Z aa) pe A cá el O a din JT x EE x O A z = OQ an un >- = . | e Y pa o 0L HR A — A , —— © Annals of the Missouri Botanical Garden Volume XXII 1935 With Twenty-one Plates and Twenty-eight Figures Published quarterly at Fulton, Missouri, by the Board of Trustees of the Missouri Botanical Garden, St. Louis, Mo. . Entered as second-class matter at the post-office at Fulton, Missouri, under the Act of March 3, 1879. Annals of the Missouri Botanical Garden A Quarterly Journal containing Scientific Contributions from the Missouri Botanical Garden and the Graduate Laboratory of the Henry Shaw Sehool of Botany of Washington University in affiliation with the Missouri Botanical Garden. Information The ANNALS Or THE MISSOURI BOTANICAL GARDEN appears four times during the calendar year: February, April, September, and November. Four numbers constitute a volume, Subscription Priee........ $6.00 per volume Single Numbers.......... 1.50 each * Contents of previous issues of the ANNALS OF THE MISSOURI BOTANICAL GARDEN are listed in the Agrieultural Index, published by the H. W. Wilson Company. * Except No. 3 of Vol. 22, containing 384 pages, price $5.00, - ue Ar. A e NM a STAFF OF THE MISSOURI BOTANICAL GARDEN Director, GEORGE T. MOORE. Assistant to the Director, KATHERINE H. LEIGH. EDGAR ANDERSON, HERMANN VON SCHRENK, athologist. Geneticist JESSE M. GREENMAN, ROBERT E. Woopson, : Be, Research Assista Curator of the Herbarium. Davin C. FAIRBURN, ERNEST S. REYNOLDS, Physiologist. Research Assistant. CARROLL M DODGE, NELL C. HORNER, Librarian ‚and Editor Mycologist. of Publications. BOARD OF TRUSTEES OF THE MISSOURI BOTANICAL GARDEN President, GEORGE C. HITCHCOCK. Vice-President, DANIEL K. CATLIN Second Vice-President, ALBERT T. PERKINS. L. Ray CARTER. GEORGE T. MOORE. SAMUEL C. Davis. THOMAS S. MAFFITT. FRED G. ZEIBIG. EX-OFFICIO MEMBERS: GEORGE R. THRO BERNARD F. DICKMANN, Mir ctore of Washington Mayor of the City of St. Louis. ROBERT J. TERR WILLIAM SCARLETT, Bishop of the Diocese of Missouri. JAMES J. FITZGERALD, President of the Board of Education of St. Louis. ence of St. Lo GERALD E. ULRICI, Secretary. President of o o dd of Sci- TABLE OF CONTENTS Observations on the Inflorescence of Apocy- naceae (With Special Reference to the American Genera of Echitoideae)....... Robert E. Woodson, Jr. A New Kallstroemia from Texas....Louis Williams $9 9 9 9 c9 9 9 9? * *9 c9 9 * * * * * € e$ * * * * A Critical Study of Certain Epappose Gen- era of the Heliantheae-Verbesininae of the Natural Family Compositae. . Ward M. Sharp Studies in the Apocynaceae. IV. The Amer- ican Genera of Echitoideae............. Robert E. Woodson, Jr. $9 9 9 9 9 9 9 9 c9 9 5 $9 c9: 9 * $ c9 o£ $ $9 i£ t$ 5 Cytology of Geotrichum versiforme Moore.. Morris Moore o9 9 9 9 c9 . €$ 9 € 9 9 c$» c 9 c3 * €» *$ c* c5 5 $ c9 f 9 * 3» $ *$ e. Head Infection Caused by a New Hemispora: H. coremiforms ......... 224 2B Morris Moore A. Morphologieal and Physiologieal Study of Two Species of Posadasia, P. capsulata (Darling) Moore and P. pyriformis Moore Morris Moore € 9 € à s « 6 6€ & 84/9 6 A 5 » 9 A4 € R An Improved Apparatus for the Determina- tion of Carbon Dioxide Production in Phys- iological Plant Studies............ F. Lyle Wynd New Species of Hydnangiaceae............ Sanford M. Zeller and Carroll W. Dodge An Annotated Catalogue of the Flowering Plants of Missouri. ES Se... PAGE 1- 48 49- 50 01-152 153-306 307-316 317-334 339-360 361-363 365-373 375-158 The Reaetions of Plants to Ultra-Violet.... datar ES. E C Rat Xr EE. De Reynolds Studies in Ultra-Violet and Respiratory Phenomena. 1. Review of Work Pub- lished before June, 1935................ Pee O ee ee F. Lyle Wynd and E. S. Reynolds Studies in Ultra-Violet and Respiratory Phenomena. II. The Effects of Ultra- Violet on Respiration and Respiratory Einzymes of Higher Plants.............. F. Lyle Wynd, Harry J. Fuller, and E. S. Reynolds Studies in Ultra-Violet and Respiratory Phenomena. III. The Influence of Vari- ous Regions of the Spectrum on the An- aerobic Fermentation of Yeast.......... NOPE err ee Te E. S. Reynolds and F. Lyle Wynd A Note on the Application of the Glass Elec- trode to the Determination of Oxidation- Reduction Potentials............. F. Lyle Wynd General Index to Volume XXIT................... 759-769 111—835 831—852 853-860 861-865 867-874 Annals of the Missouri Botanical Garden Vol. 22 FEBRUARY, 1935 No.1 OBSERVATIONS ON THE INFLORESCENCE OF APOCYNACEAE (WITH SPECIAL REFERENCE TO THE AMERICAN GENERA or ECHITOIDEAE) ROBERT E. WOODSON, JR. Research Assistant, Missouri Botanical Garden Instructor in Botany, Henry Shaw School of Botany of Washington University INTRODUCTION As one reviews the advances of plant morphology during the past quarter century and more, one can scarcely avoid recogni- tion of the fact that systematic botanists have made few recent contributions of note to our practical and theoretical knowl- edge of the structure of plants. This is in spite of the fact that the whole superstructure of plant classification rests upon an intimate knowledge of such considerations. The situation is ineongruous with the superior opportunities which systema- tists enjoy for the advancement of the very basic and elemental science of plant morphology. In the monographie method of investigation, the systematist has developed a tool of the very greatest potentiality for the exhaustive investigation of the morphology of plants. The ac- cumulating multitudes of specimens in the great herbaria of the world which are available for study, particularly by the trained systematist, constitute an unsurpassed field for ob- servations concerning the known flora in its entirety with the greatest ease. The important investigations of Solereder and ANN. Mo. Bor. GARD., VoL. 22, 1935, (1) [Vor. 22 2 ANNALS OF THE MISSOURI BOTANICAL GARDEN others upon the anatomy of flowering plants toward the close of the past century should remind us that herbarium specimens are capable of more uses in botanical research than ordinary dissection for the purpose of identification. The possibility of studying the inflorescence of Apocynaceae largely by means of herbarium specimens was first appreciated by the present writer while studying in the Royal Botanie Gar- dens and herbarium at Kew during the summer months of 1930. The accurate picture of inflorescence structure which such specimens are capable of revealing was manifest at that time upon comparison of the exsiccatae with the abundant flowering material of Allamanda Schotti Pohl found in the conserva- tories containing Victoria regia and other tropical species. The gardens at Kew are perhaps the richest in cultivated spe- cies of Apocynaceae among the botanical gardens of the world, and the preliminary comparison of desiccated and living speci- mens of numerous genera of the family found there, made pos- sible through the courtesy of the Director, Sir Arthur W. Hill, and others in authority, has proved of the utmost value in the subsequent studies of the inflorescence of which this paper is a condensed report. Since the commencement of the problem at Kew, the writer has continued the study of the apocynaceous inflorescence in the herbarium and living collections of the Mis- souri Botanical Garden at St. Louis. The taxonomy of the Apocynaceae has claimed my attention for nearly a decade during which I have examined thousands of specimens of this family deposited in the principal herbaria and botanical gardens of America and Europe. The Apocy- naceae are divisible into three subfamilies, the Apocynoideae, the Echitoideae, and the Plumeroideae. The scope of my taxo- nomic studies has included all the recognized genera and spe- cies of Apocynoideae and Echitoideae indigenous to the west- ern hemisphere, as well as certain related genera of Europe and Asia (Woodson, 1930, 1933). Earlier studies have also in- cluded certain genera of Plumeroideae (Woodson, 1928a, b). Structural advantages of the flowers of Apocynaceae which overcome the disadvantages suffered by desiccation to an ap- preciable extent in investigating the gross morphology of the 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 2 inflorescence can be indicated at this time. The flowers of the family as a whole are large, permitting even relatively small discrepancies of age to be visible in the development and size of the buds. The rule of large flowers is broken in such genera as Forsteronia and Apocynum, but even in such instances dif- ference in stages of development of the floral buds is clearly in- dicative of correlated age and potential time of blooming, very important factors in the interpretation of modes of inflores- cence. Relative age of floral members is also manifest in young fruit. A less obvious criterion in this connection is found in the accrescence of the pedicels. Since the pedicels increase in length and diameter for some time preceding abscission, even in the ease of flowers which have failed of fertilization, a third important clue is present in herbarium, as well as in living, specimens to enable us to determine the phenology of the inflo- rescence with some precision. The range of inflorescence structure exhibited throughout the family Apocynaceae comprises nearly all the more familiar types found among the flowering plants with the exception of such specialized examples as the capitulum and the ament. The most common inflorescence is the cyme, particularly the dicha- sium. At this juncture it becomes necessary to outline briefly the characteristic types of inflorescence amongst the Apocy- naceae, both as a hasty survey and as a definitive introduction to the succeeding discussion. It will be found that the defini- tions of the inflorescence follow closely those of Jackson (1900). Dichasium.—The dichasium is the most familiar type of cymose inflorescence throughout the flowering plants. It con- sists of a solitary flower terminating the primary axis of the stem or branch. Subtending this flower are normally two op- posite leaves or bracts in the axils of which arise two secondary branches, each terminated by a flower similar to the first, but blooming subsequently and usually simultaneously. Such an axis system is called a simple dichasium. A continuous, or compound dichasium is frequently observed in the branching of the secondary axes in a manner similar to that of the pri- mary, and soon. Such subsidiary components of a compound cyme are designated by the diminutive cymule. Dichasia are [Vor. 22 4 ANNALS OF THE MISSOURI BOTANICAL GARDEN frequently so composed of innumerable flowers arranged in im- pressive regularity. This type of inflorescence is found in numerous genera of Apocynaceae, including Cycladema Benth., Rhabdadenia Muell.-Arg., and Echites P. Br. Aggregate Dichasium.— A “panicle”” is strictly defined as a compound inflorescence all axes of which are indeterminate. This interpretation was original with Roeper (1826), to whom we owe the first thorough-going discussion of the inflorescence. Nevertheless the general use of the term, persisting since the time of Linnaeus, still permits its use for virtually any com- pound inflorescence, preferably pyramidal and ‘‘loose’’ (Jack- son, 1900; Gray, 1907). The exact opposite of paniculate construction, one in which all axes of a compound inflorescence are determinate, has ap- parently never received a generally accepted name, although it, too, frequently is made to masquerade as a ““panicle.”? This type of compound inflorescence differs from the typical dicha- sium in that although the latter is the ultimate constituent of its construction, more than a single pair of branches occur, ordinarily at regular, decussate intervals below the determi- nate flower of the primary axis (ef. text-fig. 3, A). For want of a better name it is called an aggregate dichasium, since the inflorescence itself consists properly not of a single dichasium but of an aggregation, just as a raceme consists not of a single flower, but of an indefinite collection of them upon a specialized shoot. This type of inflorescence is conspicuous among A pocy- naceae, as well as in certain other families, as Gentianaceae, being well organized in such familiar genera as Apocynum L., and Forsteroma G. F. W. Meyer,’ accounts of which follow shortly. Cincinnus and Bostryx.—In those eymes which consist of a single, sympodial axis, the monochasia of Eichler (1875), de- rivation from a dichasium has long been assumed by the substi- tution of a single flower for one of the equivalent inflorescence 1 The limitations of publication preclude a complete category of the inflorescence structures of the genera of Apocynaceae which number at the present time more than two hundred. 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 5 branches. A bostryx is formed when the remaining subsidiary axis of such a modified dichasium is always either to right or to left of the determinate flower. This usually results in the somewhat curled inflorescence known as ‘‘helicoid,’’ familiarly represented in many Boraginaceae. The maintenance of branching of such a highly modified dichasium alternately to right and to left, resulting in a somewhat zigzag sympodium widely known as ‘‘scorpioid”’ is characteristic of the cincinnus. In the bostryx and the cincinnus the sympodium is usually more or less curled or zigzag respectively. They may be simple throughout, or variously branched, by which they may recall their dichasial ancestry to a certain extent. Not infrequently the axis may become quite straight, however, when the nature of the sympodium must be distinguished by the usually paired pedicels in the axil of a single bract which is opposed to an ‘‘empty’’ bract upon the other side of the peduncle (ef. pl. 3, fig. 3). The peduncle is literally in the axil of the ““empty”” bract, which frequently becomes greatly dislocated in highly modified scorpioid and helicoid cymes, rendering their inter- pretation less easy. The bostryx and the cincinnus occur widely throughout the Apocynaceae, the former in such genera as Odontadenia Benth., Allamanda L., and Asketanthera Woods., and the latter in Prestonia R. Br., Temnadenia Miers, and Mesechites Muell.- Arg. Scorpioid and helieoid modifications frequently creep into the ramifications of other eymose types, particularly when multiflorous. Raceme.—Scarcely distinguishable from some highly modi- fied examples of superficially indeterminate scorpioid and heli- coid cymes is the truly indeterminate raceme. A typical ra- ceme may be defined as a simple, pluriflorous, monopodial in- florescence in which a solitary pedicel is borne in the axil of a single braet. A pronounced peduncle is usually present which is of relatively greater length than the individual pedicels. Familiar modifications of the typical raceme are the spike, the corymb, and the umbel, based upon the relative length, or es- sential suppression of peduncle and pedicels. Typical ra- cemes are infrequent in Apocynaceae, the principal genus bear- [VoL. 22 6 ANNALS OF THE MISSOURI BOTANICAL GARDEN ing them being Mandevilla Lindl., of some 108 species of the western hemisphere. Jackson (1900) defines the umbel as ‘‘properly indetermi- nate," but recognizes the use of such a term as ‘‘cymose umbel”” for those which are centrifugal and not centripetal in their floral development. Similarly, the same authority defines both a spike and a corymb as indeterminate. In these cases also it appears inevitable to recognize determinate as well as indeterminate spikes and corymbs. In such species as Fors- teroma paludosa Woods., it will further be necessary to deal with a spiciform thyrse: a typical thyrse in which the pedicels are suppressed. Corymbs, in their typical, indeterminate sense, are restricted to racemose genera, as in Mandevilla cal- lacatensis Mgf. The true umbel in Apocynaceae is unknown to this writer. Cymose, or determinate, corymbs and umbels are widely observed in such genera as Prestonia R.Br., The- nardia HBK., Malouetia A.DC., and Asketanthera Woods. The spike, the umbel, the corymb, and other similar types, are doubtless merely modes of more fundamental types of in- florescence, and it would appear more simple in the end to use them in an adjectival manner, such as ‘‘corymbose raceme,’’ and ‘‘corymbose eyme," than to limit the use of the word ““corymb”” to an indeterminate structure, in which case there would be an obvious necessity for another word to express similar, but determinate construction. Thyrse and Panicle.—The thyrse and the paniele are recog- nizable as compound inflorescences in which the primary axis is indeterminate. In the true panicle, as represented by the com- mon lilac, Syringa vulgaris L., the subsidiary axes, whatever their number, are likewise indeterminate. In the thyrse, how- ever, the ultimate ramifications of the inflorescence are deter- minate, hence the **mixed infloreseence’’ of A. P. De Candolle (1827). The modification of the thyrse is excellently shown in the apocynaceous genus Forsteronia, amongst others, in which the determinate construction is variously expressed upon branches of differing magnitude throughout the entire inflores- cence, The true panicle apparently is not found in any known Apocynaceae, although it is approached in certain species of Se luo o ALME. M sie M IM Y 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 7 Forsteronia, a detailed account of which is reserved for suc- ceeding paragraphs. Solitary Flowers.—Solitary flowers are found in both termi- nal and lateral positions in the Apocynaceae. The former is typical of such genera as Salpinctes Woods., and such excep- tional species of predominantly pluriflorous genera as Echites crassipes A. Rich. In all known instances, however, very in- conspicuous, subtending bracts are usually capable of detec- tion. In the familiar creeping myrtle, Vinca minor L., the soli- tary, lateral flowers are without the slightest evidence of sub- tending bracts. The different condition of the closely related periwinkle, Lochnera rosea (L.) Rehb., will be discussed presently. Closely correlated with the predominance of eymose inflores- cence in the family Apocynaceae is the occurrence with but few exceptions of decussate phyllotaxy. So much has been written concerning the spiral arrangement of appendages of the shoot as ‘‘primitive,’’ a term of all too vague implications in many instances, that observations to the contrary might appear to have been suppressed by the sheer weight of consensus of opinion among morphologists. In the case of Apocynaceae, however, the reverse appears obvious. Among the 132 genera recognized within the family by Schu- mann (1895) only twelve are characterized by spiral phyl- lotaxy. Of these, but one and two respectively are contained within the subfamilies Apocynoideae and Echitoideae, which have been found to have an elementary, or ‘‘primitive,’’ carpo- logical structure predominating (Woodson, 1930), while the alternate-leaved genera of Plumeroideae, characterized by the occurrence of generally more complicated, or ““advanced,”” carpologieal conditions, number nine. Whorled phyllotaxy is found in fourteen genera of Plumeroideae and four of the Echitoideae enumerated by Schumann. In this connection it appears significant that of the eighteen genera with whorled phyllotaxy, no less than seven! are known to contain at least one species with normal, decussate foliage. In this, as in other 1 Aspidosperma, Condylocarpon, Cowma, Gynopogon, Laubertia, Macrosiphonia, Pycnobothrya. [Vor. 22 8 ANNALS OF THE MISSOURI BOTANICAL GARDEN families, there is an obvious correlation of whorled and oppo- site phyllotaxy indicating the derivation of the former from the latter, possibly by the aggregation of nodes. Such deductions as the immediately preceding receive addi- tional support by observations on the young seedlings of Am- soma spp.' and Rhazya orientalis (Dene.) A. DC., the first stem-leaves of which are characteristically decussate. Two alternatives effect the adult, spiral phyllotaxy. In numerous instances among the seedlings under observation lateral buds were produced in the axils of the cotyledons. Upon subsequent development the arrangement of leaves upon such lateral branches was found in all cases to be normal for the adult con- dition of the species. The plumular shoot was short-lived. Somewhat less frequently among the seedlings a gradual change from decussate to spiral phyllotaxy was seen to occur upon the plumular shoot by the vertical translocation of nodes. The juvenile shoot was thereupon transformed directly to the adult condition. In innumerable instances more or less con- spicuous vertical translocation of individual foliar nodes may be observed in species of Apocynaceae with normally opposite phyllotaxy. Upon physiological-anatomical grounds opposite phyllotaxy and the occurrence of perfectly cymose inflorescence would ap- pear to be unavoidably correlated. To many botanists the chief, if not the only, important characteristic of the cyme 1s the fact that for each axis a solitary, terminal flower exists which may be associated with a variable number of subsidiary floral members developing in basipetal succession. A most essential feature of the perfectly compounded eyme which has been overlooked generally, even by some students of inflores- cence structure, is the almost geometrical precision which gov- erns the expansion of its floral members. In such a simple di- chasium of three flowers as has been described in a preceding paragraph, observable in innumerable genera of flowering plants in such families as Caryophyllaceae, Primulaceae, Gen- tianaceae, Apocynaceae, etc., the primary floral member termi- 1 A. Tabernaemontana Walt., A. brevifolia A. Gray, A. illustris Woods., A. pogon- osepala Woods, 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 9 nating the vegetative axis of stem or branch invariably blooms first. This is soon followed by the nearly simultaneous expan- sion of the two equivalent, lateral floral members immediately subtending. If the dichasium be compounded to include for either branch a pair of lateral floral members subtending the secondary de- terminate pedicels, these will bloom shortly thereafter, in almost clock-like concert. This very impressive similarity of composition and unison of development of equivalent members of ““perfect”” cymes may appropriately be referred to collec- tively as its characteristic, or elemental ‘‘symmetry.’’! Com- mon observation will probably lead one to object at this june- ture that many, if not most, eymose inflorescences are not pos- sessed of one or both of these ““ideal”” attributes under actual living conditions. Such is doubtless the case, and the re- mainder of this paper will be devoted to the occurrence of such irregularities among certain Apocynaceae and their possible evolutionary significance. It is a fact quite easily observed that truly ‘‘perfect’’ cymes, whether di- or pleiochasia or derivatives of these, are found only upon flowering plants with opposite or whorled phyllo- taxy. In no alternate-leaved species has this writer observed the strikingly symmetrical development of equivalent floral members so characteristic of the ““perfect”” (e. g. ‘‘ideal’’) cymes. The reason is doubtless anatomical and physiological: an inflorescence borne at the lower of two translocated nodes receives lts vascular supply from the apical meristem some- what earlier than the inflorescence immediately above. Hence subsequent development of the floral members is slightly in ad- vance of that of the cyme immediately above, in spite of the fact that the two would be **equivalent"' in the event of opposite phyllotaxy. Although observations have been made over an unusually wide assortment of specimens during a number of years, as has been indicated in a preceding paragraph, the chief trends of 1 This theory of the elementary symmetry of the ideal cyme will be found to place more emphasis upon the faetor of time in development than the somewhat similar conception of ‘‘rhythm and symmetry? propounded by Goebel (1931). [Vor. 22 10 ANNALS OF THE MISSOURI BOTANICAL GARDEN the evolutionary development of the inflorescence in Apocy- naceae may apparently be visualized from a review of a few characteristic representatives. In spite of the inevitable loss of minute detail, such an abbreviated account should possess a degree of clarity and simplieity diffieult of attainment in a more extended, if more comprehensive account. It has been a matter of considerable disappointment that anatomical obser- vations have been impossible because of the difficulty in secur- ing fresh material of the critical genera. It is hoped that such data may eventually be secured to test the wholly gross mor- phological account which follows. A final word of caution may be necessary to emphasize that perhaps no phase of the flowering plant’s gross morphology is more variable than the inflorescence. The inflorescence is es- sentially a phase of branching of the shoot system, but it is vastly more complicated than vegetative branching, from which it has been derived. Hence the examples of inflorescence used to illustrate the observations which follow must be under- stood to represent tendencies perceived after wide study: they represent by no means the whole range of variation of a genus or species. Explanation of Symbols Used in Diagrams.—In the diagrams illustrative of the sueceeding paragraphs, dimensions have been maintained constant, despite disparities of the inflores- cences in nature. The following symbols have been employed to denote floral members: O — floral bud. " — flower at anthesis or shortly thereafter. — fruit (in this and the preceding the relative size Y of symbols is indieative of the corresponding de- gree of development upon the actual specimen observed). ) — bract. ^ — leaf. 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 11 N — undeveloped bud, floral or vegetative. / — broken portion of specimen. A — whole mature inflorescence. — immature inflorescence. The limitations of diagrammatic representation render im- practical an attempt to indicate the decussate or spiral compo- sition of the inflorescence. Branches, pedicels, and bracts which appear opposite and in the same plane must be inter- preted as naturally decussate: similarly with the alternate members appearing as in the same plane which must be con- strued as naturally spiral. OBSERVATIONS Ecnurrzs P. Br. Our present knowledge of the genus Echites ( sensu stricto) is limited to seven species and a putative variety which may be separated provisionally into four sectional groups centering about E. tuxtlensis Standl., E. turbinata Woods., E. yucata- nensis Millsp., and E. umbellata Jacq., respectively. The aspect of the inflorescence of these species is reproduced in pls. 1-2. The best-known species of Echites, as well as the type of the genus, is E. umbellata Jacq., a liana occurring virtually throughout the Greater Antilles and extending to southern peninsular Florida, the Bahama Islands, and the coastal flats of northern Yucatan and British Honduras. The inflorescence of this species consists of one to several flowers, two, three, or four being the predominant numbers, borne in corymbose cymes. Tendencies of the inflorescence may be discussed briefly with the aid of five sketches provided in pl. 1. The cymes of E. umbellata are predominantly alternate-lat- eral, but may be pseudoterminal as well. Such an individual is shown in pl. 1, fig. 5. In this case the axis is terminated by a [Vor. 22 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN flower in full bloom. The two lateral branches bear one and two buds respectively, the solitary bud to the left being con- spicuously larger than either of the two upon the opposite branch. The paired bracts of the branch to the right, in the axil of one of which the smaller lateral bud is found, indicate that the present condition was probably derived from a perfect cymule of three floral buds. Detection of a minute, abortive bud in the axil of such a superficially ‘‘empty’’ bract is usually possible upon eareful dissection. At the base of solitary, lat- eral pedicels as that to the left of the figure are usually found a few rather indefinite, tiny bracts, also indicating that the uni- florous eondition has probably been derived from a lateral cymule. In fig. 2 of the same plate an inflorescence is shown in which the right lateral branch of the dichasium has failed to develop in the axil of its subtending bract which remains. The terminal flower is full blown. "The left lateral branch consists of three well-developed, and one abortive, buds. The terminal bud of this eymule is about to expand, and is the largest of the three. The lateral bud to the left of the terminal is manifestly next in size, and is undoubtedly interpretable as the remnant of a re- duced lateral eymule. The terminal bud of the right eymule is the smallest of the functional buds, and this fact appears to be correlated with the presence of the abortive, lateral bud at its base. Evidently the right cymule is slower to develop than the left because its axis has not been stripped of subsidiary axes dependent upon its vascular supply. At any rate, an examina- tion of many specimens demonstrates an indubitable associa- tion of advanced development and potential blooming with in- creased reduction of the inflorescence. Plate 1, fig. 1 illustrates a dichasium of three members, the terminal pedicel of which is bearing young fruit. The two lat- eral flowers are of approximately equal development. Two ex- planations of this eyme are possible: that the inflorescence as such is basic and elemental, or that the two lateral members have been reduced from pluriflorous, lateral eymules to the same degree, and thus expand with approximate simultaneity. Figure 4 represents a uniflorous condition of the inflorescence 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 13 of E. umbellata, not uncommonly found, in which, however, two abortive buds subtend. The truly uniflorous condition, without the presence of one or more abortive buds, has not yet been observed in this species. In E. crassipes A. Rich., however, solitary flowers are borne, and with the aid of somewhat smaller foliage serve to distin- guish this poorly understood endemic of Cuba. Even in the in- stance of this species, nevertheless, an indefinite and small number of minute bracts at the very base of the pedicel gives strong support to the assumption of reduction from a pluri- florous dichasium (pl. 1, fig. 3). The scope of variation in the relatively small inflorescence of E. wmbellata increases with the number of specimens examined, and is plainly impossible to describe adequately at this time. An examination of even a few, however, is sufficient to form the opinion that the inflores- cence of the species as a whole is undergoing reduction, and that the progressive reduction of a eymule tends to hasten the development of the remaining flower buds in relation to those of less reduced but equivalent eymules. Plate 2 represents the inflorescence of four additional spe- cies of Echites. The group centering about E. yucatanensis is confined to Central America, and is most tangibly separated from the Antillean E. umbellata by the absence of the spiral contortion of the corolla-tube of the latter. Although the di- chasium has advanced decidedly toward the umbellate condi- tion, the general tendencies of the floral development of the for- mer (pl. 2, fig. 3) are conspicuously similar to those of the latter species. In E. turrigera, known only from Guatemala, the in- florescence (pl. 2, fig. 4) is an almost typical, pluriflorous di- chasium. Here also, however, the influence of unequal reduc- tion of equivalent cymules in the disruption of the eymose symmetry is in unmistakable evidence. Conspicuously smaller flowers set apart E. tuxtlensis and E. turbinata from the other species of Echites. Here also the inflorescence is noticeably more extensive. In the former (pl. 2, fig. 1) a cyme is found in which the primary branching is di- chasial and the subsequent composition helicoid. From the previous observations upon E. umbellata one may be led to [Vor. 22 14 ANNALS OF THE MISSOURI BOTANICAL GARDEN infer that here, and perhaps in all such helicoid cymes, the single, outer flowers of the sympodium, blooming only slightly subsequent to the expansion of the median, or terminal flower, have been hastened in their relative development by their pre- vious reduction from a pluriflorous cymule. The inflorescence of E. turbinata (pl. 2, fig. 2) is of puzzling nature, the first branching being typically dichasial, but the ultimate branching apparently partaking of the nature of the aggregate dichasium, since each of the three members is sub- tended by one or two small, probably abortive, floral buds, or by paired braets. A logieal explanation is that in this species a simple dichasium is being derived from the extensive modifi- cation of a more complex system. As this species, which is an endemie of Costa Rica, is extremely rare in herbaria, it may be hoped that a fuller understanding of its inflorescence will re- sult from increased collections. APOCYNUM L. Promising clues to the origin of the aggregate dichasium are found in Apocynum, a genus of few species remarkably wide- spread over temperate North America. In A. cannabinum L., a ubiquitous weed of fields and roadsides, the inflorescence is predominantly terminal and multiflorous, consisting of a much ramified aggregate dichasium (text-fig. 1, A). Accompanying the terminal inflorescence are always two subsidiary, lateral shoots, both of which may bear a varying number of foliar nodes, in which case both eventually terminate with an inflo- rescence similar to the first. In many instances only one of such branches produces foliage; in this case the other produces directly an inflorescence similar to the first, but usually some- what smaller. Naturally this second lateral inflorescence comes to full development much sooner than the first because of the elimination of the intervening foliar nodes. In many in- stances both lateral shoots may produce inflorescences directly without the produetion of foliage, in which ease they become an integral part of the terminal inflorescence. Apocynum cannabinum is an obnoxious weed largely because of its size and long period of bloom. These attributes are the 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 15 result of its ability to produce inflorescences through a rela- tively long season by means of its dichotomous or cymose branching. A much less pestiferous weed is A. androsaemi- q— ESI F Fig. 1. Inflorescence structure: A—Apocynum cannabinum es . androsaemifolium L.; C—Forsteronia leptocarpa (Hook. & Arn.) A.DC.; D—F. paludosa Woods.; E—F. spicata (Jaeq.) G. F. W. Meyer; F—F. amblybasis Blake; G—F. ela- chista Blake. Explanation in the text. CA folium (text-fig. 1, B), in which the lateral shoots accompany- ing the strictly terminal or median inflorescence do not tend to elongate as in the sister species. The blooming period, as well as the size of this species, is consequently relatively less. [Vor. 22 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN t Aggregate dichasium"' is evidently a well-chosen name for the inflorescence of Apocynum, from the phylogenetie stand- point; for if we read the data properly, it has been derived by the aggregation of a number of terminal diehasia through the elimination of the foliar nodes subtending them, hence the de- eussate composition of its branches. Recapitulatory evidence appears to occur in the inflorescence of A. androsaemifolium and other species, in which braets of the median axis of the ag- gregate dichasium occasionally regain to a limited extent their foliar charaeter. Apocynum is apparently the only genus of Apocynaceae in which the origin of the aggregate dichasium may be read so easily. In the closely related genus Trachomitum Woods. of Eurasia, the inflorescence has become about as highly organ- ized as in Forsteronia, an account of which follows. In Poacy- num Baill., a close Asiatic relative of Apocynum and Trachom- itum, the branches of the aggregate dichasium have undergone scorpioid modification. ODONTADENIA Benth. The evolution of the inflorescence of the echitoid genus Odontadenia would appear to approach the cincinnus from both the typical and the aggregate dichasium. Significant stages in this progression, drawn from representative species, are il- lustrated in text-fig. 2. The inflorescence of O. Hoffmannseggiana (Steud.) Woods. is a variously modified, aggregate dichasium bearing few to relatively numerous flowers (text-fig. 2, A). In the specimen illustrated, which was collected by Tessmann in eastern Peru, two flowers are full-blown, one terminating the median axis, and one terminating the subsidiary axis to the right of the fig- ure. It is clear that the floral bud terminating the equivalent, secondary axis to the left has failed to keep pace with the devel- opment of the sister determinate flower, and this fact is corre- lated with the fact that the eymule is composed of four flower- ing nodes and the one to the right of but two. It is significant also that the tertiary axes of the right cymule are in different stages of relative development: that to the left, consisting of 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE E but one floral bud, is in a conspicuously more advanced stage than either of the three comprising that to the right. The same situation is found with respect to the two subsidiary eymules of N ys Fig. 2. Inflorescence structure: A- Ur nta- denia Hoff mannseg giana ‘= teud.) Woods. —0. lawifiora (Rusby) Woods.; E—O. "lines ae ) W Rmo tai A UR (A.DC.) Miers. Ex- planation in the the median axis, each consisting of four members, with abor- tion plainly in progress in the case of one and three of either eymule to the left and right respectively. The physiological result is clearly evident in the relatively advanced stage of de- velopment of the lone functional bud to the right. [Vor. 22 18 ANNALS OF THE MISSOURI BOTANICAL GARDEN The first stages of the development of a cincinnus are also observable in O. Hoffmannseggiana. In text-fig. 2, C, an in- stance is found in which the inflorescence consists of a primary, fruiting pedicel and two strongly unequal, secondary branches of one and six functional pedicels to right and left respectively. Although it happens that young fruit terminates both second- ary axes of the dichasium, the size of the symbols is expressive of a significant disparity in relative development exactly like that found under similar circumstances in floral members. An additional factor of great importance in the modification of the inflorescence is found in the effect of the vertical trans- location of nodes. The translocation of the equivalent, tertiary eymules to the left of the inflorescence illustrated in text-fig. 2, B, shows that in such cases the members of the lower node are hastened appreciably in their development, even though those of the upper node may be somewhat more reduced in numbers. This fact, again, requires but an obvious anatomical- physiological explanation, since the lower node, differentiated first from the apical meristem, is effectively joined to the fune- tioning conductive tissues of the shoot and root at a somewhat earlier time than the upper. Hastened development of its sub- tending members follows. In O. laxiflora (Rusby) Woods. a peculiar modification of the inflorescence is found in which the main axis is regularly di- chasial, but the secondary branch to the left has become helicoid by the repeated reduction of the right branch of each eymule to a solitary floral bud (text-fig. 2, D). The secondary branch to the right is compounded dichasially to the tertiary axes, which then become helicoid. The terminal flower of the secondary branch to the left naturally blooms before that of the branch to the right because of the greater degree of reduction of its axis. The modification of the dichasium to a simple cincinnus is culminated in O. polyneura (Urb.) Woods. (text-fig. 2, E), in which a superficially indeterminate axis is compounded by paired pedicels borne in alternate succession subtended by one of a pair of bracts. The flowering axis is clearly a scorpioid sympodium, although it has lost the conspicuous zigzag aspect of less highly modified scorpioid inflorescences, as that of Angadenia Sagraei (A. DC.) Miers, illustrated in text-fig. 2, F. 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 19 ForsTERONIA G. F. W. Meyer The inflorescence of Forsteronia has been described rather loosely as a panicle. A thorough study of representative spe- cies discloses the fact that a true panicle is absent in all. Va- rious modifications of the thyrse are predominant, varying from typical, pyramidal outline to structures which superfi- cially resemble the corymb and the spike, through the elonga- tion or the virtual elimination respectively of the subsidiary axes. The recent discovery of a species with the structure of an almost typical, corymbose raceme heightens the significance of the inflorescence structure of this genus, and renders its study of great value in the evolutionary history of flowering axes generally. Text-fig. 3 provides diagrams of the inflores- cence of various representatives of this genus. The inflorescence of Forsteronia is composed of many flowers of relatively small size, rendering analysis and inter- pretation somewhat difficult. Relatively young inflorescences, frequently collected separately and preserved in small packets upon herbarium sheets, have been found extremely useful for study, since they may be boiled in toto, restoring a semblance of the living posture. After study such specimens may be par- tially dried and pressed between blotters, from whence they may be returned to the herbarium packets after complete desic- cation. Such procedure serves to emphasize the convenient use of herbarium specimens for the study of gross morphology be- yond the usual practice of taxonomic identification. The elemental type of inflorescence in Forsteronia would ap- pear to be the many-flowered, aggregate dichasium, such as that of F. corymbosa (Jacq.) G. F. W. Meyer, F. floribunda (Sw.) G. F. W. Meyer, and F. portoricensis Woods. In such an inflorescence (text-fig. 3, A ), the terminal flowers of the median, as well as those of all secondary axes, bloom with approximate unison. It is possible that this rather arresting phenomenon may be in large part due to the successive reduction of the num- ber of floral members borne upon the secondary axes from bottom to top, resulting in the pyramidal outline. Focusing our attention upon the individual composition of the lateral members, it is found that the tertiary branches are [VoL. 22 20 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ch d WP X ES i B EEL EEE Ps, y 9325: > Pa ENTER E EF Fig. 3. Inflorescence structure: A—Forsteronia corymbosa (Jaeq.) G. F. W. Meyer; B—F. Velloziana (A.DC.) Woods.; C—F. thyrsoidea (Vell.) Muell.-Arg.; D—F. leptocarpa (Hook. & Arn.) A.DC.; E—F. simulans Woods. Explanation in the text. 1935] WOODSON— INFLORESCENCE OF APOCYNACEAE 21 dichasial. The slight nodal translocation has apparently not sueceeded in influencing effectively the phenology of the inflorescence. The inflorescence of F. Velloziana (A. DC.) Woods. is a corymbose structure intermediate between the aggregate di- chasium and the thyrse (text-fig. 3, B). In this inflorescence the symmetrical expansion of the terminal flowers of the second- ary axes has been disrupted effectively by the reduction of cer- tain branches, resulting in hastened development of certain of the floral members. The primary axis, however, is still determinate. The first indications of the modification of the aggregate di- chasium to the thyrse seen in F. Velloziana lead to the inflores- cence type of F. thyrsoidea (Vell.) Muell.-Arg. (text-fig. 3, C), which is more or less typical of the thyrsiform structure, since the main axis is indeterminate with respect to the secondary axes, which still retain their dichasial composition variously modified. In this extensive inflorescence the small amount of variation in flower numbers among the secondary cymules from the first to the fifteenth pair of nodes can perhaps be ex- plained by the assumption of reduction in number of floral members in basipetal suecession. This assumption would also aid in explaining the acropetal development of the secondary cymules which is evident. Typical panicles are apparently never found in Apocyna- ceae. The paniculate structure is closely approximated in the inflorescence of F. leptocarpa (Hook. & Arn.) A. DC., in which both the main and the secondary axes are indeterminate, al- though the tertiary and lesser branches remain determinate (text-fig. 3, D). The factors influencing the indeterminate modification of the primary axis of F. thyrsoidea may safely be supposed to have been identical with those effecting the in- determinacy of the secondary axes of F. leptocarpa. Our knowledge of the evolution of the inflorescence in For- steronia is culminated in F. simulans Woods., a rare liana of Colombia, in which the inflorescence is superficially a corymbose raceme with simple lateral pedicels. From the diagram of this ‘‘raceme’’ provided in text-fig. 3, E, however, have been [Vor. 22 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN omitted numerous extremely inconspicuous bracts and abortive buds oceurring rather irregularly upon the peduncle amongst the floriferous pedicels, which further exhibit a marked ten- deney of transloeation. It appears obvious that this inflores- cence has been derived from a thyrse characteristic of neigh- boring species. One can scarcely fail to be impressed by the great range of modification of the inflorescence amongst species of For- steronia which would appear to offer a most significant clue toward the solution of the questioned relationship of determi- nate and indeterminate inflorescences. 'l'he remaining known species of Forsteronia, numbering forty-one, present various gradients of the inflorescence structures of the five examples discussed in the preceding paragraphs. The great bulk of these show certain tendencies centering about the types repre- sented by F. Velloziana, F. thyrsoidea, and F. leptocarpa. Itis significant that although the inflorescence has not been em- ployed to a great extent in the taxonomie separation of the species in a recent revision by this writer, additional morpho- logical criteria used in speciation have resulted in a concurrent classification. Species of Forsteronia show a progressive transformation of vegetative to floriferous axes, distinctly recalling that pre- viously discussed in Apocynum, but in greater variety. In F. leptocarpa (text-fig. 1, C), the terminal, aggregate dichasium is regularly accompanied by two lateral, vegetative shoots which continue the extension of the massive liana characterized by this species. Each of such vegetative branches may be com- pounded similarly. A somewhat similar method of branching is found in F. paludosa Woods., in which, however, a single lateral branch is developed in alternate axils of the decussate leaves. An abortive, or dormant bud is regularly found in the axil of the opposite node, which ean become stimulated to growth upon injury of the stem above its occurrence (text- fig. 1, D). In F. elachista Blake a lateral inflorescence occurs with the strictly terminal, oceupying the alternate axils of the foliar nodes as in the vegetative branches of F. paludosa (text-fig. 1935] j WOODSON— INFLORESCENCE OF APOCYNACEAE 23 1, G). In the case of numerous tropical Apocynaceae, notably species of Odontadema, an indefinite number of cataphylls are found occurring at the base of such lateral inflorescences. These are not observed subtending vegetative branches, or they are inconspicuous. A natural assumption which pre- sents itself is that the alternate lateral inflorescences of F. elachista are homologous with the vegetative branches of F. paludosa. In F. spicata (Jacq.) G. F.W. Meyer (text-fig. 1, E), terminal inflorescences are accompanied by opposite lateral in- florescenees. In F. amblybasis Blake the tip of the branches frequently appears to produce a dichotomous inflorescence of equal halves (text-fig. 1, F). This phenomenon is shown to have been produced by the non-development of a terminal floriferous axis, traces of which can frequently be observed. Normal opposite-lateral inflorescences also occur in this species. MANDEVILLA Lindl. The large genus Mandevilla, consisting of approximately 108 known species, is typified among the American representation of the subfamily Echitoideae largely by simple, racemose in- florescence. The mode of raceme is diverse both in number of floral members and gross construction, varying from rather lax corymbs to elongate spikes amongst neighboring species. Diagrams of the chief types of inflorescence are provided in text-fig. 4. It would appear highly significant that Mandevilla and For- steronia, in which the tendency from determinate to indetermi- nate inflorescence is clearly evident, are closely neighboring genera. The foliar glands of the majority of Forsteronias are of almost the same structure and oceurrence as those of Mande- villa subgen. Eumandevilla, and the construction of the an- thers of either genus is closely similar. An additional factor of great importance is found in the stigmata, or ““clavuncles, ”” which are massive and umbraculiform in all species of Mande- villa, and in all save one species of Forsteronia are fusiform to subcapitate. The exceptional species of the latter genus is F. simulans Woods., with an umbraculiform stigma which could [Vor. 22 24 ANNALS OF THE MISSOURI BOTANICAL GARDEN scarcely be distinguished from that of a Mandevilla. It will be recalled that the inflorescence of F. simulans further ap- proaches that of Mandevilla in its virtually racemose character. NG > — PEA UEM. Se E ee w | ^Y D G E F Fig. 4. Inflorescence structure: A—Mandevilla cal- lacatensis Mgf.; B—M. "M d igit o. i Woods. scabra (R. € S8. K.Sch —M. Mone A (Stadelm.) "Woods. Explanation in the text Although the inflorescence of the greatly preponderant num- ber of Mandevillas may be characterized as simply racemose, a few, but highly significant exceptions occur. In M. subpanic- ulata Woods. (text-fig. 4, B), indubitable evidence of reduction from a compound inflorescence is found. That such composi- 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 25 tion is the effect of reduction, rather than of aggregation as was seen in Apocynum, for example, appears to be indicated in the fact that any such compounding is not in harmony with the vegetative symmetry of the shoot, but occurs somewhat irregu- larly, and often more obscurely than in the example figured. Less obvious clues leading to the conclusion of reduction are observable in the inflorescences of the closely related M. Achrestogyne Woods. and M. bogotensis (HBK.) Woods. In M. xanthostoma (Stadelm.) Woods., an herbaceous and evidently a highly evolved member of this genus formerly in- cluded in the morphologically invalid genus Dipladenia, an in- teresting series of variations in inflorescence composition oc- curs which can scarcely be viewed other than as indicative of inflorescence phylogeny. A random collection of four such variations has been illustrated by diagrams in text-fig. 4, D-G, notes upon which follow: 1. Normal, or many-flowered, inflorescences of M. zanthos- toma are of the typical, racemose type with alternate pedicels subtended by solitary bracts. Such inflorescences average roughly eleven to thirteen functional pedicels (text-fig. 4, D). 2. Abnormal, or relatively few-flowered, inflorescences tend toward a ‘‘pseudo-bostrychoid’’ racemose type with opposite pedicels, either or both subtended by more than one bract (text- fig. 4, E). 3. Multiplicity of bracts is frequently associated with op- posite pedicels and reduced inflorescences, but not invariably (text-fig. 4, F). 4. The raceme may terminate indeterminately as well as pseudo-determinately by a terminal flower expanding slightly before the bud immediately below (text-fig. 4, G). The unusual tendencies of inflorescence structure in such species as M. subpaniculata and M. xanthostoma, differing widely in their immediate intergeneric affinities, can easily be interpreted as indicating the evolution of the simple raceme characteristic of the genus as a whole from a primitive com- pound type similar to the thyrse. This evidence is entirely con- gruous with that found in the closely neighboring genus Forsteroma. [Vor. 22 26 ANNALS OF THE MISSOURI BOTANICAL GARDEN In this connection, the need of an anatomical investigation of those inflorescences, superficially racemose but with more than a single bract subtending each flowering pedicel, must be indi- cated. In the gentianaceous genus Coutoubea, it has been found that the inflorescence, described by systematists as a raceme, differs from a true raceme in the subtension of each flower by three to four inconspicuous decussate bracts. Re- cent investigation (Woodson, 1933) has disclosed the occur- rence of occasional, abortive or dormant buds in the axils of these braets which are detectable only in microscopic prepara- tions. The ‘‘pedicels’’ of this inflorescence, therefore, are anatomically determinate axes, and the ““raceme”” itself could more aceurately be described as a greatly reduced thyrse. Corymbose and spiciform modifications of the raceme in Mandevilla are illustrated by M. callacatensis Mgf. and M. scabra (R. & S.) K. Sch. respectively (text-fig. 4, A-C). ''EMNADENIA Miers The dichotomous cincinnus is a type of inflorescence char- aeteristie of a number of apocynaceous genera, in which the flowering axis consists of a more or less elongate, naked pe- duncle terminated by two bracts of various size and texture subtending roughly equal, scorpioid branches. Noting the ab- scence of any interpretable structure in the crotch of the two frequently divarieate branches, one might be tempted to sup- pose the ageney of dichotomy of the young growing initials of the inflorescenee in early stages of development. Such super- ficially ‘‘dichotomous’’ inflorescences as those of Temnadema stellaris (Lindl.) Miers (text-fig. 5, D) may occasionally be found to produce determinate flowers in the erotch of the scor- pioid branches (text-fig. 5, E), however. The suspicion that the inflorescence of Temnadenia has arisen from the elimination of the determinate flower of a con- tinuous dichasium is heightened by an examination of the in- florescence of T. stenantha Woods. (text-fig. 5, F), recently discovered in northern Colombia, in which some of the upper- most branches may be clearly dichasial, with pedicels and braets regularly disposed, although with unmistakable ten- 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 27 dencies toward the cincinnus. The lower branches of this in- florescence are of the superficially dichotomous development, even to the disappearance of the subtending bracts at the dichotomy. B and Fig.5. I (Vell.) Woods.; B—P. portobellensis (Beurl.) Woods.; C —P. mexicana A.DC.; D-E—Temnadenia stellaris (Lindl.) Miers; F—T. stenantha Woods.; G—T. violacea (Vell.) t. The inflorescence of T. violacea (Vell.) Miers is usually simple and scorpioid (text-fig. 5, G), although individuals are not diffieult to find in which derivation from such a floral axis [Vor. 22 28 ANNALS OF THE MISSOURI BOTANICAL GARDEN as that of T. stellaris is indicated by an inconspicuous subsidi- ary axis. PnEsTONIA R. Br. The genus Prestonia, consisting of approximately fifty-four known species, is largely characterized by a type of inflores- cenee rather puzzling in its complications. In such inflores- cences as those of P. agglutinata (Jacq.) Woods. (pl. 3, fig. 1) of the section Coalitae, and P. quinquangularis (Jacq.) Spreng. of the section Acutifoliae, the axis is simple and the pedicels scorpioidally arranged. These two species are selected for il- lustration since they exhibit to some extent the peculiar zigzag habit of the peduncle correlated with the scorpioid inflores- cence of Angadenia Sagraei, evidently caused by the alternate left- and right-handed modification of a primitive dichasium, to which reference has been made previously. Such species as P. portobellensis (Beurl.) Woods. (text-fig. 5, B) and P. Meg'agros (Vell.) Woods. (text-fig. 5, A) of the section Annulares exhibit a conspicuous di- or trichotomy of the inflorescence. In all such instances among species of Prestonia bracts are virtually lacking at the place of division of the floriferous from the naked branches, by whieh they lose the clue to a eymose origin found in species of Temnadenia. There appears to be little reason to doubt such dichotomous in- florescences as that of P. portobellensis to have been derived from the suppression of the determinate floral member and sub- tending bracts as in Temnadenia, indicating evolution from a continuous dichasium. If that interpretation be allowed, it would appear logical to assume the modification of the trichot- omous inflorescences, represented by that of P. Meg'agros from an ancestral aggregate dichasium, such as that of For- steronia species. The section Tomentosae of Prestonia includes numerous spe- cies varying widely in their habit of inflorescence from simple to di- or trichotomous cincinni. In general, however, the tend- ency of inflorescence of these species is toward the condensa- tion of the pedunele, producing such virtual umbels or corymbs as those of P. mezicana A. DC. (text-fig. 5, C, in which the pe- 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 29 duncle has been unnaturally extended to clarify its composi- tion) and P. brachypoda Blake. The umbellate condition is closely approached amongst the species of Thenardia as well, in which the floriferous branches of a condensed, aggregate dichasium as that of T. floribunda HBK. (pl. 3, fig. 4) become successively foreshortened until the relatively elongate pedicels have the appearance of springing from a common receptacle subtended by numerous, involucre-like bracts as in T. Galeot- tiana Baill. The remarkable umbellate cymes of the genus Malouetia (cf. M. Schomburgki Muell.-Arg., pl. 3, fig. 2) have presumably had a similar origin. MacnosiPHONIA Muell.-Arg. Reduetion from a pluriflorous condition has evidently re- sulted in the solitary, terminal flowers characteristic of the familiar North American species of Macrosiphonia. Since the evidence of reduction (1. e., position and composition of bracts and frequency of occasional lateral flowers) is used taxonom- ically in the delimitation of species, each will be discussed briefly. Macrosiphoma hypoleuca (Benth.) Muell.-Arg., a low, suf- frutescent species of the north-central Mexican plateau, is par- tially characterized by the terminal inflorescence consisting usually of a large, white, vespertine flower borne upon a rela- tively short pedicel subtended by two rather inconspicuous, paired bracts. Below the bracts is a peduncle ordinarily of about the length of the pedicel (text-fig. 6, A). The presence of the bracts reveals the compound nature of this superficially simple inflorescence, and their paired disposition would lead us to infer modification from a primitive cyme of which the pres- ent solitary flower might logically be viewed as the surviving determinate member. Happily for this view specimens are not rare in which a sec- ond flower is found accompanying the determinate flower, ob- viously in the axil of one of the subtending bracts. An occur- rence much less common is the formation of a complete three- flowered cymule (text-fig. 6, B). In at least one instance, how- ever, a specimen from southern Mexico has been found in [VoL. 22 30 ANNALS OF THE MISSOURI BOTANICAL GARDEN which the appearance of a three-flowered raceme has been pro- duced, presumably by the translocation of the normally oppo- site nodes. Macrosiphoma Macrosiphon (Torr.) Heller differs from M. hypoleuca largely in the conspicuously foliaceous calyx-lobes, more or less petalaceous in the latter, and the broader, nearly concolorous leaves, which are conspicuously paler beneath in M. hypoleuca. The inflorescence of M. Macrosiphon also dif- fers radically from that of its neighboring species, in the pre- ponderant number of specimens examined, by the pedicel of the terminal flower becoming virtually sessile through the re- EAMA Qu» A B C D E Fig. 6. Inflorescence structure: A-B— Maorosiphonia hypoleuca (Benth. ) Muell.-Arg.; C—M. Macrosiphon (Torr. ) Heller; D-E—M. Brachysiphon (Torr.) A. Gray. Explanation in the text. duction of the pedunele (text-fig. 6, C). The primitively com- pound inflorescence is scarcely recalled save by the subtending, paired bracts which remain. These also may virtually disap- pear. No instances have been observed in which floral mem- bers have been formed in the axils of the bracts. The appear- ance of a perfectly simple, terminal flower in this species through reduction evidently depends upon the impending elimi- nation of the already inconspicuous floral bracts. Macrosi- phonia Brachysiphon (Torr.) A. Gray and M. Hesperia I. M. Johnston have been collected with both one and two flowers constituting the inflorescence (cf. M. Brachysiphon, text-fig. 6, D-E), which is practically devoid of a peduncle in either. The opinion has been expressed elsewhere (Woodson, 1930) that the North and South American species of Macrosiphonia may have had independent derivation from some common stock such as that of the large and widespread genus Mandevilla or 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE əl its primitive progenitor. This conjecture receives support in the structure of the inflorescence of either group. That of South American species may be composed of either few or soli- tary flowers, which is somewhat less common. When few flowers occur, as in M. longiflora (Desf.) Muell.-Arg. (pl. 3, fig. 5), they appear in racemose disposition. Immediately sub- tending the calyx-lobes, however, is always observable a cluster of bracts which would indicate derivation from such a com- pound floral axis as that of the thyrse, for example. It has been pointed out previously that the mode of inflorescence in Man- devilla may be supposed to have been derived from a thyrse. Additional instances in which a multiplicity of bracts sup- posedly remnant from a pluriflorous inflorescence is observable directly subtending a solitary flower borne in a lateral position are illustrated by Elytropus chilensis (A. DC.) Muell.-Arg. and Macropharynz spectabilis (Stadelm.) Woods. Specimens of either have been found in which one or two supernumerary floral members have been developed. Vinca L. and LocHNera Rchb. Vinca minor L. and Lochnera rosea (L.) Rchb. are among the most familiar garden plants. The former, popularly known as “running myrtle,” is supposedly indigenous to the northern Mediterranean area, but has been so widely and so easily eultivated that it has become naturalized over a large part of western Europe and temperate North America. The latter, known as **periwinkle," has so successfully colonized the tropics of both hemispheres that considerable doubt sur- rounds the whereabouts of its original home, although it is probably from the Old World, the provenience of the two re- maining species of the genus. A tender, ever-blooming annual, it forms an almost indispensable bedding plant in parks of the United States. The inflorescence of V. minor (text-fig. 7, C) consists of soli- tary, lateral flowers. It is interesting to note, further, that the flowers appear only upon the semi-erect shoots produced in the early spring and summer from the creeping, perennial stolons. Not all the leaves of such shoots subtend flowers. The flowers [Vor. 22 32 ANNALS OF THE MISSOURI BOTANICAL GARDEN of L. rosea are produced regularly in lateral pairs subtended by the foliage leaves (text-fig. 7, A). The pedicels are ar- ranged radially with respect to the stem axis and the subtend- ing leaf ; and the central flower always expands first. As the leaves are decussate, pairs of flowers are in nearly all cases found in the axil of only one of the paired leaves, ascending the axis spirally. It is elear that the axis of the inflorescence of L. rosea is a sympodium derived from a cincinnus. Among many flowering individuals of L. rosea occasional ab- normalities are found in which only a single flower is formed in the axil of a subtending leaf (text-fig. 7, B). Considering the very close relationship of the genera V inca and Lochnera, it RN di a $ $ Pf g. 7. Inflorescence structure: A-B—Loch- nera rosea (L.) Rehb.; C—V inca minor L seems more than likely that the indeterminate flowering axis of the former, bearing solitary, lateral flowers in spiral fashion, should represent derivation from such a sympodial axis as that of the latter. Discussion The necessity of a general review of literature concerning the inflorescence is obviated by the very full historical account given by Parkin (1914), whose admirable treatment of the evolution of the inflorescence remains our most comprehen- sive. In view of the great importance of that memoir and of the somewhat different conclusions to which the present writer has arrived, it would doubtless be both appropriate and serv- iceable to combine a discussion of the apparent evolution of the inflorescence in Apocynaceae with a rather extended criticism 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 33 of the views so ably expressed by Parkin. In introduction, it is scarcely possible to recall too forcefully that this writer's opinions are based upon observations within a wide range of a single family. Although intensive study within a single natural group, even though large, entails indubitable benefits, it can scarcely be said to result in conclusions which are appli- cable to all groups equally. This will become apparent in a sub- sequent paragraph. On the other hand, it rarely happens that a general biological principle, such as that influencing the mod- ifieation of the inflorescence, can be true of a single natural group, and of none of its closely neighboring relatives. In his study of the inflorescence herbarium specimens were virtually ignored by Parkin upon the assumption of imperfect evidence offered by them. However, recourse was had to a large number of living plants selected from species commonly eultivated in English gardens. Special emphasis was placed in this selection upon dicotyledonous representatives of the so- ealled **primitive alliances," such as Magnoliales, Ranales, Papaverales, ete. . Evidently inspired by the frequent oceur- rence of solitary, terminal flowers in plants belonging to such orders of established antiquity, Parkin was led to conclude that in the primitive inflorescence **flowers were originally borne in a solitary fashion, and that the clustering of them together without intervening foliage is a later development.” Upon this assumption, he quoted the familiar Euclidean ‘‘reductio ad absurdum ”’ to dispel Church’s interpretation of the solitary flower of Pyrus Cydonia as a reduction from the many-flowered inflorescences of such species as P. Aucuparia (Church, 1908). Parkin's statement that the solitary, terminal flower is an ancient structure approaches the platitudinous when one re- fleets that the flower itself may be defined as a highly special- ized, contracted short shoot bearing a collection of sporophylls with or without an accompanying envelope. Quite likely, through past ages isolated progenitors of the modern Angio- spermous flower, occurring terminally and isolated by inter- vening foliage, became aggregated to form a specialized branch system coincident with the evolution of the flower itself. But to trace the very foundations of the evolutionary development [Vor. 22 34 ANNALS OF THE MISSOURI BOTANICAL GARDEN of the inflorescence among the extant Angiospermae is a dan- gerous undertaking none the less. No doubt the floral organi- zation of the Ranunculaceae may be considered as ““primitive”” from several points of view, but few would extend that inter- pretation to the herbaceous habit so conspicuously predomi- nant in the family. Similarly, although Papaver is clearly a genus of great antiquity, we would be slow to describe its gynoecium as ‘‘primitive’’ or elemental. It should clearly be recognized that groups of plants may possess palingenetic and cenogenetic characters side by side; our recognition of the former should not confuse our interpretation of the latter, al- though not to do so is manifestly difficult at times. In supporting his contention of the primitive nature of the solitary, terminal flower among Angiosperms, Parkin cited the existence of terminal strobili in the Bennettitalean-Cycadalean plexus, now generally assumed to represent a primordium from which the flowering plants may have evolved. He noted, how- ever, that the solitary, terminal strobili of the Cycads are oc- easionally accompanied by supernumerary members at the stem apex. The continuation of the vegetative growth is ac- complished by a lateral bud, and a sympodium results. The anatomy of the axis of Cycads discloses the existence of a tele- scoped! sympodium essentially eymose in its composition, as Miss Smith (1907) has pointed out. In the Bennettitales, also, the branching of the Triassic Williamsonia ( Anomozamites) angustifolia (Nathorst, 1902) and the Jurassic Wielandiella is of the sympodial type. The branching axis of the former, con- sisting of a system of relatively slender stems terminating in a strobilus surrounded by a rosette of leaves below which two equal branches are found similarly compounded, bears an un- mistakable similarity to the modern dichasium of Angio- * Apparently A. P. De Candolle (1827) was the first to use the metaphor of the telescope in deseribing extension and condensation of axes: ‘‘Je me ferais vind être comprendre plus complètement par une métaphore bien grossiére: en une branche florale organisée comme une lunette d’approche, qui por a un pédieelle au haut de chaeun des tubes qui la composent: que tou s les Aga soient déboités et alongés, vous aurez une grappe; repoussez ees tubes à moitié, ce sera encore une grappe, mais trös-courte; rentrez-les complótement, et vous aurez une ombelle terminale, 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 35 sperms, and the resemblance would appear to be more than fortuitous. It would appear a fruitless task to search for the earliest indications of the inflorescence among the extant flowering plants: the origin of the inflorescence is at least as remote as the origin of the flower, and a greater antiquity seems probable from the evidence of paleobotany. Development of the pluriflorous inflorescence from the soli- tary, terminal flower is accomplished according to Parkin by the appearance of subordinate, lateral flowers literally ““pushed out’’ of the axils of the subtending leaves of the orig- inal, terminal floral member. The continual ““emitting”” of sub- ordinate flowers forms the first type of compound inflores- cence: the dichasium. This process is illustrated in the genus Papaver by such species as P. Rhoeas, P. strictum, and P. pilosum in a series of floral accumulation, as we might express it for want of a better term. The naivety of this conception of floral ““emission”” is heightened by the causal agent, ascribed by Parkin (1914) with some qualification to **a superabundance of reproductive ma- terial, more than the shoot could utilize in the formation of a single flower." This somewhat feeble effort to drag the con- tinuity of the germ-plasm into the discussion of the inflores- cence serves to illustrate rather indirectly the perplexing fact that although much is known concerning the ““geometry”” of plants, little indeed is understood of the causal mechanism re- sponsible for it. Our easiest task, consequently, is to express structural modifieations of plants in terms of the geometrical pattern approaching more nearly universal application throughout specific, natural groups. One might cite the ex- ample of dichasial branching, which is infinite in its theoretical extension. Practically, however, certain internal and external factors limit the expression of the dichasium to a definite nu- merical range of floral members in one species, and to a slightly different range in another. In the end, the morphologist finds in nearly any highly de- veloped group a “reduction series” in evolutionary history much easier to read than an “ascending series." Time and again the morphologist has found himself deceived by the ap- [Vor. 22 36 ANNALS OF THE MISSOURI BOTANICAL GARDEN parent “simplicity”” of species and genera which have suffered reduetion from the more elaborate geometrical pattern of their relatives during the inserutable course of evolution. It should not be necessary to recall that our modern flowering plants represent a very ancient and specialized development whose foundations are still, and possibly always will be, exasperat- ingly obseure. The so-called ‘‘primitive alliances’’ are evi- dently searcely more ancient, from available paleobotanical records, than others from which they have been separated by the imposition of an imperfect system of classification. The herbaceous habit of such genera as Papaver, for example, might be cited as but one evidence of a long and specialized evolutionary development. From the evidence briefly reviewed for Echites umbellata, the evolution of the uniflorous condition from pluriflory through the agency of reduction appears plausible, for we have the evidence of reduction at hand in the form of ““empty”” bracts and oceasional abortive buds. The opposite assumption, which would logically include the inference that such buds are in a nascent stage, would appear preposterous. This writer finds ample sympathy for the independent and earlier views of Church (1908) respecting the evolution of the inflorescence in Pyrus to which reference has been made in a preceding paragraph. The result of reduction upon the development of the remain- ing floral members of the inflorescence described in the pre- vious paragraph is strikingly similar to that of the old horticultural practice of “picking out the suekers,"' or **disbud- ding,’’ in the cultivation of tomatoes, grapes, and other culti- vated plants, by which the growers seek to influence the subse- quent development of the remaining members. **Disbudding"' is also practiced in the foreing of cultivated chrysanthemums, ete., and the time of blooming can be controlled largely through this agency. Gardeners using the same method in the cultiva- tion of such plants as Paeonia have observed that plants which ““disbud themselves,”” e. g., by natural abortion of floral buds, bloom earlier and have larger flowers than those whose full floral complement develops unchecked. 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 5r Much the same effect can be produced experimentally by am- putating floral buds of such an inflorescence as the multiflorous thyrse of the apoeynaceous Amsonia Tabernaemontana Walt. Precaution must be taken, however, to arrest the development of the buds at a sufficiently early stage and with the slightest possible injury. All experiments made in this manner have not been wholly satisfactory, due probably at least in part to the latter factor. Since it is impossible wholly to exclude the factor of injury, even slight success in modifying the symmetry of the cyme may be regarded as significant, the more subtle in- fluence of natural abortion being beyond perfect imitation. Concerning the transition of the indeterminate from the de- terminate inflorescence, Parkin (1914) envisions a process in which **the number of lateral floral shoots increase, so that the main terminal flower no longer blooms first. The flowers then tend to open in acropetal succession from the commencement. The next step comes about through the flower-buds of the up- permost part of the inflorescence never expanding; this part, in fact, becomes arrested in its development and finally aborts, leaving a mere filament or protuberance in its place. In this way the original terminal flower disappears. The inflorescence is now racemose.”” In the light of what the author has said as previously reported concerning the supposed primitive, few- flowered inflorescence, one can scarcely avoid confusion in attempting to harmonize this last statement with the innumer- able-flowered cymes of Statice and the relatively few-flowered racemes of Plumbago among Plumbaginaceae, to cite only one example. From Parkin's statement one would be justified in expecting racemes to contain more numerous flowers than cymes. Such is scarcely the case. The flexibility of the extent of the cyme might be likened in potentiality to that of the com- pound interest table. On the other hand, failure of the apex of the peduncle to pro- duce a terminal flower is not an inalienable characteristic of the raceme. In racemes of but few flowers a terminal member fre- quently occurs. This has already been demonstrated for the genus Mandevilla of the Apocynaceae (cf. text-fig. 4, A-G). [VoL. 22 38 ANNALS OF THE MISSOURI BOTANICAL GARDEN In perhaps no other partieular are the observations of Parkin and those of this writer in greater contrast than in the im- mediate agency involved in the transformation of the indeter- minate from the determinate mode of inflorescence. Both views are based upon anatomical-physiological assumptions. The view of Parkin may be epitomized in the statement that as the lower floral members increase in number, the development of the apical members is hindered by the increasing demands upon the conductive facilities of the axis, supposedly. In the preced- ing observations among the Apocynaceae, on the other hand, a totally different process is adduced. It has been found in Echites that in the instance of two equivalent eymules of identi- cal composition the reduction of one or more floral members of one will hasten the development of the other component mem- bers, thus effectively disrupting the characteristic ‘‘sym- metry’’ of the two equivalent eymules. It has been inferred with ample evidence that the cincinnus and the bostryx have evolved from the dichasium in this family through the reduc- tion of a whole cymule to a solitary flower, thus invariably hastening its development second only to the primary deter- minate flower with which it becomes associated. It is understood that reduction has effected one of the lateral eymules of a continuous dichasium to the production of the cincinnus or the bostryx. The effect of the reduction of both lateral cymules of an aggregate dichasium may be computed artificially: its demonstration in nature is afforded in the large genus Forsteronia, by which it has been shown that closely re- lated species intergrade from aggregate dichasial inflores- cences to a raceme through various modifications of the thyrse. Having traced the development of the more important types of determinate and indeterminate inflorescences amongst the Apocynaceae, it becomes easy to extend our understanding ; by the reduction of the pedicel to derive the spike from the raceme, and by the reduction of the peduncle and the compensating elongation of the pedicel to produce the corymb and the umbel, which may evidently be derived from both determinate and indeterminate forebears. Although generalizations are fre- 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 39 quently weak in the face of detailed attack, it may be permitted to visualize the writer's conception of the evolution of the fundamental types of inflorescence in Apocynaceae as repre- sented in text-fig. 8. At this juncture prominence should be given to the fact that reduetion of floral members does not always hasten the de- thyrse solitary flower o Ae aggregate dichasium cincinnus dichasium Fig. 8. Schematic diagram of the supposed en of certain types of inflorescence in Apocynaceae. Explanation in the velopment of the remaining members of a cyme. Early in the course of these investigations recourse was had to members of other families to ascertain whether the same procedure is com- mon. Selecting families containing familiar representatives bearing cymose inflorescences, it was soon found that the same principles discovered in Apocynaceae are apparently appli- eable in members of such families as Ranunculaceae, Rosaceae, Primulaceae, Gentianaceae, ete. In the family Caryophyl- laceae, characterized by the predominance of the cyme, how- [Vor. 22 40 ANNALS OF THE MISSOURI BOTANICAL GARDEN ever, the principle of reduction in the disruption of cymose symmetry appears to be completely reversed. One needs but examine a colony of such a species as Arenaria patula to dis- cover that the terminal flower of a somewhat reduced cymule aetually blooms somewhat later than the terminal member of an equivalent, less-reduced branch. The reversal, moreover, appears widespread if not universal throughout the family, as well as in members of Convolvulaceae, and it would be odd in- deed if it were unique among the flowering plants in this respect. The solution of the enigma which is thus introduced, as well as the actual mechanism of the phenomena noted in Apocy- naceae, ete., would appear to lie in a study of the anatomy of the inflorescence, a virtually untouched field since the rather few students of the inflorescence as a whole have limited them- selves to gross morphology save in special instances. A de- cidedly hypothetical conception which must be tested by a future anatomical study of the inflorescence to explain the divergent results of reduction in Apocynaceae on the one hand and Caryophyllaceae on the other, might involve the divergent reduction of vascular tissues within the respective inflores- cences: the reduction of vascular tissues and its attendant physiological effects possibly being limited to the pedicels of discarded members in Apocynaceae, thus redistributing the full flow of the conductive elements to the fewer floral members which would benefit accordingly; the reduction of vascular tis- sues in the Caryophyllaceae on the other hand taking place in the peduncular axes, thus hindering the development of the branch system as a whole. At any event, the primitive nature of the eyme in Caryophyllaceae should be established by future studies of the inflorescence of the group; here also, as in Apocynaceae, ete., the solitary flower is evidently a condition derived from pluriflory. Minor points of criticism of Parkin’s observations, in the light of the present writer's investigations on the inflorescence of Apocynaceae, may be dismissed without discussion. It should be noted that both are in agreement concerning the primitive nature of determinate and the derived condition of 1935] WOODSON—INFLORESCENCE OF APOCYNACEAE 41 indeterminate inflorescences. It is suspected that Parkin's inferences concerning the method by which the indeterminate mode was derived from the determinate were unfortunately af- fected by the fact that his illustrations include no perfectly symmetrical representatives of the latter. Itis easily appreci- ated that the most characteristic features of the dichasium, for example, are lost in species with alternate, or spiral phyllotaxy, as has been indicated in an earlier section of this paper. SUMMARY The following conclusions have resulted from a study of the inflorescence structure of numerous representatives of Apocy- naceae throughout the distribution of the family, as revealed by both living and herbarium specimens. Special emphasis has been placed upon the American representatives of the sub- family Echitoideae, an evidently natural group lately studied by this writer (Woodson, 1933) from the combined aspects of floral anatomy and taxonomy. 1. The dichasial cyme appears to be the primitive inflores- cence of the family. 2. The symmetry characteristic of the composition and se- quence of development of equivalent cymules of the inflores- cence of Echites umbellata Jacq. is found to be profoundly modified by unequal reduction of the component floral members. Reduction of one or more lateral members of a cyme is found to accompany a somewhat hastened development of the remain- ing members. This effect of reduction appears constant, or relatively so, amongst Apocynaceae and certain other families, but appears to be reversed in Caryophyllaceae (and Convolvu- laceae) for reasons which are briefly conjectured. Such an agency is found to parallel long-established horticultural prac- tices, such as “picking out the suckers,’’ and ““disbudding.”” 3. Reduction of a single branch of a continuous dichasium always to the right or left of the determinate flower, or to the right and left alternately, results in the production of a bostryx or a cincinnus respectively. 4. The transformation of the inflorescence from an aggre- [Vor. 22, 1935] 42 ANNALS OF THE MISSOURI BOTANICAL GARDEN gate dichasium to a raceme through various modifications of the thyrse is traced amongst the species of the genus Forsteronia. This progression has evidently been actuated by the progressive reduction in acropetal fashion of both the decussate pairs of branches of an aggregate dichasium. 5. The spike is derived from the raceme; the corymb and the umbel are derivative from both determinate and indeterminate inflorescences. 6. Solitary flowers, whether terminal or lateral, are appar- ently derived from a previous condition of pluriflory through the ageney of reduction. LITERATURE CITED Church, A. H, (1908). Types of floral mechanism, pp, 10-13, Oxford. De Candolle, A. P. (1827). Organogr — végétale 1: 408-409. Paris. Eiehler, A. W. (1875). Blüthendiagrame. pp. 33-42. Leipzig. Goebel, K. (1931). Blüthenbildung e Sprossgestaltung. pp. Ls 238. Jena. Gray, A. (1907). Structural botany. pp. 151-158. ed. 6. New Jackson, B. D. (1900). A glossary of botanic terms. London. Nathorst, A. G. (1902). Beitrüge zur Kenntnis einiger Mesozoischen Cycadophyten. Handl. Kgl. Svensk. Vetensk.-Akad. 36*: 28. pl. 3 Parkin, J. (1914). The evolution of the inflorescence. Jot Linn. Soc. 42: 511-562. 1914. Roeper, J. (1826). Observationes aliquot in florum inflorescentiarumque naturam. 1: 433-466. Sehumann, K. (1895). T in Engler, A. and K. Prantl, Die Natürliehen Pflanzenfamilien. 4, Abt. 109-189. Leipzig. Smith, F. G. (1907). "nte of the trunk and development of the microspor- angium of Cycads. Bot. Gaz, 43: 187-204. Woodson, R. E., Jr. (1930). Studies in the M I. A critical study of the Apecyneiéens, fee Mo. Bot. Gard. -212. —— — — ——-, (1928a). Ibid. II. A revision of the genus Stemmadenia. Ibid. 15: 341-378. — ——-, (1928b). Ibid. III. A monograph of the genus Amsonia. Ibid. 15: 379—434 — — —————, (1933). Ibid. IV. The American genera of Echitoideae. Ibid. 20: 605—790, et seq. ——, Observations on the abnormal fibre development of certain gentia- naceous flowers. [Vor. 22, 1935] 44 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 1 Figs. 1-9, 4-5. Variation in inflorescence structure amongst individuals of Echites umbellata Jaeq. í Fig. 3. Inflorescence structure of E. crassipes A. Rich. ANN. Mo. Bor. GARD., Vor. 22, 1935 PLATE 1 WOODSON—INFLORESCENCE OF APOCYNACEAE dM di Me Pa a: EAE Pa WETTE rnit te ee *: o NE o p As J oS 3d iar Ap bul BG, is? ud + Mis ii d E. = ANN. Mo. Bor. GARD., Vor. 22, 1935 PLATE 2 WOODSON—INFLORESCENCE OF APOCYNACEAE [ Vor. 22, 1935] ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 3 Prestonia agglutinata (Jacq.) Woods. Malouetia Schomburgki Muell.-Arg. Fernaldia pandurata (A.DC.) Woods. Thenardia floribunda HBK. Macrosiphonia longiflora (Desf.) Muell.-Arg. PLATE 3 ANN. Mo. Bor. GARD., Vor. 22, 1935 AS iy N \ pA wA Wy, , X US ya AER + RÓS y MUA = WOODSON—INFLORESCENCE OF APOCYNACEAE A NEW KALLSTROEMIA FROM TEXAS LOUIS WILLIAMS Washington University Fellow in Botany Kallstroemia hirsuta n.sp.' Ascending or erect annual, 1-2 dm. high; stems 1-several from the base, densely hirsute and strigose-pubescent, the hirsute hairs with pustulate bases; leaves 24.5 em. long, oblong to ovate-oblong, petiolate, densely hirsute and strigose-pubescent on both surfaces, the petiole about as long as the lower pair of leaflets; leaflets 3-6 pairs, sessile, obliquely oblong or ovate, 7-14 mm. long, 3-8 mm. wide, obtuse or acute, about equal in size, the terminal more oblique; stipules lance-oblong, 3-4 mm. long, 1-1.5 mm. wide; calyx- lobes linear to linear-lanceolate, 10-13 mm. long, 1.5-2.5 mm. wide, hirsute and strigose-pubescent on the outer surface, the chartaceous margins inrolled in fruit; petals about 5, deltoid- ovate, 18-23 mm. long, probably white or yellowish, not mar- cescent ; fruit compressed-oval in outline, 8-10 mm. wide, 5-6 mm. high, outer surface of carpels not tuberculate but the median portion and the two edges prominent and connected by transverse ridges, densely hirsute, particularly on the raised portions ; persistent style or beak 8-10 mm. long, hirsute at the inconspicuously swollen base, stigmatic portion 1.5-2 mm. long, furcate. Texas: Langtry, Valverde County, May, 1913, C. R. Orcutt, 6126 (Mo. Bot. Gard. Herb.) TYPE. This species is most closely allied to K. grandiflora Torr., from which it may be separated by the following characters: the entire plant is densely pubescent with hirsute and strigose 1 Kallstroemia hirsuta sp. nov. Annua 1-2 dm. alta, dense hirsuta et strigoso- pubescente; foliis oblongis aut oblongo-ovatis utrinque pubescentibus; foliolis 3-6 paribus, sessilibus, 7-14 mm. longis, 3-8 mm. latis; stipulis lanceolato- oblongis, 3-4 mm. longis, 1-1.5 mm. latis; lobis calycis linearibus vel lineari- laneeolatis, 10-13 mm, longis, 1.5-2.5 mm. latis; petalis ca. 5, deltoideo-ovatis, 18-23 mm. longis; carpellis non tuberculatis parte medio et marginibus prom- inentibus; stylo 8-10 mm. longo, ad basem hirsuto. Issued March 25, 1935. ANN. Mo. Bor. GARD., VOL. 22, 1935. (49) [Vor. 22, 1935] 50 ANNALS OF THE MISSOURI BOTANICAL GARDEN hairs intermixed; the carpels are not tuberculate on the outer surface nor merely equally strigose but have the median por- tion and the outer edges raised and these connected with trans- verse ridges, and especially the raised portions hirsute with a small amount of strigose pubescence intermixed ; beak hirsute at the base only instead of strigose nearly to the stigmatic portion. EXPLANATION Or PLATE PLATE 4 Kallstroemia hirsuta Williams. From the type specimen, Charles Russell Orcutt, 6126, in the Missouri Botanical Garden Herbarium. % natural size. ANN. Mo. Bor. Garp., Vor. 22, 1935 PLATE 4 WILLIAMS—KALLSTROEMIA HIRSUTA A CRITICAL STUDY OF CERTAIN EPAPPOSE GEN- ERA OF THE HELIANTHEAE-VERBESININAE OF THE NATURAL FAMILY COMPOSITAE! WARD McCLINTIC SHARP Formerly Assistant in Botany in the Henry Shaw School of Botany of Washington University INTRODUCTION AND ACKNOWLEDGMENTS This paper endeavors to present the results of a study of certain epappose genera of the tribe Heliantheae-Verbesininae of the natural family Compositae. The genera Balsamorhiza, Zaluzania, Greenmaniella, Iostephane, Echinacea, and Rati- bida have been dealt with, first, with regards to their compara- tive morphology and probable phylogenetic relationships, and second, from a taxonomic point of view. The other genera Which are occasionally referred to have been studied only for their phylogenetie and generic relationships. The more closely related of these genera, and especially the epappose ones, are included in the generic key. The writer takes this opportunity to express his appreciation and gratitude to those who have made this work possible. Par- ticularly is he under obligation to Dr. George T. Moore for the facilities of the herbarium and library of the Missouri Botan- ical Garden. Sincere appreciation is hereby expressed to the eurators of the various herbaria from which specimens have been borrowed, thus making this study possible. He is also in- debted to Miss Nell C. Horner, Librarian, for assistance in bibliographical matters. Especial gratitude is due Dr. J. M. Greenman, under whose direction and supervision this work has been carried out, for ready and helpful suggestions given throughout this study. "An investigation carried out at the Missouri Botanical Garden in the Graduate Laboratory of the Henry Shaw School of Botany of Washington University, and submitted as a thesis in partial fulfillment of the requirements for the degree of doctor of philosophy in the Henry Shaw School of Botany of Washington University. Issued March 25, 1935. ANN. Mo. Bor. GARD., Vor. 22, 1935. (51) [Vor. 22 52 ANNALS OF THE MISSOURI BOTANICAL GARDEN Parr I. GENERAL Discussion COMPARATIVE MORPHOLOGY The genera previously mentioned, as well as Dracopis and Rudbeckia, will be here contrasted and compared. The chief distinetions are found in the involucre, the receptacle and pales, the corollas, and the achenes. The achenes and corollas present the characters of major importance in the generic differentiation. Involucre.—The involucres of the genera studied are so uni- form that it would be rather difficult to recognize any one of these genera on this character alone. The variations, however, do fall into five distinct categories: (1) the Ratibida-Dracopis type; (2) the Rudbeckia-Echinacea type; (3) the Balsamo- rhiza-Iostephane type; (4) the Zaluzania type; and (5) the Greenmaniella type. The bracts of the involucre of Ratibida and Dracopis are so similar that it would be impossible to separate them on form alone. They are biseriate, the outer series linear, reflexed or spreading, herbaceous, and two to three times longer than the inner. Those of the inner series are about as long as the pales, but they are more herbaceous in texture, have acute apices, and a plane surface. The pales are conduplicate and marked by a brownish resin duct along each margin. The character of the involucre would be sufficient to separate these genera from those that follow. The Rudbeckia-Echinacea type of involucre is of little im- portance in generic differentiation. The bracts are two- to three-seriate, herbaceous, linear to linear-lanceolate, and more or less spreading or reflexed. The series are of about equal length, thus differing from the Ratibida-Dracopis type. In Echinacea the inner series is attenuated and often shows a transition into the pales, as exemplified in E. paradoza. The involucres of Balsamorhiza and Iostephane differ from those of the other genera studied in that the outer series of bracts is usually much larger than the inner series. The bracts often become foliaceous and extend beyond the disk, a condi- tion which reaches its greatest degree of development in Bal- 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 53 samorhiza Bolanderi, B. deltoidea, and B. macrolepis. Chrys- opetala, a subgenus of /ostephane, is an exception, its involucre being of the helianthoid type. There is so much variation in the size of the bracts in this type that the character is of value only in specifie diagnosis. In Zaluzama the bracts are two or three seriate, and are graduated but never exceed the disk in length. The outer series is herbaceous, while the inner series often becomes subherba- ceous or subcoriaceous, but not membranaceous. Theinvolucre in this genus is of little value for specific characters. The involucre in the genus Greenmaniella is similar to that of Zaluzania but in the former the bracts of the inner series are distinetly membranaceous and marked by six or more conspic- uous brownish-resinous parallel nerves. This character is also present but less pronounced in the outer series. The mem- braneous nature of the inner involucral bracts, as well as the pales, presents a character by which these two genera may be readily distinguished. The Receptacle and Pales.—The pales (bracts of the recep- tacle) are quite similar in all the genera studied and of little importance in generic differentiation; hence they may be dis- eussed as a part of the receptacle rather than treated sep- arately. 'The form of the receptacle falls into three categories, namely, (1) the columnar; (2) the conical; and (3) the broadly convex. The columnar type of receptacle is typified in Ratibida, Dracopis, and also in the Laciniatae, Heliopsidae, and Laevi- gatae sections of the genus Rudbeckia. It reaches its highest degree of development in Rudbeckia maxima, where it often attains a length of 5 em.; the maximum length in Ratibida is 3 to 3.5 cm., as in R. columnaris and R. peduncularis. In Rati- bida and Dracopis the receptacle is narrowly cylindrical from the base upwards, whereas in Rudbeckia it becomes broadened somewhat towards the base. In Ratibida and Rudbeckia the pales are deciduous from the receptacle after maturity, this being more pronounced in Ratibida, and they are bordered along each margin by a large elongated brownish or purplish [Vor. 22 54 ANNALS OF THE MISSOURI BOTANICAL GARDEN resin duct. In the columnar sections of Rudbeckia the pales are absent, or if present represented by fine brownish lines. Echinacea, I ostephane, Zaluzama, Greenmamella, and a ma- jority of the species of Rudbeckia fall into the second category with conical receptacles. The maximum development is at- tained in Echinacea and the subgenus Euiostephane of Joste- phane. The receptacle in Echinacea is elongated, 1-1.5 cm. high. In Euiostephane the receptacle is also conical, but the head appears hemispherical, due to the outer pales being longer than theinner. The pales of the two latter genera are stout and terminate in straight or slightly curved spiny tips. This is more pronounced in Echinacea, where the stout slender spiny tips extend beyond the corollas, thus giving the head an echi- nate appearance. The pales of Zaluzania are without char- acters of any generic significance, but the pales of Green- maniella are thin and membranaceous and marked by several brownish parallel veins, a type of pale rather unusual for the Verbesininae. Balsamorhiza is characterized by a broadly convex recep- tacle, a type also characteristic of Wyethia, a pappose genus of the Verbesininae. In both of these genera the heads are very large and so very similar that for one not familiar with the two genera it is necessary to examine the achenes before one is certain as to the genus. The large broadly convex receptacle of Balsamorhiza may distinguish it from other epappose genera of the Verbesininae. Corollas.—The ray-flowers are of two types, styliferous, as in Balsamorhiza, Greenmaniella, and Zaluzania, or without styles, as in Echinacea, Dracopis, Iostephane, Ratibida, ete. This difference, since it is a constant character and since it re- sults in either fertile or sterile achenes of the ray-flowers, serves as a major factor in separating these genera into two main divisions. The lamina and tube present no marks of generic importance. The rays are present in all the species of these genera except Zaluzania discoidea, Z. Pringles, and Bal- samorhiza invenusta, and here the characters of the achene and disk-corolla are sufficient to identify the genus. In these 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 55 genera the ligules are commonly yellow. In Echinacea and Iostephane the rays in the majority of the species are purple, but in Ratibida they are often dark purple blotched at the base. The form and size of the disk-corollas are very uniform for each genus and present good generic characters in a majority of cases (pl. 5), but they are of minor importance for species characters. Two main groups may be recognized: (1) disk- corollas with a narrowly constricted tube which expands above into a broad tubular or tubular-campanulate throat and the 5-lobed limb; and (2) disk-corollas without a definite tube. Balsamorhiza, Greenmaniella, Dracopis, Iostephane, and Zaluzania fall within the first group. Zaluzania differs from these genera in that the base of the tube is expanded so as to completely cap the crown of the achene. This and the epappose achenes and the styliferous ray-flowers are the outstanding characters of the genus. This type of disk-corolla is also found in a few species of Viguiera. A major difference between Greenmaniella and Zaluzama is also found in the base of the corolla-tube. In the former the base of the tube is narrow and seated inside the cup-like crown of the achene. The lower por- tion of the throat of the corolla in Balsamorhiza and Dracopis is often enlarged somewhat on one side, but this varies so as to be of little generic or specific importance. Echinacea and Ratibida belong to the second group. The disk-corolla in Echinacea is enlarged at the base into a bulb- like structure which contracts above into the cylindrical, 5-lobed corolla-tube. The stamens are attached near the swollen base of the corolla. Ratibida differs from Echinacea in that the corolla is narrowed slightly at the base and then expands above into the inflated corolla-tube. The stamens, as in Echinacea, are also attached near the base of the corolla. The anthers and style appendages conform to those of the Verbesininae. Achenes.—The achenes are relatively uniform in each genus included in this study. The ray-achenes are trigonal, but in Echinacea, Ratibida, and Rudbeckia they are never fully enough developed to be of any taxonomic importance. In [Vor. 22 56 ANNALS OF THE MISSOURI BOTANICAL GARDEN Zalueania mollissima, Z. subcordata, and Z. augusta the pappus of the ray-achenes furnishes useful supplementary characters in the key to the species. The disk-achenes are more or less characteristic for each genus (pl. 5). Comparative studies are based on (1) the out- line in transverse section, and (2) the nature of the erown of the achene. The general types of outline are terete, rhom- boidal, and quadrangular. The majority of the genera fall in the two latter categories. The pappus, when present, is very rudimentary in comparison with that found in other Helian- theae. It varies from a toothed crown in Echinacea, two awn- like teeth in Ratibida, to the shallow cup- or disk-like crowns in Dracopis and Greenmaniella. Dracopis is characterized by a small, black, terete, linear- cuneate achene which is minutely striated and conspicuously transversely wrinkled. The apex of the achene is bordered by a minute, brownish, disk-like crown, this type being particu- larly characteristic of the genus. These characters, however, are based upon the mature achene. The transversely wrinkled surface is demonstrated to a lesser degree in many species of Rudbeckia. The ray-achenes in Dracopis are rhomboidal, pubescent, and also shorter and thicker than those of the disk; the crown also has a more pronounced cup-like pappus than the disk-achenes. The achene of Greenmamella also approaches the terete type of outline, In transverse section it is broadly subangulate to subterete at maturity. In longitudinal outline it is subfusi- form, that is, the broadest part is in the upper half, from which it narrows gradually towards the apex and the base. The achene is terminated by a shallow cup-like pappus with knob- like projections at the angles. The achenial character is of generie importance, since it is quite unlike those of other genera with fertile ray-flowers. In Echinacea, Iostephane, and Zaluzania the achenes have a general rhomboidal outline in transverse section. The achene of Zaluzama is capped by the expanded base of the corolla-tube, and as a result no pappus is ever present, a character which 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 57 readily distinguishes this genus from all others in this study. In Iostephane the achene is slightly rounded at the point of in- sertion of the corolla. A pappus is normally absent, but if present it is represented by one or two minute paleaceous setae. The achene in Echinacea terminates abruptly in the jagged and toothed border surrounding the concave depression at the sum- mit where the bulbous base of the corolla is seated. "The angle facing the subtending pale has developed an elongated tooth in a majority of the species. The achenes of Ratibida differ from those of the other genera in being laterally flattened or compressed-rhomboidal. In the former type, which is the more common, the pappus is com- posed of two awn-like teeth on each edge of the achene. The flattened edges are also more or less winged, as in R. pedun- cularis. Thelatter type of achene is less compressed and with- out a pappus, as in R. pinnata and R. mexicana. Balsamorhiza differs from the above genera in that the achenes are from 6 to 8 mm. long and quadrangular in cross- section. Their length is two to three times that of the average achene, which, with the quadrangular outline and the absence of a pappus, is sufficient to distinguish the achenes of this genus from any other epappose genus of the Verbesininae. PROBABLE PHYLOGENY A discussion of the evolutionary development of this group of genera, as of any large group of closely related genera, must be more or less hypothetical. Nevertheless, from the fore- going discussion a probable line of descent may be surmised. The ancestral stock of Balsamorhiza, Iostephane, Zaluzania, Ratibida, Dracopis, and Echinacea, within the subtribe Verbe- sininae, probably has been evolved from two generic complexes, namely, the Viguiera complex and the Echinacea-Rudbeckia complex. The former complex may be defined as including those pappose genera, probably extinct, which were similar to extant genera, Helianthus, Viguiera, Tithonia, ete. Bal- somarhiea, Iostephane, Zaluzania, and probably Greenmani- ella, have been derived from this complex. The Echinacea- [Vor. 22 58 ANNALS OF THE MISSOURI BOTANICAL GARDEN Rudbeckia complex probably comprised plants with a more or less columnar or conical receptacle similar to the pres- ent species of Rudbeckia, Echinacea, ete. The ancestry of Dracopis, Rudbeckia, Ratibida and Echinacea may be sought within this complex. Iostephane has taken its origin from a helianthoid ancestor. Its evolution has progressed along two lines, one terminating in the subgenus Chrysopetala, the other in Euiostephane. The former is more closely related to H elianthus in the nature of the involucre, receptacle, achenes, and the corolla. Euiostephane is less closely connected, its large conical receptacle, echinate pales, and purple rays showing a more remote relationship. It is also probably an older entity. The presence of minute setose squamellae, which is uncommon, indicates a relationship with this complex. The genus is confined to the Mexican Plateau, an old region geologically speaking. The relationships of Zaluzania are with the Viguiera com- plex, but its exact ancestral stock is somewhat in question. Evidence at present points to affinities with a Vigwera- Gymnoloma type of ancestor. The fertile ray-flowers of Zaluzama, along with other evidence such as the woody or suffruticose habit, the simple entire leaves in a majority of the species, and a distribution chiefly confined to the Mexican Plateau, all point to a primitive rather than a more recent group. The herbaceous species of this genus, with lobed or pinnatisect leaves, are of more recent origin. The absence of a pappus in the disk-flowers is probably due to displacement by the dilated corolla base. This is further borne out by the presence of a pappus in the ray-flowers, in which the corolla base is not dilated. The affinities of Greenmaniella are in doubt since its present position in the tribe is not certain. The membranaceous nature of the involucre and pales indicates affinities other than the Verbesininae, but the determination of its exact relationship must await the collection and study of additional material. Balsamorhiza is closely related to Wyethia from which it has probably been derived. The more primitive species of this 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 59 genus, as in many species of Wyethia, have entire, more or less ovate, pinnate, basal leaves, with subscapose stems. These species are also more aggressive and more widely distributed. Wyethia has affinities with the Viguiera complex, but this genus is of more recent origin. The Echinacea-Rudbeckia complex is confined chiefly to the United States. The following theory as to their origin, based on their geographical distribution, seems plausible. This group of genera probably evolved from a similar group of plants that originated on the Appalachian-Ozarkian Upland, a region ex- posed since the close of the Paleozoie. A discussion of each genus, which follows, furnishes the basis for this assumption. Echinacea is perhaps the older of these genera. Five of its six species occur on the Appalachian-Ozarkian Upland. The two most primitive species, E. laevigata and E. purpurea, are confined to this region. Echinacea laevigata occurs along the uplands east of the Alleghenies from eastern Pennsylvania to Georgia. This species is uncommon within its range. Echi- nacea purpurea ranges from Georgia north and west to Ken- tucky, and enters the Ozarkian Upland by way of southern Illinois. These two latter species are 2 to 4 feet tall, and more or less branched with leafy stems. Echinacea paradoxa is confined to the Ozarkian Upland and the Cretaceous uplands of eastern Oklahoma and Texas. Echi- nacea pallida, a more recent species, occurs on these uplands, and has also migrated into the surrounding prairies of more recent geological history. This species has given rise to E. angustifolia on the dry plains to the west, extending from the Dakotas south to western Texas. These three species are more recent, due to (1) their adaptation to a drier climate of the prairies and plains, (2) their occurrence in a more recent territory, and (3) more hispid pubescence with smaller leaves and rather low, somewhat subscapose stems. The pronounced spiny pales of Echinacea, the corolla with a bulbous base, unlike that of any other genus, are all probably primitive characters. Rudbeckia and Echinacea have evolved from a similar an- cestor, but the former is of more recent origin. The area [Vor. 22 60 ANNALS OF THE MISSOURI BOTANICAL GARDEN of present development of the later species of Rudbeckia ex- tends along the coastal plain from Virginia to Texas. The center of distribution includes the Appalachian-Ozarkian Up- land and the coastal region from Virginia to Texas. The epappose Hirtae and Bicolores sections of this genus are of more recent development. The species of the former section are aggressive and quite nascent. The center of distribution of the columnar species of Rud- beckia extends from Alabama to the Ozarkian Upland, Texas, and westward, a few species reaching the Pacific coast. It is quite evident that Rudbeckia has evolved along two lines, one giving rise to the type with the columnar receptacle, the other to the type with the conical receptacles. The former has de- veloped from the western extension of this upland, and the latter from the Appalachian region. Dracopis has been evolved from somewhere along the columnar receptacle branch. The center of distribution of Ratibida is in the region of Nebraska, since its greatest development is from the Dakotas and Montana along the dry plains to Texas, New Mexico, and northern Mexico. This is perhaps the latest genus of the Echi- nacea-Rudbeckia complex. Since Dracopis and Ratibida have many characters in common, it is probable that they have had a common ancestor. Evidence in favor of the more recent origin of this genus is to be found in (1) the distribution in a more recent territory, geologically speaking, (2) the pinnate leaves, (3) the specialized seed dispersal, and (4) the aggressive and weedy nature of many of the species. GENERAL DISTRIBUTION The geographical distribution of the several genera com- prised in this study, except in the case of Zaluzania, is co- incident for the most part with certain botanical regions in the United States and in Mexico. Balsamorhiza, Echinacea, and Ratibida are native of the United States. The first-mentioned genus is confined to the northwestern states, whereas the other two genera are confined chiefly to the central and west-central states. The geographical range of Ratibida extends southwest- ward into northern Mexico. Jostephane, Greenmaniella, and 1935] SHARP——CERTAIN EPAPPOSE GENERA OF COMPOSITAE 61 Zaluzania are mainly of Mexican distribution. Two species of Zaluzama, however, are extralimital ; Z. Grayana, for example, reaches its northern limits in A Don. while Z. Sodiroi is known only from Ecuador, South America. A more detailed distribution of each genus is given in the taxonomie treatment of the respective groups. ABBREVIATIONS The following abbreviations are used for the various her- baria cited in the taxonomie portion of this study : F - Herbarium of the Field Museum of Natural History. G = Gray Herbarium of Harvard University. K - Herbarium of the Royal Botanie Gardens, Kew. M - Herbarium of the Missouri Botanical Garden. NY - Herbarium of the New York Botanical Garden. Ph - Herbarium of the Academy of Natural Sciences, Philadelphia. RM - Rocky Mountain Herbarium, University of Wyo- ming. U CAL - Herbarium of the University of California. US - United States National Herbarium. WU = Herbarium of Willamette University. Part II. Taxonomy KEY TO THE GENERA A. Ray-flowers without styles. B. Receptacle columnar, nd or eonie-eylindrie at maturit C. Receptacle columnar; bra BR the involuere biseriate, "é inner series one-half or less as long as uter . Mature achenes terete, ie iyd wrinkled; leaves simple..... às DD. Mature achenes Ade -rhomboidal or laterally flattened, smoot pin CC. Receta al or eonie-eylindrie; bracts of the involucre bi- to tri-seriate, of about equal length..................oo........ Rudbeckia BB. B le E. Achenes with the erowns slightly rounded, epappose or with a few minute caduco u UT EU AME. cree prets pu EE. Achenes Memori abruptly into a truncate, toothed cro F. Corolla expanded below into a bulb-like base, io apa above into the cylindrical eorolla-tube; pales with stout ne pi FF. Corolla tubular; pales Aeute......o....o.ooooooommm»os er [Vor. 22 62 ANNALS OF THE MISSOURI BOTANICAL GARDEN AA. Ray-flowers styliferous. G. Pales firm, subherbaceous to subcoriaceous; mature disk- posés linear-oblong, rhomboidal or quadrangular; epappos H. Base a the disk-corolla dilated and capping the achene; re- ceptacle conical; heads small to medium (disk 0.5-1 em. prisons usually in corymbose or corymbose-paniculate [AUT 171 AAA spare ibd e's 06 40456 oD Eee Zaluzania HH. Base of disk-corolla narrow; receptacle broadly convex; heads large (disk 2-4 em. broad), usually solitary...... e EE EETA ITELT TTT Balsamorhiza GG. Pales membranaceous with several brownish-resinous parallel nerves; mature disk-achenes subfusiform, subterete; pap- pus a shallow cup-like crown with knob-like projections at |^: O EEPE E E TLT T T Greenmaniella RATIBIDA Raf. History.—The name Ratibida was proposed by Rafinesque! and a brief description of it was published in his *Flora Ludo- vieiana’ in 1817. The genus was based on Rudbeckia colum- naris Pursh? of 1814. Pursh’s characterization is identical with that of Rudbeckia columnaris Sims? in ‘Curtis’ Botanical Magazine’ which appeared in 1813. Sims’ description is ac- companied by an excellent illustration, hence there can be no doubt as to the identity of the plant concerned. Furthermore, Rafinesque in his ‘Flora Ludoviciana’ states that ‘‘all the species with naked seeds as R. pinnata & must form my genus Obelisteca, and those with a simple perianthe, such as R. columnaris Pursh, my genus Ratibida. Raf." Moreover, in 1818, he* writes briefly of this genus; and in the following year he’ published a full description of it in the ‘Journal de Phys- ique’ where Rudbeckia columnaris Pursh is given as the name- carrying synonym. The name Rudbeckia columnifera Nutt.* was first published in ‘Fraser’s Catalogue’ in 1813. The description accompany- ing it is so indefinite that the name R. columnifera may be re- 1 Rafinesque, C. S., Florula Ludoviciana. p. 73. 1817. ? Pursh, F., FL in. Sept. 2: 575. 1814. * Sims, J., in Curt. Bot. Mag. 39: pl. 1601. dee * Rafinesque, C. S., in Am. Month. Mag. 268. 1818. * Rafinesque, C. S., in Jour. Phys. € Chem. M 100. 1819. * Nuttall, T., in Fraser's Cat. no. 75. 1813 [Reprint in Pittonia 2: 118. 1890]. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 63 garded as a nomen subnudum. Nuttall’ himself in the ‘Genera of North American Plants’ in 1818 abandoned his name R. columnif era and recognized Rudbeckia columnaris Pursh. The first description of Lepachys Raf. immediately follows that of Ratibida Raf. in the ‘Journal de Physique’ in 1819. It was based upon Rudbeckia pinnata Vent. It has since been re- garded as congeneric with Ratibida Raf. Therefore, accord- ing to the International Rules of Botanical Nomenclature, Ratibida becomes the valid generic name with Ratibida colum- naris as the type species. In 1825 Cassini,’ in ‘Dictionnaire des Sciences Naturelles,’ described the genus Obeliscaria which is based on Rudbeckia pinnata Vent. DeCandolle? in the *Prodromus' in 1836 recog- nized Cassini's genus and treated Ratibida and Lepachys as sections under Obeliscaria. Don,!? in Sweet's *British Flower Garden,” in 1838, was the first worker to recognize Ratibida. Barnhart,™ in 1897, was next to recognize the validity of this name, and Lepachys was rejected for reasons similar to those given above. The genus Lepachys, with its three species and one variety, was treated by Torrey and Gray’? in their ‘Flora of North America” in 1842. Rudbeckia Tagetes James was transferred to Lepachys Tagetes by Gray?? in ‘Pacific Railroad Report’ in 1856. The last species to be described was Lepachys mexicana Watson!* in ‘Proceedings of the American Academy’ in 1888. GENERAL MORPHOLOGY Roots.—The root system of Ratibida is of two types, namely, a vertical tap-root, and a fibrous or fleshy fibrous spreading root system. The first type is represented in its best develop- ment by R. Tagetes, where the roots are long, slender, vertical, " Nuttall, T., Gen. N. Am. Pl. 2: 178. 1818. ® Cassini, M. H., in Dict. Sei. Nat. 35: 272. 1825. * DeCandolle, A., Prodr. 5: 558. 1836. 12 Don, D., in Sweet’s Brit. Fl. Gard. II, 4: 361, pl. 361. 1838. ? Barnhart, J. H., in Bull. Torr. Bot. Club 24: 410. 1897. u Torrey, J. & Gray, A., Fl. N. Am. 2: 313. 1842. ? Gray, A., in Pacif. Rail. Rept. 4: 103. 1856. * Watson, S., in Proc. Am. Acad. 23: 277. 1888. [Vor. 22 64 ANNALS OF THE MISSOURI BOTANICAL GARDEN and rather deep-seated. A less-developed form of the first type is found in R. columnaris and R. peduncularis, which have a narrow, conical tap-root soon tapering to a slender tip. In these species, the roots are shallow, the main body being usu- ally less than 1 dm. in depth. Plants with this type of root are biennials or short-lived perennials. The more extensively de- veloped root system of R. Tagetes is of perennial duration. Ratibida pinnata and R. mexicana possess the shallow hori- zontally developed or rhizome-like root from which many stout fibrous secondary roots develop. These species are perennials. Stems.—The stems are herbaceous, erect, simple, or branched, often copiously so, giving the plant a compact bushy appearance, as in R. Tagetes, The leafy branches terminate in naked peduncles of varying lengths. The strongly sulcate character of the stem and branches, with regular light green ridges and dark green grooves, is a marked feature of nearly all species of this group. Leaves.—The leaves are alternate, the lowermost long-peti- olate and the uppermost short-petiolate to sessile. The blades are pinnate-cleft, pinnate to bipinnatifid. The basal leaves in R. mexicana are lanceolate, those of the stem 3—5-pinnate-cleft. The leaves of R. pinnata and R. mexicana are distinetly triple- nerved. This character is less pronounced in the other species. Pubescence.—The hirsute to strigose-hirsute type of pubes- cence is prevalent throughout the genus. The basal leaf- petioles and the lower parts of the stems are spreading-hirsute, exemplified by R. mexicana, R. peduncularis, and R. pinnata. The upper parts of the leafy stems, as well as the long naked peduneles, are strigose-hirsute, or less frequently strigose- hirtellous. The leaves are sometimes tuberculate-strigose- hirsute, as illustrated by R. Tagetes and R. pinnata. The pu- bescence of the leaves and stems is always interspersed with resinous glands or globules. On the stems this is more pro- nounced in the grooves, while the ridges are strigose-hirsute. Inflorescence.—The heads are solitary on long naked pe- duneles terminating the leafy branches. 1935] SHARP— CERTAIN EPAPPOSE GENERA OF COMPOSITAE 65 a. Involucre.—The involucral bracts are herbaceous and bi- seriate; those of the inner series are oblanceolate, scarcely longer than the pales or often somewhat shorter; those of the outer series are linear, reflexed or spreading, and from two to three times longer than the inner series. The involuere, while characteristic for the genus, presents few characters of value in specific differentiation. b. Receptacle and Pales.—'The narrow columnar type of re- ceptacle is characteristic of all the species of Ratibida. This varies in length somewhat within the genus, but reaches its maximum development in R, columnaris. The pales are densely erowded, more or less conduplicate. The apices are broadly apiculate-obtuse, with the tip incurved and densely velutinous, thus giving the heads a velvety aspect. A narrow fusiform resin duct along the edge of each pale is present in all the species except R. mexicana. The pales closely clasp the achenes so that upon drying after maturity they sep- arate from the receptacle and fall with the ripened achene. c. Corollas.—All the species of Ratibida have heterogamous heads with neutral ligulate florets and fertile disk-florets. The 9-lobed disk-corollas are inflated-tubular. The corollas of the ray-flowers are usually yellow, but the color varies in some species from yellow to bluish-purple. This is especially dem- onstrated in R. columnaris, in which the rays range from yel- low, blotched with bluish-purple at the base or to the middle, to entirely bluish-purple. A similar color variation is also common in R. peduncularis and its variety. The rays are broadly oblong-elliptical, rotund-ovate to oblong-lanceolate, 2-3-denticulate, strongly spreading or reflexed, often cleft to the top of the achene. d. Achenes.—The achenes of the disk-flowers in Ratibida are laterally compressed, with margins acute, or prolonged into a flattened edge, with margins distinetly pectinate-fimbriate, as in R. peduncularis, or fimbriate-ciliate, as in R. columnaris and R. Tagetes; the body of the achene is glabrous. Pappus.—Pappus none, or consisting of two reduced awns, or of a small erown bordered by minute teeth. [Vor. 22 66 ANNALS OF THE MISSOURI BOTANICAL GARDEN Geographical Distribution.—The genus Ratibida is confined chiefly to the central United States, its center of distribution lying in the region of Nebraska and Kansas. The majority of the species are inhabitants of dry soils, but the species are gen- erally quite adaptable and respond readily to a more favorable environment. The species R. columnaris, native of the region from the Dakotas and Montana south to Texas and Mexico, has migrated since the development of the railroads into territory east, west, and southwest of its former range. The more favorable en- vironment, especially in Missouri and Illinois, produces more vigorous plants than those growing in Texas, Oklahoma, etc. Ratibida pinnata is confined chiefly to the prairie states, The remaining species are of a southwestern distribution; R. Tagetes, an inhabitant of dry plains, prairies, and rocky hill- sides, ranges from Kansas and Colorado south to Texas and northern Mexico. Ratibida peduncularis and its variety picta inhabit the Gulf coastal region of Louisiana and Texas. Rati- bida mexicana, the only Mexican species, inhabits cool slopes of the Sierra Madre Mountains of Chihuahua and Durango. TAXONOMY Ratibida Raf. Fl. Ludovic. p. 73. 1817; in Jour. Phys. 89: 100. 1819; D. Don in Sweet's Brit. Fl. Gard. 4: 361, pl. 361. 1838; Barnh. in Bull. Torr. Bot. Club 24: 410. 1897; Britt. & Brown, Ill. Fl. ed. 1, 3: 419. 1898, and ed. 2, 3: 474. 1913; Small, Fl. Southeast. U. S. ed. 1, 1260. 1903, and ed. 2, 1260. 1913; Coult. & Nels. Bot. Rocky Mts. 545. 1909; Woot. & Standl. in Contr. U. S. Nat. Herb. [Fl. N. Mex.] 19: 705. 1915; Rydb. Fl. Rocky Mts. 927. 1917; Small, Man, Southeast. Fl. 1428. 1933. Lepachys Raf. in Jour. Phys. 89: 100. 1819; Cass. in Diet. Sci. Nat. 26: 3. 1823; Less. Syn. Gen. Comp. 225. 1832; Torr. & Gray, Fl. N. Am. 2: 313. 1842; Gray, Manual, 225. 1848; Benth. & Hook. Gen. Pl. 2: 366. 1873; Gray, Syn. Fl. N. Am. 1?: 263. 1884; Hoffm. in Engl. & Prantl, Nat. Pflanzenfam. IV. Abt. 5, 233. 1890; Coult. in Contr. U. S. Nat. Herb. [Bot. West. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 67 Texas] 2: 215. 1892; Rob. & Fern. in Gray's Manual, ed. 7, 832. 1908; Rydb. Fl. Pr. & Pl. Cent. N. Am, 838. 1932. Obeliscaria Cass. in Dict. Sci. Nat. 35: 272. 1825; DC. Prodr. 5: 558. 1836. Herbaceous biennials or perennials arising from horizontal or vertical rootstalks. Stem simple or branched, erect or oc- casionally horizontally spreading, sulcate, marked by alternat- ing light and dark green lines. Leaves alternate, pinnate, bi- pinnate, or lyrately pinnatifid, the basal sometimes lanceolate, serrate or entire, acute, strigose-hirsute, tuberculate-strigose- hirsute and interspersed with glandular punctae, especially on the lower surface. Heads radiate, solitary on naked peduncles, cylindrical, oblong-cylindrical to globose. Bracts of the in- voluere biseriate, herbaceous, linear to linear-lanceolate, the outer series twice or more the length of the inner. Receptacle columnar. Pales more or less conduplicate, obtusely apiculate to truneately apiculate, the tip incurved, a linear or elliptical brownish resin duct usually present along each margin, apex densely puberulent or velutinous, deciduous with the mature achenes. Disk-flowers with a very short tube, throat inflated- tubular, 5-lobed, glabrous; anthers sagittate at the base with ovate appendages; style-branches recurved or spreading, apex acute or rounded, densely puberulent to hirsutulous. Ray- flowers neutral, rays yellow or bluish-purple, 2-3-dentate, the corolla-tube short or obsolete. Achenes laterally flattened, eompressed-rhomboidal in cross-section, sometimes ciliate or pectinate-fimbriate along the edges, otherwise glabrous; pap- pus when present coroniform or prolonged upwards on each edge, forming two awn-like teeth. Type species: R. columnaris (Sims) D. Don in Sweet's Brit. Fl. Gard, 4: 361, pl. 361. 1838. KEY TO THE SPECIES A. Achenes strongly compressed or flattened, inner wing fimbriate; pappus consisting of two tooth-like projections or a lacerate crown. B. Heads globular or oblong; achenial crown yellowish, thickened, bordered by minute lacerate setae. oe. essea re EEE Edere T. 2. R. Tagetes BB. Heads cylindrical or columnar; pappus consisting of two persistent tooth- like projections. [Vor. 22 68 ANNALS OF THE MISSOURI BOTANICAL GARDEN C. Plants erect, leafy throughout..........................L R. columnaris CC. Plants ereet, horizontally spreading, or ascending, leafy je ter- minating in a long naked pedunele D. Leaves bipinnatifid; achenes niii fimbriate Pap both O era 3. R. peduncularis DD. Leaves lyrately pinnate; achenes fimbriate-pectinate along the inner O 3a. R. peduncularis var. picta AA. — compressed-rhomboidal, not winged; pappus ibt. eee leaves pinnate; pales with a linear brownish resin us along argin; plants branched above.................. . pinnata EE. Bon: pen ARA entire; pales without resin > plants Pig 1) Eee aaa 5. R. mexicana 1. Ratibida columnaris (Sims) D. Don in Sweet's Brit. Fl. Gard. 4: 361, pl. 361. 1838; Britt. & Brown, Ill. Fl. ed. 1,3: 419. 1898, and ed. 2, 3: 474, fig. 4454. 1913; Small, Fl. Southeast. U. S. ed. 1, 1260. 1903, and ed. 2, 1260. 1913; Coult. & Nels. Bot. Rocky Mts. 545. 1909; Small, Man, Southeast. Fl. 1428. 1933. Rudbeckia columnaris Sims in Curt. Bot. Mag. 39: pl. 1601. 1813; Pursh, Fl. Am. Sept. 2: 575. 1814; Nutt. Gen. N. Am. Pl. 2: 178. 1818; Maund, Bot. Gard. 4: pl. 84; n. 336. 1831; Hook. Fl. Bor.-Am. 1: 311. [1834] 1840; Hemsl. Biol. Cent.-Am. Bot. 2: 167. 1881. Rudbeckia columnifera Nutt. in Fraser’s Cat. n. 75. 1813. Ratibida sulcata Raf. in Jour. Phys. 89: 100. 1819. Obeliscaria columnaris DC. Prodr. 5: 559. 1836. Lepachys columnaris (Sims) Torr. & Gray, Fl. N. Am. 2: 315. 1849; Gray, Syn. Fl. N. Am. 1?: 264. 1884; Coult. in Contr. U, S. Nat. Herb. [Bot. West. Texas] 2: 215. 1892; Rob. & Fern. in Gray's Manual, ed. 7, 833. 1908. Ratibida columnifera (Nutt.) Woot. € Standl. in Contr. U. S. Nat. Herb. [Fl. N. Mex.] 19: 706. 1915; Rydb. Fl. Rocky Mts. 927. 1917. Lepachys columnifera (Nutt.) Macbr. in Contr. Gray Herb. No. 65: 45. 1922; Rydb. Fl. Pr. & Pl. Cent. N. Am. 838. 1932. Stems one or more, erect, 2.5-12 dm. tall, branched, closely strigose-hirsute throughout with occasional interspersed res- inous glands, sometimes hirtellous-strigose, lateral branches spreading; leaves including the petiole 3-15 em. long, closely strigose-hirsute on both surfaces, interspersed by numerous punetate glands, pinnately cleft to the midrib into 5-13 linear, 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 69 narrowly lanceolate, oblong or oblong-lanceolate divisions, these entire or sometimes trifid, acute, 0.5-3 em. long, 1-10 mm. broad; heads cylindrical, 1-5.5 em. long, 7-10 mm. broad ; outer braets of the involucre linear, 4-12 mm. long, acuminate, the inner about 3 mm. long, strigose-hirsute; ray-flowers 3-7, rays yellow throughout, broadly oblong-elliptieal, 1-2 em, long, 0.6— 1 em. broad, trifid, tube and outer surface of ray puberulent ; mature achenes compressed, 2 mm. long, glabrous, except the wing on the inner side, which is ciliate-fimbriate, pappus con- sisting of two tooth-like projections on each edge of the crown. Distribution: Illinois to Minnesota and Montana, south to Texas and Mexico. The following, selected from numerous specimens, are 4 repr e: ILLINOIS: railroad, Edwardsville, Madison Co., June 26, 1891, Eggert (M). MISSOURI: dry sandy ground, Hannibal, Marion Co., Aug. 7, 1912, Davis 5114 (M); railroad tracks, Winfield, Lincoln Co., June 7, 1916, Davis 1390 (M); com- mon, introduced, Sheffield, Jackson Co., June 26, 1894, Bush 157 (M); waste ground, er Jackson Co., July 7, 1894, Bush 163 (M). ARKANSAS: rocky open ground, Brentwood, Washington Co., July 7, 1915, Palmer 8216 t ). MINNESOTA: Brown Valley, Traverse Co., Sept., 1893, Sheldon (M); Glyndon, Clay Co., Sept., 1892, Dewart (M). Nor DAKOTA: Ryder, Ward Co., July 31, 1928, Larsen 187 (M). SovrH DAKOTA: Brookings, Brookings Co., Aug. 7, 1893, Thornber (M); Iri- quois, xn us Co., Aug. 11, 1897, Thornton (M); plains and ravines, ‘‘Bad Lands,’’ Cedar Pass, near Interior, Stanley Co., June 29, 1929, ph Ge 37528 (M); hillsides, Deadwood, Lawrence Co., July 13, 1913, Carr 93 (M). NEBRASKA: railroad tracks, Haré Lancaster Co., July 13, 1915, Davis 7320 (M); field, 20 mi. west of Hastings, Adams Co., June 17, 1929, Mathias 309 (M); Kearney, Buffalo Co., June 27, 1899, Pammel (M); Orella, Sioux Co., July 6, 1912, Pool $ Folsom (M). KANSAS: prairies near Spearville, Ford Co., Aug. 24, 1902, Eggert (M); Grin- nell, Gove Co., July 15, 1872, Redfield 485 (3503) (M) ; common, near Ulysses, Grant sha Co., Jos 30, 1926, Heller 13975 (M); Paola, ru Co., Aug. 20, 1884, Oyster 4149 (M); east of Coffeyville, Montgomery Co., e 28, 1929, Rydberg $ Imler 366 (M); prairie, Riee Co., Aug. 23, 1895, xL 267 (M); Manhattan, Riley Co., June, 1891, Ellis (M). OKLAHOMA: sandy soil, 17 mi. n. Boise City, Cimarron Co., Aug. 22, 1927, Stratton 444 (M) ; Norman, T2 10, 1914, Emig 366 (M); Lawton, Comanche Co., , 1903, Duncan (M); abundant in prairies, Sapulpa, Creek Co., July 22, 1894, na. 307 (M); 615 mi. ^M p Buffalo, Harper Co., Aug. 20, 1927, Stratton 399 (Vor. 22 70 ANNALS OF THE MISSOURI BOTANICAL GARDEN (M); Arbuckle Mts., near Davis, Murray Co., June 21, 1917, Emig 772 (M); Muscogee, Muscogee Co., July, 1880, Letterman (M); prairies near Cora, Woods Co., May 28, 1913, Stevens 764 (M Texas: Claude, Armstrong Co., June 25, 1902, Reverchon 493 (M); common on ar San — Oct. 2, 1900, Bush 1234 (M); Ridge Springs, 9 mi. south of rathon, Brewster = ‚July, 1921, Ferris $ Duncan 2838 (M); foothills below MeKittriek rdi 1650 m., Guadalupe Mts., Culberson Co., July 23, 1931, Moore $ Steyer lur 2 (M); prairies, Houston, Harris Co., May 24, 1915, Fave 7746 (M); prairies near Canadian, Hemphill Co., Aug. 10, 1900, Eggert (M); rocky open ground in canyon, Davis Mts., Jeff Davis Co., June 17, 1926, Palmer 31021 (M); on prairies, Terrell, Kaufman Co., Oct, 22, 1902, Reverchon 3337 (M); near Kerrville, Kerr Co., alt. 1600-2000 ft., June 12-19, 1894, Heller 1850 (M); Lubbock, Lubbock Co., May 19, 1931, Demaree 7706 (M); along rail- way near Amarillo, Potter - , July 13, 1917, Palmer 12535 (M); dry open plains, Canyon, Randall Co., June 3, 1918, Palmer 13878 (M); sandy woods, Troupe, Smith Co., May 9, 1909, rail ats 3338 (M); Abilene, Taylor Co., May 22, 1902, Tracy 7891 (M) ; Del Rio, Valverde Co., April 17, 1930, Jones 25903 (M). MoNTANA: dry ground, Bozeman, Gallatin Co., July 26, 1905, Blankinship 283 (M); enge alt. 3000 ft., Aug. 2, 1909, M. E. Jones (M). YO : Mammoth Hot Springs, Aug. 22, 1892, Mulford (M); waste ground, lieti pedes Co., alt. 7300 ft., July 27, 1929, Goodman 226 (M). COLORADO: near Boulder, alt. 5700 ft., July 8, 1920, Hanson C139 (M); prairie near Denver, Denver Co., alt. 5300 ft., July 14, 1917, Clokey 2819 (M); Manitou, El Paso Co., Aug., 1884, Letterman (M); near Cañon City, Fremont Co., July 8, 1873, Brandegee 757 (M); Fort Collins, Larimer Co., alt. 5000 ft., July 9, 1896, Baker (M); near Pueblo, Pueblo Co., June 29, 1929, Mathias 526 (M); Ft. Lupton, June 29, 1913, Johnston 217 (M); Wray, Yuma Co., July, 1919, Eggleston (M). New Mexico: White Mountains, Lincoln Co., alt. 6700 ft., July 31, 1897, Wooton 261 (M); Nara Visa, Quay Co., Aug. 1, 1911, Fisher (M); near Pecos, San Miguel Co., Aug. 20, 1908, Standley 5124 (M); north of Glorieta, Aug. 24, 1908, Standley 6279 (M). MExico: coAHUILA—Chojo Grande, 27 miles southeast of Saltillo, Aug. 29, 1904, Palmer 345 (M) ; CHIHUAHUA—near Colonia Garcia in the Sierra Madres, alt. 750 ft., Aug. 5, 1899, Townsend $ Barber 230 (M); SAN LUIS POTOSI—Nov., 1910, Peu 4768 (M); NUEVO LEON—near Monterey, alt. 550 m., July, 1911, Arsène 2608 (M). 1a. forma pulcherrima (DC.) W. M. Sharp, comb. nov. Obeliscaria pulcherrima DC. Prodr. 5: 559. 1836. Ratibida columnaris var. pulcherrima (DC.) D. Don in Sweet's Brit. Fl. Gard. II. 4: 361, pl. 361. 1838; Coult. € Nels. Bot. Rocky Mts. 545. 1909. Lepachys columnaris var. pulcherrima (DC.) Torr. & Gray, Fl. N. Am. 2: 315. 1842; Gray, Syn. Fl. N. Am. 1?: 264. 1884; Rob, & Fern. in Gray's Manual, ed. 7, 833. 1908. 1935] SHARP-——CERTAIN EPAPPOSE GENERA OF COMPOSITAE 71 Ratibida columnifera var. pulcherrima (DC.) Woot. & Standl. in Contr. U. S. Nat. Herb. [Fl. N. Mex.] 19: 706. 1915. Habit, foliage, pubescence, ete. identical with the species; ray-flowers bluish-purple, or bluish-purple blotehed at the base. Distribution: same as the species, The following, selected from numerous specimens, are repr tative: £ MISSOURI: waste ground, Sheffield, Jackson Co., June 17, 1918, Bush 8407 (M). ARKANSAS: open ground, Fulton, Hempstead Co., June 17, 1915, Palmer 8024 (M); railroad near Mandeville, Miller Co., June 10, 1898, Eggert (M). NorTH DAKOTA: Ryder, Ward Co., July 31, 1928, Larsen 194 (M). SouTH DAKOTA: Triquois, Kingsbury Co., Aug. 1, 1892, E (M). NEBRASKA: sand hills, Cherry Co., 1892, Smith $ Pound 145 (M); Lincoln, Lan- caster Co. , July 11, 1899, Hedgcock (M); common in prairies, Sioux Co., July-Aug., 1927, Kramer 60 (M). Kansas: Ellis Co., June 26, 1885, Kellerman (M); New Ulysses, Grant Co., jos 24, 1896, pov (M). LAHOMA: Frederick, Tillman Co., July 5, 1903, vhs 28 (M). EXAS: Bryan, Brazos Co., May 27, 1915, Palmer 7792 (M); vicinity of Browns- ville, Cameron, Co., Aug. 1-5, 1921, Ferris $ Duncan bg (M); railroad near Long- view, Gregg Co., June 7, 1899, Eggert (M); near Lubbock, Lubbock Co., May, 1930, Demaree 77064 (M); Corpus Christi, Nueces Co., May, 1913, Orcutt 5821 M); sandy woods, Troupe, Smith Co., May 9, E. digi 3338 in part (M); Austin, Travis Co., Aug., 1882, Kellerman (M) ; Del Rio, April 20, 1930, T 25904 (M). Wyoming: Willow Creek, July 20, iu Nelson 570 (M); Manville, Niobrara Co., July 18, 1901, Nelson (M). COLORADO: Fort Collins, Larimer Co., alt. 5000 ft. ‚July 27, 1892, Crandall (M); Cheyenne Cañon, Colorado Springs, El Bodo Co. e. 24, 1915, Drushel 4191 (M); Denver, Denver Co., Aug. 5, 1885, Fritchey 167 (M); Basen Springs, Archuleta Co., Aug. 13, 1917, Papiol 1164 (M). New Mzxico: valley of the Pecos, Aug. 13, 1847, Fendler 387 (M); Nara Visa, Ouray Co., Sept. 19, 1910, Fisher (M); Koglin Mts., on the middle fork of the Gila ve, Socorro Co., Aug. 8, 1903, Metcalfe 415 (M) ; Carson Forest, Rio Arriba Co., 2100 m., Aug. 14, 1924, Eggleston 20562 (M); White Mts., Lincoln Co., alt. 6500 ft., Fu 30, 1897, Wooton 247 (M); near Pecos, alt. 6700 ft., Lincoln Co., Aug. 15, t Standley 4925 (M). XICO: NEUVO LEON—Carrizitos, May 28, 1847, Gregg 810 (M); COAHUILA— Matamoros, pei 12, 1906, Tracy 8947a (M). 2. Ratibida Tagetes (James) Barnh. in Bull. Torr. Bot. Club 24: 410. 1897; Britt. & Brown, Ill. Fl. ed. 1, 3: 419. 1898, and ed. 2, 3: 475. 1913; Small, Fl. Southeast. U. S. ed, 1, 1260. 1903, and ed. 2, 1260. 1913; Woot. & Standl. in Contr. U. S. Nat. [Vor. 22 12 ANNALS OF THE MISSOURI BOTANICAL GARDEN Herb. [Fl. N. Mex.] 19: 706. 1915; Rydb. Fl. Rocky Mts. 927. 1917. Rudbeckia Tagetes James in Long’s Exp. [Phila. ed.] 2: 68. 1823. Obeliscaria Tagetes (James) DC. Prodr. 5: 559. 1836. Lepachys columnaris var. Tagetes (James) Gray in Smith- son. Inst. Contr. [Pl. Wright. pt. 1] 3: 106. 1852. Lepachys Tagetes (James) Gray in Pacif. R. R. Rept. 4: 103. 1856; Gray, Syn. Fl. N. Am. 1?: 264. 1884; Coult. in Contr. U. S. Nat. Herb. [Bot. West. Texas] 2: 215. 1892; Rydb. Fl. Pr. & Pl. Cent. N. Am. 838. 1932. Stems slender, erect, the branches numerous, ascending, often giving the plant a bushy appearance, 1.54 dm. tall, stri- gose-hirsute, interspersed by numerous sessile glands espe- cially in the grooves; leaves strigose-hirsute or tuberculate- strigose-hirsute on both surfaces, interspersed by numerous dark, resinous or punctate glands; basal and lower cauline leaves lanceolate, entire, or more often pinnately or bipinnately 3-5-eleft, 5-13 em. long including the petiole, upper cauline usually 3-5-eleft into linear or linear-lanceolate segments, these 0.5-3 em. long, 1-3 mm. broad, entire, acute; heads on peduncles 2-7 em. long, numerous, often forming close clusters, globular to oblong, 0.8-1.5 cm. long, 8-10 mm. broad; ray-flowers 9-1, rays rotund-ovate to oblong-ovate, 4-6 mm. long, 3-5 mm. broad, trifid, reflexed, yellowish to bluish-purple, pubescent on the outer surface of the lamina; achenes ovate-oblong, 2-2.5 mm. long, winged on the inner edge, truncate, terminating ina thiekened, usually yellowish erown which is bordered by mi- nute laeerate setae, achenial body glabrous except along the fimbriate-ciliate wing. Distribution: dry plains, prairies, and rocky hillsides, Kan- sas and Colorado, southward to Texas and northern Mexico. KANSAS: Tribune, July 28, 1892, Reed (M); Johnson, Stanton Co., July 1, 1893, Thompson (M); prairie, Russell Co., July 13, 1895, Hitchcock 268 (M); roadside, 7 mi. west of Meade, July 10, 1929, Rydberg & Imler 807 (M); Wallace, Aug. 22, 1884, Letterman (M). OKLAHOMA: on gypsum hill near Hollis, Harmon Co., June 21, 1913, Stevens 1080 (M). v^ KR MER ir AT De VI P us. " Y sx 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 73 TEXAS: dry ground, Amarillo, Potter Co., July 13, 1917, Palmer 12534 (M); Teeline, Aug. 26-28, 1903, Griffiths 5641 (M); Tascosa, June 24, 1902, Reverchon $340 (M); Sierra Blanea, July 3, 1895, Mulford 260 (M); Amarillo, May 29, 1902, Reverchon (M); dry open ground along Rio Grande, near El Paso, El Paso Co., June 19, 1926, Palmer 31100 (M) ; dry open plains, Channing, Hartley Co., June 18, 1918, Palmer 14142 (M); near Lubbock, 1930, Demaree 75284 (M); Rio Grande Valley at Canutillo, El Paso Co., July 7, 1911, Barlow (M); near El Paso, 1849, did 52? a OLO tala Gunnison Co., July 29, 1901, Baker 667 (M); plains, July 4, 1901, uh (M) ; plains east pe pits, July 25, 1874, Engelmann (M). New MExico: near Pecos, alt. 6700 ft., Aug. 15, 1908, Standley 4950 (M) ; Mesilla Valley, Dona Ana Co., June 19, De Standley (M); vicinity of Las Vegas, San i 45 ray, Lin Co., alt. 6000-6500 ft., July, 1900, Earle $ Earle 150 (M); Santa Fe, alt. 7200 ft., July 8, 1897, Heller 3829 (M) ; Santa Fe, July 14, 1880, Rusby 78 (M) ; near Gray, Lincoln es . alt. 6000 ft., July 12, 1898, Skehan 32 (M); Las Cruces, Dona Ana Co., alt. 00 ft., June 15, 1897, Wooton 5 (M); San Miguel Co., 1897, Brandegee 12051 ped Sandia ia Mts, Allen’s Ranch, alt. 7200 ft., Aug.-Sept., 1914, Ellis 208 (M); waste fields, Santa Fe dnd eei 1847, Fendler (M); vieinity of Santa Fe, Galisteo, alt. 1865 m. ee 1926, Arséne $ Benedict 15841 (M) ; Archer Ranch, "cmt Dona Ana Co., Mer 1926, Child 5?3 (M mmon in sandy soil, * Charius,"? July 20, 1846, Wislizenus 25 (M); Mesilla Valley, Dona Ana Co., July 16, 1907, Wooton § Standley $315 (M). Mexico: State of Chihuahua, Valley of the Rio Grande, Paso del Norte, Sept. 14, 1886, Pringle 1061 (M) ; Juarez, June 27, 1891, Evans (M). 2a. var. cinerea Standley in Muhlenbergia 5: 30. 1909. Habit similar to the species; stem and peduncles cinereous, densely matted tomentulose-hirsute; leaves grayish, densely hirsute-strigose, the segments of the upper leaves about 2 mm. road. Distribution: dry hillside near Pecos, San Miguel Co., New Mexico. New Mexico: near Pecos, alt. 6700 ft., Aug. 21, 1908, Standley 5156 (M), ISOTYPE, 3. Ratibida peduncularis ( T. & G.) Barnh. in Bull. Torr. Bot. Club 24: 411. ek N Fl. Southeast. U. S. ed. 1, 1260. 1903, and ed. 2, 1260. 1913. Lepachys Torr. & Gray, Fl. N. Am. 2: 315. 1842; Gray, Syn. Fl. N. Am. 1?: 264. 1884; Coult. in Contr. U. S. Nat. Herb. [Bot. West. Texas] 2: 216. 1892. Stems erect or horizontally spreading, leafy below, terminat- [Vor. 22 74 ANNALS OF THE MISSOURI BOTANICAL GARDEN ing in a slender naked peduncle, 1-5 dm. long, spreading-hir- sute below, strigose-hirsute to strigose-hirsutulous on the pe- duncles; leaves more or less bipinnatifid; the blade and petiole 2.5-15 em. long, strigose-hirsute on both surfaces, interspersed with resinous glands below, segments 5-9, 0.5-5 em. long, the bipinnate lobes 1-4 mm. broad, acute, the terminal segment sometimes broadly pinnately cleft; heads solitary, columnar, 1.54 em. long, 8-11 mm. broad; the outer bracts of the invo- lucre linear, 6-10 mm. long, acute, hirsutulous, the inner series about 2 mm. long; rays 0.5-2.5 em. long, reflexed, yellow to blu- ish-purple, especially towards the base; achenes flat, 4 mm. long including the awn-like projections, the edges winged and conspicuously pectinate-fimbriate, otherwise glabrous. Distribution: Gulf coastal region of Louisiana and Texas. LOUISIANA: Cameron, July 4, 1903, Tracy 8548 (G, M); Cameron Parish, July, 1903, Cocks (G). TEXAS: near Corpus Christi, near sea-level, Nueces Co., May 28-31, 1894, Heller 1789 (G, M); north of Corpus Christi, March 30, 1932, Jones 29506 (M) ; Beeville, March 30, 1932, Jones 29507 (M) ; ‘‘ Felipe de Austin,’’ 1835, Drummond 108 (G); Hort. Cantab. anno 1848, Wright (G). 3a. var. picta (Gray) W. M. Sharp, comb. nov. Lepachys peduncularis var. picta Gray in Smithson. Inst. Contr. [Pl. Wright. pt. 1] 3: 107. 1852; Syn. Fl. N. Am, 1*: 264. 1884 ; Coult. in Contr. U. S. Nat. Herb. [Bot. West, Texas] 2: 216. 1892. Lepachys serratus Buckl. in Proc. Acad. Sci. Phila. 18: 457. 1861. Ratibida picta Small, Fl. Southeast. U. S. ed. 1, 1260. 1903, and ed. 2, 1260. 1913. Leaves lyrately pinnate to pinnately parted, coarsely serrate to ineised-serrate ; achenes pronouncedly winged and pectinate- fimbriate along the inner edge, outer edge entire or coarsely ciliate; other characters as in the species. Distribution: sandy woodlands of the Texas coastal region. Texas: Tyler, Tyler Co., Sept. 20, Reverchon 2229 (M); sandy woods, Hemp- stead, Waller Co., June 10, 1872, Hall 336 (G, M); gulf coast, 1849, Wright (G); Hort. Cantab., 1848, Wright (G); oak woodlands, 30 mi. west of San Antonio, Sept. 18-19, 1879, Palmer 718 (G) ; Victoria, April 18, 1905, Tracy 8914 (G, M). 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 195 4. Ratibida pinnata (Vent.) Barnh. in Bull. Torr. Bot. Club 24: 410. 1897; Britt. & Brown, Ill. Fl. ed. 1, 3: 418. 1898, and ed. 2, 8: 474. 1913; Small, Fl. Southeast. U. S. ed. 1, 1260. 1903, and ed. 2, 1260. 1913; Small, Man. Southeast. Fl. 1428. 1933. Rudbeckia pinnata, Vent, Desc. Pl. Nouv. Jard. Cels. 71, pl. 71. 1800; Sims in Curt. Bot. Mag. 49: pl. 2310. 1822. Lepachys pinnatifida Raf. in Jour. Phys. 89: 100. 1819. Obeliscaria pinnata (Vent.) Cass. in Diet. Sci. Nat, 35: 272. 1825; 46: 401. 1827; DC. Prodr. 5: 558. 1836. Lepachys pinnata (Vent.) Torr. & Gray, Fl. N. Am. 2: 314. 1842; Gray, Manual, 225. 1848; Gray, Syn. Fl. N. Am. 1?: 263. 1884; Rob. & Fern. in Gray's Manual, ed. 7, 833. 1908; Rydb. Fl. Pr. & Pl. Cent. N. Am. 838. 1932. Stems one or more from a horizontal rootstalk, stoutish, branched above, 4—12 dm. tall, spreading-hirsute below, stri- gose-hirsute to strigose-hirtellous above, intermixed with glandular globules; leaves strigose-hirsute to tuberculate- strigose-hirsute on both surfaces, interspersed with resinous glands; the basal long-petioled, pinnate, 1.5-3.7 dm. long, 3-7-pinnate, the upper short-petiolate, pinnate, 0.6-3 dm. long, leaf-segments 3-9, these linear-lanceolate to broadly lanceolate, triple-nerved, 2-9 cm. long, 0.4-2.5 cm. broad, sparsely serrate to entire, acute; heads oblong-cylindrieal, 1.2— 2.2 em. long, 1-1.5 em. broad; ray-flowers 5-10, rays oblong- lanceolate, 2-6 em. long, 0.6-1.2 em. broad, 2-3-dentate, puber- ulent on the outer surface; achenes 2-3 mm. long, glabrous; pappus absent. Distribution: dry prairies and rocky glades, etc. Ontario to Georgia, west to Minnesota and Oklahoma. The following, selected from numerous specimens, are representative: ONTARIO: Windsor, July 27, 1901, Macoun 34829 (M). GEORGIA: high sandy bank of Flint River, Sumter Co., July 11, 1901, Harper 1051 iu prairie, Adams Co., Sept. 15, 1930, Stephenson (M). KENTUCKY: Bowling Green, Warren Co., July 6, 1900, Price (M). TENNESSEE: Columbia, Maury Co., 1891, Shimek (M); railroad near Dechard, Franklin Co., July 6, 1898, Eggert (M). [Vor. 22 16 ANNALS OF THE MISSOURI BOTANICAL GARDEN INDIANA: bank of the Wabash River, 14 mi, east of Bluffton, Wells Co., July 31, 1906, Deam 1370 (M WISCONSIN: near Calla Beulah, Walworth Co., Aug. 2, 1905, Shannon 133a (M). ILLINOIS: near Tracy, Cook Co., Aug. 22, 1908, Greenman 3624 (M); prairies near Oquawka, Henderson Co., Patterson (M); 7 mi. west of Kankakee, Kankakee Co., Aug. 3, 1912, Sherff 1601 (M); prairie soil, Wabash, 6 mi. west of Decatur, Macon Co., July 12, 1915, Clokey 2406 (M) ; dry rocky open ground, Goleonda, Pope Co., Oct. 8, 1919, Palmer 17003 (M); prairies, Winnebago Co., Aug., 1859, Bebb 8508 (M). MINNESOTA: Lake Shetek, Murray Co., July, 1922, aed 1896 (M). Iowa: prairies, Armstrong, Emmet Co. . Aug. 20, 1897, Cratty (M); Ames, Story Co., July 18, 1896, jum. mel $ Ball 56 (M) ; common on prairies, near Dayton, Webster Co., Aug. 21, 1926, Pammel 17 (M); dry prairie soil, Decatur Co., Aug. 17, 1904, v e gs (M); high prairie, Black Hawk Co., July 17, 1929, Burk 556 (M). Mis : Shannon Co., July 22, 1891, Bush (M); near Independence, Jackson 2 Joly 8 o, 1894, Bush 164 (M); dry open ground, Van Buren, Carter Co., July 6, 1914 Palmer 6204 (M); in dry soil on rocky slope between Festus and Pes; Jef- ferson Co., June 20, 1929, ne... 1036 (M); near Allenton, St. Louis Co., July 20, 1893, Lotterie (M); r Mark Twain Cave, ae Marion Co., July 25, 1918, Davis (M) ; near Beglerock, June 27, 1897, Bush 1 ‘ ARKANSAS: dry open ground, near Fulton, ose Co. June 17, 1915, Palmer 8037 (M); dry rocky hillside near Cotter, Baxter Co., June 16, 1914, Palmer 6003 (M); Eureka Springs, Carroll Co., July 17, 1898, Glatfelter (M). KANSAS: prairies, Shawnee Co., July 4, 1895, Norton $ Clothier 266 (M); Johnson Co., 1890, Pellet (M). OKLAHOMA: in open shrubby woods, near Miami, Ottawa Co., Aug. 26, 1913, Stevens 2283 (M). 5. Ratibida mexicana (Wats.) W. M. Sharp, comb. nov. Lepachys mexicana Watson in Proc. Am. Acad. 23: 277. 1888. Stems one or more, unbranched, slender, 4—7 dm. tall, spread- ing-hirsute above; basal leaves lanceolate, 0.5-2.2 dm, long in- cluding the petiole, 1-3.5 cm. broad, serrate or serrate-crenu- late, acute, attenuate at the base, hirsute on both surfaces, more densely so beneath, intermixed with globular, resinous glands; cauline leaves 3-5 pinnate-cleft or parted, 6-16 em. long, the terminal lobe lanceolate, about one-half the length of the entire leaf, the lateral segments 1-3 cm. long, acute, sparsely serrate, surface as in the basal leaves; heads solitary on naked pe- duncles 1.5-5 dm. long, columnar, 1.8-2.5 cm. long, 1-1.3 cm. broad; pales without the brownish resin ducts; ray-flowers 8-15, rays yellow, oblong-lanceolate, 2-2.5 cm. long, 5-7 mm. broad, spreading, tube short, about 1 mm, long, pubescent on 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE TT the limb and outer surface; achenes about 3—4 mm. long, glab- rous, pappus absent. Distribution: cool slopes of the Sierra Madre Mts., Mexico. MEXI AHUA—cool slopes of the Sierra Madre, alt. 7000-9500 ft., Sept. 28, ine nin 1648 (M); cool slopes of the Sierra Madre, Sept. 19, 1887, Pringle 1305 (NY); Sierra Madre near Colonia Garcia, alt. 8000 ft., July 5, 1899, Townsend $ Barber 101 (M, NY); DURANGO—Sierra Madre, 30 miles north of Guanacevi, 8000—9000 ft., PE 18, aig x 4777 (NY). IosrEPHANE Benth. History.—The genus Iostephane was first described by Bentham! in “Genera Plantarum! in 1873. The species Echinacea heterophylla D. Don? was referred to and therefore must be interpreted as the type of the genus. Excellent illus- trations of this species may be seen in the *British Flower Garden’ and also in Knowles € Westeott’s ‘Floral Cabinet,’ under E. dubia. Hemsley, in ‘Biologia Centrali-Americana, * recognized Jostephane and added a new species, namely, 7. tri- lobata Hemsl. Hoffmann? in Engler and Prantl's *Die Natürlichen Pflan- zenfamilien,' in 1890, briefly characterized the genus and re- lated it to Rudbeckia and other allied genera which possess sterile ray-flowers and achenes without a pappus. At this time two species were recognized. GENERAL MORPHOLOGY Roots.—The root system consists of a fleshy spindle-shaped tap-root, The outer covering of the true tap-root is roughened by corrugated ridges of a bark-like structure, giving the root a woody appearance, but the interior tissue is fleshy. Its appear- ance, size, and structure suggest the tap-root of Balsamorhiza. Stems.—The thickened tap-root gives rise to a slender, woody, stem-like body 1—4 inches long. The herbaceous stems arise from buds at the sides, just below the surface of the ground. Alternate scars along this narrowed underground 1 Bentham, G., in Benth. & Hook. Gen. Pl. 2: 368. 1873. 2 Don, D., in Sweet's Brit. Fl. Gard. IT, 1: pl. 32. 1831. * Knowles, G. B. & Westeott, F., Fl. Cab. 3: 163. 1839. * Hemsley, W. B., Biol. Cent.-Am. Bot. 2: 168. 1881. * Hoffmann, O., in Engler € Prantl, Nat. Pflanzenfam. IV. Abt. 5. 233. 1890. (Vor. 22 78 ANNALS OF THE MISSOURI BOTANICAL GARDEN structure seem to indicate that it is an erect, perennial, under- ground stem or rootstalk which has developed from the crown of the tap-root. The stems above ground are herbaceous, erect, semiscapose, and usually unbranched. The taller plants, as I. heterophylla var. Dicksonüi, are branched. Leaves.—The basal leaves are long-petiolate and very nu- merous; those of the stem are much reduced. Sometimes only a few bracts are present, and occasionally in J. heterophylla and I. trilobata the stem is destitute of leaves or bracts. The basal leaves in all species are either pandurate, lobed, or less often ovate, the pandurate outline being the most common type. The bases of the leaf blades are decurrent along the sides of the petioles, The venation is conspicuous in that one or sometimes two prominent veins branch off from the midvein in the broad- ened basal portion of the leaf, and above and below this point the venation is less prominent. The peculiar triple venation and the decurrent blades are characteristic for all species of the genus. The stem-leaves are usually bracteiform. The low- ermost are sometimes leaf-like, and of a similar outline to the basal leaves, but they are always much reduced in size. Pubescence.—The stem bears a strigose pubescence which is less abundant on the lower parts and gradually increases above. The peduncles are often densely pubescent just below the heads, especially so in 7. heterophylla and I. trilobata. The stem in I. heterophylla var. acutiloba bears stout tuberculate- hispid hairs. The leaves are usually clothed by a tuberculate- hispid pubescence. Some specimens of Jostephane trilobata not only bear the above type of covering, but also hirsute trichomes. Involucre.—The involucral bracts are usually in two, some- times three, series. In the subgenus Euiostephane the outer bracts are longer than the innermost, often extending beyond the pales and the disk-flowers. In Chrysopetala the bracts are short, never extending beyond the disk-flowers. They are quite helianthoid, thus giving the head a very characteristic appear- ance. The bracts are foliaceous, lanceolate, or ovate-acumi- nate, the pubescence being similar to that of the foliage. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 79 Receptacle.—Two types of receptacle occur within the genus, namely, (1) the Echinacea-Euiostephane type, and (2) the helianthoid type. The receptacle in Euiostephane is strongly eonieal and about 8 mm. high, in this respect being similar to Echinacea. The pales are stout, echinulate, and terminate in a sharp straight or incurved beak. The receptacle in Chryso- petala is little more than the expanded apex of the peduncle. The pales are weak, subeoriaceous, and sharply serrated towards the apex. Ray-flowers.—The rays are neutral, with the achenes tri- angular in cross-section instead of rhomboidal as in the disk- flowers, The rays in Euiostephane are purple, oblong, and bidentate. Those of Chrysopetala, as the name signifies, are yellow and obovate. They are short, 7-12 mm. long, whereas those in the former subgenus are over 20 mm. long. Disk-flowers.—The corolla of the disk-flowers is a typical helianthoid form, as contrasted with that of the Rudbeckia and Echinacea types. The tube is constricted into a narrow cylinder, extending about one-fourth the entire length of the corolla, then it expands abruptly into the upper cylindrical throat and 5-lobed limb, similar to that of Helianthus. Geographical Distribution.—lostephane is confined chiefly tocentral Mexico. The general range extends from southwest- ern Chihuahua southwards along the Mexican Plateau, espe- cially at high elevations, to central and eastern Chiapas. The subgenus Chrysopetala occupies the extreme southeastern part of this range, with its northwestern limits in southeastern Puebla. Euiostephane extends from southern Chihuahua south to southern Puebla. Jostephane heterophylla is confined to south-central Mexico, ranging from southwestern San Luis Potosi south to Puebla, Morelos, and Michoacan. Its variety Dicksonii occurs from Chihuahua south to Puebla; whereas the variety acutiloba is known only from central Jalisco. The species of this genus, so far as available records show, occur in altitudes above 5500 feet, the average elevation being from 6000 to 8000 feet above sea-level. One collection of 7. tri- [Vor. 22 80 ANNALS OF THE MISSOURI BOTANICAL GARDEN lobata taken in the vieinity of Oaxaca was collected at an alti- tude of 9500 feet. Iostephane is chiefly confined to the oak associations. The species occur on dry rocky hills in the open oak woodlands or among the denser stands of oaks. In this respect the habitat resembles that of certain species of Echinacea. TAXONOMY Iostephane Benth. in Benth. & Hook. Gen. Pl. 2: 368. 1873; Hemsl. Biol. Cent.-Am. Bot. 2: 168. 1881; Hoffm. in Engler & Prantl, Nat. Pflanzenfam. IV, Abt. 5. 233. 1890. Herbaceous, semiscapose or leafy-stemmed perennials. Stems erect, from a thickened vertical rootstalk, simple or branched. Leaves alternate, radical or cauline, panduriform, trilobed-pandurate, ovate or broadly oblanceolate. Heads solitary, radiate, sometimes disposed in an open few-headed panicle, Involucre campanulate to hemispherical, bracts 2-3- seriate, herbaceous, lanceolate to ovate-oblong, acuminate, en- tire. Receptacle conical to slightly convex, paleaceous; pales coriaceous, entire to serrate towards the apex, broadly con- duplicate, acute, apex straight or incurved. Ray-flowers sterile, disposed in a single row; rays purple or yellow. Disk- flowers perfect; corolla-tube constricted, expanded above into a broadened cylindrical throat and 5-lobed limb; stamens 5, anthers sagittate, the apical appendages ovate. Stigmatic branches acute or acuminate, reflexed or curled, pubescent on the outer surface. Achenes of the ray-flowers sterile; those of the disk fertile, rhomboidal to obeompressed, obtuse, glabrous or pubescent; pappus usually absent, or if present of a few minute paleaceous setae. Type species: /ostephane ier ophylla (Cav.) Benth. in Benth. & Hook. Gen. Pl. 2: 368. 1873. Subgenus I. Euiostephane W. M. Sharp, new subgenus. Involuere hemispherical, 2-3 em. in diameter, 15 mm. high; receptacle conical, 5-9 mm. high, 3-6 mm. broad at base; pales rigid, echinulate, entire, acute, stout at the apex; rays purple; disk-achenes oblong-obovate, rhomboidal, cinereous or brown- 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 81 ish strigose-pubescent; pappus usually absent, when present of minute paleaceous setae. Subgenus II. Chrysopetala W. M. Sharp, new subgenus. Involuere campanulate, 1-1.5 em. in diameter, 6-10 mm. high ; receptacle convex, 1-1.5 mm. high, 1-1.5 mm. broad; pales weak, cartilaginous, subechinulate, serrate; rays yellow; disk- achenes oblong-obovate, obcompressed, glabrous; pappus absent. One species: J. trilobata Hemsl. KEY TO THE SPECIES AND VARIETIES A. Leaves broadly ovate to ovate-lanceolate, the upper little reduced........ ei ecc ec nr EN Y CUNCTI TORO la. I. heterophylla var. Dicksomii AA. Leaves ovate-pandurate to pandurate, trilobed or divided, the upper much reduced or lacking. B. Rays purple; involuere hemispherie, 2-3 em. in diameter; achenes pubescent. C. pos neos crenate, lateral lobes rounded; stems strigose- AA A A eese cl 1. I. heterophylla CC. San E -pandurate, entire or dentate, the lateral lobes acute; stems tuberculate-hispid.............. 1b. I. heterophylla var. acutiloba BB. Rays yellow; involucre campanulate, 1-1.5 cm. in diameter; achenes glabDT OU). RTT NINE TEES QT 2. I. trilobata 1. Iostephane heterophylla (Cav.) Benth. in Benth. & Hook. Gen. Pl. 2: 368. 1873; Hemsl. Biol. Cent.-Am. Bot. 2: 168. 1881. Coreopsis heterophylla Cav. Icon. Pl. 3: 34, pl. 268. 1795. Rudbeckia napif olia HBK. Nov. Gen. € Sp. 4: 244. 1820. Echinacea heterophylla (Cav.) D. Don in Sweet, Brit. Fl. Gard. II. 1: pl. 32. 1831. Echinacea dubia Knowles € Westeott, Fl. Cab. 3: 163. 1839. Stems erect, scapose to subscapose, 5-9 dm. tall, unbranched or branched in the taller specimens, striate, strigose-hirsute, densely so near the heads; basal leaves panduriform, with a long constriction between the terminal and the lateral lobes, posterior lobes roundish, crenately toothed, more conspicu- ously so between the anterior and lateral lobes, 1—3.2 dm. long, 2-6 em. broad, acute, strigose-tuberculate-hirsute especially on the upper surface, petioles 4—14 em. long; cauline leaves few, [ Vor. 22 82 ANNALS OF THE MISSOURI BOTANICAL GARDEN sessile, oblanceolate to lanceolate, 4-8 em. long, entire, reduced to narrow lanceolate bracts above; heads solitary, radiate, in- eluding the rays 4-9 em, broad; bracts of the involucre 1.5- 2 em. long, 5-6 mm. broad at the middle, oblong-lanceolate, at- tenuate, hirsute or hirsute-ciliate; pales 9-10 mm. long, cori- aceous, acute at the apex, slightly serrate on the upper mar- gins; rays 2.5-5 em. long, 7-10 mm. broad, 12-15, bi-tri-den- tate; disk-corollas sparsely pubescent on the expanded region ; achenes oblong-obovate, about 4 mm. long, obcompressed, pap- pus absent, crown and body clothed by a brown strigose-hirsute pubescence, becoming less dense towards the base. Distribution: on dry oak-covered hills, south-central Mexico. MEXICO: SAN LUIS POTOSI—Alvarez, Sept. 5-10, 1902, Palmer 65 (G, M, US); region of San Luis Potosi, Sept., 1876, Schaffner 253 (G); MEx1co—*'Santa Fe, Valley of Mexico,’’ July 26, 1865-66, Bourgeau 602 (G, US); ''Santa Fe, Valley of Mexieo,’’ Aug. 22, 1903, Rose $ Painter 6493 (G, NY, US); among oak hills, near ** El Oro,’’ alt. 9000 ft., Oct. 8, 1902, Pringle 9929 (G, M, US); dry woodland slopes, Sierra de las OR Aug. 28, 1892, D 5249 (G, US); Sierra de las ign Sept. 10, 1900, Pringle 9276 (G); near San Angel, Aug., 1855, Schaffner 58 (G); MICHOACAN—near Morelia, AN Arsène 5518 (US); near Morelia, Sul 5, 1909, Arsène 2464 (US); PUEBLA—on ‘‘cerro Tepaxuchitl,’’ alt. 6990 ft., vicinity of Puebla, Aug. 7, 1910, Arsène $ Nicolas 5408 (M, US). 1a. var. Dicksonii (Lindl.) W. M. Sharp, comb. nov. Echinacea Dickson Lindl. in Edwards’ Bot. Reg. N. S. 1: pl. 27. 1838 Similar to the species, but usually more robust ; stem 0.5-1 m. tall; basal and lower stem-leaves broadly ovate to ovate-lanceo- late, 2-4.5 dm. long, 4-15 em. broad, coarsely dentate, acute at the apex; heads including the rays occasionally 10 cm. in diameter. Distribution: mountains of Mexico, from southern Chihua- hua to Oaxaca. MEXICO: CHIHUAHUA—southwest Chihuahua, Aug., 1885, Palmer 333 (G, US); 15 mi. south of Guadalupe Calvo, alt. 7500-8000 ft., southwest Chihuahua, Aug. 22, 1898, Palmer 4824 (G, US); DURANGO—region of Bu Aug. 16, 1897, Rose 2336 (G); JALISCO—in Sierra Madre, west of Bolonas, Sept. 15-17, 1897, Rose 2969 (G, US); SAN LUIS POTOSI—region of San Luis Potosi, 22° N. lat., alt. 6000-8000 ft., 1878, diga $ Palmer 470 (G); near Huajalote, Ehrenberg 344 (G); HIDALGO—on Real del Monte, oe Coulter 374 (G); PUEBLA—vicinity of Puebla, Aug. 8, 1907, Arséne 1056 (G, M, US); Mt. Orizaba, near Esperanza, Aug. 5, 1891, Seaton 366 (G, US); vicinity of Esperanza, Sept., 1911, Purpus 5694 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 83 (G) ;MORELOS—Cuernavaca, Sept. 30, 1899, Holway 3548 (G); oaxaca—‘‘Serro San Felipe del agua," alt. 5700 ft., July 25, 1897, Conzatti $ Gonzalez 404 (G). 1b. var. acutiloba W. M. Sharp, var. nov.® Stems 4-9 dm, long, tuberculate-hispid, strigose; basal leaves trilobed-pandurate, the lateral lobes Ya as long as the termina] lobe, posterior lobes acute, entire, above tuberculate- hispid, beneath hispid; achenes oblong-obovate, 4—5 mm. long, canescent-strigose towards the summit; in all other characters similar to the species. Distribution: known only from the state of Jalisco, under oaks on rocky hills. MEXICO: JALISCO—near Guadalajara, Aug., 1893, Pringle 4480 (G, M TYPE, US); **Rio Blanco," Aug., 1896, Palmer 372 (G, NY, US). Fe Iostephane trilobata Hemsl. Biol. Cent.-Am. Bot, 2: 169. UY chrysantha Klatt in Leopoldina 23: 143. 1887. Gymnolomia scaposa Brandegee in Univ. Cal. Publ. Bot. 4: 93. 1910. An herbaceous, strigose-pubescent perennial, 1.5-7 dm. tall; stems erect, one or more from a thickened rootstalk; basal leaves panduriform to ovate, 0.5-3 dm. long, 1.5-7 em. broad, lateral lobes rounded, terminal lobe ovate to subovate, entire to sparsely serrate, hirsute-ciliate, strigose-hirsute below, stri- gose-tubereulate-hirsute to hirsute above; cauline leaves re- duced, lowermost lanceolate, 6 em. or less long, the uppermost reduced to sessile braets subtending the branches; petioles of the basal leaves more than half the total length of the blade; heads solitary or disposed in an open few-headed panicle, in- cluding the rays 2-5 em. broad, 6-13 mm. high; involucre cam- panulate; bracts of the involucre 2-3-seriate, lanceolate-acumi- nate, hirsute-pubescent; rays 6-13, yellow, 7-12 mm. long; achenes oblong-obovate, obcompressed, glabrous, epappose. * Iostephane heterophylla var. acutiloba W. M. Sharp, var. nov., caulibus 4-9 dm. longis, tuberculato- nel "re foliis basalibus trilobato- ¿A lobo terminali lobis lateralibus duplo longiore, lobis posterioribus acutis, integris, supra tuberculato-hispidis, subtus kae achaeniis oblongo-obovatis, 4-5 mm. longis, ad pieem griseo-strigosis; cetera speciei similis.—Collected at Jalisco, Mexico, near Guadalajara, Aug., 1893, Pringle 4480 (G, M TYPE, US). [Vor. 22 84 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: southern Mexico, in oak forests at elevations above 5000 feet. MEXICO: PUEBLA—in oak forests, Coxeatlan, alt. 8000-9000 ft., Sept., n Purpus 4120 (G, M); oAxaca—in Sierra de Chavellinas, alt. 8500 ft., Oct. 19, Pringle 4978 (G, M, US); 18 mi. southwest of the city of Oaxaca, ‘alt t. o ft., Sept. 10-20, 1894, Nelson 1371a (US); CHIAPAS—Oopen forests, near Fenia, July, 1925, Purpus 55 (G, US); near San Cristobal, alt. 7000-8800 feet, Sept. 18, 1895, Nelson 3223 (G, US); without locality or date, Ghiesbreght 101 (G); with- out locality, coll. of 1864-70, Ghiesbreght 561 (G, M). EcurNAcEA Moench History.—Previous to the time of Linnaeus, plants now con- sidered to belong to the genus Echinacea were discussed by Plukenet! in * Almagestum Botanicum’ in 1696 and by Catesby? in *Natural History' under the name Chrysanthemum ameri- canum. Morison? in ‘Plantarum Historia’ in 1699 described Dracunculus Virginianus latifolius. Illustrations of both Catesby and Morison are identical with what are now consid- ered to be Echinacea. Catesby’s plate is apparently that which is now interpreted as E. laevigata, but his description accom- panying the plate is somewhat vague. In ‘Species Plantarum’ Linnaeus* described an echinaceous plant under Rudbeckia purpurea. This name is based on Chrysanthemum ameri- canum of Plukenet and of Catesby and on Dracunculus Vir- ginianus latifolius of Morison. Necker* in ‘Elementa’ in 1790 described the genus Brauneria. No species are mentioned, but the generic description is practically the same as that of Rud- beckia. Moench® in *Methodus Plantarum" in 1794 described the genus Echinacea. The name is derived from the Latin word ‘‘echinus,’’ a hedge-hog, because of the prickly pales of the receptacle which give it a somewhat echinate appearance. His generic characters are clear-cut and comprehensive, the genus being based on Rudbeckia purpurea L. which, according 1 Plukenet, L., Alm. Bot. 99. 1696; ibid. pl. 21, fig. 1. 1720. 3 Catesby, M., Nat. Hist. Car. & Fla., ed. 3, 2: 59, pl. 59. 1771. * Morison, R., Pl. Hist. pt. 3. sect. 6. 42, pl. 9. 1699. * Linnaeus, C., Sp. Pl. 2: 907. 1753. 5 Necker, J. D., Elem. Bot. 1: 17. 1790. * Moench, C., Meth. Pl. 591. 1794. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 85 to the International Rules of Botanieal Nomenclature, becomes the type species of the genus Echinacea. The botanists of the early and middle nineteenth century adopted the name Echinacea, while Brauneria, although four years older, remained in obseurity. Lessing,” Endlicher,? Meisner,? Lemaire," and Bentham and Hooker! used the former name. Alphonse DeCandolle!? recognized the name Echinacea in the *Prodromus' and described under it four species. Torrey and Gray"? in the “Flora of North America’ included three species in this genus. Thomas Nuttall!* in “Transactions of the American Philosophical Society” de- seribed three species, one of which, E. pallida Nutt., is valid at the present time, while the others have been reduced to synon- ymy or excluded from the genus. Gray,** in various editions of his ‘Manual of Botany’ from 1857-1889, recognized the genus Echinacea, listing two species. In the seventh edition, however, the name Brauneria is adopted, and four species are given. In the first edition of the “Illustrated Flora of the Northern United States and Canada' Britton and Brown!* used the name Brauneria, but in the second edition this is re- placed by Echinacea. Small!” in ‘Flora of the Southeastern United States' recognized the genus Brauneria and included six species, four of which are retained in this monograph, one reduced to a variety, and the sixth being excluded. GENERAL MORPHOLOGY Stems.—The stems arise from rootstalks which die down each season and come up again in the spring, the parts above the ground thus being annual. The stems are of two types. uns: C. F., Syn. Gen. Comp. 225. 1832. ® Endlicher, S., Gen. Pl. 409. 2d ® Meisner, C. F., Pl. Vase. Gen. 202. 1839. 2 Lemaire, C., Diet. Hist. Nat. 183-184. 1844. u Bentham, G., € Hooker, J. D., Gen. Pl. 2: 366. 1873. ? DeCandolle, A. Prodr. 5: 554. 1836. * Torrey, J., € Gea y, A., Fl. N. Am. 2: 305-306. 1842. ^ Nuttall, T. in Trans, Am. Phil. Soc. II. 7: 354. 1841 ray, A., Man. of Bot., editions 1857—1889. * Britton € Brown, Ill. Fl. ed. 1,3: 419. 1898, and ed. 2, 3: 476. 1913. " Small, J. K., Fl. Southeast. U. S. 1261-1262. 1903. [Vor. 22 86 ANNALS OF THE MISSOURI BOTANICAL GARDEN The first type is usually simple, sometimes once-branched, and one to three feet high, a single rootstalk sometimes giving rise to one or several stems. The second type is usually branched above one to many times, and bears a single head on each branch. Thestems in E. purpurea commonly attain a height of four to five feet. Leaves.—The leaves of Echinacea fall naturally into two groups, namely, the ovate and the lanceolate. The latter is the most common form in the genus. "The basal leaves are usually long-petiolate, and the stem-leaves short-petiolate becoming sessile above. The leaves are normally three to five palmately veined. The primary veins run almost parallel in the narrow- leaved species. Pubescence.—' The pubescenee within the genus is variable, each species having a characteristic form. For this reason it is one of the most outstanding taxonomie characters for the group and especially important for a satisfaetory separation of the narrow-leaved species. In Echinacea paradoxa the pu- bescence is definitely strigose; in E. pallida it is hirsute; while in E. angustifoliaitis tuberculate-hirsute to tuberculate-hispid. This last species is often referred to as having shaggy pubes- cence, the tuberculate-hispid hairs giving this appearance. Echinacea laevigata is the only glabrous species in the genus. Involucral bracts.—The bracts of the involucre are usually in two to three series. In E. paradoxa the bracts show a marked transition into the pales, this transition indicating plainly the relation of pales to involucre. The pubescence on the bracts in all species is the same type as that of the leaves. When E. purpurea is grown in cultivation, abnormal heads are formed; the bracts of the involucre become leaf-like, and the receptacle somewhat elongated, thus showing a reversion per- haps to a more primitive form. Receptacle and Pales.—The receptacle in Echinacea is dis- tinetly conical. The pales on the receptacle are broad and per- sistent, terminating in coriaceous stiff or weak spines. The spiny apices extend from 1 to 4 mm. beyond the corolla, giving the head an echinate appearance. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 87 Ray-flowers.—The ray-flowers are neutral and disposed in a single series. The rays are strap-shaped, one to three toothed, and in a majority of the species are sparsely pubescent on the outer surface. In E. pallida they are 4-9 em. long and strongly reflexed. The predominating color is purple, but in E. para- doxa the rays are yellow. The Disk-corolla.—The corolla is expanded below into a bulb-like base. This enlarged portion is sessile upon the top of the achene, this being one of the most important generic char- acters. The corolla contraets above into a cylindrical tube ter- minated by a five-lobed, erect or slightly ascending limb. The median vaseular bundles of the corolla-lobes are absent, but there are five fused lateral bundles which divide at the sinuses and run along the margins of the corresponding lobes. The corolla is dark purplish as a rule. The stamens are five in number, united, and encompass the style. Each stamen is terminated by an ovate membranaceous appendage, sagittate at the base. GEOGRAPHICAL DISTRIBUTION The genus Echinacea has its center of development in the prairies and glades of the central United States, including the region from Illinois to Iowa, south to Tennessee and Oklahoma. The species are usually inhabitants of dry, rocky glades, with a preference for limestone soils. Echinacea purpurea has the widest distribution, its range extending from Georgia, north and west to Kentucky and the Ozarkian region of Missouri and Arkansas. The remaining species are less widely distributed; for example, in the region east of the Mississippi River, E. laevigata occurs along the Appalachian Upland east of the Al- leghenies from Pennsylvania to Georgia. A more restricted range is found in E. angustifolia var. tennesseensis, which is known only from the dry, rocky hills or pine barrens of Ruther- ford County, Tennessee. Echinacea pallida is an inhabitant of the prairies and glades of the central United States, and ex- tends into Oklahoma and Texas. Echinacea paradoxa is an Ozarkian species extending from southwestern Missouri to Texas, while E. angustifolia is confined to the dry prairies and barrens ranging from northern Minnesota to Texas. [Vor. 22 88 ANNALS OF THE MISSOURI BOTANICAL GARDEN Economic U ses.—E chinacea is at present of little importance in an economie way. The main value lies in horticultural pos- sibilities. The New Colwall Strain of Echinacea purpurea has been recently placed on the market in England. It is of easy eulture, quite hardy, and the handsome heads are often six inches in diameter. The most beautiful part of the head is the cone center which is green or bronze when young and later be- comes gorgeously tinted with metallic shades of copper, bright brown, or red. Other species of this genus also make attractive border plants. TAXONOMY Echinacea Moench, Meth. Pl. 591. 1794; Less. Syn. Gen. Comp. 225. 1832; DC. Prodr. 5: 554. 1836; Endl. Gen. Pl. 409. 1838; Meisner, Pl. Vase. Gen. 202. 1839; Endl. Ench. Bot. 238. 1841; Torr. & Gray, Fl. N. Am, 2: 305. 1842; Lemaire in Diet. Hist. Nat. 183-184. 1844; Benth. & Hook. Gen. Pl. 2: 366. 1873; Gray, Syn. Fl. N. Am. 1?: 258. 1884. Dracunculus Morison, Pl. Hist. pt. 3. sect. 6. 42, pl. 9. 1699. Bobartia Petiver, Herbal, 473. [1715]. 1704, Brauneria Necker, Elem. Bot. 1: 17. 1790. Helichroa Raf. Neogenyton, 3. 1825. Herbaceous perennials, arising from vertical or horizontal rootstalks. Stems erect, simple or branched, smooth to va- riously pubescent. Leaves alternate, ovate, ovate-lanceolate to lanceolate, acute or attenuated, coarsely toothed or entire, dark or light green, smooth or pubescent, 3—5-veined, medium in texture. Involueral bracts 2-3-seriate, foliaceous, linear- lanceolate to lanceolate, attenuate or acute, ciliate, smooth or pubescent, of medium texture. Receptacle distinctly conical. Pales conduplicate, terminating in sharp cartilaginous spines, exceeding the disk-flowers, straight or somewhat incurved. Ray-flowers neutral, disposed in one series, rays strap-shaped, bifid or trifid, purple or occasionally yellow. Disk-flowers fertile. Disk-corollas expanded below into a bulb-like base and sessile upon the achene, contracted above into a cylindrical tube and the five-lobed erect limb, smooth, purplish. Stamens five, anthers sagittate at the base, terminating in an ovate, 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 89 membranaceous appendage. Styles usually enlarged slightly at the base, equalling or slightly shorter than the corolla, the branches aeuminate, hairy. Achenes four-angled, in cross- section diamond-shaped at top, smooth, often sparsely pubes- cent on the angles, terminated by a toothed crown, body of the achene usually glabrous, slightly areolated. Type species: E. purpurea Moench, Meth. Pl. 591. 1794. KEY TO THE SPECIES A. Leaves ovate to ovate-lanceolate; pales slender, terminating in long weak spines. B. Stem and leaves hirsute, not glaucous.........oooooomomo... 1. E. purpurea BB. Stem and leaves smooth or glaucous........ooooooooooooo... 2. E. laevigata AA. Leaves lanceolate, attenuate at each end; pales stout, terminating in stiff spines. C. Plants hirsute or tuberculate-hirsute; stem and leaves dark green; rays purple. D. Stem and leaves tubereulate-hirsute to tuberculate-hispid; rays 2-3.5 em. long, not reflexo, -s askese 2n vé ves crec. 3. E. angustif olia DD. Stem and leaves hirsute; rays 4-9 em. long, reflexed........ 4. E. pallida CC. Plants strigose; stem and leaves light green; rays yellow....5. E. paradoza 1. Echinacea purpurea Moench, Meth. Pl. 591. 1794; DC. Prodr. 5: 554. 1836; Spach, Hist. Nat. Veg. 10: 52. 1841; Torr. & Gray, Fl. N. Am. 2: 305. 1842; Gray, Syn. Fl. N. Am. 1?: 258. 1884; Chapm., Fl. Southern U. S., ed. 3, 248. 1897; Britt. & Brown, Ill. Fl. ed. 2, 3: 475, fig. 4456. 1913; Rydb. Fl. Cent. N. Am. 837. 1932; Small, Man. Southeast. Fl. 1421. 1933. Rudbeckia purpurea L. Sp. Pl. 2: 907. 1753; Walt. Fl. Car. 214. 1788; Michx. Fl. Bor.-Am. 2: 143. 1803; Nutt. Gen. N. Am. Pl. 178. 1818; Link, Enum. Pl. 353. 1821; Barton, Fl. N. Am. 2: 84, pl. 64. 1822; Elliot, Sketch Bot. Car. & Ga. 2: 449. 1824. Chrysanthemum americanum Pluk. Alm. Bot. 99. 1696; ibid. pl. 21. fig. 1. 1720. Dracunculus Virginianus latifolius Morison, Pl. Hist. pt. 3. sect. 6. 42, pl. 9. 1699. Rudbeckia serotina Sweet, Brit. Fl. Gard. 1: pl. 4. 1823-25. Echinacea intermedia Lindl. in Paxt. Mag. Bot. 15: 79. 1849. Brauneria purpurea (L.) Britt. Mem. Torr. Bot. Club 5: 334. 1893; Britt. & Brown, Ill. Fl. ed. 1, 3: 420, fig. 3895. 1898; [Vor. 22 90 ANNALS OF THE MISSOURI BOTANICAL GARDEN Small, Fl. Southeast. U. S. ed. 1, 1261. 1903, and ed. 2, 1261. 1913; Rob. & Fern. in Gray's Manual, ed. 7, 832. 1908. Stonis erect, stout, branched, hirsute below, becoming smooth above, 6-18 dm. high; leaves ovate to ovate-lanceolate, acute, coarsely or sharply serrate below, usually entire above, hirsute on both surfaces, mature leaves often rough to the touch, 3—5- veined; basal leaves long-petiolate, 1.7-4.7 dm. long, 6-9 em. broad; cauline leaves short-petiolate, sessile above, 7-19 cm. long, 1.5-7 em. broad; involueral bracts linear-lanceolate, at- tenuate, entire, ciliate, pubescent on outer surface, texture similar to the leaf; heads 1.5-2.7 em. high, 1.1-4 cm. broad ex- clusive of the rays; pales slender, lanceolate, 11-13 mm. long, 1-1.5 mm. broad, terminating in long spines, surpassing the corolla 5-6 mm., smooth, purplish to brownish; rays 4—5.5 em. long, 5-10 mm. broad, bidentate or entire, reddish-purple; achenes 4—4.5 mm. long, 2-3 mm. broad at top, glabrous. Distribution: prairies and dry open woods from Ohio to Georgia, west to Iowa and Arkansas. GEORGIA: dry soil, Gwinnett Co., July 7, 1897, Eggert (M); ‘‘Griers Fave,’’ Randolph Co., July 17, 1903, Harper 1884 (M). OHIO: Mineral Springs, Adams Co., Oct. 5, 1930, Stephenson (M). TENNESSEE: Cumberland Mt., Franklin Co., July 21, 1897, Eggert (M); C berland Mts., Cowan, July, 1898, Ruth 646 (M); Wolf Creek, July 28, 1994 ). ALABAMA: Chambers Co., June 21, 1897, Earle (M). ILLINOIS: Wady Petra, Stark Co., July 19, 1900, Chase 685 (M); Hancock Co., em 1841 E J Progro St. Louis Co., July 27, 1898, Brownell (M); Shell Mound, Bar org 1L 1997, Palmer 32445 (M); near St. Louis, St. Louis Co., Aug. 4 1892, bind 275 (M); Windsor Springs, July 28, 1891, bugie (M); Tunis 2 July 16, 1898, Glatfelter (M); Oakville, Jefferson Co., Sept. 1, 1926, er ias (M); Dasco, Ralls er , Aug. e 1915, Davis 4864 (M); Hannibal, n uly 6, 1916, Davis 1574 (M); Webster Groves, St. Louis Co., eh CAM 4007 (M); near Allenton, ha 30, 1897, Lau Qo; ames River, Stone Co., June 3, 1914, Palmer 5843 (M); Mark Twain’s Cave, Marion Ps Aug. 24, 1915, Davis 32 284 (M); Arcadia, Iron Co., Sept., 1897, Russell (M); Pleasant Grove, Ripley Co., July 20, 1897, Mackenzie 373 (M); McDonald Co., July 24, 1892, Bus (M); Silex, Lincoln Co., Sept. 16, "1016, Davis 1844 (M); an Co., Aug. = 1884, Bush (M) ; Eagle Rock, Barry Co., June 8, 1897, Bush 26 (M); Noel, MeDon- ald Co., Sept. 10, 1913, Palmer 42569 (M ); Spring Park, July 24, 1891, Hassford (M); Oak Grove, Jackson Co., July 30, 1902, Mackenzie 64 (M); Monteer, Aug. 8, 1910, Bush 6146 (M); river bluffs, near Hannibal, Marion Co., Jan. 9, 1915, Davis 5819 (M). 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 91 ARKANSAS: Eureka Springs, July 16, 1898, Glatfelter (M); Benton Co., Plank (M); Fulton, Hempstead Co., June 19, 1915, Palmer 8040 (M). 2. Echinacea laevigata (Boynton & Beadle) Blake in Jour. Wash. Acad. Sci. 19: 273. 1929; Small, Man. Southeast. Fl. 1421. 1933. Brauneria laevigata Boynton & Beadle in Small, Fl. South- east. U. S. ed. 1, 1261. 1903, and ed. 2, 1261. 1913. Stems erect, simple or branched, glabrous, striate, 9-12 dm. high; leaves glabrous on both surfaces, palmately 3-5-veined, medium in texture; basal leaves long-petiolate, 18-36 cm. long, 3-7 em. broad, cauline short-petiolate below, sessile above, ovate to broadly lanceolate, acute, finely serrate, 7-18 em. long, 2-5 em. broad; heads 2-2.5 em. high, 3-3.5 em. broad exclusive of the rays, pales slender, linear-lanceolate, 9 mm. long, 1 mm. broad, tips eurving inwards, surpassing the corolla; rays 3.5- 8 em. long, 3-5 mm. broad, reflexed bidentate, purple; disk- corolla 5 mm. long, purplish; achenes 4 mm. long, glabrous, somewhat areolated, pappus a toothed crown of equal length. Distribution: fields and woods, Laneaster Co., Pennsyl- vania, south to South Carolina and Georgia. PENNSYLVANIA: Lancaster Co., ‘‘Caernsvron’’ township, July, without collector M). VIRGINIA: Bradford Co., July 18, 1871, Curtiss 3502 (M); near Staunton, Au- gusta Co., June 18, 1896, Murrill (NY). SOUTH CAROLINA: Keowee, Oconee Co., May 22, 1906, House 2206 (M). GEORGIA: in mountains of Georgia, Buckley (M). 3. Echinacea angustifolia DC. Prodr. 5: 554. 1836; Torr. & Gray, Fl. N. Am. 2: 306. 1842; Gray, Syn. Fl. N. Am. 1?: 258. 1884; Britt. & Brown, Ill. Fl. ed. 2, 3: 476, fig. 4457. 1913; Rydb. Fl. Pr. & Pl. Cent. N. Am. 837, fig. 570. 1932. Brauneria angustifolia (DC.) Heller in Muhlenbergia 1: 5. 1900; Small, Fl. Southeast. U. S. ed. 1, 1261. 1903, and ed. 2, 1261. 1913; Rob. & Fernald in Gray's Manual, ed. 7, 832. 1908; Rydb. Fl. Rocky Mts. ed. 1, 926. 1917, and ed. 2, 926. 1922. Echinacea sanguinea Nutt. in Trans. Am. Phil. Soc. II. 7: 354. 1841. Stems simple, 3-5 dm. high, tuberculate-hirsute to tubercu- late-hispid; leaves oblong-lanceolate to long-elliptical, entire, [Vor. 22 92 ANNALS OF THE MISSOURI BOTANICAL GARDEN dark green, tuberculate-hirsute to tuberculate-hispid, rough to the touch; basal leaves long-petiolate, 9-22 em. long, 1-2.5 cm. broad, cauline sessile, 4-14 cm. long, .6-1.5 cm. broad, acute; heads 1.9-3 em. high, 1.5-2.5 em. broad exclusive of the rays, involucral bracts lanceolate, acute, entire, 6-11 mm. long, 2- 3 mm. wide, densely tuberculate-hirsute or tuberculate-hispid; rays spreading, sparsely pubeseent on the veins below, 2— 3.5 em. long, 3-8 mm. broad, once- or twice-notched, purplish ; disk-corollas 6-7 mm. long, purplish ; achenes 5 mm. long, main body areolated by light brown and dark brown splotches, pap- pus forming a toothed erown. Distribution: dry prairies and barrens, Minnesota, south to Texas, west to Wyoming. MINNESOTA: Pike Lake, Sept. 10, 1849, Sykes (M). NORTH DAKOTA: Minot, Ward Co., Aug. 3, 1928, Larsen 198 (M). Sours DakoTA: Brookings, Brookings Co., without date, Williams (M); Dead- wood, Lawrenee Co., July 14, 1913, Carr 90 (M); Ionia, Lyman Co., July 5, 1910, Rose (M); Triquois, Kingsbury Co., Aug. 9, 1894, Thornber (M); Forestburg, San- born Co., July 3, 1910, Visher 4454 (M); near Hot Springs, Fall River Co., June 27, 1929, Palmer 37546 (M). NEBRASKA: without locality, July, 1904, Goodding 2215 (M); Lincoln, Lancaster Co., June, 1886, Webber (M). KANSAS: Ellis Co., 1908-11, Zeller (M); Manhattan, Riley Co., July 12, 1892, Clothier (M) ; prairie, June 18, 1895, Norton 265 (M); Rockport, Rooks Co., June 26, 1889, Bartholomew (M) ; east of Russell, Russell Co., July 1, 1926, Heller 13984 (M); Fort Kearney, July 10, 1856, Engelmann (M). OKLAHOMA: Cache, Comanche Co., June 25, 1913, Stevens 1341K (M); near Fort Sill, June 14, 1916, Clemens 11837 (M); near Cora, Woods Co., May 28, 1913, Stevens 737 (M); near Rocky, Washita Co., June - ane Stevens 971 (M TEXAS: near Canyon City, Randall Co., Aug. 14, 1900, Eggert (M); Dallas, Dallas Co., June 15, 1898, ... 05; Pos dite Spring, Comanche Co., May, 1849, Lindheimer 898 (M); Kerrville, Kerr Co., May 7-14, 1894, Heller 1735 (M); Leon Springs, Bexar Co., aiii $ Clemens 381 (M). WYomMING: Lake De dun. July 30, 1901, Nelson 8546 (M); Sundance, Crook Co., July 21, 1896, Nelson 2122 (M); Dome Lake, Sheridan Co., June 27, 1897, Pammel $ Bienioh 247 (M). 3a. ir em (Beadle) Blake in Jour. Wash. Acad. Sei. 19: 1929. nn tennesseensis Beadle in Bot. Gaz. 25: 359. 1898; Small, Fl. Southeast. U. S. ed. 1, 1262. 1903, and ed. 2, 1262. 1913. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 93 Echinacea tennesseensis (Beadle) Small, Man. Southeast. Fl. 1421. 1933. Stems usually simple, 1-4.5 dm. high, hirsute or tuberculate- hirsute; leaves narrowly oblong or long-elliptical, acute, en- tire, 2-13 cm. long, 3-7 mm. broad, hirsute, often tuberculate- hirsute, short-petiolate below, sessile above; involucral bracts linear, 5-7 mm. long, 2 mm. broad, hirsute-ciliate; achenes 4.5— 5 mm. long, glabrous. Distribution: pine barrens and dry hills, Tennessee. TENNESSEE: Lavergne, Rutherford Co., July 14-Aug. 18, 1897, Eggert (M); Lavergne, Sept. 7, 1898, Eggert (M); same locality, Aug. 19, 1897, Biltmore Herb. 1107 (M); cedar barrens, ‘‘middle’’ Tennessee, July 18-19, Gattinger (M). 4, Echinacea pallida Nutt. in Trans. Am. Phil. Soc. II. 7: 354. 1841; Britt. € Brown, Ill. Fl. ed. 2, 3: 476, fig. 4458. 1913; Rydb. Fl. Pr. & Pl. Cent. N. Am. 837. 1932; Small, Man. South- east. Fl. 1421. 1933. Echinacea angustifolia Hook. in Curt. Bot. Mag. 17: pl. 5281. 1861, not DC. Rudbeckia pallida Nutt. in Jour. Acad. Nat. Sci. Phila. 7: 17. 1834. Brauneria pallida Britt. in Mem. Torr. Bot. Club 5: 333. 1894; Britt. & Brown, Ill. Fl. ed. 1, 3: 420, fig. 3895. 1898; Small, Fl. Southeast. U. S. ed. 1, 1261. 1903, and ed. 2, 1261. 1913; Rob. € Fern. in Gray’s Manual, ed. 7, 832. 1908. Stems usually simple, 6-9 dm. high, sparsely hirsute below, more densely so above, pedicels leafless, becoming somewhat enlarged or flattened near the head; leaves oblong-lanceolate to long-elliptical, entire, dark green, hirsute on both surfaces, triple-veined, medium in texture; basal leaves 1.8-3.1 dm. long, 1-3.5 em. broad, the cauline 10-27 em. long, 1-2.5 em. broad, acute; involueral bracts lanceolate or narrowly oblong, 8- 17 mm. long, 2.5-4 mm. broad, strongly ciliate, hirsute, often gradually passing into the echinaceous pales ; rays strongly re- flexed, 4-9 cm. long, 5-8 mm. broad, bidentate, purplish; achenes 4 mm. long, glabrous, pappus a toothed crown. Distribution: dry prairies and barrens, Illinois and Ken- tucky west to Nebraska, southwest to Texas. [Vor. 22 94 ANNALS OF THE MISSOURI BOTANICAL GARDEN LLINOIS: Peoria, Peoria Co., June 10, 1912, Churchill (M); near Wady Petra, Stark Co., June 25, 1900, Chase 642 (M) ; near French Village, St. Clair Co., June 15, 1876, Eggert (M). KENTUCKY: Bowling Green, Warren Co., Aug., 1890, Price (M). Towa: Iowa City, Johnson Co., Hitchcock (M); Columbus Junction, Louisa Co., June, 1925, Graves (M); Blaek Hawk Co., June 18, 1929, Burk ?72 (M); ledges, State Park, Boone Co., June 30, 1926, Pammel 243 (M). MISSOURI: Oakwood, Ralls Co., June 21, 1916, Davis 4185 (M) ; Winfield, Lincoln 196 (M); Pacific, Franklin Co., June 3, 1918, Greenman 4121 (M); Gray’s Summit, Franklin Co., May 25, 1927, Kellogg (M); Jefferson Co., May 25, 1896, Eggert (M); Potosi, Washington Co., June 6, 1892, Dewart (M); Ironton, Iron Co., June 26, 1920, Palmer 18088 (M); Jerome, Phelps Co., June 1, 1914, Kellogg 438 (M); Cape Girardeau, Cape Girardeau Co., June 22, 1920, Palmer 18016 (M) ; Gainesville, Ozark Co., June 26, 1928, Palmer 34749 (M); Swan, Taney Co., June 10, 1898, Bush 149 (M); Galena, Stone Co., May 28, 1914, Palmer 5783 (M); Eagle Rock, Barry Co., May 26, 1898, Bush 261 (M); Webb City, Jasper Co., June 22, 1902, Palmer 178 (M); Nichols’ Junction, Greene Co., June 15, 1898, Bush 38 (M); Sheffield, Jackson Co., June 26, 1894, Bush 156 (M). ARKANSAS: near Mandelville, Miller Co., June 10, 1898, Eggert (M); near Tex- arkana, June 8, 1898, Eggert (M); Fulton, Hempstead Co., June 19, 1915, Palmer 8061 (M). LOUISIANA: Jennings, Jefferson Davis Parish, May 15, 1915, Palmer 7632 (M). NEBRASKA: Lineoln, Laneaster Co., June 11, 1900, Hedgcock (M KANSAS: Cowley Co., June, 1898, White (M); Caney, Mein: Co., June 29, 1929, Rydberg $ Imler 400 (M OKLAHOMA: Page, Leflore Co., June 20, 1914, Blakley 1436 (M). Texas: Willis, Montgomery Co., June, Warner (M); Houston, Harris Co., June 2, 1915, Fisher (M); near Buchanan, Bowie Co., June 13, 1898, Eggert (M); Mineola, Wood Co., June, Reverchon 2072 2. Dallas, Dallas Co., June 26, 1899, Eggert (M); Big Sandy, Upshur Co., "af 1900, Reverchon 2579 (M); Hemp- stead, Waller Co., June 10, 1872, Hall 3 wf Weatherford, Parker Co., May 29, 1902, Tracy 8331 (M); Grapeland, dde Co., May 26, 1917, Palmer 12066 (M); Gladwater, Gregg Co., June, Reverchon (M); Boerne, Kendall Co., May 22, 1916, Palmer 9853 (M). 5. Echinacea paradoxa (Nort.) Britt. & Brown, Ill. Fl. ed. 2, 3: 476, fig. 4459. 1913. Brauneria paradoxa Norton in Trans. Aead. Sci. St. Louis 12: 40. 1902; Rob. & Fern. in Gray's Manual, ed. 7, 832. 1908; Small, Fl. Southeast. U. S. ed. 1, 1261. 1903, and ed. 2, 1261. 1913. Stems simple, 3.5-8.5 dm. high, stramineous or yellowish- green, sparsely or densely strigose, sometimes tuberculate- 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 95 strigose on the peduncles; leaves oblong-lanceolate to long- elliptical, entire, light green, strigose-pubescent on both sur- faces, more densely so below, medium in texture; basal leaves long-petiolate, 1-2.5 dm. long, 1-3.2 em. broad, the cauline .6- 3 dm. long, .5-2.5 em. broad, acute; involucral bracts 7-10 mm. long, 1-4 mm. broad, lanceolate, acute, the outer foliaceous, the inner passing into the pales, smooth, strigose-ciliate; rays 3- 7 em. long, 3-8 mm. broad, yellow; achenes 4-5 mm. high, gla- brous, teeth of the pappus-erown equal in length. Distribution: rocky slopes and barrens, Missouri to Texas. Missouri: Nichols’ Junction, Greene Co., June 15, 1898, Bush 42 (M); James River, Stone Co., June 3, 1914, Palmer 5844 (M) ; Swan, Taney Co., June 10, 1898, Bush 155 (M); near Seligman, Barry Co. June 1, 1926, Palmer 30397 (M); Eagle Rock, Barry Co., June 4, 1897, Bush 76 (M); near Gainesville, Ozark Co., June 26, 1928, Palmer 34714 (M). OKLAHOMA: Idabel, McCurtain Co., May 18, 1916, Houghton 3648 (M). Texas: Willis, Montgomery Co., Wa. arner (M); Houston, Harris Co., April 29, 1916, Palmer 9617 (M); Valley of the Trinity River, June 28, 1912, Ruth 243 (M). ZALUZANIA Pers. History. —The genus Zaluzania was first published by Per- soon! in *Synopsis Plantarum’ in 1807; it was based on Anth- emis triloba Ort., a plant native of Mexico. Lagasca,? in “Genera et Species Plantarum” in 1816, de- scribed the genus Ferdinanda and included two species, F. augusta, now Zaluzania augusta, and F. eminens, now Poda- chaenium paniculatum Benth. Cassini? Kunth,* DeCandolle,5 and Endlicher* recognized Lagasca’s genus in their treatment of the Heliantheae. The generic limitations of Ferdinanda were rather broad and included species with or without a pap- pus, etc., the pappose species now being included in Podachaen- vum. Chrysophania of Kunth* is synonymous wth Ferdinanda, since its type species C. fastigiata is a synonym of Zaluzania ! Persoon, C. H., Syn. P1. 2: 473. 1807. ? Lagasea, A. M., Gen. et Sp. Nov. Pl. 31. 1816. * Endlicher, S., Gen. Pl. 408. [Vor. 22 96 ANNALS OF THE MISSOURI BOTANICAL GARDEN augusta, the species on which Ferdinanda was based. Robin- son and Greenman' in 1899 treated Ferdinanda as a subgenus under Zaluzania. This position is retained in the present revi- sion of this genus. Hybridella was described briefly by Cassini? in “Bulletin des Sciences, par la Societe Philomatique' in 1817, and was based on Anthemis globosa Ort. Hybridella was never recog- nized as a genus by other authors. Robinson € Greenman” in “Proceedings of the American Academy” in 1899 treated it as a subgenus under Zaluzania, this disposition being retained in the present paper. It comprises those palustrine, herbaceous species of Zaluzania with finely dissected leaves. Chilophyl- lum of De Candolle? described in the ‘‘Prodromus’’ in 1836 is synonymous with H ybridella. A taxonomie study of Zaluzania was published by Schultz- Bipontinus in ‘‘Flora’’ in 1861!? and 1864.1! His treatise of 1861 comprised seven species, three of which were new, namely, Z. megacephala, Z. montagnaefolia, and Z. myriophylla. His publieation on the genus in 'Flora,' 1864, included twelve species, eight of which belong to Zaluzania, three have been transferred to Viguiera, and one to Calea. Between 1864 and 1898 four new species were added to Zalu- zania, namely, Z. mollissima, Z. Coulteri, Z. resinosa, and Z. discoidea. The genus now totalled eleven species, all native of Mexico, Zaluzania Coulteri having been found to be synony- mous with Z. cinerascens. The first comprehensive revision of the genus by Robinson and Greenman” in ‘Proceedings of the American Academy” in 1899 added two new species. The genus, at this time, was known only from Mexico and Arizona. Hieronymus"? in Engler's ‘Botanische Jahrbücher’ in 1900 described three new species from Ecuador. This is the only record of Zaluzania from South America; and of these three ' Robinson, B. L., and Greenman, J. M., in Proc. Am. Acad. Sei. 34: 531-534. 1899. * Cassini, M. H., in Bull. Soc. Philom. p. 12. 1817. * De Candolle, A., in Prodr. 5: 554. 1836. 1 Schultz-Bipontinus, C. H., in Flora 44: 561-565. 1861. ? Schultz-Bipontinus, C. H., in Flora 47: 216-220. 1864. ? Hieronymus, G., in Engl. Bot. Jahrb. 29: 35-37. 1900. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 97 species Z. Sodiroiis the only one retained in the present mono- graph; the other two pass into synonymy. GENERAL MORPHOLOGY Roots.—The roots of the various suffruticose and shrubby species of the genus are little known, because, due to their bulk, they have not been taken by collectors. This also may apply to the herbaceous species that attain a height of three feet or more. The roots are mostly biennial or perennial in all the species of Zaluzania. In Z. globosa and its variety the main root is relatively short and rhizome-like with numerous secon- dary fibrous roots. Zaluzama Grayana is the only species known to the author that possesses a thickened tuberous root- stalk surmounted by a slender, lignescent, branched base which gives rise to the suffrutescent stems. Stems.—The stems of Zaluzama are terete and usually more or less striated. They vary from herbaceous to lignescent. The herbaceous type is especially exemplified in Z. globosa, Z. anthemidifolia, Z. triloba, Z. discoidea, and Z. megacephala. The suffruticose forms are lignescent at their base or upwards for half the height of the plant, the flower-bearing branches being herbaceous. This type may be observed in Z. Robin- sonu and Z. Grayana. The shrubby species are especially ex- emplified by Z. Pringlei and Z. montagnaefolia. Leaves.—Two quite different forms of leaves are displayed in the genus, namely, (1) finely dissected leaves, and (2) undi- vided or lobed leaves. The former group, comprising species that inhabit moist or swampy localities, as Z. globosa and its variety myriophylla, often have leaf-segments 3—4-pinnatiseet. The second group is typified by the subgenus Ferdinanda. Three species, namely, Z. Grayana, Z. triloba, and Z. Robin- soni, have pinnately, or more commonly palmately, lobed leaves. The remaining species display undivided leaves which are ovate, ovate-lanceolate to ovate-elliptieal in outline, acute to broadly obtuse, and entire to variously dentate or serrate. The venation is chiefly of the pinnate type, usually with three prominent veins near the base of the leaf. Leaves with pro- (Vor. 22 98 ANNALS OF THE MISSOURI BOTANICAL GARDEN nounced retieulations are exemplified in Z. Sodiroi, and to a less degree in Z. montagnaefolia. Pubescence.—The common type of pubescence in Zaluzama consists of simple unbranched hairs. The species are all more or less pubescent, except Z. anthemidif olia, which is glabrous. The degree and character of this covering vary with the differ- ent species. The hirsute type is less common, Z. globosa and Z. triloba being the best examples. The softer types of tri- chomes, varying from crispy hirtellous, velutinous, to velvety tomentose, are the most common forms within the genus. The velutinous or velvety tomentose type of pubescence is shown in Z. augusta, Z. subcordata, Z. mollissima, and Z. discoidea. Short-stiped or sessile, globular, resinous glands are present in the majority of the species. Inflorescence.—' The eorymbose to subpaniculate types of in- florescence are dominant for Zaluzania. The Hybridella sec- tion has a tendeney towards the production of heads in very loose panicles, or solitary, as in Z. anthemidifolia. The in- florescence is of little significance in separating the species. The receptacle is eonieal with persistent pales. The pales are of two sorts, namely, the linear, plane, herbaceous type characteristic of Hybridella, and (2) the complicate or boat- shaped subcoriaceous type as in the subgenus Ferdinanda. The pales are of little importance for specific delimitation but they serve as important subgeneric characters. Corollas.—All the species of Zaluzania, with the exception of Z. discoidea and Z. Pringlei, have heterogamous flower- heads, with styliferous and fertile ray-flowers, and fertile, tub- ular disk-florets. The corollas are yellow in both ray- and disk- flowers. 'The ligules are ovate, oblong-ovate, to oblong, 2—3- dentieulate or subentire, always styliferous. 'lhe tube in some species is dilated at the base so as to completely cap the achene. The disk-corollas have a narrow cylindrical tube which is about half the length of the corolla proper, and always ex- panded at the base so as to cap the achene; the throat is cam- panulate, glabrous or pubescent, with a somewhat spreading five-lobed limb. The expanded base of the corolla and the fer- 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 99 tile ray-flowers constitute the major characters of generic importance. The androecium is composed of five syngenesious stamens with ovate appendages and sagittate bases, which are similar to those of related genera. The style-branches are slender, more or less recurved, with acute or obtuse appendages. Their apices are variously pubescent. Achenes and Pappus.—The achenes of the disk-flowers are ordinarily linear-cuneate or narrowly oblong and somewhat four-angled; the ray-achenes are triangulate. The disk- achenes are glabrous, the ray-achenes variously pubescent or glabrous. The variation of the achenes within the genus is very slight and consequently of no specific importance. The disk-achenes are devoid of a pappus. The ray-achenes are epappose or erowned by a pappus varying from short-hirsute hairs to fimbriate or squamulose setae. Geographical Distribution.—The center of distribution for this genus is in the Mexican Plateau. Zaluzania Grayana is the most northerly distributed species; it occurs in the mountains of Chihuahua and southern Arizona. The most extra-limital species is Z. Sodiroi, known to occur only in the mountains of Ecuador, South America. Five of the fourteen species in- cluded in this genus have a limited distribution; for example, Z. anthemidifolia is known only from Jalisco; Z. Pringlei in- habits limestone hills near Jojutla in the state of Morelos; Z. cinerascens is known to occur only among the fir forests of the Sierra Pachuca Mountains in the state of Hidalgo; Z. mega- cephala is found in the mountains of Coahuila at an elevation of 6500-10,000 feet; whereas Z. subcordata is an inhabitant of the dry, rocky or barren hills and mountains of southeastern Puebla. Zaluzanta discoidea and Z. montagnaefolia are known to occur only in two states of Mexico; the former inhabits the dry, rocky hills and mountains of Chihuahua and Durango, whereas the latter occupies a similar habitat in the states of Puebla and Oaxaca. The remaining five species occur in cen- tral Mexieo with a general range from Durango south to the states of Mexico and Hidalgo. [Vor. 22 100 ANNALS OF THE MISSOURI BOTANICAL GARDEN The species of the subgenus Hybridella occur at relatively lower altitudes in palustrine or moist habitats. The species as- signed to the subgenus Ferdinanda occur at high altitudes, ranging usually from 5000 to 10,000 feet elevations. TAXONOMY Zaluzania Pers. Syn. Pl. 2: 473. 1807; Less. Syn. Gen. Comp. 294. 1832; DC. Prodr. 5: 553. 1836; Endl. Gen. Pl. 408. 1838; Walp. Rep. Bot. Syst. 2: 611. 1843; Schz.-Bip. in Flora 44: 561-565. 1861; Ibid. 47: 216. 1864; Benth. € Hook. Gen. Pl. 2: 362. 1873; Hemsl. Biol. Cent.-Am. Bot. 2: 159. 1881; Hoffm. in Engl. & Prantl, Nat. Pflanzenfam. IV. Abt. 5: 233. 1890; Rob. € Greenm. in Proc. Am. Acad. 34: 530. 1899; Blake in Contr. U. S. Nat. Herb. [Standley's Trees & Shrubs Mexico] 23: 1537. 1926. Ferdinanda Lag. Gen. € Sp. Nov. Pl. 31. 1816; Cass. in Dict. Sei. Nat. 46: 404. 1827; Kunth in Less. Syn. Gen. Comp. 224. 1832; DC. Prodr. 5: 552. 1836; Eindl. Gen. Pl. 408. 1838. Hybridella Cass. in Bull. Soc. Philom. 12. 1817; in Diet. Sci. Nat. 22: 86. 1821. Chrysophania Kunth in Less. Syn. Gen. Comp. 224. 1832; DC. Prodr. 5: 553. 1836; Endl. Gen. Pl. 408. 1838. Chiliophyllum DC. Prodr. 5: 554. 1836, not Chiliophyllum Phil. in Linnaea 33: 132. 1864; Endl. Gen. Pl. 408. 1838. Perennial herbs or shrubs, often striated, glabrate, velu- tinous, tomentulose, or hirsute. Leaves alternate, simple, palmately or pinnately lobed to pinnatisect, entire, serrate to dentate-mueronate, hirsute, strigose-villous, tomentulose to velutinous, frequently interspersed by resinous glands. Heads radiate or discoid, in corymbose, subeorymbose, or subpanicu- late clusters, hemispherical to subglobose. Involucre campan- ulate to subeampanulate, 2—4-seriate, graduated or subequal, the outer bracts sometimes much longer than the inner, ovate, ovate-lanceolate to oblong-lanceolate, herbaceous, the inner series becoming subcoriaceous. Receptacle conical, about 3 mm. high; pales subherbaceous to coriaceous, trilobed or acute at the apex, plane to conduplicate, not deciduous with the achenes. Disk-corollas with the base dilated and capping the 1935] SHARP——CERTAIN EPAPPOSE GENERA OF COMPOSITAE 101 erown of the achene, the tube narrowly cylindrical, expanding above into the campanulate throat and the 5-lobed, spreading or reflexed limb, tube pubescent or glabrous. Stamens sagit- tate at the base, appendages ovate; style-branches more or less recurved, with obtuse or acute appendages, pubescent at the apex. Ray-corollas yellow, styliferous, the tube often capping the ray-achene. Achenes of the disk-flowers quadrangulat obeompressed or rhomboidal in cross-section, glabrous, Ted of the ray-flowers trigonal, glabrous or pubescent; pappus ab- sent in the disk-achenes, absent or consisting of minute fimbri- ate or squamellose setae in the ray-achenes. Type species: Z. triloba (Ort.) Pers. Syn. Pl. 2: 473. 1807. KEY TO THE SUBGENERA A. Leaves finely pinnatisect; pales linear, plane, herbaceous or subherbaceous; alustrine herbs....... Subgenus Hybridella (Cass.) Rob. & Greenm. Sp. 1-2 AA. Leaves simple or lobed; pales conduplicate, coriaceous to subcartilaginous; perennial herbs or shrubs of mesophytic or dry habitats.............. RE Subgenus Ferdinanda (Lag.) Rob. & Greenm. Sp. 3-14. KEY TO THE SPECIES AND VARIETIES A. Leaves narrowly aeneis MM or bipinnatisect; pales linear, plane, herbaceous or subherbac B. Leaves finely 2-4- ru ntm the ultimate segments less than 1 mm. broad. C. Ultimate leaf segmenta Aetio iro sans nenne 1. Z. globosa CC. Ultimate leaf-segments obtuse...........-. a. Z. globosa var. myriophylla BB. Leaves narrowly pinnate to "indie the ns segments 1-2 mm. lup ee CO TO 2. Z. anthemidifolia AA. Leaves undivided or lobed; an conduplicate, coriaceous or oM B. qose palmately or pinnately C. Plants hirsute- y or rudi RÀ heads 8-20 mm. broad in- hing the . Hir china, heads 15-20 mm. broad ineluding the rays; leaves bipinnately lobed or cleft; plants herbaceous.............. 3. Z. triloba DD. poii erisp-hirtellous; heads 8-12 mm P dig including the rays; ea bed o r tripartita] plants suffruticose......... 4. Z. Robinsonti CC. Plants er hirtellous to glabrate; heads 20-50 mm. broad includ- ihe rAyB....v0 (s y E cuR desees cle oases s RE 5. Z. Grayana BB. Hare undivided. E. Heads dis oid. F. Plants ruby; leaves ovate, strigose-puberulent die ute IMPER eve eco ean EN ID Z. Pringlei FF. Plants herbaceous from a ligneous base; leaves broadly mu to eltoid-ovate, velvety tomentose beneath........... 7. Z. discoidea [Vor. 22 102 ANNALS OF THE MISSOURI BOTANICAL GARDEN EE. Heads radiate. G. Leaves entire, serrate to erenate-serrate; plants of Mexico. H. Leaves pustulate-hirsute to pustulate-hispidulous above, ere- e-serrate, base decurrent and auriculate 8. Z. montagnaefolia HH. Leaves tomentose, velutinous or hirsutulous, base not aurie- ulate. I. Plants herbaceous or suffruticose, unbranched except in the inflorescence; pappus absent on the ray-achenes. J. Leaves broadly ovate to ovate-elliptieal, 144 times as 9. Z. as broad, hirtellous................ cinerascens JJ. ium lanceolate or pa lanceolate, about já times as long as broad, velutinous.......... 10. Z. megacephala II. Plants shrubby, freely branched below; pappus of the ray- chenes a crown of minute lacerate or squamose set H 1.5 em. long; leaf-blades attenuately deeurrent to the base of the petiole............... 11. Z. mollissima KK. Heads — the rays less than 1.5 cm. broad, rays m. long; leaf-blades subeordate, not de- eurrent " ern base of the petiole. L. Leaves entire, subeordate, densely tomentose be- [o PR rr 12. Z. subcordata LL. Leaves below the branches of the Hou ere- nate-serrate to coarsely serrate, velutinous be- Mic iras ee TRETEN 13. Z. augusta GG. Leaves O plants of South America. 14. Z. Sodiroi 1. Zaluzania globosa (Ort.) Schz.-Bip. in Flora 44: 564. 1861; Hemsl. Biol. Cent.-Am. Bot. 2: 159. 1881; Rob. € Greenm. in Proc. Am. Acad. 34: 530. 1899. Anthemis globosa Ort. Desc. Pl. 46. 1797; Jacq. Desc. Rar. P1.3: 64, pl. 372. 1798. Hybridella globosa (Ort.) Cass. in Dict. Sci. Nat. 22: 86. 1821. Chiliophyllum globosum (Ort.) DC. Prodr. 5: 554. 1836; Benth. Pl. Hartweg. 17. 1839. Perennial palustrine herbs; stems 2-5 dm. tall, slender, sul- cate, hirsute to pilose-hirsute, also glandular on the peduncles; basal leaves several, oblanceolate in general outline, finely pin- nately dissected, often 3-4-pinnatisect, 1-2 dm. long, 2-4 em. broad, hirsute to pilose-hirsute, and glandular, segments 2- 3 mm. long, linear-lanceolate, acute, the petioles pilose-hirsute ; stem-leaves few subtending the pedunculate branches, 2 to 8 on naked pedunculate branches, 2-3 em. broad including the 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 103 rays, 6-10 mm. long; bracts of the involucre herbaceous, lance- olate to oblanceolate, 5-6 mm. long, 1.5-2 mm. broad, hirsute- ciliate, acute, hirsute and more or less glandular; pales weak, almost herbaceous, narrowly linear, 3-4 mm. long, 0.5 mm. or less broad, ciliate, obtuse to acute, glandular on the outer sur- face; disk-flowers densely glandular on the outer surface of the tube; ray-flowers bidentate, 6-19 mm. long including the achene; mature achenes about 2 mm. long, glabrous, pappus absent. Distribution: low wet meadows and swampy grounds of central Mexico, San Luis Potosi, southeast to the state of Mexico. MEXICO: SAN LUIS POTOSI—22° N. lat., alt. 6000-8000 ft., 1878, Parry $ Palmer 527 (M); HIDALGO—Telles, Sept. 21, 1910, Orcutt 4139 (F, M); Pachuca, July, 1903, Purpus 77 (M, U CAL, US); MEXICO—wet meadows, Valley of Mexico, alt. 7300 ft., June 7, 1901, Pringle 9395 (G, M, US) ; low meadows, Valley of Mexico, Federal Distriet, July 27, 1890, Pringle 3204 (G, M, U CAL, Mh uneultivated p ed of Mexico, June 12, 1865-66, Bourgeau 385 (G, US); damp meadows, 0 ft., Valley of Mexico, Federal District, June 25, 1897, Pringle 7440 (US); dà of Mexico, June 24, 1889, Pringle 2925 (G); Valley of Mexico, Aug., 1855, 7 le 5274 (US); near Tlalnepantla, July 6, 1905, Rose, Painter $ Rose 8417 (US); Valley of Mexico, near Tacubaya, July 30, 1901, Rose $ Hay 5816 (US). la. var. myriophylla (Schz.-Bip.) W. M. Sharp, comb. nov. Zaluzania myriophylla Schz.-Bip. in Flora 44: 565. 1861; Rob. & Greenm. in Proc. Am. Acad. 34: 530. 1899. Zaluzania globosa Schz.-Bip. in Hemsl. Biol. Cent.-Am. Bot. 2: 159. 1881, in part. Leaves with the ultimate segments finely dissected, linear, short, 1-2 mm. long, obtuse, hirsute to glabrous; habit and floral characters as in the species. Distribution: central Mexico, Durango, south to the state of Mexico, inhabiting low wet meadows and swamps. MEXICO: DURANGO—city of Durango and vicinity, 1896, Palmer 307 (F, G, M, U CAL, 2 ZACATECAS—damp hollows Pe plains at Coler Station, Sept. 1, 1904, Pringle 4 (F, G, M, U CAL, US); san LUIS POTOSI—San Luis Potosi, 1879, oie i i. (345) (G, US); MEER ad a of Mexico City, 1837, Hartweg 111 (G The short, linear, obtuse, ultimate segments and the glabrous to sparsely hirsute, glandular leaves warrant at least a varietal [Vor. 22 104 ANNALS OF THE MISSOURI BOTANICAL GARDEN recognition. The ultimate segments of Z. globosa are lance- olate, 2-3 mm. long, and acute. 2. Zaluzania anthemidifolia Rob. & Greenm. in Proc. Am. Aead. 34: 531. 1899. Plants herbaceous; stems decumbent, glabrous; leaves pin- nate to bipinnate or bipinnatisect, pilose-hirsute on the midrib and petioles, especially the young leaves, the ultimate seg- ments 1-2 mm. broad, obtusish to acute; heads solitary or some- times 2 or 3 terminating the leafy branches ; peduncles 2-5 em. long; braets of the involuere obtuse or obtusish; rays about 1 em. long, bifid, tube and outer surface of the lamina glandu- lar; disk-corollas with a narrow glandular tube; achenes gla- brous; pappus absent. Distribution: wet sandy banks of the Rio de Santiago River, near Guadalajara, Jalisco. Mexico: JALISCO— wet sandy river banks, near Guadalajara, Sept. 23, 1891, Pringle aue (G TYPE); wet sandy banks of the Rio Grande de Santiago, barranca near lajara, Oet. 12, 1895, Pringle 736? (F, M, U CAL). 3. Zaluzania triloba (Ort.) Pers. Syn. Pl. 2: 473. 1807; Kunth in Less. Syn. Comp. 225. 1832; DC. Prodr. 5: 553. 1836; Schz.-Bip. in Flora 44: 564. 1861; Hemsl. Biol. Cent.- Am. Bot. 2: 160. 1881; Rob. & Greenm. in Proc. Am. Acad. 34: 531. 1899. Anthemis triloba Ort. Desc. Pl. Rar. 72. 1800. Anthemis trilobata Willd. Sp. Pl. 3*: 2186. 1804. Anthemis sinuata LaLlave & Lexarza, Nov. Veg. Desc. 1: 26. 1824. Herbaceous plants; stems about 6 dm. tall, often irregularly bluish-purple below, branched above, hirsute-hispid; leaves dark green, broadly pinnately parted, including the petiole 2- 16 em. long, 1-5 em. broad, the segments acute to obtuse, the basal divisions cleft almost to the midrib, the terminal divisions deeply cleft; main cauline leaves long-petiolate, light green beneath and somewhat glandular, densely hirsute-hispid to hirsute especially on the veins, dark green and hirsute-hispid to strigose-hirsute above; inflorescence subcorymbose or sub- paniculate; mature heads, including the rays, 1.5-2 em. broad, 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 105 6-10 mm. high, peduncles 2-5 em. long; bracts of the involucre 2-seriate, narrowly oblong-lanceolate to lanceolate, acute, hir- sute, the outer series greenish, about twice as long as the inner, the inner ones much paler; corolla 2.5 mm. long, the tube glan- dular, the throat pubescent; rays oblong, 8-10 mm. long, 3- 5 mm. broad, pubescent on the outer surface; disk-achenes 2 mm. long, glabrous, epappose, ray-achenes pubescent, bear- ing a crown of a few bristle-like hairs. Distribution: ealeareous mesas at an elevation of 5000 to 8500 ft, San Luis Potosi and eastern Zacatecas, south to Puebla. MEXICO: SAN LUIS POTOSI—22° N. lat., alt. E 8000 ft., 1887, Parry $ Palmer 445 (F, G, M, US) ; San Luis Potosi, alt. 5950 ft., Aug. 22, 1926, Fisher 136 (US); ZACATECAS—near Guadalupe, Aug., 1855, Schafe 78 oe ney rez 1865-66, Bourgeau 704 (G, US); alt. 8000 ft., Aug. 8, 1898, Dea 7 (F, G); eity of Zacatecas, Palmer 757 (G) ; GUANAJUATO—Sept., 1891, bon Bi (G) ; Jaral, 1885, Schumann 94 (335) (US); QUERETARO—from ‘‘Cierro to San Juan,’’ Aug. 27, 905, Altamirano 1744 (US); HIDALGO—Telles, Sept. 21, 1910, Orcutt 1131 ); Pachuca, Sept. 5, 1910 Orcutt 3902 (F, M, US); Sierra de Pachuca, ; zh e G, M, US); Zontecomate, alt. 8500 ft., June 22, 1904, Pringle 13093 (F, G, US); near Tula, July 3-4, 1905, Rose, Painter $ Rose 8325 (US); PUEBLA—Orizaba Mts., Botteri 843 (G, US). 4. Zaluzania Robinsonii W. M. Sharp, sp. nov.'* Plants suffruticose; stems 3-7 dm. tall, woody below, her- baceous and branched above, sulcate by alternating light green ridges and green grooves, canescent, densely erisp-hirtellous, especially above, sparsely and minutely glandular ; leaves peti- olate, including the petiole 1.5-6 em. long, 1-4 em. broad, of medium texture, erisp-hirtellous and minutely glandular on both surfaces, palmately 3-lobed or tripartite, lobes acute or aeutish, subeordate, the anterior lobes divided or sometimes pinnatifid; heads few, in close corymbose clusters, including the rays 8-12 mm. broad, about 5 mm. high, the peduncles of 13 Z. Robinsonii Sharp, sp. nov. Plantae suffruticosae; caulibus 3-7 dm. altis, infra ligneis, supra herbaceis et ramosis, canescentibus, dense erispo-hirtellis; foliis palmater 3-lobatis vel tripartitis, petiolis pecie em. longis, 1-4 cm. latis, lobis E subcordatis, supra et subtus crispo- hirtellis; capitulis radiis in- cludentib mm. latis, 5 mm. altis; radiis 5-6 mm. longis, tubo dense villoso.— Collected 2 Cedros, Zacatecas, Mexi, F. E. Lloyd 124 (M TYPE). [Vor. 22 106 ANNALS OF THE MISSOURI BOTANICAL GARDEN the corymb short, 0.5-3 em. long; bracts of the involucre 2- seriate, linear-lanceolate, 3 mm. long, acute, ciliate, pubescent ; eorolla 2.5-3 mm. long, the tube and throat pubescent ; rays 5— 6 mm. long, tube densely villous; achenes of the disk-flowers glabrous, epappose, ray-achenes cuneate-obovate, densely vil- lous, with a erown of long-villous silky hairs. Distribution: usually in shade of larger shrubs, hills and mountain slopes of Coahuila south to Zacatecas and Hidalgo. EXICO: COAHUILA—valley of Nazar, Bolson de Mapimi, May 11, 1847, Gregg M 631 (M); valley of Nazar, Bolson de Mapimi, eolleetion of 1848—49, Gregg 447 (G); Parras, 111 miles west of Saltillo, June 18-28, 1880, Palmer 592 (G, US); 1902, Palmer 279 (F, G, M, US); Parras, 1905, Purpus 1126 (F, U CAL); Parras, Oct. 6-11, 1898, Palmer 435 (G, M, U CAL, iind Carneros Pass, Sept. 7, 1889, hg 2402 (F, G, M, U CAL, US); La Ventura, Aug. 2-5, 1896, Ben 3930 Ps US); ZACATECAS—Cedros, northern Zacatecas, June, 1 1908, Kirkwood 67 (G); shade of shrubs, vicinity of Cedros, Aug., 1908, Kirkwood 227 (9); ; in shade of larger plants, such as mesquite, Cedros, Aug., 1909, Lloyd 124 (M TYPE); SAN LUIS POTOSI—between the states of Nuevo Leon and Matehuala, June 17-18, 1898, Nelson 4520 (US); region of San Luis Potosi, Sept. 1879, Schaffner 717 (286) (F, P ; HIDALGO— mountain slopes, Ixmiquilpan, Sept. 1905, Purpus 1334 (F, G, M, U CAL). 5. Zaluzania Grayana Rob. & Greenm. in Proc. Am. Acad. 34: 531. 1899; Woot. € Standl. in Contr. U. S. Nat. Herb. [Fl. N. Mex.] 19: 708. 1915; Blake in Contr. U. S. Nat. Herb. [Stand- ley’s Trees & Berube Mexico] 23: 1537. 1926. Gymnolomia triloba Gray in Proc. Am. Acad. 17: 217. 1882. Zaluzama Grayiana Rob. € Greenm. in Proc. Boston Soc. Nat. Hist. 29: 104. 1899. Suffruticose; stems from a lignescent rootstock, 4-8 dm. high, branched above, minutely hirtellous to glabrate; leaves petiolate, ovate to 3-lobed, 3-9 em. long, 1.5-6 em. broad, the lobes sparsely incised to incised-serrate, acute, minutely hirtel- lous on both surfaces; heads in loose corymbose clusters, 2— 3 em. broad including the rays, 6-8 mm. high; bracts of the in- voluere linear-lanceolate, 4 mm. long, minutely hirtellous; pales subcoriaceous, conduplicate, trifid at the apex, ciliate, hir- sutulous along the dorsal ridges; ray-flowers 7-10, rays ob- long, 0.8-1.5 em. long, disk-flowers 2 mm. long, the tube hirsutu- lous, the limb glabrous; achenes of the disk-corollas 2 mm. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 107 long, triangular, epappose, ray-achenes bearing a crown of a few short squamellose setae. Distribution: mountains of southern Arizona south to Chi- huahua. UNITED STATES: ARIZONA— peaks south of Rucker Valley, 1881, Lemmon (G TYPE); near Fort Huachuea, high peaks of Huachuea Mts., Aug.-Sept., 1882, Lem- mon 2764 (F, G, US); Huachuca Mts., July 3, 1884, Pringle 2145 (G, US); near Ft. Huachuca, Sept., 1894, Wilcox 469 (US); high peaks of Huachuca Mts., Aug.- Sept., 1882, Lemmon (US); north slope of Huachuca Mts., Carr Peak, Aug. 25, Goodding 864 (G); Apache Pass, Chiricahua Mts., Sept., 1881, Lemmon (M). MEXICO: CHIHUAHUA—shaded ravines, Mapula Mts., alt. 6000 ft., Oct. 30, 1886, Pringle 755 (F, G, M, US); shaded slopes, La Bufa Mt. above Cusihuiriache, Sept. 2, 1887, Pringle 1310 (F, G, US). 6. Zaluzania Pringlei Greenm. in Proc. Am. Acad. 39: 101. 1903; Blake in Contr. U. S. Nat. Herb. [Standley's Trees & Shrubs Mexico] 23: 1538. 1926. Plants shrubby ; stems of the more woody parts covered with a grayish bark, the ultimate branches purplish, striate, lenticu- lar, especially so below, strigose-puberulent above, becoming glabrate below; leaves petiolate, ovate, 3—5.5 cm. long, 1.5- 3.5 em. broad, short-aeuminate, acute or submucronate-acute, entire to crenate-dentate, strigose puberulent on both surfaces, petioles slender, 0.5-1.5 em. long; heads discoid, ovate, grouped in a terminal round-topped paniculate-eorymbose cluster, 5— 7 mm. high, 5-6 mm. wide; bracts of the involucre 3-seriate, ovate-lanceolate, about 3 mm. long, acute, strigose-puberulent ; pales coriaceous, pubescent; achenes 2 mm. long, glabrous, pap- pus absent. Distribution: limestone hills of Morelos. MEXICO: MORELOS—limestone hills near Jojutla, alt. 3000 ft., Oct. 18, 1902, iacu pam (F, G, M, U CAL, US). T. Zaluzania discoidea Gray in Proc. Am. Acad. 21: 388. 1886; Rob. & Greenm. in Proc. Am. Acad. 34: 534. 1899. Plants herbaceous from a lignescent base; stems 6-10 dm. tall, simple, terminated by a corymbose Auster, striate, pur- plish, velutinous ; leaves short-petiolate, broadly ovate, round- ovate to deltoid- d 4—12 em. long, 3-10 em. broad, irregu- larly erenate to crenate-undulate, obtuse to acute, subcordate [Vor. 22 108 ANNALS OF THE MISSOURI BOTANICAL GARDEN to broadly eordate at the base, palmately veined, pale green above, strigose-velutinous to strigose-hirtellous, whitish be- neath, velvety tomentose; heads discoid, in dense corymbose clusters, subglobose, 5-7 mm. broad, 6 mm. high; bracts of the involucre herbaceous, ovate-lanceolate, about 4 mm. long, acute, hirtellous; pales coriaceous, acute, fimbriate; corolla-tube minutely puberulent; achenes about 2.5 mm. long at maturity, glabrous, epappose. Distribution: rocky hills and mountains of Chihuahua and Durango. MEXICO: CHIHUAHUA—mountains near Chihuahua, Sept. 12, 1886, Pringle 1110 (M, U CAL); rocky hills near Chihuahua, Oct. 19, ip bins id 309 (F, G, US); e“ Qosiquiriache, ’’ Sept. 25, 1846, Wislizenus 189 (M); DURANGO—EI Oro to Guana- cevi, Aug. 14-16, 1898, Nelson 4727 (US); along road Pre: ** Cerro Prieto and La Providencia," Sept. 11, 1898, Nelson 4970 (US). 8. Zaluzania montagnaefolia Schz.-Bip. in Flora 44: 563. 1861; Hemsl. Biol. Cent.-Am. Bot. 2: 160. 1881; Blake in Contr. U. S. Nat. Herb. [Standley's Trees & Shrubs Mexico] 23: 1538. 1926. Ferdinanda montagnaefolia Schz.-Bip. in Koch's Berl. Allgem. Gartenz. 179. 1858. Zaluzania asperrima Schz.-Bip. in Flora 47: 218. 1864; Hemsl. Biol. Cent.-Am. Bot. 2: 159. 1881; Rob. & Greenm. in Proc. Am. Acad. 34: 532. 1899. Zaluzania asperrima var. montagnaefolia (Schz.-Bip.) Rob. & Greenm. in Proc. Am. Acad. 34: 532. 1899 Plants shrubby; stems glabrate below, strigose-hirtellous above; leaves ovate, 2-7 em. long, 1-3 em. broad, serrate or crenate-serrate to often entire above, aeuminate to acuminate- acute, base decurrent-attenuate, auriculate, leaves of corym- bose branches sessile and auriculate, above dark green, pustu- late- hispidulous to pustulate-hirsute, beneath light green, densely resinous-glandular and hirsute especially on the veins; heads in corymbose clusters, 1-1.5 cm. broad including the rays; bracts of the involucre 3-4-seriate, ovate, 3-4 mm. long, ciliate, acute to acuminate, hirsutulous; pales subcoriaceous, strongly incurved and united above, the apex resembling the prow of a canoe; ray-flowers styliferous and fertile, 6-8 mm. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 109 long including the achene, base of tube capping the achene; achenes 1.5-2 mm. long, disk-achenes obeompressed-quadran- gular, ray-achenes trigonous, glabrous, epappose. Distribution: dry hills and barrens of Puebla and Oaxaca. MEXICO: PUEBLA—‘‘dry hills,’’ Tlacuilotepec, Aug., 1909, Purpus 3824 (F, G, M, U CAL, US); barrens near Tlacuilotepee, May, 1909, Purpus 3823 (F, G, M, US); Tehuacan, Sept., 1911, Purpus 5610 (F, M, U CAL); near Tehuacan, Aug. 1, 1901, Rose $ Hay 5954 (US); near Tehuacan, Nov. 7, 1903, Holway 5347 (G); **in vicinity of San Luis Tultitlanapa near Oaxaca,”” Aug., 1908, Purpus 2530 (F, G, M, U CAL, US); "o Po Alban,’’ near Oaxaca, alt. 5800 ft., Oct. 4, 1894, DEA 4928 (G, M, U CAL, US); ** Monte Alban near Oaxaca City,’ alt. 5500-5600 ft., Oct. 2, 1894, Buen, 612 (M); ““Distrito de Tlacolula, Magdalene Ecatepac,?” vt 1700 m., Oct. 23, 1924, Conzatti 4601 (US); **Cerro de Trujono,’’ Conzatti 2317 (F). 9. Zaluzania cinerascens Schz.-Bip. in Flora 47: 219. 1864; Blake in Contr. U. S. Nat. Herb. 26: 249. 1930. Zaluzama Coulteri Hemsl. Diagn. Sp. Nov. 33. 1879; Biol. Cent.-Am. Bot. 2: 159. 1881; Rob. € Greenm. in Proc. Am. Acad. 34: 533. 1899; Blake in Contr. U. S. Nat. Herb. [Stand- ley’s Trees & Shrubs Mexico] 23: 1538. 1926. Stems suffrutescent at base, apparently simple to the leafy- braeted corymb, conspicuously striate, densely strigulose- hirtellous; leaves broadly ovate to broadly ovate-elliptical, above dark green, strigulose-puberulent, beneath pale green, densely strigose-hirtellous ; main cauline leaves 4-13 em. long, 2-9 cm. broad, becoming smaller above, entire or sparsely ser- rulate, obtusely acute; heads in corymbose clusters, when mature 1.5-2 em. brad including the rays, 6-7 mm. high; in- volucre subhemispherical; bracts oblong-ovate, 4-5 mm. long, acute to obtusish, strigose-pubescent; pales truncate-obtuse, pubescent towards the apex; ray-flowers broadly oblong to ob- long, about 1-1.2 em. long including the achene, 4-6 em. broad, puberulent on the outer surface; achenes 2 mm. long, glabrous, epappose. Distribution: fir forests of the Sierra de Pachuca Mts. of Hidalgo at an elevation of 9000 ft. or above. MEXICO: HIDALGO—Real del Monte, Coulter 350 (G TYPE); ““El Chico,”” Sept., 1929, Lyonnet 387 (US); Sierra de Pachuca, alt. 9500 ft., Oct. 6, 1899, Pringle 7923 (G, US); Sierra de Pachuca, alt. 9500 ft., Aug. 28, 1906, Pringle 13784 (G, US); [Vor. 22 110 ANNALS OF THE MISSOURI BOTANICAL GARDEN fir forests, Sierra de Pachuca, alt. 9500 ft., Aug. 26, 1902, Pringle 999? (F, G, M, US); Sierra de Pachuca, alt. 9000 ft., Aug. 13, 1898, Pringle 6956 (F, G, M, U CAL, US). 10. Zaluzania megacephala Schz.-Bip. in Flora 44: 563. 1861; Ibid. 47: 218. 1864; Hemsl. Biol. Cent.-Am. Bot. 2: 159. 1881, in synonymy; Rob. € Greenm. in Proc. Am. Acad. 34: 533. 1899; Blake in Contr. U. S. Nat. Herb. [Standley's Trees & Shrubs Mexico] 23: 1538. 1926. Ferdinanda augusta var. megacephala Schz.-Bip. in Flora 44: 563. 1861. Plants herbaceous from a perennial rootstalk; stem 6-10 dm. tall, unbranched except in the inflorescence, striate, glabrate below, densely strigose-hirtellous or velutinous above; leaves lanceolate to ovate-lanceolate, 6-10 em. long, 2-5 cm. broad, serrate, acute, sessile or subsessile, above green, strigose-hir- tellous, beneath cinereous, densely strigose-hirtellous to velu- tinous; heads in corymbose clusters, 1.5-2.5 em. broad includ- ing the rays, 0.8-1.2 em. high; bracts of the involucre lanceo- late, 4-5 mm. long, ciliate, acute to obtuse, puberulent; pales about 5 mm. long, apex acute, slightly ineurved; ray-flowers broadly to narrowly oblong, 8-10 mm. long, 3-4 mm. broad; achenes glabrous, 3 mm. long, epappose. Distribution: mountains of Coahuila. MEXICO: COAHUILA—mountains near General Zepeda, alt. 6500 ft., Oct. 7, 1905, Pringle 10076 (F, G, M, U CAL, US); high summits, near Carneros Pass, alt. 10000 ft., Sept. 8, 1889, Pringle 2398 (F, G, M, U CAL, US); elevated portion of Sierra Madre, 12-14 leagues south of Saltillo, July 25-Aug. 1, 1880, Palmer 734 (F, G, US). 11. Zaluzania mollissima A. Gray in Proc. Am. Acad. 15: 35. 1880; Hemsl. Biol. Cent.-Am. Bot. 2: 160. 1881; Rob. & Greenm. in Proc. Am. Acad. 34: 532. 1899; Blake in Contr. U. S. Nat. Herb. [Standley's Trees & Shrubs Mexico] 23: 1538. 1926. Shrubby ; stems branched, about 0.5-1.5 m. high, bark gray- ish-brown, tomentulose above, the more woody parts glabrate; leaves ovate, ovate-lanceolate to oblong-lanceolate, sometimes tending to become trilobed, 2-5 em. long, 0.5-2.2 cm. broad, en- tire, obtuse, acute at the base and decurrent on the petiole, 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 111 above dark green, tomentulose, beneath cinereous, densely tomentulose ; heads three to several in spreading corymbose or paniculate clusters, 2-3 em. in diameter including the rays, pe- duncles 1-7 cm. long; bracts of the involucre oblong-ovate, about 4 mm. long, obtuse to acute, velvety tomentose; pales cartilaginous, slightly trilobate; ray-flowers oblong-lanceolate, 1-1.5 cm. long, 3-4 mm. broad, rays bidentate; achenes of the disk glabrous, epappose, ray-achenes slightly pubescent, bear- ing a crown of short hirsute hairs. Distribution: in moist soil in fields or among the hills, from Zacatecas to San Luis Potosi. MEXICO: ZACATECAS—low places, especially where there is relatively abundant moisture, Cedros, May, 1908, Lloyd 129 (US); hills, Cedros, June, 1908, Lloyd 32 ey vicinity of Cedros, id Kirkwood 129 (F); near Concepcion Del Oro, Aug. 904, Palmer 314 (F, G, M, U CAL, US); SAN LUIS POTOSI—vieinity of San m Potosi, Aug., 1878, pee 446 (F, G TYPE, M, US); in fields near ‘‘San Miguel,’’ Aug., 1876, Schaffner 767 (342) (F, G, US). 12. Zaluzania subcordata W. M. Sharp, sp. nov.'* Plants suffruticose; stems branched above, about 0.5-2 m. high, the bark grayish, glabrate below, densely subtomentulose in the inflorescence; leaves broadly ovate to ovate-lanceolate with slender petioles, 1.5-6 em. long, 0.6-2.5 em. broad, entire, obtuse, subeordate, above dull green, subtomentose, beneath whitish, densely velvety tomentose, subcoriaceous; heads in close corymbose clusters, 1-1.5 em. broad including the rays, 6-8 mm. high, peduncles short, 0.54 em. long; bracts of the involuere broadly oblong-lanceolate, 3—4 mm. long, densely tomentulose, whitish; pales trilobed at the summit and fim- briate-ciliate, the outer upper surface pubescent; ray-flowers ovate to oblong-ovate, 5-8 mm. long, 34.5 mm. wide, rays faintly bi- or tri-dentate; mature achenes 2-2.5 mm. long, “2. subcordata Sharp, sp. nov. Plantae i Ec es, caulibus supra ramosis, flor 0.5-2 m. altis, infra glabratis, dense subtomentosis in inflorescentia; foliis late ovatis vel ovato-lanceolatis, 1.5-6 em. longis, 0.6-2. = em. latis, integris, alaisi; sub- cordatis, sup bscure viridibus, subtus albi idis, dense velutino-tomentosis, sub- coriaceis, petiolis gracilibus; capitulis corymbosis, 1-1.5 em. latis ineludentibus radiis; radiis ovatis vel oblongo-ovatis, 5-8 mm. longis, 3-4.5 mm. latis.—Collee on rocky hills, Esperanza, Puebla, Mexico, 1907, Purpus 2580 (F, G, M TYPE, U CAL). [Vor. 22 112 ANNALS OF THE MISSOURI BOTANICAL GARDEN disk-achenes rhomboidally 4-angled, glabrous, epappose, ray- achenes trigonal, pubescent on the ridges, bearing a more or less definite fimbriate-setose or lacerate-setose pappus. Distribution: rocky or barren hills and mountains of south- eastern Puebla. MEXICO: PUEBLA—rocky hills, Esperanza, Aug., 1907, Purpus 2580 (F, G, M TYPE, U CAL, US); barren hills about Esperanza, alt. 2660 m., Aug. 17, 1905, Pit- tier 452 (US); Esperanza, alt. 2450 m., Nov. 16-19, 1907, Arséne 7093 (US); Mt. Orizaba, alt. 9000 ft., Aug. 9, 1891, Seaton 286 (F, G, US); vieinity of San Luis Tultitlanapa near Oaxaca, Aug., 1908, Purpus 3027 (F, G, M, U CAL, US); Coxeat- lan, alt. 8000-9000 ft., Sept., 1909, Purpus 4121 (U CAL). 13. Zaluzania augusta (Lag.) Schz.-Bip. in Flora 44: 562. 1861; Rob. & Greenm. in Proc. Am. Acad. 34: 532. 1899; Blake in Contr. U. S. Nat. Herb. [Standley's Trees & Shrubs Mexico] 23: 1537. 1926. Ferdinanda augusta Lag. Gen. € Sp. Nov. Pl. 31, pl. 2. 1816; DC. Prodr. 5: 552. 1836. Anthemis lutescens La Llave € Lexarza, Nov. Veg. Desc. 1: 26. 1824. Chrysophania fastigiata Kunth in Less. Syn. Gen. Comp. 224. 1832; DC. Prodr. 5: 553. 1836. Ferdinando lutescens (La Llave & Lexarza) DC. Prodr. 5: 553. 1836. Zaluzania angusta Schz.-Bip. in Hemsl. Biol. Cent.-Am. Bot. 2: 159. 1881. Plants fruticose; stems 1.5-2.5 m. tall, bark exfoliating, stri- gose-velvety and glandular on the branches above; leaves ovate to ovate-lanceolate, 2-8 cm. long, 1-3.5 em. broad, sessile to subsessile above, short-petiolate below, above densely stri- gose-velutinous, beneath densely cinereous-velvety, lower cau- line leaves crenate-serrate, acute at the base, those of the branches entire, acute to acuminately acute; heads numerous, in corymbose clusters, 5-7 mm. broad not indading the rays; involucre campanulate, bracts ovate-lanceolate, about 3 mm. long, ciliate, acute, finely strigose; pales subeoriaceous, apex acute and limbriate-lacerato to somewhat entire; ray-flowers 7-8 mm. long, oblong, bidentate ; achenes about 2 mm. long, ray- achenes trigonal, bearing a erown of squamulose setae, disk- achenes rhomboidal in cross-section, glabrous, epappose. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 113 Distribution: rocky mountain slopes and hills, Guanajuato east to Hidalgo and south to Mexico. MEXICO: GUANAJUATO—''vicinity of Guanajuato, ”” 1895, Duges 469 (G rd de las companas,’’ alt. 1850 m., July 9, 1914, Arsène 10065 ‘a M, US); collected between Cadereyta and Visaron n, Aug. 22, 1905, Rose, Painter $ Hiss 9738 (G, US) ; vicinity of Queretaro, alt. 1850 0 m., 1909-12, Arséne $ Nicolas 6162 (G, M, US); HIDALGO—between Pachuca and Real del Monte, Aug. 31, 1903, Rose $ Painter 6675 UA near Paehuea, 1899, Duges (US); roeky mountain slopes, near su Sept., 1905, Purpus 1539 (F, G, M, U CAL); hills near Tula, 680 , Oct. 14, 1902, Pringle 9996 (F, G, M, US); mExıco—hills near Lecheria urs alt. 7500 ft., Nov. 18, 1903, Pringle 11614 (G, US); hills near Lecheria, alt. 7600 ft., Aug. 23, 1904, goin PA (F, G, US); Rio Hondo Canyon, Aug. 22, 1890, Pringle 3144 (F, G, M, U CAL, US); near Tlalnepantla, uly 6, 1905, Rose, Painter $ Rose 8418 (US); ES near Miss T: Sept. 5, 1897, Pringle 7451 (M); ‘‘near Guadelupe,’’ Valley of Mexico, Sept. 23, 1903, Rose, Painter $ Rose 7278 (US); ‘‘near Guadelupe,’’ Aug. 24, 1865-66, en. 808 (G, US). 14. Zaluzania Sodiroi Hieron. in Engl. Bot. Jahrb. 29: 35. 19 Zaluzania nonensis Hieron. l.c. 29: 36. 1900. Zaluzama quitensis Hieron. l. c. 29: 37. 1900. Plants suffruticose; stems 1-2 m. tall, brownish to purplish pilose-tomentulose, besoming erisp-hirtellous to glabrate; leaves deltoid-ovate, ovate, to ovate- lanceolate, 4-10 em. long, 2-6 em. broad, triple-nerved, dentate-mucronate to subentire, acute, base cuneately decurrent on the petiole, above dark green, rugose, scabrate-hirsute, beneath light green, conspicu- ously reticulate, densely crisp-hirtellous on the veins, globular- resiniferous; heads in corymbose to subcorymbose clusters, 1— 1.2 cm. broad including the rays, pedicels 1-2 cm. long; bracts of the involucre triangulate-ovate, obtuse, erisp villous-tomen- tulose; ray-flowers yellow, oblong, 5-7 mm. long; achenes rhom- boidal in cross-section, linear, 2.5-3 mm. long, slightly ineurved atthe base, glabrous, pappus absent. Distribution: Ecuador. SOUTH AMERICA: ECUADOR—arid interandean hills, Sodiro 31/1 (US tsorypr) ; near Nono, ‘‘in silvis subandinis,’’ Sodiro 31/2 (US); in hills near Quito, Sodiro 31/3 (US); 4 ft. high, Aug. 28, 1920, near Quito, Holway 948 (US); Ficoa, near 91 , of Pichincha, between ‘‘La Ma agdalena poi Chillogallo,’’ alt. 2800 m., July 14, 1927, Firmin 140 (US); Provinee of Pic cha, between ‘‘La Ma, ER and Chil- logallo,’’ alt. 2810 m., April 19, 1928, a 471 (US); Province of Chimborazo, [ VoL. 22 114 ANNALS OF THE MISSOURI BOTANICAL GARDEN alt. 3000 m., Andre K1245 (NY); near Quito, Aug. 1898, Mille 558 (NY); near Quito, Aug., 1898, Mille 568 (NY); frequent near ‘‘Pifo,’’ alt. 2600 m., 1895, Mille 569 (US). BALSAMORHIZA Hooker History —Hooker* in “Flora Boreali-Americana’ in 1834 de- seribed two species of Heliopsis, namely, H. ?balsamorhiza and H. ?terebinthacea. However, he doubted their generic posi- tion and proposed Balsamorhiza as a more appropriate name, due to the abundance of a limpid juice, similar to the resin of Pimus balsamea, which exuded from their freshly cut or broken roots. The proposal of this genus is accompanied by a narra- tion of its outstanding characters. DeCandolle? and End- licher? treated Balsamorhiza as a subgenus under H eliopsis. Nuttall* in 1841 gave the first comprehensive study of the genus. Following his generic description he included eight species, namely, Balsamorhiza Hookeri, B. terebinthacea, B. sagittata, B. helianthoides, B. hirsuta, B. incana, B. macro- phylla, and B. deltoidea; the four last being new species. Bal- samorhiza sagittata and B. helianthoides were originally de- scribed by Nuttall® under Espeletia. All the above-named species remain valid except B. helianthoides which is synony- mous with B. sagittata. The genus was divided by Nuttall into two subgenera, namely, Artorhiza and Eubalsamorhiza. Torrey and Gray? and Walpers? followed Nuttall's treat- ment in that they recognized about seven or eight valid species as well as the two subgenera Artorhiza and Eubalsamorhiza. In 1848 Balsamorhiza Careyana was published by Asa Gray? in *Memoirs of the American Academy.” In his *Synop- tical Flora” in 1884 he recognized eight species and one va- riety. He also introduced the subgenus Kalliaetis for those 1 Hooker, W. J., Fl. Bor.-Am. 1: 310. [1834] 1840. ? DeCandolle, A., Prodr. 5: 551. 1836. * Endlicher, S., Gen. Pl. 408. 1838. * Nuttall, T., in Trans. Am. Phil. Soc. II, 7: 349. 1841. * Nuttall, T., in Jour. Acad. Nat. Sci, Phila. 7: 38-39. 1834. * Torrey, J., & Gray. A., Fl. N. Am. 2: 300. 1842. * Walpers, G. G., Rep. Bot. Syst. 2: 610. 1843. * Gray, A., in Mem. Am. Acad. [Pl. Fendl.] N.S. 4: 81. 1848. ? Gray, A., Syn. Fl. N. Am. 1*: 265. 1884. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 115 species possessing chartaceous rays. This character is not always constant, and therefore his subgenus becomes synony- mous with Artorhiza. Between 1890 and 1930 four new species were added to Balsamorhiza from the northwestern United States. GENERAL MORPHOLOGY Roots.—Balsamorhiza develops a fusiform, perennial tap- root with a rough, corrugated, bark-like covering. The inner structure is usually composed of a soft, spongy, woody tissue which secretes a terebinthine, resinous juice when freshly cut or broken. The older roots have a tendeney to become more woody. The younger roots of some species, especially in late winter or early spring, have been used as articles of food. The greatest root development is found in the subgenus Artorhiza. The tap-root is often three feet in length, as in B. sagittata and B. deltoidea. In these species the crown often de- velops several underground, horizontal branches. In Eubal- samorhiza the root system is less developed. It consists of a slender tap-root, unbranched above, and usually less than two feet in length. The extensive tap-root system, according to ecological investigations, often extends six feet or more below the surface of the soil, and enables these plants to occupy dry open uplands. Stem.—1t is difficult to say definitely from external examina- tion just where the root ends and the underground stem begins, but aecording to stelar evidence the crown of the tap-root may be designated as stem. The length of this, however, varies from a few to several inches. The lateral underground branches which are developed especially in B. sagittata and B. deltoidea originate from a region near the main crown. The development of several such branches from one root gives the plant a close bushy or clumpy habit. In a majority of the species of Balsamorhiza the crowns are covered by a dense tuft of persistent, fibrous petiole bases of previous seasons. The aerial, erect or ascending stems develop each season from the underground crowns. The average height is usually 2-4 dm. The minimum is about 1 dm., as in B. serrata and B. [Vor. 22 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN hispidula, and the maximum height of 8-10 dm. is often at- tained in B. macrophylla. The stems are usually subseapose, that is, almost entirely leafless except for a pair of small leaves near the base, as ex- emplified by a majority of the species of Eubalsamorhiza. The species with leafy or bracteate stems occur in Artorhiza. Leaves.—The leaves are chiefly opposite, sometimes alter- nate. Two types of leaf outline are manifest, namely, (1) the ovate to ovate-cordate type, and (2) the oblong-lanceolate pin- nate or bipinnatifid type. The name Artorhiza has been as- signed to the former group, and Eubalsamorhiza to the latter. In the section Artorhiza the leaves are either basal or cau- line; the cauline alternate types are exemplified in B. Bolanderi and B. invenusta with ovate or ovate-elliptical outlines. The remaining species, B. sagittata, B. deltoidea, and B. Careyana, have large, ovate-cordate basal leaves, and bracteate subop- posite cauline leaves. The leaf margins in this group are en- tire, undulate, or dentate with acute apices. The herbaceous texture is common, but in B. Bolanderi the leaves become cori- aceous and conspicuously reticulated on both surfaces. The subcoriaceous texture is also approached in the more mature leaves of B. deltoidea and B. Careyana, but the venation never becomes so prominent as in B. Bolanderi. The section Eubalsamorhiza is characterized by a pinnate or bipinnate type of leaf. The pinnate form is the more common; the bipinnate type is characteristic of B. hirsuta, and to a less degree of B. macrolepis. The subentire leaf is also present in this section. Both pinnate and undivided leaves may occur on the same plant, as in B. serrata. In this species the undivided leaves are more common, but in B. incana var. tomentosa entire or incised leaves are produced. Subentire blades also oceur oc- casionally on the outermost leaves of B. macrophylla. The leaves in this section are mostly basal, a pair of small ones be- ing produced on the lower part of the scapose stem. Pubescence.—The pubescence within the genus is quite di- verse—pilose, tomentose, hirsute, velutinous, and sericeous be- ing the common forms. The hairs in a majority of the species 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE HT are interspersed by sessile or short-stipitate glands. Fre- quently the pubescence on the stems is of the pilose nature, in- terspersed by numerous glands. Balsamorhiza incana and its variety tomentosa have a covering of long soft tomentum. The usual type of pubescence, however, is the hirsute, hirsute-his- pid, or subsericeous. Inflorescence.—' The inflorescence in Balsamorhiza consists chiefly of solitary heads which terminate the naked scapose stems. "They may be axillary, as in B. invenusta, or in raceme- like clusters as in B. Careyana. The inflorescence offers no marks of specific importance except in the latter species. Involucre.—The bracts of the involucre often present im- portant characters for species differentiation, partieularly in outline, length with reference to the disk, pubescence, texture, and venation. The linear-lanceolate to ovate-lanceolate form of involucral bract is the most common. Involucral bracts of this type usu- ally do not exceed the disk in length. An ovate form of bract abruptly constricted above the middle and attenuated to an acute apex is characteristic of Balsamorhiza hirsuta and to a less degree of B. incana. The bracts of the involucre which ex- ceed the disk in length are foliaceous and mostly oblong or ob- long-lanceolate in general outline, as in B. macrophylla, B. del- toidea, and B. macrolepis. Balsamorhiza Bolanderi is char- acterized by large, ovate, coriaceous bracts which become gla- brate and conspicuously reticulated on both surfaces. The margins of the bracts are entire except in B. macrolepis where they are often sparsely serrated or incised. The pubescence of the involucre is of four sorts, namely, (1) densely tomentose, (2) crisp-pilose, (3) subvelutinous or vel- vety pubescent, and (4) hirsute or hispid. The densely white- tomentose type of pubescence is exemplified in B. sagittata and B. incana and its variety tomentosa. Balsamorhiza serrata and B. hirsuta var. lagocephala are the best examples of a dense crisp-pilose-pubescent involucre. A majority of the species, however, are crisp-pilose-ciliate. The velutinous or subsericeous pubescence is characteristie of B. macrolepis and [Vor. 22 118 ANNALS OF THE MISSOURI BOTANICAL GARDEN B. Hookeri and its varieties. The hirsute, hispidulus to gla- brate condition is also common within the genus. Receptacle and Pales.—The receptacle in Balsamorhiza is broadly convex. The pales are conduplicate and acuminate or acuminate-acute, becoming stiff in the mature heads. They are persistent on the receptacle after the fall of the achenes. Neither the receptacle nor the pales afford any marks that may be used in specific differentiation, but only generic characters. Corollas.—The heads in Balsamorhiza are heterogamous with styliferous and fertile ray-flowers, and usually fertile tubular disk-flowers. Balsamorhiza invenusta with eradiate heads, however, is an exception. The corollas in both the ray- and disk-flowers are yellow except in Balsamorhiza Hookeri var. rosea in which the rays are occasionally purple. The rays are ovate to oblong, subentire, or 2-3-denticulate, stylif- erous, and aecompanied by one or more staminodea. The disk- corollas have a narrow eylindrical tube which expands above into the tubular-campanulate 5-lobed limb. The stamens are very similar to those described for preced- ing genera. The style-branches are slender, terete in cross- seetion, and more or less hirsutulous from base to apex. Achenes.—The achenes, for the most part, are similar throughout the genus. "They are linear-oblong, 5-8 mm. long, and usually glabrous except in Balsamorhiza Careyana and B. Hookeri var. rosea which are pubescent. The disk-achenes are quadrangular, as a rule, with a small rib alternating with each of the four larger angles. Balsamorhiza invenusta, however, has rhomboidal disk-achenes and trigonal ray-achenes. The pappus is absent, and only in rare cases in B. sagittata has there been observed one or two subulate bristles on each achene. Geographical Distribution.—The genus Balsamorhiza is con- fined to the northwestern United States. Its center of distribu- tion extends from west-central Wyoming and northern Utah west to southern Washington and northern California. Plants of this genus are mostly inhabitants of dry, usually rocky, open grounds, the deep tap-root system and the low habit enabling them to oceupy dry situations. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 119 The subgenus Artorhiza is confined chiefly to the Pacific Coast states, with the exception of B. sagittata. This species ranges from the western Dakotas and Colorado west to British Columbia and California, being the most widely distributed species of the genus. Balsamorhiza deltoidea is the species with the next widest range, extending from British Columbia southwards to Los Angeles County, California. The remain- ing species of this section are less widely distributed; for ex- ample, B. Careyana occurs on dry ground from central and southeastern Washington south to central and northeastern Oregon; while B. Bolanderi and B. invenusta are confined to central California. Eubalsamorhiza comprises the greater majority of the spe- cies in this genus, but none of the species has a very wide dis- tribution. Balsamorhiza incana, having a range from western Wyoming and southwestern Montana west to northeastern Oregon, is the most widely distributed species of the section. Balsamorhiza macrophylla and B. hispidula occur from west- ern Wyoming and southern Idaho south to Utah and eastern Nevada. The majority of the remaining species are confined to the Pacific Coast region from Washington south to central California. TAXONOMY Balsamorhiza [ Balsamorrhiza of authors] Hook. Fl. Bor.- Am. 1: 310. [1834] 1840; Nutt. in Trans. Am. Phil. Soc. II. 7: 349. 1841; Torr. € Gray, Fl. N. Am. 2: 300. 1842; Walp. Rep. Bot. Syst. 2: 610. 1843; Benth. & Hook. Gen. Pl. 2: 366. 1873; Gray, Syn. Fl. N. Am. 1?: 265. 1884; Hoffm. in Engl. € Prantl. Nat. Pflanzenf. IV. Abt. 5: 234. 1890; Howell, Fl. Northwest Am. 1: 339. 1900; Coult. & Nels. Bot. Rocky Mts. 545. 1909; Piper & Beattie, Fl. Northwest Coast, 376. 1915; Rydb. Fl. Rocky Mts. 928. 1917; Jepson, Man. Fl. Pl. Cal. 1077. 1925; Blake in Contr. U. S. Nat. Herb. [Tidestrom’s Fl. Utah € Nevada] 25: 582. 1925; Rydb. Fl. Pr. € Pl. Cent. N. Am. 839. 1932. Roots perennial, fusiform, thick, often becoming very stout in older plants, covered by a rough or corrugated bark. Stems (Vor. 22 120 ANNALS8 OF THE MISSOURI BOTANICAL GARDEN herbaceous, ereet or ascending, leafy, subscapose or scapose, usually unbranched, glabrous or variously pubescent. Leaves opposite or alternate, usually basal, sometimes cauline, ovate- cordate to oblong-lanceolate in general outline, entire, coarsely dentate, incised, pinnately divided to bipinnatifid, acute or acutish, variously pubescent or glabrate. Heads radiate or rarely discoid, usually solitary, sometimes two or more in raceme-like clusters. Bracts of the involucre 2-4-seriate, her- baceous or coriaceous, linear-lanceolate, ovate-lanceolate, ovate-attenuate to oblong, the outer series often foliaceous and longer than the disk, puberulous to densely tomentose. Re- ceptacle broadly convex; pales firm, subcoriaceous, condupli- cate, persistent after the fall of the achenes. Disk-flowers light yellow, tube eylindrical, limb tubular to tubular-campanulate, 5-lobed, the base often more strongly developed on one side, glabrous; anther appendages ovate, base sagittate; style- branches slender, attenuate, hirsutulous from base to apex. Rays usually present, fertile, yellow, rarely purplish, oblong- lanceolate, usually 2-denticulate at the apex. Achenes of the disk-flowers fertile, epappose, quadrangulate to subquadrangu- late, glabrous or rarely canescent-pubescent, those of the ray- flowers trigonal, otherwise as in the disk-achenes. Type species: Balsamorhiza Hookeri (Hook.) Nutt. in Trans. Am. Phil. Soc. IT. 7: 349. 1841 (Heliopsis? Balsam- orrhiea Hook. Fl. Bor.-Am. 1: 310. [1834] 1840). KEY TO THE SPECIES AND VARIETIES A. Leaves on or dentate-serrate, cordate-ovate, deltoid-ovate, ovate or ovate- olat B. ibus leafy to inflorescence; leaves alternate, basal and cauline similar in size. C. Heads discoid; bracts of the involucre lanceolate and -— about ong as the disk.........o.ooooooooooomoomoconsos».os . B. invenusta CC. Heads radiate; braets of the involuere ovate to ovate- eiue foli- us and much longer than the disk................... 2. B. Bolanderi BB. "hel oe bracteate; leaves Salt, basal leaves large, those of the stem reduce aves ida on both surfaces, seabrous to ees or hispid; voluere green, glabrate, or hirsute to densely erisp-p E. Leaves eordate-ovate to deltoid-ovate, entire or dena dentate, not conspicuously reticulated. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 121 F. Outer bracts of the involucre lanceolate, seldom exceeding the disk; disk 0.8-2.5 em. broad; achenes usually einereous-pubes- BIO 6s, eee evite reins eie i o True im VUES 3. B. Careyana FF. ou bracts of the involucre oblong or E lanceolate, ex- ceeding the disk; disk 2.5-4 em. broa aehenes gla- A A A | . deltoidea EE. Leaves lanceolate to ovate-lanceolate, dentate- or MS n od erous, n M reticulilod aa ee . B. serrata DD. Leaves silve e-tomentose beneath; involuere Ls per- sistently woolly tomentose. G. Leaves cordate-ovale..... 21... cies ele cles 5. B. sagittata GG. Leaves oblong- ach ge oblong- turis .e.oo.opPo.o.pposSsS$2.( ........... o... .o.o»oosso a. B. incana var. tomentosa AA. Leaves incised, pinnate or bipinnate, lanceolate 2 oblong-laneeolate in general outline. H Leaves canescent or hoary beneath, velutinous, serieeous to tomentose. I. Leaves velutinous, subsericeous to sericeous. J. Bracts of the involucre not exceeding the disk, entire. . Bracts of the involucre linear-lanceolate, Jidesofate to linear-oblong, 2-4 mm. broad, X v2 green- oon IES LE . B. Hookeri KK. Bde of the involuere triangular-aeute A ovate- cute, m. broad, usually brownish or hoary a apex, Miis iine O AAN E 7. B. platylepis JJ. m of the involuere rt ie the disk, tn dentate or incised towards the apex.............. macrolepis ll. Leaves lanate, floecose- hse to tomento Leaves lanate ; segments n bla pinnatel parted; bracts of the involucre u ...o. o. .o...s.o eo ooo. o. ]7] 0] .S$.c.r..o oo. serrate-incised; bracts of the involucre ovate-at- ate 9. B. incana HH. Leaves green on both surfaces, hispid, hirsute to crisp-pilose. M. Leaves 3-6 dm , erisp-hirsute, erisp-pilose to pilose-tox n e. prada css 5-11 em. long, entire, erisp- nip ; braets of the involucre longer than e di e d glabrous....10. B. macrophylla NN. oe -segm 3-6 cm. ‘oie, serrate-dentate or "sn mn to pilose-tomentose ; bracts of the involucre ng as or shorter h the T | to pilose- entose. 2 S MM. Dim 0.5-4. 5 = E hispid, sein ical hirsute [VoL. 22 122 ANNALS OF THE MISSOURI BOTANICAL GARDEN O. Achenes pubescent....6a. B. Hookeri var. rosea OO. Achenes glabrous. P. Leaves hispid to hispid-pilose. Q. Leaves subcoriaceous, segments serrate- subuliferous; bracts of the involucre subuliferous, pilose-tomentose....... QQ. Leaves less firm in texture, segments entire, incised or cleft, acute; bracts of the involuere acuminate-acute, his- pid-pilose-ciliate......... 12. B. hispidula PP. Leaves hirsute to strigose-hirsute. R. Involucre densely tomentose........ TIT 13a. B. hirsuta var. lagocephala RR. Involucre not lireuto cq S. Braets of the involucre ovate, abruptly attenuated into slender, erect apices....13. B. hirsuta SS. Bracts of the iin linear-lan- ceolate to ovate-lanceolate, not at- tenuated, acute. T. Leaves cleft or FPE the lobes entire......... 4. B. terebinthacea TT. Leaves dl the segments usually eleft or incised....... TOIT 13b. B. hirsuta var. neglecta 1. Balsamorhiza invenusta (Greene) Coville in Contr. U. S. Nat. Herb. 4: 130. 1893; Jepson, Man. Fl. Pl. Cal. 1077. 1925. Helianthus invenustus Greene, Pittonia, 1: 284. 1889. Stems leafy, erect or ascending, 3-6 dm. tall, pilose-hispid, becoming hispidulous below and densely glandular above; leaves petiolate, ovate, ovate-elliptical to ovate-lanceolate, 1- 2 dm. long, 4-9 em. broad, entire to undulate, acute or acumi- nate, acutish to subcordate at the base, above scabrous to his- pid-hirsute, the midrib often pilose-hispid, beneath glandular, hispid to hirsute-pilose on the veins, the petioles hispid-pilose ; heads discoid, solitary, terminal or sometimes axillary, 2.5- 4 em. broad; bracts of the involucre about as long as or longer than the disk, oblong-lanceolate, pilose-hirsute-ciliate, glandu- lar on both surfaces, occasionally sparsely hirsute-hispid along the midvein; disk-corolla about 1 em. long, the throat tubular glabrous; mature achenes rhomboidal in cross-section, 7-8 mm. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 123 long, the crown developing knob-like projections at the angles, glabrous. Distribution: forests of south-central California, Fresno, Tulare, and Kern Counties. CALIFORNIA: Milwood, Fresno Co., July 23, 1892, Brandegee (U CAL); Sequoia Nat. Park, Tulare Co., July, 1908, Davidson 2058 (U CAL); ‘ “giant forest,’’ Tulare Co., July, 1905, Brandegee (U CAL); North Fork of Middle Fork of Tule River, eastern Tulare Co., July 11, 1908, Hall 8340 (U CAL); South Fork of ah River, Tulare Co., July 21, 1904, Culbertson (F, G, M, NY, U CAL); mountains of Kern Co., 1888, Palmer 105 TYPE. This leaf y-stemmed species of Balsamorhiza is distinguished by its discoid heads, rhomboidal achenes, and tubular corolla. Its general habit resembles that of Wyethia ovata Gray. 2. Balsamorhiza Bolanderi Gray in Proc. Am. Acad. 7: 356. 1868; Syn. Fl. N. Am. 1?: 266. 1884; Jepson, Man. Fl. Pl. Cal 1078. 1925 Stems leafy, erect or ascending, unbranched, 1.5-3 dm. high, glandular above in young specimens, becoming glabrous in older plants; leaves mostly cauline, often bearing large scale- like bracts below, ovate, elliptical to broadly ovate-elliptical, 0.5-2.5 dm. long, 4—10 cm. broad, herbaceous in the early stages, becoming glabrous and leathery, conspicuously reticulated on both surfaces, entire, acute or obtuse-apiculate, cordate to acute at the base, young leaves glandular; heads solitary and terminal, 4—6.5 cm. broad including the rays; bracts of the in- voluere large, foliaceous, ovate to ovate-lanceolate, 2-3 em. long, herbaceous, becoming coriaceous and conspicuously re- ticulated in older plants, tomentose-ciliate in the early stages, later glabrous, acute, glandular to glabrate; rays yellow, about 3 em. long, glabrous; mature achenes quadrangular, 6—7 mm. long, glabrous. Distribution: along the mountains, Placer Co., southeast to Mariposa Co., California. CALIFORNIA: Amador Co., June 12, 1889, Greene (NY); ** Savage Hill," Amador Co., alt. 2200 ft., May, 1891, Hansen 168 (M, U CAL); sandy hillsides, Auburn, Placer Co., April, 1865, Bolander 4526 (G TYPE, NY); Bear Valley, Mariposa Co., May 2, 1892, Congdon (U CAL); Bear Valley, Mariposa Co., April 12, 1896, Cong- don (NY, U CAL); Bear Mountain, Mariposa Co., 1865, Torrey (NY); **Proner,'? Rich (NY). [Vor. 22 124 ANNALS OF THE MISSOURI BOTANICAL GARDEN This species may be readily distinguished by its conspicu- ously retieulated, coriaceous leaves, and large involueral bracts, which also become leathery and prominently reticulated. 3. Balsamorhiza Careyana Gray in an^ Am. Acad. [Pl. Fendl.] N. S. 4: 81. 1848; Syn. Fl. N. Am. 1?: 265. 1884; Howell, Fl. Northwest Am. 1: 339. 1900; Piper i in Contr. U. S. Nat. Herb. [Fl. State Wash.] 11: 580. 1906; Rydb. Fl. Rocky Mts., ed. 2, 929. 1922. Stems braeteate, one or more arising from the fibrous crowns, 2-6 dm. tall, strigose-hirsute to strigose-hispidulous and glandular above; basal leaves cordate-ovate, cordate-ob- long to hastate, 1.5-5 dm. long, 4-13 em. broad, entire, occasion- ally eoarsely dentate, acute, subcoriaceous, above scabrous, both surfaces green, strigose-hirsute to strigose-hispidulous, and densely to sparsely glandular; cauline leaves opposite, lanceolate to linear-lanceolate, becoming bracteate above; heads radiate, rarely solitary, usually 2 or more in raceme-like or close panieulate clusters, 0.8-2.5 em. broad excluding the rays; involuere campanulate, the outer series somewhat longer than the inner but rarely longer than the disk, not becoming foliaceous; rays yellow, oblong to oblong-ovate, 2-3 em. long, bi- or tri-dentate, becoming chartaceous as the head matures; mature achenes linear-oblong, 6 mm. long, ray-achenes 6- 7 mm. long, densely canescent-hirsute. Distribution: dry ground, eentral and southeastern Wash- ington, south to central and northeastern Oregon. WASHINGTON: Ellensberg, Kittitas Co., May 20, 1897, Piper (NY); Ephrata to — = Co., June, 1902, Griffiths $ Cotton 483 (NY); Sprague, Lincoln Co., alt. 2000-3000 ft., June, 1893, Sandberg $ Leiberg (F, G, M, NY, RM, U CAL); dry hillsides, Ritzville, — Co., alt. 1451 ft., June 6, 1893, Besdberg d Leiberg 166 (F, G, M, Ph, NY CAL); dry soil, Washtuena, Adams Co., May 18, 1903, Cotton 976 (G); aea bienes Walla Walla, May 23, 1903, Cotton 1052 (G, M, RM); Walla Walla region, May, 1883, Brandegee 891 (G, Ph); Prosser, Benton Co., May 27, 1903, Cotton 1105 (G, M) OREGON: ‘‘Clearwater,’’ Spalding (G TYPE, NY, Ph); Pendleton, Umatilla Co., May 15, 1905, Jones (NY); near Lexington, Morrow Co., June 5, 1894, Leiberg 28 (F, G, M, NY, RM); stony, basaltic slope, Bridge Creek, western slope of the Blue Mts., June 22, 1908, Cusick 3255 (F, G, M, NY, RM, U CAL); I near Pendle- ton, Umatilla Co., May-June, 1907, Cusick 3332 (F, G, M, NY, RM, U CAL); hill- sides of ‘‘ case River,’’ Crook Co., June 28, 1897, Cusick 1679 qe CAL); near 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 125 ““ Hoover Creek," Gilliam Co., June 1, 1894, Leiberg 142 (F, G, M, NY, RM); dry woods, Mt. Hood Forest Reserve, Bear Springs, Wasco Co., July 7, 1929, Thompson 6171 (Ph). This species may be distinguished from B. deltoidea by the smaller disk and pubescent achenes. The heads are usually in raceme-like clusters, whereas in B. deltoidea they are generally solitary. The pubescence of the achene is not always constant. In the region of Wasco and Gilliam Counties, Oregon, where the ranges of the two species overlap, the achenes are invari- ably glabrous as in B. deltoidea, but the other characters are typieal B. Careyana. 4. Balsamorhiza deltoidea Nutt. in Trans. Am. Phil. Soc. II. 7: 351. 1841; Torr. € Gray, Fl. N. Am. 2: 302. 1842; Walp. Rep. Syst. 2: 610. 1843; Gray, Syn. Fl. N. Am. 1?: 266. 1884; Coville in Contr. U. S. Nat. Herb. 4: 130. 1893; Howell, Fl. Northwest Am. 1: 339. 1900; Piper in Contr. U. S. Nat. Herb. [Fl. State Wash.] 11: 580. 1906; Piper & Beattie, Fl. North- west Coast, 376. 1915; Rydb. Fl. Rocky Mts., ed. 2, 928. 1922; Jepson, Man. Fl. Pl. Cal. 1078. 1925. Balsamorhiza glabrescens Benth. Pl. Hartw. 317. 1849. Stems bracteate, 2-8 dm. tall, erisp-pilose to crisp-hirsute, glandular above, glabrate; basal leaves numerous with petioles 1-4 em. long, the blades cordate-ovate, cordate-oblong to has- tate, 0.8-3 dm. long, 0.4-2 dm. broad, entire or occasionally coarsely dentate, becoming subeoriaceous in late stages, acute, both surfaces green, erisp-hirsute to more or less strigose-hir- sute, usually glandular, becoming seabrous above; cauline leaves reduced, becoming bracteate above, lanceolate to linear- lanceolate, 3-15 em. long, 1-3 cm. broad, entire, acute, surface as in the basal leaves, the lowermost pair usually opposite; heads solitary or sometimes two or more, 2.5-4 cm. broad ex- cluding the rays; involucre hemispherical to subhemispherical, bracts oblong-lanceolate to oblong, the outer series foliaceous, longer than the disk, often 3-6 em. long in later stages, acute or obtuse, the base of the outer bracts often densely crisp-pilose; rays yellow, oblong-lanceolate, 24.5 em. long; mature achenes about 8 mm. long, linear-oblong, glabrous. [Vor. 22 126 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: dry open ground, British Columbia south- wards along the Coast Ranges to Los Angeles Co., California. BRITISH COLUMBIA: bot er Island, May 10, 1875, Macoun 953 (G); Van- couver Island, 1858, Lyall (G); vicinity of Victoria, Vancouver Island, May 10, 1893, Macoun 475 (M, "m. Lake Osoyoos, June 3, — Macoun 76941 (F); Vancouver Island, May 5, 1908, Macoun 86299 (F); near “Lost Lake,’’ Victoria, May 23, 1916, Newcombe 62 (G); ‘‘ Thetis Lake,’’ Vancouver Island, open wooded hillsides, June 13, 1919, Carter (G). WASHINGTON: Whidbey Island, Clallam Co., Gardner (U CAL); Tacoma, Pierce Co., May 20, 1911, Zeller (M); Lake View, Pieces Co., May, 1928, Grant — Silesebetg, Kittitas Co., June, 1897, Piper (RM); Vancouver, Clarke Co., Jun 8, 1904, Piper 4945 (G); Klickitat River, near ** Mt. Paddo,’’ May 18, 1884, Buhe- dorf 364 (G); near Bingen, Klickitat Co., May 5, 1909, Suksdorf (NY); Yakima region, 1882, Brandegee (M) ; common on dry grounds, Klickitat Co., April 3, June, 1883, Suksdorf (F, Ph); slopes of ** Mt. Stuart,’’ July 24, 1893, Sandberg $ Lei- berg 5?3 (G, maA Painted Rocks, Yakima Co., May 20, 1923, Nelson 1476 (RM); Tacoma, Pierce Co., May 20, iei Brandell (M); Bettlósnake Hills, Yakima Co., May 2, 1901, etc 336 (Ph); Lake Chelan, Cascade Mts., Chelan Co., July 1-Sept. 15, 1915, Kanne 60 (M); Fan prairies near Ft. Lewis, Pierce Co., April 26, 1930, Thompson 5133 OREGON: ‘‘near the outlet of the Wahlamet,’’ May, 1835, Nuttall (G, NY, Ph TYPE); near Philmath, Benton Co., May 31, 1919, Holmes (M); Forest Grove, Washington Co., May, 1884, Drake $ Dickson (F); hillsides near Dalles, Wasco Co., April 19, 1903, Lunell 67 (G); hillsides and prairies, Hood River Co., April 26, 1924, Henderson 362 (M); hillsides, Deer Creek, near Selma, Josephine Co., April 11, 1926, Henderson 5702 (M, RM); west shore of Summer Lake, Lake Co., June 3, 1911, Eggleston 6820 (NY); high ground, Monmouth, Polk Co., May 20, 1893, Page 216 (F); hillsides, init cl s Village, Pone Co. | May 20, 1896, Lloyd (NY); dry ridge, 4 mi. west of Table Rock, Clac M Ob, May 30, 1924, Peck (WU); dry slope, Ashland, Jackson Co., July 14, Tt Peck (WU); dry hillsides, the Dalles, Wasco Co., April 7, 1914, Peck 5040 (WU); Wimer, Jackson Co., June 29, 1892, Hammond 209 (M, NY, Ph, RM); Salem, Marion Co., 1871, Hall 276 (F, G, M); dry ground near Hood River, Hood River Co., April 3, 1926, Thompson 661 (M); ‘‘Seraggins’’ Valley, Washington Co., May 21, 1927, Thompson 2495 (Ph); open pastures near Estacada, Clackamas Co., : une 11, 1927, Thompson 2593 (Ph); hills near Brownsboro, Jackson Co., April 10, 1927, Thompson 2163 (Ph); open sandy — Hood River, Hood River Co. er 10, 1920, Gorman (Ph); hill, one mile east of Mosier, Wasco Co., May , 1928, Gale 111 (M, Ph); open ground, Salem, + hee Co., May 2, 1917, N x 1091 (G); border of field, 3 miles south of Salem, Marion Co., April 23, 1921, Nelson 3506 (Ph); dry open woods, Grants Pass, Josephine Co. , April 20, 1912, Prescott 11 (G); Dalles, Wasco Co., April, 1882, Howell (NY); June, 1881, Howell ( Ye CALIFORNIA: Kernville, Kern Co. , May 14, 1892, Brandegee (U ERA between Auburn and Coal, Eldorado Co., April 28, 1909, pigia (U CAL); Water ie Tehachapi Mts., Kern Co., alt. 6000 ft., June 26, 1908, A din $ McGregor 3 (NY); Liebre Mts., Los Aignis es Co., alt. 4500 ft;, jas 20-23, 1908, Abrams $ pad 356 (G, NY); Cantara, Siskiyou Co., May 23, 1923, Eastwood 11927 (M); Tehachapi, Kern Co., May 13, 1913, Eastwood 3237 (G, NY); near Inde- 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 127 dones Inyo Co., alt. 5500 ft., Hall $ Chandler 7304 (M, U CAL); south slope ar Kernville En Co., alt. 5000 ft., June 1904, Hall $ Babcock 5100 (U CAL); Tejon E Lós E Co., June, 1905, Hall 6263 (U CAL); in grassy spots ong pines, Sierra Nevada foothills, Eldorado Co., April 13, 1921, Hall 11246 (U CAL); hillsides beneath pines, Yosemite Nat. Per Mariposa Co., June 10, 1911, Hall 8907 (U CAL); on Moro Creek, San Rafael Mts., Santa Barbers Co., May 18, 1907, Hall 7797 (Ph, RM, U CAL); Blue Cañon, Placer Co., alt. 4701 ft., June 24, 1908, Walker 1285 (U s Mendocino Co., 1866, Bolander 6187 (F,G, =, NY dry oon stony slopes, Back Mt., Humboldt Co., Tracy 4175 (U CAL); Sunt Ranch, Trinity Co., alt. 1500 ft., April 27, 1924, ER 6662 (U CAL); open woods, Trinity mid Valley, Humboldt Co., alt. 600 ft., May 17, 1925, Tracy 6984 (U CAL); Tehaehapi Valley, Kern Co., May 11, 1896, Davy 2185 (U CAL); open ground, * Rawhide Hill," Tuolumne Co., May 30, 1919, Williamson 185 (NY); Keene Station, Kern Co., May 1, 1905, Heller 7802 (G, M, NY, Ph); on the New- ville-Cuelo Road, Glenn Co., alt. 3000 ft., June 16, 1915, Heller 1197? (M) ; dry soil, Siskiyou Co., April 14, 1909, Butler 595 (U CAL); Klamath Hills, Siskiyou Co., April 16, 1910, Butler 1180 (M, RM, U CAL); near Yreka, Siskiyou Co., April 25, 1876, Greene 718 (F, G, M, NY); region of Tehachapi Peak, Kern Co., June 25, 1895, Dudley 363 (F, NY, U CAL); Sonoma Co., coll. of 1853-54, Bigelow (G, NY); open hills near **Freneh Flat,’’ Tuolumne Co., alt. 1450 ft., April 11-16, 1919, Ferris 1580 (NY); ** Mt. Sanhedrin,’’ Mendocino Co., June 7, 1893, Blankin- ship (G). The leaves of B. deltoidea are of relatively thin texture in the early stages, but after the flowering season they become more or less coriaceous. This occurrence is more common in the southern part of its range. 5. Balsamorhiza sagittata (Pursh) Nutt. in Trans. Am. Phil. Soc. II. 7: 350. 1841; Torr. € Gray, Fl. N. Am. 2: 301. 1842; Walp. Rep. Bot. Syst. 2: 610. 1843; Gray in Mem. Am. Acad. [Pl. Fendl.] N. S. 4: 82. 1848; Syn. Fl. N. Am. 1?: 265. 1884; Howell, Fl. Northwest Am. 1: 339. 1900; Piper in Contr. U. $. Nat. Herb. [Fl. State Wash.] 11: 580. 1906; Coult. & Nels. Bot. Roeky Mts. 545. 1909; Rydb. Fl. Rocky Mts., ed. 2, 928. 1922; Blake in Contr. U. S. Nat. Herb. [Tidestrom's Fl. Utah & Nevada] 25: 582. 1925; Jepson, Man. Fl. Pl. Cal. 1078. 1925; Rydb. Fl. Pr. & Pl. Cent. N. Am. 839. 1932 Bodl Animum sagittatum Pursh, Fl. Am. Sept. 2: 564. 1814. Balsamorhiza helianthoides Nutt. in Trans. Am. Phil. Soc. II. 7: 351. 1841; Torr. & Gray, Fl. N. Am. 2: 302. 1842; Walp. Rep. Bot. Syst. 2: 610. 1843. Espeletia helianthoides Nutt. in Jour. Acad. Nat. Sci. Phila. 7: 39. 1834. [VoL. 22 128 ANNALS OF THE MISSOURI BOTANICAL GARDEN Espeletia sagittata Nutt. Ibid. 38. pl. 4. 1834. Stems leafless or bracteate, one to several from a single erown, 2-6.5 dm. tall, tomentose below, densely tomentose above; leaves basal, cordate-ovate, cordate-oblong to hastate, 2-4 dm. long, 0.5-1.5 dm. broad, entire, acute, above cinereous- strigulose, beneath silvery tomentulose to velutinous; heads usually solitary, sometimes 2-3, 6-11 em. broad including the rays; bracts of the involucre ovate-lanceolate to lanceolate, the outer series longer than the inner, acuminate to attenuate, densely white-tomentose; pales long-attenuate, terminating in an awl-like apex, pilose-ciliate; rays yellow, oblong-lanceolate, 2-4 cm. long, 0.8-1.5 cm. broad, tube erisp-pilose; mature achenes linear-oblong, 7-8 mm. long, glabrous. Distribution: dry ground, South Dakota and Colorado, west to British Columbia and California. SouTH DAKOTA: limestone, Black Hills Nat. Forest, June 15, 1910, Murdock 4096 (F,G); EN west of Custer, Custer Co., alt. 6000-6500 ft., July 29, 1892, — 808 (G, NY). : hills near Midvale, Teton Co., June 16, 1903, Umbach 60 (F); near Great Fall, anda Co., May 2, — Anderson (U CAL, M); near Butte, Silver- bow Co., 1893, Moore (M); Bridger Mts., Gallatin Co., June 11, 1897, Rydberg 4 Bessey 5174 (F, G, NY, RM); ir Yellowstone Co., May 30, €: — 52 (G, M); near Missoula, Missoula Co., May 19, 1994, Kirkwoo U CAL); ; Helena, Lewis and Clark Co., Jue 24, 1888, Kelsey (U hs her region, Flathead Co., 1883, Ayres 63 (Ph); Flathead River, 1834, Nuttall (Ph TYPE). Wyoming: Laramie Hills, Albany Co., June, 1908, Nelson 9140 (RM); Evans- ton, Uinta Co., June 4, 1898, Nelson 4518 (RM); hills, Bush Ranch, Sweetwater Co., June 10, 1900, Nelson 7084 (RM); dry soil, Leucite Hills, Sweetwater Co., June 18, 1901, Merrill $ Wilcox 686 (G, RM); Glen Creek, Yellowstone National Park, July 1, 1899, Nelson $ Nelson 5613 (G, M, NY, RM); Little Goose Canyon, Sheri- dan Co., June 22, 1913, Sharp 356 (RM); Fremont Lake, Fremont Co., June 29, 1904, Warren (RM); near Rawlins, Carbon Co., June 30, 1899, Pammel 75 (M); dry rocky flats, Grand Teton National Park, June 17, 1933, Williams 1125 (M); dry open ground, near the ‘‘ Devil's Tower,’’ Crook Co., June 27, 1929, Palmer (G). COLORADO: Cimarron, Montrose Co., alt. 7000 ft., May 19, 1898, Crandall 2723 (F, G, NY, Ph, RM); Rio La Plata, La Plata Co., June, 1875, Brandegee 1077 (RM); Cerro Summit, Montrose Co., alt. 8000 ft., June 7, 1901, Baker 51 (G, M, NY, RM, U CAL); Norwood Hill, San Miguel Co., Aug. 16, 1912, Walker 466 (G, M, RM); open pde near Naturita, Montrose Co., alt. 7000 ft., May 25, 1914, Payson 343 (F, G, M, RM). UTAH: Ogden, i Co., July 1, 1880, Engelmann (M); hillsides, Wheelon- 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 129 Collinston, Boxelder Co., May 1, 1911, Smith 2348 (NY); Fort Douglas, Salt Lake Co., May 5, 1908, Clemens (F, G, Ph); dry hillsides, Daggett Co., May 28, 1932, Williams 415 (NY, R IDAHO: Kootenai Co. als. 1890, Leiberg E NY); Boise, Ada Co., alt. 2880 ft., May 27, 1911, Clark 28 (F, G, M, RM, U CAL); Montpelier, Bear Kaka Co., May 20, 1910, Macbride 202 (G, M, RM); near Lewiston, Nez Perees Co., April 27, 1896, Heller Ad Heller 2958 (F, M, NY, Ph); New Plymouth, Canyon Co., alt. 2200 ft., May 21, 1910, Macbride 79 (G, M, NY, RM). NEVADA: Pyramid Lake, Washoe Co., June 1, 1913, Kennedy 1985 (M, Ph, U CAL); King's Cañon, Ormsby Co., June 1, 1902, Baker 926 (G, M, NY, RM, U CAL) ; Carson E 1863, pues 173 (G BRITISH COLUMBIA: between Midway and Is Osoyoos, May 5, 1905, Macoun 76942 (NY); Acier 's Bridge, May 26, 1889, Macoun (G). WASHINGTON: Pullman, Whitman Co., rn 1, 1893, Piper 1594 (F, G, NY); Hangman Creek, Spokane ER alt. 1510 ft., May 17, 1893, Sandberg $ Leiberg 23 (F, G, M, NY, Ph, RM, U CAL); roeky ope north of Entiat, em Co., April 18, 1931, Thompson 5985 (G, NY); open slopes, Beverly Creek, Kittitas Co. , May 16, 1931, Thompson 6386 (G); hillsides near Waitsburg, Walla waa Co., April 21, 1897, Horner B277 (G); among sage-brush near Coulee City, Grant Co., May 2, o ea 6126 (G). near North Powder, Union Co., May 10, 1928, Gale 255 a. Wallowa Co. zn 30, 1927, giis ie ii near Lone Rock, Lillian Co., alt. 890 m., May 25, 1894, Leiberg 114 (F, NY, U CAL); 15 mi. n. e. of Penco, Umatilla Co., May 17, 1923, oa 372 (F, WU); dry ground, Harney Co., June 28, 1912, Peck 3071 (WU); sagebrush, Brogan, Malheur Co., 1910, tolle 1182 (WU); D ) CALIFORNIA: Alpine Co., June, 1892, Hansen 420 (M) ; American Valley, Plumas Bet J 2 1875, Austin 94 (U OAL); ; on Sierra T above Jonesville, Butte Co., 000 ft., June 16, 1915, Heller 12026 (F, G, M); open plaees in the yellow fus a E Mineral, Tehama Co., May 30, 1998, Heller 14594 (M, NY); Goose Lake Valley, Modoe Co., May, 1894, Austin 47 (U CAL); Deer Park, Placer Co. alt. 6500 ft., Walker 2016 (M, RM). 6. Balsamorhiza Hookeri Nutt. in Trans. Am. Phil. Soc. II, 7: 349. 1841; Torr. € Gray, Fl N. Am. 2: 301. 1842; Walp. Rep. Bot. Syst. 2: 610. 1843; Gray, Syn. Fl. N. Am. 12: 266. 1884; Howell, Fl. Northwest Am. 1: 340. 1900; Coult. & Nels. Bot. Rocky Mts. 546. 1909; Blake in Contr. U. S. Nat. Herb. [Tidestrom’s Fl. Uiii Nerada 25: 583. 1925, in part; Jepson, Man. Fl. Pl. Cal. 1078. 1925, in part. Heliopsis (?) Balsamorrhiza Hook. Fl. Bor.-Am. 1: 310. [1834] 1840; DC. Prodr. 5: 551. 1836. A A Balsamorrhiza (Hook.) Heller in Cat. N. Am. Pl. 7. 1898; Piper in Contr. U. S. Nat. Herb. [Fl. State Wash.] 11: 581. 1906; Piper & Beattie, Fl. Northwest Coast, 376. 1915. [Vor. 22 130 ANNALS OF THE MISSOURI BOTANICAL GARDEN Stems scapose, 1-3 dm. tall, pilose near the heads, otherwise subsericeous; leaves ovate-lanceolate to lanceolate, pinnately incised or divided, 1-3 dm. long, the divisions cleft into several spreading or ascending, acute or entire lobes, 1—4 cm. long, 0.5-1 em. broad, subvelutinous-eanescent on both surfaces, often densely so on the younger leaves, of medium texture; heads solitary, 4—7 em. broad including the rays; braets of the involuere linear-lanceolate to linear-oblong, about as long as or shorter than the disk, acute, pilose; rays yellow, oblong to ovate-oblong, 1.5-2.5 em. long, bifid or entire, tube and lamina pubescent; achenes 4-5 mm. long, glabrous. Distribution: moist grounds, Yakima and Klickitat Coun- ties, west to Clark County, Washington. WASHINGTON: near the mouth of the Columbia River, ‘‘1834-35,’’ Nuttall (G, Ph, NY); Tampico, Yakima Co., April 6, 1924, Nelson 1595 (RM); — Mts., Yakima Co., June, 1881, Howell (NY); p rairies, eae Mts., Yakim „ Howell (G); low gronni Falcon Valley, Klickitat Co. y 20, 1881, ^an p^ Ph); low grounds in open woods, Falcon Valley, ri iu Qo. May 3, 1884, Suksdorf 357 (G); Klickitat River, Klickitat Co., May 18, 1884, Suksdorf 359 (G); Falcon Valley, Kliekitat Co., May 3, 1884, Sukedorf 363 (G); Mill Plain, Clark Co., June 4, 1880, Howell (F). 6a. var. rosea (Nels. € Macbr.) W. M. Sharp, comb. nov. Balsamorhiza rosea Nels. € Macbr. in Bot. Gaz. 56: 478. Stems about 1 dm. tall, purplish; leaves .8-1.2 dm. long, stri- gose-hirsutulous; rays yellow or purplish; achenes 5-6 mm. long, pubescent; in all other characters similar to the species. Distribution: thin rocky ridges of Rattlesnake Mts., Yakima Co., Washington. WASHINGTON: ridges of Rattlesnake Mts., Yakima Co., May 8, 1902, Cotton 568 (G, M, Ph, RM TYPE) 6b. var. lanata W. M. Sharp, var. nov.!? Leaves 1-2.5 dm. long, bipinnately divided, the segments narrow, 1.5-5 em. long, 0.5-1 em. broad, lanceolate, acute, lanate " B. Hookeri var. lanata W. M. Sharp, var. nov., foliis 1-2.5 dm. longis, pin- natisectis, segmentis angustis, 1.5-5 em. longis, 0.5— . latis, lanceolatis, acutis, utrinque lanatis; petiolis dense lanatis; Pda didi lanceolatis, acuminatis, lanatis. Cetera speciei similis.—Collected near Yreka, Siskiyou Co., California, May 2, 1876, Greene 745 (F, G, M TYPE). 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 131 on both surfaces; petioles densely lanate; bracts of the in- voluere lanceolate, acuminate, lanate; other characters similar to the species. Distribution: dry hills near Yreka, Siskiyou Co., California. ae near Yreka, Siskiyou Co., May 2, 1876, Greene 745 (F, G, M TYPE); dry ear Yreka, Siskiyou Co., April 9, 1910, Butler 1157 (M, RM, U CAL); near Yreka, Siskiyou Co., April, 1903, Mooney (U CAL). 7. Balsamorhiza platylepis W. M. Sharp, sp. nov.!! Stems scapose, 1-3.5 dm. high, with cinereous-appressed pu- bescence; leaves pinnately divided, 1-3 dm. long, the divisions 2-5 em. long, 0.5-1.5 em. broad, entire, coarsely serrate to in- cised, acute, above light green, velutinous to subsericeous, be- neath silvery sericeous to velutinous; heads solitary, 4—6 cm. broad including the rays; braets of the involucre equalling or shorter than the disk, triangular to ovate, acute, 1-2 em. long, 5-8 mm. broad, outer series brownish or hoary at the apex, greenish below, sericeous to velutinous; rays yellow, broadly oblong, 2-3.5 cm. long, .5-1 em. broad, bifid or entire, tube and lamina pubescent; achenes 6-8 mm. long, glabrous, epappose. Distribution: dry gravelly or rocky ground, southern Oregon, southward to west-central Nevada and northern California. NEVADA: Marmol Station, Washoe Co., alt. 5000 ft., May 30, 1912, Kennedy 1859 (F TYPE, G, M, NY); near Verdi, Washes Co., May 5, 1895, Sonne (NY). OREGON: serpentine ridge northwest of Selma, Josephine Co., April 13, 1924, Thompson 2272 (WU); hillsides near Waldo, Josephine Co., April, 1887, Howell (M, NY); serpentine hills along Deer Creek, north of Selma, Josephine Co., March 29, 1926, Henderson 5703 (RM); Quel Valley,’’ 1893, Austin (U CAL). CALIFORNIA: Modoc Co. Meu 1898, Austin $ Bruce 2179 (NY, U CAL); Goose Lake Valley, Modoc Co., Ma 2 pum Austin (U CAL, Ph); lava beds near Egg Lake, Modoe Co., June 8, 189 aker $ Nutting (RM, U CAL); Goose Valley, Shasta Co. Se 1912, aoe 896 (G, NY); Plumas Co., 1879, Austin "B. platylepis W. M. Sharp, sp. nov., caulibus scaposis, 1-3.5 dm. altis, adpresso-cinereo-pubescentibus; foliis Xiibatisoctis, 1-3 dm. longis, segmentis 2-5 em. longis, 0.5-1.5 em. latis, integris, grosse serratis vel incisis, acutis, supra pallide Kei ciem plus minusve velutinis vel subsericeis, subtus argenteo-sericeis vel utinis; braeteis involueri disco aequalibus vel brevioribus, triangulatis vel ovatis, eco 1-2 em. longis, 5-8 mm. latis, exterioribus ad apicem fulvis aut albidis, subtus viridibus, sericeis vel velutinis.—Collected at Marmol Station, Washoe Co., Nevada, alt. 5000 ft., Kennedy 1859 (F TYPE, G, M, NY). [Vor. 22 132 ANNALS OF THE MISSOURI BOTANICAL GARDEN (G); near Truckee, Nevada Co., June 7, 1917, Hall 10327 (U CAL); Dog Valley, Sierra Co., May, 1892, Sonne 368 (M); Hobart Mills, Nevada Co., May-June, 1925, rew (RM). 8. Balsamorhiza macrolepis W. M. Sharp, sp. nov.!? Stems usually scapose, 2-6 dm. high, below appressed-pilose, above appressed or spreading pubescent; leaves pinnately di- vided to bipinnatifid, 2.5-5 dm. long, the divisions lanceolate in general outline, 3-10 cm. long, 0.5-2 em. broad, entire, incisely toothed or again pinnatifid, acute, above green, subvelutinous, beneath light green to canescent, densely subsericeous to velu- tinous, glandular; heads solitary, radiate, 4-8 em. broad includ- ing the rays; bracts of the involucre 3-seriate, the outer series large, twice as long as the disk, oblong to oblong-lanceolate, 2— 3.5 cm. long, 0.5-1.3 cm. broad, obtuse or abruptly acuminate, often dentate or incised towards the apex, minutely velutinous; rays yellow, oblong-elliptical, 2.5-3 cm. long, 1-1.5 cm. broad, tube and lamina puberulent to glabrous; achenes linear-oblong, 5-7 mm. long, quadrangular, glabrous Distribution: north-central California, Butte Co., southeast to Mariposa Co. southwest to Sonoma and Santa Clara Counties. CALIFORNIA: Clear Creek, Butte Co., May 1-15, 1897, Brown 208 (F, M, NY TYPE, Ph, RM); Brush Creek, Butte Co., 1908, Konger 122843 (U CAL); fields, Butte Co., May 1898, Bruce 2469 (N Y); sandy hillsides near Lincoln, Placer April 15, 1917, Ramaley 11109 (U CAL); Amador, Amador Co., alt. 1000 ft., May, 1895, Hansen 1069 (M) ; Sonoma, Sonoma Co., 1853-54, Bigelow (G, NY); Cedar Mt., Alameda Co., May, 1903, Elmer 4374 (M, NY, U CAL); Oakland Hills, near Oakland, Alameda Co., June 6, 1868, Kellogg $ Harford 430 (NY, G) ; San Martin, Santa Clara Co., "e 30, 1901, von 914 (U CAL); Bear Mt., Mariposa Co., 1865, Torr y (M); ; ““Coulterville Road,’’ Yosemite Nat. Park, Mariposa Co., alt. 4500 ft., June 7, 1911, Hall nio (U CAL); ‘Kirby Ranch,’’ Mariposa Co., May, 1886, Congdon (U CAL). “ B. macrolepis W. M. Sharp, sp. nov., caulibus 2-6 dm. altis, infra adpresse pilosis, supra adpresse vel laxe O foliis en vel bipinnatifidis, 2.5-5 dm. longis, segmentis laneeolatis, 3-10 em. longis, 0.5-2 em. latis, tin incisis aut pinnatifidis, acutis, supra viridibus et subvelutinis subtus pallide viri ibus vel canescentibus et plus minusve dense subsericeis vel velutinis, glandulosis; braeteis involueri 3-seriatis, exterioribus magnis, i: duplo longioribus, oblongis vel oblongo-laneeolatis, 2-3.5 em. longis 0.5-1.3 em. latis, obtusis aut acuminatis, aepe ad apice m. dentatis vel incisis, minute velutinis.—Collected at Clear Creek, Butte Co., California, May 1-15, 1897, Brown 208 (F, M, NY TYPE, Ph, RM). 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 183 This species may be readily distinguished from other species of Balsamorhiza by its large puberulous, oblong bracts 2-3 em. long, large leaves 2.5—5 dm. long, and its soft subvelutinous pu- bescenee. This is the only pinnate-leaved species of this genus oceurring in California south of Butte County. 9. Balsamorhiza incana Nutt. in Trans. Am. Phil. Soc. II. 7: 350. 1841; Torr. € Gray, Fl. N. Am. 2: 301. 1842; Walp. Rep. Bot. Syst. 2: 610. 1843; Howell, Fl. Northwest Am. 1: 340. 1900; Coult. € Nels. Bot. Rocky Mts. 546. 1909; Rydb. Fl. Rocky Mts., ed. 2, 929. 1922; Blake in Contr. U. S. Nat. Herb. [Tidestrom's Fl. Utah & Nevada] 25: 582. 1925. Balsamorrhiza Hookeri var. incana (Nutt.) Gray, Syn. Fl. N. Am. 1?: 266. 1884. Balsamorrhiza floccosa Rydb. in Bull. Torr. Bot. Club 27: 629. 1900; and Fl. Rocky Mts., ed. 2, 929. 1922. Stems subscapose, 1.5-4 dm. tall, white-tomentose to lanate- tomentose ; leaves lanceolate to oblong-lanceolate, pinnately di- vided, 1-4 dm. long, the divisions lanceolate, oblong-lanceolate to ovate-lanceolate, 1.5-6 em. long, 0.5-2.5 em. broad, entire, coarsely serrate or serrate-incised, acute, above light green, beneath cinereous, both surfaces tomentose to floccose-tomen- tose, petioles tomentose to lanate-tomentose; heads solitary, 7-11 cm. broad including the rays; bracts of the involucre 3- seriate, ovate-attenuate to lance-attenuate, about as long as the disk, densely white-tomentose to floccose-tomentose; pales at- tenuate-subulate, pilose on the upper half; rays yellow, ob- long, 34.5 em. long, 0.5-1 em. broad, bifid or entire, tube and base of lamina pilose-tomentose; disk-flowers 9-10 mm. long, glabrous; achenes linear-oblong, 5-6 mm. long, glabrous. Distribution: southwestern Montana and western Wyoming to Washington and Oregon. MONTANA: Spanish Basin, Gallatin Co., alt. 6000 ft., July 1, 1897, Rydberg $ Bessey 5175 (F, G, NY, RM); ‘‘Camp Creek,’’ June 12, 1905, Jones (U CAL). WYOMING: Rocky Mountains, 1834, Nuttall (Ph TYPE); Three Forks of the Mis- souri and Yellowstone Rivers, June 30, 1860, Hayden (M); dry soil, ** Big Sandy River near Leckie,’’ June 22, 1901, Merrill $ Wilcox 743 (G, RM, NY); eastern slope of the Big Horn Mts., headwaters of Clear Creek, Johnson Co., alt. 7000— 9000 ft, July 20-Aug. 15, 1900, Tweedy 3150 (RM, NY); Rattlesnake Mts., [Vor. 22 134 ANNALS OF THE MISSOURI BOTANICAL GARDEN Natrona Co., July 29, 1898, Nelson 4993 (M, RM, NY); near Big Horn, Sheridan Co., alt. 5000 ft., July 2, 1897, Pammel $ Stanton 322 (M); Little Goose Cr., Sheridan Co., July 16, 1896, Nelson 2342 (RM, M); ID of the Tongue River, Big Hor n Mts., Sheridan Co., July, 1898, Tweedy 11 (NY); Wind River Range, June, 1873, Parry 165 (M, Ph, NY); Little Bald Sa Big Horn Mts., Big- horn Co., alt. 9500 ft., July 13, 1900, Jack (G); Wind River Valley, May 18, 1860, Hayden (M); Big Horn Mts., Sheridan Co., alt. 7500 ft., July, 1899, Tweedy 2115 (NY). IpaHo: about Lake Waha, Nez Perces Co., alt. 3500-4000 ft., June 22, 1896, Heller $ Heller 3298 (M, Ph, NY, U CAL); Clearwater, Spalding (G, Ph); Craig Mts. at Lake Waha, Nez Perces Co., May 24, 1892, Sandberg, MacDougal $ Heller 248 in part (F, G, NY). WASHINGTON: open slopes near Lindel, Asotin Co., May 20, 1928, St. John $ Brown 3900 (Ph); Blue Mts., Columbia Co., June 11, 1887, Horner (G). OREGON: Cabbage Hill east of Pendleton, TETS Co., May 10, 1928, Gale 178 (M); i open ground near Pendleton, Umatilla Co., June 24, 1927, Sehrisien (Ph, WU); 3 miles south of Elgin, Union Co., ub 22, 1923, Bhorweod 323 (WU, F); ‘‘Camas prairie,’’ July, 1902, Griffiths $ Hunter 77 (NY). 9a. var. tomentosa (Rydb.) W. M. Sharp, comb. nov. Balsamorrhiza tomentosa Rydb. in Tem Torr. Bot. Club. 27: 628. 1900; Fl. Rocky Mts., ed. 2, 928. 192 Leaves oblong«ovaté to deca e 2.5—4 dm. long, 3- 8 em. broad, subentire, dentate-lacerate to incised, acute, above light green, tomentose to lanate-tomentose, beneath whitish, densely tomentose; bracts of the involucre linear-lanceolate, abruptly attenuate, densely tomentose; otherwise similar to the species. Distribution: western Montana, Wyoming, and Idaho. MONTANA: hills, src Gulch,’’ July 12, 1883, — 189 (Ph). WYOMING: eastern slopes e Big Horn Mts., headwaters of Clear Creek, Johnson Co., alt. 7000—9000 ^s July 20-Aug. 15, fon 3151 (NY, RM); head- waters of the Tongue River, Big Horn Mts., July, 1898, Tweedy 10 (NY TYPE); Big Horn Mts., Sheridan Co., alt. 7500 ft., July, 1899, Tweedy 214 (NY). IDAHO: hillsides, Craig Mt. at Lake Waha, Nez Perces Co., May 24, 1892, Sand- berg, MacDougal $ Heller 248 (M). The subentire to incised leaf-blades, the tomentose-pubescent leaves, and the linear-lanceolate, strongly attenuated and acute braets are the main distinguishing characters of this variety. . Balsamorhiza macrophylla Nutt. in Trans. Am. Phil. Soc. 10 II, 7: 350. 1841; Torr. € Gray, Fl. N. Am. 2: 301. 1842; Walp. Rep. Bot. Syst. 2: 610. 1843; Gray, Syn. Fl. N. Am. 1?: 266. 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 135 1884; Rydb. Fl. Rocky Mts., ed. 2, 929. 1922; Blake in Contr. U. S. Nat. Herb. [Tidestrom's Fl. Utah & N evada] 25: 583. 1925. Balsamorhiza macrophylla var. terebinthacea Nels. in Coult. & Nels. Bot. Rocky Mts., 546. 1909, not B. terebinthacea (Hook.) Nutt. Stems subscapose, 3-10 dm. high, crisp-pilose and glandular to glabrate; leaves pinnate or pinnately cleft, rarely subentire, 3-6 dm. long, 6-16 cm. broad, hirsute-ciliate on the margins, above green and sparsely crisp-hirsute to glabrous, beneath green, glandular, and crisp-hirsute on the veins, segments 5-11 em. long, entire or sparsely toothed; petioles glandular, gla- brous to sparingly pubescent ; heads solitary, including the rays 8-12 em. broad; bracts of the involucre lanceolate, oblong- lanceolate, 2.5-5 em. long, acute to acuminate, densely pilose to hirsute-pilose-ciliate, dorsal surface sparsely pilose-hirsute; rays broadly oblong-elliptical, 3.5-5.5 em. long, 1.5-2.2 cm. broad, bifid or entire, tube and outer surface of lamina puberu- lent or glabrous; achenes linear-oblong, quadrangular, with a small rib between each of the larger angles, 8-10 mm. long, glabrous. Distribution: dry open slopes and sage-brush associations, western Wyoming and eastern Idaho, south to northern Utah. NY); Beaver Creek Canyon, Fremont Co., June 27, 1895, Shear 3037 ( NY); hills, Beaver Creek Canyon, Fremont Co., June 27, Shear 3008 (NY); dry open slopes, Caribou Mts., Bonneville Co., July 17, 1923, Payson $ Armstrong 3507 (G, M, RM, Ph); Oxford, Oneida Co., May 9, 1885, Leonard (G). Wyoming: Union Pass, Wind River Mts., Fremont Co., Aug. 10, 1894, Nelson 4015 (RM); sagebrush slopes, 20 mi, east of Big Piney, Sublette Co., July 9, 1922, Payson $ Payson 2605 (G, M, Ph, RM, U CAL, NY); headwaters of the Hoback River, Lincoln Co., alt. 8000 ft., July 18, 1933, Williams 1317 (M); Gros Ventre Mts., Teton Co., alt. 8500 ft., Aug. 29-Sept. 6, 1900, Curtis (NY). UTAH: Parley's Cañon near Salt Lake City, Salt Lake Co., alt. 8000 ft., July 1, 1898, Jones 6407 (M); Parley's Canyon near Salt Lake City, June 23, 1922, Oster- [Vor. 22 136 ANNALS OF THE MISSOURI BOTANICAL GARDEN Club River, May 8, 1911, Smith 2366 (NY); Midway, Wasatch Co., June 15, 1933, Eastwood 4 Howell 507 (M). 10a. var. idahoensis W. M. Sharp, var. nov." Leaf-segments 3-6 em. long, 1-2.5 em. broad, eoarsely ser- rate-dentate or incised, acute, uniformly erisp-pilose to pilose- tomentose on both surfaces; petioles pilose-tomentose; bracts of the involucre usually shorter than the disk, lanceolate, 1- 2 em. long, pilose to pilose-tomentose; in all other characters similar to the species. Distribution: known only from Boise Co., western Idaho. IpAHO: rocky ground, Squaw Creek, Boise Co., alt. 3500 ft., May 8, 1911, Mac- bride 820 (F, G, M TYPE, NY, RM, U CAL). The variety idahoensis may be distinguished from the spe- cies Balsamorhiza macrophylla by the smaller and shallowly incised leaf-segments, the uniformly pilose leaf-surface, the pilose-tomentose petioles, and the short involueral braets. 11. Balsamorhiza serrata Nels. € Macbr. in Bot. Gaz. 56: 479. 1913. Stems scapose, unbranched, 1-3.5 dm. high, erisp-pilose, with hirsute-hispidulous hairs intermixed ; leaves chiefly basal, lan- ceolate to ovate-lanceolate, 0.6-3 dm. long, 2-10 cm. broad, dentate- to serrate-subuliferous, aeute, strigose to hispidulous on both surfaces, interspersed with minute stiff hairs, conspicu- ously retieulate-veined, eoriaceous; heads solitary, 9-8 em. broad including the rays; bracts of the involucre linear-lance- olate, about as long as the disk, subuliferous, tomentose to erisp-pilose, herbaceous; rays yellow, oblong, 2.5-3.5 em. long, 6-8 mm. broad, bidentate, tube and outer surface of lamina hirsutulous to glabrate; achenes linear-oblong, 6-7 mm. long, glabrous, epappose. Distribution: dry rocky ground, Columbia Co., Washing- ton, to Morrow and Wasco Counties, south to Harney and Lake Counties, Oregon. " B. macrophylla var. idahoensis W. M. Sharp, var. nov., segmentis folii 3-6 cm. longis, 1-2.5 em. latis, serrato-dentatis vel incisis, acutis, utrinque piloso-erispis vel piloso-tomentosis; petiolis piloso-tomentosis ; bracteis involueri disco brevioribus, lanceolatis, 1-2 em. longis, pilosis vel piloso-tomentosis. Cetera speciei similis.—Col- lected on rocky ground, Squaw Creek, Boise Co., Idaho, alt. 3500 ft., May 8, 1911, Macbride 820 (F, G, M TYPE, NY, RM, U CAL). 1935] SHARP— CERTAIN EPAPPOSE GENERA OF COMPOSITAE 137 WASHINGTON: Blue Mts., Columbia Co., July 9, 1913, Darlington 54 (G, RM). OREGON: Union Co., 1887, Cusick 595, 1031 (G) ; upper Tygh Valley, Wasco Co., April wi 1933, Bellinger (WU); dry ground near Shaniko, Waseo Co., May 5, 1928, Gale 100 (M); T stony xe 10 mi. north of Wapinitis, Waseo Co., May 29, 1933, Mes 17393 (WU); rocky hills near Rock Creek, Morrow Co., alt. 1090 m., May 19, 1894, Leiberg 83 (8, M, NY, RM TYPE); near Rhea Creek, vicinity of Lex- ington, Morrow Co., alt. 605 m., May 12, 1894, Leiberg 58 (F, G, M, NY, RM); John Day Valley, May 16, 1885, Howell (F, G, NY, Ph); dry roeky mountain top near Fossel, Wheeler Co., June 3, 1925, Henderson 5248 (G); mesa, between Warner Valley P. O. and Big Lake, Lake Co., alt. 1600 m., June 16, 1911, Eggleston 6964 (NY); dry sterile soil, Lakeview, Like Co., June 27, kir Peck 15264a (M, WU); a flat on Stein’s Mts., Harney Co., June 30, 1898, Cusick 1984 (F, G, M, CAL); Silver Creek ER aa 6 mi. west of Bo Harney Co., June 22, 1925, x 13900 (WU); near Harper Ranch, alt. 1000 m., May 22, 1896, Leiberg 2106 (G, NY, U CAL). 12. Balsamorhiza hispidula W. M. Sharp, sp. nov.!* Stems 0.6-3 dm. high, pilose-hirsute, glandular; leaves 1- 4 dm. long, pinnately divided, the segments unequally pin- nately cleft or parted, 2-4.5 em. long, acute, green on both sur- faces, above sparsely hispid, hispid-pilose to hispidulous, glan- dular, beneath globular-resinous and usually hispid, hispidu- lous or hispid-pilose on the veins; petioles pilose-hirsute to pilose-hispid, glandular; heads vadiale. solitary, 4.5-6 em. long including the rays; braets of the involucre linear to linear- lanceolate, about as long as the disk, the outer series as long as or longer than the inner series, more or less pilose-hirsute- ciliate, acute to acuminate, the outer surface interspersed by numerous glands; rays yellow, bidentate, 2.5-3 cm. long, tube and outer surface of lamina hirtellous; achenes 6-7 mm. long, glabrous, epappose. Distribution: dry grounds, southwestern Wyoming, Idaho and northern Utah, west to Nevada. Wyoming: South Butte, **thirteen mile,’’ July 17, 1897, Nelson 3552 (RM). Ipamo: vicinity of Pocatello, Bannock Co., 1921, Soth 7 (NY); hills near Oxford, * B. hispidula W. M. Sharp, sp. nov., caulibus 0.6-3 dm. altis, piloso-hirsutis et glandulosis; foliis 1-4 dm. "Per planatisestis: segmentis inaequaliter pinnati- fidis vel pinnatipartitis, 2-4.5 em. longis, acutis, utrinque viridibus, supra sparse hispido-pilosis vel hispidulis et giandulésis, subtus globuloso-resinosis et plerumque hispidis, in nerviis hispidulis vel hispido- pd petiolis piloso-hirsutis vel piloso- hispidis, glandulosis; bracteis involueri diseo aequalibus, linearibus vel lineari-lan- iam plus minusve piloso-hirsuto-ciliatis a glandulosis, acutis vel acuminatis.— eted at Lake Point, Tooele Co., Utah, alt. 4300 ft., May 20, 1880, Jones 1727 EE NY, RM, U CAL TYPE). [Vor. 22 138 ANNALS OF THE MISSOURI BOTANICAL GARDEN Bannock Co., May 16, 1885, Leonard 45 (NY); gravelly waste ground, Twin Falls, Twin Falls Co., May 9, 1912, Bennitt (RM). UTAH: Big Cottonwood Canyon, vicinity of Salt Lake City, May 23, 1908, Clemens (F, Ph); Fort Douglas, Salt Lake Co., May 5, 1908, Clemens (F); vicinity of Salt Lake City, Salt Lake Co., 1883, Meehan (M, Ph); Garfield, Salt Lake Co., alt. 4300 ft., May 30, 1884, Leonard 66 (G, NY); west of Taylorsville, Salt Lebe Co., May 4, 1916, jon 314 (G); Murray, Salt Lake Co., May 10, 1915, Jones (U CAL); Antelope Is., Davis Co., alt. 5500 ft., June, 1869, Watson 596 (NY, G); western slopes of the Wasatch Range, Springville, Utah Co., alt. 4400—5000 ft., May 17, 1913, Hill (M); Tintie, Juab Co., May 16, 1891, Jones (M, U CAL); Lake Point, Tooele Co., alt. 4300 ft., May 20, 1880, Jones 1727 (F, NY, RM, U CAL TYPE); ‘‘ Hugway,’’ May 29, 1891, Jones (NY); St. George, Washington Co., 1877, Palmer 236 T G, M, NY). on-Juniper Assoc., 3 mi. west of Baker, White Pine Co., alt. 6500 ft., June 16, prie Maguire $ Becraft 2840 (M); 1891, Jones (M); Emigrant Pun. Eureka Co., June 10, 1933, Eastwood $ Howell 203 (M). 13. Balsamorhiza hirsuta Nutt. in Trans. Am. Phil. Soc. II, 7: 349. 1841; Torr. & Gray, Fl. N. Am. 2: 301. 1842; Walp. Rep. Bot. Syst. 2: 610. 1843; Gray, Syn. Fl. N. Am. 1?: 266. 1884; Howell, Fl. Northwest Am. 1: 340. 1900; Piper in Contr. U. S. Nat. Herb. [Fl. State Wash.] 11: 581. 1906; Rydb. Fl. Rocky Mts., ed. 2, 929. 1922, in part; Blake in Contr. U. S. Nat. Herb. [Tidestrom’s Fl. Utah € Nevada] 25: 583. 1925, in part; Jepson, Manual Fl. Pl. Cal. 1078. 1925, in part. Balsamorhiza Hookeri var. hirsuta (Nutt.) Nels. in Coult. € Nels. Bot. Rocky Mts. 546. 1909. Stems subscapose, 1.5—4.5 dm. high, hirsute to hirsute-pilosu- lous, and glandular; leaves lanceolate to oblong-lanceolate, pin- nately divided, 1.5—4.5 dm. long, the divisions pinnatiseet or bipinnatisect, 2.5-6 em. long, .5-2.5 em. broad, the ultimate seg- ments linear to lanceolate, acute, both surfaces green, stri- gose-hirsute, and glandular; petioles strigose-hirsute; heads solitary, radiate, 5-9 em. broad including the rays; braets of the involucre 3—4-seriate, erect or somewhat spreading, ovate- attenuate to an acute apex, as long as or shorter than the disk, pilose-ciliate, hirsute; rays yellow, oblong-lanceolate, 2.5-4 cm. long, 7-8 mm. broad, bifid or entire, tube and outer surface of lamina pubescent ; disk-corollas 8-9 mm. long, the limb hirsutu- lous; achenes linear-oblong, 5-6 mm. long, glabrous. Distribution: dry open grounds, southeastern Washington to northeastern Oregon. 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 139 WASHINGTON: east of Walla Walla, Walla Walla Co., 1834, Nuttall (G, TYPE collection). o N: summit of Cabbage Hill, east of Pendleton, Umatilla Co., June 14, 928, Thompson 4863 (G, M); dry ground, Hot Lake, Union Co., May 19, 1923, pio 363 (F, WU); Hot Lake, Union Co., May m 1923, Sherwood 268 (WU); dry rocky hillsides, Union Co., alt. 3500 ft., June—July, 1881, Cusick 441 (F, G); northeastern Oregon, May 20, 1898, Cusick 1872 (F, G, M, RM, U CAL); stony ridge, ‘‘mouth of Ladd Canyon,’’ northeastern Oregon, June 7, 1898, Cusick 1921 (F, G, M, U CAL). 13a. var. lagocephala W. M. Sharp, var. nov.!5 Stems 0.6-2.5 dm. high, pilose, intermixed with numerous short hairs; leaves pinnately cleft or divided, 1-2 dm. long, the divisions 1-2 em. long, 0.5-1.5 em. broad, cleft into three or more ascending, acute lobes, hirsute on both surfaces, often densely interspersed with glands; bracts of the involucre lan- ceolate, shorter than the disk, the outer series progressively shorter than the inner series, acute, densely tomentose or pilose-tomentose, especially towards the base of the involucre; in other characters similar to the species. Distribution: rocky sagebrush habitats in Lincoln, Grant, and Kittitas Counties of central Washington. Was TON: Davenport, Lincoln Co., alt. 2500 ft., May 20, 1905, Jones (NY); rocky ER Ephrata, Grant Co., April 13, 1924, St. John, Pickett, Cary $ Warren 6314 (Ph); rocky sagebrush plains near Coulee City, Grant Co., May 2, 1931, Thompson 6163 (G); *'Rimrock seabland,’’ north of af ad Kittitas Co., April 23, 1932, Thompson 8229 (M TYPE, NY); sandy soil, Ellensberg, Kittitas Co., May 20, 1897, ju 2718 (G); sage-brush slopes near Vantage, Kittitas Co., April 23, 1932, Thompson 8212 (NY, RM). 13b. var. neglecta W. M. Sharp, var. nov.'® Leaf-segments linear-lanceolate to lanceolate, 2-4 em. long, » B. hirsuta var. lagocephala W. M. Sharp, var. nov., caulibus 0.6-2.5 dm. altis, pilosis et hispidulis; foliis pinnatifidis vel pinnatiseetis, 1-2 dm. longis, segmentis 1— 2 em. longis, 0.5-1.5 em. latis, adscendentibus, aeutis, utrinque hirsutis, saepe glan- dulosis; bracteis involucri lanceolatis, disco coii exterioribus interiorbus plus minusve aas |. dense tomentosis vel piloso-tomentosis, praesertim ad basem involue Cetera speciei similis pss de ki ‘‘ Rimrock seabland,’’ north p? T ni Kittitas Co., Washington, April 23, 1932, Thompson 8229 (M TYPE, NY). ? B. hirsuta var. neglecta W. M. Sharp, var. nov., segmentis folii lineari-lanceo- latis vel lanceolatis, 2-4 em. longis, 1-1.5 em. latis, integris vel pinnatisectis, a presso-hirsutis; bracteis involueri diseo aequalibus, linearibus vel anguste lineari- laneeolatis, dense piloso-tomentosis vel tomentosis. Cetera speciei similis.—Colleeted at Truckee Pass, Washoe Co., Nevada, alt. 4450 ft., May 1, 1909, Heller 9592 (F TYPE, G, NY, Ph) [Vor. 22 140 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1-1.5 em. broad, entire to pinnately parted, appressed-hirsute; braets of the involuere about as long as the disk, linear to nar- rowly linear-lanceolate, densely pilose-tomentose to tomentose; in all other characters similar to the species. Distribution: dry grounds, northeastern Utah west to Washoe Co., Nevada. UTAH: Sheep Creek Canyon, Daggett Co., June 4, 1932, Williams 520 (NY, RM); near Fort Duchesne, Uinta Co., 1913, Carter 4 (RM). NEVADA: Truckee Pass, Washoe Co., alt. 4450 ft., May 1, 1909, Heller 9592 (F TYPE, G, NY, Ph); Trinity Mts., Humboldt Co., May, 1868, Watson 595 (G, NY); near Carson City, Ormsby Co., 1864, Anderson 69 (G); north of Carson City, May, 1865, Stretch 156 (NY). 14. Balsamorhiza terebinthacea (Hook.) Nutt. in Trans. Am. Phil. Soc. II. 7: 349. 1841; Walp. Rep. Bot. Syst. 2: 610. 1843; Gray in Mem. Am. Acad. [Pl. Fendl.] N. S. 4: 82. 1848, in part; Gray, Syn. Fl. N. Am. 1?: 266. 1884; Howell, Fl. Northwest Am. 1: 340. 1900; Piper in Contr. U. S. Nat. Herb. [Fl. State Wash.] 11: 581. 1906; Rydb. Fl. Rocky Mts., ed. 2, 929. 1922, in part. Heliopsis (1) terebinthacea Hook. Fl. Bor.-Am. 1: 310. [1834] 1840; DC Prodr. 5: 551. 1836. Stems subscapose, 2.5-5 dm. tall, crisp-hirsute or crisp- pilose, glandular; leaves lanceolate or oblong-lanceolate, thick, 2-4 dm. long, 5-10 em. broad, pinnately cleft or parted, the lobes sparsely dentate-serrate or entire, acute, strigose-hirsute on both surfaces; heads solitary, 4-6 em. broad including the rays; braets of the involucre linear-lanceolate to ovate-lanceo- late, about as long as the disk, acute, crisp-pilose to crisp-hir- sute, herbaceous ; rays yellow, oblong-cuneate, 2.5-3.5 em. long, 6-9 mm. broad towards the apex, tube and outer surface of the lamina pubescent; achenes 7-8 mm. long, glabrous, epappose. Distribution: southwestern Washington, south to Siskiyou County, California. WASHINGTON: low ground in open pine woods, Falcon Valley, Klickitat Co., June 17, 1922, Suksdorf 10866 (G, M, NY, Ph, U CAL); Klickitat River, May 18, 1884, Suksdorf 360 (G); low grounds in pine woods, Falcon Valley, Suksdorf 135 (G); foothills of Cascade Mts., Yakima Region, 1882, Brandegee 96 (M, U CAL); near Fort Vancouver, **1825,*” Douglas (K TYPE). 1935] SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 141 OREGON: dry stony slope, 12 mi. west of Waldo, Josephine Co. , July 2, 1918, Peck 8456 (WU). CALIFORNIA: Shasta Valley, Siskiyou Co., May 10, 1910, Butler 1337 (U CAL). GREENMANIELLA W. M. Sharp Greenmaniella W. M. Sharp, gen. nov.! Perennial suffruticose plants with alternate ovate leaves. Heads heterogamous, medium-sized, disposed in subeorymbose clusters. Involueral bracts 2-3- -seriate, the outermost series herbaceous or subherbaceous, the two inner series membrana- ceous, marked by six or more brownish-resinous parallel nerves. Receptacle narrowly conical; pales concave to semi- eonduplieate, membranaceous with several brownish resinous parallel nerves. Ray-flowers styliferous and fertile, rays light yellow ; disk-flowers fertile, tube narrowly cylindrical, the base narrowed and seated inside the cup-like crown of the achene, limb campanulate, 5-lobed. Anther appendages ovate, base subsagittate; style-branches recurved, obtuse or acute, pubes- cent towards the apex. Achenes Sübfusiform, broadly sub- angulate, sparsely earuneulate. Pappus a shallow cup-like crown, with knob-like projections at the angles. Greenmaniella resinosa (Watson) W. M. Sharp, comb. nov. Zaluzama resinosa Watson in Proc. Am. Acad. 25: 153. 1890; Rob. € Greenm. in Proc. Am. Acad. 34: 533. 1899. Plants suffrutescent, 1.5-2.5 m. tall; stems branched above, suleate, pilose-tomentulose; leaves ovate, 0.5-3 dm. long, 2- 14 cm. broad, serrate-mucronate, acuminate, acute at the base, triple-veined, substrigose-hirsute above, subpilose on the veins below, interspersed by numerous resinous globules; heads 6- mm. high * Greenmaniella W. M. Sharp, gen. nov. Capitula heterogama, radiata, floribus radii uniseriatis et floribus disci fertilibus. Involuerum subeampanulatum, bracteis 2-3-seriatis, exterioribus herbaeeis, interioribus submembranaceis striatisque. Re- eeptaeulum anguste conicum, paleis eoneavis vel subeonduplieatis, membranaceis striatisque marginibus plus minusve seariosis. Corollae radii ligulae iL s cea - grae, apiei dentatae. Corollae disci tubulo-eampanulatae, regulares, tubo basi non dilatato, limbo late campanulato 5-dentato. Antherae basi su beagittatae, apiei ap- pendieulatae. Styli rami complanati, elongati, obtusi vel acuti. Achenia disei sub- ed subtetragona, breve coronata.—Plantae verisimilliter suffruticosae, foliis ti [Vor. 22 142 ANNALS OF THE MISSOURI BOTANICAL GARDEN Distribution: Nuevo Leon in the Sierra Madre near Monte- rey, Mexico. Mexico: Nuevo LEON—Sierra Madre, near Monterey, Aug. 24, 1889, Pringle 2412 (F, G TYPE, M, , U8); Sierra Madre, above Monterey, 3000 ft. alt., Aug. 25, 1903, Pringle 11615 (F, G, US); Sierra Madre, ‘‘Conquor,’’ near Monte- rey, July 9, 1888, Pringle 2222 (G); vicinity of Monterey, 750 m. alt., Aug. 1911, Arséne 6176 (M). List or ExsiccATAE varor numbers are indicated by italics; the collections without numbers mbers in parentheses refer to the species number The are indicated used i The genus, in each case, precedes its list of n this pi hom een exsieeatae. BALSAMORHIZA Cusiek, W. C. 595, 3255, 1679, 3332, Abrams, L. R., & E. A. McGregor. 356, 4). Anderson, C. L. 69 (13b). Anderson, F — (5). Austin, R. M. 94, 47 (5);—, — (7). Austin, R. M. & un 2179 EPR Ayres, W. B. 63 (5 Baker, C. F. 51, 926 ( J Baker, M. S. & F. P. Nutting. — (7). Bollinger, G. C. — (11). — (4); — (8). Blankinship, J. W. — (4); 52 (5); (10). Bolander, H. N. 4526 (2); 6187 (5). M. Brandell, E Brandegee, K. — 0); — (4); 208 (8); 2469 (8 ; 891 m ;—, Cooper, R. D. 1182 (5). Cotton, J. S. 1105, 1052, 976 (3); 336 (4); 568 (6a). Crandall, C. L. 2723 (5). Culbertson, J. W. 4413 (1). Curtis, C. C. — (10). (3); 1984 (11); 441, 1872, 1921 13). Darlington, H. T. 54 (11). Davidson, A. 2058 (1). D 21 avy, 85 (4) Douglas, — (14 rake & Dickson. — (4) Drew, E — Dudley, W. B. 363 . Eastwood, A. 11927, 3237 (4); 896 ( Eastwood, A. & T. Howell. 203 (12); 507 (10). Eggleston, W. W. 6820 (4); 6964 (11). Elmer, A. D. E. — P Engelmann, G. — Ferris, R. S. 1580 v Gabrielson, — (5). Gale, N. P. 111 (4); 178, — (9); 100 (11). Gardner, N. S. — (4). Gorman, — (4). Grant, J. M. — Greene, E. L. — mu 718 (4); ?45 (6 ). Griffiths, D. € J. S. Cotton. 483 (3). Griffiths, D. pot Hunter. 77 (9). Hall, E. 276 Hall, H. M. dn 8907, 11246, 7797 uin 1032 (7); 8885 (8). & G. R. 8340 (1). aa H. M. & H. D. Babeock. 5100 (4) 1935] SHARP—CERTAIN EPAPPOSE GENERA OF COMPOSITAE 143 Hall H. M. & H. P. Chandler. ?304 4). Hammond, E. W. 209 (4). Hansen, G. 168 E. 420 (5); 1069 (8). Hayden, F. V. —, — (9). Heller A. A. 11977, 7802 (4) ; 14594, 12026 (5); 9592 (13b pus A. A. & E. G. 2958 (5); 8298 PEU L. F. 362, 5902 (4); 5708 (11). — (12). Jones, M. E. — (3); 6407 a c: 1727, — (12) ; — (13a Jones, b W. 487 (8); — (9); — 314 ee leg A. L. 60 (4). Kellogg, A. & W. G. W. Harford. 430 8). Kelsey, F. D. — (5) Kennedy, P. B. M (8) 1859 (13a). Kirkwood, J. E. 2 (5). Konger, K. id ^ Leiberg, J. B. 142, 28 (3); — (5); 114 (5); 2106, 83, 58 (11). Leonard, F. E. — (10); 66, 45 (12). Lloyd, F. E. — (10). Lunell, J. 6? (4). Lyall, D. (4). Macbride, E. F. 202, 79 (5); 820 (10a). Macoun, J. 86299, 76941, 953 (4); = (5). , Maguire, B. & R. J. Beeraft. 2340 (12). Meehan, a — (12). Merrill, E. D. & E. N. Wilcox. 686 (5); 743 Si: pr (10). Mooney, W. T 6b). . E, E Murdock, J. 4096 (5). Nelson, A. 7084, 9140, 4518 (5); 2342 (9); 4015 SU $552 (12). Nelson, A. & E. 3 (5). Nelson, E. 1476 (4); 1595 (6); 4993 9 Nelson, J. C. E p ys (4). Newcombe, C. F. 62 (4). Nuttall, T. — Wo — (5) — (6); — (9); Osterhout, G. E. 6212 (10). Page, E. B. 216 (4) Palmer, E. J. — - 236 (12). Pammel, L. H. Pammel, L. H. e E. a Stanton. 322 (9). Parry, C. C. 165 (9). Payson, E. B. 343 (5). Payson, E. B. & L. B. 2605 (10). & G. M. Armstrong. Peck, M. E. 5040, 13249, 3069 (4); 15544, 3071 (5); 13900, 15264, bi aD; ; 8466 (1 4). Piper — (3); —, 4945 (4); 1594 p gus (15a). Prescott, H. S. 11 (4). Ramaley, F. Thies (10). Rich, Maj. — (2). Rydberg, P. A. 808 (5); —, 6082 (10). Rydberg, P. A. & E. A. Bessey. 5174 t. John, H. & R. H. Brown. 3900 St. Jehu, Pickett, Cary & Warren. 6314 (18a). Sandberg, J. H. & J. B. Leiberg. 166, — (3); 573 xL 23 (5). Sandberg, J. H., D. T. MacDougal & A. A. Heller. i E. 248 (9a). Sehmoll, H. M. & D. Nusbaum. 1574 5). Sharp, $. S. Shear, C. L. is. 3009 (10). Sherwood, W. 372 (5); 323 (9); 268, 363 (13). Smith, C. P. 2348 (5); 2366 (10). 368, — (7). Spalding, — (3); — (9). Streteh, 156 po Suksdorf, W. N. — (4); 357, —, 363, 359 he ' 10866, 360, 135 (14). 144 Thompson, J. W. 5171 (3); 5133, 561, 2495, 2593, 2163 (4); 6126, 6386, 5985 (5); 227 (7); 4863 (13); 6163, 8229, 8212 (13a). Torrey, J. — (2); — (8). Tracy, J. P. 4175, 6662, 6984 (4). Tweedy, F. 2115, 11, 3150 (9); 214, 66 (5). Walker, H. A. 1285 ($); 2016 (5). Warren, E. D. Watson, S. 596 (12); E (10); 595 3b). liams, L. 1125, 415 (5); 1317 (10); 520 (13b). Williamson, J. W. 185 (4). Zeller, S. M. — (4). ECHINACEA Bartholomew, E. — (3). Biltmore, Herb. 1107 (3a). Blakley, 0. W. 1436 (4). Buekley, S. B. — "T ) Burk, M. ??2 Bush, B. F. —, —, 6 (1); 38, 149, 156, = "M y^: pd 155 (5). Carr, W. P. 90 (3). Chase, V. H. p^ ay) 642 (4). Churchill, J. R. Clemens, Mrs. ie. B (3). Clemens, Mr. no Jos. 381 (3). ph 2 (2). Davis, J. 4864, 1574, 3284, 1844, 5819 (1); 4185, 4405 (4). pia ). Goodding, L. N. 2215 (3) Graves, — reenman, J. M. 4007 (1); 412 (4). Hall, E. 330 Harper, R. M. p (1). Hedgcock, G. G. — (4). Heller, A. A. 1735, 13984 (3). Hitchcock, A. 8. — (4). [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Houghton, H. 2 rs (5). House, H. D. iiid Meade, S. B. 1841 (1). Murrill, W. A. — (2). Nelson, A. 2122, pe ME — (4). Norton, J. B. 8. Palmer, E. J. nili. p 5843 (1); 37546 (3); 178, 1206, 5783, 7632, 8061, 18088, 18016, 34749, 69853 (4); 5844, 9617, 30397, 34714 (5). Pammel, L. H. 243 (4). Pammel, L. H. & Stanton, 24? (3). ` Price, S. F. Reverchon, La ==, 2579 (4). Rose, G. M. 3). Russell, C. — a). Ruth, A. 243, 646 (5). Rydberg, P. A. & R. pra 400 (4). Stephenson, B. C. — Stevens, G. W. 737, n 1341K (3). 3). Zeller, C. — GREENMANIELLA Arséne, G. 6176 (1). Pringle, C. G. 2222, 2412, 11615 (1). IOSTEPHANE Arséne, G. 2464, 5518 (1); 1056 la Arsène, G. & Nicolas. 5408 (1). Ghiesbreght, A. "We. 561 (2). Holway, E. W. D. 3543 (1a). Nelson, E. W. podre 3223 (2). 1935] DS pu s TONG- He "Tun ac SHARP—-CERTAIN EPAPPOSE GENERA OF COMPOSITAE 145 Palmer, E. 65 (1); 333, 4824 (la); 72 (1b). Parry, C. C. € E. Palmer. 470 (1a). Pringle, C. G. 5249, 9276, 9929 (1); 4480 (1b); 4978 Purpus, C. A. 5694, (1a) ; 4120, 55 (2 Rose, 3. N. 2336, 2969 (1a). Rose, J. N. & J. H. Painter. 6493 a). Schaffner, J. G. 58, 253 (1). Seaton, H. E. 366 (la). np crum Anderson, J. P. — (4) fei a. 2603 (1). Arsène, G. & A. Benedict. 15445, 15841 2). Baker, C. F. — m; 667 (2). Barlow, B Bebb, R. 35 Se J 4 3 (1). Brandegee, us ph 12051 (2). Burk, M y Bush, te F. 157, 163, 307 p (2); 8407 (la); —, 123, 164 (4). YA . 0 (4 Davis, J. 1390, 5114, 7320 a); — (4). Deam, C. C. 1370 (4) Demaree, D. 7706 a 7706a (la); rummond, T. 108 (3). Drushel, J. R. 419 (1a). Dunean, J. T. — (1); 28 (1a). Earle, F. S. & E. S. 150 (2). Eggert, H. — —, — (1); — — (la); — (2); — (4). Eggleston, W. W. — (1); 20562 Ellis, C. C. 208 (2). Ellis, L. D. — (1). Emig, W. H. 366, ?72 m Engelmann, G. — Ai Evans, W. H. — (2). Fendler, A. 378 (la); — (2). Ferris, R. S. & C. D. Duncan. 2838 (1); 3113 (la). Fisher, G. L. — (1); — (la). Fritchey, J. Q. A. 167 (1a). Glatfelter, N. M. — (4). Goodman, G. J. 226 (1). Greenman, J. M. 3624 (4). Gregg, J. 810 (1a). Griffiths, D. 5641 (2). — (1a). Heller, Fr E 13975, 1850 (1); 3829 (2); 1789 (3). edie pi S. Rs (1); 268 (2). Johnston, E. R. j^ Jones, M. 908 (1); 25904 (1a); 29506, Ze = ye Kellerman fk A. — (1a). Kramer, J. 0 (1 e Larsen, E. = es 194 mop" Letterman, G. — (1); = 14). Macoun, J. 34829 (4). Mathias, M. E. 309, 526 (1). Metealf, F. P. 1896 (4). Metcalfe, O. B. 415 (la). Moore, J. A. & J. A. Steyermark. 3618 (1). Mulford, A. I. — =; 260 (2). du dd Oyster, J. H. 4149 (1). r, E. J. 7746, 12535, 13878, 31021, 37528 (1); 7792, 8024 (la); 12534, 31100 (2); 713 (3a); 6003, 6204, 8037, 17003 s Pammel, L. H. — (1); 17 (4). Pammel, L. H. & I. C. Au 56 (4). Patterson, H. N. — (4). Payson, E. B. 1164 (1a). Pellet, H. L. ). Poole, R. J. & D. ^ am — (1). Price, S. F. Pringi c. G. bee (2); 1648, 1305 Pl C. A. 4768 (1). 146 Redfield, J. H. 458 (1). Reed, M. 2). Reverchon, J. 493, 3337, 3338 (la); —, 3340 (2); 2229 (3a). Rusby, EL. E. 78 (2). — P. A. & R. Imler. 336 (1); 8 (2). Sedis, J. I. 133a (4). 1). Skehan, J Smith, J. G. € R. Pound. 145 (la). Standley, P. C. 5124, 5279 (1); 4926 (1a); —, 4950 (2); 5156 (2a). Stephenson, B. C. — (4). Stevens, G. W. 764 (1); 1080 (2); 2283 (4). Steyermark, J. A. 1036 (4). Stratton, R. 444, 399 (1). Thompson, C. H. — (1); — (1a); — 2). ( Thornber, J. J. —, — (1); — (18). Townsend, C. H. & c. M. Barber. 230 (1); 101 (5). Tracy, S. M. 7891 (1); 8947a (la); 8548 (3); 8914 (3a). Wislizenus, A. 25 (2). Wooton, E. O. 261 (1); 247 (la); 5 (2). libro E. O. & P. C. Standley. 3315 we C. $27 (2); — (3); — (3a). ZALUZANIA Altamirano, F. 1744 Arséne, G. & Nicolas. 6162 (13). Botteri, M. 843 (3). Bourgeau, E. 803 (13); 385 (1); 704 (3). Conzatti, ek 2317, 4601 (8). Coulter, e . 350 (9). Deam, C 5? (3). Duges, 1 " (8); —, 469 (13). Firmin, G. me Bei (14). Fisher, G. L. [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Goodding, L. N. 864 (5). 631 (4). Hartweg, T. 111 (1a). Holway, E. W. D. 534? (8); 9 (14). Kirkwood, J. E. 67, (11). Lemmon, J. G. —, —, —, 2764 (5). Lloyd, F. E. 124 Ay 82 (11). (9). 227 (4); Mille, L. 568, um Qo. 4970 0 (7). Orcutt, C. 4. 4139 (1); 3902, 4181 (3). Pachano, A. 134 he 4). Palmer, E. 307 (la); 757 (3); 279, 360, 435, 592 pre 734 (10); 314 (11). Parry, C. C. 446 (11). Parry, C. C. & E. Palmer. 527 (1); (3). Pittier, H. 452 (12). Pringle, C. G. 2925, 3204, 7440, 9395, 13092 (1); 8914 (la); 5156, 7367 (2); 9481, 13093 (3); 2402 (4); 755, 1310, 2145 (5); 8710 (6); 309, 1110 (7); 4928 (8); 6956, 7923, 9997, 13784 (9); 2398, 10076, (10); 3144, 7451, 9996, 11614, 13091 (13). Purpus, C. A. 1); 1126, 1334 (4); 2530, 3823, 3824, 5610 (8); “a 3027, 4121 Qn; 1539 (13). f Hay. 5274, 5816 a): 5954 pe N. J. H. Painter. 6675 (18). Rose, J. N., J. H. Painter & J. S. Rose. 8417 Qi 8325 (3); 7278, 8418, 9738 Ma 4 G. 80 (1); 345 (1a); 78 (3); 717 (4); 767 (11). ( Wilcox, T. E. Wislizenus, A, pr? (7). (8). Sodiro, A. fi E 31/3 (14). 9 (5). 1935] SHARP— CERTAIN EPAPPOSE GENERA OF COMPOSITAE 147 INDEX TO GENERA AND SPECIES New names and new combinations are printed in bold face type; previously published accepted names in Roman type; and synonyms in italics, Ant is Page LODOS eh TESTES 102 CIO ar Rw 112 Ur LO PE EE S 104 ld EVE Ro PEERS D CI 104 irMobata- 7.2. oe on Hoses 104 ¡RISA eels LS hove ere acer DI 114 BOLAS ra o DPA 123 A o ol anor 124 delioIlde8n nn cere: ventur CETT 125 glabresoena nenne een 125 helianthotdes ..::...0esee neue 127 DITSUUR O 138 var. lagocephala ............ 139 va CUA 520 ET 139 hispidula- os ea 137 Hooker iia a 129 var.lanatá id 9 98 130 v IOS6N NS 130 Hookeri var. hirsuta .......... 138 MCNA DTe ones UE TORO 133 var. tomentosa ......2 5.95.5 184 JDVenusta «cvs eve eH ID 122 Macrolepis i ooo. de ES 132 macrophylla sesa eurae EE 134 var. idahoensis ......s..sss 136 er: var. terebinthacea 135 ge ee 131 Rao 130 ina e oe PEATE 127 ^ MCN REESE a y 136 terebinthacean -s.s 2. 92 2 Ue 140 BalsamorrhMsGa .......... ev 119 Balsamorrhiea .... or ne 129 ens du a Se g EN 133 PCENA VAI. MONA... ae ee 133 FOMENLOSO v. cr. ere 134 Hohner oer peti TA 88 E m r.i E Goes ers 88 angustifolia err e s 91 EU dn s ips Res e soie e 91 HT DESTRUERE TEC ET. 93 Page Darüdotà jo een 94 Durpurea.. 102 BÀ ener. 89 Yentesscensia A TES ee 9 92 Buphthalmum sagittatum ........ 127 ChMWliophylum |...... een 100 globomm. 20220 ib ae 102 Chrysanthemum americanum ..... 89 a ee Don re 100 FE N Er 112 cores heteros IN PU quo 81 A A 88 E oni latifolius ......... 89 Ulp en MNT een ea 84 anpuslifolia 1:7... 0 2 20 4 91 var. tennesseensis ........... 92 angustifolia onc ch ce dr adie ove 93 IDRORBONM. Ci hie ek 82 i Eolo nere ere pd 81 heterophulla bee nen 81 NRIETMOMG 1.1... ree 89 o Ear OO SES 91 (vs A Sree LII 93 BERUOOXIE s vvv ev che TT 94 DUDZUUTGHÓO he ee ee 89 SONDIMMER 4 crises se A TTT 91 TOWMESBOONDIS occ cee secs repete 93 Espeletia A A iX oc ate verbs 127 A tr. PR ST 127 A A user RO e 100 A AA re EY 112 var, megacephala ....... ess 110 CODI VERE ERR TERT UE 112 montagnaefolia ............... 108 Gree Be ON 141 re A AN 141 Gymnolomia E e O rs 83 Lin Gk A elo na 106 Helianthus IOCONUSTUS a cc pen 122 148 Page PU AAA cesas BO Heliopsis Bals e A EN 1 terebinthacea ........... 140 Hybridella .............. a 100 CRORE alv 102 A AA eee eT e TT subg. Chrysopetala ........... 81 ubg. Euiostephane ........... 80 ann! m ¿cio ros 81 . Di TUIS E 82 var, acutiloba .............. 83 a A EET 83 LODBNMER ir xs 66 ODE ernennen nen en 68 var chorrimo ............ 70 AA O 72 DOM aan 68 THU Visas xa deese VS 76 podunoularis ....oommmooo rosso. 73 CON Pi A rv x 74 A TE ET 75 is AAA Sac 75 1, A ee 74 a ER LETT TE 72 A A CEN CIIM ER 67 AN Loos. asus eR X xn 68 s as TEST TEEN 75 puloherrima ........ ere 70 TUNI Td sao IO 72 o ta LIT 66 "pdt V.P ecu T 68 ar oherrima ............ 70 forma pulcherrima ............ 70 eolumMifera .......... eee 68 var. pulcherrima ............ 71 RM TN DédUBAUSME iria 73 FOL, DIOR A Mey 73 o A O 74 [VoL. 22, 1935] ANNALS OF THE MISSOURI BOTANICAL GARDEN Page A ansehen 68 A anne 6 i | VOL, OMON eee e ars 73 Rudbeckia INNUIIIIAO visados 83 o A MEREREETETETTTTET TT 68 oolumnifera .... eee enn 68 A inserieren een 81 SOMOS obs O 93 MOTO anne 75 wo d. acini sae 89 bars 634s tea UU TU TI 89 TI ia 72 AR een bows 95 DOI Aa 112 anthemidifolia ................ 104 DEMNE A er aS 108 var. montagnaefolia ......... 108 [DoD PREET TEETE 112 (nt tnpr nn 109 EPPS TET A . 109 QNNM Sana eT TT 107 subg. Ferdinanda ............. 101 iae E ETE EET IE T EE TY T ETT ERIT 102 var myHophslla- osc cites 103 BIO ii 103 o rer 106 DANA cian Fes sik cae ae 106 subg. Mybridella .............. 101 megacephala ............ e 110 AAA . 110 montagnaefolia ............... 108 myriophylla ......omommo.m..».». 103 nro arr PENDISSE T 113 PEGI ias TEE a 107 CS Loiss ERAT WEKG XY VETE 113 TOOWONG iii a swiss 141 ob "al Iu TEES 105 o ua PETS 113 pi AA 111 SS UA 104 EXPLANATION OF PLATE PLATE 5 Disk-flower of Echinacea pallida. Corolla above, achene below. x 11. Disk-flower of Ratibida columnaris. a. Achene, side view. Nap x 9. Disk-flower of Balsamorhiza hispidula. Corolla above, achene below. Disk-flower of Iostephane heterophylla. x 7.5. Disk-flower of Zaluzania mollissima. x 9. ANN. Mo. Bor. GARD., Vor. 22, 1935 T dv NUR LU + 5 SHARP—EPAPPOSE GENERA OF COMPOSITAE PLATE 6 22, 1935 ANN. Mo. Bor. GARD., Vor. —EPAPPOSE GENERA OF COMPOSITAE SHARP Annals of the Missouri Botanical Garden Vol. 22 APRIL, 1935 No. 2 STUDIES IN THE APOCYNACEAE. IV! THE AMERICAN GENERA OF ECHITOIDEAE ROBERT E. WOODSON, JR. Research Assistant, Missouri Botanical Garden Instructor in Botany, Henry Shaw School of Botany of Washington University V. Forsteronia G. F. W. Meyer Forsteronia G. F. W. Meyer, Fl. Esseq. 133. 1818; A. DC. in DC. Prodr. 8: 436. 1844; Muell.-Arg. in Mart. Fl. Bras. 61: 95. 1860; Benth. & Hook. Gen. Pl. 2: 710. 1876; Miers, Apoc. So. Am, 242. 1878; K. Sch. in Engl. € Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Syringosma Mart. ex. Rehb. Consp. 134. 1828. Thyrsanthus Benth. in Hook. Jour. Bot. 3: 245. 1841; A. DC. loc. cit. 386. 1844; Miers, loc. cit. 93. 1878, nec Ell. nec Sehrank. Aptotheca Miers, loc. cit, 150. 1878. Lactescent, fruticose or suffruticose lianas. Stems volubile, terete, the branches opposite or alternate. Leaves opposite, or infrequently ternate or quaternate in certain species, entire, penninerved, the upper surface bearing few to several glan- dular emergences at the base of the midrib, or eglandular, the lower surface bearing with more or less constaney a single rather inconspicuous, lenticular foveum or pit in the axils of the midrib and principal primary veins ; nodes ineonspicuously * Continued from ANN. Mo. Bor. Garp. 20: 605-790. (1)-(186). 1933. Issued May 25, 1935. ANN. Mo. Bor. GARD., Vou. 22,1935. (187) (153) [Vor. 22 154 ANNALS OF THE MISSOURI BOTANICAL GARDEN stipulate. Inflorescence terminal or both terminal and lateral, aggregate diehasial to thyrsiform (simple in F. simulans). Calyx 5-parted, the lobes essentially equal, cleft nearly to the receptacle, imbrieated, bearing within one to several squa- mellae, or the squamellae evidently lacking. Corolla rotate or subrotate, the tube relatively short, exappendieulate within, the orifice not annulate, the limb actinomorphio, 5-parted, dextrorsely, or rarely sinistrorsely, convolute. Stamens 5, the anthers wholly exserted to essentially included, connivent and agglutinated to the stigma, consisting of 2 parallel, uniformly fertile sporangia borne ventrally near the apex of an enlarged, sagittate, peltate connective; pollen granular; filament free or agglutinated to the style. Carpels 2, apocarpous, or very rarely syncarpous, united at the apex by the fusiform or sub- capitate stigma; ovules many, several-seriate, borne upon an axile, binate placenta. Nectaries 5, separate or more or less concrescent. Follicles 2, apocarpous, or more or less ag- glutinated, dehiscing along the ventral suture, containing many dry, truncate, apically comose seeds. Type species: Forsteronia spicata (Jacq.) G. F. W. Meyer, Fl. Esseq. 133. 1818 KEY TO THE SUBGENERA A. Stamen filaments exappendiculate........o........ Subgen. I. EUFORSTERONIA AA. Stamen filaments with conspieuous, membranaceous > ere IA E ADA Seo Subgen. II. Pu d Subgen. I. EvronsrERoNIA Woodson, subgen. nov, Stamen filaments exappendiculate, the anthers merely acu- minate with more or less evident, hyaline tips. Spp. 1-46. KEY TO THE SPECIES a. Anther-tips exserted, often barely so, or essentially included; filaments distinet and free, not agglutinated to the style . Anthers 0.07—0.1 em. long, the tips barely exserted or essentially included. e. Ultimate branches of the inflorescences regularly and determinately composed, not indefinitely congested. d. Leaves denis usually obscurely so. e. Corolla-lobes minutely papillate or essentially glabrous within; anther-tips minutely barbellate; plants of British Guiana and DoMBern Brasil, erara vo cenar raro Ed 1. F. gracilis (188) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 155 ee. Corolla-lobes eonspieuously pilosulose within; anther- -tips p or essentially so; plants of southern Bra zil TT TES . pilosa dd. Leaves ae cordate e. Anther-tips glabrous or essentially so. f. Corolla 0.2-0.25 em. long8..........oooooooooooo». 3. F. Luschnathi ff. Corolla 0.3-0.35 em. long............o.oooooooooos.o 6. F. Gardneri ee. Anther-tips abundantly barbellate. Caly eite ovate-deltoid, obtuse; squamellae numerous, indef- ely and regularly distributed; plants of Venezuela an zn olo TC T nee 4. ff. Calyx-lobes lanceolate to ovate-lanceolate, acute to acuminate; squamellae relatively few, in alternate groups; plants of Peru F. graciloides ec. poem Eu of the inflorescences indefinitely congue’, at least F The LOWER PTT a ae . affinis bb. Anthers 1 fore em. long, the tips usually manifestly exserted. e. Stem and leaves ferruginous-tomentulose............ lesus iu... F. rufa ec. Stem and leaves glabrous or essentially so. d. Ovary manifestly 2-lobed, apoearpous. e. Inflorescences typically thyrsiform or eorymbose. f. Stigma wholly included among the connate anthers. g. Leaves glandular at the base of the midrib above. h. Calyx-lobes somewhat shorter than the corolla-tube to very slightly longer; species of Central and South America. i Anther-tips more or less barbellate; leaves 8-20 em. long. j. Leaves membranaceous; plants of the Guianas and Trinidad. «equi aequi A Bes 9. F. Acouci jj. Leaves coriaceous or subeoriaceou k. Corolla very minutely and PE papillate to 3 E glabrous without; plants of northern zil and adjaeent Peru.......... thamiana kk, Gore densely and uniformly Baberdilet- papilia e without; plants of British Honduras. .11. F. viridescens ii. Anther-tips glabrous; leaves 3-9 em. lon j. Inflorescences pyramidal; calyx-lobes ovate, aeute to acuminate; plants of the Guianas and the lower alle 12. F. Amazon Valo aa etc. voce cece 3. F. Ed a hh. = lobes much longer than the UE plants of RE arena en 14. F. Wilsonii gg. Dra eglandular. h. Leaves heavily coriaceous; calyx-lobes t > cm. long in A I HE . F. diospyrifolia hh. Leaves firmly membranaeeous; calyx- dies D 12 em. Ud cA C M . F. Riedelü ong ff. Stigma somewhat exserted beyond the connate MEM (189) [VoL. 22 156 ANNALS OF THE MISSOURI BOTANICAL GARDEN g. Inflorescences about as long as the subtending leaves, or nearly 80, — lax and diffuse; ealyx-lobes 0.075-0.1 em. long, 17. F. Duckei gg. arie diii much shorter than the subtending leaves, rela- tively congested; calyx-lobes 0.1-0.17 em. — aeute to O . F. laurifolia ee, Inflorescences compound-subspiciform, the branches very y or and nitely congested..........oooooooooooooomos» . F. paludosa dd. Ovary superficially more or less 5-lobed by the pressure = the nec- taries, more or less completely synearpous toward the base, the fruit, however, appearing normal and apocarpous. e. Braets T: conspicuous, as large as the calyx-lobes or somewhat arger, at least in part, more or less foliaceous. f. Individusl pedicels subtended by at least one braet conspicuously larger than the ealyx-lobes; plants of Bolivia....20. F. amblybasis ff. Individual pedicels subtended by bracts smaller iun the calyx- lobes; plants of Peru.......ee eee eene ntn 21. F. decipiens ee. Braets relatively ineonspieuous s throughout smaller than the ealyx- carious; plants of east-central Brazil......... . F. montana lobes aa. Anthers whalty exserted; filaments on to the style, at least above. b. Leaves glandular m the base of the midrib above; speeies of South and Central America (41 also indigenous to Cuba). e. Inflorescences aggregate dichasial to thyrsiform, obviously compound. d. Ovary a — fruit bifollicular; species of Central and South America gene e. Squamellae numerous, opposite the calyx-lobes or regularly and indefinitely distributed. i Inflorescences rather narrow, usually much longer than broad; corolla "ord ree e .28. F. — 1L Tudo recaen subeorym , broader than long; tips a-lobes de menia papi WibhoWb....ooomooonorrosisossrsrodares..» . F. cordata hh. Calyx- pres broadly obtuse or rounded............ n p. mollis gg. Leaves glabrous beneath, or merely minutely barbellate in the axils of the midrib. h. Corolla essentially glabrous without. i. Calyx-lobes acute to acuminate. j. Leaves obscurely cordate, wholly glabrous beneat plants o ad 26. F. ne D jj. Leaves obtuse or rounded at the base, minutely bar- bellate in the axils of the midrib beneath; plants of il 27. F. leptocarpa T.oscecsecosceceeceeocoonse OUNAE. Gcr rwr bx do ii. Calyx-lobes broadly obtuse or rounded. j. Peduncles and pedicels densely and conspicuously puber- ulent to tomentulose. 190) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 157 k. Leaves 7-12 em. long, 4.0-7.5 em. broad; corolla-lobes very minutely papillate or essentially R within; plants of northern Bolivia...... 25. mollis kk. Leaves 3.5-10.0 em. long, 2.0—5.5 em. qe E Kin lobes puberulent within; plants of Pos Brazil A ee SE S RITE E VS 28. F. australis jj. Peduncles and pedicels very el and indefinitely papillate to essentially gla k. Leaves rather delieately TE 4.5-8.0 em. long, 2-4 em. broad; plants of Venezuela............ ved PPP de ST 29. F. obtusiloba kk. Leaves USC pd to firmly membranaceous, 3.0— 6.5 em. , 1.2-2.7 em. broad; plants of southern Brazil Be buen: Bolivia, Mp Argentina, and UPS issues Acre eros . F. glabrescens hh. Corolla densely puberulent or puberulent- api without T PA o EI ner E F. tarapotensis ff. Corolla 0.35—0.37 em. long. g. Corolla white, the lobes minutely and rather irregularly pilosulose within; plants of Central America....32. F. myriantha gg. Corolla yellowish-green, the lobes glabrous within; plants of northwestern PUR. VII e o cosonocccrcoso»i 33. F. galbina ee. a alternate with the calyx-lobes, solitary, or infrequently in groups of 2-3, f. Bracts conspicuous, and more or less foliaceous (except in 37), persistent, at least ar g. Bracts Be as large as the calyx-lobes or larger, at least in part h. Species of South America. i. Inflorescences rather narrowly thyrsiform; P 13-35 eu JE E AU POCA . F. thyrsoidea ii. Inflorescences subeorymbose-thyrsiform ; ak 5-10 em. longe e has Weg aO E er Fea ss 35. F. Velloziana hh. Plants of Central America; inflorescences thyrsiform-sub- egg. the lateral branches virtually lacking; follicles unknown... 6 ci ee RE cc ck 36. F. chiriquensis gg. a. relatively inconspicuous, smaller than the ealyx-lobes; inflorescences thyrsiform- en the lateral branches Du laeking; plants of Ecuador........ 37. F. Pycnothyrsus ff. Bracts inconspicuous, scarious or only slightly foliaceous, aducous. g. Inflorescences rather narrowly pyramidal; anthers glabrous sr ai aor 38. F. adenobasis gg. Inflorescences broad and subcorymbose; anther-tips minutely uberulent or barbellate. h. Leaves densely and minutely tomentulose beneath; ealyx- lobes 0.15-0.17 em. al... E... 39. F. umbellata (191) [Vor. 22 158 ANNALS OF THE MISSOURI BOTANICAL GARDEN hh. Leaves very minutely and rs omg pe papillate beneath; calyx-lobes 0.3-0.35 em. long......... dd. Ovary syncarpous; fruit a bicarpellary false UE ; plante of Cuba and locally in southern Mexico, Guatemala, Niearagua and Atlantie BER OBERES O 41. F. spicata . Inflorescences ze. corymbose or subumbellate, simple....42. F. simulans bb. pen d at the base of the midrib above; species of the Greater Antilles (44 ot 1 British Honduras). e. Corolla greenish-white. d. Seeondary venation of leaves relatively dense, subhorizontal; plan ANI UV rra aaa oa 43. F. Pa dd. Secondary venation of leaves relatively distant, jor arcuate ; plants of British Honduras.......).....o.....0.... . F. peninsularis ee. Corolla deep red or somewhat purplish, very rarely pale pode d. Follieles relatively slender, acuminate, reflexed-divaricate, 14-19 em. ine Po lobes glabrous within, or the N minutely and regularly ciliolate; plants of Porto Rieo.......... 5. F. portoricensis dd. Follicles Pens stout and rigid, tiunt, sharply Piu nearly rectilinear, 11-14 em. long; corolla-lobes minutely an ge ather ir- regularly papillate within, rarely glabrate; plants of Cuba and EN AAA AA o E ERÀ 46. F. corymbosa 1. Forsteronia gracilis (Benth.) Muell.-Arg. in Mart. Fl. Bras. 6t: 101. 1860. Thyrsanthus ? gracilis Benth. in Hook. Jour. Bot. 3: 246. 1841; Miers, Apoc. So. Am. 99. 1878. Stems relatively slender, glabrous, conspicuously lenticel- late when fully mature; leaves opposite, shortly petiolate, oblong-elliptic, apex obtusely subcaudate-acuminate, base broadly and rather obscurely cordate, 5-11 em. long, 3-5 em. broad, firmly membranaceous, rather inconspicuously glandu- lar at the base of the midrib above, glabrous, or minutely bar- bellate in the axils of the midrib beneath; petioles 0.4—0.7 em. long; inflorescence rather loosely thyrsiform, both terminal and lateral, the terminal somewhat surpassing, and the lateral somewhat shorter than the subtending leaves, bearing many small white flowers; primary peduncle glabrous, ultimate branches regularly and determinately composed, densely and minutely puberulent-papillate; pedicels 0.1-0.2 em. long, min- utely puberulent-papillate ; bracts lanceolate, 0.08—0.1 em. long, scarious; calyx-lobes ovate, acute to acuminate, 0.08-0.15 cm. long, scarious, minutely papillate without, the squamellae ex- (192) 1935] j WOODSON—STUDIES IN THE APOCYNACEAE. IV 159 tremely minute, flagelliform, alternate, solitary, or evidently lacking; corolla densely and minutely puberulent-papillate without, the tube 0.05-0.11 cm. long, about 0.05-0.075 em. in diameter at the base and 0.1—0.12 em. in diameter at the orifice, minutely villous within at the insertion of the stamens, the lobes ovate-oblong, 0.11-0.2 em. long, spreading, minutely papillate or essentially glabrous within ; stamen filaments 0.02- 0.05 em. long, distinct and free, not agglutinated to the style, the anthers essentially included or the tips barely exserted, 0.07-0.1 em. long, minutely barbellate; ovary ovoid, apocar- pous, about 0.025 cm. long, minutely villosulose; nectaries somewhat shorter than the ovary ; stigma blunt, 0.03-0.07 em. long; follicles rather slender, essentially continuous, some- what falcate, 24-26 cm. long, glabrous; seeds about 2.5 cm. long, the tawny coma about 4 em. long. BRITISH GUIANA: Moruka River, Pomeroon District, July, 1927, Cruz 4584 (FM, MBG, NY, US); Wanama River, Northwest District, May 10-23, 1923, Cruz 3924 (FM, MBG, NY, US); Essequibo River, Moraballi Creek, near Bartiea, alt. near sea-level, Oct. 16, 1929, Sandwith 467 (DC, NY, 8, U, US); Curassawaka, date lacking, Schomburgk 608 (Camb., DC, FM, K, TYPE, US, V, MBG, photograph and analytical drawings) ; Rupununi River, May, 1843, Schomburgk 953 (B, BB, Camb., DC, V); Barima River, March, 1896, Jenman 6965 (NY); upper Demerara River, Sept., 1887, Jenman 4058 (NY, US). BRAZIL: AMAZONAS: Manáos, Oct., 1912, Kuhlmann 28382 (B, U, US) Uypiranga prope Manáos, ad ripas fluvii Rio Negro, Dec. 22, 1923, Kuhlmann 21863 (B, U, US). 2. Forsteronia pilosa (Vell.) Muell.-Arg. in Mart. Fl. Bras. 61: 99. 1860. Echites pilosa Vell. Fl. Flum. 112. 1830; Icon. 3: pl. 38. 1827. Thyrsanthus pilosus (Vell.) A. DC. in DC. Prodr. 8: 387. 1844; Miers, Apoc. So. Am. 106. 1878. Thyrsanthus embelioides A. DC. loc. cit. 1844; Miers, loc. eit. 106. 1878. Forsteronia embelioides (A. DC.) Muell.-Arg. loc. cit, 97. pl.50. fig. 2. 1860. Forsteronia meridionalis Muell.-Arg. loc. cit. 1860. Forsteronia minutiflora Muell.-Arg. loc. cit. 99. 1860; Miers, loc. cit. 244. 1878. (193) [Vor. 22 160 ANNALS OF THE MISSOURI BOTANICAL GARDEN Thyrsanthus meridionalis (Muell.-Arg.) Miers, loc, cit. 106. 1878. Stems relatively slender, pilosulose when young, soon be- coming glabrate and conspicuously lenticellate; leaves oppo- site, or rarely ternate, petiolate, narrowly oblong to oblong- lanceolate, apex acuminate, base very obscurely cordate, 5-9 em. long, 1.2-3.0 em. broad, membranaceous, rather incon- spieuously glandular at the base of the midrib above, above glabrous or very rarely generally and minutely puberulent- papillate, beneath glabrous or minutely barbellate in the axils of the midrib, rarely generally and minutely puberulent-papil- late; petioles 0.5-1.0 cm. long; inflorescence thyrsiform, termi- nal, infrequently both terminal and lateral, the terminal greatly surpassing, and the lateral much shorter than, the sub- tending leaves, bearing many small, white flowers; primary peduncle glabrous or infrequently minutely and rather irregu- larly puberulent-papillate, ultimate branches regularly and de- terminately composed, minutely puberulent-papillate, rarely pilosulose; pedicels 0.1-0.15 em. long, minutely puberulent- papillate, rarely pilosulose; braets lanceolate to ovate-lance- olate, 0.05-0.3 em. long, scarious; calyx-lobes ovate, acute to acuminate, 0.08-0.15 em. long, scarious, minutely puberulent- papillate without, the squamellae minute, numerous, indefi- nitely and regularly distributed, rarely evidently lacking; corolla very minutely papillate or essentially glabrous with- out, the tube about 0.1 em. long, about 0.06—0.1 em. in diameter at the base, 0.1-0.15 cm. in diameter at the orifice, villosulose within at the insertion of the stamens, the lobes oblong, 0.18— 0.2 em. long, spreading, conspieuously pilosulose within; stamen filaments about 0.05 em. long, distinet and free, not agglutinated to the style, the anthers essentially included or the tips barely exserted, 0.07-0.09 em. long, glabrous or es- sentially so; ovary ovoid, apocarpous, about 0.035 cm. long, minutely puberulent-papillate; nectaries somewhat shorter than the ovary; stigma blunt, 0.06-0.07 cm. long; follicles unknown. BRAZIL: BAHIA: data incomplete, Blanchet 1745 (BB, DC, MBG, photograph and analytical drawings); RIO DE JANEIRO: data incomplete, Glaziou 4880, 15457 (194) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 161 (B); Glaziou 3056 (Bx); MINAS GERAES: Sao João Baptista, date lacking, Sellow s.n. (B, MBG, photograph and analytical drawings). The opposite or ternate phyllotaxy is apparently a valid specifie eriterion in no known species of Forsteronia, contrary to Mueller's assumption. On different specimens of Blan- chet 1745 may be found either type of phyllotaxy upon the same branch. Similar circumstances also are to be found in other species, as in F. refracta Muell.-Arg. 3. Forsteronia Luschnathi Muell.-Arg. in Mart. Fl. Bras. 61: 98. 1860. Forsteroma acutifolia Muell.-Arg. loc. cit. 99. 1860; Miers, A poc. So. Am. 246. 1878. Forsteronia acutifolia Muell.-Arg. B. pubescens Muell.- Arg. loc. cit. 1860. Thyrsanthus Luschnatii (Muell.-Arg.) Miers, loc. cit. 106. 1878, sphalm. Forsteronia protensa Miers, loc. cit. 246. 1878. Stems relatively slender, minutely puberulent to glabrate when young, eventually glabrate and conspicuously lenticellate ; leaves opposite, shortly petiolate, oblong-lanceolate to broadly elliptic, apex acuminate, base obtuse to rounded, 4-8 cm. long, 1.54.0 em, broad, firmly membranaceous, glandular at the base of the midrib above, glabrous or very minutely puberulent- papillate above and beneath, rarely minutely puberulent gen- erally; petioles 0.3-0.8 em. long; inflorescence thyrsiform, both terminal and lateral, the terminal somewhat surpassing, and the lateral much shorter than, the subtending leaves, bear- ing many small, white flowers; primary peduncle minutely puberulent, ultimate branches regularly and determinately composed, densely and minutely puberulent-papillate ; pedicels 0.075-0.2 em. long, densely puberulent-papillate; bracts lance- olate, 0.08-0.25 em, long, searious ; ealyx-lobes ovate-lanceolate, acute to acuminate, 0.1—0.15 em. long, scarious, densely puber- ulent-papillate without, the squamellae minute, numerous, in- definitely and regularly distributed ; corolla minutely papillate without, partieularly toward the tips of the lobes, the tube 0.08-0.1 em. long, about 0.075-0.09 em, in diameter at the base (195) [Vor. 22 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN and 0.1-0,125 em. in diameter at the orifice, minutely villosulose within at the insertion of the stamens, the lobes narrowly oblong, 0.13-0.15 em. long, spreading, conspicuously pilosulose within; stamen filaments 0.03-0.05 em. long, distinet and free, not agglutinated to the style, the anthers essentially included or the tips barely exserted, 0.075-0.1 em. long, glabrous; ovary ovoid, about 0.025 em. long, minutely hirtellous, apocarpous; nectaries somewhat shorter than the ovary; stigma 0.06-0.07 em. long; follicles unknown. BRAZIL: RIO DE JANEIRO: ‘‘ Thelegraffenberg [Sebastiana],’’ Jan., 1833, Lusch- nath s.n. (Bx, TYPE, MBG, photograph and analytical drawings); ‘‘ Nouvelle Fribourg [Nova Friburgo],’’ Oct., 1842, Claussen 84 (D); circa Rio de Janeiro, date lacking, Schott 5976 (V) ; DATA INCOMPLETE, Gaudichaud 976 (B, D) ; Widgren s.n. (S); Glaziou 6818 (B). The type specimen of F. Luschnathi possesses ternate phyllotaxy. All others cited, including the types of F. acuti- folia and F. protensa (= F. acutifolia B pubescens) are char- acterized by normal, opposite foliage. As has been remarked previously, an occasional ternate specimen has been found in several species of Forsteronia, although in all other respects typieal. Abnormalities in this respect have also been found upon the same specimen with normal, opposite foliage. The occurrence of ternate phyllotaxy has therefore been inter- preted as an insignificant anomaly. 4. Forsteronia elachista Blake, Contr. U. S. Nat. Herb. 20: 029. pl.41. 1924. Stems relatively slender, glabrous, conspicuously lenticellate when fully mature; leaves opposite, petiolate, obovate-oblong, apex shortly and rather abruptly acuminate, base obtuse to rounded, 6-10 em. long, 2-5 em. broad, membranaceous, glan- dular at the base of the midrib above, glabrous above, beneath barbellate in the axils of the midrib to glabrate; petioles 0.7-0.8 em. long; inflorescence laxly thyrsiform, terminal or both terminal and lateral, the terminal greatly surpassing the subtending leaves, bearing many small, white flowers ; primary pedunele minutely puberulent to glabrate, the ultimate branches regularly and determinately composed, densely pu- (196) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 163 berulent-papillate; pedicels 0.1-0.25 em. long, minutely puber- ulent-papillate; bracts ovate or ovate-lanceolate, 0.05—0.1 cm. long, scarious; calyx-lobes ovate-deltoid, obtuse, 0.075-0.1 em. long, searious, densely puberulent-papillate without, the squamellae minute, numerous, regularly and indefinitely dis- tributed; eorolla densely puberulent-papillate without, the tube 0.08-0.1 em. long, about 0.05-0.06 cm. in diameter at the base and 0.1-0.125 em in diameter at the orifice, villosulose within at the insertion of the stamens, the lobes ovate-oblong, 0.11-0.15 em. long, spreading, conspicuously villosulose within; stamen filaments about 0.02 em. long, distinct and free, not ag- glutinated to the style, the anthers essentially included or the tips barely exserted, 0.07-0.08 cm. long, conspicuously bar- bellate; ovary ovoid, apocarpous, about 0.02 em. long, minutely hirtellous ; nectaries somewhat shorter than the ovary ; stigma 0.075 em. long; follicles unknown. COLOMBIA: BOYACA: exact locality and date lacking, Lawrance 453 (FM). VENEZUELA: CARABOBO: Upper Guaremales, road from Puerto Cabello to San Felipe, in forest and clearings, alt. 100-500 m., July 2, 1920, Pittier 8918 (G, US, TYPE, MBG, photograph and analytical drawings); DATA INCOMPLETE: Linden 267 5. Forsteronia graciloides Woodson, spec. nov. Suffruticosa volubilis; ramulis gracilibus glabris maturitate conspicue lenticellatis; foliis oppositis petiolatis obovatis vel obovato-oblongis apice breviter abrupteque subcaudato-acu- minatis basi obtusis 5-8 cm. longis 2-4 cm. latis firme mem- branaceis supra glabris nervo medio basi pauciglanduligero subtus saepe irregulariter inconspicueque puberulis saepius glabratis; petiolis 0.4-0.8 cm. longis; inflorescentiis laxe thyrsiformibus et terminalibus et lateralibus folia saepius multo superantibus flores multas albidas gerentibus; pe- duneulis omnino minute puberulo-papillatis vel basi glabratis ultimis regulariter cymoso-compositis; pedicellis 0.1—0.2 cm. longis minute puberulo-papillatis; bracteis ovatis vel ovato- laneeolatis 0.05-0.15 em. longis scariaceis; calycis laciniis laneeolatis vel ovato-lanceolatis apice acutis acuminatisve 0.075—0.1 em. longis scariaceis minute puberulo-papillatis intus eum squamellis minutissimis paucis alternatis; corollae extus [Vor. 22 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN dense puberulo-papillatae tubo 0.075-0.1 em. longo basi ca. 0.05-0.075 em. diametro faucibus ca. 0.1—0.125 em. diametro metientibus intus prope insertionem staminum villosulis lobis late oblongis obtusis 0.1-0.11 cm. longis patulis intus conspicue pilosulis ; filamentis staminum 0.01-0.02 em, longis liberis haud agglutinatis antheris paululo exsertis 0.07—0.08 em. longis conspicue barbellatis; ovario ovoideo apoearpo ca. 0.02 em. longo minute papillato; nectariis ovario paululo brevioribus ; stigmate 0.06-0.08 em, longo; follieulis erassiuseulis obscure articulatis falcatis 43-47 em. longis glabris; seminibus 2.0-2.5 em. longis como pallide aurantiaco ca. 3 em. longo. Stems relatively slender, glabrous, conspicuously lenticellate when fully mature; leaves opposite, petiolate, obovate to obovate-oblong, apex shortly and abruptly subcaudate-acu- minate, base obtuse, 5-8 em. long, 2-4 em. broad, firmly mem- branaceous, above glabrous, glandular at the base of the midrib, beneath irregularly and inconspicuously puberulent, partieularly in the axils of the midrib, frequently glabrate; petioles 0.4—0.8 cm. long; inflorescence laxly thyrsiform, both terminal and lateral, the terminal conspicuously sur- passing the subtending leaves, bearing many small, white flowers; primary peduncle minutely puberulent-papillate to glabrate at the base, the ultimate branches regularly and de- terminately composed, densely puberulent-papillate; pedicels 0.1-0.2 em. long, densely puberulent-papillate; bracts ovate to ovate-lanceolate, 0.05-0.15 em. long, searious; calyx-lobes lan- ceolate to ovate-lanceolate, acute to acuminate at the apex, 0.075-0.1 em. long, scarious, densely and minutely puberulent- papillate, squamellae very minute, relatively few, in alternate groups; eorolla densely puberulent-papillate without, the tube 0.075—0.1 em. long, about 0.05—0.075 em. in diameter at the base and 0.1-0.125 cm. in diameter at the orifice, villosulose within at the insertion of the stamens, the lobes broadly oblong, obtuse, 0.1-0.11 cm. long, spreading, conspicuously pilosulose within; stamen filaments 0.01-0.02 em. long, distinet and sepa- rate, not agglutinated to the style, the anthers essentially included or the tips barely exserted, 0.07-0.08 em. long, con- (198) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 165 spicuously barbellate; ovary ovoid, apocarpous, about 0.02 em. long, minutely papillate; nectaries somewhat shorter than the ovary; stigma 0.06-0.08 em, long; follieles rather stout, ob- seurely and rather irregularly articulated, falcate, 43-47 cm. long, glabrous; seeds 2.0-2.5 em. long, the pale tawny coma about 3 em. long. PERU: LORETO: forest, San Ramon, near Yurimaguas, Nov. 4, 1929, Williams 4547 (FM, MBG, TYPE); Yurimaguas, Oct. 24, 1929, Williams 4011 (FM, MBG); prope Tarapoto, 1855-56, Spruce 4493 (B, BB, C, D, G, V); Flutfreier Hochwald, Mündung d. Santiago, alt. 160 m., Oct. 28, 1924, Tessmann 4413 (B, D); SAN MARTIN: Pongo de Cainarachi, Rio Cainarachi, tributary of Rio Huallaga, alt. 230 m., Sept.-Oct., 1932, Klug 2761 (NY); JUNIN: La Merced im Chanchamayo- Thal, liehter Wald, alt. 1000 m., Dec., 1902, Weberbauer 1888 (B). This species differs from the complementary species of the north Atlantie coastal region of South America, F. elachista Blake, not only in the key characters, but also in such char- acters as the corolla-lobes, which are proportionally longer and narrower, and the ovary, which is minutely papillate, whilst conspicuously hirtellous in the latter species. 6. Forsteronia Gardneri (A. DC.) Muell.-Arg. in Mart. Fl. Bras. 6': 100. 1860. Thyrsanthus Gardneri A. DC. in DC. Prodr, 8: 387. 1844; Miers, Apoc. So. Am. 96. 1878. Stems relatively slender, minutely and irregularly pilosulose when young, soon becoming glabrate and conspicuously lenti- cellate; leaves opposite, shortly petiolate, narrowly oblong- elliptic, apex acuminate, base rounded, 7-14 em. long, 1.5-4.0 em, broad, firmly membranaceous to subcoriaceous, glabrous, ineonspieuously glandular at the base of the midrib above; petioles 0.3-0.6 em. long, minutely puberulent-pilosulose; in- florescence laxly thyrsiform, terminal or both terminal and lateral, in either case conspicuously surpassing the subtending leaves, bearing many small, white flowers; primary peduncle pilosulose, ultimate branches regularly and determinately com- posed, puberulent-papillate to pilosulose; pedicels 0.05-0.1 em. long, puberulent-papillate; bracts ovate-lanceolate, 0.05—0.2 em. long, scarious; calyx-lobes ovate, acute, 0.1-0.11 em. long, (199) [Vor. 22 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN scarious, puberulent-papillate without, the squamellae in groups of 2-4 opposite the calyx-lobes, or irregularly and indefinitely distributed; corolla sparsely and minutely pilosulose-papillate without, particularly toward the tips of the lobes, otherwise usually essentially glabrate, the tube 0.11-0.15 em. long, about 0.08 em. in diameter at the base and 0.15-0,175 em. in diameter at the orifice, villosulose within at the insertion of the stamens, the lobes rather narrowly oblong, 0.15-0.2 em. long, spreading, conspicuously pilosulose within; stamen filaments 0.03-0.05 em. long, distinct and free, not ag- glutinated to the style, the anthers essentially included or only the tips barely exserted, 0.1-0.12 em. long, glabrous; ovary ovoid, apocarpous, about 0.03 em. long, minutely papillate ; nectaries about half as long as the ovary ; stigma 0.1-0.12 em. long; follicles unknown. BRAZIL: GOYAZ: exact locality lacking, 1842, Gardner 3891 (B, BB, BM, Camb., D, DC, NY, V, TYPE, MBG, photograph and analytical drawings). 7. Forsteronia affinis Muell.-Arg. in Mart. Fl. Bras. 6': 100. pl. 30. 1860. Thyrsanthus affinis (Muell.-Arg.) Miers, Apoc. So. Am. 101. 1878. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, petiolate, ovate-elliptic, apex acuminate, base broadly obtuse or rounded, 8-12 em. long, 3.0-4.5 em. broad, rather delicately membranaceous, glandular at the base of the midrib above, glabrous, or very incon- spicuously and minutely barbellate in the axils of the midrib beneath; petioles 0.8-0.9 em. long; inflorescence thyrsiform, terminal, or both terminal and lateral, the terminal conspicu- ously surpassing, and the lateral somewhat shorter than, the subtending leaves, bearing many small, white flowers; primary pedunele glabrous or very minutely puberulent-papillate toward the tip, ultimate branches indefinitely congested, mi- nutely puberulent-papillate; pedicels 0.08-0.1 em. long, mi- nutely puberulent-papillate; braets lanceolate, 0.02-0.05 cm. long, scarious; calyx-lobes ovate-trigonal, acute, 0.07+0.1 em. long, scarious, minutely papillate or very minutely puberulent- (200) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 167 papillate, the squamellae evidently lacking; corolla densely papillate without, the tube 0.075—0.1 em. long, about 0.05-0.075 em. in diameter at the base and 0.1-0.125 em. in diameter at the orifice, minutely villosulose within at the insertion of the stamens, the lobes oblong-elliptie, 0.12-0.15 em. long, spread- ing, papillate within; stamen filaments 0.02-0.03 em. long, distinct and free, not agglutinated to the style, the anthers essentially included or the tips barely exserted, 0.06-0.08 cm. long, glabrous; ovary ovoid, apocarpous, about 0.03 cm. long, minutely puberulent; nectaries somewhat shorter than the ovary ; stigma 0.07-0.08 em. long; follicles unknown. BRAZIL: AMAZONAS: in sylvis ad Ega, Rio Negro, date lacking, Martius 2960 (M, TYPE, MBG, photograph and analytical drawings). 8. Forsteronia rufa Muell.-Arg. in Mart. Fl. Bras. 6': 100. pl. 31. fig. 1. 1860; K. Sch. in Engl. € Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Thyrsanthus rufus (Muell.-Arg.) Miers, Apoc. So. Am. 104. 1878. Forsteronia rufa Muell.-Arg. var. subglabra Malme, Arkiv f. Bot. 22A?: 14. 1928. Stems relatively stout, densely ferruginous-tomentulose when young, eventually becoming glabrate and conspicuously lenticellate when fully mature; leaves opposite, infrequently ternate, shortly petiolate, ovate-, oblong-, or obovate-elliptic, apex shortly and rather abruptly acuminate, base usually obscurely cordate, rarely broadly euneate, 5-11 cm. long, 2.5—6.0 em. broad, firmly membranaceous, above more or less densely ferruginous-hirtellous, particularly along the veins, fre- quently glabrate or essentially glabrous, inconspicuously glandular at the base of the midrib, beneath more or less densely ferruginous-tomentulose or -puberulent, particularly along the veins, rarely nearly glabrate; petioles 0.4-0.8 em. long, the indument as upon the stem; inflorescence laxly thyrsiform, terminal, frequently lateral as well, greatly sur- passing the subtending leaves, bearing many small, white flowers; primary peduncle ferruginous-tomentulose, ultimate branches minutely ferruginous-tomentulose, regularly and de- (201) [Vor. 22 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN terminately composed; pedicels 0.08-1.5 cm. long, minutely ferruginous-tomentulose; bracts lanceolate to ovate-lanceo- late, 0.015-0.05 em. long, scarious; calyx-lobes lanceolate to ovate-lanceolate, acuminate, minutely puberulent-papillate to ferruginous-hirtellous without, 0.13-0.2 em. long, the squamel- lae in opposite groups of 2-5; corolla glabrous without, infre- quently somewhat puberulent-papillate to minutely barbellate at the tips of the lobes, the tube 0.13-0.2 em. long, about 0.05- 0.75 em. in diameter at the base and 0.15-0.175 em. in diameter at the orifice, villosulose within, the lobes narrowly oblong, 0.2- 0.4 em. long, widely spreading, conspicuously pilosulose within; stamen filaments 0.06—0.1 em. long, distinet and free, not ag- glutinated to the style, the anthers essentially included or the tips barely exserted, 0.1-0.15 em. long, glabrous; ovary ovoid, apocarpous, about 0.03 em. long, minutely ferruginous- hirtellous; nectaries slightly shorter than the ovary; stigma 0.1-0.18 em. long; follicles relatively stout, very conspicuously and distantly moniliform, tortuous, 37-50 em. long, glabrous; seeds 0.8-1.0 em. long, the pale tawny coma 3.54.5 cm. long. BRAZIL: MINAS GERAES: exact locality lacking, 1834, Ackermann s.n. (B, Bx); -y DE JANEIRO: Organ Mts., date lacking, Gardner 530 (Camb., D, TYPE, NY, US, V, MBG, photograph); Vargem, Organ Mts., Jan., 1838, Miers 4049 (US); data ineompMt e: Glaziou 6905 (B, Bx); Glaziou 5939 (B, FM, US); Glaziou 8799 (B); Glaziou 14065 (B); SAO PAULO: sepibus sylvestribus in Serra do Mar et versus fl. mn date lacking, Martius s.n. (M); sylvis Capoes udis, Jan., year lacking, Martius s.n. (M); Butantan, Feb. 22, 1918, Hoehne 1512 (B); Iguape, April 25, 1918, Hoehne 1841 (B); PARANA: Morretes, ad marginem silvae primaevae, alt. ea, 40 m., Jan. 4, 1914, Dusen pes "s (D, US); Jacarehy, in silva primaeva, Febr. 11, 1915, posa 16662 (D, US); SANTA CATHARINA: am Waldrand, Ins. Sáo Fran- cisco, Febr., 1885, Ule 350 (B); Waldrand bei Itajahy, Jan., 1886, Ule 510 (B); am Capivary bei Tubaráo, Jan., 1889, Ule 1051 (B, US); DATA INCOMPLETE: Sellow 211 (B); Sellow 358 (B); Sellow 219 (B); Sellow 543 (B); Sellow s.n. (B); Riedel s.n. (B, BB, V). This is probably the most variable species of the genus. The dimensions of the floral parts are particularly puzzling in this regard, as well as the superficial aspect of the inflorescence as a whole. Especially noteworthy are the two specimens of Ule 1051 from the province of Santa Catharina, which vary from the norm of the species in the cuneate bases of the leaves and a somewhat ashy tinge of the ferruginous indument. (202) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE, IV 169 9. Forsteronia Acouci (Aubl.) A. DC. in DC. Prodr. 8: 439. 1844. Apocynum Acouci Aubl. Hist. Pl. Gui. Fr. 1: 274; 3: pl. 107. 1775. Apocynum paniculatum Lam. Encycl. 1: 214. 1783. Thyrsanthus Schomburgkii Benth. in Hook. Jour. Bot. 3: 249. 1841; A. DC. loc. cit. 387. 1844; Miers, Apoc. So. Am. 94. 1878. Forsteronia Schomburgkii (Benth.) Muell.-Arg. in Mart. Fl. Bras. 6': 107. 1860, not A. DC. Thyrsanthus Acouci ( Aubl.) Miers, loc. cit. 98. 1878. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong- to ovate- or obovate-elliptie, apex shortly and abruptly acu- minate, base broadly obtuse or rounded, 8-15 em. long, 3-7 em. broad, membranaceous, glabrous, inconspicuously glandular at the base of the midrib above; petioles 0.3-0.7 em, long; inflores- cence thyrsiform, both terminal and lateral, usually somewhat shorter than the subtending leaves, bearing many small, white flowers; primary peduncle minutely puberulent-papillate to glabrate, ultimate branches congested, but regularly and de- terminately composed, densely and minutely puberulent-papil- late; pedicels 0.05-0.1 em. long, very minutely puberulent- papillate; bracts ovate to ovate-lanceolate, 0.07-0.15 em. long, scarious ; calyx-lobes ovate to ovate-lanceolate, acute to broadly acuminate, 0.1-0.17 em. long, scarious, densely papillate, the squamellae solitary or rarely paired, alternate; corolla mi- nutely puberulent-papillate without, particularly toward the tips of the lobes, the tube 0.1-0.2 em. long, about 0.05-0.1 em. in diameter at the base and 0.175—0.2 em, in diameter at the orifice, villosulose within at the insertion of the stamens, the lobes oblong or ovate-oblong, 0.09-0.13 em. long, only slightly spreading, minutely papillate within ; stamen filaments 0.06—0.1 cm. long, distinct and free, not agglutinated to the style, the anthers essentially included or the tips barely exserted, 0.15- 0.2 em, long, barbellate, or very minutely papillate at the tips; ovary ovoid, apocarpous, about 0.05 em. long, minutely puber- (203) [Vor. 22 170 ANNALS OF THE MISSOURI BOTANICAL GARDEN ulent; nectaries about equalling the ovary ; stigma 0.13-0.2 em. long; follicles unknown. TRINIDAD: wet forest border, near Caroni River, south of Arima, April 11, 1921, Britton $ Britton 2900 (NY BRITISH GUIANA: data incomplete, Schomburgk 37 (D, V); Schomburgk 557 (B, BB, Camb., D, NY, US, V); Schomburgk 782 (B, BB, Camb., D, V); rip. fl. Barima, Oct., 1843, Schomburgk 1514 had DUTCH GUIANA: fl, D July, 1 Apri 90 (U); near the 2n or Oude Ryweg, Paramaribo, May 23, 6, d (B, G; ); Paramaribo, 1850, Wullschláge 454 (V); fluv. E Dee., 1903, éd td 415 (U); Jagtlust-Meerzog, fluv. Suriname, Jan. 8, 1924, Collector Indigenous 8F (B, U); DATA INCOMPLETE: Hostmann $ Kappler 569 (U); Hostmann ^s "Y D, FM, 8); Hostmann 6118 (V); Hostmann $ Kappler 1236 (B, BB, D, Y) FRENCH GUIANA: data incomplete, leBlond "n 379 (B, €). Aublet's illustration of 4pocynum Acouci leaves little doubt concerning its association with TAyrsanthus Schomburgku Benth., a common plant of the Guianas. The association of Apocynum paniculatum Lam. with these species is less secure, and is based only in part upon Lamarck’s ineonclusive diag- nosis, consideration being given to the previous disposition by A. de Candolle and Miers. 10. Forsteronia Benthamiana Muell.-Arg. in Mart. Fl. Bras. 6': 106. 1860; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. — Benthamiana (Muell.-Arg.) Miers, Apoc. So. Am. 95. 8, Stems ed stout, glabrous, conspicuously lenticellate when fully mature; liavos opposite, shortly petiolate, oblong- elliptic to broadly oval, apex shortly and abruptly acuminate to obtuse or rounded, base broadly obtuse or rounded, 8-18 em. long, 2.5-8.0 em. broad, coriaceous or subcoriaceous, somewhat nitidulous above, inconspicuously glandular at the base of the midrib, glabrous throughout; petioles 0.3-0.7 em. long; inflo- rescence thyrsiform, both terminal and lateral, about equalling, or the terminal greatly surpassing, the subtending leaves, bearing many small, white flowers; primary peduncle minutely puberulent-papillate to glabrate below, ultimate branches regularly and determinately composed, densely puberulent- papillate; pedicels 0.1-0.3 em. long, densely and minutely pu- (204) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 171 berulent-papillate; bracts ovate to ovate-lanceolate, scarious, .05-0.35 em. long; calyx-lobes ovate to ovate-lanceolate, acute to obtuse, 0.1-0.2 cm. long, scarious, minutely puberulent or puberulent-papillate without, squamellae solitary, alternate; eorolla minutely and irregularly papillate to essentially gla- brous without, the tube 0.15-0.3 em. long, about 0.1-0.15 em. in diameter at the base and 0.175—0.2 em. in diameter at the orifice, villosulose within at the insertion of the stamens, the inben oblong to ovate-oblong, 0.2-0.3 em, long, spread: very mi- nutely and irregularly papillate to essentially glabrate within; stamen filaments 0.06-0.13 cm. long, distinct and free, not ag- glutinated to the style, the anthers slightly exserted, 0.15—0.2 em. long, minutely barbellate at the tips; ovary ovoid, apo- carpous, minutely puberulent-papillate, about 0.05 em. long; nectaries much shorter than the ovary; stigma 0.1-0.18 em. long; follicles relatively stout, rather inconspicuously undu- lated, subparallel to tortuous, frequently united at the tips, 21-23 em. long, glabrous; seeds 2.0-2.3 em. long, the bright tawny coma 1.3-2.0 em. long. BRAZIL: PARA: Juruty Velho, silva locis altis, July 29, dL Ducke 21603 (B); Oriximina, Rio Trombetas infer., silva secundaria, Sept. 17, 1910, Ducke 21639 (B, US); AMAZONAS: prope San Carlos, ad Rio Negro, 1853- 4, Spruce 3481 (B, BB C, D, G, M, NY, MBG, photograph and analytical drawings); Bóa Vista, Rio Branco super., ad ripas fluminis, July, 1913, Kuhlmann 3647 (B, U, US); Manáos, ripis paludosis rivuli ad, Dec. 8, 1927, Ducke 21612 (B) ; ad ripis inter Campinho et Boa Vista, Rio Branco, Nov., 1913, Kuhlmann 3648 (B); Teffe, ad fauces lacus, June 23, 1906, Ducke 21766 (B, US); Manáos, ad ripis Rio Negro prope faucem um. Taruma, April 26, 1911, Ducke 21758 (B, US); DATA INCOMPLETE: Glaziou 9932 (C). PERU: LORETO: forest, T near Iquitos, alt. 100 m., Febr.-Mareh, 1930 Klug 1053 (FM, NY, US); omgebiet des Maranon am ae de Migbefiche, 1924, Tessmann 4805 (S); Di Urwald, June 30, 1924, Tessmann 3639 (B, D); Stromgebiet des fd Santiago-Mündung am Pon e Manseriche, 1924, T ROMA 4461 (D, S); Stromgebiet des Maranon von iios. 1924, man 5128 (B, D 11. Forsteronia viridescens Blake, Contr. Gray Herb. 52: 80, 1917. Stems relatively stout, glabrous or essentially so, conspicu- ously lenticellate when fally mature; leaves opposite, shortly petiolate, oblong-elliptic, apex very shórily and abruptly acu- (205) [Vor. 22 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN minate, base broadly obtuse or rounded, 9-13 em. long, 2.5-6.0 em, broad, coriaceous or subeoriaceous, somewhat nitidulous above, glandular at the base of the midrib above, glabrous throughout; petioles 0.3-0.6 em. long; inflorescence thyrsiform, terminal, rarely lateral as well, somewhat shorter than the sub- tending leaves, bearing many small, greenish-white flowers; primary pedunele minutely puberulent-papillate to glabrate below, ultimate branches regularly and determinately com- posed, minutely puberulent-papillate; pedicels 0.09-0.15 em. long, very minutely puberulent-papillate; bracts ovate to ovate-lanceolate, searious, 0.09-0.18 em. long; calyx-lobes ovate, acute, 0.1—0.13 em. long, scarious, very minutely puber- ulent-papillate, the squamellae minute, solitary, alternate, occasionally lacking; corolla densely and uniformly puberulent- papillate without, the tube 0.15-0.17 em. long, about 0.075 em. in diameter at the base and 0.175-0.2 em. in diameter at the orifice, villosulose within, the lobes broadly oblong to ovate- oblong, 0.18-0.2 em. long, spreading, minutely puberulent within; stamen filaments 0.05-0.08 em. long, distinct and free, not agglutinated to the style, the anthers slightly exserted, 0.15-0.17 em. long, minutely and sparsely barbellate at the tips; ovary ovoid, apocarpous, about 0.05 em. long, papillate; nec- taries much shorter than the ovary ; stigma 0.12-0.13 em. long; follicles unknown. British Honpuras: forest near Manatee Lagoon, July 7, 1906, Peck 450 (G, TYPE, MBG, photograph and analytical drawings); secondary forest, rare, Middle- sex, alt. 200 ft., Sept. 20, 1929, Schipp 360 (B, D, FM, MBG, NY). 12. Forsteronia guyanensis Muell.-Arg. Linnaea 30: 414. 1860. Thyrsanthus Guyanensis (Muell.-Arg.) Miers, Apoc. So. Am. 97. 1878. Stems relatively stout, minutely puberulent when very young, soon becoming glabrate and conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, elliptic- obovate, apex very abruptly and shortly acuminate to obtuse, base obtusely euneate, 3-9 em. long, 1.5-3.5 em. broad, cori- aceous or subcoriaceous, rather inconspicuously glandular at the base of the midrib above, glabrous throughout; petioles (206) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE, IV 173 0.2-0.5 em. long; inflorescence thyrsiform, both terminal and lateral, equalling to conspicuously surpassing the subtending leaves, bearing many small, white flowers; primary peduncle densely papillate to glabrate below, ultimate branches regu- larly and determinately composed, densely papillate; pedicels 0.09-0.2 cm. long, densely papillate; bracts ovate to ovate- lanceolate, scarious, 0.08—0.2 em. long; calyx-lobes ovate, acute to acuminate, 0.1-0.14 em. long, searious, densely and very minutely puberulent-papillate without, the squamellae usually lacking, occasionally very minute, solitary, and alternate; corolla minutely puberulent-papillate without, the tube 0.15- 0.2 em. long, about 0.075—0.1 em. in diameter at the base and 0.17-0.2 cm. in diameter at the orifice, minutely puberulent- papillate within near the orifice, the lobes broadly oblong to ovate-oblong, 0.15-0.22 em. long, spreading, minutely and densely papillate or puberulent-papillate within; stamen fila- ments 0.07—0.1 cm. long, distinct and free, not agglutinated to the style, the anthers slightly exserted, 0.15-0.18 em. long, glabrous; ovary ovoid, minutely puberulent, apocarpous; nec- taries about equalling the ovary; stigma 0.12-0.15 em. long; follicles very slender and tortuous, obscurely and indefinitely artieulated or essentially entire, 45-50 cm. long, very densely and minutely ferruginous-papillate; seeds 1.5 cm. long, the bright tawny coma 2.8-3.2 em. long. BRITISH GUIANA: rip. fl. Sururu, Sept., 1843, Schomburgk 1446 (B, ISOTYPE, MBG, photograph and analytical drawings); exact locality lacking, 1844, Schomburgk 821 (BB, Camb., D); mixed forest, Essequibo River, Moraballi Creek, near Bartica, alt. near sea-level, Oct. 9, 1929, Sandwith 405 (NY, S, U, US); ** Demarara,"? date lacking, Jenman 5091 (K, NY DUTCH GUIANA: in summus monte Brownsberg, July 28, 1924, Stahel $ Gonggrijp 6624 (U); in silva pr. Voltzberg, fluv. Coppename, July-Sept., 1920, Pulle 291 (B, U); in silva pr. Avanaverovallen, fluv. Kabalebo, July-Sept., 1920, Pulle 453 (B, NY, U); fluv. Coppename inf., Aug., 1901, Went 109 (U FRENCH GUIANA: Karouany, 1858, Sagot 1067 (BB, Bx, U); data incomplete: poss [?] 381 (B); Martin s.n. (B); Poiteau s.n. (D); Melonon 460 (B). ZIL: PARA: Peixeboi ad viam ferream Belem-Braganca, silva, Oct. 30, 1907, Poen 21623 (B, U, US). 13. Forsteronia brevifolia Mgf. Notizblatt 10: 1038. 1930. Stems relatively stout, minutely papillate when very young, soon becoming glabrate and conspicuously lenticellate when (207) [Vor. 22 174 ANNALS OF THE MISSOURI BOTANICAL GARDEN fully mature; leaves opposite, very shortly petiolate, broadly elliptie to obovate-elliptie, apex very shortly and abruptly acuminate to obtuse, base broadly obtuse or rounded, 4—7 cm. long, 2.0-3.5 em. broad, coriaceous, somewhat nitidulous above and inconspicuously glandular at the base of the midrib, gla- brous throughout; petioles 0.3-0.5 em. long; inflorescence corymbose-thyrsiform, flat-topped or only somewhat convex, both terminal and lateral, about equalling or somewhat shorter than the subtending leaves, bearing many small, white flowers; primary peduncle very densely and minutely puberulent-papil- late, ultimate branches regularly and determinately composed, densely puberulent-papillate ; pedicels 0.1-0.2 em. long, densely and minutely puberulent-papillate; braets ovate-lanceolate, 0.03-0.075 cm. long, scarious; calyx-lobes ovate to ovate-del- toid, obtuse or rounded, 0.1-0.125 em. long, scarious, minutely puberulent-papillate without, the squamellae minute, solitary or rarely paired, alternate, occasionally lacking; corolla rather irregularly puberulent-papillate without, the tube 0.13-0.15 em. long, about 0.075-0.1 em. in diameter at the base and 0.15- 0.2 em. in diameter at the orifice, villosulose within, the lobes oblong to oblong-lanceolate, 0.25-0.275 em. long, only slightly spreading, minutely puberulent to essentially glabrous within; stamen filaments 0.03-0.05 em. long, distinct and free, not ag- glutinated to the style, the anthers slightly exserted, 0.15— 0.175 em. long, glabrous; ovary ovoid, apocarpous, minutely papillate; nectaries somewhat shorter than the ovary ; stigma 0.2 em. long; follieles unknown. BRAZIL: AMAZONAS: Tocantins, silva non inundabili, Nov. 13, 1927, Ducke 21614 (B, TYPE, US, MBG, photograph and analytical drawings). PERU: LORETO: forest, Mishuyacu, near Iquitos, alt. 100 m., Dec., 1929, Klug 640 (FM, NY, US). 14. Forsteronia Wilsonii (Griseb.) Woodson, comb. nov. Thyrsanthus Wilsonii Griseb. Fl, Brit. W. I. 412. 1864. Stems relatively slender, glabrous, rather inconspicuously lenticellate when fully mature; leaves opposite, shortly petio- late, elliptie, apex rather abruptly and acutely subcaudate- acuminate, base broadly obtuse, 4-11 cm. long, 1.5-4.0 em. broad, membranaceous, glabrous, inconspieuously glandular at (208) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 175 the base of the midrib above; petioles 0.2-0.5 em. long; inflo- rescence thyrsiform, terminal, somewhat shorter than the sub- tending leaves, bearing many small, white flowers; primary peduncle minutely puberulent, ultimate branches somewhat congested, minutely puberulent to puberulent-papillate; pedi- cels about 0.1 em. long, puberulent-papillate; bracts linear to linear-lanceolate, 0.1-0.4 em. long, slightly foliaceous; calyx- lobes linear to linear-lanceolate, acuminate, 0.38-0.4 em. long, slightly foliaceous, rather irregularly and sparsely puberulent- papillate, the squamellae very minute, numerous, regularly and indefinitely distributed; corolla irregularly and sparsely puberulent-papillate without, the tube 0.12—0.15 em. long, about 0.07—0.08 em. in diameter at the base and 0.15-0.175 em. in diam- eter at the orifice, villosulose within, the lobes ovate-oblong, 0.2-0.25 cm. long, puberulent-papillate within, widely spread- ing; stamen filaments 0.05 em. long, distinet and free, not ag- glutinated to the style, the anthers slightly exserted, 0.15- 0.175 em. long, glabrous; ovary ovoid, apocarpous, about 0.05 em. long, minutely puberulent-papillate; stigma 0.12- 0.15 em. long; nectaries much shorter than the ovary ; follicles unknown. JAMAICA: on trees, Manchester, date lacking, Wilson s.n. (K, TYPE, NY, ME TYPE and analytical drawings); vicinity of Mandeville, April 26—30, 1910, S Ud 714 (NY, MBG, photograph and analytieal drawings); data incomplete, Bertero s.n. (B). This combination has been aseribed to Bentham € Hooker upon herbarium labels, although there appears to be no actual basis for such citation. 15. Forsteronia diospyrifolia Muell.-Arg. Linnaea 30: 415. 1860. Thyrsanthus diospyrifolius (Muell.-Arg.) Miers, Apoc. So. Am. 96. 1878. Stems relatively stout, glabrous or minutely puberulent- papilate when very young, conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, obovate- to elliptic-oblong, apex abruptly acute to obtuse, base broadly obtuse or rounded, 7-12 em. long, 2.5-5.0 em. broad, heavily (209) [Vor. 22 176 ANNALS OF THE MISSOURI BOTANICAL GARDEN coriaceous, glabrous, eglandular; petioles 0.4—0.6 em. long; in- florescence thyrsiform, both terminal and lateral, about equal- ling or somewhat shorter than the subtending leaves, bearing many small, white flowers; primary peduncle minutely and rather sparsely puberulent to glabrate, ultimate branches con- gested, puberulent to puberulent-papillate; pedicels 0.03- 0.05 em. long, puberulent-papillate; braets ovate, 0.05—0.35 em. long, scarious; calyx-lobes ovate, acute to obtuse, 0.175- 0.3 em. long, scarious, rather irregularly puberulent-papillate without, the squamellae solitary, alternate, minute; corolla puberulent-papillate without, the tube 0.125-0.15 em. long, about 0.1 em. in diameter at the base and 0.2 em. in diameter at the orifice, villosulose within, the lobes oblong-elliptie, 0.175— 0.2 em. long, somewhat spreading, minutely puberulent-papil- late within; stamen filaments 0.08-0.1 em. long, distinet and free, not agglutinated to the style, the anthers slightly exserted, 0.175-0.2 em. long, glabrous or very minutely barbellate; ovary apocarpous, ovoid, about 0.075 em. long, minutely papillate; stigma 0.15 em. long; nectaries somewhat shorter than the ovary ; follicles unknown. BRITISH GUIANA: in savannis, Oct., 1846, Schomburgk 854 (B, TYPE, MBG, photograph and analytical drawings); Roraima, 1842-3, Schomburgk 725 (D). 16. Forsteronia Riedelii Muell.-Arg. in Mart. Fl. Bras. 6': 103. 1860; Miers, Apoc. So. Am. 245. 1878. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong- elliptic, apex abruptly and obtusely subcaudate-acuminate, base broadly obtuse, 5-9 em. long, 2.5-3.5 em. broad, firmly membranaceous, glabrous, eglandular; petioles 0.5-0.8 cm. long; inflorescence thyrsiform, both terminal and lateral, somewhat shorter than the subtending leaves, bearing many small, white flowers; primary peduncle minutely puberulent- papillate, ultimate branches regularly and determinately com- posed, minutely puberulent-papillate; pedicels 0.2-0.3 em. long, minutely puberulent-papillate; braets ovate to ovate- lanceolate, 0.08-0.15 em. long, scarious; calyx-lobes ovate, acute, 0.1-0.12 cm. long, scarious, minutely puberulent-papil- (210) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV Lit late, the squamellae solitary, alternate, minute; corolla mi- nutely puberulent-papillate without, the tube 0.125-0.2 cm. long, about 0.075-0.1 em. in diameter at the base and 0.2-0.3 em. in diameter at the orifice, villosulose within, the lobes oblong- ovate, slightly reflexed, 0.2-0.25 em. long, puberulent within; stamen filaments 0.05 em. long, distinet and free, not agglutin- ated to the style, the anthers slightly exserted, about 0.2 cm. long, glabrous; ovary ovoid, apocarpous, about 0.075 em. long, hirtellous; stigma 0.15-0.2 em. long; nectaries somewhat shorter than the ovary; follicles unknown. - BRAZIL: AMAZONAS: silva non inundabili circa cataractas flum. Taruma su- perioris, Manáos, Oct. 17, 1929, Ducke 22429 (B); dense forest, Manáos, alt. 25 m., Oct. 17, 1929, Killip $ Smith 30151 (MBG, NY, US). The type specimen of this species has not been available for study. The specimens collected by Ducke and Killip & Smith from the vicinity of Manáos aceord well with the original de- scription of F. Riedelii, the type specimen of which, collected by Riedel, is cited as from the Serra d'Estrella. It will not be surprising, therefore, if the specimens cited from Amazonas may eventually be proved to represent a distinct species. 17. Forsteronia Duckei Mgf. Notizblatt 9: 962. 1996. Stems relatively stout, glabrous, or very minutely puber- ulent when young, conspicuously lenticellate when fully ma- ture; leaves opposite, very shortly petiolate, oblong- to ovate- elliptie, apex very shortly and abruptly acuminate to obtuse, base very broadly obtuse to rounded, 9-16 cm. long, 4-5 cm. broad, firmly membranaceous, glabrous, inconspicuously glandular at the base of the midrib above; petioles 0.3-0.35 em. long; inflorescence thyrsiform, terminal, about equalling the subtending leaves or somewhat shorter, bearing many small, white flowers; primary peduncle minutely ferruginous-puber- ulent, ultimate branches regularly and determinately com- posed, minutely and densely puberulent-papillate; pedicels 0.1-0.15 em. long, minutely puberulent-papillate; bracts ovate, 0.01-0.025 cm. long, scarious; calyx-lobes ovate-reniform, broadly obtuse or rounded, 0.075-0.1 em. long, scarious, densely and minutely puberulent-papillate, the squamellae very mi- (211) [Vor. 22 178 ANNALS OF THE MISSOURI BOTANICAL GARDEN nute, solitary, alternate; corolla densely and minutely puber- ulent-papillate without, the tube 0.08-0.1 em. long, about 0.075 em. in diameter at the base and 0.125 em. in diameter at the orifice, villosulose within, the lobes oblong-elliptie, 0.2 em. long, essentially glabrous within; stamen filaments 0.05 em. long, the anthers slightly exserted, about 0.125 em. long, gla- brous; ovary ovoid, minutely papillate, about 0.07 em. long; stigma 0.2 em. long, slightly exserted beyond the connate anthers; nectaries somewhat shorter than the ovary ; follicles unknown. BRAZIL: PARA: ad ripas fluminis Anajaz, in parte occidentali insula Marajo, Nov. 25, 1922, Ducke 17478 (B, TYPE, US, MBG, photograph and analytieal drawings). 18. Forsteronia laurifolia (Benth.) A. DC. in DC. Prodr. 8: 438. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 106. 1860. Thenardia (? ) laurifolia Benth. in Hook. Jour. Bot. 3: 246. 1841. Thyrsanthus laurifolius (Benth.) Miers, Apoc. So. Am. 94. Stems relatively stout, glabrous, inconspicuously lenticel- late when fully mature; leaves opposite, petiolate, oblong to oblong- elliptie, apex shortly acuminate, base broadly obtuse to rounded, 6-14 cm. long, 2.5—5.0 em. broad, coriaceous or sub- coriaceous, glabrous, glandular at the base of the midrib above; petioles 0.5-1.3 cm. long; inflorescence very congested, thyrsi- form, both terminal and lateral, much shorter than the sub- tending leaves, bearing relatively few small, white flowers; primary peduncle minutely puberulent-papillate to glabrate, ultimate branches congested, but regularly and determinately composed, minutely puberulent-papillate; pedicels 0.16-0.2 cm. long, minutely puberulent-papillate; bracts ovate, 0.08-1.4 cm. long, scarious; calyx-lobes ovate, acute to acuminate, 0.1- 0.17 em. long, scarious, very minutely and densely puberulent- papillate without, the squamellae minute, solitary, alternate, oecasionally evidently lacking; corolla minutely and densely puberulent-papillate without, the tube 0.15-0.18 em. long, about 0.1-0.125 em. in diameter at the base and 0.175-0.2 em. in diam- eter at the orifice, villosulose within, the lobes oblong-elliptic, (212) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 179 0.2-0.25 em. long, puberulent-papillate within, spreading; stamen filaments 0.05-0.08 em. long, distinct and free, not ag- glutinated to the style, the anthers slightly exserted, 0.16- 0.2 em. long, glabrous; ovary ovoid, apocarpous, about 0.05 em, long, glabrous or essentially so; stigma 0.18-0.25 cm. long, somewhat exserted beyond the connate anthers; nectaries somewhat shorter than the ovary; immature follicles long and slender, continuous and more or less agglutinated, 23-30 em. long, glabrous or essentially so; mature seed unknown. BRITISH GUIANA: data incomplete, Schomburgk 953 (B, BB, Camb., D, FM, K, TYPE, NY, MBG, photograph and analytical drawings). BRAZIL: AMAZONAS: in vicinibus Barra, Jan., 1851, Spruce 999 (B, BB, D, G, M, NY); prope Panure ad Rio Uaupes, Oct., 1852-Jan., 1853, Spruce 2080 (B); Rio Negro pr. Joaquim, Jan., 1902, Ule 6057 (B, D); São Joaquim, Jan., 1902, Ule 77b (B); ad ripas inundatas super cataractam Tapuruguara, Santa Izabel, Rio Negro, Dec. 8, 1929, Ducke 22431 (B). The specimens cited show this species to be unusually shrubby for the genus, and Ducke has commented upon the label of his specimen ‘‘ Arbor erecta vel frutex scandens ??' 19. Forsteronia paludosa Woodson, Ann. Mo. Bot. Gard. 21: 620. 1934. Stems relatively slender, glabrous, conspicuously lenticel- late when fully mature; leaves opposite, very shortly petiolate, narrowly oblong-lanceolate, shortly acuminate to acute, base obtuse, 5-7 em. long, 1.3-1.6 cm. broad, firmly membranaceous or subcoriaceous, above glabrous, inconspicuously glandular at the base of the midrib, beneath inconspicuously barbellate in the axils of the midrib, otherwise glabrous; petioles 0.25- 0.3 cm. long; inflorescence compound-subspiciform, terminal, about equalling or somewhat shorter than the subtending leaves, bearing several small, white flowers; primary peduncle essentially glabrous or very irregularly and sparsely pilosu- lose-papillate above, ultimate branches indefinitely congested, scarcely manifest; pedicels 0.01-0.02 em. long, very minutely puberulent-papillate; braets minutely ovate, 0.01-0.03 em. long, scarious; calyx-lobes ovate, acute, 0.1-0,12 em. long, scarious, very minutely puberulent-papillate without, the squamellae evidently lacking; corolla very minutely and ir- (213) [Vor. 22 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN regularly puberulent-papillate without, the tube 0.1-0.12 cm. long, about 0.075 em. in diameter at the base and 0.125 em. in diameter at the orifice, minutely villosulose within, the lobes narrowly oblong, about 0.15 cm. long, densely pilosulose within; stamen filaments about 0.05 em. long, distinet and free, not agglutinated to the style, the anthers slightly exserted, 0.15 em. long, minutely barbellate; ovary ovoid, apocarpous, about 0.05 em. long, minutely papillate; stigma about 0.12 em. long; nectaries somewhat shorter than the ovary; follicles unknown. BRAZIL: PARA: loeis paludosis eampos de Cupijo prope Cameta, July 22, 1916, Ducke 21627 (B, TYPE, US, MBG, photograph and analytical drawings). 20. Forsteronia amblybasis Blake, Jour. Wash. Acad. Sci. 14: 292. 1924. Stems relatively stout, glabrous, or very minutely and in- definitely puberulent when young, conspicuously lenticellate when fully mature; leaves opposite, petiolate, ovate to oblong- elliptie, apex rather abruptly acuminate, base broadly obtuse or rounded, 6.0-14.5 em. long, 2.3-7.4 em. broad, membrana- ceous, above glabrous, glandular at the base of the midrib, beneath glabrous or minutely ciliolate-barbellate in the axils of the midrib; petioles 0.6-1.3 em. long; inflorescence sub- spiciform-paniculate, both terminal and lateral, shorter than the subtending leaves, or the terminal somewhat longer, bear- ing many small, greenish-white flowers; primary peduncle pu- berulent-papillate to glabrate below, ultimate branches indefi- nitely congested and scarcely manifest, puberulent-papillate ; pedicels 0.08-0.1 em. long, minutely puberulent-papillate, greatly accrescent in fruit; bracts oblong- to obovate-spathu- late, 0.2-0.65 em. long, scarious or somewhat foliaceous ; calyx- lobes laneeolate to ovate-lanceolate, acuminate, 0.16-0.3 cm. long, scarious or very slightly foliaceous, minutely and rather irregularly puberulent-papillate without, the squamellae ap- parently lacking; corolla puberulent-papillate without, the tube 0.12-0.15 em. long, about 0.05-0.07 em. in diameter at the base and 0.175-0.2 em. in diameter at the orifice, villosulose within, the lobes ovate-oblong, 0.13-0.15 em. long, densely (214) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 181 pilosulose within, spreading; stamen filaments 0.03-0.05 em. long, distinet and free, not agglutinated to the style, the anthers 0.13-0.16 cm. long, the tips slightly exserted, glabrous or occasionally minutely and sparsely barbellate; ovary ovoid, evidently syncarpous, or the carpels closely agglutinated, about 0.04 cm. long, minutely appressed-puberulent; stigma 0.11-0.12 cm. long; nectaries equalling or slightly surpassing the ovary; follicles very stout, continuous, 15-19 em. long, gla- brous; seeds 1.5-1.7 cm. long, the pale tawny coma 3.5-5.0 cm. long. BOLIVIA: LA PAZ: Tipuani-Guani, Dec., 1892, Bang 1689 (B, BB, MBG, NY, US, TYPE); ‘‘Songo,’’ Nov., 1890, Bang 850 (B, BB, FM, M, MBG, NY, US); Polo-Polo, bei Coroico, alt. 1100 m., Oct.—Nov., 1912, Buchtien 3953 (NY); SANTA CRUZ: bosque, Buena Vista, Prov. Sara, alt. 400 m., Dec. 3, 1924, Steinbach 6717 (B, D, FM, MBG, NY, US); feuchter Wald, Buena Vista, alt. 450 m., May 5, 1928, Steinbach 8020 (B, NY). 21. Forsteronia decipiens Woodson, spec. nov. Fruticosa volubilis ; ramulis erassiuseulis glabris maturitate conspicue lenticellatis; foliis oppositis petiolatis ovato- vel oblongo-ellipticis apice abrupte breviterque subcaudato-acum- inatis basi late obtusis rotundatisve rarius obscurissime cordatis 7.5-13.0 em. longis 2.8-8.0 em. latis firme membra- naceis supra paululo nitidulis nervo medio basi pauciglandu- ligero caeterumque glabris subtus opacis glabris vel incon- spicuissime barbellatis; petiolis 0.6-1.1 em. longis glabris; in- florescentiis anguste irregulariterque subspiciformi-thyrsoi- deis et terminalibus et lateralibus foliis paululo brevioribus flores multas parvas albidas gerentibus; pedunculo irregu- lariter sparseque puberulo vel glabrato ; pedicellis 0.1-0.12 em. longis post maturitatem conspicue acerescentibus minute pu- berulo-papillatis; bracteis 0.1-0.5 em. longis minute puberulo- papillatis subfoliaceis maioribus oblongo- vel obovato- spathu- latis minoribus pedicellos subtendentibus anguste lanceolatis; calycis laciniis ovato-elliptieis acutis 0.18-0.25 em. longis sub- foliaceis minute puberulo-papillatis squamellis non visis; corollae albidae extus minute papillatae tubo campanulato 0.1-0.12 em. longo basi ca. 0.05 cm. diametro metiente faucibus ea. 0.15 em. diametro metientibus intus villosulo lobis ovato- (215) [Vor. 22 182 ANNALS OF THE MISSOURI BOTANICAL GARDEN oblongis 0.18-0.2 em. longis intus dense villosulis patulis; fila- mentis staminum 0.04-0.05 em. longis stylum haud aggluti- natis antheris 0.13-0.15 em. longis apice paulo exsertis sparse barbellatis rariusve glabratis; ovario ovoideo ut videtur syn- carpo vel carpellis inter se agglutinatis ca. 0.04 em. longo mi- nute puberulo; nectariis ovarium aequantibus vel paululo superantibus; follieulis crassiusculis rigidis continuis divari- catis 10-13 em. longis glabris; seminibus 1.6-1.7 cm. longis como dilute aurantiaco 4.0—4.5 em. longo. Stems relatively stout, glabrous, conspicuously lenticellate at maturity; leaves opposite, petiolate, ovate- to oblong-ellip- tie, apex abruptly and shortly subcaudate-acuminate, base broadly obtuse to rounded, rarely obscurely cordate, 7.5- 13.0 em. long, 2.8-8.0 em. broad, firmly membranaceous, above slightly nitidulous, glandular at the base of the midrib, other- wise glabrous, beneath opaque, glabrous or very inconspicu- ously barbellate in the axils of the midrib; petiole 0.6-1.1 em. long, glabrous; inflorescence narrowly and rather irregularly subspiciform-thyrsiform, both terminal and lateral, somewhat shorter than the subtending leaves, oceasionally conspicuously longer when terminal, bearing many small, white flowers; peduncle irregularly and sparsely puberulent to glabrate; pedicels 0.1-0.12 em. long, conspicuously acerescent in fruit, minutely puberulent-papillate; bracts 0.1-0.5 em. long, more or less subfoliaceous, minutely puberulent-papillate, the larger oblong- to obovate-spathulate, the smaller which immediately subtend the pedicels narrowly lanceolate; calyx-lobes ovate- elliptic, acute, 0.18-0.25 em. long, subfoliaceous, minutely pu- berulent-papillate, squamellae not seen; corolla whitish, mi- nutely papillate without, the tube campanulate, 0.1-0.12 em. long, about 0.05 em. in diameter at the base and about 0.15 em. in diameter at the orifice, villosulose within, the lobes ovate- oblong, 0.18-0.2 em. long, densely villosulose within, spreading; stamen filaments 0.04-0.05 em. long, distinet and free, not ag- glutinated to the style, the anthers 0.13-0.15 em. long, slightly exserted at the tips, sparsely barbellate to glabrate; ovary ovoid, evidently synearpous or the carpels agglutinated, about 0.04 em. long, minutely puberulent; nectaries about equalling (216) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 183 or slightly surpassing the ovary; follieles relatively stout and rigid, sharply divaricate, 10-13 em. long, glabrous; seeds 1.6- 1.7 cm. long, the pale tawny coma 4.0—4.5 em. long. PERU: LORETO: forest, Balsapuerto, alt. 220 m., Febr., 1933, Klug 2905 (NX; TYPE, MBG, photograph); same locality, June, 1933, Klug 3104 (NY, fruiting co- TYPE, MBG, photograph); SAN MARTIN: sandy soil, Tarapoto, alt. 360-900 m., Dec. 13, 1929, Williams 6156 (FM, MBG, US). Although this species is very closely related to the preced- ing, for which it was mistaken for some time by the writer, the size and composition of the bracts render the inflorescence quickly and accurately distinguishable. Both F. decipiens and F. amblybasis have long passed for the less frequent F. mon- tana Muell.-Arg. of eastern Brazil. 22. Forsteronia montana Muell.-Arg. in Mart. Fl. Bras. 61: 101. 1860; Miers, Apoc. So. Am. 245. 1878. Forsteronia Sellowii Muell.-Arg. loc. cit. 1860; Miers, loc. cit. 1878 Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, petiolate, elliptic-lanceo- late to oval-obovate, apex acuminate, base broadly obtuse to rounded, firmly membranaceous, above glabrous, glandular at the base of the midrib, beneath glabrous or inconspicuously barbellate in the axils of the midrib, 3.5-10.6 cm. long, 1.1— 5.4 em. broad; petioles 0.5-1.2 em. long; inflorescence thyrsi- form, terminal, or lateral as well, about equalling or somewhat surpassing the subtending leaves, occasionally shorter, par- tieularly when lateral, bearing many small, white flowers; primary peduncle minutely puberulent to glabrate below, ulti- mate branches indefinitely congested and scarcely manifest, minutely puberulent to puberulent-papillate; pedicels 0.07— 0.15 em. long, puberulent-papillate; bracts ovate-lanceolate to linear, scarious, 0.05-0.1 em. long; calyx-lobes ovate, acute to acuminate, 0.11-0.18 cm. long, scarious, puberulent-papillate without, the squamellae evidently lacking; corolla puberulent- papillate without toward the lobes, the tube 0.11—0.13 em. long, about 0.04—0.07 em. in diameter at the base and 0.12—0.14 cm. in diameter at the orifice, minutely villosulose within, the lobes (217) [Vor. 22 184 ANNALS OF THE MISSOURI BOTANICAL GARDEN ovate-oblong, 0.17-0.2 em. long, slightly spreading, minutely pilosulose within; stamen filaments 0.05-0.07 em. long, distinct and free, not agglutinated to the style, the anthers 0.16-0.18 em. long, the tips minutely barbellate, slightly exserted; ovary ovoid, syncarpous or agglutinated, minutely puberulent to hirtellous; stigma 0.12-0.14 em. long; nectaries somewhat shorter than the ovary ; follicles relatively stout and rigid, con- tinuous, 22-28 em. long, glabrous; seeds unknown, the pale tawny coma 5-6 em. long. BRAZIL: RIO DE JANEIRO: silvis caeduis, Sebastianopolis, date lacking, Martius 183 (M); data incomplete, Glaziou 7532 (B); DATA INCOMPLETE: Sellow s.n. (Bx, IsOTYPE, NY, MBG, photograph and analytical drawings); Sellow 396 (B); Riedel s.n. (BB). 23. Forsteronia pubescens A. DC. in DC. Prodr. 8: 436. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 104. 1860; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Thyrsanthus pubescens (A. DC.) Miers, Apoc. So. Am. 101. 1878. Thyrsanthus placidus Miers, loc. cit. 1878. Stems relatively stout, hirtellous when young, eventually be- coming glabrate and conspicuously lenticellate when fully ma- ture; leaves opposite, petiolate, broadly oval to narrowly ob- long-elliptic, apex obtuse to acuminate, base obtuse to acutish, 3.0-12.5 em. long, 1.5-7.5 em. broad, membranaceous, above generally puberulent to glabrate, rarely essentially glabrous, inconspicuously glandular at the base of the midrib, beneath softly and generally puberulent; petioles 0.4—0.8 em. long, densely puberulent ; inflorescence rather narrowly thyrsiform, terminal, usually equalling or somewhat surpassing the sub- tending leaves, less frequently shorter, bearing many small, white flowers; primary peduncle puberulent as upon the stem, ultimate branches regularly determinate, minutely puberulent ; pedicels 0.1—0.2 em. long, minutely puberulent; bracts ovate to lanceolate, 0.075-0.6 em. long, slightly foliaceous to scarious; calyx-lobes ovate to ovate-lanceolate, acute to acuminate, 0.11- 0.2 em. long, scarious, rather inconspicuously puberulent-papil- late without, the squamellae 5-6 times as numerous as the lobes, narrowly linear, indefinitely and regularly distributed ; corolla (218) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 185 essentially glabrous without, the tube 0.07-0.12 em. long, about 0.05-0.1 em. in diameter at the base and 0.1-0.18 em. in diameter at the orifice, densely villosulose within, the lobes narrowly ob- long, 0.3-0.5 em. long, widely spreading, pilosulose within; ann filaments 0.2-0.3 em. long, connate and agglutinated the style, the anthers 0.2-0.25 em. long, glabrous, wholly ex- serted; ovary ovoid, apocarpous, minutely puberulent-papil- late, about 0.05 em. long; stigma and style 0.11-0.17 em. long; nectaries about equalling, or slightly shorter than the ovary; follieles rather stoutish, very obseurely undulate-artieulated, usually more or less faleate, 11-27 em. long, glabrous; seeds 0.9-1.2 em. long, the bright tawny coma 2.5-3.5 em. long. BRAZIL: CEARA: data incomplete, 1837-41, Gardner 1761 (BB, TYPE, Bx, Camb., D, NY, US, MBG, Po ONE pu analytical drawings); BAHIA: Jacobina, date Making, Blanchet 2652 (B, Bx, D, NY); Catingas, date lacking, Blanchet 2452 (D); Rio Preto, date lacking, Pohl 2189 n ; in eampis ad Caitete, Nov., year lacking Martius s.n. (V); MINAS GERAES: data incomplete, Claussen s.n. (D); RIO DE JANEIRO: data incomplete, Glaziou 12939 (B, D); MATTO GROSSO: Santa Anna da Chapada, Oct. 13, 1902, Malme 2489 (D, S, US). BOLIVIA: BENI: junction of rivers Beni and Madre de Dios, Aug., 1886, Rusby 2526 (B, BB, FM, ANTA CRUZ: bosque del Rio Cuchi, Prov. Sara, alt. 500 m., Aug. 16, 1995, Feci 7782 (B, D, FM, MBG, U); Urubu, Prov. E alt. 450 m., Oet. 28, 1925, Steinbach 7295 (B, D, FM, MBG, NY, U). PARAGUAY: in regione collium **Cerros de Tobaty,’’ Sept., 1900, Hassler 6362 (B, BB); in regione lacus Ypacaray, Aug., 1913, neriti 12229 (B, BB, D, U8); in niet pr. Ita, Sept., 1885-95, Hassler 1038 (BB); in altaplani sn x declivibus ‘‘Serra de Amambay,’’ Oct., 1907, Rojas 9698 (B); Miri. Oct., 1929, Jørgen- sen 3444 (FM, MBG, NY, US); Coneepeion de Paraguay, Sept., qua Kuntze s.n. (B, NY); Colonia Risso pr. Rio Apa, Oet. 4, 1893, visio 1038 (B, S); zwischen Rio Apa und Rio Aquidaban, San Luis, March, year lacking, Fiebrig 5162 (B); bewaldetes Tal, Cordillera de Albo Oct. 11,1 1902, he 218a (B); dans les bois, Villa-Rica, Oct. 21, 1874, Balansa 1371 (BB, Bx, D) ; dans les haies, 1’Assomption, 177 1874-Oet. 25, 1875, Balansa 1371a (B, BB, Bx); data ineomplete, Morong 810 (BB, FM, MBG, NY, US). ee JUJUY: San Lorenzo, Oct. 31-Nov. 3, 1873, Lorentz $ Hieronymus 216 (B, , US); entre ore: y San Antonio, Oct. 30, 1873, Lorentz 4 i ed 358 (BD A: en bosque alto, Abra Grande, Dept. Oran, alt. 750 m., Nov. 10, 1927, DAT pur (FM, MBG, US) It appears probable that in Paraguay and northern Argen- tina hybridization may occur between this species and F. gla- brescens Muell.-Arg., also a very frequent liana of forests and wooded areas. Fiebrig 5162 is indicative of this supposition, the anthers being about the size of those of the latter species, (219) [Vor. 22 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN and eonspieuously smaller than those of the former. The num- ber and shape of the squamellae and the obtuse tips of the calyx-lobes are also very suggestive of F. glabrescens. On the other hand, the texture and pubescence of the foliage, as well as the size, conform with those of F. pubescens. The anthers of the specimens collected by Lorentz and Hieronymus, as well as the tips of the calyx-lobes, are also similar to those of F. gla- brescens, and the indument of the lower surface of the leaves, which allies them with F. pubescens, is unusually scant and somewhat more conspicuous in the axils of the midrib. A more significant difference is found in the follicles, fortunately pres- ent upon several of these specimens, which are somewhat stouter than those typieal of F. glabrescens, but more slender than those of F. pubescens and with conspicuous articulations suggestive of the moniliform fruits of the former species. The suggestion presents itself that more intimate study, partieu- larly in the field, may disclose that an intermediate species exists, probably evolved through hybridization. 24. Forsteronia cordata (Muell.-Arg.) Woodson, comb. nov. Forsteronia pubescens A. DC. p. cordata Muell.-Arg. in Mart. Fl. Bras. 6': 104. 1860, Stems relatively stout, minutely tomentulose when young, eventually becoming glabrate and conspicuously lenticellate when fully mature; leaves opposite, very shortly petiolate, oblong-elliptic to oval, apex very shortly and abruptly acumi- nate to rounded, base rounded to obscurely cordate, 2.5-7.0 em. long, 1.5-4.0 em. broad, firmly membranaceous, above minutely and rather sparsely pilosulose-strigillose, inconspicuously glandular at the base of the midrib, beneath densely tomentu- lose; petioles 0.2-0.4 em. long, minutely tomentulose; inflo- rescence broadly subeorymbose-thyrsiform, terminal, infre- quently lateral as well, about equalling or somewhat shorter than the subtending leaves, bearing many small, yellowish (dark yellowish-red in desiecation) flowers; primary pedunele minutely puberulent to tomentulose, ultimate branches regu- larly and determinately composed, minutely puberulent ; pedi- cels 0.2-0.4 em. long, minutely puberulent; bracts lanceolate, 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 187 0.08-0.25 em. long, only slightly foliaceous; calyx-lobes ovate, acute to acuminate, 0.1-0.15 em. long, scarious, minutely and rather irregularly puberulent, the squamellae about 3-4 times as many as the lobes, regularly and indefinitely distributed; corolla conspicuously puberulent-papillate without at the tips of the lobes, otherwise essentially glabrous to very minutely and indefinitely papillate, the tube 0.07-0.1 em. long, about 0.05 em. in diameter at the base and 0.13-0.17 cm. in diameter at the orifice, densely villosulose within, the lobes oblong-lingu- late, 0.35-0.43 em. long, essentially glabrous or very minutely and indefinitely papillate within, widely spreading; stamen fila- ments 0.3-0.32 em. long, connate and agglutinated to the style, the anthers wholly exserted, 0.25-0.27 em. long, glabrous; ovary ovoid, about 0.05 cm. long, minutely puberulent to puber- ulent-papillate, apocarpous; stigma 0.1-0.125 cm. long, the style 0.25—0.27 em. long; nectaries somewhat shorter than the ovary ; follieles relatively slender, rather obscurely articulated, more or less falcate, 18-25 em. long, glabrous; seeds 1.3- 1.5 em. long, the pale tawny coma 2.0-2.5 em. long. BRAZIL: RIO DE JANEIRO: in collibus inter frutiees margine locis arenosis pr. R. Jan., Nov., 1829, Lhotzky 29 (B, TYPE, MBG, photograph and analytical drawings); in der Restinga bei Copaeabana, Nov., 1896, Ule 4288 (B); same locality, May, 1897, Ule 4434 (B); data incomplete, Glaziou 14064 (B, FM); Constantino 8675 (B, U, US); DATA INCOMPLETE: Riedel s.n. (B, BB, COTYPE); Regel (Camb.). It appears reasonable to allow these plants specific rank, since they exhibit several important and conspicuous morpho- logical differences distinguishing them from true F. pubescens. There is furthermore no evidence of any degree of intergrada- tion between F. cordata and the former species. 29. Forsteronia mollis Rusby, Mem. Torrey Bot. Club 4: 218. 1895. Stem relatively stout, puberulent when young to glabrate or glabrous, conspicuously lenticellate when fully mature; leaves opposite, petiolate, oblong-elliptie to broadly oval, apex shortly and abruptly acuminate, base broadly obtuse or rounded, 7- 12 em. long, 4.0-7.5 cm. broad, membranaceous, glandular at the base of the midrib above; petioles 0.7-0.9 em. long; inflores- (221 [Vor. 22 188 ANNALS OF THE MISSOURI BOTANICAL GARDEN cence thyrsiform, terminal, about equalling or somewhat shorter than the subtending leaves, bearing many small, white or greenish flowers; primary peduncle puberulent to glabrate below, ultimate branches determinately composed, densely puberulent; pedicels 0.05-0.15 em. long, minutely and densely puberulent; bracts lanceolate to ovate-lanceolate, 0.08-0.2 em. long, only slightly foliaceous; calyx-lobes broadly ovate-sub- orbicular, broadly obtuse or rounded, 0.1-0.125 cm. long, searious, minutely and densely puberulent-papillate without, the squamellae 3-5 times as many as the lobes, regularly and indefinitely distributed; corolla essentially glabrous without, the tube 0.08-0.1 em. long, about 0.07 em. in diameter at the base and 0.15 em. in diameter at the orifice, villosulose within, the lobes 0.32-0.4 em. long, minutely puberulent-papillate to es- sentially glabrate within, widely spreading; stamen filaments 0.15-0.22 em. long, more or less connate and agglutinated to the style, the anthers 0.25 em. long, wholly exserted, glabrous; ovary apocarpous, oblong-ovoid, minutely puberulent; stigma 0.12-0.15 em. long, the style 0.15-0.175 em. long; nectaries somewhat shorter than the ovary ; follieles unknown. Var. typica. Stem puberulent when young, eventually becoming glabrate; leaves rather sparsely puberulent above, minutely tomentulose beneath ; all other essential characters as in the species. BOLIVIA: LA PAZ: ‘‘Yungas,’’ 1890, Bang 274 (B, BB, FM, MBG, NY, TYPE, US). Var. foliosa (Rusby) Woodson, comb. nov. Forsteronia folivsa Rusby, Deser. So. Am. Pl. 90. 1920. Stem and leaves glabrous; all other essential characters as in the species. LIVIA: LA PAZ: Polo-Polo bei Coroico, Nordyungas, alt. 1100 m., Oct.—Nov., Bou 1912, Buchtien 3876 (B, NY, TYPE, US, MBG, photograph and snaiytisal draw- ings); same data, Buchtien 3677 (US). This combination is made with some trepidation, for although the presence and lack of a vegetative indument is striking in these two varieties, the general aspect of the speci- mens is conspicuously similar. They certainly should not be (222) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 189 interpreted as species upon the slight basis of available specimens. 26. Forsteronia subcordata K. Sch. in herb. Frutieosa volubilis; ramulis crassiusculis glabris vel juven- tate minutissime papillatis maturitate conspicue lenticellatis ; foliis oppositis breviter petiolatis oblongo-ellipticis apice acuminatis basi obscure cordatis 5.5-11.0 em. longis 2.54.2 em. latis tenuiter membranaceis glaberrimis nervo medio supra basi pauciglanduligero; petiolis 0.3-0.45 em. longis; inflores- centiis thyrsiformibus terminalibus foliis conspicue brevior- ibus flores multas albas gerentibus; pedunculo puberulo-papil- lato; pedicellis 0.1-0.2 em. longis minute puberulo-papillatis ; braeteis ovatis 0.05-0.2 em. longis scariaceis; calycis laciniis ovatis acutis acuminatisve 0.1—0.17 em. longis scariaceis extus minute puberulo-papillatis squamellis minutissimis plus mi- nusve numerosis regulariter positis; corollae extus glabrae vel indistinctissime papillatae tubo 0.07-0.1 em. longo basi ca. 0.05- 0.07 cm. diametro metiente faucibus ca. 0.1-0.15 cm. diametro metientibus intus minute villosulo lobis 0.3-0.4 em. longis patulis prope apieem minute puberulo-papillatis; filamentis staminum 0.15-0.17 cm. longis stylum agglutinatis antheris 0.18-0.2 em. longis exsertis glabris; ovario ovoideo apocarpo ca. 0.04 em. longo minute papillato; stigmate 0.11-0.15 cm. longo; stylo ca. 0.12-0.17 em. longo; nectariis ovarium subae- quantibus ; folliculis ignotis. Stems relatively stout, glabrous or very minutely papillate when young, becoming conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptie, apex acuminate, base obseurely cordate, 5.5-11.0 em. long, 2.5- 4.2 em. broad, delicately membranaceous, glabrous, inconspicu- ously glandular at the base of the midrib above; petioles 0.3— 0.45 cm. long; inflorescence thyrsiform, terminal, conspicu- ously shorter than the subtending leaves, bearing numerous small, white flowers; primary peduncle very minutely puberu- lent, ultimate branches determinately eomposed, puberulent- papillate; pedicels 0.1-0.2 em. long, minutely puberulent-papil- late; braets ovate, 0.05-0.2 cm. long, scarious; calyx-lobes (223) [Vor. 22 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN ovate, acute to acuminate, 0.1-0.17 em. long, searious, minutely puberulent-papillate without, the squamellae very minute, numerous, indefinitely and regularly distributed; corolla es- sentially glabrous or very indefinitely papillate without, the tube 0.07-0.1 em. long, about 0.05-0.07 em. in diameter at the base and 0.1-0.15 em. in diameter at the orifice, villosulose within, the lobes oblong-lingular, 0.3-0.4 em. long, widely spreading, minutely puberulent-papillate toward the tip within; stamen filaments 0.15-0.17 em. long, connate and ag- glutinated to the style, the anthers 0.18-0.2 em. long, exserted, glabrous; ovary ovoid, apocarpous, about 0.04 em. long, mi- nutely papillate; stigma 0.11-0.15 em. long, the style 0.12- 0.17 em. long; neetaries about equalling the ovary; follicles unknown. ECUADOR: MANABI: in sylvestrib. ad El Recreo, Febr. 13, 1897, Eggers 15618 (B, TYPE, FM, M, MBG, NY). 27. Forsteronia leptocarpa (Hook. & Arn.) A. DC. in DC. Prodr. 8: 438. 1844 Parsonsia leptocarpa Hook, & Arn. in Hook. Jour Bot. 1: 287. 1834. Forsteronia brasiliensis A. DC. loc. cit. 436. 1844; Muell.- Arg. in Mart. Fl, Bras. 6': 102. 1860; K. Sch. in Engl. € Prantl, Nat. Pflanzenfam. 4°: 187. 1895. Forsteronia brasiliensis A. DC. p. bahiensis Muell.-Arg. loc. cit. 1860. Thyrsanthus Brasiliensis (A. DC.) Miers, Apoc. So. Am. 103. 1878. Laseguea leptocarpa (Hook. & Arn.) Miers, loc. cit. 254. 1878, Thyrsanthus leptocarpus (Hook. € Arn.) Griseb. Gött. Abh. 24: 224. 1879. Forsteronia leptocarpa (Hook. & Arn.) A. DC, var. glaber- rima Niederl. Bol. Mens. Prodr. Argent. 3': 315. 1890, nom. nud, Stems relatively stout, more or less puberulent or pilosulose when young, soon becoming glabrate and conspicuously lenti- cellate when fully mature; leaves opposite, shortly petiolate, (224) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 191 oblong-elliptic to broadly oval, apex rather shortly and abruptly acuminate to obtuse, base obtuse to rounded, 2.0- 10.5 em. long, 1.3-5.5 em. broad, firmly membranaceous, above glabrous, glandular at the base of the midrib, beneath minutely barbellate in the axils of the midrib; petioles 0.2-0.7 em. long, minutely puberulent-papillate to glabrous; inflorescence thyr- siform, terminal, shorter than the subtending leaves, bearing numerous, congested, small, white flowers; primary peduncle more or less minutely puberulent to glabrate below, ultimate branches determinately composed, puberulent to puberulent- papillate; pedicels 0.07-0.125 cm. long, puberulent-papillate ; braets ovate, 0.07—0.2 em. long, scarious; ealyx-lobes ovate, acute to acuminate, 0.11-0.2 em. long, scarious, puberulent- papillate without, the squamellae 3-5 times as many as the lobes, indefinitely and regularly distributed ; corolla essentially glabrous without, the tube 0.1-0.17 em. long, about 0.07-0.1 em. in diameter at the base and 0.1—0.2 em. in diameter at the orifice, villosulose within, the lobes 0.25-0.32 em. long, villosulose at the base within, otherwise essentially glabrous to indefinitely and minutely papillate, widely spreading; stamen filaments 0.11-0.2 em. long, connate and agglutinated to the style, the anthers 0.17-0.25 em. long, exserted, glabrous; ovary apocar- pous, ovoid, about 0.05 em. long, minutely papillate; stigma 0.15-0.2 em. long; style 0.08-0.15 em. long; nectaries somewhat shorter than the ovary ; follicles relatively slender, rather ob- securely articulated, usually somewhat tortuous, 16-34 cm. long, glabrous; seeds 0.8-1.3 em. long, the tawny coma 3.54.0 em. long. BRAZIL: PERNAMBUCO: Flussufer, Tapera, 1926, Pickel 1222 (B); data in- complete, Gardner 1059 (BM, Camb., FM, S, US, V); Schornbaum s.n. (Bx); BAHIA: Jacobina, date lacking, Blanchet 3229 (BB, D, DC); Vittoria, date lack- ing, Sellow 1542 (B); Sta. Anna, May, 1850, Blanchet 3431 (BB, DC); MINAS GERAES: data ineomplete, Glaziou 14063 (D); RIO DE JANEIRO: Corvocado, date lacking, Pohl s.n. (Bx, V); Lagóa de Itapemerim, Dee. 9, 1915, Frazáo 7147 (B, S, U, US); ad urbem in silvis inter Gavea et Vista Chineza, Jan. 15, 1929, Ducke 21810 (B, U); in der Restinga de Gavea, Sept., 1899, Ule 4845 (B); bei Porto das Caixas, Dec., 1897, Ule 4578 (B); Ypanema, in silva sat aperta, Oct. 10, 1904, Dusen 5097 (D, S, US); Copacabana, Jan. 25, 1887, Schenck 2269 (B); Morro do Quitambo, Jan. 4, 1887, Schenck 1875 (B) ; data ineomplete: Guillemin 168 (DC); Gaudichaud 534 (DC); Lhotzky 74 (B); Andersson s.n. (S); Glaziou 2507 (Bx); (225) [Vor. 22 192 ANNALS OF THE MISSOURI BOTANICAL GARDEN Glaziou 4206 (B); são PAULO: Barra do Juquia, Oct. 14, 1894, Hoehne 11148 (B); Santos, Jan. 15, 1875, Mosen 3434 (C, S); PARANA: Rio Cubatão, in silva fluminali, Dee. 28, 1911, Dusen 13652 (S); Jaearehy, in silva primaeva reg. lit., March 22, 1914, Dusen 14632 (FM, MBG, NY, 8); SANTA CATHARINA: Gebüsch bei Tubaráo, Dee., 1889, Ule 1543 (B); data incomplete, Pabst 557 (B); RIO GRANDE DO SUL: Porto Alegre, Dec., 1898, Reineck $ Czermak 467 (B); data incomplete, Tweedie s.n. (K, TYPE, MBG, photograph); DATA INCOMPLETE: Riedel s.n. (B, BB, D, FM, M, NY, U); Vidensis s.n. (Bx); Glaziou 2507 (C); Sellow 388 (B, Bx). Also reported by Mueller from the State of Espirito Santa. F. leptocarpa is one of the most characteristic and uniform species of the genus Forsteronia, particularly in view of its wide distribution. 28. Forsteronia australis Muell.-Arg. in Mart. Fl. Bras. 6': 103. 1860; Miers, Apoc. So. Am. 246. 1878. Thyrsanthus crebriflorus Miers, loc. cit. 105. 1878. Stems relatively stout, glabrous or minutely and indefinitely papillate when very young, becoming conspicuously lenticel- late when fully mature; leaves opposite, petiolate, elliptic, apex rather abruptly acuminate, base obtuse, 3.5-10.0 em. long, 2.0— 5.5 em. broad, firmly membranaceous, either surface glabrous, ineonspieuously glandular at the base of the midrib above; petioles 0.25-0.7 em. long, glabrous; inflorescence thyrsiform, terminal, nearly equalling or slightly surpassing the subtend- ing leaves, bearing many small, white flowers; primary pe- dunele more or less densely puberulent, ultimate branches de- terminately composed, puberulent-papillate; pedicels 0.05- 0.1 em. long, puberulent-papillate; braets ovate to linear, 0.07- 0.2 em. long, searious or only slightly foliaceous; calyx-lobes ovate-deltoid, obtuse or rounded, 0.08-0.12 em. long, scarious, densely puberulent-papillate without, the squamellae 3—5 times as many as the lobes; corolla glabrous without, the tube 0.07— 0.1 em. long, about 0.07 em. in diameter at the base and 0.12- 0.15 em. in diameter at the orifice, minutely villosulose within at the throat, the lobes oblong-elliptie, 0.3—0.35 em. long, rather irregularly puberulent within, widely spreading; stamen fila- ments 0.18-0.2 em. long, connate and agglutinated to the style, the anthers wholly exserted, 0.22-0.25 em. long, glabrous; ovary ovoid, apocarpous, about 0.04 em. long, puberulent-papil- (226) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 193 late; stigma 0.15-0.17 cm. long; style 0.1—0.2 em. long; nectaries about equalling the ovary ; follicles unknown. BRAZIL: BAHIA: S. Thomé, 1857, Blanchet 3757 (Bx, DC); Igreja Velha, 1841, Blanchet 3431 (B, DC, NY); MINAS GERAES: banks of the Rio Parahyba, 1842, Gardner 5013 (B, BM, Camb., NY, US); aovaz: Ponte Alta, au Chapadáo do Gama, date lacking, Glaziou 21724 (B, S); DATA INCOMPLETE: Sellow 460 (S). Differing from the closely related F. leptocarpa not only in the key characters but in the larger inflorescence and absolutely glabrous leaves. 29. Forsteronia obtusiloba Muell.-Arg. Linnaea 30: 413. 1860; Miers, Apoc. So. Am. 246. 1878. Stems relatively stout, glabrous, eonspicuously lenticellate when fully mature; leaves opposite, petiolate, ovate- to oblong- elliptie, apex acuminate, base broadly obtuse or rounded, 4.5- 8.0 em. long, 2-4 em, broad, membranaceous, glabrous, incon- spicuously glandular at the base of the midrib above; petioles 0.4—0.6 em. long, glabrous; inflorescence rather laxly thyrsi- form, terminal, about equalling the subtending leaves, bearing many small, white flowers; primary peduncle glabrate or very indefinitely papillate, ultimate branches determinate, puberu- lent-papillate; pedicels 0.1-0.15 em. long, very minutely puber- ulent-papillate; braets ovate, 0.07—0.12 cm. long, scarious; ealyx-lobes ovate-deltoid, broadly obtuse or rounded, 0.08- 0.1 em. long, scarious, minutely papillate or puberulent-papil- late without, the squamellae 3-5 times as many as the lobes; corolla essentially glabrous without, the tube 0.07-0.09 em. long, about 0.05 em. in diameter at the base and 0.15 em. in diameter at the orifice, villosulose within, the lobes oblong- elliptic, 0.3-0.35 em. long, densely puberulent-papillate within, widely spreading; stamen filaments 0.18-0.22 em. long, ag- glutinated to the style, the anthers 0.2 cm. long, glabrous; ovary ovoid, apocarpous, about 0.05 em. long, densely papil- late; stigma 0.18 em. long; style 0.1 em. long; nectaries about equalling the ovary ; follicles unknown. VENEZUELA: DISTRITO FEDERAL [?]: **prov. de Caracas, Curucuti,’’ alt. 2000 pp., Febr., 1846, Funck $ Schlim 310 (BB, DC, TYPE, MBG, photograph and analytical drawings). (227) [Vor. 22 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN 30. Forsteronia glabrescens Muell.-Arg. in Mart. Fl. Bras. 6': 102. 1860. Thyrsanthus glabrescens (Muell.-Arg.) Miers, Apoc. So. Am. 102. 1878. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, elliptie, apex shortly acuminate to subobtuse, base obtuse to acute, fre- quently more or less cuneate, 3.0-6.5 em. long, 1.2-2.7 em. broad, firmly membranaceous to subeoriaceous, glabrous; pet- ioles 0.2-0.6 em. long, glabrous; inflorescence amply thyrsi- form, terminal, about equalling or somewhat surpassing the subtending leaves, bearing many small, white flowers; primary pedunele minutely puberulent, ultimate branches puberulent- papillate, determinately composed; pedicels 0.05-0.1 em. long, puberulent-papillate; bracts ovate, 0.05-0.25 em. long, scarious or only slightly foliaceous ; calyx-lobes ovate, acute to obtuse, 0.1-0.12 em. long, scarious, very minutely papillate to essen- tially glabrous without, the squamellae 3-5 times as many as the lobes, indefinitely and regularly distributed; corolla gla- brous without, the tube 0.08-0.1 em. long, about 0.05-0.07 em. in diameter at the base and 0.1-0.15 em. in diameter at the ori- fice, minutely villosulose within, the lobes 0.25-0.34 cm. long, pilosulose within, widely spreading; stamen filaments 0.13- 0.2 em. long, agglutinated to the style, the anthers 0.15—0.2 em. long, glabrous, wholly exserted; stigma 0.11-0.14 cm. long; style 0.08-0.11 em. long; ovary ovoid, minutely papillate, apo- carpous, about 0.04 em. long; neetaries somewhat shorter than the ovary ; follicles relatively slender, rather distantly monili- form, subparallel or more or less tortuose, 15-33 em. long, gla- brous ; seeds 0.65-0.75 em. long, the pale tawny coma 2.7-3.2 em. long. BRAZIL: SAO PAULO: Batataes, March, 1849, Regnell III 882 (S); PARANA: Fortaleza, in silvula, Febr. 26, 1910, Dusen, 9457 (D, FM, MBG, S, US) ; RIO GRANDE DO SUL: Uferwald des Rio Camaquam, 1887, Ihering 8 (B); in margine silvula, loeo sat sicco, Cachoeira, Febr. 20, 1893, Malme 602 (S); Ilha dos Marinheiros pr. Rio Grande, in arena plus minusve mobilis, Nov. 24, 1892, Malme 364 (S); in margine silvularum, loco sat sicco, Porto Alegre, Nov. 3, 1892, Malme 248 (S); in Waldringen an der Avenida Ernesto, Porto Alegre, Jan., 1899, Reineck 467 (D); data incomplete, Ihering 300 (B); DATA INCOMPLETE: Sello 1473 (B). (298) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 195 BOLIVIA: TARIJA: Bermejo, alt. 1400 m., Nov. 30, 1903, Fiebrig 2310 (B, D, M, 8, U, US) PARAGUAY: common in woods and hedges, Villarica, Nov. 9, 1928, Jørgensen 3445 (FM, MBG, NY, US); Cordillere de EN cT au dessus de us Nov. 1, 1883, Balansa 4606 (BB, D); dans les foré raguari, Oct. 1876-July, 1877, Balansa 1369 (B, BB, Bx, D, m bewaldetes sd Yes ra de Kw Oet. 11, 1902, Fiebrig 218 (B, D, FM, M); zwischen Rio Apa und Rio Aquidaban, San Luis, Oct. 31, 1908, Fiebrig 4157 (B, D, MO; in silva pr. Cerro Leon, June, 1895, Hassler 417 (BB); in dumetis = Campo Duarte, Oet., 1885, Hassler 1350 (BB); San Ber- nardino, June, year lacking, Hassler 3032 (BB); ad marg, silvae, Cordillera de Altos, Oet., year lacking, Hassler 3345 (B, BB); in regione fluminis Corrientes, ept., year lacking, Hassler 4528 (B, BB); in altaplanitie et decliviis ** Sierra de Maracayu,’’ Oct., year lacking, Hassler 5136 (BB); in altaplanitie ‘‘Sierra de Amambay,’’ Sept., 1912, Hassler 11962 (B, D, FM); in regione lacus Ypacaray, Oet., 1913, Hassler 12326 (B, D, MBG, US) ARGENTINA: FORMOSA: Guayculee, Febr., 1918, Jørgensen 3573 (BA, MBG); exact locality laeking, Nov., 1891, Niederlein 67b (B); cHaco: Mocovi, Nov. 13, 1903, Venturi 46 (BA, D, MBG, US); Las Palmas, de 1917, Jørgensen 1956 (BA, BG, US); Rio de Oro, Dee. 7, 1892, Niederlein 72 (B); MISIONES: Santa Ana, Sept. 20, 1912, Rodriguez 3588 (BA, MBG); San Ignacio, Sept. 7, 1919, Munies 3586 (BA, MBG); Posadas, Bonpland, in silva ad occid. versus, Jan. 23, us Ekman 1586 (NY, S); same locality, in fruticeto ad praed. **La Granja," 1907, Ekman 1585 (S) ; CORRIENTES: Wald bei Riachuelo, Jan. 19, 1883, de 83 (B). URUGUAY: ARTIGAS: Montes del Cuareim, May, 1901, Arechavaleta 18 (B). 31. Forsteronia tarapotensis K. Sch. in Engl. Bot. Jahrb. 40: 411. 1908, nom. nud Frutiees volubiles; ramulis crassiusculis glabris maturitate conspicue lenticellatis ; foliis oppositis breviter petiolatis lance- olatis oblongo- dccus apice acuminatis basi obtusis 4.5-10.6 em. longis 1.3-3.0 em. latis membranaceis glaberrimis supra nervo medio pauciglanduligero; petiolis 0.3-0.5 em. longis ; in- florescentiis dense subspicato-thyrsiformibus terminalibus folis conspicue brevioribus; pedunculo omnino puberulo; pedicellis 0.04—0.05 em. longis minute puberulis ; bracteis ovatis vel ovato-lanceolatis 0.1-0.2 em. longis scariaceis vel paululo foliaceis; calycis laciniis ovatis acutis acuminatisve 0.18-0.2 em. longis scariaceis extus dense minuteque puberulis squa- mellis minutis multis plus minusve regulariter positis ; corollae tubo ca. 0.15 em. longo basi ea. 0.08-0.1 cm. diametro metiente faucibus ea. 0.175-0.2 em. diametro metientibus extus minute puberulo-papillatis intus villosulis lobis oblongo-ovatis 0.35- (229) [Vor. 22 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN 0.55 em. longis patulis extus omnino denseque puberulis intus villosulis; filamentis staminum 0.25-0.3 em. longis stylum ag- glutinatis antheris 0.2-0.25 em. longis glabris omnino exsertis ; ovario ovoideo apocarpo ca. 0.07 em. longo puberulo-papillato ; stigmate ca. 0.18-0.2 em. longo; stylo ca. 0.075 em. longo; nec- tariis ovario conspicue brevioribus ; folliculis ignotis. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, lanceo- late to oblong-elliptie, apex acuminate, base obtuse, 4.5-10.6 em. long, 1.3-3.0 em. broad, firmly membranaceous, glabrous, above inconspicuously glandular at the base of the midrib; petioles 0.3-0.5 em. long; inflorescenee densely thyrsiform-subspici- form, terminal, conspicuously shorter than the subtending leaves, bearing numerous small, white flowers; peduncle rather densely puberulent, ultimate branches rather indefinitely con- gested, minutely puberulent; pedicels 0.04—0.05 em. long, minutely puberulent; ealyx-lobes ovate to ovate-lanceolate, acute to acuminate, 0.18-0.2 em. long, scarious or only slightly foliaceous, the squamellae minute, numerous, regularly and in- definitely distributed; corolla-tube about 0.15 em. long, about 0.08-0.1 em. in diameter at the base and 0.175-0.2 em. in di- ameter at the orifice, without minutely puberulent-papillate, villosulose within, the lobes oblong-ovate, 0.35-0.55 em. long, densely puberulent without, villosulose within, widely spread- ing; stamen filaments 0.25-0.3 em. long, agglutinated to the style, the anthers 0.2-0.25 em. long, glabrous, wholly exserted; ovary ovoid, apocarpous, about 0.07 cm. long, puberulent- papillate; stigma about 0.18-0.2 em. long; style about 0.075 em. long; nectaries much shorter than the ovary ; follicles unknown. PERU: LORETO: im Buschwald, Tarapoto, Nov., 1902, Ule 6561 (B, TYPE, D, MBG, photograph and analytical drawings); prope Tarapoto, 1855—6, Spruce 4908 (Bx, Camb.); forest, Yurimaguas, Oct. 22, 1929, Williams 3891 (FM); Tarapoto, Dec. 5, 1929, Williams 5611 (FM, MBG). Well distinguished from its immediate relatives by the small foliage and puberulent corollas. Spruce 4908 was referred to F. Pavonu A. DC. (= Apocynum cannabinum L. var. glaber- rimum A. DC.) by Miers (Apoc. So, Am. 243. 1878). (230) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 197 32. Forsteronia myriantha Donn. Sm. Bot. Gaz. 2T: 435. 1899. Stems relatively stout, minutely puberulent to papillate when very young, soon becoming glabrate and conspicuously lenticellate; leaves opposite, shortly petiolate, elliptic to oval, apex acute to acuminate, occasionally obtuse, base obtuse to broadly acute, 4.5-10.0 cm. long, 2.0-4.5 em. broad, membra- naceous, inconspicuously glandular at the base of the midrib above, beneath glabrous, or infrequently rather sparsely and irregularly pilosulose to glabrate; petioles 0.2—0.45 em. long; inflorescence rather densely and broadly thyrsiform, terminal, conspicuously surpassed by the subtending leaves, bearing many small, white flowers; primary peduncle minutely puber- ulent-papillate, rarely densely and conspicuously puberulent, ultimate branches determinate, puberulent-papillate to densely puberulent; pedicels 0.1—0.21 cm. long, puberulent to puber- ulent-papillate; bracts ovate-trigonal, acuminate, 0.1—0.3 em. long, scarious; calyx-lobes ovate, broadly acute to obtuse, 0.1-0.12 cm. long, scarious, puberulent-papillate or rarely tomentulose without, the squamellae minute, many, indefinitely distributed and apparently adnate to the base of the lobes; corolla glabrous to very minutely papillate without, the tube 0.07-0.09 em. long, about 0.05-0.07 em. in diameter at the base and 0.15-0.17 cm. in diameter at the orifice, villosulose within, the lobes 0.25—0.3 em. long, minutely pilosulose within, widely spreading; stamen filaments 0.12-0.18 em. long, agglutinated to the style, the anthers 0.18-0.23 em, long, widely exserted, gla- brous; ovary ovoid, apocarpous, minutely puberulent-papil- late, about 0.03 em. long; stigma 0.08-0.13 em. long; style 0.08-0.12 cm. long; nectaries somewhat shorter than the ovary ; follieles unknown. GUATEMALA: SANTA ROSA: Naranjo, alt. 1100 m., May, 1893, Heyde $ Lux 4533 (B, BB, FM, MBG, NY, US); ZACATEPEQUEZ: Embaulada, alt. 1800 m., Dec., 1889, Heyde $ Lux 4534 (B, BB, M, MBG, US); PETEN: La Libertad, April 6, 1933, Lundell 2540 (FM). CosTA RICA: ALAJUELA: collines de Santiago, prés S. Ramon, alt. 1100-1200 m., April 25, 1901, Brenes 14272 (B, FM, US); CARTAGO: bords du rio de Las Vueltas, Tucurrique, alt. 635 m., May, 1899, Tonduz 13355 (B, D, FM, M, US); SAN JOSE [?]: Rio Hondo, May 8, 1903, Pittier 16622 (B, US). (231) [Vor. 22 198 ANNALS OF THE MISSOURI BOTANICAL GARDEN PANAMA: CANAL ZONE: Gatun Sta., Panama R. R., Febr. 11, 1860, Hayes s.n. (G, MBG, NY). Heyde € Lux 4533 is conspicuous because of the densely puberulent inflorescence, The inflorescence of the species as a whole is densely puberulent-papillate, however, and since but a single specimen exists of the more conspicuously pubescent type, no taxonomie segregation has been attempted, other characters being quite in harmony with those other recorded speeimens from Guatemala, Costa Rica, and Panama. 33. Forsteronia galbina Woodson, spec. nov. Frutieosa volubilis; ramulis erassiuseulis glabris vel juven- tate minutissime papillatis maturitate conspicue lenticellatis ; foliis oppositis petiolatis ovalibus vel obovato-elliptieis apice obtusis vel brevissime acuminatis basi obtusis 4.5-8.0 em. longis 2.7-4.0 em. latis membranaceis supra glabris nervo medio basi inconspicue glanduligero subtus in axillis nervi medii minute barbellatis caeterumque glabris; petiolis 0.3-0.8 em. longis; inflorescentiis latissime thyrsiformibus paene sub- eorymbosis terminalibus folia subaequantibus flores multas minutas galbinas gerentibus; pedunculo ramulisque puberulo- papillatis; pedicellis 0.12-0.15 em. longis minute puberulo- papillatis; bracteis lanceolatis acuminatis 0.05-0.1 em. longis scariaceis; calycis laciniis ovatis acutis acuminatisve 0.1-0.125 cm. longis extus dense puberulo-papillatis squamellis minutis numerosis basi plus minusve connatis; corollae tubo 0.07- 0.1 em. longo basi ca. 0.06-0.07 em. diametro metiente faucibus ea. 0.15-0.175 em. diametro metientibus extus minutissime papillato intus laxe villosulo lobis 0.25 em. longis patulis extus minute puberulis puberulo-papillatisve intus glabriusculis; filamentis staminum ca. 0.09 em. longis stylum agglutinatis antheris 0.19-0.21 cm. longis glabris omnino exsertis; ovario ovoideo apocarpo ca. 0.05 em. longo puberulo-papillato ; stigmate ca. 0.11 cm. longo; stylo ca. 0.09 em. longo; nectariis ovarium paene aequantibus ; follieulis desiderantibus, Stems relatively stout, glabrous or minutely papillate when very young, soon becoming glabrate and conspicuously lenti- eellate when fully mature; leaves opposite, petiolate, oval to (232) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 199 obovate-elliptie, apex obtuse to very shortly acuminate, base obtuse, 4.5-8.0 em. long, 2.7-4.0 em. broad, membranaceous, above glabrous, inconspicuously glandular at the base of the midrib, beneath minutely barbellate in the axils of the midrib, otherwise glabrous; petioles 0.3-0.8 cm. long; inflorescence very broadly thyrsiform, almost subeorymbose, terminal, somewhat shorter than or about equalling the subtending leaves, bearing many small, yellowish-green flowers; peduncle puberulent-papillate, ultimate branches determinate, minutely puberulent-papillate; pedicels 0.12-0.15 em. long, minutely puberulent-papillate; bracts lanceolate, acuminate, 0.05-0.1 em. long, scarious; calyx-lobes ovate, acute to acuminate, 0.1— 0.125 cm. long, scarious, densely puberulent-papillate without, the squamellae minute, numerous, in groups more or less con- nate at the base; corolla-tube 0.07—0.1 em. long, about 0.06-0.07 cm. in diameter at the base and 0.15-0.175 em. in diameter at the orifice, very minutely papillate without, laxly villosulose within, the lobes about 0.25 cm. long, widely spreading, mi- nutely puberulent-papillate or puberulent without, essentially glabrous within; stamen filaments about 0.09 em. long, ag- glutinated to the style, the anthers 0.19-0.21 em. long, wholly exserted, glabrous; ovary apocarpous, ovoid, about 0.05 em. long, puberulent-papillate; stigma about 0.11 em. long; style about 0.09 em. long; nectaries almost equalling the ovary ; fol- licles unknown, PERU: LORETO: Flutfreier Hochwald, Mündung d. Santiago, alt. 160 m., Sept. 24, 1924, Tessmann 3967 (B, TYPE, D, MBG, photograph and analytieal drawings). This species is perhaps too closely related to the foregoing, of which it might be considered a variety. The geographical isolation, together with the rather slight morphological distine- tions, however, has prompted its interpretation as a specific entity. Fruit of both F. myriantha and F. galbina is awaited with interest. 34. Forsteronia thyrsoidea (Vell.) Muell.-Arg. in Mart. Fl. Bras. 6': 105. 1860; Miers, Apoc. So. Am. 247. 1878. Stems relatively stout, more or less densely pilose or puber- (233) [Vor. 22 200 ANNALS OF THE MISSOURI BOTANICAL GARDEN ulent when young, eventually becoming glabrate and conspicu- ously lenticellate when fully mature; leaves opposite, petiolate, oblong-elliptie to broadly oval, apex acuminate to acute or obtuse, base broadly obtuse to rounded, 4-12 cm. long, 1.5—5.0 cm. broad, firmly membranaceous, above minutely pilose to gla- brous, inconspicuously glandular at the base of the midrib, beneath more or less densely puberulent generally, or barbel- late in the axils of the midrib and otherwise glabrous; petioles 0.3-0.9 em. long, indument as upon the stem; inflorescence rather narrowly and densely thyrsiform, terminal, some- what shorter than the subtending leaves, bearing many small, greenish-yellow flowers; primary peduncle puberulent, ultimate branches determinate, puberulent to puberulent- papillate; pedicels 0.05-0.1 em. long, puberulent to puberulent- papillate; bracts lanceolate to linear-lanceolate, 0.4—1.0 cm. long, eonspicuously foliaceous ; calyx-lobes lanceolate to ovate- lanceolate, long-acuminate, 0.1-0.3 em. long, minutely puber- ulent-papillate without, conspicuously foliaceous, the alternate squamellae solitary or infrequently paired; corolla-tube 0.13- 0.2 em. long, about 0.07—0.125 em. in diameter at the base and 0.15-0.2 em. in diameter at the orifice, essentially glabrous without, villosulose within, the lobes oblong to oblong-ovate, 0.25-0.38 em. long, widely spreading, papillate without, mi- nutely villosulose at the base within; stamen filaments 0.18-0.3 em. long, agglutinated to the style, the anthers 0.2-0.28 em. long, glabrous, wholly exserted; ovary ovoid, apocarpous, about 0.05 em. long, papillate; stigma 0.11-0.15 em. long; style 0.1-0.25 em. long; neetaries somewhat shorter than the ovary. Var. typica. Echites thyrsoidea Vell. Fl. Flum. 111. 1850; Icones 3: pl. 37. 1827, except the fruit. Forsteronia multinervia A. DC. in DC, Prodr. 8: 437. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 104. 1860. Forsteronia multinervia A. DC. 5 ovalifolia Muell.-Arg. loc. cit. 105. 1860. Thyrsanthus multinervius (A. DC.) Miers, Apoc. So. Am. 103. 1878. (234) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 201 Stems conspicuously pubescent when young, eventually be- coming glabrate; leaves rather inconspicuously and irregularly pubescent to glabrate above, beneath more or less puberulent generally; follicles relatively stout, continuous or essentially so, sharply divaricate, 13-25 cm. long, glabrous or essentially so; seeds 1.0-1.6 cm. long, the tawny coma 3.2-4.7 em. long; other characters as in the species. BRAZIL: MINAS GERAES: Lagóa Santa, date lacking, Warming s.m. (C, NY); RIO DE JANEIRO: in fruticetis, collibus, pr. civ. R. Jan., Febr., 1829, Lhotzky 55 (B); ad urbem, in silvis inter Gavea et Vista rp p 15, 1929, Ducke 21810 (US); locality lacking, 1865, Allemáo s.n. (DC); s ULO: ad ripas Rio Pardo, eg ae 1917, Frazäo 8674 (B, U); Ilha da E Grande, Nov. 6, 1920, Gehrt 4 5 (B) data incomplete, Dec. 5, 1875, Mosen 4271 (S); RIO Mon DO SUL: Ed dos Duridos, Munieipio Rio e Nov., 1928, Jürgens 29 (B); DATA INCOMPLETE: Riedel s.n. (B, BB, G, Uys etos 17138 (B, Bx, C); Luschnath s.m. (C); Sellow 635 (B). PARAGUAY: dans les bois, Paraguari, Oct., 1874, Balansa 1370b (D, DC); Grande Pieada de Caaguazu, April, 1876, Balansa "18704 (BB); in regione fluminis Alto Parana, 1909-10, Fiebrig 6267 (B); in silva pr. Cordillera de Altos, Dec., 1885, Hassler 1577 (BB); same locality, Nov., year lacking, Hassler 3494 (B RGENTINA: MISIONES: Santa Ana, Sept. 19, 1912, Rodriguez 605 (B A, MBG); Piearda a San Pedro, Oct. 27, 1886, Niederlein 2219 (B); data incomplete, Oct. 8, 1896, Bettfreund 943 (B). A very distinctive species because of the narrow inflores- cence with conspicuous, foliaceous bracts. There appears to be little room for doubt that our specimens correspond to Vel- lozo's illustration with the exception of the three pairs of sub- parallel, moniliform follicles. Since the combination of such an inflorescence and foliage with such follicles, which closely resemble those of F. pubescens or F. glabrescens, is unknown in the flora as represented in our herbaria, the assumption appears to be justified that the illustration is an accidental composite. Var. glabriuscula (A. DC.) Woodson, comb. nov. Forsteronia mutinervia A. DC. B glabriuscula A. DC. in DC. Prodr. 8: 437. 1844; Muell.-Arg. in Mart. Fl. Bras. 61: 105. 1860. Forsteronia obscura Rusby, Mem. Torrey Bot. Club 4: 2 (235) [Vor. 22 202 ANNALS OF THE MISSOURI BOTANICAL GARDEN Stems minutely and inconspicuously puberulent when very young, soon becoming glabrate;leaves glabrous above, beneath rather ineonspieuously barbellate in the axils of the midrib; follieles unknown; other characters as in the species. BRAZIL: RIO DE JANEIRO: inter Cabo Frio et Campos, 1829, V idensis s.n. (Bx); data incomplete, Glaziou 4092 (B); SANTA CATHARINA: an der Garciastrasse bei Blumenau, Nov., 1888, Ule 1161 (B). Borrvi: ““Songo,”” Nov., 1890, Bang 855 (B, BB, D, FM, M, MBG, NY, US). It appears possible that this entity may represent an arti- fieial eolleetion of spontaneous variations assuming a like aspect independently. Although such a view may be supported by the rather incongruent distribution of the few specimens cited in the foregoing paragraphs, the plants themselves are remarkably similar, and conspicuous intergradation to the typical variety is lacking as yet. 35. Forsteronia Velloziana (A. DC.) Woodson, Ann. Mo. Bot. Gard. 21: 622. 1934. Echites bracteata Vell. Fl. Flum. 112. 1830; Icones 3: pl. 41. 1827, not HBK. Echites Velloziana A. DC. in DC. Prodr. 8: 474. 1844. Forsteroma multinervia A. DC. y microphylla Muell.-Arg. in Mart, Fl. Bras. 6': 105. 1860, Forsteronia ? bracteata (Vell.) Muell.-Arg. loc. cit. 106. 1860. Thyrsanthus bracteatus (Vell.) Miers, Apoc, So. Am. 102. 1878 Forsteronia microphylla (Muell.-Arg.) Handel-Mzt. Denkschr. K.K. Akad. Wiss. Wien 79: 388. 1931 (re- print, 12. 1910). Stems relatively stout, conspicuously ferruginous-hirsutu- lose when young, eventually becoming glabrate and conspicu- ously lenticellate when fully mature; leaves opposite, very shortly petiolate, nearly subsessile, oblong-elliptie to broadly oval, apex shortly and abruptly acuminate to obtuse, base broadly obtuse or rounded, oceasionally almost subcordate, 2.5-7.0 em. long, 1-4 em. broad, firmly membranaceous, above rather sparsely and minutely hirtellous to glabrate, incon- (236) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 203 spicuously glandular at the base of the midrib, beneath finely and generally puberulent ; petioles 0.15-0.35 em. long, indument as upon the stem; inflorescence densely subeorymbose-thyrsi- form, terminal, conspicuously shorter than the subtending leaves, bearing few to numerous small, white or greenish flowers; peduncle hirtellous throughout; pedicels 0.08-0.1 em. long, minutely hirtellous ; bracts lanceolate to linear-lanceolate, somewhat foliaceous, 0.1-0.8 em. long; calyx-lobes lanceolate, acuminate, 0.25—0.4 cm. long, minutely puberulent without, the alternate squamellae solitary or infrequently paired; corolla- tube 0.1-0.12 em. long, minutely papillate or essentially gla- brous without, puberulent within, about 0.1-0.13 em. in di- ameter at the base and 0.15-0.23 em. in diameter at the ori- fice, the lobes oblong-ovate, 0.3-0.4 em. long, only slightly spreading, puberulent-papillate without, papillate within; sta- men filaments 0.1-0.15 em. long, agglutinated to the style, the anthers 0.23-0.29 cm. long, glabrous, wholly exserted; ovary apocarpous, ovoid, about 0.45 em. long, minutely puber- ulent-papillate; stigma 0.12-0.17 em. long; style about 0.1 em. long ; nectaries somewhat shorter than the ovary; follicles rela- tively short and stout, 5-10 em. long, glabrous, or sparsely and minutely hirtellous when immature, divaricate; seeds 0.6-0.8 em. long, the tawny coma 2.3-2.6 em. long. BRAZIL: RIO DE JANEIRO: data ineomplete, Glaziou 7757 (B); MINAS GERAES: Caete, Nov., 1915, Hoehne 6623 (B); S. de Antonio Pereira, Oct. 27, 1892, Schwacke 8754 (B); in virgultis, Ouro Preto, Oct. 20, 1895, Schwacke 11854 (B); data incomplete: Widgren 52 (Bx, 8); Widgren 51 (B, S); Regnell III 877 (B, FM, 8, US); são PAULO: Botucatu, Nov., 1896, Novaes 384 (US); Rio Pequeno, Butantan, Oct. 28, 1918, Hoehne 2549 (B); Morro-Pellado, Ityrapina, Jan., 1901, Edwall 11147 (B); PARANA: ad marg. silvulae, J aguariahyva, Nov. 5, 1910, Dusen 10366 (D, FM, MBG, NY, US); DATA INCOMPLETE: Sellow 36 (B). There can be no question that this entity is entitled to full specific rank. Its characteristic inflorescence, virtually erect corolla-lobes, vegetative indument, and short follicles are quite uniform throughout the specimens examined. 36. Forsteronia chiriquensis Woodson, spec. nov. Fruticosa volubilis; ramulis sat erassiusculis juventate laxe ferrugineo-hirtellis maturitate glabratis conspicue lenticel- (237) [Vor. 22 204 ANNALS OF THE MISSOURI BOTANICAL GARDEN latis; foliis oblongo- vel obovato-ellipticis apice abrupte breviterque subeaudato-acuminatis basi late obtusis 7-10 cm. longis 3.2-3.8 em. latis membranaceis supra glabris glabratisve nervo medio basi pauciglanduligero subtus in axillis nervi medii inconspicue barbellatis; petiolis 0.3-0.5 em. longis ut in ramulis vestitis; inflorescentiis subspiciformi-thyrsiformibus terminalibus folia subaequantibus flores multas viridi-albidas (?) gerentibus ; peduneulo hirtello ramulis vix manifestis glom- eratis; pedicellis vix manifestis subnullis; bracteis lanceolatis vel ovato-lanceolatis 0.1-0.45 em. longis subfoliaceis; calycis laciniis ovato-lanceolatis acuminatis 0.25-0.3 em. longis minute irregulariterque puberulo-papillatis squamellis alternatis 1-2; corollae tubo 0.1 em. longo basi ea. 0.05 em. diametro metiente faucibus ca. 0.175 em, diametro metientibus extus glabriusculo intus villosulo lobis oblongis 0.28-0.3 em. longis patulis extus minute papillatis intus minute puberulo-papillatis; filamentis staminum 0.17-0.2 em. longis stylum agglutinatis antheris ca. 0.25 em. longis omnino exsertis glabris; ovario apocarpo ovoideo ea. 0.06 em. longo minute hirtello; stigmate 0.15 em. longo; stylo 0.18 em. longo; nectariis ovario brevioribus; fol- liculis ignotis. Stems relatively stout, rather laxly ferruginous-hirtellous when young, eventually becoming glabrate and conspicuously lenticellate when fully mature; leaves opposite, shortly peti- olate, oblong- to obovate-elliptie, apex shortly and abruptly subcaudate-acuminate, base broadly obtuse, 7-10 em. long, 3.2-3.8 em. broad, membranaceous, above glabrous or glabrate, glandular at the base of the midrib, beneath inconspicuously barbellate in the axils of the midrib; petioles 0.3-0.5 em. long, indument as upon the stem; inflorescence subspiciform-thyrsi- form, terminal, about equalling the subtending leaves, bearing numerous small, greenish-white (?), subsessile flowers; pe- dunele hirtellous, the ultimate branches extremely reduced and indefinitely manifest; pedicels scarcely manifest; bracts lan- ceolate to ovate-lanceolate, 0.1—0.45 em. long, subfoliaceous ; calyx-lobes ovate-lanceolate, acuminate, 0.25-0.3 em. long, mi- nutely and irregularly puberulent-papillate without, the alter- (238) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 205 nate squamellae solitary or paired; corolla-tube about 0.1 em. long, about 0.05 em. in diameter at the base and 0.175 em. in diameter at the orifice, essentially glabrous without, villosulose within, the lobes oblong, 0.28-0.3 em. long, widely spreading, minutely papillate without, minutely puberulent-papillate within; stamen filaments 0.17-0.2 em. long, agglutinated to the style, the anthers about 0.25 em. long, wholly exserted, gla- brous; ovary apocarpous, ovoid, about 0.06 em. long, minutely hirtellous; stigma 0.15 em. long; style 0.18 cm. long; nectaries somewhat shorter than the ovary ; follicles unknown. PANAMA: CHIRIQUI: data incomplete, May, 1858, Wagner s.n. (M, TYPE, MBG, photograph and analytieal drawings). This species suggests F. thyrsoidea of southeastern South America, but differs notably in the subspiciform construction of the inflorescence. Additional material is greatly to be desired. 37. Forsteronia Pycnothyrsus K. Sch. in herb. Frutieosa volubilis; ramulis sat crassis juventate dense laxeque ferrugineo-hirtellis maturitate glabratis conspicue lentieellatis; foliis oppositis breviter petiolatis oblongo- ovalibus apice acutis obtusisve basi late obtusis rotundatisve 4.0-4.5 em. longis 1.8-2.2 cm. latis membranaceis supra sparse minuteque hirtellis nervo medio basi inconspicue glanduligero subtus dense puberulis ; petiolis 0.35-0.4 em. longis ut in ramulo vestitis; inflorescentiis subspiciformi-thyrsiformibus termi- nalibus folia conspieue superantibus flores multas parvas viridi-albidas (?) gerentibus; pedunculo minute ferrugineo- hirtello ramulis glomeratis vix manifestis ; pedicellis 0.03-0.05 em. longis minute puberulis; bracteis ovatis 0.1-0.2 em. longis scariaceis; calycis laciniis late ovatis obtusis rotundatisve 0.13-0.15 em. longis scariaceis extus dense minuteque puberulis squamellis alternatis solitariis; corollae tubo 0.1 em. longo basi ca. 0.07 em. diametro metiente faucibus ca. 0.15 em. diametro metientibus extus glabriusculo vel indistincte papillato intus villosulo lobis oblongis 0.25 em. longis patulis extus puberulo- papillatis intus dense pilosulis; filamentis staminum 0.18 cm. (239) [Vor. 22 206 ANNALS OF THE MISSOURI BOTANICAL GARDEN longis antheris ca. 0.2 em. longis omnino exsertis glabris ; ovario apocarpo ovoideo ca. 0.035 em, longo minute hirtello; stigmate 0.14-0.15 em. longo ; stylo 0.1 em. longo; nectariis ovario brevi- oribus ; folliculis ignotis, Stems relatively stout, densely and laxly ferruginous- hirtellous when young, eventually becoming glabrate and con- spicuously lenticellate when fully mature; leaves opposite, petiolate, oblong-oval, apex acute to obtuse, base broadly obtuse or rounded, 4.0—4.5 cm. long, 1.8-2.2 em. broad, mem- branaceous, above sparsely and minutely hirtellous, inconspic- uously glandular at the base of the midrib, beneath densely puberulent; petioles 0.35-0.4 em, long, indument as upon the stem; inflorescence subspiciform-thyrsiform, terminal, con- spicuously surpassing the subtending leaves, bearing many small, greenish-white (?) flowers; primary peduncle minutely ferruginous-hirtellous, the ultimate branches glomerate, scarcely manifest; pedicels 0.03-0.05 em. long, minutely puber- ulent; bracts ovate, 0.1-0.2 em. long, scarious; calyx-lobes broadly ovate, obtuse to rounded, 0.13-0.15 em. long, densely and minutely puberulent without, the alternate squamellae soli- tary; corolla-tube 0.1 em. long, about 0.07 em. in diameter at the base and 0.15 em. in diameter at the orifice, minutely and indistinctly papillate or essentially glabrous without, vil- losulose within, the lobes oblong, 0.25 em. long, widely spread- ‘ing, puberulent-papillate without, densely pilosulose within; stamen filaments 0.18 em. long, the anthers about 0.2 em. long, wholly exserted, glabrous; ovary apocarpous, ovoid, about 0.035 em. long, minutely hirtellous; stigma 0.14—0.15 em. long; style 0.1 em. long; nectaries somewhat shorter than the ovary ; follicles unknown. CoLoMBIA or ECUADOR: data incomplete, Lehmann 7885 (B, TYPE, FM, MBG, photograph and analytical drawings). Closely related to the preceding, from which it may be dis- tinguished by its smaller foliage, smaller, ovate calyx-lobes, and inconspicuous, searious bracts. It is guessed that Ecuador may be the true provenience of this species, as the apocyna- ceous flora of upper Panama and Ecuador have shown several (240) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE. IV 207 notable affinities. Additional specimens and reliable data are greatly to be desired for this species, 38. Forsteronia adenobasis Muell-Arg. Linnaea 30: 412. 1860. Thyrsanthus adenobasis (Muell.-Arg.) Miers, Apoc. So. Am. 96. 1878. Stems relatively stout, minutely ferruginous-tomentulose when very young, soon becoming glabrate and conspicuously lenticellate when fully mature; leaves opposite, petiolate, broadly oval, apex very abruptly and shortly acuminate to ob- tuse, base obscurely and broadly cordate, 5.0-10.5 em. long, 2.8-7.0 em. broad, firmly membranaceous, above glabrous or minutely and very inconspicuously pubescent along the veins, glandular at the base of the midrib, beneath minutely and densely tomentulose; petioles 0.8-1.0 em. long; inflorescence rather narrowly pyramidal-thyrsiform, terminal, about equal- ling to conspicuously surpassing the subtending leaves, bearing many small, white flowers; primary peduncle minutely ferru- ginous-tomentulose, ultimate branches determinate, minutely tomentulose; pedicels 0.13-0,2 em. long, minutely tomentulose; braets ovate, 0.1-0.25 cm. long, searious; calyx-lobes ovate, acute, 0.1-0.2 em. long, densely and minutely puberulent-papil- late without, the alternate squamellae very minute, solitary; corolla-tube 0.1 em. long, about 0.07-0.1 em. in diameter at the base and 0.15-0.2 cm. in diameter at the orifice, without mi- nutely and irregularly puberulent-papillate, essentially gla- brous toward the base, within minutely villosulose, the lobes ovate-oblong, 0.25-0.3 em. long, somewhat spreading, densely puberulent-papillate without, densely pilosulose within; sta- men filaments 0.12-0.15 em. long, agglutinated to the style above, the anthers 0.17-0.19 em. long, wholly exserted, gla- brous; ovary apocarpous, ovoid, about 0.03 em. long, minutely hirtellous; stigma 0.15-0.17 cm. long; style 0.05—0.08 em. long; nectaries about equalling the ovary, minutely pilosulose at the tips; follieles unknown. BRITISH GUIANA: ad ripas fl. Pomeroon, Sept., 1843, Schomburgk 1438 (B, TYPE, MBG, photograph and analytieal drawings); in mixed forest, Moraballi Creek, (241) [Vor. 22 208 ANNALS OF THE MISSOURI BOTANICAL GARDEN near Bartica, alt. near sea-level, Oct. 10, 1929, Sandwith 416 (K, NY, S, U, US); ““Roraima,”* data incomplete, 1842-3, Schomburgk 707 (B, BB, D, COTYPES). 39. Forsteronia umbellata (Aubl.) Woodson, comb. nov. Apocynum umbellatum Aubl. Hist. Pl. Gui. Fr. 1: 275; 3: pl. 108. 1775. Thenardia umbellata (Aubl.) Spreng. Syst. 1: 636. 1825. Thenardia (?) corymbosa Benth, in Hook. Jour. Bot. 3: 246. 1841. Forsteronia Schomburgku A. DC. in DC. Prodr. 8: 438. 1860, not Muell.-Arg. Forsteronia Schomburgkü A. DC. P umbellata (Aubl.) A. DC. loc. cit. 1860, Forsteronia macrophylla Muell.-Arg. Linnaea 30: 411. 1860. Thyrsanthus macrophyllus (Muell.-Arg.) Miers, Apoc, So. Am. 96. 1878. Thyrsanthus Aubletianus Miers, loc. cit. 98. 1878. Thyrsanthus corymbiferus Miers, loc. cit, 1878. Stems relatively stout, minutely and inconspicuously puber- ulent when very young, soon becoming glabrate and conspicu- ously lenticellate when fully mature; leaves opposite, petiolate, broadly oval to oblong-oval, apex very abruptly and shortly acuminate-subcuspidate to obtuse, base rounded, frequently obscurely cordate, 7.8-15.0 em. long, 4.5-9.2 em, broad, firmly membranaceous, above glabrous, or minutely and indefinitely puberulent along the veins, glandular at the base of the midrib, beneath very minutely and generally puberulent or tomentu- lose; petioles 0.6-1.3 em. long; inflorescence broadly sub- corymbose-thyrsiform, terminal, shorter than the subtending leaves, bearing many small, white flowers; primary peduncle puberulent-papillate, ultimate branches determinate, puber- ulent-papillate throughout ; pedicels 0.2—0.3 em. long, minutely puberulent; bracts ovate, 0.1-0.25 em. long, scarious or only slightly foliaceous; calyx-lobes ovate, acuminate, scarious or only slightly foliaceous, densely and minutely puberulent with- out, 0.15-0.17 em. long, the alternate squamellae solitary, very minute; corolla-tube 0.125-0.15 em. long, about 0.07-0.1 em. in diameter at the base and 0.15-0.2 em, in diameter at the ori- (242) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 209 fiee, irregularly and minutely puberulent-papillate without, villosulose within, the lobes broadly oblong, somewhat spread- ing, 0.3-0.35 em. long, densely and minutely puberulent-papil- late without, within densely and minutely pilosulose; stamen filaments 0.15-0.25 cm. long, the anthers 0.25 em. long, barbel- late at the tips, wholly exserted; ovary apocarpous, ovoid, about 0.07 em. long, minutely hirtellous; stigma 0.18-0.2 cm. long; style 0.07-0.15 em. long; nectaries nearly as long as the ovary, minutely barbellate; follicles unknown. FRENCH GUIANA: Cayenne, date lacking, collector unknown (DC, MBG, photo- graph and analytical drawings). BRAZIL: PARA: silva paludosa, Belem do Para, Oct. 31, 1926, Ducke 21598 (B, U, US); silva non inundata, Braganca, Jan. 14, 1923, Ducke 17482 (B). The specimens cited above agree remarkably well with Aublet's illustration, partieularly that from French Guiana, the type specimen of F. macrophylla Muell.-Arg. 40. Forsteronia Sandwithiana Woodson, spec. nov. Frutieosa volubilis; ramulis sat crassiusculis, juventate minute inconspicueque ferrugineo-tomentulosis mox glabratis maturitate conspicue lenticellatis; foliis oppositis breviter petiolatis oblongo-elliptieis vel ovalibus apice abrupte brevis- simeque acuminatis basi obseure cordatis 4.5-10.0 em. longis 1.6-5.8 cm. latis firme membranaceis subcoriaceisve supra glabris nervo medio basi pauciglanduligero subtus nervo medio venisque minutissime puberulo-papillatis caeterumque glabris ; petiolis 0.4-0.9 em. longis minute puberulo-papillatis; inflo- rescentiis dense subcorymboso-thyrsiformibus terminalibus foliis multo brevioribus flores multas parvas virides gerenti- bus; pedunculo minutissime puberulo-papillato; pedicellis 0.12-0.2 cm. longis minute puberulo-papillatis ; bracteis ovatis 0.1-0.15 em. longis scariaceis; calycis laciniis lanceolatis acu- minatis 0.3-0.35 em. longis subfoliaceis extus minute puberulo- papillatis squamellis alternatis minutissimis solitariis; corol- lae tubo 0.13-0.15 em. longo basi ca. 0.1 em. diametro metiente faucibus ca. 0.13 cm. diametro metientibus extus minute papil- latis intus lanulosis lobis late oblongis 0.4—0.42 em. longis vix patulis extus puberulo-papillatis intus conspicue pilosulosis; (243) [Vor. 22 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN filamentis staminum 0.2-0.25 cm. longis antheris ca. 0.3 cm. longis omnino exsertis apice minute sparseque barbellatis; ovario apocarpo ovoideo ca. 0.05 em. longo minute hirtello; stigmate 0.2 cm. longo; stylo 0.15-0.2 cm. longo; nectariis ovarium vix aequantibus glabris; folliculis ignotis. Stems relatively stout, minutely and inconspicuously fer- ruginous-tomentulose when very young, soon becoming gla- brate and conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptie to oval, apex abruptly and very shortly acuminate, base obseurely cordate, 4.5-10.0 em. long, 1.6-5.8 em. broad, firmly membranaceous to subeoriaceous, above glabrous or essentially so, inconspic- uously glandular at the base of the midrib, beneath very minutely puberulent upon the veins and midrib, otherwise gla- brous ; petioles 0.4-0.9 em, long, minutely puberulent-papillate ; inflorescence densely subeorymbose-thyrsiform, terminal, much shorter than the subtending leaves, bearing many small, green flowers; primary peduncle very minutely puberulent-papillate, ultimate branches determinate, minutely puberulent-papillate ; pedicels 0.12-0.2 cm. long, minutely puberulent-papillate ; bracts ovate, 0.1-0.15 em. long, searious; calyx-lobes lanceo- late, long-acuminate, 0.3-0.35 em, long, subfoliaceous, minutely puberulent-papillate without, the squamellae very minute, alternate, solitary; corolla-tube 0.13-0.15 em. long, about 0.1 cm. in diameter at the base and 0.13 em. in diameter at the ori- fice, minutely papillate without, within lanulose, the lobes broadly oblong, 0.4—0.42 cm. long, slightly spreading, puber- ulent-papillate without, within conspicuously and densely pilosulose; stamen filaments 0.2-0.25 em. long, agglutinated to the style, the anthers 0.3 em. long, wholly exserted, minutely and sparsely barbellate at the tips; ovary apocarpous, ovoid, about 0.05 em. long, minutely hirtellous; stigma 0.2 em. long; style 0.15-0.2 em. long; nectaries somewhat shorter than the ovary, glabrous; follicles unknown. BRITISH GUIANA: in mora forest, Moraballi Creek, near Bartica, alt. near sea- level, Sept. 24, 1929, Sandwith 334 (K, NY, TYPE, U, MBG, photograph and analyti- cal drawings); data incomplete, Jenman 6316 (K, NY). (244) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 211 F. umbellata, F. adenobasis, and F. Sandwithiana form an extremely natural and closely knit group of species centering in the Guianas and northeastern Brazil. The group is dis- tinguished from the neighboring species of Forsteronia having stamens wholly exserted by the larger leaves of somewhat firmer texture, the slightly spreading corolla-lobes, and the somewhat slighter tendency for the staminal filaments to be- come agglutinated to the style. As a matter of fact, the ag- glutination may not actually exist in F. umbellata, as the herbarium specimens available for dissection have been incon- clusive in this regard. In F. adenobasis agglutination of the staminal filaments occurs only toward the apex of the style. F. Sandwithiana is distinguished from its immediately related congeners by means of its practically glabrous foliage, small, densely subcorymbose-thyrsiform inflorescence, and relatively long, subfoliaceous calyx-lobes. Mr. Sandwith describes the aspect of the liana as follows: **Bush-rope in mora forest. Fl. green, but hairs in the throat and at tops of anthers are white. ?” 41. Forsteronia spicata (Jacq.) G. F. W. Meyer, Fl. Esseq. 135. 1818; A. DC. in DC. Prodr. 8: 437. 1844. Echites spicata Jacq. Enum. Pl. Carib. 13. 1760; Select. Stirp. Am. Hist. 1: 34;2: pl. 29. 1763. Parsonsia spicata (Jacq.) R. Br. Mem. Wern. Soc. 1: 65. 1811. Thyrsanthus corylifolia Griseb. Mem. Amer. Acad. N. $. 8: 519. 1863. Forsteronia corylifolia Griseb. Cat. Pl, Cub. 172. 1866. Thyrsanthus spicatus (Jacq.) Miers, Apoc. So. Am. 95. 1878. Aptotheca corylifolia (Griseb.) Miers, loc. cit. 150, pl. 21B. 1878. Stems relatively stout, minutely ferruginous-tomentulose when young, eventually becoming glabrate and conspicuously lenticellate when fully mature; leaves opposite, shortly peti- olate, broadly oval to obovate-elliptic, apex very abruptly and shorty subcaudate-acuminate, infrequently to acute or obtuse (245) [VoL. 22 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN upon the lower, base broadly obtuse or rounded, 6-16 em. long, 3.5-9.0 em. broad, firmly membranaceous, above very sparsely and minutely pilosulose, more densely so along the veins, oc- easionally glabrate, inconspicuously glandular at the base of the midrib, beneath minutely and rather sparsely tomentulose, rarely glabrate; petioles 0.4—1.0 em. long, indument as upon the stem; inflorescence subspiciform-thyrsiform, both terminal and lateral at the upper nodes, much shorter than the subtend- ing leaves, bearing numerous small, white flowers; primary pedunele minutely ferruginous-tomentulose, ultimate branches extremely reduced; pedicels usually less than 0.05 em. long, scarcely manifest; bracts ovate, 0.1-0.5 em. long, somewhat foliaceous ; calyx-lobes ovate, acute to acuminate, 0.25-0.4 em. long, densely and minutely tomentulose without, slightly foliaceous, the alternate squamellae solitary, or rarely in groups of 2-3; eorolla-tube 0.15-0.21 em. long, about 0.1-0.12 cm. in diameter at the base, 0.2-0.25 em. in diameter at the orifice, without essentially glabrous or minutely papillate above, villosulose within, the lobes oblong-ovate, 0.35—0.4 em. long, widely spreading, puberulent-papillate without, densely pilosulose within; stamen filaments 0.25-0.3 em. long, agglu- tinated to the style, the anthers 0.23-0.28 em. long, widely ex- serted, glabrous; ovary synearpous, ovoid, about 0.065 em. long, densely tomentulose; stigma 0.12-0.16 em. long; style 0.16-0.23 em. long; nectaries much shorter than the ovary; follieles syncarpous, relatively stout and rigid, continuous, 12-19 em, long, minutely and densely ferruginous-tomentulose when immature, eventually becoming glabrate; seeds 0.9—1.0 em. long, the brilliantly tawny coma 3.1-4.5 em. long. CUBA: HAVANA: near mangrove swamps at Boea Ciega between Tarara and Guanabo, in thickets a little farther inland, Oct. 15, 1921, Ekman 13334 (B, S); Tarapaste, Lomas de la Jaula, in frut. scand., es 11, 1914, Ekman 1297 (B, 8); MATANZAS: near mouth of the Bueyvaca, dou , 1903, Britton $ Wilson 10 (NY); SANTA CLARA: climbing on trees, ataco to bees, La Bisita, alt. 500—800 ft., July 9, 1929, pe 7397 (AA, FM, NY, S, US); elimbing on bushes and trees, bog pasture, Limones, Soledad, Fi. July 16, 1930, Jack 8013 (AA, FM, S, US); climbing on trees, Guabairo, Soledad, Cienfuegos, Nov. 26, 1928, Jack 6730 (AA, FM, 8) ; ad Pozo Azul, alt. 150 m., Ane, 1889, agora 5347 (B); ORIENTE: in pascuis, vulgar, Santiago (ad Santiago Bay), Oct. 21, 1916, Ekman 8020 (S); in dumetis ad Rio Jagua, Bayate, July 10, 1915, Ekman 6221 (S); in (246) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 213 sylva, Sabanaso (prope Mir ad epu dapes unge 26, 1915, Ekman 6514 (8); ad Rio Grande prope Sevilla, J ia^ 1; , Ekm 314 (S); DATA INCOMPLETE: Wright 1664 (B, BB, Bx, Camb., D, d Po) NY, E US) MEXICO: GUERRERO: xcci, date laeking, Henke s.n. (M); Llanos de Temal- huacan, alt. 50 m., July 31, 1898, Langlassé 261 (B, D, US); CHIAPAS: banks of ereek near Monserrat July, 1925, Purpus 1034? (NY). GUATEMALA: ZACAPA: Gualan, alt. 620 ft., June 20, 1909, Deam 6368 (FM, G, MBG, NY, US NICARAGUA: MANAGUA: exact loeality lacking, Oct. 8, 1927, Chaves 313 (FM, US). SAN SALVADOR: SANTA ANA: exact locality lacking, 1924, Calderon 2176 er COLOMBIA: MAGDALENA: Santa Marta, alt. 100 ft., May, 1898-1901, H. H. Sm $84 (B, Bx, D, FM, G, MBG, NY, S, U, US); ATLANTICO: Barranquilla, June, sr Elias 222 (US); DATA INCOMPLETE: (Rio Magdal., April, 1801,’’ Humboldt $ Bonpland 1296 (B); ‘‘Nova Granata,’’ date ih Triana 1920 (B). This species breaks down the carpological distinetion be- tween the American Forsteronia and the Asiatie Parsonsia. The affinity of F. spicata is distinctly with the remaining American species, however. It appears probable that the only absolute morphological distinetion of Forsteroma and Par- sonsia capable of superficial detection may lie in the presence or absence of the dorsal foliar foveae, The use of this char- acter among certain genera of Apocynaceae indigenous to the Old and the New Worlds is intended for publication in the near future. 42. Forsteronia simulans Woodson, Ann. Mo. Bot. Gard. 21: 621. 1934. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong- elliptie, apex acute, base obtuse to somewhat cuneate, 2-5 cm. long, 0.7-2.2 em. broad, coriaceous or subcoriaceous, glabrous throughout, above inconspieuously glandular at the base of the midrib, beneath paler, the secondary venation subhorizontal; petioles 0.3-0.5 em. long; inflorescence corymbose-subumbel- late, evidently simple (or very obscurely compound ?), ter- minal, about equalling the subtending leaves, bearing numerous small, yellowish-green flowers; peduncle minutely papillate; pedicels 0.5-0.6 cm. long, minutely papillate; bracts ovate- lanceolate, 0.15-0.2 em. long, slightly foliaceous; calyx-lobes ovate, obtuse, 0.15-0.2 cm. long, glabrous or very minutely (247) [Vor. 22 214 ANNALS OF THE MISSOURI BOTANICAL GARDEN papillate without, the alternate squamellae solitary; corolla- tube about 0.25 em. long, about 0.125 em. in diameter at the base, about 0.15 em. in diameter at the orifice, essentially gla- brous without, minutely puberulent within, the lobes oblong- ovate, obtuse, about 0.2 em. long, spreading, essentially gla- brous without and within, the margins minutely and rather sparsely ciliolate; staminal filaments about 0.15 em. long, ag- glutinated to the style; anthers about 0.2 em. long, wholly ex- serted, glabrous; ovary apocarpous, ovoid, about 0.06 em. long, glabrous; stigma about 0.2 cm. long; style about 0.2 em. long; neetaries about equalling the ovary ; follieles unknown. COLOMBIA: SANTANDER DEL NORTE: eastern slope of Paramo del Hatieo, en route from Toledo to Pamplona, alt. 2300 m., edge of woods, March 12, 1927, Killip $ Smith 20568 (NY, TYPE, MBG, photograph and analytical drawings). 43. Forsteronia floribunda (Sw.) G. F. W. Meyer, Fl. Esseq. 135. 1818; A. DC. in DC. Prodr. 8: 437. 1844; Miers, Apoc. So. Am. 243. 1878. Echites floribunda Sw. Prodr. 52. 1788. Parsonsia floribunda (Sw.) R. Br. Mem, Wern. Soc. 1: 65, 1811. Forsteronia Alexandri Griseb. Fl. Brit. W. I. 412. 1861. Stems relatively stout, glabrous or essentially so, very in- conspicuously and minutely lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptie to broadly oval, apex very shortly and abruptly acuminate to obtuse, base broadly obtuse, 4-11 em. long, 0.8-5.0 em. broad, coriaceous, glabrous throughout, the secondary venation subhorizontal and relatively dense, eglandular, somewhat nitidulous above, conspicuously paler beneath; petioles 0.35-0.8 em. long; inflo- rescence aggregate dichasial to thyrsiform, terminal, some- what shorter than the subtending leaves, bearing many small, greenish-white flowers; peduncle very minutely papillate; pedicels 0.3-0.4 em. long, somewhat accrescent in fruit, mi- nutely papillate; braets ovate, 0.05-0.15 em. long, scarious, persistent ; calyx-lobes ovate, acute to obtuse, 0.21-0.3 em. long, essentially glabrous or very minutely and indefinitely papil- late without, the alternate squamellae solitary or in groups of 2-3; corolla-tube 0.2-0.3 em. long, about 0.08-0.125 em. in diam- (248) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 215 eter at the base and 0.175-0.25 em. in diameter at the orifice, glabrous or very minutely and indefinitely papillate without, villosulose within, the lobes oblong-lanceolate, acute to obtuse, 0.39-0.45 em. long, widely spreading, essentially glabrous with- out and within; staminal filaments 0.3-0.4 cm. long, exappen- diculate, agglutinated to the style, the anthers 0.25-0.3 em. long, wholly exserted, glabrous; ovary apocarpous, ovoid, mi- nutely and sparsely pilosulose to essentially glabrate, about 0.08 em. long; stigma 0.1-0.12 cm. long; style 0.3—0.4 em. long; nectaries somewhat shorter than the ovary ; follicles relatively long and slender, continuous or only indefinitely undulated, subparallel or somewhat tortuous to sharply divaricate, 16— 30 em. long, glabrous; seeds 0.7—0.9 em. long, the pale tawny coma 1.5-2.1 em. long. JAMAICA: woodlands near Newport, Parish of Manchester, Sept. 3-7, 1908, Britton 3196 (B, FM, NY, US); rocky wooded hill, Troy, Cockpit Country, Sept. 13-18, 1906, Britton 641 (NY); near Troy, alt. 2000 ft., April, 1916, Perkins 1100 (B); same locality, March 23, 1917, Perkins 1378 (B, D); Farm Pen, alt. 200 ft., Aug. 23, 1895, Campbell 5831 (B, FM, NY); growing over bushes, road to Hagley Gap, alt. 1800 ft., July 10, 1903, Harris 5903 (B, FM, NY); Stony Hill, Jan. 22, 1903, Harris 8440 (B); Hope Gardens, June 3, 1903, Harris 8599 (B, FM, US); Sehwallenburgh, St. Ann, Jan. 26, 1898, Harris 7042 (B, NY); data incomplete: Alexander s.n. (B, NY); Bertero s.n. (B); Swartz s.n. (S, TYPE, MBG, photo- graph and analytieal drawings). Popular names for this species are said to be ‘‘ Milk Withe’’ and “Rubber Withe." Because of its isolation in Jamaica, it is remarkably constant and free of conspicuous variability. In this regard it resembles Mandevilla torosa (Jacq.) Wood- son, also the only species of its genus upon the island. 44, Forsteronia peninsularis Woodson, spec. nov. Frutieosa volubilis altitudine ignota; ramulis crassiusculis juventate minutissime puberulo- papillatis mox glabratis ma- turitate cortice brunneis conspicue lenticellatis ; foliis oppositis longiuscule petiolatis oblongo-elliptieis apice anguste acutis acuminatisve basi obtusis 4.5-5.5 em. longis 1.7-2.2 em. latis firme membranaceis glaberrimis utrinque opacis eglandulosis nervis secundariis sat remotis arcuatis; petiolis 0.5—0.8 cm. longis glabris; inflorescentiis subthyrsiformibus terminalibus foliis paululo brevioribus flores multas parvas viridi-albidas (249) [Vor. 22 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN gerentibus; pedunculo minute puberulo-papillato; pedicellis 0.2-0.25 em. longis ut in pedunculo vestitis; calycis laciniis ovatis obtusis 0.15 em. longis extus puberulo-papillatis ; corol- lae tubo 0.2 em. longo basi ca. 0.08 cm. diametro metiente fauci- bus ea. 0.1—0.125 em. extus prope orifieium minute puberulo- papillato eaeterumque glabro intus villosulo lobis oblongo- ellipticis obtusis 0.4—0.425 em. longis ca. 0.125 em. latis extus puberulo-papillatis intus papillatis reflexis; antheris 0.25 em. longis valde exsertis filamentis stylum agglutinatis laxe pilosu- lis; ovario ovoideo ca, 0.1 em. longo dense minuteque puberulo; nectariis haud concrescentibus ovarium ca. dimidio aequanti- bus; stigmate ca. 0.09 em. longo; follieulis teretibus gracilibus glabris plus minusve falcatis maturitate non visis; seminibus ignotis. Stems relatively stout, very minutely puberulent-papillate when young, soon becoming glabrate, conspicuously lenticel- late when fully mature; leaves opposite, petiolate, oblong-ellip- tie, apex narrowly acute to acuminate, base obtuse, 4.5-5.5 em. long, 1.7-2.2 em. broad, firmly membranaceous, glabrous, opaque, eglandular, secondary venation relatively distant, arcuate; petiole 0.5-0.8 cm. long, glabrous; inflorescence sub- thyrsiform, broadly pyramidal in outline, terminal, somewhat shorter than the subtending leaves, bearing numerous green- ish-white flowers; peduncle minutely puberulent-papillate; pedicels 0.2-0.25 em. long, clothed with a similar indument to that of the peduncle; calyx-lobes ovate, obtuse, 0.15 em. long, puberulent-papillate without; corolla-tube 0.2 em. long, about 0.08 em. in diameter at the base and 0.1—0.125 em. at the orifice, minutely puberulent-papillate toward the orifice without, gla- brous to glabrescent below, villosulose within, the lobes oblong- elliptie, obtuse, 0.4—0.425 cm. long, about 0.125 em. broad, without puberulent-papillate, within papillate, reflexed; anthers 0.25 cm. long, glabrous, the filaments exappendiculate, laxly pilosulose, agglutinated to the style; ovary ovoid, apocar- pous, about 0.1 cm. long, densely and minutely puberulent; stigma 0.09 em. long; nectaries about half equalling the ovary ; immature follicles terete, relatively slender, more or less fal- eate, glabrous; seeds unknown. (250) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 217 Britis HoNpURAS: Maskall, Northern River, July 16, 1934, Gentle 1281 (MBG, TYPE). Differing from F. floribunda (Sw.) G. F. W. Meyer, which it closely simulates, in the firmly membranaceous texture and arcuate venation of the foliage, and conspicuously puberulent- papillate corolla. 45. Forsteronia portoricensis Woodson, Ann. Mo. Bot. Gard. 21: 618. 1934. Stems relatively stout, glabrous, or very minutely papillate when young, eventually becoming inconspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong- elliptie to oval, apex very shortly and abruptly acuminate to acute or obtuse, base broadly obtuse, 3.2-9.0 em. long, 1.3- 4.7 em. broad, coriaceous, the margin somewhat revolute upon desiecation, secondary veins areuate and relatively distant, wholly glabrous, eglandular, nitidulous above, somewhat paler beneath; petioles 0.4-0.6 cm. long; inflorescence aggregate dichasial, terminal, about equalling or somewhat shorter than the subtending leaves, bearing many small, red or flesh-colored flowers; peduncle minutely puberulent-papillate to glabrate; pedicels 0.2-0.3 cm. long, papillate or infrequently minutely and sparsely puberulent-papillate; bracts ovate, 0.1-0.25 em. long, scarious; calyx-lobes ovate, acute to obtuse, 0.18-0.3 cm. long, scarious, minutely papillate to essentially glabrous with- out, the alternate squamellae in groups of 2-4; corolla-tube 0.15-0.2 em. long, about 0.12-0.15 em. in diameter at the base and 0.18-0.2 cm. in diameter at the orifice, glabrous without, glabrous or very indistinctly papillate within, the lobes oblong- ligular, obtuse, 0.4-0.5 em. long, widely spreading, glabrous within and without, the margin frequently minutely and sparsely ciliolate; staminal filaments 0.3-0.32 em. long, exap- pendieulate, agglutinated to the style, the anthers 0.25-0.3 em. long, wholly exserted, glabrous; ovary apocarpous, ovoid, about 0.07 cm. long, minutely puberulent to essentially gla- brous; stigma 0.1-0.13 em. long; style 0.25-0.3 em. long; nec- taries about equalling the ovary; follicles relatively slender, ` acuminate, somewhat flexuose, reflexed-divarieate, more or (251) [Vor. 22 218 ANNALS OF THE MISSOURI BOTANICAL GARDEN less falcate, 14-19 em. long, glabrous; seeds 0.9-1.0 em. long, the pale tawny coma 1.8-2.0 em. long. Porto Rico: prope Humacao in fruticetis litoralibus ad ‘‘Candelero,’’ Sept. 29, 1886, Sintenis 5195 (B, TYPE, MBG, photograph and analytical drawings); Sierra de Naguabo ad Rio Blanco, in silv. prim., Sept. 5, 1886, Sintenis 5357 (B, Bx, S); pro ros in sylv. prim. mont. ‘‘Guayava,’’ sl ““La Pandura,’’ Oct. 12, 1886, Sintenis 5306 (B); sia pame in Barrio del Pasto, Febr. 25, 1885, Sintenis 2866 (B, adi Marieao, in sylvis montis ‘‘ Alegrillo,’’ Nov. 26, 1884, Sintenis 273 (B, M, 8, US); Sa. de Luquillo, in monte Jimenes, July, 1885, Sintenis 1622 (B, D, FM, e pei Playa de Humaeao in fruticetis paludosis, June, 1881, Eggers 438 (B, BB, D); pr. Mayaguez, 1875, Krug 709 (B); pr. Ciales in fruticetis ad “Torre e May 9, 1887, Bindenia 6823 (B); prope Bayamon in fruticetis, Oet., 1887, Stahl 743 (B) ; Luquillo Mountains, July 14, 1902, Wilson 214 (B, NY), data incomplete, Wydler 209 (B, DC, NY); Sierra de Ligia, 1854, Blauner 181 (DC); Rio Piedras, May 7, 1912, Johnston 357 (NY). 46. Forsteronia corymbosa (Jacq.) G. F. W. Meyer, Fl. Esseq. 133. 1818; A. DC. in DC. Prodr. 8: 437. 1844; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Echites ciriimbasa Jacq. Enum. Pl. Carib. 13. 1760; Select. Stirp. Am. Hist. 1: 34;2: pl. 30. 1763. Periploca umbellata Aubl. Hiat. Pl. Gui. Fr. 1: 273. 1775, fide Urban. Parsonsia corymbosa (Jacq.) R. Br. Mem. Wern. Soc, 1 Thyrsanthus corymbosus (Jacq.) Miers, Apoc. So. Am, 97. 1 Stems relatively stout, glabrous or essentially so, rather in- conspicuously lenticellate when fully mature; leaves opposite petiolate, obovate to broadly oval, apex obtuse or rounded to very shortly and abruptly submucronulate-acuminate, base broadly obtuse, 3-7 em. long, 1.0-3.5 cm. broad, coriaceous, eglandular, margins somewhat revolute upon desiccation, sec- ondary venation arcuate, relatively distant, glabrous and nit- idulous above, beneath paler, glabrous, or very minutely and inconspicuously barbellate in the axils of the midrib; petioles 0.3-1.1 cm. long; inflorescence aggregate dichasial, terminal, about equalling or somewhat shorter than the subtending leaves, bearing many small, red, or infrequently flesh-colored flowers; primary peduncle minutely and rather irregularly puberulent-papillate to essentially glabrate, the ultimate (252) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 219 branches determinate, very minutely puberulent-papillate to essentially glabrate ; pedicels 0.2-0.5 em. long, minutely puber- ulent-papillate to essentially glabrate; bracts ovate to ovate- lanceolate, 0.1-0.38 cm. long, scarious or only slightly foli- aceous ; ealyx-lobes ovate, obtuse, 0.17-0.25 em. long, scarious, papillate to essentially glabrous without, the alternate squa- mellae in groups of 2-4; corolla-tube 0.19-0.25 cm. long, about 0.1-0.125 em. in diameter at the base and 0.125—0.25 em. in diam- eter at the orifice, essentially glabrous without and within, the lobes oblong-ligular, 0.4-0.52 em. long, widely spreading, mi- nutely but definitely papillate within, rarely glabrate; staminal filaments exappendiculate, 0.28-0.35 em. long, agglutinated to the style, the anthers 0.25-0.32 cm. long, widely exserted, gla- brous; ovary apocarpous, ovoid, about 0.08 cm. long, minutely puberulent to essentially glabrous; stigma 0.07—0.1 cm. long; style 0.3 em. long; nectaries about equalling the ovary ; follicles relatively short and stout, sharply divaricate, rigid and nearly rectilinear, not acuminate, 11-14 em. long, glabrous when fully mature; seeds 0.65-0.7 cm. long, the pale tawny coma 1.9— 2.1 cm. long. CUBA: HAVANA: in rupibus caleareis ad flum. Cojimar, June 14, 1914, Ekman 1355 (S); MATANZAS: prope Mantanzas, Pri ned $43 (B, NY); SANTA CLARA: Calicita, July 24, 1895, Ph 524 (B, FM, MBG, NY); Soledad, pes fuegos, climbing on trees, Nov. 6, 1928, Jack e (AA, FM, S); PINAR DEL R Bay 0f Mariel, Sept. 21, 1910, ud $ Earle 7652 (FM, NY); CAMAGUEY; vicin- ity of Tiffin, Oct. 30-31, 1909, Shafer 2872 (NY, US); ORIENTE: Yateras et Mt. y, 184 Holguin, April 18, 1909, Shafer 1447 (B, NY, US); Florida, in mont. su Daiquiri, alt. 750 m., June 28-29, 1914, Ekman 1548 (S); Sabana Miranda prope Bayate ad margin. ‘‘savannas,’’ July 11, 1914, Ekman 1937 (S); Sierra de Nipe ad Rio Piedra ad marg. sylvae, Oct. 4, 1919, Ekman 9827 (S); Baraeoa, in collibus, Jan. 8, 1915, Ekman 4151 (S); data incomplete, Wright 398 (B, Bx, Camb. MBG, NY, 8). HISPANIOLA: HAITI: Dep. du Sud, inter Cabaillon et Aux Cayes, T 7, 1917, Ekman 847 (S); wooded hill, alt. 650 m., fle La Gonave, July 28, 1927, Ekm 8730 (B, S); in Morne Coudré, Massif du No rd, Marmelade, alt. a m., May 29, 1927, Ekman 8289 (B, US); Schattige Stellen, Passe David, Gonaives, Na. 1900, Buch 339 (B); prope ed Lis July, 1899, Buch 176 (B); twining on shrub, trail north of ** Digue Puits,’’ vicinity of La Vallé, Tortue Island, May 3-10, 1929, Leonard $ Leonard 15543 (MBG, US); sommet du Morne Bellevue, ouest de Naneivet, alt. 600—700 m., Aug. 31, 1908, Christ ps (B); prope nn alt. 400-500 m., Aug. 17, 1891, Picarda 813 (B); prope Payan ad viam ad fuent (253) [Vor. 22 220 ANNALS OF THE MISSOURI BOTANICAL GARDEN versus, Jan., 1891, Picarda 154 (B) ; SANTO DOMINGO: Sierra del Palo Quemado, alt. 500 m., May 10, 1887, Eggers 1894 (B, D, M) ; in sylva montis, ** Izabel de la Torée,”” alt. 300 m., July 30, 1887, Eggers 1894b (B); prope Puerto-Plata, July 2, 1887, Eggers 1894c (B) ; in pineto, prope Constanza, alt. 1190 m., July, 1910, Tuerckheim 3476 (B, D, M, I prope Maniel de Ocoa, alt. 300 m., Oct., 1910, Tuerckheim 3636 (B, D, M, NY); semi-arid pine region, Moncion, Pun. Monte Cristy, alt. 375 m., Oct. 9, Tw Valeur 226 (D, MBG, NY, US); prope Barahona, July, 1910, Fuertes 410 (B, D, FM, MBG, NY, 8, US). An interesting account of the aspect of these plants is re- corded by Christ: **Liane trés-longue, ligneuse, affectionnant, semble-t-il, de Manguier, sur les branches les plus élevés duquel elle s’étale. Fleurs rouges foncés, nombreux, ne poussant qu'à la partie superieure de la tige étalée. Les fruits ont une double position: au début relevés l'un vers l'autre et se touchant aux extremités. A maturité écartés completement l'un de l'autre, sur une méme ligne horizontal. ”” Ekman 8730 bears flowers described as ‘‘pallide carneis’’ by the collector. These plants may eventually be found to repre- sent a color form. The typical color of the flowers is dark red. The dried specimens show the flowers to be much paler than all others examined. Subgen. II. PrERANTHERA Woodson, subgen. nov. Stamen filaments bearing a conspicuous, alate, membrana- ceous appendage just below the insertion of the anther. Spp. 47—48. KEY TO THE SPECIES a. Inflorescence glabrous or essentially so; von firmly membranaceous, the not revolute in desiecation, 7-15 € loNE......1....» 47. F. refracta aa. flee minutely puberulent to M t at least in part; leaves eoriaeeous, the margin somewhat revolute in desieeation, 6—7 em. (cr! Serre EA AA 48. F. fallax 47. Forsteronia refracta Muell.-Arg. in Mart. Fl. Bras. 6': 97. 1860; Miers, Apoc. So. Am. 244. pl. 35 B. 1878. Forsteronia floribunda Muell.-Arg. loc. cit. 96. 1860, not G. F. W. Meyer. Forsteronia lagoënsis Muell.-Arg. in Warming, Kjoeb. Vidensk. Meddel. 108. 1869. Thyrsanthus myrianthus Miers, loc. cit. 105. 1878. (254) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 223 Forsteroma refracta Muell.-Arg. var. contracta Taub. in Engl. Bot. Jahrb. 21: 448. 1896, Stems relatively stout, glabrous, rather inconspicuously lenticellate; leaves opposite or infrequently ternate, shortly petiolate, rather narrowly elliptic-oblanceolate to broadly oval, apex shortly and abruptly acuminate to acute, infrequently obtuse upon older specimens, 7-15 em. long, 2.5-8.0 em. broad, firmly membranaceous, above glabrous, inconspieuously glan- dular at the base of the midrib, glabrous beneath, or very mi- nutely and sparsely ciliolate in the axils of the midrib; petioles 0.2-0.9 cm. long; inflorescence laxly thyrsiform, terminal, usu- ally greatly surpassing the subtending leaves, occasionally somewhat shorter, bearing many small, white flowers; primary peduncle glabrous, ultimate branches glabrous, determinate; pedicels 0.17-0.35 cm. long, glabrous; bracts ovate to ovate- lanceolate, deciduous, 0.05-0.2 em. long, scarious; calyx-lobes broadly ovate, acute to obtuse, 0.09—0.11 em. long, scarious, gla- brous without, the squamellae 3-4 times as many as the lobes, regularly and indefinitely distributed ; corolla-tube 0.1-0.12 em. long, about 0.05-0.07 cm. in diameter at the base and 0.2- 0.225 em. in diameter at the orifice, glabrous without, or essen- tially so, puberulent-papillate within, the lobes ovate, 0.2- 0.28 cm. long, widely spreading, densely and conspicuously pilosulose within; staminal filaments 0.1-0.12 cm. long, dis- tinctly bi-alate toward the insertion of the anthers, the anthers about 0.1 em. long, glabrous, exserted; ovary apocarpous, ovoid, about 0.07 cm. long, glabrous, or very minutely and sparsely pilosulose toward the tips; stigma 0.05-0.08 em. long; style about 0.2 cm. long; follicles relatively long and stout, con- spicuously and rather distantly moniliform, relatively rigid and subparallel, occasionally somewhat divaricate, 28-52 em. long, glabrous ; seeds 2.7-2.9 em. long, the brilliant tawny coma 5.0-5.2 em. long. VENEZUELA: MERIDA: prope Coloniam Tovar, 1854-5, Fendler 2381 (MBG). BRAZIL: PARA: understory in cut-over wood, Japanese Colony, Thomé Assu, Distrieto Acara, July 16, 1931, Mexia 5915 in part (MBG); RIO DE JANEIRO: Theresopolis, alt. 900 m., Oct. 11, 1929, Brade 9672 (B); data incomplete: Glaziou 7756 (B); Glaziou 2091 (Bx); MINAS GERAES: in shade of forest, road to Sáo (255) [Vor. 22 222 ANNALS OF THE MISSOURI BOTANICAL GARDEN Miguel, alt. 710 m., frequent, Oct. 29, 1930, Mexia 5234 (MBG) ; Lagóa Santa, Aug. 29, 1864, Engle s.n. (C); data incomplete: Lhotzky s.n. (B); Araujo 11935 (B); Gardner 5012 (B, DC, $ ); PARANA: in silva primaeva, Porto de Cima, Dec. 23, 1908, Dusen 7445 (Bx, D, MBG, NY, US); DATA INCOMPLETE: Sellow s.n. (B, Bx, COTYPE, MBG, photograph and analytical drawings). PARAGUAY: in altaplanitie et declivibus ‘‘Sierra de Amambay,’’ Oct., 1907, Rojas 10652 (B) ; in regione fluminis, Alto Parana, 1909-10, Fiebrig 5380 (M). ARGENTINA: MISIONES: San Pedro, Rosados de los Indios, Nov. 9, 1886, Nieder- lein 1965 ( Considerable doubt remains concerning the identity of Fendler 2381 from Venezuela and Mexia 5915 from the state of Para, Brazil. Both specimens bear fruit, without flowers. In either case the fruit corresponds closely to that of authentic specimens of F. refracta; but the occurrence of the species in Venezuela and northern Brazil remains to be verified by more perfeet records. 48. Forsteronia fallax Taub. in herb. Frutieosa volubilis; ramulis sat crassiusculis glaberrimis maturitate conspicue lenticellatis; foliis oppositis brevissime petiolatis elliptieis apice obtusis roimadatutvs basi obtusis sub- euneatis 6-7 em. longis 2.0-3.1 em. latis coriaceis vel subcori- aceis margine post exsiecationem paulo revolutis glaberrimis nervo medio supra inconspicue glanduligero; petiolis 0.3-0.35 em, longis; inflorescentiis laxe thyrsiformi-paniculatis termi- nalibus folia superantibus flores parvas multas albidas gerent- ibus; peduneulo minute puberulo-papillato vel basi glabriusculo ramulis minute puberulo-papillatis; pedicellis 0.07-0.1 cm. longis minute puberulo-papillatis; bracteis ovatis vel ovato- lanceolatis deeiduis 0.08-0.15 em. longis scariaceis; calycis laciniis late ovato-deltoideis obtusis rotundatisve ca. 0.08 em. longis seariaceis minutissime irregulariterque puberulo-papil- latis squamellis alternatis solitariis minutissimis ; corollae tubo ca. 0.07 em. longo basi ca. 0.04 cm. diametro metiente faucibus ca. 0.15-0.17 em. diametro metientibus extus glabris glabrius- eulisve intus minute villosulis lobis ovatis ca. 0.17 em. longis patulis extus glabriusculis intus dense conspicueque pilosulis ; filamentis staminum ca. 0.08 em. longis apice appendiculatis ; antheris ca. 0.8 em. longis glabris exsertis; ovario oblongo- ovoideo apocarpo ea. 0.04 em. longo minute papillato; stigmate (256) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 223 0.05-0.07 cm. longo; stylo ca. 0.01 em. longo; nectariis ovario paululo brevioribus ; folliculis ignotis. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, very shortly petiolate, el- liptie, apex obtuse or rounded, base obtuse, subeuneate, 6—7 em. long, 2.0-3.1 em. broad, coriaceous or subcoriaceous, margin somewhat revolute after desiccation, glabrous throughout, in- conspicuously glandular above at the base of the midrib; pet- ioles 0.3-0.35 cm. long; inflorescence laxly thyrsiform-panicu- late, terminal, somewhat surpassing the subtending leaves, bearing many small, white flowers; peduncle minutely puberu- lent-papillate to glabrous below, lato branches determi- nate, minutely puberulent-papillate ; pedicels 0.07—0.1 em. long, minutely puberulent-papillate; braets ovate or ovate-lanceo- late, 0.08-0.15 cm. long, searious, deciduous; calyx-lobes broadly ovate-deltoid, obtuse or rounded, about 0.08 em. long, scarious, minutely and irregularly puberulent-papillate with- out, the squamellae very minute, alternate, solitary; corolla- tube about 0.07 em. long, about 0.04 em. in diameter at the base and 0.15-0.17 em. in diameter at the orifice, essentially gla- brous without, minutely villosulose within, the lobes ovate, about 0.17 em. long, widely spreading, essentially glabrous without, densely and conspicuously pilosulose within ; staminal filaments about 0.08 em. long, appendiculate above, the anthers about 0.08 cm. long, glabrous, wholly exserted; ovary oblong- ovoid, apocarpous, about 0.04 em. long, minutely papillate; stigma 0.05-0.07 cm. long; style about 0.01 cm. long; nectaries somewhat shorter than the ovary ; follicles unknown. BRAZIL: RIO DE JANEIRO: data incomplete, Glaziou 4080 (B, TYPE, MBG, photo- graph and analytieal drawings). EXCLUDED SPECIES Forsteronia panniculata Casar. ex K. Sch. in Mart. Fl. Bras. 6*: 200. 1889, nom. nud. in synon. - Molopanthera paniculata Turez. Bull. Soc. Nat. Mose. 21!: 581. 1848. Forsteronia Pavoni A. DC. in DC. Prodr. 8: 438. 1844 = Apocynum cannabinum L. var. glaberrimum A. DC. loc. cit. 439. 1844. (257) [Vor. 22 224 ANNALS OF THE MISSOURI BOTANICAL GARDEN The type specimen of Forsteronia Pavoni, an unnumbered collection by Pavon in the herbarium of Boissier at Geneva, shows itself typical of the glabrous variety of the frequent and widespread Apocynum cannabinum L. of temperate North America. Pavon's collection label is without locality. Just how the plant became confused with the tropical genus For- steroma may be explained variously. The ultimate reason, naturally, is the failure of de Candolle to observe the conspieu- ous, sagittate enations at the base of the corolla-tube alternat- ing with the staminal filaments, easily perceived from a simple disseetion. The accidental inclusion of the plant, perhaps from the col- lections of another botanist from temperate North America, with those of Pavon from Mexico and Peru may have completed the circumstantial evidence misplacing the plant systemati- cally, for it is quite true that a superficial similarity exists be- tween Apocynum and Forsteronia in the thyrsiform inflores- cence and small flowers of either. A photograph accompanied by analytical drawings of the type specimen has been incorpo- rated in the herbarium of the Missouri Botanical Garden. Miers (Apoc. So. Am. 243. 1878), without easy access to Pavon's specimen, erroneously referred to this species Spruce 4908, more correctly referable to F. tarapotensis K. Sch. (cf. p. (230) ). VI. Seconvarıa A. DC. Secondatia A. DC. in DC. Prodr. 8: 445. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 107. 1860; Benth. & Hook. Gen. Pl. 2: 123. 1876; Miers, Apoc. So. Am. 226. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4°: 165. 1895. Orthechites Urb. Symb. Ant. 6: 36. 1909. Lactescent, fruticose or suffruticose lianas. Stems volubile (or erect in S. Macnabu ?), terete, the branches opposite or alternate above. Leaves opposite, petiolate, entire, penni- nerved, eglandular; nodes minutely appendiculate. Inflores- cence terminal, or both terminal and lateral, thyrsiform, brae- teate, several-flowered. Calyx 5-parted, the lobes equal to sub- equal, cleft nearly to the receptacle, imbricated, bearing within (258) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 225 solitary, or rarely paired, alternate squamellae. Corolla sal- verform, the tube straight, exappendieulate within, the limb actinomorphic, 5-parted, dextrorsely convolute. Stamens 5, the anthers connivent and agglutinated to the stigma, consist- ing of 2 parallel, linear sporangia borne ventrally near the apex of an enlarged, sagittate, narrowly 2-lobed, peltate connective; pollen granular; filaments short. Carpels 2, united at the apex by a common stylar shaft surmounted by the fusiform stigma (sessile in S. floribunda and S. Duckei) ; ovules numerous, sev- eral-seriate, borne upon an axile, binate placenta. Nectaries 9, more or less concrescent at the base. Follicles 2, apocarpous, broadly fusiform, dehiscent along the ventral suture, contain- ing many dry, truncate, apically comose seeds. Type species: Secondatia densiflora A. DC. in DC. Prodr. 8: 445. 1844. KEY TO THE SPECIES a. Corolla-tube 0.5—0.82 em. long; speeies of South Ameriea. b. Stigma supported by a short style; PAES nearly equalling the ovary. c. Corolla-lobes obliquely obovate, shorter than the tube. . Anthers SREE puberulent raid eiu HET E S. densiflora ad. Anthers glabrous. c SEMBRA E e o as sei . S. peruviana Ces pi lobes bloss: -dolabriform, equalling or slightly surpassing the p YO ct tito Joa RM NE 3. S. Schlimiana bb. Stigma sessile; nectaries much shorter than the ovary. €. Corolla- TM obliquely elliptie, 0.7-1.2 em. long; ovary glabrous...... e E O A T SFO 4. S. floribunda ec. Corolla-lobes oblong-linear, 2.2-2.5 em. long; ovary p nes and arsely puberulant. u... ste aes op RET a d RÀ . S. Duckei aa. erie tube about 1.5 em. long; plants of Jamaica............ 6. T Macnabü 1. Secondatia densiflora A. DC. in DC. Prodr. 8: 445. 1844; Muell.-Arg. in Mart. Fl. Bras. 61: 108. pl. 32. fig. 2. 1860; Miers, Apoc. So. Am. 226. pl. 32. 1878; K. Sch. in Engl. & Pra Nat. Pflanzenfam. 4?: 165. 1895. Secondatia densiflora A. DC. B parvifolia Muell.-Arg. loc. eit. 1860. Secondatia densiflora A. DC. var. paraguariensis Hassl. in Fedde, Rep. Sp. Nov. 12: 263. 1913 Secondatia densiflora A. DC. a genuina Muell.-Arg. ex Hassl. loc. cit. 264. 1913, sphalm. (259) [Vor. 22 226 ANNALS OF THE MISSOURI BOTANICAL GARDEN Stems relatively stout, minutely papillate when very young, soon becoming glabrate and conspicuously lenticellate when fully mature; leaves broadly ovate-, obovate-, or oblong-ellip- tic, apex abruptly subcaudate-acuminate, base broadly obtuse or rounded, 3.5-11.5 em. long, 1.5-6.5 cm. broad, firmly mem- branaceous to subcoriaceous, glabrous, essentially concolorous or only slightly paler beneath, the vein-ends immersed and relatively obscure; petioles 0.7-1.5 em. long; inflorescence sub- eorymbose, terminal, or infrequently lateral as well, much shorter than the subtending leaves, bearing numerous small, white flowers; pedicels 0.35—0.6 cm. long, greatly acerescent in fruit, minutely papillate or essentially glabrous; bracts ovate or ovate-oblong, 0.1-0.3 em. long, scarious or only slightly foli- aceous; calyx-lobes ovate to ovate-oblong, obtuse, 0.15-0.25 cm. long, scarious, minutely papillate to essentially glabrous; corolla salverform, glabrous without, the tube 0.65-0.8 em. long, about 0.1-0.12 cm. in diameter at the base, not conspicu- ously dilated at the insertion of the stamens, slightly narrow- ing toward the orifice, densely puberulent within, the lobes obliquely obovate, obtuse, 0.35-0.5 em. long, glabrous, spread- ing; stamens inserted at about the lower 14 of the corolla-tube, the anthers 0.35—0.4 em. long, minutely puberulent dorsally; stigma 0.175-0.25 em. long, surmounting a style of about equal length; ovary ovoid, about 0.07 em. long, glabrous; nectaries concrescent at the base, slightly shorter than the ovary; fol- licles broadly fusiform, somewhat compressed laterally, 12— 13 em. long, about 2.0-2.5 cm. in greatest diameter, glabrous; seeds about 2 em. long, the pale tawny coma 3.5—4.0 em. long. BRITISH GUIANA: Pirara, etc., 1841-2, Schomburgk 421 (V); same locality, Schomburgk 681 (B); data ineomplete, Schomburgk 599 (V, COTYPE); Schom- burgk 665 (D) DUTCH GUIANA: fluv. Lawa, Oct., 1903, Versteeg 451 (U). BRAZIL: PIAUHY: exact locality lacking, 1883, Netto 46 (US); MINAS GERAES: data ineomplete, Claussen 341 (B, COTYPE); RIO DE JANEIRO: Sumidouro, 1845, Pohl s.n. (V); data incomplete, Sellow s.n. (Bx); GOYAZ: data incomplete, Gardner 8325 (B, D, corYPE, NY, V); MATTO GROSSO: Cuyaba, Nov. 21, 1893, Malme 1118 (B, D, S) ; Cuyaba, 1834, Manso s.n. (Bx) ; Rio Arimos, margins do rio, Dec., 1914, Kuhlmann 1234 (US); Uferwald am oberen Paranatinga, Sept. 28, 1899, Pilger 799 (B); são PAULO: Rib. da Lagóa, Rio Feio, cerrado, Nov., 1905, Edwall (260) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE. IV 221 11242 (B); PARA: campos do Ariramba, prope flum. Jaramaearu, Dee. 2, 1910, Ducke 21622 (B, US); Obidos, silva secundaria, Dec. 11, 1903, Ducke pr (B); AMAZONAS: in E des campo bei der Serra de Pracama, Rio Braneo, Jan., 1909, Ule 7940 (B); silva humosa non inundabili, Sáo Paulo de "inp Auf. 19, 1929, Ducke 22430 (B); DATA INCOMPLETE: Riedel s.n. (B, BB, NY, V); Pohl 1845 (V); Martius 967 (B, D, TYPE, M, NY, V, MBA, photograph and analytical drawings); Glaziou 11179 (B). BOLIVIA: SANTA CRUZ: Bañado I Rio Moreno, Prov. Cereado, alt. 450 m., Oct. 28, 1925, Steinbach 7301 (B, D, FM, MBG, U); Bañado del Rio Perdiz, Prov. ing alt. 450 m., ee 30, 1916, ade 3113 (B, D); LA PAZ: Millaguaya, alt. m., Dec, 1 T RON 4371 (B, US); BENI: Riberalta, Sept. 28, 1923, sd 21873 e U, US). PARAGUAY: in altaplanitie ‘‘Sierra de Amambay,” Nov., 1912, Hassler 11420 (D, V). 2. Secondatia peruviana Poeppig, Nov. Gen. 3: 71. pl. 281. 1845; Miers, Apoc. So. Am. 228. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 165. 1895. Stems relatively slender, glabrous or essentially so, conspic- uously lenticellate when fully mature; leaves ovate-elliptic, apex abruptly subcaudate-acuminate, base broadly obtuse or rounded, 5.2-9.5 em. long, 2.2-5.0 cm. broad, membranaceous, glabrous, essentially concolorous or only slightly paler be- neath, the vein-ends immersed and relatively obscure; petioles 0.7-1.5 em. long; inflorescence subcorymbose, both terminal and lateral, much shorter than the subtending leaves, bearing numerous small, white flowers; pedicels 0.2-0.5 em. long, gla- brous or very minutely papillate; bracts ovate, 0.1-0.2 cm. long, scarious; calyx-lobes ovate, acute to obtuse, 0.1-0.15 em. long, scarious, glabrous or very minutely ciliolate; corolla sal- verform, glabrous without, the tube 0.7-0.75 em. long, about 0.125-0.15 em. in diameter at the base, not conspicuously di- lated at the insertion of the stamens, slightly narrowing to- ward the orifice, puberulent within, the lobes obliquely obovate, obtuse or rounded, 0.4—0.55 em. long, glabrous, spreading; stamens inserted at about the lower 14 of the corolla-tube, the anthers 0.4—0.45 cm. long, glabrous; stigma 0.18-0.2 em. long, supported by a style of about equal length; ovary ovoid, some- what less than 0.1 cm. long, glabrous; nectaries concrescent at the base, slightly shorter than the ovary; mature follicles unknown. (261) [Vor. 22 228 ANNALS OF THE MISSOURI BOTANICAL GARDEN PERU: LORETO: Cuchero et in sylvis montanis ad Pampayaro, Dee., 1829, Poeppig 1582 (B, BB, MBG, V, TYPE); Stromgebiet des Maranon, Santiago-Mündung, Oct. 24, 1924, Tessmann 4359 (B, D). The immature fruit of this species is figured by Poeppig (loc. cit. pl. 281. fig. 9. 1845) as broadly ovoid, and sharply divaricate, and described as about 1 inch in length. Bentham and Hooker (Gen. Pl. 2: 723. 1876) favored the rejeetion of S. peruviana from the genus Secondatia upon this evidence, be- lieving the fruit of the genus to be narrowly linear. The only follicles known of the type species (Martius 967 in Herb. Vindob. and Manso s.n. in Herb. Brux.), however, are broadly fusiform, and are suggested plainly by Poeppig’s drawings of the immature mericarps of S. peruviana. It appears wholly probable that the fruits of S. densiflora and S. peruviana are no more than specifically distinct when mature. 3. Secondatia Schlimiana Muell.-Arg. Linnaea 30: 416. 1860; Miers, Apoc. So. Am. 227. 1878. Stems relatively stout, minutely puberulent-papillate when very young, soon becoming glabrate and conspicuously lenticel- late when fully mature; leaves broadly oval to ovate-elliptie, apex abruptly subcaudate-acuminate, base broadly obtuse or rounded, 3.7-6.0 em. long, 1.7-3.0 em. broad, membranaceous, glabrous, essentially concolorous or only slightly paler be- neath, the vein-ends immersed and relatively obscure; petioles 0.7-1.2 em. long; inflorescence subeorymbose, terminal, or lat- eral as well, much shorter than the subtending leaves, bearing numerous small, white flowers; pedicels 0.2-0.4 em. long, mi- nutely puberulent-papillate to essentially glabrous; bracts ovate-oblong, 0.1-0.2 cm. long, scarious or only slightly foli- aceous; calyx-lobes ovate-suborbicular, broadly obtuse or rounded, 0.15-0.2 em. long, minutely puberulent-papillate with- out, scarious; corolla salverform, glabrous without, the tube 0.7-0.8 cm. long, about 0.175 cm. in diameter at the base, not conspicuously dilated at the insertion of the stamens, slightly narrowing toward the orifice, puberulent within, the lobes ob- long-dolabriform, obtuse or rounded, 0.9-1.0 em. long, gla- brous, reflexed or widely spreading; stamens inserted at about the lower 14 of the corolla-tube, the anthers 0.4—0.45 em. long, (262) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE. IV 229 minutely puberulent dorsally; stigma 0.17-0.2 em. long, sup- ported by a style of about equal length; ovary ovoid, about 0.1 em. long, glabrous; nectaries concrescent at the base, about as long as the ovary ; follicles unknown. COLOMBIA: SANTANDER DEL NORTE: Ocana, alt. 4000 pp., May, 1846-52, Schlim 510 (BB, TYPE, BM, Bx, D, MBG, photograph and analytical drawings). 4. Secondatia floribunda A. DC. in DC. Prodr. 8: 446. 1844; Muell.-Arg. in Mart, Fl. Bras. 6': 109. 1860; Miers, Apoc. So. Am. 227. 1878. Secondatia foliosa A. DC. loc. cit. 1844; Muell.-Arg. loc. cit, pl. 32. fig. 1. 1860; Miers, loc. cit. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 165. 1895. Secondatia foliosa A. DC. p. Gardneri A. DC. loc, cit. 1844; Muell.-Arg. loc. cit. 1860. Secondatia foliosa A, DC. y. petiolaris Muell.-Arg. loc. cit. 1860. Secondatia foliosa A. DC. 9. lanceolata Muell.-Arg. loc. cit. 1860. Stems relatively slender, glabrous or essentially so, conspic- uously lenticellate when fully mature; leaves ovate-elliptic to broadly oval, abruptly acuminate to subcaudate-acuminate, base broadly obtuse or rounded, 3-9 em. long, 1.5—4.5 em. broad, firmly membranaceous to subcoriaceous, glabrous, conspicu- ously pallid beneath, the vein-ends very conspicuous and retic- ulate; petioles 0.4—1.4 cm. long; inflorescence subeorymbose, terminal, or lateral as well, somewhat shorter than the subtend- ing leaves, bearing numerous small, white flowers; pedicels 0.6—0.8 em. long, glabrous or very minutely puberulent-papil- late; braets ovate to ovate-lanceolate, 0.075-0.125 em. long; calyx-lobes ovate, acute, 0.15-0.2 em. long, scarious, minutely puberulent-papillate to essentially glabrous without; corolla salverform, glabrous without, the tube 0.55—0.65 em. long, about 0.1-0.125 em. in diameter at the base, conspicuously dilated at the insertion of the stamens, slightly narrowing toward the orifice, the lobes obliquely elliptie, broadly acute, 0.7-1.2 em. long, glabrous, widely spreading; stamens inserted barely above the base of the corolla-tube, the anthers 0.33-0.35 em. long, long-acuminate, minutely puberulent dorsally; stigma (263) [Vor. 22 230 ANNALS OF THE MISSOURI BOTANICAL GARDEN sessile, 0.15—0.18 em. long; ovary ovoid, about 0.125 em. long, glabrous or very minutely papillate; nectaries concrescent at the base, much shorter than the ovary ; follicles unknown. BRAZIL: CEARA: data incomplete, Gardner 1762 (NY, US, V); BAHIA: foréts, — date laeking, Blanchet 3370 (B, Bx, DC, TYPE, NY, V, MBG, photo- h and analytical drawings); Jacobina, date lacking, Blanchet 3635 (DC, M, V); Capáo bei Sineora, Nov., 1906, Ule 7118 (B); MINAS GERAES: Vespasiano, Nov., 1915, Hoehne 6251 (B); cut-over woods on hill, Fazenda de Jose Alexandre, alt. 725 m., Nov. 20, 1930, Mexia 5334a (MBG); data incomplete, Claussen s.n. (V); Catingas, locis aridis ad fl. Rio Fermozo, Sept., year lacking, Martius s.n. M); Lagóa Santa, Oct. 1864, Warming s.n. (C, NY); DATA INCOMPLETE: Glaziou 21720 (B, Bx); Glaziou 12941 (B, Bx); Pohl 1846 (B, V). Secondatia foliosa differs from the type specimen of S. flori- bunda only in slight and insignificant differences in dimension of the flowers and foliage. It appears significant that both were collected in the vicinity of Jacobina, state of Bahia, the type specimens being Blanchet 3635 and Blanchet 3370 respec- tively. One can scarcely restrain the opinion that varietal or even specific distinctions were frequently made by A. de Can- dolle and Mueller upon characteristies which are usually inter- preted as individual] variations among a wide selection of study specimens. 9. Secondatia Duckei Mef. Notizblatt 11: 338. 1932. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves broadly ovate-lanceolate to oblong- elliptie, apex rather gradually acuminate to subcaudate-acu- minate, base broadly obtuse or rounded, 7.5-12.5 cm. long, 3.0- 9.2 em. broad, firmly membranaceous to subcoriaceous, gla- brous, conspicuously pallid beneath; petioles 0.8-1.0 em. long; inflorescence much shorter than the inblenditig leaves, lax and relatively few-flowered; pedicels 0.4-0.6 cm. long, glabrous; bracts very minute; calyx- lobes ovate-trigonal, acute, 0.1- 0.12 cm. long, slahrous, scarious; corolla salverform, glabrous without, the tube 0.8-0.82 em. long, somewhat inflated at the in- sertion of the stamens, narrowing toward the orifice, puberu- lent within, the lobes linear or oblong-linear, 2.2-2.5 em. long, glabrous, spreading; stamens inserted near the base of the corolla-tube, the anthers 0.43-0.45 em. long, minutely puberu- lent dorsally; stigma sessile, 0.15-0.17 cm. long; ovary ovoid, (264) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 231 0.1-0.12 em. long, sparsely and minutely puberulent; nectaries concrescent at the base, much shorter than the ovary; follicles unknown. BRAZIL: AMAZONAS: Rio Negro super confluentiam flum. Curicuriary, silva non inundabili, Nov. 24, 1929, Ducke 22432 (B, TYPE, US, MBG, photograph and analytical drawings). 6. Secondatia Macnabii (Urb.) Woodson, Ann. Mo. Bot. Gard. 19: 385. 1932. Orthechites Macnabú Urb, Symb. Ant. 6: 37. 1909. Stems relatively stout, glabrous or essentially so, inconspic- uously lenticellate when fully mature; leaves lanceolate to ob- long-lanceolate, apex acuminate, base acute to obtuse, 5-7 cm. long, 1.3-2.3 cm. broad, firmly membranaceous or chartaceous, glabrous; petioles 0.3-0.4 em. long; inflorescence subcorym- bose, terminal, bearing relatively few (3-6) small, white flowers; pedicels 0.3-0.4 em. long, glabrous; bracts minute; calyx-lobes 0.2 cm. long, acute, minutely ciliolate; corolla sal- verform, glabrous without, the tube about 1.5 em. long, about 0.15 em. in diameter at the base, puberulent within, the lobes obliquely elliptie, acute, 0.7-0.8 em. long, spreading; stamens inserted somewhat below midway within the corolla-tube, the anthers linear-lanceolate; stigma about 0.05 em. long; ovary ovoid, glabrous; nectaries much shorter than the ovary ; fol- licles unknown. JAMAICA: exact locality and date lacking, Macnab s.n. (B, drawing of TYPE). It has been impossible to examine the type specimen of this species, which was sent with other specimens from the herba- rium at Edinburgh to the British Museum (Natural History) for examination by Fawcett and Rendle and has evidently been lost or destroyed. The drawing, together with Urban's deseription, presents no valid reason for not including the species with Secondatia, where it constitutes an element no more foreign to the type species than do S. floribunda and S. Duckei. Fora relatively small island which has been known to botanical collectors for such a long time, Jamaica has pro- dueed a surprising number of mysterious species of plants known from single, insufficiently recorded specimens. (265) [Vor. 22 232 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXCLUDED SPECIES Secondatia difformis (Walt.) Benth. & Hook. ex Miers, Apoc. So. Am. 99. 1878, in synon. (Echites difformis Walt. Fl. Carol. 98. 1788) = Trachelospermum difforme (Walt.) A. Gray, Syn. Fl. N. Am. 2: 84. 1878. Secondatia ferruginea (A. Rich.) Miers, loc. cit. 227. 1878 (Echites ferruginea A. Rich. in Sagra, Fl. Cub. 2: 92. 1853) = Angadenia Lindeniana (Muell.-Arg.) Miers, loc. cit. 180. 1878 (Rhabdadenia Lindeniana Muell.-Arg. Linnaea 30: 438. 1860). Secondatia stans (A. Gray) Standl. Contr. U. S. Nat. Herb. 23: 1165. 1924 (Trachelospermum stans A. Gray, Proc. Am. Acad. 21: 394. 1886) = Mandevilla foliosa (Muell.-Arg.) Hemsl. Biol. Centr.-Am. Bot. 2: 316. 1882 (Amblyanthera foliosa Muell.-Arg. Linnaea 30: 427. 1860). VII. TRACHELOSPERMUM Lem. Trachelospermum Lem. Jard. Fleur. 1: pl. 61. 1851; A. Gray, Syn. Fl. N. Am. 2: 84. 1878. Lactescent, suffruticose lianas. Stems volubile, terete, the branches alternate, occasionally becoming opposite after in- jury. Leaves opposite, petiolate, entire, penninerved, eglan- dular; nodes very ineonspieuously stipulate. Inflorescence lateral and alternate in our species, thyrsiform, aggregate dichasial in our species. Calyx 5-parted, the lobes essentially equal, cleft nearly to the receptacle, imbricated, bearing within alternate pairs of minute, glandular squamellae in our species. Corolla salverform, in our species the tube relatively short, slightly dilated above the insertion of the stamens, exappendic- ulate within, the orifice not annulate, the limb aetinomorphie, 5-parted, dextrorsely convolute. Stamens 5, the anthers in- cluded or the tips barely exserted in our species, connivent and agglutinated to the stigma, consisting of 2 parallel, uniformly fertile sporangia borne ventrally near the apex of an enlarged, sagittate, peltate connective; pollen granular; filaments free from the style. Carpels 2, apocarpous, united at the apex by a common style surmounted by the capitate stigma; ovules many, several-seriate, borne upon an axile, binate placenta. Nec- taries 5, essentially separate or somewhat conerescent at the (266) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 233 base. Follicles 2, apocarpous, terete and relatively slender in our species, dehiscing along the ventral suture, containing many dry, truncate, apically comose seeds. Type species: Trachelospermum jasminoides (Lindl.) Lem. Jard. Fleur. 1: pl. 61. 1851. The genus Trachelospermum includes numerous species of southeastern Asia, among which is the type species. In the western hemisphere the genus is represented by a single spe- cies, T. difforme (Walt.) A. Gray (= Echites difformis Walt.). As has already been indicated more briefly (p. (19) ante), rea- sons for placing this species in the Asiatic genus Trachelosper- mum rather than in the tropical American Secondatia may be reduced to the consideration that the distribution of E. dif- formis in the southeastern United States has been found in numerous instances to be strongly correlated with southeastern Asiatic affinities not only in another genus of Apocynaceae (Amsonia, cf. Woodson, R. E., Jr., Ann. Mo. Bot, Gard. 15: 388. 1928) but in certain other families of vascular plants as well. Morphological reasons for assigning the species to Trache- lospermum rather than to Secondatia are found in the fusiform stigma and solitary squamellae of the latter genus. The con- trasting capitate or subcapitate stigma and geminate squamel- lae of E. difformis, however, are not characters which are maintained rigidly throughout the Asiatic representation of Trachelospermum. As may be inferred from the foregoing con- siderations, the whole problem boils down to our ignorance of adequate characters to distinguish Secondatia and Trachelo- spermum as genera. Should the suspected congenericity of the two be acted upon nomenclatorially, the former name should have to be preserved by reason of priority. Thus a more famil- iar name would be displaced by one much less generally known, and a fairly large number of new nomenclatorial combinations necessitated. Such radical changes are always deplorable un- less activated by conclusive observations, which are yet lacking in the present instance. The preferred course in the situation at hand, then, would appear to be the rather arbitrary reten- tion of both Trachelospermum and Secondatia for the present, (267) [Vor. 22 234 ANNALS OF THE MISSOURI BOTANICAL GARDEN and the support of Gray's disposition of Echites difformis Walt.: 1. Trachelospermum difforme (Walt.) A. Gray, Syn. Fl. N. Am. 2: 85. 1878; Britton & Brown, Illustr. Fl. ed. 1. 3: 4. fig. 2899. 1898; Robinson & Fernald in A. Gray, Man. ed. 7. 662. 1908. Echites difformis Walt. Fl. Carol. 98. 1788. Echites puberula Michx. Fl. Bor. Am. 1: 120. 1803. Tabernaemontana populifolia Poir. Diet. Suppl. 1: 276. 1811; A. DC. in DC, Prodr. 8: 374. 1844. Echites salicifolia Raf. New Fl. N. Am. 4: 59. 1836, not Willd. Forsteronia difformis (Walt.) A. DC, loc. eit. 437. 1844. Secondatia difformis (Walt.) Benth. in Benth. & Hook. Gen. Pl. 2: 710. 1876; K. Sch, in Engl. & Prantl, Nat. Pflanzenfam. 4?: 165. 1895. Thyrsanthus populifolius (Poir.) Miers, Apoc. So. Am, 99. 1878. Thyrsanthus difformis (Walt.) Miers, loc. cit. 1878. Stems relatively slender, glabrous, or minutely and sparsely pilosulose when very young, rather inconspicuously lenticel- late when fully mature; leaves opposite, shortly petiolate, ellip- tie to obovate-elliptie, less frequently elliptie-linear to sub- orbieular, apex acuminate, occasionally very shortly and abruptly so, base acute to obtuse, occasionally rounded, 2.7- 12.0 em. long, 0.4—7.2 em. broad, delicately membranaceous, above glabrous or essentially so, beneath softly puberulent to essentially glabrous; petioles 0.1-1.5 cm. long; stipular ap- pendages interpetiolar, minute, flagelliform to narrowly den- ticulate, several; inflorescence thyrsiform, lateral, alternate, bearing numerous small, pale yellow flowers; peduncle sparsely pilosulose to essentially glabrous, ultimate branches determi- nate, minutely and sparsely puberulent-papillate to essentially glabrous; pedicels 0.4—0.7 em. long, minutely puberulent-papil- late to nearly glabrate, conspicuously accrescent in fruit; braets narrowly lanceolate to linear, 0.1—0.5 em. long, more or less foliaceous; calyx-lobes ovate-lanceolate, long-aeuminate, 0.3-0.37 em. long, only slightly foliaceous, minutely and rather (268) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 235 sparsely pilosulose-barbellate at the tips, otherwise essentially glabrous, the squamellae usually geminate, occasionally accom- panied by smaller, supernumerary individuals ; corolla salver- form or subinfundibuliform, the tube 0.55-0.65 em. long, about 0.08-0.1 cm. in diameter at the base, slightly constricted at the insertion of the stamens, then somewhat inflated, about 0.2— 0.3 em. in diameter at the orifice, essentially glabrous without, minutely hirtellous within somewhat above the insertion of the stamens, the lobes obliquely obovate, shortly acuminate, 0.3— 0.37 em. long, somewhat spreading to reflexed, glabrous with- out, minutely and indistinctly papillate to essentially glabrous within; stamens inserted somewhat below midway within the corolla- tube, the filaments 0.1-0.12 cm. long, very minutely tomentulose, the anthers narrowly sagittate, acuminate, acutely auriculate, 0.32—0.45 cm. long, minutely papillate dor- sally; ovary ovoid, about 0.7 em. long, glabrous; stigma sub- eapitate, broadly oblongoid, 0.08-0.1 cm. long; style 0.25- 0.3 em. long, deciduous at the insertion to the ovary ; nectaries compressed-ovoid, separate, much shorter than the ovary ; fol- licles relatively slender, obseurely undulate-articulate or es- sentially continuous, tortuose to somewhat divaricate, fre- quently more or less falcate, acuminate, 15-23 cm. long, gla- brous; seeds 0.7-1.0 em. long, the pale yellowish-ashy coma 1.5-1.8 em. long. UNITED STATES: DELAWARE: moist thicket, borders of cedar swamp, near Col- lins Beach, July 6, 1874, Commons s.n. en MBG, NY, US); New Castle City, on borders of a cedar swamp, Aug. 23, 1865, Commons s.n. (NY, US); VIRGINIA: in fruticetis pr. Portsmouth, LANA, 1840, Rugel s.n. (BB); Virginia Beach, July 3, 1892, Britton Britton $ Vail s.n. (NY); NORTH CAROLINA: Burgaw, June, 1880 Hyams s.n. (MBG); ‘‘middle North Carolina,’’ May 28, year he Ashe 2354 (NY, US); near Gaston, Northampton Co., June 15, 1895, Ward s.n. (US); Eden- ton, Chowan Co., July 29-30, 1898, Kearney 1874 (US); SOUTH CAROLINA: Aiken, date lacking, Ravenel s.n. (FM); near Charleston, 1865, Stewart s.n. (FM); Bluff- ton, date laeking, Mellichamp s.n. (NY, ); GEORGIA: edge of Ogeechee River 4 , Burke Co., date lacking, Chapman s.n. (MBG); Rome, date laeking, Chapman s.n. (US); vicinity of Louisville, date lacking, Hopkins s.n. (NY); FLORIDA: bank of Apalach- icola River, near Chattahoochee, June 16, 1897, Curtiss 5893 (FM, MBG, US); lola, date lacking, Chapman s.n. Po ; near Tallahassee, summer, year lacking, Berg n. (NY); ALABAMA: low wooded banks, Cullman, June, 1891, Mohr s.n. (MBG); bass A: G. B. B. R. Valley Head, July, 1898, Ruth 468 (MBG, NY, US); woods, (269) [VoL. 22 236 ANNALS OF THE MISSOURI BOTANICAL GARDEN near Attalla, Etowah Co., June 30, 1897, Eggert s.n. (MBG); near Tuscaloosa, date lacking, Johnson s.n. (NY); Calera, date lacking, Everts s.n. (NY); Pratt Mines, Aug. 16, 1879, Mohr s.n. (US); Busby's Ford, Walker Co., Sept. 10, 1879, Mohr s.n. (US); Gadsden, 1878, Vasey s.n. (FM, US); banks of Alabama River, Chestang's Bluff, Sept. 1, 1879, Mohr s.n. (US); Cedar Plains, swampy copses, May 21, 1891, Mohr $ Sudworth s.n. (US); MISSISSIPPI: Centerville, Aug. 4, 1897, Tracy 5468 (MBG, NY); Jackson, May 23, 1888, Tracy s.n. (NY); Taylorville, Aug. 23, 1903, Tracy s.n. (US); Brookhaven, April 25, 1882, Flint s.n. (US); fie diteh, Forest, May 26, 1925, nee s.n. (US); LOUISIANA: near New Orleans, date er Ingalls s.n. (NY); e data, Drummond s.n. (Camb.) ; Atakapas, May, 1883, Langlois s.n. (FM); an June 11, 1915, Paini 7958 (MBG); East u. Rouge Par., May 20, 1874, Joor s.n . (FM); TEXAS: Polk, Bowie Co., June 13, 1898, Eggert s.n. (MBG); near Howien, April, 1842, Lindheimer s.n. (MB G); Big Sandy, c in swamp, June 27, 1901, Reverchon 2558 (MBG); climbing on small treen Montgomery Co., July 18-21, 1909, Dixon 491 (FM, NY); Harrisburg, July 14, 1876, s.n. (US); ILLINOIS: low woods bordering swamps, Karnak, do den te, b 71 1919, Palmer 16542 (MBG, US); INDIANA: frequent on the high bank of a slough in the ‘‘bottoms,’’ about % mi. southeast of Yankeetown, rare, Aug. 13, 1922, Deam 37581 (NY); TENNESSEE: low woods, Haywood Co., June, 1893, Bain 428 (NY); Covington, June, 1889, Byars s.n. (US) ; MISSOURI: border of Creve Coeur Lake, St. Louis Co., February, 1931, Woodson s.n. (MBG) ; swamps, Campbell, Sept. 7, 1910, Bush 6238 (MBG, NY); in Sümpfen, Dunklin Co., July 27-Sept. 26, 1893, Eggert s.n. (MBG); swamps, Butler Co., Oct. 17, 1905, Bush 3742 (MBG, NY, US); between bii at [Ark.] and Kennett, July 27, 1893, Eggert s.n. (FM, MBG, NY, US); low wet woods, Neeleyville, Butler Co., Sept. 16, 1919, Palmar 16460 (MBG, UB); thiekets, fence-rows, etc., Kennett, Dunklin Co., June 8, 1930, Kellogg 15263 (MBG); ARKANSAS: low rocky banks of Ouachita Bv; near Hot Springs, Garland Co., Oct. 11, 1925, Palmer 29164 (MBG); thiekets along small rocky ereek, near Hot Springs, Garland Co., May 9, 1925, Palmer 27116 (MBG); low woods, Corning, Clay Co., a 2 1914, Palmer 6063 (MBG); low open ground, Arkadelphia, Clark Co., Jun ‚1915, Palmer 8089 (MBG); Little Rock, 1881, Letterman s.n. (MBG); froda Pott, Sept. 24, 1909, Kellogg s.n. (MBG); climbing on bushes, Judsonia, June 12, 1877, Reynolds s.n. (FM, US); swamps, Moark, Oct. 17, 1905, Bush 3751 (MBG); Paragould, Sept. 26, 1893, Eggert s.n. (MBG); low woods, Greene Co., July 27, Sept. 26, er Eggert s.n. (MBG, NY); ; along Bridge Creek, near MeNab, Hempstead Co., 1923, Greenman 4316 (MBG); St. Franeis Club (w. of Memphis), April 5, t Trelease s.n. (MBG); wet sandy dp Gum Springs, Clark Co. ied 21, 1916, Palmer 10537 (MBG) ; Conway, May 15, 1923, Wheeler s.n. (FM); common, wooded I1 Pike Co., Sept. 29, 1932, dl 9392 (MBG); in CLP BUR rubble on sed mountain eni Arkan nsas National Forest, Sebastian Co., May 24, 1931, Palmer densa (M ; 4 mi, southeast of Prescott, P 7, 1912, Hollister 46 (US); OKLAHOMA: at sri ja a pond, Talihina, June 19, 1898, Glatfelter s.n. (MBG); Kiamiehi River, LeFlore Co., June, 1927, bct s.n. (US). The striking coextension of the ranges of T. difforme and Taxodium distichum, as symbolic of the characteristic flora of (270) iis WOODSON—STUDIES IN THE APOCYNACEAE. IV 237 the ancient Cretaceous strand, also indicated, is illustrated in text-fig. 2. The most remarkable extension of the species at present recorded is from the banks of Creve Coeur Lake, St. Louis County, Missouri, where a single plant with several mature follicles containing viable seeds was found by the writer during February, 1931. Although in winter condition, the plant was proved correctly identified by the germination — T T = SS / ) f aa, E S R $ ¿A NAAA AS xe be V ; LES AE We Y ! PANA A ie CARA Cl - e ^ DURS > x + AS ; eee f. AS t ^ Y END HA Ya A v = / M y A 4 MO x ^ 4 NY 5 ( i : np NE to EN u > pes NS L — sa EV.“ SEP A n AS J f 4 vd I~ s E y Cf ‘Sal j ! Are 7 É ) AM - eqq E i \ ER —-— £s i | q NA 7 LT x ais 7 | Li. > e DEN i 2 j Nk IN x | e \ - / Am wot 7 WEG EXC f | . e de ES [2 / e ; ? N A A ^ / / e d " Y ` S | E y + d Fig. 2. Solid line: approximate shore-line of the Middle Late Cretaceous period in the southeastern United States (after Schuchert & Dunbar); broken approximate northern limit of Tazodium distichum (after Berry); dots: line: Explanation in the text. known distribution of Trachelospermum difforme and subsequent growth of seedlings from the fruit. Creve Coeur Lake is one of the familiar **ox-bow"' lakes of the Mis- souri River a few miles west of its junction with the Missis- sippi. Although repeated search has been made, particularly by local botanists of the vicinity of Saint Louis including the original finder, no further evidence of plants has as yet been found. This station is more than 125 miles north of the low- (271) | [VoL. 22 238 ANNALS OF THE MISSOURI BOTANICAL GARDEN land of Missouri south of the ancient Cretaceous shore-line of the Mississippi embayment (popularly known by the inhabi- tants of the district as the ‘‘ Poplar Bluff’’ for the occurrence of Liriodendron tulipifera) where the species is common, to- gether with Taxodium distichum and the associated southeast- ern Atlantic coastal plain flora. Additional interest in the iso- lated occurrence of T. difforme in the lower Missouri River Valley is found in persistent reports of logs of Taxodium washed from time to time by the unruly river from the border- ing beds of clay and loess. Although the writer has never been so fortunate as to see actual specimens of such ancient wood, it is generally believed that the former distribution of T. distichum in North America was much wider than at present. Although the flowers and inflorescence of T. difforme are rather uniform throughout the distribution of the species, vegetative characters are found to be variable. The leaves are variable in shape and size, and the presence, or relative abun- danee, of the foliar indument is manifestly inconstant. Par- tieularly striking are occasional plants with very narrowly elliptie to almost linear foliage which are found toward the southwestern limits of the species. In this area, however, are also found plants bearing leaves which are among the broadest of the species. The outline and width of foliage also have been found inconstant upon single specimens. Consequently it has been considered inadvisable to erect varieties for the expres- sion of such fluctuations of vegetative characters. EXCLUDED SPECIES Trachelospermum stans A. Gray, Proc. Am. Acad. 21: 394, 1886 = Mandevilla foliosa (Muell.-Arg.) Hemsl. Biol. Centr.- Am. Bot. 2: 316. 1882 (Amblyanthera foliosa Muell.-Arg. Lin- naea 30: 427. 1860). VIII. Matoverta A. DC. Malouetia A. DC. in DC. Prodr. 8: 378. 1844; Muell.-Arg. in Mart. Fl. Bras. 61: 89. 1860; Benth. € Hook. Gen. Pl. 2: 708. 1876; Miers, A poc. So. Am. 86. 1878; K. Sch. in Engl. € Prantl, Nat. Pflanzenfam. 4?: 186. 1895. (272) er WNOTT S lo. cial d »* ><] 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 239 Robbia A. DC. loc, cit. 444. 1844; Miers, loc. cit. 107. 1878; K. Sch. loc. cit. 187. 1895. Lactescent shrubs or small trees. Stems ligneous, erect, terete, the branches dichotomous or opposite, rarely alternate. Leaves opposite, entire, penninerved, the upper surface eglan- dular, the lower surface bearing with more or less frequency a single lenticular or pustulate foveum or pit in the axils of the midrib and primary veins; nodes exstipulate or virtually so. Inflorescence terminal, occasionally lateral as well, umbellate, few- to several-flowered. Calyx 5-parted, the lobes essentially equal, cleft nearly to the receptacle, bearing within alternate, solitary, or rarely geminate, glandular squamellae. Corolla salverform, the tube relatively short in certain species, cylin- drical to flask-form, exappendiculate within, the orifice con- spieuously callose-thiekened! in certain species, the limb acti- nomorphie, 5-parted, dextrorsely convolute. Stamens 5, the anthers almost completely exserted to wholly included in some species, connivent and agglutinated to the stigma, consisting of 2 parallel, basally protuberant sporangia borne ventrally near the apex of an enlarged, sagittate, peltate connective; pollen granular; filaments free. Carpels 2, apocarpous, united at the apex by a common stylar shaft surmounted by the fusiform- subcapitate stigma ; ovules numerous, 2-4-seriate, borne upon an axile, binate placenta. Nectaries 5, separate or more or less conerescent. Follicles 2, occasionally solitary by abortion, apocarpous, narrowly terete to broadly fusiform, usually more or less divaricate or falcate, dehiscing along the ventral suture; containing numerous dry, ecomose, glabrous or generally pu- bescent seeds. Type species: Malouetia Tamaquarina (Aubl.) A. DC. loc. cit. 378. 1844. The question relative to the congenericity of Malouetia and Robbia is scarcely nearer solution than in 1860 when the two were combined by Mueller, whose position was later endorsed by Bentham. Miers and Schumann maintained the generic status of Robbia; while more recently the broader interpreta- ! Cf. p. (291), et seq. (273) [Vor. 22 240 ANNALS OF THE MISSOURI BOTANICAL GARDEN tion of the plexus -— been reaffirmed by Markgraf (in Pulle, Fl. Surinam 4: 56. 1932). The distinction rs ve two genera evidently rests solely upon the character of the seeds, whether glabrous, as in typical Malouetia, or pubescent, as in the species referred to Robbia by de Candolle. No other distinguishing characters have been advanced by any competent student of the genus. Although such a character as that of the seeds is doubtless of genuine bio- logical significance, it can scarcely be interpreted as more so than, for example, the absence of latex in Asclepias tuberosa, which would properly be frowned upon as the basis of a generic segregation largely on the basis of practice. Similarly in the instance of Robbia and Malouetia, fruit is so rare in herbarium collections that separation of the genera upon that single character would prove extremely difficult in the absence of ad- ditional criteria. For a number of species, fruit is unknown at the present writing. Intergradation of the seminal indument of Robbia further weakens the claim of that character as of generic, or even sub- generic or sectional value. The seeds of Malouetia Tamaquar- ma (Aubl) A. DC. are absolutely glabrous: those of M. furfuracea Muell.-Arg. are very sparsely villous. Yet these two species are scarcely separable in flowering material, and are indigenous to an almost identical area. The seeds of M. ces- troides (Nees) Muell.-Arg., not distantly related to the pre- ceding, are densely lanate-villous. In view of our imperfect understanding of the group, the most practical policy would appear to be the inclusion of Robbia within Malouetia. KEY TO THE SECTIONS A. Anthers conspicuously exserted (except in M. Mexiae and M. cestroides), inserted near the orifice of the corolla-tube......... ect. 1. TAMAQUARINAE AA. Anthers included, or essentially so, inserted about midway, or deeper, within the eorolla-tube.......... ccc cece cece nne Sect. 2. GRACILES Sect. 1. Tamaquarinaz Woodson. Corolla-tube cylindrical to broadly flask-form, gradually narrowing toward the inser- tion of the stamens; anthers ovate to elliptic, conspicuously ex- (274) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 241 serted (essentially included, or barely exserted in M. Mewiae and M. cestroides), inserted near the orifice of the corolla-tube. Spp. 1-15. KEY TO THE SPECIES a. Calyx-lobes delicate and foliaceous in texture, not closely imbricated at anthesis, spreading; corolla-tube flask-form, eonspieuously inflated below the middle. b. Anthers conspicuously exserted; eorolline faueal tube 0.05-0.075 em. long. e. Corolla without a eallose faueal annulus ;* plants of the "F Amazon CESTE) cx ree A A TERN . M. flavescens ec. Corolla with a definite faueal annulus d. Corolla-tube 0.8-1.3 em. long; ab -lobes laneeolate to linear-lance- olate, 0.25—0.5 cm. long. e. oie 0.8-1.0 em. long, the lobes 0.9-1.2 em. long; faucal nnulus relatively indefinite, consisting of numerous minute, MIS. protuberances; plants of the Antilles (St. Erw Cuba?) . cubana ee. Corolla-tube 1.1-1.3 em. long, the lobes 1.2—1.6 em. a ; faucal annulus conspicuous, 5-lobed; plants of British Guiana........ PPP vue em Et 3. M. Schomburgki dd. Corolla-tube 0.6-0.7 em. long; calyx-lobes ovate to ovate-lanceolate, 0.15-0.25 em. long; plants of the lower Amazon Valley ( eo o ooos$onn.................. oo... ......oo.. ooo bb. zd E barely surpassing or attaining the corolline orifice; faucal .25 em. long; plants of the Amazon Valley be and (ML M aa. Calyx-lobes relatively fleshy and coriaceous in texture, closely isi at an thesis; corolla-tube cylindrical, or only somewhat flask-f in certain spee b. Corolla Se delieate in texture, the tube 0.4-0.9 em. long. e. Leaves wholly glabrous. d. Anthers barely pem or the tips barely exserted...... 6. M. cestroides dd. Anthers conspicuously exserted. Leaves membranaceous, conspicuously foveate in the axils of the midrib; plants of southeastern Brazil (Rio de Janeiro, Minas Gernes, Sho Panlo). wm sees Dunn 7. M. arborea ee. Leaves subcoriaceous, inconspicuously and rarely foveate in the of the midrib; Lage: of northern Brazil (Para, ner "e en Colonia sahen . M. Duckei ee. Leaves softly puberulent nci Lg OU OS O. 9. E pubescens bb. Cor en edd anc. in texture, the tube 0.9—1.5 em. long. e. Leaves wholly glabrou d. Leaves definitely imr e. Corolla-lobes puberulent within, at least toward the base. * Cf. p. (291), et seq. (275) [Vor. 22 242 ANNALS OF THE MISSOURI BOTANICAL GARDEN f. Calyx-lobes 0.2-0.3 em. long, 4-4 (rarely %) as long as the corolla-tube, obtuse, occasionally broadly acute; seeds glabrous SAC EV OREO ISS a4 00S EN AEREOS 0. M. Tamaquarina ff. Calyx-lobes 0.3—0.5 em. long, 4—% as long as the corolla-tube, acute to acuminate; seeds sparsely villous...... 11. M. furfuracea ee. Corolla-lobes about uniformly papillate to glabrate within and without. f. Leaves oblong-oblanceolate, obtuse or rounded; eorolla-tube 0.9 2. M LELO mu, DAB. ara teris 13. M. peruviana dd. Leaves sessile or subsessile, and more or less amplexicaul.......... TELTET rA 4461214 IAE 14. M. amplexicaulis ce. Leaves softly and minutely puberulent beneath............. 15. M. Killipü 1. Malouetia flavescens Muell.-Arg. in Mart. Fl. Bras. 6*: 95. 0 60. Small shrubs (fide Ducke); stems relatively slender, gla- brous, reddish-brown, very inconspieuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-ellip- tic, subeaudate-aeuminate, base obtuse, 7-11 cm. long, 2.5- 4.5 cm. broad, membranaceous, glabrous throughout, light olivaceous in desiccation, somewhat subnitidulous above, opaque beneath, infrequently and obscurely foveate in the axils of the midrib; petioles 0.1-0.3 em. long, glabrous; umbels terminal, few-flowered; peduncle 0.1-0.3 em. long, glabrous; pedicels 1.8-2.0 em. long, somewhat accrescent in fruit, gla- brous; ealyx-lobes lanceolate, acuminate, spreading at anthesis, not elosely imbricated, 0.3-0.35 em. long, rather delicately foli- aceous, puberulent-papillate within, minutely papillate with- out; corolla broadly flask-form, greenish, the tube 1.0-1.1 em. long, about 0.15 cm. in diameter at the base, dilated to about 0.3 em. in diameter below the middle, faucal tube 0.075 em. long, about 0.175 em. in diameter at the orifice, indefinitely papillate to essentially glabrate without, minutely puberulent generally within, the lobes obliquely oblong-lanceolate, long-acuminate, 1.1-1.2 em. long, about 0.25 em. broad, sharply reflexed, the tips usually more or less turbinate, minutely papillate without, minutely puberulent toward the base within; anthers conspicu- ously exserted, ovate-elliptie, 0.3-0.325 em. long, minutely pu- berulent dorsally; ovary about 0.1 em. long, minutely puberu- (276) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 243 lent; stigma 0.15 em. long; nectaries less than half equalling the ovary; follicles terete, 18-20 cm. long, about 0.4 cm. in diameter, falcate, glabrous; seeds 3-4 em. long, 0.2-0.3 em. in diameter, glabrous. RAZIL: PARA: ad cataractam infimam loco Maria Luiza, silva, Rio Tapajoz, Sept. 27, 1922, Ducke 11394 (B, S, US). The type specimen of this species (Hoffmannsegg s.n. in herb. Willdenow, fide Muell.-Arg.) has not been available for examination. However, the specimen cited agrees so well with the satisfyingly complete original description that its relega- tion to M. flavescens appears to be well substantiated. 2. Malouetia cubana A. DC. in DC. Prodr. 8: 379. 1844; Miers, Apoc. So. Am. 92. 1878. Malouetia retroflexa Muell.-Arg. Linnaea 30: 408. 1860. Stems relatively slender, glabrous, reddish-brown, inconspic- uously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptie, apex shortly and obtusely subcau- date-acuminate, base obtuse to broadly acute, 5-11 em. long, 1.54.5 em. broad, membranaceous, glabrous throughout, oliva- ceous in desiccation, somewhat subnitidulous above, opaque beneath, minutely foveate in the axils of the midrib; petioles 0.15-0.5 em. long; umbels terminal, few-flowered; peduncle 0.2-0.4 cm. long; pedicels 2.0-2.3 cm. long, glabrous; calyx- lobes lanceolate to ovate-lanceolate, acuminate, spreading at anthesis, not closely imbricated, 0.3-0.45 em. long, rather deli- cately foliaceous, glabrous without, minutely and sparsely pu- berulent-papillate toward the tips within; corolla broadly flask-form, greenish-white (?), the tube 0.8-1.0 cm. long, about 0.1-0.15 em. in diameter at the base, dilated to about 0.2- 0.25 em. below the middle, faucal tube 0.05 cm. long, orifice about 0.15 em. in diameter, faucal annulus relatively indefinite, consisting of numerous minute, callose protuberances, glabrous without, very minutely puberulent within, the lobes obliquely laneeolate to ovate-lanceolate, acuminate, 0.9-1.2 em. long, about 0.2 cm. broad, sharply reflexed, the tips usually more or less turbinate, minutely papillate without, minutely puberu- lent toward the base within; anthers conspicuously exserted, (277) [Vor. 22 244 ANNALS OF THE MISSOURI BOTANICAL GARDEN ovate-elliptie, 0.3-0.325 em. long, minutely puberulent dorsally ; ovary about 0.125 cm. long, minutely puberulent ; stigma 0.125- 0.15 em. long; nectaries about 14 equalling the ovary ; follicles unknown. St. VINCENT: exact locality lacking, 1822, Guilding s.n. (K, MBG, photograph and analytieal drawings). I am strongly inclined to the opinion expressed by Miers (loc. cit. 1878) that the original descriptions of M. cubana and M. retroflexa can scarcely be interpreted as applying to dis- tinet species. The type specimen of the former, de la Ossa s.n., should presumably be deposited in the herbarium of de Can- dolle, but it was not found during my examination of the col- lection in 1930. At any rate, the species has evidently not been collected in Cuba since the discovery of the type. Ambiguity likewise surrounds the present status of M. retroflexa in St. Vincent. Beside the type colleetion, made over a hundred years ago, a specimen with only the data '' Echites guianensis; West-Indien,”” has been found in the herbarium at Vienna, and another, presumably a duplicate of the type, at Berlin with the notation **St. Vincent; Lindley misit 1827.’ 3. Malouetia Schomburgki Muell.-Arg. Linnaea 30: 409. 1860; Miers, Apoc. So. Am. 88. 1878. Shrubs or small trees, 1.8-3.0 m. tall (fide de la Cruz) ; stems relatively slender, glabrous, reddish-brown, inconspicuously lenticellate when fully mature; leaves opposite, petiolate, ob- long- to ovate-elliptie, apex obtusely subcaudate-acuminate, base obtuse to broadly acute, 4.5-13.0 em. long, 2.0-5.5 em. broad, firmly membranaceous to somewhat subcoriaceous, gla- brous throughout, light olivaceous in desiccation, nitidulous above, opaque, minutely foveate in the axils of the midrib be- neath; petioles 0.2-0.5 em. long, glabrous; umbels terminal, bearing several white (or brownish ?) flowers; peduncle 0.2- 0.5 em. long; pedicels 0.15-0.3 em. long, glabrous; ealyx-lobes lanceolate to linear-lanceolate, acuminate, 0.25—0.5 em. long, glabrous to very minutely papillate without, minutely puberu- lent-papillate toward the tips within, rather delicately foli- aceous, spreading at anthesis, not closely imbricated; corolla (278) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 245 broadly flask-form, the tube 1.1-1.3 em. long, about 0.1- 0.125 em. in diameter at the base, dilated to about 0.25-0.3 em. below the middle, faucal tube 0.05-0.075 em. long, about 0.1— 0.15 em. in diameter at the orifice, faucal annulus conspicuous, 9-lobed, glabrous without, minutely puberulent within, the lobes obliquely oblong-lanceolate, acuminate, 1.2-1.6 cm. long, 0.175-0.25 cm. broad, sharply reflexed, the tips usually more or less turbinate, minutely papillate or puberulent-papillate with- out, puberulent toward the base within; anthers conspicuously exserted, ovate-elliptic, 0.25—0.3 em. long, minutely puberulent dorsally; ovary about 0.1 cm. long, minutely puberulent; stigma 0.1-0.125 cm. long; nectaries 14—Y equalling the ovary; follicles terete, 16-36 cm. long, 0.3-0.4 em. in diameter, falcate, glabrous; seeds 2.5-4.0 cm. long, glabrous. BRITISH GUIANA: Ufer der fl. Pomeroon, Aug., 1843, Schomburgk 1386 (B, SYNTYPE, MBG, photograph); Waini River, Northwest Distriet, Sept. 13, 1921, Cruz 1116 (G, NY, US); Cart Market, Moruka River, Pomeroon Distriet, Sept. 20, 1921, Cruz 1157 (NY, US); Anabisi River, Northwest District, Febr. 15, 1922, Cruz 1360 (FM, NY, US); upper Rupununi River, near Dadanawa, June 3, 1922, Cruz 1476 (MBG, NY, US); vicinity of Bartiea, Essequibo River, Sept. 3-12, 1922, Cruz 1874 (FM, G, MBG, NY, US); Kabakaburi, Pomeroon District, Febr. 10-15, 1923, Cruz 3242 (FM, MBG, NY, US); Wanama River, Northwest Distriet, May 10-23, 1923, Cruz 3935 (MBG, NY, US); Kamakusa, upper Mazaruni River, July 11-12, 1923, Cruz 4232 (MBG, NY, US); data incomplete, Schomburgk 830 (V, PARATYPE). 4. Malouetia lata Mgf. Notizblatt 10: 1037. 1930. Shrubs (fide Ducke) ; stems relatively stout, glabrous, red- dish-brown, inconspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, broadly oblong- to ovate-el- liptie, apex obtusely subcaudate-acuminate, base obtuse to rounded, 5-12 cm. long, 3-7 cm. broad, rather delicately mem- branaceous, glabrous throughout, olivaceous in desiccation, slightly nitidulous above, opaque, very inconspicuously foveate in the axils of the midrib beneath; petioles 0.2-0.5 em. long, glabrous; umbels terminal, bearing relatively few, greenish- white flowers; peduncle 0.1-0.3 em. long; pedicels 0.7-0.8 cm. long, glabrous; calyx-lobes ovate to ovate-lanceolate, acumi- nate, 0.15-0.25 cm. long, spreading at anthesis, not closely im- brieated, rather delicately foliaceous in texture, glabrous with- (279) [Vor. 22 246 ANNALS OF THE MISSOURI BOTANICAL GARDEN out, minutely and rather sparsely puberulent-papillate at the tips within; corolla broadly flask-form, the tube 0.6-0.7 cm. long, about 0.1 cm. in diameter at the base, dilated to 0.25- 0.3 em. below the middle, faucal tube 0.05 cm. long, about 0.125 em. in diameter at the orifice, faucal annulus conspicuous, 9-lobed, without minutely papillate below, otherwise glabrous, within minutely puberulent to puberulent-papillate, the lobes obliquely oblong-elliptic, acuminate, 0.7-0.8 em. long, sharply reflexed, minutely papillate without, minutely puberulent to- ward the base within; anthers conspicuously exserted, ovate- elliptic 0.25 em. long, minutely puberulent dorsally; ovary about 0.1 em. long, minutely puberulent; stigma 0.1 em. long; nectaries about Ya equalling the ovary; follicles terete, 40- 45 em. long, 0.4—0.5 em. in diameter, faleate, glabrous; seeds 0.35—0.4 em. long, about 0.2 em. in diameter, glabrous. BRAZIL: PARA: Parana de Juruty Velho, silva ab Amazonium fluvio periodice inundata, Febr. 26, 1926, Ducke 21591 (B, TYPE, S, US, MBG, photograph and analytieal drawings); Caeaval Imperial prope Obidos, silva ab Amazonium fluvio periodiee inundata, Sept. 12, 1910, Ducke 21784 (B); AMAZONAS: Rio Antimary (Purus), silva paludosa, April 1, 1904, Huber 21764 (B). 9. Malouetia Mexiae Woodson, spec. nov. Arbuseulae vel frutices; ramulis glaberrimis cortice atro- brunneis maturitate inconspicue lenticellatis; foliis oppositis breviter petiolatis oblongo-ellipticis apice obtuse subcaudato- acuminatis basi obtusis 6-12 em. longis 0.15-0.4 em. latis cori- aceis glaberrimis supra subnitidulis subtus opacis in axillis nervi medii inconspicue foveatis; petiolis 0.2-0.4 em. longis glabris; umbellis terminalibus flores 4—10 pallide viridulas gerentibus; pedunculo 0.2-0.5 em. longo; pedicellis 1.5-2.5 em. longis glaberrimis; calycis laciniis ovatis vel ovato-lanceolatis acuminatis 0.25—0.35 em. longis tenuiter foliaceis ex apice patu- lis haud imbricatis extus minute papillatis intus minute puber- ulis; corollae ampulliformis tubo 0.9-1.2 em. longo basi ca. 0.1- 0.15 em. diametro metiente deinde sub medio usque 0.275- 0.35 em. dilatato faucibus tubularibus 0.2-0.25 em. longis ostio ca. 0.15 em. diametro metiente conspicue calloso-annulato extus glabro intus prope insertionem staminum puberulo-papillato lobis anguste et oblique elliptieo-oblanceolatis acuminatis 0.9— (280) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 247 1.4 em. longis 0.25—0.3 em. latis patentibus extus papillatis intus prope basem puberulis; antheris inclusis vel paululo exsertis ovato-elliptieis 0.25—0.3 em. longis dorso minute puberulis puberulo-papillatisve; ovario ca. 0.1 em. longo minutissime pu- berulo; stigmate 0.1 em. longo; neetariis ovario ca. dimidio brevioribus ; folliculis ignotis. Shrubs or small trees; stems glabrous, reddish-brown, in- conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptic, apex obtusely subcaudate- acuminate, base obtuse, 6-12 em. long, 0.15—0.4 em. broad, cori- aceous, glabrous throughout, above subnitidulous, beneath opaque, inconspicuously foveate in the axils of the midrib; pet- ioles 0.2-0.4 cm. long; umbels terminal, bearing 4-10 pale greenish flowers; peduncle 0.2-0.5 em. long; pedicels 1.5- 2.5 cm. long, glabrous; calyx-lobes ovate to ovate-lanceolate, acuminate, 0.25-0.3 cm. long, delicately foliaceous, spreading at anthesis, not closely imbricated, minutely papillate without, within minutely puberulent ; corolla-tube flask-form, 0.9-1.2 em. long, about 0.1-0.15 cm. in diameter at the base, dilated to 0.275—0.35 em., faucal tube 0.2-0.25 em. long, about 0.15 cm. in diameter at the orifice, faucal annulus conspicuous, 5-lobed, the lobes narrowly and obliquely elliptic-oblanceolate, acumi- nate, 0.9-1.4 cm. long, 0.25-0.3 cm. broad, sharply reflexed, without minutely papillate, puberulent toward the base within; anthers ovate-elliptie, 0.25-0.3 em. long, included, or the tips barely exserted, minutely puberulent to puberulent-papillate dorsally; ovary about 0.1 em. long, very minutely puberulent ; stigma 0.1 cm. long; nectaries about 1% equalling the ovary; follieles unknown. BRAZIL: PARA: Thomé Assu, Distrieto Acara, Rio Acara, overflow bank in dense shade, July 28, 1931, Mexia 5994 (MBG, TYPE); inter Mosquiero et Carananduba, in ripis inundatis fluvii Para, May 13, 1923, Ducke 17465 (B); ad ripas inundatas fluminis Jaburuzinho, regione Breves, aestuarii amazonici, July 12, 1923, Ducke 17466 (B); Cameta, silvula secundaria, Sept. 13, 1903, Siquiera 21771 (B); AMAZONAS: Rio Negro infer., loco Terra Preta, silva paludosa, Dec. 31, 1923, Kuhlmann 21868 (B). Sharply differentiated from its relatives of the Schomburgki plexus by the included or barely exserted anthers, a position caused by the extension of the faucal tube of the corolla. The (281) [Vor. 22 248 ANNALS OF THE MISSOURI BOTANICAL GARDEN texture of the leaves, in addition, is more heavy than that of its related eongeners. Siquiera 21771 and Kuhlmann 21868 are placed within this species somewhat doubtfully because of a lack of fully expanded floral buds. 6. Malouetia cestroides (Nees) Muell.-Arg. in Mart. Fl. Bras. 61: 94. pl. 29. fig. 2. 1860. Tabernaemontana cestroides Nees, ex Mart. Nov. Act. Nat. Cur. 11: 83. 1823. * Robbia cestroides (Nees) A. DC, in DC. Prodr. 8: 445. 1844; Deles. Icon. 5: pl. 52. 1846; Miers, Apoc. So. Am. 107. 1878; K. Sch, in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Shrubs or small trees; stems relatively stout, glabrous, red- dish-brown, inconspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong- to ovate-elliptie, apex obtusely acuminate, base obtuse to broadly acute, 6— 12 em. long, 2.0-4.5 em. broad, firmly membranaceous, glabrous throughout, above somewhat nitidulous, beneath opaque, mi- nutely foveate in the axils of the midrib; petioles 0.2-0.6 cm. long; umbels terminal, relatively few-flowered; peduncle 0.2- 0.4 em. long; pedicels 0.8-1.0 em. long, glabrous; calyx-lobes ovate, obtuse to rounded, occasionally broadly acute, 0.1- 0.125 em. long, relatively thick in texture, closely imbricated at anthesis, glabrous without, papillate toward the tips within; corolla salverform, only slightly inflated toward the base, the tube 0.75-0.8 em. long, about 0.075-0.1 em. in diameter at the base, faucal tube 0.125-0.15 em. long, about 0.1 em. in diameter at the orifice, faucal annulus conspicuous, 5-lobed, glabrous without, minutely puberulent above within, the lobes obliquely elliptic-lanceolate, acuminate, 0.6-0.8 em. long, reflexed, papil- late without, minutely puberulent toward the base within; anthers barely included or the tips barely exserted, ovate-ellip- tic, 0.2 em. long, very minutely pilosulose dorsally ; ovary about 0.15 cm. long, minutely puberulent; stigma 0.1 cm. long; nec- taries about Y2 equalling the ovary; follicles terete, 11-12 cm. long, about 0.5 em. in diameter, somewhat divaricate, glabrous; seeds 2.5-3.0 em. long, densely lanate-villous. (282) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 249 BRAZIL: BAHIA: Vittoria, date lacking, Sello s.n. (B); Nazareth, date lacking, Sello s.n. (B); data ineomplete, Blanchet 1738 (MBG); Blanchet 1578 (MP); RIO DE JANEIRO: Gavea, Jan. 3, 1872, Glaziou 5934 (B, NY); MINAS GERAES: data incomplete, Aug.-April, 1840, Claussen s.n. (Camb.); DATA INCOMPLETE: Martius 105 (Bx, TYPE, V, MBG, photograph) ; Riedel s.n. (B, V) ; Regel s.n. (Camb.). T. Malouetia arborea (Vell.) Miers, Apoc. So. Am. 89. 1878. Echites arborea Vell. Fl. Flum. 114. 1830; Icon. 3: pl. 47. 1827. Tabernaemontana laeta Marts according to A. DC. in DC. Prodr. 8: 364. 1844, as to synonymy, in part. Malouetia Tamaquarina A. DC. y. Brasiliensis A. DC. loc. cit. 379. 1844. Malouetia lanceolata Muell.-Arg. in Mart. Fl, Bras. 6!: 93. pl. 29. fig. 3. 1860; Miers, loc. cit. 88. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Malouetia Martii Muell.-Arg. loc. cit. 94. pl. 29. 1860; Miers, loc, cit. 90. 1878. Secondatia ? arborea (Vell) Muell.-Arg. loc. cit. 110. 1860. Robbia gossipina Miers, loc. cit. 108. pl. 12. fig. B. 1878. Small trees (12 m. tall, fide Kuhlmann); stems relatively stout, glabrous, reddish-brown, rather inconspicuously lenticel- late when fully mature; leaves opposite, petiolate, oblong-ellip- tie, apex obtusely subcaudate-acuminate, base obtuse to broadly acute, 4-12 em. long, 1.3-4.0 em. broad, membrana- ceous, glabrous throughout, above somewhat subnitidulous, beneath opaque, conspicuously foveate in the axils of the mid- rib; petioles 0.3-0.5 cm. long; umbels terminal, bearing several small, white flowers; peduncle 0.2-0.4 em. long; pedicels 0.9- 1.0 em. long, glabrous; calyx-lobes ovate, acute to obtuse, 0.15- 0.2 em. long, relatively thick in texture, closely imbricated at anthesis, without glabrous to minutely papillate, within papil- late at the tips; corolla salverform, only slightly inflated at the base, the tube 0.7—0.8 em. long, about 0.1 em. in diameter at the base, faucal tube 0.05—0.1 em. long, about 0.1-0.125 em. in diam- eter at the orifice, faucal annulus conspicuous, 5-lobed, without glabrous or rarely very minutely papillate, within very sparsely and minutely puberulent above, the lobes obliquely (283) [Vor. 22 250 ANNALS OF THE MISSOURI BOTANICAL GARDEN elliptic-lanceolate, acuminate, 0.8-0.95 em. long, 0.25-0.3 cm. broad, reflexed, without minutely papillate, within minutely puberulent toward the base; anthers conspicuously exserted, ovate-elliptie, 0.2 em. long, minutely puberulent dorsally ; ovary 0.1 em. long, minutely puberulent; stigma 0.08-0.1 em. long; nectaries about 1% equalling the ovary; follicles terete, 9— 14 em. long, 0.7-0.8 em. in diameter, divaricate, glabrous; seeds 3.5-4.0 em. long, densely lanate-villous. LÀ BRAZIL: RIO DE JANEIRO: S. Luiz, Aug. 9, 1867, Glaziou 1989 (MP); Gavea, Jan. 3, 1872, Glaziou 5934 (MP); Fazenda de Sta. Cruz, Aug. 8, 1877, Glaziou 9511 (B, MP); Careadura, Dec. 23, 1869, Glaziou 4077 (B, MP); ad urbem in silvis .. do Fonseca, Jan. 3, 1922, Occhioni 4459 (B, MP); Itatiaia, Dec., 1918, Porto 8671 (B); ad urbem in loeo Sta. Theresa, silva secundaria, Dec. 15, 1925, nn 2187? (B); data inedmplsbé, Weddell s.n. (MP); Schott s.m. (V); Mikan s.n. (V); Gardner 5547 (B, Camb., , V); MINAS GERAES: Ilheos, date lacking, Martius 966 (B, M, MBG, NY, V); Organ Mts., Jan., 1838, Miers 4027 (US); SAO PAULO: in campis editoribus prope S. Paulo civitatem, Dec.-Jan., year laeking, Martius 324 (M); DATA INCOMPLETE: Riedel s.n. (B, MP, NY); McRae s.n. (Camb.). That Miers was correct in interpreting Echites arborea Vell. as a member of the genus Malouetia must be agreed after an examination of the plate and text by any one familiar with the characters of the group. Vellozo's plate ( pl. 47) not only indi- eates the foliar foveae, so pronounced in M. lanceolata and M. Martii, the type specimens of which appear clearly conspe- cific, but those extremely characteristic structures of the leaves are specifically deseribed as **poris in divisione nervorum ad costam,’’ which could possibly apply only to species of Malou- etia (or the liana Forsteronia) in the American flora. The general habit, relative size of the flowers, exserted anthers, and distribution (Mendanha, D. F.) completely clarify the natural affinities of this species. Miers also called attention to the erroneous inclusion of a comose seed in Vellozo's plate, pos- sibly explaining Mueller's relegation of the species to Secondatia. 8. Malouetia Duckei Mgf. Notizblatt 9: 962. 1926. Trees (20 m, tall, fide Ducke); stems relatively stout, gla- brous, reddish-brown, inconspicuously lenticellate when fully mature; leaves opposite, petiolate, oblong-elliptic, apex shortly (284) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 251 and obtusely acuminate, base obtuse to broadly acute, 6-13 cm. long, 1.5-5.5 em. broad, subcoriaceous, glabrous throughout, nitidulous above, beneath opaque, rarely and very inconspicu- ously foveate in the axils of the midrib; petioles 0.3-0.6 cm. long, glabrous; umbels terminal, bearing relatively numerous greenish-white flowers; peduncle 0.2-0.4 cm. long; pedicels 0.3-0.5 cm. long, minutely papillate to puberulent-papillate ; calyx-lobes ovate, acute, 0.15-0.2 em. long, closely imbricated at anthesis, papillate within and without; corolla salverform, only slightly broadening toward the base, 0.75-0.9 em. long, about 0.1 cm. in diameter at the base, faucal tube 0.1 em. long, about 0.15 em. in diameter at the orifice, faucal annulus rela- tively conspicuous, very minutely puberulent or puberulent- papillate without, papillate within, the lobes obliquely oblong- elliptie, acute to acuminate, 0.7—0.8 em. long, 0.25—0.3 em. broad, reflexed, papillate to puberulent-papillate within, particularly toward the base, minutely papillate without; anthers conspic- uously exserted, ovate-elliptic, 0.25 em. long, very minutely and sparsely pilosulose to puberulent-papillate dorsally; ovary 0.1 em. long, puberulent-papillate ; stigma 0.125-0.15 em. long; nectaries 14-24 equalling the ovary; follicles unknown. COLOMBIA: CAQUETA: Rio Caqueta, super cataractas Cupaty, silva non in- undabilis, Nov. 18, 1912, Ducke 21822 (B, US). BRAZIL: PARA: Itaituba, Rio Tapajoz, silva non inundata, Aug. 26, 1923, Ducke 17464 (B, TYPE, S, US, MBG, photograph and analytical drawings); Faro, silva non inundabilis, Aug. 26, 1907, Ducke 21625 (B, US); AMAZONAS: Manáos, silva non inundabili ultra locum Flores, Aug. 10, 1932, Ducke 23952 (B). The specimen from Amazonian Colombia is placed with this species rather reluctantly because of the smaller foliage leaves, which are much more conspicuously foveate beneath than among the typical specimens from lower downstream. It is hoped that the discovery of fruit may enable a sharper distine- tion between this species and the preceding which it closely resembles. 9. Malouetia pubescens Mgf. Notizblatt9: 88. 1924. Shrubs or small trees (3-8 m. tall, fide Ule) ; stems relatively stout, very minutely puberulent when young, soon becoming glabrate, dark reddish-brown, rather inconspicuously lenticel- (285) [VoL. 22 252 ANNALS OF THE MISSOURI BOTANICAL GARDEN late when fully mature; leaves opposite, shortly petiolate, ob- long-elliptie, apex obtusely acuminate, base obtuse, 5-10 em. long, 1.5-4.0 em. broad, firmly membranaceous, above some- what nitidulous, beneath finely and densely puberulent; pet- ioles 0.3-0.4 em. long, minutely puberulent; umbels terminal, bearing relatively few small, white flowers; peduncle 0.2- 0.3 em. long; pedicels 0.4-0.5 em. long, minutely puberulent ; calyx-lobes ovate, acute, 0.15-0.17 em. long, minutely puberu- lent; corolla salverform, only somewhat swollen toward the base, the tube 0.4—0.45 em. long, about 0.125 em. in diameter at the base, faueal tube 0.075 em. long, about 0.1 em. in diameter at the orifice, glabrous without, or essentially so, within mi- nutely puberulent-papillate above, the lobes ébliqnaly ovate- oblong, obtuse or broadly acute, 0.55-0.6 cm. long, reflexed, papillate without, minutely puberulent within, particularly to- ward the base; anthers conspicuously exserted, ovate-elliptic, 0.175 em. long, minutely pilosulose dorsally; ovary 0.1 em. long, minutely puberulent; stigma 0.05 cm. long; nectaries about equalling the ovary ; follicles unknown. BRAZIL: AMAZONAS: in Campwald bei S. Marcos, Rio Branco, Dec., 1908, Ule 7827 (B, TYPE, US, MBG, photograph and analytical drawings). A peculiar and characteristic species whose association with the cestroides-arborea complex must be confirmed by fruiting specimens. 10. Malouetia Tamaquarina (Aubl.) A. DC. in DC. Prodr. 8: 378. 1844; Muell.-Arg. in Mart. Fl. Bras. 6*: 92. pl. 26. fig. 2. 1860; Miers, Apoc. So. Am. 87. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Cameraria Tamaquarina Aubl. Hist, Pl. Gui. Fr. 1: 260; 3: pl. 102. 1775. Cameraria Guyanensis Aubl. loc. cit. 262. 1775. Cameraria lutea Lam. Encycl, 1: 573. 1783. Tabernaemontana odorata Vahl, Eclog. 2: 22. 1798. Malouetia Tamaquarina A. DC. p. minor A. DC. loc. cit. 319. 1844. Malouetia odorata (Vahl) A. DC. loc, cit. 1844; Miers, (286) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 253 Apoc. So. Am. 87. 1878 (where erroneously attributed to himself). Malouetia obtusiloba A. DC. loc. cit, 1844; Miers, loc. cit. 88. 1878 (where erroneously cited as obtusifolia). Malouetia albiflora Miq. Stirp. Surinam. Sel. 161. 1851. Malouetia Tamaquarina A. DC. p. brasiliensis Muell.-Arg. loc, cit. 1860, not A. DC., in part as to specimens cited. Malouetia Tamaquarina A. DC. y. lancifolia Muell.-Arg. loc. cit. 1860. Malouetia Guianensis ( Aubl.) Miers, loc. cit. 87. 1878. Malouetia Tarumensis Benth. ex Miers, loc. cit. 1878, sphalm. Shrubs and small trees (3-10 m. tall, fide de la Cruz and Mon- teiro da Costa); stems relatively stout, glabrous, inconspicu- ously lenticellate when fully mature; leaves opposite, shortly petiolate, narrowly oblong- to ovate-elliptie, apex subcaudate- acuminate, base obtuse, 4—15 cm. long, 1.3-7.0 em. broad, mem- branaceous, glabrous throughout, opaque, inconspicuously foveate in the axils of the midrib beneath; petioles 0.2-0.4 em. long, glabrous; umbels terminal, bearing relatively few yellow- ish- or greenish-white flowers (or purplish according to cer- tain collectors); peduncle 0.2-0.4 cm. long; pedicels 1-2 cm. long, glabrous; calyx-lobes ovate, obtuse to broadly acute, 0.2- 0.3 em. long, coriaceous, closely imbricated at anthesis, gla- brous to minutely papillate without, minutely papillate within; corolla salverform, occasionally somewhat inflated at the base, relatively fleshy in texture, the tube 0.9-1.5 em. long, about 0.1- 0.15 cm. in diameter at the base, occasionally dilating to as much as 0.3 em. below the middle, faucal tube 0.1—0.15 em. long, about 0.15-0.25 cm. in diameter at the orifice, faucal annulus conspicuous, 5-lobed, glabrous without, minutely puberulent toward the insertion of the stamens within, the lobes obliquely oblong- to obovate-elliptie, obtusish to shortly acuminate, 0.9- 1.7 em. long, reflexed, papillate to minutely puberulent-papil- late without, minutely puberulent or pilosulose, particularly toward the base within; anthers conspicuously exserted, ovate-elliptie, 0.25-0.4 cm. long, minutely puberulent dorsally ; ovary 0.15-0.2 cm. long, minutely puberulent to puberulent- (287) [ VoL. 22 254 ANNALS OF THE MISSOURI BOTANICAL GARDEN papillate; stigma 0.1-0.15 em. long; nectaries 15-24 equalling the ovary ; follicles terete, 10-35 em. long, 0.4—0.5 em. in diam- eter, faleate, glabrous; seeds 3.5—4.5 em. long, 0.35-0.4 cm. in diameter, glabrous. BRITISH GUIANA: Pomeroon River, Pomeroon District, Dee. 17-24, 1922, Cruz $135 (FM, G, US) ; Kamakusa, upper Mazaruni River, July 11-22, 1923, Cruz 4083 (FM); between the Demerara and Berbice Rivers, July 15-19, 1922, Cruz 1660 (MBG); Amakura i die Distriet, March 23-30, 1923, Cruz 3473 (FM, US); Aruka R., Barima R., Northwest District, June, 1908, Anderson 60 (FM); on edes Moraballi Creek, near judi alt. near sea-level, Sept. 18, 1929, Sandwith à (N > rip. fl. Pomeroon, Aug., 1843, Schomburgk 1378 (B); data incomplete, ee 39 (V); Schomburgk 788 (Camb., MP); Schomburgk 800 (V) een 951 (B, V). DUTCH GUIANA: fluv. Coppename sup., Aug., 1901, Boon 1044 (B, U); Kaboerie, fl. Corantyne, June 21, 1916, Gonggrijp 2135 (B, US); ad fl. Marowyne med., Aug., 1847, Kappler 1825 (MP, S, V); ad fl. Marowyne super. in rupestribus, Sept., year oo pé a (B); data incomplete, Kappler 444a (B, M, MP, S, V); Kappl 5 (MP, S); Hostmann 269 (Camb., D, FM, M, NY, §, V). cb do Karouany, 1855, Sagot 392 (B, MP, V); Sagot 1143 (B, MP, V); environs de Godebert, date lacking, Wackenheim 119 (FM, MP, NY, US); rer 1819, Martin s.n. bs. MP); data incomplete, 1820, Perrottet s.n. (MP); —64, Melinon s.n. (B, MP) pue PARA: Ilha "e Oncas, prope Belem do Para, ripis inundatis, Oct. 2, 1903, Huber 21770 (B); Lago de Alter do Cháo, prope Santarem, ripis arenosis, June 22, 1910, Ducke rr (B); in ripis, Rio Oyapoe, June 4, 1904, Ducke 21775 (B); Bóa Vista, on the eum EN May-June, 1929, Dahlgren 4 pes 96 (FM); low and high land, Aramanahy, Jan. 11, 1932, Costa 253 (FM, MBG); CEARA: data ineomplete, 1842, dd 951 (V); AMAZONAS: Barcellos, Rio T ad rivulum silvestrem, June 13, 1905, Ducke 21763 p Rio Purus, loco Bom Logar, May 14, 1904, Huber 21762 (B); in fruticetis arenosis siecis ad Ega, Sept., 1831, Poeppig 2547 (V); prope E ad Rio Vaupes, Oct. 1852-Jan. 1853, Spruce 2435 (B, Camb., G, MP, NY, V). Monteiro da Costa reports that the wood of this species is used for fashioning small utensils, such as spoons, ete. It is said to be light and soft. Aublet reported his specific adjective derived from the vernacular name used by the Indians of French Guiana. Other names used by the native population of the species’ area of distribution are said to be: **Oonsé balli’’; '*Jaramiloerang" (Dutch Guiana); ‘‘Molongo de Colher” (Para). 11. Malouetia furfuracea Spruce, ex Muell.-Arg. in Mart. Fl. Bras. 6*: 93. 1860; Miers, Apoc. So. Am. 91. 1878. (288) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 255 Malouetia furfuracea Spruce B. grandifolia Muell.-Arg. loc, cit. 1860. Shrubs (fide Ducke and Ule); stems relatively stout, gla- brous, inconspieuously lenticellate when fully mature; leaves opposite, shortly petiolate, narrowly elliptie to broadly oblong- elliptie, apex obtusely subeaudate-acuminate, base obtuse to acute, 5-17 cm. long, 1.5—6.5 em. broad, membranaceous, gla- brous throughout, usually somewhat subnitidulous above, be- neath opaque, foveate in the axils of the midrib; petioles 0.3— 0.5 cm. long; umbels terminal, bearing relatively few white flowers; peduncle 0.2-0.4 cm. long; pedicels 0.8-2.0 cm. long, glabrous; calyx-lobes ovate, acute to acuminate, 0.3-0.5 cm. long, rather fleshy and coriaceous in texture, closely imbricated at anthesis, glabrous or minutely papillate without, papillate at the tips within, margins minutely ciliolate; corolla salver- form, frequently somewhat inflated at the base, rather fleshy in texture, the tube 1.0-1.4 em. long, about 0.1-0.125 em. in diam- eter at the base, faucal tube 0.1-0.125 em. long, about 0.15 cm. in diameter at the orifice, faucal annulus conspicuous, 5-lobed, glabrous without, minutely puberulent above within, the lobes obliquely ovate- or obovate-elliptie, obtuse to shortly acumi- nate, 1.3-2.3 em. long, 0.3-0.45 em. long, reflexed, papillate to puberulent-papillate without, minutely puberulent toward the base within; anthers conspieuously exserted, ovate-elliptie, 0.3-0.4 cm. long, minutely puberulent dorsally; ovary 0.1- 0.125 em. long, puberulent-papillate; stigma 0.1-0.15 em. long; nectaries 15-24 equalling the ovary; follicles terete, 15-30 em. long, 0.3-0.4 cm. in diameter, faleate, glabrous; seeds 0.35- 0.4 em. long, about 0.25-0.3 em. in diameter, rather sparsely pilose or villous. DUTCH GUIANA: fluv. Suriname sup., July 1, 1921, Boschwesen 5302 (B, U); ad fl. Lawa, Sept., year lacking, Kappler 2089 (B). FRENCH GUIANA: data incomplete, Richard s.n. (MP). BRAZIL: PARA: Lago de Faro, ad ripas, Aug. 20, 1907, Ducke 21774 (B); Rio Trombetas infer., ad ripas arenosas loco Caipuru, Sept. 16, 1910, Ducke 21773 (B); RIO DE JANEIRO: data incomplete, Glaziou 14082 (B); AMAZONAS: Cachoeira Grande, bei Manäos, May, 1910, Ule 8952 (B); am Strande, Rio Negro, Manäos, Jan., 1901, Ule 5339 (B); in vicinibus Barra, April, 1851, Spruce 1003 (B, TYPE, M, NY, V, MBG, photograph and analytical drawings); ad oram meridionalem Rio (289) [Vor. 22 256 ANNALS OF THE MISSOURI BOTANICAL GARDEN Negro, usque ad concursum flum. Solimoes, May, 1851, Spruce 1566 (B, Camb., G, M, NY, V); prope S. Gabriel do Caehoeira, ad Rio Negro, 1852, Sores 2305 (B, Cami, G, MP, NY, V). The uncertainty which I have encountered in identifying specimens relegated to this species is decidedly conducive to the view that they may constitute merely a variety, or perhaps only a phase, of M. Tamaquarina. On the other hand, it cannot be denied that the seeds of the few available fruiting specimens are more or less villous, and that the follieles are somewhat more slender than those of M. Tamaquarina. The calyx-lobes alone are certainly insufficient evidence upon which to base the distinetion of the species, as Markgraf has recently pointed out (in Pulle, Fl. Surinam 4: 57. 1932). Nevertheless the latter author has maintained the specifieity of M. furfuracea. The question is clearly one which will await solution by a student familiar with the plants in the field. 12. Malouetia glandulifera Miers, Apoc. So. Am. 90. pl. 134. 1878. Malouetia Tamaquarina A. DC. B brasiliensis Muell.-Arg. in Mart. Fl. Bras. 6': 92. 1860, not A. DC. as to speci- mens cited, in part, Stems relatively stout, glabrous, inconspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong- oblaneeolate, apex obtuse or rounded, base broadly acute to obtuse, 5-9 em. long, 1.5-3.0 em. broad, subcoriaceous, glabrous throughout, foveate in the axils of the midrib beneath ; petioles 0.2-0.4 cm. long; umbels terminal, bearing 5-8 flowers; pe- duncle 0.2-0.3 em. long; pedicels 1.3-2.0 em. long, acerescent in fruit, glabrous; calyx-lobes ovate, broadly acute to obtuse, 0.15—0.175 cm. long, relatively fleshy in texture, closely im- bricated at anthesis, generally puberulent-papillate within and without; corolla salverform, the tube 0.9 em. long, about 0.1 cm. in diameter at the base, faucal tube 0.1 em. long, about 0.15 em. in diameter at the orifice, conspieuously annulate (the lobes in desiccation triangular, not divided), glabrous without, very minutely puberulent-papillate within, the lobes obliquely obo- vate-elliptie, acute, 1.5 em. long, 0.4-0.5 cm. broad, reflexed, about uniformly papillate to glabrate within and without; (290) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 257 anthers rather inconspicuously exserted (in desiccation), ovate-elliptic, 0.25 cm. long, essentially glabrate dorsally; ovary 0.1 em. long, minutely puberulent; stigma 0.075 em. long ; nectaries about 25 equalling the ovary; follicles terete, 8-10 cm. long, 0.35-0.4 em. in diameter, divaricate, glabrous; seeds 2.0— 2.5 em. long, glabrous. VENEZUELA: AMAZONAS: ad flumina Casiquiari, Vasiva et Pacimoni, 1853-4, Spruce 3305 (B, Camb., G, NY, V, isorvyPES, MBG, photograph and analytical drawings). Although the status of this species is somewhat ambiguous in view of its poor representation in herbaria and the manifest variability of the closely neighboring M. Tamaquaria and M. furfuracea, it appears necessary to recognize it specifically because of its unusual foliage, almost uniformly papillate co- rolla-lobes, relatively deep insertion of anthers, and construc- tion of the faueal annulus. The nature of the faucal annulus itself is one which appears to have been widely misinterpreted in past literature. In in- augurating the genera Malouetia and Robbia, A. de Candolle made no reference to the annulus with regard to the former, on the other hand stating quite definitely (in DC. Prodr. 8: 378. 1844) ‘‘fauce exappendiculata.’’ With regard to the latter, the lobes were described, in part, as ‘‘basi intus in media parte calloso dentatis, appendicibus praetera nullis’’ [loc. cit. 445. 1844]. In 1860, however, Mueller consolidated Malouetia and Robbia and emended the description of the aggregate genus, including the phrase ‘‘Corolla . . . intus fauce 5-appendicu- latus [tubus], appendieulae parvae subsquamulaeformes, lobis corollae oppositae [in Mart. Fl. Bras. 6t: 90. 1860]." Still later, Miers, usually an acute and intuitive observer, described the corolla-throat of Malouetia as **saepius squamulis 5 prae- ditus" [Apoc. So. Am. 86. 1878]; and later, with regard to Robbia, censures Delessert [Icon. 5: pl. 52. 1846] for omitting to depict ‘‘the 5 scales which close the mouth of the corolla" of R. cestroides (Nees) A. DC. More recently, Schumann has spoken, with regard to Malouetia of the **Blkr. . . . in der Mitte verengter, am Schlunde 5-10 schuppiger Róhre" [in Engl. € Prantl, Nat. Pflanzenfam. 4?: 186. 1895]; and simi- (291) (Vor. 22 258 ANNALS OF THE MISSOURI BOTANICAL GARDEN larly of **im Sehlunde beschuppter Róhre”” [loc. cit. 187. 1895] of the corolla of Robbia. Markgraf has maintained the tradi- tional deseription of an appendiculate corolla in Malouetia by describing the throat as ‘‘closed by 5-10 short, triangular scales”” [in Pulle, Fl. Surinam 4: 57. 1932]. One might expect the “scales”? of the corolla-throat of Malouetia to provide an easy criterion of specific entities, as those of Prestonia, for example, and such might be thought to be the case after only a few dissections. After many dissec- tions, however, this hope fades. Not only is a striking varia- bility found among individuals from specimens of the same collector’s number, but a more significant phenomenon is found in recently collected specimens which upon careful soaking re- fuse to divulge the ‘‘squamulae”’ or “scales?” at all! Almost exactly identical specimens, made by older collectors, how- ever, usually demonstrate very pronounced ‘‘faucal scales. ”” The genus Laubertia, of the American Echitoideae, is typi- fied in part by a corolline structure which appears in well- pressed specimens as a slightly thickened, but still membrana- ceous, ring or ‘‘annulus’’ at the orifice of the throat. Prestonia and Rhodocalyx are similarly provided. Quite recently Mr. William A. Schipp, of Stann Creek, British Honduras, has sent me a vial of aleoholie preservative containing several flowers of an apoeynaceous genus which was at first examination a per- plexing novelty. After careful serutiny, however, it was recog- nized as a new species of Laubertia. The first failure of recog- nition was caused by the fact that the corolline orifice, which in well-desiccated material appears to have a well-defined mem- branaceous annulus, is in fresh material represented by a uni- form, callose thickening of the orifice grading indefinitely into the more delicate texture of the corolla-tube and lobes. In other words, the faucal ‘‘ring,’’ so manifest in exsiccatae, is not an artifact in itself, but its true nature 1s greatly altered in desiecation, caused by the drying and consequent collapse of the fleshy tissues of the orifice. Subsequent pressure completes the illusion of the membranaceous ““annulus. ”” After examining a large series of specimens of Malouetia previously described as provided with five to ten “scales,” a (292) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 259 similar interpretation to that arrived at with regard to Lau- bertia appears plausible. It is well to acknowledge, however, that if such be the case, wide knowledge of the living specimens, or at any rate liquid preserves, is necessary for accurate in- formation. From an examination of numerous specimens of recent collection, in which the tissues are relatively resilient upon soaking and heating, it appears that the corolline orifice of Malouetia species, as in Laubertia, is rather uniformly thiekened. The high insertion of the anthers, however, gives to the orifice a five-lobed contour. The anthers, which alternate with the corolla-lobes, also protrude beyond the faucal annulus in most species. "These factors, together, would appear to be responsible for the configuration of the ‘‘squamulae’’ or ‘‘fau- cal scales?” in material which has been long desiccated; whose tissues, as every systematic botanist can testify, have conse- quently lost their earlier powers of resilience. In Malouetia glandulifera the **faucal scales" of the her- barium material available (collected in 1853-54) are definitely acute-triangular, entire, and quite unlike those of M. Tama- quarina specimens of comparable age, which are broader, lower, rounded, and usually more or less bifid. It is not meant to imply that faucal differences do not exist between these two species: such is probably the case. Nevertheless it will prob- ably be agreed that difficulty would be encountered in an at- tempt either to reconstruct the living condition from the desic- cated, or to recognize the living from a description of the desiccated. 13. Malouetia peruviana Woodson, spec. nov. Arbuscula (3-7 m. alta, fide Klug); ramulis teretibus cras- siusculis glaberrimis maturitate inconspicue lenticellatis; foliis oppositis breviter petiolatis anguste oblongo- vel ovato- ellipticis aut acute aut subeaudate acuminatis 5-20 em, longis 1.5-8.5 em. latis subcoriaceis glaberrimis supra subnitidulis subtus opacis in axillis nervi medii conspicuissime foveatis ; petiolis 0.3-0.5 em. longis; umbellis terminalibus flores 3-8 viridi-albidas gerentibus; pedunculo 0.2-0.5 em. longo; pedi- eellis 1.0-1.5 em. longis glaberrimis; calycis laciniis ovatis (293) [ Vor. 22 260 ANNALS OF THE MISSOURI BOTANICAL GARDEN obtusis vel late acutis 0.25—0.4 em. longis subeoriaceis valde im- brieatis extus intusque minutissime papillatis glabratisve; corollae salverformis cereolae tubo 1.2-1.5 em. longo basi ca. 0.15-0.3 em, diametro metiente supra saepius paululo dilatato faucibus tubulari-campanulatis ea. 0.1 em. longis ostio ca. 0.25-0.3 em. diametro metiente ibique conspicue calloso- annulato extus glabro intus prope insertionem staminum minute puberulo-papillato lobis oblique ovato- vel obovato- ellipticis acutis 1.2-1.7 em. longis 0.5-0.7 em. latis patentibus intus extusque minute papillatis glabratisve; antheris con- spleue exsertis ovato-elliptieis 0.4-0.5 cm. longis, glabris glabratisve; ovario 0.15 em. longo minute puberulo; stigmate 0.15 em. longo; nectariis ovarium 15—24 aequantibus ; folliculis ignotis. Small trees (3-7 m. tall, fide Klug) ; stems terete, relatively stout, glabrous, inconspicuously lenticellate when fully mature; birds opposite, shortly petiolate, lance- to ovate-elliptie, apex acutely and frequently subeaudate-acuminate, 5-20 em, long, 1.5-8.5 em. broad, subcoriaceous, glabrous throughout, above somewhat subnitidulous, beneath opaque, very conspicuously foveate in the axils of the midrib; petioles 0.3-0.5 em. long; umbels terminal, bearing 3-8 greenish-white flowers ; peduncle 0.2-0.5 em. long; pedieels 1.0-1.5 em. long, glabrous; calyx- lobes ovate, obtuse to broadly acute, 0.25-0.4 cm. long, sub- coriaceous, closely imbricated at anthesis, minutely papillate to glabrate throughout; corolla salverform, rather thick and waxy in texture, the tube 1.2-1.5 em. long, about 0.15-0.3 em. in diameter at the base, very slightly dilated above if at all, faucal tube campanulate, 0.1 cm. long, about 0.25—0.3 em. in diameter at the orifice, conspicuously callose-annulate, glabrous without, minutely puberulent-papillate within toward the insertion of the stamens, the lobes obliquely ovate- or obovate-elliptie, acute, 1.2-1.7 em. long, 0.5-0.7 em. broad, reflexed, about uni- formly papillate to glabrate within and without; anthers con- spieuously exserted, ovate-elliptie, 0.4—0.5 em. long, glabrous to glabrate; ovary 0.15 em. long, minutely puberulent; stigma 0.15 em. long; nectaries about !5—24 equalling the ovary; fol- licles unknown. (294) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 261 PERU: LORETO: woods, Iquitos, alt. about 100 m., Aug. 3-11, 1929, Killip ¢ Smith 27004 (FM, MBG, TYPE, NY, US); Aug. 2-8, 1929, Killip $ Smith 27371 (MBG, NY, US); Mishuyacu, near Iquitos, forest, alt. 100 m., Febr.-Mareh, 1930, Klug, 972 (FM, NY, US); Caballo-Cocha, on the Amazon River, Aug., 1929, Wil- liams 2089 (FM, MBG); Timbuchi, on the Rio Nanay, June-July, 1929, Williams, 928 (FM, MBG); Williams 973 (FM, MBG). There can be little doubt that this species is distinct from M. Tamaquarina and M. furfuracea for which it has previously been mistaken. Its salient characteristic, as indicated in the key to species, are supplemented by many minor ones which may be detected through a comparison of the foregoing descriptions. 14, Malouetia amplexicaulis Muell.-Arg. in Mart. Fl. Bras. 61: 91. 1860; Miers, Apoc. So. Am. 91. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Stems relatively stout, glabrous, rather inconspicuously lenticellate when fully mature; leaves opposite, sessile or sub- sessile, appearing more or less amplexicaul, broadly oblong- elliptic, apex obtusely, and almost subcaudately, acuminate, base obtuse to rounded, or slightly cordate, 9-23 em. long, 3.5- 10 em. broad, firmly membranaceous, glabrous throughout, somewhat subnitidulous above, beneath opaque and rather in- conspicuously foveate in the axils of the midrib; umbels termi- nal, relatively few-flowered; peduncle 0.15-0.4 cm. long; pedi- cels 1.0-1.5 em. long, glabrous; calyx-lobes ovate, obtuse, 0.2— 0.3 em. long, somewhat subcoriaceous, closely imbricated at anthesis, minutely papillate within and without; corolla salver- form, the tube 1.2-1.3 em. long, about 0.125 em. in diameter at the base, faucal annulus about 0.075 em. long, about 0.15 cm. in diameter at the orifice, conspicuously callose-annulate, gla- brous without, minutely puberulent within toward the inser- tion of the stamens, the lobes obliquely and broadly oblong-el- liptie, obtuse to acute, 0.8-0.9 em. long, reflexed, very minutely and indefinitely papillate without, minutely puberulent or pu- berulent-papillate within; anthers conspicuously exserted, ob- long-elliptie, 0.3-0.35 em. long, minutely puberulent dorsally; ovary 0.1 em. long, densely puberulent-papillate; stigma 0.075 em. long; nectaries about Ya equalling the ovary; follicles unknown. (295) [ Vor. 22 262 ANNALS OF THE MISSOURI BOTANICAL GARDEN BRAZIL: AMAZONAS: prope Panure ad Rio Vaupes, Oct. 1852-Jan, 1853, Spruce 2595 (Camb., G, MP, NY, V, isorvPEs, MBG, photograph and analytical drawings). The chief, if not the only, distinetion of M. amplexicaulis ap- pears to be the rather large, nearly sessile and amplexicaul foliage. However, but a single collection is known to us, and this, together with the fact that all specimens are rather meagre, renders the status of the species somewhat dubious. It is to be hoped that the renewed botanical activity in the upper Amazon Valley may lead to the rediseovery of this in- teresting plant. 15. Malouetia Killipii Woodson, Ann. Mo. Bot. Gard. 18: 901. 1931. Small trees (5-10 m. tall, fide Klug; Killip & Smith) ; stems relatively stout, terete, minutely puberulent when very young, soon becoming glabrate, rather inconspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, nar- rowly oblong- to broadly ovate-elliptie, apex obtusely and narrowly subeaudate-acuminate, base obtuse to rounded, 15- 25 em. long, 5-12 em. broad, membranaceous, above somewhat subnitidulous, glabrous, beneath densely and minutely veluti- nous-puberulent; petioles 0.3-0.7 cm. long; umbels terminal, infrequently lateral as well, bearing several greenish-white flowers; peduncle 0.3-0.5 em. long; pedicels 0.5-1.0 em. long, very minutely papillate to essentially glabrous; calyx-lobes ovate, obtuse to broadly acute, 0.15—0.25 cm. long, somewhat subcoriaceous, closely imbrieated at anthesis, minutely papil- late within and without; corolla salverform, somewhat thick and waxy in texture, the tube 1.1-1.2 em. long, about 0.1 em. in diameter at the base, faucal tube 0.1 em. long, about 0.15-0.2 em. in diameter at the orifice, conspicuously callose-annulate, gla- brous to minutely papillate without, minutely puberulent with- in toward the insertion of the stamens, the lobes obliquely ovate, acute to obtuse, 0.9-1.2 em. long, 0.35-0.5 em. broad, re- flexed, papillate without, minutely puberulent within toward the base; anthers conspicuously exserted, ovate-elliptie, 0.25— 0.3 em. long, minutely puberulent dorsally; ovary 0.1 em. long, minutely puberulent; stigma 0.15 em. long; nectaries 15-34 equalling the ovary ; follieles unknown. (296) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE. IV 263 PERU: LORETO: woods, Iquitos, alt. about 100 m., Sept. 26, 1929, Killip $ Smith 29860 (FM, MBG, TYPE, NY, US); forest, Mishuyaeu, near Iquitos, alt. 100 m., Oct.—Nov., 1929, Klug 79 (B, FM, US). Sect. 2. Gracies Woodson. Corolla-tube cylindrical, abruptly constricted at the insertion of the stamens; anthers narrowly oblong- to linear-elliptic, wholly included, inserted about midway or deeper within the corolla-tube. Spp. 16-20. KEY TO THE SPECIES a. Corolla-tube 0.7-1.5 em. long; species of eastern tropical South America. b. Corolla-lobes ovate-oblong; ovary glabrous; plants of British Guiana.... bb. Corolla-lobes linear to oblong-lanceolate; ovary pubescent; species of northern Brazil. e. Corolla-lobes about as long as the tube or somewhat ag d stamens inserted at somewhat below midway within the corolla- d. Corolla-lobes oblong- to ovate-lanceolate; eorolla-tube not narrowing toward the orifice; anthers glabrous; leaves p or sub- AP CM S REL C SCHOEN ER . M. virescens dd. Coretta: ‘ches linear-lanceolate; corolla-tube extremely cae nar- rowing toward the orifice; anthers minutely mie e ; leaves delicately membranaceous........oooooooomoooo.o.o.. . M. gracillima ec. Corolla-lobes about twice as long as the tube, or nearly so; is) in- rted about midway within the eorolla-tube.............. 19. nitida aa. RUE 0.4—0.55 em. long; plants of Central Ameriea..20. M. guatemalensis 16. Malouetia gracilis (Benth.) A. DC. in DC. Prodr. 8: 380. 1844; Miers, Apoc. So. Am. 87. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 187. 1895. Tabernaemontana gracilis Benth. in Hook. Jour. Bot. 3: 244. 1841 Stems relatively slender, terete, glabrous, rather inconspicu- ously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptie, apex acuminate, base obtuse, 3.5- 8.0 em. long, 1-3 em. broad, membranaceous, glabrous through- out, above subnitidulous, beneath opaque, pallid; petioles 0.3- 0.5 em. long, glabrous; umbels terminal, few-flowered; pe- dunele 0.1-0.3 em. long; pedicels 0.3-0.5 em. long, glabrous; calyx-lobes ovate, acute, closely imbricated, 0.225—0.25 em. long, glabrous without, minutely puberulent-papillate within toward the tips; corolla salverform, the tube cylindrical, abruptly con- stricted at the insertion of the stamens, 1.4—1.5 em. long, about (297) [Vor. 22 264 ANNALS OF THE MISSOURI BOTANICAL GARDEN 0.125 em. in diameter at the base, about 0.15 em. in diameter at the orifice, glabrous without, minutely puberulent within above the insertion of the stamens, somewhat callose-thickened gener- ally above the insertion of the stamens (appearing as 5-10 linear welts in desiccation), the lobes ovate-oblong, acute, 0.9— 1.0 em. long, densely puberulent-papillate without and within, widely spreading; anthers included, inserted at somewhat be- low midway within the corolla-tube, narrowly oblong- to linear- elliptie, 0.35 em. long, glabrous; ovary 0.1 em. long, glabrous; stigma 0.075 cm. long; nectaries about 1⁄2 equalling the ovary; follieles terete, relatively short and stout, 5.0—5.5 em. long, rigidly divaricate, glabrous ; seeds 0.7-0.9 em. long, glabrous. BRITISH GUIANA: upper Essequibo River, Jan., 1842, Schomburgk 39 (B, Camb., G, NY, isotypes, MBG, photograph and analytical drawings). 17. Malouetia virescens Spruce, ex Muell.-Arg. in Mart. Fl. Bras. 6': 92. 1860; Miers, Apoc. So. Am. 90. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4*: 187. 1895. Small trees (according to Ducke); stems relatively stout, terete, glabrous, rather inconspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptic, apex acute to obtuse, base obtuse to broadly acute, 4.5-10.0 em. long, 1.5-3.0 em. broad, coriaceous or subcoriaceous, glabrous throughout, nitidulous above, opaque and inconspicuously foveate in the axils of the midrib beneath; petioles 0.3—0.4 cm. long; umbels terminal, few-flowered ; peduncle 0.1-0.2 em. long; pedicels 0.5-0.8 cm. long, glabrous; calyx-lobes ovate, broadly obtuse, 0.125-0.2 em. long, somewhat coriaceous, closely im- brieated at anthesis, glabrous without, very minutely and in- definitely papillate within; corolla salverform, the tube cylin- drical, abruptly constricted at the insertion of the stamens, 0.8-1.0 em. long, about 0.1 em. in diameter at the base, about 0.125-0.15 em. in diameter at the orifice, somewhat callose- thickened generally in the region of the orifice (appearing as 5-10 linear welts in desiccation), glabrous without, minutely and indefinitely papillate above the insertion of the stamens within, the lobes oblong- to ovate-lanceolate, acute, 0.7—1.1 em. long, 0.25-0.3 em. broad, widely spreading, minutely papillate (298) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 265 to essentially glabrous without, puberulent-papillate toward the base within; anthers included, inserted somewhat below midway within the corolla-tube, narrowly oblong- to linear- elliptic, 0.325-0.375 cm. long, glabrous or essentially so; stigma 0.15 em. long; ovary 0.1 em. long, very minutely puberulent; nectaries about Ya equalling the ovary; follicles unknown. BRAZIL: AMAZONAS: prope Panure ad Rio Vaupes, Oet. 1852-Jan. 1853, Spruce 2472 (B, TYPE, Camb., G, NY, MBG, photograph and analytical drawings); Rio Curicuriary, afl. R. Negro super., in ripis profunde inundatis, Dec. 23, 1931, Ducke 23956 (B) 18. Malouetia gracillima Woodson, spec. nov. Ramulis teretibus gracilibus glaberrimis; foliis oppositis petiolatis ellipticis apice acute obtuseve acuminatis basi ob- tusis 5-10 cm. longis 2.8-4.0 em. latis tenuiter membranaceis omnino glaberrimis supra nitidis et eleganter venosis subtus opacis in axillis nervi medii obseure foveatis; petiolis 0.3- 0.6 em. longis; umbellis et terminalibus et lateralibus flores paucas parvas gracillimas gerentibus; pedunculo 0.1—0.15 em. longo; pedicellis 0.4-0.6 em. longis glabris; calycis laciniis ovatis vel ovato-lanceolatis acutis 0.09-0.15 cm. longis extus intusque minute puberulo-papillatis margine ciliolatis; corol- lae graciliter salverformis tubo 1.0-1.1 em. longo basi ca. 0.1 em. diametro metiente faucibus ca. 0.195 em. diametro metientibus vix incrassatis extus glabro intus prope fauces minute puberulo-papillato lobis oblique lineari-lanceolatis acuminatis 0.8-1.0 em. longis 0.08-0.1 em. latis extus minute papillatis intus basi puberulo-papillatis patulis; antheris in- clusis paulo sub medio corollae tubi insertis lineari- ellipticis 0.3 em. longis sparse minutissimeque pilosulis; ovario 0.05 em. longo minutissime puberulo; stigmate 0.1 em. longo ; nectariis ovario multo brevioribus; follieulis ignotis. Stems terete, relativas slender, glabrous; leaves opposite, petiolate, elliptic, apex acutely or obtusely acuminate, base ob- tuse, 5-10 em. long, 2.8—4.0 cm. broad, delicately membrana- ceous, glabrous throughout, nitid and elegantly veined above, beneath opaque, obscurely foveate in the axils of the midrib; petioles 0.3-0.6 em. long; umbels both terminal and lateral] (299) (Vor. 22 266 ANNALS OF THE MISSOURI BOTANICAL GARDEN bearing relatively few, small, graceful flowers; peduncle 0.1- 0.15 em. long; pedicels 0.4—0.6 em. long, glabrous; calyx-lobes ovate or ovate-lanceolate, acute, 0.09-0.15 em. long, minutely puberulent-papillate without and within, the margin ciliolate; corolla gracefully salverform, the tube 1.0-1.1 em. long, about 0.1 em. in diameter at the base, about 0.125 em. in diameter at the orifice, abruptly constricted at the insertion of the stamens, glabrous without, minutely puberulent-papillate toward the throat, the lobes obliquely linear-lanceolate, acuminate, 0.8— 1.0 em. long, about 0.08-0.1 em. broad, minutely papillate with- out, puberulent-papillate toward the base within, spreading; anthers included, inserted somewhat below midway within the corolla-tube, narrowly oblong- or linear-elliptic, 0.3 em. long, sparsely and minutely pilosulose dorsally ; ovary 0.05 em. long, very minutely puberulent ; stigma 0.1 em. long; nectaries much shorter than the ovary (scarcely manifest) ; follicles unknown. BRAZIL: AMAZONAS: Lages, April 8, 1879, Traill 517 (G, TYPE, K, MBG, photo- graph and analytical drawings). This is the most distinetive of the South American species of sect. Graciles, and is unmistakable because of its extremely narrow corolla-tube. Traill reports the plant as the “Ygapo tree.” 19. Malouetia nitida Spruce, ex Muell.-Arg. in Mart. Fl. Bras. 6': 94. 1860; Miers, Apoc. So. Am. 91. 1878. Stems relatively stout, terete, glabrous, greyish-brown, rather inconspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-elliptie, apex acutely acumi- nate, base obtuse to rounded, 8-15 em. long, 3-6 em. broad, membranaceous, glabrous throughout, nitid above, beneath opaque, obseurely and rather infrequently foveate in the axils of the midrib; petioles 0.2-0.5 em. long; umbels terminal, oc- easionally lateral as well, bearing several flowers; peduncle 0.3-0.5 em. long; pedicels 1.2-1.5 em. long, glabrous; calyx- lobes ovate, acute, 0.2-0.25 em. long, minutely puberulent-papil- late at the tips without and within, otherwise essentially gla- brous; corolla salverform, the tube cylindrical, abruptly con- stricted at the insertion of the stamens, 0.7-0.75 em. long, about (300) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 267 0.125 cm. in diameter at the base and orifice, very indefinitely papillate without, minutely papillate to puberulent-papillate within toward the throat, the lobes linear-oblong, acute to acuminate, 1.3-1.4 cm. long, widely spreading to reflexed, gla- brous without, minutely puberulent toward the base within; anthers included, inserted at about midway within the corolla- tube, linear-elliptic, 0.3 em. long, essentially glabrous; ovary 0.1 em. long, minutely puberulent-papillate; stigma 0.12 cm. long; nectaries about Ya equalling the ovary ; follicles unknown. BRAZIL: AMAZONAS: prope Barra, Prov. Rio Negro, Aug., 1851, Spruce 1672 (B, Camb., G, ISOTYPES, MBG, photograph and analytical drawings). 20. Malouetia guatemalensis (Muell.-Arg.) Standl. Jour. Wash. Acad. Sei. 15: 459. 1925. Stemmadenia guatemalensis Muell.-Arg. Linnaea 30: 410. 1860. Malouetia panamensis Heurck & Muell.-Arg. in Heurck, Obs. Bot. 185. 1871. Small trees (5-10 m. tall, fide Pittier; “large trees,” fide Schipp); stems relatively stout, terete, reddish-brown, gla- brous, inconspicuously lenticellate when fully mature; leaves opposite, petiolate, oblong- to ovate-elliptic, obtusely subeau- date-acuminate, base obtuse, 6-25 cm. long, 2-10 cm. broad, firmly membranaceous, glabrous throughout, somewhat nitidu- lous above, beneath opaque, conspicuously foveate in the axils of the midrib; petioles 0.5-1.0 em. long, glabrous; umbels ter- minal, frequently lateral as well, bearing relatively numerous small, greenish-white flowers; peduncle 0.1-0.3 em. long; pedi- cels 0.3-0.5 em. long, glabrous or very minutely papillate; calyx-lobes ovate, acute to obtuse, subcoriaceous, closely im- bricated, 0.15-0.25 cm. long, minutely puberulent-papillate within and without; corolla salverform, the tube abruptly con- stricted at the insertion of the stamens, somewhat inflated above the base, 0.4—0.55 em. long, about 0.125-0.15 em. in diam- eter at the base, orifice about 0.15 em. in diameter, callose- thiekened (appearing as 5 linear, double welts in desiecated material), glabrous without, or minutely papillate toward the orifice, essentially glabrous within, the lobes obliquely lanceo- (301) [Vor. 22 268 ANNALS OF THE MISSOURI BOTANICAL GARDEN late- to ovate-oblong, acuminate, 0.7-1.2 em. long, 0.2-0.3 em. broad, widely spreading, puberulent-papillate within, papillate without; anthers included, inserted slightly below midway within the corolla-tube, narrowly oblong-elliptic, 0.2 cm. long, very minutely puberulent-papillate dorsally; ovary 0.1 cm. long, minutely papillate; stigma 0.075 em. long; nectaries Ya to about equalling the ovary; follicles stout, fusiform, rigidly di- varicate, 10-13 em. long, 1-2 em. in diameter, glabrous; seeds 2.5-3.0 em. long, glabrous. BRITISH HONDURAS: brackish swamps, South p Creek, Sept. 17, 1930, Schipp 653 (B, FM, MBG, NY); data incomplete, Peck 6 GUATEMALA: IZABAL: Mniogalpa, 1841, aM 138 (V, TYPE, MBG, un and analytieal drawings); Livingston, June, 1906, Tuerckheim II 1233 (FM, US); same loeality, Febr. 17, 1905, Deam s.n. (NY, U HONDURAS ATLANTIDA: vicinity of Tela, at sea-level, Dec. {i March 15, 1928, Standley 54737 (FM, US); forest trail west of Tela River, Puerto Sierra, Febr. 15, 1903, Wilson 422 (FM, NY NICAR : data ineomplete, Wright s.n. (B, US). COSTA iin MON: Cienequita, prés Limon, litt. Atlantique, July, 1911, Pit- tier 1613? d PANAMA: BOCAS DEL TORO: Laguna de Chiriqui and its neighborhood, Nov.-Dee., 1885, Hart 128 (US); Changuinola Valley, 1927, Cooper $ Slater 113 (FM, US); CANAL ZONE: wooded swamp, common, between France Field and Catival, Prov. of Colon, Jan. 9, 1924, Standley 30184 (US); near Fort Randolph, swampy woods, common, Dee. 28, 1923, Standley 28636 (US); in low, wet woods near Frijoles Sta., zs R., Febr., 1861, Hayes 55 (B, G); Chagres, Jan.-Mareh, 1850, Fendler 184 MBG, US); bail along the Rio Indio de Gatun, near sea-level, Febr. 17, 1911, Pittier 2813 (FM, NY, US); around Frijoles, in forest and thickets, alt. 25-30 m., July 1,1911, Pittier 3752 (NY, US). This truly distinctive species is probably entitled to sectional rank in the genus Malouetia, because of its large, fusiform fol- licles and peculiar corolla-tube construction. It is difficult to base classification of the genus upon fruit characters at the present time, however, as follicles are unknown for certain of the species, as has already been indicated. Although seemingly distant from the South American species, the relationships of M. guatemalensis are evidently most closely with those of sect. Graciles, as evidenced not only by the deep insertion of the anthers, but the character of the callose thickening of the co- rolla throat as well. (302) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 269 EXCLUDED OR UNCERTAIN SPECIES Malouetia asiatica Sieb. & Zuce. Abh. Akad. München 43: 163. 1846 = Trachelospermum asiaticum (Sieb. & Zucc.) Nakai, in Mori, Pl. Cor. 293. 1922. Malouetia gutanensis Klotzsch, in Schomb. Faun. & Fl. Guian. 952. 1841, nom. nud. This may possibly refer to Cam- eraria Guyanensis Aubl. Hist. Pl. Gui. Fr. 1: 262. 1775, which is synonymous with Malouetia Tamaquarina (Aubl.) A. DC. in DC. Prodr. 8: 378. 1844. Malouetia jasminoides HBK. Nov. Gen. 3: 226. 1819 = Tab- ernaemontana amygdalifolia Jacq. Select. Stirp. Am, Hist. 1: 39. pl. 181. fig. 15. 1763. Malouetia lactiflua Miers, Apoc. So. Am. 88. 1878. Based upon Schomburgk 168. Impressed by the notation of the col- lection with regard to a milky exudation, Miers expressed the unequivocable interpretation of it as a Malouetia. The speci- men is fragmentary and vegetative only, and as the leaves show no indication of the familiar foveae in the axils of the midrib beneath, its relegation to an undetermined species of Zschok- kea by Sandwith (in herb. Kew.) appears far better substanti- ated by our present knowledge than that of Miers. Malouetia puberula Klotzsch, in Schomb. loc. cit. 1841, nom. nud. Malouetia riparia (HBK.) A. DC. loc. cit. 380. 1844 (Tab- ernaemontana riparia HBK. loc. cit. 228. 1819) = Stemmadenia grandiflora (Jacq.) Miers, loc. cit. 75. 1878 (Tabernaemontana grandiflora Jacq. Enum. Pl. Carib. 14. 1760). Malouetia ? sessilis (Vell.) Muell.-Arg. in Mart. Fl. Bras. 6!: 96. 1860 (Echites sessilis Vell. Fl. Flum. 111. 1830; Icon. 3: pl. 35. 1827). The plate cited indicates the plant to be a liana. Since no characters of a Malouetia are indicated save a sessile, umbellate inflorescence and exserted anthers of a small flower, it appears impossible to hazard a suggestion concerning the generic affinities of this species. Malouetia tetrastachya (HBK.) Miers, loc. cit. 92. 1878 = Tabernaemontana tetrastachya HBK. loc. cit. 227. 1819. Robbia macrocarpa (Rich.) Miers, loc. cit. 108. 1878 (Echites (303) [Vor. 22 210 ANNALS OF THE MISSOURI BOTANICAL GARDEN ? macrocarpa Rich. Fl. Cub. 2: 86. 1853, not Wall.) = Catalpa macrocarpa (Rich.) Ekman, in Urb. Symb. Ant. 9: 254. 1924. IX. ODoNTADENIA Benth. Odontadenia Benth. in Hook. Jour. Bot. 3: 242. 1841; A. DC. in DC. Prodr. 8: 359. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 116. 1860; Benth. & Hook. Gen. Pl. 2: 723. 1876; Miers, Apoc. So. Am. 126. 1878; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4°: 169. 1895. Amsolobus A. DC. loc. cit. 395. 1844; Muell.-Arg. loc. cit. 110. 1860; Miers, loc. cit. 168. 1878, Cylicadenia Lem. in Van Houtte, Illustr. Hort. 2: Misc. 9. 1855. Angadenia Miers, loc. cit. 173. 1878, in part. Perictenia Miers, loc. cit. 182. 1878. Mitozus Miers, loc. cit, 217. 1878, in part. Codonechites Mgf. Notizblatt 9: 80. 1924. Echites of early authors, in part, not P. Br. Lactescent, fruticose or suffruticose lianas, very rarely be- coming suberect. Stems volubile, terete, the branches oppo- site, or opposite below becoming alternate above. Leaves op- posite, entire, penninerved, eglandular; nodes stipulate or ex- stipulate. Inflorescence lateral, or both terminal and lateral, thyrsiform to simply scorpioid, bracteate, multiflorous to sub- uniflorous. Calyx 5-parted, the lobes essentially equal to con- spicuously unequal, cleft nearly to the receptacle, imbricated, bearing within 5 to many alternate or indefinitely distributed squamellae. Corolla infundibuliform or infrequently salver- form, the tube straight or very slightly gibbous, exappendicu- late within, the limb actinomorphie, 5-parted, dextrorsely con- volute. Stamens 5, the anthers connivent and agglutinated to the stigma, consisting of 2 parallel sporangia borne ventrally near the apex of an enlarged, sagittate, narrowly 2-lobed, pel- tate connective; pollen granular; filaments short, subcylin- drical, usually densely pilosulose. Carpels 2, united at the apex by a common stylar shaft surmounted by the fusiform or subeapitate, inconspicuously maniculate or 5-digitate stigma; (304) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 271 ovules many, several-seriate, borne upon an axile, binate pla- centa. Nectaries 5, usually concrescent, rarely more or less separate. Follicles 2, apocarpous, terete or more or less flat- tened laterally, dehiscing along the ventral suture, containing many dry, truncate, apically comose seeds. Type species: Odontadema Hoffmannseggiana | (Steud.) Woodson, in Gleason & A. C. Smith, Bull. Torrey Bot. Club 60: 392. 1933. KEY TO THE SUBGENERA AND SECTIONS A. Inflorescence thyrsiform; calyx-lobes unequal; stipules present, ag ae stems conspicuously lenticellate at maturity........ Subgen pg eon B. pear definitely infundibuliform, the throat conspicuously Fols id (ex- n O. gracilipes) ; stipules 2, laminate, searious...Sect. l. VERRUCOSAE BB. an salverform to subsalverform, the tube dilating but slightly; stipules several, filiform, subfoliaceous............... ect. 2, ANOMALAE AA. Inflorescence not thyrsiform; calyx-lobes essentially equal; stipules absent or al indefinite; stems not lenticellate, or very inconspicuously so A ad Up eno ubgen. II. EUODONTADENIA B. mie 2.5-8 em. long; stigma fusiform, frequently maniculate. C. Inflorescence both terminal and lateral; nectary deeply and indefi- nitely multifid; anthers densely hirsutulose dorsally.............. Sect. 3. HOFFMANNSEGGIANAE CC. Inflorescence lateral; nectary 5-lobed, the divisions entire or merely crenulate; anthers glabrous to puberulent-papillate dorsally...... on roit ro cM rt PEU. d et. 4. NITIDAE BB. Corolla 1.8-2.1 em. long; stigma subcapitate, digitate...Sect. 5. LAXIFLORAE Cro ux.............0.-.<.«< 0... 0.001... .0+.0..0.0.... Subgen. I. AnısoLogus (A. DC.) Woodson, comb. nov. Anisolobus A. DC. in DC. Prodr. 8: 395. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 110. 1860; Miers, Apoc. So. Am. 168. 1878, pro gen. Corolla-tube straight, or slightly gibbous or arcuate; calyx- lobes unequal, the outer shorter and broader than the inner ; in- florescence thyrsiform; stipules present, interpetiolar, cadu- cous; stems conspicuously lenticellate. Sects. 1-2. Sect. 1. VERRUCOSAE Woodson. Corolla definitely infundi- buliform, the throat conspicuously and abruptly dilated (ex- cept in O. gracilipes where subsalverform) ; inflorescence ter- minal, or both terminal and lateral, the subtending leaves op- posite; stipules 2, laminate, searious. Spp. 1-17. (305) [Vor. 22 212 ANNALS OF THE MISSOURI BOTANICAL GARDEN KEY TO THE SPECIES a. Corolla infundibuliform. Stamens inserted within the corolla-tube proper below the dilation of the roat. e. Calyx-lobes 0.5—0.9 em. long. d. Corolla densely and minutely velutinous-papillate without, at least 1. O. cognata in 34 DUA ii eR EX PA dd. Corolla glabrous without, very rarely with inconspicuous, scattering trichomes. e. Corolla-throat about 1.5 em. in diameter, abruptly T from pose A Re unigera ee. Corolla-throat 0. y. 7 em. in med, gradually on "wt the proper tube. f. Corolla-throat 2.6-3.0 em. long, 0.3-0.4 em. in diameter, the lobes 0.7—0.9 em. long; leaves 12-15 em. long......... 3. O. surinamensis ff. Corolla-throat 1.9-2.1 em. long, 0.7 em. in diameter, the lobes 1.8-2.0 cm. pig leaves 6-10 em. long.............. 4. O. neglecta ec, Calyx-lobes 0.2-0.55 e d. Leaves not cordate; ectary fleshy, essentially entire or obseurely and irregularly 5- cleft, or multifid. e. Corolla densely and fien velutinous-papillate without, at least O. verrucosa E. PR A ee. Corolla glabrous without. rescence lateral; corolla-throat narrowly conical, about iu GIMMONOE irr var . O. affinis ff. Inflorescence terminal; corolla-throat broadly conical, ape 1 nb a MIMO A ahaa ees GE owe . O. Killipú dd. Leaves obscurely cordate; nectary membranaceous, cuiu IOUS ciu aaa RA . O. cordigera bb. Stamens inserted at the base of the corolla-throat e. Corolla-throat campanulate to conical- eotaoleta: leaves ii uri a PP y MER a kiA CERA VE Yu O. lut ec. Corolla-throat tubular to subtubular; leaves never cordate. d. Corolla ANS: velutinous-papillate without, at least in the bud. utea e. Leaves glabrous throughout.................o..... 10. O. puncticulosa ee. =p minutely tomentulose beneath...........o.o.ooo... . O. spoliata dd. Corolla glabrous without. e. Calyx-lobes as long as the corolla-tube proper, or nearly so...... VAR EP EE a TERT Evo UD I d ee bd GNE 2. O. Perrottetü ee. Calyx-lobes manifestly shorter than the — dg pt T. Calya giabrons wi... ana en . O. Sandwithiana Calyx more or less densely puberulent- an wi d t. g. Outer ealyx-lobes nearly equalling the inner, the latter inis ló the length of the corolla-tube proper; species of Sout meriea. h. Inflorescence terminal, many-flowered — ; corolla- throat about 3.5 em. long.................. 4. O. lauretiana (306) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE. IV 219 hh. Inflorescence lateral, few-flowered (2-6); corolla-throat 2.0— AD um. JON. oct hese coa ERA 15. O. Kochii gg. Outer calyx-lobes much shorter than the inner, the latter about 2% the length of the corolla-tube proper; plants of British 1. Odontadenia cognata (Stadelm.) Woodson, Ann. Mo. Bot. Gard. 18: 546. 1931. Echites cognata Stadelm. dun 24': Beibl. 79. 1841; A. DC. in DC. Prodr. 8: 470. Anisolobus cognatus Cue EMEN: in Mart. Fl. Bras. 6*: 113. 1860. Angadenia cognata (Stadelm.) Miers, Apoc. So. Am. 176. 1878, Odontadenia augusta Woodson, loc. cit. 548. 1931. Stems relatively stout, dull reddish-brown, finely puberu- lent-papillate when young, glabrate and conspicuously lenticel- late when fully mature; leaves opposite, petiolate, broadly el- liptie to oblong-elliptic, occasionally to obovate-elliptie, apex acute to acuminate, base obtuse to attenuate, 8-17 cm. long, 3- 9 em. broad, firmly membranaceous to chartaceous, either sur- face glabrous, opaque; petioles 0.9-2.0 em. long; stipules 2, ovate-trigonal, acuminate, 0.3-0.4 cm. long, scarious, caducous; inflorescenee corymbose-thrysiform, terminal, about equalling or somewhat surpassing the subtending leaves, bearing 20—50 yellow, reddish-tinged flowers; pedicels 1.0-1.5 cm. long, some- what accrescent in fruit, densely and minutely puberulent; bracts ovate-trigonal 0.2-0.4 em. long, scarious, caducous; calyx-lobes broadly ovate to ovate-oblong, obtuse or rounded, unequal, 0.55-0.8 em. long, searious, densely puberulent-papil- late without, the internal squamellae in alternate groups of 2- 6; corolla infundibuliform, densely velutinous-papillate with- out, the proper-tube 1.2-2.0 em. long, about 0.35 em. in diameter at the base, the throat narrowly conical to conical-campanulate, 2-3 em. long, 0.5-1.0 em. in diameter at the orifice, the lobes broadly dolabriform, obtuse or rounded, 1.5-1.8 cm. long, widely spreading; stamens inserted in the corolla-tube proper below the dilation of the throat, the anthers narrowly elliptic to linear, 0.4—0.6 cm. long, papillate dorsally; ovary broadly (307) [Vor. 22 274 ANNALS OF THE MISSOURI BOTANICAL GARDEN ovoid, 0.1-0.15 cm. long, minutely velutinous to glabrate; stigma fusiform, usually broadly 2-lobed, 0.2-0.3 em. long; nec- taries concrescent, entire or more or less undulate or crenulate, l5 to % as long as the ovary; mature follicles unknown. PANAMA: COLON: high hills back = Puerto Obaldia, San Blas Coast, alt. 50-200 m., Aug., 1911, Pittier 4329 (B, 8, BRAZIL: PARA: Belem do Para, a secundaria, loco humido, Febr. 1, 1928, Ducke 21585 (B); Santa Izabel, ad viam ferream, Belem-Braganca, silva paludosa, Sept. 22, 1908, Ducke 21624 (B); vicinity of Para, Dec. 28, 1907, Baker 84 (D); AMAZONAS: in sylvis Japurensibus, Jan., year lacking, Martius s.m. (M, TYPE, MBG, photograph and analytical drawings); campo, Fortaleza, Oct., 1901, Ule 5923 (B). PERU: LORETO: Flutfreier Hochwald, Mündung de Santiago, alt. 160 m., Sept. 11, 1924, Tessmann 4009 (B, D, S); same locality, Oct. 21, 1924, Tessmann 4356 (B, D); lower Rio Huallaga, alt. 155-210 m., Oct.-Nov., 1929, Williams 4024 (B, FM); forest, Mishuyacu, near Iquitos, alt. 100 m., Oet.-Nov., 1929, Klug 422 (FM) ; same locality, Dec., 1929, Klug 657 (FM, US). The rather disintegrated geographical distribution of this species has not been clarified by the discovery of morphological characters which might be used for taxonomie subdivision. In the variability of leaf outline and dimensions of the corolla . cognata vividly recalls O. Hoff mannseggiana, which has a very similar distribution. 2. Odontadenia funigera Woodson, spec. nov. Suffruticosa volubilis altitudine ignota; ramulis teretibus crassiusculis cortice rubro-brunneis conspicue lenticellatis ; foliis oppositis petiolatis late ovalibus apice abrupte AURA basi obtusis 6-25 em. longis 4.5-12.0 em. latis utrinque opacis ; petiolis 1-2 em. longis; stipulis ovatis acutis 0.2-0.4 em. longis scariaceis caducis; inflorescentiis corymboso-thyrsiformibus terminalibus lateralibusque 6-20-floris pedunculo petiolos ca. bis terve superante; pedicellis 1.3-1.5 em. longis post maturi- tatem paulo accrescentibus minute denseque papillatis; brac- teis ovatis 0.2-0.3 em. longis scariaceis caducis; calycis laciniis ovato- oblongis obtusis rotundatisve inaequalibus 0.6-0.8 em. longis scariaceis extus minute papillatis intus cum sequentibus alternatis 2-4-glanduligeris; corollae infundibuliformis ful- gide flavae extus omnino glabrae tubo proprio 1.5-1.7 em. longo basi ca. 0.25 em. diametro metiente faucibus anguste campanu- (308) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 219 latis vel late tubulosis abrupte dilatatis plus minusve ventri- cosis 2.7-3.0 em. longis ostio ca. 1.5 em. diametro metiente lobis oblique obovatis obtusis rotundatisve ea. 3 em. longis patulis; staminibus sub dilatatione faucium insertis antheris linearibus anguste aurieulatis 0.6 em. longis dorso minute papillatis; ovario latissime ovoideo ca. 0.15 em. alto minute puberulo- papillato; stigmate fusiformi ca. 0.3 em. longo; nectariis con- crescentibus subintegris ovario ca. bis brevioribus; folliculis ignotis. Stems relatively stout, dull reddish-brown, glabrous, con- spicuously lenticellate when fully mature; leaves opposite, petiolate, broadly oval, apex abruptly acute, base obtuse, 6- 25 em. long, 4.5-12.0 em. broad, either surface opaque; petioles 1-2 em. long; stipules 2, ovate, acute, 0.2-0.4 em. long, scarious, caducous; inflorescence corymbose-thyrsiform, terminal and lateral, bearing 6-20 bright yellow flowers; peduncle about 2-3 times longer than the subtending petioles; pedicels 1.3-1.5 cm. long, slightly accrescent after maturity, minutely and densely papillate; bracts ovate, 0.2-0.3 em. long, scarious, caducous; ealyx-lobes ovate-oblong, obtuse or rounded, unequal, 0.6- 0.8 em. long, scarious, without minutely papillate, the internal squamellae in alternate groups of 2-4; corolla infundibuliform, glabrous without, the proper-tube 1.5-1.7 cm. long, about 0.25 em. in diameter at the base, the throat narrowly campanu- late or broadly tubular, abruptly dilated, more or less ventri- cose, 2.7-3.0 em. long, about 1.5 em. in diameter at the orifice, the lobes obliquely obovate, obtuse or rounded, about 3 cm. long, widely spreading ; stamens inserted below the dilation of the eorolla-throat, the anthers linear, narrowly auriculate, 0.6 em, long, minutely papillate dorsally; ovary broadly ovoid, about 0.15 em. long, minutely puberulent-papillate ; stigma fusi- form, about 0.3 em. long; nectaries concrescent, subentire, about 1% as long as the ovary; follicles unknown. RAZIL: AMAZONAS: sandy woods, Manáos, alt. 25 m., Oct. 13, 1929, Killip $ Smith 30048 (MBG, TYPE, US). The relationships of this species, previously confused with O. cognata, are set forth in the key to species. (309) [Vor. 22 216 ANNALS OF THE MISSOURI BOTANICAL GARDEN 3. Odontadenia surinamensis Woodson, spec. nov. Suffrutieosa volubilis omnino glabra altitudine igmota; ramulis teretibus gracilibus cortice rubro-brunneis conspicue lenticellatis; foliis oppositis breviter petiolatis ovalibus apice acutis vel brevissime acuminatis basi obtusis 12-15 em. longis 9-7 em. latis utrinque opacis; petiolis 1.0-1.5 em. longis; stipu- lis haud visis; inflorescentiis corymboso-thyrsiformibus termi- nalibus ca. 20-40-floris pedunculo petiolos subaequante; pedi- cellis 0.8-1.0 em. longis post maturitatem paulo aceres- centibus; bracteis ovato-lanceolatis acuminatis 0.2-0.4 cm. longis scariaceis caducis; calycis laciniis oblongis obtusis rotundatisve inaequalibus 0.6-0.7 em. longis extus minute papillatis intus eum sequentibus alternatis 2-4-glanduligeris ; eorollae infundibuliformis viridi- vel flavidulo-albidae extus glabrae vel indistinete papillatae tubo proprio 1.0-1.3 cm. longo basi ca. 0.1 em. diametro metiente faucibus angustis- sime conicis 2.6-3.0 em. longis ostio ca. 0.3-0.4 em. diametro metiente lobis late dolabriformibus obtusis 0.7—0.9 em, longis patulis; staminibus sub dilatatione faucium insertis antheris anguste elliptico-oblongis acuminatis anguste auriculatis 0.45 em. longis dorso minute papillatis; ovario late ovoideo ca. 0.15 em. longo glabro; stigmate fusiformi apice obtuse 2-lobato 0.2 em. longo; nectariis concrescentibus subintegris ovario ca. bis brevioribus; follieulis ignotis, Plants glabrous; stems relatively slender, dull reddish- brown, conspieuously lenticellate when fully mature; leaves opposite, shortly petiolate, oval, apex acute to very shortly acuminate, base obtuse, 12-15 em. long, 5-7 em. broad, either surface opaque; petioles 1.0-1.5 em. long; stipules caducous, not seen; inflorescence corymbose-thyrsiform, terminal, bear- ing about 20—40 greenish- or yellowish-white flowers; peduncle about as long as the subtending petioles; pedicels 0.8-1.0 cm. long, somewhat accrescent after maturity; bracts ovate-lan- ceolate, acuminate, 0.2-0.4 em. long, scarious, caducous; calyx- lobes oblong, obtuse or rounded, unequal, 0.6-0.7 em. long, mi- nutely papillate without, the internal squamellae in alternate groups of 2-4; corolla infundibuliform, glabrous or indis- tinetly papillate without, the proper-tube 1.0-1.3 em. long, (310) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 971 about 0.1 em. in diameter at the base, the throat very narrowly conical, 2.6-3.0 em. long, about 0.3-0.4 em. in diameter at the orifice, the lobes broadly dolabriform, 0.7—0.9 em. long, widely spreading; stamens inserted below the dilation of the corolla- throat, the anthers narrowly elliptic-oblong, acuminate, nar- rowly auriculate, 0.45 em. long, minutely papillate dorsally; ovary broadly ovoid, about 0.15 em. long, glabrous; stigma fusiform, bluntly 2-lobed, 0.2 em. long; nectaries concrescent, nearly entire, about half as tall as the ovary ; follicles unknown. DUTCH GUIANA: fluv. Tapanahoni, July, 1904, Versteeg 680 (U, TYPE, MBG, photograph and analytical drawings). Differs from O. cognata chiefly in the much shorter inflores- cence, narrower, glabrous corolla, and narrower calyx-lobes, as set forth in the key to species. This species has previously been confused with O. puncticulosa. 4. Odontadenia neglecta Woodson, spec. nov. Suffruticosa volubilis; ramulis teretibus crassiusculis cor- tice rubro-brunneis conspicue lenticellatis glabris; foliis op- positis breviter petiolatis ovato-ellipticis apici acutis basi ob- tusis subrotundatisve 6-10 cm. longis 2.5-6.0 em. latis firme membranaceis omnino glaberrimis opacis subtus nervo medio venisque conspicuis subverrucosis; petiolis 0.7-0.9 em. longis; stipulis interpetiolaribus 2 late oblongo-lanceolatis scariaceis caducis; inflorescentiis thyrsiformibus plurifloris et terminali- bus et lateralibus folia superantibus; pedicellis 0.5-0.7 cm. longis post maturitatem paulo accrescentibus glabris; bracteis oblongo-lanceolatis 0.1-0.3 em. longis scariaceis caducis ; calycis laciniis conspicue inaequalibus oblongo-ovatis rotundatis arcte imbrieatis exterioribus 0.5-0.6 em. longis interioribus 0.7—0.9 em. longis scariaceis extus glabris vel marginibus minute cilio- latis intus basi in marginibus 2-glanduligeris ; corollae infundi- buliformis luteae extus omnino glabrae tubo proprio gracile eylindrieo 1.0-1.2 cm. longo basi ca. 0.15 em. diametro metiente faucibus gradatim dilatatis anguste eylindrico-conieis 1.9-2.1 em. longis ostio ea. 0.7 em. diametro metiente lobis oblique obovatis brevissime acuminatis 1.8-2.0 em. longis patulis; sta- minibus sub dilatatione faucium insertis antheris oblongo- (311) [Vor. 22 278 ANNALS OF THE MISSOURI BOTANICAL GARDEN linearibus anguste 2-auriculatis acuminatis 0.5 cm. longis apice minutissime papillatis caeterum glabris; ovario late ovoideo minute papillato ca. 0.1 em. longo; stigmate anguste fusiforme 0.2 em. longo; nectario alte annulato subintegro ovarium sub- aequante; follieulis ignotis. Stems relatively stout, glabrous, reddish-brown, conspicu- ously lenticellate when fully mature; leaves opposite, shortly petiolate, ovate-elliptie, apex acute, base obtuse to rounded, 6— 10 em. long, 2.5-6.0 em. broad, firmly membranaceous, gla- brous, opaque, the midrib and veins conspicuously subverru- cose beneath ; petioles 0.7-0.9 em. long; stipules 2, interpetiolar, broadly oblong-lanceolate, scarious, caducous; inflorescence thyrsiform, several-flowered, both terminal and lateral, some- what surpassing the subtending leaves; pedicels 0.5-0.7 em. long, somewhat acerescent at maturity, glabrous; bracts ob- long-lanceolate, 0.1—0.3 em. long, scarious, caducous; calyx- lobes conspicuously unequal, oblong-ovate, rounded, closely imbricated, the outer 0.5-0.6 em. long, the inner 0.7-0.9 cm. long, scarious, glabrous without, or the margins minutely cilio- late, the internal squamellae in alternate pairs; corolla infundi- buliform, glabrous without, the proper-tube 1.0-1.2 em. long, about 0.15 em. in diameter at the base, the throat narrowly eylindrieal-eonieal, 1.9-2.1 em. long, about 0.7 em. in diameter at the orifice, the lobes obliquely obovate, very shortly acumi- nate, 1.8-2.0 cm. long, spreading; stamens inserted somewhat below the dilation of the corolla-throat, the anthers oblong- linear, narrowly 2-auriculate, 0.5 em. long, very minutely papil- late toward the tips, otherwise glabrous; ovary broadly ovoid, minutely papillate, about 0.1 em. long; stigma narrowly fusi- form, 0.2 em. long; nectary annular, subentire, about equalling the ovary ; follieles unknown. BRAZIL: AMAZONAS: prope Panure ad Rio Uaupes, Oct. 1852-Jan. 1853, Spruce 2748 (K, TvPE, MBG, photograph and analytical drawings). 5. Odontadenia n (R. & S.) K. Sch. ex Mgf. in Pulle, Fl. Surinam 4: 53. 1932. Echites verrucosa R. € S. Syst. 4: 795. 1819; A. DC. in DC. Prodr. 8: 475. 1844. (312) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE. IV 279 Echites amazonica Stadelm. Flora 24': Beibl. 50. 1841; A. DC. loc. cit. 464. 1844. Amsolobus amazonicus (Stadelm.) Muell.-Arg. in Mart. Fl. Bras. 6': 114. 1860. Echites bicornis R. Spruce, ex Muell.-Arg. loc. cit. 1860, nom. nud. in synon. Anisolobus amazonicus (Stadelm.) Muell.-Arg. f. lati- folus Muell.-Arg. loc. cit. 1860. Amisolobus Sprucei Muell.-Arg. loc. cit, 1860. Angadenia Amazonica (Stadelm.) Miers, Apoc. So. Am. 175. 1878. Angadenia latifolia (Muell.-Arg.) Miers, loc. cit. 176. 1878. Angadenia Sprucei (Muell.-Arg.) Miers, loc. cit. 1878. Odontadema Sprucei (Muell.-Arg.) K. Sch. in Engl, & Prantl, Nat. Pflanzenfam. 4?: 169. 1895. Odontadenia amazonica (Stadelm.) Malme, Arkiv f. Bot. SLA 16. 1925 Stems relatively stout, minutely puberulent when very young, glabrate and conspicuously lenticellate when fully ma- ture; leaves opposite, petiolate, rather narrowly obovate-ellip- tie, apex shortly and obtusely acuminate, base acute, somewhat cuneate, 9-16 cm. long, 3.5-6.5 em. broad, subcoriaceous, either surface glabrous, opaque, the lower somewhat paler; petioles 0.7—1.0 cm. long; stipules 2, broadly dentiform, about 0.1 em. long, scarious, caducous; inflorescence thyrsiform, terminal, densely canescent, somewhat shorter than the subtending leaves, bearing 12-30 greenish, cream-colored flowers; pedicels 1.0-1.7 cm. long, somewhat accrescent after maturity; bracts minutely ovate, 0.1-0.2 em. long, scarious, caducous; calyx- lobes ovate to broadly ovate-oblong, obtuse or rounded, un- equal, 0.3-0.5 em. long, scarious, minutely and densely papillate without, the internal squamellae in alternate groups of 2-3; corolla infundibuliform, densely and minutely velutinous-pap- illate without, the proper-tube 0.8-1.0 em. long, about 0.2 em. in diameter at the base, the throat narrowly tubular-conical, 1.5-2.3 em. long, about 0.5 em. in diameter at the orifice, the lobes obliquely obovate-oblong, obscurely acuminate, 0.9- (313) [Vor. 22 280 ANNALS OF THE MISSOURI BOTANICAL GARDEN 1.5 em. long, widely spreading ; stamens inserted just below the dilation of the corolla-throat, the anthers elliptic-linear, nar- rowly auriculate, 0.4-0.6 em. long, minutely hirtellous dorsally ; ovary oblongoid, about 0.15 em. long, densely puberulent-pap- illate; stigma fusiform-capitate, with 5 inconspicuous basal projeetions, 0.2 em. long; nectaries conerescent, subentire or minutely erenulate, about as long as the ovary ; follicles rela- tively stout, divaricate, 12-15 em. long, densely canescent with- out; seeds about 1 cm. long, the pale tawny coma about 2 cm. long. BRAZIL: PARA: insulae Brenes aestuarii amazonici, Sept. 6, 1901, Guedes 21800 (B); in ripa Ygarape et Tagipuru, Aug., year lacking, Martius 2663 (M, MBG, photograph and analytieal drawings); AMAZONAS: prope Panure ad Rio Uaupes, Oct., 1852-Jan., 1853, Spruce 2503 (B, Bx, Camb., D); ad flum. Guiania v. Rio Negro supra ostium fluminis Casiquiari, 1854, Spruce 3550 (Bx, Camb., V, MBG, photograph and analytical drawings); Panure ad Rio Uaupes, Oct., 1852-Jan., 1853, Spruce 2555 (C, Camb., G, MBG, photograph and analytical drawings). Spruce 2555 and Spruce 2503, referred to Anisolobus Sprucei and A. amazonicus respectively by Mueller-Argoviensis, are virtually indistinguishable except for the very slightly more copious indument of the former. The collector’s data do not contradict the assumption that both were from the same local- ity, if not from the same plant or colony. 6. Odontadenia affinis Woodson, Ann. Mo. Bot. Gard, 18: 549. 1931. Stems relatively slender, glabrous, reddish-brown, rather in- conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, oblong-ovate, apex shortly and obtusely sub- cuspidate-acuminate, base acutish, 6-8 cm. long, 3.5-4.0 em. broad, subcoriaceous, glabrous, opaque; petioles 0.5-0.8 em. long; inflorescence lateral, thyrsiform, relatively few-flowered, somewhat shorter than the subtending leaves; bracts very in- conspicuous; pedicels about 0.5 em. long, Abro calyx-lobes ovate, obtusish, more or less conspicuously unequal, 0.4- 0.55 em. long, atrodi: or the margins very minutely ciliolate, the squamellae in alternate groups of 3—4; corolla infundibuli- form, glabrous without, the proper tube 1.5-1.7 cm. long, about (314) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 281 0.25 cm. in diameter at the base, the throat conical, 2.0-2.3 em. long, about 0.55 em. in diameter at the orifice, the lobes ob- liquely obovate, 1.5-2.0 em. long, spreading ; stamens inserted somewhat below the dilation of the corolla-throat, the anthers narrowly lanceolate-sagittate, 0.5 em. long, very minutely pap- illate dorsally; ovary broadly ovoid, about 0.15 em. long, gla- brous; stigma fusiform, about 0.2 cm. long; nectary annular, erenulate, much shorter than the ovary ; follieles unknown. PERU: LORETO: dense forest, Balsapuerto, alt. 150—350 m., Aug. 28—30, 1929, Killip $ Smith 28609 (US, TYPE, MBG, photograph and analytical drawings). 7. Odontadenia Killipii Woodson, Ann. Mo. Bot. Gard. 18: 546. 1931. Stems relatively stout, glabrous, conspicuously lenticellate when fully mature; leaves opposite, petiolate, broadly elliptic- oblong, apex abruptly and obtusely acuminate, base obtuse to acutish, 6-8 cm. long, 3-5 cm. broad, subcoriaceous, glabrous; petioles 1.0-1.5 cm. long, glabrous; inflorescence terminal, thyrsiform, somewhat surpassing the subtending leaves, bear- ing 15-20 showy, yellowish flowers, glabrous throughout; pedi- cels 0.5-0.7 cm. long; bracts ovate, 0.2-0.3 em. long, scarious; calyx-lobes conspicuously unequal, ovate to ovate-oblong, ob- tuse, 0.2-0.3 cm. long, glabrous or essentially so, the internal squamellae solitary or in alternate pairs; corolla infundibuli- form, glabrous without, the proper tube 1.0-1.5 em. long, about 0.2 em. in diameter at the base, the throat conical, 2.0-2.5 em. long, about 1 em. in diameter at the orifice, the lobes obliquely obovate-dolabriform, 2.5-3.0 em. long, spreading; stamens in- serted somewhat below the dilation of the corolla-throat, the anthers linear-sagittate, 0.3-0.4 em. long, minutely puberulent- papillate dorsally ; ovary ovoid-oblongoid, about 0.15 cm. long, glabrous; stigma fusiform, 0.2 em. long; nectary tubular, en- tire or minutely crenulate, somewhat surpassing the ovary; follicles unknown. PERU: LORETO: woods, Iquitos, alt. about 100 m., Sept. 26, 1929, Killip $ Smith 29847 (MBG, TYPE, US); Flussfreier Urwald, Iquitos, alt. 100 m., May 9, 1925, Tessmann 5100 (B). (315) [Vor. 22 282 ANNALS OF THE MISSOURI BOTANICAL GARDEN 8. Odontadenia cordigera Woodson, spec. nov. Suffruticosa volubilis omnino glaberrima altitudine ignota ; ramulis crassiusculis maturitate cortice rubro-brunneis dense lenticellatis; foliis oppositis breviter petiolatis obovatis apice rotundatis non rarius sensim emarginatis basi obscure cordatis 7-8 em. longis 4-6 cm. latis utrinque opacis ; petiolis 0.5-0.7 em. longis ; stipulis 2 late reniformibus 0.2-0.3 em. longis scariaceis caducis; inflorescentiis corymboso-thyrsiformibus lateralibus ea. 15-30-floris folia subaequantibus; pedicellis 0.4—0.5 em. longis, post maturitatem paulo aeerescentibus ; bracteis minute ovatis ca. 0.1 em. longis seariaceis caducis; calycis laciniis ovatis vel late trigonalibus obtusis rotundatisve inaequalibus 0.3-0.4 em, longis scariaceis extus glabris intus basi cum se- quentibus alternatis 1—2-glanduligeris; corollae infundibuli- formis fulgide flavae extus glabrae tubo proprio 0.9-1.1 em. longo basi ea. 0.15 em. diametro metiente faucibus anguste conicis 2.0-2.3 em, longis ostio ca. 1.2 cm. diametro metiente lobis oblique obovatis ca. 1.7 em. longis patulis; staminibus sub dilatatione faucium insertis antheris linearibus acuminatis anguste auriculatis ca. 0.4 cm. longis dorso indistinete papil- latis; ovario oblongoideo ca. 0.1 em. longo glabro ; stigmate fusi- formi ca. 0.2 em. longo; nectariis imperfecte conerescentibus manifeste inaequalibus tenuibus ovario multo brevioribus; follieulis desiderantibus. Plants eompletely glabrous; stems relatively stout, reddish- brown, densely lenticellate when fully mature; leaves opposite, shortly petiolate, obovate, apex rounded, not infrequently some- what emarginate, base obscurely cordate, 7-8 em. long, 4—6 em. broad, either surface opaque ; petioles 0.5-0.7 em, long; stipules 2, broadly reniform, 0.2-0.3 cm. long, scarious, caducous; in- florescence corymbose-thyrsiform, lateral, about equalling the subtending leaves, bearing 15-30 bright yellow flowers; pedi- cels 0.4—0.5 em. long, somewhat acerescent at maturity; bracts minutely ovate, about 0.1 em. long, scarious, caducous; calyx- lobes ovate or broadly trigonal, obtuse or rounded, unequal, 0.3-0.4 em. long, scarious, glabrous without, the internal alter- nate squamellae solitary or paired; corolla infundibuliform, glabrous without, the proper-tube 0.9-1.1 em. long, about 0.15 (316) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 283 em. in diameter at the base, the throat narrowly conical, 2.0- 2.3 em. long, about 1.2 cm. in diameter at the orifice, the lobes broadly obovate, about 1.7 em. long, widely spreading; sta- mens inserted somewhat below the dilation of the corolla- throat, the anthers linear, acuminate, narrowly auriculate, about 0.4 em. long, indistinetly papillate dorsally; ovary oblongoid, about 0.1 em. long, glabrous; stigma fusiform, about 0.2 cm. long; nectaries imperfectly concrescent, manifestly unequal, tenuous, much shorter than the ovary; follicles unknown. PBRU: LORETO: dense forest, Mishuyacu, near Iquitos, alt. about 100 m., Sept. 24-28, 1929, Killip $ Smith 29916 (MBG, TYPE, US). 9. Odontadenia lutea (Vell.) Mgf. in Fedde, Rep. Sp. Nov. 20: 24. 1924. Echites lutea Vell. Fl. Flum, 109. 1830; Icon. 3: pl. 25. 1827; A. DC. in DC. Prodr. 8: 467. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 159. 1860; Miers, Apoc. So. Am. 194. 1878. Echites Zuccariniana Stadelm. Flora 241: Beibl. 76. 1841; A. DC. loc. cit. 471. 1844, Anisolobus hebecarpus Muell.-Arg. loc. cit. 111. pl. 33. 1860; Miers, loc. cit. 170. 1878. Amsolobus hebecarpus Muell.-Arg. a. tomentosus Muell.- Arg. loc. cit. 112. 1860. Amisolobus hebecarpus Muell.-Arg. a. tomentosus Muell.- Arg. a. erectus Muell.-Arg. loc. cit. 1860. Amsolobus hebecarpus Muell.-Arg. a. tomentosus Muell.- Arg. b. scandens Muell.-Arg. loc. cit. 1860. Amsolobus hebecarpus Muell.-Arg. B. glabratus Muell.- Arg. loc. cit. 1860. Echites pulcherrima Pohl, ex Muell.-Arg. loc. cit. 1860, nom. nud. in synon, Echites hebecarpa Benth, ex Muell.-Arg. loc. cit. 1860, nom. nud. in synon. Anisolobus Zuccarinianus (Stadelm.) Miers, loc. cit. 171. 1878. Amisolobus pulcherrimus Miers, loc. cit. 1878. (317) [VoL. 22 284 ANNALS OF THE MISSOURI BOTANICAL GARDEN Odontadenia Zuccariniana (Stadelm.) K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 169. 1895. Odontadenia Zuccarimana (Stadelm.) K. Sch. f. angusti- folia Malme, Bihang till K. Sv. Vet. Akad. Handl. Afd. III. 241": 23. 1899. Odontadenia Zuccariniana (Stadelm.) K. Sch. f. ovalifolia Malme, loc, cit. 1899. Odontadema Zuccarimana (Stadelm.) K. Sch. var. tomen- tosa Muell.-Arg. f. scandens Muell.-Arg, ex Mgf. Notiz- blatt 9: 80. 1924, sphalm. Plants densely and minutely hirtellous to glabrate or glabrous; stems relatively stout, becoming conspicuously lenti- cellate when fully mature; leaves opposite, the blade subcori- aceous, broadly oval to narrowly oblong-elliptie, apex obtuse or rounded to broadly acuminate, base narrowly and rather ob- seurely eordate, 7-15 em. long, 3-6 em. broad, drying dark brownish-green, either surface opaque, the lower somewhat paler, the petiole 0.5-0.75 em. long; stipules 2, oblong, 0.5-0.75 em. long, caducous; inflorescence lateral to subterminal, con- tracted-thyrsiform, many-flowered, usually somewhat sur- passed by the subtending leaves, densely canescent to glabrate; pedicels 1.5-2.0 em. long, somewhat acerescent at maturity, the subtending bracts ovate-lanceolate, 0.4—0.6 em. long, caducous; calyx-lobes conspicuously unequal, ovate to broadly oblong, broadly obtuse or rounded, 0.75-1.25 em. long, the squamellae solitary or rarely geminate, narrowly liguliform; corolla in- fundibuliform, whitish tinged with yellow, glabrous without, the proper-tube cylindrical, somewhat inflated at the base, con- tracting toward the insertion of the stamens, 1.75-2.0 em. long, about 0.4 em. in diameter at the base, the throat broadly conical- campanulate, 2.0-2.25 em. long, about 1.25 em. in diameter at the orifice, the lobes obliquely obovate, shortly acuminate, 2.25-3.0 em. long, widely spreading; stamens inserted at the base of the corolla-throat, the anthers narrowly elliptic-sagittate, 0.8 cm. long, minutely hirtellous dorsally ; carpels oblong-ovoid, 0.2 em. long, minutely papillate; stigma fusiform-capitate, with 5 ob- scure basal lobes, 0.2 em. long; nectary annular, deeply 5-lobed, (318) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 285 about twice surpassing the ovary; follicles relatively stout, divaricate, 15-20 em. long, glabrous to minutely velutinous (ac- cording to Muell.-Arg.) ; seeds unknown. BRAZIL: PARA: in vieinibus Santarem, June, 1850, Spruce s.n. (C, Camb., V, MBG, photograph); RIO DE JANEIRO: environs de Rio de Janeiro, Febr., 1882, Glaziou 12957 (C, K): MINAS GERAES: Trahiras, date lacking, Pohl 1899 (V); Corallino, date lacking, Pohl 1383 (V); sho PAULO: inter Canna Verde et S. João, Febr., 1849, Regnell III 880 (S); Batataes, Febr., 1849, Regnell III 850 (S); ing Grosso: Serra do Itapirapuan, in campo dumetoso (‘‘cerrado’’), April 29, 4, Lindmann A3337 (S); Cuyaba, in ‘‘cerrado,’’ April 10, 1894, Malme 1536 (S); Sta. Anna da Chapada, in ‘‘cerrado,’’ July 29, 1902, Malme s.n. (S). Odontadenia lutea is extremely variable in leaf size and out- line, and particularly so in indument. Mueller recognized sev- eral subspecifie categories in dealing with this species, and justification may exist for such a treatment, Nevertheless, the numerous specimens examined during the course of this study have revealed such a complicated range of intergradations that it has been resolved to postpone subspecific division until the genus ean be studied more intimately, preferably by a stu- dent resident in the South American tropies. 10. Odontadenia puncticulosa (A. Rich.) Pulle, Enum. Pl. Surinam 383. 1906.1 Echites ee Rich. Act. Soc. Hist. Nat. Paris 1: 107. ; E. Mey. Nov. Aet. Acad. Nat. Cur. 12: 782. TA Echites Cururu Mart. in Buchn. Rep. Pharm. 101. 1830; Stadelm. Flora v Beibl. 78. 1841; A. DC. in DC. Prodr. 8: 470. 184 Echites Cururu T var. a. angustifolia Stadelm. loc. cit. 1841 Echites Cururu Mart. var. P. grandifolia Stadelm. loc. cit. 79. 1841; A. DC. loc. cit. 1844. Anisolobus Kappleri Miq. Linnaea 18: 737. 1844. * Originally peas by Pulle citing E. Mey. parenthetically. Meyer, however, in the work cited above, gives Richard as authority for E. puncticulosa, and his extended c Rare should be construed, therefore, as supplementing the meagre diagnosis of the earlier author. (319) [Vor. 22 286 ANNALS OF THE MISSOURI BOTANICAL GARDEN Anisolobus puncticulosus (A. Rich.) Miq. Stirp. Surinam. 158. 1851; Miers, Apoc. So. Am. 172. 1878. Anisolobus Fockei Miq. loc. cit. 159. 1851; Miers, loc. cit. 1878. Anisolobus Hostmanni Miq. loc. cit. 1851; Miers, loc, cit. 1878. Anisolobus Cururu (Mart.) Muell.-Arg. in Mart. Fl. Bras. 61: 112. pl. 34. 1860. Anisolobus Cururu (Mart.) Muell.-Arg. B. grandifolius (Stadelm.) Muell.-Arg. loc. cit. 113. 1860. Angadenia Cururu (Mart.) Miers, loc. cit. 175. 1878, Angadenia grandifolia (Stadelm.) Miers, loc. cit. 1878. Odontadenia Cururu (Mart.) K. Sch, in Engl. & Prantl, Nat. Pflanzenfam. 4?: 169. 1895. Stems relatively stout, glabrous or minutely puberulent when young, conspicuously lenticellate when fully mature; leaves opposite, the blade subcoriaceous, ovate- to oblong-el- liptie, apex shortly and obtusely acuminate, base obtuse to broadly acute, 8-20 em. long, 3.5-10.0 em. broad, either surface opaque, glabrous, drying dark brown, the lower somewhat paler, the petiole 0.5-1.0 em. long; stipules 2, broadly ovate, 0.2-0.3 em. long, caducous; inflorescence terminal, thyrsiform, many-flowered, somewhat surpassing the subtending leaves, minutely canescent; pedicels 0.75-1.0 em. long, somewhat ac- erescent at maturity, the subtending braets minutely ovate- reniform ; ealyx-lobes conspicuously unequal, ovate to broadly ovate-oblong, obtuse or rounded, 0.3-0.5 em. long, the squamel- lae geminate; corolla infundibuliform, minutely papillate to glabrate without, the proper-tube cylindrical, slightly gibbous- inflated toward the base, sharply constricted toward the inser- tion of the stamens, 1.0-1.5 em. long, about 0.3 em. in diameter at the base, the throat narrowly tubular-conical, 2.0-2.75 em. long, 0.6-0.75 em. in diameter at the orifice, the lobes obliquely obovate-oblong, rather obscurely acuminate, 1.75-2.25 em. long, widely spreading; stamens inserted at the base of the corolla- throat, the anthers narrowly oblong-elliptic, 0.5-0.6 em. long, densely hirtellous dorsally ; ovary oblongoid, 0.2 em. long, gla- brous; stigma obtusely fusiform with 5 inconspicuous, acute, (320) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 287 basal projections, 0.2 em. long; nectary tubular, completely concealing the ovary; follicles unknown. DUTCH GUIANA: e regione Surinam. super., date lacking, Wullschlägel 321 (V); in virgultis, date lacking, Hostmann $ Kappler 1203 (Camb., MBG, S); data in- complete, Hostmann 629 (S). AZIL: AMAZONAS: Vista Alegre, Rio Branco, Jan. 9, 1924, Kuhlmann 21858 (S); Ega, date lacking, Poeppig 2657 (V); PARA: Santa Izabel, ad viam ferream Belem-Braganca, Oct., 1906, Goeldi 21777 (S). 11. Odontadenia spoliata Malme, Arkiv f. Bot. 21A*: 16. 1927. Stems relatively stout, dull reddish-brown, minutely puberu- lent-papillate when very young, glabrate and conspicuously lenticellate when fully mature; leaves opposite, shortly petio- late, broadly ovate-elliptic, apex acute, base broadly obtuse or rounded, 11-14 em. long, 6-9 em. broad, firmly membranaceous to chartaceous, either surface opaque, the upper glabrous, the lower minutely tomentulose; petioles 1.0-1.5 em. long; stipules 2, ovate-oblong, acute, 0.3-0.4 em. long, scarious, caducous; in- florescence thyrsiform, terminal, somewhat shorter than the subtending leaves, bearing 6-12 cream-colored flowers; pe- duncle about twice as long as the petioles, densely canescent ; pedicels 1.5-1.7 em. long, somewhat accrescent after Bahr densely and minutely canescent; braets ovate-lanceolate, 0.2— 0.4 em. long, scarious, caducous; calyx-lobes ovate, obtuse, un- equal, 0.45-0.5 em. long, scarious, papillate without, the inter- nal squamellae in alternate groups of 3-6; corolla infundibuli- form, minutely velutinous-papillate without, the proper-tube 1.3-1.6 cm. long, about 0.5 em. in diameter at the base, the throat broadly tubular, 2.7-3.3 em. long, about 1 em. in diam- eter at the orifice, the lobes obliquely obovate, obtuse or rounded, 1.3-1.6 em. long, widely spreading; stamens inserted at the base of the corolla-throat, the anthers narrowly oblong- elliptie, acuminate, narrowly auriculate, about 0.6 em. long, mi- nutely papillate dorsi ovary ovoid, about 0.5 cm. long, gla- brous or indistinctly puberulent-papillate; nectaries concres- cent, irregularly erose, somewhat surpassing the ovary; fol- licles unknown. BRAZIL: MATTO GROSSO: in silva ripa rivi, Sta. Anna da Chapada, July 20, 1902, Malme 2036 (S, TYPE, MBG, photograph and analytical drawings). (321) [Vor. 22 288 ANNALS OF THE MISSOURI BOTANICAL GARDEN The unusual foliar indument appears to be the chief mark of distinetion between this species and O. puncticulosa, of which it might better be interpreted as a variety. 12. Odontadenia Perrottetii (A. DC.) Woodson, Ann. Mo. Bot. Gard. 18: 546. 1931. Anisolobus Perrottetii A. DC. in DC. Prodr. 8: 395. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 115. 1860; Miers, Apoc. So. Am. 169. 1878. Amsolobus Perrottetii A. DC. p. obtusus Muell.-Arg. loc. eit. 1860. Stems relatively stout, dull yellowish-brown, glabrous, con- spicuously lenticellate when fully mature; leaves opposite, pet- iolate, broadly oval to oblong-elliptic, apex rather abruptly acuminate, base broadly obtuse or rounded, 7-13 em. long, 2- 6 em. broad, subeoriaceous, either surface glabrous, opaque, or the upper somewhat nitidulous; petioles 0.8-1.5 em. long; sti- pules ovate-trigonal, acute, 0.3-0.5 em. long, searious, cadu- cous; inflorescence corymbose-thyrsiform, both terminal and lateral, bearing 2-10 pale yellowish flowers; peduncle glabrous, somewhat less than half as long as the subtending leaves ; pedi- cels 1.0-1.5 em. long, glabrous; bracts ovate, 0.2-0.3 em. long, scarious, caducous; calyx-lobes ovate to ovate-oblong, broadly obtuse to rounded, 0.8-1.2 em. long, scarious, glabrous without, the internal squamellae in alternate groups of 4—6; corolla in- fundibuliform, glabrous without, the proper-tube 0.9-1.5 cm. long, about 0.3 em. in diameter at the base, the throat tubular- conical, 2.0-2.2 em. long, about 0.6 em. in diameter at the orifice, the lobes obliquely obovate, rounded, 1.2-1.5 em. long, widely spreading; stamens inserted at the base of the corolla-throat, the anthers narrowly oblong, acuminate, narrowly and sharply auriculate, 0.6 em. long, finely papillate dorsally; ovary ovoid, about 0.15 em. long, glabrous; stigma fusiform, about 0.2 em. long; nectaries concrescent, irregularly lacerate, concealing the ovary, about 0.2 cm. long; immature follicles relatively stout, divaricate, minutely puberulent-papillate. FRENCH GUIANA: exact locality lacking, 1821, Perrottet 270 (DO, TYPE). BRAZIL: PARA: campos do Ariramba, regione fl. Trombetas, Oct. 9, 1913, Ducke (322) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 289 216647 (B); MARANHÃO: exact locality and date lacking, Martius s.n. (Bx); MATTO GROSSO: Chiquitos, 1842, d'Orbigny 879 (Bx, D). 13. Odontadenia Sandwithiana Woodson, Ann. Mo. Bot. Gard. 18: 547. 1931. Plants completely glabrous; stems terete, relatively slender, dull yellowish-brown, conspicuously lenticellate when fully mature; leaves opposite, shortly petiolate, lanceolate- to ob- long-elliptie, abruptly and obtusely subcaudate-acuminate, base acute, 6-15 cm. long, 2-6 em. broad, subcoriaceous, oliva- ceous, the lower surface somewhat pallid; petioles 0.8-2.0 cm. long; stipules 2, broadly dentiform, 0.1-0.2 em. long, scarious, caducous; inflorescence corymbose-thyrsiform, lateral (occa- sionally subterminal ?), bearing 6-18 yellowish flowers; pe- duncle 2-3 times longer than the subtending petioles; pedicels 1.8-3.0 cm. long, somewhat accrescent after maturity; calyx- lobes ovate to obovate-oblong, obtuse to rounded, unequal, 1.0- 1.5 em. long, scarious, glabrous, the internal squamellae in alternate groups of 2-4; corolla infundibuliform, the proper- tube 1.5-2.0 em. long, about 0.2 em. in diameter at the base, the throat tubular-conical, 1.4—1.6 cm. long, about 0.2-0.3 em. in diameter at the orifice, the lobes obovate-dolabriform, obtuse or rounded, 0.7-1.0 cm. long, spreading; stamens inserted at the base of the corolla-throat, the anthers narrowly oblong, acuminate, narrowly auriculate, 0.5-0.6 cm. long, glabrous; ovary oblongoid, about 0.2 em. long, glabrous; stigma fusiform, 0.2 em. long; nectary tubular, subentire or minutely crenulate, somewhat surpassing the ovary ; follieles unknown. BRITISH GUIANA: Essequibo River, Moraballi Creek, near Bartica, alt. near sea- level, Nov. 2, 1929, Sandwith 552 (K, TYPE, MBG, photograph and analytical drawings); Amakura River, Northwest District, Lat. 8” 10' N., Long. 60? W., March 23-30, 1923, Cruz 3542 (MBG, NY). 14. Odontadenia lauretiana Woodson € Steyermark, spec. nov. Suffruticosa volubilis ; ramulis teretibus crassiusculis cortice obscure flavo-brunneis juventate minute sparseque puberulis maturitate inconspicue lenticellatis; foliis oppositis petiolatis elliptico-lanceolatis apiee acuminatis basi acutis 9-19 em. (323) [VoL, 22 290 ANNALS OF THE MISSOURI BOTANICAL GARDEN longis 3-4 em. latis firme membranaceis omnino glabris opacis obseure olivaeeis subtus pallidioribus; petiolis 1.2-1.7 em. longis glabris; stipulis haud visis; inflorescentiis thyrsiformi- bus subeorymbosis terminalibus 30—50-floris; pedunculo glabro folia subaequante; pedicellis 1.5 em. longis post maturitatem paulo aeerescentibus glabris; bracteis ovato-lanceolatis 0.1- 0.3 em. longis scariaceis caducis; calycis laciniis ovato-oblongis rotundatis paulo inaequalibus 0.6-0.65 em. longis minutissime papillatis margine ciliolatis intus basi cum sequentibus alter- natis uniglanduligeris; corollae infundibuliformis pallide gilvae extus minute sparseque papillatae tubo proprio 1.2- 1.4 em. longo basi ea. 0.2 em. diametro metiente prope apicem sensim angustato ibique staminigero faucibus tubulo-conicis 3.5 em. longis ostio ca. 0.6 cm. diametro metiente lobis oblique obovatis ea. 1.5-1.8 em. longis patulis; antheris anguste ob- longo-elliptieis acuminatis anguste auriculatis ca. 0.5 cm. longis dorso glabris; ovario ovoideo minute papillato ca. 0.2 em. longo; stigmate fusiformi ca. 0.2 em. longo; nectariis concrescentibus margine minute erenulatis ovario ca. dimidio brevioribus ; folliculis ignotis. Stems terete, relatively stout, minutely and sparsely puberu- lent when young, glabrate, dull yellowish-brown, inconspicu- ously lenticellate when fully mature; leaves opposite, petiolate, elliptic-lanceolate, apex acuminate, base acute, 9-12 em. long, 3-4 em. broad, firmly membranaceous, glabrous, dull olive- green, opaque, the lower surface somewhat paler; petioles 1.2- 1.7 em. long, glabrous; stipules not seen; inflorescence thyrsi- form, subeorymbose, terminal, bearing 30-50 cream-colored flowers; peduncle glabrous, about as long as the subtending leaves; pedicels about 1.5 em. long, somewhat accrescent after maturity, glabrous; braets ovate-lanceolate, 0.1-0.3 em. long, scarious, caducous; calyx-lobes ovate-oblong, rounded, some- what unequal, scarious, minutely papillate without, margin ciliolate, the internal squamellae alternate, solitary ; corolla in- fundibuliform, minutely and sparsely papillate without, the proper-tube 1.2-1.4 em. long, about 0.2 em. in diameter at the base, markedly tapering toward the apex, the throat tubular- conical, about 3.5 em. long, about 0.6 em. in diameter at the ori- (324) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 291 fice, the lobes obliquely obovate, 1.5-1.8 cm. long, widely spreading; stamens inserted at the base of the corolla-throat, the anthers narrowly oblong-elliptic, acuminate, narrowly auriculate, about 0.5 em. long, glabrous; ovary ovoid, about 0.2 em. long, minutely papillate; stigma fusiform, about 0.2 em. long; nectaries concrescent, minutely crenulate, about half equalling the ovary ; follicles unknown. PERU: LORETO: edge of forest, Fortaleza, Yurimaguas, alt. 155-210 m., Oet. 30, 1929, Williams 4385 (B, FM, TYPE, MBG, photograph and analytical drawings). In many respects O. lawretiana provides a connecting link be- tween subgen. Anisolobus and Ewodontademia. The calyx- lobes are nearly equal, although closely imbricated as in other species of Anisolobus. The form of the corolla and the aspect of the foliage are likewise similar to those found in that sub- genus. On the other hand, the inflorescence is thyrsiform, thus combining structural characters of the two subgenera. If stipules are proved to be absent, an additional link to Euodon- tadenia will be provided. 15. Odontadenia Kochii Pilger in Koch-Gruenberg, Zwei Jahre unter den Indianern 2: 371. 1910; in Fedde, Rep. Sp. Nov. 8: 151. 1910. Stems relatively stout, glabrous, dark reddish-brown, con- spieuously lenticellate when fully mature; leaves opposite, petiolate, oblong- to ovate-elliptie, apex abruptly and shortly acuminate, base obtuse to rounded, 6-10 cm. long, 3-6 em. broad, firmly membranaceous to chartaceous, either surface glabrous, opaque, the lower somewhat paler; petioles 0.8- 1.0 em. long; stipules 2, broadly ovate, 0.1-0.2 cm. long, sca- rious, caducous; inflorescence corymbose, lateral, bearing 2— 6 pale yellowish flowers; peduncle somewhat longer than the subtending petioles, glabrous; pedicels 1.3-1.5 cm. long, some- what accrescent after maturity, glabrous; bracts ovate-lanceo- late, 0.1—0.2 em. long, scarious, caducous; calyx-lobes ovate to ovate-oblong, obtuse to rounded, 0.7-0.9 em. long, scarious, mi- nutely puberulent-papillate to glabrate without, the internal squamellae in alternate groups of 3-4; corolla infundibuliform, glabrous without, the proper-tube 1.4-2.0 em. long, about (325) [Vor. 22 292 ANNALS OF THE MISSOURI BOTANICAL GARDEN 0.3 em. in diameter at the base, the throat tubular-conical, 2.0- 2.6 em. long, about 0.7 em. in diameter at the orifice, the lobes obliquely obovate, inconspicuously acuminate, 1.3-1.6 em. long, widely spreading; stamens inserted at the base of the corolla- throat, the anthers oblong-elliptie, acuminate, narrowly auricu- late, 0.5 em. long, minutely papillate dorsally; ovary ovoid, about 0.15 em. long, glabrous; stigma fusiform-capitate, about 0.15 em. long; nectaries concrescent, irregularly 5-lobed, about as long as the ovary; follicles relatively stout, divaricate, 8- 12 em. long, densely and minutely velutinous to glabrate, seeds unknown. BRAZIL: AMAZONAS: in Pflanzung neben Maloka, Dec., 1903, Koch 70, 73 (B, TYPE, MBG, photograph and analytical drawings); Camanáos, Rio Negro, Dec. 22-93, 1930, Holt $: Blake 592 (MBG, US); same data, Holt § Blake 594 (US). 16. Odontadenia Schippii Woodson, spec. nov. Frutieosa volubilis alte scandens; ramulis crassiusculis juventate minutissime puberulis cortice flavo-brunneis con- spicue lenticellatis; foliis oppositis petiolatis late ellipticis : ovalibusve apice in acumen brevissimum obtusum abrupte produetis basi obtusis 6.5-12.0 em. longis 3.0-5.5 em. latis firme membranaceis glabris supra subnitidulis subtus opacis pal- lidioribusque; petiolis 1.0-1.5 cm. longis glabris; inflores- centiis thyrsiformibus plurifloris et terminalibus et lateralibus folia aequantibus vel paulo superantibus; pedunculo brevis- sime puberulo-papillato; pedicellis 2.25-2.5 em. longis minute denseque puberulo-papillatis; calycis laciniis manifeste in- aequalibus exterioribus late ovatis obtusis rotundatisve 0.55- 0.6 em. longis eoriaceis extus minute denseque puberulo-papil- latis interioribus late oblongis late obtusis 0.9-1.0 em. longis firme membranaceis extus minute sparseque puberulis; corol- lae speciosae gilvae extus omnino glabrae intus prope inser- tionem staminum puberulae caeterumque glabrae ut dicitur paululo suaveolentis tubo proprio 1.5-1.7 em. longo basi ca. 0.35—0.4 cm. diametro metiente faucibus tubulo-conicis 2.5— 2.7 em. longis ostio ca. 0.5-0.6 em. diametro metiente lobis ob- lique lateque dolabriformibus breviter acuminatis 1.3-1.5 cm. longis patulis; staminibus prope basem faucium insertis (326) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 293 antheris oblongo-linearibus acuminatis 0.6 em. longis dorso minute hirtellis; ovariis ovoideis ca. 0.1 em. longis glabris; stigmate fusiformi ca. 0.225 em. longo; nectariis concrescenti- bus carnosis obfuniformibus ca. 0.3 em. longis; folliculis ignotis. Stems relatively stout, minutely puberulent when young, deep brown, conspicuously lenticellate; leaves opposite, peti- olate, broadly elliptie to oval, apex shortly and obtusely acumi- nate, base obtuse, 6.5-12.0 em. long, 3.0-5.5 em. broad, firmly membranaceous, glabrous, above somewhat subnitidulous, opaque and paler beneath; petioles 1.0-1.5 em. long, glabrous; inflorescence thyrsiform, both terminal and lateral, equalling or somewhat surpassing the subtending leaves, bearing several handsome, creamy-white flowers said to be slightly fragrant; pedunele puberulent-papillate; pedicels 2.25-2.5 em. long, pu- berulent-papillate; ealyx-lobes manifestly unequal, the outer broadly ovate, obtuse or rounded, 0.55-0.6 em. long, coriaceous, minutely and densely puberulent-papillate without, the inner broadly oblong, obtuse or rounded, 0.9-1.0 em. long, firmly membranaceous, minutely and sparsely puberulent without; corolla infundibuliform, glabrous without, puberulent within at the insertion of the stamens, otherwise glabrous, the proper tube 1.5-1.7 cm. long, 0.35—0.4 em. in diameter at the base, the throat tubular-conical, 2.5-2.7 em. long, about 0.5-0.6 em. in diameter at the orifice, the lobes obliquely and broadly dolabri- form, shortly acuminate, 1.3-1.5 em. long, spreading; stamens inserted at the base of the corolla-throat, the anthers oblong- linear, acuminate, 0.6 cm. long, minutely hirtellous dorsally; ovaries ovoid, about 0.1 em. long, glabrous; stigma fusiform, about 0.225 em. long; nectaries concrescent, fleshy, obfuniform, about 0.3 em. long, deeply enclosing the ovary; follicles unknown. British HONDURAS: Camp 36, Guatemala-B.H. survey, alt. 2800 ft., June 18, 1934, Schipp S-709 (MBG, TYPE). Mr. Schipp describes this species as follows: ‘‘Tall vine growing in mountain forest, fairly common, but difficult to col- lect as it grows on the largest of forest trees. Flowers creamy (327) [ VoL. 22 294 ANNALS OF THE MISSOURI BOTANICAL GARDEN white with slight odor. 60 ft., stem 3 inches in diameter.’’ Odontadenia Schippiiis of great interest not only because it is by far the northernmost representative of its genus, but be- cause of its close affinities with several South American species of rather limited distribution, as indieated in the key to species. 17. Odontadenia gracilipes (Stadelm.) Woodson, comb. nov. Echites gracilipes Stadelm. Flora 24!: Beibl. 22. 1841; A. DC. in DC, Prodr. 8: 455. 1844. Anisolobus ? gracilipes (Stadelm.) Muell.-Arg. in Mart. Fl. Bras. 6!: 115. 1860. Mitozus gracilipes (Stadelm.) Miers, Apoc. So. Am. 220. 1878. Odontadenia goyazensis Glaziou, Bull. Soc. Bot. France 57: Mem. 3e.: 455. 1910, nom. nud. Stems terete, relatively slender, ferruginous-pilosulose when young, eventually glabrate and inconspicuously lenticellate when fully mature ; leaves opposite, petiolate, oval, apex acumi- nate, base acute to obtuse, 7-12 cm. long, 2-5 em. broad, sub- eoriaeeous or firmly chartaceous, opaque, glabrous above, beneath rather irregularly and generally pilosulose; petioles 0.4-1.2 em. long, pilosulose; stipules minute, flagelliform, numerous; inflorescence corymbose-thyrsiform, lateral, bear- ing 3-8 pale yellowish flowers; peduncle about 2 to 3 times longer than the subtending petioles, pilosulose; pedicels 1.3- 1.4 em. long, somewhat accrescent after maturity, minutely pu- berulent-papillate to minutely pilosulose; bracts minutely ovate-reniform, scarious, caducous; calyx-lobes ovate-oblong, unequal, rounded, 0.5-0.8 em. long, scarious, glabrous or essen- tially so, the internal squamellae solitary, or in alternate pairs; corolla subsalverform, glabrous without, the tube 1.0-1.4 cm. long, about 0.2 em. in diameter at the base, the lobes obovate- dolabriform, 0.7-0.8 em. long, widely spreading; stamens in- serted slightly below midway within the corolla-tube, the anthers narrowly oblong-elliptie, shortly acuminate, narrowly aurieulate, about 0.4 em. long, glabrous; ovary ovoid, about 0.15 em. long, glabrous ; nectaries concrescent, subentire, nearly equalling the ovary ; follieles unknown. (328) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 295 BRAZIL: MINAS GERAES: in sylvis prope Rio Piranga, April, 1818, Martius 1034 (M, TYPE, MBG, photograph and analytical drawings); DATA INCOMPLETE: Sellow 692 (B); Glaziou 20416 (B). Sect. 2. ANoMaALAE Woodson. Corolla salverform to subsal- verform, the tube gradually dilating toward the orifice, but without a conspieuously expanded throat; inflorescence ter- minal, the subtending leaves indefinitely congested or sub- verticillate ; stipules numerous, filiform, subfoliaceous. Sp. 18. 18. Odontadenia anomala (Heurck € Muell.-Arg.) Macbr. Field Mus. Publ. Bot. 11: 35. 1931. Anisolobus anomalus Heurck € Muell.-Arg. in Heurck, Obs. Bot. 160. 1870. i Perictenia stipellaris Miers, Apoc. So. Am. 183. pl. 28. 1878. Echites stipellaris Spruce ex Miers, loc. cit. 1878, nom. nud. in synon. Stems relatively stout, densely ferruginous-hirtellous to glabrate; leaves opposite or irregularly verticillate at the ends of branches, the blade firmly membranaceous, broadly obovate to obovate-oblong, apex rounded with an abrupt and short acumen, base broadly obtuse, 10-20 em. long, 6.0-13 em. broad, above drying dark brownish-green, minutely and rather sparsely hirtellous to glabrate, beneath much paler, densely and persistently tomentulose, the petiole 1.0-1.5 em. long; stip- ules numerous, filiform, 0.75-1.0 em. long, subfoliaceous; in- florescence terminal, densely fasciculate-thyrsiform, many- flowered, somewhat shorter than the subtending leaves, mi- nutely and densely hirtellous; pedicels 0.75 cm. long, somewhat accrescent at maturity, the subtending bracts ovate to ovate- lanceolate, 0.5-0.75 em. long, persistent; calyx-lobes conspicu- ously dissimilar, broadly ovate, obtuse or rounded, 0.3-0.4 cm. long, densely puberulent without, the squamellae solitary, deeply and irregularly cleft and divided; corolla deep yellow flushed with orange, salverform, glabrous, or minutely and sparsely pilosulose above, the tube narrowly cylindrical, some- what dilated at the orifice, 2.5—3.0 em. long, about 0.2 em. in diameter at the base, the lobes obliquely obovate, acuminate, 3.25-4.0 em. long, widely spreading; stamens inserted nearly (329) [Vor. 22 296 ANNALS OF THE MISSOURI BOTANICAL GARDEN at the base of the corolla-tube, the anthers narrowly oblong- linear, 0.9-1.0 em. long, densely puberulent dorsally toward the tip; earpels ovoid, about 0.1 em. long, densely lanulose; stigma fusiform, 0.15—0.2 em. long; nectary tubular, irregularly erose, about twice as long as the ovary, sparsely and minutely pilosu- lose ; follicles unknown. PERU: LORETO: Tarapoto, 1855-56, Spruce 4900 (B, C, Camb., K, V, MBG, photograph and analytical drawings); Stromgebiet des Maranon, Santiago- Mündung am Pongo de Manseriche, 1924, Tessmann 4034 (S). Subgen. II. EivoroxTApENIA Woodson, subgen. nov. Corolla-tube straight, never gibbous or arcuate; calyx-lobes equal or essentially so; inflorescence variously cymose, but not thyrsiform; stipules absent or extremely indefinite; stems not lenticellate, at least above, or very inconspicuously so. Sects. 3-5. Sect. 3. HoFFMANNSEGGIANAE Woodson. Inflorescence both terminal and lateral, a variously compounded dichasium, fre- quently aggregate; corolla broadly infundibuliform, the proper tube eonspieuously shorter than the throat; anthers densely hirsutulose dorsally; nectary deeply and indefinitely multifid. Spp. 19-20. KEY TO THE SPECIES a. Corolla 5-8 em. long, the throat longer than broad..... 19. 0. Hoff mannseg giana aa. Corolla 3—4 em. long, the throat about as broad as long.............. a a ee EEE 20. O. stemmadeniaef olia 19. Odontadenia Hoffmannseggiana (Steud.) Woodson, ex Gleason & A. C, Smith, Bull. Torrey Bot. Club 60: 392. 1933. Echites grandiflora G. F. W. Meyer, Fl, Esseq. 131. 1818, not Roxb., Roth, Stadelm., ete. Echites macrantha R. & S. Syst. 4: 795. 1819, not Spreng. Echites Hoff mannseggiana Steud. Nomencl. ed. 2. 1: 539. 1840. Echites grandiflora Hoffmsg. ex Steud. loc. cit. 1840, nom. nud. in synon. Odontadenia speciosa Benth. in Hook. Jour. Bot. 3: 242. 1841; A. DC. in DC. Prodr. 8: 360. 1844; Muell.-Arg. in Mart. Fl. Bras. 6': 117. 1860; Miers, Apoc. So. Am. 126, pl. 16. 1878. (330) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 297 Haemadictyon ? grandiflorum (G. F. W. Meyer) A. DC. loc. cit. 426. 1844. Echites Meyeriana R. & S. ex A. DC. loc. cit. 1844, nom. nud. in synon. Echites sylvestris A. DC. loc, cit. 464. 1844. Odontadenia grandiflora Miq. Stirp. Surinam. 166. 1851. Dipladenia Harris Purdie in Hook. Bot. Mag. IIT, 11: pl. 4825. 1855. Cylicadenia Harristi Lem. in Van Houtte, Illustr. Hort. 2: Misc. 9. 1855. Odontadenia sylvestris (A. DC.) Muell.-Arg. loc. cit. 1860. Dipladenia Harrisonii Purdie ex Muell.-Arg. Linnaea 30: 446. 1860, sphalm. Odontadenia grandiflora (G. F. W. Meyer) Miers, loc. cit. 127. 1878. Odontadenia formosa Miers, loc, cit. 1878. Odontadenia Harrisú (Purdie) Miers, loc. cit. 128. 1878. Cycladenia Harrison; Lemaire, ex Miers, loc cit. 1878, sphalm in synon. Angadema sylvestris (A. DC.) Miers, loc. cit. 174. 1878. Odontadenia grandiflora (G. F. W. Meyer) O. Ktze. Rev. Gen. 416. 1891, sphalm. Odontadenia grandiflora (Q. F. W. Meyer) K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 169. 1895, sphalm. Plants completely glabrous; stems relatively stout; leaves opposite, the blade firmly membranaceous or chartaceous, broadly elliptie to obovate-lanceolate, apex rather abruptly and shortly subcaudate-acuminate, infrequently acute in older leaves, base acute to obtuse, 13-22 em. long, 4-10 em. broad, either surface opaque, or the upper somewhat glossy, the petiole 1.25-2.0 em. long; stipules obsolete; inflorescence lat- eral, infrequently subterminal, dichasially eymose, frequently much reduced, about equalling the subtending leaves, not rarely much shorter, bearing few to many flowers; pedicels 2.0-2.5 em. long, somewhat aecrescent at maturity, the sub- tending bracts minutely ovate, persistent; calyx-lobes es- sentially equal, broadly ovate, obtuse, 0.4-0.6 cm. long, the squamellae solitary, or occasionally geminate; corolla infundi- (331) [Vor. 22 298 ANNALS OF THE MISSOURI BOTANICAL GARDEN buliform, deep yellow tinged with red or orange, the proper- tube 0.5-1.0 em. long, 0.3-0.5 em. in diameter at the base, con- spieuously constricted at the insertion of the stamens, the throat conical, 2.5-3.5 em. long, 1.25-1.75 em. in diameter at the orifice, the lobes obliquely obovate, broadly acuminate, 2.0-3.25 em. long, widely spreading ; stamens inserted at the base of the eorolla-throat, the anthers elliptie-sagittate, 1.25—1.3 em. long, densely hirtellous dorsally; ovary ovoid, about 0.2 cm. ne; glabrous; stigma fusiform, 0.4 em. long; nectary annular, deeply and indefinitely multifid, about half equalling the ovary ; follicles relatively stout, more or less falcate, 15-30 em. long, 1-2 em, in diameter, glabrous ; seeds 3-4 em. long, the abundant, pale yellowish coma of about equal length. CosTA RICA: foréts sur les bords du Rio Coto, March, 1896, Pittier 9881 (Bx). PANAMA: PANAMA: Trinidad River, alt. 40-80 m., 1911, Pittier 3976 (G, US); in dense woods, Twin Hill, May, 1862, Hayes 663 (BB, Bx, Camb., M, V); COLON: Perme, San Blas district, > 24, 1933, Cooper 261 (MBG, NY RITISH GUIANA: upper Rupununi River, near eig May 30, 1922, Cruz 1418 (MBG NY); d data, Schomburgk 309 (B, € D, V). DurcH Qua: ad fl. Marowyne mediam, date DAE Xs 1989 (B, 8); e reg. Para, date lacking, Wullschlägel 1971 (Bx, V). FRENCH GUIANA: Acarouany, 1857, Sagot 383 (D, S, V); Cayenne, date lacking, Martin s.n. (B) BRAZIL: AMAZONAS: Ilha do Frio, Rio Branco, March, 1913, Kuhlmann 3119 (B, S, U); PARA: Parana do Mirity-pueu (prope faueem fluvii Tocantins), ad ripas io vam May 1, 1924, Kuhlmann 21856 (S); ad oram mer. fl. Amazonum ad ostium flum. Solimoes, June, 1851, Spruce 1615 (Bx); MATTO GROSSO: On varzia land, river shore, near Tabajaza, upper Machado River region, Nov.-Dee., 1913, Krukoff 1457 (MBG, NY). PERU: LORETO: forest, Mishuyacu, near Iquitos, alt. 100 m., Febr.-March, 1930, Klug 1026 (US). In founding the genus Odontadema upon O. speciosa (= O. Hoff mannseggiana), Bentham (loc. cit.) remarked that the im- mature seeds available for his study were without an apical coma. Sagot, however, noted upon the labels of his specimen cited above: ** Flor. aureo lutei ampli, semina papposa!”” In all other species of Odontadenia of which seed is known, furthermore, the usual apical coma has been observed. Never- theless, Bentham's opinion concerning the lack of a coma in O. speciosa has been maintained, even by recent authors (ef. Markgraf, in Pulle, Fl. Surinam 4: 48. 1932). Several fruit- (332) 1935] WOODSON—-STUDIES IN THE APOCYNACEAE. IV 299 ing specimens have been examined during the course of this study, and it may be stated unequivocably that the seeds of all were found surmounted by an abundant coma. A particularly handsome specimen upon which the seminal coma may be ob- served is Kuhlmann 3119 (in Herb. Berol.). 20. Odontadenia stemmadeniaefolia Woodson, Ann. Mo. Bot, Gard. 18: 548. 1931. Plants completely glabrous; stems relatively stout; leaves opposite, shortly petiolate, obovate-oblong, apex shortly and obtusely acuminate, base more or less conspicuously cuneate, 15-20 em. long, 8-10 em. broad, firmly membranaceous, opaque; petioles 1.0-1.5 em. long; inflorescence rather irregularly dichasial or aggregate, lateral, somewhat shorter than the sub- tending leaves, bearing 3-5 showy, reddish-yellow flowers; pedicels 1.0-1.3 em. long; calyx-lobes equal or subequal, ovate- reniform, about 0.2 cm. long, the squamellae in alternate groups of 2-3; corolla infundibuliform, the proper-tube 0.8-1.0 em. long, about 0.3 em. in diameter at the base, the throat broadly conical, 0.7-1.0 em, long, about 1 em. in diameter at the orifice, the lobes obliquely obovate-dolabriform, 1.5-2.0 cm. long, widely spreading; anthers narrowly oblanceolate-sagit- tate, 0.8 em. long, densely hirtellous dorsally; ovary ovoid, about 0.15 cm. long, glabrous; nectary annular, deeply and in- definitely multifid, somewhat surpassed by the ovary ; follicles unknown. PERU: LORETO: forest, Mishuyaeu, near Iquitos, alt. 100 m., Jan., 1930, Klug 782 (US, TYPE, MBG, photograph and analytical drawings). Sect. 4. Nrrinag Woodson. Inflorescence lateral, scorpioid, usually simple; corolla salverform or narrowly infundibuli- form, the proper tube scarcely narrower than the throat; anthers glabrous to puberulent-papillate dorsally; nectary 9-lobed, the divisions entire to erenulate. Spp. 21-25. KEY TO THE SPECIES a. Stamens inserted well above the base of the corolla-tube; species of South America (including i da b. Leaves obscurely e . Corolla narrowly are leaves glabrous throughout. .21. O. nitida [ Vor. 22 300 ANNALS OF THE MISSOURI BOTANICAL GARDEN bb. Leaves not cordate. e. Corolla salverform, 5-6 em. long; calyx-lobes 0.4-0.5 em. long...... TEITTE TTT ETES cee 111213122001 11 1T TX PREIS unter ec. Corolla narrowly infundibuliform, 2.5—3.5 em. long; calyx-lobes 0.1 em. a VETTDITIQITELZCII III UI SY . aa. Stamens inserted nearly at the base of the corolla- ms Ergo of REMMI a is is . O. polyneura 21. Odontadenia nitida (Vahl) Muell.-Arg. in Mart. Fl. Bras. 6': 118. 1860; K. Sch. in Engl. € Prantl, Nat. Pflanzenfam. 42: 169. 1895. Echites nitida Vahl, Eclog. 2: 19. f. 13. 1798; A. DC. in DC. Prodr. 8: 453. 1844. Echites lucida R. € S. Syst. 4: 795. 1819; A. DC. loc. cit. 415. 1844, not Wall, Odontadenia cordata A. DC. loc. cit. 360. 1844. Odontadema angustifolia A. DC. loc. cit, 1844. Odontadenia nitida (Vahl) Muell.-Arg. a. oblongifolia Muell.-Arg. loc. cit. 119. 1860. Odontadema nitida (Vahl) Muell-Arg. f. acuminata Muell.-Arg. loc. cit, 1860. Odontadenia nitida (Vahl) Muell.-Arg. y. angustifolia (A DC.) Muell.-Arg. loc, cit. 1860, Odontadenia lucida (R. € S.) Muell.-Arg. loc. cit. 190. 1860. Angadenia nitida (Vahl) Miers, Apoc. So. Am. 177. 1878. Rhabdadenia ? lucida (R. & S.) Miers, Apoc. So. Am. 123. 1878. Mitozus tenellus Miers, loc. eit. 220. 1878. Odontadenia Dusendschoenii K, Sch. ex Ule, in Engler's Jahrb. 40: 403. 1908, nom. nud. Plants completely glabrous; stems relatively slender ; leaves opposite, the blade subcoriaceous, broadly oval to narrowly oblong-elliptie, obtuse to abruptly and shortly aeuminate, base broadly and rather obseurely eordate, 7-18 em. long, 2.5-9.0 em. broad, drying dark olive-green to brownish, shining above, paler and glaucous beneath, the petiole 1.0-1.25 cm. long; stipules obsolete or essentially so; inflorescence lateral, scor- pioid, simple, few- to many-flowered, somewhat surpassing the (334) 1935] WOODSON—STUDIES IN THE APOCYNACEAE, IV 301 subtending leaves; pedicels 0.75—1.0 em. long, somewhat ac- crescent at maturity, the subtending bracts minutely ovate- lanceolate, persistent; calyx-lobes essentially equal, ovate to ovate-lanceolate, acuminate, 0.4-0.5 cm. long, the squamellae geminate; corolla infundibuliform, glabrous without, white or pale cream, the proper tube 0.7-1.0 em. long, about 0.15 em. in diameter at the base, the throat tubular-conical, 1.7-2.3 em. long, about 0.6 em. in diameter at the orifice, the lobes obliquely obovate-dolabriform, obscurely acuminate, 1.5-1.8 cm. long, widely spreading; stamens inserted somewhat below midway within the proper tube of the corolla, the anthers narrowly elliptie-sagittate, 0.7 em. long, minutely hirtellous dorsally; ovary ovoid, about 0.15 em. long, glabrous; stigma fusiform, 0.3 cm, long; nectary annular, deeply 5- to many-lobed, about half equalling the ovary ; follicles relatively short and stout, parallel to slighty falcate, 8-12 em. long; seeds about 0.8 em. long, the pale tawny coma about 2 em. long. TRINIDAD: Chancellor’s Road, St. Ann's, March 24, 1924, Broadway 5243 (D, MBG) ; Pitch Lake, 1891-92, Warming 239 (C) ; Piarco Savanna, south of Dabadie, March 18, 1920, Britton $ Hazen 717 (G). VENEZUELA: Caura-La Botellas, 1902, Passarge $ Selwyn 287 (B). BRITISH GUIANA: locality lacking, 1858, Schomburgk 200 (B). DUTCH GUIANA: in campis (savannis), distr. Para, Febr.-Apr., 1844, Kappler 1449 (BB, MBG, S); Paramaribo, date lacking, Wullschlägel 322 (Bx, V); Re- publiek, Oct. 13, 1911, Kuiper 40 (B, U). FRENCH GUIANA: Cayenne, date lacking, Jelski s.n. (B). BRAZIL: PARA: Alemquer, prope flumen, July 28, 1903, Ducke 21644 (B); MATTO GROSSO: Cuyaba, in silvula ripa rivulis, July 17, 1902, Malme 1766 (8); Cuyaba, in dumetis ruderalibus (‘‘charravasco’’) alte volubilis, July 13, 1894, Lindmann 43563 (S, MBG, photograph) PERU: LORETO: Yurimaguas, Aug., 1902, Ule 6272 (B, D); Tarapoto, Sept., 1902, Ule 6390 (B, D). 22. Odontadenia hypoglauca (Stadelm.) Muell-Arg. in Mart. Fl. Bras. 6': 118. pl. 35, fig. 1. 1860; K. Sch. in Engl. & Prantl, Nat. Pflanzenfam. 4?: 169. 1895. Echites hypoglauca Stadelm. Flora 24!: Beibl. 23. 1841; A. DC. in DC. Prodr. 8: 448. 1844. Angadema hypoglauca (Stadelm.) Miers, Apoc. So. Am. 173. 18798. Angadenia majuscula Miers, loc, cit. 174. 1878. (335) [Vor. 22 302 ANNALS OF THE MISSOURI BOTANICAL GARDEN Stems relatively stout, glabrous; leaves opposite, the blade coriaceous, broadly oval to oblong-elliptie, apex obtuse to rounded, base broadly and obscurely cordate, 7-15 em. long, 3- 8 em. broad, upper surface glabrous, shining, lower surface glaucous, minutely and densely puberulent-papillate, the pet- iole 0.75—1.0 em. long; stipules obsolete; inflorescence lateral, simply scorpioid, glabrous, about equalling or somewhat sur- passing the subtending leaves, many-flowered; pedicels 1.0— 1.25 em. long, somewhat acerescent in fruit, the subtending braets minutely ovate-reniform, caducous; calyx-lobes essen- tially equal, ovate-oblong, acute to obtuse, 0.4—0.6 em. long, the squamellae geminate ; corolla white or pale cream, salverform, glabrous without, the tube narrowly cylindrical, slightly dilat- ing toward the orifice, 2.5—3.0 em. long, about 0.3 em. in diam- eter at the base, the lobes obliquely obovate, shortly acuminate, 2.0-2.5 em. long, widely spreading; stamens inserted slightly below midway within the corolla-tube, the anthers narrowly elliptie-sagittate, about 1 em. long, minutely papillate dorsally ; carpels ovoid, 0.2 em. long, glabrous; stigma fusiform, 2-lobed, 0.3-0.4 em. long; nectary erown-shaped, deeply 5-lobed, about half as long as the ovary ; follicles relatively stout, parallel or slightly falcate, 12-15 em. long, glabrous ; seeds unknown. BRAZIL: PARA: in vicinibus Santarem, Apr., 1850, Spruce 680 (Camb., M, V); campo non inundabili, Prainha, May 11, 1903, Ducke 21645 (B); BAHIA: Maraeas, date lacking, Martius s.n. (M); in sepibus ad Caiete, date lacking, Martius s.n. (M); Goyaz: data incomplete, Pohl 1898 (M); MATTO GROSSO: Sta. Anna da Chapada, in ora silva, Jan. 12, 1903, Malme 3305 (8, MBG, photograph and ana- lytical drawings); Cuyaba, in ‘‘cerrado,’’ May 25, 1894, Malme 1642 (8). Also reported from Pernambueo by Mueller. 23. Odontadenia geminata (R. & S.) Muell.-Arg. in Mart. Fl. Bras. 6': 118. 1860; K. Sch. in Engl. € Prantl, Nat. Pflanzen- fam. 4?: 169. 1895. Echites geminata R. & S. Syst. 4: 796. 1819; A. DC. in DC. Prodr. 8: 475, 1844. Echites elegans Benth. in Hook. Jour. Bot, 3: 249. 1841; A. DC. loc. cit. 1844. Echites coriacea Benth. loc. cit, 1841; A. DC. loc. cit. 467. 1844, not Blume. (336) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 303 Odontadenia Poeppigii Muell.-Arg. loc. cit. 119. 1860. Odontadenia coriacea (Benth.) Muell.-Arg. Linnaea 30: 450. 1860. Angadenia coriacea (Benth.) Miers, Apoc. So. Am. 177. 1878. Angadema elegans (Benth.) Miers, loc. cit. 178. 1878. Angadenia geminata (R. & S.) Miers, loc. cit. 1878. Plants eompletely glabrous; stems relatively slender; leaves opposite, the blade coriaceous or subcoriaceous, broadly oval to narrowly oblong-elliptie, apex obtuse to subacuminate, base obtuse to rounded, 6-12 em. long, 2.5—7.0 em. broad, above shin- ing, beneath paler, glaucescent; petiole 0.75-1.0 em. long; stip- ules obsolete; inflorescence lateral, simply scorpioid, about equalling or somewhat shorter than the subtending leaves, 2- 7-fowered ; pedicels 1.5-1.75 em. long, somewhat accrescent in fruit, the subtending bracts minutely ovate-lanceolate, per- sistent; ealyx-lobes essentially equal, broadly ovate, obtuse or broadly acute, 0.4—0.5 em. long, the squamellae solitary or occa- sionally geminate; corolla white or pale cream tinged with yel- low, salverform, glabrous without, the tube narrowly cylin- drieal, slightly dilated toward the orifice, 2.5-3.0 cm. long, about 0.3 em. in diameter at the base, the lobes obliquely obo- vate, shortly acuminate, 2.5-3.0 em. long, widely spreading; stamens inserted about midway within the corolla-tube, the anthers narrowly elliptic-sagittate, 0.9—1.0 em. long, minutely papillate dorsally ; ovary ovoid, about 0.15 em. long, glabrous; stigma fusiform, 0.3 em. long; nectary annular, deeply 5-lobed, somewhat shorter than the ovary ; follicles relatively short and stout, parallel to slightly faleate, 8-10 em. long, glabrous; seeds unknown. BRITISH GUIANA: Kamakusa, upper Mazaruni River, July 11-22, 1923, Cruz 4211 (MBG, NY); Kakburi, Pomeroon District, Febr. 10-15, 1923, Cruz 3251 (US); Waini River, Northwest District, April 3-18, 1923, Cruz 3703 (US). DUTCH GUIANA: Wonotobo, Oct., 1916, Stahel $ Gonggrijp 2862 (B, U). COLOMBIA: CAQUETA: Rio Caqueta ad cataractas Cupati, in ripis, Dec. 1, 1912, Ducke 21821 (B). BRAZIL: AMAZONAS: Vista Alegre, Rio Branco, in campis, March, 1913, Kuhl- mann 2917 (B, S, MBG, photograph and analytical drawings); prope San Gabriel da Cachoeira, ad Rio Negro, Jan.-Aug., 1852, Spruce 2083 (Camb., V, MBG, photo- (337) [VoL. 22 304 ANNALS OF THE MISSOURI BOTANICAL GARDEN graph); prope San Carlos, ad Rio Negro, 1853-54, Spruce 3152 (M, V); PARA: Bóa Vista on the Tapajos River, May-June, 1929, Dahlgren $ Sella 74 (B, FM, MBG, 8); Faro, eampina arenosa, Sept. 24, 1907, Ducke 21650 (B); on varzia land, near Cassipa, Tapajos River region, Sept., 1931, Krukoff 1247 (MBG, NY). 24, Odontadenia glauca Woodson, Ann. Mo. Bot. Gard. 18: 990. 1931. Plants glabrous throughout ; stems relatively slender; leaves opposite, shortly petiolate, obovate-oblong, apex shortly and acutely acuminate, base obtuse or rounded, 6-8 em. long, 3.5- 4.5 em. broad, coriaceous, lustrous above, glaucous beneath; petioles 0.3-0.5 em. long; inflorescence lateral, seorpioid, some- what shorter than the subtending leaves, bearing 3-5 white or. eream-eolored flowers; pedicels 1 em. long; calyx-lobes sub- equal, ovate-trigonal, acutish, about 1 em. long, scarcely imbri- cated, the alternate squamellae solitary; corolla infundibuli- form, glabrous without, the proper tube 0.8-1.0 em. long, about 0.1 em. in diameter at the base, the throat tubular-conical, 0.7— 1.0 em. long, the lobes obliquely dolabriform, 1.0-1.5 em. long, spreading; stamens inserted about midway within the corolla- tube, the anthers narrowly oblong-sagittate, 0.6 cm. long, gla- brous; ovary ovoid, about 0.1 em. long, glabrous; stigma 0.15 em. long; neetary deeply 5-lobed, somewhat surpassing the ovary ; follicles unknown. VENEZUELA: AMAZONAS: Cerro Yapaeana, upper Rio Orinoco, alt. about 100 m., April, 1931, Holt $ Blake 750 (US, TYPE, MBG, photograph and analytical drawings). 25. Odontadenia polyneura (Urb.) Woodson, Ann. Mo. Bot. Gard. 18: 546. 1931. Rhabdadema polyneura Urb. Symb. Ant. 7: 337. 1912. Plants completely glabrous; stems relatively slender, red- dish-brown, not lenticellate or very inconspicuously so; leaves opposite, petiolate, oblong- to obovate-elliptie, apex acute to acuminate, base obtuse to rounded, 3.5-8.0 em. long, 1-4 em. broad, firmly membranaceous, above olivaceous, slightly nitid- ulous, beneath paler and more or less glaucous, the midrib and veins somewhat verrucose; petioles 0.5-1.0 em. long; stipules obsolete; inflorescence scorpioid, simple, lateral, opposite, somewhat surpassing the leaves, bearing 6-10 white or eream- (338) 1935] WOODSON—STUDIES IN THE APOCYNACEAE. IV 305 colored, showy flowers; pedicels 0.9—1.0 em. long, somewhat ac- crescent after maturity; bracts ovate-lanceolate, 0.1—0.3 cm. long, scarious, caducous; calyx-lobes ovate, broadly acute, es- sentially equal, 0.3-0.4 em. long, glabrous without, the squamel- lae in alternate groups of 3-6; corolla infundibuliform, gla- brous without, the proper-tube 0.4—0.5 em. long, about 0.3 em. in diameter at the base, the throat narrowly campanulate, 1.3- 1.5 em. long, about 0.7-0.8 cm. in diameter at the orifice, the lobes obliquely obovate, shortly acuminate, 1.5-1.7 em. long, widely spreading; stamens inserted nearly at the base of the corolla-tube, the anthers narrowly elliptic-lanceolate, nar- rowly auriculate, 0.8 cm. long, glabrous; ovary broadly ovoid, about 0.15 em. long, glabrous; stigma fusiform, 0.2 em. long; nectaries 5, concrescent at the base, truneate or somewhat emarginate, about half as long as the ovary ; follicles unknown. SANTO DOMINGO: inter Constanza et Rio Jimenoa in Loma del Hato quemado, alt. 1400 m., in sylvis, June, 1910, Tuerckheim 3341 (B, TYPE, MBG, 8). Also reported from Haiti by Urban (loc. eit.). Sect. 5. LaxiFLoRAE Woodson. Inflorescence both terminal and lateral, laxly and irregularly compound, dichasial to scor- pioid or helicoid; corolla infundibuliform, relatively small; anthers glabrous; nectary annular, essentially concrescent. Sp. 26. 26. Odontadenia laxiflora (Rusby) Woodson, Ann. Mo. Bot. Gard. 19: 386. 1932. Laubertia (?) laxiflora Rusby, Bull. N. Y. Bot. Gard. 4: 408. 1907 Codonechites paniculata Mgf. Notizblatt 9: 80. fig. 2 A-E. 1924. Plants completely glabrous; stems relatively slender, light greenish- to reddish-brown, not lenticellate above or very in- conspicuously so; leaves opposite, shortly petiolate, oblong- elliptie, apex shorii? subcaudate-acuminate, base acute to ob- tuse, 8-13 cm. long, 2.5—4.0 cm. broad, membranaceous to sub- chartaceous, above pale olivaceous, nitidulous, beneath some- what paler, opaque; petioles 0.4—0.6 cm. long; stipules obso- lete; inflorescence laxly and irregularly compound, dichasial (339) [ Vor. 22, 1935] 306 ANNALS OF THE MISSOURI BOTANICAL GARDEN to scorpioid or helicoid, both terminal and alternate-lateral, about twice surpassing the leaves, bearing 20-30 white or cream-colored flowers; pedicels 1.0-1.2 em. long, somewhat ac- erescent in fruit; bracts minutely ovate-trigonal, scarious; calyx-lobes ovate, broadly acute, 0.2-0.25 em. long, glabrous without, the squamellae in alternate groups of 2-4; corolla in- fundibuliform, glabrous without, the proper-tube 0.6-0.7 cm. long, about 0.15 em. in diameter at the base, the throat rather narrowly eampanulate, 0.7—0.8 em. long, about 0.5 em. in diam- eter at the orifice, the lobes obliquely obovate, shortly acumi- nate, 0.5—0.6 em. long, widely spreading ; stamens inserted at the base of the corolla throat, the anthers elliptic-lanceolate, acuminate, narrowly auriculate, 0.35 em. long, glabrous; ovary broadly ovoid, about 0.1 em. long, glabrous; stigma fusiform- subcapitate, obscurely digitate below, 0.15 em. long; nectaries essentially conerescent, truncate, fleshy, about half as long as the ovary ; follieles unknown. Bouivia: data incomplete, Bang 2056 (NY, TYPE, MBG, photograph and ana- lytical drawings). BRAZIL: Rio Acre, Seringal bei S. Francisco, March, 1911, Ule 9698 (B, MBG, photograph). It does not appear possible to recognize O. laxiflora as con- stituting a distinct genus without raising to generic rank also other entities of the inclusive genus Odontadenia interpreted as seetions in this revision. EXCLUDED OR UNCERTAIN SPECIES Odontadema cuspidata Rusby, Deser. So. Am. Pl. 89. 1920 = Mandevilla cuspidata (Rusby) Woodson, Ann. Mo. Bot. Gard. 20: 730. 1933. Odontadema glandulosa (R. € P.) K. Sch. in Engl. € Prantl, Nat. Pflanzenfam. 4?: 169. 1895 = Mandevilla glandulosa (R. & P.) Woodson, Ann. Mo. Bot. Gard. 19: 66. 1932. Odontadenia gracilis K. Sch. in Glaziou, Bull. Soc. Bot. France 57, Mem. 3e.: 455. 1910, nom. nud. Amsolobus lancifolius K. Sch. in Glaziou, loc. cit. 1910, nom. nud. (To be continued) (340) CYTOLOGY OF GEOTRICHUM VERSIFORME MOORE! MORRIS MOORE Mycologist to The Barnard Free Skin and Cancer Hospital, St. Louis Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany of Washington University INTRODUCTION The purpose of this paper is to report a study of the cytology and nuclear phenomena in the development of Geotrichum versiforme, a fungus which was described morphologically and culturally in a previous report by the author (Moore, 734). As far as the writer is aware, there has been no cytological study of the genus Geotrichum, except for the suggestion that the arthrospores were possibly uninucleate cells. Guillier- mond (700), in describing the structure of Oidium lactis, gave indications of the mycelial structure of that organism, while Jannin (713) referred to the cytology in Guilliermond's paper as representing the type present in Myeodermata. Inasmuch as Geotrichum has a structure and development often confused with and similar to Mycoderma, Oidium, and perhaps Oospora and Monilia, the work of the above two writers may be taken to represent the previous work on the subject. This may further be emphasized by the fact that certain Oidia, Mycodermata, and others have been shown to be synonymous. However, the life eyele of Geotrichum versiforme seemed, on superficial ex- amination, to differ in several factors from that of the other genera. In addition, a newer technique for work of this sort led the author to the study here described. MATERIALS AND TECHNIQUE It has been shown that the fungus has different aspects on various media, and that maltose increased the growth of the ! Preliminary report presented before the Myeologieal Society of America, De- cember 30, 1933, at the winter meeting of the American Association for the Advancement of Science, held at Boston, Massachusetts. Issued May 25, 1935. ANN. Mo. Bor. GARD., Vor. 22, 1935. (307) [ Vor. 22 308 ANNALS OF THE MISSOURI BOTANICAL GARDEN organism. Consequently, Sabouraud's glucose and maltose agar were used as substrates. The former develops all the features found in the life cycle, and the latter, because of the maltose, produced many of the irregularities in morphology, such as large selerotie cells. Cultures on agar slants of these media were fixed with Hermann's fluid, embedded in celloidin, seetioned, and stained with iron-alum haematoxylin as de- scribed by the author (Moore, ’33) for Endomyces. For the observation of the cellular constituents, as fat, gly- cogen, volutin, and the like, several chemicals were used which will be described under their respective headings. CYTOLOGY Since there has been no definite relationship established with the Ascomycetes, by means of ascospores, the arthrospore must be considered as the germ of the colony. It would thus seem ad- visable to trace the development of a mycelium with that cell as the initial stage. The single arthrospore (pl. 7, fig. 1) is uni- nucleate, with a fairly thick wall or membrane. Germination proceeds, usually laterally, by sending out a thin tube, the germ-tube, which at first has the appearance of a small bud but later becomes a thin-walled hypha. This contains many gran- ules near its apex which are probably volutin. The nucleus is seen as a large granular structure with a heavily staining nucleolus. With the germination of the arthrospore, nuclear division takes place, by simple fission as far as could be de- termined (pl. 7, fig. 3). Although many nuclear divisions were examined, it cannot be stated definitely that mitosis was pres- ent. The process continues until there may be as many as eight nuclei in the germinating cell, but usually there are only two or three (pl. 7, fig. 2). The germ cell itself contains a granular and retieulated pro- toplasm of metachromatie material. In many of the germ- tubes, there seems to be a vacuolated area near the apex (pl. 7, fig. 3). After a number of nuclei have apparently entered into the germ-tube, the hypha is ready to form cross-walls. In the young filaments this is accomplished by a pinching in or a transverse abscission of the walls of the hypha. This may take 1935] MOORE—CYTOLOGY OF GEOTRICHUM VERSIFORME 309 place between a dividing nucleus as in pl. 7, fig. 4, or between separated nuclei. The cell elongates and nuclear division con- tinues, chiefly at the terminal portion. The nuclei may divide laterally (pl. 7, figs. 6, 8), giving the appearance in a later stage of alternate nuclei (pl. 7, figs. 8, 11, 12), or longitudinally (pl. 7, figs. 9, 12). The young filaments elongate and may be simple or branched (pl. 7, figs. 8-12; pl. 8, fig. 22) as explained in the previous paper (Moore, 34). When these become mature, eross-walls are again laid down, but at this time they are formed by a thickening of the region where the partition is to develop (pl. 7, figs. 9-10). These may be simple or collar-like (pl. 8, figs. 22, 24). With their formation, the resulting cells become uninucleate (pl. 7, figs, 13, 16) and are now the arthro- spores. In addition to the reticulated network in the cell noted previously, there is a heavy granulation on the inner surface of the wall (pl. 7, figs. 16, 18; pl. 8, fig. 24) which seems to be noticed more often in the arthrospores. An important feature in the filaments is the presence of clear, non-granulated, thin-walled cells, apparently devoid of cytoplasm (pl. 7, fig. 19; pl. 8, figs. 20-22, 24, 26-27, 32, 48, 53). What this lack of protoplasm indieates cannot be explained un- less it is that the contents were used up in the nourishment of the adjoining cells. On maltose agar the cells are much enlarged, especially those submerged. Here the peculiar condition arises of many long hyphae devoid of protoplasm with clusters of short branches of arthrospores as in pl. 8, fig. 22. Also, there are series of empty cells and arthrospores (pl. 8, fig. 24). These intercalary arthrospores give rise to chains of small arthrospores while still attached to the filament. In addition to the regular development of the fungus, there are several structures which must be given consideration. The first of these is the chlamydospore. This particular organ which can generally be distinguished by its apparent large size and thick wall is found here (pl. 7, figs. 7, 17, 19; pl. 8, figs. 20-21, 23) in much the same condition as the arthrospores. It is a coenocytie structure which has a heavily granulated, retieu- lar network and may oceur terminally as a spherical body. [Vor. 22 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN They may be found as spherical, cylindrieal, intercalary struc- tures (pl. 8, figs. 27, 48, 53) or in chains as sclerotie, thick-walled eells (pl. 7, fig. 19; pl. 8, figs. 20-21). Chlamydospores germi- nate (pl. 7, figs. 7, 17) and give rise to mycelium. The so-called blastospores are seen as granulated, nucleated cells, the nuclei varying in number from one to three, fre- quently one. The conidia, considered by many to be possibly blastospores, have been found to have a heavily granulated protoplasm with no indieation of a nucleus. A nucleus if pres- ent would be masked probably by the accumulated, heavily stained material within the cell. All indications, however, point to the absence of a nucleus, which would therefore justify the retention of the term conidium for that cell, as has been pointed out by the author. Cellular contents.—Not all the constituents of the cell have been identified, but those that have will be considered here. 1t has been shown by many authors that fungi tend to store food in the cell, particularly in the older mycelium. These reserve products are usually in the form of glycogen, lipoids, oil glob- ules, metachromatic granules of Guilliermond or nuclear de- composition products, as volutin, nucleic acid substances, and probably other protein derivatives and carbohydrates. Since most of these substances have been demonstrated and discus- sed by the author for Endomyces, it will suffice to mention here merely their presence and quantity and the technique employed. Volutin.—V olutin or metachromatic material is very easily demonstrated with methylene blue or even iron-alum haema- toxylin, as substances within the cell, usually along the inner surface of the cell wall (pl. 7, figs. 16, 18) or along the reticu- lated network. A pinch of benzidine sulphate added to a water mount of living material reveals a number of granules (pl. 8, figs. 33-41), particularly the so-called **daneing bodies”” which are precipitated volutin in a state of Brownian movement. These take a blue coloration. According to the work of Ber- trand (cited in Guilliermond, Mangenot and Plantefol, ’33), benzidine produces with peroxidases a blue coloration. 1935] MOORE—CYTOLOGY OF GEOTRICHUM VERSIFORME BEL Glycogen.—With neutral red and iodine potassium iodide, glycogen may be easily demonstrated. Neutral red shows this material as large, pink to red drops (pl. 8, figs. 25-27). The older cells have larger and more deeply colored drops than do the younger cells. With saturated iodine potassium iodide (pl. 8, figs. 28-32) glycogen is seen as orange-brown, ir- regular bodies spread throughout the cell. As in the case of neutral red, there is a greater amount in the older than in the younger cells. Vacuoles.—The presence of vacuoles is easily demonstrated with methylene blue or haematoxylin, as surrounded by the reticulum (pl. 7, fig. 19). Iodine potassium iodide (pl. 8, figs. 28, 31) shows the vacuoles very clearly, while benzidine sul- phate (pl. 8, figs. 36-41), as used previously, shows them to best advantage, most of them containing the ““dancing bodies. ”” Neutral red also brings out the vacuoles. Chondriosomes.—Chondriosomes or mitochondria may be demonstrated with iodine potassium iodide. They are seen as light yellow, refractile droplets of various sizes. With benzi- dine sulphate they are supposed to take a light blue color and are distributed as droplets throughout the cell (pl. 8, figs. 33- 35), and where vacuoles are present surround the vacuolar membrane (pl. 8, figs. 36, 40). Guilliermond has demonstrated them in Oidium lactis with benzidine. These bodies are few in young cells and abundant in older cells. Fat, lipoidal substances.—In addition to the substances listed above, lipoidal substances, fats, and various other re- serve materials, as well as secretion and excretion materials, can be demonstrated easily. They are found in varying amounts in the cells. These have also been discussed previ- ously by the author. Several agents can be used very favorably to study these ma- terials, each showing some degree of difference. With 2 per cent osmic acid (pl. 8, figs. 42-48) these substances are reduced and take a black coloration. They are found as small globules or droplets in the young mycelium (pl. 8, fig. 47) and as larger, heavier masses usually in the center of the older cells. Much [Vor. 22 812 ANNALS OF THE MISSOURI BOTANICAL GARDEN the same picture is presented with 5 per cent platinie chloride solution (pl. 8, figs. 49-53). With iodine potassium iodide as applied for glycogen and chondriosomes, fats and lipoidal sub- stances are seen, with careful focusing, as very small, highly refractile and hyaline bodies. DISCUSSION AND CONCLUSIONS It is not intended in this paper to formulate any new theories as to cytological pictures, but merely to interpret the phenom- ena involved in the development of Geotrichum versiforme. To summarize briefly, the uninucleate arthrospore serves as the spore for new mycelia. It germinates, forming a thin- walled tube which becomes multinucleate and when well elon- gated develops eross-walls to form coenocytic cells. These cells mature and secondary partitions are formed by a thicken- ing and subsequent gelatinization of a partieular portion of the hypha. "This process therefore gives rise to cells which are the uninucleate arthrospores. The division of the nucleus, as has been pointed out previ- ously, is apparently direct or amitotie and may take place lon- gitudinally or transversely in the cell. In the former ease, cross-walls can be seen forming at the point of division, while in the latter the nuclei appear alternately on the sides of the hypha, due to the growth of the filament. Nuclear phenomena in Oidium lactis as described by Guilliermond (700) are re- vealed by simple division into two masses within an areola. Dangeard, Janssens and Leblanc, and a host of others cited by the present author considered this division as mitotie, with the phenomena masked. In Geotrichum, as observed here, the light region or areola surrounding the nucleus is lacking. Darkly staining central portions may be seen which represent the nucleoli, but definite karyokinesis with ehromosomal forma- tion cannot be distinguished. Chlamydospores are found as coenocytic, enlarged, sclerotic, thick-walled cells. So-called blastospores are present as nu- cleated structures, while granular, apparently non-nucleated conidia can be distinguished. 1935] MOORE—-CYTOLOGY OF GEOTRICHUM VERSIFORME 313 Volutin, glycogen vacuoles, chondriosomes, and additional storage or reserve materials, as well as fat and lipoidal sub- stances, can be demonstrated in varying amounts by means of several agents. ACKNOWLEDGMENTS The author acknowledges gratefully the courtesies extended him by Dr. Carroll W. Dodge, Professor of Botany in the Henry Shaw School of Botany of Washington University, and Dr. George T. Moore, Director of the Missouri Botanical Gar- den, St. Louis, Missouri. BIBLIOGRAPHY Guilliermond, A. ('00). Etude sur le developpement et la structure de l'Oidium lactis. Rev. Gén. Bot. 12: 465—479. 1900. G. Mangenot, et L. Plantefol (733). "Traité de cytologie végétale. 1195 pp. Librairie E. Le Francois. Paris, 1933. Jannin, L. (13). Les **Mycoderma.”? Leur role en pathologie. Thèse Fac. Méd., ey . 1913. Moore, M. (’33). A study of Endomyces capsulatus Rewbridge, Dodge and Ayers: A causative agent of fatal cerebrospinal meningitis. Ann. Mo. Bot. Gard. 20: 471-568. 1933 — — — — —, (734). A new Geotrichum from a bronchial and pulmonary infec- tion, uotum versiforme Moore, n. sp. Ibid. 21: 349—366. 1934. [Vor. 22, 1935] 314 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION Or PLATE PLATE 7 Geotrichum versiforme AU figures drawn as correctly as possible with the aid of a camera lucida. All grown on Sabouraud’s glucose agar. Figure 10 drawn at x 1440, all others drawn at x 2300; plate reduced approximately one-half. 1-4. Germinating arthrospores. Fig. 5. Cells formed from a germinating arthrospore. Fig. 6. Arthrospore showing type of nuclear division. Fig. 7. Germinating chlamydospore Fig. 8. Advanced stage of fig. 5, showing cell formation and multinucleate condition, 8. 9-10, Coenoeytie cells with the beginning of eross-wall formation. Ties 11-12. Branching hyphae showing young coenoeytie cells becoming uni- nucleate arthrospores. Fig. 13. Filament of cells with arthrospore formation. Fig. 14. Cells showing nuclear condition and division. Fig. 15. Branching condition. Fig. 16. Chain of arthrospores. Fig. 17. Germinating ehlamydospore. Fig. 18. Arthrospore showing granular appearance of inner wall. Fig. 19. Chain of cells, probably ehlamydospores. "C". rre : à ANN. Mo. Bor. GARD., VOL. 22, 1935 PLATE 7 D S4, Ea I DA (E a MOORE—GEOTRICHUM VERSIFORME [Vor. 22, 1935] 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 8 Geotrichum versiforme All figures drawn as accurately as possible with the aid of a camera lucida. Figures 25-27 drawn at x 1440, all others drawn at x 2300; plate reduced approxi- mately one-half. Figs. 20-21. Coenoeytie cells, probably chlamydospores, on Sabouraud's glucose Fig.22. Branching condition of mycelium submerged in Sabouraud's maltose agar. Fig.23. Large round chlamydospores on the same medium. Fig. 24. Mycelium on the same medium Figs. 25-27. Whole cells mounted in neutral red, showing volutin eontent. Figs. 28-32. Cells mounted in iodine potassium iodide, showing glycogen as the heavily stained material, lipoidal substances as small granular hyaline bodies, and probable ehondriosomes as small dark bodies. Figs. 33-41. Living cells mounted in distilled water to which was added a pinch of benzidine sulphate, showing vacuoles with dancing bodies, probably volutin. Fi . Living cells mounted in 2 per cent osmie acid, showing fat content. Figs. 49-53. Cells mounted in 5 per cent platinie chloride, showing fat content, ANN. Mo. Bor. GARD., VOL. 22, 1935 PLATE 8 EL O Y Ra K S/A a => = A gl W I me | a fe) MOORE—GEOTRICHUM VERSIFORME HEAD INFECTION CAUSED BY A NEW HEMISPORA: H. COREMIFORMIS! MORRIS MOORE Mycologist to The Barnard Free Skin and Cancer Hospital, St. Louis Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany of Washington University INTRODUCTION Hemisporosis, as a clinical entity, has been established since 1909, when Gougerot and Caraven (’09) cultured an organism from fragments of an infected tibia. It resembled and was at first diagnosed as a syphilitic periostitis, but was found to be an osteo-periostitis. Later cultural studies (Gougerot and Caraven, ’10) on various media proved that the organism was similar to Hemispora stellata, a species which Vuillemin had isolated from Aspergillus repens and had described in detail (Vuillemin, '06). Since then Castellani has isolated another species, Hemispora rugosa, from cases of bronchitis and tonsil- litis and from the vulva (Castellani, '25). He also described a third species, H. pararugosa Castellani, Douglas and Thomp- son, but stated that it was probably only a variety of H. rugosa. The disease was later confirmed by several writers, notably Auvray (709), who described a gummatous infection or infiltra- tion of the neck and face. De Beurmann, Clair, and Gougerot (709) reported a case which showed cold abscesses of the penis, and there have also been cases of nasal involvement, cheek in- fections, and subcutaneous ulcerating gummas on the back (Balzer and Belloir, '13). Porcelli (22) found an infection of the nose, the first case of hemisporosis from Italy. Other cases have been reported, but since the literature has been sum- marized (Grütz-Elberfeld in the Handbuch der Haut- und 1 Presented before the Mycological Society of America, December 27, 1934, at the winter meeting of the American Association for the Advancement of Science, held at Pittsburgh, Pennsylvania. Issued May 25, 1935. ANN. Mo. BoT. GARD., VOL. 22, 1935. (317) [VoL. 22 318 ANNALS OF THE MISSOURI BOTANICAL GARDEN Geschlechtskrankheiten, '28), a complete review here would be unnecessary. The organism described in this paper, morphologically, eul- turally, and physiologically, was isolated from an infeetion of the head by Peña Chavarria of Costa Rica. An abstract of the case, which is in press, in the Archiv für Schiffs- und Tropen- hygiene, Hamburg, was kindly sent the author by Chavarria. CASE REPORT ‚a farmer, 34 years of age, living at Turrialba (Atlantic slope), a region frorn which a patient with a similar infection had been treated six years previously. Patient entered the hospital on June 13, 1932, with an infection due to scratching bee bites with soiled hands. A small vesicle formed at the center of his forehead, which when seratehed exuded a clear liquid. He received medical treatment quite early (injeetions of Antimosan, an antimony preparation), but new lesions which followed the course of the former one appeared on his forehead, back of the neck, and on the upper extremity of the right ear. The patient showed no history of syphilis or tubereulosis, but of several of the tropical, parasitie diseases, including malaria. Wassermann negative. The patient is well-built, in a good state of health on a general physieal exami- nation, The skin infection, however, shows a large lesion on his forehead, 4 x 5 inches, with the surface and edges slightly raised and irregular. The color is brass = simulating a syphilitie lesion. The tissues are Vini with no pain to the touch. There is an occasional pruritus. Along the left eyebrow there are four sur which tend to evolve as did the other on the forehead. On the edge of the right ear there are several small papulo-verrueoid lesions, some of which show suppuration. There are also lesions on the angle of the right jaw. Scars on the “n region of the same side correspond (according to the patient’s state- nts) to the above lesions. Repeated examinations of material from the lesions an always yeast-like organisms. The patient was treated with iodine and tartar emetie intravenously and anti- septies locally, with healing resulting in two months. TECHNIQUE Cultural studies were made of the organism from a fairly wide variety of substrates. The morphology was studied in distilled water mounts, in Amann's lacto-phenol, and mounts in erystal violet in glycerine (1 per cent erystal violet to the de- sired intensity in glycerine). Details of the cellular and spore development were observed from hanging-drop and slide mounts in 2 per cent bacto-peptone, laetose broth (Difco), and beef extract broth (Difeo). 1935] MOORE—HEMISPORA COREMIFORMIS 319 DESCRIPTION The microbe studied was obtained on a Sabouraud's glucose agar slant, growing as a yeast-like organism and appearing culturally as a vermiculate growth. The parasite exists in the host as a yeast-like organism with budding cells (pl. 9, fig. 1) 4-6 » in diameter. These cells are thick-walled and appear double-contoured. When transferred to an artificial substrate they elongate to form selerotie cells, simulating germinating spores (pl. 9, fig. 1). The organism exists in this state for a variable length of time, depending on the medium in which it is grown, the temperature, and other growth factors. With repeated subculturing in most cases of yeast-like or- ganisms, it is possible to obtain a filamentous type of fungus. In this ease, the mierobe was subeultured twice, and showing no appreciable change morphologieally or culturally, was stored in the ice-box for approximately five months. It was then subcultured twice, after which it underwent several changes, physiologically and morphologically. This develop- ment may be attributed to the adaptation of the organism to a saprophytie form of life on the artificial medium. The reac- tion involved, in addition to the different substrate, a changed pH, nitrogen and carbohydrate source. On most media the cells elongate to form long filamentous hyphae (pl. 9, figs. 2, 10). The cells may be long and thin, or short and narrow (pl. 9, fig. 35), or they may be thick (pl. 9, fig. 36). On the other hand, the mycelium may consist of simple or branched chains of yeast-like cells simulating oidia or arthrospores as on Richards” (pl. 9, fig. 21) and Raulin's agar (pl. 9, fig. 22), or there may be combinations of both types (pl. 9, fig. 28; pl. 10, fig. 73). On wort agar, a medium which is particularly adapted to yeast cultivation, the organism de- velops, in addition to cylindrical cells, spherical or ovoid struc- tures with a heavy gelatinous wall or envelope, from which the cell either grows or slips out (pl. 9, fig. 19; pl. 10, figs. 51—52, 54-59, 61-62, 66, 68, 70). This type of structure is also evident on malt-extract agar (pl. 9, fig. 26) which is a modification of wort agar. [VoL. 22 320 ANNALS OF THE MISSOURI BOTANICAL GARDEN With the formation of hyphae, which may vary from 2 to 4 a in diameter, there are formed also interealary chlamydospores (pl. 9, fig. 9) either spherical, 6-10 » in diameter, or ovoid or elongate, 7-9 x 12-15 » or larger. Usually, however, there may be seen, in mounts of mycelium, large spherical cells (pl. 9, fig. 32) floating freely. On wort agar, large cells 6-17 » in diameter (pl. 10, fig. 51) are in abundance. Conidia (pl. 9, fig. 9), usually pyriform, at times spherical, 4-6 » in diameter, are found frequently and germinate (pl. 9, figs. 23-25) to form hyphae. Blastospores (pl. 9, fig. 10; pl. 10, fig. 73) are also found. The formation of the hyphae is further emphasized by the production of spherical, globose or sub-clavate to clavate cells at the apex of the filament (pl. 9, figs. 26-30, 33-34; pl. 10, figs. 37-88, 41-44, 49-50, 60, 63-64, 68-69, 72-73). These cells may also be ampulliform (pl. 10, figs. 45-46) and Vuillemin con- siders them to be protoconidia or hemispores which later di- vide to form several spore-like cells termed deuteroconidia. In the growth of the filament, eross-walls are laid down much as in the arthrospore-produeing organisms, which accounts par- tially for the systematic position of the fungus and also for its confusion with other genera. The hyphae may branch and give rise to several filaments which undergo the same process of laying down eross-walls by a growth inward of the cell wall. The formation of the primary partitions is followed by that of secondary walls which form the catenulate conidia or arthrosporous cells (pl. 10, fig. 31). These cells, which vary in number and size, are formed by an almost simultaneous eross-wall production. They are usually preceded by a thickening of the filament wall, giving the ap- pearance of a constriction, so-called by Vuillemin and Castel- lani (pl. 9, figs. 30, 34; pl. 10, figs. 44, 50). Where there are a number of adjacent branches, coremia are formed (pl. 10, fig. 70), with large heads of many of the arthrosporous spherical or ovoid cells. The presence of coremia is noted on most solid media, but not in liquid substrates. The cells of the filaments, as stated above, may be arthro- sporous, spherical, cylindrical, or ovoid. In addition, there are 1935] MOORE—HEMISPORA COREMIFORMIS 321 sections of the hypha which are thin-walled and have no ap- parent cellular material (pl. 9, figs. 14, 19-20, 26-30; pl. 10, figs. 38, 41-50, 70, 72-73). This makes the intercalary cells appear as isolated chlamydospores, comparable to akinetes in the algae as pointed out for Geotrichum in a previous paper (Moore, *34). When the partitions are finally laid down the cells mature rapidly, and the last step in their cycle consists of the forma- tion of echinulate spores. The spores develop on filaments or hyphal branches, as do conidiospores, and the carriers of these cells have been termed conidiophores. Such a terminology would place the organism in the Conidiophorales as has already been done. There is apparently no sexuality here. The con- fusion, however, is due to the fact that the resulting spores ap- pear to be formed in a manner which would take them from the above group, the Fungi Imperfecti, and place them in a better- known division of fungi, the Ascomycetes. This change is sub- ject to the further investigation of the organism. Vuillemin observed a thickening and granulation of the ex- ternal spore wall which makes the spore appear dark. Here, the dark echinulate spore develops from within the cell (pl. 10, fig. 65). When mounted in a lacto-phenol preparation or some aniline dye, the internal cytoplasm appears as a much-thick- ened and darker-staining material surrounded by a faintly staining cell wall. As the spore matures it becomes thicker, showing small striations which at first appear as deep corru- gations, and later developing fine prickles. In the meantime the cell wall degenerates, becomes irregular, loses its consist- eney and finally is devoid of life. At this time it 1s a disinte- grating brownish material (pl. 10, fig. 74). The fine prickles are now more apparent and finally develop into small spines. At times the hemispores may sporulate simultaneously (pl. 9, fig. 26), so that two spores may appear attached (pl. 10, fig. 66). However, the usual procedure consists of the dispersal and pro- duetion of these structures at the apex and proceeding down towards the base of the hypha. The type of spore development is highly suggestive of uni-spored asci and the hypha may then be ealled ascogenous. Further evidence will be derived from [VoL. 22 322 ANNALS OF THE MISSOURI BOTANICAL GARDEN eytological investigations now in progress. In coremia the de- veloping spores simulate roses in a bouquet, and the whole structure, superficially, appears like that found in Briosia of the Phaeostilbaceae of the Fungi Imperfecti. CULTURAL DESCRIPTIONS The organism presented a yeast-like appearance at first, but after repeated subculturing and storage in the ice-box, the morphology was changed. This suggested an adaptation to a saprophytie mode of life, and consequently it was decided to grow the mierobe in a number of different media. These sub- strates form a general list commonly employed in examination of fungi and involve a range of hydrogen-ion concentrations, different amounts of protein or protein decomposition produets as peptones, and also varying amounts of carbohydrate and nitrogen. The following media used for the description of the organism are arranged in the order of decreasing concentration of hydro- gen ions. All cultures were grown at approximately 25? C. Raulin’s Agar (pH 4.1).—Colony submerged in agar and seen only by allowing a light to pass through the medium. Diameter approximately 3.5 em. after 32 days. Chains of cells spherical to ovoid, 5-6 y in diameter; coremia with slightly elongated cells 4 x 6 y, and elongated cells 4-6 x 7-9 »; long narrow hyphal cells 3 x 15 »; large spherical cells 9 » in diameter; chains of cells somewhat cylindrieal, terminating in a spherieal or ovoidal cell approximately 7 » in diameter. Richards’ Agar (pH 4.3).—Colony submerged as above, showing the branched growth with a diameter of approxi- mately 3.5 em. after 32days. Cells in chains, spherical to ovoid, 6-8 „in diameter; cylindrical cells in series, 4 x 9 a, thick-walled or double-contoured; many young filaments tapering off from a series of yeast-like cells, to longer finer cells of a smaller diameter, 2-3 a; filaments of cells with the ultimate structure thiek-walled, enlarged, and showing a eross-wall to form 2 hemispherieal cells. Cezapek's Agar (pH 4.4).—Colony as above, diameter 4 cm. after 32 days. Chains of cells; round cells 4-8 u; elongated 1935] MOORE—HEMISPORA COREMIFORMIS 323 ovoid cells 3-4 x 7-9 », arthrosporous; series of subelavate cells with ultimate cell approximately 6 » in diameter. Wort Agar (Product of Digestive Ferments Co., pH 4.8).— Colony vermiculate, heaped up, light cinnamon in color, ap- proximately 2 em. in diameter after 32 days. Growth mucoid, thick and tenacious. Many large yeast-like cells 6-8 » in diam- eter, with a thick gelatinous or mucilaginous sheath enlarging the cell diameter to 8-15 »; coremia of bundles of chains of cells 34.5 » in diameter, 7-21 » in length. Gelatin-encased cells borne at tips of filaments in short chains, developing from hemispores or partitioned-off cells by a process simulating arthrospore formation. Blastospores approximately 6 » in diameter; large spherical thick-walled cells (intercalary) ap- proximately 10 » in diameter; ovoid cells 8-9 x 12-14 u; echinu- late spores approximately 5 » in diameter. Malt Extract Agar (pH 5.2).—Colony vermiculate at periph- ery and center with radiate creamy-buff to yellow ridges. Cul- ture heaped up at center, point of inoculation approximately 3 em. in diameter after 32 days. Chains of spherical cells ap- proximately 10 » in diameter. Mucoid secretion present, but not in such great abundance as above. Characters otherwise similar to those on wort agar. Sabouraud's Agar (pH 5.6).—Colony cerebriform at point of inoculation with a coremiform, vermiculate ring of growth a short distance from the center, merging into a flat growth. Color that of medium, creamy-buff to amber. Diameter ap- proximately 5 cm. after 29 days. Coremia of chains of cells 3-4 x 8-20 »; elongate blastospores 4 x 6 y, spherical 5 » in diam- eter; large clavate cells terminal on filaments; elongated uni- cellular filaments and sclerotic cells 3 » in diameter; many bud- ding spherical cells approximately 6 » in diameter; echinulate dark spores approximately 5 » in diameter. Sabouraud’s Broth (The above minus the agar ).—Organism in flakes and groups of cells at bottom of flask. Chains of cells as above; no coremia; spherical cells 4-15 » in diameter; arthrosporous cells 4 x 6 »; hyphae of long cells 4 x 30 »; echinu- late spores approximately 5 y in diameter. Corn-Meal Agar (Product of Digestive Ferments Co., pH [Vor. 22 324 ANNALS OF THE MISSOURI BOTANICAL GARDEN 6.0 ).—Growth similar to that on Richards’ agar except for a finely visible surface formation. Colony approximately 3.5 em. in diameter after 32 days, color light Isabella. Thick-walled spherical cells in chains, 8-10 » in diameter; arthrosporous cells 4-6 x 6-10 », many cells of varying dimensions; hemispo- rous clavate cells 7-10 x 15-18 a; hyphae 3-4 » in diameter. Potato-Dextrose Agar (pH 6.2) —Colony 6 em. in diameter after 32 days, with many radiate branched cerebriform stria- tions or grooves from a highly peaked center. Color ereamy- buff. Coremia present; hyphae 3-4 » in diameter; chains of spherical cells 5-6 » in diameter, and arthrosporous cells 4 x 6-8 »; large cells 12-17 » in diameter; clavate triseptate cells 12-15 x 15-18 a; hemisporous clavate cells 9 x 15 a; elongated cells 4-6 x 6-15 a, in chains; eitriform to subovoid cells 4—6 x 6-8 a; sclerotic cells numerous, as well as racquet or clavate cells in short chains. Lactose Broth (Product of Digestive Ferments Co., pH 6.8). —Maeroscopie appearance similar to that in Sabouraud's broth. Chains of spherical cells 5-6 » in diameter, occurring terminally on filaments. Cells catenulate, simulating those found in Oospora; hyphae approximately 3 & in diameter; spherical terminal cells 9 » in diameter; clavate hemisporous cells 7 x 12 »; conidium-like cells 6-9 » in diameter, somewhat pyriform to ovoid; interealary spherical cells 6-8 » in diameter. Lactose Agar (The above plus 2 per cent agar ).—Colony raised and convolute in center, with radiate striations to pe- riphery, approximately 5 em. in diameter after 32 days. Cells in chains as in broth; fine eoremia appearing hyaline to white on surface, consisting of spherical, subovoid and arthrosporous cells; hyphae 3 » in diameter; echinulate spores approximately 9 p in diameter, appearing black with lacto-phenol (cotton blue). Nutrient Agar (Product of Digestive Ferments Co., pH 7.0). — Colony flat except for a slightly raised center, approximately 4 em. in diameter after 32 days. Culture appears asteroid or stellate, color that of medium. Hyphae 4 y in diameter; ter- minal spherical cells approximately 6 u in diameter; terminal clavate hemisporous cells and intercalary spherical cells; large 1935] MOORE—HEMISPORA COREMIFORMIS 325 spherical cells 10-12 „in diameter ; cells in chains, 6-8 » in diam- eter. General characteristics similar to those on lactose agar. Glycerine Agar (Nutrient agar plus 6 per cent glycerine, pH 7.0).—Cultural appearance similar to that on potato-dex- trose agar. Very fine prickles on surface of colony, coremia. Colony approximately 4 cm. in diameter after 32 days. Color creamy-buff, with a hyaline sheen. Hyphae 3-4 » in diameter; many large multi-septate filaments 5-7 » in diameter; conid- ium-like cells approximately 5 » in diameter; clavate cells, di- and tri-septate, 9-12 » in diameter; large cells 9 » in diameter; small spherical cells 4-7 » in diameter; cells of filaments 4 x 12 »; arthrosporous cells 4 x 8-9 u. Many clumps of cells 6-7 y in diameter held by a gelatinous matrix; clavate hemisporous cells 9x 12-15 a. Serum Agar (Nutrient agar plus 1 per cent bacto-beef blood serum, pH 7.2).—Colony macroscopically simulates that on nutrient agar except for the greater number of radiations or ridges. Diameter approximately 3 em. after 32 days. Color creamy-buff to amber. Cells of filaments approximately 6 » in diameter; terminal cells 7-8 » in diameter; hyphae 3-4 » in diameter; ovoid cells 4 x 7-8 4; many small spherical cells ap- proximately 4 » in diameter. Cells in general seem to be in a simple yeast condition due perhaps to the serum in the medium. Endo's Agar (Product of Digestive Ferments Co., pH 7.5).— Culture macroscopically similar to that on glycerine except that here the mycelium has taken up the red dye of the medium. Diameter approximately 3.5 cm. after 32 days. Characters similar to those on Sabouraud's agar. Gelatine (Beef extract broth plus 10 per cent bacto gelatine). —Liquefaction starts on twelfth day at surface and proceeds downward. Litmus Milk.—Acid and curdling start on the third day and are complete on the seventeenth day. Carbohydrate Reactions.—Acid and no gas production with l-arabinose, l-xylose, glucose, galactose, d-mannose, levulose, lactose, maltose. No acid or gas, but an alkaline reaction with rhamnose, saccharose, raffinose, amygdalin, and salicin. [Vor. 22 326 ANNALS OF THE MISSOURI BOTANICAL GARDEN DISCUSSION The genus Hemispora was created by Vuillemin in 1906, with a single species stellata, so called because of the presence of star-like growths on Aspergillus repens. The surface of the organism was powdery and the whole growth was brown in color. On microscopic examination, the powder or dust was found to be a mass of spores. The mierobe was later found in a number of conditions existing as a saprophyte. As a para- site the fungus or a closely related organism has been isolated from a number of cases which resembled clinically sporotricho- sis, and the cultivation of the mierobe was required in order to arrive at a better diagnosis. On the basis of the type of spore formation, that is, the pro- duction of branches of hyphae which have at their apex a large cell termed a protoconidium or hemispore, the fungus was clas- sified with the Conidiophorales. The protoconidia give rise almost simultaneously to cells which approximate each other in size and are termed deuteroconidia. Several years after Vuillemin's el tion and identifica- tion of Hemispora stellata, Castellani in 1910 isolated a new species, H. rugosa, from cases of bronchitis and a case of tonsil- litis. This organism differs from the former in the type of cul- tural growth, which is abundant on glucose agar, erinkled, and at times cerebriform. The color varies from amber to brown. The main difference, according to Vuillemin, seems to be that it grows in gelatine and liquefies it slowly, while gelatine lique- faetion is negative with H. stellata. Milk is not changed gen- erally, but may be somewhat peptonized with a small coagulum. The sugar reactions are as follows: acid with saccharose, glu- cose, arabinose and levulose, and doubtful for maltose; no acid with laetose, dulcite, mannite, dextrin, raffinose, adonite, inulin, starch, salicin, galactose, and glycerine. Litmus milk is negative. A third species, H. pararugosa Castellani, Douglas and Thompson, mentioned by Castellani (725), differs from H. rugosa in being a rapid liquefier of gelatine. This organism may be only a variety of the former, or perhaps identical with 1935] MOORE—HEMISPORA COREMIFORMIS 327 that species. It is generally recognized, however, that both species are unlike H. stellata, and Ciferri and Redaelli ( 34), in a foot-note, make the following statement: ‘‘Hemispora rugosa Cast. and H. pararugosa Cast., Douglas and Thompson, are Trichosporon- ( Trichosporum-) like fungi, in no way com- parable to Hemispora stellata, in spite of the strange affirma- tion of Vuillemin (1931) that the differences pending [sie] on the liquefaction of gelatine.’’ These two authors in the same paper, which is a summary of a paper dealing with a comparative study of twenty-one strains of fungi referred to as H. stellata Vuill., Oospora d'Agatae Sacc., Torula sacchari Corda, etc., place the genus Hemispora in synonymy with the genus Sporendonema of Desmaziéres (1827). Although the type species of the latter genus should be S. casei (Desmaziéres, 1827), they create a new combination with the species Torula epizoa Corda, Sporendonema epizoum (Corda) Ciferri and Redaelli, and establish that as the type Species. In spite of the elaborate deseription and combination by these two authors, it seems that the interpretation of previ- ously existing genera is almost purely personal. On the other hand, it is difficult to compare favorably the generic characters based entirely on morphologie features with the structures ex- isting in Vuillemin's Hemispora. It is very easy to assume similarities, but genera cannot be based entirely on assump- tions especially if the diagnostic features are lacking. The author is satisfied that the genus Sporendonema shows insuf- ficient similarity in characteristics to replace Hemispora. Further, there is no apparent certainty that the fungus orig- inally described by Vuillemin was the one which these or other authors have been able to obtain for comparison. In addition, because of the type of spore production which suggests itself as ascogenous, the genus Hemispora should stand, with a posi- tion in the Ascomycetes. In this regard, the organism described in this paper simu- lates closely characters attributed to H. stellata. After care- fully comparing these characters with those of the monotypic [Vor. 22 328 ANNALS OF THE MISSOURI BOTANICAL GARDEN genus, Hemispora, it appears to the author that the fungus de- seribed in this paper has several differences which require for it a separate classification as a new species. Hemispora coremiformis Moore, n. sp. Growth in host of yeast-like cells 4-6 » in diameter. On arti- ficial media cultures show abundant mycelium of branching septate hyphae 2-4 » in diameter, terminating in clavate, spher- ical or ampulliform cells (hemispores) 7-15 x 12-18 a. Sexu- ality absent. Coremia formed on most solid media. Cultures stellate, vermiculate, coremiform, or cerebriform; color gray- ish-white, ereamy-buff, light einnamon, light Isabella. Cells of coremia 3-6 x 5-21 a; large spherical cells 6-17 » in diameter; conidia 4-6 » in diameter; blastospores 4-6 » (spherical), 4x6 y (pyriform); spherical intercalary chlamydospores 6-10 » in diameter, ovoid 8-9 x 12-14 p. Echinulate spores resulting from apical cells of mature filaments 3-6 » in diameter, in gen- eral 5. Acid and no gas with l-arabinose, l-xylose, glucose, galactose, d-mannose, levulose, lactose, maltose. No acid or gas with rhamnose, saccharose, raffinose, amygdalin, salicin. Gelatine liquefies slowly, starting on the twelfth day. Litmus milk becomes acid and curdles starting on the third day, com- plete on the seventeenth day. Hemispora iformis Moore, n. sp. Mycelium in culturis abundans sed in hospite cellulae sin- gulae diametro 4-6 » sunt. Hyphae ramulosae septataeque, diametro 2-4 y, cellulae terminales clavatae, sphericae vel ampulliformes (hemisporae) 7-15 x 12-18 a. Sexus deest. Coremia in mediis solidiis. Culturae stellatae, vermiculatae, coremiformes cerebriformesve; color albidus, subalutaceus, dilute cinnamomeus vel dilute isabellinus. Cellulae coremiae 3-6 x 21 »; conidia diametro 4-6 y, blastosporae pyriformes, 4 x 6 »; chlamydosporae, sphericae intercalarae diametro 6-10 », ovoideae 8-9 x 12-14 y. Sporae echinulatae ex cellulis apicali- bus hypharum maturarum diametro 3-6 » generatim diametro 5 a. Acidus in saecharo ; **l-arabinose, 1-xylose, glucose, galac- tose, d-mannose, levulose, lactose, maltose." Fermentatio nulla in saccharo ; **rhamnose, saccharose, raffinose, amygdalin, 1935] MOORE—HEMISPORA COREMIFORMIS 329 salicin.?? Gelatinum per duodecim diebus fluidificans. Lac concretus, acidum per tribus diebus absolute per septendecim diebus faciens. SUMMARY AND CONCLUSIONS 1. A brief historical review of hemisporosis is given, with an abstract of an infection of the head oceurring on a man in Costa Rica. 2. The organism exists as a yeast-like organism in the host and changes to a filamentous fungus on artificial media. 3. The mierobe is characterized by the formation of coremia on most solid media, not on liquid media. 4. The mycelium consists of arthrosporoid cells, elongated forms and short structures which occur either as simple or branched filaments. These hyphae usually terminate in spher- ical or clavate cells, the protoconidia, which later divide to form deuteroconidia and develop echinulate spores. 9. The organism is described on several media, culturally and microscopically. 6. Gelatine is liquefied starting on the twelfth day. 7. Litmus milk is acidified and eurdled starting the third day and completed on the seventeenth day. 8. Acid and no gas with 1-arabinose, 1-xylose, glucose, galac- tose, d-mannose, levulose, lactose, maltose. No acid or gas with rhamnose, saccharose, raffinose, amygdalin, and salicin. 9. As a result of the characteristics and properties of the organism, it is described as a new species of Hemispora, H. coremiformas. ACKNOWLEDGMENTS Thanks are due Dr. Carroll W. Dodge, Professor of Botany in the Henry Shaw School of Botany of Washington Univer- sity, and Dr. George T. Moore, Director of the Missouri Botan- ieal Garden, for the courtesies extended in the use of the labo- ratories, equipment, and library; and Dr. Pefia Chavarria of Costa Rica for the organism and case report. BIBLIOGRAPHY Auvray [M.] (709). A propos d'une nouvelle myeose observée chez l'homme (1’Hemispora stellata). Bull. Mém. Soe. Chirurg. 35: 686-695. 1909. [Vor. 22, 1935] 330 ANNALS OF THE MISSOURI BOTANICAL GARDEN Balzer, F., et Belloir. ('13). Un cas d'hémisporose. Soc. Franc. Dermatol. Syphil., Bull. 24: 129-132. 1913 Castellani, A., and A. J. Chalmers (719). Manual of tropical medicine. 2436 pp. illiam Wood & Co., New York. 3rded. 1919. a, (2). Torker observations on — Monilias and vaginal ——— Jour. Trop. . & Hyg. 28: 242-249. 1925. Ciferri, R., and P. Redaelli ( ^90. Sporendonema epizoum (Corda) Cif. et Red.; An entity including Hemispora stellata and Oospora d’ Agatae. Ibid. 37: 167— 170. 1934. De Beurmann, Claire, et Gougerot (’09). Une nouvelle mycose: L'hémisporose. Un *hémisporose de la verge. Mém. Soc. Med. Hop. Paris, Bull. 17: 917-919. Desmaziéres, J. B. H. J. (1827). Observation sur le Sporendonema casei, nouveau genere de Mucedinées. Ann. Sci. Nat. Bot. 11: 246-249, 1827. Gougerot et Caraven (709, ’10). Hémisporose humaine. Rev. Chirurg. 40: 896—920. 1909; Ibid. (nouvelle mycose) ; Ibid. 41: 66-89. 1910. Grütz-Elberfeld, O. ('28). Sporotrichosen und verwandte Krankheiten. Hand- buch der Haut- und Geschlechtskrankheiten 11: 724—824. (pp. 805-809.) Moore, M. (7/34). A new Geotriehum from a bronchial and pulmonary infection, Geotrichum versiforme Moore, n. sp. Ann. Mo. Bot. Gard. 21: 349-366. 1934. Porcelli, R. (22). Micosi Pr da , (Vuillemin). Prim caso in Italia [Con una tavola]. Giorn. Ital. Mal. Ven. Pelle 63: 698-709. 1998. Redaelli, P., and R. Ciferri (7/34). Studio comparativa di ventum ceppi di € donema epizoum (Corda) nobis (Hemispora stellata Vuillemin; Oospor d'Agatae Saecardo; eec.). Atti dell’ Ist. Bot. dell’ Univ. Pavia 5: 145- 198. Votimi, P. (’06). Un nouveau genre de Mucedínées: Hemispora stellata. Soc. Myc., Bull. 22: 125-129. 1906 (731). Les champignons parasites et les mycoses de l'homme. En- cyclopédie myeologique. 291 pp. Lechevalier. Paris. 1931. [Vor. 22, 1935] 332 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 9 Hemispora coremiformis All figures drawn with the aid of a camera lucida at a magnification of x 1440 and reduced to approximately x 750. Fig. 1. Group of yeast-like cells, first subeulture on Sabouraud's agar. Fig.2. Filament in Sabouraud's broth. Fig.3. Germinating spore on Raulin's agar Fig.4. Filament with ehlamydospore on Raulin's agar. Fig.5. Filament with chlamydospore on Richards” agar. Figs. 6-7. Type of cells on Raulin's agar. Fig. 8. Oidia-like cells in Sabouraud’s broth. Fig. 9. Mycelium with chlamydospore and conidium on lactose agar. Fig. 10. Hypha showing m on Sabouraud’s agar Fig. 11. Oidia-like cells on malt-extract agar Figs. 12-13. Types of cells on Baulin^ s agar. Fig. 14. Type of mycelium in lactose ehe Figs. 15-16. Hemispores on corn-m ag Fig. 17. Cells mounted in lacto- Edu. as ^ii 16 from EM agar. Fig. 18. Thick-walled spherical cell in Sabouraud 's bro Fig. 19. Filament showing gelatinous secreted cells on pm agar. Fig. 20. Mycelium on potato-dextrose agar. Fig. 21. Mycelium on Richards’ Fig. 22. Smaller cells as seen on Raulin’ s agar. Figs. 23-25. Germinating conidia on Sabouraud’s agar. Fig. 26. Cells on malt-extract agar, showing a terminal hemispore heavily encased. Fig. 27. Oidia-like cells separated by clear cells in lactose broth. Fig. 28. Type of m — on Czapek's agar Fig. 29. Mycelium as fig. 2 Fig. 30. Terminal cell EUM of a filament on serum agar. Fig. 31. Deuteroconidia formed on potato-dextrose agar. Fig. 32. Group of spherieal cells mounted in lacto-phenol from Sabouraud's agar. Fig. 33. Terminal spherical cell on lactose agar. Fig. 34. Terminal deuteroconidial condition on lactose agar. Fig. 35. Hypha on glycerine agar. Fig. 36. Hypha in Sabouraud's broth. PLATE 9 0 DB (N VC MOORE—HEMISPORA COREMIFORM IS ANN. Mo. Bor. GARD., VOL. 22, 1935 i i 334 [VoL, 22, 1935] ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 10 Hemispora coremiformis All figures drawn with the ai dof a camera lucida at a magnification of x 1440 and reduced to approximately x 750, except fig. 70, which was drawn at x 960 and reduced to x 510. Fig. 47. Fig. 48. Fig. 49. . Cross-wall formation in s in laetose broth. . Mycelium on serum agar . Cells on Richards? agar. b renes ii on malt-extraet agar. Richards? agar. 42. Terminal Are cell on potato-dextrose agar. . Terminal cells on Raulin’s agar . Young filament showing tapering of cells from a terminal spherical cell. 6. Ampulliform cells on potato- ee — Deuteroconidial cells in Sabouraud’s bro Spherical cell in Sabouraud’s broth d in lacto-phenol. Terminal spherieal cell on malt-extract a ilament showing terminal eell eondition on , Baulin 's agar. ig. Figs. 51-52, 54—59, 61-62, 68, 70. Cells on wort agar showing gelatinous seeretion. Fig. 70 shows a eoremiu Fig. A Hemisporous pom on eorn-meal agar. . Terminal cells on Richards” agar. Fig. Figs. "Ps Mycelium showing terminal hemisporous condition on Richards? agar. Fig. 65. Filament on malt-extraet agar showing formation of echinulate spores from y end. Fi . Adjoining echinulate spores formed from a hemispore on wort. Mycelium on Czapek ’s r. . Terminal cell condition on malt-extract agar. . Spherical cell on Richards? agar 2. Filaments from a coremium on shale -dextrose agar. . Mycelium on Czapek’s Fig. 74. Series showing p Ae a of echinulate spore. ANN. Mo. Bor. GARD., VOL. 22, 1935 TU a £o do Ws ae WOE SCH Un Se a SON (x = = N amp P aus ee A T; a" MOORE—HEMISPORA COREMIFORMIS A MORPHOLOGICAL AND PHYSIOLOGICAL STUDY OF TWO SPECIES OF POSADASIA P. caPsULATA (DARLING) Moore AND P. PYRIFORMIS MOORE MORRIS MOORE Mycologist to The Barnard Free Skin and Cancer Hospital, St. Louis Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany of Washington University INTRODUCTION The etiological agent of histoplasmosis has for a number of years been incorrectly determined. The disease, made well- known in a series of papers by the original describer of the in- fection, Darling (706, *08, 09), was attributed to a protozoon. Later investigators considered it a Cryptococcus (da Rocha- Lima, *12, "13; Castellani and Chalmers, ’19; Vuillemin, ’31) or an Oidium. This confusion may have been due to the inability of the earlier workers to grow the organism, all systematic classification being based on smears or tissue sections. The present author, upon receipt of an organism from a proven case of histoplasmosis, sought to determine the correct position of the fungus. The disease itself is a serious and apparently fatal infectious condition which is present in America and resembles kala-azar of India and Tropical America. It is characterized clinically by emaciation, severe anemia, with a marked leucopenia, splen- omegaly, enlargement of the liver, and irregular pyrexia. Pathologically, the organism is found to invade the endothelial cells in the smaller lymph and blood vessels and capillaries. The affected organs become necrosed and the liver develops cirrhosis. The lungs and small and large intestines are studded with pseudotubercles, giving the appearance of miliary tuber- culosis. The peribronchial lymph nodes usually are enlarged, showing some tubercles most of which become ulcerated. Issued May 25, 1935. ANN. Mo. Bor. GARD., Vor. 22, 1935. (335) [Vor. 22 336 ANNALS OF THE MISSOURI BOTANICAL GARDEN The first case that came under Darling's observation was that of a negro, a native of Martinique, 27 years of age, who was mildly delirious and incoherent in his speech. The patient showed an enlarged spleen. The autopsy picture was typical of what has been pointed out previously. The second case was also that of a Martiniquan negro. The third case was a Chinese shopkeeper, who had lived on or near the Isthmus of Panama for 15 years. The organism of Posadasia capsulata, studied in this paper, was received from Charles Thom, Principal Mycologist of the United States Department of Agriculture. It had been sent to him by DeMonbreun of Vanderbilt University, who isolated it from a case of histoplasmosis, deseribed by Dodd and Tomp- kins (734). DeMonbreun (734) apparently proved the patho- genicity of the organism by carrying it through experimental animals and recovering the pathogen. Dodd and Tompkins (733) and DeMonbreun (733) reported their findings at the meeting of the Society for Tropical Medicine at Richmond, Virginia, November 15, 1933. The second organism, P. pyriformis, was received from G. H. Hansmann, formerly of Iowa City and now of Georgetown Uni- versity School of Medicine. Hansmann and Schenken (733) had reported a case before the American Association of Pathol- ogists and Baeteriologists at their Washington meeting on May9,1933. The patient was a white male, 43 years old at time of death. He had had a refractory skin ailment for the last 16 years of his life. This was a scaly inflammation underneath which the skin was thick and red, becoming more intense during 1929. In June of 1932 the skin showed hard nodules, and in July the patient developed a high fever, signs of pleurisy, and died on August 7. The microbe was cultured from dermal ele- vated spots and an enlarged lymph node. The fungus was con- sidered an Oidium. TECHNIQUE Hanging-drop, water mounts, and also simple water mounts on a slide revealed the general outline of the organism clearly. Mounts made with Amann’s lacto-phenol also gave favorable 1935] MOORE—POSADASIA CAPSULATA AND P. PYRIFORMIS 337 results, but those in erystal violet in glycerine were only fair. The mycelium of these fungi is such that only the large cells and the asci show any indication of cellular construction. The hyphae are fairly small, and the details are obscured by the wall, which is quite thick here. In order to trace the development of a single ascus, beef-ex- tract gelatine mounts were made with a suspension of myce- lium, gelatine being used in place of agar because of its greater transparency. Many of the cell interpretations are based on a study of sectioned materials. DESCRIPTION The life cycles of yeast-like fungi are usually complicated, because different habitats change the type of growth. In many cases, a different mycelium is linked with the loss of infectivity, or the change from a parasitic to a saprophytic mode of life. In any case, the organisms involved here existed as simple yeast-like cells, spherical or ovoid, varying from 1 to 4 in long axis, usually 3 » According to Darling, these cells are sur- rounded by a clear refractile and non-staining rim which equals in thickness about one-sixth the total diameter of the cell. The structure of these cells is found to be non-homogeneous, having a granular protoplasm which may be vacuolated. These cells multiply by elongation and fission according to da Rocha-Lima (712, 713), which led him to consider these fungi as members of the genus Cryptococcus. When transferred to an artificial substrate, the organism changes to a mycelial growth with aerial and submerged hyphae. When kept in blood and serum cultures at 37° C., how- ever, and transferred at short intervals the yeast-like form persists or predominates, according to observations by DeMon- breun (734) and also by Thom. All studies made here are from cultures which had already formed mycelium, having no evi- dence of yeast-like cells. The filamentation consists of multi-celled hyphae 1-5 » in diameter, with various types of morphology (pls. 11-14). There finally results a tubercled ascus containing a number of small spores. The yeast-like cells thus undergo a state of [Vor. 22 338 ANNALS OF THE MISSOURI BOTANICAL GARDEN elongation which gives them a sclerotie appearance, to form filaments. To deseribe the later development of the fungus on an arti- ficial substrate, it seems best to start with the ascospore which supposedly is the germ of the colony. "These spores (pl. 11, fig. 1; pl. 13, fig. 72) are small, and when set free from the ascus germinate to form a mycelium such as is shown in the illustra- tions. The hyphae are multi-celled and, from indications of eytological work in progress, coenocytic. These cells may be elongated and narrow (pl. 11, fig. 2) or short and thick, usually depending on the type of medium. "The filaments may develop peculiar swellings (pl. 11, figs. 32, 38, 42; pl. 12, fig. 55; pl. 13, figs. 75, 82, 90), or they may give rise to the well-known raequet mycelium (pl. 11, figs. 4, 8, 16-17; pl. 13, figs. 77, 97). The hyphae may further be distorted and present peculiar types of development (pl. 11, figs. 40, 47 ; pl. 13, fig. 101). In addition, there are a number of types of cells which play an important róle both in propagation and conservation of the organism. These organs include chlamydospores, conidia, and asci. The chlamydospores vary in size and proportions in Posadasia capsulata and P. pyriformis. They may be inter- calary, 3-10 » in diameter, and may occur singly (pl. 11, fig. 23) or in groups or chains (pl. 13, fig. 96) and may also be found laterally (pl. 11, fig. 30; pl. 12, fig. 64; pl. 14, fig. 109), sessile or pedieellate. They may also occur terminally as enlarged forms, 3-10 x 6-20 y. On serum agar in the moist region of the colony are series of cells (pl. 13, fig. 95) which simulate chlamy- dospores but are actually yeast-like. Conidia are cells of great importance in propagation. They occur laterally, are sessile or pedicellate, spherical or pyriform, 2-8 » in diameter (pl. 11, figs. 23, 28; pl. 12, figs. 60, 64). They are present on most media, especially on substrates with a low pH. On serum agar in P. capsulata they are found at just above or just under the surface, while in P. pyriformis they are mostly above the surface. Conidia break off easily from the hyphae and germinate (pl. 11, fig. 1; pl. 13, fig. 72) to give rise to a new colony serving much the same purpose as do the ascospores. 1935] MOORE—POSADASIA CAPSULATA AND P. PYRIFORMIS 339 In all the observations of the mierobes presented here, no indication of sexuality has made itself evident. Instead, the non-sexual ascus develops terminally, laterally, or as an inter- calary organ. Asci form first as globose or clavate cells on short or long pedicels or on one- to several-celled filaments. These cells enlarge and may become 5-18 » in diameter or in long axis. The pedicels may be perpendicular to the hyphae (pl. 14, figs. 106-107). The presumptive asci have thick walls (pl. 11, figs. 35-36; pl. 12, fig. 53) and when placed in a water mount show numerous oil globules, representing a large food- storage supply. In P. capsulata these cells are spherical or be- come so to form the spherical ascus typical of the species, while in P. pyriformis they may be spherical or pyriform. When young the presumptive asci are smooth-surfaced and thick- walled. Asthey grow older those present at or on the surface of the medium become somewhat pitted, then spinose (pl. 12, figs. 56, 65; pl. 14, figs. 113-115), while those below the surface or submerged in the agar remain smooth. The spines develop into variously shaped tubercles in the adult ascus. In the case of P. capsulata, on some of the substrates there are more asci at or on the surface than within the medium, whereas in P. pyri- formis these cells form a heavy and deep layer on the surface with smaller smooth forms just below the surface. The tubercles of the asci, mentioned above, vary in number and size and may become long finger-like projections simulat- ing germ-tubes (pl. 12, fig. 66). These may also be spherical emergences (pl. 11, fig. 51; pl. 12, figs. 58, 67 ; pl. 14, fig. 122) or short blunt outgrowths (pl. 12, figs. 62-63, 68-70), or they may be situated almost diametrically opposite as large blunt mem- branous growths (pl. 12, figs. 57, 61). In most cases, however, there are combinations of nearly all types, particularly long and short tubercles. These are comparatively smaller on the pyriform asci in P. pyriformis (pl. 14, figs. 118, 125, 130). The so-called germ-tubes may vary from approximately .5 to 7 » in the adult aseus. The asci themselves, exclusive of the tu- bercles, vary in size in both species. The spherical forms vary from 5 to 25 » in diameter, while the pyriform asci, present only in P. pyriformis, are 6-12 x 12-26 », usually 10 x 22 u. [Vor. 22 340 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cytologieal investigations show that the single aseus con- tains a number of spherical spores which are set free by a rup- ture of the aseus wall to germinate and commence another eycle. When in a nutrient condition the aseus may germinate with few to several germ-tubes which develop into a mycelium (pl. 12, fig. 71). Apparently, the tubereles play no part in re- produetion or propagation, for a study of their strueture shows them to be clear and void of any protoplasm. CULTURAL DESCRIPTIONS The cultures obtained were transferred to Sabouraud's glu- cose agar. Since, as previously emphasized, a study of an organism should not be limited to a single substrate, a number of standard media were inoculated and grown at room tempera- ture, approximately 22? C. It was desirable to make a compar- ative study of both organisms, and like media were therefore seeded with the two species. The following artificial substrates are arranged in the order of their deereasing hydrogen-ion concentration. POSADASIA CAPSULATA Raulin’s Agar (pH 4.1). —Growth poor, diameter approxi- mately 2 cm. after 43 days. Colony white and cottony. Myce- lium variable in size and shape; hyphae 2-3 » in diameter, with intercalary chlamydospores approximately 10 » in diameter; racquet mycelium; terminal hypnospores 9 x 18 a; large round cells at end of a thin filament approximately 10 » in diameter; conidia 3-5 » in diameter; tuberculate asci lacking. Richards’ Agar (pH 4.3). —Colony approximately 2.5 em. in diameter after 43 days. Point of inoculation somewhat heaped up, white and cottony. Mycelium mostly submerged in the agar. Culture consists chiefly of long branching hyphae 2-3 a in diameter; many small round cells approximately 3 » in diam- eter, large round cells terminal on long thin filaments, 5-10 » in diameter; conidia ovoid or pyriform, 5 » in diameter; inter- calary ehlamydospores 5-10 » in diameter, terminal hypno- spores as above; racquet mycelium present; asci with tubercles not apparent. 1935] MOORE—POSADASIA CAPSULATA AND P. PYRIFORMIS 341 Ceapek's Agar (pH 4.4).—Colony approximately 3 em. in diameter after43 days. Growth loose and cottony, color white. Mycelium similar to that found on the above two media. Malt Extract Agar (Product of Digestive Ferments Co., pH 4.6).—Colony approximately 4.2 cm. in diameter after 43 days. Development of ridges from point of inoculation to woolly periphery. Color light Isabella. Abundant mycelium of hyphae 2-3 y in diameter; racquet mycelium; conidia 3-5 » in diameter, spherieal on short pedicels, or pyriform; inter- ealary ehlamydospores 5-8 » in diameter, singly or in chains; terminal clavate and elongate cells varying in size and shape with the age of the organism; many small round cells approxi- mately 3 » in diameter, probably conidia; asci numerous, tuber- culate, 15-20 »; round cells on long thin filaments, approxi- mately 1.5 „in diameter, probably young asci. Wort Agar (Product of Digestive Ferments Co., pH 4.8).— Colony approximately 3 em. in diameter after 43 days. Growth thick and cottony. Color light Isabella. Racquet mycelium abundant, with hyphae approximately 3 y in diameter; inter- calary chlamydospores 5-8 » in diameter; terminal spherical cells 5-10 » in diameter; conidia numerous, 5 y in diameter; many asci 10-20 y with tubercles or elub-like projections up to approximately 7 » in length. Sabouraud’s Agar (pH 5.6).—Colony approximately 4.7 cm. in diameter after 43 days. Growth cottony, showing clumps similar to pleomorphie changes found in cultures of derma- tophytes, color light Isabella. Hyphae 2-3 y in diameter; conidia 3-5 » in diameter, mostly spherical; intercalary chlamy- dospores spherical, 5-7 » in diameter; racquet mycelium pres- ent; asci numerous, 15-18 y in diameter, with many tubercles projecting from the surface of the ascus. Sabouraud’s Broth (The above minus the agar, pH 5.6).— Growth of flakes or clumps, chiefly at bottom of flask. Surface shows a folded growth, cottony, with upright hyphae on sur- face macroscopically similar to columellae. Mycelium on sur- face white to light Isabella in color, similar to that found on agar. Broth becoming dark with growth of organism. Sub- merged growth of long branching hyphae 1.5-2 » in diameter. [Vor. 22 342 ANNALS OF THE MISSOURI BOTANICAL GARDEN Clavate cells terminal on filaments, varying in proportion and size. Corn-Meal Agar (Product of Digestive Ferments Co., pH 6.0 ).—Colony cottony, slightly heavy at point of inoculation, white in color, with a diameter of approximately 1.5 em. after 43 days. Hyphae 2-3 » in diameter, showing many swellings; racquet mycelium very much enlarged; conidia thick-walled, approximately 5 » in diameter; intercalary chlamydospores ap- proximately 8 » in diameter; terminal hypnospores also pres- ent; terminal clavate cells approximately 10 » in diameter; asci 10-15 » in diameter. Potato-Dextrose Agar (pH 6.2).—Colony very much heaped- up in center, cottony, white to light Isabella in color, 3 cm. in diameter after 43 days. Cells larger than on the above media. Hyphae 3-4 » in diameter, some larger; racquet mycelium very characteristic, swollen portions much enlarged ; conidia 5-7 a in diameter; intercalary chlamydospores 8-10 » in diameter, spherical lateral forms on 2- or 3-celled pedicels, 6 » in diam- eter, many occurring singly as well as in chains; clavate sessile cells; asci larger than others, up to 25 » in diameter. Lactose Broth (Product of Digestive Ferments Co., pH 6.8). —Flakes of cottony growth submerged in the medium with surface growth. Mycelium of long branching hyphae approxi- mately 2 » in diameter; very few swollen cells or racquet forma- tion; very few conidia ; terminal clavate cells present; tubercu- late asci absent in submerged mycelium. Aerial growth simi- lar to that found on an agar substrate of the same constituents. Lactose Agar (The above plus 2 per cent agar ).—Colony cot- tony, white to light Isabella in color, showing radiating ridges from the inoculum, attaining a diameter of approximately 3 em. after 43 days. Hyphae small, many 1-1.5 y in diameter, some 2-3 p; conidia numerous, 3-5 » in diameter; racquet mycelium present; intercalary chlamydospores 5-8 » in diameter; asci approximately 15 » in diameter. Nutrient Agar (Product of Digestive Ferments Co., pH 7.0). —Growth fair, colony sparsely cottony, somewhat flat, with a diameter of approximately 2.8 em. after 43 days. Color light Isabella. Hyphae 2-3 » in diameter; conidia sessile and pedi- 1935] MOORE—POSADASIA CAPSULATA AND P. PYRIFORMIS 343 cellate, approximately 5 » in diameter; racquet mycelium pres- ent; clavate terminal cells and interealary chlamydospores as on lactose agar; asci as large as 22 „in diameter with tubercles. Glycerine Agar ( Beef-extract agar plus 6 per cent glycerine, pH 7.2).—Thick cottony growth with an elevated inoculum ap- proximately 2.5 cm. in diameter after 43 days. Color light Isabella. Hyphae 2-3 y in diameter; sclerotic cells present; many conidia, spherical and pyriform, 3-5 » in diameter; ter- minal clavate cells, spherical cells on long thin filaments, 5-10 y in diameter; chlamydospores 5-8 „ in diameter; asci approxi- mately 15 » in diameter; many hyphal swellings. Serum Agar (Bacto-beef blood serum, product of Digestive Ferments Co., plus 2 per cent agar, pH 7.3).—Colony approxi- mately 2.5 cm. in diameter after 43 days, with deep radiating ridges from a somewhat white cottony point of inoculation to a moist and flat periphery. Hyphae tend to be thick and heavier and appear coremioid. Mycelium generally thicker, with the various cells present ; conidia spherical and pyriform, sessile or pedicellate, 5 » in diameter; chlamydospores approximately 8 a in diameter ; many large round cells in chains; asci up to 15 » in diameter, plus the tubercles. Endo’s Agar (Product of Digestive Ferments Co., pH 7.5).— Culture assumes the pink color from the medium, cottony in appearance, with a diameter of approximately 3 cm. after 43 days. Microscopic appearance similar to that on nutrient agar. POSADASIA PYRIFORMIS Raulin’s Agar (pH 4.1).—Colony approximately 2 cm. after 43 days. Culture white and cottony at inoculum, submerged in the medium. Mycelium shows raequet formation with swel- lings; hyphae 2-3 y in diameter; conidia sessile or pedicellate, spherical or pyriform, 5-7 » in diameter; intercalary chlamydo- spores spherical or slightly elongated, 3-10 » in diameter; clavate or spherical cells on pedicels, 6-10 » in diameter; asci below the agar without tubercles, above the surface multi-tu- bereulate, spherical 6-18 y, pyriform 8-10 x 18-26 y, in general 9 x 24 u. [Vor. 22 344 ANNALS OF THE MISSOURI BOTANICAL GARDEN Richards” Agar (pH 4.3).—Colony approximately 2 em. in diameter after 43 days. Culture cottony, white, and heaped-up at point of inoculation. Hyphae 3-4 y in diameter; conidia ap- proximately 5 y in diameter; racquet mycelium present; inter- calary chlamydospores 5-8 y in diameter; clavate cells on long filaments; asci terminal on long several-celled pedicels, or lateral on short pedicels, spherical approximately 12 y in diam- eter, pyriform 8-10 x 16-24 u. Czapek’s Agar (pH 4.4).— (Growth loose and cottony, approx- imately 4.5 em. in diameter after 43 days. Color white. Gen- eral characters similar to those on the above media. Hyphae 2-3 p in diameter; intercalary chlamydospores 5-7 a in diameter. Malt Extract Agar (Product of Digestive Ferments Co., pH 4.6 ).—Colony approximately 3 cm. in diameter after 43 days. Culture eottony and somewhat flat with a heaped-up center and showing concentrie circles of growth, Isabella color. Racquet mycelium in abundanee with numerous swellings; hyphae 2-3 » in diameter, some larger; conidia sessile or pedi- cellate, spherical or pyriform, 5-8 » in diameter; intercalary chlamydospores 3-8 » in diameter; terminal clavate cells pres- ent; asci relatively few in number, spherical predominating, approximately 12 » in diameter. Wort Agar (Product of Digestive Ferments Co., pH 4.8).— Growth thick and cottony, appearing cerebriform and con- volute at point of inoculation. Diameter approximately 2 cm. after 43 days” growth. Color light Isabella. Racquet myce- lium abundant; hyphae 24 y in diameter; intercalary chlamy- dospores 3-8 » in diameter; conidia as on the above medium; mycelium somewhat resembling chains of oidia, with the cells 6-8 x 7-12 a; spherical asci predominating, approximately 12 a in diameter. Sabouraud’s Agar (pH 5.6).—Growth thick and cottony, ap- proximately 6 cm. in diameter after 43 days. Colony Isabella- colored in center, approximately 1 em. high, with a concentric ridge surrounded by a lighter woolly periphery. Hyphae 2-4 » in diameter; racquet mycelium present; intercalary 1935] MOORE—-POSADASIA CAPSULATA AND P. PYRIFORMIS 345 chlamydospores 3-5 » in diameter; conidia approximately 5 y in diameter, both spherical and pyriform; asci numerous, on surface of agar, spherical 6-22 » in diameter, pyriform 6-10 x 12-24 u. Sabouraud’s Broth (The above minus the agar ).—Broth has turned dark with increased growth of flakes of mycelium. Sur- face shows a thick growth of Isabella-colored mycelium similar to that on the agar. Submerged mycelium of long hyphae ap- proximately 2 » in diameter with some variations; few swell- ings or racquet formations; clavate cells reduced in size. Corn-Meal Agar (Product of Digestive Ferments Co., pH 6.0).—Growth poor, colony approximately 1.5 em. in diameter after 43 days. Culture cottony with an irregular periphery, Isabella-colored. Cells and hyphae thicker, with the filaments 2-5 » in diameter; conidia spherical or pyriform, approxi- mately 5 » in diameter; racquet mycelium present; spherical asci predominating, 6-22 » in diameter; many thick-walled cells. Potato-Dextrose Agar (pH 6.2).—Colony 5 em. in diameter after 43 days. Culture cottony and somewhat flat with con- centric rings of growth, color Isabella. Hyphae 2-4 » in diam- eter; racquet mycelium and hyphal swellings; intercalary chlamydospores 5-7 » in diameter; conidia 3-8 » in diameter, both pyriform and spherical; many long filaments, approxi- mately 100 », bearing a clavate cell terminally, the presumptive ascus ; asci both spherical and pyriform, 6-22 » in diameter. Lactose Broth (Product of Digestive Ferments Co., pH 6.8). —Growth of submerged flakes of mycelium, with some Isabella- colored lumpy cottony colonies on the surface. Mycelium on surface similar to that on the agar; submerged mycelium of long hyphae approximately 2 « in diameter; few hyphal swell- ings or racquet mycelium; few conidia or intercalary chlamy- dospores ; many spherical and clavate terminal cells; cells much reduced in size. Lactose Agar (The above plus 2 per cent agar).—Colony 3 em. in diameter after43 days. Culture cottony and somewhat flat except for a woolly periphery, color Isabella. Hyphae 2-3 a [Vor. 22 346 ANNALS OF THE MISSOURI BOTANICAL GARDEN in diameter; conidia 3-5 » in diameter, both spherical and pyri- form; intercalary ehlamydospores 3-7 y in diameter; racquet mycelium and hyphal swelling present; spherical asci 6-20 a in diameter, pyriform 6-12 x 10-22 u. Nutrient Agar (Product of Digestive Ferments Co., pH 7.0). — Growth similar to that on lactose agar. Glycerine Agar (Beef extract agar plus 6 per cent glycerine, pH 7.2) —Colony approximately 4 em. in diameter after 43 days. Culture cottony with center showing convolutions and a splitting of the agar. Color Isabella. Cells short and thick; racquet mycelium abundant with hyphal swellings; intercalary chlamydospores 5-8 » in diameter; conidia spherical and pyri- form, approximately 5 » in diameter; spherical cells 8-9 « in diameter present in series in a hypha; spherical asci up to 22 y in diameter, pyriform 6-10 x 10-24 y, with well-defined tu- bereles mostly spherical or blunt. Serum Agar (Bacto-beef blood serum, product of Digestive Ferments Co., plus 2 per cent agar, pH 7.3).—Colony approxi- mately 2.5 em. in diameter after 43 days. Culture cottony, light Isabella, with a flat moist periphery suggestive of a yeast- like condition. Hyphae 2-3 » in diameter; racquet mycelium abundant; conidia sessile or pedicellate, spherical or pyriform, approximately 5 » in diameter; terminal subelavate to clavate cells; spherical asci 6-15 » in diameter, and pyriform of vary- ing proportions; interealary chlamydospores 5-7 y in diameter. Endo's Agar ( Product of Digestive Ferments Co., pH 7.5).— Colony approximately 3 cm. in diameter after 43 days. Cul- ture cottony with the central portion red and a pink coloration spread throughout due to the absorption of the dye in the medium by the mycelium. Characteristics similar to those found on nutrient agar. Gelatine.—No liquefaetion by either species, or if present extremely slow. Litmus Milk.—No acid or curdling, but an alkaline reaction which is interpreted as negative. Carbohydrate Reactions.—No fermentation of any sugar used. No acid or gas production. 3 ü 1935] MOORE—POSADASIA CAPSULATA AND P. PYRIFORMIS 347 Discussion The causative agent of histoplasmosis, as has been pointed out, has passed through a period of nomenclatorial change, due in all cases to the failure of the organism to grow on an arti- fieial substrate. The original name Histoplasma capsulatum was assigned to it by Darling on the basis that it was a proto- zoon. He described the microbe as small, round or oval, 1-4 y in diameter, possessing a polymorphous chromatin nucleus, basophilic cytoplasm, and achromatic spaces all enclosed within an achromatie refractile capsule. In smears, the para- site to him represented certain features found in protozoa by Leishman, Donovan, and several others. Furthermore, he thought he observed some forms possessing flagellae, and con- sequently he placed the organism in the Flagellidia, or the flagellates as we now know them. This view was held for a number of years. In 1912 and 1913 da Rocha-Lima published papers dealing with lymphangitis in horses due to Cryptococcus farciminosus. He also studied ma- terial taken from cases of Darling's histoplasmosis, using the Romanowsky stain. On the basis of the differential staining reaction he concluded that the organism was a fungus of the genus Cryptococcus and closely related to C. farciminosus. His opinion was based also on the fact that the type of lesion produced simulated that of the above infection. Furthermore, with the Romanowsky dye, protozoa show up with a blue proto- plasm within which is a large red clump called the macronucleus and also a smaller clump, the micronucleus or blepharoblast. This latter inclusion is lacking in Darling's organism. In ad- dition, the type of division of the cell was suggestive of Crypto- coccus. Da Rocha-Lima's idea was further emphasized by Castellani, Neveu-Lemaire (quoted by Vuillemin), and adhered to by most text-books. The unfortunate cireumstance in all this nomenclature was the fact that the organism was not cultured. The problem of naming a microbe of this kind, therefore, depends on the kind of growth obtained on an artificial substrate, in addition to the form in which it exists as a parasite. The morphology of an [VoL, 22 348 ANNALS OF THE MISSOURI BOTANICAL GARDEN organism is usually the first criterion that taxonomists con- sider in classification. In these two fungi, an association of these facts as described led to placing the two species in the genus Posadasia Canton, Posadasia pyriformis being a new species, and P. capsulata a new combination as published in a previous paper (Moore, ’34). The genus Posadasia was created by Canton in 1898 as the representative etiological agent of coccidioidal granuloma of South America. Da Fonseca and Areà Leão (28) supposedly examined Posada's organism and found that the spores de- hisced through the membrane. Such an observation might easily confuse an investigator and lead to an erroneous con- clusion. Illustrations seem to show the organism in tissue to be similar to the asci here described on agar, a point further emphasized by Magalhäes (732) who illustrated the organism with tubereles and named it Neogeotrichum pulmoneum. A series of papers by Floriano de Almeida (’33, 733a, *34, 784a) brings out a relationship between the organisms de- scribed here and Coccidioides immitis and Paracoccidioides brasiliensis as found in the tissues of individuals in South America (Brazil). He deseribed particularly the latter, il- lustrating the typieal club formation found in cultures of Posadasia and present in Paracoccidioides brasiliensis. It is quite evident that these organisms are closely related and that some, such as Coccidioides immitis and Paracoccidioides brasiliensis, as described by Almeida, have a specialized life cycle (Moore, ’32). Posadasia capsulata and P. pyriformis, on the other hand, show a yeast-like stage in the host, a reduced condition which develops on artificial media to the type of re- productive body identical with that of Paracoccidioides. Because of several observations made by various investiga- tors, the genus Posadasia was made synonymous with Coccidi- oides by the author (Moore, ’32). However, after examining Posadasia capsulata and P. pyriformis, and since the work of Almeida, it seems quite apparent that the genus should stand asanentity. Itis on this basis that the two above species are established, showing a relationship to Coccidioides immitis of the family Coccidioideaceae and its relative Paracoccidioides 1935] MOORE—POSADASIA CAPSULATA AND P, PYRIFORMIS 349 brasiliensis which should constitute a genus of the same family. Further, reviewing all data at hand, it seems that Posadasia capsulata and P. pyriformis form the connecting link between the family Coceidioideaceae and the family Endomycetaceae of the Endomycetales. These two fungi, therefore, occupy a posi- tion in the Coceidioideaceae, closely related to the family En- domycetaceae. On the basis of these views, the family Coc- cidioideaceae has the following members : Coccidioideaceae Posadasia Posadasia capsulata Posadasia pyriformis Coccidioides Coccidioides immitis Paracoccidioides Paracoccidioides brasiliensis Rhinosporidium Rhinosporidium Seeberi In recent publications, Redaelli and Ciferri (734; Ciferri and Redaelli, '34) have created the family Histoplasmaceae and in- cluded it with the families Nectaromycetaceae Cif. and Red. and Torulopsidaceae Cif. in the super family Adelosacchar- omycetaceae Guilliermond, comprising all the non spore-bear- ing yeasts. Basing their evidence on the pathology of the host, cultural, morphological and biochemical reactions of the organ- isms, they place Cryptococcus farciminosus Rivolta and Micel- lone, 1883, and Cryptococcus muris Shortt, 1923, in the genus Histoplasma, with H. capsulatum as the type species. Itis dif- ficult to conceive of a relationship existing between these three fungi, and furthermore the presence of asci in Histoplasma (Posadasia) eliminates a family classification in the non spore- forming yeasts. SUMMARY 1. Two cases of Darling's histoplasmosis yielded two spe- cies, Posadasia capsulata, P. pyriformis. 2. The organisms were studied on many substrates covering [Vor. 22 350 ANNALS OF THE MISSOURI BOTANICAL GARDEN a fairly wide range of hydrogen-ion concentration and with a varying amount of carbohydrate and nitrogen. 3. Posadasia capsulata differs from P. pyriformis in its smaller size and amount of growth; spherical tubereulate asci as compared with both spherical and pyriform asci of the lat- ter; white to light Isabella color as compared with Isabella to light einnamon for the latter. 4. The organism oceurs in the host as a small cell 1-4 a in diameter. On an artificial substrate a mycelium is developed which is aerial and also submerged in the medium, giving rise to many characteristic cells as chlamydospores, conidia, rac- quet mycelium. Tuberculate asci develop from globose or clavate cells in the absence of a sexual act. 5. With these two species, the family Coccidioideaceae now has four genera and five species: Posadasia capsulata, P. pyriformis, Coccidioides immitis, Paracoccidioides brasilien- sis, and Rhinosporidium Seeberi. ACKNOWLEDGMENTS The author wishes to express his thanks to Dr. Carroll W. Dodge, Professor of Botany in the Henry Shaw School of Botany of Washington University, and Dr. George T. Moore, Director of the Missouri Botanical Garden, for the courtesies extended; Dr. Charles Thom, Principal Mycologist, United States Department of Agriculture, for the culture of Posadasia capsulata; and Dr. George H. Hansmann of Georgetown Uni- versity, School of Medicine, for the culture of P. pyriformas. BIBLIOGRAPHY — Floriano P. de (733). Notas sobre a Morena de Beppe teo ds iliensis’’ nos tecidos parasitados. Rev. Biol. Hyg. pp. April 1 > (988 Disstosiiooses- em geral e sua niniin. ond e elumitiéagko das blastomieoses. Rev. Assoc. Paul. Med. 3: 270-277. 1933. , (7034) e blastomicosi nel Brasile. Fol. Clin. Biol. 6*: 15 pp. 1934 ; (734a). Formagoes radiadas de membrana dos cogumelos parasitos. Ann. Facul. Med. Univ. d. Paulo 10: 163-174. 7 pl. 14 figs. 1934. Castellani, A., and A. J. Chalmers ('19). Manual of I medicine. pp. 1669- 1670. William Wood & Co., New York, 3rd ed. Ciferri, R., and P. Redaelli (734). Sulla posizione Q^ dell agente patogeno del farcino equino. Boll. Ist. Sier. Mil. 10: Oct. 1934. 1935] MOORE—POSADASIA CAPSULATA AND P. PYRIFORMIS 351 Da Fonseca, O., et A. E. Aréa Leäo (’28). Sur la granulome coccidioidal. Formes d'évolution du parasite dans les tissus, dans le pus des ganglions lymphatiques t dans les cultures. Vue systematique de Coccidioides immitis. Compt. Bend. Soe. Biol. 98: 619-621. 1928. Da Rocha-Lima, H. (’12). Histoplasmosis und epizootische Lymphangitis. Arch. f. Schiffs- u. Tropenhyg. Beih. 16: 79-85. ; (13). Beitrag z er der Blastomykosen. Lymphangitis en und Historia Centralbl. f. Bakt. I. 67: 233-249. 1913. Darling, S. T. (’06). A protozoon Barn A producing cepudotubsteles in the lungs and focal necroses the liver, spleen and lymph nodes. Am. Me Assoc., Jour. 46: 1283-1285. 1906. , (708). m eus A fatal infectious disease resembling kala- azar Found among natives of Tropical America. Arch. Int. Med. 11: 107-123 , (709). The morphology of the parasite (Histoplasma capsulatum ) and the lesions of histoplasmosis. A fatal disease of Tropical America. Jour. p. Med. & Hyg. 11: 515-531. 1909. DeMonbreun, W. A. (733). pees of the micro-organism of Darling’s histo- plasmosis. [title only.] South. Med. Jour. 26: 1005. 1933. , (734). The cultivation and cultural characteristics of Darling’s 934 Hist Dodd, K., and E. H. Tompkins (’33). A case of Darling’s histoplasmosis (so-called Cryptococeus infection) in an infant. [title only.] South. Med. Jour. 26: 1005. 1933. ; (7384). A case of histoplasmosis of Darling in an infant. Amer. Jour. Tog. Med, 14: 127-138. 1934 Hansmann, G. H., and J. R. Schenken (733). New disease caused by a yeast-like organism. and 77: 460. 1933. Magalhäes, O. de (732). elassifieacáo do **Oidiwm brasiliense’’ Neogeotrichum pulmoneum, n.g. (O. Magalhäes, Code emend. O. Magalhäes, 1931. Ins swaldo Cruz, Mem. 26: 151-168. Moore, M. (’32). Coceidioidal granuloma: A classification of e causative agent, Coccidioides "donde Ann. Mo. Bot. Gard. 19: 397—428. '84). Posadasia ren an . capsu c de causative or- ganisms of Darling's histoplasmosis in the United States. Ibid. 21: 347-348 1934, Redaelli, P., and R. Ciferri (734). Affinité entre les agents de l'histoplasmose an du farein aaup et d’une mycose spontanée des muridées. Boll, Sez. Ital. Soc. Int. Micro. 10: Oct. 1934. Vuillemin, P. (’31). Les Tnn parasites et les mycoses de l'homme, En- cyclopédie mycologique. 291 pp. Loco Paris. 1931. [Vor. 22, 1935] 352 ANNALS OF THE MISSOURI BOTANICAL GARDEN ExPLANATION OF PLATE PLATE 11 Posadasia capsulata AU figures drawn as eorrectly as possible with the aid of a camera lucida at a magnification of x 1440 and reduced to x 750. Fig. 1. Group of germinating spores and conidia on lactose agar. Fig. 2. Hypha with elavate SOIN cell on Sabouraud's agar Fig. 3. Mycelium on Raul agar. Fig. 4. Racquet Pr aig on i cel 's agar. Fig. 5. Terminal clavate cell on wort agar Fig. 6. Terminal elavate cell on potato-dextrose agar. Fig. 7. Terminal clavate cell on eorn-meal a Fig. 8. Raequet mycelium with terminal wer cell on malt extraet agar. Fig. 9. Racquet mycelium on wort a, Fig. 10. Spherical hyphal a on malt extract agar. Fig. 11. Blastosporoid cell on Sabouraud’s agar. Fig. 12. Type of mycelium on malt extraet agar. Fig. 14, Racquet formation on lactose agar. Fig. 15, Raequet formation on eorn-meal agar. Fig. 16. Racquet formation on serum agar. Fig. a Raequet formation on serum agar. Fig.20. Lateral elavate cell, presumptive aseus on potato-dextrose agar. Fig. 21. Terminal clavate cell, as fig. 20 on Fig. 22. Young terminal clavate swelling on einn diia agar. Figs. 23-25. Mycelium on Raulin's agar showing swellings, chlamydospores, and conidia. Fig. 26. Young spherical cells, showing oil p on potato-dextrose agar. Fig. 27. Presumptive ascus on glycerine agar Fig. 28. Conidia on corn-meal agar. Fig. 29. Young cell on thin filament, possibly a young ascus on Sabouraud 's agar. Fig.30. Lateral thick-walled resting cell on corn-meal agar. Fig. 31. nnde! clavate cell with oil globules on potato-dextrose agar. Fig.32. Type o celium on malt extract agar Fig. 33. bus d hypnospore on wort agar. Fig. 34. Blastosporoid cell on corn-meal agar. Fig. 35. "Lore cell in laeto-phenol on Richards" agar. Fig. 36. Young aseus in lacto- m. showing internal network on wort agar. Fig. 37. a. on wort a Fig. 38. Mycelium on Raulin y agar. Fig. 39. as on lactose agar Fig. 40. Lateral outgrowth Se win probably develop an ascus on glycerine Fig. 41. Hyphal swellings on potato-dextrose agar. [Vor. 22, 1935] 354 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 11 (Continued) Posadasia capsulata Fig. 42. Hyphal swellings on corn-meal agar. Fig. 43. Young globose cell on lateral filament, on malt extract agar. Fig. 44. Lateral pyriform conidium on potato-dextrose agar. Fig. 45. Lateral spherical conidium on malt extract agar. Fig. 46. Lateral chlamydospore on potato-dextrose agar. Fig. 47. Lateral outgrowth as fig. 40, on Richards” agar. Fig. 48. Lateral pyriform conidium on Richards? agar. Fig. 49. Mycelium on the same agar. Fig. 50. Broken young ascus showing thick wall, on the same medium. Fig. 51. Adult ascus in a water mount, showing oil globules on potato-dextrose i J MOORE—POSADASIA CAPSULATA [Vor. 22, 1935] 356 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 12 Posadasia capsulata All figures drawn as correctly as possible with the aid of a camera lucida at a magnifieation of x 1440 and reduced to x 750, except fig. 71, whieh was drawn at a magnification of x 960 and reduced to x 510. Fig. 52. Mycelium on Richards” agar Fig. 53. Thick-walled cell below rin of Sabouraud 's agar, probably a young Fig. 54, Mycelium on potato-dextrose agar, showing lateral young asci. li 55. Mycelium with hyphal swellings on wort agar Fig. 56. Young ascus with fine prickles on Saboursud 's agar. Figs. 57-58. Asei with blunt tubereles on wort agar. Fig.59. Large spherical cell on nutrient agar mounted in lacto-phenol, showing internal vaeuolation. ig ycelium on potato-dextrose agar, showing asci, conidia, and clavate terminal cell. Figs. 61-63. Types of asci on wort agar Figs. 64-65. Mycelium with asci on filaments, on Cc ped "a Figs. 66-68. Types of asci with different tubereles on wort Fig. 69. Ascus showing oil globules in a water mount from bi agar. Fig. 70. Ascus mounted in laeto-phenol, showing network of cell on nutrient agar. Fig.71. Ascus which had germinated in a gelatine slide eulture, 10 days after inoculation, ERAT "T Cup amm eO WE Y. ds i cas "T E e ) = 7 A * me” zd ad ui , e "pet: TIT AE. ANN. Mo. Bor. GARD., VOL. 22, 1935 PLATE 12 MOORE—POSADASIA CAPSULATA 358 All figure [Vor. 22, 1935] ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 13 Posadasia pyriformis s drawn as correctly as possible with the aid of a camera lucida at a magnifieation of x 1440 and reduced to x 750 Fig. Babouraud 's Fig. 73. Filament with a elavate terminal portion on Raulin's agar. . Hypha on serum agar. . Myeelium with swellings on wort agar . Peeuliar type of lateral branehing on Richards? agar. agar 72. Gr pm of conidia, and small round cells, probably ascospores, on acquet mycelium on potato-dextrose . Branching of hypha on glycerine agar. . Interealary — on Sabouraud's agar. . Myeelium on Richards? agar. . Raequet formation on malt extract agar. Figs 82-83. Enlarged cells on wort agar. . 84, Mycelium on glycerine agar. . Myeelium on eorn-meal agar. . Thick hypha on Richards’ agar. . Swollen interealary cell on Raulin's agar. . Mycelium on wort agar . Interealary chlamydospores on Raulin's agar. aga Figs. 90-91. Mycelium on wor Fig. 92. Lateral outgrowth, showing . an interealary chlamydospore on Richards’ agar. Fig. 93. Terminal clavate cells on Raulin’s agar. Fig. 94. Figs. 95-96. Chains of spherical cells on serum agar. Fig. 97. Fig. 98. Terminal clavate cell, Kg oil RE on Raulin’s agar. Fig. 99. Mycelium on Richards’ ag Fig. 100. Hyphal swellings on qoi Fig. 101 Lateral pedieel with a terminal elavate cell on potato-dextrose agar. acquet mycelium on corn-me agar. . Mycelium showing snake- like processes which are presumptive asci agar bearers on Richards” ig. 102. Terminal Jarabo cell on Raulin's agar. Fig. 103. Mycelium showing intercalary chlamydospore on Raulin's agar. A "TT RM. S ANN. Mo. Bor. GARD., VOL. 22, 1935 —POSADASIA PYRIFORMIS MOORE Ea po. i , T puer e art ttti i m [Vor. 22, 1935] 360 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 14 Posadasia pyriformis All figures drawn as correctly as possible with the aid of a camera lucida at a magnifieation of x 1440 and reduced to x 750 Fig. 104. Enlarged and thickened mycelium on corn-meal agar. Fig. 105. Mycelium on wort agar Fig. 106. Mycelium showing nm _— clavate cell on Sabouraud’s agar. Fig. 107. Same as fig. 106, on eorn-meal a Fig. 108. Hyphal swellings on malt ext E agar. Fig.109. Lateral outgrowth on potato-dextrose agar. Fig. 110. Mycelium on malt extract agar. Fig. 111. Terminal pyriform aseus on malt extract agar. Fig. 112. Probably a conidium on Richards” agar. Fig. 113. Terminal pyriform aseus on corn-meal agar. Fig. 114. Same as fig. 113, on Sabouraud 's agar. Fig. 115. Lateral Ar aseus on corn-meal agar. .116. Same as fig. 115, on the same mediu Figs. 117-131. Various types of asci on rmn media. Figs. 117, 119-120, 124, 126. On Richards? agar. Figs.118, 121, 125, 127, 129—130. On Raulin's agar. Fig. 122. On glycerine agar. Fig.123. On Sabouraud's agar. Fig. 128. On Czapek's agar. Fig. 131. On potato-dextrose agar. OO EE ER ANN. Mo. Bor. GARD., VOL. 22, 1935 PLATE 14 MOORE-—POSADASIA PYRIFORMIS PT. ee AN IMPROVED APPARATUS FOR THE DETERMINA- TION OF CARBON DIOXIDE PRODUCTION IN PHYSIOLOGICAL PLANT STUDIES F. LYLE WYND Henry Shaw School of Botany of Washington University The method of determining carbon dioxide production de- seribed previously! has been used in this laboratory for various types of experiments over a period of two years, particularly in determining the respiration of plants in water culture. At first sight the construction of the apparatus seems complicated, but once assembled it is extremely simple. The manipulation con- sists of adjusting the stopeocks M and W and detaching and weighing the Fleming-Martin bulb. Five units attached to a single water-pump suction line have been successfully oper- ated. One charging of the various chambers with the necessary chemieals is sufficient for months of use. Experience with the apparatus has suggested the advisabil- ity of several modifications of the original construction. The calcium chloride Tower B has been added to prevent the atmospheric moisture from wetting the Ascarite in C. Stop- cock A, through which the air enters, is closed when the ap- paratus is not in use. C is a large test-tube, 32 x 300 mm., and its inlet should terminate near the stopper and the outlet should be from the bottom, in order that the air may be drawn down through the entire mass of Ascarite and thus prevent channelling. Two calcium-chloride towers, H and I, have replaced the former tube of concentrated sulphurie acid for the first stage of the dehydration of the air passing through the bell jar serv- ing as the respiration chamber, This prevents the negative pressure of about one pound per square inch which existed in the carbon-dioxide absorbing part of the system when sul- * Wynd, F. Lyle. An apparatus for the determination of carbon dioxide produe- tion in physiologieal plant studies. Ann. Mo. Bot. Gard. 19: 499-501. 1932. ANN. Mo. Bor. GARD., Vor. 22, 1935. (361) [ VoL. 22 362 ANNALS OF THE MISSOURI BOTANICAL GARDEN phurie acid was used. Even this small negative pressure greatly inereases the likelihood of leaks and limits the number of units which may be attached to the common suction line V. It has been found also that volatile substances from plants are oxidized when drawn through concentrated sulphurie acid, and that the resulting earbon dioxide and sulphur dioxide some- times produce large errors. Fig. 1. Explanation in text. The final stage of drying the air from the respiration cham- ber is aecomplished by the tower J, which is filled with Desicchlora. This drying agent absorbs ammonia as well as water and therefore eliminates the use of a separate structure to remove this omnipresent contamination of laboratory air. The use of phosphorus pentoxide at this stage is very unsatis- factory because of its tendency to channel. The carbon-dioxide absorption bulb is the Fleming-Martin type which has been found very satisfactory for this purpose. The upper chamber is filled with Desicchlora, the lower with Ascarite. Tower L is filled with Desiechlora. M is a three-way stopcock. P is half filled with water and serves as an indicator for the regulation of the flow of air through the apparatus. 1935] WYND—A CARBON DIOXIDE APPARATUS 363 In the operation of the unit, cock W is elosed and chamber N is connected to tower L through the three-way stopcock, M. After the suction has been turned on, eock W is slowly opened until the desired rate of flow is obtained. In stopping the ex- periment the three-way stopcock M must be turned so that air enters through its side arm, then the cock W is closed and the suction stopped. After the suction stops, W may be opened and water will not back up in the system and ruin the drying agent in L. Cock M closes off the tower L from the air and from N when the apparatus is not in use. The apparatus may be operated for periods of twelve hours and even longer, and amounts of carbon dioxide up to several grams may be determined with an experimental error of less than one milligram if the usual analytical precautions are taken. All joints should be sealed with shellac. The detach- able rubber connections at D and E should be sealed with grease. The rubber connections of the Fleming-Martin bulb should be carefully wired with copper wire. The contact of the respiration chamber with the glass plate G should be sealed by vaseline. The absorption bulb must be handled with cheese- cloth, and a counterpoise, preferably another Fleming-Martin bulb, should be used in weighing. The tower J must be com- pletely filled with the drying agent and the short glass-tube outlet should be loosely filled with cotton. If these precautions are not taken, sufficient carbon dioxide, moisture, or ammonia may accumulate in the lower part of J, when the Fleming- Martin bulb is detached for weighing, to cause an appreciable error in the next determination. Since calcium chloride always contains some calcium oxide, the towers containing this substance must be saturated with earbon dioxide before the apparatus is used for the first time. Further details are contained in the author's original paper. NEW SPECIES OF HYDNANGIACEAE SANFORD M. ZELLER Plant Pathologist, Oregon State Agricultural College and Experiment Station Formerly Visiting Fellow, Henry Shaw School of Botany of Washington University AND CARROLL W. DODGE Mycologist to the Missouri Botanical Garden Professor in the Henry Shaw School of Botany of Washington University Since the publication of our monograph of the Hydnangia- ceae has been unavoidably delayed, it seems desirable to record the following new species and new combinations in this group. ARCANGELIELLA africana (Lloyd) Zeller & Dodge, comb. nov. Octaviania africana Lloyd, Myc. Notes 67: 1142. 1922. Octaviana africana Verwoerd, S. Afr. Jour. Sci. 22: 164. ARCANGELIELLA alveolata (Cooke & Massee) Zeller € Dodge, comb. nov. Octaviania alveolata Cooke & Massee, Grevillea 16: 2. 1887. Octavianina alveolata O. Kuntze, Rev. Gen. Pl. 3?: 501. 1898. Gymnomyces pallidus Massee & Rodway, Kew Bull. Misc. Inf. 1898: 125. 1898 Elasmomyces russuloides Setchell, Jour. Mycol. 13: 240- 241. : Hydnangium glabrum Rodway, Papers € Proc. Roy. Soc. Tasmania 1920: 157. 1921. ARCANGELIELLA ambigua Zeller & Dodge, sp. nov. Fruetifieationes pyriformes vel irregulares, lobatae, ad 5 em. diametro metientes, sordide albidae tactu caerulescentes nigrescentesque, ““honey-yellow*? vel ‘‘light olive?” siccatae, Bi glabra, rimosa; basis sterilis prominens, sti- DE suberosa; columella prominens percurrensque ; peridium erassissimum, ad 1500 u EA E hyphis magnis, dense contextis, 4—7.5 y diametro, laetiferis, eum hyphis et funieulis RN a (hyphis tenuioribus ad gle- bam); gleba ‘‘chocolate’’ vel ‘‘wine color,’’ ‘‘blackish brown" vel ‘‘antique nt et ‘‘auburn’’ siccata, locellis parvis, sinuosis; septa 35-45 y crassitudine, Issued May 25, 1935. ANN. Mo. Bor. GARD., Vor. 22, 1935. (365) [Vor. 22 366 ANNALS OF THE MISSOURI BOTANICAL GARDEN fragilia, hyphis magnis dense contextis, lactiferis eum hyphis; basidia 20-24 x 6-8 y, ` clavata, 2- vel 4-spora, sterigmatibus brevibus; sporae obseure brunneae, ovoideae vel ovoideo-citriformes, utrieulo inconscipuo, 9—10-rugoso, immaturae 13-15 x 8-9 u, maturae 11-12.5 x 8-11 u. CALIFORNIA: Santa Clara County, Saratoga, Dale Parks, type (in Univ. Cal. Herb., as H. E. Parks 825 and Z 31; also in Dodge Herb. and Zeller Herb.). ARCANGELIELLA asterosperma (Vittadini) Zeller € Dodge, eomb. nov. Octaviania asterosperma Vittadini, Monogr. Tuberac. 17. 1831. var. depauperata (Tulasne) Zeller € Dodge, comb. nov. Octaviania asterosperma var. depauperata Tulasne, Fung. Hypog. 78. 1851. Octaviania vacua Tulasne, herb. nom. var. hololeuca (Hesse) Zeller € Dodge, comb. nov. Octaviania asterosperma var. hololeuca Hesse, Hypog. Deutschl. 1: 74. 1891. ARCANGELIELLA Beccari (Petri) Zeller & Dodge, comb. nov. Clathrogaster Beccari Petri, Malpighia 14: 126. 1900. ARCANGELIELLA Behrii (Harkness) Zeller & Dodge, eomb. nov. Splanchnomyces Behrii Harkness, Bull. Cal. Acad. Sei. 1: 30. 1884. Hymenogaster Behrii De Toni in Sace. Syll. Fung. 7: 174. 888 var. caudata Zeller & Dodge, comb. nov. Arcangeliella caudata Zeller & Dodge, Ann. Mo. Bot. Gard. 6: 49-52. 1919. ARCANGELIELLA brunneola (Harkness) Zeller & Dodge, comb. nov. Octaviania brunneola Harkness, Proc. Cal. Acad. Sci. Bot. III. 1: 251. 1899, Octaviama microsporium Mattirolo, herb. nom. ARCANGELIELLA Campbellae Berkeley € Broome, sp. nov. Hymenogaster Campbellii Berkeley € Broome, nom. herb. Fructificationes subsphericae, obovoideae vel lobatae, inferne attenuatae, 1-2.5 cm. diametro metientes, caespitosae, violascentes, **elay color”? vel **tawny olive?” 1935] ZELLER AND DODGE—HYDNANGIACEAE 361 siccatae, superficie glabra; basis sterilis conica, in glebam prominens vel percurrens; columella adest, inferne ed Mea ign. spongi osa, Ayi o nehymatie de phis hyalinis; peridium 100-115 , crassitudine, hyphis gelificatis hyalinis cu dd vire in superfieie, emailen in as extero, oes periclinalibus in trato medio et dense periclinalibus in strato intero; gleba ‘‘ ochraceous tawny?’ sic- Fr locellis ab basi sterili radiantibus; septa seissilia, hyalina, prosenchymatica, 1 n crassitudine; basidia non visa; sporae ellipsoideae, apice obtuso, sub- pedieellatae, Pei: 8-10 (-11) x 4.4-5. 6 (-7) m, minute foveolat AUSTRALIA: Victoria, Melbourne (Lilydale), F. Campbell 27a (Mrs. asbl 429 i (in Kew Herb., Lloyd Mus. 0229, and N. Y. Bot. Gard. Herb. marked ‘‘fro Massee Herb.’ ») ARCANGELIELLA Cremea Zeller & Dodge, sp. nov. Fruetifieationes irregulares, subreniformes, 1 x 2 x 2 em., ‘‘light buff’’ vel ““cinnamon buff?! siecatae; columella eremea percurrens; peridium variabile crassitudine ad 250 uy, hyphis magnis, periclinalibus, prosenchymaticum, lactiferis e yphis; gleba firma, **einnamon buff ””; septa 120—130 y erasitudine, pseudo- parenehymatiea, paueis cum hyphis magnis periclinalibus in strat dio; idi brevia, cylindrica, 14 x 8 yw, sterigmatibus tenuibus, ad 4 u Ion ruhen sporae pi luteae, alveolatae, dein ad 24 echinis in circulo magno, 9-11 y diametro. ON: Benton County, Scott's Hill south of Corvallis, R. idrages, type (in Zeller 2 7927, and in Dodge Herb.). ARCANGELIELLA Curtisii Zeller & Dodge, sp. nov. Hydnangium Ravenelii Farlow in Foerste, Bot. Gaz. 19: 37. 1894, non aliorum. a 0.7—1.4 em. diametro, ‘‘isabella color?” vel ‘‘brownish olive?” ae; peridium 100-300 u erai MCA hyphis never s u a in fibrillis eum hyphis contextis; gleba ‘‘pinkish buff”? vel ‘‘cinnamon buff? sic- cata; septa 30-40 y crassitudine, stupposa, seissilia; basidia non visa; sporae sub- sphericae vel late ellipsoideae, 8-11 x 7.4-9 yw, aspere reticulatae. SOUTH CAROLINA: Darlington County, Society Hill, M. 4. Curtis, type (Mo. Bot. Gard. Herb. 5647, and in Curtis Herb. at Farlow Herb.). ARCANGELIELLA ellipsoidea Zeller & Dodge, sp. nov. Fruetifieationes 0.7-3.0 cm. diametro et 1-2 cm. altitudine, o globosae, pyriformes vel reniformes, ‘‘honey-yellow’’ vel tieabella color”? sieeatae; basis sterilis prominens; columella percurrens; peridium 170-350 y erassitudine, prosen- chymaticum, cellulis me eum funieulis E pphisque e lactiferis; gleba ‘‘cinnamon rufous’ vd *hazel,?? locellis magnis; septa 30-65 y crassitudine, BRE ES facile scissilia; basidia non visa; pts: Mi 5 immaturae, ellipsoideae maturae, obseure brunneae, minute alveolatae, 10-12 x 6-7.4 u. TASMANIA: Hobart, L. Rodway 1286, type (in Dodge Herb.) ARCANGELIELLA Gardneri (Zeller & Dodge) Zeller & Dodge, comb. nov. [Vor. 22 368 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gymnomyces Gardneri Zeller & Dodge, Ann. Mo. Bot. Gard. 6: 54—55. 1919 ARCANGELIELLA glabrella Zeller € Dodge, sp. nov. Fructificationes 1-1.5 em. diametro (0.5—1.0 em. sieeatae) metientes, subspheri- e vel depresso-globosae, glabrae, ees vel brunneseentes, sordide albidae vel ** buekthorn-brown" siccatae; columella percurrens, fibrosa, laetiferis eum hyphis insulisque parenehymatieis; peridium ad 640 u crassitudine superne vel 80-130 Pe 1 pot pen dogs gg magnis superfieie E zie contextis t lactiferis eum hyphis; gleba alba, ‘‘ivory yellow”? vel ‘‘cream buff" siecata; Mon iin scissillima, locellis Vubeghenelo hyphis rc lactiferisque; eystidia in finibus hypharum lactiferarum, eylindrica, rupta, spheras laticis exuden- tia; basidia non visa; A hyalinae, laeves vel superne subrugosae, pedicellatae, subsphericae, 4-6 u TASMANIA: Kingston, us Road, L. Rodway 1111, type (in Lloyd Mus., Dodge Herb., and Zeller Herb.) ARCANGELIELLA krjukowensis (Bucholtz) Zeller & Dodge, comb. nov. Secotium (Elasmomyces) krjukowense Bucholtz, Hedwigia 40: 314-315. 1901. var. michailowskjana (Bucholtz) Zeller & Dodge, comb. nov. Secotium (Elasmomyces) michailowskjanum Bucholtz, Hed- wigia 40: 315. 1901. ARCANGELIELLA laevis (Hesse) Zeller & Dodge, comb. nov. Octaviania laevis Hesse, Hypog. Deutschl. 1: 80-81. 1891. Octavianina laevis O. Kuntze, Rev. Gen. Pl. 32: 501. 1898. Octaviania levis ace. Syll. Fung. 11: 169. 1895. ARCANGELIELLA nana (Massee & Rodway) Zeller € Dodge, us nov. menogaster nanus Massee & Rodway, Kew Bull. Misc. Inf. 1899: 180. 1899. ARCANGELIELLA Occidentalis (Harkness) Zeller & Dodge, comb. nov. Octaviania occidentalis Harkness, Proc. Cal. Acad. Sci. Bot. III. 1: 253. 1899 ARCANGELIELLA pilosa Zeller & Dodge, sp. nov. Fructificationes 3—4.5 em. diametro, 1-4 em. altitudine metientes, sphericae - depresso-pyriformes, contractae siceatae, albidae, rufescentes, ‘‘citrine drab,’ 1935] ZELLER AND DODGE—HYDNANGIACEAE 369 ““light brownish olive?” vel ‘‘olive’’ siceatae, laeves, deci basis sterilis prom- inens; eolumella ramosa, percurrens, hyphis laxe implexis; peridium 500-1000 u erassitudine recens (240-320 y siecatum), prosenchymaticum, gelificatum, lactiferis cum hyphis, superficie hyphis radialibus, 30-40 „ longitudine, pilosa; gleba firma, gelifieata, ‘‘buff,’’ dura, ‘‘cinnamon’’ fe **Saecardo's umber?” siccata, locellis parvis, vacuis collabentibus; Mom 20-35 „ erassitudine, hyphis tenuibus, laxe im- plexis, lactiferis cum hyphis; basidia ken, ee sporae hyalinae, sub- sphericae, minute alveolatae, Mure: —12 y diametr CALIFORNIA: Santa Clara County, pe H. E. Parks 4th, Z340, type, 524, 525 (in Zeller Herb. and Dodge Herb.). ARCANGELIELLA Scissilis Zeller & Dodge, sp. nov. Fruetifieationes irregulares, ad 3 cm. diametro, superficie glabra, lutescentes, ““tawny”” siecatae; columella tenuis, alba, mediam fructificationis attingens; peri- dium lentum, faro secernibile, 320-400 u crassitudine siccatum, pseudoparenchy- maticum, superficie prosenchymatica, lactiferis cum hyphis; gleba cremea, ‘‘an- timony yellow?! siccata, densa; septa 25-35 yu crassitudine, prosenchymatica ee eum hyphis; basidia non visa; sporae sphericae, minute ¿relistas, echinulatae, obscure luteae, 11-1 rH idis virginianae. "pen IFORNIA: Humboldt County, Trinidad, H. E. Parks 4125, type (in Zeller Herb.). ARCANGELIELLA Socialis (Harkness) Zeller € Dodge, comb. nov. Octaviania socialis Harkness, Proc. Cal. Acad. Sci. Bot. III. 1: 252. 1899 ARCANGELIELLA tasmanica (Kalehbrenner) Zeller & Dodge, comb. nov. Hydnangium tasmanicum Kalchbrenner in Massee, Grevillea 19: 95. 1891. ARCANGELIELLA Violacea (Massee & Rodway) Zeller & Dodge, comb. nov. Hymenogaster violaceus Massee € Rodway, Kew Bull. Misc. Inf. 1898: 127. 1898. ARCANGELIELLA Vulvaria (Petri) Zeller & Dodge, comb. nov. Clathrogaster vulvarius Petri, Malpighia 14: 126. 1900 MACOWANITES magnus Parks, sp. nov. Pileus 3-14 em. diametro agariciformis, ee expansus, conicus imma- turus, in maturus, marginibus irregularibus vel sinuosis; superfieies laevis, viscida, ‘‘ pale tan" vel obscure brunnea e ““fawn-brown?? vel ‘‘ M inis. i -brown ?” el ‘‘cinnamon-buff’’ siccata; stipes brevis, 3-7 em. longitudine, , ad 3 cm., albus, fragilis, ie lidere ti: subglaber A innato- Sic d hyphis [Vor. 22 310 ANNALS OF THE MISSOURI BOTANICAL GARDEN longitudinalibus, tenuibus; gleba alba, ** warm buff" vel vinacea siccata, spongiosa ut in Gautieria morchelliformá, locellis ab dnas €— liberis vel adnexis, inferne aperta, superne peridio teeta; peridium duplex, strato intero 250-380 u erassitudine, laxe stupposo, funiculis hyphis idi. strato extero 70-120 u, com- pacto, stupposo, hyphis gelificatis, viscido, locellis magnis, labyrinthiformibus; septa ad 170 y (ad 95 „ inter hymenia) crassitudine, stupposa, strato subhymeniali pseudoparenehymatieo; basidia magna, cylindriea vel subclavata, bisporigera, 20-24 x 10-11 y, sterigmatibus brevibus; sporae subsphericae, echinis brevibus, minutis, e pao T inj 10 y diametro. Odor saporque Lactarii dein pungens CALIFO : San ara County, Call-of-the-Wild, H. E. Parks 208, ‘one (in Univ. Cal. Hab, Doles Herb., and Zeller Herb. 2804). ELasmomyces borneensis (Petri) Zeller & Dodge, comb. nov. Octaviania borneensis Petri, Malpighia 14: 128. 1900 Kiiasmomycss echinosporus Zeller & Dodge, comb. nov. Macowanites echinosporus Zeller & Dodge, Ann. Mo. Bot. Gard. 6: 57-58. 1919 SCLEROGASTER Broomeianus Zeller & Dodge, sp. nov. Octaviania compacta Massee, Ann. Bot. 4: 32-33. 1889 [often cited as Monog. Brit. Gast. 32-33. 1889]. SCLEROGASTER Candidus (Tulasne) Zeller & Dodge, comb. nov. Hydnangium candidum Tulasne, Ann. Sci. Nat. Bot. II. 19: 376. 1843. SCLEROGASTER hysterangioides (Tulasne) Zeller € Dodge, comb. nov. "ca dad hysterangioides Tulasne, Fung. Hypog. 76. 1851 Octavian hysterangioides Lloyd, Myc. Notes 67: 1141. 22. SCLEROGASTER luteocarneus (Bresadola) Zeller € Dodge, comb. nov. Octaviama luteocarnea Bresadola, Ann. Mye. 18: 54. 1920. SCLEROGASTER pacificus Zeller & Dodge, sp. nov. Fructificationes oso. gene 0.8-1.0 cm. diametro metientes, albidae; basis sterilis adest; me visa; peridium evanescens, ad 100 , crassitudine, setius Nis nmm cellulis 20 x 30 u; gleba ““ochraceous buff,’’ firma, deinde friabilis, locellis polyhedrieis, sporis impletis; septa tenuia, 20-30 , erassitudine, hyphis tenuibus, laxe implexis; basidia clavata, ^as sporae spherieae, verrueis magnis, 9-10 in cireulo magno, 7-8 „ diametro 1935] ZELLER AND DODGE—HYDNANGIACEAE 311 OREGON: Coos County, Bandon, S. M. Zeller 7425 (in Zeller Herb. and Dodge Herb.). SCLEROGASTER Siculus Zeller & Dodge, sp. nov. Sclerogaster lanatus Mattirolo, Malpighia 14: 85-86. 1900, non Hesse. Fruetifieationes 0.6 x 0.4 em. diametro metientes, depresso-globosae, albidae, floceosae; peridium duplex, strato extero variabili erassitudine, hyphis tenuibus ipM strato intero ad 90 „ erassitudine, hyphis periclinalibus dense contex- tum; gleba ochraceo-fulva, locellis sporis impletis; septa a: p inter hymenia, hyphis perielinalibus; basidia evaneseentia; sporae brunneae, sparse minuteque echinatae, episporio erasso. ITALY: Sicily, Fanfani a Cefalú, forest of S. Cosimo, O. Mattirolo (in Dodge Herb.). Hyonancıum Archeri (Berkeley) Zeller & Dodge, comb. nov. Octaviania Archeri Berkeley in J. D. Hooker, Bot. Antarctic Voy. III. Fl. Tasmaniae 2: 263-265. 1859. Hyvpnancıum citrinum (Harkness) Zeller € Dodge, comb. nov. Octaviania citrina Harkness, Proc. Cal. Acad. Sci. Bot. III. 1: 252. 1899; Sace. & Sydow in Sace. Syll. Fung. 16: 248-249. 1902. Hypnancıum Eisenii (Harkness) Zeller & Dodge, comb. nov. Melanogaster Eisenii Harkness, Proc. Cal. Acad. Sci. Bot. III. 1: 259. 1899. Hypnancium Gilkeyae Zeller & Dodge, sp. nov. Fruetifieationes oblongae vel subsphericae, ad 4.5 x 3 x 3 em. metientes, fragiles, superficie glabra, venis innato-reticulatis, ‘‘buff,’’? brunneo-maculatae, ‘‘pinkis ff’? vel **tawny olive?” erigens venis obscurioribus; basis tesis so non visa; peridium 150-200 „ crassitudine, 70-85 u siccatum, prosenchymaticum, stupposum siccatum; gleba alba vel cremea recens, ‘‘ pale orange yellow" vel ‘‘maize-yellow’’ siccata; septa 35-40 y crassitudine, seissilia in angulis, hyphis magnis laxe im- plexis; basidia uni- vel bispora, elavata; sporae subsphericae, luteo-brunneae, echinis magnis, 3-3.5 u T episporio 0.7—1.0 „ crassitudine, pedicellatae, 18-22 X 14.5-18.5 „ echinis inclus OREGON: Linn County, near er Peoria road, Helen M. Gilkey, type (in Oregon State College Herb., Dodge Herb., and Zeller Herb. 2334). Hyvpnancıum Hessei (O. Kuntze) Zeller & Dodge, comb. nov. Octavianina Hessei O. Kuntze, Rev. Gen. Pl. 3?: 501. 1898. [Vor. 22 312 ANNALS OF THE MISSOURI BOTANICAL GARDEN Hypnanerum laeve (Hesse) Zeller & Dodge, comb. nov. Octaviania laevis Hesse, Hypog. Deutschl. 1: 80-81. 1891. Octavianina laevis O. Kuntze, Rev. Gen. Pl. 32: 501. 1898. Octaviania levis Sace. Syll. Fung. 11: 169. 1895 Hypnanerum lanigerum (Hesse) Zeller & Dodge, comb. nov. Octaviania lamigera Hesse, Hypog. Deutschl. 1: 79-80. 1891; Sacc. Syll. Fung. 11: 169. 1895 Octavianina lanigera O. Kuntze, Rev. Gen. Pl. 3?: 501. 1898. Hyonancıum luteum (Hesse) Zeller € Dodge, comb. nov. Octaviania lutea Hesse, Jahrb. f. wiss. Bot. 16: 255. 1885. Hypnanerum mistiforme (Mattirolo) Zeller & Dodge, comb. nov. Martellia mistiformis Mattirolo, Malpighia 14: 78-82. 1900. Hypnanorum monticola (Harkness) Zeller & Dodge, comb. nov. Octaviania monticola Harkness, Proc. Cal. Acad. Sci. Bot. III. 1: 254. 1899 Hyonancıum pusillum Harkness, sp. nov. Octaviania pusilla Harkness, herb. nom. Fructificationes 1 em. diametro metientes, cinnamomeae; basis sterilis prominens, hemispheriea, eum stipite tenui, brevi; peridium ad mw crassitudine, hyphis hyalinis, perielinalibus; gleba ** Viibthorna bur ’’ locellis magnis, ir repelstibws, vacuis; septa 100 y crassitudine, hyphis tenuibus, hyalinis, gelificatis; basidia late clavata, bispora, sterigmatibus 3-4 u longis; sporae 15 u diametro, echinis longis, tenuibus. CALIFORNIA: H. W. Harkness 282, type (in Dudley Herb. at Leland Stanford Jr. Univ.). HypxaweruM Soehneri Zeller & Dodge, sp. nov. Fructifieationes spherieae vel reniformes, 0.6 x0.9 em., nigro-brunneae vel nigrae siccatae; basis sterilis columellaque non Pari sittin d 0-60 u crassitudine, prosenchymaticum, cellulis parvis, gelificatis; gleba **Sudan eru locellis mag- nis, vaeuis; septa tenuia, subseissilia, du s magnis, brunneis, laxe implexis; basidia — cylindrica, celeriter collabentia; sporae ellipsoideae, obscure brun- neae, 15.4—18.2 2 » MANY: Bayern, Wolfratshausen, Pupplinger Heide, E. Soehner, type (in Boshnar Herb. and Dodge Herb.). Hyvonancıum Thaxteri Zeller € Dodge, sp. nov. Fructificationes sphericae, rubrae, **eapueine yellow" sieeatae; columella incon- spieua si vere adsit; peridium 125-130 , crassitudine, strato extero 30 y crassitudine, 1935] ZELLER AND DODGE—HYDNANGIACEAE 373 hyphis tenuibus dense contextum, strato intero 100 „ erassitudine, hyphis ide. ddp laxe implexis, subpericlinalibus; gleba **orange buff?! vel ‘‘light ochraceous buff,’ locellis minutis; septa 50-60 y crassitudine, hyphis tenuibus contextis, subhymenio Cae tiet plc basidia brevia, eylindrica, sterigmatibus longis; sporae metro, spherieae, echinis 20-94, tenuibus, in circulo magno. cune goccia R. Thaxter (in Farlow Herb.). Hypnancium tuberculatum (Hesse) Zeller € Dodge, comb. nov. Octaviania tuberculata Hesse, Hypog. Deutschl. 1: 75-77. 1891; Sace. Syll. Fung. 11: 169. 1895. UN ales tuberculatum O. Kuntze, Rev. Gen. Pl. 3*: 501. MELANOGASTER AMBIGUUS (Vittadini) Tulasne, Ann. Sci. Nat. Bot. II. 19: var. eurysperma Zeller & Dodge, var. nov. Fruetifieationes spherieae vel irregulares, ad 3.5 em. diametro metientes, ‘‘ snuff- brown? vel ‘‘bister,’’ immutabiles siceatae, superficie laevi; fibrillae paucae, con- colores, non radiantes, adnato-appressae ; dug erassum, 320—400 „ erassitudine, hyphis obscure brunneis, subvesieulosis, Br magnitudine variabilibus; T a septis albis, Per siecatis, loc tin gelificatis, ad 3 mm. dia- metro; septa hyphis tenuibus, ad 2.5 u ai pee sae basidia pyriformia, 5 x a n, pedi- cellis longis, 2.5 u diametro, a sterigmatibus brevibus ad 3 u Vo de eam sporae Be vel eitriformes, nigrae, 10-11 x p. Odor vini rhenan ORE : Polk Cou us Riekreall, Etta pd i (in Zeller Herb. 2690, and in paige Herb.). By a regrettable error the new combination Hymenogaster viscidus was not so designated in our previous paper (Ann. Mo. Bot. Gard. 21: 625-708. 1934). The revised synonymy should read as follows: HyYMENOGASTER viscipus Zeller & Dodge, Ann. Mo. Bot. Gard. 21: 642. 1934. Hysterangium viscidum Massee € Rodway, Kew Bull. Misc. Inf. 1898: 127. 1898, Protoglossum luteum Massee, Grevillea 19: 97. 1891, fide Cunningham, Proc. Linn. Soc. N. S. Wales 59: 169. 1934. Hymenogaster luteus Cunningham, Proc. Linn. Soc. N. Wales 59: 169. 1934, non aliorum. Annals of the . Missouri Botanical Garde Vor. 22 SEPTEMBER, 1935 No.3 AN ANNOTATED CATALOGUE OF THE FLOWERING PLANTS OF MISSOURI* ERNEST J. PALMER Collector and Research Assistant, Arnold Arboretum of Harvard University, Formerly Collector for the Missowri Botanical Garden AND JULIAN A. STEYERMARK Formerly Rufus J. Lackland Research Fellow in the Henry Shaw School of Botany of Washington University PREFACE In 1886 Professor S. M. Tracy of the Department of Botany of the University of Missouri compiled and published a list of the native and introduced plants that had been collected or reported as growing spontaneously in the state. This work, entitled ‘A Catalogue of the Phaenogamous and Vascular Cryptogamous Plants of Missouri,’ listed 1785 species and varieties and constituted at the time a valuable contribution to the botanical knowledge and literature of the state. However, it was far from complete; little collecting had been done in numerous sections, and many of the records were taken from previously compiled lists or from verbal reports of local col- lectors, some of which proved to be unreliable or at least based upon erroneous determinations of material. For these rea- An enumeration of ‘‘The ferns and fern allies of Missouri," which may be eonsidered supplementary to this eatalogue, was published by the authors in the American Fern Journal 22: 105-122. 1932. Issued September 30, 1935. ANN. Mo. Bor. GARD., Vol. 22, 1935. —— (375) [Vor. 22 316 ANNALS OF THE MISSOURI BOTANICAL GARDEN sons, as well as on account of the many changes in nomenclature and in the understanding of species, a considerable number of the plant names found in Professor Tracy's Catalogue do not appear in the present list. The total number of plants now re- corded is, however, much larger, due to new discoveries and to the segregation of many species and varieties not recognized at that time. Since the appearance of the Tracy Catalogue many sections of the state have been made accessible by the building of rail- roads, eleetrie lines, and more recently of automobile high- ways, making it possible now for the collector to get into every county, even into the most rugged and thinly settled parts that were quite unexplored botanically at that time. An inereasing number of students have become interested in the flora and have eoóperated in making it better known through their work in different localities, and a considerable amount of literature, dealing wholly or in part with Missouri plants, has been pub- lished, all of which have contributed to a greatly inereased knowledge of the state flora. Perhaps the most noteworthy change has been in the methods of studying plants and in the attitude of students towards them. Fifty years ago plant catalogues and floras generally were little more than lists of species and genera arranged in families, and not much attention was given to plant associations, succession, ecology, or to the causes determining the distribution of plants and the development of floras, some understanding of which opens up one of the most interesting and significant fields of botanical investigation. In the preface to his ‘Catalogue’ the only previous publica- cations Tracy mentioned as sources of information were “A Partial Catalogue of the Plants of Illinois and Missouri,” based upon collections of Mr. Geyer, and published by Dr. Engelmann in 1884; *The Flora of Jackson County,' by B. F. Bush; and such general works as Gray's * Manual of Botany,' and the gov- ernment reports of King and Wheeler. He acknowledged notes, collections, and assistance received from Professor Swallow, State Geologist, Dr. Pech of Louisiana, Mo., Mr. Geyer, Dr. Engelmann, and others of St. Louis, Professor 1935] PALMER € STEYERMARK—— PLANTS OF MISSOURI 377 G. C. Broadhead of Pleasant Hill, Mr. George W. Letterman of Allenton, Mr. B. F. Bush of Independence, Mr. B. T. Gallo- way of Columbia, and Professor Trelease of Washington Uni- versity. Only a few other works dealing particularly with the flora of Missouri or containing references to it had appeared up to that time, and the list of other collectors, besides those mentioned by Professor Tracy, was not a long one. One of the earliest scientific travelers to leave an account of his botanieal observations in Missouri was Mr. John Brad- bury, an Englishman who traveled extensively in America from 1809 to 1811. In the preface to the account of his journeys, published in Liverpool in 1819, he states that he ar- rived in St. Louis on December 31, 1809, and during the ensuing year made extensive excursions into the wilderness for a dis- tance of 80 or 100 miles for the purpose of collecting plants. The following year he accompanied an expedition, in company with Thomas Nuttall, up the Missouri River and to the Pacific Northwest. There are references in his *Journal' to a number of the trees and other plants observed along the river in Mis- souri, and at the end of the work there is a ** Catalogue of some of the more rare and valuable plants discovered in the vicinity of St. Louis or on the Missouri." It appears to have been the intention of Bradbury to publish a flora or a full list of his botanical collections, but this inten- tion was not carried out for several reasons. Part of his col- lection was lost or damaged in transit to England, and his travels in America were cut short by the War of 1812. Upon his return to England he discovered that the part of his col- lection that had arrived safely had been submitted to Mr. Frederick Pursh, who had described a number of the new species in an appendix to his ‘Flora Americae Septentri- onalis. ^ Thus Pursh, who had traveled in other parts of North America but who had never been west of the Mississippi River, became the first botanical author to describe plants collected in Missouri. In 1818 and 1819 Henry R. Schooleraft made a journey through the almost unexplored Ozark region for the purpose of scientific study. He has left an interesting account of his [Vor. 22 378 ANNALS OF THE MISSOURI BOTANICAL GARDEN adventures and of the country, and although he was more con- cerned with the geology and other features of the natural his- tory than with the flora, he mentions some of the trees and other plants, and his general description of the country gives us a valuable picture of it at that time. About the same time Thomas Nuttall set out on his journey to the Arkansas Territory, traveling down the Mississippi to the mouth of the Arkansas River and making notes on some of the plants seen on the way. Nuttall's direct contributions to the study of Missouri botany are not large, but a number of Ozark species extending into the state from the south are men- tioned or were first described by him. Several of the early reports of the State Geological Survey contain references to the flora or give lists of trees and shrubs. B. F. Shumard, G. C. Swallow, and G. C. Broadhead were amongst the early state geologists who paid some attention to the flora, Broadhead collecting many plants in Cass County, where he lived, and in other parts of the state. Amongst the more important lists of Missouri plants since the publication of Professor Tracy's Catalogue are a ‘Cata- logue of the Phaenogamous and Vascular Cryptogamous Plants in the Vicinity of St. Louis,’ by Henry Eggert, 1891, ‘A Manual of the Flora of Jackson County,’ by K. K. Mackenzie and B. F. Bush, 1902, ‘The Flora of Columbia and Vicinity," by Francis P. Daniels, 1907, and a recent revision of this plant list by H. W. Rickett, *A Preliminary Check List of the Crypto- gams and Phanerogams in the Vicinity of St. Louis," by mem- bers of the Engelmann Botanical Club, 1911, *A Catalogue of the Plants of Jasper County,’ by E. J. Palmer, 1916, and ‘A List of Missouri Trees,’ by F. Dunlap, 1929. Besides these there are numerous papers dealing with special families or genera of Missouri plants or with the flora or ecology of local areas, a complete list of which will be found in the bibliography. In the process of working out the details of the flora of a large and diverse region such as the state of Missouri, it is unavoidable that certain parts should receive more attention or should become known earlier than others. "This is partly due to differences in accessibility or nearness to centers of popula- 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 379 tion or to institutions of learning, and partly because certain students or collectors study the flora of the particular vicinity in which they live, while that of the rest of the state is investi- gated only sporadically by itinerant collectors, usually on hasty expeditions. St. Louis, Jefferson, Franklin, Boone, Jackson, Jasper, and Greene counties seem to be the only ones in the state in which anything like an exhaustive study of the flora has so far been made. In several others extensive collecting has been done and the flora is fairly well known. Amongst these are Clark, Shannon, Dunklin, Butler, Atchison, and McDonald. There has also been considerable exploration in the counties of Ralls, Marion, Pike, Warren, Madison, Iron, St. Francois, Reynolds, Wright, Saline, Stone, Taney, Barry, and Newton, but many of the inconspicuous plants and probably some of the rare ones have never been collected or recorded. The flora of many of the other counties of the state appears to be quite inade- quately known. On the whole, much more botanical work has been done in the area south of the Missouri River than in the northern part of the state, partly due to the fact that the Ozark region is more varied and contains many more rare plants. However, the counties to the north of the great river also hold much of botanical interest, and they would certainly repay a more intensive study. It is quite possible that a number of species of which the range is given as central and southern Missouri, because of lack of other records, will be found to extend also into the northern part of the state. In this catalogue we have attempted to give the range within the state, so far as it is known, for each species, and also to note the type of soil on which it is usually found. The 114 counties of the state have been taken as convenient units for working out the distribution, and these are cited in addition to the general range for many of the less common plants or those confined to certain sections of the state. Common plants that grow in all parts of the state are listed as of general range, and in many cases they can be found in every county. For the purpose of giving the general geographical range of species the state has been divided into three nearly parallel [Vor. 22 380 ANNALS OF THE MISSOURI BOTANICAL GARDEN zones of different width, which are called northern, central, and southern Missouri. Three zones are also recognized from east to west: eastern, middle, and western Missouri. How- ever, since the east and west boundaries of the state are so irregular, it is not practicable to follow meridian lines, and a sketch map is given to show the general extent of these di- visions. County lines are also irregular, and the entire county may be eonsidered as belonging to the division in which the major part of it lies. In the case of a few counties that are approximately evenly divided by these lines, Reynolds and Crawford may be placed with eastern Missouri, Hickory with western Missouri, and St. Louis with central Missouri. We have endeavored to bring the nomenelature up to date according to the International Rules, and in this we have fol- lowed the decisions of the Gray Herbarium for herbaceous plants and that of the Arnold Arboretum for woody species. No complete synonymy is attempted, but to aid the reader in locating unfamiliar names some synonyms are given, and where the name used is one that has not yet appeared in the general manuals, reference is given to the place of publication and in most eases to the name used in the seventh edition of Gray’s ‘Manual’ (G), in the second edition of Britton and Brown's ‘Illustrated Flora’ (B € B), Rydberg's ‘Flora of the Prairies and Plains’ issued in 1932 (R), or in Small's ‘Manual of the Southeastern Flora’ issued in 1933 (S). The abbrevia- tion Rh., which is frequently used, refers to *Rhodora,' and auth. to author or authors. Other abbreviations for the titles of publieations and for geographical names will be obvious, we believe, and most of those used for the names of authors may be found by consulting the list given in the seventh edition of Gray's ‘Manual.’ Popular names of plants are of considerable interest both to botanists and laymen and those known generally or locally are given in this catalogue, but it has not been thought worth while to coin names nor to translate scientific ones. Some of the popular names are quite characteristic, and not all of them seem to have found their way into literature. Amongst examples of these are bodare, chinquapin (oak), yanquapin, 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 381 sheep-sour, tonguegrass, and pursley. Any Missouri farmer, at least of an older generation, can tell you about his bodare hedge—often pronounced ‘‘bodock,’’ and evidently a corrup- tion of the French bois d? arc. Chinquapin and yanquapin are WESTERN \ za \ LU NORTHERN EN CENTRAL rn — Ó— Fig. l. Map of Missouri, divided into zones to show geographieal range of species used in this catalogue. of Indian origin, and the former, commonly used in some sec- tions for Quercus Muhlenbergu without the additional word oak, is sometimes confused with the native Castanea ozarkensis found in the extreme southern part of the state, and to which [Vor. 22 382 ANNALS OF THE MISSOURI BOTANICAL GARDEN it properly belongs. The Missouri schoolboy or girl is familiar with sheep-sour and tonguegrass, although he might not recog- nize sheep sorrel and peppergrass. Likewise the thrifty house- wife in search of spring greens ean tell you the difference between narrow-leaf dock, which is good to eat, and broad-leaf dock, which is bitter, and she knows the excellent qualities of the young sprouts of poke-weed and lamb's quarter (not quarters). The farmer too knows his pursley or ‘‘pusley,’’ but only the city chap has heard of purslane. Crab-apple and not erab is the name given to the native species of Malus, and the woodsman might sniff at hearing a red haw called a thorn or a hawthorn as quickly as he would at the mention of a brook or a marsh, although he knows all about a branch or a slough. Old-man's-beard and Dutchman’s pipe show the whimsical turn of mind, and there is a bit of slightly grim humor in such names as shin oak, beggar's lice, devil's claws, tear-blanket, and wait-a-bit. The principal sources drawn upon in compiling this cata- logue and in working out the distribution of species are, first, the collections in the herbarium of the Missouri Botanical Gar- den, the Gray Herbarium, the Arnold Arboretum, the National Herbarium for the grasses, and in some cases other public and private collections, including those of the University of Missouri, Columbia, the State Museum collection, Jefferson City, the herbaria, mainly local, of the following colleges: Central Wesleyan College, Warrenton; Cotty College, Nevada; Drury College, Springfield; Hannibal-Lagrange College, Han- nibal; Missouri Valley College, Marshall; Park College, Park- ville; State Teachers College, Springfield ; and William Jewell College, Liberty; second, lists and monographs in which Mis- souri specimens are cited ; and third, the unpublished notes and correspondence of the two authors, supplemented by notes and correspondence from other collectors. Free use has also been made of manuals and general works on botany, and these have been cited only where necessary for the sake of clearness, but a full list of the titles will be found in the bibliography. In ad- dition to our previous work, extending in the case of the senior author over a period of thirty-five years, much intensive col- 1935] PALMER & DLEIERMANK PLANTS OF MISSOURI 383 lecting has also been done during the progress of this catalogue. In the summer of 1933 the authors spent several weeks together collecting in various parts of the state; and during the past three years the junior author has made several extended tours, visiting many counties where collections were especially needed to complete the records. As a result of these explora- tions a number of species not previously known in the state have been added to the flora ; others have been found to extend into sections where they were not formerly known, and much valuable information has been secured in regard to the distri- bution and the composition of the flora in the different regions of the state. There seems to be always a temptation in preparing lists and floras to make them as large as possible, and plants of doubtful occurrence or distinctness are often included. We have en- deavored to avoid this error in the present catalogue and to follow a conservative course in admitting species. Many plants that have previously been reported as growing in the state have been omitted because we were unable to find speci- mens of them or to verify their occurrence. In some of the published lists, especially the earlier ones, species were re- ported that we now know to be quite foreign to our flora, and these errors in range have been copied repeatedly. It has been necessary to be on guard for such examples, and it is hoped that one result of this catalogue will be to dispose permanently of the report of plants that have been mistakenly included in the state flora. In preparing the catalogue it has been found necessary to make a number of new combinations in plant names ; one new species, three new varieties, and six new forms are also de- seribed, and names are proposed for four hybrid oaks. All new names and combinations are by the two authors, except where otherwise stated, as in the genus Crataegus, which is treated by the senior author. While the two authors have cooperated and consulted with each other as fully as possible in the preparation of this cata- logue, much of the work has had to be done individually and at two widely separated bases. "This has necessitated a certain [Vor. 22 384 ANNALS OF THE MISSOURI BOTANICAL GARDEN division of labor and considerable independent investigation. The introduction and preface are largely the work of the senior author, the latter being in part a summary of an unpublished paper on the origin and distribution of the Ozark flora begun several years previously. He is also responsible for the re- vision of the genus Crataegus and has done most of the work on the other groups of woody plants. Most of the work of examin- ing collections in herbaria other than that of the Arnold Arbo- retum has devolved upon the junior author, who has checked the large collections of Missouri plants in the herbarium of the Missouri Botanical Garden and has visited numerous other scientific institutions in the state and in other parts of the country, including the Gray Herbarium and the National Herbarium. In the course of this work he has found it nec- essary to question or change the determinations of many specimens and to eliminate a number of plants that had been credited to the state on misidentified specimens. The compil- ing of the bibliography of Missouri botany is also largely the work of the junior author. Due recognition should be given to a number of collectors since the publication of Tracy's Catalogue whose assiduous efforts have contributed much towards extending our knowl- edge of the flora of Missouri. Amongst these are the late Henry Eggert of St. Louis, and G. W. Letterman of Allenton, who worked out so carefully the flora in the vicinity of St. Louis, as well as doing some work in other parts of the state; Mr. John H. Kellogg of St. Louis, who has carried on their work and extended it into many other counties, including par- ticularly Franklin, Jefferson, Phelps, Texas, and Dent coun- ties; Mr. B. F. Bush, the veteran collector and botanieal author of Courtney, who in addition to his long study of the flora of Jackson County, has covered a wide field and done much towards making known the flora of the state; the late Rev. John Davis of Hannibal, who for several years industriously col- lected the plants of Ralls, Marion, and Pike counties, and also added new records from a number of other counties in the northeastern part of the state; the late Professor E. M. Shep- ard of Drury College, Springfield, who through his own collec- tions and those of his students, especially Mr. Paul C. Standley 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 385 and Mr. J. W. Blankenship, helped make known the flora of Greene and several adjoining Ozark counties; and Dr. H. W. Rickett, Dr. Francis Drouet, and others of the botanical de- partment of the University of Missouri, who are reviving an interest there in a state-wide study of the flora and in reorgan- izing the herbarium at the University of Missouri. The list might be further extended to include many others who have made local collections or botanical expeditions into various parts of the state. We wish to express our sincere thanks to Mr. B. F. Bush, who has furnished us many records and specimens for examination. We are under obligation also to the curators and custodians of the botanical collections of the various colleges and museums of the state who have forwarded them to us for examination or placed them at our disposal. Our grateful appreciation is also extended to Mr. C. A. Weatherby, of the Gray Herbarium, for valuable assistance, to Professor Alfred Rehder, Curator of the Herbarium of the Arnold Arboretum, for advice on questions of nomenclature, to Miss Nell C. Horner, Librarian of the Missouri Botanical Garden, for her bibliographie as- sistance, and to Dr. George T. Moore, Director of the Missouri Botanical Garden, for interest shown in the work and for aid and suggestions. INTRODUCTION LOCATION AND AREA Missouri lies somewhat east and a little north of the geo- graphical center of the United States. The northern boundary of the state is along parallel 40° 30’ north; parallel 36? 30’ north forms most of the southern boundary, but a small sec- tion in the southeastern corner extends south to 36?. On the east the state is bounded by the irregular course of the Mis- sissippi River, except for a short distance near the north- eastern corner, where the Des Moines River separates it from Iowa. About two-thirds of the western boundary is a line running from the southwestern corner of the state to the Mis- sourt River in longitude 94? 38' west. North of this, the Missouri River, flowing in a generally southeasterly direction, forms the boundary. [Vor. 22 386 ANNALS OF THE MISSOURI BOTANICAL GARDEN The direct distance from north to south over most of the state 1s about 300 miles, but the southeastern corner extends 35 miles farther south. It is about 390 miles from the most easterly to the most westerly point in the state, but nowhere is the distance in a direct line so great. At the widest place near the southern border the distance is about 325 miles, while at the narrowest, from just above Hannibal to St. Joseph, it is scarcely 200 miles. It will thus be seen that the state extends farther from east to west than from north to south, although a contrary impression might be gained by a cursory glance at the map. Compared with the Atlantie coast, the state lies approxi- mately between the latitudes of Newark, N. J., and Norfolk, Va., or between those of Lisbon and Gibraltar in the Old World. The central and north-central parts of California parallel it on the Pacific side. The area of the state is 69,415 square miles, and it is therefore considerably larger than that of the six New England states. Geographically, Missouri lies in the middle Mississippi val- ley and on the west bank of the great river. The states of Illinois, Kentucky, and Tennessee lie to the east, Iowa to the north, Nebraska, Kansas, and Oklahoma to the west, and Ar- kansas to the south. Thus it touches eight states, which is true of only one other state in the union, the neighboring common- wealth of Tennessee. The state may be broadly described as an undulating plain rising gradually, or abruptly in places, from the Mississippi River towards the west and northwest, where it merges into the Great Plains. This plain is diversified by several pro- nounced features. On the east is the alluvial and recently formed valley of the Mississippi, varying from a narrow strip of sand and gravel at the foot of rocky cliffs to a maximum width of several miles of flood plain and fertile terraces. And across the state a little north of the center, the Missouri River has carved a similar valley, running irregularly, but mainly from west to east. Many of the larger tributaries of these two great rivers dissect the plain and divide the uplands with a network of alluvial bottoms of varying widths. The greater 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 387 part of southern Missouri is occupied by the northern part of the Ozark plateau, which is the most conspicuous and extensive relief feature between the two great mountain systems of the Appalachians and the Cordilleras. North of the Missouri River and west of the Ozark region, the plain is less diversified, and the rolling surface, interrupted only by the usually shallow valleys of the streams, rises gradually and rather uniformly towards the northwest. Finally, in the southeastern corner, abruptly separated from the Ozark plateau, there is a small area of low relief, which is a northward extension of the Coastal plain. Thus, three principal physiographic regions, the South- eastern lowlands, the Ozark region, and the Prairie region, may be recognized in the state, and since the boundaries of the main plant provinces correspond generally with these, it will be worth while to describe them in some detail. ELEVATION AND DRAINAGE 'The rise in the elevation of the land surface from the level of the Mississippi Riveris very irregular. In the Southeastern lowlands the altitude above tide level over most of the area is between 280 and 350 feet, and the generally level surface is re- lieved by only one slight elevation along Crowleys Ridge, in Dunklin, Stoddard, and Scott counties. This reaches a maxi- mum height of about 450 feet. In the Ozark region the rise is generally abrupt, and the river, except sometimes for a narrow strip of valley, is bordered by precipitous bluffs. In the Iron Mountain sub-region, about 75 miles west of the river, the sur- face rises in many places to over 1,000 feet above tide level, and the altitude of the highest points at Tom Sauk and Wild Cat Mountains is approximately 1,800 feet. Most of the area of the Ozark uplands farther west has an elevation of between 1,000 and 1,300 feet and it reaches a maximum of slightly over 1,700 feet in Wright County. On the northern and north- western sides there is a gradual falling off towards the valleys of the Missouri and Osage rivers. Between these two streams a local area in Morgan County again rises to over 1,000 feet, and in Barry County, near the southwestern corner of the state, the highlands reach an altitude of over 1,500 feet. In the [Vor. 22 388 ANNALS OF THE MISSOURI BOTANICAL GARDEN Prairie region, north of the Missouri, the altitude rises from about 600 feet in St. Charles County to a little over 1,200 feet at the northwestern corner of the state. The drainage of the entire state is into the Mississippi- Missouri River system, but the considerable differences in elevation and in the hardness and strike of the underlying rocks, and other features of its complex topography have caused the tributary streams to run in various and sometimes circuitous directions before reaching the great rivers. In the northern part of the state the general direction of the streams is southward into the Missouri River, except along the eastern quarter, where they flow mostly southeast into the Mississippi. Most of the streams of the Prairie region are slow-flowing or somewhat sluggish, and in very few places have they developed bluffs or banks more than a few feet high. This is due to the generally soft charaeter of the formations through which they have worn their channels, which is usually glacial drift or Pennsylvanian sandstones and shales. None of the streams in this part of the state attain much size, the largest being the Grand and Chariton rivers. South of the Missouri River, the drainage from the Ozark plateau, where nearly all of the streams have their sources, is in various directions. In the southeastern part and to beyond the center of the state along the southern border, most of them have a generally southerly course. The St. Francis, Black River, Current River, and the North Fork of White River are the most important streams in this area. Along the northern side of the plateau the Meramec flows northeast, east, and finally southeast before reaching the Mis- sissippi; the Gasconade and its tributaries flow north and northeast into the Missouri, and the Osage, the largest tribu- tary of that river in the state, flows in a circuitous course, in the main eastward and along the northern side of the plateau, before joining it a little east of the center of the state. In its eastward course the Osage captures a number of north-flowing streams. White River, one of the larger tributaries of the Mississippi, which has its sources in the Boston Mountains of western 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 389 Arkansas, through a great curve to the north flows for a short distance through southern Missouri, in Barry, Stone, and Taney counties, before again turning southeast to cut its nar- row valley through some of the most rugged parts of the southern Ozarks, and finally to emerge into the broad lowlands and to join the Mississippi far to the south in Arkansas. In Missouri it receives one large tributary, the James River, and several smaller ones from the north. The influence of this stream on the flora is very great, as it seems to have been the gateway by which many plants from the southwest and others from the southeast have entered. In the southwestern corner, in the counties of McDonald, Newton, Jasper, and the southern part of Barton, the drainage is westward through Spring River, Shoal Creek, and Elk River, which join the Neosho farther west; and through that stream their waters find their way into the Arkansas and finally into the Mississippi. The direction of these streams has also had an influence on the flora, which is evident in the presence of many species from the Southwest in the area which they drain. Many of the Ozark streams have their sources in large springs, and the majority are fed by springs along their courses. Most of them are perennial, but often when not at flood stage the water may disappear under the beds of coarse gravel and boulders for some distance, only to emerge again where a ledge of solid rock comes to the surface. Usually they are swift-flowing and subject to rapid rises, sometimes with disastrous results after heavy rains. Only the larger ones have reached the stage of forming alluvial valleys. These are usually bounded by precipitous bluffs, and on one side or other of the valley the stream in its meanderings will often closely approach one of these. Narrow V-shaped valleys and cañons are common along the smaller streams in the more rugged parts of the region, and these, as well as the high bluffs, afford protection to many rare plants, which are generally species of northern range or of the southern Appalachians, although in some situations they are plants of southwestern distribution. The meandering courses of many of the Ozark streams, and sometimes their general direction for considerable distances, [Vor. 22 390 ANNALS OF THE MISSOURI BOTANICAL GARDEN seem to have been influenced or determined by differences in the hardness of the rocks, or by the structure and pitch of the strata through which they have cut their channels, and in some cases there is evidence that their original courses have been much changed or reversed by geological forces during the last cycle of elevation of the region. All the streams that flow through the Southeastern lowlands have their sources in the Ozark plateau, but in their lower eourses they are sluggish, with scarcely defined valleys, and they are often bordered with wide swampy areas. Ox-bow bends and shallow bayous, sometimes widening into shallow lakes, are frequent along their courses, and before the present system of artifieial drainage was inaugurated, in times of flood their waters spread over almost all of the area, often ris- ing to a height of two or three feet on the trunks of the forest trees. Such eonditions naturally had a marked effect upon the flora, completely inhibiting many species and restricting others to the low ridges and knolls, while favoring aquatic and palustral plants, which formed a large percentage of the flora. CLIMATE The climate of Missouri, like that of all the central Missis- sippi Valley, is marked by great extremes and is subject to sudden changes of temperature. Cyclonic storms and occa- sionally destructive tornadoes, the latter generally very local, occur throughout the spring and summer months. These some- times terminate periods of extreme heat and humidity, and are followed by longer intervals of mild pleasant weather. The storms are frequently attended by electrical discharges and by rain, which falls in torrential quantities ; and there is some- times also hail, that may do great damage over limited areas. The winters are generally mild and open for the most part, but they are interrupted by occasional severe storms or ‘‘bliz- zards’’ that originate in high altitudes and sweep down across the plains, bringing sudden drops of temperature, sometimes of 30° or 40° or more in a few hours. The total annual rainfall averages from 40 to 45 inches over the southern part of the state, and it diminishes to 30 or 35 inches in the northwestern 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 391 part. However, it is often too unevenly distributed throughout the season to be of the greatest value to plant life. Most of it falls during the spring and early summer months, and there is sometimes a short rainy season in September or October; but protraeted droughts, that often occur in the late summer and sometimes earlier, have a destructive or limiting effect upon many plants. And this is one of the causes for the absence of some speeies found in similar or more northern latitudes nearer to the coast or where more equable conditions prevail. The direction of the winds is quite variable. In the spring and summer the prevailing direction is from the south and west, and in the autumn or winter it is often from the north or northwest. Although the severe winter storms come from these direetions, the west and northwest winds of summer are often hot and dry and at other times storm-laden, while those from the south, which originate in the Gulf-Caribbean cyclone center, because of the distance and the altitude at which they travel, are usually cool and refreshing. The east winds of spring and early summer are frequently chilly and damp. In the spring and fall months the winds are often strong and blow steadily, sometimes for several days. The winds influence the flora in several ways. As carriers of moisture or as agents in its evaporation, their effects are most important. The strong dry winds are at least one of the faetors that have prevented the spread of forests and have kept the western plains and prairies as open grass lands. They are also an important agency in the transportation of seeds, and as an erosive force and as carriers of soil, dust, and sand, their effect, though not so conspieuous as in drier or more sandy regions, is by no means negligible. The generally small size and stocky character of the trees on open uplands is at least in part due to the strong winds. The winter and early spring blizzards are generally accom- panied by snow or sleet. The amount of snowfall varies greatly in different years. Some winters pass with only a few light snows, aggregating not more than two or three inches in thiekness, while at rare intervals there may be a season in which there are repeated falls, and in small areas in more [Vor. 22 392 ANNALS OF THE MISSOURI BOTANICAL GARDEN northern parts a maximum of two or three feet on the level is sometimes formed. Memory of such exceptional seasons gives rise to the popular belief in a changing climate and to the tradition of the ‘‘old-fashioned’’ winters. Snows gen- erally melt rapidly, but when they remain on the ground for some time they have a favorable influence upon vegetation, especially upon grasses and herbaceous plants. One of the most unfavorable climatie influences upon the flora in Missouri is the comparatively late dates at which killing frosts occur. Sometimes these come as late as the first week in May, and their effect then upon tender plants is most disastrous, often de- stroying the fruit or the entire plant for the season and in some cases killing them outright. This has probably been one of the most effective barriers to the northward spread of many tender or less hardy species. The influence of climate upon the vegetation is very great, and climatie eonditions operating in the past have doubtless been one of the principal factors in determining the present composition of our flora. Rich and varied as the flora of Mis- souri is, its diversity is due largely to other causes, such as the variety of soils, drainage, exposures, or protection afforded in different localities. For it is the extremes of moisture and temperature and not the averages that affect tender plants and work destruetion upon them, and the temperatures oc- casionally recorded in Missouri winters are lower than those on the Atlantic coast far up into New England. The long se- vere droughts also prevent the spread of many mesophytie or hydrophytie species, limiting them to restricted areas or en- tirely excluding them from the region. Also the torrential rains and resulting floods in the stream valleys cause rapid erosion on steep slopes and prevent the accumulation of soil in roeky areas, as well as submerging large tracts of lowlands for considerable periods. The following tables showing the average and the maximum and minimum temperature and rainfall by months, taken from the records of the United States Weather Bureau, will illus- trate some of the extremes of climate in Missouri. 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 393 CLIMATE OF COLUMBIA, BOONE COUNTY, MISSOURI YEARS” RECORD)* Elevation above sea level 784 feet. Mean Mean Mean Mean Average MONTH temp. max. min. rainfall | snowfall Deine i aia ip BS inches inches inds from December 33 42 24 1.86 3.8 SW January 29 39 20 1.92 5.8 S February 32 41 22 2.04 6.0 NW Winter 31 41 22 5.82 154] SW March 43 54 33 2.94 37 S April 55 65 44 5 brin 0.5 S May 64 75 53 4.46 E S Spring 54 65 43 11.17 4.9 S June 73 83 62 4.76 0 S July 7T 88 66 3.49 0 S August 76 87 65 3.60 0 S Summer 75 86 64 11.85 0 S September 68 80 58 4.35 0 SE Oetober 57 69 46 2.61 0.1 S November 43 54 34 2.20 1.0 S Autumn 56 68 46 9.16 1.1 S YEAR 54 65 44 38.00 20.4 S Highest temperature in 41 years, 111; lowest i y Hi years, —26. Average date last killing frost in spring, April 1 Average date first killing frost in —— mn, Okicher 18. Latest date killing frost in spring, May 9. on TE er in a P mn, September verage length growing season (K- -frost to K: Di), 188 rn Fe annual number of pred with 0.01 inch mms in mo verage number days 0.01 inch or more rain, April y cade. am 64. nehes. ehes. The wettest single month, June, 1928, with 14.86 EN. The driest single month, "August, 1008, Pa 0.06 i ide Sy M ges veloeity, Janua ary 9 mil y Praia lo miles; March 10 miles; Apr miles; May 8 miles; June 7 miles; July 7 miles; August 7 werd September 7 miles; October 8 miles; November 9 miles; December 9 miles. 18. wind ki of yr ram per hour or more, about 2 a yea ind velocities of 47 miles per hour or over, about 1 n 10. yea Highest elocity ever ents d th or o pe of 5 ah vA 50 pi Tes Extreme or one minute, 57 miles per The pe cd known va de frost pue pe "y the ground, 36 inches Authority for ground frost penetration city engineers, grave diggers, and telephone-hole diggers. * Furnished by the United States Weather Bureau Office. (VoL. 22 394 ANNALS OF THE MISSOURI BOTANICAL GARDEN GEOLOGY AND TOPOGRAPHY Since plants grow in and are entirely dependent upon the soils, it is important to consider them and the rock formations from which they are derived in a general study of the flora. Moreover, clues to many perplexing facts in regard to the pres- ent distribution of native plants can often be found only in the past geological history of the region as well as in a study of its present topography and ecology. All of the state of Missouri, with the exception of the South- eastern lowlands, has been above sea level since late Paleozoic time. Parts of the Ozark region are amongst the oldest lands on the continent, the Archaean and Algonkian rocks of the Iron Mountain sub-region having been a part of the original land axis of North America. The Iron and St. Francois Mountains in the southeastern part of the state are remnants of an old system, the peaks of which doubtless once stood much higher than they do at persent, and the granitic and porphyritic rocks of which they are composed have during ages of erosion fur- nished material for the formation of the sedimentary beds which were later laid down in the seas about their bases. They may at one time have been entirely submerged and buried by sediments from higher land areas, and they are now in the process of being excavated by erosion. About this nucleus, beds of sandstones, limestones, and dolomites, and less gen- erally of shales and conglomerates, were being formed in con- centrie bands and in overlapping layers down to the close of the Carboniferous or the beginning of the Permian period, when the whole region was gradually uplifted above sea level. Although no traces of them now remain, we can safely as- sume that the earliest forms of land plants once occupied this region, and that the whole long pageant of the development of plant life has passed in review here. The luxuriant fern forests of the Coal period flourished in the lowlands and marshes bordering the seas, as the abundant fossil relies found in many places bear witness, and as the higher classes of flower- ing plants appeared at least in the Cretaceous and succeeding ages they found in the Ozark region a land surface already old 1935] PALMER € STEYERMARK—— PLANTS OF MISSOURI 395 and with a diversity of soils and ecological conditions suited to a varied flora. So our present flora has an ancient lineage, and there is reason to believe that many of the rarer plants found isolated in parts of the region may have survived from remote times. During the Cambrian and Ordovician periods most of the present area of Missouri was covered by seas of varying depths. Thick beds of magnesian limestone or dolomite were deposited in the deep seas and sandstones nearer the shores. These deposits now lie deeply buried under later strata over most of the state, but they come to the surface over much of the Ozark region. Narrow belts of Silurian and Devonian limestones and shales are found along the eastern and north- eastern sides of the present Ozark plateau, and more locally to the west and southwest, but most of the region seems to have been a land surface during those times. Stratified rocks of the Mississippian series, mostly pure limestones with interbedded chert and some more local beds of sandstone and shale, also encircle the Ozark plateau in a wider belt, coming to the sur- face over considerable areas in northeastern Missouri and in the west-central and southwestern parts of the state. Farther north, as well as west, and eastward beyond the borders of the state, they are buried beneath the sandstones and shales of the Pennsylvanian series. The shallow seas of the late Carbon- iferous period probably at one time covered nearly the entire state, as outlying areas of the rock deposits are found in many parts of the Ozark region, though in most places they have been removed by subsequent erosion. But over most of northern Missouri and south of the Missouri River on the western side of the state they come to the surface, except where buried under transported glacial material. Limestones formed in the more open seas at this time are also found in some places. Nearly all of Missouri was a land surface undergoing erosion through the middle ages (Mesozoic) of geologic time, although there were probably considerable changes of level during the late Cretaceous, when igneous rocks were being thrust up in the Ouachita Mountain region of central Arkansas. At that [VoL. 22 396 ANNALS OF THE MISSOURI BOTANICAL GARDEN time the waters of the Gulf of Mexico came up to the south- eastern border of the present Ozark region, and deposits made along the shores come to the surface in Scott and Stoddard counties in parts of Crowleys Ridge in the southeastern Mis- souri lowlands.? The upward movement of land surfaces at that time probably affected the drainage of the entire region and changed the directions of many streams. The upper Mis- sissippi River may not have assumed its present course until about this time.? The Mississippi Embayment of southeastern Missouri re- mained until a much later time, at least until the last elevation of the Ozark region, towards the close of the Tertiary period. The Southeastern lowlands now occupy the area of this old embayment. Prior to this time the surface of the Ozark up- lands was being worn down until most of the region was re- duced to a low, comparatively level plain, with sluggish streams and probably large areas of swamps, covered with a forest in many respects similar to that which now occupies the South- eastern lowlands. The probably slow and long-continued movement that again elevated the region resulted in a rejuve- nation of the streams, which, because of the steep grades, began eutting new valleys and producing the rough topography that is now found especially along the steep slopes of the dome. The cold climate and the several southward movements of the polar ice sheets during the Pleistocene period must have profoundly affected the flora of the entire northern and central United States. In Missouri during the Kansan epoch, one of the earliest advances of the ice, the ice sheet covered a large part of what is now the Prairie region north of the Missouri River, bringing down from the north great quantities of gravel and boulders, mostly of igneous and metamorphie rocks such as are not found in place nearer than Minnesota, the Dakotas, and Canada. As the ice melted and retreated the region it had * Matthes, F. E. Cretaceous sediments in Crowleys Ridge, southeastern Missouri. Bull, Am. Assoc. Petrol. Geol. 17: 1003-1009. 1933; also Farrar, W. The geology and bleaching clays of southeast Missouri. Part I. The Cretaceous and Tertiary Geology. Mo. Geol. Surv. Bien. Rept. Appendix IL. pp. 7-20. 1935. * Marbut, C. F., Physical features of Missouri. Mo. Geol. aso. 10: 76. 1896. 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 397 covered was left with these, as well as a deep mantle of soil and clay, the result of the grinding and pulverizing of the rocks over which it had passed. Later, much of this material was taken up and redistributed over wider areas by the waters of the flooded streams. The ice sheet may have come as far south at this time as Clinton, Carroll, and Randolph counties.* In the later Iowan epoch the ice seems to have again advanced to just within the northern borders of Missouri. The thick deposits of loess, which are so conspicuous above the bluffs of the Missouri River from Atchison County to be- low Kansas City and which also occur along the Mississippi River on the eastern side, were probably deposited in shallow lakes or sluggish streams during this time of the retreating ice. The most recent effects of geological forces in the state can be seen in the alluvial valleys of streams, in eroded surfaces and ravines that result from every flood, and in the cumulative effects of frost, wind, and rain, all of which go on every year, modifying the surface of the land and influencing the vegetation. The present topography and surface geology of the state is the result of this long geologic history, every cycle and change of which has had its influence in determining the present char- acter and distribution of the flora. Some plants are restricted to special kinds of soil and others show a preference for certain ones. And since most of the soils in Missouri, except over the glaciated areas of the Prairie region and in the alluvial valleys of streams, are residual and have resulted from the breaking down of the local rocks, the relation between the underlying geological formation and the plants that grow above them is often very close. Plants that grow on alkaline soils, or those containing a large percentage of lime, are called ealeiphiles. Soils and plants of this class are usually found in limestone areas. Acid or sub- acid soils, deficient in lime, are generally found in areas where ‘Todd, J. E., Formation of the quaternary deposits of Missouri. Mo. Geol. Surv. 10: 213. 1896. [Vor. 22 398 ANNALS OF THE MISSOURI BOTANICAL GARDEN sandstones, cherts, and other siliceous or granitie rocks come to the surface. But acidity may result locally from other causes, and in the Ozark region this type of soil is often due to the leaching out of the lime and the accumulations of chert that was originally present as lenses or nodules in the limestone strata, or to the coneentration of magnesium in dolomite areas. Plants that grow on acid soils are called oxylophiles, but most plants are not absolutely restricted to one type of soil, though showing a more or less decided preference for it. Another group of plants, including by far the greatest number, are found on soils of not too pronounced a character, but which may be either neutral, sub-acid, or sub-alkaline. Such plants are described as circumneutral. Others, such as many of the common weeds, grow on almost any sort of soil, where other conditions are not too unfavorable, and these are said to be indifferent. Tue FLORA In its broadest sense the term flora embraces all forms of vegetation found within a region, including, besides the higher or flowering plants, mosses, lichens, fungi, algae, and other less conspicuous and mieroscopie groups. But it is generally used in a much more restricted way, and in speaking of the flora most writers and students have in mind only the more con- spicuous groups of the phanerogams and vascular eryptogams, and it is in this sense that the term is used in this catalogue. The character of the vegetation in any region is determined by the physical conditions, and the requirements of different sorts of plants vary widely. Some grow directly in water or require a constant and abundant water supply; others grow in drier situations or can live only in places that are usually very dry and well drained. Some require shade or the support or protection of other species; and some thrive only with abun- dant light and sunshine. Many forms of vegetation are very sensitive to frost and cold and are limited to tropical or sub- tropical regions; while others, through long periods of dor- mancy, have become inured to severe cold. Within every region 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 399 of any considerable extent a wide variation in many or all of these conditions is found, giving opportunity for plants of dif- ferent requirements within certain limits to find a place in the flora. But the limits of a region and its flora are defined more or less sharply by physical barriers or by differences in ecologic conditions. Climatic zones of temperature or rainfall, geo- graphic barriers, such as oceans, high mountain ranges, deserts, or wide river valleys, or sudden changes in the char- acter of the soil or drainage, are amongst the things that often determine the boundaries of floras. Some of these barriers, however, are not absolute. The effect of climate asserts itself only gradually and in different degrees upon different sorts of plants, so that the boundaries between different floras or sub- divisions of floras are generally not sharp and clear, but there is a border line along which one merges into another. This is particularly true when speaking of the flora of a limited area. In considering the flora of Missouri, it should be remembered that it is only a part of the flora of eastern and middle North America, and that while several different plant regions can be recognized within the state, the boundaries between none of them are absolute, and that the range of many species extends from one region to adjoining ones and often through- out the entire state. But at the same time each of the principal plant regions, and to a less extent each subdivision of them, is distinguished by the presence of a number of species restricted to it, and by certain characteristics in the type and composition of the flora. PLANT REGIONS The principal plant regions of the state, corresponding in general to the physiographic regions previously described, are the Southeastern lowlands, the Ozark region, and the Prairie region, but the boundaries of the physiographic and phytogeo- graphic regions are not quite identical. In places the flora of the lowlands pushes for short distances up the river valleys into the Ozark plateau, and the flora of the latter extends north- ward beyond the plateau and is the dominant element along the . [Vor. 22 400 ANNALS OF THE MISSOURI BOTANICAL GARDEN bluffs north of the Missouri River and on the hills bordering that stream as far west as the mouth of the Chariton. On the western side it also occupies the river bluffs and rocky uplands in what is mainly the Prairie region, while, on the other hand, wedge-like extensions of the prairie flora penetrate the Ozark region, and colonies of prairie plants occupy glades and open- ings in the forest. Lists of characteristic plants are given after the description of each of the plant regions. These are intended to include such species as are wholly or mainly confined to the region, and many of the most common and conspicuous plants may not be mentioned because of the fact that they are equally common in other regions. SOUTHEASTERN LOWLANDS REGION In the Southeastern lowlands there are no outerops of solid rocks except a few outlying remnants along the border of the Ozarks. Much of the soil has been transported as alluvium by streams, and in places it has been enriched by humus from the decay of the forests. Tn other parts it has become acid through leaching, lack of drainage, or the influence of the decaying vege- tation. On some of the slight elevations the soil is sandy, and this may have been washed down locally into depressions, forming sandy bogs or bayous, with an acid-loving type of vegetation. Most of the soils in this region are either sub-acid or neutral, with areas of a definitely acid type and others that are mildly alkaline. The drainage is generally poor, except on Crowleys Ridge, and although there are many plants here not found in other parts of the state, the flora is rather uniform over most of the region. The characteristic and dominant plants are those of the Gulf Coastal Plain, and the region as a whole is Lower Austral, and with the southern phase of the Carolinian flora. Nearly all of the region was originally forested with a dense and luxuriant growth of deciduous trees. In the swampy or more frequently inundated areas, bald eypress, tupelo (Nyssa aquatica), pumpkin ash (Fraxinus profunda), red maple, swamp hickory, water locust, swamp cottonwood (Populus 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 401 heterophylla), water elm (Planera aquatica), overcup oak, itea, button-bush, and a few other trees and shrubs are dominant. Oo ojo y Ó "o )) 080 oo 0 00 5 O o co f Ó Olo 0 O oro O oo fe} o OOo o Ol O doo Ojoo o o pao o s o o 9 el! o (JO O xis of 2 A +, = "77. PPPA le / O gjo e, S ES Zs 223s SS A oA IB: 00 E 4% 7 ps YZ 9 ESA C 771227 RG BEE AS de C/ 444: 1000 "v SKY AAP 444444 099.198 SA i E PA FF. Os + NANG NY Prairie Region (Loess mounds) Prairie Region (unglaciated) Prairie Region (glaciated) Ozark Region j| Southeastern Lowlands Fig. 2. Map of Missouri, showing the plant regions. There are comparatively few herbaceous plants except aquatics, although coarse sedges and grasses and some others grow along the margins of the sloughs and swamps. On the [Vor. 22 402 ANNALS OF THE MISSOURI BOTANICAL GARDEN slightly higher ground above the flood stage, or where it is seldom inundated, the flora is much richer in species. The forest here contains many sorts of oaks, white hickory, sweet gum, black gum, winged elm, white ash, flowering dogwood, and other trees, besides a variety of shrubs and vines and a large array of herbaceous plants. Yellow pine (Pinus echinata) is found rarely on the higher sandy ground, but there are no other conifers and seldom any ericaceous plants. Most of the forest species found on the drier parts of the lowlands are also found on Crowleys Ridge, and growing with them are many others that are practically restricted to this elevation in the region. Some of the latter are species of distinetly eastern or northeastern distribution which are near their western limits here, at least in this latitude. Beech, Spanish oak (Quercus falcata), tulip-tree, cucumber-tree, sweet gum, linden (Tilia glabra), and sugar maple are amongst the commonest forest trees, and the scarlet oak and beaked hazelnut are also rarely found. Geological evidence indicates that Crowleys Ridge lay to the east of the Mississippi River until comparatively re- cently, when the channel of the river changed, and the flora would seem to lend support to this. The flora of the Southeastern lowlands, with the exception of the eastern element found on Crowleys Ridge, is probably a very ancient one. Except for the loss of some of its more tropi- cal species, it may have changed only slightly since middle or early Tertiary time, when it extended far beyond its present limits. In Missouri it probably occupied the lowlands to the north and west before the elevation of the Ozark plateau, where traces of it are still to be found, and it may have covered most of the state before the incursion of the northern ice sheet.’ * Evidence is rapidly aceumulating that seems to make it necessary to revise the ideas previously held as to the intensity of the cold that accompanied the ad vanee of the glaciers in the Quaternary era, and of its completely destructive effects upon the flora. It now seems more probable that fragments of the forest and associated herbaceous flora continued to flourish in parts of the generally glaciated regions, especially toward the southern limits of the ice, although doubtless losing many of the less hardy species. Similar conditions are found today in regions where glaciers are present and where a varied flora is sometimes found growing almost up to the edge of the ice. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 403 Fossil remains of the bald cypress, which now reaches its northern limits here, have been found in Tertiary and Pleisto- cene deposits over a large part of North America, reaching as far north as the St. Lawrence valley. And other fossil as well as living plants prove the former wide extent of this southern flora. The physiography of the Ozark region and of regions surrounding it was at that time very different, and it has since also undergone great climatic changes in the course of which plants have invaded the region from different directions. But it is interesting to find that a number of species which are gen- erally of coastal distribution and that are now mainly confined to the Southeastern lowlands in Missouri reappear in isolated stations far up the Mississippi River and sometimes in remote parts of the Ozark region. The most extensive colonies of such plants are along White River and its tributaries, and some species extend to the western boundaries of the state. As the White River country belongs to the Ozark region, of which it is a rather distinct subdivision, it will be described later, but a few examples of the extension of lowland species into other parts of the state may be mentioned here. The green haw (Crataegus viridis) is a common species throughout the Coastal plain from Virginia and Florida to eastern Texas, and in Missouri it is abundant in wet woods in the Southeastern lowlands. But it also extends up the Missis- sippi and Missouri Rivers to Marion and Cole counties, and it is found in the valleys of some of the other larger streams across the southern part of the state, and northward again along the western side of the state to the Marais des Cygnes bottoms in Bates County. Although usually a species of low or swampy woods, it is found both in northeastern Missouri and in the southwestern part of the state, on high limestone hills where there is an abundant supply of seepage water along the ledges in rainy seasons. Rhamnus carolimiana and Ilex de- cidua, both of which have a general range similar to that of the green haw, also have much the same range as that species in Missouri, although the former does not extend quite so far north. And it is an interesting fact that all of them have [Vor. 22 404 ANNALS OF THE MISSOURI BOTANICAL GARDEN adapted themselves to the same unusual conditions in the lime- stone areas. Forestiera acuminata is another lowland species, although its general range is more southwestern, that in Mis- souri extends from the Southeastern lowlands up the Missis- sippi River to Pike County, and in isolated stations westward along White River and Spring River in Jasper County. Quercus lyrata and Trachelospermum difforme, typical Coastal Plain species, besides being common in the South- eastern lowlands, are found locally in St. Louis County. Ampelopsis arborea extends north to Jefferson and St. Louis counties, and Gleditsia aquatica is also found in the former county. Hottomia inflata has been collected in St. Louis County, and Mitchella repens as far north as Lincoln County. The pubescent form of the sugar maple, Acer saccharum var. Schnecku, which has a well-defined range in the lowlands of the central Mississippi Valley, including southeastern Missouri, has been found in an isolated locality in Benton County. Other southern plants that apparently skip across the state from the Southeastern lowlands to the drainage basin of Spring River in southwestern Missouri are Paspalum floridanum, Scirpus carinatus, Agalimis viridis, Mecardonid acuminata, and Pluchea petiolata. The unusual distribution of some of these plants may pos- sibly be accounted for on the theory of recent extensions of range, but in the case of some of those which are found over such wide areas, as well as of others found in the White River sub-region, it seems more reasonable to believe that they are survivals from a former more general distribution in the region and examples of successful adaptation to changing conditions. The Southeastern lowlands is the most distinet plant region in the state and its flora is the most uniform. But in recent years lumbering, drainage, and cultivation have worked wide- spread destruetion upon the native flora, and it is rapidly losing its distinetive character, while many of the plant species seem doomed to early destruction here. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 405 CHARACTERISTIC PLANTS OF THE SOUTHEASTERN LOWLANDS REGION Woodwardia areolata Wisteria macrostachya Botrychium dissectum var. tenuifolium Sesbania macrocarpa Taxodium distichum Amorpha orooccolanata Sagittaria platyphylla Croton Engelman Echinodorus radicans Crotonopsis nea Panicum hians Ilex opaca Arundinaria gigantea Evonymus americana Cyperus Gatesü Acer floridanum perus compressus Acer rubrum var. Drummondii Cyperus dipsaciformis Aesculus discolor var. mollis Rynchospora macrostachya Sida Elliottü Carex cherokeensis Vitis rotundifolia Carex Joorti itis palma Carex oxylepis ypericum petiolatum Carex louisianica Hypericum virginianum Wolfia papulifera Viola lanceolata Wolfiella floridana Rhexia mariana milax glauca var. leurophylla Jussiaea decu Smilax rotundifolia var. quadrangularis Ludvigia vani ij m TN occidentalis Aralia spinosa ris fulv Eryngium prostratum il ee Cynosciadum digitatum Populus heterophylla Steironema radicans Leitneria floridana Cornus femina Carya aquatica Nyssa aquatica Quercus Prinus Bumelia lycioides Quercus lyrata Styrax americana Quercus nigra Fraxinus profunda Quercus Phellos Frazinus profunda var. Ashei Quercus Nuttallü olypremum procumbens Quercus falcata Trachelospermum difforme Quercus falcata var. pagodaefolia Asclepias perennis Planera aquatica Asclepias variegata Phoradendron flavescens Hydrolea ovata Polygonum densiflorum Hydrolea affinis Brunnichia cirrhosa Lippia nodiflora Cabomba caroliniana Agalinis heterophylla Ranunculus pusillus Veronica serpyllifolia Itea virginica Bignonia capreolata Liquidambar Styraciflua Dianthera ovata Crataegus Marshallü Diodia virginiana Rosa palustris Cephalanthus occidentalis var. pubescens Gleditsia aquatica Oldenlandia uniflora assia Tora Cayaponia grandifolia Cassia occidentalis Lobelia puberula [VoL. 22 406 ANNALS OF THE MISSOURI BOTANICAL GARDEN Mikania scandens Spilanthes americana var. repens Eupatorium incarnatum Gaillardia lutea Eupatorium cuneifolium var. serratum Helianthus angustifolius Solidago suaveolens Pluchea foetida Solidago leptocephala CHARACTERISTIC PLANTS OF CROWLEYS RIDGE IN SOUTHEASTERN LOWLANDS* Erianthus strictus Malus ioensis var. spinosa Carex nigro-marginata Nemophila microcalyz Jorylus cornuta Hydrophyllum canadense Fagus grandif olia Blephilia ciliata Fagus grandifolia var. caroliniana Pycnanthemum incanum — zen Fraxinus biltmoreana Magnolia acumin Catalpa speciosa poseo por m Epifagus virginiana Hydrastis canadensis Mitchella repens Actaea brachypoda Helianthus microcephalus Ribes Cynosbati * It will be noted that several species of this list are of wider range in other regions of the state, but they are practically restricted to this part of the lowlands. THE OZARK REGION Magnesian limestones and sandstones are the surface rocks over a large part of the Ozark region. The strata are usually horizontal or only slightly tilted, although in plaees there are local faults and disturbances. Chert, which is associated with the magnesian limestone in many places, often covers the up- lands as a residual deposit after the less resistant limestone has been removed by erosion or solution. The magnesian lime- stone or dolomite produces an alkaline soil except where an unusual amount of magnesium is present or where the lime has been leached out on exposed slopes. In such places, as well as on the cherty uplands, the soil may be more or less acid. e sandstones, of which the Roubidoux and St. Peter are the most important, outcrop along bluffs and ledges and locally over more level areas. They are composed of rather pure quartz and weather into distinctly acid soils, on which a num- ber of characteristic oxylophytes are found. The granites, syenites, and other igneous rocks of the Iron 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 407 Mountain sub-region also produce acid soils, but on account of the hardness of the rocks they are usually not so pro- nouncedly acid as are those of the sandstone areas. Other local areas of sandstone, with resulting acid soils, are found on the eastern side of the region, where the La Motte sandstone outerops along the east side of the Iron Mountains and the Ste. Genevieve sandstone near the Mississippi River, and also towards the western side where there are numerous small areas of Pennsylvanian sandstone. East of the Iron Mountains and along the Mississippi River local outerops of Silurian, Devonian, and Lower Carboniferous age occur; and limestones of the Mississippian series also underlie considerable areas along the west side of the Ozark dome and extend beyond into the Prairie region. The lime- stones of this series are nearly pure calcium carbonate, and they usually produce fertile alkaline soils. But chert is also present in these beds, sometimes in large quantities, and acid or sub-acid soils are found locally on eroded ridges or slopes. Shales are not abundant in most parts of the Ozark region, but they sometimes occur as separating layers between the strata of dolomite or limestone, and some thicker beds of Devonian or earliest Mississippian age outerop locally on both sides of the plateau. The shales disintegrate into stiff soils or clays, which are generally acid, and on account of poor drainage, dripping banks or barrens, according to the location and slope, may be found in such places. The flora of the Ozark region is more complex and varied than that of either of the other plant regions of the state, and several rather distinct sub-regions and plant colonies can be recognized in it. Considered as a botanical region, the bound- aries both on the north and the west side extend somewhat beyond the Ozark plateau. The richness and diversity of the flora can be attributed to the present topography, drainage, and surface geology, and to the geologie history of the region. The geographical position of the Ozark region on the Ameri- ean continent and its long geologic history both point to the probability of its having been a center of plant development [Vor. 22 408 ANNALS OF THE MISSOURI BOTANICAL GARDEN and distribution from early times, and it is extremely likely that some of the Ozarkian-Alleghenian species originated here and were dispersed from this center instead of the migration having been in the opposite direction, and the same may be true of some of the plants that now range widely to the southwest. Most of the trees that now form the Ozark forests seem to have invaded the region from the south in comparatively recent times, although their ancestors may have occupied it before they were driven out by the cold climate of the Pleiostocene. A few woody species and many herbaceous ones now confined to the Ozark region or found also throughout a wider range only in special habitats often far apart may have first appeared here. As examples of such woody plants may be mentioned Castanea ozarkensis, Hamamelis vernalis, Cladrastis lutea, Andrachne phyllanthoides, and Cotinus americanus. It is in- teresting to note that nearly all of these belong to ancient genera common to both eastern Asia and North America. The scattered distribution of many Ozark plants and the presence in the region of numerous relie species and colonies of plants, some of which are described later, can readily be under- stood when it is r bered that it has been a continuous land surface but with changing boundaries, elevations, climate, and other ecological conditions since late Paleozoie time, and it is not surprising that of the numerous waves of plant life that have passed over it during this time, some of the later ones at least should have left recognizable traces in the present flora. Viewed broadly, the flora of the Ozark region is a forest flora, with the oak-hiekory type of woods prevailing on the uplands, and with mixed or pure stands of yellow pine on some of the higher ridges. The hard-wood trees are usually of small or medium size and in places they are distinctly stunted. The forest in the alluvial valleys is much richer in species and the trees attain a considerably larger size. Only traces of the forest now remain in the valleys, as most of the land has been cleared for agricultural use. The forest, with its associated herbaceous plants, belongs to the great Carolinian flora, and 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 409 floristically it is intermediate between the austral and boreal phases, with a slight preponderance of southern species. The herbaceous plants of general distribution on the uplands are usually species that range from the Appalachian plateau to the grassy plains. In the protection of bluffs along the larger streams and in the deep V-shaped valleys of the more rugged parts of the region, many plants of more northern range are found, and in various places, especially towards the western side of the plateau, there are numerous prairie openings and rocky glades and barrens in which plants of western or southwestern distri- bution are an important or dominant element. In a number of other local areas scattered throughout the region, generally where some unusual ecological condition prevails, small col- onies or single species of plants are found that do not seem to belong to the general flora. The presence of some of these relic species is interesting and significant and may help to throw light on the geological and botanical history of the region. Sink-hole ponds.—In portions of the Ozark region occur what are locally known as ‘‘sink-hole ponds." These are small, shallow, usually circular bodies of water which occupy natural depressions in the level upland plateau, and are best developed in certain of the flatter portions of the Ozark Plateau, partic- ularly in Dent, Shannon, Texas, and Howell counties. The sink-holes become gradually built up with soil and humus until only a shallow depression remains which fills up with water that usually stands throughout the year. Floristically, these natural ponds are extremely interesting. They are gen- erally surrounded by an upland oak-hickory forest of Quercus palustris, Q. coccinea, Q. marilandica, Q. stellata associated with Carya cordiformis, C. alba, C. Buckleyi var. arkansana, C. ovalis var. obcordata, and sometimes other trees. Bushes of Cephalanthus occidentalis, and Hibiscus lasiocarpus often oc- cur, standing in one to three feet of water. The lower crotches of these shrubs may support large tussocks of Carex decom- posita, among which or separately may be growing plants of Viola lanceolata, Lycopus rubellus, Steironema lanceolatum, [Vor. 22 410 ANNALS OF THE MISSOURI BOTANICAL GARDEN and Galium Claytom. Aquatic species in great variety at- tain a luxuriant development in these ponds, among which are Potamogeton pulcher and P. diversifolius, Glyceria acutiflora, Eleocharis acicularis, Spirodela polyrhyza, H eteranthera reni- formis and H. limosa, Brasenia Schreberi, Ranunculus aqua- tilis var. capillaceus, R. pusillus, R. oblongifolius, Callitriche heterophylla, Scirpus americanus, Polygonum hydropiper- oides, Myriophyllum heterophyllum, Sagittaria heterophylla and S. graminea, and Utricularia gibba. Along the pond mar- gins may occur Gratiola lutea and G. virginiana, Acorus Cala- mus, and other species. Certain ponds support distinct species. Glyceria acutiflora and Carex decomposita have been found only in such ponds, while Viola lanceolata is known from only two isolated stations in southeastern Missouri, one of which is in a sink-hole pond. CHARACTERISTIC PLANTS OF GENERAL DISTRIBUTION IN THE OZ ARK REGION Adiantum Capillus-V eneris Crataegus collina Polypodium polypodioides Rosa subserrulata Pteridium latiusculum Lespedeza Stuevei Asplenium Trichomanes Pih intermedia Camptosorus rhizophyllus Lespede a Pinus echinata Cassia nictitans Aristida purpurascens Baptisia vespertina Panicum sphaerocarpon Vicia caroliniana Paspalum circulare Clitoria mariana Commelina virginica Polygala verticillata var. ambigua Quercus Shumardii var. Schneckii Crotonopsis elliptica Quercus velutina var. missourtensis Euphorbia commutata Celtis laevigata var. texana Vaceinium arboreum var. glaucescens Celtis pumila var. georgiana Vaccinium stamineum Silene virginica Vaccinium stamineum var. neglectum Arenaria patula Vaceinium vacillans var. crinitum Ranunculus Harveyi idua Aristolochia tomentosa Rhamnus caroliniana Hepatica americana Vitis rupestris Hamamelis vernalis Ascyrum hypericoides Cocculus carolinus Passiflora lutea Calycocarpum Lyoni Angelica villosa Hydrangea arborescens Eulophus americanus Aruncus sylvester Ligusticum canadense Gillenia stipulata Nyssa sylvatica 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 411 Bumelia lanuginosa Triosteum angustifolium Steironema quadriflorum Silphium Asteriscus Amsonia illustris Parthenium hispidum Vincetoxicum carolinense Eupatorium coelestinum Heliotropium tenellum Brickellia grandiflora Onosmodium subsetosum Solidago Gattingeri Scutellaria incana Solidago hispida Cunila origanoides Solidago petiolaris var. Wardii Monarda. labiis Solidago arguta var. Boottü Aureolaria pectina Aster patens Aureolaria Mice var. cinerea Aster linariifolius Agalinis Skinneriana Aster ptarmicoides Buchnera americana Grindelia lanceolata Ruellia pedunculata Rudbeckia speciosa Galium virgatum Rudbeckia missouriensts Galium arkansanum Helianthus occidentalis Spermacoce glabra Coreopsis lanceolata Houstonia longifolia Coreopsis pubescens onicera flava Hieracium longipilum Viburnum rufidulum Most of the plants on the above list are well distributed throughout the region and a few of them are found occasionally beyond the limits of the Ozarks either in the Southeastern low- lands or in the Prairie region; others are comparatively rare and local but are not confined to any one of the well-defined subdivisions. Many of the commonest trees, shrubs, and her- baceous plants of the Ozark region are found throughout the state and for that reason are not on this list. These include amongst woody plants the red cedar, several species of willows, cottonwood, ironwood, hop hornbeam, most of the oaks and hickories, red and American elm, hackberry, wild cherry, plums, red haws, roses, sassafras, sumachs, maples, lindens, mulberry, persimmon, ash, and many others. The Granitic-siliceous sub-region.—Although the Iron and St. Francois Mountains, with smaller outlying areas of igneous rocks, constitute one of the most distinct sub-divisions of the Ozark region, the boundaries of this floral sub-region must be drawn more widely to include the surrounding sandstone areas, with a belt extending northward to beyond the Missouri River. This may be called the Granitic-siliceous sub-region. While there are many other places in the Ozark region where chert or [Vor. 22 412 ANNALS OF THE MISSOURI BOTANICAL GARDEN sandstone on the surface has given an acid charaeter to the soil, this is by far the largest area and the one in which the influence on the flora of this type of soil can best be seen. The sand- stones, being more friable and therefore disintegrating more rapidly, as well as being more permeable to water, furnish more favorable habitats for many oxylophytes than the gra- nitic or porphyritic areas, and most of the local and characteris- tic species are found on or near sandstone outerops. A rather stunted forest growth covers most of the region, but there are numerous open places in glades or where the bare rock comes to the surface on slopes or where it stands out as domes and peaks in the igneous areas or as bluffs and ledges of sandstone. Pine is found on some of the granite or trachyte hills and on the out- crops of sandstone as far north as Franklin County. The red cedar (Juniperus virginiana) is sometimes found growing on igneous or siliceous rocks, but it is much more frequent in limestone areas. Some of the trees of the Southeastern low- lands, such as Spanish oak, willow oak, black gum, and sweet gum, as well as a few herbaceous species, are found in the extreme southern part, but the dominant and characteristic element over the whole area is Alleghenian-Ozarkian, and a number of eastern plants reach their western limit of distribu- tion here. Ericaceous plants are more abundant than in any part of the state, and the rosy azalea (Rhododendron nudi- florum var. roseum) is a characteristic species in this sub- region. PLANTS CHARACTERISTIC OF THE GRANITIC-SILICEOUS SUB-DIVISION OF THE OZARK REGION Polypodium virginianum Cyperus refractus Asplenium pinnatifidum Carex glaucodea Athyrium acrostichoides Carex hystricina Thelypteris palustris var. pubescens Allium cernuum Thelypteris spinulosa Malaxis unifolia Thelypteris spinulosa var. intermedia Habenaria peramoena Dennstaedtia punctilobula Salix sericea Lycopodium lucidulum irn longifolia Lycopodium lucidulum var. porophilum Cerastium arvense var. oblongifolium Lycopodium complanatum var. flabelli- usn diphylla forme Saxifraga virginiensis 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 413 Saxifraga pensylvanica var. Forbesü Phlox maculata Hamamelis virginiana Trichostema dichotomum Crataegus uniflora Salvia lyrata Euphorbia obtusata Collinsonia canadensis Linum striatum Pedicularis lanceolata Ilez verticillata var. padifolia Aureolaria calycosa Frasera caroliniensis Houstonia patens Rhododendron nudiflorum var. roseum Houstonia coerulea Rhexia virginica Solidago suaveolens Vincetoxicum obliquum Cirsvum virginianum Phlox glaberrima The White River sub-region and the bald knobs.—In de- scribing the drainage and the geology of the state mention has already been made of White River, which makes an incursion into the counties of Barry, Stone, and Taney, along the south- ern border. Another loop of this river also nearly touches the state boundary a little farther east, near the southwest corner of Ozark County, and all of the south-flowing streams as far east as Howell County are tributary to it. The extremely rugged country bordering this river and the lower courses of some of its tributaries comprises one of the most interesting and distinet subdivisions of the Ozark region. The river in its circuitous course has cut a deep narrow valley through the nearly horizontal strata of the Ordovician rocks. Thick solid beds of dolomite, separated by softer layers of shale, underlie most of the area. Hard nodules or masses of chert are sometimes present in the dolomite or are strewn in broken fragments over the surface of the hills, and ledges or local outerops of quartzite and sandstone appear in some of the bluffs and hills. The river bluffs sometimes have a height of 200 or 300 feet, and the stream in places impinges against them or is separated only by a steep rocky talus. On the op- posite side there may be a narrow alluvial or gravelly valley, usually subject to occasional overflows. The uplands a little back from the river are covered with the typical oak-hickory forest of the Ozarks, but along the river bluffs and in the deep cañons of the tributary streams, as well as on the bald knobs, many rare and characteristie plants are found. Along the talus or alluvial strip at the base of the bluff there is usually a dense [Vor. 22 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN growth of trees. The wider parts of the valley that were orig- inally wooded with large trees have nearly all been cleared for eultivation, but floods often work destruction on erops and strip off the surface soil. Where there are irregular ledges or clefts along the face of the cliffs many small trees and shrubs as well as herbaceous plants have found lodgment. The character of the vegetation depends upon the exposure. Where this is to the north or northeast a number of eastern Appalachian species are present. Amongst these are the butternut, north- ern red oak, cucumber-tree (Magnolia acuminata), yellow- wood (Cladrastis lutea), sugar maple, nettle-tree (Celtis oc- cidentalis var. canina), linden, staff-tree, and hydrangea, be- sides a luxuriant growth of ferns and other herbaceous plants. This association of plants is not, however, confined to this sub- region, but most of the species are also found in similar situa- tions throughout the Ozarks, as will be shown later. On south and west exposures the assemblage of plants is quite different, especially on the higher levels of the bluff. Fantastically twisted and gnarled junipers often crown the erest of the cliffs or cling precariously to crevices and ledges along its face. The red cedar (Jumiperus virginiana) is the common species, but in Barry County as well as farther south in Arkansas, another species, called the white cedar by the local inhabitants on aecount of its thick white sap-wood, has been found. In open situations this tree has a different habit from that of the red cedar, with usually several boles or branches arising from the base instead of a single trunk. It is also dis- tinguished by the usually single seed of the fruit and by the minutely serrulate margins of the leaves. This was named Juniperus Asher Buchholz? for the late Mr. W. W. Ashe of the U. S. Forest service, who first called attention to it, but a careful study of this species and a comparison with Juniperus mexicana Sprengel seem to indicate clearly that the two are identieal and that the latter and older name should be applied to the white cedar of the Ozarks. The range of Juniperus mexicana, as given in the second edition of Sargent's “Manual,” * Bot. Gaz. 90: 329. 1930. 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 415 is from Brazos County over the low limestone hills of western and southern Texas and southward into Mexico. It is very abundant on the limestone hills of the Edwards Plateau of central Texas and has recently been collected in the Arbuckle Mountains of Oklahoma, and as a number of other south- western plants of similar range are associated with it in the White River sub-region there seems to be conclusive evidence that this flora at one time extended eastward to the Ozarks. In similar situations is also found the smoke-tree ( Cotinus ameri- canus ), one of the rarest of American trees, and known only from a few other isolated localities in Alabama, Kentucky, Arkansas, Oklahoma, and Texas. It is most abundant and at- tains its largest size here along the White River bluffs, where it sometimes reaches a height of 35 or 40 feet and a trunk diam- eter of over a foot. The soap-berry (Sapindus Drummond) and Acacia angustissima var. hirta, both species of the South- west, are sometimes found in protected situations along the talus at the base of the bluff, and a succulent vine, Cissus incisa, has been found in Taney and McDonald counties, where it grows in hot dry exposures on the face of the bluff. The Mis- souri currant (Ribes odoratum) grows on ledges or in clefts along the bluffs, usually in partial shade. Characteristic also of such places, although of wider distribution in the state, are Celtis pumila var. georgiana, Celtis laevigata var. texana, Bumelia lanuginosa, Fraxinus quadrangulata, Rhus cana- densis var. serotina, and Parthenocissus quinquefolia; and amongst herbaceous plants, selecting situations with various degrees of exposure, are Woodsia obtusa, Pellaea glabella, Asplenium resiliens, Notholaena dealbata, Cheilanthes Feei, and rarely C. alabamensis, Elymus virginicus var. glabriflorus, Uniola latifolia, Commelina erecta var. crispa, M entzelia oligo- sperma, Scutellaria ovata, Aster oblongifolius, and Solidago radula. It is evident that this is mainly a southwestern flora, and it contrasts strikingly with that of the north exposures in which Alleghenian species predominate. But even more interesting and distinct than the flora of the river bluffs is that of the bald knobs, as the eroded hills oc- [Vor. 22 416 ANNALS OF THE MISSOURI BOTANICAL GARDEN eupying the divides between the valleys are locally known. The peculiar topography found here is due to the long-con- tinued action of erosional forces upon the alternating harder and softer horizontal strata, consisting of compact layers of fine, porous dolomite sometimes two or three feet in thickness, separated by bands of shale or clay. The thicker ledges stand out as low bluffs encircling the terraced hills, separated by wide slopes of rather sterile soil which becomes thoroughly satu- rated with seepage water during rainy times and extremely dry later in the season. Asa result of this unequal distribution of available moisture and other factors only a few small trees and shrubs have, at least until recently, been able to establish themselves on the hills, which stand out in striking contrast to the heavily wooded valleys. In parts of the region some of the higher hills are capped with remnants of the purer Missis- sippian limestone, and in such places there is usually a much more abundant woody growth as well as many herbaceous species not found on the dolomite. A few small trees and a variety of shrubs are usually found along the outstanding ledges. Amongst the commonest species are Quercus Shumardü var. Schnecku, Q. Muhlenbergu, Ulmus alata, Celtis laevigata var. texana, Cercis canadensis, Rhamnus carolimiana, Ilex decidua, Diospyros virginiana, Bumelia lanu- ginosa, Fraxinus americana, and the woody twiner Berchemia scandens. On some of the steeper slopes Cotinus americanus is also found, as well as various other trees and shrubs from the surrounding forests. A number of herbaceous plants are also confined to the narrow protected strip at.the foot of the ledges, but most of these are not restricted to the bald knobs but belong to the flora of the surrounding woods. The flora on the open slopes is not rich in the number of species, as most of the plants of the region cannot endure the conditions, and as a result those that are able to do so have a practical monopoly of the area. Some of these are so abundant locally and are so conspicuous when in flower that at such times the hillsides present a most brilliant and striking spectacle. Almost the only trees which flourish on these open slopes are the junipers 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 417 (Juniperus virginiana and more locally J. mexicana, and some- times the former is abundant). Such low shrubs as Rosa caro- lina, Ceanothus ovatus var. pubescens, Acacia angustissima var. hirta, Andrachne phyllanthoides, and a few others may be found here. The Andrachne, locally called buck-brush, is the most abundant and characteristic, and it is interesting both because of its range, which extends through northern and central Arkansas and eastern Oklahoma to central Texas, and because it is the only woody member of the large and generally southern family of the Euphorbiaceae to extend so far north. The herbaceous plants of these slopes are mostly either short-lived annuals or deep-rooted perennials. Amongst the most abundant and conspicuous are the tall larkspur ( Del- phinium Treleasei), an endemic species, the large-flowered evening primrose (Oenothera missourensis, locally known as glade lily), the pink- and the yellow-rayed cone-flowers ( Echi- nacea pallida and E. paradoxa), blue wild indigo (Baptisia vespertina), black-eyed-Susan (Rudbeckia missouriensis), and Hymenopappus scabiosaeus. In certain places the purple- bracted horse-mint (Monarda citriodora) and the large- flowered purple beard-tongue (Pentstemon Cobaea var. purpu- reus) are abundant, and both are very showy when in bloom. A number of other less conspicuous plants are also character- istic of the bald knobs, and many of them are found nowhere else in the state while others occur elsewhere locally in glades. These are included in the fuller list given in a later paragraph. An examination of this list will show that here also, as on the bluffs with south and west exposures, the predominant element in the flora is of southwestern range, but that mingled with such plants are also a number of species that have come from the eastern Appalachian region. In both cases they seem to be relic colonies isolated on these bald knobs and to include many species that have survived through adaptation to the peculiar conditions found here, while they are unable to compete with the more widespread and aggressive plants making up the pres- ent flora of the general region. The significance of such colonies in indicating the climatic changes that have occurred [Vor. 22 418 ANNALS OF THE MISSOURI BOTANICAL GARDEN in the region and in contributing evidence on its geologic his- tory is very great, and it is interesting to see whether we can determine through their aid something of the sequence of events that brought these successive invasions of plants from different directions into the region. In describing the Southeastern lowlands it was stated that the flora now confined to that section in Missouri probably occupied much of the area of the Ozark region before its last elevation, and it was suggested that the distribution of several species now scattered throughout parts of the state beyond this region may be accounted for in this way. Some of the plants found in the White River country and on the bald knobs seem to furnish even more conclusive evidence of this. The small form of the cane ( Arundinaria gigantea), a typical species of the Coastal Plain from Maryland and Florida to eastern Texas and extending with other southern plants into the Mississippi Embayment area of southeastern Missouri, where it grows in low wet woods and alluvial banks of streams and bayous, is also found along White River and its tributaries in southwestern Missouri. In this part of its range it usually grows at the foot of rocky bluffs, often in widely separated stations and under very different conditions from those found in the lowlands. The southern red oak (Quercus falcata) is a tree of wide distribution in the Coastal Plain, and it is a char- acteristic species of the Southeastern lowlands, growing on the low elevations or in the better-drained bottom lands. It is also found along White River as far west as Barry County, Missouri, and Carroll County, Arkansas, where it has taken to the dry rocky hills. The fringe-tree (Chionanthus virginica) has a similar range, although more restricted in Missouri, where it is known locally in the lowlands of Mississippi County and in dry rocky uplands along the North Fork of White River, in Ozark County. Similarly, the herbaceous Hymenopappus scabiosaeus, a common plant of sandy areas in the Coastal Plain and found in the lowlands in Scott and Mississippi counties, reappears as a characteristic species of the glades and bald knobs along White River. Other southern plants with 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 419 a somewhat similar distribution in Missouri are the cucumber tree (Magnolia acuminata) and the leguminous Rhynchosia latifolia. The widely scattered distribution of Forestiera acuminata, Rhamnus caroliniana, and Ilex decidua has already been mentioned. All of them are found along White River, the first growing along the river margins or along sloughs and old channels, and the others along rocky ledges of the bald knobs. Most remarkable of all, however, is the supple-jack or rattan vine (Berchemia scandens), both for its abundance and the way in which it has adapted itself to conditions entirely dif- ferent from those of its usual habitats. In the Coastal Plain, where it is found from Virginia and Florida to eastern Texas and in the Southeastern lowlands of Missouri, it usually grows in swamps or low wet woods, and there it is a slender twiner, winding its tough stems about the trunks of shrubs and trees, which it sometimes ultimately strangles in its strong embrace. On the bald knobs, to which it is confined in the White River sub-region, it generally has a somewhat shrubby habit in the absence of any support, although the slender branches find support on the rocky ledges or on shrubs or small trees where they are at hand. It has also adapted itself to the irregular water supply afforded by the run-off from the hills after rains and the seepage from the rocky ledges that continues for some time afterwards. After this is exhausted the ground often becomes very dry and sterile for a considerable period. In tliis ease, since the plant is not found at all in the alluvial valley or along the bluffs, it is scarcely reasonable to believe that it could have surmounted the present wide barrier of the rocky uplands between the old Mississippi Embayment area and the bald knobs or that it may have been introduced on these hills by some unaccountable circumstance, have survived in so unusual an environment, and have spread from one isolated glade to another. The only reasonable explanation of its pres- ence here seems to be that it is a relie of former general distri- bution through the region prior to its last elevation, and that it is one of the species that, finding itself stranded in the grad- [Vor. 22 420 ANNALS OF THE MISSOURI BOTANICAL GARDEN ually rising Ozark region, succeeded in accommodating itself to the eonditions that have slowly developed in the glades of the bald knobs, where competition with other species has not been too keen. But before considering further the significance of these re- markable relie colonies it may be well to mention others of a somewhat similar character found in other parts of the Ozark region. CHARACTERISTIC PLANTS OF THE BALD KNOBS, SOME OF WHICH ARE ALSO FOUND IN ROCKY LIMESTONE OR DOLOMITE GLADES IN OTHER PARTS OF THE REGION Ophioglossum Engelmanni Evolvulus argenteus Isoetes Butleri Scutellaria Bushii Juniperus virginiana Satureja glabra Juniperus mexicana Monarda citriodora Sporobolus neglectus Amsonia ciliata var. tenuifolia Carex Crawei Pentstemon Cobaea var. purpureus Carex Meadii Heliotropium tenellum Allium stellatum Centaurium texense astylis acuta 4 een scabiosaeus Agave virginica a paradoxa Arenaria patula soups Gat ttingeri Talinum calycinum Aster 8 Leavenworthia uniflora Aster TER Delphinium Treleasei Aster hat Por var. rigidulus Acacia angustissima var. hirta Andrachne phyllanthoides Marshallia peca var. platyphylla erchemia scandens Palafoxia callosum Hypericum pseudomaculatum Rudbeckia missouriensis Oenothera missourensis Coreopsis lanceolata var. villosa Stenosiphon linifolius Glades and barrens of the Ozark region.—Rocky glades and open hillsides upon which ledges of rock come to the surface are found scattered throughout the Ozark region, and many characteristic plants, some of which are remote from the main area of their range, are found in such places. A considerable number of the species found on the bald knobs also grow in some of the rocky glades. Some of the characteristic plants are common to most of the glades, while others are restricted to a few localities or to a single station. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 421 The term glade is used here for small open areas in the forest where rocks outeropping on the surface prevent the encroachment of most trees and shrubs. The name barren has also sometimes been given to such areas, but it should perhaps be restricted to larger areas or to those with a different type of soil and ecology, and in the Ozark region it is used col- loquially for the larger prairie openings which properly are extensions of the Prairie region. Glades are most abundant in limestone and dolomite areas, but they are also frequently found on sandstone outerops or on those of igneous rocks in the Iron Mountain sub-region, and in southwestern Missouri on exposures of chert. The type of vegetation differs somewhat with the nature of the rocky out- erop. Selaginella rupestris, Cheilanthes lanosa, Crotonopsis elliptica, Hypericum gentianoides, and Trichostema dichot- omum are characteristic plants of the igneous or siliceous rocky barrens, and other species of Hypericum and shrubby species of Vaccinium are usually also present. The chert glades of southwestern Missouri are so distinct in their flora from any other part of the Prairie region that they should be treated separately. They are found in the valleys of Turkey Creek and Shoal Creek, in Jasper and Newton counties, and lie beyond the borders of the Ozark plateau. Solid beds of the Grand Falls Chert of the Mississippian series come to the surface in several places along the borders of streams, and in the depressions of the irregular hummocks there is an accumulation of soil which is generally acid but enriched with alluvium. As the rock is impermeable except where fractured, water also fills the depressions after rains and often remains there until evaporated by the sun. This combination has produced unusual ecological conditions, and a number of plants are found here that are known nowhere else in the state, and others that are known only here and in the traehyte or granite glades of the Iron Mountain sub-region. Most of the characteristic species are of southern or south- western distribution. [Vor. 22 422 ANNALS OF THE MISSOURI BOTANICAL GARDEN CHARACTERISTIC PLANTS OF THE CHERT GLADES Cheilanthes lanosa Saxifraga texana Selaginella rupestris Lathyrus pusillus Isoetes melanopoda Crotonopsis elliptica Cyperus inflexus Hypericum pseudomaculatum Scirpus carinatus Opuntia macrorhiza Allium mutabile Opuntia Rafinesquii Rumex hastatulus Chaerophyllum texanum Polygonum tenue Spermolepis echinata Arenaria patula Cynosciadium pinnatum Talinum parviflorum Phacelia hirsuta Talinum calycinum Ruellia caroliniensis var. parviflora Portulaca parvula Specularia leptocarpa Portulaca retusa Coreopsis lanceolata Selenia au Coreopsis tinctoria Sedum N aho SR Krigia occidentalis Limestone and dolomite glades.—The list of plants found in the limestone and dolomite glades is a long one, but many of the species are common also to the rocky woods or prairies. Some are found generally in the larger prairie openings or barrens of the Ozark region or on the uplands of the adjoining Prairie region, and a few of them also grow in glades of sand- stone or igneous areas. The glades often merge gradually into the rocky woods, and shrubs or small trees may encroach and establish themselves in parts of the glade or about its margins. An invasion of herbaceous woodland species soon follows in such places. Many of the species of the following list are nearly always present in glades of this class, while other are comparatively rare and restricted. CHARACTERISTIC PLANTS OF LIMESTONE AND DOLOMITE GLADES Ophioglossum Engelmanni Agave virginica Isoetes Butleri Nemastylis acuta Juniperus virginiana Oxybaphus albidus Sporobolus asper var. pilosus Arenaria patu Sporobolus ozarkanus Portulaca parvula Bouteloua curtipendula Talinum calycinu Carex Crawei Clematis Fremontii Carex Meadii Sedum pulchellum Allium mutabile Delphinium Penardi Allium stellatu Delphinium azureum var. Nortonianum Nothoscordum uds Draba cuneifolia Camassia hyacinthina Erysimum asperum 1935] PALMER & STEYERMARK— PLANTS OF MISSOURI 423 Lesquerella gracilis Polytaenia Nuttallii Lesquerella angustifolia Cogswellia daucifolia Leavenworthia uniflora Phacelia hirsuta Baptisia vespertina Heliotropium tenellum Psoralea esculenta Onosmodium hispidissimum Psoralea tenuiflora Onosmodium subsetonim Petalostemum purpureum Dodecatheon Me Petalostemum albidum Asclepiodora viridis Astragalus distortus Acerates viridiflora tragalus mexicanas Isanthus brachiatus Euphorbia zygophylloides Monarda citriodora Tragia urticaefolia Galium virgatum Rhus canadensis var. serotina Houstonia angustifolia Rhamnus caroliniana Echinacea pallida Opuntia Rafinesquii Solidago Gattingeri Mentzelia oligosperma er sericeus Malvastrum angustum Aster oblongifolius Oenothera missourensis icoides enothera speciosa Thelesperma trifidum Chaerophyllum texanum Silphium terebinthinaceum In the deep narrow valleys and along the bases of the high bluffs with a northern or eastern exposure, and especially in the rich woods formed by the talus, soil accumulations, and humus, many plants of northern and Appalachian range are found. A few of these were mentioned as growing in such situ- ations along White River, and many of them extend through- out the Ozark region and sometimes beyond along the higher stream bluffs in the Prairie region, or they may grow in more open situations farther north. CHARACTERISTIC PLANTS OF NORTH-FACING BLUFFS OF THE OZARK REGION Adiantum pedatum Smilacina racemosa Athyrium asplenioides aac canaliculatum Athyrium angustum var. rubellum Erythronium americanum Athyrium angustifolium rn albidum Cystopteris bulbifera Trillium sessile Cystopteris fragilis Trillium recurvatum Brachyelytrum erectum Cypripedium parviflorum and var. pu- rex Jamesú bescens Carex Sr Sg Juglans cinerea Carez eburn Asarum canadense Arisaema EEE Asarum canadense var. reflexum Uvularia grandiflora Thalictrum dioicum [Vor. 424 ANNALS OF THE MISSOURI BOTANICAL GARDEN Hepatica americana Aquilegia canadensis Isopyrum biternatum Actaea brachypoda Caulophyllum thalictroides Sanguinaria canadensis Corydalis flavula Hydrangea arborescens Ribes Cynosbat olygala Senega v var. latifolia bov Tilia heterophylla var. Michauxii Acer saccharum var. glaucum Dirca palustris e Thaspium trifoliatum var. flavum Hybanthus concolor Hydrophyllum appendiculatum Viburnum molle Solidago caesia Solidago latifolia Eupatorium urticaefolium Staphylea trifolia Ace Senecio obovatus var. rotundus r saccharum Before summarizing this sketch of the Ozark flora, mention must be made of another interesting plant colony where a num- ber of rare species are found, some of which are not known elsewhere in the state. Mr. B. F. Bush, while collecting in Shannon County many years ago, came upon a locality along Jacks Fork of Current River where high bluffs of magnesian limestone, in places capped by beds of Roubidoux sandstone border the stream. A most unusual assemblage of plants was found here, many of them growing on the dolomite, either at the base of the bluff or along ledges, and others growing on the sandstone either along the top of the bluff or along out- crops a little way back from the stream. Mr. Bush visited this locality a number of times, exploring it thoroughly and col- lecting and distributing many plants from it. Amongst species of special interest that have been found near Jam-up Bluff, as the high point along the river is known, are Scirpus plani- folius, Carex Careyana, C. leptalea, C. laxiculmis var. copulata, Zigadenus elegans, Cypripedium Reginae, Spiranthes lucida, Berberis canadensis, Potentilla canadensis var. villosissima, Parnassia grandifolia, Hypericum gymnanthum, Galium boreale var. hyssopifolium, Campanula rotundifolia, Senecio obovatus var. umbratilis, Trautvetteria carolinensis, Artemi- sia campestris var. caudata, Solidago rugosa var. aspera, and 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 425 S. juncea. Growing in the same locality or near by are the following plants, most of which, although known in other local- ities in the state, are found here and seem to be part of the association: Asplenium cryptolepis, Veratrum Woodu, Cypri- pedium candidum, Ribes Cynosbati, Evonymus obovatus, Dirca palustris, Cornus alternifolia, and Solidago caesia. While collecting during 1934 and 1935 in Texas County, the junior author came upon a locality in a deep limestone cañon where a number of these rare species were growing under similar conditions, and the two localities are evidently closely associated phytogeographically. The range of most of these species (excluding those that are southern Appalachian) is in the glaciated areas in northern Missouri or farther north beyond the borders of the state, and their presence in the heart of the Ozark region far beyond the recognized limits of glaciation would clearly indicate a relic colony that has survived from the time when this northern flora extended much farther south than it does at present in Missouri. They have held out in peculiarly protected or favored spots after changing ecological conditions had brought about the gen- eral destruction or migration of the floras of which they were a part and had replaced them generally with others more adapted to the new conditions. It has been pointed out that a number of species from the Coastal Plain flora, which in its full development is now found in Missouri only in the Southeastern lowlands, are isolated in the bald knobs of the White River sub-region and in other parts of the Ozarks; and these plants probably constitute one of the oldest recognizable elements in the present plant life of the region, harking back to conditions that prevailed before its last elevation in late Tertiary time. This does not of course necessarily mean that they have occupied continuously the exact locations where they are now found, but merely that they were never entirely exterminated in the region. Many of the plants of general northeastern and Appalachian distribution now found in the deep cañons and along north- facing bluffs may also have persisted in the region since pre- [Vor. 22 426 ANNALS OF THE MISSOURI BOTANICAL GARDEN glacial times. For the possibility should not be overlooked that some of these species, as well as those of earlier and later periods, may have been developed in the Ozark region and have been dispersed from this center. And it is scarcely likely that extensive immigration could have taken place from the east after the development of the wide flood plains of the Mis- sissippi River, which offers an effective barrier to many up- land species. Finally, the mainly xerophytie plants of southwestern range, which are now found in the glades, on the bald knobs, and along the south-facing bluffs of White River and other streams, and which are the best preserved and most obvious relic col- onies in the region, are probably of still more recent origin, and are remnants of a prairie or plains flora that penetrated into eentral North America during a period of increasing aridity some time subsequent to the final retreat of the glaciers, perhaps within the last few millenia. Relies of this flora are still recognizable as far east as the cedar glades of middle Tennessee and in several adjoining states. It would seem, however, that the climate at this time, while greatly influencing and changing the character of the flora on the uplands, was not sufficiently arid to destroy or drive out all of the northern flora nor even all traces of the old Tertiary forest flora from the protecting valleys in the Ozark region. Conditions at the maxi- mum of this cycle may perhaps have been comparable to those which prevail at present in the Edwards plateau of central Texas, where a number of the peculiar species of the Ozark glades and bald knobs are now found on the limestone hills, and relie colonies of eastern mesophytie species survive in the canons." There is abundant evidence of various kinds that such a period of widespread aridity occurred about the time we have indicated, both in America and in the Old World. That it pro- foundly affected the flora and through it, as well as in other ways, the development and culture of man, groping his way T See e » J. Canyon flora of the Edwards Plateau of Texas. Jour. Arnold Arb. 1: 288. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 427 up through savagery or the first stages of civilization, is prob- able. And so botanical evidence may aid in clarifying or con- firming the theories of the elimatologists and geologists in working out the story of the past. THE PRAIRIE REGION The flora of the Ozark region and that of the Prairie region merge gradually into each other in many places, and the boundary between them is nowhere so clearly drawn as that between the Southeastern lowlands and the former. While the generally open treeless character of the uplands distinguishes and gives name to the region, it is really one of transition from the eastern forest that has now reestablished itself over most of the Ozark plateau, and the western prairies. The alluvial valleys of the larger streams were all occupied by narrow strips of forest until largely cleared off since the settlement of the country. The forests are of similiar type to those along the Ozark streams, although a number of the southern species begin to disappear towards the north and a few northern ones to come in. As in the Ozark region, the larger valleys may have remained wooded through the period of more arid climate. The Prairie region may be separated into two main divi- sions: the Glaciated and the Unglaciated sub-divisions. The first occupies most of the area north of the Missouri River, ex- cept a narrow strip along that stream and along the Mississippi River near their junction, and it also includes small areas to the south of the Missouri, in St. Louis, Saline, Cooper, Lafayette, and Jackson counties. In the small unglaciated areas north of the Missouri River, the flora, as well as the topography along the river bluffs, has more in common with that of the Ozarks, and it has been considered as part of that region. Stratified rocks of Silurian, Devonian, and Lower Carbon- iferous age underlie the glacial drift in several of the eastern counties. These earlier Paleozoic rocks are mostly limestones, sometimes carrying chert, and with more limited areas of shale and sandstone. Westward and throughout most of this sub- region the underlying deposits are the sandstones and shales [Vor. 22 428 ANNALS OF THE MISSOURI BOTANICAL GARDEN of the Pennsylvanian series, with local beds of limestone in a few places. Glacial drift, till, or loess covers most of the up- lands, but the older stratified rocks come to the surface in places where the drift has been removed, and especially along bluffs of streams. Much of the material brought in by the glaciers has been redistributed both as alluvium in the valleys and gravel bars along the streams and as loess in several parts of the sub-region. As the flora found on the loess hills in the northwestern part of the state is so distinct, it will be described in a separate paragraph. There is considerable mechanical as well as chemical differ- ence in the soils of the different parts of the glaciated uplands, and they vary greatly in fertility. However, in general they are more uniform over wide areas than are the residual soils of the Ozark region and of the unglaciated prairies. Judging by the character of the flora, neutral or subalkaline soils are most common. A typical prairie flora, consisting largely of grasses and perennial herbs in which legumes and composites played a conspicuous part, covered the uplands before they were brought under agriculture. Only a few traces of this now remain, the best examples being along the fenced right-of-way of the railroads. Most of the species of this flora are also found in the unglaciated areas farther south. "There are few or no trees along the margins of most of the smaller streams, but sometimes there are a few cottonwoods or willows or small thickets of shrubs alternating with the tall grass and ranker growth of herbs. Amongst common species here, besides the black willow and cottonwood, are Salix cordata, S. missouri- ensis, S. mterior, Prunus americana, P. virginiana, Amorpha fruticosa, Vitis vulpina, Cornus asperifolia, Cephalanthus oc- cidentalis, and Viburnum prunifolium. Of this list only the choke cherry (Prunus virginiana) is practically restricted to the sub-region. In addition to this widely spread prairie flora of the uplands, which eenters in the western plains, and the wooded strips of the alluvial valleys, evident extensions of the eastern and 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 429 southern forest, there is a sprinkling of distinctly northern plants that are generally distributed over the glaciated areas of the middle and eastern states. These plants, although forming a comparatively small percentage of the flora, are the most characteristic species in distinguishing this sub-region. Less detailed study of the flora has been done in this sub-region than in any other part of the state, and it is probable that the small list of characteristic species could be considerably in- creased if systematic collecting were undertaken in all of the counties. The flora of the loess areas, or where the glacial drift has been redistributed, has little to distinguish it from that of the drift areas except on the terraced bluffs and mound-like hills along the Missouri River in the northwestern corner of the state. So many peculiar plants are found in that locality that it must be regarded as a small but distinct sub-region. CHARACTERISTIC PLANTS OF THE GLACIATED SUB-DIVISION F THE PRAIRIE REGION Ulmus Thomasi Acer nigrum Acer Negundo var. violaceum Vitis v Pteretis nodulosa Phalaris arundinacea Phragmites communis var. Berlandieri Calamagrostis canadensis var. Macouni- ulpina ana 4eolepiae Sullivantti Allium tricoccum spermum caroliniense ilaci ellata Phlox pilosa var. pee Collomia linear Chelone bua var. speciosa Fraxinus pen Sambucus racemosa Prunus virginiana Quercus ellipsoidalis Viburnum Lentago Viburnum afine var. hypomalacum Lonicer ica var. glaucescens Senecio pauperculus Prenanthes alba Artemisia vulgaris var. gnaphalodes Antennaria neglecta Several of the species of this list are quite local within the area and others are not strictly confined to the glaciated areas, but they are typical plants of the northern flora that in a few cases have spread somewhat farther south. [Vor. 22 430 ANNALS OF THE MISSOURI BOTANICAL GARDEN The loess is best developed and attains its greatest thickness above the bluffs of the Missouri River, interruptedly, from the northwest corner of the state to below Kansas City. Because of the rapid and charaeteristie way in which erosion has worked upon it, a peeuliar type of topography, with terraced hills or mounds separated by deep, steep-sided ravines, has been developed. The best examples of this topography are found in Atchison and Holt counties, and a number of plants from the northwestern plains are found here and are restricted to this sub-region. The alluvial lands along the river are heavily wooded, and many of the forest species extend up the deeper ravines, but rapid erosion and wind action have kept the slopes bare in most places, although a few woody plants are found at the tops of some of the mounds. Occasionally a few cottonwoods and willows and a stunted form of the bur oak grow in such places or on the slopes where they are not too steep. Grasses and other herbaceous plants constitute the bulk of the flora, and some of the plains species reach their eastern limit here. The bear grass (Yucca glauca) is one of the most characteristic species. Amongst the few other shrubs are the prairie willow (Salix humilis), the lead plant (Amorpha canescens), and the downy- leaved red-root (Ceanothus ovatus var. pubescens). The low oak, Quercus macrocarpa var. olivaeformis, is abundant in places. It is usually a stout spreading shrub, but may become a small tree in protected situations. The fruit is small and elongated and the leaves are extremely variable in shape. The incursion of this northwestern flora into the state may have taken place about the same time that the plants of the southwestern plains overran the Ozark region. Due to the somewhat drier conditions that still prevail here and to the rapid rate of erosion, this flora has not had to encounter the competition of the advancing forest that has pressed upon and almost engulfed the xerophytic plains flora to the south, so that it has remained dominant over the small favorable areas of the loess hills. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 431 CHARACTERISTIC PLANTS OF THE LOESS HILLS Muhlenbergia cuspidata Euphorbia serpens Bouteloua hirsuta Euphorbia marginata Bouteloua gracilis Euphorbia glyptosperma porobolus airoides Oenothera serrulata ucca glauca Gaura coccinea Anemone cylindrica Teucrium occidentale Chenopodium dakoticum Pentstemon grandiflorus Atriplex patula Castilleja sessiliflora Quercus macrocarpa var. olivaeformis | Symphoricarpos occidentalis Cleome serrulata Liatris acidota Rosa polyanthema Heterotheca subazillaris Rosa conjuncta Artemisia vulgaris var. gnaphalodes Oxytropis plattensis Cirsium discolor Astragalus lotiflorus Lygodesmia juncea Psoralea argophylla Lactuca pulchella Dalea enneandra Chrysopsis Berlandieri Glycyrrhiza lepidota Salt licks and saline springs.—Natural saline areas occur in the form of salt ‘‘licks’’ and saline springs, and are found in portions of the prairie region, particularly the glaciated prairie, in areas adjacent to streams and usually in territory of low relief. They reach their greatest development in middle Missouri in Saline, Randolph, Howard, and Cooper counties. The soil immediately around such springs is heavily charged with concentrated alkaline salts, and supports an unusual as- semblage of plants, some of which are known only from such places. Typha angustifolia and Eleocharis parvula have been found in marshy ground around certain saline springs, and are known in Missouri only from these stations. Floating in the water of the springs or branches from them are clumps of Zan- nichellia palustris, Potamogeton pectinatus, Callitriche heter- ophylla, and species of Chara and Nitella. Other species which are found around the saline springs include Scirpus ameri- canus, S. fluviatilis, Typha latifolia, Diplachne fascicularis and D. acuminata, Atriplex rosea and A. patula var. hastata, Polyg- onum aviculare and var. angustissimum, Echinochloa pungens, and many others. The Unglaciated sub-division of the Prairie region.—There is no even approximate natural boundary between the Ozark [Vor. 22 432 ANNALS OF THE MISSOURI BOTANICAL GARDEN region and the unglaciated part of the Prairie region to the west of it as is formed by the Missouri River on the north, and the line of demareation is often very irregular and indefinite, with the Ozark flora dominant on broken rocky ground along streams and the prairie flora occupying the more level and open parts of the uplands over a wide transition zone. This is particularly noticeable in areas of Mississippian limestone and chert, which is the surface formation along most of the border line, with an increasing width towards the south. As the soils of the uplands are all residual, it is easy to account for this rather sharp contrast between the flora growing on different geological formations. West of this belt of limestone and over most of the sub-region the sandstones and shales of the Penn- sylvanian series are the surface rocks. Because of the com- paratively soft character of these formations some of the streams have developed broad shallow valleys, subject to fre- quent overflow, and with numerous small shallow lakes and sloughs in which many aquatie and semi-aquatie plants flourish. These wide flood plains are particularly well developed along the Grand and Marais des Cygnes rivers and some of their small tributaries. Typha latifolia, Leersia virginica, L. oryzoides, Spartina pectinata, Nelumbo pentapetala, Nympho- zanthus advena, Ceratophyllum demersum, Jussiaea diffusa, Myriophyllum scabratum, Proserpinaca palustris var. crebra, and species of Potamogeton and Sagittaria are characteristic of such places. The pin oak (Quercus palustris) is abundant along the smaller streams and in the borders of low woods, and the pecan (Carya illinoensis) in somewhat drier situations. The true prairie flora of the uplands is quite similar to that of the Glaciated sub-division just described, but with an in- creasing proportion of southwestern species towards the south. Other conspicuous differences in the two sub-divisions are the absence of most of the characteristic northern plants to which attention was ealled in the last list, and in the greater variety of trees and shrubs found along the streams. The prairie flora has been better preserved here than in the northern part of the state, due to the fact that large tracts have 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 433 been kept as grazing lands as well as for putting up hay of the native grasses. In the big prairie hay fields of Vernon, Barton, and Jasper counties, as well as along the railways, fine ex- amples of the native flora can be seen. The prairies here are very attractive at different seasons, with a profusion of flower- ing plants, many of them of brilliant colors. With the first warm days of spring little drabas, March lily (Erythronium mesochoreum), spring beauty, prairie anemone (Anemone caroliniana), buttercups, violets, and bluets (Houstonia min- ima) begin to appear; and these are followed quickly by wild hyacinth (Camassia hyacinthina), spiderwort (Tradescantia canaliculata), wild roses, blue-, yellow-, and white-flowered wild indigo (Baptisia vespertina, B. leucophaea, and B. leucan- tha), bush clover, prairie clover (Petalostemum purpureum and P. candidum), wild senna, shooting star, phlox, Indian blanket, larkspur (Delphinium Penardi), gromwell (Lithosper- mum canescens and L. angustifolium), and coreopsis (Coreop- sis grandiflora). Later come butterfly-weed, milk-weed, horse- mint, gerardias, wine cups (Callirhoë digitata), blazing star (Liatris scariosa and L. pycnostachya), ironweeds, rosin- weeds, and many species of asters and solidagos. On the Barton upland, in the southwestern part of Barton County and the northwestern part of Jasper County, a number of peculiar plants have been found, some of which are known nowhere else in the state. The upland prairies here have a sandy soil, with blocks of residual sandstone scattered over the surface and ledges sometimes outcropping, producing little glades or barrens; and along the small intermittent streams are small boggy areas with an acid type of soil. Some of the plants found here are Andropogon ternarius, Setaria gemic- ulata, Panicum scoparium, Rynchospora capitellata, R. cy- mosa, Scleria ciliata, Scirpus carinatus, Carex arkansana, Juncus interior, J. effusus var. solutus, J. validus, J. nodatus, Geocarpon minimum, Anemone caroliniana, Ranunculus ob- longifolius, Crotonopsis elliptica, Saxifraga texana, Rhexia interior, Marshallia caespitosa, and Chrysopsis pilosa. [Vor. 22 434 ANNALS OF THE MISSOURI BOTANICAL GARDEN OHARACTERISTIC PLANTS OF THE UNGLACIATED SUB-DIVISION OF THE PRAIRIE REGION Sagittaria ambigua Rubus Bushii Andropogon saccharoides Ammannia auriculata Sporobolus ozarkanus Lotus americanus dotati dactyloides Rotala ramosior var. interior icum scoparium Rhexia interior Schedonnardus paniculatus Asclepias syriaca var. kansana Sabatia campestris Fimbristylis autumnalis Lobelia spicata var. hirtella Carex oklahomensis Agalinis fasciculata Carex Bicknellii Gutierrezia dracunculoides Juncus validus Solidago gymnospermoides Erythronium mesochoreum Helianthus salicifolius Sisyrinchium campestre var. kansanum Artemisia vulgaris var. ludoviciana Anemone caroliniana Marshallia caespitosa Draba reptans Aster paludosus Many of the most abundant and striking plants of the un- glaciated prairies are of wide distribution throughout the state and therefore cannot be included amongst the distinctive plants of the sub-division, although, since they often dominate the flora and give character to it, they are in a sense characteristic. Most conspicuous amongst these may be mentioned Andropo- gon scoparius var. frequens, A. provincialis, Sorghastrum nutans, Tradescantia canaliculata, Camassia hyacinthina, Ra- nunculus fascicularis, Delphinium Penardi, Rosa setigera var. tomentosa, Desmanthus illinoensis, Schrankia uncinata, Cassia fasciculata, Baptista leucophaea, B. leucantha, Petalostemum purpureum, P. candidum, Astragalus mexicanus, Amorpha canescens, Desmodium illinoense, D. canadense, D. sessilifo- lium, Lespedeza capitata, Linum medium, Polygala incarnata, Croton capitatus, Euphorbia corollata, Callirhoé digitata, Viola sagittata, Jussiaea diffusa, Gaura biennis, Oenothera biennis, Eryngium yuccifolium, Ptilimnium Nuttallii, Ci- cuta maculata, Polytaenia Nuttallii, Apocynum cannabinum, Asclepiodora viridis, Asclepias tuberosa, A. incarnata, A. verticillata, Acerates viridiflora, A. hirtella, Phlox pilosa, Lithospermum canescens, L. angustifolium, Verbena stricta, V. canadensis, Physostegia virginiana, Monarda mollis, Cas- tilleja coccinea, Pentstemon tubiflorus, P. Digitalis, Houstonia 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 435 minima, Vernonia crinita, V. Baldwini, V. missurica, Liatris scariosa, L. pycnostachya, Solidago glaberrima, S. speciosa, S. altissima, Aster praealtus, A. ericoides var. prostratus, Antennaria plantagimif olia, Silphium laciniatum, S. integri- folium, Rudbeckia subtomentosa, Echinacea pallida, Ratibida pinnata, Helianthus rigidus, H. mollis, H. grosse serratus, H. hirsutus, Coreopsis tripteris, C. palmata, and Cacalia tuberosa. Besides the main plant regions and their sub-divisions and the isolated colonies of relic plants which have been described, a number of species have been found only in a single locality or in a few widely separated localities in the state, and several others seem to be confined to a single geological formation. Thelypteris spinulosa, Saxifraga pensylvanica var. Forbesu, and Sullivantia renifolia have been found only on outerops of St. Peter sandstone, near the Mississippi and Missouri Rivers from Jefferson to Montgomery counties. Somewhat farther south in the Iron Mountain area, Dennstaedtia punc- tilobula, Lycopodium complanatum var. flabelliforme, Good- yera pubescens, and Viola pallens are known only very locally on the La Motte sandstone. Pedicularis lanceolata, a species which is usually known only from much farther north than Mis- souri, west of the Mississippi, has been found in acid bogs in Iron, Shannon, Morgan, and Greene counties. In the central part of the Ozark region Silene Wherryi, a plant of the southern Appalachians, has been collected in Phelps, Pulaski, Dent, Douglas, Shannon, Texas, and Laclede counties, where it grows in cherty woods. Zigadenus Nuttallii and Eriogonum longifolium have been found only in Oregon County. In the limestone glades of Jackson and Cass counties Cogswellia daucifolia is locally abundant, although it is known from no other locality in the state. The general range of this species is westward through Nebraska to Texas, and it is a curious fact that Clematis Fremontit, with a similar general range, has been found only in limestone glades on the opposite side of the state. Geocarpon minimum, a very interesting little plant both because of its rarity and also because it seems to be in some respects a connecting link between the plant families [Vor. 22 436 ANNALS OF THE MISSOURI BOTANICAL GARDEN Aizoaceae and Caryophyllaceae, was discovered in sandstone glades in Jasper County, and it is so far known from no other loeality. Epipactis latifolia has been found in the same county, and this is apparently the only record for this species west of Pennsylvania. There seems to be no doubt that it is native at this station. Amsonia ciliata var. tenuifolia is growing abun- dantly in glades of the bald knob type in Ozark and Douglas counties, and these are also the only known localities for it in the state. Mention has already been made of a number of other similar instances in describing the Jam-up Bluff loeality of Shannon County and the glades and bald knobs of other parts of the state, and the list of such species could probably be considerably lengthened. Several distinctly southern or coastal species of the South- eastern lowlands again appear in the extreme southwestern part of the state. Amongst these are Scirpus carinatus, Paspalum floridanum, Triodia stricta, Leersia lenticularis, Me- lothria pendula, Mecardonia acuminata, and Agalinis viridis. Some of these wide gaps in distribution may be filled in by future discoveries, and in some cases at least they would seem to indicate a former wider distribution of the species in the intervening areas. More than 10 per cent of the plants included in the present catalogue have been introduced into the state since the settle- ment of the country by Europeans. Most of these are weeds or grasses that grow in waste places or in cultivated ground. Many of them have become thoroughly established and com- mon, and in some cases they are crowding out to a great extent the native species which find themselves unable to compete with them. Some of our introduced plants were formerly cultivated for their supposed medicinal value or for other uses, and others were first grown as ornamental plants in old gardens. The day-lily, blackberry lily, Queen Anne’s lace, hoarhound, catnip, motherwort, matrimony-vine, Gill-over- the-ground, butter-and-eggs, bouncing Bet, and teasel are familiar examples of these. Most of the introduced plants are very adaptable and con- 1935] PALMER & STEYERMARK— PLANTS OF MISSOURI O + Rain OT NS ions 437 r falcata ] E 4 Nn! [m a X Smilax glauca var. leurophylla 48 | SR X a |^ L. e F—— —4 Oxytropis plattensis ENSIS: el Fig. 3. in the Whit the So ar lowlan four four maps). Since the cine were made, Bar Showing occupation of three species in the ee nr and e and Current ay er systems (first three maps) ; bo in the b. M (lower inaria gigantea has been found in Pemiscot, Dunklin, Barry, and ebene ar and Quercus falcata in rry Co. (Vor. 22 438 ANNALS OF THE MISSOURI BOTANICAL GARDEN Pteretis nodulosa Calamagrostis canadensis Macouniana w N N e = da \ r 1 Populus tremuloide Rosa blanda x x = = = — ) "m. 1 Prunus virginíana Artemisia vulgaris vår. gnaphalodes L [^L [1/1 Anemone caroliniana Agalinis fasciculata XT pa Fia hs Fig. 4. Showing occupation of species in the glaciated prairie region of north- ern and central Missouri (upper six maps); of two species in the unglaciated region (lowest two maps). 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 439 Taxodium distichum halia dealbata C m] LRA im & EN DEOS, AAT TEN PESTA ERES Leitneria floridara^Liquidambar Styraci Ada X R y : Nyssa aquatica Asclepias perenni j 24 ie — 4 Quercus lyrata Porestiera acuminata a ATI | XXI 7 LL TU Peer” 1) Fig. 5. Showing occupation of species in the Southeastern lowlands (upper six maps); species extending outside the limits of this region (lowest two maps). [VoL. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN 440 Pinus echinata Pteridium latiusculum ETA, 3 e D LX. LX PU 178 X) NS Meet: var. eN Schneckii e o #22 E Tox o x = E R po AX g £ [^] ri u oi — o M mn a > o 26 364 75. ~ 7972 z è: A H e y XE 3 x IV al E 1 Fe gi q aA E S 4 k^ S = iN ri T u cala >, Y Pr E Cunila origanoides Vaccinium vacillans var. ab. f LA IA “ay, F m Se E A X) E mab x! X T s = PN a) $ x XX A, E 1% A x DX ST] LX | T de CIO [X | E é M «Es a L eX i. [X 106098 e FRI X TX t lr p PR Dx AL Ire ER L^ DEO p Y ACD e sube 55 s LL DI HERE ai E TOUS Fig 6. Distribution of eight species oceupying the Ozark region. 1935] PALMER & STEYERMARK——PLANTS OF MISSOURI 441 sequently they are not restricted to any type of soil, and their distribution has little signifieance or relation to the native flora. Ina few cases, however, they seem to be limited to cer- tain areas or environments and to show little disposition to spread beyond them. Amongst such plants are the Japanese clover (Lespedeza striata), common only in rocky ground in the Ozark region although sometimes found elsewhere, Helio- tropium indicum and Perilla frutescens that seem inclined to spread only in the southern counties, the teasel, which seems to establish itself mainly along rocky stream beds in the eastern and southern counties, and the recently introduced Tamarix gallica, which, judging by its behavior in other sections, is likely to spread along the margins of the great rivers but scarcely beyond them. Southern species, such as Arundo Donax and the water hyacinth (Eichhornia crassipes ), if they persist will probably be confined to the Southeastern lowlands. The number and relative proportion of introduced plants tend constantly to increase as the country becomes more thickly settled and as means of transportation and communica- tion multiply. Many of them have been brought in by rail- roads, either in ballast or merchandise or in the cleanings of cars, and new immigrants are often first noticed along railways or about railroad centers. Although some of them show little tendency to spread far beyond the place of introduction, others extend their range rapidly and in a few years cover large sections or all parts of the state. The rapid spread in recent years of the white- and yellow-flowered sweet clover and of Torilis Anthriscus and Helenium tenuifolium is a good ex- ample of this. The great majority of our weed immigrants are of European origin; others came originally from Asia, and a few from South America or other parts of the world. Some weedy species, and others that would scarcely be called weeds, have been introduced into the state from the West or from other parts of North America, and in some cases species that are native in one part of the state have been introduced into other sections where they are now established. This is sometimes [Vor. 22 442 ANNALS OF THE MISSOURI BOTANICAL GARDEN confusing in considering the natural distribution of species and its significance, and attention is called to some of these in the notes accompanying the catalogue. Species which are found native in some part of the United States are printed in bold-face type, while the introduced species (1. e., native of some country other than the United States) appear in Roman capitals. The maps and descriptions of the different plant regions of the state, with the lists of characteristic plants found in them, will serve to give some idea of the diversity and richness of the flora of Missouri and of the varied elements that enter into it. BIBLIOGRAPHY OF PHANEROGAMIC Botany or Missouri* Adams, G. I. (01). en and geology of the Ozark region. U. S. Geol. Surv. Ann, Rept. 22°: 1901 Aellen, P. (’29). Beitrag zur raniti der Chenopodium Arten-Amerikas. Fedde’s Rep. Sp. Nov. Veg. 26: 119-16 29. Allen, T. (’76). Missouri, its history, dirseteristios resources, and present con- ditions. 29 pp. h Anderson, 5 (729). Variation in Aster anomalus. Ann. Mo. Bot. Gard. 16: 129— i: ’31). The chromosome complements of Allium stellatum and Nothos- iin bivalve. Ibid. 18: 465-468. 1931 (733). Variation in flower color in Ramaneti vernalis. Jour. Arn. Arb. 14: 253-257. 1933. ehl, D. = Ki Contributions to the Tradescantia problem. Anonymous, (’95a). n nn thistle. Independence Sentinel. Jan. 4, 1895. ——, (’95b). The Russian thistle. Jackson Co. Tribune. June 29, 1895; also tagen Sentinel. June 24, 1895, ——— A orge W. Letterman. Mo. Bot. Gard. Bull. 1: 102-103. 1913. — — — , (?20a). Collecting native plants for the Garden, nk 8: 33-35. 1920 , (20b). Native plants suitable for gardens of Missouri and adjoining , (721). Unusual late-flowering period of plants. Ibid. 9: 126-128. 1921. ————, (722a). Medicinal plants of our local flora. Ibid. 118-123, 137-142, 153-157. 1922. , (722b). Plant immigrants of our local flora. Ibid. 48-52, 84-92. 1922. *Although this bibliography is intended to be as complete as possible, there are probably additional titles which have not been brought to the authors” attention. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 443 , (722e). Plants of our local flora which have been used by man as food. Ibid. 102-110. 1922 ————, (’22d). Plants of very limited local distribution. Ibid. 166-170. 1922. ———, ('22e). Vanished plants of our local flora. Ibid. 10: 28-33. 1922. ————, (’22f). White red-bud. Ibid. 110. 1922 , (723). The hawthorn—the official state flower. Ibid. 11: 45-55. 1923. , (724). Local oaks and their uses. Ibid. 12: 25-32. 19 , (725). List of Missouri wild flowers according to time of blooming. . 1925 , (7262). The conifers of Missouri. Ibid. 14: 25-34. 1926. , (726b). The river birch. Ibid. 67-69. 1926. , (26e). Spring flowers in winter. Ibid. 155-157. 1926. , (727a). The earliest St. Louisian of botanical note. Ibid. 15: 147-152, , (727b). The landscape of St. Louis in 1727. Ibid. 96-102. 1927. , (270). Two botanists of frontier days. Ibid. 42-54. 1927. 28a). Check-list of the ferns and flowering plants indigenous to the Mwoa Botanical Garden Extension, Gray Summit, Missouri. Ibid 1928 74, 83-91. : , (728b). A notable native plant (Sabatia angularis). Ibid. 93-94. 1928. E N i Early spring flowers in Missouri pasture lands. Ibid. 17: 41- 51. ; lb A winter-flowering shrub native to Missouri. Ibid. 30-32. 1929, , (730a). Records M first blooming of wild flowers in the vicinity of Saint Louis. Ibid. 18: 82—84. 0. , (732). Native ud in flower at the Missouri Botanieal Garden Arbore- tum, Gray Summit, Mo., May 10, 1933. Ibid. 21: 91-92. 1933. Ashe, W. W. (700a). Contributions from my herbarium. Bull. N. C. Agr. Exp. Sta. 175: 110. 1900. , (700b). Ibid. VI. New East American species of Crataegus. Jour. Elisha pio Sei. Soe. 16: 72-75. 1900. , (700e). Ibid. X. Some East American species of Crataegus. Ibid. 17: 12. : — — ——, (01). Ibid. XI. Ibid. 17°: 8-9. 1901. , (703). New North American thorns, Ibid. 19: 16. 1903. , (704). East American thorns. Ibid. 20: 1. 1904. , (724). Notes on woody plants. Ibid. 40: 43-48. 1924. , (725a). Notes on woody plants. a 41: 269. 1925. , (725b). Further notes on woody plants. Torreya 25: 10-11. 1925. , (731). Notes on Vacciniaceae. 2 33: 193-198. 1931. Bailey, L. H. (32). North American blackberries. Gentes Herbarum 2: 271-423. figs. 147-182. 1932. Barnes, c. R. (77). Notes on the flora of Clinton Co., Missouri. Bot. Gaz. 2: 138- 139. 1877 Beck, L. C. (1826-28). Contributions nos the botany of the states of Illinois and Missouri. Am. Jour. Sci, 10: 257-264. 1826; 11: 167-182. 1826; 14: 112-121. 1828. [Vor. 22 444 ANNALS OF THE MISSOURI BOTANICAL GARDEN Beilmann, A. P. (732). Big trees in Missouri, Mo. Bot. Gard. Bull. 20: 149-151. Benke, H. C. (’28). Two new varieties of early spring plants. Rhodora 30: 200- 201. 1928 — — —, (732). Earliest spring flowering plants. Am. Midl. Nat. 13: 89-109. 1932. Berry, E. W. (’11). Notes on the ancestry of the bald cypress, Plant World 14: 9—45. 1911. ( Notes on the geologieal history of the walnuts and hickories. —, (12a). Ibid. 15: 225-240, — — ——, (12b). Some ehe of the persimmon. Ibid. 15-21. 1912. , (720). The geological history of the sweet gum and witch hazel, Ibid. : 345-354, 1920 Blake, 8. F. (718). On some species of Viburnum. Rhodora 20: 14. 1918. Bourne, A. (1820). Prairies and barrens of the West. Am. Jour. Sei. 2: 30-34. 1820 Boynton, C. L., and Beadle, C. D. ('02). Notes on certain cone flowers. Biltmore Bot. Stud. 1: 11-18. 1902 Braekenridge, H. M. (1814). Views of Louisiana, Pittsburgh, 1814. , (1816). Journal of a voyage up the Missouri River in 1811. 2nd ed. Baltimore, 1816. [In Thwaites, Early Western Travels 6: 23-166. 1904]. Bradbury, J. (1815). A description of the minerals and plants found at the lead mines in the Missouri territory. New York, 1815. [In Medical Repository of — Nee and Intelligence, etc. N. S. 3: 135-138, 1817]. 819). Travels in the interior of America in - yn 1809-11. 2nd ed. London, n. [In Thwaites, Early Western Travels 5: 9—320. 1904]. Bratton, S. T. (726). The geography of the St. Francis u Univ. Mo. Stud. 1: 1-54. 1926. i Broadhead, G. C. Manuscript list of Missouri plants. , (’67a). Botany and geology of Cass County. Mo. State Bd. Agr. Rept. 2: 226-229. 1807. ; (767b). Distribution of trees and shrubs in Missouri. Ibid. 97-99. ————7, (74). Geol. Surv. Mo. Rept. 1873-1874: 734 pp. Jefferson City, 1874. ————-, (76). Quercus heterophylla, Bot. Bull. 1: 9-10. 7 ; (7782). On the distribution of certain plants in Missouri. Bot. Gaz. 3: 51-53, 58-61. 1878 ; (778b). The range of the common huckleberry in Missouri. Ibid. 24. 79. , (80). Melanthium nian Bot. Gaz. 5: 232. 1880. , (84). Relations of soils of Missouri to geology. Mo. State Bd. Agr. Ann. 2 j^ 159-168. 1884. — . B., and pagi E F. (773). Geol. Surv. Mo. Rept. 1855- 1871: T vn. Jefferson City, 1 Buchholz, J. T. ('30). The Ozark zu cedar, Bot. Gaz. 90: 326-332, figs. 1—2. 30 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 445 Bush, B. F. (782). Notes from Independence, Missouri. Ibid. 7: 24. 1882. —————, (^84). Missouri Notes. Ibid. 9: 63. 1884. , (85). Flora of Jackson Co. 16 pp. Independence, 1882; 1st suppl. 8 pp. 1885 , (789). Report on the botany of Jackson County. Mo. Hort. Soc. Rept. [1888]. 31: 370-372. 1889. ————, (791). Distribution of the trees, shrubs, and vines of J ackson County, Mont Kansas City Seientist 5: 161—164. 1891 , (794). Notes on a list of ghe collected in southeastern Missouri in 1893. Ann. Rept. Mo. Bot. Gard. 5: 139—153. 1894. , (795a). Hybrid oaks in western Missouri. Garden & Forest 8: 32-33. 1895. , (795b). A list of the trees, shrubs, and vines of Missouri. Rept. Mo. Hort. Soc. [1894]. 37: 353-393. 1895. , (795e). Notes on the mound flora of Atchison County, Missouri. Ann. 1895. a ). Quercus Phellos x callis in Missouri. Garden & Forest 8: 379. b. , (797). Notes on the botany of some southern swamps. Garden and Mide 10: 514-516. (700). In Davis, K. D, hes and garden delphiniums of North Amer- i 1900. y 3). A new genus E grasses. Trans. Acad. Sci. St. Louis 13: 175- 1903 , (705). The American species of Fuirena. Ann. Rept. Mo. Bot. Gard. 16: 87-99. 1905. , (709). The Missouri saxifrages. Ann. Rept. Mo. Bot. Gard. 20: 138- 09 , 11). The Missouri rhexias. Rhodora 13: 166-168. 1911. , (713). Notes on box elders. Am. Midl. Nat. 3: 148-150. 1913. , (714). A new Antennaria. Ibid. 3: 352-353. 19 , (716). The Missouri agrimonies. Ann. Mo. Bot. died 3: 309-318. , (718). The genus Euthamia in Missouri. Am. Midl. Nat. 5: 157-177. 8 x re The Missouri muhlenbergias. Ibid. 6: 17-28, 33-49, 57-77, 81- 97. 1919. , (726a). The Missouri species of Elymus, Ibid. 10: 49- 88. 1926. ————, (’26b). Notes on trees and shrubs of Missouri. Ibid. 133- 158. 1926. , (27a). The glabrate species of Tilia, Bull. Torr. Bot. Club 54: 231- 248. 1927. , (27b). A great blunder. Am. Midl. Nat. 10: 400—403. 1927. id. 2 1 . 38 , (28a). The Missouri artemisias. Ibid. 11: 25-39. - , (728b). Notes on trees and shrubs of Missouri. Ibid. 101—124. 1928. , (728e). Some species of Saxifraga. Ibid. 213-235. 1928. , (730). Notes on Aesculus species. Ibid. 12: 19-26. 1930, [VoL. 22 446 ANNALS OF THE MISSOURI BOTANICAL GARDEN ———, ('3la). Francis Daniels’ Carices. Ibid. 185-194. 1931. , (31b). Francis Daniels’ grasses, Ibid. 343-362. 1931. , (31e). Shepard's Greene County Flora—An unpublished list of plants of Greene County, Missouri, by Prof, Edward M. Shepard, Springfield, Missouri. Ibid. 488—498. 1931 — — ——, (31d). Some Lacinaria species. Ibid. 312-318. 1931. — — ——, (31e). The sugar maples. Ibid. 499—503. 1931. — — ——, (34a). More about Acer saccharum Marshall. Ibid. 15: 784. 1934. , ('34b). Prof. Shepard's trees of Greene County. Ibid. 581-585. 1934. — — —, (735a). Francis Daniels’ Flora of Columbia, Mo. Ibid. 16: 83-93. 1935. ————, (35b). The identity of Prunus lanata M. & B. Ibid. 254. 1935. Call, R. E. ('89). Notes on the forest trees of Crowleys’ Ridge region. Ark. Geol. Surv. Ann. Rept. 2: 183-202. 1889, Campbell, R. A. (75). Gazetteer of Missouri. 794 pp. St. Louis, 1875. Canby, W. M. ('85). An pegar ip pen some reminiscences of the late August endler. I. Bot. Gaz. 10: 285-2 Pis S. M. ('04). An ecological cota. x some typieal swamp areas. Ann. Rept. Mo. Bot. Gard. 15: 39. 4 Cutak, L. (33). Prickly pear in fruit. Mo. Bot. Gard. Bull. 21: 141. 1933. , (734). Talinum calycinum. Desert Life 6: 52-53. 1934 , (7342). Talinums—lovely gems for our rock gardens. Garden Gossip 1934 9: 3, » Daniels, F. (98). A preliminary oecological study of the native and introduced plants of a werd E — Boone Co., Mo. Mo. Agr. Exp. Sta. Ann. Rept. 1898: 194—156. 1898. A CN. po Msc in the vicinity of Columbia, Mo. Science N. S. 19: 170. 1904. — ——, (07). Flora of Columbia, Missouri, and vicinity. Univ. Mo. Stud. Sci. Ber. 1: 1-319. 1907. Deane, W., and Robinson, B. L. ('96). A new Viburnum from Missouri. Bot. Gaz. 22: 166- 167. pl. 8. 1896. Drouet, F. (733). Notes on the flora of Columbia, Missouri. Rhodora 35: 359- 933. — ———, (735). Additional notes on the flora of Columbia, Missouri. Ibid. 37: 189-196. 1935 , and Jeffrey, L. (734). Grass flora of Columbia, Mo. Ibid. 36: 415-417. 1934. Duke, W. W., and Durham, O. C. ('24). A botanie survey of Kansas City, Mo., and neighboring rural distriets (with reference to the flora responsible for hay- fever, asthma, and dermatoses). Jour. Am. Med. Assoc. 82: 939-944. 1924. Dunlap, F. ('21). Growth of oak in the Ozarks. Mo. Agr. Exp. Sta. Res. Bull. 41: 6 , (739). List of Missouri trees. Mo. State Bd. Agr. Bull. 27: 1-15. Eggert, H., (791). Catalogue of the phaenogamous and vascular eryptogamous plants in the vicinity of St. Louis, Missouri. 16 pp. 1891. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 447 Elmore, C. J. (26). Manual of the spring flowering plants of Liberty, Missouri, and vicinity. 116 pp. 1926 Emerson, T. V. (712). dudrdphy of Missouri. Univ. Mo. Stud. Educ. Ser. 1*: 1- Engelmann, G. (44). Catalogue of a collection of plants in Illinois and Missouri, by Charles A. Geyer, with critical remarks, ete. Am. Jour. Sei. 46: 94-104. 1844. 48). Sketch of the botany of Dr. A. Wislizenus's expedition from Mis- souri ide Santa Fe, Chihuahua, Parras, Saltillo, Monterey, and Matamoras. Washington, 1848. , (759). Systematic arrangement of species of the genus Cuscuta. Trans. Acad. Sei. St, Louis 1: 453-523. 1859. ; (760a). Notes on the grape-vines of Missouri. Ibid. 660-662. 1860. —— — ——, (760b). Remarks on Nelumbium luteum. Ibid. 2: 136-139. 1860. , (761). On dimorphism of Draba brachycarpa. Ibid. 154-155. 1861. ; (765). On the fruit and seed of different species of Viburnum. Ibid. , (780a). pe speciosa. Bot. Gaz. 5: 1-3. 1880. , (80b). Fraxinus quadrangulata. Ibid. 63. 1880. ————, (83). Vitis palmata Vahl. Ibid. 8: 254-255. 1883. Eeieclnens Botanical Club, Check-list Committee of (711). Preliminary check-list of the eryptogams and phanerogams in the vicinity of St. Louis, Missouri. 63 p 11 Erlanson, E. W. (728). Ten new American species and varieties of Rosa. Rhodora 30: 109-121. 1928, Fenneman, N. M. dida Physiography of the St. Louis area. Bull. Ill. Geol. Surv. 12: 83. 190 Flint, T. (1832). Er and geography of the Missouri Valley. 1-2. Cineinnati, nd ed. en Ss p (785). Grasses of Missouri. Mo. State Bd. Agr. Ann. Rept. 18: 3. 1885. edie N. M. (’94a). A study of the relations of Salix nigra and Salix amygda- loides, together with the hybrids arising from them as these species exhibit them- selves in the vicinity of St. Louis. Trans. Acad. Sci. St. Louis 6: 427-431. 1894 ‚(’94b). A study of the venation of the species E Salix IT. in Gray's Manual, with reference to their determination. . Rept. Mo. Gard. 5: 46—60. 1894. , (795). Salix Wardii. Science N. S. 2: 582-584. 1895. , (7962). Relations of Salix missouriensis to S. cordata. Trans. Acad. Bei. St. Louis 7: 137-144. 1896. ; (796b). Salix cordata x sericea. Bot. Gaz. 22: 392—400. 1896. , (98). Notes on Salix longipes Shuttlw. and its relating to S. nigra. n. Rept. Mo. Bot. Gard. 9: 43-51. 1898. Harris, J. A. (709). Correlation in the inflorescence of Celastrus scandens. Ann. Rept. Mo. Bot. Gard. 20: 116-122. 1909. 24 Viriato and correlation in the inflorescence of Manfreda vir- ginica, Ann. Mo. Bot. Gard. 11: 411—459. 1924 [Vor. 22 448 ANNALS OF THE MISSOURI BOTANICAL GARDEN Harvey, F. L. (80). Southwestern plants. Bot. Gaz. 5: 84. 1880, '81). Leavenworthia in 8. W. Missouri and N. W. Arkansas. Ibid. 6: 1881. Hilgard, E. W. (’84). General features of the alluvial plain of the wen River below the mouth of the Ohio. Rept. U. S. Bur. Census 10: pt. Cot n i . S. 1884, Hooker, W. J. (1834-1836). Notice concerning Mr. Drummond's p pa in the southern and western parts of the United States. Hook. Jou 50—60, 183-202. ony P MA to Bot. Mag. 1: 21-26, 39-49, 05-101, 170- à ; 2: 60- Horsford, F. H. fot sae — the Missouri Botanieal Garden. Forest 4: 225. 1891. , (791b). Notes from the St. Louis Botanical Gardens. Ibid. 284. 1891. ————, (91e). ‘Some early native flowers. Ibid. rid 1891. — — ——, (91d). Some native plants. Ibid. 356. ——, (91e). Wild flowers around St. Louis. e 260— 261. 1891. Houck, úl (708). History of Missouri 1: 26. Chicago, 1908, Hus, H. (’07a). Germination of Hydrastis canadensis. Ann. Rept. Mo. Bot. Gard. 18: 85-94. 1907. , (707b). Virescence of Oxalis stricta. Ibid. 99-108. 1907. , (708). Ecological eross-section of b Mississippi River in the region of St. Sl Missouri. Ibid. 19: 127- Irish, H. C. (797). Comparative sbsaclesied — 325. 1897. ; (799). Street trees for St. Louis. Garden & Rept. Mo. State Bd. Hort, 4: Colman's Rural World 52: 379. Irwin, J. T. Manuscript list of trees and shrubs of the northeastern part of Mercer County, Missouri. 3 pp. Jeffrey, L., and Drouet, F. (734). Grass flora of Columbia, Mo. Rhodora 36: 415- 19 Jensen, L. P. C '13). Native and ricos plants of Missouri suitable for orna- mental planting. Special Bull. Mo. State Bd. Hort. 56: 13 » ( Wild flowers at the = Summit Extension. Mo. Bot. Gard. Bull, 18: 126- 127. 1930. , (31). A white-flowering partridge pea. Ibid. 19: 38-39. 1931. ———, (32a). New plant copos in pa d Summit Extension of the Gardan, Mo. Bot. Gard. Bul 41- , (732b). Ozarks are a dies flower il 32. The Blue Diamond 27: 14- 15 1 i (732e). The purple cone-flowers. Mo. Bot. Gard. Bull. 20: 68—69. ——— , (783). Some interesting primroses. Ibid. 21: 132—134. 1933. — J. H., and Hogstad, A., Jr. (730). I zn Extension, the home of h of wild flowers, Ibid. 18: 113-115. Kien. x ja History of seienee in St. Louis. 79-129. 1914. La Mance, L. S. (’91). Our negleeted native flowers. Garden and Forest 4: 358- 359. 1891. pen Acad. Sei, St. Louis 23: 1935] PALMER & STEYERMARK-—PLANTS OF MISSOURI 449 —— —————, (792a). A pretty native vine. Ibid. 5: 418. 1892. — — —— , (92b). Spiraea Aruncus. Ibid. 431. 1892. —, (793). The reappearance of wild flowers. Ibid. 6: 219. 1893. IG on. M. (731). The pim Bottom and the nde: d. plants of the region. Mo. Bot. Gari. Bull. 19: 99-109. 1931. Lewis, F. (’03). The Meramec tu: and the poison in their plants. St. Louis Daily Globe Democrat. May 3, 1903. Mackenzie, K. K. ('99). Spring flora of Kansas City and vicinity. 189 — — —— (assisted by B. F. Bush and others), (702). Manual of the Hid of Jack- son County, Missouri. 242 pp. , (705). Onosmodium. Bull. diet, Bot. Club 32: 495—506. 1905. , (706). Ranunculus sicaeformis. Torreya 6: 123. 1906. —— ————, (113). Western allies of Carex pennsylvanica. Ibid. 13: 14-16. 1913. —— — ——, (14a). A new genus from Missouri. Ibid. 14: 67-68. 1914. — — ——, (14b). A new southwestern sedge. Ibid. 125-127. 1914. — — —-, and Bush, B. F., (02a). The Lespedezas of Missouri. Trans. Acad. Sci. St. boite 12: 11-19. ph 1-4. 1902. ——————, —— ———— (’02b). New plants from Missouri. Ibid. 79-89. 1902. ———— —, —— — ——, (705). New plants from Missouri. Ann. Rept. Mo. Bot. Gard. 16: 102-108. 1905. Marbut, C. F. (704a). Geology and physiography in the state of Missouri. pp. 63- 70. ee 1904. ; . Physiography of Ozark region in Missouri. Abstract in Seienee 1904 "E (710); zi vrelinitan report on the general character of the soils and agriculture of the Missouri Ozarks. Mo. Agr. Exp. Sta. Res. Bull. 3: 151-273. 1910. , (711). Soil reconnaissance of the Ozark region of Missouri and Arkan- sas. U.S. Dept. Agr. Bur. Soils Rept. 13: 1727-1875 (especially pages 1842- 1844, 1864-1865, and discussion of vegetation in connection with soils). 1911. Maximilian, Prince of Wied neu-Wied, (1843). Systematic view of plants collected n a tour on the Missouri River. London, 1843 Metcalf, F. P. (22). Notes on marsh and aquatic plants of Missouri. Jour. Wash. Acad. S 4 Mische, E. (’95). Notes from the Midi Botanical Garden. Garden and Forest Nolen, J. H. (713). Missouri’s swamp and overflowed lands. Rept. to 47th Mo. Gen. Assembly. 141 pp. 1913. Nuttall, T. (1813). Fraser’s Catalogue.—A catalogue of new and interesting ien penna in Upper Louisiana and principally on the Missouri River, North America. London, 1813. [Reprinted in Pittonia 2: 114-119. 1889-92]. , (1821). Journal of travels into the Arkansa Territory during the year 1819. Philadelphia, 1821. [In Thwaites, Early Western Travels 13: 27-366. 1905]. Palmer, E. J. (*10). Flora of the Grand Falls Chert Barrens. Trans. Acad. Sci. St. Louis 19: 97-112. 1910. [Vor. 22 450 ANNALS OF THE MISSOURI BOTANICAL GARDEN — —— ——, (16). Catalogue of plants of Jasper Co., Mo. Ann. Mo. Bot, Gard. 3: 345—401. 1916. , (722). Forest flora of the Ozark region. Jour. Arn. Arb. 2: 216-232. 22. — — ——, (735). Synopsis of North American Crataegi. Ibid. 6: 5-128, 1925. — ——, (26). Leaves from a colleetor’s note book. Ibid. 7: 136-145. 1926. ——————, (31). Conspectus of the genus Amorpha, Ibid. 12: 157-197. pl. 36, and 31 — — ——, (32). Leaves from a eolleetor's note book. Ibid. 13: 417—437. 1932. ————, (84). Quercus ellipsoidalis in Missouri, Ibid. 15: 89. 1934. Pammel, L. H. (*11). Poisonous and medicinal plants of Missouri. Mo. State Bd. ort. Bull. 14: 46 pp. 1911. Peeh, F. (766). Catalogue of the United States plants in the Department of Agri- ure, 1866, Pursh, F. (1814). Flora Americae Septentrionalis. London, 1814. Record, S. J. (*10). Forest conditions of the Ozark region of Missouri, Mo. Agr. Exp. Sta. Bull. 89: 195-280. 1910. Rehder, A. (11). Trees and shrubs 2: 141-143. (Notes on Malus). 1911. — — ——, (13). Ibid. 229-232. 1913. Rickett, H. W. ig? A list of plants from the Missouri Ozarks. Am. Midl. Nat. 11: 243-2 ————, noto Flora of Columbia, Missouri. Univ. Mo. Stud. Sci. Ser. 6: 1-84, (31b). Notes on the vegetation of Columbia, Missouri, Am, Midl. Nat. 931. , (734a). Cornus amomum and C. candidissima, Rhodora 36: 269-274. , (34b). Specimens collected by Bradbury in Missouri Territory. Roy. Bot. Gard, Kew, Bull. Misc. Inf. 1934: 49-61. 1934. — m A (1835). Synopsis of the flora of the western states. 16 pp. Cincin- 835. [Review by L. H. Bailey in Bot. Gaz. 8: 269-271. 1883.] M "à J. (721). Precipitation and the growth of oaks at Columbia, Missouri. Mo. Agr. Exp. Sta. Res. Bull. 44: 19 pp. 1921. Rydberg, P. A. (’30). Taxonomie notes on the flora of the prairies and plains of central North America. Brittonia 1: 79-104. 1930. — (8 Flora of the prairies and plains of central North Ameriea, New York, 1932. Saeger, A. C. (729). The flowering of Lemnaceae. Bull. Torr. Bot. Club 56: 351- ; (34). Spirodela oligorhiza collected in Missouri. Ibid. 61: 233-236. 1934. Sargent, C. S. ('84—86). Botanical papers of George Engelmann. Bot. Gaz. 9: 69—74. 1884; 11: 192, 286. 6. , (784). Report on the forests of North America mem of Mexico). U.S. rtm Rept. (Missouri) 10: 560- y Washingto 03). "Trees and shrubs 1: 57, 89 ( Prone Seat): 63-65, pl. 32-33 (Crataegus Treleasei and C. eco 109, pl. 55 (Crataegus disjuncta) ; 113, pl. 57 (Crataegus lanuginosa); 117, pl. 59 (Crataegus Kellogg). 1903. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 451 , (705). Manual of trees of North America. 1905. , (707—13). Trees and shrubs 2: 3-13, pls. 102, 103, 104, 105, 106, 107 (Crataegus incaedua, C. ludoviciensis, C. rubicundula, C. Neo- ti, tri- anthophora, C. oil): 59-61, pls. 127, 128 (Crataegus severa, C. villiflora) ; 65-69, pls. 130, 131, 132 (Crataegus lanceolata, C. aspera, C. magnifolia) ; 75— 77, pls. 185, 136 (Crataegus padifolia, C. mollita) ; 137, pl. 156 (Hamamelis Hiis do 206—209 Pr on hiekories); 239-245 (Crataegus seclusa, C. mida, C. scabra); 247, pl. 192 (Prunus Palmeri); 261-262 Notes on es. 1907-1913. ( — ————, (708). Crataegus in Missouri. Ann. Rept. Mo. Bot. Gard. 19: 35-126. 1908. —— — ——, (1). Ibid. 22: 67-97. 1911. , (18). Notes on North American Tre I. Quercus. Bot. Gaz. 65: 423-459; II. Carya. Ibid. 66: 20-254; III. "Tila. Ibid. 421—438, 494, 504- 507. v 20-22). Notes on North American trees. VI. Jour. Arn. Arb. 1: 253. 1920; VIII. Ibid. 2: 164-174. 1921; IX. Ibid. 3: 5, 7; X. Ibid. ya 183. 1922. , (725). Notes on Crataegus. Ibid. 6: 4-5. 1925. Sauer, C. O. (7/20). The geography of the Ozark niet of Missouri. Bull. Geogr. Soe. Chieago 7: 52-59. 1920. Schooleraft, H. R. (1819). View of the lead mines of Missouri. 294 pp. New York, 1819. , (1821). Journal of a tour into the interior of Missouri and Arkansaw. 102 S London, 1821. — ————, (753). Scenes and adventures into the ceri e region of Ozark Modai of Missouri and mene ete. Philadelphi , (755). Summary narrative of an exploratory expwaltio’ to the sources of the TE River in 1820. Pilindsiphia, 1855. Seott, I. T. (’34). oody plants of Missouri [mimeographed]. State Resources Mus. Bull. 7. M Seavey, F. C. (797). Uncultivated erops in the Ozark Mountains. Garden and Forest 10: 465—466. T. ai J. I. Manuseript list of plants from St. Louis Co., collected in 1905-6. p. [in Mo. Bot. Gard. library]. Es m E. M. (793). Manuscript list of the plants of Greene County, Missouri. 1893 _— de 05). The New Madrid earthquake. Jour. Geol. 13: 45-62. 1905. a " E. (712). A new variety of Rudbeckia subtomentosa. Rhodora 14: 164. "s È S. Manual of Kansas City trees. Kansas City Teachers’ Training School Bull. 4: 4 pp. Small, J. K. (’96). Studies in the botany of pari e van United State, XV. (Oxalis Bushii). Bull. Torr. Bot. Club 25: 896. Spaulding, P. ('08). A biographieal history i oe at St. Louis, Missouri. [Baldwin, Beck, Bradbury, James, Lewis, Michaux]. Pop. Sei. Month. 73: 488—499. 1908. [Vor. 22 452 ANNALS OF THE MISSOURI BOTANICAL GARDEN — ———, (709). Ibid. II. [Drummond, Maximilian, Nuttall]; Ibid. 74: 48-57; III. Tühngeimansı, Geyer, Hilgard, Riehl] ; Ibid. 124-133; IV. [Eggert, Fendler, Glatfelter, Harris, Hedgeock, Hus, Irish, James, Kellogg, Letterman, Murt- feldt, Nelson, Norton, Pammel, Riley, Shaw, Spaulding, Thompson, Trelease, von Schrenk, Webber, Wislizenus]. Ibid. 240—258. 09. Starr, F. (’98). Academy of Natural Science of St. Louis. Ibid. 52: 629-647. Steyermark, J. A. (31a). Notes on Missouri plants found on certain geological formations. Mo. Bot. Gard. Bull. 19: 52-56. 1931. » (78 Some new and atera plants from Missouri, Ibid. 56-57. 1931. » (31e). A study of plant distribution in relation to the acidity of various soils in Missouri. Ann. Mo. Bot. Gard. 18: 41-55, 31 ————, (’33). Notes on Missouri arr Rhodora 35: 283-291. 1933 — ———, (94a). Epifagus virginiana in Missouri. Ibid. 36: 352-353. 1934. ; (734b). A grass new to Missouri, Ibid. 313-314. 1 —————, (34e). Hamamelis virginiana in Missouri. Ibid. 97-100. 1934. — ———, (34d). Recent additions to the flora of St. Louis County, Missouri, Ibid. 375-376. 1935. — — ——, (34e). Some features of the flora of the Ozark region in Missouri. Ibid. 214- 233. 1934 Sullivan, K. C. (728). A survey of the animal and plant life of the Niangua River and of the Bennett Spring Branch at Bennett Springs State = in Missouri. State Game and Fish Dept., Leaflet Hatchery Div. pp. 1-14. Swallow, G. C. (55). Catalogue of the trees and shrubs of dfi ven Rept. Mo. Geol. Surv. 2: 221-226. Appendix C. 5. ý A '58). Grape culture in Missouri. Trans. Acad. Sei. St. Louis 1: 156- 858. 168. i '59). Geological report of the country along the line of the southwest- ern iind of the Pacific Railroad, St. Louis. 93 pp. — — ^, (767). Trees, shrubs, and vines of Missouri. Rept. to 1st Mo. Gen. As- debi. 112 pp.; and Ann. Rept. State Bd. Agr. 2: 79-96. Torrey, J. (1845). emba of plants collected by Charles Geyer, under the direc- tion of I. N. Nicollet, during his exploration of the region between the Mis- sissippi yy Pra rivers. Rept. 28th Congress, 2nd session, House Doe. 52: 143-165, 1845. Tracy, S. M. oin Catalogue of the phaenogamous and vascular eryptogamous plants of Missouri. 106 pp. Jefferson City, 1886; and Ann. Rept. Mo. State Bd. Agr. 18: 397-500. 1885, and Broadhead, G. C. (786). Distribution of plants in Missouri. 1886. Trelease, w. (’85). Blooming times for flowers. Science Almanae. 5-7. 1885. ‚(’95). Leitneria floridana. An n. Rept. Mo. Bot. Gard. 6: 65-90. 1895. — — ——, (797). The swamps of southeastern Missouri. Garden and Forest 10: 370-371. 1897. — — —, (798). The Missouri dogbanes. Ann. Rept. Mo. Bot. Gard. 9: 147. 1898, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 453 —— ———, ('03). The Academy of Science of St. Louis. Pop. Sei. Month. 64: 117-130. 1903. , and Gray, A. (787). The botanical works of the late George Engelmann. 548 pp. Cambridge, 1887. Turner, J. A. (7/31). A new plant for Missouri. Mo. Bot. Gard. Bull. 19: 121. Ulrich, 2 G. (’34). New or noteworthy plants for St. Louis. VI. The native spider lily. Ibid. 22: Uphof, J. C. (’21). Das Yorker von Neviusia alabamensis im süden von Mis- souri. Deut. Dendrol. Ges. 1921: 282-283. fig. 1921. , (722). vine relations of plants in southeastern Missouri. Am. Jour. Bot. 9: 1-17. von Sehrenk, H. (798). T den of St. Louis as influenced by the tornado of . Trans. Acad. Sci. St. Louis 8: 25-41. 1898, Waddell, E. (729). Posies (wild flowers) in ‘‘Ozarks.’’ [Booklet sponsored by Springfield Lions Club]. 1929 Ward, L. F. (778). St. Louis and Botany. Field and Forest 3: 1878. i ases H. A Sketch Ne p of Academy of Science of St. Louis. Trans. cad. Sei. St. Louis 16: 15-99, 1906. Nus C. A (796). -a icc Biogr. Mem. Nat. Acad. Sci. 4: 3-21. 1896. Whitney, J. D. (776). Plain, prairie, and forest. Am. Nat. 10: 577-588, 656-667. Whitten, š C. (794). Phenologieal notes for 1892 and 1893. Ann. Rept. Mo. Bot. ard. 5: 123-135, 1894, Wight, W. F. er Native American species of Prunus. U. S. Dept. Agr. Bull. 179: 73p 1915. Williams, W. Du. The State of Missouri. 229—240. Columbia, 1904. Woodson, R. E., Jr. (730). The dogbane: roadside weed and future staple. Mo. Bot. Gard. Bull. 18: 87-104. , (732). The most interesting tree in Missouri. Ibid. 20: 145-149. 1932. ANNOTATED List or Missouri PLANTS SPERMATOPHYTA Fam. TO Pinus echinata Mill. Southern Yellow Pin Dry uplands, roeky slopes and bluffs, ET sometimes alluvial woods along PA in sandstone, chert, granitie or porphyritie trachyte areas, in mixed growth with deciduous trees or loeally in LI stands. Oxylophile. Southern Mo., south of a dur ne from Franklin to MeDonald counties: Franklin, Jefferson, Wash ingto e. Genevieve, isi se Madison, Wayne, St. Francois, Butler, ron, "Ra ds, Shannon, Carter, Ripley Texas, Dent, Phelps, Pulaski " Wright, Doblas dk. Taney, Stone, Christian, Barry, and McDonald counties. [Vor. 22 454 ANNALS OF THE MISSOURI BOTANICAL GARDEN The Austrian pine, Pinus nigra Arnold, has been found loeally established in woods in St. Louis Co., and the scrub pine, Pinus virginiana Mill, has been found in woods and open slopes in Callaway Co. The Scotch pine, Peut sylvestris L., and the white pine, Pinus Strobus L., are often planted but do not appear to establish themselves away from eultivation, and should not be considered as additions to the state flora. Taxodium distichum (L.) Richard. Bald Cypress. Swamps, ""- and flood plains of streams. Oxylophile to circumneutral. Southeastern Mo.: Cape Girardeau, Bollinger, Seott, Mississippi, Stoddard, New rid, Planos, Dunklin, Butler, and Ripley counties. Juniperus virginiana L. Red Cedar. Glades, bluffs, and borders of woods, on limestone or dolomite, and rarely on silieeous or granitie rocks. Calciphile to cireumneutral. General, but eom- monest in the Ozark regio Juniperus virginiana var. crebra Fernald € Griscom, Rh. 37: 133. 1935. Borders of woods and glades or bluffs. Circumneutral. Eastern Mo.: Lewis and Jefferson counties. Juniperus mexicana Spreng. White Cedar, Juniperus Ashei Buchholz, Bot. Gaz. 90: 329, figs. 1, 2. 1930. Juniperus sabinoides Nee Glades and dolomite blue of White River. Calciphile. Southwestern Mo.: Barry Co, Fam. TYPHACEAE Typha latifolia L. Common Cat-tail. Swamps and muddy margins of ponds and streams. Cireumneutral. General. Typha angustifolia L. Narrow-leaved Cat-tail. Muddy borders and in shallow water po saline spring-fed lake. Caleiphile. Local: Saline Co. This has often been confused with narrow-leaved and slender- spiked forms of T. latifolia. Fam. SPARGANIACEAE Sparganium eurycarpum Engelm. Bur-reed. Swamps and borders of ponds and streams. Cireumneutral. General but not common: St. Charles, St. Louis, Clay, and Jackson counties. Bparganium americanum Nutt. See Rh. 24: 27,33. 1922, Bur-reed. Sparganium androcladum of auth., not Morong (B& B Sparganium americanum var. ira Fernald & Sinai: not S. simplex var. marae Engelm. (G). n spring branches pe margins of streams and m Cireumneutral to ealeiphile. Southern and central Mo.: St. Louis, Jefferson, St. Franeois, Bol- linger, Ripley, New Madrid, Dunklin, Reynolds, ‘Gud , Phelps , Dent, Shannon, Texas, Howell, Camden, Jackson, Greene, Jasper, and Kelle counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 455 onem androcladum (Engelm.) Morong. See Rh. 24: 27, 32. 1922. Bur- eed. Sparganium lucidum Fernald & Eames (G), (B & B). Borders of ponds and swales. Cireumneutral. Southern and central Mo.: St. Louis, Jefferson, Iron, St. Francois, Scott, Dunklin, Boone, Cass, Greene, and Jackson counties. Fam. NAJADACEAE Potamogeton pulcher Tuckerm. ien Pondweed. ow streams and ponds. Cireumneutral. Eastern Mo., south of the Missouri River: St. Louis, Jefferson, bin , Dent, Texas, Pinno, Iron, and Ripley counties. Yir a podios has ¿cesa been confused with Potamogeton natans which is not known in Mo. ee amplifolius Tuckerm. Large-leaved Pondweed. oughs, streams, ii spring branches, and in ids fed by cold springs. mos eutral. Southern and eentral Mo.: Dallas, Morgan, Camden, Howell, Texas, Douglas, Ries ps counties. Potamogeton illinoensis Morong. Sloughs and slow waters along streams. Circumneutral. Southeastern Mo., local: Ripley Co. This species has sometimes been confused with Sobeiodets icanus, a common species in Mo. Ii americanus Cham. € Schlecht. Long-leaved Pondweed. Ponds and rivers. Cireumneutral Southern and central Mo.: Lou Jefferson, Franklin, St. Francois, Iron, New Madrid, Dunklin, Ripley, Gasconade, M run. Camden, Miller, Laclede, Taney, Jackson, Jasper, and Newt "heroe lucens L. Shining Pondweed. Slow-flowing streams, springs, and spring branches, and in lakes fed by eold springs. Cireumneutral to ealeiphile. Southeastern and middle Mo.: Pe Iron, Wayne, Dunklin, Reynolds, Shannon, Carter, Pulaski, and Laclede baitis. POTAMOGETON CRISPUS L. Curly Muck-weed. Ponds, sloughs, and lakes fed by cold springs. Circumneutral. Southern Mo.: Crawford and Newton counties. Potamogeton foliosus Raf. var. genuinus Fernald, Mem. Am. Acad. 17: 43. 1932. treams, springs, and ponds. Circumneutral to caleiphile. General: St. Louis, Wayne, Butler, Shannon, Carter, Reynolds, Oregon, Howell, Maeon, Boone, Morgan, Camden, Pulaski, Laclede, Dallas, Wright, Mood er Ozark, Douglas, Platte, dur. Jackson, Bates, Jasper, and Barry co Potamogeton foliosus var. macellus Fernald, Mem. Am. Acad. 17: 46. 1932. Potamogeton foliosus Raf. in part (G). Similar piso to the que Cireumneutral to calciphile. Southern, central, and w n Mo. ouis, Butler, Crawford, Reynolds, Boone, Miller, aclede, Howell, ae ma AM and Christian counties. Potamogeton panormitanus Bivona-Bernardi var. major G. Fisc Slow streams and ponds. Cireumneutral Middle Mo., beat! | Benton Co. [Vor. 22 456 ANNALS OF THE MISSOURI BOTANICAL GARDEN Potamogeton panormitanus var. minor Biv. Slow streams. Circumneutral. Southern Mo.: Perry and Laclede counties. Potamogeton diversifolius Raf. See Mem. Am. Acad. 17: 105-106. 1932. Pond- eed. Potamogeton dimorphus of Am. auth. in part, not Raf. (G), (B & B). Potamogeton Spirillus of Am. auth. in part, not Tuckerm. De hybridus of Am. auth. in part, not Michx. (G Streams and ponds. Circumneutral. General: Scotland, SER TER Audrain, Mud St. Louis, Bollinger, Wayne, Butler, Dent, Shannon, Oregon, Caldwell, Ray, Clay, Jaekson, Henry, Greene, Wright, Webster, Ozark, Taney, and Jasper counties. Potamogeton pectinatus L. Fennel-leaved Pondweed. Ponds and streams. Caleiphile to cireumneutral General but scattered: St. — Perry, Ripley, Osage, Phelps, Saline, Atchison, Buchanan, and Platte counti Fio rotundatus Hagst. Ponds and slow streams. Circumneutral. Local: St. Louis Co. Najas guadalupensis (Spreng.) Morong. See Rh, 25: 107. 1923. Naiad. Marshes and shallow water of streams. Cireumneutral. Central and southern t. Louis, St. Franeois, Iron, Reynolds, Ripley, Phelps, Dent, Shannon, Tossa. Boone, Laelede, Greene, Platte, Jaekson, and Vernon eounties. Zannichellia palustris L. Horned Pondweed. arshy and springy ground, and in ponds and shallow streams, frequently in saline marshes and springs. — to eireumneutral. General but scat- tered: Clark, St. Louis, Jefferson, Perry, Randolph, Cooper, Saline, Maries, Miller, Dent, Reynolds, Carter, die Texas, Crawford, Laclede, Dallas, Jack- son, and Newton counties. Ruppia maritima L. var. rostrata Agardh. Ditch-grass. Ponds and shallow streams. Local: Henry Co. Fam. ALISMACEAE Sagittaria longirostra (M. Micheli) J. G. Smith. pane -beaked Arrow-head. Seattered and uncommon: Henry and Jasper countie All of the Missouri species of Sagittaria are ti in shallow water or on muddy borders of sloughs, ponds, and sluggish streams, most frequently in eireumneutral or alkaline soils. Sagittaria latifolia Willd. Broad-leaved Arrow-head. General and common. Sagittaria latifolia f. obtusa (Muhl.) Robinson. Frequent with the typical form. General. Sagittaria brevirostra Mack. & Bush. Short-beaked Arrow-head. General and common. Sagittaria ambigua J. G. Smith. Lance-leaved Sagittaria. Southwestern Mo.: Barton, Jasper, and Newton counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 457 Sagittaria heterophylla Pursh. Scattered: Callaway, Boone, Jackson, Dent, Texas, Howell, Wright, Ozark, Webster, and Greene counties, Sagittaria heterophylla var. elliptica Engelm. Seattered: St. Louis and Boone eounties. Sagittaria heterophylla var. RI dH em Scattered: Greene and Texas counti Sagittaria graminea Michx. Grass- pem dites General but scattered: St. Louis, Butler, Texas, Carroll, Jackson, Jasper, Lawrence, Greene, and Taney counties. Sagittaria platyphylla Ma vu J. G. Smith. Southeastern Mo. unklin and Butler counties. Lophotocarpus ayeni LAS J. G. Sm hallow water and muddy margins of a" ponds, and sluggish streams. Caleiphile to cireumneutral. General. Echinodorus tenellus (Martius) Buchenau. Bur-head. et ground about ponds and sloughs. Circumneutral. Local: St. Louis Co. Echinodorus cordifolius (L.) Griseb. Bur-hea wamps and muddy banks. Obiin. Scattered, mostly along the larger rivers: Lincoln, St. Louis, Franklin, Mississippi, Ripley, and Jackson counties. Echinodorus cordifolius var. lanceolatus (Engelm.) Mack. & Bush. Similar situations to the preceding. Circumneutral. Scattered: Clay and Jackson counties. Echinodorus radicans AN "pd Creeping Bur-head. Sw Cireumneutral. Southeastern Mo., and north along Mississippi River: B Louis, Jefferson, Stoddard, New Madrid, Dunklin, and Butler counties. Alisma subcordatum Raf. See Arkiv f. Bot. 24A': 19. 1932. Water Plantain. lisma Plantago-aquatica of Am. auth. in part, not L. (G). Swamps and muddy i ie of sloughs, ponds, and sluggish streams. Cir- cumneutral, General. This has been confused with Alisma Plantago-aquatica and subsp. brevipes (as defined 1 by Samuellson) which do not occur in Mo Vallisneria americana Michx. See Rh. 20: 108. 1918. Eel-grass. Vallisneria spiralis of Am. auth., not L. (G), (B& B Slow streams. Cireumneutral. Fast ern Mo., south of the Missouri River: St. Louis, Jefferson, Franklin, Gasconade, Phelps, Iron, Butler, Carter, and Ripley unties. Fam. HYDROCHARITACEAE a occidentalis pig Bw ‘Contr. Lab. Bot. Univ. Montreal, No. WELT also Rh. 14-116. 1932. Water-weed. Eon canadensis idea in part (G) Philotria canadensis (Michx.) Britton in part (B & B). Ponds, pools, and sloughs, on the surfaee of stagnant water, in springs and spring branches, and margins of streams. Circumneutral to calciphile. South- [Vor. 22 458 ANNALS OF THE MISSOURI BOTANICAL GARDEN ern and central Mo.: Boone, wg St. Charles, St. Louis, J apti Crawford, Madison, Reynolds, Carter, Butler, Dunklin, Ripley, Oregon, Sha , Texas, Osage, Cole, Miller, Phelps, Eae Dallas, Camden, Laclede, Wright, "Douglas, Ozark, Jackson, and Jasper counties. Limnobium Spongia (Bosc) Richard. American Frog-bit Streams and sloughs. Cireumneutral. Southeastern Mo., local: Dunklin Co. Fam. GRAMINEAE Arundinaria gigantea (Walt.) Chapm. See Am. Jour. Bot. 21: 127. 1934; also Bull. Torr. Bot. Club 56: 315-318. 1929. Switch Cane, Giant Cane. Arundinaria macrosperma Michx. (G), (B & B), in part. Arundinaria tecta (Walt.) Muhl, (G), (B & B), in part. Low wet woods and moist ground along —€— Cireumneutral. Southern Mo., mostly in the southeastern lowlands and along White River and its tribu- tape: Bollinger, Cape Girardeau, Scott, bene Stoddard, New Madrid, Pemiscot, Dunklin, Butler, Wayne, Ripley, Carter, Reynolds, Oregon, Shannon, Howell, Douglas, Ozark, Taney, Stone, Barry, and MeDonald counties. Bromus breviaristatus Buckley. Introduced in Jackson Co. Bromus catharticus Vahl. See Am. Jour. 22 21: 127. 1934. Bromus unioloides (Willd.) HBK. (B Waste ground. Western Mo., local: is edt Co. BROMUS INERMIS Leyss. Hungarian Brome Grass. Introduced in fields, waste ground, and roadsides. Seattered: St. Louis, Boone, J E and MeDonald counties BROMUS TECTORUM L. Int Diisi in fields and waste ground. Indifferent. General. Bromus purgans L. Wild Chess. Rocky or rich woods and bluffs. Circumneutral to calciphile. General. Bromus latiglumis (Scribn.) Hitehe. See Rh. 35: 316. 1933. Bromus purgans var. latiglumis (Seribn.) Shear. Bromus altissimus Pursh, not Gilib. (G). Woods and bluffs. Calciphile. Eastern Mo.: Clark, St. Louis, and Jefferson counties. Bromus latiglumis f, incanus (Shear) Fernald, Rh. 35: 316. 1933. Bromus altissimus f. incanus (Shear) Wiegand. Wooded banks. Caleiphile. Western Mo.: Indos Co. BROMUS SECALINUS L. Chess, Cheat. Fields and waste ground. Indifferent. General. BROMUS MOLLIS L. See Am. Jour. Bot. 21: 128. 1934. Soft Chess. Bromus hordeaceus of auth., not L. (G), (B & B). Introduced in waste ground and along railways. Scattered: St. Louis and Jackson counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 459 BROMUS RACEMOSUS L. See Rh. 24: 90. 1922. Introduced along Da railways, and in waste ground. Indifferent. Scattered: St. Louis, Jackson, Vernon, and Jasper counties. Introduced in fields and waste ground. Indifferent. Seattered: Iron, Boone, Clay, Jackson, Webster, and Jasper counties. BROMUS ARVENSIS L. Field Chess. Introduced in fields and waste ground. Indifferent. Scattered: Lewis, Au- drain, Jefferson, Boone, Dekalb, Jackson, and Jasper counties BROMUS JAPON Thunb. ntro duced in fields, waste ground, and roadsides. Indifferent. Scattered: Boone and Jackson counties. Festuca octoflora Walt. Slender Fescue Grass. Fields, prairies, glades, rocky open or dry woods, and waste ground. Oxy- lophile. Southern and eentral Mo. south and east of a line drawn from St. Louis and Morgan counties to Vernon Festuca octoflora var. tenella (Willd.) Fernald, Rh. 34: 209. 1952. Similar situations to the preeeding. Oxylophile. General and common. This is the commonest variety found in Mo Festuca octoflora var. glauca (Nutt.) ag a, an 34: 209. 1932. Similar situations to the preceding. Ox ile. Southern and central Mo. south and east of a line drawn from St. NE od and Morgan counties to Vernon Co Festuca sciurea Nutt. Rocky or sandy open ground. Oxylophile to eireumneutral. Southern Mo., scattered: Dunklin and Jasper counties. FESTUCA ELATIOR L. Meadow Fescue. eadows, open woods, and waste ground. Cireumneutral. General. Festuca obtusa Spren Festuca mutans Spreng. (G), (B & B Roeky or rieh woods, bluffs, alluvial ground along streams, and waste ground. Cireumneutral. General. Festuca Shortii Kunth. Moist open ground and woods. Circumneutral. General but scattered: Clark, St. Louis, Jefferson, Dent, Shannon, Boone, Jackson, and Jasper counties. Glyceria Scu MOTA Torr. Mar of ponds, in shallow water. oo Southern Mo.: Dent, Texas, Howell, Wright, Webster, and Ozark countie Glyceria arkansana Fernald, Rh. 31: 49. 1929. Swampy ground. Cireumneutral Southeastern Mo.: Dunklin, Butler, and Ripley counties. Glyceria septentrionalis Hitche. Floating Manna Grass. Borders of sloughs, wet open woods, and meadows. Circumneutral. Northern and central Mo.: Scotland, St. Louis, Jefferson, a , and J counties. 460 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Glyceria pallida (Torr.) Trin. Pale Manna Grass. Swampy ground. Circumneutral. Southeastern Mo. : Scott and Butler eounties. Glyceria striata (Lam.) Hitehe. Proc. Biol. Soc. Wash. 41: 157. 1928. Fowl M Grass. Glyceria nervata (Willd.) Trin. (G). Panicularia nervata (Willd.) Kuntze (B & B). Sloughs, bogs, wet woods, and meadows. Circumneutral. General. Poa ANNUAL. Low Spear Grass, Annual Blue Grass Fields, gardens, and waste ground. Indifferent. General and eommon. Poa Chapmanniana Seribn. Prairies, glades, and waste ground. Cireumneutral. Central and southern Mo.: St. Louis, Jefferson, Dent, St. Franeois, Shannon, Texas, Boone, Jaekson, Polk. Greene, Christian, Douglas, Ozark, Stone, Jasper, and Newton counties. PoA COMPRESSA L. Canada Blue Grass, English Blue Grass. ields and rocky waste ground. Indifferent. General. Poa nemoralis L, Introduced along railroads. Jackson Co. Poa pratensis L. Kentucky Blue Grass. Fields, meadows, glades, open woods, and waste ground. Circumneutral to calciphile. General and common Rh. 18: 235. 1916. Poa triflora Gilib. (G), (B & B). Wet places and open ground. Cireumneutral. Seattered: St. Louis, Ste. Genevieve, and Jackson counties. Poa sylvestris Gray. Rich or alluvial woods. Poa Wolfii Scribn. Rocky woods, moist c and along streams. Calciphile to eireumneutral thern Central and sou : Franklin, Shannon, Texas, Boone, Jackson , Jasper, MeDonald, Taney, and =. eounties. Cireumneutral. General, — hypnoides (Lam.) BSP. See Rh. 28: 114. 1926. Creeping Love Gras Wet alluvial river banks and margins of sloughs and ponds. Cireumneutral. General. Eragrostis reptans (Michx.) Nees. See Rh. 28: 114-115, 1926. Eragrostis Weigeltiana (Reichenb.) Bush (B & B Moist alluvial ground along the Missouri River. Circumneutral. Western Mo., local: Jackson Co. Eragrostis capillaris (L.) Nees. Love Gra Fields, S rocky ledges, glades, ef waste ground, Indifferent. Gen- eral and commo Eragrostis Frankii pe Mey. & Lall.) Steud Waste alluvial and cultivated ground. Circumneutral. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 461 ERAGROSTIS CILIANENSIS (All) Link ex Lutati, Malpighia 18: 386. 1904; see Hitchcock, Man. Grasses U. S. Misc. Publ. U. S. Dept. Agr. 200: 154. 1935. ragrostis megastachya TER Link (G). Eragrostis major Host (B Waste and eultivated Br Indifferent. General. ERAGROSTIS POAEOIDES (L.) Beauv. Nes Hiteheoek, Man. Grasses U. S. Mise. Publ. U. S. Dept. Agr. 200: 156. Eragrostis minor Host (G), (B "e Waste ground. Indifferent. Introduced in Stone Co. Eragrostis pilosa (L.) Beauv. Indian Love Grass. Moist banks, rocky open or alluvial or waste ground. Indifferent. General. Eragrostis En = ) Nees. See Hitchcock, Man. Grasses U. 8. Misc. Publ. U. S. De r. 200: 151. 1935. Eragrostis o Wisi Seribn. Eragrostis Purshii Sehrad. (B & Alluvial or open ground sud o and waste places. Indifferent. - General. Eragrostis diffusa Buckl. See Hitchcock, Man. Grasses U. 8. Misc. Publ. U. $. Dept. Agr. 200: 151. 1935. Open ground. Introduced in Jaekson Co. Eragrostis hirsuta (Michx.) N Open ground. roe en Southwestern Mo.: Jasper Co. Eragrostis intermedia Hitche. Jour. Wash. Acad. Sci. 23: 450. 1933. Prairies and open ground. Cireumneutral. Western Mo., seattered: Jackson and Newton counties. This has been confused with Eragrostis lugens which does not reach Missouri Eragrostis spectabilis (Pursh) pape See Hitchcock, Man. Grasses U. 8. Misc. Publ, U. S. Dept. Agr. 200: 163-164. 1935. Purple Love Grass. Eragrostis pectinacea of Am, auth., not Miehx. (G), (B & B). Eragrostis pectinacea var. Wee is Gray (G). Sandy or rocky open or waste ground. Oxylophile to circumneutral. General and common. Eragrostis trichodes (Nutt.) Nash. Moist open, sandy or rocky ground. Oxylophile. Central and southern Mo.: Ste. Genevieve, Dent, Ozark, Carell and Jackson countie Diarina festucoides Raf. Diarrhena diandra P deo 2 Wood (G), (B&B). Diarrhena americana Low, rich or uae dine and river banks. Cireumneutral. General but scattered: St. Jefferson, Franklin, Shannon, Boone, Camden, Gentry, Clinton, Clay, m Hickory, Jasper, McDonald, Stone, and Taney counties. Distichlis stricta (Torr.) Rydb. Salt Grass, Alkali Grass. Along railroads and waste ground. Indifferent. Western Mo.: introduced in Buchanan and Jaekson counties. [VoL. 22 462 ANNALS OF THE MISSOURI BOTANICAL GARDEN Uniola latifolia Michx. Spike Gra Rocky woods, bluffs, moist Fh and alluvial or rich low woods. Circum- neutral to ealeiphile. General. DACTYLIS GLOMERATA L. Orchard Grass. Fields, waste ground, and roadsides. Indifferent, General. ARUNDO Donax L. Giant Reed. Open ground. Escaped from cultivation in Dunklin Co. Phragmites communis Trin. var. Berlandieri (Fournier) Fernald, Rh. 34: 211. 1932. Reed. Phragmites communis of Am. auth. in part (G), (B € B). ow ground along streams and ponds, Circumneutral. Northern and west- central Mo.: Maeon, Nodaway, Platte, Clay, Jackson, Lafayette, Saline, and Bates counties. Melica Porteri Scribn. Rocky wooded bluffs, Calciphile. Western Mo., local: Jackson Co. Melica mutica Walt. Melic Grass. Rocky woods and bluffs, Calciphile. General: St. Louis, Jefferson, Madi- son, Iron, Wayne, Carter, Shannon, Texas, Miller, Hickory, Callaway, Mercer, Dekalb, Jackson, and Jasper counties. Melica nitens Nutt. Melic Grass. Rocky woods, glades, and bluffs. Caleiphile. General but scattered: St. Louis, Jefferson, Wayne, Oregon, Boone, Camden, Gentry, Clinton, Clay, Jack- son, Hiekory, J bcp MeDonald, Stone, and Taney counties. Triodia flava (L.) Smyth. See Hitchcock, Man. Grasses U. S. Mise. Publ. U. S. Dept. Agr. 200: 213, "Ns 1935. "Tall a Top, Purple Top. Tridens flavus (L.) Hitehe. (G), (B € B). Tricuspis flava (L.) Hubbard. Open woods, meadows, glades, and waste ground. Cireumneutral General. Triodia stricta (Nutt.) Benth. See Hitchcock, Man. Grasses U. S. Mise. Publ. U. 8. Dept. Agr. 200: 215 and 971. 1935 Tridens strictus (Nutt.) Nash (G), (B & B). Tricuspis stricta (Nutt.) Gray. Prairies and moist open ground. Circumneutral to oxylophile. Southern Mo.: Mississippi, Dunklin, Butler, Dade, Barton, J asper, and Newton counties. Triodia elongata (Buckl.) Seribn. See Em Man. Grasses U. S. Misc. ubl. U. S. Dept. Agr. 200: 217, 968, 193 ridens elongatus (Buckl.) Nash (B ^ B). Tricuspis elongata (Buckl.) Heller. Rocky bluffs and glades. Calciphile. Southwestern Mo.: Henry, Barry, and McDonald counties, Triplasis purpurea (Walt.) Chapm. Sand Grass Sandy open ground. Oxylophile, Central and southeastern Mo.: St. Louis, Jefferson, Dunklin, and Jackson counties. AGROPYRON REPENS (L.) Beauv. var. SUBULATUM (Scribn.) Reichenb. See Rh. 35: 183-184, 1933. Quitek Grass, Couch Grass, Quack Grass. Introduced in fields and waste ground. Indifferent. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 463 Agropyron Smithii Rydb. Blue-joint. Prairies, fields, and waste ground. Indifferent. General but scattered. Agropyron do (Link) Steud. var. typicum Fernald, Rh. 35: 169, 1933, Awned Whea Agropyron as Vasey (G), (B&B). Agropyron caninum var. tenerum (Vasey) Pease & Moore. Introduced in fields and waste ground. Indifferent. Scattered in western ckson and Jasper counties. This has been confused with Agropyron Ei eue which does not occur in Mo. AEGILOPS CYLINDRICA Host. Triticum Aegilops Beauv. riticum cylindricum Ces., Pass. & Gi nodu along ibat Jackson Co. Elymus glaucus Buckley. Blue Wild Rye. Elymus Mackenzii Bush. Rocky wooded bluffs. Calciphile. Southwestern Mo.: Jasper, Barry, and Ozark counties. This has been confused with Elymus Macounii which does not reach Mo Elymus villosus Muhl. See Rh. 35: 195. 1933. Elymus striatus of Am. auth., not Willd. (G), (B & B). Open or rocky woods and prairies. Circumneutral to calciphile. General. Elymus villosus f. arkansanus (Scribn. & Ball) Fernald, Rh. 35: 195. 1933. Rocky woods. Circumneutral to maed Southern Mo.: Oregon, Greene, Barry, and Jasper counties. ro canadensis L. Sce Rh. 35: 191. 1933. Nodding Wild Rye. Elymus robustus var. vestitus Wiegand. Rocky woods and prairies, and waste ground. Circumneutral. General. Elymus canadensis f. glaucifolius (Muhl.) Fernald, Rh. 35: 191. 1933. Elymus glaucifolius Muhl Elymus glaucifolius var. robustus (Seribn. & Sm.) Bush. Elymus sowie var. crescendus (Ramaley) Bush. Elymus robustus Seribn. & Sm. (G). Rocky open E alluvial ground, prairies, and waste places. Cireumneutral, General. Elymus virginicus L. See Rh. 35: 198. 1933. Wild Rye. Elymus striatus Willd., but not of most recent authors. Alluvial ground, open woods, and prairies. Cireumneutral. General. Elymus virginicus f. hirsutiglumis (Scribn.) Fernald, Rh. 35: 198. 1933. Elymus virginicus var. hirsutiglumis (Seribn.) Hitche. (G). Elymus hirsutiglumis Seribn. (B & B). Open woods and prairies. Cireumneutral. General but seattered. Elymus virginicus var. jejunus (Ramaley) ps Am. Midl. Nat. 10: 65. 1926. Elymus jejunus (Ramaley) Rydb. (B y Fields, open woods, and waste ground. Eu General. [Vor. 22 464 ANNALS OF THE MISSOURI BOTANICAL GARDEN Elymus ze. var. submuticus Hoo Open woods. Cireumneutral. Sonti: Jefferson, Oregon, and Jackson Ri Elymus virginicus var. glabriflorus (Vasey) Bush, Am. Midl. Nat. 10: 62. 1926. Elymus glabriflorus (Vasey) Seribn. & Ball (B & B). Elymus australis var. glabriflorus (Vasey) Wiegand. Open, rocky, rich or alluvial woods, and prairies. Cireumneutral. General. Elymus bg pong var. glabriflorus f. australis (Scribn. € Ball) Fernald. See Rh. 35: 935. Pili on Seribn, & Ball (G), (B & B). Rocky open woods and prairies. Circumneutral. Central and southern Mo. Sitanion hystrix (Nutt.) J. G. Smith Introduced along railroads. Jackson Co. Hystrix patula Moench. See Am. Jour. Bot. 21: 133-134. 1934. Bottle-brush. er Hystrix (L.) Millsp. (B & B). Open pond woods and dry open or waste ground. Cireumneutral to oxy- lophile. Gener Hordeum jubatum L. Squirrel-tail Grass. Fields, roadsides, and waste ground. Indifferent. General, except in the Ozark region, but commonest in northern Mo., where native, and probably intro- dueed elsewhere, Hordeum pusillum Nutt. Cultivated and waste ground. Indifferent. General. Hordeum nodosum L. Meadow Barley. Dry open ground. Indifferent. Eastern Mo., scattered: St. Louis, Jefferson, Franklin, and Phelps counties. HORDEUM VULGARE L, var. TRIFURCATUM (Schlecht.) Alefeld. Beardless Barley. Waste ground. Indifferent. Introduced in Jackson Co. LOLIUM PERENNE L. Perennial Rye Grass. Introduced in fields, meadows, and waste ground. Indifferent. General. LOLIUM MULTIFLORUM Lam, Italian Rye Grass. Introduced in similar situations to the preceding, but less common. Indif- ferent. Scattered: Boone, Saline, and Jackson counties. LOLIUM TEMULENTUM L. Darnel. Introduced in fields and waste ground. Indifferent. Jackson Co. Koeleria cristata (L.) Pers. Crested Hair Grass. Prairies and fields. Circumneutral. General. Sphenopholis obtusata (Michx.) Seribn. Early Bunch Gra Open rocky ground and bluffs. oe to caleiphile. General. Sphenopholis obtusata var. pubescens (Seribn. & Merr.) Seribn. ocky open ground. Scattered. Southern and east-central Mo.: St. Louis, Jefferson, Shannon, Seott, and Taney counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 465 Sphenopholis obtusata var. lobata (Trin.) Scribn. Occasionally found with the typical form. Sphenopholis intermedia Rydb. See Bartonia 14: 34. 1932; also Am. Jour. Bot. 21: 134. 1934. Sphenopholis pallens of auth., not Aira pallens Spreng. Moist rocky open ground. Circumneutral. General. Sphenopholis nitida (Spreng.) Scribn. Rocky woods. Cireumneutral Southern Mo., scattered: Ste. Genevieve, Ripley, Reynolds, Carter, Douglas, and Wright counties. Trisetum flavescens (L.) Beauv. Yellow False Oats. Introduced along railroads. Jackson Co AVENA SATIVA L. Oats. Commonly rg in waste ground and along railroads and roadsides. In- different. Gene AVENA FATUA L. Wild Oats. Oceasionally introduced in waste ground. Scattered. ARRHENATHERUM ELATIUS (L.) Beauv. Oat Gra Fields and waste ground. Introduced. eg Jasper Co. HOLCUS LANATUS L. Velvet Grass. Notholous lanatus (L.) Nash (B & B). Introduced along railroads. Jackson Co. Danthonia spicata (L.) Beauv. Wild Oat Gra Rocky prairies, Mass woods, and glades. Oxylophile. General, but commonest in central and souther Calamagrostis canadensis TP Nutt. var. typica Stebbins, Rh. 32: 40. 1930. Blue-joint Grass. Wet meadows and prairie swales. Cireumneutral. Northern and central Mo.: Mereer, Sar, Linn, Ralls, St. Louis, Dekalb, and Jackson counties. Calamagrostis canadensis var. Macouniana (Vasey) Stebbins, Rh. 32: 41. 1930. edi Macouniana Vasey (B & B). S. Circumneutral. rthern and central Mo.: Clark, Lewis, m Pin Adair, Grundy, Harrison, and Jackson counties. Calamagrostis inexpansa Gray var. brevior (Vasey) Stebbins, Rh. 32: 50. 1930. mf inexpansa Gray, in part (G), (B & B). w wet woods. Cireumneutral Rare and local. Southern and central Mo.: see and Texas counties Calamovilfa longifolia (Hook.) Hack. Long-leaved Reed Grass. Low sandy open ground. Oxylophile to circumneutral. Central Mo.: St. Louis, Jefferson, Clay, and Jackson counties. AGROSTIS SPICA-VENTI L. a spica-venti (L.) Beauv. (G), (B & B). Waste ground. Introduced in Jackson Co. AGROSTIS INTERRUPTA L. Waste ground. Introduced in St. Louis Co. [Vor. 22 466 ANNALS OF THE MISSOURI BOTANICAL GARDEN AGROSTIS STOLONIFERA L. See Rh. 35: E 1933. grostis alba of auth., not L. (G), (B MM and waste iud: A General and common. AGROSTIS TENUIS Sibth. Agrostis alba var. vulgaris (With.) Thurb. (G). In similar situations to the preceding. Indifferent. General and common. Agrostis Elliottiana Schultes. en sandy or cherty ground, sterile fields, prairies, and glades. Oxylophile. East-central and southern Mo.: Lincoln, Montgomery, St. Louis, Jefferson, St. Francois, Iron, Ste. Genevieve, Madi ison, Bollinger, Dunklin, Butler, Wayne, Shannon, Dent, Crawford, Washington, Phelps, Maries, Pulaski, Gascona s Franklin, — Miller, Laclede, Christian, Webster, Barry, Jasper, pe MeD ald countie Agrostis scabra Willd. See Rh. 35: 207. 1933. Hair Grass, Tickle Grass. is hyemalis of auth., not Walt. (G), (B & B). Fields, meadows, and open woods. dria to circumneutral. General. Agrostis perennans (Walt.) Tuckerm. Thin Gra Fields, prairie swales, margins of streams wg oh low or rieh woods, and moist open ground. Cireumneutral. General Agrostis ee ans var. aestivalis Vasey. See Rh. 35: 318. 1933. Agrostis Schweinitzii Trin, ut banks, slopes, rich shaded woods, and along streams. Cireumneutral. General. . Cinna arundinacea L. Wood Reed Grass. Wet woods and meadows. Cireumneutral. General. ALOPECURUS PRATENSIS L. Meadow Fox-tail Grass. s, meadows, and waste ground. Indifferent, Scattered: St. Louis and Boone counties. Alopecurus carolinianus Walt. See Am. Jour. Bot. 21: 136. 1934. Alopecurus geniculatus of auth., not L. (G), (B & B). Alopecurus geniculatus L. var. ramosus (Poir.) St. John, Rh. 19: 167. 1917. eadows, waste ground, borders of ponds, sloughs, wet ground, and wet depressions in rocky glades. Circumneutral to ealeiphile. General. Typieal A. geniculatus does not reach Mo. This species is much commoner than the next following. Alopecurus aequalis Sobol. See Rh. 27: 196. 1925, Floating Fox-tail Grass. Alopecurus geniculatus var. aristulatus (Michx.) Torr. (G). Alopecurus aristulatus Michx. (B & B). Swampy ground, borders of ponds and sloughs, and wet open ground. Circum- neutral. General. PHLEUM PRATENSE L. "Timothy. Meadows, roadsides, and waste ground. Circumneutral. General. Muhlenbergia cuspidata (Torr. Rydb. See Rh. 18: 233. 1916. Sporobolus brevifolius (Nutt.) Seribn., in part jn Dry banks, rocky bluffs, and loess hills. Cireumneutral to calciphile. Western 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 467 and southern Mo.: Crawford, Phelps, Pulaski, Miller, Camden, Atchison, Holt, Buchanan, Jackson, Henry, and McDonald counties. Muhlenbergia asperifolia (Nees & Meyen) Parodi. See Hitchcock, Man. Grasses U.S. Mise. Publ. U. S. Dept. Agr. 200: 366. 1935. Scratch Grass. Sporobolus asperifolius (Nees € Meyen) Thurber r (B & B). Moist sandy ground. Oxylophile. Western Mo.: Holt and Jackson counties. Muhlenbergia sobolifera (Muhl.) Trin. Rocky woods, shaded slopes, limestone bluffs, and prairies. Calciphile to eireumneutral. General. Muhlenbergia tenuiflora (Willd.) BSP ocky woods, glades, dry open oi atta: and prairies. Oxylophile to circum- neutral. General Muhlenbergia racemosa (Michx.) BSP. Moist prairies, alluvial banks, meadows, and open woods. Circumneutral. General. E ev mexicana (L.) Trin. Satin Gra Ro s and moist woods. Ci lcd. General. um is mexicana f. commutata (Scribn.) Wiegand, Rh. 26: 1. 1924. In similar situations to the typical form. Local: Lincoln Co. Muhlenbergia glabriflora Scribn. oist open ground. Circumneutral. Local: St, Louis Co. de erm sylvatica Torr. See Rh. 32: 118. 1930. ergia umbrosa Serib FE rocky woods and slopes along streams. Circumneutral. General. Muhlenbergia sylvatica f. attenuata (Scribn.) Palmer € Steyermark, comb. nov. gia wmbrosa f. attenuata (Scribn.) Deam, Publ. Indiana Dept. Conserv. 82: 171. 1 Oceasionally found in aliia situations to the preceding. Scattered: Gentry, St. Louis, and Greene counties. Muhlenbergia foliosa Trin. Thickets and open IR and moist ground. Cireumneutral Western Mo.: Jackson and Jasper countie Muhlenbergia Schreberi J. F. Gmel. Drop-seed Grass, Nimble Will. Moist open and waste ground; often about dwellings. Circumneutral. General. Eam curtigetosa (Beribn. ) Bush. ia Sc var. curtisetosa Scribn. ae on Cireumneutral. Southwestern Mo.: Barry Co. € mn. Bush, Am. Midl. : 41. 1919. Prairie ban low moist woods. a Southwestern Mo.: RN ud silia counties. Muhlenbergia capillaris (Lam.) Trin. Hair Gra ry open ground and cherty woods. B ai: Southern Mo.: Shannon, Ozark, Taney, Henry, Barry, Greene, Jasper, Newton, and McDonald counties. [ Vor. 22 468 ANNALS OF THE MISSOURI BOTANICAL GARDEN Muhlenbergia I (DC.) Trin. ons Hitcheock, Man. Grasses U. S. Misc. Publ. U. S. Dept. Agr. 200: 386, MuMebbergia trichopodes pom Dry roeky ground. Cireumneutral. Southwestern Mo., loeal: MeDonald Co. SPOROBOLUS PoigETII Roem. & Schult. See Bartonia 14: 32. 1932, and Am. Jour. ot. 21: 136. 1934, Smut Grass. Sporobolus Berteroanus (Trin.) Hitche. € Chase. Sporobolus indicus of auth., not R. Br. (G). Introdueed in cotton fields, Cireumneutral. Southeastern Mo.: Dunklin Co. Sporobolus vaginiflorus (Torr.) Wood. Prairies, fields, and waste places, sterile soils. Oxylophile. General. — e neglectus Nash. open woods, glades, fields, and waste places. Oxylophile to eireum- a. General but scattered: St. Louis, Jefferson, Shannon, Montgomery, Boone, Macon, Falak Henry, Jasper, Barry, Stone, and Ozark counties. Sporobolus ozarkanus Fernald, Rh. 25: 109. 1933. Chert glades. Oxylophile. Southwestern Mo., local: Jasper Co. Sporobolus asper (Michx.) Kunth. Bush Grass. ry prairies, rocky open woods and glades, and waste ground. Oxylophile to circumneutral. General but — St. Louis, Franklin, Macon, Sullivan, Boone, Cooper, Henry, Caldwell, Jackson, Jasper, and Newton counties. — asper var. pilosus po Hitehe, Proc. Biol. Soc. Wash. 41: 161. 928. Sporobolus pilosus Vasey (B € B). Rocky open P ro dace Southern Mo.: Jefferson, Washington, Iron, Jasper, and Barry co Sporobolus asper var. Hookeri (Trin.) Vasey. I a Man. Grasses . S. Misc. Publ. U. S. Dept. Agr. 200: 398, 958. Sporobolus Drummondii dip Vasey (B & E. Sporobolus attenuatus Nash Rocky open woods, glades, = prairies. Oxylophile to eireumneutral. Cen tral, western, and southern Mo.: St. Louis, Phelps, Jackson, Caldwell, and Jasper countie Minis clandestinus (Spreng.) Hitche. Sporobolus canovirens Nash (G), (B & B). Rocky open woods, glades, open ground, and prairies. Oxylophile. East- central ag gemis Mo.: Lineoln, Boone, Montgomery, Warren, St. Louis, Jefferson, . Genevieve, St. Francois, Washington, Crawford, Iron, Scott, Dunklin qe Dent, Shannon, Ozark, Taney, Barry, Jasper, Newton, and McDonald countie Sporobolus einen Gray. Dropseed Gra Prairies, glades, and rocky cliffs. Onlcipátis to circumneutral. General, though appare My absent from much of the southwestern part: Clark, Lewis, Marion, Ralls, Pike, St. Louis, Jefferson, Washin , St , Madison, Pi Francois 0 Tron, Reynolds, Carter, Shannon, Texas, Dent, Phelps, Maries, Pulaski, Cal- laway, Shelby, Schuyler, Randolph, Macon, Linn, Sullivan, Worth, Daviess, Dekalb, Clinton, Caldwell, Jackson, Johnson, Henry, and Taney counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 469 Sporobolus argutus (Nees) Kun ry open ground. EA in Jackson Co. Sporobolus cryptandrus (Torr.) Gray. Sand Dropseed. Dry or moist open ground, usually in sandy soil, and loess mounds, also intro- duced along railroads. Oxylophile. Central and western Mo., mostly along the larger rivers: St. Louis, Jefferson, Montgomery, Boone, Monlitexn, Atchison, Holt, Andrew, Ba Clay, and Jackson counties. Sporobolus airoides Torr. Alkali Sacato oess hills. Caleiphile. Netia en Mo.: Atchison Co., and also intro- duced in Jackson Co HELEOCHLOA SCHOENOIDES (L.) Host. Introduced in meadows. Local: St. Louis Co. Brachyelytrum erectum (Schreb.) Beauv. Roeky woods and bluffs. Veo: M to calciphile. General, but eom- monest in southern Mo. Oryzopsis racemosa (Sm.) Ricker. Black-seeded Mountain Ric Woods ea: Eastern Mo., local: Clark and ^u counties. Stipa en Trin. De Grass. Dr dre iud Oxy ile. Northern and western Mo., uncommon: Macon, pd Schu PX Daviess, Dekalb, Nodaway, Jaekson, Barton, and Jasper e. p reported from Greene Co Aristida basiramea Engelm. Triple-awned Gras Dry open ground. Oxylophile. Central Mo, local: Montgomery and Jack- son counties. Aristida dichotoma Michx. Poverty Gra ry prairies, | ae open upland olli and waste ground. Oxylophile. General and com ee aires TN Nash. See Contr. U. S. Nat. Herb. 22: 535—536. 1924. istida dichotoma var. Curtissü Gray In rand situations to the preceding. Oxiviophile, Central and southern s, Jefferson, Madison, Crawford, Montgomery, Polk, Barry, and Newton SU ipie oligantha Michx. Triple-awned Grass Dry open ground, glades, sterile fields, P waste ground. Oxylophile. General. Aristida ramosissima Engelm. Branching Poverty Grass. Dry prairies, glades, and sterile soils. Oxylophile. oe and southern Mo.: St. Louis, St. Francois, Dunklin, Butler, and Greene e Aristida longespica Poir. See Contr. U.S. Nat. Herb. 22: 538-539. 1924. Aristida gracilis Ell. (G), (B & B Rocky prairies, EPET diario fields, add waste ground. Oxylophile. Eastern, central, and southern Mo.: Marion, St. Louis, Jefferson, St. Francois, Ste. Genevieve, Dunklin, bee Boone, Cooper, Jackson, Greene, Barry, Jasper, and McDonald counties. [Vor. 22 470 ANNALS OF THE MISSOURI BOTANICAL GARDEN Aristida adscensionis L. See Contr. U. S. Nat. Herb. 22: 541-545. 1924. Aristida fasciculata Torr. (B & B). Aristida bromoides HBK. Upland prairies and sterile open ground. Oxylophile. Central and southern Mo.: Maries, Madison, Jackson, and Jasper counties. Aristida intermedia Scribn. & Ball. ry rocky woods, glades, and sterile prairies. Oxylophile. Central and southern Mo.: St. Louis, Washington, Iron, Jackson, and Jasper counties. Aristida lanosa Muhl. Sandy open ground. Oxylophile. Southeastern Mo., loeal: Stoddard Co. Aristida purpurascens Poir. Ro € .. hid glades, and open ground. Oxylophile. Central and uther , St. Louis, Jefferson, Ste. Genevieve, Madison, Scott, Stoddard, Dunklin, an Dent, d Montgomery, Polk, Henry, Ozark, Taney, Barry, Jasper, Newton, and d counties. Leptochloa filiformis (Lam.) Beauv. Red yrs Top. Leptochloa attenuata (Nutt.) Steud. (B & B). Leptochloa mucronata Kunth. Fields, low cultivated ground, and alluvial ground along the larger rivers. Cireumneutral. Central and southern Mo.: St. Charles, St. Louis, Jefferson, Perry, Mississippi, Dunklin, iin Ripley, Crawford, Carroll, Jackson, Greene, Ozark, Taney, Vernon, Barton, Jasper, and Newton counties, Diplachne acuminata Nash. Wet open ground along the Missouri River, and wet soil about saline springs. Caleiphile to eireumneutral. Western Mo.: Atchison, Holt, Jackson, Carroll, Saline, Randolph, Howard, Cooper, and Jaekson counties. Diplachne fascicularis (Lam.) Beauv. Leptochloa fascicularis (Lam.) Gray (G Wet open ground along the larger streams, saline springs, and depressions in rocky glades. Caleiphile to eireumneutral. Central and southern Mo.: St. Louis, Jefferson, New Madrid, Ripley, Randolph, Saline, Carroll, Clinton, Holt, Buchanan, Clay, Jackson, Greene, Jasper, and Newton counties. ELEUSINE INDICA (L.) Gaertn. Goose Grass, Yard Grass. Waste and cultivated ground. Indifferent. General and common, CYNODON DACTYLON (L.) Pers. Bermuda Grass. Open, waste, sandy, and cultivated ground. Oxylophile to eireumneutral. Central pu southern Mo.: St. Charles, St. Louis, Jefferson, Mississippi, Dunklin, Butler, Boone, Jackson, Vernon, and Jasper counties Schedonnardus paniculatus (Nutt.) Trel. Tumble Grass. Dry open lg ein glades, and waste ground. Oxylophile to eireum- neutral. Middl d western Mo.: Boone, Andrew, Wright, Greene, Jackson, Jasper, and Newton counties. Beckmannia Syzigachne (Steud.) Fernald, Rh. 30: 27. 1928. American Slough Grass. Beckmannia erucaeformis of Am. auth. in part, not Host (G), (B & B). Wet open ground. Introduced in Jackson Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 471 Spartina pectinata Link, Sec Rh. 35: 258-260. 1933. Slough Grass. Spartina Michauxiana Hitche. in part (G). Wet meadows, sloughs, and margins of ponds. Cireumneutral to calciphile. General. Spartina pectinata var. Suttiei (Farwell) Fernald, Rh. 35: 260. 1933. Commonly found with the typical form but more frequent. Gymnopogon ambiguus (Michx.) BSP Roeky open woods, sandy open adi, and glades. Oxylophile. Southern and east-central Mo.: St. Charles, Jefferson, Scott, Stoddard, Shannon, Dent, Craw- ford, Jasper, Newton, and MeDonald counties. Chloris verticillata Nutt. Windmill Grass. Upland prairies and waste ground; introduced from farther west. Middle and western Mo.: Boone, Jackson, and Jasper counties. Chloris virgata Sw Chloris sui die HBK. (G), (B & B). Introduced in Jackson Co. Bouteloua curtipendula (Michx.) Torr. Rocky open woods and glades. Cireumneutral to ealeiphile. General. Bouteloua gracilis (HBK.) Lag. See Contr. U. S. Nat. Herb. 14°: 375. 1912. Mesquite Grass, Blue Grama. Bouteloua oligostachya (Nutt.) Torr. (G), (B land prairies and loess hills. Cir Auri ede to ieie. Western Mo.: Atchison, Holt, and Jackson counties Bouteloua hirsuta Lag. Hai y Mesie Grass, Hairy Grama Loess hills. Caleiphile. Nor Mo.: Atchison = Holt eounties. Buchloé E (Nutt.) Engelm. Buffalo Grass ills. Caleiphile. Northwestern Mo.: Atchison and Holt counties, and ina introduced in Jackson Co. ANTHOXANTHUM ODORATUM L. Sweet Vernal Grass. Introdueed along railroad: Jackson Co., and also reported from St. Louis Co. PHALARIS CANARIENSIS L. Canary Grass Waste ground, and along railroad and roadsides. Indifferent. Scattered: St. Louis, Madison, Jackson, and Jasper counties Phalaris arundinacea L. Reed Canary Grass. Wet meadows and ditches along roadsides, Circumneutral. Northern and western Mo., uncommon: Macon, Holt, Andrew, Clay, and J ackson counties. diac caroliniana Walt. Canary Grass. ist open ground. Circumneutral. gar and southern Mo., scattered and uncommon: Scott, Dunklin, Butler, Jackson, Johnson, Greene, and Jasper counties. Leersia lenticularis Michx. Catch-fly Grass Wooded swamps and borders of Ms pe streams. Circumneutral. Gen- eral, but commonest in southern Mo.: Lincoln, St. Louis, Jefferson, Mississippi, New Madrid, Dunklin, ti a cae and Jasper counties. Leersia oryzoides (L.) Sw. Rice Cut Gra amps and wet meadows. irem General. (VoL. 22 472 ANNALS OF THE MISSOURI BOTANICAL GARDEN Leersia virginica Willd. Cut Grass, White Grass Swamps, wet woods, and low meadows. Circumneutral. General. Zizania aquatica L. var, interior Fassett, Rh. 26: 158. 1924. Water Rice, Indian Rice, Swamps and borders of ponds and streams. Cireumneutral. Scattered and uncommon: St. Charles, New Madrid, Dunklin, Butler, Clay, Jaekson, Greene, and Newton counties. TN miliacea (Michx.) Dóll. & Asch. ampy ground and borders of ditches. Cireumneutral. Southeastern Mo.: "bienes and Dunklin eounties, DIGITARIA SANGUINALIS (L.) Scop. Crab Grass, Finger Gras Waste and cultivated ground. Often a troublesome "eal. Indifferent. General, DIGITARIA SANGUINALIS var, MARGINATA (Link) Fernald, Rh. 22: 103, 1920. Syntherisma marginata (Link) Nash (B & B). Oceasionally found with the typical form. Scattered: Dunklin Co. DIGITARIA ISCHAEMUM Schreb. See Rh, 18: 231. 1916. Small Crab Grass. Digitaria humifusa Pers. (G Syntherisma linearis (Kroek.) Nash (B & B). Waste and eultivated ground, Indifferent. General. Digitaria filiformis (L.) Koeler. Slender Finger Grass Prairies, gs and sandy fields. Oxylophile to eireumneutral. Central and southern Mo.: Boone, Gasconade, Dent, Maries, Madison, Jackson, Jasper, Newton, McDonald, and Stone counties. Digitaria filiformis var. villosa (Walt.) Fernald, Rh. 36: 19-20, 1934. Digitaria villosa (Walt.) Ell. (G). Prairies and glades. Oxylophile. Southern and east- pd Mo.: St. Louis, St. Francois, Iron, Dunklin, Jasper, and McDonald e Leptoloma cognatum (Schultes) Chase. Fall Witch Grass. Prairies, glades, and waste ground. Oxylophile, General. Eriochloa contracta Hitche. Dotted Millet, Prairie Cup Grass. Eriochloa punctata of auth., not Hamilton (G), (B & B). Introduced in cultivated and waste ground. Uncommon. Cireumneutral. Boone and Jackson counties. Paspalum repens Bergius. See Contr. U, 8. Nat. Herb. 28: 31-32. 1929. Paspalum mucronatum Muhl. (G), (B 1 B). Wet alluvial ground along streams and margins of ponds and lakes. Cireum- neutral. Eastern, central, and southern Mo.: Marion, St. Louis, Jefferson, New Madrid, Dunklin, Butler, Howell, Chariton, J ackson, Johnson, Vernon, Jasper, and Barry counties. Paspalum dissectum L. uddy banks of streams and ditches. Cireumneutral to oxylophile. South- ern Mo., scattered: Dunklin and Jasper counties. Paspalum ciliatifolium Michx. Prairies and borders of woods. Oxylophile to cireumneutral. Southwestern J 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 473 Paspalum ciliatifolium var. stramineum (Nash) Fernald, Rh. 36: 20. 1934, Paspalum stramineum Nash (G), (B & B Paspalum Bushii Nash (G). Prairies and open woods. Oxylophile. General: Putnam, Jefferson, Dent, Pemiscot, Dunklin, Clay, Jackson, and Polk counties. Paspalum ciliatifolium var. Muhlenbergii (Nash) Fernald, Rh. 36: 20. 1934. Paspalum Muhlenbergii Nash (G), Ps & B). Paspalum pubescens Muhl. (G), (B & B). Prairies, dud banks, and open duin Oxylophile to eireumneutral. General: Putnam, Warren, St. Louis, Jefferson, Franklin, Ste. Genevieve, St. Franeois, Mississippi, bei Dunklin, Oregon, Pulaski, Boone, Saline, Jackson, Barton, Jasper, and Greene counties. Paspalum floridanum Michx. Moist meadows and prairie swales, Oxylophile to circumneutral. Southern Mo.: Mississippi, Barton, and Jasper counties. Paspalum pubiflorum Rupr. var. glabrum ipo See Rh. 36: 22, 1934. Paspalum laeviglume Een (G), Moist open ground, meadows, and ES swales; sometimes a weed in waste ground. Circumneutral. Southern Mo.: Jefferson, Ste. Genevieve, Dunklin, Butler, Taney, Jasper, and McDonald counties. Paspalum laeve Michx. var. circulare (Nash) Stone. See Rh. 36: 22. 1934, Paspalum circulare Nash (G), (B & B Prairies, fields, and moist open ground. Cireumneutral to oxylophile. Central and southern Mo.: Lincoln, St. Louis, Jefferson, Seott, Mississippi, New Madrid, Dunklin, Butler, Randolph, Boone, Callaway, Dent, Shannon, Oregon, Texas, Howell, Wright, Greene, Jaekson, Barton, Jasper, and MeDonald counties. Paspalum laeve var. pilosum Scribn. See Rh. 36: 22. 1934. Paspalum plenipilum Nash (G), (B & B). Paspalum praelongum Nash. Low ground, and borders of lakes and bayous. Cireumneutral. Southern Mo., seattered: New Madrid and Wright counties. Panicum depauperatum Muhl. See Rh, 23: 194. 1921. Prairies, glades, open sterile woods, and waste ground. Oxylophile. Gener al, but commonest in central and southern Mo.: Se ell Lincoln, St. Louis, Jef- ferson, St. Francois, Ste. Genevieve, Bollinger, Wayn hannon, Ripley, E right, Webster, Ozark, Stone, Jasper, Newton, and ‘McDonald counties. Panicum perlongum Nas cky prairies, Jut and open woods, chiefly in ehert, gravel, or sand. Oxy- lophile. General. Panicum linearifolium Seribn. Prairies, glades, and rocky open woods. Oxylophile. General. Panicum linearifolium var. Werneri (Scribn.) Fernald, Rh. 23: 194. 1921. Panicum Werneri Seribn. (G), (B Rocky prairies, glades, and open woods. Oxylophile. Southern Mo.: Ste. Genevieve, St. Francois, Foti Ozark, Douglas, Webster, Christian, Taney, and Jasper counties. [Vor. 22 414 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panicum xalapense HBK Low woods and wet rocky or sandy banks. Oxylophile to circumneutral. Central and southern Mo.: Boone, St. Louis, Franklin, Jefferson, St. Francois, Ste. Genevieve, Bollinger, Iron, Wayne, Dunklin, Ripley, Texas, Douglas, Taney, Stone, and MeDonald counties. Panicum Bicknellii Nash. See Contr. U. S, Nat. Herb. 15: 177. 1910. Panicum Bushii Nash (B & B). Dry rocky woods and prairies, Oxylophile to eireumneutral. Southern Mo.: Dent, Wayne, Bates, Christian, Barry, and McDonald counties. Panicum microcarpon Muhl. Wet woods and low open ground. Oxylophile to cireumneutral. Southern Mo.: St. Francois, Madison, Jefferson, Ste. Genevieve, Perry, Bollinger, Dunklin, Pemiseot, Butler, Ripley, Barry, and MeDonald counties. Panicum nítidum Lam. wet ground. D Southeastern Mo.: Carter Co. (according to Hitcheoek & Chas l Panicum annulum pe Dry woods. Oxylophile to eireumneutral. Southeastern Mo.: Carter Co. Panicum dichotomum L Rocky open woods. Oxylophile. Southern and east-central Mo.: Warren, St. Louis, Jefferson, St. Francois, Madison, Bollinger, Iron, Dunklin, Wayne, Ripley, Oregon, Crawford, Dent, Shannon, Texas, Miller, Morgan, Laclede, Hickory, Douglas, Taney, Greene, Barry, and Jasper counties, x. Rocky open woods and glades. Oxylophile. Southern Mo.: Jeffers son, Ste. Genevieve, Perry, Bollinger, Wayne, Iron, Dunklin, Ripley, Carter, Shanno Oregon, Gasconade, Texas, Howell, Christian, Taney, Barry, and MeDonald eounties Panicum m" Schult. Alluvial woods, prairies, and sandy open ground. Cireumneutral. Southern Mo.: New Madrid and Jasper counties Panicum lanuginosum Ell, var. Lindheimeri (Nash) Fernald, Rh. 36: 77. 1934. Panicum Lindheimeri Nash (G), (B & B). Sandy or sterile woods and prairies. Oxylophile to eireumneutral General. Panicum lanuginosum var. septentrionale Fernald, Rh. 36: 77. 1934. Panicum tennesseense Ashe, in part (G), (B & B Panicum Lindheimeri var. septentrionale (Fern.) Fernald, Rh. 23: 227. 1922, Dry open woods and barrens. í— — and southern Mo.: Boone, Miller, Madison, Shannon, and Taney cou Panicum lanuginosum var. fasciculatum mans Fernald, Rh. 36: 77. 1934. Panicum huachucae Ashe (G), (B & B) Panicum tennesseense Ashe, in part (G), (B & B). Panicum huachucae var. silvicola Hitehe. & Chase (G). Panicum Lindheimeri var. fasciculatum (Torr.) Fernald, Rh. 23: 228. 1922. Dry open woods. Oxylophile to circumneutral. General. 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 475 Panicum lanuginosum var. implicatum (Scribn.) Fernald, Rh. 36: 77. 1934. Panicum implicatum Seribn. (G), (B & B). Panicum Lindheimeri var. implicatum (Seribn.) Fernald, Rh. 23: 228. 1922. Wet woods and prairie swales. Oxylophile to cireumneutral. Eastern, central, and southern Mo.: Schuyler, Knox, Boone, St. Louis, Jefferson, Ste. Genevieve, Dent, Shannon, Hickory, Polk, and Jasper counties. Panicum subvillosum Ashe. Dry rocky woods. Oxylophile. Southwestern Mo.: Jasper and Newton counties. Panicum praecocius Hitche. € Chase. Dry open woods, prairies, glades, and thickets. Oxylophile to eireumneutral. General. Panicum villosissimum Nash. ry open woods and thickets, usually in cherty, sandy, or granitie soils. Oxylophile. East-central and southern Mo.: Montgomery, Warren, St. Louis, Jefferson, Dunklin, Ripley, Carter, Crawford, Shannon, Texas, Christian, Barry, and MeDonald counties. Panicum sphaerocarpon Ell. Dr en woods, glades and prairies, and alluvial or moist ground along streams, and th ickets, ES rocky or sandy ground. Oxylophile. Central and southern Mo.: Lincoln, St. Louis, Jefferson, Franklin, Washington, Ste. Genevieve, Perry, Du St. Francois, Madison, Iron, Dent, Shannon, Carter, Phelps, Texas, Morgan, Douglas, Ozark, Taney, Boone, Hickory, Polk, St. Clair, Cedar, Jackson, Jasper, Newton, McDonald, and Barry counties. Panicum sphaerocarpon var. inflatum (Seribn. & Smith) Hitehe. Rh. 13: 68. 1911. Moist sandy open ground. Oxylophile. Southeastern Mo.: Shannon Co. Panicum polyanthes Schultes. ow wet woods, banks of streams, and prairie swales. Oxylophile to circum- neutral. Eastern and middle Mo.: Lincoln, St. Charles, St. Louis, Franklin, Ste. Genevieve, Perry, Bollinger, Iron, Mississippi, Dunklin, Butler, Ripley, Craw- ford, Boone, and Phelps counties. Panicum malacophyllum Nash. Open sandy woods. Oxylophile. Local: Johnson and Jasper counties. Panicum oligosanthes Schultes var. Scribnerianum (Nash) Fernald, Rh. 36: 80. 1934 Panicum Scribnerianum Nash (G), (B & B Dry open woods, barrens, rocky banks, and dry prairies. Oxylophile. General. noe N ag var. Helleri (Nash) Fernald, Rh. 36: 80. 1934. m Helleri Nash (B & B). itia open woods and prairies. Oxylophile. Western Mo.: Jackson, Jasper, Newton, and MeDonald counties Panicum Ravenelii Scribn. € Merr. Dry ground. Circumneutral to oxylophile. Southern Mo.: Ste. Genevieve, Bollinger, Dent, Shannon, Ripley, Texas, Polk, Cedar, Taney, Barry, and Donald counties, [VoL. 22 476 ANNALS OF THE MISSOURI BOTANICAL GARDEN Panicum Leibergii (Vasey) Seribn. Prairies and open woods. Circumneutral to oxylophile. Central and southern Mo.: St. Louis, Phelps, Shannon, and Jackson counties. Panicum sgun Lam. swales and banks, chiefly on wet sands. Oxylophile. Southern Mo.: Mississippi, Howell, Henry, Vernon, Barton, and Jasper counties. Panicum commutatum Schultes. Open or alluvial rich woods and thickets. Cireumneutral. Southern Mo.: Bollinger, Dunklin, Pemiscot, er Ripley, Ste. Genevieve, Carter, onte: Taney, Stone, and MeDonald eounties. Panicum commutatum var. Ashei (Pearson) Fernald, Rh. 36: 83. 1934. Panicum Ashei Pearson (G), (B & B Dry open woods and rocky banks. Cireumneutral to oxylophile. Southern Mo.: Ripley, Shannon, Texas, Douglas, Taney, Stone, Christian, and MeDonald counties. Panicum clandestinum Open upland "i and thickets. Circumneutral to oxylophile. General. Panicum latifolium L. Rich or rocky woods and thickets. Circumneutral. General. Panicum Boscii Poir. Panicum Porterianum Nas Dry open woods and thickets, Cireumneutral to oxylophile. General. Panicum Boscii var, molle (Vasey) Hitche. & Chase. See Contr. U. S, Nat. Herb. 15: 319-320. 1910. Panicum pubifolium Nash. In similar situations to the typical form. Southern Mo.: Shannon, Wright, Taney, Barry, and McDonald counties. Panicum dichotomiflorum Michx, Cultivated and waste ground, sometimes on glades and in alluvial soils, often a weed on dumps and ballast. Indifferent. General Panicum flexile (Gattinger) Seribn, Glades and rocky open ground. Oxylo — to eireumneutral. Central and southern Mo. and locally north in Gentry Co.: St. Louis, Jefferso on, Franklin, Ste. Genevieve, Dunklin, Crawford, Dent, Reynolds, Shannon, Marin, Pulaski, Camden, Boone, Randolph, Jackson, Polk, Greene, Ozark, Taney, Barry, Jasper, McDonald, and Gentry counti Panicum Gattingeri Nash. Dry rocky ground and barrens, and sometimes in wet ground along stream ncois, iin Shannon, Boone, Jackson, Greene, Taney, Jasper, and "MeDonald pha Panicum philadelphicum Bernh. Glades and rocky open ground. Cireumneutral. Southern and east-central 0.: St. Louis, Jefferson, St. Francois, Dent, duse Wayne, Maries, Texas, Howell, Henry, Jasper, and Newton countie 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 477 Panicum capillare L. Witch Gra Open eultivated, waste erg alluvial ground. Indifferent. General and common Panicum capillare var. occidentale Rydb. See Rh. 21: 111. 1919. Occasionally found with the typical form: Putnam, Macon, Marion, Ralls, Pike, St. Louis, Cole, and Jackson counties. PANICUM MILIACEUM L. Broom-corn Millet. Sparingly escaped in cultivated and waste ground. Western Mo.: Jackson Co. Panicum virgatum L. Switch Grass. Prairie swales, wet open woods, moist glades and ledges, and gravel bars along streams. Indifferent. General Panicum agrostoides Spreng. Wet prairies, open woods, and borders of sloughs and ponds. Circumneutral. Central and southern Mo.: Lincoln, St. Louis, St. Charles, Franklin, Jefferson, Stoddard, New Madrid, Dunklin, Butler, Wayne, Oregon, Shannon, Gasconade, Howell, Jackson, Vernon, Greene, and Jasper counties. Panicum stipitatum Nash. Panicum elongatum Pursh, not Salisb. Wet banks. Circumneutral. Southeastern Mo.: Wayne, Shannon, and Dunk- imn counties. Panicum anceps Michx. Moist sandy ground, wet prairies, borders of streams, and low open woods. Oxylophile. Central and southern Mo.: St. Lo ouis, Jefferson, Franklin, St. Francois, Iron, Perry, Cape Girardeau, Seott, Dunklin, Butler, Ripley, Craw- ford, Shannon, Pulaski, Barton, Greene, and Jasper counties. Panicum hians Ell. See Rh. 13: 67. 1911. Steinchisma hians (Ell.) Nash (G), (B & B). Swamps and wet woods. Oxylophile. a Mo.: Dunklin, Butler, and Ripley counties. Panicum obtusum HBK. Vine Mesquite. Introdueed in sandy soil along the Missouri River. Western Mo.: Jackson Co. zen CRUS-GALLI (L.) Beauv. See Rh. 23: 58. 1921. Barnyard Grass. open and eultivated ground. er General but not common: Clark, Beo, and Jackson counties. This has often been confused with Echino- chloa pungens and varieties. DR CRUS-GALLI f. LONGISETA (Trin.) Farwell. Alluvial and waste ground. EM Seattered in central Mo.: St. Charles, St. Louis, Miller, and Jackson eounti — pungens (Poir. Rydb. See Rh. 37: 137. 1935. Barnyard Grass. E chloa muricata (Michx.) Fernald. mi unten, and alluvial rn borders of streams, ponds, and sloughs, and thiekets. Indifferent. General and common. rer pungens var. occidentalis (Wiegand) Fernald & Griscom, Rh. 37: som) occidentalis (Wiegand) Rydb. (R). [Vor. 22 478 ANNALS OF THE MISSOURI BOTANICAL GARDEN aste, cultivated and alluvial ground. Indifferent. General: Putnam, Schuyler, Lincoln, Marion, St. Louis, Jefferson, Saline, and Taney counties. Echinochloa pungens var. Minds. (Wiegand) Fernald & Griscom, Rh. 37: 137. 1935. shee muricata var, microstachya oa. nochloa microstachya (Wiegand) Rydb. (R) In tain situations to the preceding. Indifferent, and Holt countie Echinochloa pungens var. ludoviciana (Wiegand) Fernald & Griscom, Rh. 37: 137. 1935. Echinochloa muricata var. ludoviciana Wiegand. Alluvial ground. Circumneutral. Scattered in central Mo.: and Jaekson counties. ECHINOCHLOA COLONUM (L.) Link. Jungle Rice. ground. Cireumneutral, Southeastern Mo., local: Pemiscot Co. Yellow Foxtail Grass. Western Mo.: Jackson St. Louis, Clay, SETARIA LUTESCENS (Weigel) Hubbard, Rh. 18: 232. 1916. Setaria glauca (L.) Beauv. (G), (B & B). Waste and cultivated ground. Indifferent. General and common. Setaria geniculata Beauv. Prairie Foxtail Grass. Setaria imberbis R. & S. (G), (B& B Prairies, usually in sandy or cherty soil. “Cason le. General but scattered : Putnam, Randolph, Linn, St. Louis, Mississippi, Dunklin, Shannon, Phelps, Hickory, Vernon, Barton, and Jasper counties, SETARIA VERTICILLATA (L.) Beauv. Foxtail Grass. Waste and cultivated ground. Circumneutral. General but less common than the following. SETARIA VIRIDIS (L.) Beauv. Green Foxtail Grass. e and cultivated ground. Indifferent. General and common. SETARIA ITALICA (L.) Beauv. Italian Millet. Escaped from cultivation, in waste ground and along railroads and roadsides. Seattered SETARIA ITALICA var. GERMANICA (Mill.) Richter. Hungarian Millet. Occasionally escaped from cultivation, in waste ground and along railways and roadsides. Scattered. Cenchrus pauciflorus Benth. See Contr. U. S. Nat. Herb. 22%: 67. 1920. Bur Grass, Sand-burs. Cenchrus carolinianus of some auth., not Walt. (G), (B & B). aste ground and along roadsides and railways. Oxylophile. General but scattere Erianthus strictus Baldw. Beard Grass. Swampy ground and wet open woods. Dunklin Co Erianthus alopecuroides (L.) Ell. Beard Grass. Erianthus divaricatus (L.) Hitehe. (G), (B & B). Circumneutral, Southeastern Mo.: See Am. Jour. Bot. 21: 139. 1934. Woolly 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 479 Moist rocky open woods and thickets. Oxylophile. Southern Mo.: Iron, Shannon, Carter, Dunklin, Ozark, Taney, Christian, Stone, and Barry counties. Eres HISPIDUS (Thunb.) Merr. var. CRYPTATHERUS (Hack.) Honda. Moist grassy banks. Cireumneutral Introduced in St. Louis Co. Andropogon scoparius Michx. var. genuinus Fernald & Griscom, Rh. 37: 144. 1935. Blue-stem, Prairie Beard Grass ndropogon scoparius var. ela Kea Prairies, glades, open and waste ground. Nue General. Andropogon scoparius var. frequens Hubbard, Rh. 19: 103. 1917. Similar situations to the preceding. Oxylophile. General. Andropogon scoparius var. divergens Hack. See Rh. 37: 144. 1935. Prairies, glades, and sandy open ground. Oxylophile. Eastern Mo.: Lewis, St. Louis, and Scott counties. Andropogon virginicus L. var. genuinus Fernald & Griscom, Rh. 37: 142. 1935. Old Field Beard Grass, Broom Sedge. Prairies, glades, old fields, and waste ground. Oxylophile. Central and southern Mo. south and east of a line drawn from Lincoln, Boone, and Cooper counties to Henry and Vernon counties. Andropogon virginicus var. tetrastachyus (Ell) Hack. See Rh. 37: 142. 1935. Sandy open and waste ground. Oxylophile. Southeastern Mo.: Madison and Mississippi counties. Andropogon eta Lam. See Rh. 37: 146. 1935. Big Blue-stem, Blue- joint Turkeyfoo Andropogon neh Muhl. (G), (B & B). Prairies, glades, and open rocky woods. Oxylophile to eireumneutral. General. Andropogon provincialis var. chrysocomus (Nash) Fernald € Griscom, Rh. 37: 47. 1935 Andropogon chrysocomus Nash (B & B Prairies and waste ground. Oxylophile. Southwestern Mo.: Jackson, Stone, and Jasper eounties. Andropogon ternarius Michx. Silvery Beard Grass. Prairies and rocky open slopes, usually in sandy or cherty soils. Oxylophile. Southern Mo.: Ste. Genevieve, Madison, Stoddard, Dunklin, Jasper, and Me- Donald MON Andropogon Elliottii Chapm ones big and db open woods. Oxylophile. Southern Mo.: Jeffer- , Madison, Shannon, and Barry counties. Mois Ellioti var. projectus Fernald $ Griscom, Rh. 37: 139. 1935 Similar situations to the preceding. Oxylophile. Southern Mo.: Shannon and Taney counties Andropogon saccharoides Sw. Beard Grass. ndropogon Torreyanus Steud. Amphilophis saccharoides (Sw.) Nash (B & B). Prairies, glades, and rocky ledges. Oxylophile. Southwestern Mo.: Jasper and MeDonald eounties, and also adventive in Jaekson Co (Vor. 22 480 ANNALS OF THE MISSOURI BOTANICAL GARDEN SoRGHUM HALEPENSE (L.) Pers. Johnson Grass. Introduced and established in low meadows and fertile cultivated ground. ten a pernicious weed. Cireumneutral, Scattered but becoming more general. SORGHUM VULGARE L. Sorghum Car Escaped in waste ground and d railways and roadsides. Circumneutral, Scattered. Sorghastrum nutans (L.) Nash. Indian Grass. Prairies, glades, and open rocky woods. Oxylophile. General. ` Manisuris cylindrica (Michx.) Kuntze. See Hitchcock, Man. Grasses U. 8. Mise. Publ. U. S. Dept. Agr. 200: 761 and 881. 1935 Rottboellia cylindrica (Michx.) Torr., not Willd, (G). Coelorachis cylindrica (Michx.) Nach (B&B). Prairies and sandy open ground. Oxylophile. Seattered in southeastern and western Mo.: Mississippi and Jackson eounties. The common cultivated cereals, wheat (Triticum aestivum), rye (Secale sereale), and barley (Hordeum sativum), as well as Indian corn (Zea Mays), are frequently found growing as waifs along roads and railroads and on dumps, but as they do not self-seed nor show any sign of persisting they can scarcely be considered as additions to the flora. Sometimes Zea Mays is found self-seeding in rich or rocky woods where probably carried by man or lower animals, such as squirrels, and appears as a spontaneous part of the flora. Rice (Oryza sativa) is infrequently found as a waif in eultivated or waste ground, as an escape from eultivation. Fam. CYPERACEAE Cyperus flavescens L. Low wet ground. Cireumneutral. Central and southern Mo., mostly in the eastern counties: St. Louis, Jefferson, Washington, St. Franeois, Iron, Madison, Dunklin, ee Shannon, Boone, Miller, and Webster eounties. Cyperus diandrus Tor Low alluvial nn along streams and sloughs. Circumneutral, General. Cyperus compressus L. Coco Gra Swamps and moist eultivated pm Circumneutral. Eastern Mo. south of the Missouri River: St. Louis, Dunklin, and Butler counties. Cyperus rivularis Kunth. Wet open ground. Circumneutral. General. Cyperus Gatesii Torr. Low woods and swamps. Circumneutral. Southeastern Mo., local: Dunklin 0. Cyperus inflexus Muhl. Li in aristatus Rottb. (G depressions in glades, -— and sand bars along streams, alluvial and iuit Hi see gen Oxylophile. General, but commonest in central an southern Mo.: Marion, Ralls, Lincoln, St. Louis, Jefferson, Iron, Madison, Perry, Mississippi, Dunklin, Butler, Marion , Pulaski, Montgomery, Miller, Laclede, oone, Saline, Clinton, > dan J a: nts. Cedar, Greene, Jasper, New- ton, and MeDonald counti 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 481 Cyperus Schweinitzii Torr. Low ground along streams and about EX Cireumneutral. Seattered: St. Louis, Jefferson, and Jackson eounties Cyperus acuminatus Torr. & Hook. Low open ground. Cireumneutral. General. Cyperus pseudovegetus Steud. Wet prairies and borders of Mida: and ponds. Cireumneutral Central and southern Mo.: Boone, New Madrid, Mississippi, Stoddard, Butler, Ripley, Jasper, Newton, and McDonald counties. Cyperus Hallii Britton. Missouri River bottoms. Cireumneutral. Western Mo.: Jackson Co. Cyperus esculentus L. Yellow Nut-grass. t woods, sand and gravel bars along streams, prairie swales, and moist cultivated ground. Cireumneutral. General Cyperus esculentus var. longispicatus Boeckl. Occasionally found with the preceding, in similar situations. Cyperus esculentus var. angustispicatus Britton. w wet ground. Cireumneutral. Jasper Co. Cyperus erythrorhizos Muhl. Including Cyperus erythrorhizos var. pumilus Engelm., merely a dwarf form. erus Halei Torr. Wet woods, low and alluvial ground along streams and lakes. Cireumneutral. General but seattered: Lineoln, St. Louis, Perry, Dunklin, Butler, Putnam, Boone, Saline, Atchison, Holt, Clinton, Dekalb, Clay, Carroll, and Jackson counties. Cyperus ferruginescens Boeckl. See Rh. 37: 150. 1935. Y Linn ferax of auth. in part, not Rich. Cyperus speciosus of auth. in part, not Vahl (B & B). Low E along streams and sloughs, especially the larger rivers. Circum- neutral. General. This has been confused with Cyperus feraz which is known only in southeastern Mo. Cyperus ferax Rich. See Rh. 37: 150. 1935. Swampy woods along St. Francis River. Circumneutral. Ws osi Mo., local: Dunklin Co. This has been confused with Cyperus ferruginesce Cyperus Engelmanni Steud. Low alluvial ground. Cireumneutral. East-central Mo., loeal: St. Louis Co. Cyperus strigosus L. See Rh. 37: 150-151. 1935. Cyperus strigosus var. compositus Britton (G). Cyperus strigosus var. elongatus Britton (G). Cyperus strigosus var. capitatus Britton, wrongly aseribed to Boeckeler (G). Low wet ground, along streams, ponds, and sloughs, sometimes in waste or eultivated soils. Cireumneutral. General, Cyperus strigosus var. robustior Britton. See Rh. 37: 151. 1935. Commonly found with the typical form, in similar situations. [Vor. 22 482 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cyperus refractus Engelm. n woods or open ground, usually in dry sandy or sterile soils, also in low ground along streams, and on gravel bars. Oxylophile. Eastern and southern Mo., south of the Missouri River: St. Louis, Jefferson, Ste. Genevieve, St. Fran- eois, Wayne, Stoddard, Butler, Reynolds, Ripley, Pulaski, Crawford, Dent, Shan- non, Oregon, Howell, and Barry counties. Cyperus lancastriensis Porter. Prairie swales and moist rocky slopes and glades. Oxylophile to cireum- neutral, Southern and east-central Mo.: St. Louis, Madison, Jasper, and New- ton countie Cyperus retrofractus (L.) Torr. Sandy open ground, in woods or fields. Oxylophile. lin, Ripley, and Carter counties. Cyperus dipsaciformis Fernald. Cyperus retrofractus of auth. in part (B& B Sandy open woods or fields. Oxylophile. Southeastern Mo.: Mississippi and Dunklin counties. Cyperus ovularis (Michx.) Torr. Hedgehog Club-rush. Sandy soil along streams, dry open woods, glades, and prairies. Oxylophile to circumneutral. Central and southern Mo. south of a line drawn from St. Louis, Montgomery, and Boone counties to Jaekson Co. Cyperus ovularis var. robustus Boec Oceasionally found with the typical form, in similar situations. Cyperus globulosus Aubl. See Rh. 37: 153-154. 1935. Cyperus echinatus (Ell.) Wood (G). Open woods and prairies. Circumneutral. Southeastern Mo.: Dunklin Co. Cyperus filiculmis Vahl. See Rh, 37: 153. 1935. Cyperus Bushii Britton (B & B), in part. Dry open woods, prairies, and glades. "Hood to oxylophile. General. Cyperus filiculmis var. macilentus Fernald. See Rh. 37: 153. Í Sandy open E glades, and banks of streams. Oxylophile. Eastern, central, and southern Mo.: Shelby, St. Charles, Jefferson, Wayne, Bollinger, Dunklin a Hiekory, Cedar, and Saline counties. Kyllinga pumila Michx. Wet open woods and prairies. Cireumneutral. General but not common. um arundinaceum (L.) Britton. Borders of sloughs and ponds. Cireumneutral. General but uncommon: St. Charles, St. Louis, Iron, Shannon, Howell, Boone, Jackson, Cass, and Newton Southeastern Mo.: Dunk- ndis. Eleocharis —— (Miehx.) R. & + See Rh. 31: 132-133. 1929. ow muddy banks of ponds an eams. Circumneutral. Central and southern Mo., uneommon: St. geh Im, Stoddard, New Madrid, Ripley, Shan- non, Texas, Greene, and Newton cou Eleocharis obtusa (Willd.) Schultes. M Rush. Muddy borders of ponds and streams. Cireumneutral. General and common. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 483 Eleocharis Engelmanni Steud. Muddy borders of streams, sloughs, and ponds. Circumneutral. Easte southern, and central Mo.: Lincoln, St. Louis, Wayne, Iron, Shannon gora Howell, Greene, Jasper, Barry, heb cual and Jackson n. Eleocharis Engelmanni var. robusta Fernald. Occasionally found with the typical form, in similar situations: Iron, Shan- non, Lawrence, and Greene counties. Eleocharis mamillata Lindb. f. See Rh. 31: 66—67. 1929. Spike Rush. Eleocharis macrostachya Britton (B & B). Prairie swales and borders of streams and ponds. Circumneutral. Central and southern Mo.: Lincoln, St. Charles, St. Louis, Franklin, Washington, La- fayette, Jackson, and Jasper counties. Eleocharis lanceolata Fernald. t sandy open ground and glades. Oxylophile to circumneutral. Southern Mo.: Carter, Jasper, and Barton counties. Eleocharis Smallii Britton. See Rh. 31: 64-66. 1929, Wet meadows, prairies, and sloughs. Circumneutral. General. Eleocharis calva Torr. See Rh. 31: 68-70. 1929. Creeping Spike Rush. Eleocharis palustris (L.) R. € S. var. glaucescens of Am. auth, (G), (B € B). t prairies, DENM n and sloughs. Circumneutral. General and common. Electa lucis (L.) R. var. verrucosa Svenson, Rh. 34: 202. 1932. aris tenuis of ln in un (G), (B & B). n postes and swampy open ground. Circumneutral. General, Eleocharis compressa Sulliv. See Rh. 34: 215. 1932. Bleocharis acuminata (Muhl.) Nees (G), (B&B). Wet prairies, depressions in glades, and borders of er s nr and ditches. Calciphile d circumneutral. Eastern, central, and southe : Lincoln, Mont- gomery, St. Louis, Franklin, Jefferson, Washington, Ste. donee rm Texas, Shannon, Mississippi, Miller, Camden, Maries, Pulaski, Laelede, Wright, Greene, Douglas, Vernon, der Barry, and Jackson counties. Eleocharis parvula (R. € 8.) Link. zm Rh. 31: 168. 1929. cirpus nanus Eur (G), (B & B). Marshy ground about saline springs and ponds. Middle Mo.: Cooper and Saline counties. Eleocharis acicularis (L.) R. & S. var. typica Svenson, Rh. 31: 184. 1929. s of ponds and sloughs. Circumneutral. General but seattered: Put- Louis, Jefferson, Franklin, Dent, Carter, Shannon, Howell, Holt, Jackson, Webster, pons Tann and Greene counties. Eleocharis acicularis var. gracilescens Svenson, Rh. 31: 191. 1929. prings and muddy borders of ponds and slou nd Cireumneutral. Central and southern Mo., scattered: St. Louis, Perry, Ripley, Phelps, Laclede, Texas, Ozark, Greene, and Jasper counties. Stenophyllus capillaris (L.) Britton. Prairie swales, glades, and barrens. Oxylophile. General but commonest in Ozark region. B 5 os MES. O Iz = a et [Vor. 22 484 ANNALS OF THE MISSOURI BOTANICAL GARDEN Stenophyllus capillaris var. cryptostachys Fernald, Rh. 19: 154. 1917. andy open woods. Oxylophile. Southern Mo., loeal: Shannon Co. Fimbristylis castanea (Michx.) Vahl. Prairies and wet depressions in glades. Cireumneutral to oxylophile. Central and southern Mo.: St. Louis, Shannon, Benton, Dallas, Cedar, Jaekson, and Jasper counties. Fimbristylis castanea var. puberula (Michx.) Britton. imbristylis puberula (Michx.) Vahl (8). High sandy prairies, been meadows, and glades. Oxylophile to eireum- neutral. Central and southern Mo.: Montgomery, St. Louis, Jefferson, Wash- ington, Franklin, E Phelps, Maries, Madison, Iron, er e: Ozark, — — Hickory, Cedar, St. Clair, Barry, Barton, Jasper, and Newton coun —. = Vahl. is Baldwiniana Torr. (B & B). itm 2 = — and moist plaees in glades, and Fai or sand bars along streams. Oxylophile. Central and southern Mo. Mis- sissippi, Duskiin Cooper, Pine Jaekson, Vernon, Greene, uide Nen: and MeDonald counties. Fimbristylis Vahlii (Lam.) Link Moist sandy open promi.. Oxylophile. Southern Mo., scattered: Wayne, Dunklin, and Barry counties. Fimbristylis mucronulata (Michx.) Blake, Rh. 20: 25. 1918. Fimbristylis autumnalis of Am. auth. (G B) Prairies and sandy fields. Oxylophile to lrctinmen tá. Central and southern t. Francois, Perry, New Madrid, Dunklin, Howell, Jackson, Jasper, and N ein counties, lai autumnalis (L.) R. & S. See Rh. 20: 24. 1918. Fimbristylis Frankii Steud. (G). an geminata (Nees) Kunth (B & B). Sandy fields, sandy open ground along streams, and prairies. Circumneutral to oxylophile. General. Scirpus planifolius Muhl. ry open woods and shaded sandstone ledges. Oxylophile. Southern Mo.: Shannon and Douglas counties. Scirpus americanus Pers. Three-square. Wet woods, sloughs, swamps, ua alluvial banks of streams. Circumneutral to oxylophile. General and common. Scirpus validus Vahl. Great Bulru Wet prairies and borders of ind and ponds. Circumneutral. General. Scirpus acutus Muhl. See Rh. 22: 55-56, 1920. Scirpus occidentalis (Wats.) Chase (G), (B & B). Ponds and sloughs. Calciphile to cireumneutral. Seattered in northern and western Mo.: Adair, Saline, Holt, Buchanan, Andrew, Platte, Jackson, and Greene u, 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 485 Scirpus campestris Britton. Bayonet Gras Sandy open ground. Calciphile to | Northwestern Mo., local: Holt Co. Scirpus fluviatilis (Torr.) Gray. River Bulrush. et ipee: along streams, and borders of ponds and sloughs. Cireumneutral. Genera Scirpus atrovirens Muhl. Common Bulrush. Boggy or swampy open ground. Cireumneutral. General, Scirpus atrovirens : var. georgianus (Harper) Fernald, Rh. 23: 134. 1921. Low wet woods. Circumneutral is oxylophile. Southeastern Mo.: Iron Co. Scirpus pallidus Pen Fernald. Wet open d. Cireumneutral. Western Mo.: Dekalb and Jasper eounties. Scirpus divaricatus Ell. Swamps and low wet woods. Circumneutral. Southeastern Mo.: Dunklin, Butler, and Ripley counties. Scirpus lineatus Michx. Wet prairies and bogs. Circumneutral. General. Scirpus pedicellatus Fernald. Sloughs and ponds. Circumneutral. Northern Mo.: Putnam and Adair counties. Scirpus Eriophorum un wamps and low oods. Cireumneutral. General but seattered: Lineoln, Iron, Stoddard, tbe Butler, Caldwell, and Jackson counties. Scirpus carinatus Gray. Glades, barrens, and sandy open ground. Oxylophile. Southern Mo., rare: Dunklin, Jasper, Newton, and MeDonald counties, Fuirena simplex Vahl. Umbrella grass. Bogs and swampy open ground. si to ealeiphile. Northwestern, eentral, ue southern Mo., mostly in the eastern Ozark region: Washington, . Franeois, Iron, Dent, Wayne, "did Shannon, Texas, Holt, and Jaekson a Hemicarpha micrantha (Vahl) Pax. Sand bars along streams, borders of ponds, and wet depressions in glades. Cireumneutral to oxylophile. General but seattered: Clark, Lincoln, St. Louis, Jefferson, Franklin, Perry, Bollinger, Mississippi, Dunklin, Butler, Wayne, Reynolds, Oregon, Howell, Miller, Pulaski, Holt, Jackson, Jasper, Barry, and Stone eounties. Hemicarpha micrantha var. aristulata Coville. Sandy borders of ponds, lakes, and streams, and wet depressions in glades. Western Mo., scattered: Buchanan, Barry, and Stone counties. Rynchospora corniculata (Lam.) Gray. Horned Rush. Swamps, low wet woods, and along drainage canals. Circumneutral to oxy- lophile. Eastern Mo., mostly in the lowland counties: Audrain, St. Louis, Iron, [Vor. 22 486 ANNALS OF THE MISSOURI BOTANICAL GARDEN Cape Girardeau, Stoddard, New Madrid, Mississippi, Dunklin, and Butler counties, Rynchospora macrostachya Tor Low swampy woods eje Southeastern Mo.: Bollinger, Stoddard, Pemiseot, Butler, and Ripley countie Rynchospora cymosa Ell. Prairie — meadows, and wet rocky banks. Oxylophile to eireumneutral. Southern Madison, Shannon, St. Clair, Vernon, Barton, and Jasper coun- ties. This ci sometimes been til with R. compressa which does not occur in Mo. — capillacea Tor Moist rocky ground and wet ledges along bluffs. Calciphile. Eastern Mo., south » the Missouri River: Jefferson, Washington, St. Francois, Ripley, Shan- non, and Texas counties. Rynchospora capitellata (Michx.) Vahl. See Rh, 20: 27. 1918. False Bog Rush. Rynchospora glomerata of auth., not Vahl (G), (B € B). igh prairies and rocky or sandy banks of streams. Oxylophile. Southern Mo.: St. Francois, Ste. Genevieve, Perry, Madison, Iron, Reynolds, Ripley, Dent, Howell, Shannon, Greene, nis and Jasper counties. — pera Michx. Tall Nut-grass. open woods, glades, ES prairies. Oxylophile. General but seattered: M Lewis, Macon, St. Louis, Jefferson, Iron, Reynolds, Dent, Shannon, Phelps, Howell, Mercer, Linn, Stone, Jackson, Barton, Jasper, Newton, MeDonald, and Barry counties. Scleria pauciflora Muhl. Sandy prairies and barrens. Oxylophile. Southern Mo.: Jefferson, Iron, Shannon, Wright, Ozark, Jasper, and Barry counties. Scleria pauciflora var. caroliniana (Willd.) Wood. Rocky woods and glades. Oxylophile. Southwestern Mo.: Hickory, Jasper, and Newton counties. Scleria oligantha Michx. Rocky open woods and glades. Cireumneutral. Southwestern Mo.: Ozark and Barry counties. Scleria ciliata Michx. Scleria Elliottii Chapm. (G). High sandy prairies and open rocky woods. Oxylophile. Southern Mo.: Franklin, pr. ngton, vinh arter, Shannon, Dent, Taney, Barry, Barton, Jasper, and McDonald countie Scleria sotacos Poi apon Michx. var. pubescens Britton (G). Die ground. Southwestern Mo., local: Greene Co. Scleria verticillata Muhl. Moist sandy open ground. Oxylophile. Southern Mo., scattered: Jefferson and McDonald counties. 1935] PALMER € STEYERMARK——PLAN T$ OF MISSOURI 487 Carex praegracilis Boott. Carez camporum Mack. B). Dry open ground. Cireumneutral. Introduced in Jackson Co. This has sometimes been eonfused with Carez siccata whieh does not oceur in Mo. Carex Sartwellii Dewey. ows, Cireumneutral. Western Mo.: Jackson Co. Carex retroflexa Muhl. z open woods. Cireumneutral. Sag Schuyler, Shelby, Monroe, St. , Jefferson, Franklin, Ste. Genevieve, Wayne, Butler, Dunklin, Shannon, Phelps, Lad Bone, Clay, Jackson, Senne and Bane counties, Carex texensis (Torr.) Bailey. Carex retroflexa var. texensis (Torr.) Fernald (G). Swampy open ground. Cireumneutral. Southeastern Mo.: Cape Girardeau Co Carex rosea Schkuhr. Carex rosea var. minor Boott (G). Dry rocky woods and prairies. Circumneutral. General. Carex convoluta Mack. Bull. Torr. Bot. Club 43: 428. 1916. Carex rosea of auth., not Schkuhr (G), (B & B). Dry rocky woods. Circumneutral. General. Carex cephalophora Muhl. Prairies, swales, meadows, and open woods. Circumneutral. General. Carex Leavenworthii Dewey. Prairies and open rocky woods. Cireumneutral. General. Carex mesochorea Mack. Bull. Torr. Bot. Club 37: 246. 1910. Central and southern Mo.: Lincoln, St. Louis, Jackson, Greene, and Stone counties. Carex arkansana Bailey. rairie swales and open woods. Cireumneutral Southwestern Mo.: Jasper Co. Carex Muhlenbergii Schkuhr. Dry open woods and prairies. Circumneutral. General. Carex 2 var. enervis Boott. ex plana Mack. Bull. Torr. Bot. Club 50: 350. 1923. Be meadows, and open ground. Circumneutral. General. Carex austrina (Small) Mack. eky prairies and glades. Cireumneutral to ealeiphile. Western and south- ern Mo.: Shannon, Harrison, Jackson, Christian, Taney, and Jasper counties. Carex gravida Bailey. Carex gravida var. laxifolia Bailey, in part (G). Prairies and dry open banks. Cireumneutral. General, Carex Lunelliana Mack. Bull. Torr. Bot. Club 42: 615. 1915. Carex gravida var. laxifolia Bailey, in part (G). Prairies and dry open banks. Cireumneutral. General, [Vor. 22 488 ANNALS OF THE MISSOURI BOTANICAL GARDEN Carex aggregata Mack. Bull. Torr. Bot. Club 37: 246. 1910. Prairie swales. Cireumneutral to caleiphile. Central and western Mo.: Wash- ington, Osage, Jaekson, and Jasper eounties. Carex cephaloidea Dewey. ry open qm and hillsides. Circumneutral. Seattered: St. Louis and Jackson counties Carex se poen Muhl. Margins of ponds, bogs, and swampy woods. Cireumneutral. General but scattered: Sehuyler, Putnam, Knox, St. Louis, Jefferson, Washington, Shan- non, Boone, Linn, Harrison, Atchison, Jackson, Dallas, Barton, Jasper, and Lawrenee counties. Carex triangularis Boeckl. Swampy open ground. Oxylophile. Southeastern Mo.: Wayne and Dunklin counties. Carex annectens Bicknell, Bull. Torr. Bot, Club 35: 492. 1908. Carex xanthocarpa var. annectens Bicknell. Carex annectens var. xanthocarpa Wiegand. Carex setacea var. ambigua (Barratt) Fernald, in part (G), (B & B). Carez brachyglossa Mack. Bull. Torr. Bot. Club 50: 355. 1923. Prairies, dry open woods, and open soils. Cireumneutral to oxylophile. eneral. Carex vulpinoidea Michx. Fox Sec Wet prairies, meadows, and en of sloughs. Cireumneutral. General. Carex decomposita Muhl. Swamps and boggy ground. Cireumneutral. Southeastern Mo.: Dunklin, Dent, Shannon, and Texas counties. Carex oklahomensis Mack. Torreya 14: 126. 1914. Prairie swales and borders of ponds and ditches. Cirtumneutral to ealeiphile. Western Mo.: Jackson, Jasper, and Newton counties. Carex stipata Muhl. Prairie swales and borders of ponds and ditehes. Cireumneutral. General but seattered: Shelby, Lineoln, St. Louis, Jefferson, Iron, Madison, Wayne, Butler, Shannon, Boone, Jackson, Jasper, and Newton counties. Carex € (Kükenth.) Mack. in B. & B. Ill. Fl. ed. 2, 1: 371. 1913. t meadows and open bogs. Circumneutral. Eastern Mo., local: St. Louis "d Carex crus-corvi Shuttlw. Low wet =. nm and swampy prairies. Cireumneutral. General. Carex conjuncta Boo Wet m E swamps, and prairie swales. Circumneutral, General: Seot- land, Adair, Sullivan, Boone, St. Louis, Franklin, Howell, Stone, Clay, Jaekson, and Jasper counties. Carex interior Bailey. Carex seirpoides Schkuhr (G). Wet open ground. Circumneutral to oxylophile. Southeastern Mo.: Iron and Shannon eounties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 489 Carex scoparia Schkuhr. Corea deca var. yá di Tuekerm. (G), in and w eadows. Circumn dei pue but seattered: Sat ce s rel St. Louis, Franklin, Howell, Boone, Jackson, Benton, Wright, Greene, and Jasper counties. Carex tenera Dewey. See Rh. 23: 235. 1921. Straw Sedge. Carez straminea of Am. auth., not Willd. (G), (B & B) Dry open woods and prairies. Cireumneutral. General. Carex normalis Mack. Bull. Torr. Bot. Club 37: 244, 1910, Carex mirabilis Dewey, not Host (G). Carex mirabilis var. perlonga Fernald (G), in part. Prairies and open woods. Circumneutral. Genera Carex festucacea Schkuhr. Dry roeky ground, meadows, and prairies. Cireumneutral. General. Carex brevior ieu Mack. Am. Midl. Nat. 4: 235. 1915. Carex molesta Carex pain var. brevior (Dewey) Fernald (G). Dry rocky ground, meadows, and prairies. Circumneutral. General, Carex hormathodes Fernald. n swampy woods and prairies, Circumneutral. Southwestern Mo., local: Jasper Co. Carex projecta Mack. Low swampy ground. Cireumneutral Northeastern Mo.: Adair Co. Carex Bicknellii Britton. Dry open woods and prairies. Circumneutral. General. Carex straminea Willd. l Prairie swales, meadows, and swampy open woods. Circumneutral to oxy- lophile. General. Carex tribuloides Wahlenb. Prairie swales and swampy open woods. Circumneutral. General. Carex tribuloides var. sangamonensis Clokey, Rh. 21: 84. 1919. Oceasionally found with the typical form. St. Louis, Boone, and Jackson counties Carex cristatella Britton. cristata Schwein., not Clairy (G). aes swales, wet woods, and thiekets. Cireumneutral. Northern and een- tral Mo.: qn Adair, Knox, Monroe, Lincoln, St. Louis, Jefferson, Boone, Saline, Clay, and Jackson counties. Carex ub. Sehwein. Wet woods and swampy prairies. Cireumneutral. Northern and central Mo.: Clark, Ralls, St. Louis, Jefferson, Adair, Livingston, and Jackson counties. Carex Jamesii Sehwein. or rieh woods and along base of bluffs and alluvial soils of river banks and ponds and sloughs. Cireumneutral General [Vor. 22 490 ANNALS OF THE MISSOURI BOTANICAL GARDEN Carex physorhyncha Liebm. Low wet woods. Circumneutral. Southeastern Mo., local: Dunklin Co. This has been confused with Carex albicans of some authors and Carex Emmonsii which do not oceur in Missouri, Carex pennsylvanica Lam. var. digyna Boeckl. Carex pennsylvanica of auth. in part (G). Carex heliophila Mack. Torreya 13: 15. 1913. Prairies, dry woods, and rocky open banks. Oxylophile. Northern and central Mo.: Schuyler, St. Louis, Daviess, Clay, and Jackson counties. Carex heliophila cannot be considered as specifically distinet from Carez pennsylvanica, and the Mo. material should be treated as a variety of the latter species Carex artitecta Mack. See N. Am. Fl. 18: 189. 1935. Carex varia Muhl., not Lumnitzer nor "dn (G), (B & B). Carex varia var, peleeats Bailey, in par Dry rocky woods and prairies. REM General. Carex nigro-marginata Schwein. pen woods. Cireumneutral. Southeastern Mo.: Crowleys Ridge, Dunklin o Carex umbellata Schkuhr, Carex microrhyncha Mack. Bull. Torr. Bot. Club 40: 548. 1913. Rocky prairies, dry rocky or upland woods, and glades. Oxylophile. General: known north of the Missouri River in St. Charles, Warren, Montgomery, Calla- way, Boone, Audrain, and Grundy counties. The Missouri material has been referred = Mackenzie to Carez microrhyncha, but this does not appear to be distinet from Carex umbellat Carex hirtifolia Mack. Bull. Torr. Bot. Club 37: 244. 1910. Carex pubescens Muhl., not Poir. (G), (B € B). Rich woods, prairies, and meadows, Circumneutral, General. Carex eburnea Boott. Shaded limestone bluffs, and rarely on sandstone. Calciphile to circumneutral. Eastern, central, and southern Mo., south and east of a line from Lincoln, Mon- roe, Boone, Morgan, Benton, and Hickory counties to MeDonald Co. Carex tetanica Sehkuhr. Carex tetanica var. Woodii (Dewey) Bailey (G). Carex Woodii Dewey Carex colorata Mack. (B € B). Open ground. Cireumneutral. Northwestern Mo.: Atchison Co. Carex Meadii Dewey. Carex tetanica var. Meadü (Dewey) Bailey (G). Upland prairies, glades, and meadows. Caleiphile to eireumneutral. General and common. This has frequently been confused with Carex Crawei. Carex Careyana Torr. Rich or rocky woods. Cireumneutral. East-central and southern Mo.: St. Louis, Shannon, Taney, and Stone counties, Carex digitalis Willd. r rocky open woods and moist ledges. Cireumneutral. Southern Mo.: Jefferson, Dunklin, Shannon, Douglas, Wright, and Taney countie 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 491 Carex "picos Sehw woods. B CIMA Southeastern Mo., loeal: Ripley Co. Carex laxiculmis var. copulata (Bailey) Fernald. Rich or rocky woods. Cireumneutral. Local: Shannon Co. Carex albursina Sheldon. See Rh. 24: 193. 1922. Carex laxiflora var. latifolia Boott (G). Rich woods. Circumneutral. General but not common. Carex blanda Dewey. See Rh, 24: 194. Carez laxiflora var. blanda eee up G). Carez laxiflora var. varians of auth., not Bailey (G). Rich or rocky woods. Cireumneutral. General. Carex laxiflora Lam. See Rh. 24: 195. 1922. arez laxiflora var. gracillima Boott (G). Rich woods and along shaded bluffs. Cireumneutral. General. Carex anceps Willd. Carex laxiflora var. patulifolia To Carey (G). Rich woods and along rocky bluffs and ledges. Cireumneutral. Western Mo.: Jackson, Jasper, Barry, and Stone counties. Carex AER Muhl. ist woods and meadows. Circumneutral. General. Carex granularis var. Haleana i2 Porter. Carex Shriveri Britton (B & B). Moist meadows and low open ground. Circumneutral. Southern Mo., seat- tered: Wayne and Phelps counties. Carex Crawei Dewey. et plaees in glades. RR East-eentral and southern Mo.: Jefferson, Franklin, and Taney counties Carex oligocarpa Schkuhr. Rich rocky woods and ledges along bluffs. Cireumneutral. Central and southern Mo.: St. Louis, Jefferson, Franklin, Iron, Wayne, Shannon, Miller, Laelede, Boone, Clay, Jackson, Taney, Stone, Jasper, Newton, and MeDonald counties. Carex Hitchcockiana Dew Rieh woods and Milk bluffs. Cireumneutral. General but seattered: St. Louis, Washington, Shannon, Mercer, and Jackson counties, and also recorded from Boone Co. Carex amphibola Steud. Carez grisea var. angustifolia Boott (G). Rich alluvial or wet woods. Cireumneutral. Southeastern and central Mo.: St. Louis, Jefferson, Iron, Mississippi, Dunklin, Butler, Shannon, and Jackson counties. Carex grisea Wahlenb. Prairie swales and moist woods. Circumneutral. General. Carex grisea var. rigida Bailey. Occasionally found with the typical form. [Vor. 22 492 ANNALS OF THE MISSOURI BOTANICAL GARDEN Carex grisea var. globosa Bailey. In similar situations to the typical form. Local: Shannon Co. Carex glaucodea Tuckerm. open ground and open woods, Cireumneutral. Central and southern Mo.: St. Louis, Jefferson, Ste. Genevieve, St. Franeois, Bollinger, Iron, Dunk- lin, 2m Ripley, Shannon, Dent, Texas, Phelps, Boone, Douglas, and Webster counties Carex acide Dewey. S py woods. Cireumneutral. Southeastern Mo., local: Dunklin Co. Carex p Sehw Wet de td Southern and eentral Mo., seattered: St. Louis and diis eounties. Carex oxylepis Torr. & Hook. Rich low woods. Cireumneutral. Southeastern Mo.: Dunklin and Butler counties. Carex Davisii Schwein. & Torr oist or rieh woods, ió and borders of streams. General: _ Marion, Ralls, Lincoln, St. Charles, St. Louis, Franklin, Stoddard, Shan Phelps, Boone, Sehuyler, Seotland, fotum. Mercer, Harrison, Gentry, Clay: Jackson, Johnson, Greene, Jasper, and Newton counties. Carex cherokeensis Schwein. wampy ground. Cireumneutral. Southeastern Mo.: Butler Co. Carex virescens Muhl. woods and swampy open ground. Circumneutral. Southeastern Mo.: sas, Dunklin, Butler, and Ripley countie Carex hirsutella Mack. Bull. Torr. Bot. Club 50: 349, 1923. Carex triceps var. hirsuta (Willd.) Bailey (G). Carex complanata Torr. (B € B). Dry rocky woods and prairies. Oxylophile. General. Carex caroliniana Schwein. Carex triceps var. Smithü Porter (G). Wet woods and prairie swales. Cireumneutral. Central and southern Mo.: St. Louis, Jefferson, St. Francois, and Jasper counties. Carex Bushii Mack. Bull. Torr. Bot. Club 37: 241. 1910. Wet open ground. Circumneutral. Northern = western Mo.: Adair, Jack- son, and Jasper counties. Carex In bg Moist b and wet open ground. Calciphile to cireumneutral. Southern Mo., local: aa Washington, and Shannon countie Carex Joorii Bailey. Carez macrokolea Steud. (G). Swamps and low wet woods. Circumneutral. Southeastern Mo.: Dunklin Co. Carex Shortiana Dewey. Swales, meadows, and borders of ponds and streams. Circumneutral. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 493 Carex Buxbaumii Wahlenb. Swampy ground and prairies. Circumneutral. Scattered: Marion, Butler, and Clinton counties. Carex stricta Lam. Tussock Sedge Carez stricta var. a (Boott) Bailey, in part. mpy woods and prairies. Cireumneutral. Scattered: Shannon, Jackson, Jasper, and Greene mec Carex Haydeni Dewey. Carez stricta var. decora Bailey (G). Prairie swales. Cireumneutral. Northern and western Mo., local: Jackson and Adair counties, Carex scabrata Schw t open or outs M Cireumneutral. Southwestern Mo., loeal: Greene Carex torta Boott. Swamps and banks of streams. Oxylophile to eireumneutral. Southern Mo.: Washington, Ste. Genevieve, Ripley, Iron, Shannon, and Webster counties. Carex Emoryi Dewey. Wet woo às and prairie swales. Circumneutral, Scattered: Jackson and Mercer counties Carex crinita Lam. Fringed Sedge. Prairie swales, sloughs, and margins of ponds and streams. Cireumneutral. General. Carex crinita var. d (Schwein.) Schwein. & Torr. Carex gynandra Schwei Prairie swales, nim d margins of ponds. Circumneutral. Seattered: Ripley and Jaekson counties. Carex lacustris Willd. Carez riparia Curtis (G). Carex riparia var. lacustris (Willd.) Kükenth. Low wet woods and swamps. Circumneutral. General. Carex hyalinolepis Steud. Carex impressa (S. H. Wright) Mack. Bull. Torr. Bot. Club 37: 236. 1910. (B & B). Carex riparia var. impressa S. H. Wright. Swampy woods and borders of sloughs. Circumneutral. General. Carex lanuginosa Michx. Prairie swales and boggy ground. Cireumneutral. General. Carex laeviconica Dewey. Carex trichocarpa var. Deweyi Bailey (G), in part. Carez trichocarpa var. imberbis Gray (G), in part. Carez trichocarpa Muhl. (B & B), in p Low wet ground. Cireumneutral. Local: Schuyler and Jackson eounties. Carex atherodes Spreng. Carez trichocarpa var. aristata (R, Br.) Bailey (G). Low swampy ground. Cireumneutral. Local: Jackson Co. [Vor. 22 494 ANNALS OF THE MISSOURI BOTANICAL GARDEN Carex vesicaria L. Meadows and low ground. Cireumneutral. Local: St. Louis Co. Carex vesicaria var. monile (Tuckerm.) Fernald. Carex monile Tuckerm Meadows and low anani Cireumneutral. Northern and central Mo.: Scot- land, Adair, St. Louis, Saline, and Jackson counties. Carex lurida Wahlenb. Carex lurida var. flaccida Bailey, in part. Wet woods, swales, and borders of sloughs and streams. Cireumneutral. General. Carex comosa Boott. Swampy ground. Circumneutral. Central-western and southeastern Mo.: Platte, Dunklin, and Butler counties. Carex hystricina Muhl. Rich moist woods and swales. Circumneutral. Central and southern Mo.: St. Louis, Jefferson, Washington, Iron, Shannon, Texas, Ripley, Saline, Jackson, and Greene counties. This species and Carex lurida have been confused with Carex Schweinitzii which is not known to occur in Mo, Carex Frankii Kunth. Prairie swales and borders of sloughs and streams. Cireumneutral. General. Carex squarrosa L. Wet woods and prairies, Cireumneutral. General. Carex typhina Michx. Carex typhinoides 2 (G). Wet woods and swamps. Cireumneutral. Northern, central, and eastern Mo.: Scotland, Adair, Shelby, Audrain, St. Louis, Franklin, Phelps, Bollinger, Wayne, Butler, Ripley, Boone, Linn, Livingston, and Jackson counties. Carex intumescens Rudge. Low woods and swampy open ground. Cireumneutral. Southeastern Mo.: Cape Girardeau, Dunklin, Butler, and Ripley counties. Carex Asa-Grayi Bailey. Carez Grayii var. hispidula Gray (G), in part. Low wet woods and sloughs. Circumneutral. General. Carex louisianica Bailey. Carez Halei Carey, not Dewey (G). Swamps and wet woods. Cireumneutral. Southeastern Mo.: New Madrid, Dunklin, and Butler counties, Carex lupuliformis Sartwell. ampy ground. Cireumneutral. Northern and central Mo., rare and scat- tered: St. Louis and Livingston countie Carex lupulina Muhl. Hop Sedge. Carex lupulina var. pedunculata Dewey (G), in Wet — swampy woods, and borders e pa” Circumneutral. Genera 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 495 Carex gigantea Rudge. Swampy woods and wet open ground. Circumneutral. Southeastern Mo.: Dunklin, Butler, and Ripley counties. Fam. ARACEAE Arisaema triphyllum (L.) Schott. Indian Turnip, Jack-in-the-pulpit. Rich or rocky woods and alluvial thickets. Cireumneutral. General. Arisaema Dracontium (L.) S Green Dragon. woods and dcn in one situations than the preceding. Circum- Mos General Peltandra virginica (L.) Kunth. Arrow Arum. Boggy ground and borders of sloughs and ponds. Cireumneutral. East- central Mo.: St. Charles, St. Louis, and Jefferson counties. Peltandra virginica f. hastifolia Blake, Rh. 14: 105. 1912. Wet or boggy ground. Circumneutral. Southeastern Mo.: Dunklin Co. Acorus Calamus L. Sweet Flag. Wet prairies and meadows, and borders of sloughs and ponds. Cireumneutral. General. Fam. LEMNACEAE een polyrhiza (L.) Sehleid. Duckweed. onds and sloughs. General but not common. sre oligorhiza (Kurz) Hegelm. See Bull. Torr. Bot. Club 61: 233-236. p -flowing streams and sloughs. Local: Jackson Co. Lemna trisulca L. Ivy-leaved Duckweed. Stagnant pools, slow streams, and deep springs. Circumneutral. Central and southern Mo., seattered: Shannon, Oregon, and Jackson counties. Lemna valdiviana Philippi. See Rh. 37: 75. 1935. Lemna cyclostasa (Ell.) Chev. (B & B). Ponds and sloughs. Circumneutral. Central and southern Mo., scattered: St. Francois, Dunklin, Oregon, and Jackson counties. Lemna dandi Torr. onds and sloughs. Central and southern Mo.: St. Louis, Dent, Boone, Raine, J ae , Cass, and Greene countie Lemna perpusilla var. trinervis Aust. Ponds. Occasionally found with the typical form. Jackson Co. Lemna minor L. Little Duckweed. Ponds, sloughs, and stagnant streams. Circumneutral. General. Wolffia columbiana Karst. Ponds and sloughs. Cireumneutral. Central Mo., and perhaps more general: St. Louis, Boone, and Jackson counties. Wolffia papulifera C. H. Thom Sloughs, ponds, and rivers. pm E Seattered: Dunklin and Boone eountie [VoL. 22 496 ANNALS OF THE MISSOURI BOTANICAL GARDEN Wolffiella floridana (J. D. Sm.) C. H. Thompson. Stagnant water. Circumneutral. Southeastern Mo.: Dunklin Co., in Varney and St. Francis rivers. Fam. CoMMELINACEAE Tradescantia subaspera Ker-Gawl, See Contr. Arn, Arb. 9: 48. 1935. Tradescantia pilosa Lehm. (G), (B & B). Rieh woods, shaded ger: and banks of streams. Cireumneutral. Eastern and middle Mo.: Marion, Monroe, "e Pike, Boone, Callaway, Montgomery, Warren, St. Louis, lios , Franklin, Washington, Crawford, Ripley, Oregon, Shannon, Texas, Maries, Pulaski, coo Miller, Moniteau, Morgan, and Ozark counties, Tradescantia canaliculata Raf. See Contr. Arn. Arb. 9: 74. 1935, Tradescantia reflera Raf. (G), (B & B). sd prairies, moist banks, thickets, and open woods. Cireumneutral. Genera Tradescantia virginiana L. See Contr. Arn. Arb. 9: 60. 1935, Spiderwort. Tradescantia brevicaulis Raf., not of recent auth. Rich moist ledges along bluffs, and alluvial woods. Cireumneutral to eal- eiphile. Eastern and central Mo.: Pike, Lincoln, St. Charles, St. Louis, Jeffer- son, Washington, Ste. Genevieve, Iron, Madison, Wayne, Carter, Ripley, Mont- gomery, Boone, Clinton, and Jaekson counties, Tradescantia bracteata Small. See Contr. Arn. Arb. 9: 85 Prairies, thickets, and meadows, and also often nmi along railway embankments and roadsides. Cireumneutral to oxylophile. Northern and central Mo.: Scotland, Pike, Lincoln, St. Charles, St. Louis, Jefferson, Atchison, Jackson, Cass, and Henry counties liae mts longipes Anderson & Woodson, Contr. Arn. Arb. 9: 91. 1935. radescantia brevicaulis of auth. in part, not Raf. (G), (B & B). ihi hillsides and sandy or rocky open woods. Oxylophile. Eastern Ozark region: Crawford, Washington, Madison, St. Francois, wies Reynolds, Wayne, Carter, Shannon, Dent, Texas, Howell, uni Douglas countie Tradescantia Tharpii Anderson & Woodson, Contr. Arn. Arb. 9: 70. 1935. Tradescantia brevicaulis of auth. in part, not Raf. (G), (B & B). Rocky prairies and hillsides. Cireumneutral Southwestern Mo.: Greene, Lawrenee, and Jasper counties. Tradescantia ozarkana Anderson & Woodson, Contr. Arn. Arb. 9: 56. 1935. Rich rocky woods and moist ledges along bluffs. Cireumneutral to ealeiphile. Southwestern Mo., local: Barry Co. Tradescantia Ernestiana Anderson & Woodson, Contr. Arn. Arb. 9: 58. 1935. Rocky wooded hillsides and ledges along bluffs. Cireumneutral. Southwest- ern Mo.: Douglas, Taney, Barry, Jasper, and Newton counties Commelina communis L. Creeping Day-flower. Wet alluvial eultivated or waste ground, and in woods and thickets. General. Sometimes becoming a troublesome weed. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 497 Commelina caroliniana Walt. ields and waste ground. Indifferent. Introduced in Jackson Co. Commelina virginica L. See Bull. Torr. Bot. Club 43: 97-111. 1916, Commelina hirtella of auth., not Vahl (G), (B & B). Moist alluvial ground along streams, and margins of swamps and sloughs. Central and odes Mo.: St. Louis, Jefferson, Bollinger, Scott, — New Madrid, Pemiscot, Dunklin, Butler, Wayne, Carter, Ripley, Oregon, Sha non, Phelps, Pulas "| Ta elede, Jackson, Cedar, St. Clair, Great Barry, Pind Newton, and McDonald counties. Commelina erecta L. Day-flower. Commelina virginica of auth., not L. (G), (B & B). Bluffs and hillsides, sandy al rocky woods, alluvial banks, and gravel and sand bars along Ozark streams. Oxylophile to eireumneutral. General: Mercer, Linn, Warren, Montgomery, Maries, Franklin, Jefferson, Dent, Reynolds, Shan- non, Oregon, Texas, Pulaski, Miller, Camden, Laclede, Dallas, Morgan, Hickory, Polk, Cedar, Douglas, Ozark, Andrew, Jackson, Vernon, Jasper, Barry, and McDonald counties. Commelina nin var. crispa s Palmer & Steyermark, comb. nov. crispa Wooto & B). ed p ledges and glades. Circumneutral to calciphile. General but scattered: Putnam, Mississippi, Ripley, Jackson, Jasper, and McDonald counties. Commelina longicaulis Jacq. See Bull. Torr. e ra 43: 96. 1916. Commelina nudiflora of Am. auth., not L. (B Moist open or eultivated pal in sandy or Bu soil, frequently a weed in low fields. Cireumneutral. General: Shelby, Lincoln, Randolph, Saline, St. Charles, St. Louis, Jefferson, Madison, Scott, New Madrid, Pemiscot, Dunklin, Ripley, Oregon, Howell, Shannon, Miller, Henry, Pettis, Douglas, Barry, Vernon, and Jasper counties. Fam. XYRIDACEAE Xyris flexuosa Muhl. Moist borders of streams and ponds. Cireumneutral. Local: Greene Co. Fam. PONTEDERIACEAE Eichhornia crassipes (Mart.) Schlecht. Water Hyacinth. Introduced along drainage canals. Butler Co Pontederia cordata L. Pickerel-weed. Ponds and sloughs. "aad utral Central and southern Mo.: Lincoln, St. Charles, Jae Crawford, Stoddard, Ray, Jackson, Barton, Jasper, and Newton counties. Heteranthera limosa (Sw.) Willd. Slow streams, ponds, and sloughs. Cireumneutral. Central and southern o.: Montgomery, St. Louis, Jefferson, New Madrid, Dent, Ray, Jackson, Greene, Ozark, Taney, Jasper, and McDonald counties. E [Vor. 22 498 ANNALS OF THE MISSOURI BOTANICAL GARDEN Heteranthera reniformis R. & P. Mud Plantain. Ponds and sloughs. — ake General but seattered, and mainly along the larger rivers: Lincoln, St. Charles, St. Louis, Jefferson, Dunklin, Shannon, Boone, Holt, Clay, eng and "nine counties. — dubia (Jaeq.) MaeM. Water Sta ams and sloughs. Circumneutral. Central, pem and 1 northwestern b pineda: in the Ozark region, and seattered elsewhere: St. Louis, Jefferson, Franklin, Gasconade, Crawford, Dent, Reynolds, Wayne, Butler, Denkita, Ripley, Oregon, Carter, Texas, Pulaski, Osage, Miller, Camden, vtto Laclede, Douglas, Ozark, Taney, Stone, Greene, Holt, and Jackson countie Fam. JUNCACEAE Juncus bufonius L. Toad Rush Sandy and gravelly ground along the larger rivers. Oxylophile. West- central Mo.: Clay and Jackson counties. Juncus macer S. F. Gray. See Jour. Bot. 68: 366, 1930. Slender Rush. Juncus tenuis of auth., not Willd. (G), (B Juncus monostichus Bartlett (G). Fields, roadsides, waste ground, and wet places. Indifferent. General and common. True Juncus tenuis Willd. (Juncus dichotomus Ell. a auth.) has been misidentified in Missouri and is not known from the state, Junows monos- tichus is considered only an abnormal form of Juncus macer and is treated here as identical with the latter species. Juncus macer var, anthelatus (Wiegand) Fernald, Jour. Bot. 68: 367. 1930. Juncus tenuis var. anthelatus Wiegand (G). Occasionally found with the typical form. Scotland, Shelby, Boone, Scott, Wayne, Ripley, Mercer, Saline, and Jasper counties. Juncus interior Wiegand. Upland prairies and meadows. Circumneutral to oxylophile. General. Juncus kansanus Hermann, Papers Mich. Acad. Sci. 20: 41. Dry rocky open ground. Circumneutral. Northeastern Mo., local: Pike Co. Juncus secundus Beauv. Dry upland prairies, sandy or rocky ground. Oxylophile. Eastern and south- ern Mo.: Lincoln, Jefferson, St. Francois, Dent, Phelps, Texas, and Laelede countie Juncus ÓN Wie Moist prairies, id ws, and wet places along streams. Circumneutral. Central and southern Mo.: Crawford, Iron, Shannon, Phelps, Dallas, and Jack- son eounties. Juncus effusus L. var. solutus Fernald & Wiegand, Rh. 12: 90. 1910. Rush. Prairie swales and borders of ponds, sloughs, and streams. Circumneutral. General, Juncus balticus Willd. var. littoralis Engelm. Sand bars and alluvial ground along Missouri River. Cireumneutral. Local: Jaekson Co. 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 499 Juncus canadensis J. Ga Moist ground along — Southern Mo., loeal: Howell Co. Juncus subcaudatus (Engelm.) Coville & Blake, Proc. Biol. Soc. Wash. 31: 45. 1918 Juncus canadensis var. subcaudatus Engelm. (G). : Moist ee along rocky beds of streams and slopes. Calciphile. Southeast- ern Mo.: Texas and Iron counties. Juncus nodosus L. Knotted Rush. Sand bars of Missouri River. Introduced in Jackson Co. Juncus Torreyi Coville. Borders of ponds, sloughs, and eroded banks. Circumneutral. General. Small-headed forms of this have been confused with J. nodosus. Juncus brachycarpus Engelm Prairies, swales, wet sant ows, and wet sandy borders of small stroams. Cir- cumneutral to oxylophile. Southern and central Mo.: Boone, St. Louis, Jeffer- son, Crawford, Iron, Scott, Mississippi, Wayne, Shannon, Phelp Howell, Ozark, Webster, Greene, Vernon, Jasper, McDonald, and Barry counties. Juncus validus Coville. Juncus polycephalus of auth. in part, not Michx. (G), (B & B). Prairie swales and swampy ground. Oxylophile to circumneutral. South- western Mo.: Jasper and Vernon counties. This species has been confused with Juncus polycephalus which is not known in Mo. Juncus scirpoides Lam Wet open or swampy ground. Circumneutral. Central and southern Mo.: St. Louis, Mississippi, Wayne, Howell, and Greene counties. Juncus acuminatus Michx. Knotty-leaved Rush. Borders of ponds, sloughs, and ditches. Cireumneutral. General. Juncus bonn Vill. var. fuscescens Fernald. w sandy ground along Missouri River. Circumneutral. Western Mo., local: J a Co. Juncus diffusissimus Buckley. Wet meadows, bogs, and borders of ponds, streams, and sloughs. Cire neutral to oxylophile. Central and southern Mo.: Miller, Crawford, need Ripley, Howell, Benton, Henry, Cedar, Vernon, Barton, Jasper, and MeDonald counties. Juncus nodatus Coville. Juncus robustus (Engelm.) Coville, not Wats. (G). andy open ground, borders of streams and sloughs. Oxylophile. Southern and east-central Mo.: St. Louis, Bollinger, Wayne, Butler, Howell, Barton, and Jasper counties. Juncus marginatus Rostk. ES Rh. 37: 156. 1935. uncus aristulatus Mic Prairies, meadows, = open sandy ground. Circumneutral to oxylophile. Eastern, eentral, and southern Mo.: Clark, Marion, Audrain, Boone, St. Louis, Jefferson, St. Francois, Mississippi, Dunklin, Wayne, Iron, ttis: Butler, [Vor. 22 500 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ripley, Crawford, Phelps, Shannon, Howell, Henry, Cedar, Greene, Vernon, Barton, Jasper, Newton, McDonald, and Barry counties. Juncus marginatus var. setosus Coville, Occasionally found with the typical form. Southern Mo.: Shannon, Greene, Jasper, and Newton eounties. Juncus biflorus Ell. See Rh. 37: 156. 1935. Juncus aristulatus of Bieknell and later auth., not Miehx. (G). Moist sandy open ground. Oxylophile. Southern and east-central Mo.: St. Louis, Butler, Iron, Wayne, Phelps, Texas, Greene, Jasper, Barry, and MeDonald counties. Juncus biflorus f. adinus Fernald € Griscom, Rh, 37: 157. 1935. Wet meadows and swampy ground. Oxylophile. Seattered: St. Louis and Jasper counties. Luzula campestris (L.) DC. var. bulbosa A. Wood. Dry roeky prairies, meadows, glades, and dry open woods, sometimes in sandy alluvial ground. Oxylophile. Central and southern Mo. south and east of a line drawn from St. Louis, Warren, Montgomery, and Callaway counties to Henry and Vernon counties, Fam. LILIACEAE Amianthium muscaetoxicum (Walt.) Gray. Fly Poison. Sandy woods. Oxylophile. Southern Mo., rare and seattered: Carter, Ripley, and Greene counties, Stenanthium gramineum (Ker) Kunth. oist meadows and rich woods along bluffs. Cireumneutral. Eastern and middle Mo.: St. Louis, Jefferson, St. Francois, Madison, Ripley, Shannon, and Phelps counties, Stenanthium gramineum f. robustum (Wats.) Palmer & Steyermark, comb. nov. Stenanthium robustum Wats. (G), (B & B). Moist rocky woods and thickets. Cireumneutral. East-central and PP” Mo., seattered: St. Louis, Lawrence, and Jasper eounties. Zigadenus — Pursh. See Rh. 37: 256-257. 1935. Moist shaded limestone bluffs. Bae cine Local: Shannon Co. This species has been oe ith Zigade glaucus Nutt. (Z. chloranthus Richards.), which is not known to occur in Mo 0. Zigadenus Nuttallii (Gray) Wat Toxicoscordion Nuttallii (Gray) Rydb. (B & B). Dry rocky limestone slopes and glades. Caleiphile. Local: Oregon Co. Melanthium virginicum L. Bunch-flowe Wet meadows, thickets, and iunii ground below bluffs. Circumneutral. General but seattered:: Linn, Adair, Knox, Pike, St. Louis, Jefferson, Franklin Washington, Phelps, Iron, Shannon, Boone, Webster, Greene, dk nis Johnson, and Jasper counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 501 Veratrum Woodii Robbins. False Hellebore. Rich or rocky woods. Cireumneutral. Eastern, middle, and southern Mo.: vili Sullivan, Adair, Schuyler, Knox, Shelby, Marion, Ralls, Boone, Callaway, ineoln, Warren, St. Louis, Jefferson, Franklin, St. Francois, Madison, Bol- E Iron, od Dent, Shannon, Reynolds, Oregon, Texas, Howell, Taney, Wright, and Barry counti Uvularia grandiflora Sm. Straw-flower, Large Bellw Rich alluvial or rocky woods. Cireumneutral. ale but absent from most of the western prairie counties. This ia has sometimes been eonfused with U. perfoliata, which is not known in Mo. Oakesia sessilifolia (L.) Wats. Small Bellwort, Wild Oats. Rich alluvial or low woods, and along shaded banks and bluffs. Circum- neutral, Scattered, ie ap in northern Mo.: Harrison, Boone, Knox, Au- drain, and Jasper co Allium cernuum Roth. an Wild Onion. Rocky ledges along bluffs. Caleiphile. East-eentral and southern Mo.: Jef- ferson, Franklin, Washington, Crawford, Dent, Phelps, Pulaski, Shannon, and Greene counties. Allium stellatum Ker. Glades and B ipM ledges along bluffs. E Central and south- ern Mo.: Lincoln, St. s, Jefferson, Franklin . Genevieve, St. Francois, Madison, Wess ion. virt Phelps, Maries, nd Dent, Shannon, Texas, Laclede, Boone, Henry, Benton, Dallas, Greene, Wright , Webster, Douglas, Ozark, Taney, and Barry counties. Allium canadense L. Wild Garlic. Moist open woods, meadows, “prairies, and thickets. Indifferent. General. Allium Nuttallii Wats. | : | Glades and dry rocky ledges. Caleiphile. Southwestern Mo., local: McDonald Co. Allium mutabile Miehx. Wild Onion. mestone glades, barrens, and prairies. Caleiphile. General but mostly in Pise and southern Mo.: Montgomery, Jefferson, St. Franeois, Madison, Iron, Dent, Maries, Laclede, Ozark, Wright, Webster, ie Boone, Daviess, Jack- son, Vernon, Jasper, Newton, and MeDonald countie ~ ALLIUM SATIVUM L ields, waste ground, and along railroads and roadsides. Indifferent. General. f ALLIUM VINEALE L. Crow Garlic. Moist fields and meadows. Indifferent. Scattered: St. Louis, Franklin, Boone, and Jasper counties. Í Allium tricoccum Ait. d Lee Low rich woods and slopes along bluffs. Circumneutral. Northern and cen- tral Mo., scattered and rare: Mercer, Boone, and St. Louis counties. jema bivalve (L.) Britton. False Garlic. En. and prairies. Circumneutral. General but commonest in the ed regio [Vor. 22 502 ANNALS OF THE MISSOURI BOTANICAL GARDEN HEMEROCALLIS FULVA L. Day Lily. Escaped from gardens to fields and roadsides. Indifferent. General. LILIUM TIGRINUM Ker. Tiger Lily. Occasionally found as an escape from cultivation. Lilium michiganense Farwell, Bull. Torr. Bot. Club 42: 353. 1915. Turk’s-cap Lily Lilium superbum of auth., not L. (G), (B & B). Rich open woods, prairie swales, and meadows, Circumneutral. General but not common: Knox, Adair, Livingston, Chariton, Mercer, Boone, Callaway, St. Louis, Jefferson, Franklin, Washington, Crawford, Dent, Iron, Saline, Jaekson, ohnson, Benton, Webster, Greene, Jasper, and Newt eountie Erythronium americanum Ker. Yellow Adder’s-tongue, Yellow Dog-tooth Violet. Rieh alluvial woods =s bluffs and streams. Circumneutral. Southern and east-central Mo.: Boone, St. Louis, Franklin, Ste, Genevieve, Iron, Shannon, Howell, Taney, a may, Jasper, and McDonald counties. Erythronium albidum Nutt. White Adder’s-tongue, White Dog-tooth Violet. Rich woods along bluffs and streams. Cireumneutral. General. Erythronium mesochoreum Knerr. Prairie Dog-tooth Violet. ocky prairies, glades, and dry open woods. Cireumneutral to calciphile, Central and southern Mo. Camassia hyacinthina (Raf.) Palmer & Steyermark, comb. nov. Wild hyacinth. Lemotris hyacinthina Raf, Camassia esculenta (Ker) Robinson, not Lindl. (G), (B & B). Rocky glades and prairies. Caleiphile to eireumneutral. General. SCILLA BIFLORA L Occasionally escaped from cultivation; in open woods and fields. Pike Co. MUSCARI BOTRYOIDES (L.) Mill. Grape Hyacin Escaped from cultivation; in fields and is ground. Seattered: Jefferson and Taney counties. Yucca glauca Nutt. Bear Grass. oess hills, Caleiphile, Northwestern Mo.: Atchison and Holt counties. Yucca arkansana Trelease. Rocky open woods and bluffs. Cireumneutral. Southwestern Mo.: Stone Co. Yucca filamentosa L. Needle-and-thread, Spanish Bayonet. Frequently escaped from cultivation; in open woods, thickets, and along rail- roads and roadsides. General but scattered. ORNITHOGALUM NUTANS L. Star of Bethlehem. Occasionally escaped from gardens to fields and lawns. Boone Co. ORNITHOGALUM UMBELLATUM L. Star of Bethlehem. Escaped from cultivation; in fields, meadows, and on shaded banks. Scat- tered: Marion, St, Louis, Jefferson, Perry, Callaway, Boone, Cole, Jackson, and Vernon counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 503 ASPARAGUS OFFICINALIS L. Asparagus. er l0 from eultivation; along roads and in fields and waste ound.. General. Smilacina racemosa (L.) Desf. False Spikenard, False Solomon's Seal. Rich or rocky woods. Circumneutral. General. Smilacina stellata (L.) Desf. False Solomon's Seal. Moist rich woods. Circumneutral. Northern and central Mo., scattered: Shelby, Boone, and Jackson counties. Medeola virginiana L. Indian Cueumber-ro Moist shaded banks along streams. leia Northern Mo., local: Linn Co. Qu cdm canaliculatum (Muhl.) Pursh. ix Solomon's Seal. Polygonatum giganteum Dietr., in part Eoi commutatum (R. € S.) Dietr., in part (G), (B & B). Rich or rocky woods and alluvial thickets ind banks. Cireumneutral. General. Polygonatum canaliculatum var. ovatum (Farwell) Palmer € Steyermark, comb. nov. Polygonatum biflorum ovatum Farwell, Bull. Torr. Bot. Club 42: 255. 1915. Polygonatum commutatum ovatum (Farwell) Gates. Polygonatum ovatum (Farwell) Bush. Rich woods and thiekets. Cireumneutral Western Mo.: Jaekson Co. Trillium sessile L. Wake-Robin. Rich alluvial or rocky woods. Circumneutral, Central and southern Mo.: St. Louis, Jefferson, Franklin, Shannon, Dent, Iron, Wayne, Oregon, Warren, andolph, G Cole, Phelps, Pulaski, Laclede, Texas, Howell, Morgan, Cooper, Chariton, Jack- son, Johnson, Bates, Benton, Hiekory, Polk, St. Clair, Greene, Christian, Taney, Douglas, Wright, Webster, ni Jasper, Newton, and McDonald counties. Trillium sessile f. luteum Peatt Trillium sessile eratis pe Oceasionally found with the typical form: St. Louis and Moniteau counties. Trillium viride B Open roeky iiid and banks. Cireumneutral. Eastern Mo.: Pike, Callaway, Warren, St. Louis, Franklin, and Iron counties. Trillium viridescens Nutt. ocky or alluvial open woods. Cireumneutral Southwestern Mo.: Ozark, Taney, Stone, and Barry counties. Trillium recurvatum Beck. Rich alluvial woods and thickets and rocky slopes or base of bluffs, Circum- neutral to queam East tern and central Mo., and loeally southwestwards to ouglas t. Louis, Jefferso | Frankie Washington, St. Francois, Ste. Genevieve, ary, Cans Oasis: "Belinea, Dunklin, Butler, Ripley, Wayne, Iron, Carter, Oregon, Boone, Howard, Cooper, Saline, Howell, and Douglas counties. [Vor. 22 504 ANNALS OF THE MISSOURI BOTANICAL GARDEN Trillium recurvatum f, Shayi Palmer & Steyermark, nom. nov Trillium recurvatum f. luteum Friesner, Butler pe Bot. “ie 3: 31. 1929, not Trillium recurvatum f. luteum Clute, Am. Bot. 28: Rich woods. Cireumneutral. East-eentr ai Mo., loeal: St. > Co. A typo differt petalis staminibusque luteis vel viridi-luteis. This form appears identical with that described by Friesner. The Trillium reourvatum forma luteum described by Clute has the elaws, the petals, and the stamens brownish-red, and therefore cannot be considered the same as the yellow- flowered form in Missouri in which the petals, including the elaws, are through- out yellow or nr: as are the stamens. This form is dedieated to Mr. W. F. Shay, Head of the Biology een at Normandy High School, “sete who first called attention to Collected at Monarch Hill, near Monarch, St. Louis Co., Mo. ei 29, 1935, by W. F. Shay (Mo. Bot. Gard.). Trillium Gleasoni Fernald, Rh. 34: 21—22. 1932, Trillium declinatum (Gray) Gleason, not Raf. (G), (B & B). Rich rocky or alluvial woods. Cireumneutral. Eastern and middle Mo.: Ralls, Boone, Callaway, Warren, St. Charles, St. Louis, Jefferson, Franklin, St. rancois, Perry, Scott, and Wayne counties. Trillium ozarkanum Palmer & Steyermark, s Roeky or alluvial ground. PREIS to A Southwestern Mo., local: Barry Co. Planta rhizomate crasso elongato fere horizontale; caule purpureo vel inter- dum in parte superiore viridi, 1-3 dm. alto; foliis lanceolatis elliptico-oblongis vel oblongo-ovatis, firme m reale apie o obtusis basi late euneatis vel subito in petiolum brevissimum eontractis, 4-8 . longis, 1-2.5 em. latis, manifeste quinque-nerviis; pedunculo ereeto, mes em. Pe sepalis oblongo-linearibus vel oblongo-ovatis, 1.5—4 em. longis, 0.5-1.6 em. latis, quinque-nerviis (vel raro tri-nerviis), petalis plerumque longioribus; petalis oblongo-laneeolatis vel anguste ovatis, intus eandidis ad roseis vel rubro-purpureis, extus plerumque magis coloratis vel interdum utrinque intense roseo-purpureis; filamentis 2.5—5 mm. longis; antheris oblongo-linearibus, 4.5-9 mm. longis; stigmatibus basi connatis apice reflexis; fruetu ovoideo vel subgloboso, 0.8-1 em. longo, 0.7—0.8 em. lat «ete thickened and elongated; stems 1-3 dm. tall, dark purple below and sometimes green in upper portion; leaves forest-green to grass-green, firmly membranaeeous, dull above, lustrous beneath, lanceolate or oblong-elliptic to oblong-ovate, obtuse at apex, broadly euneate at base, a: or abruptly eon- traeted into very short petiolules, 4—8.5 (rarely 3.5) em. long, 1-2.5 em. broad; pedunele ereet, 1.5-3 cm. long; sepals spreading, LM veined with 5 (rarely 3) prineipal veins, linear-oblong to broadly oblong-ovate, obtuse or rounded at apex, 1.5—4 em. long, 0.5-1.6 em. broad, usually exceeding the petals; petals varying from pure white to > or rose-purple within, usually more deeply colored without, or sometimes rich rose-purple on both sides, ge or slightly reeurved at tips, oblong- MR to ovate, broadest near the base, obtuse to acute at apex, 1.6-3.4 cm. long, 0.5-1.8 em. broad; A in 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 505 anthesis equalling or somewhat exceeding the stigmas, the filaments short, 2.5-5 mm. long, anthers straight, 4.5-9 mm. long; stigmas united at base, spreading and recurved at apex; sad white or pale green; fruit ovoid to subglobose, .8-1 em. long, 0.7-0.8 e road. Distribution: Cherty sol of low wooded slopes and along small draws in upland oak Mop western Ozark region, northwestern Arkansas, and south- western Misso new species adds another to the numerous endemies of the Ozark region, Mis of which are confined to the E ern portion. While elosely related to Trillium pusillum Michx., of the Atlantic Coastal Plain, and to Trillium texanum Buckley, of canteen: Texas, id ozarkanum has several characters that mark it as distinct from either of these species. The leaves and sepals average much broader and have normally five instead of three principal veins, which stand out more prominently; the petals are also much longer and broader on the average, and the peduncles are distinctly longer. The habitat, in com- p dry cherty soil on wooded slopes of deciduous upland woods, is quite pee in cherty soil along slopes of a draw in upland oak-chinquapin woods, 3 mi. south of Cassville, Barry Co., April 20, 1935, J. A. Steyermark 18628 (Mo. Bot. Gard. Herb. TYPE); 5 mi. s. of Cassville, April 13, 1930, C. Shoop, (Shoop Herb. and Univ. Mo. Herb.) ; ARKANSAS: in woods, dry SE chert soil, Boone Co., March 28, 1921, J. T. Buchholz (Mo. Bot. Gard. Herb. and Univ. Ark. Herb.) ; id pis. in ehert hills southwest of Elm Springs, Wash- ington Co., June 1, 1923, J. T. Buchholz (Mo. Bot. Gard. Herb. and Univ. Ark. Herb.) ; in Boone det flinty limestone soil, Osage, April 20, May 10, 1922, J. T. Buchholz E Herb); Withrow Springs, Madison Co., April 15, 1928, Moore $ Demaree 4803 (Mo. Bot. Gard. Herb.) ; same Shealites May 10, 1924, E. J. Palmer $1 (Palmer Herb.). Trillium nivale Riddell. Snowy Wake-Robin. Rich and rocky woods along streams and moist shaded ledges. Cireumneutral. Southern and central Mo.: Callaway and Texas eounties. Rare and local Smilax herbacea L. Carrion-flower. ich alluvial woods and thickets. Cireumneutral. General. Smilax herbacea var. pulverulenta (Michx.) Gray. Rich or rocky woods and thickets. Circumneutral. General, but scattered: Adair, Ralls, Pike, Jefferson, Shannon, Texas, Boone, Jackson, Greene, Lawrence, and Jasper counties Smilax herbacea var. lasioneuron (Hook.) A. DC. Occasionally found with the typical form. Cireumneutral General but seattered: St. Louis, Jefferson, Shannon, Wright, Adair, Mercer, Jackson, Cass, Greene, and Jasper counties. Smilax ecirrhata (Engelm.) Wats. Rocky woods. Cireumneutral. General but not common: Harrison, Gentry, Putnam, Mem Clark, Macon, Shelby, St. dc Jefferson, Franklin, Phelps, Miller, Morgan, Boone, and Jasper countie [Vor. 22 506 ANNALS OF THE MISSOURI BOTANICAL GARDEN Smilax ee L. var. quadrangularis (Muhl.) Wood. Cat-bri Moist alluvial grounds, woods, and thickets. Circumneutral P" oxylophile. E Mo.: Madison, Bollinger, Mississippi, New Madrid, Dunklin, But- ler, Ripley, and Oregon counties. Forms of S. hispida and S. Bona-nox have often been confused with this. The typical var. is probably not found in Mo. Smilax hispida Muhl. Bristly Green-brier. Woods, thiekets, and banks of streams. Cireumneutral. General. Smilax Bona-nox L. Stretch-berry, Green-brier. Rocky open woods, lu and fields. Oxylophile to eireumneutral. South- ern and east-central .: Bt. Louis, Jefferson, Perry, Cape Girardeau, Missis- sippi, New Madrid, Madison, Bollinger, Iron, Wayne, roni "ins Ripley, Oregon, Howell, Carter, Shannon, Reynolds, Dent, Phe Te Douglas Wright, Webster, Ozark, Taney, Stone, Barry, Jasper, "n m “McDonald counties, Smilax glauca Walt. var. genuina Blake, Rh. 20: 18, y open woods and wet open ground. Oxylophile. Southeastern Mo.: Madison, Carter, and Dunklin counties, Less common than the next following Smilax glauca var. leurophylla Blake, Rh. 20: 80. 1918. Rocky open woods, fields, and wet open ground. Oxylophile. Southeastern Mo.: Ste. Genevieve, St. Francois, Cape Girardeau, Bollinger, Madison, Iron, Wayne, Caries, Stoddard, Scott, Mississippi, New Madrid, Dunklin, Butler, Ripley, and Oregon counties. Fam. DIOSCOREACEAE Dioscorea — un Gmel. See Rh. 36: 345. 1934. Dioscorea glauca L (8). Rich nti and "nh rocky ground at base of bluffs, and talus slopes. Je Cireumneutral. Southern Mo.: Jefferson, Madison, Bollinger, Ste. Genevieve, ayne, Carter, Iron, vigilum Crawford, Ripley, Oregon, Texas, Taney, and Ozark countie Dioscorea cdm L. See Rh.20: 48. 1918. Wild Yam. Rich rocky woods and thiekets, Circumneutral. General. Dioscorea villosa var. glabrifolia (Bartlett) Blake, an 20: 49. 1918. Occasionally found with the typical form, Seattered: Perry, Daviess, Macon, Boone, Jackson, Cass, Henry, Jasper, Newton, vain ld, and Stone counties. Fam. AMARYLLIDACEAE NARCISSUS POETICUS L. Poet’s Narcissus. — escaped from gardens into abandoned fields, meadows, and waste place aac pine (LeConte) Kunth. Spider-lily. wet woods. Circumneutral. Southeastern Mo.: Pemiscot, Phoen Hitler, and Ripley counties. Agave virginica L. False Aloe. Rocky open woods and glades. Calciphile. Southern and east-central Mo., 1935] Ä PALMER € STEYERMARK—PLANTS OF MISSOURI 507 south of a line drawn from St. Louis to the southwestern corner of the state: St. Louis, Jefferson, Franklin, Ste. Genevieve, Perry, Washington, Madison, Bdbingsr] Dunklin, Ripley, Orten, Oregon, Dent, Howell, Shannon, Texas, Pulaski, Douglas, Ozark, Taney, Stone, Wright, Greene, Barry, and McDonald counties. Agave virginica f. tigrina (Engelm.) Palmer & Steyermark, comb. n Agave virginica var. tigrina Engelm. Trans. Acad. Sei. St. Louis " MS. 1876. Occasionally oceurring with the species in Missouri. Hypoxis hirsuta (L.) Coville. Yellow Star-grass. Dry open woods, thickets, glades, and rocky prairies. Oxylophile. General. Hypoxis hirsuta var. leptocarpa (Engelm. & Gray) Brackett. Rh. 25: 127. 1923 Moist sandy ground. Oxylophile. Local: Ste. Genevieve Co. Fam. InrpACEAE Iris virginica L. Wild Blue Flag. Iris Shrevei Small (S). Iris caroliniana Wats. (G), in part. Iris versicolor of auth., not L. (G), (B & B), in par Swamps and borders of ponds and sloughs. E d General. Iris fulva Ker. Red Iris. Swampy woods, and borders of sloughs and bayous. Circumneutral. South- eastern Mo.: D E Dunklin, Butler, and Ripley counties. Iris foliosa Mack. & B Sloughs, wet al and borders of low woods. Circumneutral. Central Mo., along the larger rivers: Marion, St. Louis, Jefferson, Washington, Boone, and Jackson counties. IRIS PSEUDACORUS L. Yellow Flag. Cultivated in old pori and oceasionally escaped to fields and roadsides. IRIS PALLIDA Lam. Blue Flag, Fleur-de-lis. Cultivated and escaping to roadsides and waste ground. Iris cristata Ait. Dwarf Wild Iris. Rocky woods, sandy banks and ledges along bluffs. Oxylophile to circum- neutral. Usually growing on sandstone, chert, or granitie rocks, rarely on lime- stone. Southeastern Mo.: Reynolds, Wayne, Carter, Ripley, Shannon, and Ore- gon counties. IRIS PUMILA L. ed from cultivation. Fields, meadows, and wet glades. Scattered: Wand and Boone counties. Nemastylis acuta (Bart.) Herb. Limestone glades and rocky prairies. Calciphile. Central and southern Mo., scattered and rare: Franklin, Washington, Ste. Genevieve, Iron, Wright, Taney, and Cass counties. BELAMCANDA CHINENSIS (L.) DC. Blackberry Lily. Cultivated in old gardens, and escaped and well established in thiekets, glades, and rocky open woods. Circumneutral, General. [Vor. 22 508 ANNALS OF THE MISSOURI BOTANICAL GARDEN Sisyrinchium albidum Raf. Meadows and open woods. Cireumneutral. Eastern and middle Mo.: Ran- dolph, St. Louis, Jefferson, Franklin, Washington, Ste. Genevieve, and Iron ounties. This and other species of Sisyrinchium have sometimes been con- fused with S. angustifolium which is not known in Mo. Sisyrinchium flaviflorum Bicknell. Post-oak woods. Oxylophile. Western Mo., loeal: Jaekson Co. Sisyrinchium campestre Bicknell. Blue-eyed Grass. Prairies, meadows, glades, and open rocky woods. Cireumneutral. General. Sisyrinchium campestre var. kansanum Bicknell. ocky prairies, glades, and open woods. Cireumneutral Southwestern Mo.: Jasper, Newton, and Barry counties. Sisyrinchium graminoides Bicknell. Sisyrinchium gramineum Curtis, not Lam. (G). Rocky open woods, meadows, and prairies, Cireumneutral General. Fam. MARANTACEAE Thalia dealbata Roscoe Swamps. boumseutral, Southeastern Mo., loeal: Butler Co. Fam. ORCHIDACEAE Cypripedium parviflorum Salisb. Yellow Lady 's Slipper. Rich rocky ground, and moist shaded hillsides. Cireumneutral. General but scattered and commonest in the western half of the state: Boone, Callaway, Crawford, Phelps, Shannon, Nodaway, Grundy, men J — Johnson, Greene, one, Lawrence, Jasper, Newton, and McDonald cou — parviflorum var. pubescens (Willd. ) E Large Yellow Lady 's Sli Moist alluvial banks and rich rocky woods. Cireumneutral. General but scattered and commonest in eastern and northern Mo.: Warren, Pike, St. Louis, Jefferson, Franklin, Washington, Crawford, Dent, Wayne, Carter, Ripley, Schuyler, Adair, Putnam, Mercer, Harrison, Jackso , and Greene counties, Cypripedium candidum Muhl. Small White Lady’s ena f. Rich rocky woods at the base of bluffs. Circumneutral. Seattered and rare: St. Louis, Shannon, Atehison, and Andrew counties. Cypripedium Reginae Walt. Showy Lady's Slipper. Cypripedium hirsutum of some Am. auth., not Mill. (G). Rich rocky or moist woods and bluffs. Caleiphile to um Seat- tered: Putnam, Iron, Shannon, Texas, and Ozark countie Orchis spectabilis L. Showy Orchis. Rich moist ds. Cireumneutral. General but uneommon: St. Louis, Jef- ferson, Franklin, Bollinger, Iron, ynolds, Dunklin, Schuyler, Adair, Sullivan, arren, Boone, Ray, Clay, Platte, Jackson, Greene, and Taney counties, 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 509 Habenaria flava (L.) Gray var. virescens (Muhl.) Fernald. See Rh. 23: 148. 1921. Yellow Orchis. Wet woods, borders of swamps and bogs. Cireumneutral to oxylophile. Gen- eral but scattered and mostly in eastern and northern Mo.: Schuyler, Living- ston, St. Louis, Jefferson, Iron, Butler, Dunklin, Shannon, and Howell counties. Habenaria clavellata (Michx.) Spreng. Green Wood Orchis. orders of swamps and bayous. Oxylophile. Southeastern Mo., local: Dunklin Co Habenaria ciliaris (L.) R. Br. Yellow Fringed Orchis. Pine woods and moist wooded slopes. Oxylophile. Southeastern Mo.: Iron and Ripley counties. Habenaria lacera (Michx.) R. Br. Ragged Orchis. Moist meadows and prairies. Cireumneutral. Scattered: Audrain, Boone, Callaway, St. Louis, and Shannon counties, and also reported from Greene Co. Habenaria peramoena Gray. Purple Fringed Orchis. Low borders of lakes, wet rocky ground about springs, and low rich woods along streams. Oxylophile. Southeastern Mo.: Iron, Carter, Bollinger, Ripley, Butler, and Dunklin counties. Habenaria leucophaea (Nutt.) Gray. White-flowered Prairie Orchis. Prairies and meadows. Circumneutral. Seattered: Ralls, St. Louis, Jeffer- son, Madison, Clinton, Jackson, Vernon, and Greene counties Pogonia trianthophora (Sw.) BSP. Nodding Log Orchis. Ri woods. Cireumneutral. Central and southern Mo.: Boone, St. Louis, Jefferson, Franklin, Bollinger, Mississippi, Dunklin, Butler, Shannon, Jackson, and Jasper counties. Isotria affinis (Aust. j =. Pogonia affinis t. (G). Wooded NL "ills. Caleiphile. Southeastern Mo., local: Bollinger Co. Calopogon pulchellus (Sw.) R. Br. Grass Pink. Moist rich woods. Cireumneutral, Scattered and rare: Shannon and Greene counties. xao latifolia (L.) All. Helleborine. Serapias Helleborine of auth., not L. (G), (B & B). E gravelly soil at base of bluff. Oxylophile. Southwestern Mo., local: Jasper Co. m. Beckii Lindl. Little Ladies’ Tresses. P 8, open rocky hago and dry meadows. Oxylophile to circumneutral. East- piked and southern Mo.: St. Louis, Ste. Genevieve, Scott, Franklin, Gas- conade, Crawford, Dent, Sine. Oregon, Texas, Greene, and Jasper counties. Spiranthes gracilis (Bigel.) Beck. Slender Ladies’ Tresses. Dry open woods. Oxylophile. Central and southern Mo.: St. Louis, Jeffer- son, Franklin, Ste. Genevieve, Iron, Reynolds, Dent, Shannon, Oregon, Texas, Randolph, Hocus, Jae akeon n, Dade, Greene, Tene’ Stone, Barry, Jasper, Newton, and McDonald counti [Vor. 22 510 ANNALS OF THE MISSOURI BOTANICAL GARDEN Spiranthes vernalis Engelm. & Gray. Prairies and dry meadows. Cireumneutral. Central and southern Mo., scat- tered: St. Louis, Jefferson, Jackson, Barton, and Jasper counties, Spiranthes lucida (H. H. Eaton) Ames, Moist sandstone Mise along ereek and springy ground. Oxylophile. South- eastern Mo.: Texas and Shannon counties. This speeies blooms during May. Spiranthes ovalis Lindl. Rich or low wet woods. Cireumneutral. Central and southern Mo., rare and seattered: St. Louis, Mississippi, and Jackson countie Spiranthes cernua (L.) Richard. Nodding Ladies' Tresses. Prairies, meadows, and moist places in glades. Cireumneutral. General. Spiranthes cernua var. ochroleuca (Rydb.) Ames, Prairies and glades. Cireumneutral. Central and southern Mo.: Boone, Shannon, Taney, Stone, Barry, Jasper, and Newton counties. Goodyera pubescens (Willd.) R. Br. Downy Rattlesnake Plantain. Epipactis pubescens (Willd.) A. A. Eaton n (G). Peramium pubescens (Willd.) MacM. (B & B). Moist, rich, sandy soil, in wooded ravines. Oxylophile. Southeastern Mo., local: Ste. Genevieve Co. Corallorrhiza Wisteriana Conrad. Coral-root. Rich or rocky open woods. Circumneutral. Central and southern Mo. Louis, Jefferson, Franklin, St. Francois, Madison, Boone, Jackson, e Lawrence, and Jasper ns, Corallorrhiza odontorhiza Nutt. Small Coral-root. Moist rich woods. Cireumneutral. East-eentral and southern Mo.: St. Louis, Jefferson, Franklin, Ste. Genevieve, Dunklin, Shannon, and Jasper eounties. Corallorrhiza trifida Chatelain. Yellow Coral-root. Moist rich hillsides. Cireumneutral. Southwestern Mo., rare and local: Lawrence Co. Malaxis unifolia Michx. See Rh. -= 176. 1926. Adder’s-mouth. Microstylis unifolia (Michx.) BSP. (G). Dry rich woods. Oxylophile. East- central and southern Mo.: St. Louis, Jef- ferson, Franklin, Ste. Aside Shannon, Howell, and Greene counties. Liparis liliifolia (L.) Richard. Large Twayblade. Dry open woods and hillsides. Oxylophile to eireumneutral, General, but uncommon and apparently absent from much of western Mo.: d Sullivan, Schuyler, Clark, Lewis, Boone, Callaway, St. Lou is, Jefferson, Franklin, Ste. Genevieve, Cape Girardeau, Dunklin, Phelps, Beynolda, Shannon, Texas, Howell, ouglas, Ozark, Greene, Jasper, and McDonald counties. Aplectrum hyemale (Muhl.) Torr. px bs -root, Adam-and-Eve. ich or alluvial woods. Cireum eutrai, Central and southern Mo.: Boone, Callaway, Warren, St. Charles, rg Lou , Jefferson, Franklin, Ste. Genevieve, Washington, Madison, Iron, Wayne, Clay, Jackson, and Greene counties. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 511 Hexalectris spicata (Walt.) Barnh. See Rh. 17: 136. 1915, Crested Coral-root. Hezalectris aphylla (Nutt.) Raf. (G), (B & B). Rieh roeky woods. Oxylophile to eireumneutral. Eastern Mo. south of the Missouri River, rare: St. Louis, Jefferson, Franklin, Dunklin, Phelps, Pulaski, and Texas counties. Fam. PIPERACEAE Saururus cernuus L. Lizard 's-tail. Swampy woods and borders of sloughs and sluggish streams. Circumneutral. Central and southern Mo.: Gasconade, Franklin, St. Louis, Jefferson, Bollinger, Mississippi, New Madrid, Pemiscot, Dunklin, Butler, Ripley, Carter, Wayne, Iron, Oregon, Reynolds, Shannon, Dent, Crawford, Texas, Phelps, Maries, Osage, Camden, Pulaski, Laclede, Punk Greene, Barry, Barton, Jasper, Newton, and MeDonald counties, Fam. SALICACEAE Salix dn Marsh. Blaek Willow woods and borders of cla sloughs, and ponds. Circumneutral, d Salix longipes Shuttlw. var. Wardii (Bebb) Schneider, Bot. Gaz. 65: 22. 1918. Ward's Willow. Saliz Wardii Bebb (G), (B & B). cky banks of streams. Oxylophile to circumneutral. Southern and central Mo. and loeally northward along Missouri and Mississippi rivers. Salix amygdaloides Anders. Peach-leaved Willow. Low woods and alluvial banks of streams, especially along the larger rivers. Cireumneutral. Northern, central, and eastern Mo., absent from most of the ark region. SALIX ALBA L. White Willow. Moist ground along banks of streams and ponds. Cireumneutral. Often planted and escaping. General but scattered. SALIX ALBA Var. VITELLINA (L.) Koch. In similar situations to the typical form and more common in eultivation and as an escape SALIX ALBA var. CALVA G. F. W. Mey. aliz rulea (Sm h. niin Joost. as an escape, in similar situations to the last. SALIX FRAGILIS L. Crack Willow, Brittle Willow ow grounds along streams. Seattered: St. Louis, Atehison, and Dekalb counties. Salix interior Rowlee. Sand-bar Willow. Saliz longifolia Muhl., not Lam. (G) Wet alluvial banks and sand-bars along streams and ponds. Cireumneutral. General. [Vor. 22 512 ANNALS OF THE MISSOURI BOTANICAL GARDEN Salix interior var. Wheeleri Rowlee, Bull. Torr. Bot. Club 27: 253, pl. 9. 1900. banks and sand bars along streams. Circumneutral. Scattered: lark, Monroe, Clay, and Jackson counties. SALIX BABYLONICA L. Weeping Willow. Escaped from cultivation in Jackson Co. Salix cordata Muhl. Heart-leaved Willow. et open ground along streams and about springs. Oxylophile to circum- neutral. General but not eommon: Clark, Lewis, Knox, St. Louis, Ste. Gene- vieve, Perry, Washington, Iron, Madison, Wayne, Carter, iinctalls, Shannon, Texas, Howell, Wright, Douglas, Behuyler, Adair, Putnam, Sullivan, Linn, Sa- line, Boone, Cooper, Benton, Polk, Greene, Taney, Mercer, Gentry, Livingston, Harrison, Worth, Atchison, Jackson, Bates, Vernon, Jasper, and Lawrence counties. Salix cordata var. angustata Anders. Moist ground along streams. Oxylophile to eireumneutral. Scattered, mainly in northern Mo.: Clark, Knox, Schuyler, Adair, Wayne, Sullivan, Grundy, Har- rison, Nodaway, Saline, and Jaekson eounties. Salix missouriensis Bebb. Saliz cordata var. myricoides (Muhl.) Carey (G), in part. Low wet ground, mainly along the larger rivers. Cireumneutral. Northern, central, and eastern Mo.: Clark, Marion, Boone, Warren, St. Louis, Cape Girar- deau, Worth, Atchison, Holt, Clay, and Jackson counties. Salix discolor Muhl. Pussy Willow. Low ground along streams. Circumneutral. Northeastern Mo., local: Clark 0. Salix petiolaris Sm. Low wet ground along streams. Cireumneutral. Northeastern Mo.: Marion and Pike counties. Salix humilis Marsh. Prairie Willow. Prairies, thickets, and open woods. Oxylophile to cireumneutral. General. Salix humilis var. rigidiuscula Anders. In similar situations to the preceding. Circumneutral. Scattered: Shannon, Atchison, Holt, and Jackson counties. Salix tristis Ait. Dwarf Gray Willow, Dwarf Pussy Willow Dry prairies and rocky or gravelly banks. Circumisentgal to oxylophile. Scattered: Pike, Dent, Phelps, Putnam, Sullivan, Harrison, and Jackson counties. Salix sericea Marsh. Silky Willow. Along small rocky streams and in boggy ground about springs. Oxylophile. Eastern Ozark region: St. Louis, Washington, St. Francois, Perry, Cape Girar- deau, Bollinger, Madison, Iron, Carter, and Texas counties. SALIX PURPUREA L. Purple Willow. oist or wet ground. Cultivated and occasionally escaped. Pike Co. Salix myricoides Muhl. A hybrid ow Salix cordata and S. sericea. Local: St. Louis Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 513 Salix humilis x missouriensis. Wet ground along streams. Jackson Co. Salix longipes var. Wardii x sericea. Wet rocky ground. St. Francois Co. Salix cordata x longipes var. Wardii. Along rocky streams. Taney Co. Salix cordata x nigra, Low open ground. Atchison Co. Salix cordata x humili Low ground along small streams. Putnam Co. Salix longipes var. Wardii x nigra. Along rocky streams. St. Louis, St. Francois, Iron, Dunklin, and Jasper counties, Salix amygdaloides x nigra. ow ground along river. St. Louis Co. Populus m Marsh. Cotton VE: Populus balsamifera of rece P auth., and perhaps of L., in pa Moist or un usually alluvial, xn along streams. Circumneutral. Gen- eral and probably in every county. Populus deltoides f. pilosa (Sarg.) Sudw. Populus balsamifera var. pilosa Sarg Swamps and low woods. Cireumneutral Southeastern Mo., local: Dunklin o POPULUS CANADENSIS Moench. var. EUGENEI (Simon-Louis) Schelle. Carolina Poplar. Frequently planted under the name of Carolina Poplar and occasionally escaping in thickets and along streams. Jasper Co. Populus tremuloides Michx. Quaking Aspen. hickets and borders of upland woods. Circumneutral to oxylophile. North- eastern Mo.: Clark and Adair counties, and also reported from Sullivan Co. Populus grandidentata Michx. Large-toothed Aspen. In similar situations to the last. Local: Clark Co. Probably introduced or escaped from cultivation. POPULUS ALBA L. Silver Poplar. Commonly planted and escaping freely into thickets and along fence rows and borders of small streams. Scattered: Ralls, Pike, St. Louis, Franklin, Reynolds, Dent, Boone, Mercer, Vernon, and Jasper counties. Populus heterophylla L. Downy Poplar, Swamp Cottonwood. Swamps and low wet woods. Circumneutral. Southeastern Mo.: Cape Gi- rardeau, Bollinger, Scott, Mississippi, New Madrid, Pemiscot, Dunklin, Stod: ard, Wayne, Butler, and Ripley counties. Fam. LEITNERIACEAE Leitneria floridana Chapm. Cork Wood. Deep swamps. Cireumneutral. Southeastern Mo.: Dunklin, Butler, and Ripley counties. [Vor. 22 514 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fam. JUGLANDACEAE Juglans nigra L. Black Walnut. Rich or open woods, along streams, Circumneutral to ealeiphile. General and probably found in every county. Juglans cinerea L. Butternut, ich woods and along bluffs. Cireumneutral. General except in extreme western part of state, but nowhere common: Clark, Sehuyler, Sullivan, Adair, Lewis, Macon, Shelby, "Rien Monroe, Ralls, Pike, Lincoln, Audrain, St. Louis, Jefferson, Franklin, St. Fra ron, Madison, Bollinger, Cape Girardeau, Scott, Mississippi, Stoddard, Dunkin, Butler, Wayne, Carter, Ripley, Reynolds, Shannon, Oregon, Texas, Crawford, Dent, Phelps, Pulaski, Maries, Boone, How- ard, Cooper, inline, Lafayette, Henry, Morgan, Miller, Camden, Benton, Hickory, Dallas, Laclede, Wright, Douglas, Ozark, Taney, Greene, and Stone counties. Carya Pecan (Marsh.) Engl. & Graebn. Pecan. Carya illinoensis (Wang.) K. Koeh (G). Hicoria Pecan (Marsh.) Britton (B & B). w woods and alluvial ground along streams. Circumneutral. General but most frequent along the M rivers: Clark, Lewis, Pike, Lincoln, Montgom- ery, St. Charles, St. Louis, Jefferson, Ste. Genevieve, Perry, Cape Girardeau, Bollinger, Mississippi, inihi t, Dunklin , Ripley, Ozark, Gasconade, Maries, Cole, Morgan, Cooper, Howard, Carroll aie, i vines, Platte, Clay, Jackson, Henry, Bates, Vernon, Jasper, i MeDonald counties. Carya aquatica (Michx. f.) Nutt. Bitter Pecan, Swamp Hickory Swamps and low wet woods. Circumneutral. s oon Se; Bollinger, Scott, Dunklin, and Butler counties. Carya cordiformis (Wang.) K. Koch. Pignut Hiekory, Bitternut Hickory. Hicoria cordiformis (Wang.) Britton (B & B). Woods, usually in moist ground along streams. Circumneutral. General. Carya cordiformis var. latifolia Sarg. Trees € Shrubs 2: 206. 1913. In similar situations to the typical form. Cireumneutral. General but not common: Clark, Marion, Shelby, Shannon, Mercer, Harrison, Nodaway, Jack- son, Jasper, and MeDonald counties. Carya ovata (Mill) K. Koch. Shag- yi or Bhell-bark Hickory. Hicoria ovata (Mill. Britton (B & B). Woods, limestone hills, and fuels of streams. Caleiphile. General and found in nearly all if not all eounties of the states, though uncommon in some parts of the Ozark regio Carya ovata var. ellipsoidalis Sarg. Bot. Gaz. 66: 235. 1918. Occasiona lly found with the typical form. Seattered: Marion, Ralls, and Johnson counties Carya laciniosa (Michx. f.) Loud. igs bed Shell-bark Hickory. Low alluvial woods along strea Cir eutral. General except in most of the Ozark region: Clark, iie Praha Putnam, Mercer, Adair, Living- ston, Linn, Macon, Shelby, Chariton, Monroe, Audrain, Lincoln, St. Louis, Jef- 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 515 ferson, Cape Girardeau, Bollinger, Madison, Scott, Mississippi, Pemiscot, Dunklin, Stoddard, Wayne, Butler, Ripley, Maries, Cole, Moniteau, Boone, Gentry, Atchison, Jackson, Johnson, St. Clair, Hickory, Cedar, Lawrence, Vernon, Barton, and Jasper counties. Carya tomentosa Butt, Mocker Nut, White Hickory. Carya alba (L.) K. Koch, not Nutt. (G), (B & B). Dry upland woods. Oxylophile. Usually on sandstone, chert, or igneous rocks. General. Carya tomentosa f. ficoides (Sarg.) Palmer € Steyermark, comb. nov. Carya alba var. ficoides Sarg. Trees & Shrubs 2: 206. 1913 Dry upland woods. Oxylophile. Southwestern Mo.: Jasper Co. Carya tomentosa f. ovoidea (Sarg.) Palmer & Steyermark, comb. now. Carya alba var. ovoidea Sarg. Bot. Gaz. 66: 238. 1918. Rocky upland woods. Oxylophile. tartera Mo.: MeDonald Co. Carya tomentosa var. subcoreacea (Sarg.) Palmer & Ss - comb. nov. Carya alba var. subcoreacea Sarg. Trees € Shrubs 207. Dry upland woods. Oxylophile. Southern Mo.: louis: hina, Texas, Barry, and MeDonald counties. Carya Buckleyi Durand var. arkansana Sarg. Bot. Gaz. 66: 249. 1918. a and rocky upland woods. Oxylophile. Eastern, central, and southern : Marion, Ralls, Pike, St. Louis, Jefferson, Ste. Genevieve, Perry, Dunklin, Bu Hi Wayne, Carter, Ripley, Oregon, Howell, Shannon, Texas, Iron, Dent, hi lps, i teau, Pettis, Jackson, Henry, Benton, Camden, Hickory, Dallas, Polk, Cedar, Vernon, Dade, Douglas, Ozark, Taney, Stone, Barry, Jasper, Newton, and Me- Donald counties. Carya Buckleyi var. villosa Sarg. Bot. Gaz. 66: 251. 1918. Carya glabra var. villosa (Sarg.) Robinson (G). Hicoria villosa (Sarg.) Ashe (B & B). Dry and rocky upland woods. Oxylophile. Eastern, central, and southern Mo.: Marion, Ralls, St. Louis, Ste. Genevieve, Iron, Bollinger, Dunklin, Howell, Phelps, Pettis, J pest rom and MeDonald counties. Carya glabra (Mill.) S pland woods. mem Southeastern Mo.: Cape Girardeau and Dunklin countie Carya ovalis (Wang.) Sarg. Dry or rocky upland woods. Oxylophile. Southeastern Mo.: Ste. Genevieve and Dunklin counties. Carya ovalis var. obovalis Sarg. Trees & Shrubs 2: 209. 1913. Carya glabra (Mill.) Spach, in part (G). Hicoria glabra (Mill.) Britton, in part (B & B). Dry and rocky upland woods. Oxylophile. Eastern and southern Mo.: Marion, Ralls, Pike, St. Louis, Washington, Iron, Madison, Cape Girardeau, Mississippi, Dunklin, Butler, Wayne, Shannon, Ozark, ins. Stone, Greene, enton, Hickory, St. Clair, Japa and Newton counti [Vor. 22 516 ANNALS OF THE MISSOURI BOTANICAL GARDEN Carya ovalis var. odorata Sarg. Trees & Shrubs 2: 208. 1913. arya microcarpa Nutt., in part (G). Hicoria microcarpa (Nutt.) Britton, in part (B Rocky upland woods. Oxylophile sub-region: Iron and Madison countie Carya ovalis var. obcordata Sarg. Trees & Shrubs 2: 208. 1913. Carya mierocarpa Nutt., in part (G). Hicoria microcarpa (Nutt.) Britton, in part (B & B). Dry and rocky uplands. Oxylophile. Eastern, central, and southern Mo.: Marion, St. Louis, Iron, Madison, m Dunklin, Ripley, Shannon, Boone, Howard, Cooper, Saline, Pettis, Johnso dati e 8, Ozark, Taney, Stone, Cedar, Vernon, Barton, Jasper, and MeDonald eoun Carya ovalis var. obcordata f. vestita Sarg. Bot. Gaz. 66: 246. 1918. Dry upland woods. Oxylophile. Henry counties MA Mo., in the Iron Mountain Scattered: Montgomery, Gasconade, and Fam. BETULACEAE Corylus americana Walt. Hazelnut. Thickets and borders of woods. Cireumneutral. General and probably in every eounty of the state. Corylus americana f. missouriensis (A. DC.) Fernald, Rh. 34: 96. 1932. Occasionally found with the typical form. Seattered. Corylus americana var. indehiscens Palmer & Steyermark, var lekets, fence-rows, and borders of woods. Cireu KM. Seattered : Jackson, Johnson, Jasper, and Newton counties, and perhaps more widely distributed. A typo differt involuero fruetus indehiscente vel dehiseente ad unum marginem Differs from the typieal form in having the involueres of the fruit completely united on one side and usually partly united, towards the base, on the other side North Carolina to Missouri and Arkansas. It has sometimes been confused with C. cornuta Marsh. (C. rostrata Ait.), which is rare in Missouri and which seems to = I to the southeastern eounti H CAROLINA: Sept. 19, 1897, Biltmore Herb. 1244b; MISSOURI: Gree E jd. Co., Sept. k 1924, B. F. Bush 10332 (Herb. Arn. Arb. Mig same locality and date E A 26035 (Herb. Arn. Arb.) ; ARKANSAS: Mar- tinsville, Faulkner Co., Oct. , 1924, E. J. Palmer 26537 (Herb. Arn. Arb.) ; Hot Springs, Garland Co., di 11, 1925, E. J. Palmer 29160 (Herb. Arn. Arb.) Corylus cornuta Marsh. Beaked Hazelnut. orylus rostrata Ait. (G), (B & B). Thiekets and open woods. Cireumneutral to oxylophile. Southeastern Mo.: Butler Co. Ostrya virginiana (Mill) K. Koch. Hop Hornbeam Borders of upland woods and along roeky bluffs and hillsides. to calciphile. General and probably in every county. Ostrya virginiana var. glandulosa (Spach) Sarg. Bot. Gaz. 67: 21 6. 1919 In similar situations to the preceding. Known in Clark, Jackson, ini Jasper counties, and probably more widely distributed, though uncommon, Cireumneutral 1935] PALMER & STEYERMARK—-PLANTS OF MISSOURI 517 Carpinus caroliniana Walt. Ironwood, Blue Beech. Along streams and bluffs, and in low woods. Cireumneutral. Eastern, central, and southern Mo.; apparently absent from most of the northern and western counties: Clark, ie Ralls, Pike, Lincoln, Warren, St. Charles, St. Louis, Jef- ferson, bes t. Francois, Perry, Cape Girardeau, lio prend Madison, Iron, Reyno n Wayne, cott, Mississippi, Pemiscot, Dunklin, Stoddard, Butler, Ripley, n Dent, Shannon, Oregon, Howell, Texas, nd Phelps, Calla- way, Boone, Howard, Carroll, Saline, Buehanan, Cole, Miller, Morgan, Camden, Benton, Hickory, Douglas, Ozark, Taney, Stone, Barry, and MeDonald counties. Betula nigra L. River Birch. Banks of rivers and creeks. Circumneutral. General but apparently absent from parts of the Ozark region and some of the northwestern eounties. Alnus rugosa (Du Roi) Spreng. Smooth Alder. Banks of doe and about springs and bogs. Oxylophile. Eastern, central, and southern Mo.: Lewis, Marion, Pike, em Warren, St. Louis, Jefferson, Franklin, rt Washington, St. Francois, Ste. Genevieve, Perry, Bollinger, Madison, Iron, Reynolds, Wayne iddia, Bipkey, Carter, Oregon, Howell, n , Shannon, Texas, Dent, Pulaski, Miller, Cole, Boone, Benton, St. Clair, Cedar, Douglas, Ozark, Taney, and N MN. PNE Fam. FAGACEAE Fagus grandifolia Ehrh, Beech. ds, low hills, and along streams. Cireumneutral. Southeastern Mo.: Cape Girardeau and Stoddard counties. Fagus grandifolia var. caroliniana (Loud.) Fernald & Rehder In similar situations to the preceding and more common. Southeastern Mo.: Perry, Cape Girardeau, Bollinger, Scott, Stoddard, Dunklin, and Butler counties. en ozarkensis Ashe, Bull. Torr. Bot. Club 50: 360. 1923. Chinquapin. Castanea pumila of auth. in part, not Mill. (G), (B & B). "in woods. Oxylophile. Southern Mo.: Howell, Barry, and McDonald counties. Quercus alba L. White Oak. oods, chiefly on dry uplands. Cireumneutral. General and probably in every county. Quercus alba f. latiloba (Sarg.) Palmer € Steyermark, comb. nov. Quercus alba var. latiloba Sarg. In similar situations to the preceding and more common in some sections. A complete series of gradations are found between the two forms, and it scarcely seems distinct enough to be regarded as a variety. Quercus stellata Wang. Post Oak. Upland woods and along small streams. Oxylophile to circumneutral. Gen- eral and probably in every county. Quercus lyrata Walt. Overcup Oak. Low « or swampy woods. Circumneutral. East-central and southeastern Mo.: St. Louis, Jefferson, Cape Girardeau, Bollinger, Scott, Mississippi, New Madrid, P Dunklin, Stoddard, Butler, Wayne, and Ripley counties. [Vor. 22 518 ANNALS OF THE MISSOURI BOTANICAL GARDEN Quercus macrocarpa Miehx. Bur Oak. Low woods, chiefly in alluvial soil along streams. Cireumneutral to ealeiphile. General. Quercus macrocarpa var. olivaeformis (Michx f.) Gra Loess hills along the Missouri River, ey a: Mo.: Atchison Co. Quercus bicolor Willd. Swamp White Oak. Low hills and alluvial ground along streams. Cireumneutral. Northern, east- ern, and central Mo., commonest north of the Missouri River and absent from most of the Ozark region. Quercus Prinus L. See Rh. 17: 40. de Pgs Oak, Basket Oak. Quercus Michauxii Nutt. (G), (B Low woods along streams. LM buio Southeastern Mo.: Iron, Madison, Bollinger, Cape Girardeau, Scott, Mississippi, Stoddard, Dunklin, Butler, and Ripley counties. Quercus Muhlenbergii Engelm. Chinquapin Oak, Yellow Oak. Upland woods, glades, and along bluffs. Caleiphile to eireumneutral. Most frequent in limestone areas, but also found sometimes on sandstone or igneous rocks, General and probably in every county. Quercus en f. Alexanderi (Britton) Trelease. Quercus Alexanderi Britton, In tuns situations to the preceding into whieh it passes by gradations. General but scattered. Quercus prinoides Willd. Shin Oak, Serub Oak. orders of woods, rocky bluffs, and glades. Calciphile. Central, southern, and western Mo., mostly in the western part: St. Louis, Shannon, Howell, Mercer, Harrison, Daviess, Caldwell, Worth, Gentry, Nodaway, Atchison, Holt, Andrew, Clinton, Jackson, Cass, Johnson, St. Clair, Vernon, Polk, Dade, Greene, and Stone counties. Quercus falcata Michx. Spanish Oak, Southern Red Oak, Quercus rubra L., in part, not of most recent auth. Quercus triloba Michx. (B & B), in part. Mir both alluvial and rocky ground. Oxylophile to — South- n Mo.: Perry, Cape Girardeau, Bollinger, Seott, Mississippi, New Madrid, Pesci, Dunklin, Stoddard, But) er, Wayne, Carter, Oregon, Ripley, Howell, ark, Stone, and Bar unt hen falcata P triloba rias Palmer € Steyermark, comb. nov. Quercus triloba Miehx., in part (B& B Upland or lowland woods. Southeastern Mo.: Ripley and Dunklin counties. Quercus falcata var. pagodaefolia Ell. Quercus falcata Michx., in part (G). Quercus pagodaefolia (Ell.) Ashe (B € B). In similar situations to the last. Southeastern Mo.: Dunklin and Butler counties Quercus falcata var. leucophylla (Ashe) Palmer & Steyermark, comb. nov. Quercus rubra var, leucophylla Ashe. Low woods. Southeastern Mo.: New Madrid Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 519 Quercus borealis Michx. f. Red Oak. rcus rubra var. ambigua (Michx. f.) Fernald (G). Rarely occurring in our region with the following and common form. Jackson Co. Quercus borealis var. maxima Ashe, Proc. Soc. Am. Foresters 11: 90. 1916; and Rh. 18: 48. 1916. Red Oak. Quercus rubra of auth., not L. (G), (B & B). Upland woods. Frequent along bluffs and steep hillsides in limestone regions, but also found on silieeous and igneous rocks. Cireumneutral. General, and probably in every county Quercus palustris Muench. Pin Oak. Low and swampy woods. Circumneutral. General, but absent or uncommon in mueh of the Ozark region, where it is sometimes found in sinks or shallow ponds and is known as Turkey Oak Quercus coccinea Muench. Scarlet Oak. Upland woods. Oxylophile. Southeastern Mo., mostly in the eastern Ozarks: Ste. Genevieve, Washington, Crawford, Dent, ines Iron, Madison, Wayne, wisis Dunklin, Ripley, Carter, Reynolds, Shannon, Oregon, Texas, and Howell eo Quercus coccinea var. tuberculata Sarg. Bot. Gaz. 65: 426. 1918. Oceasionally found with the typieal form. Southwestern Mo.: Butler Co. Quercus ellipsoidalis E. J. Hill. Jack Oak, Yellow Oak. Upland woods. Cireumneutral. Northern Mo., local: Harrison Co. as i Shumardii Buckley. Red Oak. upland woods. Circumneutral. Central and southern Mo.: Jefferson, ballet Dunklin, Ripley, Maries, Pulaski, Dent, Shannon, Howell, Montgomery, Ozark, Pettis, Jackson, Bates, Jasper, and McDonald counties. Quercus Shumardii var. Schneckii (Britton) Sarg. Red Oak. Quercus Schneckii Britton (B & Quercus texana of auth., not a (G). Upland woods, bluffs, RS alluvial ground along streams. Circumneutral to ealeiphile. Central E souther 0. Quercus Nuttallii Palmer, Jour. Arnold Arb. 8: 52. 1927. Low wet woods. Cireumneutral. Southeastern Butler Co. Quercus nigra L. Water Oak. Low woods. Circumneutral. Southeastern Mo.: Pemiscot, Dunklin, Butler, and Ripley counties. Quercus Phellos L. Willow Oak. pes niin Mo. and loeally north along Mississippi River: St. Louis, Madi- Bollinger, Scott, DIIS New Madrid, Pemiscot, Dunklin, Stoddard, du. and Bil dc eo Quercus marilandica Muench. Black Jack Oak. Dry or rocky upland woods. Oxylophile to cireumneutral. General, but com- monest in the Ozark region. [Vor. 22 520 ANNALS OF THE MISSOURI BOTANICAL GARDEN Quercus imbricaria Michx. Shingle Oak. Upland woods and along small streams. Cireumneutral. General but ap- parently absent from a number of the Ozark and southwestern counties. Quercus velutina Lam. Black Oak. Upland woods and well-drained ground along streams. Oxylophile to eireum- neutral. General and probably in every eounty. Quercus velutina var. missouriensis Sarg. In similar situations to the last. General but scattered and commonest in the Ozark region: Clark, Audrain, Shannon, Dunklin, Ozark, Taney, Stone, Barry, Jackson, Jasper, and MeDonald eounties Quercus leiodermis Ashe, Jour, Elisha Mitchell Sei, Soc. 40: 43. 1924. Upland woods. Cireumneutral. Jackson and St. Clair counties. Perhaps only a form of Q. velutina. x Quercus Hillii Trelease, Proc. Am, Phil. Soc. 56: 49. 1917. A hybrid between Q. macrocarpa and Q. Muhlenbergü. Holt and Jackson counties. X Quercus Bebbiana Schneider, Handb. Laubholz. 1: 201. 1904. A hybrid between Q. alba and Q. macrocarpa. Marion and Jackson counties. x Quercus Fernowii Trelease, Proc. Am. Phil. Soc. 56: 49. 1917. A hybrid between Q. alba and Q. stellata. St. Louis Co. x Quercus Leana Nutt. A hybrid between Q. imbricaria and Q. velutina. General but seattered: Adair P Boone, Jefferson, Washington, Iron, Madison, Saline, Dekalb, Jaekson, Dade, and Christian counties X Quercus Bushii Sarg. Bot. Gaz. 65: 453. 1918. A hybrid between Q. marilandica and Q. velutina. Jaekson, Johnson, and Jasper counties, X Quercus Porteri Trelease, Proc. Am. Phil. Soc. 56: 51. 1917. A hybrid between Q. borealis var. maxima and Q. velutina. Clark, Dunklin, and Jaekson counties. x Quercus exacta Trelease, Proc. Am. Phil. Soc. 56: 49. 1917. A hybrid beca i imbricaria and Q. palustris. ius Jefferson, Sullivan, and Dekalb coun x Quercus 2 Sarg. Trees & Shrubs 2: 223. 1913. x Quercus subfalcata Trelease. A hybrid between Q. falcata and Q. Phellos. Carter and Butler counties. x Quercus Richteri Baenitz, Allg. Bot. Zeit. 9: 85. 1903. A hybrid between Q. borealis var. maxima and Q. palustris. Osage and Benton counties, x Quercus runcinata Engelm. A hybrid between Q. borealis var. maxima and Q. imbricaria. St. Louis, Wayne, and Jackson counties. X Quercus tridentata Engelm. A hybrid between Q. imbricaria and Q. marilandica. St. Louis Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 521 x Quercus heterophylla Michx. f. A hybrid between Q. borealis var. maxima and Q. Phellos. Dunklin Co. x Quercus vaga Palmer & Steyermark, hyb. nov. Hybrida intermedia Q. palustrem et Q. velutinam foliis, gemmis hyemalibus fructuque. Intermediate in character of foliage, winter buds, and fruit, between Q. palustris and Q. velutina, the supposed parent species, both of which were grow- ing in close proximity to the type tree. Alluvial ground along small upland streams. Northeastern Missouri, and to be expected elsewhere in the range of the parent species. MISSOURI: Maryville, Nodaway Co., June 13, 1924, E. J. Palmer 25421 (Arn. Arb. TYPE) x Quercus inaequalis Palmer & Steyermark, hyb. nov. Hybrida intermedia Q. Phellos et Q. velutinam foliis, gemmis hyemalibus fructuque. Intermediate in character of foliage, winter buds, and fruit, between Q. Phellos and d elutina, the supposed parent species, both of which were growing near the type tree. The leaves in this hybrid are extremely variable and many of them unsymmetrical, Low hills bordering lowlands. Southeastern Missouri to Arkansas and Louisiana MISSOURI: Poplar Bluff, Butler Co., Sept. 11, 1919, E. J. Palmer 16342 (Arn. Arb. TYPE); same locality, Oct. 5, 1921, swa 20676 (Arn. 2. ad type); ARKANSAS: Hot Springs, Garland Co., Oct. jJ Pal 29098 (Arn. Arb.) ; same locality, Nov. 5, et ind 29577 d jeny Har ciconia Faulkner Co., Oct. 13, 1924, E. J. Palmer 26505 (Arn. Arb.); OUISIANA : r Sardis, July 30, 1915, R. S. n 814 paid Arb.) ; same lo- eality, Sept. dg 1915, R. S. Cocks 938 (Arn. Arb.). x Quercus Shirlingii Bush, hyb. nov. Hybrida intermedia Q. imbricariam et Q. Shumardü var. Schneckii foliis fruetuque Foliage and fruit intermediate between Q. imbricaria and Q. Shumardii var. p N the alid parent species, both of whieh were growing with t type species is named in honor of Professor A. E. Shirling, pe E ias College at Kuskas City, Missouri. MISSOURI: Oak Grove, Jackson Co., Oct. 22, 1927, B. F. Bush 11637 (Arn. Arb. pie x Quercus mutabilis Palmer & Steyermark, hyb. n Hybrida intermedia Q. palustrem et Q. omar var. Schneckii foliis fructuque. Intermediate in foliage and frs between Q. palustris and Q. Shumardii var. Low hills bordering river valley. Western Missouri. MISSOURI: Montieth Junction, Bates Co., Sept. 10, 1924, E. J. Palmer 26069 (Arn. Arb. TYPE). [VoL. 22 522 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fam. ULMACEAE Ulmus americana L. White Elm, American Elm. Woods, principally along streams. Circumneutral. General, and probably in every county. Ulmus Thomasi Sarg. Cork Elm, Rock Elm. Ulmus racemosa a not Borkh. (G). Woods along streams. Circumneutral. Northern, central, and southeastern : Clark, Monroe, Boone, Stoddard, Dunklin, Atehison, and Jackson eounties. Ulmus alata Michx. Winged Elm. Low and upland woods. Oxylophile to cireumneutral. East-central and south- ern Mo.: Lincoln, Warren, St. Louis, Jefferson, Franklin, Gasconade, St. Francois, Ste. Genevieve, Perry, Cape Girardeau, Bollinger, Madison, Iron, Reynolds, Wayne, Stoddard, "a , ty New Madrid, Pemiseot, Dunklin, Butler, Ripley, Carter, Oregon t, Howell, Shannon, Texas, Phelps, Pulaski, Callaway, Cooper, Cole, Do Maite ee Taney, Christian, Greene, Stone, Barry, and MeDonald counties, Ulmus fulva Michx. Slippery Elm, Red Elm. Upland woods and along bluffs. Calciphile to cireumneutral. General and probably in every county. Planera aquatica (Walt.) J. F. Gmel. Water Elm, Planer Tree. Swamps and low wet woods. Cireumneutral. Southeastern Mo.: Mississippi, New Madrid, Pemiscot, Dunklin, Butler, and Ripley counties, Celtis occidentalis L. Hackberry, Sugarberry. d upland woods, mostly along streams. Circumneutral. General but less common than the varieties that follow: Mercer, Putnam, Marion, Madison, Iron, Texas, Daviess, Gentry, Dekalb, Nodaway, Buchanan, St. Clair, and Jasper counties. Celtis occidentalis var. crassifolia (Lam.) Gray. Low and upland woods along streams. Cireumneutral. General: Clark, Seot- land, Knox, Adair, Sullivan, Carroll, Boone, Pike, e Girardeau, Butler, Oregon, Shannon, Ozark, Taney, Stone, Harrison, Grundy, Praise Alison, Johnson, Jasper, and McDonald counties. Celtis occidentalis var. canina (Raf.) Sarg. Bot. Gaz. 67: 217. 1919. Low and upland woods along streams. Cireumneutral. General and perhaps in every eounty. Celtis laevigata Willd. Southern Hackberry. Celtis mississippiensis Bose, in part (G), 2 & dn Low wet woods. ZI eentral, and so n Mo.: Marion, Pike, Mont- gomery, Gasconade, St. Charles, St. Louis, J Gesn Perry, Cape Girardeau, Seott, Mississippi, New e liri, Dunklin, Butler, Ripley, Shannon, Iron, Cole, Boone, Howard, Jackson, Bates, St. Clair, Jasper, McDonald, Stone, and Taney counties Celtis laevigata var. texana (Scheele) Sarg. Bot. Gaz. 67: 223. 1919. Celtis mississippiensis Bose, in part (G), (B & B). Rocky woods, glades, and bluffs. Circumneutral, Central and southern Mo., 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 523 mainly in the Ozark region: St. Charles, Perry, Stoddard, Butler, Pres “eit Shannon, Ozark, Taney, Stone, Douglas, ee ae Pulaski, Cam en Miller, Moniteau, Montgomery, Boone, Jackson, , Greene, St. Cali, Cedar, Lawrence, Jasper, Newton, and MeDonald iuc Celtis pumila Pursh. Dwarf Hackberry. Rocky woods. Calciphile to eireumneutral Central and southern Mo.: St. Louis, Madison, Carter, Shannon, Dent, Jackson, Benton, Taney, Barry, Vernon, and Newton counties. Celtis pumila var. georgiana (Small) Sarg. Bot. Gaz. 67: 227. 1919. Celtis georgiana Small (B & B). eky woods, aes and bluffs. Central and southern Mo.: Ralls, Lincoln, ORAT St. Francois, Ste. Genevieve, Madison, Iron, Wayne, Carter, Ore- gon, Shannon, Dent, pape. Maries, Miller, Camden, Benton, Polk, Dallas, Henry, Hiekory, Ozark, Taney, Stone, St. Clair, Cedar, Vernon, Jasper, New- ton, MeDonald, and Barry counties. i Celtis "eeu var. Deamii Sarg. Bot. Gaz. 67: 22 cky woods and glades. Cireumneutral. bouibenstorn Mo., Iron Mountain "e -region: St. Francois a Iron counties. Fam. MORACEAE Maclura pomifera (Raf.) Sehneider. Osage Orange, Bodare. Pastures, Mna and borders of woods. Caleiphile to circumneutral. Prob- ably not native in the state, but formerly much planted for hedges and freely escaping. deed but yid and T from many of the Ozark counties: Adair, Shelby, Audrain, St. Louis, Franklin, Gasconade, Iron, New Madrid, Pemiscot, Howell, Laclede, Living. crc s Dekalb, mec de Holt, Jack- son, Johnson, Bates, Vernon, St. Clair, Greene, and Jasper counties. BROUSSONETIA PAPYRIFERA (L.) Vent. Paper Mulberry. Sometimes planted as a shade tree and occasionally escaping. Johnson and Butler counties. Morus rubra L. Red Mulberry. Low and upland woods. Cireumneutral. General and probably in every county. MORUS ALBA L. White Mulberry. Commonly planted and freely escaping. Indifferent. big: E but seattered: Clark, Marion, Ralls, Pike, Lincoln, n Louis, Franklin, Mercer, Harrison, Sulli- van, Chariton, Boone, Atchison, Holt, adii Dekalb, J aekson, Johnson, Bates, ernon, Barton, Jasper, and MoDonail counties. CANNABIS SATIVA L. mp. Thickets, open and waste ground. Circumneutral. General. Humulus Lupulus L. Hop. Fertile open and waste ground. Circumneutral, General, HUMULUS JAPONICUS Sieb. € Zuec. Japanese Ho Fertile open ground and waste plaees. different. Scattered: Jackson and Jasper counties. [VoL. 22 524 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fam. URTICACEAE URTICA URENS L. Dwarf Nettle. Introduced in waste ground. Indifferent. Western Mo., local: Jackson Co. Urtica procera Muhl. See Rh. 28: 192-195. 1926. ay Nettle. Urtica gracilis of most Am. auth., not Ait. (G), (B & B). Waste or cultivated ground, a tel or alluvial Zi and fertile soils. Cir- cumneutral. Northern and central Mo.: Marion, Lincoln, St. Charles, St. Louis, Boone, Chariton, Saline, roig Livingston, Mercer, Harrison, Worth, Knox, Adair, Shelby, Macon, Gentry, Atchison, Holt, Buchanan, Andrew, Platte, Clay, and Jaekson counties. Urtica chamaedryoides Pursh. Low open ground. Cireumneutral. Southeastern Mo.: Pemiscot Co. Laportea canadensis (L.) Gaud. Wood Nettle. Low rich woods. Cireumneutral. General. Pilea pumila (L.) Gray. Clearweed, Richweed Low woods and boggy ground along streams, and about ponds. Circumneutral. General, cylindrica (L.) Sw. False Nettle. Open woods, thickets, and moist open ground. Cireumneutral. General. Parietaria pennsylvanica Muhl. Pellitory. Open woods, thickets, and moist open ground. Cireumneutral. General. Parietaria obtusa Rydb. Rocky open woods and glades. Cireumneutral. Local: Ozark Co. Fam. SANTALACEAE Comandra Richardsiana Fernald. Bastard Toadfla Dry roeky woods, prairies, and barrens. Oxylophile to eireumneutral. Gen- eral, but commonest in central and southern Mo. This has been long confused with C. umbellata, a northern species which does not reach Mo. Fam. LORANTHACEAE Phoradendron flavescens (Pursh) Nutt, zu Parasitie on deciduous trees; low wo and banks of streams. Southern Mo.: Seott, Mississippi, New Madrid, head — Butler, Ripley, Carter, Reynolds, Shannon, Oregon, Ozark, and Stone eoun Fam. AnISTOLOCHIACEAE Asarum canadense L. var. acuminatum Ashe. Wild Ginger. Rich woods and along base of rocky bluffs. Circumneutral. General. Asarum canadense var. reflexum (Bicknell) Robinson. In similar situations to the preceding. General Aristolochia Serpentaria L. Virginia Snakeroot. ey woods and alluvial thickets. Circumneutral. Central and southern St. Louis, Jefferson, Franklin, Cape Girardeau, Bollinger, New Madrid, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 525 Dunklin, Butler, Ripley, Carter, Iron, Oregon, Shannon, Dent, Gasconade, Howell, Texas, Maries, Pulaski, Boone, Saline, Jackson, Henry, Wright, Webster, Greene, Christian, Douglas, Ozark, Taney, Stone, Barry, wane and MeDonald counties. Aristolochia Serpentaria var. hastata (Nutt.) D Low swampy woods. Circumneutral. South iare Mo. local: Dunklin Co. Aristolochia tomentosa Sims. Pipe Vine, Dutehman's Pipe. and thickets along streams. Cireumneutral. East-central and eR Mo: St. Louis, Jefferson, Franklin, Madison, Wayne, Butler, Dunklin, Ripley, Oregon, Shannon, Phelps, Pulaski, Texas, Maries, Wright, Douglas, Ozark, Greene, Stone, Lawrence, Jasper, Newton, and MeDonald eounties. Fam. POLYGONACEAE Eriogonum longifolium Nutt. Roeky xc woods and glades. Oxylophile. Southern Mo., rare and loeal: Oregon RUMEX ELONGATUS Gus Waste open Spe Indifferent. Scattered: Boone, Clay, and Jackson counties. RuMEX PATIENTIA L. var. KURDICUS Boiss. Patience Dock. Waste ground. Indifferent. Introduced in Clay and Jackson counties. RuMEX BRITANNICA L. Great Water Dock. Low ground along Missouri River. Cireumneutral. Local: St. Louis Co. RUMEX cRISPUS L. Narrow-leaved Dock, Curle k. Fields and waste ground. Indifferent. General and common. exicanus Meisn. Moist alluvial ground. Indifferent. Seattered: Clark, Knox, St. Louis, Saline, Mercer, Carroll, Clay, Jaekson, and Holt counties. Rumex altissimus Wood. Pale Dock. Fields, thickets, waste, and alluvial ground. Circumneutral. General, Rumex verticillatus L. Swamp Dock. amps and low wet woods. Cireumneutral. General, RUMEX SANGUINEUS L. Waste Miis. Indifferent. Locally introduced in St. Louis Co. RUMEX OBTUSIFOLIUS L. Bitter Dock, Broad-leaved Dock, Fields, waste ground, and about dwellings. Indifferent. General. Rumex maritimus L. var. fueginus (Philippi) Dusén. See Rh. 17: 80-81. Sand bars and alluvial ground along the larger rivers. Circum dor o pss - tral and northwestern Mo.: St. Louis, Franklin, Perry, Boone, Saline, Carroll, Holt, Buchanan, Clay, and Jackson counties. This has been confused with Rumex persicaroides, which is not known in Mo. Rumex hastatulus Baldw. ocky prairies, glades, and sandy open ground. Oxylophile. Scattered in east-central and southern Mo.: St. Louis, Ste. Genevieve, Scott, Dunklin, and Newton counties. [Vor. 22 526 ANNALS OF THE MISSOURI BOTANICAL GARDEN RUMEX ACETOSELLA L. Sour Dock, Sheep Sorrel. Fields and waste ground. Oxylophile. General. RUMEX CRISPUS X OBTUSIFOLIUS. Sometimes occurring locally with the parent species. St. Louis, Jefferson, and Carroll counties. Polygonum exsertum Small. ow alluvial meadows, prairies, and muddy banks. Cireumneutral. General. Polygonum prolificum (Small) Robinson. Open ground. Indifferent. Local: Clay and Jackson eounties. Polygonum aviculare L. Knotweed, Knot-grass. Fields, waste ground, and about dwellings. Indifferent. General and common. Polygonum aviculare var. vegetum Ledeb In similar situations to the typical form. Indifferent. Scattered. Polygonum aviculare var. angustissimum Meisn. Polygonum neglectum Bess. (B € B). With the typical form and often commoner. Indifferent. General. Polygonum buxiforme Small. Polygonum littorale of auth., not Link (B € B). Polygonum aviculare var. littorale (G). Fields and waste ground. Indifferent. General. Polygonum erectum L. Low wet woods and alluvial banks of streams and ponds. Circumneutral. General. Polygonum ramosissimum Michx. Prairies and open ground, usually in moist alluvial soil. Cireumneutral. General. Polygonum achoreum Blake, Rh. 19: 232. 1917. Introduced in waste ground. Jackson Co. Polygonum triangulum Bicknell in Britton & Brown, Ill. Fl. ed. 2. 1: 664, fig. 1626. 1913. Fields and waste ground. Indifferent. Jackson Co. Polygonum tenue Michx. Rocky prairies, glades, and gravel bars along streams. Oxylophile. Central and southern Mo.: Lincoln, Boone, Montgon "i Warren, St. Charles, Gasconade, Crawford, Jefferson, Ste. Genevieve, St. Francois, Perry, Ma ied Bollinger, Iron, Reynolds, Stoddard, Dunklin, Shan, Fotos, Howell, Pulaski, Maries, Camden inten Polk, St. Clair, Greene, Jackson, Jasper, Newton, MeDonald, a nd Barry counties. Polygonum vs RR L. See Rh. 23: 258-259. 1921. olygonum lapathifolium var. nodosum (Pers.) Weinmann (G). Wet open ground about ponds and sloughs, and in waste ground. Indifferent. General, Polygonum densiflorum Meisn. Low open ground. Cireumneutral. Southeastern Mo.: Scott, Dunklin, and Carter counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 527 Polygonum coccineum Muhl. See Rh. 27: 162. 1925. Polygonum Muhlenbergii (Meisn.) Wats. (G). Persicaria Muhlenbergii (Wats.) Small (B & B). Prairies and open woods, and borders of streams, ponds, and sloughs. Circum- neutral. General. Polygonum coccineum var. pratincola (Greene) Stanford, Rh. 27: 165. 1925. In similar situations to the typical form. Local: Jackson Co. Polygonum natans Eat. Polygonum amphibium of Am. auth., not L. (G). Persicaria amphibia S. F. Gray (B & B In shallow water. Local: Greene Co. Polygonum pensylvanicum L. Wet ground along borders of streams, ponds, and sloughs, and in waste ground. Indifferent. General Polygonum pensylvanicum var. laevigatum Fernald, Rh. 19: 73. 1917. Frequently found with the typieal form. Polygonum longistylum Small. See Rh. 27: 182. 1925. Wet or moist open ground along prairie streams. Cireumneutral. Central and southern Mo.; seattered: New Madrid, Dunklin, Clay, Jackson, and Jasper counties. Polygonum Hydropiper L. var. projectum Stanford, Rh. 29: 86. 1927. Smart- weed, Water Pepper. Polygonum Hydropiper of auth. in part, not L. (G). Persicaria Hydropiper of auth., not Opiz ( B) et woods and prairies, along sloughs and streams. Cireumneutral. General. Polygonum punctatum Ell. Smartweed, Water Pepper. Polygonum acre HBK, not HEN ., in part (G). Polygonum acre var. leptostachyum vu in part (G). Persicaria punctata (Ell.) em (B & B). Wet ground, borders of ponds and epa Circumneutral. General. Polygonum eigen (Small) Fernald, Rh. 23: woods and borders of sloughs. as Southeastern Mo.: Dunklin ii POLYGONUM ORIENTALE L. Prince’s Feather. Waste ground; escaped from cultivation. Indifferent. Scattered: St. Louis, Boone, Miller, Greene, Taney, Clay, Jackson, Cass, and Jasper counties, POLYGONUM PERSICARIA L. Lady’s Thumb. aste and cultivated ground. Cireumneutral. General. Polygonum setaceum Baldw. Swampy open ground and low open woods. Cireumneutral. Southeastern Mo.: Dunklin, Butler, and Ripley counties. Polygonum hydropiperoides Michx. See Rh. 28: 24. 1926. pen ground about ponds, streams, and spring branches. Cireumneutral. General. [Vor. 22 528 ANNALS OF THE MISSOURI BOTANICAL GARDEN Polygonum hydropiperoides f. strigosum (Small) Stanford, Rh. 28: 26. 1926. olygonum hydropiperoides var. e Small (G). Occasionally found with the typical for Polygonum M" Riddell. See Rh. 28: 27. 1920. Persi B). " woods. Cireumneutral. Southern Mo.: Dunklin, Butler, Ripley, Jasper, Newton, and MeDonald pe ies. Polygonum virginian Rich woods. dsl General. Polygonum arifolium L. Halberd-leaved Tear-thumb. et or moist open ground. Circumneutral. Southeastern Mo., local: Stod- dard Co. Polygonum sagittatum L. Arrow-leaved Tear-thumb. Boggy ground about springs and streams. Oxylophile. General but scattered: Putnam, Shelby, Audrain, Boone, Randolph, Lincoln, St. Charles, St. Louis, Jef- ferson, St. Francois, Bollinger, Mississippi, Dunklin, Shannon, Crawford, Saline, Jackson, St. Clair, Greene, Cedar, Vernon, Barton, and Jasper counties. PoLYGoNUM CONVOLVULUS L. Climbing Buckwheat, Black Bindweed. Thickets, waste and cultivated ground. Circumneutral. General but seattered. Polygonum scandens L. Climbing False Buckwhea Thickets, waste and cultivated ground. did General. POLYGONUM CUSPIDATUM Sieb. & Zuce. See Rh. 32: 223. 1930. Japanese Knot- Cultivated and rarely escaped into thickets and waste places. St. Louis Co. FAGOPYRUM ESCULENTUM Moench. Buckwheat. Rich cultivated and waste ground. Escaped from cultivation. General but scatter Polygonella americana (Fisch. & Mey.) Small. Jointweed. Glades and open gee — Oxylophile. Southeastern Mo.: Iron, Stod- dard, and Dunklin coun — re ne Ladies’ Ear-dro w wet woods, and wet thickets. odd Southeastern Mo.: Bollinger, Mississippi, New Madrid, Pemiscot, Dunklin, Stoddard, Butler, and Ripley counties. Fam. CHENOPODIACEAE — atriplicifolium (Spreng.) Coult. Winged Pigweed. ndy fields, rocky or sandy open ground, and along roadways and railways "eie f General but seattered: Clark, Pike, St. Charles, St. Louis, Franklin, St. Franeois, Perry, Stoddard, Dunklin, Randolph, Carroll, Saline, Clay, Jackson, and Jasper counties. KOcHIA TRICHOPHYLLA Stapf. Fire-bush, Summer Cypress Cultivated and occasionally escaped, in fields and along railroads. Scattered: St. Louis, Callaway, McDonald, Gentry, and Andrew counties. 1935] i PALMER € STEYERMARK— PLANTS OF MISSOURI 529 KocHIA scoparia (L.) Schrad. Standing Cypress, Belvedere Escaped from cultivation into waste ground and along railrdada Seattered: St. Louis, Jefferson, Franklin, Andrew, Buchanan, and Jackson counties. KOCHIA PROSTRATA (L.) Schrad. Waste ground. Indifferent. Introduced in Jackson Co, CHENOPODIUM AMBROSIOIDES L. Mexican Tea. Fields, rich waste ground, and about dwellings. Indifferent. General. CHENOPODIUM AMBROSIOIDES var. ANTHELMINTICUM (L.) Gray. Commonly found with the oo CHENOPODIUM CARINATUM R. Waste ground, fields, rar P dwellings, often in roeky or gravelly ground. Indifferent. Central and southern Mo., seattered: Bollinger, Ripley, Reynolds, Shannon, Oregon, Dent, Howell, Texas, Ozark, Douglas, Jackson, and Vernon counties. CHENOPODIUM Botrys L. Feather Geranium, Jerusalem Oak. Waste ground. Indifferent. Scattered: St. Louis, Shannon, Ozark, Stone, and Jackson counties Chenopodium dota (L.) Aseh. Strawberry Blite. Open ground. Cireumneutral. Local: Pike Co. CHENOPODIUM GLAUCUM L. Oak-leaved Goosefoot. aste open ground and moist alluvial ground along ponds and the larger rivers. Calciphile pu cireumneutral. Scattered: St. Louis, Clinton, Holt, and Jackson counties. Chenopodium gigantospermum Aellen in Fedde, Rep. Spec. Nov. 26: 144. 1929. Maple-leaved Goosefoot. Chenopodium hybridum of Am. auth., not L. (G), (B € B). Rich waste and cultivated ground. Indiffe rent. General. Chenopodium gigantospermum var. Standleyanumi Aellen in Fedde, Rep. Spec. Nov. 26: 147. 1929 Waste paak Indifferent. St. Louis Co. CHENOPODIUM ALBUM L. Pigweed, Lamb's Quarter aste and cultivated ground. Indifferent. General and common. The young shoots eaten as greens. CHENOPODIUM ALBUM f. VIRIDE (L.) Aellen in Fedde, Rep. Spec. Nov. 26: 129. 1929. Chenopodium album var. viride (L.) Moq., in part (G). Chenopodium album L., in part (B & B). Waste ground, in rich or moist soil. Indifferent. General. CHENOPODIUM ALBUM f. LANCEOLATUM (Muhl.) Aellen in Fedde, Rep. Spec. Nov. 26: 130. 1929. aste and low alluvial ground. Sometimes found with the typical form. Scattered. CHENOPODIUM PAGANUM Reichenb. "vec album var. viride (L.) Moq., in ei (GF Chenopodium album of auth., not L., in part Waste and cultivated ground. Indifferent. nuls bat scattered. [Vor. 22 530 ANNALS OF THE MISSOURI BOTANICAL GARDEN Chenopodium pratericola Rydb. Bull. Torr. Bot. Club 39: 310. 1912. Waste and cultivated ground. Indifferent. Jackson Co. Chenopodium Berlandieri Moq. Waste ground. Indifferent. General but scattered. Chenopodium Berlandieri subsp. Zschackei var. typicum f. latifolium (Ludwig) Aellen in Fedde, Rep. Spee. Nov. 26: 55. 1929. Waste ground. Jackson Co. Chenopodium Berlandieri subsp. aren var. typicum f. angustius (Ludwig) Aellen in Fedde, Rep. Spee. Noy. 26: 55. 1929. Occasionally found with the het pos Indifferent, Scattered: Jackson and Jasper, and probably in many other countie Chenopodium Berlandieri subsp. Zschackei var. —-—— f. pedunculare Aellen in Fedde, Rep. Spec. Nov. 26: 55. 1929 Waste ground. Indifferent, Jackson Co. Chenopodium Berlandieri ioi Zschackei var. typicum f. neglectum Aellen. in Fedde, Rep. Spec. Nov. 26: 55. 1929, Waste ground. deni Jackson Co. Chenopodium Berlandieri subsp. Zschackei var. farinosum Aellen in Fedde, Rep. Spec. Nov. 26: 56. 1929, Waste ground. Indifferent. Jackson Co. Chenopodium Berlandieri subsp. Zschackei var. foetens (Ludwig) Aellen in Fedde, Rep. Spec. Nov. 26: 57. 1929. Waste ground. Indifferent. Jackson Co. ne en subsp. Boscianum (Moq.) Aellen in Fedde, Rep. Spec. Nov. 26: 51. 1929, i Boscianum Moq. (G), (B & B). ods and thickets. Indifferent. Southern and central Mo., scattered: St. Louis, Jackson, Greene, and Jasper counties. Chenopodium linde Aellen in Fedde, Rep. Spee. Nov. 26: 153. 1929. Chenopodium Boscianum Moq., in part e ground. Indifferent. "cin and western Mo.: Shannon, Holt, Jackson, Greene, and Jasper counties. Chenopodium missouriense Aellen, Bot. Not. 1928: 206, figs. 1928. Chenopodium paganum Standley, N. Am. Fl. 21: 1, 1916, not Reichenb. Waste and cultivated ground. Indifferent. General, but seattered: St. Louis, Jackson, and Jasper counties, and probably in many others. M missouriense var. Bushianum Aellen in Fedde, m» Spec. Nov. 26: 929. poe ground. Indifferent. Jackson Co. Chenopodium glaucophyllum Aellen in Fedde, Rep. Spec. Nov. 26: 155. 1929. Waste ground. Indifferent. Jackson Co. Chenopodium Bushianum Aellen in Fedde, Rep. Spec. Nov. 26: 63. 1929. Rocky open and waste ground. Indifferent. Western Mo.: Jackson, St. Clair, Greene, and Ozark eounties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 531 Chenopodium leptophyllum Nutt. ocky or sandy open ground. pee Western Mo.: Jackson and Jasper counties, and probably in many o Chenopodium leptophyllum var. scio (Murr.) Thellung & Aellen in Fedde, Rep. Spec. Nov. 26: 134. 1929 Chenopodium leptophyllum var. oblongifolium Wats. (G). E leptophyllum Nutt. in part (B & B). and eultivated ground. Indifferent. General but scattered: St. Louis, J oibus Jasper, and probably in many other counties. Chenopodium incanum (Wats.) Heller. Open waste ground. Indifferent. Introduced from the western plains. Jackson Co. Chenopodium dacoticum Standley, N. Am. Fl, 21: 22, 1916. Prairies and fields. Indifferent. Western Mo.: Atchison and Jackson counties. CHENOPODIUM MURALE L. Waste ground and about dwellings. Indifferent. General but seattered: Warren, St. Louis, Texas, Boone, Jackson, Greene, Taney, and Jasper counties. CHENOPODIUM URBICUM L. Waste ground, dumps, and about dwellings. Indifferent. General. CHENOPODIUM URBICUM Var. INTERMEDIUM (Mert. € Koch) Koch. Commonly occurring with the typical form. x Chenopodium variabile var. Murrii Aellen in Fedde, Rep. Spec. Nov. 26: 157. 1929 A Spp hybrid between Chenopodium album and C. Berlandieri subsp. Zschackei var. ty Waste ground. J x kb Co. ATRIPLEX ROSEA L. Red Sea Waste and saline im " Caleiphile. Central Mo.: Howard and Jackson counties. Atriplex patula L. Spear Scale. Rich cultivated or waste ground. Indifferent. Northern and central Mo.: Knox, St. Louis, Putnam, and Jackson counties. ae mn var. hastata (L.) G eky o te ground, and 2 saline springs and licks. Indifferent. a "Bt Louis, Callaway, Boone, Randolph, Jackson, and Jasper counties. a argentea Nutt. Silver Scale, open ground. Indifferent. Introduced in Jackson Co. "ut Nuttalliana (R. & S.) Wats. Introduced in waste ground. Jackson Co. AXYRIS AMARANTOIDES L. See Rh. 29: 223. 1927. Introduced in waste ground. Jackson C Corispermum nitidum Kit. Bug-seed. andy open ground. absent. Introduced in Jackson Co. [Vor. 22 532 ANNALS OF THE MISSOURI BOTANICAL GARDEN Corispermum hyssopifolium L. Bug-seed. Sand bars of the large rivers. ir AN Beattered: St. Louis, Perry, Clay, and Jaekson counties. Corispermum orientale L. var. emarginatum (Rydb.) Maebride, Contr. Gray Herb. N. S. 53: 13. 1918 Sandy open ground. O Introduced in Jackson Co. Suaeda depressa (Pursh) Wats. a Blite. open ground. Oxyl Me Introdueed in Jackson Co. Salsola Kali L. Open waste ground. Introduced in Jackson Co. SALSOLA KALI var. TENUIFOLIA G. F. W. Mey. Russian Thistle. Dry or rocky open ground and waste places. Caleiphile to eireumneutral. Introduced from farther west. Seattered: St. Louis, St. Francois, Saline, Carroll, Atehison, Holt, Andrew, Buchanan, Platte, Clay, Jackson, and Jasper counties. Fam. AMARANTHACEAE Amaranthus Palmeri Wats. Waste ground and along railroads. Indifferent. Introduced in Jackson Co. AMARANTHUS HYBRIDUS L. Pig-weed, Green Amaranth. Fields, gardens, and waste ground. Indifferent. General and common. The young shoots eaten as greens. AMARANTHUS HYBRIDUS var. HYPOCHONDRIACUS (L.) Robinson. Prince’s Feather. Cultivated and rarely escaped. Scattered AMARANTHUS RETROFLEXUS L. Pig-weed, Green Amaranth. Cultivated and waste ground. Indifferent. General AMARANTHUS PANICULATUS L. Purple Amaranth. Escaped from gardens to roadsides and waste ground. Uncommon and scat- tered: St. Louis, Saline, and Jackson counties, and reported from Boone Co. Amaranthus graecizans L. Tumble-weed. Fields, waste places, and rocky open ground. Indifferent. General. Amaranthus graecizans var. pubescens Uline & Bray. Waste and cultivated ground. Occasionally growing with the typical form. Jackson Co. Amaranthus blitoides Wats. Spreading Pig-weed Waste and cultivated pie Indifferent. General. AMARANTHUS SPINOSUS L. Spi Waste and cultivated mood. Fairs General. Acnida tuberculata Moq. Water Hemp. Wet ground about sloughs, en and streams. Circumneutral. Scattered, but eommonest along the large rivers. Acnida tuberculata var. prostrata Wana & Bray) Robinson. and bars and mud flats along streams, and in waste ground. Circumneutral. Scattered: Pike, St. Louis, and Jackson counties, 1935] PALMER & STEYERMARK— PLANTS OF MISSOURI 533 Acnida tuberculata var. subnuda Wats. ommonly occurring with the typical form. Acnida tamariscina (Nutt.) Wood. Water Hemp. Cultivated fields and waste ground, mostly in moist alluvial soil. Cireum- neutral. General. Cladothrix lanuginosa Nutt. Introduced from the Southwest. Waste ground. Jackson Co. Iresine paniculata (L.) Ktze. Blood-leaf, Juba's Bush. w wet woods and thiekets. Central and southern Mo., seattered: St. Louis, Jefferson, Franklin, Ripley, Dent, Pulaski, Maries, Ozark, Jasper, Newton, and MeDonald counties. Froelichia floridana (Nutt.) M Froelichia campestris ET (B & B Sandy open ground and along REISEN Oxylophile. Eastern and central Mo., mostly near the larger rivers: Lewis, St. Louis, Madison, Scott, Mississippi, unklin, Butler, Reynolds, Boone, and Jackson counties. Froelichia gracilis Moq. Dr de ground and sandy barrens. Oxylophile. Eastern, central, and southern : Marion, Lincoln, St. Charles, St. Francois, Perry, Bollinger, ayne, Dunklin, Howard, Monltenks Jackson, and Jasper counties. Fam. PHYTOLACCACEAE Phytolacca americana L. See Rh. 17: 180. 1915. Pokeweed, Pokeberry. Phytolacca decandra L. (G). Fields, roadsides, and waste ground. Indifferent. General and probabl every county. The young shoots eaten as greens by the country people, and in juice of the ripe berries formerly used as ink. Fam. NYCTAGINACEAE Oxybaphus nyctagineus (Michx.) Sweet. Wild Four-o’clock. Allionia nyctaginea Michx. (B & Open and waste ground. Oben General. Oxybaphus floribundus Chois. Umbrella-wort. Rocky open slopes and glades. Oeleiph ile. General but scattered: Clark, St. Louis, aa Wayne, Ripley, Dent, Holt, Carroll, Jackson, and Jasper eountie bs hirsutus (Pursh) Sweet. y open ground and glades. Circumneutral. Scattered: Nodaway, Boone, Ozark, and Jackson counties. Oxybaphus hirsutus var. at Ap ae Chois. Ozybaphus pilosus (Nutt.) S Glades and rocky open P ' Caleiphile, Southwestern Mo.: MeDonald 0. (VoL. 22 534 ANNALS OF THE MISSOURI BOTANICAL GARDEN €— albidus (Walt.) IO Umbrella-wort. ia albida Walt. (B & B). a bracteata Rydb. ` Allionia lanceolata of Standley, not Rydb. n woods, prairies, glades, and open ground. Calciphile to circum- neutral olt, Worth, bey, Dekalb, St uis, Warren, Montgo ery, Mississippi, Stoddard, Ripley, Shannon, Maries, Crawford, Dent, Phelps, aski, Boone, Clinton, Jackson, Dallas, Polk, Hickory, Laclede, Johnson, Henry, Greene, Ozark, Vernon, Jasper, and McDonald counties, Oxybaphus Unearis (Pursh) Robinson. phus ü Bri ocky prairies and nc Caleiphile. Central and southern Mo.: St. Louis, Ste. Genevieve, Washington, Boone, Pulaski, Wright, Taney, Jaekson, Benton, Jasper, Newton, and Barry counties. Fam. ILLECEBRACEAE Anychia polygonoides Raf. Forked Chickweed Rocky open woods and glades. Oxylophile. General: Mercer, Sullivan, Linn, Randolph, Shelby, Monroe, Audrain, Boone, Lincoln, St. Louis, Jefferson, St. Francois, Mississippi, Pemiscot, Dunklin, Ripley, Orii Carter, Shannon, Reynolds, Iron, Dent, Texas, Phelps, Pulaski, Maries, Miller, Camden, Laclede, Hickory, Dallas, Greene, Ozark, Jackson, Henry, Cedar, Jasper, Newton, and McDonald counties. Anychia canadensis (L.) BSP. open woods. Oxylophile to cireumneutral. General: Sehuyler, Putnam, Sullivan, Knox, Shelby, Ralls, Boone, Warren, St. Louis, Jefferson, Ste. Gene- vieve, St. leni Bollinger, Wayne, Ripley, Shannon, Reynolds, Dent, Phelps, Howell, Douglas, Taney, Christian, Dallas, Webster, Greene, Hickory, Dekalb, Clinton, Jaekson, Johnson, Vernon, Jasper, MeDonald, and Barry counties. Fam. AIZOACEAE MOLLUGO VERTICILLATA L. Carpet-weed. Waste and cultivated ground. Indifferent. General and common. Geocarpon minimum Mack. Torreya 14: 67. 1914. Sandy barrens. Oxylophile. Rare and local: Jasper Co. Fam. CARYOPHYLLACEAE ligne ARVENSIS. Corn Spurrey. Waste and cultivated ground. Indifferent. Scattered: Jackson, Greene, and Jasper yo Sagina decumbens (Ell.) T. & G. Pearlwort. Waste ground, fallow fields, and rocky open or sandy ground, usually in alluvial or sandy soils. Oxylophile to circumneutral. Central and southern Mo.: Boone, Callaway, St. Louis, Jeff Franklin, Ste. Genevieve, St. Francois, Bollinger, Madison, Dunklin, Butler, Binley, Ore, Reynolds, Wayne, 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 535 Dent, Shannon, Texas, Osage, Phelps, Maries, Pulaski, Miller, Camden, Dallas, Laclede, Polk, Webster, Henry, Douglas, Greene, Cedar, Saline, Vernon, Jasper, and Newton countie ARENARIA SERPYLLIFOLIA L. Thyme-leaved Sandwort. Sandy or rocky soil, nn an ledges, and open places. Caleiphile to eir- eumneutral. General and co Arenaria patula Michx. Sandwort. Glades and roeky prairies. Caleiphile to cireumneutral. Central and southern Mo.: Boone, Callaway, St. Louis, Jefferson, Franklin, Gaseonade, Osage, Wash- S St. Franeois, Ste. Genevieve, Perry, Iron, Reynolds, Dunklin, Ripley, Ore- mden gon, Shannon, Maries, Texas, Phelps, Pulaski, Laclede, Camden, Miller, Cole, Hic cad d bg. Webster, Douglas, Christian, Taney, Omk, Stone, Polk, t. Clair, Cedar, Barry, Jui, Vernon, Jasper, Newton, and eDo udin counties. Arenaria stricta Michx. var. texana Robinson. Rock Sandwort. Roeky slopes, ledges along bin and glades. Calciphile to ann astern, central, and southern Mo.: Monroe, Lincoln, Jefferson, St. Francois, Washington, Boone, Camden, ona Greene, Christian, and McDonald na Stellaria ipsius Miehx. var. silvatica (Béguinot) Weatherby, Rh. 26: 171. 1924. reat Chickw Alsine ME (Miehx. ) Britton, in part (B & B). Shaded slopes along bluffs. Circumneutral. Eastern Mo., local: Franklin Co. STELLARIA MEDIA (L.) Cyrill. Chickweed. Alsin B&B) media L. ( : Waste ground, "ims and cultivated ground. Indifferent. General. Stellaria longifolia Muh g-leaved Stitchwort. Alsine longifolia nb piv! n (B & B). Low open woods and moist meadows. Cireumneutral. Eastern Mo.: Seotland, St. Louis, Jefferson, Madison, and Cape Girardeau counties. STELLARIA GRAMINEA L. Narrow-leaved Stitehwort. ne graminea (L.) Britton (B & B). n meadows, and waste ground. Indifferent. Introduced in Boone Co. CERASTIUM VULGATUM L. var. HIRSUTUM Fries. See Rh. 22: 178. 1920. Mouse- ckweed Cerastium vulgatum of auth., not L. (G), (B & B). Waste and cultivated ground, and moist meadows. Indifferent. General and common. sag brachypodum (Engelm.) Robinson Rich open woods, rocky slopes, moist nn; waste and eultivated ground. Cireumneutral. Commonest in eastern, central, and southern Mo. and locally north in Sehuyler and Atchison counties. Cerastium nutans Raf. Nodding Chickweed. Moist meadows, rich rocky woods, cultivated and waste ground. Indifferent. General. A [VoL. 22 536 ANNALS OF THE MISSOURI BOTANICAL GARDEN CERASTIUM VISCOSUM L. Clammy Chickweed. ‘ Moist alluvial and cultivated ground, and wet rocky woods. Oxylophile to eireumneutral. General Cerastium arvense L. var. oblongifolium ( Torr.) Holliek & Britton Low wet ground, and on moist siliceous or granitic rocks. Oxylo phile. East- ern Mo., south of Missouri River: St. Louis, Jefferson, St. Francois, Ste. Gene- vieve, Madison, Iron, and Wayne counties. AGROSTEMMA GITHAGO L. Corn Cockle. Cultivated and waste ground, and along roadsides and railroads. Indifferent. eneral, DIANTHUS DELTOIDES L. Maiden Pink. Introduced in waste ground. Franklin Co. DIANTHUS ARMERIA L. Deptford Pink. Roeky open woods, sandy open and waste ground. Oxylophile to cireum- neutral. Central and iban Mo. south and east of a line from Lineoln, Mont- gomery, Boone, and Henry counties to McDonald Co, DIANTHUS BARBATUS L. Sweet William. Cultivated and rarely escaped to roadsides and waste ground. Jasper Co. LYCHNIS ALBA Mill, White Campion. Waste and ge > nd. Indifferent. Seattered: Harrison, Marion, Lin- coln, Warren, St, , Franklin, Mio Pulaski, Howell, Texas, Boone, Saline, Clay, er d Jasper counties LYCHNIS DIOICA L. Red Campion. Escaped into waste ground. Jackson Co. This has sometimes been mistaken for Silene noctiflora. Silene antirrhina L. Sleepy Catehfly Field, open woods, and cultivated and waste ground. Indifferent. General. Silene antirrhina var. divaricata Robinson. Occasionally occurring with the typical form. Holt and Jackson counties. Silene nivea (Nutt.) Otth. Moist or wet banks of streams, and also introduced in waste ground and along railroads. Cireumneutral. Northern and central Mo.: Sullivan, Boone, and Audrain counties. SILENE LATIFOLIA (Mill) Britten & Rendle. Bladder Campion. Introduced in Platte and Jackson counties. SILENE NOCTIFLORA L. Night- SIME Catehfly. Waste and eultivated ground. Indifferent. Scattered: St. Louis, Franklin, Boone, Saline, and Jasper eounties. SILENE GALLICA L. Old fields and waste ground. Indifferent. Cape Girardeau and Jackson counties, Silene virginica L. Indian Pink, Fire Pink. Rocky woods and along moist cliffs and banks. Oxylophile to T East-central and southern Mo., mostly in the Ozark region: Mont mery, Jef- ferson, Ste, Genevieve, St. Francois, Washington, Crawford, Bollinger, Madison, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 537 Wayne, Carter, Butler, Iron, Reynolds, Shannon, Dent, Oregon, Howell, Texas, Wright, Webster, Greene, Douglas, Ozark, Taney, Stone, Barry, and McDonald counties. Silene regia Sims. Royal Catchfly. Rocky prairies, rocky open woods, and thickets. Circumneutral. East-central and southern Mo., Ozark region: St. Louis, Jefferson, Franklin, Washington, Crawford, Dent, Shannon, Texas, a Pulaski, Maries, Miller, Cole, Greene, Barry, Jasper, and McDonald counties Silene stellata (L.) Ait. f. Starry Campio Rich upland woods and thickets. IL ental General, Silene Wherryi Small, Torreya 26: 66, fig. Rocky chert woods and bluffs. S to eireumneutral. Central Ozark region, southern Mo.: Phelps, Dent, Texas, Shannon, Pulaski, Laclede, and uglas counties. Silene Wherryi x virginica. Rocky chert woods, deb with the parent species. Local: Shannon Co. SAPONARIA OFFICINALIS L. Bouncing Bet, Soapwort. Cultivated in old gardens ep freely escaped into fields, por bars, and waste ground, and along roadsides and railroads. Indifferent. Genera SAPONARIA VACCARIA L. Cow-herb. n waste ground, and along railroads and roadsides. Indifferent. Scat- Nis Marion, Ralls, Greene, Jackson, and Jasper counties. Fam. PoRTULACACEAE Claytonia virginica L. Spring Beauty. Rich or roeky woods and prairies. Cireumneutral. General. Claytonia virginica T robu sta eines à Palmer & Steyermark, comb. nov. Occasionally found with the typical form, growing in rich woods. Circum- neutral Scattered: Ste. Genevieve, Madison, Boone, Cooper, Jackson, Douglas, and Jasper counties. Talinum parviflorum L. Glades and barrens. Oxylophile, usually on drag or chert and occa- sionally on leached limestone. Southern Mo.: St. , St. Louis, Jefferson, Ste. Genevieve, Madison, Shannon, Texas, Polk, b Cedar, Dade, Lawrenee, Vernon, Jasper, and Newton counties. Talinum calycinum Engelm. Rock Pink, Flame Flower Glades and exposed rocky ledges. Oxylophile to circumneutral; usually found on sandstone, loin or granitie rocks, but oceasionally on limestone. Central and southern Mo.: Montgomery, Lincoln, St. Louis, Jefferson, Franklin, St. Francois, Ste. Genevieve Madison, Iron, Ripley, Shannon, Texa asconade, Wright Laclede, Pulaski, Maries, Miller, Dallas, Cooper, uae, Polk, Hickory, St. Clair, Cedar, Me Ozark, dioit Stone, Barry, Jaekson, Vernon, Jasper, and Newton coun [Vor. 22 538 ANNALS OF THE MISSOURI BOTANICAL GARDEN — OLERACEA L. Purslane, Pursley. ated and waste ground. Indifferent. General and common. pura E Mack. & Bus alluvial fields and prairies. Western Mo.: Jackson, Johnson, and Jasper Md Portulaca retusa Engelm. Notched Purslane. Limestone and chert glades. Oxylophile to eireumneutral. Southwestern Mo.: Jasper and Newton counties. Portulaca parvula Gray. See N. Am. Fl. 21: 334. 1922, Portulaca pilosa of auth., not L. (G), (B & B). Rocky prairies and gla ten. Oxylophile to eireumneutral. Western Mo., south of Missouri River: Jackson, Polk, Greene, Barton, Jasper, and Newton counties. PORTULACA GRANDIFLORA Hook. Portulaea, ‘Rose Moss. Common in cultivation and occasionally escaped into waste or open ground. Fam. CERATOPHYLLACEAE Ceratophyllum demersum L. Hornwort. Sloughs, sluggish streams, and ponds. Cireumneutral. General but scattered: St. Charles, St. Louis, Perry, Dunklin, Reynolds, Oregon, Pulaski, Texas, Boone, Saline, Jackson, Barton, and Jasper counties. Fam. NyMPHAEACEAE Nymphozanthus advena (Ait.) Fernald, Rh. 21: 186. 1919. Spatter Dock, Yel- low Pond-lily. Nymphaea advena Ait. (G), (B & B). Ponds, sloughs, and slow streams. General but scattered: Lincoln, St. Charles, St. Louis, Jefferson, Franklin, Dunklin, Butler, Laclede, Bates, Vernon, Barton, Jasper, and Newton counti Nymphozanthus ozarkanus (Miller & Standley) Palmer & Steyermark, comb, n Nymphaea ozarkana Miller & Standley, Contr. U. S. Nat. Herb. 16: 91. 1918 Ponds, sloughs, and streams. En Southern Mo., mostly in the eastern Ozark region: Washington, Dent, Iron, Reynolds, Carter, Ripley, Ore- gon, Shannon, Texas, Howell, Douglas, Dallas, Greene, and Taney counties. Nymphaea tuberosa Paine, See Rh. 18: 120. 1916. "Tuberous Water-lily. Castalia tuberosa (Paine) Greene (G), (B & B). Slow streams and ponds. Circumneutral. Eastern Mo., scattered: St. Louis and Dunklin counties. Nymphaea odorata Ait. Water-lily, sweet-scented biet -lily. Castalia odorata (Ait.) Woodville & Wood (G), (B Ponds and sloughs. Circumneutral. Scattered: ^ ER Butler, Boone, Ray, Vernon, Barton, and Jasper counties. Often cultivated and escaped in aces Nelumbo pentapetala lod Fernald, Rh. 36: 23-24. 1934. Yanquapin, Water Chinquapin, Yellow Nelumbo lutea 2; y Pers. (G), (B & B). 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 539 Lakes, ponds, and sloughs. Cireumneutral. General but scattered: Harrison, Adair, Audrain, Boone, Lineoln, St. Charles, des Louis, Stoddard, Dunklin, Carter, Shannon, Phelps, Laclede, Camden, Webster, Mp Dade, Buchanan, Platte, Jackson, Cass, Bates, Barton, Jasper, and Nord counties. NELUMBO NUCIFERA Gaertn. Oriental Lotus. Introduced in a lake in Iron Co. Cabomba caroliniana Gray. Carolina Water-shield. Swamps and sloughs. Cireumneutral Eastern Mo.: St. Charles, St. Louis, Wayne, Dunklin, and Butler counties. Brasenia Schreberi Gmel. Water-shield. Ponds and sloughs. Cireumneutral. East-central and southern Mo., scat- ed: Audrain, St. Charles, St. Louis, St. Francois, Iron, Texas, Taney, Barton, and Jasper counties. Fam. RANUNCULACEAE Ranunculus aquatilis L. var. capillaceus DC. White Water Crowfoot Sloughs, ponds, and streams. Circumneutral. Central and s onthe n Mo.: St. Louis, Crawford, Butler, Ripley, Oregon, Shannon, Pulaski, Howell, Bii: Dallas, Greene, Jackson, Jasper, and Newton counti Ranunculus longirostris Godro Ponds. Cireumneutral. Local: St, Louis Co. Ranunculus delphinifolius Torr. Yellow Water Crowfoot. Ranunculus delphinifolius var. terrestris (Gray) Farwell, a mere terrestrial state of the above Swamps, bayous, ponds, and ar ig in shallow water or on muddy banks. Cireumneutral. Seattered, mostly near the bc rivers and absent from most of the Ozark region: Shelby, Ralls, St. ps , St. Louis, Jefferson, Madison, Bollinger, Scott, Dunklin, Butler, Livingston, Pisis and Jackson counties. Ranunculus Cymbalaria Pursh f. hebecaulis Fernald, Rh. 16: 162. 1914. Sea- side Crowfoot. Sand bars and sandy open ground. Calciphile to circumneutral. Western Mo., northward from the Missouri River: olt, Clay, and Jackson counties. Ranunculus laxicaulis (T. & G.) Darby. "Water Plantain Spearwort. Moist open ground and borders of sloughs and ditches. Circumneutral. Scattered: Montgomery and Greene counties. Ranunculus oblongifolius Ell. Spearwort. ow wet woods and swamps. Oxylophile to eireumneutral. East-central and southern Mo.: St. Charles, St. Louis, Jefferson, Mississippi, New Madrid, Dunk- lin, Howell, Butler, Webster, Greene, Barton, and Jasper counties. Ranunculus pusillus Poir. wamps and low wet woods. Circumneutral. Eastern Mo., south of the Missouri River: St. Louis, Iron, Mississippi, New Madrid, Dunklin, Butler, and Texas counties. Ranunculus sceleratus L. Cursed Crowfoot. Moist alluvial open ground and sand bars. Northern and central Mo., mostly [VoL. 22 540 ANNALS OF THE MISSOURI BOTANICAL GARDEN near the larger rivers: Marion, Montgomery, St. Charles, St. Louis, Ste. Gene- vieve, Boone, Howard, Saline, Gentry, Holt, Platte, Jackson, and Vernon counties. Ranunculus — Nutt. Open rocky woods, and moist or alluvial open ground. Circumneutral. East- ern, bee and southern Mo.: Adair, Pike, St. Louis, Jefferson, Franklin, St. Francois, Oregon, Shannon, Crawfo rd, Phelps, Howell, Boone, Moniteau, Mont- gomery, Clay, parani Greene, Taney, Barry, Jasper, Newton, and McDonald counties Ranunculus abortivus L. Small-flowered Crowfoot. Moist alluvial or rocky woods, and open or waste ground. Cireumneutral. General Ranunculus Harveyi (Gray) Britton. ocky open woods and ledges. Oxylophile to eireumneutral. Central and Bere Mo.: Montgomery, Gasconade, Osage, St. Louis, Jefferson, Wash- ington, Franklin, Crawford, Dent, Madison, Wayne, Iron, Reynolds, Carter, Maries, Phelps, Pulaski, Texas, Camden, Dallas, Laclede, Greene, Barry, and eDonald counties. Ranunculus Harveyi f. pilosus (Benke) Palmer & Steyermark, comb. nov. Ranunculus Harveyi var. pilosus Benke, Rh. 30: 200. 192 ometimes occurring with the typical form, but scarcely distinet enough to de- serve varietal rank. Ranunculus recurvatus Poir. Hooked Crowfoot. Rich or rocky open woods. Circumneutral. General. Ranunculus fascicularis Muhl. Prairie Buttercup. eky prairies, open woods, banks, and glades. Cireumneutral to oxylophile. General. Ranunculus septentrionalis Poir. Swamp Buttercup. Rich or wet woods, and borders of sloughs and ponds. Cireumneutral. General Ranunculus caricetorum Greene. Ranunculus d Mack. € Bush. Low wet prair Cire prone Local: Jackson Co. Similar to the last and perhaps not prod disti inc Ranunculus hispidus Michx. eky open woods. Cireumneutral to oxylophile. General but commonest in ion: ion, St. Louis, Jefferson, Franklin, Bollinger, Iron, bises Ripley, Shannon, Texas, Warren, Boone, Cole, Morgan, Camden, Howell Greene, Webste er IR Stone, Barry, Platte, Jackson, Jasper, New- ton, ai Melionsla counti Ranunculus hispidus var. falsus Fernald, Rh, 22: 30. 1920, Sometimes found with the typical form RANUNCULUS REPENS L. See Rh. 21: 169. 1919. Creeping Buttercup. w wet woods and open ground. Cireumneutral. Local: St, Louis Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 541 RANUNCULUS PARVIFLORUS L. Swampy woods and open ground. Oxylophile. Southeastern Mo.: Butler and Dunklin counties. RANUNCULUS ACRIS L. Meadow Buttercup. dows and waste ground. Circumneutral. Introduced in St. Louis and Jackson countie Myosurus wa L. Mouse-tail. Low open woods and moist fields. Cireumneutral. General, Trautvetteria carolinensis (Walt.) Vail. False Bugban Moist limestone bluffs. Caleiphile. Rare and local: dl Co. Thalictrum dioicum L. Early Meadow Rue Rich rocky woods and ledges along iade bluffs. Cireumneutral to calciphile. General, but seattered: Adair, St. Louis, Jefferson, Franklin, Carter, Reynolds, Shannon, Oregon, Howell, Morgan, J ioni Benton, and Stone counties. Thalictrum dasycarpum Fisch. & Lall. Meadow Rue. Rich or rocky woods, and moist open ground. Cireumneutral, General. Thalictrum revolutum DC. Rich or roeky woods. Cireumneutral. General but less common than the last species. Anemonella thalictroides (L.) Spach. Rue Anemone. Rich or rocky woods. Circumneutral. General. Hepatica americana (DC.) Ker. See Rh. 19: 45. 1927. Hepatica, Liver-leaf. Hepatica triloba of Am. auth., not Chaix. (G). epatica Hepatica (L.) art, (B&B). Rocky hillside woods and bluffs. Oxylophile to cireumneutral. Southern Mo.; Iron, Reynolds, Dent, Shannon, Oregon, Howell, Texas, 2e Douglas, Ozark, Webster, Greene, Christian, Taney, Stone, and pni eountie Hepatica americana f. candida Fernald, Rh. 6. 1917. Commonly found throughout the range of pi typical form in Missouri. Hepatica acutiloba DC. Hepatica, Liver-leaf. Rocky hillside woods and bluffs. Caleiphile to eireumneutral. Eastern and middle Missouri: Sehuyler, Adair, Clark, Marion, Pike, Boone, Callaway, War- ren, Montgomery, St. Charles, St. Louis, Jefferson , Franklin, mich, Wash- ington, St. Francois, Ste. Genevieve, Perry, Bolinger , Cape Girardeau, Iron, Wayne, Carter, Reynolds, and Shannon countie Hepatica acutiloba f. albiflora Hoffmann, Proc. Bost. Soc. Nat. Hist. 36: 268. 1922 Commonly found throughout the range of the typical form in Mo. Hepatica acutiloba f. rosea Hoffmann, Proc. Bost. Soc. Nat. ES 36: 268. 1922. Commonly found throughout the range of the typieal form in Anemone caroliniana Walt. Prairie Anem Prairies, meadows, glades, waste dl and along roadsides. Circum- neutral. estern and middle Mo.: Boone, Pulaski, Morgan, Dallas, Johnson, Henry, St. Clair, Cedar, ¿PANA Sd Mises Cass, Bates, Vernon, Barton Greene, Jasper, Newton, and MeDonald eounti à [Vor. 22 542 ANNALS OF THE MISSOURI BOTANICAL GARDEN Anemone cylindrica Gray. Thimbleweed. Prairies and open woods. Circumneutral to calciphile. Northwestern and west-central Mo.: rerit Nodaway, Holt, and Jaekson counties, Anemone virginiana L. Thimbleweed. Prairies and open woods. Cireumneutral. General, Anemone virginiana f. leucosepala Fernald, Rh. 19: 140, 1917, In similar situations to the typical form. Found throughout the range of the species in Mo Anemone canadensis L. White Anemone. ow open woods and meadows. Circumneutral. Northern, central, and east- Mo., mainly near the larger rivers: Clark, Marion , St. Louis, Jefferso Franklin, Ste. Genevieve, Madison, Butler, Re Saline, pedo Holt, Andrew, Platte, Clay, and Jackson counties, Clematis virginiana L. Virgin's Bower. Woods and thickets along streams. Circumneutral. General. Clematis virginiana var. missouriensis (Rydb.) Palmer & Steyermark, comb. nov. tis missouriensis Rydb. (G), (B& B Roeky woods and thiekets along streams. — eutral. Central, southern, and western Mo.: Scott, Butler, Shannon, Maries, Cole, Saline, eed N Greene, Christian, Taney, Stone, Barry, J asper, and MeDonald eounti Clematis Viorna L. Leather Flower. Rich woods and bluffs. Circumneutral. Southern Mo.: Bollinger and Taney counties. Clematis versicolor Small. Rocky woods d bluffs. Cireumneutral. Southern Mo.: Oregon, Ozark, Taney, Greene, Stone, Barry, and MeDonald counties. Clematis Pitcheri T. & G. Leather Flower, Bluebell. Viorna Pitcheri (T. & G.) Britton (B & B). Thiekets and rocky woods. Cireumneutral to ealeiphile. General. Clematis crispa L. Swamp Leather Flower. Low wet woods and borders of swamps. Circumneutral. Southern Mo.: Scott, Mississippi, Pemiseot, Dunklin, Stoddard, Butler, Ripley, Ozark, and New- ton counties, Clematis Fremontii Wats. Limestone glades. Caleiphile. Eastern Mo., south of the Missouri River: St. Louis, Jefferson, Franklin, St. Franeois, and Washington counties Isopyrum biternatum (Raf.) T. & G. False Rue Anemone. Rich woods. Cireumneutral to calciphile. General. Aquilegia canadensis L. Wild Columbine. Aquilegia coccinea Small (8). Shaded rich woods, rocks and bluffs; usually on limestone. Caleiphile to cir- cumneutral. General. Aquilegia canadensis var. flaviflora (Tenney) Britton. Occasionally occurring with the species. Northwestern Mo., local: Bu- chanan Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 543 DELPHINIUM AJACIS L. Larkspur. Commonly cultivated in old gardens and often escaping. General, Delphinium tricorne Michx. Dwarf Larkspur oeky woods and thickets. Cireumneutral. General, Delphinium Treleasei Bush. Glade Larkspur Glades and rocky slopes of bald isthe. Calciphile. Southwestern Mo.: Greene, Taney, Stone, and Barry counties. Delphinium azureum Michx. Rocky open woods, glades, and prairies. Calciphile to circumneutral, Central and southern Mo.: Franklin, Crawford, Dent. Shannon, Carter, Phelps, Pulaski, Maries, Cole, Texas, Howell, Laclede, Wright, Webster, Taney, Miller, Douglas Ozark, Greene, and Barry counties. This species and Delphinium Penardi sates: grade in color considerably. Delphinium azureum var. Nortonianum (Mack. & Bush) Palmer & Steyermark, comb. nov. Delphinium Nortonianum Mack. & Bush (G). Rocky upland open woods, glades, and prairies. Oxylophile to eireumneutral. Southern Mo.: Crawford, Carter, Shannon, Oregon, Texas, Howe ougla Ozark, Stone, Taney, Wright, Barry, a and McDonald counties. The ae id upon which Delphinúwm Nortonianum were based have been found to merge with typical Delphinium azurewm in certain degrees, and it appears best treated as a variety of the latter species. Delphinium Penardi Huth. Prairie Larkspur. Delphinium camporum Greene (B & B). Rocky prairies and glades. Cireumneutral. Northern, central, and southwest- ern Mo., apparently "be from much of the Ozark region and from the South- eastern lowlands: Clark, Adair, Macon, Linn, Ralls, Lincoln, St. Louis, Boone, Saline, Miller, ud Lacie e, Dallas, Atchison, Grundy, Daviess, Caldwell, Jackson, Cass, Bates, Vernon, Henry, St. Clair, Greene, Webster, Stone, Dade, Lawrence, Barry, Jasper, Newton, and MeDonald counties Cimicifuga racemosa (L.) Nutt. Bugbane, Black Snakeroot. ocky woods and bluffs. Circumneutral to oxylophile. I central and southern Mo.: St. Louis, St. Francois, Bollinger, Madison, Dent, Iron, Wayne, Reynolds, te. rae Dies, pra , Shannon, Howell, vers Wright, Web- ster, Greene, Douglas, Ozark, Stone, Barry, and Jasper counties Actaea brachypoda Ell. White PEDE Actaea alba (L.) Mill. (G), & B). Rieh woods and banks of 2 streams. Cireumneutral. Eastern, middle, and igno dod and doni in northwestern Mo.: I. Adair, Clark, Marion, , St. Louis, Jefferson, Franklin, St. Francois, Perry, Scott, erui] as Texas, Dent, Crawford, Phelps, Pulaski, Camden Laclede, Atchison, piss and Barry counties. piece canadensis L. Golden Seal. Rie s and thickets. Cireumneutral. Central and southern Mo.: well "rade but nowhere common: St. , Jefferson, Franklin, St. eise jid Bollinger, Dunklin, MU Cain, os, ers), Phelps, Miller, Cole, e, Saline, Jackson, Johnson, Polk, Greene, Taney, Lawrence, Barry, and J aois counties. [Vor. 22 544 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fam. MAGNOLIACEAE Magnolia acuminata L. Cucumber Tre Low hills and rocky bluffs. renal Southern Mo.: Cape Girardeau, Scott, Stoddard, Dunklin, Butler, and Ozark counties. Magnolia acuminata var, ozarkensis Ashe, Jour. Elisha Mitchell Sci. Soc. 41: 269, 1926, Along rocky bluffs. Cireumneutral. Southwestern Mo.: Barry Co. Liriodendron Tulipifera L. Tulip > Yellow Poplar. Upland woods and along small ams, Ci tral. Southeastern Mo.: Perry, Cape Girardeau, Bollinger, Pen Mississippi, Dunklin, Stoddard, and Butler counties. Fam. ANNONACEAE Asimina triloba Dunal. Papaw. Rich woods and along bluffs and streams. Cireumneutral. General. Fam. MENISPERMACEAE Cocculus carolinus (L.) DC. Fishberry. Roeky woods and thiekets. Cireumneutral. Southern Mo., and loeally north- ward along the large rivers: Marion, 8, Cole, Iron, Reynolds, Wayne, Mis- sissippi, Dunklin, Ripley, Oregon, Shannon, Howell, Onak, Douglas, Taney, Stone, Barry, Greene, Cedar, Vernon, Barton, Jasper, Newton, and McDonald counties. Menispermum canadense L. Moonseed, Yellow-root. Moist woods and thickets. Cireumneutral. General. Calycocarpum Lyoni (Pursh) Nutt. Cup-seed. Thickets and low alluvial fields. f Aoc East-central and southern Mo.: St. Louis, Jefferson, Franklin, Madison, Scott, Mississippi, Stoddard, Dunklin, Ripley, Carter, Oregon, Dent, Shannon, Pulaski, — Ozark, Taney, Stone, Cedar, Barry, Jasper, Newton, and MeDonald eountie Fam. BERBERIDACEAE Podophyllum peltatum L. May Apple, Mandrake. Open woods and thickets. Cireumneutral to ealeiphile. General. Caulophyllum thalictroides (L.) Michx. Blue Cohosh. Rich woods and moist banks along bluffs. i e to m Gen- eral but uncommon: Mereer, Sehuyler, Marion, B , St. Louis, Jef- ferson, Franklin, Shannon, Texas, Howell, Phelps, piss y rim Laclede, Ozark, Greene, Clay, J Mis and Jasper counties. Berberis canadensis Mill. American Barberry. Rocky woods and bluffs, on limestone and sandstone. Oxylophile. Southeast- ern Mo., local: Shannon, Texas, and Howell counties. BERBERIS VULGARIS L. European Barberry. Cultivated pe rarely escaped into pastures and open woods. Clark and Marion Oomi ies 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 545 BERBERIS THUNBERGI DC. Japanese Barberry. Cultivated and rarely escaped into open woods and open ground. Marion Co. Fam. LAURACEAE Sassafras officinale Nees & Eberm. Sassafras. Sassafras variifolium (Salisb.) Ktze. (G). Sassafras Sassafras (L.) Karst. (B € B). Borders of woods and Pls dry or sterile soil. Oxylophile to eireumneutral. Eastern, central, and s n Mo.; south of a line drawn from Marion to Lafayette and Cass pa A Sassafras officinale var. albidum (Nutt.) Blake, Rh. 20: 99. 1918. Similar situations and range to those of the typical form, but less common. Benzoin aestivale (L.) Nees. Spice Bush. oods and thickets, in moist alluvial or rocky soil. Circumneutral. Central and southern Mo.; commonest in the Ozark region. Benzoin aestivale var. pubescens Palmer & Steyermark, var. Similar situations to the last. Cireumneutral. irl a southern Mo.: Montgomery, St. Louis, Jefferson, New Madrid, Stoddard, Iron, Shannon, Dent. Ozark, Moniteau, Morgan, Hickory, Stone, Jasper, Newton, and MeDonald counties. A typo recedit ramulis annotinis pubescentibus; foliis subtus ad nervos a Vide margine ciliatis. Diff m the typical form in the pubescent young branchlets and in the a Ber are more or less pubescent, at least along the veins beneath, eiliate on the margins, and with pubescent petioles. outh Carolina, Georgia, Alabama, Mississippi, Louisiana, Tennessee, Ken- tucky, southern Illinois, and central and southern Missouri, to southeastern Kan- as, Oklahoma, and eastern Texas. The occurrence of this undescribed pubescent uiri FR ae a large area has been the eause of eonsiderable confusion and as been responsible for the wide range given in manuals to Benzoin melissae- da (Walt.) Nees, which has been credited to Missouri, but which appears to be a rare species confined to the coastal plain and Piedmont regions of the south- eastern states. MISSOURI: Alba, Jasper Co., Sept. 12, 1922, E. J. Palmer 21923 (Arn. Arb. TYPE). Fam. PAPAVERACEAE Sanguinaria canadensis L. Bloodro oeky woods and moist rich cl along bluffs. Circumneutral to calciphile. General. Sanguinaria canadensis var. rotundifolia (Greene) Fedde. Wooded hillsides and bluffs along the Missouri River. Circumneutral to calci- phile. Northwestern Mo., local: Atchison Co. CHELIDONIUM MAJUS L. Celandin Introduced in waste ground. J ackson Co. [Vor. 22 546 ANNALS OF THE MISSOURI BOTANICAL GARDEN Stylophorum diphyllum (Miehx.) Nutt. Celandine Poppy. Rocky woods, rich ground, and bluffs. Cireumneutral. East-eentral and southern Mo., mostly in the eastern Ozark region: Warren, St. Charles, St. Louis, Jefferson, Washington, St. Francois, Madison, Stoddard, "Wayne, Carter, Shan- non, and Tamer counties. Argemone intermedia Sweet. Prickly Poppy. Open en Oxylophile to eireumneutral. Central and southern Mo., scat- tered: St. Louis, Dunklin, Butler, Jackson, and Greene counties. Introduced ex- cept perhaps in E lowlan Argemone alba Lestiboudois. Waste and open ground. Indifferent. Southeastern Mo.: Butler Co., and also introduced in Jackson Co. Argemone mexicana L. Mexican Poppy. Waste open ground and along railroads. Introduced in Knox, Phelps, Jack- son, and Jasper counties. PAPAVER DUBIUM L. Blind Eyes. Waste ground. Escaped from cultivation in Adair and Jackson counties. PAPAVER RHOEAS L. Corn Poppy. Waste ground. Escaped in Adair and Cole counties. PAPAVER SOMNIFERUM L. Common Poppy. Cultivated in gardens and escaped into waste ground: Jaekson Co., and also reeorded from Greene Co. Fam. FUMARIACEAE Dicentra Cucullaria (L.) Bernh. Dutchman’s Breeches. ich woods and moist rocky ground along bluffs. Cireumneutral to calciphile, General, Dicentra canadensis nd Walp. Squirrel Cor ieh b shaded bluffs, Rd Rare. Northern and central Mo.: zone "Warren, St. Louis, Jefferson, Boone, and Jaekson counties. Corydalis flavula (Raf.) DC. Rich woods and moist rocky banks or ledges along bluffs, Cireumneutral. General Corydalis micrantha (Engelm.) Gray. Rich oup, and along open or shaded banks and bluffs. Cireumneutral. Western Mo. Apres lin Clay, Jaekson, Henry, Webster, Wright, Stone, Jasper, N SE d counties. Corydalis id: ti Open woods, prairies, and barrens. Circumneutral. Western Mo.: Livingston, Pettis, Cass, Bates, c Henry, St. Clair, Greene, Lawrenee, Jasper, Newton, and MeDonald eoun Corydalis campestris c Buchholz & Palmer, Trans. Acad. Sci. St. Louis 25: 115. 192 Capnoides campestre Britton (B & B). Rocky woods. Circumneutral. Southwestern Mo.: MeDonald Co. 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 547 Corydalis aurea Willd. Open or rocky woods, prairies, v AE fallow fields, ledges, and roadsides. Caleiphile to cireumneutral. General: Livingston, Grundy, Daviess, Adair, Ralls, St. Louis, Jefferson, Ste. khan Mississippi, Dunklin, Wayne, Carter, Phelps, Texas, ‘Candie, Saline, Randolph, Wright, Polk, Clay, Jackson, Cass, Vernon, Greene, Jasper, and McDonald counties. Corydalis aurea var. occidentalis Engelm Corydalis montanum (Engelm.) Britton (B & Open or rocky woods, glades, and prairies. f MENOR Central and south- ern Mo.: St. Louis, Jefferson, Madison, Gasconade, Boone, Phelps, Crawford, Pulaski, Moniteau, Henry, St. Clair, Cedar, Jackson, and Jasper counties. Fam. CRUCIFERAE DRABA VERNA L. Whitlow Grass. Fields and waste ground. psc iden Eastern and middle Mo., scattered: Marion, St. Louis, Boone, and Cole counties Draba reptans (Lam.) Fernald, Rh. 36: 368. 1934. Draba caroliniana Walt. (G), (B & B). Fields, glades, rocky ledges, and waste ground. Cireumneutral to oxylophile; frequently on sandstone or cherty soils. General. Draba reptans var. micrantha (Nutt.) Fernald, Rh. 36: 368. 1934. Draba caroliniana var. micrantha (Nutt.) Gray (G). Sandy bottoms and rocky glades. Oxylophile to eireumneutral. Local: Jack- son Co. Draba cuneifolia Nutt. eky open woods, glades, and rocky ledges. Calciphile to cireumneutral, Central and southern Mo., south of a line drawn from Ralls and Boone counties to Jackson Co Draba brachycarpa Nutt. Prairies, eultivated and fallow fields, and low open ground or glades. Cir- cumneutral. Central and southern Mo. south of a line drawn from St. Charles, Audrain, and Randolph counties to Jackson Co BERTEROA INCANA (L.) DC. Hoary Alyssum. open ground. Cireumneutral Introduced in Greene Co. Lesquerella gracilis (Hook.) Wats. Slender Bladder-pod. Glades and rocky open woods. Calciphile. Southwestern Mo.: Greene, Dade, Lawrence, and Jasper counties, and also introduced in Bollinger Co. knee angustifolia (Nutt.) Wats. s and barrens. Caleiphile. Southwestern Mo., local: Greene Co. THLASPI ARVENSE L. ny Cress aste ground, and along roadsided and railroads. Indifferent. Northern and eentral Mo., and perhaps more general: Macon, St. Charles, St. Louis, Jeffer- son, Franklin, Boone, Cooper, Chariton, Carroll, Gentry, Daviess, Atehison, Noda- way, and Jackson counties. THLASPI PERFOLIATUM L. Along roeky stream bed. Introduced in Jefferson Co. [Vor. 22 548 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lepidium virginicum L. Tonguegrass, Peppergrass. Lepidium texanum Buckl. (R) Fields, roadsides, and waste ground. Indifferent. General and common. LEPIDIUM DENSIFLORUM Sehrad. L m neglectum Thell. Lepidium apetalum of auth., not Willd. (G). Fields, waste ground, and about dwellings. Indifferent. General but scattered. LEPIDIUM PERFOLIATUM L. Waste ground and along railroads and roadsides. Indifferent. Scattered: St. Louis, Madison, and Jackson counties. LEPIDIUM CAMPESTRE (L.) R. Br. Waste ground and along railroads. Indifferent. Northern, central, and eastern Mo., and occasional elsewhere: Adair, Knox, Ralls, Pike, Lincoln, St. Charles, St. Louis, Franklin, Madison, Iron, Laclede, Livingston, Carroll, Daviess, and Jackson counties. LEPIDIUM DRABA L. Waste ground. Circumneutral. Introduced in St. Louis Co. CoRONOPUS DIDYMUS (L.) Sm. aste ground. Indifferent. Southeastern Mo.: Dunklin and Butler counties. CAPSELLA BURSA-PASTORIS (L.) Medie. Shepherd’s Purse. ields, roadsides, waste ground, and about dwellings. Indifferent. General common CAMELINA SATIVA (L.) Crantz. False Flax aste ground and along railroads. Indifferent. Seattered: St. Louis and Jackson counties CAMELINA MICROCARPA Andrz. imilar situations to the e preceding and more common. Indifferent. General but scattered: St. "itid: Franklin, Iron, Cole, Clay, Jackson, Law- renee, and Jasper nn RAPHANUS SATIVUS L. Radish. Introduced in fields and waste ground. Indifferent. Scattered: Boone and J ae counties ERUCA SATIVA Mill. Garden Rocket. Introduced along railroads. Indifferent. Scattered: Clark and Jackson countie ERUCASTRUM POLLICHII Spenn aste ground. Indifferent. Introduced in Jackson Co. BRASSICA ALBA (L.) Boiss. White Mustard. Fields and waste ground. i ean General. BRASSICA ARVENSIS (L.) Ktze. Charlock. Fields, roadsides, and waste ground. Indifferent. General. BRASSICA JUNCEA (L.) Cosson. Indian Mustard. elds and waste ground. Indifferent. General. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 549 BRASSICA JAPONICA Siebold. Curled Mustard. Fields and waste ground. Indifferent. General but scattered: Pike, St. Louis, Boone, Jackson, and Jasper eounties. BRASSICA NIGRA (L.) Koch. Black Mustard. ields, waste ground, and roadsides. Indifferent. General. BRASSICA CAMPESTRIS L. Rutaba Open and waste ground. ea General. CONRINGIA ORIENTALIS (L.) Dumort. Hare’s-ear Mustard. Waste Aires and along roadsides and railroads. Indifferent. Scattered: St. viec St. Franeois, Iron, Randolph, Boone, Daviess, Clay, and J Moni Bod SISYMBRIUM OFFICINALE (L.) Seop. Hedge Mus Fields, waste ground, and along railroads. Cop General. SISYMBRIUM OFFICINALE var. LEIOCARPUM DC. Frequently found with the typieal form. SISYMBRIUM ALTISSIMUM L. Tumble Mustard. ields, waste ground, and along railroads. Indifferent. General. DESCURAINIA SoPHIA (L.) Wetts. Sisymbrium Sophia L. (G). Sophia Sophia (L.) Britton (B & B). Prairie banks and waste ground. Cireumneutral. Western Mo.: Jackson and Nodaway counties. Descurainia intermedia (Rydb.) Daniels. Tansy Mustard. Sisymbrium canescens Nutt. var. brachycarpon Bn ) Wats. (G). Sophia brachycarpa (Richards. ; Rydb. (R). Sophia pinnata (Walt.) Howell (B & B), in part. Glades, open banks, and prairies, roadsides, and dry rocky ledges along bluffs. Caleiphile to eireumneutral. General. is and the previous species have some- times been confused with Descurainia incisa (Sisymbrium incisum) and Des- curainia canescens (Sisymbrium canescens) which are not known to occur in Mo. ARABIDOPSIS THALIANA (L.) Heynh. Mouse-ear Cress. Sisymbri ) Rocky or alluvial open ground and fields. Circumneutral. East-central and southern Mo., seattered: Callaway, St. Louis, Ste. Genevieve, Cape Girardeau, Stoddard, Pemiseot, and Ozark counties. ERYSIMUM REPANDUM L Fields, waste ground, and along railroads. Indifferent. Central Mo., seattered: Jefferson, Franklin, Cole, Montgomery, Boone, and Jackson eounties. Erysimum asperum DC. Western Wall-flower. Erysimum arkansanum Glades and along EN bluffs. Calciphile. Central and southern Mo.: St. Charles, St. Louis, Jefferson, Franklin, Osage, Maries, Miller, Camden, Ben- ton, Cooper, Jackson, and Taney countie Erysimum cheiranthoides L. Worm-seed uim Fields and waste ground. Cireumneutral. Seattered: Clark, St. Louis, Pulaski, and Jackson counties. [Vor. 22 550 ANNALS OF THE MISSOURI BOTANICAL GARDEN Roripa Nasturtium-aquaticum (L.) Schinz & Thell. Fl. Schweiz, ed. 3, 240. 1909. Water Cress. Radicula Nasturtium-aquaticwm (L.) Britten & Rendle (G). Sisymbrium Nasturtium-aquaticum L. (B € B). In springs and clear running water. Caleiphile to eireumneutral. Abundant in central and southern Mo., and perhaps found throughout the state. Roripa sinuata (Nutt.) Hitche. Radicula sinuata (Nutt.) Greene (G), (B& B ow open ground and alluvial banks of streams. Cireumneutral. General but seattered: Marion, Ralls, St. Louis, Jefferson, Perry, Pemiscot, Camden, Boone, Cooper, Daviess, Atchison, Clay, and Jaekson eounties. Roripa sessiliflora (Nutt.) Hitche. Radieula sessiliflora (Nutt.) Greene (G), (B&B). Wet open woods and borders of sloughs, ebd and streams, Circumneutral. General. RORIPA SYLVESTRIS (L.) Bess. Yellow Cress, Radicula sylvestris (L.) Druee (G), (B & B). t meadows and banks of streams, Circumneutral. Scattered: St. Louis, Boone, Stone, and McDonald counties. Roripa obtusa (Nutt.) Britt Radicula obtusa (Nutt) es (G), (B& Wet woods and borders of sloughs, diri and ditches. Cireumneutral. General. Roripa 7 (Desv.) Britton. € Rh. 30: 131. 1928. Marsh Water Cress. di palustris var. hispida (Desv.) Robinson ( pecie flats along the MS River. Circumneutral, Local: Perry Co. Roripa hispida var. glabrata Lunell. See Rh. 30: 133, 1928. Marsh Water Cress. Radicula palustris of Am. auth., not Moench (G), (B & B). Wet woods, borders of sloughs a and ponds, and alluvial soils of streams and rivers. Cireumneutral. Genera Roripa aquatica (Eat.) Palmer & NU comb. nov. Lake Cress. Sloughs and swampy woo Cire eutral. Southern Mo., scattered: St. Louis, Jefferson, Ripley, Phelps, and Tan asper countie RoRiPA ARMORACIA (L.) Hitche. Horseradish. Radicula Armoracia (L. ) Robinson (G). Armoracia Armoracia (L.) Britton (B & B Cultivated and occasionally escaped into fields and waste ground. Indifferent. Scattered: Boone, Jackson, and Taney counties. Barbarea vulgaris R. Br. Yellow Rocket, Winter Cress Wet fields, moist meadows, and alluvial ground. — n" General. Barbarea vulgaris var. longisiliquosa Carion. See Rh. 11: 139. 1909 Barbarea stricta of auth. in part, not Andrz. (G), (B € B). 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 551 Moist meadows and alluvial ground. Cireumneutral. Western Mo., local: Jackson Co. This has sometimes been confused with the preceding variety. Selenia aurea Nutt. Rocky prairies, chert, sandstone and leached limestone glades, sandy fields, and roadsides. Oxylophile. Southwestern Mo.: Benton, Polk, Webster, Greene, St. Clair, Cedar, Lawrence, Vernon, Jasper, and Newton counties. Iodanthus pinnatifidus (Michx.) Steud. Purple Rocket. ods and moist alluvial thickets. Circumneutral. General, Leavenworthia uniflora (Michx.)Britton. Wet rocky ledges and depressions in glades. beiden ile. East-central and southern Mo.: St. Louis, Franklin, Jefferson, St. Francois, Washington, Craw- ord, Madison, Shannon, Maries, Gaseonade, a, Phelps, Pulaski, Laelede, icd , Camden, Dallas, Polk, Wright, Webster, Douglas, Taney, Stone, and Barry coan iien Dentaria laciniata Muhl. Toothwort, Pepper-root. Rich or rocky woods. Cireumneutral. General. Dentaria laciniata var. integra (Schulz) Fernald. imilar situations to the typical form. Circumneutral. Scattered throughout the range of the typical form. Cardamine bulbosa (Schreb.) BSP. Spring Cress. Wet woods and meadows, and along wet sandstone cliffs. Circumneutral to oxylophile. Eastern, central, and southern Mo., south of a line drawn from ouis, Montgomery, Audrain, and Randolph, to Camden and Jackson counties Cardamine pennsylvanica Muhl. Wet woods, bogs, and springy ground. Circumneutral. General. Cardamine parviflora L. var. arenicola (Britton) O. E. Schulz. See Rh. 29: 192. 927. Cardamine parviflora of Am. auth., not L. (G), (B & B), in part. Cardamine arenicola Britton (B & B), in part. Wet open woods, fields, wet rocky ledges, and depressions in glades. Cireum- neutral to oxylophile. General. inae N Britton. Spring Cress. al and moist woods. Cireumneutral. Eastern and northern Mo.: Adair, pe Shelby, cud) dud Cape Girardeau counties. Arabis lyrata L. Cress. Rocky bluffs or ledges, on limestone or sandstone. Caleiphile to eireumneutral. Eastern Mo., south of the Missouri River: St. Louis, Jefferson, Washington, Crawford, Shannon, and Texas counties. Arabis dentata T. & G. Rieh woods, moist rocky ledges, and borders of streams. Cireumneutral. Cen oone, War ton, St. Francois, Ste. Genevieve, parda toddard, Shannon, Camden, Texas, Laclede, Morgan, Polk, Saline, Jackson, Benton, St. Clair, Wright, Douglas, Taney, Stone, and Jasper counties. [Vor. 22 552 ANNALS OF THE MISSOURI BOTANICAL GARDEN Arabis virginica (L.) Poir. See Rh. 29: 192. 1927. Open alluvial ground, fallow fields, and waste ground, Cireumneutral to oxy- lophile. Central and southern Mo., south of a line drawn from Ralls and Chari- ton eounties to Jackson Co, Arabis hirsuta (L.) Scop. ocky ledges and bluffs, usually on limestone. Caleiphile to cireumneutral. Eastern, central, and southern Mo., but apparently absent from much of the Ozark region: Clark, Lewis, Marion, Ralls, Boone, St. Louis, Jefferson, Frank- lin, Washington, Cape Girardeau, Carter, Shannon, Oregon, Webster, Christian, Taney, Stone, Jackson, Jasper, Newton, and McDonald counties. Arabis viridis Harger, Rh. 13: 38. 1911. Dry rocky woods, rocky open ground and bluffs. Oxylophile. Southern and east-central Mo.: Ste. Genevieve, Madison, Wayne, Stoddard, Ripley, Carter, Iron, Washington, Franklin, Crawford, Dent, Reynolds, Ore egon, Shannon, Texas, Phelps, Pulaski, Douglas, Stone, Ozark, Jasper, Newton, and MeDonald impen es. This species has previously been confused with Arabis Drummondii, A. patens, A. brachycarpa, and A. glabra, none of which occur in Missouri, and has also been confused with A. laevigata. Arabis laevigata (Muhl.) Poir. Rich woods, along base of bluffs, alluvial ground along streams, and along bluffs and ledges. Circumneutral. Eastern, central, and Southern Mo., east and south of a line drawn from Marion, Shelby, Boone, and Henry iude to Ver- non Co., and locally northwest in Platte Co. Arabis canadensis L. Sickle-pod Rocky woods and bluffs. Circumneutral. General. Fam. CAPPARIDACEAE Polanisia graveolens Raf. Clammy-weed. Rocky open ground, waste ground, and gravel bars along streams. Circum- neutral to oxylophile. General but not common. Polanisia trachysperma T. & G. In similar situations to the preceding. Cireumneutral. Southwestern Mo.: Greene, Taney, Stone, Barry, and McDonald counties. Cleome serrulata Pursh. Rocky Mountain Bee-plant. Loess hills and waste ground. Cireumneutral Northwestern Mo., and intro- duced in central Mo.: Atchison, Jackson, and St. Louis counties. CLEOME SPINOSA L. Spider-flower. Cultivated in gardens and sometimes escaped into waste ground. Indifferent. Seattered: St. Louis, Pemiscot, Dunklin, Taney, Stone, Barry, and J asper counties. Fam. CrassuLACEAE Penthorum sedoides L. Ditch Stoneerop. ow wet woods, swamps, and along sloughs and ditches. Circumneutral. General, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 553 Sedum Nuttallianum Raf. Yellow-flowered Stonecrop. Chert glades. Oxylophile to circumneutral. Southwestern Mo., local: Jasper and Newton counties. Sedum pulchellum Michx, Widow’s Cross. Glades, rocky ledges, and bluffs. Calciphile to cireumneutral; most frequently on limestone, but also found on granitic and siliceous dis Central and south- ern Mo., but commonest in the western Ozark region: Moniteau, Boone, Lincoln, Oregon, s Dallas, Polk, ppe Wright, Webster, ibus S Stone, Ja P son, dye enry, St. Clair Bes r, awrence, Barry, Jasper, Newton, an ald counties. This pilis ira with the eastern Sedum Nevii im is not known in E Sedum ternatum Michx. Rocky ledges and ary Calciphile to cireumneutral. Scattered: Marion, Greene, and Barry countie SEDUM TRIPHYLLUM uis S. F. Gray. See Rh. 11: 46. 1909. Live-forever, Orpine. Sedum purpureum Tausch (G). anted in old gardens and escaped to roadsides and waste ground. Indif- ferent. Scattered: Clark, Scotland, Marion, St. Louis, Boone, and Jasper counties. Fam. SAXIFRAGACEAE pedum renifolia Rosendahl, Univ. Minn. Studies Biol. Sci, 6: 410, pl. 43. 927. Sullivantia Sullivantii of auth., not (T. & G.) Britton (G), (B & B). On moist shaded bluffs of St. Peter sandstone. Oxylophile. East-central Mo., loeal: Warren and Jefferson counties. Saxifraga pensylvanica L. var. Forbesii (Vasey) Engl. & Irmsch. Pflanzenreich 4, Fam. 117, 1: 66. 1916. Saxifraga Forbesti Vasey (G). Micranthes pennsylvanica (L.) Haw., in part (B & B). Moist shaded sandstone bluffs. Oxylophile. East-central Mo.: Montgomery, Warren, Lincoln, St. Charles, St. Louis, and Jefferson counties. Saxifraga virginiensis Michx. Early Saxifrage. Micranthes virginiensis (Michx.) Small (B & B). Moist ledges along bluffs; on sandstone or granitic rocks. Oxylophile. East- central Mo., and southern io . in the eastern Ozark region: St. Louis, Jefferson, Ste. Sab, Madison, Iron, Phelps, Texas, and De pues owe Saxifraga texana Buckley. Micranthes texana (Buckl.) Small. eky cm and er ‘a a Southwestern Mo.: Polk, Greene, Heuchera dins L. Alum Boot. Ro wd ^" woods. Oxylophile to circumneutral. Southern and east-central rson, Washington, Madison, Iron, Dunklin, Butler, Ozark, Jasper, and J ur er [Vor. 22 554 ANNALS OF THE MISSOURI BOTANICAL GARDEN Heuchera hirsuticaulis (Wheelock) Rydb. Roeky bluffs and banks. Cireumneutral. Southern Mo., mostly in the eastern Ozark region: Jefferson, Washington, Cape Girardeau, Dak Madison, Reynolds, Carter, Ded Shannon, Dent, Pulaski, Camden, and Taney countie Heuchera Richardsonii R. Br. var. Passa Rosendahl, Butters & Bo Rh. 35: 117. 1933. Heuchera hispida of auth., not Pursh (G), (B & B). Open rocky woods, prairies, and along bluffs and embankments. Circum- neutral, de eral. This is the common species in Mo., and it has frequently been confused with H. hirsuticaulis Heuchera puberula Mack. € Bush. Heuchera parviflora of auth., not Bartl. (G), (B & B). Shaded limestone bluffs. Caleiphile. Southern Mo.: Ste. Genevieve, Ripley, Carter, Shannon, Oregon, Texas, Howell, Ozark, and Douglas counties, Heuchera villosa Michx. Shaded limestone bluffs. Calciphile. Southeastern Mo.: Iron Co. Heuchera macrorhiza Small. Shaded limestone bluffs. Calciphile. Southeastern Mo.: Iron Co. Mitella diphylla L. Miterwort. Moist shaded bluffs; on limestone or sandstone. Caleiphile to eireumneutral. East-central and southeastern Mo.: Montgomery, Warren, St. Charles, Jeffer- son, Perry, ys Oregon, Howell, and Texas counties. Parnassia grandifolia DC. Grass of Parnassus. Moist ee bluffs and moist rocky or boggy ground along streams. Cir- eumneutral to ealeiphile. Southern Mo., rare and seattered: Washington, Shan- non, Texas, Douglas, and Ozark counties. Hydrangea arborescens L. Wild Hydrangea. Shaded rocky banks and bluffs. Circumneutral; often on limestone but also found on siliceous and granitie rocks. Central and southern Mo. Common in the Ozark regio Hydrangea arborescens f. grandiflora Rehder, Rarely found with the typieal form. Stoddard Co. Hydrangea arborescens var. Deamii St. John, Rh. 23: 208. 1922. ar situations to the typical form. Scattered in southern Mo.: Bollinger, Shannon, Phelps, Pulaski, Dallas, Hickory, and Newton counties. Hydrangea arborescens var. Deamii f. acarpa St. John, Rh. 23: 208. 1902. Hydrangea cinerea f. sterilis Rehder. Rarely found with the typieal form. Shannon Co. Hydrangea arborescens var. oblonga T. & In similar situations to the typical form. East-eentral and southern Mo.: St. Louis, Cape Girardeau, Reynolds, Shannon, Ozark, and Barry counties. Hydrangea arborescens var. oblonga f. sterilis St. John, Rh. 23: 208. 1922, Rarely found with the typical form. Ozark Co Itea virginica L. Virginia Willow. Swamps, borders of bayous, and low wet woods. Circumneutral. Southeast- e 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 555 ern Mo., in the lowlands: Cape Girardeau, Bollinger, New Madrid, Dunklin, and Butler counties Ribes Cynosbati L. Prickly Gooseberry. Ro woods and bluffs. endi Eastern and southern Mo.: Clar Lewis, Gasconade, St. Louis, Jefferson, Perry, Cape Girardeau, Scott, E Oregon, Shannon, Dent, Texas, Howell, Polk, and Stone counties. Ribes PONS ense Nutt. Wild Gooseberry. Ribes gracile Michx. (G). istos missouriensis (Nutt.) Coville & Britton (B & B). Rocky open woods and bluffs. Cireumneutral to calciphile. General. Ribes odoratum Wendl. See Rh. 11: 47. 1909. Flowering Currant. Ribes aureum of auth., not Pursh (G). Along high rocky bluffs of White River and its tributaries. Caleiphile. South- western Mo.: Taney, Stone, and Barry counties Fam. HAMAMELIDACEAE Hamamelis virginiana L. Witch-hazel. Rocky woods and low ground along streams. Oxylophile to circumneutral. Southeastern Mo., in the Iron Mountain sub-region: Iron, Shannon, and Rey- nolds counties Hamamelis vernalis Sarg. Trees € Shrubs 2: 137, pl. 156. 1911. Ozark Witch- hazel. Margins and pie bars of des streams. Cireumneutral. Southern Mo.: Franklin, Washington, St. Francois, Ste. Genevieve, Bollinger, Stoddard, Madi- son, Iron, Reynolds, eut "esc Ripley, Oregon, Shannon, Texas, Phelps, Pulaski, Dallas, Douglas, Ozark, Christian, Taney, Stone, and Barry counties. Hamamelis vernalis f. tomentella Rehder, Jour. Arnold Arb. 1: 256. 1920. Sometimes found with the typical form. Taney and Barry counties. Hamamelis vernalis f. carnea Rehder, Jour. Arnold Arb. 9: 30. 1928. Oeeasionally found with the typical form. Iron and Madison counties. E cups Styraciflua L. Sweet Gum. woods and along streams. Oxylophile to eireumneutral Southeastern Mon in the lowlands: Perry, Cape Girardeau, Bollinger, Madison, Scott, Missis- sippi, New Madrid, Pemiscot, Dunklin, Stoddard, Butler, Wayne, Ripley, and Oregon counties. Fam. PLATANACEAE Platanus occidentalis L. Sycamore, Woods along streams. Circumneutral. General and probably in every county. Platanus occidentalis var. glabrata (Fernald) Sarg. Bot. Gaz. 67: 230. 1919. Occasionally found with the typical form. Madison and Jasper counties. Platanus occidentalis f. attenuata Sarg. Bot. Gaz. 67: 229. 1919. Occasionally found with the typical form. Shannon and Jasper counties. [Vor. 22 556 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fam. ROSACEAE Physocarpus opulifolius (L.) Maxim var. intermedius (Rydb) Robinson. Nine- Thickets, and rocky banks and bluffs of streams. migrado Eastern, central, and southern Mo.; commonest in the Ozark regio Spiraea alba Du Roi. Meadow-sweet. Spiraea salicifolia of auth. in part, not L. (G). Moist meadows and open ground along streams. Oxylophile to eireumneutral. Rare. Northern Mo.: Grundy Co., and also reported from Harrison Co. Spiraea tomentosa L. var. rosea (Raf.) Fernald, Rh. 14: 190. 1912. Hardhack. Rocky and sandy open ground. Oxylophile. Southeastern Mo., local: Dunk- lin Co, Aruncus sylvester Kostel. Goat’s Beard. Rocky or moist woods, and along base of bluffs. Circumneutral. Eastern, central, and southern Mo., south and east of a line drawn from Lewis, Mont- gomery, Saline, Benton, and Greene counties to McDonald Co, Gillenia stipulata (Muhl.) Trelease. American Ipecac. Rocky woods. Circumneutral to oxylophile. Eastern, central, and southern Mo.; eommonest in the Ozark region. PYRUS COMMUNIS L. Pear. Planted and sometimes escaped in thickets. Seattered: St. Louis, Boone, and Jasper counties. MALUS PUMILA Mill. Apple. Pyrus Malus L. (G). Malus Malus (L.) Britton (B & B). Cultivated and eseaped, in thickets, neri and waste ground. Scattered: St. Louis, Boone, Jackson, and Jasper counti Malus ioensis (Wood) Bailey. Wild Crab- aly Pyrus ioensis (Wood) Britton (G). Thiekets along small streams, pastures, and borders of woods. Cireumneutral to ealeiphile. General, but eommonest in northern Mo. Malus ioensis var. Palmeri Rehder in Sarg. Trees & Shrubs 2: 142, Thiekets and woods along small streams. Cireumneutral. aw p com- monest in the Ozark region. Malus ioensis var. Bushii Rehder in Sarg. Trees € Shrubs 2: 232. 1913. Open woods. Circumneutral. Southeastern Mo.: Shannon, Wayne, and Dunk- lin eounties. Malus ioensis var. spinosa Rehder in Sarg. Trees & Shrubs 2: 231. 1913. Open woods and thickets. Cireumneutral Southeastern Mo.: dl and Oregon counties Malus lancifolia Rehder in Sarg. Trees & Shrubs 2: 141. 1911. Open woods and thickets. Cireumneutral Central and southern Mo.: St. Francois, Iron, Bollinger, Wayne, Boone, and Jackson counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 557 Malus coronaria (L.) Mill. American Crab. rus coronaria L. (G) Thickets along streams. Circumneutral to calciphile. Scattered: Madison, Butler, and Jackson counties. Malus bracteata Rehder in Sarg. Trees & Shrubs 2: 230. 1913. Open woods and thiekets. Cireumneutral Southeastern Mo.: Dunklin Co. x Malus Soulardi (Bailey) Britton. oulardi Bailey (G). A er between Malus pumila and M. ioensis. Occasionally found in the range of the latter. Amelanchier canadensis (L.) Medie. See Rh. 14: 150. 1912. Juneberry, Shad Bush, Service Berry. Am AUF canadensis var. Botryapium (L.f.) T. & G. (G). Bocky woods and along bluffs. Oxylophile to circumneutral, General, but apparently absent from some of the northern counties Amelanchier humilis Wiegand, Rh. 14: 126, 141. 1912. Open ground and prairie banks along railroad. Western Mo.: Pettis and Law- renee eounties. Probably introduced. Crataegus crus-galli L. Cockspur Thorn. Crataegus albanthera Sarg. Crataegus efferta Sarg. Crataegus effulgens Sarg. Crataegus po Sarg. Crataegus severa Sarg. rataegus a eR Sarg. Crataegus tardiflora Sarg. Thickets, pastures, and borders of woods, rocky or alluvial ground. Cireum- neutral to calciphile. General and commo ds: crus-galli f. truncata (Sarg.) Pues. comb. nov. Crataegus truncata igit found with the typical form. Southern Mo.: Taney Co. Crataegus crus-galli var. exigua (Sarg.) Eggleston. 8- Occasionally found with the typieal form. Shannon Co. Crataegus crus-galli var. macra (Beadle) Palmer, eomb. nov. rataegus macra Beadle Crataegus consueta Sarg. Crataegus monosperma. Sarg. Crataegus permera Sarg. Found in similar situations to the typical form. Calciphile to cireumneutral. [Vor. 22 558 ANNALS OF THE MISSOURI BOTANICAL GARDEN Central and southern Mo.: Boone, Butler, Shannon, Ozark, Taney, and Greene counties. Crataegus crus-galli var, barrettiana (Sarg.) Palmer, comb. nov. Crataegus barrettiana Sar Found in similar situations to the typical form. Eastern Mo.: St. Louis Co. Crataegus crus-galli var. leptophylla (Sarg.) Palmer, comb. nov. Crataegus leptophylla Sar Found in similar iiuiticts to the typical form, Scattered: Osage, Madison, and Wayne counties. Crataegus crus-galli var. oe (Sarg.) Palmer, comb. nov. Crataegus pachyphylla Sar Thickets and open wonder along streams. Caleiphile to circumneutral. Scattered: Osage and St. Francois counties. Crataegus crus-galli var. bellica (Sarg.) Palmer, comb. nov. arg. Thiekets and borders of woods; ne hills and alluvial ground. Cireum- neutral to calciphile. Donibane Mo.: 0, Crataegus lawrencensis Sarg. Thickets along streams. Cireumneutral to caleiphile, Southwestern Mo.: Lawrence, Jasper, and Newton counties, Crataegus rotunda Sarg. Prairies and thickets along small streams. Cireumneutral to ealeiphile. Southwestern Mo.: Jasper, Newton, and Barry eountie Crataegus discolor Sarg. arg. Thiekets, pastures, and borders of woods, along small streams. Caleiphile to eireumneutral. Southern Mo.: Butler, Carter, Texas, Douglas, Dallas, Barry, and Jasper counties. Crataegus acutifolia Sarg. ataegus erecta Sarg. Crataegus ludoviciensis Sarg. en woods and borders of woods along streams. Cireumneutral to calciphile. Northern and east-central Mo.: Clark, Daviess, Pike, St. Louis, and Osage counties. Crataegus Palmeri Sarg. Thickets, pastures, and open woods; on limestone = or along small streams. Caleiphile to cireumneutral, v Mo.: Vernon, Lawrence, Jasper, Newton, and MeDonald counti Crataegus a: Sarg. hickets, pastures, and limestone hills, Calciphile to circumneutral. Eastern Mo., north pe the Missouri River: Marion, Ralls, and Pike counties, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 559 Crataegus regalis Beadle. Rocky open woods and thickets. Cireumneutral to calciphile. Southern Mo.: Oregon Co. Crataegus regalis var. paradoxa (Sarg.) Palmer, comb. nov. Crataegus paradoza Sarg. Prairies and thickets along small streams. Circumneutral to calciphile. Southwestern Mo.: Jasper and Newton counties. Crataegus pyracanthoides Beadle var. arborea (Beadle) Palmer, comb. nov. Crataegus arborea Beadle. Crataegus tenuispina Sarg. Open woods and banks of small streams. -— s to b on e South- eastern Mo.: Dunklin, Wayne, Carter, and Shannon countie T A eqs Rr Jour. Arnold Arb. 16: 353. 1935. , borders of woods, and along bluffs. Caleiphile to eireumneutral. NE nd eastern Mo.: Schuyler, Clark, Marion, Ralls, Pike, Monroe, St. Francois, Mercer, Harrison, and Clinton counties. Crataegus Danielsi Palmer, Jour. Arnold Arb. 16: 355. Thickets, limestone hills, and bluffs. Mo.: Boone Co. 1935. Calciphile to circumneutral. Central Ons Crataegus Danielsi f. glabra Palmer, Jour. Arnold Arb, 16: 357. 1935. Rarely found with the typical form. Central Mo.: Boone Co. Crataegus vallicola Sarg. Crataegus phaneroneura Sarg. Thickets and borders of woods, limestone hills, and along bluffs. Calciphile to eireumneutral. Eastern and middle Mo.: Maco ent counties n, Pike, Lincoln, St. Louis, and Crataegus Engelmannii Sarg. Crataegus barbata Sarg. Crataegus hirtella Sarg. Crataegus munita Sarg. Crataegus pilifera Sarg. Crataegus setosa Sarg. Crataegus villiflora Sarg. Thickets and open woods, glades, and limestone hills. Calciphile to aid neutral. Central and southern Mo.: St. Louis, Franklin, S t. Francois, Ste. Gen vieve, Iron, Wayne, Butler, Dunklin, Carter, Texas, Dent, Phelps, -— Cole, ind Polk, pue Christian, Taney, Stone, Barry, Jasper, Newto Donald e Eod 2 el f. nuda Palmer, f. nov. Crataegus infesta Sarg. Cr da => Sarg. Oeeasionally found with the typical form. Scattered: St. Louis, Franklin, and Jasper an A typo differt foliis inflorescentiis ramulisque novellis glabris. Crataegus fecunda Sarg. Thiekets and open ground along streams. Caleiphile to cireumneutral. Cen- [VoL. 22 560 ANNALS OF THE MISSOURI BOTANICAL GARDEN tral and southeastern Mo.: Boone, St. Louis, Ste. Genevieve, Cape Girardeau, and Wayne counties. Crataegus collina Chapm. Crataegus angustata Sarg. Crataegus sordida var. p Sarg. Crataegus vicina Sar Thickets and open ob on limestone hills or along small streams. Calei- phile to eireumneutral. Central and southern Mo.: Boone, St. Louis, Franklin, Washington, St. Francois, Ste. Genevieve, Cape Girardeau, Iron, Wayne, Butler, Ripley, Carter, Oregon, Shannon, Texas, Phelps, Maries, Osage, Benton, Morgan, Greene, Taney, Stone, Barry, Jasper, Newton, and MeDonald counties. Crataegus collina var. succincta (Sarg.) Palmer, comb, nov. Crataegus succincta Sarg. Thickets, limestone hills, and glades. Caleiphile to circumneutral. Eastern Mo., south of the Missouri River: St. Louis, Franklin, St. Francois, and Wayne counties. Crataegus collina var. secta (Sarg.) Palmer, comb. nov. Crataegus secta Sar Thiekets, limestone hills, and along small streams. Central and southern Mo., scattered: Miller, Greene, and Jasper counties. Crataegus collina var. sordida (Sarg.) Palmer, comb. nov. Crataegus sordida Sarg. Thickets, limestone hills. — to eireumneutral. Southern Mo.: Ripley, Carter, Shannon, and Greene counties Crataegus verruculos Thickets and oh of woods. Caleiphile to eireumneutral. Central and southern Mo., scattered: Boone, Cape Girardeau, Bollinger, Shannon, and Greene counties. Crataegus hirtiflora Sarg. Thiekets and borders of woods. Calciphile to eireumneutral. Southern Mo., loeal: Taney and Stone counties. Crataegus latebrosa Sar Thiekets and open woods along rocky streams. Caleiphile to eireumneutral. Southwestern Mo., local: MeDonald county. — Lettermanii Sarg. Open woods and thickets along streams. Cireumneutral to caleiphile. Cen- tral ind southern Mo., seattered: St. Louis, Ste. Genevieve, and Jasper counties. Probably a I idt Crataegus collina and C. mollis. Crataegus viridis Green Haw. Crataegus eb El. Crataegus D ii Sarg Crataegus furcata Sarg. Crataegus larga Sarg. Crataegus Pechiana Sarg. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 561 Low wet woods, and limestone hills and bluffs with seepage water. Calciphile to eireumneutral. Eastern, central, and southern Mo.: Marion, Ralls, Pike, St. Louis, Gasconade, Osage, Cole, Mississippi, Pemiscot, Dunklin, Stoddard, Butler, Ripley, Howell, Jasper, and Newton counties. Crataegus viridis var. ovata (Sarg.) Palmer, comb. nov. Crataegus ovata Sarg. Crataegus nitens Sarg. Occasionally found with the typical form. Marion, Ralls, St. Louis, Stoddard, and Jasper eounties. Crataegus viridis var. lanceolata (Sarg.) Palmer, comb. nov. Crataegus lanceolata Sarg. Occasionally found with the typical form. Marion, Ralls, St. Louis, Dunklin, and Jasper counties, Crataegus viridis var. . Iutensis (Sarg.) Palmer, comb. nov. Crataegus lutensis Sarg. woods and Bee glades. Circumneutral. Southwestern Mo.: Bates, Vernon, St. Clair, Jasper, and Newton counties. Crataegus viridis var. atrorubens (Ashe) Palmer, comb. nov. Crataegus atrorubens Ashe. Crataegus Dawsoniana Sarg. Low wet or alluvial woods. East-central Mo.: St. Louis Co. Crataegus nitida (Engelm.) S iekets and borders of mc limestone hills and bluffs along the Mississippi River. Caleiphile. Northeastern Mo.: Marion and Ralls counties. Crataegus padifolia Sarg. Glades and rocky hillsides. Cireumneutral to calciphile. Southwestern Mo.: Ozark, Taney, and Stone counties. Crataegus padifolia var. incarnata Sarg. ocky open woods. Cireumneutral to caleiphile, Southwestern Mo.: Stone Co. Crataegus neobushii Sarg. Crataegus leioclada Sarg. Glades, thickets, and rocky open woods. Cireumneutral to caleiphile. South- ern Mo.: Iron, Ripley, Shannon, Dallas, Taney, and Stone counties. Glades and rocky hillsides. Circumneutral to — Southern Mo.: Shannon, Taney, Stone, and Barry counties. Crataegus Margaretta Ashe Thickets, borders of woods, and rocky open ground. Circumneutral to cal- ciphile. Eastern and southern Mo.: Putnam, Schuyler, Adair, Scotland, Clark, wis, Knox, Shelby, Marion, Ralls, Pike, Lineoln, St. Louis, St. Francois, Madi- son, Dallas, Polk, Cedar, Greene, and Christian counties. Crataegus cpm Sarg. . Crataegus glabrifolia Sarg. Thickets and rocky open woods. Circumneutral to calciphile. Eastern and [Vor. 22 562 ANNALS OF THE MISSOURI BOTANICAL GARDEN middle Mo., south of the Papa E Franklin, Washington, Bollinger, Wayne, Carter, Cole, and Laclede e Crataegus pruinosa nn. 4 K. Koch. Crataegus conjuncta S Crataegus depressa pal si Presl, in part. Crataegus patrum Sarg. Open woods and thickets, limestone hills and along small streams. Circum- neutral. General, but commonest in northern and eastern Mo. Crataegus pruinosa f. angulata (Sarg.) Palmer, comb. nov. Crataegus arcana of Sarg., in part, not Beadle. Crataegus angulata Sarg. Occasionally found with the typical form. Carter Co. Crataegus pruinosa var. brachypoda (Sarg.) Palmer, comb. nov. Crataegus brachypoda Sarg. Thickets and open woods. Circumneutral to calciphile. Occasionally found with the typical form. Southeastern Mo.: Carter and Ripley counties. Crataegus rugosa Ashe. Crataegus onusta Ashe Thickets, pastures, and borders of woods. Cireumneutral to calciphile. North- ern Mo.: Shelby, Mercer, and Harrison counties Crataegus disjuncta Sarg. Crataegus tumida Sarg. Thiekets and open woods. Cireumneutral to calciphile. East-central and southern Mo.: Franklin, Stoddard, Dent, Shannon, and Jasper counties. Crataegus disjuncta var. magnifolia (Sarg.) Palmer, comb. nov. Crataegus magnifolia Sarg. Thiekets and open woods, limestone hills and along small streams. Circum- neutral to ealeiphile. Southwestern Mo.: Cedar, Dade, Lawrence, Barton, and Jasper counties. Crataegus Mackenzii Sarg. Crataegus depressa Ashe, not Presl, in part. En rubicundula Sarg. ckets, pastures and glades. Circumneutral to calciphile. General: inre Adair, St. Louis, Franklin, Washington, Iron, Cape dns Bol- linger, Butler, Miller, TE enton, Greene, and Stone countie Crataegus Mackenzii var. bracteata (Sarg.) Palmer, eomb. nov. re ag bracteata fa Crataegus aperta Crataegus catia ta Barg. Crataegus rigida Sarg. Crataegus seclusa Sarg. Crataegus seducta Sarg. ege open woods, aye and pastures. Circumneutral to calciphile. Cen- ral and southern Mo.: Boone, Franklin, St. Francois, Madison, Bollinger, rein Ripley, Phelps, Pulaski, Miller, Camden, Laelede, m Ozark, Taney, Stone, Barry, Johnson, Jasper, Newton, and MeDonald countie 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 563 Crataegus Gattingeri Ashe Rocky open woods and thickets. Circumneutral. Southeastern Mo.: Madi- son, Iron, and Wayne counties. Crataegus gun Sarg. Open woods and thickets. Circumneutral to calciphile. Eastern Mo.: Schuyler, ‘Lincoln, Iron, and Wayne counties. Crataegus aspera Sarg. Rocky open woods, thickets, and pastures. Circumneutral to calciphile. Southern Mo.: Ripley, Taney, Stone, Barry, and Jasper counties. Crataegus locuples Sarg. Open woods and thickets. Cireumneutral to calciphile. East-central and Southeastern Mo.: St. Louis, Wayne, and Carter counties. Probably a hybrid between Crataegus mollis and C. pruinosa or a related species. Crataegus mollis (T. & G.) Scheele. Summer Haw, Turkey Apple. Crataegus macrophylla Sarg Ashe. Open woods and thickets along streams. Cireumneutral to caleiphile. Gen- eral, but commonest in northern and central Mo., and apparently absent from some of the Ozark counties. Crataegus mollis f. dumetosa (Sarg.) Palmer, comb. nov. Crataegus dumetosa Sar Occasionally found with the typieal form. Jasper and Taney counties. iodo noelensis Sarg. aegus transmississippiensis Sarg. Bor open woods and thickets along streams. Circumneutral to calciphile. Western Mo., south of the Missouri River: Webster, Benton, Dallas, Greene, Barry, Stone, and MeDonald counties. Crataegus dispessa Ashe. Crataegus pyriformis Britton. lata Treleasei Sarg. cky open woods along streams. Cireumneutral to calciphile. Southern Mo.: St. Francois, Shannon, Howell, and Dallas counties. Crataegus lanuginosa S Crataegus Pew. Eus Open woods, thickets, and pastures; limestone hills and along small streams. Cireumneutral to calciphile. Southwestern Mo.: Jasper, Newton, and Barry counties Crataegus Kelloggi Sar Thickets along = streams. Circumneutral to ealeiphile. Scattered: Put- nam, Lincoln, St. Louis, and Greene counties. Probably a hybrid between Crataegus parta and C. mollis. [Vor. 22 564 ANNALS OF THE MISSOURI BOTANICAL GARDEN Crataegus declivitatis Sar Rocky open woods €. bluffs. Cireumneutral to ealeiphile. East-central Mo., local: St. Louis and St. Francois counties. Probably a hybrid between Crataegus mollis and C. pruino Crataegus coccinioides Ashe. Crataegus callicarpa Sarg. Crataegus speciosa Sarg. Thickets and open woods; limestone hills. Cireumneutral to calciphile. South- ern Mo.: St. Louis, Franklin, St. Clair, Cedar, Dade, Jasper, Newton, and Barry counties. Crataegus spathulata Michx. Crataegus apiifolia var. flavanthera Sarg. Crataegus spathulata var. flavanthera Sarg. ex Palmer. Open woods and thickets. Circumneutral. Southwestern Mo.: Taney and Jasper counties. Crataegus Marshallii Eggleston. Parsley Haw. Crataegus apiifolia (Marsh.) Michx., not Medic. Low swampy woods. Circumneutral. Southeastern Mo.: Dunklin and Butler counties. Crataegus Phaenopyrum i f. Medie. Washington Thorn. Crataegus cordata. Thickets and a E woods, Circumneutral to calciphile. Central and southern Mo.: Boone, Maries, Washington, Crawford, St. Francois, Wayne, Carter, Ripley, Shannon, Texas, Howell, Greene, Stone, and Jasper counties. Crataegus uniflora Muench. Crataegus trianthophora Sar Crataegus tomentosa PN not L. (G). Rocky open woods and glades. Cireumneutral to oxylophile. Southern Mo.: Ripley, Shannon, Wayne, Carter, Texas, Douglas, and Ozark countie Crataegus calpodendron (Ehrh.) Medic, Crataegus tomentosa of auth., not L. (1). Thickets, open woods, and rocky ground along streams and bluffs. Circum- neutral to ealeiphile. General. Crataegus calpodendron var. obesa (Ashe) Palmer, comb. nov. Crataegus obesa Ashe. Crataegus globosa Sarg. Il & B). Crataegus molli Sar Rocky woods, thickets, pm — of streams. Circumneutral to calciphile. Central and southern Mo.: St, 8, Iron, Madison, Wayne, Carter, Ripley, Shannon, Oregon, Jackson, leida "Hiekor ory, Polk, Greene, Christian, Ozark, Taney, Stone, Barry, Jasper, and MeDonald countie Crataegus De var. hispidula (Sarg.) E comb. nov. Crataegus hispidula Sarg. Crataegus spinulosa sid 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 565 Thickets, open woods, and banks or bluffs of streams, Cireumneutral to cal- ciphile. Southern Mo., seattered: St. Francois, Polk, Stone, and Jasper counties. Crataegus succulenta Schrad. Crataegus neofluvialis Ashe (B & B). Crataegus macracantha var. neofluvialis (Ashe) Eggleston (G), in part. Crataegus ensifera S Thickets and borders pe woods, along streams. Circum Wee | to calciphile. Seattered: Harrison, Montgomery, Benton, and Christian eounti Crataegus succulenta var. pertomentosa ( iao Palmer, comb. nov. Crataegus pertomentosa Ashe (G), B & B). Crataegus campestris Britton. ky open woods, glades, and rocky ground along streams. Calciphile to eireumneutral. Scattered, mostly in northern and central Mo.: Mercer, Macon, Clark, Marion, Ralls, Wayne, and Jackson counties. Crataegus nuda Sarg. Glades and rocky ground along streams. Circumneutral to calciphile. South- western Mo., local: Taney Co. Perhaps a hybrid between Crataegus crus-galli and C. succulenta. Crataegus simulata Sar Open woods and thickets along streams. Circumneutral to ealeiphile. South- western Mo., local: Jasper Co. Probably a hybrid between Crataegus cal- podendron var. obesa and C. Palmeri. Crataegus incaedua Sarg. Crataegus pudens Sarg. Crataegus swanensis Sarg. Rocky open woods, glades, and banks of streams. Circumneutral to calciphile. Southern Mo., scattered: Shannon, Carter, Dallas, and Taney counties. Crataegus Vailiae Britton. Crataegus missouriensis Ashe. Bluffs and rocky banks of streams. Cireumneutral Southeastern Mo.: Shan- non and Ripley counties. Crataegus collina x viridis Thickets, low hills bordering lowlands. Local: Cape Girardeau Co. Crataegus Engelmannii x uniflora. Rocky glades. Local: Ozark Co. Waldsteinia fragarioides (Michx.) Trattinick. Shaded cherty or sandstone upland slopes. Oxylophile. Southern Mo.: Texas and Douglas counties. Fragaria virginiana Duchesne. Wild Strawberry. Prairies and open eleared lands. Cireumneutral General but scattered. Fragaria virginiana var. illinoensis (Prinee) Gray. Fragaria Grayana Vilmorin (S) Similar situations to the b General and more common, Fragaria vesca L. v ana Porter Prairies and Bodas ap ien M thickets. Cireumneutral. Scattered: Jasper Co. [Vor. 22 566 ANNALS OF THE MISSOURI BOTANICAL GARDEN FRAGARIA CHILOENSIS Duchesne var. ANANASSA (Duchesne) Hort. ex Bailey. Gar- den Strawberry. Fragaria grandiflora Ehrh. Cultivated and occasionally escaped. Scattered: Andrew Co. DUCHESNEA INDICA (Andr.) Focke. Indian Strawberry. Prairies, — woods, and waste ground. eni i Seattered and un- common: Dunklin and Jackson counties. Potentilla arguta Pursh. Rocky woods and prairies. Oxylophile to circumneutral, General but scattered: Schuyler, Linn, Lewis, St. Louis, Carter, Worth, Clinton, Jackson, Greene, and Barry counties. Potentilla norvegica L. var. hirsuta (Michx.) Lehm. See Rh. 28: 213-214. Potentilla monspeliensis L. (G), (B & B). Prairies, meadows, and waste ground. Circumneutral. General. Potentilla rivalis Nutt. Prairies and open banks. Cireumneutral. St. Louis, Jefferson, and Jackson counties. Potentilla rivalis var. millegrana (Engelm.) Wats. Prairies, meadows, and waste ground. Circumneutral. Central Mo., and per- haps more general: St. Louis, Jefferson, Boone, and Jackson counties, Potentilla rivalis var. pentandra (Engelm.) Wats. s and waste ground. Circumneutral, Local: Jackson Co. Potentilla Salad Nutt. Rocky open woods, upland prairies, and banks of streams, Circumneutral. Central and southern Mo.: St. Louis, Perry, Boone, Baline, Carroll, Clay, Jaek- son, and Holt counties. Potentilla Nicolletii (Wats.) Sheldon. andy bottoms along Missouri and Mississippi Rivers. Oxylophile. Scat- u Perry, Carroll, and Jackson counties. POTENTILLA RECTA L. Fields, meadows, and waste ground. Indifferent. Introduced and becoming more common. General but seattered: St. Louis, Jefferson, Franklin, Wash- ington, St. Francois, Madison, Iron, Crawford, Phelps, Boone, Saline, Jackson, and Jasper counties. Potentilla canadensis L. var. villosissima Fernald, Rh. 33: 187. 1931. Potentilla caroliniana of Rydb., not Poir. (G), (B & B). Dry open woods, prairies, and open banks. Oxylophile. Southern Mo., east- ern Ozark region: Dent, Reynolds, Shannon, Texas, and Howell counties. Potentilla simplex Michx. See Rh. 33: 188, 1931. Cinquefoil. Potentilla canadensis of auth., not L. (G), (B 4 B). y open woods, prairies, and waste ME. Oxylophile to circumneutral. General and common. Potentila simplex var. calvescens Fernald, Rh. 33: 189. Potentilla canadensis var. simplex of auth., not (ies) ES & G. (G). Potentilla simplex of auth., not Miehs: (B & B 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 567 ocky open woods and prairies. Circumneutral: St. Louis, Boone, Jackson, Douglas, Greene, and Jasper countie Potentilla simplex var. argyrisma nl Rh. 33: 191. 1931. Prairies and dry open ground. Cireumneutral. Seattered in southwestern o.: Barton and Pettis counties. d Geum canadense Jacq. Rich woods. Circumneutral. Scattered: Lewis, Monroe, Dallas, Hickory, and Miller counties. Geum canadense var. camporum (Rydb.) Fernald & Weatherby, Rh. 24: 49. 1922. Red Root Geum canadense of auth. in part (G), (B & B). Woods and meadows. Cireumneutral General and common. Geum laciniatum Murr. var. trichocarpum Fernald, Rh. 37: 293. 1935. Rough Ave eu virginianum of auth. in part, not L. (G). Moist woods. Circumneutral. Northern and central Mo.: Scotland, Adair, Marion, Ralls, Monroe, Audrain, and Jackson counties. Geum vernum (Raf.) T. & G. Early Water Avens Moist woods, thickets, and open boggy pui Circumneutral. General. Rubus idaeus L. var. strigosus (Michx.) Maxim. Red Raspberry. Rubus idaeus var. aculeatissimus (C. A. Mey.) Regel & Tiling (G). s hills in northwestern Mo., and also in Jackson Co., where it is probably E Atchison, Holt, and Jackson counties. Rubus occidentalis L. Black Raspber Thickets and open woods. Cireihipen bul. General and probably in every eounty. RUBUS PROCERUS P. J. Muell Cultivated and occasionally escaped. Waste ground. Scattered: Jasper Co. Rubus allegheniensis Porter. High-bush Blackberry. open woods, thickets, and bluffs. Cireumneutral. General but scattered. Rubus alumnus Bailey, Gent. Herb. 1: 191, fig. 88. 1923. Thickets, and banks and bluffs of streams. Circumneutral to oxylophile. Eastern, central, and southern Mo.: Marion, St. Francois, Iron, Madison, Bol- linger, Cape Girardeau, Scott, mih wins Butler, Wayne, Carter, Shannon Phelps, Morgan, Jaekson, Stone, Jasper, and Newton sion 8. Rubus laudatus Berger. Rocky thickets and borders of woods. Circumneutral. Central and southern Mo., seattered: Boone, Wayne, Saline, Jackson, Barry, and MeDonald counties. Rubus Deamii Bailey, Gent. Herb. 2: 463, fig. 203. 1932. Thiekets and old fields. Cireumneutral. Southern Mo.: Texas Co. Rubus 64 Sap s Rydb. High-bush Blackberry. Ru Andrewsianus Blanchard (G). Sube argutus of auth., not Link. (B & B). Woods, thiekets, fields, and fence rows. Circumneutral. General, and prob- ably in every eounty [VoL. 22 568 ANNALS OF THE MISSOURI BOTANICAL GARDEN Rubus Bushii Bailey, Gent. Herb. 2: 403. 1932. High rocky prairies, thickets, and open ground. Circumneutral to oxylophile. Southwestern Mo.: Barton and Jasper counties. Rubus frondosus Bigel. Thiekets and woods along streams. Cireumneutral. General but not eommon: Clark, Marion, Macon, Iron, Phelps, Boone, Cooper, Morgan, Jasper, and Newton eounties. — reg Bailey, Gent. Herb. 2: 452, figs. 196, 197, 198. 1932. , pastures, and waste ground. Cireumneutral. Scattered: Boone, Neon = Clay eountie Rubus heterophyllus Wi na. Rubus recurvans Blanchard (G). Dry open ground and borders of woods. Cireumneutral Northwestern Mo.: Nodaway and Jackson counties. RUBUS LACINIATUS Willd. Cultivated ground. Introduced in Barry Co. Rubus missouricus Bailey, Gent. Herb, 2:459, fig. 200. 1932. Prairies and thickets. Circumneutral. Western Mo.: Jackson Co. Rubus trivialis Michx. Rubus rubrisetus ideae e (B & B), in part. Rubus continentalis Moist thickets and open Ludi. Cireumneutral. Central and southern Mo.: St. Louis, Jefferson, Perry, Cape Girardeau, Bollinger, Madison, Iron, Missis- sippi, Pemiseot, Butler, Ripley, Carter, Shannon, Phelps, Jackson, Jasper, and McDonald counties. Rubus nefrens Bailey, Gent. Herb. 1: 239, fig. 111. 1925 Prairies, old fields, and open banks. Cireumneutral. Central and southern Mo., scattered: Bollinger, Iron, Texas, Jackson, and Jasper counties. Rubus flagellaris Willd. See Gent. Herb. 2: 317. 1932. Dewberry. Rubus villosus Ait. = Rubus procumbens Muhl. (B & B). Open rocky woods, delis, prairies, and railway embankments. Cireum- neutral. General and probably in every eounty. Rubus flagellaris var. occidualis Bailey, Gent. Herb. 2: 318, 1932, Thiekets and banks along prairie streams. Cireumneutral. General but seattered: Clark, Marion, Audrain, Texas, Saline, Jackson, Vernon, and Jasper counties Rubus invisus Bailey. iekets, prairies, and rocky banks. Circumneutral. Jackson and Morgan counties. Agrimonia pubescens Wallr. See N. Am. Fl. 22: 393. 1913. Agrimonia mollis (T. & G.) Britton (G), (B & B). Rocky open woods and thickets. Cireumneutral. General. Agrimonia mer. Wallr. Low ground along streams. Cireumneutral. Eastern Mo., south of the Mis- souri Diver, scattered: St. Louis and St. Francois counties 1935] PALMER € STEYERMARK——PLANTS OF MISSOURI 569 nia gryposepala Wallr. ocky open woods. Cireumneutral. Scattered: Clark, Putnam, Macon, and Douglas countie Agrimonia parto Ait. L pen ds, mo p and boggy ground along prairie streams. uu HE to siat General. Agrimonia rostellata Wallr. Open woods, rocky slopes, and thickets. Circumneutral. Southern and cen- tral Mo., south of a line from St. Louis, Warren, and Boone counties to Jack- son Agrimonia platycarpa Wallr. Low or rocky woods. Circumneutral. General, but seattered: Marion, Ralls, St. Louis, St. Francois, Shannon, Texas, Greene, Taney, Clay, Jackson, Jasper, and Newton counties. Rosa setigera Michx. Prairie Rose, Climbing Ros Low open woods and moist rocky ground ira reams, Circumneutral. Cen- tral and southern Mo., seattered: Bollinger, Scott, Dunklin, Butler, Shannon, Pulaski, Saline, Pettis, and Jasper counties. Rosa setigera f. inermis Palmer & Steyermark, f. n en woods and rocky ground along streams. Boetius found with the typieal form. Cireumneutral. Southeastern Mo., seattered: Bollinger and Shannon counties. A typo recedit ramulis inermibus vel raro paucis tenuibus spinis. The unarmed glabrous form appears to be comparatively rare and we have seen specimens only from southeastern Missouri. It differs from the type only in the canes and branches being entirely unarmed or rarely with a few small scattered spines. MISSOURI: Patton, Bollinger Co., April 26, 1931, E. J. Palmer 39093 (Arn. Arb. TYPE); Monteer, Shannon Co., July 30, 1930, B. F. Bush 11907 (Arn. Arb.). Rosa setigera var. tomentosa T. & G. Rocky open woods, verb and prairies. Cireumneutral. General and prob- ably in every county. Rosa setigera var. serena Palmer and a var. pen woods, and rocky banks or bluffs of streams. C eutral. Southern Mo.: Iron, R D. Carter, Ripley, Shannon, Ozark, Taney, Christian, Stone, and Barry counti A typo recedit foliia subter pubescentibus; ramulis inermibus vel raro paucis spinis. This variety is distinguished from var. tomentosa 'T. & G., which it resembles, in the pubescent under surface of the leaves, and by its smooth spineless canes and branehes, although rarely a few small teed thorns may be found plants in inen eel all of the branches are entirely unarmed. It appears to be more dis and constant in its distinguishing characters, as well as more isolated erorapical than the variety based solely on the more or less pubes- cent character of the leaves. In parts of the Ozark region, in southern Missouri and northern ca it is the prevailing form of Rosa setigera. [Vor. 22 570 ANNALS OF THE MISSOURI BOTANICAL GARDEN MissourI: Campbell, Dunklin Co., Oct. 7, 1910, B. F. Bush 6385 (Arn. Arb.); Ponder, Ripley Co., July 2, 1933, Palmer $ Steyermark 41642 (Arn. Arb. TYPE, Mo. Bot, Gard. ISOTYPE) ; Monteer, Shannon Co., Oet. 6, 1920, E. J. Palmer 1 (Arn. Arb.); Galena, Stone Co., Sept. 25, 1923, E. J. Palmer 23844, and Sept. 15, 1934, 26135 (Arn. Arb.); Roark, Stone Co., Sept. 28, 1920, E. J. Palmer 19213 (Arn. Arb.) ; Melva, Stone Co., Sept. 17, 1924, E. J. Palmer 26194 (Arn. Arb.) ; Branson, Taney Co., June 8, 1914, E. J. Palmer 5897 (Arn. Arb.) ; Swan, Taney i June 9, 1899, B. F. Bush 47 (Arn. Arb.) ; Teeumseh, Ozark Co., Oct, 7. 1927, E. J. Palmer 32902 (Arn. Arb.); Gainesville, Ozark Co., June 26, 1928, E. J. Palmer 34724 (Arn. Arb.); Isabella, Ozark Co., June 27, 1928, E. J. Palmer 3477 (Arn. Arb.); Eagle Rock, Barry Co. Joly 27, 1926, E. J. Palmer 31439 (Arn. Arb.). ARKANSAS: Cotter, Baxter Co., Sept. 19, 1924, E. J. Palmer 26216 (Arm. Arb.); opposite Cotter, Marion Co., June 14, 1914, E. J. Palmer 5978 (Arn. Arb.); Eureka Springs, Carroll Co., Sept. 17, 1921, E. J. Palmer 20486 (Arn. Arb.); Beaver, Carroll Co., April 30, 1926, E. J. Palmer 29843 (Arn. Arb.). Rosa rudiuscula Greene, Leaflets Bot. Obs. & Crit. 2: 134. 1911. Prairies and thickets. Cireumneutral. Northern and western Mo.: Shelby, Pettis, Benton, Gentry, Nodaway, Jackson, Cass, Bates, Vernon, and Jasper counties. Rosa suffulta Greene. See N. Am. Fl. 22: 504. 1913; also Bull. Torr. Bot. Club 50: 65. 1923. Rosa heliophila Greene Rosa pratincola euis, not R. Br. (G). Prairies and thickets. Cireumneutral to ealeiphile. Northern, central, and western Mo., mostly in the prairie region: Seotland, Knox, Boone, Saline, Pettis, Worth, Nodaway, Atchison, Clinton, Platte, Ray, Jackson, Cass, Bates, and Vernon eounties. Rosa suffulta var. valida Erlanson, Rh. 30: 114. 1928. Occasionally found with the typical form. Northwestern Mo.: Atchison Co. Rosa blanda Ait. Meadow Rose. Thickets and open woods. Circumneutral. Northeastern Mo.: Clark and Schuyler counties. Rosa relicta Erlanson, Rh. 30: 116. 1928. Prairies and thickets. Cireumneutral. Northern and central Mo.: Schuyler, behets D Cooper, Morgan, Lafayette, Johnson, Henry, and $t. Clair eo Rosa m Rydb. N. Am. Fl. 22: 501. 1918. kets and open woods. Cireumneutral. Northern and central Mo., scat- Tien Clark, Jackson, Cooper, and Morgan counties. Rosa is viget Rydb. N. Am. Fl. 22: 505. 1918. n banks, loess hills, pe bluffs. Caleiphile to eireumneutral. Northwest- ern Mo.: : Atchison, Nodaway, Dekalb, and Jackson counties, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 571 Rosa Bushii Rydb. N. Am. Fl. 22: 506. 191 Thiekets and open banks. Binti: Northern and central Mo.: Put- nam, Jackson, and Lafayette counties. Rosa Palmeri Rydb. N. Am. Fl. 22: 502. 1918. Prairies and thickets. Circumneutral. Middle and western Mo.: Clinton, Randolph, Jasper, and Stone counties. Rosa subserrulata Rydb. N. Am. Fl. 22: 500. 1918. cky woods and glades. Circumneutral to oxylophile. Central and southern Mo., mostly in the Ozark region and locally north to St. Louis, Pike, and Jackson counties. Rosa Le Rag Rydb. N. Am. Fl. 22: 501. 1918. ets and prairies. Circumneutral. Northeastern Mo.: Clark Co. Rosa polyanthema Lunell, Am. Midl. Nat. 3: 138. 1913. Thiekets, open banks, loess hills, and prairies. Circumneutral to caleiphile. Northwestern Mo.: Grundy, Caldwell, Nodaway, Atchison, Holt, and Buchanan unties. Rosa carolina L. See Rh. 20: 91. 1918. Pasture Rose. Rosa humilis Marsh. (G). Rosa virginiana of auth. in part, not L. (B & B). Thickets, open woods, and prairies. Cireumneutral. General and probably in every county. x Rosa carolina var. grandiflora (Baker) Rehder in Bailey, Stand. Cyclop. Hort. 5: 2991. 1916. Rosa obovata Raf. open woods and thickets. Circumneutral. General but scattered: Clark, Lewis, Marion, Putnam, Adair, ipd Daviess, Shannon, Texas, Wright, Cedar, Hickory, Jasper, and Newton coun Rosa carolina var. glandulosa (Crepin) Nod Rept. Mich. Acad. Sei. 21: 366. 1920. Rosa serrulata Open woods and dud Cireumneutral. Seattered: Maries, Schuyler, Clin- ton, and Jasper eounties. Rosa carolina var. Lyoni (Pursh) Palmer € Steyermark, comb. nov. Rosa Lyoni Pursh. Open woods, thickets, and rocky prairies. Circumneutral. General. Rosa palustris Marsh. See Rh. 20: 91. 1918. Swamp Rose. osa carolina of auth., not L. (G), ( 007 Low swampy woods. Circumneutral. Southeastern Mo., in the lowlands: Cape Girardeau, Bollinger, Madison, Iron, Scott, Mississippi, Stoddard, Dunklin, Butler, Wayne, and Ripley counties. Rosa palustris var. dasistema (Raf.) Palmer € Steyermark, comb. nov. Rosa dasistema Raf. Swamps and low wet woods. Cireumneutral. Southeastern Mo.: in the low- lands: Scott, Stoddard, Dunklin, Butler, Wayne, and Iron counties. [Vor. 22 572 ANNALS OF THE MISSOURI BOTANICAL GARDEN ROSA RUGOSA Thunb, ltivated and escaped into fields and along railroads. Southwestern Mo., local: Greene Co. Rosa EGLANTARIA L. Sweet-brier. sa rubiginosa L. (G), (B & B). Formerly planted in gardens and escaped and established in pastures and waste ground. Seattered: Clark, Marion, Montgomery, St. Louis, St. Franeois, Boone, Dekalb, Clinton, Jackson, Johnson, and Jasper counties. Rosa CANINA L. Dog Rose. Occasionally escaped from gardens to roadsides and waste ground. Dade Co. ROSA SPINOSISSIMA L, Occasionally escaped to roadsides and waste ground. Boone Co. ROSA CENTIFOLIA L. garden rose, occasionally found as an escape in pastures and along is. Clark Co. Prunus serotina Ehrh. Wild Cherry. Upland woods and along streams. Circumneutral. General and probably in every county. Prunus virginiana L. Choke Cherry. Thickets and borders of woods. Circumneutral. Northern and central Mo., mostly north of the Missouri River: Clark, Marion, Shelby, Schuyler, Putnam, dair, Mercer, Harrison, Caldwell, Worth, Gentry, ge Nodaway, Atchison, Buchanan, Sullivan, Boone, Saline, and Benton eounti Prunus angustifolia Marsh. Chickas lum. Thickets and pastures. a General but scattered. Perhaps in troduced by the Indians in prehistoric times: Grundy, Bollinger, Madison, Butler, Shannon, Jackson, Cass, Johnson, Pettis, Ozark, Stone, and Jasper counties Prunus angustifolia var. varians Wight & Hedrick, Plums of N. Y. p. 87. 1911. Thickets, roadsides, and along prairie streams. Circumneutral, Central and southern Mo.: Dunklin, Ripley, Oregon, Johnson, Benton, Jasper, and Newton counties. Prunus Munsoniana Wight € Hedrick, Plums of N. Y. p. 88. 1911. Wild Goose Thiekets, prairies, borders of streams, and waste ground. Cireumneutral. General but scattered: Adair, Knox, Shelby, Marion, Monroe, Audrain, Boone, St. Louis, e Girardeau, Madison, Iron, Dunklin, Butler, Ripley, Shannon, Putnam, Mercer, Harrison, Caldwell, Jackson, Bates, Vernon, St. Clair, Dade, Taney, Stone, Jasper, Newton, and MeDonald counties, Prunus hortulana Bailey. Thickets, open woods, and along streams. Cireumneutral. General but com- monest in central and southern Mo. Prunus hortulana var. pubens Sarg. Trees € Shrubs 2: 248. 1913. Thiekets and open woods. Cireumneutral. General but scattered: Marion, Pike, Boone, Howard, Shannon, Ozark, Jackson, and Jasper counties, 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI. 573' Prunus americana Marsh. Wild Plum. Thickets, open woods, and pastures. Circumneutral. General. Prunus lanata (Sudw.) Mack. & Bush. Prunus arkansana Sarg. ] | Thickets and open woods. Circumneutral. General. Prunus Palmeri Sarg., found in southwestern Mo. and later referred to this species, may be distinct or possibly a hybrid between this and P. hortulana. Prunus mexicana Wats. Big Tree Plum ; Rocky open woods. Obamas Southern Mo.: Dunklin, Butler, Shan- non, Ozark, xt and Stone counties. aba SPINOSA L. aan in thickets. Jasper Co. PRUNUS MAHALEB L. Perfumed Cherry. Thickets, pastures, and roadsides. Used as grafting stock by nurserymen and often escaped. Scattered: Marion, Gasconade, Jackson, and Jasper counties. PRUNUS CERASUS L. Sour Cherry. Thickets and roadsides. Cultivated and occasionally escaped. Scattered: Marion, Holt, Jackson, and Barry counties Fam. LEGUMINOSAE Acacia angustissima (Mill.) Ktze. Prairie Acacia. Glades and rocky open ground. Circumneutral to caleiphile. Southwestern Mo., local: Barry Co. Acacia angustissima var. hirta (Nutt.) Robinson Glades, rocky hillsides, and exposed odios. along bluffs. Circumneutral to ealeiphile. Southwestern Mo.: Jasper, Newton, McDonald, Barry, Stone, and Taney counties. : Desmanthus illinoensis (Michx.) MacM. False Sensitive Plan ) open ground, thickets, and prairies. Circumneutral a oxylophile. Gen- eral but apparently absent from much of the Ozark region Desmanthus leptolobus T. & G Low open ground. Circumneutral. Southeastern Mo.: Cape Girardeau Co. Perhaps introduced. Schrankia uncinata Willd. Sensitive Brier. Rocky prairies, open banks, and glades. Cireumneutral to ealeiphile. General but absent from many of the eastern and northern eounties: Putnam, Marion, Shelby, Macon, Audrain, Boone, Cole, Gasconade, Jefferson, Washington, Craw- ford, Iron, Wayne, Carter, Ripley, Oregon, Shannon, Howell, Dent, Osage, Phelps, Pulaski, Maries, Moniteau, Morgan, Laclede, Dallas, Wright, Webster, Hiekory, Ozark, Douglas, Greene, Taney, Stone, Jackson, Johnson, Vernon, Bar- ton, Jasper, Newton, and McDonald counties. Gymnocladus dioica (L.) Koch. Kentucky Coffee-tree. Woods, chiefly along small streams: sometimes in alluvial bottoms. Circum- neutral, General but nowhere very common. [Vor. 22 574 ANNALS OF THE MISSOURI BOTANICAL GARDEN Gleditsia triacanthos L. Honey Locust. bes chiefly along small streams; sometimes in low alluvial ground. Cir- cumneutral, General and probably in every county. und triacanthos var. inermis Pursh. T abend found with the common form, mostly in eastern and southern : Harrison, Mercer, Shelby, Monroe, St. Louis, Jefferson, Franklin, Wash- “a Iron, Mississippi, and MeDonald ed ies. — I: Marsh. Water Locust. woods and swamps. Circumneutral. Southeastern Mo., in the low- itn] us pes north along the Mississippi River: St. Charles, St. Louis, Jef- ferson, Cape — — Scott, Mississippi, New Madrid, Dunklin, Butler, and Ripley coun Hoffmanseggia Jamesii T. & G. Dumps and waste ground. Found as a waif in Jasper Co. Cassia Medsgeri Shafer. Thickets, open woods, and along rocky branches. Circumneutral. General. This has often been confused with Cassia marilandica, an eastern species which it resembles Cassia Tora L. Rocky open ground "e streams, and sandy bottoms. Oxylophile. South- eastern Mo.: Dunklin and also introduced in Jackson Co. CASSIA OCCIDENTALIS L, Coffee Wee ow alluvial ground. Deest ten Seattered: Introduced in Jackson and New Madrid counties. Cassia fasciculata Michx, Partridge Pea. Cassia Chamaecrista of auth., not L. (G). Chamaecrista fasciculata (Michx.) Greene (B & B Thickets, glades, and rocky prairies. Cireumneutral to oxylophile. General. Cassia fasciculata f. Jenseni Palmer & Steyermark, f. nov. Rarely occurring with the typical form. Local: Franklin Co. A typo recedit petalis albis. MISSOURI: Gray Summit, Franklin Co., Aug. 20, 1928, L. p pu (Mo. Bot. Gard, TYPE). See Jensen, Mo. Bot. Gard. Bull, 19: 38-39. Differs from the typical form in having white instead of rom petals. Cassia fasciculata var. robusta (Pollard) Macbride, Contr. Gray Herb. N. $. 59: 24. 1919 Cassia Chamaecrista var. robusta Pollard (G). j Chamaecrista fasciculata (Michx.) Greene, in part (B & B). Occasionally found with the typical form. Circumneutral. Scattered: Lin- coln, St. Charles, St. Louis, Perry, Pemiscot, Boone, and Polk counties. Cassia fasciculata var. depressa (Pollard) Macbride. Cassia depressa Pollard (G). Chamaecrista depressa (Pollard) Greene (B & B Rocky open and alluvial ground. Clrearmeatest, Eastern Mo.: Washing- ton Co 1935] PALMER € STEYERMARK—— PLANTS OF MISSOURI 575 Cassia nictitans L. Sensitive Pea Rocky woods, thickets, and prairies. Oxylophile to eireumneutral. Central and southern Mo, south and east of a line from Lincoln, Montgomery, Boone, and St. Clair counties to Vernon Co. Cercis canadensis L. Redbud. Thickets and borders of woods, along rocky streams and bluffs. Cireum- neutral, General and probably in every county. ne leucantha T.& G. White Wild Indigo. Prairies, glades, and rocky hillsides. Cireumneutral. General, NUM vespertina Rydb. Blue False Indigo. australis of auth., not R. Br. B& ES prairies and glades. Caleiphile to iai? Central and southern Mo.: St. Louis, Jefferson, Franklin, St. Francois, Ste. Genevieve, Ozark, Taney, Stone, Barry, Greene, Hickory, Cedar, Vernon, Jasper, and Newton counties, and also introduced in Jaekson Co. Baptisia leucophaea Nutt. Baptisia bracteata of auth., not (Muhl.) Ell. (G), (B & B). Rocky prairies, meadows, and open woods. Circumneutral. General. Baptisia leucophaea var. laevicaulis Gray. Sometimes found with the typieal form. Central and southern Mo. RN leucophaea x vespertina. rely found with the parent species. Jasper Co. "en sphaerocarpa Nutt Introduced from farther south, along railways. Circumneutral. Scattered: St. Louis and Barry counties. en lutea (Michx. f.) Koch. Yellow Woo ong high rocky bluffs of White River and its tributaries, Circumneutral to Sn outhwestern Mo.: Taney, Stone, and Barry counties Crotalaria sagittalis L. Rattle-box. Prairies, glades, and open sandy or rocky ground. Circumneutral to oxylo- phile. General TRIFOLIUM ARVENSE L. Rabbit-foot Clover. Waste ground and along roadsides and railroads. Indifferent. Scattered: St. Louis, Phelps, Texas, Greene, Barry, and Jasper counties. TRIFOLIUM INCARNATUM L. Crimson Clover. Fields and waste ground. Indifferent. Scattered in middle and southeastern Mo.: Boo St. Louis, Jefferson, Ste. Genevieve, Bollinger, Iron, Dunklin, Ripley, d Dun eountie TRIFOLIUM PRATENSE L. Red Clover. Fields, meadows, waste and cultivated ground. Indifferent. General and common. Trifolium reflexum L. Buffalo Clover. Rocky open woods, glades, and prairies. Oxylophile to cireumneutral. North- western, eentral, and southern Mo.: St. Louis, Jefferson, St. Franeois, Ste. Gene- vieve, Iron, Madison, Bollinger, Wayne, Carter, Ripley, Oregon, Washington, [Vor. 22 576 ANNALS OF THE MISSOURI BOTANICAL GARDEN Howell, Shannon, Texas, Dent, Crawford, Maries, Phelps, Cooper, Morgan, Web- ster, Douglas, WI Johnson, Greene, and Jasper counties. Trifolium reflexum var. glabrum Lojacono. ently EC with the typical form, in similar situations. Central, ME. and wester : Warren, St, Louis, Jefferson, Franklin, Phelps, Cole, Atehison, Jackson, Cann, "Webster, Greene, Barry, and Jasper counties. Trifolium stoloniferum Muhl. Running Buffalo Clover. Open woods and prairies. Circumneutral. Central and southern Mo., seat- tered: Boone, St. Louis, Wayne, and Jasper counties. Trifolium repens L. White Clover. Fields, pastures, and waste ground. Indifferent. General and common. TRIFOLIUM HYBRIDUM L. Alsike Clover. Fields, roadsides, and waste ground. General and common, Trifolium carolinianum Michx. es and rocky ictus Cireumneutral to oxylophile, Southwestern Mo.: Phasa and Newton counti TRIFOLIUM AGRARIUM L. Yellow Hop Clover Fields and waste ground. Scattered: St. Charles and Christian counties, TRIFOLIUM PROCUMBENS L. Low Hop Clover. ocky open ground, roadsides, and waste ground: Cireumneutral. Eastern, central, and southern Mo. south of a line from Marion and Boone counties to Caldwell Co. TRiFOLIUM DUBIUM Sibth. Little Hop Clover. Roadsides and waste ground. Indifferent. Seattered: Boone, Johnson, Greene, and Jasper counties. TRIFOLIUM RESUPINATUM aste and eultivated — Introduced in Johnson Co. MELILOTUS OFFICINALIS (L.) Lam. Yellow Sweet Clover. long roadsides, punitus and in waste ground. Indifferent, General, Re- cently heredad ta laces MELILOTUS ALBA Desf. White Sweet Clover. Roadsides, railroads, and waste ground. Indifferent. General and common. MEDICAGO SATIVA L. Alfalfa. ommonly cultivated in recent years, and often escaping along railroads and roadsides. Indifferent. Genera MEDICAGO LUPULINA L. Black Medick. Roadsides and waste ground. Indifferent. General. MEDICAGO HISPIDA Gaertn. Bur Clover. Introduced along railroads and in waste ground. Jackson Co. — en tus L. Ground Honeysuckle e and eultivated ied troll: in Boone and Clay counties. Lotus americanus (Nutt.) Bisch. Prairie Trefoil. Hosackia americana (Nutt.) Piper (G), (B & B). Prairie swales and glades, and also frequently adventized along roadsides and 1935] PALMER & STEYERMARK— PLANTS OF MISSOURI 577 railroads. Cireumneutral. Central and southern Mo.; mostly in the western prairie region: Jefferson, Saline, Jackson, Henry, Vernon, Barton, Jasper, New- ton, and McDonald counties. Psoralea Onobrychis Nutt. French Gra Open woods, low alluvial piri m" river banks. Circumneutral. North- eastern and east-central Mo.: Clark, Marion, Ralls, Shelby, St. Louis, Jeffer- son, and Franklin counties. Psoralea pedunculata (Mill.) Vail. pen rocky woods, glades, and prairies. Circumneutral to oxylophile. Central and southern Mo.: St. Louis, Jefferson, Franklin, St. Francois, Washington, Crawford, Iron, Dent, Shannon, Reynolds, Phelps, Maries, Pulaski, Texas, Laclede, Howell, Camden, Wright, Webster, Douglas, Hiekory, Greene, Lawrence, Bates, Vernon, Jasper, Newton, and MeDonald counties. Psoralea tenuiflora Pursh. Scurfy Pea Psoralea tenuiflora var. bene (Nutt.) Rydb. (G). soralea floribunda Nutt. (B Glades, rocky or open a ae open woods. Caleiphile to circumneutral. Southern and central Mo. north to Monroe, Daviess, and Clinton counties. Psoralea argophylla Pursh. Prairies and loess hills. Caleiphile to cireumneutral Northwestern and west- central Mo.: Atchison, Holt, and Jackson counties. Psoralea esculenta Pursh. Prairie Turnip, Indian Bread-ro Rocky prairies and glades. Caleiphile to cireumneutral. y and southern Mo.: St. Louis, Jefferson, Franklin, Miller, Dallas, Greene, Ozark, Taney, Cedar, Jackson, and Bates counties. Dalea alopecuroides Willd. poi leporina (Ait.) Rydb. var. alba (Michx.) Maebride, Contr. Gray Herb. 6 el odo (L. ) Britton (B & B). Loess hills, sandy open ground, and alluvial dien along rivers. Circum- neutral Northern, central, and eastern Mo., seattered: Putnam, Boone, St. Louis, Jefferson, Mississippi Atchison, Holt, in Buchanan, Clay, Jack- son, Cass, and Johnson eountie Dalea enneandra Nutt. Parosela enneandra (Nutt.) Britton (B & B). Loess hills and Bened Calciphile to eireumneutral Northwestern Mo.: Atchison and Holt cou Petalostemum purpureum eas Rydb. Red Tassel-flower, Prairie Clover. Prairies, glades, and rocky hillsides. ik ai to eireumneutral General. Petalostemum purpureum var. pubescens Gr Oeeasionally found with the ea MN Seattered: Shannon, Taney, Barry, Jasper, and McDonald counties. Petalostemum multiflorum Nutt. Rocky prairies and along railroads. Adventized from farther west. Jack- son Co. [Vor. 22 518 ANNALS OF THE MISSOURI BOTANICAL GARDEN Petalostemum candidum Michx. White Tassel-flower, White Prairie Clover. Prairies, glades, and rocky hillsides. Calciphile to eireumneutral. General, Tephrosia virginiana (L.) Pers. Goat’s Rue, Hoary Pea Rocky open woods, prairies, and glades, Circumneutral to oxylophile. € eral: common in the Ozark region and seattered northward; known north o the Missouri River in Putnam, Monroe, Warren, rw and Howard counties. Tephrosia virginiana var. holosericea (Nutt.) T. & G. Oceasionally found with the typical form. Avi and Cedar counties. Sesbania macrocarpa Muhl. Low sandy fields and sand bars of streams. Oxylophile, Southern Mo., in the southeastern lowlands and loeally along White River: Mississippi, New Madrid, Pemiscot, Dunklin, and Barry counties. Wisteria macrostachya Nutt. Low woods along streams, and borders of swamps and bayous. Cireumneutral. Southeastern Mo.: Scott, Mississippi, Pemiscot, — Butler, and Ripley counties, and also escaped from cultivation in Adair Wisteria frutescens (L.) Poir. Cultivated, and escaped in Daviess Co. Robinia Pseudo-Acacia L. Black Locust. Rocky upland woods, pastures, and waste ground. Circumneutral. General; native in parts of the Ozark region and escaped from cultivation elsewhere. Astragalus caryocarpus Ker. Ground Plum. Prairies and glades. Circumneutral to yas AN Western Mo., scattered: Atchison, Jackson, Vernon, Greene, and Jasper counties Astragalus mexicanus A. DC. Ground Plum. cky prairies, glades, ledges along bluffs, and rocky open woods. Caleiphile to eireumneutral. Central and southern Mo., south and east of a line drawn from St. Charles, Montgomery, Boone, Morgan, and Henry counties to Jasper Co. Astragalus canadensis L. Rattle Weed. Open woods, thickets, and prairies. _Circumneutral, General: Schuyler, Knox, Marion, Ralls, St. Charles, St. Louis, Jefferson, Mississippi, Ripley, Oregon, Shannon, esse, Howell, Phelps, Pulaski Miller, Morgan, Boone, Sullivan, Linn, Mereer, Harrison, Atchison, Dekalb, Jackson, St. Clair, Dallas, Greene, Cedar, Stone, and Jasper counties. Astragalus distortus T. & G. Roeky prairies and glades. pri pd to circumneutral, General but scat- tered: Marion, Ralls, Monroe, St. Louis, Jefferson, Franklin, Scott, Butler, Shannon, Moniteau, Miller, iiem, Boone, Lavine’ on, Jackson, Cass, Johnson, Hickory, Polk, Vernon, Dade, Greene, Jasper, and MeDonald counties. Astragalus lotiflorus Hook. Loess hills and plains. Calciphile to cireumneutral. Northwestern Mo.: Atchison and Holt counties. oe plattensis Nutt. Loco Weed. ropis Lamberti Pursh (G), (B & B). bu hills. Calciphile. Northwestern Mo.: Atchison and Holt counties. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 579 Amorpha fruticosa L. False Indigo. Low open woods, thickets, and banks of streams. Circumneutral. General and probably in every county. Amorpha fruticosa var. tennesseensis (Shuttlw.) Palmer, Jour. Arnold Arb. 12: i Z 20b. orpha ied Shuttlw. nd banks and moist ground along prairie streams. Cireumneutral. South- ern Mo.: Shannon, Ozark, Taney, Stone, Barry, Jasper, and McDonald counties. Amorpha fruticosa var. angustifolia Pursh. Wet ground along streams and borders of ponds. Circumneutral. Central and southern Mo. Amorpha fruticosa var. oblongifolia Palmer, Jour. Arnold Arb. 12: 192, fig. 20c. Wet ground, thickets, and along prairie streams. Cireumneutral Western Mo.: Atchison and Jasper counties. Amorpha croceolanata Wats. Sandy open ground. Cireumneutral to oxylophile. Southeastern Mo.: Dunk- o Amorpha canescens Pursh. Lead Plant. Rocky open woods, prairies, and glades. Oxylophile to circumneutral. General. Amorpha canescens var. glabrata Gray. Rocky open woods, oe and glades. Oxylophile to circumneutral, Central and southern Mo.: ries, a Macon, Benton, Hickory, Dallas, Morgan, Ozark, "ipiis and Jasper countie Amorpha brachycarpa Palmer, Jour. Arnold Arb. 12: 171, pl. 36. fig. 6. 1931. Rocky open woods and glades. Cireumneutral. Southwestern Mo.: Stone Co. Glycyrrhiza lepidota (Nutt.) Pursh. Wild Liquorice. Loess hills, prairies, and waste ground. Calciphile to cireumneutral Native in northwestern Mo. and introduced in other parts of the state. Scattered: Marion, St. Louis, Jefferson, Moniteau, Gentry, Atchison, Holt, Andrew, Jack- son, and Jasper counties CORONILLA VARIA L. Crown Vetch. Roadsides and waste ground. Cireumneutral St. Louis and Taney counties. Introduced near St. Louis by Fendler. Desmodium acuminatum (Michx.) DC. Beggar's Lice, Beggar's Tieks. Desmodium grandiflorum (Walt.) DC. (G). Meibomia grandiflora (Walt.) Ktze. (B € B). Rocky open or rich low woods and bluffs. Cireumneutral General. Desmodium nudiflorum (L.) DC. Tick Trefoil. oeky woods. Oxylophile. Central and southern Mo., south of a line drawn from Ralls and Randolph counties to Jaekson Co. —€— € — (Nutt. i Rich or rocky woods and thi H Cireumneutral. Central and southern Mo.: Boone, Franklin, Bt ph uis, filer Scott, Mississippi, New Madrid, Dunklin, Stoddard, Butler, Ripley, Pulaski, Greene, Jasper, and Newton counties. [Vor. 22 580 ANNALS OF THE MISSOURI BOTANICAL GARDEN Desmodium rotundifolium (Michx.) DC. Roeky open woods. Oxylophile. Central and southern Mo., south and east of a line drawn from St. Louis, Warren, Boone, Morgan, and Hickory counties to Jasper Co. Desmodium ochroleucum M. A. Curtis. Open woods and low open ground. Cireumneutral. Southeastern Mo.: Stod- dard and Shannon counties, Desmodium canescens (L.) DC. Beggar's Lice, Beggar's Ticks. ry open woods and thiekets. Circumneutral to oxylophile. General. Desmodium canescens var. hirsutum (Hook.) Robinson. In similar situations to the typical form. Cireumneutral to oxylophile. Scat- tered: St, Louis, Jefferson, Butler, Boone, Texas, Greene, Jasper, and Jackson counties. Desmodium bracteosum (Michx.) DC, Dry open woods and thickets. Cireumneutral to ede General. Desmodium bracteosum var. longifolium (T. € G.) Robinso In similar situations to the typical form. Seatte uh “Marion, St. Louis, Gentry, Boone, Jackson, and Jasper countie Desmodium illinoense Gray. Prairies, glades, - — open woods. Circumneutral to A North- ern, central, and western Mo., absent from the Ozark r : Adair, Lewis, Marion, Ralls, iini veto. Boone, Saline, Pettis, Worth, sae "Dekalb, Clinton, Caldwell, Jackson, Greene, and Jasper counties. Desmodium laevigatum (Nutt. Rocky open woods, istnd to oxylophile. Eastern and southern Mo.: Ralls, Ste. Genevieve, Iron, Reynolds, Shannon, Phelps, Pulaski, Douglas, Ozark, Taney, Barry, and Jasper counties. Desmodium viridiflorum (L.) Beck. Open woods, glades, and rocky prairies. Cireumneutral to oxylophile. Southern Mo.: Scott, Dunklin, Ripley, Shannon, Howell, Barry, and MeDonald counties. Desmodium Dillenii Darl. oc , open woods, thickets, and glades. Oxylophile to cireumneutral. Genera Desmodium — um (L.) DC. Rich woods and thickets and alluvial ground. Circumneutral to calciphile. General. Desmodium paniculatum var. pubens T. & G. In similar situations to the typical form. General. Desmodium paniculatum var. angustifolium T. € G. In similar situations to the typical form. General. Desmodium sessilifolium (Torr.) T. € G. Rocky open woods, prairies, and glades. Cireumneutral to calciphile. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 581 Desmodium canadense (L.) DC. Prairies and open thickets. Circumneutral. Northern, central, and western Mo., apparently absent from the Ozark region: Clark, Schuyler, St. Louis, Mercer, Saline, Jackson, and Jasper counties. Desmodium rigidum (Ell.) DC. oeky woods, glades, and prairies. Oxylophile to circumneutral. Southern and central Mo.: Lineoln, St. Louis, Jefferson, St. Francois, Dunklin, Butler, Iron, Dent, Crawford, Pulaski, Cole, Howell, Moniteau, Boone, Douglas, Taney, Barry, Jasper, Newton, and rey counties. Desmodium obtusum (Muhl.) D Dry open woods, ip S sandy or roeky open ground. Oxylophile. Cen- tral and southern Mo.: Boone, St. Louis, Jefferson, Scott, Mississippi, Dunklin, Stoddard, Reynolds, ae Dent, Phelps, Pulaski, Howell, Maries, Dallas, Ze Henry, Pettis, Greene, Ozark, Stone, Barry, Jasper, and MeDonald eounties Desmodium marilandicum (L.) DC. Dry open woods and thickets. Oxylophile. Central and southern Mo.: Mont- gomery, Warren, Lineoln, St. Louis, Jefferson, Franklin, Iron, Wayne, Ripley, Reynolds, Dent, Crawford, Maries, Phelps, Pulaski, Camden, Greene, Taney, Stone, Barry, Jackson, Jasper, Newton, and MeDonald eounties. Desmodium canadense x Dillenii. Rarely found with the parent speeies. Loeal: St. Louis Co. Lespedeza procumbens Michx. Bush Clover. Dry rocky woods. Oxylophile to eireumneutral. Southern and central Mo. south and east of a line drawn from St. Louis, Boone, and Hickory counties to Jasper Co. rs repens (L.) Bar open woods. A PUR to eireumneutral. Southern and central Mo. "Ns of a line drawn from Lineoln, Boone, and Saline counties to Cass Co. Lespedeza violacea (L.) Pers. Bush Clover. Lespedeza violacea var. prairea Mack. & Bush ocky or dry open woods, thiekets, and ior prairies. Circumneutral. General. Lespedeza Manniana Mack. & Bush. RE woods and glades. Cireumneutral. Scattered in central and southern : erson, Newton, and Jackson counties. Probably a hybrid between EI G virginica and T capitata. TI Stuevei Nutt. See Rh. 26: 28. 1924. open mh thickets, and prairies. Oxylophile to circumneutral. No Mo.: Ste. Genevieve, een Mississippi, Stoddard, Dunklin, Shannon, Phelps, Pulaski, Howell, Ozark, Taney, Vernon, Barry, Jasper, Newton, and MeDonald counties. — Stuevei var. angustifolia Britton. See Rh. 26: 29. 1924. a Stuvei var. neglecta Britton (G). Occasional found with the typical form. Scattered in southern and central Mo.: St. Louis, Iron, and Pulaski counties. Probably a hybrid between Les- RER virginica and Lespedeza Stuevei (Vor. 22 582 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lespedeza virginica (L.) Britton. Bush Clover. Rocky or dry open woods, thickets, prairies, and glades. Cireumneutral to oxylophile. General and common, Lespedeza intermedia (Wats.) Britton. See Rh. 26: 29-31. 1924. espedeza frutescens Britton, not Hedysarum frutescens L. (G), (B & B). Rocky open woods and thickets. Cireumneutral to oxylophile. Southern and east-central Mo.: St. Louis, Ste. Genevieve, Bollinger, Iron, Butler, Reynolds, — Dent, Phelps, Henry, Taney, Stone, Barry, Jasper, and McDonald counties Lespedeza dei Mack. € Bush. Dry open woods and rocky prairies. Le Southwestern Mo.: Barry, Jasper and Newton counties. ro ya = rid between Lespedeza virginica or Lespedeza Stuevei and Sun capita Lespedeza acuticarpa Mack. & Bush. cky open woods, prairies, and glades. Circumneutral. Southern and west- ern Mo.: Iron, Shannon, Barry, and Jackson counties. Probably a hybrid between Lespedeza virginica or Lespedeza frutescens and Lespedeza violacea. Lespedeza Nuttallii Darl. ry roeky woods and thiekets. A to "— Southern Mo.: Iron, Phelps, Miller, Wright, and Jasper cou Lespedeza hirta (L.) Hornem. Hairy Bush Clover. ... open woods, thickets, and glades. Oxylophile. Central and southern Mo. south and east of a line drawn from St. Louis, Warren, Morgan, and Cedar venie ds to Barton Co, Lespedeza capitata Michx. Rocky prairies, thickets, and glades. Oxylophile to eireumneutral. General. Lespedeza capitata var. longifolia (DC.) T. & G. ometimes found with the typical form. Scattered: St. Louis, Jackson, and Jasper counties. Lespedeza capitata var, sericea H. € A. In similar situations to the typical form, and in many places more common. Cireumneutral. General, LESPEDEZA STRIATA (Thunb.) H. & A. Japanese Clover. woods and roadsides; angen, in rocky or gravelly ground. Oxy- lophile. Eastern, southern, and cen LESPEDEZA STIPULACEA Maxim. Ba Clover. Open woods, roadsides, and waste ground. Indifferent. General. Recently introdueed and being planted in many places along — for a ground eover, and likely to spread widely as an introduced speci Stylosanthes biflora (L.) BSP. cky open woods, Mure and glades. Oxylophile to eireumneutral. Cen- tral and southern Mo.: Boone, Lincoln, St. Charles, St. Louis, Jefferson, Frank- lin, Washington, he. Reynolds, Dunklin, Ripley, Oregon, "e Dent, Maries, Texas, Howell, Laclede, Wright, Taney, Stone, Greene, Polk, Hickory, Ozark, Jackson, Vernon, Barton, Jasper, Newton, MeDonald, and Barry counties 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 583 Stylosanthes biflora var. hispidissima (Michx.) Pollard € Ball. Similar situations to the preceding. Oxylophile to cireumneutral. Central and southern Mo.: Mis E Iron, Reynolds, Carter, Oregon, Howell, Douglas, Dent, Maries, Pulaski, en , Henry, Laelede, Dallas, Hiekory, Greene, Stone, Barry, and McDonald coun VICIA SATIVA L. Spring Vetch. Waste ground and roadsides. Escaped from cultivation. Indifferent. Seat- tered: St. Louis, Jefferson, and Jackson counties. VICIA VILLOSA Roth. Winter Vetch. Fields, waste ground, and roadsides. Indifferent. Seattered: Boone, St. Louis, Jefferson, Texas, Howell, Livingston, Clay, and Jasper counties. Vicia ludoviciana Nutt. Rich rocky woods. Cireumneutral Southwestern Mo.: Greene and Mce- Donald counties. Vicia caroliniana Walt. Rich woods and alluvial banks. Oxylophile to eireumneutral. East-central and southern Mo.: Franklin, Washington, Bollinger, Madison, Wayne, Reynolds, Carter, Ripley, Oregon Howell, Texas, Phelps, Pulaski, Dallas, Ma Doug- las, Greene, Christian, Taney, Barry, Newton, and MeDonald coun Vicia americana Muhl. . Open woods and thickets. Circumneutral. Northern and central Mo.: St. Louis, Jefferson, Atchison, Holt, Platte, Jackson, and Bates counties. VICIA ANGUSTIFOLIA (L.) Reichard. Fields and waste ground. Indifferent. Introduced in Boone and Jackson counties. VICIA MICRANTHA Nutt. Thickets and prairies. Circumneutral. Introduced in Jackson Co. VICIA TETRASPERMA (L.) Moench. oadsides and waste ground. Indifferent. Introduced in Jackson Co. Lathyrus venosus Muhl. var. intonsus Butters & St. John, Rh. 19: 158. 1917. Bushy Vetch. Rocky open ods. Circumneutral. East-eentral and southern Mo.: St. Louis, Crawford, Wayne, Carter, Shannon, Howell, and Barry counties. Lathyrus pusillus Ell. Rocky open woods and chert glades. Oxylophile. Southwestern Mo.: Jasper and Newton countie Lathyrus palustris L. See Kh. 13: 50. 1911. Marsh Pea. Open woods and moist banks. Circumneutral. Central Mo., scattered: Boone, Ray, and Jackson counties. Lathyrus palustris var. myrtifolius (Muhl.) Gray. Open woods and moist banks. Oxylophile to eircumneutral. Central and southeastern Mo.: Boone, St. Louis, Jefferson, Scott, and Iron counties. Apios americana Medic. See Rh. 36: 88-89. 1934. Groundnut. A uberosa Moench (G), (B & B cea and borders of streams. clara to oxylophile. General. [Vor. 22 584 ANNALS OF THE MISSOURI BOTANICAL GARDEN Phaseolus polystachyus (L.) BSP. Wild Bean. cky open woods and thickets. Circumneutral. East-central and southern Mo.: St. Louis, Jefferson, Iron, vpn — Phelps, Pulaski, Dallas, Doug- las, (ema, Jasper, and McDonald eo VIGNA SINENSIS (L.) Endl. Cow Pea. Occasionally escaped from cultivation, Jackson Co. — pies helvola (L.) Britton. Ri cky woods and thickets. Cireumneutral. General. edis helvola var. missouriensis (Wats.) Britton. Moist alluvial ground, woods, and thickets. Cireumneutral. General but scattered: Clark, Boone, Gasconade, Lincoln, St. Charles, St. Louis, Jefferson, Mississippi, New Madrid, Pemiscot, Dunklin, Butler, Clay, Jackson, and Dekalb counties, Strophostyles umbellata (Muhl.) Britton. Moist woods and thickets. Cireumneutral to oxylophile. General but scat- tered: Clark, Marion, Boone, St. Louis, Jefferson, Ste. Genevieve, Iron, Dunklin, Butler, Shannon, Phelps, Taney, Clay, and Jasper counties, Strophostyles pauciflora (Benth.) Wats. Open woods, thickets, and prairies. Circumneutral to oxylophile. General. Olitoria mariana L. Turkey Pea, Butterfly Pea Rocky open woods and glades. Oxylophile. Southern Mo.: Ste. Genevieve, Madison, Iron, Wayne, Reynolds, Carter, Shannon, Howell, Douglas, Taney, Stone, Barry, Newton, and MeDonald counties. Amphicarpa bracteata (L.) Fernald, Rh. 35: 276. 1933. Hog Peanut. Amphicarpa monoica (L.) Ell. (G). Falcata comosa (L.) I (B & B). Open woods and thickets. Cireumneutral to oxylophile. General. — comosa (L.) G. Don. See Rh. 35: 276. 1933. mphicarpa Pitcheri T. & G. (G). ye Pitcheri (T. & G.) Ktze. (B & B Moist rich woods - thickets. Olreumientral, General but seattered: Marion, Ralls, Pike, St. Louis, Jefferson, Iron, Reynolds, Shannon , Phelps, Cole, Boone, pum a” taba "Clinton, Platte, Jackson, Cass, vine, Taney, and Jasper counties Galactia regularis (L.) BSP. Milk Pea. Thickets and open woods along streams. —— to oxylophile. South- eron Mo.: Dunklin, Butler, Shannon, and Barry counties, Galactia volubilis (L.) Britton var. mississippiensis Vail. woods, slopes, and glades. dt ile. Eastern and southern Mo., Nic. in the Ozark region: Knox, St. Charles, Jefferson, Ste. Genevieve, Perry, aep Butler, Scott, Bollinger, Ripley, Oregon, Warne, Shannon, Reynolds, Dent, Phelps, Pulaski, Texas, Howell, Douglas, Taney, Barry, Dallas, Jasper, and Ld counties. — tomentosa (L.) H. & A. Dolicholus tomentosus (L.) Vail B). Sandy open ground. Oxylophile. Southeastern Mo.: Dunklin Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 585 Rhynchosia latifolia Nutt. Dolicholus latifolius (Nutt.) Vail (B & B). andy Ber thickets, and rocky open woods along streams. Oxylophile. Re n Mo.: Mississippi, Stoddard, Dunklin, Ripley, Ozark, and MeDonal Fam. OXALIDACEAE Oxalis violacea L. Sheep Sour, Wood Sorrel. Rocky open woods, glades, and rocky prairies. Circumneutral. General. en florida Salisb. var. strigosifolia Wiegand, Rh. 27: 134. 1925. Ozalis filipes Small, in part (G). Open woods and thiekets. Oxylophile. Southeastern Mo., local: Ripley Co. Oxalis europaea Jord. See Rh. 27: 134. 1925. n Sorrel. Oxalis corniculata of Am. auth., not L. ( Xanthozalis corniculata (L.) Small (B& B). Open woods, thickets, fields, and waste ground. Circumneutral. General and common. Oxalis europaea f. pilosella Wiegand, Rh. 27: 135. 1925. imilar situations to the last. Oxylophile. Scattered: St. Louis, Boone, and Jackson countie Oxalis europaea f. villicaulis Wiegand, Rh. 27: 135. 1925. Frequently oceurring with the other forms. St. Louis, Jefferson, and Doug- las counties. Oxalis europaea f. cymosa (Small) Wiegand, Rh. 27: 135. 1925. Oxalis cymosa Small. Xanthozalis cymosa Small (B € B). Rocky open woods, fields, and waste ground. Circumneutral. General. Oxalis europaea var. Bushii (Small) Wiegand, Rh. 27: 135. 1925. Oxalis Bushii Small. Xanthozalis Bushii Small (B € B). Oxalis interior (Small) Fedde. Xanthozalis interior Small (B & B). Open woods and rocky waste ground. Circumneutral. Southern and central efferson, Boone, Jackson, Johnson, Newton, and Jasper counties Oxalis europaea var. Bushii f. subglabrata Wiegand, Rh. 27: 136. 1925. Occasionally found with the typical and other forms. Jackson Co. Oxalis stricta L. Sheep Sorrel, Yellow Wood Sorrel. y open woods, prairies, fields, and glades. Circumneutral. General, Oxalis stricta var. piletocarpa Wiegand, Rh. 27: 123. 1925 Prairies, open thickets, and glades. Circumneutral. General. Oxalis corniculata L. See Rh. 27: 120. 1925. Ozalis repens Thunb. (G). Xanthozalis corniculata (L.) Small, in part (B & B). Fields and waste ground, and frequently about greenhouses and gardens. Indifferent. Scattered: St. Louis, Jefferson, St. Francois, Boone, Jackson, and asper counti [Vor. 22 586 ANNALS OF THE MISSOURI BOTANICAL GARDEN Oxalis corniculata var. viscidula Wiegand, Rh. 27: 121, Commonly oceurring with the typical form. Ouylophiie. St. Louis and Jackson counties, Oxalis pilosa Nutt. var. Wrightii (Gray) Wiegand, Rh. 28: 67. 1926. pen and waste ground. Oxylophile. Introduced in Jackson Co. Fam. GERANIACEAE Geranium maculatum L. Wild Geranium, Cranesbill Rich or rocky open woods, and thickets. Cireumneutral . General. Geranium carolinianum Meadows, prairies, slides and waste ground. Cireumneutral to oxylophile. General, Geranium carolinianum var. confertiflorum Fernald, Rh. 37: 300. 1935. Similar situations to the preeeding. Oxylophile. General. Geranium pusillum Burm. f. Rocky open ground, Cireumneutral. Southwestern Mo., local: MeDonald Co. ERODIUM CICUTARIUM (L.) L'Her. Pink Needle, Storksbill. Open and waste ground. Cireumneutral. Introduced in Boone and Platte counties. Fam. LrNACEAE Linum medium (Planch.) Britton. Prairies, rocky woods, and glades. Bir rati Central and southern Mo.: Boone, Callaway, St. Louis, Jefferson, Franklin, Washington, Ste. Genevieve, St. Francois, Madison, Iron, Reynolds, Ripley, pA egon, Shannon, Dent, Phelps, Camden, Wright, Polk, Hickory, Cedar, St. Clair, Greene, Ozark, Taney, Barry, Vernon, Barton, Jasper, Newton, and McDonald eountie Linum rigidum Pursh Rocky and waste ground. Cireumneutral. Introduced in Jackson Co. Linum sulcatum Riddell. Rocky prairies, glades, and open woods. Circumneutral, General. Linum striatum Walt. Moist woods, bogs, and wet sandy open ground. Oxylophile to eireumneutral. Eastern Mo., south of the Missouri River; Franklin, Jefferson, Ste. Genevieve, Perry, Iron, Bollinger, Scott, Dunklin, Butler, and Ripley counties. LINUM USITATISSIMUM L. Flax, Linseed. Cultivated and frequently adventive along railroads and roadsides. Indif- erent. Seattered: Boone, Warren, St. Louis, Franklin, Saline, Clay, Jaekson, Greene, Barton, and Jasper counties. Fam. ZYGOPHYLLACEAE TRIBULUS TERRESTRIS L. Caltrop. aste ground. Indifferent. Scattered: St. Louis, Franklin, Boone, Moni- teau, Atehison, Andrew, and Jaekson counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 587 Kallstroemia intermedia Rydb. Kallstroemia maxima (L.) T. & G. (G). Waste ground. Oxylophile. Seattered: Jefferson, Jackson, and Jasper counties. Fam. RUTACEAE Zanthoxylum americanum Mill. Prickly Ash. Thickets and open rocky woods. Circumneutral to calciphile. General, although apparently absent from parts of the Ozark region RuTA GRAVEOLENS L. Rue. Planted in old gardens and occasionally escaped into waste ground. Indif- ferent. Franklin Co Ptelea trifoliata L. Hop Tree, Wafer Ash. Rocky thickets and glades. Cireumneutral to oxylophile. Eastern, central, and southern Mo., and loeally in Atehison Co., in the northwestern part. Com- monest in the Ozark region. Fam. SIMARUBACEAE AILANTHUS ALTISSIMA (Mill) Swingle, Jour. Wash. Acad. Sci. 6: 495. 1916. Tree of Heav poe glandulosa Desf. (G), (B & B). A Chinese tree commonly planted and eseaping in many places. Thickets and along streams. Scattered: St. Louis, Ripley, Jackson, Greene, and Jasper counties. Fam. PoLyGALACEAE Polygala Senega L. Seneca Snakeroot. Rocky open woods, thickets, and clearings. Oxylophile to circumneutral. Seattered: Schuyler, Sullivan, Dent, and Ozark counties. lern ut var. latifolia T. & G. In similar a. to the pes form and more common. Oxylophile to Pint Southern Mo.: n, Reynolds, Carter, Ripley, Oregon, Shan- non, Dent, eun Wright, Wed Douglas, Ozark, Taney, Dallas, Hickory, Polk, Greene, Stone, Barry, and MeDonald counties Polygala incarnata L. Pink Milkwort. Prairies and glades. Circumneutral to oxylophile. Southern and western Mo.: Dent, Texas, Greene, Clinton, Jackson, Cass, Henry, Vernon, Jasper, Newton, and MeDonald countie Polygala sanguinea L. Field Milkwort. rairies, glades, and meadows. Circumneutral. General. Polygala verticillata L. Open woods, mete and glades Cireumneutral to oxylophile. General: Sullivan, Cla de e, St. Charles, St. Louis, Jefferson, Washington, Crawford, Wayne, a rl Dent, Wright, Pri Stone, Polk, p Saline, Atchison, Holt, Jackson, Jasper, Newton, and MeDonald counti [Vor. 22 588 ANNALS OF THE MISSOURI BOTANICAL GARDEN Polygala verticillata var. ambigua (Nutt.) Wood. ocky woods and glades. Oxylophile. Southern Mo.: Washington, Iron, Dent, Shannon, Texas, Howell, Greene, Stone, Barry, and McDonald counties, Polygala verticilata var. sphenostachya Pennell, Bartonia 13: 12, 1932, Limestone glades. Caleiphile. Local: Jackson Co Fam. EUPHORBIACEAE Croton glandulosus L. var. septentrionalis Muell. Arg. Prairies, open woods, eultivated and waste ground. Oxylophile to cireum- neutral. General but commonest in central and southern Mo. Croton capitatus Michx. Hogwort. Prairies, e... and waste ground. Circumneutral to oxylophile. General and commo Croton iens ANA * Dry prairies, bri. "ide and waste ground. Circumneutral to oxylophile. General, except perhaps in parts of northern Mo. Croton Engelmanni Ferg. Open ground. Circumneutral to oxylophile. Southern Mo.: Scott, Missis- sippi, New Madrid, Pemiscot, Stoddard, Butler, Ripley, and Howell counties, and also adventized along railways in St. Louis and Moniteau counties. Croton texensis (Klotzsch) Muell. Arg. Introduced along railroade. Oxylophile. Jackson Co. Crotonopsis elliptica Willd. See Bull. Torr. Bot. Club 45: 478-479. 1919. is lineari (B & B). cky open woods and glades. Oxylophile. East-eentral and southern Mo.: Lincoln, Montgomery, Warren, St. Charles, Gasconade, Franklin, St. Louis, Jef- ferson, St. Francois, Ste. Genevieve, Perry, Bollinger, Madison, Iron, Wa ayne, Khan, Dent, Phelps, Crawford, Pulaski, Texas, Greene, Polk, St. Clair, Barry, Vernon, Jasper, N ae and McDonald counties. Crotonopsis linearis Mic Open sandy mcg "Oxylophie Southeastern Mo.: Stoddard and Dunklin counties. MERCURIALIS ANNUA L. Mercury. Introduced in waste d St. Louis Co. Acalypha virginica L. Three-seeded Mercury. Prairies, thickets, open woods, and int and waste ground; usually in moist or wet ground. Cireumneutral. General. Acalypha digyneia Raf. See Rh. 29: 198. 1927. In similar pipi to the preceding. Cireumn eutral. Eastern, central, and southern Mo.: Marion, St. Charles, St. Louis, Jefferson, St. Francois, Perry, Butler, "Dent, "E Pulaski, Cole, Dallas, Mestqoniebhs Boone, Randolph, Saline, Linn, Clinton, Jaekson, Barton, and Jasper counties, Acalypha gracilens Gray var. monococca Engelm. See 2 = 203. 1927. Acalypha gracilens of auth. in part, not Gray (G), (B Rocky prairies, open woods, and glades. E Eastern, central, and southern Mo., commonest in the Ozark region, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 589 Acalypha ostryaefolia Riddell. Moist ina and eultivated ground. Cireumneutral. East-central and southern Mo.: Gasconade, St. Louis, a“ Perry, Mississippi, Reynolds, Howell, Ou, f x and Jasper cou RICINUS COMMUNIS L. Castor-oil Plant. Cultivated and sometimes adventive in waste ground. Indifferent. Scattered: Lineoln, St. Louis, Dunklin, Boone, Jaekson, and Jasper counties. Tragia urticaefolia Michx. Tragia nepetaefolia of auth., not Torr. (B & B), in part. Tragia ramosa of auth., not Torr. (B € B), in part. Rocky prairies, open ground, and glades. Cireumneutral. Central and south- ern Mo., mostly in the Ozark region. Tragia cordata Michx. ba pci cad d. (G), (B & B). en woods, du and glades. Oxylophile to circumneutral. nl Mo.: Oregon, Stone, Taney, Greene, Barry, and McDonald counties. Phyllanthus caroliniensis Walt. ields, thickets, and moist alluvial open ground. Circumneutral. Central and southern Mo.: Boone, St. Louis, Franklin, St. Francois, Scott, New Madrid, Dunklin, Ripley, scq Shannon, Phelps, Barry, Vernon, Jasper, Newton, and MeDonald co Andrachne ONIS (Nutt.) Muell. Arg. Rocky ledges, glades, and bald knobs. Calciphile to cireumneutral. Southern Mo.: Phelps, Shannon, Texas, Ozark, Christian, Taney, Stone, and Barry counties. Euphorbia zygophylloides Boiss. Prairie Spurge. d rocky open ground. Caleiphile to eireumneutral Western Mo., scattered: Jackson, Cass, Henry, Barry, and Greene counties. Euphorbia serpens HBK. Open ground, E in inm alluvial soil Caleiphile to circumneutral. Northern and central Mo., tered: St. Charles, St. Louis, Perry, Miller, Sa- line, Atehison, ajo nt ers Jaekson, wal Henry counties. Euphorbia glyptosperma Engelm. ase EE bars, and la ys ground. Circumneutral. Northwestern and 1 Mo., scattere t. Louis, Jefferson, Atchison, Holt, Buchanan, Pine ers and Jackson Ba, —— nutans Lag. Nodding Spurge. orbia Preslii Guss. ( mia fields, and waste pee Cireumneutral. General and common. — maculata L. Milk Purslane. Cultivated and waste ground, and in glades. Indifferent. General and Euphorbia humistrata Engelm. Rocky open ground. Oxylophile. Eastern, central, and southern Mo., scat- tered: Marion, Lincoln, Boone, Cooper, Cole, Laclede, St. Louis, Jefferson. Franklin, Perry, Pemiscot, New Madrid, Barry, and Jackson counties. [Vor. 22 590 ANNALS OF THE MISSOURI BOTANICAL GARDEN Euphorbia marginata Pursh. Snow-on-the-mountain. Native on loess hills and plains in northwestern Mo. and introduced in other parts of the state: Atchison, Holt, Jackson, Lafayette, Boone, Greene, Stone, Barton, and Jasper counties. Euphorbia corollata L. Flowering Spurg Prairies, glades, and open roeky sui Cireumneutral. General. Euphorbia dentata Michx. Prairies, glades, and waste ground. Circumneutral. General. Euphorbia heterophylla L. Painted-leaf. Open rocky woods, thickets, glades, and alluvial soils. Circumneutral. General. Euphorbia obtusata Pursh. Warty Spurge. Moist woods and thickets. Cireumneutral. East-eentral and southeastern Mo. b mostly in the eastern Ozark region: Boone, St. Louis, Jefferson, Franklin, St. Franeois, Washington, and Shannon counties. Euphorbia dictyosperma Fisch. & Mey. Tithymalus missouriensis (Norton) Small (B & B), in part. Tithymalus arkansanus (Engelm . & Gray) Kl. & Garcke (B & B), in part. alarrh (8), (R). Galarrhoeus arkansanus (Engelm. € Gray) Small (S), (R). Roeky ledges, glades, and open ground. Caleiphile to eireumneutral. Seat- tered in western and middle Mo.: Boone, Clay, Jackson, St. Clair, Barry, Jasper, and MeDonald eounties, Euphorbia commutata Engelm. Rich rocky woods, and moist open ground. D East-central and southern Mo.: Montgomery, Warren, St. Lou 8, Jefferson, Franklin, Ste. Gene- vieve, St. Francois, Bollinger, Weshiueton, "ahi d, Iron, Reynolds, Wayne, Carter, Ripley, Oregon, Shannon, Texas, Dent, Phelps, tation, Pulaski, Howell Laclede, Wright, Douglas, Taney, Stone, Barry, and MeDonald counties. — CYPARISSIAS L. Cypress Spurge. monly planted in cemeteries and sometimes escaping to roadsides and dada. pr ie Seattered: Marion, is Louis, Boone, and Jackson counties. , Fam. CALLITRICHACEAE Callitriche deflexa A. Br. var. Austini (Engelm.) Hagelm. et or damp open ground, around borders of ponds and swales, fields and waste places, mostly on alluvial soils. Circumneutral to oxylophile. Central and southern Mo.: Boone, Lincoln, St. Louis, Jefferson, Franklin, Ste. Genevieve, Dunklin, Shannon, Phelps, n Texas, Webster, Laelede, Douglas, Greene, Jackson, Henry, Dade, Lawrence, Barton, and Jasper counties. Callitriche heterophylla Pursh. Water Starwort. Ponds, — shallow water of slow streams, and in springs and spring branches. ireumneutral. General but scattered: Livingston, Clark, Ran- dolph, Boone, Os law , St. Louis, Jefferson, Perry, Wayne, Butler, Ripley, Dunklin, Reynolds, ue Iron, Oregon, Howell, Texas, Dent, Crawford, 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 591 Pulaski, Dallas, Laclede, Greene, Ozark, Barry, Wright, Webster, Jackson, Barton, and Jasper counties. Fam. ANACARDIACEAE Rhus glabra L. Smooth Sumach. Upland prairies, thickets, and borders of woods. Circumneutral. General and probably in every county. Rhus copallina L. an thickets, and open woods. Cireumneutral to oxylophile. General, ut commonest in southern Mo Rhus Toxicodendron L. Poison Ivy. Thickets and open woods; commonest on alluvial soil and along streams. Cir- eumneutral General and probably in every county. Very variable in foliage and habit, but the shrubby and seandent forms are not clearly distinguishable. Rhus quercifolia (Michx.) Steud. Poison Oak. Rocky open woods and glades. Oxylophile. Southern Mo., rare and scat- tered: Mississippi and Ozark counties. Rhus canadensis L. Pole-cat Bush, Fragrant Sumac Thickets, glades, and open woods. iO am to ealeiphile. General. Rhus ee var. serotina (Greene) Palmer & Steyermark, comb. nov. Pole- cat Bus at serotina Gree Rhus canadensis v ner of auth., not Rhus trilobata Nutt. (G). aipin glades, and rocky bluffs. Cireumneutral to calciphile. General, but commonest in the Ozark region: Mercer, Putnam, Knox, Shelby, Ralls, Audrain, Boone, St. Francois, Wayne, Butler, Shannon, Texas, Phelps, Pulaski, Laclede, Maries, Cole, Moniteau, Morgan, Miller, Camden, Douglas, Ozark, aney, pp Polk, D al hia dius Henry, St. Clair, Cedar, Barry, Jasper, Newt and Me ou A tont, a "ba AP bd much later than R. canadensis, and after the leaves are half gro It differs from Rhus trilobata Nutt. in its larger, thinner, more obtusely lobed leaflets and larger fruit clusters, as well as in habit of growt Rhus canadensis var. illinoensis (Greene) Fernald. Rocky bluffs and en Calciphile to eireumneutral. Scattered: Putnam and Barry counties Cotinus americanus Nutt. Smoke Tree. Rhus cotinoides Nutt. (G). Rocky bluffs, glades, and bald knobs. Calciphile to eireumneutral. South- western Mo.: Polk, Ozark, Taney, and Stone counties. Fam. AQUIFOLIACEAE Ilex efc Walt. Possum Haw. w woods and glades. Caleiphile to circumneutral. Eastern, central, and ana Mo.: Lewis, Ralls, Pike, Lincoln, Montgomery, St. Charles, St. Louis, (Vor. 22 592 ANNALS OF THE MISSOURI BOTANICAL GARDEN Jefferson, Franklin, Gaseonade, Perry, Cape Girardeau, Bollinger, Scott, Mis- sissippi, New Madrid, Pemiscot, Dunklin, Stoddard, Butler, Wayne, Madison, Iron, Ripley, irse Carter, Shannon, Cole, Miller, Howell, Ozark, Taney, Stone, Barry, St. Clair, Bates, and Jasper counties. Dex verticillata (L.) Gray var. padifolia (Willd.) T. & G. Black Alder, Winter- berry. Sandy bogs and along rocky streams. Oxylophile. Southeastern Mo.: Iron, Madison, and Reynolds counties. Ilex opaca Ait. Christmas Holly. Low wet woods and sandy bogs. Oxylophile. Southeastern Mo.: Cape Girar- deau, Mississippi, New Madrid, Dunklin, Stoddard, Butler, and Wayne counties. Fam. CELASTRACEAE Evonymus atropurpureus Jaeq. Burning Bush, Wahoo Thickets, banks of streams, and open woods. — I" General, Evonymus americanus L. Strawberry Bus Low woods and moist banks of small deta, 2 to eireumneutral. Southeastern Mo.: Dunklin, Butler, and Ripley eoun Evonymus obovatus Nutt. Running Strawberry Bus Shaded rocky bluffs. Cireumneutral to ealeiphile. Ale Mo.: Madison, Shannon, Texas, Howell, and Stone counties. Celastrus scandens L. Bitter-sweet. Thickets and fence rows. Cireumneutral. General, Fam. STAPHYLEACEAE Staphylea trifolia L. Bladder-nut, Rattle-box. Rich woods, thickets, and along bluffs. Circumneutral, General. Fam. ACERACEAE Acer saccharum Marsh. Sugar Maple. Bluffs and woods along streams. Circumneutral. General, but less common in Missouri than the following variety: Daviess, Sullivan, Adair, Saline, Cal- laway, St. Louis, Washington, Ste. Genevieve, Iron, Scott, Butler, Ozark, Gentry, Clay, Jackson, Benton, Cedar, Greene, Barry, and McDonald counties. Acer saccharum f. Rugelii (Pax) Palmer € Steyermark, comb. nov. peel — Pax. accharum var. Rugelíi (Pax) Rehder. iss found with the typical form. Cireumneutral. Eastern and south- ern Mo., scattered: St. Louis, Washington, St. Francois, D Iron, Shan- non, Wayne, Butler, Texas, and MeDonald counties. A f form sometimes looking distinct, but inconstant. Acer saccharum var. glaucum (Pax) Sarg. Bot. Gaz. 67: 233. 1919. In similar situations to the typical form and more common, especially in the Ozark region. Eastern, central, and southern Mo, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 593 Acer saccharum var. Schneckii Rehder in Sarg. Trees € Shrubs 2: 356. 1913. Alluvial woods. Cireumneutral. Southeastern Mo. and locally in central Mo.: Perry, Cape Girardeau, Bollinger, Madison, Iron, Scott, Mississippi, Stoddard, Dunklin, Wayne, and Benton counties. Acer floridanum (Chapm.) P woods. v dem to oxylophile. Southeastern Mo.: Wayne and Dunklin counties. Acer nigrum Marsh. Black Maple. Upland woods and along small streams. Cireumneutral. Northern and central Mo., and occasionally in the southern part of state: Mercer, Grundy, Putnam, Schuyler: Adair, Clark, Knox, Marion, Shelby, Monroe, Randolph, ard, Boone, Cooper, Moniteau, Montgomery, Warren, St. Louis, Washing- ton, Butler, Taney, Holt, Andrew, ‘Cla. ay, Jackson, and Lafayette counties. Acer nigrum var. Palmeri Sarg. Jour. Arnold Arb. 2: 166. 1921. Occasionally found with the typical form: Clark, Lewis, Grundy, and Jackson counties. Acer saccharinum L. Silver Maple, Soft Maple. wet woods and alluvial banks of streams. Circumneutral. General and os in every county. Acer rubrum L. Red Maple. Low wet woods, and rocky hillsides and bluffs in the Ozark region. Circum- neutral Central and southern Mo., south and east of a line from St, Louis, Warren, Boone, and Morgan counties to MeDonald Co. Acer rubrum var. Drummondii (Hook. $ Arn.) T. & G Low wet woods and swamps. Circumneutral. Southeastern Mo.: Perry, Cape Girardeau, Seott, Mississippi, New Madrid, Stoddard, Dunklin, Butler, and Ripley counties. Acer rubrum var. Drummondii f. rotundata Sarg. Bot. Gaz. 67: 237. 1919. Rarely Mai with the preceding form. Southeastern Mo., local: Dunklin and Ripley eounties. Acer rubrum var. tridens Wood. Sometimes found with the typical form. Cireumneutral to oxylophile. East- ern Mo., south of the Missouri River: St. Louis, Jefferson, Franklin, Iron, Stod- dard, and Dunklin counties Acer Negundo L. Box Elder. Low alluvial woods. Cireumneutral. General. Acer Negundo var. texanum Pax Woods along streams, and sow rocky bluffs. Cireumneutral. Southwestern Mo.: Lawrence, Jasper, and McDonald counties. Acer Negundo var. RAMS (Britton) Sarg. Bot. Gaz. 67: 239. Woods along streams. Circumneutral. Northwestern s HER Platte, Clay, and Jackson counties. Acer Negundo var. violaceum Kirch luvi Cireumneutral Northern Mo.: Scotland, Schuyler, Adair, Putnam, Sullivan, Linn, Mercer, Grundy, Livingston, Harrison, — — n. o [=] B [Y t5 Es o iz] nR m i c $ B un (VoL, 22 594 ANNALS OF THE MISSOURI BOTANICAL GARDEN Daviess, Mni m Dekalb, Clinton, Lafayette, Nodaway, Atchison, and uchanan co Fam. SAPINDACEAE Sapindus Drummondi Hook. € Arn. Soapberry, Wild China Tree. Glades, and along the base of bluffs with south or west exposures. Calciphile to circumneutral. Southwestern Mo.: Taney, Stone, Barry, Jasper, Newton, and MeDonald counties. CARDIOSPERMUM HALICACABUM L. Balloon Vin Infrequently eseaped from gardens to picem and waste ground. St. Louis Fam. HiPPOCASTANACEAE ger degit Willd. Ohio Buekeye. alluvial woods along streams. Cireumneutral. Scattered: Putnam, en Pike, Jackson, Barry, and MeDonald counties. The typical form of the species is quite rare in Mo. Aesculus — var. Sargentii Rehder, Jour. Arnold Arb. 7: 241. 1926. us glabra var. arguta Robinson, in part (G). art. Thiekets and glades. Cireumneutral. General but scattered: Lewis, Marion, Pike, Washington, Perry, Bollinger, St. Clair, Jaekson, Cass, Vernon, and Jasper counties, Previously confused with Aesculus arguta Buckley, a southwestern speeies which does not oeeur in Mo Aesculus glabra var. leucodermis Sarg. Trees & Shrubs 2: 262. 1913. eky woods and along small rocky streams. Cireumneutral. Southern and east-central Mo.: St, Louis, Maries, Dallas, Webster, Douglas, Ozark, and Taney counties. Aesculus glabra var. pallida (Willd.) Kirehner. See Jour. Arnold Arb. 2: 118. 1920 n woods, thickets, and low or rocky ground along streams. Circumneutral. General, This is the common variety of buckeye found in the state Aesculus discolor Pursh. Red-flowered Buckeye. Aesculus Pavia of auth., not L. (G), in part (B & B). Low open woods. tiva Southeastern pd Cape Girardeau and Dunklin counties. Aesculus discolor var. mollis pes Trees & Shrubs 2: 267. 1913. Aesculus austrina Small (B & B). Low open woods and SUE ml streams. Southeastern Mo., in the low- lands and locally oret to Phelps Madison, Bollinger, Cape Girardeau, 2: Stoddard, Dunklin, Butler, Wayne, Ripley, Carter, Oregon, and Phelps unties Fam. BALSAMINACEAE Impatiens pallida Nutt. Pale-flowered Touch-me-not. Low wet woods, bogs, and springy banks. Cireumneutral. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 595 Impatiens biflora Walt. Spotted-flowered Touch-me-not. Similar situations to the last. Cireumneutral. General. IMPATIENS BALSAMINA L. Cultivated and sometimes eseaped. Dunklin Co. Fam. RHAMNACEAE Berchemia scandens (Hill) Trelease. Supple-Jack. Low swampy woods, and ledges and slopes of bald knobs and bluffs with seep- age water. Calciphile to cireumneutral. Southern Mo.: Mississippi, New Madrid, Pemiscot, Dunklin, Stoddard, Butler, Ripley, Dent, Shannon, Oregon, Texas, Howell, Douglas, Ozark, Taney, Stone, and Barry counties. Rhamnus lanceolata Pursh. Glades and thiekets. m General exeept in the southeastern lowlands where it is apparently a Rhamnus caroliniana Walt. Indian Cherry. Rocky open woods, thickets, and glades. eo. to cireumneutral. Central and southern Mo.: Boo one, Moniteau, St. Charles, Osage, Maries, Phelps, Gas- D Texas, Howell, Wright, Douglas, Ozark, Taney, Stone, Bab. and Jasper counties. i Rhamnus caroliniana var. mollis Fernald, Rh. 12: 79. 1910. In si Charles, St. Louis, Jefferson, Franklin, St. Francois, Bollinger, Iron, Wayne, Reynolds, Carter, "ab Oregon, "uade Crawford, Gasconade, Ozark, Miller, and Hickory counties RHAMNUS CATHARTICA L. Bue , Thickets and waste P4 jana in Marion and Boone counties. Ceanothus americanus L. New Jersey Tea. Rocky open woods, thickets, and glades. Cireumneutral General but com- monest in southern Mo. Ceanothus ovatus Desf. Red-r woods and glades. ym to oxylophile. Scatterred: Iron, J icu St. ii Jasper, chibi and McDonald counties. Ceanothus ovatus var. pubescens T. & G. ocky prairies, pa a loess hills. Caleiphile to eireumneutral. General. Fam. VITACEAE Parthenocissus quinquefolia (L.) Planch. Virginia Creeper. Psedera c folia (L.) Greene ( pe S, thiekets, fence rows, ind along rocky bluffs. Cireumneutral. General hin hy in every county. Parthenocissus quinquefolia var. hirsuta (Donn) Planch. Psedera quinquefolia var. hirsuta (Donn) Rehder (G). Rocky open woods, thickets, and glades. Cireumneutral General but seat- [Vor. 22 596 ANNALS OF THE MISSOURI BOTANICAL GARDEN tered: Adair, Monroe, Ralls, Boone, Iron, Seott, Ripley, Dallas, Pulaski, Atchi- son, Gentry, Jaekson, Johnson, Jasper, and Newton counties. Parthenocissus quinquefolia var. Saint-Paulii (Koehne € Graebn.) Rehder. sedera quinquefolia var. Saint-Paulii (Koehne & Graebn.) Rehder (G). Rocky ¢ open woods, thickets, and bluffs. Cireumneutral. Southern and w ern Mo.: Mississippi, Ripley, Pulaski, Stone, Jaekson, Cedar, Jasper, Newton, and MeDonald counties. Parthenocissus vitacea (Knerr) Hitche. Psedera vitacea (Knerr) Greene (G). Thickets, open woods, and along bluffs and fence rows. Circumneutral. Gen- eral but steh pede Le Holt, Jackson, Cooper, Christian, Stone, Jasper, and New Puoi cordata mul. Raccoon Grape, False Grape. Cissus Ampelopsis Pers. (G). Open woods, thickets, and fence rows. Circumneutral. General. Ampelopsis arborea (L.) Rusby. Pepper- vine. Ci (G). Low open woods and thickets. Circumneutral. Eastern Mo., south of the Missouri River: St. Louis, Jefferson, St. Francois, Perry, Cape Girardeau, Mississippi, New Madrid, Pemiscot, Dunklin, and Butler counties. Cissus incisa (Nutt.) Des Moulins. Rocky ledges of bluffs with south or west exposure. Calciphile. Southwest- ern Mo.: Taney and McDonald counties. Vitis labrusca L. Northern Fox Grap — ted and occasionally dime into thickets and waste ground. Scat- : St. Louis and Jasper counties. Vitis aestivalis itin Summer Grape. Upland woods and thickets. Circumneutral to oxylophile. General. Vitis Lincecumii Buckley var. glauca Munson. See Bailey, Gent. Herb. 3: 193. 1934. Post Oak Grape. Rocky upland woods, glades, and thickets. Circumneutral to oxylophile, Cen- oddard I e 8 Maries, Laelede, dh , Douglas, Taney, Stone, Barry, Polk, Hickory, Jackson, Cedar, Barry, eke Jasper, Newton, and McDonald counties. This has sometimes been confused with Vitis Lecontiana, or V. bicolor as it was formerly Vitis cinerea Engelm. Winter Grape, Bird Grape. Low alluvial woods, thickets, and fence rows. Circumneutral. General and common. Vitis cinerea var. canescens (Engelm.) Bailey. Occasionally found with the typical form. Scattered: St. Louis, Boone, and Jasper counties. Vitis cordifolia Michx. Frost Grape. Alluvial woods and thiekets along streams. Cireumneutral General and common, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 597 Vitis cordifolia var. foetida Engelm. rowing with the typical form. Local: St. Louis Co. Vitis vulpina L. River-bank Gra Alluvial woods, thiekets, and Pal of streams. Circumneutral. gg and central Mo., and locally south to Madison Co.: Pike, St. Charles, St. Louis, Madison, sy ease Seotland, Knox, Shelby, Moneta, fe rs Callaway, oone, Warren, Cooper, Morgan, Pettis, Schuyler, Adair, Putnam, Sullivan, Linn, Mercer, Grundy, Livingston, Carroll, Harrison, Daviess, Caldwell, Worth, Gentry, Dekalb, Clinton, Atchison, Nodaway, Holt, Andrew, Buchanan, Platte, Jackson, Bates, and Vernon counties Vitis vulpina var. praecox Bailey. similar situations to the typical form. Cireumneutral. Central Mo.: St. Louis, St. Charles, Boone, Carroll, Clay, and Jackson counties. Vitis palmata d Red Grape. Low wet woods and borders of swamps and bayous. Cireumneutral East- central and southeastern Mo.: St. Charles, St. Louis, Jefferson, Bollinger, New Madrid, Pemiseot, Dunklin, and Butler counties. Vitis rupestris Scheele. Sand Grape. Ro anks and beds of streams. I aoe to oxylophile. Central and southern Mo.: Montgomery, Fran efferson, Washington, Madison, Iron, Reynolds, Carter, Oregon, Shannon, Howell, Texas, Phelps, Pulaski, Laclede, Dallas, Miller, Morgan, Stone, m Jasper, Newton, and McDonald counties. Vitis rupestris var. dissecta Egger Roeky ground along streams. pu hile. Eastern Mo., south of the Missouri River: St. Louis, Jefferson, and Carter counties. Vitis rotundifolia Michx. Muscadine. Low open woods and borders of swamps and bayous. Oxylophile. South- eastern Mo.: Madison, Pemiscot, and Dunklin counties. Fam. TILIACEAE Tilia glabra Vent. See Bot. Gaz. 66: 424. 1918. Linn, Linden. Tilia americana of auth., and ais L., in bdo (G), (B & B). Woods and bluffs doug strea Cire eutral. General, but apparently absent nien some of the oiii ond vid counties. Tilia floridana Ashe. Bluffs and rocky woods along streams. Circumneutral. Southern Mo. and along the larger streams to the northern boundary: Clark, Ralls, St. Louis, Ste. Genevieve, Madison, Iron, Wayne, Butler, Carter, Shannon, Douglas, Ozark, Taney, Greene, Stone, Barry, Holt, Jackson, Dade, Newton, and McDonald counties. Tilia floridana var. hypoleuca Sarg. Bot. Gaz. 66: 436. 1918. Rocky woods and bluffs. Cireumneutral. Southern Mo.: Ozark, Taney, Stone, and Barry counties. Sometimes looks very Airline but intergrades with the typical for Tilia heterophylla Vent. var. Michauxii (Nutt.) Sarg. Bot. Gaz. 66: 506. 1918. Tilia Michauxii Nutt. (G), (B& [Vor. 22 598 ANNALS OF THE MISSOURI BOTANICAL GARDEN Rocky woods and bluffs, Cireumneutral. Eastern Mo.: Marion, St. Francois, and Butler counties. Fam. MALVACEAE ABUTILON THEOPHRASTI Medie, Velvet Leaf. Cultivated fields and waste ground. Indifferent. General. Malvastrum angustum Gray. Yellow False Mallow. Rocky prairies and glades. eene to eireumneutral. Central, ~~" and western Mo.: Montgomery, Lincoln, St. I , Jefferson, Franklin ington, Shannon, Ozark, Taney, SK ake Polk, Daila, Tidit, Cass, Henry, St. Clair, and Jasper counties. Sida Elliottii T. & G. Sandy open ground. Oxylophile. Southeastern Mo.: Scott, Stoddard, and Dunklin counties. SIDA SPINOSA L. Cultivated and waste ground. Circumneutral, General. ANODA CRISTATA Schlecht. var. BRACHYANTHA (Reichb.) Hochr. Ann. Conserv. & Jard. Bot. Genève 20: 47. 1916. Introduced in waste ground. McDonald Co. ALTHAEA ROSEA (L.) Cav. Hollyhock. Cultivated in gardens and sometimes escaping to roadsides, fields, and waste round. Boone, Saline, and Greene counties. MALVA ROTUNDIFOLIA L. Common Mallow, Cheeses, Fields, lawns, and waste ground. Indifferent. General. MALVA PARVIFLORA L. Fields and waste ground. Indifferent. Scattered: Atchison and Jackson counties. MALVA MOSCHATA L. Musk Mallo elds and waste ground. Indifferent. Seattered: Boone Co. MALVA SYLVESTRIS L. High Mallow. Roadsides and waste ground. Indifferent. Scattered: Warren, Jackson, Greene, and Stone counties. MALVA CRISPA L. Curled Mallow Waste ground. Indifferent. Seattered: Osage and Jackson counties, and also reported from Greene Co. Callirhoé triangulata (Leavenworth) Gray. Clustered Poppy Mallow. Rocky open woods, prairies, and glades. Cireumneutral. East-eentral and southeastern Mo.: St. Louis, Franklin, Scott, and Mississippi counties. Callirhoé involucrata (T. & G.) Gray. Wine Cups, Purple Poppy Mallow. Upland prairies, fields, and roadsides. Cireumneutral to oxylophile. Scat- tered: Putnam, St. Louis, Linn, Boone, Jackson, and Taney counties. Callirhoé Papaver (Cav.) Gray. Rocky prairies and glades. Calciphile to eireumneutral. Scattered: Pulaski, Pettis, Jackson, and Barry counties, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 599 Callirhoé Bushii Fernald, Rh. 11: 51. 1909, Rocky open woods and glades. Calciphile. Southwestern Mo.: Taney and Barry counties. Callirhoé digitata Nutt. Fringed Poppy Mallow. ocky prairies, meadows, and glades. Circumneutral to oxylophile. Central and southwestern Mo.: Boone, Clay, Polk, Greene, Christian, Taney, Stone, Dade, Lawrence, Barry, Barton, Ja asper, Newton, and McDonald counties. Perhaps introduced in central Mo. Callirhoé alcaeoides (Michx.) Gray. Poppy Mallow Prairies, open woods, and waste ground. eonek Scattered: Shelby, Boone, St. Louis, Dent, Clay, Jackson, Greene, and Jasper counties. Hibiscus lasiocarpos Cav. w open woods, and borders of sloughs, ponds, and bayous. Circumneutral. TAS and southern Mo.: Platte, Lincoln, St. Charles, St. Louis, Jefferson, Franklin, Cape Girardeau, Perry, Scott, Sttddiestpni, New Madrid, Pemiseot, unklin, Butler, Ripley, Wayne, Dent, Shannon, Pulaski, Oregon, Howell, Gas- conade, Jasper, and Newton counties. This has sometimes been confused with ibiscus incanus which is not known to occur in Mo. Hibiscus militaris Cav Prairie sloughs, a borders of ponds and slow streams. Circumneutral. HiBiscus TRIONUM L. Flower-of-an-hou Fields ; TORRES, waste and edi ground. Indifferent. General, but onion d in northern and central Mo HIBISCUS SYRIACUS L. Rose of Sharon, Shrubby Althea. Commonly planted and rarely escaped to thickets and roadsides. Jasper Co. Fam. HYPERICACEAE Ascyrum hypericoides L. St. Andrew's Cross. Rocky open woods. Oxylophile to eireumneutral. East-central and southern Mo., mainly in the Ozark region. Hypericum Ascyron L. Great St. John Moist open woods and thiekets. dio roba to eireumneutral. Northern and central Mo., scattered: Clark, Atchison, and Jackson counties. HYPERICUM PERFORATUM L. St. John’s-wort. Mad —— and waste ground. Cireumneutral to m General but red: Mereer, Clark, Shelby, Marion, Ralls, Boone, St. Louis, Jefferson, ssi "Duas Dent, Shannon, Tiin. Eie. Clay, Jackson, and Greene counties. air — iim Lam pen woods, thickets: and prairie swales. Circumneutral. General. — ji aa Bush Rocky prairies and glade Oxylophile to circumneutral. Southern Mo.: Phelps, enun. Ozark, Barry, Jasper, Newton, and MeDonald eounties. [Vor. 22 600 ANNALS OF THE MISSOURI BOTANICAL GARDEN Hypericum densiflorum Pursh. Sandy open ground and glades. Oxylophile. Southeastern Mo., seattered: Madison and Ripley counties. Hypericum prolificum L. Shrubby St. John 's-wort. oeky ground along streams and bluffs. Cireumneutral to oxylophile. East- ern, middle, and southern Mo., and oeeasional in the western counties. Hypericum cistifolium Lam Rocky prairies, open woods, and glades. Oxylophile to eireumneutral. General. Hypericum mutilum L. Dwarf St. John’s-wort. Open woods, prairie swales, and borders of ponds and ditehes, Cireumneutral. Eastern, central, and southern Mo. Hypericum gymnanthum Engelm. & Gray. Moist, sandy, open ground. Oxylophile. Southern Mo., seattered: Missis- sippi, Shannon, Greene, and MeDonald countie Hypericum gentianoides (L.) BSP. Pine-wee Sterile open ground, prairies, and glades. p A usually on siliceous or granitic rocks. East-central and southern Mo.: Warren, Lincoln, St. Charles, Jefferson, Franklin, Washington, St. Francois, Ste. Genevieve, Perry, Madison, Iron, Dunklin, Ripley, Reynolds, Shannon, Texas, Dent, Crawford, Gasconade, Maries, Pulaski, Greene, Stone, Cedar, Dade, Barton, Jasper, and MeDonald counties. Hypericum Drummondii (Grev. & Hook.) T. € G. Fields, prairie swales, barrens, and open woods. Oxylophile. Eastern, central, and southern Mo.: Knox, Marion, Randolph, Boone, Montgomery, Lincoln Charles, St. Louis, Jefferson, St. Franeois, Ste. Genevieve, Perry, Madison, fik; Wayne, Stoddard, Dunklin, Butler, Franklin, gorii Crawford, Maries, Phelps, Dent, Shannon, Texas, Howell, Ozark, Polk, Gree » Bt. Clair, Dade, Barry, Vernon, Barton, m Newton, and McDonald counti Hypericum lobocarpum Gatti andy open ground. n AN Southeastern Mo.: Dunklin Co. Hypericum petiolatum Walt. Low wet woods, swamps, bayous, and wet pe — sandstone bluffs. lophile to eireumneutral. Southeastern Mo.: Ste. Genevieve, Bollinger, Mis- sissippi, New Madrid, Dunklin, Butler, and Ripley edd 8. Hypericum virginicum L. Low wet woods, swamps, and bayous. Cireumneutral, Southeastern Mo.: Dunklin and Butler counties. , Fam. ELATINACEAE Elatine americana (Pursh) Arn. See Rh. 19: 12. 1917. Waterwort. Ditches and borders of sloughs and swales. Circumneutral. Western Mo., local: Jackson Co. Bergia texana A Seub. d of the Missouri River. Oxylophile to eireumneutral. Western Mo., loeal: Julii Co. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 601 Fam. TAMARICACEAE TAMARIX GALLICA L. Tamarisk Recently introduced along sand bars and banks of the Missouri and Missis- sippi Rivers. Oxylophile to eireumneutral. Northwestern and central Mo.: St. Louis, Warren, Carroll, Jackson, and Holt eounties. Fam. CISTACEAE Helianthemum Bicknellii Fernald, Rh. 21: 36. 1919. Rock Rose. Helianthemum majus Bicknell, not BSP. (G). Crocanthemum majus (L.) Britton (B & B). Rocky prairies and dry open woods. Oxylophile. General: St. Louis, Jef- ferson, Dent, Shannon, Crawford, Oregon, Texas, Howell, Shelby, Putnam, Sullivan, pou Boone, Mercer, Jackson, Greene, Barry, Jasper, and MeDonald eounties. s has mer confused with Helianthemum canadense which is not known to occur in Lechea villosa Ell. Pinweed Rocky open woods and glades. Oxylophile. Eastern, central, and southern Mo.: Knox, Shelby, Boone, Lincoln, Warren, St. Charles, St. Louis, Jefferson, Perry, Iron, Madison, Bollinger, Mississippi, Dunklin, Butler, Ripley, Oregon, Howell, Carter, Shannon, Texas, Reynolds, Dent, Crawford, Phelps, Camden, Polk, E Taney, Barry, Jaekson, Vernon, Jasper, Newton, and MeDonald counties Lechea tenuifolia Michx. Pinweed. Rocky open woods and glades. Oxylophile. General, except in the South- eastern lowlands. Fam. VIOLACEAE Hybanthus concolor (Forster) Spreng. Green Violet. Rich woods and along shaded bluffs. Cireumneutral to calciphile. General. Viola pedata L. Bird-foot Violet, Wild Pansy. cky open woods and prairies. Oxylophile. General, but commonest in east- central and southern Mo. Viola Page L. . lineariloba DC. milar Moli to the typieal form and much eommoner in most parts of pen ion Viola ‚nisse nsis Gre w woods, obti and moist alluvial ground along streams. Circum Bond General but seattered: Clark, Adair, Howard, Boone, Audrain, St. Charles, Gaseonade, Franklin, St. Louis, Jefferson, Ste. Gori, Mississippi, Dunklin, Butler, Wayne, Madison; Ripley, Carter, Moniteau, Dallas, Hickory, Polk, Howell, Greene, Stone, Linn, Grundy, Daviess, Livingston, Platte, Clay, Jackson, Henry, Bates, Jasper, and Newton counties. Viola papilionacea Pursh. See Bull. Vt. Agr. Exp. Sta. 224: 22. 1921. Viola pratincola Greene. Moist meadows, thickets, open woods, and waste or alluvial ground. Circum- [Vor. 22 602 ANNALS OF THE MISSOURI BOTANICAL GARDEN neutral. General. This L— has often been confused with Viola cucullata which is not known in Miss Viola triloba Schwein. Roeky or dry open woods and thiekets. Oxylophile to eireumneutral. Central and southern Mo, north to St. Louis, Warren, Boone, and Jackson counties. This species and the following variety have often bein eonfused with Viola palmata which is not known to oceur in Missouri, Viola triloba var. dilatata (Ell.) Brainerd, Bull. Torr. Bot. Club 37: 587. 1910. imilar situations to the preceding and often found with it. Oxylophile to eireumneutral. Central and southern Mo. south and east of a line drawn from St. Louis, Warren, Boone, Moniteau, and Polk counties to Jasper Co., and locally northwest in Atchison Co., northwestern Mo, Viola Lovelliana Brainerd, Bull. Torr. Bot. Club 37: 526. 1910. Moist rocky ground. Circumneutral. Southern Mo., local: Ozark Co. Viola sororia Willd. pen woods, rocky or dry slopes, and thickets. Oxylophile to eireumneutral. General, Viola sagittata Ait. Upland prairies, open banks, and glades. Oxylophile to cireumneutral. Gen- eral, but seattered: Putnam, Schuyler, Adair, Macon, Scotland, Knox, Shelby, Monroe, Audrain, Marion, Ralls, Pike, Randolph, Boone, Callaway, Franklin Crawford, Washington, Iron, Madison, Carter, Phelps, Moniteau, Greene, Henry, Vernon, Barton, Jasper, Newton, Barry, and McDonald counties. A bet form of this species ( V. subsagittata Greene) has sometimes been confused with Viola fimbriatula which is not known in Missouri. Viola emarginata (Nutt.) LeConte. Prairies, glades, and open banks. Oxylophile to eireumneutral. East-central and southern Mo.: St. Louis, Madison, Iron, Reynolds, Shannon, Oregon, Moni- teau, Jasper, Newton, and MeDonald counties. Viola pedatifida G. Don. Prairie Violet, Prairies, borders of woods, and loess hills. Cireumneutral. Northern, cen- tral, and western Mo.: Putnam, Schuyler, Scotland, Adair, Knox, St. Louis, Daviess, Atehison, Pettis, Cedar, Greene, Barry, Jaekson, Cass, and Jasper counties. Viola viarum Pollard. Moist woods, banks of streams, open ground along roads, and crevices of rocks along streams. Circumneutral. General but scattered: Schuyler, Sul- livan, St. Louis, St. Francois, Crawford, Shannon, Oregon, zark, Pulaski, acids, Texas, Saline, Daviess, Worth, Atchison, Jackson, Henry, Hicko ory, Stone, Barry, and Jasper counties. Viola Bo. L. Narrow-leaved Violet. t sandy prairies, en open ground, and moist hummocks, Oxylophile, "Wt que 0.: Mississippi and Dent counties. Viola er (Banks) nn. Northern White Violet. an streams and moist ledges along bluffs of LaMotte sandstone. Oxy- lophile. educi Mo., loeal: Ste. Genevieve Co. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 603 Viola pubescens Ait. Yellow Vio Rich woods and thickets. pl neutral. Northern and eastern Mo.: Har- rison, Mercer, Putnam, Sullivan, Schuyler, Shelby, Ralls, St. Louis, St. Francois, Btoddard, and Butler eounties. This has frequently been confused with the next followi Viola A Schwein. See Rh. 23: 275. 1922; also Bull. Torr. Bot. Club 38: 194. 1911. Viola scabriuscula Sehwein., in part (G), (B & B). Rich, alluvial or rocky woods. Cireumneutral. General. Viola eriocarpa var. leiocarpa Fernald & Wiegand, Rh. 23: 275. 1922. Viola scabriuscula Sehwein., in part (G), (B & B). Rieh, alluvial or rocky woods. Cireumneutral. General and eommoner than the species. Viola striata Ait. Pale Violet. Moist rich or rocky woods, and alluvial ground along streams. Circum- neutral. Eastern, middle, and southern Mo. south and east of a line drawn from Shelby, Randolph, Boone, and Hickory counties to Barry Co. Viola Rafinesquii Greene. Wild Pansy, Johnny Jump-up. Prairies, glades, roadsides, and waste ground. Indifferent. General and common. VIOLA ARVENSIS Mur Fields, roadsides, and waste ground. Indifferent. Introduced in Johnson 0. Viola papilionacea x pedatifida Brainerd, Bull. Torr. Bot. Club 40: 249, pl. 15. 1913 plis and thickets. Western Mo., seattered: Jackson, Pettis, and Jasper counties. Viola papilionacea x triloba Brainerd, Bull. Torr. Bot. Club 39: 90. 1912, Woods, thickets, and open banks. Jasper Co. Viola pedatifida x sagittata Brainerd, Bull. Torr. Bot. Club 40: 252, pl. 16. 1913. Prairies. Jasper Co Viola a x sororia Brainerd, Bull. Torr. Bot. Club 40: 253, pl. 17. 1913. Loess hills. Atehison Co. Viola sororia x triloba Brainerd, Bull. Torr. Bot. Club 39: 92. 1912. pen woods and thickets. Southern Mo., seattered: Oregon, Greene, and Jasper counties. Viola emarginata x sororia Dowell, Proc. Staten Isl. Assoc. 3: 162. 1912. Prairies, thiekets, and open banks. Jasper Co. Viola sagittata x sororia Brainerd, Bull. Vt. Agr. Exp. Sta. 239: 193. 1924. Thiekets and open banks. Jasper Co. Viola Neue a x sororia Brainerd, Bull. Torr. Bot. Club 37: 178. 1910. oist woods, thiekets, and open ground along roads and banks. any found throughout the state where the parent species occur. Viola missouriensis x sororia. Low open E n banks. Sometimes occurring with the parent species. [Vor. 22 604 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fam. PASSIFLORACEAE Passiflora lutea L. Thickets and low or open rocky woods. Circumneutral. East-central and southern Mo. south and east of a line drawn from St, Charles, Warren, Mont- gomery, Morgan, Hickory, and Cedar counties to Jasper Co. Passiflora incarnata L. Maypops, Passion Flower. Fields, thiekets, low alluvial woods, and along railroads and roadsides. Cir cumneutral to oxylophile. Southern Mo.: Scott, Mississippi, New Madrid, Stoddard, Pemiseot, Dunklin, Butler, Oregon, Greene, Lawrenee, Barry, Jasper, and MeDonald eounties, Probably native in southeastern Mo. and introduced in some of the southwestern counties, Fam. LoASACEAE Mentzelia oligosperma Nutt. Stick Leaf. xposed ledges along bluffs, glades, roeky open soil, and €— embank- ments. Calciphile to eireumneutral. Central and southern Mo.: St. Charles, St. Louis, ones LES Boone, Saline, Jackson, Henry, Deane, Jasper, and McDonald counti Fam. CACTACEAE Opuntia Rafinesquii Engelm. Prickly Pear. oeky prairies, glades, and open woods. Cireumneutral to ealeiphile. South- ern and eentral Mo. south of a line drawn from Ralls, Monroe, Boone, and Saline counties to Caldwell Co. Opuntia macrorhiza Engelm. Rocky ledges and glades. Oxylophile. Southwestern Mo.: Jasper, Newton, McDonald, and Barry counties. Mamillaria missouriensis Sweet var. caespitosa (Engelm.) Wats. Nipple Cactus. Rocky bluffs. Cireumneutral to ealeiphile. Found in Pulaski Co., where perhaps native. Fam. TuvMELAEACEAE Dirca palustris L. Leatherwood, Shaded roeky bluffs, banks of streams, and low alluvial woods. Oxylophile to men Central and southern Mo., and loeally in extreme northern rcer, Callaway, Warren, Perry, RAIN Wayne, Madison, Iron, Reynolds, Shannon, Oregon, Washington, Crawford, Phelps, Maries, Osage, Morgan, Benton, Laclede, Texas, Taney, Stone, and Barry counties. Fam. ELAEAGNACEAE ELAEAGNUS ANGUSTIFOLIA L. Russian Olive, Cultivated and rarely escaped to thickets and waste ground. Pike and St, Louis counties. Fam. LYTHRACEAE Peplis diandra Nutt. Water Purslane. Didiplis diandra (Nutt.) Wood (G), (B & B). 1935] | PALMER € STEYERMARK—PLANTS OF MISSOURI 605 Borders of ponds, sloughs, and ditehes. Circumneutral. General but scat- tered: Livingston, Sullivan, Shelby, Audrain, St. Charles, St. Louis, Jefferson, Butler, Carter, Oregon, Greene, Clay, Jackson, and Jasper counties. Rotala ramosior (L.) Koehne var. interior Fernald € Griscom, Rh. 37: 169. 1935. Tooth-cu Pp. Rotala ramosior of auth. (G), (B & B), in part. Wet prairies, ditches, sloughs, and margins of ponds and streams. Circum- neutral. Central and southern Mo.: pc eae Boone, Lincoln, St. Charles, Gasconade, St. Louis, Jefferson, Perry, Wayne, New Madrid, Butler, Dunklin, on, Madison, Dent, Laclede, Howell, Ozark, Taney, Cedar, Greene, Jackson, Barton, Jasper, Newton, and Me Donald counties. Ammannia coccinea Rottb. Tooth-cup. Ditches, sloughs, and muddy margins of ponds and slow streams. Circum- neutral. General Ammannia auriculata Willd. Similar situations to the preceding. Cireumneutral Western Mo., south of the Missouri River: Jackson, Taney, Jasper, and McDonald counties. Lythrum alatum Pursh. Winged Loosestrife. Wet prairies and alluvial margins of small streams and ponds. Circum- neutral. General. LYTHRUM SALICARIA L. Purple Loosestrife. Introduced in waste ing Franklin Co. Cuphea petiolata (L.) K ne. n woods, ES prairies, and glades. Cireumneutral. General. Fam. MELASTOMACEAE Rhexia er L. var. leiosperma Fernald & Griscom, Rh. 37: 171. 1935. Meadow Beauty Moist M open ground, and borders of bogs. Oxylophile. Southeastern Mo.: Mississippi, Stoddard, Dunklin, Butler, and Ripley counties. Rhexia virginica L. Deer-grass, Meadow Beauty. Moist open ground and borders of bogs. Oxylophile. Southeastern Mo.: St. Francois, Iron, Dunklin, Butler, and Dent counties. "e Interior P nell. es Bush, not Aubl, See Rh. 13: 167. 1911. bri virginica of auth., not L. (B & B), in part. Moist sandy soil along teni streams, Oxylophile. Southwestern Mo.: Greene, Lawrenee, Vernon, Barton, and Jasper counties. Fam. ONAGRACEAE — decurrens (Walt.) DC. Primrose-willow , bayous, and still streams. nom tS Southern Mo.: Wayne, iig Bndseippi, New Madrid, Pemiseot, Dunklin, Butler, and Barry counties. Jussiaea diffusa Forsk. Floating Primrose-willow Sloughs, ponds, and slow streams; in iul water or on mud. Circum- neutral. General. [Vor. 22 606 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ludvigia alternifolia L. Seed-box Prairie swales, low open woods, and borders of streams. Circumneutral. General, Ludvigia polycarpa Short € Peter, False Loosestrife. et open woods and along prairie streams and sloughs. Circumneutral. Gen- eral but seattered: Putnam, Clark, Shelby, Dekalb, Randolph, Boone, St. Louis, Jefferson, Iron, Ripley, Ray, Jackson, and Jasper counties. Ludvigia palustris (L.) Ell. var. americana (DC.) Fernald & Griscom, Rh. 37: 176 Ludvigia palustris of auth. in part (G), (B& B Low wet woods, and borders of slow streams, orm and ditches. Circum- neutral. General. Ludvigia natans Ell. var. typica Fernald & Griscom, Rh. 37: 175. 1935. Border of pond. Southwestern Mo., loeal: Greene Co. Ludvigia glandulosa Walt. Low wet woods, swamps, and bayous. Cireumneutral to oxylophile, South- eastern. Mo.: Bollinger, Stoddard, New Madrid, Pemiseot, Dunklin, Butler, and Ripley counties. Epilobium coloratum Muhl. Willow Herb. Bogs and wet ground along small streams. Cireumneutral. General but scat- tered: Putnam, Boone, St. Louis, Jefferson, Franklin, Ste. Genevieve, Washing- ton, Iron, Shannon, Pat Phelps, Greene, Taney, Stone, Barry, Jackson, Newton, and MeDonald eounties Epilobium densum Raf. Bogs and wet ground along streams. Cireumneutral. Local: Jackson Co. Oenothera rhombipetala Nutt. Evening Primrose. ndy open ground. Oxylophile. Eastern and central Mo., scattered: Clark, Ralls, St. Louis, Phelps, and Jackson counties. Oenothera parviflora L. See Rh. 26: 4. 1924. Oenothera muricata L. (G), (B& B Open woods, thickets, prairies, and waste ground. Circumneutral, General and common. Oenothera I (Rydb.) Mack. & Bush. Oen muricata var. canescens (T. & G.) zen (G). "A prairies, and waste ground. Cireumneutral. General: Putnam, Adair, Jefferson, Phelps, ELE Worth Gentry, ced Grundy, Randolph, Jack- son, Henry, and er counties Oenothera laciniata Hill. Fields, prairies, roadsides, and waste end Cireumneutral. General. Oenothera laciniata var. grandiflora (Wats.) R Occasionally found with the typical don poem Jackson and Jasper counties. Oenothera linifolia Nutt. Sundrops. Fields, prairies, and glades. Oxylophile. Central and southern Mo.: Lincoln, Montgomery, Warren, St. Charles, St. Louis, Jefferson, Franklin, Washington, 1935] PALMER & STEYERMARK-——PLAN TS OF MISSOURI 607 Crawford, St. Franeois, Ste. Genevieve, Perry, Madison, Bollinger, Iron, Wayne, Dunklin, Ripley, Carter, Reynolds, Shannon, Texas, Dent, Oregon, Gasconade, Maries, Phelps, Pulaski, Laclede, St. Clair, Greene, Lawrence, Jackson, Vernon, Barton, Jasper, and Newton counties. Oenothera fruticosa L. See Rh. 20: 51. 1918. Sundrops. Oenothera linearis Michx. (G). Kneiffia linearis (Michx.) Spach (B & B). Rocky and sandy open a UE Southern Mo., seattered: Jeffer- son, Shannon, and Barry cou Oenothera hybrida Michx. See Rh. 24: 177. 1922. Oenothera fruticosa var. hirsuta Nutt. (G). Rocky and sandy open ground. Oxylophile. Western Mo., local: Henry Co. Oenothera pratensis (Small) Robinson. neifia pratensis Small (B & B). Moist alluvial ground. Cireumneutral. Eastern and southern Mo.: Marion, St. Louis, Jefferson, Wayne, Phelps, and Howell counties. Oenothera speciosa Nutt. White Evening Primrose. Hartmannia speciosa (Nutt.) Small (B & B). Limestone glades and prairies. Calciphile. General but seattered: Adair, Boone, St. Louis, Crawford, Dent, Wayne, Butler, Wright, Greene, Jackson, Johnson, pos e St. Mas Vernon, Jasper, and Newton counties. Oenothera trilo ba L cue ciao Spach (B € B). Lois glades and prairies. Calciphile. Southern Mo., seattered: Iron and McDonald counties, and also introduced in Benton and Jasper counties. Oenothera missourensis Sims. Glade Lily, Missouri Primrose. Megapterium missouriense (Sims) Spach (B € B). Glades, bald knobs, and rocky prairies. Caleiphile. Central and southern Mo.: St Pen. Jefferson, Franklin, Washington, Crawford, Ste. Genevieve, Stod- Shannon, Texas, Phelps, Maries, Pulaski, Miller, Laclede, Camden, Dallas, Cedar, Wright, Webster, Greene, Douglas, Ozark, Taney, Stone, visits Lawrence, and Barry counties. The specific name was originally spelled mis sourensis by the author. Oenothera m Nutt. Mer errulata (Nutt.) Walp. (B & B). Jun ps fields, and prairies. Cireumneutral to ealeiphile. Northern and central Mo.: Marion, St. Louis, Boone, Atchison, Holt, and Jackson counties. p biennis L. elds, thickets, and waste ground. Circumneutral. General and common. Gaura biennis L Rocky prairies, thickets, and glades. Circumneutral. General. Gaura filipes Spach. Ro open woods. Circumneutral. Southern Mo., local: Phelps Co. Gaura parviflora Dougl. Prairies, waste ground, and along railroads and roadsides. Circumneutral. Northern, central, and western Mo., seattered: Shelby, Marion, St. Louis, Boone, [VoL. 22 608 ANNALS OF THE MISSOURI BOTANICAL GARDEN Atehison, Holt, Andrew, Platte, Clay, Jackson, Carroll, and Jasper counties. Introduced from farther west except perhaps in northwestern Mo. Gaura parviflora var. lachnocarpa Weatherby, Rh. 27: 14. 1925. Occasionally found with the typical form. Jackson and Jasper counties. Gaura coccinea Pursh, Plains and loess hills. Caleiphile. Northwestern Mo.: Atchison and Holt counties, and also introduced along railroads in Clay and Jackson counties. Stenosiphon linifolius (Nutt.) Britton. Bald knobs and glades. Calciphile. Southern Mo.: Ozark and Taney counties, Circaea latifolia Hill. See Rh. 19: 87. 1917. Enchanter’s Nightshade, Circaea lutetiana of Am. auth., not L. (G), Rich woods and thickets, Cireumneutral. General. Fam. HALORAGIDACEAE MYRIOPHYLLUM PROSERPINACOIDES Gill. Water Feather hallow water and muddy margins of ponds. RT Scattered : Crawford, Texas, and Jasper counties. A Chilean species often cultivated in aquaria and pools. Myriophyllum scabratum Michx. Slow streams, slo "seid gio and ditehes, Circumneutral. General but scat- tered: Seotland, Monro , St. Charles, St. Louis, Butler, Carter, Howell, Jackson, and Jasper nn Myriophyllum heterophyllum Michx. Water Milfoil. Slow streams, sloughs, ponds, and spring branches. Cireumneutral. East- central and southern Mo.: St. Charles, St, Louis, Jefferson, Perry, Wayne, Dunklin, Butler, Ripley, Carter, Oregon, Crawford, Phelps, Pulaski, Texas, Howell, Ozark, Laclede, Dallas, Greene, and E counties. Proserpinaca palustris L. See Rh. 37: 177. 1935. Ponds, sloughs, and slow streams. ED dini. Southeastern Mo.: New Madrid, Dunklin, and Butler counties. Proserpinaca palustris var. amblyogona Fernald, Rh. 11: 120. 1909. In similar situations to the typical form. Cireumneutral. Southeastern Mo.: Butler and Dunklin counties, Proserpinaca palustris var. crebra Fernald & Griscom, Rh. 37: 177. 1935. Swampy woods, ponds, and sloughs. Cireumneutral. Seattered: Livingston, St. Louis, Jefferson, Shannon, Barton, and Jasper counties. "This is the com- monest variety in Missouri. Fam. ARALIACEAE Aralia spinosa L. Tear Blanket, Hercules” Club. Low woods and thickets. Circumneutral. Southeastern Mo.: Cape Girar- deau, Bollinger, Madison, Wayne, Stoddard, Dunklin, Butler, AL Ripley eounties, and also introduced in St. Louis Co 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 609 Aralia racemosa L. American Spikenard. Rich wooded hillsides, and along shaded bluffs. Cireumneutral to oxylophile, iddle Mo.: Sulliv son, Franklin, Ste. Genevieve, Perry, Bollinger, Stoddard, Scott, Wayne, Rey- nolds, Shannon, Carter, Oregon, Howell, Texas, Dent, Phelps, Morgan, Benton, Laclede, Douglas, and Ozark counties. Panax quinquefolium L. Ginseng. Rich woods and along the base of rocky bluffs. Cireumneutral. General but uneommon and being exterminated in ied seetions through persistent digging by natives: Clark, Pike, Boone, St. Louis, Jefferson, Franklin, Bol- linger, Shannon, Jackson, and Greene countie Fam. UMBELLIFERAE Eryngium yuccifolium Miehx. Button Snakeroot, Rattlesnake Master. Roeky open woods, prairies, and glades. Cireumneutral. General. Eryngium prostratum Nutt. Low wet woods, sandy prairies, and borders of swamps and ponds. Circum neutral to oxylophile. Southeastern Mo.: Scott, ^ pa Stoddard, Dunklin, Butler, Ripley, Shannon, Texas, and Howell coun Sanicula gregaria Bicknell. Black Snakeroot. Roeky open woods and thickets. Circumneutral. General. Sanicula canadensis L. Rocky open woods and thickets. Cireummeutral. General. Sanicula Smallii Bicknell. Rich woods. Cireumneutral. Southeastern Mo.: Ripley Co. D bulbosa (Michx.) Nutt. Harbinger-of-sprin ieh woods, alluvial jos and along the base of rocky bluffs. Circum- Nc p ealeiphile. East-central and southern Mo.: Randolph, Boone, Audrain, Callaway, PIE SR Warren, St. qe Jefferson, Franklin, Gas- eonade, Washington, St. Franeois, Ste. Genevieve, Madison, Bollinger, Wayne, Shannon, Phelps, Pulaski, Moniteau, Laclede, Greene, Taney, Douglas, and Jasper counties. Chaerophyllum procumbens (L.) Crantz. Wild Chervil. Rich or ~ bd woods, thickets, glades, and open alluvial ground. Cireum- neutral. Gene Chaerophyllum no. var. Shortii T. & G Chaerophyllum Shortii (T. & G.) Bush (8). Similar situations to the preceding. Seattered: St. Louis, Taney, and Jack- son counties. Chaerophyllum texanum Coult. & Rose. See Rh. 11: 52. 1909. Ro prairies and glades. Cireuibibeu eL Central and southern Mo.: Bol linger, Boone, Taney, Jackson, St. Clair, Greene, Barry, Vernon, dica, and Newton counties. [Vor. 22 610 ANNALS OF THE MISSOURI BOTANICAL GARDEN Chaerophyllum Tainturieri Hook, var. floridanum Coult. & Ros Glades and rocky waste ground. Southern Mo., er Dunklin, Butler, Barry, and Jasper counties. Osmorhiza Claytoni (Michx.) Clarke. Woolly Sweet Cicely. Rich woods and thiekets. Cireumneutral. General. Osmorhiza longistylis (Torr.) DC. Smooth Sweet ND Rich woods and thiekets. Cireumneutral. Gener Osmorhiza longistylis var. villicaulis Fernald. Sometimes found with the IM form. Scattered. Spermolepis patens (Nutt.) Robinso Rocky prairies and glades. a General but seattered: Macon, Boone, St. Louis, Jefferson, Scott, Clay, Jackson, Greene, Taney, Barry, Jasper, and Newton counties, Spermolepis echinata (Nutt.) Heller. Rocky prairies, glades, and sandy or gravelly ground along streams. Oxy- lophile. Bollinger, Seott, Iron, Carter, Ripley, Polk, Jasper, and Newton counties. CONIUM MACULATUM L. Poison Hemlock. Waste ground and along railroads and roadsides. Indifferent. Central and southern Mo., scattered: St. Louis, Washington, Madison, Cole, Saline, Law- rence, Jasper, and Newton eounties. Ptilimnium capillaceum (Michx.) Raf. Mock Bishop's-weed. Swamps and prairie swales. Cireumneutral. East-eentral and southern Mo., scattered: St. Louis, Jefferson, New Madrid, Butler, Ripley, Greene, and Jasper counties. ilimnium Nuttallii (DC.) Britton. Prairie swales, wet meadows, and "pie in glades. en to oxylophile. Central and southern s, Jefferson, Franklin, Iron, Wayne, Bollinger, Mississippi, Dunklin, P Ripley, Pu enl Howell, Greene, Jaekson, Vernon, Barry, Jasper, Newton, and MeDonald counties. Cicuta maculata L. Spotted Cow Borders of ponds, sloughs, ngs prairie streams. Cireumneutral. General. CARUM Carvi L. Caraway. Occasionally escaped from gardens to roadsides and waste ground. Indif- ferent. Scattered: Boone and Jackson counties. Sium suave Walt. See Rh, 17: = 1915. hno Parsley. Sium cicutaefolium J. F. . (G), (B . Swamps and wet a nn. edid. Northern, eentral, and eastern Mo.: Shelby, St. Charles, St. Louis, Mississippi, New Madrid, Dunklin, Ripley, Boone, Livingston, and Jackson counties. Cryptotaenia canadensis (L.) DC. Honewort. oeky woods and ledges along bluffs. Cireumneutral. General. Zizia aurea (L.) Koch. Golden Alexanders, Meadow Parsnip. Rich rocky woods and thickets. Circumneutral to calciphile. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 611 Zizia cordata (Walt.) DC. Rich rocky woods, thickets, and prairies. Cireumneutral. Central and south- ern Mo.: St. Louis, Jefferson, Franklin, Madison, Iron, Reynolds, Carter, Ore- gon, Shannon, Boone, Hickory, Taney, Stone, Barry, Jackson, Jasper, and McDonald counties Taenidia integerrima (L.) Drude. Yellow Pimpernel. Rocky open woods and ledges along bluffs. Caleiphile to eireumneutral. Gen- eral, but commonest in the Ozark region. Eulophus americanus Nut ocky open woods id glades. Caleiphile to eireumneutral. East-central and southern Mo.: Franklin, Jefferson, Washington, Phelps, Shannon, Texas, Greene, Wright, Webster, Taney, Barry, Jasper, and Newton counties. BUPLEURUM ROTUNDIFOLIUM L. Thorough-wax Fields and waste ground. irede, Scattered: Jefferson, Franklin, Washington, and Clay counties. Cynosciadium pinnatum DC. Pools and wet depressions in glades. Oxylophile. Southwestern Mo.: Newton and MeDonald counties. en digitatum D wet woods and swamps. Cireumneutral to oxylophile. Southeastern Mo., e nat Co Ligusticum canadense (L.) Britton. Angelico. Rocky woods and moist ground along streams. Cireumneutral to oxylophile. Southern Mo.: Iron, Bollinger, Ripley, Oregon, Shannon, Texas, Howell, Wright, Webster, Douglas, Stone, and McDonald counties. CORIANDRUM SATIVUM L. Coriander. Occasio m escaped from gardens to roadsides and waste ground. Indif- ferent. Scattered: St. Louis, Boone, and Jackson counties. FOENICULUM VULGARE Hill. Fennel. Sometimes escaping from gardens to roadsides and waste ground. Indifferent. Seattered: St. Louis, Boone, and Jackson eounties. Thaspium barbinode (Michx.) N oeky open woods, and on mm and bluffs of streams. Cireumneutral to oxylophile. Eastern, southern, and central Mo. Thaspium trifoliatum (L.) Gray var. edi Blake, Rh. 20: 53. 1918. Thaspium aureum of auth., not Nutt. (G). Thaspium trifoliatum (L.) Ie in n pax (B & B). Thiekets and rocky prairies. Circumneutral, General. Cogswellia daucifolia (Nutt.) M. E. Jones, Contr. West. Bot. 12: 34. 1908. Lomatium daucifolium (Nutt.) Coult. & Rose (G). Limestone glades. Caleiphile. West-central Mo.: Jackson and Cass counties. Polytaenia Nuttallii DC. Prairie Parsley Rocky prairies and glades. Osleinhile to eireumneutral. General, PASTANICA SATIVA L. Parsnip. caped from eultivation and eommon in field, waste ground, and along road- sides and railroads. Indifferent. General [VoL, 22 612 ANNALS OF THE MISSOURI BOTANICAL GARDEN Heracleum lanatum Michx, Cow Par Moist rich woods and thickets. po ac oiii Seattered: St. Louis, Cass, and Jasper counties. LEVISTICUM OFFICINALE (L.) Koch. Lovage. Escaped from cultivation to waste ground and roadsides. Washington Co. ANETHUM GRAVEOLENS L. Dill. Cultivated and rarely escaped to roadsides and waste ground. Jackson Co. Oxypolis rigidior (L.) Coult. € Rose. Cowbane. Moist shaded bluffs and wet ground along streams. Calciphile to circum- neutral. Eastern, central, and southern Mo.: y, St. Louis, Jefferson, Franklin, Washington, Iron, Bollinger, Reynolds, gd ipley, annon, Dent, Pulaski, Laelede, Dallas, Texas, Polk, Howell, Douglas, Greene, Taney, Barry, Jasper, and Jackson counties. Oxypolis rigidior var. ambigua (Nutt.) Robins In similar situations to the typical form. Onleiphile to eireumneutral. East- ern and southern Mo.: Shelby, Dent, Iron, Shannon, Texas, and Laclede counties, Conioselinum chinense (L.) BSP. Hemloek Parsley. aste ground and along small streams. Indifferent. Southwestern Mo.: Christian, Lawrence, and Jasper counties. Angelica villosa (Walt.) BSP. Wood Angelica. ocky woods and glades. Cireumneutral to oxylophile, Southern Mo.: Craw- ford, Iron, Carter, Shannon, Oregon, Dent, Phelps, Texas, Howell, Ozark, Pulaski, Wright, Webster, Douglas, Greene, Barry, and McDonald counties. TorILIS ANTHRISCUS (L.) Bernh. Hemlock Chervil. oadsides and waste ground. Indifferent. Recently introduced in many places and becoming general: Schuyler, Jefferson, Carter, Oregon, Howell, Boone, Saline, Camden, Pulaski, Phelps, Benton, Dallas, Wright, Douglas, Greene, Clay, Jackson, Cass, Johnson, Cedar, Vernon, Jasper, and McDonald counties Daucus pusillus Michx. Rocky prairies and glade Cireumneutral to oxylophile. Scattered, but commonest in southwestern Mo.: Schuyler, Linn, Benton, Ozark, Taney, Stone, Barry, Jasper, Newton, and MeDonald counties. Daucus CAROTA L. Queen Anne's Lace, Carrot. Fields, waste ground, and along railroads and roadsides. Indifferent. Es- eaped from eultivation. General Fam. CorNACEAE Cornus florida L. Flowering Dogwood ocky open woods, bluffs, and thickets, en to oxylophile. Eastern, central, and southern Mo.; common in the Oza Cornus florida f. rubra (West) Palmer & an comb. nov. Cornus florida var. rubra West. Rarely found with the typical form. Jasper Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 613 Cornus alternifolia L. f. Pigeon Berry, Alternate-leaved Dogwood. Along bluffs and banks of streams. Circumneutral to oxylophile. Eastern and middle Mo., mostly along the larger rivers and locally in the eastern Ozark region: Clark, Lewis, Boone, Callaway, St. Louis, Jefferson, Dent, Shannon, and Texas counties. Cornus obliqua Raf. Kinnikinnik, Swamp Dogwood. Cornus Purpusi Koehne. Cornus Amomum of auth. in part, not Mill. (G), (B & B). Thickets, wet prairies, bogs, and along beds of small streams. Circumneutral. General, and probably in every county. Cornus asperifolia Michx. Rough-leaved Dogwood. hickets and borders of woods; in dry rocky, or sometimes in moist, alluvial ground. Cireumneutral. General, and probably in every count Cornus femina Mill. Stiff Dogwood. Cornus stricta Lam. (G). (B & B). Swamps and low open ground. Cireumneutral. Southeastern Mo.: Cape Girar- deau, Bollinger, Seott, Mississippi, New Madrid, Dunklin, Stoddard, Butler, Ripley, Carter, Reynolds, and Iron eounties. Cornus racemosa Lam. Gray Dogwood. Low woods and thickets, and wet rocky pon along streams. Circumneutral. General but seattered Nyssa sylvatica Marsh. Black Gum. Rocky or alluvial woods along streams and hillsides. Circumneutral to oxy- lophile. Central and southern Mo., south and east of a line drawn from Jefferson, Franklin, Colé, and Benton counties to Newton Co. Nyssa et L. Tupelo. Swamps and low wet woods. Circumneutral. Southeastern Mo.: Bollinger, Cape pridie) Scott, Mississippi, New Madrid, Pemiscot, Dunklin, Stoddard, Butler, and Ripley counties. Fam. ERICACEAE Monotropa uniflora L. Indian Pipe, Ghost La Woods and thickets. Oxylophile to circumneutral. General but seattered: Macon, Boone, Lincoln, St. Louis, Ste. Genevieve, > Franklin, Jefferson, Washing- ton, Crawford, Stoddard, Dunklin, Delle Carter, Shannon, Howell, Ozark, Webst mo Stone, Harrison, Platte, Jackson, Jasper, Newton, and Me- Donald counties. Monotropa S io. L. Pinesap. Rich or roeky woods, and gravelly banks. Oxylophile. East-central and southern Mo., scattered: St. Lou pc ip Ste. Genevieve, Shannon, Greene, Jasper, Newton, and MeDonald co Rhododendron omae T Torr. var. roseum (Lois.) Wiegand, Rh. 26: 4. 1924. Wild Honey Rhododendron mcm Torr., in part (G). [Vor. 22 614 ANNALS OF THE MISSOURI BOTANICAL GARDEN Azalea nudiflora L., in part (B € B). Rocky wooded hillsides and bluffs, Oxylophile: usually on siliceous or granitic rocks. Southern Mo., mainly in the eastern Ozark region: Ste. Gen vieve, Cape Girardeau, Bollinger, Madison, Iron, Reynolds, Wayne, Ripley, Douglas, and Ozark counties, Vaccinium arboreum Marsh. Farkleberry. Rocky woods and bluffs: peared Southern Mo.: Perry, Bollinger, Ore- gon, Gasconade, Shannon, and Howell countie Vaccinium arboreum var. glaucescens bris Sarg. Batodendron glaucescens Greene (8) Batodendron andrachnaef orme Small (8). Rocky woods and bluffs. Oxylophile; usually on siliceous or granitic rocks. East-central and southern Mo.: Lincoln, St. Louis, Jefferson, Ste. Genevieve, St. Francois, Perry, Bollinger, Madison, Iron, Reynolds, Wayne, Butler, Dunklin, Washington, Carter, Ripley, Oregon, Shannon, Dent, Crawford, Gasconade, Phelps, Pulaski, Texas, Howell, Douglas, Ozark, Taney, Stone, Barry, Jasper, d counties. Vaccinium stamineum L. High-bush Huckleberry, Deerberry. Rocky woods and glades. Oxylophile. Southern Mo.: Washington, St. Franeois, Ste. Genevieve, Iron, Madison, Bollinger, Wins, Carter, Ripley, Oregon, Shannon. Texas, Howell, Douglas, Webster, Ozark, Greene, Christian, aney, Stone, Barry, Newton, and MeDonald eounties. Vaccinium stamineum var. neglectum (Small) Deam, Indiana Dept. Conserv. 19 1 (8). Vaccinium neglectum (Small) Fernald (G). Rocky open woods and glades, Oxylophile. Southern Mo.: Ste. Genevieve, Bollinger, Madison, Iron, Reynolds, Wayne, Dunklin, Ripley, Carter, Oregon, hannon, Dent, Texas, Howell, Douglas, Webster, Ozark, Taney, Stone, Cedar, Barry, and McDonald counties, Vaccinium stamineum var. interius (Ashe) Palmer & —— e nov. Polycodium interius Ashe, Jour. Elisha Mitehell Sei. Soc. 46: Nis Rocky open woods. Oxylophile. Southern Mo., scattered: iind and Barry eounties. Vaccinium melanocarpum Mohr. High-bush Huekleberry, Southern Gooseberry. Ro open woods, Oxylophile. Southern Mo.: Wayne, Carter, Barry, and MeDonald counties Vaccinium may Ait. var. tenellum (Ait.) Gra oist sandy or rocky ground. Oxylophile. Edwin Mo.: Ste. Genevieve, Bollinger, Madison, and Butler counties. Vaccinium vacillans Kalm. Low-bush Huckleberry, Low-bush Blueberry. Roeky open woods, glades, and bluffs. Oxylophile. Southern Mo., seattered and mostly in the eastern Ozark region: Jefferson, St. Franeois, alison, Iron, Dunklin, Shannon, and McDonald counties. Vaccinium vacillans var. crinitum Fernald, Rh. 13: 236. 1911. Vaccinium missouriense Ashe. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 615 Rocky open woods, glades, and bluffs. Oxylophile. Central and southern Mo., south and east of a line drawn from Lineoln, Boone, and Henry counties to Jasper Co. This is the eommonest form in Missouri and throughout the Ozark region. Fam. PRIMULACEAE Androsace occidentalis Pursh. Rocky prairies, fields, glades, open woods, and bluffs. ws np to circum- neutral. Eastern, southern, and central Mo., south and east o line drawn from Marion and Boone counties to Jackson Co., and locally is in Daviess Co. Hottonia inflata Ell. Water Violet. wamps, sloughs, and bayous. Cireumneutral to oxylophile. Eastern Mo., south of the Missouri River: St. Louis, Jefferson, Bollinger, Wayne, and Dunklin counties. Samolus parviflorus Raf. Water Pimpernel. Samolus floribundus HBK. (G), (B & B). Low wet woods, iss n spring branches, and wet ledges along bluffs. Cal- ciphile to eireumneutral. East-eentral and southern Mo.: Ralls, Randolph, Boone, Callaway, Mis ery, Warren, St. Louis, Jefferson, Franklin, Ste. Genevieve, St. Franeois, Perry, Bollinger, Madison, Mississippi, Dunklin, Butler, Wayne, Ripley, Carter, Reynolds, Oregon, Dent, Shannon, Howell, Texas, Craw- ford, Phelps, Miller, Morgan, Benton, St. Clair, Hiekory, Polk, Dallas, Laclede, Douglas, Ozark, Taney, Greene, Stone, Barry, Jasper, and MeDonald counties. Lysimachia thyrsiflora L. Tufted Loosestrife. Swamp and bogs. Circumneutral to oxylophile. Western Mo., local: Jack- son : LYSIMACHIA NUMMULARIA L. Moneywort. Low wet woods and boggy open ground. Cireumneutral. Scattered: Marion, Boone, St. Louis, Perry, Madison, Oregon, Gasconade, Platte, Clay, and Jackson counties, Steironema ciliatum (L.) Raf open woods, prairie mid and banks of ponds and streams. Circum- Mero General ue un radicans (Hook.) Tr t woods, and borders of streams and bayous. Circumneutral. Bol- "mw Stoddard, ^ Madrid, Poda Dunklin, Butler, and Ripley counties. Steironema lanceolatum (Walt.) Gray. Wet or rocky dry woods, thickets, en and banks of streams. Oxylophile to cireumneutral. Eastern, southern, and central Mo.: Knox, Shelby, Randolph, Boone, Audrain, Callaway, Montgomery, Lincoln, St. Louis, Jefferson, Franklin, Wagilastod Ste. Genevieve, Bollinger, Madison, Wayne, Iron, Reynolds, Carter, pr iius. Crawford, Dent, d i MU ud s da Webster, Greene, Ozark, Jackson, Cass, Jasper, and MeDonald e "Hu Meum var. ipu E ray. Occasionally found with the typical form. Scattered: Jefferson, Shannon, and Greene counties. [Vor. 22 616 ANNALS OF THE MISSOURI BOTANICAL GARDEN — lanceolatum var. hybridum (Michx.) Gra w open ground. Circumneutral. Seattered: Knox, St. Louis, Mississippi, Mud and Jaekson countie Steironema quadriflorum (Sims) Hitehe. Crosswort, Whorled Loosestrife. Low wet woods and rocky banks of streams. Caleiphile to cireumneutral. East-central and southern Mo.: St. Louis, Jefferson, Washington, Iron, Butler, Ripley, Oregon, Shannon, Dent, Crawford, Phelps, Laelede, Texas, Howell, Douglas, Ozark, Greene, and Taney counties. ANAGALLIS ARVENSIS L. Poor Man's Weatherglass, Scarlet Pimpernel. Fields, glades, and waste ground. Mog ion Central and southern Mo.: St. Louis, Jefferson, Franklin, Washington, te en Madison, Boone, Phelps, Ozark, Miller, Polk, Greene, er onald eountie ANAGALLIS ARVENSIS var. CAERULEA (Sehreb.) Ledeb. Similar situations to the typical form. Seattered: St. Louis, Franklin, Gas- conade, and Miller counties. — minimus L. Chaffweed. ws, prairies, and glades. Cireumneutral. Central and southern Mo., nid: Boone, St. Charles, St. Louis, Jefferson, Crawford, Shannon, Greene, Barry, Jackson, Jasper, and McDonald counties. Dodecatheon Meadia L. Shooting Sta Prairies, glades, bluffs, and ei wool hillsides. Cireumneutral to oxy- lophile. General but uncommon in many localities. Fam. SAPOTACEAE Bumelia lycioides (L.) Pers. Southern Buckthorn. Low woods. Cireumneutral. Southeastern Mo.: Pemiscot and Butler counties, Bumelia lanuginosa (Michx.) Pers. Chittim-wood, Gum-elastic. roeky woods, bluffs, and glades. Cireumneutral. Central and southern Mo., south of a line drawn from Lineoln, Boone, Pettis and St. Clair to Barton Co. Fam. EBENACEAE Diospyros virginiana L. Persimmon. Prairies, borders of woods, and — small streams. Cireumneutral. General but commonest in central and souther 0 Diospyros virginiana f. pumila Palmer & Steyermark, f. Rocky open woods and glades. West-central Mo.: J icons and Bates counties, Frutex 2—4 m. altus vel raro arbor ad 7-8 lta; a typo differt foliis minoribus dense pubescentibus; ramulis iut: "petiolis sepalisque dense pubeseentibu This form differs from the variety pubescens in its usually shrubby habit, in the generally smaller leaves, and the more densely tomentose un foliage, and calyx. It differs ies the variety platycarpa in the ller, mo pubescent leaves, and in the smaller fruit, which is oblong or qut quin and not flattened or depressed. The leaves of this proposed form are oblong or oblong- 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 617 lanceolate, rounded or abruptly pointed at the apex, usually rounded or sub- cordate at the base, ciliate on the margins, dark green and seabrate above when young, but becoming glabrous, permanently densely pubescent on the pale under surface. Pu short stout petioles (8-15 mm. long) are densely pubescent, as is also the ca MISSOURI: E DOIN June 6, 1913, B. F. Bush 7041 (Arn. Arb. TYPE); West- port, June 10, 1900, B. F. Bush 787 (Arn. Arb.); Greenwood, June 5, 1923, B. F. Bush 10092 (Arn. Arb.) ; Greenwood, Sept. 5, 1924, E. J. Palmer 26030 (Arn. Arb.); Monteith Junction, Bates Co., Sept. 10, 1924, E. J. Palmer 26080 (Arn. Arb.); ARKANSAS: Hot dapi defina Co., April 21, Oct. 9, 1925, E. J. Palmer 26840, 29109 (Arn. Arb. Although the American persimmons vary greatly in foliage and fruit, it is difficult to find constant characters on which to separate the different forms. The one here described looks so well marked that it might well be regarded as a distinet species or variety, but since a specimen cultivated at the Arnold Arboretum from seed collected at the type locality does not retain the distinctive shape of the small leaves, it is perhaps best to regard it as an ecological for Diospyros virginiana var. platycarpa Sarg. Jour. Arnold Arb. 2: 168. 1921. Cireumneutral to calciphile. East-central and southern Mo., scattered: St. Louis, Dunklin, Carter, Shannon, Christian, Barton, and Jasper counties. Diospyros virginiana var. pubescens (Pursh) Dippel, Handb. Laubholzk. 1: 306. Diospyros pubescens Pursh Frequently found with the. typical form. Scattered: Marion, Ralls, Boone, Callaway, St. Louis, Wayne, Oregon, Ozark, Johnson, Greene, Stone, and Me- Donald counties. Fam. STYRACACEAE Styrax americana Lam. Smooth Storax. Swamps and low wet woods. Cireumneutral to oxylophile. Southeastern Mo.: Bollinger, Mississippi, New Madrid, Pemiscot, Dunklin, Butler, and Ripley counties Fam. OLEACEAE Fraxinus americana L. White Ash. Upland woods, glades, and borders of streams. Circumneutral. General, and probably in every eounty. Fraxinus americana var. subcoriacea Sarg. Bot. Gaz. 67: 241. 1919. Low woods. Circumneutral. Southeastern Mo.: Dunklin Co. Fraxinus pennsylvanica Marsh. Red Ash. Woods along streams, and borders of lakes and ponds. Circumneutral. North- ern and central Mo., seattered: Scotland, Adair, Marion, St. Louis, Saline, Atch- ison, and Jackson counties Fraxinus pennsylvanica var. lanceolata (Borkh.) Sarg. Green Ash. Low woods, swamps, and borders of streams. Circumneutral. General, and probably in every county. [Vor. 22 618 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fraxinus profunda Bush. Pumpkin Ash. Low wet woods and swamps. Cireumneutral. Southeastern Mo.: Cape Girar- deau, Mississippi, New Madrid, Pemiscot, Dunklin, Stoddard, Butler, and Ripley counties Fraxinus profunda var. Ashei Palmer, Jour. Arnold wie 13: 417-418. 1932. Oceasionally found with the typical form, Dunklin Co Fraxinus quadrangulata Michx. Blue Ash. Bluffs, glades, rocky upland woods, and rarely in alluvial ground along streams, Calciphile to eireumneutral. Eastern, central, and southern Mo. an locally north to Livingston Co.: Marion, Ralls, Pike, Monroe, Randolph, Liv- ingston, Saline, Howard, Cooper, Boone, St, Charles, St. Louis, Franklin, Jeffer- son, Crawford, Washington, Iron, Madison, Bollinger, Mississippi, Butler, Ore- gon, Dent, Shannon, Texas, Phelps, Laelede, Miller, Benton, Hickory, Dallas, Howell, Ozark, Greene, Christian, Taney, Stone, Barry, Jasper, and MeDonald counties. Fraxinus biltmoreana Beadle. Woods and hillsides. Cireumneutral. Southeastern Mo.: Dunklin Co. Forestiera acuminata (Michx.) Poir. Swamp Privet. Swamps, low woods, and borders of streams and bayous. Circumneutral. Eastern and southern Mo., Son along the Mississippi and White River: Marion, Pike, Lincoln, St. Charles, St. Louis, Jefferson, Ste. Genevieve, Perry, Mississippi, New Madrid, Stoddard, Dunklin, Pemiscot, Butler, Taney, Stone, and Jasper counties. Chionanthus virginica L. Old Man’s Beard, Fringe Tree. ow open woods and along bluffs. Cireumneutral to oxylophile. Southern Mo., scattered and rare: Mississippi and Ozark counties, Fam. LOGANIACEAE Spigelia marilandica L. Pink-root. ow open woods and moist thiekets. Cireumneutral to oxylophile. South- eastern Mo.: St. Francois, Cape Girardeau, Dunklin, Butler, Ripley, and Oregon counties. Polypremum procumbens L. Sandy open ground. Oxylophile. Southeastern Mo.: Mississippi and Dunklin counties. Fam. GENTIANACEAE Sabatia campestris Nutt. Prairie Pink. Prairies, ena and glades. Cireumneutral to ealeiphile. Central and south- ern Mo., scattered: Boone, Montgomery Dent, Shannon, Greene, Clay, Jackson, Wais, orien and Newton counties, Sabatia angularis (L.) Pursh. Rose Pink. Rocky open woods, thickets, and glades. Cireumneutral to ealeiphile, East- ern, central, and southern Mo.: Shelby, Randolph, Boone, Callaway, Mont- gomery, Lincoln, St. Charles, St. Louis, Jefferson, Franklin, Gasconade, Ste. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 619 Genevieve, Perry, Bollinger, Stoddard, New Madrid, Dunklin, Butler, Ripley, Oregon, Shannon, Howell, Texas, Phelps, Pulaski, Laclede, Camden, Miller, Cole, Polk, Greene, Taney, Stone, Barry, Vernon, Jasper, Newton, and MeDonald counties. Centaurium texense (Griseb.) Fernald. Centaury. Glades and bald knobs. Calciphile. ea and southwestern Mo.: Miller, Greene, Christian, Taney, and Barry e ies. Centaurium calycosum (Buckley) Fernald. Introduced along railways. Jackson Co. Gentiana quinquefolia L. var. occidentalis en Hitehe. Ague Wee oeky wooded banks and bluffs of stre Oxylophile to eireumneutral. Eastern Mo.: Clark, Adair, Jefferao , Ste. pu enevieve, Iron, Reynolds, Carter, and Shannon counties. This has de confused with the typical form which is not known to occur in Mo. Gentiana puberula Michx. Purple Gentia Prairies and glades. Cireumneutral to poem General, but not common: Sehuyler, Adair, Maeon, Randolph, Shelby, Marion, Monroe, Ralls, Pike, Audrai i ster, Greene, Taney, Lawrence, Barry, D Misc Saline, Randolph, Jackets, Cass. Hickory, Cedar, Jasper, and eben ld counties. Gentiana Andrewsii Griseb. Closed Gentian. Wet open woods, and borders of ponds and streams. Cireumneutral. Scat- tered: St. Louis, Ste. Genevieve, Dent, Boone, Jackson, and Jasper counties. Gentiana clausa Raf. See Rh. 19: 149. 1917. Closed Gentian. Prairie swales, moist banks, and low ground along streams. Cireumneutral. General but scattered: Schuyler, Adair, Macon, Osage, Iron, Butler, Oregon, Texas, Douglas, Saline, Jackson, Cass, and Polk counties. This has sometimes been eonfused with the last ana and with Gentiana Saponaria. Gentiana flavida Raf. Pale Gen Roeky prairies, thiekets, and id Olreumneutral. General but seattered: Schuyler, Sullivan, Ralls, Warren, St. Louis, Franklin, Scott, Shannon, Texas, Phelps, Maries, za Jackson, Cass, nda, Raney, Stone, "usi Jasper, and Newton counties. Frasera caroliniensis s Kindes Columbo. Ro Oxylophile. Southeastern Mo., mostly in the eastern Ozark region: ario , Jefferson, Ste. Genevieve, St. Penta, Bol- linger, Washington, Madison, Wayne , Iron, and Scott counties NYMPHOIDES PELTATUM (S. P. Gmel.) Britten & Rendle. Floating Heart. Introduced in ponds. Scattered: St. Louis, Iron, and Newton counties. Fam. APOCYNACEAE Amsonia Tabernaemontana Walt. Rich woods and thickets. Circumneutral to oxylophile. East-central and southern Mo.: St. Louis, Jefferson, Franklin, Polk, Greene, Webster, Douglas, aney, Stone, Lawrence, Barry, Jasper, Newton, and McDonald counties. [Vor. 22 620 ANNALS OF THE MISSOURI BOTANICAL GARDEN Amsonia Tabernaemontana var. salicifolia Woodson, Ann. Mo. Bot. Gard. 15: 406. 1928 Alluvial thickets and rich ground along bluffs. Cireumneutral. Scattered: — St. Louis, Jefferson, Dent, Ripley, Greene, Christian, and MeDonald countie ES AA var. Gattingeri Woodson, Ann. Mo. Bot. Gard. 15: Rich open woods along bluffs and streams. Cireumneutral. East-central and southern Mo.: Lincoln, St. Louis, Lawrence, and Jasper counties. Amsonia illustris Woodson, Ann. Mo. Bot. Gard. 16: 407. 1929, Roeky open rdiet and gravel bars of streams. Cireumneutral. Central and southern Mo.: Lineoln, St. Louis, Franklin, Jefferson, Crawford, Carter, Ripley, Oregon, Dent, een Osage, a Phelps, Maries, Pulaski, Laclede, Hick- ory, Dallas, Stone, Jasper, Newton, and MeDonald counties. Amsonia ciliata Walt. var. nde. (Raf.) Woodson, Ann. Mo. Bot. Gard. 15: 400. 1928. Glades and bald knobs. Cireumneutral to ealeiphile. Southern Mo.: Douglas and Ozark counties. Trachelospermum difforme (Walt.) Gra Swamps and borders of bayous. Cireumneutral. Boutheastern Mo.: Stod- dard, Dunklin, Butler, and Ripley counties, and also in St. Louis Co. — androsaemifolium L. Pink-flowered Dogbane. Apocynum sylvaticum Greene. Thickets and dry rocky woods. Cireumneutral to ne General but scattered: Mercer, Putnam, Sullivan, Adair, St. Louis, Jefferson, Iron, Shan- non, Howell, Boone, Miller, Morgan, Dallas, Wright, Douglas, Webster, Greene, Lawrenee, Barry, and Jasper counties. Apocynum cannabinum L. Indian Hemp, Dogbane. Rocky prairies, open woods, and glades. Circumneutral. East-central and southern Mo., scattered: St. Louis, Iron, Oregon, Phelps, Stone, and McDonald counties. Apocynum cannabinum var. glaberrimum A. DO. Rocky open woods, thickets, and waste ground. Cireumneutral. General. Apocynum cannabinum var. pubescens (Mitchell) A. DC. ocky open woods, thickets, and waste ground. Circumneutral. General. Apocynum medium Greene. Rocky open woods and thickets. Cireumneutral to oxylophile. General. Apocynum medium var, leuconeuron (Greene) Woodson, Ann. Mo. Bot. Gard. 17: 112. 1930 Similar situations to the preceding. Cireumneutral to oxylophile. General but scattered. Apocynum hypericifolium Ait. Apocynum cannabinum var. hypericifolium (Ait.) Gray (G). Rocky stream beds and alluvial soil. Cireumneutral to calciphile. Northern and central Mo., and loeally south in Texas Co.: Mercer, Boone, St. Charles, St. Louis, Andrew, Carroll, Jackson, and Texas counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 621 Apocynum hypericifolium var. cordigerum (Greene) Beg. € Bel. Atti R. Accad. Lincei, V. 9: 114. 1913 In similar situations to the typical form. Circumneutral to ealeiphile. North- rn and centra o., scattered: Scotland, Dekalb, Livingston, Macon, and Jackson counties. VINCA MINOR L. Periwinkle. Cultivated and occasionally escaped. Indifferent. Scattered: Marion, St. Charles, St. Louis, Jefferson, Scott, Texas, and Clay counties. Fam. ASCLEPIADACEAE Asclepiodora viridis (Walt.) Gra ocky prairies and glades. Caleta to eireumneutral. Central and southern Mo.: St. Louis, Jefferson, Washington, St. Francois, Ste. Genevieve, Bollinger, Butler, Ripley, Oregon, Carter, Shannon, Phelps, Miller, Douglas, Wright, Ozark, Taney, Stone, Barry, Vernon, Barton, Jasper, dino E and MeDonald countie Asclepias tuberosa L. Butterfly-weed. Rocky open woods, prairies, and po Cireumneutral. General. Asclepias tuberosa f. lutea Clute, Am. Bot. 18: 73. Occasionally found with the typical form. J us Co. Asclepias purpurascens L. Purple Milkweed. Rocky open woods, thickets, and glades. Cireumneutral. General This has sometimes been confused with Asclepias exaltata (A. phytolaccoides) which is not known to occur in Mo. Asclepias incarnata L. Swamp Milkweed. Low wet woods, and borders of ponds, sloughs, and prairie streams. Circum- neutral. General. Asclepias speciosa Torr. Open ground. Cireumneutral. Local: St. Louis Co. Asclepias syriaca L. var. kansana (Vail) Palmer € Steyermark, comb. nov. Asclepias kansana Vail (B & B Asclepias syriaca of auth., in ind (G), (B & B). Open woods, fields, and vano ground. Circumneutral. General and common, Typical Asclepias syriaca with few and short prickles on the seed pods is more eastern and has not been found in Mo. Asclepias Sullivantii Engelm. Prairies vis thiekets. Cireumneutral. Northern and central Mo.: Scotland, ipn Gentry, Linn, Livingston, Chariton, Randolph, Lincoln, St. Charles, St. Louis, Atchison, Dekalb, Jackson, and Henry counties. Asclepias humistriata Walt. Asclepias ee Miehx., not Sm. (G), (B & B). Rocky open woods, prairies, and glades. Circumneutral. Central and southern Mo., seattered: St. Louis, Scott, Mississippi, Shannon, Dent, Jackson, Greene, and Jasper counties. [Vor. 22 622 ANNALS OF THE MISSOURI BOTANICAL GARDEN Asclepias Meadii Torr. Dry open woods and prairies. Cireumneutral. Central and southern Mo., scattered: St. Louis, Iron, and Cass counties. Asclepias variegata L. Sandy open ground and dry roeky woods. Oxylophile to eireumneutral. Southeastern Mo.: Seott, Dunklin, and Ripley counties. — mignon (Torr.) Raf. amesü Torr. elie Introduced in Jackson Co. Sometimes confused with Asclepias humistriata which it resembles. Asclepias quadrifolia Jacq. ocky open woods. Oxylophile to eireumneutral. Eastern, middle, and south- ern Mo.: Sehuyler, Marion, Ralls, Monroe, Pike, Boone, Callaway, Cole, St. Louis, Jefferson, Franklin, St. Franeois, Bollinger, Wayne, Iron, Madison, Rip- ley, Shannon, Texas, Phelps, Maries, Wright, Greene, Lm Taney, Stone, Lawrenee, Barry, Jasper, Newton, and MeDonald counti Asclepias perennis Walt. Low wet woods and borders of bayous, Circumneutral. Southeastern Mo.: Mississippi, New Madrid, Pemiscot, Dunklin, Stoddard, Butler, and Ripley counties, Asclepias verticillata L. Whorled Milkweed. Dry prairies, glades, and rocky bluffs. Circumneutral to oxylophile. General, but commonest in northern and central Mo. Asclepias stenophylla Gray. Acerates angustifolia (Nutt. E Dene. (B & B). Dry prairies, Tom and roc uffs. Calciphile to circumneutral. Genera but scattered: Pike, Lincoln, st. Charles, Jefferson, Pulaski, Hickory, Dallas, Ozark, Greene, Stone, Barry, Jackson, and Jasper inco . Acerates hirtella Pennell, Bull. Torr. Bot. Club 46: 184-185. 1919. Acerates floridana of auth., not Asclepias floridana Lam. (G), (B € B). ocky prairies and glades. Circumneutral to oxylophile. General. Acerates viridiflora Ell. Green Milkweed. Roeky prairies and glades. Oli General. Acerates viridiflora var. lanceolata (Ives) Gray. In similar situations to the preceding. Cireumneutral. General but scattered : Boone, Callaway, St. Louis, —— NN inp Howell, Pettis, Cedar, Atehison, Jackson, Jasper, and MeDonald counti Acerates viridiflora var. linearis Gray. Sometimes found with the typical form. Seattered: Ste. Genevieve, Pulaski, Saline, Atchison, Holt, Buchanan, and Greene eounties. Gonolobus laevis Michx. Angle-pod, Sand V Moist alluvial woods, thiekets, and pu lt fields. Circumneutral to oxy- lophile. General. Vincetoxicum gonocarpos Walt. Rocky woods and thiekets along streams. Cireumneutral. Southern Mo.: Mis- sissippi, Dunklin, Ripley, Ozark, Barry, and Jasper counties. 1935] i PALMER € STEYERMARK—PLANTS OF MISSOURI 623 hin cam obliquum (Jacq.) Britto woods and thickets. Cod reri Eastern Mo., south of the Mis- souri i Ber i Rt , Madison, and Sha counties Vincetoxicum rs TS Britton. I ou Milkweed. Roeky open woods, ad and pus Cireumneutral to P Cen- tral and southern Mo.: Warren, St. Louis, Jefferson, Frankl t. Franeois, ington, Ste. Genevieve, "Wa ayne, ond ipley, annon, ends elps, Miller, Howell, Douglas, Wright, Benton, Greene, Barry, Jasper, and Newton counties. Vincetoxicum Baldwinianum (Sweet) Britton. Kia woods and thickets along streams. Circumneutral. Southern Mo.: , Shannon, Taney, Stone, Lawrence, Barry, Jasper, Newton, and en counties. Fam. CONVOLVULACEAE Evolvulus argenteus Pursh. Glades and bald knobs. Caleiphile to cireumneutral. East-central and south- ern Mo.: Montgomery, St. Louis, Jefferson, Franklin, Washington, IT Oregon, Shaanan, Douglas, Ozark, Taney, Greene, Stone, and Barry countie Evolvulus alsinioides L. Rocky open ground. Local: St. Louis Co. Probably introduced. a COCCINEA L. Red Morning Glory. s, banks of streams, and waste ground. Indifferent. Seattered: Monroe, PA Cole, St. Louis, Seott, Reynolds, Greene, Jaekson, and Jasper counties. IPoMoEA QUAMOCLIT L. Cypress Vine. Cultivated and rarely escaped into waste ground. Greene Co. Ipomoea pandurata (L.) G. F. W. Mey. Man-of-the-earth. Fields, REN and waste ground. Cireumneutral. General: St. Louis, Franklin, Jefferson, Morgan, Benton, Greene, and Jasper counties. Ipomoea pandurata var. rubescens Chois. See Rh. 20: 65. 1918. Similar situations to the preceding. Cireumneutral. General. This is the common form in Missouri. Ipomoea lacunosa L. Small White Morning Glory, Bindweed. E iekets, roadsides, waste ground, and alluvial soils along streams and ana Cireumneutral. General: Linn, Boone, Lincoln, St. Charles, St. Louis, a efferson, Madison, Scott, Mississippi, New Madrid, Pemiscot, Osage, Clay, o Tun Cass, Johnson, Vernon, Greene, Jasper, and Newton counties. IPOMOEA PURPUREA (L.) Roth. Common Morning Glory. en fields and waste ground. Indifferent. General, and probably in every co IPOMOEA HEDERACEA Jaeq. Blue Morning Glory. gei fields and waste ground. Indifferent. General, and probably in every coun Convolvulus e L. Dwarf Morning Glory. Rocky open woods and banks. Cireumneutral. Eastern and central Mo.: Clark, St. Louis, Washington, Iron, and Jackson counties. [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN 624 Convolvulus sepium L. Hedge Bindweed Fields, roadsides, and waste ground. Indifferent. General Convolvulus sepium var. pubescens (Gray) Fernald. In similar situations to the preceding. Scattered: Schuyler, Scotland, Marion, Ralls, Monroe, St. Louis, Clay, Jackson, and Jasper counties ek, ush Indifferent. Central and southern Mo.: St. General. Seattered: Schuyler, St. Louis, Waste and cultivated ground Convolvulus sepium var. fraterniflorus Mack Louis, Jefferson, Osage, Jackson, Cass, and Jasper counties L. Small Bindweed. , Open banks, roadsides, and waste ground. Circumneutral Escaped from cultivation CONVOLVULUS ARVENSI CONVOLVULUS ARVENSIS var. OBTUSIFOLIUS Chois. eeasionally found with the Mire form. d Jackson countie CONVOLVULUS JAPONICUS Thunb. one Morning Glory, German Ros Field Washington, Iron, an Roadsides and waste ground about dwellings ed: St. j Cuscuta — Engelm. Smartweed Dodder a obtusifolia HBK. Moist pde along streams and ponds, prairies, and thickets; on species of ral. See Univ. Ill. Biol. Monogr. 6**: 50. 1921. shi Seattered: St. Louis, Platte, Jackson, and Johnson counties Gene on various herbs, especially Compositae. Cuscu Polygonum and other herbs. Cuscuta pentagona Engelm. der Cuscuta arvensis Beyrich (G), (B & General Jackson Co odder. Thickets, prairies, and glades; General but Moi Cuscuta Coryli Engelm. Hazel Dodder. Thickets and open gr seattered: Knox, Montgomery, Bt. Barry, Jasper, and McDonald counti d. r. vulgivaga addo ove vine, Dodder t. nidos Jefferson, Iron, General Cuscuta pentagona var. calycina Enge Occasionally found with the typical vn oist ground, dile ur low woods; on various herbs and shrubs ouis, Gasconade, Jackson, Polk, Stone, Piro — will 121. 1921. beg Gronovii of auth., in part, not wina. (G), (B & B). Moist ground, thickets and prairies; on various herbs and shrubs woods, and borders of prairie streams; on Ambrosia and various 7 B round, on herbs and shrubs, often on hazel. See Univ. Ill. Biol. Monogr Cuscuta Cephalanthi Enge General Cuscuta cuspidata Engelm. Low open other herbs and shrubs. Central and southern Wayne, Dunklin, Greene, Jackson, and Jasper counties Cuscuta — Chois. ground, od swales, and along small streams; on Compositae and other tall "iii Gener 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 625 Cuscuta compacta Juss. Wet ground, low open woods and thickets; usually on shrubs. Eastern and southern Mo., south of the Missouri River: St. Louis, Jefferson, Franklin, Scott, New Madrid, Iron, Reynolds, Carter, Shannon, Howell, and Douglas counties. Fam. POLEMONIACEAE Phlox amplifolia Britton. See Bartonia 15: 15. 1933. Rocky wooded hillsides. Cireumneutral Eastern Mo., south of the Missouri River: St. Louis and Reynolds counties. Phlox paniculata L Moist woods and thickets, and gravel bars and banks along streams. Circum- neutral Central and southern Mo. south and east of a line from St. Louis, Warren, Montgomery, Cole, Camden, and Hiekory to Vernon Co., and also locally north in Marion, Jackson, and Holt counties. Phlox maculata L. var. odorata (Sweet) Wherry, Bartonia 14: 26. 1932. Wild Sweet William. Rich moist woods. Circumneutral. Southeastern Mo., local: Iron Co. Phlox glaberrima L. var. melampyrifolia (Salisb.) Wherry, Bartonia 14: 19. 1932. es and open woods. Cireumneutral. Southeastern Mo.: Bollinger, B Dunklin, Butler, Wayne, Iron, and Ripley counties. Phlox glaberrima var. interior Wherry, Bartonia 14: 19. 1932. Low open ground. Circumneutral. East-eentral Mo.: St. Louis and Jeffer- son counties. Phlox pilosa L. var. virens (Michx.) Wherry, Bartonia 12: 47. 1931. ocky open woods, thickets, and prairies. Cireumneutral to oxylophile. East- ern, central, and southern Mo., commonest in the Ozark region. Phlox pilosa var. fulgida Wherry, Bartonia 12: 47. 1931. Rocky open woods, thickets, and prairies. Cireumneutral to oxylophile. Northern and western Mo.: Lewis, Marion, Ralls, Monroe, Shelby, Randolph, Maeon, Adair, Putnam, Mereer, Daviess, ebur, Atehison, Platte, Jackson, Cass, and Vernon counties Phlox WF var. fulgida f. albiflora (MaeM.) Standley, Rh. 34: 176. 1932. Oeeasionally found with the typical p Scattered: B Adair, arion, Randolph, Macon, and Putnam eounti Phlox pilosa var. ozarkana Wherry, Am. Midl. Nat. 16: 413-416. 1935. po rocky woods. Oxylophile to eireumneutral. Southern and east-central . Louis, ioi Iron, Carter, Wayne, Shannon, Texas, Christian, m RR 5 5 d McDonald eounties Phlox divaricata L. var. Laphamii Wood. Blue Phlox, Wild Sweet William. Rich or rid woods and thiekets. Cireumneutral. General Phlox bifida Beck. hlox pve Gray (G), (B & B), in part. Rocky woods and open banks. Oxylophile to circumneutral. Scattered in outhern Mo., and locally north in "EE Co.: Randolph, Madison, Iron, Reyn- sn Doug an and Taney countie [VoL. 22 626 ANNALS OF THE MISSOURI BOTANICAL GARDEN Phlox Drummondii Hook. Introdueed along railway. Southeastern Mo., local: Dunklin Co. Gilia rubra (L.) Heller. Standing Cypress. Plains and waste ground. Caleiphile to eireumneutral. Cultivated and escaped, except in northeastern Mo., where possibly native. Scattered: Clar Holt, and Greene counties. Collomia linearis Nutt. See Rh. 23: 288, 1921. ilia linearis (Nutt.) Gray (G). Meadows and along bluffs. Circumneutral. Northeastern Mo., local: Marion Co. Polemonium reptans L. Jacob's Ladder, Greek Valerian. Rich alluvial or rocky woods and thickets. Cireumneutral General. Fam. HyproPHYLLACEAE Hydrophyllum canadense L. Waterleaf. Rich woods and moist shaded hillsides. Cireumneutral Eastern Mo.: War- ren, Franklin, St. Louis, Jefferson, and Stoddard counties. Hydrophyllum virginianum L. Rich moist woods and thickets and alluvial ground along streams. Circum- neutral. General. Hydrophyllum appendiculatum Michx. Woollen Breeches. Rich woods and thickets. Cireumneutral to ealeiphile, General but scattered: Schuyler, Adair, Macon, Lewis, Marion, Ralls, Pike, Randolph, Boone, Callaway, Warren, St. Charles, St. Louis, Titans. Franklin, Washington, St. Francois, Ste. Genevieve, Perry, Stoddard, Wayne, Dent, Reynolds, nies, Phelps, Pulaski, Cole, Camden, Laelede, Dallas, Morgan, Greene, Stone, Barry, Grundy, Platte, Clay, Jackson, Lafayette, and Cass counties Nemophila microcalyx (Nutt.) Fisch. & Mey. Moist open woods. Cireumneutral Southeastern Mo.: Dunklin and Butler counties. Ellisia Nyctelea L. oist woods, thickets, alluvial soils, and cultivated or waste ground. Cireum- neutral, General but eg Mercer, je oid Shelby, Marion, Randolph, Boone, Cooper, Cole, Lincoln, St. Louis, Jefferson, Franklin, Wa ne, Shannon, Phelps, Pulaski, Laelede Cs Howell, Saline, "Grands, Daviess, Platte, Clay, Jaekson, Cass, Henry, Polk, Greene, Barry, Jasper, and MeDonald counties. Phacelia neruis Miehx. rocky woods and banks. Cireumneutral. Eastern and middle Mo., south of e Missouri River: ^ Lou uis, Jefferson, Crawford, Shannon, Carter, Phelps, Pulaski, and Texas counties. Phacelia Purshii Buckley. Miami Mist. Moist open woods, alluvial thiekets, and along rocky banks, bluffs, and open places. Cireumneutral to ealeiphile. Eastern and middle Mo. and scattered southwestward, mostly south of the Missouri River: Boone, Callaway, Cole, Montgomery, Warren, Lineoln, St. Louis, Jefferson, Franklin, Washington, St. Francois, Ste. Genevieve, Madison, Bollinger, Iron, Wayne, Reynolds, Carter, 1935] : PALMER € STEYERMARK—PLANTS OF MISSOURI 627 Ripley, Shannon, Gasconade, Maries, Osage, Crawford, Dent, Oregon, Phelps, Pulaski, Ozark, Miller, Camden, Laclede, and Cedar counties, and also introduced in Jackson Co. Phacelia hirsuta Nutt. ocky prairies, glades, ledges along bluffs, low rich woods, and alluvial ground along streams. Calciphile to eireumneutral. Southern Mo.: Shannon, Oregon, Douglas, Wright, Webster, Ozark, Greene, Stone, Lawrence, Barry, Jasper, Newton, and MeDonald counties. This has frequently been confused with Phacelia dubia, an eastern species not known in Missouri. Hydrolea ovata Nutt. Swamps and bayous. Circumneutral to oxylophile. Southeastern Mo.: Dunk- lin and Butler counties. Hydrolea affinis Gray. Swampy woods, sloughs, and bayous. Circumneutral to oxylophile. South- eastern Mo.: Cape Girardeau, Bollinger, Stoddard, New Madrid, Dunklin, But- ler, and Ripley counties. Fam. BORAGINACEAE Heliotropium tenellum (Nutt.) Torr. ky prairies and glades. Calciphile to circumneutral. Eastern, central, and southern Mo., mostly in the Ozark region: Ralls, Montgomery, St. Charles, St. Louis, Jefferson, Franklin, Washington, St. Francois, Ste. Genevieve, Iron Rey- nolds, Ripley, Shannon, Dent, Osage, Maries, Phelps, Pulaski, Camden, Laclede, Wright, Douglas, Ozark, ced Stone, Greene, Polk, Hickory, Dallas, Barry, Jasper, and MeDonald eo HELIOTROPIUM INDICUM L. Indian Heliotrope. Low moist alluvial woods and waste ground. Central and southern Mo.: Lincoln, St. Charles, St. Louis, Jefferson, Perry, Cape Girardeau, Bollinger, Scott, Missi mi^ New Madrid, Pemiseot, Dunklin, Butler, Ripley, Oregon, Shannon, Reynolds, Gasconade, ns Maries, Jackson, Henry, Greene, Taney, Stone, podes and MeDonald counti CYNOGLOSSUM OFFICINALE L. Hound’s s Fields and rocky waste ground. Cireumneutral to oxylophile. Central and southern Mo.: Ralls, Pike, Lincoln, Warren, St. Louis, Jefferson, Washington, St. Francois, Madison, Dent, Carter, Iron, Reynolds, Shannon, Oregon, Boone, Jackson, Greene, and Taney counties. Cynoglossum virginianum L. Wild Comfrey. Rich or rocky woods and alluvial thiekets. Cireumneutral. Central and southern Mo.: Boone, Cole, Warren, St. Louis, Jefferson, Franklin, Washington, i Francois, Ste. Genevieve, Cape Girardeau, Bollinger, Madini, ‘Wises, Iron, ynolds, Dunklin, Ripley, Oregon, Dent, Osage, Greene, Shannon, and Bar we Lappula virginiana (L.) Greene. Beggar’s Liee. Open woods, thiekets, and waste ground. Cireumneutral. General. Lappula Redowskii (Hornem.) Greene var, occidentalis (Wats.) Rydb. Waste ground along railroads. Introduced from farther west. Jackson Co. [Vor. 22 628 ANNALS OF THE MISSOURI BOTANICAL GARDEN LAPPULA ECHINATA Gilib. Stickseed. Waste and cultivated ground. Indifferent. General but scattered: Marion, St. Louis, St. Francois, Iron, Dent, Atchison, Jackson, and Barry counties. Amsinckia lycopsoides Lehm. Introduced along railroads. Jackson Co. SYMPHYTUM OFFICINALE L. Comfrey. Waste ground and along railways. Eseaped from eultivation. Washington and Jackson counties. Myosotis virginica (L.) BSP. Scorpion Grass. Open woods, prairies, and waste ground. Circumneutral. General. Myosotis virginica var. macrosperma (Engelm.) Fernald. In similar situations to the typieal form. Southern Mo., scattered: Dunklin, Shannon, and Jasper counties. MYOSOTIS SCIRPOIDES L. Forget-me-not. haded ground along spring branch. Introduced in St. Louis Co. Mertensia virginica L. Bluebells, Virginia Cowslip. Moist rich woods and slo sometimes on low rocky slopes and bluffs, usually in alluvial soils. Circum ipid to UN General: Knox, Adair, Marion, Pike, Audrain, Callaway, Warren, St. fferson, Franklin cois, Madison, Wayne, Washington, Bollinger, fina, ulaski, Miller, Daviess, Sulli- van, Grun ay, Jackson, Johnson, Laclede, Greene, Christian, Taney, and Stone Moin LITHOSPERMUM ARVENSE L. Corn Gromwell. Fields, waste ground, and along roadsides and railroads. Indifferent. General, Lithospermum latifolium Michx. Rich woods and thickets. Circumneutral. Scattered: St. Louis and Jackson counties. Lithospermum caroliniense (Walt.) MacM. See Rh. 17: 131. 1915. Lithospermum Gmelini (Michx.) Hitehe. (G). Prairies and thickets. Oxylophile. Northern and east-central Mo., scattered: St. Louis, Jefferson, and Nodaway counties. Lithospermum canescens (Michx.) Lehm. Orange Puccoon. Rocky open woods, glades, and prairies. Calciphile to eireumneutral. Gen- eral, but commonest in the Ozark regio Lithospermum angustifolium Michx. Yellow Puccoon. ry open woods, ape and prairies. Caleiphile to circumneutral. General : Cl no Jackson, a. Carroll, Pettis, Cedar, Greene, Stone, Vernon, Jasper, and Newton countie Onosmodium occidentale Mack. False Gromwell. ocky prairies, glades, pesce and loess hills. Cireumneutral. General. Onosmodium occidentale var. sylvestre Mack. Open woods and thickets. dial; Seattered: St. Louis and Jackson counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 629 Onosmodium hispidissimum Mack. Dry rocky prairies, thickets, and glades. Circumneutral. Central and western Mo.: St. Louis, Jefferson, Franklin, Randolph, Boone, Holt, Clay, Jackson, and Jasper counties. Onosmodium hispidissimum var. macrospermum Mack, € Bush. Rich woods and moist prairies. Cireumneutral. Western Mo., local: Jackson Co. Onosmodium subsetosum Mack. & Bush. Dry rocky prairies, thickets, and glades. Circumneutral to calciphile. Central and southern Mo.: Boone, St. ipe Jefferson, Washington, St. Francois, Ste. Genevieve, Dent, Shannon, Tex elps, Miller, Camden, Greene, Christian, Douglas, Ozark, Stone, Barry, = s asper counties. ECHIUM VULGARE L. Blue-wee e ground, gravel E and roadsides. Calciphile. Seattered: Marion, Ralls, x Louis, Jefferson, St. Francois, Reynolds, Shannon, Clay, and Jackson a Fam. VERBENACEAE Verbena urticifolia L. White Vervain. Fields, roadsides, and waste ground. Indifferent. General. Verbena simplex Lehm. See Ann. Mo. Bot. Gard. 20: 282. 1933. Verbena angustifolia Michx., not Mill. (G Prairies, fields, and glades. Caleiphile to circumneutral. Eastern, central, and southern Mo., south and east of a line drawn from Marion and Boone coun- ties to Jackson Co. Verbena hastata L. Blue Verva w open woods, wet prairies, SET and waste ground. Circumneutral. Gen- eral, but commonest in northern and central Mo. Verbena stricta Prairies, dn glades, and waste ground. Circumneutral, General. idees bracteata Lag. & Rodr. See u nr Bot. Gard. 20: 304. 1933. a bracteosa Miehx. (G), (B Fields, roadsides, and waste ground. otl AN General. T bipinnatifida Nut long railroads and in kn ground. Introdueed from farther west. Jackson Co. Verbena canadensis (L.) Britton. Wild Verbena. Prairies, fields, glades, and rocky slopes. Calciphile to eireumneutral. Gen- eral, but commonest in central and southern Mo. Verbena simplex x stricta. Occasionally found with the parent species. St. Louis, Jefferson, Franklin, Phelps, Greene, a and Jasper eounties Verbena bracteata x s Oceasionally is xd the parent species. Jasper Co. [Vor. 22 630 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lippia lanceolata Michx. var. recognita Fernald € Griscom, Rh. 37: 178. 1935. Fog-fruit. Prairie swales, ditches, and borders of ponds and streams. Cireumneutral to ealeiphile. General. me Lippia lanceolata is found along the coastal plain and does not reach Mis Lippia nodiflora (L.) Michx. Low swampy woods. Cireumneutral. Southeastern Mo., local: Dunklin Co. Fam. LABIATAE Teucrium canadense L. var. virginicum (L.) Eat. See Rh. 35: 395. 1933. eucrium canadense of auth., not L. (G), (B € B). Thiekets, fields, and waste ground. Cireumneutral. General. Teucrium occidenta le Pra "sein it Western Mo., loeal: Jackson Co. Perhaps intro- Isanthus brachiatus (L.) BSP. False Pennyroyal. Rocky banks, prairies, and glades. Circumneutral to ealeiphile. General but (— Clark, Macon, Boone, Montgomery, St. Charles, St. Louis, Crawford, ... Bollinger, Reynolds, Shannon, Oregon, Texas, Dent, Ozark, Pulaski, en Mas, Jackson, Barry, Jasper, Newton, and MeDonald counties. Teichostema dichotomum L. Blue Curls. Rocky open woods and glades. Oxylophile. Eastern and middle Mo., mostly in the eastern Ozark region: Montgomery, St. Louis, Jefferson, Ste. Genevieve, St. Franeois, Iron, Wayne, Dunklin, Reynolds, Crawford, Phelps, Shannon, Texas, and Howell counties. Scutellaria lateriflora L. Mad-dog Skullcap. et woods and borders of streams Circumneutral, General but scattered: Marion, Boone, Lincoln, St. Charles, St. Lou rin , St. Francois, Perry, Bollinger, Mississippi, Naw Madrid, Pemiscot, Dunklin, Butle r, Ripley, Carter, Reynolds, Iron, Pulaski, Oregon, Howell, Ache Greene, Taney, Atchison, Pelias, Clay, Jackson, Cass, Jasper, and McDonald counties. Scutellaria lateriflora f. rhodantha ign Rh. 23: 86. 1921. Occasionally found with the common m of the species. Scutellaria ovata Hill. See Contr. U. S. Nat. Herb. 22: 734. 1927, Scutellaria versicolor Nutt. (G). Scutellaria cordifolia Muhl. (B € B). Rocky open woods, bluffs, and glades. Caleiphile to cireumneutral, General. Scutellaria ovata var. pilosior (Benth.) Leonard, Contr. U. S. Nat. Herb. 22: 735. 1927 Beetellaria versicolor var. minor Chapm Roeky bluffs and ledges. Central and debio Mo.: St. Louis, Iron, Maries, Phelps, Pulaski, Laclede, Ozark, Taney, and Barry counties. Scutellaria ovata var. bracteata (Benth.) Blake, Rh. 17: 134. 1915; also U. Nat. E 221: 735. 1927. Rocky open or shaded ground. Circumneutral to ealeiphile. East-central and Berna Mo., seattered: St. Louis and Barry counties. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 631 Scutellaria montana Chapm. See Contr. U. S. Nat. Herb. 22%: 740. Rock en woods. Cireumneutral. Southeastern Mo., local: sa Co. This has Nn confused with Scutellaria serrata, a species not known to occur in Mo. Scutellaria incana Spreng. See Contr. U. S. Nat. Herb. 22%: 743. 1927. Scutellaria canescens Nutt. (G). Rocky pes woods and thickets. Cireumneutral. East-central and southern o.: Ralls, Pike, Lincoln, Warren, St. Louis, Jefferson, St. Francois, 3 Cape wae Iron, Wayne, Bollinger, Butler, Ripley, Oregon Reynolds, kaanon, Texas, Dent, Maries, Phelps, Pulaski, Miler, Howell, ad las, Hickory, Greene, Stone, Jasper, Newton, and McDonald counties. Vice punctata (Chapm.) Leonard. Scutellaria canescens var. punctata Chapm. spi open woods and thickets. Cireumneutral Southern Mo.: Iron, Rey- nolds, Texas, Douglas, Howell, and Ozark counties Scutellaria ovalifolia Pers. See Contr. U. S. Nat. Herb. 22?: 741. 1927. cutellaria pilosa Michx., not Hill (G), (B & B). Roeky woods and bluffs. Cireummeutral to oxylophile. Southern Mo.: Ste. Genevieve, x mii Maie Reynolds, Shannon, Texas, Laclede, Wright, Ozark, Stone, and Bar Scutellaria vcn Zr. See Contr. U. S. Nat. Herb. 22": 723-724. 1927; and Rh. 23: 85. 1921. Scutellaria galericulata of Am. auth., not L. (G), (B & B). w wet woods. Circumneutral. Werten Mo., local: Jackson Co. Scutellaria Bushii Britt es and bald en Caleiphile. Southern Mo.: Carter, Dent, Shannon, Pulaski, Douglas, and Ozark counties. Scutellaria parvula Michx Rocky prairies and glades. Circumneutral. General but commonest in the Ozark region. Scutellaria parvula var. ambigua (Nutt.) Fernald. Frequently found with the typieal form. General but seattered, and com- moner in northern and central Mo. ee nervosa Pursh, oist woods and thiekets. Cireumneutral. Eastern and middle Mo.: someto, helby, Boone, Lincoln, St. Charles, St. Louis, Cape Girardeau, eot, and Dunklin counties. MARRUBIUM VULGARE L. Horehound. Fields, roadsides, and P. ground, especially about old dwellings. Indif- ferent. General and probably in every county. Agastache nepetoides (L.) Ktze. Giant Hyssop. Thiekets and borders of woods. Cireumneutral. General. Agastache scrophulariaefolia (Willd.) Ktze. s, open woods, and alluvial ground. Cireumneutral. Northern and west-central Mo.: Scotland, Mercer, Jackson, and Cass counties. [Vor. 22 632 ANNALS OF THE MISSOURI BOTANICAL GARDEN Agastache scrophulariaefolia var. mollis (Fernald) Helle Similar situations to the preceding. boi -festiered in northern and west-eentral Mo.: Scotland and Jackson counties. NEPETA CATARIA L. tnip Fields, roadsides, aa — ground about old dwellings. Indifferent. Gen- eral and probably in every eounty. NEPETA HEDERACEA (L.) Trevisan, See Rh. 23: 289. 1921. Ground Ivy, Gill-over- the-ground. Fields, waste ground, and open banks. Planted in old gardens and freely escaping. Indifferent. Seattered in eastern Mo.: Marion, Pike, St. Louis counties. The typical form of Nepeta hederacea with larger lovin i is not às common in Missouri as the next following. NEPETA HEDERACEA var, PARVIFLORA Benth. See Rh. 23: 289. 1921. Similar situations to the preceding, and more common. Indifferent. Gen- eral. Dracocephalum parviflorum Nutt. Dragon Head. Along railroads and in waste ground. Eseaped from cultivation. Indifferent. Seattered: St. Louis and Jackson counties, PRUNELLA VULGARIS L. Self-heal, Heal-all. Thickets, fields, and waste ground. Cireumneutral, General. Prunella vulgaris var. lanceolata (Barton) Fernald, Rh. 15: 183. 1 Low woods, thickets, and along streams. In similar situations to the typieal form. General. Prunella vulgaris var. lanceolata f. iodacalyx Fernald, Rh. 15: 184. 1913. Occasionally found with the typical form. Scattered: Ralls, St. Louis, Frank- lin, Stone, and Jasper counties. Physostegia virginiana (L.) Benth. Obedient Plant, False Dragon-head. Moist ground, along prairie streams, and in glades. Circumneutral. General. —— intermedia (Nutt.) Engelm. & Gra wet woods and swamps. Cireuemnentral. Southeastern Mo.: Dunklin, "eed and Ripley counties. Physostegia formosior Lunell, Bull. Leeds Herb. 2: 7. 1908. Rocky open ground. Cireumneutral, Central and southern Mo., seattered: St. Louis, Iron, Jackson, and Cass counties. LAMIUM AMPLEXICAULE L. Henbit, Lawns, meadows, and eultivated or waste ground. Indifferent. General. LAMIUM PURPUREUM L. Dead Nettle. Roadsides and waste ground. Indifferent. Scattered: St. Charles, St. Louis, Saline, Barry, Jasper, and Jackson counties, LEONURUS CARDIACA L. Motherwort. Fields, roadsides, and waste ground about dwellings. Indifferent. General. Stachys tenuifolia Willd. Hedge N i Rich woods, thickets, and roadsides. Circumneutral. General. Stachys aspera Michx. Low moist ground. Circumneutral. Local: St. Louis Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 633 Stachys hispida Pursh. Stachys tenuifolia Willd. var. aspera of auth., not S. aspera Michx, (G). Stachys aspera of auth., not Miehx. (B & B Low alluvial ground. Circumneutral. Korien; central, and eastern Mo.: Perry, Audrain, Ralls, Sullivan, Worth, Clay, and Jackson counties. Stachys palustris L. Thiekets and moist ground along prairie streams. Circumneutral, Northern Mo., loeal: Daviess Co Stachys ambigua Sm tachys palire of auth. in part, not L. Open or moist ground along streams. Circumneutral. General but scattered : Clark, Scotland, Knox, Adair, St. Charles, St. Louis, New Madrid, Pemiscot, Put- nam, Atchison, Holt, us Ray, and Jaekson eounties. Salvia viene L. Cancer-weed. Rieh or roeky woe “thicke ets, and moist open ground. Cireumneutral to oxylophile. Eat. -central and southern Mo., commonest in the eastern Ozark region: St. Louis, Jefferson, Washington, St. Francois, Ste. Genevieve, Perry, Bollinger, Madison, Iron, Wayne, Butler, Dunklin, sg Carter, Oregon, Shannon, Texas, Howell, Douglas, Taney, and Barry counti Salvia azurea Lam. var. grandiflora Benth. Blue Sage. Rocky prairies, n pug and glades. Cireumneutral to calciphile. West- eentral and southwestern Mo.: Jackson, Cass, Johnson, Henry, Greene, Ozark, Taney, Stone, Barry, a Jasper, Newton, and McDonald counties, nn also iode d in St. Louis Co. Salvia lanceaefolia Poi Plains, fields, and ds ground. Indifferent. Perhaps native in northwestern , but introduced in most of the state. General but scattered: Monr uni ain a Warren, St. Charles, St. Louis, Mississippi, Soldat, Ripley, Texas, Greene, bean Caldwell, Atchison, Holt, Andrew, Gentry, Dekalb, Jackson, Johnson, Jasper, and McDonald sialon. SALVIA SCLAREA L. Clear-eye. Cultivated and rarely eseaped to roadsides and waste ground. St. Louis Co. Monarda didyma L. Bee Balm. Moist open woods. Circumneutral. Pike and Clay counties. Probably eseaped from eultivation. Monarda pectinata Nutt. Along bia and in waste ground. Introduced in Jaekson Co. Monarda fistulos Wild Bergamot oeky prairies, ‘thickets, and roads: Cireumneutral to — General but scattered: Schuyler, Scotland, Sullivan, co Marion, Ralls, Randolph, Boone, St. Louis, Jefferson, Ste. Gen e, St. Francois, Du db. Butler, Ripley, Shannon, Maries, Dent, Phelps, Pa. "Howell, San Barry, Buchanan, Jack- son, Barton, Jasper, and MeDonald counties. Monarda fistulosa var. mollis (L.) Benth. is L. (G), (B & B). Roeky prairies, thiekets, and roadsides. Cireumneutral. General. [Vor. 22 634 ANNALS OF THE MISSOURI BOTANICAL GARDEN Monarda Bradburiana Beck. Open rocky woods, thickets, and glades. Cireumneutral to ealeiphile. Eastern, central, and southern Mo., south and east of a line from Marion, Boone, and Benton counties to Jasper Co. Monarda punctata L. Horse Mint. Prairies and open ground. Cireumneutral to oxylophile. Southeastern Mo., seattered: Seott and Dunklin counties. Monarda punctata var. villicaulis Pennell, Bull. Torr. Bot. Club 46: 186. 1919. Sandy open ground. Circumneutral to oxylophile. Eastern Mo., scattered: Clark, Pike, and Dunklin counties. Monarda punctata var. occidentalis (Epling) Palmer & rhe icone ze nov. Monarda punctata subsp. occidentalis Epling, Madrofio 3: 25. Open ground. Cireumneutral. Western Mo., local: Jackson E Monarda citriodora Cerv. Monarda dispersa Small (B & B). Rocky prairies, ps and bald knobs. Caleiphile to eireumneufral. Central and en Mo.: St. Louis, Boone, Jackson, St. Clair, Greene, Taney, Stone, Barry, Jasper, ind MeDonald counties Blephilis hirsuta (Pursh) Benth. Wood Min Rich open woods and thickets. Clremtadttinl; General. Blephilia ciliata (L.) Raf. Ohio Horse Mint. Open woods and thickets. Cireumneutral. General. Hedeoma pulegioides (L.) Pers. Pennyroyal. Rocky open woods, hiela, and glades. Cireumneutral. General. Hedeoma hispida Pursh. Rocky prairies and glades. Cireumneutral to oxylophile. General but seat- y i Reynolds — Wright, Douglas, Webster, 8t Clair, Harrison, Platte, Clay, Jackson, s, Jasper, and Newton counties, HEDEOMA ACINIOIDES Schee Waste ground. In en in Jasper Co. Satureja glabra (Nutt.) Fernald. Savory, Calamint. Rocky open ledges and glades. Caleiphile. Central and southern Mo.: St. Louis, Jefferson, St. Francois, Ste. Genevieve, Washi ington, Iron, Reynolds, Carter, Ripley, Oregon, Shannon, Texas, Phelps, Maries, Pulaski, Miller, Douglas, Wright, Ozark diee i Benton, Hickory, St. Clair, Polk, Grae, Stone, Barry, and MeDonald. coun MELISSA OFFICINALIS L. Lemon Balm. Thickets, Zn. and waste idos Indifferent. Central and southern Mo., seattered: Boone, St. Louis, Jefferson, Greene, Stone, Barry, Jackson, Jasper, Newton, "e MeDonald counties Pycnanthemum Torrei Benth. Dry open ground. Oxylophile. Southeastern Mo.: Dunklin Co. Pycnanthemum flexuosum (Walt.) BSP. Mountain Mint. Rocky open woods and thiekets. Cireumneutral General and common. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 635 Pycnanthemum pilosum Nutt. Prairies, thiekets, and open woods. Cireumneutral. General. Pycnanthemum virginianum (L.) Durand & Jackson. Rocky woods, ereviees of bluffs, and thickets. Cireumneutral to ealeiphile. Central and southern Mo.: Boone, Audrain, St. Louis, Jefferson, St. Fra E Iron, Butler, Carter, Oregon, Shannon, Texas, Dent, Howell, Saline, Jackson Benton, ronis. Hickory, Caney: and Stone counties des image incanum (L.) Michx. open woods. x Southern Mo., seattered: Dunklin, Butler, Nun foe and Barry countie Pycnanthemum albescens T. & G. Rocky open woods. Oxylophile. Southern Mo.: Madison, Ste. Genevieve, Perry, Wayne, Dunklin, Butler, xA Dent, Shannon, Phelps, Pulaski, Texas, Howell, Oui Douglas, Barry, Newton, and McDonald counties, This has often been eonfused with the preceding, es is the commoner of the two species. er muticum (Michx.) Pers. cky open woods. Circumneutral. Southeastern Mo.: Dunklin and Butler Begin Cunila origanoides (L.) Britton. Dittany. Roeky woods. Cireumneutral to oxylophile. Central and southern Mo., south and east of a line drawn from Lincoln, Boone, Morgan, and Hickory counties to Jasper Co. Lycopus virginicus L. Bugle Weed. Low wet woods and open ground. Circumneutral. General. Lycopus rubellus Moench. Water Horehoun Low woods and borders of streams and TN Cireumneutral. Central and southern Mo.: Boone, St. Louis, Jefferson, Mississippi, Dunklin, Butler, Carter, Shannon, Dent, Taxan, Howell, Ozark, Taney, Greene, Barry, Jasper, Newton, and McDonald counties. Lycopus lucidus Turez. var. americanus Gray. ycopus asper Greene (B & B). Low sandy ground along Missouri River. Oxylophile. West-central Mo., local: Jackson Co. Lycopus americanus Muhl. et woods, margins of ponds, and along prairie streams. Circumneutral. General. MENTHA LONGIFOLIA Huds. See Rh. 26: 171-175. 1924. a sylvestr Escaped from cultivation to waste ground and roadsides. Indifferent. Scat- tered: Clay, Jackson, and Jasper counties. MENTHA ALOPECUROIDES Hull. Woolly Mint. e ground, fields, and roadsides. Indifferent. Introduced in Jackson Co. MENTHA CARDIACA Gerarde. Waste ground and borders of springs and brooks. Indifferent. Introduced in Jasper Co. [Vor. 22 636 ANNALS OF THE MISSOURI BOTANICAL GARDEN MENTHA SPICATA L. Spea Was nd and aee especially about dwellings. Indifferent. Gen- eral, but scattered : ls, Boone, Franklin, Jefferson, Dunkli ent, Shannon, Phelps, Howell, Cedar, Gren, Stone, Jasper, and Newton eit MENTHA PIPERITA L. Peppermint. Waste ground and margins of springs or spring branches. Cireumneutral. Pun MENTHA CITRATA Ehrh. Bergamot Mint. Waste vi and along spring branehes. Introduced in Jasper Co. MENTHA GENTILIS L. wisis Indifferent. Introduced in Jackson Co. Min arvensis L. Low ground along streams. Cireummeutral. Southeastern Mo., local: Ripley Co. Mentha arvensis L. var. canadensis (L.) Briq. Corn Mint, Field Mint. Moist ground along small streams and about springs. Circumneutral. Gen- eral, but less common than the next. Typical Mentha arvensis is rare in Mo. Mentha arvensis var. glabrata (Benth.) Fernald. In similar situations to the preceding. Cireummeutral. General and com- mon. This is the commonest variety found in Missouri. Collinsonia canadensis L. Horse Balm. Rich or rocky woods. Circumneutral to oxylophile. Eastern Ozark region: Iron, Wayne, Reynolds, and Shannon counties. PERILLA FRUTESCENS (L.) Britton. Beef-steak Plant. oods, rich or alluvial soil along streams, and waste ground. Circum aui. Eseaped and becoming a troublesome weed along some of the Ozark streams, Eastern, central and southern Mo. north to St. Louis, Montgomery, and Jackson counties, Fam. SOLANACEAE SOLANUM DULCAMARA L. Climbing Nightshade. Thickets and roadsides. Indifferent. Scattered: Adair, Boone, St. Louis, Jefferson, Jackson, and Greene counties Solanum triflorum Nu Waste ground he along railroads. Introduced from farther west. Jack- son Co. Solanum nigrum L. Black Nightshade. Waste and eultivated ground. Indifferent. General and eommon. Solanum carolinense L. Bull or Horse Nettle. Rocky prairies, roadsides, and waste ground. Cireumneutral. General. Solanum —Á— Cav. White Horse Ber Sagi roadsides , and waste und. Cireumneutral. Central and Bed Mo.: St. + Belles Boone, Balin, ackson, Barry, Jasper, Newton, and MeDonald ae. Perhaps native in southwestern Mo. and introduced in other sections, 1935] PALMER & STEYERMARK— PLANTS OF MISSOURI 637 Solanum Torreyi Gray. Rocky open ground and along railroads. Cireumneutral to oxylophile. Seat- ed: St. Louis, Iron, Jackson, and Barry counties. Apparently native in southwestern Mo., and HP elsewhere. Solanum rostratum Dunal. Buffalo aste ground and roadsides. s General. Introduced from the Southwest and recently spreading over the state. Physalis ixocarpa Brotero. Tomatillo. Cultivated and occasionally escaped into waste ground. Jaekson Co. Physalis angulata L Cultivated and waste ground, usually on alluvial soil. Circumneutral. Seat- tered: Pemiseot, St. Louis, Henry, and Jasper counties. Physalis pendula Rydb. Alluvial cultivated and waste ground. Cireumneutral. Western Mo.: Jack- son and Jasper counties. Physalis pubescens L. Ground Cherry. Cultivated and waste ground, moist open fields and ledges along streams. Cireumneutral Southern and central Mo., north to St. Louis and Jackson counties. Physalis pruinosa L. Strawberry Tomato Rocky open ground, roadsides, rich alluvial and waste ground. Cireumneutral. General but seattered: Marion, Pm. Boone, St. Louis, Wayne, Shannon, Taney, Jackson, Barry, and Jasper counti Physalis barbadensis Jacq. Moist rocky ground along streams, and ledges along bluffs. Circumneutral. Western and southern Mo.: Ozark, Taney, Jackson, and Jasper counties. Physalis barbadensis var. obscura (Michx.) Rydb. Rich or rocky woods, bluffs, sandy open ground, and banks along streams. Cireumneutral. Central and southern Mo.: St. Louis, New Madrid, Dunklin, Butler, Shannon, Taney, Jackson, and Barry a Physalis missouriensis Mack. & Bus ich woods along bluffs and real Cireumneutral. Eastern, southern, and central Mo.: Marion, Boone, St. Louis, Jefferson, Shannon, Oregon, Greene, Taney, Barry, Platte, Jackson, Jasper, and MeDonald counties. This has been eonfused with Physalis Lagascae, a southern RE not found in Mo Physalis heterophylla Nees. Roadsides, fields, and waste ground. Cireumneutral. General. Physalis heterophylla var. ambigua (Gray) R re found with the typical form. E St. Louis, Boone, and Web- ster countie Physalis u... var. nyctaginea (Dunal) Rydb. Oeeasionally found with the typical form. Seattered: Scotland, Cole, Miller, Oregon, J ackson, and Greene counties. [Vor. 22 638 ANNALS OF THE MISSOURI BOTANICAL GARDEN Physalis pumila Nutt. Dry or rocky open ground, prairies, and glades. Cireumneutral. General, but scattered : Macon, St. Louis, Phelps, Barry, Greene, Jackson, Clay, Vernon, and asper counti Physalis a Mack. & Bush. Thiekets, prairies, and eultivated and waste ground. Cireumneutral. General. m longifolia Nutt. s and waste ground. Cireumneutral. Eastern Mo.: Marion and St. lesus eountie Physalis eis lata E Mill. Ground Cherry. Dry prairies, rocky open woods, thickets, and waste ground. Cireumneutral to oxylophile. General. Physalis lanceolata Michx. rairies and sandy or rocky open ground. Cireumneutral. Western Mo., loeal: Jaekson Co, PHYSALIS PERUVIANA L. Waste ground. Introduced in St. Louis Co. LYCOPERSICUM ESCULENTUM Mill. See Rh. 11: 56. 1909. "Tomato. Cultivated and escaping in many pla Seattered: St. Louis, Dunklin, Boone, Jackson, and Jasper counties. Well est tablished ding gravel and sand bars of streams and in other situations in some sections. NICANDRA PHYSALODES (L.) Pers. Apple-of-Peru. Physalodes physalodes (L.) Britton (B & B). Cultivated fields and waste ground. Circumneutral. Scattered: Marion, Boone, St. Louis, Crawford, Phelps, Greene, Stone, Jackson, and Newton eounties. LYcIUM HALIMIFOLIUM Mill. Matrimony Vine. Waste ground, roadsides, and about old dwellings. Indifferent. General but seattered: Shelby, Marion, Boone, St. Louis, Jefferson, St. Francois, Cole, Atchi- son, Clay, Jaekson, Greene, Jasper, and Newton counties. DATURA STRAMONIUM L. Jimson Weed. Cultivated and waste ground. Indifferent. General and common. DATURA TATULA L. Jimson Weed, Purple Thorn Apple. Cultivated and waste ground. Indifferent. General. DATURA METEL L. Waste ground and roadsides. Indifferent. Seattered: Boone, St. Louis, Phelps, Vernon, and Jasper eounties, NICOTIANA RUSTICA L. Wild Tobacco. Cultivated and waste ground. Indifferent. St. Louis Co. NICOTIANA LONGIFOLIA Cav Cultivated and escaping into waste ground. Indifferent. St. Louis Co. Fam. SCROPHULARIACEAE VERBASCUM THAPSUS L. Mullein. Pastures, roadsides, and waste ground. Indifferent. General and common. 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 639 VERBASCUM BLATTARIA L. Moth Mullein. Pastures, roadsides, and waste ground. Indifferent. General. VERBASCUM BLATTARIA f. ALBIFLORUM (G. Don) House, N. Y. Mus. Bull. 243- 244: 45. 1923. Commonly found with the typical form. General. LINARIA VULGARIS Hill. Butter and E ields, waste ground, and along roadsides and railroads. Oxylophile to cir- eumneutral. General Linaria canadensis (L.) Dumont var. texana (Scheele) Pennell, Proc. Acad. Nat. Sci. Phila. 73: 502. 1921. Blue Toadflax Rocky ee, glades, a and sanity or cherty open ground. Oxylophile. ud central, and southern Mo.: Marion, Lineoln, ps ox Warren, St. Charles, St. Lou CR Franklin, Ste. Genevieve, St. Francois, Wash- ington, ubi Det Shannon, Oregon, Iron, "ue yp "nnm Mis- sissippi, Dunklin, Butler, Wayne, Jaekson, St. Clair, Polk, Greene, Lawrence, Vernon, Jasper, Newton, and MeDonald dE Typical ue canadensis has a more eastern range and does not exten LINARIA CYMBALARIA (L.) Mill. Kenilworth I ste ground and roadsides. Indifferent. Introdueed in Marion and St. Louis eounties. LINARIA SPURIA (L.) Mill. Cancerwort. Rocky waste ground, and along sand and gravel bars of streams. Seattered: Boone, Barry, and MeDonald counties. KıckxIA ELATINE (L.) D Md See Torreya 22: 81-84. 1922. Cancer-root. Linaria Elatine (L.) Mill. (G). Waste ground, and along mes and gravel bars of streams. Oxylophile. Cen- tral and southern Mo., scattered: St. Louis, Jefferson, Franklin, Gasconade, Perry, Bollinger, Reynolds, Shannon, Oregon, Texas, Laclede, Douglas, Clay, nd Jackson countie Collinsia verna Nutt. Blue-eyed-Mar Rich woods and thickets. rias : prt Central and southern Mo., seattered southward: Warren, St. Charles, St. Louis, Jefferson, St. Fran- eois, Seott, Audrain, Callaway, Boone, ea ede, Monte mery, Howard, Cooper, Saline, Clay, Jackson, Cass, J olei Res and MeDonald eounties. Collinsia violacea Nutt Rocky open woods. Circumneutral to oxylophile. Southern Mo.: Jefferson, Dunklin, ee ag St. Clair, Cedar, Polk, Greene, Bates, Vernon, Jasper, and Newton tie Scrophularia marilandica L. Figwo Open woods and thickets. Geet E General. Minore lanceolata Pursh. See Torreya 22: 81-84. 1922. Scrophularia leporella Bieknell (G), (B & B nds woods and thickets. Circumneutral. edis Mo., local: Jackson Co. Pentstemon pallidus Small. Beard-tongue entstemon canescens of auth., not Britton (G), in part. Prairies, open woods, and bluffs. Oxylophile to eireumneutral. [Vor. 22 640 ANNALS OF THE MISSOURI BOTANICAL GARDEN Pentstemon calycosus Small, Rocky woods and bluffs. Cireumneutral. Eastern and southern Mo., seat- tered: St. Louis and Texas counties. Pentstemon tubiflorus Nutt. rairies and open woods. Oxylophile to cireumneutral. General, but e monest in southwestern - and absent from much of the Ozark epe and the southeastern lowlands: Putnam, Marion, Shelby, Monroe, Boone, Franklin, Cape Girardeau, Dunklin, Pulaski, Camden, Laclede, Douglas, Wright, Webster, Greene, Taney, Christian, Barry, Hickory, Clay, Jackson, Vernon, Barton, Jasper, Newton, and McDonald counties. Pentstemon arkansanus Pennell, Proc. Acad. Nat. Sci. Phila. 73: 493. 1921. Rocky open woods, bluffs, and glades. Cireumneutral to oxylophile. South- ern Mo.: Madison, Ozark, Taney, Stone, Barry, Newton, and MeDonald counties. Pentstemon arkansanus var. pubescens Pennell, Proc. Acad. Nat. Sci. Phila. 73: 494. 1921. Occasionally found with the typical form. Barry Co. Pentstemon Digitalis (Sweet) Nutt. Pentstemon laevigatus var. Digitalis (Sweet) Gray (G). Open woods, thickets, prairies, and meadows, Cireumneutral. General. Pentstemon grandiflorus Nutt. ren hills and plains. Cireumneutral to caleiphile, Northwestern Mo.: Atchison Co a Cobaea Nutt. var. purpureus Pennell, Proc. Acad. Nat. Sci. Phila. 73: 490. 1921 Glades and bald knobs. NUM to cireumneutral. Southwestern Mo.: Ozark, Stone, and Taney counti Chelone glabra L. Turtlehead. Wet rocky or boggy a" along small streams and bluffs. Oxylophile. Southeastern and central Mo.: Boo ne, St. Louis, Jefferson, — Bollinger, Stoddard, Dunklin, E Clinton, Clay, and Jackson counties Chelone obliqua L. var. speciosa Pennell & Wherry, Bartonia 10: 19, pl. 2, fig. 1. 1929. Swamps and borders of streams. Oxylophile to cireumneutral. Northeastern and eentral Mo.: Lewis and Chariton counties. PAULOWNIA TOMENTOSA (Thunb.) Steud. Empress Tree. Planted and occasionally escaped. Marion, Perry, and Ripley counties. Mimulus ringens L. Monkey Flower. Bogs, low woods, and borders of streams, commonest in alluvial soils along the larger rivers. Circumneutral. General. Mimulus alatus Ait. Bogs, low woods, and borders of streams. Cireumneutral. General. Com- moner than the preceding species. M glabratus HBK. var, Fremontii (Benth.) Grant, Ann. Mo. Bot. Gard. 11: 190. 1924, Wet ledges along bluffs, and about springs. Cireumneutral to anne Southern Mo., scattered: Ste. Genevieve, Greene, and Barry counties 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 641 Leucospora multifida (Michx.) Nut onobea multifida (Michx.) Se (G), (B&B Low open woods and fields. Cireumneutral. trad and southern Mo. Mecardonia acuminata (Walt.) Small. Water Hyssop. Bacopa acuminata bind ) Robinson Low wet woods, mps, and sloughs. Oxylophile to eireumneutral. South- ern Mo.: Minisipni, Donki in, Jasper, and Newton counties. Macuillamia rotundifolia (Michx.) Raf. See Torreya 22: 79. 1922. Bacopa rotundifolia (Michx.) Wettst. (G). Bramia rotundifolia (Michx.) Britton (B & B). Low wet woods, and borders of ponds, sloughs, and streams. Cireumneutral to ER Central and southern Mo., scattered: Boone, Lincoln, St. Charles, t. Louis, Jefferson, Dent, Maries, Ray, Jackson, Cass, Greene, Jasper, Newton, and MsDonald dx Ilysanthes dubia (L.) Barnhart. False Pimpernel. Low wet woods and borders of sloughs and ponds. Cireumneutral. General: ape am, Marion, Boone, Lineoln, St. Charles, St. Louis, Jefferson, Franklin, e. Genevieve, Iron, Missins sippi, Dunklin, Butler, Harrison, Buchanan, Clay, J a Bates, Vernon, Greene, Taney, Stone, Jasper, and Newton mn Ilysanthes inaequalis (Walt.) Pennell, Torreya 19: 149. 1919. Ilysanthes anagallidea of auth. in part, not Gratiola anagallidea Michx. (G). Ilysanthes dubia of auth., not Gratiola dubia L. (B & B). et ground goce streams, and borders of ponds and sloughs. Cireumneutral to om! phile. General but scattered: Linn, Monroe, Ralls, Boone, St. Louis, Ste. Genevieve, St. Y Missis Dunklin, Buchanan, Jackson, Henry, Greene, Taney, Barton, J ya and Newton counties. Gratiola lutea Raf. See Rh. 34: 147. 1932. Clammy Hedge Hyssop. Gratiola virginiana of y not L. prs (B & B). Gratiola neglecta 'Tor Low cay and on borders of ponds, sloughs, and slow streams. Cireum- neutral. General. Gratiola virginiana L. See Rh. 20: 65. 1918. Gratiola sphaerocarpa Ell. (G), (B & B). ddy ers of ponds, sloughs, and ditches. Circumneutral. General but scattered : er Adair, Shelby, Marion, Monroe, Audrain, Boone, Callaway, St. Louis, Washington, Shannon, Dent, Ripley, Texas, Laclede, Howell, Ozark, Wright, Webster, Linn, Livingston, Saline, Jackson, Benton, Greene, Jasper, and Newton counties. Veronicastrum virginicum (L.) Farwell, Drug. Cire. 61: 231. 1917. Culver’s- ot. Veronica virginica L. (G). Leptandra virginica (L.) Nutt. (B & B). Open woods and thickets. Cirene General. Veronicastrum virginicum f. villosa (Raf.) Pennell, Rh. 23: 6. 1921. In similar situations to the preceding. Cireumneutral. "Genéral and eom- monly oecurring with the species. [Vor. 22 642 ANNALS OF THE MISSOURI BOTANICAL GARDEN Veronica serpyllifolia L. Thyme-leaved Speedwell. Swampy woods and waste ground. Circumneutral to oxylophile. Southeast- ern Mo.: Cape Girardeau, Bollinger, pe Stoddard counties. Veronica peregrina L. Neckweed. Cultivated and waste ground. Indifferent. General and common. Veronica peregrina var. xalapensis (HBK.) Pennell, Torreya 19: 167. 1919. Occasionally found with the typical form. Waste ground. Indifferent. Seat- tered: St. Louis and Pemiseot counties. VERONICA ARVENSIS L. Corn Speedwell. Cultivated and waste ground, and open rocky woods, glades, and ledges along bluffs. Cireumneutral to calciphile. General. Veronica connata Raf. Water Speedwell. onica Anagallis-aquatica of auth. in part, not L. (G), (B ^ and along streams and spring branches. Cireumneutral to otylophile. West-central and southern Mo., seattered: Perry, Bollinger, Reynolds, Ripley, Oregon, Shannon, Dent, Camden, erm Howell, Laelede, Dallas, Benton, Doug- las, Ozark, Barry, and Jackson countie Veronica americana Sehwein. American Brooklime. et open ground. Indifferent. Local: Boone Co Limosella subulata Ives. See Rh. 20: 160. 1918; also Torreya 22: 83-84. 1922, Mudwort Limosella aquatica var. tenuifolia of auth., not L. tenuifolia Wolf (G). Limosella aquatica of auth., not L. (B & B). Sand bars of Missouri River. Loeal: Jaekson Co. Dasistoma macrophylla (Nutt.) Raf. See Rh. 20: 68. 1918. False Foxglove. eymeria macrophylla Nutt. (G). Afzellia macrophylla (Nutt.) Ktze. (B & B). Open woods and thiekets. Oxylophile. General. Aureolaria grandiflora M Pennell var. cinerea Pennell, Proc. Acad. Nat. Sei. Phila. 73: 510. Gerardia jee do var. serrata (Torr.) Robinson, in part (G). Dasystoma serrata (Benth.) Small, in part (B € B). oeky woods and thickets. Oxylophile. Middle and western Mo., mostly south of the Missouri River: Crawford, Dent, Gasconade, Boone, Moniteau, Miller, Pulaski, Phelps, Benton, Camden, Polk, Greene, Webster, Wright, Doug- las, Ozark, Christian, Taney, Stone, Barry, Henry, Cedar, Vernon, Jasper, New- ton, and McDonald counties. Aureolaria grandiflora var. pulchra Pennell, Proc. Acad. Nat. Sci. Phila, 80: 392. 1928 i Rich or rocky open woods. Oxylophile. Northeastern and east-central Mo.: lark, Knox, Macon, Adair, Schuyler, Sullivan, Shelby, Marion, Ralls, Monroe, Lincoln, St. Charles, St. Louis, Jefferson, Franklin, Crawford, Gasconade, and Maries counties. it pa —— Pennell, Bull. Torr. Bot. Club 40: 414. 1913 w False Fox m meti he var. pectinata Nutt., in part (G). 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 643 Dasystoma pectinata (Nutt.) Benth., in part (B & B). Rocky T woods and borders of glades. Oxylophile. East-central and cim da Mo.: St. Louis, Jefferson, Franklin, Iron, Madison, Wayne, Oregon, , De ki, Crawford, Maries, Pulaski, Texas, Wright, Webster, Greene, a di, Do ouglas, Ozark, Stone, and Barry counties. Aureolaria flava (L.) Farwell var. macrantha Pennell, Proc. Acad. Nat. Sci. Phila. 73: 511. 1921. Gerardia flava L., in part (G). Dasystoma flava (L.) Wood, in part (B & B). Rocky woods. Oxylophile. Southern Mo., mainly in the eastern Ozark region: Phelps, Reynolds, Iron, Madison, Wayne, Scott, and Oregon counties Aureolaria calycosa (Mack. € Bush) Pennell, Proc. Acad. Nat. Sci. Phila. 73: 512. 1921 Dasystoma calycosa Mack. € Bush (B & B). Rocky open woods and borders of glades. Oxylophile. Southern Mo., mostly in the eastern Ozark region: St. Francois, Madison, Bollinger, Cape Girardeau, Iron, Wayne, Carter, Ripley, Shannon, Oregon, Texas, Howell , Douglas, an Ozark counties. Otophylla auriculata Rena Small. See Rh. 20: 137. 1918. G ia auriculata Miehx. (G). Prairies, thickets, and open woods. Circumneutral to oxylophile. General but seattered: Clark, Adair, St. Louis, Shannon, Ozark, Greene, Barton, Jasper, Newton, and MeDonald counties. Agalinis purpurea (L.) Pennell. Gerar G Prairie swales and moist open ground along small streams. Circumneutral to oxylophile. Southern and west-central Mo., scattered: Stoddard, Shannon, Jackson, and Greene counties. Agalinis aspera (Dougl.) n NE rdia aspera Dou G). Prairie swales and Be ground along small streams. Circumneutral. West- ern Mo., and loeally eastward in Crawford Co.: Atchison, Holt, Jackson, Jasper, and Crawford counties. e p (Ell) Raf. G ia. fasciculata Ell. (G). id swales and borders of woods and thiekets. Circumneutral. South- western Mo.: Bates, Barton, Jasper, and Newton counties. Agalinis tenuifolia (Vahl) Raf. Gerardia tenuifolia Vahl (G). rairie swales and moist open banks. Circumneutral to oxylophile. Central and southern Mo.: Boone, St. Louis, Jefferson, Franklin, St. Francois, Dent, Shannon, B. Oregon, Jackson, Benton, Christia and J asper counties. Agalinis tenuifolia var. macrophylla (Benth.) Blake, Rh. 20; 7141918. erardia tenuifolia var. macrophylla Benth. (G). Gerardia Besseyana Britton (B & B). Prairies and moist open ground. Cireumneutral to oxylophile. General. This is the commonest form throughout the state. [Vor. 22 644 ANNALS OF THE MISSOURI BOTANICAL GARDEN Agalinis tenuifolia var. parviflora (Nutt.) Pennell, Proc. Acad. Nat. Sci. Phila 73: 525. 1921. Prairie swales and open banks. Cireumneutral to oxylophile. Middle and western Mo., seattered: Boone, Jackson, Douglas, Greene, and Jasper eounties. Agalinis Gattingeri Small. Gerardia tenuifolia of auth., not Vahl (G), in p Oxylophile. d HB v and southern Mo.: Clar Shannon, Texas, Miller, ardia Rocky open woods. Franklin, Crawford, Ma Ozark, Taney, Greene, Barry, Jasper, Newton, and McDonald counties on, oone, Osage, Agalinis Skinneriana (Wood) Britton. Gerardia Skinneriana Wood open woods and thiekets. Oxylophile. West-central and southern Mo.: Iron, Dunklin, Shannon, Taney, Barry, Jackson, Jasper, Newton, and MeDonald Oxylophile. Southern Mo., co a, viridis I Ben Gerardia viridis Sm hickets, open en and borders of woods scattered: Shannon, Dunklin, and Jasper count Agalinis heterophylla (Nutt.) Small Moist nn open ground. Oxylophile. Southeastern Mo.: Dunklin Co. — americana L. Blue Hearts. and prairies and glades. Circumneutral to oxylophile. East-eentral and His unn Mo.: St. Louis, Jefferson, Iron, Dent, Shannon, Ripley, Howell, Greene, Stone, Barton, Jasper, iic and MeDonald counti Castilleja coccinea (L.) Spre Indian Blanket, Painted Cup. Upland prairies, P i moist open woods. Circumneutral. General but uncommon in most se Castilleja coccinea f. lutescens Farwell, Am. Midl. Nat. 8: 276. kv n do found with the typieal form. Washington, daa Jasper, and Atchison and Holt Northwestern Mo Newton countie — sessiliflora Pursh. ess hills and aid, Calciphile. local: Greene Co. Mora Castilleja purpurea G. Don. open ground. Circumneutral, Southwestern Mo., Pedicularis canadensis L. Wood Betony, Lousew Open woods, thickets, and prairies u ec ical to oxylophile. General. Pedicularis lanceolata Michx. We ods and bogs. Oxylophile. Southern Mo., scattered: Iron, Shannon, Morgan, and Greene counties. Fam. LENTIBULARIACEAE var. americana Gray. =>.” Bladderwort. mneutral. General but scattered: Utricularia vulgaris L. Sloughs, ponds, and sluggish streams. Cire Putnam, St. Charles, St. Louis, Teftereon, ar Butler, Dunklin, Oregon, 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 645 Platte, Clay, Jackson, Greene, and Jasper counties. This has been confused with Utricularia minor, which is not known in Mo. Utricularia gibba L. Stagnant water of ponds, lakes, and sloughs. Circumneutral. Central and southern Mo., scattered: Boone, St. Charles, St. Louis, Iron, Reynolds, Dunklin, Oregon, Texas, Greene, Jackson, and Jasper counties, This species has been con- fused ROBA. with Utrioularia pumila Walt. (U. biflora Lam.) which is not known to occur in Mo. Fam. OROBANCHACEAE Orobanche uniflora L. Small Cancer-r Rocky open woods and fields. roue e. General but seattered: Ralls, Boone, Cole, St. Louis, Jefferson, Franklin, Ste. Genevieve, Iron, Carter, Craw- ford, Phelps, Maries, Miller, Camden, Greene, Christian, Platte, Jackson, John- son, Henry, Vernon, Jasper, Barry, and McDonald counties. Epifagus ee (L.) Bart. Beech-drops. oods, under beech trees. Circumneutral to oxylophile. Crowleys Ridge, Nr bM M. local: Seott Co Fam. BIGNONIACEAE inpr m radicans (L.) Seem. "Trumpet Creeper, Trumpet-vine. a radicans (L.) Juss. (G). ra radicans L. (B " DB): Open woods, thiekets, banks of streams, and along road and railroad embank- ments, Indifferent. General, but scattered northward. Native in the Ozark region and introduced or escaped in other sections. Bignonia capreolata L. Cross-vine ow alluvial woods and thickets along streams. Cireumneutral to oxylophile. Southeastern Mo.: Iron, Bollinger, Cape Girardeau, Mississippi, Pemiseot, Dunk- lin, Butler, and Ripley counties. Catalpa speciosa Warder. Catalpa, Cigar Tre Low or upland woods, and also introduced ins streams and in waste ground. Eastern and southern Mo.: Pike, Iron, Madison, Rollinger, Cape Girardeau, Scott, a h New Madrid, Stoddard, Pemiscot, Dunklin, Oregon, Howell, Dallas, Greene, Jasper, and Newton counties. Native in southeastern Mo. and OREN, from pruni era Catalpa bignonioides Walt. Along streams and roadsides and in waste ground. Indifferent. Scattered: St. Louis, Boone, Jackson, Jasper, and Newton counties. Introduced Fam. PEDALIACEAE SESAMUM ORIENTALE L. es i Sesamum indicu Sandy open cay Introduced in Jackson Co. [Vor. 22 646 ANNALS OF THE MISSOURI BOTANICAL GARDEN Fam. MARTYNIACEAE Martynia louisiana Mill. Devil's-claws, Unicorn Plant a eultivated and waste ground. Caleiphile to eireumneutral. Seat- tered: Boone, St. Louis, St. Francois, New Madrid, Dunklin, Ozark, Jackson, Johnson, Dade, and emo counties. Fam, ACANTHACEAE Dianthera americana L. Water Willow. Borders of shallow streams, sloughs, and ditches. Circumneutral. Central and southern Mo.: Boone, Montgomery, Warren, St. Louis, Jefferson, Franklin, Washington, St. Francois, Ste. Genevieve, Perry, Cape Girardeau, ap Madison, i Reynolds, Wayne, heb Ripley, Oregon, Shannon, How Texas, Dent, Crawford, Gasa ade, Osage, Cole, Miller, Maries, Phelps, P se Laclede, Mu Webster, Douglas, Oladk, (rans, Christian, Taney, Stone, Morgan, Camden, Miller, Dallas, Hiekory, Polk, Barry, Jackson, Jasper, Newton, and McDonald counties. — age Walt. woods and borders of swamps. Circumneutral. Southeastern Mo.: u. pa Madrid, Pemiscot, Dunklin, Butler, and Ripley counties. Ruellia caroliniensis (Walt.) Steud. See Rh. 17: 134. 1915. Wild Petunia. Ruellia ciliosa Pursh (G), (B & B Rocky prairies, hillsides, and glades. Oxylophile to cireumneutral. General. Ruellia caroliniensis var. parviflora (Nees) Blake, Rh. 17: 134. 1915. Ruellia ciliosa var. parviflora (Nees) Britton (G). Ruellia ri a Britton (B & B). Glades and roe edges. Oxylophile. West-central and southern Mo.: Ste. vind M pen Jaekson, Jasper, and Newton counties. Ruellia strepens L. Rich or rocky open woods and thickets. Cireumneutral to calciphile, General. Ruellia pedunculata Tor open woods a glades. Cireumneutral to calciphile. East-central and southern Mo.: St. Louis, Jefferson, Franklin, Ste. Genevieve, Iron, Madison, Dunklin, Butler, E Reynolds, Dent, Oregon, Shannon, Texas, Howell, Phelps, Pu laski, Wright, Webster, Douglas, Ozark, Taney, Stone, Barry, and McDonald count Mirac brachiata E Spreng. See Rh. 11: 57. 1909. m brachiatum (Pursh) Ktze. (B & B). Qm alluvial woods and thickets. Cireumneutral. East-eentral and southern St. Louis, Gasconade, Shannon, Ripley, Ozark, Taney, Stone, Barry, oni Newton, and MeDonald counties. Fam. PHRYMACEAE Phryma Leptostachya L. Lopseed Rocky or alluvial woods and thiekets. Circumneutral. General. 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 647 Fam. PLANTAGINACEAE Plantago cordata Lam. Swampy woods, sloughs, and rocky stream beds. Cireumneutral. Northern, central, and southeastern Mo.: Adair, Boone, St. Louis, Jefferson, Dent, Iron, Carter, and Shannon counties Plantago major L. Plantain. ields, waste ground, and door yards. Indifferent. General, but less com- mon than the next species. Plantago Rugelii Dene. Plant Fields, waste ground, M" door yards. Indifferent. General and in every eount PLANTAGO LANCEOLATA L. Rib Grass. Pastures, roadsides, and waste ground. Indifferent. General. Broad-leaved specimens of this species have sometimes been confused with Plantago media which has not been found in Mo Plantago Purshii R. € S. Salt-and-pepper Plant. Loess ae glades, dry prairies, and waste ground. Cireumneutral. Central and northwestern Mo.: St. Louis, Holt, Clay, and Jackson counties. Probably native in Reese Mo. and introduced elsewher Plantago spinulosa Dene Waste ground. Indifferent. Introduced in Jackson Co. Plantago aristata Michx. ields, glades, barrens, and waste ground. Oxylophile to eireumneutral. 1. Plantago virginica Fields, glades, ae ded waste ground. Oxylophile to eireumneutral. General. Plantago rhodosperma Den aste ground. nist Introduced in Jackson Co. Plantago heterophylla Nutt andy open ground. Oxylophile. Southeastern Mo.: Seott Co. Plantago elongata Pursh. Plantago pusilla Nutt. (B & B). Fallow fields, prairies, and glades. Oxylophile to eireumneutral. General but most abundant southward: Boone, Audrain, Marion, TER. Montgomery, St. Louis, Jefferson, Franklin, Ste. Genevieve, Iron, Madison, Cape Girardeau, Dunklin, Butler, Wayne, Ripley, Carter, Shannon, Texas, rad Laclede, Greene, Taney, Jaekson, Lawrence, Jasper, and Newton counties. Fam. RUBIACEAE Galium virgatum N Rocky n is and prairies. Caleiphile to ee Central and southern Mo.: St. Louis, Jefferson, Franklin, Washingto Ste. Genevieve, Ripley, de ac Shannon, Texas, Laclede, Daten Miller, Camden, Benton, Hickory, Polk, St. Clair, Greene, Webster, Ozark, "yu Stone, Barry, Jasper, Newton, Er phum counties. [ Vor. 22 648 ANNALS OF THE MISSOURI BOTANICAL GARDEN Galium Aparine L. Cleavers, Goose Grass. Rich woods, thickets, alluvial and waste ground. Cireumneutral. General. Galium Aparine var. Vaillantii (DC.) Koch. In similar situations to the typical form. Jackson and Jasper counties. Galium pilosum Ait. Rocky woods and thiekets. Cireumneutral to oxylophile. Central and south- ern Mo. Galium pilosum var. puncticulosum (Miehx.) T. € G. Oceasionally found with the typieal form. Southern Mo., seattered: Jeffer- son and Jasper counties Galium circaezans Michx. Wild Licorice. ocky woods and thickets. Circumneutral. General. Galium arkansanum Gray. Rocky woods. Oxylophile to eireumneutral. Southern Mo.: Washington, Ste. Genevieve, Seott, Iron, Reynolds, Carter, Ripley, Dent, Shanno on, Oregon, Howell, Texas, Wright, Douglas, Ozark, Taney, Greene, Stone, Barry, Newton, and MeDonald counties Galium boreale L. var. hyssopifolium (Hoffm,) DC. See Rh, 30: 106-107. 1928. Northern Bedstraw Ledges along high north-faeing sandstone and dolomite bluffs. Oxylophile to eireumneutral. Southeastern Mo., loeal: Shannon Co. Galium tinctorium L. Wild Madder. Swamps and low wet woods and thiekets. Circumneutral. General, Galium m Michx, mpy places and wet ground among hummocks in sink-hole ponds. Cir- Bici Southeastern Mo.: Mississippi, Dent, and Texas counties. Galium concinnum T. & G. Shining Bedstraw Prairie swales and moist open woods. Cireumneutral. General. Galium triflorum Michx. Fragrant Bedstraw. Rich or moist woods and low open e Cireumneutral. General. —— glabra Michx, Smooth Button- wet woods, fields, and borders of iei sloughs, and streams. Circum- ail to oxylophile. Central and southern Mo.: Lin coln, St. Charles, St. Louis, Jefferson, Perry, Mississippi, New Madrid, Fadik: Dunklin, Butler, Ripley, Reynolds, Oregon, Howell, Gasconade, Maries, Miller, Pulaski, pm ede, Dallas, Henry, Greene, Taney, Barry, Jasper, Newton, and McDonald counties. Diodia teres Walt. Rough Button-weed. Prairies, glades, fields, and waste ground. Oxylophile to eireumneutral. Gen- eral but commonest south of the Missouri River Diodia virginiana L. Button-weed. Low wet woods and borders of streams and bayous. Cireumneutral to o oxylo- phile. Southeastern Mo.: Bollinger, si vest d New Madrid, Pemiscot, Dunk- lin, Stoddard, Wayne, Butler, and Ripley countie Mitchella repens L. Partridge Berry. Moist sandy banks of streams, sandy bogs, and moist ledges of sandstone 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 649 bluffs. Oxylophile. Eastern Mo., mostly south of the Missouri River: Lincoln, Jefferson, Ste. Genevieve, Perry, Madison, New Madrid, Dunklin, Butler, and Ripley counties. poa occidentalis L. Button-bush. Low swampy woods and borders of streams and sloughs. Circumneutral. General zd probably in every ah Cephalanthus occidentalis var. pubescens Raf. Swampy woods and borders of RO Cireumneutral. Southeastern Mo.: Seott, Dunklin, lr and Ripley counties. Houstonia caerulea L. Bluets, Innocence. Moist open woods, open banks, and moist sandstone or granitie ledges. Oxylo- phile. Southeastern Mo., mostly in the Iron Mountain sub-region: Ste. Gene- vieve, St. Francois, Iron, Madison, Wayne, Butler, Dunklin, Ripley, Carter, Shan- non, Dent, and Texas counties Houstonia patens Ell. Star Violet Dry open ground, glades, and rdi ledges, usually in sandstone or granitie areas. Oxylophile. Eastern and middle Mo., mostly south of the Missouri River: Lineoln, Warren, Jefferson, Franklin, Orawford, Ste. Genevieve, St. Fra Iron, Madison, Wayne, Butler, Dunklin, Ripley. Carter, Reynolds, Ain Dent, Phelps, Dallas, Douglas, and Wright counties. Houstonia minima Beck. Small Bluets Prairies, pastures E rocky ledges, and open soils. Oxylophile to eireum- neutral. Central aid hern Mo. south of a line from Lincoln, Audrain, Ran- dolph, en and ae counties to Jackson Co. Houstonia purpurea L. Moist open woods and ledges along bluffs. Oxylophile. Southwestern Mo.: Webster, Ozark, Greene, Stone, Bates, Vernon, Barton, Jasper, and McDonald counties. Houstonia tenuifolia Nutt. Rocky open woods and glades. Oxylophile. Southeastern Mo., local: Madi- son Co. Houstonia ar Gaertn. Rocky open woods, glades, and ledges along bluffs. Circumneutral. East- central ‘and southern Mo., south and east of a line drawn from St. Charles, ES and Camden counties a Newton Co. This has been confused with H. cilio uid does not occur in Missouri. Houstonia lanceolata (Poir.) Britton eky open woods. Oxylophile i circumneutral. East-central and southern Mo., Roses ed: St. Louis, Jefferson, Taney, and Barry counties. Houstonia E Miehx. eky ledges, glades, bluffs, and loess hills. Circumneutral to an Cen- southern, and northwestern Mo.: Macon, Howard, Boone, Montgomery, Warren, St. Charles, St. Louis, Jetterson, Franklin, lacio St. Francois, Ste. Genevieve, Perry, Cape Girardeau, Bollinger, Madison, Dunklin, ag Reynolds, Wayne, AS, Ripley, Ore Mg Howell, Shannon, Texas, Dent, ford, Phelps, Gasconade, Osage, Maries, Cole, Pulaski, Miller, Morgan, nt [Vor. 22 650 ANNALS OF THE MISSOURI BOTANICAL GARDEN Henry, Camden, Laclede, Dallas, Hickory, Polk, Greene, Douglas, Wright, Web- ster, Ozark, Taney, Stone, Barry, Cedar, Atchison, Holt, Buchanan, Bates, Jasper, Newton, Christian, and MeDonald counties. Oldenlandia uniflora L. Moist sandy ground. Oxylophile. Southeastern Mo., local: Dunklin Co. Fam. CAPRIFOLIACEAE LONICERA JAPONICA Thunb. Japanese Honeysuekle. Open woods, thiekets, and roadsides. Escaped from eultivation; seattered but becoming more widespread: Boone, Pike, St. Charles, St. Louis, Cape Girardeau, — Reott, nm Dunklin, Butler, Shannon, Jaekson, Vernon, Bar- ton, and Jasper countie Lonicera sempervirens L. Trumpet Honeysuckle. Thickets and roadsides. Escaped from cultivation in our area. Scattered: St. Louis, Greene, Barton, and Jasper counties. Lonicera prolifera (Kirch.) Rehder, Rh. 12: 166. 1910. Grape Honeysuckle. Lonicera Sullivantii Gray (G), (B & B). Thiekets and wooded bluffs. Calciphile. Northern, central, and western Mo.: Mercer, Sullivan, Schuyler, Adair, Clark, Knox, Shelby, Warren, Boone, Saline, Pettis, Benton, Morgan, Clay, Jackson, Cass, Dade, Lawrence, Greene, and J Me counties. Lonicera dioica L. var. glaucescens (Rydb.)-Butters, Minn. Trees & Shrubs. 289. 1912 Lonicera glaucescens Rydb. (G), (B & B). Thiekets, bluffs, and banks of streams, Circumneutral to ealeiphile. North- ern Mo., south locally in Morgan Co.: Clark, Lewis, Ralls, Atehison, and Morgan counties. Lonicera flava Sims, Yellow Honeysuckle. Rocky woods and bluffs. Oxylophile to eireumneutral. Southern Mo.: Madi- son, Iron, Wayne, Reynolds, Carter, Ripley, Be Pigg Gasconade, Maries, Pu Crawford, Dent, Texas, Howell, Phelps, Pulaski, n, Laclede, Dallas, Miller, Hickory, Webster, Wright, old Ozark, Raney, pes Barry, and MeDonald counties. Lonicera flavida Cockerell, Rh. 12: 167. 1910. Bluffs and rocky ground along streams. Caleiphile to eireumneutral. South- western Mo.: Greene, Taney, Stone, and Barry counties, Symphoricarpos orbiculatus Moench. Buck Brush, Indian Curran Thiekets, roeky bluffs, and banks of streams. MM to oxylophile. General and probably in every county. Symphoricarpos occidentalis Hook. Wolfber Loess hills, thickets, and open banks. Caiciphile. Northern and west-central Mo.: Adair, Nodaway, Atchison, and Jackson counties Triosteum perfoliatum L. Horse Gen Open woods and thickets. ern A General. 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 651 Triosteum aurantiacum Bicknell var. illinoense (Wieg.) Palmer € Steyermark, comb. nov. Triosteum illinoense (Wiegand) Rydb. (R). Triosteum perfoliatum var. illinoense Wiegand. Rich or rocky woods and bluffs. Cireumneutral. Eastern and middle Mo.: Putnam, Schuyler, Sullivan, Adair, Monroe, Audrain, Boone, Warren, Carter, Crawford, Texas, Pulaski, Shannon, Morgan, and Dallas counties. Triosteum angustifolium L. Rocky open woods and ledges. Cireumneutral to calciphile. i el and southern Mo.: St. Louis, Jefferson, Franklin, Carter, Ripley, Den non, Howell, Texas, Morgan, pon Wright, Hickory, Webster, Douglas, Tr ERN and MeDonald counties. Viburnum molle L. Arrow-wood. Woods along streams and at base of high bluffs. Calciphile. Central and southern Mo., seattered: Warren, Crawford, Texas, Howell, Benton, Christian, and Stone counties. Viburnum molle f. leiophyllum Rehder, Jour. Arnold Arb. 5: 57. 1924. In similar situations to the typieal form. Western Mo., south of the Missouri River, scattered: Benton, Taney, Stone, and MeDonald counties. Viburnum pubescens (Ait.) Pursh var. Deamii Rehder; Jour. Arnold Arb. 5: 58. 19 Low woods along river. Cireumneutral Northeastern Mo., local: Shelby Co. Viburnum affine Bush ex Rehder. See Rh. 20: 14. 1918. Viburnum evade of auth., not Pursh (G), (B & B). Roeky or open woods and bluffs, Circumneutral to ealeiphile. General but scattered: Mötesr, Sullivan, Sehuyler, Clark, Shelby, Adair, Boone, Phelps, arter, Oregon, Shannon, Howell, Benton, Taney, Stone, Barry, and MeDonald counties. Viburnum affine var. hypomalacum Blake, Rh. 20: 14. 1918. In similar situations to the typical form. Cireumneutral to ealeiphile. North- eastern and central Mo.: Clark, Lewis, and Morgan counties. Viburnum prunifolium L. Black Haw. oods, thickets, and banks of streams. Circumneutral. General and prob- ably in every county. Viburnum prunifolium var. Bushii (Ashe) Palmer & dci 5 comb. nov. Viburnum Bushii Ashe, Jour. Elisha Mitchell Sci. Soc. 40: 1924. Rocky or open woods and thickets. Circumneutral. Central ee southern Mo.: Cape Girardeau, Washington, Crawford, Iron, Reynolds, Carter, Ripley, Shan- non, Texas, Dent, Phelps, Laclede, Camden, Boone, Randolph, Cooper, Moniteau, Saline, Lafayette, Clay, Jackson, Henry, Hickory, Polk, Cedar, Lawrence, Barton, and Jasper counties. This has sometimes been confused with V. Lentago. Viburnum rufidulum Raf. Black Haw. Rocky woods, thickets, glades, and banks of streams. Cireumneutral to eal- eiphile. Central and southern Mo.: Common south of the Missouri River, and north to Ralls, Monroe, and Clay counties. [Vor. 22 652 ANNALS OF THE MISSOURI BOTANICAL GARDEN Viburnum Lentago L. Nannyberry Woods and bluffs. Cireumneutral. Northeastern Mo., loeal: Sehuyler Co. Sambucus canadensis L. Elderberry Open woods and thickets. Cireumneutral. General and probably in every county. Sambucus canadensis var. submollis ie in Sarg. Trees & Shrubs 2: 188. 1911. In similar situations to the typical f Circumneutral. General but scat- tered: Clark, St. Louis, Stoddard, LS Sewell, oen Atchison, Jackson, Jasper, Newton, and Barry countie — — Michx. Red dd racemosa of auth., not L. (G), (B & B). aed vein and rocky banks of streams. Circumneutral. Northeastern Mo., loeal: Marion Co. Fam. VALERIANACEAE Valerianella radiata (L.) Dufr. Beaked Corn Salad. Prairies, fields, and roadsides. Cireumneutral. General. Valerianella radiata var. leiocarpa (T. & G.) Krok. Fields, glades, and roeky open or waste ground. Frequently found with the typieal form. General but seattered: Marion, Lineoln, St. Louis, Jefferson, Franklin, Ste. Genevieve, Iron, Wayne, Jackson, Henry, e Jasper counties. bigger stenocarpa (Engelm.) Krok. cky eot and glades. Cireumneutral, Scattered: Pike, Jackson, and Jasper coun Valerianella e (T. € G.) Walp. Rocky open woods and glades. Cireumneutral to oxylophile. Southern Mo.: Madison, Taney, Stone, Barry, and MeDonald counties, Fam. DIPSACACEAE DIPSACUS SYLVESTRIS Huds. Teasel. Along roadsides, railroads, and gravel bars of small streams. ‚ Indiffe rent. entral and southern Mo.: Boone, Montgomery, Warren, St. Lou , Jefferson, Franklin, Osage, Clay, Webster, Greene, Barry, Jasper, and Kemon | ea Fam. CUCURBITACEAE Cucurbita foetidissima HBK. Wild Gourd, Missouri Gourd. Fields, waste ground, and along railroad and road embankments. Circum- neutral to ealeiphile. Central and southern Mo., scattered: St. Louis, Jefferson, Crawford, Boone, Jackson, and Greene counties. Cayaponia grandifolia (T. & G.) Small. w woods and borders of bayous. Cireumneutral. Southeastern Mo.: Mis- sissippi and New Madrid counties. Sicyos angulatus L. One-seeded Bur Cueumber. Low alluvial woods, rich ground at base of bluffs, fields, thiekets, and waste ground. Cireumneutral to caleiphile. Eastern, southern, and central Mo.: 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 653 Marion, Boone, Warren, St. Louis, Jefferson, Franklin, Bollinger, Crawford, Dent, Phelps, Pulaski, St. Francois, Mississippi, Reynolds, at coe Clay, Jaekson, Johnson, Hiekory, Webster, Jasper, and MeDonald eountie it lobata (Miehx.) T. & G. Wild Balsam-apple, Wild E os Vine. Thickets, dae alluvial and waste ground. Cireumneutral. General but scattered: Sullivan, Shelby, Boone, St. Charles, St. Louis, Harrison, Gentry, Dekalb, Jackson, E Johnson, Greene, and Jasper counties. Melothria pendula L. Rich woods and thickets. Cireumneutral. Southern Mo.: Stoddard, Dunklin, Butler, Barry, and McDonald counties. The watermelon (Citrullus vulgaris Sehrad.), muskmelon (Cucumis Melo L.), eueumber (Cucumis sativus L.), pumpkin (Cucurbita Pepo L.), and the gourd (Lagenaria vulgaris Ser.), are often found growing spontaneously on dumps, waste ground, and along roadsides; but since none of them show any tendency to self-seed or persist they eannot properly be eonsidered as additions to the state flora. Fam. CAMPANULACEAE Specularia perfoliata (L.) A. DC. Venus's Looking-glass. Prairies, fields, and waste ground. Cireumneutral. General. cae biflora (R. & S.) Fisch. & Me Fields, roadsides, and waste pnh Circumneutral. Southern Mo.: Bol- "is Dunklin, Ripley, Oregon, Iron, Dent, Wayne, Shannon, Texas, Laelede, Ozark, Christian, Taney, Barry, Jasper, and Newton counties. EUR leptocarpa (Nutt.) Gray. « cky or gravelly open ida and glades. Cireumneutral to oxylophile. Cent am and southwestern t. Louis, Boone, Benton, Jackson, Cass, St. Clair, Jasper, and Newton CODE Campanula americana L. Tall Bellflower. Thickets and borders of woods. Cireumneutral. General. ERA rotundifolia L. Bluebell, Harebell. Moist ledges along bluffs. Cireumneutral to caleiphile. Southeastern Mo., loeal: poss Co CAMPANULA RAPUNCULOIDES L. Creeping Bellflower. Fields and waste ground. Introduced in Warren Co. Fam. LoBELIACEAE Lobelia cardinalis L. Cardinal Flow wet woods, and borders of Me and streams. Cireumneutral. General. Lobelia siphilitica L. Blue Lobelia. Wet ledges along bluffs, and borders of ponds, sloughs, and small streams. Cireumneutral. General Lobelia ede: Miehx. Moist sandy open ground and open woods. Oxylophile. Southeastern Mo.: Dunklin and Ripley counties. [Vor. 22 654 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lobelia leptostachys A. DC. Open woods, glades, and prairies. Cireumneutral. Central and southern Mo.: Cole, Warren, St. Louis, Jefferson, Franklin, Washington, — Madison, Iron, Shannon, Dent, Phelps, Pulaski, Miller, Wright, Ozark, Polk, MN runs, Taney, Barry, Jackson, Jasper, Newton, and DE asintió Lobelia spicata Lam. Open woods and prairies. Cireumneutral General: Schuyler, Adair, "d Linn, Kno helby, Marion, Montgomery, Boone, Callaway, St. St. Louis, Reynolds, Oregon, Texas, Polk, Saline, Jackson, Bates, Vernon, = Jasper counties Lobelia spicata var. hirtella Gray. rairies and glades. Cireumneutral. Western Mo.: Nodaway, Jackson, Bates, Greene, Jasper, and Newton counties. Lobelia inflata L. Indian Tobacco Rocky open woods, thickets, dl alluvial ground along streams. Circum- neutral. General. Fam. COMPOSITAE Vernonia crinita Raf, Iro ed. Open woods, thickets, bn of sloughs, and gravel and sand bars along Ozark streams, Circumneutral to oxylophile. Central and southern Mo. south of a line from St. Louis, Warren, Montgomery, Boone, and Pettis counties to Clay Co. Vernonia fasciculata Michx, Prairies, meadows, alluvial soils along streams, and waste places in rich moist soil. Cireumneutral. General, commonest in northern and central Mo., and ap- parently absent from most of the Ozark region: Clark, Lewis, Marion, Pike, Lincoln, St. Charles, St. Louis, Butler, Shannon, Boone, Sullivan, Linn, Living- ston, Gentry, Nodaway, Holt, Andrew, Buchanan, Platte, Jackson, Cass, John- son, Henry, and Jasper counties Vernonia altissima Nutt. Thiekets, prairies, meadows, and waste ground, eommon in alluvial and low woods. Cireumneutral. General, but commonest in eastern, southern, and cen- tral Mo Vernonia altissima var. taeniotricha Blake, Rh. 19: s 1917. In similar situations to the typical form. Gene Vernonia missurica Raf. Vernonia illinoensis Gleason. (G). Meadows, prairies, moist open ground along streams, fields and waste places. Cireumneutral. General, but commonest in northern, central, and western Mo. Vernonia Baldwini Torr. Vernonia Baldwini var. parthenoides (Daniels) Rickett, Univ. Mo. Stud. Sei. Ser. 6: 81. 1931. Fields, prairies, meadows, and waste ground. Circumneutral. Eastern, southern, and central Mo. south and east of a line drawn from Marion, Boone, and Saline counties to Jackson Co. 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 655 Vernonia interior Small. Dry prairies, pastures, and waste ground. Circumneutral. General, Vernonia interior var. Drummondii (Shuttlw.) Mack. & Bush. Prairies and rocky open ground. Circumneutral. Scattered: St, Francois and Jackson counties. Vernonia Baldwini x crinita. Commonly found with the parent species. Rocky woods and glades and along streams. Caleiphile to cireumneutral. Central and southern Mo.: Jefferson, Washington, Crawford, Reynolds, Ripley, Dent, Shannon, Howell, Oregon, Gas conade, Texas, Dallas, Webster, Polk, Pettis, Stone, Clay, and Jasper counties. Vernonia altissima x missurica. Occasionally found with the parent species. Scattered: Gentry, Dekalb, Boone, Crawford, Ripley, and Laclede counties. Vernonia crinita x fasciculat Rarely found with the m species. St. Louis Co. Vernonia crinita x interior. Rarely found with the parent species. Phelps Co. Vernonia crinita x missurica. Occasionally found with the parent species. Crawford and Dent counties. Vernonia fasciculata x interior Occasionally found with (lis parent species. Northwestern Mo.: Buchanan and Andrew counties. Vernonia interior x missurica Sometimes found with iis parent species. Clark, St. Louis, and Crawford counties. Vernonia altissima x crini Rarely found with me Bst speeies. Crawford, Cedar, and Hickory counties. Elephantopus carolinianus Willd. Elephant’s-foot. Open woods and thiekets. Circumneutral. General. Eupatorium falcatum Michx. See Rh. 22: 62-68. 1920. Eupatorium purpureum of auth. in part, not L. (G), (B & B). Eupatorium Holzingeri Rydb. Bi Moist denn egi. and margins of streams and sloughs. Circumneutral. General and c This has E confused with E. maculatum, which is of more diu range nd ce not reach Mo. Eupatorium purpureum L. See Rh. 22: 62-67. 1920. Low open woods and moist open ground. Circumneutral. Northern and east- ern Mo.: Livingston, Marion, Butler, and Scott counties. Eupatorium cuneifolium Willd. var. semiserratum (DC.) Fernald & Griscom, Rh. 37: 179. 1935. Eupatorium semiserratum DC. (G). Swamps, wet prairies, and open woods. Cireumneutral. Southeastern Mo.: Dunklin and Butler counties, and also collected in St. Louis Co. where perhaps introduced. [Vor. 22 656 ANNALS OF THE MISSOURI BOTANICAL GARDEN Eupatorium serotinum Michx. rairies, fields, thiekets, and waste ground. Cireumneutral. General. Eupatorium sessilifolium L. Upland Boneset. y open woods, bluffs, and banks. Circumneutral to caleiphile. Eastern, eentral, and southern Mo.: n, Ralls, Boone, Jefferson, St. Francois, Iron, Reynolds, Crawford, Shannon, tad Pulaski, Camden, Laclede, Greene, Taney, Barry, and McDonald counties. Eupatorium altissimum Prairies, rocky hills, amd thickets. Circumneutral. General. Eupatorium perfoliatum L. Boneset, Thoroughwort. ow wet woods, and along sloughs and prairie streams. Oxylophile. General. Eupatorium perfoliatum var. cuneatum Engelm. Occasionally found with the typical form. Seattered: Marion, Ralls, St. Louis, Perry, Jackson, and Greene counties. Eupatorium urticaefolium Reichard. White Snakeroo Rich woods, thickets, and along base of bluffs. u General. Eupatorium urticaefolium var. villicaule Fernald. Oeeasionaly found with the typical form. Seattered: Putnam, Boone, St. Louis, Jefferson, Oregon, Ozark, Greene, Christian, Lafayette, and Clinton counties. Eupatorium incarnatum Walt. Low woods and swamps. Cireumneutral. Southeastern Mo.: Dunklin Co. Eupatorium coelestinum L. Mist-flower. wampy woods, and wet banks of streams and bayous. Circumneutral. East- ern, central, and southern Mo., south and east of a line from Shelby, Boone, and Saline eounties to Jackso s Ok Mikania dere (L.) Willd. Climbing Hemp-weed. Swampy woods and borders of bayous. Cireummeutral Southeastern Mo.: Seott, peara New Madrid, Stoddard, Dunklin, and Butler counties. Brickellia grandiflora (Hook.) Nutt. Tassel Flower. Rocky woods and along shaded bluffs. Calciphile to cireumneutral. Central and southern Mo.: St. Louis, Reynolds, Shannon, Texas, Osage, En Pulaski, Miller, Camden, Ozark, Taney, Stone, Henry, and Barry eounti Kuhnia eupatorioides L. False Boneset. Ro "d open woods, prairies, and thickets. Circumneutral to oxylophile. Genera Kuhnia eupatorioides var. corymbulosa G. Occasionally found with the typical FR Seattered: Carroll, Boone, Seott, Atchison, Holt, Jackson, Henry, and Greene counties, Kuhnia eupatorioides var. angustifolia Raf. — ledges and glades. Cireumneutral to oxylophile. Central and southern Mo.: St. Francois, Dent, Shannon, Oregon, Phelps, Maries, Pulaski, Laclede, Greene, Shannon, Texas, Ozark, Barry, Jackson, Jasper, and MeDonald suidiin. Liatris squarrosa Willd. Blazing Star. iatris hirsuta Rydb. (R). 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 697 Rocky open woods and glades, and ledges along bluffs. Oxylophile to cireum- neutral General, but eommonest in the Ozark region: Mercer, Linn, Adair, Macon, Randolph, Boone, Monroe, Audrain, Pike, Montgomery, Lincoln, St. Louis, Stoddard, Iron, Reynolds, Shannon, Dent, Crawford, 15 Pulaski, Laclede, Dallas, Saline, Pettis, Henry, Hickory, Greene, Christia n, Stone, Barry, ackson, Cass, Vernon, Jasper, Newton, and McDonald counti: Liatris squarrosa var. intermedia (Lindl.) DC. Liatris glabrata Rydb., in part (R). Rocky prairies and E Won to eireumneutral. Eastern, central, and southern Mo.: n, Boone, St. Louis, Crawford, Dunklin, Iron, Shan Dent, LM Greene, pis Stone, Taney, Jackson, Barry, and ud counties Liatris le Miehx. Liatris glabrata Rydb., in part (R). Rocky open woods, prairies, and glades. Circumneutral. Eastern, central, and southern Mo.: Lewis, Pike, Warren, Montgomery, St. , Jeffer ranklin, Washington, Ste. Genevieve, Reynolds, Carter, d rand pote 4 Den ab Crawford, Maries, Phelps , Pulaski, Texas, Howell, Wright, Douglas, Ozark, Taney, Greene, Stone, Barry, and MeDonald counties. Liatris acidota Engelm. & Gra Liatris punctata of auth., aot Hook. (G), (B & B). Lacinaria angustifolia Bush. Loess hills, prairies, glades, and open woods. Circumneutral to calciphile. Western Mo.: Atchison, Holt, Jackson, Barry, and Taney counties. Liatris scariosa Willd. Gay Feather. iatris sphaeroidea Michx. (R). Liatris aspera (Michx.) resis ys Rocky prairies, meadows, glades, and bald knobs. Calciphile to cireumneutral. General. Liatris pycnostachya Mic Prairies, meadows, a ae open ground. Circumneutral. General. Liatris spicata (L.) Willd Rocky open woods and prairies. Cireumneutral. Southeastern Mo., local: Oregon Co. E et re Dunal. Gum Plan es, low alluvial and waste ground. m to circumneutral. Eastern Ne re I an. Marion, St. Louis, Perry, Carroll, and Jackson counties. ud ee var. serrulata (Rydb.) Steyermark, Ann. Mo. Bot. Gard. 21: a d fields and dd ies: introduced from farther west. Circum- neutral n ealeiphile. Clark and Boone counties. Grindelia squarrosa var. nuda (Wood) ps Sometimes found with the typical form. St. Louis and Jackson counties. Grindelia lanceolata Nut ocky prairies, bins and bald knobs, and also sometimes introduced along [Vor. 22 658 ANNALS OF THE MISSOURI BOTANICAL GARDEN railways or in fields. Caleiphile. Central and southern Mo.: St. Louis, Jeffer- son, Butler, Camden, Wright, Douglas, Ozark, Henry, DN Taney, Barry, Vernon, Barton, Jasper, Newton, and MeDonald countie Grindelia lanceolata f. latifolia Steyermark, Ann. Mo. Bot. Gard. 21: 515. 1934. Rarely oceurring with the species. Southwestern Mo., local: Stone Co. Gutierrezia dracunculoides (DC.) Blake, Contr. U. S. Nat. Herb. 22: 592. 1924. ugust piola dracunculoides (DC.) Nutt. (G), (B € B). Dry upland prairies. Circumneutral to calciphile. Western Mo.: Atchison, Jackson, Cass, Cedar, Greene, Jasper, Newton, and MeDonald counties, and also introduced in St, Louis Co. —— subaxillaris (Lam.) Britton & Rusby. piero and waste ground. Calciphile to cireumneutral. Northwestern on Co., and also introduced in Jaekson Co. Chrysopsis camporum Greene. Golden Aster. Chrysopsis villosa of auth., not Nutt. (G), (B & B). Sandy open ground. Oxylophile. Eastern Mo., south of the Missouri River: St. Louis, en Ste. Genevieve, Seott, Mississippi, Stoddard, Dunklin, and Wayne eountie Chrysopsis sid Nutt. ocky or sandy prairies and open banks. Oxylophile. Southern Mo.: Dunk- lin and Jasper counties, Chrysopsis Berlandieri Greene. Chrysopsis villosa var. canescens (T. & G.) Gray. Loess hills, Caleiphile to eireumneutral. Northwestern Mo.: Holt Co. — ciliatus (Nutt.) DC. es, glades, open alluvial and waste ground. Calciphile. Western Mo.: Pi 5 — and Stone counties. ze petiolaris Ait. cky prairies, — and hed open woods. Circumneutral to oxylophile. Central and southern . Louis, Jefferson, Franklin, Ste. A St. Francois, Madison, Tron, aen Butler, Shannon Desk; Crawford, eonade, Wright, Taney, Greene, Stone, Jackson, Jasper, "p Newton counties. Solidago petiolaris var. Wardii (Britton) Fernald. Rocky open woods and thickets. Oxylophile. East-central and southern Mo.: St. Louis, Jefferson, Crawford, Callaway, Montgomery, Boone, inv Phelps, Pulaski, Henry, Greene, Stone, Jasper, Newton, and MeDonald counti Solidago caesia L. Blue-stemmed Goldenrod. ocky woods, bluffs, and ledges. Cireumneutral to oxylophile. Southern Mo.: Perry, Cape Girardeau, Bollinger, Iron, Wayne, Ripley, Reynolds, Shannon, Oregon, Texas, Howell, Douglas, Oni; and Barry counties. — latifolia L. 1 rocky woods and shaded bluffs. Circumneutral. General but commonest in mdi and southern Mo.: Putnam, Marion, Ralls, Monroe, Boone, Callaway, Warren, Lineoln, St. Charles, St. Louis, Seliseion. Franklin, Cape Girardeau, 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 659 Bollinger, Madison, Iron, Wayne, Carter, Shannon, Oregon, Morgan, Camden, Dallas, Ozark, Taney, Stone, Barry, Jackson, Lafayette, Pettis, Hickory, Greene, and Jasper counties. Solidago Lindheimeriana Scheele. Rocky woods and bluffs. Cireummeutral East-central and southern Mo.: Jefferson, Franklin, Carter, Barry, and McDonald counties. Solidago hispida Muhl. Roeky open woods and ledges. Oxylophile. East-central and southern Mo.: St. Louis, Jefferson, Franklin, Gasconade, Crawford, Ste. Genevieve, Bollinger, Madison, Iron, Reynolds, Wayne, Carter, Ripley, Oregon, Shannon, Dent, Texas, Howell, Douglas, Ozark, Taney, Stone, Barry, Jasper, Newton, and MeDonald counties Solidago HEN Nut Rocky open Des thiekets, prairies, and meadows. Circumneutral. General but scattered. . Solidago speciosa var. angustata T. & G. Frequently found with the typical form. Southern and central Mo. and locally north in Worth, Putnam, and Buchanan counties. Solidago patula Muhl. Swampy open ground. Circumneutral. Southeastern Mo.: Bollinger, Iron, and Scott counties. Solidago arguta Ait. Rocky open woods and bluffs. Oxylophile to circumneutral. Southern Mo.: Ste. Genevieve, Madison, Reynolds, Carter, Shannon, Oregon, Texas, Howell, Ozark, Taney, Barry, Jasper, Newton, and McDonald counties. Solidago arguta var. Boottii ( Hook.) d & Steyermark, comb. nov. cons Boottii Hook. P (B 6 Dr en woods and thickets. GR to oxylophile. Southern Mo.: Ste. RUP Reynolds, Ripley, Shannon, Texas, Howell, Douglas, Ozark, Barry, Jasper, Newton, and MeDonald counties. Solidago juncea Ait. Early Goldenrod. Open woods, thickets, and along prairie streams. Circumneutral to oxylophile. Central and southern Mo.: Boone, St. Louis, Seott, Iron, Carter, Shannon, Taney, Barry, Newton, and McDonald counties. Solidago os var. scabrella (T. € G.) Gray. Occasionally found with the typical form. Southwestern Mo.: Taney, Barry, and Newton countie Solidago suaveolens Blot Ed Rh. 21: 70. 1919. Fragrant Goldenrod. ic od si Ait. (G), (B & B). Rocky o open Lh and thiekets. Oxylophile. Southeastern Mo.: Scott, Dunklin, ra and Ripley counties. Solidago microphylla Engelm. Rocky woods and bluffs. Cireumneutral Southern Mo., local: Ozark Co. Solidago ulmifolia Muhl. Open woods, thiekets, and alluvial banks. Cireumneutral. General. [VoL. 22 660 ANNALS OF THE MISSOURI BOTANICAL GARDEN Solidago rugosa Mill. Rough-leaved Goldenrod. Rocky prairies, glades, and open woods along streams. Cireumneutral to oxylo- phile. Central and southern Mo.: St. Louis, Jefferson, Ste. Genevieve, Scott, Iron, Wayne, Butler, Dunklin, Texas, Howell, Jackson, Greene, Jasper, and New- ton counties. Solidago rugosa var. "dp (Ait.) Fernald, Rh. 17: 7. 1915. Solidago aspera In similar bn to the typieal form. East-central and southeastern Mo.: Lincoln, Perry, Iron, Wayne, Butler, Dunklin, and Shannon counties. Solidago glaberrima Martens. Solidago missouriensis of auth., not Nutt. (G). Open woods, thickets, and prairies. Cireumneutral. General. Solidago glaberrima var. moritura (Steele) Palmer & Steyermark, comb. nov. Solidago moritura Steele (R). Thickets, prairies, limestone hills, and loess mounds. — m to circum- neutral. Western Mo.: Atchison, Holt, Henry, and Vernon countie Solidago Gattingeri Chapm. oeky open ground, Lao and bald knobs. Calciphile. East-eentral and southern Mo.: St. Lou "aat Franklin, Camden, Dallas, Ozark, Taney, Barry, and McDonald sedis Solidago nemoralis Ait. Gray I Open woods, thickets, fields, prairies, and waste ground. Oxylophile. General. iren nemoralis var. longipetiolata (Mack. & Bush) Palmer & Steyermark, mb. n "Solidago longipetiolata Mack. & Bush (R). Rocky open woods, glades, prairies, and loess hills, Oxylophile to circum- neutral. General but scattered: Clark, Boone, St. Louis, Jefferson, St. Francois, Iron, annon, Maries, — Worth, Atchison, Holt, Beds, Jackson, Jasper, and Barry counties Solidago radula Nutt. Rocky woods, bluffs, and glades. Cireumneutral to calciphile. Eastern, cen- tral and southern Mo. south and east of a line drawn from Marion, Monroe, Baline, and Henry counties to Cedar and Jasper counties. Solidago altissima L. Fields, prairies, thickets, and waste ground. Cireumneutral. General. This is one of the commonest goldenrods in the state, and it has often been confused with Solidago canadensis, the typical var. of which is not found in Mo Solidago altissima var. procera (Ait.) Fernald. Occasionally found with the typical form. idago canadensis L. var. gilvocanescens Rydb. s and prairies. Circumneutral. Seattered: Linn, Pike, Boone, Saline, Pe and Greene counties. Solidago serotina Ait. Moist open no and thickets, prairie swales, and banks of streams. Cir- cumneutral. Gene 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 661 Solidago serotina var. gigantea (Ait.) Gra Oeeasionally found with the typical ER Seattered: St. Louis, Dunklin, Boone, Jackson, and Cass counties. Solidago rigida L. Prairies, thiekets, and rocky open ground. Cireumneutral to calciphile. General. Solidago Drummondii T. & G. Dry limestone or dolomite bluffs and ledges. Calciphile. Eastern and middle Mo., mostly south of the Missouri River: Monroe, Boone, laway, Montgomery, St. Charles, St. Louis, Jefferson, Franklin, Gasconade, Crawford, Washington, St. Francois, Ste. Genevieve, Perry, Bollinger, Madison, Iron, Reynolds, Carter, Oregon, Shannon, pin Er Moniteau, Pulaski, Laelede, Texas, Howell, Douglas, and Ozark e Solidago graminifolia sy Xu Flat-topped Goldenrod. Euthamia glutinosa Rydb. (R) Fields and prairies. Circumneutral. General and common. Solidago gymnospermoides (Greene) Fernald. Euthamia gymnospermoides Greene (B & B). Rocky or upland prairies. Circumneutral to ealeiphile. Middle and western Mo.: Audrain, Boone, Miller, Pettis, Greene, Daviess, Dekalb, Jackson, Cass, Vernon, Barton, Jasper, Newton, and MeDonald counties. Er leptocephala T. & G. open ground. Oxylophile. Southeastern Mo.: Mississippi, Dunklin, and me counties. Bellis integrifolia Michx. Western Daisy. ocky open woods and glades. Calciphile. Southwestern Mo.: Taney and MeDonald counties. Chaetopappa asteroides DC. Rocky open woods, prairies, and glades. Oxylophile. Southwestern Mo.: Vernon, Jasper, and Newton counties. Boltonia asteroides (L.) L'Her. False Starwort Moist open woods and thickets. Cireumngetral: General. Forms of this have been confused with Boltonia diffusa, which is not known in Boltonia asteroides var. decurrens (T. & G.) Engelm. Frequently found with the typical form. General but seattered. Boltonia latisquama Gray. airie swales and borders of streams and sloughs. Circumneutral North- ern, central, op western Mo.: ped ii. Shelby, Monroe, Ralls, Pike, A udrain, Ripley, Boone, Randolph, Macon, Linn, Carroll, Livingston, Chariton, Caldwell, Bo M Lafayette, Pettis, Dons Henry, Vernon, Greene, and Jasper counties. Aster paludosus Ait. Prairies and along railroads. Cireumneutral to oxylophile. Southwestern Mo. and loeally east in Marion Co. where probably introduced: Marion, Barton, Jasper, Newton, and Barry counties. [Vor. 22 662 ANNALS OF THE MISSOURI BOTANICAL GARDEN Aster furcatus Burges Shaded rocky mdi and bluffs of streams. East-central and southern Mo.: Jefferson, Shannon, and Texas counties, Aster oblongifolius Nutt. oeky prairies, open slopes, and glades. Caleiphile to eireumneutral. General. Aster oblongifolius var. rigidulus Gray. Frequently found with the typical form, Seattered: St. Louis, Jefferson, Atehison, Holt, Henry, Greene, Ozark, wi Jasper, and McDonald counties. Aster on L. New England Ast Thickets, prairie swales, and moist open a along streams, Circumneutral, General lat scattered. Aster sericeus Vent. Roeky prairies, glades, and loess hills. Is us mdi to eircumneutral. Central, southern, and northwestern Mo.: St. Jefferson, Was —€— Iron, Dent, Shannon, Oregon, Texas, Howell, Died im Dallas, iekory, ar, Greene, aney, Stone, Barry, Atehison, Holt, Jackson, Cass, and MeDonald pomis Aster patens Ait. Purple Daisy. Rocky open woods, thickets, and glades. Circumneutral to oxylophile. General. Aster patens var. gracilis Hook. Occasionally found with the typical form. Jasper and MeDonald counties. Aster anomalus Engelm. Rocky open woods and thiekets. Oxylophile to eireumneutral. Eastern, cen- tral, and southern Mo., south and east of a line from Lewis, Boone, and Henry to Jasper Co. Aster azureus Lindl. Blue Devil. Rocky open woods, borders of prairies, and glades. Calciphile to cireum- neutral. Genera Aster azureus x lae Sometimes occurring with the parent species. oliin and dry open ground. amor Scattered in northern and central Mo.: Monroe, Adair, Ran- dolph, Putnam, Daviess, and Caldwell po Aster cordifolius L. Heart-leaved Aste Moist alluvial woods, and along shaded bluffs. Cireumneutral to oxylophile. General. Aster cordifolius var. polycephalus Porter. Occasionally found with the typical form. Boone Co. Aster sagittifolius Wedemey Rocky woods and gla ing "emendi to oxylophile. General. Aster Drummondii Lindl. Aster hirtellus Lindl. (R). Rocky open woods and thiekets. Circumneutral. General. Aster turbinellus Lindl. Rocky open woods, thickets, and glades. Circumneutral to oxylophile. East- ern, eentral, and southern Mo., south and east of a line from Putnam, Maeon, Pettis, and Henry counties to Jasper Co. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 663 Aster laevis L. Rocky open woods, thickets, glades, prairies, and loess hills. Cireumneutral to calciphile. General Aster concinnus Willd. open woods, prairies, and glades. Calciphile to eireumneutral. Central and southern Mo., seattered: St. Louis, Jefferson, Franklin, Maries, Shannon, Ozark, Cooper, Jackson, Jasper, and Newton counties Aster polyphyllus Willd. Rocky open ground. Cireumneutral to calciphile. East-central Mo., local: St. Louis Co. Aster pilosus Willd. See Rh. 32: 139. 1930. White Heath Aster. Aster ericoides var. villosus T. & G. (G). Aster ericoides of auth. in part, not L. (B & B). Fields, prairies, and rocky open ground. Cireummeutral to oxylophile. General Aster pilosus var. demotus Blake, Rh. 32: 139. 1930. Aster ericoides of auth., not L. (G), (B & B). Rocky open woods, thickets, and prairies. Circumneutral to oxylophile. General. Aster Ir var. platyphyllus (T. & G.) Blake, Rh. 32: 139. 1930. Aster ericoides var. platyphyllus T. & G. (G). iba found with the typical form, Seattered: Ralls, St. Louis, and Jackson counties. Aster ericoides L. See Rh. 32: » 1930, and Rh. 80: 227. 1928. Aster multiflorus Ait. (G), (B Aster multiflorus var. exiguus bun Rocky open woods and prairies. Cireasa: Northern, central, and west- ern Mo., apparently absent from most of the Ozark region: Clark, Lewis, Marion, Ralls, Pike, Lincoln, St. Charles, St. Louis, Warren, Montgomery, Audrain, Mon- roe, Shelby, Knox, Scotland, Schuyler, Adair, Macon, Randolph, Boone, Howard, Chariton, Tini, ullivan, Putnam, Mercer, Grundy, is es Carroll, Saline, Cooper, Morgan , Pettis, Greene, Hay, Johnson, Caldwell, Daviess, Harrison, Worth, Gentry, Dekalb, Clinton, Clay, Jackson, Bates, J ami Platte, Buchanan, Andrew, Nodaway, Holt, and Atchison counties. Aster ericoides var. prostratus (Ktze.) Blake, Rh. 32: 138. 1930. Aster exiguus (Fernald) Rydb., not pa eg var. exiguus Fernald. Aster multiflorus var. pansus Blake, . 30: 227. Prairies, dry open ground, along id and na: or t gravelly ES along streams. Circumneutral. Northern, central, and weste : Knox, Macon, Marion, Lineoln, St. Louis, Cole, Montgomery, Randolph, rd Linn, 2 bun. Jackson, and Jasper counties. Aster parviceps (Burgess) Mack. & Bush. See Rh. 11: 59. 1909. Aster depauperatus var. parviceps (Burgess) Fernald (G) Prairies and open woods. Eastern, central, and southwestern Mo.: Marion, Ralls, Pike, Audrain, St. Louis, Bollinger, Warren, Montgomery, Callaway, Boone, Jackson, and J asper counties. (Vor. 22 664 ANNALS OF THE MISSOURI BOTANICAL GARDEN Aster parviceps x pilosus var. demotus. Sometimes occurring with the parent species. Circumneutral. Northern Mo.: Knox, Marion, Putnam, and Grundy counties, Aster amethystinus Nutt. Prairies. Circumneutral. Central Mo., local: Randolph Co. Probably a hybrid between Aster praealtus and Aster novae-angliae with which species it was found growing. Aster commutatus (T. & G.) Gray. White Prairie Ast ields and prairies. Circumneutral. obli ‘St. Louis and Jackson counti Aster JL" Lam. var. subdumosus Wiegand, Rh. 30: 171. 1998, Frost Flow poni vimineus of auth. in part (G), (B & B). Open woods and prairie swales. Circumneutral. General but scattered: Mon- roe, Warren, St. Louis, Jefferson, Seott, Stoddard, Dunklin, Butler, Wayne, Audrain, Randolph, bien. Henry, Greene, and Jasper counties, "This has been confused with Aster dumosus and varieties which are not known to occur in issouri. Aster tetra (Lindl.) T. & G. en. loess hills. Cireumneutral to ealeiphile. Northwestern Mo., iosal; Atchi Aster pantotrichus Blake, Jour. Wash. Acad. py 21: 327. 1931. Aster missouriensis Britton, not Ktze. (B Jé Aster lateriflorus var. thyrsoideus (Gray) -ipi (G), in par Rocky open woods. Cireumneutral to oxylophile, Eastern, cum and south- ern Mo.: Ralls, Pike, Lincoln, St. Louis, Ste. Genevieve, Iron, Taney, Boone, and Jackson counties, Aster lateriflorus (L.) Britton var. pendulus (Ait.) Burgess. See Rh. 30: 173. 1928, Aster lateriflorus of auth. in part (G), (B & B). Low wet woods and borders of ponds, sloughs, and swales. Circumneutral. General and common. Typical Aster lateriflorus is of more northern and eastern range and does not reaeh Missour Aster paniculatus Lam. Tall White Aster. w wet woods and prairie swales. Circumneutral. General. Aster paniculatus var. bellidiflorus ( Willd.) Burgess. Occasionally found with the typical form. Boone and Jackson counties, Aster Sr var. simplex (Willd.) Burgess. Oeeasionally found with the typieal form. Boone, Randolph, and Jasper Pine Aster praealtus Poir. See Rh. 35: 21. 1933. Willow-leaved Aster. Aster salicifolius Ait., in part (G), (B & B). Wet meadows, piedi. roadsides, and borders of small streams. Cireum- neutral. General. Aster praealtus var. subasper (Lindl.) Wiegand, Rh. 35: 24. 1933. Aster salicifolius var. subasper (Lindl.) Gray (G). 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 665 Oceasionally found with the typical form. Mississippi, Boone, and Jackson counties. Aster interior Wiegand, Rh. 35: 35, 312. 1933. Michaelmas Daisy. Aster Tradescanti of auth. in part, not L. (G), (B 4 B). Low wet woods, thickets, and prairie swales. Circumneutral. General. Aster linariifolius L. open woods and glades. Oxylophile. East-central and southern Mo.: St. xod Jefferson, Franklin, Ste. Genevieve, Perry, Bollinger, Madison, Iron, Wayne, Ripley, Carter, Reynolds, Oregon, Dent, Shannon, Crawford, Maries, Phelps, Pulaski, Laclede, Texas, Howell, Douglas, Ozark, Greene, Taney, Barry, Jasper, Newton, and MeDonald counties. Aster ptarmicoides T. & G. Rocky open woods and glades. Calciphile. Central and southern Mo.: Jef- ferson, Washington, Crawford, Dent, Iron, Shannon, Phelps, Pulaski, Jackson, Greene, Barry, Newton, and MeDonald counties. Aster lateriflorus var. pendulus x pantotrichus. Occasionally found with the parent species. St. Louis Co. Aster lateriflorus var. pendulus x praealtus. Occasionally found with the typical form. St. Louis Co. Erigeron pulchellus Michx. Robin’s Plantain ocky open woods and thickets. Oxylophile to circumneutral. General. Erigeron philadelphicus L. Fields, open woods, and waste ground. Cireumneutral. General. Erigeron annuus (L.) Pers. Daisy Fleabane. Fields, prairies, and waste Su Indifferent. General. Erigeron ramosus (Walt.) B ocky prairies, fields, x ds Calciphile to cireumneutral. General. Erigeron ramosus var. Beyrichii (T. € G.) Trelease. Occasionally found with the typical form. Jasper Co. Erigeron tenuis T. & G. inne: fields, and waste ground. Circumneutral. Central and southern Mo., scattered: St. Louis, Jasper, Newton, and McDonald countie Erigeron canadensis L. Fields, roadsides, and waste ground. Indifferent. General and common. pages divaricat atus Michx. Dwarf Fleaban Prairies, fields, glades, aad waste ground. enm General. Pluchea foetida (L.) DC. Marsh Fleabane. Swampy open woods and borders of bayous. Cireumneutral. Southeastern Mo.: Butler Co. Pluchea petiolata Cass. Low wet woods, swamps, and borders of ap d and bayous. Cireumneutral. Southern Mo.: Bollinger, Scott, Mississippi, New Madrid, Pemiscot, Dunklin, Butler, "Wayne, Ripley, Oregon, Howell, Taney, HE and Jasper que and also introduced in Jaekson Co Antennaria plantaginifolia (L.) Richards. Ladies? Tobacco, Pussy's Toes. Rocky open woods and thickets. Oxylophile. General. [Vor. 22 666 ANNALS OF THE MISSOURI BOTANICAL GARDEN Antennaria fallax Greene. Indian Tobacco. Rocky open woods, glades, and prairies. Oxylophile. General. This is mon species in Missouri, and it has often been confused with A. an Antennaria calophylla Greene. Antennaria ampla Bush. Rocky open woods and prairies. Oxylophile to eireumneutral. General but seattered: Shelby, sso e Audrain, Shannon, Ripley, Greene, Christian, Taney, Jasper, and Newton countie Antennaria neglecta Greene. Antennaria longifolia Greene. Prairies and meadows. Cireumneutral. Northern, central, and western Mo.: Sehuyler, Scotland, Knox, Shelby, Lewis, Marion, Ralls, os Lineoln, Audrain, Boone, Randolph, Macon, Adair, Sullivan, Linn, Mercer, Grundy, Daviess, Platte, Moniteau, Morgan, Pettis, Hickory, Heals, Cedar, Jackson, latas and Jasper counties. Anaphalis margaritacea (L.) Benth. £ Hook. Pearly Everlasting. Fields and waste ground. Introduced in Boone Co. Gnaphalium purpureum L. Cudweed. Fields, prairies, iene thickets. Cireumneutral to oxylophile. General. Gnaphalium obtusifolium L. See Rh. 20: 71. 1918. Everlasting. naphalium polycephalum Miehx. (G). Rocky prairies, fields, and thickets. Cireumneutral to en General. Gnaphalium obtusifolium var. micradenium Weatherby, Rh. 25: 22. 1923. Rocky open woods. Oxylophile. Southeastern Mo., =. ve region: Ste. Genevieve and Shannon counties. INULA HELENIUM L, Elecampane Wet open ground and waste Jibi Scattered: Monroe, Boone, and Jackson counties, and also E from Greene Co. Polymnia canaden Rich rocky an and talus slopes. Cireumneutral to ealeiphile. Eastern, central, and southern Mo., south and east of a line drawn from Marion, Warren, and Boone counties to Jackson Co, — canadensis var. radiata Gray. Commonly as with the typical form. Scattered in eastern, central, and diea Mo.: Clark, Marion, St. Louis, Carter, Dent, Phelps, Pulaski, Shannon, Taney, D Ma Boone, Jackson, and MeDonald counties. Polymnia Uvedalia y var. genuina Blake, Rh. 19: 47. 1917. Leaf-cup Rich woods and along bluffs, Calci ipid. Scattered in central and southern ne and iei counties, en Uvedalia var. densipilis Blake, Rh. 19: 48. 1917. Rıch woods, gp n and along bluffs. Caleiphile to cireumneutral. Centr and southern Mo.: Boone, St. Louis, Jefferson, Dunklin, Butler, Ripley, Shan- non, Ozark, Barry, Der Newton, and McDonald counties. Silphium laciniatum L. Compass Plant. Prairies and glades. Circumneutral to calciphile. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 667 Silphium terebinthinaceum Jacq. Prairie Dock. Prairies, glades, and bald knobs. Caleiphile to eireumneutral. Eastern, cen- tral, and southern Mo., south and east of a line drawn from Marion, Boone, Ben- on Greene counties to Barry Co pM Asteriscus L. Starry Rosin-weed. Roc woods and glades. Cireumneutral to oxylophile. Southern Mo.: uis. d Ripley, Oregon, Shannon, Texas, Howell, Douglas, Ozark, Stone, and Barry counties Silphium integrifolium Michx. Rosi Roeky open woods, glades, and prairies. Cireumneutral to ealeiphile. General. Silphium perfoliatum L. Cup Plant. Moist prairies, thickets, and borders of streams and ponds. Circumneutral. General. Berlandiera texana DC. Rocky open woods and thickets. Circumneutral to calciphile. Southern Mo.: Phelps, Pulaski, Carter, Dent, Shannon, Oregon, Howell, Douglas, Ozark, Taney, Stone, Greene, Lawrence, Barry, Jasper, Newton, and McDonald counties. Engelmannia pinnatifida T. & G. Waste ground and along railroads. Introduced from farther west. Jack- son Co. Parthenium integrifolium L. Rocky woods, thickets, and glades. Calciphile to circumneutral. General, Parthenium hispidum Raf. Parthenium repens Eggert (G). Prairies and glades. Caleiphile. Central and southern Mo.: St. Louis, Jeffer- son, Franklin, St. Francois, Iron, Washington, Crawford, Dent, Shannon, Oregon, T , Phelps, Maries, Pulaski, Miller, Camden, Laclede, Dallas, Wright, Web- ster, Douglas, Ozark, Taney, Stone, Greene, Polk, Hickory, Cedar, Barry, Jasper, Newton, and McDonald counties. Parthenium Hysterophorus L. Santa Maria. Waste ground and along railroads. Introduced in Jackson Co. Iva ciliata Willd. Marsh Elder. Prairies, fields, and waste ground. Circumneutral to oxylophile. General but scattered, and apparently ti from much of the Ozark region: Clark, Linn, o arle sissippi, New Madrid, "^ pus nade, Miller, Morgan, Saline, Livingston, Clay, Jackson, laria Belt Greene, Bates, Vernon, Barton, Jasper, and Newton counties. Iva xanthifolia Willd. Fields, alluvial and waste ground. Indifferent. Northwestern and west-cen- tral Mo.: Atchison, Buchanan, and Jackson counties. Introduced from the Northwest. Ambrosia bidentata Michx. Rocky prairies, fields, and waste ground. Circumneutral to oxylophile. General. [Vor. 22 668 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ambrosia bidentata x trifida. Rarely occurring with the parent species. Local: Morgan Co. Ambrosia trifida L. Horse-weed. Ambrosia striata Rydb. (R). Alluvial fields, borders of woods, and waste ground. Circumneutral, General and common Ambrosia trifida var. integrifolia € T. & G. Sometimes found with the typical for Ambrosia artemisiaefolia L. var. elatior (Lo) Dese. See Rh. 37: 185. 1935. Ragweed, Roman Wormwood mbrosia artemisiifolia of auth., not L. (G), (B & B). Fields and waste ground. Indifferent. General and common. Ambrosia artemisiaefolia var. elatior f. villosa Fernald & Griscom, Rh. 37: 185. 1935 Commonly oceurring with the preceding variety. General. Ambrosia psilostachya DC. oeky prairies, pastures, and glades. Cireumneutral to oxylophile. Northern, central, and western Mo., absent from most of the Ozark region: Clark, Marion, Pike, Montgomery, St. ts , St. Francois, Randolph, Boone, Carroll, — Atebion; Holt, Andrew, ra Dekalb, Clinton, Jackson, Barton, and Jas per counties. Ambrosia aptera DC. Introduced in waste ground. St. Louis Co. Xanthium acerosum Greene. Cockl k Introdueed in waste ground. "edt Co. Xanthium italicum Mor. Xanthium — Greene d Xanthiw mmune Britton (G), (B & B). Fields, ci and waste ground. ea General. Xanthium speciosum Kearney. Cultivated fields and waste ground. Indifferent. Western Mo., scattered: Jackson, Cass, and Jasper counties. Xanthium inflexum Mack. € Bush. Low eultivated ground. Indifferent. Western Mo: Jackson Co. Xanthium echinatum Murr. Cultivated and waste ground. Scattered: St. Louis, Jefferson, Jackson, and Jasper counties. Xanthium globosum Shull, Bot. Gaz. 59: 482, figs. 1, 4, 5, 7. 1915. Low alluvial fields, roadsides, and waste ground. Indifferent. Scattered: Randolph, Cass, and Jasper counties. XANTHIUM SPINOSUM Fields and waste ae | Indifferent, Eastern Mo.: St. Louis Co. Xanthium pennsylvanicum Wallr. Cultivated fields, roadsides, and waste ground. Indifferent. General. Xanthium chinense Mill. Xanthium canadense of auth., not Mill. (G). 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 669 Fields, roadsides, waste and alluvial ground, often in moist soil along ponds, sloughs, streams, and sw open woods. Indifferent. General: Schuyler, Sullivan, Boone, Race dic Warren, St. Charles, St. Louis, Mississippi, New Madrid, Jefferson, Dent, Maries, Pettis, Linn, Jackson, Greene, Jasper, and MeDonald counties. Xanthium chinense x pennsylvanicum. Oceasionally found with the parent species. Knox and St. Louis counties. Xanthium oe x pennsylvanicum Oceasionally foi with the vient species. Montgomery and St. Louis Won Hands scabra Dunal. Rough Ox-eye. Rocky woods and thiekets. Cireumneutral. General. Heliopsis helianthoides (L.) Sweet. Ox-eye. R s and thickets. ena Eastern and middle Mo. Eclipta alba i Hassk. Moist fields, open ground, and borders of sloughs and ponds. Circumneutral. General, but commonest along the "n rivers. Rudbeckia triloba L. Brown-eyed Sus Rocky woods, thickets, and open decis Cireumneutral. General. Rudbeckia subtomentosa Pursh. open woods, thickets, and borders of prairie streams. Circumneutral. General. Rudbeckia subtomentosa var. Craigii Sherff, Rh. 14: 164. 1912. Oceasionally found with the typical form. St. Loui Rudbeckia speciosa Wenderoth. Rocky open woods, glades, and thiekets. Cireumneutral. Southern Mo.: Iron, Shannon, Jasper, and Newton counties. Rudbeckia palustris Eggert. Wet ground about springs, wet ledges and banks. Caleiphile to circumneutral. Southern Mo.: Iron, Wayne, Ripley, Dent, Shannon, Texas, Douglas, Ozark, and Rudbeckia hirta L. Black-eyed Susan, Nigger Head. Rudbeckia sericea Moore (S). Rudbeckia longipes Moore (8). Open Min thickets, rocky prairies, and waste ground. Circumneutral. Gen- eral and common Rudbeckia missouriensis Engelm. Rudbeckia fulgida of auth. in part, not Ait. (G), (B € B). Rocky prairies, glades, and bald knobs. Calciphile. Southern and central Mo. south and east of a line drawn from St. Charles, Montgomery, Boone, Morgan, I 2 and Polk eounties to Benton Co. This species has been confused with Rudbeckia fulgida which is not known to occur in Mo. en. a Gmel. Dr en ground. Eastern Ozark region: Washington and Shannon counties, and MR) introduced in Boone and Jackson counties. [Vor. 22 670 ANNALS OF THE MISSOURI BOTANICAL GARDEN Rudbeckia laciniata L. Low open woods, alluvial thickets, and banks of small streams. Cireumneutral. General. Rudbeckia maxima Nutt. )pen and waste ground. Introdueed from the southwest. Jackson Co. Rudbeckia amplexicaulis Vahl. Roadsides and waste ground. Cireumneutral to caleiphile. Introduced in western Mo.: Jackson, Jasper, and Newton counties. Echinacea purpurea (L.) Moench. Purple Cone-flower. rauneria purpurea (L.) Britton (G). Roeky open woods, thiekets, and glades. Caleiphile to eireumneutral. Gen- eral but commonest in southern and central Mo. Echinacea angustifolia DC. pland prairies. Cireumneutral. Northeastern Mo., loeal: Shelby Co. Echinacea pallida Nutt Brauneria pallida (Nutt.) Britton (G). Rocky prairies and glades. Caleiphile to cireumneutral. General. Echinacea paradoxa (Norton) Britton. rauneria paradoxa Norton (G). Rocky prairies, glades and bald knobs. Southern Mo., eentral and western Ozark region: Phelps, Pulaski, Miller, Camden, Laclede, Bs. Dallas, Cedar, Greene, Ozark, Taney, Stone, and Barry counties. Ratibida pinnata (Vent. ) Barnhart. Lepachys pinnata (Vent.) T. & G. Rocky prairies, thickets, and borders of woods. Circumneutral. General. Ratibida columnaris (Sims) D. Don. Lepachys columnaris (Sims) T. & G. rairies, waste ground, and along roadsides and railroads. Cireumneutral. Seattered: Marion, Lineoln, St. Louis, Clay, son, Greene, and Jasper counties, Generally and perhaps everywhere Aipa fro i durer west. xc m f. pulcherrima (DC.) W. M. Sharp, Ann. Mo. Bot. Gard. EL duda var. pulcherrima (DC.) T. & G. (G). Waste ground. Introdueed in Jackson Co. — americana (Mutis) Hieronymus var. repens (Walt.) A. H. Moore. wet woods, and borders of swamps and bayous. Cireumneutral. South- d Mo.: Bollinger, New Madrid, Pemiscot, Dunklin, Butler, and Ripley counties. Helianthus annuus L. Sunflower. aste and cultivated ground. Indifferent. General but scattered. Helianthus petiolaris Nutt. Prairie Sunflower, Kansas Sunflow Fields, roadsides, and waste ground. Cireumneutral to bci Seat- tered: Marion, St. Louis, Seott, Oregon, Jackson, and Jasper counties. Helianthus divaricatus L. Rocky woods and thickets. Cireumneutral. Eastern Mo.: Marion, Ralls, Pike, St. Louis, Bollinger, New Madrid, Reynolds, and Oregon counties. 1935] PALMER € STEYERMARK—— PLANTS OF MISSOURI 671 Helianthus lenticularis Dougl. Waste ground. Introduced in Jackson Co. Helianthus salicifolius A. Dietr. See A Mich. Acad. Sei. 9: 333. 1929. Helianthus orgyalis DC. (G), (B & B). Upland prairies and limestone Piu. A s Western Mo.: Jackson, Henry, Bates, Vernon, and Jasper countie Helianthus atrorubens L. var. hist B See Rh. 34: 1-2. 1932. Dry open ground. Cireumneutral. Southern Mo.: Scott, Gtiddsrd, Dunklin, Butler, Wayne, pem Bipley, Oregon, Lem and Ozark counties. Helianthus rigidus (Cass.) Desf. See Papers Mich. Acad. Sci. 9: 344. 1929. Helianthus scaberrimus Ell. (G B & B). foy open woods, thiekets, ee prairies. Circumneutral. General except in Southeastern lowlands and parts of the Ozark region. Helianthus laetiflorus Pers. Prairies, thickets, and open banks. Cireumneutral. Scattered: Ralls, St. Louis, Jefferson, and Jackson counties. Helianthus occidentalis Riddell. Rocky open e prairies, and glades. Circumneutral to calciphile. Central and southern Mo.: St. Louis, Franklin, Washington, St. Francois, Crawford, grim Phelps, Pulaski, Dent, Shannon, Oregon, Howell, Texas, Wright, s, Camden, Dallas, Bicis, Vernon, Jasper, and MeDonald counties. Helianthus mollis Lam rairies, fields, iste. and roadsides. Cireumneutral. General. Helianthus grosseserratus Marten Fields, Virus and PERA of small streams. Circumneutral. p central, u stern Mo. Known south of the Missouri River in St. Louis, Jef- ferson, Iron, Mu Hiekory, Cooper, Saline, Pettis, Lafayette, ne, Henry, Greene, Jackson, Cass, Bates, Barton, and Jasper counties, and in a the northern eounties Helianthus Maximiliani Schrad. cky Mir e) mies: loess hills, and bald knobs. Cireumneutral to caleiphile. Winters and s ern Mo., scattered: Atchison, Jackson, Ozark, Taney, Stone, and Barry ud Helianthus microcephalus T. & T Small Woods Sunflow Moist open woods and thickets, epa to MEERE 7 Southeastern Mo.: Iron, Seott, and Dunklin eountie Helianthus doronicoides Lam Open w a thickets, ind prairies. Circumneutral. Scattered: Schuyler, Boone, St. Louis, Shannon, and Jasper counties. Helianthus formosus E. E. Wats. Papers Mich. Acad. Sci. 9: 445, pl. 7 10) oods and along bluffs. Circumneutral to calciphile. DE M We cur TMo.: : Marion, St. Louis, Jefferson, Shannon, and Greene counties. Helianthus angustifolius L. Sa open woods and prairies. Oxylophile. Southeastern Mo.: Scott, Mis- sissippi, and Butler counties. [VoL. 22 672 ANNALS OF THE MISSOURI BOTANICAL GARDEN Helianthus hirsutus Raf, Helianthus leptocaulis (Wats.) Blake. Rocky open woods, thiekets, and prairies. Cireumneutral. General. Helianthus hirsutus var. trachyphyllus T. € G. Prairies and thickets, Occasionally found with the typical form. Jasper Co. Helianthus hirsutus var. leoninus = E. Wats.) Palmer & Steyermark, comb. nov. Helianthus leoninus E. E. W Helianthus virilis E. E, ring Open woods and roadsides. Cireumneutral. Scattered: Marion, Lafayette, Clay, Jackson, and Buchanan counties. Helianthus strumosus L. Prairie swales and moist ground along streams. Cireumneutral. General but scattered: Marion, Ralls, Pike, St. Louis, Crawford, Dunklin, Boone, Taney, Jackson, and Jasper counties. Helianthus tracheliifolius Mill. Introduced in waste ground. Jackson Co. Helianthus decapetalus L. Open woods, renta and prairies. General but seattered: Marion, Ralls,. Boone, Dunklin, Jasper counties. Helianthus MS L. Jerusalem Artichoke. Moist woods and thickets, wet prairies, and waste ground. Circumneutral. General. Helianthus tuberosus var. subcanescens Gray. Helianthus subcanescens pod = E. Wats. Helianthus mollissimus E. Commonly found with the win dl Seattered: Audrain, St. Louis, Jef- ferson, Jackson, and Jasper counties. Helianthus instabilis E. E. Wats. Papers Mich. Acad. Sci. 9: 423. pl. 65. 1929. Moist prairies and roadsides. Cireumneutral. Scattered. Verbesina encelioides (Cav.) Benth. & Hook. var. exauriculata Robinson € Greenman. Introdueed in waste ground. St. Louis and Jackson counties. Verbesina virginica L. White Crown-beard. eky open woods and thiekets. parc ER East-central and southern : Bt. Louis, Jefferson, Franklin, Washington, Dent, Iron, Scott, Stoddard, tien Shannon, Texas, Howell, Phelps, Pulaski, Douglas ney, Greene, Webster, nins. Barry, Vernon, Barton, Jasper, Newton, and McDonald counties. Verbesina helianthoides Michx. Rocky open woods and thickets. Cireumneutral. General. Actinomeris alternifolia (L.) DC. en woods and thiekets. Cireumneutral. General. Coreopsis tinctoria Nutt. Glades and sandy or rocky open ground. Oxylophile to eireumneutral. Seat- tered: Schuyler, Boone, Cole, Phelps, Butler, Ripley, Oregon, Jackson , Jasper, and Newton counties. Native in southern Mo., and escaped from eutiteation northward. 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI 673: Coreopsis lanceolata L. Tickseed Coreopsis. ocky prairies, glades, and fields. Oxylophile. Central and southern Mo. north to St. Louis, Saline, and Clay counties, and escaped from cultivation in Pike Co. Coreopsis lanceolata var. villosa Michx. reopsis crassifolia Ait. (B & B), (R). Rocky prairies and glades. Oxylophile. East-central and southern Mo.: St. Louis, Jefferson, Franklin, Shannon, Texas, Howell, Ozark, Barry, cdm and MeDonald counties. Coreopsis grandiflora Hogg. Prairies, meadows, glades, and thickets. Cireumneutral to oxylophile. West- ern and southern Mo.: Wayne, Ripley, Jaekson, Henry, St. Clair, Cedar, Bates, Vernon, Greene, Jasper, and Newton countie Coreopsis pubescens Ell. Star Tickseed. Roeky open woods, thiekets, and along gravelly beds of streams. kj ee to eireumneutral. Eastern, central, and southern Mo.: Marion, St. Lou Jef- ferson, BEN Washington, Crawford, Iron, Reynolds, Ripley, Ala Ore- gon, l, Texas, Pulaski, Miller, Dallas, Hickory, Polk, Wright, St. Clair, Cedar, Ca aney, Douglas, Shae Newton, and McDonald counties. Coreopsis palmata Nutt. Open woods, glades, thickets, and rocky prairies. Cireumneutral to oxylophile. General, but commonest in the Ozark region Coreopsis tripteris L. Tall Tickseed. Rocky open woods, bia. thickets, and prairies. Circumneutral to calciphile. General, but commonest in the Ozark ips Coreopsis tripteris var. Deamii Standley, R 1930. Occasionally found with the typical dr, phon St. Louis, Oregon, War- ren, Boone, Dekalb, Harrison, Mereer, and Saline counties. Bidens discoidea (T. € G.) Britton. Wet ground along streams, and Ant eges Circumneutral. Eastern Mo.: Lineoln, St. Louis, Jefferson, Cape G Butler counties. Bidens roncon L. Beggar-ticks. st woods, thickets, and waste ground. Cireumneutral. General. Bidens td Greene. Moist ground along streams and about ponds, fields, thiekets, and waste ground. Circumneutral. General but scattered: Marion, Pike, St. Louis, Shan- non, Atehison, and Jackson counties. Bidens vulgata var. puberula (Wiegand) Greene. d found with the typieal form. Western Mo.: Atchison and Jack- son eounties Bidens comosa (Gray) Wiegand. et ground, borders of aa sloughs, and ponds. Cireummeutral. Gen eral but scattered: Putnam, Boone, Pike, St. Louis, Saline, Atchison, Clay, Jackson, and Greene counties. Bidens comosa var. acuta Wiegand. Occasionally found with the typical form. St. Louis, Perry, Carroll, Gentry, and Jackson counties. [Vor. 22 674 ANNALS OF THE MISSOURI BOTANICAL GARDEN Bidens connata Muhl. Swamp Beggar-ticks. et borders of streams, ponds, and sloughs. Circumneutral. General but scattered: Marion, Pike, Boone, St. Louis, Saline, Atchison, Jackson, and Green counties Bidens cernua L. See Rh. 24: 206. 1922, Sticktight. Muddy borders of slow streams, sloughs, and ponds. Cireumeutral. General. Bidens cernua var. elliptica Wiegand. Occasionally found with the typical form. Western Mo.: Jackson and Jasper counties Bidens cernua var, dabii Wiegand. Frequently found throughout the range of the typical form. Bidens «line (L.) BSP. ground about sloughs and ponds. Cireumneutral. Eastern Mo., south of the Missouri River: St. Louis, Jefferson, and Dunklin counties, Bidens bipinnata L. Spanish Needles Open woods, thickets, fields, and isi ground. Cireumneutral. General. Bidens aristosa (Michx.) Britton. Prairies, cultivated fields, and waste ground. Cireumneutral. General, but most common in prairie regions and in alluvial Él Bidens aristosa var. Fritcheyi Fernald, Rh. 78. 1913 Oeeasionally found with the typical id deste: Marion, St. Louis, and Jasper counties. Bidens aristosa var. mutica (Gray) Gattinger. See Rh. 15: 78. 1913. Sometimes found with the typical form. Scattered: Lewis, Marion, Ralls, Pike, St. Louis, Phelps, Jackson, and Greene counties. Bidens involucrata (Nutt.) Britton. et prairies, cultivated or fallow fields, and waste ground. Cireumneutral. General. Bidens Beckii Torr. Water Marigold. Wet ground about ponds and sloughs. Cireumneutral. Eastern Mo., loeal: St. Louis Co. Thelesperma trifidum (Poir.) Britton. Rocky prairies and glades. Caleiphile. Southwestern Mo., local: Greene Co. Thelesperma gracile (Torr.) Gray. Waste ground and along railroads. Introduced in Jackson Co. Marshallia caespitosa Nutt Rocky or upland prairies and glades. Calciphile to cireumneutral. South- western Mo.: Jasper and Newton counties. — obovata ( Walt.) Beadle € Boynton var. platyphylla (Curtis) Beadle &B n. False Sea = and bald knobs. ' Caleiphile. Southern Mo., local: Ozark Co. — PARVIFLORA Cav. w and cultivated ground. Indifferent. Scattered. GALINSOGA CILIATA (Raf.) Blake, Rh. 24: 35. 1922. Galinsoga parviflora var. hispida DC. (G). Waste and cultivated ground. Indifferent. General. 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 675 Flaveria O Johnston and waste ground. raue from farther west. Marion and Jackson xu Hymenopappus cipem L’Her. Hymenopappus carolinensis (Lam.) Porter irn (B & B), (R). Rocky prairies, Ps ald knobs, and sandy open woods. Circumneutral to oxylophile. Southern Mo.: Seott, E Taney, Stone, and Barry counties. This has "iis eonfused with Hymenopappus corymbosus whieh is not known in Mo. Palafoxia ae (Nutt.) T. : ris callosa (Nutt.) Gray (G). DAE callosum (Nutt.) Bush (B & B). Glades, bald knobs, rocky open ground, and along gravelly beds of streams. Basi rd iba Mo.: Douglas, Ozark, Greene, Taney, Stone, Barry, and MeDonald counties. Helenium nudiflorum Nutt. Sneezeweed. Helenium campestre Small (8). Helenium polyphyllum Small (R). Moist ground, meadows and fields, and borders of ponds, sloughs, and streams. Oda General, but commonest in central and southern Mo. Helenium autumnale Helenium cR Mill. (R), (S). Helenium altissimum Link ( Moist dien MEE swales, — borders of ponds and streams. Cireum- neutral. Gene mmon. Helenium tenuifolium Nutt. Bitterweed, Yellow Dog Fennel. Fields and waste ground. Oxylophile. General, but commonest in the Ozark ion. Generally and perhaps everywhere introdueed from farther south, and sometimes becoming a troublesome weed Gaillardia lutea Greene. Moist sandy open ground. Oxylophile to eireumneutral. Southeastern Mo., loeal: Dunklin Co. dope pulchella Foug. Indian Blanket. elds and waste ground. Cireummeutral Introduced or escaped from culti- cs Pike, Platte, and Jackson counties. TAGETES ERECTA L. sides and waste ground. Escaped from cultivation. Scattered: Marion, St. all and St. Francois counties. Dyssodia papposa (Vent.) Hitche. Fetid Marigold. Dry prairies, fields, and waste ground. Cireumneutral to oxylophile. General. Achillea Millefolium L. Yar Prairies, thiekets, and us ine" Indifferent. General and common. Achillea Millefolium f. rosea Rand & Redfield. Commonly oceurring with the typieal form. General. Achillea lanulosa Nutt. Fields and waste ground. Cireumneutral. Scattered: Sullivan, Marion, Monroe, St. Louis, Linn, Moniteau, Wright, and Vernon counties. [VoL. 22 676 ANNALS OF THE MISSOURI BOTANICAL GARDEN ANTHEMIS CoTULA L. Dog Fennel. Fields and waste ground; frequently about dwellings. Indifferent. General. ANTHEMIS ARVENSIS L. var. AGRESTIS (Wallr.) DC. Corn Chamomile. Fields and waste ground. Indifferent. Seattered: Jackson and Jasper counties. ANTHEMIS TINCTORIA L. Yellow Chamomile. seaped from cultivation. Local: Greene Co. MATRICARIA CHAMOMILLA L. Wild Chamomile. Introduced in waste ground. Indifferent. Seattered: St. Louis and Phelps counties, MATRICARIA MATRICARIOIDES (Less.) Porter. neo Weed. Matricaria suaveolens (Pursh) Buchenau (G), (B & B). Rocky waste ground, fields, and roadsides. Oxylophile to eireumneutral. Cen- tral and southern Mo., scattered: St. Louis, Jefferson, Franklin, Jasper, and Newton counties. — LEUCANTHEMUM L. Ox-eye Dais 8, esed ion ads and waste ground. — St. Louis, Boone, bite and Jackson coun CHRYSANTHEMUM "Hoppe var. PINNATIFIDUM Lecoq & Lamotte. Fields, rocky pastures, and waste ground. Indifferent. General and more common than the typieal form. TANACETUM VULGARE L. Tansy. Fields and waste ground. Indifferent. General but scattered. TANACETUM VULGARE var. CRISPUM DC, Rarely found with the typieal form. Boone Co. Artemisia dracunculus L. var. glauca Jepson. rtemisia dracunculoides Pursh (G), (B & B). Prairies, rocky or alluvial banks, and loess hills. Caleiphile to cireumneutral. Northern Mo., seattered: Clark, Holt, Buchanan, and Jackson counties. Artemisia ei L. var. caudata (Michx.) Palmer € Steyermark, comb. nov. Wild Wormwood. did caudata Michx. (G), rtemisia campestris subsp. Spada (Miehx.) Hall & Clements. des, bluffs, and gravel bars of streams. Oxylophile. Eastern Mo. . south of m Missouri River: St. Louis, M Dent, and Shannon counties Artemisia frigida Willd. Wormwood Sa Waste ground. Introduced from her west. Jackson Co. ARTEMISIA ANNUA L. Wormwood. Fields, and rocky or waste ground. ee Eastern, central, and south- ern Mo., commonest in the Ozark regio Artemisia biennis Willd. aste ground and gravel bars of streams. Circumneutral. Scattered: St. Louis, Iron, Clay, and Jackson counties. Artemisia vulgaris L. var. ee (Nutt.) Ktze. Mugwort. Artemisia ludovieiana Nutt. (G) in part, (B&B). Rocky prairies, a and glades. Caleiphile to eireumneutral. General but seattered. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 677 Artemisia vulgaris var. . Wrightii (Gray) Palmer & Steyermark, comb. nov. Artemisia vulgaris subsp. Wrighti Hall & Clements. Artemisia kansana Britton (G), (B & B). Artemisia Carruthi Wood. Waste ground. Iutrodueed from the Southwest. Jaekson Co. Artemisia vulgaris var. gnaphalodes (Nutt.) Ktze. Artemisia inito Nutt. )" Artemisia ludov a of auth. in part, not Nutt. (G) in par Upland prairies p glades. Cireumneutral to calciphile. Northern, central, and western m. Clark, Knox, Adair, Marion, Ralls, St. Charles, St. Lou Mereer, Worth, Grundy, Carroll, Saline, Harrison, T i Dekalb. Atchison, bis Clinton, Clay, Jackson, and Dade c Artemisia vuigaris var. mexicana (Willd.) Torr. & n. Scheele. Rocky prairies and glades. en to eireumneutral. Western and middle Mo.: Phelps, Macon, Chariton, Boone, Saline, Henry, Jackson, Barton, Jasper, and Newton counties. Erechtites hieracifolia (L.) Raf. var. intermedia Fernald, Rh. 19: 27. 1917. Fireweed. Erechtites hieracifolia of auth. in part (G), (B & B). Rocky open woods, thickets, and waste ground; also often in burned-over woods. Indifferent. General. The typical form is of more northern range and is not known in Missouri. Cacalia atriplicifolia L. Open woods and thickets. Circumneutral. General, Cacalia suaveolens L. Rich or wet open woods. Cireumneutral. East-central Mo.: Warren, St. Louis, and Jefferson counties Cacalia reniformis Muhl. ich or moist rocky woods. Cireumneutral. Eastern, middle, and southern Mo.: Putnam, Sullivan, 'Bshujler , Adair, Clark, Ralls, Boone, Montgomery, Warren, St. Louis, Jefferson, Franklin, Ste. Genevieve, Cape Gi irardeau, Bol- linger, Shannon, Texas, Howell, Douglas, Barry, and MeDonald counties. Cacalia tuberosa Nutt. Indian Plantain. Prairies and limestone glades. Caleiphile to eireumneutral. General but commonest in the glades of the Ozark region. Senecio glabellus Poir. Butter-weed. uvial una along and borders of sloughs and swamps. Cir- cumneutral. Northern, Eds and central Mo., mostly along the larger rivers: Shelby, a Lincoln, St. , Jeffer Ste. Genevieve, Bollinger, Mis- sissippi, Pemiseot, Dui reine ds EM ue Ripley, RN Linn, Livingston, Boone, hse Saline, Clay, and Jackson counti tis rh Muhl. Squaw oods and moist ras kid bluffs. Cireumneutral to ealeiphile. Cen- E ih pier a Mo., seattered: Jefferson, Boone, Phelps, Shannon, Greene, Taney, Vernon, Jasper, and McDonald counties. (Vor. 22 678 ANNALS OF THE MISSOURI BOTANICAL GARDEN Senecio obovatus var. rotundus Britton. In similar situations to the preceding, but more common. Circumneutral. Central and southern Mo. south and east of a line drawn from St. Louis, Mont- gomery, Boone, and Morgan counties to Jasper Co. Senecio obovatus var. umbratilis Greenman. Occasionally found with the typical form. Circumneutral. Scattered. Shan- non and Moniteau counties. Senecio aureus L. (Golden Ragwort, Squaw-weed. Rich or rocky woods, and along spring brooks and streams. Cireumneutral. General, Senecio aureus var. semicordatus (Mack. € Bush) Greenman, Ann, Mo. Bot. Gard. 3: 129, 1916 Senecio aureus x Balsamitae Greenman. Prairies swales. Cireumneutral. Western Mo,, local: Jackson Co. Senecio aureus var. gracilis (Pursh) Brit Moist woods and prairie swales. ME Scattered: Randolph and Douglas counties. Senecio plattensis Nutt. Prairie Ragwort. Senecio pseudotomentosus Mack. & Bush (B & B). Upland prairies, glades, rocky woods, and ledges. Circumneutral to ealeiphile. General. Senecio pauperculus Michx. See Ann. Mo, Bot. Gard. 3: 159. 1916. Senecio Balsamitae Muhl. (G B & B). Senecio pauperculus var. BoleeiMiae (Muhl.) Fernald. Prairies and meadows. Cireumneutral. Northeastern and central Mo.: Adair, Maeon, Knox, Shelby, Marion, Ralls, Pike, Audrain, Randolph, Boone, Callaway, Moniteau, Pettis, and Henry counties. ARCTIUM MINUS Bernh rdock. Waste ground, frequently about dwellings. Indifferent. General and common. ARCTIUM TOMENTOSUM Mill. See Rh. 12: 44-45. 1 Waste ground. Introduced in Jackson Co, CARDUUS CRISPUS L. Curly Thistle. Waste ground and roadsides. Introduced in St. Louis Co. CIRSIUM LANCEOLATUM (L.) Hill. Bull Thistle. Fields, pastures, and waste ground. Indifferent. General. Cirsium undulatum (Nutt.) Spreng. var. megacephalum (Gray) Fernald. Loess hills and prairies. Scattered: Wayne, Atchison, and Jackson counties. Cirsium discolor (Muhl.) Spreng. Prairies, thickets, and open woods. Cireumneutral. General but scattered: Mercer, Marion, Boone, St. Louis, Dunklin, Vernon, Barton, and Jasper counties Cirsium odoratum — Petrak, Beih. z. Bot. Centralbl. II. 35: 378-381. 1917. Cirsium pumilum Spreng. (G). Cirsium Hillit ( Canby) Fernald (G), (B & B). Waste ground. Introduced in St. Louis Co. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 679 Cirsium ochrocentrum Gray var. Helleri (Small) Petrak, Beih. z. Bot. Centralbl. II. 35: 418-419. 1917. Waste ground. Introduced in Jackson Co. Cirsium altissimum (L.) Spreng. See Beih. z. Bot. Centralbl. IT. 35: 396-400. 1917 Cirsium iowense (Pammel) Fernald (R). Thickets, dry open or low alluvial woods, and rocky slopes. Cireumneutral to oxylophile. General. The characters used to distinguish Cirsium iowense do not seem constant and it intergrades freely with true C. altissimum. Cirsium virginianum (L.) Michx Rocky open woods and thiekets. Oxylophile to cireumneutral. Southern Mo., eastern Ozark region: St. Francois, Iron, Madison, Wayne, Carter, Ripley, Dent, and Phelps counties. Cirsium muticum Miehx. Swamp Thistle Swampy ground. Circumneutral. Takt dii Mo., south of the Missouri River: St. Louis and Shannon counties, CIRSIUM ARVENSE (L.) Seop. Canada This Fields and waste ground. Indiffer den u Adair, St. Louis, Jeffer- son, St. Franeois, and Jackson counties. nn ACANTHIUM L. Cotton Thistle, Seoteh Thistle. ground. ed rent. Southeastern Mo.: Iron, St. Francois, and Cape phis coun CENTAUREA CYANUS L. Ragged Robbin, Bachelor's Button. Waste ground and roadsides. Escaped from cultivation. Indifferent. Scat- tered: Boone, Jackson, and Jasper counties. CENTAUREA SOLSTITIALIS L. Barnaby’s Thistle. and waste ground. Introduced in Boone Co. CENTAUREA MELITENSIS L. aste ground. Introduced in Jackson Co. ee VOCHINENSIS Bernh. s and waste ground. Introduced in Boone and St. Louis counties. CENTAUREA MACULOSA Lam Old fields and ub dunt Seattered. Introdueed in Boone and Howell countie CENTAUREA PICRIS Pall. aste ground. Introduced in Jackson Co. Centaurea americana Nutt. dai Roeky open woods, glades, and bald knobs. Southwestern Mo.: Barry and McDonald counties, and also introduced in St. Louis and Jackson counties. Serinia oppositifolia p )K Prairies, meadow AL and glades or sandy open ground. Oxylophile. Southern and end Mo. and loeally northwestward in Livingston Co.: St. Francois, Bollinger, Cape Girardeau, Wayne, Seott, Dent, Oregon, Maries, Cass, Henry, Livingston, Bates, Lawrence, Jasper, Newton, and McDonald counties. [VoL. 22 680 ANNALS OF THE MISSOURI BOTANICAL GARDEN CICHORIUM INTYBUS L. Chicory. Fields, pastures, and waste ground. Indifferent. General but commonest in eastern Mo. Krigia virginica (L.) Willd. Dwarf Dandelio Roeky open ground, glades, and fallow folds. — Eastern and south- ern Mo.: Montgomery, Lincoln, Jefferson, St. Fra s, Ste. Genevieve, Madi- son, Wayne, Iron, Gasconade, Texas, Polk, Henry, orat Ozark, Lawrence, Jasper, Newton, and MeDonald counties, Krigia biflora (Walt.) Blake, Rh, 17: 135. 1915. Krigia amplexicaulis Nutt. (G). Cynthia virginica (L.) D. Don (B & B), in part. Rocky open woods, low ground along streams, thickets, and prairies. Circum neutral to oxylophile. General and common Krigia Dandelion (L.) Nutt. Prairies, glades, borders of woods, sandy fields, and alluvial or low sandy woods. Oxylophile. Central and southern Mo.: Montgomery, St. Louis, Jef- ferson, Franklin, Washington, St. Francois, Ste. Genevieve, Madison, ee Crawford, Iron, Wayne, Ripley, Dunklin, u Reynolds, pose Carter, Gas conade, Sh annon, Phelps, Maries, Pulaski, Texas, Laclede, Douglas, Howell, Henry, Polk, Cedar, Webster, Vernon, Greene, Jasper, isl and MeDon ald counties. Krigia occidentalis Nutt. mbia occidentalis (Nutt.) Standley (B & B). Rocky open woods and glades. Oxylophile. Southwestern Mo.: Jasper and MeDonald counties. This species has often been confused with K rigia virginica and Serinia oppositifolia. Pickis SPRENGERIANA Poir. See Bull. Mo. Bot. Gard. 19: 121. 1931. aste ground. Introduced in St. Louis Co. TRAGOPOGON PRATENSIS L. Goat’s Beard. Fields, meadows, and bad een and railroads. Indifferent. General: Lineoln, St. Charles, St. Louis, Jeffers St. Francois, Madison, Maries, Schuyler, Adair, Mercer, doc Laclede, e diam Nodaway, and Platte coun- ties, and spreading state-wide. er PORRIFOLIUS L. Vegetable Oyster, Salsify. and waste ground. Escaped from eultivation, Indifferent, Seattered: Boone, st. Louis, Planta amden, and Jackson counties, TARAXACUM PALUSTRE as Lam. & DC. var. VULGARE (Lam.) Fernald, Rh. je 380. 1933. Dandelion. a officinale bie (G). ee Taraxacum L. Fields and waste ground; common | abut dwellings. Indifferent. General. TARAXACUM LAEVIGATUM (Willd.) DC. See Rh. 35: 379. 1933. Red-seeded Dandelion Fue erythrospermum Andrz. (G). Leontodon erythrospermum (Andrz.) Britton (B & B). Fields and waste ground about dwellings. Indifferent. General. 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 681 Crepis setosa Hall. Hawk's Beard. Fields and waste ground. Introduced in Boone Co. CREPIS CAPILLARIS (L.) Wallr. pen ground. Introduced in St. Louis Co. SONCHUS ARVENSIS L. Sow Thistle. Waste ground. Indifferent. Introduced in Jackson Co. SoNcHUS OLERACEUS L. Common Sow Thistle. Fields and waste ground. Indifferent. General. SONCHUS ASPER (L.) Hill. Fields and waste ground. Indifferent. General. LACTUCA SALIGNA L. Roadsides and waste ground. Indifferent. Seattered: Marion, Pike, Monroe, St. Louis, Maries, Phelps, Pulaski, Laclede, Saline, Pettis, Dallas, Wright, Web- ster, Douglas, Greene, and Barry counties. LACTUCA SCARIOLA L. Priekly Wild Lettuce Roadsides zi waste ground. Indi mee) General. LACTUCA SCARIOLA var. INTEGRATA Gren, & Godr. Commonly found with the typieal form. General. Lactuca canadensis L. var. typica Wiegand, Rh. 22: 10. 1920. L. canadensis L., in part (G), (BG B e thickets, and waste ground. Indifferent: General but scattered. Lactuca canadensis var. latifolia Ktze. Occasionally found with the typical form. Scattered: St. Charles, St. Louis, Jefferson, Boone, Clinton, and Jasper counties. Lactuca NOS var, latifolia f. exauriculata Wiegand, Rh. 22: 11. 1920. Sometimes found with the typical and other varieties. Scattered: St. Fran- eois, Howell, and Hickory counties. Lactuca canadensis var. integrifolia T )T.&G. Lactuca sagittifolia Ell. (G), (B & B). actuca canadensis var. montana ee (B&B). Thickets and waste ground. Indifferent. General but scattered: Boone, St. Louis, Dunklin, Jackson, Henry, and Jasper counties. Lactuca canadensis var. integrifolia f. Red picis Rh. 22: 11. 1920. Oeeasionally found with the other forms. St. Louis Co. Lactuca canadensis var. obovata Wiegand, E Rh. 22: 11. 1920. Lactuca integrifolia of auth., not Bigel. (G). Thickets, fields and waste ground. Beatieréd : St. Louis, Ste. Genevieve, Mis- sissippi, Boone, Jackson, and Taney counties. Lactuca canadensis obovata = ne oda y nein Rh. 22: 11. 1920. Occasionally occurring with t ariety. Lin Lactuca ludoviciana (Nutt.) Riddell. Plains and waste ground. Cireumneutral Northwestern Mo.: Atchison Co. Lactuca hirsuta Muhl. open woods and waste ground. Cireumneutral. Southwestern Mo., loeal: Jasper Co. [Vor. 2 ba ly 682 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lactuca villosa Jacq. pen woods and thickets. Cireumneutral. General. Lactuca ze (Pursh) DC. Moist open ground and loess hills. Caleiphile, Western Mo.: Atchison and Bein eounties Lactuca floridana (L.) Gaertn. pen woods and thickets. Cireumneutral. General. Lactuca graminifolia Michx. Introduced along railroads. Butler Co. Lygodesmia juncea (Pursh) D Loess hills. Caleiphile, poo ME Mo.: Atehison, Holt, and Buehanan counties. Agoseris cuspidata (Pursh) Steud. Prairies and glades. Calciphile. West-central Mo., local: Jackson Co. Pyrrhopappus carolinianus (Walt.) DC. False Dandelion. Prairies, fields, and waste ground. Cireumneutral. General. Prenanthes altissima L. var, cinnamomea Fernald. ocky woods and bluffs. Cireumneutral. Eastern and middle Missouri, mostly in the eastern Ozark region: Boone, Warren, St. Louis, Jefferson, Ste. tenevieve, Wayne, Iron, Dunklin, Carter, Shannon, Texas, and Ozark counties. Prenanthes alba L. Rattlesnake-root, White Snake-root. Rich or rocky woods. Crowd to es Eastern Mo.: Clark, Marion, St. Louis, Franklin, and Gaseonade counties Prenanthes aspera Michx. Rocky open woods and prairies. Cireumneutral. General but seattered. Prenanthes crepidinea Michx. pen woods and thickets. Cireumneutral. East-eentral and southern Mo.: Audrain, Boone, St. Louis, Washington, Wayne, Greene, and Taney counties, Hieracium oa Michx. Hawkweed. Rocky woods and thickets. Oxylophile. Eastern, middle, and southern Mo.: Adair, Boone, Callaway, Lincoln, St. Loui uis, Jefferson, St. Francois, Ste. Gen vieve, Scott, Iron, Reynolds, Shannon, Dent, Wright, Greene, Stone, Jasper, oy Newton countie Hieracium ane L. See Rh. 37: 185. 1935 Roeky open woods, thiekets, and fields. Oxylophi General. Several forms of this species have been confused with both Hieracium venosum and Hieracium Traillii (Hieracium Greenii) neither of ng oeeur in Missouri. Hieracium Gronovii var. foliosum Michx. See Rh. 37: 185. 1935. Similar eg to pi Metern Oxylophile. Seattered in eastern, cen- tral, and s ern Mo.: St. Louis, Jefferson, Shannon, Phelps, Ozark, Jasper, and Marion peri Hieracium longipilum Torr Me ded bo woods and prairies. Oxylophile. Eastern, central, and southern , Boone, Callaway, Iron, Reynolds, Dent, Shannon, Texas, Morgan, iig LS Polk, Greene, Howell, Vernon, Barton, Jasper, and Newton countie 1935] PALMER € STEYERMARK— —PLANTS OF MISSOURI TABULAR List or FAMILIES Genera Species Varieties Forms z "d à m : a Families > 3 in E = 3 E 3 5 = S| & 5 * S43 BE zZzis8|iZ|8iIizZ |3|lz 19] SPERMATOPHYTA GYMNOSPERMAE Pinaceae 3 4 2 1 ANGIOSPERMAE MONOCOTYLEDONEAE Typhaceae 1 2 Sparganiaceae 1 3 Najadaceae 4 11 1 4 Alismae 5 13 3 1 Hydrocharitaceae 2 9 Gramineae 59 130 be 51 5 7 Cyperaceae 12 166 22 Araceae 3 4 1 Lem 4 9 1 Commelinaceae 2 13 1 Xyridae 1 1 Pontederiaceae 3 5 Juneace 2 18 6 1 Liliacea 18 5 39 9 5 3 Dioscoreaceae al 2 1 Amaryllidaceae 3 1 3 1 al I Tridaceae 3 ji 9 4 1 Marantaceae 1 1 Orchidaceae 14 27 3 DICOTYLEDONEAE Piperaceae 1 1 Salicaceae 2 14 5 4 3 1 Leitneriaceae il 1 Juglandaceae 2 10 8 3 Betulac 5 6 2 1 Fagaceae 3 23 8 3 Ulmaceae 3 8 5 orae 3 2 3 4 Urticaceae 5 7 1 Santalaceae 1 ak Loranthae j T Aristolochiaceae 2 2 3 Polygonaceae 5 ji siert 6 1 1 Chenopodiaceae 7 2 19 | 13 8 3 4 Amaranthaceae 5 9 4 3 1 Phytolaceaceae d 1 Nyctaginaceae 1 5 L Illecebraceae 1 2 Aizoaceae 1 il al J 684 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABULAR List or FAMILIES (Continued) [Vor. 22 Families Species Varieties | Forms Nati Introd. Native Introd. Native Introd. Hybrids Caryophyllaceae Portulacaceae Ceratophyllaceae Nymphaeaceae Ranunculaceae Magnoliaceae Annonaceae Capparidaceae Crassulaceae Zygophyllaceae Rutaceae Simarubaceae Hippocastanaceae Balsaminaceae Violaceae ore — ul A SS A m mo m m SETS THE OU A oe oe l bo i — t0 l2 pe — = DAA HH U OO SID DV pS O A r2 "e [wn oe t2 Da > e ji — NX»4ODO0OOaNNnN AS a No bi — T3 02 hd e m C mI pu — Ha pd oR bo ft w 1935] PALMER & STEYERMARK— PLANTS OF MISSOURI 685 TTABULAR List or Famurs (Continued) Genera Species Varieties Forms | : o 4 o - Q 4 Q an: Families E E Z E E E 5 E s E: + e + E > q 2 Pa 21/61/44 ar ra IC |E Passifloraceae 1 2 Loasaceae 1 1 Cactaceae 2 2 1 Thymelaeaceae 1 1 Elacagnacea 1 1 Lythrac 5 5 1 1 Melastomaceae 1 2 I Onagrae fj 27 3 Haloragidaceae 2 3 1 2 Araliaceae 2 3 Umbelliferae 23 9 32 10 5 Cornaceae 2 8 1 Ericace 3 6 6 Primulaceae 7 1 10 2 2 1 Sapotaceae 1 2 Ebenaceae 1 1 2 1 Styracaceae 1 1 Oleaceae 3 7 3 Loganiaceae 2 2 Gentianaceae 4 1 9 1 1 Apocynaceae 3 1 7 1 7 Asclepiadaceae 5 21 3 1 Convolvulaceae 4 13 6 5 1 Polemoniaceae 4 7 7 1 Hydrophyllaceae 5 10 Boraginaceae 8 2 13 7 4 Verbenaceae 2 8 1 4 Labiatae 18 6 49 | 18 14 1 2 Solanaceae 2 5 19 10 3 Serophulariaceae 21 3 44 8 11 2 1 Lentibulariaceae 1 1 1 Orobanchaceae 2 2 Bignoniaceae 3 4 Pedaliaceae 1 1 Martyniaceae 1 1 Acanthaceae B 6 i Phrymaceae 1 1 Plantaginaceae 1 10 al Rubiaceae 7 23 4 Caprifoliaceae 5 15 1 6 1 Valerianaceae 1 3 1 ipsacaceae 1 1 Cueurbitaceae 5 5 Campanulaceae 2 5 1 Lobeliaceae 1 6 1 Compositae 65 | 15 | 243 | 35 (Al 5 7 15 [ Vor. 22 686 ANNALS OF THE MISSOURI BOTANICAL GARDEN SUMMARY By DIVISIONS AND CLASSES Genera Species Varieties Forms " E z|28l|l$34|2|3|* A E E E A E E s| 5 * 5 3 E * $ * sel” Z a | 2 |9|2 || 2 |8|5 Pteridophyta 26 52 13 Spermatophyta 850 [109 |1785 |336 | 442 24 69 4 | 60 iymnospermae 3 4 2 1 Angiospermae 847 |109 |1781 |334 | 441 | 24 69 4 Monoeotyledoneae 139 | 20 | 509 | 68 89 5 14 1 Dieotyledoneae 708 89 |1272 |266 | 352 19 55 3 | 60 Summary By Minor Groups Lx BE PPM 131 Genera VE ERA re O e 876 a a A A AAA 109 TUM A A NO 985 Species A nee eT EN 1837 A AA O vn a el rines. 336 BEE. mise ee er et rise 2173 Varieties ET RE Y SSK RR SNR a RO FOR aie oho ewe ai k 455 er De a arriero 24 TUM A A A O 479 Forms a EA A O 74 ai AA A A A 4 WOME A E PE N TS EPVa LU. 78 BEINE: A A A 60 Total number of plants (species, varieties, forms, and hybrids) 2790 Of the 479 varieties listed 108 (such as Utricularia vulgaris var. americana, Lonicera dioica var. glaucescens) should be more properly counted in with the regular species, since they are the only representatives of particular species in the flora. This would bring the total number of species up to 2,281. 1935] PALMER € STEYERMARK—— PLANTS OF MISSOURI INDEX 687 Valid names of plants in ordinary Roman type, synonyms or otherwise invalid names in italic new species varieties, forms, hybrids, and new combinatio tals. Sy bold face, and family names in Roman capit Page Abutilon Theophrasti ........... 598 Acacia angustissima ............ 573 angustissima var. hirta ........ 57 IPBraipe A A 573 Acalypha: digyneig es... $e 588 Oractiens Of anth: aa oo 588 gracilens var. monococea ....... 588 ee es RE IAN 589 VIrgINICH 05 1 sie Sis eter te tue 588 ara ade 646 Acer fioridanüum re a. 593 Negundo cxkiu lon A 593 Negundo var. interior ......... 593 Negundo var. texanum ........ 593 Negundo var. violaceum ....... 593 OS OE 593 nigrum var. Palmen 22.0.0... 593 rubrum... DE Td er ee 593 rubrum var. Drummondii ...... 593 rubrum var. Drummondii f. ro- tundasla. sea eS 593 rubrum var. tridens ........... 593 UDCA: re 592 SACCHATINUM cic lo 593 Baccharum 5200509 050 sais oe 592 saecharum var. glaueum ....... 592 saecharum f. Rugelii .......... 592 accharum var. Rugelü ........ 592 saccharum var. Schneckii ...... 593 AOERAOBAHB Ma Meme 592 Acerates angustifolia ............ 622 floridana of auth. ............. 6223 irtela ar nee, 622 viridiflora. ee ne 622 viridiflora var. lanceolata ...... 622 viridiflora var. linearis ........ 622 Achillea lanulosa ............... 675 letolium: A een 675 Millefohum f. rosea ........... 675 Acnida D Hu tO RIO ERR 533 tubereulatag a ao 532 ns in Page tuberculata var. prostrata ...... 532 tubereulata var. subnuda ...... 533 Acorus Calamus a loe 495 EAT AA A O Ses 543 rae A RA wets 543 Aetinomeris alternifolia ......... 672 Adam-and-Eve ................. 510 Adder:3-month 2:22. 0.5: 510 Adder ’s-tongue Kite: 2:5... c benc MU 502 A A O 502 egilops cylindrica ............. 463 Aesculus arguta 3. sige dees ss 594 BUSTHUNG: ad crete ate mars Oe sce 594 a rol) N. en. ee ee 594 diseolor var. mollis ............ 594 A oe eurn usse 594 glabra var. arguta ......o..... 594 glabra var. Buckley .......... 594 glabra var. leucodermis ........ 594 labra var. pallida ...... sse 594 img var. Sargentil ........:- 594 OA d ads ele E ee 594 Afzellia iia PI E PEST 642 Agalinis TO IO qe 643 A A O 643 A lr N 644 heteropbylia i eee ere 644 DUPGS NE nne I Ae ad xs 643 A AN 644 tenulfola as a 643 tenuifolia var macrophylla ..... 643 tenuifolia var. parviflora ...... 644 A A A 644 Agastache N M ope Mts 631 serophulariaefolia ............ 31 serophular Fe var. mollis 632 Agave virginica .: B. ..-.ee 06 VIPQHUOR T. tigrina de 2.2.0 oe seo 507 virginica var. tigrina .......... 507 Agoseris euspidata .............. 682 [Vor. 22 688 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Page — ze. VEA SADIO. ¿a ana 50 iU 2 4 TT ERE ER 568 BEEN. coax aa ae 501 "M abba 568 "joo are 501 aa 569 on io RTT n A ek E tc 569 A oves dvo 501 TN Qik Daa oats 568 v a OA A PA A AR 517 Agropyron caninum var. tenerum .. 463 Aloe, False .........o.ooooooooo.. 506 pseudorepens ......ooooooo.o... 463 — aequalis ............. 466 repens var. a AAA 462 MOM anne 466 A A does ee 463 inercia PM 466 rt nina 463 goniculatus .................. 466 trachycaulum var. typicum ..... 463 geniculatus of auth. ........... 466 Agrostemma Githago ............ 536 geniculatus var. aristulatus .... 466 re DA ne 466 geniculatus var. ramosus ....... 466 RAS Lu anne 466 RR 466 e EVA A R49 EUR 466 lsine graminea .....oooooooo.o»» 535 Ayomalis 66006. rr 466 uU DELLI 2:33: iria 535 JENEPUDIA. ae 465 Di a rad 535 errada 466 u LE DELE T ET |. perennans var, aestivalis ....... 400. Alhan TOMÁA i: pav rar ed hx» 598 PEDIR ERA 466 Alles, Bhrabby PEE A d 9 vlla ad 599 Bohwenitsü ...... oe eee oo 460 ADUD Boot arena 553 DUUM a cr rn 465 Alyssum, Hoary ................ 547 A aaa 466 Amaranth li iua d Dex eb dx br s Ua 466 era 532 Ars 619 Aei A Ailanthus A sense 587 c4 Mer nr as pt 532 glandulosa nennen 587 ns PEN 532 Aira palloni ci 465 Amaranthus blitoides ............ 532 ABOGAUSAB sn 534 Po! Loose n 532 Alder graecizans var. pubescens ...... 532 MESE naar 592 ISDHOUN voy con tib o ade 532 A RARA 517 hybridus var. hypochondriacus .. 532 A A ee 576 IIA. range 532 Alisma Plantago-aquatica of Am Dnlenlablus ¿¿ 2.1 2950) ER 532 RAO dar ac Rc det a N 457 retroflexus ii oO sie A i57 CORO? diia 532 ALISMACBANE pu as 456 AMARYLLIDACEAE PEPE 506 AIKAI SACBUOR SEPARARE 4690 Ambrosia aptera coisas es RAS 68 Al DII ion Saha ELE heii 53 E var. elatior ..... 668 b EO ci SUR ar 534 emisiaefolia var. elatior f. lanceolata .......... eee ub AAA cope a RR ta 668 BASQUE Lies riis na pcd 534 ria of NEU. ad 668 NE DR 533 A er dT Alium un ausu dran cerdos s 501 bidentata x trifida ............ 668 1935] PALMER € STEYERMARK—— PLANTS OF MISSOURI a var. integrifolia ...... Amelanchier canadensis ........ canadensis var. Botryapium ... humilis re ee ee American Barberry ........... oklime P Spikonardı zen OP SIS Amianthium museaetoxieum .... Ammannia auriculata 222725. e... o. o. o 9 o 9 9 9 à n n 9 n n n5 frutieosa var. angustifolia ... frutieosa var. oblongifolia ... frutieosa var. tennesseensis .... Ampelopsis arborea ........... cordata ne ES Amphiachyris dracunculoides ... Amphiearpa braeteata ........ COMORAS Giese ee DNUS MNONOLCE CES © oe IRIS RS EPULORGTY,. DM: dee OH Amphilophis saccharoides ...... Amsinekia lycopsoides ......... msonia eiliata var. tenuifolia ... ontana REITER var. Gattingeri 620 Taber re var. salieifolia 620 591 ANACARDIACEAE .......... An es occidentalis Pa ex e. ae Se a Tet LREEET ER] 689 Page ne var. projeetus ........ 479 pU. tors LI 479 o ai TS Er PELO 479 provineialis var. chrysocomus ... 479 saccharoides «9 em ele 479 se var. divergens ....... 479 seoparius var. frequens ........ 479: scoparius var. genuinus ........ 479 seoparius var. villosissimus ..... 479 ternarius „onen 479 A etn 479 virginieus v EHUINUR 2. tt. > 479 virginicus var . totrasinehgus 479 Androsace occidentalis ..........-. 615 Anemone canadensis ............ 542 caroliniana ........ 4e eene 541 cylindrica .........«oo.oooo.o... 542 O RU A. DOO 541 virginiana ........o.ooo.oo..o... 542 E f. leucosepala ....... 542. RUM SCRA ORE ts CECI TER 542 Adis thalietroides ......... 541 Anethum graveolens ............. 612 Angelica villosa ................ 612 A ee ee RU 612 Angeles en oe ie ce 610 Anglepod ............ enne 622 ANNONACEAE .............-* 544 Anoda eristata var. brachyantha .. 598 Antennaria ampla .....c.......... 666 89Jopliy la. secsec en 666 ETE: O ees uis est OR 666. DOOR Sere eee qe ue pris 666. t|; ——————— E 666 plantaginifolia ............... 665: Anthemis arvensis var. agrestis ... 676 Dota aoi ne 76 A +... EUR S 676 Mb lem odoratum. v. 471 Anyehia canadensis ............. 534 polygonoides ............. es 534 Apera spica-venti ....ooooooo..o.. 465 Apios americana .........o..o..... 583 PUDEN er a a ta eek > 583 Apleetrum hyemale ............. 510 Aplopappus ciliatus ............. 658 ABOCHNAOBAER 6265 $5. vc ee 619 690 ANNALS OF THE Page Apoeynum androsaemifolium ..... 620 SERERE rinnen 620 eannabinum var. glaberrimum .. 620 cannabinum var. hypericifolium 620 cannabinum var. pubescens 620 hypericifolium ................ 620 hypericifolium var. cordigerum . 621 o AR CET 620 medium var. leuconeuron ...... 620 sylvaticum ....... eee 620 Mica ando UE La 556 Apsleof-Poera- AAA 638 PAA Tn ee ir 591 Aquilegia canadensis ............ 542 canadensis var. faviðora PEPEE 542 COCO: ¿cars 542 Arabidopsis Thaliana ............ 549 Arabis braehyearpa ............. 552 AURORA un 552 A A 551 Drummondii ................. 552 DIR iiia rrr rnb YER 553 OUCH. co kint cane A 552 ee ee ere er 552 1i MP 551 PASIOA AA RU 552 o A 552 o A nas Ce E 552 AR A O are ee 495 Aralia raeemosa ................ 609 INE IP Test 608 ARALIACEAE ................ 608 Arctium minus ................. 678 tomentosum .................. 678 Arenaria patula ................ 535 — ec 535 ricta var. texana ............ 535 Precisa BA A seas 546 intermedia ................... 546 OADE as ii REEL 546 pira Dracontium ........... 95 trip UN si 495 Ari stida u re 470 balBirAmeR ee 469 bromoides RN 470 DATEN + ecc adas 469 dichotoma ................... 469 MISSOURI BOTANICAL GARDEN [VoL. 22 Page dichotoma var. Curtissii ........ 469 fasoioulata ..... eee 470 EA 469 a sau 470 I MITTIT 470 EEE aiii oss 469 5 oir est 469 purpuraseans iii ad 470 ramosissima .................. 469 Aristolochia Serpentaria ......... 524 — var. hastata ....... 525 vicio ds 525 ARISTOLOCHIA CHA Ciena es 524 Armoracia Armoracia ........... 550 Arrhenatherum dite re ene 465 SPO A 495 Arrow-he Broad-leaved ................. 456 Long-beaked ................. 456 Short-beaked ................. 456 ARVOR WOO i... royal 651 Artemisia annua ................ 676 BUE O A stet 676 campestris subsp. — PET 676 campestris var. caudata ....... 676 E d REX x hs 677 caudata oossoo 676 dracunculoides .......... luu. 676 en var. glauca ........ 676 A — hx) 676 apa PA oda 677 TEN FERFIERE TET 677 Lindheimeria Via iT res Ys 677 ludoviciana of auth. ........... 676 cS eese 677 vulgaris var. gnaphalodes ...... 677 vulgaris var. ludoviciana ....... 676 vulgaris var. mexicana ........ 677 vulgaris subsp. Wrightü ........ 677 vulgaris var. Wrightii ......... 677 A re rn 677 Arthraxon hispidus var. erypta- BENE APA RE a TEE 479 Aruneus sylvester ............... 556 T gigantea ........... 458 crosperma II cece 458 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 691 Page Page Arundo Donax aa nti € p wise 462 Azureus X laevi o o sae es 662 Asarum eanadense var. acuminatum 524 commutatus- esine na 664 eanadense var. reflexum ........ CONCINNUS: oia cs a ae 663 ASCLEPIADACEAE ........... 621 COTOILOUUS: eres RR. 662 a amplexicaulis .......... 621 cordifolius var. polycephalus .... 662 Pen SO a 621 depauperatus var. parviceps ... 663 a IE O ME 622 Drümmondu- 0 o ue. 662 hnmistmata cn ee 621, 622 GUMOSUS: nia o en 664 incarnata oe ae ee En AE NIC a ericoldes cias are BR ee ae oe 663 Se ENS 622 enicordes of, Auth... denia. 663 UA E SS O E 621 ericoides var. platyphyllus .... 663 latifolia: ©... a ee 622 ericoides var. prostratus ....... 663 Mead 55.5 t TBI 622 ericoides var. villosus .......... 663 Derenms. a e AE a 622 OUVOWILS een ee TE ES 663 nhtolaccoides siese sean e ee 621 RUECALUS ne Mow ee 662 UT PULASCENS te ao 621 Matas 658 quadriroha c mE ene 622 LA EA 599v 662 A Oo er 621 a AAA E n 662 Btenophy.lla en ee sen: 622 O Winvere ee AO 665 Sullıvantıı 5: 0999 sonne 621 NZ RER RE SR 663 A O O OOO ISO Se 621 Laa A A «e 664 syriaca of auth. in part ........ 621 lateriflorus anth- Li. se ees 664 a var. kansana .......... 621 lateriflorus var. pendulus ...... 664 ABUS pod n ori oro Unione: 621 T var. pendulus x panto- chi f. uten. uem eis A NAAA AA s sea, 65 ari m e ea ONT 622 ee var. pendulus x prae- lata seen 622 AL ee EE 65 sele a VAT ICIS 224 50s ee 621 lateriflorus var. thyrsoideus 664 fenis hyperieoides ........... 599 BIHATETONUR IO 665 Ash PULSU Loue dene T 664 Bl ten nero stet 618 TRULTULOTHS ole see 663 GTO tada emus 617 multiflorus var. exiguus ........ 663 Prickly «c hene DS 587 multiflorus var. pansus ......... 663 Pumpkin- EM ee 618 Noy: England ........ shale oe 662 mc p D 617 pe ecu adc c nis AL y= Water tef e HD SS 587 OBlonmrfolns 1o lorc ele 662 TOS TRI 617 oblon z. var. rigidulus 662 'ASUNINA triloba oorsese tas eaaa 544 JA o MERSER EE deti CE 661 ASDAISQUS to eeN 503 paniculatus -eere eet 664 cama soei NAE a 503 E var. bellidiflorus ... 664 Aspe ieulatus var. simplex ....... 664 Er toothed ime micara ta cotes 513 i cH DUS 4... CRI o 664 Mi ucc O a 513 parviceps u... Se ee y Au 663 he oe DICT PIN 664 sd x pilosus var. demotus. 664 hnousbusc a ee, 664 1 CN un co: eo E rs e 662 anomalus senaat eaae Deae 662 ne Var TEN SE: Bean 662 AZUTEUB (c c m MN 662 BHORUS duce rco TS 663 692 ANNALS OF THE pilosus var. demotus ........... pilosus var. ty h polyphyllu nn sagittifolius salicifolius salicifolius var. subasper 8 RER: iod vd vx Panes oy Pp ee 4 Pests Flower Aureolaria calycosa flava var. macrantha ........... grandiflora var. cinerea ........ o9 9o» 9o o] 9 9 9 9 9 9 3 o n n | n n eer t] ] |! | | | |) d |n ]| ]|) |) | n "ONT. Axyris amaranthoides Azalea nudiflora wr] n ot ] || á À ...o........ +... +... Bacopa acuminata .............. rotundifolia (Vor. 22 MISSOURI BOTANICAL GARDEN Page Balsam-apple, wild .............. 653 BALSAMINACEAE ............ 544 Baneberry, white ............... 543 Baptisia australis of auth. ....... 575 bracteata of auth.............. 575 P yrtttasg POETE 575 CIT PED 575 leucophaea x vespertina ........ 575 leucophaea var. laevieaulis ..... 575 MBUNBFOGRYDE. iret erre Cos 575 0 silks kid RE dates 575 Barbarea stricta ................ 550 PESE anne TT 550 vulgaris var. longisiliquosa ..... 550 Barberr BS MM dor 544 BENE rr rere 544 DOANDO ireexiasMrsy vati 544 BST arce ERE do thes 480 DORÉ ea en 464 BEE anne 464 Banyasd Grass ns 470 Bastard Toadflax ............... 524 Batodendron andrachnaeforme 614 giauoesoen8 ...oomomm ent 614 Bayonet Grass ...... eere 485 BENE "Id ouisexkre»SEeretAv EA 584 Hee ATOR T 02 HEU TARE cir 478, 479 o RI EeePC 479 BERE Loupe ERRANT ii 479 A wxrk «Y edes 479 BED ausus siue DIE ed e ees 478 A A 639 Beckmannia erucaeformis of Am. BED A Ras aM 470 o aunt ek sass tev 470 straw TEIDE. Lisez xax on É 648 RUPTA. o ica lr cia 648 A 648 AAA ee 633 AA A va 517 A EIEEE TTT 517 Ds E LAHT MESE MN 645 Beef-steak Plant ................ 636 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 693 Page Beggar's Liee .......... 579, 580, 627 d ed o LOU 645 Beggars Ticks aan. en. A 7 E ice 645 E IA ANON 673 BIGNONIACERE E AUGE E 645 SWAMP! NN SE SEE SS 674 mdweed «oa RENNES 623 Belameanda chinensis ........... 507 Ee TCI E IRL 528 Bellflower Hedge sense ee 624 Creeping s o 653 mall £54 AR eee ars 624 lue AR 693% "Birch, River seis vss ee 517 Bellis integro 661 Bitter-sweet ..... 4 Es 592 Bellwort Bilterweed nun mI 675 AFRO: atada UE 501 Blbkbery. High-bush $. 4. 52$... 567 Small as AA 501 Black-eyed Susan ............... 664 Belvedere. ains a TEN EIS 029. »Bládder-mut -isece ode neh. 592 BOM Pee eaters E E 645 Bladder-pod, Slender ........... 644 Benzoin aestivale ........... e. 545 Bladderwort, Large ............. 547 aestivale var. pubescens ....... S45. Behr Biar ........ sys. 656 melissaefolium, oca 545 Blephilia ciliata .....25.......-.. 634 BERBERIDACEAE ............ 544 RUS UGE Lo secoes O ER 634 erberis canadensis ............. E MM A 546 en O IEEE c ols A PP 533 mE DAL- :BloodzO0U Waived uec rese DAD een scandens. un ee 595 ue Orgia TEXAN o E 600 a AAN AR 630 Bergamot; wild een o es 633 i: AA SEE 662 Berlandiera texana .............. 667 A EAS EM PEST 44 ermuda TAR eere ons 470: BNG ann dre vs 542, 653 Berterog!icana S edes 547 BiuebellB van anne Pees ee 628 Betula nigra Dra da 517 Blueberry, Low-bush ............ 614 BETUBACEA RM sas 516 as Grass: nenn 508 Bidens aristosa ninia atado rers 674 Blue-eyed-Mary ................. 639 aristosa var. Friteheyi ......... 674: Biue-Joint... sec. eo es 463 aristosa var. mutiea .......... 674-- Blue-Joit- TA 6e en 465 Beck? citer teh A 674 MO BOUL? ores: sow A M AT 479 bipinnata sea ovate 074. Blue-weed ........... 9 eee 629 Garinus A OO CIE A Ce E EA 648 cernua var. elliptica ........... 674 (SLUT LACREET weenie 648 cernua var. integra ............ 0140 AAA ER CCCo UIS 523 COMOBA™ ni Gud ol 673 Boehmeria cylindrica ............ 524 comosa var. acuta... nur. 9.06 673 Boltonia asteroides .............. 661 CORALA ta ld No 674 asteroides var. decurrens ...... 661 discoidea: er QR. 673 A A cs 661 DIT A a EL 673 an AA ee dere 661 involucrata er 2 ede 074 E AAA E Et 656 VIS uoo EE SENI m 674 a S Arne 656 vulgata ee A 673 A E 656 vulgata var. puberula .......... 673 )BORAQGINAORBAE ¿eric o... 627 [Vor. 22 694 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Page Potts- RUN Pes meer 464 Brown-eyed Susan .............. 66 Bonnes Bot IA 537 Brunniehia cirrhosa ............. 528 Bouteloua eurtipendula .......... 471 Buchloé dactyloides ............. 471 Co AR O 471 Buchnera americana ............ 644 a 2:40.40 10 eae ews SSeS ee E 7 ow . c per 649 eg — TTC 471 Buckeye reis 593 AI a en | nf penis erectum .......... 469 Red-flowered ................. 594 Bramia rotundifolia ............. 641 Buekthorn .............. sees 595 Brasenia Schreberi .............. 539 A AA 616 Brassica alba ................... 548. Hudkwhenb uses 528 ASPEN ir eR Ea à 548 o A diras xr oe aca d 528 eampestris .......oooooooo.oo.». 549 Climbing TUE Serer rere rrr ee 528 b RR ME TER Dur iris 637 A EEE nere nce d eX 548 M rare 1 RIEF 549 Bugeeed ............ ee 531, 532 Brauneria pallida ............... 070. - Bugbano der paradoxa ..... sees 670 = False ............. cece eee eee 541 DWUFDWM VG sis hnnc 670 gl Uh iii 635 Bread-root, Indian .............. 070 INI Ne ¿rior ores 636 Brickellia grandiflora ............ 656 Bulrush A 485 IAEA RETA 506 SA aac Se ERBEN. E mh 00088 SONA 506 ROWED 650 Kea eyed 485 Sensitive ...... eoo n 573 Bumelia lanuginosa ............. 616 Brome .. — EEEE 458 anl MAA 616 Bromus altissimus .............L. 458 Bunch-flower ................... 500 altissimus t incamus ... eese 458 Bupleurum rotundifolium ........ 611 BREVE RV Cy. KGa is 459 NER ns E en REEL d 458 Br Grass .....ooooooooomomososs = bari ee 458 Bur- head, pond ISA as commutatus ...........o..o.o.o... 400 BE oe ht 454, n hordeaceus of auth. ............ 458 Burdock ...eoeooceo sir nnl 678 IAN qur cir oe Ea ACER CES 458 Burning Bush .................. 592 A ee XE TT 458 Butter and Eggs ................ 639 JOD ara 459 Buttercup latiglumis ........... 0202000 458 CODO ee 540 latiglumis f. incanus .......... 458 BESBDOUW. erahnen 541 200 D vaa cce p RE ob See eee cr 458 ni A d b) T 44m bred 540 A su E messes 458 MN PA A 540 purgans var. latiglumis ........ 458 Mind WOOD. A 621 AQOMOBUB i eari ees cece eens 400 AA 514 DOADOR anna 458 Butter-weed .................... 677 o AR sao Wie 458 Button-bush ..........o..o.ooooo... 649 unioloides ....oooommooopossor.» 458 Button-weed ..........o.o.ooooo... 648 Brooklime, American ............ 642 AA TIL AT TERT TIT 648 Broussonetia papyrifera ......... 523 BIBODUID. ar aaa 648 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI C Page Cabomba earoliniana ............ 539 Cacalia atriplieifolia ............ 677 TENILOFMIR socere (ee dis v vie 677 BUAVEOIENS ....... «4. terno 677 tuberosa 2209: 4 980 ner 677 a we cag EOC ae 604 Cactus, Nipple .......o.ooo.o..o.... 604 EU canadensis var. Ma- COUDIADE in visiere aseo are ele iu 465 canadensis var. typica ......... 465 MELDANEE 5 occ esse es slate hs io eis 465 inexpansa var. brevior ......... 465 GCOWMAOnMO cle tn 465 Waban iit ice aaa 634 Calamovilfa longifolia ........... 465 allirhoë aleaeoldes 2.0 we eec 599 A A Y ARN 599 DEI ee See mee eh 599 involucrata ......... ree 598 O TT 598 triangulata ae cites M ee 598 HACHAD ee 590 deflexa var. Austini 590 LODO ee era 590 culopos pulehiollus ee are 509 Caltrop ciao v Vo» Syn 586 ea AM LI O Ud. 544 Camelina microcarpa ............ 548 AA 2 O OS 548 Camassia esculenta .............. 502 LEN A s Met a 502 Campanula americana ........... 653 rapunculoides ........oo.....: 653 OLUNGIL OMA canis. se ee zn 653 CAMPANULACEAE .......i... 653 Campion EOS ee 536 Td ono ee IET 536 Spin dto Attac O ete hp 537 White. a MEC 536 Campsie TATIONE crc a 645 Canary Gras err cet cate ie ots 471 Cancenroot. 0 en cote Soe 639 Mal 0 d ir ren tr 645 Cancer-weed nme een ees 633 WANCONWONG corno eere soie 639 695 Page Cane Giant ..... nes «Meer « 458 Sorghum .......... ed ern 480 witeh .....2 e CASAN eere 458 Cannabis sativa ...... de e eere 523 Capnoides campestris ........... 546 CAPPARIDACEAD EV. 552 CAPRIFOLIACEAE ..2..... 650 Capsella Bursa-pastoris .......... 548 CATAWAY .....99 es Rer eade 610 Cardamine arenicola ............ 551 DUIDOSa M nen 3 ose elo eer is 551 DouglasSl ....... petens 551 parviflora ...... cce beth nn 551 parviflora var. arenieola ....... 551 pennsylvanica .........o....... 551 Cardinal Flower .....« 488 696 $99 9 9 t] m om o 9 n 9 £9 9 9 t] 9 n n o n e... o oo... 39 9 oo à 9 à 9 $ t n n n, n Davisii decomposita NINE eek DNE eec ] | ||] 9] c] o£ n .Loo a er a .......... eo o oo. .osxp> són + +. +... .... "| ]|] ee 5 1|] | |]| | |] |] 9 n n ett | t] | |n EEE VEN WERE: HOLAAA AAA Ass eos. o s$s$a $$... .e.oo...o.oo.. 9 9 à à * d 9 o n n5 A eee re eee TE gravida var. laxifolia TS PTF Gray var. hispidula .......... dial, NP VaR PSCC RC SOHR OR OR OREO HE DS ee | | 9 n [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page BENDUM i 4h 05s 004000000004 493 N ddes bs tens dae 491 Le var. copulata ....... 491 n EET TRITT EIS EEEETTTT 491 "6 var, DIMIDO 5 isss 491 laxiflora var. gracillima ........ 491 laxiflora var. latifolia .......... 491 laxiflora var. — — 491 laxiflora var. varians .......... 491 lov illt TIER TEE 487 UT EEE TTTS 492 nau MTTTTTTTERERITTETT 494 A geom T 487 certi VIRA Creer yy T 494 Pe ree Peer eee re 494 pi var. pedunculata ...... 494 aa is 494 ride var. MOOG ea 494 MEUM .iossdoteeciskasncuns 492 i Ways apMRSERATAAAM TET 490 o PA 487 mierorhyneha .......iooooom»... 490 EMEN A vectus 489 mirabilis var. perlonga ........ 489 ind MEA AI 489 unc N RE 494 weed ho alu 487 Muhlenbergii var. enervis ...... 487 muskingumensis .............. 489 nigro-marginata .............. 490 o 555655 IRAE EO SCR 489 oklahomensis ................. 488 DNE PR vise 491 EE ipai A 492 pennsylvanica of auth. in part .. 490 pennsylvaniea var. digyna ...... 490 — $3239 244354 Pond cs 490 TREE 487 E a ee in Kee 487 s A t cc eee 489 a A cepa 359 346 Xia 490 y EA P 487 retroflexa var. texensis ........ 487 Be a 493 riparia var. impressa .......... 493 riparia var. lacustris .......... 493 oe rr A O 487 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 697 Page ‚Page rosea of auth, ....... eer er 48 Carpinus earoliniana ............ 517 TOS0G VAT. MNOF vegas eee e se 487 Carrion-flower .....-.9B8 eissis» 505 Martwellil. 5459599. SIRO ASIA URITOL na SNR Re nod 612 SCHDIAUA | cca cee see te eum Sen 493 Carum Caryi ......-23 M e cesses 610 Sehweinitzil--$05 2942 co 494 Carya alba ......... even 515 J ET RE OAA, 488 alba var. flooides E: NN 515 BCOPRTIB A ee 489 Alba. var. ovoidea. een 515 scoparia var. moniliformis ..... 489 alba var. subcoreacea .......... 515 setacea var. ambigua .......... 488 SQUALIOR aaa ote SINE 514 ShOruüang Nee ee es 492 Buckleyi var. arkansana ....... 515 a AS IO E 491 Buckleyi var. villosa .......... 515 BICCALE Nc eet rete eee oie sie E 487 GOTAIEOTIDIS Teresa oe ees 514 ET entente e e e m 488 alt var. latifolia ....... 514 A A une 494 BINDER e e 515 Fa sje SERVIS Teide ve eoe re 488 Eo var. villosa ............ 515 stramimea- coc eB Te 489 PROON ¿ic neen 514 straminea of Am. auth. ........ 489 feines AT erem e T e eerte 514 EULICUR Ao ee ris 493 HHNCPODOFDO EAS: MA A 516 stricta var. angustata .......... 493 111 heart A NDR TER TER 515 stricta var. decora ............ 493 ovalis var. obeordata .......... 516 GOTH v o DU cols SIUE 489 ovalis var. obeordata f. vestita 516 TOLANICH ote es 490 ovalis var. obovalis ........ 5... 515 tetanica var. Meadü ........... 490 Via var. odoraia $e s 516 tetanica var. Woodü .......... AD) = “COVED LU Coe ces ote aep sere es 514 toxenmsd TE REESE 487 E var. ellipsoidalis ......... 514 (LU b T ROCK IO IU 493 A O c 514 AQUA Ve cier a's ee 488 ud A. AA 515 a ODA MS 489 tomentosa f. ficoides .......... 515 tribuloides var. sangamonensis .. 489 ve. f. 0voldega. 22.0.0..0% 515 triceps var. hirsuta ............ 492 entosa var. subcoreacea .... 515 triceps var. Smith ............ 492 CARYOPIYLLA CEAH se. 34 irichocarpa oro a) ote sts ele ater 493 € aecrista of auth. ..... 574 ichocarpa var. aristata AN e 493 E us var. robusta ...... 574 trichocarpa var. Deweyi ........ 493 MOGI serlo eremi ber 574 trichocarpa var. imberbis ...... 493 IascioulBta D... Bee 574 OPENA oen 494 fasciculata var. depressa ....... 574 TY DANOS ii TA. 494 fasciculata ensem A A 574 umbellata s S90 coe eek ters ore 490 asciculata var. robusta ........ 574 ORO 490 marilandiea sse co Rees ober 574 varia var. colorata ............ 490 Medageri ee 574 VOBICATIA Tee tere 494 CLIO MN ee 575 vesicaria var. monile ........... 494 MCCINOUUALIG. 0. E E eet 574 VIFGSCONG een ec seele A AS NDS 574 vulbinoidon tes. a le A 488 Castalia OUoraia EEES: EI 538 sic var. annectens . 488 AA A REN 538 o O AO 490 Castanea ozarkensis ............. 917 hes ined PS IPLE E PES 53 pumila of auth, in part ........ 517 [Vor. 22 698 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Page Castilleja coccinea .............. 644 NENNEN PR ese 679 coccinea f. lutescens ........... 644 NRORIDABNA- ras riera 679 A T" 644 Centaurium ealyeosum ........... 619 Ctr iibi: MEENE POEET TET 644 texense ......cccccccccecccece 619 Castor-oil Plant ................ 589 Phe O L TT ee 619 COME SPrrereeye Tyee err ee ere es 645 Centunculus minimus ............ 616 bignonioides ........+..+.++55- 645 — occidentalis ........ 649 "uo rio eC A 645 occidentalis var. pubescens ..... 649 a MPH ae 506 Cerastium arvense var. oblongi- Catehfl FRUI S IS TELE Rs 536 Night-flowering ..........+...- 536 braehypodum ........... sess 535 Royal ......... cece eee eee 537 pg vr APO Cn 535 BROWN ee 536 viseosum ......ccccccccccccece 536 A D'UN AS 471 en N 535 S a ea OT 632 vulgatum var. hirsutum ........ 535 Cat-tail CERATOPHYLLACEAE ........ 538 COMMON anna 454 Ceratophyllum demersum ........ 538 Narrow-leaved ................ 454 Qereis canadensis ..........o.o... 375 Caulophyllum thalictroides ....... 544 Chaerophyllum procumbens MEHR 609 Cayaponia grandifolia ........... 652 procumbens var. Shortii ........ 609 Ceanothus americanus ........... 595 NR Sra aera ee tr 609 UEM Loisuecsoséesecusseseces 595 Tainturieri var. floridanum 610 ovatus var. pubescens .......... 595 bore d MERE REST CET TTE TT 609 Cedar DENM acidity 616 TETAS 454 T depressa ........... 574 BH viai ae 454 [asciculata een 574 Celandine ............. sees 545 Chnetopapa asteroides .......... 661 CELASTRACEAE .............. 592 Chamom Celastrus seandens .............. 592 EE TETTETETT 676 Celtis vc EN T TT 523 A O 676 BOUM RR 522 A ERAEDI ERES SE RÉ 676 a VAT. LOZADA .......... E AMS 548 mississippiensis ....oooooooo o... 522 a ON da 458 DOCIGENTANIG: «¿ac us» FER E A dar 598 occidentalis var, canina ........ 522 Chelidonium majus .............. 545 eer var. crassifolia .... 522 Chelone glabra ................. 640 PNEU ads Tom 523 obliqua var. speeiosa .......... 640 umila var: DOMO iaces 523 CHENOPODIACEAE ........... 528 pumila var. georgiana ......... 523 Chenopodium album ............. 529 Cenchrus carolinianus ........... 478 a "Tm 529 DANSE 200a i nr a CR RR 478 album f. laneeolatum .......... 529 Centaurea amerieana ............ 679 inum T. viride 2.0.5. en 529 NE ILI P T VE 679 A AAA 529 O 679 BMUPONGIGGE: acid 529 DEDI AA 679 ambrosioides var. anthelminticum 529 PEE. AA PA 89 0 PRA 1935] PALMER € STEYERMARK—-PLANTS OF MISSOURI Page Berlandieri subsp. Boscianum .. 530 Be Po Zschaekei var. PT eee WR OM 530 LN = Zschackei var trs MERERI 30 a subsp. Sk var. typieum f. angustius ........ 30 E ERA Zahn var. typicum ni o er 530 Berlandieri eodd pars var. m f. negleetum ........ 30 pd subsp. Zschackei var. typicum f. peduneulare ...... 530 BOIAN ores ee irse 6 530 MACAO ee eR E sonore 529 Bushisnum 9.9955 iio inte 530 capitatum ....... eee ero 529 BATUM 5909-19 SS 529 ER o 531 gigantospermum ....o.ooooo.... 29 an var. Standley- Vive CXV s wa Va ROCA 529 glaueophyilam sdb oc tie tote 530 A CR 529 desc of Am. auth. .. 2... 0%. 529 incanum ....... 0 toot 531 leptophyllum re see ee 531 TeDtopnYllum i See eee sen 531 leptophyllum var. leptophylloides 531 531 leptophyllum var. oblongifolium. 531 MIBSOUTIONBE ER 00395 er seen ee: 53 missouriense var. Bushianum ... 530 murale ve reo da 531 PALANCA o TEE 529 GJR AOS 530 pratenicola sao a aa o 530 Standleyanum .....oooooo..o...o. 530 oi RD 531 urbicum var. intermedium ...... 531 x variabile var. Murrii ........ 531 Cherr Choko on PECES 57 Ground ie 637, 638 Indian cae TT 595 A A IE 573 NOU lalo led 573 O S UNLESS 572 699 Page Chervil Hemlock ..... eris 612 Wald oia ea 609 CHESS 36.6 eos woes o rta 458 Field i27. 19 e an EE RS 459 Soft... vvv E S 458 Wald V. cover 458 Chiekweed ........ pe der n 535 lammy "ccoo da vae ann 536 Forked 2 4 ve een Dev ele 534 TORE 2002504 RICE ISI 535 Nodding ......... tton 535 A A DAL. 680 Chinquapin ............ nn 517 Cfr MIA E PU CEP 538 a virginica ........... 618 Ghittim-wood .......... e er es 616 Chloris Eon: WER Hee 471 ILU wmv O eee 471 y | 5. cis sve ON 471 Piin Holly peca pe ues ees 592 Chrosperma muscaetorieum ....... 500 don ost Leucanthem . 676 Leu mum var. . pinnatitdum 676 cimi T Berlandie > AR Ae LEE 58 mporum ....... nnt 658 he A RR ass 658 villosa of auth, ............o... 658 llosa var. canescens ......... 658 Cichorium Intybus .............. 680 Cicuta maculata .....2.. een 610 CURE A valve days es 645 imie iis uga racemosa ............ 543 Cinna arundinaeea .............. 466 Cinquefoll ..........21 4 eene 566 Cireaca sms A. EOS 3 608 lu ma of Am. auth, 22.534... 608 Died. seus E UAE 679 AVOIR 4.224.295 SE 679 E 2.2.2... eee I rts 678 Hill CIS: ET Dr 678 LU OA 679 lanceolatum PONES aeo e cree 678 A eee Seer Ope 679 en var. Hölleri ...... 679 o O ERRE eR TS 678 DUAE erred es 678 700 Page undulatum var. megacephalum .. 678 a AAA 679 Cissus Ampelopsis .............. 596 ch, EEE 596 MUN. en 596 GUTAORBAR. rar ae 601 Citrullus vulgaris ............... 653 Cladothrix lanuginosa ........... 533 Cladrastis lutea ................. 575 Clammy Hedge Hyssop .......... 641 Clammy-weed ........ooooooo.... 552 — multicaulis var. robusta . 537 AITE LEA KE d RE 537 ie A TER 537 virginiea f. robusta ........... 537 ke’, SER ET TEC ECCT Tee 633 Clearweed ..............seeeses 524 RI is TETTE TT T 648 Clematis erispa ................. 542 A AMA 542 missouriensis 6.6.6... eee eee 542 FEE ee er ee ye 542 VEN N ns d wa 542 cr hr eerie aren re te 542 TE ree 542 virginiana var. missouriensis 542 Cleome serrulata ................ 52 onec TTET ETT 552 Clitoria mariana ................ 584 Clover A EP YT TITO 576 onn e 575 ncn RR TOS 576 NN A 576 MB Leere 581 bo EN TT 582 DNE ae 582 DENEN. ea cleans. 575 enu NETTE T TERTIO TET T 582 A EIE TT TET TT TEE ETT TT 576 DEDE Lobos pnkPes e E 576 — Ur E Des OPE 577 hl PP 578 Rabbit Ps Taupe Pe es MS 575 ANA AN 575 er AA 576 MEM A pec ER 576 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page SENE AAA 576 sinad au era 576 Club-rush, Hedgehog ............ 482 pair aerolineas ces 544 BT ren 668 oos duo: VEVRTTTTEEITERTTS 557 oo Eoi AA 480 Coelorachis cylindrica ........... 480 Coffee-tree, Kentucky ........... 573 QUEE - Well ici 574 Cogswellia daucifolia ............ 611 CO DE rr rr 544 Collinsia Li ee ses 639 OEECATRTVITTTTIDIILTOL Y 639 Collinsonia eanadensis ........... 636 Collomia linearis ................ 626 Columbine, Wild ................ 542 Columbo, American ............. 619 Comandra Richardsiana .......... 524 KENN arena 524 CEDE AA 628 Commelina caroliniana ........... 497 A diui ae 496 N 497 BEE NT 497 éreeta VAr. erispa ............. 497 hirtella of auth. ............... 497 MUN sedas 497 VLA BERET CT CULT TPE TCT e TT. 497 Vlil iiie er e kir 497 virginica of auth. ............. 497 COMMELINACEAE ............ 496 Compass Plant SA 666 COMPOSITA B uns 654 Cone-flower, Purple ............. 670 Ben chinense ........... 612 m maculatum .............. 610 en alude. CERT XO CORO NO 641 Conringia orientalis ............. 549 CONVOLVULACEAE .......... 623 Convolvulus arvensis ............ 624 arvensis var. obtusifolius ...... 624 REN are ET 624 | RA 624 sepium var. fraterniflorus ...... 624 sepium var. pubescens ......... 624 o Loi MMC 623 1935] Page COTA TOOL EE Te eme Ime 510 rontod ER 511 Sa A THE M 510 NE EpL dr RE, 510 Cora odontorhiza ........ 510 Wo E cum cce 510 copa ER Mec ADS 510 Coreopsis crassifolia ...........- 673 ADAMO x foros = eters oe ote 673 lanceolata. eee ee 673 lanceolata var. villosa ......... 673 palmata .....ooooooomoomo.»..» 673 pubescens ......o.ooooooooo»..» 673 SIRCIOEIB ee ee ee eie 672 triptenB: 2.2: onde eio eoe 673 tripteris var. Deamii .......... 673 QGoriander ee ee ae 611 Coriandrum sativum ............. 611 Corispermum an porke 532 O OS er 531 orientale var. emarginatum 532 Cor 000 CENTER eade eei 513 Corn, Indian e 955 9s 480 GornoGOcklo: vo EE eem 536 Gromwell $e eee cies eee een 628 Salad, Beaked ...............- 652 OO e 534 CORNAGCRAD australis. er noms 612 Cornus alternifolia .............. 613 Amomum of auth. ............. 613 M aim noe Pa oor 613 ED ESSEN METTE 613 PER of auto e eet: 613 Borda vete sees due esr 612 florida. f. IUDIS sc. 612 florida var. rubra ............. 612 obliqua d mS E a es 613 pantoulalta usais le 613 PUG a as quae 613 TACOMOSA o aja ae 613 A T S D A 613 abide Varna 579 Coronopus didymus ............. 548 Corydalis aurea .......... eee 547 ea var. occidentalis ......... 547 Gampo tria cs er 546 SOS EE 546 701 Page flavula ss scarcer tees Re 546 micrantha .... v.e es 546 lanum ea ER Oe 547 Corylus americana .............. 516 americana var. inca O americana f. missouriensis ..... 516 eornuta- 6s. cbt sis os ERST IT 516 rostrata o OAM BONS S 516 Cotinus americanus ......:.... 0: 591 Cottonwood VV UTERE TAS 513 A A I 513 Couch Grass 7.220. ico Bete SS 462 Cow Parsnip «2.6.9 nte ven 12 Cowbane occ cc 0.0 f ES 610, 612 Gow-herb 1.2.0149 NE eder s. 537 Cowslip, Virginia ..............- 628 Crab, American .......v.......: 557 Crab-apple, Wild ...............- 556 BI A O 586 ORASSULACEAE ..........:%. 552 Crataegus acutifolia ............. 558 DIDOBTROVYO «o. «cd ¿cr gs 557 A pene cr IA OR 562 Tc MOOD Rd 560 di Ax e duree Vl es i NN TS 562 pb xxr c ME T 564 apiifolia var. flavanthera ...... 564 ORO il A ON 559 OPDOFOBCERE ooo cc eye ee re 560 e A O T 562 A ne SI PES LEE 563 A ee 561 A . A is 559 Darrottiana A A 558 E TN E es 558 biltmoreana AAA 0 nn 561 liypoda- ive ca wees 562 COMETA: aa 562 CUalltaniha. ee EIL 562 OGUUCONDA ss Aree. T 564 ODIA: o uio esr 557 ealpodendron ............. 64 ealpodendron var. hispidula .... 564 ealpodendron var. obesa ....... 564 A olii vin ew eh eene 565 A A S ee rmn 557 coseinioldes ro ree 564 [Vor. 22 702 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Page RN A 560 incaedua ............ es Irae collina var. secta ............ . 560 infesta. EE I 559 collina var. so ie. VOD 560 ns Au a TELESE PEET 564 collina var. succincta ......... 560 ca T PEARERA 558 DOLINA x viridis (i.i sr a 565 RD Lu a anne TT TT 558 n b LS NER 562 O A TTC TETOTTT T 563 [1,77 MEME 557 lanceolata ............ eee 561 hiuc 8s RC ER ICT ETT ee T. 564 n aut E TU TEETTTTTU DTP TOT TT 563 RI vado Wess was 557 o PA QM KA 563 erus-galli var. — TEET 558 BR O 560 crus-galli var. bellica ......... 558 lasiantha EE E T COT TERRE REET 563 erus-galli var. ben T 557 sucus ver 560 erus-galli var. I OTT 558 — CETT ERE TT 558 erus-galli var. macra .......... 557 PEN. irrita 561 erus-galli var. shi e... OOS it Aa 558 erus-galli f. truncata ....... 557 AA 560 BEEN AAA en ene EPI 559 AR 563 Danielsi f. glabra ............. 559 ludoviciensis ..........oooooo.o.. 558 COTA IA 563 lut PEET O eee | AAA ERAS 560 A BOB Dawsoniana ...... eee 561 Mackenzii var, bracteata ...... 562 declivitatis ................... 564 A Ete icis 557 NITET 562 macracantha war. neofluvialis 565 CI TP 558 macrophylla ........... sese GENS. OT TTT TCT ee 562 BONUM ann rare 560 disjuneta var, magnifolia ...... 562 magnifolia .... eee 562 ONE 563 A EET TE T TTTTTTTTTTTTTT 561 DUNEDIN vedi eX dS ENS 563 BENIN Senn 564 SU EI 557 missouriensis ......oooooo oo... 565 A ARA, 557 mollicula .......oooooooooo... 564 LA TT 559 eee eius o MN ingelmannii f, nuda .......... 559 mollis f. dumetosa ............ 563 Engelmannii x uniflora ........ 565 MONOEPOTMNG su nennen 7 ONE aa 565 a ee 559 vss OTe ee E TT 558 DINI ae 561 LL o o a Rd A 557 neoffwialia ......oooo momo...» 565 A TETTE TT 559 c E TTE T, 561 ia Ra 557 A BE ors een 560 DOMO isa 563 o A A 562 nee 565 glabrifolia euere 561 En ER O A 564 A a ee 564 a E A o NÉ A A dA p sad aa x 557 a EEE NET TIEREN ER 562 AN T 559 CLA EET ITTTTTTT TTE TO T 561 bus Oks dart us rue hs 559 pachyphylla .................. 558 a IW eee ds cha 560 AAA 561 a ERNEST SEEN 564 padifolia var. inearnata ........ 561 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI ........ 9 m 9 9 9 9 t9 9 n t n ong n UU o .. PIC UNOS ee ee pruinosa Ed angulata pruinosa var. brachypoda pudens rubrifolia rubrisepala spa spathulata var. flavanthera .... speciosa ........... +... ct | n CLOSER e Enn SDUMIORG SS A OE succule e... oo... 39 9 c 9 9 9 3 9 9 3 9 3 n tardiflora viridis var. viridis var. lanceolata viridis var. lutensis Cro ort Gvtataria sagittalis Croton WM d ..o no... o... ......+. +... +. ..oo....o..»............. ...o..o.......o..+..... ..o..o.............o.os .e...o.......o.oo.o.....».o e...—Á.o.... o... ..... o... ..os. 942849 545059o065»$099686695899€99 ..........+.. || css t] | n9 E c t 9 9 o n 9 3 3 Ss... 9 9 9 9 9 e 9 9o» 9 o9 o o9 9 9 9 o9 om t9 o n 9 9 n 9 n 9 n ..oo ..oo..s. eec 2 2 9 f. ..........».».». .... +... .......... ee ESASE -AYARLA ...oo.o....o.o.o..o.o.......s. n e 9 9 $ n o9 o 9 o» 9 9 * 3 n 9 9 s n o9 9 t n PI .......o...s eo .n..o............».»..s ...o... o. à à a 9 c n o n n n ngelman Seld var. septentrionalis . monanthogynus texensis .oo...... +... +...» & o9 o... 3» 9 9 9 P 39 9 9 3 9» 9 9 * 9 9 n [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Cynoglossum offieinale ........... 627 A iiiésseecitecevbss 627 Cynoseiadium digitatum ......... 611 A MAA 611 Cynthia virginica .......oo.o.o... 680 OXFEBACBAN ¿0.100.030 480 Cyperus acuminatus ............. 481 NE E TET TETTE T CETTE Te 480 IPP TIT11015. 482 SOMDIOBEUS imss. ...5...0. 480 Minis nd cs 480 dipsaciformis ................ 482 OS. Sunset 482 DA ici ra id n 481 ar IP 481 th ia var. pumilus ...... 481 DIN ¿oa Y esculentus var. angustispicatus .. 481 esculentus var, longispicatus .... 481 f 227 ara aa ITO: 481 ferae MR A A 481 Fiir iE Tee aa ee 481 EHI, id O 482 fflieuinis var. macilentus ...... 482 A Loi ER epe 480 Used io ev us dh V Oe 480 BENE AR eee eee ee eee 482 UO PP OT 481 O a 3S 481 O E T as eee 480 lancastriensis IRTE 482 N aci 2*8 482 cloro var, robustus .......... 482 pseudovegetus ................ 481 o MERERETUR EET T 482 a IUE A vie ee as 482 rofractus of auth. .......... 482 rad erry SEEN 480 Bohweinitzli ........... nn 481 8 Ma ERR 481 o t E T ICE T T TTE TT E E T EE 481 i dali var. capitatus ........ 481 trig . compositus ...... 481 nr var. elongatus ........ 481 strigosus var. robustior ........ 481 Cypress (D Al 454 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI Pa Blandhig. .. ty 529, 626 Bummer. 2.5. 69 e ST 528 Cypress Vine: in ee ota deed 623 ida dd candidum ssr errei 508 hirsutum iaa ees 508 a d cr ORIG ede LION 508 parviflorum var. pubeseens ..... 508 Regiis T. celos ee 508 D Dactylis glomerata .....com...... 462 Michgelmas meeer oe tS 665 (0 a A E RETURN O 676 UDC om ae ee 662 WEBLELN ee APPLE Sn 661 Dalea alopeeuroides ............. 577 enneandra 4 wee ane een ie 577 Dandelion Se res ee erem 680 SWEDE 0 eir er 680 MAI aaa eee 682 Ned 900066 A 680 Danthonia spicata .............. 465 RE TE 464 mani macrophylla .......... 642 xoti T RUNS e ys 643 NU SORTE OR 643 pectinata E EIE T 643 ALA pco E E E 642 Br Moto ia ti eee 638 SLAM O ee ee 638 Ul&- ui. 638 Daucus, Carota ne... tete ado 612 ALUMS Se ee 612 DAY flower” Sure 497 Oreopimp sr cate css NIE 496 Day. Lily: ees Vale ee ee 502 Deer:grüss hoes ieee ie 605 Deerberry naaa cei ates oe xt 614 an ATA iis UE ste one 543 o E EET 543 azureum var. Nortonianum 543 BONBONS a 543 OFTOMARUM cir 54 OUR OU ics A ns 543 705 Page Treleasel une wen ter 543 trI1Corne... 9. eS woe ee 543 Dentaria laciniata... P o u: 551 laeiniata var. integra .......... 551 Deseurainia canescens ........... 549 a o. v BIS x ee 549 miermedis. 4:220. Vers 549 Mia... PIN. 549 Desmanthus illinoensis ........... 573 Jeptolobus'......... Sen are 573 yi vds aeuminatum ......... 579 DOUM A ENS 580 en var. longifolium 580 BEDRÄENSE A A T 581 eanadense x Dillenii .......... 581 CHnONCODB E A es 580 eaneseens var. hirsutum ........ 580 A A Eee 580 O A. e es 579 in A A 580 ITALIA ina 580 Iarliandicum E EEEE ET 581 IRA Drum: ae ae 579 po oos ecu ire ns 581 oshroleneum occorre e 580 panieui&tum 5:8 5.75 e o Ev a 580 panieulatum var. angustifolium . 580 panieulatum var. pubens ....... 580 paucorum ...:...4 he 579 THidume entonado es 581 FOUUBOQ £o mm. «es ooo ee ev s 580 BeseiBbfolium es. A AO 580 YIPHBHOPUm too 580 A O deo eM 646 o A US EN O 568 pistes AMETIGANA ¿orinar imss 646 A e A RE 646 Dianthus ATmerla..... poema es 536 DSrDALDUS cio ee 536 as E eM EA eR ole 536 Diapedium brachiatum ........... 646 Diarina- féstueoides urn“ 461 Diarrhena americana ....oo.o.o.... 461 UIS En nee eae ae 461 Dieentra eanadensis ............. 546 Cuocullarig- uos cc NOE Ins 546 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Dodeeatheon Meadia ............ 616 ae Er ee 620 Pink. a EIS Hob PENES 620 Dog Hl o PET PI ee 676 uA A T 675 Do Alternate-leaved .............. 612 LE TEE. 612 E E 612 Rough: ORTE. ee 612 AROSE PN ead COS 612 PT eer rere Ra 612 Dotichotus 2 rd 585 a PIT 585 a racer ae 547 EE E 547 dips var. micrantha ..... 547 MAMMA IAN 547 HESS ir 547 reptans var. mierantha ........ 547 ar rn 547 Draeocephalum parviflorum ...... 632 PRON HOM ia 632 eer TT TER EET T 632 Dropseed Grass ............ 467, 468 Duchesnea indiea ............... 566 o ee 495 rx io REEETITTTTOZUZTTTO T 495 ZEN ira 495 Dulichium arundinaceum ........ 482 Dutehman ü Breeches ...::.. 5 546 eer eT TEE EN 525 Da, DADPDER ici csee8 675 E HBENAOCEAN eaaet us 616 Echinaeea angustifolia .......... 670 EN. AD S 670 DE BUGXA- 12.2239 8 T va vv ao 670 MESA. ¿ini es dx vex» 670 giovani. A AO 478 A oca Po eRES 477 erus- "T f: = Deke nea 477 rere er eiTe 478 e OUTRE 477 1935] PALMER & STEYERMARK— PLANTS OF MISSOURI 707 ge Page muricata var. microstachya ..... 478 d vus ER ae 52 ocotdentiala See ccs os eco tins wee Bock |... 208 UTE Oxo 522 PENGONS 4... Rx A ied 477 Slippery 2.2.2000 Seren. 522 pungens var. ludovieiana ...... 478 Aller ll O 522 pungens var. microstachya ..... 478 16 462 oie e cr OSO DE 522 pungens var. occidentalis ...... 477 Winged) 2... 000 0. ees 522 Eehinoeystis lobata ............. 653 Elodea canadensis of auth. ....... 457 Echinodorus cordifolius O O O. 457 Eleusine indica .......+......... 470 cordifolius var. lanceolatus ..... 457 lymus arkansanus ......... e 463 Indieans 223 9 0 01) eive 457 australis .....ooocoosoosorsso» 464 AAA OOO ao ao 457 australis var. glabriflorus ...... 464 chium vulgare T aan ee ee 629 CANRdOnsis8 5.0.25. NEUSS oe 463 Belipta AIDA ceci 0, 000 669 eanadensis f. glaueifolius ...... 463 Helgrass: ....:10 1 vc v een 457 QUGDTE LOTUS... cites on ele ieee ole 464 Eichhornia crassipes ............ 497 glaucwolius anche 463 ABA SS ers rer 604 glaucifolius var. crescendus ..... 463 Elaeagnus angustifolia .......... 604 en var. robustus ...... 463 Blatine americana ...... 9. 995 AUS EIER RESO ORO MIO EI MOL 463 EPATENA CIA orn 600 hiroutigtumis ER MUS IS HM 463 der; “Marsh ets 00 9 Se ele 667 A E 260s 91 8 Er 463 Elderporry Pre See CI er 652 SVE EC GUILT Su care 463 ed are a hanes MOOD DCN aces WOB Vd e. ao cM ss costar 463 Blecampaney Sry 666 pobubtua: oe eve New oe cs 463 Biophant’s fool... urn 655 robustus var. vestitus ......o.... 463 Eleocharis acicularis var. graciles SIUE es abe. wats ne 463 O O E eek ae ee striatus var. arkansanus ....... 463 acicularis var. typica .......... A ETA O O 463 ACUMMALA oe ee ces nee 483 villosus f. arkansanus ......... 463 ED Deor asis 483 VISIONE Lares das OR eapitata var. verrueosa ........ 483 virginicus var. glabriflorus ..... 464 COMPTE a IPIS 483 virginicus var. glabriflorus f. aus- Engelmanni Coe A ops 483 A AO 464 Engelmanni var. robusta ...... 483 virginieus f. hirsutiglumis ...... 63 ]anceolata ord sies eto 483 us var. hirsutiglumis ... 463 macrostach ya a ae 483 virginieus var. jejunus ........ 463 lata T re: 483 virginicus var. submuticus ..... 464 biusam. oce EE MORIS 482 Empress Tree ........4 Pee 640 palustris var. glaucescens ...... 483 Enblaater? 8 Nightshade AES TE 608 parvula sr scence seca san ea eee 483 Engelmannia pinnatifida ........ 667 quadrangulata s. on aee 482 Epifagus virginiana ............ 645 A roue vds 483 Epilobium densum .............. 606 lonas. SO IT TO 483 eoloratam ...:... AA 606 Elephantopus earolinianus ....... 655 Mores Intifolin- .. cite ce an 509 Tlisia^Nyctelea d. ern 626 —— pubescens -assii aes cess 510 Elm pales la capillario Feit qe C 460 American te 500 0,0009 iss «fs 522 COLOLUMIOMO Hoje ces ed Seca tene 461 Cork O Re 522 E O A dere ve 461 708 Pa os eer ery rT TT CORTE S 461 ZH ere rr eT Ter ee rere ys 460 UN kgs TTTTRTUT ITO ETT T 461 MEN. an ara en 460 2 ana es biee e's os 461 J) aa ers are T T TT TEE 461 A AOS 461 megasiachya EE ET TTT 461 COMO aa i rie 461 ses N DEAN eu ere re 461 pectinacea of Am. auth, ........ 461 pectinacea var. spectabilis ...... 461 BENE. RETTET 461 DOREEN E T ER ETT ETTETTITTTT 461 I: EE TETTSITIEUTETTETTT 461 oh! A T 460 inte utes AAA 461 AR 61 Weigeltiana .................. 460 — hieracifolia of auth. in VIT LTES LUT TET 677 un var. intermedia ..... 677 Erianthus alopecuroides ......... 478 DUM iria 478 min WETTE EEEF TT T 478 DECRG ¿ri Cote) 613 Erigenia bulbosa ................ 609 Erigeron annuus ................ 665 IE ET TT TTT IO 665 OVEM. PT 665 philadelphieus ................ 665 A Lisci cresi eestor 665 O 2425421: CE X CIN 665 ramosus var. Beyrichii ......... 665 cr E Vaud XA CER AN 665 Eriochloa eontraeta ............. 472 DER oecvest th aa A TEE 472 Eriogonum longifolium .......... 525 Erodium eieutarium ............. 586 MR MED. oues vias ci E 548 Erucastrum Polliehii ............ 548 Eryngium prostratum ........... 609 A MAA 609 Erysimum arkansanum ........... 549 A Oe Rae ee ae re 549 cheiranthoides ................ 549 a PAPA 549 [VoL. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Erythronium albidum ........... 502 A E T S ETT T T EE E TE TT T ET 502 mnesochoreum ......... nnn 502 Eulophus americanus ............ 611 en altissimum .......... 656 eoelestinum ........oo.ooo...».... 656 lien var. semiserratum 655 nno naar ae 655 MODO: ee 655 et rosae 656 o anni 656 perfoliatum var. euneatum ..... 656 BRSDOFOUM Mi S 655 eméserratum ¿....oooooo.o.oo.o». 655 EUN icone voor 656 o MAA 656 urticaefolium ................. 656 urticaefolium var. villicaule . 656 uphorbia commutata ........... 590 "0 E 590 A AA 590 AT 590 dietyosperma ................. 590 PIV OCOSDETING rr da 589 |ohtahé zn LI +. ¿;+0 ine an 590 o MAPA 589 BEA 5 ip E ee VERTES 589 RRA 590 BENE ci ViEREPLED 589 simi EDT 590 ds A IS ERIT TT 589 o MERE ETETETT ESETET ETET 589 d uri i iaaio 589 EUPHORBIACEAE ............ 588 Euthamia glutinosa .............. 661 gymnospermoides ............ s. 661 ee TT 666 o D e Ae 666 Evolvulus alsinioides ............ 623 o nennen 623 Evonymus americanus ........... 592 ig AMA 592 a BERT TET TT TTTITTTT T TT TT 592 F FABACRBABR diari cid 517 Fagopyrum esculentum .......... 528 1935] PALMER € STEYERMARK— PLANTS OF MISSOURI 709 Page Page Fagus grandifolia .............. 517 Ts ere er us 48 grandifolia var. caroliniana .... 517 mucronulata re.as de one 484 Faleata comosa .......... eee 584 puborua a us vM es 484 OME. acca vs ERAI ee 584 lH re vo RA eee 484 False Aloe ocres e etis poo . Finger Grass 2... Re 472 ONEBEL ce o dU 656 lender... o0 es 472 Bugbane: ee len ¿RRA AAA 528 Dandelion ct. een eon 8 Fireweed ...... 5. Mu 677 it Hest ass eU $32 Fire Pink u... rts 536 MM S TUAE LSU IE P48 Tishberry A bass MM Jibi M IUD NI GU DNE 642 lag (CC RM EU aa cue 501 Bine ee ds 507 A EISQNE DESI 596 |. ON UN 27, 495 mwell su E IN ES 628 Lo Bl mec" T 507 Hellobóre ec ee O 501 BIDS O A 507 A RE IE MU RA hys 537 [JOOSOSULEIÉG Pri ee Merle 606 Flaveria campestris ............. 675 ust RR M cq M ps 524 II DEME BennyroyaD rer 630 Fleabane Pinpernel 222 25m ened 641 bo BRAZIL A CERERI 665 Rue Anemone SUE toD 542 MUAY BATS Oye Sp A ee erem es 665 N add duc aa 675 A ILI ann Saaie Plant 4 e 573 Tier. LOSING. coe A A 507 Bolomon^se-Seal ......... se. Bs Fou He .:: rng 619 biais AS ROC RIO ae 503 Flower-of-an-hour .............. 599 WENGER A EVE xxx 661 TM Currant RA N ER 555 egi elo corona tere CO c. 5$ IN PT A 500 Kennel Seius ne 611 Foenicalam vulgare... eene 611 Dog rud ce ETE T UT 010, SO. o RA erf PRÉVUS 630 MERCUOLZTASB voee etre rs amie oper Forestiera acuminata ............ 618 E is CERES CS I FFugeébeenot 2.2... en 628 FHestucacelatioros evo o ni ore 459 Four-o’clock, Wild .............. 533 > NE $69. Nosyow, Fable error 642 nu A a O OE C 459 E S ERIBe 92.5 5-5 AO x 642 Coon A RIT EU Qe OMM a e euos RA 478 octoflora var. glauca .......... 459 A Y AA IL 466 octoflora var. tenella .......... 459 VM CPC INS re 478 BOIUTOR e ace cers oes 459 MORIUN. ierre O E 466 Bhor 243285 0022850895 459 les a EEEE 4039 ae a 478 Kugworle sense re 000 vos 639 AL A UEM. 478 Fimbristylis autumnalis ......... 484 Fragaria chiloensis var. Ananassa . 565 autumnalis of Am. auth. ....... 484 Urandiftora ¿coc VS 565 Baldwiniana ............. eese 484 IT vol o's) cisely MSS 565 oüNtanen- c ro cro veis 484 vesea var. amerieana .......... 565 eastanea var. puberula ........ 484 VIPQHHADA O vr es 565 NW EN earn ne iste ERU 484 virginiana var. illinoensis ...... 565 geminata 2... ee teres 484 Frasera caroliniensis ............ 619 [Vor. 22 MISSOURI BOTANICAL GARDEN 110 ANNALS OF THE Page Fraxinus americana ............. 617 americana var. subcoriacea ..... 617 PM T T arra etsoéevocvr 618 pennsylvanica ................ 617 pennsylvanica var. laneeolata ... 617 profunda ...ooommoooo roo...» 618 funda var. Ashei ........... 618 quadrangulata ................ 618 E AAA 577 Fringe TIER sire) rete ew En v 618 Froelichia campestris ............ 533 A I 533 gracilóa .....oooommmosor.sso.o» 533 Frog-bit, American ............. 458 Frost NER A Ern xd 664 Fuirena simplex ................ 485 FUMARIACEAE ............... 546 G Gaillardia luton „css 675 OA EEEE CER 675 Galactia regularis ..............-. 584 volubilis var. mississippiensis ... 584 Pocas arkansamus ......... 590 méissouriensis ....ooomooonoo.oo.. 590 Galinsoga ciliata ................ 674 DAFTIÉOIR- rias 674 er. var. hispida ......... 674 Glum Aparine iia sas 648 seien var. . Vaillantii ak PIS 648 BERADBANUM Siria cn 648 dr: var. RN ER eee 648 Cog) E 648 DN A. EEE 648 CONC 250er 648 Mii 648 pilosum var. puncticulosum 648 febrium ir riders id 648 THHorum 1 $4 rS 648 Wirgautum ia. hie 24244225997 647 Garden Rocket ................. 548 ie O TUNER 501 ZEN A ra ENS 501 AAA CEU IRE 501 Page GENTE Dinig ¿ricino aa. 607 EMEN AI 608 Serer ee rare re on 607 o aaa ae eer. 607 parviflora var. lachnocarpa ..... 608 Gay Pestheor ...452 oorr 657 Gentian SOC aaa 619 BON ¿EA 650 SRR oe S 619 re ee eee eee 619 Gentiana Andrewsii ............. 619 ON hs 619 BANN raras 619 o aaa 619 quinquefolia var. occidentalis 619 Baponaria ..... rn nnn 619 GENTIANACEAE ............. 618 eocarpon minimum ...........- 534 GEBANIACEAB .............. 586 Geranium earolinianum .......... 86 earolinianum var. confertiflorum 586 T MUTO n MEET RO T ET ET E LIT 586 DOSDUM ui: era scans 586 Geranium, Feather .............. 529 Vt reer et MARA 586 Gerardia aspera ......... ee 643 Mutoulalà occ cece rsen 643 O 643 A A ova 643 OOS RACER RADAR 643 randiflora var. serrata ........ 642 pedicularis var. la TE 642 MOE: 65555008505 odes Rand 643 Bkinncriana iaa kt seas 644 TOM OG A Ea 643 tenuifolia of auth. ............ 644 tenuifolia var. macrophylla 643 A ere eT errr ee Try 644 CNOA Boso oii thea ei ES 624 deum cán&dense 4.2.2292 acne 567 anadense of auth, ............ 567 canadense var. eamporum ...... 567 laeiniatum var. triehocarpum 567 IIA 567 virginianum of auth. .......... 567 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI 711 Page Page host: Flower. es opens einer 818 Grama, Blue ........ 4 e een 471 gie URGE RM T. 626 Hairy: .. cos P Eee 471 a VM I ERC UE 626 Grape am. -over-the-ground ......... 632 Bhd .... Do ale 596 Gillenia stipulata ..............-. 556 A Do de 596 Ginger, Wild ............ eee 524 Frost esee cateo BU ar 596 InBOng Do DONNE e i TIO. 609 Northern For. E aw ee 596 Gleditsia aquatica ...........-.. 574 Post: Oak 1.25 TON ar: 596 San NEN UN EI 574 BACCHON Jesuit SI a lo 596 triacanthos var. inermis ........ 574 Bed. A eS ROS a 597 Glyceria acutiflora .............. 459 Biver-bank .. 245.0 de ee. 597 BTKANBANA (sion cero 459 udo. B D 597 NEPAL rU ED E 460 nr REN IPR REE ce 596 palida. ui x exar dE BE S E 1. dA AS 596 septentrionalis ............... 459 Grape os ers USE pe ae 502 rA e ne. ea ee 460 Glyeyrrhiza lepidota ............ 579 Alkali Ee eR EOD DE 461 Gonolobus laevis ...:25..5......- 622 American ia SRR Oe s 470 naphalium obtusifolium ........ 666 Wheat rs. secede ae es 463 obtusifolium var. micradenium .. 666 A 05. ore sce ee 470 polycephalum ........ eee 666 Inv onset oos ret E 485 RO ONO 666 A A ROS lO 02 Gosti A RS 556, 680 e AA ST 478, 479 M CINE S 578 Qo RE. A E A © Golden Alexanders .............. 610 A ee. S oe 479 TU n EE cem. 543 al A TOIT Y dodo d Y S S runde ERO e 478 Se De ee 658 A A ee ei 470 A O ee 659 Nus Annual .......]. «0»... 400 Flat. LU A LS. 661 Dada oi 60 6029 RS Ses 460 e OS 659 A A Per ee re 460 Gry ais ER sls Ore ee ate 660 ERY 1... EUR xm rs 460 Ro M Joaved Al 660 MOTO Ea ern 508 Goodyera pubescens ............. 510 a AAA NO 465 EA A AR O o 648 Do C a ee runes OE 471 Gooseberry Hund Barly 2... ccm oos 464 jckly rs ron nr 555 lib td PCR MS rus coc 478 Southern: oS CARO oe 614 ODA ascii v haere ater T 468 ld nt RIO u 555 a ei o ON 471 Goosefoot «Bae. O OA ADEST 471 Maple-leaved ................. 529 Gabel o e uve Br EC 471 Oak-enved. are aaa 529 MT SERE TENOR SORORE ROO ODE 480 Gourd (UU esi sche CINE: Toe n 462 Missouri OS D ler 652 A O T 472 O NE 652 AS C DTE 472 GRAMINEAE tonos nicole 458 O 0200: WEBER NS 471 [Vor. 22 112 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page DEMNM .Leeeeoeeireoio vt 467, 468 A DD T TET TM 408 EAE WEIMAR PI DE. LOW ee san 460 Feseue, Meadow .............. 459 A vir en 462 x ME nennen 459 ee) AAA 464 BE AAA 472 o A A TEPPE 477 MODOS circo 472 A PA 466 Dom Meadow o... .socosi 460 MEN IP rear 466 c ARA 478 A AAA 469 FERMEN eee eee eee ee eee ee 466 SEN E NO 470 A RT 478 o TRO oe ere 485 A AA 466 a Ser 465 FEMME PA 478 Versu, ERS riscos... o 471 ZUM ana 478 o ee ee reer 472 i A 577 a A 547 UN. eiie] ee errr ee 470 "WU UN ee eer adeb 465 AA 466, 467 PAE AAA 471 EEE: A 464 co AA A 477 Hungarian Brome ............ 458 1 ana TTL UEPET TOT. 472 o A 480 I Bl. na 466 PIDEN AN tv eX MEER MO DENN AAA 470 MN IN 460 im FH Tee ee ee eee ee Pree 509 CI eer er eee eer ee 460 Grass of Parnassus .............. 554 AS 461 Gratiola anagall PARAGRAFO 5 x Meee Teer Ter Tr or 461 e METRE ECT. CEUTA PTET CRT 641 E TS A reer ek a 641 Ln” PA 460 BEN iden 641 Tete ene ee EN 460 sphaerocarpa ........... sss. G41 NM ood our 64x Ea ia 462 o AAA 641 nr AAA 471 virginiana of auth. ............ 641 A RI 471 Greek Valerian ................. 626 MI A ee eee 465 Green Dragon .....oooooooomoo.. 495 A ERE IET TIT re 462 "un A A 506 a BP AA 469 nc METITITIT ITE TT TETTITTTT. 506 E iussvessss»és enu AREA 469 dad. lanceolata ............. 658 Pu OUD iria 472 lanceolata f. latifolia .......... 658 AR 462 OQURTTOMA ¿ici ta a 657 DENN. PEN 462 squarrosa var. nuda ........... 657 Reed, Long-leaved ............ 465 squarrosa var. cn CREE E 657 NU éd ChO ev eoe XR 647 Gromwell DM eee eres rer eee Tees 628 MEN Lote eee re ere 464 AA TT TII T TO 628 a Sense 464 Grossularia missouriensis ........ 555 NULLE O 461 d o EPR IA 462 BENE Ee EE rir vL (éd sas bis 637 ane S TEE TE ET TS Te eT TT TT 467 pene EME AIR 576 A 4 NE E PE 632 AAA A 471 FR PS Te: 578 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI Page Groundnut as 0 0 are 58 SEA AAA o a 613 Bweet ine nen ee re 555 Gum Plant oo ters 657 Gum-elastie eu... 1-2 O 616 Gutierrezia dracunculoides ....... 658 Gymnocladus dioica ............. 573 Gymnopogon ambiguus .......... 471 H Habenaria ciliaris’ orcos 509 clavellata a iaa 509 AVB y ar VITGSCODAS er se cn ae ee 509 MYCOJ EM A T 509 leucophaea 7... au. $e Deeds 509 Dersmoensa an viele cies: le 509 Hackberry Pates nes 522 Wario e caen UN 523 Southern). a ee leere 522 Hair Grasse... d eec rer ae 466, 467 Tonic acia IS alo 608 HAMAMELIDACEAE .......... 555 Hamamelis vernalis ............. 555 vernabls fe CHINES 5.2.7. 555 vernalis f. tomentella os 555 Irginiuna oo esae Wee Qr 555 Harbinger ot Dacia Sater 609 Hardlisik iseset veneesi eraann ns 556 iHarebellos cov NE OU saree S 653 Hartmannia speciosa ............ 607 Black: 7 pU 651 o eA UNE A CCP UD 560 Parado an E A cele cs 564 Possum 2344 92819 ala aha ee 591 mmer 5.20 AN Eo VUE 563 Hank a Beard aa near 681 TETSU W EW OCC cis: etinm ss 682 Harout TAO 516 Bosko: sonata asses vee ae 516 ¡Henao cree 632 Hedeoma acinioides ............. 634 hispida. eere eu EC tetas 634 Dulegioidos c LU Oe T 634 Hedge: Nettle a see 632 Hedysarum frutescens Helenium altissimum polyphyllum tenuifolium e. o.oo.o..s....o.o..o EEEE ILERI EE, "t! t] |] EI) ecc tr t |] |o) ..oooooo....o.......s. tracheliifolius tuberosus tuberosus var. subcanescens .. ilis ee +... .... Bo9 9 9 9 9 9 9 9 9] 3 £9 9 9 9 9 9 ot t t n t1 n9 114 Page Heliotropium indieum ........... 627 o (iussi aaa C4 ENS 627 Helebore, DEE | ¿rr 501 Helleborine ..............ioo... 509 Hemerocallis fulva .............. 502 Hemiearpha mierantha .......... 485 micrantha var, aristulata ...... 485 Memon Chervil 1.2... iae 612 SEMI yaa TUVO EO EX AE US 612 NEN ON 523 ns aa LU TCT Te 620 a $233 532, 533 Sin ied Climbing ........... 656 ea 632 <1 a. aan einer 541 RR ras eirans 541 acutiloba f. albiflora .......... 541 acutiloba f. rosea ............. 541 a A 405d A3 TRIER 541 americana f. candida .......... 541 BS ETETE EEIT PELIS 541 triloba of Am. auth. ........... 541 Heraeleum lanatum ............. 612 Hercules?” Club 5... s 608 Heteranthera dubia ............. 498 es rare eer ee 497 BONITO ae oii chats os tae 498 Heterotheca subaxillaris ......... 658 Heuchera americana ............ 553 DIMUDICAMÍÓS 4: ae 554 hispi RR 048 554 A een 554 parviflora of auth. ............. 554 DUDITRA. sonen naco XA 554 een var. Grayana ..... 554 a AS T dE 554 Hoxalstrs aphylla ............. 511 aaa ses ewe 511 Hibiscus DIEM 4 455 bo oa er 599 MEOURIDUS +. diras 599 1 ha) EEEE EEEE 599 TAO ee 599 a AA eter pns 599 Hiekor Big Bhell-bark ............... 514 IA rr aaa 514 BME: G4 cs bes CEVREXERHE Cx 514 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page E O erac deem 514 er > LEBEN a daa 514 GU SNP La C23 17 2) 514 White NT ERE EE 515 Hicoria cordiformis ............. 514 IA RA 514 a AS 514 A iy en's ee 514 coo NT 514 DEA A A 514 Hieracium Greenü .............. 682 A TITO TIT 682 ronovii var. foliosum ......... 682 o AE 682 BATON ¿ii 682 is ETET ETTE TEET 682 eae PEE di EUR 682 HIPPOCASTANACEAE AR 594 MANET ALIGNS WE 547 Hoftmanseggia JAM 6545050008 574 A APRA 584 MOONS METATI O 588 Zoe lanati ¿cc ceia 465 o IN O T 598 MONS ae cause sawn 610 Honeysuckle DA «cri aa 650 A A écran 576 MADAREMÓ ¿o carrer candace 650 qua A anne 650 | Seer O AN 613 (fun. AA TOY TEST 650 ser 523 SENE A TT 523 T e. E PI 516 A Y Y 587 Hordeum JUNGEN rad 464 A TP 464 BEEN a 464 BRORUnI 2:0, ee 480 vulgare var. trifureatum ....... 464 A RP Tr eTererr ee 631 DMARD cursa de S aa Os banal 635 o 5.744 eG.anen ETA an SEE 538 Noe Bolm 22. eagssassi eas 636 CODD en 650 ER TTE 634 1935] PALMER & STEYERMARK Page Nettle soets enoar US 636 White Horse Nettle ........... 636 EOP SO WOU, near ito wes eras 668 Lforseradish oo 000. ee ee ore 550 sackia americana ......... ese 576 Hottonia mntiata se. 2. een 615 "HN TORDO ES ee 627 Bouton pata DNA Ad 649 er T TIS 649 Sols TOU NR ET dE IUS 649 Ianceolata ee een 649 Iongitola Gs. oi 649 OO a S 649 A O O GO Goa 649 purpurea 1... ao 649 A o AA OE 649 Huckleberry Lu Usher T TCU 614 «Mowsbush ee ot Nee cre cre 614 Humulus japonicus .............. 523 US elects eerie oe abe 523 Hungarian Brome Grass ......... 458 Hyacinth TADO da LIE ee 502 AA ETE OOO OO 497 ida nr Tete s 502 H anthus concolor ............. 601 Hydrangea arborescens .......... 554 arborescens var. Deamii ........ 55 arborescens var. Deamii f. acarpa E. arborescens f. grandiflora ...... arboreseens var. oblonga ....... arborescens var. oblonga f. sterilis 554 RS MO 55 Hydrangea, Wilde Nw as 554 Hydrastis canadensis ............ 543 HARBRITACEAHB........ 457 dehesa AMAIA a E T 627 SN i RE RSI CORE 627 baby SEN CoE "oe 626 Hydrophyllum appendiculatum ... 626 Canadense ....secsecsesesesess 626 virginianum e UE 626 Hymenoeallis oecidentalis ........ 506 Den Scc carolinensis ...... 675 OFVINDOSUS. e e Yes eee net 675 pe ii NI IDA 675 PLANTS OF MISSOURI 715 Page HYPEBICACEBAM '... $1... eese 599 Hyperieum Ascyron ............. 599 eistifolium, 7:22. SS eese 600 ensiflorum- un... due: eis 600 Drummondü in. ern 600 gentianoides ....... «eee ne 600 mnanthum-.... ies 600 loboearpum seansas seras 600 mutilum -.::.. 5... SR. 600 perforatum. 2 0.0 Ee mes 599 petiolatum ............ eee 600 CUM LL. o ces 600 pseudomaeulatum ............. 599 punctatum. ......... 2... 0e e$ ee 599 vVirgihloum ¿dede Uem vate = © 600 VUDIA hirsuta ...... ES 507 hirsuta var. leptocarpa ........ 507 pisi. Clammy Hedge .......... 641 NEN E Se A ore vee e 631 o AS E ER T 641 Hor Hystrid. ...... RE T 464 A es vereri. ee 464 I Moa y AAA p ow tee hee 591 O PME s OL CAE RO ORI 592 dois. var. padifolia ...... 592 ILLECEBRACEAE ............ 534 EU anagallidea ........... 641 SPORIS ES NC c o 641 foeda BOSE SNP. PEE EUR 641 Impatiens Balsamina ............ 595 DIA ias. o UR 595 HI o ee 94 Indian Blanket ....... Ho... 644, 675 Bronüd-foo0t ca ee 576 nid crc Dee none 595 a a 480 on AAA LOOG 11. ieee ey UR o 503 CHEND: scene see 650 ee TER 480 Eien. DISC i ht de 627 DI S ata cci IR 620 DOON PASS) ¿os B ES 461 MOA olco epe 548 PUE cnin A 536 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Jerusalem Artichoke ............ 672 TURION Weed 0 ee 638 Johnny Jump-up ................ 603 LS e ers 480 a o (coser an a e YA 528 PAT O ta TT. 533 JUGLANDACEAE ............. 514 J po QGIÉ6fOR au ee 514 ——— PICS 514 J Ion cid BER A PENE EEEE ET 498 Juncus acuminatus .............. 499 alpinus var. fuscescens ........ aristulatus ........... ees 499, 500 baltieus var, littoralis .......... 498 O qud aan F 500 biflorus f. adinus ............. 500 brachycarpus ............. s 499 n loni MERO 498 O an Ra 499 nadensis var. subcaudatus .... 499 Oioholomus ses tots 498 A ee 499 DAA EE s uf E 498 effusus var. solutus ............ 498 ar vor) A E T 498 c —X vU Tree eis rigare 498 errr eee reer ere 498 macer var. anthelatus .......... 498 A CITTA 499 E Mus var. setosus ........ 500 "Il 10 SUCRE ONE RETI 498 BODRUM A a RO 499 S DL UT EFLENT iras LIT 499 polysenhaluB os os iss o o 499 dan MA ne 499 o ii 499 cd daca ar id 499 DA eae OS eee 498 eo PAM 499 TERPEN 498 tenis of auth. oss. ie ces sees 498 tenuis var. anthelatus .......... 498 TOMAYl 2. are 499 TM ET EEE ET 499 WEIT a, 557 e. AI IT TZLIEEIL TY 454 s ti.Dl 0 O EE 454 1935] PALMER € STEYERMARK—PLANTS OF MISSOURI Page ROE A A 454 ya A O 454 virginiana var. crebra ......... 454 Jussiaea decurrens .............o. 605 CIM ccu Mero C O 605 K Kallstroemia intermedia ......... 587 MORE cies RO ele; aie DOS 587 Kenilworth Ivy ................. 639 entucky B (erri ME ma e 460 Coffae-tree rU edens uM cars 573 Kiekxia Elatine ......... 9. 639 idee Ur UU 514 Kinnikinnikrs ese eri 613 Kneifia linearis .......... ees 607 pratensis ae eI rei 607 Knot-grass ........ eee eren 526 Ienot-weed ee een 526 Japanese ........ e ertt 528 Kochia prostrata olaa eve 529 Aria 5 TR eis n teris 529 trichophy lla nee ee 528 Koeleria cristata ................ 464 Krigia amplexicaulis ............ 680 HOLAS a vene 680 Dandehon arte 680 occidentalis u erster 05503 680 Virgimies Decisis 20e sie Slate ee 680 Kuhnia eupatorioides ............ 656 eupatorioides var. angustifolia .. 656 eupatorioides var. corymbulosa 656 Kyllingia punilg 2099.93 8 L LABIATAE $21 2 554] 630 Lacinaria lee ER 657 Laetuea ca of auth. in part 681 E potea Are 081 . integrifolia f. an- 6 He deae rere 81 eanadensis var. latifolia ....... 81 canadensis var. latifolia f. exau- Jeulata a na eR eru 681 canadensis var. montana ....... 681 canadensis var. obovata ........ 681 7117 age canadensis var. obovata f. steno- poda ........ eee ento ern 681 canadensis var. typiea ......... 681 idana 2002 Eee 682 graminifolia .........o.o....... 682 IBrBULH- 05 ee: 681 integrifolia of auth. A ones 681 ludoviciana 1.5.26 e Ee canes 681 pulchella .......ooooooooooo”. 682 sagittifolia 5s. al aja s sures 681 E A O ege eese © 681 Beariola "u. 0000 o id ss 681 scariola var. integrata ......... 681 1 tes. tS enis 682 Ladies? Ear-drops .............. 528 Aut eA TOU ERI AQUIS. 1o 665 ME 2 NIME LC 509 A A cmi quee 509 NET A uc 510 IV EEEE ROS em coe 509 Lady *s Slipper AAC AA. AAA 508 A A A nus 508 ISA A erm ere 508 SU PA AE 508 SA E O eae 585 lo AA d... tmn 527 Lagenaria me — t Don 653 Jumbi Quarter... edet ers 529 Lamium anie M erstes 632 DUPGUm. 2.2.5.4 A cows 632 Laportea canadensis ............. 524 Lappula echinata ............... 628 Redowskii var. occidentalis ..... 627 VIRADA nenn ernennen une 627 TGP AUIS ae 543 Dd AA. A ey rt» 543 Glade O Ex Wes 543 a R TEE EL AT 543 an palustris scan 583 pa alu ustris var. ya M er 583 AA CES D 583 venosus var. intonsus .......... 583 LAUBACHEAB 5. AE D Ee. 545 Lavoaucta iriloda .....oooo«.o.o... 607 LendrBlant Pi een 579 Bash A te 666 718 Page AA APA 542 i | AA T. 542 Leatherwood sarah 604 Leavenworthia uniflora .......... 551 hea ME erara era 601 DOGO eu 601 ER 6a 444) ca VERTRETEN 600 uU WEE ias 501 Leersia lenticularis .............. 471 AAA i i rrr 471 vi NÉ AAA SERIEN 472 LEGUMINOSAE ............... 573 Leitneria floridana .............. 513 ERIACEAE ............ 513 na cycl Se ee re oe 495 Bee ET TCT ER TTC POET TT TTT 495 a UATE E TOOT ET TTS 495 perpusilla var. trinervis ........ 495 _ APP 0000s 495 VAIVANA AAA 495 DLDERNACBAB coo seitas 495 Lemo = er 634 Lemotris hyacinthina ............ 502 periods atajo ES 644 Leontodon erythrospermum ...... 680 Tarazaown ......... een 680 Leonurus Cardiaca .............. 632 Lepachys columnaris ............ 670 columnaris var. pulcherrima .... 670 IAN Losada xa FALE 670 Lepidium apetalum ............. 548 A ae wank 548 Ve 548 > AAA E TRU E 548 teglectum ..... ecce eene 548 pov errada hen 548 ML E 548 A O T 548 tandra virginica ............. 641 — attenuata AT E 470 FUNEM A 470 era Serer eee 470 e ELE ES 470 Leptoloma cognatum ............ 472 Lespedeza aeutiearpa ............ 582 A AA 582 capitata var. longifolia ........ 582 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page capitata var. sericea .......... 582 a A PP 582 a E EAE TTAN 582 ei mL AiE eina n OTT 582 RA 581 AAA 582 procumbens .................. 581 a VU a as ers 581 BEE eee ON 582 s A 582 NEN. EEL TE ON 582 DA adi 581 Stuevei var. angustifolia ....... 581 Stuvei var. neglecta ........... 581 WEN A ach 581 violacea. var. prairea ........... 581 NEM ee 582 Lesquerella angustifolia ......... 547 A A D T YETRPITT 547 Lettuce, Priekly Wild ............ 681 Leucospora multifida ............ 641 Levisticum officinale ............. 612 T adio PODES Rr TEN 657 EUA cac eT 657 eylindracea E EII RESET 657 AUTRES E AO ETE. 657 wis. LELLE E EEEE TT 656 punctata of auth. ............. 657 pyenostaehya .......ooooooo... 657 BEAR cosoeceallisasauweseérexa 657 e a OP CORT TEE 657 on nase 657 MENTRE. ann 656 squarrosa var. intermedia ...... 657 Licorice, Wild .................. 648 Ligustieum canadense ........... 611 ud Kar 7% ¿ocio ricos 500 Lilium michiganense ............ 502 a O a a 502 em PETELE RN 502 Bato ch Oe ED iE one wa 507 EEROR TITRE tees VPE AP 502 x AA da 607 xo. MM TIT TY TERN 502 TUTTI RA 502 1935] PALMER & STEYERMARK—PLANTS OF MISSOURI 119 Page Page Limnobium Spongia ............ 458 Locust Limosella aquatica of auth. ...... 642 Black... ar ie 578 aquatica var. ara of auth. 642 Honey eee. ek ee 574 tata ... A eee 649 Cle AERE OUT RU EVI 574 tenufolia-. aii en 642 LOGANIACEAE .....1 sses 618 BEINACHAD See 586 Lolium multiflorum ............. 464 Linaria canadensis var. texana 639 perenne: A. ue yrs 464 br Non eu 639 temulentum .:... 3-95 0. 464 HiGtue Gare ne E sie cesis 639 Lomatiwm daueifolium .......... 611 spuria .....oooooommoooooooo.o»..o 639 Lonicera dioica var. glaucescens ... 650 vulgaris 2.6... eee eee reece eee 639 aT, ree i e Re 650 AA A ein tras 597 HavidA ¿canas EDU S 650 e Ter HE TER 597 glauoesoons Lecce pent 650 DS C NEM M PE 586 japonica ....ooocmcococcoccros.. 650 Linum medium ...............s 586 prolifera PT O ee 650 er A IO AE 586 BIRDErVITENE ..... $9.91 59 ev. 650 VD 586 EU V << ns AO ce 650 A xu d 586 Loosestrife ih za cese duco 586 cn 606 Liparis liliifolia ..............0:4 510 Purple .....-seeeeseeeeeeeees 605 Lippia lanceolata .........0..... ed UN he rari S leche ui ds M 03 630 A E A reru A bo es 616 : g : MOGINOT He. ra ee Se 630 “hy edie ee ee ee m Liquidambar Styraciflua ......... 555 Bf m calycinls ......... 457 «Exquorice, Wild Re. oe en 570: POE re so cle nr ie 646 Liriodendron Tulipifera ......... 544 LORANTHACEAE saene uasa 524 Lithospermum angustifolium ..... 628 Lotus americanus ............... 576 AYVONEO: oe seine ass om 628 pornieulatus v... os. deren A 576 e CILE E aia MEE 628 A er re 539 CATOUNIENBO acesso opcs 628 MQOW 1. voe UR ee 538 Mr O E E E E One: Winusawort vende adta esi 644 a 1129303 652 e 028 Lovage ......... eadh n n n 612 Füve-loroevor oe ya 0s ke ee 553 Love Gras ......... 4o eer e 460 Liver-leaf ............ e eee 541 Creeping .......oooooommoooo... 460 Lizard s-tail cn ao 29. 0 oH 511 Ddian. rM Uer 461 L ACHABIU TITLE 604 AN Ue ce 461 Lobelia cardinalis ............. 4 GBR Love-wie IA O O 624 iniata sees A E 654 Ludvigia alternifolia ............ 606 dT —— E 654 glan cum 606 LAGER ex ee 653 atans var. typica ............. 606 "hii RER EOS: 653 palustris of auth, 2... ees 606 U c O OS 654 palustris var. amerieana ....... 606 ía var hiriella-— o. sm 654 sid. iar MES SCC le « 606 Lobola Buena ee 653 Luzula pt var. bulbosa ... 500 LO A ER 668 avons AA ech es 536 a .....+.+. +... ee 120 Page Lyeium halimifolium ............ 638 Lyeopersieum esculentum ........ 63 gd Di BTE TETTE 635 UNLOAD E Qux 635 lueldus var. americanus ........ 635 FEM aos ae 635 TONOS srianan uide 635 Lygodesmia juncea .............. 682 Lysimaehia Nummularia ......... 615 o BERTIE UTENT TET TEE TT 615 BYIBEBAUBABE 2a nasa eua 604 Lythrum alatum ................ 605 PETER 605 M Maclura pomifera .............. 523 Macuillamia rotundifolia ........ 641 Mad-dog Skulleap ............... 630 Maalat Wa VELTTETTTCTTTETTITU 648 Magnolia acuminata ............ 544 acuminata var. ozarkensis ...... 544 MAGNOLIACEAE ............. 544 LEGE A 536 Malaxis unifolia ................ 510 Clustered Poppy .............. 598 bo AREA TT T EN 598 AAA 598 Fiagéd Poppy irtiri ent 599 EMEN O a 598 n lA QVI ele te 598 O E TE 599 ecl PORD MPP eee eee 598 TUDO TOMA soii 598 Malus bracteata ................ 557 a AAA 557 O en TT 556 ioensis var. Bushii ............ 556 ioensis var. Palmeri ........... 556 ioensis var. spinosa ........... 556 Qus A A le 556 ne IT. 556 RR TT 556 22. EEE 557 Malva erispa ................ 598 RA 598 eh PE TT T T TT TT T TET 598 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Fotundifoll ¿Es O 598 EDS ER bor Vv ya 598 MAETACBAD sr cess 598 Malvastrum angustum ........... 598 T missouriensis var. caes- a: ran ere 604 Man-of- tho- GATUD- brevi Env pU 623 MEUM ARI 544 Manisuris eylindrica ............ 480 anna Grass RR a riit 460 PEE AP ER o akenes 460 Maple oO A ee er re ere er 593 RETTET DOC PTT ITI 593 AA ee t 593 BE $05 x5500 RS 593 2 IMA AA 592, 593 MARANTACEAE .............. 50 Marigold FE PRA 675 ee ce eee ee 674 Marrubium -— io 631 Marshallia caespitosa ........... 674 obovata var. Ims FÉ 674 Martynia louisiana .............. 646 MARTYNIACEAE ............. 646 Matricaria Chamomilla .......... 676 matriearioldes ................ 676 A A 676 "cien TH reso veo videxava 638 A 544 mmis A E Fes RN 604 Meadow Beauty ................ 605 a A PNE 541 any rra 541 Meadow-Sweet ................. 556 Meeardonia acuminata .......... 641 Medeola virginiana ............. 503 Medicago hispida ............... 576 A ci 576 a MENEER TET tT DETTO. 576 NEM DEN A ceondzs kv a o 576 Megapterium missouriense N 607 grandiflora ........... 579 Melanthium virginieum .......... 500 MELASTOMACEAE ........... 605 1935] Page Mele (TABS o... ssi sos o UP an 462 Melica mutica .......- ecce ers 46 iir mee pr NE 462 Porteri 25022900 VEO EO 462 Melilotus alba... 2.2.8 576 Officina s oe evi» corr eife eia 576 1838 OfHelnALIB u: ea. 634 Melothria pendula ss: coc. geese 653 MENISPERMACEAE .......... 544 Menispermum canadense ......... 544 Mentha alopecuroides EA. 635 BIVOUHIS A NA SL 636 arvensis var. canadensis ........ 636 arvensis var. glabrata .......... 636 GS PETIT ori edo ee 635 CILFALRA nos 90529019951 636 eh et OH 636 longifola haa nee 635 piperita a e o 636 ICAÍA: ve Er rung 636 sSyloesirta Jor vd es os de i 635 Mentzelia oligosperma ........... 604 Mercurialis annua .....oooooo...3 588 OFeUry 575559 ee 588 Three-seeded $$ ccr ved 588 Merioliz serrulata ......... eei 607 Mertensia virginiea ............. 628 Mesquite Grass ......... esed 471 OSqUii6, VING e resisti 477 MOXJORD ROA 9€ bes cos S 529 Miami Mist 2c swe ss ect see 626 22 pennsylvanica ........ 553 OS 553 tirgintondia A A 553 ERE netiis A tas slave 510 Mikania scandens ............... 656 Milkweed Chmbing ao cra e coto e 623 Green sa ee 622 Purple eis erresis tos oo 621 WAND tiara Dr 621 Mhorled (4.5 cui ds ets cio ee 622 Milkwort Wield o REO UE 58 Pink «uy eed re nern 587 Millet Broom-eom 5 uod me cns y 477 721 Page O TO Ue es 472 Hungarian a 478 Hallan A ass se 478 Mimulus alatus ......3.. 5. ee 640 glabratus var. Fremontii ....... 640 Iingons ¿ao s 640 int Bergamot A TA s es 636 Om... OO Hk. 636 Field '....-5—12. e DURS 636 HOrSOó...--9 REN 634 Ohio 1... NOSE S 634 Mountain eesse souk 049 Wer es 634 LO0Q A BI ES chs 634 ds UI TIG 635 Minotti Gouri s$. iisas 652 a A od 656 A A EROS DECIDES 524 Mit behala A OA woes 648 Mitella ua TOP DTI en 554 A A let 554 Mock ee NEWbed Shed nek ee ee 610 E A cd Te 515 Mollugo verticillata ............. 534 Monarda Bradburiana ........... 634 A ene 634 o X MENO MEME. eves 633 GUDOPEd osa ESS deas s. 634 HNDUIONG + aaa di c 633 fistulosa var. mollis ............ 633 AMT SERERE ERE INI DUE 633 D DERE. RqUIAN ORC OT 633 . DUIS CAU v EAEE S ener 634 pun sie var. occidentalis ..... 634 punctata subsp. occidentalis .... 634 punetata var. villieaulis ........ 634 Moneywort: spares saage eer ote 615 Monkey Flower ................. 640 Monolepis Nuttalliana ........... 531 Monotro - Hypopitys ........... 613 Unior e. Be la voor CMS. 613 an Pe eee d op d^ 544 BRACEAD 012528 a rotor... 523 Morning Glory AE Y AN 623 Commun ES ARI ner 623 VOU DONAR A EE 623 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page o a ere Se eee ee oe 54 m ER CES CET TET TET T TIT 549 VEN. AA Ah 548 d ni EET T T TET T EET TTITET 549 Myosotis seirpoides ............. 628 o MM 628 virginica var. macrosperma ..... 628 Myosurus minimus .............. 541 Myriophyllum heterophyllum ..... 608 proserpinaeoides .............. 608 MENU cci opea rris 608 N TOP 456 NAJADAOEAE TESTER 455 Najas guadalupensis ............ 456 o bL EET TETTTETTET TIT 652 Narcissus poetieus .............. 506 PINO PP 506 ss AAA 643 Needle-and-thread ............... 502 Nelumbo lutea .................. 538 sonata E TOT TE TESTE T 539 m a nn 538 Nemastylis acuta ............... 507 Nemophila microcalyx ........... 626 Neobeckia aquatica ............. 550 Nepsta Ostaris .......0...0.0.0... 632 NOON S TETTE TET 632 hederacea var. parviflora ....... 632 c NDS CIT TERRE 636 BE A AR 632 BER ern 524 PARA 524 AO 632 A E 636 nn EI 524 BER EON ana en 636 PEPER CSTE (20 TET TQ. 524 Nettle. Clo O 414 Now Jersey Ton PA 595 Nieandra Physalodes ............ 638 Nicotiana longifolia ............. 638 SEER TS eee TTE TET 638 1935] Page Nightshade IYO) NO EDD DEUS 636 AO ers ee wars 636 Nimble“ Wil 2255.5 22.00.0082 ss. « 46 ININGDATE A DEUS IE US 556 Nipple Cactus ..52-24 vna 604 Nodding Wild Onion ............ 501 Notholcus lanatus ......... ese. 465 Nothoseordum bivalve ........... 501 Nac MUN CHUTE IRE 486 EEE TORE 481 ke eee P 533 Nymphaea advena ....ooooooo.o... 538 E coo O OECI EIE 538 OKOT PONA ZT E 538 tuberosa oo A IE 538 NYMPHABAORAR ME annor 538 Nymphoides peltatum ........... 619 Nymphozanthus advena .......... 538 OSATKADUS nr o 538 Nyssa aquatica +... cis. oe ves 613 TAE no ER 613 (0) Oak IBASKOL ee Abou A 518 ME ch betale en cme oe 520 BICE Tack: Ba 519 MEL odes caste ini TE. 518 Ohinquapin4 2390659 995 902. 518 AO OSO OD 518 O E 519 Jerusalem- a eje os 529 MNSICUO 09920202055 sla 517 ihn aR OCH eei 519 VATERS ee ee er N 517 Beda uoo voie ee a 519 A oc O ee 519 BoruD hebt C Dc MT 518 eater opio ed 518 Shingle ea a de isa 520 ipo Beda ocu 518 EC EON LR AE 518 ii White ore er 518 Turkey ee e ee 519 O OD Ra USO: 519 123 Page White esas 00002 DI 0 e? 517 ‚Willow: euer eae sis eee ats 19 ellow: 0.0... E NOT 518, 519 ON gessilifolià am e 501 o REED RE 465 pun PERDU e CEP 465, 501 Yellow False ..... ev. 465 Obedient Plant nenn 632 Oenothera biennis .... 4 75 C9 . 607 ETULICOSA iu ee lo 607 fruticosa var. hirsuta .......... 607 ERN Dr ARX DD 607 JACIDIALA «dues voa EE S 606 acininta var. grandiflora ...... 606 MO OS AD oe 607 Hnifola M A T2 11 5C 606 OL S Se rer Go vies en 607 VIUA. eee rrr TETT: OR 606 muricata var. canescens ........ 606 A SM Sacre re rores 606 pra i Mes s D ARP REM MT 607 Ada A A os 606 ogg A A O 607 O NO ON EGE 607 A eT det var e 607 A A A AO orem in 607 ld a BOSrd ic o setrs «6 618 Oldenlandia uniflora ............ 650 AI: AAA PP E 617 ONAGRACEAE AN DU TIE 605 oda MM POR PEE 501 O T ET 501 olo rdum Acanthium .......... 679 Onosmodium epica 239911 628 hispidissimum var. maero- spermum 35 2.5 Sa era Eo 628 mecidentale Mi. a ES res 628 occidentale var. sylvestre ....... 628 aubsetosum 5.5... see. e 628 Opuntia Pl rA Fs a te 604 DALMAU Sa er 604 ehard GTA. O one vers 462 ORCELDA OMAR o Ec S rece 508 as WN OOM O AO T 509 A A AP 509 124 Pa Purple Fringed IAN 509 MTOR TET TTY DIR 120 2 509 o AR O 508 inti flowered Prairie ........ 509 a CA IT 509 E ins. EMRETITITETTTITOT 509 Orchis spectabilis ............... 508 Ornithogalum nutans ............ 502 Umselgtum uoces ctio à ees 502 OROBANCHACEAE ............ 645 Orobanche uniflora .............. 645 i Pe eeRe eT Te eee Tee T 553 A ETIRTTETUTTT S 480 Oryzopsis racemosa ............. 469 Osage Orange .................. 523 Osmorhiza Claytoni ............. 610 o AA 610 longistylis var. villicaulis ...... 610 Ostrys virginians ......... s 516 virginiana var, glandulosa ...... 516 Othake callosum ............ Ls. 675 Oto + BUrIODIBES i315 e 643 a A IMA 585 pm A td 585 a ST TE 585 corniculata of Am. auth. ....... 585 corniculata var. viscidula ...... 586 sis a dis I T TO 585 a ae (uda 585 europaea var. Bushii ........... 585 europaea var. Bushii f. subgla- | LTTE TL TRE E TT 585 europaea f, cymosa ............ 585 europaea f. pilosella ........... 585 europaea f, villicaulis .......... 585 ni io a a T 585 florida var. strigosifolia ....... 585 DUMP eiii oa E ERO EU 585 pilosa var. Wrightii ........... 586 ick’ E E IA 585 ik, E CETTE TT 585 stricta var. piletocarpa ........ 585 cam PP 585 A A ERAI es 669 BUM RER 669 Ces DINE ¿ri 676 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page a. an Veiis 534 Cork ERR REV UN 534 void EISEN 533 hirsutus var. integrifolius ...... 533 EEE Keen da 533 BUB E A 534 A san 533 A E S 533 Oxypolis rigidior ............... 612 rigidior var. ambigua .......... 612 Oxytropis Lamberti ............. 578 SEMEN AO 578 P ess D PEE EEEE 644 EEE PTET T 590 Ar WEA RITE TTEETOS 675 Panax quinquefolium ............ 609 Panicularia nervata ............. 460 Panieum agrostoides ............ 477 A ITT COLT eee ere 477 er T TIS TET T ETT 474 BE ARPA A 476 o N ee i 474 MORAO O VER UN 474 C. NA eee or re rrr 476 Boscii var. molle .............. 476 A CTE Tore Te 474 AA 477 eapillare var. occidentalis ...... 477 clandestinum ................. 476 sommutatum. ea 476 commutatum var. Ashei ........ 476 depauperatum ................ 473 dichotomiflorum ............... 476 MIGNOTOMUM. sonen 474 elongatum 0... cccccccnsccsves 477 BENE LL eee cade erased Nba 476 AAA 476 NEP A RN 475 APP 477 o ann 474 nn var. silvicola ........ 474 Implicatum ...... eere 75 lanuginosum var. faseieulatum .. 474 lanuginosum var. implieatum ... 475 1935] Page lanuginosum var. Lindheimeri .. 474 lanuginosum var. septentrionale . 474 Igtifoliun |. sext eva 476 dIeibergi 5 2 nA REOS 476 Tindheimert Sib teste Ee epe s 474 Lindhei var. fasciculatum .. 474 PRAE Nd var. implicatum ... 475 Lindheimeri var. septentrionale . 474 Imearıtohum meos 478 linearifolium var. Werneri ..... 473 Malacophylam A ee 475 DINELOCALPON 55:593 99:999 92 055 474 nmüliacenm a ee 477 A IO s 474 TED t a TS 477 oligosanthes var. Helleri ....... 475 oligosanthes var. Seribnerianum . 475 Dperlonpum. N... oir. 980205228 473 philadelphieum ............... 476 Olyanthes ao IE 475 orterianum ¿e 476 DTAOCDCIUS Se ee 475 DUDAS ORY nennen 476 Havenell 2.29209 906 eee 475 BEOPSLIUM cue erste iaa 476 Soribnerianum ...icommmo.o.om... 475 SDNZPLOCRTTON 5. inde sero des 475 sphaerocarpon var. inflatum .... 475 pretum 5525292023292. 98 c 1970 474 AIDA CUM er sions coors a 477 BUD villosumsc ce ee 475 TONNESSCONEO .......-.. v enn 474 VallOSISSIMUM: vae eese ro ee 475 vVirgatum sie IS NEN 477 ER T E N RITE 473 lapense 204 900009 0999 9 474 Papaver Sub: Serv eue E eris 546 Wes NEN Pr nC DN 546 AES MN CIE T 546 ee USD RS 545 RDEV Wa AS CRUS Uere dst s he 544 Panetaria obtusa 22... 524 Sy IVIDICA nee 524 Parnassia grandifolia ........... 554 Parosela Dalea ....... ecce 577 NEGNGNG E ee 577 leporina var. alba ............. 577 125 Page Parsley Hemlock: „a Nd e 612 Prairie 0000. 09 2 EAS 611 Paripe seo us 611 on AME rper MOD D 612 Ws sca CEP. o AP 610 n edd hispidum Ein. vies 667 Hysterophorus .... Riva cade. 667 integrifolium +... 24 s 667 TEDENE Sesaro oa BEER S 667 Parthenoeissus quinquefolia ...... 595 quinquefolia var. hirsuta ....... 595 Rm t. var. Saint-Paulii .. 596 PLUTO aoe 596 Partridge Berry |... sr 648 Paspslum.Bushié -... ea oc os 473 PETT A O & 472 ciliatifolium var. Muhlenbergii .. 473 ciliatifolium var. stramineum ... 473 Vals T PEN EMEN Ame ES 473 neis o Antio. HEMOS aD MR 472 Horis A ce es 473 laeve var. cireulare ............ 473 laeve var. pilosum ............. 473 PRUE a A deese 473 UONI eer TR ss 472 MUNENDELIW << eer ee 473 Mr Ey on 5 52 E 0 473 DIGSIOS UM Wy ere rrr vo cette os 473 A A der oid ye ai 478 AOS var. glabrüm serios 473 AO O « 472 Pos: d dii Bass ea A e RR 473 prona ancafnata ¿Da 604 O A A E 604 pa abbas qe ern cit 604 PAOLO WEN. a ass ees 604 Pastinaea sa in COS AP Pe 611 Paulownia io a D M T E 640 Pea Ser T URP ar 584 e 584 Hoa; PES OC A ESEL RE ect 578 NEIN A. A SE. d 583 jui PR. ct Mors EE ERE 584 Durtridge: She omes crees 574 a A. A CA ree 577 126 Pa A A ETT IT 575 | | ... «eo»... r. «soso. 584 Peanut, Hog .........o..«...... 584 O raros asa ad 55 EIU d. DV une 534 early Everlasting .............. 666 Mesias EN RAT COS 514 ico is a as ieee re 514 PEDALIAOEAB ........oo..... 645 pase canadensis ........... 644 AOR a eee ee ea 644 eres rest. ER CU ECL ER Te 524 Peltandra virginica ............. 495 virginica f. hastifolia .......... 495 Pennyroyal ............o........ 634 MN a Quad RATE CERIAA V LEY YT 630 Penthorum sedoides ............. 552 Pentstemon arkansanus .......... 640 arkansanus var, pubeseens ...... 640 GALyGONUSB esse eorr oon 640 canescens of auth. ............. 639 Cobaea var. purpureus ......... 640 A 640 Srandifloru ¿ico a 640 laevigatus var. Digitalis ....... 640 pallidus ........ eer nnn 639 t ^ MRSTEEERERSTEDEDOIU DS. 640 Peplis diandra .....e eorr 002 604 PebperdO0b viren tikina 551 o ml. sun 596 s.l AM 548 PODDOIMING. erar VY 636 Peramium pubescens ............ 510 Perilla frutescens 2. 4n 636 PONIA xro wx oe or RE 621 Persicaria amphibia ............. 527 ISR ¿a ici aan na 527 Muhlenbergü ..........ooooo.. 527 OD OUPDBR (ior rrr RA 528 o M PECES 527 o A 616 Petalostemum candidum ......... 578 ds AR ee 577 EDITO MMC 577 purpureum var. pubescens ...... 577 A A S 646 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Page Phaeelia bipinnatifida ........... 626 A RT: 627 A eive s 627 A tede T 626 Phalaris arundinaeea ............ 471 canariensis epee NI AY a 471 AI EA ee 471 Phaseolus polystachyus NA 584 Philotria canadensis ............. 457 Phleum Meine Is 466 BhlX atplifollá £252. viser vs 625 ca a ae 625 A ery ee re rer gree 625 divaricata var. Laphamii ....... 625 DEUBIOBUIM na 626 glaberrima var, interior ........ 25 glaberrima var. melampyrifolia . 625 culata v lL) A 625 A AA 625 pilosa var. fulgida ............ 625 pilosa var. fulgida f. albiflora ... 625 pilosa var. ozarkana ........... 625 pilosa var. virens .............. 625 A PORRAS 625 Phoradendron flavescens ......... 24 Phragmites communis of Am. auth. 462 communis var. Berlandieri ...... 462 Phryma Leptostachya ........... 646 PHRYMAOBAB ............+.0. 646 Phyllanthus caroliniensis ........ 589 Physalis angulata ¿cr coro ces 637 DAFDEOONEIN y «citar o Tih 637 barbadensis var. obscura ....... 637 ÜDbYHA A A bode 637 heterophylla var. ambigua ...... 637 heterophylla var. nyetaginea .... 637 icono Q0 PEIUS er st 637 A une 637 ATA A yo vaio P CARCER 638 IBRIIEONS Sy ccs S bans es eee: 638 uii peti i MEMINI: 637 DEBGDIA 244.0 0 ARE c nr 637 DUNARE vorher eb Ye 638 a 200 CRAT RA Ek ER 637 o AAA 637 A IT T REL. 638 1935] Page dbi Sede eld" T 638 A see e v Rn 638 Duis p RALES as 638 Physoearpus opulifolius var. inter- NAO 556 Physostegia formosior ........... 632 Intermedia o 0 9 eH o 632 VAEO TIAS A ole ie gene 632 Prts 1 AMOLICALA 20005 een 533 ORTI 533 PHYTOLACCACEAE PR ROE 533 Piekerel:weed caes ee 497 Picris Sprengeriana E ERE 680 IPI CON Berry OO OS 613 Bened A EI 529, 532 PIPTOAGING rare icons 532 DOG: Wr O ee 528 Pilea pumila. is ors oles) toes en 524 Pimpernel RISC cctv vcr eee D E ore ate 641 SCAT ICL. 25 MEL OOO 616 ide EORUM RS ENS IPM 615 A UM E ATHE SUE CES RUN. 611 PINACHAB MODI SORT 454 pem a IIS Ir 454 roce CRM NES 454 A ETE SIUS 454 Southern Yellow mic se sea nee 453 do UR ER 454 Pme- weed eea a as a 600 Pineapple Weed 5... 2.» ee 676 Pinesap oon occa EUIS e ters 613 Dentfordt- 5 AO arenas 536 ome pM M 536 SITO S Er BOUT 509 E Ao aae UPS 536 Mardon v certe E ar 536 LS OMA PIRA IT, 618 C] Gen cO Poo JO AO 537 A US eem UE 618 Pink Nec m he oe DER 586 Pink root iia 989» 618 Pinus AEG GRE PI RC PPS 453 MITA O O O e ois) 454 727 Page ODUS s.s. srian esi eroi Eee 454 sylvestris... sence eed 454 virginian& una. 454 Pmweed ....... RI eos « 601 ipe VIDO ..«. vers n RETO ee 525 PIPERACEAE ...... 53:54. 511 laner Tree cesare Bevor es 522 Planera aquatica ...... ooo...» 522 Plantago aristata ..... vae rere 647 COLAALA ....: cule s o S n even 647 elongata ....... oye sucess © 647 heterophylla ..-....- 3... 647 Innéeolata 7.5.99. Ee Re ae 647 AA sonne see 647 bl RE ER ET Ok. 647 O ve. rennen ee. 647 een ee 647 PHOGUBSDOEIDEOR eere erre 647 Blu een. IE Ts vy 647 x nni "A O SEO Oe 647 e A A eee s 647 PLANTAGINACEAE CU citur 647 Puteo ose olet 647 Downy Rattlesnake ............ 510 A. ee eene rer 677 Mudo as erras 498 a rig uec mop Sere Trea Ora 665 EU P MP Jun cp 457 a TW EE Ni 555 Platanus occidentalis ............ 555 occidentalis f. attenuata ....... 555 cidentalis var. glabrata ....... 555 Pluchea e AA O cs 665 Y A, i A scate 665 lum AI O a re 573 CINIGKASAW otero ev pea sso ss 572 irouind ^. Qe. oO ev hayes 578 A AA Mo RU Pre 573 Wild Goose Tis. oc e ec 572 LOA Annua. en ee sa 460 hapmanmafna s... c redis 460 GOmprogsn e. s Wa cen EUM 460 NEMOFAHR. os te Bro ier rrr 460 a cO cis: «mia pco n. 460 igo T Ee. Coa PEE MT PIS 460 728 Page 2 f A TU TT 460 n oa 460 VE: esse 460 Podophyllum peltatum ........... 544 DE ol | ... «cis :.0.»..... 506 Pogonia affinis .................. 509 trianthophora ................. 509 Poison Hemlock ................ 610 P E EEEE nee ne ees IT TT Do 591 TEPER TEE E ETT ET OTe 591 ss ETIESE TR IT TTE TEES 533 ig AAA 533 Polanisia graveolens ............. 552 ri gy PITT E a6 Ua e CR 552 Psleen AA 591 ee NADA 625 Polemonium reptans ............. 626 Polycodium interius ............. 614 ai RER 614 Polygala incarnata .............. 587 ET 587 o oix Erat EAR EREED 587 — var, latifolia .......... 587 LL. 517 Pm 587 vertieilista var. ambigua ....... 88 verticillata var. oce edicit 588 POLYGALACEAE ............. 587 POLYGONACEAE ............. 525 Polygonatum biflorum ovatum .... 503 eanalleulatlum |... devis 503 canaliculatum var. ovatum 503 commutatum ............ ees 503 commutatum ovatum .......... 503 rn MERERETUR TT TT 503 BEER EEE 503 Polygonella amerieana ........... 528 bó am aahoreum ¿Li tons 526 EEE RR ee 527 acre var. leptostachyum ........ 527 amphibium of Am. auth. ....... 527 MENON: «a6. situado sS 538 aviculare A eS ee ay 526 culare var. angustissimum ... 526 poda var. littorale ......... 526 aviculare var. vegetum ......... 526 aa ol A A 526 A rs o CER 527 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN s.s... res] | |! | | | » | À ] 5 |] | ns) 9 5 H scii of a Hydropiper var. projectum hydropiperoides hydropiperoides f. strigosum . ene i var. strigosum .. lapathifo la pathifotiun s var. nodosum .... littorale of a Mirren PIPA $$4&65)656€404046*29445 e. ...oo.on.o.....» .eonn..osSX.. .............. ............. o... +... As dm AAA canadensis var. radiata ........ Uvedalia var. genuina ......... DX .... ++. |! | |]! c! ] |||] c] ong ng echan eo. cortes. De ee ee 527 1935] PALMER & STEYERMARK Page OVO ETE ETT ETE EE TT IT EET 455 Pontederia cordata .............. 497 PONTEDERIACEAE ........... 497 Poor Man's Weatherglass ........ 616 Poplar Caroling or vedete oo 513 DA O ao ADO. 513 A A A E 513 A A ie rp 544 Poppy RAMMED: A O O 546 A 1.559 ET RE 546 OUT D CEPS rA e 546 MEXICAN nre senos PEU rere S 546 EIERN Sense tee ae 546 POPPY MALO Wi ere. 599 CU ee le 598 FHIged ... O ates 599 Purple messen 598 Populus O PO 513 balsamifera qo er en 513 balsamifera var. pilosa ......... 513 eanadensis var. Eugenei ........ 513 deltoido8 ee een ee 513 deltoides f. pilosa ............. 513 ri ARMA 513 heterophyHa ista ias ses 513 tremuloides = oa ch sen. een 51 Porcupine (Grass an ie eee 469 PEoriulaen e LEE 538 PANICINOLA scien eels d err. 538 MC ZIOCEA Cs ve cicero ete oar otters 538 OLETACOR elos cies ee e rd 538 DRITEIS een 538 YET MOERS ciao aa 538 eer cn HU eM 538 PORTULACAOEAE AAA Ee 537 AS ncc 591 Bert americanus ......... 455 O oen ee 455 EINIG A E iere d 455 dimorphus of Am. auth. ........ 456 QR C CIT RET ERO AO 456 NIME I gaa Goce eere stus tipo 455 foliosus var. genuinus .......... 455 foliosus var. maeellus .......... 455 hybridus of Am, auth. ......... 456 PLANTS OF MISSOURI 729 Page ilinoensis ¿ds WR «ieee is 455 Tuéons: ;:...90 8 042 te ciety 5s 455 DALANB ..2.. O PS 455 panormitanus var. major ....... 455 panormitanus var. minor ....... 456 POCUINATUS << ice o's ins er 456 pulcher .....cscecdimrwsevness 455 rotundatus ......: 9232 ees 456 iri Am. SUMAR ose. =. 456 Potentilla argu A rms 566 ub 242790. 566 nk var. seimplez ........ 566 canadensis var. villosissima .... 566 A Loser eo 566 Tonenelensig A. A 566 c. EP eee er 566 norvegiea var. hirsuta ......... 566 "ucc iobSRAES A Apt uer ME 566 ur V CONSTANCE RN AO NR 566 PAV HOM I E 5 ss ders ors 566 rivalis var. millegrana ......... 566 rivalis var. pentandra .......... 566 pr Co de eet A enieve 566 Swipes of ADE. .......... sese 566 simplex var. argyrisma ........ 567 simplex var. calvescens ........ 566 POVOLUY OLAS LT. . le eei cid < 469 AA AOS RE 469 buo Dog. AAA A PS 618 Prenanthes SIDA .....94. eren 682 ltissima var. cinnamomea ..... 682 BETH SS qaae desees wrasse 682 pes a A A: nen 682 AA A 604 tuc RI da ds 681 Primrose oon. RN PERA TET 606 MISEQUEL da io o ES nn 607 hito Evening... Ense 607 Primrose-willow ... Gera sans 605 LOREM oa o 605 PRIMA CHAS NER en 15 Princet Feather. rdg tme. 585 ist ES DM RR T 585 — me rp e 585 i Rin EM SLT 497 AES ROM. Quasi dh mr ER 497 Y Xam Wald. onen... ee 506 Yauguupm lis aj seine ees 538 E O O O de 470 ETS ren 675 A Ad een 575 Yellow-flowered Stoneerop ........ 553 MXaloWw se OCKO Ub ints os «nets oor e 550 AM SS CERE III 544 Yuce APEADSADA o... ¿ele eos enn 502 filgmenbosh A ee 502 O ev aede c pni K 502 Z Zannichellia palustris ............ 456 EN americanum ........ 587 So Re RR 480 mtd chloranthus ........... 500 II AA A ae E & 500 Pa AA: E O 500 Natalie»... Ben e rn 500 Zizania aquatica var. interior ..... 472 Zizaniopsis NILIABER . «Uh. «+ ses 472 Zizina AUTOR N IS ETT. 610 MAD. COMPE gir RU. m 611 "roo ES SOY T 586 [Vor. 22, 1935] 146 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 15 Fig. 1. St. Francis River, Wayne Co., showing gravel bars m of many Ozark streams, and deeiduous oak-hickory forest on granitie substrat Fig. 2 olomite bluffs of the White River, Barry Co., ME FREE mexicana near top and rich talus slope below. Fig. 3. Mississippian limestone with interbedded chert, the latter producing acid soils when left as a surface deposit after erosion and solution of the limestone, Jas- per Co. Pb 4. Erosion in loess, Clark Co. orphyry ‘‘shut-in’’ along Little St. Francis River, Madison Co., show- ing Pinui echinata along stream course. 35 PLATE 15 ANN. Mo. Bor. GARD., Vor. 22, 19 PALMER € STEYERMARK—PLANTS OF MISSOURI [Vor. 22, 1935] 748 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 16 Fig. 1. Bald knobs, Ozark Co., showing the advance of the forest upon the glades. Fi Porphyritic trachyte glade, summit of Hughes Mountain, St. Francois Co., showing weathering of the formation into block-like structures. Fig. 3. Grand Falls ehert I Pie Co., habitat of several southwestern species not found elsewhere in the s ig. 4. St. Peter sandstone dra on Juniperus virginiana and deciduous forest along = Jefferson Co ig. 5. Porp iens traehyte glade, typieal of the Iron Mountain area in the eastern Mi Hughes Mountain, St. Francois Co. ANN. Mo. Bor. GARD., Vor. 22, 1935 PLATE 16 PALMER € STEYERMARK—PLANTS OF MISSOURI [VoL. 22, 1935] 750 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 17 Fig. 1. La Motte sandstone bluff in wooded ravine, Ste. Genevieve Co., western limit of a number of eastern and Appalachian speci ig. 2. St. Peter sandstone bluff with slough at ik Jefferson Co. Fig. 3. Porphyritie trachyte ‘‘shut-in’’ along Little St. Francis River, Madison Co., with forest of Pinus echinata on Fig. 4. La Motte sandstone bluffs vil Pinus echinata above, Ste. Genevieve Co. Fig. 5. Sandstone outerops on high prairies, Jasper Co., loeality for Geocarpon minimum and other rare species. ANN. Mo. Bor. GARD., Vor. 22, 1935 PLATE 17 PALMER & STEYERMARK-—PLANTS OF MISSOURI [Vor. 22, 1935] 752 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 18 Fig. 1. Chert glades, outcrops of Grand Falls chert, Jasper Co. Fig. 2. Dolomite ledges near top of bald knob, with Delphinium Treleasei, Barry Co. Fig. 3. Loess mounds, Holt Co., showing high plains and prairie — on exposed ridges and slopes, and iia of forest in depressions and r ood ze of the influenee of exposure to winds and diu. on p formati Fig. ^" " Allavial bottoms of Missouri River, with lake in distanee and loess mounds on sky-line in background, Holt Co. Fig. 5. Glaciated prairies topography, Schuyler Co. ANN. Mo. Bor. GARD., Vor. 22, 1935 PLATE 18 PALMER € STEYERMARK—PLANTS OF MISSOURI [Vor. 22, 1935] 754 ANNALS OF THE MISSOURI BOTANICAL GARDEN EXPLANATION OF PLATE PLATE 19 Fig. 1. Densely forested country typical of the Ozark region near the influence of streams, and p accordant levels of hills earved by erosion from old pene- plain, Shannon Fig. 2. Beier Bluff, pom Co., locality for Cheilanthes alabamensis and other Merten speci ig. Po rn trashy bluffs of ‘‘shut-in’’ near mouth of Little St. Francis ra Madi Fi hrs pd (Joachim limestone), Jefferson Co., with typical de- velopment of + dab virginiana in background and clumps of Clematis Fremontii on exposed p Fig. 5. ““Shut- n along Little St. Francis River, Madison Co., typical of the Iron Mountain area of igneous rocks in the eastern Osark region vhi a number of eastern and Appalachian species, such as Ham is virginiana, Ilex verticillata var. padifolia, Rhododendron nudiflorum var. roseum, and Pedicularis lanceolata, are found PLATE 19 ANN. Mo. Bor. GARD., Vor. 22, 193: PY . T Bea Ni ae PALMER & STEYERMARK—PLANTS OF MISSOURI EXPLANATION OF PLATE PLATE 20 M MM nu A TEE er Ce ler Co. echinata, Amelanchier canadensis (large tree form), Schuyler Co. ANN. Mo. Bor. GARD., Vor. 22, 1935 PLATE 2 PALMER € STEYERMARK—PLANTS OF MISSOURI $ E $ 4 " n N 4 EXPLANATION OF PLATE PLATE 21 Rhododendron nudiflorum var. roseum, on wooded granitic slope, Madi- eron pulchellus. rson Clematis Fremontii, in limestone glade, Jefferson Co. Sedum pulchellum, in limestone glade, Jasper Co. ANN. Mo. Bor. GARD., Vor. 22, 1935 PLATE 21 PALMER & STEYERMARK-—PLANTS OF MISSOURI Annals of the Missouri Botanical Garden Vol. 22 NOVEMBER, 1935 No. 4 THE REACTIONS OF PLANTS TO ULTRA-VIOLET E. S. REYNOLDS Physiologist to the Missouri Botanical Garden Associate Professor of Botany in the Henry Shaw School of Botany of Washington University GENERAL CONSIDERATIONS It has been recognized for a considerable time that in gen- eral,as Ellis and Wells (725) observe, there are two effects of ultra-violet radiations upon organisms, designated as stimu- lative and lethal, with the dividing line approximately at 2900 À. If, however, various types of organisms and their dif- ferent functions are considered separately both the exact sig- nificance of these terms and the limits of the regions concerned must be defined more explicitly. In addition, the numerous environmental factors so modify the effects of any one of them that unless the conditions are exactly stated there is a conflict of results such as has been emphasized by Popp and Brown (733). It must be recognized, however, that such conflict of re- sults is due to the different conditions under which the experi- ments have been carried on, the differences in physiologic action of various combinations of ultra-violet regions, and the specificity of organisms, rather than to a non-responsiveness of plants to ultra-violet. The terms ‘‘stimulative’’ and ‘‘lethal’’ must be defined in relation to definite standards, and when we are dealing with a complex organism and especially with one of our higher plants ANN. Mo. Bor. GARD., Vor. 22, 1935. (759) [Vor. 22 760 ANNALS OF THE MISSOURI BOTANICAL GARDEN which is not so completely integrated as are the higher animals, we must recognize that these terms may be applied either to the whole organism or to one or more of its parts or functions. As is well known, the lack of a proper amount of light results in the etiolation of green plants. Under such conditions the use of ultra-violet irradiation cannot be expected to give a stimula- tion which will reveal itself in greater elongation. In fact the opposite result, as might be expected from the known action of the violet end of the spectrum, actually occurs. Under condi- tions of reduced visual illumination tomato plants irradiated with the mercury arc are shorter than those which are un- irradiated, although with a moderate amount of irradiation from the mercury arc the dry weight of tissue produced may be the same as in the somewhat etiolated, unirradiated plants (table 1). More strongly irradiated plants under the same conditions develop less tissue as determined by its dry weight, and correlated with this reduced vegatative growth there may be much more abundant formation of green fruit. TABLE I REACTION OF dias => PLANTS TO ULTRA-VIOLET IRRADIATION UNDER EDUCED VISIBLE ILLUMINATION Set A * Set B* Set C* Average increase in height i h 28.9 cm. 21.6 cm. 20.9 cm. Average dry weight 0.73 gm. 0.71 gm. 0.57 gm. Weight of green fruit pro-| (26 plants) (26 plants) (20 plants) duced 26.3 gms. 23.1 gms. 55.7 gms. * Set A (30 plants) unirradiated controls. Set B (29 plants) and C (21 plants), irradiated at 50 inches from the mercury vapor are through a water filter of Vita glass, 9 minutes and 18 minutes daily, respectively. Thus it is evident that the action of ultra-violet on these plants may result at the same time in reduction of elongation and in an increase in fruit formation. On the other hand, a reduction in the amount of ultra-violet attained by increasing air absorption through greater distance, concurrent with a 1935 REYNOLDS—REACTIONS OF PLANTS TO ULTRA-VIOLET 761 more normal amount of visible illumination as evidenced by lack of etiolation in checks, results in actual increase in dry weight, elongation, and mineral content (Fuller, ’31). Similar results have been obtained in other series of experiments in our laboratory more recently and have also been reported by others (Benedict, 34), all of which indicate clearly that the criteria of ““stimulation”” and “injury?” vary with the species and with the different conditions under which they are studied. We cannot determine at the present time how much of the effect is due to direct action and how much to indirect. Thus it is prob- able that the greater fruit formation in plants showing strongly retarded vegetative growth is due largely to the dis- turbed nutritional balances as may be inferred from the work of Kraus and Kraybill (’18), Murneek (726), and Harvey (’31), yet some direct stimulation is possible. As will be demon- strated in later papers in this series, various phases of res- pirational activities react differently to the same ultra-violet regions. Enzymic activity may be stimulated as shown by Fuller (732), v. Euler and Günther (’33), Bersin (733), and from studies to be reported in a later paper in this series, and this may occur when there is coincident evidence of injury to other systems. It is especially important therefore to examine the applica- tions of our criteria in order to avoid much needless discussion of results based upon uncertainty in their use and to determine the conclusions which may properly be drawn. In the study of the effects of radiations upon organisms many indefinite fac- tors exist, not the least of which is the variety of material used in the experiments. Itis now well understood that while there are certain broad fundamental ways in which all green plants are similar in their physiology, yet each species and variety has a physiology of its own both qualitatively and quantita- tively different from that of any other. We do not know as yet the experimental limits of this specificity in any given case ex- eept as expressed in the general results which give us the indistinet concept of the species. This one variable factor has given rise to many pages of useless discussion in the papers [Vor. 22 162 ANNALS OF THE MISSOURI BOTANICAL GARDEN on radiations. To expect all plants to respond uniformly to such a complex environmental factor as ultra-violet is to ignore their specifie character as well as the specific action of different types and quantities of energy. Another general factor which frequently is forgotten is the historical, evolutionary development of the plant kingdom. The physical concept of a smooth cumulative curve of effects from a given kind of force, such as increasing heat causing an inereasing expansion of an iron bar, has restricted value when applied to plants. They have been subjected through eons of time to cir ibed ranges or intensities of certain physical and chemical conditions to which they have become adjusted. Therefore in subjecting them experimentally to the entire range of a given force we should expect them to respond in one manner in the range to which they have become adjusted and in another manner in the ranges outside of their historical ex- perience. Thus we find that within a certain restricted range of temperature most plants inerease in size in a direct ratio to increased temperature, while outside of this range they react quite differently. It is necessary of course to take into consid- eration in such a statement a certain elastie adaptability which enables living organisms to extend their responses somewhat normally even beyond the usual range of the physical factors. These two variables, plant specificity and specific adjusted re- action to certain ranges of each environmental factor, are probably more important in influencing attempts to obtain generalizations than many other factors which have been so greatly stressed by some. As concerns growth related to any external stimulus, there are evidently three possible general conditions: (1) it may be definitely increased over that under established standards due to dominance of stimulative over retarding influences; (2) it may be definitely reduced below the established standards, due to dominance of retarding factors over stimulative ones; and (3) there may be such a balance of these two sets of effects that no significant increase or decrease can take place. It is pos- sible also that in this last case certain physiological activities 1935] REYNOLDS—REACTIONS OF PLANTS TO ULTRA-VIOLET 763 which are not reflected in growth may be stimulated or re- tarded. From the above considerations it is clear that growth cannot be a complete index of stimulative or lethal reaction of plants. Tn the field of radiations growth as a criterion of stimulation must be earefully defined. Size, wet weight, dry weight, and number of leaves must all be given careful consideration and balanced against one another, for no one of them can be used exclusively to indieate stimulation. In the phase of reproduc- tion radiations are coming to be recognized as more and more important, and we expect that stimulation will be evidenced in the number, form, and size of the fruiting structures at times when vegetative features fail to give evidence of reaction. Oc- casionally the phrase ‘‘beneficial effects” (Popp and Brown, '33) is used more or less as synonymous with stimulation, but it is liable to have a subjective rather than an objective content, and a very careful assessing of the values of the various stimuli and of criteria must be made before it can have a quantitative value. Thus a stimulated growth may be beneficial under cer- tain circumstances and stages of growth, but positively injur- ious at other times, while the reverse may also be true. Let us now consider some of the ways in which growth has been used as a eriterion in radiation experiments on plants. Experiments, such as those of Popp and Brown (728), in which the early seedling stages have been used as a test for stimulation, ean hardly be used as a proof against the concept of stimulation as applied to older plants. It is a well-known faet in general physiology that the physiology of the young organism is often quite different from that of the more mature individual. Moreover, the growth curve of an individual shows an exceedingly rapid rise during the early seedling stages, and it is doubtful whether at this time all stimulative factors could be expected to affect the rate of growth even if they could at a later, slower period. On the other hand, retarding and limiting factors may become especially effective during the period of normally active enlargement. Thus a small decrease in the water supply at such a time may have a much greater effect [Vor. 22 164 ANNALS OF THE MISSOURI BOTANICAL GARDEN than at a later stage and hence have a greater antagonistic action toward the tendencies of reaction to stimulative factors. It is entirely probable that one of the reasons why growth is so active during the early seedling stages is that there is stored away in the seed a maximum amount of all sorts of food sub- stanees, including the vitamines, stimulative to rapid growth. As a seedling matures it tends to exhaust the stored supplies, or distribute them throughout more and more tissues, or re- move them from further utilization by combining them in more or less stable, permanent compounds unless renewed by favor- able physiological processes. If, as may well be, the stimula- tive action of ultra-violet radiations is due to the manufacture in the plant of stimulative compounds, some of which may be stored in the seed, adequate to the maximum stimulation of the developing seedling, the stimulative aetion of ultra-violet would not become manifest until after the essential exhaustion of the seed reserves. Fora similar reason, if plants are grow- ing under daylight conditions in which ultra-violet adequate to maximum stimulation for a given set of conditions has been supplied naturally, then additional raying from an artificial source would not be evidenced in stimulation. Hence experi- ments performed in one season of the year may not be at all comparable with those at another season, and likewise experi- ments under different aerial conditions such as arise from varying quantities of smoke, moisture, cloudiness, ete., cannot be fairly compared, nor equal value attached to negative re- sults as to positive ones. As illustrating the seasonal effects upon ultra-violet the fol- lowing may be cited: In summer about 1 per cent of solar radiation is in the ultra-violet and about .04 per cent in the region 290 ma to 320 ma. In winter the figures are .2 per cent total ultra-violet and .013 per cent in the region 290-320 my (Laurens, ’33, p. 54). While these values were obtained for a different region than ours, comparisons of numerous other de- terminations indicate that these are fair approximations for many other regions. It has been our common experience in this laboratory that even injury from irradiation by an unscreened mereury vapor 1935] REYNOLDS—REACTIONS OF PLANTS TO ULTRA-VIOLET 769 are is much more difficult to demonstrate on tomato plants which have developed during the summer months in the green- house than upon those developed during the winter months. Older plants in the summer are likewise less easily injured by ultra-violet than younger ones. Some of these differences may no doubt be attributed to certain anatomical changes which oceur in plants when rayed with ultra-violet such as described by Eltinge (28). From the various considerations just discussed it is evident that experiments showing lack of stimulation under ultra- violet eannot be used to condemn positive results unless it is certain that the other environmental and biological conditions have been duplicated. Tf one compares the investigations re- porting lack of stimulation with those most carefully controlled which report stimulation (Fuller, '31; Benedict, 34), it is evident that no careful attempt has been made in the former to duplicate the conditions under which stimulation has taken place. It should be pointed out here that there is a decided difference in the conditions of irradiation between those in which there is given a long-time or close exposure of seeds, seedlings, and more mature plants to artificial irradiation sources and those in which short-time or long-distance expo- sures are made. Short injurious ultra-violet waves are much more fully absorbed by air than are the longer ultra-violet, and in short-time exposures the variable of visible light is elimi- nated in a practical manner and the variable of infra-red is also greatly reduced. Although both of these variables have been stressed recently (Popp and Brown, 733), as seriously in- fluencing conclusions concerning the ultra-violet stimulation, it should be noted that there is a great difference between adding a little more visible light and some extra heat to the environ- ment of a plant well supplied with these factors and adding ultra-violet to an environment almost devoid of such a factor. Shirley (729), for example, records that the rise in temperature due to the heat from his 1500-watt lamps at 24 inches distance was less than 0.5? C., while Fuller's lamp was at 100 inches and a large electrie fan was constantly used during irradiation to remove so far as possible any excess heating. [Vor. 22 166 ANNALS OF THE MISSOURI BOTANICAL GARDEN Since for all of the experiments reported from our labora- tories the plants were grown in the winter or spring in the greenhouses, and much of the ultra-violet which is accepted as stimulative in character is sereened out by greenhouse glass and reduced by climatie and seasonal conditions, it is evident that the plants would be in a condition to be stimulated by the ultra-violet from an artificial source. Such actual increased growth has been recorded in our laboratories for a variety of plants irradiated by a mercury vapor arc. Such distances and such screens were used as would remove injurious ultra-violet, and essentially eliminate the variables of added visible light and of infra-red since they constituted but a minute fraction, an average of nine minutes a day exposure, of the total of such energy received by the plants during the experimental growth period. In these experiments (Fuller, '31) where there were ten sets of one hundred plants each, both the wet weight and dry weight of all the sets rayed through filters which cut off the injurious ultra-violet were significantly greater than those of the unrayed controls. In six out of the eight rayed sets also significant increases in height occurred. The pertinent question may be raised as to how one may explain the stimulative action of small amounts of ultra-violet on organisms when we are inclined to discount the larger amounts of visible and infra-red radiations from the same artificial source. It is rather generally accepted that within certain limits growth increases quantitatively with light and heat. Onthe other hand, the changes in plants caused by ultra- violet seem to be qualitatively different from those caused by other radiations (Jacobi, '28) and certainly cannot be esti- mated in terms of the quantity of energy available in the ultra- violet compared with that present in visible and infra-red radiations. Or we may say that a small quantity of energy added experimentally is more effective in this range of radia- tions than in the others, due perhaps to a more complete ab- sorption and to the fact that the amount of ultra-violet added is proportionately very large during winter and early spring. Whether, as has been suggested (Laurens, '83), these shorter 1935] REYNOLDS—REACTIONS OF PLANTS TO ULTRA-VIOLET 767 wave lengths cause special intimate molecular or atomie re- arrangements we cannot at present know. It may be also that something akin to the rigor or tetanus produced in organisms by consecutive applications of stimuli might prevent stimula- tion in the presence of larger amounts of the same forces, but in the absence of natural ultra-violet of certain wave lengths an artificial applieation might be especially effective, particu- larly when added in successive, short doses. Johnston (732) has recently used infra-red radiations in conjunction with light and found that when the latter was well below the optimum for tomato plants the near infra-red stimulated growth, but with a higher degree of illumination the infra-red stimulation was much less. This seems to indicate that stimulative action of the infra-red is much less effective as normal illumination is approached. In conclusion we may summarize the discussion as follows: Physiological response includes both increased and decreased activities. Growth is a resultant of numerous physical and chemical reactions. Stimulation or increased growth may therefore be a resultant of some increases and some decreases of physieal and chemical activities. Injury and retarded growth may indicate either a total resultant reaction of all the physical and chemical reactions or interference with some one or several specific reactions. Ultra-violet and X-radiations cannot be expected to affect every one of the numerous physical and chemical activities in an accelerative fashion or, on the other hand, all in a retarding fashion. Evidences are found all through the numerous studies on radiations to substantiate these statements. It is to be expected, therefore, that there will be numerous apparently contradictory results as long as growth alone is taken as a criterion for stimulation and inter- preted as the result of purely quantitative applications of energy. Thisis especially true if the results from studies upon different species and varieties are massed together and com- pared. Itis evident therefore that experimental studies which have failed to produce acceleration of growth cannot be used to impugn those which have demonstrated stimulation. [VoL. 22 168 ANNALS OF THE MISSOURI BOTANICAL GARDEN The element of critical surprise evidenced in Popp and Brown's (733) summary of ultra-violet work, that such con- tradietory results are published, and the implieation of un- trustworthy data are not wholly justifiable in view of the considerations discussed above. We should expect just such a condition, and instead of attempting at the present time to amalgamate all such work into a consistent unit and to evaluate it in terms of uniformity of growth results, it would be more profitable to recognize the existenee of numerous separate problems and allocate to each the data properly belonging to it. In some of the earlier work where only a reconnaissance of the field has been attempted conditions have not been com- pletely controlled, and it is indeed true that in other studies unjustifiable deductions have been made, or poor technique has made the work essentially valueless as soon as more carefully controlled experiments are reported. Finally, the historical development of the plant kingdom has led to adjustments of plants to restricted ranges of environmental factors. Within these ranges any green plant will react quite differently than it will outside of them, and therefore even opposite reactions may be expected from the same plant in these different ranges, as, for example, stimulated growth in some radiations and retarded in others. A series of investigations has therefore been initiated and is in progress in which separate physiological activities are being studied in relation to ultra-violet, in an effort to define the conditions under which stimulative and lethal actions occur. LIST OF REFERENCES Benedict, H. M. (*34). Effect of ultra-violet radiation on growth and on the calcium phosphorus content of plants. Bot. Gaz. 96: 330-34 ent T. (733). Über die Einwirkung von Oxydations- und Reduktions-mitteln auf Papain. II. Die ger ussung dureh Licht, Organoarsenverbindun- gen und Aseorbinsaure. Zeitschr. f. physiol. Chem. 222: 177-186. Ellis, C., and Wells, A. A. (725). “of chemical action of ultraviolet rays. New York, 95. Eltinge, Ethel T. (728). The effect of ultra-violet radiation upon higher plants. Ann. Mo, Bot. Gard. 15: 169-240, v, Euler, H., and Günther, G. (733). Enzy deine und Enzymbildung in lebenden Zellen. Zeitschr. f. physiol. Chem. 220: 69- 1935] REYNOLDS—REACTIONS OF PLANTS TO ULTRA-VIOLET 769 arta H. J. (731). Stimulatory effects of radiation from a quartz mercury vapor upon higher plants. Ann. Mo. Bot. Gard. 18: 17-40. , (732). Some effects of radiations from a mercury vapor are in quartz upon enzymes. Ibid. 19: 505—531. valid E. M. (731). A preliminary report on the vegetative growth of okra (Hibis- us esculentus Linn.) in relation to the RR of varying amounts of re- 19 pp. Jaeobi, G. (728). Untersuchungen über die Wirkung dis ultravioletten Lichtes auf Keimung und Waehstum. Beitr. Biol. Pflanzen [Cohn] 16: 405—464. jdn. E. S. (732). The EN Uns of radiation in the physiology of -" LE Some effects of near infra-red radiation on plants. Smithson. Mise. Coll. 87'*: 1-15, pl. 1-4. Kraus, E. J., and rar H. R. (718). Vegetation and reproduetion with special reference to the tomato. Oregon Agr. Exp. Sta. Bull. 149: 90 pp. Laurens, H. (733). The physiologieal effeets of radiant energy. Am. Chem. Soc. Monogr. 62: 610 pp. Murneek, A. E. (726). Effeets of correlation between vegetative and reproduetive functions in the tomato (Lycopersicon esculentum Mill). Plant Physiol. 1: H Popp, H. W., and Brown, Florence T Is ultraviolet radiation stimulating to plants? AB. Jour. Bot. 15: 623-6 ’33). is review of recent work on the effect of ultra- violet allen upon desd plants. Bull. Torr. Bot. Club 60: 161-210. Shirley, H. L. (29). The influence of light intensity and light quality upon growth of plants. Am. Jour. Bot. 16: 354-390 STUDIES IN ULTRA-VIOLET AND RESPIRATORY PHENOMENA. I! Review or Work PUBLISHED BEFORE JUNE, 1935 F. LYLE WYND Henry Shaw School of Botany of Washington University Formerly Assistant in Botany in the Henry Shaw School of Botany AND E. S. REYNOLDS Physiologist to the Missouri Botanical Garden Associate Professor of Botany in the Henry Shaw School of Botany of Washington University T. Introduction 2.22... v deter RE ES © e ES u RR I ec) e Mie aj ie ere 772 II; Basal metabolism of animals eseni. ados a 010 0's ale aaa oe 773 A. TOWER LOMB on. erinan do WO DERI WIE ass ee ee 773 BISVertebrBie8 ce... ee OUI Me une eis sra e 66 715 1. Entire organisms sree oo. 9 casaca soe nies ole e inre 775 2. “Individual. tissues 2... «3.9 sS a ea ee eis eem es 778 3. en, Studies. 2 0 ee re ee er een ses 779 €. Human ‘subjects ....... ooo «ecosmecajco none cco senden neuen 781 TIT TA ka artificial botra foc 6 se 6 5 oes ols os ev eer aa 781 . Irradiation as a climatic factor «...oomoooroooonmo.rrrorrrcnonos 784 D. Theoretical’ considérations ......- 0: ee lemas sp ne neben ees sic 787 III. Metabolism of plants ..........ooonorroporenororaracionor.rsones.s 790 Tung A a wear Hanne cS eee PRAE 790 1. Bacteria osos seas só "XT MESURES de alas ole. seele ero MEE 790 PE BOYD det oo core O I Ou QU GC UD D IO IQUIQUE 792 O e eer v oie. siehe dr UON E s eee e^t 6.0 6 AO 793 B. Green plants ...... oce o eee seed hhoTarubessesevocccoevecos 800 C. Theoretical eee Be ee OSU ODN SOO ee on 801 IV. Oxidizing enzymes .......-. «v9 tence net sesso sie ees se seno 802 A-'OntalüNe sal eoe c ble A A RECO Ue adhere tee 802 1. In vivo studies maosna e cic ose A A Ve des ce 802 ADDRESS ere eae APPA OOO RIO 802 pog cd LC EG ERES XU BOE 805 9. In vitro studiò co... nee ee aie FS eder ee pA d eee 805 9. Theoretical considerations .... aes 95932 9 3] 4 2 US an ee 810 1 The bibliographieal study reported in this paper was carried on through aid from the Science Research Fund of Washington University, St. Louis, provided by the Rockefeller Foundation. ANN. Mo. Bor. GARD., Vor. 22, 1935. CUL) [Vor. 22 172 ANNALS OF THE MISSOURI BOTANICAL GARDEN AI eT Ce ere err er en ee co dcos 812 De AEN Sii pao ce ok eir en Ios tas 812 Bx o dar ER 812 i SM EA esos Cit ede rods 813 VEM 440 IS O aC HE ER a RR 814 : slc A MI AP a 814 UO RR A Y ON 814 $ pise: atras A IA 814 CIO a 815 ks Pisone TURTRET ELE Cee T ee ee Were ad wxa 3 EEE 815 a ARO NR EEE Sree: VE 815 N EEE py a! ike er aa cae | P 816 PNE Goby 0404.5 IO ews eie Era, 816 IC ENON cd C ko hone TTD 816 NN ess ee IIA E ES 817 3, Theoretic loot tenia. ¡ES AKG S KARA OR EE ZEE EE VE AR 819 M oan AA A TN 820 1. Animal um TELET a ETN TEE TE 20000 RS 820 «o. RP A P S 821 a EE TT ERU EPA T EPE ei 821 an a TE ETE E E E S 821 ANM QUNM E RTTA NEE OT 822 4. Theoretical considerations .............cccccccccccccccccccces 824 D WENN IND a a bebe era enl 824 mM qM isa II T E 824 B. oe TO iia rd 826 E. AA a od 826 VII. o CEP rra reis Ad 827 I. INTRODUCTION The study of the relationships between ultra-violet light and biological materials has become enormously extended in recent ears. The nature of the problems treated is so diverse that the field is already breaking up into smaller divisions, each of which is expanding so rapidly that it fully occupies the energies of students who wish to keep abreast of the developments. As soon as any field of research reaches these proportions, the need of critical reviews and extensive published bibliographies becomes increasingly apparent. The present paper is a critical review covering the effect of ultra-violet radiation on respiration phenomena. It is scarcely possible that no paper has been overlooked, yet it is felt that the bibliography is essentially complete. Only papers giving data indicating the significant presence of ultra-violet light 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 773 have been included, and many studies on sunlight and on light from indefinite sources have not been mentioned. Reviews of ultra-violet physiology containing material on respiratory phenomena have been published by Euler and Lindberg (712), Pincussen (21a, ’30), Meyer (726), Lippmann (28), and by Laurens (733), all of which are either outdated or inadequate. The best general review, especially from the viewpoint of biological theory, is that of Pineussen (730), while the most recent work is that of Laurens, which, because of its wide scope, cannot give an adequate survey of all the pertinent literature. II. Basan METABOLISM OF ANIMALS A. LOWER FORMS Merker (725a) was the first to report the respiratory activi- ties of irradiated lower animal forms. In his earlier experi- ments he used a 1000-watt filament lamp, and although he makes no mention of ultra-violet rays, a light operating at such a wattage must have emitted a considerable amount of the longer ultra-violet. Merker was primarily interested in the problem of photosensitization by fluorescent substances. He treated living earthworms (Lumbricus terrestris) for several hours with a .001 per cent neutral-red solution, and then rayed them in a constant-temperature water bath. By determining the loss of dissolved oxygen from the water inhabited by the organisms, he calculated the oxygen consumption and found that it was markedly increased, but he concluded that the met- abolie effects of the increased movement of the organisms when rayed could not be separated from the specific effect of irradia- tion. Later (25b) he repeated these experiments with Allolo- bophora caliginosa, but extended the period of irradiation to 6 hours. Anincreased use of oxygen was evident for the first 200 minutes, after which there was a decrease. These worms lived until the next day, but showed definite injury. In yet another publication (25d) these results are duplicated, and in the following year (726) apparently the identical data were republished. [Vor. 22 774 ANNALS OF THE MISSOURI BOTANICAL GARDEN These papers were followed by another (Merker and Brau- nig, 27), which reported the results of raying a number of species of moist-skinned animals with the quartz mercury are at a distance of 18.5 em., for various periods. Both Allolo- bophora and Lumbricus, which were rayed for about 20 min- utes, showed a decided decrease in oxygen use, and the raying of Gammarus pulex L., a swimming amphipod crustacean, for 54 minutes resulted in a very great decrease. In all cases of injury to the oxygen consumption, the termination of the ir- radiation allowed the return to about the normal rate. This return to normality was temporary, showing that the organ- isms had been permanently injured. Studies with filters showed that the injurious region was between 3660 Á and 3840 Á and that red light, 7800 À to 6500 À, had no effect. Merker's experiments were not subject to definite interpre- tation because the organisms were stimulated to active move- ment, but this difficulty was satisfactorily met by Sonne (726, '29). He used the pupae of meal worms, which show no move- ment when irradiated and have no disturbing food factors. In his earlier paper, he reports that the unsereened mercury are produced a 40 per cent increase of oxygen use with an exposure of 45 minutes, and that the pupae turned black and died in a few days. Pupae rayed under a glass filter or through solu- tions of methylene-blue or potassium bichromate apparently were uninjured, although the oxygen stimulation was 20, 10, and 0 per cent, respectively. The respiration of excised tissues responded to heat, as did living tissue, but there was no re- sponse to irradiation. Sonne concluded that the action of light must depend on the presence of nervous and vascular tracts, and hence he made the unwarranted assumption that mineral metabolism must have been concerned in some way. This work was carefully repeated three years later. The pupae rested on a thermo-needle which indicated any rise in temperature during the raying. The temperature of the water bath was so varied that no significant temperature change oc- eurred in the organism. The open flame of a Kromayer lamp was used, as well as various filters, the irradiation lasting 30 minutes to 1 hour. The full are gave a 40 per cent increase in 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 775 the oxygen uptake, a blue filter gave a 10 per cent increase, and a glass filter passing the visible and near ultra-violet gave an increase of 20 per cent. Sonne (’29) stated again that an in- crease in metabolism must bear some relation to the beneficial effects of raying on mineral metabolism. There is no evidence to support this assumption, for, as will be shown later, there is no undoubted metabolic change of human patients when rayed with the usual therapeutic doses. Wickwire and Burge (’27) studied the effect of irradiation on the sugar assimilation of Paramecium caudatum. They used a Cooper-Hewitt mercury are, operating on 170 volts and 3.5 amperes. Five cc. of the washed and centrifuged organisms were added to 100 cc. of a .1 per cent solution of dextrose, levu- lose, or galactose. The suspensions were rayed for 10 hours at various distances, after which the amount of sugar remaining in the solution was determined. When the raying was carried out at a distance of 50 em., all the cells were killed and the sugar use was lowered 80 per cent. When rayed at 400 cm., the lowering was 35 to 40 per cent. In the latter instance, the cells were as active as the controls and appeared to be uninjured. In view of the fact that the activity of insulin is destroyed by ultra-violet light, these authors added insulin periodically to the solution and found that the lowering of the sugar use was prevented. It was concluded that the ultra-violet light affected the Paramecia by destroying their insulin. Absorption spectra showed that the cells absorbed some of the shorter waves, but that no bands were completely removed. Ultra-violet is ab- sorbed so variously by organic substances that no physiological significance can be attributed to the specific absorption that these authors obtained. Roskin and Schischliajewa (’34) obtained irregular results when they studied the effect of ultra-violet light on the oxygen uptake of Paramecium, although they concluded that in gen- eral the oxygen consumption was lessened. B. VERTEBRATES 1. Entire organisms.—Various vertebrates were studied by Merker in connection with the problem of photosensitization. [Vor. 22 116 ANNALS OF THE MISSOURI BOTANICAL GARDEN He treated ('25a) the overwintered larvae of Alytes, a genus of obstetrical toads, the summer carp, and ‘‘Schlammpeitz- ger," an indefinite term for the Cobitidae, an Old-World family of small fishes, for several hours in .001 per cent neutral-red, and found that the illumination by a 1000-watt light gave no- table increases in oxygen use. In a series of papers he reported this generally true. However, irradiation from the mereury arc always gave an important decrease in oxygen uptake. The paper by Merker and Braunig (27) describes experiments with Gammarus pulex (young white fish), Gasterosteus aculeatus (a stickleback fish), Rana esculenta, and larvae of Molge cristata and M. taeniata (both belonging to the Old-World salamanders). Harris (25a) rayed rats in a small quartz respiration cham- ber with either a mercury are operating on 15 kilowatt, or a carbon are operating on Y, kilowatt. He used the full arc, the visible only which he obtained by a plate glass filter, and ultra- violet only obtained by a “Blue Uviol” filter which passed rays between 2910 A and 4360 A. A constant temperature was maintained and the carbon-dioxide elimination was estimated at 15-minute intervals with an accuracy of * .5 per cent. The results were calculated in terms of mgms. of carbon dioxide eliminated per square em. of body surface. In one experiment neither the unscreened are nor the visible alone gave a sig- nifieant effect, but ultra-violet produced a definite stimulation. This author concluded that visible light nullified ultra-violet stimulation by a physiologieal antagonism, and not by any physieal interference, and he made the interesting comment that this is the reason that human subjects suffer less in sun- light than when irradiated by equivalent dosages of pure ultra-violet. It is not possible at the present time to conclude definitely concerning the physiological antagonism of different wave lengths of light. This phase of the problem must wait for the fuller development of monochromatie studies. The work of Harris was questioned by Campbell (26). He studied mice and rats in a constant-temperature chamber. The 1935] WYND & REYNOLDS—ULTRA-VIOLET. I ui unsereened illumination of a Cooper-Hewitt mercury arc, oper- ating on 110 volts and 3.5 amperes, at a distance of 25 cm., was used for the first 114 hours, a “Blue Uviol"' filter for the next 115 hours, and then an opaque screen for the next similar period. He found no metabolie effect, either immediate or de- layed, of any of these spectrum regions. The question was reopened by Crofts (728a), who criticized previous students because they did not keep the rayed animals under observation for a sufficiently long period after the ir- radiation. She used canaries, because of their marked physio- logical sensitivity, and observed their oxygen use for a period of 6 days after irradiation. A Hanovia mercury are was the source of illumination, at a distance of 25 cm., for a period of 10 minutes, this being equivalent to 14 lithopone units. The feathers were removed from the under side of the body to expose the skin. There was usually an abrupt drop in oxygen use immediately after raying, which averaged about 14.1 per cent. Single heavy doses caused a belated rise after the preliminary drop, averaging 21.6 per cent. Still heavier initial doses extended the period of depression. Daily rayings gave the usual depression, but no subsequent rise. It is par- ticularly significant that there was no relationship between size of dosage and the amount of depression or elevation. This agrees with the present authors’ results on yeast, which is re- ported in the third paper in this series, but it is improbable that similar mechanisms are involved. Hardy (’28) extended the work of Crofts to rabbits, and in- creased the period of observation to 12 weeks. She used single doses of 24, 60, and 90 lithopone units, and also daily doses of 20 and 30 lithopone units which gave a total of 120 and 180 lithopone units respectively. She found no effect on the rest- ing metabolism. During the same year, Vasetko (’28) rayed white mice for 3 minutes with a mercury arc of the artificial sun type at a dis- tance of 30 em., and measured the heat production in a calo- rimeter for 10 hours. "The heat production inereased slightly. No observations were made on gas exchange. [Vor. 22 778 ANNALS OF THE MISSOURI BOTANICAL GARDEN 2. Individual tissues.—The first observation on the effect of ultra-violet on the respiration of individual tissues was made by Gottschalk and Nonnenbruch (723). They ground 2 gms. of frog muscle in 10 ce. of water to which .2 gms. of m-dinitro- benzol had been added. The preparation was rayed with a mercury are at a distance of 70 cm. for periods of 15 to 90 minutes. Exposures of 45 minutes or less inereased the oxy- gen use, the maximum stimulation of 24 per cent occurring with 30-minute exposures. Dosages of 60 minutes or more were depressing. Wohlgemuth and Szörényi (*33c) placed the tissues of liver, kidney and skin in Ringer's solution, and studied the gas ex- change by the manometrie method of Warburg. A 75-watt lamp was suspended in the water bath 10 to 12 cm. from the vessel containing the tissue. In the absence of photosensitizers there was either no effect on the oxygen absorption or a slight depression, but in the presence of haematoporphyrin, there was a well-defined stimulation. This photosensitized fraction of the oxygen uptake apparently was not normal respiration, since it was not affected by temperature and since M/300 KCN not only did not inhibit it but actually had a stimulatory effect. Other experiments were carried out with a carbon are drawing 30 amperes and with a quartz mercury are. In these instances the light was removed 50 em. from the vessels containing the tissues. In the presence or absence of sensitizers (haemato- porphyrin or Bengal-rose) there was a decrease of the oxygen use. 'lhe sensitizers alone caused a decrease of 6 to 22 per cent, but when rayed with sensitizers the decrease was 17 to 43 per cent. Guinea pigs that had been injured by raying usually showed an inereased tissue respiration of the excised organs, and also occasionally an increase of anaerobic glycolysis. The authors postulated that these effects were probably due to the forma- tion of toxie substances during the raying. "They also (733a) studied the effect of raying normal tissue of liver, kidney, skin, brain, and retina of guinea pigs. In the presence of sensitizers, irradiation with a carbon arc decreased oxygen use, probably because of a photo-oxidative destruction of protoplasm. If 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 779 sensitized guinea pigs were rayed until they showed acute photo-dynamic injury, the excised tissues showed an increased oxygen use, probably due to the formation of a toxic substance. Since Ellinger had shown that ultra-violet irradiation changed histidin to histamin both in vitro and in vivo, Wohlge- muth and Szórényi (33b) attempted to show that histamin would eall forth the same physiological effect as irradiation. Histamin stimulated the oxygen use by kidney, liver, brain, and skin, but Jensen sarcoma was unaffected. 3. Biochemical studies.—Pineussen (21a) attempted to show an analogy between the action of ultra-violet on oxalic acid in vitro and in vivo. If oxalic acid is destroyed in vivo, then determinations of this substance in the urine can give no picture of oxidation within the body, and this uncertainty is further increased by the doubtful significance of its presence in the urine. In studying this problem, Pincussen injected oxalate subcutaneously into the rabbit, and then rayed the ani- mal with a carbon are. In control animals, 75 per cent of the added oxalate was recovered in the urine, but when the animals were sensitized by eosin or by the sodium salt of dichlor- anthracenedisulphonie acid, only about 45 per cent of the oxalic acid could be recovered. The conclusion was that oxalie acid was an end product of normal metabolism, and that irradiation caused the oxidation of the acid. By chemical analogy, the end produet of the raying must be carbon dioxide and water. Biochemieal changes connected with metabolism were studied particularly by Pineussen and Zuckerstein (29). Guinea pigs were fasted several hours, with and without previous eosin injections, then rayed, and killed 30 minutes later. A study of the individual organs showed that fat markedly increased in the serum but cholesterin did not change, while in the heart and liver, fat decreased but cholesterin increased. Eosin injection inereased the magnitude of these effects. In the kidney, fat changed but little and cholesterin increased. The previous injection of eosin diminished cholesterin. In muscle, there was but little change in the absence of the eosin, but with it present fat was lowered. [Vor. 22 180 ANNALS OF THE MISSOURI BOTANICAL GARDEN In a further study of the biochemical changes in separate organs, Pincussen and Kawakami (729) used a mercury are operating on 140 volts and 4 amperes. Rats were rayed at a distanee of 15 em. for 30 minutes, after which they were killed and the organs removed. The liver and muscle showed an in- crease in glycogen and a decrease in lactic acid. The tables included in their paper show that the ratio, glycogen: lactic acid, is increased in the liver about 55 per cent and in the muscle about 45 per cent. In the heart no significant difference was found. The increase of carbohydrate in the organs of rayed animals had already been reported by Jacoby. The change in the glycogen: lactic acid ratio involves, according to these authors, a lower catabolism of carbohydrate, and probably also an inereased synthesis, and the relation of these results to diabetes was suggested. Raying induces for a short period a small increase of blood sugar followed by a decrease. In diabeties, the sugar in the urine is also deereased, this being explained not by an inereased oxidation of sugar but by its re- tention and storage in the liver and muscle as glycogen. In later experiments, organs were removed from normal rats, mineed, suspended in a thin layer, and then rayed with a mercury arc. The fact that results were not consistent indi- cates the possibility that the specific reaction of living animals to ultra-violet might not be due to the action of the irradiation as such, but rather to secondary reactions due to a generally disturbed metabolism. Wohlgemuth and Szórényi (33a) reported that ultra-violet increased the anaerobie lactic acid formation in the absence of sensitizers, but diminished it in their presence. The source of radiation was either a carbon are or mercury are. Anaerobie glycolysis by the tissues of guinea pigs in the presence of sensi- tizers was shown (33e) to be injured by intensive raying with a carbon arc. In the absence of sensitizers the glycolysis re- action was increased. Since ultra-violet irradiation had been shown to change histidin to histamin in vivo, they ('33b) studied the effect of histamin directly. Although tissue res- piration was increased, anaerobic lactic acid formation was not changed. 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 781 C. HUMAN SUBJECTS 1. Irradiations from artificial sources.—Hasselbaleh (705) was the first to record the effect of ultra-violet on human me- tabolism. This worker rayed himself under carefully con- trolled conditions with a carbon are, operating on 65 volts and 150 amperes. Relatively heavy exposures of about 1 hour were used, with the result that the frequency of inhalation was lowered, but the amplitude increased, the total ventilation therefore remaining unchanged. This condition may last for several days, and frequently longer than the erythema. Durig, Schrötter, and Zuntz (712) also found a slower rhythm of inhalation. The effect on actual metabolism was not definitely demonstrable, although their tables do show an in- ereased use of oxygen by some subjects. The results of raying patients suffering from various heart diseases were reported by Rubow and Sonne (712). The carbon are eaused reduced frequency of inhalation and an inereased amplitude, which were progressive in magnitude as the daily rayings continued. These results agree with those obtained by Hasselbaleh on healthy patients. Rubow and Sonne ob- served that the development of erythema was necessary for the changes in inhalation and that super-sensitive patients who de- veloped erythema with 5- to 10-minute exposures exhibited the typical reactions. These data were verified by Austgen (719), who rayed tubercular patients twice a week with the mercury arc at a dis- tance of 70 em., the dosage being increased from 3 to 20 minutes at each exposure. His results were very irregular. Of 70 patients, 35 showed no change, in 21 the rhythm of inhalation was increased, and in 14 it was lowered. Éderer (722) also studied this problem and reported that short exposures to the mercury arc had no effect on alveolar carbon-dioxide tension, but that moderate doses increased it, and large doses lowered it. These effects were found to be in- dependent of the development of erythema. Kestner, Peemóller, and Plaut (723) observed an increase in oxygen use unless erythema developed. Their subjects had [Vor. 22 782 ANNALS OF THE MISSOURI BOTANICAL GARDEN - fasted 12 hours, and were then rayed with a mercury are at a distance of 50cm. Animals with hair showed a reaction similar to the human subjects who developed pigment. These authors point out that the increase in oxygen absorption depends on the activity of the chemically active rays, and on the absence of heat rays. Since the application of heat to warm-blooded ani- mals decreases their heat production this factor would counter- act any stimulatory effect of the ultra-violet. This problem will be discussed more fully under the topic, ‘‘Theoretical considerations. ”” Several papers by Kestner and his co-workers ('23, 25, ’26, ’27) reported increased metabolism due to intense sunlight. Contrary to Kestner, Kroetz (’24) found that even though ery- thema always developed physiological changes occurred. The blood showed a temporary acidosis, followed by an alkalosis which was maintained for about a day. He attributed a stim- ulation of the respiratory center to the general effect of raying the organism. The presence of eosin did not influence these results. Fries’ (25) experiments with children were carried out from December to March, since the ultra-violet from the sun is not strong enough during these months to confuse the re- sults. Three children were kept on a controlled diet for one month, during which time their basal metabolism was deter- mined three times a week. They were then rayed with a Han- ovian mercury are under various conditions, and the basal metabolism again determined three times a week. Each de- termination followed the raying by 20 to 70 hours. One patient was rayed 20 times in 24 days, one at irregular intervals, and one 7 timesin16 days. The exposures were at 24 inches at first, then lowered to 16, and began at 3 minutes and were gradually increased to 15. All three subjects became pigmented. Two other children not under controlled conditions were rayed. None of these 5 patients showed any increased metabolism. In two children rayed in the month of April to May, the metab- olism rose after the third and fifth treatment and then fell 13 per cent and 11 per cent below normal. The ultra-violet 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 183 from the sun may have confused these results. The work of Fries shows that any prolonged rise in metabolism is not usual, and that although individual variation may influence the re- sults, certainly the clinical improvement in malnourished chil- dren as a result of raying does not depend on an increase in metabolism. Fries published later (27) on the same problem and again found that the metabolism of rayed children did not vary more than 10 per cent. "This is an insignificant variation in view of the inaccuracies of determining basal metabolism. In the same year, Fries and Topper (727) extended the study. The patients were rayed 3 times each week at 20 inches with a mer- cury arc. The first exposure was 3 to 5 minutes, on the chest and back, and each succeeding exposure was increased 2 to 4 minutes. All patients developed good pigmentation. In the previous experiments, the metabolism determinations were made 20 to 70 hours after the irradiation, but in order not to lose immediate effects the determinations were now made be- fore and after each treatment and also 2 and 6 hours later. There was no change in metabolism, pulse rate, or blood pres- sure, that exceeded experimental error. A more extended study was made by Mason and Mason (726) who rayed 10 patients with the unsereened mercury arc. Eight showed a notable lowering of heat production and a lower pulse rate. In no instance was an increased metabolism observed. The subjects showing a depressed metabolism were referable to two groups: (1) normal patients who pigmented well; (2) patients having an increased amount of circulating bilirubin. The 5 individuals of the first group showed a lowered metab- olism of 13.8, 15.6, 14.5, 7.8, and 10.5 per cent, which was pro- portional to the degree of pigmentation. Sometimes this lowered rate persisted as long as 23 days after the raying ceased. The 3 patients constituting the second group showed inhibitions of 24.1, 24.5, and 8.0 per cent. The interesting theoretical significance of these results are discussed under the heading, ‘‘Theoretical considerations. ”” Eichelberger (726) rayed 9 women and 2 men in order to de- [VoL. 22 784 ANNALS OF THE MISSOURI BOTANICAL GARDEN termine whether creatinine was connected with respiratory metabolism. The fact that changes in creatinine excretion had no effect on metabolism convinced this author that they were unrelated. 2. Irradiation as a climatic factor.—Hasselbalch and Lind- hard (711) reported that with increasing altitude there is an increasing sensitivity of the respiratory center. This effect was attributed to the increased ultra-violet reaching the sub- jects from the rays of the sun. During the first day at a high altitude, there is an abnormally great sensitivity of the respira- tory center, but physiological adaptation soon allowed a re- covery. The final condition was a subnormal sensitivity. These authors believe that the lowered frequency and deepened amplitude of inhalation were due to the reduced tonus of the vessels in the skin. A lengthy report, which unfortunately was based on scanty material, was published by Durig and Zuntz ('12). Ata higher altitude the respiratory quotient remained constant, although the alveolar carbon dioxide and oxygen tension de- creased proportionally to the altitude. The body temperature was not affected. An increased frequency of breathing was shown by 2 subjects, while 1 showed an increase. The ampli- tude of inhalation also was decreased in 2 subjects, but in- creased inl. These results are obviously unconclusive. In the same year, Durig, Schrótter, and Zuntz (12) attempted to cor- relate high altitude effects with those produced by artificial raying. The result of raying depended on the individual and on the intensity of exposure. The alveolar carbon dioxide was reduced, and many subjects showed a decreased frequency but a greater amplitude of inhalation. It was concluded that climatie effects had no correlation with those of raying. Kestner and his co-workers have maintained in a series of papers that the short-wave irradiation in high-altitude sun- light stimulated human respiratory metabolism. Kestner, Peemöller and Schadow (727) reported the results of 5 men ascending a mountain, each person making metabolism tests on himself. The locality was chosen so as to obtain the most in- 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 185 tense solar rays possible, but at an altitude of less than 3000 m. so that the oxygen pressure would not be a disturbing factor. The intensity of the radiation was measured by a cadmium cell and an electrometer. This cell almost exclusively reacts to wave lengths less than 3200 Á. Dorno had shown that this length is the most physiologically active on man. This climatie study showed that this spectrum region stimulated respiration and gas exchange. "Through over-heating the metabolism was greatly reduced. Wave lengths less than 3200 À were called by these authors the ‘‘Ra’’ rays. This is an unfortunate term, since it might reasonably be confused with radium emanations if the student were not familiar with the details of the experi- mental procedure. During the same year, Kestner and Schadow (27) reported a similar study carried out at the Jungfrau Pass. The rays shorter than 3200 Á again stimulated the oxygen use. These authors state that the diminished oxygen pressure at this altitude caused a deepened amplitude of inhalation. The “Ra”” rays appeared to penetrate relatively thick masses of clouds, and the reflection from the snow was also found very important. Crofts (28b) rayed 2 subjects with a **Uviare" which gave 6.6 lithopone units of ultra-violet in 8 minutes at 18 inches. The dosages were as follows: 5 L. U., 615 L. U., a total of 20 L. U. distributed over a series of rayings, incrementally re- peated rayings giving a total of 1715 L. U., 3 large doses giving a total of 19.8 L. U., inerementally repeated rayings giving a total of 44.9 L. U. distributed over a period of 6 days. Even though these dosages were in excess of usual clinical treat- ments, there was no appreciable effect on metabolism. A very excellent review by Lippman (728) contains the gen- eralization that ultra-violet in low erythema doses inereases the oxygen use during the period of exposure 5 to 10 per cent. After 30 minutes, the oxygen consumption falls, often below normal. A report based on copious clinical material by Lippmann and Völker (728) is one of the most important contributions in this field. Dosages varied between 1.5 and 6.5 erythema units. [Vor. 22 186 ANNALS OF THE MISSOURI BOTANICAL GARDEN During the actual raying there was an inerease in metabolism of 10 to 18 per cent, this stimulation not depending on the size of dose nor upon the intensity of the developing erythema. The fact that mere uncovering of the patients produces some increased metabolism lessens the significance of the results. The respiratory quotient showed a slight lowering, perhaps due to oxidation of the non-carbohydrate substances which Pin- cussen considers to be proteins. These authors believe that the variations which they obtained in the respiratory quotient were too small and variable to be significant. The volume and frequency of breathing also remained unchanged. There was no time effect during the 24 hours of observation after the ray- ing. There was no significant change in nitrogen metabolism, and the slight decrease in the blood sugar was within the limit of usual variations, being from .32 to .27, .30 to .25, and .24 to .20. Lehmann and Szakáll (32a) reported that raying laborers with a mereury are at weekly intervals lowered their basal me- tabolism 10 to 15 per cent and increased the capacity for work up to 60 per cent. There was a decrease in the oxygen debt. These effects were thought to be similar to those of muscular training. Another experiment (732b) reported that when the same subjects were rayed, but with a sereen interposed which eliminated the ultra-violet without their knowledge, the in- crease in work capacity did not take place. Rubow and Sonne (712) had reported previously that patients suffering with heart diseases reacted favorably to irradiation and that their capacity for muscular work increased. In a more complete study, Lehmann (33) rayed a patient for 9 minutes and then measured the heat production hourly for 22 hours. There was a small increase until 7 hours after raying, at which time erythema developed. From this point onward there was a progressive decrease, although sometimes the value was still above normal for several days. In another experiment, weekly averages were obtained for 17 weeks. Dur- ing the fourth week, the ultra-violet dosages were 3 minutes, and in succeeding weeks they were increased to 6, 10, 15, 20, 30, 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 787 and 40 minutes. There was no raying during the remaining 5 weeks. The results are very definite, and the curves repre- senting them are among the best that have appeared. The elimination of carbon dioxide showed no important change, while the oxygen use decreased continuously until the rayings were stopped and then showed a progressive return to the normal during the succeeding 5 weeks. The curve representing heat production parallels very closely the above oxygen changes. The oxygen debt also was parallel to that of heat production. D. THEORETICAL CONSIDERATIONS It is difficult to come to positive conclusions concerning the relation of animal metabolism to ultra-violet light. At first glance the experiments of Merker and his co-workers on lower forms show that a 1000-watt filament lamp inereases the oxy- gen uptake, but that a mercury are always decreases it. How- ever, the presence of eosin confuses any determination of what wave lengths are effective, and, as he himself admits, the in- ereased movement of the rayed organisms may have caused the inereased oxygen uptake. Also, the method by which he determined oxygen use is open to criticism, and the curves which he published are extremely unsteady. However, if we give full credence to Merker's results, raying eosin-treated organisms with a 1000-watt light caused an inerease of oxygen uptake which resembled that produced by increased tempera- ture. These experiments contribute but little to an ultimate theory of ultra-violet physiology. The fact that Sonne used pupae, which showed no movement when rayed, and that he used a mercury arc, a much more potent source of ultra-violet, makes his experiments of more significance. He obtained an important increase in oxygen use. Although his organisms were injured, making it impos- sible to draw conclusions concerning the effects of ultra-violet itself, yet this author's work is the best evidence available that lower forms are stimulated in their oxygen uptake by ultra- violet light. [Vor. 22 788 ANNALS OF THE MISSOURI BOTANICAL GARDEN The work on vertebrates may be summarized with greater assurance. In the absence of factors depending on pigmenta- tion, it is probable that such sensitive organisms as canaries, and, to a lesser extent, white mice, etc., may show increased metabolism. But even here the results are irregular. It is to be noted emphatically that the dosages given in such experi- ments are greatly in excess of any that might be given thera- peutieally to human beings, and any suggestion that ultra- violet light owes part of its clinical effect to increased respira- tional metabolism is without foundation. The above statement is made with full knowledge of the very positive opinion of Warnshius (733). This author states that ultra-violet irradiation of human patients ‘‘is the most valu- able and perhaps the most positive means for the improvement of oxidative processes.’’ This paper contains a large amount of loosely organized theory purporting to show that ultra- violet light has a greater anti-rachitie power than vitamin D, which he attributed to the improved oxidative power of the blood enzymes. He further states that vitamin D is probably only another oxidative enzyme or a preeursor or an intermedi- ary of other oxidases. Ultra-violet acts, then, by activating the oxidizing enzymes and this affects the mineral metabolism. This theory cannot be substantiated by data in the literature. Ultra-violet experiments with human beings have not always been above criticism. Scanty clinical material, poorly con- trolled conditions, and faulty technique have thrown an un- deserved appearance of futility in this field of research. After evaluating the various published results on the basis of the conditions under which they were obtained it is fairly certain that ultra-violet irradiation gives no stimulation of respira- tory metabolism. If the effects of irradiation are influenced by pigments, such as melanin, bilirubin, etc., there is a decrease in oxygen use because the pigments degrade the incident ir- radiation into the longer wave lengths. The total effect is, then, the same as the addition of heat to the body, and conse- quently there is need of less intracellular oxidation. This viewpoint might be conclusively checked by raying negro and 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 789 white patients. The difference in pigmentation is here so great that there should be a marked drop in oxygen use by the colored patients long before the pigments could develop in the white patients, and the usual unavoidable variations should become insignificant. Merker (’25c), in a long review of the biological effects of fluorescent dyes, suggests that they may act either as oxygen carriers, or under other conditions they may destroy cellular oxidizing mechanism. "These suggestions are worthy of con- sideration, but it is possible that fluorescent substances inhibit by some physiological effect, depending on their emission spectra. The mechanism by which photo-sensitizers affect oxidation is still controversial. Gaffron (’33) believes that the oxidizing effect of irradiated dye solutions depends on an activation of the oxidizable substance. An extended study of this problem was carried out by Kautsky, Bruijn, Neuwirth, and Baumeister (733). They believe that the photodynamic effect of a fluoresc- ing dye is due to a meta-stable condition of oxygen. The oxy- gen absorbs energy from the fluorescing dye molecule and thereby causes fluorescence to disappear. Since the meta-stable oxygen has a finite time existence it may diffuse away from the point of activation to an oxidizable body which is itself not irradiated. This mechanism was convincingly demonstrated by irradiating tryptoflavine and observing the resultant oxi- dation of leuco triphenylmethane and also leuco malachite green. These reduced dyes were maintained in silica jell some distance from the point of irradiation. The statement of Wickwire and Burge (’27) that ultra-violet inhibits the oxygen use of Paramecia by destroying the activity of insulin is suggestive, but it seems a far cry to assume that these organisms are so similar to higher forms as to possess an identical mechanism of sugar utilization. The reports covering the stimulation of metabolism by solar rays and under various climatic conditions are subject to too much criticism of technique and are based on insufficient ma- terial to be of theoretical importance. This is also the attitude of Flickinger (’26). [Vor. 22 190 ANNALS OF THE MISSOURI BOTANICAL GARDEN III. METABOLISM or PLANTS A. FUNGI 1. Bacteria.—Schnitzler and Henri (’09) were the first to observe the effect of radiations from a mercury are on bacterial metabolism. They studied acetic acid fermentation of various wines, red Algerian, white Algerian, and a mixture consisting of 100 parts of wine, 200 parts of water, and 4 parts of glacial acetic acid. The solutions were irradiated at a distance of 15 em. in 50-ce. samples, 1 cm. deep, and in 200-cc. samples, 4 em. deep. An exposure of 30 minutes completely stopped acetic acid fermentation, while shorter exposures were inhibitory to a less degree. Special tests indicated that the ozone formed by the ultra-violet light was without effect, and that wave lengths shorter than 30214 constituted the effective spectrum region. It is particularly significant, as will be pointed out later, that if the wine were rayed in the absence of oxygen the ultra-violet was without effect. This paper was followed by a more complete report the fol- lowing year (*10). The irradiation was carried out as before, and the cultures watched for 3 to 9 days in a water bath main- tained at 25? C. The experimental solution consisted of 100 parts of red wine, 200 parts of water, and 4 parts of glacial acetie acid. Irradiation for 30 minutes deereased the amount of acetic acid from the control value of 5.90 to 5.05 per cent. Shorter dosages were proportionally less inhibitory. The fact that even 40-minute rayings were without effect in the absence of oxygen emphasizes their previously published results. Traces of hydrogen peroxide gave not only similar inhibitory results but also produced the same color changes in the wine as did irradiation. Hence it was concluded that the toxic effect of ultra-violet irradition by wave lengths shorter than 3021 À depends on the formation of hydrogen peroxide in the presence of atmospheric oxygen. The aetual possibility of hydrogen peroxide developing in irradiated water solutions will be more fully diseussed in the section on peroxidase. The first data concerning the effects of ultra-violet light on 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 191 the respiration of baeteria in pure culture were reported by Cook and Stephenson (728). "They studied primarily the rela- tion of viability of the bacterial cell to its aerobie oxidation of glucose and certain of its fermentation products, and irradia- tion with ultra-violet was only incidentally introduced as a means of reducing viability. "They grew Bacillus coli on tryptic broth agar to which .5 per cent sodium lactate had been added. After 24 hours, the cells were suspended in phosphate buffer at pH = 7.4, centrifuged, then washed twice in Ringer's solution, and finally suspended in 100 ec. of Ringer's solution, and aerated for 15 minutes in order to obtain an even suspen- sion. This stock suspension was diluted with an equal volume of Ringer's solution, and 1 ec. was used for each determina- tion. Glucose, 1/200 M, was used as the control culture. The substrates, lactate and pyruvate, were studied in 1/100 M con- centration, and acetate, ethyl aleohol, acetaldehyde, formate, oxalate, and glycollate were also studied. In order to decrease the viability of the cells, they were rayed in quartz test-tubes at a distance of 6 inches from a mercury are for 20 and 30 min- utes. Super heating was avoided by protecting the culture tube by another and larger test-tube. The suspensions rayed for 20 minutes showed 2.74 viable cells in each 1000; those rayed 30 minutes showed 2 viable cells in each 1000. The effects of ultra-violet on the respiration process, aside from the mere killing of most of the cells, can only be inferred. At first, there was a slight decrease of oxygen use, but later there was no significant difference. This preliminary period of slight inhibition was attributed to an injury to the respira- tory enzymes, although no actual tests were made. Since a re- duetion of the number of cells by dilution gave a rate of oxygen use proportional to the number of cells, and since a re- duetion of living cells by irradiation did not materially affect the respiration, these authors eoncluded that living and dead cells oxidize alike. The oxygen uptake was measured by differential manom- eters of the Haldane-Bareroft type. It was assumed that if the control culture were placed in the vessel on one arm and the [Vor. 22 792 ANNALS OF THE MISSOURI BOTANICAL GARDEN experimental eulture in the vessel on the other arm, the move- ment of the manometer liquid would indicate the difference in rates. If all the forces acting on each limb of the manometer were perfectly symmetrical, then the above assumption would be valid. This can rarely be the case in actual experimental procedure, as can easily be determined by placing duplicate aliquots in each vessel. Experiments by Wynd have shown the unsuitability of this method, and errors from this cause alone might easily be of such magnitude as to account for any of the differences between the normal and rayed cells in the experiments reported above. Lohmann's (*34) study of Bacillus coli, tubercle bacilli, and Streptococcus is the only one that has been direeted specifically to determining the effect of ultra-violet on the respiration of bacteria. Bacillus coli was grown in a buffered Ringer solu- tion which contained sugar. This medium allowed no repro- duetion, and the cultures exhibited a constant rate of oxygen uptake as determined by the Warburg method. A bouillon medium which allowed reproduction was also used. Tubercle bacilli were cultured on Lockemann’s medium. The most com- plete data reported in this paper concerned Streptococcus. The culture was diluted with 9 volumes of colorless Ringer solu- tion at pH = 7.4. An exposure in a quartz flask to a mercury are for 45 minutes lessened the oxygen uptake about 50 per cent, and this rate was maintained for several hours. After 7 or 8 hours new growth caused an increase in oxygen use and in elimination of carbon dioxide. An exposure of 80 minutes stopped growth completely and almost all of the respiration, but new growth began after 12 hours. Rayings as long as 3.5 hours did not completely sterilize the cultures. Experiments on B. coli showed that exposures of 3 minutes resulted in a constant use of oxygen, showing that growth had been stopped although metabolism remained uninjured. 2. Botrytis.—Fazi ('23) reported that fermentation of grapes infected by Botrytis cinerea was stimulated by exposure to the mercury arc. Although it cannot be said that this result is not valid, yet, as will be shown later, all of the work reporting 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 193 inereased fermentation in yeast by ultra-violet is open to serious criticism. 3. Yeast.—The effect of irradiating yeast on respirational activities was first observed by Maurain and Warcollier (709). They used a mercury arc of the Poulene type, operating on 110 volts and 2.5 amperes. Sweet cider was rayed in layers 1 mm. deep, at a distance of 5 cm. An exposure of 3 minutes killed fermentation. If the cider were diluted with 20 parts of water, then 2 minutes stopped fermentation. If the layer were re- duced to 14 mm., 2 to 3 minutes were completely inhibitory. In the following year, these authors (710) extended their observations to foaming white wine in an effort to determine if irradiation might be used to inhibit the undesirable late stages of fermentation. The wine was in layers 5 mm. deep, and was 4 em. removed from the mercury arc. Irradiation for 10 seconds completely inhibited fermentation. The greater sensitivity of the white wine was due to its greater transpar- ency for the effective radiations. The most prolifie writer in this field of research is Fazi. His name first appears in the literature in the announcement of a British patent (714) for carrying out commercial fermentation under the influence of ultra-violet light. He points out that either sunlight or ‘‘Uviol’’ lamps may be used as the source of irradiation. The fermentation vats were to be made of “Uviol’’ glass, or the lamps immersed directly in the liquids. The announcement of this patent was followed by a long series of papers, all purporting to show great stimulation of fermen- tation by ultra-violet light. The first paper reported that beer yeast could be rayed profitably even for as long a period as 12 hours, and that it was not injured by 14 hours of exposure. The irradiation was carried out in quartz flasks, 20 cm. from the mercury arc. Fazi and Fazi (716) describe experiments showing that Saccharomyces opunitiae lives, reproduces, and ferments more actively when stimulated by irradiation. Many hours of ex- posure were needed to cause injury. Also, the must of India figs forms abundant film on the surface in ordinary fermenta- [vor. 22 194 ANNALS OF THE MISSOURI BOTANICAL GARDEN tion, but under the influence of ultra-violet there was no trace of film. These authors later ('17) mixed Saccharomyces cerevisiae with S. opunitiae and found that the rate of fermen- tation of fig must was doubled by ultra-violet light. The change of sugar to alcohol and carbon dioxide was almost com- plete, while in the unrayed cultures there remained a residue of sugar. Pure cultures of S. cerevisiae also were rayed in fig must, and at the end of the experiment the samples contained less acid, about 10 per cent more alcohol, and a smaller residue of sugar. Fazi (21) published new data in order to contradict a paper by Feuer and Tanner (720), who had stated that their yeast could not endure more than a 1-minute exposure. Fazi not only states that this result is opposed to his own, but also to the observation of many others. He mentions partieularly Henri and Stodel (’09), Cernovodeanu and Henri (710), Vallet (710), Lombard (710), Van Aubel (709). Fazi elaims to have done much work on bacteria and on yeast, although there is no ex- tensive publication of data. In the above paper he described experiments in which brewer's yeast was rayed for 12 hours with a 1200-candle power lamp at 20 cm. The yeast was not in- jured and showed an increased fermentation, but all the bae- teria in the suspension were destroyed. This method of clear- ing yeast from bacteria was instituted in 1918 by the Peroni Brewery, in Rome. The significance of Fazi's paper was completely nullified by the reply of Feuer and Tanner (721) in which they state that Fazi misquoted them in saying that the yeast died with 1 minute of raying, since they actually had said that some samples endured exposure for 7 minutes. It is significantly pointed out that the suspensions were very dilute, consisting of a single loopful of inoculum in 9 ec. of water, and then rayed in thin layers at a distance of 25 em. It is a surprising fact that none of the 5 papers cited above by Fazi in support of his results mentions yeast in any connection whatever! Fazi and Fazi (’22), in support of their previous contentions, say that tests on an industrial scale confirm their results in the 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 795 laboratory, that yeast previously treated by ultra-violet ex- hibited stimulation of fermentation activities. The possibility of commercial applications was pointed out for the third time two years later (724). This paper claimed that ultra-violet reduced the time for proper fermentation of beer 25 per cent, reduced time of maturation 40 per cent, and that it improved the flavor and storage qualities. The temperature of the fer- menting liquid might be held as low as 4? to 6” R., and the fermentation was still greater than at the usual warmer temperatures. The last paper by Fazi (727) describes the effect of raying glucose solutions before inoculation by Saccharomyces cere- visiae. Although the yeast itself was not rayed, it showed an increased fermentation and produced a smaller amount of acid. He concludes that the preliminary exposure of the glucose solution imparts bactericidal properties, which, however, do not inhibit the yeast. Fewer bacteria were found in solutions which had been rayed. Lindner (722) was one of the earliest supporters of Fazi. He used bottom brewer’s yeast and found that the velocity of fermentation was increased by ultra-violet to an even greater degree than originally claimed by Fazi for top yeast. For in- stance, 30 gms. of glucose in 300 cc. of water yielded in 24 hours 119 cc. of carbon dioxide, while a similar solution rayed with a mercury are gave 2743 cc.! The original and the rayed yeast contained but little glycogen, but signifieant amounts were present in yeast cells that were allowed to ferment in the absence of ultra-violet. This is eonsistent with the results on fermentation. A most interesting observation in this paper is that 20 to 30 per cent of the cells in the rayed solutions had died. The present writers? experiments on yeast, which are reported in the third number of this series, show that fermentation is in- hibited by ultra-violet even before a significant number of the cells show mortality. Minor variations in the substrate were shown by Lindner to be ineffectual in disturbing the results, since the use of wort gave similar effects to that of glucose. A more scientifie approach is that of Gronchi who showed [Vor. 22 196 ANNALS OF THE MISSOURI BOTANICAL GARDEN (732a, b) that the shorter wave lengths were toxic to fermenta- tion of yeast. He reported irradiating Saccharomyces cere- visiae for 45 minutes at 40 cm. Wood's light caused an in- ereased production of carbon dioxide which continued some time after the irradiation stopped. The shorter ultra-violet excited fermentation, but this increase was of short duration and stopped when the irradiation did, and from this point onward the elimination of carbon dioxide was diminished. The next attempt to show a favorable effect by ultra-violet light on yeast fermentation is that of Owen (733) and his co- workers. He used a National Carbon Company carbon arc, which is rich in the wave lengths 2300 to 3100 À. Seven grams of yeast cake were added to 100 ec. of 5 per cent glucose and exposed in a shallow pan. Exposures of only 10 or 30 seconds imparted an increased fermentative power. The greatest dif- ference between the control and experimental cultures occurred about 1 hour after irradiation. Owen reported a commercial application of irradiated yeast to bread baking, since 24 per cent less yeast is needed if it be previously rayed 10 seconds. However, itis highly probable that the quantity of yeast ordi- narily used by bakers is in excess of that needed, and the fact that 24 per cent less gives proper leavening does not prove any beneficial effect of irradiation. The most extensive report on stimulatory effects is that of Owen and Mobley (733). These workers rayed seed yeast in the same type of solution as was to be fermented, the wort itself and colonies on molasses agar. The wort was prepared from Cuban Black Strap molasses to which was added 1 ec. of sul- phurie acid and 1 gm. of ammonium-sulphate per liter. Pure cultures of Magne yeast were used. The raying was accom- plished by dripping the wort through a funnel at 15 em. from a carbon arc. In one experiment type ‘‘C’’ carbons of the National Carbon Company were used, which gave radiations from 2700 to 3000 A. The inoculated wort was run through the funnel three times before the arc. After 72 hours the efficiency of the alco- holic fermentation was as follows : 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 797 nao CA 80.62 per cent in PA 83.20 per cent o e a shade 83.10 per cent These results are not impressive evidence for stimulation. Another experiment was carried out with type ‘‘B’’ carbons, and the fluid passed 5 times through the funnel. After 72 hours the results were: Comro ny tenon 88.9 per cent e eee tenn 90.8 per cent A third experiment differed from the above in that the sus- pension was passed 6 times through the funnel. After 48 hours the efficiency of the alcoholic fermentation was: COntrO A 83.3 per cent Rayed ir ea eee ee 85.2 per cent It was further reported that raying the yeast on agar RES and then transferring it to the wort were almost as sti ti as raying the inoculated wort. Irradiation of the media was also held to be beneficial. ‘‘B’’ carbons were used, and the distance is not given. The results were: A o. t 0 ae 80.3 per cent CONTON RAA EE 82.8 per cent Colonies rayed....... 83.4 per cent Colonies rayed....... 80.3 per cent Seed rayed in wort....87.5 per cent Seed rayed in wort....84.0 per cent The scant data and the obvious variation in the duplicates are not convincing evidence for stimulation. These authors concluded with the statement that irradiation by the carbon are was beneficial because of the development of some ‘‘ growth- promoting”” substance which stimulated the fermentation of sugar. It is obvious that a ‘‘growth-promoting’’ substance is something that promotes growth and not necessarily the fer- mentation of sugar. This improper terminology is more plainly demonstrated by the values given for the actual num- [VoL. 22 798 ANNALS OF THE MISSOURI BOTANICAL GARDEN ber of cells per ec. as related to the period of irradiation. These values after 32 hours were: o 0 RE IAN? 168,000,000 Rayed 1 minute........ 116,000,000 Rayed 2 minutes....... 182,000,000 Rayed 3 minutes. ...... 48,000,000 Rayed 4 minutes. ...... 82,000,000 Rayed 5 minutes. ...... 12,000,000 Rayed 6 minutes....... 16,000,000 In spite of the erratic values, which vary from about 30 to over 100 per cent from a smooth curve, there is evidently no ‘‘growth-promoting’’ substance acting as a result of irradiation. Surányi and Vermes ('29), employing the Warburg tech- nique, were the first to observe the effect of irradiation on the use of oxygen by yeast. The nutrient medium was the Tyrade solution at pH = 7.2. Exposures to ultra-violet light gave an increase in oxygen use of about 50 per cent. Cyanide inhibited the inereased fraetion of the total as well as normal respira- tion, and this effect was reversible. Avian erythrocytes gave similar results. The evidence of the inhibitory influence of ultra-violet on fermentation is more convincing. Söhngen and Coolhaas (*23) were the first to differentiate between decrease of reproduction and of fermentative power. "They observed that neither fer- mentation nor reproduction was affected unless the distanee from the lamp was great, the volume of solution large, and the time of exposure short. Tanner and Ryder (723) extended their earlier work in view of the objections raised by Fazi. They rayed 20 different species of yeast with a Cooper-Hewitt lamp operating on 110 volts and 4 amperes, at 20 em. from the cultures. One loopful of each was inoculated in 100 cc. of sterile water, and 1 ec. of this suspension was added to 99 cc. of sterile water. The final suspension was rayed in layers 8 mm. deep in Petri dishes, after which 1 cc. was plated on acid Sabouraud's agar. Ex- 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 799 posures of 1.5 to 20 minutes were required to kill the various species. The loss in weight from 30-day cultures of Saccharo- myces ellipsoideus showed that 1- to 10-minute exposures had no effect on fermentation, but that longer exposures inhibited proportionally. Since bacilli were killed under similar condi- tions in 1.5 to 6 minutes, it was concluded that the size of the cell determines its resistance to ultra-violet. Abderhalden (727) raised the next dissenting voice. He rayed fresh yeast, dry yeast, macerated yeast, and press juice with a mercury arc and with a magnesium light. He added both rayed and unrayed egosterin and cholesterin, and under no condition did he find stimulation of fermentation. He con- cluded that fats are not concerned in yeast fermentation, and from his data it may be inferred also that ultra-violet itself was not stimulatory. Tanner and Byerley (734) reported the inhibitory effect of ultra-violet on Saccharomyces cerevisiae and on Fleisch- mann's commercial pressed yeast cake. They rayed glucose- broth eultures, aqueous suspensions before seeding, ferment- ing cultures and glucose-broth before its inoculation, with a Cooper-Hewitt mercury are at a distance of 25 cm. Oster (734), in the most careful study yet made on the sub- ject, reported the effect on respiration in terms of oxygen use. He used monochromatie light from the mereury are, including all those lines which are of sufficient intensity to be practically applied. The results were reported in terms of the wave length used, the total energy in terms of ergs per sq. mm. per second. The oxygen use was observed by the Warburg technique. The respiration eurves were run for 215 hours. At the end of the experiment, the suspension was removed and plated on agar to discover any morphological changes induced by the raying. The conclusions were that there was no stimulation of growth and no change in the oxygen use until the cells were injured as shown by morphological studies when there was only a de- crease. About 22 per cent decrease in oxygen uptake occurred when the total energy exceeded 81.6 x 10* ergs per sq. mm. per second, this value being obtained by exposure to the line 2536 À [VoL. 22 800 ANNALS OF THE MISSOURI BOTANICAL GARDEN for 60 minutes. The region 2482 Á to 3132 Á showed about the same degree of inhibition, although the total energy was 408 x 10* ergs per sq. mm. per second. The effect is not one of total energy alone but also one of individual wave lengths, since itis proportional to total energy for any one wave length, but different total energies were needed for the different wave lengths to produce a given effect. This paper is one of funda- mental importance and contains too much material to be re- viewed completely here. B. GREEN PLANTS The only study on the effects of ultra-violet light on the respiration of higher plants is that of Masure (’32). He used a Cooper-Hewitt mercury are operating on 110 volts and 4 amperes, and a Corning G586AW ‘‘Ultraglass’’ filter, 9 mm. thiek, and passing wave lengths from 3334 À to 3690 À. Be- cause of the characteristics of the mercury are, this filter showed a peak of transmission in the region 3650 Á to 3663 Á. The mereury are triplet at 3663 À, 3665 À and 3650 À consti- tuted about Y, of the total radiant energy passed. Since the line at 3650Ä was about 1% of the total intensity of the triplet, the light was practically monochromatic at 3650 Ä. This line has been shown to be within the transmission region of certain seed coats. Three varieties of pea seeds were rayed in Petri dishes for periods of 15, 30, 60, 120, and 265 minutes, and planted im- mediately after. The hypocotyls were measured after 4 days to determine the effect of raying on growth. Respiration was observed in etiolated seedlings rayed at 27 em. through the G586A W filter. The flask was in a water bath, which caused the elimination of the longer infra-red. Since window glass trans- mits down to 3200 À, the thin glass of the flask probably passed most of the band at 3650 À. The seedlings were rayed 62 minutes, then after 50 minutes they were rayed a second period of 49 minutes. The technique of measuring the oxygen uptake was not so refined as might be wished, but it appears from these experiments that the absorption of oxygen was tem- porarily increased by irradiation. 1935] ! WYND & REYNOLDS—ULTRA-VIOLET. I 801 C. THEORETICAL CONSIDERA TIONS The work of Cook and Stephenson might be interpreted as in- dicating that Bacillus coli was unaffected, but, as pointed out above, the balancing of one limb of the differential manometer against the other introduces errors that might conceal small differences. It should be noted, however, that this error need not be present if the two vessels were identical in every way. This could be determined experimentally by putting duplicate aliquots in both vessels; and should the manometer liquid re- main at a constant level, then the method of Cook and Stephen- son would be sound. But these authors specifically state that they ““assumed”” this condition. The present writers have never been able to obtain symmetric behavior of pairs of ma- nometer vessels. The work of Lohmann shows only inhibitory effects. The work on acetic fermentation is even more difficult to interpret, since no data were given that would make possible the necessary distinction between inhibiting the fermentation itself and the actual killing of the fermenting organisms. It is significant that acetic acid fermentation was not inhibited if the irradiation occurred in the absence of oxygen. This indicates that the inhibiting action does not depend on the specific effect of the ultra-violet in destroying the oxidizing mechanisms, but - rather on some toxic oxidation that ultra-violet induced by the presence of oxygen. The earlier work on yeast fermentation is open to the same objection, since no distinetion was made between fermentative activity and the killing of the cells. The evidence for the stimu- lation of yeast fermentation centers around the work of Fazi and of Owen and their co-workers. When we note that Fazi's name appears twice in the announcements of commercial patents, and three times in artieles emphasizing the desir- ability of commercial application, the actual amount of new scientific evidence advanced to support his belief is small. The work of Owen and co-workers is to be eriticized even more severely. Their actual experimental results, which have been partly quoted above, are sufficient indication that no significant differences were found. [Vor. 22 802 ANNALS OF THE MISSOURI BOTANICAL GARDEN The favorable effects of irradiating the nutritional medium reported by both Fazi and by Owen are worthy of further study. Woodrow, Bailey, and Fulmer (727) found that the ir- radiation of sucrose, glucose, calcium gluconate, and glycerol imparted toxic properties to yeast. Euler and Lindberg (12) found that rayed glucose solutions develop a gas mixture con- taining about 15 per cent carbon dioxide, 40 per cent earbon monoxide, and 40 per cent hydrogen. "They also showed that ultra-violet light anaerobically split lactic acid in water solu- tions into alcohol and carbon dioxide. These reactions are all purely photochemical, and such effects must be considered in interpreting results. The work of Bierry (*26) is particularly interesting because he also found that ultra-violet could act on carbohydrates similarly to certain enzymes, making possible an ““ultra-violet fermentation." He found that d-fructose, and to a less extent d-glucose, in the presence of air is degraded by ultra-violet to formic aldehyde and carbon dioxide. Sucrose is hydrolyzed by the irradiation and then degraded to carbon dioxide and formic aldehyde. The evidence against the stimulation of yeast fermentation is based on more stable foundations. The present writers re- port their results in the third paper of the series. There is not yet sufficient evidence in the literature concern- ing the effect of radiation on the higher plants to form definite conclusions. The present writers report in the second num- ber of this series their work on tomato and bean plants. IV. Oxipizixa ENZYMES A. CATALASE 1. In vivo studies. a. Animals.—Ostwald (’08) rayed larvae of Porthesia chrysorrhoea with sunlight and found that both violet and white light reduced the amount of catalase activity below that found in larvae treated with yellow light or with darkness. The amount of ultra-violet light reaching the or- ganisms in these experiments is very uncertain. The first in vivo study of the effects of relatively intense 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 803 ultra-violet light on animal catalase appeared in 1926. Pin- cussen, in a series of papers on the changes in the enzymatic content of the blood, points to the probability that ultra-violet affects the organism by increasing oxidation. This assump- tion was based on previously published data showing that ultra-violet exhibited an oxidative effect when studied ?n vitro. Recent developments, particularly in the chemistry of the enzymes, show that in vitro studies are not necessarily appli- cable to the interpretation of in vivo phenomena. Pineussen approached the problem by observing the catalase activity of the blood of rabbits after they had been exposed to bright sunshine from 10 o'clock a. m. to 4 o'clock p. m., and also after being irradiated by a mercury arc. Animals in the sunlight always showed a lower reaction constant of the catalase than those kept in darkness. The magnitude of this deerease was not changed materially when 2 cc. of 2 per cent erythrosin were injected subcutaneously. Neither were 10 ce. of 2 per cent potassium iodide administered orally effective in altering the effect of sunlight. Rabbits irradiated for 12 minutes by a mercury are at a distance of 100 cm. showed a lowering of the reaction constant from .0621 to 0.410. That this decrease was not a transitory effect was shown by raying rabbits 12 minutes at a distance of 90 em., and withdrawing blood samples periodically until 120 minutes had elapsed. The reaction constant showed a progressive decline as long as the observations were continued. The blood catalase of animals rayed by the mercury are was markedly more decreased in the presence of potassium iodide and eosin than in those treated with sunlight. This was ex- plained by assuming that the eosin was not acting as a photo- sensitizer for catalase, but by increasing the amount of free iodine splitting from potassium iodide. In these experiments there was also a progressive decrease during the 2 hours of observation after raying. Somewhat different results were obtained by Castagna (27a). After keeping white mice in the dark, he rayed them with the mercury arc, using both the visible rays plus the ultra- [Vor. 22 804 ANNALS OF THE MISSOURI BOTANICAL GARDEN violet and also the ultra-violet alone from 3900 Á to 3190 À. The catalase of the blood was determined at intervals of one- half hour, and it was found to inerease during the first 3 hours of raying when the open arc was used and then to drop to nor- mal after several hours. When the wave lengths 3900 to 3190 Á were used alone, there was no significant change. This author's conclusion, that visible light inereased the enzyme or protected it from destruction by some natural process, and that irradiations shorter than 3190 À were destructive, does not adequately explain these results. It should be noted that these dosages are very much greater than those used by Pineussen. It will be remembered that Harris ('25a), in his study of the effect of light on the carbon-dioxide elimination of rats, found that the open mereury arc had no effect, while the ultra-violet of 2910 A to 4360 A gave a stimulation. He concluded there- fore that visible rays nullified the ultra-violet stimulation by physiological antagonism. Koldaev and Al’Tshulla (730a) published data that is com- patible with those of Pincussen. They rayed 20 rabbits with a mercury are and observed the changes in catalase, lipase, and amylase in the blood. Single rayings gave a temporary decrease in catalase, while successive doses produced a pro- gressive decrease. The above data appeared in Russian and was apparently republished the same year in German (730b). Pineussen and Tanino (731) rayed white rats, which had been starved for 16 hours, with a mercury are operating on 220 volts at a distance of 50 em. The animals were then killed and 10 ec. of the diluted tissue extract were added to 10 ce. of 1 per cent hydrogen peroxide. The mixture was buffered at pH = 6.8 by phosphate. The results were as follows: CUBIC CENTIMETERS OF OXYGEN EVOLVED IN 5 MINUTES 1 ec. blood 1 gm. liver 1 gm. spleen 1 gm. heart Control 1427 7129 606 178 Rayed 30 minutes 1534 5659 722 182 Rayed 60 minutes 1454 5173 665 172 Rayed 90 minutes 1648 7104 700 157 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 805 The significance of these data will be discussed under the topic ‘Theoretical considerations. ’’ 1. In vivo studies. b. Plants.—In vivo studies on plants con- cerning the effect of irradiation on the catalase are less numer- ous than those on animals. The effect of sunlight on yeast catalase was observed by Euler and Laurin (18). They mixed 15 cc. of a suspension of Saccharomyces thermantitonum con- taining .0885 gms. of fresh yeast with 100 ec. of phosphate buffer at pH = 6.1 and with 50 ec. of .01 N hydrogen peroxide. Exposure of the yeast to sunlight uniformly lessened catalase activity. Novikov and Herber (733) soaked the seeds of the tau-sagyz rubber plant in water for 18 hours and then irradiated them by a mercury arc. The seeds were kept at 25? C. for 2 days and the catalase of the germinated seedlings then determined. The radiation gave an abrupt rise in eatalase. The paper by Fuller (732) is the only publication to date re- porting the effects of raying on the catalase activity of growing plants. Tomato plants and Red Kidney field beans were grown in individual 4-inch pots to a height of about 20 em. They were then rayed 3 minutes daily for 7 days at a distance of 50 em. from the arc, the rays being filtered through a layer of 1.5 em. of water in a quartz cell. Serious injury was produced, as . shown by the bronzing and curling of the leaves. At the end of the 7-day period, catalase was determined by a modification of the Appleman method. The averages showed that the catalase activity of the beans had been stimulated 43.2 per cent and that of the tomatoes 49.7 per cent. 2. In vitro studies.—Catalase was studied in its relation to ultra-violet light by in vitro studies much earlier than it was approached by i» vivo methods. It is surprising that the first of these reports was also the first one concerning catalase and ultra-violet in any connection and was carried out by mono- ehromatie radiation. Hertel ('04), in a preliminary study of several sources of ultra-violet, noted the effect of the mag- nesium line 280 ma on peroxidase and catalase. In the cata- [Vor. 22 806 ANNALS OF THE MISSOURI BOTANICAL GARDEN lase experiments, he rayed fragments of fungus mycelium, milk, and fibrin. When a drop of hydrogen peroxide was then added, the evolution of oxygen bubbles under the microscope seemed to show that the irradiation had been without effect. But when these substances were rayed in the droplets of per- oxide solution, there was a great increase in the evolution of oxygen over that in the unrayed droplets. Since drops of pure peroxide solution foamed when rayed, Hertel correctly as- sumed that his experiment did not demonstrate increased cata- lase activity. It is regrettable that his admirably arranged monochromatie irradiation could lead to no significant results because of the very poor method of determining the effect of raying. His results have been quoted by several authors as an indieation that catalase activity was increased. A reading of the original “paper shows this to be in error. Catalase preparations from Dytiscus marginalis were ex- posed to sunlight by Ostwald (708). The activity was deter- mined by titrating the hydrogen peroxide that remained in the test preparation after the lapse of a definite period of time. The enzyme from the larvae exposed to sun used up 94.7 per cent of the available hydrogen peroxide, and the control de- stroyed 96.8 per cent. While these values show a slight injury to the catalase, it is probable that they are insignificant. The present writers have found that the titration of duplicate aliquots of eatalase by the method of Ostwald gives very erratie results unless the amount of peroxide added as a sub- strate is sufficiently great to insure at least 15 per cent of the original amount still present at the time of titration. Catalase from rabbit and from human blood was found by Lockemann, Thies, and Wichern (’08) to be greatly injured by 2-hour exposures to strong sunlight. The preparations were exposed while they were reacting with the hydrogen peroxide substrate. Controls containing peroxide alone allowed a cor- rection to be made for the amount of peroxide destroyed by sunlight itself. The relation of photosensitizers to catalase was studied par- tieularly by Zeller and Jodlbauer ('08). An exposure of 25 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 807 minutes to sunlight filtered through water injured the catalase activity 68 per cent if oxygen were present. In the absence of oxygen there was only 10 per cent injury. Exposure to a mer- eury are for 30 minutes showed 29 per cent injury if oxygen were present and 25 per cent if absent. These figures show that oxygen is necessary for the injurious action of the visible light and is unnecessary for the ultra-violet. Increasing alka- linity was found to increase the injurious effects of the sunlight, while corresponding increases of acidity up to 1/1500 N sulphuric acid were without effect. The sunlight was increased in its destructive action by 1/5000 N eosin only if the light were passed through a glass plate to remove the short ultra-violet. The eosin had no influence on the irradiation from the mercury are unless a glass plate was interposed as a filter. Essentially the same results were obtained by the use of sodium-dichlor- anthracene, methylene-blue, and Bengal-rose. The injurious waves of the emission spectrum of the fluo- rescent substances might depend on the nature of the incident light, but why this should be prevented by the mere additional presence of ultra-violet is difficult to explain. It would seem that the destruction of both types of rays should be additive. If, however, the above results are authentic they furnish an in- teresting instance of antagonism. It is not possible to surmise whether this antagonism is one of physical interference or of physiological reaction. Some phases of the work of Zeller and Jodlbauer were criti- cized by Battelli and Stern (10). They found that the destruc- tion of catalase by the visible sun rays proceeded equally fast in the presence or absence of oxygen. Although the sun can rapidly and completely destroy catalase, a protective action is exerted by traces of alcohol, aldehyde, and certain other chem- ical agents. Agulhon (’11) studied the behavior of a number of enzymes to the radiation from a Heraeus mercury are operating on 110 volts and 2 to 3 amperes. The enzyme preparations were rayed in layers about 1 mm. deep in quartz tubes at a distance of 15 to 20 cm. The activity of catalase obtained from pork fat was [VoL. 22 808 ANNALS OF THE MISSOURI BOTANICAL GARDEN reduced to 83 per cent of the control by an exposure of 1 hour, to 39 per cent in 3 hours, and to 8 per cent in 6.5 hours. A more complete report was published by Agulhon (712) the following year. He found, contrary to Battelli and Stern, that catalase from pork liver was destroyed in the absence or the presence of oxygen by visible light, although to a less extent in vacuum, and, in eontrast with Zeller and Jodlbauer, he found that the destruetion by ultra-violet was notably lessened in vacuum. These inconsistencies were explained by the probable traces of oxygen in evacuated tubes which might be ample to effect the destructive oxidation of the comparatively minute traces of actual enzyme material. Waentig and Steche (*12) rayed with a mercury are catalase prepared from fat, liver, blood serum, alcohol, precipitate of blood, larvae of Sphinx ligustris, pupae of S. ligustris, and from fungi. In all cases the catalase activity was lowered. The injury was small when the raying took place in glass tubes. In some instances, nitrogen was bubbled through the solution for a long time to eliminate the oxygen, but injury was not pre- vented. It was concluded that the injury was not oxidative, but probably an effect on the solubility of the enzyme, since more or less precipitate accompanied the destruction of the enzyme. $Sinee alkaline solutions increased the injury while acid did not, these authors concluded that probably all catalase reacted the same. Zeller and Jodlbauer found similar results with fat and blood catalase. The remarkable stability of the catalase activity of red blood corpuscles was demonstrated by Peemóller and Franke (726). They suspended washed human blood corpuscles in a .9 per cent sodium ehloride solution. One cc. of the suspension was added to 10 ce. of 3 per cent hydrogen peroxide, and aliquots of 1.5 to 2.0 ce. were rayed in thin layers at a distance of 50 em. Under these eonditions an 8-minute exposure gave 1 HED (Haut- erythemdosis) unit. Exposures up to 10 HED units (80 minutes) did not produce any significant injury. It is important to distinguish in studies on the catalase ac- tivity of blood between the activity of hemoglobin and that of the blood catalase proper. The great and unexpected stability 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 809 of the catalytic action of ‘‘Blutkatalase’’ of Peemöller and Franke is not concerned with the true enzymatic activity. This distinetion was taken into consideration by Pincussen and Seligsohn (726). These authors point out that errors due to the activity of hemoglobin can be lessened by proper buffering, since the optimum pH for blood catalase differs from that of hemoglobin. The experimental preparations of Peemóller and Franke were obtained by adding .02 ce. of blood to 50 cc. of water. Aliquots of 5 cc. of this dilution were mixed with 100-ec. por- tions of 1/2000 hydrogen peroxide. The reaction was stopped by adding 2 ce. of 20 per cent sulphurie acid. The remaining peroxide was titrated with N/100 potassium permanganate. Aliquots of 25 cc. were removed for titration after 6, 15, and 20 minutes. Irradiation for 20 minutes at a distance of 10 centimeters greatly injured the catalase. This injury was -greatest at the pH that is optimum for catalase activity, and was also proportional to the degree of dilution. The presence of potassium iodide and sodium chloride exerted some pro- tective action against destruetion of the enzymatie activity by enlarging the colloidal particles sensitive to light. Sodium chloride was itself injurious in addition to its effect on the light injury. In the presence of this salt, about 50 per cent of the activity was lost on raying for 10 minutes, while in its absence the injury was 25 per cent. The action of potassium iodide is complicated. There is an injury due to the splitting off of free iodine, but in an excess of the salt the protective ac- tion of the salt itself exceeds this injurious effect. It is im- possible in this instance to clearly distinguish the effect of the salt as a protector, the injurious effect of the halogen ion, and the injurious effect of the liberated iodine. The toxicity of liberated iodine can be demonstrated by raying catalase with alival. Blood from dogs was exposed in quartz tubes to ultra-violet by Castagna (’27b), and the catalase was progressively in- activated, becoming almost zero after 11% hours. Sunlight gave a very slight diminution. Morgulis (729), working with kidney catalase at various hy- [VoL. 22 810 ANNALS OF THE MISSOURI BOTANICAL GARDEN drogen-ion concentrations, found complete inactivation at all degrees of acidity, but the required time varied. At pH - 6.0, 7.0, and 8.0 the inactivation curves were straight lines as plotted against the time of exposure. At other acidities the inactivation was faster at first, but became slower as the time of exposure increased. The minimum inactivation occurred at pH - 6.0. This relation to pH was studied further by Morgulis (730). He disagreed with the opinion of Pincussen that catalase was most injured at its optimum pH =6.8. Using a Hanovian mer- eury are, operating on 110 volts and 5 amperes at a distance of 25 or 50 em., he was able to show that the greatest injury oc- eurred between pH =8.0 and 9.0, which is on the alkaline side of the optimum. The stability of his preparation was greatest toward ultra-violet and heat at pH = 6.0, which is on the acid side of the optimum. This pH is suggested as the probable iso- electric point. This paper contains excellent graphs relating. activity to pH and time of exposure. 3. Theoretical considerations.—The literature contains a surprisingly great amount of trivial discussion concerning the significance of catalase in tissues. The accumulated authentic data are of permanent value, but their significance is not well understood. A very important statement was published by Pineussen and Tanino (731), calling attention to two diametri- eally opposed interpretations of the relation of catalase to respiration. In their study they found a decrease of liver cata- lase on irradiating the live rats. Assuming that catalase develops proportionally as it is needed to destroy excess hy- drogen peroxide, this decrease may be interpreted either as due to an increased oxidation in which hydrogen peroxide is used or to a lessened production of the peroxide by some de- creased respirational process. The paper by Ranjan and Mallik (’31) is one of the most im- portant contributions on the significance of catalase. They show very convincingly that catalase activity parallels the formation of hexose, whether this formation results from hydrolysis of disaccharides or from photosynthetic activity. 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 811 Wynd (734) has suggested that this parallelism may signify a transitory peroxide group during the mobilization of the hex- ose substrate for respiration, and that some of this peroxide is available to peroxidase. Ostwald (708), finding that sunlight decreased catalase of larvae when rayed both in vivo and in vitro, and also inciden- tally that peroxidase was increased under similar conditions, concluded that these enzymes were connected with photot- ropism. He also observed that the rayed larvae increased in weight over the controls, but he wisely admitted that these phenomena need not be related necessarily. The statement of Lockemann, Thies, and Wichern ('08), that the effects of high mountain sun and of sunbaths in general were connected with the depression of the catalase, was made on the unwarranted assumption that in vivo effects were sim- ilar to those obtained in their i» vitro raying of blood. The eorrelation of many apparent discrepancies in the behavior of catalase to irradiation in in vivo studies necessitates a consid- eration of the relative dosages. From the available data it ap- pears that heavy and therefore toxie doses increase catalase of animal tissues while lesser dosages are depressing. The data supporting this generalization are yet comparatively meager and need further verification. Both of the in vivo plant studies show inereased eatalase. In one instance the rayed seeds produced more vigorous growth, and in the other the growing plants were rayed until they ex- hibited definite injury. Further data concerning this problem are included in the second paper of this series. In the in vitro studies on animals it is particularly important to separate the catalase activity of hemoglobin and that of the true enzymes, as has been adequately pointed out by Pincussen and Seligsohn (726). The widely different degrees of injury of catalase solutions by sunlight and by the mercury arc need not be considered incompatible since many known factors, such as dilution, presence of other substances, light intensity, ete., exert powerful influences. It is clear that catalase is injured by light when irradiated in vitro. [Vor. 22 812 ANNALS OF THE MISSOURI BOTANICAL GARDEN B. OXYGENASE 1. Animals.—The general destructive action of ultra-violet on oxygenase of animal origin was reported by Vedder (’24). He found that temperatures greater than 62° C. caused the disappearance of the oxidase-containing granules of leuco- cytes, but ultra-violet light caused their disappearance in the absence of any temperature change. The most recent contribution on this subject is that of Wohlgemuth and Sugihara (’25). It had previously been pointed out by Wohlgemuth and Yamasaki (’24) and by Yamasaki (’24) that skin is rich in the enzymes, diastase, lipase, catalase, and oxygenase (phenolase). This oxygenase could oxidize the dihydroxy derivatives of adrenalin, dioxyphenyl- alanine, and pyro-catechol. Wohlgemuth and Sugihara studied the diastase, lipase, gelatinase, and oxygenase in the skin of different animals and found that oxygenase was highest in the rat and frog. In general, all the animals exhibited greater enzymatic activity of the skin than did man. There was no parallel between the enzymatic content of the blood and that of the skin. Irradiation of guinea pigs over a sufficiently long period with natural sun, artificial sun, and artificial ultra-sun caused a considerable increase of oxygenase, and a decrease of diastase and lipase. In no animal was there any change in the aetivity of these enzymes in the blood. 2. Plants.—Agulhon (11) made the observation that the oxygenase (laccase) extracted in glycerin from the fungus Russula was extremely stable towards irradiation. Even 6 hours of raying by a Heraeus mercury are at a distance of 15 to 20 em. only rarely produced a detectable lessening of its subsequent action on 1 per cent guaiacol. However, if the glycerin extract were diluted with 4 parts of water then oxy- genase was notably destroyed in 3 hours of irradiation. Wave lengths longer than 3022 À were found to be almost inactive. The following year (712) he published a more extended re- port. 'The method of raying was similar to that of his earlier work, and from the identity of the numerical data it may be 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 813 inferred that the previously obtained data were the basis of his later paper. Besides restating the observations of his earlier paper, this publication describes a possible mechanism for the action of the ultra-violet rays. Since oxygenase is not attacked in a glycerin extract in vacuum, and but a slight decrease becomes apparent when the glycerin extract is diluted with water, it is therefore probable that the destructive action is one of oxidation. It is known that ultra-violet light decomposes water to a slight degree, liberating hydrogen. In the presence of oxygen, this hydrogen forms hydrogen peroxide, which injures the enzyme. Experiments showed that .5 per cent hydrogen per- oxide destroys completely the oxygenase of Russula extracts in 4 hours. Visible light was thought to owe its destructive action to the same mechanism, although in this instance actual air was necessary, while with the ultra-violet the slight traces of oxy- gen unavoidably present in vacuum sufficed. 3. Theoretical considerations.—Because of the small amount of work done on the behavior of the oxygenases to ultra-violet, it is diffieult to arrive at definite conclusions. If the interpre- tation of Agulhon is verified, then it forms a significant basis upon which we may interpret various of the activities of ultra- violet light. The authenticity of the formation of hydrogen peroxide by the action of ultra-violet on water is undoubted, but whether this could act destructively on the oxygenases is a question, although the fact that .5 per cent hydrogen peroxide completely inhibited Russula oxygenase in 4 hours cannot be ignored. More discussion of the problem will follow in the theoretical consideration of peroxidase. The work of Wohlgemuth and Sugihara is suggestive, since it is not impossible that the oxygenase increase in the skin of rayed animals might play a role in the development of melanin in spite of the fact that the specifie enzyme preparation used in these experiments did not affect tyrosine. The distinction between oxygenase and the tyrosinase may be assailed on theo- retical grounds, and the entire problem of the nature of the ac- [VoL. 22 814 ANNALS OF THE MISSOURI BOTANICAL GARDEN tivity of oxygenase and tyrosinase is not as firmly supported as many writers suppose. A review of this interesting problem is not appropriate in the present paper. C. TYROSINASE 1. Animals.—Narayanamurti and Ramaswami (729, 30) ir- radiated the larvae of Dolichos and found consistently in- creased tyrosinase activity of extracts. It was shown by Lignac (30) that human skin, after death, would develop melanin as a result of raying with a Kromayer mercury arc. Several hours of exposure of dead skin were needed to produce the same degree of pigmentation as an ex- posure of 4 minutes of live skin. The difference was attributed to the constant oxygen supply of live skin. One might assume very reasonably that the inerease of melanin in skin after ray- ing is due to an increase of the activity of tyrosinase. Pin- cussen and Hammerich (731a), acting on this supposition, mod- ified the method of Raper and Wormall for the estimation of tyrosinase, so that it could be used as a miero method. En- zymes prepared from meal worms were rayed in vitro with a mercury arc. Tyrosinase, as most other enzymes, was greatly injured, and they concluded that the formation of melanin in the skin by ultra-violet was not due to an increase of tyrosinase. 2. Plants.—The papers of Agulhon ('11, ’12), mentioned above, state that the extracts from Russula contain tyrosinase and that this enzyme is more stable towards irradiation than is oxygenase. Like oxygenase, it was not destroyed by ultra- violet in glycerin in vaeuum, and only slightly injured when rayed in a water solution in vacuum. Likewise, it was inacti- vated in 4 hours by .5 per cent hydrogen peroxide. It was thought to be destroyed by the same mechanism as suggested for oxygenase. 3. Theoretical considerations.—The conclusions of Pineus- sen and Hammerich may be attacked on several grounds. In the first place, as has been clearly demonstrated by Fuller (32), the behavior of an enzyme under in vitro raying bears no relation to its behavior when rayed in vivo. Moreover, the 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 815 total exposure, 1 hour at 50 cm., might well have obscured the effect of less toxic exposures. But the most arresting fact of all is that these authors used dioxyphenylalanine, as well as tyrosine, as a substrate, with identical results. Wohlgemuth and Sugihara found that dioxyphenylalanine was oxidized more by the skin oxidase (called phenolase by them) extracted from guinea pigs which had been rayed while living with a mercury are. Since enzymes are defined in terms of their sub- strates, it may be assumed that whatever enzymes actually at- tack this particular substance, they reacted differently to in vivo and in vitro irradiation. These authors did not, however, obtain any oxidation of tyrosine by their skin extracts, and yet we know that tyrosinase should have been present in the frac- tion which they called **phenolase.'" As mentioned above, the oxygenase and tyrosinase reactions are surprisingly complex and confused in the literature. It seems reasonable to conclude that tyrosinase, as most en- zymes, is injured by irradiation in vitro, but that its behavior in vivo to exposures has not yet been reported authentically. D. PEROXIDASE 1. Animals. a. Milk.—The paper of Hertel (704), already cited as the earliest study on the effect of ultra-violet on cata- lase, occupies a similar position in respect to peroxidase. Milk peroxidase was studied by mixing 2 cc. of fresh milk with 2 ce. of .1 per cent aqueous solution of guaiacol and 1 drop of hydro- gen peroxide. Irradiating the milk with the 2800 À magnesium line produced a notable diminution of activity even with as brief exposures as 5 minutes. Römer and Sames (’10) materially extended the observa- tions of Hertel. They used 5 ec. of cow's milk mixed with 2 drops of guaiacum tincture and 6 drops of 2 per cent hydrogen peroxide. If the milk were rayed with a Heraeus mercury are at a distance of 15 em. for 1 hour, the peroxidase activity was completely inhibited. Exposures of 14 hour allowed a positive test to appear, although less than the control. When milk whey was rayed, a trace of peroxidase activity remained after 1 hour of exposure, but 114 hours were completely inhibitory. These [Vor. 22 816 ANNALS OF THE MISSOURI BOTANICAL GARDEN authors state that this destruction of the peroxidase (called ““oxidase”” by them) could have no hygienic significance in infant feeding, since human milk normally contains no peroxidase. In view of the great similarities between cow's milk and human milk, it seems strange that such a difference could exist in this particular. Asakura (732) has shown that human milk lacks peroxidase if it also lacks vitamin B. Heavy feeding of vitamin B causes the peroxide reaction to appear. Reinle (21) extended the study of the peroxidase of cow's milk to include the effect of Róntgen rays, the y-rays from radium chloride, and ultra-violet. The X-rays and y-rays had no effect. When the milk was rayed with a mercury arc oper- ating on 110 volts and 4 amperes at a distance of 15 cm., there was a temporary increase for the first 20 to 30 minutes. This stimulation is not significant since mere warming of the milk will also give this effect. This apparent stimulation could not be due to the destruction of the dehydrogenase system by the irradiation, since, as will be shown later, these enzymes are notably stable towards ultra-violet. The effect of warming could very well depend, on the other hand, on the inhibition of dehydrogenating systems. Such an effect has been suggested by Keilin (729) to account for the fact that fresh baker’s yeast exhibits no peroxidase test while warmed yeast does. 1. Animals. b. Tissue.—' The relationship between animal tropisms and the light-sensitive enzymes was studied exten- sively by Ostwald ('08). He found that strong sunlight in- creased the peroxidase activity of extracts from the larvae of Porthesia chrysorrhoea. Irradiation of the living larvae also produced the inerease, from which it was concluded that pho- totropism was connected with cellular respiration. The per- oxidase activity of irradiated Paramecia was studied by Roskin and Schischliajewa ('34), but no conclusive results were obtained. 2. Plants. a. In vivo studies.— There have been no data re- ported on the effect of raying living plants on the activity of 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 817 peroxidase. The authors report studies of this problem in the next number of this series. 2. Plants. b. In vitro studies.—The effects of strong sun- light were reported first by Bach (708). The preparations were exposed in Erlenmeyer flasks, and hence only the longer ultra-violet reached the solution. The exposures lasted from 4 to 76 hours, with a progressive decrease in the activity of the enzyme. Jamada and Jodlbauer (708), in an extensive study on the effect of irradiating peroxidase of vegetable origin, reported that horseradish peroxidase was injured by 15-minute ex- posures to sunlight even when the light was filtered through 20 em. of water to remove the infra-red. The injurious effect of ultra-violet was shown by the fact that the activity of the enzyme was lessened more by 25-minute exposures to sunlight filtered through quartz than through ordinary glass. Expo- sures to a mereury are with 20 em. of water used as a filter were made. When ordinary glass was interposed there was some injury, but when a quartz plate was substituted the injury was very marked. It was also shown that the visible portion of the spectrum, either from the sun or from a mercury are, had no effect in the absence of oxygen. The rays from an unsereened mereury are gave great injury in absence of oxygen, which was a little inereased if oxygen were present. From these data the authors concluded that ultra-violet exerted its injurious effect by a fundamentally different mechanism than did the visible light. If eosin were present in concentrations of 1/2000 to 1/5000 N, the peroxidase was sensitized notably towards the visible light. The results after 75 minutes were: Bata D tals an 0 per cent injury Rayed Oye terse oe 41 per cent injury Rayed with eosin... .58 per cent injury This paper contains excellent data illustrating the fact that fluorescent substances may act as sensitizers by changing in- cident light to more harmful wave lengths, or they may act as protectors by degrading toxic to harmless wave lengths. Thus, [Vor. 22 818 ANNALS OF THE MISSOURI BOTANICAL GARDEN in the present study, eosin and Bengal-red (1/10,000 N) sensi- tized peroxidase to the visible light, when methylene-blue and the sodium salt of dichlorantl disulphurie acid pro- tected the enzyme. In the case of methylene-blue, the injury by visible light was lessened 50 per cent. Eosin in the ultra- violet had no sensitizing action. In the absence of the visible rays, it had a protective influence against ultra-violet, since it absorbed some of the toxic waves and rendered them harmless. In the absence of oxygen the visible could not act, hence eosin also protected. The fact that Karamitsas ('07) found that eosin did not significantly sensitize peroxidase may be due to his use of thinner glass tubes. If the ultra-violet were transmitted, then eosin would exert a protective action toward these waves which might cancel the added destructive power of the visible, thereby giving the false appearance of inaction on the part of the fluorescent compound. - Bering ('12) reported a stimulation of horseradish peroxi- dase by irradiation. Exposure to strong sunlight in dosages of .7 to 1.0 **Finsen"' units stimulated the subsequent forma- tion of purpurogallin about 6 per cent. The unscreened mer- cury are gave only inhibition, but when filtered through a rabbit's skin the rays stimulated peroxidase again about 6 per cent. Experiments with filters showed that the red region of the spectrum had no effect; the yellow and also the green regions were stimulative; the blue plus the long ultra-violet stimulated about 12 per cent. The author concluded that those rays which penetrate the skin stimulate the respiratory en- zymes and therefore are physiologically important. Later Bering and Meyer (712a, b) verified these results, and pointed out that ultra-violet filtered through rabbit's skin stimulated or inhibited peroxidase, depending on the concentration of the enzyme. The unfiltered sunlight stimulated when the exposure was short. Capelli (26) rayed a purified vegetable peroxidase and found the mercury are injurious. The solutions developed ozone which may have caused an initial apparent increase in 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 819 oxidative power. During this same year, Meyer (726) pub- lished a review in which he restated the belief that light acti- vated the oxidizing enzymes, and was of therapeutie value on that account. New data were given verifying the stimulation of peroxidase by exposure to the mercury are, but only when the shorter irradiations were removed by a rabbit's-skin filter. Pincussen and Hammerich ('31b) prepared a very concen- trated horseradish peroxidase according to the method of Will- státter. They mixed .4 ec. of the preparation with .2 em. of .5 per cent hydrogen peroxide, and then diluted to 400 cc. with water containing 1 gm. of pyrogallol. Aliquots of 20 cc. were removed for the determination of purpurogallin. The reaction was stopped by adding 10 cc. of 10 per cent sulphuric acid. The color was extracted by 20 ec. of ether, and the ether solution compared colorimetrically with a .57 per cent solution of chromic acid. The enzyme solutions were diluted with 5 parts of water. Acetate buffers were used to obtain pH = 3.4, 4.0, 4.6, 5.2, 6.0, and 6.4. The exposures lasted 30 and 60 minutes. The light source was a ‘‘ Vitaluxlampe’’ mecury arc with a glass cell to filter out the heat rays. The enzyme was greatly injured by irradiation. The pH had no great influence on the degree of injury, although there was a greater diminution of activity when the solutions were acid. 3. Theoretical considerations.—The effect of raying on per- oxidase is subject to several sources of error. The ability of ultra-violet rays to produce hydrogen peroxide in water has been demonstrated by a number of students among whom we may mention Tian (’11). If sub-optimum concentrations of peroxide are used in experiments, this additional substrate for the enzyme may cause an increase in oxidation. This is especially true of oxygenase, in which the slight amount of peroxide formed would enable the otherwise inactive peroxi- dase to act, obscuring the activity of oxygenase itself. Per- oxides may also develop when oils are irradiated, as has been worked out by Delore (729) and others. These peroxides not only act as an additional source of substrate for the enzyme, but also cooperate with ultra-violet on their own account to ac- [Vor. 22 820 ANNALS OF THE MISSOURI BOTANICAL GARDEN complish oxidation of the dye indicator. The peroxide also is destroyed by ultra-violet, and Koeppe (’30) has shown that the liberated oxygen is in the nascent condition and easily oxidizes guaiacol. The oxygen that develops when hydrogen peroxide is decomposed by heat does not oxidize guaiacol. From Koeppe's data, it is seen that false results may be obtained if the enzyme is irradiated in the presence of the indicator. The influence of ultra-violet on tautomerie equilibria is well known. It is not improbable that substances which exist in a keto-peroxide tautomeric equilibrium may have oxidative powers due to this fact when affected by irradiation. The ex- istence of an enol and a keto-peroxide form in cholesterin is particularly significant, since Remesow (732) has shown that this keto form interferes, because of its oxidative power, with the conventional Nadi test for oxygenases. Pincussen (730), in the most adequate discussion that has ap- peared concerning the physiological effects of ultra-violet, has pointed out that the mechanism of color formation in the per- oxidase tests is very complex, and that the observed stimula- tion may not be connected with the enzyme itself. From the discussion above, it is seen that this is indeed probable. The few instances of stimulated peroxidase activity result- ing from irradiation are probably due to some of the disturbing factors discussed above. The theory that light, particularly ultra-violet, owes any of its physiological effect to stimulated oxidizing enzymes is to be disregarded. It is not known whether the thermo-stable peroxidases react differently than the labile type. E. DEH Y SE 1. Animal tissues.—Krestownikoff (’27) prepared a dehy- drogenase solution from horse muscle, using potassium suc- cinate as the hydrogen donater. Evacuated tubes containing the enzyme-succinate-methylene blue mixture were rayed in a water bath by a carbon are. Since glass tubes were used, only the longer ultra-violet reached the solutions. Exposure to the carbon are stimulated the rate of reduction of the methylene- 1935] WYND € REYNOLDS—ULTRA-VIOLET. I - 821 blue. Control solutions containing no enzyme showed no re- ducing power. Junghagen (728) found that the succinic-acid and glycero- phosphoric-acid dehydrogenases extracted from horse muscle were rapidly destroyed by ultra-violet irradiation. Bertrac- eini (729) found that dehydrogenase of animal origin, as studied by the reduction of m-dinitrobenzol, was increased by small exposures to ultra-violet, but depressed by larger doses. The work of Palmieri (733) was more extensive. He rayed rabbit eyes for various periods, and then studied the reducing power of the lenses by Thunberg's method. Immediately after raying, there was a slight inerease in the rate of reduction of methylene-blue, followed by a decrease, and finally, 3 days after the irradiation there was a progressive increase. 2. Plants. a. Bacteria.—The only data concerning the be- havior of bacterial dehydrogenases depend on inferences drawn from the work on Bacillus coli, already quoted, by Cook and Stephenson (728). These authors studied the relation of the viability of the cell to its ability to oxidize aerobically glu- cose and its fermentation products. Cell suspensions exposed 20 minutes to a mercury are contained but 2.74 living cells per 1000, and exposures of 30 minutes killed all but 2 per 1000. The substrates used were glucose, acetate, lactate, and formate. When glucose was the substrate, the cells rayed 20 minutes showed only about one-half as much oxygen use after a period of 30 minutes, but after one hour the total oxygen use was about equal to the control. The same was true of lactate and acetate, and only in the case of formate was an undoubted inhibition of oxygen use evidenced. The authors concluded, as has been mentioned above, that no significant difference was found be- tween the oxidation of dead and living cells. 1t should be men- tioned, however, that the objections to their technique, already described, are pertinent also to this interpretation. 2b. Higher Plants.—The dehydrogenases of plants have not been as well studied as those of animals, and therefore the extensive paper by Lehmann (722) is of particular importance. [Vor. 22 822 ANNALS OF THE MISSOURI BOTANICAL GARDEN The seeds of Phaseolus vulgaris were ground very fine and .05 gm. plaeed in Thunberg tubes. Besides the meal, each tube received .1 ec. of 1/5000 methylene-blue, .1 cc. of 1/10 N potas- sium biphosphate as a buffer, .5 cc. of potassium malate as a hydrogen donator. Water was added to give a total of 1 cc. The potassium malate solution was made by dissolving 346 mgms. in 10 cc. of water. The tubes were evacuated and the time required for the reduction of the methylene-blue recorded. The dry powder was rayed with a carbon are at a distance of 40 em., with the following results: 4 7 1 PTS NETTE required 21 minutes for reduction Rayed 100 minutes. ...required 19 minutes for reduction Rayed 240 minutes... .required 43 minutes for reduction It is significant, as will be pointed out later, that a stimulation oceurred when the sample was rayed 100 minutes. 3. Xanthine dehydrogenase.—Some authors consider the aldehyde oxidizing system in milk to be catalyzed by an enzyme distinct from that concerned in the oxidation of xanthine. It seems logical that substrates differing from each other as greatly as do aldehyde and xanthine should be activated by dif- ferent enzymes. Most writers believe, however, that a single enzymic system is concerned. Romer and Sames (710), in the paper referred to above, re- ported the effect of raying cow’s milk on the dehydrogenase (reductase). They used methylene-blue as an indicator in the presence of formaldehyde. Exposures to a Heraeus mercury are at a distance of 15 em. for various periods gave a small in- crease at shorter exposures and then a decrease, although even 2 hours of irradiation did not completely destroy the enzyme, as is shown by the table: Control ........... reduced methylene-blue in 6 minutes Rayed 15 hour...... reduced methylene-blue in 4 minutes Rayed 1 hour....... reduced methylene-blue in 6 minutes Rayed 2 hours...... reduced methylene-blue in 30 minutes 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 823 This same reaction was studied by Reinle (721) and the re- sistance of the dehydrogenase to ultra-violet verified. His ab- breviated results are: Fresh milk....... reduced methylene-blue in 715 minutes Rayed 10 minutes. reduced methylene-bluein 8 minutes Rayed 30 minutes. reduced methylene-blue in 12 minutes Rayed 1 hour..... reduced methylene-bluein 15 minutes Rayed 2 hours... .reduced methylene-blue in 1915 minutes Rayed 3 hours. . . .reduced methylene-bluein 24 minutes Rayed 4 hours....reduced methylene-blue in 31 minutes Injurious effects of the same order of magnitude were again obtained by Pineussen and Oya (729). They mixed 1 part fresh cow's milk with 1 part of M/3 phosphate buffer and 1 part of water, and added 5 cc. of this mixture to .3 cc. of a methylene- blue solution. The methylene-blue solution was prepared by mixing 5 cc. of saturated alcoholic methylene-blue with 5 ec. of formalin, and diluting to 200 ce. with water. The milk-buffer mixture was rayed in thin layers at a distance of 20 cm. from a mercury are for 1 hour. The dehydrase (aldehydrase) was weakened about 50 per cent. Bernheim and Dixon (728) studied the effect of light on xan- thine oxidase of fresh milk, the easeinogen preparation of Dixon and Thurlow, and on the whey preparation of Dixon and Kodama. Hypoxanthine or aldehyde were the hydrogen do- nators, and methylene-blue or nitrate were the hydrogen ac- ceptors. The reduction of the methylene-blue was followed in Thunberg tubes. Moderate irradiation by a carbon are or by a 100-watt filament bulb gave great stimulation when traces of oxygen were present. Long exposures were inhibitory. Both of these effects were found to be due to the formation of traces of hydrogen peroxide. The first stage of oxidation of the oxi- dase by the peroxide caused an increase in its activity, while its further oxidation lessened its activity. This interpretation is made probable by the fact that 1/100,000,000 M hydrogen per- oxide gave stimulation. Higher concentrations than these destroy the enzyme. The intensity of the effect depends on the amount of enzyme present. [VoL. 22 824 ANNALS OF THE MISSOURI BOTANICAL GARDEN 4. Theoretical considerations.—It is interesting to note that small increases of dehydrogenating activity, when the period of irradiation was short, were observed in milk by Rómer and Sames and by Krestownikoff, and in animal tissues by Bertrac- cini, by Krestownikoff, and by Palmieri. Despite the few published papers on the subject, it is evident that dehydrogenases are particularly resistant to ultra-violet destruction. It would be interesting to compare the resistance of the oxidases under similar in vivo conditions. The work on xanthine dehydrogenase (known more commonly as ‘‘xanthine oxidase") furnishes the most convincing evidence at hand of the stimulation of an oxidizing enzyme by ultra-violet light. The fact that methylene-blue can aet as the hydrogen acceptor in the absence of oxygen, and that formaldehyde ean be sub- stituted for xanthine or hypoxanthine as the hydrogen donator suggests an explanation for the several instances of dehydro- genase stimulation deseribed above. In many of these ex- periments, the preparations, particularly those of indefinite composition, may have been exhibiting the typical xanthine- oxidase reaction rather than that of the supposed dehydro- genases. This seems the more probable when it is recalled that . xanthine and its derivatives, and xanthine- activating systems, are widely distributed in nature. Meldrum (733) has summarized the characteristies of this interesting enzyme. He points out that it differs from the usual dehydrogenases in that the dehydrogenase inhibitors, such as vanillin, urethane, diethylurea, propionirile, etc., do not lessen its activity, nor is it inhibited by hydrogen cyanide or hydrogen sulphide. It contains neither eytochrome nor an oxidase. From these facts it is seen that the enzyme is funda- mentally different from dehydrogenase of the succinate de- hydrogenase type. V. SuLPHYDRYL COMPOUNDS A. EXPERIMENTAL The first reversibly oxidizable substance of known composi- tion to be isolated from tissues was glutathione. In later years 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 825 it has assumed great importance in respiration chemistry, although eomparatively few students have studied it in con- nection with the problems of ultra-violet biology. Glutathione bears an intimate relationship to many enzyme systems, and it has also been considered to be a factor con- cerned in the proliferation of cancer tissue. In view of the ef- fect of raying cancer tissue with ultra-violet, Woodward (733) irradiated glutathione in pure solution. She used a Uviare at a distance of 25 cm., and irradiated the solution in thin layers on an ice bath to eliminate destruction by the heat. The destruc- tion of the glutathione, as well as the oxidation of sulphydryl group generally, was found to depend on the pH, on the time of exposure, and on the concentration of the sulphydryl com- pound. At pH - 7.0, the oxidation of the sulphydryl compound to the disulphide proceeded more rapidly than destruction, and this process is therefore probably a primary step in the de- structive effect of ultra-violet on glutathione. The destruction seemed to be a progressive oxidizing effect rather than the lib- eration of inorganie sulphide or free sulphur. Bersin (733) based his study of the sulphydryl compounds on the fact that papain had been shown to be activated by hy- drogen sulphide or by such thiol compounds as eystein or glutathione. This activation involves a reduction. In opposi- tion to Woodward, Szendro had shown that cystein developed from cystine when exposed to ultra-violet light. The work of Szendro was repeated by Bersin (733) and found to be correct. He rayed the substance, HOOC.CH (CH3).S.S.C (CH3). COOH and found that it went to the reduced form. This experiment took place in a hydrogen atmosphere, while Woodward ir- radiated her solution in air. Bersin assumed that papain was a disulphide, PaSSPa, which became activated when reduced to the PaSH form by irradiation in hydrogen. Pineussen and Takahashi (733) removed the organs from animals 30 minutes after raying them and determined their gluthathione content. These authors irradiated with infra- red, blue, orange-red, and yellow. The latter was obtained by a sodium-vapor lamp and a significant amount of ultra-violet [Vor. 22 826 ANNALS OF THE MISSOURI BOTANICAL GARDEN was present. The sodium lamp produced in general a small inerease in the glutathione content, with no significant dif- ference when the ultra-violet was removed by filtration. Wels and Jokiseh (733), Wels (733), and Wels and Hesse (734) have shown that egg albumen when rayed in nitrogen with ultra-violet developed a reversibly oxidizable body, prob- ably a sulphydryl eompound. It is very probable that irradia- tion of living tissue produces some similar result which eannot but be of importance to the respiratory processes. B. THEORETICAL CONSIDERA TIONS The apparently contradictory results of Woodward and Bersin are to be explained by the fact that Woodward rayed her solution in air, while Bersin used a hydrogen atmosphere. Wels and his co-workers state that when egg albumen is rayed in air, the oxidized form of the reversible oxidizable body oc- curs, while in nitrogen it is the reduced form. This difference of effect, depending on the presence of oxy- gen, has a possible explanation in the fact that ultra-violet rays cause water to split off hydrogen in small quantities. If air be present, then hydrogen peroxide results, and this peroxide is responsible for an oxidizing effect. In the absence of oxygen, this ‘‘active’’ hydrogen could be taken up by any reducible body, causing reduced forms to develop. While these reactions undoubtedly occur, there is also the direct effect of the ultra- violet itself to be considered. Hence, the ultra-violet may cause the reversibly oxidizable body to develop from protein, but the presence or absence of oxygen would determine, by the above suggested mechanism, whether this substance would appear in the oxidized or reduced state. VI. CONCLUSIONS A review of the published data concerning the effect of ultra- violet on oxidizing enzymes in vitro shows, with the exception of xanthine oxidase, generally destructive effect. Agulhon (^12) has described three classes of enzymes depending on the part that ozygen plays in their photodestruction. 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 827 l. Invertase, oxygenase (laecase), and tyrosinase are de- stroyed only in the presence of oxygen by visible rays and are less rapidly attacked by ultra-violet in vacuum, the rapidity depending on the amount of hydrogen peroxide that develops. 2. Catalase and emulsin are destroyed in vacuum by all radiations, but less actively in vacuum than in oxygen. 3. Rennet is insensible to the visible radiations, but is at- tacked by ultra-violet with equal intensity in vaeuum and in oxygen. Are these three classes of enzymes dependent on their in- herent differences, or are these divisions only apparent, de- pending on some activity of impurities, or on the nature of the substrate? To these questions Agulhon wisely replies that the present state of knowledge is too incomplete to allow defi- nite answers. The various degrees of injury reported by different workers are not to be considered as inconsistent nor surprising, since many factors both known and unknown are cooperating to give the finally observed effect. This is particularly true of studies carried out in vivo. Despite the efforts of a few writers to correlate physiological phenomena due to raying with the effects on individual en- zymes, it may be said that such correlations are based upon very insecure evidence. The profound difference between ray- ing enzymes in vivo and in vitro itself argues against any such conclusion. The few data reporting similar reactions in vivo and in vitro of peroxidase and catalase require further verification. Finally, we must conclude that it is not possible at the present time to analyze the effect of ultra-violet light on res- pirational metabolism into its various components. VII. BIBLIOGRAPHY Abderhalden, E. (’27). Untersuchungen über die alkoholische Gárung mittels Hefe- zellen unter verschiedenen Bedingungen. Fermentforschung 9: 195-198. Agulhon, H. (11). Action des rayons ultraviolets sur les diastases. Compt. rend. Acad. Paris 152: 398—401. , (712). Action de la lumière sur les diastases. Ann. Inst. Pasteur 26: 38-47. [Vor. 22 828 ANNALS OF THE MISSOURI BOTANICAL GARDEN Asakura, K. (732). Relation between Arakawa's reaetion and suitability of human milk, Tohoku Jour. Exp. Med. 19: 275-281. Austgen, H. G. (719). Uber einige Wirkungen der Quarzlampenbestrablungen bei Lungentuberkulose. Beitr. z. Klin. d. Tuberk. 42: 18-24. Bach, A. (708). Über das Verhalten der Peroxydase gegen Licht. Ber. deutsch. chem, Ges. 41: 225. Battelli, F. ('10). Action de la lumiére sur la catalase. Compt. rend. Soc. Biol. 68: 1040-1042. Bering, F. (12). Beiträge zur Wirkung des Lichtes. Münch. med. Wochenschr. 51: 2795-2797 — und Meyer, H. (’12a). Methoden zur Messung der Wirksamkeit violetter d vilttetelettee Strahlenquellen. Strahlenther. 1: -207. , (’12b). Experimentelle Studien über die Wirkung des Lichtes, Ibid, 411-437. Bernheim, F., and Dixon, M. (’28). Studies on xanthine oxidase. X. The action of light Bio chem. Jour. 22: 113-124. Bersin, T. (733). Über die Einwirkung von Oxydations- n Reduktionsmitteln auf Papain. II. Die Aktivitütsbeeinflussung dureh Licht, Organoarsenverbindun- gen und Ascorbinsäure. Hoppe-Seyler's Zeitschr. f. yhyulei. Chem. 222: 177- 1 Bertraceini, G. (729). Ricerche sui processi deidrogenativi nella cute irradiata eon u. v. Giorn. Ital. Dermatol. e Sifilol, 70: 799-809. [Abstract in Biol. Abs. 4: ] Bierry, H. (726). Action des acd ultra-violets sur les hydrates de carbone. ompt. rend. Soc. Biol. 94: 330-33 Campbell, J. A. (26). Ultra-violet ls and metabolism, with a new method for estimating metabolism. Roy. Soc. London, Proc. B. 99: 451-461. Capelli, G. (726). Comportamento di perossidasi vegetali (Behavior of vegetable peroxidase). Sperimentale 80: 529—539 Castagna, S. (27a). Il eontenuto di ont nel sangue in animali esposti ai raggi ultravioletti. Arch. des Sei. Biol. 10: 118-124. [Abstract in Ber. u. d. ges. Biol. Abt. II. 44: 786. 1928. ; (27b). (The catalase content of blood exposed to ultra-violet rays and to sunlight.) Ibid. 125-127. [Abstraet in Chem. Abs. 22: 97. 1928. Cernovodeanu, P., et Henri, V. (*10). Etude de l’action iu rayons ultraviolets sur les rohe Compt. rend. Acad. Paris 150: 52-54, Cook, R. P., and Stephenson, M. (’28). Bacterial oxidations by molecular oxygen. z The fone oxidation of glueose and certain of its fermentation er in relation to the viability of the organism. Biochem. Jour. 22: 1368-1386. MS Elisabeth ('28a). Studies on basal and resting metabolism 2s radiation with ultra-violet light. I. The effect of ultra-violet radiation on the resting metabolism of birds. Am. Jour. Hre, 8: 1014-1019. 28b). Ibid. II. The eie of ultra-violet radiation on the basal metabolism of human subjects. Ibid. 1020-1023. Delore, P. (729). Sur la capacité d'oxydation de l'huile de foie de morue et sa teneur en oxygéne peroxydique. Influence des rayons ultra-violets. Soc. Chim. Biol, Bull. 11: 74-91. [Abstract in Chem. Abs. 24: 1239. 1931.] 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 829 Durig, - Wd n H. v., und Zuntz, N. (712). Über die Wirkung intensiver Beliehtung auf den TH CM uud die Atemmechanik. Biochem. Zetitschr. v pep. ‚und Zuntz, N. (12). Beobachtungen über die Wirkung des Höhen- klimas aud Teneriffa. Ibid. 435-468 nae S. (722). Die Wirkun a euuatlichen Lichtes auf die alveolare Koh- nsáurespannung. Ibid. 1 103-109. Bichelberger, Marietta (726). p effect of light on creatinine and creatine exere- tion and basal metabolism. Jour. Biol. Chem. 69: 17-2 Essinger, R., und György, P. (724). Beiträge zum Chemismus der Strahlenwirkung. Biochem. Zeitsehr. 149: 344-362. Euler, H. v., und Laurin, I. (718). Verstärkung der Katalasewirkung in Hefezellen. LE Mite: Hoppe- Ped s Zeitschr. f. physiol. Chem. 106: 312- —— — — ——, und Li dy . (712). Über biochemische Reaktionen im Lt I. Biochem. Zeitschr. 39: 410—421. Fazi, R. de (14). British m 1335, January 17, 1914. ———— — ——, (15). (Action of ultra-violet rays upon alcoholic fermentation.) Ann. di en Appl. 4: 301-329. [Abstraet in Chem. Abs. 10: 950. 1916. — The i of ultra-violet rays on the Saccharomycetes. Jour. Tn X Eng. Chem , (728). ye p raggi ultravioletti nella fermentazione aleooliea da Botrytis cinerea. Atti Accad. Lincei Rendie. V. 32: 235-236. [Abstract in Chem. Abs. 17: 3400. — — — ———, (727). Sulla fermentazione alcoolica di soluzioni di glucosio esposte all A. dei raggi m M Ibid. VI. 5: 344-348. [Abstract in Chem Abs. 21: 2045. 1927.] ‚and Fazi, R. de (716). (Action of ultra-violet rays on the alcoholic fommentation of India fig must.) Ann. di Chim. Appl. 6: 221-246. [Abstract in Chem. Abs. 11: 683. 1917.] 717). (Action of ultra-violet rays upon alcoholic fervientation of the mat of India fig.) Ibid. 8: 93-101. [Abstract in Chem Abst. 12: 402. 1918.] , (722). (Action of ultra-violet rays upon Saccharo- myces cerevisiae.) Giorn. di Chim. Ind. e Appl. 4: 463-464. [Abstract in Chem. Abs. 17: 1527. 1923.] (724). (Action of ultra-violet rays on alcoholic fermentation and on yeast.) Atti Cong. Naz. Chim. Ind. 1924: 449—450. [Ab- straet in Chem. Abs. 19: 1470. 1925.] Feuer, B., and Tanner, F. W. (720). The action of ultraviolet light on the yeast- like fungi. I. Jour. Ind. & Eng. Chem. 12: 740-741 Ah The action of ultraviolet rays on the Sac- charom Feet. Ibid. 13: E 266. ren R. (’26). Fragen der Stofechelinderung nach Bestrahlung. Deutsch. ochensehr. 36: 1501-1 TEE sieht E. (725). pe of ultra- y radiations on basal metabolism in children. Proc. Exp. Biol. € Med. 22: 431-433. , (27). Effect of therapeutic E of ultraviolet radiation on basal melaboliam | in children. Am. Jour. Diseases Children 34: 159-165. [VoL. 22 830 ANNALS OF THE MISSOURI BOTANICAL GARDEN ‚and Topper, Anne (’27). Effect of therapeutic doses of ultraviolet n bas piera in children. Ibid. 166-175. Puller, H. J. (732). Some effects of radiations from a mereury vapor are in quartz nenzymes. Ann. Mo. Bot. Gard. 19: 505-531. Gaffron, H. (733). Über den Mechanismus - emi die Do tre durch belich- tete Farbstoffe. Biochem. Zeitsehr. 264: -271. ey A., und Nonnenbruch, W. (723). ^ Wirkung von Strahlenenergie auf ewebsatmang tierischer Zellen. Strahlenther. 15: 98-102, Gronehi, V. (732a). (The action of ultra-violet rays on aleoholie fermentation by Saccharomyces cerevisiae. I.) Soe. Ital. Biol. Sper. Boll. 7: 957-960. [Ab- straet in Chem. Abs. 27: 1447. 1933. , (732b). (Ibid. II.) Ibid. 960-963. [Abstract in Chem. Abs. 27: 1447. 1933.] Hardy, Mary (28). Studies on basal and resting metabolism after radiation with ultra-violet light. III. The effect of ultra-violet radiation on the resting metab- olism of normal rabbits. Am. Jour. Hyg. 8: 1024-1029, Harris, D. T. (25a). Studies on the biological action of light. I. The effect of light on the metabolie rate of small animals. Roy. Soc. London, Proc. B. 98: 171-178. ——— — — ——, (235b). Ibid. II. The part played by pigment in the skin of ani- mals. Ibid. 178-187. Hasselbalch, K. A. (*05). Die Wirkungen des chemischen Lichtbades auf Respira- tion und Blutdruck. Skand. Arch, f. Physiol, 17: 431-472 ‚und Lindhard, J. (*11). Analyse des Höhenklimas in seinen Wirkun- gen auf die Respiration. Ibid. 25: 361-408 Henri, V., et Stodel, G. (’09). er au lait par les rayons ultraviolets. Compt. rend. Acad. Paris 148: 582 Hertel, E. (04). Uber pri b: —— dureh Licht, speziele dureh die chemisch wirksamen Strahlen. Zeitschr. f. allg. Physiol. 4: 1-43. Jamada, K., und Jodlbauer, A. ("08). Die Wirkungen des Lichtes auf die Per eroxy- dase "^ ihre Sensibilisierung durch fluoreseierende Stoffe. Biochem, Zeitschr. : 61-83. Junghagen, S. (’28). Über die Wirkung des ultravioletten Lichtes auf die Dehydro- genosen der Bernsteinsüure und der Glyzerinphosphorsüure. Skand. Arch. f. Physiol, 54: 115-119. Karamitsas, J. (07). Uber die Wirkung des Lichtes auf das Ferment Peroxydase. Inaug. Diss. Univ. München, Kautsky, H., deBruijn, H., Neuwirth, R., und Baumeister, W. (733). Photosensibil- erg Oxydation als Wirkung eines —— metastabilen Zustandes des Sauer- -Molekiils, Ber. deutsch. chem. Ges. 66: 1588-1599. Me D. (729). Cytochrome and uu enzymes, Roy. Soc. London, Proc. B. 04: 206—252. Nube. O. (726). Die physiologischen Wirkung des en Im Handbuch der normalen und pathologischen Physiologie 17: 498-56 meyer, F., Peemöller, F., qe irem R. M db Die Heilwirkung des Hoi, Klin. os. 19: ————————, Hüberlin, C., Lehmann, F., e. E, und Georges, B. (723). Die Heilwirkeng Pd Nordsoekiinas. Klin, Wosheuadie, 44: 2020-2024, 1935] WYND & REYNOLDS—ULTRA-VIOLET. I 831 ————————, Peemöller, F., u Plaut, R. (’23). Die Einwirkung der Strahlung auf Menschen. Ibid. 44: —2019 des RE EDS (22724 mage einer Klimaex- pedition nach Teneriffa. Pflüger’s Arch. 217: 473-49 und Schadow, H. (27). pres Fon, und Gaswechsel Ver- suche am an Ibid. 217: Koeppe, H. (728). Angriffspunkt und DM der ultravioletten Strahlen in der Nahrung und im Kórper des Kindes. VIII. Über das Vermógen der U. V. Strahlen, Oxydation zu fórdern. Archiv. f. Kinderheilk. 85: 81-86. —, (230). U. V.-strahlen und Katalase. Ibid. 89: 3-72. Koldaev, B. M, and Al'Tshulla, M. M. (730a). (Effect of light of a quartz lamp on blood tems ) [In Russian, with German summary.] Jour. Biol. et Méd. Exp. 14: 33-38. (Translation of title. Abstract in Biol. Abs. 7: 1559. 1933.) , (730b). Uber die Beeinflussung der Blutfermente durch Quir iue edle Hoppe Seyler's Zeitschr. f. physiol. Chem. 186: 223-228. ninh in as Abs. 24: 1872. 19 Krestownikoff ‚A. (’27). Die Wirkung des Lichtes auf den Ent ee in inf Deiyarogenasf en ausystem. Skand. Arch. f. Physiol. 52: Kroetz, C. (’24). Zur Biochemie der Strahlenwirkungen. I. Der Einfluss ultra- violetten und ee auf die aktuelle a und auf die Erreg- barkeit des Atemzentrums. ochem. Zeitschr. 151: Laurens, H. (’ he bap rus effects of radiant energy. Am. Chem. Soc. onogr. 62: 61 Lehmann, G. (733). Enida der Ultraviolettbestrahlung auf den Arbeitsstoff- wechsel und die Arbeitsfähigkeit des Menschen. Strahlenther. 48: 364—372 —, and Szakáll, A. (732a). (The influence of ultraviolet radiation on the boe of muscular work and on the capacity of man for work.) Arbeitsphysiol. 5: 278-341. [Abstract in Chem. Abs. 28: 2396. 1934.] , (732b). (Further investigations on the influence of ultra- violat radiation and the metabolism of ir: x on the capacity of man bid. 6: 84-89. [Abstract in Chem s. 28: 2396. 1934. Lehmann, J. (’22). Über die Einwirkung B m Faktoren auf Oxydations- enzyme im Samen von Phaseolus -~ Ein Beitrage zur Kenntnis der De- Basen Bot. Not. 1922: 289-31 Lignae, G. O. E. (’30). (The effect of violet Bi on the formation and ONSE of the skin pigment, melanin.) Neder . Maandschr. Geneesk. 16: : has ire P. (722). (Action of ultra-violet rays on fermentation and yeast.) ochenschr. f. Brauerei 39: 166-167. eint in Chem. Abs. 17: 1686. im Lippmann, A. (728). Licht und Stoffwechsel. Ergeb. d. med. Strahlenforsch. 3: 643—662 ‚und Völker, H. (28). Beiträge zur Frage der Stoffwechselbeein- flussung dureh mW cut pya Wochenschr. 7: 213-214 Loekemann, G., Thies, J., und Wichern, W. (708). eitráge zur Kenntnis der atalase des Blutes. Hoppe ar s Zeitschr. f. physiol. Chem. 58: 390-431. [Vor. 22 832 ANNALS OF THE MISSOURI BOTANICAL GARDEN Lohmann, Ruth (34). Manometrische Untersuchungen über Stoffwechsel und den Bedingungen der Entzündung. Klin. Wochenschr. II. 1934: 1112-1116 Lombard, M. (*10). Sur les effets chimiques et biologiques des rayons ultraviolets. Compt. rend. Acad. Paris 150: 227-229, Mason, E. H., and Mason, H. H. (26). The effect of ultra-violet light upon oxygen consumption and total metabolism, Assoc. Amer, Physicians, Trans. 41: 76-79. ; (27). Effect of ultra-violet light on oxygen con- m and on total Mebéboliun. Arch, Internal Med. 39: 317-329. Masure, M, P. (’32). Effects of ultra-violet radiation on growth and respiration of pea seeds, with notes on statistics. Bot. Gaz. 93: 21-41. Maurain et Wareollier (09). Action des rayons ultra-violets sur le cidre en fer- mentation. Comp. rend. Acad. Paris 149: 155-157. y ; (710). Action des rayons ultraviolets sur le vin en fermentation. Ibid. 150: 343-344. Meldrum, N. U. (*33). Cellular respiration. Methuen and Co. Ltd., London. Merker, E. (25a). Die Atemnot beim Belichten neutralroter Feuchthäuter. Ober- hess. Ges. f. Natur- und Heilk. zu Giessen, Ber. N. F. 10: 41-48. ; (725b). Die Atemnot beim Belichten feuehthüutiger Tiere. Ibid. 67-73 ————— —, ('25e). Die p scade feuchthäutiger Tiere im Lichte. Zool. Jahrb. Abt. f. allg. Zool. u. Physiol. —174. =, (85d). Die Atemnot buts Beliehten neutralroter Feuchthäuter. Zool, "WW 65: 34-40, , (726). Licht und Atmung. Ibid. Supplementband 2: 129-136, und Braunig, G. (’27). Die Empfindlichkeit feuchthäutiger Tiere im Liehte. III. Die Atemnot feuchthiutiger Tiere im Lichte der Quarzqueck- silberlampe. Zool. Jahrb. Abt. f. allg. Zool. u. Physiol. 43: 275-338. Meyer, H. (726). Alte und neue experimentelle Untersuchungen zur Wirkung des Lichtes. Strahlenther. 23: 369—384. Morgulis, S. (729). Inactivation of catalase by ultra-violet radiation. Amer. Jour. Physiol. ad 455 . The inactivation of catalase. II. Inactivation by ultra-violet radiation at different hydrogen-ion concentrations. Jour. Biol. Chem. 86: 75-85. Narayanamurti, D., and Ramaswami, ©. V. (729), Indian -— Sei. Jour. 12 (A): 109—129. rimase in Chem. Zentralbl. II. 1930: 3585. 1930.) y A (30). Irradiation of Dolichos tyrosinase. Biochem. Jour. 24: 1655-1658. Novikov, V. A., and Herber, E. K. (733). (Indueing rubber formation in plants by ultra-violet rays.) Compt. rend. Acad. Sei. U. R. S. 8. (N. S.) 1933: 131-133. [Abstract in Chem. Abs. 28: 1740. 1934. Oster, R. H. ('34). Results of irradiating Saccharomyces with monochromatic ultra-violet light. I. Morphological and respiratory changes. Jour. Gen Physiol. 18: 71-88. Ostwald, W. (*08). Uber die Lichtempfindlichkeit tierischer Oxydasen und über die Beziehungen dieser Eigensehaft zu den Erscheinungen des tierischen Photo- tropismus, Biochem. Zeitschr, 10: 1-130, 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 833 Owen, W. L. (733). Ultra-violet irradiation stimulates yeast aetivity. Food In- dustries 5: 252-254. and Mobley, R. L. (733). The effect of ultra-violet rays upon the fe RE efficiency of yeast in the alcoholic fermentation of molasses. Zentralbl. f. Bakt. Parasitenk. II. Abt. 88: 273-286. Palmieri, L. (733). Sui processi deidrogenativi del cristallino in rapporto all’azione die raggi ultravioletti. (Effect of ultra-violet rays on the dehydrases in the lens.) Riv. di Patol. Sper. 11: 99-120. [Abstract seen in Physiol. Abs. 19: 345. Peemöller, F., und Franke, H. (726). Die Einwirkung ultravioletter Strahlen auf die Blutkatalase. Seraiienther. 21: 165-170. Pineussen, L. (’13a). Über die Einwirkung des Lichtes auf den Stoffwechsel. Ibid. 3: 644—650. —— ———————, (13b). Über die Wirkung des Lichtes auf den tierischen Organis- mus. Dine med. Wochensehr. 44: 2143-2145. —— ————————, (’2la). Zum Verhalten der Oxalsäure im Tierkörper. Biochem. Zeitschr. 126: 82-85. IND: Biologische Lichtwirkungen, ihre physikalischen und chem- ischen een. Ergeb. d. Physiol. 19: 79-289. 6). Über Veränderungen des Fermentgehaltes des Blutes. III. Mitt. Die Beeinflussung der Katalase dureh Strahlung. Biochem. Zeitschr. 168: 474—480. , (730). Photobiologie; Grundlagen, Ergebnisse, Ausblicke. 543 pp. Leipzig. und Hammerich, T. (731a). E dna und Licht. XVII. Über Tyro- sinase. Mundum Zeitschr. 239: 273-2 wo LD) Sin XIX. Über Peroxydase. Ibid. 241: , 384-391. und Ka mean T. (729). Uber Veründerungen des Stoffweehsels unter Bestrahlung . Mitt. Veränderungen in Kohlenhydratstoffwechsel. II Ibid. 208: 0. a be Oya, T. (729). Fermente und Licht. XVI. Über das pH Opti- mum der Milchaldehydrase und die Beeinflussung dieses Fermentes dureh Licht. Ibid. 215: 398—401. und Roman, W. (730). Ibid. XVII. Über den Einfluss des sicht- baren uud ultravioletten en auf die Suceinodehydrogenase des Pferde- muskelfleisches. Ibid. 299: und Seligsohn, F. pron Ibi. VIII. Katalase I. Ibid. 168: 457- 463. und Bos. S. oe Uber ia des Stoffwechsels unter MAN ung. XII. Mit Über den Einfluss verschiedener Strahlen- qualitáten auf den urna der ^in bestrahlter Tiere. 1 265: 64—68. —— — — ———, und Tanino, F. i ne Ibid. VII. Mitt. Veränderungen des Kat- E Ibid. 234: — und Yokota, $. pr qd. IX. Mitt. Der Glutathiongehalt der Organe. mia. 239: 303-309. [VoL. 22 834 ANNALS OF THE MISSOURI BOTANICAL GARDEN — 777, und Zuckerstein, E. (729). Ibid. IV. Mitt. Untersuchungen über den Fettgehalt der Organe. Ibid. 207: 426-431. [Abstract in Ber. u. d. ges. Biol. Abt. II. 51: 182. 1929. Ranjan, S., and Mallik, A. K. ('31). A study of the catalase reaetion, with special reletence to respiration in plants. New Phytol. 30: 355-381. Reinle, H. (21). Uber die Wirkung der Beequerel- und Róntgenstrahlen sowie des ultravioletten Liehtes auf die Peroxydase und Methylenblau-Formalin-Reduk- tase-Reaktion der Kuhmileh. Biochem. Zeitschr. 115: 1-21. Remesow, I. (32). Uber Nadi Reaktion. Arch. f. path. Anat. u. Physiol. 285: 591-598. 1938. Römer, P. H., und Sames, T. (10). Notizen zur Frage der Milchsterilisierung dureh ultraviolettes Lis ht. Hyg. Rundschau 20: 873-877. Roskin, G., und Schischliajewa, S. ('34). Zur vergleichenden Untersuchung der Ultraviolettwirkung auf die lebende Substanz. (Beobachtungen an Para- mecium caudatum). Strahlenther. 49: 596-617 Rubow, V., und Sonne, C. (712). Untersuehungen über die Wirkung des universellen Lichterythems auf die Respiration bei Herzkrankheiten. Zeitschr, f. klin, Med. 75: 33-44. Sehnitzler, J., et Henri, V. (709). Action des rayons ultra-violets sur la fermenta- tion acetique du vin. Compt. rend. Acad. Paris 149: 312-314, ; (10). Die Wirkung der ultravioletten Strahlen auf die NHesigskuregkrung des Weines. Biochem. Zeitschr. 25: 263-271 Söhngen, N. L., and Coolhaas, C. (23). (Influence of ultra-violet light on alcoholic fermentation.) Wochenschr, f. Brauerei 40: 187-188. [Abstract in Chem. Abs. 18: 1029. 1924. Sonne, C. (726). Investigations of the action of light upon oxygen eonsumption. cta Radiol. 5: 419—456 , (729). The bisleghud effeets of the ultra-violet rays and investiga- tions as to what part of the speetrum they lie in. Arch. Physieal Therapy, X-ray, Radium 10: 239-252 Surányi, G., and Vermes, M. (29). The effect of ultra-violet radiation on = resbitetion of avian erythrocytes and yeast cells. Magyar Orvosi Arch. 30 585-590. [Abstract in Chem. Abs. 25: 2162. 1931 " Tanner, F. W., and Byerley, J. R. ('34). The effect of der violet light on the fer- menting ability of yeasts, Archiv. f. Mikrobiol. 5: 349-357 and Ryder, E. (723). Action of ultra- violet light. on yeast-like fungi. II. Bot. ben, 75: 309-317 Tian, A. (11). Sur la disons de l'eau par la lumiére ultra-violette. Compt. 14. Vallet, G. ('10). Pénétration et action bactéricide des rayons ultra-violets par rap- port à la constitution chimique des milieux, Ibid. 150: 632-634. Van Aubel, E. (^09). Sur la production d'ozone sous l'influence de la lumière ultra- violette. Ibid. 149: 983-985. Vasetko, N. (728). (Anderung des Würmehaushaltes bei weisen vp eai er influss der Róntgen- und Ul traviolettenstrahlen.) Ukrain. med. Vistnik. 4: 801-803. [Translated from Ukranian. Abstract in Ber. u. d. ges. Biol. p B. 50: 537. 2 1935] WYND € REYNOLDS—ULTRA-VIOLET. I 835 pe U. (724). (The influence of ultra-violet light and heat upon the oxidase anules in leucocytes.) Nederland. Tijdschr. Geneesk. 68: 2357-2364. [Ab- inn in Chem. Abs. 19: 1143. 1925.] Waentig, P., und Steche, O. (12). Über die fermentative Hydroperoxydzersetzung. gp Mitteilung. Hoppe-Seyler’s Zeitschr. f. physiol. ehem. 76: 177-213. Warnshius, G. J. (733). Influenee of ultraviolet in the role t oxygen in mineral metabolism and immunity reactions. Arch. phys. Ther. 14: 220—224. Wels, P. (733). Die Wirkung der Strahlen auf einige elementare Lebensvorgänge. Sfrahlenthor: 47: 401-407. und Hesse, R. (7/34). Uber einen Atmungskórper im bestrahlten Eiweiss. Naturwiss. 22: 648. , und Jokiseh, Margot (733). Die Entstehung eines reversibel oxy- dierbaren Kórpers dureh Bestrahlung von Eiweiss? Naunyn-Schmiedebergs Arch. 171: 480—495. [Abstract in eN u. d. ges. Biol. Abt. B. Ber. u. d. ges. Physiol. u. exp. Pharm. 76: 397. 34.] Wickwire, G. C., and Hage W. E. (727). The mode of ultra-violet radiation in de- ereasing sugar metabolism. Am. Jour. Physiol. 81: 514-515. Wohlgemuth, J., und Sugihara, N. (725). Uber die Fermente in der Haut. III. Vergleichende PS úber den Fermentgehalt frischer Haut von Menseh und Tier und über den Einfluss verschiedener Lichtarten auf die Haut. Be N 163: umm , und Szörényi, E. (733a). Uber die Wirkung des Lichtes auf den Chemismus der Zelle. I. Mitt. Versuche an Gewebsschnitten. Ibid. 264: , (33b). Ibid. III. Mitt. Einfluss des Histamins. , Ibid. 406-11. , (330). Uber die Wirkung des Lichtes auf den Chem- ismus "m qm unter doni Einfluss von Sensibilisatoren. Klin. Wochenschr. II. 1933: ii Yamasaki, Y. (724). Über die Fermente in der Haut. Ibid. 25: 1113-1114, Woodrow, J. W., Bailey, A. C., and Fulmer, E. J. (727). The effect of ultra-violet radiation upon yeast eulture media. Plant Physiol. 2: 171-175. Woodward, Gladys Estelle (33). Effect of ultra-violet, radium and X-ray radia- tion on glutathione in pure solution. Biochem. Jour. 27: 1411-1414. Wynd, F. L. (734). The effects of increasing the iodine content of the tomato plant on respiration and enzymatic activity. Ann. Mo. Bot. Gard. 21: 367—432. Yamasaki, Y. (24). Uber die Fermente in der Haut. Biochem. Zeitschr. 147: 203- 215. Zeller, M., and Jodlbauer, A. (708). Die Sensibilisierung der Katalase. Ibid. 8: 84-97 STUDIES IN ULTRA-VIOLET AND RESPIRATORY PHENOMENA. II THE EFFECTS or ULTRA-VIOLET ON RESPIRATION AND RESPIRATORY Enzymes or HIGHER PLANTS? F. LYLE WYND Henry Shaw School of Botany of Washington University Formerly Assistant in Botany in the Henry Shaw School of Botany HARRY J. FULLER Associate in Botany, University of Illinois Formerly Instructor in Botany, Henry Shaw School of Botany of Vashington University ERNEST S. REYNOLDS Physiologist to the Missouri Botanical Garden Associate Professor in the Henry Shaw School of Botany of Washington University The present paper reports observations on the respiratory rate and activity of oxidizing enzymes of tomato and bean plants as affected by ultra-violet radiation, and offers sugges- tions concerning the relationship of oxygenase and peroxidase to the respiratory mechanism. The pertinent literature has been discussed in the first paper of the series (Wynd and Reynolds, ’35). EXPERIMENTAL METHODS From several hundred plants of Bonny Best tomatoes and Red Kidney field beans, those of uniform size and vigorous physiological condition were selected. They were irradiated under the conditions of the individual experiments by a Burdick quartz mercury vapor are suspended above the plants. The distances reported were measured from the are to the surface of the soil. Two types of filters were used : one a quartz-plate 1 The investigations reported in this paper were carried on through aid from the Science Research Fund of Washington University, St. Louis, provided by the Rocke- feller Foundation. ANN. Mo. Bor. GARD., Vor. 22, 1935. (837) [VoL. 22 838 ANNALS OF THE MISSOURI BOTANICAL GARDEN tray 5 mm. thick, containing 2 em. of distilled water; the other, a tray made of Vita glass, containing a similar depth of water. The quartz water cell, according to Jones (728), absorbs all wave lengths longer than 1.4 », and the Vita glass water cell eliminates not only this infra-red but also the ultra-violet shorter than 2894 A. Respiration was determined by weighing the amount of car- bon dioxide eliminated per gram of fresh weight over a given period (Wynd, *32, '85). The intrinsie accuracy of this ap- paratus is very great. However, the respirational rates of the plants themselves were found to vary about + 5 per cent in du- plieate controls, and so this must be regarded as the maximum accuracy of the data obtained. The plants were eut off at the surface of the soil in the evening and placed in beakers of water in the respiration chambers, and the next morning the carbon dioxide absorption bulbs were weighed. Five plants were used for each determination, and observations on the control and the experimental plants were carried on simultaneously. Since all of the enzymatie determinations were made on juice pressed from the plants after they had been rayed, the values obtained represent in vivo reactions. The necessary distinc- tion between in vivo and in vitro experiments has been particu- larly pointed out by Fuller (732) and by Wynd and Reynolds (735). The upper leaves and the upper 2 inches of the stems were ground to a paste in a mortar with a little quartz sand, then squeezed through 2 layers of cheese-cloth, and aliquots of this solution used for the enzymatie determinations. Since the usual methods of measuring acidity are disturbed by oxidizing enzymes or by hydrogen peroxide, the final pH of all experi- mental solutions was checked by the glass electrode. Extended experimentation by Wynd has shown that the de- termination of catalase activity can be more closely duplicated by titration than by gasometrie methods. However, the gaso- metric method is entirely satisfactory as a micro procedure if the usual precautions are taken. Experiments involving rela- tively large amounts of material can be duplicated by titration with an error of 1 or 2 per cent, which is not possible by macro gasometrie procedures. The experimental solutions contained 1935] WYND, FULLER € REYNOLDS—ULTRA-VIOLET. II 839 50 ec. of phosphate buffer at pH = 7.00, 20 ec. of 3 per cent com- mercial hydrogen peroxide, and 1 cc. of the expressed juice. The reaction time varied from 8 to 20 minutes in the different experiments, and it was stopped by adding 20 cc. of a 25 per cent solution of sulphurie acid. The time of adding the plant juice and of the subsequent addition of acid was carefully con- trolled by a stop watch. The amount of hydrogen peroxide re- maining was then titrated with N/5 potassium permanganate. The reaction times of all the solutions were spaced 1 minute apart to allow time for the pipettings. Peroxidase was determined by the method of Guthrie (731). The substrate for the enzyme was prepared by adding 200 ce. of a citrate buffer at pH = 4.5 to 200 cc. of water. To this were added 1 gm. of p-phenylenediamine hydrochloride and 20 cc. of a 4 per cent solution of alpha-naphthol in 50 per cent aleohol. The final mixture was filtered and used immediately. At pH = 4.5, atmospherie oxidation of the substrate is slow, and catalase is so far depressed as not to seriously interfere. The experi- mental solutions consisted of 25 ce. of substrate, 1 cc. of com- mercial 3 per cent hydrogen peroxide, and .5 cc. of the plant juice. After 10 to 20 minutes, depending on the intensity of the color which developed, the reaction was stopped by the ad- dition of 5 ec. of a.1 per cent potassium cyanide solution. Since the cyanide inhibits the enzyme but does not interfere with the atmospherie oxidation, the indophenol produced was deter- mined as quickly as possible. This was done by shaking each experimental mixture with 50 cc. of toluene until the water layer was clear. The upper layer of toluene, after being poured off and centrifuged to remove bubbles and tissue fragments, was examined in a colorimeter. The error of this method is about 3 per cent. Oxygenase was determined similarly to peroxidase except that the hydrogen peroxide was not added to the substrate. Due to the feeble activity of oxygenase as compared with that of peroxidase, 1 cc. of plant juice was used for each determina- tion, and the reaction time was usually extended to 20 minutes. The indophenol was extracted in 25 ec. of toluol. The error of this method is about 3 per cent. [VoL. 22 840 ANNALS OF THE MISSOURI BOTANICAL GARDEN Since the temperature, reaction periods, etc. varied from day to day, the absolute values of data obtained at different times may not be directly compared, and the tables contain values calculated as percentages of their corresponding con- trols. These percentage values obtained at different times may be compared directly. In the figures, the numbers on the ordinate represent the per cent of stimulation or depression, the graphical value of the control therefore being zero. The horizontal line at zero divides the stimulation from the depres- sion values. The numbers on the abscissa represent minutes of daily exposure. EXPERIMENTAL RESULTS Experiment I.—Bonny Best tomato seeds were planted in greenhouse flats January 23, 1934, and were transplanted to 4-inch pots February 8. The raying began March 4, and was continued daily for 5 days. The distance was 24 inches and the dosages 4, 8, 16, and 32 minutes daily. The unscreened mer- cury arc was used as a source of illumination. TABLE I ENZYMATIC ACTIVITY OF TOMATO PLANTS RAYED AT 24 INCHES WITH THE UNSCREENED MERCURY ARC, CALCULATED IN PERCENTAGES OF THE CONTROL b Daily Dosage Control | y g | 4 minutes 8 minutes | 16 minutes | 32 minutes After 5 days of irradiation Oxygenase 100 96 107 96 78 Peroxidase 100 97 159 166 213 Catalase 100 123 134 119 125 xygenase 10 95 92 88 74 Peroxidase 100 189 263 345 385 Catalase 100 115 146 194 165 All plants showed definite injury at the end of the 5-day period of irradiation. The plants submitted to the heavier 1935] WYND, FULLER € REYNOLDS—ULTRA-VIOLET. II 841 dosages were very severely burned. One set of determinations was made at the end of the irradiation period and another 5 days later. The plants were not rayed during this intervening period. From table 1 and figs. 1 and 2, it is seen that with in- creasing amounts of daily irradiation the peroxidase activity greatly increased. Catalase also increased but less than the Peroxidase 80 60 40 20 Catalase [o EA Control —— te UNIX Value Oxygenase 20 40 2 à 4 — Minutes 4 8 16 32 Fig. 1. Determination of enzymatic activities of tomato plants afte ee irradiated daily for 5 days at 24 inches, through quartz water- cell filte peroxidase, while oxygenase was progressively inhibited. These results were of greater magnitude 5 days after irradia- tion ceased. Experiment II.—Bonny Best tomatoes were planted Febru- ary 15, 1934, and transplanted from the flats to 4-inch pots on [Vor. 22 842 ANNALS OF THE MISSOURI BOTANICAL GARDEN April5. The raying began April 16, at a distance of 100 inches, and continued in 10-minute doses daily for 5 weeks. According to Fuller (731), these are the optimum conditions for growth stimulation. Some of the plants were rayed through the quartz Peroxidase Catalase Oxygenase +M 4 8 16 32 Minutes 2. Determination of enzymatie activities of tomato plants 5 days after irradiation ceased. water cell, and others through the Vita glass water cell. Two determinations of enzymes and respiration were made on con- secutive days at the end of the period of irradiation. The data in table 11, graphically represented in fig. 3, indi- cate that the plants rayed through the Vita glass water cell 1935] WYND, FULLER € REYNOLDS—ULTRA-VIOLET. II 843 TABLE II gud e OF IRRADIATING 5 TOMATO PLANTS AT 100 INCHES FOR 10 MINUTES DAILY FOR 5 WEEKS, CALCULATED IN PERCENTAGES OF THE CONTROL One day after irradiation ceased Fresh wt. in Tun piene percentage of Treatment gms. of tops CO, elimination nzymatie aetivity . | Per gm. y Oxy- Per- Control 95.6 | 100 .00098 100 100 100 100 Rayed through Vita glass water cell 107.5 | 112 .00128 131 164 104 100 Rayed through quartz water 11 93.5 98 .00102 104 190 117 103 Two days after irradiation ceased Control 84.5 | 100 .00200 100 100 100 100 Rayed through Vita glass water cell 102.0 | 121 .00218 109 142 103 100 m de Mig ar water 90.0 | 107.0 .00208 104 118 106 102 were stimulated in growth from 12 to 21 per cent. Since only 5 plants were used for the determination, this result is not statistically significant, but there is qualitative agreement with the results published by Fuller (7/31) on the stimulation of growth under these conditions. Respiration is also increased. Catalase showed a very great increase, while oxygenase was but slightly increased, and peroxidase was not affected. The variations given for oxygenase probably are not far beyond the experimental error. There was qualitative agreement be- tween the data obtained on successive days, and consequently on different samples of tissue. Irradiation through the quartz water cell, which allows much of the short, injurious ultra-violet rays to pass, had no con- [Vor. 22 844 ANNALS OF THE MISSOURI BOTANICAL GARDEN sistent effect on growth, although, as noted above, the number of plants was too small to present convincing evidence. Res- piration was stimulated only slightly, less than when the Vita glass water cell was used as the filter. Again catalase was greatly stimulated, while oxygenase showed a smaller increase. Peroxidase was slightly inereased, although the increase is too close to expeet experimental error to be accepted as significant. It is thus seen that the physiological effects of rayings through Control GROWTH Vita glass — Quartz Contr RESPIRATION vita giass Quartz if Ls Control PEROXIDASE Vita glass de L_ Quartz Con OXYGENASE Vita glass ae ontrol a CATALASE Vita glass [ Quartz Fig. 3. Determinations made on tomato plants day after irradia- tion eeased. Plants had been irradiated at 100 inches for 10 minutes daily for 5 weeks. Linear values caleulated as percentages of the contro the quartz water cell were very similar to those obtained by the use of the Vita glass water cell, with an indication of a possible injury of the plants. Experiment III.—The tomato plants used for this experi- ment were planted February 15, 1934, and transplanted from the flats to 4-inch pots on April 5. The raying began April 20, at a distance of 36 inches from the arc. The quartz water cell was used as a filter. One group was rayed 15 minutes, and another group 30 minutes, daily for 7 days. Determinations 1935] WYND, FULLER € REYNOLDS—ULTRA-VIOLET. II 845 were made daily for 4 days following the irradiation period. The data are included in table 11, and are represented graphi- cally in figs. 4—7 inclusive. All of the rayed plants in this experiment were greatly in- jured. The first set of determinations, made the day after the 60 Respiration oo Peroxidase 40 20 20 Peroxidase HE SUPE o edntrofevalue o —— — —— — Control Value Catalase “au ae Growth 20 Growth 0 on pude 40 xygenase 40 1g go Minutes is — 30 Minutes 2 4. Determinations made on bon Fig. 5. Determinations made on to- o plants after being rayed at 3 mato plants on the second day after s for 7 days with quartz Ls irradiation ceased. cell filter 80 Catalase E idas 80 erox e 60 60 40 Peroxidase 40 20 20 ey ud : Catalase | Qnm C Y NT Y alue 3 Oxygenase ee 0 > ut ontrol Value oxygenase SS 20 ee Growth Growth 40 40 15 E 15 Pla vis 6. Determinations made on to- DE 7. Determinations made on to- mato plants on the third day after mato plants on the fourth day after n T adictas ceased. irradiations ceased, showed a heightened rate of respiration that was proportional to the length of exposure. On subse- quent days the amount of carbon dioxide eliminated per gram of fresh weight remained at the heightened level in those plants rayed for 15 minutes daily, but there was a great drop in respiration on subsequent days in those plants which [Vor. 22 846 ANNALS OF THE MISSOURI BOTANICAL GARDEN TABLE III RESULTS OF IRRADIATING 5 TOMATO PLANTS THROUGH ¡mail WATER ELL AT 36 INCHES DAILY FOR 7 DAYS, CALCULATED PERCENTAGES OF THE CONTROL Relative percentage of Fresh wt. in gms. n ej of tops CO, elimination enzymatie aetivity Treatment 1 Per gm. , Oxy- Per- Actual | Relative | fresh wt,| Relative | Catalase genase | oxidase First determination Control 60.2 100 .00130 100 100 100 100 Rayed 15 minutes 46.5 77 .00165 127 77 85 92 Rayed 30 minutes 49.0 81 .00209 161 87 67 119 Second determination Control 57.4 100 .00219 100 100 100 100 Rayed 15 minutes 53.9 94 .00235 108 75 67 102 Rayed 30 minutes 43.8 76 .00223 107 104 62 135 Third determination Control 80.4 100 .00174 100 100 100 100 Rayed 15 minutes 64.2 79 .00208 129 117 94 115 Rayed 30 minutes 58.7 73 .00153 89 112 108 179 Fourth determination Control 86.7 100 .00228 100 100 100 100 Rayed 15 minutes 54.4 63 .00264 116 149 103 115 Rayed 30 minutes 59.1 67 .00248 109 180 91 145 1935] WYND, FULLER € REYNOLDS—ULTRA-VIOLET. II 847 had been rayed for 30 minutes daily. This drop is only ap- parent, because there were progressively larger amounts of withered and dead tissue in these greatly injured plants on each succeeding day. Since these non-respiring tissues were un- avoidably included in the recorded fresh weight, a false value is obtained. This was not an important source of error in those TABLE IV RESULTS po IRRADIATING 5 BEAN PLANTS AT 70 INCHES FOR 10 MINUTES Y FOR 5 WE EOS eres IN PERCENTAGES THE CONTRO Fresh wt. in gms. A. gie pereentage of of tops CO, elimination matie activity Treatment : Per gm. 1 ' Actual | Relative | fresh wt.| Relative | Peroxidase | Catalase First determination Control 92.5 100 .0053 100 100 100 Rayed through ita glass water cell 74.5 81 .0053 100 150 91 Rayed through ien artz water ell 59.8 64 .0049 92 237 100 Second determination Control 70.0 100 .0062 100 100 100 Rayed through Vita glass water cell 58.0 83 .0057 92 119 95 Rayed through quartz water 1 45.0 63 .0068 109 145 108 plants rayed 15 minutes daily. Since the enzymes were deter- mined in the extracted sap, these values represent more truly the comparative activities of the actually living tissue. The peroxidase, with the exception of a single determination, fol- lowed the upward trend of respiration. Oxygenase was greatly decreased during the first 2 days after the irradiations ceased, [Vor. 22 848 ANNALS OF THE MISSOURI BOTANICAL GARDEN the decrease being proportional to the length of exposure, but normal values were regained on the third and fourth day. Catalase showed a progressive increase for the four days fol- lowing the termination of the raying. The activity of catalase shows no apparent relationship to respiration nor to the activi- ties of peroxidase and oxygenase. Experiment IV.—For this experiment, Red Kidney field beans were planted in flats January 23, 1934, and transplanted Control Quartz Control | RESPIRATION Vita puni Quartz Control | CATALASE Vita glass glass Quartz RAR adorna Control | PEROXIDASE Vita glass Quartz Fig. 8. Determinations made the day after irradiation eeased on bean plants rayed at 70 inches for 10 minutes daily for 5 weeks. Linear values caleulated in pereentages of the controls into 4-inch pots February 8. The irradiation began February 15, and continued for 5 weeks. The distance was 70 inches, and the daily exposure was 10 minutes. One group was rayed through the quartz water cell, and one through the Vita glass water cell. One set of determinations was made the day after the irradiations ceased, and another 4 days later. The results are presented numerically in table rv and graphically in figs. 8 and 9. All irradiated plants showed injury. The injury and stunt- ing were slight in those plants rayed through the Vita glass water cell, but quite pronounced in those rayed through the quartz water cell. Respiration and catalase were not signifi- 1935] WYND, FULLER € REYNOLDS—ULTRA-VIOLET. II 849 cantly affected, but peroxidase was very markedly increased in both rayed groups, particularly when the quartz water cell was used as the filter. Five days after irradiation ceased these dif- ferences were still evident although of lesser magnitude. The tomato plants in experiment II were rayed under approxi- mately similar conditions, but they were stimulated in growth, respiration was slightly increased, peroxidase was insignifi- cantly affected, and catalase was very greatly stimulated. Control GROWTH Vita glas | Control RESPIRATION Vita glass Quartz m Control CATALASE Vita glass | Quartz L Control PEROXIDASE Vita elass | Quartz Fig. 9. Determinations made 5 days after irradia- tion ceased on bean plants rayed at 70 inches for 10 minutes daily for 5 weeks. Linear distances calculated in percentages of the controls. DISCUSSION These experiments on irradiated tomato plants show that the rate of respiration as indicated by the amount of carbon dioxide eliminated per gram of fresh weight markedly in- creases proportionally to the extent of injury. However, it is not possible to obtain accurate numerical data on plants in the most advanced state of injury because the development of dead, non-respiring tissue confuses the weights. This increased rate of respiration is approximately paralleled by the activitiy of peroxidase in the expressed juice. Oxygenase, on the other hand, is inhibited proportionally to the extent of raying. Ex- periment III shows that normal oxygenase activity is regained [Vor. 22 850 ANNALS OF THE MISSOURI BOTANICAL GARDEN soon after irradiation stops, although the plants remain in a very disturbed physiological condition. Catalase reacts vari- ously. In experiment I it was stimulated, yet in experiment III determinations made the day after raying ceased gave sub- normal values. On successive days its activity increased until a considerable degree of stimulation was reached. "These re- sults agree with those of Fuller (32). The irradiated bean plants reacted somewhat differently. Their growth was not stimulated under conditions stimulative to tomatoes. In experiment IV, the only decided effect was a greatly stimulated peroxidase activity, while the tomato plants under the approximately similar conditions of experiment II showed a greatly stimulated catalase activity as the chief physiologieal reaction. Bean plants do not give the colori- metric test for oxygenase, and this is itself sufficient evidence that the physiology of beans is different from that of tomatoes. It is not surprising, then, that beans and tomatoes react dif- ferently in regard to other features of their physiology. The data obtained in these experiments have a bearing on the nature of the oxidation processes in plants. The numerous contributions of Bach and Chodat on the intracellular oxida- tion mechanism have been responsible for the concept that oxy- genase unites molecular oxygen to some cellular component to form a peroxide. This peroxide is then activated towards some oxidizable substance in the protoplasm by peroxidase. It is obvious that if this relationship be true, then oxygenase and peroxidase would necessarily be related quantitatively to each other. This is indeed suggested by numerous experiments upon diseased plant tissues in which both enzymes were markedly greater in activity than in normal plants. A sum- mary of these experiments has been published by Wynd (734). The necessity of oxygenase for the respiratory process is, however, open to several serious objections. Onslow (’21, ’31) has shown that only about 60 per cent of the higher green plants give the oxygenase reaction and that any fundamental component of the respiratory mechanism would be more uni- versally distributed. Kertesz (734) has shown that fresh tis- 1935] WYND, FULLER & REYNOLDS—ULTRA-VIOLET. II 851 sue of some varieties of peaches does not discolor upon ex- posure to the air. Qualitative and quantitative studies showed the lack of catechol compounds in the non-discoloring varieties. This indicates that the oxygenase-catechol system may be ab- sent even in very closely related plants, and consequently could not be of fundamental importance. Wynd (734) has shown, in an extended study of the tomato plant, that there is no quantitative relationship between the ac- tivities of oxygenase and peroxidase. Plants stimulated to great respirational activity by the addition of potassium iodide to the nutrient solution showed a corresponding increase of peroxidase but oxygenase was greatly inhibited. These re- sults, together with those of the present paper, tend to indicate that the oxygenase-catechol-peroxidase system does not rep- resent a fundamental respiratory mechanism in the tomato plant. SUMMARY 1. Tomato and bean plants were rayed under various condi- tions by a mereury are, and the effect on respiration and oxi- dizing enzymes determined. 2. In tomato plants, injurious raying stimulated respiration and peroxidase. Oxygenase was inhibited, and catalase re- acted variously but in general was stimulated. The most pro- nounced effect of non-injurious raying was greatly stimulated catalase. 3. In bean plants, comparatively weak dosages produced greatly stimulated peroxidase as the chief reaction. Bean plants did not exhibit oxygenase activity, which indicates a different chemical physiology consistent with their different reactions to ultra-violet light. 4. The comparative activities of peroxidase and oxygenase show that the oxygenase-peroxide-peroxidase system as pro- posed by Bach and Chodat does not represent a fundamental respiratory mechanism in the tomato plant. BIBLIOGRAPHY Fuller, H. J. (*31). Stimulatory effects of radiation from a quartz mercury vapor are upon higher plants. Ann. Mo. Bot. Gard. 18: 17-40. [Vor 22, 1935] 852 ANNALS OF THE MISSOURI BOTANICAL GARDEN ; (732). Some effects of radiations from a mercury vapor are in quartz upon enzymes. Ibid. 19: -531. Guthrie, J. D. (731). A method for the determination of peroxidase activity. Am. Chem. Soc. Jour. 53: 242-244. Jones, L. A. (’28). Light filters for the isolation of narrow speetral regions. Opt. oc. Am. Jour. 16: 259-271. Kates, Z. I. (733). The oxidase system of a non-browning yellow peach. New York Agr. Exp. Sta. Tech. Bull. 219: 1-14. Onslow, M. W. (721). Oxidizing enzymes. IV. The distribution of oxidizing en- zymes among the higher plants. Biochem, Jour. 15: 107-112, ). The prineiples of plant biochemistry. Part I. 326 pp. Cam- bridge. Wynd, F. L. (732). An apparatus for the determination of carbon dioxide produe- tion in physiological plant studies. Ann. Mo. Bot. Gard. 19: 499-501 ; (734). The effeets of inereasing the ine eontent of the tomato plant on respiration me enzymatic ein i Ibid. 7—432 —————— mproved apparatus for the Pesca of carbon dioxide production i in afi tritio Cien studies. y bid. 22: 361-363. , and Reynolds, E. S. (735). Studies in ultra-violet and respiratory phenomena. I. Review of work published before J une, 1935. Ibid. 22: 771-835. STUDIES IN ULTRA-VIOLET AND RESPIRATORY PHENOMENA. III THE INFLUENCE OF VARIOUS REGIONS OF THE SPECTRUM ON THE ANAEROBIC FERMENTATION OF YEAST! E. S. REYNOLDS Physiologist to the Missouri Botanical Garden Associate Professor in the Henry Shaw School of Botany of Washington University AND F. LYLE WYND Henry Shaw School of Botany of Washington University Formerly Assistant in Botany in the Henry Shaw School of Botany As was pointed out in the first paper of this series,? the claim that ultra-violet irradiation stimulates the fermentation of yeast is based on very insecure foundations. The recent re- vival of this claim warrants a new and more critical approach to the problem. The present paper reports the results of an attempt to duplicate the essential features of the work of Owen? but under more accurately controlled experimental conditions. EXPERIMENTAL PROCEDURE The CO» elimination was determined by observing the posi- tive pressure developing in the Barcroft differential manom- eter. Similar vessels were attached to each arm, one vessel re- ceiving the yeast suspension while the other remained empty. Both were thoroughly swept out with nitrogen and the manom- eter taps immediately closed. After shaking for 30 minutes in 1 The Fe reported in this paper were carried on through aid from the Science Research Fun d of Washington University, St. Louis, provided by the er ller Bie : nd F. Lyle, Dr fuod s, E. S. Studies in ultra- ve and respiratory phe- na. I. Review of work published before June, 1935. . Mo. Bot. Gard. 22: pos pes 1935. 3 Owen, W. L. Ultra-violet irradiation stimulates yeast activity. Food Industries 5: 252-254. 1933. ANN. Mo. Bor. GARD., VOL. 22, 1935. (853) [VoL. 22 854 ANNALS OF THE MISSOURI BOTANICAL GARDEN the water bath, the taps were opened and the manometer liquid allowed to come to equal levels in each arm. The taps were then closed, and the positive pressure plotted by frequent readings. The vessels were of the type shown in fig. 1. The total vol- ume was about 12 cc. Chamber A received the culture suspen- sion and Chamber B an alkali when CO» absorption was desired. A metal shield protected the manometer from the heat of the mercury arc. The water bath was of the Warburg type in which the tem- perature was held to 25° C., + .001? C. by a sensitive Thyratron Fig. l. Respirometer vessel: A, culture chamber; B, alkali chamber; C, quartz cover glass. relay. The manometers were shaken at the rate of 100 excur- sions of 4 em. in length per minute. The source of irradiation was a Burdick quartz mercury are suspended immediately above the respiration vessel and re- moved 25 em. from the thin quartz plate C. The quartz plate C was immersed 3 em. below the surface of the water of the water bath. A conveniently built rack allowed the various filters to be placed several em. above the water surface. A quartz tray, containing water 1 cm. deep, was suspended just below the are itself. This was necessary to prevent the heat from the are from cracking the filters, some of which were very fragile. 1935] REYNOLDS € WYND——ULTRA-VIOLET. III Metal shield One limb at tie Filter di Mi al nometer KOH Yeast Culture Water bath Fig. 2. Arrangement of the apparatus for raying eas eul- tures. The alkali is omitted in fermentation experimen [Vor. 22 856 ANNALS OF THE MISSOURI BOTANICAL GARDEN Through this set-up also the infra-red radiations were elim- inated. The following light filters were used: Red Purple Corex A, No. 986—about 4.0 mm. Chemical Pyrex Plate—2.1-2.6 mm. Blue Purple Ultra, No. 585—4.4-4.6 mm. Red Purple Ultra, No. 597—4.5-5.4 mm. Nultra, No. 385 Noviol Shade ‘‘O,’’ No. 306 Vita glass—2.6-3.0 mm. The transmissions of these filters are indicated in fig. 3. Pasteur's yeast medium was used as the substrate. Its com- position was : 150.0 gms. glucose 10.0 gms. ammonium tartrate 2.0 gms. mono-potassium phosphate 0.2 gms. calcium phosphate 0.2 gms. magnesium sulphate 1000.0 cc. water Since autoclaving causes this medium to turn brown if sugar is present, aliquots of 100 cc. were sterilized without the sugar, and 15 gms. of glucose were added just before use. Fleisch- man's commercial baker's yeast was obtained in pound cakes, only the center portion being used, and 7 gms. of it were seeded in the 100-ce. portions. The seeded medium was vigorously aerated at 25? C. for 30 minutes. Three drops, or about 0.1 of a cc., were then delivered into chamber A of the vessel shown in The cultures were not pure in the bacteriological sense, but preliminary tests showed that at 25° C. the medium allowed no significant bacterial growth during the short time that each experiment lasted. As may be seen from the graph in figure 4 for the control eulture, the fermentation curve over the period represented was practically a straight line. This period is the second hour of the life of the culture, since half an hour was spent in the preliminary aeration, and half an hour in tempera- ture equilibration. 1935] REYNOLDS € WYND—ULTRA-VIOLET. III 857 Each experiment was repeated three times with identical re- sults. Almost every determination was made on a separate cake of yeast, thus eliminating the possibility of the results being due to some specifie physiological eondition of the sample. RESULTS No effect on anaerobic fermentation was produced by the ir- radiation passed through any of the following filters: Blue Purple Ultra No. 585; Red Purple Ultra No. 597; Nultra No. 385; Noviol Shade ‘‘O,’’ No. 306. All negative experiments were duplicated with a total of only 1.5 em. of water intervening between the culture and the light, in order to obtain the maxi- mum intensity of irradiation, and in no instance were the re- sults altered. By referring to fig. 3, it is seen that these ineffective regions consist of wave lengths longer than 3000 À. A comparison of figs. 3 and 4 shows that spectrum regions containing wave lengths shorter than 3000 À produce an in- hibition progressively greater as the wave lengths become shorter. Probably the first inhibitory line is 2967 Ä, from which point the irradiation becomes increasingly toxic. It should be noted, however, that possibly some of the mild in- hibition under the chemical Pyrex filter may be due to the mer- cury are line of 3024 Á, since it is excluded by the Blue Purple Ultra filter which gave no inhibition. In all instances, this inhibitory effect on the CO: elimination was detectable man- ometrically within five minutes. It is interesting to note that the Red Purple Corex A, No. 986 filter passed radiations that are conspicuously more effec- tive than did the Vita glass. This Corex glass differs qualita- tively from Vita glass chiefly in its transmission of the strong line at 2537 Ä. Irradiation of this wave length apparently is very toxic, since only about 5 per cent of its intensity calls forth an inhibition about 50 per cent greater than the combined ef- feet of all longer wave lengths. It should be noted, however, that the Corex filter passes these radiations at a somewhat greater intensity than does Vita glass. The increase in area under the graph line (fig. 3), for wave lengths less than 3000 À, 858 Per cent transmission [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Ultra- Violet Visible E Middle Eryth Near [Violet | Blue | Green Ius ^ Red 5 Ki 2lo E 6 . 40 45 «SO 5 o 65 7p 80 amm 6o NO 40 \ 20 LL \ / Mie 80 pet 6 PT Y » e 50 60 /. " ‘a à / P. to a " N in / Z eo A 40 \ 20 Eod Fi © / 4o / 20 / Fá 80 60 A 40 E 20 E Fig. 3. Transmission of various filters. Relative in- tensity of the emission lines of the Hg spectrum Red Purple Corex Vita glass Pyrex (2 mm.) Blue Purple Ultra Red Purple Ultra Nultra Noviol “O” din REYNOLDS € WYND—ULTRA-VIOLET. III 859 for Vita glass as contrasted with Pyrex is about nine times, while the increase in inhibition is only about 60 per cent. This lack of proportionate relationship is probably due to the rela- tively low intensity of the radiations in this portion of the mer- cury arc. The corresponding comparison between Vita glass and the Corex filters is less than twice the area for the latter Cubic millimeters CO, eliminated El 3 «A t2 e P 70 5 10 20 30 20 50 60 Minutes . 4. Curve 1, Control; 2, irradiated through var and 4 em. water; 3, late through Vita glass em. water; 4, irradiated through Red Purple Corex A and 1 Py em. water; 5, irradi ated through Red pis big x A and 4 em. water; 6, irradiated by the open are through 4 em. with an 80 per cent increase in inhibition. This close relation- ship is due no doubt in part to the relatively great intensity of the radiation of 2537 À. "Thus it appears that this increased inhibition associated withe the Corex filter may be due to some extent to qualitative difference in the radiations transmitted and partly also to quantitative difference. Curves 4 and 5 in fig. 4 are as close together as experimental error in the duplication of cultures would permit, and they in- [Vor 22, 1935] 860 ANNALS OF THE MISSOURI BOTANICAL GARDEN dicate that the difference in intensity produced by 4 and by 1.5 em. of water was not sufficiently great to influence the results. Comparison of figs. 3 and 4 reveals that the inhibitory irradia- tion begins with the most effective erythemal region. At the end of each experiment, the cells were treated with 1/2000 methylene-blue (final concentrations) to test for injured or dead cells. The unsereened are produced complete staining at the end of 50 minutes of irradiation. No staining was ob- served when any of the filters described were used. If the staining of yeast by methylene-blue is accepted as evidence of injury this shows that the observed inhibition is a specifie ef- fect and not dependent upon a general injury to the cells. The inhibition due to death of the cells is much more profound and sudden as shown by the curve for the open arc in fig. 4. CONCLUSIONS The results of this investigation indicate that ultra-violet between 3000 and 2500 À has an inhibiting effect upon the fer- mentative activity of suspensions of two-hour-old cultures of baker's yeast when made in the very favorable Pasteur medium. This inhibition of fermentation is a specific effect and is not dependent on a general injury to the cell. The degree of inhibi- tion of fermentation is both a qualitative and a quantitative function of ultra-violet. Irradiations of longer wave length than 3000 À are without effect. A NOTE ON THE APPLICATION OF THE GLASS ELECTRODE TO THE DETERMINATION OF OXIDATION-REDUCTION POTENTIALS F. LYLE WYND Henry Shaw School of Botany of Washington University Formerly Assistant in Botany in the Henry Shaw School of Botany The electromotive force of the chain consisting of a glass eleetrode and a bare noble metal is influenced by the presence of oxidation-reduetion systems in the solution. It has been frequently stated that such an electrode system may be used to determine oxidation-reduetion potentials independently of simultaneous effects of pH, but this statement must be exten- sively qualified, as will be seen from the following discussion. Ignoring the small asymmetry potential existing between the two glass surfaces, there are four sources of electrical activity in the chain. They are the half cell inside the glass electrode, the liquid-glass interface potential on the inner sur- face of the glass, the glass-liquid interface potential on the outer surface of the glass, and the oxidation-reduction poten- tial imparted to the platinum. Any solution containing an oxidation-reduction system in which the glass electrode and the platinum electrode are immersed will affect the electro- motive force of the chain in two ways. The oxidation-reduction system will impart a potential to the platinum, and the hydro- gen ions present will modify the glass-liquid interface poten- tial on the outer surface of the glass. If the pH of the solution remains constant, then only changes in the oxidation-reduction potential will modify the E.M.F. of the chain. The true E» of the oxidation-reduction system may be calculated by applying a calibration correction obtained by observing the E.M.F. of the chain when immersed in some known system at the same pH as that of the unknown ANN. Mo. Bor. GARD., VOL. 22, 1935. (861) [Vor. 22 862 ANNALS OF THE MISSOURI BOTANICAL GARDEN system. If the pH of the solution changes during the course of oxidation-reduction potential measurement, either of two con- ditions may exist. A consideration of the formula is necessary for their explanation: RT (reduetant) RT (reductant) RT E. n E, = E, a Dee = n ———— nF (oxidant) nF (oxidant) nF (H*) +K, The oxidation-reduction potential is seen to be a function of the ratio of total reductant to total oxidant, and also of pH. First, if the system is such that the slope of the curve relating changes in pH to changes in oxidation-reduction potential is the same as that relating changes in pH to changes of the glass- liquid interface potential, then these two effects of pH cancel each other and the potentials of the system would correspond to those obtained at a constant pH. It is this that has given rise to the statement that the glass electrode records oxidation- reduction potential independently of pH changes. The falsity of this view comes from the fact that the observed reading must be corrected by an empirical factor obtained by observing the potential given by a known oxidation-reduction system at the same pH as that of the unknown. "The magnitude of this calibration correction changes with pH, and hence its applica- tion to the observed reading will not give the E» of the solution if the pH has changed from that of the calibrating system. In order to obtain the true En of a system of changing pH, deter- minations of pH must be made simultaneously with those of oxidation-reduction potential, from which new calibration cor- rections can be calculated. On the other hand, if the system is such that the slope of the curve relating changes in pH to changes in oxidation-reduc- tion potential is unlike that relating changes of pH on the glass- liquid interface potential, then the two effects of pH do not cancel each other. In order to convert observed potentials to E, one would need to take simultaneous measurements of pH changes, and then apply as a correction factor only that part of the pH effect that is cancelled by the response of the glass electrode to the same pH changes. This would be a complicated 1935] WYND—GLASS ELECTRODE 863 and tedious process—and only possible if the reaction of the redox system to changes in acidity were accurately known. Clark and Cohen! were the first to point out the various theoretieally possible relationships between pH and oxidation- reduction potentials. They show how very complex is the ef- fect of pH on many oxidation-reduction systems. For this reason, it is impossible to use this chain to determine the En of such complex biological systems as growing bacterial cultures, if the pH is changing simultaneously. The use of the glass eleetrode in the determination of oxi- dation-reduetion potentials is limited to buffered solutions, and its advantage lies solely in its high resistance and conse- quent laek of polarization. A high resistance (10 megohms) inserted in the usual ealomel-platinum chain would be equally advantageous. It may be pointed out that if the potential of the oxidation- reduction system changes with pH to the same degree as does the glass-liquid interface potential, then the observed E.M.F. of the oxidation-reduction system will be in terms of the pH inside the glass electrode. "This apparently anomalous phe- nomenon has a simple explanation. If the outside solution contains no oxidation-reduction sys- tem and is at the same pH as the fluid inside the glass electrode, then, ignoring the asymmetry potential, the potentials on the glass surfaces cancel each other, since they are functions of pH and are exerted in opposite directions. Again, for simplicity, ignoring certain possible ionic effects on the platinum, there remains in the chain but a single effective source of electro- motive force, the half cell inside the glass electrode. Since a single source of electrical potential cannot be measured, there will be no E.M.F. detected in the system. Now, if quinhydrone be added to the outer liquid, a potential will be imparted to the platinum whose magnitude in comparison with the normal hy- drogen electrode will be defined by the equation given above. 1 Clark, W. M., and Cohen, B. Studies on oxidation-reduction. IT. An analysis of the theoretical relations between reduetion potentials and pH. U. S. Publie Health Reports 38: 666—683. 1923. [Vor. 22 864 ANNALS OF THE MISSOURI BOTANICAL GARDEN The effective sources of electromotive force in the glass eleetrode-platinum chain consist of only the half cell inside the glass electrode and the oxidation-reduction potential imparted to the platinum, since the interface potentials on the glass sur- faces cancel each other. The potential of the half cell inside of the glass electrode can be expressed in terms of the normal hydrogen electrode (En) by applying a suitable correction, and then the system reads the En of the oxidation-reduction system in terms, of course, of the pH, which under the conditions de- fined above is the same on both sides of the glass. Now, if the pH of the solution in which the oxidation-reduc- tion system is dispersed is changed by one pH unit, then the oxidation-reduction potential on the platinum changes by 59 millivolts (at 25? C.). However, this effect on the electromo- tive force of the oxidation-reduction system is the same quanti- tatively as that on the glass-liquid interface potential. These effects, therefore, cancel each other and the total E.M.F. of the chain will read as before. If theoriginal pH equals 1.00 on both sides of the glass membrane, then it can be seen that no matter how the pH of the oxidation-reduction system outside be changed, the reading will still represent the En of the system at pH=1.00. Butif the pH outside the glass electrode remains at 1.00, and the pH inside is changed by 1 pH unit, the pH effect is no longer cancelled and the total E.M.F. of the chain will change 59 millivolts. The chain again does not indicate the En of the oxidation-reduction system, since that obviously has not changed. But the magnitude of this change, which is the pH effect on the inner surface of the glass, is the same as the change of oxidation-reduction potential if it were varied by this same pH unit. It is now clear that the observed potential of all oxidation- reduction systems which vary 59 millivolts with a change of acidity of one pH unit will depend on the pH inside of the glass electrode, and curiously enough not that of the liquid in which the system is actually dispersed. The validity of the above discussion depends on the assump- tion that the half cell inside the glass electrode is a metal in 1935] WYND—GLASS ELECTRODE 865 equilibrium with its ions such as Ag: AgCl or Hg: HgCl. If this electrode is a noble metal in equilibrium with an oxidation- reduction system, e. g., platinum: quinhydrone, then the ob- served potential of the system will be the difference in milli- volts between the oxidation-reduction potentials of the system inside and that outside the glass electrode, both dispersed in solutions of equal hydrogen ion concentration. Reconsidera- tion of the above diseussion will show that this will be true re- gardless of how the actual pH of the two solutions differ from each other, providing only that the pH of each is such that both systems vary 59 millivolts per pH unit change. GENERAL INDEX TO VOLUME XXII New scientific names of plants and the final members of new combinations are printed in bold face type; synonyms and page numbers having reference to figures and plates, in italics; and previously published names and all other matter, in ordinary type. A Adelosaccharomycetaceae, 349 rol y Ur xL dt of, in Mis a ren 232, 238 Amelanchier canadensis in Schuyler Co., 756 Anemone err ee Papa aes of, in Missouri, 438, of cylindrica, 4 Angadeni Lindeniana, 232; Boni inflorescence structure in, 17, 1 Angad > mica, 279; cog- nata, ela fw Cururu, 286; elegans, 4% a grandi- 01 3 nitida, 300; ae €i, i, 379; sylvestr 297 t of ‘ultraviolet, ys "inpr for of human subjects, 781, of vertebrate, 775, on oxidizing enzymes of Anisolobus, 270, 271; amazonicus, 270, p. latifolius, 279; anomalus, 295; E BER a Cururu, 286, folius 286; Fockei, 286; gracilipes, 294; hebecarpus 283, glabra I tomentosus, 283, a erectus, 288: b. a 283; stmanni, n Ka appleri, 285; loncifois 5005 288, E 2 S pru Am "globe sa, 102; sinuata, 104; triloba, 104; trilobata, 104 Apoeynaeeae: Observations on the in- florescence of, (with special reference i nera of Echi- posed relationship of certain types inflorescence in, 39 ; Studies in the, lv, 153 Apocynum, inflorescence structure in, Acouci, 169; andro osaemifolium, inflorescence strueture in, 15, 15; ean- ANN. Mo. Bor. GARD., VOL. 22, 1935. nabinum, inflorescence run in, anicu med te ERR of carbon dioxide produc- e on the applieation of the glass eleetrode to the det terminati ion of oxi- ials, 861 da Aptotheca, 153; ia, 211 Arcangeliella E 365; alveolata, 365 3 Campbellae, cremea, 367; Curtisii, soidea, 367; Gardneri, s. 368; krjukowensis, 368, v owskjana, 368; laevis, 368; nana, 368; occidentalis, 368; pilosa, 368; un a A = i= eg [45] der) e un o o e. o £5 Qu m n et a | T c = . e^ Pe. o B TO Th — B 1B 438 m uM Pra "distribution of, n Mis 437 Asclepias perennis distribution of, in Mis reis ne 317, 326 B Bacteria: effect of ultra-violet, on illa of, 790, on respirational of, 821 Bolanderi, 123; Careyana, 124; doe daa, 125; gla- brescens, 125; helianthoides, 127; hir- ta, 130, var. rosea, 130; Hookeri var. hirsuta, 138; incana, 133, var. tomentosa, 134; in- (867) 868 venusta, 122; macrolepis, 132; maero- sagittata, nd te deti Fe. ai 140 119; Ba alsamorrhiza, ; floccosa, 133; poe var. in- 133; tom Basal — 9 of En 773, of human subjects, 781, of lower forms, 713, of cec es, 775 Bean plants, effect of ultra-violet on respiration and respiratory enzymes 0 Berchemia seandens, distribution of, in Missouri, Bioehemieal studies with ultra-violet, 77 Bobartia, 88 Bostryx in Apocynaceae, 4, 39 Botry EM effeet of ultra-violet on metab- olism of, Bra ver 88; angustifolia, 91; laevi- gata, so pallida, Sal paradoza, 94; pur. , 89; ten 92 Bumelia lanuginosa, Bateibation of, in Missou Buphihalmum sagittatum, 127 € iei frs eanadensis var. Macoun of, in Missouri, 438 , distribution Cameraria Guyanensis, 252, 269; lutea, 25 aqua , 252 Gebiet andre of Hemispora coremiformis, 325, of Posadasia cap- sulata, 346 Carbon dioxide: elimination of, 853; an apparatus for the det ermination of, in pen cleat studies, 361, 36 Catalase, effect of ultra-violet on: of — 802, of baeteria, 821, of plants, 805, of bean plants, 849, of higher plants, 821, of tomato plants, a. M en of the flower- 5 ing plants of Misso Catalpa eg OMI Aia , 100; glo bosu MI. EO alabamensis, in D aepusastá Mo., ? en american 89 omes, 100; fastigiat, 112 cin n Apocynaceae, 4 , 49 Daireler Beccari, 366; vulvarius, 369 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Clematis Fremontii in Jefferson Co., Mo., Tue» irradiation as a factor of, 764, Coeeidioidenceae , 349 Coecidioides immitis, didi. , 270; panicu 30 Compositae, A critical study of certain epappose gene: f the Heliantheae- Verbesininae of “the natural family, 51 Conidiophorales, 326 Coreopsis heterophylla, 81 Coutoubea, inflorescence structure in, 26 ks (qp: us m 335; fareiminosus, 347, 349; , 94 Cunila. origanoides, distribution of, in Mis Cycladenia' H ario 297 Cylicaden arrisü, 297 Cytology of decisa versiforme, 307 348, 349 iculata, D Mr pron effect of ultra-violet on: mal tissues, 820, of plan ants, 821 Dichaslum in peers cynaceae, 3, 39; ag- gre 9 Dip e ps Harrisii, 297; Harrisonii, 297 A Carroll W., Sanford M. Zeller and. New species of Hydnangiaceae, Dracunculus Virginianus latifolius, 89 E Echinacea, 84; angustifolia, 91, var. 81; intermedia, 89; pallida, 93, p paradoxa, 94; pur- E anguinea, 91; tennes- 6; tuxtlensis, Ll, st; umbellata, n Hi yueatensis, 11, Echites, 270; amazonica, 279; Aa. 49; bicornis, 279; bracteata, 202; cog , 273; coriacea, 302; corym osa, 218; veda sd 285, var ee an- D 5, B. ndif olia, ; difformis, 532, 234; ; elegans, 302; gi f a, 914; geminata, 302; gracilipes, 294; grand- a, hebecarpa, mannseggiana, 296 ; hypoglauca, 301; lucida, 300; lutea, 283; macrant tha, 296; macrocarpa, 270; Meyeria 1935] INDEX 297; nitida, 300; pilosa, 159; ei ula, 234; ulcherrima, 283; punc losa, 285; salicifolia, 234 P ERU. 269; spicata, 211; stipellaris, 295; sylvestris, 297; thyrsoidea, 200; Y el- pies: 202; werrucosa, 278; Zuc- Eichitoidbae, American genera o 25 uu Observations on the infloresce of ee with Special le Plasmomyees borneensis, 370; echino- ESOS rita 365 s, A note on the applica- oxidation-reduction potentials, 861 re oe eae, 34 Enzym of animals, effect of ultra- violet. on, 802; of as P res- n ja E tory, Effects of 837, of beans, 848, of tomatoes, 837, 841 Erigeron pulchellus in Jefferson Co., Espeletia helianthoides, 127; sagittata, 127 F E grandiflora var. Ed in w woods, Butler Co., Mo. M in cells of Geotrichum prts Ferdinanda, 100; augusta, 112, var. e 110; lutescens, 112; 108 t of ne idee? on, easts, Infinen of various regions of the tan on ine; 853. iria dard inflorescence strue- Flower, TOU in Apocynaceae, 7, 39 ores Tante distribution of, in Miss 439 Porsteronia 153, inflorescenee structure 19; ci, 169; acutifolia, 161, E que EM TOT adenobasis, 207; af- finis, 166; Alex zandri, 214 ; amblybasis, 180, inflore escenee structure in, 15, 23; ea Is onthamiana, 170; bracteata, , B. bahiensis, 190; jare ird 178; uie quensis, 203; cordata, 186; coryli- folia, 211; corymbosa, 218, inflo ores- cence structur ; 19, £0; dec ecipiens, 181 difformis, "234, diospyrifolia, uckei, 177; elachista, 162, in- nee structure in, 15, 22; em- 869 belioides, 159; subg. Euforsteronia, 154; fallax Tau b., 222; floribu EU 214, inflorescence structure in, 220; foliosa, 188; = ; Gardneri nn glabrescens, aciloides, 163; a, leptocarpa, inflo- rescence structure in, 15, 20, 21; leptoca var. glaberrima, 11903 Luschnathi, 161; ron 208; mer idionalis, 159; 202; minutiflora, 159; mollis, var scu v m y insularis, TA pilosa, 159; por- Herak a Riedelii, 176; rufa v subglabra, 167; Sandwi ith: iana, 209; Schomburgkit, 169, 208, g umbellata, 208; Sellow 6 183; simu- lans, 213, in nflore escence "true ure in, 20, 21; spicata, 211, ed pibe structure in, ia 23; subcordata K. Sch., 189; tarapote nsis, 195; thyr- soidea, 199, inflorescenee structure in, 20, 21, var. glabriuscula, 201, var. typica, Sr ` umbe llata, 208; Vellozi a, orescence structu 20, 21; 2 e 171: Wilsonii, 174 Fuller, H. Ae. le Wynd, and Ernest eyno s in ultra-violet ena I. The and respiratory enzymes of higher plants, 837 Fungi: effect of ultra-violet on metab- olism of, 790; yeast-like, 307, 317, 337 G Geocarpon minimum, in Jasper Co., Mo., 750 eror Mugs Moore, Cytology of, 307, , 916 Glass eai A note on the applica- the, to the determination of tion of Bua reduetion potentials, 861 870 — agar, growth of Geotrichum rsiforme on, 3 Glutathione, 8 Gly h^ am in cells of Geotrichum versi- bs Pines i ella, 141; resinosa, 141, Growth in per Ple to external hern um 7 Gymnolomia scaposa, 83; triloba, 106 Gy ve myces Gardneri, 368; pallidus, H adietyon grandiflorum, 297 Head ge — by a new Hemi- spora, H. coremiformis, 317 Heliantheae- drid of the natural ily Compositae, A critical var of certain epappose genera of the, Helianthus invenustus, 122 terebin- inacea, — head infeetion eaused by a Hemispora 326; coremiformis, 328, 332, par arugosa, 317, 326; rugosa, 817. 326; Mr cae 317, 326 Hemisporo 317 Histo uiu. capsulatum, 347, 349 Histoplasmacea e, Histoplasmosis, Hybridella, 100; qon Wa 102 heri, 371; candidum, 370; citrinum, 371; Hisenii, 371; Gilkeyae, 371; glabrum, 365; He ssei, g 372; forme, 372; dee Po 372; iden. 372; Ravenelii, 367; So ehneri, 372; tasmanicum, 369; Thaxteri, 372; tu- berculatum TO arboreseens, distribution of, n Missouri, 44 Eire! ion "con centration, relation of, to oxidation-reduction sys stem s, 861 Hymenogaster Behrii, 366; Campbelli, Ls lut 373; nanus, 368; vio s, 369; 'viscidus, 3 73 d viscidum, 373 I Infections, fungous, 307, me. 337 peer of Apoeynaeeae: Observa- he, 1; ne of, 15, 17, 20, "" "2, 40, 32 [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Iostephane, 77; subg. Chrysopetala, 81; ubg. Euiostephane, 80; heterophylla, LN var. m artificia i animals to, 773, of human subjects, 781, of lower forms, 773, o 90, of plan tomato plants, 760, 837, of yeast, 793, 853 J Juniperus mexicana, on dolomite bluffs of White River, Barry Co., Mo. 746, 756 ; virginiana, in Jefferson Co., 9 Mo., ?48 K Kallstroemia, A new, from Texas, 49 Kallstroemia hirsuta, 49, 50 L Laseguea poete ud 190 Laubertia laxiflo Leitneria florida ana, distribution of, in Missouri, 439 Lepachys, 66; columnaris, 68, var. pul- cherrima, 70, var. Tagetes, 12; yo ont laris, 73, var. picta, 74; pinnata, 75; vcr B Pa 75. serratus, 74; Tagetes, 72 Light, reaetion bis plants to, 760 T substances in cells of Geo- richum venian me, Ligedamber Styraciflua, distribution in Missouri e -violet and respira- phenom bo y nenas of, 771 — rosea, inflorescence. structure ie areas of Missouri, 429; plants of, 431; views in, 746, 752 Lygodesmia juncea, distribution of, in Missouri, 437 M Macowanites echinosporus, 370; magnus, 369 n bar deere — 39, 30: achysiphon, 30, iss, 29, A longiflora, 31, ry Macrosiphon, 30, 1935] INDEX Malouetia, 238; albiflora, 253; amplexi- B. grandif olia, 255; glandulifera, 256; sect. bier gra 65; 263; gracilis, pu cillima, 2 esse , guianensis, 269; Guianensis, asminoides, 269; Killipii, 268; ducti, 269; O lata, 249; lat Martii, 249; Mexiae, 246; nda, 06; obtusifolia, 253; _obtusiloda, a. odorato, 252; pana is, 267; ruvian 259; ia 269; pubescena 951, retro- flexa, 243; ripa ria, 269; Schomburgki, 2 inflorescence structure in, sessilis , 269; Tamaquarina, 252, 269 ina B. brasiliensis, 253, 256, 269; virescens, Mandevilla: inflorescence structure in, 23, in Achrestogyne, 25, in bogo- ids. 25, an Free 24, 26; cuspidata, 6; folio 232, 38; glandulosa, 306. Peer EM A structure in , 24, 26, in subpaniculata, 24, 24, in xanthost toma, 24, 25 : divided nt zones to e of species, ing the plant regions, 401 Marta alone Med growth of Geotrichum versi- me on various, Pen of eoremiformis, 322, f Pos sulata, 340, of P. Mere, 343 Melanogaster ns bae 373, var. eury- sperma, 373; Eis Metabolism, effect Pn ultra-violet on: of , 773, of human subjects, 781, KEN of vertebrates, Milk, effect of S on ee activity of, Missouri: 2 en catalogue of e flowering plants of, 375; anno- 2 elevat draina 387; flora, 398; geology m i A Se ta 394; location and area, 385; map, showing Mie EN. ran abe of species, 381; sho plan egions, 401; One Be 406, views in, 746, 756 granitie-silieeous sub-region of, 411, sink-hole ponds, 409, White River desd region and bald knobs of, 413, lim 871 stone and dolomite glades of, 422; prairie region, ko glac aciated sub- division o ess hills of, 431, 746, 752, unglaciated sub- division of, 431, views in, 746, 752, 756 ; southe ast- ern lowl nis Fro die 400; tabular list of familie Mitozus, 270. poo 294; tenellus, 3 MD am. panieulata, 223 Moni Moore, Morris. ES of Geotrichum versiforme Moo Ad Head infec- ois caus u AM w Hemispor HT e pa morp hological Me physiological pues of two speeies of Posadas Morphological A physiologieal P of two species of Posadasia, 335 Mycoderma, 307 N Nectaromycetaceae, 349 Neogeotrichum pulmoneum, 348 Nyssa Wr S aistributiha of, in Mis- souri, 439 O Obeliscaria, 67 ; columnaris, 68; pinnata, 10; pulcherrima, 7 70 E ge 72 ia ic veo ys pusila, 372; ; socialis, 369; tuberculata, 3 vacu Octavianina alveolata, 365; Hessei, 371; laevis, 368, 372 ; lanigera, 372; levis, 372; tuberculatu ; Odontadenia, 270, inflorescenee struc- ture in, 6; affini , 280; amazonica, 279; angustifolia, 300; subg. Ani- solobus, ae ano ne 295; woes Anomalae, 295; augus 273; cog- nata, 273; cordata, $007 cordigera, 282; Mirror pos Cu ururu, 286; cus- pidat usendschoenäü, 300 subg. daa 297; funigera, 274; geminata, 302; 306; grandiflora, 297; Harris, 297; 872 priae MERE, YE - inflorescenee structure in, 16, 17; t. Hoffma seggianae, "996; ee term 301; Killipii, 281; Kochii, 291; lauretiana, 289; laxiflora, 305, inflorescence struc- ture in, 17, 18; sect. Laxiflorae, 305; lucida, 300; lute tea, 283; El 277; nitida, 300, a. senec 300, B. acumi nata, 300, ustifo lia, 300; sect. Nitidae, 299; grinch errr 288 ; Poeppigii, 303; E ar 304, 297; Sprucei, 279; stemmadeniaefolia, 299; surinamensis, 276; syw estris, 297; — 978; sect. Verru riniana, 284, f. folia, 284, E ovalifolia, tomentosa, 284 buat 1 missourensis in Barry Co., an er 284, var Mo. Oidiu m, 335; lactis, 307 Oospora a, 307; d’Agatae, 327 Orthechites, 224; —' 231 era! A no the application of the glass ek trode to the determination of, 861 Oxygenase, effect of ultra-violet on, 812, Oxytropis es distribution of, in Missouri, Ozark regio 3 Missouri, 406; char- acteristic plata of, 410; bald knobs, 415, plants of, 420; chert glades, 421, plants of, 422; granitie — sub- on, 411 , plan t $ views in, 746-758; White River = “regi ion, 413 P Palmer, Ernest J. and Julian A. Steyer- ceae, Paracoeeidioides, 349; brasiliensis, 348, 3 Parsonsia corymbosa, 218; MIN 214; leptocarpa, 190; spicata, Pentstemon Cobaea var. Ama er in Taney Co., Mo., 758 Perictenia, 270; stipellaris 295 Periploca umbellat , 218 Peroxidase, effect “Of very violet on of animal tissues, 815, of plants, 816, E zn ‚848, of m 839, 844, [VoL. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN Pinus noH along Little St. dis ou River, Mo., 746; in EON 756; gistzIbat B4 i 440 Populus tremuloides, distribution P in Missouri, 438 Posadasia, A morphological and physio- of, ap — "€ of two species of, P. e sulata (Darlin g) Moore and P. pyri premi ioar Posadasia, sis; capauats, 336, 340, 348, 352, 354, ; pyriformis, "336, 343 348, 358, $6 0° Prairie region of Missouri, 427; glac ated sub-division of, 427, plants of, 29, 746, 752, plants ; views in, 7 mexicana, 27, 28, in portobellensis, 27, 28, in qu inquangularis 28 Pr do luteum, Prunus rig T qa Missouri, 438 Pteridium er distribution of, n Misso Pteretis nodulosa, distribution of, in Missouri, tión of, in Q Quartz mercury are, in irradiation ex- periments, 837, 843, 854, 857 Quercus falcata, distribution of, in Mis- souri, 437, of lyrata, 439, of Shu- mardii var. Schneckeii, 440 R Raceme in Apocynaceae, 5, 3 Ratibida, 66; columnaris, 68, 150, var. Ich errima, 70,8 pulche rrima, 70; columnifera, 68, var. pulcherrima, 71; me — 76; peduncularis, 73, var. picta, 7 picta, 74; pinn ata, 75; sulcata, i Tagetes, 71; Tagetes var. cinerea. Reactions of oe wi oo bl 759 Respiration: of plan for violet on, of beans, 848, of fungi, 790, of ine plante, 800, of poi x 837, f yeast , 79 Respiratory e xe es of higher plants, Effects of P asia violet on respiration and, 800, 83 Respiratory die Studies in ultra- 1935] INDEX violet and. I. Review of work pub- lished before dios 1935 yt: The effects of ultra r us of the o the ana Te fermentation yeast, 853 Review of ultra-violet work published before June, 1935, 771 Reynolds, E. The UR BE plants to ultra- violet, 759; and F. Lyl u in ultra- violet s P respiratory phe . Revi k pu lished otis June, 1935, E T Lyle regions of th aerobie Piep et- of yeast, 853 Rhabdadenia Lindeniana, 232; lucida, 300; polyneura, 304 ISA Ri men 349; Seeberi, 349 hodode ee nudiflorum var. roseum, in Madison Co., Mo., 758 Robbia, 239; cestroides, 248; gossipina, 249; acrocar a, 26 Rosa blanda, distribution of, in Missouri, Rudbeckia chrysantha, 83; columnaris, 68; columnifera, 68; napifolia, 81; pallida, 93; pinnata, 75; purpurea, 89; serotina, 89; Tagetes, 72 S Selerogaster Broomeianus, 370; didus, 37 an 0; hysterangioides, 370; lan- atus, 371; luteocarneus, 370; pa acifi- cus, 370; sic Secondatia, 224; arborea, 249; VAM ra, ch d a genui ensiflora 225 var. paraguariensis, 225, g ds 225; diff ormis, 232, 234; Duckei, 230; ferrugineo 232; Beben, Sl foli osa, 229, g. Ga rdne lata, 229, y. he geo. Machabii, 231; peruviana, 227; Schlimiana, 228; stans, 232 Secotium (Elasmomyces) krjukowense, 368; michailowskyanum , 968 Sedum pulehellum, in J asper Co., Mo., Sharp, Ward M. A critical study of cer- e genera o the Heli- e of the natural family Comporta, 51- 873 ec De distribution of, in Mis- Smilax foem var. dpt in distribu- of, in Misso Band Lowlands Ni of Mis- of various waves of, ence of cux regio robie Ed of yeast, Spla Ran fos Behrii, 366 Sporendonema, 327; casei, 327; epizoum, 327 ns of the, on the 853 Miss p Ag grandiflora, 269; guate- mal leal, Julian A., Ernest J. Palmer "pud annotated vea pd ES the pus ing plan Missouri, Stimulation of dd under aise alee he Apocynaceae, IV. The American genera of Echitoideae, 153 Studies En ultra-violet and respiratory ena, I. Review r - lished before dir 1935, 771; II. The ets of ar ge effect of pare violet on, 8 Syringosma, 153 T Tabernaemontana amygdalifolia, 269; cestroides, 248; gracilis, 263; grandi- flora, PRA: laeta, 249; odorata, 252; populifolia, , 234; riparia, 269; tetra- staehya. Taxodium distie pup Dre of, in Missouri, 439; wampy s, Mis- sissippi Co. Mo. 7256 s ie TONES no orthern limit of, 237 Temnadenia, inflorescence structure in, 26, 27, in stellaris, 26, T in stenantha, 26, 27, in violae 97,2 Saum in fatia is growth of plant Texas, a troemia from, 49 Thalia dealbata distribution of, in Mis- souri, ee floribunda, inflorescence structure in, 48 tee ant 208; laurif olia, 178; umbellata, 2 repr 15 3; erry D basis, 207; affinis, 166; PAN 208; ' Benthamiana, 170; bracteatus, 874 202; Brasiliensis, 190; 211; ande der ro? m corymbosus 218; crebriflorus ; difformis, 934; aaaea, UM p. lioides, 159; Gardneri, 165; glabrescens, 194; gra- Thyrse in Apocynaceae, 6, 39 Tomato plants, Effects of "ultra- violet on respiration and respiratory enzymes of, 837; reaction of, to ultra-violet ir- radiation Ji Torula epizoa, 327; sacchari, 327 Cordon die, 349 Trachelospermum, 232; asiaticum, 269; Pw orme, 232, 234, distribution of, jasminoides, 233; stans, 238 een 2 Gyrains, effect of ultra-violet on, 814 U Ultra-violet: reaction to, of imals, 773, of bacteria, 790, of dh sy 848, of Botrytis, 792, of plants, 759, of green plants, 800, of tomatoes, 760, of yeas "* 793; and — ory phe nomena, Studies w of work published DES ue. 1935, 771; The effects of ultra-violet on — respiratory en of , 837; e The Pise the speetrum on the anaerobie fpecies Of yeast, 853 bd Vaccinium vacillans var. erinitum, dis- tribution of, in Missour e... in cells of Geotrichum versi- forme, 31 Pa dues. inflorescence strueture in, 81, 32 Vita. glass in ultra-violet experiments, 42, 8 Volutin in eells of Geotriehum versi- forme, 310 WwW A new Kallstroemia Williams, Louis. from Tex as, 49 corylif eoi [Vor. 22 ANNALS OF THE MISSOURI BOTANICAL GARDEN coo Robert E. Jr. Observations ocynaceae genera of eiae cci. 1; Studies in the Apocyna IV. The Am ean genera of Echitoideae, 153 Wynd, F. Lyle n improved apparatus nt sexed in ultra- eset and respiratory phen ew of work pub- lished “belate Tuo. 1935, 771; Fuller, Reyn olds, E. S. Ibid. E JJ, E The — of aded violet on res- piration respir higher siue 837; E. S. regions of the spec on the an- aerobic amó. en wy yeast, 853 X Xanthine dehydrogenase, effect of ultra- violet on, 822 Y Yeast: anaerobie fermentation of, m e n he, 853; violet on fermentation of, 793; 56 dium used, Yeast-like fungi, 307, 317, 3 Yucca or er. distribution of, in Mis- souri, Z Zaluzania, 95; angusta, 112; — 4 'asperrima, 108, Ro 113; subcordata, 111; triloba, 10 Zeller, Sanford M., and Carroll W. = cd P species of Hydnangi- Zachokkes, se VU CMM A WAT d quA Li e ESRA Y Number X. y 'of the Missouri Botanical Garden RR .. Contents rey CUPS ND hs A February, 1935. PORE on the Inflorescence of Apocynaceae (With Special Ref- -erence to me American Grace DERERKONEREN. se s dv te e i : ¿Robert E. Woodson, Jr. 1-48 CEA New Kallstroemia from Texas; 20 arca Ceu Williams 49-50 a A KTA Study of Certain URN Genera. of the Heliantheae- a g ie ......... “e's eee ee erbes sininae of the Natural gies or , Ward M. Sharp 51-152 PUBLISHED. QUARTERLY. AT FULTON, MISSOURI, _ BY. THE BOARD OF TRUSTEES OF THE MISSOURI BOTANICAL. GARDEN ST. LOUIS, MISSONAL : od Entered : as second-class matter BEUTE post ostofico at Hallen, Missouri, under the Act of March , 1879. " $ 2 9 e y T ee ^ ] re M ; Fox x poa 1 Be $ * Lu ene ty > T2. x 5 & Lilo du Annals of the Missouri Botanical Garden A Gunter Journal containing Seientifie Contributions from | si Missouri begun Garden and the Graduate Laboratory of the Henry Shaw School of Botany of Mud ir teren in affiliation with the Missouri Botanical Garden I Hem NNALS OF THE MISSOURI BOTANICAL GARD: appears four times during the ealendar year: viet Ka, opte $ and November. Four numbers constitute a volun Subscription Prhb r5 Pe T n pa volume Single Numbers.......... ach Contents of previous issues of the ANNALS OF THE MISSOURI BOTANICAL GARDEN are listed in the ap aus pa Index, published by the H. W. Wilson Co. 5 SUN. STAFF OF THE MISSOURI BOTANICAL GARDEN Director, GEORGE T. MOORE Assistant to ihe Director, KATHERINE H. LEIGH. CARROLL W. DODGE, HERMANN VON SCHRENE, Pathologist. Mycologist. JESSE M. GREENMAN, ROBERT E. WOODSON, JR., Curator of the Herbarium. ; Research Assistant. Ernest B. REYNOLDS, Davip C, FAIRBURN, Physiologist. Research Assistant. Neu C. HORNER, Librarian and Editor of Publications, BOARD OF TRUSTEES OF THE MISSOURI BOTANICAL GARDEN President, GEORGE C. HITCHCOCK. Fice-President, SAMUEL C. Davis. Second. Vice-President; Danie, K, CATLIN. L. RAY CARTER. ALBERT T. PERKINS. THOMAS 8, MAFFITT, EUGENE 8. GEORGE T. MOORE. ETHAN A, H. SHEPLEY. Frep G, ZEIBIG. EX-OFFICIO MEMBERS: BERNARD F. Dick Mayor of the Dod ot St. Louis. GEORGE R. THROOP, icone of Washington Univers a Amm ROBERT J. Terr TEE Ade President dh "The Academy oi Bi Bishop of the BR of Missouri. ence of St. Louis. SoLoN CAMERON, President of the Board of Education of St. Louis. GERALD E. UrRicr Secretary.