from what had been lumped as P. sub- * Reprints may be obtained by application to the Botanical Department, Pur- due University Agricultural Experiment Station, Lafayette, Indiana, under whose auspices the studies here reported were carried out. 411 412 ARTHUR AND MAINS: GRASS RUSTS striata. Then the telia were found in P. Chaseana on a host belonging to a family preceding the Paniceae. The final triumph came in the discovery of telia with the much-studied form, Uredo pallida (Puccinia pallescens) on Tripsacum, a grass belonging to the tribe Maydeae. These telia would doubtless not have been discovered through the usual method of examination by scraping the spores from the leaf with a scalpel. Sections show the sori to be small, indehiscent, and filled with pale, compacted spores (Fic. 2A) that are thin-walled and practically without pedicels. Many of the teliospores are three-celled, especially toward the center of the sorus. The whole appearance of the telia and telio- spores is quite unlike that of the ordinary grass rusts, and reminds one of those in the group of melampsoraceous rusts represented by Phakopsora. Even with this experience the telia of still another species of rust on Olyra latifolia were passed over fora time. The uredinia had been referred to Puccinia deformata Berk. & Curt., on the same host, a species with prominent telia, known for over fifty years, but with no described uredinia.* Thin-walled, nearly colorless urediniospores, surrounded by incurved paraphyses, were found, and on the same leaves were discovered telia by the sectioning method. So unusual was the appearance of these telia, however, that they were tentatively referred to the genus Phaksopsora, and might have remained there longer had the grasses been known to harbor species of this genus. It was then thought there might have been an error in determination of the host, and that it was a bamboo, instead of the Olyra. After much study the following description of a new species of rust was evolved, and incidentally the true urediniospores of P. deformata were found. Puccinia phakopsoroides sp. nov. II. Uredina amphigenous, small, round, cinnamon-brown, early naked, ruptured epidermis inconspicuous; paraphyses peripheral, incurved, clavate, 10-12 by 35-50 u, the wall cinna- mon-brown, I yu thick, usually thickened up to 2-3 u on the convex side; urediniospores ellipsoid or broadly obovoid, 19-26 by 27-35 #3 wall slightly brownish or nearly colorless, 1-1.5 « thick, closely *See Arthur & Johnston, Uredinales of Cuba. Mem. Torrey Club 17: 136. 1918. ARTHUR AND MAINS: GRASS RUSTS 413 echinulate, the pores very indistinct, probably 2 and equatorial. III. Telia hypophyllous, scattered, squarish-oblong, 0.5—1 mm. long, covered by the epidermis, compact with the spores adhering laterally, brownish-black due to discolored host-tissue; teliospores I-, 2- or 3-celled, short-cylindric, 8-14 by 24-39 un, rounded or depressed above and below, somewhat constricted at septa; wall pale-brown or colorless, smooth, thin, uniformly 1 u or less in thickness, but with an outer colorless layer swelling in in water to 2-3 uw above, demonstrable by staining; pedicel wanting or extremely short and colorless. On Olyra latifolia L., Mayagiiez, Porto Rico, January 30, 1890, II, III, A. A. Heller 4443, communicated by P. L. Ricker; San German, Porto Rico, December 12, 1913, II, F. L. Stevens 5849, 5855; Manati, Porto Rico, July 2, 1915, II, F. L. Stevens 7700; Soledad, Cienfuegos, Cuba, November 7, 1915, ii, J. R. Johnston 247; Isle of Pines, Cuba, February 20, 1916, II, Britton, Britton & Wilson 14648; Ceballos, Cuba, March 24, 1916, II, J. R. Johnston 509; Guantanamo, Cuba, February 7, 1918, II, Ill, J. R. Johnston 1028 (TYPE). Fic. 1. Puccinia phakopsoroides: A, section showing part of a telial sorus and the adjoining host tissues; B, teliospores after treatment with potassic hydrate. X 625 diameters. There is a marked gradation evident in some tropical rusts, especially the grass rusts as indicated above, toward the produc- tion of thin-walled, pale or colorless spores. In the uredinia these spores are frequently accompanied with a fringe of incurved, hyphoid paraphyses. In the telia the spores are small and often compacted, long remaining covered by the epidermis. As usual in subepidermal forms, these teliospores have short pedicels. 414 ARTHUR AND MAINS: GRASS RUSTS . Puccinia phakopsoroides illustrates an extreme development in this general direction, most strongly seen in the telia. Drawings of the telial sorus of both P. phakopsoroides and P. pallescens (Fics. 1A, 2A) show the spores as cylindrical bodies pressed very closely together, as if restricted by the unyielding epidermis. The spores appear to be less highly differentiated than in most grass rusts, having a variable number of cells with a quite uniform wall, and a pedicel so short and delicate that one is left in doubt whether there is a truly pedicellate cell, or only a hyphal connec- tion with the hymenial layer of the fungus. Sections of the sorus give a superficial resemblance to forms of Phakopsora, especially in the denseness, the pale color and trans- lucency, and in the evenly disposed, vertical rows of apparently similar cells, more cells being in a row at the center of the sorus Soe Aq ' 7s Cee py iS, =~" if i. B Fic. 2. A, section showing part of a telial sorus of Puccinia pallescens, with adjoining host tissues; B, a teliospore of P. phakopsoroides treated with eosin to show the outer hygroscopic layer of the spore-wall. 625 diameters. than at the edge. In Puccinia each row of cells forms a single spore; in Phakopsora there are as many independent spores as there are cells. In P. phakopsoroides this resemblance is em- phasized when the sections are treated with potassic hydrate, and the spores pressed out of the sorus (Fic. 1B). The cells then seem ARTHUR AND MAINS: GRASS RUSTS 415 to separate and appear like one-celled spores in rows, the envelop- ing outer layer of the spore-wall being invisible, having been swollen and partly dissolved by the potash. However, if sections are stained with eosin, and‘.the spores pressed apart, it is easy to discern a delicate outer layer of wall, swelling in water and be- coming thicker above than at the sides of the spore (Fic. 2B). The presence of this hygroscopic layer explains why the cells remain in chains, even after macerating in potash, and doubtless why the cells adhere laterally with so much tenacity in an ordinary mount. In P. pallescens there is no such hygroscopic layer. In the above paragraphs comparison has been drawn especially between three tropical rusts of similar but extreme development, which are found on grasses having slight relationship, according to the classification used by modern agrostologists: Puccinia pallescens on Tripsacum, the first genus in Hitchcock and Chase’s “Grasses of the West Indies,’’ P. phaksopsoroides on Olyra, the fifty-second genus in the same work and Uredo ignava on Bambos, the one hundred and tenth and last genus of that work. The telia of the last species have not yet been found, but it is con- fidently believed that when discovered they will resemble those of the other two species named. PURDUE UNIVERSITY, AFAYETTE, INDIANA INDEX TO AMERICAN BOTANICAL LITERATURE 1917-1919 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, iidicdtiare, pc aaa products of vegetable origin, or laboratory methods are not included, and attempt is made to index the literature of bacteriolo ogy. An occasional exception is ‘si 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 spondence relating to the card issue should be addressed to the Treasurer of the Torrey Botanical Club Abrams,L.R A new California Cypress. Torreyatg: 92. 9 Jl 1919. Cupressus nevadensis sp. nov. Arthur, J.C. Acorrection. Torreya 19: 83. 25 Je 1919. Syntherisma pruriens, error in publication. Ashe, W. W. Notes on trees and shrubs in the vicinity of Washington. Bull. Torrey Club 46: 221-226. 8 Jl 1919. Amelanchier sera and A. micropetala (Robinson), spp. nov., are described. Bailey, I. W. Structure development, and distribution of so-called rims or bars of Sanio. Bot. Gaz. 67: 449-468. pl. 13-15. 21 Je I9I9. Barnhart, J. H. Brackenridge and his book on ferns. Jour. N. Y. Bot Gard. 20: 117-124. Je 1919. Benedict, R. C. The simplest fern in existence. Am. Fern Jour. 9: 48-50. pl. 3. Jl igrog. Bigelow,H. C. A fern from the battle-ground. Am. Fern Jour. 9: 58. Jl 1919. Bisby, G. R. Studies on Fusarium diseases of potatoes and truck crops in Minnesota. Minnesota Agr. Exp. Sta. Bull. 181: 1-47 f. I-30. Mr 1919. Blakeslee, A. F. A unifoliolate mutation in the Adzuki bean. Jour. Heredity 10: 153-155. f. 1, 2. 26 My 1919. 417 418 INDEX TO AMERICAN BOTANICAL LITERATURE Boynton, K. R. Eupatorium coelestinum. Addisonia 4: 39, 40. pl. 140. 30 Je 1919. Britton, N. L. & Rose, J. N. The Cactaceae. Descriptions and illus- trations of plants of the Cactus family. i-vii + 1-236. pl. 1-36 + f. t-3or. Washingtan. 1919. Carnegie Publ. No. 248, Vol. I; includes the new genus Tacinga and new species in Opuntia (30), Pereskia (4), Pterocactus (3), Nopalea (1), and Tacinga (1). Burt, E. A. Merulius in North America, supplementary notes. Ann. Missouri Bot. Gard. 6: 143-145. Ap rg1g. Clark, H. W. Dwarf shore floras. Trans. Illinois Acad. Sci. 10: 145- 159, pl. 1-5. 1917. Clayton, E.E. Hydrogen cyanide fumigation. Bot. Gaz. 67: 483-500. fe 2, 22:22% Je toto. Clute, W. W. Defining double flowers. Gard. Chron. Am. 23: 189. Je 1919. Cockerell, T. D. A. Sir Joseph Hooker. Science II. 49: 525-530. 6 Je 1919. Collins, G. N. A fossil ear of maize. Jour. Heredity 10: 170-172. f.7. 26 My 1919. Coville, F. V. The threatened extinction of the box huckleberry, Gaylussacia brachycera. Science II. 50: 30-34. 11 Jl 19109. Dalbey, N. E. Phyllachora as the cause of a disease of corn, and a general consideration of the genus Phyllachora. Trans. Illinois Acad. Sci. 10: 230-248. f. 3-9. 1917 Duggar, B. M., & Davis, A. W. Seed disinfection for pure culture work: the use of hypochlorites. Ann. Missouri Bot. Gard. 6: 159-170. Ap I9gI9. Ensign, M. R. Gas injury to wheat. Phytopathology 9: 266. 24 Je , I9I9Q. Evans, A. W. A new Riccia from Peru. Torreya 19: 85-88. f. I- 9 Jl 1919. Riccia bistriata sp. nov. Falk, K. G., McGuire, G., & Blount, E. Studies on enzyme action— XVII. The oxidase, peroxidase, catalase, and amylase of fresh and dehydrated vegetables. Jour. Biol. Chem. 38: 229-244- Je 1919. Farwell, O. A. Tsuga americana (Mill.) Farwell, a final word. Rho- dora 21: 108, 109. Je 1919. INDEX TO AMERICAN BOTANICAL LITERATURE 419 Fernald, M.L. Panicum § Capillaria in New England. Rhodora 21: 110-114. Je 1919. Paricum Tuckermani sp. nov. is described. Fromme, F. D., & Wingard,.S. A. Bean rust. Virginia Agr. Exp. Sta. Bull. 220: 1-18. pl. 1-5. N 1918. Gleason, H. A. What is ecology? Torreya 19: 89-91. 9 Jl 1919. Graves, E. W. The botrychiums of Mobile County, Alabama. Am. Fern Jour. 9: 56-58. Jl 1919. Griffiths, D. New and old species of Opuntia. Bull. Torrey Club 46: 195-206. pl. 9g, 10. 8 Jl 1919. Eight new species are described. Giissow, H. T. Establishment of an imperial (British) Bureau of Mycology. Phytopathology 9: 265. 24 Je 1919. Haas, A.R.C. Anesthesia and respiration. Science II. 46: 462-464. 9 N 1917. , Harper, R. M. A forest reconnaissance of the Delaware Peninsula. Jour. Forestry 17: 546-555. My 1919. [Map] Haskell, R. J. Fusarium wilt of potato in the Hudson River valley, _ New York. Phytopathology 9: 223-260. pl. 13-15. 24 Je 1919. Higgins, B. B. Gum formation with special reference to cankers and decays of woody plants. Georgia Agr. Exp. Sta. Bull. 127: 23-51. pl. 1-6. + f. 1, 2. . Ja 1919. Higley, R. Notes on Cephaleuros virescens. Trans. Illinois Acad. Sci. 10: 256-258. 1917. Hughes, F. T. Botany in the city high schools. Torreya 19: 57-65. 25 Je 1919. Hunnicutt, B. H. A forage plant from the Solanaceae family. Jour. Heredity 10: 185-187. f. 14, 15. 26 My 1919. Johnson, J. The influence of heated soil on seed germination and plant growth. Soil. Sci. 7: 1-103. pl. 1-8. Ja 1919. Johnson, J., & Hartman, R. E. Influence of soil environment on the rootrot of tobacco. Jour. Agr. Research 17: 41-86. pl. 1-7. 15 My I9IQ. King, C.A. Changes in teaching biology in our high schools. fee 19: 65-71. 25 Je 1919. Kylin, H., & Skottsberg, C. Zur Kenntnis der subantarktischen und antarktischen Meeresalgen—II. Rhodophyceen. Wissenschaft- liche Ergebnisse der Schwedischen Siidpolar Exp. 1901-1903. 4: I-88. pl. 1 + f. 1-38. 1919. 420 INDEX TO AMERICAN BOTANICAL LITERATURE Lillo, M. Segunda contribucion al conocimiento de los arboles de la Argentina. 1-69. Tucuman. 1917. Long, B. Jasione montana a conspicuous weed near Lakewood, New Jersey. Rhodora 21: 105-108. Je 1919. Lutman, B. F. Osmotic pressures in the potato plant at various stages of growth. Am. Jour. Bot. 6: 181-202. f. 1, 2. My 1919. MacDougal, D. T. Growth in organisms. Science II. 49: 599-605: 27 Je 1919. Mackenzie, K. K. LEupatorium maculatum. Addisonia 4: 23, 24. pl. 132. 30 Je 1919. Mann, P. B. The relation of first year botany to advanced work, with reference to certain applications and by-products. Torreya 19: 72-78. 25 Je 1919. Maxon, W. R. Ferns of the District of Columbia. Am. Fern Jour. 9: 38-48. Jl 1919. Maxon, W. R. A new Alsophila from Guatemala and Vera Cruz. Proc. Biol. Soc. Washington 32: 125-129. 27 Je 1919. Alsophila scabriuscula sp. nov. Macinnes, F. J. The occurrence of Alternaria in a characteristic apple spot, and an apple rot caused by Gliocladium viride. Trans. Illinois Acad. Sci. 10: 218-229. pl. 1-4. 1917. McAtee, W.L. Summary of notes on winter blooming at Washington, D.C. Proc. Biol. Soc. Washington 32: 129-132. 27 Je 1919. McLean, R. C. Studies in the ecology of tropical rain-forests: with ' special reference to the forests of south Brazil. Jour. Ecol. 7: 5-54. pl. 1 +f. 1-27. My 19109. Melhus, I. E., & Rhodes, L.L.. A quick method of eliminating seed- borne organisms of grain. Science II. go: 21. 4 Jl 1919. Miles, L. E. Some new Porto Rican fungi. Trans. Illinois. Acad. Sci. 10: 249-255. f. I-3. 1917. Includes new species in Mycosphaerella (9), Helminthosporium (1), Cercospora (1). Nash, G. V. Crataegus macrosperma. Addisonia hc 38. Pl 138, 30 Je 1919. Nash, G. V. The digger pine in the botanical garden. Jour. N. Y. Bot. Gard. 20: 127, 128. Je 1919. Nash, G.V. Malus Halliana. Addisonia 4: 27. pl. 134. 30 Je 1919. Nash, G. V. Oxydendrum arboreum. Dh stinaecnas 4: 37; 38. pl. 139- 30 Je 1919. INDEX TO AMERICAN BOTANICAL LITERATURE 421 Payson, E. B. The North American species of Agquilegia. Contr. U.S. Nat. Herb. 20: 133-157. pl. 8-15. 1919. Four new species are described. Pellegrin, F. Quelques remarques sur les Dioscoréacées du Paraguay. Bull. Soc. Bot. Genéve 10: 383-388. 1918. Pennell, F. W. Penstemon calycosus. Addisonia 4: 31, 32. pl. 136. 30 Je 1919. Pepoon, H. S. The primrose rocks of Illinois. Trans. Illinois Acad. Sci. 10: 159-162. 1917. [Illust.] Petch, T. Citrus mildew: a correction. Phytopathology 9: 266. 24 Je 1919. Pipal, F. J. White top and its control. Purdue Univ. Agr. Exp. Sta. Circ. 85: 1-12. f. 1-8. D 1918. Pring, G. H. Orchids of the Missouri Botanical Garden, St. Louis. Mo. Gard. Chron. Am. 23: 208, 209. Je 1919. [Illust.| Roberts, H. F. The founders of the art of breeding—II. Jour. Heredity 10: 147-152. f. 1 + frontispiece. 26 My 1919. Roberts, R. H. ‘‘Crinkle’’ on northwestern greening. Phytopath- ology 9: 261-263. pl. 16,17. 24 Je 1919. Rock, J. F. Cyrtandreae Hawaiienses, sect. Microcalyces Hillebr. Am. Jour. Bot. 6: 203-216. pl. 29-32. My 1919. Cyrtandra Giffardii, C. limosiflora, C. montis Loa, C. ramosissima, and C. Hashi- motoi, spp. nov., are described wee Sayre, J. D. Comparative transpiration of tobacco and mullein. Ohio Jour. Sci. 19: 422-426. f. 1. My 1919. Schaffner, J. H. The nature of the dioecious condition in Morus alba and Salix amygdaloides. Ohio Jour. Sci. 19: 409-416. My 1919. Schmitz, H. Studies in the physiology of the fungi—VI. The relation of bacteria to cellulose fermentation induced by fungi, with special reference to the decay of wood. Ann. Missouri Bot. Gard. 6: 93-136. Ap Igig. Selby, A. D. Apple blotch, a serious fruit disease. Bull. Ohio Agr. Exp. Sta. 333: 491-505. f. 1-5. F 1919. Seward, A.C. Recent paleobotany in Great Britain. Science II. 50: 43-48. 11 Je 1919. Small, J. K. Heliotropium Leavenworthii. Addisonia 4: 29, 30. pl. 135. 30 Je 1919. Small, J. K. Heliotropium polyphyllum. Addisonia 4: 25, 26. pl. 133- 30 Je 1919. 492 INDEX TO AMERICAN BOTANICAL LITERATURE Small, J. K. Rhabdadenia corallicola. Addisonia 4: 33, 34. pl. 137- 30 Je 1919. Small, J. K. Urechites pinetorum. Addisonia 4: 21, 22. pl. 131. 30 Je 1919. Smith, E. F., Jones, L. R., & Reddy, C.S. The black chaff of wheat. Science II. 59: 48. 11 Je 1919. Southworth, W. Twining in alfalfa. Jour. Heredity 10: 182. f. 12, 13- 26 My 1919. Stakman, E. C. Destroy the common barberry. U.S. Dept. Agr. Farm. Bull. 1058: 1-12. f. 1-6. My 1919. Steil, W. N. Apospory in Pieris suleata L. Bot. Gaz. 67: 469-482. pl. 16, 17. 21 Je 1919. Stevens, F.L. An apple canker due to Cytospora. Illinois. Agr. Exp. Sta. Bull. 217: 367-379. f. 1-15. Mgitgr1g. [Illust.] Stevens, F.L. Porto Rican fungi, old and new. Trans. Illinois Acad. Sci. ro: 162-218. f. I-13. 1917. Includes Dimeriopsis, Hyalosphaera, Borinquenia, Dexteria, Monogrammia, and Microclava, gen. nov., and 53 new species in various genera Stevens, F. L., & True, E. Y. Black spot of onion sets. Illinois Agr. Exp. Sta. Bull. 220: 507-532. f. 1-19. My 1919. Describes Cleistothecopsts gen. nov. Tehon, L. R. Studies of some Porto Rican fungi. Bot. Gaz. 67: eat pl. 18. 21 Je 1919 Ne ies are described in Méliole (4), Phyllachora (2), Stigmatea (1), Phaeo- Sphaerella 1. Coniothyrium (1), Pestalozzia (1), Acrothecium (1), and Trichostroma (1). Tracy, S. M. Rhodes grass. U. S. Dept. Agr. Farm. Bull. 1048: I-14. f. 1-3. ‘Je 1919. Trelease, W. The chestnut in I!inois. Trans. Itlinois Acad. Sci. 10: 143-145. I987. Tuttle, G. M. Induced changes in reserve materials in evergreen herbaceous leaves. Ann. Bot. 33: 201-210. f. 1-7. Ap 1919. Waterman, W. G. Plant ecology and its relation to agriculture- Trans. Illinois Acad. Sci. 10: 123-129. 1917. Williams, R. S. The genus Desmatodon in North America. Bull. Torrey Club 46: 207-220. pl. 11. 8 Je 1919. Winslow, E. J. Early days of the American Fern Society. Am. Fern Jour. 9: 33-38. Jl 1919. Woodward, R. W. Some Connecticut plants. Rhodora 21: 114-116- Je 1919. Vol. 46 No. 11 BULLETIN TORREY BOTANICAL CLUB NOVEMBER, I919 The ferns and flowering plants of Nantucket-—XX EUGENE P. BICKNELL CONCLUSION The plants of Nantucket enumerated in the catalogue, brought to a close with these pages, number 1,136, including thirty-three natural hybrids. Other probable hybrids and some sixty varieties, mainly such as have received names, have been referred to in the text. Some of these plants that are listed under binomial names are without general botanical consent to be taken as com- pleted species. These, however, are all plants of marked attributes and, however opinions may differ as to the precise status of this one or that, in each case a binomial name has been allowed its sufficient claim unmistakably to denote the plant referred to. This dissembles nothing and has seemed to avoid whatever of factitious precision may inhere in a needlessly composite name. On the other hand forms that pass current under varietal names that seem unduly to honor certain of those variations which, in undefinable number, are common to plants in general, have had only their passing word. Omitting the hybrids the number of plants formally listed is I,103.*. Of these 746 are native to the island and 362 are in- troduced. : * This number includes a few mere varieties admitted in the earlier parts of this series, and allows for two species that should be omitted and four here to be _ added. The eliminations are Panicum linearifolium (see Supplementory Notes, antea) and Bartonia iodandra. The Nantucket plant discussed under this latter name should be referred to B. paniculata—a contracted form in which the anthers [The BULLETIN for October (46: 389-422) was issued November 5, 1919.] 423 ‘ 424 BICKNELL: FERNS AND Included among the introduced plants are some thirty species common in cultivation which, although not to be taken as es- tablished, have been found persisting in a wild state. The tend- ency shown by these plants to stray away from their home gardens may well thus be put on record. Should any of them in time come to take a wider place in the flora the beginning of their history as wild plants would become a matter of interest. Nearly one half of the introduced species are well naturalized and more than two-thirds of these are widespread, including in their number many of the island’s abundant plants. About sev- enty-five species may be accounted common, forty as frequent, seventy as of occasional occurrence. Almost everywhere where introduced plants abound a much larger proportion of their species than those of the native plants are confined to a single station or to a very few places, and this is notably the case on Nantucket, notwithstanding that an unusual number of native plants are there thus localized. One hundred and seventy of Nantucket’s introduced plants have been found at not more than two or three localities, eighty of them at a single station only. The town of Nantucket itself with its old gardens, its resting wharves, its streetsides and habitable places for plants that have had a quiet past, has come into a flora very interestingly its own, are sometimes deep purple, a character hitherto regarded as diagnostic of B. iodan- dra. ome examples, as already intimated, seem to pass into more typical forms of B. paniculata, and appear to be similar to the Massachusetts plant reported and figured as B. iodandra by Williams (Rhodora 2: 55-57. pl. 15, f. 5. 1900), now re- garded, I understand, as B. paniculata. Professor Robinson who, with Professor Fernald, has recently examined my Nantucket specimens advises me that they are not B. ioandra, and specimens of the nae from Newfoundland which were kindly sent to me, show a plant very unlike anything I have net wt ith on Nantucket and beyond question, as I now see it, a strikingly distinct species. I am indebted to Miss Albertson for Sen of the eile additions to the list, one native and three introduced species, as follow STEIRONEMA LANCEOLATUM (Walt.) A. eae Monomoy, August 22, 1918, in man HUS MAXIMILIANI Schrad. Field i in Madequet, August 15,1919. Col- lected in full flower by Miss Ober. CENTAUREA JACEAL. Field near Franklin Fountain, August, 1915, and July. 1918. CENTAUREA MACULOSA Lam. Near Big Mioxes, August 23, 1919. Collected in full flower by Miss Milne. FLOWERING PLANTS OF NANTUCKET 425, many of whose plants would be looked for in vain elsewhere on the island. So it is that more than one hundred introduced plants have been reported only from the general district of the town. Additions to this domesticated flora now follow on almost from year to year. Freshly seeded lawns and modern gardens add their new comers, and suburban fields where rubbish has been cast encourage the growth of adventitious plants that take their foothold either transiently or tenaciously as time shall prove. Among introduced species not everywhere found that are well established in the town flora but are rarely if at all seen elsewhere on the island are the following: Pucctnellia distans Viola laetecaerulea Bromus sterilis Aethusa Cynapium Carex hirta Marrubium vulgare Iris pseudacorus Physalodes Physalodes Lychnis dioica Solanum peregrinum Chelidonium majus Tragopogon pratense Geranium pusillum Sonchus arvensis Some rather unusual alien plants have made their appear- ance within recent years in different parts of the island about abandoned spots where poultry has been kept, their source of introduction doubtless being the prepared chicken feed now in general use. The mixed growth of weeds that throng such places: have yielded the following species, very few of which have been. found elsewhere: Vaccaria Vaccaria Conringia orientalis Lepidium neglectum Lappula Lappula Thlaspi arvense Amsinckia arenaria Nesha paniculata Lithospermum arvense Erysimum cheiranthoides Centaurea melitensis In addition to those abundant naturalized plants that every- where enrich our flora Nantucket has adopted into its vegetation some less usual species that now, common and wide-spread, assert themselves as characteristic island plants. The following may especially be named: Alopecurus pratensis Festuca myuros a geniculatus “ capillata 426 BICKNELL: FERNS AND Aira caryophyllea T rifolium dubium Carex muricata Epilobium hirsutum Salix Smithiana Centaurium spicatum Cerastium semidecandrum A pargia autumnalis Ranunculus repens Artemisia Stelleriana Cytisus scoparius Arctium tomentosum The great majority of the introduced species are herbaceous plants of dry and open ground, many of them succeeding in poor and sandy soils. Less than twenty species are plants of moist or wet places and only one is conditioned by saline influences. Eighteen herbaceous species are twining or climbing plants, three species are woody vines and twelve are shrubs. The introduced trees number thirty-two, although only one has become a strong struc- tural element in the flora, this being our native pitch pine which, history tells us, was first planted on Nantucket in the year 1847. Few other introduced trees have made much response to the con- ditions that Nantucket has offered, although the cockspur thorn is making itself at home there, and the apple, the pear and the hybrid willow (Salix Smithiana) are sparingly more or less wide- spread. The Scotch pine and the European larch have long formed an extensive and increasing growth at the locality where they were. originally set out, and at a few places the locust and the silver poplar are well established, but most of the other trees are not much to be considered, and some number only a few examples that have appeared spontaneously and grown up in out-of-the-way places. At different times in the past collections of trees were brought to Nantucket and set out in certain places which now, long neg- lected and apart from cared for land, appear like wild tracts covered with their native growth. Both European and American species were used in these plantings and many of them survive, although few have made much growth. Most of these trees have been noted in the text. Not many of the introduced plants are from slinwiseed than Europe and Eurasia, these numbering some two hundred and seventy-five species. Twelve species belong to Asia and nine are tropical American. Japan, China, India, New Zealand and Mex- ico have each contributed a single species which is, however, either i FLOWERING PLANTS OF NANTUCKET 427 a scarce weed ora casual estray from cultivation. Included among the introduced plants are also some sixty North American species, all of which are quite certainly not of the island’s indigenous flora. The families represented by all these plants number sixty- three, eleven being foreign to the indigenous Nantucket flora. As many as nineteen have only a single member and the same number not more than two or three. The most numerous family is the Gramineae with fifty or more members, a number subject to con- siderable increase by taking account of all named varieties. Among the grasses, as in other groups, the absence will be re- marked of some species that might well be looked for. As an instance, so common a port and streetside immigrant as Eleusine indica has not yet made its way to Nantucket where, it might be thought, the shipping activities of earlier days might long ago have brought it in. The family most noteworthy in respect of greater representa- tion by alien than by native members is the Cruciferae with thirty- four introduced species and only four that are native. A less marked preponderance of immigrant species is found in the Caryophyllaceae and Fabaceae, these families having respectively ninteeen and twenty-five alien members against eight and eighteen, in corresponding order, that are indigenous. The number of introduced species belonging to other well-represented families are as follows: Compositae, 28; Labiatae, 18; Cichoriaceae, IT; Scrophulariaceae, 10; Salicaceae, 10; Solanaceae, 9; Polygonaceae, 9; Chenopodiaceae, 8, and Rosaceae, Ammiaceae and Borragi- naceae, 7 each. The largest of the native plant families, the Cyperaceae, numbering eighty-seven species, has only two species that are introduced, both Carices, and the third largest native family, the Compositae, to which so many of our immigrant weeds belong, numbers but thirty-two introduced species against seventy- two that are indigenous. The Rosaceae and Polygonaceae also preponderate as native families, possessing respectively twenty- nine and twenty indigenous members and only seven and nine that have come by immigration. Of important native families the Juncaceae and Orchidaceae are, as would be expected, totally un- represented in the introduced flora, and there is only a single in- 428 BICKNELL: FERNS AND troduced fern, the common polypody, this having somehow found lodgment on the island at a single station. Among the introduced plants yellow flowers and white flowers greatly predominate, their ratios to the total number being re- spectively over 31 per cent and 28 per cent. White flowers and yellow flowers likewise predominate in the indigenous flora, but here white-flowered plants are relatively, as absolutely, much more numerous, their corresponding ratio being 39 per cent, that of the yellow-flowered 22 per cent. Next in order among the introduced plants come pink-flowered species, including purple-pinks, 19 per cent; purple and blue-purple, 12 per cent; blue, over 6 per cent. Orange and red are each found in only four species, and green- petaled flowers are no less rare. Eliminating those introduced species that seem to have little chance of permanency the resulting percentages show not much change beyond about 4 per cent de- crease in the purple- and blue-flowered species and a corresponding increase in the white-flowered. As among the introduced plants, so in the native flora flowers of some shade of pink, including those scarcely assignable shades lying between pink and purple, are next most numerous after whites and yellows, making up’nearly 17 per cent of the whole Purple and blue-purple follow with over 8 per cent; blue with nearly 4 per cent; red and orange, the one something over, the other a little under 1 per cent. A wide percentage disparity is found between the native and the introduced plants that bear flowers fairly to be described as green, these numbering less than I per cent of the introduced species and over 8 per cent of those of the native flora. From the nature of the case all these percentages are somewhat approximate, not being susceptible of very exact rendering. Petalous flowers only have been taken into the reckoning. * * * * The indigenous flora of Nantucket has its most interesting side to the botanist in its many species of plants that more es- pecially abound southward on the coastal plain. This is its domi- nating note. But mixing with these plants of southward range are others which trace through the flora a strain of northward re- lationship that is all the more sharply defined by the emphasis of contrast. FLOWERING PLANTS OF NANTUCKET 429 In its more local aspect two interesting features of the flora that will not long escape the exploring botanist are the large proportion of its plants that are confined to the eastern side of the island and the number of species siete that are strictly localized. The eastern side of the island is the more diversified in its topography and natural features, abounding in knolls and hollows, in damp open grounds and straggling swamps beset with crowded shrubbery, and hillsides and banks with their herbage and woody growth descending to numerous small ponds and mossy or peaty bogs. It has also more varied and mature tree growths than are found on the western side and reveals better evidences of former forestation. Accompanying these conditions are more varied and richer soils with their responding plant life, and many of the more southern and more northern plants that belong to the flora have their place only here. Here, too, surviving in the thickets and tree groupings, are little colonies of woodland plants, vestiges, we may suppose, of an earlier flora that had its day in that unrecorded period before the woodlands were destroyed. Extensive dry plains clothed with low herbaceous vegetation spread over much of the southern side of the island, invaded to- wards the east by barrens of scrub oak and midway in the island by open formations of young pitch pines advancing from denser growths that earlier made their conquest. Westward towards Hummock Pond are veritable tracts of pine barrens which, how- ever, as a modern feature of the island’s vegetation, have merely adopted their plants from the general plains flora, not contributing any distinctive species of their own.* Certain reaches of these plains of darker soil call to mind the Hempstead Plains of Long Island. Like the Hempstead Plains, too, these Nantucket plains have their widely distributed assemblage of particular plants, very few of which do not belong as well to neighboring territory. * It should be noted that among the Nantucket pines are found a few woodland plants that either do not occur at all elsewhere on the island or are nowhere else at home. It would seem to follow that the advent of these plants, or some of them, must have been subsequent to the introduction of the pines. Four of them that have been reported from only one or two localities are Polypodium vulgare, cee rhiza maculata, Linnaea americana, and Hypopitys americana; three others, t are sparingly scattered, are Hypopitys lanuginosa, Pyrola chlorantha and hse mac 430 BICKNELL: FERNS AND And with few exceptions their characteristic plants are also plants of the Hempstead Plains.* Among the more interesting are the following: Schizachyrium scoparium ence week propinquum Panicum depauperatum majus meridionale tHudsonia ericoides Aristida purpurascens Lechea maritima Agrostis antecedens Viola pedata Carex pennsylvanica tEpigaea repens “tonsa tArctostaphylos Uva-urst Juncus Greenet Asclepias amplexicaulis Salix tristis Agalints acuta Aletris farinosa +Houstonia coerulea Sisyrinchium. arenicola Nabalus serpentarius Baptisia tinctoria Chrysopsis falcata Cracca virginiana Solidago memoralis Lespedeza Bicknellii Euthamia tenuitfolia Linum intercursum Sericocarpus lintfolius Polygala polygama Tonactis linartifolius Sarothra gentianoides | Aster concolor Helianthemum dumosum “* dumosus Only four of these plants, those marked with a dagger, appear to be wanting on the Hempstead Plains, although the arbutus and the bearberry occur at outlying points. The Houstonia, an abundant Nantucket plant, is apparently unknown on Long Island and the Hudsonia is a rare plant there. * * * * * Over one half of Nantucket’s native plants may fairly be accounted as prevailingly more southern in their general distri- bution. More than one hundred of these reach their northern or eastern limits in Massachusetts, mainly in the Cape Cod region and in the southeastern quarter of the state, and a considerable number of others that pass beyond Massachusetts are reported from no further than southwestern Maine or are of only casual occurrence at more northern or eastern points. Altogether about * For a very full discussion of the Hempstead Plains and their vegatation, see Harper, The Vegetation of the s. Hempetend Plains. - Mem. Torrey Club 17: 262- 286. pl. 7. June, 1918. FLOWERING PLANTS OF NANTUCKET 431 one hundred and fifty of these plants extend their northeastern limits, in many cases only locally, into Vermont, New Hampshire or Maine, and over seventy others, some of them unknown in Maine, occur as far to the east as the Canadian Maritime Prov- inces. A smaller group, very noteworthy in its isolation, are found at the extreme eastern extension of the continent in New- foundland.* The following plants of Nantucket do not appear to have been reported from any more northern or eastern point, and a considerable number of others have only within recent years been added to the known flora of the eastern Massachusetts main- land. The names used follow those of the catalogue, although some changes have since been adopted. Lycopodium alopecuroides Boehmeria scabra Naias guadalupensis Polygonum pennsylvanicum var. Schizachyrium villosissimum nesophilum 2 littorale Persicaria setacea Panicum virgatum cubense Amaranthus pumilus “ Bicknellii Cardamine arenicola “ albemarlense Rubus flagellaris “ auburnae Aronia arbutifolia Chaetochloa versicolor ‘ Ascyrum hypericoides Agrostis elata Lechea Leggettit Elymus halophilus Opuntia Eleocharis tricostata Vaccinium atlanticum Scirpus Eriophorum 0 vicinum Rynchospora Torreyanat Apocynum pubescens Carex debilis Setiscapella subulata “ incomperta Viburnum venosum Arisaema pusillum Solidago aestivalis Juncus dichotomus Euthamia minor “ aristulatus Doellingeria humilus Quercus pagodaefolia The similarity of the Nantucket flora to that of southern New especially, Fernald, A botanical expedition to Soka aR and i a Labrador (Rhodora 13: 109-162 pl. 86-91. 1911), a paper narrative interest apart from its botanical importance + One other New England station for this species is known at East Wash- ington, New Hampshire. 432 BICKNELL: FERNS AND Jersey, which has often been remarked, is strikingly brought out by a mere accounting of the number of coastwise plants common to the two regions. More than one hundred and ninety Nan- tucket plants belonging to the more southern element of the flora occur in the New Jersey Pine Barrens, and over three hundred are plants of the Coastal Plain elsewhere in that state; while all the maritime plants of Nantucket of general southward range, about forty species, are also of the New Jersey flora.* Over one half of Nantucket’s native plants may be classed as common, about one hundred and forty as frequent and about fifty as of occasional occurrence. As many as one hundred and eighty-two have been found at not more than one to three places and, as far as known, nearly ninety of these occur at only a single station, these categories of rarest plants constituting almost one quarter of the native flora. Nearly three-quarters of these have been found only on the eastern side of the island. The total number of species that are thus restricted number one hundred and seventy-five, and forty-five others are mainly of the same local distribution, these together making up nearly 30 per cent of the flora. Not more than thirty-seven species, or 5 per cent of the flora, are found exclusively on the western side of the island. Nearly 45 per cent of Nantucket’s native plants, about three hundred and thirty species, are plants of low grounds, swamps and pond shores, while little more than one quarter are species that thrive best in dry open ground. Less than twenty species are plants primarily of low or wet woods. Other woodland plants number about ninety species, here for the most part keeping to copses and thickets but also, many of them, appearing on the plains and commons, some having very successfully adapted themselves to these unaccustomed dwelling places. Fogs and moisture from the sea from whatever direction the wind may blow must have had an influence in this, and a curious contributing cause may be seen in the habit of growth of one of the plants them- selves that belong to these exposed tracts, the abounding bear- berry, whose shining evergreen carpet lies broadspread on hill * See Stone, The Plants of southern New Jersey with especial referencé to the flora of the Pine Barrens and the geographic distribution of the species. Rep. New Jersey State Mus., 1910. The statistics above given have been made pos- sible only through consultation of this work FLOWERING PLANTS OF NANTUCKET 433 and plain. This gives winter protection to the soil and in sum- -Mer a continuous shade and coolness under its low covering for such woodland plants as here and there may find space enough to make their growth. Nantucket has been called a treeless island and, apart from the town and the scattered farms, the casual tourist might be- lieve this to be true except for the naturalized pines which are now widely in the landscape. Nevertheless, twenty-five to thirty kinds of trees are native to the island, the larger reckoning allow- ing for those that, shrub-like on Nantucket, are trees in their full growth. In general, however, the Nantucket trees are not promi- nent in the vegetation, not many rising above a very moderate height, although there are copses and groupings in low grounds where they attain a good woodland size, and in the seclusion of dense thickets is to be found here and there a beech or an oak little noticeable for height but of a girth of trunk that reports a venerable age. Shrubs abound, and in swamps and low grounds are massed into thickets of the most dense and impenetrable character. The number of species that are botanically shrubs is seventy-two, many of them, however, bearing little distinction of aspect from the herbaceous vegetation amid which they grow. There are eight woody climbing vines and the same number of twining herbaceous species. Of other Nantucket plants having their particular habitats fifty-five belong to the salt marshes and twenty are of the coastal sands. Of thirty-three aquatics three only are exclusively maritime. The number of native families represented in the Nantucket flora is one hundred and thirteen. Thirty-eight have only one species, twenty-seven two or three species, twenty-six four to nine species and seventeen ten to twenty species. Only four families contain over twenty species, the Rosaceae with twenty- nine and the three predominant families, the Cyperaceae, Grami- neae and Compositae with eighty-seven, seventy-eight and seventy-two members. Including in the Rosaceae twenty-four hybrid blackberries that have been described (and other combina- tions among these occur) its actual membership would approach that of the three highly preponderant families. The ferns number twenty-six, belonging to three families, the Ophioglossaceae, five 434 BICKNELL: FERNS AND species, the Osmundaceae, three species, and the Polypodiaceae, eighteen members, including three commonly regarded as strongly marked varieties and one hybrid. When discussing the introduced plants it was remarked that they included some sixty North American species that were not indigenous on Nantucket. Some forty of these are common plants of the New England mainland and seventeen are of the west and north. Not more than two or three immigrant native species have come to Nantucket by way of the south, and these seem to be little at home and have not spread from their original stations. Here is perhaps a hint that plants of southern conformities may léss readily adapt themselves on Nantucket than do species from the north and west, as if the present climatic conditions of the island might not be encouraging to the advance of more southern types. Should this indeed be true it would seem to reveal that those southern affinities now in such clear aspect in the flora of the island are to be referred back to influences not the same as those of today. And there would be accordance in this with those now well understood evidences of an extensive flora of southern derivation belonging to the New England seaboard of Tertiary time, a flora lost to our later day with these broad coastal tracts that now lie beneath the sea.* Yet not wholly lost. We find it still, much of it, we may believe, in the less disturbed flora of our more southern coastal plain, and we find its remnants persisting as the merest fringe along the withdrawn more northern coast- lines of the present day. And isolated on Nantucket it has been preserved to us in that assemblage of southward ranging plants, now a primary element in the general composition of the flora. Thus, understanding the far back origin of this relationship, we may the more readily believe that Nantucket’s possession of southern plants may be little attributable to influences operative at the present day. There is other evidence than the general absence of immigrant southern plants on the island that would seem to denote conditions less favorable to more southern than to more northern plants. * See Fernald, loc. cit. Also, The geographic affinities of the vascular floras of New England, the Maritime Provinces and Newfoundland. Am. Jour. Bot. 5: 219-236. pl. 12-14. 1918. FLOWERING PLANTS OF NANTUCKET 435 It lies open to the eye on Nantucket that the island is not at all inhospitable to plants of far northern and eastern range, and not a few such species have placed here their outlying southward colonies. And this corresponds in its import to what is true of such of our naturalized plants as have adopted anything like a definite range. Many more Nantucket plants are species whose main colonization in this country is to the northward rather than to the southward. And evidence of like bearing may be seen in this, that many of the island’s more southern plants are strictly localized, and, going back, as we may do in certain cases, even to the time of its earliest botanical explorers, have spread not at all from the localities where they were originally found. In the case of coastal plain species an explanation of this is hardly to be sought in soil conditions, and some repressive influence may be suspected in the notably delayed spring on this island and the average low temperature during the growing season that is a feature of its climate. * * ok oe * The northern element in the Nantucket native flora comes to view in its broader aspect in something over one hundred and fifty species that are at least prevailingly more northern in their general distribution. Many of these plants are to be accounted more northern partly in a distributional sense which allows for that equivalency in altitude which has permitted a far southward extension along the Appalachian highlands. Thus while probably less than fifteen Nantucket plants are nowhere found at a more southern point some three times that number are on this island at or near the southern limit of their coastwise range. Other Nan- tucket northward plants have taken some further way toward the south, a few to be stayed on Long Island, a larger number passing on, not without wide intervals, to find their southern limits in the Pine Barrens and Coastal Plain region of New Jersey. It might be supposed of the maritime species, their way open along the coast as far as had been their bent to follow it, that a freer progress had marked their course. But this has been not at all the case. Some of these plants also have made little or no advance beyond Nantucket, others in like manner with the more’ inland species have been held at various more southern points. 436 BICKNELL: FERNS AND And it is pertinent here to note that no northward maritime plant of New Jersey is not also a plant of Nantucket, for with the in- land species the case is different. Turning again to Dr. Stone’s admirable and enlightening analysis of the flora of southern New Jersey, wherein is much.that now enables us better to understand the flora of Nantucket, we find a list of fifty-nine species that range south to New Jersey from the Canadian Maritime Provinces. Forty-one of these plants belong to the Nantucket flora. Of the eighteen that are not found there, ten are also wanting on Long Island and most of the others are all but absent there. And only three of these eighteen plants have been found on Martha’s Vineyard. These species are the following: Lycopodium inundatum Carex limosa Schizaea pusilla Salix lucida Scheuchzeria palustris Dalibarda repens Phalaris arundinacea Geum strictum Panicularia obtusa Hypericum ellipticum Scirpus subterminalis ~ Ascyron Carex trichocarpa Polanisia graveolens * extlis Aster nemoralis “ livida Xanthium commune It is to be noted of these plants whose course has taken them to southern New Jersey, while withholding them from the re- lated floras of Nantucket, Martha’s Vineyard and Long Island, that with few exceptions they are of pronounced general northern range. At their eastern limits they approach the seaboard at a relatively high latitude, most of them ranging westward and bearing south as elevation of land or favoring conditions may allow. Their approach to the New Jersey littoral would thus appear to have been by an inland route by way of the Alleghanian highlands that cross the Hudson and the northwestern part of that state. From their places of abundance to the northeast the drift of their range westward and southward has passed to the north of southeastern Massachusetts and borne them on their highway along the hills whence, at a lower latitude, they have found access to the region of the Coastal Plain. All this the evi- dence would seem to allow us to believe. FLOWERING PLANTS OF NANTUCKET 437 Missing thus certain plants not debarred by climatic condi- tions Nantucket has received into its flora other northern species even less to be expected there. There is some reason to believe that these plants may not have come to Nantucket directly from the north but from a more eastern part of their range. Just as Nantucket’s coastal plain flora partly reappears in the Maritime Provinces and even in Newfoundland, so, conversely, do many northward species characteristically associated in these regions find themselves together on Nantucket. Whether this association of their plants, many of which are of wide distribution, points back to some common heritage in the floras of these regions our present knowledge does not assure us. But something of affinity between these far separated floras seems to sketch itself in outline and not without features more clearly drawn. Such plants of Nantucket as Fragaria terrae-novae, Ribes oxyacanthoides var. calcicola and Antennaria petaloidea var. subcorymbosa would scarcely be looked for from elsewhere than far to the east. If an ancient land connection has conducted southern forms to these far eastern fields, some counter extension of northern forms, at least in the later age of that one time highway, may well have taken place. It is in point that almost directly to the north of Nantucket in eastern Massachusetts, at no greater distance than Essex County, nowhere of much elevation, many northern plants prevail that have obtained no foothold on Nantucket, even though, as we have seen, the island offers locally soils and conditions that are acceptable to northern woodland plants. And nearer at hand, in the Cape Cod region, where many of the conditions repeat those of Nantucket and many features of the flora are the same, there are, well in place, northern plants that have failed to cross the narrow strip of water to Nantucket. And, emphasizing in an unexpected way this slight rift of disconnection between the similar floras of these coastwise tracts, there are many southern plants of the Cape Cod region, not a few of them well established there, although at the extreme northern limit of their range, that on Nan- tucket are unknown.* * See, wom Gam S. Collins, Notes on the Flora of lower Cape Cod. Rhodora II: 125-133. 1909. Also, Flora of lower Cape Cod; supplementary note. Rhodora 12:8-10. 1910. Also, Flora of lower Cape Cod; third note. Rhodora 13: 19-22. 911. Also Sinnott, The pond flora of Cape Cod. Rhodora 14: 25-34. 1912. 438 BICKNELL: FERNS AND If we may picture this region in the formative period of its present flora as enriched with a vegetation crowded along the shore lines of bays and inlets from sound and sea, just as today in such situations plants have assembling places that delight the botanist, then today’s surprises in this inward coastal flora have their explanation. For Nantucket, then perhaps little more than a headland flanked with bleak sand wastes along an exposed outer coast, must have proved a difficult and impermanent refuge to many southern species that would find encouragement and per- petuation along the quiet inward shore lines further to the north. Fragmentary and unclear as these seeming evidences may be they unite in suggesting a closer affinity in Nantucket’s flora with the flora of a more eastern region than with that of the northward mainland more nearly at hand. And if there be revealed in this the broken ties of an ancient relationship it traces itself con- formably with the general trend of the coast and with that diagonal northeastward-southwestward sweep of distribution that has given their geographic lines to so many of our plants in their eastern range. The following northward species of Nantucket are unknown in the coastal region of New Jersey, not many of them passing on even to Long Island. Phegopteris Phegopteris Myrica Gale Isoétes Tuckermani Corylus rostrata Panicularia grandis Persicaria Hartwrightii Scirpus occidentalis Tissa canadensis ‘““ rubrotinctus Coptis trifolia “ pedicellatus Ribes oxyacanthoides var. calct- Eriophorum viride-carinatum cola es utriculata Fragaria terrae-novae Goodenovii Argentina litoralis ‘* monile Rubus strigosus ‘“ subloliacea “ triflorus “ prairea Lathyrus pilosus “ diandra Ilex bronxensis Lemna trisulca oe palustre Juncus balticus lineare *“* bufonius var. halophilus xe strictum © FLOWERING PLANTS OF NANTUCKET 439 Epilobium adenocaulon Agalinis paupercula Ligusticum scoticum: Galium palustre Coelopleurum actaetfolium Linnaea americana Chamaepericlymenum canadense Antennaria neodioica var. attenu- Chiogenes hispidula ata Oxycoccus Oxycoccos Antennaria petaloidea var. sub- Pneumaria maritima : corymbosa Mentha glabrata * es * * * A word remains to be said of the degree of completeness with which the flora of Nantucket is now probably known. The botany of many another region of even greater extent might be very thoroughly chronicled from far less investigation than has been bestowed on Nantucket. Here, as we have seen, are found conditions that ask for close work by the explorer and notwith- standing what has already been accomplished the island today remains a broad and an inviting botanical field. I do not in the least doubt that a systematic exploration continued through an entire season would bring to light many unexpected things and add materially to the list of Nantucket plants. Many a bog, or shrubby patch, or little space of open hillside or plain is alone the home of some noteworthy Nantucket species, and the difficult thickets, especially, cannot yet have yielded all their secrets. When we recall how easily, out of their particular season, many plants may miss the eye, and that there is already a long list of Nantucket species, each one having its single spot somewhere in the fifty square miles that make up the island’s area, the possibili- ties of new discoveries may well stir the later explorer with some- thing of the enthusiasm which animated those who were pioneers. I should suppose that it were well within the possibilities that IO per cent or more of the native flora remained to be brought to light, and that an addition of not less than 5 per cent to the number of plants already known might be confidently looked for. y own visits to the island have been never for more than brief periods and have wholly missed the important seasons of mid- summer in the last weeks of July and of August. Nor have I col- lected on Nantucket in mid-August since 1904, nor in the autumn since 1907, nor explored there at all since 1912. It may be further 440 BICKNELL: FERNS AND FLOWERING PLANTS OF NANTUCKET useful to the future explorer to speak of certain parts of the island where, especially, careful work remains to be done. In the southwestern quarter that not extensive tract known as ‘‘The Woods” which, although treeless, must at some unrecorded period have justified its name, has probably not been visited in every season, and Trot’s swamp and the maze of thickets, wet and dry, in Squam, as well as parts of Polpis, are places which not every _ collector may have cared to penetrate. At Coskaty there is a thickly wooded tract where it is probable few botanists have ever been. Only once and too hurriedly have I gone amid the thick undergrowth of this piece of timber and its complete ex- ploration has remained an object unattained. Nor has my hope yet been realized of sometime traversing the long sand strip be- - tween Coskaty and Great Point. Reading again the pages of Mrs. Owen’s catalogue through its perspective of more than thirty years, we are reminded anew of the - singular rarity on Nantucket of many of its most interesting plants, and of their close seclusion in those chosen spots that have given them protection. Some of these plants, many of them, indeed, that were discovered by Mrs. Owen, or announced through her by the active group of collectors which she inspired, have rarely been encountered since. And as many as fifteen or twenty species then reported, which we have no reason to believe are not growing somewhere on Nantucket today, remain to be redis- covered by those whose pleasure it shall be to continue the study of the island’s botany so long ago begun. Sex in the Conjugatae and the relative frequency of the different types of conjugation H. W. Tuurston, Jr. It has been much disputed in the past whether there is a true sexual differentiation between the cells that fuse in the process of zygospore formation as found in the filamentous Conjugatae. Bessey (2) and others have argued that the process probably is sexual in cases of scalariform conjugation at least. Bennett (1) says: ‘‘I do not myself entertain any doubt that even in lateral conjugation there is an incipient differentiation of sex, although this differentiation extends only as far as the individual cells.’ The Wests (6) have gone even farther, and say, ‘‘against the sexuality of the Zygnemeae only two plausible objections can be raised; these are the phenomena of /ateral and cross-conjugation,”’ but they conclude that there is no reason even on these grounds to regard the Zygnemeae as other than sexual. Regarding the Mesocarpeae however, they say further that ‘indications of sexu- ality . . . are much less marked than in the Zygnemeae,”’ and, ‘‘as these scarcely appreciable indications of sexuality are often absent, we may regard the Mesocarpeae as having lost almost all traces of differentiation into male and female gametes.”’ Still later, G. S. West (7, p. 135) discusses the question as follows: “‘ The term ‘sex- ual’ is often used to embrace all forms of gamogenesis, and is in this sense firmly established in botanical literature. It would, how- ever, be more scientifically accurate to distinguish between gamo- genesis (or the mere fusion of gametes) and sexual reproduction in the narrower sense (which should be restricted to those cases where there is a fusion of clearly differentiated 7 and 9 gametes). From this point of view, therefore, gamogenesis although including sexual reproduction is not identical with it. It must be remarked, however, that the gradation is so fine, especially in the Chloro- phyceae, that the distinction is scarcely worth making. Some- times, as in many of the Zygnemaceae, the gametes are morpho- logically’ indistinguishable but physiologically differentiated, 441. 442 THURSTON: SEX IN THE CONJUGATAE and in these cases there is often a morphological differentiation of the gametangia.”” The question certainly arises whether we ever find cases of reproduction by cell fusion as contrasted with reproduction by cell division in which the process is not. to be considered as involving the essential features at least of sexual reproduction. We have obviously come to think of sexuality as involving some fundamental difference between the cells that fuse or conjugate, and consequently when we can find no. visible difference between the gametes and can not in any way tell the male from the female, we find ourselves inclined to doubt the so-called sexual nature of such a conjugation. This is the case for all species of filamentous Conjugatae that form their spores midway between the two conjugating cells (TABLEs II and IV) and the common doubt as to the sexual nature of this conjugation is expressed by the above quotation from West. Naturally among low plant forms of this kind we should expect to find the most primitive kind of sexuality in its most fundamental expression. Many of our fundamental ideas regarding the nature of sex pro- cesses we owe to the study of these very algal forms. Schmitz in 1879 saw two nuclei in a zygospore of Spirogyra ‘approach each other and fuse to a single nucleus.’ This was the first description of the nuclear behavior in the sexual process in the plant kingdom and was made before any such process was known to occur in any of the higher plants. It is certainly possible that by the intensive study of these primitive forms we may further clarify our ideas as to the nature and origin of sex. Wittrock (8) has defined a zygospore as ‘‘a spore formed by an act of fecundation in which two or more cells of the same kind, not sexually different, have participated.” Can we accept such a definition, and at the same time accept conjugation in these forms as being essentially a fecundating reproduction? If we can, the formation of gametes can involve only the difference - between somatic cells as such and gametes, as such, rather than any essential difference between the two gametes themselves. Perhaps, after all, cells fuse rather because they are alike than because of an inherent difference between them. Certainly both of a pair of fusing cells in Spirogyra differ in some way from the | somatic cells from which they arose. Such differences between THURSTON: SEX IN THE CONJUGATAE 443 gametes as those of size and motility are perhaps to be considered as mere adaptations introduced in evolution to facilitate the bringing together of the twoelements. Tréndle (5), for example, claims to have proved that the vegetative cells of the Spirogyra filament are haploid; then the difference, in the case of Spirogyra at least, between somatic cells and gametes can not be one of chromosome number, but some difference there must be to inhibit further vegetative division in any given cell of a filament and cause instead a union of that cell with another which has under- gone a similar change. The gametic nature of any cell in a fila- ment would then be fixed when it ceases to have the power to divide but instead has a new found power to unite with some one of its fellows. At that moment the cell ceases to be a somatic cell and becomes a gamete. Cunningham (3) has reviewed the literature of the sexuality of Spirogyra rather thoroughly, but he throws out the cytological - evidence of Tréndle when he compares the filament of Spirogyra with the sporophyte of higher plants. He concludes that ‘reduction may occur in the zygote, in which case a filament wholly of one sex arises, or reduction may occur just previous to reproduction, in Which: Casey cs a es filaments of a bisexual nature are produced, which would conjugate either laterally or by cross-conjugation.”” Two of the species described by Tréndle as having the reduction take place in the zygote, would therefore according to Cunningham produce filaments ‘“‘wholly of one sex,’’ which however are known to conjugate in both the lateral and scalariform manners (see TABLES I and III). These are S. Jongata and S. neglecta. The real evidence of fecundation in these forms is not to be found in a visible or measurable difference between the filaments or even between the cells that fuse, nor is it to be found in the method of conjugation, whether lateral, scalariform, or cross conjugation, ' but it lies in the newly achieved possibility of fusion itself with resultant doubling of the nuclear material and the subsequent reduction division, which as far as the evidence extends at present takes place on the germination of the zygospore. If we believe that similarity between gametes favors rather than hinders fusion, then there is nothing at all startling in considering lateral con- _ jugation as a fertilization even if the cells taking part are sister 444 THURSTON: SEX IN THE CONJUGATAE cells, and nothing startling in considering the conjugation in all the species of the Mesocarpeae as fecundation. Even a case such as that described by Petit (4) for S. mirabilis, where he says that the contents of a single cell separate into two parts which reunite forming a spore, might be considered as a fertilization. There may also be an explanation here for many of the so-called aplano- spores and parthenospores, which when the facts are fully known may be found to go through some process similar to that described by Petit for Spirogyra or by Woodruff for Paramoecium. My purpose in gathering together the subtended tables has been to show the relative frequency of the different types of conjugation as shown in the literature. It is obvious that lateral conjugation is very common. This fact, together with occasional observations of cross conjugation, and the fact that both lateral and scalariform conjugation are often found not only in the same species, but in one and the same filament, go to prove that the individual cell rather than the filament should be regarded as the unit when the fusions in these forms are being considered. It is of course quite impossible in the present state of our knowledge to estimate finally the validity of many of these species. No claim is made for the completeness of the tables nor that all synonyms have been eliminated. In my opinion, however, they show approximately the proportions in which the different types of conjugation have so far been described in the literature and may be useful to students of the general problems of sex. The principal sources of the tables are such standard works as Wolle, Hassall, Cooke, Petit, De Toni, West and Collins (as indicated in parentheses), although numerous shorter papers have also been consulted. Acknowledgment is due Dr. R. A. Harper for suggestions leading to the compilation of the tables. Species which do not appear are omitted because no drawing or definite statement as to the method of conjugation could be found in the literature. TABLE I CONJUGATION SCALARIFORM, SPORE IN ONE OF THE CELLS Zygnema _ (Hassall) Zygnema cylindricum (Transeau) nomalum (Wolle) cruciatum (Wolle) oe és chalvbeieketwiens (West) 2 cyanospermum (Cleve) Collinsianum (Transeau) pe ericetorum (West) THURSTON: SEX IN THE CONJUGATAE Zygnema aries (Wolle) Spirogyra pth (Peti t) leiospermum (Cooke) Juergensti re? s pit hardener (Wittrock) ae plain (Cooke) “ purpureum (Wolle) ty Lagerheimii (Wittrock) ef rhynchonema (West) S. laxa tit de eum (West) <4 longata (Fritsch) St stellinum (Wolle) ~ lutetiana (Wolle) a Vaucherit (Cooke) ue olle) Spirogyra —— (Wolle) Re 3 Fritsch) = Pi pesto (Transeau) i angolensis (West) = neglecta (Pe ss (Wes ie itida (C a bellis (Petit) as orbicularis (Cooke) z Borgeana (Transeau) “s orthospira (Wolle te calospora (Wolle) a parvispora (Wolle) i catenaeformis (Petit) y porticalis ie Ys condensata olle) z phone a- (Wolle) e cr olle) os quadrata (Petit) iy cylindros por. re quinina (Cooke my daedalea (Transeau) " rectangularis (Transeau) ; decimina (Petit x eflexa (Tra a) dil d) as rivularis (Wolle) = dubia (Wolle) iy setiformis (Wolle) 3 ellipsospora (Transeau) a serratum (Hassall) ny Farlowii (Transeau) = Spreeiana (Petit) Sflavescens (Wolle) pte subaequa (Wolle) c Sluviatilis (Wolle) s subsalsa (Wolle) fuscoatra (Wolle) “ (Transeau) es gallica (Petit) Ne tenuissima (Petit) 2 graci Ile) s ternata (Petit) as Grevilleana (Petit) varians (Wolle) " Hantzschii (Wolle) . lata (Wi es Hassallii (West) ion Weberi (Petit) of hydrodictya (Transeau) ay Welwitschii (West) rr tllinoiensis (Transeau) Plagiospermum tenue (Wolle) es inflata (Cunningham) Sirogonium stricta (Wolle) TABLE II CONJUGATION SCALARIFORM, SPORE IN THE CONJUGATION TUBE Zygnema Collinsianum (Transeau) Mougeoiia date (Wolle) i parvulum (Wolle) cillima (Wittrock) «f pectinatum (Wolle) - haere (West) 1 Ralfsit Wolle) i laetevirens (West) Zygogonium psa (Wolle) bs laevis (Cooke) ardhit (Wolle) minnesotensis (Wolle) s © sh aes (Wolle) numuloides (Wolle) zs ericetorum (Cooke) : parvula (Wolle) Ate (Cooke) “t pulchella (Wittrock) ie yynchonema (West) i quadrata (West) As Saeare (Wolle) se sphaerocarpa (Wolle) Mougeotia angolensis (West) ds tenuis (Wittrock) ‘0 : ' tumidula (Transeau) % alcarea (Wittrock) he S a (West) iy ca est) ~ verrucosa (Wolle) ne delicatula (Wolle) is (Wittrock) o divaricata (Wolle) Mesocaras crass: us (Wolle) “ genuflexa (Collins) aiis (Cooke) 445 446 THURSTON: SEX IN THE CONJUGATAE Mesocarpus macrosporus (Wolle) Debarya desmidioides (West) radicans olle) L ptosperma (Wittrock) A recurvus olle) ies ardyi (West) - peesion (Wolle) = laevis (West) aris (Wolle) immersa (West) Debaroa arican (West) ay reticulata (West) mericana (Transeau) . Pyxispora mirabilis (West) % pais bore est Temnogametum heterosporum (West) decussata (Transeau) TABLE III CONJUGATION LATERAL, SPORE IN ONE OF THE CELLS Zygnema ee _- sall) Spirogyra pabtie 57 (Petit) lei m (Cooke) salliit (Wol stell um. m (Wolle ) At acannon (Transeau) Mongoose nummuloides (West) es inflata (Cunningham) a (West) sa intermedia (Hassall) Spirogyra afins (Fritsch) 1 insignis (Cooke) gulare (Hassall) tt Juergensii (West) a bellis e) = longata (Tréndle) ty tanaeformis (Petit) i mirabilis ) ie commu Petit) ae neglecta (T. le) ie condensata (Cooke) e at. aot ag Mes crassa (Cook n quadrata (Co ug — (West) s rectangu lave Gransean c a (West) . Spreeiana (Pet os Page (Cooke) tenuissima esas acilis est) m varians (Petit sf groenlandica (West) - Weberi (Cooke) TABLE IV CONJUGATION LATERAL, SPORE IN THE CONJUGATION TUBE Mesocarpus pleuro oe (Cooke) Mougeotia mirabilis (Wolle) Mougeotia genuflexa (West) Temnogametum heterosporum (West) LITERATURE CITED 1. Bennett, A. W. Sexuality among Conjugatae. Jour. Bot. 29: 172... 1891. 2. Bessey,C.E. Sexuality in Zygnemaceae. Am. Nat. 18: 421, 422. 1884. 3. Cunningham, B. Sexuality of filament of Spirogyra. Bot. Gaz. 63: 486-500. pi. 23-25. AOI7. 4. Petit, P. ‘Spirogyra des environs de Paris. Pals 1880. 65. Trndle, A. Uberdie Reductionsteilung in den Zygoten von Spiro- gyra und tiber die Bedeutung der Synapsis. Zeitschr. Bot. 3: 593- 619. pl. 5 +f. 1-20. IQII. 6. West, W. & G. S. Observations on the Conjugatae. Ann. Bot. 12: 29-58. pl. 4,5. 1898. 7- West, G. S. Algae 1. Cambridge. 1916. 8. Wittrock, V. B. On the spore-formation of the Wecuaeee and especially of the new genus Gonatonema. Bih. Kongl. Svensk. Akad. Handl. 5°: 1-180. 1 pl. 1878. : Studies on plant cancers—l, The mechanism of the formation of the leafy crown gall* MICHAEL LEVINE (WITH PLATES I7 AND 18) Smith in 1916 (1) announced a new type of crown gall, con- sisting of leafy shoots, which was produced by inoculating the leaf axils of a plant, where a dormant bud was present, with Bac- terium tumefaciens. Similar leafy crown galls were produced by inoculating the midvein of the leaf of the tobacco, with the bacterial organism. Smith considered this type of crown gall identical with the atypical teratoid embryomata found in the animal. In 1917 (2) he showed further evidence of the power of this organism to produce leafy shoots in fifteen different families of plants. He contends that the leafy tumor is produced by inocu- lating Bacterium tumefaciens into the tissue of a susceptible species in the vicinity of totipotent cells. Levin and Levine in 1918 (3) indicated that these leafy shoots are always secondary and that the crown gall develops first and then a group of crown gall cells become differentiated and give rise to a tissue, an organ, or potentially an entire plant, the leafy shoot. According to these authors, such differentiation of cells of a malignant tumor does not occur in animal cancer. Crown gall represents only one type in the large group of pathological processes known under the general term ‘‘cancer.”’ It occurred to the writer that if, as Smith claims, Bacterium tumefaciens inoculated into the epidermis (epithelium) of a plant gives rise to an epithelioma and a similar inoculation into the cortex or vascular bundles (connective tissue) produces a sarcoma, then the inoculation of a plant in any region of totipotent cells (bud Anlage), which are known to produce leafy shoots under normal conditions should produce them under the added stimulus of Bac- terium tumefaciens much more readily and in greater abundance. *From the Department of Cancer Research of the Montefiore Hospital, New York, Dr. Isaac Levin, Chief. 447 448 LEVINE: STUDIES ON PLANT CANCERS—I It is well known that when Bryophyllum calycinum leaves are detached from the mother plant and are put on moist soil the marginal notches of the leaves, at which totipotent cells are found, develop into leafy shoots and eventually form new plants. For this reason, leaves and stems of B. calycinum were used to study the effect of the bacterium on the leafy shoot formation. METHOD AND MATERIAL The leaves of B. calycinum were detached from the plant and were placed on moist soil in pots in the greenhouse. The marginal notches of one side of the leaf, right or left, upper or lower, were inoculated, by pricking the tissue five to ten times with a deli- cate needle containing a culture of Bacterium tumefaciens from five to forty days old. The uninoculated notches of the opposite side of the leaf, served as controls, after they had been pricked with a sterile needle. Entire leaves with each notch pricked five to ten times with a sterile needle were also used as controls. Leaves in all stages of development were used. The veins of leaves and growing regions of stems of the Bryophyllum were also inoculated with Bacterium tumefaciens. In all over a thousand inoculations were made. It may be stated that thin young leaves did not lend themselves to these experiments, because they dried out too rapidly. It was likewise found that normal embryos develop better in the greenhouse than in the open. As a rule, two days after inoculation with Bacterium tume- faciens both the infected and control notches showed necrotic areas in those regions. It appeared, however, that the wounds infected showed greater areas of dead tissue, which subsequently caused deeper indentations at the margin of the leaf. This was also observed by Levin and Levine (3) for a number of plants, The uninfected or control notches recovered readily and although slight scars were formed, the bud Anlagen in the notches developed into normal embryos forty to seventy days after injury. Inocula- tions were made: (1) into the notches of Bryophyllum calycinum leaves; (2) into the leaf in the vicinity of the notches; (3) into the midveins of the leaf; and (4) into the growing regions of the young stems of this plant. LEVINE: STUDIES ON PLANT CANCERS—I 449 OBSERVATIONS 1. Inoculation of Bacterium tumefaciens into the notches of Bryo- bhyllum calycinum leaves The study of this material shows that in the great majority of cases, the notches infected with Bacterium tumefaciens, instead of causing the development of leafy shoots, formed ordinary crown galls. Fic. 1 represents a young detached leaf forty-five days after having been placed on soil and having had the basal notches inoculated with the bacterium. The apical notches were pricked with a sterile needle and served as controls. The inoculated notches show well-developed ordinary crown galls without leafy shoots, while the control notches on the ventral surface show the beginning of the development of leafy shoots. Fic. 2 represents an older leaf, in which the basal notches were inoculated forty-five days previously. In this case no galls or shoots have as yet been formed at the notches but the control notches are beginning to proliferate and the uppermost notch of the leaf has produced a shoot. Fics. 3 and 4 represent the ventral and dorsal surfaces of a detached leaf seventy days after inoculating the basal notches with the bacterium. All the basal, infected notches show well-developed galls without leafy shoots on the ventral surface (Fic. 3). The majority of the control, apical notches have already developed leafy shoots. Fic. 5 represents an old detached leaf grown on soil, 140 days after the basal notches were infected. The mother leaf is seen in the center of the figure with a number of well-developed shoots coming from the apical notches. The basal, inoculated notches all show well-developed crown galls. In one instance a poorly developed shoot, visible in Fic. 5 over the largest gall, made its appearance. A similar condition is shown in Fic. 7. These leafy shoots appeared much later in the development of the gall. This seems to show the dwarfing and inhibiting effect the crown gall organism has on the growth of the bud Anlage. 2. Inoculation in the vicinity of the notch When the inoculation is made near the notch instead of in it, a crown gall is developed alongside of a poorly developed leafy 450 LEVINE: STUDIES ON PLANT CANCERS—I shoot. This is shown in Fic. 6, which represents a leaf forty- five days after inoculation with Bacterium tumefaciens. The control notch developed a much larger and more vigorous leafy | shoot. FIG. 7 represents an old detached leaf, 140 days after inocula- - tion. Here again several of the inoculations were made near the bud Anlagen. To the left are seen the large plants which have developed from the control notches. At the right, in the fore- ground, are seen three galls; next to the lower ones there appear small dwarfed plants. It appears that the galls have interfered with the normal development of these leafy shoots when compared with the large normal plants seen to the left. A: STdondon in the midvein It may be assumed that the midvein may have totipotent cells which by the inoculation of Bacterium tumefaciens can be stimulated to develop leafy shoots. Forty leaves of B. calycinum, both young and old, detached from and attached to the mother plants, were inoculated with Bacterium tumefaciens by means of pricking the midvein of the leaf with a fine needle. All produced crown galls within a month after the inoculation. Fics. 2 and 3 show the appearance of such galls on young leaves forty days after the inoculations were made. Fic. 8 represents an old leaf attached to the mother plant. A large gall has been formed on the midvein by inoculating it with the bacterium five months previously. The tumor is a charcteristic crown gall consisting of a great number of tubercles. This leaf was carefully guarded in the hopes that these tubercles would produce leafy shoots. Fic. 9 represents the gall shown in Fic. 8, nine months after inoculation. The leaf became detached and withered. The gall has grown considerably larger, taking on a cylindrical shape, and has become covered with numerous tuberosities. No leafy shoots were formed. 4. Inoculation of the growing region of the stem A large number of B. calycinum plants were also inoculated with Bacterium tumefaciens in the growing region of the stem with the object of stimulating there the possible totipotent cells to leafy shoot formation. Fic. 10 (a, b,c, e, f) represents a few of the LEVINE: STUDIES ON PLANT CANCERS—I 451 young plants in which the growing regions had been inoculated four months previously. In one case only has a small leafy shoot been formed (Fic. 10, 0). This, however, appeared after the crown gall had been well established.' The plants are all dwarfed, as may be seen by comparing Fic. 10, a, b, c, e, and f, with d, one of the control plants. SUMMARY AND CONCLUSIONS 1. Bacterium tumefaciens inoculated by pricks of a delicate needle into the marginal notches of a leaf of Bryophyllum calyci- num, where totipotent cells are present, results in the formation of a crown gall as readily as in other plants used for inoculation but without leafy shoots. 2. Inoculation of Bacterium bammefaitens into the tissue of a leaf of B. calycinum in the vicinity of a small bud causes the forma- tion of a gall and interferes with the normal development of the bud or leafy shoot. 3. Inoculation of Bacterium tumefaciens into the midvein of a young or old leaf detached from or attached to the mother plant results in the development of a large gall without the develop- ment of leafy shoots. 4. Inoculation of Bacterium tumefaciens into the growing region of the stem of a young plant causes the development of the ordinary crown gall with the occasional and subsequent devel- opment of a leafy shoot. 5. Bacterium tumefaciens does sit cause the formation of leafy shoots in Bryophyllum calycinum but rather inhibits and retards their normal development, when inoculated into the totipotent cells which appear at the notches of the leaf. LITERATURE CITED . Smith, E. F. Crowngall studies showing changes in plant struc- tures due to a changed stimulus. Jour. Agr. Research 6: 179-182. pl. 18-23. 1916. —— Embryomasin plants. (Produced by bacterial inoculations.) The John Hopkins De Bull. 28: 277-294. pl. 26-53. 1917. Levin, I., & Levine, M. Malignancy of the crown gall and its analogy to Heabre cancer. Proc. Soc. Exp. Biol. and Med. 16: 21-22. 1918. Lael hes - 452 LEVINE: STUDIES ON PLANT CANCERS—I Description of plates 17 and 18 PLATE 17 Fic. . Young leaf of Bryophyllum calycinum, showing well-developed crown galls at ess basal notches, inoculated with Bacterium tumefaciens forty-five days previously. Fic. 2. Older leaf, control notches showing leafy shoots, none appearing at the inoculated notches forty-five days later. The midvein, also inoculated, shows ordinary crown gall. S. 3, 4. Ventral and dorsal a of a leaf, the basal notches and mid- vein 2 which were inoculated with Bac m tumefaciens, showing well-developed galls without leafy shoots. The control, velo notches show normal leafy shoo Fic. 6. Young leaf, in which the inoculations were made near the apical notches, showing a dwarfed leafy shoot with a crown gall attached to it. One of the control notches shows a well developed leafy shoot. Fic. 10. Young plants. In a, b, c, e, and f, the growing regions were inocu- lated; in d, there was no inoculation PLATE 18 1G. 5. Old leaf, inoculated at the basal notches 140 days previously, showing a large crown gall. The control notches show normally developed plants coming from them. Fic. 7. Old leaf, 140 days after inoculations d the notches, showing dwarfed leafy shoots eae in contact with crown galls. “The control notches have a normal plan Fic. 8. Old leaf roe to mother plant, showing a large gall on the midvein spot . inoculating it with Bacterium tumefaciens five months previously. IG. 9. Same leaf now detached, nine months after inoculation. INDEX TO AMERICAN BOTANICAL LITERATURE 1914-1919 m 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 be gt used in the broadest sense. s, and papers that relate exclusively to forestry, agriculture, horticulture, beabiatactaned products of vegetable origin, or laboratory methods are not included, and no attempt i 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 oem 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 ages must take 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 Boteics! Club Apolinar Maria, Hno. Curiosidades botanicas. Bot., Soc. Cien. Nat. Inst. La Salle 2: 85, 86. 1 Ap 1914. Apolinar Maria, Hno. Notes pour servir a une étude comparative entre la flore de la Savane et celle de la France. Bol. Soc. Cien. Nat. Inst. La Salle 1: 9-11. 1 F 1913; 39-43. 1 My 1913; 80-83. 16 Jl 1913; 153-156. 1 N 1913; 2: 17-19. 1 F 1914; 48-51. 1 Mr 1914; 123-125. 1 My 1914; 249-252. O 1914; 295-297. N 1914. Apolinar Maria, Hno. Notes pour servir a une étude comparative entre la flore de la Savane et celle de la France. Bol. Soc. Cien. Nat. Inst. La Salle 3: 9-11. F 1915; 20-22. Mr 1915; 120— 124. S 1915; 4: 29-31. Mr 1916; 121-124. S 1916; 5: 9-13. F 1917. Atkinson, G. F. Collybia ani Peck, and its near relatives in the eastern United States. N. Y. State Mus. Bull. 205, 206: 61- 65. I919. Ball, E. D. The potato leafhopper and the hopperburn. Phyto- pathology 9: 291-293. Jl 1919. : Bean, W. J. Malus rivularis. Curt. Bot. Mag. IV. 15: pl. 8798. Je 1919. A plant found in western North America. ! 454 INDEX TO AMERICAN BOTANICAL LITERATURE Berry, E. W. Fossil.plants from Bolivia and their bearing upon the age of uplift of the eastern Andes. Proc. U. S. Nat. Mus. 54: 103— 164. pl. 15-18 +f. 1,2. 27 O 1917 Ninetéen new species in various genera are described. Berry, E. W. The fossil plants from Vero, Florida. Jour. Geol. 25: 661-666. WN 1917. Berry, E. W. Geologic history indicated by the fossiliferous deposits of the Wilcox group (Eocene) at Meridian, Mississippi. U. S. Geol. Surv. Prof. Paper 108 E: 61-66. pl. 24-26 +f.17, 18. 22 Je 1917, RT sleoeahin Bent leston spp. nov., are described Berry, E. W. Missa fossil iat from northern Peru. Proc. U. S. Nat. Mus. 55: 279-294. pl. 14-17. 1919. Includes Iriartites gen. noy. and one new species in Iriartites, Ficus, Anona, Banisteria and Mespilodaphne. Berry, E. W. Rilly, a fossil lake. Sci. Mo. 5: 175-185. f. 1-3. Au 1917. Brunson, A. M. The first generation cross between two strains of corn bred for high and low ears. Trans. Illinois Acad. Sci. 9: 87-91. f. I-3. 1916. te Chamberlain, C. J. Structure of the adult cycad stem. Trans. IIli- nois Acad. Sci. 4: 145, 146. I9II. Cheyney, E. G., & Wentling, J. P. The farm woodlot. i-xii + I- 343. f. I-62.-° T915. Cole, A. H. Demonstration of the movement of the water in leaves. Trans Illinois Acad. Sci. 4: 145-147. I9QI1. Collins,G. N. Structure of the maize ear as indicated in Zea- Euchlaena hybrids. Jour. Agr. Research 17: 127-135. pl. 16-18 + f. I. 16 Je 1919. Cooper, W. S. Reproduction by layering in the balsam fir and other conifers. Trans. Illinois Acad. Sci. 4: 132. I9gI1I. (Abstract.) Coulter, M. C. A corn-pollinator. Bot. Gaz. 68: 63, 64. f.1. 18 Ji 1919. Dearness, J., & House, H. D. New or noteworthy species of fungi. N. Y. State Mus. Bull. 205, 206: 43-59. I9I19. Shree new species in Anthostoma (1), Asterella (1), Aylographum (1), Dendro- ma (1), Diaporthe (1), Dothiorella (1), Gloniella (1),Glonium (1), i. ella (I), oe (2), Leptostromella (1), Phyllosticta (1), Septoria (1), and S m (1 Dorsey, M. J. A note on the dropping of flowers in the potato. 5 our. | Heredity 10: 226-228. f. 19. 15 Jl 1919. INDEX TO AMERICAN BOTANICAL LITERATURE 455 Dorsey, M. J. Relation of weather to fruitfulness in the plum. Jour. Agr. Research 17: 103-126. p!. 133-15 +f.1. 16 Je 1919. Drechsler, C. Cotyledon infection of cabbage seedlings by Pseudo- monas campestris. Phytopathology 9: 275-282. f. 1-6. Jl 1919. Drieberg, C. A freak papaw (Carica Papaya). Jour. Heredity. 10: 207. 15 Jl 1919. Egan, W. C. The Russian May day tree: Prunus padus var. commu- | tata. Jour. Internat. Gard. Club 3: 277, 278. Je1g19. [Illust.] Eikenberry, W. L. Some notes on the forests of Ogle County. Trans. Illinois Acad. Sci. §: 121-125. f. I. 2. 1912. Ekblaw, W. E. The plant life of northwest Greenland. Nat. Hist. 19: 272-291. Mrigig._ [Illust.] Eldredge, A. G. Photographing flowers and insects. Trans. IIli- nois Acad. Sci. 9: 33-35. 1916. Fernald, M. L. Two new myriophyllums and a species new to the United States. Rhodora 21: 120-124. 19 Jl 1919. M. exalbescens and M. magdalense, spp. nov. Fred, E.B. The growth of higher plants in soils free of microdrganisms. Jour. Gen. Physiol. 1: 623-629. f. I-3. 20 Jl 1919. Fred, E. B., & Haas, A. R. C. The etching of marble by roots in the presence and absence of bacteria. Jour. Gen. Physiol. 1: 631- 638. f. 1-3: 20 JL 1919. Fromme, F. D. Plant diseases in Virginia in 1915 and 1916. Ann. Rep. Virginia Agr. Exp. Sta. 1915 & 1916: 187-192. j. is. Je IQI7. Fromme, F. D., & Thomas, H. E. Black rootrot of the apple. Jour. Agr. Research 10: 163-174. pl. 15-17 +f. 1. 23 Jl 1919. Fuller, G. D. A comparison of certain Rocky Mountain grasslands with the prairie of Illinois. Trans. Illinois Acad. Sci. 8: 121-130. IQI5. Fuller, G. D. Evaporation and plant succession on the sand dunes of Lake Michigan. Trans. Illinois Acad. Sci. 4: 119-125. f. 2-4. IQII. Fuller, G. D. Germination and growth of the cottonwood upon the sand dunes of Lake Michigan near Chicago. Trans. Illinois Acad. Bei. §: .437--143.. f-. 2-6. . 1912; Fuller, G. D. Reproduction by layering in the black spruce. Trans. Illinois Acad. Sci. 6: 91, 92. 1913- 456 INDEX TO AMERICAN BOTANICAL LITERATURE Fuller, G. D. Soil moisture and plant succession. Trans. Illinois Acad. Sci. 7: 68-73. 1914. Gates, F. C. Relic dunes, a life history. Trans. Illinois Acad. Sci. 3: 110-116. 1910. [Illust.] Gleason, H. A. Taxonomic studies in Vernonia and related genera. Bull. Torrey Club 46: 235-252. 31 Jl 1919. Includes the new genus Epmania and 6 new species in Vernonia. Gleason, H. A. The vegetational history of a river dune. Trans. Illinois Acad. Sci. 2: 19-26. 1909. Grove, W. B. Species placed by Saccardo in the genus Phoma. Kew Bull. Misc. Inf. 4: 177-201. 1919. [Illust.] Groves, J. F. Evaporation and soil moisture in forests and cultivated fields. Trans. Illinois Acad. Sci. 7: 59-67. f. 1-5. 1914. Groves, J. F. A method of prophesying the life duration of seeds. Trans. Illinois Acad. Sci. 8: 133-136. Fs Ee, AGTE: Halsted, B. D. Report of the department of botany. New Jersey Agr. Exp. Sta. Ann. Rep. 37: 433-463. pl. I-1r, 1916. Harshberger, J. W. The diversity of ecologic conditions and its in- fluence on the richness of floras. Proc. Nat. Acad. Sci. Phila- delphia 67: 419-425. 24 Au 1915. Harvey, E.M. Evaporation and soil moisture on the prairies of Illi- nois. Trans. Illinois Acad. Sci. 6: 92-99. 1913. Hauman, L. Notes floristiques, Quelques cryptogames, gymnospermes et monocotylédones de l’Argentine. An. Mus. Nac. Hist. Nat. Buenos Aires 29: 391-443. pl. 1-4 + f. 1-3. 10917. Hauman, L. Quelques Orchidées de l’Argentine. An. Mus. Nac. Hist. Nat. Buenos Aires 29: 353-389. f. 1-8. 1917. Four new species are described. Hauman, L., & Vanderveken, G. Catalogue des Phanérogames de l’Argentine. An. Mus. Nac. Hist. Nat. Buenos Aires 29: 1-350- IQI7. Haynes, C. C. List of French Hepaticae collected by Major George H. Conklin, M. R. C. Bryologist 22: 27. pl.z. 15 Jl 1919. » B. B. A Colletotrichum Beas of ss Jour. Agr. Research 10: 157-162. pl. 13, tao 23 Jl a Hitchcock, A. S. Botanical field-work in oe southwestern United States. Smithsonian Misc. Col. 70°: 50-61. f. 57-67. 1919. a INDEX TO AMERICAN BOTANICAL LITERATURE 457 House, H. D. Report of the State Botanist 1917. N. Y. State Mus. Bull. 205, 206: 1-169. pl. 1-23 +f. 1-6. Includes papers by Dearness & House, G. F. Satie and L. O. Overholts, here indexed separately. Hurd, A. M. Some orienting effects of monochromatic lights of equal intensities on Fucus spores and rhizoids. Proc. Nat. Acad. Sci. 5: 201-206. Je 1919. Jones, F. R. The leaf-spot diseases of alfalfa and red clover caused by the fungi Pseudopeziza medicaginis and Pseudopesiza trifolii re- spectively. U.S. Dept. Agr. Bull. 759: 1-38. pl. 1-3 +f. I-4. 19 Je 1919. Jones, L. R., & McKinney, H. H. The influence of soil temperature on potato scab. Phytopathology 9: 301, 302. Jl 1919. Kempton, J. H. Inheritance of waxy endosperm in maize. U. S. Dept. Agr. Bull. 754: 1-99. f. 1-14. 26 Je 1919. Kezer, A. & Boyack, B. Mendelian inheritance in wheat and barley crosses with probable error studies on class frequencies. Colorado Agr. Exp. Sta. Bull. 249: 1-139. pl. 7-9 + f. 1-10. O 1918 Knight, H. G. Effect of altitude upon the composition of forage plants. Trans. Illinois Acad. Sci. 9: 91-95. 1916. Knowlton, F. H. Contributions to the geology and paleontology of San Juan County, New Mexico—y4. Flora of the Fruitland and Kirtland Formations. _U. S. Geol. Surv. Prof. Paper 98S: 327- 353- pl. 84-91. 18 D 1916. Sixteen new species in various genera are described. Knowlton, F. H. The flora of the Fox Hills egies U.S. Geol. Surv. Prof. Paper 98H: 85-93. pl. 15-18. 3Je19 Sequoia magnifolia, seria coloradenis, Po heads stantoni, Rhamnus willardi, Aristolochia coloradensis, Viburnum vulpinum and Phyllites Cockerelli, spp. nov., are described. a F. H. Principles governing ie use of fossil plants in geologic correlation. Bull. Geol. Soc. Am. 27: 525-530. 1 S 1916. Lankester, C. H. Orchids of Costa Rica. Orchid Rev. 27: 84. Je IQI9. Levy, D. J. Preliminary list of mosses collected in the neighborhood of Hulett’s Landing, Lake George, N. Y. Bryologist 22: 23-26. 15 Jl 1919. Lloyd, C. G. Mycological notes No. 57:, 830-844. f. 1388-1412. Ap 1919; No. 58: 814-828. f. 1358-1387. Mr 1919. 458 INDEX TO AMERICAN BOTANICAL LITERATURE Loeb, J. The physiological basis of morphological polarity in regenera- tion—II. Jour. Gen. Physiol. 1: 687-715. f. I-23. 20 Jl 1919. Love, H. H., & Craig, W. T. Fertile wheat-rye hybrids. Jour. Heredity 10: 195-207. f. 1-17. 15 Jl 1919. Lowe, R. L. Collecting in Oklahoma. Bryologist 22: 21, 22. Jl 1919. McDougall, W. B. Some _ interesting hous of Champaign County. Trans. Illinois Acad. Sci. 9: 125-128. f. 1-7. 1916. Mehlhop, M. A Florida smut, Ustilago sieglingiae, in Illinois. Trans. Illinois Acad. Sci. 8: 140-142. 1915. Meyer, R. Die Bestachelung des Echinocactus Corniger P. DC. Monats. Kakteenk. 26: 46,47. Mr 1916. Miller, H. G. Relation of sulphates to plant growth and soma poeiiole Jour. Agr. Research 17: 87-102. pl. g-12. 16 Je 1919. M’Nutt, W., & Fuller, G. D. The range of evaporation and soil moisture in the oak-hickory forest association of Illinois. Trans. Illinois Acad. Sci. 5: 127-137. 1912. Moore, G. T. The Missouri Botanical Garden. Jour. Internat. Gard. Club 3: 281-291. Je 1g1g. _ [Illust.] Mosher, E. The grass flora of Illinois. Trans. Illinois Acad. Sci. 8: 137-139. I915. Orton, C. R., & McKinney, W.H. Notes on some tomato diseases. Ann. Rep. Pennsylvania Agr. Exp. Sta. 1915-1916: [1-9]. 1917. Osterhout, W. J. V. Apparatus for the study of photosynthesis and respiration. Bot. Gaz. 68: 60-62. f. rz. 18 Jl 1919. Overholts, L. O. The species of Poria described by Peck. N. Y.- State Mus. Bull. 205, 206: 67-120. pl. 1-23. 1919. Pepoon, H. S. The cliff flora of Jo Daviess County. Trans. Illinois Acad. Sci. 2: 32-37. 1909. Pepoon, H. S. Peculiar examples of plant distribution a problem in phyto-geography. Trans. Illinois Acad. Sci. 8: 136. 1915. Pepoon, H. S. Peculiar plant distribution. Trans. Illinois Acad. Sci. 9: 128-136. 1916. Abstract. Pretz, H. W. Discovery of Trisetum spicatum in Pennsylvania. Rhodora 21: 128-132. 19 Jl 1919. . Pritchard, F. J., & Clark, W. B. Effect of spraying on early ripening of tomato fruit. Phytopathology 9: 289-291. Jl 1919. INDEX TO AMERICAN BOTANICAL LITERATURE 459 Purdy, C. Pacific coast wild flowers: their history and cultivation. Jour. Internat. Gard. Club 3: 211-231. Je 1919. [Illust.] Ramsey, G. B. Studies on the viability of the potato blackleg or- ganism. Phytopathology 9: 285-288. Jl 19109. Record, S. J. Storied or tier-like structure of certain dicotyledonous woods. Bull. Torrey Club 46: 255-273. 31 Jl 1919. Reed, H. S. Growth and variability in Helianthus. Am. Jour. Bot. 6: 252-271. f. 1-3. Jl 1919. Rehder, A. _ New species, varieties and combinations from the her- barium and the collections of the Arnold Arboretum. Jour. Arnold Arboretum 1: 44-60. Jl 1919. Taxus chinensis and Carpinus Handelii, spp. nov., are described. Roberts, H. F. The founders of the art of breeding—III. Jour. Heredity 10: 229-239. f. 20. 15 Jl 1919;—IV. Jour. Heredity ro: 257-270. 15 Au 1919. Rolfe, R. A. »The true mahoganies. Kew Bull. Misc. Inf. 4: 201- 207. I919. Rolfe, R. A. Wittia panamensis. Curt. Bot. Mag. IV. 15: pl. 8799- Je 1919. A plant from Panama. : {Rose, J. N.] Botanical exploration in Ecuador. Smithsonian Misc. Col. '70?: 43-50. f. 45-56. 1919. Sampson, H.C. An ecological survey of the vegetation of the Illinois prairie—a preliminary report. Trans. Illinois Acad. Sci. 9: 123- 125; 1916, Sando, C. E. Endothia pigments—II. Endothine red. Am. Jour. Bot. 6: 242-251. f. 1-3. Jl 1919. Sargent, C. S. Notes on North American trees—V. Jour. Arnold Arboretum 1: 61-65. Jl 1919. Savage, T. E. On the conditions under which the vegetable matter of the Illinois coal beds accumulated. Trans. Illinois Acad. Sci. 7: 100-110. 1914. Schneider, C. Bemerkungen zur Systematik der Gattung Betula L. Osterreichische Bot. Zeit. 65: 305-312. D 1915.. Schneider, C. Notes on American willows—V. The species of the lotr ta group. Jour. ise Arboretum. 1: 1-32. Jl 1919. isonti sp. nov. is descri CBS C. Uber die ue Gliederung der Gattung Salix. Osterreichische Bot. Zeit. 65: 273-278. D 1915. 460 INDEX TO AMERICAN BOTANICAL LITERATURE Sherff, E. E. Competition and general relationships among the sub- terranean organs of marsh plants. Trans. Illinois Acad. Sci. 5: 125-127. 1912. : Sinnott, E. W. A botanical criterion of the antiquity of the Angio- sperms. Jour. Geol. 24: 777-782. D 1916. Sinnott, E. W., & Bailey, I. W. The evolution of herbaceous plants and its bearing on certain problems of geology and climatology. Jour. Geol. 23: 289-306. Je 1915. Skinner, J. J., & Noll, C. F. Botanical composition of a permanent pasture as influenced | by fertilizers of different compositions. Soil Sci. 7: 161-178. pl. 1,2 +f. 1-4. F 1919. Smith, I. S. Native trees of Morgan County. Trans. Illinois Acad. Sci. 2: 15-18. 1909. Smith, L. H., & Andronescu, D. I. The artificial germination of maize pollen. Trans. Illinois Acad. Sci. 9: 95-101. ‘ 1916. Speare, A. T. The fungus parasite of the periodical cicada. Science II. 50:116,117. 1 Au1gi9 ; Spegazzini, C. Revisién de las Laboulbeniales Argentinas. An. Mus. Nac. Hist. Nat. Buenos Aires 29: 445-668. f. 1-213. 1917. Standley, P.C. A new locality for Senecio Crawfordii. Rhodora 21: 177-120. 19 }l 1910. Stevens, F. L. & Dalby, N. Some phyllachoras from Porto Rico. Bot. Gaz. 68: 54-59. pl. 6-8. 18 Jl 19109. Ten new species are described. Stevens, N. E., & Morse, F. W. The effect of the endrot fungus on cranberries. Am. Jour. Bot. 6: 235-241. f. 1-3. Je 1919. Stone, R. E. A new stem-rot and wilt of tomatoes. pieces 9: 2906-208. Juz, 2. 711619, Transeau, E. N. The occurrence of the rare alga, Gloeotaenium, in Illinois. Trans. Illinois Acad. Sci. 4: 143. 1911. Trelease, W. Botany and commerce. Trans. Illinois Acad. Sci- 2: 84-89. 1909. : Trelease, W. Two leaf-fungi of Cyclamen. Trans. Illinois Acad- Sci. 9: 143-146. Ramularia cyclaminicola and Phyllosticta cyclaminicola, spp. nov. Zeller, S. M., Schmitz, H., & Duggar,B.M. Studiesin the physiology of the fungi—VII. Growth of wood-destroying fungi on liquid media. Ann. Missouri Bot. Gard. 6: 137-142. Ap 1919. BULL. T PCr f ORREY CLUB VOLUME 46, PLATE 17 LEVINE: PLANT CANCERS BULL. TORREY CLUB VOLUME 46, PLATE 18 LEVINE: PLANT CANCERS Vol, 46: No. 12 BULLETIN OF THE TORREY BOTANICAL CLUB DECEMBER, I919 Preliminary notes on the embryology of Reboulia hemisphaerica WILLIAM L. WooDBURN (WITH PLATE 19) The object of this paper is to present a part of a more extended study on the embryology and cytology of Reboulia hemisphaerica (L.) Raddi. Only certain stages in the process of fertilization and in the early development of the embryo sporophyte will be con- sidered at this time. The description of various other stages in the life history together with a summary of the previous literature on Reboulia will be left for a subsequent report. FERTILIZATION The male gamete in the bryophytes may undergo two series of marked morphological changes. The first series includes the steps leading from the non-motile condition of the androcyte (the cell which directly becomes transformed into the mature sperm) through the formation of the actively motile free-swim- ming sperm. The second series includes the reverse steps and occurs in those cases where a sperm reaches an egg. In this second case the motile sperm becomes again non-motile with a - resting nucleus similar to that of the androcyte or androcyte mother-cell. Details of the main stages in the first series have been carefully described for numerous liverworts and mosses. € processes accomplished in the second series, after the sperm reaches the egg, are not so well known. However, soon after [The ButteTin for November (46: 423-460. pl. 17, 18) was issued December 8, 1919.] 461 462 WoopBURN: EMBRYOLOGY OF REBOULIA HEMISPHERICA reaching the egg the sperm nucleus is found to have the form and structure of a nucleus in the resting stage (Fic. 1). Quite similar conditions have been reported by Black (1) and Garber (3) for Riccia and by K. Meyer (4) for Corsinia. The form and structure of the egg and sperm nuclei at this stage are to a certain extent similar (Fic. 1). The egg nucleus is larger and shows a coarser and more open disposition of the chromatin. An unstained area surrounds the nucleolus. A smaller nucleolus is present in the sperm nucleus. The latter as a whole stains somewhat more heavily than does the nucleus of the egg. The accompanying figures are drawn as nearly as possible in that position which is occupied in nature. For instance, the necks of the archegonia at this stage of development project obli- quely downward or toward the substratum. Consequently it will be seen that, in Fic. 1, the sperm nucleus lies in contact with that side of the egg nucleus which is toward the neck of the archegonium. The membrane of the egg nucleus is somewhat infolded along the surface of contact. Compared with corresponding stages in the life history of gymnosperms and angiosperms we know very little concerning the details of nuclear behavior in this and further stages of fer- tilization. EARLY DIVISIONS OF THE EMBRYO The first division wall of the zygote is laid down transversely. Fic, 2 represents the telophase of this first division, with the cell plate in the process of formation. The spindle lies parallel with the longitudinal axis of the venter of the archegonium; conse- quently a transverse basal wall results (Fics. 2 and 3). In Fic. 3 the nuclei are in the prophase of subsequent division; ‘‘a”’ represents the hypobasal cell or that one next tothe base of the archegonium, and b, the epibasal cell or that one next to the neck of the archegonium. The second division is transverse and occurs in the epibasal cell (compare Frcs. 3, 4, 5 and 6). The epibasal cell (Fic. 3, “‘b”’), with the chromosomes clearly dif- . ferentiated, is ina more advanced stage of prophase than the hypo- basal cell. There is also evidence of centrosomes or centrosome- like structures connected with the nucleus. Fic. 4 represents the second division completed. Later stages (compare FiGs- WOODBURN: EMBRYOLOGY OF REBOULIA HEMISPHERICA 463 3, 4, 5,6 and 11) indicate that from the hypobasal cell is devel- oped the foot and from the epibasal cell the stalk and sporangium of the mature sporophyte. A third division follows in either the middle or apical cell of the tier of three (Fic. 4) which results from the first and second divisions. Compare Fic. 5, which shows a tier or series of four cells, with Fic. 4. Then follows (Fic. 6) a division in the basal or foot cell at right angles to the first three division walls. The foot at this stage has become quite dense in protoplasmic contents. The order of divisions just described seems to represent the usual conditions. Compare, however, Fics. 2-6 with Fics. 7-10. In each section represented by Fics. 7 and 9 there is a triangular- shaped apical cell, while in Fics. 8 and 10 both apical and basal cells of triangular shape are present. FG. 11 represents a slightly different condition, in which the foot has become divided into an irregular group of cells. In no case do we find the same sequence of early divisions as described by Cavers (2). In speaking of the early divisions of the sporophyte of Reboulia, he says, “The transverse basal wall is followed by two sets of nearly equal vertical walls which intersect each other at right angles, so that the embryo shows a regular octant stage.”’ SUMMARY. The egg and sperm nuclei are both in a resting condition in the earliest stages of fusion. Among the Bryophytes little is known concerning the details of nuclear behavior during the stages of fertilization. The earliest divisions of the zygote are transverse. A longi- tudinal series or tier of four cells may be formed. Occasionally, however, both apical and foot cells of triangular outline may be formed, or the basal cell may divide into an irregular group which constitutes the foot. Of the first two cells formed, the hypobasal cell evidently produces the foot, and the epibasal cell the stalk and sporangium of the mature sporophyte. ~ NORTHWESTERN UNIVERSITY, Evanston, ILLINOIS 464 WoopsurRN: EMBRYOLOGY OF REBOULIA HEMISPHERICA LITERATURE CITED 1. Black, Caroline A. The morphology of Riccia Frostii Aust. Ann. Bot. 27: 511-532. pl. 37, 38. 1912. 2. Cavers, F. Contributions to the biology of the Hepaticae. Part > I.—Targionia, Reboulia, Preissia, Monoclea. Leeds and London. March, 1904. 3. Garber, J. F. The life history of Ricciocarpus natans. Bot. Gaz. 37: 161-177. pl. 9, 10 +f. I-g. 1904. 4. Meyer, K. Untersuchungen iiber den Sporophyt der Lebermoose. Entwickelungsgeschichte des Sporogon der Corsinia marchantioides. ull. Soc. Imp. Nat. Moscou 1911: 263-286. gh. at + fi 4-22: 1912. Explanation of plate 19 The figures were drawn by Miss Mary C. Blair. Fic. 1. Early stage of fusion of the egg and sperm nuclei, X 1775. Fic. 2. Telophase of first division of zygote, X 565. Fic. 3. First divisio& of zygote completed; hypobasal cell ‘‘a,’’ and epibasal cell ““b” in prophase of subsequent division, x Fic. 4. Division of epibasal cell completed, sliewne tier of three cells, X 565- Fic. 5. Young embryo 0 consisting of tier of four cells, X 565 Fic. 6. Hypobasal cell divided longitudinally; cell next above in process of longitudinal division, x 505 F Young embryo itt wedge-shaped apical cell, K 735. Fic. 8. Young embryo with both basal and apical cells triangular in shape, FiG.0; a older than in any of the preceding figures, with wedge-shaped aye cell <5 Fic. 10. . more advanced embryo than in Fic. 9, showing both apical and basal cells triangular in outline, X 565. 1G. 11. Embryo showing early differentiation of foot “a,” seta “‘b,” and capsule ‘‘c,” X 565. / The development of the endosperm in Vaccinium corymbosum NeiLt E. STEVENS (WITH FOUR TEXT FIGURES) The occurrence of a “‘chambered” embryo sac, formed by the development of a transverse wall following the first division of the primary endosperm nucleus, has been reported in four genera of the Ericaceae. In recent studies of the high bush blueberry, Vacci- nium corymbosum L., the writer has noted a variation from this of sufficient interest to warrant brief publication. In 1849 Hofmeister (2) published an account of the develop- ment of Monotropa Hypopitys. According to his descriptions and figures (pl. r2, f. 11) the embryo sac in this species is first cut in two by a cross wall near the middle; afterwards cross walls are formed in each half and the micropylar end is cut off by a wall, thus resulting in an embryo sac of five superimposed cells. Nine years after the publication cited above Hofmeister described the development of Vaccinium Myrtillus and Pyrola rotundifolia (3), in both of which he found that the development of the endosperm took place as in the Monotropa without free nuclear division. Koch (4), in his studies of Monotropa Hypopitys, largely con- firms the work of Hofmeister. His figure (p/. ro, f. 12) closely resembles that of Hofmeister except that he finds no cross wall near the micropylar end and thus figures a four-chambered, in- stead of a five-chambered, embryo sac. In the trailing arbutus, Epigaea repens, the writer (5) found a condition much like that reported in other genera. His figure of the Epigaea (f. 3) very closely resembles Koch's figure of the Monotropa, a fact which the writer apparently overlooked in his earlier paper (5, p. 540). The writer also reported the occurrence in the Epigaea of haustoria extending out into the tissues of the integument from the ends of the developing endosperm. The slides of Vaccinium corymbosum, on which the present notes are based, were all made from material collected at East 466 STEVENS: ENDOSPERM IN VACCINIUM CORYMBOSA Wareham, Massachusetts, during June, 1916. The material was fixed in a solution of equal parts glacial acetic acid and absolute alcohol, imbedded in paraffin, cut and stained in the usual way. From a study of this material it is apparent that the development of the endosperm of this species may begin in two quite different ways, either by the formation of a cross wall following the first division of the primary endosperm nucleus as has been reported in the other genera of the Ericaceae, or by a period of free nuclear division as has been described in a very large number of species. Fic. 1 shows a typical two-chambered embryo sac much like those described in Monotropa and Epigaea. The material ex- amined also showed the four-chambered stage which characteris- tically follows this. Fic. 2, on the other hand, illustrates an embryo sac which has developed by free nuclear division. The stage represented by Fic. 3 may obviously have resulted from free nuclear division followed by the beginning of wall formation near the center of the embryo sac, or there may have been some free nuclear division after the formation of the cross walls. Fic. 4 shows a somewhat more advanced stage, in which the irregular arrangement of the cell walls suggests that there was a period of free nuclear division. In this figure the layer of small cells with dense protoplasm (shaded) represents the ‘tapetum” mentioned in Epigaea (5, f. 4). The developing haustoria may also be noted. The antipodal haustorium consists of only one cell, while the micropylar haustorium contains three. In the mature seed of Vaccinium corymbosum the haustoria are larger than in the seed of Epigaea repens; their development also begins relatively early. In Epigaea when the embryo is in the eight- celled stage and the endosperm well differentiated both haustoria are still small and consist of but a single cell, whereas in Vaccinium — the haustoria have attained a considerable size before any divisions of the fertilized egg are apparent. Hofmeister found that Vaccinium (3, p. 141) differed from the other genera described by him in that after the formation of the first wall across the embryo sac, endosperm developed only in the antipodal chamber, while in Monotropa and Pyrola endosperm developed in both chambers (see also Coulter and Chamberlain, _ I,p. 177). The condition found by the writer in V. —— STEVENS: ENDOSPERM IN VACCINIUM CORYMBOSA oy Se 10t@ 4 - ey eo a; v; : eo i. ovat “DD, , i? ef fe) fs Sma iy 8 Fa) Ks ae S, a <3 oe ee eee foe () a Ne sf ee eo Ee 2 i CE - ae ae aS, SR G. 1. Longitudinal section of embryo sac of Vaccinium corymbosum in two- celled stage; the fertilized egg and the remains of the pollen tube may be seen at the upper end and the remains of the antipodal cells at the lower end. 290. Fic. 2. Embryo sac of V. corymbosum which shows only free nuclear division. x 290. Fic. 3. Embryo sac from the same ovary as that in Fic. 2, showing two cross walls and evidence of free nuclear division. X 290. 1G. 4. Longitudinal section of ovule of V. corymbosum showing few-celled en- dosperm with haustoria developing at either end. The inner cells of the integument are specialized to form a tapetum, being more densely crowded with protoplasm and generally smaller than the other cells of the integument. The walls of the epidermal cells are already somewhat thickened. 170. 467 468 STEVENS: ENDOSPERM IN VACCINIUM CORYMBOSA represents a still further variation and suggests the desirability of a careful study of the development of the endosperm in other genera of this family. The writer has, however, been able to examine only one. Slides of Kalmia latifolia made from material collected at Vienna, Virginia, on May 28, 1916, showed embryo sacs before division and in the two-celled and four-celled stages. Material collected August 13, 1916, showed the endosperm with haustoria at either end, the whole condition very closely re- sembling that described in Epigaea. The significance of the condition described in Vaccinium corymbosum lies of course in the fact that here occur in a single species types of development hitherto associated with different genera or even different families. All the figures were drawn from material collected from a single plant and Fics. 2 and 3 represent ovules in the same ovary. Examination of a large amount of material of the other genera described might result in finding variations as marked as those in the Vaccinium. BUREAU OF PLANT INDUSTRY WASHINGTON, D. C LITERATURE CITED . Coulter, J. M., & Chamberlain, C. J. Morphology of angiosperms. New York. 1903. _ 2. Hofmeister, Wilhelm. Die Entstehung des Embryo der Phanero- gamen. Leipzig. 18409. —_—— Neuere Beobachtungen iiber Embryobildung der Phanerogamen. Jahrb. Wiss. Bot. 1: 82-188. pl. 7-10. 1858. 4. Koch, Ludwig. Die Entwicklung des Samens von Monotropa Hypopitys L. Jahrb. Wiss. Bot. 13: 202-252. 1882. 5. Stevens, Neil E. Dioecism in the trailing arbutus, with notes on the morphology ‘of the seed. Bull. Torrey Club 38: 531-543- j. 24. > WOU. Three South American species of Asterella* ALEXANDER W. Evans The genus A sterella has many representatives on the American continent. Most of the northern species are widely distributed, but the southern and tropical species tend to be more restricted in range. Of the fifteen North American species recognized by the writer in his recent revisionf only two are known to extend into South America. These are A. venosa, which was based on material from Brazil, and A. lateralis, which Spruce collected in Ecuador. No other South American stations for either species have been recorded. In the present paper three species which are not known to extend into North Americaare discussed. Four other South American species have been described by Stephani in his Species Hepaticarum, but no material of these is available at the present time. 1. Asterella chilensis (Mont.) comb. nov. Fimbriaria chilensis Nees & Mont.; Montagne, Ann. Sci. Nat. Bot. II. 9: 41. 1838. Hypenantron chilense Trevis. Men. R. Ist. Lomb. III. 4: 441. 1877. Thallus green, becoming brownish or purplish with age, the ventral scales usually more deeply pigmented, mostly 0.5~1 cm. long and 1.5-2.5 mm. wide, in fertile branches broadening out somewhat at the apex, plane or somewhat concave, the thin and slightly wavy margins not incurved when dry, branching normally by forking, ena ey (according to published descriptions) innovating at the apex, keel broad and rounded; epidermis com- posed of cells with toh ity thickened walls, sometimes with more or less evident trigones, averaging about 35 x 25 uw; pores slightly * Contribution from the Osborn Botanical Laboratory. .? The North American species of Asterella. Contr. U. S. Nat. Herb. 20: 247- 312. 1919. In this paper full descriptions may be found of the following species referred toin the present article: A, elegans (Spreng.) Trevis., A. lateralis M. A. Howe, A. Lindenbergiana (Corda) Lindb., A. Ludwigii (Schwaegr.) Underw., A. tenella (L.) Beauv., and A. venosa (Lehm. & Lindenb.) Evans. - 469 470 Evans: THREE SOUTH AMERICAN SPECIES OF ASTERELLA elevated, measuring (with their surrounding cells) mostly 90-100 u in length and 80-90 p in width, surrounded usually by six (more rarely five or seven to nine) radiating series of cells with two or three cells in each series, radial walls slightly thickened; cells with oil bodies as in A. tenella; green tissue loose especially toward the margins, air chambers in two or three layers in the median portion, those of the dorsal layer sparingly subdivided by supple- mentary partitions and considerably larger than the deeper cham- bers; compact tissue occupying about two thirds the thickness of the thallus in the median portion, thinning out gradually on the sides and extending about halfway to the margins, composed of thin-walled cells without pits (so far as observed); mycorrhiza present; ventral scales ovate to lunulate, reddish purple throughout or sometimes with bleached appendages, marginal slime-papillae very short-lived, cells containing oil-bodies mostly three to five, scattered, appendages usually borne singly, rarely in pairs, nar- rowly subulate, scarcely or not at all constricted at the base, mostly 0.3-0.45 mm. long and 0.06—-0.12 mm. wide, acuminate, entire (or sometimes, according to Stephani, with a basal spine), the cells mostly 40-60 » long and 15-35 u wide: inflorescence paroicous, the antheridia forming a small and vaguely defined group close to the peduncle of the female receptacle: female inflorescence borne on a leading branch, peduncle mostly I-1.5 cm. high, more or less purple, with small clusters of narrow paleae at base and apex but otherwise naked or nearly so; disc of recep- tacle green or yellowish, mostly 2-3 mm. across, bluntly conical and usually three- or four-lobed to about the middle, the lobes extending downward and outward, mostly eight-cleft, the divisions subulate, becoming free with age: capsule wall not studied (hyaline, according to Stephani); spores yellow to brownish yellow, mostly 80-90 in diameter, with a thin, wavy, and minutely crenulate wing, 10-16 « wide along the edges, whole surface covered over with a very fine and irregular reticulum, the meshes mostly I-5 # across, formed by a system of delicate anastomosing lines, spherical face showing in addition a coarse reticulum with meshes 16-20 # across, formed of ridges similar to the wings, plane faces without ridges (so far as observed),-margins of wings and ridges more or less broadened; elaters:pale yellow, mostly 120-200 » long and about 6 u wide, tapering slightly toward the blunt ends, usually with two loosely twisted spirals throughout the entire length. _ Known only from Chile; the following specimen has been examined: | Evans: THREE SOUTH AMERICAN SPECIES OF ASTERELLA 471 CHILE: near Santiago, 1915, NV. Costes (N. Y.).* The type material was collected by Bertero near Quillota; Stephani has since reported the species from Pelaquén, P. Dusén (in Bih. K. Svensk. Vet.-Akad. Hand]. 26 (3°): 17. 1900). According to Montagne’s account the capsules of the original material were immature, making it impossible to give any data about the spores and elaters. The description of the gametophyte, however, is unusually full and discusses certain histological features which were usually ignored at that time. It calls atten- tion, among other things, to the thallus broadening out from a narrow linear base; to the elevated pores, making the epidermis appear undulate in cross section; to the large air chambers in the green tissue, arranged in a single layer; to the coarsely tuberculate receptacle, three- or, rarely, four-lobed to the middle, with veiny, truncate lobes; to the apical paleae of the peduncle; to the rela- tively short pseudoperianth, with six to eight divisions, free at maturity.. There is little to criticise in his account, except that the undulate appearance of the epidermis is not always striking and that the air chambers are in more than one layer in the median portion of the thallus. As a matter of fact they are in one layer toward the margin, and the more deeply situated median cham- bers are difficult to demonstrate in dried material. Stephani, in his description, assigns a paroicous inflorescence to the species, noting the androecium at the base of the peduncle, and adds that the spores are yellow, rough, and broadly winged and that the elaters are hyaline and bispiral. He places the species in the same group as A. tenella, A. macropoda, and A. Lindenbergiana» on account of the shape of the receptacle, which he describes as shortly conical and obtuse at the apex. In general appearance A. chilensis bears a strong resemblance to A. tenella and A. Ludwigit, and the species agree further in their dichotomous branching; in their paroicous inflorescence; in their normally eight-cleft pseudoperianths, the divisions of which become free at maturity; and in their yellow spores with broad wings along the edges and a coarse surface-reticulum, at least on the spherical faces. Of course the structure of the green tissue * In the citation of specimens “‘N. Y." signifies the herbarium of the New - York Botanical Garden and “‘ Y," the herbarium of Yale University. _ 472 Evans: THREE SouTH AMERICAN SPECIES OF ASTERELLA will at once distinguish the Chilean species, the dorsal air chambers being subdivided by supplementary partitions, while those of A. tenella and A. Ludwigii remain undivided. The basal portions of the ventral scales, moreover, have fewer cells with oil-bodies and the appendages are much more slender. The deeply lobed female receptacle is also a distinctive feature, the lobes being much shorter in A. tenella and scarcely evident in A. Ludwigii. In A. venosa, with which the species was compared by Montagne, the thallus is far more delicate, the disc of the female receptacle is flatter, the spores are smaller, and there is no coarse reticulum on the surface. 2. Asterella macropoda (Spruce) comb. nov. Fimbriaria macropoda Spruce, Trans. Bot. Soc. Edinburgh 15: 564. 1885. Fimbriaria canalensis Spruce, l.c. 564. 1885. Fimbriaria Mandoni Steph. Bull. Herb. Boissier 7: 207. 1899. Thallus green above and sometimes throughout but usually with the margin and ventral surface more or less pigmented with purple, mostly 2-3 tm. long and 5-8 mm. wide, plane or nearly so with undulate and often crispate margins, branching by forking and also by apical innovations and intercalary ventral outgrowths, keel narrow but rounded: epidermis composed of cells of somewhat thickened walls, sometimes with distinct trigones, averaging about 28 x 24 mu (exceptional cells sometimes 60 u long); pores somewhat elevated, measuring (with their surrounding cells) mostly 120-140 u in length and 100-120 p in width, surrounded by eight (sometimes seven, nine or, rarely, ten) radiating series of cells with four (sometimes three or five) cells in each series, radial walls distinctly thickened, becoming thinner toward the opening; cells containing oil-bodies as as in A. tenella; green tissue loose, the air-chambers in three or four layers (in the median portion), those of the dorsal layer sparingly subdivided by supplementary partitions and thus appearing about as large as the deeper chambers; compact tissue occupying about half the thickness of the thallus in the median portion, thinning out gradually on the sides but extending scarcely more than one-tenth the distance to the margin, composed of cells with slightly thickened, pitted walls: mycorrhiza sometimes present; ventral scales ovate and long-decurrent, purple through- out or with the appendages and margins more or less bleached, cells containing oil-bodies mostly eight to twelve, scattered, : Evans: THREE SOUTH AMERICAN SPECIES OF ASTERELLA 473 slime-papillae short-lived and inconspicuous, marginal cells smaller and more irregular than the median cells, appendages one or two, usually distinctly constricted at the base, narrowly ovate to lanceolate, mostly 0.35-0.75 mm. long and 0.15-0.35 mm. wide, the apex rarely rounded, usually obtuse, acute or api- culate, the margin entire or vaguely crenulate from projecting cells, cells averaging about 45 x 25 uw, one or two smaller cells with oil-bodies often present: inflorescence autoicous: male inflores- cence borne on a very short ventral branch (so far as observed), consisting of a cluster of antheridia, variable in number, and destitute of marginal paleae, ostioles short: female inflorescence variable in position, sometimes borne on a branch of a dichotomy, sometimes on a ventral branch, variable in length; peduncle mostly I.5-5 cm. long, with very long scattered paleae and an apical cluster, more or less pigmented with purple; disc of receptacle often purple, mostly 4-7 mm. across, delicate in texture, the center depressed-hemispherical, mostly four-lobed to about the middle, the lobes spreading almost horizonfally, broadening out and separated by sharp sinuses, upper surface covered over with coarse tubercles, making the margins appear crenate, involucres green to purple, undivided, entire or nearly so, not reaching the margins of the lobes; pseudoperianth extending downward and outward, yellowish brown at the base and rarely throughout, usually for the most part deep vinous purple, mostly twelve- to sixteen-cleft, the divisions becoming filiform upon drying, coherent at the apex: capsule brown to purple, circumscissile by an irregular line; spores brown, translucent, mostly 80-90 in diameter, wit thin and wavy, minutely crenulate wings 12-14 wu wide along the edges, entire surface minutely and closely punctulate or with short and irregular lines, spherical face showing in addition a coarse and usually regular reticulum, the meshes mostly 20-30 u across, enclosed by the marginal wings and a series of similar anastomosing ridges, each plane face with a similar but often incomplete reticulum, margins of wings and ridges darker and somewhat thicker, marked by subparallel lines, and often showing minute interstices especially at points of junction; elaters brown, variously curved, mostly 300-360 u long and 8-10 u wide, tapering slightly to the blunt ends, mostly bispiral throughout. On rocks and banks of streams; known only from the Andes. The following specimens have been examined: Ecuapor: Quito, W. Jameson (N. Y., listed by Mitten, as Fimbriaria elegans, in Jour. Bot. & Kew Misc. 3: 361. 1851); Pichincha, R. Spruce (N. Y.; type of Fimbriaria macropoda, 474 Evans: THREE SoUTH AMERICAN SPECIES OF ASTERELLA distributed in Hep. Spruceanae); Canelos, R. Spruce (N. Y., Y.; type of F. canelensis, distributed in Hep. Spruceanae). Stephani lists Fimbriaria Mandoni from the following locality: Bo.ivia: Sorata, G. Mandon. In proposing Fimbriaria macropoda as a new species Spruce cited no specimens except those which he himself had collected in rivuli ripis montis Pichincha.” In Stephani’s monograph the species is still restricted to the ‘‘Andes quitenses,”” but Jame- son, Lindig and Wallis are mentioned as collectors, in addition to Spruce. Stephani states, in fact, that he had not seen Spruce’s original material at all. Unfortunately he gives no further infor- mation about the specimens which he cites, but it is possible to draw certain inferences from the fact that he does not allude to Jameson, Lindig or Wallis elsewhere in connection with the genus. There is a probability, for example, that Jameson’s specimens are those from Quito, listed"by Mitten under the name F. elegans. There is a further probability that Lindig’s specimens are those which Gottsche* referred to F. Lindenbergiana on account of their violet-colored capsules. They were collected at Boqueron and Tocarema, in the province of Bogoté, Colombia; and, although these stations are not actually in the ‘Andes quitenses,’’ they are assuredly in the same general mountainous district. There is also a possibility that Wallis’s specimens may have come from Colom- bia, where most of his South American Hepaticae were collected. Unfortunately the writer has been unable to consult the specimens in question, so that it is impossible to support these inferences by direct evidence. In Spruce’s original description many of the distinctive features of the species are clearly brought out. The delicacy and translucency of the thallus, for example, are emphasized and attention is called to the purple pigmentation of the margin and ventral surface; to the small number of epidermal pores present; to the narrow midrib; to the very long peduncle of the female receptacle, with scattered paleae and a denser cluster at the apex; to the deeply four-lobed disc, covered over with tubercles; to the membranous involucre, narrower than the lobes; to the rose- purple segments of the pseudoperianth, connate eae . * Ann. Sci. Nat. Bot. V. r: 187. 1864. | EvANs: THREE SoUTH AMERICAN SPECIES OF ASTERELLA 475 to the large spores with the outer wall pellucid and “‘laxe celluloso”’ (in apparent allusion to the coarse surface-network). Spruce ascribed a dioicous inflorescence to the species, probably because the male branches escaped his notice, and stated further that ventral branches were lacking, that apical dichotomies were very rare, that the female inflorescence was terminal (presumably on the main thallus or on one of its innovations), and that the pseudo- perianth was definitely twelve-cleft. Stephani, in his description, states that the sexual branches are ventral in position and at least implies that forking is less uncommon than Spruce indicates. He definitely assigns an autoicous inflorescence to the species and emphasizes the minute- ness of the male branches. He also describes certain structural features omitted by Spruce, such as the green tissue with narrow air chambers and the elevated epidermal pores, each surrounded ~ by six radiating series of cells with four cells in a series. In this last characterization no allowance is made for variability, and the same criticism would apply to his account of the appendages of the ventral scales. According to his statements these are borne in pairs and are approximate, elongated, parallel, lanceolate, and composed of very irregular cells. In the writer’s experience the appendages are often borne singly—the only condition men- tioned by Spruce—and, although the parallel position of paired scales is sometimes striking, it is by no means constant., Steph- ani’s description of the spores as 63 uw in diameter, yellowish, and broadly ‘‘lobate cristate’’ might also be amplified to advantage. Spruce’s F. canalensis was based on material which grew on wet and shaded rocks. As emphasized in the original description the plants are extremely delicate, and the thallus shows almost no signs of pigmentation except on the appendages of the ventral scales. Stephani throws doubt upon the constancy of these features, suggesting that plants of less sheltered situations might perhaps be more robust, and the writer feels that these doubts are amply justified. It may be further shown that the points of resemblance brought out in the descriptions of F. canalensis and F. macropoda are many and important, while the differences are either insignificant or inconstant, this being true not only of those drawn from the texture and color but also of those drawn from 476 Evans: THREE SOUTH AMERICAN SPECIES OF ASTERELLA other structural features. The writer therefore feels compelled to regard the two species as synonyms and maintains the specific name macropoda because F. macropoda precedes F. canalensis in Spruce’s work. A careful study of the specimens listed above (which include the types of both species) has also shown that this reduction to synonymy is warranted. Among the resemblances mentioned by Spruce the following may be cited as important: the narrow midrib; the terminal female receptacle; the slender paleae of the peduncle, clus- ‘tered at the apex but scattered elsewhere; the tuberculate disc, four-lobed to the middle; and the violet-colored divisions of the pseudoperianth. Stephani adds that the inflorescence in both is autoicous, that the minute male inflorescences are borne on short ventral branches, and that the elaters are bispiral. Among the differences brought out by Spruce it will be sufficient to note the following: in A. canalensis the thallus is elongated, the appendages of the ventral scales are lanceolate-subulate, the peduncle is short and the pseudoperianth is sixteen- (or seventeen-) cleft; while in A. macropoda the thallus is ovate-oblong, the appendages of the ventral scales are obliquely triangular and acuminate, the peduncle is long, and the pseudoperianth is twelve-cleft. Stephani describes or implies. certain further differences in the epidermal pores, in the ventral scales, in the discs of the female receptacles, and in the spores; but a careful scrutiny of his statements, as well as those quoted from Spruce will at once make it evident that these differences would easily come within the range of vari- ability to be expected in the organs concerned. The writer regrets that no specimens of F. Mandoni have been available for study and that the reduction of this species to synonymy might therefore be considered unjustifiable. A careful comparison of Stephani’s descriptions, however, will show that the characters separating it from F. macropoda are exceedingly questionable, and that most of the organs are described in essen- tially equivalent phrases. Perhaps the most important differ- ences indicated are those drawn from the ventral scales and the spores. In F. Mandoni the appendages of the scales are said to be lanceolate, strongly attenuate, and filiform at the apex, and the spores are described as 90u in diameter and broadly lobate- EvANS: THREE SOUTH AMERICAN SPECIES OF ASTERELLA 477 winged; in F. macropoda (as already noted) the appendages are said to be merely lanceolate (nothing further being stated about the apices), and the spores are described as 63 u in diameter and lobate-cristate. It has aleady been shown that the spores of F, macropoda usually (if not always) exceed 63 u in diameter; and it will at once be obvious that the other differences noted are of very slight significance. A. macropoda occupies a somewhat unique position in the genus. The structure of the green tissue and the frequent occurrence of a female inflorescence on a long branch derived from a dichotomy indicates a relationship with A. Lindenbergiana and A. venosa, while the short ventral male branches and the not unusual occur- rence of a female inflorescence on a more or less abbreviated ven- tral branch indicates a relationship with A. elegans. Perhaps the relationship to A. Lindenbergiana is as close as any, the deep purple pseudoperianths being a very striking feature which both species exhibit. The spores, however, are essentially unlike, those of A. macropoda being brown and covered over with a coarse network, while those of A. Lindenbergiana are purple and covered over with a much finer and more irregular network. A. Linden- bergiana is further distinguished by its more extensive compact tissue and sharper keel, by its smaller and usually narrower scale- appendages, by its frequently paroicous inflorescence, and by the shorter lobes of its female receptacle. From A. venosa, with which Spruce compares his species, it differs in the possession of lateral intercalary branches, in its larger epidermal pores and less extensive compact tissue, in its autoicous inflorescence, in its purple pseudo- perianths with more divisions, and in its darker, larger and coarsely reticulate spores. 3. Asterella boliviana (Steph.) comb. nov. Fimbriaria boliviana Steph. Spec. Hepat. 6: 11. 1917. Thallus yellowish green above, usually purple on the ventral surface and along the margin, mostly I—1.5 cm. long and 4-6 mm. wide, more or less concave, especially when dry, the margins vaguely undulate-crispate, sometimes erect or incurved when dry, branching intercalary and lateral (so far as observed), keel broad and rounded: epidermis composed of cells with distinctly thickened walls, sometimes with indefinite trigones, averaging about 48 x 478 Evans: THREE SOUTH AMERICAN SPECIES OF ASTERELLA 24 4; pores elevated, measuring (with their surrounding cells) mostly 100-120 » long and 70-100 uw wide, surrounded by seven or eight ‘(rarely six or nine) radiating series of cells with three (rarely four or two) cells in each series, radial walls more or less thickened; cells containing oil bodies as in A. elegans; green tissue compact, the air chambers narrow, in three or four layers in the median portion of the thallus, those of the dorsal layer abundantly subdivided by vertical supplementary partitions not reaching the epidermis in the vicinity of the pores; compact tissue occupying - little more than half the thickness of the thallus in the median portion, thinning out rather abruptly in the sides and extending about halfway to the margin, composed of cells with slightly thickened, pitted walls; mycorrhiza not observed; ventral scales ovate to lunulate, pigmented throughout with a reddish purple or with more or less bleached appendages, cells containing oil- bodies numerous, mostly thirty to sixty, usually scattered but sometimes in pairs or small clusters, tending to be more numerous scarcely or not at all constricted at the base, mostly 0.7-1.3 mm. long and 0.3-0.35 mm. wide, margin entire or vaguely crenulate or denticulate from projecting cells, acuminate or abruptly api- culate or cuspidate, often tipped with a filament variable in length and variously curved, hooked or contorted, the cells very irregular usually including several with oil-bodies, averaging in the median portion about 70 x 30 yw: inflorescence autoicous: male inflorescence borne on a short and slightly expanded ventral branch, the anthe- ridia forming an irregular median cluster without marginal paleae (so far as observed), ostioles low; female inflorescence borne on a slightly longer and more expanded ventral branch; peduncle more or less pigmented with purple, 1-2 cm. long when well de- veloped, bearing a loose cluster of filiform paleae at the apex and scattered paleae elsewhere; disc of receptacle about 4 mm. wide, bluntly conical, more or less purple, shortly four-lobed, the lobes extending obliquely outward, surface bearing very low and coarse tubercles, especially on the lobes, giving the latter a crenate appearance, involucre not examined; pseudoperianth more or less splotched with purple, mostly eight- to ten-cleft, the divisions subulate, loosely connate at the apex and apparently becoming — free at maturity: capsule wall not examined; spores brown OF yellowish brown, mostly 90-110 » in diameter, with wavy, ™- nutely crenulate wings 12-14 u wide along the edges, spherical face covered over with a coarse and fairly regular network, the m mostly 16-20 pu across, formed by a series of anastomosing ridges Evans: THREE SouTH AMERICAN SPECIES OF ASTERELLA 479 similar to the marginal wings, plane faces with similar but more irregular. networks, surface otherwise irregularly punctate and marked with fine and irregular lines, especially along the broadened out and darker margins of the wings and ridges, sometimes tending to form secondary and finer networks within the meshes of the coarse network; elaters brown or reddish brown, somewhat curved or often nearly straight, mostly 180-240 uw long and 12-16 pu wide, tapering toward the rounded ends, usually with two or three spirals in the median portion and one or two at the ends Known only from Bolivia; the following specimens have been examined: BoLtviA: without definite localities or dates, M. Bang (N. Y.; two specimens, one numbered 1869, the other not numbered). The type locality is described by Stephani as follows: ‘‘ Hab. Bolivia. (Bong. legit).””. In all probability ‘Bong.”’ is a mis- print for ‘‘Bang,’’ and the specimens listed above represent a portion of the original material, but this cannot be proved at the present time. The original description of this distinct and interesting species is far from complete and makes no mention whatever of the epi- dermis, the green tissue, the spores or the elaters. The author emphasizes the robust and rigid thallus, the ventral and short sexual branches, and the large disc (6 mm. in diameter, according to his statements), semiglobose at the vertex and slightly con- stricted. He notes further the presence of four campanulate involucres, with large truncate mouths, without saying anything about the lobes of the receptacle, and he describes the ventral scales as large and purple, bearing a large, broadly oval appendage with a long hooked bristle at the apex. The material examined by the writer shows that the ventral scales are more variable than this description indicates and that the hooked apical bristle, although sometimes distinct, is usually represented by a straight or variously contorted or curved bristle. It shows, moreover, the frequent presence of two appendages. Unfortunately the specimens are insufficient to establish an autoicous inflorescence _ beyond a doubt, only one male branch having been observed; . the features of the involucre have likewise been left undetermined _in order to avoid the sacrifice of one of the few receptacles, but _ there is no reason for supposing that the involucre is in any _ way distinctive. | 480 Evans: THREE SOUTH AMERICAN SPECIES OF ASTERELLA In the structure of the thallus, in the restriction of the sexual organs to ventral branches, and in many of the characters derived from the female receptacles and the spores, A. boliviana shows a close relationship to A. elegans and especially to A. lateralis. It differs from both of these species in the broader and usually longer appendages of the ventral scales, which are usually much more abruptly contracted into an apical cilium and which are further distinguished by shorter cells and by the frequent occurrence of cells with oil-bodies. In the basal portions of the scales the cells with oil-bodies are unusually abundant, numbering at least twice as many as in A. elegans and often five to ten times as many. The elaters of A. boliviana, moreover, usually show two spirals at the ends, while those of A. elegans and A. lateralis usually show only one. In distinguishing A. boliviana from A. elegans further charac- - ters of importance may be drawn from the sexual branches, the androecia, and the female receptacles. In A. boliviana the sexual branches (so far as known) are invariably short; the antheridia form a vaguely defined group without marginal paleae; and the female receptacle is covered over with very short tubercles, scarcely apparent in the central portion: in A. elegans the sexual branches vary greatly in length; the antheridia are in a clearly defined and elevated group, surrounded by marginal paleae; and the female receptacle is hemispherical in the center and covered over with longer and more conspicuous tubercles. Some of the characters which separate A. boliviana from A. elegans are shared by A. lateralis, but aside from the important differences derived from the ventral scales and elaters certain other differences derived from the spore markings deserve mention. In A. Jaéeralis, except for the coarse reticulum, the spore surface is covered over with crowded and minute dots but not with lines; in A. boliviana the surface shows both dots and lines, the latter sometimes anastomosing and thus forming secondary reticula within the meshes of the coarse reticulum. SHEFFIELD SCIENTIFIC SCHOOL, YALE UNIVERSITY, INDEX TO AMERICAN BOTANICAL LITERATURE 1914-1919 The aim of this Index is to include all current botanical literature written by Americans, published in seas: or based upon American material ; the word Amer- ica being used in the broadest s Reviews, and papers that oe 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 de 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; eac 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 Acree, S. F. Destruction of wood and pulp by fungi and bacteria. Pulp & Paper Mag. 17: 569-571. 17 Jl 1919. Allen, C. E. The basis of sex inheritance in Sphaerocarpos. Proc. Am. Philos. Soc. 58: 289-316. f. 1-27. 1919. Allen, C. L. Bulbs and tuber-rooted plants. i-vi + 1-311. New York. 1919. [Illust.] Arthur, J.C. Errors in double nomenclature. Bot. Gaz. 68: 147, 148. 15 Au 1919. Bachmann, F. M. Vitamine requirements of certain yeasts. Jour. Biol. Chem. 39: 235-257. S 1919. Bailey, I.W. Phenomena of cell division in the cambium of arborescent gymnosperms and their cytological significance. Proc. Nat. Acad, Sci: §: 283, 284. fi 7. Jl r9r9. Bassler, H. A sporangiophoric lepidophyte from the Carboniferous. Bot. Gaz. 68: 73-108. pl. 9-11. 15 Au 191 Includes 7 new species in Cantheliophorus. Beaumont, A. B. Studies in the reversibility of the colloidal condition ee gee ae Agr. Exp. Sta. Mem. 21: 479-524. Ap 1919. 481 482 INDEX TO AMERICAN BOTANICAL LITERATURE Berry, E. W. Upper Cretaceous floras of the eastern gulf region in Tennessee, Mississippi, Alabama, and Georgia. U.S. Geol. Surv. Dept. Int. Prof. paper 112: 1-177. pl. 1-33 +f. I-12. 1919. Includes descriptions of 42 new species and 3 new genera in various families, several new varieti 1es, and new combinations. Blasdale, W. C. A preliminary list of the Uredinales of California. Univ. Calif. Publ. Bot. 7: 101-157. 14 Au 1919. Bonazzi, A. On nitrification—III. The isolation and description of the nitrite ferment. Bot. Gaz. 68: 194-207. pl. 14. 16S 1919. Bérgesen, F. A new moss collected in Blue Mountains, Jamaica. Bot. Tidsk. 36: 279, 280. 1919. [Illust.] Anoectangium incrassatum sp. nov. Britton, E. G. Mosses from Florida collected by Severin Rapp. Bryologist 21: 27, 28. Jl 19109. Britton, N. L. A large tulip tree struck by lightning. Jour. N. Y. Bot. Gard. 20: 160. Au 1919. Burger, O. F. Sexuality in Cunninghamella. Bot. Gaz. 68: 134-146. 15 Au 1919. Burnham, S. H. Hepaticae of the Lake George flora. Bryologist 22: 32-37. 8S 1919. Burnham, S. H. The sedges of the Lake George flora. Torreya 19: 125-136. 10:5). 1916, Byars, L. P., Johnson, A. G., & Leukel, R. W. The wheat nemotode Tylenchus tritici, attacking rye, oats, spelt, and emmer. Phyto- pathology 9: 283, 284. pl. 18. 29 Jl 1919. Card, F. W. Bush-fruits. i-xiii + 1-409. pl. 1-16 +f. 1-58. . New York. 1919. Carleton, M. A. The small grains. i-xxxii + 1-699. f. 1-181. New York. 1919. Chamberlain, E. B. A herbarium note. Bryologist 22: 39, 40. 85 1919. Clute, W. N. Causes that produce the colors of plants. Gard. Chron. Am. 23: 232, 233. Jl 1919. Clute, W. N. The structure of plants.. Gard. Chron. Am. 23: 475) _ 476, 478. Au IgI9.- Cockerell, T. D. A. The girasole or Jerusalem artichoke. Mo. Bull. : State Comm. Hort. Calif. 8: 243-250. f. ro8-114. My 1919. : INDEX TO AMERICAN BOTANICAL LITERATURE 483 Cockerell, T. D. A. Some western columbines. Torreya 19: 137-141. Io S 1919. Condit, I. J. Bits of fig history in California. Mo. Bull. State Comm. Hort. Calif. 8: 260-265. f. rr8-120. My 1919. Davidson, A. Lupinus subhirsutus n. sp. Bull. So. Calif. Acad. Sci. 18: 80. Jl 1919. Davidson, J. Douglas fir sugar. Canadian Field Nat. 33: 6-9. Ap 1919. [Illust.] Denslow, H. M. Reminiscences of orchid- binds Torreya 19: 152-156. 17S 1919. Dondlinger, P. T. ©The book of wheat. i-xii + 1-369. New York. 1919. [Illust.] Doolittle, S. P., & Gilbert, W. W. Seed transmission of cucurbit mosaic by the wild cucumber. Phytopathology 9: 326, 327. Au 1919. Eames, E. H. Another exceptional specimen of Daucus Carota. Rhodora 21: 147, 148. 7 Au 1919. Earle, F.S Varieties of sugar cane in Porto Rico. Jour. Dept. Agr. & Labor Porto Rico 3: 15-54. Ap 1919. Eastwood, A. Early spring at the Grand Cafion near El Tovar. Plant World 22: 95-99. f. 7, 2. Ap I9g19. Emerson, P. Inoculation of legumes. Idaho Agr. Exp. Sta. Circ. 7: 1-8. Mr 191g. _ [lIllust.] Fairchild, D. The Barbour Lathrop bamboo grove. Jour. Heredity 10: 243-249. f. 1-4 + frontispiece. Je 1919. Fairchild, D. The dramatic careers of two plantsmen. Jour. Heredity 10: 276-280. Je1gig9. [lIllust.] Farlow, W. G., Thaxter, R., & Bailey, L.H. George Francis Atkinson. Am. Jour. Bot. 6: 301, 302. Au 1919. Farwell,O. A. Panicum lineare Linn. Am. Mid. Nat. 6: 49-51. Jl 1919. Fellers, C. R. The longevity of B. radicicola on legume seeds. Soil Sci. 7: 217-232. Mr 1919. Fernald, M. L. A new Polygonum from southeastern Massachusetts. - Rhodora 21: 140-142. a +s 1919. Polygonum puritanorum F 484 INDEX TO AMERICAN BOTANICAL LITERATURE Fernald, M.L. Theidentity of Angelica lucida. Rhodora 21: 144-147. 7 Au 1919. Fernald, M. L. The white-flowered bird’s eye primrose. Rhodora 21: 148. ““ Primula mistassinica Michx., forma leucantha, n. {., corolla lactea.”’ Fernald, M. L., and others. Field trips of the New England Botanical Club 1919. Rhodora 21: 143. 7 Au 1919. Fitzpatrick, H. M. Publications of George Francis Atkinson. Am. Jour. Bot. 6: 303-308. Au 1919. Fitzpatrick, H. M. Rostronitschkia, a new genus of Pyrenomycetes. Mycologia 11: 163-167. pl. rr. 30 Au 1919. Fraser, A. C. The inheritance of the weak awn in certain Avena crosses and its relation to other characters of the oat grain. Cornell Agr. Exp. Sta. Mem. 23: 635-676. f. rog-109. Je 1919. Garrett, A. O. Smuts and rusts of Utah—III. Mycologia 11: 202-215. 30 Au 1919. Georgia, A. E. A’ manual of weeds. i-xi + 1-593. f. 1-386. New York. 1919. Gleason, H. A. Rhamnus dahurica in Michigan. Torreya 19: 141, 142. 10S 1919. Grout, A. J. Moss notes—II. Two pogonatums. Bryologist 22: 37, 38. 8S 1919. [Illust.] Guba, E. F., & Anderson, P. J. Puhyllosticta leaf spot and damping off of snapdragons. Phytopathology 9: 315-325. f. 1-7. Au 1919. Harper, R. A. The structure of protoplasm. Am. Jour. Bot. 6: 273-300. Au 1919. Harper, R.M. Twumion taxifolium in Georgia. Torreya 19: 119-122. 7 Au 1gi9. Harper, R. M. Some vanishing scenic features of the southeastern United States. Nat. Hist. 19: 192-204. 1919. . [IIlust.] Harris, F. S. The sugar-beet in America. i-xviii + 1-342. fl. 1-32 + f. 1-38. New York. 1919. Harshberger, J. W. Alpine fell-fields of eastern North America, Geog. Rev. 7: 233-255. f. 1-12. Ap 1919. Hill, A. F. The vascular flora of the eastern Penobscot Bay region, - Maine. Proc. Portland Soc. Nat. Hist. 3: 199-304. f. 1-6. 1919+ INDEX TO AMERICAN BOTANICAL LITERATURE 485 Hill, J.B. Anatomy of Lycopodium reflexum. Bot. Gaz. 68: 226~231. Re ee Ramm imme (op tt Hoerner, G. R. Biologic forms of Puccinia coronata on oats. Phyto- pathology 9: 309-314. pl. 19, 20. Aug 1918. Hunt, T. F. The cereals of America. i-xxvii + 1-421. New York. 1919. . Illick, J.S. When trees grow. Forest Leaves “a 60-64. Au IgI9. Also published in Am. Forest. 25: 1386-1390. O1 Jennings, D.S. The effect of certain colloidal substances on the growth of wheat seedlings. Soil Sci. 7: 201-215. Mr. 1919. Johnson, A. G., & Dickson, J. G. Stem rust of grains and the bar- berry in Wisconsin. Wisconsin Agr. Exp. Sta. Bull. 304: 1-16 Fo 9-9 5 Au 1919. Johnston, I. M. The flora of the pine belt of the San Antonio Moun- tains of southern California. Plant World 22: 105-122. Ap 1919. Johnson, J. An improved strain of Wisconsin tobacco. Jour. Hered- ity 10: 281-288. f. 8, 9. Je 1919. Kempton, J. H. Inheritance of spotted aleurone color in hybrids of Chinese maize. Genetics 4: 261-274. f. 1-3. My 1919. Kirkconnell, T. W. The flora of Kapuskasing and vicinity. Canadian Field-Nat. 33: 33-35. My 1919. [lIllust.] Knudson, L. Viability of detached root-cap cells. Am. Jour. Bot. 6: 309, 310. Au 1919. Kopeloff, N., & Kopeloof, L. The deterioration of cane sugar by fungi. Louisiana Agr. Exp. Sta. Bull. 166: 1-72. f.z. F 1919. Lange, D. Mysteries and revelations of the plant world. Am. Forest. 25: 1273-1280. Au 1919. _ [Illust.] Lloyd, C. G. Mycological notes 59: 846-860. f. 1413-1443 + frontis- piece. Je 1919. ; The specific characters of Eragrostis peregrina and its two allies. Rhodora 21: 133-140. 7 Au 1919. Long, C. A. E. Notesfrom Matinicus. Rhodora 21:148. 7 Au IgI9. Long, F.L. The quantitative determination of photosynthetic activity in plants. Physiol. Researches 2: 277-300. Au 1919. Marsh, C. D. The loco-weed disease. u. S. Dept. Agr. Farmers’ Bull. 1054: I-19. f. 1-11. Au 1919. 486 INDEX TO AMERICAN BOTANICAL LITERATURE Matz, J. Citrus spots and blemishes. Porto Rico Dept. Agr. Exp. Sta. Circ. 16: 1-8. My 1919. [Illust.] McKinney, H. H. Nomenclature of the potato scab organism. Phy- topathology 9: 327-329. Au 1919. Meech, W. W. Quince culture. 1-180. f. 1-145. New York. 1919. Merrill, E. D., & Wade, H. W. The validity of the name Discomyces for the genus of fungi variously called Actinomyces, Streptothrix, and Nocardia. Philip. Jour. Sci. 14: 55-69. Ja 1919. Miller, W.L. Polyxylic stem of Cycas media. Bot. Gaz. 68: 208-221. jc. f-T1. 165 for, Moxley, G. L. Petalody of the stamens in Eschscholtzia. Bull. So. Calif. Acad. Sci. 18:79. Jl 1919. Murrill, W. A. Bahama fungi. Mycologia 11: 222, 223. 30Au 1919. Includes Polyporus Bracei sp. nov. Murrill,W.A. Fungifrom Ecuador. Mycologia 11: 224. 30 Au 1919. Murrill, W.A. A new species of Lentinus from Minnesota. Mycologia II: 223, 224. 30 Au Ig19. Lentinus Freemanii Murrill. Murrill, W. A. Queer fungus growths. Mycologia 11: 225, 226. f. I. 30 Au I919. Myrick, H. The American sugar industry. 1-v + 1-216. New York. 1919. [Illust.] Nash, G. V. Hardy woody plants in the New York Botanical Garden. Jour. N. Y. Bot. Gard. 20: 11-14. Ja 1919; 41-45. F 1919; 67-71; Mr 1919; 87-90. Ap 1919; 107-111. My 1919; 128-132. Je 1919; 144-148. Jl 1919. Nichols, G. E. Additions to the list of Bryophytes from Cape Breton. Bryologist 21: 28. Jl 1919. Orton, C. R. Notes on some polemoniaceous rusts. Mycologia 11: 168-180. 30 Au 1919. Payne, T. The California wild garden in Exposition Park, its history and objects. Bull. So. Calif. Acad. Sci. 18: 55-77- Jl 1919. Pearson, C. H. The sea grape or uvero tree. Cuba Rev. 17: 14-19. Au 1919. [Illust.] Peltier, G. L. Snapdragon rust. Illinois Agr. Exp. Sta. Bull. 221: 535-548. f. 1-5. Au 1919. INDEX TO AMERICAN BOTANICAL LITERATURE © 487 Pennell, F. W. Scrophulariaceae of the local flora—I. Torreya 19: 107-119. 7 Au 1919;—II. Torreya 19: 143-152. 17 S 1919;—III. Torreya 19: 161-171. S$ 1919. Includes Veronica Brittonit Porter and V. glandifera Pennell, spp. nov., and several new varieties and combinations. Powell, O. Insect enemies and diseases of the tomato. U.S. Dept. Agr. Circ. 40: 1-18. f. 1-23. Je 1919. Reed, H. S. Certain relationships between the flowers and fruits of the lemon. Jour. Agr. Research 17: 153-165. f. rz. 15 Jl 1919. Reinking, O. A. Phytophthora faberi Maubl: the cause of coconut bud rot in the Philippines. Philip. Jour. Sci. 14: 131-151. pl. 1-3. Ja 1919. Rock, J. F. The arborescent indigenous legumes of Hawaii. Hawaii Agr. & Forest. Bot. Bull. 5: 1-53. pl. 1-18. 9 Je 1919. Rock, J. F. The Hawaiian genus Kokia a relative of the cotton, Hawaii Agr. & Forest. Bot. Bull. 6: 1-22. pl. 1-7. 9 Je 1919. Rose, J. N. Botanical explorations in Ecuador in 1918. Jour. N. Y. Bot. Gard. 20: 155-160. pl. 234, 235. Au 1919. Russell, G. A. Effect of removing the pulp from camphor seed on germination and the subsequent growth of the seedlings. Jour. Agr. Research 17: 223-238. pl. 20, 21 +f.1-5. 15 Au 1919. Sampson, A. W. Plant succession in relation to range management. U. S. Dept. Agr. Bull. 791: 1-76. pl. 1,2 +f. 1-26. 27 Au 1919. Sayre, J.D. Factors controlling variations in the rate of transpiration. Ohio Jour. Sci. 19: 491-509. f. 1-9. Je 1919. Schaffner, J. H. Unusual dichotomous branching in Vernonia. Ohio Jour. Sci. 19: 487-490. f. 1. Je 1919. Schneider, C. Notes on American willows—III. A conspectus of American species and varieties of sections Reticulatae, Herbaceae, Ovalifoliae, and Glaucae. Bot. Gaz. 57: 27-64. 18 Ja 1919. Shaw, T. Forage crops other than grasses. 1-287. f. I-29. New York. 1919. Shear, C. L., & Stevens, N. E. The mycological work of Moses Ashley Curtis. Mycologia 11: 181-201. 30 Au 1919. Shreve, E. B. A thermo-electrical method for the determination of leaf temperature. Plant World 22: 100-104. f. 1,2. Ap 1919. 488 INDEX TO AMERICAN BOTANICAL LITERATURE Smith, E. F., & McCulloch, L. Bacterium solanacearum in beans. Science IT. 50: 238. 5S 1919. Segazzini, C. Relique mycologicae tropical. Bol. Acad. Nac. Cien. Cérdoba 23: 365-609. 1919. Includes a number of new species in various genera. Stakman, E.C. The black stem rust and the barberry. Yearbook U.S. Dept. Agr. 1918: 75-100. pl. 1-9 + f.7. 1919. Standley, P. C. Studies of tropical American phanerogams. No. 3 Contr. U. S. Nat. Herb. 20: 173-220. 1919. Includes the new genus Randia and 65 new species in various genera. Stevens, F. L., & Dalbey, N. A parasite of the tree fern ( Cyathea). Bot. Gaz. 68: a ‘a 35, 16... 16:3 1010. Griggsia gen. nov. is descri Stout, A. B.° Intersexes in Plantago lanceolata. Bot. Gaz. 68: 109-133. pl. 12, 13. 15 Au Igio. Stover, W. G., & Coons, G. R. St. Louis Conference on take-all and flag smut of wheat. Phytopathology 9: 330-332. Au 1919. Taylor, A. Mosses as formers of tufa and of floating islands. Bryolo- gist 22: 38, 39. 8S 1919. Téllez, U. Resefia histérica de los estudios botanicos en Colombia. Bot. Soc. Cien. Nat. Inst. La Salle 1: 106-109. 15S 1913; 220. I Au 1914. Thurston, H. W. Jr. Puccinia antirrhini. Phytopathology 9: 330. 1919. Tracy, W. W. Tomato culture. i-x + 1-150. f. 1-43. New York. I9IQ. The chapter on tomato diseases contributed by W. A. Orton. Not indexed separately. Transeau, E. N. Science of plant life. i-ix + 1-336. f. I-Tog. Yonkers-on-Hudson. 1919. Twiss, W. C. A study of plastids and mitochondria in Preissia and corn. Am. Jour. Bot. 6: 217-234. pl. 33, 34. Jl 1919. Van Fleet, W. Progress in breeding freesias. Jour. Internat. Gard. Club 3: 232-239. Je 1919. [Illust.] Vestal, A. G. Phytogeography of the eastern mountain-front in Colorado—I. Physical geography and distribution of vegetation. Bot. Gaz. 68: 153-193. f. 1-17. 16S 1919 INDEX TO VOLUME 46 New names and the final numbers of new combinations are in bold face type. ei concolor, 320; grandis, 314 rospora cervina, 25; chlorophana, 25; 7 haga 2 Acer Douglasii, 315; rubrum, Acerates angustifolia, be flridana, 183; Ila, 184; viridifl Peas fusca, 3195 inal, 47, 309 el thera, 201 i ta, 430; paupercula, 439 Agaricus campestris, 57; The sporadic appeara 5 he of non-edible mushrooms in cultures of, 57 Agoseris ati 310; frondifera, 56; mon- tana, 56; glauca, 311; soe baty ce purpurea, 48 Agropyron, 107; biflorum, 43; lance latum, 319; molle, 43; peeudlorepens, : chardsoni, 43; Scribneri, 43; Smithii, 39, 311; spicatum, 43, 49; tener 43 Agrostis antecedum, 430; asperifolia, 300; elata, 431; foliosa, 315; grandis, 300; humilis, 312; hyemalis, 302; idahoensis, 312; oregonensis, 318; oreop! , 305; Thurberiana £7; variabilis, 310 Aira caryophyllea, 426 Albi pr , 263 osa, 44 Allium Cepa, 338; briltum, 3195 ae aia: -f higpareigt ie 44; si e, 44; ursinum, 347; validum, re Alnus iianinta: ee tenuifolia, Alopecurus ari Se geniculatus, 425; pallescens, = 9 pratensis, 425 Alsine eae at borealis, 300; ; calycantha, 3105 | 489 rassifolia, 306; Ses 316; Jamesii, aeta, 31 = gels stricti- re eruslctes 45; occidentalis, 317 Amaranthus Po well, 320; pumilus, 431 ala, 318; Amarella anisosepa monantha, 322; strictiflora, 31 llis formosissima, 353 Ambrosia psilostachya, 47 Amelanchier alnifolia, 307; Botrya apium, 221; canadensis, 221 - he canadensis intermedia, 221; media, 221; laevis, 221, 5 es opetala, 223; n shpnieitelis prema a guinea, 222 — fs) aize—a ool to criticism, 275 Andia a ae da inermis, 264 NDREWS, F. M., & BEA The val Oeeg oe tect in soaking in water and of aération the growth of Zea Mays, 91 fe Saitoh 39; Hallii, 43; laniger, 111; provincialis, 43; n eget coparius, 111; stolonifer, 110, 11 Androsace di filiformis, ‘ties occidentalis, 46; septentrionalis, 303; subumbellata, 46, 30 Anemone cylindrica, 108; Drummondii, 316; globosa, 311 onan Lyallii, 318; Piperi, 320; Rose- 324 Ascites Fetemecoearar bag latifolia, see Ante ennari anaphaloi S, 47; hed aprica, Au; bern Me obits botnet 313; racemosa, 316; rose! ene simplex, 315 iclea . Antiphylla ha cee 304 A pargia au . 42 Apinus alicauis, 315i rapege 308 | Apocynum Se aeRse 490 folium, 306; pubescens, 431; pumilum, 31 cree in Camptosorus rhizophyllus, peed praecox, 264 Aquilegia coerulea, 308; columbiana, deflexus, 45; gracilis, 324; Lambertii, 45, on ich- ardsonii, 45s eitatae” she AE 324; villosus, 3 Aus nical og ctium tomentosum Arctostaphylos Seale: 323; Uva- ursi, ty) Aeetarie "Barked. 311; cephaloidea, 319; congesta, Rats, ie eee i, 317; Fend- leri ie Argentina Anserina, 302; argentea, 312; gala Arisaema fo lad Aristida arizonica, as longiseta, 43; purpura ° Arnica ar f cana, 324; cordifolia, ae dive siflora, 316; et Ong; STF 5.312 acilis, 318; longifoli 10; Menzi nrg 3rs5: mollis, 313; Parryi, 313; pedunculata, 313 . gnaphaloides, ompta, 311 chauxiana, 313; Pattersoni, 47; pean we 426; triden- tata, 255, 263, 311; tripartita a, 313 car gracile, 264 i Gos of Uredineae C., & Mar - B., Grass rusts of unusual Poswiecteg 4tt rata maculatum, lig Poke us acuminatus, Asclepias am plexi ‘eunlia 430; pumila, 46 scyrum gl agorenceny, | 431 n trees and shrubs nn ‘ei zinc sulphate on the wt A comparison of two strains of fung INDEX Asplenium septentrionale, 300; Tricho- ma 00 Aster amplifolius, 324; apricus, 313; 2. ae) os | =] = “gaa: 473 meses 320; conspicuus, 318; diaboli- ; dumosus, 430; Geyeri, 312; Seagtbenicc. 316; Jessicae, 318; laevis, gag Sy g ganus, 37 Porteri, 47; Sayianna, 3193 Wilso: ‘Aunerstia. "Three South American species of, 469 ‘ Asterella boliviana, 477, 480; chilensis, 4690, 4713 elegans, 478, 480; lateralis, AT2, 4775 tealla, 47%, 472; venosa, 469, 472, 47 Astragalus goniatus, 313; striatus, 313; sulphurescens, 45 Atelophragma Forwoodii, 324; glab- riuscula, 324 thyrium ‘alpestre; 309; cyclosorum, 309 | Atragene columbiana, 312; grosseserrata, 319 Avena orm striata, 3 ‘Aeshna albiflorum, 314 43; Mortoniana, 43; oe rae oaxacana, 108; sarothroides, erium tumefaciens, 447-4 st sg ar Careyana, 320; deltoidea, 315; sagittata, Sea ee seacntethachal 316 Bambos 2 latifolia, prengelii, 217; 430 ait; u 209; obtusifolia, 213: S sub- carnifolia, 21 Bartonia iodandra, 423; paniculata, 424 Batrachium Drouetii, 302; flaccidum, 3013; ee 301 auhinia = ata, 264; reticulata, 264 EALS; C.€ ws, ,& effect ih fraps in water and of aération on ike growth of Zea Mays, QI Beckmannia erucaeformis, Ase Belairia mucronata, 264; — Ammonilla, 271; “quinqucloculatis, Vi BW. A EET with tribution of the Matoniacea new Matonidium from ks on the dis- Aspidium viride, 303 INDEX 491 BICKNELL, , The ferns a — plants of Naniucket—XX Bicuculla uni | poren ee eyathophorum. 263; insigne, 263; Botecebian "Eoakat. 318; Lunaria, 300; neglectum, 306; _ lium, 300; sim- plex, 306; virgin + 300 Bouteloua, 39; peewee 43; gracilis, - & oe. “a PX Brief = cara of the species of Kneif- fia, with the characterization of a new A, 36 Bromus brizaeformis, 43, 49; ciliatus, 301; polyanthus, 313; Pumpellianus, ilis, 425 Brown, E. D. W., Apogamy in Camp- tosorus rhizophyllus, 27 Brown, F. B. H., The preparation and treatment of woods for microscopic d na a 250, 43 Bulnesia arborea, 272; ere 272 Bursa Bursa-pastoris, 3 i mucilage or slime formation in, Caesalpinia coriaria, 264; Ebano, 264; echinata, 264; melanocarpa, 264; Sap- pan, aaa: tinctoria, 2 26 Calamagrostis canadensis, 302; Cusickii, rfii, 300; h } pur- purasec Or; Aegremgaa 316; scopulorum, 323i Suksdorfii, Calamovilfa, rake Boe autumnalis, 302; palustris, 302 unnl pad ala, Campanula petiolata, a 49, 311 Campe americana, 310 pi Pn rhizophyllus, Apogamy in, cordifolia, 322; multifolia, 323; oligo- al QO os) tal o * g o = uw ° A © S @ a = ve Et 5 ‘ 438; duritolla, 305; ebu urnea , 307; elynoides, 39, 44, 49; ae sets 320; exilis, 436; filifolia, 313, fissuricola, 320; flava, 316; iKelloeeit 310; lacustris, 304; : 6: : A ’ uginosa, . ; laeviculmis, 314; limosa, livida, ‘ luzulina, 316; oe nsii, 315; rop- , 318; militaris, 305; monile, 43 muricata, 426; nervina, 316; nubicola, obtusata, 44, 311; oreocharis, 44; pachystachya, 313; P. rryana, : paupercula, 0; Peckii, 303, 4; ec pedunculata, 307; pennsylvanica, pa: phacocepala 311; Pipeii — 8; pra 438; raticola, 301; eslii, pine Raynoldsii, 310; retrorsa, heb Richard- 73 1, 44, 311, 312; a i lorum, 313; — 39» 44 ; rum Carui, Carya glabra Cow 225; ovalis hirsuta, 225 Cassia Fistula, 2 265 Castalia Leibergii, Castanospermum australe, 265 tilleja cervina, Oo; exilis, 312; hispida, 313; lancifolia, 313; 1 13; linariaefolia, as lutea, 319; dentalis, 47; ms, 318; pine- orum 314; sesailifiore: pi Suksdorfii, 3 r ii Catabrosa aquatica, 300 Ceanothus Fendleri, 322; prostratus, 314; ve sect 321 Cedrela Ceiba onde. 5 263 Cen Se ce 424; maculosa, 424; niienen. < cones: Studies on plant,— mechanism of the forma oe ore the ___ leafy crown gall, 457 ‘Capnodes aureum, 302 Carapa guianensis, ae vata, 260 Cardamine arenicola, 43 = Srewesi, 3163 Centaurium fas 426 492 — patinense, 265; robustum, 265; 265 tats - Prcatiae Beeringianum, 45; dum mide- m, igen age — eee ndrum, 426; Cercis Pec ch a — 53 Te 26 Cercocarpus ledifolius, BAO: Cetraria islandica c crispa, Cha eis sc ies - oe Cham Chamachatiarta Mei tole. 323 riclym Chamae menum canadense, 200, 434; naa schense, 320 Chaetochloa ah caieaol 431 Characea e, Prelim nary note on a differ- ential sain ning Zi the cytoplasm of, 375 hara coronata, Cheilan ies Fendieri, , 323; gracillima, 315 caries inia as sa a, 4 perri lanceolata, 45; Wheele eri, 4 Chel Gdontane majus, 425 Chenopodium Botrys, 302; desiccatum, a, 45; 0 oe tabularis, 269 ee ma nape 429; Menziesii, ccidentalis Chiogens hispidula, 307, 439 Chlor on Swi nied - thee C I2I Chrysopsis asprella, 47, foliosa, 47; fal- cata, 430; fulcrata, 48; hispida, 48; villosa, 48, 306 URcH, M. B., The oe =~ in Cooperi pac ifica i, 324: Eaton, 322: nii, gta PP ed igen Cissus inci Cladonia asd chordalis, 23; pyxidata, 21 Claytonia Semaepri. 319; lanceolata, 0; virginica at Clevea Rousselian a, 167 Clinopodium corcineuns, 187; vulgare, 306 Ciitorshe dealbata, 60; dealbata defor- mata, one — minor, 60; dealbata sudori Coelo Aas ‘. cteatum, 305 Cobloghiae a actaeifolium, 439; Gmelini, Be Cogswellia macrocarpa, 46; orientalis, 46 Co! san autumnale, 3 Coleosanthus SPP a 313 Collinsia seb eta nae Collomia linearis, 46, 3 Col! _ Additions Fe ek ora of, 5. Colorado, A new nee @aseaeeaa: from, ee INDEX Colorado, nee gh grasslands at dif- i 37 marum palus re optis trifoliata, as trifolia, 438 —e rrhiza Corallorrhiza, 305; macu- lat i peace, 314; multi- hee pacar rum suffruticosum, 318 ah tinctoria, 48 ies rostrata, 438 Coryphantha missouriensis, 46; radiosa, 4 Coumarouna odorata, 265; panamensis, 265; sp, Cracca ——— ra Cratae apposita, 225; Smithii, 225 Crescentia sn 262; Cujete, 262 Crunocallis C 5 Es thess Sulton: 300; occidentalis, 314 Cytisus scoparius, 426 Dactylis glomerata, 302 albergia Brownei, 265; hupeana, 265; latifolia, 265; fe 265; retusa, 265; Sissoo, 265; sp., 265 Dalea akties, . 255, 256 D ibarda repens, 4. anthonia californica, 3 intermedia 3; Parryi, 43; nee ba uniapicets. to Dasiphora fruticosa, 299 ta | Dasystephana Bigelovii, 46; calycosa, 316; monticola, 318; oregana, 315; Porphyrio, 183, 184; tenuifolia, 183, 184 Debarya africana, 446; americana, 446 cal decussata, — des- ; Har- dyi, 446; 1 , 446; imm ro 446; reticulata, 44 phinium Besa) 318, — depaupet~ atum, 315; multiflorum. N 2; Nelsonii, — Sis: uttallianum, ate “occidentale, INDEX 322; Penardii, 45; reticulatum, 322; virescens, Deschampsia rage eel sip aewNy pre North reels The genus, 207 Desmatedon arenaceus, 207, 213; Bushii, pto thecius £ btorque rectus, 208, a i 208, ars; eer egroesee ig pony ges on North ale Ss, 20 Development and structure of the bulb in Co a Drumm gir The, 337 Development of endosperm inium sr ill The, 465 The in 317; 3195 conjugens, iba Draba andina, 323; auriformis, 322; coloradensis, 45; ‘lutea, 311; nemorosa, 302; ni ssxipent cei Dracaena a ede langifotia,, ee poieseae nies 304 is corymbosa ; fissa, 321 Srna: ao glandulosa, 45> a 312; pumila, padbenaainrth aie 307; dita, ip 304; fragrans, 305; 267; denudata 167; hirsuta, mit £793 ior, hid _ nter- pie one ‘hirsu 1773 irrigua, 167-1773, ao det 167-181; Spathysii, Peis phala, 167-178; velutina, 167-1 Eburophyton Austinae, 314 Echinopan sie Pegg 308 Effect of rok anata te: water and of aéra- 493 tion on the growth of Zea Mays, The, I Ekmania, 250; lepidota, 250 Eleo sine acicularis, 301; palustris, 301; ostata, 431 Cader Beene 6 came Elephantopus caroli us, ober elatus, 251; elatus i intermedi, 252; nudatus, 251; tomentosus, Elymus condensa halophilus 18; si us. 311; glaucus, 308; wellii, 319; nitidus, t Enterolobium Schomber: oki, oe sp., mie ebe pubesce Rolaes AREA 465 Epilobium adenneaulon, 3073; 4393 alpi- asp 48; label, 319; Howellii, = acranth salsuginosus, 310 24 effus vutifellces phyllum, 319; stellatum, 311; amy ides, 319 bellatum, 44, 3 Eriophorum pala aa 300; callitrix, 305; Chamissonis, 303; gracile, 301 opacum, 305; Ngavomerant 303; viride- carinatum, 438 Erocallis triphylla, 320 Erxlebenia min eenee Erysimum shetvanik na aerate pts ‘Tiesie. 2653 indica, 265; monosperma, 265; velutina, 26 Erythronium grandiflorum, 319 Euploca sg etre 186; racemosa. 186 Eurotia Janata, 311 Euce ~~ elegans, 3 E "a3 aie uifolia, EVANS, a w.. 'A axonomic ph of Dumortiera, 167; Then So uth Ameri- can s' of Asterella, ea: Evolvulus sericeus, 185 494 Fendlerella he eg 323 Ferns and flowering plants of Nantucket Salts guyanensis, 2 260, fee te chee. £35 Festuca arizo capellata, as ingrata, 43; minutiflora, pe S, ; ‘ctofiora, 43, 302; ontana, 43; sega ny pein Thasbed. 3 114; viridula, Ficus altissima, 269; Benjamina, 269 Filix bulbifera, 300; fragilis, 300; mon- tana, 300 Fimbriaria boliviana canalensis, garia a, 306; bracteata, 308, ovalis, ppg ge ion sees are terrae- novae, 437, 438 Fraxinus caroliniana, 226 Gagea arvensis, 352; stenopetala, 352 48 6 47, 300; ha oie 310; palustre, ‘4303 triflorum INDEX — D., New and old species of Opu 195 Grind erecta, 48; perennis, 48; squar- subalpina, 48 Grosstaria ‘eptanthe, 322 teri, 272; officinale, 272; Gua oe ae yeni Siilokies 269 Guazu ifolia, ~ Gutierrezia Sarothrae Gymnolomia coy sey ie oe sill aaa 302 Ham calyco ° Harbeusis rip se 46 ala a hispida, 47 Hed ci ee um, 307 i ea etn haat dumosum, 430; majus, 430; propinquum, 4. Helianthus Maximiliani, 424; petiolaris, 48, 49, 311 : eliocarpus americanus, 271; appendi- culatus, 271 emieva ranunculifolia, 319 Heracleum lanatum, 300 Heritiera littoralis, 270; minor, 270 H a 319; PLA StdN Rd hispida, 3073 rubescens 269; si imilis, ri Uliacea 269 Hicoria glabra hir 225 Hieracium albert 309; cyno elena a vulg “a um, 318; albiflorum, 318; griseus, 318 HitTcHcock “Pe ais inary n presale ys staining of the piv of Characeae, pas yx Balan Fisliascen ‘eee Gametogenesis and fecu Biation j in Zea | Hes Mays ba of nia and heredity in the endosperm, acini humifusa, 30 ura coccinea, 46; glabra sea ohytum diffusum, 37 intermedium, 311; pumilum, 317; racemosum, 31 Ey amosissi + Sit Genista canariensis, 2 Gentiana angustifolia, 183 Geoprumnon a um, 45 m Fremontii, 46; Parryi, e ~~ — Richardsonii, 308; sissimu Holoc Geu um macrophyilum, 304; oregonense, ari -_ ncisum, 307; rivale, 306; strictum, 43 Genus eae in North America, Gite ‘agaregata, Bens pulchelia, 313; scariosa, 46; sp’ a, 4! Fg 302 ea decorticans, Graphephorum pears 310; muticum, pelea aL sts of unusual structure, 411 Grewia parte 271; laevigata, 271. ge studies in H sulphur- gi pium arboreum, 268; drynarioides, pei us et - Hexuons, 45 Hordeum jubat 2; murinum, 8, 344 Hydrophyilum area 321; capitatum, veropyla irrigua, 171, 177; nepalensis, Tac eae us filifolius, 4 Hyophila fragilis, 216; Se ae 218 Hypenantron chil Hy opivs americana, 429; lanuginosa, 429 Rolfei, 271; tiliaefolia, 271 Ibidium porrifolium, 312; strictum, 303 INDEX Ichanthus pallens, 115 Ichthyomethia piscipula, 255, 266 Tlex nig scares sis, 438 o American Botanical Literature, i } TOL) 151) °180,/'229,2 920, S8te I, 417, 453, 481 phe Zollingeriana, 266 In — ra, 2 edulis, 254, 266 Tonactis Hnaritous, 430 , 310; pseudacorus, 42 315; Nuttallii, 315; Tuckermani, 438 Juncoides comosum, 311; glabrum, 319; intermedium, nig parviflorum, 301; spicatum, Juncus ptt ae pee ater, wah balticus, 438; bufonius, ; bufonius halo- philus, 438; confusus, 313; preta eaa 4313 — ss rior, 44; longi- stylis, 311 Mertensianus, 310 neveciennis, ee Regal 318; Richard- sonianus, 303, 304; Torreyi, 306; Tweedyi, 3233 oo 306 Juniperus siberic Kalmia microphylla, 309 Kalmiella hirsuta, 183 Kelloggia galioides, 316 Kentrophyta tegetaria, 320 Khay 68 ita, 270 ee ok A brief conspectus of the species Kacifia’ 3 ; Alleni, 366, 371; angusti- folia, ery "arenicola, 364, 367; brevi- 372; (iL By 03 chrysantha, enn raseri, 371; fruticosa, | a humifusa. 366, 3713 ligentifolias sox nifolia, 3 eae | xii hybrida, 365, 371; tetragona lon ipata, 365, 371; velutina, 363, as: Koeleria gracilis, 4 Koompassia excelsa, 266 es a 266 na pti pct 48, 49, ge Lactuca multifida, 320; spicata, 3 Lappula usa, 317; ee a Bir: Lappula, 425 365, | | | | | | | 495 Larix occidentalis, 3 Larrea cuneifolia, selon divaricata, 272 $, 438 calcarea, fruticulosa 22; lent sca, 23; thamnoplaca, 22, 25; tar Ree ej L mea Lace, A3i; spent 430 Lecidia amylacea, 24; armeniaca, 24; fect steed. oncaay elobifera, my granu- osa, 21; tessellata, 2 Ledum glandulosum, 31 e Lemn pa , 310; minor, 301; trisulca, 301, jee scopulorum, 322 Lepargyraea c a ateliiie: 209 pete buxifolia, 241 — den siflorum, 45, 306; monta- ep sity 5 Lep aesees Be | Levianteyion Niitellt. ae 1; pungens, 311 Lesquerella m Leucocrinum » 44 tae M., Th he sporadic appearance of no: e in cultures of xaut plan The the bcunien - the leafy crown gall, 44 Lewisia redeviva, 311 Lichens, Notes on some w Ligusticum Canbyi, 320; Leiber, 320; , 312 artagon, 338 _Limn ia scart, 317; perfoliata, 317; sibirica, 317 Limnobotrya montigena, 310 Limnorchis borealis, 312; stricta, viridiflora, 312 Limosella aquatica, 301 : ; 200 312; ; tenuifolia, 301 , 306, ; nnaea americana, 429, 439 longiflora, 314 Linum intercursum, 430; Lewisii, 46, Lithophragma oe 311, 312; par- viflora, 31 , 44 pus ations, 266 Luetkea pectinata Lupinus, Studies in a genus,—1V. The Pusi Lupinus ‘argenteus, 3 1; argillaceus aio 393, eal saevicaali. 380, 390, 396; Bur 318, caespitosus, ea vet 38 pone ; concinnus, densiflorus, ; dispersus, 389, aon 496 flavoculatus, 380, 392, 404; inter- montanus, 389, 392, 408; Kingii, 389, 391, 392, 393; Kingii argillaceus, 3 laxiflorus, 316; leucophyllus, 317; leu- copsis, sl mmalacophy illus, 389, 390; microcar ruber. transmontanus, 390; , 3II Lycopodiu nu site bscurum, 304; aay 304 Lygodesmia juncea, Lysias Menziesii, o orbiculata, 305 tvglets obtusata, 3 Machaeranthera aspera, 48; varians, 48; viscosa, 48, acrocalyx Nyctelea, 306 é Macronema discoideum, 321; suffruti- cosum, 311 Madia See 311 Marns Son Grass . B., ARTHUR, = yuats ~ ‘unusual structur Maize, The ancestry span ne of criti- ism archantia hirsuta, I : 73 trichocephala irrigua, 177; 287— m. Althausii, 285, Fa ameri- 287; Goepperti, 285 Melnoeran lineare, 307 Melica bella, 313; sige 303; — bilis, 313; subulata, 3 Melochia indica esia ferruginea, 317; glabella, 317 Meriolix serrulat a, 46 Mertensia Bakeri, 47, 49; Cana, 47; Clokeyi, ay; lanctolate, 47, 553 lateri: flora, 47; lineariloba, 47; media, 55; osporus, 446; pleuro- carpus, 446; icans, 446 _ 446; robustus, 446; scalaris Mezon gaan gg 266 Mi aestivalis, 319; arguta, 310; Greenei, oti rhomboidea, rant Milletia senaoa: 366 INDEX Mi sein! Breweri, - = Langsdorfii, 310; Lewisii, 310; moschatus, 301; nasutus, 3 eects uloides, ye 5 tella nuda, Mibcincere pee aoe 314 Moehringia latifolia, 301; macrophylla, Monarda menthaefolia, sai pectinata, ata immdcu- ulis, 186 ES Moneses retic ulata, iS uniflora, 300 Monoclea Forsteri Monotropa sera 3s, “gk gene: ie Moringia pterygosperma, 269; sp., Mougeotia angolensis, 445; Hootie: si calcarea, 445; capucina, 445; delicatula, 445; divaricata, 445; genuflexa, 445, 446; glyptosperma, 445; gracillima, 445; irregularis, 445; laetevirens, Back laevis, 445; minnesotensis, ae ira- bilis, 446; nummuloides, 445, 44! vula, 445, 446; pulchelia, 4453 een. 445; sphaerocarpa, 445; tenuis — — la, 445; uberosperma, 4453.9 T- rucosa, 445; viridis, ia or slime formation in the cacti, Mabieibergin comata, 300; pane 309; diffusa, 113; gracilis, $i gracillim a, 43; racemosa, ty eae sonis, 43, 302; subalpina, 4 epermonige divaricatum, 46; * reduanates tum, Myos roan cee me 310 yrica Gale, Myrocarpus 267 Ne helaios 267; frondosus, M mum frutescens, 267 Myroxylon toluiferum, 267 ATahal rs Naias guadalu upensis, 431 Aebsivciie parvifolia, 316 es The ferns and flowering plants 2 rs Matt rages 22 Neslia paniculata, 42 New and old species + oe, 195 New ee from Colorado, with remarks the ‘distribution of, the Mato: jutacene, A, 2 New species of ee XI, 107 itella opaca, 378 ee rth America, The genus Desmatodon 207 Notes on dees western Lichens, 21 e plants of the southern ae states —V, Csi Notes on galt the vicinity of Wash- __ ington, 2 Notholacna Fendt, 323 INDEX Nuttallia hastata, 53 nuda, 53; Rusbyi, 54; stricta, 53 Nymphaea polysepala, 310 — + bra 308 nothera ua, 370; canadensis, wae: pkevees sat 342° florida, 368; Fraseri, 371; fruticosa, 367, 370, 372; fruticosa humifusa, 68; fruticosa incana, phe fruticosa Dayllopis, S715 glauca Ho oa hy brida, a, 369; serotina, pie hana, ne tetragona, 370 yea efirop neaige tohgtielind a 184 MIneya tesota, 256, 2 ye said 3 es ws opordum m, 56 ynosmodium ‘occidentale, 47 50 6.0:00250 Oplismenus hirtellus, 115 ulaster monogynus, 323 oe praecox, 264; sp., 264 { Opun amarilla, 205; Bartrami, 201; ats, 199; cyanea, 196; diversispina. 197; effulgia, 195; elongata, 199; ee 750; iy ragilis, 46; - ocarpa, 199; geen anage ov inermis, pea Maideni a, 204; obovata, ae atarax. 46; strep- ticantha, oe O puntia, New d old eae of, 195 Oreobroma radiate. nosperma, oe thyrsiflora, 147 46 Ormosia calavensis, 267; rubusta, 267 Orophaca tridactylica Orthocarpus luteus, 47, 49, 313; Tolmiei, Oryzopsis asperifolia, 305; Bloo 308; pungens, 303, 3045 Webber Fe 20 Osmorrhiza brevipes, 31 303, 304; — 320; ees pe aon: OSTERHOUT, ic E. Additions to the Flora of Co se Ougei einia dale giokiee 267 305; Oxycoccus, 305, 439 Pachystima Myrsinites, 308 Paeonia Brownii, 317 Panaeolus campanulatus, 59; retirugis, 59; 497 438; nervata, 300; obtusa, 436; septen- trionalis, 300 Panicum albemarlense, 431; auburnae, 431; Bicknellii, 431; co mj T8335 depauperatum, 430; Huachucae, 303; hy m, 23; linearifolium, 423; meridionale, 430; thermale, 316; vir- t oa physod ssia fimbriata, eae: palustris, 307; saivtiion Seonvciix: it 45 =m thia pate soto engeteanes 308 gar adh bracteo: 310 nsis, 306; contorta, abe: aes eo anven pana 04 Peltigera aphthosa, 2 oy are 273% ‘inifotium, 373 ‘species ‘of Kneifia, cr the characteri- of ied genus, 363; Noe on olnate ok Aci southern United States—V., 183 inn polyantha, 271 Pen mon crassifolius, " yallii, pine noaban s5Ft} Rydbergii, Yaa Ae aca. Peramium ion 300; ophioides, 305 I : I a pga um Pereskia Pereskia, 16 Perito tie serrulatum, 313 iyi coccinea, 301; Hartwrightii, 8; a ection a, 431 Peeuinaaeon oligophyllus, 45; purpureus, Petasites nodulosa, 307; palmata, 307; sagittata, brit as dome caespitosum, 311, 312 Peziza ston ian haca a a, 62 a, yreve Hookeriana, 320; serpens a ri ’Sileriana, 323 celia Bakeri, 54; crenulata, 54; formosula, 543 slandulosa, 54; hetero- phylla, 47; sericea, 47, 313 Pha chlorac a; eee" Phalari inacea, 30 di es | Phegopters Phegopteris, ee polypo- pitchaphes Lewisii, 319 Phleum alpinum, 300; pratense, 302 caespitosa, 313; oe ar05 floridana, 185; multiflora, 46 Phoenocaulis cheira nthoides, 315 venenosus, 57 Panicularia borealis, 300, grandis; 301, ountain regi a the Montane plants, 498 Picea canadensis, 124, 307; Engelmannii, mariana, 307; pungens, 321 celsa, 270 ta, Banksiana, Mitiaeen a, 308; Aa dcioea, 408, 355; 1 osa scopulorum, 321 elegans, 3143 multiflora, 314; | unalaschensis, 308 Piptocoma rufescens latifolia, 251 306; major, Purshii, _ me of “the one United States, —V, 18 pe bani dubium, dd > 267 Platypodium Maxonian : tan, — 143, 320; pedunci eo agin eyae, t . 1, 1%; a, triflora, 302; Vaseyochloa, 3:9 Polanisia poeestebie Polemonium confertum, 46; occidentale, 309 Polycodium floribundum, 183 Polygala polygama, 430 iforme, sed, Douglasii, 3033 nnii, 44; pennsylvanicum nesopbilum, ying Pe atc Pg 302; watchense, 44; scandens, 117; Palacall, 310 Polypodium hesperium, 309; vulgare, Polyatichum imple Bat Lonchitis, M, 315; scopulinum, 321 Pongamia ates, ees saith is, 267 Populus angustifolia, 321; balsamifera, 299; tremuloides, 299; Wislizenii, 321, 22 Porliera hygrometra, 272 Bipriarts. gun alpinus, 301; foliosus, 301; Tat a, 45; dichro 43; Dr ndii, 319; e AS) caulis, 322; glomerata, 316 ap gunn Ee 45; Nutt » 313; Ovina, 324; pro- pinqua, 322; strigosa, 45 Preliminary note on a_ differ ential bist of ae cytoplasm of Shae 375 cick notes on the embryology of Reboulia hemisphaerica, 461 295; | 302; | INDEX Preparation _ treatment “ woods for TO scopic study, The, 127 ee mistassinica, 307; Colbbeloust si a 122; deformata 412; farinacea, 118; Oe aa spi et fr cola, 109; ge tilis s, 118; Helianthi, 119; imposita, 112, 411; inclita, 115, 411; invelata, 119; bachii, 110; longi- ornis, 122; m s, 119; mammillata, 117; missouriensis, 108; mitrata, 118; obtecta, 107; pallescens, III, 412, 344: 415; pallida, 111; parca, 117; phakop- soroides, 412-415; Polygoni, 117; prospera, 118; Scribnerianum, 109; rigs nr Bs substriata, I16, vahidtnrcaay — nsis, 107 Pucciniastrum sai es 25 Pulsatilla fone trier 303; occiden- talis, 315 Pyrola Borage 300; bracteata, 320; to} es entata, 316; » 309; perenen 446 amoclidion multifiorum, 320 Quercus pagoda, 225; pagodaefolia, 431; Schumardii, 225 Xeri ophytic c grasslands at epee pr in Colorado, 37 Ramischia . Raminculus’ alismaefoliug, 342, 3105 ongardii, 308; dio: tg iti 312; Douglasii, 308; oltzii, 310; intertextus, 322; micranthus, 3065 INDEX limosus, ht eee 302; recurvatus, rep a saxicola, 323; sceleratus Ratibida ae oes. 48, 4 Razoumofskia am agar mH Douglasii, 312; Laricis, 3 Reboulia Ee ohaweeas -peoaemnld notes on the embryology 61 RECORD, S. J., ried or late struc- Sto: ture of certain a enous woods, 253 ter Houlletiana, sige date 1615 mbea, 159, I 160, ca) ; pet raeum Ribes ee 307; Hudsonianum, 304; ne se, 314; laxiflorum, 315; ongeanthoides ealcicola, 437, pas ; aes! , 319; viscosissimum, 308 iccia, 4 Rinodina cerca aena, 25 Robinia Pseude-Acacia, 143 cky ountain gion, Phytogeo- aot notes on the,—VIII. Dis- ributi nts, 295 Rosa ecedtaatid 303; Bourgeauiana, 307; nutkana, 3 betes orum, 308 Rubus melanolasius, 312; nivalis, 314; a2: en nus, x. pe bepneatts 302 Ruppia oor gia cee P,; went Roe Distbation ee the Montane plants, So eesoors alba, 304; Torreyana, 431 Sabatia Elliottii, 183 Sabi ina horizontalis, 303 saginoides go oS latifolia, 3 316; Bebbi Pp commutata, 319; cordata, _ exigua, dii 312; Fe iy: S10; riana, 318; | glaucops, 309; id Neo tg 318; Lem- mi, 314; lucida, Pst icellaris, 304; pellita, 307; eriana, 308; sitchensis, 314; ie 426; tristis, 4303 Wolfii, 322 Salvia mosana, microphylla, 118 Sambucus coerulea, — melanocarpa, 321; microbo Santalum Freyeinetianan 134 pindus senegalensis, 2 118; fulgens, 121; Sa pecta- | Seric 499 — vermiculatus, 311 ussurea amer 320 Saxifraga simulata, 322 Schedonnardus paniculatus, 44 Scheuchzeria palustris, 436 Schizachyrium littorale, 431; 44; 430, ie nen ga 431 Schizaea pusilla, Schoutenia ovata, se Scirpus americanus, none Scacar: 304; osus 4 scoparium, caes 8 300; Erio ; ntal “ 438; pauc rai 300; pedicelats Bott rub- pumilus, 305; subt Remnant ig 304, pre sie egal Scrophularia occidentalis, 321 Sedum Douglasii, 315; Sa: 319; stenosepalum, 45, 49, spe os densa, 43, ye 3; montanensis, ica, = oe ae 306; Un derw oodii, 321; Watso Senecio filicifolige, 321; pet Mii, 318; oblanceolatus, 48; pauciflorus, 301; perplexus, 48, 322; plattensis, 48; udaureus, 300; ifolius, 322; 1ra, 313; spartioides, 48, 322; sub- gy triangularis, 310; wer- escens, aa 45; mb 318 45; Ly ys 318: steitlaaetlan 318; oregana, em ear 270; a 270 445 Si asim Nasturtium-aquaticum, 302 yrin nchium arenicola, 430 Sitanion vtec oides, 44; montanum, 318 Sium cicutaefolium, = Ow: li Solidago aestivalis, 431; concinna, 48; decu . 48; ma, 306; mis- , 48; nemoralis 430; nana, 48; phila, 48 is mchus arvensis, 425 ora chrysophylla, 268; secundiflora, 500 Sorbus ocsheatalin 3195, scopulina, anal Sorghastrum nutans, 44 South Nps oe species of Asterella, Thre Spuithers United ‘States, Notes on plants | of the,—V, I Sparganium Ser cetlcliaris 301; 300 mini- Ss a perfoliata eae doautees: rales lucida, 317 iro, , various ies of, 445, 44) Sp Sporadic eaginriaieee es no n-edible mus rooms in c pestris, The, 57 Spraguea multiceps, 318 Sprekelia formosissima, 353 TEINBERG, R. A., A of tw ains of t Steironema food eolatum, 424 oe foetida, aa%s populifolia, 271 , N. E., The development of ee pgp in Vacctin corymbos desu se FE. G., Mucilage or slime for- ion in the cacti, 157 Stipa ton Ta 44; Elmeri, 316; minor, As; Nelsonii, 44; oreganensis, 3 316; specios T ar eaeary indica, 315; Tweedyi, py , 322; viridula 320; Thor iana, aseyi, 322; viridula, 44 : Storied or tier-like structure of certain =] ism - the foxthation of ‘he le 1, 44 sg car in gt genus Lupinus—IV. The Pus 8 INDEX 262; pentaphylla, 262; Schumanniana, 262 268 endron, 271; javanica, a Argyrod ag sumatrana, 271; sylvatica, 271 axonom studies in Vernonia and ic rela — — a, 235 axono tudy of Dumortiera, A, 167 Taxus preritolta’ 314 Tecoma araliacea, 262; chrysantha, ~ leucoxylon, 262; obtusata, 262; spe- ciosa, 263; stans, 263, sp., 263 Temnogamet heterosporum, 446 Tessa: ei lum arpum ; Fen cidentale, 3175 sparsiflorum, 308-310 Thelesperma gracilis, 322 s Dryopteris, che Oreopteris, Phe opteris, pest obertiana, 307 313. ea. apie pat coh Ae 317; = aoapriea 45; Nuttallii, 313; purpur- scen rs) Three Soutls American species of Aster- ella, 469 huja plicata, 314 TON, H. W., JR. ee jugatae and the felative frequency of the different ome of c Fe eedon ia Tiarella unifoliata, 316 Tilia americana, 23 cordata, 272; heter- 272; THU pe andshurica, 272; Mich- 225;n neglect, 2 226; pot a ns, 272 Uiseaniuas aqua m, 302 Tiniaria scande Tipuana speciosa, 268; sp., 268 Eta signing 46 00; Drummondii, 46, 4: ofeldia Hemet ast A, 316; ceienta te Stud ase some factors canine cing the stimulative action of z sai on | T nebo pygmaeus, 48 th f Aspergillus Il. | Torresia odorata, 301 A a of two seratas of the! Tortula bryoid 219; Gu con 2IT; fungu obtusifolia, 213; ae a, Sabilnen: ey 301 Townsendia egetanter ee 48; Sullivantia Hapemanii, 306 scapigera, 31 iana maritima, 27 Toxicoscordion gramineum, 44 Svida instolonea, 321 Tragopogon pratense, 425 Swartzia Gaillardi, 268; panamensis, | Trautvetteria grandis, 314 tomentosa 268 Chelan ma melaleucum, 61 ertia congesta, Trichostomum meg oe Peeves 19 Swietenia Mahagoni, 269 Trifolium Beckwithii, 316 um, 426; Syntherisma Ischaemum, 302 longipes, 3155 pleesaeen, pie Rusbyi, Synthyris laciniata, 324; major, 320 320; scariosum, 324 Triglochin mari maritima; 301; palustris, 300 Tabebuia Donnell-Smithii, 262; Guay-| Trillium ovatum, 3 acan, 255, 262; nodosa, 262; Palmeri, | Trisetum majus, 44; — 44 INDEX Tripsacum dactyloides, 112; lanceolatum, 2p ee atifolium, III 6 hetero Tulipa sylvestria, 353 Unifolium dilatatum, 314 Uredineae, new species of, —XI, 10 Ured a Sots Wh 21. pre ar I21; Fuirenae, 100; oe Tet, ATE, : ser me hie hlenbergiae, 112; Olyrae, I15, 411; pallida, 111, 412; pas aspalicola, , 122, 411; Stevensiana, 411 Urgine aritima, Uromyces Aristidae, 109; cdg 120; osus, ah aocmapnis 20. Urtica gracilis, Utricularia ae 302; vulgaris, 302 ) Vaccaria ecgicvelon 425 ium Beira Bae develop- m, 308; viscinum, 431 ee oe 308; lilacina, 308; stellata, 305; trifoliata, 30 Pacer pels phylla, 318; Scouleri, 319; septentrionalis, 303; sitchensis, 320 Veratrum speciosum, 3 Verbena bracteosa, 47; "venosa, 186 bear montanofolia, gina. 245; alti ssima » 248; brevi : i 501 240; salamana, 242; shore 337, 239; Shaferi, 238; = uerc cee Veronica ameri pero ‘ick i: ce bahar sein 300 ; Wormabjoli 30 Vi va) ame i ee oregana, 310; sparsiflora, 310; trifida, 306 Viola a ,» 301; Beckwithii, 321; canadensis, 306; laetecaerule ae: lin folia, 3113 acloskeyi, 316; nephrophylla, 302; Nuttallii, 46; or- biculata, 319; palustris, 306; pedata, 430; datifda, 306; renifolia, 306; rugulosa, 308; Selkirkii, ae n- trionalis, 302; ee 321; vallicola, 313; venosa, 31 Ghee ae celebicum, 268 otes on trees and shrubs setae and s the basis osperm a: Mot stry of maize—a reply to critici 215 Weisia Ber rteri anag 217 4; — denudata, 168 ane , R. S., The genus Desmatodon fo) America, a Notes on cai peepee Lichens, 2 Woopsurn, W. L., Pre he tiary notes the embryology of Reboulia he sphaerica Woods, Sorted or tier-like structure of certain dicotyledonous, oods, Prepara and treatment of, ooh Woodsia glabella, 305; oregana, 311; scopulina, 311 ; | Wyethia amplexicaulis, 311 i atc Lg anthi mune, 436 pms tee tial "Douglas 319; tenax, 316 Xerophytic asslands at different altitudes in Cceaaites 37 Xylophacos Parryi, 46; Purshii, 46; Shortianus, 46 Xy losma hawaiiensis, 148 as Youngia nana, 303, 304 cicula i 247; pon St Yueca glauca, 44, 49 folia, 237; iifolia pla 238; utivena, 244; icosantha, 238;| Zea Mays, The effect of soaking in interior, 247; illinoensis, 244; j water and of Pcie on the growth ; lepidota, 250; longifolia, 2393 of, or michiganensis, 245; Milleri, 251; mis- Zea Mays, gametogenesis and fecun- surica, 244; uri austr dation in, 7 2453 241; mollis, 243;| Zygnema affine, ; ‘ ovalifo : chaly um, 444; Collinsianum a | | | INDEX . a = ; + , , aren aequale, 445; Agana. 4453 _leios Lestesarinpe, 445; ericetorum, “aasi 445; purpur oS ae gracile, i : : ect 4453, steiinum, 445, 446; ie ee Vaucherii, 44 , 3 | | VOLUME 46, PLATE I9 BULL. TORREY CLUB EMBRYOLOGY OF REBOULIA WOODBURN: